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Title: The Geographical Distribution of Animals, Volume I - With a study of the relations of living and extinct faunas - as elucidating the past changes of the Earth's surface
Author: Wallace, Alfred Russel
Language: English
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* * * * *
[Illustration: Zoogeographical Regions]
THE GEOGRAPHICAL
DISTRIBUTION OF ANIMALS
_WITH A STUDY OF THE RELATIONS OF LIVING AND EXTINCT FAUNAS AS ELUCIDATING
THE PAST CHANGES OF THE EARTH'S SURFACE._
BY ALFRED RUSSEL WALLACE,
AUTHOR OF "THE MALAY ARCHIPELAGO," ETC.
WITH MAPS AND ILLUSTRATIONS.
_IN TWO VOLUMES.--VOLUME I._
London: MACMILLAN AND CO. 1876.
[_The Right of Translation and Reproduction is Reserved._]
LONDON:
R. CLAY, SONS, AND TAYLOR, PRINTERS,
BREAD STREET HILL.
PREFACE.
The present work is an attempt to collect and summarize the existing
information on the Distribution of Land Animals; and to explain the more
remarkable and interesting of the facts, by means of established laws of
physical and organic change.
The main idea, which is here worked out in some detail for the whole earth,
was stated sixteen years ago in the concluding pages of a paper on the
"Zoological Geography of the Malay Archipelago," which appeared in the
_Journal of Proceedings of the Linnean Society_ for 1860; and again, in a
paper read before the Royal Geographical Society in 1863, it was briefly
summarized in the following passage:--
"My object has been to show the important bearing of researches into the
natural history of every part of the world, upon the study of its past
history. An accurate knowledge of any groups of birds or of insects and
of their geographical distribution, may enable us to map out the islands
and continents of a former epoch,--the amount of difference that exists
between the animals of adjacent districts being closely related to
preceding geological changes. By the collection of such minute facts,
alone, can we hope to fill up a great gap in the past history of the
earth as revealed by geology, and obtain some indications of the
existence of those ancient lands which now lie buried beneath the ocean,
and have left us nothing but these living records of their former
existence."
The detailed study of several groups of the birds and insects collected by
myself in the East, brought prominently before me some of the curious
problems of Geographical Distribution; but I should hardly have ventured to
treat the whole subject, had it not been for the kind encouragement of Mr.
Darwin and Professor Newton, who, about six years ago, both suggested that
I should undertake the task. I accordingly set to work; but soon became
discouraged by the great dearth of materials in many groups, the absence of
general systematic works, and the excessive confusion that pervaded the
classification. Neither was it easy to decide on any satisfactory method of
treating the subject. During the next two years, however, several important
catalogues and systematic treatises appeared, which induced me to resume my
work; and during the last three years it has occupied a large portion of my
time.
After much consideration, and some abortive trials, an outline plan of the
book was matured; and as this is, so far as I am aware, quite novel, it
will be well to give a few of the reasons for adopting it.
Most of the previous writings on Geographical Distribution appeared to me
to be unsatisfactory, because they drew their conclusions from a more or
less extensive _selection_ of facts; and did not clearly separate groups of
facts of unequal value, or those relating to groups of animals of unequal
rank. As an example of what is meant, I may refer to Mr. Andrew Murray's
large and valuable work on the Geographical Distribution of Mammalia, in
which an immense number of coloured maps are used to illustrate the
distribution of various groups of animals. These maps are not confined to
groups of any fixed rank, but are devoted to a selection of groups of
various grades. Some show the range of single species of a genus--as the
lion, the tiger, the puma, and a species of fox; others are devoted to
sections of genera,--as the true wolves; others to genera,--as the hyænas,
and the bears; others to portions of families,--as the flying squirrels,
and the oxen with the bisons; others to families,--as the Mustelidæ, and
the Hystricidæ; and others to groups of families or to orders,--as the
Insectivora, and the opossums with the kangaroos. But in no one grade are
all the groups treated alike. Many genera are wholly unnoticed, while
several families are only treated in combination with others, or are
represented by some of the more important genera.
In making these observations I by no means intend to criticise Mr. Murray's
book, but merely to illustrate by an example, the method which has been
hitherto employed, and which seems to me not well adapted to enable us to
establish the foundations of the science of distribution on a secure basis.
To do this, uniformity of treatment appeared to me essential, both as a
matter of principle, and to avoid all imputation of a partial selection of
facts, which may be made to prove anything. I determined, therefore, to
take in succession every well-established family of terrestrial
vertebrates, and to give an account of the distribution of all its
component genera, as far as materials were available. Species, as such, are
systematically disregarded,--firstly, because they are so numerous as to be
unmanageable; and, secondly, because they represent the most recent
modifications of form, due to a variety of often unknown causes, and are
therefore not so clearly connected with geographical changes as are the
natural groups of species termed genera; which may be considered to
represent the average and more permanent distribution of an organic type,
and to be more clearly influenced by the various known or inferred changes
in the organic and physical environment.
This systematic review of the distribution of families and genera, now
forms the last part of my book--Geographical Zoology; but it was nearly the
first written, and the copious materials collected for it enabled me to
determine the zoo-geographical divisions of the earth (regions and
sub-regions) to be adopted. I next drew up tables of the families and
genera found in each region and sub-region; and this afforded a basis for
the geographical treatment of the subject--Zoological Geography--the most
novel, and perhaps the most useful and generally interesting part of my
work. While this was in progress I found it necessary to make a careful
summary of the distribution of extinct Mammalia. This was a difficult task,
owing to the great uncertainty that prevails as to the affinities of many
of the fossils, and my want of practical acquaintance with Palæontology;
but having carefully examined and combined the works of the best authors, I
have given what I believe is the first connected sketch of the relation of
extinct Mammalia to the distribution of living groups, and have arrived at
some very interesting and suggestive results.
It will be observed that man is altogether omitted from the series of the
animal kingdom as here given, and some explanation of this omission may
perhaps be required. If the genus _Homo_ had been here treated like all
other genera, nothing more than the bare statement--"universally
distributed"--could have been given;--and this would inevitably have
provoked the criticism that it conveyed no information. If, on the other
hand, I had given an outline of the distribution of the _varieties_ or
_races_ of man, I should have departed from the plan of my work for no
sufficient reason. Anthropology is a science by itself; and it seems better
to omit it altogether from a zoological work, than to treat it in a
necessarily superficial manner.
The best method of illustrating a work of this kind was a matter requiring
much consideration. To have had a separate coloured or shaded map for each
family would have made the work too costly, as the terrestrial vertebrates
alone would have required more than three hundred maps. I had also doubts
about the value of this mode of illustration, as it seemed rather to
attract attention to details than to favour the development of general
views. I determined therefore to adopt a plan, suggested in conversation by
Professor Newton; and to have one general map, showing the regions and
sub-regions, which could be referred to by means of a series of numbers.
These references I give in the form of diagrammatic headings to each
family; and, when the map has become familiar, these will, I believe,
convey at a glance a body of important information.
Taking advantage of the recent extension of our knowledge of the depths of
the great oceans, I determined to give upon this map a summary of our
knowledge of the contours of the ocean bed, by means of tints of colour
increasing in intensity with the depth. Such a map, when it can be made
generally accurate, will be of the greatest service in forming an estimate
of the more probable changes of sea and land during the Tertiary period;
and it is on the effects of such changes that any satisfactory explanation
of the facts of distribution must to a great extent depend.
Other important factors in determining the actual distribution of animals
are, the zones of altitude above the sea level, and the strongly contrasted
character of the surface as regards vegetation--a primary condition for the
support of animal life. I therefore designed a series of six maps of the
regions, drawn on a uniform scale, on which the belts of altitude are shown
by contour-shading, while the forests, pastures, deserts, and perennial
snows, are exhibited by means of appropriate tints of colour.
These maps will, I trust, facilitate the study of geographical distribution
as a science, by showing, in some cases, an adequate cause in the nature of
the terrestrial surface for the actual distribution of certain groups of
animals. As it is hoped they will be constantly referred to, double folding
has been avoided, and they are consequently rather small; but Mr. Stanford,
and his able assistant in the map department, Mr. Bolton, have taken great
care in working out the details from the latest observations; and this,
combined with the clearness and the beauty of their execution, will I trust
render them both interesting and instructive.
In order to make the book more intelligible to those readers who have no
special knowledge of systematic zoology, and to whom most of the names with
which its pages are often crowded must necessarily be unmeaning, I give a
series of twenty plates, each one illustrating at once the physical aspect
and the special zoological character of some well-marked division of a
region. Great care has been taken to associate in the pictures, such
species only as do actually occur together in nature; so that each plate
represents a scene which is, at all events, not an impossible one. The
species figured all belong to groups which are either peculiar to, or very
characteristic of, the region whose zoology they illustrate; and it is
hoped that these pictures will of themselves serve to convey a notion of
the varied types of the higher animals in their true geographical
relations. The artist, Mr. J. B. Zwecker, to whose talent as a zoological
draughtsman and great knowledge both of animal and vegetable forms we are
indebted for this set of drawings, died a few weeks after he had put the
final touches to the proofs. He is known to many readers by his vigorous
illustrations of the works of Sir Samuel Baker, Livingstone, and many other
travellers,--but these, his last series of plates, were, at my special
request, executed with a care, delicacy, and artistic finish, which his
other designs seldom exhibit. It must, however, be remembered, that the
figures of animals here given are not intended to show specific or generic
characters for the information of the scientific zoologist, but merely to
give as accurate an idea as possible, of some of the more remarkable and
more restricted types of beast and bird, amid the characteristic scenery of
their native country;--and in carrying out this object there are probably
few artists who would have succeeded better than Mr. Zwecker has done.
The general arrangement of the separate parts of which the work is
composed, has been, to some extent, determined by the illustrations and
maps, which all more immediately belong to Part III. It was at first
intended to place this part last, but as this arrangement would have
brought all the illustrations into the second volume, its place was
changed,--perhaps in other respects for the better, as it naturally follows
Part II. Yet for persons not well acquainted with zoology, it will perhaps
be advisable to read the more important articles of Part IV. (and
especially the observations at the end of each order) after Part II., thus
making Part III. the conclusion of the work.
Part IV. is, in fact, a book of reference, in which the distribution of all
the families and most of the genera of the higher animals, is given in
systematic order. Part III. is treated somewhat more popularly; and,
although it is necessarily crowded with scientific names (without which the
inferences and conclusions would have nothing solid to rest on), these may
be omitted by the non-scientific reader, or merely noted as a certain
number or proportion of peculiar generic types. Many English equivalents to
family and generic names are, however, given; and, assisted by these, it is
believed that any reader capable of understanding Lyell's "Principles," or
Darwin's "Origin," will have no difficulty in following the main arguments
and appreciating the chief conclusions arrived at in the present work.
To those who are more interested in facts than in theories, the book will
serve as a kind of dictionary of the geography and affinities of animals.
By means of the copious Index, the native country, the systematic position,
and the numerical extent of every important and well established genus of
land-animal may be at once discovered;--information now scattered through
hundreds of volumes.
In the difficult matters of synonymy, and the orthography of generic names,
I have been guided rather by general utility than by any fixed rules. When
I have taken a whole family group from a modern author of repute, I have
generally followed his nomenclature throughout. In other cases, I use the
names which are to be found in a majority of modern authors, rather than
follow the strict rule of priority in adopting some newly discovered
appellation of early date. In orthography I have adopted all such modern
emendations as seem coming into general use, and which do not lead to
inconvenience; but where the alteration is such as to completely change the
pronunciation and appearance of a well-known word, I have not adopted it. I
have also thought it best to preserve the initial letter of well-known and
old-established names, for convenience of reference to the Indices of
established works. As an example I may refer to _Enicurus_,--a name which
has been in use nearly half a century, and which is to be found under the
letter _E_, in Jerdon's Birds of India, Blyth's Catalogue, Bonaparte's
Conspectus, and the Proceedings of the Zoological Society of London down to
1865. Classicists now write _Henicurus_ as the correct form; but this seems
to me one of those cases in which orthographical accuracy should give way
to priority, and still more to convenience.
In combining and arranging so much detail from such varied sources, many
errors and omissions must doubtless have occurred. Owing to my residence at
a distance from the scientific libraries of the metropolis, I was placed at
a great disadvantage; and I could hardly have completed the work at all,
had I not been permitted to have a large number of volumes at once, from
the library of the Zoological Society of London, and to keep them for
months together;--a privilege for which I return my best thanks to Mr.
Sclater the Secretary, and to the Council.
Should my book meet with the approval of working naturalists, I venture to
appeal to them, to assist me in rendering any future editions more
complete, by sending me (to the care of my publishers) notes of any
important omissions, or corrections of any misstatements of fact; as well
as copies of any of their papers or essays, and especially of any lists,
catalogues, and monographs, containing information on the classification or
distribution of living or extinct animals.
To the many friends who have given me information or assistance I beg to
tender my sincere thanks. Especially am I indebted to Professor Newton, who
not only read through much of my rough MSS., but was so good as to make
numerous corrections and critical notes. These were of great value to me,
as they often contained or suggested important additional matter, or
pointed out systematic and orthographical inaccuracies.
Professor Flower was so good as to read over my chapters on extinct
animals, and to point out several errors into which I had fallen.
Dr. Günther gave me much valuable information on the classification of
reptiles, marking on my lists the best established and most natural genera,
and referring me to reliable sources of information.
I am also greatly indebted to the following gentlemen for detailed
information on special subjects:--
To Sir Victor Brooke, for a MS. arrangement of the genera of Bovidæ, with
the details of their distribution:
To Mr. Dresser, for lists of the characteristic birds of Northern and
Arctic Europe:
To Dr. Hooker, for information on the colours and odours of New Zealand
plants:
To Mr. Kirby, for a list of the butterflies of Chili:
To Professor Mivart, for a classification of the Batrachia, and an early
proof of his article on "Apes" in the Encyclopedia Britannica:
To Mr. Salvin, for correcting my list of the birds of the Galapagos, and
for other assistance:
To Mr. Sharpe, for MS. lists of the birds of Madagascar and the Cape Verd
Islands:
To Canon Tristram, for a detailed arrangement of the difficult family of
the warblers,--Sylviidæ:
To Viscount Walden, for notes on the systematic arrangement of the
Pycnonotidæ and Timaliidæ, and for an early proof of his list of the birds
of the Philippine Islands.
I also have to thank many naturalists, both in this country and abroad, who
have sent me copies of their papers; and I trust they will continue to
favour me in the same manner.
An author may easily be mistaken in estimating his own work. I am well
aware that this first outline of a great subject is, in parts, very meagre
and sketchy; and, though perhaps overburthened with some kinds of detail,
yet leaves many points most inadequately treated. It is therefore with some
hesitation that I venture to express the hope that I have made some
approach to the standard of excellence I have aimed at;--which was, that my
book should bear a similar relation to the eleventh and twelfth chapters of
the "Origin of Species," as Mr. Darwin's "Animals and Plants under
Domestication" does to the first chapter of that work. Should it be judged
worthy of such a rank, my long, and often wearisome labours, will be well
repaid.
MARCH, 1876.
CONTENTS OF THE FIRST VOLUME.
PART I.
THE PRINCIPLES AND GENERAL PHENOMENA OF DISTRIBUTION.
CHAPTER I.
INTRODUCTORY.
CHAPTER II.
THE MEANS OF DISPERSAL AND THE MIGRATIONS OF ANIMALS.
Means of Dispersal of Mammalia (p. 10)--Climate as a Limit to the Range
of Mammals (p. 11)--Valleys and Rivers as Barriers to Mammals (p. 12)--
Arms of the Sea as Barriers to Mammals (p. 13)--Ice-floes and drift-wood
as aiding the Dispersal of Mammals (p. 14)--Means of Dispersal of Birds
(p. 15)--Dispersal of Birds by Winds (p. 16)--Barriers to the Dispersal
of Birds (p. 17)--The Phenomena of Migration (p. 18)--Migrations of Birds
(p. 19)--General remarks on Migration (p. 25)--Means of Dispersal of
Reptiles and Amphibia (p. 28)--Means of Dispersal of Fishes (p. 29)--
Means of Dispersal of Mollusca (p. 30)--Means of Dispersal of Insects and
the Barriers which limit their Range (p. 32) 10-34
CHAPTER III.
DISTRIBUTION AS AFFECTED BY THE CONDITIONS AND CHANGES OF THE EARTH'S
SURFACE.
Land and Water (p. 35)--Continental Areas (p. 36)--Recent Changes in the
Continental Areas (p. 38)--The Glacial Epoch as affecting the
Distribution of Animals (p. 40)--Changes of Vegetation as affecting the
Distribution of Animals (p. 43)--Organic Changes as affecting
Distribution (p. 44) 35-49
CHAPTER IV.
ON ZOOLOGICAL REGIONS.
Principles upon which Zoological Regions should be formed (p. 53)--Which
class of Animals is of most importance in determining Zoological Regions
(p. 56)--Various Zoological Regions proposed since 1857 (p. 58)--
Discussion of proposed Regions (p. 61)--Reasons for adopting the Six
Regions first proposed by Mr. Sclater (p. 63)--Objections to the system
of Circumpolar Zones (p. 67)--Does the Arctic Fauna characterise an
independent Region (p. 68)--Palæarctic Region (p. 71)--Ethiopian Region
(p. 73)--Oriental Region (p. 75)--Australian Region (p. 77)--Neotropical
Region (p. 78)--Nearctic Region (p. 79)--Observations on the series of
Sub-regions (p. 80) 50-82
CHAPTER V.
CLASSIFICATION AS AFFECTING THE STUDY OF GEOGRAPHICAL DISTRIBUTION.
Classification of the Mammalia (p. 85)--Classification of Birds (p. 92)--
Classification of Reptiles (p. 98)--Classification of Amphibia (p. 100)--
Classification of Fishes (p. 101)--Classification of Insects (p. 102)--
Classification of Mollusca (p. 104) 83-104
PART II.
ON THE DISTRIBUTION OF EXTINCT ANIMALS.
CHAPTER VI.
THE EXTINCT MAMMALIA OF THE OLD WORLD.
Historic and Post-pliocene Period (p. 110)--Pliocene Period (p. 112)--
General Conclusions as to the Pliocene and Post-pliocene Faunas of Europe
(p. 113)--Miocene Period (p. 114)--Extinct Animals of Greece (p. 115)--
Miocene Fauna of Central and Western Europe (p. 117)--Upper Miocene
Deposits of India (p. 121)--General Observations on the Miocene Faunas of
Europe and Asia (p. 123)--Eocene Period (p. 124)--General Considerations
on the Extinct Mammalian Fauna of Europe (p. 126) 107-128
CHAPTER VII.
EXTINCT MAMMALIA OF THE NEW WORLD.
North America--Post-pliocene Period (p. 129)--Remarks on the Post-
pliocene Fauna of North America (p. 130)--Tertiary Period (p. 132)--
Primates (p. 132)--Insectivora (p. 133)--Carnivora (p. 134)--Ungulata
(p. 135)--Proboscidea (p. 138)--Tillodontia (p. 139)--Rodentia (p. 140)--
General Relations of the Extinct Tertiary Mammalia of North America and
Europe (p. 140)--South America (p. 143)--Fauna of the Brazilian Caves
(p. 143)--Pliocene Period of Temperate South America (p. 146)--Pliocene
Mammalia of the Antilles (p. 148)--Eocene Fauna of South America (p. 148)
--General Remarks on the Extinct Mammalian Fauna of the Old and New
Worlds (p. 148)--The Birth-place and Migrations of some Mammalian
Families and Genera (p. 153) 129-156
CHAPTER VIII.
VARIOUS EXTINCT ANIMALS;--AND ON THE ANTIQUITY OF THE GENERA OF
INSECTS AND LAND-MOLLUSCA.
Extinct Mammalia of Australia (p. 157)--Mammalian Remains of the
Secondary Formations (p. 159)--Extinct Birds (p. 160)--Palæarctic Region
and North India (p. 161)--North America (p. 163)--South America,
Madagascar, New Zealand (p. 164)--Extinct Tertiary Reptiles (p. 165)--
Antiquity of the Genera of Insects (p. 166)--Antiquity of the Genera of
Land and Fresh-water Shells (p. 168) 157-170
PART III.
ZOOLOGICAL GEOGRAPHY: A REVIEW OF THE CHIEF FORMS OF LIFE
IN THE SEVERAL REGIONS AND SUB-REGIONS, WITH THE INDICATIONS
THEY AFFORD OF GEOGRAPHICAL MUTATIONS.
CHAPTER IX.
THE ORDER OF SUCCESSION OF THE REGIONS.--COSMOPOLITAN GROUPS OF
ANIMALS.--TABLES OF DISTRIBUTION.
Order of succession of the Regions (p. 173)--Cosmopolitan Groups (p. 175)
--Tables of Distributions of Families and Genera (p. 177) 173-179
CHAPTER X.
THE PALÆARCTIC REGION.
Zoological Characteristics of the Palæarctic Region (p. 181)--Summary of
Palæarctic Vertebrata (p. 186)--Insects (p. 187)--Land-shells (p. 190)--
The Palæarctic Sub-regions (p. 190)--Central and Northern Europe (p. 191)
--North European Islands (p. 197)--Mediterranean Sub-region (p. 199)--
The Mediterranean and Atlantic Islands (p. 206)--The Siberian Sub-region,
or Northern Asia (p. 216)--Japan and North China, or the Manchurian
Sub-region (p. 220)--Birds (p. 223)--Insects (p. 227)--Remarks on the
General Character of the Fauna of Japan (p. 230)--General Conclusions as
to the Fauna of the Palæarctic Region (p. 231)--Table I. Families of
Animals inhabiting the Palæarctic Region (p. 234)--Table II. List of the
Genera of Terrestrial Mammalia and Birds of the Palæarctic Region
(p. 239) 181-250
CHAPTER XI.
THE ETHIOPIAN REGION.
Zoological Characteristics of the Ethiopian Region (p. 252)--Summary of
Ethiopian Vertebrates (p. 255)--The Ethiopian Sub-regions (p. 258)--The
East African Sub-region, or Central and East Africa (p. 258)--The West
African Sub-region (p. 262)--Islands of the West African Sub-region
(p. 265)--South African Sub-region (p. 266)--Atlantic Islands of the
Ethiopian Region;--St. Helena (p. 269)--Tristan d'Acunha (p. 271)--
Madagascar and the Mascarene Islands, or the Malagasy Sub-region (p. 272)
--The Mascarene Islands (p. 280)--Extinct Fauna of the Mascarene Islands
and Madagascar (p. 282)--General Remarks on the Insect Fauna of
Madagascar (p. 284)--On the probable Past History of the Ethiopian Region
(p. 285)--Table I. Families of Animals inhabiting the Ethiopian Region
(p. 294)--Table II. List of Genera of Terrestrial Mammalia and Birds of
the Ethiopian Region (p. 300) 251-313
CHAPTER XII.
THE ORIENTAL REGION.
Zoological Characteristics of the Oriental Region (p. 315)--Summary of
Oriental Vertebrata (p. 318)--The Oriental Sub-regions (p. 321)--
Hindostan, or Indian Sub-region (p. 321)--Range of the Genera of Mammalia
which inhabit the Sub-region of Hindostan (p. 322)--Oriental, Palæarctic,
and Ethiopian Genera of Birds in Central India (p. 224)--Sub-region of
Ceylon and South India (p. 326)--The Past History of Ceylon and South
India, as indicated by its Fauna (p. 328)--Himalayan or Indo-Chinese
Sub-region (p. 329)--Islands of the Indo-Chinese Sub-region (p. 333)--
Indo-Malaya, or the Malayan Sub-region (p. 334)--Malayan Insects (p. 341)
--The Zoological Relations of the several Islands of the Indo-Malay
Sub-region (p. 345)--Philippine Islands (p. 345)--Java (p. 349)--Malacca,
Sumatra, and Borneo (p. 353)--Probable recent Geographical Changes in the
Indo-Malay Islands (p. 357)--Probable Origin of the Malayan Fauna
(p. 359)--Concluding Remarks on the Oriental Region (p. 362)--Table I.
Families of Animals inhabiting the Oriental Region (p. 365)--Table II.
Genera of Terrestrial Mammalia and Birds in the Oriental Region (p. 371)
314-386
CHAPTER XIII.
THE AUSTRALIAN REGION.
General Zoological Characteristics of the Australian Region (p. 390)--
Summary of the Australian Vertebrata (p. 397)--Supposed Land-connection
between Australia and South America (p. 398)--Insects (p. 403)--
Land-shells (p. 407)--Australian Sub-regions (p. 408)--Austro-Malayan
Sub-region (p. 409)--Papua, or the New Guinea Group (p. 409)--The
Moluccas (p. 417)--Insects--Peculiarities of the Moluccan Fauna (p. 420)
--Timor Group (p. 422)--Celebes Group (p. 426)--Origin of the Fauna of
Celebes (p. 436)--Australia and Tasmania, or the Australian Sub-region
(p. 438)--The Pacific Islands, or Polynesian Sub-region (p. 442)--Fiji,
Tonga, and Samoa Islands (p. 443)--Society and Marquesas Islands (p. 443)
--Ladrone and Caroline Islands (p. 444)--New Caledonia and the New
Hebrides (p. 444)--Sandwich Islands (p. 445)--Reptiles of the Polynesian
Sub-region (p. 448)--New Zealand Sub-region (p. 449)--Islets of the New
Zealand Sub-region (p. 453)--Reptiles, Amphibia, and Fresh-water Fishes
(p. 456)--Insects (p. 457)--The Ancient Fauna of New Zealand (p. 459)--
The Origin of the New Zealand Fauna (p. 459)--Causes of the Poverty of
Insect-life in New Zealand: its Influence on the Character of the Flora
(p. 462)--Concluding Remarks on the Early History of the Australian
Region (p. 464)--Table I. Families of Animals inhabiting the Australian
Region (p. 468)--Table II. Genera of Terrestrial Mammalia and Birds of
the Australian Region (p. 473) 387-485
Index to Vol. I 489-503
MAPS AND ILLUSTRATIONS IN VOL. I.
1. Map of the World, showing the Zoo-Geographical Regions and the
contour of the Ocean-bed _Frontispiece_
_To face page_
2. Map of the Palæarctic Region 181
3. Plate I. The Alps of Central Europe with Characteristic
Animals 195
4. Plate II. Characteristic Mammalia of Western Tartary 218
5. Plate III. Characteristic Animals of North China 226
6. Map of the Ethiopian Region 251
7. Plate IV. Characteristic Animals of East Africa 261
8. Plate V. Scene in West Africa with Characteristic Animals 264
9. Plate VI. Scene in Madagascar with Characteristic Animals 278
10. Map of the Oriental Region 315
11. Plate VII. Scene in Nepaul with Characteristic Animals 331
12. Plate VIII. A Forest in Borneo with Characteristic Mammalia 337
13. Plate IX. A Malacca Forest with some of its Peculiar Birds 340
14. Map of the Australian Region 387
15. Plate X. Scene in New Guinea with Characteristic Animals 415
16. Plate XI. The Characteristic Mammalia of Tasmania 439
17. Plate XII. The Plains of New South Wales with Characteristic
Animals 442
18. Plate XIII. Scene in New Zealand with some of its Remarkable
Birds 455
ERRATA IN VOL. I.
I have detected several misprints and small errors in the final impression,
and Dr. Meyer, who has translated the work into German, has kindly
communicated all that he has noticed. It is not thought necessary to give
here all the smaller orthographical errors, most of which will be corrected
in the Index. The following seem, however, to be of sufficient importance
to justify me in asking my readers to correct them in their copies.
Page 93, 12 lines from foot, _for_ Hocco _read_ Hoazin.
" 97, line 2, _for_ Hocco _read_ Hoazin.
" 147, 13 lines from foot, _for_ three-handed _read_ three-banded.
" 177, line 6, _for_ Lycænidæ _read_ Zygænidæ.
" 183, line 20, _for_ third _read_ fourth.
" 238, line 18, _for_ Spirigidea _read_ Sphingidea.
" 242, _insert_ | 92a | Tamias | 1 | All Northern Asia | N. America.
" 245, last line, _insert in 2nd column_ (6).
" 309, line 20, _for_ Motacilla _read_ Budytes.
" 327, 12 lines from foot, _after_ Hindostan _read_ and.
" 331, last line, for _Icthyopsis_ read _Icthyophis_.
" 340, line 15, for _Edolius_ read _Bhringa_.
" 348, line 17, _for_ Flores _read_ New Guinea.
" 371, 11 lines from foot, _for_ and Borneo _read_ Borneo and
Philippines.
" 391, 10 lines from foot, _after_ Celebes _add_ and the Papuan
Islands.
" 391, 9 lines from foot, _omit_ New Guinea or.
" 414, 6 lines from foot, for _Epimachus_ read _Seleucides_.
" 415, line 10 _for_ ditto _read_ ditto.
" 427, line 20, _after_ Celebes _add_ and on some of the Philippine
Islands.
" 427, 5 lines from foot, _for_ tusks _read_ jaw.
" 462, 15 lines from foot, _for_ p. 156 _read_ p. 166.
" 474, 9 lines from foot, _after_ Celebes _add_ Papua.
THE
GEOGRAPHICAL DISTRIBUTION
OF ANIMALS.
PART I.
_THE PRINCIPLES AND GENERAL PHENOMENA OF DISTRIBUTION._
{3}CHAPTER I.
INTRODUCTORY.
It is a fact within the experience of most persons, that the various
species of animals are not uniformly dispersed over the surface of the
country. If we have a tolerable acquaintance with any district, be it a
parish, a county, or a larger extent of territory, we soon become aware
that each well-marked portion of it has some peculiarities in its animal
productions. If we want to find certain birds or certain insects, we have
not only to choose the right season but to go to the right place. If we
travel beyond our district in various directions we shall almost certainly
meet with something new to us; some species which we were accustomed to see
almost daily will disappear, others which we have never seen before will
make their appearance. If we go very far, so as to be able to measure our
journey by degrees of latitude and longitude and to perceive important
changes of climate and vegetation, the differences in the forms of animal
life will become greater; till at length we shall come to a country where
almost everything will be new, all the familiar creatures of our own
district being replaced by others more or less differing from them.
If we have been observant during our several journeys, and have combined
and compared the facts we have collected, it will become apparent that the
change we have witnessed has been of two distinct kinds. In our own and
immediately surrounding districts, particular species appeared and
disappeared because {4}the soil, the aspect, or the vegetation, was adapted
to them or the reverse. The marshes, the heaths, the woods and forests, the
chalky downs, the rocky mountains, had each their peculiar inhabitants,
which reappeared again and again as we came to tracts of country suitable
for them. But as we got further away we began to find that localities very
similar to those we had left behind were inhabited by a somewhat different
set of species; and this difference increased with distance,
notwithstanding that almost identical external conditions might be often
met with. The first class of changes is that of _stations_; the second that
of _habitats_. The one is a _local_, the other a _geographical_ phenomenon.
The whole area over which a particular animal is found may consist of any
number of _stations_, but rarely of more than one _habitat_. Stations,
however, are often so extensive as to include the entire range of many
species. Such are the great seas and oceans, the Siberian or the Amazonian
forests, the North African deserts, the Andean or the Himalayan highlands.
There is yet another difference in the nature of the change we have been
considering. The new animals which we meet with as we travel in any
direction from our starting point, are some of them very much like those we
have left behind us, and can be at once referred to familiar types; while
others are altogether unlike anything we have seen at home. When we reach
the Alps we find another kind of squirrel, in Southern Italy a distinct
mole, in Southern Europe fresh warblers and unfamiliar buntings. We meet
also with totally new forms; as the glutton and the snowy owl in Northern,
the genet and the hoopoe in Southern, and the saiga antelope and collared
pratincole in Eastern Europe. The first series are examples of what are
termed _representative species_, the second of distinct groups or _types_
of animals. The one represents a comparatively recent modification, and an
origin in or near the locality where it occurs; the other is a result of
very ancient changes both organic and inorganic, and is connected with some
of the most curious and difficult of the problems we shall have to discuss.
{5}Having thus defined our subject, let us glance at the opinions that have
generally prevailed as to the nature and causes of the phenomena presented
by the geographical distribution of animals.
It was long thought, and is still a popular notion, that the manner in
which the various kinds of animals are dispersed over the globe is almost
wholly due to diversities of climate and of vegetation. There is indeed
much to favour this belief. The arctic regions are strongly characterised
by their white bears and foxes, their reindeer, ermine, and walruses, their
white ptarmigan, owls, and falcons; the temperate zone has its foxes and
wolves, its rabbits, sheep, beavers, and marmots, its sparrows and its song
birds; while tropical regions alone produce apes and elephants, parrots and
peacocks, and a thousand strange quadrupeds and brilliant birds which are
found nowhere in the cooler regions. So the camel, the gazelle and the
ostrich live in the desert; the bison on the prairie; the tapir, the deer,
and the jaguar in forests. Mountains and marshes, plains and rocky
precipices, have each their animal inhabitants; and it might well be
thought, in the absence of accurate inquiry, that these and other
differences would sufficiently explain why most of the regions and
countries into which the earth is popularly divided should have certain
animals peculiar to them and should want others which are elsewhere
abundant.
A more detailed and accurate knowledge of the productions of different
portions of the earth soon showed that this explanation was quite
insufficient; for it was found that countries exceedingly similar in
climate and all physical features may yet have very distinct animal
populations. The equatorial parts of Africa and South America, for example,
are very similar in climate and are both covered with luxuriant forests,
yet their animal life is widely different; elephants, apes, leopards,
guinea-fowls and touracos in the one, are replaced by tapirs,
prehensile-tailed monkeys, jaguars, curassows and toucans in the other.
Again, parts of South Africa and Australia are wonderfully similar in their
soil and climate; yet one has lions, antelopes, zebras and giraffes; the
other only kangaroos, wombats, {6}phalangers and mice. In like manner parts
of North America and Europe are very similar in all essentials of soil
climate and vegetation, yet the former has racoons, opossums, and
humming-birds; while the latter possesses moles, hedgehogs and true
flycatchers. Equally striking are the facts presented by the distribution
of many large and important groups of animals. Marsupials (opossums,
phalangers &c.) are found from temperate Van Diemen's land to the tropical
islands of New Guinea and Celebes, and in America from Chili to Virginia.
No crows exist in South America, while they inhabit every other part of the
world, not excepting Australia. Antelopes are found only in Africa and
Asia; the sloths only in South America; the true lemurs are confined to
Madagascar, and the birds-of-paradise to New Guinea.
If we examine more closely the distribution of animals in any extensive
region, we find that different, though closely allied species, are often
found on the opposite sides of any considerable barrier to their migration.
Thus, on the two sides of the Andes and Rocky Mountains in America, almost
all the mammalia, birds, and insects are of distinct species. To a less
extent, the Alps and Pyrenees form a similar barrier, and even great rivers
and river plains, as those of the Amazon and Ganges, separate more or less
distinct groups of animals. Arms of the sea are still more effective, if
they are permanent; a circumstance in some measure indicated by their
depth. Thus islands far away from land almost always have very peculiar
animals found nowhere else; as is strikingly the case in Madagascar and New
Zealand, and to a less degree in the West India islands. But shallow
straits, like the English Channel or the Straits of Malacca, are not found
to have the same effect, the animals being nearly or quite identical on
their opposite shores. A change of climate or a change of vegetation may
form an equally effective barrier to migration. Many tropical and polar
animals are pretty accurately limited by certain isothermal lines; and the
limits of the great forests in most parts of the world strictly determine
the ranges of many species.
Naturalists have now arrived at the conclusion, that by some {7}slow
process of development or transmutation, all animals have been produced
from those which preceded them; and the old notion that every species was
specially created as they now exist, at a particular time and in a
particular spot, is abandoned as opposed to many striking facts, and
unsupported by any evidence. This modification of animal forms took place
very slowly, so that the historical period of three or four thousand years
has hardly produced any perceptible change in a single species. Even the
time since the last glacial epoch, which on the very lowest estimate must
be from 50,000 to 100,000 years, has only served to modify a few of the
higher animals into very slightly different species. The changes of the
forms of animals appear to have accompanied, and perhaps to have depended
on, changes of physical geography, of climate, or of vegetation; since it
is evident that an animal which is well adapted to one condition of things
will require to be slightly changed in constitution or habits, and
therefore generally in form, structure, or colour, in order to be equally
well adapted to a changed condition of surrounding circumstances. Animals
multiply so rapidly, that we may consider them as continually trying to
extend their range; and thus any new land raised above the sea by
geological causes becomes immediately peopled by a crowd of competing
inhabitants, the strongest and best adapted of which alone succeed in
maintaining their position.
If we keep in view these facts--that the minor features of the earth's
surface are everywhere slowly changing; that the forms, and structure, and
habits of all living things are also slowly changing; while the great
features of the earth, the continents, and oceans, and loftiest mountain
ranges, only change after very long intervals and with extreme slowness; we
must see that the present distribution of animals upon the several parts of
the earth's surface is the final product of all these wonderful revolutions
in organic and inorganic nature. The greatest and most radical differences
in the productions of any part of the globe must be dependent on isolation
by the most effectual and most permanent barriers. That ocean which has
remained broadest and deepest from the most remote geological epoch {8}will
separate countries the productions of which most widely and radically
differ; while the most recently-depressed seas, or the last-formed mountain
ranges, will separate countries the productions of which are almost or
quite identical. It will be evident, therefore, that the study of the
distribution of animals and plants may add greatly to our knowledge of the
past history of our globe. It may reveal to us, in a manner which no other
evidence can, which are the oldest and most permanent features of the
earth's surface, and which the newest. It may indicate the existence of
islands or continents now sunk beneath the ocean, and which have left no
record of their existence save the animal and vegetable productions which
have migrated to adjacent lands. It thus becomes an important adjunct to
geology, which can rarely do more than determine what lands have been
raised above the waters, under what conditions and at what period; but can
seldom ascertain anything of the position or extent of those which have
sunk beneath it. Our present study may often enable us, not only to say
where lands must have recently disappeared, but also to form some judgment
as to their extent, and the time that has elapsed since their submersion.
Having thus briefly sketched the nature and objects of the subject we have
to study, it will be necessary--before entering on a detailed examination
of the zoological features of the different parts of the earth, and of the
distribution of the orders, families, and genera of animals--to examine
certain preliminary facts and principles essential for our guidance. We
must first inquire what are the powers of multiplication and dispersal of
the various groups of animals, and the nature of the barriers that most
effectually limit their range. We have then to consider the effects of
changes in physical geography and in climate; to examine the nature and
extent of such changes as have been known to occur; to determine what
others are possible or probable; and to ascertain the various modes in
which such changes affect the structure, the distribution, or the very
existence of animals.
{9}Two subjects of a different nature must next engage our attention. We
have to deal with two vast masses of facts, each involving countless
details, and requiring subdivision and grouping to be capable of
intelligible treatment. All the continents and their chief subdivisions,
and all the more important islands of the globe, have to be compared as
regards their various animal forms. To do this effectively we require a
natural division of the earth especially adapted to our purpose; and we
shall have to discuss at some length the reasons for the particular system
adopted,--a discussion which must to some extent anticipate and summarize
the conclusions of the whole work. We have also to deal with many hundreds
of families and many thousands of genera of animals, and here too a true
and natural classification is of great importance. We must therefore give a
connected view of the classification adopted in the various classes of
animals dealt with.
And lastly, as the existing distribution of animals is the result and
outcome of all preceding changes of the earth and of its inhabitants, we
require as much knowledge as we can get of the animals of each country
during past geological epochs, in order to interpret the facts we shall
accumulate. We shall, therefore, enter upon a somewhat detailed sketch of
the various forms of extinct animals that have lived upon the earth during
the Tertiary period; discuss their migrations at various epochs, the
changes of physical geography that they imply, and the extent to which they
enable us to determine the birthplace of certain families and genera.
The preliminary studies above enumerated will, it is believed, enable us to
see the bearing of many facts in the distribution of animals that would
otherwise be insoluble problems; and, what is hardly less valuable, will
teach us to estimate the comparative importance of the various groups of
animals, and to avoid the common error of cutting the gordian knot of each
difficulty by vast hypothetical changes in existing continents and
oceans--probably the most permanent features of our globe.
{10}CHAPTER II.
THE MEANS OF DISPERSAL AND THE MIGRATIONS OF ANIMALS.
All animals are capable of multiplying so rapidly, that if a single pair
were placed in a continent with abundance of food and no enemies, they
might fully stock it in a very short time. Thus, a bird which produces ten
pairs of young during its lifetime (and this is far below the fertility of
many birds) will, if we take its life at five years, increase to a hundred
millions in about forty years, a number sufficient to stock a large
country. Many fishes and insects are capable of multiplying several
thousandfold each year, so that in a few years they would reach billions
and trillions. Even large and slow breeding mammals, which have only one at
a birth but continue to breed from eight to ten successive years, may
increase from a single pair to ten millions in less than forty years.
But as animals rarely have an unoccupied country to breed in, and as the
food in any one district is strictly limited, their natural tendency is to
roam in every direction in search of fresh pastures, or new hunting
grounds. In doing so, however, they meet with many obstacles. Rocks and
mountains have to be climbed, rivers or marshes to be crossed, deserts or
forests to be traversed; while narrow straits or wider arms of the sea
separate islands from the main land or continents from each other. We have
now to inquire what facilities the different classes of animals have for
overcoming these obstacles, and what kind of barriers are most effectual in
checking their progress.
_Means of Dispersal of Mammalia._--Many of the largest mammalia are able to
roam over whole continents and are hardly {11}stopped by any physical
obstacles. The elephant is almost equally at home on plains and mountains,
and it even climbs to the highest summit of Adam's Peak in Ceylon, which is
so steep and rocky as to be very difficult of ascent for man. It traverses
rivers with great ease and forces its way through the densest jungle. There
seems therefore to be no limit to its powers of wandering, but the
necessity of procuring food and its capacity of enduring changes of
climate. The tiger is another animal with great powers of dispersal. It
crosses rivers and sometimes even swims over narrow straits of the sea, and
it can endure the severe cold of North China and Tartary as well as the
heats of the plains of Bengal. The rhinoceros, the lion, and many of the
ruminants have equal powers of dispersal; so that wherever there is land
and sufficient food, there are no limits to their possible range. Other
groups of animals are more limited in their migrations. The apes, lemurs,
and many monkeys are so strictly adapted to an arboreal life that they can
never roam far beyond the limits of the forest vegetation. The same may be
said of the squirrels, the opossums, the arboreal cats, and the sloths,
with many other groups of less importance. Deserts or open country are
equally essential to the existence of others. The camel, the hare, the
zebra, the giraffe and many of the antelopes could not exist in a forest
country any more than could the jerboas or the prairie marmots.
There are other animals which are confined to mountains, and could not
extend their range into lowlands or forests. The goats and the sheep are
the most striking group of this kind, inhabiting many of the highest
mountains of the globe; of which the European ibex and mouflon are striking
examples. Rivers are equally necessary to the existence of others, as the
beaver, otter, water-vole and capybara; and to such animals high
mountain-ranges or deserts must form an absolutely impassable barrier.
_Climate as a Limit to the Range of Mammals._--Climate appears to limit the
range of many animals, though there is some reason to believe that in many
cases it is not the climate itself so much as the change of vegetation
consequent on climate which produces the effect. The quadrumana appear to
be limited by climate, {12}since they inhabit almost all the tropical
regions but do not range more than about 10° beyond the southern and 12°
beyond the northern tropic, while the great bulk of the species are found
only within an equatorial belt about 30° wide. But as these animals are
almost exclusively fruit-eaters, their distribution depends as much on
vegetation as on temperature; and this is strikingly shown by the fact that
the _Semnopithecus schistaceus_ inhabits the Himalayan mountains to a
height of 11,000 feet, where it has been seen leaping among fir-trees
loaded with snow-wreaths! Some northern animals are bounded by the
isothermal of 32°. Such are the polar bear and the walrus, which cannot
live in a state of nature far beyond the limits of the frozen ocean; but as
they live in confinement in temperate countries, their range is probably
limited by other conditions than temperature.
We must not therefore be too hasty in concluding, that animals which we now
see confined to a very hot or a very cold climate are incapable of living
in any other. The tiger was once considered a purely tropical animal, but
it inhabits permanently the cold plains of Manchuria and the Amoor, a
country of an almost arctic winter climate. Few animals seem to us more
truly inhabitants of hot countries than the elephants and rhinoceroses; yet
in Post-tertiary times they roamed over the whole of the northern
continents to within the arctic circle; and we know that the climate was
then as cold as it is now, from their entire bodies being preserved in ice.
Some change must recently have occurred either in the climate, soil, or
vegetation of Northern Asia which led to the extinction of these
forerunners of existing tropical species; and we must always bear in mind
that similar changes may have acted upon other species which we now find
restricted within narrow limits, but which may once have roamed over a wide
and varied territory.
_Valleys and Rivers as Barriers to Mammals._--To animals which thrive best
in dry and hilly regions, a broad level and marshy valley must often prove
an effectual barrier. The difference of vegetation and of insect life,
together with an unhealthy atmosphere, no doubt often checks migration if
it is attempted. Thus {13}many animals are restricted to the slopes of the
Himalayas or to the mountains of Central India, the flat valley of the
Ganges forming a limit to their range. In other cases, however, it is the
river rather than the valley which is the barrier. In the great Amazonian
plains many species of monkeys, birds, and even insects are found up to the
river banks on one side but do not cross to the other. Thus in the lower
part of the Rio Negro two monkeys, the _Jacchus bicolor_ and the
_Brachiurus couxiou_, are found on the north bank of the river but never on
the south, where a red-whiskered _Pithecia_ is alone found. Higher up
_Ateles paniscus_ extends to the north bank of the river while _Lagothrix
humboldtii_ comes down to the south bank; the former being a native of
Guiana, the latter of Ecuador. The range of the birds of the genus
_Psophia_ or trumpeters, is also limited by the rivers Amazon, Madeira, Rio
Negro and some others; so that in these cases we are able to define the
limits of distribution with an unusual degree of accuracy, and there is
little doubt the same barriers also limit a large number of other species.
_Arms of the Sea as Barriers to Mammals._--Very few mammals can swim over
any considerable extent of sea, although many can swim well for short
distances. The jaguar traverses the widest streams in South America, and
the bear and bison cross the Mississippi; and there can be no doubt that
they could swim over equal widths of salt water, and if accidentally
carried out to sea might sometimes succeed in reaching islands many miles
distant. Contrary to the common notion pigs can swim remarkably well. Sir
Charles Lyell tells us in his "Principles of Geology" that during the
floods in Scotland in 1829, some pigs only six months old that were carried
out to sea, swam five miles and got on shore again. He also states, on the
authority of the late Edward Forbes, that a pig jumped overboard to escape
from a terrier in the Grecian Archipelago, and swam safely to shore many
miles distant. These facts render it probable that wild pigs, from their
greater strength and activity, might under favourable circumstances cross
arms of the sea twenty or thirty miles wide; and there are facts in the
distribution of this tribe of animals which seem to indicate that they have
sometimes done so. Deer {14}take boldly to the water and can swim
considerable distances, but we have no evidence to show how long they could
live at sea or how many miles they could traverse. Squirrels, rats, and
lemmings often migrate from northern countries in bands of thousands and
hundreds of thousands, and pass over rivers, lakes and even arms of the
sea, but they generally perish in the saltwater. Admitting, however, the
powers of most mammals to swim considerable distances, we have no reason to
believe that any of them could traverse without help straits of upwards of
twenty miles in width, while in most cases a channel of half that distance
would prove an effectual barrier.
_Ice-floes and Driftwood as Aiding the Dispersal of Mammals._--In the
arctic regions icebergs originate in glaciers which descend into the sea,
and often bear masses of gravel, earth, and even some vegetation on their
surfaces; and extensive level ice-fields break away and float southwards.
These might often carry with them such arctic quadrupeds as frequent the
ice, or even on rare occasions true land-animals, which might sometimes be
stranded on distant continents or islands. But a more effectual because a
more wide-spread agent, is to be found in the uprooted trees and rafts of
driftwood often floated down great rivers and carried out to sea. Such
rafts or islands are sometimes seen drifting a hundred miles from the mouth
of the Ganges with living trees erect upon them; and the Amazon, the
Orinoco, Mississippi, Congo, and most great rivers produce similar rafts.
Spix and Martius declare that they saw at different times on the Amazon,
monkeys, tiger-cats, and squirrels, being thus carried down the stream. On
the Parana, pumas, squirrels, and many other quadrupeds have been seen on
these rafts; and Admiral W. H. Smyth informed Sir C. Lyell that among the
Philippine islands after a hurricane, he met with floating masses of wood
with trees growing upon them, so that they were at first mistaken for
islands till it was found that they were rapidly drifting along. Here
therefore, we have ample means for carrying all the smaller and especially
the arboreal mammals out to sea; and although in most cases they would
perish there, yet in some favourable instances strong winds or {15}unusual
tidal currents might carry them safely to shores perhaps several hundred
miles from their native country. The fact of green trees so often having
been seen erect on these rafts is most important; for they would act as a
sail by which the raft might he propelled in one direction for several days
in succession, and thus at last reach a shore to which a current alone
would never have carried it.
There are two groups of mammals which have quite exceptional means of
dispersal--the bats which fly, and the cetacea, seals, &c., which swim. The
former are capable of traversing considerable spaces of sea, since two
North American species either regularly or occasionally visit the Bermudas,
a distance of 600 miles from the mainland. The oceanic mammals (whales and
porpoises) seem to have no barrier but temperature; the polar species being
unable to cross the equator, while the tropical forms are equally unfitted
for the cold polar waters. The shore-feeding manatees, however, can only
live where they find food; and a long expanse of rocky coast would probably
be as complete a barrier to them as a few hundred miles of open ocean. The
amphibious seals and walruses seem many of them to be capable of making
long sea journeys, some of the species being found on islands a thousand
miles apart, but none of the arctic are identical with the antartic
species.
The otters with one exception are freshwater animals, and we have no reason
to believe they could or would traverse any great distances of salt water.
In fact, they would be less liable to dispersal across arms of the sea than
purely terrestrial species, since their powers of swimming would enable
them to regain the shore if accidentally carried out to sea by a sudden
flood.
_Means of Dispersal of Birds._--It would seem at first sight that no
barriers could limit the range of birds, and that they ought to be the most
ubiquitous of living things, and little fitted therefore to throw any light
on the laws or causes of the geographical distribution of animals. This,
however, is far from being the case; many groups of birds are almost as
strictly limited by barriers as the mammalia; and from their larger numbers
and the avidity with which they have been collected, they furnish
{16}materials of the greatest value for our present study. The different
groups of birds offer remarkable contrasts in the extent of their range,
some being the most cosmopolite of the higher animals, while others are
absolutely confined to single spots on the earth's surface. The petrels
(_Procellariidæ_) and the gulls (_Laridæ_) are among the greatest
wanderers; but most of the species are confined to one or other of the
great oceans, or to the arctic or antarctic seas, a few only being found
with scarcely any variation over almost the whole globe. The sandpipers and
plovers wander along the shores as far as do the petrels over the ocean.
Great numbers of them breed in the arctic regions and migrate as far as
India and Australia, or down to Chili and Brazil; the species of the old
and new worlds, however, being generally distinct. In striking contrast to
these wide ranges we find many of the smaller perching birds, with some of
the parrots and pigeons, confined to small islands of a few square miles in
extent, or to single valleys or mountains on the mainland.
_Dispersal of Birds by Winds._--Those groups of birds which possess no
powers of flight, such as the ostrich, cassowary, and apteryx, are in
exactly the same position as mammalia as regards their means of dispersal,
or are perhaps even inferior to them; since, although they are able to
cross rivers by swimming, it is doubtful if they could remain so long in
the water as most land quadrupeds. A very large number of short-winged
birds, such as toucans, pittas, and wrens, are perhaps worse off; for they
can fly very few miles at a time, and on falling into the water would soon
be drowned. It is only the strong-flying species that can venture to cross
any great width of sea; and even these rarely do so unless compelled by
necessity to migrate in search of food, or to a more genial climate. Small
and weak birds are, however, often carried accidentally across great widths
of ocean by violent gales. This is well exemplified by the large numbers of
stragglers from North America, which annually reach the Bermudas. No less
than sixty-nine species of American birds have occurred in Europe, most of
them in Britain and Heligoland. They consist chiefly of migratory birds
which in autumn {17}return along the eastern coasts of the United States,
and often fly from point to point across bays and inlets. They are then
liable to be blown out to sea by storms, which are prevalent at this
season; and it is almost always at this time of year that their occurrence
has been noted on the shores of Europe. It may, however, be doubted whether
this is not an altogether modern phenomenon, dependent on the number of
vessels constantly on the Atlantic which afford resting-places to the
wanderers; as it is hardly conceivable that such birds as titlarks,
cuckoos, wrens, warblers, and rails, could remain on the wing without food
or rest, the time requisite to pass over 2,000 miles of ocean. It is
somewhat remarkable that no European birds reach the American coast but a
few which pass by way of Iceland and Greenland; whereas a considerable
number do reach the Azores, fully half way across; so that their absence
can hardly be due to the prevailing winds being westerly. The case of the
Azores is, however, an argument for the unassisted passage of birds for
that distance; since two of the finches are peculiar 'species,' but closely
allied to European forms, so that their progenitors must, probably, have
reached the islands before the Atlantic was a commercial highway.
_Barriers to the Dispersal of Birds._--We have seen that, as a rule, wide
oceans are an almost absolute barrier to the passage of most birds from one
continent to another; but much narrower seas and straits are also very
effectual barriers where the habits of the birds are such as to preserve
them from being carried away by storms. All birds which frequent thickets
and forests, and which feed near or on the ground, are secure from such
accidents; and they are also restricted in their range by the extent of the
forests they inhabit. In South America a large number of the birds have
their ranges determined by the extent of the forest country, while others
are equally limited to the open plains. Such species are also bounded by
mountain ranges whenever these rise above the woody region. Great rivers,
such as the Amazon, also limit the range of many birds, even when there
would seem to be no difficulty in their crossing them. The supply of food,
and the kind of vegetation, soil, and climate {18}best suited to a bird's
habits, are probably the causes which mark out the exact limits of the
range of each species; to which must be added the prevalence of enemies of
either the parent birds, the eggs, or the young. In the Malay Archipelago
pigeons abound most where monkeys do not occur; and in South America the
same birds are comparatively scarce in the forest plains where monkeys are
very abundant, while they are plentiful on the open plains and campos, and
on the mountain plateaux, where these nest-hunting quadrupeds are rarely
found. Some birds are confined to swamps, others to mountains; some can
only live on rocky streams, others on deserts or grassy plains.
_The Phenomena of Migration._--The term "migration" is often applied to the
periodical or irregular movements of all animals; but it may be questioned
whether there are any regular migrants but birds and fishes. The annual or
periodical movements of mammalia are of a different class. Monkeys ascend
the Himalayas in summer to a height of 10,000 to 12,000 feet, and descend
again in winter. Wolves everywhere descend from the mountains to the
lowlands in severe weather. In dry seasons great herds of antelopes move
southwards towards the Cape of Good Hope. The well-known lemmings, in
severe winters, at long intervals, move down from the mountains of
Scandinavia in immense numbers, crossing lakes and rivers, eating their way
through haystacks, and surmounting every obstacle till they reach the sea,
whence very few return. The alpine hare, the arctic fox, and many other
animals, exhibit similar phenomena on a smaller scale; and generally it may
be said, that whenever a favourable succession of seasons has led to a
great multiplication of any species, it must on the pressure of hunger seek
food in fresh localities. For such movements as these we have no special
term. The summer and winter movements best correspond to true migration,
but they are always on a small scale, and of limited extent; the other
movements are rather temporary incursions than true migrations.
The annual movements of many fishes are more strictly analogous to the
migration of birds, since they take place in large bodies and often to
considerable distances, and are {19}immediately connected with the process
of reproduction. Some, as the salmon, enter rivers; others, as the herring
and mackerel, approach the coast in the breeding season; but the exact
course of their migrations is unknown, and owing to our complete ignorance
of the area each species occupies in the ocean, and the absence of such
barriers and of such physical diversities as occur on the land, they are of
far less interest and less connected with our present study than the
movements of birds, to which we shall now confine ourselves.
_Migrations of Birds._--In all the temperate parts of the globe there are a
considerable number of birds which reside only a part of the year,
regularly arriving and leaving at tolerably fixed epochs. In our own
country many northern birds visit us in winter, such as the fieldfare,
redwing, snow-bunting, turnstone, and numerous ducks and waders; with a
few, like the black redstart, and (according to Rev. C. A. Johns) some of
the woodcocks from the south. In the summer a host of birds appear--the
cuckoo, the swifts and swallows, and numerous warblers, being the most
familiar,--which stay to build their nests and rear their young, and then
leave us again. These are true migrants; but a number of other birds visit
us occasionally, like the waxwing, the oriole, and the beefeater, and can
only be classed as stragglers, which, perhaps from too rapid multiplication
one year and want of food the next, are driven to extend their ordinary
range of migration to an unusual degree. We will now endeavour to sketch
the chief phenomena of migration in different countries.
_Europe._--It is well ascertained that most of the birds that spend their
spring and summer in the temperate parts of Europe pass the winter in North
Africa and Western Asia. The winter visitants, on the other hand, pass the
summer in the extreme north of Europe and Asia, many of them having been
found to breed in Lapland. The arrival of migratory birds from the south is
very constant as to date, seldom varying more than a week or two, without
any regard to the weather at the time; but the departure is less constant,
and more dependent on the weather. Thus the swallow always comes to us
about the middle {20}of April, however cold it may be, while its departure
may take place from the end of September to late in October, and is said by
Forster to occur on the first N. or N.E. wind after the 20th of September.
Almost all the migratory birds of Europe go southward to the Mediterranean,
move along its coasts east or west, and cross over in three places only;
either from the south of Spain, in the neighbourhood of Gibraltar, from
Sicily over Malta, or to the east by Greece and Cyprus. They are thus
always in sight of land. The passage of most small birds (and many of the
larger ones too) takes place at night; and they only cross the
Mediterranean when the wind is steady from near the east or west, and when
there is moonlight.
It is a curious fact, but one that seems to be well authenticated, that the
males often leave before the females, and both before the young birds,
which in considerable numbers migrate later and alone. These latter,
however, seldom go so far as the old ones; and numbers of young birds do
not cross the Mediterranean, but stay in the south of Europe. The same rule
applies to the northward migration; the young birds stopping short of the
extreme arctic regions, to which the old birds migrate.[1] When old and
young go together, however, the old birds take the lead. In the south of
Europe few of the migratory birds stay to breed, but pass on to more
temperate zones; thus, in the south of France, out of 350 species only 60
breed there. The same species is often sedentary in one part of Europe and
migratory in another; thus, the chaffinch is a constant resident in
England, Germany, and the middle of France; but a migrant in the south of
France and in Holland: the rook visits the south of France in winter only:
the _Falco tinnunculus_ is both a resident and a migrant in the south of
France, according to M. Marcel de Serres, there being two regular passages
every year, while a certain number always remain.
{21}We see, then, that migration is governed by certain intelligible laws;
and that it varies in many of its details, even in the same species,
according to changed conditions. It may be looked upon as an exaggeration
of a habit common to all locomotive animals, of moving about in search of
food. This habit is greatly restricted in quadrupeds by their inability to
cross the sea or even to pass through the highly-cultivated valleys of such
countries as Europe; but the power of flight in birds enables them to cross
every kind of country, and even moderate widths of sea; and as they mostly
travel at night and high in the air, their movements are difficult to
observe, and are supposed to be more mysterious than they perhaps are. In
the tropics birds move about to different districts according as certain
fruits become ripe, certain insects abundant, or as flooded tracts dry up.
On the borders of the tropics and the temperate zone extends a belt of
country of a more or less arid character, and liable to be parched at the
summer solstice. In winter and early spring its northern margin is verdant,
but it soon becomes burnt up, and most of its birds necessarily migrate to
the more fertile regions to the north of them. They thus follow the spring
or summer as it advances from the south towards the pole, feeding on the
young flower buds, the abundance of juicy larvæ, and on the ripening
fruits; and as soon as these become scarce they retrace their steps
homewards to pass the winter. Others whose home is nearer the pole are
driven south by cold, hunger, and darkness, to more hospitable climes,
returning northward in the early summer. As a typical example of a
migratory bird, let us take the nightingale. During the winter this bird
inhabits almost all North Africa, Asia Minor, and the Jordan Valley. Early
in April it passes into Europe by the three routes already mentioned, and
spreads over France, Britain, Denmark, and the south of Sweden, which it
reaches by the beginning of May. It does not enter Brittany, the Channel
Islands, or the western part of England, never visiting Wales, except the
extreme south of Glamorganshire, and rarely extending farther north than
Yorkshire. It spreads over Central Europe, through Austria and Hungary to
Southern Russia and the warmer parts of Siberia, {22}but it nevertheless
breeds in the Jordan Valley, so that in some places it is only the surplus
population that migrates. In August and September, all who can return to
their winter quarters.
Migrations of this type probably date back from at least the period when
there was continuous land along the route passed over; and it is a
suggestive fact that this land connection is known to have existed in
recent geological times. Britain was connected with the Continent during,
and probably before, the glacial epoch; and Gibraltar, as well as Sicily
and Malta, were also recently united with Africa, as is proved by the
fossil elephants and other large mammalia found in their caverns, by the
comparatively shallow water still existing in this part of the
Mediterranean while the remainder is of oceanic profundity, and by the
large amount of identity in the species of land animals still inhabiting
the opposite shores of the Mediterranean. The submersion of these two
tracts of land (which were perhaps of considerable extent) would be a slow
process, and from year to year the change might be hardly perceptible. It
is easy to see how the migration that had once taken place over continuous
land would be kept up, first over lagoons and marshes, then over a narrow
channel, and subsequently over a considerable sea, no one generation of
birds ever perceiving any difference in the route.
There is, however, no doubt that the sea-passage is now very dangerous to
many birds. Quails cross in immense flocks, and great numbers are drowned
at sea whenever the weather is unfavourable. Some individuals always stay
through the winter in the south of Europe, and a few even in England and
Ireland; and were the sea to become a little wider the migration would
cease, and the quail, like some other birds, would remain divided between
south Europe and north Africa. Aquatic birds are observed to follow the
routes of great rivers and lakes, and the shores of the sea. One great body
reaches central Europe by way of the Danube from the shores of the Black
Sea; another ascends the Rhone Valley from the Gulf of Lyons.
{23}_India and China._--In the peninsula of India and in China great
numbers of northern birds arrive during September and October, and leave
from March to May. Among the smaller birds are wagtails, pipits, larks,
stonechats, warblers, thrushes, buntings, shrikes, starlings, hoopoes, and
quails. Some species of cranes and storks, many ducks, and great numbers of
_Scolopacidæ_ also visit India in winter; and to prey upon these come a
band of rapacious birds--the peregrine falcon, the hobby, kestrel, common
sparrowhawk, harrier, and the short-eared owl. These birds are almost all
natives of Europe and Western Asia; they spread over all northern and
central India, mingling with the sedentary birds of the oriental fauna, and
give to the ornithology of Hindostan at this season quite a European
aspect. The peculiar species of the higher Himalayas do not as a rule
descend to the plains in winter, but merely come lower down the mountains;
and in southern India and Ceylon comparatively few of these migratory birds
appear.
In China the migratory birds follow generally the coast line, coming
southwards in winter from eastern Siberia and northern Japan; while a few
purely tropical forms travel northwards in summer to Japan, and on the
mainland as far as the valley of the Amoor.
_North America._--The migrations of birds in North America have been
carefully studied by resident naturalists, and present some interesting
features. The birds of the eastern parts of North America are pre-eminently
migratory, a much smaller proportion being permanent residents than in
corresponding latitudes in Europe. Thus, in Massachusetts there are only
about 30 species of birds which are resident all the year, while the
regular summer visitors are 106. Comparing with this our own country,
though considerably further north, the proportions are reversed; there
being 140 residents and 63 summer visitors. This difference is clearly due
to the much greater length and severity of the winter, and the greater heat
of summer, in America than with us. The number of permanent residents
increases pretty regularly as we go southward; but the number of birds at
any locality during the breeding season seems to increase as we go
{24}northward as far as Canada, where, according to Mr. Allen, more species
breed than in the warm Southern States. Even in the extreme north, beyond
the limit of forests, there are no less than 60 species which breed; in
Canada about 160; while in Carolina there are only 135, and in Louisiana,
130. The extent of the migration varies greatly, some species only going a
few degrees north and south, while others migrate annually from the tropics
to the extreme north of the continent; and every gradation occurs between
these extremes. Among those which migrate furthest are the species of
_Dendroeca_, and other American flycatching warblers (_Mniotiltidæ_), many
of which breed on the shores of Hudson's Bay, and spend the winter in
Mexico or the West Indian islands.
The great migratory movement of American birds is almost wholly confined to
the east coast; the birds of the high central plains and of California
being for the most part sedentary, or only migrating for short distances.
All the species which reach South America, and most of those which winter
in Mexico and Guatemala, are exclusively eastern species; though a few
Rocky Mountain birds range southward along the plateaux of Mexico and
Guatemala, but probably not as regular annual migrants.
In America as in Europe birds appear in spring with great regularity, while
the time of the autumnal return is less constant. More curious is the fact,
also observed in both hemispheres, that they do not all return by the same
route followed in going northwards, some species being constant visitors to
certain localities in spring but not in autumn, others in autumn but not in
spring.
Some interesting cases have been observed in America of a gradual
alteration in the extent of the migration of certain birds. A Mexican
swallow (_Hirundo lunifrons_) first appeared in Ohio in 1815. Year by year
it increased the extent of its range till by 1845 it had reached Maine and
Canada; and it is now quoted by American writers as extending its annual
migrations to Hudson's Bay. An American wren (_Troglodytes ludovicianus_)
is another bird which has spread considerably northwards since {25}the time
of the ornithologist Wilson; and the rice-bird, or "Bob-o'-link," of the
Americans, continually widens its range as rice and wheat are more
extensively cultivated. This bird winters in Cuba and other West Indian
Islands, and probably also in Mexico. In April it enters the Southern
States and passes northward, till in June it reaches Canada and extends
west to the Saskatchewan River in 54° north latitude.
_South Temperate America._--The migratory birds of this part of the world
have been observed by Mr. Hudson at Buenos Ayres. As in Europe and North
America, there are winter and summer visitors, from Patagonia and the
tropics respectively. Species of _Pyrocephalus_, _Milvulus_, swallows, and
a hummingbird, are among the most regular of the summer visitors. They are
all insectivorous birds. From Patagonia species of _Tænioptera_,
_Cinclodes_, and _Centrites_, come in winter, with two gulls, two geese,
and six snipes and plovers. Five species of swallows appear at Buenos Ayres
in spring, some staying to breed, others passing on to more temperate
regions farther south. As a rule the birds which come late and leave early
are the most regular. Some are very irregular in their movements, the
_Molothrus bonariensis_, for example, sometimes leaves early in autumn,
sometimes remains all the winter. Some resident birds also move in winter
to districts where they are never seen in summer.
_General Remarks on Migration._--The preceding summary of the main facts of
migration (which might have been almost indefinitely extended, owing to the
great mass of detailed information that exists on the subject) appears to
accord with the view already suggested, that the "instinct" of migration
has arisen from the habit of wandering in search of food common to all
animals, but greatly exaggerated in the case of birds by their powers of
flight and by the necessity for procuring a large amount of soft insect
food for their unfledged young. Migration in its simple form may be best
studied in North America, where it takes place over a continuous land
surface with a considerable change of climate from south to north. We have
here (as probably in Europe and elsewhere) every grade of migration, from
species which merely shift the northern and southern {26}limits of their
range a few hundred miles, so that in the central parts of the area the
species is a permanent resident, to others which move completely over 1,000
miles of latitude, so that in all the intervening districts they are only
known as birds of passage. Now, just as the rice-bird and the Mexican
swallow have extended their migrations, owing to favourable conditions
induced by human agency; so we may presume that large numbers of species
would extend their range where favourable conditions arose through natural
causes. If we go back only as far as the height of the glacial epoch, there
is reason to believe that all North America, as far south as about 40°
north latitude, was covered with an almost continuous and perennial
ice-sheet. At this time the migratory birds would extend up to this barrier
(which would probably terminate in the midst of luxuriant vegetation, just
as the glaciers of Switzerland now often terminate amid forests and
corn-fields), and as the cold decreased and the ice retired almost
imperceptibly year by year, would follow it up farther and farther
according as the peculiarities of vegetation and insect-food were more or
less suited to their several constitutions. It is an ascertained fact that
many individual birds return year after year to build their nests in the
same spot. This shows a strong local attachment, and is, in fact, the
faculty or feeling on which their very existence probably depends. For were
they to wander at random each year, they would almost certainly not meet
with places so well suited to them, and might even get into districts where
they or their young would inevitably perish. It is also a curious fact that
in so many cases the old birds migrate first, leaving the young ones
behind, who follow some short time later, but do not go so far as their
parents. This is very strongly opposed to the notion of an imperative
instinct. The old birds have been before, the young have not; and it is
only when the old ones have all or nearly all gone that the young go too,
probably following some of the latest stragglers. They wander, however,
almost at random, and the majority are destroyed before the next spring.
This is proved by the fact that the birds which return in spring are as a
rule not more numerous than those which came the {27}preceding spring,
whereas those which went away in autumn were two or three times as
numerous. Those young birds that do get back, however, have learnt by
experience, and the next year they take care to go with the old ones. The
most striking fact in favour of the "instinct" of migration is the
"agitation," or excitement, of confined birds at the time when their wild
companions are migrating. It seems probable, however, that this is what may
be called a social excitement, due to the anxious cries of the migrating
birds; a view supported by the fact stated by Marcel de Serres, that the
black swan of Australia, when domesticated in Europe, sometimes joins wild
swans in their northward migration. We must remember too that migration at
the proper time is in many cases absolutely essential to the existence of
the species; and it is therefore not improbable that some strong social
emotion should have been gradually developed in the race, by the
circumstance that all who for want of such emotion did not join their
fellows inevitably perished.
The mode by which a passage originally overland has been converted into one
over the sea offers no insuperable difficulties, as has already been
pointed out. The long flights of some birds without apparently stopping on
the way is thought to be inexplicable, as well as their finding their
nesting-place of the previous year from a distance of many hundreds or even
a thousand miles. But the observant powers of animals are very great; and
birds flying high in the air may be guided by the physical features of the
country spread out beneath them in a way that would be impracticable to
purely terrestrial animals.
It is assumed by some writers that the breeding-place of a species is to be
considered as its true home rather than that to which it retires in winter;
but this can hardly be accepted as a rule of universal application. A bird
can only breed successfully where it can find sufficient food for its
young; and the reason probably why so many of the smaller birds leave the
warm southern regions to breed in temperate or even cold latitudes, is
because caterpillars and other soft insect larvæ are there abundant at the
proper time, while in their winter home the {28}larvæ have all changed into
winged insects. But this favourable breeding district will change its
position with change of climate; and as the last great change has been one
of increased warmth in all the temperate zones, it is probable that many of
the migratory birds are comparatively recent visitors. Other changes may
however have taken place, affecting the vegetation and consequently the
insects of a district; and we have seldom the means of determining in any
particular case in what direction the last extension of range occurred. For
the purposes of the study of geographical distribution therefore, we must,
except in special cases, consider the true range of a species to comprise
all the area which it occupies regularly for any part of the year, while
all those districts which it only visits at more or less distant intervals,
apparently driven by storms or by hunger, and where it never regularly or
permanently settles, should not be included as forming part of its area of
distribution.
_Means of Dispersal of Reptiles and Amphibia._--If we leave out of
consideration the true marine groups--the turtles and sea-snakes--reptiles
are scarcely more fitted for traversing seas and oceans than are mammalia.
We accordingly find that in those oceanic islands which possess no
indigenous mammals, land reptiles are also generally wanting. The several
groups of these animals, however, differ considerably both in their means
of dispersal and in their power of resisting adverse conditions. Snakes are
most dependent on climate, becoming very scarce in temperate and cold
climates and entirely ceasing at 62° north latitude, and they do not ascend
very lofty mountains, ceasing at 6,000 feet elevation in the Alps. Some
inhabit deserts, others swamps and marshes, while many are adapted for a
life in forests. They swim rivers easily, but apparently have no means of
passing the sea, since they are very rarely found on oceanic islands.
Lizards are also essentially tropical, but they go somewhat farther north
than snakes, and ascend higher on the mountains, reaching 10,000 feet in
the Alps. They possess too some unknown means (probably in the egg-state)
of passing over the ocean, since they are found to inhabit many islands
where there are neither mammalia nor snakes.
{29}The amphibia are much less sensitive to cold than are true reptiles,
and they accordingly extend much farther north, frogs being found within
the arctic circle. Their semi-aquatic life also gives them facilities for
dispersal, and their eggs are no doubt sometimes carried by aquatic birds
from one pond or stream to another. Salt water is fatal to them as well as
to their eggs, and hence it arises that they are seldom found in those
oceanic islands from which mammalia are absent. Deserts and oceans would
probably form the most effectual barriers to their dispersal; whereas both
snakes and lizards abound in deserts, and have some means of occasionally
passing the ocean which frogs and salamanders do not seem to possess.
_Means of Dispersal of Fishes._--The fact that the same species of
freshwater fish often inhabit distinct river systems, proves that they have
some means of dispersal over land. The many authentic accounts of fish
falling from the atmosphere, indicate one of the means by which they may be
transferred from one river basin to another, viz., by hurricanes and
whirlwinds, which often carry up considerable quantities of water and with
it fishes of small size. In volcanic countries, also, the fishes of
subterranean streams may sometimes be thrown up by volcanic explosions, as
Humboldt relates happened in South America. Another mode by which fishes
may be distributed is by their eggs being occasionally carried away by
aquatic birds; and it is stated by Gmelin that geese and ducks during their
migrations feed on the eggs of fish, and that some of these pass through
their bodies with their vitality unimpaired.[2] Even water-beetles flying
from one pond to another might occasionally carry with them some of the
smaller eggs of fishes. But it is probable that fresh-water fish are also
enabled to migrate by changes of level causing streams to alter their
course and carry their waters into adjacent basins. On plateaux the sources
of distinct river systems often approach each other, and the same thing
occurs with lateral tributaries on the lowlands near their mouths. Such
changes, although small in extent, and occurring only at long intervals,
would {30}act very powerfully in modifying the distribution of fresh-water
fish.
Sea fish would seem at first sight to have almost unlimited means of
dispersal, but this is far from being the case. Temperature forms a
complete barrier to a large number of species, cold water being essential
to many, while others can only dwell in the warmth of the tropics. Deep
water is another barrier to large numbers of species which are adapted to
shores and shallows; and thus the Atlantic is quite as impassable a gulf to
most fishes as it is to birds. Many sea fishes migrate to a limited extent
for the purpose of depositing their spawn in favourable situations. The
herring, an inhabitant of the deep sea, comes in shoals to our coast in the
breeding season; while the salmon quits the northern seas and enters our
rivers, mounting upwards to the clear cold water near their sources to
deposit its eggs. Keeping in mind the essential fact that changes of
temperature and of depth are the main barriers to the dispersal of fish, we
shall find little difficulty in tracing the causes that have determined
their distribution.
_Means of Dispersal of Mollusca._--The marine, fresh-water, and land
mollusca are three groups whose powers of dispersal and consequent
distribution are very different, and must be separately considered. The
_Pteropoda_, the _Ianthina_, and other groups of floating molluscs, drift
about in mid-ocean, and their dispersal is probably limited chiefly by
temperature, but perhaps also by the presence of enemies or the scarcity of
proper food. The univalve and bivalve mollusca, of which the whelk and the
cockle may be taken as types, move so slowly in their adult state, that we
should expect them to have an exceedingly limited distribution; but the
young of all these are free swimming embryos, and they thus have a powerful
means of dispersal, and are carried by tides and currents so as ultimately
to spread over every shore and shoal that offers conditions favourable for
their development. The fresh water molluscs, which one might at first
suppose could not range beyond their own river-basin, are yet very widely
distributed in common with almost all other fresh water productions; and
Mr. Darwin has shown that this is {31}due to the fact, that ponds and
marshes are constantly frequented by wading and swimming birds which are
pre-eminently wanderers, and which frequently carry away with them the
seeds of plants, and the eggs of molluscs and aquatic insects. Fresh water
molluscs just hatched were found to attach themselves to a duck's foot
suspended in an aquarium; and they would thus be easily carried from one
lake or river to another, and by the help of different species of aquatic
birds, might soon spread all over the globe. Even a water-beetle has been
caught with a small living shell (_Ancylus_) attached to it; and these fly
long distances and are liable to be blown out to sea, one having been
caught on board the _Beagle_ when forty-five miles from land. Although
fresh water molluscs and their eggs must frequently be carried out to sea,
yet this cannot lead to their dispersal, since salt water is almost
immediately fatal to them; and we are therefore forced to conclude that the
apparently insignificant and uncertain means of dispersal above alluded to
are really what have led to their wide distribution. The true land-shells
offer a still more difficult case, for they are exceedingly sensitive to
the influence of salt water; they are not likely to be carried by aquatic
birds, and yet they are more or less abundant all over the globe,
inhabiting the most remote oceanic islands. It has been found, however,
that land-shells have the power of lying dormant a long time. Some have
lived two years and a half shut up in pill boxes; and one Egyptian desert
snail came to life after having been glued down to a tablet in the British
Museum for four years!
We are indebted to Mr. Darwin for experiments on the power of land shells
to resist sea water, and he found that when they had formed a membranous
diaphragm over the mouth of the shell they survived many days' immersion
(in one case fourteen days); and another experimenter, quoted by Mr.
Darwin, found that out of one hundred land shells immersed for a fortnight
in the sea, twenty-seven recovered. It is therefore quite possible for them
to be carried in the chinks of drift wood for many hundred miles across the
sea, and this is probably one of the most effectual modes of their
dispersal. Very young shells would also {32}sometimes attach themselves to
the feet of birds walking or resting on the ground, and as many of the
waders often go far inland, this may have been one of the methods of
distributing species of land shells; for it must always be remembered that
nature can afford to wait, and that if but once in a thousand years a
single bird should convey two or three minute snails to a distant island,
this is all that is required for us to find that island well stocked with a
great and varied population of land shells.
_Means of Dispersal of Insects and the Barriers which Limit their
Range._--Winged insects, as a whole, have perhaps more varied means of
dispersal over the globe than any other highly organised animals. Many of
them can fly immense distances, and the more delicate ones are liable to be
carried by storms and hurricanes over a wide expanse of ocean. They are
often met with far out at sea. Hawk-moths frequently fly on board ships as
they approach the shores of tropical countries, and they have sometimes
been captured more than 250 miles from the nearest land. Dragon-flies came
on board the _Adventure_ frigate when fifty miles off the coast of South
America. A southerly wind brought flies in myriads to Admiral Smyth's ship
in the Mediterranean when he was 100 miles distant from the coast of
Africa. A large Indian beetle (_Chrysochroa ocellata_) was quite recently
caught alive in the Bay of Bengal by Captain Payne of the barque _William
Mansoon_, 273 miles from the nearest land. Darwin caught a locust 370 miles
from land; and in 1844 swarms of locusts several miles in extent, and as
thick as the flakes in a heavy snowstorm, visited Madeira. These must have
come with perfect safety more than 300 miles; and as they continued flying
over the island for a long time, they could evidently have travelled to a
much greater distance. Numbers of living beetles belonging to seven genera,
some aquatic and some terrestrial, were caught by Mr. Darwin in the open
sea, seventeen miles from the coast of South America, and they did not seem
injured by the salt water. Almost all the accidental causes that lead to
the dispersal of the higher animals would be still more favourable for
insects. Floating trees could carry hundreds of insects for one bird or
mammal; and so many of the larvæ, eggs, {33}and pupæ of insects have their
abode in solid timber, that they might survive being floated immense
distances. Great numbers of tropical insects have been captured in the
London docks, where they have been brought in foreign timber; and some have
emerged from furniture after remaining torpid for many years. Most insects
have the power of existing weeks or months without food, and some are very
tenacious of life. Many beetles will survive immersion for hours in strong
spirit; and water a few degrees below the boiling point will not always
kill them. We can therefore easily understand how, in the course of ages
insects may become dispersed by means which would be quite inadequate in
the case of the higher animals. The drift-wood and tropical fruits that
reach Ireland and the Orkneys; the double cocoa-nuts that cross the Indian
ocean from the Seychelle Islands to the coast of Sumatra; the winds that
carry volcanic dust and ashes for thousands of miles; the hurricanes that
travel in their revolving course over wide oceans; all indicate means by
which a few insects may, at rare intervals be carried to remote regions,
and become the progenitors of a group of allied forms.
But the dispersal of insects requires to be looked at from another point of
view. They are, of all animals, perhaps the most wonderfully adapted for
special conditions; and are so often fitted to fill one place in nature and
one only, that the barriers against their permanent displacement are almost
as numerous and as effective as their means of dispersal. Hundreds of
species of lepidoptera, for example, can subsist in the larva state only on
one species of plant; so that even if the perfect insects were carried to a
new country, the continuance of the race would depend upon the same or a
closely allied plant being abundant there. Other insects require succulent
vegetable food all the year round, and are therefore confined to tropical
regions; some can live only in deserts, others in forests; some are
dependent on water-plants, some on mountain-vegetation. Many are so
intimately connected with other insects during some part of their existence
that they could not live without them; such are the parasitical hymenoptera
and diptera, and those mimicking species whose welfare depends upon their
being {34}mistaken for something else. Then again, insects have enemies in
every stage of their existence--the egg, the larva, the pupa, and the
perfect form; and the abundance of any one of these enemies may render
their survival impossible in a country otherwise well suited to them. Ever
bearing in mind these two opposing classes of facts, we shall not be
surprised at the enormous range of some groups of insects, and at the
extreme localization of others; and shall be able to give a rational
account of many phenomena of distribution that would otherwise seem quite
unintelligible.
{35}CHAPTER III.
DISTRIBUTION AS AFFECTED BY THE CONDITIONS AND CHANGES OF THE EARTH'S
SURFACE.
The distribution of animals over the earth's surface, is evidently
dependent in great measure upon those grand and important characteristics
of our globe, the study of which is termed physical geography. The
proportion of land and water; the outlines and distribution of continents;
the depth of seas and oceans; the position of islands; the height,
direction, and continuity of mountain chains; the position and extent of
deserts, lakes, and forests; the direction and velocity of ocean currents,
as well as of prevalent winds and hurricanes; and lastly, the distribution
of heat and cold, of rain, snow, and ice, both in their means and in their
extremes, have all to be considered when we endeavour to account for the
often unequal and unsymmetrical manner in which animals are dispersed over
the globe. But even this knowledge is insufficient unless we inquire
further as to the evidence of permanence possessed by each of these
features, in order that we may give due weight to the various causes that
have led to the existing facts of animal distribution.
_Land and Water._--The well-known fact that nearly three-fourths of the
surface of the earth is occupied by water, and but a little more than
one-fourth by land, is important as indicating the vast extent of ocean by
which many of the continents and islands are separated from each other. But
there is another fact {36}which greatly increases its importance, namely,
that the mean height of the land is very small compared with the mean depth
of the sea. It has been estimated by Humboldt that the mean height of all
the land surface does not exceed a thousand feet, owing to the comparative
narrowness of mountain ranges and the great extent of alluvial plains and
valleys; the ocean bed, on the contrary, not only descends deeper than the
tops of the highest mountains rise above its surface, but these profound
depths are broad sunken plains, while the shallows correspond to the
mountain ranges, so that its mean depth is, as nearly as can be estimated,
twelve thousand feet.[3] Hence, as the area of water is three times that of
the land, the total cubical contents of the land, above the sea level,
would be only that of the waters which are below that level. The important
result follows, that whereas it is scarcely possible that in past times the
amount of land surface should ever greatly have exceeded that which now
exists, it is just possible that all the land may have been at some time
submerged; and therefore in the highest degree probable that among the
continual changes of land and sea that have been always going on, the
amount of land surface has often been much less than it is now. For the
same reason it is probable that there have been times when large masses of
land have been more isolated from the rest than they are at present; just
as South America would be if North America were submerged, or as Australia
would become if the Malay Archipelago were to sink beneath the ocean. It is
also very important to bear in mind the fact insisted on by Sir Charles
Lyell, that the shallow parts of the ocean are almost always in the
vicinity of land; and that an amount of elevation that would make little
difference to the bed of the ocean, would raise up extensive tracts of dry
land in the vicinity of existing continents. It is almost certain,
therefore, that changes in the distribution of land and sea must have taken
place more frequently by additions to, or {37}modifications of pre-existing
land, than by the upheaval of entirely new continents in mid-ocean. These
two principles will throw light upon two constantly recurring groups of
facts in the distribution of animals,--the restriction of peculiar forms to
areas not at present isolated,--and on the other hand, the occurrence of
allied forms in lands situated on opposite shores of the great oceans.
_Continental Areas._--Although the dry land of the earth's surface is
distributed with so much irregularity, that there is more than twice as
much north of the equator as there is south of it, and about twice as much
in the Asiatic as in the American hemisphere; and, what is still more
extraordinary, that on a hemisphere of which a point in St. George's
Channel between England and Ireland is the centre, the land is nearly equal
in extent to the water, while in the opposite hemisphere it is in the
proportion of only one-eighth,--yet the whole of the land is almost
continuous. It consists essentially of only three masses: the American, the
Asia-African, and the Australian. The two former are only separated by
thirty-six miles of shallow sea at Behring's Straits, so that it is
possible to go from Cape Horn to Singapore or the Cape of Good Hope without
ever being out of sight of land; and owing to the intervention of the
numerous islands of the Malay Archipelago the journey might be continued
under the same conditions as far as Melbourne and Hobart Town. This curious
fact, of the almost perfect continuity of all the great masses of land
notwithstanding their extremely irregular shape and distribution, is no
doubt dependent on the circumstances just alluded to; that the great depth
of the oceans and the slowness of the process of upheaval, has almost
always produced the new lands either close to, or actually connected with
pre-existing lands; and this has necessarily led to a much greater
uniformity in the distribution of organic forms, than would have prevailed
had the continents been more completely isolated from each other.
The isthmuses which connect Africa with Asia, and North with South America,
are, however, so small and insignificant compared with the vast extent of
the countries they unite that {38}we can hardly consider them to form more
than a nominal connection. The Isthmus of Suez indeed, being itself a
desert, and connecting districts which for a great distance are more or
less desert also, does not effect any real union between the luxuriant
forest-clad regions of intertropical Asia and Africa. The Isthmus of Panama
is a more effectual line of union, since it is hilly, well watered, and
covered with luxuriant vegetation; and we accordingly find that the main
features of South American zoology are continued into Central America and
Mexico. In Asia a great transverse barrier exists, dividing that continent
into a northern and southern portion; and as the lowlands occur on the
south and the highlands on the north of the great mountain range, which is
situated not far beyond the tropic, an abrupt change of climate is
produced; so that a belt of about a hundred miles wide, is all that
intervenes between a luxuriant tropical region and an almost arctic waste.
Between the northern part of Asia, and Europe, there is no barrier of
importance; and it is impossible to separate these regions as regards the
main features of animal life. Africa, like Asia, has a great transverse
barrier, but it is a desert instead of a mountain chain; and it is found
that this desert is a more effectual barrier to the diffusion of animals
than the Mediterranean Sea; partly because it coincides with the natural
division of a tropical from a temperate climate, but also on account of
recent geological changes which we shall presently allude to. It results
then from this outline sketch of the earth's surface, that the primary
divisions of the geographer correspond approximately with those of the
zoologist. Some large portion of each of the popular divisions forms the
nucleus of a zoological region; but the boundaries are so changed that the
geographer would hardly recognise them: it has, therefore, been found
necessary to give them those distinct names which will be fully explained
in our next chapter.
_Recent Changes in the Continental Areas._--The important fact has been now
ascertained, that a considerable portion of the Sahara south of Algeria and
Morocco was under water at a very recent epoch. Over much of this area
sea-shells, identical with those now living in the Mediterranean, are
abundantly scattered, {39}not only in depressions below the level of the
sea but up to a height of 900 feet above it. Borings for water made by the
French government have shown, that these shells occur twenty feet deep in
the sand; and the occurrence of abundance of salt, sometimes even forming
considerable hills, is an additional proof of the disappearance of a large
body of salt water. The common cockle is one of the most abundant of the
shells found; and the Rev. H. B. Tristram discovered a new fish, in a salt
lake nearly 300 miles inland, but which has since been found to inhabit the
Gulf of Guinea. Connected with this proof of recent elevation in the
Sahara, we have most interesting indications of subsidence in the area of
the Mediterranean, which were perhaps contemporaneous. Sicily and Malta are
connected with Africa by a submerged bank from 300 to 1,200 feet below the
surface; while the depth of the Mediterranean, both to the east and west,
is enormous, in some parts more than 13,000 feet; and another submerged
bank with a depth of 1,000 feet occurs at the straits of Gibraltar. In
caves in Sicily, remains of the living African elephant have been found by
Baron Anca; and in other caves Dr. Falconer discovered remains of the
_Elephas antiquus_ and of two species of _Hippopotamus_. In Malta, three
species of elephant have been discovered by Captain Spratt; a large one
closely allied to _E. antiquus_ and two smaller ones not exceeding five
feet high when adult. These facts clearly indicate, that when North Africa
was separated by a broad arm of the sea from the rest of the continent, it
was probably connected with Europe; and this explains why zoologists find
themselves obliged to place it along with Europe in the same zoological
region.
Besides this change in the level of the Sahara and the Mediterranean basin,
Europe has undergone many fluctuations in its physical geography in very
recent times. In Wales, abundance of sea-shells of living species have been
found at an elevation of 1,300 feet; and in Sardinia there is proof of an
elevation of 300 feet since the human epoch; and these are only samples of
many such changes of level. But these changes, though very important
locally and as connected with geological problems, need not be further
noticed here; as they were not of a {40}nature to affect the larger
features of the earth's surface or to determine the boundaries of great
zoological regions.
The only other recent change of great importance which can be adduced to
illustrate our present subject, is that which has taken place between North
and South America. The living marine shells of the opposite coasts of the
isthmus of Panama, as well as the corals and fishes, are generally of
distinct species, but some are identical and many are closely allied; the
West Indian fossil shells and corals of the Miocene period, however, are
found to be largely identical with those of the Pacific coast. The fishes
of the Atlantic and Pacific shores of America are as a rule very distinct;
but Dr. Günther has recently shown that a considerable number of species
inhabiting the seas on opposite sides of the isthmus are absolutely
identical. These facts certainly indicate, that during the Miocene epoch a
broad channel separated North and South America; and it seems probable that
a series of elevations and subsidences have taken place uniting and
separating them at different epochs; the most recent submersion having
lasted but a short time, and thus, while allowing the passage of abundance
of locomotive fishes, not admitting of much change in the comparatively
stationary mollusca.
_The Glacial Epoch as affecting the Distribution of Animals._--The
remarkable refrigeration of climate in the northern hemisphere within the
epoch, of existing species, to which the term Glacial epoch is applied,
together with the changes of level that accompanied and perhaps assisted to
produce it, has been one of the chief agents in determining many of the
details of the existing distribution of animals in temperate zones. A
comparison of the effects produced by existing glaciers with certain
superficial phenomena in the temperate parts of Europe and North America,
renders it certain that between the Newer Pliocene and the Recent epochs, a
large portion of the northern hemisphere must have been covered with a
sheet of ice several thousand feet thick, like that which now envelopes the
interior of Greenland. Much further south the mountains were covered with
perpetual snow, and sent glaciers down every valley; and all the {41}great
valleys on the southern side of the Alps poured down streams of ice which
stretched far out into the plains of Northern Italy, and have left their
débris in the form of huge mountainous moraines, in some cases more than a
thousand feet high. In Canada and New Hampshire the marks of moving ice are
found on the tops of mountains from 3,000 to 5,000 feet high; and the whole
surface of the country around and to the north of the great lakes is scored
by glaciers. Wherever the land was submerged during a part of this cold
period, a deposit called boulder-clay, or glacial-drift has been formed.
This is a mass of sand, clay, or gravel, full of angular or rounded stones
of all sizes, up to huge blocks as large as a cottage; and especially
characterized by these stones being distributed confusedly through it, the
largest being as often near the top as near the bottom, and never sorted
into layers of different sizes as in materials carried by water. Such
deposits are known to be formed by glaciers and icebergs; when deposited on
the land by glaciers they form moraines, when carried into water and thus
spread with more regularity over a wider area they form drift. This drift
is rarely found except where there is other evidence of ice-action, and
never south of the 40th parallel of latitude, to which in the northern
hemisphere signs of ice-action extend. In the southern hemisphere, in
Patagonia and in New Zealand, exactly similar phenomena occur.
A very interesting confirmation of the reality of this cold epoch is
derived from the study of fossil remains. Both the plants and animals of
the Miocene period indicate that the climate of Central Europe was
decidedly warmer or more equable than it is now; since the flora closely
resembled that of the Southern United States, with a likeness also to that
of Eastern Asia and Australia. Many of the shells were of tropical genera;
and there were numbers of large mammalia allied to the elephant,
rhinoceros, and tapir. At the same time, or perhaps somewhat earlier, a
temperate climate extended into the arctic regions, and allowed a
magnificent vegetation of shrubs and forest trees, some of them evergreen,
to flourish within twelve degrees of the Pole. In the Pliocene period we
find ourselves {42}among forms implying a climate very little different
from the present; and our own Crag formation furnishes evidence of a
gradual refrigeration of climate; since its three divisions, the Coralline,
Red, and Norwich Crags, show a decreasing number of southern, and an
increasing number of northern species, as we approach the Glacial epoch.
Still later than these we have the shells of the drift, almost all of which
are northern and many of them arctic species. Among the mammalia indicative
of cold, are the mammoth and the reindeer. In gravels and cave-deposits of
Post-Pliocene date we find the same two animals, which soon disappear as
the climate approached its present condition; and Professor Forbes has
given a list of fifty shells which inhabited the British seas before the
Glacial epoch and inhabit it still, but are all wanting in the glacial
deposits. The whole of these are found in the Newer Pliocene strata of
Sicily and the south of Europe, where they escaped destruction during the
glacial winter.
There are also certain facts in the distribution of plants, which are so
well explained by the Glacial epoch that they may be said to give an
additional confirmation to it. All over the northern hemisphere within the
glaciated districts, the summits of lofty mountains produce plants
identical with those of the polar regions. In the celebrated case of the
White Mountains in New Hampshire, United States (latitude 45°), all the
plants on the summit are arctic species, none of which exist in the
lowlands for near a thousand miles further north. It has also been remarked
that the plants of each mountain are more especially related to those of
the countries directly north of it. Thus, those of the Pyrenees and of
Scotland are Scandinavian, and those of the White Mountains are all species
found in Labrador. Now, remembering that we have evidence of an exceedingly
mild and uniform climate in the arctic regions during the Miocene period
and a gradual refrigeration from that time, it is evident that with each
degree of change more and more hardy plants would be successively driven
southwards; till at last the plains of the temperate zone would be
inhabited by plants, which were once confined to alpine heights or to the
arctic regions. {43}As the icy mantle gradually melted off the face of the
earth these plants would occupy the newly exposed soil, and would thus
necessarily travel in two directions, back towards the arctic circle and up
towards the alpine peaks. The facts are thus exactly explained by a cause
which independent evidence has proved to be a real one, and every such
explanation is an additional proof of the reality of the cause. But this
explanation implies, that in cases where the Glacial epoch cannot have so
acted alpine plants should not be northern plants; and a striking proof of
this is to be found on the Peak of Teneriffe, a mountain 12,000 feet high.
In the uppermost 4,500 feet of this mountain above the limit of trees, Von
Buch found only eleven species of plants, eight of which were peculiar; but
the whole were allied to those found at lower elevations. On the Alps or
Pyrenees at this elevation, there would be a rich flora comprising hundreds
of arctic plants; and the absence of anything corresponding to them in this
case, in which their ingress was cut off by the sea, is exactly what the
theory leads us to expect.
_Changes of Vegetation as affecting the Distribution of Animals._--As so
many animals are dependent on vegetation, its changes immediately affect
their distribution. A remarkable example of this is afforded by the
pre-historic condition of Denmark, as interpreted by means of the peat-bogs
and kitchen-middens. This country is now celebrated for its beech-trees;
oaks and pines being scarce; and it is known to have had the same
vegetation in the time of the Romans. In the peat-bogs, however, are found
deposits of oak trees; and deeper still pines alone occur. Now the
kitchen-middens tell us much of the natural history of Denmark in the early
Stone period; and a curious confirmation of the fact that Denmark like
Norway was then chiefly covered with pine forests is obtained by the
discovery, that the Capercailzie was then abundant, a bird which feeds
almost exclusively on the young shoots and seeds of pines and allied
plants. The cause of this change in the vegetation is unknown; but from the
known fact that when forests are destroyed trees, of a different kind
usually occupy the ground, we may suppose that some such change as a
temporary submergence might cause an entirely {44}different vegetation and
a considerably modified fauna to occupy the country.
_Organic Changes as affecting Distribution._--We have now briefly touched
on some of the direct effects of changes in physical geography, climate,
and vegetation, on the distribution of animals; but the indirect effects of
such changes are probably of quite equal, if not of greater importance.
Every change becomes the centre of an ever-widening circle of effects. The
different members of the organic world are so bound together by complex
relations, that any one change generally involves numerous other changes,
often of the most unexpected kind. We know comparatively little of the way
in which one animal or plant is bound up with others, but we know enough to
assure us that groups the most apparently disconnected are often dependent
on each other. We know, for example, that the introduction of goats into
St. Helena utterly destroyed a whole flora of forest trees; and with them
all the insects, mollusca, and perhaps birds directly or indirectly
dependent on them. Swine, which ran wild in Mauritius, exterminated the
Dodo. The same animals are known to be the greatest enemies of venomous
serpents. Cattle will, in many districts, wholly prevent the growth of
trees; and with the trees the numerous insects dependent on those trees,
and the birds which fed upon the insects, must disappear, as well as the
small mammalia which feed on the fruits, seeds, leaves, or roots. Insects
again have the most wonderful influence on the range of mammalia. In
Paraguay a certain species of fly abounds which destroys new-born cattle
and horses; and thus neither of these animals have run wild in that
country, although they abound both north and south of it. This inevitably
leads to a great difference in the vegetation of Paraguay, and through that
to a difference in its insects, birds, reptiles, and wild mammalia. On what
causes the existence of the fly depends we do not know, but it is not
improbable that some comparatively slight changes in the temperature or
humidity of the air at a particular season, or the introduction of some
enemy might lead to its extinction or banishment. The whole face of the
country would then soon be changed: new species would {45}come in, while
many others would be unable to live there; and the immediate cause of this
great alteration would probably be quite imperceptible to us, even if we
could watch it in progress year by year. So, in South Africa, the
celebrated Tsetse fly inhabits certain districts having well defined
limits; and where it abounds no horses, dogs, or cattle can live. Yet
asses, zebras, and antelopes are unaffected by it. So long as this fly
continues to exist, there is a living barrier to the entrance of certain
animals, quite as effectual as a lofty mountain range or a wide arm of the
sea. The complex relations of one form of life with others is nowhere
better illustrated than in Mr. Darwin's celebrated case of the cats and
clover, as given in his _Origin of Species_, 6th ed., p. 57. He has
observed that both wild heartsease and red-clover are fertilized in this
country by humble-bees only, so that the production of seed depends on the
visits of these insects. A gentleman who has specially studied humble-bees
finds that they are largely kept down by field-mice, which destroy their
combs and nests. Field-mice in their turn are kept down by cats; and
probably also by owls; so that these carnivorous animals are really the
agents in rendering possible the continued existence of red-clover and wild
heartsease. For if they were absent, the field-mice having no enemies,
would multiply to such an extent as to destroy all the humble-bees; and
these two plants would then produce no seed and soon become extinct.
Mr. Darwin has also shown that one species often exterminates another
closely allied to it, when the two are brought into contact. One species of
swallow and thrush are known to have increased at the expense of allied
species. Rats, carried all over the world by commerce, are continually
extirpating other species of rats. The imported hive-bee is, in Australia,
rapidly exterminating a native stingless bee. Any slight change, therefore,
of physical geography or of climate, which allows allied species hitherto
inhabiting distinct areas to come into contact, will often lead to the
extermination of one of them; and this extermination will be effected by no
external force, by no actual enemy, but merely because the one is slightly
better {46}adapted to live, to increase, and to maintain itself under
adverse circumstances, than the other.
Now if we consider carefully the few suggestive facts here referred to (and
many others of like import are to be found in Mr. Darwin's various works),
we shall be led to conclude that the several species, genera, families, and
orders, both of animals and vegetables which inhabit any extensive region,
are bound together by a series of complex relations; so that the increase,
diminution, or extermination of any one, may set in motion a series of
actions and reactions more or less affecting a large portion of the whole,
and requiring perhaps centuries of fluctuation before the balance is
restored. The range of any species or group in such a region, will in many
cases (perhaps in most) be determined, not by physical barriers, but by the
competition of other organisms. Where barriers have existed from a remote
epoch, they will at first have kept back certain animals from coming in
contact with each other; but when the assemblage of organisms on the two
sides of the barrier have, after many ages, come to form a balanced organic
whole, the destruction of the barrier may lead to a very partial
intermingling of the peculiar forms of the two regions. Each will have
become modified in special ways adapted to the organic and physical
conditions of the country, and will form a living barrier to the entrance
of animals less perfectly adapted to those conditions. Thus while the
abolition of ancient barriers will always lead to much intermixture of
forms, much extermination and wide-spread alteration in some families of
animals; other important groups will be unable materially to alter their
range; or they may make temporary incursions into the new territory, and be
ultimately driven back to very near their ancient limits.
In order to make this somewhat difficult subject more intelligible, it may
be well to consider the probable effects of certain hypothetical conditions
of the earth's surface:--
1. If the dry land of the globe had been from the first continuous, and
nowhere divided up by such boundaries as lofty mountain ranges, wide
deserts, or arms of the sea, it seems probable that none of the larger
groups (as _orders_, _tribes_, or {47}_families_,) would have a limited
range; but, as is to some extent the case in tropical America east of the
Andes, every such group would be represented over the whole area, by
countless minute modifications of form adapted to local conditions.
2. One great physical barrier would, however, even then exist; the hot
equatorial zone would divide the faunas and floras of the colder regions of
the northern and southern hemispheres from any chance of intermixture. This
one barrier would be more effectual than it is now, since there would be no
lofty mountain ranges to serve as a bridge for the partial interchange of
northern and southern forms.
3. If such a condition of the earth as here supposed continued for very
long periods, we may conceive that the action and reaction of the various
organisms on each other, combined with the influence of very slowly
changing physical conditions, would result in an almost perfect organic
balance, which would be manifested by a great stability in the average
numbers, the local range, and the peculiar characteristics of every
species.
4. Under such a condition of things it is not improbable that the total
number of clearly differentiated specific forms might be much greater than
it is now, though the number of generic and family types might perhaps be
less; for dominant species would have had ample time to spread into every
locality where they could exist, and would then become everywhere modified
into forms best suited to the permanent local conditions.
5. Now let us consider what would be the probable effect of the
introduction of a barrier, cutting off a portion of this homogeneous and
well-balanced world. Suppose, for instance, that a subsidence took place,
cutting off by a wide arm of the sea a large and tolerably varied island.
The first and most obvious result would be that the individuals of a number
of species would be divided into two portions, while others, the limits of
whose range agreed approximately with the line of subsidence, would exist
in unimpaired numbers on the new island or on the main land. But the
species whose numbers were diminished and whose original area was also
absolutely diminished by the portion now under the sea, would not be able
to hold their {48}ground against the rival forms whose numbers were intact.
Some would probably diminish and rapidly die out; others which produced
favourable varieties, might be so modified by natural selection as to
maintain their existence under a different form; and such changes would
take place in varying modes on the two sides of the new strait.
6. But the progress of these changes would necessarily affect the other
species in contact with them. New places would be opened in the economy of
nature which many would struggle to obtain; and modification would go on in
ever-widening circles and very long periods of time might be required to
bring the whole again into a state of equilibrium.
7. A new set of factors would in the meantime have come into play. The
sinking of land and the influx of a large body of water could hardly take
place without producing important climatal changes. The temperature, the
winds, the rains, might all be affected, and more or less changed in
duration and amount. This would lead to a quite distinct movement in the
organic world. Vegetation would certainly be considerably affected, and
through this the insect tribes. We have seen how closely the life of the
higher animals is often bound up with that of insects; and thus a set of
changes might arise that would modify the numerical proportions, and even
the forms and habits of a great number of species, would completely
exterminate some, and raise others from a subordinate to a dominant
position. And all these changes would occur differently on opposite sides
of the strait, since the insular climate could not fail to differ
considerably from that of the continent.
8. But the two sets of changes, as above indicated, produced by different
modes of action of the same primary cause, would act and react on each
other; and thus lead to such a far-spreading disturbance of the organic
equilibrium as ultimately perhaps to affect in one way or another, every
form of life upon the earth.
This hypothetical case is useful as enabling us better to realize how
wide-spreading might be the effects of one of the simplest changes of
physical geography, upon a compact mass of mutually {49}adapted organisms.
In the actual state of things, the physical changes that occur and have
occurred through all geological epochs are larger and more varied. Almost
every mile of land surface has been again and again depressed beneath the
ocean; most of the great mountain chains have either originated or greatly
increased in height during the Tertiary period; marvellous alterations of
climate and vegetation have taken place over half the land-surface of the
earth; and all these vast changes have influenced a globe so cut up by seas
and oceans, by deserts and snow-clad mountains, that in many of its more
isolated land-masses ancient forms of life have been preserved, which, in
the more extensive and more varied continents have long given way to higher
types. How complex then must have been the actions and reactions such a
state of things would bring about; and how impossible must it be for us to
guess, in most cases, at the exact nature of the forces that limit the
range of some species and cause others to be rare or to become extinct! All
that we can in general hope to do is, to trace out, more or less
hypothetically, some of the larger changes in physical geography that have
occurred during the ages immediately preceeding our own, and to estimate
the effect they will probably have produced on animal distribution. We may
then, by the aid of such knowledge as to past organic mutations as the
geological record supplies us with, be able to determine the probable
birthplace and subsequent migrations of the more important genera and
families; and thus obtain some conception of that grand series of
co-ordinated changes in the earth and its inhabitants, whose final result
is seen in the forms and the geographical distribution of existing animals.
{50}CHAPTER IV.
ON ZOOLOGICAL REGIONS.
To the older school of Naturalists the native country of an animal was of
little importance, except in as far as climates differed. Animals were
supposed to be specially adapted to live in certain zones or under certain
physical conditions, and it was hardly recognised that apart from these
conditions there was any influence in locality which could materially
affect them. It was believed that, while the animals of tropical,
temperate, and arctic climates, essentially differed; those of the tropics
were essentially alike all over the world. A group of animals was said to
inhabit the "Indies;" and important differences of structure were often
overlooked from the idea, that creatures equally adapted to live in hot
countries and with certain general resemblances, would naturally be related
to each other. Thus the Toucans and Hornbills, the Humming-Birds and
Sun-Birds, and even the Tapirs and the Elephants, came to be popularly
associated as slightly modified varieties of tropical forms of life; while
to naturalists, who were acquainted with the essential differences of
structure, it was a never-failing source of surprise, that under climates
and conditions so apparently identical, such strangely divergent forms
should be produced.
To the modern naturalist, on the other hand, the native country (or
"habitat" as it is technically termed) of an animal {51}or a group of
animals, is a matter of the first importance; and, as regards the general
history of life upon the globe, may be considered to be one of its
essential characters. The structure, affinities, and habits of a species,
now form only a part of its natural history. We require also to know its
exact range at the present day and in prehistoric times, and to have some
knowledge of its geological age, the place of its earliest appearance on
the globe, and of the various extinct forms most nearly allied to it. To
those who accept the theory of development as worked out by Mr. Darwin, and
the views as to the general permanence and immense antiquity of the great
continents and oceans so ably developed by Sir Charles Lyell, it ceases to
be a matter of surprise that the tropics of Africa, Asia, and America
should differ in their productions, but rather that they should have
anything in common. Their similarity, not their diversity, is the fact that
most frequently puzzles us.
The more accurate knowledge we have of late years obtained of the
productions of many remote regions, combined with the greater approaches
that have been made to a natural classification of the higher animals, has
shown, that every continent or well-marked division of a continent, every
archipelago and even every island, presents problems of more or less
complexity to the student of the geographical distribution of animals. If
we take up the subject from the zoological side, and study any family,
order, or even extensive genus, we are almost sure to meet with some
anomalies either in the present or past distribution of the various forms.
Let us adduce a few examples of these problems.
Deer have a wonderfully wide range, over the whole of Europe, Asia, and
North and South America; yet in Africa south of the great desert there are
none. Bears range over the whole of Europe, Asia, and North America, and
true pigs of the genus Sus, over all Europe and Asia and as far as New
Guinea; yet both bears and pigs, like deer, are absent from Tropical and
South Africa.
Again, the West Indian islands possess very few Mammalia, all of small size
and allied to those of America, except one {52}genus; and that belongs to
an Order, "Insectivora," entirely absent from South America, and to a
family, "Centetidæ," all the other species of which inhabit Madagascar
only. And as if to add force to this singular correspondence we have one
Madagascar species of a beautiful day-flying Moth, _Urania_, all the other
species of which inhabit tropical America. These insects are gorgeously
arrayed in green and gold, and are quite unlike any other Lepidoptera upon
the globe.
The island of Ceylon generally agrees in its productions with the Southern
part of India; yet it has several birds which are allied to Malayan and not
to Indian groups, and a fine butterfly of the genus _Hestia_, as well as
several genera of beetles, which are purely Malayan.
Various important groups of animals are distributed in a way not easy to
explain. The anthropoid apes in West Africa and Borneo; the tapirs in
Malaya and South America; the camel tribe in the deserts of Asia and the
Andes; the trogons in South America and Tropical Asia, with one species in
Africa; the marsupials in Australia and America, are examples.
The cases here adduced (and they might be greatly multiplied) are merely to
show the kind of problems with which the naturalist now has to deal; and in
order to do so he requires some system of geographical arrangement, which
shall serve the double purpose of affording a convenient subdivision of his
subject, and at the same time of giving expression to the main results at
which he has arrived. Hence the recent discussions on "Zoological Regions,"
or, what are the most natural primary divisions of the earth as regards its
forms of animal life.
The divisions in use till quite recently were of two kinds; either those
ready made by geographers, more especially the quarters or continents of
the globe; or those determined by climate and marked out by certain
parallels of latitude or by isothermal lines. Either of these methods was
better than none at all; but from the various considerations explained in
the preceding chapters, it will be evident, that such divisions must have
often been very unnatural, and have disguised many {53}of the most
important and interesting phenomena which a study of the distribution of
animals presents to us.
The merit of initiating a more natural system, that of determining
zoological regions, not by any arbitrary or _a priori_ consideration but by
studying the actual ranges of the more important groups of animals, is due
to Mr. Sclater, who, in 1857, established six primary zoological regions
from a detailed examination of the distribution of the chief genera and
families of Birds. Before stating what these regions are, what objections
have been made to them, what other divisions have been since proposed, and
what are those which we shall adopt in this work, it will be well to
consider the general principles which should guide us in the choice between
rival systems.
_Principles on which Zoological Regions should be formed._--It will be
evident in the first place that nothing like a perfect zoological division
of the earth is possible. The causes that have led to the present
distribution of animal life are so varied, their action and reaction have
been so complex, that anomalies and irregularities are sure to exist which
will mar the symmetry of any rigid system. On two main points every system
yet proposed, or that probably can be proposed, is open to objection; they
are,--1stly, that the several regions are not of equal rank;--2ndly, that
they are not equally applicable to all classes of animals. As to the first
objection, it will be found impossible to form any three or more regions,
each of which differs from the rest in an equal degree or in the same
manner. One will surpass all others in the possession of peculiar families;
another will have many characteristic genera; while a third will be mainly
distinguished by negative characters. There will also be found many
intermediate districts, which possess some of the characteristics of two
well-marked regions, with a few special features of their own, or perhaps
with none; and it will be a difficult question to decide in all cases which
region should possess this doubtful territory, or whether it should be
formed into a primary region itself. Again, two regions which have now
well-marked points of difference, may be shown to have been much more alike
at a comparatively recent geological epoch; {54}and this, it may be said,
proves their fundamental unity and that they ought to form but one primary
region. To obviate some of these difficulties a binary or dichotomous
division is sometimes proposed; that portion of the earth which differs
most from the rest being cut off as a region equal in rank to all that
remains, which is subjected again and again to the same process.
To decide these various points it seems advisable that convenience,
intelligibility, and custom, should largely guide us. The first essential
is, a broadly marked and easily remembered set of regions; which
correspond, as nearly as truth to nature will allow, with the distribution
of the most important groups of animals. What these groups are we shall
presently explain. In determining the number, extent, and boundaries of
these regions, we must be guided by a variety of indications, since the
application of fixed rules is impossible. They should evidently be of a
moderate number, corresponding as far as practicable with the great natural
divisions of the globe marked out by nature, and which have always been
recognized by geographers. There should be some approximation to equality
of size, since there is reason to believe that a tolerably extensive area
has been an essential condition for the development of most animal forms;
and it is found that, other things being equal, the numbers, variety and
importance of the forms of animal and vegetable life, do bear some
approximate relation to extent of area. Although the possession of peculiar
families or genera is the main character of a primary zoological region,
yet the negative character of the absence of certain families or genera is
of equal importance, _when this absence does not manifestly depend on
unsuitability to the support of the group_, and especially _when there is
now no physical barrier preventing their entrance_. This will become
evident when we consider that the importance of the possession of a group
by one region depends on its absence from the adjoining regions; and if
there is now no barrier to its entrance, we may be sure that there has once
been one; and that the possession of the area by a distinct and well
balanced set of organisms, which must have been slowly {55}developed and
adjusted, is the living barrier that now keeps out intruders.
When it is ascertained that the chief differences which now obtain between
two areas did not exist in Miocene or Pliocene times, the fact is one of
great interest, and enables us to speculate with some degree of probability
as to the causes that have brought about the present state of things; but
it is not a reason for uniting these two areas into one region. Our object
is to represent as nearly as possible the main features of the distribution
of existing animals, not those of any or all past geological epochs. Should
we ever obtain sufficient information as to the geography and biology of
the earth at past epochs, we might indeed determine approximately what were
the Pliocene or Miocene or Eocene zoological regions; but any attempt to
exhibit all these in combination with those of our own period, must lead to
confusion.
The binary or dichotomous system, although it brings out the fundamental
differences of the respective regions, is an inconvenient one in its
application, and rather increases than obviates the difficulty as to
equality or inequality of regions; for although _a_, _b_, _c_, and _d_, may
be areas of unequal zoological rank, _a_ being the most important, and _d_
the least, yet this inequality will probably be still greater if we first
divide them into _a_, on one side, and _b_, _c_, and _d_, on the other, and
then, by another division, make _b_, an area of the second, and _c_, and
_d_, of the third rank only.
Coming to the second objection, the often incompatible distribution of
different groups of animals, affords ground for opposition to any proposed
scheme of zoological regions. There is first the radical difference between
land and sea animals; the most complete barriers to the dispersal of the
one, sometimes offering the greatest facilities for the emigration of the
other, and _vice versa_. A large number of marine animals, however,
frequent shallow water only; and these, keeping near the coasts, will agree
generally in their distribution with those inhabiting the land. But among
land animals themselves there are very great differences of distribution,
due to certain specialities {56}in their organization or mode of life.
These act mainly in two ways,--1stly, by affecting the facilities with
which they can be dispersed, either voluntarily or involuntarily;--2ndly,
by the conditions which enable them to multiply and establish themselves in
certain areas and not in others. When both these means of diffusion are at
a maximum, the dispersal of a group becomes universal, and ceases to have
much interest for us. This is the case with certain groups of fungi and
lichens, as well as with some of the lower animals; and in a less degree,
as has been shown by Mr. Darwin, with many fresh-water plants and animals.
At the other extreme we may place certain arboreal vertebrata such as
sloths and lemurs, which have no means of passing such barriers as narrow
straits or moderately high mountains, and whose survival in any new country
they might reach, would be dependent on the presence of suitable forests
and the absence of dangerous enemies. Almost equally, or perhaps even more
restricted, are the means of permanent diffusion of terrestrial molluscs;
since these are without any but very rare and accidental means of being
safely transported across the sea; their individual powers of locomotion
are highly restricted; they are especially subject to the attacks of
enemies; and they often depend not only on a peculiar vegetation, but on
the geological character of the country, their abundance being almost in
direct proportion to the presence of some form of calcareous rocks. Between
these extremes we find animals possessed of an infinite gradation of powers
to disperse and to maintain themselves; and it will evidently be impossible
that the limits which best define the distribution of one group, should be
equally true for all others.
_Which class of Animals is of most importance in determining Zoological
Regions._--To decide this question we have to consider which groups of
animals are best adapted to exhibit, by their existing distribution, the
past changes and present physical condition of the earth's surface; and at
the same time, by the abundance of their remains in the various tertiary
formations will best enable us to trace out the more recent of the series
of changes, both of the earth's surface and {57}of its inhabitants, by
which the present state of things has been brought about. For this purpose
we require a group which shall be dependent for its means of dispersal on
the distribution of land and water, and on the presence or absence of lofty
mountains, desert plains or plateaux, and great forests; since these are
the chief physical features of the earth's surface whose modifications at
successive periods we wish to discover. It is also essential that they
should not be subject to dispersal by many accidental causes; as this would
inevitably in time tend to obliterate the effect of natural barriers, and
produce a scattered distribution, the causes of which we could only guess
at. Again, it is necessary that they should be so highly organized as not
to be absolutely dependent on other groups of animals, and with so much
power of adaptation as to be able to exist in one form or another over the
whole globe. And lastly, it is highly important that the whole group should
be pretty well known, and that a fairly natural classification, especially
of its minor divisions such as families and genera, should have been
arrived at; the reason for which last proviso is explained in our next
chapter, on classification.
Now in every one of these points the mammalia are preeminent; and they
possess the additional advantage of being the most highly developed class
of organized beings, and that to which we ourselves belong. We should
therefore construct our typical or standard Zoological Regions in the first
place, from a consideration of the distribution of mammalia, only bringing
to our aid the distribution of other groups to determine doubtful points.
Regions so established will be most closely in accordance with those
long-enduring features of physical geography, on which the distribution of
all forms of life fundamentally depend; and all discrepancies in the
distribution of other classes of animals must be capable of being
explained, either by their exceptional means of dispersion or by special
conditions affecting their perpetuation and increase in each locality.
If these considerations are well founded, the objections of those who study
insects or molluscs, for example,--that our regions are not true for their
departments of nature--cannot be {58}maintained. For they will find, that a
careful consideration of the exceptional means of dispersal and conditions
of existence of each group, will explain most of the divergences from the
normal distribution of higher animals.
We shall thus be led to an intelligent comprehension of the phenomena of
distribution in all groups, which would not be the case if every specialist
formed regions for his own particular study. In many cases we should find
that no satisfactory division of the earth could be made to correspond with
the distribution even of an entire class; but we should have the
coleopterist and the lepidopterist each with his own Geography. And even
this would probably not suffice, for it is very doubtful if the detailed
distribution of the Longicornes, so closely dependent on woody vegetation,
could be made to agree with that of the Staphylinidæ or the Carabidæ which
abound in many of the most barren regions, or with that of the Scarabeidæ,
largely dependent on the presence of herbivorous mammalia. And when each of
these enquirers had settled a division of the earth into "regions" which
exhibited with tolerable accuracy the phenomena of distribution of his own
group, we should have gained nothing whatever but a very complex mode of
exhibiting the bare facts of distribution. We should then have to begin to
work out the causes of the divergence of one group from another in this
respect; but as each worker would refer to his own set of regions as the
type, the whole subject would become involved in inextricable confusion.
These considerations seem to make it imperative that one set of "regions"
should be established as typical for Zoology; and it is hoped the reasons
here advanced will satisfy most naturalists that these regions can be best
determined, in the first place, by a study of the distribution of the
mammalia, supplemented in doubtful cases by that of the other vertebrates.
We will now proceed to a discussion of what these regions are.
_Various Zoological Regions proposed since 1857._--It has already been
pointed out that a very large number of birds are limited by the same kind
of barriers as mammalia; it will therefore not be surprising that a system
of regions formed to suit the {59}one, should very nearly represent the
distribution of the other. Mr. Sclater's regions are as follows:--
1. The Palæarctic Region; including Europe, Temperate Asia, and N. Africa
to the Atlas mountains.
2. The Ethiopian Region; Africa south of the Atlas, Madagascar, and the
Mascarene Islands, with Southern Arabia.
3. The Indian Region; including India south of the Himalayas, to South
China, and to Borneo and Java.
4. The Australian Region; including Celebes and Lombock, eastward to
Australia and the Pacific Islands.
5. The Nearctic Region; including Greenland, and N. America, to Northern
Mexico.
6. The Neotropical Region; including South America, the Antilles, and
Southern Mexico.
This division of the earth received great support from Dr. Günther, who, in
the _Proceedings of the Zoological Society_ for 1858, showed that the
geographical distribution of Reptiles agreed with it very closely, the
principal difference being that the reptiles of Japan have a more Indian
character than the birds, this being especially the case with the snakes.
In the volume for 1868 of the same work, Professor Huxley discusses at
considerable length the primary and secondary zoological divisions of the
earth. He gives reasons for thinking that the most radical primary
division, both as regards birds and mammals, is into a Northern and
Southern hemisphere (Arctogæa and Notogæa), the former, however, embracing
all Africa, while the latter includes only Australasia and the Neotropical
or Austro-Columbian region. Mr. Sclater had grouped his regions primarily
into Palæogæa and Neogæa, the Old and New Worlds of geographers; a division
which strikingly accords with the distribution of the passerine birds, but
not so well with that of mammalia or reptiles. Professor Huxley points out
that the Nearctic, Palæarctic, Indian, and Ethiopian regions of Mr. Sclater
have a much greater resemblance to each other than any one of them has to
Australia or to South America; and he further suggests that New Zealand
alone has peculiarities which might entitle it to rank as a primary region
{60}along with Australasia and South America; and that a Circumpolar
Province might be conveniently recognised as of equal rank with the
Palæarctic and Nearctic provinces.
In 1866, Mr. Andrew Murray published a large and copiously illustrated
volume on the _Geographical Distribution of Mammals_, in which he maintains
that the great and primary mammalian regions are only four: 1st. The
Palæarctic region of Mr. Sclater, extended to include the Sahara and Nubia;
2nd. the Indo-African region, including the Indian and Ethiopian regions of
Mr. Sclater; 3rd. the Australian region (unaltered); 4th. the American
region, including both North and South America. These are the regions as
_described_ by Mr. Murray, but his coloured map of "Great Mammalian
Regions" shows all Arctic America to a little south of the Isothermal of
32° Fahr. as forming with Europe and North Asia one great region.
At the meeting of the British Association at Exeter in 1869, Mr. W. T.
Blanford read a paper on the Fauna of British India, in which he maintained
that a large portion of the peninsula of India had derived its Fauna mainly
from Africa; and that the term "Indian region" of Mr. Sclater was
misleading, because India proper, if it belongs to it at all, is the least
typical portion of it. He therefore proposes to call it the "Malayan
region," because in the Malay countries it is most highly developed. Ceylon
and the mountain ranges of Southern India have marked Malay affinities.
In 1871 Mr. E. Blyth published in _Nature_ "A suggested new Division of the
Earth into Zoological Regions," in which he indicates seven primary
divisions or regions, subdivided into twenty-six sub-regions. The seven
regions are defined as follows: 1. The Boreal region; including the whole
of the Palæarctic and Nearctic regions of Mr. Sclater along with the West
Indies, Central America, the whole chain of the Andes, with Chili and
Patagonia. 2. The Columbian region; consisting of the remaining part of
South America. 3. The Ethiopian region; comprising besides that region of
Mr. Sclater, the valley of the Jordan, Arabia, and the desert country
towards India, with all the plains and table lands of India and the
northern {61}half of Ceylon. 4. The Lemurian region; consisting of
Madagascar and its adjacent islands. 5. The Austral-Asian region; which is
the Indian region of Mr. Sclater without the portion taken to be added to
the Ethiopian region. 6. The Melanesian region; which is the Australian
region of Mr. Sclater without New Zealand and the Pacific Islands, which
form 7. the Polynesian region. Mr. Blyth thinks this is "a true
classification of zoological regions as regards mammalia and birds."
In an elaborate paper on the birds of Eastern North America, their
distribution and migrations (_Bulletin of Museum of Comparative Zoology,
Cambridge, Massachusetts_, Vol. 2), Mr. J. A. Allen proposes a division of
the earth in accordance with what he terms, "the law of circumpolar
distribution of life in zones," as follows: 1. Arctic realm. 2. North
temperate realm. 3. American tropical realm. 4. Indo-African tropical
realm. 5. South American tropical realm. 6. African temperate realm. 7.
Antarctic realm. 8. Australian realm. Some of these are subdivided into
regions; (2) consisting of the American and the Europæo-Asiatic regions;
(4) into the African and Indian regions; (8) into the tropical Australian
region, and one comprising the southern part of Australia and New Zealand.
The other realms each form a single region.
_Discussion of proposed Regions._--Before proceeding to define the regions
adopted in this work, it may be as well to make a few remarks on some of
the preceding classifications, and to give the reasons which seem to render
it advisable to adopt very few of the suggested improvements on Mr.
Sclater's original proposal. Mr. Blyth's scheme is one of the least
natural, and also the most inconvenient. There can be little use in the
knowledge that a group of animals is found in the Boreal Region, if their
habitat might still be either Patagonia, the West Indies, or Japan; and it
is difficult to see on what principle the Madagascar group of islands is
made of equal rank with this enormous region, seeing that its forms of life
have marked African affinities. Neither does it seem advisable to adopt the
Polynesian Region, or that comprising New Zealand alone (as hinted at by
Professor Huxley and since adopted by {62}Mr. Sclater in his Lectures on
Geographical Distribution at the Zoological Gardens in May 1874), because
it is absolutely without indigenous mammalia and very poor in all forms of
life, and therefore by no means prominent or important enough to form a
primary region of the earth.
It may be as well here to notice what appears to be a serious objection to
making New Zealand, or any similar isolated district, one of the great
zoological regions, comparable to South America, Australia, or Ethiopia;
which is, that its claim to that distinction rests on grounds which are
liable to fail. It is because New Zealand, in addition to its negative
merits, possesses three families of birds (Apterygidæ living, Dinornithidæ
and Palapterygidæ extinct), and a peculiar lizard-like reptile, _Hatteria_,
which has to be classed in a distinct order, Rhynchocephalina, that the
rank of a Region is claimed for it. But supposing, what is not at all
improbable, that other Rhynchocephalina should be discovered in the
interior of Australia or in New Guinea, and that Apterygidæ or
Palapterygidæ should be found to have inhabited Australia in Post-Pliocene
times, (as Dinornithidæ have already been proved to have done) the claims
of New Zealand would entirely fail, and it would be universally
acknowledged to be a part of the great Australian region. No such reversal
can take place in the case of the other regions; because they rest, not
upon one or two, but upon a large number of peculiarities, of such a nature
that there is no room upon the globe for discoveries that can seriously
modify them. Even if one or two peculiar types, like Apterygidæ or
_Hatteria_, should permanently remain characteristic of New Zealand alone,
we can account for these by the extreme isolation of the country, and the
absence of enemies, which have enabled these defenceless birds and reptiles
to continue their existence; just as the isolation and protection of the
caverns of Carniola have enabled the _Proteus_ to survive in Europe. But
supposing that the _Proteus_ was the sole representative of an order of
Batrachia, and that two or three other equally curious and isolated forms
occurred with it, no one would propose that these caverns or the district
containing them, should form one of the {63}primary divisions of the earth.
Neither can much stress be laid on the negative characteristics of New
Zealand, since they are found to an almost equal extent in every oceanic
island.
Again, it is both inconvenient and misleading to pick out certain tracts
from the midst of one region or sub-region and to place them in another, on
account of certain isolated affinities which may often be accounted for by
local peculiarities. Even if the resemblance of the fauna of Chili and
Patagonia to that of the Palæarctic and Nearctic regions was much greater
than it is, this mode of dealing with it would be objectionable; but it is
still more so, when we find that these countries have a strongly marked
South American character, and that the northern affinities are altogether
exceptional. The Rodentia, which comprise a large portion of the mammalia
of these countries, are wholly South American in type, and the birds are
almost all allied to forms characteristic of tropical America.
For analogous reasons the Ethiopian must not be made to include any part of
India or Ceylon; for although the Fauna of Central India has some African
affinities, these do not preponderate; and it will not be difficult to show
that to follow Mr. Andrew Murray in uniting bodily the Ethiopian and Indian
regions of Mr. Sclater, is both unnatural and inconvenient. The
resemblances between them are of the same character as those which would
unite them both with the Palæarctic and Nearctic regions; and although it
may be admitted, that, as Professor Huxley maintains, this group forms one
of the great primary divisions of the globe, it is far too extensive and
too heterogeneous to subserve the practical uses for which we require a
division of the world into zoological regions.
_Reasons for adopting the six Regions first proposed by Mr. Sclater._--So
that we do not violate any clear affinities or produce any glaring
irregularities, it is a positive, and by no means an unimportant, advantage
to have our named regions approximately equal in size, and with easily
defined, and therefore easily remembered, boundaries. All elaborate
definitions of interpenetrating frontiers, as well as regions extending
over three-fourths of the land surface of the globe, and including places
which are {64}the antipodes of each other, would be most inconvenient, even
if there were not such difference of opinion about them. There can be
little doubt, for example, that the most radical zoological division of the
earth is made by separating the Australian region from the rest; but
although it is something useful and definite to know that a group of
animals is peculiar to Australia, it is exceedingly vague and
unsatisfactory to say of any other group merely that it is
extra-Australian. Neither can it be said that, from any point of view,
these two divisions are of equal importance. The next great natural
division that can be made is the separation of the Neotropical Region of
Mr. Sclater from the rest of the world. We thus have three primary
divisions, which Professor Huxley seems inclined to consider as of
tolerably equal zoological importance. But a consideration of all the
facts, zoological and palæontological, indicates, that the great northern
division (Arctogæa) is fully as much more important than either Australia
or South America, as its four component parts are less important; and if
so, convenience requires us to adopt the smaller rather than the larger
divisions.
This question, of comparative importance or equivalence of value, is very
difficult to determine. It may be considered from the point of view of
speciality or isolation, or from that of richness and variety of animal
forms. In isolation and speciality, determined by what they want as well as
what they possess, the Australian and Neotropical regions are undoubtedly
each comparable with the rest of the earth (Arctogæa). But in richness and
variety of forms, they are both very much inferior, and are much more
nearly comparable with the separate regions which compose it. Taking the
families of mammalia as established by the best authors, and leaving out
the Cetacea and the Bats, which are almost universally distributed, and
about whose classification there is much uncertainty, the number of
families represented in each of Mr. Sclater's regions is as follows:
I. Palæarctic region has 31 families of terrestrial mammalia.
II. Ethiopian " " 40 " " "
III. Indian " " 31 " " "
IV. Australian " " 14 " " "
V. Neotropical " " 26 " " "
VI. Nearctic " " 23 " " "
{65}We see, then, that even the exceedingly rich and isolated Neotropical
region is less rich and diversified in its forms of mammalian life than the
very much smaller area of the Indian region, or the temperate Palæarctic,
and very much less so than the Ethiopian region; while even the
comparatively poor Nearctic region, is nearly equal to it in the number of
its family types. If these were united they would possess fifty-five
families, a number very disproportionate to those of the remaining two.
Another consideration is, that although the absence of certain forms of
life makes a region more isolated, it does not make it zoologically more
important; for we have only to suppose some five or six families, now
common to both, to become extinct either in the Ethiopian or the Indian
regions, and they would become as strongly differentiated from all other
regions as South America, while still remaining as rich in family types. In
birds exactly the same phenomenon recurs, the family types being less
numerous in South America than in either of the other tropical regions of
the earth, but a larger proportion of them are restricted to it. It will be
shown further on, that the Ethiopian and Indian, (or, as I propose to call
it in this work, Oriental) regions, are sufficiently differentiated by very
important groups of animals peculiar to each; and that, on strict
zoological principles they are entitled to rank as regions of equal value
with the Neotropical and Australian. It is perhaps less clear whether the
Palæarctic should be separated from the Oriental region, with which it has
undoubtedly much in common; but there are many and powerful reasons for
keeping it distinct. There is an unmistakably different facies in the
animal forms of the two regions; and although no families of mammalia or
birds, and not many genera, are wholly confined to the Palæarctic region, a
very considerable number of both have their metropolis in it, and are very
richly represented. The distinction between the characteristic forms of
life in tropical and cold countries is, on the whole, very strongly marked
in the northern hemisphere; and to refuse to recognise this in a
subdivision of the earth which is established for the very purpose of
expressing such contrasts more clearly and concisely than by ordinary
geographical terminology, would be both illogical and {66}inconvenient. The
one question then remains, whether the Nearctic region should be kept
separate, or whether it should form part of the Palæarctic or of the
Neotropical regions. Professor Huxley and Mr. Blyth advocate the former
course; Mr. Andrew Murray (for mammalia) and Professor Newton (for birds)
think the latter would be more natural. No doubt much is to be said for
both views, but both cannot be right; and it will be shown in the latter
part of this chapter that the Nearctic region is, on the whole, fully as
well defined as the Palæarctic, by positive characters which differentiate
it from both the adjacent regions. More evidence in the same direction will
be found in the Second Part of this work, in which the extinct faunas of
the several regions are discussed.
A confirmation of the general views here set forth, as to the distinctness
and approximate equivalence of the six regions, is to be found in the fact,
that if any two or more of them are combined they themselves become
divisions of the next lower rank, or "sub-regions;"--and these will be very
much more important, both zoologically and geographically, than the
subdivisions of the remaining regions. It is admitted then that these six
regions are by no means of precisely equal rank, and that some of them are
far more isolated and better characterized than others; but it is
maintained that, looked at from every point of view, they are more equal in
rank than any others that can be formed; while in geographical equality,
compactness of area, and facility of definition, they are beyond all
comparison better than any others that have yet been proposed for the
purpose of facilitating the study of geographical distribution. They may be
arranged and grouped as follows, so as to exhibit their various relations
and affinities.
Regions.
{ NEOTROPICAL Austral zone Notogæa.
Neogæa {
{ NEARCTIC } }
} Boreal zone }
{ PALÆARCTIC } }
{ } Arctogæa.
{ ETHIOPIAN } }
Palæogæa { } Palæotropical zone }
{ ORIENTAL } }
{
{ AUSTRALIAN Austral zone Notogæa.
The above table shows the regions placed in the order followed in the
Fourth Part of this work, and the reasons for which are {67}explained in
Chapter IX. As a matter of convenience, and for other reasons adduced in
the same chapter, the detailed exposition of the geographical distribution
of the animals of the several regions in Part III. commences with the
Palæarctic and terminates with the Nearctic region.
_Objections to the system of Circumpolar Zones._--Mr. Allen's system of
"realms" founded on climatic zones (given at p. 61), having recently
appeared in an ornithological work of considerable detail and research,
calls for a few remarks. The author continually refers to the "_law of the
distribution of life in circumpolar zones_," as if it were one generally
accepted and that admits of no dispute. But this supposed "law" only
applies to the smallest details of distribution--to the range and
increasing or decreasing numbers of _species_ as we pass from north to
south, or the reverse; while it has little bearing on the great features of
zoological geography--the limitation of groups of _genera_ and _families_
to certain areas. It is analogous to the "_law of adaptation_" in the
organisation of animals, by which members of various groups are suited for
an aerial, an aquatic, a desert, or an arboreal life; are herbivorous,
carnivorous, or insectivorous; are fitted to live underground, or in fresh
waters, or on polar ice. It was once thought that these adaptive
peculiarities were suitable foundations for a classification,--that whales
were fishes, and bats birds; and even to this day there are naturalists who
cannot recognise the essential diversity of structure in such groups as
swifts and swallows, sun-birds and humming-birds, under the superficial
disguise caused by adaptation to a similar mode of life. The application of
Mr. Allen's principle leads to equally erroneous results, as may be well
seen by considering his separation of "the southern third of Australia" to
unite it with New Zealand as one of his secondary zoological divisions. If
there is one country in the world whose fauna is strictly homogeneous, that
country is Australia; while New Guinea on the one hand, and New Zealand on
the other, are as sharply differentiated from Australia as any adjacent
parts of the same primary zoological division can possibly be. Yet the
"_law of circumpolar distribution_" leads to the division of {68}Australia
by an arbitrary east and west line, and a union of the northern two-thirds
with New Guinea, the southern third with New Zealand. Hardly less unnatural
is the supposed equivalence of South Africa (the African temperate realm)
to all tropical Africa and Asia, including Madagascar (the Indo-African
tropical realm). South Africa has, it is true, some striking peculiarities;
but they are absolutely unimportant as compared with the great and radical
differences between tropical Africa and tropical Asia. On these examples we
may fairly rest our rejection of Mr. Allen's scheme.
We must however say a few words on the zoo-geographical nomenclature
proposed in the same paper, which seems also very objectionable. The
following terms are proposed: _realm_, _region_, _province_, _district_,
_fauna and flora_; the first being the highest, the last the lowest and
smallest sub-division. Considering that most of these terms have been used
in very different senses already, and that no means of settling their
equivalence in different parts of the globe has been even suggested, such a
complex system must lead to endless confusion. Until the whole subject is
far better known and its first principles agreed upon, the simpler and the
fewer the terms employed the better; and as "region" was employed for the
primary divisions by Mr. Sclater, eighteen years ago, and again by Mr.
Andrew Murray, in his Geographical Distribution of Mammals; nothing but
obscurity can result from each writer using some new, and doubtfully
better, term. For the sub-divisions of the regions no advantage is gained
by the use of a distinct term--"province"--which has been used (by
Swainson) for the primary divisions, and which does not itself tell you
what rank it holds; whereas the term "sub-region" speaks for itself as
being unmistakably next in subordination to region, and this clearness of
meaning gives it the preference over any independent term. As to minor
named sub-divisions, they seem at present uncalled for; and till the
greater divisions are themselves generally agreed on, it seems better to
adopt no technical names for what must, for a long time to come, be
indeterminate.
_Does the Arctic Fauna characterize an independent {69}Region._--The
proposal to consider the Arctic regions as constituting one of the primary
zoological divisions of the globe, has been advocated by many naturalists.
Professor Huxley seems to consider it advisable, and Mr. Allen
unhesitatingly adopts it, as well as an "antarctic" region to balance it in
the southern hemisphere. The reason why an "Arctic Region" finds no place
in this work may therefore be here stated.
No species or group of animals can properly be classed as "arctic," which
does not exclusively inhabit or greatly preponderate in arctic lands. For
the purpose of establishing the need of an "arctic" zoological region, we
should consider chiefly such groups as are circumpolar as well as arctic;
because, if they are confined to, or greatly preponderate in, either the
eastern or western hemispheres, they can be at once allocated to the
Nearctic or Palæarctic regions, and can therefore afford no justification
for establishing a new primary division of the globe.
Thus restricted, only three genera of land mammalia are truly arctic:
_Gulo_, _Myodes_, and _Rangifer_. Two species of widely dispersed genera
are also exclusively arctic, _Ursus maritimus_ and _Vulpes lagopus_.
Exclusively arctic birds are not much more numerous. Of land birds there
are only three genera (each consisting of but a single species),
_Pinicola_, _Nyctea_, and _Surnia_. _Lagopus_ is circumpolar, but the genus
has too wide an extension in the temperate zone to be considered arctic.
Among aquatic birds we have the genus of ducks, _Somateria_; three genera
of Uriidæ, _Uria_, _Catarractes_, and _Mergulus_; and the small family
Alcidæ, consisting of the genera _Alca_ and _Fratercula_. Our total then
is, three genera of mammalia, three of land, and six of aquatic birds,
including one peculiar family.
In the southern hemisphere there is only the single genus _Aptenodytes_
that can be classed as antarctic; and even that is more properly south
temperate.
In dealing with this arctic fauna we have two courses open to us; we must
either group them with the other species and genera which are common to the
two northern regions, or we {70}must form a separate primary region for
them. As a matter of convenience the former plan seems the best; and it is
that which is in accordance with our treatment of other intermediate tracts
which contain special forms of life. The great desert zone, extending from
the Atlantic shores of the Sahara across Arabia to Central Asia, is a
connecting link between the Palæarctic, Ethiopian, and Oriental regions,
and contains a number of "desert" forms wholly or almost wholly restricted
to it; but the attempt to define it as a separate region would introduce
difficulty and confusion. Neither to the "desert" nor to the "arctic"
regions could any defined limits, either geographical or zoological, be
placed; and the attempt to determine what species or genera should be
allotted to them would prove an insoluble problem. The reason perhaps is,
that both are essentially unstable, to a much greater extent than those
great masses of land with more or less defined barriers, which constitute
our six regions. The Arctic Zone has been, within a recent geological
period, both vastly more extensive and vastly less extensive than it is at
present. At a not distant epoch it extended over half of Europe and of
North America. At an earlier date it appears to have vanished altogether;
since a luxuriant vegetation of tall deciduous trees and broad-leaved
evergreens flourished within ten degrees of the Pole! The great deserts
have not improbably been equally fluctuating; hence neither the one nor the
other can present that marked individuality in their forms of life, which
seems to have arisen only when extensive tracts of land have retained some
considerable stability both of surface and climatal conditions, during
periods sufficient for the development and co-adaptation of their several
assemblages of plants and animals.
We must also consider that there is no geographical difficulty in dividing
the Arctic Zone between the two northern regions. The only debateable
lands, Greenland and Iceland, are generally admitted to belong respectively
to America and Europe. Neither is there any zoological difficulty; for the
land mammalia and birds are on the whole wonderfully restricted to their
respective regions even in high latitudes; and the aquatic forms {71}are,
for our present purpose, of much less importance. As a primary division the
"Arctic region" would be out of all proportion to the other six, whether as
regards its few peculiar types or the limited number of forms and species
actually inhabiting it; but it comes in well as a connecting link between
two regions, where the peculiar forms of both are specially modified; and
is in this respect quite analogous to the great desert zone above referred
to.
I now proceed to characterize briefly the six regions adopted in the
present work, together with the sub-regions into which they may be most
conveniently and naturally divided, as shown in our general map.
_Palæarctic Region._--This very extensive region comprises all temperate
Europe and Asia, from Iceland to Behring's Straits and from the Azores to
Japan. Its southern boundary is somewhat indefinite, but it seems advisable
to comprise in it all the extra-tropical part of the Sahara and Arabia, and
all Persia, Cabul, and Beloochistan to the Indus. It comes down to a little
below the upper limit of forests in the Himalayas, and includes the larger
northern half of China, not quite so far down the coast as Amoy. It has
been said that this region differs from the Oriental by negative characters
only; a host of tropical families and genera being absent, while there is
little or nothing but peculiar species to characterize it absolutely. This
however is not true. The Palæarctic region is well characterized by
possessing 3 families of vertebrata peculiar to it, as well as 35 peculiar
genera of mammalia, and 57 of birds, constituting about one-third of the
total number it possesses. These are amply sufficient to characterize a
region positively; but we must also consider the absence of many important
groups of the Oriental, Ethiopian, and Nearctic regions; and we shall then
find, that taking positive and negative characters together, and making
some allowance for the necessary poverty of a temperate as compared with
tropical regions, the Palæarctic is almost as strongly marked and well
defined as any other.
_Sub-divisions of the Palæarctic Region._--These are by no means {72}so
clearly indicated as in some of the other regions, and they are adopted
more for convenience than because they are very natural or strongly marked.
The first, or European sub-region, comprises Central and Northern Europe as
far South as the Pyrenees, the Maritime and Dinaric Alps, the Balkan
mountains, the Black Sea, and the Caucasus. On the east the Caspian sea and
the Ural mountains seem the most obvious limit; but it is doubtful if they
form the actual boundary, which is perhaps better marked by the valley of
the Irtish, where a pre-glacial sea almost certainly connected the Aral and
Caspian seas with the Arctic ocean, and formed an effective barrier which
must still, to some extent, influence the distribution of animals.
The next, or Mediterranean sub-region, comprises South Europe, North Africa
with the extra-tropical portion of the Sahara, and Egypt to about the first
or second cataracts; and eastward through Asia Minor, Persia, and Cabul, to
the deserts of the Indus.
The third, or Siberian sub-region, consists of all north and central Asia
north of Herat, as far as the eastern limits of the great desert plateau of
Mongolia, and southward to about the upper limit of trees on the Himalayas.
The fourth, or Manchurian sub-region, consists of Japan and North China
with the lower valley of the Amoor; and it should probably be extended
westward in a narrow strip along the Himalayas, embracing about 1,000 or
2,000 feet of vertical distance below the upper limit of trees, till it
meets an eastern extension of the Mediterranean sub-region a little beyond
Simla. These extensions are necessary to avoid passing from the Oriental
region, which is essentially tropical, directly to the Siberian sub-region,
which has an extreme northern character; whereas the Mediterranean and
Manchurian sub-regions are more temperate in climate. It will be found that
between the upper limit of most of the typical Oriental groups and the
Thibetan or Siberian fauna, there is a zone in which many forms occur
common to temperate China. This is especially the case among the pheasants
and finches.
{73}_Ethiopian Region._--The limits of this region have been indicated by
the definition of the Palæarctic region. Besides Africa south of the tropic
of Cancer, and its islands, it comprises the southern half of Arabia.
This region has been said to be identical in the main characters of its
mammalian fauna with the Oriental region, and has therefore been united
with it by Mr. A. Murray. Most important differences have however been
overlooked, as the following summary of the peculiarities of the Ethiopian
region will, I think, show.
It possesses 22 peculiar families of vertebrates; 90 peculiar genera of
mammalia, being two-thirds of its whole number; and 179 peculiar genera of
birds, being three-fifths of all it possesses. It is further characterized
by the absence of several families and genera which range over the whole
northern hemisphere, details of which will be found in the chapter treating
of the region. There are, it is true, many points of resemblance, not to be
wondered at between two tropical regions in the same hemisphere, and which
have evidently been at one time more nearly connected, both by intervening
lands and by a different condition of the lands that even now connect them.
But these resemblances only render the differences more remarkable; since
they show that there has been an ancient and long-continued separation of
the two regions, developing a distinct fauna in each, and establishing
marked specialities which the temporary intercommunication and immigration
has not sufficed to remove. The entire absence of such wide-spread groups
as bears and deer, from a country many parts of which are well adapted to
them, and in close proximity to regions where they abound, would alone mark
out the Ethiopian region as one of the primary divisions of the earth, even
if it possessed a less number than it actually does of peculiar family and
generic groups.
_Sub-divisions of the Ethiopian Region._--The African continent south of
the tropic of Cancer is more homogeneous in its prominent and superficial
zoological features than most of the other regions, but there are
nevertheless important and {74}deep-seated local peculiarities. Two
portions can be marked off as possessing many peculiar forms; the luxuriant
forest district of equatorial West Africa, and the southern extremity or
Cape district. The remaining portion has no well-marked divisions, and a
large proportion of its animal forms range over it from Nubia and
Abyssinia, to Senegal on the one side and to the Zambesi on the other; this
forms our first or East-African sub-region.
The second, or West African sub-region extends along the coast from Senegal
to Angola, and inland to the sources of the Shary and the Congo.
The third, or South African sub-region, comprises the Cape Colony and
Natal, and is roughly limited by a line from Delagoa Bay to Walvish Bay.
The fourth, or Malagasy sub-region, consists of Madagascar and the adjacent
islands, from Rodriguez to the Seychelles; and this differs so remarkably
from the continent that it has been proposed to form a distinct primary
region for its reception. Its productions are indeed highly interesting;
since it possesses 3 families, and 2 sub-families of mammals peculiar to
itself, while almost all its genera are peculiar. Of these a few show
Oriental or Ethiopian affinities, but the remainder are quite isolated.
Turning to other classes of animals, we find that the birds are almost as
remarkable; but, as might be expected, a larger number of genera are common
to surrounding countries. More than 30 genera are altogether peculiar, and
some of these are so isolated as to require to be classed in separate
families or sub-families. The African affinity is however here more
strongly shown by the considerable number (13) of peculiar Ethiopian genera
which in Madagascar have representative species. There can be no doubt
therefore about Madagascar being more nearly related to the Ethiopian than
to any other region; but its peculiarities are so great, that, were it not
for its small size and the limited extent of its fauna, its claim to rank
as a separate region might not seem unreasonable. It is true that it is not
poorer in mammals than Australia; but that country is far more isolated,
and cannot be so decidedly and {75}naturally associated with any other
region as Madagascar can be with the Ethiopian. It is therefore the better
and more natural course to keep it as a sub-region; the peculiarities it
exhibits being of exactly the same kind as those presented by the Antilles,
by New Zealand, and even by Celebes and Ceylon, but in a much greater
degree.
_Oriental Region._--On account of the numerous objections that have been
made to naming a region from the least characteristic portion of it, and
not thinking "Malayan," proposed by Mr. Blanford, a good term, (as it has a
very circumscribed and definite meaning, and especially because the "Malay"
archipelago is half of it in the Australian region,) I propose to use the
word "Oriental" instead of "Indian," as being geographically applicable to
the whole of the countries included in the region and to very few beyond
it; as being euphonious, and as being free from all confusion with terms
already used in zoological geography. I trust therefore that it may meet
with general acceptance.
This small, compact, but rich and varied region, consists of all India and
China from the limits of the Palæarctic region; all the Malay peninsula and
islands as far east as Java and Baly, Borneo and the Philippine Islands;
and Formosa. It is positively characterized by possessing 12 peculiar
families of vertebrata; by 55 genera of land mammalia, and 165 genera of
land birds, altogether confined to it; these peculiar genera forming in
each case about one half of the total number it possesses.
_Sub-divisions of the Oriental region._--First we have the Indian
sub-region, consisting of Central India from the foot of the Himalayas in
the west, and south of the Ganges to the east, as far as a line drawn from
Goa curving south and up to the Kistna river; this is the portion which has
most affinity with Africa.
The second, or Ceylonese sub-region, consists of the southern extremity of
India with Ceylon; this is a mountainous forest region, and possesses
several peculiar forms as well as some Malayan types not found in the first
sub-region.
{76}Next we have the Indo-Chinese sub-region, comprising South China and
Burmah, extending westward along the Himalayan range to an altitude of
about 9,000 or 10,000 feet, and southward to Tavoy or Tenasserim.
The last is the Indo-Malayan sub-region, comprising the Peninsula of
Malacca and the Malay Islands to Baly, Borneo, and the Philippines.
On account of the absence from the first sub-region of many of the forms
most characteristic of the other three, and the number of families and
genera of mammalia and birds which occur in it and also in Africa, it has
been thought by some naturalists that this part of India has at least an
equal claim to be classed as a part of the Ethiopian region. This question
will be found fully discussed in Chapter XII. devoted to the Oriental
region, where it is shown that the African affinity is far less than has
been represented, and that in all its essential features Central India is
wholly Oriental in its fauna.
Before leaving this region a few words may be said about Lemuria, a name
proposed by Mr. Sclater for the site of a supposed submerged continent
extending from Madagascar to Ceylon and Sumatra, in which the Lemuroid type
of animals was developed. This is undoubtedly a legitimate and highly
probable supposition, and it is an example of the way in which a study of
the geographical distribution of animals may enable us to reconstruct the
geography of a bygone age. But we must not, as Mr. Blyth proposed, make
this hypothetical land one of our actual Zoological regions. It represents
what was probably a primary Zoological region in some past geological
epoch; but what that epoch was and what were the limits of the region in
question, we are quite unable to say. If we are to suppose that it
comprised the whole area now inhabited by Lemuroid animals, we must make it
extend from West Africa to Burmah, South China, and Celebes; an area which
it possibly did once occupy, but which cannot be formed into a modern
Zoological region without violating much more important affinities. If, on
the other hand, we leave out all those areas which undoubtedly belong to
other regions, we reduce Lemuria to Madagascar and its adjacent
{77}islands, which, for reasons already stated, it is not advisable to
treat as a primary Zoological region. The theory of this ancient continent
and the light it may throw on existing anomalies of distribution, will be
more fully considered in the geographical part of this work.
_Australian Region._--Mr. Sclater's original name seems preferable to
Professor Huxley's, "Austral-Asian;" the inconvenience of which alteration
is sufficiently shown by the fact that Mr. Blyth proposed to use the very
same term as an appropriate substitute for the "Indian region" of Mr.
Sclater. Australia is the great central mass of the region; it is by far
the richest in varied and highly remarkable forms of life; and it therefore
seems in every way fitted to give a name to the region of which it is the
essential element. The limits of this region in the Pacific are somewhat
obscure, but as so many of the Pacific Islands are extremely poor
zoologically, this is not of great importance.
_Sub-divisions of the Australian Region._--The first sub-region is the
Austro-Malayan, including the islands from Celebes and Lombock on the west
to the Solomon Islands on the east. The Australian sub-region comes next,
consisting of Australia and Tasmania. The third, or Polynesian sub-region,
will consist of all the tropical Pacific Islands, and is characterized by
several peculiar genera of birds which are all allied to Australian types.
The fourth, consists of New Zealand with Auckland, Chatham, and Norfolk
Islands, and must be called the New Zealand sub-region.
The extreme peculiarities of New Zealand, due no doubt to its great
isolation and to its being the remains of a more extensive land, have
induced several naturalists to suggest that it ought justly to form a
Zoological region by itself. But the inconveniences of such a procedure
have been already pointed out; and when we look at its birds as a whole
(they being the only class sufficiently well represented to found any
conclusion upon) we find that the majority of them belong to Australian
genera, and where the genera are peculiar they are most nearly related to
Australian types. The preservation in these islands {78}of a single
representative of a unique order of reptiles, is, as before remarked, of
the same character as the preservation of the _Proteus_ in the caverns of
Carniola; and can give the locality where it happens to have survived no
claim to form a primary Zoological region, unless supported by a tolerably
varied and distinctly characterized fauna, such as never exists in a very
restricted and insular area.
_Neotropical Region._--Mr. Sclater's original name for this region is
preserved, because change of nomenclature is always an evil; and neither
Professor Huxley's suggested alteration "Austro-Columbia," nor Mr.
Sclater's new term "Dendrogæa," appear to be improvements. The region is
essentially a tropical one, and the extra-tropical portion of it is not
important enough to make the name inappropriate. That proposed by Professor
Huxley is not free from the same kind of criticism, since it would imply
that the region was exclusively South American, whereas a considerable
tract of North America belongs to it. This region includes South America,
the Antilles and tropical North America; and it possesses more peculiar
families of vertebrates and genera of birds and mammalia than any other
region.
_Subdivisions of the Neotropical Region._--The great central mass of South
America, from the shores of Venezuela to Paraguay and Eastern Peru,
constitutes the chief division, and may be termed the Brazilian sub-region.
It is on the whole a forest country; its most remarkable forms are highly
developed arboreal types; and it exhibits all the characteristics of this
rich and varied continent in their highest development.
The second, or Chilian sub-region, consists of the open plains, pampas, and
mountains of the southern extremity of the continent; and we must include
in it the west side of the Andes as far as the limits of the forest near
Payta, and the whole of the high Andean plateaus as far as 4° of south
latitude; which makes it coincide with the range of the Camelidæ and
Chinchillidæ.
The third, or Mexican sub-region, consists of Central America and Southern
Mexico, but it has no distinguishing {79}characteristics except the absence
of some of the more highly specialized Neotropical groups. It is, however,
a convenient division as comprising the portion of the North American
continent which belongs zoologically to South America.
The fourth, or Antillean sub-region, consists of the West India islands
(except Trinidad and Tobago, which are detached portions of the continent
and must be grouped in the first sub-region); and these reproduce, in a
much less marked degree, the phenomena presented by Madagascar. Terrestrial
mammals are almost entirely wanting, but the larger islands possess three
genera which are altogether peculiar to them. The birds are of South
American forms, but comprise many peculiar genera. Terrestrial molluscs are
more abundant and varied than in any part of the globe of equal extent; and
if these alone were considered, the Antilles would constitute an important
Zoological region.
_Nearctic Region._--This region comprises all temperate North America and
Greenland. The arctic lands and islands beyond the limit of trees form a
transitional territory to the Palæarctic region, but even here there are
some characteristic species. The southern limit between this region and the
Neotropical is a little uncertain; but it may be drawn at about the Rio
Grande del Norte on the east coast, and a little north of Mazatlan on the
west; while on the central plateau it descends much farther south, and
should perhaps include all the open highlands of Mexico and Guatemala. This
would coincide with the range of several characteristic Nearctic genera.
_Distinction of the Nearctic from the Palæarctic Region._--The Nearctic
region possesses twelve peculiar families of vertebrates or one-tenth of
its whole number. It has also twenty-four peculiar genera of mammalia and
fifty-two of birds, in each case nearly one-third of all it possesses. This
proportion is very nearly the same as in the Palæarctic region, while the
number of peculiar families of vertebrata is very much greater. It has been
already seen that both Mr. Blyth and Professor Huxley are disposed to unite
this region with the Palæarctic, while Professor Newton, in his article on
birds in the new edition of the {80}Encyclopædia Britannica, thinks that as
regards that class it can hardly claim to be more than a sub-region of the
Neotropical. These views are mutually destructive, but it will be shown in
the proper place, that on independent grounds the Nearctic region can very
properly be maintained.
_Subdivisions of the Nearctic Region._--The sub-regions here depend on the
great physical features of the country, and have been in some cases
accurately defined by American naturalists. First we have the Californian
sub-region, consisting of California and Oregon--a narrow tract between the
Sierra Nevada and the Pacific, but characterized by a number of peculiar
species and by several genera found nowhere else in the region.
The second, or Rocky Mountain sub-region, consists of this great mountain
range with its plateaus, and the central plains and prairies to about 100°
west longitude, but including New Mexico and Texas in the South.
The third and most important sub-region, which may be termed the
Alleghanian, extends eastward to the Atlantic, including the Mississippi
Valley, the Alleghany Mountains, and the Eastern United States. This is an
old forest district, and contains most of the characteristic animal types
of the region.
The fourth, or Canadian sub-region, comprises all the northern part of the
continent from the great lakes to the Arctic ocean; a land of pine-forests
and barren wastes, characterized by Arctic types and the absence of many of
the genera which distinguish the more southern portions of the region.
_Observations on the series of Sub-regions._--The twenty-four sub-regions
here adopted were arrived at by a careful consideration of the distribution
of the more important genera, and of the materials, both zoological and
geographical, available for their determination; and it was not till they
were almost finally decided on, that they were found to be equal in number
throughout all the regions--four in each. As this uniformity is of great
advantage in tabular and diagrammatic presentations of the distribution of
the several families, I decided not to disturb it unless very strong
reasons should appear for adopting a greater or less number in any
particular case. Such however have not {81}arisen; and it is hoped that
these divisions will prove as satisfactory and useful to naturalists in
general as they have been to the author. Of course, in a detailed study of
any region much more minute sub-division may be required; but even in that
case it is believed that the sub-regions here adopted, will be found, with
slight modifications, permanently available for exhibiting general results.
I give here a table showing the proportionate richness and speciality of
each region as determined by its _families_ of vertebrates and _genera_ of
mammalia and birds; and also a general table of the regions and
sub-regions, arranged in the order that seems best to show their mutual
relations.
COMPARATIVE RICHNESS OF THE SIX REGIONS.
+-------------+-----------------+
| | VERTEBRATA. |
| REGIONS. |Families|Peculiar|
| | |families|
+-------------+--------+--------+
| Palæarctic | 136 | 3 |
| Ethiopian | 174 | 22 |
| Oriental | 164 | 12 |
| Australian | 141 | 30 |
| Neotropical | 168 | 44 |
| Nearctic | 122 | 12 |
+-------------+--------+--------+
+-------------+-----------------------+-----------------------+
| | MAMMALIA. | BIRDS. |
| REGIONS. |Genera|Peculiar| Per |Genera|Peculiar| Per |
| | |genera |centage| |genera |centage|
+-------------+------+--------+-------+------+--------+-------+
| Palæarctic | 100 | 35 | 35 | 174 | 57 | 33 |
| Ethiopian | 140 | 90 | 64 | 294 | 179 | 60 |
| Oriental | 118 | 55 | 46 | 340 | 165 | 48 |
| Australian | 72 | 44 | 61 | 298 | 189 | 64 |
| Neotropical | 130 | 103 | 79 | 683 | 576 | 86 |
| Nearctic | 74 | 24 | 32 | 169 | 52 | 31 |
+-------------+------+--------+-------+------+--------+-------+
TABLE OF REGIONS AND SUB-REGIONS.
+--------------+-----------------------------+--------------------------+
| Regions. | Sub-regions. | Remarks. |
+--------------+-----------------------------+--------------------------+
|I. Palæarctic |1. North Europe. | |
| |2. Mediterranean (or S. Eu.) |Transition to Ethiopian. |
| |3. Siberia. |Transition to Nearctic. |
| |4. Manchuria (or Japan) |Transition to Oriental. |
| | | |
|II. Ethiopian |1. East Africa. |Transition to Palæarctic. |
| |2. West Africa. | |
| |3. South Africa. | |
| |4. Madagascar. | |
| | | |
|III. Oriental |1. Hindostan(or Central Ind.)|Transition to Ethiopian. |
| |2. Ceylon. | |
| |3. Indo-China (or Himalayas) |Transition to Palæarctic. |
| |4. Indo-Malaya. |Transition to Australian. |
| | | |
|IV. Australian|1. Austro-Malaya. |Transition to Oriental. |
| |2. Australia. | |
| |3. Polynesia. | |
| |4. New Zealand. |Transition to Neotropical.|
| | | |
|V. Neotropical|1. Chili (or S. Temp. Am.) |Transition to Australian. |
| |2. Brazil. | |
| |3. Mexico (or Trop. N. Am.) |Transition to Nearctic. |
| |4. Antilles. | |
| | | |
|VI. Nearctic |1. California. | |
| |2. Rocky Mountains. |Transition to Neotropical.|
| |3. Alleghanies (or East U.S.)| |
| |4. Canada. |Transition to Palæarctic. |
+--------------+-----------------------------+--------------------------+
{83}CHAPTER V.
CLASSIFICATION AS AFFECTING THE STUDY OF GEOGRAPHICAL DISTRIBUTION.
A little consideration will convince us, that no inquiry into the causes
and laws which determine the geographical distribution of animals or plants
can lead to satisfactory results, unless we have a tolerably accurate
knowledge of the affinities of the several species, genera, and families to
each other; in other words, we require a natural classification to work
upon. Let us, for example, take three animals--_a_, _b_, and _c_--which
have a general external resemblance to each other, and are usually
considered to be really allied; and let us suppose that _a_ and _b_ inhabit
the same or adjacent districts, while c is found far away on the other side
of the globe, with no animals at all resembling it in any of the
intervening countries. We should here have a difficult problem to solve;
for we should have to show that the general laws by which we account for
the main features of distribution, will explain this exceptional case. But
now, suppose some comparative anatomist takes these animals in hand, and
finds that the resemblance of _c_ to _a_ and _b_ is only superficial, while
their internal structure exhibits marked and important differences; and
that _c_ really belongs to another group of animals, _d_, which inhabits
the very region in which _c_ was found--and we should no longer have
anything to explain. This is no imaginary case. Up to a very few years ago
a curious Mexican animal, _Bassaris astuta_, was almost always classed in
the civet family (Viverridæ), a group entirely {84}confined to Africa and
Asia; but it has now been conclusively shown by Professor Flower that its
real affinities are with the racoons (Procyonidæ), a group confined to
North and South America. In another case, however, an equally careful
examination shows, that an animal peculiar to the Himalayas (_Ælurus
fulgens_) has its nearest ally in the _Cercoleptes_ of South America. Here,
therefore, the geographical difficulty really exists, and any satisfactory
theory of the causes that have led to the existing distribution of living
things, must be able to account, more or less definitely, for this and
other anomalies. From these cases it will be evident, that if any class or
order of animals is very imperfectly known and its classification
altogether artificial, it is useless to attempt to account for the
anomalies its distribution may present; since those anomalies may be, to a
great extent, due to false notions as to the affinities of its component
species.
According to the laws and causes of distribution discussed in the preceding
chapters, we should find limited and defined distribution to be the rule,
universal or indefinite distribution to be the exception, in every natural
group corresponding to what are usually regarded as families and genera;
and so much is this the case in nature, that when we find a group of this
nominal rank scattered as it were at random over the earth, we have a
strong presumption that it is not natural; but is, to a considerable
extent, a haphazard collection of species. Of course this reasoning will
only apply, in cases where there are no unusual means of dispersal, nor any
exceptional causes which might determine a scattered distribution.
From the considerations now adduced it becomes evident, that it is of the
first importance for the success of our inquiry to secure a natural
classification of animals, especially as regards the families and genera.
The higher groups, such as classes and orders, are of less importance for
our purpose; because they are almost always widely and often universally
distributed, except those which are so small as to be evidently the nearly
extinct representatives of a once more extensive series of forms. We now
proceed to explain the classification to be adopted, as low down as the
series of families. To these, equivalent English {85}names are given
wherever they exist, in order that readers possessing no technical
knowledge, may form some conception of the meaning of the term "family" in
zoology.
The primary divisions of the animal kingdom according to two eminent modern
authorities are as follows:
HUXLEY. CARUS AND GERSTAEKER.
Classification of Animals Handbuch der Zoologie
(1869). (1868).
1. Protozoa }
} 1. Protozoa.
2. Infusoria }
3. Coelenterata 2. Coelenterata.
4. Annuloida 3. Echinodermata.
{ 4. Vermes.
5. Annulosa {
{ 5. Arthropoda.
6. Molluscoida 6. Molluscoida.
7. Mollusca 7. Mollusca.
8. Vertebrata 8. Vertebrata.
For reasons already stated it is only with the fifth, seventh, and eighth
of these groups that the present work proposes to deal; and even with the
fifth and seventh only partially and in the most general way.
The classes of the vertebrata, according to both the authors above quoted,
are: 1. Mammalia. 2. Aves. 3. Reptilia. 4. Amphibia. 5. Pisces, in which
order they will be taken here.
The sub-classes and orders of mammalia are as follows:
MAMMALIA.
HUXLEY (1869), FLOWER (1870). CARUS (1868).
{ 1. Primates.
{ 1. Primates {
{ { 5. Prosimii.
{ 2. Chiroptera 2. Chiroptera.
{ 3. Insectivora 3. Insectivora.
{ { 6. Carnivora.
{ 4. Carnivora {
{ { 7. Pinnipedia.
Monodelphia { 5. Cetacea }
{ } 12. Natantia.
{ 6. Sirenia }
{ {10. Artiodactyla.
{ 7. Ungulata {
{ {11. Perissodactyla.
{ 8. Proboscidea 9. Proboscidea.
{ 9. Hyracoidea 8. Lamnungia.
{ 10. Rodentia 4. Rodentia.
{ 11. Edentata 13. Bruta.
{
Didelphia 12. Marsupialia 14. Marsupialia.
Ornithodelphia 13. Monotremata 15. Monotremata.
{86}The above series of orders is arranged according to Professor Flower's
_Osteology of Mammalia_, and they will follow in this succession throughout
my work. Professor Huxley arranges the same orders in a different series.
In determining the manner in which the several orders shall be subdivided
into families, I have been guided in my choice of classifications mainly by
the degree of attention the author appears to have paid to the group, and
his known ability as a systematic zoologist; and in a less degree by
considerations of convenience as regards the special purposes of
geographical distribution. In many cases it is a matter of great doubt
whether a certain group should form several distinct families or be united
into one or two; but one method may bring out the peculiarities of
distribution much better than the other, and this is, in our case, a
sufficient reason for adopting it.
For the Primates I follow, with some modifications, the classification of
Mr. St. George Mivart given in his article "Apes" in the new edition of the
_Encyclopædia Britannica_, and in his paper in the _Proceedings of the
Zoological Society of London_, 1865, p. 547. It is as follows:
Order--PRIMATES, divided into two Sub-orders:
I. Anthropoidea.
II. Lemuroidea.
Sub-order--ANTHROPOIDEA.
Fam.
Hominidæ Man.
{ 1. Simiidæ Anthropoid Apes.
Simii { 2. Semnopithecidæ Old-world Monkeys.
{ 3. Cynopithecidæ Baboons and Macaques.
Cebii { 4. Cebidæ American Monkeys.
{ 5. Hapalidæ Marmosets.
Sub-order--LEMUROIDEA.
Fam.
6. Lemuridæ Lemurs.
7. Tarsiidæ Tarsiers.
8. Chiromyidæ Aye-ayes.
Omitting man (for reasons stated in the preface) the three first families
are considered by Professor Mivart to be sub-families of Simiidæ; but as
the geographical distribution of the Old World apes is especially
interesting, it is thought {87}better to treat them as families, a rank
which is claimed for the anthropoid apes by many naturalists.
As no good systematic work on the genera and species of bats has been yet
published, I adopt the five families as generally used in this country,
with the genera as given in the papers of Dr. J. E. Gray and Mr. Tomes. A
monograph by Dr. Peters has long been promised, and his outline arrangement
was published in 1865, but this will perhaps be materially altered when the
work appears.
Order--CHIROPTERA.
Fam.
Frugivora 9. Pteropidæ Fruit-eating Bats.
{Istiophora { 10. Phyllostomidæ Leaf-nosed Bats.
Insectivora { { 11. Rhinolophidæ Horse-shoe Bats.
{
{Gymnorhini { 12. Vespertilionidæ True Bats.
{ 13. Noctilionidæ Dog-headed Bats.
The genera of Chiroptera are in a state of great confusion, the names used
by different authors being often not at all comparable, so that the few
details given of the distribution of the bats are not trustworthy. We have
therefore made little use of this order in the theoretical part of the
work.
The osteology of the Insectivora has been very carefully worked out by
Professor Mivart in the _Jounral of Anatomy and Physiology_ (Vol. ii., p.
380), and I follow his classification as given there, and in the
_Proceedings of the Zoological Society_ (1871).
Order--INSECTIVORA.
Fam.
14. Galeopithecidæ Flying Lemurs.
15. Macroscelididæ Elephant Shrews.
16. Tupaiidæ Squirrel Shrews.
17. Erinaceidæ Hedgehogs.
18. Centetidæ Tenrecs.
19. Potamogalidæ Otter Shrew.
20. Chrysochloridæ Golden Moles.
21. Talpidæ Moles.
22. Soricidæ Shrews.
The next order, Carnivora, has been studied in detail by Professor Flower;
and I adopt the classification given by him in the _Proceedings of the
Zoological Society_, 1869, p. 4.
{88}Order--CARNIVORA.
Fam.
{ 23. Felidæ Cats, Lion, &c.
{ 24. Cryptoproctidæ Cryptoprocta.
{ Æluroidea { 25. Viverridæ Civets.
{ { 26. Protelidæ Aard-wolf.
{ { 27. Hyænidæ Hyænas.
{
Fissipedia { Cynoidea 28. Canidæ Dogs, Foxes, &c.
{
{ {29. Mustelidæ Weasels.
{ {30. Procyonidæ Racoons.
{ Arctoidea {31. Æluridæ Pandas.
{32. Ursidæ Bears.
{33. Otariidæ Eared Seals.
Pinnipedia {34. Trichechidæ Walrus.
{35. Phocidæ Seals.
The Cetacea is one of those orders the classification of which is very
unsettled. The animals comprising it are so huge, and there is so much
difficulty in preserving them, that only a very few species are known with
anything like completeness. A considerable number of genera and species
have been described or indicated; but as many of these are founded on
imperfect specimens of perhaps a single individual, it is not to be
wondered at that those few naturalists who occupy themselves with the study
of these large animals, cannot agree as to the proper mode of grouping them
into natural families. They are, however, of but little importance to us,
as almost all the species inhabit the ocean, and of only a few of them can
it be said that anything is accurately known of their distribution. I
therefore consider it best to follow Professor Carus, who makes a smaller
number of families; but I give also the arrangement of Dr. Gray in his
British Museum catalogue of whales and seals, as modified subsequently in
the _Proceedings of Zoological Society_, 1870, p. 772. The Zeuglodontidæ, a
family of extinct tertiary whales, are classed by Professors Owen and Carus
between Cetacea and Sirenia, while Professor Huxley considers them to have
been carnivorous and allied to the seals.
{89}Order--CETACEA.
Fam. (CARUS). Fam. (GRAY).
Sub-order I.-- { Balænidæ 36. Balænidæ.
Mystaceti. { Balænopteridæ 37. Balænopteridæ.
{ Catodontidæ 38. Catodontidæ.
{
{ { Hyperoodontidæ.
{ Hyperoodontidæ 39. { Epiodontidæ.
{ { Xiphiadæ.
{
Sub-order II.-- { Monodontidæ 40. (Part of Delphinidæ.)
Odontoceti {
{ { Platanistidæ.
{ { Iniadæ.
{ { Delphinidæ.
{ Delphinidæ 41. { Globiocephalidæ.
{ Orcadæ.
{ Belugidæ.
{ Pontoporiadæ.
Extinct family Zeuglodontidæ.
Order--SIRENIA.
The order Sirenia, comprising the sea-cows, consists of a single family:
Family 42. Manatidæ.
The extensive order Ungulata comprises the three orders Pachydermata,
Solidungula, and Ruminantia of the older naturalists. The following
classification is that now generally adopted, the only difference of
opinion being as to whether some of the groups should be classed as
families or sub-families, a matter of little importance for our purpose:
Order--UNGULATA.
Fam.
Perissodactyla or } 43. Equidæ Horses.
Odd-toed Ungulates } 44. Tapiridæ Tapirs.
45. Rhinocerotidæ Rhinoceros.
{ Suina { 46. Hippopotamidæ Hippopotamus.
{ { 47. Suidæ Swine.
{
Artiodactyla or { Tylopoda 48. Camelidæ Camels.
Even-toed Ungulates {
{ Tragulina 49. Tragulidæ Chevrotains.
{
{ { 50. Cervidæ Deer.
{ Pecora { 51. Camelopardidæ Giraffes.
{ 52. Bovidæ { Cattle, Sheep,
{ Antelopes, &c.
{90}The two next orders consist of but a single family each, viz.:
Order Fam.
PROBOSCIDEA 53. Elephantidæ Elephants.
HYRACOIDEA 54. Hyracidæ Rock-rabbits.
We now come to the Rodentia, a very extensive and difficult order, in which
there is still much difference of opinion as to the details of
classification, although the main outlines are pretty well settled. The
foundations of a true classification of this order were laid by Mr. G. R.
Waterhouse more than thirty years ago, and succeeding authors have done
little more than follow his arrangement with unimportant modifications.
Professor Lilljeborg, of Upsala, has however made a special study of this
group of animals, and has given an original and detailed classification of
all the genera. (_Systematisk Öfversigt af de Gnagande Däggdjuren, Glires.
Upsala, 1866._) I follow this arrangement with a few slight modifications
suggested by other naturalists, and which make it better adapted for the
purposes of this work.
Order--RODENTIA.
Fam.
{ 55. Muridæ Rats.
{ 56. Spalacidæ Mole-rats.
{ 57. Dipodidæ Jerboas.
{ 58. Myoxidæ Dormice.
{ Murina { 59. Saccomyidæ Pouched Rats.
{ (Waterhouse) { 60. Castoridæ Beavers.
{ { 61. Sciuridæ Squirrels.
Simplicidentati { { 62. Haploodontidæ Sewellels.
{
{ { 63. Chinchillidæ Chinchillas.
{ { 64. Octodontidæ Octodons.
{ Hystricina { 65. Echimyidæ Spiny Rats.
(Waterhouse) { 66. Cercolabidæ Tree Porcupines
{ 67. Hystricidæ Porcupines.
{ 68. Caviidæ Cavies.
Duplicidentati { Leporina { 69. Lagomyidæ Pikas.
{ (Waterhouse) { 70. Leporidæ Hares.
The Edentata have been classified by Mr. Turner, in the _Proceedings of the
Zoological Society_ (1851, p. 205), by Dr. Gray in the British Museum
Catalogue, and by Professor Carus in his _Handbuch_. The former takes a
middle course between {91}the numerous families of Dr. Gray, seven in
number, and the two families to which Professor Carus restricts the
existing species. I therefore follow Mr. Turner.
Order--EDENTATA.
Fam.
Bradypoda 71. Bradypodidæ Sloths.
{ 72. Manididæ Scaly Ant-eaters.
{ 73. Dasypodidæ Armadillos.
Entomophaga { 74. Orycteropodidæ Ant-bears.
{ 75. Myrmecophagidæ Ant-eaters.
The Marsupials have been well classified and described by Mr. Waterhouse in
the first volume of his _Natural History of Mammalia_, and his arrangement
is here followed. The suborders adopted by Professor Carus are also given.
Order--MARSUPIALIA.
Fam.
{ 76. Didelphidæ Opossums.
{ 77. Dasyuridæ Native Cats.
Rapacia (Wagner) { 78. Myrmecobiidæ Native Ant-eater.
{ 79. Peramelidæ Bandicoots.
Poephaga (Owen) 80. Macropodidæ Kangaroos.
Carpophaga (Owen) 81. Phalangistidæ Phalangers.
Rhizophaga (Owen) 82. Phascolomyidæ Wombats.
Order--MONOTREMATA.
The last order, the Monotremata, consist of two families, which Professor
Carus combines into one, but which it seems more natural to keep separate.
Fam.
83. Ornithorhynchidæ Duckbill.
84. Echidnidæ Echidna.
{92}BIRDS.
Birds are perhaps the most difficult to classify of all the divisions of
the vertebrata. The species and genera are exceedingly numerous, and there
is such a great uniformity in general structure and even in the details of
external form, that it is exceedingly difficult to find characters by which
orders and families can be characterised. For a long time the system of
Vigors and Swainson was followed; but this wholly ignored anatomical
characters and in many cases plainly violated well-marked affinities.
Characters derived from the form of the sternum, the scutellation of the
tarsi, and the arrangement of the feathers, have all assisted in
determining natural groups. More recently Professor Huxley has applied the
variations of the bony palate to the general arrangement of birds; and
still more recently Professor Garrod has studied certain leg-muscles for
the same purpose. The condition of the young as regards plumage, and even
the form, texture, and coloration of the egg, have also been applied to
solve doubtful cases of affinity; yet the problem is not settled, and it
will probably remain for another generation of ornithologists to determine
with any approach to accuracy what are the most natural divisions of the
class into orders and families. In a work like the present it is evidently
not advisable to adopt all the recent classifications; since experience has
shown that no arrangement in which one set of characters is mainly relied
on, long holds its ground. Such modifications of the old system as seem to
be well established will be adopted; but the older groups will be adhered
to in cases where the most recent classifications are open to doubt, or
seem inconvenient as separating families, which, owing to their similarity
in general structure, form and habits are best kept together for the
purposes of geographical distribution.
The old plan of putting the birds of prey at the head of the class, is now
almost wholly given up; both because they are not {93}the most highly
organised, but only one of the most specialised forms of birds, and because
their affinities are not with the Passeres, but rather with the cormorants
and some other of the aquatic groups. The Passeres therefore are placed
first; and the series of families is begun by the thrushes, which are
certainly the most typical and generally well-organised form of birds.
Instead of the Scansores and Fissirostres of the older authors, the order
Picariæ, which includes them both, is adopted, but with some reluctance; as
the former are, generally speaking, well marked and strongly contrasted
groups, although certain families have been shown to be intermediate. In
the Picariæ are included the goat-suckers, swifts, and humming-birds,
sometimes separated as a distinct order, Macrochires. The parrots and the
pigeons form each a separate order. The old groups of Grallæ and Anseres
are preserved, as more convenient than breaking them up into widely
separated parts; for though the latter plan may in some cases more strictly
represent their affinities, its details are not yet established, nor is it
much used by ornithologists. In accordance with these views the following
is the series of orders and families of birds adopted in this work:
Class--AVES.
Orders.
1. Passeres { Including the great mass of the smaller birds--Crows,
{ Finches, Flycatchers, Creepers, Honeysuckers, &c., &c.
2. Picariæ { Including Woodpeckers, Cuckoos, Toucans, Kingfishers,
{ Swifts, &c., &c.
3. Psittaci Parrots only.
4. Columbæ Pigeons and the Dodo.
5. Gallinæ Grouse, Pheasants, Curassows, Mound-builders, &c.
6. Opisthocomi The Hoazin only.
7. Accipitres Eagles, Owls, and Vultures.
8. Grallæ Herons, Plovers, Rails, &c.
9. Anseres Gulls, Ducks, Divers, &c.
10. Struthiones Ostrich, Cassowary, Apteryx, &c.
The Passeres consist of fifty families, which may be arranged and grouped
in series as follows. It must however be remembered that the first family
in each series is not always that which is most allied to the last family
of the preceding series. All extensive natural groups consist of divergent
or branching alliances, which renders it impossible to arrange the whole in
one continuous series.
{94}A.--TYPICAL OR TURDOID PASSERES.
1. Turdidæ Thrushes.
2. Sylviidæ Warblers.
3. Timaliidæ Babblers.
4. Panuridæ Reedlings.
5. Cinclidæ Dippers.
6. Troglodytidæ Wrens.
7. Chamæidæ
8. Certhiidæ Creepers.
9. Sittidæ Nuthatches.
10. Paridæ Tits.
11. Liotrichidæ Hill-tits.
12. Phyllornithidæ Green Bulbuls.
13. Pycnonotidæ Bulbuls.
14. Oriolidæ Orioles.
15. Campephagidæ Caterpillar-shrikes.
16. Dicruridæ Drougos.
17. Muscicapidæ Flycatchers.
18. Pachycephalidæ Thick-heads.
19. Laniidæ Shrikes.
20. Corvidæ Crows.
21. Paradiseidæ Paradise-birds.
22. Meliphagidæ Honey-suckers.
23. Nectarineidæ Sun-birds.
B.--TANAGROID PASSERES.
24. Dicæidæ Flower-peckers.
25. Drepanididæ
26. Cærebidæ Sugar-birds.
27. Mniotiltidæ Wood-warblers.
28. Vireonidæ Greenlets.
29. Ampelidæ Waxwings.
30. Hirundinidæ Swallows.
31. Icteridæ Hangnests.
32. Tanagridæ Tanagers.
33. Fringillidæ Finches.
C.--STURNOID PASSERES.
34. Ploceidæ Weaver-birds.
35. Sturnidæ Starlings.
36. Artamidæ Swallow-shrikes.
37. Alaudidæ Larks.
38. Motacillidæ Wagtails.
D.--FORMICAROID PASSERES.
39. Tyrannidæ Tyrants.
40. Pipridæ Manakins.
41. Cotingidæ Chatterers.
42. Phytotomidæ Plant-cutters.
43. Eurylæmidæ Broad-bills.
44. Dendrocolaptidæ American Creepers.
45. Formicariidæ Ant-thrushes.
46. Pteroptochidæ {95}
47. Pittidæ Pittas.
48. Paictidæ
E.--ANOMALOUS PASSERES.
49. Menuridæ Lyre-birds.
50. Atrichidæ Scrub-birds.
The preceding arrangement is a modification of that proposed by myself in
the _Ibis_ (1874, p. 406). The principal alterations are adding the
families Panuridæ and Sittidæ in series A, commencing series B with
Dicæidæ; bringing Vireonidæ next to the allied American family Mniotiltidæ;
and placing Motacillidæ in series C next to Alaudidæ. At the suggestion of
Professor Newton I place Menuridæ and Atrichidæ apart from the other
Passeres, as they both possess striking peculiarities of anatomical
structure.
The heterogeneous families constituting the order Picariæ may be
conveniently arranged as follows:
{ 51. Picidæ Woodpeckers.
{ 52. Yungidæ Wrynecks.
{ 53. Indicatoridæ Honey-guides.
Sub-order-- { 54. Megalæmidæ Barbets.
Scansores. { 55. Rhamphastidæ Toucans.
{ 56. Musophagidæ Plantain-eaters.
{ 57. Coliidæ Colies.
{ 58. Cuculidæ Cuckoos.
Intermediate 59. Leptosomidæ The Leptosoma.
{ 60. Bucconidæ Puff-birds.
{ 61. Galbulidæ Jacamars.
{ 62. Coraciidæ Rollers.
{ 63. Meropidæ Bee-eaters.
{ 64. Todidæ Todies.
{ 65. Momotidæ Motmots.
{ 66. Trogonidæ Trogons.
Sub-order-- { 67. Alcedinidæ Kingfishers.
Fissirostres. { 68. Bucerotidæ Hornbills.
{ 69. Upupidæ Hoopoes.
{ 70. Irrisoridæ Promerops.
{ 71. Podargidæ Frog-mouths.
{ 72. Steatornithidæ The Guacharo.
{ 73. Caprimulgidæ Goatsuckers.
{ 74. Cypselidæ Swifts.
{ 75. Trochilidæ Humming-birds.
{96}The Psittaci or parrot tribe are still in a very unsettled state of
classification; that recently proposed by Professor Garrod differing widely
from the arrangement adopted in Dr. Finsch's monograph of the order. Taking
advantage of the researches of these and other authors, the following
families are adopted as the most convenient in the present state of our
knowledge:
76. Cacatuidæ The Cockatoos.
77. Platycercidæ The Broad-tailed Paroquets of Australia.
78. Palæornithidæ The Oriental Parrots and Paroquets.
79. Trichoglossidæ The Brush-tongued Paroquets and Lories.
80. Conuridæ The Macaws and their allies.
81. Psittacidæ The African and South American Parrots.
82. Nestoridæ The Nestors of New Zealand.
83. Stringopidæ The Owl-parrots of New Zealand.
The Columbæ, or pigeons, are also in a very unsatisfactory state as regards
a natural classification. The families, sub-families, and genera proposed
by various authors are very numerous, and often quite irreconcilable. I
therefore adopt only two families; and generally follow Mr. G. R. Gray's
hand-list for the genera, except where trustworthy authorities exist for a
different arrangement. The families are:
84. Columbidæ Pigeons and Doves.
85. Dididæ The extinct Dodo and allies.
The Gallinæ, or game-birds, may be divided into seven families:
Fam. Sub-fam.
86. Pteroclidæ Sand-grouse.
87. Tetraonidæ Partridges and Grouse.
{ Pavoninæ Peafowl.
{ Lophophorinæ Tragopans, &c.
{ Phasianinæ Pheasants.
88. Phasianidæ { Euplocaminæ Fire-backed Pheasants, &c.
{ Gallinæ Jungle-fowl.
{ Meleagrinæ Turkeys.
{ Numidinæ Guinea-fowl.
89. Turnicidæ Hemipodes.
90. Megapodiidæ Mound-makers.
{ Cracinæ Curassows.
91. Cracidæ { Penelopinæ Guans.
{ Oreophasinæ Mountain-pheasant.
92. Tinamidæ Tinamous.
{97}The Opisthocomi consist of one family containing a single species, the
"Hoatzin" of Guiana.
Family 93. Opisthocomidæ.
The Accipitres, or birds of prey, which were long considered to be the
highest and most perfect order of birds, are now properly placed lower down
in the series, their affinities being more with the aquatic than with the
perching birds. The following is the arrangement adopted by Mr. Sharpe in
his recently published British Museum catalogue of diurnal birds of prey:--
Sub-orders. Fam. Sub-families.
{ Vulturinæ Vultures.
{ 94. Vulturidæ { Sarcorhamphinæ Turkey-buzzards.
{
Falcones { 95. Serpentariidæ
{
{ { Polyborniæ Caracaras.
{ { Accipitrinæ Hawks.
{ 96. Falconidæ { Buteoninæ Buzzards.
{ Aquilinæ Eagles.
{ Falconinæ Falcons.
Pandiones 97. Pandionidæ Fishing-eagles.
Striges 98. Strigidæ Owls.
The Grallæ or Grallatores are in a very unsettled state. The following
series of families is in accordance with the views of some of the best
modern ornithologists:
99. Rallidæ Rails, &c.
100. Scolopacidæ Sandpipers and Snipes
101. Chionididæ Sheath-bills.
102. Thinocoridæ Quail-snipes.
103. Parridæ Jacanas.
104. Glareolidæ Pratincoles.
105. Charadriidæ Plovers.
106. Otididæ Bustards.
107. Gruidæ Cranes.
108. Cariamidæ Cariamas.
109. Aramidæ Guaraunas.
110. Psophiidæ Trumpeters.
111. Eurypygidæ Sun-bitterns.
112. Rhinochoetidæ Kagus.
113. Ardeidæ Herons.
114. Plataleidæ Spoonbills and Ibis.
115. Ciconiidæ Storks.
116. Palamedeidæ Screamers.
117. Phænicopteridæ Flamingoes.
{98}The Anseres or Natatores are almost equally unsettled. The flamingoes
are usually placed in this order, but their habits best assort with those
of the waders.
Fam.
118. Anatidæ Duck and Geese.
119. Laridæ Gulls.
120. Procellariidæ Petrels.
121. Pelecanidæ Pelicans.
122. Spheniscidæ Penguins.
123. Colymbidæ Divers.
124. Podicipidæ Grebes.
125. Alcidæ Auks.
The last order of birds is the Struthiones or Ratitæ, considered by many
naturalists to form a distinct sub-class. It consists of comparatively few
species, either living or recently extinct.
Fam.
{ 126. Struthionidæ Ostriches.
Living { 127. Casuariidæ Cassowaries.
{ 128. Apterygidæ Apteryx.
{ 129. Dinornithidæ Dinornis.
Extinct { 130. Palapterygidæ Palapteryx.
{ 131. Æpyornithidæ Æpyornis.
REPTILES.
In reptiles I follow the classification of Dr. Günther as given in the
_Philosophical Transactions_, vol. clvii., p. 625. He divides the class
into five orders as follows:--
Sub-classes. Orders.
{ 1. Ophidia Serpents.
I. Squamata { 2. Lacertilia Lizards.
{ 3. Rhyncocephalina The Hatteria.
II. Loricata 4. Crocodilia Crocodiles.
III. Cataphracta 5. Chelonia Tortoises.
In the arrangement of the families comprised in each of these orders I also
follow the arrangement of Dr. Günther and Dr. J. E. Gray, as given in the
British Museum Catalogue, or as modified by the former gentleman who has
kindly given me much personal information.
{99}The Ophidia, or Snakes, form the first order and are classified as
follows:--
Fam.
{ 1. Typhlopidæ }
{ 2. Tortricidæ }
{ 3. Xenopeltidæ } Burrowing Snakes.
{ 4. Uropeltidæ }
{ 5. Calamaridæ Dwarf ground-snakes.
{ 6. Oligodontidæ.
{ 7. Colubridæ Colubrine Snakes.
{ 8. Homalopsidæ Fresh-water Snakes.
{ 9. Psammophidæ Desert-snakes.
Innocuous Snakes { 10. Rachiodontidæ.
{ 11. Dendrophidæ Tree-snakes.
{ 12. Dryiophidæ Whip-snakes.
{ 13. Dipsasidæ Nocturnal tree-snakes.
{ 14. Scytalidæ
{ 15. Lycodontidæ Fanged ground-snakes.
{ 16. Amblycephalidæ Blunt-heads.
{ 17. Pythonidæ Pythons.
{ 18. Erycidæ Sand-snakes.
{ 19. Acrochordidæ Wart-snakes.
{ 20. Elapidæ Cobras, &c.
Venomous Colubrine { 21. Dendraspididæ.
Snakes { 22. Atractaspididæ.
{ 23. Hydrophidæ Sea-snakes.
{ 24. Crotalidæ Pit-vipers.
Viperine Snakes { 25. Viperidæ True vipers.
The second order, Lacertilia, are arranged as follows:--
Fam.
26. Trogonophidæ }
27. Chirotidæ }
28. Amphisbænidæ } Amphisbænians.
29. Lepidosternidæ }
30. Varanidæ Water Lizards.
31. Helodermidæ.
32. Teidæ Teguexins.
33. Lacertidæ } Land Lizards.
34. Zonuridæ }
35. Chalcidæ.
36. Anadiadæ.
37. Chirocolidæ.
38. Iphisadæ.
39. Cercosauridæ.
40. Chamæsauridæ.
41. Gymnopthalmidæ Gape-eyed Scinks.
42. Pygopodidæ Two-legged Lizards.
43. Aprasiadæ.
44. Lialidæ. {100}
45. Scincidæ Scinks.
46. Ophiomoridæ Snake-lizards.
47. Sepidæ Sand-lizards.
48. Acontiadæ.
49. Geckotidæ Geckoes.
50. Iguanidæ Iguanas.
51. Agamidæ Fringed Lizards.
52. Chameleonidæ Chameleons.
The third order, Rhyncocephalina consists of a single family:--
53. Rhyncocephalidæ The Hatteria of New Zealand.
The fourth order, Crocodilia or Loricata, consists of three families:--
54. Gavialidæ Gavials.
55. Crocodilidæ Crocodiles.
56. Alligatoridæ Alligators.
The fifth order, Chelonia, consists of four families:--
57. Testudinidæ Land and fresh-water Tortoises.
58. Chelydidæ Fresh-water Turtles.
59. Trionychidæ Soft Turtles.
60. Cheloniidæ Sea Turtles.
AMPHIBIA.
In the Amphibia I follow the classification of Professor Mivart, as given
for a large part of the order in the _Proceedings of the Zoological
Society_ for 1869. For the remainder I follow Dr. Strauch, Dr. Günther, and
a MSS. arrangement kindly furnished me by Professor Mivart.
The class is first divided into three groups or orders, and then into
families as follows:--
{101}Order I.--PSEUDOPHIDIA.
Fam.
1. Cæciliadæ Cæcilia.
Order II.--BATRACHIA URODELA.
2. Sirenidæ Siren.
3. Proteidæ Proteus.
4. Amphiumidæ Amphiuma.
5. Menopomidæ Giant Salamanders.
6. Salamandridæ Salamanders and Newts.
Order III.--BATRACHIA ANOURA.
7. Rhinophrynidæ }
8. Phryniscidæ }
9. Hylaplesidæ } Toads.
10. Bufonidæ }
11. Xenorhinidæ }
12. Engystomidæ }
13. Bombinatoridæ }
14. Plectromantidæ } Frogs.
15. Alytidæ }
16. Pelodryadæ }
17. Hylidæ } Tree Frogs.
18. Polypedatidæ }
19. Ranidæ } Frogs.
20. Discoglossidæ }
21. Pipidæ } Tongueless
22. Dactylethridæ } Toads.
FISHES.
These are arranged according to the classification of Dr. Günther, whose
great work "The British Museum Catalogue of Fishes," has furnished almost
all the material for our account of the distribution of the class.
In that work all existing fishes are arranged in six sub-classes and
thirteen orders. A study of the extraordinary _Ceratodus_ from Australia
has induced Dr. Günther to unite three of his sub-classes; but as his
catalogue will long remain a handbook for every student of fishes, it seems
better to follow the arrangement there given, indicating his later views by
bracketing together the groups he now thinks should be united.
-----------------------------------------+---------+--------------- {102}
Sub-class. Order. |Families.| Remarks.
-----------------------------------------+---------+------------------
{ 1. Acanthopterygii | 47 | Gasterosteidæ to
{ | | Notacanthi.
{ 2. Do. Pharyncognathi | 5 | Pomacentridæ to
{ | | Chromidæ.
Ganoidei { 3. Acanthini | 6 | Gadopsidæ to
======== { | | Pleuronectidæ.
{Teleostei { 4. Physostomi | 29 | Siluridæ to
{ { | | Pegasidæ.
{ { 5. Lophobranchii | 2 | Solenostomidæ and
{ { | | Syngnathidæ.
{ { 6. Plectognathi | 2 | Sclerodermi and
{ { | | Gymnodontes.
{ | |
{Dipnoi 7. Sirenoidei | 1 | Sirenoidei.
{ | |
{ { 8. Holostei | 3 | Amiidæ to
{ { | | Lepidosteidæ.
{Ganoidei { 9. Chondrostei | 2 | Accipenseridæ and
{ | | Polydontidæ.
| |
Chondropterygii { 10. Holocephala | 1 | Chimæridæ.
{ 11. Plagiostomata | 15 | Carchariidæ to
| | Myliobatidæ.
| |
Cyclostomata 12. Marsipobranchii | 2 | Petromyzontidæ and
| | Myxinidæ.
| |
Leptocardii 13. Cirrhostomi | 1 | Cirrhostomi.
+---------+
Total 116 families.
-----------------------------------------------------------------------
INSECTS.
The families and genera of insects are so immensely numerous, probably
exceeding fifty-fold those of all other land animals, that for this cause
alone it would be impossible to enter fully into their distribution. It is
also quite unnecessary, because many of the groups are so liable to be
transported by accidental causes, that they afford no useful information
for our subject; while others are so obscure and uninteresting that they
have been very partially collected and studied, and are for this reason
equally ineligible. I have therefore selected a few of the largest and most
conspicuous families, which have been so assiduously collected in every
part of the globe, and so carefully studied at home, as to afford valuable
materials for comparison with the vertebrate groups, when we have made due
allowance for the dependence of many insects on peculiar forms of
vegetation, and the facility with which many of them are transported either
in the egg, larva, or perfect state, by winds, currents, and other less
known means.
I confine myself then, almost exclusively, to the sixteen families of
Diurnal Lepidoptera or butterflies, and to six of the most extensive,
conspicuous, and popular families of Coleoptera. {103}The number of species
of Butterflies is about the same as that of Birds, while the six families
of Coleoptera selected, comprise more than twenty thousand species, far
exceeding the number of all other vertebrates. These families have all been
recently catalogued, so that we have very complete information as to their
arrangement and distribution.
LEPIDOPTERA DIURNA, OR BUTTERFLIES.
Fam.
1. Danaidæ.
2. Satyridæ.
3. Elymniidæ.
4. Morphidæ.
5. Brassolidæ.
6. Acræidæ.
7. Heliconidæ.
8. Nymphalidæ.
9. Libythæidæ.
10. Nemeobiidæ.
11. Eurygonidæ.
12. Erycinidæ.
13. Lycænidæ.
14. Pieridæ.
15. Papilionidæ.
16. Hesperidæ.
COLEOPTERA, OR BEETLES.
Fam.
1. Cicindelidæ Tiger-beetles.
2. Carabidæ Ground-beetles.
3. Lucanidæ Stag-beetles.
4. Cetoniidæ Rose-chafers.
5. Buprestidæ Metallic Beetles.
6. Longicornia Long-horned Beetles.
The above families comprise the extensive series of ground beetles
(Carabidæ) containing about 9,000 species, and the Longicorns, which are
nearly as numerous and surpass them in variety of form and colour as well
as in beauty. The Cetoniidæ and Buprestidæ are among the largest and most
brilliant of beetles; the Lucanidæ are pre-eminent for remarkable form, and
the Cicindelidæ for elegance; and all the families are especial favourites
with entomologists, so that the whole earth has been ransacked to procure
fresh species.
Results deduced from a study of these will, therefore, fairly represent the
phenomena of distribution of Coleoptera, and, as they are very varied in
their habits, perhaps of insects in general.
{104}MOLLUSCA.
The Mollusca are usually divided into five classes as follows:--
Classes.
I. Cephalopoda Cuttle-fish.
II. Gasteropoda Snails and aquatic Univalves.
III. Pteropoda Oceanic Snails.
IV. Brachiopoda Symmetrical Bivalves.
V. Conchifera Unsymmetrical Bivalves.
The Gasteropoda and Conchifera alone contain land and freshwater forms, and
to these we shall chiefly confine our illustrations of the geographical
distribution of the Mollusca. The classification followed is that of Dr.
Pfeiffer for the Operculata and Dr. Von Martens for the Helicidæ. The
families chiefly referred to are:--
Class II.--GASTEROPODA.
Order 2.--Pulmonifera.
Fam.
{ 1. Helicidæ.
{ 2. Limacidæ.
{ 3. Oncidiadæ.
In-operculata { 4. Limnæidæ.
{ 5. Auriculidæ.
{ 6. Aciculidæ.
{ 7. Diplommatinidæ.
Operculata { 8. Cyclostomidæ.
{ 9. Helicinidæ.
PART II.
_ON THE DISTRIBUTION OF EXTINCT ANIMALS._
{107}CHAPTER VI.
THE EXTINCT MAMMALIA OF THE OLD WORLD.
Although it may seem somewhat out of place to begin the systematic
treatment of our subject with extinct rather than with living animals, it
is necessary to do so in order that we may see the meaning and trace the
causes of the existing distribution of animal forms. It is true, that the
animals found fossil in a country are very generally allied to those which
still inhabit it; but this is by no means universally the case. If it were,
the attempt to elucidate our subject by Palæontology would be hopeless,
since the past would show us the same puzzling diversities of faunas and
floras that now exist. We find however very numerous exceptions to this
rule, and it is these exceptions which tell us of the past migrations of
whole groups of animals. We are thus enabled to determine what portion of
the existing races of animals in a country are descendants of its ancient
fauna, and which are comparatively modern immigrants; and combining these
movements of the forms of life with known or probable changes in the
distribution of land and sea, we shall sometimes be able to trace
approximately the long series of changes which have resulted in the actual
state of things. To gain this knowledge is our object in studying the
"Geographical Distribution of Animals," and our plan of study must be
determined, mainly, by the facilities it affords us for attaining this
object. In discussing the countless details of distribution we shall meet
with in our survey of the zoological regions, we shall often find it useful
to refer to the evidence we possess of the range of the group in question
in {108}past times; and when we attempt to generalise the phenomena on a
large scale, with the details fresh in our memory, we shall find a
reference to the extinct faunas of various epochs to be absolutely
necessary.
The degree of our knowledge of the Palæontology of various parts of the
world is so unequal, that it will not be advisable to treat the subject
under each of our six regions. Yet some subdivision must be made, and it
seems best to consider separately the extinct animals of the Old and of the
New Worlds. Those of Europe and Asia are intimately connected, and throw
light on the past changes which have led to the establishment of the three
great continental Old World regions, with their various subdivisions. The
wonderful extinct fauna recently discovered in North America, with what was
previously known from South temperate America, not only elucidates the past
history of the whole continent, but also gives indications of the mutual
relations of the eastern and western hemispheres.
The materials to be dealt with are enormous; and it will be necessary to
confine ourselves to a general summary, with fuller details on those points
which directly bear upon our special subject. The objects of most interest
to the pure zoologist and to the geologist--those strange forms which are
farthest removed from any now living--are of least interest to us, since we
aim at tracing the local origin or birthplace of existing genera and
families; and for this purpose animals whose affinities with living forms
are altogether doubtful, are of no value whatever.
The great mass of the vertebrate fossils of the tertiary period consist of
mammalia, and this is precisely the class which is of most value in the
determination of zoological regions. The animals of the secondary period,
though of the highest interest to the zoologist are of little importance to
us; both because of their very uncertain affinities for any existing
groups, and also because we can form no adequate notion of the distribution
of land and sea in those remote epochs. Our great object is to trace back,
step by step, the varying distribution of the chief forms of life; and to
deduce, wherever possible, the physical changes which must have accompanied
or caused such changes. {109}The natural division of our subject therefore
is into geological periods. We first go back to the Post-Pliocene period,
which includes that of the caves and gravels of Europe containing flint
implements, and extends back to the deposit of the glacial drift in the
concluding phase of the glacial epoch. Next we have the Pliocene period,
divided into its later portion (the Newer Pliocene) which includes the
Glacial epoch of the northern hemisphere; and its earlier portion (the
Older Pliocene), represented by the red and coralline crag of England, and
deposits of similar age in the continent. During this earlier epoch the
climate was not very dissimilar from that which now prevails; but we next
get evidence of a still earlier period, the Miocene, when a warmer climate
prevailed in Europe, and the whole fauna and flora were very different.
This is perhaps the most interesting portion of the tertiary deposits, and
furnishes us with the most valuable materials for our present study.
Further back still we have the Eocene period, with apparently an almost
tropical climate in Europe; and here we find a clue to some of the most
puzzling facts in the distribution of living animals. Our knowledge of this
epoch is however very imperfect; and we wait for discoveries that will
elucidate some of the mystery that still hangs over the origin and
migrations of many important families. Beyond this there is a great chasm
in the geological record as regards land animals; and we have to go so far
back into the past, that when we again meet with mammalia, birds, and
land-reptiles, they appear under such archaic forms that they cease to have
any local or geographical significance, and we can only refer them to
wide-spread classes and orders. For the purpose of elucidating geographical
distribution, therefore, it is, in the present state of our knowledge,
unnecessary to go back beyond the tertiary period of geology.
The remains of Mammalia being so much more numerous and important than
those of other classes, we shall at first confine ourselves almost
exclusively to these. What is known of the birds, reptiles, and fishes of
the tertiary epoch will be best indicated by a brief connected sketch of
their fossils in all parts of the globe, which we shall give in a
subsequent chapter.
{110}_Historic Period._--In tracing back the history of the organic world
we find, even within the limits of the historical period, that some animals
have become extinct, while the distribution of others has been materially
changed. The _Rytina_ of the North Pacific, the dodo of Mauritius, and the
great auk of the North Atlantic coasts, have been exterminated almost in
our own times. The kitchen-middens of Denmark contain remains of the
capercailzie, the _Bos primigenius_, and the beaver. The first still
abounds farther north, the second is extinct, and the third is becoming so
in Europe. The great Irish elk, a huge-antlered deer, probably existed
almost down to historic times.
_Pleistocene or Post-Pliocene Period._--We first meet with proofs of
important changes in the character of the European fauna, in studying the
remains found in the caverns of England and France, which have recently
been so well explored. These cave-remains are probably all subsequent to
the Glacial epoch, and they all come within the period of man's occupation
of the country. Yet we find clear proofs of two distinct kinds of change in
the forms of animal life. First we have a change clearly traceable to a
difference of climate. We find such arctic forms as the rein-deer, the
musk-sheep, the glutton, and the lemming, with the mammoth and the woolly
rhinoceros of the Siberian ice-cliffs, inhabiting this country and even the
south of France. This is held to be good proof that a sub-arctic climate
prevailed over all Central Europe; and this climate, together with the
continental condition of Britain, will sufficiently explain such a
southward range of what are now arctic forms.
But together with this change we have another that seems at first sight to
be in an exactly opposite direction. We meet with numerous animals which
now only inhabit Africa, or South Europe, or the warmer parts of Asia. Such
are, large felines--some closely related to the lion (_Felis spelæa_),
others of altogether extinct type (_Machairodus_) and forming the extreme
development of the feline race;--hyænas; horses of two or more species; and
a hippopotamus. If we go a little further back, to the remains furnished by
the gravels and brick-earths, we still find the same association of forms.
The reindeer, the glutton, {111}the musk-sheep, and the woolly rhinoceros,
are associated with several other species of rhinoceros and elephant; with
numerous civets, now abundant only in warm countries; and with antelopes of
several species. We also meet here with a great extension of range of forms
now limited to small areas. The Saiga antelope of Eastern Europe occurs in
France, where wild sheep and goats and the chamois were then found,
together with several species of deer, of bear, and of hyæna. A few extinct
genera even come down to this late period, such as the great sabre-toothed
tiger, _Machairodus_; _Galeotherium_, a form of Viverridæ; _Palæospalax_,
allied to the mole; and _Trogontherium_, a gigantic form of beaver,
We find then, that even at so early a stage of our inquiries we meet with a
problem in distribution by no means easy to solve. How are we to explain
the banishment from Europe in so short a space of time (geologically
speaking) of so many forms of life now characteristic of warmer countries,
and this too during a period when the climate of Central Europe was itself
becoming warmer? Such a change must almost certainly have been due to
changes of physical geography, which we shall be better able to understand
when we have examined the preceding Pliocene period. We may here notice,
however, that so far as we yet know, this great recent change in the
character of the fauna is confined to the western part of the Palæarctic
region. In caves in the Altai Mountains examined by Prof. Brandt, a great
collection of fossil bones was discovered. These comprised the Siberian
rhinoceros and mammoth, and the cave hyæna; but all the others, more than
thirty distinct species, are now living in or near the same regions. We may
perhaps impute this difference to the fact that the migration of Southern
types into this part of Siberia was prevented by the great mountain and
desert barrier of the Central Asiatic plateau; whereas in Europe there was
at this time a land connection with Africa. Post-pliocene deposits and
caverns in Algeria have yielded remains resembling the more southern
European types of the Post-pliocene period, but without any admixture of
Arctic forms; showing, as we might expect, that the glacial cold did not
{112}extend so far south. We have here remains of _Equus_, _Bos_,
_Antilope_, _Hippopotamus_, _Elephas_, _Rhinoceros_, _Ursus_, _Canis_, and
_Hyæna_, together with _Phacochoerus_, an African type of swine which has
not occurred in the European deposits.
It is perhaps to the earlier portion of this period that the
_Merycotherium_ of the Siberian drift belongs. This was an animal related
to the living camel, thus supporting the view that the _Camelidæ_ are
essentially denizens of the extra-tropical zone.
PLIOCENE PERIOD.
_Primates._--We here first meet with evidence of the existence of monkeys
in Central Europe. Species of _Macacus_ have left remains not only in the
Newer Pliocene of the Val d'Arno in Italy, but in beds of the same age at
Grays in Essex; while _Semnopithecus_ and _Cercopithecus_, genera now
confined to the Oriental and Ethiopian regions respectively, have been
found in the Pliocene deposits of the South of France and Italy.
_Carnivora._--Most of the genera which occurred in the Post-Pliocene are
found here also, and many of the same species. Few new forms appear, except
_Hyænarctos_, a large bear with characters approaching the hyænas, and
_Pristiphoca_, a new form of seal, both from the Older Pliocene of France;
and _Galecynus_, a fox-like animal intermediate between _Canis_ and
_Viverra_, from the Pliocene of Oeninghen in Switzerland.
_Cetacea._--Species of _Balæna_, _Physeter_, and _Delphinus_ occur in the
Older Pliocene of England and France, and with these the remains of many
extinct forms, _Balænodon_ and _Hoplocetus_ (Balænidæ); _Belemnoziphius_
and _Choneziphius_ (Hyperoodontidæ), and _Halitherium_, an extinct form of
the next order--Sirenia, now confined to the tropics, although the recently
extinct _Rytina_ of the N. W. Pacific shows that it is also adapted for
temperate climates.
_Ungulata._--The Pliocene deposits are not very rich in this order. The
horses (_Equidæ_) are represented by the genus _Equus_; and here we first
meet with _Hipparion_, in which small lateral toes appear. Both genera
occur in British deposits of this age. {113}A more interesting fact for us
is the occurrence of the genus _Tapirus_ in the Newer Pliocene of France
and in the older beds of both France and England, since this genus is now
isolated in the remotest parts of the eastern and western tropics. The
genera _Rhinoceros_, _Hippopotamus_, and _Sus_, occur here as in the
preceding epoch.
We next come to the deer genus (_Cervus_), which appears to have been at
its maximum in this period, no less than eight species occurring in the
Norwich Crag, and Forest-beds. Among the Bovidæ, the antelopes, ox, and
bison, are the only forms represented here, as in the Post-Pliocene period.
Passing on to the Proboscidea, we find three species of elephants and two
of _Mastodon_ preserved in European beds of this period, all distinct from
those of Post-Pliocene times.
_Rodentia._--In this order we find representatives of many living European
forms; as _Cricetus_ (hamster), _Arvicola_ (vole), _Castor_ (beaver),
_Arctomys_ (marmot), _Hystrix_ (porcupine), _Lepus_ (hare), and _Lagomys_
(pika); and a few that are extinct, the most important being _Chalicomys_,
allied to the beaver; and _Issiodromys_, said to come nearest to the
remarkable _Pedetes_ of South Africa, both found in the Pliocene formations
of France.
_General Conclusions as to Pliocene and Post-Pliocene Faunas of
Europe._--This completes the series of fossil forms of the Pliocene
deposits of Europe. They show us that the presence of numerous large
carnivora and ungulates (now almost wholly tropical) in the Post-Pliocene
period, was due to no exceptional or temporary cause, but was the result of
a natural succession from similar races which had inhabited the same
countries for long preceding ages. In order to understand the vast periods
of time covered by the Pliocene and Post-Pliocene formations, the works of
Sir Charles Lyell must be studied. We shall then come to see, that the
present condition of the fauna of Europe is wholly new and exceptional. For
a long succession of ages, various forms of monkeys, hyænas, lions, horses,
hipparions, tapirs, rhinoceroses, hippopotami, elephants, mastodons, deer,
and antelopes, together {114}with almost all the forms now living, produced
a rich and varied fauna such as we now see only in the open country of
tropical Africa. During all this period we have no reason to believe that
the climate or other physical conditions of Europe were more favourable to
the existence of these animals than now. We must look upon them, therefore,
as true indigenes of the country, and their comparatively recent extinction
or banishment as a remarkable phenomenon for which there must have been
some adequate cause. What this cause was we can only conjecture; but it
seems most probable that it was due to the combined action of the Glacial
period, and the subsidence of large areas of land once connecting Europe
with Africa. The existence, in the small island of Malta, of no less than
three extinct species of elephant (two of very small stature), of a
gigantic dormouse, an extinct hippopotamus, and other mammalia, together
with the occurrence of remains of hippopotamus in the caves of Gibraltar,
indicate very clearly that during the Pliocene epoch, and perhaps during a
considerable part of the Post-Pliocene, a connection existed between South
Europe and North Africa in at least these two localities. At the same time
we have every reason to believe that Britain was united to the Continent,
what is now the German Ocean constituting a great river-valley. During the
height of the Glacial epoch, these large animals would probably retire into
this Mediterranean land and into North Africa, making annual migrations
northwards during the summer. But as the connecting land sank and became
narrower and narrower, the migrating herds would diminish, and at last
cease altogether; and when the glacial cold had passed away would be
altogether prevented from returning to their former haunts.
MIOCENE PERIOD.
We now come to a period which was wonderfully rich in all forms of life,
and of which the geological record is exceptionally complete. Various
lacustrine, estuarine, and other deposits in Europe, North India, and North
America, have furnished such a {115}vast number of remains of extinct
mammalia, as to solve many zoological problems, and to throw great light on
the early distribution and centres of dispersal of various groups of
animals. In order to show the bearing of these remains on our special
subject, we will first give an account of the extinct fauna of Greece, of
the Upper Miocene period; since this, being nearest to Africa and Asia,
best exhibits the relations of the old European fauna to those countries.
We shall then pass to the Miocene fauna of France and Central Europe; and
conclude with the remarkable Siwalik and other Indian extinct faunas, which
throw an additional light on the early history of the animal life of the
great old-world continents.
_Extinct Animals of Greece._
These are from the Upper Miocene deposits at Pikermi, near Athens, and were
collected by M. Gaudry a few years ago. They comprise ten living and
eighteen extinct genera of mammalia, with a few birds and reptiles.
_Primates._--These are represented by _Mesopithecus_, a genus believed to
be intermediate between the two Indian genera of monkeys, _Semnopithecus_
and _Macacus_.
_Carnivora._--These were abundant. Of _Felis_ there were four species,
ranging from the size of a cat to that of a jaguar, a large _hyæna_, and a
large weasel (_Mustela_). Besides these there were the huge _Machairodus_,
larger than any existing lion or tiger, and with enormously developed
canine teeth; _Hyænictis_ and _Lycæna_, extinct forms of Hyænidæ;
_Thalassictis_=_Ictitherium_, an extinct genus of Viverridæ but with
resemblances to the hyænas, represented by three species, some of which
were larger than any existing Viverridæ; _Promephytis_, an extinct form of
Mustelidæ, having resemblances to the European marten, to the otters, and
to the S. African _Zorilla_; and lastly, _Simocyon_, an extraordinary
carnivore of the size of a small panther, but having the canines of a cat,
the molars of a dog, and the jaws shaped like those of a bear.
_Ungulata._--These are numerous and very _interesting_. The Equidæ are
represented by the three-toed _Hipparion_, which {116}continued to exist
till the Older Pliocene period. There are three large species of
_Rhinoceros_, as well as a species of the extinct genus _Leptodon_ of
smaller size. Remains of a very large wild boar (_Sus_) were found. Very
interesting is the occurrence of a species of giraffe (_Camelopardalis_) as
tall as the African species but more slender; and also an extinct genus
_Helladotherium_, not quite so tall as the giraffe but much more robust,
and showing some approach to the Antilopidæ in its dentition. Antelopes
were abundant, ranging from the size of the gazelle to that of the largest
living species. Three or four seem referable to living genera, but the
majority are of extinct types, and are classed in the genera _Palæotragus_,
_Palæoryx_, _Tragocerus_, and _Palæoreas_; while _Dremotherium_ is an
ancient generalized form of _Cervidæ_ or deer.
_Proboscidea._--These are represented by two species of _Mastodon_, and two
of _Dinotherium_, an extraordinary extinct form supposed to be, to some
extent, intermediate between the elephants and the aquatic manatees
(_Sirenia_.)
_Rodentia._--This order is represented by a species of _Hystrix_, larger
than living porcupines.
_Edentata._--This order, now almost confined to South America, was
represented in the Miocene period by several European species.
_Ancylotherium_ and _Macrotherium_, belonging to an extinct family but
remotely allied to the African ant-bear (_Orycteropus_), occur in Greece.
_Birds._--Species of _Phasianus_ and _Gallus_ were found; the latter
especially interesting as being now confined to India.
_Reptiles._--These are few and unimportant, consisting of a tortoise
(_Testudo_) and a large lizard allied to _Varanus_.
_Summary of the Miocene Fauna of Greece._--Although we cannot consider that
the preceding enumeration gives us by any means a complete view of the
actual inhabitants of this part of Europe during the later portion of the
Miocene period, we yet obtain some important information. The resemblance
that appeared in the Pliocene fauna of Europe, to that of the open country
of tropical Africa, is now still more remarkable. We {117}not only find
great felines, surpassing in size and destructive power the lions and
leopards of Africa, with hyænas of a size and in a variety not to be
equalled now, but also huge rhinoceroses and elephants, two forms of
giraffes, and a host of antelopes, which, from the sample here obtained,
were probably quite as numerous and varied as they now are in Africa.
Joined with this abundance of antelopes we have the absence of deer, which
probably indicates that the country was open and somewhat of a desert
character, since there were deer in other parts of Europe at this epoch.
The occurrence of but a single species of monkey is also favourable to this
view, since a well-wooded country would most likely have supplied many
forms of these animals.
_Miocene Fauna of Central and Western Europe._
We have now to consider the Miocene fauna of Europe generally, of which we
have very full information from numerous deposits of this age in France,
Switzerland, Italy, Germany, and Hungary.
_Primates._--Three distinct forms of monkeys have been found in Europe--in
the South of France, in Switzerland, and Wurtemberg; one was very like
_Colobus_ or _Semnopithecus_; the others--_Pliopithecus_ and
_Dryopithecus_--were of higher type, and belonged to the anthropomorphous
apes, being nearest to the genus _Hylobates_ or gibbons. Both have occurred
in the South of France. The _Dryopithecus_ was a very large animal (equal
to the gorilla), and M. Lartet considers that in the character of its
dentition it approached nearer to man than any of the existing anthropoid
apes.
_Insectivora._--These small animals are represented by numerous remains
belonging to four families and a dozen genera. Of _Erinaceus_ (hedgehog)
several species are found in the Upper Miocene; and in the Lower Miocene of
Auvergne two extinct genera of the same family--_Amphechinus_ and
_Tetracus_--have been discovered. Several species of _Talpa_ (mole) occur
in the Upper Miocene of France, while the extinct _Dinylus_ is from
Germany, and _Palæospalax_ from the Lower Miocene of the Isle of
{118}Wight. The Malayan family Tupaiidæ or squirrel-shrews, is believed to
be represented by _Oxygomphus_, a fossil discovered in South Germany
(Wiesenau) by H. von Meyer. The Soricidæ or shrews, are represented by
several extinct genera--_Plesiosorex_, _Mysarachne_ and _Galeospalax_; as
well as by _Amphisorex_ and _Myogale_ still living. _Echinogale_, a genus
of Centetidæ now confined to Madagascar, is said to occur in the Lower
Miocene of Auvergne, a most interesting determination, if correct, as it
would form a transition to the _Solenodon_ of the Antilles belonging to the
same family; but I am informed by Prof. Flower that the affinities of the
animals described under this name are very doubtful.
_Carnivora._--Besides _Felis_ and _Machairodus_, which extend back to the
Upper Miocene, there are two other genera of Felidæ, _Pseudælurus_ in the
Upper Miocene of France, and _Hyænodon_, which occurs in the Upper and
Lower Miocene of France, named from some resemblance in its teeth to the
hyænas, and considered by some Palæontologists to form a distinct family,
Hyænodontidæ. The Viverridæ, or civets, were very numerous, consisting of
the living genus _Viverra_, and three extinct
forms--_Thalassictis_=_Ictitherium_, as large as a panther, and
_Soricictis_, a smaller form, occurring both in France and Hungary. Of
_Hyænidæ_, there was the living genus _Hyæna_, and the extinct _Hyænictis_,
which has occurred in Hungary as well as in Greece. The Canidæ, or wolf and
fox family, were represented by _Pseudocyon_, near to _Canis_; _Hemicyon_,
intermediate between dogs and gluttons; and _Amphicyon_, of which several
species occur in the Upper and Lower Miocene of France, some of them larger
than a tiger. The Mustelidæ, or weasels, were represented by five genera,
the existing genera _Lutra_ (otter) and _Mustela_ (weasel);
_Potamotherium_, an extinct form of otter; _Taxodon_, allied to the badger
and otter; _Palæomephitis_ in Germany, and the _Promephytis_ (already
noticed) in Greece. The bears were represented only by _Hyænarctos_, which
has been noticed as occurring in the Pliocene, and first appears in the
Upper Miocene of France. Seals are represented by a form resembling the
Antarctic _Otaria_, remains of which occur in the Upper Miocene of France.
{119}_Cetacea_ (whales).--These occur frequently in the Miocene deposits,
four living, and five extinct genera having been described; but these
marine forms are not of much importance for our purpose.
_Sirenia_ (sea-cows).--These are represented by two extinct genera,
_Halitherium_ and _Trachytherium_. Several species of the former have been
discovered, but the latter has occurred in France only, and its affinities
are doubtful.
_Ungulata._--Horses are represented by _Hipparion_ and _Anchitherium_, the
latter occurring in both Upper and Lower Miocene and Eocene; while
_Hipparion_, which is more nearly allied to living horses, first appears in
the Upper Miocene and continues in the Pliocene.
_Hippotherium_, in the Upper Miocene of the Vienna basin, forms a
transition to _Paloplotherium_, an Eocene genus of Tapiridæ or
Palæotheridæ. Tapirs, allied to living forms, occur in both Upper and Lower
Miocene. Rhinoceroses are still found in the Upper Miocene, and here first
appear the four-toed hornless rhinoceros, _Acerotherium_. The Suidæ (swine)
are rather numerous. _Sus_ (wild boar) continued as far back as the Upper
Miocene; but now there first appear a number of extinct forms which have
been named _Hyotherium_, _Palæochoerus_, _Choeromorus_, all of a small or
moderate size; _Hyopotamus_, nearly as large as a tapir; and
_Anthracotherium_, nearly the size of a hippopotamus and, according to Dr.
Leidy, the type of a distinct family. _Listriodon_, from the Upper Miocene
of the Vienna basin, is sometimes classed with the tapirs.
We now come to a well-marked new family of Artiodactyle or even-toed
Ungulata, the _Anoplotheriidæ_, which consisted of more slender long-tailed
animals, allied to the swine but with indications of a transition towards
the camels. The only genera that appear in the Miocene formation are,
_Chalicotherium_, nearly as large as a rhinoceros, of which three species
have been found in Germany and France; and _Synaphodus_, known only from
its teeth, which differ somewhat from those of the _Anoplotherium_ which
appears earlier in the Eocene formation. Another extinct family,
_Amphimericidæ_ or _Xiphodontidæ_, is represented by two {120}genera,
_Cainotherium_ and _Microtherium_, in the Miocene of France. They were of
very small size, and are supposed to be intermediate between the Suidæ and
Tragulidæ.
The Camelopardalidæ, or giraffes, were represented in Europe in Miocene
times by the gigantic _Helladotherium_, which has been found in the south
of France, and in Hungary, as well as in Greece. The chevrotains
(Tragulidæ) are represented by the extinct genus _Hyomoschus_.
The Cervidæ do not seem to have appeared in Europe before the Upper Miocene
epoch, when they were represented by _Dorcatherium_ and _Amphimoschus_,
allied to _Moschus_, and also by true _Cervus_, as well as by small allied
forms, _Dremotherium_, _Amphitragalus_ (in the Lower Miocene),
_Micromeryx_, _Palæomeryx_, and _Dicrocerus_.
The Bovidæ, or hollow-horned ruminants, were not well represented in
Central Europe in Miocene times. There were no sheep, goats, or oxen, and
only a few antelopes of the genus _Tragocerus_, and one allied to
_Hippotragus_; and these all lived in the Upper Miocene period, as did the
more numerous forms of Greece.
_Proboscidea._--The true elephants do not extend back to the Miocene
period, but they are represented by the Mastodons, which had less complex
teeth. These first appear in the Upper Miocene of Europe, five species
being known from France, Germany, Switzerland, and Greece. _Dinotherium_,
already noticed as occurring in Greece, extended also to Germany and
France, where remains of three species have been found.
_Rodentia._--A considerable number of generic forms of this order have been
obtained from the Miocene strata. The principal genera are _Cricetodon_,
allied to the hamsters, numerous in both the Upper and Lower Miocene period
of France; _Myoxus_ (the dormice) in France, and an allied genus,
_Brachymys_, in Germany. The beavers were represented by the still living
genus _Castor_, and the extinct _Steneofiber_ in France. The squirrels by
the existing _Scuirus_ and _Spermophilus_; and by extinct forms, _Lithomys_
and _Aulacodon_, in Germany, the latter resembling the African genus
_Aulacodes_. The hares, by _Lagomys_ and an {121}extinct form _Titanomys_.
Besides these, remains referred to the South American genera, _Cavia_
(cavy) and _Dasyprocta_ (agouti), have been found, the former in the Upper
Miocene of Switzerland, the latter in the Lower Miocene of Auvergne.
_Palæomys_, allied to the West Indian _Capromys_, has been found in the
same deposits; as well as _Theridomys_, said by Gervais to be allied to
_Anomalurus_ and _Echimys_, the former now living in W. Africa, the latter
in S. America.
_Edentata._--These are only represented by the _Macrotherium_ and
_Ancylotherium_ of the Grecian deposits, the former occurring also in
France and Germany in Upper Miocene strata.
_Marsupials_.--These consist of numerous species related to the opossums
(_Didelphys_), but separated by Gervais under the name _Peratherium_. They
occur in both Upper and Lower Miocene beds.
_Upper Miocene Deposits of the Siwalik Hills and other Localities in N. W.
India._
These remarkable fresh-water deposits form a range of hills at the foot of
the Himalayas, a little south of Simla. They were investigated for many
years by Sir P. Cautley and Dr. Falconer, and add greatly to our knowledge
of the early fauna of the Old World continent.
_Primates_.--Remains of the genera _Semnopithecus_ and _Macacus_ were
found, with other forms of intermediate character; and some teeth indicated
animals allied to the orang-utan of Borneo, and of similar size.
_Carnivora_.--These consisted of species of _Felis_ and _Machairodus_ of
large size; _Hyæna_, _Canis_, _Mellivora_, and an allied genus _Ursitaxus_;
_Ursus_, in the deposits of the Nerbudda valley (of Pliocene age);
_Hyænarctos_ as large as the cave bear; _Amphicyon_ of the size of a polar
bear (in the deposits of the Indus valley, west of Cashmere); _Lutra_, and
an extinct allied genus _Enhydrion_.
_Ungulata_.--These are very numerous, and constitute the most important
feature of this ancient fauna. Horses are represented by a species of
_Equus_ from the Siwalik Hills and the Irawaddy {122}deposits in Burmah,
and by two others from the Pliocene of the Nerbudda Valley; while
_Hippotherium_--a slender, antelope-like animal, found in the Siwalik Hills
and in Europe--is supposed to form a transition from the Equidæ to the
Tapiridæ. These latter are found in the Upper Indus deposits, where there
is a species of _Tapirus_, and one of an extinct genus _Antelotherium_. Of
_Rhinoceros_, five extinct species have been found--in the Siwalik Hills,
in Perim Island, and one at an elevation of 16,000 feet in the deserts of
Thibet. _Hippopotamus_ occurs in the Pliocene of the Nerbudda, and is
represented in the older Miocene deposits by _Hexaprotodon_, of which three
species have been found in various parts of India. Another remarkable
genus, _Merycopotamus_, connects _Hippopotamus_ with _Anthracotherium_, one
of the extinct European forms allied to the swine. These last are
represented by several large species of _Sus_, and by the extinct European
genus _Choerotherium_.
The extinct Anoplotheridæ are represented by a species of the European
genus _Chalicotherium_, larger than a horse.
An extinct camel, larger than the living species, was found in the Siwalik
Hills.
Three species of deer (_Cervus_) have been found in the Siwaliks, and one
in the Nerbudda deposits.
A large and a small species of giraffe (_Camelopardalis_) were found in the
Siwalik Hills and at Perim Island.
The Bovidæ are represented by numerous species of _Bos_, and by the extinct
genera _Hemibos_ and _Amphibos_. There are also three species of antelopes,
one of which is allied to the African _Alcephalus_.
We now come to an extraordinary group of extinct animals, probably forming
a new family intermediate between the antelope and the giraffe. The
_Sivatherium_ was an enormous four-horned ruminant, larger than a
rhinoceros. It had a short trunk like a tapir, the lower horns on the
forehead were simple, the upper pair palmated. The _Bramatherium_, an
allied form from Perim Island, showed somewhat more affinity for the
giraffe.
_Proboscidea._--No less than seven species of elephants and four {123}of
mastodons ranged over India, their remains being found in all the deposits
from the Siwalik Hills to Burmah. A large _Dinotherium_ has also been found
at Perim Island.
_Reptiles._--Many remains of birds were found, but these have not been
determined. Reptiles were numerous and interesting, the most remarkable
being the huge tortoise, _Colossochelys_, whose shell was twelve feet long
and head and neck eight feet more. Other small tortoises of the genera
_Testudo_, _Emys_, _Trionyx_ and _Emydida_ were found, the Emys being a
living species. There were three extinct and one living species of
crocodile, and one of them was larger than any now living. The only other
reptile of importance was a large lizard of the genus _Varanus_.
_General Observations on the Miocene faunas of Europe and Asia._--Comparing
the three faunas of approximately the same period, and allowing for the
necessarily imperfect record of each, we find a wonderful similarity of
general type over the enormous area between France on the west and the
Irawaddy river in Burmah on the east. We may even extend our comparison to
Northern China, where remains of _Hyæna_, _Tapir_, _Rhinoceros_,
_Chalicotherium_, and _Elephas_, have been recently found, closely
resembling those from the Miocene or Pliocene deposits of Europe or India,
and showing that the Palæarctic region had then the same great extent from
west to east that it has now. Of about forty genera comprised in the Indian
Miocene fauna, no less than twenty-seven inhabited Central and Western
Europe during the same epoch. The Indian Miocene fossils are much what we
should expect as the forerunners of the existing fauna, the giraffes and
hippopotami being the only additions from the present Ethiopian fauna. The
numerous forms of the restricted bovine type, show that these probably
originated in India; while the monkeys appear to be altogether of Oriental
types.
In Europe, however, we meet with a totally different assemblage of animals
from those that form the existing fauna. We find apes and monkeys, many
large Felidæ, numerous civets {124}and hyænas, tapirs, rhinoceros,
hippopotamus, elephants, giraffes, and antelopes, such as now characterise
the tropics of Africa and Asia. Along with these we meet with less familiar
types, showing relations with the Centetidæ of Madagascar, the Tupaiidæ of
the Malay Islands, the _Capromys_, of the West Indies, and the _Echimys_ of
South America. And besides all these living types we have a host of extinct
forms,--ten or twelve genera allied to swine; nine genera of tapir-like
animals; four of horses; nine of wolves; with many distinct forms of the
long-extinct families of Anoplotheridæ, Xiphodontidæ, and the edentate
Macrotheridæ. It is almost certain that during the Miocene period Europe
was not only far richer than it is now in the higher forms of life, but not
improbably richer than any part of the globe now is, not excepting tropical
Africa and tropical Asia.
EOCENE PERIOD.
The deposits of Eocene age are less numerous, and spread over a far more
limited area, than those of the Miocene period, and only restricted
portions of them furnish any remains of land animals. Our knowledge of the
Eocene mammalian fauna is therefore very imperfect and will not occupy us
long, as most of the new types it furnishes are of more interest to the
zoologist than to the student of distribution. Some of the Eocene mammalia
of Europe are, however, of interest in comparison with those of North
America of the same age; while others show that ancestral types of groups
now confined to Australia or to South America, then inhabited Europe.
_Primates._--The only undoubted Eocene examples of this order, are the
_Cænopithecus lemuroides_ from the Jura, which has points of resemblance to
the South American marmosets and howlers, and also to the Lemuridæ; and a
cranium recently discovered in the Department of Lot (S.W. France),
undoubtedly belonging to the Lemuridæ, and which most resembles that of the
West African "Potto" (_Perodicticus_). This discovery has led to another,
for it is now believed that remains formerly {125}referred to the
Anoplotheridæ (_Adapis_ and _Aphelotherium_ from the Upper Eocene of Paris)
were also Lemurs. Some remains from the Lower Eocene of Suffolk were at
first supposed to be allied to _Macacus_, but were subsequently referred to
the Ungulate, _Hyracotherium_. There is still, however, some doubt as to
its true affinities.
_Chiroptera._--In the Upper Eocene of Paris remains of bats have been
found, so closely resembling living forms as to be referred to the genus
_Vespertilio_.
_Carnivora._--The only feline remains, are those of _Hyænodon_ in the Upper
Eocene of Hampshire, and _Pterodon_, an allied form from beds of the same
age in France; with _Ælurogale_, found in the South of France in deposits
of phosphate of lime of uncertain age, but probably belonging to this
period. Viverridæ (civets) are represented by two genera, _Tylodon_, the
size of a glutton from the Upper Eocene, and _Palæonyctis_, allied to
_Viverra_, from the Middle Eocene of France. The Canidæ (wolves and foxes)
appear to have been the most ancient of the existing types of Carnivora,
five genera being represented by Eocene remains. Of these, _Galethylax_ and
_Cyotherium_ were small, and with the existing genus _Canis_ are found in
the Upper Eocene of France. _Arctocyon_, about the size of a wolf, is a
very ancient and generalised form of carnivore which can not be placed in
any existing family. It is found in the Lower Eocene of France, and is thus
the oldest known member of the Carnivora.
_Ungulata._--These are more numerous. Equidæ (horses) are represented by
the Miocene _Anchitherium_ in the Lower, and by a more ancient form,
_Anchilophus_, in the Middle Eocene of France. Tapiridæ and Palæotheridæ
were very numerous. _Palæotherium_ and the allied genus _Paloplotherium_,
were abundant in France and England in Upper Eocene times. They somewhat
resembled the tapir, with affinities for the horse and rhinoceros. A new
genus, _Cadurcotherium_, allied to the rhinoceros and equally large, has
been found in the same deposits of phosphate of lime as the lemur and
_Ælurogale_. In the Middle Eocene of both England and France are found
_Lophiodon_ allied to the tapir, {126}but in some of the species reaching a
larger size; _Propalæotherium_ and _Pachynolophus_ of smaller size and
having affinities for the other genera named; and _Plagiolophus_, a small,
slender animal which Professor Huxley thinks may have been a direct
ancestor of the horse. In the Lower Eocene we meet with _Coryphodon_, much
larger than the tapir, and armed with large canine teeth; _Pliolophus_, a
generalised type, allied to the tapir and horse; and _Hyracotherium_, a
small animal from the Lower Eocene of England, remotely allied to the
tapir.
Among the Artiodactyla, or even-toed ungulates, the swine are represented
by several extinct genera, of moderate or small size--_Acotherium_,
_Choeropotamus_, _Cebochoerus_ and _Dichobune_, all from the Upper and the
last also from the Middle Eocene of France; but _Eutelodon_, from the
phosphate of lime deposits is large. The _Dichobune_ was the most
generalised type, presenting the characters of many of the other genera
combined, and was believed by Dr. Falconer to approach the musk-deer. The
_Cainotherium_ of the Miocene also occurs here, and an allied genus
_Plesiomeryx_ from the same deposits as _Euteledon_.
The Eocene Anoplotheridæ were numerous. The _Anoplotherium_ was a two-toed,
long-tailed Pachyderm, ranging from the size of a hog to that of an ass;
the allied _Eurytherium_ was four-toed; and there are one or two others of
doubtful affinity. All are from the Upper Eocene of France and England.
_Rodentia._--Remains referred to the genera _Myoxus_ (dormouse) and
_Sciurus_ (squirrel) have been found in the Upper Eocene of France; as well
as _Plesiarctomys_, an extinct genus between the marmots and squirrels. The
Miocene _Theridomys_ is also found here.
_Marsupials._--The _Didelphys_ (opossum) of Cuvier, now referred to an
extinct genus _Peratherium_, is found in the Upper Eocene of France and
England.
_General Considerations on the Extinct Mammalian Fauna of Europe._--It is a
curious fact that no family, and hardly a genus, of European mammalia
occurs in the Pliocene deposits, without extending back also into those of
Miocene age. There are, {127}however, a few groups which, seem to be late
developments or recent importations into the Palæarctic region, as they
occur only in Post-Pliocene deposits. The most important of these are the
badger, glutton, elk, reindeer, chamois, goat, and sheep, which only occur
in caves and other deposits of Post-Pliocene age. Camels only occur in the
Post-Pliocene of Siberia (_Merycotherium_), although a true _Camelus_ of
large size appears to have inhabited some part of Central Asia in the Upper
Miocene period, being found in the Siwalik beds. The only exclusively
Pliocene genera in Europe are _Ursus_, _Equus_, _Hippopotamus_, _Bos_,
_Elephas_, _Arvicola_, _Trogontherium_, _Arctomys_, _Hystrix_ and _Lepus_;
but of these _Equus_, _Hippopotamus_, _Bos_, and _Elephas_ are found in the
Miocene deposits of India. Owing, no doubt, in part to the superior
productiveness of the various Miocene beds, large numbers of groups appear
to have their origin or earliest appearance here. Such are Insectivora,
Felidæ, Hyænidæ, Mustelidæ, _Ursus_, Equidæ, _Tapirus_, Rhinocerotidæ,
Hippopotamidæ, Anthracotheridæ (extinct), _Sus_, Camelopardidæ, Tragulidæ,
Cervidæ, Bovidæ, Elephantidæ, and Edentata.
Groups which go back to the Eocene period, are, Primates allied to South
American monkeys, as well as some of the Lemuridæ; bats of the living genus
_Vespertilio_; Hyænodontidæ, an ancestral form of Carnivore; Viverridæ;
Canidæ (to the Upper Eocene), and the ancestral Arctocyonidæ to the Lower
Eocene; _Hyænarctos_, an ancestral type of bears and hyænas; Anchitheridæ,
ancestral horses, to the Middle Eocene; Palæotheridæ, comprising numerous
generalised forms, ancestors of the rhinoceros, horse, and tapir; Suidæ,
with numerous generalised forms, to the Middle Eocene; Anoplotheridæ and
Xiphodontidæ, ancestral families of even-toed Ungulates, connecting the
ruminants with the swine; and lastly, several groups of Rodents, and a
Marsupial, in the Upper Eocene. We thus find all the great types of
Mammalia well developed in the earliest portion of the tertiary period; and
the occurrence of Quadrumana, of the highly specialized bats
(_Vespertilio_), of various forms of Carnivora, and of Ungulates, clearly
differentiated into the odd and even-toed series, associated with such
lower forms as {128}Lemurs and Marsupials--proves, that we have here hardly
made an approach towards the epoch when the mammalian type itself began to
diverge into its various modifications. Some of the Carnivora and Ungulates
do, indeed, exhibit a less specialised structure than later forms; yet so
far back as the Upper Miocene the most specialised of all carnivora, the
great sabre-toothed _Machairodus_, makes its appearance.
The Miocene is, for our special study, the most valuable and instructive of
the Tertiary periods, both on account of its superior richness, and because
we here meet with many types now confined to separate regions. Such facts
as the occurrence in Europe during this period of hippopotami, tapirs,
giraffes, Tragulidæ, Edentata, and Marsupials--will assist us in solving
many of the problems we shall meet with in reviewing the actual
distribution of living forms of those groups. Still more light will,
however, be thrown on the subject by the fossil forms of the American
continent, which we will now proceed to examine.
{129}CHAPTER VII.
EXTINCT MAMMALIA OF THE NEW WORLD.
The discoveries of very rich deposits of mammalian remains in various parts
of the United States have thrown great light on the relations of the faunas
of very distant regions. North America now makes a near approach to Europe
in the number and variety of its extinct mammalia, and in no part of the
world have such perfect specimens been discovered. In what are called the
"Mauvaises terres" of Nebraska (the dried-up mud of an ancient lake),
thousands of entire crania and some almost entire skeletons of ancient
animals have been found, their teeth absolutely perfect, and altogether
more resembling the preparations of the anatomist, than time-worn fossils
such as we are accustomed to see in the museums of Europe. Other deposits
have been discovered in Oregon, California, Virginia, South Carolina,
Texas, and Utah, ranging over all the Tertiary epochs, from Post-Pliocene
to Eocene, and furnishing a remarkable picture of the numerous strange
mammalia which inhabited the ancient North American continent.
NORTH AMERICA--POST-PLIOCENE PERIOD.
_Insectivora._--The only indications of this order yet discovered, consists
of a single tooth of some insectivorous animal found in Illinois, but which
cannot be referred to any known group.
_Carnivora._--These are fairly represented. Two species of _Felis_ as large
as a lion; the equally large extinct _Trucifelis_, found only in Texas;
four species of _Canis_, some of them larger {130}than wolves; two species
of _Galera_, a genus now confined to the Neotropical region; two bears, and
an extinct genus, _Arctodus_; an extinct species of racoon (_Procyon_), and
an allied extinct genus, _Myxophagus_--show, that at a very recent period
North America was better supplied with Carnivora than it is now. Remains of
the walrus (_Trichechus_) have also been found as far south as Virginia.
_Cetacea._--Three species of dolphins belonging to existing genera, have
been found in the Eastern States; and two species of _Manatus_, or sea-cow,
in Florida and South Carolina.
_Ungulata._--Six extinct horses (_Equus_), and one _Hipparion_; the living
South American tapir, and a larger extinct species; a _Dicotyles_, or
peccary, and an allied genus, _Platygonus_; a species of the South American
llamas (_Auchenia_), and one of a kind of camel, _Procamelus_; two extinct
bisons; a sheep, and two musk-sheep (_Ovibos_); with three living and one
extinct deer (_Cervus_), show an important increase in its Herbivora.
_Proboscidea._--Two elephants and two mastodons, added to this remarkable
assemblage of large vegetable-feeding quadrupeds.
_Rodentia._--These consist mainly of genera and species still living in
North America; the only important exceptions being a species of the South
American capybara (_Hydrochoerus_) in South Carolina; and _Praotherium_, an
extinct form of hare, found in a bone cave in Pennsylvania.
_Edentata._--Here we meet with a wonderful assemblage, of six species
belonging to four extinct genera, mostly of gigantic size. A species of
_Megatherium_, three of _Megalonyx_, and one of _Mylodon_--huge terrestrial
sloths as large as the rhinoceros or even as the largest elephants--ranged
over the Southern States to Pennsylvania, the latter (_Mylodon_) going as
far as the great lakes and Oregon. Another form, _Ereptodon_, has been
found in the Mississippi Valley.
_Marsupialia._--The living American genus of opossums, _Didelphys_, has
been found in deposits of this age in South Carolina.
_Remarks on the Post-Pliocene fauna of North America._--The assemblage of
animals proved, by these remains, to have {131}inhabited North America at a
comparatively recent epoch, is most remarkable. In Europe, we found a
striking change in the fauna at the same period; but that consisted almost
wholly in the presence of animals now inhabiting countries immediately to
the north or south. Here we have the appearance of two new assemblages of
animals, the one now confined to the Old World--horses, camels, and
elephants; the other exclusively of South American type--llamas, tapirs,
capybaras, _Galera_, and gigantic Edentata. The age of the various deposits
in which these remains are found is somewhat uncertain, and probably
extends over a considerable period of time, inclusive of the Glacial epoch,
and perhaps both anterior and subsequent to it. We have here, as in Europe,
the presence and apparent co-existence in the same area, of Arctic and
Southern forms--the walrus and the manatee--the musk-sheep and the gigantic
sloths. Unfortunately, as we shall see, the immediately preceding Pliocene
deposits of North America are rather poor in organic remains; yet it can
hardly be owing to the imperfection of the record of this period, that _not
one_ of the South American types above numerated occurs there, while a
considerable number of Old World forms are represented. Neither in the
preceding wonderfully rich Miocene or Eocene periods, does any _one_ of
these forms occur; or, with the exception of _Morotherium_, from Pliocene
deposits _west_ of the Rocky Mountains, any apparent ancestor of them! We
have here unmistakable evidence of an extensive immigration from South into
North America, not very long before the beginning of the Glacial epoch. It
was an immigration of types altogether new to the country, which spread
over all the southern and central portions of it, and established
themselves sufficiently to leave abundance of remains in the few detached
localities where they have been discovered. How such large yet defenceless
animals as tapirs and great terrestrial sloths, could have made their way
into a country abounding in large felines equal in size and destructiveness
to the lion and the tiger, with numerous wolves and bears of the largest
size, is a great mystery. But it is nevertheless certain that they did so;
and the fact that no such {132}migration had occurred for countless
preceding ages, proves that some great barrier to the entrance of
terrestrial mammalia which had previously existed, must for a time have
been removed. We must defer further discussion of this subject till we have
examined the relations of the existing faunas of North and South America.
TERTIARY PERIOD.
When we get to remains of the Tertiary age, especially those of the Miocene
and Eocene epochs, we meet with so many interesting and connected types,
and such curious relations with living forms in Europe, that it will be
clearer to trace the history of each order and family throughout the
Tertiary period, instead of considering each of the subdivisions of that
period separately.
It will be well however first to note the few American Post-Pliocene or
living genera that are found in the Pliocene beds. These consist of several
species of _Canis_, from the size of a fox to that of a large wolf; a
_Felis_ as large as a tiger; an Otter (_Lutra_); several species of
_Hipparion_; a peccary (_Dicotyles_); a deer (_Cervus_); several species of
_Procamelus_; a mastodon; an elephant; and a beaver (_Castor_). It thus
appears that out of nearly forty genera found in the Post-Pliocene
deposits, only ten are found in the preceding Pliocene period. About twelve
additional genera, however, appear there, as we shall see in going over the
various orders.
_Primates._--Among the vast number of extinct mammalia discovered in the
Tertiary deposits of North America, no example of this order had been
recognized up to 1872, when the discovery of more perfect remains showed,
that a number of small animals of obscure affinities from the Lower Eocene
of Wyoming, were really allied to the lemurs and perhaps also to the
marmosets, the lowest form of American monkeys, but having a larger number
of teeth than either. A number of other remains of small animals from the
same formation, previously supposed to be allied to the Ungulata, are now
shown to {133}belong to the Primates; so that no less than twelve genera of
these animals are recognized by Mr. Marsh, who classes them in two
families--Limnotheridæ, comprising the genera _Limnotherium_, (which had
larger canine teeth), _Thinolestes_, _Telmatolestes_, _Mesacodon_,
_Bathrodon_, and _Antiacodon_ of Marsh, with _Notharctos_, _Hipposyus_,
_Microsyops_, and _Palæacodon_ previously described by Leidy;--and
Lemuravidæ, consisting of the genera _Lemuravus_ (Marsh) and _Hyopsodus_
(Leidy). The animals of the latter family were most allied to existing
lemurs, but were a more generalized form, _Lemuravus_ having forty-four
teeth, the greatest number known in the order. These numerous forms ranged
from the size of a small squirrel to that of a racoon. It is especially
interesting to find these peculiar lemuroid forms in America, just when a
lemur has been discovered of about the same age in Europe; and as the
American forms are said to show an affinity with the South American
marmosets, while the European animal is most allied to a West African
group, we have evidently not yet got back far enough to find the primeval
or ancestral type from which all the Primates sprang.
About the same time, in the succeeding Miocene formation, true monkeys were
discovered. Mr. Marsh describes _Laopithecus_ as an animal nearly the size
of the largest South American monkeys, and allied both to the Cebidæ and
the Eocene Limnotheridæ. Mr. Cope has described _Menotherium_ from the
Miocene of Colorado, as a lemuroid animal, the size of a cat, and perhaps
allied to _Limnotherium_. More Miocene remains will, no doubt, be
discovered, by which we shall be enabled to trace the origin of some of the
existing forms of South American monkeys; and perhaps help to decide the
question (now in dispute among anatomists) whether the lemurs are really
Primates, or form an altogether distinct and isolated order of mammalia.
_Insectivora._--This order is represented by comparatively few forms in the
tertiary beds, and these are all very different from existing types. In the
Upper Miocene of Dakota are found remains indicating two extinct genera,
_Lepictis_ and _Ictops_. In the Miocene of Colorado, Professor Cope has
recently discovered four new genera, _Isacis_; allied to the preceding, but
as large as a {134}_Mephitis_ or skunk; _Herpetotherium_, near the moles;
_Embasis_, more allied to the shrews; and _Dommina_, of uncertain
affinities. Two others have been found in the Eocene of Wyoming; _Amomys_,
having some resemblance to hedgehogs and to the Eastern _Tupaia_; and
_Washakius_, of doubtful affinities.
Far back in the Triassic coal of North Carolina has been found the jaw of a
small mammal (_Dromotherium_), the teeth of which somewhat resemble those
of the Australian _Myrmecobius_, and may belong either to the Insectivora
or Marsupials; if indeed, at that early period these orders were
differentiated.
_Carnivora._--The most ancient forms of this order are some remains found
in the Middle Eocene of Wyoming, and others recently described by Professor
Cope (1875) from the Eocene of New Mexico, of perhaps earlier date. The
former consist of three genera, _Patriofelis_, _Uintacyon_, and
_Sinopa_,--animals of large size but which cannot be classed in any
existing family; and two others, _Mesonyx_ and _Synoplotherium_, believed
by Mr. Cope to be allied to _Hyænodon_. The latter consist of four
genera,-- _Oxyæna_, consisting of several species, some as large as a
jaguar, was allied to _Hyænodon_ and _Pterodon_; _Pachyæna_, allied to the
last; _Prototomus_, allied to _Amphicyon_ and the Viverridæ; and
_Limnocyon_, a civet-like carnivore with resemblances to the Canidæ.
In the Miocene formations we find the Feline type well developed. The
wonderful _Machairodus_, which in Europe lived down to Post-Pliocene times,
is found in the Upper Miocene of Dakota; and perfect crania have been
discovered, showing that the chin was lengthened downwards to receive and
protect the enormous canines. _Dinyctis_ was allied both to _Machairodus_
and to the weasels. Three new genera have been lately described by
Professor Cope from the Miocene of Colorado,--_Bunælurus_, with characters
of both cats and weasels; _Daptophilus_, allied to _Dinyctis_; and
_Hoplophoneus_, more allied to _Machairodus_. The Canidæ are represented by
_Amphicyon_, which occurs in deposits of the same age in Europe; and by
_Canis_, four species of which genus are recorded by Professor Cope from
the Miocene of Colorado, and it also occurs in the Pliocene. The _Hyænodon_
is represented by three species in the Miocene of Dakota and Colorado. It
occurs {135}also in the European Miocene and Upper Eocene formations, and
constitutes a distinct family Hyænodontidæ, allied, according to Dr. Leidy,
to wolves, cats, hyænas and weasels. The Ursidæ are represented by only one
species of an extinct genus, _Leptarchus_, from the Pliocene of Nebraska.
From the Pliocene of Colorado, Prof. Cope has recently described
_Tomarctos_, as a "short-faced type of dog;" well as species of _Canis_ and
_Martes_.
_Ungulata._--The animals belonging to this order being usually of large
size and accustomed to feed and travel in herds, are liable to wholesale
destruction by floods, bogs, precipices, drought or hunger. It is for these
reasons, probably, that their remains are almost always more numerous than
those of other orders of mammalia. In America they are especially abundant;
and the number of new and intermediate types about whose position there is
much difference of opinion among Palæontologists, renders it very difficult
to give a connected summary of them with any approach to systematic
accuracy.
Beginning with the Perissodactyla, or odd-toed ungulates, we find the
Equine animals remarkably numerous and interesting. The true horses of the
genus _Equus_, so abundant in the Post-Pliocene formations, are represented
in the Pliocene by several ancestral forms. The most nearly allied to
_Equus_ is _Pliohippus_, consisting of animals about the size of an ass,
with the lateral toes not externally developed, but with some differences
of dentition. Next come _Protohippus_ and _Hipparion_, in which the lateral
toes are developed but are small and functionless. Then we have the allied
genera, _Anchippus_, _Merychippus_, and _Hyohippus_, related to the
European _Hippotherium_, which were all still smaller animals,
_Protohippus_ being only 2½ feet high. In the older deposits we come to a
series of forms, still unmistakably equine, but with three or more toes
used for locomotion and with numerous differentiations in form,
proportions, and dentition. These constitute the family Anchitheridæ. In
the Miocene we have the genera _Anchitherium_ (found also in the European
Miocene), _Miohippus_ and _Mesohippus_, all with three toes on each foot,
and about the size of a sheep or large goat. In the Eocene of {136}Utah and
Wyoming, we get a step further back, several species having been discovered
about the size of a fox with four toes in front and three behind. These
form the genus _Orohippus_, and are the oldest ancestral horse known. Prof.
Marsh points out the remarkably perfect series of forms in America, which,
beginning with this minute ancient type, is gradually modified by gaining
increased size, increased speed by concentration of the limb-bones,
elongation of the head and neck, the canine teeth decreased in size, the
molars becoming longer and being coated with cement--till we at last come
to animals hardly distinguishable, specifically, from the living horse.
Allied to these, are a series of forms showing a transition to the tapirs,
and to the _Palæotherium_ of the European Eocene. In the Pliocene we have
_Parahippus_; in the Miocene _Lophiodon_, found in the same formation and
in the Eocene of Europe, and allied to the tapir; and in the Eocene,
_Palæosyops_, as large as a rhinoceros, which had large canines and was
allied to the tapir and _Palæotherium_; _Limnohyus_, forming the type of a
family Limnohyidæ, which included the last genus and some others mentioned
further on; and _Hyrachyus_, allied to _Lophiodon_, and to _Hyracodon_ an
extinct form of rhinoceros. Besides these we have _Lophiotherium_ (also
from the Eocene of Europe); _Diplacodon_ allied to _Limnohyus_, but with
affinities to modern Perissodactyla and nearly as large as a rhinoceros;
and _Colonoceras_, also belonging to the Limnohyidæ, an animal which was
the size of a sheep, and had divergent protuberances or horns on its nose.
A remarkable genus, _Bathmodon_, lately described by Professor Cope, and of
which five species have been found in the Eocene of New Mexico and Wyoming,
is believed to form the type of a new family, having some affinity to
_Palæosyops_ and to the extinct Brontotheridæ. It had large canine tusks
but no horns.
The Rhinocerotidæ are represented in America by the genus _Rhinoceros_ in
the Pliocene and Miocene, and by _Aceratherium_ and _Hyracodon_ in the
Miocene. Both the latter were hornless, and _Hyracodon_ was allied to the
Eocene _Hyrachyus_, one of the Lophiodontidæ. In the Eocene and Miocene
deposits of Utah, and Oregon, several remarkable extinct rhinoceroses have
been {137}recently discovered, forming the genus _Diceratherium_. These had
a pair of nasal horns placed side by side on the snout, not behind each
other as in existing two-horned rhinoceroses, the rest of their skeleton
resembling the hornless _Aceratherium_. They were of rather small size.
Next to these extinct rhinoceroses come the Brontotheridæ, an extraordinary
family of large mammalia, some of which exceeded in bulk the largest living
rhinoceros. They had four toes to the front and three to the hind feet,
with a pair of large divergent horns on the front of the head, in both
sexes. Professor Marsh and Dr. Leidy have described four genera,
_Brontotherium_, _Titanotheium_, _Megacerops_, and _Anisacodon_,
distinguished by peculiarities of dentition. Though most nearly allied to
the rhinoceroses, they show some affinity for the gigantic Dinocerata of
the Eocene to be noticed further on. Professor Cope has since described
another genus, _Symborodon_, from the Miocene of Colorado, with no less
than seven species, one nearly the size of an elephant. He thinks they had
a short tapir-like proboscis. The species differ greatly in the form of the
cranium and development of the horn-bearing processes.
We commence the Artiodactyla, or even-toed Ungulates, with the hog tribe.
These are represented by species of peccaries, (_Dicotyles_) from the
Pliocene of Nebraska and Oregon; and by an allied form _Thinohyus_, very
like _Dicotyles_, but having an additional premolar tooth and a much
smaller brain-cavity. From the Miocene are three allied genera, _Nanohyus_,
_Leptochoerus_, and _Perchoerus_. Professor Cope, however, thinks
_Leptochoerus_ may be Lemuroid, and allied to _Menotherium_. The
Anthracotheridæ, a family which connects the Hippopotamidæ and Ruminants,
and which occurs in the Miocene of Europe and India, are represented in
America by the genus _Hyopotamus_ from the Miocene of Dakota, and
_Elotherium_ from the Miocene of Oregon and the Eocene of Wyoming; the
latter genus being sometimes classed with the preceding family, and lately
placed by Professor Marsh, in the new order, Tillodontia. Professor Cope
has since described three other genera from the Eocene of New {138}Mexico:
_Meniscotherium_, having resemblances to _Palæosyops_, _Hyopotamus_, and
the Limnotheridæ; _Phenacodus_, the size of a hog, of doubtful position,
but perhaps near _Elotherium_; and _Achænodon_, as large as a cow, but more
hog-like than the preceding. Another new genus from the Miocene of
Colorado--_Pelonax_--is said by Professor Cope to come between _Elotherium_
and _Hippopotamus_.
The Camelidæ are very abundant, and form one of the most striking features
of the ancient fauna of America. _Procamelus_, _Homocamelus_, and
_Megalomeryx_, are extinct genera found in the Pliocene formation; the
first very closely allied to the Old World camel, the last smaller and more
sheep-like. In the Miocene two other genera occur, _Poebrotherium_ and
_Protomeryx_, the former allied to both the camel and the llama.
Deer are represented by a single species of _Cervus_ in the Pliocene, while
two extinct genera, _Leptomeryx_ and _Merycodus_, are found in the Miocene
deposits, the latter indicating a transition between camels and deer. Two
other genera, _Hypisodus_ and _Hypertragulus_, of very small size, are said
by Professor Cope to be allied to the Tragulidæ and to _Leptomeryx_.
The Bovidæ, or hollow-horned ruminants, are only represented in the Newer
Pliocene by a single species of an extinct genus, _Casoryx_, said to be
intermediate between antelopes and deer.
We now come to an exclusively American family, the _Oreodontidæ_, which
consisted of small animals termed by Dr. Leidy, "ruminating hogs," and
which had some general structural resemblances to deer and camels. They
abounded in North America during the Pliocene, and especially during the
Miocene epoch, no less than six genera and twenty species having been
discovered. _Merychus_ contains the Pliocene forms; while _Oreodon_,
_Eporeodon_, _Merychochoerus_, _Leptauchenia_, and _Agriochoerus_ are
Miocene. The last genus extends back into the Eocene period, and shows
affinity to the European Anoplotheridæ of the same epoch.
_Proboscidea._--The Elephantidæ are only represented in America by one
species of _Mastodon_ and one of _Elephas_, in the Newer Pliocene deposits.
In the Older Pliocene, Miocene, {139}and Upper Eocene, no remains of this
order have been found; and in 1869, Dr. Leidy remarked on the small average
size of the extinct North American mammalia, which were almost all smaller
than their living analogues. Since then, however, wonderful discoveries
have been made in deposits of Middle Eocene age in Wyoming and Colorado, of
a group of huge animals not only rivalling the elephants in size, but of so
remarkable and peculiar a structure as to require the formation of a new
order of mammals--Dinocerata--for their reception.
This order consists of animals with generalised Ungulate and Proboscidean
affinities. The lower jaw resembles that of the hippopotamus; they had five
toes on the anterior feet and four on the posterior; three pairs of horns,
the first pair on the top of the head, large and perhaps palmated, the
second pair above the eyes, while the third and smallest stood out sideways
on the snout. They had enormous upper canines, of which the roots entered
the middle horn cores, no upper incisors, and small molars. Professor Marsh
believes that they had no trunk. The remains discovered indicate four
genera, _Dinoceras_ (3 sp.), _Tinoceras_ (2 sp.), _Uintatherium_ (1 sp.),
and _Eobasileus_ (2 sp.). Many other names have been given to fragments of
these animals, and even those here given may not be all distinct.
Another new order, Tillodontia, recently established by Professor Marsh, is
perhaps yet more remarkable in a zoological point of view, since it
combines the characters of Carnivora, Ungulata, and Rodents. These animals
have been formed into two families, Tillotheridæ and Stylinodontidæ; and
three genera, _Tillotherium_, _Anchippodus_, and _Stylinodontia_. All are
from the Eocene of Wyoming and New Jersey. Perhaps to these must be added
_Elotherium_ from the Miocene of Dakota, the other forms being all Eocene.
They were mostly animals of small size, between that of the capybara and
tapir. The skull resembled in form that of a bear; the molar teeth were of
Ungulate type, and the incisors like those of a Rodent; but the skeleton
was more that of the Ursidæ, the feet being plantigrade. Professor Cope has
since described three new genera from the Eocene of New Mexico,
_Ectoganus_, _Calamodon_, and _Esthonyx_, comprising {140}seven species
allied to _Tillotherium_ and _Anchippodus_, and having also relations, as
Professor Cope believes, with the South American Toxodontidæ.
_Rodentia._--This order is represented in the Pliocene by a beaver, a
porcupine, and an American mouse (_Hesperomys_), all extinct species of
living genera, the _Hystrix_ being an Old World type; and Professor Cope
has recently described _Panolax_, a new genus of hares from the Pliocene of
New Mexico. The Miocene deposits have furnished an extinct genus allied to
the hares--_Palæolagus_; one of the squirrel family--_Ischyromys_; a small
extinct form of beaver--_Palæocastor_; and an extinct mouse--_Eumys_. The
Eocene strata of Wyoming have lately furnished two extinct forms of
squirrel, _Paramys_ and _Sciuravus_; and another of the Muridæ (or mouse
family), _Mysops_.
_Cetacea._--Numerous remains of dolphins and whales, belonging to no less
than twelve genera, mostly extinct, have been found in the Miocene deposits
of the Atlantic and Gulf States, from New Jersey to South Carolina and
Louisiana; while seven genera of the extinct family, Zeuglodontidæ, have
been found in Miocene and Eocene beds of the same districts. Some remains
associated with these are doubtfully referred to the Seal family (Phocidæ)
among the Carnivora.
_Edentata._--Till quite recently no remains of this order have occurred in
any North American deposits below the Post-Pliocene; but in 1874 Prof.
Marsh described some remains allied to _Megalonyx_ and _Mylodon_, from the
Pliocene beds of California and Idaho, and forming a new genus,
_Morotherium_. As these remains have only occurred to the west of the Rocky
Mountains, and in Pliocene deposits whose exact age is not ascertained,
they hardly affect the remarkable absence of this group from the whole of
the exceedingly rich Tertiary deposits in all other parts of North America.
_General Relations of the extinct Tertiary Mammalia of North America and
Europe._--Having now given a sketch of the extinct Mammalia which inhabited
Europe and North America during the Tertiary period, we are enabled by
comparing them, {141}to ascertain their relations to each other, and to see
how far they elucidate the problem of the birth-place and subsequent
migrations of the several families and genera. We have already pointed out
the remarkable features of the Quaternary (or Post-Pliocene) fauna of North
America, and now proceed to discuss that of the various Tertiary periods,
which is closely connected with the extinct fauna of Europe.
The Tertiary Mammalia of North America at present described belong to from
eighty to one hundred genera, while those of Europe are nearly double that
number; yet only eighteen genera are common to the two faunas, and of these
eight are living and belong chiefly to the Pliocene period. Taking first,
the genera which in America do not go back beyond the Pliocene period (ten
in number), we find that eight of them in Europe go back to the Upper
Miocene. These are _Felis_, _Pseudælurus_, _Hipparion_, _Cervus_,
_Mastodon_, _Elephas_ (in India), _Castor_ and _Hystrix_; while another,
_Canis_, goes back to the Upper Eocene and the tenth, _Equus_, confined to
the newer Pliocene or perhaps to the Post-Pliocene in America, extends back
to the older Pliocene in Europe. Of the seven European genera which are
confined to the Miocene period in America, three, _Hyænodon_,
_Anchitherium_, and _Lophiodon_ go back to the Eocene in Europe; three
others, _Machairodus_, _Rhinoceros_, and _Aceratherium_, are also of
Miocene age in Europe; _Amphicyon_ goes back to the Lower Miocene of
Europe. _Lophiotherium_ belongs to the Eocene of both countries.
If we turn now to families instead of genera, we find that the same general
rule prevails. Mustelidæ (weasels), Ursidæ (bears), true Equidæ (horses),
and Bovidæ (oxen &c.), go no further back in America than the Pliocene,
while they all go back to the Miocene in Europe. Suidæ (swine) and
Anoplotheridæ (extinct) are found in the American Miocene and in the
European Eocene. Anchitheridæ (extinct) reach the Upper Eocene in America,
while in Europe they range through Upper, Middle, and Lower Eocene. Cervidæ
(deer) alone are Miocene in both countries. There remain two families in
which America has the preeminence. Camelidæ (camels) were wonderfully
developed in {142}the American Pliocene and Miocene periods, abounding in
genera and species; whereas in Europe the group only exists in the
Post-Pliocene or Lower Pliocene, with one Upper Miocene species of
_Camelus_ in N. India. The Anthracotheridæ (extinct), found only in the
Upper Miocene of France and India, reach even the Lower Eocene in America.
These facts may be due, in part, to a want of strict co-ordination between
the Tertiary deposits of Europe and North America,--in part to the
imperfection of the record in the latter country. Yet it does not seem
probable that they are altogether due South America and well marked
differences to imperfect knowledge; yet we find such important families as
the Civets, Hyænas, Giraffes, and Hippopotami absent from America, with the
Weasels, and Antelopes almost so; while America possesses almost all the
Camelidæ, two peculiar orders, Dinocerata and Tillodontia, and four
remarkably peculiar families, Limnotheridæ, Lemuravidæ, Oreodontidæ and
Brontotheridæ. If then the facts at present known represent approximately
the real time-relations of the groups in question on the two continents,
they render it probable that weasels, bears, true horses, swine, oxen,
sheep and antelopes, originated on the Old World continent, and were
transmitted to America during some part of the Miocene period; while camels
originated in the New World, and somewhere about the same time passed over
to Europe. Of the extinct families common to the two hemispheres, the
Anthracotheridæ alone seem to have had an American origin. Of the genera
common to the two countries, almost all seem to have had a European origin,
the only genera of equal date being the two rhinoceroses and three
Anchitheridæ; but if the Brontotheridæ are allied to the Rhinocerotidæ,
these latter may have originated in America, although now an exclusively
Old World type. These conclusions are not improbable when we consider the
much greater size of the Old World continents, extending far into the
tropics and probably {143}always more or less united to the tropical areas;
while the evidence of the extinct mammalia themselves shows, that South
America has been for the most part isolated from the northern continent,
and did not take part in the development of its characteristic Tertiary
fauna.
Before speculating further on this subject, it will be well to lay before
our readers a summary of South American palæontology, after which we shall
be in a better position to draw correct inferences from the whole body of
the evidence.
SOUTH AMERICA.
Unfortunately, our knowledge of the interesting fossil fauna of this
continent, is almost wholly confined to the Post-Pliocene and Pliocene
periods. A few remains have been discovered in deposits believed to be of
Eocene age, but nothing whatever representing the vast intervening period,
so rich in peculiar forms of animal life both in North America and Europe.
_Fauna of the Brazilian caves._--What we know of the Post-Pliocene period
is chiefly due to the long-continued researches of Dr. Lund in the caves of
Central Brazil, mostly situated in a district near the head waters of the
San Francisco river in the Province of Minas Geraes. The caves are formed
in limestone rocks, and are so numerous that Dr. Lund visited thousands,
but only sixty contained bones in any quantity. These caves have a floor of
reddish earth, often crowded with bones. In one experiment, half a cubic
foot of this earth contained jaws of 400 opossums, 2,000 mice, besides
remains of bats, porcupines and small birds. In another trial, the whole of
the earth in a cavern was carried out for examination, amounting to 6,552
firkins; and, from a calculation made by measured samples, it was estimated
to contain nearly seven millions of jaw-bones of cavies, opossums,
porcupines, and mice, besides small birds, lizards, and frogs. This immense
accumulation is believed to have been formed from the bodies of animals
brought into the cavern by owls; and, as these are unsocial birds, the
quantity found implies an {144}immense lapse of time, probably some
thousands of years. More than 100 species of Mammalia, in all, were
obtained in these caves. Some were living species or closely allied to
such; but the majority were extinct, and a considerable number, about
one-fourth, belonged to extinct genera, or genera not now inhabiting South
America. Stone implements and human remains were found in several of the
caves with extinct animals. The following enumeration of these remains is
from the corrected list of M. Gervais.
_Primates._--Extinct species of _Cebus_, _Callithrix_, and _Jacchus_--South
American genera of monkeys; with an extinct genus, _Protopithecus_--an
animal of large size but belonging to the American family Cebidæ.
_Chiroptera._--Species belonging to the South American Phyllostomidæ, and
to two South American genera of other families.
_Carnivora._--Five species of _Felis_, some allied to living animals,
others extinct; a species of the widespread extinct genus _Machairodus_;
and a small species referred to _Cynælurus_, the genus containing the
hunting leopard now found only in Africa and India. Canidæ are represented
by _Canis_ and _Icticyon_ (a living Brazilian species of the latter genus),
and the extinct genus _Speothos_. Mustelidæ are represented by extinct
species of the South American genera _Mephitis_ and _Galictis_. Procyonidæ,
by a species of _Nasua_. Ursidæ, by _Arctotherium_, a genus closely
resembling, if not identical with, that containing the "spectacled bear" of
Chili.
_Ungulata._--_Equus_, _Tapirus_, _Dicotyles_, _Auchenia_, _Cervus_,
_Leptotherium_, and _Antilope_, are the cave-genera of this order. _Equus_
and _Antelope_ are particularly interesting, as representing groups forming
no part of existing South American zoology; while the presence also of
_Leptotherium_, an extinct genus of antelopes, shows that the group was
fairly represented in South America at this comparatively recent period.
_Proboscidea._--A species of _Mastodon_, found also in the Pliocene of La
Plata, represents this order.
_Rodentia._--These abound. _Dasyprocta_, _Cælogenys_, _Cavia_, _Kerodon_,
all living genera of Caviidæ, are represented by {145}extinct species.
_Cercolabes_, the 'tree porcupine' (Cercolabidæ) has two species, one as
large as a peccary; _Myopotamus_, _Loncheres_, _Carterodon_, are existing
genera of spiny rats (Echimyidæ); and there are two extinct genera of the
same family, _Lonchophorus_ and _Phyllomys_. _Lagostomus_ (Chinchillidæ),
the viscacha of the Pampas, is represented by an extinct species. There is
also an extinct species of _Lepus_; several species of _Hesperomys_ and
_Oxymycterus_; and a large _Arvicola_, a genus not living in South America.
_Edentata._--These, which constitute the great feature of the existing
South American fauna, were still more abundant and varied in the Cave
period, and it is remarkable that most of them are extinct _genera_. The
armadillos are alone represented by living forms, _Dasypus_, and _Xenurus_;
_Eurydon_ and _Heterodon_, are extinct genera of the same family, as well
as _Chlamydotherium_--huge armadillos the size of a tapir or rhinoceros,
and _Pachytherium_, which was nearly as large. The ant-eaters are
represented only by _Glossotherium_, an extinct form allied to
_Myrmecophaga_ and _Manis_. The sloths were more numerous, being
represented by the extinct genera _Cælodon_, _Sphenodon_ and _Ochotherium_,
the last of large size. The huge terrestrial sloths--Megatheridæ, also
abounded; there being species of _Megatherium_ and _Megalonyx_, as well as
the allied _Scelidotherium_, supposed to have some affinity for the African
_Orycteropus_.
_Marsupials._--No new forms of these appear, but numerous species of
_Didelphys_, all closely allied to opossums still living in South America.
The preceding sketch of the wonderful cave fauna of Central Brazil, is
sufficient to show that it represents, in the main, a period of great
antiquity. Not only are almost the whole of the species extinct, but there
are twenty extinct genera, and three others not now inhabitants of South
America. The fact that so few remains of the living animals of the country
are found in these caves, indicates that some change of physical conditions
has occurred since they were the receptacles of so many of the larger
animals; and the presence of many extinct genera of {146}large size,
especially among the Edentata and American families of Rodents, are
additional proofs of a very high antiquity. Yet many of these cave animals
are closely allied to those which are found in North America in the
Post-Pliocene deposits only, so that we have no reason to suppose the
cave-fauna to be of much earlier date. But the great amount of organic
change it implies, must give us an enlarged idea of the vast periods of
time, as measured by years, which are included in this, the most recent of
all geological epochs.
_Pliocene Period of Temperate South America._--We have now to consider the
numerous remains of extinct animals found in various deposits in the
Pampas, and in Patagonia, and a few in Bolivia. The age of these is
uncertain; but as they are very similar to the cave-fauna, though
containing a somewhat larger proportion of extinct genera and some very
remarkable new forms, they cannot be _very_ much older, and are perhaps
best referred at present to the newer portion of the Pliocene formation.
_Carnivora._--The genus _Machairodus_ or sabre-toothed tigers, represents
the Felidæ. There are several species of wolves (_Canis_); a weasel
(_Mustela_); two bears of the Brazilian cave-genus _Arctotherium_; and the
extinct European genus _Hyænarctos_.
_Ungulata._--There are two species of _Equus_, found in the Pampas, Chili,
and Bolivia; two of _Macrauchenia_, an extraordinary extinct group allied
to the tapir and _Palæotherium_, but with the long neck, and general size
of a camel. A second species found on the highlands of Bolivia is much
smaller.
A more recent discovery, in Patagonia, is the almost perfect series of
teeth of a large animal named _Homalodontotherium_; and which is believed
by Professor Flower, who has described it, to have been allied to
_Rhinoceros_, and still more to the Miocene _Hyracodon_ from North America;
and also to present some resemblances to _Macrauchenia_, and though much
more remotely, to the curious genus _Nesodon_ mentioned further on.
The Artiodactyla, or even-toed Ungulates, are represented by a species of
_Dicotyles_, or peccary, found in the deposits of the {147}Pampas; by
_Auchenia_, or llama, of which three extinct species inhabited Bolivia, in
which country two allied but extinct genera, _Palæolama_ and
_Camelotherium_, have also been found. Three species of deer (_Cervus_),
from the Pampas deposits, complete the list of Pliocene Ungulates.
_Proboscidea._--The cave species of _Mastodon_ is found also in the Pampas
deposits, and another in the Andes of Chili and Bolivia.
_Rodents._--These are not so numerous as in the caves. There are species of
the existing genera, _Kerodon_ and _Cavia_ (Caviidæ); _Lagostomus_
(Chinchillidæ); _Ctenomys_ (Octodontidæ); _Lepus_ (hare); _Hesperomys_ and
_Oxymycterus_ (Muridæ); _Arvicola_, a genus not living in South America;
and an extinct genus, _Cardiodus_. There is also a remarkable extinct form,
_Typotherium_, larger than the capybara, and having affinities to Edentates
and Ungulates. Three species have been found in the Pampas deposits.
_Edentata._--These are as abundant and remarkable as in the cave deposits.
_Scelidotherium_, _Megatherium_, _Megalonyx_, _Glossotherium_ and
_Dasypus_, have already been noticed as from the Brazilian caves. We have
here, in addition, the huge _Mylodon_ allied to the _Megatherium_, and the
allied genera--_Gnathopsis_ and _Lestodon_. We then come to the huge
extinct armadillos, _Glyptodon_ and _Schistopleurum_, the former consisting
of numerous species, some of which were as large as an elephant. Another
genus, _Eutatus_, is allied to the living three-banded armadillos; and a
species of the existing genus _Euphractus_ has been found in Bolivia.
_Toxodontidæ._--There remain a number of huge animals rivalling the
Megatherium in size, and forming the genera _Toxodon_ and _Nesodon_, but
whose position is doubtful. Several species have been found in the deposits
of the Pampas and Patagonia. They are allied at once to Ungulates, Rodents,
Edentates, and the aquatic Sirenia, in so puzzling a manner that it is
impossible to determine to what order they belong, or whether they require
a new order to be formed for their reception. Some are believed to date
back to the Miocene period, and they indicate what strange forms may still
be discovered, should any {148}productive deposits be found in South
America of middle Tertiary age.
_Pliocene Mammalia of the Antilles._--These may be noticed here, as they
are of special interest, proving the connection of the larger West Indian
Islands with the Continent some time in the later Tertiary period. They
consist of remains of two large animals belonging to the South American
Chinchillidæ, found in cave deposits in the island of Anguilla, and forming
two new genera, _Amblyrhiza_ and _Loxomylus_; and remain allied to
_Megalonyx_ from Cuba, which have been named _Megalocnus_ and _Myomorphus_.
_Eocene fauna of South America._--The few remains yet discovered in the
Tertiary deposits of the Pampas which are believed to be of Eocene age, are
exceedingly interesting, because they show us another change in the scenery
of the great drama of life; there being apparently a considerable
resemblance, at this epoch, between South America and Europe. They consist
of a large extinct feline animal, _Eutemnodus_; of _Palæotherium_ and
_Anoplotherium_, the well-known extinct Ungulates of the European
Tertiaries, and which have never been found in North America; and of three
genera of Rodents,--_Theridromys_, allied to _Echimys_, and found also in
the Eocene and Miocene of France; _Megamys_, allied to the living
_Capromys_ of the Antilles, and also to _Palæomys_, an extinct form of the
French Miocene; and a very large animal referred to _Arvicola_, a genus
found also in the Pliocene deposits of South America, and abundant in the
northern hemisphere. No Edentates have been found.
The resemblances of this fauna to that of Europe rather than to any part of
America, are so strong, that they can hardly be accidental. We greatly
want, however, more information on this point, as well as some
corresponding evidences as to the condition of West and South Africa about
the same epoch, before we can venture to speculate on their bearing as
regards the early migrations of organic forms.
_General Remarks on the Extinct Mammalian Fauna of the Old {149}and New
Worlds._--Leaving the more special applications of palæontological evidence
to be made after discussing the relations of the existing fauna of the
several regions, we propose here to indicate briefly, some of the more
general deductions from the evidence which has now been laid before our
readers.
The first, and perhaps the most startling fact brought out by our
systematic review, is the very recent and almost universal change that has
taken place in the character of the fauna, over all the areas we have been
considering; a change which seems to be altogether unprecedented in the
past history of the same countries as revealed by the geological record. In
Europe, in North America, and in South America, we have evidence that a
very similar change occurred about the same time. In all three we find, in
the most recent deposits--cave-earths, peat-bogs, and gravels--the remains
of a whole series of large animals, which have since become wholly extinct
or only survive in far-distant lands. In Europe, the great Irish elk, the
_Machairodus_ and cave-lion, the rhinoceros, hippopotamus, and
elephant;--in North America, equally large felines, horses and tapirs
larger than any now living, a llama as large as a camel, great mastodons
and elephants, and abundance of huge megatheroid animals of almost equal
size;--in South America these same megatheroids in greater variety,
numerous huge armadillos, a mastodon, large horses and tapirs, large
porcupines, two forms of antelope, numerous bears and felines, including a
_Machairodus_, and a large monkey,--have all become extinct since the
deposition of the most recent of the fossil-bearing strata. This is
certainly not a great while ago, geologically; and it is _almost_ certain
that this great organic revolution, implying physical changes of such vast
proportions that they must have been due to causes of adequate intensity
and proportionate range, has taken place since man lived on the earth. This
is proved to have been the case in Europe, and is supported by much
evidence both as regards North and South America.
It is clear that so complete and sudden a change in the higher forms of
life, does not represent the normal state of things. Species and genera
have not, at all times, become so rapidly extinct. The time occupied by the
"Recent period," that is the {150}time _since_ these changes took place is,
geologically, minute. The time of the whole of the Post-Pliocene period, as
measured by the amount of physical and _general_ organic change known to
have taken place, is exceedingly small when compared with the duration of
the Pliocene period, and still smaller, probably, as compared with the
Miocene. Yet during these two periods we meet with no such break in the
continuity of the forms of life, no such radical change in the _character_
of the fauna (though the number of specific and generic changes may be as
great) as we find in passing from the Post-Pliocene to recent times. For
example, in Central Europe numerous hyænas, rhinoceroses, and antelopes,
with the great _Machairodus_, continued from Miocene all through Pliocene
into Post-Pliocene times; while hippopotami and elephants continued to live
through a good part of the Pliocene and Post-Pliocene periods,--and then
all suddenly became extinct or left the country. In North America there has
been more movement of the fauna in all the periods; but we have similar
great felines, horses, mastodons, and elephants, in the Pliocene and
Post-Pliocene periods, while _Rhinoceros_ is common to the Miocene and
Pliocene, and camels range continuously from Miocene, through Pliocene, to
Post-Pliocene times;--when all alike became extinct. Even in South America
the evidence is, as far as it goes, all the same way. We find
_Machairodus_, _Equus_, _Mastodon_, _Megatherium_, _Scelidotherium_,
_Megalonyx_, and numerous gigantic armadillos, alike in the caves and in
the stratified tertiary deposits of the Pampas;--yet all have since passed
away.
It is clear, therefore, that we are now in an altogether exceptional period
of the earth's history. We live in a zoologically impoverished world, from
which all the hugest, and fiercest, and strangest forms have recently
disappeared; and it is, no doubt, a much better world for us now they have
gone. Yet it is surely a marvellous fact, and one that has hardly been
sufficiently dwelt upon, this sudden dying out of so many large mammalia,
not in one place only but over half the land surface of the globe. We
cannot but believe that there must have been some physical cause for this
great change; and it must have been a cause capable of acting almost
simultaneously over large {151}portions of the earth's surface, and one
which, as far as the Tertiary period at least is concerned, was of an
exceptional character. Such a cause exists in the great and recent physical
change known as "the Glacial epoch." We have proof in both Europe and North
America, that just about the time these large animals were disappearing,
all the northern parts of these continents were wrapped in a mantle of ice;
and we have every reason to believe that the presence of this large
quantity of ice (known to have been thousands of feet if not some miles in
thickness) must have acted in various ways to have produced alterations of
level of the ocean as well as vast local floods, which would have combined
with the excessive cold to destroy animal life. There is great difference
of opinion among geologists and physicists as to the extent, nature, and
duration of the Glacial epoch. Some believe it to have prevailed
alternately in the northern and southern hemispheres; others that it was
simultaneous in both. Some think there was a succession of cold periods,
each lasting many thousands of years, but with intercalated warm periods of
equal duration; others deny that there is any evidence of such changes, and
maintain that the Glacial epoch was one continuous period of arctic
conditions in the temperate zones, with some fluctuations perhaps but with
no regular alternations of warm periods. Some believe in a huge ice-cap
covering the whole northern hemisphere from the pole to near 50° north
latitude in the eastern, and 40° in the western hemisphere; while others
impute the observed effects either to glaciers from local centres, or to
floating icebergs of vast size passing over the surface during a period of
submersion.
Without venturing to decide which of these various theories will be
ultimately proved to be correct, we may state, that there is an increasing
belief among geologists in the long duration of this ice-period, and the
vast extent and great thickness attained by the ice-sheet. One of the most
recent, and not the least able, of the writers on this question (Mr. Belt)
shows strong reasons for adopting the view that the ice-period was
simultaneous in both hemispheres; and he calculates that the vast amount of
water abstracted from the ocean and locked up {152}in mountains of ice
around the two poles, would lower the general level of the ocean about
2,000 feet. This would be equivalent to a general elevation of the land to
the same amount, and would thus tend to intensify the cold; and this
elevation may enable us to understand the recent discoveries of signs of
glacial action at moderate elevations in Central America and Brazil, far
within the tropics. At the same time, the weight of ice piled up in the
north would cause the land surface to sink there, perhaps unequally,
according to the varying nature of the interior crust of the earth; and
since the weight has been removed land would rise again, still somewhat
irregularly; and thus the phenomena of raised beds of arctic shells in
temperate latitudes, are explained.
Now, it is evident, that the phenomena we have been considering--of the
recent changes of the mammalian fauna in Europe, North America, South
Temperate America, and the highlands of Brazil--are such as might be
explained by the most extreme views as to the extent and vastness of the
ice-sheet, and especially as to its simultaneous occurrence in the northern
and southern hemispheres; and where two such completely independent sets of
facts are found to combine harmoniously, and supplement each other on a
particular hypothesis, the evidence in favour of that hypothesis is greatly
strengthened. An objection that will occur to zoologists, may here be
noticed. If the Glacial epoch extended over so much of the temperate and
even parts of the tropical zone, and led to the extinction of so many forms
of life even within the tropics, how is it that so much of the purely
tropical fauna of South America has maintained itself, and that there are
still such a vast number of forms, both of mammalia, birds, reptiles, and
insects, that seem organized for an exclusive existence in tropical
forests? Now Mr. Belt's theory, of the subsidence of the ocean to the
extent of about 2,000 feet, supplies an answer to this objection; for we
should thus have a tract of lowland of an average width of some hundreds of
miles, added to the whole east coast of Central and South America. This
tract would, no doubt, become covered with forests as it was slowly formed,
would enjoy a perfectly {153}tropical climate, and would thus afford an
ample area for the continued existence and development of the typical South
American fauna; even had glaciers descended in places so low as what is now
the level of the sea, which, however, there is no reason to believe they
ever did. It is probable too, that this low tract, which all round the Gulf
of Mexico would be of considerable width, offered that passage for
intermigration between North and South America, which led to the sudden
appearance in the former country in Post-Pliocene times, of the huge
Megatheroids from the latter; a migration which took place in opposite
directions as we shall presently show.
_The birth-place and migrations of some mammalian families and genera._--We
have now to consider a few of those cases in which the evidence already at
our command, is sufficiently definite and complete, to enable us to
pronounce with some confidence as to the last movements of several
important groups of mammalia.
_Primates._--The occurrence in North America of numerous forms of
Lemuroidea, forming two extinct families, which are believed by American
palæontologists to present generalized features of both Lemuridæ and
Hapalidæ, while in Europe only Lemurine forms allied to those of Africa
have occurred in deposits of the same age (Eocene), renders it possible
that the Primates may have originated in America, and sent one branch to
South America to form the Hapalidæ and Cebidæ, and another to the Old
World, giving rise to the lemurs and true apes. But the fact that apes of a
high degree of organization occur in the European Miocene, while in the
Eocene, a monkey believed to have relations to the Lemuroids and Cebidæ has
also been discovered, make it more probable that the ancestral forms of
this order originated in the Old World at a still earlier period. The
absence of any early tertiary remains from the tropical parts of the two
hemispheres, renders it impossible to arrive at any definite conclusions as
to the origin of groups which were, no doubt, always best developed in
tropical regions.
_Carnivora._--This is a very ancient and wide-spread group, the families
and genera of which had an extensive range in very {154}early times. The
true bears (_Ursus_) are almost the only important genus that seems to have
recently migrated. In Europe it dates back to the Older Pliocene, while in
North America it is Post-Pliocene only. Bears, therefore, seem to have
passed into America from the Palæarctic region in the latter part of the
Pliocene period. They probably came in on the north-west, and passed down
the Andes into South America, where one isolated species still exists.
_Ungulata._--Horses are very interesting. In Europe they date back under
various forms to the Miocene period, and true _Equus_ to the Older
Pliocene. In North America they are chiefly Pliocene, true _Equus_ being
Post-Pliocene, with perhaps one or two species Newer Pliocene; but numerous
ancestral forms date back to the Miocene and Eocene, giving a more perfect
"pedigree of the horse" than the European forms, and going back to a more
primitive type--_Orohippus_. In South America, _Equus_ is the only genus,
and is Post-Pliocene or at most Newer Pliocene. While, therefore, the
ancient progenitors of the Equidæ were common to North America and Europe,
in Miocene and even Eocene times, true horses appear to have arisen in the
Palæarctic region, to have passed into North America in the latter part of
the Pliocene period, and thence to have spread over all suitable districts
in South America. They were not, however, able to maintain themselves
permanently in their new territory, and all became extinct; while in their
birth-place, the Old World, they continue to exist under several varied
forms.
True tapirs are an Old World group. They go back to the Lower Miocene in
Europe, while in both North and South America they are exclusively
Post-Pliocene. They occur in France down to the Newer Pliocene, and must,
about that time, have entered America. The land connection by which this
and so many other animals passed between the Old and New Worlds in late
Tertiary times, was almost certainly in the North Pacific, south of
Behring's Straits, where, as will be seen by our general map, there is a
large expanse of shallow water, which a moderate elevation would convert
into dry land, in a sufficiently temperate latitude.
{155}The peccary (_Dicotyles_), now a characteristic South American genus,
is a recent immigrant from North America, where it appears to have been
developed from ancestral forms of swine dating back to the Miocene period.
Antelopes are an Old World type, but a few of them appear to have entered
North, and reached South America in late Pliocene times. Camels, strange to
say, are a special North American type, since they abounded in that
continent under various ancient forms in the Miocene period. Towards the
end of that period they appear to have entered eastern Asia, and developed
into the Siberian _Merycotherium_ and the North Indian _Camelus_, while in
the Pliocene age the ancestral llamas entered South America.
_Cervidæ_ are a wide-spread northern type in their generalized form, but
true deer (_Cervus_) are Palæarctic. They abounded in Europe in Miocene
times, but only appear in North and South America in the later Pliocene and
Post-Pliocene periods.
True oxen (_Bovinæ_) seem to be an Oriental type (Miocene), while they
appear in Europe only late in the Pliocene period, and in America are
confined to the Post-Pliocene.
Elephants (_Elephantidæ_) are an Old World type, abounding in the Miocene
period in Europe and India, and first appearing in America in Post-Pliocene
or later Pliocene times. Ancestral forms, doubtfully Proboscidean
(_Dinocerata_), existed in North America in the Eocene period, but these
became extinct without leaving any direct descendants, unless the
_Brontotheridæ_ and rhinoceroses may be so considered.
Marsupials are almost certainly a recent introduction into South and North
America from Asia. They existed in Europe in Eocene and Miocene times, and
presumably over a considerable part of the Old World; but no trace of them
appears in North or South America before the Post-Pliocene period.
_Edentata._--These offer a most curious and difficult problem. In South
America they abound, and were so much more numerous and varied in the
Post-Pliocene and Pliocene, that we may be sure they lived also in the
preceding Miocene period. A few living Edentates are scattered over Africa
and Asia, and {156}they flourished in Europe during the Miocene
age--animals as large (in some species) as a rhinoceros, and most allied to
living African forms. In North America no trace of Edentata has been found
earlier than the Post-Pliocene period, or perhaps the Newer Pliocene on the
west coast. Neither is there any trace of them in South America in the
Eocene formations; but this may well be owing to our very imperfect
knowledge of the forms of that epoch. Their absence from North America is,
however, probably real; and we have to account for their presence in the
Old World and in South America. Their antiquity is no doubt very great, and
the point of divergence of the Old World and South American groups, may
take us back to early Eocene, or even to Pre-Eocene times. The distribution
of land and sea may then have been very different from what it is now; and
to those who would create a continent to account for the migrations of a
beetle, nothing would seem more probable than that a South Atlantic
continent, then united parts of what are now Africa and South America.
There is, however, so much evidence for the general permanence of what are
now the great continents and deep oceans, that Professor Huxley's
supposition of a considerable extension of land round the borders of the
North Pacific Ocean in Mesozoic times, best indicates the probable area in
which the Edentate type originated, and thence spread over much of the Old
World and South America. But while in the latter country it flourished and
increased with little check, in the other great continents it was soon
overcome by the competition of higher forms, only leaving a few small-sized
representatives in Africa and Asia.
{157}CHAPTER VIII.
VARIOUS EXTINCT ANIMALS;--AND ON THE ANTIQUITY OF THE GENERA OF INSECTS AND
LAND MOLLUSCA.
EXTINCT MAMMALIA OF AUSTRALIA.
These have all been obtained from caves and late Tertiary or Post-Tertiary
deposits, and consist of a large number of extinct forms, some of gigantic
size, but all marsupials and allied to the existing fauna. There are
numerous forms of kangaroos, some larger than any living species; and among
these are two genera, _Protemnodon_ and _Sthenurus_, which Professor Garrod
has lately shown to have been allied, not to any Australian forms, but to
the _Dendrolagi_ or tree-kangaroos of New Guinea. We have also remains of
_Thylacinus_ and _Dasyurus_, which now only exist in Tasmania; and extinct
species of _Hypsiprymnus_ and _Phascolomys_, the latter as large as a
tapir. Among the more remarkable extinct genera are _Diprotodon_, a huge
thick-limbed animal allied to the kangaroos, but nearly as large as an
elephant; _Nototherium_, having characters of _Macropus_ and
_Phascolarctos_ combined, and as large as a rhinoceros; and _Thylacoleo_, a
phalanger-like marsupial nearly as large as a lion, and supposed by
Professor Owen to have been of carnivorous habits, though this opinion is
not held by other naturalists.
Here then we find the same phenomena as in the other countries we have
already discussed,--the very recent disappearance of a large number of
peculiar forms, many of them far surpassing in size any that continue to
exist. It hardly seems probable that in this case their disappearance can
have been due to the direct effects of the Glacial epoch, since no very
extensive {158}glaciation could have occurred in a country like Australia;
but if the ocean sank 2,000 feet, the great eastern mountain range might
have given rise to local glaciers. It is, however, almost certain that
during late Tertiary times Australia must have been much more extensive
than it is now. This is necessary to allow of the development of its
peculiar and extensive fauna, especially as we see that that fauna
comprised animals rivalling in bulk those of the great continents. It is
further indicated by the relations with New Guinea, already alluded to, and
by the general character of the various faunas which compose the Australian
region, details of which will be found in the succeeding part of this work.
The lowering of the ocean during the Glacial period would be favourable to
the still further development of the fauna of such a country; and it is to
the unfavourable conditions produced by its subsequent rising--equivalent
to a depression of the land to the amount of two thousand feet--that we
must impute the extinction of so many remarkable groups of animals. It is
not improbable, that the disappearance of the ice and the consequent
(apparent) subsidence of the land, might have been rapid as compared with
the rate at which large animals can become modified to meet new conditions.
Extensive tracts of fertile land might have been submerged, and the
consequent crowding of large numbers of species and individuals on limited
areas would have led to a struggle for existence in which the less adapted
and less easily modifiable, not the physically weaker, would succumb.
There is, however, another cause for the extinction of large rather than
small animals whenever an important change of conditions occurs, which has
been suggested to me by a correspondent,[4] but which has not, I believe,
been adduced by Mr. Darwin or by any other writer on the subject. It is
dependent on the fact, that large animals as compared with small ones are
almost invariably slow breeders, and as they also necessarily exist in much
smaller numbers in a given area, they offer far less materials for
favourable variations than do smaller animals. In such an extreme case as
that of the rabbit and elephant, the {159}young born each year in the world
are probably as some millions to one; and it is very easily conceivable
that in a thousand years the former might, under pressure of rapidly
changing conditions, become modified into a distinct species, while the
latter, not offering enough favourable variations to effect a suitable
adaptation, would become extinct. We must also remember the extreme
specialization of many of the large animals that have become extinct--a
specialization which would necessarily render modification in any new
direction difficult, since the inherited tendency of variation would
probably be to increase the specialization in the same directions which had
heretofore been beneficial. If to these two causes we add the difficulty of
obtaining sufficient food for such large animals, and perhaps the injurious
effects of changes of climate, we shall not find it difficult to understand
how such a vast physical revolution as the Glacial epoch, with its
attendant phenomena of elevations and subsidences, icy winds, and sudden
floods by the bursting of lake barriers, might have led to the total
extinction of a vast number of the most bulky forms of mammalia, while the
less bulky were able to survive, either by greater hardiness of
constitution or by becoming more or less modified. The result is apparent
in the comparatively small or moderate size of the species constituting the
temperate fauna, in all parts of the globe.
It is much to be regretted that no mammalian remains of earlier date have
been found in Australia, as we should then see if it is really the case
that marsupials have always formed its highest type of mammalian life. At
present its fossil fauna is chiefly interesting to the zoologist, but
throws little light on the past relations of this isolated country with
other parts of the globe.
MAMMALIAN REMAINS IN THE SECONDARY FORMATIONS.
In the oldest Tertiary beds of Europe and North America, we have (even with
our present imperfect record) a rich and varied mammalian fauna. As
compared with our living or recent highly specialized forms, it may be said
to consist of generalised types; but as compared with any primeval
mammalian type, it must be pronounced highly specialised. Not only are such
diversified {160}groups as Carnivora, Perrissodactyle and Artiodactyle
Ungulates, Primates, Chiroptera, Rodents, and Marsupials already well
marked, but in many of these there is a differentiation into numerous
families and genera of diverse character. It is impossible therefore to
doubt, that many peculiar forms of mammalia must have lived long anterior
to the Eocene period; but there is unfortunately a great gap in the record
between the Eocene and Cretaceous beds, and these latter being for the most
part marine continue the gap as regards mammals over an enormous lapse of
time. Yet far beyond both these chasms in the Upper Oolitic strata, remains
of small mammalia have been found; again, in the Stonesfield slate, a
member of the Lower Oolite, other forms appear. Then comes the marine Lias
formation with another huge gap; but beyond this again in the Upper Trias,
the oldest of the secondary formations, mammalian teeth have been
discovered in both England and Germany, and these are, as nearly as can be
ascertained, of the same age as the _Dromatherium_ already noticed, from
North America. They have been named _Microlestes_, and show some
resemblance to those of the West Australian _Myrmecobius_. In the Oolitic
strata numerous small jawbones have been found, which have served to
characterise eight genera, all of which are believed to have been
Marsupials, and in some of them a resemblance can be traced to some of the
smaller living Australian species. These, however, are mere indications of
the number of mammalia that must have lived in the secondary period, so
long thought to be exclusively "the age of reptiles;" and the fact that the
few yet found are at all comparable with such specialised forms as still
exist, must convince us, that we shall have to seek far beyond even the
earliest of these remains, for the first appearance of the mammalian type
of vertebrata.
EXTINCT BIRDS.
Compared with those of mammalia, the remains of birds are exceedingly
scarce in Europe and America; and from the wandering habits of so many of
this class, they are of much less value {161}as indications of past changes
in physical geography. A large proportion of the remains belong to aquatic
or wading types, and as these have now often a world-wide range, the
occurrence of extinct forms can have little bearing on our present inquiry.
There are, however, a few interesting cases of extinct land-birds belonging
to groups now quite strangers to the country in which they are found; and
others scarcely less interesting, in which groups now peculiar to certain
areas are shown to have been preceded by allied species or genera of
gigantic size.
_Palæarctic Region and N. India._--In the caves and other Post-Pliocene
deposits of these countries, the remains of birds almost all belong to
genera now inhabiting the same districts. Almost the only exceptions are,
the great auk and the capercailzie, already mentioned as being found in the
Danish mounds; the latter bird, with _Tetrao albus_, in Italian caverns;
and a species of pheasant (_Phasianus_) said to have occurred in the
Post-Pliocene of France, considerably west of the existing range of the
genus in a wild state.
In the preceding Pliocene deposits, but few remains have been found, and
all of existing genera but one, a gallinaceous bird (_Gallus bravardi_)
allied to the domestic fowl and peacock.
The Miocene beds of France and Central Europe have produced many more
remains of birds, but these, too, are mostly of existing European genera,
though there are some notable exceptions. Along with forms
undistinguishable from crows (_Corvus_), shrikes (_Lanius_), wagtails
(_Motacilla_), and woodpeckers (_Picus_), are found remains allied to the
Oriental edible-nest swift (_Collocalia_) and _Trogon_; a parrot resembling
the African genus _Psittacus_; an extinct form _Necrornis_, perhaps allied
to the plantain-eaters (_Musophaga_); _Homalophus_, doubtfully allied to
woodpeckers, and _Limnatornis_ to the hoopoes. The gallinaceous birds are
represented by three species of pheasants, some very close to the
domesticated species; _Palæoperdix_ allied to the partridges; and
_Palæortyx_, small birds allied to the American genus _Ortyx_, but with
larger wings. There are also species of _Pterocles_ allied to living birds,
and a small pigeon. There are numerous living genera of Accipitres; such as
eagle (_Aquila_), {162}kite (_Milvus_), eagle-owl (_Bubo_), and screech-owl
(_Strix_); with the African secretary-bird (_Serpentarius_), and some
extinct forms, as _Palæocercus_, _Palæohierix_ and _Palæetus_.
Aquatic and wading birds were abundant, including numerous rails, bustards,
herons, sandpipers, gulls, divers, and pelicans. There were also many
ducks, some allied to the genus _Dendrocygna_; the Oriental genus of
storks, _Leptoptilus_; _Ibidipodia_, a remarkable form allied to _Ibis_ and
_Ciconia_; _Elornis_, near _Limosa_; _Pelagornis_, a large bird allied to
gannets and pelicans; _Hydrornis_, allied to the ducks and petrels;
_Dolichopterus_, allied to plovers. Perhaps the most interesting of these
extinct birds are, however, the flamingoes, represented by forms hardly
distinguishable from living species, and by one extinct genus _Palælodus_,
which had very long toes, and probably walked on aquatic plants like the
tropical jacanas.
The Miocene beds of North India have furnished few birds; the only one of
geographical interest being an extinct species of ostrich, not very
different from that now inhabiting Arabia.
On the whole, the birds of Europe at this period were very like those now
living, with the addition of a few tropical forms. These latter were,
however, perhaps more numerous and important than they appear to be, as
they belong to inland and forest-haunting types, which would not be so
frequently preserved as the marsh and lake-dwelling species. Taking this
into consideration, the assemblage of Miocene birds accords well with what
we know of the mammalian fauna. We have the same indications of a luxuriant
vegetation and subtropical climate, and the same appearance of Oriental and
especially of African types. _Trogon_ is perhaps the most interesting of
all the forms yet discovered, since it furnishes us with a central point
whence the living trogons of Asia, Africa, and South America might have
diverged.
In the Eocene we find ourselves almost wholly among extinct forms of birds.
The earliest known Passerine bird is here met with, in _Protornis_,
somewhat similar to a lark, found in the Lower Eocene of Switzerland; while
another Passerine form, _Palægithalus_, and one allied to the nuthatch
(_Sitta_), have been {163}discovered in the Upper Eocene of Paris. Picariæ
of equal antiquity are found. _Cryptornis_, from the Paris Eocene, and
_Halcyornis_ from the Lower Eocene of the Isle of Sheppey, were both allied
to kingfishers; while a form allied to _Centropus_ a genus of cuckoos, or,
as Milne-Edwards thinks, to the Madagascar _Leptosomus_, has been found in
the Upper Eocene of France. Several _Accipitres_ of somewhat doubtful
affinities have been found in the same country; while _Lithornis_, from the
Lower Eocene of the Isle of Sheppey, was a small vulturine bird supposed to
be allied to the American group, _Cathartes_. Among the waders, some
extinct forms of plovers have been found, and a genus (_Agnopterus_),
allied to the flamingoes; while there are many swimming birds, such as
pelicans, divers, and several extinct types of doubtful affinities. Most
intersting of all is a portion of a cranium discovered in the Lower Eocene
of Sheppey, and lately pronounced by Professor Owen to belong to a large
Struthious bird, allied to the New Zealand _Dinornis_ and also perhaps to
the ostrich. Another gigantic bird is the _Gastornis_, from the Lower
Eocene of Paris, which was as large as an ostrich, but which is believed to
have been a generalised type, allied to wading and swimming birds as well
as to the Struthiones.
Beyond this epoch we have no remains of birds in European strata till we
come to the wonderful _Archæopteryx_ from the Upper Oolite of Bavaria; a
bird of a totally new type, with a bony tail, longer than the body, each
vertebra of which carried a pair of diverging feathers.
_North America._--A number of bird-remains have lately been found in the
rich Tertiary and Cretaceous deposits of the United States; but here, too,
comparatively few are terrestrial forms. No Passerine bird has yet been
found. The Picariæ are represented by _Uintornis_, an extinct form allied
to woodpeckers, from the Eocene of Wyoming. Species of turkey (_Meleagris_)
occur in the Post-Pliocene and as far back as the Miocene strata, showing
that this interesting type is a true denizen of temperate North America.
The other birds are, _Accipitres_; waders and aquatics of existing genera;
and a number of extinct forms of the two latter orders--such as,
_Aletornis_ an Eocene wader; {164}_Palæotringa_, allied to the sandpipers,
and _Telmatobius_ to the rails, both Cretaceous; with _Graculavus_, allied
to _Graculus_; _Laornis_ allied to the swans; _Hesperornis_ a gigantic
diver; and _Icthyornis_ a very low form, with biconcave vertebra, such as
are only found in fishes and some reptiles--also from Cretaceous deposits.
_South America._--The caverns of Brazil produced thirty-four species of
birds, most of them referable to Brazilian genera, and many to still
existing species. The most interesting were two species of American ostrich
(_Rhea_), one larger than either of the living species; a large
turkey-buzzard (_Cathartes_); a new species of the very isolated South
American genus _Opisthocomus_; and a _Cariama_, or allied new genus.
_Madagascar and the Mascarene Islands._--We have here only evidence of
birds that have become extinct in the historical period or very little
earlier. First we have a group of birds incapable of flight, allied to
pigeons, but forming a separate family, _Dididæ_; and which, so far as we
yet know, inhabited Mauritius, Rodriguez, and probably Bourbon.
_Aphanapteryx_, an extinct genus of rails, inhabited Mauritius; and another
genus, (_Erythromachus_), Rodriguez. A large parrot, said by Prof. Milne
Edwards to be allied to _Ara_ and _Microglossus_, also inhabited Mauritius;
and another allied to _Eclectus_, the island of Rodriguez. None of these
have been found in Madagascar; but a gigantic Struthious bird, _Æpyornis_,
forming a peculiar family distinct both from the ostriches of Africa and
the _Dinornis_ of New Zealand inhabited that island; and there is reason to
believe that this may have lived less than 200 years ago.
_New Zealand._--A number of extinct Struthious birds, forming two families,
_Dinornithidæ_ and _Palapterygidæ_, have been found in New Zealand. Some
were of gigantic size. They seem allied both to the living _Apteryx_ of New
Zealand and the emu of Australia. They are quite recent, and some of them
have probably lived within the last few centuries. Remains of _Dinornis_
have also been found in a Post-Pliocene deposit in Queensland, N. E.
Australia[5]--a very important discovery, as it {165}gives support to the
theory of a great eastward extension of Australia in Tertiary times.
EXTINCT TERTIARY REPTILES.
These will not occupy us long, as no very great number are known, and most
of them belong to a few principal forms of comparatively little
geographical interest.
Tortoises are perhaps the most abundant of the Tertiary reptiles. They are
numerous in the Eocene and Miocene formations both in Europe and North
America. The genera _Emys_ and _Trionyx_ abound in both countries, as well
as in the Miocene of India. Land tortoises occur in the Eocene of North
America and in the Miocene of Europe and India, where the huge
_Colossochelys_, twelve feet long, has been found. In the Pliocene deposits
of Switzerland the living American genus _Chelydra_ has been met with.
These facts, together with the occurrence of a living _species_ in the
Miocene of India, show that this order of reptiles is of great antiquity,
and that most of the genera once had a wider range than now.
Crocodiles, allied to the three forms now characteristic of India, Africa,
and America, have been found in the Eocene of our own country, and several
species of _Crocodilus_ have occurred in beds of the same age in North
America.
Lizards are very ancient, many small terrestrial forms occurring in all the
Tertiary deposits. A species of the genus _Chamæleo_ is recorded from the
Eocene of North America, together with several extinct genera.
Snakes were well developed in the Eocene period, where remains of several
have been found which must have been from twelve to twenty feet long. An
extinct species of true viper has occurred in the Miocene of France, and
one of the Pythonidæ in the Miocene brown coal of Germany.
Batrachia occur but sparingly in a fossil state in the Tertiary deposits.
The most remarkable is the large Salamander (_Andreas_) from the Upper
Miocene of Switzerland, which {166}is allied to the _Menopoma_ living in
North America. Species of frog (_Rana_), and _Palæophryus_ an extinct genus
of toads, have been found in the Miocene deposits of Germany and
Switzerland.
Fresh water fish are almost unknown in the Tertiary deposits of Europe,
although most of the families and some genera of living marine fish are
represented from the Eocene downwards.
ANTIQUITY OF THE GENERA OF INSECTS.
Fossil insects are far too rarely found, to aid us in our determination of
difficult questions of geographical distribution; but in discussing these
questions it will be important to know, whether we are to look upon the
existing generic forms of insects as of great or small antiquity, compared
with the higher vertebrates; and to decide this question the materials at
our command are ample.
The conditions requisite for the preservation of insects in a fossil state
are no doubt very local and peculiar; the result being, that it is only at
long intervals in the geological record that we meet with remains of
insects in a recognisable condition. None appear to have been found in the
Pliocene formation; but in the Upper Miocene of Oeninghen in Switzerland,
associated with the wonderfully rich fossil flora, are found immense
quantities of insects. Prof. Heer examined more than 5,000 specimens
belonging to over 800 species, and many have been found in other localities
in Switzerland; so that more than 1,300 species of Miocene insects have now
been determined. Most of the orders are represented, but the beetles
(Coleoptera) are far the most abundant. Almost all belong to existing
genera, and the majority of these genera now inhabit Europe, only three or
four being exclusively Indian, African, or American.
In the Lower Miocene of Croatia there is another rich deposit of insects,
somewhat more tropical in character, comprising large white-ants and
dragon-flies differently marked from any {167}now inhabiting Europe. A
butterfly is also well preserved, with all the markings of the wings; and
it seems to be a _Junonia_, a tropical genus, though it may be a _Vanessa_,
which is European, but the fossil most resembles Indian species of
_Junonia_.
The Eocene formations seem to have produced no insect remains; but they
occur again in the Upper Cretaceous at Aix-la-Chapelle, where two
butterflies have been found, _Cyllo sepulta_ and _Satyrites Reynesii_, both
belonging to the Satyridæ, and the former to a genus now spread over
Africa, India, and Australia.
A little earlier, in the Wealden formation of our own country, numerous
insects have been found, principally dragon flies (_Libellula_, _Æshna_);
aquatic Hemiptera (_Velia Hydrometra_); crickets, cockroaches, and cicadas,
of familiar types.
Further back in the Upper Oolite of Bavaria--which produced the wonderful
long-tailed bird, _Archæopteryx_--insects of all orders have been found,
including a moth referred to the existing genus _Sphinx_.
In the Lower Oolite of Oxfordshire many fossil beetles have been found
whose affinities are shown by their names:--_Buprestidium_,
_Curculionidium_, _Blapsidium_, _Melolonthidium_, and _Prionidium_; a wing
of a butterfly has also been found, allied to the Brassolidæ now confined
to tropical America, and named _Palæontina oolitica_.
Still more remote are the insects of the Lias of Gloucestershire, yet they
too can be referred to well-known family types--Carabidæ, Melolonthidæ,
Telephoridæ, Elateridæ, and Curculionidæ, among beetles; Gryllidæ and
Blattidæ among Orthoptera; with _Libellula_, _Agrion_, _Æshna_, _Ephemera_,
and some extinct genera. When we consider that almost the only vertebrata
of this period were huge Saurian reptiles like the _Icthyosaurus_,
_Plesiosaurus_, and _Dinosaurus_, with the flying Pterodactyles; and that
the great mass of our existing genera, and even families, of fish and
reptiles had almost certainly not come into existence, we see at once that
types of insect-form are, proportionately, far more ancient. At this remote
epoch we find the chief family types (the _genera_ of the time of Linnæus)
perfectly differentiated {168}and recognisable. It is only when we go
further back still, into the Palæozoic formations, that the insect forms
begin to show that generalization of type which renders it impossible to
classify them in any existing groups. Yet even in the coal formation of
Nova Scotia and Durham, the fossil insects are said by competent
entomologists to be "allied to _Ephemera_," "near _Blatta_," "near
_Phasmidæ_;" and in deposits of the same age at Saarbrück near Trèves, a
well-preserved wing of a grasshopper or locust has been found, as well as a
beetle referred to the Scarabeidæ. More remarkable, however, is the recent
discovery in the carboniferous shales of Belgium, of the clearly-defined
wing of a large moth (_Breyeria borinensis_), closely resembling some of
the Saturniidæ; so that we have now all the chief orders of
Insects--including those supposed to be the most highly developed and the
most recent--well represented at this very remote epoch. Even the oldest
insects, from the Devonian rocks of North America, can mostly be classed as
Neuroptera or Myriapoda, but appear to form new families.
We may consider it, therefore, as proved, that many of the larger and more
important genera of insects date back to the beginning of the Tertiary
period, or perhaps beyond it; but the family types are far older, and must
have been differentiated very early in the Secondary period, while some of
them perhaps go back to Palæozoic times. The great comparative antiquity of
the _genera_ is however the important fact for us, and we shall have
occasion often to refer to it, in endeavouring to ascertain the true
bearing of the facts of insect distribution, as elucidating or invalidating
the conclusions arrived at from a study of the distribution of the higher
animals.
ANTIQUITY OF THE GENERA OF LAND AND FRESH-WATER SHELLS.
The remains of land and fresh-water shells are not much more frequent than
those of insects. Like them, too, their forms are very stable, continuing
unchanged through several geological {169}periods. In the Pliocene and
Miocene formations, most of the shells are very similar to living species,
and some are quite identical. In the Eocene we meet with ordinary forms of
the genera _Helix_, _Clausilia_, _Pupa_, _Bulimus_, _Glandina_,
_Cyclostoma_, _Megalostoma_, _Planorbis_, _Paludina_ and _Limnæa_, some
resembling European species, others more like tropical forms. A British
Eocene species of _Helix_ is still living in Texas; and in the South of
France are found species of the Brazilian sub-genera _Megaspira_ and
_Anastoma_. In the secondary formation no true land shells have been found,
but fresh water shells are tolerably abundant, and almost all are still of
living forms. In the Wealden (Lower Cretaceous) and Purbeck (Upper Oolite)
are found _Unio_, _Melania_, _Paludina_, _Planorbis_, and _Limnæa_; while
the last named genus occurs even in the Lias.
The notion that land shells were really not in existence during the
secondary period is, however, proved to be erroneous by the startling
discovery, in the Palæozoic coal measures of Nova Scotia, of two species of
Helicidæ, both of living genera--_Pupa vetusta_, and _Zonites priscus_.
They have been found in the hollow trunk of a _Sigillaria_, and in great
quantities in a bed full of Stigmarian rootlets. The most minute
examination detects no important differences of form or of microscopic
structure, between these shells and living species of the same genera!
These mollusca were the contemporaries of Labyrinthodonts and strange
Ganoid fishes, which formed almost the whole vertebrate fauna. This
unexpected discovery renders it almost certain, that numbers of other
existing genera, of which we have found no traces, lived with these two
through the whole secondary period; and we are thus obliged to assume as a
probability, that any particular genus has lived through a long succession
of geological ages. In estimating the importance of any peculiarities or
anomalies in the geographical distribution of land shells as compared with
the higher vertebrates, we shall, therefore, have to keep this possible,
and even probable high antiquity, constantly in mind.
We have now concluded our sketch of Tertiary Palæontology as a preparation
for the intelligent study of the Geographical {170}Distribution of Land
Animals; and however imperfectly the task has been performed, the reader
will at all events have been convinced that some such preliminary
investigation is an essential and most important part of our work. So much
of palæontology is at present tentative and conjectural, that in combining
the information derived from numerous writers, many errors of detail must
have been made. The main conclusions have, however, been drawn from as
large a basis of facts as possible; and although fresh discoveries may show
that our views as to the past history of some of the less important genera
or families are erroneous, they can hardly invalidate our results to any
important degree, either as regards the intercommunications between
separate regions in the various geological epochs, or as to the centres
from which some of the more important groups have been dispersed.
PART III.
_ZOOLOGICAL GEOGRAPHY:_
_A REVIEW OF THE CHIEF FORMS OF ANIMAL LIFE IN THE SEVERAL REGIONS AND
SUB-REGIONS, WITH THE INDICATIONS THEY AFFORD OF GEOGRAPHICAL MUTATIONS._
{173}CHAPTER IX.
THE ORDER OF SUCCESSION OF THE REGIONS.--COSMOPOLITAN GROUPS OF
ANIMALS.--TABLES OF DISTRIBUTION.
Having discussed, in our First Part, such general and preliminary matters
as are necessary to a proper comprehension of our subject; and having made
ourselves acquainted, in our Second Part, with the most important results
of Palæontology, we now come to our more immediate subject, which we
propose to treat first under its geographical aspect. Taking each of our
six regions in succession, we shall point out in some detail the chief
zoological features they present, as influenced by climate, vegetation, and
other physical features. We shall then treat each of the sub-regions by
itself, as well as such of the islands or other sub-divisions as present
features of special interest; endeavouring to ascertain their true
relations to each other, and the more important changes of physical
geography that seem necessary to account for their present zoological
condition.
_Order of Succession of the Regions._--We may here explain the reason for
taking the several regions in a different succession from that in which
they appear in the tabular or diagrammatic headings to each family, in the
Fourth, and concluding part of this work. It will have been seen, by our
examination of extinct animals (and it will be made still clearer during
our study of the several regions) that all the chief types of animal life
appear to have originated in the great north temperate or northern
continents; while the southern continents--now represented by {174}South
America, Australia, and South Africa with Madagascar--have been more or
less completely isolated, during long periods, both from the northern
continent and from each other. These latter countries have, however, been
subject to more or less immigration from the north during rare epochs of
approximation to, or partial union with it. In the northern, more
extensive, and probably more ancient land, the process of development has
been more rapid, and has resulted in more varied and higher types; while
the southern lands, for the most part, seem to have produced numerous
diverging modifications of the lower grades of organization, the original
types of which they derived either from the north, or from some of the
ancient continents in Mesozoic or Palæozoic times. Hence those curious
resemblances in the fauna of South America, Australia, and, to a less
extent, Madagascar, which have led to a somewhat general belief that these
distant countries must at one time or other have been united; a belief
which, after a careful examination of all the facts, does not seem to the
author of this work to be well founded. On the other hand, there is the
most satisfactory evidence that each southern region has been more or less
closely united (during the tertiary or later secondary epoch) with the
great northern continents, leading to numerous resemblances and affinities
in their productions.
In endeavouring to present at a glance in the most convenient manner, the
distribution of the families in the several regions and sub-regions, it was
necessary to arrange them, so that those whose relations to each other were
closest should stand side by side; the first and last being those between
which the relations were least numerous and least important. Influenced by
the usual opinions as to the relations between Australia and South America,
the series was at first begun with the Nearctic, and terminated with the
Australian and Neotropical regions; and it was not till the whole of the
vertebrate families had been gone through, and their distribution carefully
studied, that these last two regions were seen to be really wider apart
than any others of the series. It was therefore decided to alter the
arrangement, beginning with the Neotropical, and ending with the Australian
{175}regions; and a careful inspection of the diagrams themselves, taken in
their entirety, will, it is believed, show that this is the most natural
plan, and most truly exhibits the relations of the several regions.
In the portion of our work now commencing, we are not, however, by any
means bound to begin at either end of this series. Each region is studied
by itself, but reference will often have to be made to all the other
regions; and wherever we begin, we must occasionally refer to facts which
will be given further on. As, however, the great northern continents form
the central mass from which the southern regions, as it were, diverge, and
as the Palæarctic region is both more extensive and much better known than
any other, it undoubtedly forms the most convenient starting-point for our
proposed survey of the zoological history of the earth. We thus pass from
the better known to the less known--from Europe to Africa and tropical
Asia, and thence to Australia, completing the series of regions of the
Eastern Hemisphere. Beginning again with the Neotropical region, we pass to
the Nearctic, which has such striking relations with the preceding and with
the Palæarctic region, that it can only be properly understood by constant
reference to both. We thus keep separate the Eastern and Western
hemispheres, which form, from our point of view, the most radical and most
suggestive division of terrestrial faunas; and as we are able to make this
also the dividing point of our two volumes, reference to the work will be
thereby facilitated.
_Cosmopolitan Groups._--Before proceeding to sketch the zoological features
of the several Regions it will be well to notice those family groups which
belong to the earth as a whole, and which are so widely and universally
distributed over it that it will be unnecessary, in some cases, to do more
than refer to them under the separate geographical divisions.
The only absolutely cosmopolitan families of Mammalia are those which are
aerial or marine; and this is one of the striking proofs that their
distribution has been effected by natural causes, and that the permanence
of barriers is one of the chief {176}agencies in the limitation of their
range. Even among the aerial bats, however, only one family--the
Vespertilionidæ--is truly cosmopolitan, the others having a more or less
restricted range. Neither are the Cetacea necessarily cosmopolitan, most of
the families being restricted either to warm or to cold seas; but one
family, the dolphins (Delphinidæ), is truly so. This order however will not
require further notice, as, being exclusively marine the groups do not
enter into any of our terrestrial regions. The only other family of mammals
that may be considered to be cosmopolitan, is the Muridæ (rats and mice);
yet these are not entirely so, since none are known to be truly indigenous
in any part of the Australian region except Australia itself.
In the class of Birds, a number of families are cosmopolites, if we reckon
as such all which are found in each region and sub-region; but several of
these are so abundant in some parts, while they are so sparingly
represented in others, that they cannot fairly be considered so. We shall
confine that term therefore, to such as, there is reason to believe,
inhabit every important sub-division of each region. Such are, among the
Passerine birds the crows (Corvidæ), and swallows (Hirundinidæ); among the
Picariæ the kingfishers (Alcedinidæ); among other Land birds the pigeons
(Columbidæ), grouse and partridges (Tetraonidæ), hawks (Falconidæ), and
owls (Strigidæ); among the Waders the rails (Rallidæ), snipes
(Scolopacidæ), plovers (Charadriadæ), and herons (Ardeidæ); and among the
Swimmers the ducks (Anatidæ), gulls (Laridæ), petrels (Procellariidæ),
pelicans (Pelecanidæ), and grebes (Podicipidæ).
In the class of Reptiles there are few absolutely cosmopolitan families,
owing to the scarcity of members of this group in some insular sub-regions,
such as New Zealand and the Pacific Islands. Those which are most nearly so
are the Colubridæ among snakes, and the Scincidæ among lizards.
There is no cosmopolitan family of Amphibia, the true frogs (Ranidæ) being
the most widely distributed.
Neither is any family of Freshwater Fishes cosmopolitan, the Siluridæ,
which have the widest range, being confined {177}to warm regions, and
becoming very scarce in the temperate zones.
Among the Diurnal and Crepuscular Lepidoptera (butterflies and sphinges)
the following families are cosmopolitan:--Satyridæ, Nymphalidæ, Lycænidæ,
Pieridæ, Papilionidæ, Hesperidæ, Zygænidæ, and Sphingidæ.
Of the Coleoptera almost all, except some of the small and obscure
families, are cosmopolitan.
Of the terrestrial Mollusca, the Helicidæ alone are true cosmopolites.
_Tables of Distribution of Families and Genera._--Having been obliged to
construct numerous tables of the distribution of the various groups for the
purposes of the descriptive part of the work, I have thought it well to
append the most important of them, in a convenient form, to the chapter on
each region; as much information will thereby be given, which can only be
obtained from existing works at the cost of great labour. All these tables
are drawn up on a uniform plan, the same generic and family names being
used in each; and all are arranged in the same systematic order, so as to
be readily comparable with each other. This, although it seems a simple and
natural thing to do, has involved a very great amount of labour, because
hardly two authors use the same names or follow the same arrangement. Hence
comparison between them is impossible, till all their work has been picked
to pieces, their synonymy unravelled, their differences accounted for, and
the materials recast; and this has to be done, not for two or three authors
only, but for the majority of those whose works have been consulted on the
zoology of any part of the globe.
Except in the two higher orders--Mammalia and Birds--materials do not exist
for complete tables of the genera brought down to the present time. We have
given therefore, first, a complete table of all the families of Vertebrata
and Diurnal Lepidoptera found in each region, showing the sub-regions in
which they occur, and their range beyond the limits of the region. Families
which are wholly peculiar to the region, or {178}very characteristic and
almost exclusively confined to it, are in _italics_. The number prefixed to
each family corresponds to that of the series of families in the Fourth
Part of this work, so that if further information is required it can be
readily referred to without consulting the index. Names inclosed in
parentheses--( . . . ) thus--indicate families which only just enter a
region from an adjacent one, to which they properly belong. The eye is thus
directed to the more, and the less important families; and a considerable
amount of information as to the general features of the zoology of the
region, is conveyed in the easiest manner.
The tables of genera of Mammalia and Birds, are arranged on a somewhat
different plan. Each genus is given under its Family and Order, and they
follow in the same succession in all the tables. The number of species of
each genus, inhabiting the region, is given as nearly as can be
ascertained; but in many cases this can only be a general approximation.
The distribution of the genera within the region, is then given with some
detail; and, lastly, the range of the genus beyond the region is given in
general terms, the words "Oriental," "Ethiopian," &c., being used for
brevity, to indicate that the genus occurs over a considerable part of such
regions. Genera which are restricted to the region (or which are very
characteristic of it though just transgressing its limits) are given in
_italics_; while those which only just enter the region from another to
which they really belong, are enclosed in parentheses--( . . . ) thus. The
genera are here numbered consecutively, in order that the number of genera
in each family or each order, in the region, may be readily ascertained (by
one process of subtraction), and thus comparisons made with other regions
or with any other area. As the tables of birds would be swelled to an
inconvenient length by the insertion in each region of all the genera of
Waders and Aquatics, most of which have a very wide range and would have to
be repeated in several or all the regions, these have been omitted; but a
list has been given of such of the genera as are peculiar to, or highly
characteristic of each region.
As this is the first time that any such extensive tables of
{179}distribution have been constructed for the whole of the Mammalia and
Birds, they must necessarily contain many errors of detail; but with all
their imperfections it is believed they will prove very useful to
naturalists, to teachers, and to all who take an intelligent interest in
the wider problems of geography and natural history.
{180}CHAPTER X.
THE PALÆARCTIC REGION.
This region is of immense extent, comprising all the temperate portions of
the great eastern continents. It thus extends from the Azores and Canary
Islands on the west to Japan on the east, a distance not far short of half
the circumference of the globe. Yet so great is the zoological unity of
this vast tract, that the majority of the genera of animals in countries so
far removed as Great Britain and Northern Japan are identical. Throughout
its northern half the animal productions of the Palæarctic region are very
uniform, except that the vast elevated desert-regions of Central Asia
possess some characteristic forms; but in its southern portion, we find a
warm district at each extremity with somewhat contrasted features. On the
west we have the rich and luxuriant Mediterranean sub-region, possessing
many peculiar forms of life, as well as a few which are more especially
characteristic of the Ethiopian region. On the east we have the fertile
plains of Northern China and the rich and varied islands of Japan,
possessing a very distinct set of peculiar forms, with others belonging to
the Oriental region, into which this part of the Palæarctic region merges
gradually as we approach the Tropic of Cancer. Thus, the countries roughly
indicated by the names--Northern Europe, the Mediterranean district,
Central and Northern Asia, and China with Japan--have each well-marked
minor characteristics which entitle them to the rank of sub-regions. Their
boundaries are often indefinable; and those here adopted have been fixed
upon to some extent by considerations of convenience, dependent on custom
and on the more or less perfect knowledge we possess of some of the
intervening countries.
[Illustration: PALAEARCTIC REGION]
{181}_Zoological Characteristics of the Palæarctic Region._--The Palæarctic
region has representatives of thirty-five families of mammalia, fifty-five
of birds, twenty-five of reptiles, nine of amphibia, and thirteen of
freshwater fishes. Comparing it with the only other wholly temperate
region, the Nearctic, we find a much greater variety of types of mammalia
and birds. This may be due in part to its greater area, but more, probably,
to its southern boundary being conterminous for an enormous distance with
two tropical regions, the Ethiopean and Oriental; whereas the Nearctic has
a comparatively short southern boundary conterminous with the Neotropical
region only. This is so very important a difference, that it is rather a
matter of surprise that the two north temperate regions should not be more
unequal in the number of their higher vertebrate forms, than they actually
are.
It is also to the interblending of the Palæarctic with the two adjacent
tropical regions, that we must attribute its possession of so few peculiar
family groups. These are only three; two of reptiles, _Trogonophidæ_ and
_Ophiomoridæ_, and one of fishes, _Comephoridæ_. The number of peculiar
genera is, however, considerable, as the following enumeration will show.
_Mammalia._--The monkey of Gibraltar and North Africa, and an allied
species found in Japan, are now considered to belong to the extensive
eastern genus _Macacus_. The former, however, is peculiar in the entire
absence of the tail, and has by many naturalists, been held to form a
distinct genus, _Inuus_, confined to the Palæarctic region.
Of bats there are one or two genera (_Barbastellus_, _Plecotus_) which seem
to be mainly or wholly Palæarctic, but the classification of these animals
is in such an unsettled state that the distribution of the genera is of
little importance.
In the next order, Insectivora, we have almost the entire family of the
Moles confined to the region. _Talpa_ just enters Northern India; and
_Urotrichus_ is common to Japan and {182}North-Western America, but the
remaining genera, six in number, are all exclusively Palæarctic.
Among Carnivora we have _Nyctereutes_, the curious racoon-dog of Japan and
North-Eastern Asia; _Lutronectes_, an otter peculiar to Japan; and the
badger (_Meles_), which ranges over the whole region, and just enters the
Oriental region as far as Hongkong; _Æluropus_, a curious form of the
Himalayan panda, inhabiting the high mountains of Eastern Thibet; and
_Pelagius_, a genus of seals, ranging from the shores of Madeira to the
Black Sea.
The Ungulata, or hoofed animals, are still more productive of forms
peculiar to this region. First we have the Camels, whose native home is the
desert region of Central and Western Asia and Northern Africa, and which,
even in their domesticated condition, are confined almost wholly within the
limits of the Palæarctic region. Of Deer we have six peculiar genera,
_Dama_ and _Capreolus_ found in Europe, with _Elaphodus_, _Lophotragus_,
_Hydropotes_, and _Moschus_, confined to Northern China and Mongolia. The
great family Bovidæ--comprising the oxen, sheep, goats and
antelopes--furnishes no less than seven peculiar Palæarctic genera. These
are _Poephagus_, the yak of Thibet; _Addax_, a well-known antelope of
Northern Africa and Syria; _Procapra_, _Pantholops_ and _Budorcas_,
antelopine genera peculiar to Thibet and Mongolia; with _Rupicapra_ (the
chamois), and the extraordinary large-nosed antelope _Saiga_, confined to
Europe and Western Asia. Besides these we have _Capra_ (the wild sheep and
goats), all the numerous species of which, except two, are exclusively
Palæarctic.
Coming to the Rodents, we have again many peculiar forms. Of Muridæ (the
mouse and rat tribe), we have six peculiar genera, the more important being
_Cricetus_, _Rhombomys Sminthus_, and _Myospalax_. Of Spalacidæ (mole-rats)
both the Palæarctic genera, _Ellobius_ and _Spalax_, are peculiar.
_Ctenodactylus_, a genus of the South American family Octodontidæ, is found
only in North Africa. To these we may add _Myoxus_ (the dormice) and
_Lagomys_ (the pikas or tail-less hares) as essentially Palæarctic, since
but one species of each genus is found beyond the limits of the region.
_Birds._--It appears to have been the opinion of many {183}naturalists that
the Palæarctic region could not be well characterised by its peculiar
genera of birds. In Mr. Sclater's celebrated paper already referred to, he
remarks, "It cannot be denied that the ornithology of the Palæarctic region
is more easily characterised by what it has not than by what it has," and
this has been quite recently quoted by Mr. Allen, in his essay on the
distribution of North American birds, as if it represented our present
knowledge of the subject. But, thanks to the labours of Dr. Jerdon, Mr.
Swinhoe, Père David and others, we have now learnt that a large number of
birds included in the Indian list, are either mere winter emigrants from
Central Asia, or only inhabit the higher ranges of the Himalayas, and thus
really belong to the Palæarctic region. The result is, that a host of
genera are now seen to be either exclusively or characteristically
Palæarctic, and we have no further difficulty in giving positive
ornithological characters to the region. In the tables appended to this
chapter, all these truly Palæarctic genera will be found printed in
_italics_, with an indication of their distribution, which will sometimes
be found more fully given under the respective families in the fourth part
of this work. Referring to this table for details we shall here summarise
the results.
Of the Sylviidæ or warblers, no less than fourteen genera are either
exclusively or characteristically Palæarctic, of which _Locustella_,
_Sylvia_, _Curruca_ and _Erithacus_ are good examples. Of the oriental
family Timaliidæ, the genus _Pterorhinus_ is Palæarctic. Of Panuridæ, or
reedlings, there are four peculiar genera (comprising almost the whole
family); of Certhiidæ, or creepers, one--_Tichodroma_--which extends
southward to the Abyssinian highlands. Of Paridæ, or tits, one--_Acredula_;
of Corvidæ, or crows, four--_Pica_ (containing our magpie) being a good
example; of Fringillidæ, or finches and buntings, twelve, among which
_Acanthis_, _Pyrrhula_ and _Emberiza_ are good illustrations; of Alaudidæ,
or larks, there are two peculiar genera. Leaving the Passeres we next come
to peculiar forms among the gallinaceous birds: _Syrrhaptes_ among the
Pteroclidæ or sand grouse; four genera of Tetraonidæ or grouse and
partridges, and five of Phasianidæ or pheasants, comprising some of the
most magnificent birds in the world. Lastly {184}among the far-wandering
aquatic birds we have no less than five genera which are more especially
Palæarctic,--_Ortygometra_, the corn-crake, and _Otis_, the great bustard,
being typical examples. We may add to these, several genera almost confined
to this region, such as _Garrulus_ (jays), _Fringilla_ (true finches),
_Yunx_ (wrynecks) and some others; so that in proportion to its total
generic forms a very large number are found to be peculiar or
characteristic.
This view, of the high degree of speciality of the Palæarctic region, will
no doubt be objected to by some naturalists, on the ground that many of the
genera reckoned as exclusively Palæarctic are not so, but extend more or
less into other regions. It is well, therefore, to consider what principles
should guide us in a matter of this kind, especially as we shall have to
apply the same rules to each of the other regions. We may remark first,
that the limits of the regions themselves are, when not formed by the
ocean, somewhat arbitrary, depending on the average distribution of a
number of characteristic forms; and that slight local peculiarities of
soil, elevation, or climate, may cause the species of one region to
penetrate more or less deeply into another. The land boundary between two
regions will be, not a defined line but a neutral territory of greater or
less width, within which the forms of both regions will intermingle; and
this neutral territory itself will merge imperceptibly into both regions.
So long therefore as a species or genus does not permanently reside
considerably beyond the possible limits of this neutral territory, we
should not claim it as an inhabitant of the adjacent region. A
consideration of perhaps more importance arises, from the varying extent of
the range of a genus, over the area occupied by the region. Some genera are
represented by single species existing only in a very limited area; others
by numerous species which occupy, entirely or very nearly, the whole extent
of the region; and there is every intermediate grade between these
extremes. Now, the small localised genera, are always reckoned as among the
best examples of types peculiar to a region; while the more wide-spread
groups are often denied that character if they extend a little beyond
{185}the supposed regional limits, or send one or two, out of a large
number of species, into adjacent regions; yet there is some reason to
believe that the latter are really more important as characterising a
zoological region than the former. In the case of a single isolated species
or genus we have a dying-out group; and we have so many cases of
discontinuous species of such groups (of which _Urotrichus_ in Japan and
British Columbia, _Eupetes_ in Sumatra and New Guinea are examples), that
it is quite as probable as not, that any such isolated species has only
become peculiar to the region by the recent extinction of an allied form or
forms in some other region. On the other hand, a genus consisting of
numerous species ranging over an entire region or the greater part of one,
is a dominant group, which has most likely been for some time extending its
range, and whose origin dates back to a remote period. The slight extension
of such a group beyond the limits of the region to which it mainly belongs,
is probably a recent phenomenon, and in that case cannot be held in any
degree to detract from its value as one of the peculiar forms of that
region.
The most numerous examples of this class, are those birds of the temperate
regions which in winter migrate, either wholly or partially, into adjacent
warmer countries. This migration most likely began subsequent to the
Miocene period, during that gradual refrigeration of the temperate zones
which culminated in the glacial epoch, and which still continues in a
mitigated form. Most of the genera, and many even of the species of birds
which migrate southwards in winter, have therefore, most likely, always
been inhabitants of our present Palæarctic and Nearctic regions; permanent
residents during warm epochs, but only able now to maintain their existence
by migration in winter. Such groups belong truly to the temperate zones,
and the test of this is the fact of their not having any, or very few,
representatives, which are permanent residents in the adjacent tropical
regions. When there are such representative species, we do not claim them
as peculiar to the Northern regions. Bearing in mind these various
considerations, it will be found that we have been very moderate in our
estimate of the number of genera {186}that may fairly be considered as
exclusively or characteristically Palæarctic.
_Reptiles and Amphibia._--The Palæarctic region possesses, in proportion to
its limited reptilian fauna, a full proportion of peculiar types. We have
for instance two genera of snakes, _Rhinechis_ and _Halys_; seven of
lizards, _Trigonophis_, _Psammodromus_, _Hyalosaurus_, _Scincus_,
_Ophiomorus_, _Megalochilus_, and _Phrynocephalus_; eight of tailed
batrachians, _Proteus_, _Salamandra_, _Seiranota_, _Chioglossa_,
_Hynobius_, _Onychodactylus_, _Geotriton_, and _Sieboldia_; and eight of
tail-less batrachians, _Bombinator_, _Pelobates_, _Didocus_, _Alytes_,
_Pelodytes_, _Discoglossus_, _Laprissa_, and _Latonia_. The distribution of
these and other Palæarctic genera will be found in our second vol. chap.
xix.
_Freshwater Fish._--About twenty genera of freshwater fishes are wholly
confined to this region, and constitute a feature which ought not to be
overlooked in estimating its claim to the rank of a separate primary
division of the earth. They belong to the following families:--Percidæ
(three genera), _Acerina_, _Percarina_, _Aspro_; Comephoridæ (one genus),
_Comephorus_, found only in Lake Baikal; Salmonidæ (three genera),
_Brachymystax_, _Luciotrutta_, and _Plecoglossus_; Cyprinodontidæ (one
genus), _Tellia_, found only in Alpine pools on the Atlas Mountains;
Cyprinidæ (thirteen genera), _Cyprinus_, _Carassus_, _Paraphoxinus_,
_Tinca_, _Achilognathus_, _Rhodeus_, _Chondrostoma_, _Pseudoperilampus_,
_Ochetebius_, _Aspius_, _Alburnus_, _Misgurnus_, and _Nemachilus_.
_Summary of Palæarctic Vertebrata._--Summarising these details, we find
that the Palæarctic region possesses thirty-five peculiar genera of
mammalia, fifty-seven of birds, nine of reptiles, sixteen of amphibia, and
twenty-one of freshwater fishes; or a total of 138 peculiar generic types
of vertebrata. Of these, 87 are mammalia and land-birds out of a total of
274 genera of these groups; or rather less than one-third peculiar, a
number which will serve usefully to compare with the results obtained in
other regions.
In our chapter on Zoological Regions we have already pointed out the main
features which distinguish the Palæarctic from the Oriental and Ethiopian
regions. The details now given will {187}strengthen our view of their
radical distinctness, by showing to how considerable an extent the former
is inhabited by peculiar, and often very remarkable generic types.
_Insects: Lepidoptera._--The Diurnal Lepidoptera, or butterflies, are not
very abundant in species, their number being probably somewhat over 500,
and these belong to not more than fifty genera. But no less than fifteen of
these genera are wholly confined to the region. Nine of the families are
represented, as follows:--1. _Danaidæ_; having only a single species in
South Europe. 2. _Satyridæ_; well represented, there being more than 100
species in Europe, and three peculiar genera. 3. _Nymphalidæ_; rather
poorly represented, Europe having only about sixty species, but there is
one peculiar genus. 4. _Libytheidæ_; a very small family, represented by a
single species occurring in South Europe. 5. _Nemeobiidæ_; a rather small
family, also having only one species in Europe, but which constitutes a
peculiar genera. 6. _Lycænidæ_; an extensive family, fairly represented,
having about eighty European species; there are two peculiar genera in the
Palæarctic region. 7. _Pieridæ_; rather poorly represented with thirty-two
European species; two of the genera are, however, peculiar. 8.
_Papilionidæ_; very poorly represented in Europe with only twelve species,
but there are many more in Siberia and Japan. No less than five of the
small number of genera in this family are wholly confined to the region, a
fact of much importance, and which to a great extent redeems the character
of the Palæarctic region as regard this order of insects. Their names are
_Mesapia_, _Hypermnestra_, _Doritis_, _Sericinus_, and _Thais_; and besides
these we have _Parnassius_--the "Apollo" butterflies--highly
characteristic, and only found elsewhere in the mountains of the Nearctic
region. 9. _Hesperidæ_; poorly represented with about thirty European
species, and one peculiar genus.
Four families of _Sphingina_ occur in the Palæarctic region, and there are
several peculiar genera.
In the _Zygænidæ_ there are two exclusively European genera, and the
extensive genus _Zygæna_ is itself mainly Palæarctic. The small family
_Stygiidæ_ has two out of its three genera {188}confined to the Palæarctic
region. In the _Ægeriidæ_ the genus _Ægeria_ is mainly Palæarctic. The
_Sphingidæ_ have a wider general range, and none of the larger genera are
peculiar to any one region.
_Coleoptera._--The Palæarctic region is the richest portion of the globe in
the great family of _Carabidæ_, or predacious ground-beetles, about 50 of
the genera being confined to it, while many others, including the
magnificent genus _Carabus_, have here their highest development. While
several of the smaller genera are confined to the eastern or western
sub-regions, most of the larger ones extend over the whole area, and give
it an unmistakable aspect; while in passing from east to west or
_vice-versâ_, allied species and genera replace each other with
considerable regularity, except in the extreme south-east, where, in China
and Japan, some Oriental forms appear, as do a few Ethiopian types in the
south-west.
Cicindelidæ, or tiger-beetles, are but poorly represented by about 70
species of the genus _Cicindela_, and a single _Tetracha_ in South Europe.
Lucanidæ, or stag-beetles, are also poor, there being representatives of 8
genera. One of these, _Æsalus_ (a single species), is peculiar to South
Europe, and two others, _Cladognathus_ and _Cyclopthalmus_, are only
represented in Japan, China, and Thibet.
Cetoniidæ, or rose-chafers, are represented by 13 genera, two of which are
peculiar to South Europe (_Tropinota_ and _Heterocnemis_), while
_Stalagmosoma_, ranging from Persia to Nubia, and the fine
_Dicranocephalus_ inhabiting North China, Corea, and Nipal, may also be
considered to belong to it. The genera _Trichius_, _Gnorimus_, and
_Osmoderma_ are confined to the two north temperate regions.
Buprestidæ, or metallic beetles, are rather abundant in the warmer parts of
the region, 27 genera being represented, nine of which are peculiar. By far
the larger portion of these are confined to the Mediterranean sub-region. A
considerable number also inhabit Japan and China.
The Longicorns, or long-horned beetles, are represented by no less than 196
genera, 51 of which are peculiar. They are {189}much more abundant in the
southern than the northern half of the region. Several Oriental genera
extend to Japan and North China, and a few Ethiopian genera to North
Africa. Thirteen genera are confined, to the two north temperate regions.
Several large genera, such as _Dorcadion_ (154 species), _Phytæcia_ (85
species), _Pogonochærus_ (22 species), _Agapanthia_ (22 species), and
_Vesperus_ (7 species), are altogether peculiar to the Palæarctic region;
and with a preponderance of _Leptura_, _Grammoptera_, _Stenocorus_, and
several others, strongly characterise it as distinct from the Nearctic and
Oriental regions.
The other families which are well developed in the Palæarctic regions, are,
the Staphylinidæ or rove-beetles, Silphidæ or burying-beetles, Histeridæ or
mimic-beetles, Nitidulidæ, Aphodiidæ, Copridæ (especially in South Europe),
Geotrupidæ or dung-beetles, Melolonthidæ or chafers, Elateridæ or
click-beetles, the various families of Malacoderms and Heteromera,
especially Pimeliidæ in the Mediterranean sub-region, Curculionidæ or
weevils, the Phytophaga or leaf-eaters, and Cocinellidæ or lady-birds.
The number of species of Coleoptera in the western part of the Palæarctic
region is about 15,000, and there are probably not more than 2,000 to add
to this number from Siberia, Japan, and North China; but were these
countries as well explored as Europe, we may expect that they would add at
least 5,000 to the number above given, raising the Palæarctic Coleopterous
fauna to 20,000 species. As the total number of species at present known to
exist in collections is estimated (and perhaps somewhat over-estimated) at
70,000 species, we may be sure that were the whole earth as thoroughly
investigated as Europe, the number would be at least doubled, since we
cannot suppose that Europe, with the Mediterranean basin, can contain more
than one-fifth of the whole of the Coleoptera of the globe.
Of the other orders of insects we here say nothing, because in their case
much more than in that of the Coleoptera and Lepidoptera, is the
disproportion enormous between our knowledge of the European fauna and that
of almost all the rest of the globe. {190}They are, therefore, at present
of comparatively little use for purposes of geographical distribution, even
were it advisable to enter into the subject in a work which will, perhaps,
be too much overburdened with details only of interest to specialists.
_Land Shells._--These are very numerous in the warmer parts of the region,
but comparatively scarce towards the North. South Europe alone possesses
over 600 species, whereas there are only 200 in all Northern Europe and
Asia. The total number of species in the whole region is probably about
1,250, of which the great majority are Helicidæ; the Operculated families
being very poorly represented. Several small genera or sub-genera are
peculiar to the region, as _Testacella_ (West Europe and Canaries);
_Leucochroa_ (Mediterranean district); _Acicula_ (Europe); _Craspedopoma_
(Atlantic Islands); _Leonia_ (Algeria and Spain); _Pomatias_ (Europe and
Canaries); _Cecina_ (Mongolia). The largest genera are _Helix_ and
_Clausilia_, which together comprise more than half the species; _Pupa_,
very numerous; _Bulimus_ and _Achatina_ in moderate numbers, and all the
rest small. _Helix_ is the only genus which contains large and handsome
species; _Bulimus_ and _Achatina_, so magnificent in tropical countries,
being here represented by small and obscure forms only. _Daudebardia_ is
confined to Central and South Europe and New Zealand; _Glandina_ is chiefly
South American; _Hyalina_ is only American and European; _Buliminus_ ranges
over all the world except America; and the other European genera of
Helicidæ are widely distributed. Of the Operculata, _Cyclotus_,
_Cyclophorus_, and _Pupina_ extend from the Oriental region into Japan and
North China; _Tudoria_ is found in Algeria and the West Indies; _Hydrocena_
is widely scattered, and occurs in South Europe and Japan. The genera of
freshwater shells are all widely distributed.
THE PALÆARCTIC SUB-REGIONS.
The four sub-regions which are here adopted, have been fixed upon as those
which are, in the present state of our knowledge, at once the most natural
and the only practicable ones. {191}No doubt all of them could be
advantageously again subdivided, in a detailed study of the geographical
distribution of _species_. But in a general work, which aims at treating
all parts of the world with equal fulness, and which therefore is confined
almost wholly to the distribution of families and genera, such further
subdivision would be out of place. It is even difficult, in some of the
classes of animals, to find peculiar or even characteristic genera for the
present sub-regions; but they all have well marked climatic and physical
differences, and this leads to an assemblage of species and of groups which
are sufficiently distinctive.
_I. Central and Northern Europe._
This sub-region, which may perhaps be termed the "European," is
zoologically and botanically the best known on the globe. It can be pretty
accurately defined, as bounded on the south by the Pyrenees, the Alps, the
Balkans, the Black Sea, and the Caucasus range; and by the Ural Mountains,
or perhaps more correctly the valley of the Irtish and Caspian Sea, on the
east; while Ireland and Iceland are its furthest outliers in the west. To
the north, it merges so gradually into the Arctic zone that no demarcation
is possible. The great extent to which this sub-region is interpenetrated
by the sea, and the prevalence of westerly winds bringing warmth and
moisture from an ocean influenced by the gulf-stream, give it a climate for
the most part genial, and free from extremes of heat and cold. It is thus
broadly distinguished from Siberia and Northern Asia generally, where a
more extreme and rigorous climate prevails.
The whole of this sub-region is well watered, being penetrated by rivers in
every direction; and it consists mainly of plains and undulating country of
moderate elevation, the chief mountain ranges being those of Scandinavia in
the north-west, and the extensive alpine system of Central Europe. But
these are both of moderate height, and a very small portion of their
surface is occupied either by permanent snow-fields, or by barren uplands
inimical to vegetable and animal life. It is, in {192}fact, to these, and
the numerous lesser mountains and hills which everywhere diversify the
surface of Europe, that the variety and abundance of its animal life is
greatly due. They afford the perennial supplies to rivers, and furnish in
their valleys and ever varying slopes, stations suited to every form of
existence. A considerable area of Central Europe is occupied by uplands of
moderate elevation, a comparatively small portion being flat and marshy
plains.
Most of the northern and much of the central portions of Europe are covered
with vast forests of coniferous trees; and these, occupying as they do
those tracts where the winter is most severe, supply food and shelter to
many animals who could not otherwise maintain their existence. It is
probable that the original condition of the greater part, if not the whole,
of temperate Europe, except the flat marshes of the river valleys and the
sandy downs of the coast, was that of woodland and forest, mostly of
deciduous trees, but with a plentiful admixture of such hardy evergreens as
holly, ivy, privet, and yew. A sufficient proportion of these primeval
woods, and of artificial plantations which have replaced them, fortunately
remain, to preserve for us most of the interesting forms of life, which
were developed before man had so greatly modified the surface of the earth,
and so nearly exterminated many of its original tenants. Almost exactly in
proportion to the amount of woodland that still remains in any part of
Europe, do we find (other things being equal) the abundance and variety of
wild animals; a pretty clear indication that the original condition of the
country was essentially that of a forest, a condition which only now exists
in the thinly inhabited regions of the north.
Although the sub-region we are considering is, for its extent and latitude,
richly peopled with animal life, the number of genera altogether peculiar
to it is not great. There are, however, several which are very
characteristic, and many species, both of the smaller mammalia and of
birds, are wholly restricted to it.
_Mammalia._--The genera wholly confined to this sub-region are {193}only
two. _Myogale_, the desman, is a curious long-snouted Insectivorous animal
somewhat resembling the water-rat in its habits. There are two species, one
found only on the banks of streams in the French Pyrenees, the other on the
great rivers of Southern Russia. The other peculiar genus, _Rupicapra_ (the
chamois of the Alps), is found on all the high mountains of Central Europe.
Almost peculiar are _Spalax_ (the mole-rat) found only in Eastern Europe
and Western Siberia; and _Saiga_, an extraordinary large-nosed antelope
which has a nearly similar distribution. Highly characteristic forms, which
inhabit nearly every part of the sub-region, are, _Talpa_ (the mole),
_Erinaceus_, (the hedgehog), _Sorex_ (the shrew), _Meles_ (the badger),
_Ursus_ (the bear), _Canis_ (the wolf and fox), _Mustela_ (the weasel),
_Lutra_ (the otter), _Arvicola_ (the vole), _Myoxus_ (the dormouse), and
_Lepus_ (the hare and rabbit); while _Bos_ (the wild bull) was, until
exterminated by man, no doubt equally characteristic. Other genera
inhabiting the sub-region will be found in the list given at the end of
this chapter.
_Birds._--It is difficult to name the birds that are most characteristic of
this sub-region, because so many of the most familiar and abundant are
emigrants from the south, and belong to groups that have a different range.
There is perhaps not a single genus wholly confined to it, and very few
that have not equal claims to be placed elsewhere. Among the more
characteristic we may name _Turdus_ (the thrushes), _Sylvia_ (the
warblers), _Panurus_ (the reedling), _Parus_ (the tits), _Anthus_ (the
pipits), _Motacilla_ (the wagtails), which are perhaps more abundant here
than in any other part of the world, _Emberiza_ (the buntings),
_Plectrophanes_ (the snow buntings), _Passer_ (the house sparrows), _Loxia_
(the crossbills), _Linota_ (the linnets), _Pica_ (the magpies), _Tetrao_
(grouse), _Lagopus_ (ptarmigan) and many others.
I am indebted to Mr. H. E. Dresser, who is personally acquainted with the
ornithology of much of the North of Europe, for some valuable notes on the
northern range of many European birds. Those which are characteristic of
the extreme Arctic zone, extending beyond 70° north latitude, and tolerably
abundant, are two falcons (_Falco gyrfalco_ and _F. peregrinus_); {194}the
rough-legged buzzard (_Archibuteo lagopus_); the snowy owl (_Nyctea
scandiaca_); the raven (_Corvus corax_); three buntings (_Emberiza
schæniculus_, _Plectrophanes nivalis_ and _P. calcarata_); a lark
(_Otocorys alpestris_); several pipits, the most northern being _Anthus
cervinus_; a wagtail (_Budytes cinereocapilla_); a dipper (_Cinclus
melanogaster_); a warbler (_Cyanecula suecica_); the wheatear (_Saxicola
oenanthe_); and two ptarmigans (_Lagopus albus_ and _L. salicetus_). Most
of these birds are, of course, only summer visitors to the Arctic regions,
the only species noted as a permanent resident in East Finmark (north of
latitude 70°) being the snow-bunting (_Plectrophanes nivalis_).
The birds that are characteristic of the zone of pine forests, or from
about 61° to 70° north latitude, are very numerous, and it will be
sufficient to note the genera and the number of species (where more than
one) to give an idea of the ornithology of this part of Europe. The birds
of prey are, _Falco_ (three species), _Astur_ (two species), _Buteo_,
_Pandion_, _Surnia_, _Bubo_, _Syrnium_, _Asio_, _Nyctala_. The chief
Passerine birds are, _Corvus_ (two species), _Pica_, _Garrulus_ (two
species), _Nucifraga_, _Bombycilla_, _Hirundo_ (two species), _Muscicapa_
(two species), _Lanius_, _Sturnus_, _Passer_ (two species), _Pyrrhula_,
_Carpodacus_, _Loxia_ (two species), _Pinicola_, _Fringilla_ (eight
species), _Emberiza_ (five species), _Alauda_, _Anthus_, _Turdus_ (five
species), _Ruticilla_, _Pratincola_, _Accentor_, _Sylvia_ (four species),
_Hypolais_, _Regulus_, _Phylloscopus_ (two species), _Acrocephalus_,
_Troglodytes_, and _Parus_ (six species). Woodpeckers are abundant, _Picus_
(four species), _Gecinus_, and _Yunx_. The kingfisher (_Alcedo_),
goatsucker (_Caprimulgus_), and swift (_Cypselus_) are also common. The
wood-pigeon (_Columba_) is plentiful. The gallinaceous birds are three
grouse, _Tetrao_ (two species) and _Bonasa_, and the common quail
(_Coturnix_).
The remaining genera and species of temperate or north-European birds, do
not usually range beyond the region of deciduous trees, roughly indicated
by the parallel of 60° north latitude.
Plate I.
[Illustration]
THE ALPS OF CENTRAL EUROPE, WITH CHARACTERISTIC ANIMALS.
{195}_Plate I.--Illustrating the Zoology of Central Europe._--Before
considering the distribution of the other classes of vertebrata, it will be
convenient to introduce our first illustration, which represents a scene in
the Alps of Central Europe, with figures of some of the most characteristic
Mammalia and Birds of this sub-region. On the left is the badger (_Meles
Taxus_) one of the weasel family, and belonging to a genus which is
strictly Palæarctic. It abounds in Central and Northern Europe and also
extends into North Asia, but is represented by another species in Thibet
and by a third in Japan. The elegantly-formed creatures on the right are
chamois (_Rupicapra tragus_), almost the only European antelopes, and
wholly confined to the higher mountains, from the Pyrenees to the
Carpathians and the Caucasus. The chamois is the only species of the genus,
and is thus perhaps the most characteristic European mammal. The bird on
the left, above the badgers, is the Alpine chough, (_Fregilus
pyrrhocorax_). It is found in the high mountains from the Alps to the
Himalayas, and is allied to the Cornish chough, which is still found on our
south-western coasts, and which ranges to Abyssinia and North China. The
Alpine chough differs in having a shorter bill of an orange colour, and
vermilion red feet as in the other species. In the foreground are a pair of
ruffs (_Machetes pugnax_) belonging to the Scolopacidæ or snipe family, and
most nearly allied to the genus _Tringa_ or sandpiper. This bird is
remarkable for the fine collar of plumes which adorns the males in the
breeding season, when they are excessively pugnacious. It is the only
species of its genus, and ranges over all Europe and much of Northern Asia,
migrating in the winter to the plains of India, and even down the east
coast of Africa as far as the Cape of Good Hope; but it only breeds in the
Palæarctic region, over the greater part of which it ranges.
_Reptiles and Amphibia._--There are no genera of reptiles peculiar to this
sub-region. Both snakes and lizards are comparatively scarce, there being
about fourteen species of the former and twelve of the latter. Our common
snake (_Tropidonotus natrix_) extends into Sweden and North Russia, but the
viper (_Viperus berus_) goes further north, as far as Archangel (64° N.),
and in Scandinavia (67° N.), and is the most Arctic of all known
{196}snakes. Of the lizards, _Lacerta stirpium_ (the sand lizard) has the
most northerly range, extending into Poland and Northern Russia; and
_Anguis fragilis_ (the blind or slow-worm) has almost an equal range.
Amphibia, being more adapted to a northern climate, have acquired a more
special development, and thus several forms are peculiar to the North
European sub-region. Most remarkable is _Proteus_, a singular eel-like
aquatic creature with small legs, found only in the subterranean lakes in
Carniola and Carinthia; _Alytes_, a curious toad, the male of which carries
about the eggs till they are hatched, found only in Central Europe from
France to the east of Hungary; and _Pelodytes_, a frog found only in
France. Frogs and toads are very abundant all over Europe, the common frog
(_Rana temporaria_) extending to the extreme north. The newts (_Triton_)
are also very abundant and widely spread, though not ranging so far north
as the frogs. The genera _Bombinator_ (a toad-like frog), and _Hyla_ (the
tree frog) are also common in Central Europe.
_Freshwater Fish._--Two genera of the perch family (Percidæ) are peculiar
to this sub-region,--_Percarina_, a fish found only in the river Dniester,
and _Aspro_, confined to the rivers of Central Europe. Of the very
characteristic forms are, _Gasterosteus_ (stickle-back), which alone forms
a peculiar family--Gasterosteidæ; _Perca_, _Acerina_ and _Lucioperca_,
genera of the perch family; _Silurus_, a large fish found in the rivers of
Cenrtal Europe, of the family Siluridæ; _Esox_ (the pike), of the family
Esocidæ; _Cyprinus_ (carp), _Gobio_ (gudgeon), _Leuciscus_ (roach, chub,
dace, &c.), _Tinca_ (tench), _Abramus_ (bream), _Alburnus_ (bleak),
_Cobitis_ (loach), all genera of the family Cyprinidæ.
_Insects--Lepidoptera._--No genera of butterflies are actually confined to
this sub-region, but many are characteristic of it. _Parnassius_, _Aporia_,
_Leucophasia_, _Colias_, _Melitæa_, _Argynnis_, _Vanessa_, _Limenitis_, and
_Chionobas_, are all very abundant and widespread, and give a feature to
the entomology of most of the countries included in it.
_Coleoptera._--This sub-region is very rich in Carabidæ; the genera
_Elaphrus_, _Nebria_, _Carabus_, _Cychrus_, _Pterostichus_, _Amara_,
{197}_Trechus_ and _Peryphus_ being especially characteristic. Staphylinidæ
abound. Among Lamellicorns the genus _Aphodius_ is most characteristic.
Buprestidæ are scarce; Elateridæ more abundant. Among Malacoderms
_Telephorus_ and _Malachius_ are characteristic. Curculionidæ abound:
_Otiorhyuchus_, _Omias_, _Erirhinus_, _Bagous_, _Rhynchites_ and
_Ceutorhynchus_ being very characteristic genera. Of Longicorns
_Callidium_, _Dorcadion_, _Pogonochærus_, _Pachyta_ and _Leptura_ are
perhaps the best representatives. _Donacia_, _Crioceris_, _Chrysomela_, and
_Altica_, are typical Phytophaga; while _Coccinella_ is the best
representative of the Securipalpes.
_North European Islands._--The British Islands are known to have been
recently connected with the Continent, and their animal productions are so
uniformly identical with continental species as to require no special note.
The only general fact of importance is, that the number of species in all
groups is much less than in continental districts of equal extent, and that
this number is still farther diminished in Ireland. This may be accounted
for by the smaller area and less varied surface of the latter island; and
it may also be partly due to the great extent of low land, so that a very
small depression would reduce it to the condition of a cluster of small
islands capable of supporting a very limited amount of animal life. Yet
further, if after such a submergence had destroyed much of the higher forms
of life in Great Britain and Ireland, both were elevated so as to again
form part of the Continent, a migration would commence by which they would
be stocked afresh; but this migration would be a work of time, and it is to
be expected that many species would never reach Ireland or would find its
excessively moist climate unsuited to them.
Some few British species differ slightly from their continental allies, and
are considered by many naturalists to be distinct. This is the case with
the red grouse (_Lagopus scoticus_) among birds; and a few of the smaller
Passeres have also been found to vary somewhat from the allied forms on the
Continent, showing that the comparatively short interval since the glacial
period, and the slightly different physical conditions dependent on
{198}insularity, have sufficed to commence the work of specific
modification. There are also a few small land-shells and several insects
not yet found elsewhere than in Britain; and even one of the smaller
Mammalia--a shrew (_Sorex rusticus_). These facts are all readily explained
by the former union of these islands with the Continent, and the alternate
depressions and elevations which are proved by geological evidence to have
occurred, by which they have been more than once separated and united again
in recent times. For the evidence of this elevation and depression, the
reader may consult Sir Charles Lyell's _Antiquity of Man_.
Iceland is the only other island of importance belonging to this
sub-region, and it contrasts strongly with Great Britain, both in its
Arctic climate and oceanic position. It is situated just south of the
Arctic circle and considerably nearer Greenland than Europe, yet its
productions are almost wholly European. The only indigenous land mammalia
are the Arctic fox (_Canis lagopus_), and the polar bear as an occasional
visitant, with a mouse (_Mus islandicus_), said to be of a peculiar
species. Four species of seals visit its shores. The birds are more
interesting. According to Professor Newton, ninety-five species have been
observed; but many of these are mere stragglers. There are twenty-three
land, and seventy-two aquatic birds and waders. Four or five are peculiar
species, though very closely related to others inhabiting Scandinavia or
Greenland. Only two or three species are more nearly related to Greenland
birds than to those of Northern Europe, so that the Palæarctic character of
the fauna is unmistakable. The following lists, compiled from a paper by
Professor Newton, may be interesting as showing more exactly the character
of Icelandic ornithology.
1. Peculiar species.--_Troglodytes borealis_ (closely allied to the common
wren, found also in the Faroe Islands); _Falco islandicus_ (closely allied
to _F. gyrfalco_); _Lagopus islandorum_ (closely allied to _L. rupestris_
of Greenland).
2. European species resident in Iceland.--_Emberiza nivalis_, _Corvus
corax_, _Haliæetus albicilla_, _Rallus aquaticus_, _Hæmatopus ostralegus_,
_Cygnus ferus_, _Mergus_ (two species), _Phalacocorax_ (two {199}species),
_Sula bassana_, _Larus_ (two species), _Stercorarius catarractes_,
_Puffinus anglorum_, _Mergulus alle_, _Uria_ (three species), _Alca torda_.
3. American species resident in Iceland.--_Clangula islandica_,
_Histrionicus torquatus_.
4. Annual visitants from Europe.--_Turdus iliacus_, _Ruticilla tithys_,
_Saxicola ænanthe_, _Motacilla alba_, _Anthus pratensis_, _Linota linaria_,
_Chelidon urbica_, _Hirundo rustica_, _Falco æsalon_, _Surnia nyctea_,
_Otus brachyotus_, _Charadrius pluvialis_, _Ægialites hiaticula_,
_Strepsilas interpres_, _Phalaropus fulicarius_, _Totanus calidris_,
_Limosa_ (species), _Tringa_ (three species), _Calidris arenaria_,
_Gallinago media_, _Numenius phæopus_, _Ardea cinerea_, _Anser_ (two
species), _Bernicla_ (two species), _Anas_ (four species), _Fuligula
marila_, _Harelda glacialis_, _Somateria mollissima_, _Oedemia nigra_,
_Sterna macrura_, _Rissa tridactyla_, _Larus luecopterus_, _Stercorarius_
(two species), _Fratercula artica_, _Colymbus_ (two species), _Podiceps
cornutus_.
5. Annual visitant from Greenland.--_Falco candicans_.
6.--Former resident, now extinct.--_Alca impennis_ (the great auk).
_II.--Mediterranean Sub-region._
This is by far the richest portion of the Palæarctic region, for although
of moderate extent much of it enjoys a climate in which the rigours of
winter are almost unknown. It includes all the countries south of the
Pyrenees, Alps, Balkans, and Caucasus mountains; all the southern shores of
the Mediterranean to the Atlas range, and even beyond it to include the
extra-tropical portion of the Sahara; and in the Nile valley as far as the
second cataract. Further east it includes the northern half of Arabia and
the whole of Persia, as well as Beluchistan, and perhaps Affghanistan up to
the banks of the Indus. This extensive district is almost wholly a region
of mountains and elevated plateaus. On the west, Spain is mainly a
table-land of more than 2000 feet elevation, deeply penetrated by extensive
valleys and rising into lofty mountain chains. Italy, Corsica, Sardinia,
and Sicily, are all very {200}mountainous, and much of their surface
considerably elevated. Further east we have all European Turkey and Greece,
a mountain region with a comparatively small extent of level plain. In Asia
the whole country, from Smyrna through Armenia and Persia to the further
borders of Affghanistan, is a vast mountainous plateau, almost all above
2000, and extensive districts above 5000 feet in elevation. The only large
tract of low-land is the valley of the Euphrates. There is also some
low-land south of the Caucasus, and in Syria the valley of the Jordan. In
North Africa the valley of the Nile and the coast plains of Tripoli and
Algiers are almost the only exceptions to the more or less mountainous and
plateau-like character of the country. Much of this extensive area is now
bare and arid, and often even of a desert character; a fact no doubt due,
in great part, to the destruction of aboriginal forests. This loss is
rendered permanent by the absence of irrigation, and, it is also thought,
by the abundance of camels and goats, animals which are exceedingly
injurious to woody vegetation, and are able to keep down the natural growth
of forests. Mr. Marsh (whose valuable work _Man and Nature_ gives much
information on this subject) believes that even large portions of the
African and Asiatic deserts would become covered with woods, and the
climate thereby greatly improved, were they protected from these
destructive domestic animals, which are probably not indigenous to the
country. Spain, in proportion to its extent, is very barren; Italy and
European Turkey are more woody and luxuriant; but it is perhaps in Asia
Minor, on the range of the Taurus, along the shores of the Black Sea, and
to the south of the Caucasus range, that this sub-region attains its
maximum of luxuriance in vegetation and in animal life. From the Caspian
eastward extends a region of arid plains and barren deserts, diversified by
a few more fertile valleys, in which the characteristic flora and fauna of
this portion of the Palæarctic region abounds. Further east we come to the
forests of the Hindoo Koosh, which probably form the limit of the
sub-region. Beyond these we enter on the Siberian sub-region to the north,
and on the outlying portion of the Oriental region on the south.
{201}In addition to the territories now indicated as forming part of the
Mediterranean sub-region, we must add the group of Canary Islands off the
west coast of Africa which seem to be an extension of the Atlas mountains,
and the oceanic groups of Madeira and the Azores; the latter about 1,000
miles from the continent of Europe, yet still unmistakably allied to it
both in their vegetable and animal productions. The peculiarities of the
faunas of these islands will be subsequently referred to.
It seems at first sight very extraordinary, that so large and wide a sea as
the Mediterranean should not separate distinct faunas, and this is the more
remarkable when we find how very deep the Mediterranean is, and therefore
how ancient we may well suppose it to be. Its eastern portion reaches a
depth of 2,100 fathoms or 12,600 feet, while its western basin is about
1,600 fathoms or 9,600 feet in greatest depth, and a considerable area of
both basins is more than 1,000 fathoms deep. But a further examination
shows, that a comparatively shallow sea or submerged bank incloses Malta
and Sicily, and that on the opposite coast a similar bank stretches out
from the coast of Tripoli leaving a narrow channel the greatest depth of
which is 240 fathoms. Here therefore is a broad plateau, which an elevation
of about 1,500 feet would convert into a wide extent of land connecting
Italy with Africa; while the same elevation would also connect Morocco with
Spain, leaving two extensive lakes to represent what is now the
Mediterranean Sea, and affording free communication for land animals
between Europe and North Africa. That such a state of things existed at a
comparatively recent period, is almost certain; not only because a
considerable number of identical _species_ of mammalia inhabit the opposite
shores of the Mediterranean, but also because numerous remains of three
species of elephants have been found in caves in Malta,--now a small rocky
island in which it would be impossible for such animals to live even if
they could reach it. Remains of hippopotami are also found at Gibraltar,
and many other animals of African types in Greece; all indicating means of
communication between South Europe and North Africa which no longer exist.
(See Chapter VI. pp. 113-115.)
{202}_Mammalia._--There are a few groups of Palæarctic Mammalia that are
peculiar to this sub-region. Such are, _Dama_, the fallow deer, which is
now found only in South Europe and North Africa; _Psammomys_, a peculiar
genus of Muridæ, found only in Egypt and Palestine; while _Ctenodactylus_,
a rat-like animal classed in the South American family Octodontidæ,
inhabits Tripoli. Among characteristic genera not found in other
sub-regions, are, _Dysopes_, a bat of the family Noctilionidæ;
_Macroscelides_, the elephant shrew, in North Africa; _Genetta_, the civet,
in South Europe; _Herpestes_, the ichneumon, in North Africa and (?) Spain;
_Hyæna_, in South Europe; _Gazella_, _Oryx_, _Alcephalus_, and _Addax_,
genera of antelopes in North Africa and Palestine; _Hyrax_, in Syria; and
_Hystrix_, the porcupine, in South Europe. Besides these, the camel and the
horse were perhaps once indigenous in the eastern parts of the sub-region;
and a wild sheep (_Ovis musmon_) still inhabits Sardinia, Corsica, and the
mountains of the south-east of Spain. The presence of the large feline
animals--such as the lion, the leopard, the serval, and the hunting
leopard--in North Africa, together with several other quadrupeds not found
in Europe, have been thought by some naturalists to prove, that this
district should not form part of the Palæarctic region. No doubt several
Ethiopian groups and species have entered it from the south; but the bulk
of its Mammalia still remains Palæarctic, although several of the species
have Asiatic rather than European affinities. The _Macacus innuus_ is
allied to an Asiatic rather than an African group of monkeys, and thus
denotes an Oriental affinity. Ethiopian affinity is apparently shown by the
three genera of antelopes, by _Herpestes_, and by _Macroscelides_; but our
examination of the Miocene fauna has shown that these were probably derived
from Europe originally, and do not form any part of the truly indigenous or
ancient Ethiopian fauna. Against these, however, we have the occurrence in
North Africa of such purely Palæarctic and non-Ethiopian genera as _Ursus_,
_Meles_, _Putorius_, _Sus_, _Cervus_, _Dama_, _Capra_, _Alactaga_; together
with actual European or West Asiatic species of _Canis_, _Genetta_,
_Felis_, _Putorius_, _Lutra_, many bats, _Sorex_, _Crocidura_, _Crossopus_,
_Hystrix_, {203}_Dipus_, _Lepus_, and _Mus_. It is admitted that, as
regards every other group of animals, North Africa is Palæarctic, and the
above enumeration shows that even in Mammalia, the intermixture of what are
now true Ethiopian types is altogether insignificant. It must be
remembered, also, that the lion inhabited Greece even in historic times,
while large carnivora were contemporary with man all over Central Europe.
_Birds._--So many of the European birds migrate over large portions of the
region, and so many others have a wide permanent range, that we cannot
expect to find more than a few genera, consisting of one or two species,
each, confined to a sub-region; and such appear to be, _Lusciniola_ and
_Pyrophthalma_, genera of Sylviidæ. But many are characteristic of this, as
compared with other Palæarctic sub-regions; such as, _Bradyptetus_,
_Aedon_, _Dromoloea_, and _Cercomela_, among Sylviidæ; _Crateropus_ and
_Malacocercus_, among Timaliidæ; _Telophonus_ among Laniidæ; _Certhilauda_
and _Mirafra_ among larks; _Pastor_ among starlings; _Upupa_, the hoopoe;
_Halycon_ and _Ceryle_ among kingfishers; _Turnix_ and _Caccabis_ among
Gallinæ, and the pheasant as an indigenous bird; together with _Gyps_,
_Vultur_ and _Neophron_, genera of vultures. In addition to these, almost
all our summer migrants spend their winter in some part of this favoured
land, mostly in North Africa, together with many species of Central Europe
that rarely or never visit us. It follows, that a large proportion of all
the birds of Europe and Western Asia are to be found in this sub-region, as
will be seen by referring to the list of the genera of the region.
Palestine is one of the remote portions of this region which has been well
explored by Canon Tristram, and it may be interesting to give his summary
of the range of the birds. We must bear in mind that the great depression
of the Dead Sea has a tropical climate, which accounts for the presence
here only, of such a tropical form as the sun-bird (_Nectarinea osea_).
The total number of the birds of Palestine is 322, and of these no less
than 260 are European, at once settling the question of the general
affinities of the fauna. Of the remainder eleven belong to North and East
Asia, four to the Red Sea, and {204}thirty-one to East Africa, while
twenty-seven are peculiar to Palestine. It is evident therefore that an
unusual number of East African birds have extended their range to this
congenial district, but most of these are desert species and hardly true
Ethiopians, and do not much interfere with the general Palæarctic character
of the whole assemblage. As an illustration of how wide-spread are many of
the Palæarctic forms, we may add, that seventy-nine species of land birds
and fifty-five of water birds, are common to Palestine and Britain. The
Oriental and Ethiopian genera _Pycnonotus_ and _Nectarinea_ are found here,
while _Bessornis_ and _Dromolæa_ are characteristically Ethiopian. Almost
all the other genera are Palæarctic.
Persia is another remote region generally associated with the idea of
Oriental and almost tropical forms, but which yet undoubtedly belongs to
the Palæarctic region. Mr. Blanford's recent collections in this country,
with other interesting information, is summarised in Mr. Elwes's paper on
the "Geographical Distribution of Asiatic Birds" (_Proc. Zool. Soc._ 1873,
p. 647). No less than 127 species are found also in Europe, and
thirty-seven others belong to European genera; seven are allied to birds of
Central Asia or Siberia, and fifteen to those of North-East Africa, while
only three are purely of Indian affinities. This shows a preponderance of
nearly nine-tenths of Palæarctic forms, which is fully as much as can be
expected in any country near the limits of a great region.
_Reptiles and Amphibia._--The climatal conditions being here more
favourable to these groups, and the genera being often of limited range, we
find some peculiar, and several very interesting forms. _Rhinechis_, a
genus of Colubrine snakes, is found only in South Europe; _Trogonophis_,
one of the Amphisbænians--curious snake-like lizards--is known only from
North Africa; _Psammosaurus_, belonging to the water lizards (Varanidæ) is
found in North Africa and North-West India; _Psammodromus_, a genus of
Lacertidæ, is peculiar to South Europe; _Hyalosaurus_, belonging to the
family Zonuridæ, is a lizard of especial interest, as it inhabits North
Africa while its nearest ally is the _Ophisaurus_ or "glass snake" of North
America; the family of {205}the scinks is represented by _Scincus_ found in
North Africa and Arabia. Besides these _Seps_, a genus of sand lizards
(Sepidæ) and _Agama_, a genus of Agamidæ, are abundant and characteristic.
Of Amphibia we have _Seiranota_, a genus of salamanders found only in Italy
and Dalmatia; _Chioglossa_, in Portugal, and _Geotriton_, in Italy,
belonging to the same family, are equally peculiar to the sub-region.
_Freshwater Fish._--One of the most interesting is _Tellia_, a genus of
Cyprinodontidæ found only in alpine pools in the Atlas mountains.
_Paraphoxinius_, found in South-East Europe, and _Chondrostoma_, in Europe
and Western Asia, genera of Cyprimidæ, seem almost peculiar to this
sub-region.
_Insects--Lepidoptera._--Two genera of butterflies, _Thais_ and _Doritis_,
are wholly confined to this sub-region, the former ranging over all
Southern Europe, the latter confined to Eastern Europe and Asia Minor.
_Anthocharis_ and _Zegris_ are very characteristic of it, the latter only
extending into South Russia, while _Danais_, _Charaxes_, and _Libythea_ are
tropical genera unknown in other parts of Europe.
_Coleoptera._--This sub-region is very rich in many groups of Coleoptera,
of which a few only can be noticed here. Among Carabidæ it possesses
_Procerus_ and _Procrustes_, almost exclusively, while _Brachinus_,
_Cymindis_, _Lebia_, _Graphipterus_, _Scarites_, _Chlænius_, _Calathus_,
and many others, are abundant and characteristic. Among
Lamellicorns--Copridæ, Glaphyridæ, Melolonthidæ, and Cetoniidæ abound.
Buprestidæ are plentiful, the genera _Julodis_, _Acmæodera_, _Buprestis_,
and _Sphenoptera_ being characteristic. Among Malacoderms--Cebrionidæ,
Lampyridæ, and Malachiidæ abound. The Tenebrioid Heteromera are very varied
and abundant, and give a character to the sub-region. The Mylabridæ,
Cantharidæ, and Oedemeridæ are also characteristic. Of the immense number
of Curculionidæ--_Thylacites_, _Brachycerus_, _Lixus_, and _Acalles_ may be
mentioned as among the most prominent. Of Longicorns there are few genera
especially characteristic, but perhaps _Prinobius_, _Purpuricenus_,
_Hesperophanes_, and _Parmena_ are most so. Of the remaining families, we
may mention Clythridæ, Hispidæ, and Cassididæ as being abundant.
{206}_The Mediterranean and Atlantic Islands._--The various islands of the
Mediterranean are interesting to the student of geographical distribution
as affording a few examples of local species of very restricted range, but
as a rule they present us with exactly the same forms as those of the
adjacent mainland.[6] Their peculiarities do not, therefore, properly come
within the scope of this work. The islands of the Atlantic Ocean belonging
to this sub-region are, from their isolated position and the various
problems they suggest, of much more interest, and their natural history has
been carefully studied. We shall therefore give a short account of their
peculiar features.
Of the three groups of Atlantic islands belonging to this sub-region, the
Canaries are nearest to the Continent, some of the islands being only about
fifty miles from the coast of Africa. They are, however, separated from the
mainland by a very deep channel (more than 5,000 feet), as shown on our
general map. The islands extend over a length of 300 miles; they are very
mountainous and wholly volcanic, and the celebrated peak of Teneriffe rises
to a height of more than 12,000 feet. The small Madeira group is about 400
miles from the coast of Morocco and 600 from the southern extremity of
Portugal; and there is a depth of more than 12,000 feet between it and the
continent. The Azores are nearly 1,000 miles west of Lisbon. They are quite
alone in mid-Atlantic, the most westerly islands being nearer Newfoundland
than Europe, and are surrounded by ocean depths of from 12,000 to 18,000
feet. It will be convenient to take these islands first in order.
_Azores._--Considering the remoteness of this group from every other land,
it is surprising to find as many as fifty-three species of birds inhabiting
or visiting the Azores; and still more to {207}find that they are of
Palæarctic genera and, with one exception, all of species found either in
Europe, North Africa, Madeira, or the Canaries. The exception is a
bullfinch peculiar to the islands, but closely allied to a European
species. Of land birds there are twenty-two, belonging to twenty-one
genera, all European. These genera are _Cerchneis_, _Buteo_, _Asio_,
_Strix_, _Turdus_, _Oriolus_, _Erithacus_, _Sylvia_, _Regulus_, _Saxicola_,
_Motacilla_, _Plectrophanes_, _Fringilla_, _Pyrrhula_, _Serinus_,
_Sturnus_, _Picus_, _Upupa_, _Columba_, _Caccabis_, and _Coturnix_. Besides
the bullfinch (_Pyrrhula_) other species show slight differences from their
European allies, but not such as to render them more than varieties. The
only truly indigenous mammal is a bat of a European species. Nine
butterflies inhabit the Azores; eight of them are European species, one
North American. Of beetles 212 have been collected, of which no less than
175 are European species; of the remainder, nineteen are found in the
Canaries or Madeira, three in South America, while fourteen are peculiar to
the islands.
Now these facts (for which we are indebted to Mr. Godman's _Natural History
of the Azores_) are both unexpected and exceedingly instructive. In most
other cases of remote Oceanic islands, a much larger proportion of the
fauna is endemic, or consists of peculiar species and often of peculiar
genera; as is well shown by the case of the Galapagos and Juan Fernandez,
both much nearer to a continent and both containing peculiar genera and
species of birds. Now we know that the cause and meaning of this difference
is, that in the one case the original immigration is very remote and has
never or very rarely been repeated, so that under the unchecked influence
of new conditions of life the species have become modified; in the other
case, either the original immigration has been recent, or if remote has
been so frequently repeated that the new comers have kept up the purity of
the stock, and have not allowed time for the new conditions to produce the
effect we are sure they would in time produce if not counteracted. For Mr.
Godman tells us that many of the birds are modified--instancing the
gold-crested wren, blackcap, and rock dove--and he adds, that the
{208}modification all tends in one direction--to produce a more sombre
plumage, a greater strength of feet and legs, and a more robust bill. We
further find, that four of the land-birds, including the oriole,
snow-bunting, and hoopoe, are not resident birds, but straggle accidentally
to the islands by stress of weather; and we are told that every year some
fresh birds are seen after violent storms. Add to this the fact, that the
number of species diminishes in the group as we go from east to west, and
that the islands are subject to fierce and frequent storms blowing from
every point of the compass,--and we have all the facts requisite to enable
us to understand how this remote archipelago has become stocked with animal
life without ever probably being much nearer to Europe than it is now. For
the islands are all volcanic, the only stratified rock that occurs being
believed to be of Miocene date.
_Madeira and the Canaries._--Coming next to Madeira, we find the number of
genera of land birds has increased to twenty-eight, of which seventeen are
identical with those of the Azores. Some of the commonest European
birds--swallows, larks, sparrows, linnets, goldfinches, ravens, and
partridges, are among the additions. A gold-crested warbler, _Regulus
Maderensis_, and a pigeon, _Columba Trocaz_, are peculiar to Madeira.
In the Canaries we find that the birds have again very much increased,
there being more than fifty genera of land birds; but the additions are
wholly European in character, and almost all common European species. We
find a few more peculiar species (five), while some others, including the
wild canary, are common to all the Atlantic Islands or to the Canaries and
Madeira. Here, too, the only indigenous mammalia are two European species
of bats.
_Land Shells._--The land shells of Madeira offer us an instructive contrast
to the birds of the Atlantic Islands. About fifty-six species have been
found in Madeira, and forty-two in the small adjacent island of Porto
Santo, but only twelve are common to both, and all or almost all are
distinct from their nearest allies in Europe and North Africa. Great
numbers of fossil shells are also found in deposits of the Newer Pliocene
period; and {209}although these comprise many fresh species, the two faunas
and that of the continent still remain almost as distinct from each other
as before. It has been already stated (p. 31) that the means by which land
mollusca have been carried across arms of the sea are unknown, although
several modes may be suggested; but it is evidently a rare event, requiring
some concurrence of favourable conditions not always present. The diversity
and specialization of the forms of these animals is, therefore, easily
explained by the fact, that, once introduced they have been left to
multiply under the influence of a variety of local conditions, which
inevitably lead, in the course of ages, to the formation of new varieties
and new species.
_Coleoptera._--The beetles of Madeira and the Canaries have been so
carefully collected and examined by Mr. T. V. Wollaston, and those of the
Azores described and compared by Mr. Crotch, and they illustrate so many
curious points in geographical distribution, that it is necessary to give
some account of them. No less than 1,480 species of beetles have been
obtained from the Canaries and Madeira, only 360 of which are European, the
remainder being peculiar to the islands. The Canaries are inhabited by a
little over 1,000 species, Madeira by about 700, while 240 are common to
both; but it is believed that many of these have been introduced by man. In
the Azores, 212 species have been obtained, of which 175 are European;
showing, as in the birds, as closer resemblance to the European fauna than
in the other islands which, although nearer to the continent, offer more
shelter and are situated in a less tempestuous zone. Of the non-European
species in the Azores, 19 are found also in the other groups of islands, 14
are peculiar, while 3 are American. Of the European species, 132 are found
also in the other Atlantic islands, while 43 have reached the Azores only.
This is interesting as showing to how great an extent the same insects
reach all the islands, notwithstanding the difference of latitude and
position; and it becomes of great theoretical importance, when we find how
many extensive families and genera are altogether absent.
The Madeira group has been more thoroughly explored than {210}any other,
and its comparatively remote situation, combined with its luxuriant
vegetation, have been favourable to the development and increase of the
peculiar forms which characterize all the Atlantic islands in a more or
less marked degree. A consideration of some of its peculiarities will,
therefore, best serve to show the bearing of the facts presented by the
insect fauna of the Atlantic islands, on the general laws of distribution.
The 711 species of beetles now known from the Madeira group, belong to 236
genera; and no less than 44 of these genera are not European but are
peculiar to the Atlantic islands. Most of them are, however, closely allied
to European genera, of which they are evidently modifications. A most
curious general feature presented by the Madeiran beetles, is the total
absence of many whole families and large genera abundant in South Europe.
Such are the Cicindelidæ, or tiger beetles; the Melolonthidæ, or chafers;
the Cetoniidæ, or rose-chafers; the Eumolpidæ and Galerucidæ, large
families of Phytophagous, or leaf-eating beetles; and also the extensive
groups of Elateridæ and Buprestidæ, which are each represented by but one
minute species. Of extensive genera abundant in South Europe, but wholly
absent in Madeira, are _Carabus_, _Rhizotrogus_, _Lampyris_, and other
genera of Malacoderms; _Otiorhynchus_, _Brachycerus_, and 20 other genera
of Curculionidæ, comprising more than 300 South European and North African
species; _Pimelia_, _Tentyra_, _Blaps_, and 18 other genera of Heteromera,
comprising about 550 species in South Europe and North Africa; and
_Timarcha_, containing 44 South European and North African species.
Another most remarkable feature of the Madeiran Coleoptera is the unusual
prevalence of apterous or wingless insects. This is especially the case
with groups which are confined to the Atlantic islands, many of which
consist wholly of wingless species; but it also affects the others, no less
than twenty-two genera which are usually or sometimes winged in Europe,
having only wingless species in Madeira; and even the same species which is
winged in Europe becomes, in at least three cases, wingless in Madeira,
without any other perceptible change having taken place. But there is
another most curious fact noticed by {211}Mr. Wollaston; that those species
which possess wings in Madeira, often have them rather larger than their
allies in Europe. These two facts were connected by Mr. Darwin, who
suggested that flying insects are much more exposed to be blown out to sea
and lost, than those which do not fly (and Mr. Wollaston had himself
supposed that the "stormy atmosphere" of Madeira had something to do with
the matter); so that the most frequent fliers would be continually weeded
out, while the more sluggish individuals, who either could not or would not
fly, remained to continue the race; and this process going on from
generation to generation, would, on the well-ascertained principles of
selection and abortion by disuse, in time lead to the entire loss of wings
by those insects to whom wings were _not a necessity_. But those whose
wings were essential to their existence would be acted upon in another way.
All these must fly to obtain their food or provide for their offspring, and
those that flew best would be best able to battle with the storms, and keep
themselves safe, and thus those with the longest and most powerful wings
would be preserved. If however all the individuals of the species were too
weak on the wing to resist the storms, they would soon become extinct.[7]
Now this explanation of the facts is not only simple and probable in
itself, but it also serves to explain in a remarkable manner some of the
peculiarities and deficiencies of the Madeiran insect fauna, in harmony
with the view (supported by the distribution of the birds and land shells,
and in particular by the immigrant birds and insects of the Azores) that
all the insects have been derived from the continent or from other islands,
by {212}immigration across the ocean, in various ways and during a long
period. These deficiencies are, on the other hand, quite inconsistent with
the theory (still held by some entomologists) that a land communication is
absolutely necessary to account for the origin of the Madeiran fauna.
First, then, we can understand how the tiger-beetles (Cicindelidæ) are
absent; since they are insects which have a short weak flight, but yet to
whom flight is necessary. If a few had been blown over to Madeira, they
would soon have become exterminated. The same thing applies to the
Melolonthidæ, Cetoniidæ, Eumolpidæ, and Galerucidæ,--all flower and
foliage-haunting insects, yet bulky and of comparatively feeble powers of
flight. Again, all the large genera abundant in South Europe, which have
been mentioned above as absent from Madeira, are wholly apterous (or
without wings), and thus their absence is a most significant fact; for it
proves that in the case of all insects of moderate size, flight was
essential to their reaching the island, which could not have been the case
had there been a land connection. There are, however, one or two curious
exceptions to the absence of these wholly apterous European genera in
Madeira, and as in each case the reason of their being exceptions can be
pointed out, they are eminently exceptions that prove the rule. Two of the
apterous species common to Europe and Madeira are found always in ants'
nests; and as ants, when winged, fly in great swarms and are carried by the
wind to great distances, they may have conveyed the minute eggs of these
very small beetles. Two European species of _Blaps_ occur in Madeira, but
these are house beetles, and are admitted to have been introduced by man.
There are also three species of _Meloe_, of which two are European and one
peculiar. These are large, sluggish, wingless insects, but they have a most
extraordinary and exceptional metamorphosis, the larvae in the first state
being minute active insects parasitic on bees, and thus easily conveyed
across the ocean. This case is most suggestive, as it accounts for what
would be otherwise a difficult anomaly. Another case, not quite so easily
explained, is that of the genus _Acalles_, which is very abundant in all
the Atlantic {213}islands and also occurs in South Europe, but is always
apterous. It is however closely allied to another genus, _Cryptorhynchus_,
which is apterous in some species, winged in others. We may therefore well
suppose that the ancestors of _Acalles_ were once in the same condition,
and that some of the winged forms reached Madeira, the genus having since
become wholly apterous.
We may look at this curious subject in another way. The Coleoptera of
Madeira may be divided into those which are found also in Europe or the
other islands, and those which are peculiar to it. On the theory of
introduction by accidental immigration across the sea, the latter must be
the more ancient, since they have had time to become modified; while the
former are comparatively recent, and their introduction may be supposed to
be now going on. The peculiar influence of Madeira in aborting the wings
should, therefore, have acted on the ancient and changed forms much more
powerfully than on the recent and unchanged forms. On carefully comparing
the two sets of insects (omitting those which have almost certainly been
introduced by man) we find, that out of 263 species which have a wide
range, only 14 are apterous; while the other class, consisting of 393
species, has no less than 178 apterous; or about 5 per cent in the one
case, and 45 per cent in the other.[8] On the theory of a land connection
as the main agent in introducing the fauna, both groups must have been
introduced at or about the same time, and why one set should have lost
their wings and the other not, is quite inexplicable.
Taking all these singular facts, in connection with the total absence of
all truly indigenous terrestrial mammalia and reptiles from these
islands--even from the extensive group of the Canaries so comparatively
near to the continent, we are forced to reject the theory of a land
connection as quite untenable; and this view becomes almost demonstrated by
the case of the Azores, which being so much further off, and surrounded by
such a vast expanse of deep ocean, could only have been {214}connected with
Europe at a far remoter epoch, and ought therefore to exhibit to us a fauna
composed almost entirely of peculiar forms both of birds and insects. Yet,
so far from this being the case, the facts are exactly the reverse. Far
more of the birds and insects are identical with those of Europe than in
the other islands, and this difference is clearly traced to the more
tempestuous atmosphere, which is shown to be even now annually bringing
fresh immigrants (both birds and insects) to its shores. We here see nature
actually at work; and if the case of Madeira rendered her mode of action
probable, that of the Azores may be said to demonstrate it.
Mr. Wollaston has objected to this view that "storms and hurricanes" are
somewhat rare in the latitude of Madeira and the Canaries; but this little
affects the question, since the _time_ allowed for such operations is so
ample. If but one very violent storm happened in a century, and ten such
storms recurred before a single species of insect was introduced into
Madeira, that would be more than sufficient to people it, as we now find
it, with a varied fauna. But he also adds the important information that
the ordinary winds blow almost uninterruptedly from the north-east, so that
there would be always a chance of a little stronger wind than usual
bringing insect, or larva, or egg, attached to leaves or twigs. Neither Mr.
Wollaston, Mr. Crotch, Mr. A. Murray, nor any other naturalist who upholds
the land-connection theory, has attempted to account for the fact of the
absence of so many extensive groups of insects that ought to be present, as
well as of all small mammalia and reptiles.
_Cape Verd Islands._--There is yet another group of Atlantic islands which
is very little known, and which is usually considered to be altogether
African--the Cape Verd Islands, situated between 300 and 400 miles west of
Senegal, and a little to the south of the termination of the Sahara. The
evidence that we possess as to the productions of these islands, shows
that, like the preceding groups, they are truly oceanic, and have probably
derived their fauna from the desert and the Canaries to the north-east of
them rather than from the fertile and more truly {215}Ethiopian districts
of Senegal and Gambia to the east. There is a mingling of the two faunas,
but the preponderance seems to be undoubtedly with the Palæarctic rather
than with the Ethiopian. I owe to Mr. R. B. Sharpe of the British Museum, a
MS. list of the birds of these islands, twenty-three species in all. Of
these eight are of wide distribution and may be neglected. Seven are
undoubted Palæarctic species, viz.:--_Milvus ictinus_, _Sylvia
atricapilla_, _S. conspicillata_, _Corvus corone_, _Passer salicarius_,
_Certhilauda desertorum_, _Columba livia_. Three are peculiar species, but
of Palæarctic genera and affinities, viz.:--_Calamoherpe brevipennis_,
_Ammomanes cinctura_, and _Passer jagoensis_. Against this we have to set
two West African species, _Estrilda cinerea_ and _Numida meleagris_, both
of which were probably introduced by man; and three which are of Ethiopian
genera and affinities, viz.:--_Halcyon erythrorhyncha_, closely allied to
_H. semicærulea_ of Arabia and North-east Africa, and therefore almost
Palæarctic; _Accipiter melanoleucus_; and _Pyrrhulauda nigriceps_, an
Ethiopian form; but the same species occurs in the Canaries.
The Coleoptera of these islands have been also collected by Mr. Wollaston,
and he finds that they have generally the same European character as those
of the Canaries and Madeira, several of the peculiar Atlantic genera, such
as _Acalles_ and _Hegeter_, occurring, while others are represented by new
but closely allied genera. Out of 275 species 91 were found also in the
Canaries and 81 in the Madeiran group; a wonderful amount of similarity
when we consider the distance and isolation of these islands and their
great diversity of climate and vegetation.
This connection of the four groups of Atlantic islands now referred to,
receives further support from the occurrence of land-shells of the subgenus
_Leptaxis_ in all the groups, as well as in Majorca; and by another
subgenus, _Hemicycla_, being common to the Canaries and Cape Verd islands.
Combining these several classes of facts, we seem justified in extending
the Mediterranean sub-region to include the Cape Verd Islands.
{216}_III.--The Siberian Sub-region, or Northern Asia._
This large and comparatively little-known subdivision of the Palæarctic
region, extends from the Caspian Sea to Kamschatka and Behring's Straits, a
distance of about 4,000 miles; and from the shores of the Arctic Ocean to
the high Himalayas of Sikhim in North Latitude 29°, on the same parallel as
Delhi. To the east of the Caspian Sea and the Ural Mountains is a great
extent of lowland which is continued round the northern coast, becoming
narrower as it approaches the East Cape. Beyond this, in a general E.N.E.
direction, rise hills and uplands, soon becoming lofty mountains, which
extend in an unbroken line from the Hindu Koosh, through the Thian Shan,
Altai and Yablonoi Mountains, to the Stanovoi range in the north-eastern
extremity of Asia. South of this region is a great central basin, which is
almost wholly desert; beyond which again is the vast plateau of Thibet,
with the Kuenlun, Karakorum, and Himalayan snow-capped ranges, forming the
most extensive elevated district on the globe.
The superficial aspects of this vast territory, as determined by its
vegetable covering, are very striking and well contrasted. A broad tract on
the northern coast, varying from 150 to 300 and even 500 miles wide, is
occupied by the Tundras or barrens, where nothing grows but mosses and the
dwarfest Arctic plants, and where the ground is permanently frozen to a
great depth. This tract has its greatest southern extension between the
rivers Obi and Yenesi, where it reaches the parallel of 60° north latitude.
Next to this comes a vast extent of northern forests, mostly of conifers in
the more northern and lofty situations, while deciduous trees preponderate
in the southern portions and in the more sheltered valleys. The greatest
extension of this forest region is north of Lake Baikal, where it is more
than 1,200 miles wide. These forests extend along the mountain ranges to
join those of the Hindu Koosh. South of the forests the remainder of the
sub-region consists of open pasture-lands and vast intervening deserts, of
which the Gobi, and those of Turkestan between the Aral and Balkash lakes,
are the most {217}extensive. The former is nearly 1,000 miles long, with a
width of from 200 to 350 miles, and is almost as complete a desert as the
Sahara.
With very few exceptions, this vast territory is exposed to an extreme
climate, inimical to animal life. All the lower parts being situated to the
north, have an excessively cold winter, so that the limit of constantly
frozen ground descends below the parallel of 60° north latitude. To the
south, the land is greatly elevated, and the climate extremely dry. In
summer the heat is excessive, while the winter is almost as severe as
further north. The whole country, too, is subject to violent storms, both
in summer and winter; and the rich vegetation that clothes the steppes in
spring, is soon parched up and replaced by dusty plains. Under these
adverse influences we cannot expect animal life to be so abundant as in
those sub-regions subject to more favourable physical conditions; yet the
country is so extensive and so varied, that it does actually, as we shall
see, possess a very considerable and interesting fauna.
_Mammalia._--Four genera seem to be absolutely confined to this sub-region,
_Nectogale_, a peculiar form of the mole family (Talpidæ); _Poephagus_, the
yak, or hairy bison of Thibet; with _Procapra_ and _Pantholops_, Thibetan
antelopes. Some others more especially belong here, although they just
enter Europe, as _Saiga_, the Tartarian antelope; _Sminthus_, a desert
rat;, and _Ellobius_, a burrowing mole-rat; while _Myospalax_, a curious
rodent allied to the voles, is found only in the Altai mountains and North
China; and _Moschus_, the musk-deer, is almost confined to this sub-region.
Among the characteristic animals of the extreme north, are _Mustela_, and
_Martes_, including the ermine and sable; _Gulo_, the glutton; _Tarandus_,
the reindeer; _Myodes_, the lemming; with the lynx, arctic fox, and polar
bear; and here, in the Post-pliocene epoch, ranged the hairy rhinoceros and
Siberian mammoth, whose entire bodies still remain preserved in the
ice-cliffs near the mouths of the great rivers. Farther south, species of
wild cat, bear, wolf, deer, and pika (_Lagomys_) abound; while in the
mountains we find wild goats and sheep of several species, and in the
plains and deserts wild horses {218}and asses, gazelles, two species of
antelopes, flying squirrels (_Pteromys_), ground squirrels (_Tamias_),
marmots, of the genus _Spermophilus_, with camels and dromedaries, probably
natives of the south-western part of this sub-region. The most abundant and
conspicuous of the mammalia are the great herds of reindeer in the north,
the wolves of the steppes, with the wild horses, goats, sheep, and
antelopes of the plateaus and mountains.
Among the curiosities of this sub-region we must notice the seal, found in
the inland and freshwater lake Baikal, at an elevation of about 2,000 feet
above the sea. It is a species of _Callocephalus_, closely allied to, if
not identical with, one inhabiting northern seas as well as the Caspian and
Lake Aral. This would indicate that almost all northern Asia was depressed
beneath the sea very recently; and Mr. Belt's view, of the ice during the
glacial epoch having dammed up the rivers and converted much of Siberia
into a vast freshwater or brackish lake, perhaps offers the best solution
of the difficulty.[9]
_Plate II.--Characteristic Mammalia of Western Tartary._--Several of the
most remarkable animals of the Palæarctic region inhabit Western Tartary,
and are common to the European and Siberian sub-regions. We therefore
choose this district for one of our illustrative plates. The large animals
in the centre are the remarkable saiga antelopes (_Saiga Tartarica_),
distinguished from all others by a large and fleshy proboscis-like nose,
which gives them a singular appearance. They differ so much from all other
antelopes that they have been formed into a distinct family by some
naturalists, but are here referred to the great family Bovidæ. They inhabit
the open plains from Poland to the Irtish River. On the left is the
mole-rat, or sand-rat (_Spalax murinus_). This animal burrows under ground
like a mole, feeding on bulbous roots. It inhabits the same country as the
saiga, but extends farther south in Europe. On the right is a still more
curious animal, the desman (_Myogale Muscovitica_), a long-snouted
water-mole. This creature is fifteen inches long, including the tail; it
burrows in the banks of streams, feeding on insects, {219}worms, and
leeches; it swims well, and remains long under water, raising the tip of
the snout, where the nostrils are situated, to the surface when it wants to
breathe. It is thus well concealed; and this may be one use of the
development of the long snout, as well as serving to follow worms into
their holes in the soft earth. This species is confined to the rivers Volga
and Don in Southern Russia, and the only other species known inhabits some
of the valleys on the north side of the Pyrenees. In the distance are
wolves, a characteristic feature of these wastes.
Plate II.
[Illustration]
CHARACTERISTIC MAMMALIA OF WESTERN TARTARY.
_Birds._--But few genera of birds are absolutely restricted to this
sub-region. _Podoces_, a curious form of starling, is the most decidedly
so; _Mycerobas_ and _Pyrrhospiza_ are genera of finches confined to Thibet
and the snowy Himalayas; _Leucosticte_, another genus of finches, is
confined to the eastern half of the sub-region and North America;
_Tetraogallus_, a large kind of partridge, ranges west to the Caucasus;
_Syrrhaptes_, a form of sand-grouse, and _Lerwa_ (snow-partridge), are
almost confined here, only extending into the next sub-region; as do
_Grandala_, and _Calliope_, genera of warblers, _Uragus_, a finch allied to
the North American cardinals, and _Crossoptilon_, a remarkable group of
pheasants.
Almost all the genera of central and northern Europe are found here, and
give quite a European character to the ornithology, though a considerable
number of the species are different. There are a few Oriental forms, such
as _Abrornis_ and _Larvivora_ (warblers); with _Ceriornis_ and _Ithaginis_,
genera of pheasants, which reach the snow-line in the Himalayas and thus
just enter this sub-region, but as they do not penetrate farther north,
they hardly serve to modify the exclusively Palæarctic character of its
ornithology.
According to Middendorf, the extreme northern Asiatic birds are the Alpine
ptarmigan (_Lagopus mutus_); the snow-bunting (_Plectrophanes nivalis_);
the raven, the gyrfalcon and the snowy-owl. Those which are characteristic
of the barren "tundras," but which do not range so far north as the
preceding are,--the willow-grouse (_Lagopus albus_); the Lapland-bunting
(_Plectrophanes {220}lapponica_); the shore-lark (_Otocorys alpestris_);
the sand-martin (_Cotyle riparia_), and the sea-eagle (_Haliæetus
albicilla_).
Those which are more characteristic of the northern forests, and which do
not pass beyond them, are--the linnet; two crossbills (_Loxia Leucoptera_
and _L. Curvirostra_); the pine grosbeak (_Pinicola enucleator_); the
waxwing; the common magpie; the common swallow; the peregrine falcon; the
rough-legged buzzard; and three species of owls.
Fully one-half of the land-birds of Siberia are identical with those of
Europe, the remainder being mostly representative species peculiar to
Northern Asia, with a few stragglers and immigrants from China and Japan or
the Himalayas. A much larger proportion of the wading and aquatic families
are European or Arctic, these groups having always a wider range than land
birds.
_Reptiles and Amphibia._--From the nature of the country and climate these
are comparatively few, but in the more temperate districts snakes and
lizards seem to be not uncommon. _Halys_, a genus of Crotaline snakes, and
_Phrynocephalus_, lizards of the family Agamidæ, are characteristic of
these parts. _Simotes_, a snake of the family Oligodontidæ, reaches an
elevation of 16,000 feet in the Himalayas, and therefore enters this
sub-region.
_Insects._--_Mesapia_ and _Hypermnestra_, genera of Papilionidæ, are
butterflies peculiar to this sub-region; and _Parnassius_ is as
characteristic as it is of our European mountains. Carabidæ are also
abundant, as will be seen by referring to the Chapter on the Distribution
of Insects in the succeeding part of this work. The insects, on the whole,
have a strictly European character, although a large proportion of the
species are peculiar, and several new genera appear.
_IV.--Japan and North China, or the Manchurian Sub-region._
This is an interesting and very productive district, corresponding in the
east to the Mediterranean sub-region in the west, or rather perhaps to all
western temperate Europe. Its limits are not very well defined, but it
probably includes all Japan; the Corea and Manchuria to the Amour river and
to the lower {221}slopes of the Khingan and Peling mountains; and China to
the Nanlin mountains south of the Yang-tse-kiang. On the coast of China the
dividing line between it and the Oriental region seems to be somewhere
about Foo-chow, but as there is here no natural barrier, a great
intermingling of northern and southern forms takes place.
Japan is volcanic and mountainous, with a fine climate and a most luxuriant
and varied vegetation. Manchuria is hilly, with a high range of mountains
on the coast, and some desert tracts in the interior, but fairly wooded in
many parts. Much of northern China is a vast alluvial plain, backed by
hills and mountains with belts of forest, above which are the dry and
barren uplands of Mongolia. We have a tolerable knowledge of China, of
Japan, and of the Amoor valley, but very little of Corea and Manchuria. The
recent researches of Père David in Moupin, in east Thibet, said to be
between 31° and 32° north latitude, show, that the fauna of the Oriental
region here advances northward along the flanks of the Yun-ling mountains
(a continuation of the Himalayas); since he found at different altitudes
representatives of the Indo-Chinese, Manchurian, and Siberian faunas. On
the higher slopes of the Himalayas, there must be a narrow strip from about
8,000 to 11,000 feet elevation intervening between the tropical fauna of
the Indo-Chinese sub-region and the almost arctic fauna of Thibet; and the
animals of this zone will for the most part belong to the fauna of
temperate China and Manchuria, except in the extreme west towards Cashmere,
where the Mediterranean fauna will in like manner intervene. On a map of
sufficiently large scale, therefore, it would be necessary to extend our
present sub-region westward along the Himalayas, in a narrow strip just
below the upper limits of forests. It is evident that the large number of
Fringillidæ, Corvidæ, Troglodytidæ, and Paridæ, often of south Palæarctic
forms, that abound in the higher Himalayas, are somewhat out of place as
members of the Oriental fauna, and are equally so in that of Thibet and
Siberia; but they form a natural portion of that of North China on the one
side, or of South Europe on the other.
{222}_Mammalia._--This sub-region contains a number of peculiar and very
interesting forms, most of which have been recently discovered by Père
David in North and West China and East Thibet. The following are the
peculiar genera:--_Rhinopithecus_, a sub-genus of monkeys, here classed
under _Semnopithecus_; _Anurosorex_, _Scaptochirus_, _Uropsilus_ and
_Scaptonyx_, new forms of Talpidæ or moles; _Æluropus_ (Æluridæ);
_Nyctereutes_ (Canidæ); _Lutronectes_ (Mustelidæ); _Cricetulus_ (Muridæ);
_Hydropotes_, _Moschus_, and _Elaphodus_ (Cervidæ). The _Rhinopithecus_
appears to be a permanent inhabitant of the highest forests of Moupin, in a
cold climate. It has a very thick fur, as has also a new species of
_Macacus_ found in the same district. North China and East Thibet seem to
be very rich in Insectivora. _Scaptochirus_ is like a mole; _Uropsilus_
between the Japanese _Urotrichus_ and _Sorex_; _Scaptonyx_ between
_Urotrichus_ and _Talpa_. _Æluropus_ seems to be the most remarkable mammal
discovered by Père David. It is allied to the singular panda (_Ælurus
fulgens_) of Nepal, but is as large as a bear, the body wholly white, with
the feet, ears, and tip of the tail black. It inhabits the highest forests,
and is therefore a true Palæarctic animal, as most likely is the _Ælurus_.
_Nyctereutes_, a curious racoon-like dog, ranges from Canton to North
China, the Amoor and Japan, and therefore seems to come best in this
sub-region; _Hydropotes_ and _Lophotragus_ are small hornless deer confined
to North China; _Elaphodus_, from East Thibet, is another peculiar form of
deer; while the musk deer (_Moschus_) is confined to this sub-region and
the last. Besides the above, the following Palæarctic genera were found by
Père David in this sub-region: _Macacus_: five genera or sub-genera of bats
(_Vespertilio_, _Vesperus_, _Vesperugo_, _Rhinolophus_, and _Murina_);
_Erinaceus_, _Nectogale_, _Talpa_, _Crocidura_ and _Sorex_, among
Insectivora; _Mustela_, _Putorius_, _Martes_, _Lutra_, _Viverra_, _Meles_,
_Ælurus_, _Ursus_, _Felis_, and _Canis_, among _Carnivora_; _Hystrix_,
_Arctomys_, _Myospalax_, _Spermophilus_, _Gerbillus_, _Dipus_, _Lagomys_,
_Lepus_, _Sciurus_, _Pteromys_, _Arvicola_, and _Mus_, among Rodentia;
_Budorcas_, _Nemorhedus_, _Antilope_, _Ovis_, _Moschus_, _Cervulus_ and
_Cervus_ among Ruminants; and the wide-spread _Sus_ or wild boar. The
following Oriental genera are also {223}included in Père David's list, but
no doubt occur only in the lowlands and warm valleys, and can hardly be
considered to belong to the Palæarctic region: _Paguma_, _Helictis_,
_Arctonyx_, _Rhizomys_, _Manis_. The _Rhizomys_ from Moupin is a peculiar
species of this tropical genus, but all the others inhabit Southern China.
A few additional forms occur in Japan: _Urotrichus_, a peculiar Mole, which
is found also in north-west America; _Enhydra_, the sea otter of
California; and the dormouse (_Myoxus_). Japan also possesses peculiar
species of _Macacus_, _Talpa_, _Meles_, _Canis_, and _Sciuropterus_.
It will be seen that this sub-region is remarkably rich in Insectivora, of
which it possesses ten genera; and that it has also several peculiar forms
of Carnivora, Rodentia, and Ruminants.
_Birds._--To give an accurate idea of the ornithology of this sub-region is
very difficult, both on account of its extreme richness and the
impossibility of defining the limits between it and the Oriental region. A
considerable number of genera which are well developed in the high
Himalayas, and some which are peculiar to that district, have hitherto
always been classed as Indian, and therefore Oriental groups; but they more
properly belong to this sub-region. Many of them frequent the highest
forests, or descend into the Himalayan temperate zone only in winter; and
others are so intimately connected with Palæarctic species, that they can
only be considered as stragglers into the border land of the Oriental
region. On these principles we consider the following genera to be confined
to this sub-region:--
_Grandala_, _Nemura_ (Sylviidæ); _Pterorhinus_ (Timaliidæ); _Cholornis_,
_Conostoma_, _Heteromorpha_ (Panuridæ); _Cyanoptila_ (Muscicapidæ);
_Eophona_ (Fringillidæ); _Dendrotreron_ (Columbidæ); _Lophophorus_,
_Tetraophasis_, _Crossoptilon_, _Pucrasia_, _Thaumalea_, and _Ithaginis_
(Phasianidæ). This may be called the sub-region of Pheasants; for the above
six genera, comprising sixteen species of the most magnificent birds in the
world, are all confined to the temperate or cold mountainous regions of the
Himalayas, Thibet, and China; and in addition we have {224}most of the
species of tragopan (_Ceriornis_), and some of the true pheasants
(_Phasianus_).
The most abundant and characteristic of the smaller birds are warblers,
tits, and finches, of Palæarctic types; but there are also a considerable
number of Oriental forms which penetrate far into the country, and mingling
with the northern birds give a character to the Ornithology of this
sub-region very different from that of the Mediterranean district at the
western end of the region. Leaving out a large number of wide-ranging
groups, this mixture of types may be best exhibited by giving lists of the
more striking Palæarctic and Oriental genera which are here found
intermingled.
PALÆARCTIC GENERA.
SYLVIIDÆ.
Erithacus.
Ruticilla.
Locustella.
Cyanecula.
Sylvia.
Potamodus.
Reguloides.
Regulus.
Accentor.
CINCLIDÆ.
Cinclus.
TROGLODYTIDÆ.
Troglodytidæ.
CERTHIIDÆ.
Certhia.
Sitta.
Tichodroma.
PARIDÆ.
Parus.
Lophophanes.
Acredula.
CORVIDÆ.
Fregilus.
Nucifraga.
Pica.
Cyanopica.
Garrulus.
AMPELIDÆ.
Ampelis.
FRINGILLIDÆ.
Fringilla.
Chrysomitris.
Chlorospiza.
Passer.
Coccothraustes.
Pyrrhula.
Carpodacus.
Uragus.
Loxia.
Linota.
Emberiza.
STURNIDÆ.
Sturnus.
ALAUDIDÆ.
Otocorys.
PICIDÆ.
Picoides.
Picus.
Hyopicus.
Dryocopus.
YUNGIDÆ.
Yunx.
PTEROCLIDÆ.
Syrrhaptes.
TETRAONIDÆ.
Tetrao.
Tetraogallus.
Lerwa.
Lagopus.
VULTURIDÆ.
Gypaëtus.
Vultur.
FALCONIDÆ.
Archibuteo.
ORIENTAL GENERA.
SYLVIIDÆ.
Suya.
Calliope.
Larvivora.
Tribura.
Horites.
Abrornis.
Copsychus.
TURDIDÆ.
Oreocincla.
TIMALIIDÆ.
Alcippe.
Timalia.
Pterocyclus.
Garrulax.
Trochalopteron.
Pomatorhinus. {225}
Suthora.
PANURIDÆ.
Paradoxornis.
CINCLIDÆ.
Enicurus.
Myiophonus.
TROGLODYTIDÆ.
Pnoepyga.
LIOTRICHIDÆ.
Liothrix.
Yuhina.
Pteruthius.
PYCNONOTIDÆ.
Microscelis.
Pycnonotus.
Hypsipetes.
CAMPEPHAGIDÆ.
Pericrocotus.
DICRURIDÆ.
Dicrurus.
Chibia.
Buchanga.
MUSCICAPIDÆ.
Xanthopygia.
Niltava.
Tchitrea.
CORVIDÆ.
Urocissa.
NECTARINEIDÆ.
Æthopyga.
MOTACILLIDÆ.
Nemoricola.
DICÆIDÆ.
Zosterops.
FRINGILLIDÆ.
Melophus.
Pyrgilauda.
PLOCEIDÆ.
Munia.
STURNIDÆ.
Acridotheres.
Sturnia.
PITTIDÆ.
Pitta.
PICIDÆ.
Vivia.
Yungipicus.
Gecinus.
CORACIIDÆ.
Eurystomus.
ALCEDINIDÆ.
Halcyon.
Ceryle.
UPUPIDÆ.
Upupa.
PSITTACIDÆ.
Palæornis.
COLUMBIDÆ.
Treron.
Ianthænas.
Macropygia.
PHASIANIDÆ.
Phasianus.
Ceriornis.
STRIGIDÆ.
Scops.
In the above lists there are rather more Oriental than Palæarctic genera;
but it must be remembered that most of the former are summer migrants only,
or stragglers just entering the sub-region; whereas the great majority of
the latter are permanent residents, and a large proportion of them range
over the greater part of the Manchurian district. Many of those in the
Oriental column should perhaps be omitted, as we have no exact
determination of their range, and the limits of the regions are very
uncertain. It must be remembered, too, that the Palæarctic genera of
Sylviidæ, Paridæ, and Fringillidæ, are often represented by numerous
species, whereas the corresponding Oriental genera have for the most part
only single species; and we shall then find that, except towards the
borders of the Oriental region the Palæarctic element is strongly
predominant. Four of the more especially Oriental groups are confined to
Japan, the southern {226}extremity of which should perhaps come in the
Oriental region. The great richness of this sub-region compared with that
of Siberia is well shown by the fact, that a list of all the known
land-birds of East Siberia, including Dahuria and the comparatively fertile
Amoor Valley, contains only 190 species; whereas Père David's catalogue of
the birds of Northern China with adjacent parts of East Thibet and Mongolia
(a very much smaller area) contains for the same families 366 species. Of
the Siberian birds more than 50 per cent, are European species, while those
of the Manchurian sub-region comprise about half that proportion of
land-birds which are identical with those of Europe.
Japan is no doubt very imperfectly known, as only 134 land-birds are
recorded from it. Of these twenty-two are peculiar species, a number that
would probably be diminished were the Corea to be explored. Of the genera,
only nine are Indo-Malayan, while forty-three are Palæarctic.
_Plate III.--Scene on the Borders of North-West China and Mongolia with
Characteristic Mammalia and Birds._--The mountainous districts of Northern
China, with the adjacent portions of Thibet and Mongolia, are the
head-quarters of the pheasant tribe, many of the most beautiful and
remarkable species being found there only. In the north-western provinces
of China and the southern parts of Mongolia may be found the species
figured. That in the foreground is the superb golden pheasant (_Thaumalea
picta_), a bird that can hardly be surpassed for splendour of plumage by
any denizen of the tropics. The large bird perched above is the eared
pheasant (_Crossoptilon auritum_), a species of comparatively sober plumage
but of remarkable and elegant form. In the middle distance is Pallas's sand
grouse (_Syrrhaptes paradoxus_), a curious bird, whose native country seems
to be the high plains of Northern Asia, but which often abounds near Pekin,
and in 1863 astonished European ornithologists by appearing in considerable
numbers in Central and Western Europe, in every part of Great Britain, and
even in Ireland.
Plate III.
[Illustration]
CHARACTERISTIC ANIMALS OF NORTH CHINA.
The quadruped figured is the curious racoon dog (_Nyctereutes
procyonoides_), {227}an animal confined to North China, Japan, and the
Amoor Valley, and having no close allies in any other part of the globe. In
the distance are some deer, a group of animals very abundant and varied in
this part of the Palæarctic region.
_Reptiles and Amphibia._--Reptiles are scarce in North China, only four or
five species of snakes, a lizard and one of the Geckotidæ occurring in the
country round Pekin. The genus _Halys_ is the most characteristic form of
snake, while _Callophis_, an oriental genus, extends to Japan. Among
lizards, _Plestiodon_, _Maybouya_, _Tachydromus_, and _Gecko_ reach Japan,
the two latter being very characteristic of the Oriental region.
Amphibia are more abundant and interesting; _Hynobius_, _Onychodactylus_,
and _Sieboldtia_ (Salamandridæ) being peculiar to it, while most of the
European genera are also represented.
_Fresh-water Fish._--Of these there are a few peculiar genera; as
_Plecoglossus_ (Salmonidæ) from Japan; _Achilognathus_, _Pseudoperilampus_,
_Ochetobius_, and _Opsariichthys_ (Cyprinidæ); and there are many other
Chinese Cyprinidæ belonging to the border land of the Palæarctic and
Oriental regions.
_Insects._--The butterflies of this sub-region exhibit the same mixture of
tropical and temperate forms as the birds. Most of the common European
genera are represented, and there are species of _Parnassius_ in Japan and
the Amoor. _Isodema_, a peculiar genus of Nymphalidæ is found near Ningpo,
just within our limits; and _Sericinus_, one of the most beautiful genera
of Papilionidæ is peculiar to North China, where four species occur, thus
balancing the _Thais_ and _Doritis_ of Europe. The genus _Zephyrus_
(Lycænidæ) is well represented by six species in Japan and the Amoor,
against two in Europe. _Papilio paris_ and _P. bianor_, magnificent insects
of wholly tropical appearance, abound near Pekin, and allied forms inhabit
Japan and the Amoor, as well as _P. demetrius_ and _P. alcinous_ belonging
to the "Protenor" group of the Himalayas. Other tropical genera occurring
in Japan, the Amoor, or North China are, _Debis_, _Neope_, _Mycalesis_,
_Ypthimia_ (Satyridæ); _Thaumantis_ (Morphidæ), at Shanghae; _Euripus_,
_Neptis_, _Athyma_ (Nymphalidæ); _Terias_ (Pieridæ); and the
above-mentioned Papilionidæ.
{228}_Coleoptera._--The beetles of Japan decidedly exhibit a mixture of
tropical forms with others truly Palæarctic, and it has been with some
naturalists a matter of doubt whether the southern and best known portion
of the islands should not be joined to the Oriental region. An important
addition to our knowledge of the insects of this country has recently been
made by Mr. George Lewis, and a portion of his collections have been
described by various entomologists in the _Transactions of the
Entomological Society of London_. As the question is one of considerable
interest we shall give a summary of the results fairly deducible from what
is now known of the entomology of Japan; and it must be remembered that
almost all our collections come from the southern districts, in what is
almost a sub-tropical climate; so that if we find a considerable proportion
of Palæarctic forms, we may be pretty sure that the preponderance will be
much greater a little further north.
Of Carabidæ Mr. Bates enumerates 244 species belonging to 84 genera, and by
comparing these with the Coleoptera of a tract of about equal extent in
western Europe, he concludes that there is little similarity, and that the
cases of affinity to the forms of eastern tropical Asia preponderate. By
comparing his genera with the distributions as given in _Gemminger and
Harold's Catalogue_, a somewhat different result is arrived at. Leaving out
the generic types altogether peculiar to Japan, and also those genera of
such world-wide distribution that they afford no clear indications for our
purpose, it appears that no less than twenty-two genera, containing
seventy-four of the Japanese species, are either exclusively Palæarctic,
Palæarctic and Nearctic, or highly characteristic of the Palæarctic region;
then come thirteen genera containing eighty-seven of the species which have
a very wide distribution, but are also Palæarctic: we next have seventeen
genera containing twenty-four of the Japanese species which are decidedly
Oriental and tropical. Here then the fair comparison is between the
twenty-two genera and seventy-four species whose affinities are clearly
Palæarctic or at least north temperate, and seventeen genera with
twenty-four species which are Asiatic and tropical; and this seems to prove
that, although South {229}Japan (like North China) has a considerable
infusion of tropical forms, there is a preponderating substratum of
Palæarctic forms, which clearly indicate the true position of the islands
in zoological geography. There are also a few cases of what may be called
eccentric distribution; which show that Japan, like many other
island-groups, has served as a kind of refuge in which dying-out forms
continue to maintain themselves. These, which are worthy of notice, are as
follows: _Orthotrichus_ (1 sp.) has the only other species in Egypt;
_Trechichus_ (1 sp.) has two other species, of which one inhabits Madeira,
the other the Southern United States; _Perileptus_ (1 sp.) has two other
species, of which one inhabits Bourbon, the other West Europe; and lastly,
_Crepidogaster_ (1 sp.) has the other known species in South Africa. These
cases diminish the value of the indications afforded by some of the
Japanese forms, whose only allies are single species in various remote
parts of the Oriental region.
The Staphylinidæ have been described by Dr. Sharp, and his list exhibits a
great preponderance of north temperate, or cosmopolitan forms, with a few
which are decidedly tropical. The Pselaphidæ and Scydmenidæ, also described
by Dr. Sharp, exhibit, according to that gentleman, "even a greater
resemblance to those of North America than to those of Europe," but he says
nothing of any tropical affinities. The water-beetles are all either
Palæarctic or of wide distribution.
The Lucanidæ (_Gemm. and Har. Cat._, 1868) exhibit an intermingling of
Palæarctic and Oriental genera.
The Cetoniidæ (_Gemm. and Har. Cat._, 1869) show, for North China and
Japan, three Oriental to two Palæarctic genera.
The Buprestidæ collected by Mr. Lewis have been described by Mr. Edward
Saunders in the _Journal of the Linnæan Society_, vol. xi. p. 509. The
collection consisted of thirty-six species belonging to fourteen genera. No
less than thirteen of these are known also from India and the Malay
Islands; nine from Europe; seven from Africa; six from America, and four
from China. In six of the genera the Japanese species are said to be allied
to those of the Oriental region; while in three they are allied to European
forms, and in two to American. Considering {230}the southern latitude and
warm climate in which these insects were mostly collected, and the
proximity to Formosa and the Malay Islands compared with the enormous
distance from Europe, this shows as much Palæarctic affinity as can be
expected. In the Palæarctic region the group is only plentiful in the
southern parts of Europe, which is cut off by the cold plateau of Thibet
from all direct communication with Japan; while in the Oriental region it
everywhere abounds and is, in fact, one of the most conspicuous and
dominant families of Coleoptera.
The Longicorns collected by Mr. Lewis have been described by Mr. Bates in
the _Annals of Natural History for 1873_. The number of species now known
from Japan is 107, belonging to sixty-four genera. The most important
genera are _Leptura_, _Clytanthus_, _Monohammus_, _Praonetha_,
_Exocentrus_, _Glenea_, and _Oberea_. There are twenty-one tropical genera,
and seven peculiar to Japan, leaving thirty-six either Palæarctic or of
very wide range. A number of the genera are Oriental and Malayan, and many
characteristic European genera seem to be absent; but it is certain that
not half the Japanese Longicorns are yet known, and many of these gaps will
doubtless be filled up when the more northern islands are explored.
The Phytophaga, described by Mr. Baly, appear to have a considerable
preponderance of tropical Oriental forms.
A considerable collection of Hymenoptera formed by Mr. Lewis have been
described by Mr. Frederick Smith; and exhibit the interesting result, that
while the bees and wasps are decidedly of tropical and Oriental forms, the
Tenthredinidæ and Ichneumonidæ are as decidedly Palæarctic, "the general
aspect of the collection being that of a European one, only a single exotic
form being found among them."
_Remarks on the General Character of the Fauna of Japan._--From a general
view of the phenomena of distribution we feel justified in placing Japan in
the Palæarctic region; although some tropical groups, especially of
reptiles and insects, have largely occupied its southern portions; and
these same groups have in many cases spread into Northern China, beyond the
{231}usual dividing line of the Palæarctic and Oriental regions. The causes
of such a phenomenon are not difficult to conceive. Even now, that portion
of the Palæarctic region between Western Asia and Japan is, for the most
part, a bleak and inhospitable region, abounding in desert plateaus, and
with a rigorous climate even in its most favoured districts, and can,
therefore, support but a scanty population of snakes, and of such groups of
insects as require flowers, forests, or a considerable period of warm
summer weather; and it is precisely these which are represented in Japan
and North China by tropical forms. We must also consider, that during the
Glacial epoch this whole region would have become still less productive,
and that, as the southern limit of the ice retired northward, it would be
followed up by many tropical forms along with such as had been driven south
by its advance, and had survived to return to their northern homes.
It is also evident that Japan has a more equable and probably moister
climate than the opposite shores of China, and has also a very different
geological character, being rocky and broken, often volcanic, and
supporting a rich, varied, and peculiar vegetation. It would thus be well
adapted to support all the more hardy denizens of the tropics which might
at various times reach it, while it might not be so well adapted for the
more boreal forms from Mongolia or Siberia. The fact that a mixture of such
forms occurs there, is then, little to be wondered at, but we may rather
marvel that they are not more predominant, and that even in the extreme
south, the most abundant forms of mammal, bird, and insect, are
modifications of familiar Palæarctic types. The fact clearly indicates that
the former land connections of Japan with the continent have been in a
northerly rather than in a southerly direction, and that the tropical
immigrants have had difficulties to contend with, and have found the land
already fairly stocked with northern aborigines in almost every class and
order of animals.
_General Conclusions as to the Fauna of the Palæarctic Region._--From the
account that has now been given of the fauna {232}of the Palæarctic region,
it is evident that it owes many of its deficiencies and some of its
peculiarities to the influence of the Glacial epoch, combined with those
important changes of physical geography which accompanied or preceded it.
The elevation of the old Sarahan sea and the complete formation of the
Mediterranean, are the most important of these changes in the western
portion of the region. In the centre, a wide arm of the Arctic Ocean
extended southward from the Gulf of Obi to the Aral and the Caspian,
dividing northern Europe and Asia. At this time our European and Siberian
sub-regions were probably more distinct than they are now, their complete
fusion having been effected since the Glacial epoch. As we know that the
Himalayas have greatly increased in altitude during the Tertiary period, it
is not impossible that during the Miocene and Pliocene epochs the vast
plateau of Central Asia was much less elevated and less completely cut off
from the influence of rain-bearing winds. It might then have been far more
fertile, and have supported a rich and varied animal population, a few
relics of which we see in the Thibetan antelopes, yaks, and wild horses.
The influence of yet earlier changes of physical geography, and the
relations of the Palæarctic to the tropical regions immediately south of
it, will be better understood when we have examined and discussed the
faunas of the Ethiopian and Oriental regions.
{233}TABLES OF DISTRIBUTION.
In constructing these tables showing the distribution of various classes of
animals in the Palæarctic region, the following sources of information have
been chiefly relied on, in addition to the general treatises, monographs,
and catalogues used in compiling the fourth part of this work.
_Mammalia._--Lord Clement's Mammalia and Reptiles of Europe; Siebold's
Fauna Japonica; Père David's List of Mammalia of North China and Thibet;
Swinhoe's Chinese Mammalia; Radde's List of Mammalia of South-Eastern
Siberia; Canon Tristram's Lists for Sahara and Palestine; Papers by
Professor Milne-Edwards, Mr. Blanford, Mr. Sclater, and the local lists
given by Mr. A. Murray in the Appendix to his Geographical Distribution of
Mammalia.
_Birds._--Blasius' List of Birds of Europe; Godman, On Birds of Azores,
Madeira, and Canaries; Middendorf, for Siberia; Père David and Mr. Swinhoe,
for China and Mongolia; Homeyer, for East Siberia; Mr. Blanford, for Persia
and the high Himalayas; Mr. Elwes's paper on the Distribution of Asiatic
Birds; Canon Tristram, for the Sahara and Palestine; Professor Newton, for
Iceland and Greenland; Mr. Dresser, for Scandinavia; and numerous papers
and notes in the Ibis; Journal für Ornithologie; Annals and Mag. of Nat.
History; and Proceedings of the Zoological Society.
_Reptiles and Amphibia._--Schreiber's European Herpetology.
{234}TABLE I.
_FAMILIES OF ANIMALS INHABITING THE PALÆARCTIC REGION._
EXPLANATION.
Names in _italics_ show families peculiar to the region.
Names inclosed thus (......) barely enter the region, and are not
considered properly to belong to it.
Numbers are not consecutive, but correspond to those in Part IV.
---------------------+-------------------+-------------------------------
| Sub-regions |
| 1=Europe. |
Order and Family | 2=Mediterranean. | Range beyond the Region.
| 3=Siberian. |
| 4=Japan. |
---------------------+----+----+----+----+-------------------------------
| 1. | 2. | 3. | 4. |
---------------------+----+----+----+----+-------------------------------
| | | | |
MAMMALIA. | | | | |
PRIMATES. | | | | |
3. Cynopithecidæ | | -- | | -- |Ethiopian, Oriental
| | | | |
CHIROPTERA. | | | | |
9. (Pteropidæ) | | | | -- |Tropics of E. Hemisphere
11. Rhinolophidæ | -- | -- | -- | -- |Warmer parts of E. Hemis.
12. Vespertilionidæ | -- | -- | -- | -- |Cosmopolite
13. Noctilionidæ | | -- | | |Tropical regions
| | | | |
INSECTIVORA. | | | | |
15. Macroscelididæ | | -- | | |Ethiopian
17. Erinaceidæ | -- | -- | -- | -- |Oriental, S. Africa
21. Talpidæ | -- | -- | -- | -- |Nearctic, Oriental
22. Soricidæ | -- | -- | -- | -- |Cosmopolite, excl. Australia
| | | | | and S. America
| | | | |
CARNIVORA. | | | | |
23. Felidæ | -- | -- | -- | -- |All regions but Australian
25. Viverridæ | | -- | | |Ethiopian, Oriental
27. Hyænidæ | | -- | | |Ethiopian, Oriental
28. Canidæ | -- | -- | -- | -- |All regions but Australian
29. Mustelidæ | -- | -- | -- | -- |All regions but Australian
31. Æluridæ | | | | -- |Oriental
32. Ursidæ | -- | -- | -- | -- |Nearctic, Oriental, Andes
33. Otariidæ | | | | -- |N. and S. temperate zones
34. Trichechidæ | -- | | -- | |Arctic regions
35. Phocidæ | -- | -- | -- | -- |N. and S. temperate zones
| | | | |
CETACEA. | | | | |
36 to 41. | | | | |Oceanic
| | | | |
SIRENIA. | | | | |
42. Manatidæ | -- | | -- | |Tropics, from Brazil to
| | | | | N. Australia
| | | | |
UNGULATA. | | | | |
43. Equidæ | | -- | -- | |Ethiopian
47. Suidæ | -- | -- | -- | -- |Cosmopolite, excl. Nearctic
| | | | | reg. and Australia
48. Camelidæ | | -- | -- | |Andes
50. Cervidæ | -- | -- | -- | -- |All regions but Ethiopian and
| | | | | Australian
52. Bovidæ | -- | -- | -- | -- |All regions but Neotropical and
| | | | | Australian
| | | | |
HYRACOIDÆ. | | | | |
54. (Hyracidæ) | | -- | | |Ethiopian family
| | | | |
RODENTIA. | | | | |
55. Muridæ | -- | -- | -- | -- |Almost Cosmopolite
56. Spalacidæ | -- | -- | -- | |Ethiopian, Oriental
57. Dipodidæ | | -- | -- | -- |Ethiopian, Nearctic
58. Myoxidæ | -- | -- | -- | -- |Ethiopian
60. Castoridæ | -- | | -- | |Nearctic
61. Sciuridæ | -- | -- | -- | -- |All regions but Australian
64. Octodontidæ | | -- | | |Abyssinia, Neotropical
67. Hystricidæ | | -- | | |Ethiopian, Oriental
69. Lagomyidæ | | | -- | |Nearctic
70. Leporidæ | -- | -- | -- | -- |All regions but Australian
| | | | |
BIRDS. | | | | |
PASSERES. | -- | -- | -- | -- |
1. Turdidæ | -- | -- | -- | -- |Cosmopolite
2. Sylviidæ | | -- | | -- |Cosmopolite
3. Timaliidæ | -- | -- | -- | -- |Ethiopian, Oriental, Australian
4. Panuridæ | -- | -- | -- | -- |Nearctic, Oriental
5. Cinclidæ | -- | -- | -- | -- |Oriental
6. Troglodytidæ | -- | -- | -- | -- |American, Oriental
8. Certhiidæ | -- | -- | -- | -- |Oriental, Nearctic
9. Sittidæ | -- | -- | -- | -- |Nearctic, Oriental, Australian,
| | | | | Madagascar
10. Paridæ | -- | -- | | -- |Nearctic, Oriental, Australian
| | | | | [?]
13. Pycnonotidæ | | -- | | -- |Oriental, Ethiopian
14. Oriolidæ | -- | -- | -- | -- |Ethiopian, Oriental, Australian
17. Muscicapidæ | | | | |Eastern Hemisphere
19. Laniidæ | -- | -- | -- | -- |Eastern Hemisphere and N.
| | | | | America
20. Corvidæ | -- | -- | -- | -- |Cosmopolite
23. (Nectariniidæ) | | -- | | |Ethiopian, Oriental, Australian
24. (Dicæidæ) | | | | -- |Ethiopian, Oriental, Australian
29. Ampelidæ | -- | -- | -- | -- |Nearctic
30. Hirundinidæ | -- | -- | -- | -- |Cosmopolite
33. Fringillidæ | -- | -- | -- | -- |All regions but Australian
35. Sturnidæ | -- | -- | -- | -- |Eastern Hemisphere
37. Alaudidæ | -- | -- | -- | -- |All regions but Neotropical
38. Motacillidæ | -- | -- | -- | -- |Cosmopolite
47. (Pittidæ) | | | | -- |Oriental, Australian, Ethiopian
| | | | |
PICARIÆ. | | | | |
51. Picidæ | -- | -- | -- | -- |All regions but Australian
52. Yungidæ | -- | -- | -- | -- |N. W. India, N. E. Africa,
| | | | | S. Africa
58. Cuculidæ | -- | -- | -- | -- |Almost Cosmopolite
62. Coraciidæ | -- | -- | -- | -- |Ethiopian, Oriental, Australian
63. Meropidæ | -- | -- | | |Ethiopian, Oriental, Australian
67. Alcedinidæ | -- | -- | -- | -- |Cosmopolite
69. Upupidæ | | -- | | -- |Ethiopian, Oriental
73. Caprimulgidæ | -- | -- | -- | -- |Cosmopolite
74. Cypselidæ | -- | -- | -- | -- |Almost Cosmopolite
| | | | |
COLUMBÆ. | | | | |
84. Columbidæ | -- | -- | -- | -- |Cosmopolite
| | | | |
GALLINÆ. | | | | |
86. Pteroclidæ | -- | -- | -- | -- |Ethiopian, Indian
87. Tetraonidæ | -- | -- | -- | -- |Nearctic, Ethiopian, Oriental
88. Phasianidæ | | -- | -- | -- |Oriental, Ethiopian, Nearctic
89. Turnicidæ | | -- | | -- |Ethiopian, Oriental, Australian
| | | | |
ACCIPITRES. | | | | |
94. Vulturidæ | -- | -- | -- | -- |All regions but Australian
96. Falconidæ | -- | -- | -- | -- |Cosmopolite
97. Pandionidæ | -- | -- | -- | -- |Cosmopolite
98. Strigidæ | -- | -- | -- | -- |Cosmopolite
| | | | |
GRALLÆ. | | | | |
99. Rallidæ | -- | -- | -- | -- |Cosmopolite
100. Scolopacidæ | -- | -- | -- | -- |Cosmopolite
104. Glareolidæ | -- | -- | -- | -- |Ethiopian, Oriental, Australian
105. Charadriidæ | -- | -- | -- | -- |Cosmopolite
106. Otididæ | -- | -- | -- | -- |Ethiopian, Oriental, Australian
107. Gruidæ | -- | -- | -- | -- |Eastern Hemisphere, and
| | | | | N. America
113. Ardeidæ | -- | -- | -- | -- |Cosmopolite
114. Plataleidæ | -- | -- | -- | -- |Almost Cosmopolite
115. Ciconiidæ | -- | -- | -- | -- |Nearly Cosmopolite
117. Phænicopteridæ | | -- | | |Neotropical, Ethiopian, Indian
| | | | |
ANSERES. | | | | |
118. Anatidæ | -- | -- | -- | -- |Cosmopolite
119. Laridæ | -- | -- | -- | -- |Cosmopolite
120. Procellariidæ | -- | -- | -- | -- |Cosmopolite
121. Pelecanidæ | -- | -- | -- | -- |Cosmopolite
123. Colymbidæ | -- | | -- | -- |Arctic and N. Temperate
124. Podicipidæ | -- | -- | -- | -- |Cosmopolite
125. Alcidæ | -- | | -- | -- |N. Temperate zone
| | | | |
REPTILIA. | | | | |
OPHIDIA. | | | | |
1. Typhlopidæ | | -- | | -- |All regions but Nearctic
5. Calamariidæ | | -- | | |All other regions
6. Oligodontidæ | | | | -- |Oriental and Neotropical
7. Colubridæ | -- | -- | -- | -- |Almost Cosmopolite
8. Homalopsidæ | | -- | -- | -- |Oriental, and all other regions
9. Psammophidæ | | -- | | |Ethiopian and Oriental
18. Erycidæ | | -- | | |Oriental and Ethiopian
20. Elapidæ | | | | -- |Australian and all other
| | | | | regions
24. Crotalidæ | | | -- | -- |Nearctic, Neotropical, Oriental
25. Viperidæ | -- | -- | -- | -- |Ethiopian, Oriental
| | | | |
LACERTILIA. | | | | |
26. _Trogonophidæ_ | | -- | | |
28. Amphisbænidæ | | -- | | |Ethiopian, Neotropical
30. Varanidæ | | -- | | |Oriental, Ethiopian, Australian
33. Lacertidæ | -- | -- | -- | -- |All continents but American
34. Zonuridæ | | -- | | |America, Africa, N. India
41. Gymnopthalmidæ | -- | -- | -- | |Ethiopian, Australian,
| | | | | Neotropical
45. Scincidæ | -- | -- | -- | -- |Almost Cosmopolite
46. _Ophiomoridæ_ | | -- | | |
47. Sepidæ | | -- | | |Ethiopian
49. Geckotidæ | | -- | -- | -- |Almost Cosmopolite
51. Agamidæ | | -- | -- | -- |All continents but America
52. Chamæleonidæ | | -- | | |Ethiopian, Oriental
| | | | |
CHELONIA. | | | | |
57. Testudinidæ | -- | -- | | -- |All continents but Australia
59. Trionychidæ | | | | -- |Ethiopian, Oriental, Nearctic
60. Cheloniidæ | | | | |Marine
| | | | |
AMPHIBIA. | | | | |
URODELA. | | | | |
3. Proteidæ | -- | | | |Nearctic
5. Menopomidæ | | | | -- |Nearctic
6. Salamandridæ | -- | -- | -- | -- |Nearctic to Andes of Bogota
| | | | |
ANOURA. | | | | |
10. Bufonidæ | -- | -- | -- | -- |All continents but Australia
13. Bombinatoridæ | -- | -- | | |Neotropical, New Zealand
15. Alytidæ | -- | | | |All regions but Oriental
17. Hylidæ | -- | -- | -- | |All regions but Ethiopian
18. Polypedatidæ | | | -- | -- |All the regions
19. Ranidæ | -- | -- | -- | -- |Almost Cosmopolite
20. Discoglossidæ | -- | -- | -- | -- |All regions but Nearctic
| | | | |
FISHES (FRESH-WATER).| | | | |
ACANTHOPTERYGII. | | | | |
1. Gasterosteidæ | -- | -- | -- | -- |Nearctic
3. Percidæ | -- | -- | -- | -- |All regions but Australian
12. Scienidæ | -- | -- | | -- |All regions but Australian
26. _Comephoridæ_ | | | -- | |
37. Atherinidæ | -- | -- | | |N. America and Australia
| | | | |
PHYSOSTOMI. | | | | |
59. Siluridæ | -- | -- | -- | -- |All warm regions
65. Salmonidæ | -- | -- | -- | -- |Nearctic, New Zealand
70. Esocidæ | -- | -- | | |Nearctic
71. Umbridæ | -- | | | |Nearctic
73. Cyprinodontidæ | | -- | | |All regions but Australia
75. Cyprinidæ | -- | -- | -- | -- |All regions but Australian and
| | | | | Neotropical
| | | | |
GANOIDEI. | | | | |
96. Accipenseridæ | -- | -- | -- | |Nearctic
97. Polydontidæ | | | | -- |Nearctic
| | | | |
INSECTS. | | | | |
LEPIDOPTERA (PART). | | | | |
DURINI (BUTTERFLIES).| | | | |
1. Danaidæ | -- | -- | | -- |All tropical regions
2. Satyridæ | -- | -- | -- | -- |Cosmopolite
8. Nymphalidæ | -- | -- | -- | -- |Cosmopolite
9. Libytheidæ | -- | -- | | |All continents but Australia
10. Nemeobeidæ | -- | | | |Absent from Nearctic region and
| | | | | Australia
13. Lycænidæ | -- | -- | -- | -- |Cosmopolite
14. Pieridæ | -- | -- | -- | -- |Cosmopolite
15. Papilionidæ | -- | -- | -- | -- |Cosmopolite
16. Hesperidæ | -- | -- | -- | -- |Cosmopolite
| | | | |
SPHINGIDEA. | | | | |
17. Zygænidæ | -- | -- | -- | -- |Cosmopolite
21. Stygiidæ | -- | -- | -- | -- |Neotropical
22. Ægeriidæ | -- | -- | -- | -- |Absent only from Australia
23. Sphingidæ | -- | -- | -- | -- |Cosmopolite
---------------------+----+----+----+----+-------------------------------
COLEOPTERA.--Of about 80 families into which the Coleoptera are divided,
all the more important are cosmopolite, or nearly so. It would therefore
unnecessarily occupy space to give tables of the whole for each region.
LAND SHELLS.--The more important families being cosmopolite, and the
smaller ones being somewhat uncertain in their limits, the reader is
referred to the account of the families and genera under each region, and
to the chapter on Mollusca in the concluding part of this work, for such
information as can be given of their distribution.
{239}TABLE II.
_LIST OF THE GENERA OF TERRESTIAL MAMMALIA AND BIRDS INHABITING THE
PALÆARCTIC REGION._
EXPLANATION.
Names in _italics_ show genera peculiar to the region.
Names inclosed thus (...) show genera which just enter the region, but
are not considered properly to belong to it.
Genera which undoubtedly belong to the region are numbered consecutively.
_MAMMALIA._
-------------------+-------+----------------------+----------------------
Order, Family, and | No. of| Range within | Range beyond
Genus. |Species| the Region. | the Region.
-------------------+-------+----------------------+----------------------
| | |
PRIMATES. | | |
SEMNOPITHECIDÆ. | | |
| | |
(Semnopithecus | 1 |Eastern Thibet) |Oriental genus
| | |
CYNOPITHECIDÆ. | | |
| | |
1. Macacus | 4 |Gibraltar, N. Africa, |Oriental
| | E. Thibet to Japan |
| | |
CHIROPTERA. | | |
PTEROPIDÆ. | | |
| | |
(Pteropus | 2 |Egypt, Japan) |Tropics of the E. .
| | | Hemis
(Xantharpyia | 1 |N. Africa, Palestine) |Oriental, Austro-
| | | Malayan
| | |
RHINOLOPHIDÆ. | | |
| | |
2. Rhinolphus | 9 |Temperate & Southern |Warmer parts E.
| | parts of Region | Hemisphere
(Asellia | 1 |Egypt) |Ethiopian, Java
(_Rhinopoma_ | 1 |Egypt, Palestine) |[?] India
(Nycteris | 1 |Egypt) |Nubia, Himalaya
| | |
VESPERTILIONIDÆ. | | |
| | |
3. Vesperugo | 1 |Siberia, Amoorland |[?]
4. _Otonycteris_ | 1 |Egypt |[?]
5. Vespertilio | 35 |The whole region |Cosmopolite
(Kerivoula | 1 |N. China) |Oriental, S. Africa
6. Miniopteris | 1 |S. Europe, N. Africa, |S. Afric. Malaya,
| | Japan | Austral.
7. Plecotus | 1 |S. Europe |Himalayas
8. Barbastellus | 2 |Mid. and S. Europe, |Darjeeling, Timor
| | Palestine |
| | |
NOCTILIONIDÆ. | | |
| | |
9. Molossus | 2 |S. Europe, N. Africa |Ethiop., Neotrop.,
| | | Australian
| | |
INSECTIVORA. | | |
ERINACEIDÆ. | | |
| | |
10. Erinaceus | 4 |The whole region; excl.|Oriental, Africa.
| | Japan |
| | |
TALPIDÆ. | | |
| | |
11. _Talpa_ | 5 |The whole region |N. India
12. _Scaptochirus_ | 1 |N. China |
13. _Anurosorex_ | 1 |N. China |
14. _Scaptonyx_ | 1 |N. China |
15. _Myogale_ | 2 |S. E. Russia, Pyrenees |
16. _Nectogale_ | 1 |Thibet |
17. Urotrichus | 1 |Japan |N. W. America
18. _Uropsilus_ | 1 |E. Thibet |
| | |
SORICIDÆ. | | |
| | |
19. Sorex | 10 |The whole region |Absent from Australia
| | | & S. America
20. Crocidura | 4 |W. Europe to N. China | [?]
| | |
CARNIVORA. | | |
FELIDÆ. | | |
| | |
21. Felis | 12 |The whole region; excl.|All regions but
| | extreme North | Austral.
22. Lyncus | 9 |S. Europe to Arctic sea|America N. of 66° N.
| | | Lat.
| | |
VIVERRIDÆ. | | |
| | |
(Viverra | 1 |N. China) |Oriental and Ethiopian
23. Genetta | 1 |S. Europe & N. Africa, |Ethiopian
| | Palestine |
(Herpestes | 1 |N. Africa, Spain [?], |Oriental and Ethiopian
| | Palestine) |
| | |
HYÆNIDÆ. | | |
| | |
24. Hyæna | 1 |N. Africa and S. W. |Ethiopian, India
| | Asia |
| | |
CANIDÆ. | | |
| | |
25. Canis | 4 |The whole region |All reg. but Austral.
| | | [?]
26. _Nyctereutes_ | 1 |Japan, Amoorland, N. |
| | China |
| | |
MUSTELIDÆ. | | |
| | |
27. Martes | 7 |N. Europe and Asia, E. |Oriental, Nearctic
| | Thibet |
28. _Putorius_ | 3 |W. Europe to N. E. Asia|
29. Mustela | 10 |The whole region |Nearctic, Ethiop.,
| | | Himalayas, Peru
30. Vison | 2 |Europe and Siberia |N. America, N. India,
| | | China
31. Gulo | 1 |The Arctic regions |Arctic America
32. Lutra | 2 |The whole region |Oriental
33. _Lutronectes_ | 1 |Japan |
34. Enhydris | 1 |N. Asia and Japan |California
35. _Meles_ | 2 |Cen. Europe, Palestine,|China to Hongkong
| | N. China, Japan |
| | |
ÆLURIDÆ. | | |
| | |
36. Ælurus | 1 |S. E. Thibet |Nepal
37. _Æluropus_ | 1 |E. Thibet |
| | |
URSIDÆ. | | |
| | |
38. Thalassarctos | 1 |Arctic regions |Arctic America
39. Ursus | 4 |The whole region |Oriental, Nearctic,
| | | Chili
| | |
OTARIIDÆ. | | |
| | |
40. Callorhinus | 1 |Kamschatka and |
| | Behring's Straits |
41. Zalophus | 1 |Japan |California
42. Eumetopias | 1 |Japan, Behring's |California
| | Straits |
| | |
TRICHECHIDÆ. | | |
| | |
43. Trichechus | 1 |Polar Seas |Arctic America
| | |
PHOCIDÆ. | | |
| | |
44. Callocephalus | 3 |North Sea, Caspian, |Greenland
| | Lake Baikal |
45. Pagomys | 2 |North Sea, Japan |N. Pacific
46. Pagophilus | 2 |Northern Seas |N. Pacific
47. Phoca | 2 |Northern Seas |N. Pacific
48. Halichærus | 1 |North Sea and Baltic |Greenland
49. _Pelagius_ | 2 |Madeira to Black Sea |
50. Cystophora | 2 |N. Atlantic |N. Atlantic
| | |
SIRENIA. | | |Tropics & Behring's
| | | Strts.
| | |
CETACEA. | | |Oceanic
| | |
UNGULATA. | | |
EQUIDÆ. | | |
| | |
51. Equus | 4 |Cent. and W. Asia & N. |Ethiopian
| | Africa |
| | |
SUIDÆ. | | |
| | |
52. Sus | 2 |The whole region |Oriental, Austro-
| | | Malayan
| | |
CAMELIDÆ. | | |
| | |
53. _Camelus_ | 2 |Deserts of Cent. and W.|
| | Asia and N. Africa |
| | |
CERVIDÆ. | | |
| | |
54. Alces | 1 |North Europe and Asia |N. America
55. Tarandus | 1 |Arctic Europe and Asia |Arctic America
56. Cervus | 8 |The whole region |All regions but
| | | Austral.
57. _Dama_ | 1 |Mediterranean district |
58. _Elaphodus_ | 1 |N. W. China |
59. _Lophotragus_ | 1 |N. China |
60. _Capreolus_ | 2 |Temp. Europe and W. |
| | Asia and N. China |
61. _Moschus_ | 1 |Amoor R., N. China, to |
| | Himalayas |
62. _Hydropotes_ | 1 |N. China |
| | |
BOVIDÆ. | | |
| | |
63. { Bos | 1 |Europe, (not wild) |Oriental
64. { Bison | 1 |Poland and Caucasus |Nearctic
65. { _Poephagus_ | 1 |Thibet |
66. _Addax_ | 1 |N. Africa to Syria |
67. Oryx | 1 |N. Africa to Syria |Ethiopian deserts
68. { Gazella | 12 |N. Africa to Persia, |S. Africa, India
{ | | and Beloochistan |
69. { _Procapra_ | 2 |W. Thibet and Mongolia |
70. {_Saiga_ | 1 |E. Europe and W. Asia |
71. {_Pantholops_ | 1 |W. Thibet |
(Alcephalus | 1 |Syria) |Ethiopian genus.
72. _Budorcas_ | 2 |E. Himalayas to E. |
| | Thibet |
73. _Rupicapra_ | 2 |Pyrenees to Caucasus |
74. Nemorhedus | 7 |E. Himalayas to E. |Oriental to Sumatra,
| | China and Japan | Formosa
75. Capra | 20 |Spain to Thibet and |Nilgherries, Rocky
| | N.E. Africa | Mtns.
| | |
HYRACOIDEA. | | |
HYRACIDÆ. | | |
| | |
(Hyrax | 1 |Syria) |Ethiopian genus
| | |
RODENTIA. | | |
MURIDÆ. | | |
| | |
76. Mus |?15 |The whole region |E. Hemisphere
77. _Cricetus_ | 9 |The whole region |
78. _Cricetulus_ | 3 |N. China |
79. Meriones | 8 |W. and Central Asia to |Ethiopian, Indian.
| | N. China, N. Africa |
80. _Rhombomys_ | 6 |E. Europe, Cent. Asia, |
| | N. Africa |
81. _Psammomys_ | 3 |Egypt and Palestine |
82. _Sminthus_ | 3 |East Europe, Siberia |
83. Arvicola |?21 |The whole region |Himalayas, Nearctic
84. Cuniculus | 1 |N. E. Europe, Siberia |Arctic America
85. Myodes | 1 |North of region |Nearctic
86. _Myospalax_ | 3 |Altai Mountains and N. |
| | China |
| | |
SPALACIDÆ. | | |
| | |
87. _Ellobius_ | 1 |S. Russia and S. W. |
| | Siberia |
88. _Spalax_ | 1 |Hungary and Greece to |
| | W. Asia, Palestine |
| | |
DIPODIDÆ. | | |
| | |
89. Dipus |?15 |S. E. Europe and N. |Africa, India
| | Africa to N. China |
| | |
MYOXIDÆ. | | |
| | |
90. Myoxus | 12 |Temperate parts of |Ethiopian
| | whole region |
| | |
CASTORIDÆ. | | |
| | |
91. Castor | 1 |Temperate zone, from |N. America
| | France to Amoorland |
| | |
SCIURIDÆ. | | |
| | |
92. Sciurus | 8 |The whole region |All regions but
| | | Austral.
92a. Tamias | 1 |All Northern Asia |N. America
93. Sciuropterus | 4 |Finland to Siberia and |Oriental, Nearctic
| | Japan |
94. Pteromys | 3 |Japan and W. China |Oriental
95. Spermophilus | 10 |E. Europe to N. China |Nearctic
| | and Kamschatka |
96. Arctomys. | 4 |Alps to E. Thibet and |Nearctic
| | Kamschatka |
| | |
OCTODONTIDÆ. | | |
| | |
97._Ctenodactylus_ | 1 |N. Africa |
| | |
HYSTRICIDÆ. | | |
| | |
98. Hystrix | 2 |S. Europe, Palestine, |Ethiopian, Oriental
| | N. China. |
| | |
LAGOMYIDÆ. | | |
| | |
99. Lagomys | 10 |Volga to E. Thibet and |Nearctic
| | Kamschatka |
| | |
LEPORIDÆ. | | |
| | |
100. Lepus | 12 |The whole region |All regions but
| | | Austral.
_BIRDS._
PASSERES. | | |
TURDIDÆ. | | |
| | |
1. Turdus | 18 |The whole region |Almost cosmopolite
| |(excluding Spitsbergen)|
2. Oreocincla | 1 |N. E. Asia and Japan, |Oriental and
| | straggler to Europe | Australian
3. Monticola | 3 |S. Europe, N. Africa, |Oriental and
| | Palestine, N. China | S. African
(Bessornis | 1 |Palestine) |Tropical and
| | | S. Africa
| | |
SYLVIIDÆ. | | |
| | |
4. Cisticola | 1 |S. W. Europe, N. |Ethiop., Orient.,
| | Africa, Japan | Austral.
5. {Acrocephalus | 10 |W. Europe to Japan |Orient., Ethiop.,
{ | | | Austral.
6. {_Dumeticola_ | 4 |Nepaul, Lake Baikal, |
{ | | E. Thibet high |
7. {_Potamodus_ | 3 |W. and S. Europe, |
{ | | N. Africa, E. Thibet |
8. {_Lusciniola_ | 1 |S. Europe |
9. {_Locustella_ | 7 |W. Europe and N. Africa|India, winter
{ | | to Japan | migrants(?)
10. {Bradyptetus | 2 |S. Europe and Palestine|E. and S. Africa
11. {_Calamodus_ | ?3 |Europe, N. Africa, |
| | Palestine |
12. {Phylloscopus | 6 |The whole region |Oriental
{ | | (excluding western |
{ | | islands) |
13. {Hypolais | 9 |Europe, N. Africa, |China, Moluccas,
{ | | Palestine, China | India, Africa
14. {Abrornis | 2 |Cashmere, E. Thibet |Oriental region
15. {Reguloides | 2 |Europe and China |N. India, Formosa
16. {Regulus | 4 |The whole region |N. and Central America
| | (excluding Iceland, |
| | &c.) |
17. {Aedon | 2 |S. Europe, W. Asia, |E. and S. Africa
{ | | N. Africa |
18. {_Pyrophthalma_ | 2 |E. Europe and Palestine|
19. {_Melizophilus_ | 2 |W. and S. Europe, |
{ | | Sardinia |
20. {_Sylvia_ | 6 |Madeira to W. India, |N. E. Africa, Ceylon
{ | | N. Africa | migrants(?)
21. {_Curruca_ | 7 |Madeira to India, |E. Africa, India,
| | N. Africa | migrants
22. {_Luscinia_ | 2 |W. Europe, N. Africa, |
{ | | Persia |
23. {_Cyanecula_ | 3 |Europe and N. Africa to|Abyssinia and India
{ | | Kamschatka | migrants
24. {_Calliope_ | 2 |N. Asia, Himalayas, |Centl. India
{ | | China | (? migrant)
25. {_Erithacus_ | 3 |Atlantic Islands to |
{ | | Japan |
26. {_Grandala_ | 1 |High Himalayas and |
| | E. Thibet |
27. { Ruticilla | 10 |Eu. to Japan, N. Afr., |Abyssinia, India
{ | | Himalayas |
28. { Larvivora | 2 |E. Thibet, Amoor, Japan|Oriental
29. Dromolæa | 3 |S. Europe, N. Africa, |Ethiopian
| | Palestine |
30. Saxicola | 10 |The whole region |E. and S. Africa,
| | | India
31. Cercomela | 2 |Palestine (a desert |N. E. Africa, N. W.
| | genus) | India
32. Pratincola | 3 |W. Europe, N. Africa to|Ethiopian to Oriental
| | India |
33. _Accentor_ | 12 |W. Europe to Japan; |Himalayas(?) in winter
| | high Himalayas |
| | |
TIMALIIDÆ. | | |
| | |
34. _Pterorhinus_ | 3 |Thibet and N. W. China |
(Malacocercus | 1 |Palestine) |Oriental genus
(Crateropus | 2 |N. Africa, Persia) |Ethiopian genus
(Trochalopteron | 3 |E. Thibet) |Oriental genus
(Ianthocincla | 3 |E. Thibet) |Oriental genus
| | |
PANURIDÆ. | | |
| | |
(Paradoxornis | 3 |Himalayas and |(?)Oriental genus
| | E. Thibet) |
35. _Conostoma_ | 1 |High Himalayas |
| | E. Thibet) |
36. Suthora | 3 |E. Thibet |Himalayas, China,
| | | Formosa
37. _Panurus_ | 1 |W. Europe to W. Siberia|
38. _Heteromorpha_ | 1 |Nepaul and E. Thibet, |
| | from 10,000 feet |
| | altitude |
39. _Cholornis_ | 1 |E. Thibet |
| | |
CINCLIDÆ. | | |
| | |
40. Cinclus | 5 |The whole region |American highlands
| | (Atlantic Islands |
| | excluded) |
(Myiophonus | 1 |Turkestan, Thian-Shan |Oriental genus
| | Mountains, 6,000 feet|
| | |
TROGLODYTIDÆ. | | |
| | |
41. Troglodytes | 3 |Iceland and Britain to |Neotropical and
| | Japan | Nearctic, Himalayas
(Pnoepyga | 2 |E. Thibet) |Oriental genus
| | |
CERTHIIDÆ. | | |
| | |
42. Certhia | 2 |W. Europe to N. China |Himalayas, Nearctic
43. _Tichodroma_ | 1 |S. Europe to N. China |Abyssinia, Nepaul,
| | | high
| | |
SITTIDÆ. | | |
| | |
44. Sitta | 7 |W. Europe to Himalayas India, Nearctic
| | and Japan |
| | |
PARIDÆ. | | |
| | |
45. Parus | 20 |W. Europe to Kamschatka|Nearctic, Oriental,
| | N. Africa | Ethiopian
46. Lophophanes | 6 |Europe and high |Nearctic
| | Himalayas |
47. _Acredula_ | 6 |W. Europe to N. China |
| | and Kamschatka |
48. Ægithalus | 1 |S. E. Europe |Ethiopian
| | |
LIOTRICHIDÆ. | | |
| | |
(Proparus | 4 |Moupin, in E. Thibet) |Oriental genus and
| | | fam.
| | |
PYCNONOTIDÆ. | | |
| | |
49. Microscelis | 1 |Japan |Oriental genus
50. Pycnonotus | 2 |Palestine, N. China, |Oriental and Ethiopian
| | Japan |
| | |
ORIOLIDÆ. | | |
| | |
51. Oriolus | |S. Europe, China |Ethiopian and Oriental
| | |
MUSCICAPIDÆ. | | |
| | |
52. Muscicapa | 2 |W. and Central Europe |Ethiopian.
53. Butalis | 2 |W. Europe to Japan and |E. and S. Africa,
| | China | Moluccas
54. Erythrosterna | 3 |Central Europe to N. |Oriental & Madagascar
| | China and Japan |
(Xanthopygia | 1 |Japan) |Oriental genus
(Eumyias-- | 1 |E. Thibet) |Oriental genus
(Cyanoptila | 1 |Japan and Amoor) |Oriental genus
(Siphia | 1 |Moupin, E. Thibet) |Oriental genus
55. Tchitrea | 2 |N. China and Japan |Ethiopian and Oriental
| | |
LANIIDÆ. | | |
| | |
56. Lanius | 11 |The whole region (excl.|Nearctic, Ethiopian,
| | Atlantic Islands) | Oriental
(Telephonus | 1 |N. Africa) |Ethiopian genus
| | |
CORVIDÆ. | | |
| | |
57. Garrulus | 7 |W. Europe, N. Africa, |Himalayas, Formosa
| | to Japan |
58. Perisoreus | 1 |N. Europe and Siberia |N. America
(Urocissa | 2 |Cashmere, Japan) |Oriental genus
59. _Nucifraga_ | 3 |W. Europe to Japan, |Himalayan pine forests
| | and Himalayas |
60. _Pica_ | 5 |W. Europe to China and |S. China and Formosa
| | Japan | migrants[?]
61. _Cyanopica_ | 2 |Spain, N. E. Asia and |
| | Japan |
62. Corvus | 12 |The whole region |Cosmopolite (excl.
| | | S. Am.)
63. _Fregilus_ | 3 |W. Europe to N. China, |Abyssinian mountains
| | Himalayas |
| | |
NECTARINIIDÆ. | | |
| | |
(Arachnecthra | 1 |Palestine) |Oriental genus
| | |
DICÆIDÆ. | | |
| | |
(Zosterops | 1 |Amoor and Japan) |Ethiop., Orien.,
| | | Austral.
| | |
AMPELIDÆ. | | |
| | |
64. Ampelis | 2 |Northern half of region|North America
| | |
HIRUNDINIDÆ. | | |
| | |
65. Hirundo | 2 |The whole region |Cosmopolite
66. Cotyle | 2 |The whole region (excl.|Nearctic, Ethiop.,
| | Atlan. Is.) | Orien.
67. Chelidon | 3 |The whole region |Oriental
| | |
FRINGILLIDÆ. | | |
| | |
68. Fringilla | 6 |The whole region |Africa
69. _Acanthis_ | 3 |Europe and N. Africa to|
| | Central Asia |
70. _Procarduelis_ | 1 |High Himalayas and |
| | E. Thibet |
71. Chrysomitris | 2 |W. Europe to Japan |N. and S. America
72. _Dryospiza_ | 4 |Atlantic Islands to |
| | Palestine, N. Africa |
73. _Metoponia_ | 1 |N. E. Europe to |
| | W. Himalayas |
74. Chlorospiza | 5 |W. Europe, N. Africa |China, E. Africa
| | to Japan |
75. Passer | 8 |The whole region |Ethiopian, Oriental
76. Montifringilla | 4 |Europe to Cashmere and |
| | Siberia |
77. _Fringillauda_ | 1 |N. W. Himalayas to |
| | E. Thibet, high |
78. Coccothraustes | 3 |W. Europe, High |N. America
| | Himalayas to Japan |
79. _Mycerobas_ | 2 |Central Asia & High |
| | Himalayas |
80. Eophona | 2 |E. Thibet, China, and |China
| | Japan |
81. _Pyrrhula_ | 9 |Azores to Japan, High |Alaska
| | Himalayas |
(Crithagra | 1 |Palestine) |Ethiopian genus
82. Carpodacus | 12 |Cent. Eu. to Japan, |India & China,
| | High Himalayas | N. Amer.
83. _Erythrospiza_ | 4 |N. Africa to |
| | Afghanistan and |
| | Turkestan |
84. _Uragus_ | 2 |Turkestan & E. Thibet |
| | to Japan |
85. Loxia | 3 |Europe, High Himalayas |N. America
| | to Japan |
86. Pinicola | 1 |N. Europe, Siberia |N. America
87. _Propyrrhula_ | 1 |High Himalayas |Darjeeling in winter
88. _Pyrrhospiza_ | 1 |Snowy Himalayas |
89. Linota | 6 |The whole region |N. America
90. Leucosticte | 4 |Turkestan to Kamschatka|N. W. America
| | |
Emberizinæ | | |
| | |
91. {Euspiza | 4 |E. Europe to Japan |N. America
92. {_Emberiza_ | 25 |Europe to Japan |N. India, China
93. {Fringillaria | 2 |S. Europe, N. Africa |African genus
94. {Plectrophanes | 2 |Northern half of region|N. America
| | |
STURNIDÆ. | | |
| | |
95. Pastor | 1 |East Europe, Central |India
| | Asia |
96. Sturnia | 2 |Amoor, Japan, N. China |Oriental
97. Sturnus | 3 |The whole region (excl.|India, China
| | Atlantic Islands) |
(Amydrus | 1 |Palestine) |N. E. African genus
98. _Podoces_ | 3 |Cen. Asia, Turkestan, |
| | Yarkand |
| | |
ALAUDIDÆ. | | |
| | |
99. Otocorys | 6 |N. Europe to Japan, |India, N. America,
| | N. Africa, Arabia | Andes
100. Alauda | 7 |The whole region (excl.|India, Africa
| | Iceland) |
101. Galerita | 2 |Central Europe to |India, Central Africa
| | N. China, N. Africa |
102. Calandrella | 4 |Central Europe to |India
| | N. China, N. Africa |
103. _Melanocorypha_| 5 |S. Eu., N. Africa, |N. W. India
| | N. & Cen. Asia |
104. _Pallasia_ | 1 |Mongolia |
(Certhilauda | 1 |N. Africa) |S. African genus
(Alaemon | 1 |N. Africa, Arabia) |Ethiopian genus
105. Ammomanes | 3 |S. Europe, N. Africa, |Africa, India
| | to Cashmere |
| | |
MOTACILLIDÆ. | | |
| | |
106. Motacilla | 6 |The whole region |Oriental, Ethiopian
107. Budytes | 4 |Europe to China |Oriental, Moluccas
108. Calobates | 2 |Atlantic Is., W. |Malaisia, Madagascar
| | Europe, to China |
| | |
PITTIDÆ. | | |
| | |
(Pitta | 1 |Japan) |Oriental & Austral.
| | | genus
| | |
PICARIÆ. | | |
PICIDÆ. | | |
| | |
109. Picoides | 3 |N. and Cen. Europe to |North America
| | Thibet & E. Asia |
110. Picus | 16 |The whole region (excl.|India, China, N. and
| | Atlantic Islands) | S. America
111. Hypopicus | 1 |N. China |Himalayas
(Yungipicus | 1 |N. China) |Oriental genus
112. Dryocopus | 1 |N. & Cen. Europe to |Neotropical
| | N. China |
113. Gecinus | 6 |W. Europe to Thibet, |Oriental
| | Amoor & Japan |
| | |
YUNGIDÆ. | | |
| | |
114. Yunx | 2 |W. Europe to N. W. |N. E. Africa,
| | India, Thibet and | S. Africa
| | Japan |
| | |
CUCULIDÆ. | | |
| | |
115. Cuculus | 2 |The whole region (excl.|Ethiop., Oriental,
| | Atlantic Islands) | Austral.
116. Coccystes | 1 |S. Europe and N. Africa|Ethiopian and Oriental
| | |
CORACIIDÆ. | | |
| | |
117. Coracias | 1 |Cent. Europe to Cent. |Ethiopian, Oriental
| | Asia |
(Eurystomus | 1 |Amoor in summer) |Oriental & Austral.
| | | genus
| | |
MEROPIDÆ. | | |
| | |
118. Merops | 2 |S. Europe to Cashmere, |Ethiopian and Oriental
| | N. Africa |
| | |
ALCEDINIDÆ. | | |
| | |
(Halcyon | 3 |W. Asia, N. China, |Ethiop., Orien.,
| | Japan) | Austral.
119. Alcedo | 2 |Europe, N. China |
120. Ceryle | 2 |S. E. Europe, Japan |Africa, India, America
| | |
UPUPIDÆ. | | |
| | |
121. Upupa | 1 |S. Europe, N. China |Ethiop. & Oriental
| | | genus
| | |
CAPRIMULGIDÆ. | | |
| | |
122. Caprimulgus | 5 |Europe to Japan |Ethiopian and Oriental
| | |
CYPSELIDÆ. | | |
| | |
123. Cypselus | 4 |The whole region (excl.|Ethiopian, America
| | Iceland) |
124. Chætura | 2 |N. China, Dauria |Africa, India
| | |
COLUMBÆ. | | |
COLUMBIDÆ. | | |
| | |
125. Columba | 6 |The whole region |Africa, Asia, America
126. Turtur | 4 |W. Europe to Japan |Ethiopian and Oriental
(Alsæcomus | 1 |E. Thibet) |Oriental genus
| | |
GALLINÆ. | | |
PTEROCLIDÆ. | | |
| | |
127. Pterocles | 2 |S. Europe, N. Africa, |Ethiopian genus
| | to W. India |
128. _Syrrhaptes_ | 2 |Central Asia, N. China |
| | |
TETRAONIDÆ. | | |
| | |
129. Francolinus | 1 |Borders of |Ethiopian, Oriental
| | Mediterranean |
130. _Perdix_ | 2 |Europe to Mongolia |
131. Coturnix | 1 |Central and S. Europe |Ethiop., Orien.,
| | to Japan | Austral.
132. _Lerwa_ | 1 |Snowy Himalayas to |
| | E. Thibet |
133. _Caccabis_ | 5 |Cen. Europe and N. |Abyssinia, Arabia
| | Africa to N. W. |
| | Himalayas |
134. _Tetraogallus_ | 4 |Caucasus to E. Thibet |
| | and Altai Mountains |
135. Tetrao | 4 |Europe and N. Asia |N. America
136. Bonasa | 1 |Europe and N. Asia |N. America
137. Lagopus | 4 |Iceland, W. Europe to |N. America, Greenland
| | Japan |
| | |
PHASIANIDÆ. | | |
| | |
138. _Crossoptilon_ | 4 |Thibet, Mongolia, |
| | N. China |
139. _Lophophorus_ | 3 |Cashmere to E. Thibet |
| | (highest woods) |
140. Tetraophasis | 1 |E. Thibet |E. Thibet(?)
141. Ceriornis | 1 |N. W. Himalayas (high) |Himalayas to W. China
142. Pucrasia-- | 3 |N. W. Himalayas to |Himalayas
| | N. W. China |
143. _Phasianus_ | 10 |Western Asia to Japan |W. Himalayas, Formosa
144. _Thaumalea_ | 3 |E. Thibet to Amoor, |West China
| | N. China |
145. _Ithaginis_ | 2 |Nepaul to E. Thibet |
| | (high) |
| | |
TURNICIDÆ. | | |
| | |
146. Turnix | 2 |Spain and N. Africa, |Ethiop., Orien.,
| | N. China | Austral.
| | |
ACCIPITRES. | | |
VULTURIDÆ. | | |
| | |
147. _Vultur_ | 1 |Spain and N. Africa to |
| | N. China |
148. Gyps | 1 |S. Europe, Palestine, |E. Africa, India
| | Cen. Asia |
149. Otogyps | 1 |S. Europe, N. Africa |S. Africa, India
150. Neophron | 1 |Atlantic Isds. to |Africa, India
| | Palestine |
| | |
FALCONIDÆ. | | |
| | |
151. Circus | 5 |Europe to Japan |Almost Cosmopolite
152. Astur | 1 |Europe to N. China |Almost Cosmopolite
153. Accipiter | 2 |Europe to Japan |Almost Cosmopolite
154. Buteo | 4 |Europe to Japan |Cosmopolite (excl.
| | | Australia)
155. Archibuteo | 1 |N. Europe to Japan |N. America
156. Gypaetus | 1 |S. Europe, N. Africa |Abyssinia, Himalayas
157. Aquila | 5 |Europe to Japan |Nearctic, Ethiop.,
| | | Orien.
158. Nisaetus | 2 |E. Europe, N. Africa, |India, Australia
| | W. Asia |
159. Circaetus | 1 |E. and S. Europe, N. |Africa, India
| | Africa, W. Asia |
160. Haliæetus | 3 |Iceland and S. Europe |Cosmopolite (excl.
| | to Japan | Neotropical region)
161. Milvus | 4 |Europe to Japan, |The Old World &
| | N. Africa | Austral.
162. Elanus | 2 |N. Africa, N. China to |Cosmopolite (excl.
| | Amoor | East U. S.)
163. Pernis | 1 |Europe to Japan |Ethiopian and Oriental
164. Falco | 5 |The whole region |Cosmopolite (excl.
| | | Pacific Islands)
165. Hierofalco | 5 |The whole region |N. America
166. Cerchneis | 4 |Atlantic Islands to |Cosmop. (excl.
| | Japan | Oceania)
| | |
PANDIONIDÆ. | | |
| | |
167. Pandion | 1 |Europe to Japan |Cosmopolite
| | |
STRIGIDÆ. | | |
| | |
168. Surnia | 1 |N. Europe and Siberia |North America
169. Nyctea | 1 |Arctic regions |Arctic America
170. Athene | 4 |Central and S. Europe |Ethiop., Orien.,
| | to Japan | Austral.
(Ninox | 1 |N. China and Japan) |Oriental genus
171. Glaucidium | 1 |Europe to N. China |America
172. Bubo | 2 |Europe to N. China |Africa, India,
| | | America
173. Scops | 3 |S. Europe to Japan |African, Orien.,
| | | Austral.
174. Syrnium | 5 |Europe to Japan |African, Oriental,
| | | Amer.
175. Otus | 2 |Europe to Japan |Almost Cosmopolite
176. Nyctala | 1 |N. Europe to E. Siberia|N. America
177. Strix | 1 |Europe and N. Africa |All warm & temp.
| | | regions
_Peculiar or very characteristic Genera of Wading and Swimming Birds._
GRALLÆ. | | |
RALLIDÆ. | | |
| | |
_Ortygometra_ | 8 |Europe, N. E. Africa |
| | |
SCOLOPACIDÆ. | | |
| | |
_Ibidorhyncha_ | 1 |Cashmere & Cen. Asia, |Himalayan Valleys
| | N. China |
Terekia | 1 |N. E. Europe and |India, Australia
| | Siberia | (migrant)
_Helodromas_ | 1 |E. and N. Europe, |
| | N. India |
_Machetes_ | 1 |N. and Cen. Europe, |India in winter
| | Cen. Asia |
_Eurinorhynchus_ | 1 |N. E. Asia |Bengal
| | |
GLAREOLIDÆ. | | |
| | |
_Pluvianus_ | 1 |N. Africa, Spain |
| | |
CHARADRIIDÆ. | | |
| | |
Vanellus | 8 |Europe to the Punjaub |S. America
| | |
OTIDIDÆ. | | |
| | |
_Otis_ | 2 |W. Europe to Mongolia, |
| | N. Africa |
ANSERES. | | |
ANATIDÆ. | | |
| | |
Aix | 1 |N. China to Amoor |N. America
Bucephala | 3 |Iceland, N. Europe, and|N. America
| | Asia |
Histrionicus | 1 |Iceland, N. Siberia |N. America
Harelda | 1 |North of whole region |Arctic America
Somateria | 3 |North of whole region |N. America
Oedemia | 3 |North of whole region |N. America
| | |
LARIDÆ. | | |
| | |
Rissa | 1 |North coasts of whole |N. America
| | region |
| | |
COLYMBIDÆ. | | |
| | |
Colymbus | 3 |North of whole region |N. America
| | |
ALCIDÆ. | | |
| | |
Alca | 2 |North coasts of whole |N. America
| | region |
Fratercula | 3 |North coasts of whole |N. America
| | region |
Uria | 3 |North coasts of whole |N. America
| | region |
Mergulus | 1 |Iceland and Arctic |Arctic America
| | coasts |
--------------------+-----+-----------------------+----------------------
[Illustration: ETHIOPIAN REGION]
{251}CHAPTER XI.
THE ETHIOPIAN REGION.
This is one of the best defined of the great zoological regions, consisting
of tropical and South Africa, to which must be added tropical Arabia,
Madagascar, and a few other islands, all popularly known as African. Some
naturalists would extend the region northwards to the Atlas Mountains and
include the whole of the Sahara; but the animal life of the northern part
of that great desert seems more akin to the Palæarctic fauna of North
Africa. The Sahara is really a debatable land which has been peopled from
both regions; and until we know more of the natural history of the great
plateaus which rise like islands in the waste of sand, it will be safer to
make the provisional boundary line at or near the tropic, thus giving the
northern half to the Palæarctic, the southern to the Ethiopian region. The
same line may be continued across Arabia.
With our present imperfect knowledge of the interior of Africa, only three
great continental sub-regions can be well defined. The open pasture lands
of interior tropical Africa are wonderfully uniform in their productions; a
great number of species ranging from Senegal to Abyssinia and thence to the
Zambesi, while almost all the commoner African genera extend over the whole
of this area. Almost all this extensive tract of country is a moderately
elevated plateau, with a hot and dry climate, and characterised by a grassy
vegetation interspersed with patches of forest. This forms our first or
East African sub-region. The whole of the west coast from the south side of
the Gambia River to about 10° or 12° south latitude, is a very
{252}different kind of country; being almost wholly dense forests where not
cleared by man, and having the hot moist uniform climate, and perennial
luxuriance of vegetation, which characterise the great equatorial belt of
forest all round the globe. This forest country extends to an unknown
distance inland, but it was found, with its features well marked, by Dr.
Schweinfurth directly he crossed the south-western watershed of the Nile;
and far to the south we find it again unmistakably indicated, in the
excessively moist forest country about the head waters of the Congo, where
the heroic Livingstone met his death. In this forest district many of the
more remarkable African types are alone found, and its productions
occasionally present us with curious similarities to those of the far
removed South American or Malayan forests. This is our second or West
African sub-region.
Extra-tropical South Africa possesses features of its own, quite distinct
from those of both the preceding regions (although it has also much in
common with the first). Its vegetation is known to be one of the richest,
most peculiar, and most remarkable on the globe; and in its zoology it has
a speciality, similar in kind but less in degree, which renders it both
natural and convenient to separate it as our third, or South African
sub-region. Its limits are not very clearly ascertained, but it is probably
bounded by the Kalahari desert on the north-west, and by the Limpopo
Valley, or the mountain range beyond, on the north-east, although some of
its peculiar forms extend to Mozambique. There remains the great Island of
Madagascar, one of the most isolated and most interesting on the globe, as
regards its animal productions; and to this must be added, the smaller
islands of Bourbon, Mauritius and Rodriguez, the Seychelles and the Comoro
Islands, forming together the Mascarene Islands,--the whole constituting
our fourth sub-region.
_Zoological Characteristics of the Ethiopian Region._--We have now to
consider briefly, what are the peculiarities and characteristics of the
Ethiopian Region as a whole,--those which give it its distinctive features
and broadly separate it from the other primary zoological regions.
{253}_Mammalia._--This region has 9 peculiar families of mammalia.
Chiromyidæ (containing the aye-aye); Potamogalidæ and Chrysochloridæ
(Insectivora); Cryptoproctidæ and Protelidae (Carnivora); Hippopotamidæ and
Camelopardalidæ (Ungulata); and Orycteropodidæ (Edentata). Besides these it
possesses 7 peculiar genera of apes, _Troglodytes_, _Colobus_,
_Myiopithecus_, _Cercopithecus_, _Cercocebus_, _Theropithecus_, and
_Cynocephalus_; 2 sub-families of lemurs containing 6 genera, confined to
Madagascar, with 3 genera of two other sub-families confined to the
continent; of Insectivora a family, Centetidæ, with 5 genera, peculiar to
Madagascar, and the genera _Petrodromus_ and _Rhynchocyon_ belonging to the
Macroscelididæ, or elephant-shrews, restricted to the continent; numerous
peculiar genera or sub-genera of civets; _Lycaon_ and _Megalotis_,
remarkable genera of Canidæ; _Ictonyx_, the zorilla, a genus allied to the
weasels; 13 peculiar genera of Muridæ; _Pectinator_, a genus of the South
American family Octodontidæ; and 2 genera of the South American Echimyidæ
or spiny rats. Of abundant and characteristic groups it possesses
_Macroscelides_, _Felis_, _Hyæna_, _Hyrax_, _Rhinoceros_, and _Elephas_, as
well as several species of zebra and a great variety of antelopes.
The great speciality indicated by these numerous peculiar families and
genera, is still farther increased by the absence of certain groups
dominant in the Old-World continent, an absence which we can only account
for by the persistence, through long epochs, of barriers isolating the
greater part of Africa from the rest of the world. These groups are,
Ursidæ, the bears; Talpidæ the moles; Camelidæ, the camels; Cervidæ, the
deer; Caprinæ, the goats and sheep; and the genera _Bos_ (wild ox); and
_Sus_ (wild boar). Combining these striking deficiencies, with the no less
striking peculiarities above enumerated, it seems hardly possible to have a
region more sharply divided from the rest of the globe than this is, by its
whole assemblage of mammalia.
_Birds._--In birds the Ethiopian region is by no means so strikingly
peculiar, many of these having been able to pass the ancient barriers which
so long limited the range of mammalia. {254}It is, however, sufficiently
rich, possessing 54 families of land birds, besides a few genera whose
position is not well ascertained, and which may constitute distinct
families. Of these 6 are peculiar, Musophagidæ (the plantain eaters);
Coliidæ (the colies); Leptosomidæ, allied to the cuckoos; Irrisoridæ,
allied to the hoopoes; and Serpentaridæ, allied to the hawks. Only one
Passerine family is peculiar--Paictidæ, while most of the other tropical
regions possess several; but _Euryceros_ and _Buphaga_, here classed with
the Sturnidæ, ought, perhaps, to form two more. It has, however, many
peculiar genera, especially among the fruit-thrushes, Pycnonotidæ;
flycatchers, Muscicapidæ; shrikes, Lanidæ; crows, Corvidæ; starlings,
Sturnidæ; and weaver-birds, Ploceidæ; the latter family being very
characteristic of the region. It is also rich in barbets, Megalæmidæ (7
peculiar genera); cuckoos, Cuculidæ; rollers, Coraciidæ; bee-eaters,
Meropidæ; hornbills, Bucerotidæ; and goat-suckers, Caprimulgidæ. It is poor
in parrots and rather so in pigeons; but it abounds in _Pterocles_ and
_Francolinus_, genera of Gallinæ, and possesses 4 genera of the peculiar
group of the guinea-fowls, forming part of the pheasant family. It abounds
in vultures, eagles, and other birds of prey, among which is the anomalous
genus _Serpentarius_, the secretary-bird, constituting a distinct family.
Many of the most remarkable forms are confined to Madagascar and the
adjacent islands, and will be noticed in our account of that sub-region.
_Reptiles._--Of the reptiles there are 4 peculiar Ethiopian families;--3 of
snakes, Rachiodontidæ, Dendraspidæ, and Atractaspidæ and 1 of lizards,
Chamæsauridæ.
Psammophidæ (desert snakes) are abundant, as are Lycodontidæ (fanged
ground-snakes), and Viperidæ (vipers). The following genera of snakes are
peculiar or highly characteristic:--_Leptorhynchus_, _Rhamnophis_,
_Herpetethiops_ and _Grayia_ (Colubridæ); _Hopsidrophis_ and _Bucephalus_
(Dendrophidæ); _Langalia_ (Dryophidæ); _Pythonodipsas_ (Dipsadidæ);
_Boedon_, _Lycophidion_, _Holuropholis_, _Simocephalus_ and _Lamprophis_
(Lycodontidæ); _Hortulia_ and _Sanzinia_ (Pythonidæ); _Cyrptophis_,
_Elapsoidea_ and _Poecilophis_ (Elapidæ); and _Atheris_ (Viperidæ). The
following genera {255}of lizards are the most
characteristic:--_Monotrophis_ (Lepidosternidæ); _Cordylus_,
_Pseudocordylus_, _Platysaurus_, _Cordylosaurus_, _Pleurostichus_,
_Saurophis_ and _Zonurus_ (Zonuridæ); _Sphænops_, _Scelotes_,
_Sphænocephalus_ and _Sepsina_ (Sepidæ); _Pachydactylus_ (Geckotidæ);
_Agama_ (Agamidæ); and _Chameleon_ (Chameleonidæ). Of tortoises, _Cynyxis_,
_Pyxis_ and _Chersina_ (Testudinidæ), and _Cycloderma_ (Trionychidæ) are
the most characteristic.
_Amphibia._--Of the 9 families of amphibia there is only 1 peculiar, the
Dactylethridæ, a group of toads; but the Alytidæ, a family of frogs, are
abundant.
_Fresh-water Fish._--Of the 14 families of fresh-water fishes 3 are
peculiar: Mormyridæ and Gymnarchidæ, small groups not far removed from the
pikes; and Polypteridæ, a small group of ganoid fishes allied to the
gar-pikes (Lepidosteidæ) of North America.
_Summary of Ethiopian Vertebrates._--Combining the results here indicated
and set forth in greater detail in the tables of distribution, we find that
the Ethiopian region possesses examples of 44 families of mammalia, 72 of
birds, 35 of reptiles, 9 of amphibia, and 15 of fresh-water fishes. It has
23 (or perhaps 25) families of Vertebrata altogether peculiar to it out of
a total of 175 families, or almost exactly one-eighth of the whole. Out of
142 genera of mammalia found within the region, 90 are peculiar to it; a
proportion not much short of two-thirds. Of land birds there are 294
genera, of which 179 are peculiar; giving a proportion of a little less
than three-fifths.
Compared with the Oriental region this shows a considerably larger amount
of speciality under all the heads; but the superiority is mainly due to the
wonderful and isolated fauna of Madagascar, to which the Oriental region
has nothing comparable. Without this the regions would be nearly equal.
_Insects: Lepidoptera._--11 out of the 16 families of butterflies have
representatives in Africa, but none are peculiar. Acræidæ is one of the
most characteristic families, and there {256}are many interesting forms of
Nymphalidæ, Lycænidæ, and Papilionidæ. The peculiar or characteristic forms
are _Amauris_ (Danaidæ); _Gnophodes_, _Leptoneura_, _Bicyclus_,
_Heteropsis_ and _Coenyra_ (Satyridæ); _Acræa_ (Acræidæ); _Lachnoptera_,
_Precis_, _Salamis_, _Crenis_, _Godartia_, _Amphidema_, _Pseudacræa_,
_Catuna_, _Euryphene_, _Romalæosoma_, _Hamanumida_, _Aterica_, _Harma_,
_Meneris_, _Charaxes_, and _Philognoma_ (Nymphalidæ); _Pentila_, _Liptena_,
_Durbania_, _Zeritis_, _Capys_, _Phytala_, _Epitola_, _Hewitsonia_ and
_Deloneura_ (Lycænidæ); _Pseudopontia_, _Idmais_, _Teracolus_, _Callosune_
(Pieridæ); _Abantis_, _Ceratrichia_ and _Caprona_ (Hesperidæ). The total
number of species known is about 750; which is very poor for an extensive
tropical region, but this is not to be wondered at when the nature of much
of the country is considered. It is also, no doubt, partly due to our
comparative ignorance of the great equatorial forest district, which is the
only part likely to be very productive in this order of insects.
_Coleoptera._--In our first representative family, Cicindelidæ or
tiger-beetles, the Ethiopian region is rather rich, having 13 genera, 11 of
which are peculiar to it; and among these are such remarkable forms as
_Manticora_, _Myrmecoptera_ and _Dromica_; with _Megacephala_, a genus only
found elsewhere in Australia and South America.
In Carabidæ or carnivorous ground beetles, there are about 75 peculiar
genera. Among the most characteristic are _Anthia_, _Polyrhina_,
_Graphipterus_ and _Piezia_, which are almost all peculiar; while
_Orthogonius_, _Hexagonia_, _Macrochilus_, _Thyreopterus_, _Eudema_, and
_Abacetus_ are common to this and the Oriental region; and _Hypolithus_ to
the Neotropical.
Out of 27 genera of Buprestidæ, or metallic beetles, only 6 are peculiar to
the region, one of the most remarkable being _Polybothrus_, confined to
Madagascar. _Sternocera_ and _Chrysochroa_ are characteristic of this
region and the Oriental; it has _Julodis_ in common with the Mediterranean
sub-region, and _Belionota_ with the Malayan.
The region is not rich in Lucanidæ, or stag-beetles, possessing only 10
genera, 7 of which are peculiar, but most of them {257}consist of single
species. The other three genera, _Cladognathus_, _Nigidius_, and _Figulus_,
are the most characteristic, though all have a tolerably wide range in the
Old World.
In the elegant Cetoniidæ, or rose-chafers, this region stands preeminent,
possessing 76 genera, 64 of which are peculiar to it. The others are
chiefly Oriental, except _Oxythræa_ which is European, and _Stethodesma_
which is Neotropical. Preeminent in size and beauty is _Goliathus_,
comprising perhaps the most bulky of all highly-coloured beetles. Other
large and characteristic genera are _Ceratorhina_, _Ischnostoma_,
_Anochilia_, _Diplognatha_, _Agenius_, and many others of less extent.
In the enormous tribe of Longicorns, or long-horned beetles, the Ethiopian
is not so rich as the other three tropical regions; but this may be, in
great part, owing to its more productive districts having never been
explored by any competent entomologists. It nevertheless possesses 262
genera, 216 of which are peculiar, the others being mostly groups of very
wide range. Out of such a large number it is difficult to select a few as
most characteristic, but some of the peculiarities of distribution as
regards other regions may be named. Among Prionidæ, _Tithoes_ is a
characteristic Ethiopian genus. A few species of the American genera
_Parandra_ and _Mallodon_ occur here, while the North Temperate genus
_Prionus_ is only found in Madagascar. Among Cerambycidæ, _Promeces_ is the
most characteristic. The American genera _Oeme_ and _Cyrtomerus_ occur;
while _Homalachnus_ and _Philagathes_ are Malayan, and _Leptocera_ occurs
only in Madagascar, Ceylon, Austro-Malaya, and Australia. The Lamiidæ are
very fine; _Sternotomis_, _Tragocephala_, _Ceroplesis_, _Phryncta_,
_Volumnia_, and _Nitocris_, being very abundant and characteristic. Most of
the non-peculiar genera of this family are Oriental, but _Spalacopsis_ and
_Acanthoderes_ are American, while _Tetraglenes_ and _Schoenionta_ have
been found only in East and South Africa and in Malaya.
_Terrestrial Mollusca_.--In the extensive family of the Helicidæ or snails,
13 genera are represented, only one of which, _Columna_, is peculiar. This
region is however the metropolis of _Achatina_, some of the species being
the largest land-shells {258}known. _Buliminus_, _Stenogyra_, and _Pupa_
are characteristic genera. _Bulimus_ is absent, though one species inhabits
St. Helena. The operculated shells are not very well represented, the great
family of Cyclostomidæ having here only nine genera, with but one peculiar,
_Lithidion_, found in Madagascar, Socotra, and Arabia. None of the genera
appear to be well represented throughout the region, and they are almost or
quite absent from West Africa.
According to Woodward's _Manual_ (1868) West Africa has about 200 species
of land-shells, South Africa about 100, Madagascar nearly 100, Mauritius
about 50. All the islands have their peculiar species; and are, in
proportion to their extent, much richer than the continent; as is usually
the case.
THE ETHIOPIAN SUB-REGIONS.
It has been already explained that these are to some extent provisional;
yet it is believed that they represent generally the primary natural
divisions of the region, however they may be subdivided when our knowledge
of their productions becomes more accurate.
_I. The East African Sub-region, or Central and East Africa._
This division includes all the open country of tropical Africa south of the
Sahara, as well as an undefined southern margin of that great desert. With
the exception of a narrow strip along the east coast and the valleys of the
Niger and Nile, it is a vast elevated plateau from 1,000 to 4,000 feet
high, hilly rather than mountainous, except the lofty table land of
Abyssinia, with mountains rising to 16,000 feet and extending south to the
equator, where it terminates in the peaks of Kenia and Kilimandjaro, 18,000
and 20,000 feet high. The northern portion of this sub-region is a belt
about 300 miles wide between the Sahara on the north and the great
equatorial forest on the south, extending from Cape Verd, the extreme
western point of Africa, across the northern bend of the Niger and Lake
Tchad to the mountains of Abyssinia. The greater part of this tract has a
{259}moderate elevation. The eastern portion reaches from about the second
cataract of the Nile, or perhaps from about the parallel of 20° N.
Latitude, down to about 20° S. Latitude, and from the east coast to where
the great forest region commences, or to Lake Tanganyika and about the
meridian of 28° to 30° E. Longitude. The greater part of this tract is a
lofty plateau.
The surface of all this sub-region is generally open, covered with a
vegetation of high grasses or thorny shrubs, with scattered trees and
isolated patches of forest in favourable situations. The only parts where
extensive continuous forests occur, are on the eastern and western slopes
of the great Abyssinian plateau, and on the Mozambique coast from Zanzibar
to Sofala. The whole of this great district has one general zoological
character. Many species range from Senegal to Abyssinia, others from
Abyssinia to the Zambesi, and a few, as _Mungos fasciatus_ and
_Phacochoerus æthiopicus_, range over the entire sub-region. _Fennecus_,
_Ictonyx_, and several genera of antelopes, characterise every part of it,
as do many genera of birds. _Coracias nævia_, _Corythornis cyanostigma_,
_Tockus nasutus_, _T. erythrorhynchus_, _Parus leucopterus_, _Buphaga
africana_, _Vidua paradisea_, are examples of _species_, which are found in
the Gambia, Abyssinia and South East Africa, but not in the West African
sub-region; and considering how very little is known of the natural history
of the country immediately south of the Sahara, it may well be supposed
that these are only a small portion of the species really common to the
whole area in question, and which prove its fundamental unity.
Although this sub-region is so extensive and so generally uniform in
physical features, it is by far the least peculiar part of Africa. It
possesses, of course, all those wide-spread Ethiopian types which inhabit
every part of the region, but it has hardly any special features of its
own. The few genera which are peculiar to it have generally a limited
range, and for the most part belong, either to the isolated
mountain-plateau of Abyssinia which is almost as much Palæarctic as
Ethiopian, or to the woody districts of Mozambique where the fauna has more
of a West or South African character.
{260}_Mammalia._--The only forms of Mammalia peculiar to this sub-region
are _Theropithecus_, one of the Cynopithecidæ confined to Abyssinia;
_Petrodromus_ and _Rhynchocyon_, belonging to the insectivorous
Macroscelididæ, have only been found in Mozambique; the Antelopine genus
_Neotragus_, from Abyssinia southward; _Saccostomus_ and _Pelomys_ genera
of Muridæ inhabiting Mozambique; _Heterocephalus_ from Abyssinia, and
_Heliophobius_ from Mozambique, belonging to the Spalacidæ; and
_Pectinator_ from Abyssinia, belonging to the Octodontidæ. _Cynocephalus_,
_Rhinoceros_, _Camelopardalis_, and antelopes of the genera _Oryx_,
_Cervicapra_, _Kobus_, _Nanotragus_, _Cephalophus_, _Hippotragus_,
_Alcephalus_, and _Catoblepas_, are characteristic; as well as _Felis_,
_Hyæna_, and numerous civets and ichneumons.
_Birds._--Peculiar forms of birds are hardly to be found here; we only meet
with two--_Hypocolius_, a genus of shrikes in Abyssinia; and _Balæniceps_,
the great boat-billed heron of the Upper Nile. Yet throughout the country
birds are abundant, and most of the typical Ethiopian forms are well
represented.
_Reptiles._--Of reptiles, the only peculiar forms recorded are
_Xenocalamus_, a genus of snakes, belonging to the Calamariidæ; and
_Pythonodipsas_, one of the Dipsadidæ, both from the Zambesi; and among
lizards, _Pisturus_, one of the Geckotidæ, from Abyssinia.
_Amphibia and Fishes._--There are no peculiar forms of amphibia or of
fresh-water fishes.
_Insects._--Insects are almost equally unproductive of peculiar forms.
Among butterflies we have _Abantis_, one of the Hesperidæ, from Mozambique;
and in Coleoptera, 2 genera of Cicindelidæ, 8 of Carabidæ, 1 or 2 of
Cetoniidæ, and about half-a-dozen of Longicorns: a mere nothing, as we
shall see, compared with the hosts of peculiar genera that characterise
each of the other sub-regions. Neither do land-shells appear to present any
peculiar forms.
The fact that so very few special types characterise the extensive area now
under consideration is very noteworthy. It justifies us in uniting this
large and widespread tract of country as forming essentially but one
sub-division of the great Ethiopian region, and it suggests some curious
speculations as to the former history of that region, a subject which must
be deferred to the latter part of this chapter. In none of the other great
tropical regions does it occur, that the largest portion of their area,
although swarming with life, yet possesses hardly any distinctive features
except the absence of numerous types characteristic of the other
sub-regions.
Plate IV.
[Illustration]
CHARACTERISTIC ANIMALS OF EAST AFRICA.
{261}_Plate IV._--_Illustrating the Zoology of East Africa._--Although this
sub-region has so little speciality, it is that which abounds most in large
animals, and is, perhaps, the best representative of Africa as regards
zoology. Some of the most distinctive of African animals range over the
whole of it, and as, from recent explorations, many parts of this wide area
have been made known to the reading public, we devote one of our plates to
illustrate the especially African forms of life that here abound. The
antelopes represented are the koodoo (_Tragelaphus strepsiceros_) one of
the handsomest of the family, which ranges over all the highlands of Africa
from Abyssinia to the southern districts. To the left is the aardvark, or
earth pig, of North Eastern Africa (_Orycteropus æthiopicus_) which, to the
north of the equator in East Africa, represents the allied species of the
Cape of Good Hope. These Edentata are probably remnants of the ancient
fauna of Africa, when it was completely isolated from the northern
continents and few of the higher types had been introduced. The large bird
in the foreground is the secretary-bird, or serpent-killer (_Serpentarius
reptilivorus_), which has affinities both for the birds-of-prey and the
waders. It is common over almost all the open country of Africa, destroying
and feeding on the most venomous serpents. The bird on the wing is the
red-billed promerops (_Irrisor erythrorhynchus_), a handsome bird with
glossy plumage and coral-red bill. It is allied to the hoopoes, and feeds
on insects which it hunts for among the branches of trees. This species
also ranges over a large part of east and central Africa to near the Cape
of Good Hope. Other species are found in the west; and the genus, which
forms a distinct family, _Irrisoridæ_, is one of the best marked Ethiopian
types of birds. In the distance is a rhinoceros, now one of the
characteristic features of African {262}zoology, though there is reason to
believe that it is a comparatively recent intruder into the country.
_II. The West-African Sub-region._
This may be defined as the equatorial-forest sub-region, since it comprises
all that portion of Africa, from the west coast inland, over which the
great equatorial forests prevail more or less uninterruptedly. These
commence to the south of the Gambia River, and extend eastwards in a line
roughly parallel to the southern margin of the great desert, as far as the
sources of the upper Nile and the mountains forming the western boundary of
the basin of the great lakes; and southward to that high but marshy
forest-country in which Livingstone was travelling at the time of his
death. Its southern limits are undetermined, but are probably somewhere
about the parallel of 11° S. Latitude.[10]
This extensive and luxuriant district has only been explored zoologically
in the neighbourhood of the West coast. Much, no doubt, remains to be done
in the interior, yet its main features are sufficiently well known, and
most of its characteristic types of animal life have, no doubt, been
discovered.
_Mammalia._--Several very important groups of mammals are peculiar to this
sub-region. Most prominent are the great anthropoid apes--the gorilla and
the chimpanzee--forming the genus _Troglodytes_; and monkeys of the genera
_Myiopithecus_ and _Cercocebus_. Two remarkable forms of lemurs,
_Perodicticus_ and _Arctocebus_, are also peculiar to West Africa. Among
the Insectivora is _Potamogale_, a semi-aquatic animal, forming a distinct
family; and three peculiar genera of civets (Viverridæ) have been
described. _Hyomoschus_, a small, deer-like animal, belongs to the
Tragulidæ, or chevrotains, a family otherwise {263}confined to the Oriental
region; and in the squirrel family is a curious genus, _Anomalurus_, which
resembles the flying squirrels of other parts of the world, without being
directly allied to them.
_Birds._--In this class we find a larger proportionate number of peculiar
forms. _Hypergerus_ and _Alethe_, belonging to the Timaliidæ, or babblers,
are perhaps allied to Malayan groups; _Parinia_, a peculiar form of tit, is
found only in Prince's Island; _Ixonotus_ is an abundant and characteristic
form of Pycnonotidæ; _Fraseria_, _Hypodes_, _Cuphopterus_, and
_Chaunonotus_, are peculiar genera of shrikes; _Picathartes_ is one of the
many strange forms of the crow family; _Cinnyricinclus_ is a peculiar genus
of sun-birds; _Pholidornis_ is supposed to belong to the Oriental Dicæidæ,
or flower-peckers; _Waldenia_ is a recently-described new form of swallow;
_Ligurnus_, a finch, _Spermospiga_, a weaver bird, and _Onychognathus_ a
starling, are also peculiar West African genera. Coming to the Picariæ we
have _Verreauxia_, a peculiar woodpecker; three peculiar genera of barbets
(Megalæmidæ); the typical plantain-eaters (Musophaga); _Myioceyx_, a
peculiar genus of kingfishers; while _Berenicornis_ is a genus of crested
hornbills, only found elsewhere in Malaya. The grey parrots, of the genus
_Psittacus_, are confined to this sub-region, as are two peculiar genera of
partridges, and three of guinea-fowl. We have also here a species of
_Pitta_, one of the Oriental family of ground-thrushes; and the Oriental
paroquets, _Palæornis_, are found here as well as in Abyssinia and the
Mascarene Islands.
We thus find, both in the Mammalia and birds of West Africa, a special
Oriental or even Malayan element not present in the other parts of tropical
Africa, although appearing again in Madagascar. In the Mammalia it is
represented by the anthropoid apes; by _Colobus_ allied to _Semnopithecus_,
and by _Cercocebus_ allied to _Macacus_; and especially by a form of the
Malayan family of chevrotains (Tragulidæ). The Malayan genus of otters,
_Aonyx_, is also said to occur in West and South Africa. In birds we have
special Oriental and Malayan affinities in _Alethe_, _Pholidornis_,
_Berenicornis_, _Pitta_, and _Palæornis;_ while the Oriental genus _Treron_
has a wide range in Africa. We shall {264}endeavour to ascertain the
meaning of this special relation at a subsequent stage of our inquiries.
_Plate V._--_River Scene in West Africa, with Characteristic Animals._--Our
artist has here well represented the luxuriance and beauty of a tropical
forest; and the whole scene is such as might be witnessed on the banks of
one of the rivers of equatorial West Africa. On the right we see a red
river-hog (_Potamochoerus penicillatus_), one of the handsomest of the
swine family, and highly characteristic of the West African sub-region. In
a tree overhead is the potto (_Perodicticus potto_), one of the curious
forms of lemur confined to West Africa. On the left is the remarkable
_Potamogale velox_, first discovered by Du Chaillu,--an Insectivorous
animal, with the form and habits of an otter. On the other side of the
river are seen a pair of gorillas (_Troglodytes gorilla_), the largest of
the anthropoid apes.
The bird on the wing is the Whydah finch (_Vidua paradisea_), remarkable
for the enormous plumes with which the tail of the male bird is decorated
during the breeding season. The crested bird overhead is one of the
beautiful green touracos (_Turacus macrorhynchus_), belonging to the
Musophagidæ, or plantain-eaters, a family wholly African, and most abundant
in the western sub-region.
_Reptiles._--In this class we find a large number of peculiar forms; 13
genera of snakes, 3 of lizards, and 2 of tortoises being confined to the
sub-region. The snakes are _Pariaspis_, _Elapops_, and _Prosymna_
(Calamariidæ), _Rhamnophis_, _Herpetethiops_, and _Grayia_ (Colubridæ),
_Neusterophis_ and _Limnophis_ (Homalopsidæ), _Simocephalus_ and
_Holurophis_ (Lycodontidæ); _Pelophilus_ (Pythonidæ); _Elapsoidea_
(Elapidæ); and _Atheris_ (Viperidæ). The lizards are _Dalophia_
(Lepidosternidæ); _Otosaurus_ (Scincidæ); _Psilodactylus_ (Geckotidæ). The
tortoises, _Cinyxis_ (Testudinidæ) and _Tetrathyra_ (Trionichidæ).
_Amphibia._--Of Amphibia, there are 2 peculiar genera of tree-frogs,
_Hylambatis_ and _Hemimantis_, belonging to the Polypedatidæ.
Plate V.
[Illustration]
SCENE IN WEST AFRICA, WITH CHARACTERISTIC ANIMALS.
{265}Here, too, we find some interesting relations with the Oriental region
on the one side, and the Neotropical on the other. The snakes of the family
Homalopsidæ have a wide range, in America, Europe, and all over the
Oriental region, but are confined to West Africa in the Ethiopian region.
_Dryiophis_ (Dryiophidæ) and _Dipsadoboa_ (Dipsadidæ) on the other hand,
are genera of tropical America which occur also in West Africa. The family
of lizards, Acontiadæ, are found in West and South Africa, Ceylon, and the
Moluccas. The family of toads, Engystomidæ, in West and South Africa and
the whole Oriental region; while the Phryniscidæ inhabit tropical Africa
and Java.
_Insects._--We have here a large number of peculiar genera. There are 10 of
butterflies, _Lachnoptera_, _Amphidema_, and _Catuna_ belonging to the
Nymphalidæ, while four others are Lycænidæ. The genus _Euxanthe_ is common
to West Africa and Madagascar.
Of Coleoptera there are 53 peculiar genera; 20 are Carabidæ, 2 Lucanidæ, 12
Cetoniidæ, 3 Prionidæ, 16 Cerambycidæ, and 34 Lamiidæ. Besides these there
are 4 or 5 genera confined to West Africa and Madagascar.
_Land Shells._--West Africa is very rich in land shells, but it does not
appear to possess any well-marked genera, although several of the smaller
groups or sub-genera are confined to it. Helicidæ of the genera _Nanina_,
_Buliminus_ and _Achatina_ are abundant and characteristic.
_Islands of the West African Sub-region._--The islands in the Gulf of
Guinea are, Fernando Po, very near the main land, with Prince's Island and
St. Thomas, considerably further away to the south-west. Fernando Po was
once thought to be a remarkable instance of an island possessing a very
peculiar fauna, although close to the main land and not divided from it by
a deep sea. This, however, was due to our having obtained considerable
collections from Fernando Po, while the opposite coast was almost unknown.
One after another the species supposed to be peculiar have been found on
the continent, till it becomes probable, that, as in the case of other
islands similarly situated, it contains no peculiar species whatever. The
presence of numerous mammalia, among which are baboons, lemurs, _Hyrax_,
and {266}_Anomalurus_, shows that this island has probably once been united
to the continent.
Prince's Island, situated about 100 miles from the coast, has no mammals,
but between 30 and 40 species of birds. Of these 7 are peculiar species,
viz., _Zosterops ficedulina_, _Cuphopterus dohrni_ (a peculiar genus of
Sylviidæ), _Symplectes princeps_, _Crithagra rufilata_, _Columba
chlorophæa_, _Peristera principalis_, and _Strix thomensis_.
In the Island of St. Thomas, situated on the equator about 150 miles from
the coast, there are 6 peculiar species out of 30 known birds, viz., _Scops
leucopsis_, _Zosterops lugubris_, _Turdus olivaceofuscus_, _Oriolus
crassirostris_, _Symplectes sancti-thomæ_ and _Aplopelia simplex_; also
_Strix thomensis_ in common with Prince's Island. The remainder are all
found on the adjacent coasts. It is remarkable that in Prince's Island
there are no birds of prey, any that appear being driven off by the parrots
(_Psittacus erithacus_) that abound there; whereas in St. Thomas and
Fernando Po they are plentiful.
_III. South-African Sub-region._
This is the most peculiar and interesting part of Africa, but owing to the
absence of existing barriers its limits cannot be well defined. The typical
portion of it hardly contains more than the narrow strip of territory
limited by the mountain range which forms the boundary of the Cape Colony
and Natal, while in a wider sense it may be extended to include Mozambique.
It may perhaps be best characterised as bounded by the Kalahari desert and
the Limpopo river. It is in the more limited district of the extreme south,
that the wonderful Cape flora alone exists. Here are more genera and
species, and more peculiar types of plants congregated together, than in
any other part of the globe of equal extent. There are indications of a
somewhat similar richness and specialization in the zoology of this
country; but animals are so much less closely dependent on soil and
climate, that much of the original peculiarity has been obliterated, by
long continued interchange of species with so vast an area as {267}that of
Africa south of the equator. The extreme peculiarity and isolation of the
flora must not, however, be lost sight of, if we would correctly interpret
the phenomena afforded by the distribution of animal life on the African
continent.
_Mammalia._--A much larger number of peculiar forms of mammals are found
here than in any of the other sub-regions, although it is far less in
extent than either of the three divisions of the continent. Among
Insectivora we have the Chrysochloridæ, or golden moles, consisting of two
genera confined to South Africa; while the Macroscelididæ, or elephant
shrews, are also characteristically South African, although ranging as far
as Mozambique and the Zambezi, with one outlying species in North Africa.
The Viverridæ are represented by three peculiar genera, _Ariela_,
_Cynictis_, and _Suricata_. The Carnivora present some remarkable forms:
_Proteles_, forming a distinct family allied to the hyænas and weasels; and
two curious forms of Canidæ--_Megalotis_ (the long-eared fox) and _Lycaon_
(the hyæna-dog), the latter found also in parts of East Africa. _Hydrogale_
is a peculiar form of Mustelidæ; _Pelea_ one of the antelopes; _Dendromys_,
_Malacothrix_, and _Mystromys_ are peculiar genera of the mouse family
(Muridæ); _Bathyerges_ one of the mole-rats (Spalacidæ); _Pedetes_, the
Cape-hare, a remarkable form of jerboa; and _Petromys_, one of the
spiny-rats (Echimyidæ). The remarkable _Orycteropus_, or earth-pig, has one
species in South and one in North East Africa. We have thus eighteen genera
of mammalia almost or quite peculiar to South Africa.
_Birds._--These do not present so many peculiar forms, yet some are very
remarkable. _Chætops_ is an isolated genus of thrushes (Turdidæ).
_Lioptilus_, one of the fruit-thrushes (Pycnonotidæ). _Pogonocichla_, one
of the fly-catchers; _Urolestes_, a shrike; _Promerops_, a sun-bird;
_Philetærus_ and _Chera_, weaver-birds; and three peculiar genera of
larks--_Spizocorys_, _Heterocorys_, and _Tephrocorys_, complete the list of
peculiar types of Passeres. A wood-pecker, _Geocolaptes_, is nearly allied
to a South American genus. The Cape-dove, _Oena_, is confined to South and
East Africa and Madagascar; and _Thalassornis_ is a peculiar form of duck.
Several genera are also confined to West and South {268}Africa;--as
_Phyllastrephus_ (Pycnonotidæ), _Smithornis_ (Muscicapidæ), _Corvinella_
(Laniidæ); _Barbatula_ and _Xylobucco_ (Megalæmidæ); _Ceuthmochares_, also
in Madagascar, (Cuculidæ); _Typanistria_ (Columbidæ). Other remarkable
forms, though widely spread over Africa, appear to have their metropolis
here, as _Colius_ and _Indicator_. Others seem to be confined to South
Africa and Abyssinia, as the curious _Buphaga_ (Sturnidæ); and _Apaloderma_
(Trogonidæ). _Machærhamphus_ (Falconidæ) is found only in South-West
Africa, Madagascar, and the Malay Peninsula.
_Reptiles._--There are 4 peculiar genera of snakes,--_Typhline_, belonging
to the blind burrowing snakes, Typhlopidæ; _Lamprophis_ (Lycodontidæ);
_Cyrtophis_ and _Pæcilophis_ (Elapidæ), a family which is chiefly Oriental
and Australian. Of Lizards there are 10 peculiar genera; _Monotrophis_
(Lepidosternidæ), but with an allied form in Angola; _Cordylus_,
_Pseudocordylus_, _Platysaurus_, _Cordylosaurus_, _Pleurostichus_, and
_Saurophis_, all peculiar genera of Zonuridæ; _Chamæsaura_, forming the
peculiar family Chamæsauridæ; _Colopus_ and _Rhopitropus_ (Geckotidæ).
_Amphibia._--Of Amphibia there are 4 peculiar genera: _Schismaderma_
(Bufonidæ); _Brachymerus_ (Engystomidæ); _Phrynobatrachus_ and
_Stenorhynchus_ (Ranidæ). These last are allied to Oriental genera, and the
only other Engystomidæ are Oriental and Neotropical.
_Fresh-water Fish._--Of fresh-water fishes there is 1
genus--_Abrostomus_--belonging to the carp family, peculiar to South
Africa.
_Insects._--South Africa is excessively rich in insects, and the number of
peculiar types surpasses that of any other part of the region. We can only
here summarize the results.
_Lepidoptera._--Of butterflies there are 7 peculiar genera; 2 belonging to
the Satyridæ, 1 to Acræidæ, 3 to Lycænidæ, and 1 to Hesperidæ. _Zeritis_
(Lycænidæ) is also characteristic of this sub-region, although 1 species
occurs in West Africa.
_Coleoptera._--These are very remarkable. In the family of Cicindelidæ, or
tiger-beetles, we have the extraordinary _Manticora_ and _Platychile_,
forming a sub-family, whose nearest allies are in North America; as well as
_Ophryodera_ and _Dromica_, the latter an extensive genus, which ranges as
far north as Mozambique {269}and Lake Ngami. Another genus of this family,
_Jansenia_, is common to South Africa and South India.
In the large family of Carabidæ, or ground-beetles, there are 17 peculiar
South African genera, the most important being _Crepidogaster_,
_Hytrichopus_, _Arsinoë_, and _Piezia_. Three others--_Eunostus_,
_Glyphodactyla_, and _Megalonychus_--are common to South Africa and
Madagascar only. There is also a genus in common with Java, and one with
Australia.
Of Lucanidæ, or stag-beetles, there are 3 peculiar genera; of Cetoniidæ, or
rose-chafers, 14; and of Buprestidæ, 2.
In the great family of Longicorns there are no less than 67 peculiar
genera--an immense number when we consider that the generally open
character of the country, is such as is not usually well suited to this
group of insects. They consist of 5 peculiar genera of Prionidæ, 25 of
Cerambycidæ, and 37 of Lamiidæ.
_Summary of South-African Zoology._--Summarizing these results, we find
that South Africa possesses 18 peculiar genera of Mammalia, 12 of Birds, 18
of Reptiles, 1 of Fishes, 7 of Butterflies, and 107 of the six typical
families of Coleoptera. Besides this large amount of speciality it contains
many other groups, which extend either to West Africa, to Abyssinia, or to
Madagascar only, a number of which are no doubt to be referred as
originating here. We also find many cases of direct affinity with the
Oriental region, and especially with the Malay districts, and others with
Australia; and there are also less marked indications of a relation to
America.
_Atlantic Islands of the Ethiopian Region._ _St. Helena._--The position of
St. Helena, about 1,000 miles west of Africa and 16° south of the equator,
renders it difficult to place it in either of the sub-regions; and its
scanty fauna has a general rather than any special resemblance to that of
Africa. The entire destruction of its luxuriant native forests by the
introduction of goats which killed all the young trees (a destruction which
was nearly completed two centuries ago) must have led to the extermination
of most of the indigenous birds and insects. At present there is no land
bird that is believed to be really indigenous, and but one {270}wader, a
small plover (_Ægialitis sanctæ-helenæ_) which is peculiar to the island,
but closely allied to African species. Numerous imported birds, such as
canaries, Java sparrows, some African finches, guinea-fowls, and
partridges, are now wild. There are no native butterflies, but a few
introduced species of almost world-wide range. The only important remnant
of the original fauna consists of beetles and land shells. The beetles are
the more numerous and have been critically examined and described by Mr. T.
V. Wollaston, whose researches in the other Atlantic islands are so well
known.
_Coleoptera of St. Helena._--Omitting those beetles which get introduced
everywhere through man's agency, there are 59 species of Coleoptera known
from St. Helena; and even of these there are a few widely distributed
species that may have been introduced by man. It will be well, therefore,
to confine ourselves almost wholly to the species peculiar to the island,
and, therefore, almost certainly forming part of the endemic or original
fauna. Of these we find that 10 belong to genera which have a very wide
range, and thus afford no indication of geographical affinity; 2 belong to
genera which are characteristic of the Palæarctic fauna (_Bembidium_,
_Longitarsus_); 3 to African genera (_Adoretus_, _Sciobius_,
_Aspidomorpha_); and two species of _Calosoma_ are most allied to African
species. There are also 4 African species, which may be indigenous in St.
Helena. The peculiar genera, 7 in number, are, however, the most
interesting. We have first _Haplothorax_, a large beetle allied to
_Carabus_ and _Calosoma_, though of a peculiar type. This may be held to
indicate a remote Palæarctic affinity. _Melissius_, one of the Dynastidæ,
is allied to South African forms. _Microxylobius_, one of the Cossonides (a
sub-family of Curculionidæ) is the most important genus, comprising as it
does 13 species. It is, according to Mr. Wollaston, an altogether peculiar
type, most allied to _Pentarthrum_, a genus found in St. Helena, Ascension,
and the south of England, and itself very isolated. _Nesiotes_, another
genus of Curculionidæ, belongs to a small group, the allied genera forming
which inhabit Europe, Madeira, and Australia. A third peculiar and isolated
genus is _Trachyphlæosoma_. The Anthribidæ are represented by {271}2
genera, _Notioxenus_ and _Homoeodera_, which are altogether peculiar and
isolated, and contain 9 species. Thus no less than 27 species, or more than
half of the undoubtedly indigenous beetles, belong to 5 peculiar and very
remarkable genera of Rhyncophora.
It appears from this enumeration, that the peculiar species as a whole,
exhibit most affinity to the Ethiopian fauna; next to the South European
fauna; and lastly to that of the islands of the North Atlantic; while there
is such a large amount of peculiarity in the most characteristic forms,
that no special geographical affinity can be pointed out.
_Land Shells._--These consist of about a dozen living species, and about as
many extinct found in the surface soil, and probably exterminated by the
destruction of the forests. The genera are _Succinea_, _Zonites_, _Helix_,
_Bulimus_, _Pupa_, and _Achatina_. The _Bulimi_ (all now extinct but one)
comprise one large, and several small species, of a peculiar type, most
resembling forms now inhabiting South America and the islands of the
Pacific. _Zonites_ is chiefly South European, but the other genera are of
wide range, and none are peculiar to the island.
The marine shells are mostly Mediterranean, or West Indian species, with
some found in the Indian Ocean; only 4 or 5 species being peculiar to the
island.
_Tristan d'Acunha._--This small island is situated nearly midway between
the Cape of Good Hope and the mouth of the La Plata, but it is rather
nearer Africa than America, and a little nearer still to St. Helena. An
island so truly oceanic and of whose productions so little is known, cannot
be placed in any region, and is only noticed here because it comes
naturally after St. Helena. It is known to possess three peculiar land
birds. One is a thrush (_Nesocichla eremita_) whose exact affinities are
not determined; the other a small water-hen (_Gallinula nesiotis_) allied
to our native species, but with shorter and softer wings, which the bird
does not use for flight. A finch of the genus _Crithagra_ shows African
affinities; while another recently described as _Nesospiza acunhæ_ (Journ.
für Orn. 1873, p. 154) forms a new genus said to resemble more nearly some
American forms.
{272}The only known land-shells are 2 peculiar species of _Balea,_ a genus
only found elsewhere in Europe and Brazil.
_IV. Madagascar and the Mascarene Islands, or the Malagasy Sub-region._
This insular sub-region is one of the most remarkable zoological districts
on the globe, bearing a similar relation to Africa as the Antilles to
tropical America, or New Zealand to Australia, but possessing a much richer
fauna than either of these, and in some respects a more remarkable one even
than New Zealand. It comprises, besides Madagascar, the islands of
Mauritius, Bourbon, and Rodriguez, the Seychelles and Comoro islands.
Madagascar itself is an island of the first class, being a thousand miles
long and about 250 miles in average width. It lies parallel to the coast of
Africa, near the southern tropic, and is separated by 230 miles of sea from
the nearest part of the continent, although a bank of soundings projecting
from its western coast reduces this distance to about 160 miles. Madagascar
is a mountainous island, and the greater part of the interior consists of
open elevated plateaus; but between these and the coast there intervene
broad belts of luxuriant tropical forests. It is this forest-district which
has yielded most of those remarkable types of animal life which we shall
have to enumerate; and it is probable that many more remain to be
discovered. As all the main features of this sub-region are developed in
Madagascar, we shall first endeavour to give a complete outline of the
fauna of that country, and afterwards show how far the surrounding islands
partake of its peculiarities.
_Mammalia._--The fauna of Madagascar is tolerably rich in genera and
species of mammalia, although these belong to a very limited number of
families and orders. It is especially characterized by its abundance of
Lemuridæ and Insectivora; it also possesses a few peculiar Carnivora of
small size; but most of the other groups in which Africa is especially
rich--apes and monkeys, lions, leopards and hyænas, zebras, giraffes,
antelopes, elephants and rhinoceroses, and even porcupines and squirrels,
are wholly wanting. No less than 40 distinct families of land {273}mammals
are represented on the continent of Africa, only 11 of which occur in
Madagascar, which also possesses 3 families peculiar to itself. The
following is a list of all the genera of Mammalia as yet known to inhabit
the island:--
PRIMATES.
LEMURIDÆ.
Indrisinæ. Species.
_Indris_ 6
Lemurinæ.
_Lemur_ 15
_Hapalemur_ 2
_Microcebus_ 4
_Chirogaleus_ 5
_Lepilemur_ 2
CHIROMYIDÆ.
_Chiromys_ 1
BATS--(Chiroptera).
PTEROPIDÆ.
Pteropus 2
RHINOLOPHIDÆ.
Rhinolophus 1
VESPERTILIONIDÆ.
Vespertilio 1
Taphozous 1
NOCTILIONIDÆ.
Nyctinomus 1
INSECTIVORA.
CENTETIDÆ.
_Centetes_ 2
_Hemicentetes_ 2
_Ericulus_ 2
_Oryzorictes_ 1
_Echinops_ 3
SORICIDÆ.
Sorex 1
CARNIVORA.
CRYPTOPROCTIDÆ.
_Cryptoprocta_ 1
VIVERRIDÆ.
_Fossa_ 2
_Galidia_ 3
_Galidictis_ 2
_Eupleres_ 1
UNGULATA.
SUIDÆ.
Potamochoerus 1
RODENTIA.
MURIDÆ.
_Nesomys_ 1
_Hypogeomys_ 1
_Brachytarsomys_ 1
We have here a total of 12 families, 27 genera, and 65 species of Mammals;
3 of the families and 20 of the genera (indicated by italics) being
peculiar. All the species are peculiar, except perhaps one or two of the
wandering bats. Remains of a _Hippopotamus_ have been found in a sub-fossil
condition, showing that this animal probably inhabited the island at a not
very remote epoch.
The assemblage of animals above noted is remarkable, and seems to indicate
a very ancient connection with the southern portion of Africa, before the
apes, ungulates, and felines had entered it. The lemurs, which are here so
largely developed, are {274}represented by a single group in Africa, with
two peculiar forms on the West coast. They also re-appear under peculiar
and isolated forms in Southern India and Malaya, and are evidently but the
remains of a once wide-spread group, since in Eocene times they inhabited
North America and Europe, and very probably the whole northern hemisphere.
The Insectivora are another group of high antiquity, widely scattered over
the globe under a number of peculiar forms; but in no equally limited area
represented by so many peculiar types as in Madagascar. South and West
Africa are also rich in this order.
The Carnivora of Madagascar are mostly peculiar forms of Viverridæ, or
civets, a family now almost confined to the Ethiopian and Oriental regions,
but which was abundant in Europe during the Miocene period.
The _Potamochoerus_ is a peculiar _species_ only, which may be perhaps
explained by the unusual swimming powers of swine, and the semi-aquatic
habits of this genus, leading to an immigration at a later period than in
the case of the other Mammalia. The same remark will apply to the small
_Hippopotamus_, which was coeval with the great Struthious bird Æpiornis.
Rodents are only represented by three peculiar forms of Muridæ, but it is
probable that others remain to be discovered.
_Birds._--Madagascar is exceedingly rich in birds, and especially in
remarkable forms of Passeres. No less than 88 genera and 111 species of
land-birds have been discovered, and every year some additions are being
made to the list. The African families of Passeres are almost all
represented, only two being absent--Paridæ and Fringillidæ, both very
poorly represented in Africa itself. Among the Picariæ, however, the case
is very different, no less than 7 families being absent, viz.--Picidæ, or
woodpeckers; Indicatoridæ, or honey-guides; Megalæmidæ, or barbets;
Musophagidæ, or plantain-eaters; Coliidæ, or colies; Bucerotidæ, or
hornbills; and Irrisoridæ, or mockers. Three of these are peculiar to
Africa, and all are well represented there, so that their absence from
Madagascar is a very remarkable fact. The number of peculiar genera in
Madagascar constitutes one of the main features of its ornithology, and
many of these are so {275}isolated that it is very difficult to classify
them, and they remain to this day a puzzle to ornithologists. In order to
exhibit clearly the striking characteristics of the bird-fauna of this
island, we shall first give a list of all the peculiar genera; another, of
the genera of which the species only are peculiar; and, lastly, a list of
the species which Madagascar possesses in common with the African
continent.
GENERA OF BIRDS PECULIAR TO MADAGASCAR, OR FOUND ELSEWHERE ONLY IN THE
MASCARENE ISLANDS.
SYLVIIDÆ. Species.
1. Bernieria 2
2. Ellisia 1
3. Mystacornis 1
4. Eroessa 1
5. Gervasia 1
TIMALIIDÆ.
6. Oxylabes 2
CINCLIDÆ(?).
7. Mesites 1
SITTIDÆ.
8. _Hypherpes_ 1
PYCNONOTIDÆ(?)
9. Tylas 1
ORIOLIDÆ.
10. Artamia 3
11. Cyanolanius 1
MUSCICAPIDÆ.
12. Newtonia 1
13. Pseudobias 1
LANIIDÆ.
14. Calicalicus(?) 1
15. Vanga 4
NECTARINIIDÆ.
16. Neodrepanis 1
HIRUNDINIDÆ.
17. Phedina 1
PLOCEIDÆ.
18. Nelicurvius 1
STURNIDÆ.
19. Euryceros(?) 1
20. Hartlaubia 1
21. Falculia 1
PAICTIDÆ.
22. Philepitta 1
CUCULIDÆ.
23. Coua 9
24. Cochlothraustes 1
LEPTOSOMIDÆ.
25. Leptosomus 1
CORACIIDÆ.
26. Atelornis 2
27. Brachypteracias 1
28. Geobiastes 1
PSITTACIDÆ.
29. Coracopsis 2
COLUMBIDÆ.
30. _Alectrænas_ 1
TETRAONIDÆ.
31. _Margaroperdix_ 1
FALCONIDÆ.
32. Nisoides 1
33. Eutriorchis 1
--
Total species of peculiar genera 50
ÆPYORNITHIDÆ(extinct).
34. Æpyornis 1
{276}ETHIOPIAN OR ORIENTAL GENERA WHICH ARE REPRESENTED IN MADAGASCAR BY
PECULIAR SPECIES.
TURDIDÆ. Species.
1. Bessonornis 1
SYLVIIDÆ.
2. Acrocephalus 1
3. _Copsychus (Or.)_ 1
4. Pratincola 1
PYCNONOTIDÆ.
5. _Hypsipetes (Or.)_ 1
6. Andropadus 1
CAMPEPHAGIDÆ.
7. Campephaga 1
DICRURIDÆ.
8. Dicrurus 1
MUSCICAPIDÆ.
9. Tchitrea 1
LANIIDÆ.
10. Laniarius 1
NECTARINIIDÆ.
11. Nectarinia 1
PLOCEIIDÆ.
12. Foudia 2
13. Hypargos 1
14. Spermestes 1
ALAUDIDÆ.
15. Mirafra 1
MOTACILLIDÆ.
16. Motacilla 1
CUCULIDÆ.
17. Ceuthmochares 1
18. Centropus 1
19. Cuculus 1
CORACIIDÆ.
20. Eurystomus 1
ALCEDINIDÆ.
21. Corythornis 1
22. Ispidina 1
UPUPIDÆ.
23. Upupa (?) 1
CAPRIMULGIDÆ.
24. Caprimulgus 1
CYPSELIDÆ.
25. Cypselus 2
26. Chætura 1
PSITTACIDÆ.
27. Poliopsitta 1
COLUMBIDÆ.
28. Treron 1
29. Columba 1
30. Turtur 1
PTEROCLIDÆ.
31. Pterocles 1
TETRAONIDÆ.
32. Francolinus 1
PHASIANIDÆ.
33. Numida 1
TURNICIDÆ.
34. Turnix 1
FALCONIDÆ.
35. Polyboroides 1
36. Circus 1
37. Astur 3
38. Accipiter 1
39. Buteo 1
40. Haliæetus 1
41. Pernis 1
42. Baza 1
43. Cerchneis 1
STRIGIDÆ. {277}
44. Athene 1
45. Scops 1
RALLIDÆ.
46. Rallus 3
47. Porzana 1
SCOLOPACIDÆ.
48. Gallinago 1
PLATALEIDÆ.
49. Ibis 1
PODICIPIDÆ.
50. Podiceps 1
--
Total peculiar species of Eth. }
or Or. genera } 56
SPECIES OF BIRDS COMMON TO MADAGASCAR AND AFRICA OR ASIA.
1. Cisticola cursitans.
2. Corvus scapulatus.
3. Crithagra canicollis.
4. Merops superciliosus.
5. Collocalia fuciphaga.
6. Oena capensis.
7. Aplopelia tympanistria.
8. Falco minor.
9. Falco concolor.
10. Milvus ægyptius.
11. Milvus migrans.
12. Strix flammea.
These three tables show us an amount of speciality hardly to be found in
the birds of any other part of the globe. Out of 111 land-birds in
Madagascar, only 12 are identical with species inhabiting the adjacent
continents, and most of these belong to powerful-winged, or wide-ranging
forms, which probably now often pass from one country to the other. The
peculiar species--49 land-birds and 7 waders, or aquatics--are mostly
well-marked forms of African genera. There are, however, several genera
(marked by italics) which have Oriental or Palæarctic affinities, but not
African, viz.--_Copsychus_, _Hypsipetes_, _Hypherpes_, _Alectrænas_, and
_Margaroperdix_. These indicate a closer approximation to the Malay
countries than now exists.
The table of 33 peculiar genera is of great interest. Most of these are
well-marked forms, belonging to families which are fully developed in
Africa; though it is singular that not one of the exclusively African
families is represented in any way in Madagascar. Others, however, are of
remote or altogether doubtful affinities. _Sittidæ_ is Oriental and
Palæarctic, but not Ethiopian. _Oxylabes_ and _Mystacornis_ are of doubtful
affinities. _Artamia_ and _Cyanolanius_ still more so, and it is quite
undecided what family they belong to. _Calicalicus_ is almost equally
obscure. _Neodrepanis_, one of the most recent discoveries, seems to
connect the Nectariniidæ with the Pacific {278}Depanididæ. _Euryceros_ is a
complete puzzle, having been placed with the hornbills, the starlings, or
as a distinct family. _Falculia_ is an exceedingly aberrant form of
starling, long thought to be allied to _Irrisor_. _Philepitta_, forming a
distinct family, (Paictidæ), is most remarkable and isolated, perhaps with
remote South American affinities. _Leptosoma_ is another extraordinary
form, connecting the cuckoos with the rollers. _Atelornis_,
_Brachypteracias_, and _Geobiastes_, are terrestrial rollers, with the form
and colouring of _Pitta_. So many perfectly isolated and remarkable groups
are certainly nowhere else to be found; and they fitly associate with the
wonderful aye-aye (_Chiromys_), the insectivorous Centetidæ, and
carnivorous _Cryptoprocta_ among the Mammalia. They speak to us plainly of
enormous antiquity, of long-continued isolation; and not less plainly of a
lost continent or continental island, in which so many, and various, and
peculiarly organized creatures, could have been gradually developed in a
connected fauna, of which we have here but the fragmentary remains.
_Plate VI.--Illustrating the characteristic features of the Zoology of
Madagascar._--The lemurs, which form the most prominent feature in the
zoology of Madagascar, being comparatively well-known from the numerous
specimens in our zoological gardens; and good figures of the Insectivorous
genera not being available, we have represented the nocturnal and
extraordinary aye-aye (_Chiromys madagascariensis_) to illustrate its
peculiar and probably very ancient mammalian fauna; while the river-hogs in
the distance (_Potamochoerus edwardsii_) allied to African species,
indicate a later immigration from the mainland than in the case of most of
the other Mammalia. The peculiar birds being far less generally known, we
have figured three of them. The largest is the _Euryceros prevosti_, here
classed with the starlings, although its remarkable bill and other
peculiarities render it probable that it should form a distinct family. Its
colours are velvety black and rich brown with the bill of a pearly grey.
The bird beneath (_Vanga curvirostris_) is one of the peculiar Madagascar
shrikes whose plumage, variegated with green-black and pure white is very
conspicuous; while that in the right hand corner is the _Leptosoma
discolor_, a bird which appears to be intermediate between such very
distinct families as the cuckoos and the rollers, and is therefore
considered to form a family by itself. It is a coppery-green above and
nearly white beneath, with a black bill and red feet. The fan-shaped plant
on the left is the traveller's tree (_Urania speciosa_), one of the
peculiar forms of vegetation in this marvellous island.
Plate VI.
[Illustration]
SCENE IN MADAGASCAR, WITH CHARACTERISTIC ANIMALS.
{279}_Reptiles._--These present some very curious features, comparatively
few of the African groups being represented, while there are a considerable
number of Eastern and even of American forms. Beginning with the snakes, we
find, in the enormous family of Colubridæ, none of the African types; but
instead of them three genera--_Herpetodryas_, _Philodryas_, and
_Heterodon_--only found elsewhere in South and North America. The
Psammophidæ, which are both African and Indian, are represented by a
peculiar genus, _Mimophis_. The Dendrophidæ are represented by _Ahætulla_,
a genus which is both African and American. The Dryiophidæ, which inhabit
all the tropics but are most developed in the Oriental region, are
represented by a peculiar genus, _Langaha_. The tropical Pythonidæ are
represented by another peculiar genus, _Sanzinia_. The Lycodontidæ and
Viperidæ, so well developed in Africa, are entirely absent.
The lizards are no less remarkable. The Zonuridæ, abundantly developed in
Africa, are represented by one peculiar genus, _Cicigna_. The wide-spread
Scincidæ by another peculiar genus, _Pygomeles_. The African Sepsidæ, are
represented by three genera, two of which are African, and one,
_Amphiglossus_, peculiar. The Acontiadæ are represented by a species of the
African genus _Acontias_. Of Scincidæ there is the wide-spread _Euprepes_.
The Sepidæ are represented by the African genera _Seps_ and _Scelotes_. The
Geckotidæ are not represented by any purely African genera, but by
_Phyllodactylus_, which is American and Australian; _Hemidactylus_, which
is spread over all the tropics; by two peculiar genera; and by _Uroplatis_,
_Geckolepis_, and _Phelsuma_, confined to Madagascar, Bourbon, and the
Andaman Islands. The Agamidæ, which are mostly Oriental and are represented
in {280}Africa by the single genus _Agama_, have here three peculiar
genera, _Tracheloptychus_, _Chalarodon_, and _Hoplurus_. Lastly, the
American Iguanidæ are said to be represented by a species of the South
American genus _Oplurus_. The classification of Reptiles is in such an
unsettled state that some of these determinations of affinities are
probably erroneous; but it is not likely that any corrections which may be
required will materially affect the general bearing of the evidence, as
indicating a remarkable amount of Oriental and American relationship.
The other groups are of less interest. Tortoises are represented by two
African or wide-spread genera of Testudinidæ, _Testudo_ and _Chersina_, and
by one peculiar genus, _Pyxis_; and there are also two African genera of
Chelydidæ.
The Amphibia are not very well known. They appear to be confined to species
of the wide-spread Ethiopian and Oriental genera--_Hylarana_,
_Polypedates_, and _Rappia_ (Polypedatidæ); and _Pyxicephalus_ (Ranidæ).
_Fresh-water Fishes._--These appear to be at present almost unknown. When
carefully collected they will no doubt furnish some important facts.
_The Mascarene Islands._
The various islands which surround Madagascar--Bourbon, Mauritius,
Rodriguez, the Seychelles, and the Comoro Islands--all partake in a
considerable degree of its peculiar fauna, while having some special
features of their own.
Indigenous Mammalia (except bats) are probably absent from all these
islands (except the Comoros), although _Lemur_ and _Centetes_ are given as
natives of Bourbon and Mauritius. They have, however, perhaps been
introduced from Madagascar. _Lemur mayottensis_, a peculiar species, is
found in the Comoro Islands, where a Madagascar species of _Viverra_ also
occurs.
Bourbon and Mauritius may be taken together, as they much resemble each
other. They each possess species of a peculiar genus of Campephagidæ, or
caterpillar shrikes, _Oxynotus_; while the remarkable _Fregilupus_,
belonging to the starling family, inhabits Bourbon, if it is not now
extinct. They also have {281}peculiar species of _Pratincola_,
_Hypsipetes_, _Phedina_, _Tchitrea_, _Zosterops_, _Foudia_, _Collocalia_,
and _Coracopsis_; while Mauritius has a very peculiar form of dove of the
sub-genus _Trocaza_; an _Alectrænas_, extinct within the last thirty years;
and a species of the Oriental genus of parroquets, _Palæornis_. The small
and remote island of Rodriguez has another _Palæornis_, as well as a
peculiar _Foudia_, and a _Drymoeca_ of apparently Indian affinity.
Coming to the Seychelle Islands, far to the north, we find the only mammal
an Indian species of bat (_Pteropus edwardsii_). Of the twelve land-birds
all but one are peculiar species, but all belong to genera found also in
Madagascar, except one--a peculiar species of _Palæornis_. This is an
Oriental genus, but found also in several Mascarene Islands and on the
African continent. A species of black parrot (_Coracopsis barklayi_) and a
weaver bird of peculiar type (_Foudia seychellarum_) show, however, a
decided connection with Madagascar. There are also two peculiar pigeons--a
short-winged _Turtur_ and an _Alectrænas_.
Most of the birds of the Comoro Islands are Madagascar species, only two
being African. Five are peculiar, belonging to the genera _Nectarinia_,
_Zosterops_, _Dicrurus_, _Foudia_, and _Alectrænas_.
Reptiles are scarce. There appear to be no snakes in Mauritius and Bourbon,
though some African species are said to be found in the Seychelle Islands.
Lizards are fairly represented. Mauritius has _Cryptoblepharus_, an
Australian genus of Gymnopthalmidæ; _Hemidactylus_ (a wide-spread genus);
_Peropus_ (Oriental and Australian)--both belonging to the Geckotidæ.
Bourbon has _Heteropus_, a Moluccan and Australian genus of Scincidæ;
_Phelsuma_ (Geckotidæ), and _Chameleo_, both found also in Madagascar; as
well as _Pyxis_, one of the tortoises. The Seychelles have _Theconyx_, a
peculiar genus of Geckotidæ, and _Chameleo_. Gigantic land-tortoises, which
formerly inhabited most of the Mascarene Islands, now only survive in
Aldabra, a small island north of the Seychelles. These will be noticed
again further on. Amphibia seem only to be recorded from the Seychelles,
where two genera of tree-frogs of the family Polypedatidæ are found; one
(_Megalixalus_) peculiar, the other (_Rappia_) found also in Madagascar and
Africa.
{282}The few insect groups peculiar to these islands will be noted when we
deal with the entomology of Madagascar.
_Extinct fauna of the Mascarene Islands and Madagascar._--Before quitting
the vertebrate groups, we must notice the remarkable birds which have
become extinct in these islands little more than a century ago. The most
celebrated is the dodo of the Mauritius (_Didus ineptus_), but an allied
genus, _Pezophaps_, inhabited Rodriguez, and of both of these almost
perfect skeletons have been recovered. Other species probably existed in
Bourbon. Remains of two genera of flightless rails have also been found,
_Aphanapteryx_ and _Erythromachus_; and even a heron (_Ardea megacephala_)
which was short-winged and seldom flew; while in Madagascar there lived a
gigantic Struthious bird, the _Æpyornis_. Some further details as to these
extinct forms will be found under the respective families, Dididæ, Rallidæ,
and Æpyornithidæ, in the fourth part of this work; and their bearing on the
past history of the region will be adverted to in the latter part of this
chapter. Dr. Günther has recently distinguished five species of fossil
tortoises from Mauritius and Rodriguez,--all of them quite different from
the living species of Aldabra.
_Insects._--The butterflies of Madagascar are not so remarkable as some
other orders of insects. There seems to be only one peculiar genus,
_Heteropsis_ (Satyridæ). The other genera are African, _Leptoneura_ being
confined to Madagascar and South Africa. There are some fine _Papilios_ of
uncommon forms. The most interesting lepidopterous insect, however, is the
fine diurnal moth (_Urania_), as all the other species of the genus inhabit
tropical America and the West Indian Islands.
The Coleoptera have been better collected, and exhibit some very remarkable
affinities. There is but one peculiar genus of Cicindelidæ, _Pogonostoma_,
which is allied to the South American genus, _Ctenostoma_. Another genus,
_Peridexia_, is common to Madagascar and South America. None of the
important African genera are represented, except _Eurymorpha_; while
_Meglaomma_ is common to Madagascar and the Oriental region.
In the Carabidæ we have somewhat similar phenomena on a {283}wider scale.
Such large and important African genera as _Polyhirma_ and _Anthia_, are
absent; but there are four genera in common with South Africa, and two with
West Africa; while three others are as much Oriental as African. One genus,
_Distrigus_, is wholly Oriental; and another, _Homalosoma_, Australian.
_Colpodes_, well developed in Bourbon and Mauritius, is Oriental and South
American. Of the peculiar genera, _Sphærostylis_ has South American
affinities; _Microchila_, Oriental; the others being related to widely
distributed genera.
The Lucanidæ are few in number, and all have African affinities. Madagascar
is very rich in Cetoniidæ, and possesses 20 peculiar genera. _Bothrorhina_,
and three other genera belonging to the _Ichnostoma_ group, have wholly
African relations. _Doryscelis_ and _Chromoptila_ are no less clearly
allied to Oriental genera. A series of eight peculiar genera belong to the
Schizorhinidæ, a family the bulk of which are Australian, while there are
only a few African forms. The remaining genera appear to have African
affinities, but few of the peculiarly African genera are represented.
_Glyciphana_ is characteristic of the Oriental region.
The Buprestidæ of Madagascar consist mainly of one large and peculiar
genus, _Polybothris_, allied to the almost cosmopolite _Psiloptera_. Most
of the other genera are both Ethiopian and Oriental; but _Polycesta_ is
mainly South American, and the remarkable and isolated genus _Sponsor_ is
confined to the Mauritius with a species in Celebes and New Guinea.
The Longicorns are numerous and interesting, there being no less than 24
peculiar genera. Two of the genera of Prionidæ are very isolated, while a
third, _Closterus_, belongs to a group which is Malayan and American.
Of the Cerambycidæ, _Philematium_ ranges to Africa and the West Indies;
_Leptocera_ is only found eastward in Ceylon and the New Hebrides; while
_Euporus_ is African. Of the peculiar genera, 2 are of African type; 3
belong to the _Leptura_ group, which are mostly Palæarctic and Oriental,
with a few in South Africa; while _Philocalocera_ is allied to a South
American genus.
Among the Lamiidæ there are several wide-ranging and 7 {284}African genera;
but _Coptops_ is Oriental, and the Oriental _Praonetha_ occurs in the
Comoro Islands. Among the peculiar genera several have African affinities,
but _Tropidema_ belongs to a group which is Oriental and Australian;
_Oopsis_ is found also in the Pacific Islands; _Mythergates_, _Sulemus_,
and _Coedomæa_, are allied to Malayan and American genera.
_General Remarks on the Insect-fauna of Madagascar._--Taking the insects as
a whole, we find the remarkable result that their affinities are largely
Oriental, Australian, and South American: while the African element is
represented chiefly by special South African or West African forms, rather
than by such as are widely spread over the Ethiopian region.[11] In some
families--as Cetoniidæ and Lamiidæ--the African element appears to
preponderate; in others--as Cicindelidæ--the South American affinity seems
strongest; in Carabidæ, perhaps the Oriental; while in Buprestidæ and
Cerambycidæ the African and foreign elements seem nearly balanced. We must
not impute too much importance to these foreign alliances among insects,
because we find examples of them in every country on the globe. The reason
they are so much more pronounced in Madagascar may be, that during long
periods of time this island has served as a refuge for groups that have
been dying out on the great continents; and that, owing to the numerous
deficiencies of a somewhat similar kind in the series of vertebrata in
Australia and South America, the same groups have often been able to
maintain themselves in all these countries as well as in Madagascar. It
must be remembered too, that these peculiarities in the Malagasy and
Mascarene insect-fauna are but exaggerations of a like phenomenon on the
mainland. Africa also has numerous affinities with South America, with the
Malay countries, and with Australia; but they do not bear anything like so
large a proportion to the whole fauna, and do not, therefore, attract so
much attention. The special conditions of existence and the long-continued
isolation of Madagascar, will account for much of this difference; and it
will evidently not be necessary {285}to introduce, as some writers are
disposed to do, a special land connection or near approach between
Madagascar and all these countries, independently of Africa; except perhaps
in the case of the Malay Islands, as will be discussed further on.
_Land-shells._--Madagascar and the adjacent islands are all rich in
land-shells. The genera of Helicidæ are _Vitrina_, _Helix_, _Achatina_,
_Columna_ (peculiar to Madagascar and West Africa), _Buliminus_, _Cionella_
(chiefly Oriental and South American, but not African), _Pupa_,
_Streptaxis_, and _Succinea_. Among the Operculata we have _Truncatella_
(widely scattered, but not African); _Cyclotus_ (South American, Oriental,
and South African); _Cyclophorus_ (mostly Oriental, with a few South
African); _Leptopoma_ (Oriental); _Megalomastoma_ (Malayan and South
American); _Lithidion_ (peculiar to Madagascar, Socotra, and South-West
Arabia); _Otopoma_ (with the same range, but extending to West India and
New Ireland); _Cyclostomus_ (widely spread but not African); and
_Omphalotropis_ (wholly Oriental and Australian). We thus find the same
general features reproduced in the land-shells as in the insects, and the
same remarks will to a great extent apply to both. The classification of
the former is, however, by no means so satisfactory, and we have no
extensive and accurate general catalogues of shells, like those of
Lepidoptera and Coleoptera, which have furnished us with such valuable
materials for the comparison of the several faunas.
_On the probable Past History of the Ethiopian Region._
Perhaps none of the great zoological regions of the earth present us with
problems of greater difficulty or higher interest than the Ethiopian. We
find in it the evidence of several distinct and successive faunas, now
intermingled; and it is very difficult, with our present imperfect
knowledge, to form an adequate conception of how and when the several
changes occurred. There are, however, a few points which seem sufficiently
clear, and these afford us a secure foundation in our endeavour to
comprehend the rest.
Let us then consider what are the main facts we have to account for.--1. In
Continental Africa, more especially in the south {286}and west, we find,
along with much that is peculiar, a number of genera showing a decided
Oriental, and others with an equally strong South American affinity; this
latter more particularly showing itself among reptiles and insects. 2. All
over Africa, but more especially in the east, we have abundance of large
ungulates and felines--antelopes, giraffes, buffaloes, elephants, and
rhinoceroses, with lions, leopards, and hyænas, all of types now or
recently found in India and Western Asia. 3. But we also have to note the
absence of a number of groups which abound in the above-named countries,
such as deer, bears, moles, and true pigs; while camels and
goats--characteristic of the desert regions just to the north of the
Ethiopian--are equally wanting. 4. There is a wonderful unity of type and
want of speciality in the vast area of our first sub-region extending from
Senegal across to the east coast, and southward to the Zambezi; while West
Africa and South Africa each abound in peculiar types. 5. We have the
extraordinary fauna of Madagascar to account for, with its evident main
derivation from Africa, yet wanting all the larger and higher African
forms; its resemblances to Malaya and to South America; and its wonderful
assemblage of altogether peculiar types.
Here we find a secure starting-point, for we are sure that Madagascar must
have been separated from Africa before the assemblage of large animals
enumerated above, had entered it. Now, it is a suggestive fact, that all
these belong to types which abounded in Europe and India about the Miocene
period. It is also known, from the prevalence of Tertiary deposits over the
Sahara and much of Arabia, Persia, and Northern India, that during early
Tertiary times a continuous sea from the Bay of Bengal to the British Isles
completely cut off all land communication between Central and Southern
Africa on the one side, and the great continent of the Eastern hemisphere
on the other. When Africa was thus isolated, its fauna probably had a
character somewhat analogous to that of South America at the same period.
Most of the higher types of mammalian life were absent, while lemurs,
Edentates, and Insectivora took their place. At this period Madagascar was
no doubt united with Africa, {287}and helped to form a great southern
continent which must at one time have extended eastward as far as Southern
India and Ceylon; and over the whole of this the lemurine type no doubt
prevailed.
During some portion of this period, South Temperate Africa must have had a
much greater extension, perhaps indicated by the numerous shoals and rocks
to the south and east of the Cape of Good Hope, and by the Crozets and
Kerguelen Islands further to the south-east. This would have afforded means
for that intercommunion with Western Australia which is so clearly marked
in the flora, and to some extent also in the insects of the two countries;
and some such extension is absolutely required for the development of that
wonderfully rich and peculiar temperate flora and fauna, which, now crowded
into a narrow territory, is one of the greatest marvels of the organic
world.
During this early period, when the great southern continents--South
America, Africa, and Australia--were equally free from the incursions of
the destructive felines of the north, the Struthious or ostrich type of
birds was probably developed into its existing forms. It is not at all
necessary to suppose that these three continents were at any time united,
in order to account for the distribution of these great terrestrial birds;
as this may have arisen by at least two other easily conceivable modes. The
ancestral Struthious type may, like the Marsupial, have once spread over
the larger portion of the globe; but as higher forms, especially of
Carnivora, became developed, it would be exterminated everywhere but in
those regions where it was free from their attacks. In each of these it
would develope into special forms adapted to surrounding conditions; and
the large size, great strength, and excessive speed of the ostrich, may
have been a comparatively late development caused by its exposure to the
attacks of enemies which rendered such modification necessary. This seems
the most probable explanation of the distribution of Struthious birds, and
it is rendered almost certain by the discovery of remains of this order in
Europe in Eocene deposits, and by the occurrence of an ostrich among the
fossils of the Siwalik hills; but it is just possible, also, that the
{288}ancestral type may have been a bird capable of flight, and that it
spread from one of the three southern continents to the others at the
period of their near approach, and more or less completely lost the power
of flight owing to the long continued absence of enemies.
During the period we have been considering, the ancestors of existing apes
and monkeys flourished (as we have seen in Chapter VI.) along the whole
southern shores of the old Palæarctic continent; and it seems likely that
they first entered Africa by means of a land connection indicated by the
extensive and lofty plateaus of the Sahara, situated to the south-east of
Tunis and reaching to a little north-west of Lake Tchad; and at the same
time the elephant and rhinoceros type may have entered. This will account
for the curious similarity between the higher faunas of West Africa and the
Indo-Malay sub-region, for owing to the present distribution of land and
sea and the narrowing of the tropical zone since Miocene times, these are
now the only lowland, equatorial, forest-clad countries, which were in
connection with the southern shores of the old Palæarctic continent at the
time of its greatest luxuriance and development. This western connection
did not probably last long, the junction that led to the greatest incursion
of new forms, and the complete change in the character of the African
fauna, having apparently been effected by way of Syria and the shores of
the Red Sea at a somewhat later date. By this route the old
South-Palæarctic fauna, indicated by the fossils of Pikermi and the Siwalik
Hills, poured into Africa; and finding there a new and favourable country,
almost wholly unoccupied by large Mammalia, increased to an enormous
extent, developed into new forms, and finally overran the whole continent.
Before this occurred, however, a great change had taken place in the
geography of Africa. It had gradually diminished on the south and east;
Madagascar had been left isolated; while a number of small islands, banks,
and coral reefs in the Indian Ocean alone remained to indicate the position
of a once extensive equatorial land. The Mascarene Islands appear to
represent the portion which separated earliest, before any carnivora had
{289}reached the country; and it was in consequence of this total exemption
from danger, that several groups of birds altogether incapable of flight
became developed here, culminating in the huge and unwieldy Dodo, and the
more active Aphanapteryx. To the same cause may be attributed the
development, in these islands, of gigantic land-tortoises, far surpassing
any others now living on the globe. They appear to have formerly inhabited
Mauritius, Bourbon, and Rodriguez, and perhaps all the other Mascarene
islands, but having been recklessly destroyed, now only survive in the
small uninhabited Aldabra islands north of the Seychelle group. The largest
living specimen (5½ feet long) is now in our Zoological Gardens. The only
other place where equally large tortoises (of an allied species) are found,
is the Galapagos islands, where they were equally free from enemies till
civilized man came upon the scene; who, partly by using them for food,
partly by the introduction of pigs, which destroy the eggs, has greatly
diminished their numbers and size, and will probably soon wholly
exterminate them. It is a curious fact, ascertained by Dr. Günther, that
the tortoises of the Galapagos are more nearly related to the extinct
tortoises of Mauritius than is the living tortoise of Aldabra. This would
imply that several distinct groups or sub-genera of _Testudo_ have had a
wide range over the globe, and that some of each have survived in very
distant localities. This is rendered quite conceivable by the known
antiquity of the genus _Testudo_, which dates back to at least the Eocene
formation (in North America) with very little change of form. These
sluggish reptiles, so long-lived and so tenacious of life, may have
remained unchanged, while every higher animal type around them has become
extinct and been replaced by very different forms; as in the case of the
living _Emys tectum_, which is the sole survivor of the strange Siwalik
fauna of the Miocene epoch. The ascertained history of the genus and the
group, thus affords a satisfactory explanation of the close affinity of the
gigantic tortoises of Mauritius and the Galapagos.
The great island of Madagascar seems to have remained longer united with
Africa, till some of the smaller and more active {290}carnivora had reached
it; and we consequently find there, no wholly terrestrial form of bird but
the gigantic and powerful _Æpyornis_, well able to defend itself against
such enemies. As already intimated, we refer the South American element in
Madagascar, not to any special connection of the two countries
independently of Africa, but to the preservation there of a number of
forms, some derived from America through Africa, others of once almost
cosmopolitan range, but which, owing to the severer competition, have
become extinct on the African continent, while they have continued to exist
under modified forms in the two other countries.
The depths of all the great oceans are now known to be so profound, that we
cannot conceive the elevation of their beds above the surface without some
corresponding depression elsewhere. And if, as is probable, these opposite
motions of the earth's crust usually take place in parallel bands, and are
to some extent dependent on each other, an elevation of the sea bed could
hardly fail to lead to the submergence of large tracts of existing
continents; and this is the more likely to occur on account of the great
disproportion that we have seen exists between the mean height of the land
and the mean depth of the ocean. Keeping this principle in view, we may,
with some probability, suggest the successive stages by which the Ethiopian
region assumed its present form, and acquired the striking peculiarities
that characterise its several sub-regions. During the early period, when
the rich and varied temperate flora of the Cape, and its hardly less
peculiar forms of insects and of low type mammalia, were in process of
development in an extensive south temperate land, we may be pretty sure
that the whole of the east and much of the north of Africa was deep sea. At
a later period, when this continent sank towards the south and east, the
elevation may have occurred which connected Madagascar with Ceylon; and
only at a still later epoch, when the Indian Ocean had again been formed,
did central, eastern, and northern Africa gradually rise above the ocean,
and effect a connection with the great northern continent by way of
Abyssinia and Arabia. And if this last change took place with
{291}tolerable rapidity, or if the elevatory force acted from the north
towards the south, there would be a new and unoccupied territory to be
taken possession of by immigrants from the north, together with a few from
the south and west. The more highly-organised types from the great northern
continent, however, would inevitably prevail; and we should thus have
explained the curious uniformity in the fauna of so large an area, together
with the absence from it of those peculiar Ethiopian types which so
abundantly characterise the other three sub-regions.
We may now perhaps see the reason of the singular absence from tropical
Africa of deer and bears; for these are both groups which live in fertile
or well-wooded countries, whereas the line of immigration from Europe to
Africa was probably always, as now, to a great extent a dry and desert
tract, suited to antelopes and large felines, but almost impassable to deer
and bears. We find, too, that whereas remains of antelopes and giraffes
abound in the Miocene deposits of Greece, there were no deer (which are
perhaps a somewhat later development); neither were there any bears, but
numerous forms of Felidæ, Viverridæ, Mustelidæ, and ancestral forms of
_Hyæna_, exactly suited to be the progenitors of the most prevalent types
of modern African Zoology.
There appears to have been one other change in the geography of Africa and
the Atlantic Ocean that requires notice. The rather numerous cases of close
similarity in the insect forms of tropical Africa and America, seem to
indicate some better means of transmission, at a not very remote epoch,
than now exists. The vast depth of the Atlantic, and the absence of any
corresponding likeness in the vertebrate fauna, entirely negative the idea
of any union between the two countries; but a moderate extension of their
shores towards each other is not improbable, and this, with large islands
in the place of the Cape Verd group, St. Paul's Rocks, and Fernando
Noronha, to afford resting places in the Atlantic, would probably suffice
to explain the amount of similarity that actually exists.
Our knowledge of the geology and palæontology of Africa {292}being so
scanty, it would be imprudent to attempt any more detailed explanation of
the peculiarities of its existing fauna. The sketch now given is, it is
believed, founded on a sufficient basis of facts to render it not only a
possible but a probable account of what took place; and it is something
gained to be able to show, that a large portion of the peculiarities and
anomalies of so remarkable a fauna as that of the Ethiopian region, can be
accounted for by a series of changes of physical geography during the
tertiary epoch, which can hardly be considered extreme, or in any way
unlikely to have occurred.
{293}TABLES OF DISTRIBUTION.
In drawing up these tables showing the distribution of various classes of
animals in the Ethiopian Region, the following sources of information have
been chiefly relied on, in addition to the general treatises, monographs,
and catalogues, used for the Fourth Part of this work:--
_Mammalia._--Blanford's Abyssinia; Peters's Mozambique; Heuglin and
Schweinfurth for North East Africa; Grandidier Schlegel, &c., for
Madagascar; the local lists given by Mr. Andrew Murray; numerous papers by
Fraser, Gray, Kirk, Mivart, Peters, Sclater, and Speke; and a MS. list of
Bovidæ from Sir Victor Brooke.
_Birds._--Finsch and Hartlaub for East Africa; Heuglin for North-East
Africa; Blanford for Abyssinia; Layard for South Africa; Hartlaub for West
Africa; Dohrn for Princes Island; Andersson for Damaraland; and papers by
Gurney, Hartlaub, Kirk, Newton, Peters, Sharpe, Sclater, Schlegel, and
Pollen and a MS. list of Madagascar Birds from Mr. Sharpe.
{294}TABLE I.
_FAMILIES OF ANIMALS INHABITING THE ETHIOPIAN REGION._
EXPLANATION.
Names in _italics_ show families peculiar to the region.
Names inclosed thus (......) barely enter the region, and are not
considered properly to belong to it.
Numbers are not consecutive, but correspond to those in Part IV.
---------------------+-------------------+-------------------------------
| Sub-regions |
| 1=East Africa. |
Order and Family | 2=West Africa. | Range beyond the Region.
| 3=South Africa. |
| 4=Madagascar. |
---------------------+----+----+----+----+-------------------------------
| 1. | 2. | 3. | 4. |
---------------------+----+----+----+----+-------------------------------
| | | | |
MAMMALIA. | | | | |
PRIMATES. | | | | |
1. Simiidæ | | -- | | |Oriental
2. Semnopithecidæ | -- | -- | | |Oriental
3. Cynopithecidæ | -- | -- | -- | |Oriental, Palæarctic
6. Lemuridæ | -- | -- | -- | -- |Oriental
8. _Chiromyidæ_ | | | | -- |
| | | | |
CHEIROPTERA. | | | | |
9. Pteropidæ | -- | -- | -- | -- |Oriental, Australian
11. Rhinolophidæ | -- | -- | -- | -- |The Eastern Hemisphere
12. Vespertilionidæ | -- | -- | -- | -- |Cosmopolite
13. Noctilionidæ | -- | -- | -- | -- |All Tropical regions
| | | | |
INSECTIVORA. | | | | |
15. Macroscelididæ | -- | | -- | |South Palæarctic
17. Erinaceidæ | | | -- | |Palæarctic, Oriental
18. _Centetidæ_ | | | | -- |Greater Antilles
19. _Potamogalidæ_ | | -- | | |
20. _Chrysochloridæ_ | -- | | -- | |
22. Soricidæ | -- | -- | -- | -- |All regions but Australian and
| | | | | Neotropical
| | | | |
CARNIVORA. | | | | |
23. Felidæ | -- | -- | -- | -- |All regions but Australian
24. _Cryptoproctidæ_ | | | | -- |
25. Viverridæ | -- | -- | -- | -- |Oriental, S. Palæarctic
26. _Protelidæ_ | | | -- | |
27. Hyænidæ | -- | -- | -- | |S. Palæarctic, India
28. Canidæ | -- | -- | -- | |Almost cosmopolite
29. Mustelidæ | -- | -- | -- | |All regions but Australian
33. Otariidæ | | | -- | |All temperate regions
| | | | |
CETACEA. | | | | |
36 to 41. | | | | |Oceanic
| | | | |
SIRENIA. | | | | |
42. Manatidæ | -- | -- | | |Neotropical, Oriental,
| | | | | Australian
| | | | |
UNGULATA. | | | | |
43. Equidæ | -- | -- | -- | |Palæarctic
45. Rhinocerotidæ | -- | -- | -- | |Oriental
46. _Hippopotamidæ_ | -- | -- | -- | |
47. Suidæ | -- | -- | -- | -- |Cosmopolite; excl. Australia
49. Tragulidæ | | -- | | |Oriental
51. _Camelopardidæ_ | -- | | -- | |
52. Bovidæ | -- | -- | -- | |All regions but Neotrop. and
| | | | | Australian
| | | | |
PROBOSCIDEA. | | | | |
53. Elephantidæ | -- | -- | -- | |Oriental
| | | | |
HYRACOIDEA. | | | | |
54. Hyracidæ | -- | -- | -- | |Syria
| | | | |
RODENTIA. | | | | |
55. Muridæ | -- | -- | -- | -- |Cosmopolite; excl. Oceania
56. Spalacidæ | -- | -- | -- | |Palæarctic, Oriental
57. Dipodidæ | -- | -- | -- | |Palæarctic, Nearctic
58. Myoxidæ | -- | -- | -- | |Palæarctic
61. Sciuridæ | -- | -- | -- | |All regions but Australian
64. Octodontidæ | -- | | | |N. Africa, Neotropical
65. Echimyidæ | -- | | -- | |Neotropical
67. Hystricidæ | -- | -- | -- | |S. Palæarctic, Oriental
70. Leporidæ | -- | | -- | |All regions but Australian
| | | | |
EDENTATA. | | | | |
72. Manididæ | -- | -- | -- | |Oriental
74. _Orycteropodidæ_ | -- | | -- | |
| | | | |
BIRDS. | | | | |
PASSERES. | | | | |
1. Turdidæ | -- | -- | -- | -- |Almost Cosmopolite
2. Sylviidæ | -- | -- | -- | -- |Cosmopolite
3. Timaliidæ | -- | -- | -- | -- |Oriental, Australian
5. Cinclidæ? | | | | -- |Widely scattered
6. Troglodytidæ | -- | -- | -- | |Almost Cosmopolite
9. Sittidæ | | | | -- |Palæarctic, Oriental,
| | | | | Australian
10. Paridæ | -- | -- | -- | |All regions but Australian
13. Pycnonotidæ | -- | -- | -- | -- |Oriental
14. Oriolidæ | -- | -- | -- | -- |Oriental, Australian
15. Campephagidæ | -- | -- | -- | -- |Oriental, Australian
16. Dicruridæ | -- | -- | -- | -- |Oriental, Australian
17. Muscicapidæ | -- | -- | -- | -- |The Eastern Hemisphere
19. Laniidæ | -- | -- | -- | -- |The Eastern Hemisphere and
| | | | | North America
20. Corvidæ | -- | -- | -- | -- |Cosmopolite
23. Nectariniidæ | -- | -- | -- | -- |Oriental, Australian
24. Dicæidæ | -- | -- | -- | -- |Oriental, Australian
30. Hirundinidæ | -- | -- | -- | -- |Cosmopolite
33. Fringillidæ | -- | -- | -- | -- |Cosmopolite, except Australian
| | | | | region
34. Ploceidæ | -- | -- | -- | -- |Oriental, Australian
35. Sturnidæ | -- | -- | -- | -- |Eastern Hemisphere
37. Alaudidæ | -- | -- | -- | -- |Eastern Hemisphere and North
| | | | | America
38. Motacillidæ | -- | -- | -- | -- |The Eastern Hemisphere
47. Pittidæ | | -- | | |Oriental, Australian
48. _Paictidæ_ | | | | -- |
| | | | |
PICARIÆ. | | | | |
51. Picidæ | -- | -- | -- | |Cosmopolite, excl. Australian
| | | | | region
52. Yungidæ | -- | | -- | |Palæarctic
53. Indicatoridæ | -- | -- | -- | |Oriental
54. Megalæmidæ | -- | -- | -- | |Oriental, Neotropical
56. _Musophagidæ_ | -- | -- | -- | |
57. _Coliidæ_ | -- | -- | -- | |
58. Cuculidæ | -- | -- | -- | -- |Cosmopolite
59. _Leptosomidæ_ | | | | -- |
62. Coraciidæ | -- | -- | -- | -- |Oriental, Australian
63. Meropidæ | -- | -- | -- | -- |Oriental, Australian
66. Trogonidæ | -- | -- | -- | |Oriental, Neotropical
67. Alcedinidæ | -- | -- | -- | -- |Cosmopolite
68. Bucerotidæ | -- | -- | -- | |Oriental and to N. Guinea
69. Upupidæ | -- | -- | -- | -- |Palæarctic, Oriental
70. _Irrisoridæ_ | -- | -- | -- | |
73. Caprimulgidæ | -- | -- | -- | -- |Cosmopolite
74. Cypselidæ | -- | -- | -- | -- |Almost Cosmopolite
| | | | |
PSITTACI. | | | | |
78. Palæornithidæ | -- | -- | | -- |Oriental
81. Psittacidæ | -- | -- | -- | -- |Neotropical
| | | | |
COLUMBÆ. | | | | |
84. Columbidæ | -- | -- | -- | -- |Cosmopolite
85. _Dididæ_ | | | | -- |(Extinct)
| | | | |
GALLINÆ. | | | | |
86. Pteroclidæ | -- | | -- | -- |Palæarctic, Oriental
87. Tetraonidæ | -- | -- | -- | -- |Eastern Hemisphere and
| | | | | N. America
88. Phasianidæ | -- | -- | -- | -- |Old World and N. America
89. Turnicidæ | -- | -- | -- | -- |Eastern Hemisphere.
| | | | |
ACCIPITRES. | | | | |
94. Vulturidæ | -- | -- | -- | |All the continents but
| | | | | Australia
95. _Serpentariidæ_ | -- | -- | -- | |
96. Falconidæ | -- | -- | -- | -- |Cosmopolite
97. Pandionidæ | -- | -- | -- | -- |Cosmopolite
98. Strigidæ | -- | -- | -- | -- |Cosmopolite
| | | | |
GRALLÆ. | | | | |
99. Rallidæ | -- | -- | -- | -- |Cosmopolite
100. Scolopacidæ | -- | -- | -- | -- |Cosmopolite
103. Parridæ | -- | -- | -- | -- |Tropical
104. Glareolidæ | -- | -- | -- | -- |Eastern Hemisphere
105. Charadriidæ | -- | -- | -- | -- |Cosmopolite
106. Otididæ | -- | -- | -- | |Eastern Hemisphere
107. Gruidæ | -- | -- | -- | |All regions but Neotropical
113. Ardeidæ | -- | -- | -- | -- |Cosmopolite
114. Plataleidæ | -- | -- | -- | -- |Almost Cosmopolite
115. Ciconiidæ | -- | -- | -- | -- |Almost Cosmopolite
117. Phoenicopteridæ | -- | -- | -- | -- |Oriental and Neotropical
| | | | |
ANSERES. | | | | |
118. Anatidæ | -- | -- | -- | -- |Cosmopolite
119. Laridæ | -- | -- | -- | -- |Cosmopolite
120. Procellariidæ | -- | -- | -- | -- |Cosmopolite
121. Pelecanidæ | -- | -- | -- | -- |Cosmopolite
122. Spheniscidæ | | | -- | |South temperate regions
124. Podicipidæ | -- | -- | -- | -- |Cosmopolite
126. Struthionidæ | -- | | -- | |Temperate S. America
131. _Æpyornithidæ_ | | | | -- |(Extinct)
| | | | |
REPTILIA. | | | | |
OPHIDIA. | | | | |
1. Typhlopidæ | -- | -- | -- | -- |All regions but Nearctic
5. Calamariidæ | -- | -- | -- | |Warm parts of all regions
7. Colubridæ | -- | -- | -- | -- |Almost Cosmopolite
8. Homalopsidæ | | -- | | |Oriental, and all other regions
9. Psammophidæ | -- | -- | -- | -- |Oriental and S. Palæarctic
10. _Rachiodontidæ_ | | -- | -- | |
11. Dendrophidæ | -- | -- | -- | -- |Oriental, Australian,
| | | | | Neotropical
12. Dryiophidæ | | -- | | -- |Oriental, Neotropical
13. Dipsadidæ | -- | -- | -- | |Oriental, Australian,
| | | | | Neotropical
15. Lycodontidæ | -- | -- | -- | |Oriental
17. Pythonidæ | -- | -- | -- | -- |All tropical regions
18. Erycidæ | | -- | | |Oriental, S. Palæarctic
20. Elapidæ | -- | -- | -- | |Tropical regions, S. U. States
| | | | | and Japan
21. _Dendraspididæ_ | -- | -- | | |
22. _Atractaspididæ_ | | -- | -- | |
23. Hydrophidæ | | | | -- |Oriental, Australian, Panama
25. Viperidæ | -- | -- | -- | -- |Oriental, Palæarctic
| | | | |
LACERTILIA. | | | | |
28. Amphisbænidæ | -- | -- | | |S. Europe, Neotropical
29. Lepidosternidæ | | -- | -- | |N. America
30. Varanidæ | -- | -- | -- | |Warm parts of E. Hemisphere
33. Lacertidæ | -- | -- | -- | |All continents but America
34. Zonuridæ | -- | -- | -- | -- |All America, N. India,
| | | | | S. Europe
40. _Chamæsauridæ_ | | | -- | |
41. Gymnopthalmidæ | | -- | | -- |Palæarctic, Australian,
| | | | | Neotropical
45. Scincidæ | -- | -- | -- | -- |Almost Cosmopolite
47. Sepidæ | -- | -- | -- | -- |South
48. Acontiadæ | | -- | -- | -- |Ceylon and Moluccas
49. Geckotidæ | -- | -- | -- | -- |Almost cosmopolite
51. Agamidæ | -- | -- | -- | -- |Oriental, Australian,
| | | | | S. Palæarctic
52. Chamæleonidæ | -- | -- | -- | -- |Oriental, S. Palæarctic
| | | | |
CROCODILIA. | | | | |
55. Crocodilidæ | -- | -- | -- | -- |Oriental, Neotropical
| | | | |
CHELONIA. | | | | |
57. Testudinidæ | -- | -- | -- | -- |All continents but Australia
58. Chelydidæ | -- | -- | -- | -- |Australia, S. America
59. Trionychidæ | -- | -- | -- | |Oriental, Japan, E. United
| | | | | States
60. Cheloniidæ | | | | |Marine
| | | | |
AMPHIBIA. | | | | |
PSEUDOPHIDIA. | | | | |
1. Cæciliadæ | | -- | | |Oriental, Neotropical
| | | | |
ANOURA. | | | | |
7. Phryniscidæ | -- | -- | | |Neotropical, Australia, Java
9. Bufonidæ | -- | -- | -- | |All regions but Australian
11. Engystomidæ | | -- | -- | |All regions but Palæarctic
14. Alytidæ | -- | -- | -- | |All regions but Oriental
17. Polypedatidæ | -- | -- | -- | -- |All the regions
18. Ranidæ | -- | -- | -- | -- |Almost Cosmopolite
19. Discoglossidæ | | -- | -- | |All regions but Nearctic
21. _Dactylethridæ_ | -- | -- | -- | |
| | | | |
FISHES (FRESH-WATER).| | | | |
ACANTHOPTERYGII. | | | | |
3. Percidæ | -- | | | |All regions but Australian
12. Scienidæ | -- | -- | -- | |All regions but Australian
35. Labyrinthici | | | -- | -- |Oriental, Moluccas
38. Mugillidæ | -- | -- | -- | -- |Australian, Neotropical
52. Chromidæ | -- | -- | -- | -- |Oriental, Neotropical
| | | | |
PHYSOSTOMI. | | | | |
59. Siluridæ | -- | -- | -- | -- |All warm regions
60. Characinidæ | -- | -- | | |Neotropical
68. _Mormyridæ_ | -- | -- | | |
69. _Gymnarchidæ_ | -- | -- | | |
73. Cyprinodontidæ | -- | -- | | -- |Palæarctic, Oriental, American
75. Cyprinidæ | -- | -- | -- | -- |Absent from Australia and
| | | | | S. America
78. Osteoglossidæ | -- | -- | | |All tropical regions
82. Notopteridæ | | -- | | |Oriental
| | | | |
GANOIDEI. | | | | |
92. Sirenoidei | -- | -- | | |Neotropical, Australian
94. _Polypteridæ_ | -- | -- | | |
| | | | |
INSECTS. | | | | |
LEPIDOPTERA (PART). | | | | |
DIURNI (BUTTERFLIES).| | | | |
1. Danaidæ | -- | -- | -- | -- |All warm countries and Canada
2. Satyridæ | -- | -- | -- | -- |Cosmopolite
3. Elymniidæ | | -- | | |Oriental, Moluccas
6. Acræidæ | -- | -- | -- | -- |All tropical regions
8. Nymphalidæ | -- | -- | -- | -- |Cosmopolite
9. Libytheidæ | | -- | | -- |Absent from Australia only
10. Nemeobiidæ | | -- | | -- |Absent from Australia and
| | | | | Nearctic region
13. Lycænidæ | -- | -- | -- | -- |Cosmopolite
14. Pieridæ | -- | -- | -- | -- |Cosmopolite
15. Papilionidæ | -- | -- | -- | -- |Cosmopolite
16. Hesperidæ | -- | -- | -- | -- |Cosmopolite
| | | | |
SPHINGIDEA. | | | | |
17. Zygænidæ | -- | -- | -- | -- |Cosmopolite
19. Agaristidæ | -- | -- | -- | -- |Australian, Oriental
20. Uraniidæ | | | | -- |All tropical regions
22. Ægeriidæ | -- | -- | -- | -- |Cosmopolite, excl. Australia
23. Sphingidæ | -- | -- | -- | -- |Cosmopolite
---------------------+----+----+----+----+-------------------------------
{300}TABLE II.
_LIST OF GENERA OF TERRESTRIAL MAMMALIA AND BIRDS INHABITING THE ETHIOPIAN
REGION._
EXPLANATION.
Names in _italics_ show genera peculiar to the region.
Names inclosed thus (......) show genera which just enter the region, but
are not considered properly to belong to it.
Genera which undoubtedly belong to the region are numbered consecutively.
_MAMMALIA._
-------------------+-------+----------------------+----------------------
Order, Family, and | No. of| Range within | Range beyond
Genus. |Species| the Region. | the Region.
-------------------+-------+----------------------+----------------------
| | |
PRIMATES. | | |
SIMIIDÆ. | | |
| | |
1. _Troglodytes_ | 2 |W. Africa to Western |
| | Nile Sources |
| | |
SEMNOPITHECIDÆ. | | |
2. _Colobus_ | 11 |Abyssinia to West |
| | Africa |
| | |
CYNOPITHECIDÆ. | | |
| | |
3. _Myiopithecus_ | 1 |West Africa |
4. _Cercopithecus_| 24 |Tropical Africa |
5. _Cercocebus_ | 5 |West Africa |
6. _Theropithecus_| 2 |North-east Africa, |Palestine
| | Arabia |
7. _Cynocephalus_ | 10 |Nubia to Cape, |
| | W. Africa, Arabia |
| | |
(Sub-Order) | | |
_LEMUROIDEA._ | | |
LEMURIDÆ. | | |
| | |
8. _Indris_ | 6 |Madagascar |
9. _Lemur_ | 15 |Madagascar |
10. _Hapalemur_ | 2 |Madagascar |
11. _Microcebus_ | 4 |Madagascar |
12. _Chirogaleus_ | 5 |Madagascar |
13. _Lepilemur_ | 2 |Madagascar |
14. _Perodicticus_ | 1 |Sierra Leone |
15. _Arctocebus_ | 1 |Old Calabar |
16. _Galago_ | 14 |Tropical and S. Africa |
| | |
CHIROMYIDÆ. | | |
| | |
17. _Chiromys_ | 1 |Madagascar |
| | |
CHIROPTERA. | | |
PTEROPIDÆ. | | |
| | |
18. Pteropus | 7 |Africa and Madagascar |Tropics of Eastern
| | | Hemisphere
19. Xantharpya | 1 |All Africa |Oriental, Austro-
| | | Malayan
20. Cynopterus | 1 |Tropical Africa |Oriental
21. _Epomophorus_ | 6 |Tropical Africa and |
| | Abyssinia |
22. _Hypsignathus_ | 1 |W. Africa |
| | |
RHINOLOPHIDÆ. | | |
| | |
23. Rhinolophus | 6 |Africa and Madagascar |Warmer parts of
| | | Eastern Hemisphere
24. _Macronycterys_| 1 |W. Africa |
25. Phyllorhina | 4 |Tropical Africa |Indo-Malaya, Austro-
| | | Malaya
26. Asellia | 1 |Nubia |Indo-Malaya, Austro-
| | | Malaya
27. Megaderma | 1 |Senegal, Upper Nile |Oriental, Moluccas
28. Nycteris | 3 |All Africa |Java
| | |
VESPERTILIONIDÆ. | | |
| | |
29. Vespertilio | 14 |Africa and Madagascar |Cosmopolite
30. Kerivoula | 1 |S. Africa |Oriental
31. Miniopteris | 1 |S. Africa |Indo-Malaya
32. Nycticejus | 7 |Tropical Africa |India
33. Taphozous | 2 |Africa and Madagascar |Oriental, Austro-
| | | Malayan, Neotropical
| | |
NOCTILIONIDÆ. | | |
| | |
34. Nyctinomus | 1 |Madagascar |Oriental, American,
| | | S. Palæarctic
35. Molossus | 3 |Africa, Bourbon |Neotropical,
| | | S. Palæarctic
| | |
INSECTIVORA. | | |
MACROSCELIDIDÆ. | | |
| | |
36. _Macroscelides_| 2 |South and East Africa |N. Africa
37. _Petrodromus_ | 1 |Mozambique |
38. _Rhynchocyon_ | 1 |Mozambique |
| | |
ERINACEIDÆ. | | |
| | |
39. Erinaceus | 2 |Cen. and South Africa |Palæarctic, N. India
| | |
CENTETIDÆ. | | |
| | |
40. _Centetes_ | 2 |Madagascar and |
| | Mauritius |
41. _Hemicentetes_ | 2 |Madagascar |
42. _Ericulus_ | 2 |Madagascar |
43. _Oryzorictes_ | 1 |Madagascar |
44. _Echinops_ | 3 |Madagascar |
| | |
POTAMOGALIDÆ. | | |
| | |
45. _Potamogale_ | 1 |Old Calabar |
| | |
CHRYSOCHLORIDÆ. | | |
| | |
46. _Chrysochloris_| 3 |Cape to Mozambique |
| | |
SORICIDÆ. | | |
| | |
47. Sorex | 15 |All Africa and |Palæarc., Nearc., Ori
| | Madagascar |
| | |
CARNIVORA. | | |
FELIDÆ. | | |
| | |
48. Felis | 8 |All Africa |All reg. but
| | | Australian
49. Lynx[?] | 1 |N. and S. Africa |Palæarctic and
| | | Nearctic
50. Cynælurus | 1 |Cape of Good Hope |
| | |
CRYPTOPROCTIDÆ. | | |
| | |
51. _Cryptoprocta_ | 1 |Madagascar |
| | |
VIVERRIDÆ. | | |
| | |
52. Viverra | 1 |Tropical Africa |Oriental
53. Genetta | 4 |Tropical and S. Africa |S. Palæarctic
54. _Fossa_ | 2 |Madagascar |
55. _Poiana_ | 1 |W. Africa |
56. _Galidia_ | 3 |Madagascar |
57. _Nandinia_ | 1 |W. Africa |
58. _Galidictis_ | 2 |Madagascar |
59. Herpestes | 13 |All Africa |S. Europe, Oriental
60. _Athylax_ | 3 |S. and E. Africa(?) |
| | Madagascar |
61. Calogale | 9 |Tropical and S. Africa |Oriental
62. _Galerella_ | 1 |E. Africa |
63. _Ariela_ | 1 |S. Africa |
64. _Ichneumia_ | 4 |E. Africa, Senegal, |
| | S. Africa |
65. _Bdeogale_ | 3 |Tropical Africa |
66. _Helogale_ | 2 |E. and S. Africa |
67. _Cynictis_ | 3 |S. Africa |
68. _Rhinogale_ | 1 |E. Africa |
69. _Mungos_ | 3 |Tropical and S. Africa |
70. _Crossarchus_ | 1 |Tropical Africa |
71. _Eupleres_ | 1 |Madagascar |
72. _Suricata_ | 1 |S. Africa |
| | |
PROTELIDÆ. | | |
| | |
73. _Proteles_ | 1 |S. Africa |
| | |
HYÆNIDÆ. | | |
| | |
74. Hyæna | 3 |All Africa |S. Palæarctic, India
| | |
CANIDÆ. | | |
| | |
75. _Lycaon_ | 1 |S., Central, and |
| | E. Africa |
76. Canis | 5 |All Africa |Almost Cosmopolitan
77. _Megalotis_ | 1 |S. Africa |
| | |
MUSTELIDÆ. | | |
| | |
78. Mustela | 1 |Angola |Palæarctic, Nearctic
79. Gymnopus[?] | 1 |S. Africa |Oriental
80. Aonyx | 1 |S. and W. Africa |Oriental
81. _Hydrogale_ | 1 |S. Africa |
82. Mellivora | 2 |South and Tropical |India
| | Africa |
83. _Ictonyx_ | 2 |Tropical and S. Africa |
| | |
OTARIIDÆ. | | |
| | |
84. Arctocephalus | 1 |Cape of Good Hope |South Temperate Zone
| | |
SIRENIA. | | |
MANATIDÆ. | | |
| | |
85. Manatus | 1 |W. Africa |Tropical America
86. Halicore | 1 |E. Africa |Oriental and
| | | Australian
| | |
UNGULATA. | | |
EQUIDÆ. | | |
| | |
87. Equus | 3 |Tropical and S. Africa |Palæarctic
| | |
RHINOCEROTIDÆ. | | |
| | |
88. Rhinoceros | 4 |All Tropical and |Oriental
| | S. Africa |
| | |
HIPPOPOTAMIDÆ. | | |
| | |
89. _Hippopotamus_ | 2 |Great Rivers of Africa |
| | |
SUIDÆ. | | |
| | |
90. _Potamochoerus_| 3 |Tropical Africa and |
| | Madg. |
91. _Phacochoerus_ | 2 |Abyssinia to Caffraria |
| | |
TRAGULIDÆ. | | |
| | |
92. _Hyomoschus_ | 1 |W. Africa |
| | |
CAMELOPARDALIDÆ. | | |
| | |
93._Camelopardalis_| 1 |All open country |
| | |
BOVIDÆ. | | |
| | |
94. Bubalus | 3 |Trop. and S. Africa |India
95. _Oreas_ | 2 |Africa S. of Sahara |
96. _Tragelaphus_ | 8 |Africa S. of Sahara |
97. Oryx | 3 |Arabian and African |S. Palæarctic
| | deserts |
98. Gazella | 12 |Africa N. of Equator |Palæarctic Deserts
| | and S. Africa |
99. _Æpyceros_ | 1 |S. E. Africa |
100. _Cervicapra_ | 4 |All Tropical Africa |
101. _Kobus_ | 6 |Pastures of all Africa |
102. _Pelea_ | 1 |South Africa |
103. _Nanotragus_ | 9 |Africa S. of Sahara |
104. _Neotragus_ | 1 |Abyssinia and |
| | N. E. Africa |
105. _Cephalophus_ | 22 |All tropical Africa |
106. _Hippotragus_ | 3 |Gambia, Central Africa |
| | to Cape |
107. _Alcephalus_ | 9 |All Africa |
108. _Catoblepas_ | 2 |Africa S. of Equator |
(Capra | 1 |Abyssinia, high) |Palæarctic genus
| | |
PROBOSCIDEA. | | |
| | |
ELEPHANTIDÆ. | | |
| | |
109. Elephas | 1 |Tropical and S. Africa |Oriental
| | |
HYRACOIDEA. | | |
HYRACIDÆ. | | |
| | |
110. Hyrax | 10 |Tropical and S. Africa |Syria
| | |
RODENTIA. | | |
MURIDÆ. | | |
| | |
111. Mus | 26 |All Africa |E. Hemis. excl.
| | | Oceania
112. _Lasiomys_ | 1 |W. Africa |
113. Acanthomys | 4 |Tropical Africa |India
114. _Cricetomys_ | 1 |Tropical Africa |
115. _Saccostomus_ | 2 |Mozambique |
116. _Dendromys_ | 2 |S. Africa |
117. _Nesomys_ | 1 |Madagascar |
118. _Steatomys_ | 2 |East and S. Africa |
119. _Pelomys_ | 1 |Mozambique |
120. _Otomys_ | 6 |S. and E. Africa |
121. Meriones | 14 |Africa |Palæarctic, India
122. _Malacothrix_ | 2 |S. Africa |
123. _Mystromys_ | 1 |S. Africa |
124._Brachytarsomys_| 1 |Madagascar |
125. _Hypogeomys_ | 1 |Madagascar |
126. _Lophiomys_ | 1 |S. Arabia and |
| | N. E. Africa |
| | |
SPALACIDÆ. | | |
| | |
127. Rhizomys | 4 |Abyssinia |Oriental to Malacca
128. _Bathyerges_ | 1 |S. Africa |
129. _Georychus_ | 6 |E. Central, and |
| | S. Africa |
130. _Heliophobius_ | 1 |Mozambique |
| | |
DIPODIDÆ. | | |
| | |
131. Dipus | 7 |N. and Central Africa |Central Palæarctic
132. _Pedetes_ | 1 |S. Af. to Mozambique |
| | and Angola |
| | |
MYOXIDÆ. | | |
| | |
133. Myoxus | 1 |Africa to Cape |Palæarctic
| | |
SCIURIDÆ. | | |
| | |
134. Sciurus | 18 |All woody districts of |All regions but
| | Africa | Australia
135. _Anomalurus_ | 5 |W. Africa and Fernando |
| | Po. |
| | |
OCTODONTIDÆ. | | |
| | |
136. _Pectinator_ | 1 |Abyssinia |
| | |
ECHIMYIDÆ. | | |
| | |
137. _Petromys_ | 1 |S. Africa |
138. _Aulacodes_ | 1 |W., E., and S. Africa |
| | |
HYSTRICIDÆ. | | |
| | |
139. Hystrix | 1 |Africa to Cape |S. Palæarctic Oriental
140. Atherura | 1 |W. Africa |Palæarctic
| | |
LEPORIDÆ. | | |
| | |
141. Lepus | 5 |East and South Africa |All regions but
| | | Australian
| | |
EDENTATA. | | |
MANIDIDÆ. | | |
| | |
142. Manis | 4 |Sennaar to W. Africa |Oriental
| | and Cape |
| | |
ORYCTEROPODIDÆ. | | |
| | |
143. _Orycteropus_ | 2 |N. E. Africa to Nile |
| | Sources, and S. Africa|
_BIRDS._
PASSERES. | | |
TURDIDÆ. | | |
| | |
1. Turdus | 13 |The whole reg. (excl. |Almost Cosmopolite
| | Madagas.) |
2. Monticola | 2 |S. Africa |Palæarctic and
| | | Oriental
3. _Chætops_ | 3 |S. Africa |
4. _Bessonornis_ | 15 |The whole region |Palestine
| | |
SYLVIIDÆ. | | |
| | |
5. { _Drymoeca_ | 70 |The whole region |Palestine
6. { Cisticola | 13 |The whole region |Palæarc., Orien.,
{ | | | Austral.
7. { Sphenoeacus | 1 |S. Africa |Australian
8. { _Camaroptera_| 5 |Africa |
| | |
9. { Acrocephalus | 8 |The whole region |Palæarc., Orien.,
{ | | | Austral.
10. { Bradyptetus | 8 |Abyssinia and S. Africa|S. Europe, Palestine
11. { _Catriscus_ | 3 |All Africa |
12. { _Bernieria_ | 1 |Madagascar |
13. { _Ellisia_ | 1 |Madagascar |
14. { _Mystacornis_| 1 |Madagascar |
| | |
15. { Phylloscopus | 1 |S. Africa |Palæarctic, Oriental
16. { _Eremomela_ | 16 |All Africa |
17. { _Eroessa_ | 1 |Madagascar |
18. { Hypolais | 2 |S. Africa |Palæarctic, Oriental
| | |
19. { Aedon | 8 |E. and S. Africa |Palæarctic
20. { Sylvia | 3 |N. E. Africa, Gambia, |Palæarctic, Oriental
{ | | Cape Verd Ids. |
21. { Curruca | 2 |S. Africa |Palæarctic
| | |
22. { Ruticilla | 2 |Abyssinia and Senegal |Palæarctic, Oriental
23. { Cyanecula | 2 |N. E. Africa |Palæarctic
| | |
24. { Copsychus | 2 |Madagascar and |Oriental
{ | | Seychelle Ids. |
25. { Thamnobia | 7 |All Africa |Oriental
26. {_Cercotrichas_| 2 |W. and N. E. Africa |
27. { _Poeoptera_ | 1 |W. Africa |
28. { _Gervasia_ | 2 |Madagascar and |
{ | | Seychelle Ids. |
29. { Dromolæa | 13 |All Africa |S. Palæarctic, India
30. { Saxicola | 14 |Central, E. and |Palæarctic, India
{ | | S. Africa |
31. { Cercomela | 3 |N. E. Africa |Palestine, N. W. India
32. { Pratincola | 7 |Africa and Madagascar |Palæarctic, Oriental
| | |
TIMALIIDÆ. | | |
| | |
33. Chatarrhæa | 1 |Abyssinia |Oriental, Palestine
34. _Crateropus_ | 17 |All Africa |N. Africa, Persia
35. _Hypergerus_ | 1 |W. Africa |
36. _Cichladusa_ | 3 |W. and E. Africa |
37. _Alethe_ | 4 |W. Africa |
38. _Oxylabes_ | 2 |Madagascar |
| | |
CINCLIDÆ.[?] | | |
| | |
39. _Mesites_ | 1 |Madagascar |
| | |
TROGLODYTIDÆ.[?] | | |
| | |
40. Sylvietta | 2 |Central, E. and |
| | S. Africa |
| | |
SITTIDÆ. | | |
| | |
41. _Hypherpes_ | 1 |Madagascar |
| | |
PARIDÆ. | | |
| | |
42. Parus | 5 |All Africa |Palæarc., Orien.,
| | | Nearc.
43. _Parisoma_ | 5 |All Africa |
44. Ægithalus | 4 |W., Central, and |Palæarctic
| | S. Africa |
45. _Parinia_ | 1 |W. Africa, Prince's |
| | Island |
| | |
PYCNONOTIDÆ. | | |
| | |
46. Pycnonotus | 8 |All Africa |S. Palæarctic,
| | | Oriental
47._Phyllastrephus_| 4 |W. and S. Africa |
48. Hypsipetes | 4 |Madagascar and |Oriental
| | Mascarene Ids. |
49. _Tylas_ | 1 |Madagascar |
50. Criniger | 14 |W. and S. Africa |Oriental
51. _Ixonotus_ | 8 |W. Africa |
52. _Andropadus_ | 9 |Africa and Madagascar |
53. _Lioptilus_ | 1 |S. Africa |
| | |
ORIOLIDÆ. | | |
| | |
54. Oriolus | 10 |All Africa |Palæarctic, Oriental
55. _Artamia_[?] | 3 |Madagascar |
56._Cyanolanius_[?]| 1 |Madagascar |
| | |
CAMPEPHAGIDÆ. | | |
| | |
57. _Lanicterus_ | 5 |All Africa |
58. _Oxynotus_ | 2 |Mauritius and Bourbon |
59. Campephaga | 5 |The whole region |Celebes to New
| | | Caledonia
| | |
DICRURIDÆ. | | |
| | |
60. Dicrurus | 11 |The whole region |Oriental, Australian
| | |
MUSCICAPIDÆ. | | |
| | |
61. Butalis | 3 |All Africa |Palæarctic,
| | | N. Oriental
62. Muscicapa | 10 |All tropical Africa |Palæarctic
63. Alseonax | 4 |S. Africa |Oriental
64. _Newtonia_ | 1 |Madagascar |
65. _Hyliota_ | 2 |W. Africa |
66. _Erythrocercus_| 2 |Tropical Africa |
67. _Artomyias_ | 2 |W. Africa |
68. _Pseudobias_ | 1 |Madagascar |
69. _Smithorius_ | 2 |W. and S. Africa |
70. _Megabias_ | 1 |W. Africa |
71. _Cassinia_ | 2 |W. Africa |
72. _Bias_ | 1 |Tropical Africa |
73. _Elminia_ | 2 |Tropical Africa |
74. _Platystira_ | 12 |All Africa |
75. Tchitrea | 18 |The whole region |Oriental
76. _Pogonocichla_ | 1 |S. Africa |
77. _Bradyornis_ | 7 |All Africa |
| | |
LANIIDÆ. | | |
| | |
78. _Parmoptila_[?]| 1 |W. Africa. |
79. _Calicalicus_ | 1 |Madagascar |
80. Lanius | 15 |All Africa |Palæarc., Orien.,
| | | Nearc.
81. _Hypocolius_ | 1 |Abyssinia |
82. _Corvinella_ | 1 |S. and W. Africa |
83. _Urolestes_ | 1 |S. Africa |
84. _Fraseria_ | 2 |W. Africa |
85. _Hypodes_ | 1 |W. Africa |
86. _Cuphoterus _ | 1 |Prince's Island |
87. _Nilaus_ | 1 |All Africa |
88. _Prionops_ | 9 |All Africa |
89. _Eurocephalus_ | 2 |N. E. and S. Africa |
90. _Chaunonotus_ | 1 |W. Africa |
91. _Vanga_ | 4 |Madagascar |
92. _Laniarius_ | 38 |All Africa, |
| | Madagascar[?] |
93. _Meristes_ | 2 |W. and S. E. Africa |
94. _Nicator_ | 1 |E. Africa |
95. _Telephonus_ | 10 |All Africa |N. Africa
| | |
CORVIDÆ. | | |
| | |
96. _Ptilostomus_ | 2 |W. and E. Africa |
97. Corvus | 7 |All Africa and |Cosmop., excl.
| | Madagascar | S. Amer.
98. _Corvultur_ | 2 |N. E. to S. Africa |
99. _Picathartes_ | |W. Africa |
(Fregilus | 1 |Abyssinia) |Palæarctic genus
| | |
NECTARINIIDÆ. | | |
| | |
100. _Nectarinia_ | 55 |The whole region |
101. _Promerops_ | 1 |S. Africa |
102._Cinnyricinclus_| 4 |W. Africa |
103. _Neodrepanis_ | 1 |Madagascar |
| | |
DICÆIDÆ. | | |
| | |
104. Zosterops | 23 |The whole region |Oriental and
| | | Australian
105. _Pholidornis_ | 1 |W. Africa |
| | |
HIRUNDINIDÆ. | | |
| | |
106. Hirundo | 17 |The whole region |Cosmopolite
107. _Psalidoprogne_| 10 |The whole region |
108. _Phedina_ | 2 |Madagascar and |
| | Mauritius |
109. Petrochelidon | 1 |S. Africa |Neotropical
110. Chelidon | 1 |Bogos-land |Palæarctic, Oriental
111. Cotyle | 6 |All Africa |Palæarctic, Oriental
112. _Waldenia_ | 1 |W. Africa |
| | |
FRINGILLIDÆ. | | |
| | |
113. Dryospiza | 8 |All Africa |S. Palæarctic
114. Chlorospiza | 4 |Abyssinia to Cape |Palæarctic
115. Passer | 18 |All Africa |Palæarctic, Oriental
116. _Crithagra_ | 12 |All Africa |N. Africa, Syria
117. _Ligurnus_ | 2 |W. Africa |
(Erythrospiza | 1 |Nubia, Arabia) |S. Palæarctic genus
118. Pinicola[?] | 1 |Cameroons, W. Africa |N. Temperate genus
119. _Fringillaria_ | 9 |All Africa |South Palæarctic
| | |
PLOCEIDÆ. | | |
| | |
120. _Textor_ | 5 |All Africa |
121. _Hyphantornis_ | 32 |Tropical and S. Africa |
122. _Symplectes_ | 8 |Tropical and S. Africa |
123. _Malimbus_ | 9 |W. and E. Africa |
124. Ploceus | 2 |W. and E. Africa |Oriental
125. _Nelicurvius_ | 1 |Madagascar |
126. _Foudia_ | 11 |Tropical Africa, |
| | Madagascar, &c. |
127. _Sporopipes_ | 1 |Tropical and S. Africa |
128. _Pyromelana_ | 12 |Tropical and S. Africa |
129. _Philetærus_ | 1 |S. Africa |
130. _Nigrita_ | 7 |W. and N. E. Africa |
131. _Plocepasser_ | 4 |E. and S. Africa |
132. _Vidua_ | 6 |Tropical and S. Africa |
133. _Colliuspasser_| 9 |Tropical and S. Africa |
134. _Chera_ | 1 |S. Africa |
135. _Spermospiza_ | 2 |W. Africa |
136. _Pyrenestes_ | 6 |Tropical and S. Africa |
137. Estrilda | 16 |Tropical and S. Africa |Oriental
138. _Pytelia_ | 20 |Tropical and S. Africa |
139. _Hypargos_ | 2 |E. Africa, Madagascar |
140. _Amadina_ | 6 |Tropical and S. Africa |
141. _Spermestes_ | 7 |The whole region |
142. _Amauresthes_ | 1 |E. and W. Africa |
143. _Hypochera_ | 2 |Tropical and S. Africa |
| | |
STURNIDÆ. | | |
| | |
144. _Dilophus_ | 1 |S. Africa, Loanda, |
| | Sennaar |
145. _Buphaga_ | 2 |Trop. and S. Africa |
| | ([?] a family) |
146. _Euryceros_ | 1 |Madagascar ([?] a |
| | family) |
147. _Juida_ | 5 |Tropical and S. Africa |
148. _Lamprocolius_ | 16 |Tropical and S. Africa |
149._Cinnyricinclus_| 2 |Tropical and S. Africa |
150. _Onychognathus_| 2 |W. Africa |
151. _Spreo_ | 5 |Tropical and S. Africa |
152. _Amydrus_ | 5 |N. E. Africa |Palestine
153. _Hartlaubius_ | 1 |Madagascar |
154. _Falculia_ | 1 |Madagascar |
155. _Fregilupus_ | 1 |Bourbon |
| | |
ALAUDIDÆ. | | |
| | |
156. Alauda | 3 |Abyssinia and |Palæarctic, Indian
| | S. W. Africa |
157. _Spizocorys_ | 1 |South Africa |
158. Galerida | 4 |North of tropical |Palæarctic, Indian
| | Africa |
159. _Calendula_ | 2 |Abyssinia, S. Africa |
(Melanocorypha | 1 |Abyssinia) |Palæarctic genus
160. Certhilauda | 3 |South Africa |S. Europe
161. Alaemon | 3 |South Africa |S. Palæarctic
162. _Heterocorys_ | 1 |South Africa |
163. Mirafra | 10 |South Africa, |Oriental, Australian
| | Madagascar |
164. Ammomanes | 4 |African deserts |S. Palæarctic, Indian
165. _Megalophonus_ | 5 |Tropical and S. Africa |
166. _Tephrocorys_ | 2 |S. Africa |
167. Pyrrhulauda | 6 |Tropical and S. Africa |Oriental, Canary
| | | Islands
| | |
MOTACILLIDÆ. | | |
| | |
168. Budytes | 8 |The whole region |Palæarctic, Oriental,
| | |Australian
| | |
169. Anthus | 10 |Tropical and S. Africa |All regions, exc.
| | | Australia
170. _Macronyx_ | 4 |Tropical and S. Africa |
| | |
PITTIDÆ. | | |
| | |
171. Pitta | 1 |W. Africa |Oriental, Australian
| | |
PAICTIDÆ. | | |
| | |
172. _Philepitta_ | 2 |Madagascar |
| | |
PICARIÆ. | | |
| | |
PICIDÆ. | | |
| | |
173. _Verreauxia_ | 1 |W. Africa |
174. _Dendropicus_ | 14 |Tropical and S. Africa |
175. _Campethera_ | 14 |Tropical and S. Africa |
176. _Geocolaptes_ | 1 |South Africa |
| | |
YUNGIDÆ. | | |
| | |
177. Yunx | 1 |N. E. Africa, S. Africa|Palæarctic
| | |
INDICATORIDÆ. | | |
| | |
178. Indicator | 8 |Tropical and S. Africa |Oriental
| | |
MEGALÆMIDÆ. | | |
| | |
179._Pogonorhynchus_| 14 |Tropical and S. Africa |
180. _Buccanodon_ | 1 |West Africa |
181. _Stactolæma_ | 1 |West Africa |
182. _Barbatula_ | 9 |West and South Africa |
183. _Xylobucco_ | 3 |West and South Africa |
184. _Gymnobucco_ | 3 |West Africa |
185. _Trachyphonus_ | 6 |Tropical and South |
| | Africa |
| | |
MUSOPHAGIDÆ. | | |
| | |
186. _Musophaga_ | 2 |West Africa |
187. _Turacus_ | 10 |Tropical and S. Africa |
188. _Schizorhis_ | 6 |Tropical and S. Africa |
| | |
COLIIDÆ. | | |
| | |
189. _Colius_ | 7 |Tropical and S. Africa |
| | |
CUCULIDÆ. | | |
| | |
190. _Ceuthmochares_| 2 |Africa and Madagascar |
191. _Coua_ | 9 |Madagascar |
192._Cochlothraustes_ 1 |Madagascar |
193. _Centropus_ | 8 |Africa and Madagascar |Oriental, Australian
194. Cuculus | 10 |Africa and Madagascar |Palæarc., Orien.,
| | | Austral.
195. Chrysococcyx | 7 |Tropical and S. Africa |Oriental, Australian
196. Coccystes | 6 |Tropical and S. Africa |S. Palæarctic,
| | | Oriental
| | |
LEPTOSOMIDÆ. | | |
| | |
197. _Leptosomus_ | 1 |Madagascar |
| | |
CORACIIDÆ. | | |
| | |
198. Coracias | 5 |Africa and Madagascar |S. Palæarctic,
| | | Oriental
199. Eurystomus | 3 |Africa and Madagascar |Oriental, Australian
200. _Atelornis_ | 2 |Madagascar |
201._Brachypteracias_ 1 |Madagascar |
202. _Geobiastes_ | 1 |Madagascar |
| | |
MEROPIDÆ. | | |
| | |
203. Merops | 11 |Africa and Madagascar |S. Palæar., Orien.,
| | | Austral.
204. _Melittophagus_| 5 |Tropical and S. Africa |
| | |
TROGONIDÆ. | | |
| | |
205. _Apaloderma_ | 2 |Tropical and S. Africa |
| | |
ALCEDINIDÆ. | | |
| | |
206. Alcedo | 2 |W. Africa, Abyssinia, |Palæar., Orien.,
| | Natal | Austral.
207. _Corythornis_ | 3 |Africa and Madagascar |
208. Ceryle | 1 |W. Africa, Abyssinia, |American, Palæarctic
| | Natal |
209. _Myioceyx_ | 2 |West Africa |
210. _Ispidina_ | 4 |Africa and Madagascar |
211. Halcyon | 10 |Africa, Prince's Is., |S. Palæar., Orien.,
| | St. Thomé | Austral.
| | |
BUCEROTIDÆ. | | |
| | |
212. Berenicornis | 1 |West Africa |Malaya
213. _Tockus_ | 12 |Tropical and S. Africa |
214. _Bycanistes_ | 6 |Tropical and S. Africa |
215. _Bucoreus_ | 2 |Tropical and S. Africa |
| | |
UPUPIDÆ. | | |
| | |
216. Upupa | 3 |Africa and Madagascar |S. Palæarctic,
| | | Oriental
| | |
IRRISORIDÆ. | | |
| | |
217. _Irrisor_ | 12 |Africa and Madagascar |
| | |
CAPRIMULGIDÆ. | | |
| | |
218. Caprimulgus | 18 |Africa and Madagascar |Palæarc., Orien.,
| | | Austral.
219. _Scortornis_ | 3 |Tropical Africa |
220. _Macrodipteryx_| 2 |W. Africa to Abyssinia |
221. _Cosmetornis_ | 1 |Tropical Africa to the |
| | Zambesi |
| | |
CYPSELIDÆ. | | |
| | |
222. Cypselus | 6 |The whole region |Palæarctic, Oriental
223. Collocalia | 1 |Mascarene Ids., |Oriental, Australian
| | Madagascar |
224. Chætura | 4 |Tropical Africa and |Cosmop., exc.
| | Madagascar | W. Palæarctic
| | |
PSITTACI. | | |
PALÆORNITHIDÆ. | | |
| | |
225. Palæornis | 3 |W. Africa to Abys. & |Oriental
| | Mauritius |
| | |
PSITTACIDÆ. | | |
| | |
226. _Coracopsis_ | 5 |Madagascar and |
| | Seychelle Ids. |
227. _Psittacus_ | 2 |W. Africa |
228. _Poeocephalus_ | 9 |Tropical and S. Africa |
229. _Agapornis_ | 4 |Tropical and S. Africa |
230. _Poliopsitta_ | 2 |Trop. Africa and |
| | Madagascar |
| | |
COLUMBÆ. | | |
COLUMBIDÆ. | | |
| | |
231. Treron | 6 |Africa and Madagascar |Oriental
232. _Alectrænas_ | 5 |Madagascar and Masc. |
| | Ids. (extct. in |
| | Mauritius and |
| | Rodriguez) |
233. Columba | 12 |Africa and Madagascar |Palæarctic, Oriental
234. _Oena_ | 1 |Tropical and S. Africa |
235. Turtur | 10 |Africa, Madagascar, |Palæarctic, Oriental
| | Comoro and Seychelle |
| | Islands |
236. _Aplopelia_ | 4 |Abyssinia, S. Africa |
| | and West African |
| | Islands |
237. _Chalcopelia_ | 3 |Tropical and S. Africa |
| | |
DIDIDÆ. (extinct) | | |
| | |
238. _Didus_ | 5 |Mascarene Islands |
| | |
GALLINÆ. | | |
PTEROCLIDÆ. | | |
| | |
239. Pterocles | 9 |Africa and Madagascar |S. Palæarctic, Indian
| | |
TETRAONIDÆ. | | |
| | |
240. _Ptilopachus_ | 1 |West Africa |
241. Francolinus | 30 |Africa and Madagascar |S. Palæarctic, Indian
242. _Peliperdix_ | 1 |West Africa |
243. _Margaroperdix_| 1 |Madagascar |
244. Coturnix | 2 |Tropical and S. Africa |Palæar., Orient.,
| | | Austral.
(Caccabis | 1 |Abyssinia) |Palæarctic genus
| | |
PHASIANIDÆ. | | |
| | |
245. _Phasidus_ | 1 |West Africa |
246. _Agelastes_ | 1 |West Africa |
247. _Acryllium_ | 1 |West Africa |
248. _Numida_ | 9 |Africa to Natal and |
| | Madagascar |
| | |
TURNICIDÆ. | | |
| | |
249. Turnix | 4 |S. Africa and |Palæarc., Orient.,
| | Madagascar | Austrl.
250. _Ortyxelos_ | 1 |Africa |
| | |
ACCIPITRES. | | |
VULTURIDÆ. | | |
| | |
251. Gyps | 2 |Africa, except W. |Palæarctic, Oriental
| | sub-region |
252. Pseudogyps | 1 |N. E. Africa to Senegal|Oriental
253. Otogyps | 1 |N. E. and S. Africa |Palæarctic, Oriental
254. _Lophogyps_ | 1 |N. E. and S. Africa and|
| | Senegal |
255. Neophron | 2 |Africa, excl. west |S. Palæarctic,
| | coast | Oriental
| | |
FALCONIDÆ. | | |
| | |
256. _Polyboroides_ | 2 |Africa and Madagascar |
257. Circus | 4 |Africa and Madagascar |Almost Cosmopolite
258. _Urotriorchis_ | 1 |W. Africa |
259. _Melierax_ | 5 |Africa, excl. west |
| | coast |
260. Astur | 5 |Africa and Madagascar |Almost Cosmopolite
261. _Nisoides_ | 1 |Madagascar |
262. _Eutriorchis_ | 1 |Madagascar |
263. Accipiter | 8 |Africa and Madagascar |Almost Cosmopolite
264. Buteo | 5 |Africa and Madagascar |Cosmop., excl.
| | | Austral.
265. Gypaëtus | 1 |N. E. and S. Africa |S. Palæarctic
266. Aquila | 5 |All Africa |Nearc., Palæarc.,
| | | Indian
267. Nisaëtus | 1 |W. Africa |S. Palæarctic,
| | | Oriental, Australia
268. Spizaëtus | 3 |All Africa |Neotropical, Oriental
| | | to N. Guinea
269. _Lophoætus_ | 1 |All Africa |
270. _Asturinula_ | 1 |Tropical Africa |
271. _Dryotriorchis_| 1 |W. Africa |
272. Circaëtus | 5 |All Africa |Palæarctic, Oriental
273. Butastur | 1 |N. E. Africa |Oriental to New Guinea
274. _Helotarsus_ | 2 |Tropical and S. Africa |
275. Haliæetus | 2 |The whole region |Cosmopolite, excl.
| | | Neotropical region
276. _Gypohierax_ | 1 |West and East Africa |
277. _Elanoides_ | 1 |West and N. E. Africa |
278. Milvus | 1 |The whole region |The Eastern Hemisphere
279. Elanus | 1 |Africa |India to Australia
280. Machærhamphus | 1 |S. W. Africa and |Malacca
| | Madagascar |
281. Pernis | 1 |S. Africa and |Palæarctic, Oriental
| | Madagascar |
282. Baza | 3 |Africa and Madagascar |India to N. Australia
283. Poliohierax | 1 |East Africa |Burmah
284. Falco | 4 |All Africa |Almost Cosmopolite
285. Cerchneis | 8 |The whole region |Almost Cosmopolite
| | |
SERPENTARIIDÆ. | | |
| | |
286. _Serpentarius_ | 1 |The greater part of |
| | Africa |
| | |
PANDIONIDÆ. | | |
| | |
287. Pandion | 1 |All Africa |Cosmopolite
| | |
STRIGIDÆ. | | |
| | |
288. Athene | 5 |Africa and Madagascar, |Palæarctic, Oriental,
| | Rodriquez (extinct) | Australian
289. Bubo | 8 |Africa and Madagascar |Cosmopolite
290. _Scotopelia_ | 2 |West and S. Africa to |
| | Zambesi |
291. Scops | 3 |W. and S. Africa, |Almost Cosmopolite
| | Madagascar, Comoro |
| | Islands |
292. Syrnium | 2 |Africa |Palæarctic, Oriental,
| | | American
293. Asio | 1 |N. E. and S. Africa |Cosmopolite
294. Strix | 4 |Africa and Madagascar |Cosmopolite
_Peculiar or very Characteristic Genera of Wading or Swimming Birds._
GRALLÆ. | | |
RALLIDÆ. | | |
| | |
_Himantornis_ | 1 |West Africa |
Podica | 3 |Africa |Burmah
| | |
GLAREOLIDÆ. | | |
| | |
Cursorius | 8 |All Africa |S. Europe, India
| | |
OTIDIDÆ. | | |
| | |
Eupodotis | 16 |All Africa |India, Australia
| | |
GRUIDÆ. | | |
| | |
_Balearica_ | 2 |All Africa |
| | |
ARDEIDÆ. | | |
| | |
_Balæniceps_ | 1 |Upper Nile |
| | |
PLATALEIDÆ. | | |
| | |
_Scopus_ | 1 |Tropical and S. Africa |
| | |
ANSERES. | | |
ANATIDÆ. | | |
| | |
_Thalassornis_| 1 |South Africa |
| | |
STRUTHIONES. | | |
STRUTHIONIDÆ. | | |
| | |
295. _Struthio_ | 2 |All Africa |Syria
| | |
ÆPYORNITHIDÆ. | |(Extinct) |
| | |
296. _Æpyornis_ | 3[?]|Madagascar |
{314}CHAPTER XII.
THE ORIENTAL REGION.
This region is of comparatively small extent, but it has a very diversified
surface, and is proportionately very rich. The deserts on the north-west of
India are the debatable land that separates it from the Palæarctic and
Ethiopian regions. The great triangular plateau which forms the peninsula
of India is the poorest portion of the region, owing in part to its arid
climate and in part to its isolated position; for there can be little doubt
that in the later Tertiary period it was an island, separated by an arm of
the sea (now forming the valleys of the Ganges and Indus) from the
luxuriant Himalayan and Burmese countries. Its southern extremity, with
Ceylon, has a moister climate and more luxuriant vegetation, and exhibits
indications of a former extension southwards, with a richer and more
peculiar fauna, partly Malayan and partly Mascarene in its character. The
whole southern slopes of the Himalayas, with Burmah, Siam and Western
China, as well as the Malay peninsula and the Indo-Malay islands, are
almost everywhere covered with tropical forests of the most luxuriant
character, which abound in varied and peculiar forms of vegetable and
animal life. The flora and fauna of this extensive district are essentially
of one type throughout; yet it may be usefully divided into the
Indo-Chinese and the Malayan sub-regions, as each possesses a number of
peculiar or characteristic animals. The former sub-region, besides having
many tropical and sub-tropical types of its own, also possesses a large
number of peculiarly modified temperate forms on the mountain ranges of its
northern boundary, which are wholly wanting in the Malayan sub-region. The
Philippine islands are best classed with the Indo-Malay group, although
they are strikingly deficient in many Malayan types, and exhibit an
approach to the Celebesian division of the Austro-Malay sub-region.
[Illustration: ORIENTAL REGION]
{315}_Zoological Characteristics of the Oriental Region._--The Oriental
Region possesses examples of 35 families of Mammalia, 71 of Birds, 35 of
Reptiles, 9 of Amphibia, and 13 of Fresh-water Fishes. Of these 163
families, 12 are peculiar to the region; namely, Tarsiidæ, Galeopithecidæ,
and Tupaiidæ among Mammalia, while Æluridæ, though confined to the higher
Himalayas, may perhaps with more justice be claimed by the Palæarctic
region; Liotrichidæ, Phyllornithidæ, and Eurylæmidæ among birds;
Xenopeltidæ (extending, however, to Celebes), Uropeltidæ, and Acrochordidæ
among reptiles; Luciocephalidæ, Ophiocephalidæ and Mastacembelidæ among
fresh-water fishes. A number of other families are abundant, and
characteristic of the region; and it possesses many peculiar and
characteristic genera, which must be referred to somewhat more in detail.
_Mammalia._--The Oriental region is rich in quadrumana, and is especially
remarkable for its orang-utans and long-armed apes (_Simia_, _Hylobates_,
and _Siamanga_); its abundance of monkeys of the genera _Presbytes_ and
_Macacus_; its extraordinary long-nosed monkey (_Presbytes nasalis_); its
Lemuridæ (_Nycticebus_ and _Loris_); and its curious genus _Tarsius_,
forming a distinct family of lemurs. All these quadrumanous genera are
confined to it, except _Tarsius_ which extends as far as Celebes. It
possesses more than 30 genera of bats, which are enumerated in the lists
given at the end of this chapter. In Insectivora it is very rich, and
possesses several remarkable forms, such as the flying lemur
(_Galeopithecus_); the squirrel-like Tupaiidæ consisting of three genera;
and the curious _Gymnura_ allied to the hedgehogs. In Carnivora, it is
especially rich in many forms of civets (Viverridæ), possessing 10 peculiar
genera, among which _Prionodon_ and _Cynogale_ are remarkable; numerous
Mustelidæ, of which _Gymnopus_, _Mydaus_, _Aonyx_ and _Helictis_ are the
most conspicuous; _Ælurus_, a curious animal, cat-like in appearance but
{316}more allied to the bears, forming a distinct family of Carnivora, and
confined to the high forest-districts of the Eastern Himalayas and East
Thibet; _Melursus_ and _Helarctos_, peculiar forms of bears; _Platanista_,
a dolphin peculiar to the Ganges and Indus. Among Ruminants it has the
beautiful chevrotain, forming the genus _Tragulus_ in the family Tragulidæ;
with one peculiar genus and three peculiar sub-genera of true deer. The
Antilopinæ and Caprinæ are few, confined to limited districts and not
characteristic of the region; but there are everywhere wild cattle of the
genera _Bibos_ and _Bubalus_, which, with species of _Rhinoceros_ and
_Elephas_, form a prominent feature in the fauna. The Rodents are less
developed than in the Ethiopian region, but several forms of squirrels
everywhere abound, together with some species of porcupine; and the
Edentata are represented by the scaly manis.
_Birds._--The families and genera of birds which give a character to
Oriental lands, are so numerous and varied, that we can here only notice
the more prominent and more remarkable. The Timaliidæ, represented by the
babblers (_Garrulax_, _Pomatorhinus_, _Timalia_, &c.), are almost
everywhere to be met with, and no less than 21 genera are peculiar to the
region; the elegant fork-tailed _Enicurus_ and rich blue _Myiophonus_,
though comparatively scarce, are characteristic of the Malayan and
Indo-Chinese faunas; the elegant little "hill-tits" (Liotrichidæ) abound in
the same part of the region; the green bulbuls (_Phyllornis_) are found
everywhere; as are various forms of Pycnonotidæ, the black and crimson
"minivets" (_Pericrocotus_), and the glossy "king-crows" (_Dicrurus_);
_Urocissa_, _Platylophus_ and _Dendrocitta_ are some of the interesting and
characteristic forms of the crow family; sun-birds (Nectariniidæ) of at
least three genera are found throughout the region, as are the beautiful
little flower-peckers (Dicæidæ), and some peculiar forms of weaver-birds
(_Ploceus_ and _Munia_). Of the starling family, the most conspicuous are
the glossy mynahs (_Eulabes_). The swallow-shrikes (_Artamus_) are very
peculiar, as are the exquisitely coloured pittas (Pittidæ), and the gaudy
broad-bills (Eurylæmidæ). Leaving the true Passeres, we find woodpeckers,
barbets, and cuckoos everywhere, often of peculiar and {317}remarkable
forms; among the bee-eaters we have the exquisite _Nyctiornis_ with its
pendent neck-plumes of blue or scarlet; brilliant kingfishers and strangely
formed hornbills abound everywhere; while brown-backed trogons with red and
orange breasts, though far less frequent, are equally a feature of the
Ornithology. Next we have the frog-mouthed goatsuckers (_Battrachostomus_),
and the whiskered swifts (_Dendrochelidon_), both wide-spread, remarkable,
and characteristic groups of the Oriental region. Coming to the parrot
tribe, we have only the long-tailed _Palæornis_ and the exquisite little
_Loriculus_, as characteristic genera. We now come to the pigeons, among
which the fruit-eating genera _Treron_ and _Carpophaga_ are the most
conspicuous. The gallinaceous birds offer us some grand forms, such as the
peacocks (_Pavo_); the argus pheasants (_Argusianus_); the fire-backed
pheasants (_Euplocamus_); and the jungle-fowl (_Gallus_), all strikingly
characteristic; and with these we may close our sketch, since the birds of
prey and the two Orders comprising the waders and swimmers offer nothing
sufficiently remarkable to be worthy of enumeration here.
_Reptiles._--Only the more abundant and characteristic groups will here be
noticed. In the serpent tribe, the Oligodontidæ, a small family of
ground-snakes; the Homalopsidæ, or fresh-water snakes; the Dendrophidæ, or
tree-snakes; the Dryiophidæ, or whip-snakes; the Dipsadidæ, or nocturnal
tree-snakes; the Lycodontidæ or fanged ground-snakes; the Pythonidæ, or
rock-snakes; the Elapidæ, or venomous colubrine snakes (including the
"cobras"); and the Crotalidæ, or pit-vipers, are all abundant and
characteristic, ranging over nearly the whole region, and presenting a
great variety of genera and species. Among lizards, the Varanidæ or
water-lizards; the Scincidæ or "scinks;" the Geckotidæ, or geckoes; and the
Agamidæ, or eastern iguanas; are the most universal and characteristic
groups. Among crocodiles the genus _Crocodilus_ is widely spread,
_Gavialis_ being characteristic of the Ganges. Among Chelonia, or shielded
reptiles, forms of fresh-water Testudinidæ and Trionychidæ (soft tortoises)
are tolerably abundant.
_Amphibia._--The only abundant and characteristic groups of {318}this class
are toads of the family Engystomidæ; tree-frogs of the family Polypedatidæ;
and several genera of true frogs, Ranidæ.
_Fresh-water Fishes._--The more remarkable and characteristic fishes
inhabiting the fresh waters of the Oriental region belong to the following
families: Nandidæ, Labyrinthici, Ophiocephalidæ, Siluridæ, and Cyprinidæ;
the last being specially abundant.
The sketch here very briefly given, must be supplemented by an examination
of the tables of distribution of the genera of all the Mammalia and Birds
inhabiting the region. We will now briefly summarize the results.
_Summary of the Oriental Vertebrata._--The Oriental region possesses
examples of 163 families of Vertebrata of which 12 are peculiar, a
proportion of a little more than one-fourteenth of the whole.
Out of 118 genera of Mammalia 54 seem to be peculiar to the region, equal
to a proportion of 9/20 or a little less than half. Of Land-Birds there are
342 genera of which 165 are peculiar, bringing the proportion very close to
a half.
In the Ethiopian region the proportion of peculiar forms both of Mammalia
and Birds is greater; a fact which is not surprising when we consider the
long continued isolation of the latter region--an isolation which is even
now very complete, owing to the vast extent of deserts intervening between
it and the Palæarctic region; while the Oriental and Palæarctic were,
during much of the Tertiary epoch, hardly separable.
_Insects._
_Lepidoptera._--We can only glance hastily at the more prominent features
of the wonderfully rich and varied butterfly-fauna of the Oriental region.
In the first family Danaidæ, the genera _Danais_ and _Euploea_ are
everywhere abundant, and the latter especially forms a conspicuous feature
in the entomological aspect of the country; the large "spectre-butterflies"
(_Hestia_) are equally characteristic of the Malayan sub-region. Satyridæ,
though abundant are not very remarkable, _Debis_, _Melanitis_, _Mycalesis_,
and _Ypthima_ being the most characteristic {319}genera. Morphidæ are well
represented by the genera _Amathusia_, _Zeuxidia_, _Discophora_, and
_Thaumantis_, some of the species of which almost equal the grand South
American Morphos. The Nymphalidæ furnish us with a host of characteristic
genera, among the most remarkable of which are, _Terinos_, _Adolias_,
_Cethosia_, _Cyrestis_, _Limenitis_, and _Nymphalis_, all abounding in
beautiful species. Among the Lycænidæ are a number of fine groups, among
which we may mention _Ilerda_, _Myrina_, _Deudoryx_, _Aphneus_, _Iolaus_,
and _Amblypodia_, as characteristic examples. The Pieridæ furnish many fine
forms, such as _Thyca_, _Iphias_, _Thestias_, _Eronia_, _Prioneris_, and
_Dercas_, the last two being peculiar. The Papilionidæ are unsurpassed in
the world, presenting such grand genera as _Teinopalpus_ and _Bhutanitis_;
the yellow-marked _Ornithopteræ_; the superb "Brookiana;" the elegant
_Leptocercus_; and _Papilios_ of the "Coon," "Philoxenus," "Memnon,"
"Protenor," and especially the 'green-and-gold-dusted' "Paris" groups.
The Moths call for no special observations, except to notice the existence
in Northern India of a number of forms which resemble in a striking manner
some of the most remarkable of the above mentioned groups of the genus
_Papilio_, especially the "Protenor" group, which there is reason to
believe protected by a peculiar smell or taste like the _Heliconias_ and
Danaidæ.
_Coleoptera._--The most characteristic Oriental form of the Cicindelidæ or
tiger beetles, is undoubtedly the elegant genus _Collyris_, which is found
over the whole region and is almost confined to it. Less abundant, but
equally characteristic, is the wingless ant-like _Tricondyla_. Two small
genera _Apteroessa_ and _Dromicidia_ are confined to the Indian Peninsula,
while _Therates_ only occurs in the Malayan sub-region.
The Carabidæ, or ground carnivorous beetles, are so numerous that we can
only notice a few of the more remarkable and characteristic forms. The
wonderful _Mormolyce_ of the Indo-Malay sub-region, stands pre-eminent for
singularity in the entire family. _Thyreopterus_, _Orthogonius_,
_Catascopus_, and _Pericallus_ are very characteristic forms, as well as
_Planetes_ and {320}_Distrigus_, the latter having a single species in
Madagascar. There are 80 genera of this family peculiar to the region, 10
of which have only been found in Ceylon.
Among the Lucanidæ, or stag-beetles, _Lucanus_, _Odontolabris_, and
_Cladognathus_ are the most characteristic forms. Sixteen genera inhabit
the region, of which 7 are altogether peculiar, while three others only
extend eastward to the Austro-Malayan sub-region.
The beautiful Cetoniidæ, or rose-chafers, are well represented by
_Rhomborhina_, _Heterorhina_, _Clinteria_, _Macronota_, _Agestrata_,
_Chalcothea_ and many fine species of _Cetonia_. There are 17 peculiar
genera, of which _Mycteristes_, _Phædimus_, _Plectrone_, and _Rhagopteryx_,
are Malayan; while _Narycius_, _Clerota_, _Bombodes_, and _Chiloloba_ are
Indian.
In Buprestidæ--those elongate metallic-coloured beetles whose elytra are
used as ornaments in many parts of the world--this region stands
pre-eminent, in its gigantic _Catoxantha_, its fine _Chrysochroa_, its
Indian _Sternocera_, its Malayan _Chalcophora_ and _Belionota_, as well as
many other beautiful forms. It possesses 41 genera, of which 14 are
peculiar to it, the rest being generally of wide range or common to the
Ethiopian and Australian regions.
In the extensive and elegant group of Longicorns, the Oriental region is
only inferior to the Neotropical. It possesses 360 genera, 25 of which are
Prionidæ, 117 Cerambicidæ, and 218 Lamiidæ;--about 70 per cent. of the
whole being peculiar. The most characteristic genera are _Rhaphidopodus_
and _Ægosoma_ among Prionidæ; _Neocerambyx_, _Euryarthrum_, _Pachyteria_,
_Acrocyrta_, _Tetraommatus_, _Chloridolum_, and _Polyzonus_ among
Cerambycidæ; and _Coelosterna_, _Rhytidophora_, _Batocera_, _Agelasta_, and
_Astathes_ among Lamiidæ.
Of remarkable forms in other families, we may mention the gigantic horned
_Chalcosoma_ among Scarabæidæ; the metallic _Campsosternus_ among
Elateridæ; the handsome but anomalous _Trictenotoma_ forming a distinct
family; the gorgeous _Pachyrhynchi_ of the Philippine Islands among
Curculionidæ; _Diurus_ {321}among Brenthidæ; with an immense number and
variety of Anthotribidæ, Heteromera, Malacoderma, and Phytophaga.
THE ORIENTAL SUB-REGIONS.
The four sub-regions into which we have divided the Oriental region, are
very unequal in extent, and perhaps more so in productiveness, but they
each have well-marked special features, and serve well to exhibit the main
zoological characteristics of the region. As they are all tolerably well
defined and their faunas comparatively well-known, their characteristics
will be given with rather more than usual detail.
_I. Hindostan, or Indian Sub-region._
This includes the whole peninsula of India from the foot of the Himalayas
on the north to somewhere near Seringapatam on the south, the boundary of
the Ceylonese sub-region being unsettled. The deltas of the Ganges and
Brahmaputra mark its eastern limits, and it probably reaches to about
Cashmere in the north-west, and perhaps to the valley of the Indus further
south; but the great desert tract to the east of the Indus forms a
transition to the south Palæarctic sub-region. Perhaps on the whole the
Indus may be taken as a convenient boundary. Many Indian naturalists,
especially Mr. Blyth and Mr. Blanford, are impressed with the relations of
the greater part of this sub-region to the Ethiopian region, and have
proposed to divide it into several zoological districts dependent on
differences of climate and vegetation, and characterized by possessing
faunas more or less allied either to the Himalayan or the Ethiopian type.
But these subdivisions appear far too complex to be useful to the general
student, and even were they proved to be natural, would be beyond the scope
of this work. I agree, however, with Mr. Elwes in thinking that they really
belong to local rather than to geographical distribution, and confound
"station" with "habitat." Wherever there is a marked diversity of surface
and vegetation the productions of a country will correspondingly differ;
the groups peculiar to forests, for example, will be absent from open
{322}plains or arid deserts. It happens that the three great Old World
regions are separated from each other by a debatable land which is chiefly
of a desert character; hence we must expect to find a resemblance between
the inhabitants of such districts in each region. We also find a great
resemblance between the aquatic birds of the three regions; and as we
generally give little weight to these in our estimate of the degree of
affinity of the faunas of different countries, so we should not count the
desert fauna as of equal weight with the more restricted and peculiar types
which are found in the fertile tracts,--in the mountains and valleys, and
especially in the primeval forests. The supposed preponderance of
exclusively Ethiopian groups of Mammalia and Birds in this, sub-region,
deserves however a close examination, in order to ascertain how far the
facts really warrant such an opinion.
_Mammalia._--The following list of the more important genera of Mammalia
which range over the larger part of this sub-region will enable naturalists
to form an independent judgment as to the preponderance of Ethiopian, or of
Oriental and Palæarctic types, in this, the most important of all the
classes of animals for geographical distribution.
RANGE OF THE GENERA OF MAMMALIA WHICH INHABIT THE SUB-REGION OF HINDOSTAN.
1. Presbytes Oriental only.
2. Macacus Oriental only.
3. Erinaceus Palæarctic genus.
4. Sorex Widely distributed.
5. Felis Almost Cosmopolitan.
6. Cynælurus Ethiopian and S. Palæarctic.
7. Viverra Ethiopian and Oriental to China and Malaya.
8. Viverricula Oriental only.
9. Paradoxurus Oriental only.
10. Herpestes Ethiopian, S. Palæarctic, and Oriental to Malaya.
11. Calogale Ethiopian, Oriental to Cambodja.
12. Tæniogale Oriental.
13. Hyæna Palæarctic and Ethiopian (a Palæarctic species.)
14. Canis Palæarctic and Oriental to Malaya.
15. Cuon Oriental to Malaya.
16. Vulpes Very wide range.
17. Lutra Oriental and Palæarctic.
18. Mellivora Ethiopian.
19. Melursus Oriental only; family not Ethiopian.
20. Sus Palæarctic and Oriental, not Ethiopian.
21. Tragulus Oriental. {323}
22. Cervus Oriental and Palæarctic; family not Ethiopian.
23. Cervulus Oriental; family not Ethiopian.
24. Bibos Palæarctic and Oriental.
25. Portax Oriental.
26. Gazella Palæarctic and Ethiopian.
27. Antilope Oriental.
28. Tetraceros Oriental.
29. Elephas Oriental species.
30. Mus Cosmopolite nearly.
31. Platacanthomys Oriental.
32. Meriones Very wide range.
33. Spalacomys Oriental.
34. Sciurus Almost Cosmopolite.
35. Pteromys Palæarctic and Oriental to China and Malaya,
36. Hystrix Wide range.
37. Lepus Wide range.
38. Manis Ethiopian and Oriental to Malaya,
Out of the above 38 genera, 8 have so wide a distribution as to give no
special geographical indications. Of the remaining 30, whose geographical
position we have noted, 14 are Oriental only; 5 have as much right to be
considered Oriental as Ethiopian, extending as they do over the greater
part of the Oriental region; 2 (the hyæna and gazelle) show Palæarctic
rather than Ethiopian affinity; 7 are Palæarctic and Oriental but not
Ethiopian; and only 2 (_Cynælurus_ and _Mellivora_) can be considered as
especially Ethiopian. We must also give due weight to the fact that we have
here Ursidæ and Cervidæ, two families entirely absent from the Ethiopian
region, and we shall then be forced to conclude that the affinities of the
Indian peninsula are not only clearly Oriental, but that the Ethiopian
element is really present in a far less degree than the Palæarctic.
_Birds._--The naturalists who have adopted the "Ethiopian theory" of the
fauna of Hindostan, have always supported their views by an appeal to the
class of birds; maintaining, that not only are almost all the
characteristic Himalayan and Malayan genera absent, but that their place is
to a great extent supplied by others which are characteristic of the
Ethiopian region. After a careful examination of the subject, Mr. Elwes, in
a paper read before the Zoological Society (June 1873) came to the
conclusion, that this view was an erroneous one, founded on the fact that
the birds of the plains are the more abundant and more {324}open to
observation; and that these are often of wide-spread types, and some few
almost exclusively African. The facts he adduced do not, however, seem to
have satisfied the objectors; and as the subject is an important one, I
will here give lists of all the genera of Passeres, Picariæ, Psittaci,
Columbæ, and Gallinæ, which inhabit the sub-region, leaving out those which
only just enter within its boundaries from adjacent sub-regions. These are
arranged under four heads:--1. Oriental genera; which are either wholly
confined to, or strikingly prevalent in, the Oriental region beyond the
limits of the Indian peninsula. 2. Genera of Wide Range; which are fully as
much entitled to be considered Oriental or Palæarctic as Ethiopian, and
cannot be held to prove any Ethiopian affinity. 3. Palæarctic genera; which
are altogether or almost absent from the Ethiopian region. 4. Ethiopian
genera; which are confined to, or very prevalent in, the Ethiopian region,
whence they extend into the Indian peninsula but not over the whole
Oriental region. The last are the only ones which can be fairly balanced
against those of the first list, in order to determine the character of the
fauna.
1. ORIENTAL GENERA IN CENTRAL INDIA.
_Geocichla_, _Orthotomus_, _Prinia_, _Megalurus_, _Abrornis_, _Larvivora_,
_Copsychus_, _Kittacincla_, _Pomatorhinus_, _Malacocercus_, _Chatarrhæa_,
_Layardia_, _Garrulax_, _Trochalopteron_, _Pellorneum_, _Dumetia_,
_Pyctoris_, _Alcippe_, _Myiophonus_, _Sitta_, _Dendrophila_, _Phyllornis_,
_Iora_, _Hypsipetes_, _Pericrocotus_, _Graucalus_, _Volvocivora_, _Chibia_,
_Chaptia_, _Irena_, _Erythrosterna_, _Hemipus_, _Hemichelidon_, _Niltava_,
_Cyornis_, _Eumyias_, _Hypothymis_, _Myialestes_, _Tephrodornis_,
_Dendrocitta_, _Arachnechthra_, _Nectarophila_, _Arachnothera_, _Dicæum_,
_Piprisoma_, _Munia_, _Eulabes_, _Pastor_, _Acridotheres_, _Sturnia_,
_Sturnopastor_, _Artamus_, _Nemoricola_, _Pitta_, _Yungipicus_,
_Chrysocolaptes_, _Hemicircus_, _Gecinus_, _Mulleripicus_, _Brachypternus_,
_Tiga_, _Micropternus_, _Megalæma_, _Xantholæma_, _Rhopodytes_,
_Taccocoua_, _Surniculus_, _Hierococcyx_, _Eudynamnis_, _Nyctiornis_,
_Harpactes_, _Pelargopsis_, _Ceyx_, _Hydrocissa_, _Meniceros_,
_Batrachostomus_, _Dendrochelidon_, _Collocalia_, _Palæornis_, _Treron_,
_Carpophaga_, _Chalcophaps_, _Ortygornis_, _Perdix_, _Pavo_, _Gallus_,
_Galloperdix_;--87 genera; and {325}one peculiar genus, _Salpornis_, whose
affinities are Palæarctic or Oriental.
2. GENERA OF WIDE RANGE OCCURRING IN CENTRAL INDIA.
_Tardus_, _Monticola_, _Drymoeca_, _Cisticola_, _Acrocephalus_,
_Phylloscopus_, _Pratincola_, _Parus_, _Pycnonotus_, _Criniger_, _Oriolus_,
_Dicrurus_, _Tchitrea_, _Lanius_, _Corvus_, _Zosterops_, _Hirundo_,
_Cotyle_, _Passer_, _Ploceus_, _Estrilda_, _Alauda_, _Calandrella_,
_Mirafra_, _Ammomanes_, _Motacilla_, _Anthus_, _Picus_, _Yunx_,
_Centropus_, _Cuculus_, _Chrysoccocyx_, _Coccystes_, _Coracias_,
_Eurystomus_, _Merops_, _Alcedo_, _Ceryle_, _Halcyon_, _Upupa_,
_Caprimulgus_, _Cypselus_, _Chætura_, _Columba_, _Turtur_, _Pterocles_,
_Coturnix_, _Turnix_;--48 genera.
3. PALÆARCTIC GENERA OCCURRING IN CENTRAL INDIA.
_Hypolais_, _Sylvia_, _Curruca_, _Cyanecula_, _Calliope_, _Chelidon_,
_Euspiza_, _Emberiza_, _Galerita_, _Calobates_, _Corydalla_;--11 genera.
4. ETHIOPIAN GENERA OCCURRING IN CENTRAL INDIA.
_Thamnobia_, _Pyrrhulauda_, _Pterocles_, _Francolinus_;--4 genera.
A consideration of the above lists shows us, that the Hindostan sub-region
is by no means so poor in forms of bird-life as is generally supposed (and
as I had myself anticipated, it would prove to be), possessing, as it does,
151 genera of land-birds, without counting the Accipitres. It must also set
at rest the question of the zoological affinities of the district, since a
preponderance of 88 genera, against 4, cannot be held to be insufficient,
and cannot be materially altered by any corrections in details that may be
proposed or substantiated. Even of these four, only the first two are
exclusively Ethiopian, _Pterocles_ and _Francolinus_ both being Palæarctic
also. It is a question, indeed, whether anywhere in the world an outlying
sub-region can be found, exhibiting less zoological affinity for the
adjacent regions; and we have here a striking illustration of the necessity
of deciding all such cases, not by _examples_, which may be so chosen as to
support any view, but by carefully weighing and contrasting the whole of
the facts on which the solution of the {326}problem admittedly depends. It
will, perhaps, be said that a great many of the 88 genera above given are
very scarce and very local; but this is certainly not the case with the
majority of them; and even where it is so, that does not in any degree
affect their value as indicating zoo-geographical affinities. It is the
_presence_ of a type in a region, not its abundance or scarcity, that is
the important fact; and when we have to do, as we have here, with many
groups whose habits and mode of life necessarily seclude them from
observation, their supposed scarcity may not even be a fact.
_Reptiles and Amphibia._--Reptiles entirely agree with Mammalia and Birds
in the main features of their distribution. Out of 17 families of snakes
inhabiting Hindostan, 16 range over the greater part of the entire region,
and only two can be supposed to show any Ethiopian affinity. These are the
Psammophidæ and Erycidæ, both desert-haunting groups, and almost as much
South Palæarctic as African. The genus _Tropidococcyx_ is peculiar to the
sub-region, and _Aspidura_, _Passerita_ and _Cynophis_ to the peninsula and
Ceylon; while a large number of the most characteristic genera, as
_Dipsas_, _Simotes_, _Bungarus_, _Naja_, _Trimeresurus_, _Lycodon_ and
_Python_, are characteristically Oriental.
Of the six families of lizards all have a wide range. The genera _Eumeces_,
_Pentadactylus_, _Gecko_, _Eublepharis_, and _Draco_, are
characteristically or wholly Oriental; _Ophiops_ and _Uromastix_ are
Palæarctic; while _Chamæleon_ is the solitary case of decided Ethiopian
affinity.
Of the Amphibia not a single family exhibits special Ethiopian affinities.
_II. Sub-region of Ceylon and South-India._
The Island of Ceylon is characterised by such striking peculiarities in its
animal productions, as to render necessary its separation from the
peninsula of India as a sub-region; but it is found that most of these
special features extend to the Neilgherries and the whole southern
mountainous portion of India, and that the two must be united in any
zoo-geographical {327}province. The main features of this division
are,--the appearance of numerous animals allied to forms only found again
in the Himalayas or in the Malayan sub-region, the possession of several
peculiar generic types, and an unusual number of peculiar species.
_Mammalia._--Among Mammalia the most remarkable form is _Loris_, a genus of
Lemurs altogether peculiar to the sub-region; several peculiar monkeys of
the genus _Presbytes_; the Malayan genus _Tupaia_; and _Platacanthomys_, a
peculiar genus of Muridæ.
_Birds._--Among birds it has _Ochromela_, a peculiar genus of flycatchers;
_Phoenicophaës_ (Cuculidæ) and _Drymocataphus_ (Timaliidæ), both Malayan
forms; a species of _Myiophonus_ whose nearest ally is in Java;
_Trochalopteron_, _Brachypteryx_, _Buceros_ and _Loriculus_, which are only
found elsewhere in the Himalayas and Malayana. It also possesses about 80
peculiar species of birds, including a large jungle fowl, one owl and two
hornbills.
_Reptiles._--It is however by its Reptiles, even more than by its higher
vertebrates, that this sub-region is clearly characterised. Among snakes it
possesses an entire family, Uropeltidæ, consisting of 5 genera and 18
species altogether confined to it,--_Rhinophis_ and _Uropeltis_ in Ceylon,
_Silybura_, _Plecturus_ and _Melanophidium_ in Southern India. Four other
genera of snakes, _Haplocercus_, _Cercaspis_, _Peltopelor_, and _Hypnale_
are also peculiar; _Chersydrus_ is only found elsewhere in Malaya; while
_Aspidura_, _Passerita_, and _Cynophis_, only extend to Hindostan; and
species of _Eryx_, _Echis_, and _Psammophis_ show an affinity with
Ethiopian and Palæarctic forms. Among lizards several genera of _Agamidæ_
are peculiar, such as _Otocryptis_, _Lyricoephalus_, _Ceratophora_,
_Cophotis_, _Salea_, _Sitana_ and _Charasia_. In the family Acontiadæ,
_Nessia_ is peculiar to Ceylon, while a species of the African genus
_Acontias_ shows an affinity for the Ethiopian region.
_Amphibia._--The genera of Amphibians that occur here are generally of wide
range, but _Nannophrys_, _Haplobatrachus_, and _Cacopus_ are confined to
the sub-region; while _Megalophrys_ is Malayan, and the species found in
Ceylon also inhabit Java.
{328}_Insects._--The insects of Ceylon also furnish some curious examples
of its distinctness from Hindostan, and its affinity with Malaya. Among its
butterflies we find _Papilio jophon_, closely allied to _P. antiphus_ of
Malaya. The remarkable genus _Hestia_, so characteristic of the Malay
archipelago, only occurs elsewhere on the mountains of Ceylon; while its
_Cynthia_ and _Parthenos_ are closely allied to, if not identical with,
Malayan species. Among Coleoptera we have yet more striking examples. The
highly characteristic Malayan genus _Tricondyla_ is represented in Ceylon
by no less than 10 species; and among Longicorns we find the genera
_Tetraommatus_, _Thranius_, _Cacia_, _Praonetha_, _Ropica_, and _Serixia_,
all exclusively Malayan or only just entering the Indo-Chinese peninsula,
yet all represented in Ceylon, while not a single species occurs in any
part of India or the Himalayas.
_The Past History of Ceylon and South-India as indicated by its Fauna._--In
our account of the Ethiopian region we have already had occasion to refer
to an ancient connection between this sub-region and Madagascar, in order
to explain the distribution of the Lemurine type, and some other curious
affinities between the two countries. This view is supported by the geology
of India, which shows us Ceylon and South India consisting mainly of
granitic and old metamorphic rocks, while the greater part of the
peninsula, forming our first sub-region, is of tertiary formation, with a
few isolated patches of secondary rocks. It is evident therefore, that
during much of the tertiary period, Ceylon and South India were bounded on
the north by a considerable extent of sea, and probably formed part of an
extensive southern continent or great island. The very numerous and
remarkable cases of affinity with Malaya, require however some closer
approximation to these islands, which probably occurred at a later period.
When, still later, the great plains and table-lands of Hindostan were
formed, and a permanent land communication effected with the rich and
highly developed Himalo-Chinese fauna, a rapid immigration of new types
took place, and many of the less specialised forms of mammalia and birds
(particularly those of ancient Ethiopian type) became extinct. Among
reptiles and insects the competition was less severe, or the older forms
were too well {329}adapted to local conditions to be expelled; so that it
is among these groups alone that we find any considerable number, of what
are probably the remains of the ancient fauna of a now submerged southern
continent.
_III. Himalayan or Indo-Chinese Sub-region._
This, which is probably the richest of all the sub-regions, and perhaps one
of the richest of all tracts of equal extent on the face of the globe, is
essentially a forest-covered, mountainous country, mostly within the
tropics, but on its northern margin extending some degrees beyond it, and
rising in a continuous mountain range till it meets and intercalates with
the Manchurian sub-division of the Palæarctic region. The peculiar
mammalia, birds and insects of this sub-region begin to appear at the very
foot of the Himalayas, but Dr. Gunther has shown that many of the reptiles
characteristic of the plains of India are found to a height of from 2,000
to 4,000 feet.
In Sikhim, which may be taken as a typical example of the Himalayan portion
of the sub-region, it seems to extend to an altitude of little less than
10,000 feet, that being the limit of the characteristic Timaliidæ or
babbling thrushes; while the equally characteristic Pycnonotidæ, or
bulbuls, and Treronidæ, or thick-billed fruit-pigeons, do not, according to
Mr. Blanford, reach quite so high. We may perhaps take 9,000 feet as a good
approximation over a large part of the Himalayan range; but it is evidently
not possible to define the line with any great precision. Westward, the
sub-region extends in diminishing breadth, till it terminates in or near
Cashmere, where the fauna of the plains of India almost meets that of the
Palæarctic region, at a moderate elevation. Eastward, it reaches into East
Thibet and North-west China, where Père David has found a large number of
the peculiar types of the Eastern Himalayas. A fauna, in general features
identical, extends over Burmah and Siam to South China; mingling with the
Palæarctic fauna in the mountains south of the Yang-tse-kiang river, and
with that of Indo-Malaya in Tenasserim, and to a lesser extent in Southern
Siam and Cochin China.
{330}_Zoological Characteristics of the Himalayan Sub-region._--Taking this
sub-region as a whole, we find it to be characterised by 3 genera of
mammalia (without counting bats), and 44 genera of land-birds, which are
altogether peculiar to it; and by 13 genera of mammalia and 36 of birds,
which it possesses in common with the Malayan sub-region; and besides these
it has almost all the genera before enumerated as "Oriental," and several
others of wide range, more especially a number of Palæarctic genera which
appear in the higher Himalayas. The names of the more characteristic genera
are as follows:--
PECULIAR HIMALO-CHINESE GENERA.
Mammalia.--_Urva_, _Arctonyx_, _Ælurus_.
Birds.--_Suya_, _Horites_, _Chæmarrhornis_, _Tarsiger_, _Oreicola_,
_Acanthoptila_, _Grammatoptila_, _Trochalopteron_, _Actinodura_, _Sibia_,
_Suthora_, _Paradoxornis_, _Chlenasicus_, _Tesia_, _Rimator_,
_Ægithaliscus_, _Cephalopyrus_, _Liothrix_, _Siva_, _Minla_, _Proparus_,
_Cutia_, _Yuhina_, _Ixulus_, _Myzornis_, _Erpornis_, _Hemixus_, _Chibia_,
_Niltava_, _Anthipes_, _Chelidorhynx_, _Urocissa_, _Pachyglossa_,
_Heterura_, _Hæmatospiza_, _Ampeliceps_, _Saroglossa_, _Psarisomus_,
_Serilophus_, _Vivia_, _Hyopicus_, _Gecinulus_, _Aceros_, _Ceriornis_.
GENERA COMMON TO THE HIMALO-CHINESE AND MALAYAN SUB-REGIONS.
Mammalia.--_Hylobates_, _Nycticebus_, _Viverricula_, _Prionodon_,
_Arctitis_, _Paguma_, _Arctogale_, _Cuon_, _Gymnopus_, _Aonyx_, _Helictis_,
_Rhinoceros_, _Nemorhedus_, _Rhizomys_.
Birds.--_Oreocincla_, _Notodela_, _Janthocincla_, _Timalia_, _Stachyris_,
_Mixornis_, _Trichastoma_, _Enicurus_, _Pnoepyga_, _Melanochlora_,
_Allotrius_, _Microscelis_, _Iole_, _Analcipus_, _Cochoa_, _Bhringa_,
_Xanthopygia_, _Hylocharis_, _Cissa_, _Temnurus_, _Crypsirhina_,
_Chalcostetha_, _Anthreptes_, _Chalcoparia_, _Cymbirhynchus_, _Hydrornis_,
_Sasia_, _Venilia_, _Indicator_, _Carcineutes_, _Lyncornis_, _Macropygia_,
_Argusianus_, _Polyplectron_, _Euplocamus_, _Phodilus_.
Plate VII.
[Illustration]
SCENE IN NEPAUL, WITH CHARACTERISTIC ANIMALS.
{331}_Plate VII. Scene in Nepal, with Characteristic Himalayan
Animals._--Our illustration contains figures of two mammals and two birds,
characteristic of the higher woody region of the Himalayas. The lower
figure on the left is the _Helictis nepalensis_, confined to the Eastern
Himalayas, and belonging to a genus of the weasel family which is
exclusively Oriental. It is marked with white on a grey-brown ground. Above
it is the remarkable Panda (_Ælurus fulgens_), a beautiful animal with a
glossy fur of a reddish colour, darker feet, and a white somewhat cat-like
face. It is distantly allied to the bears, and more nearly to the American
racoons, yet with sufficient differences to constitute it a distinct
family. The large bird on the tree, is the horned Tragopan (_Ceriornis
satyra_), one of the fine Himalayan pheasants, magnificently spotted with
red and white, and ornamented with fleshy erectile wattles and horns, of
vivid blue and red colours. The bird in the foreground is the
_Ibidorhynchus struthersii_, a rare and curious wader, allied to the
curlews and sandpipers but having the bill and feet red. It frequents the
river-beds in the higher Himalayas, but has also been found in Thibet.
_Reptiles._--Very few genera of reptiles are peculiar to this sub-region,
all the more important ranging into the Malay islands. Of snakes the
following are the more characteristic genera:--_Typhline_, _Cylindrophis_,
_Xenopeltis_, _Calamaria_, _Xenelaphis_, _Hypsirhina_, _Fordonia_, several
small genera of Homalopsidæ (_Herpeton_ and _Hipistes_ being characteristic
of Burmah and Siam), _Psammodynastes_, _Gonyosoma_, _Chrysopelea_,
_Tragops_, _Dipsas_, _Pareas_, _Python_, _Bungarus_, _Naja_, _Callophis_,
and _Trimeresurus_. _Naja_ reaches 8,000 feet elevation in the Himalayas,
_Tropidonotus_ 9,000 feet, _Ablabes_ 10,000 feet, and _Simotes_ 15,000
feet.
Of lizards, _Pseudopus_ has one species in the Khasya hills while the other
inhabits South-east Europe; and there are two small genera of Agamidæ
peculiar to the Himalayas, while _Draco_ and _Calotes_ have a wide range
and _Acanthosaura_, _Dilophyrus_, _Physignathus_, and _Liolepis_ are found
chiefly in the Indo-Chinese peninsula. There are several genera of
Scincidæ; and the extensive genus of wall-lizards, _Gecko_, ranges over the
whole region.
Of Amphibia, the peculiar forms are not numerous. _Ichthyophis_ {332}a
genus of Ceciliadæ, is peculiar to the Khasya Hills; _Tylotritron_
(Salamandridæ) to Yunan in Western China, and perhaps belongs to the
Palæarctic region.
Of the tail-less Batrachians, _Glyphoglossus_ is found in Pegu; _Xenophys_
in the Eastern Himalayas; while _Callula_, _Ixalus_, _Rhacophorus_,
_Hylurana_, _Oxyglossus_, and _Phrynoglossus_, are common to the
Himalo-Chinese and Malayan sub-regions.
Of the lizards, _Colotes_, _Barycephalus_, and _Hinulia_,--and of the
Batrachia, _Bufo_,--are found at above 11,000 feet elevation in the
Himalayas.
_Insects._--So little has been done in working out the insect faunas of the
separate sub-regions, that they cannot be treated in detail, and the reader
is referred to the chapter on the distribution of insects in the part of
this work devoted to Geographical Zoology. A few particulars may, however,
be given as to the butterflies, which have been more systematically
collected in tropical countries than any other order of insects. The
Himalayan butterflies, especially in the eastern portions of the range--in
Assam and the Khasya Hills--are remarkably fine and very abundant; yet all
the larger groups extend into the Malayan sub-region, many to Ceylon, and a
considerable proportion even to Africa and Austro-Malaya. There are a large
number of peculiar types, but most of them consist of few or single
species. Such are _Neope_, _Orenoma_, and _Rhaphicera_, genera of Satyridæ;
_Enispe_ (Morphidæ); _Hestina_, _Penthema_, and _Abrota_ (Nymphalidæ);
_Dodona_ (Erycinidæ); _Ilerda_ (Lycænidæ); _Calinaga_, _Teinopalpus_, and
_Bhutanitis_ (Papilionidæ). Its more prominent features are, however,
derived from what may be termed Malayan, or even Old World types, such as
_Euplæa_, among Danaidæ; _Amathusia_, _Clerome_, and _Thaumantis_, among
Morphidæ; _Euripus_, _Diadema_, _Athyma_, _Limenitis_, and _Adolias_, among
Nymphalidæ; _Zemeros_ and _Taxila_ among Erycinidæ; _Amblypodia_,
_Miletus_, _Ilerda_, and _Myrina_, among Lycænidæ; _Thyca_, _Prioneris_,
_Dercas_, _Iphias_, and _Thestias_ among Pieridæ; and Papilios of the
"_Amphrisius_," "_Coon_", "_Philoxenus_," "_Protenor_," "_Paris_," and
"_Sarpedon_" groups. In the Himalayas there is an unusual abundance of
large and gorgeous species of the genus _Papilio_, {333}and of large and
showy Nymphalidæ, Morphidæ, and Danaidæ, which render it, in favoured
localities, only second to South America for a display of this form of
beauty and variety in insect life.
Among the other orders of insects in which the Himalayas are remarkably
rich, we may mention large and brilliant Cetoniidæ, chiefly of the genus
_Rhomborhima_; a magnificent Lamellicorn, _Euchirus macleayii_, allied to
the gigantic long-armed beetle (_E. longimanus_) of Amboyna; superb moths
of the families Agaristidæ and Sesiidæ; elegant and remarkable Fulgoridæ,
and strange forms of the gigantic Phasmidæ; most of which appear to be of
larger size or of more brilliant colours than their Malayan allies.
_Islands of the Indo-Chinese Sub-region._--A few important islands belong
to this sub-region, the Andamans, Formosa, and Hainan being the most
interesting.
_Andamans._--The only mammalia are a few rats and mice, a _Paradoxurus_,
and a pig supposed to be a hybrid race,--all of which may have been
introduced by man's agency. The birds of the Andaman Islands have been
largely collected, no less than 155 species having been obtained; and of
these 17, (all land-birds) are peculiar. The genera are all found on the
continent, and are mostly characteristic of the Indo-Chinese fauna, to
which most of the species belong. Reptiles are also tolerably abundant;
about 20 species are known, the majority being found also on the continent,
while a few are peculiar. There are also a few Batrachia, and some
fresh-water fishes, closely resembling those of Burmah. The absence of such
mammalia as monkeys and squirrels, which abound on the mainland, and which
are easily carried over straits or narrow seas by floating trees, is
sufficient proof that these islands have not recently formed part of the
continent. The birds are mostly such as may have reached the islands while
in their present geographical position; and the occurrence of reptiles and
fresh-water fishes, said to be identical in species with those of Burmah,
must be due to the facilities, which some of these animals undoubtedly
{334}possess, for passing over a considerable width of sea. We must
conclude, therefore, that these islands do not owe their existing fauna to
an actual union with the mainland; but it is probable that they may have
been formerly more extensive, and have then been less distant from the
continent than at the present time.
The Nicobar Islands, usually associated with the Andamans, are less known,
but present somewhat similar phenomena. They are, however, more Malayan in
their fauna, and seem properly to belong to the Indo-Malay sub-region.
_Formosa._--This island has been carefully examined by Mr. Swinhoe, who
found 144 species of birds, of which 34 are peculiar. There is one peculiar
genus, but the rest are all Indo-Chinese, though some of the species are
more allied to Malayan than to Chinese or Himalayan forms. About 30 species
of mammalia were found in Formosa, of which 11 are peculiar species, the
rest being either Chinese or Himalayan. The peculiar species belong to the
genera _Talpa_, _Helictis_, _Sciuropterus_, _Pteromys_, _Mus_, _Sus_,
_Cervus_, and _Capricornis_. A few lizards and snakes of continental
species have also been found. These facts clearly indicate the former
connection of Formosa with China and Malaya, a connection which is rendered
the more probable by the shallow sea which still connects all these
countries.
_Hainan._--The island of Hainan, on the south coast of China, is not so
well known in proportion, though Mr. Swinhoe collected 172 species of
birds, of which 130 were land-birds. Of these about 20 were peculiar
species; the remainder being either Chinese, Himalayan, or Indo-Malayan.
Mr. Swinhoe also obtained 24 species of mammalia, all being Chinese,
Himalayan, or Indo-Malayan species except a hare, which is peculiar. This
assemblage of animals would imply that Hainan, as might be anticipated from
its position, has been more recently separated from the continent than the
more distant island of Formosa.
_IV. Indo-Malaya, or the Malayan Sub-region._
This sub-region, which is almost wholly insular (including only the Malayan
peninsula on the continent of Asia), is equal, if {335}not superior, in the
variety and beauty of its productions, to that which we have just been
considering. Like Indo-China, it is a region of forests, but it is more
exclusively tropical; and it is therefore deficient in many of those
curious forms of the temperate zone of the Himalayas, which seem to have
been developed from Palæarctic rather than from Oriental types. Here alone,
in the Oriental region, are found the most typical equatorial forms of
life--organisms adapted to a climate characterised by uniform but not
excessive heat, abundant moisture, and no marked departure from the average
meteorological state, throughout the year. These favourable conditions of
life only occur in three widely separated districts of the globe--the Malay
archipelago, Western Africa, and equatorial South America. Hence perhaps it
is, that the tapir and the trogons of Malacca should so closely resemble
those of South America; and that the great anthropoid apes and crested
hornbills of Western Africa, should find their nearest allies in Borneo and
Sumatra.
Although the islands which go to form this sub-region are often separated
from each other by a considerable expanse of sea, yet their productions in
general offer no greater differences than those of portions of the
Indo-Chinese sub-region separated by an equal extent of dry land. The
explanation is easy, however, when we find that the sea which separates
them is a very shallow one, so shallow that an elevation of only 300 feet
would unite Sumatra, Java, and Borneo into one great South-eastern
prolongation of the Asiatic continent. As we know that our own country has
been elevated and depressed to a greater amount than this, at least twice
in recent geological times, we can have no difficulty in admitting similar
changes of level in the Malay archipelago, where the subterranean forces
which bring about such changes are still at work, as manifested by the
great chain of active volcanoes in Sumatra and Java. Proofs of somewhat
earlier changes of level are to be seen in the Tertiary coal formations of
Borneo, which demonstrate a succession of elevations and subsidences, with
as much certainty as if we had historical record of them.
It is not necessary to suppose, nor is it probable, that all these
{336}great islands were recently united to the continent, and that their
separation took place by one general subsidence of the whole. It is more
consonant with what we know of such matters, that the elevations and
depressions were partial, varying in their points of action and often
recurring; sometimes extending one part of an island, sometimes another;
now joining an island to the main land, now bringing two islands into
closer proximity. There is reason to believe that sometimes an intervening
island has sunk or receded and allowed others which it before separated to
effect a partial union independently of it. If we recognise the probability
that such varied and often-renewed changes of level have occurred, we shall
be better able to understand how certain anomalies of distribution in these
islands may have been brought about. We will now endeavour to sketch the
general features of the zoology of this interesting district, and then
proceed to discuss some of the relations of the islands to each other.
_Mammalia._--We have seen that the Indo-Chinese sub-region possesses 13
species of mammalia in common with the Indo-Malay sub-region, and 4 others
peculiar to itself, besides one Ethiopian and several Oriental and
Palæarctic forms of wide range. Of this latter class the Malay islands have
comparatively few, but they possess no less than 14 peculiar genera, viz.
_Simia_, _Siamanga_, _Tarsius_, _Galeopithecus_, _Hylomys_, _Ptilocerus_,
_Gymnura_, _Cynogale_, _Hemigalea_, _Arctogale_, _Barangia_, _Mydaus_,
_Helarctos_, and _Tapirus_. The islands also possess tigers, deer, wild
pigs, wild cattle, elephants, the scaly ant-eater, and most of the usual
Oriental genera; so that they are on the whole fully as rich as, if not
richer than, any part of Asia; a fact very unusual in island faunas, and
very suggestive of their really continental nature.
Plate VIII.
[Illustration]
A FOREST IN BORNEO, WITH CHARACTERISTIC MAMMALIA.
{337}_Plate VIII. Scene in Borneo with Characteristic Malayan
Quadrupeds._--The Malayan fauna is so rich and peculiar that we devote two
plates to illustrate it. We have here a group of mammalia, such as might be
seen together in the vast forests of Borneo. In the foreground we have the
beautiful deer-like Chevrotain (_Tragulus javanicus_). These are delicate
little animals whose body is not larger than a rabbit's, thence often
called "mouse-deer." They were formerly classed with the "musk-deer," owing
to their similar tusk-like upper canines; but their anatomy shows them to
form quite a distinct family, having more resemblance to the camels. On the
branch above is the curious feather-tailed Tree-Shrew (_Ptilocerus lowii_),
a small insectivorous animal altogether peculiar to Borneo. Above this is
the strange little Tarsier (_Tarsius spectrum_), one of the lemurs confined
to the Malay islands, but so distinct from all others as to constitute a
separate family. The other small animals are the Flying Lemurs
(_Galæopithecus volans_) formerly classed with the lemurs, but now
considered to belong to the Insectivora. They have a very large expansion
of the skin connecting the fore and hind limbs and tail, and are able to
take long flights from one tree to another, and even to rise over obstacles
in their course by the elevatory power of the tail-membrane. They feed
chiefly on leaves, and have a very soft and beautifully marbled fur.
In the distance is the Malayan tapir (_Tapirus indicus_), a representative
of a group of animals now confined to the larger Malay islands and tropical
America, but which once ranged over the greater part of temperate Europe.
_Birds._--Owing to several of the families consisting of very obscure and
closely allied species, which have never been critically examined and
compared by a competent ornithologist, the number of birds inhabiting this
sub-region is uncertain. From the best available materials there appear to
be somewhat less than 650 species of land-birds actually known, or
excluding the Philippine Islands somewhat less than 600. The larger part of
these are peculiar species, but mostly allied to those of Indo-China; 36 of
the genera, as already stated, being common to these two sub-regions. There
are, however, no less than 46 genera which are peculiarly or wholly Indo
Malayan and, in many cases, have no close affinity with other Oriental
groups. These peculiar genera are as follows:--_Timalia_, _Malacopteron_,
_Macronus_, _Napothera_, _Turdinus_, and _Trichixos_--genera {338}of
Timaliidæ; _Eupetes_, a most remarkable form, perhaps allied to _Enicurus_,
and _Cinclus_; _Rhabdornis_ (Certhiidæ) found only in the Philippines;
_Psaltria_, a diminutive bird of doubtful affinities, provisionally classed
among the tits (Paridæ); _Setornis_ (Pycnonotidæ); _Lalage_ (Campephagidæ)
extending eastward to the Pacific Islands; _Pycnosphrys_, _Philentoma_
(Muscicapidæ); _Laniellus_, a beautiful bird doubtfully classed with the
shrikes (Laniidæ); _Platylophus_ and _Pityriasis_, the latter a most
anomalous form--perhaps a distinct family, at present classed with the
jays, in Corvidæ; _Prionochilus_, a curious form classed with Dicæidæ;
_Erythrura_ (Ploceidæ), extending eastwards to the Fiji Islands; _Gymnops_,
_Calornis_, (Sturnidæ); _Eurylæmus_, _Corydon_, and _Calyptomena_
(Eurylæmidæ); _Eucichla_, the longest tailed and most elegantly marked of
the Pittidæ; _Reinwardtipicus_ and _Miglyptes_ (Picidæ); _Psilopogon_ and
_Calorhamphus_, (Megalæmidæ); _Rhinococcyx_, _Dasylophus_, _Lepidogrammus_,
_Carpococcyx_, _Zanclostomus_, _Poliococcyx_, _Rhinortha_, (Cuculidæ);
_Berenicornis_, _Caldo_, _Cranorhinus_, _Penelopides_, _Rhinoplax_,
(Bucerotidæ); _Psittinus_, (Psittacidæ); _Ptilopus_, _Phapitreron_,
(Columbidæ); _Rollulus_, (Treronidæ); _Machærhamphus_, (Falconidæ). Many of
these genera are abundant and wide-spread, while some of the most
characteristic Himalayan genera, such as _Larvivora_, _Garrulax_,
_Hypsipetes_, _Pomatorhinus_, and _Dendrocitta_, are here represented by
only a few species.
Among the groups that are characteristic of the Malayan sub-region, the
Timaliidæ and Pycnonotidæ stand pre-eminent; the former represented chiefly
by the genera _Timalia_, _Malacopteron_, _Macronus_, and _Trichastoma_, the
latter by _Criniger_, _Microscelis_, and many forms of _Pycnonotus_. The
Muscicapidæ, Dicruridæ, Campephagidæ, Ploceidæ, and Nectariniidæ are also
well developed; as well as the Pittidæ, and the Eurylæmidæ, the limited
number of species of the latter being compensated by a tolerable abundance
of individuals. Among the Picariæ are many conspicuous groups; as,
woodpeckers (Picidæ); barbets (Megalæmidæ); trogons (Trogonidæ);
kingfishers (Alcedinidæ); and hornbills (Bucerotidæ); five families which
are perhaps the most conspicuous in the whole fauna. Lastly come the
pigeons {339}(Columbidæ), and the pheasants (Phasianidæ), which are fairly
represented by such fine genera as _Treron_, _Ptilopus_, _Euplocamus_, and
_Argusianus_. A few forms whose affinities are Australian rather than
Oriental, help to give a character to the ornithology, though none of them
are numerous. The swallow-shrikes (_Artamus_); the wag-tail fly-catchers
(_Rhipidura_); the green fruit-doves (_Ptilopus_); and the mound-makers
(_Megapodius_), are the chief of these.
There are a few curious examples of remote geographical alliances that may
be noted. First, we have a direct African connection in _Machærhamphus_, a
genus of hawks, and _Berenicornis_, a genus of hornbills; the only close
allies being, in the former case in South, and in the latter in West
Africa. Then we have a curious Neotropical affinity, indicated by
_Carpococcyx_, a large Bornean ground-cuckoo, whose nearest ally is the
genus _Neomorphus_ of South America; and by the lovely green-coloured
_Calyptomena_ which seems unmistakably allied to the orange-coloured
_Rupicola_, or "Cock of the rock," in general structure and in the
remarkable form of crest, a resemblance which has been noticed by many
writers.
In the preceding enumeration of Malayan genera several are included which
extend into the Austro-Malay Islands, our object, at present, being to show
the differences and relations of the two chief Oriental sub-regions.
_Plate IX. A Malayan Forest with some of its peculiar Birds._--Our second
illustration of the Malayan fauna is devoted to its bird-life; and for this
purpose we place our scene in the Malay peninsula, where birds are perhaps
more abundant and more interesting, than in any other part of the
sub-region. Conspicuous in the foreground is the huge Rhinoceros Hornbill
(_Buceros rhinoceros_), one of the most characteristic birds of the Malayan
forests, the flapping of whose wings, as it violently beats the air to
support its heavy body, may be heard a mile off. On the ground behind, is
the Argus pheasant (_Argusianus giganteus_) whose beautifully ocellated
wings have been the subject of a most interesting description in Mr.
Darwin's _Descent of Man_. The wing-feathers are here so enormously
{340}developed for display (as shown in our figure) that they become
almost, if not quite, useless for their original purpose of flight; yet the
colours are so sober, harmonizing completely with the surrounding
vegetation, and the bird is so wary, that in the forests where it abounds
an old hunter assured me he had never been able to see a specimen till it
was caught in his snares. It is interesting to note, that during the
display of the plumage the bird's head is concealed by the wings from a
spectator in front, and, contrary to what usually obtains among pheasants,
the head is entirely unadorned, having neither crest nor a particle of
vivid colour,--a remarkable confirmation of Mr. Darwin's views, that gayly
coloured plumes are developed in the male bird for the purpose of
attractive display in the breeding season. The long-tailed bird on the
right is one of the Drongo-shrikes (_Bhringa remifer_), whose long bare
tail-feathers, with an oar-like web at the end, and blue-black glossy
plumage, render it a very attractive object as it flies after its insect
prey. On the left is another singular bird the great Broad-bill (_Corydon
sumatranus_), with dull and sombre plumage, but with a beak more like that
of a boat-bill than of a fruit-eating passerine bird. Over all, the
white-handed Gibbon (_Hylobates lar_) swings and gambols among the topmost
branches of the forest.
_Reptiles and Amphibia._--These are not sufficiently known to be of much
use for our present purpose. Most of the genera belong to the continental
parts of the Oriental region, or have a wide range. Of snakes _Rhabdosoma_,
_Typhlocalamus_, _Tetragonosoma_, _Acrochordus_, and _Atropos_, are the
most peculiar, and there are several peculiar genera of Homalopsidæ. Of
Oriental genera, _Cylindrophis_, _Xenopeltes_, _Calamaria_, _Hypsirhina_,
_Psammodynastes_, _Gonyosoma_, _Tragops_, _Dipsas_, _Pareas_, _Python_,
_Bungarus_, _Naja_, and _Callophis_ are abundant; as well as _Simotes_,
_Ablabes_, _Tropidonotus_, and _Dendrophis_, which are widely distributed.
Among lizards _Hydrosaurus_ and _Gecko_ are common; there are many isolated
groups of Scincidæ; while _Draco_, _Calotes_, and many forms of Agamidæ,
some of which are peculiar, abound.
Plate IX.
[Illustration]
A MALAYAN FOREST, WITH ITS CHARACTERISTIC BIRDS.
{341}Among the Amphibia, toads and frogs of the genera _Micrhyla_,
_Kalophrynus_, _Ansonia_, and _Pseudobufo_, are peculiar: while the
Oriental _Megalophrys_, _Ixalus_, _Rhacophorus_, and _Hylorana_ are
abundant and characteristic.
_Fishes._--The fresh-water fishes of the Malay archipelago have been so
well collected and examined by the Dutch naturalists, that they offer
valuable indications of zoo-geographical affinity; and they particularly
well exhibit the sharply defined limits of the region, a large number of
Oriental and even Ethiopian genera extending eastward as far as Java and
Borneo, but very rarely indeed sending a single species further east, to
Celebes or the Moluccas. Thirteen families of fresh-water fishes are found
in the Indo-Malay sub-region. Of these the Scienidæ and Symbranchidæ have
mostly a wide range in the tropics. Ophiocephalidæ are exclusively
Oriental, reaching Borneo and the Philippine islands. The Mastacembelidæ
are also Oriental, but one species is found as far as Ceram. Of the
Nandidæ, 3 genera range over the whole region. The Labyrinthici extend from
Africa through the Oriental region to Amboyna, The single species
constituting the family Luciocephalidæ is confined to Borneo and the small
islands of Biliton and Banca. Of the extensive family Siluridæ 17 genera
are Oriental and Malayan, and 11 are Malayan exclusively; and not one of
these appears to pass beyond the limits of the sub-region. The Cyprinidæ
offer an equally striking example, 23 genera ranging eastward to Java and
Borneo and not one beyond; 14 of these being exclusively Malayan. It must
be remembered that this is not from any want of knowledge of the countries
farther east, as extensive collections have also been made in Celebes, the
Moluccas, and Timor; so that the facts of distribution of fresh-water
fishes come, most unexpectedly, to fortify that division of the archipelago
into two primary regions, which was founded on a consideration of mammalia
and birds only.
_Insects._--Few countries in the world can present a richer and more varied
series of insects than the Indo-Malay islands, and we can only here notice
a few of their more striking peculiarities and more salient features.
{342}The butterflies of this sub-region, according to the best estimate
that can be formed, amount to about 650 described species, a number that
will yet, no doubt, be very considerably increased. The genera which appear
to be peculiar to it are _Erites_ (Satyridæ); _Zeuxidia_ (Morphidæ);
_Amnosia_, _Xanthotænia_, and _Tanæcia_ (Nymphalidæ). The groups which are
most characteristic of the region, either from their abundance in
individuals or species, or from their size and beauty, are--the rich
dark-coloured _Euplæa_; the large semi-transparent _Hestia_; the
plain-coloured _Mycalesis_, which replace our meadow-brown butterflies
(_Hipparchia_); the curious _Elymnias_, which often closely resemble
Euplæas; the large and handsome _Thamantis_ and _Zeuxidia_, which take the
place of the giant Morphos of South America; the _Cethosia_, of the
brightest red, and marked with a curious zigzag pattern; the velvety and
blue-glossed _Terinos_; the pale and delicately-streaked _Cyrestis_; the
thick-bodied and boldly coloured _Adolias_; the small wine-coloured
_Taxila_; the fine blue _Amblypodia_; the beautiful _Thyca_, elegantly
marked underneath with red and yellow, which represent our common white
butterflies and are almost equally abundant; the pale blue _Eronia_, and
the large red-tipped _Iphias_. The genus _Papilio_ is represented by a
variety of fine groups; the large _Ornithoptera_, with satiny yellow
under-wings; the superb green-marked "_brookeana_;" the "_paradoxa_" group,
often closely resembling the Euplæas that abound in the same district; the
"_paris_" group richly dusted with golden-green specks; the "_helenus_"
group with wide-spreading black and white wings; the black and crimson
"_polydorus_" group; the "_memnon_" group, of the largest size and
richly-varied colours; and the "_eurypilus_" group, elegantly banded or
spotted with blue or green: all these are so abundant that some of them are
met with in every walk, and are a constant delight to the naturalist who
has the privilege of observing them in their native haunts.
The Coleoptera are far less prominent and require to be carefully sought
after; but they then well repay the collector. As affording some measure of
the productiveness of the tropics in insect life it will not be out of
place to give a few notes of the {343}number of species collected by myself
in some of the best localities. At Singapore 300 species of Coleoptera were
collected in 15 days, and in a month the number had increased to 520; of
which 100 were Longicorns and 140 Rhyncophora. At Sarawak in Borneo I
obtained 400 species in 15 days, and 600 in a month. In two months this
number had increased to about 850, and in three months to 1,000 species.
This was the most prolific spot I ever collected in, especially for
Longicorns which formed about one-fifth of all the species of beetles. In
the Aru Islands in one month, I obtained only 235 species of Coleoptera,
and about 600 species of insects of all orders; and this may be taken as a
fair average, in localities where no specially favourable conditions
existed. On the average 40 to 60 species of Coleoptera would be a good
day's collecting; 70 exceptionally good; while the largest number ever
obtained in one day was 95, and the majority of these would be very minute
insects. It must be remembered, however, that many very common species were
passed over, yet had every species met with been collected, not much more
than 100 species would ever have been obtained in one day's collecting of
four or five hours. These details may afford an interesting standard of
comparison for collectors in other parts of the world.
Of Cicindelidæ the most peculiarly Malayan form is _Therates_, found always
on leaves in the forests in the same localities as the more widely spread
_Collyris_. Five genera of this family are Indo-Malayan.
The Carabidæ, though sufficiently plentiful, are mostly of small size, and
not conspicuous in any way. But there is one striking exception in the
purely Malayan genus _Mormolyce_, the largest and most remarkable of the
whole family. It is nocturnal, resting during the days on the under side of
large _boleti_ in the virgin forest. _Pericallus_ and _Catascopus_ are
among the few genera which are at all brillantly coloured.
Buprestidæ are abundant, and very gay; the genus _Belionota_ being perhaps
one of the most conspicuous and characteristic. The giant _Catoxantha_ is,
however, the most peculiar, though comparatively scarce. _Chrysochroa_ and
_Chalcophora_ are also {344}abundant and characteristic. Out of the 41
Oriental genera 21 are Malayan, and 10 of these are not found in the other
sub-regions.
In Lucanidæ the Malay islands are rich, 14 out of the 16 Oriental genera
occurring there, and 3 being peculiar. There are many fine species of
_Odontolabris_, which may be considered the characteristic genus of the
sub-region.
The Cetoniidæ are well represented by 16 genera and about 120 species. The
genera _Mycteristes_, _Phædimus_, _Plectrone_, _Euremina_, _Rhagopteryx_
and _Centrognathus_ are peculiar, while _Agestrata_, _Chalcothea_, and
_Macronota_ are abundant and characteristic.
The Longicorns, as in all continental forest regions near the equator, are
very abundant and in endlessly varied forms. No less than 55 genera
containing about 200 species are peculiar to this sub-region, the
Cerambycidæ being much the most numerous. _Euryarthrum_, _Coelosterna_,
_Agelasta_, and _Astathes_ may be considered as most characteristic; but to
name the curious and interesting forms would be to give a list of half the
genera. For the relations of the Longicorns of the Indo-Malay, and those of
the Austro-Malay region, the reader is referred to the chapter on the
distribution of insects in the succeeding part of this work.
_Terrestrial Mollusca._--The Philippine islands are celebrated as being one
of the richest parts of the world for land shells, about 400 species being
known. The other islands of the sub-region are far less rich, not more than
about 100 species having yet been described from the whole of them. _Helix_
and _Bulimus_ both abound in species in the Philippines, whereas the latter
genus is very scarce in Borneo and Java. Ten genera of Helicidæ inhabit the
sub-region; _Pfeifferia_ is found in the Philippines and Moluccas, while
the large genus _Cochlostyla_ is almost peculiar to the Philippines. Of the
Operculata there are representatives of 20 genera, of which _Dermatoma_ and
_Pupinella_ are peculiar, while _Registoma_ and _Callia_ extend to the
Australian region. _Cyclophorus_, _Leptopoma_, and _Pupina_ are perhaps the
most characteristic genera.
{345}_The Zoological Relations of the Several Islands of the Indo-Malay
Sub-region._
Although we have grouped the Philippine islands with the Indo-Malay
sub-region, to which, as we shall see, they undoubtedly belong, yet most of
the zoological characteristics we have just sketched out, apply more
especially to the other groups of islands and the Malay peninsula. The
Philippine islands stand, to Malaya proper, in the same relation that
Madagascar does to Africa or the Antilles to South America; that is, they
are remarkable for the absence of whole families and genera which
everywhere characterise the remainder of the district. They are, in fact,
truly insular, while the other islands are really continental in all the
essential features of their natural history. Before, therefore, we can
conveniently compare the separate islands of Malaya[12] with each other, we
must first deal with the Philippine group, showing in what its speciality
consists, and why it must be considered apart from the sub-region to which
it belongs.
_Mammals of the Philippine Islands._--The only mammalia recorded as
inhabiting the Philippine Islands are the following:--
QUADRUMANA. 1. Macacus cynomolgus.
2. Cynopithecus niger. Dr. Semper doubts this
being a Philippine
species.
LEMUROIDEA. 3. Tarsius spectrum.
INSECTIVORA. 4. Galeopithecus philippinensis.
5. Tupaia (species). On Dr. Semper's authority.
CARNIVORA. 6. Viverra tangalunga.
7. Paradoxurus philippensis.
UNGULATA. 8. Sus (species). On Dr. Semper's authority.
9. Cervus mariannus.
10. Cervus philippensis.
11. Cervus alfredi.
12. Bos (species). Wild cattle; perhaps
introduced.
RODENTIA. 13. Phlæomys cummingii.
14. Scuirus philippinensis.
Also 24 species, belonging to 17 genera, of bats.
{346}The foregoing list, although small, contains an assemblage of species
which are wholly Oriental in character, and several of which (_Tarsius_,
_Galeopithecus_, _Tupaia_) are characteristic and highly peculiar Malayan
forms. At the same time these islands are completely separated from the
rest of Malaya by the total absence of _Semnopithecus_, _Hylobates_,
_Felis_, _Helarctos_, _Rhinoceros_, _Manis_, and other groups constantly
found in the great Indo-Malay islands and peninsula of Malacca. We find
apparently two sets of animals: a more ancient series, represented by the
deer, _Galeopithecus_, and squirrel, in which the species are distinct from
any others; and a more recent series, represented by _Macacus cynomolgus_,
and _Viverra tangalunga_, identical with common Malayan animals. The former
indicate the earliest period when these volcanic islands were connected
with some part of the Malayan sub-region, and they show that this was not
geologically remote, since no peculiar generic types have been preserved or
differentiated. The latter may indicate either the termination of the
period of union, or merely the effects of introduction by man. The reason
why a larger number of mammalian forms were not introduced and established,
was probably because the union was effected only with some small islands,
and from these communicated to other parts of the archipelago; or it may
well be that later subsidences extinguished some of the forms that had
established themselves.
_Birds of the Philippine Islands._--These have been carefully investigated
by Viscount Walden, in a paper read before the Zoological Society of London
in 1873, and we are thus furnished with ample information on the relations
of this important portion of the fauna.
The total number of birds known to inhabit the Philippines is 219, of which
106 are peculiar. If, however, following our usual plan, we take only the
land-birds, we find the numbers to be 159 species, of which 100 are
peculiar; an unusually large proportion for a group of islands so
comparatively near to various parts of the Oriental and Australian regions.
The families of birds which are more especially characteristic of the
Indo-Malay sub-region are about 28 in number, and examples {347}of all
these are found in the Philippines except four, viz., Cinclidæ,
Phyllornithidæ, Eurylæmidæ, and Podargidæ. The only Philippine families
which are, otherwise, exclusively Austro-Malayan are, Cacatuidæ and
Megapodiidæ. Yet although the birds are unmistakably Malayan, as a whole,
there are, as in the mammalia (though in a less degree), marked
deficiencies of most characteristic Malayan forms. Lord Walden gives a list
of no less than 69 genera thus absent; but it will be sufficient here to
mention such wide-spread and specially Indo-Malay groups as,--_Eurylæmus_,
_Nyctiornis_, _Arachnothera_, _Geocichla_, _Malacopteron_, _Timalia_,
_Pomatorhinus_, _Phyllornis_, _Iora_, _Criniger_, _Enicurus_, _Chaptia_,
_Tchitrea_, _Dendrocitta_, _Eulabes_, _Palæornis_, _Miglyptes_, _Tiga_, and
_Euplocamus_. These deficiencies plainly show the isolated character of the
Philippine group, and imply that it has never formed a part of that
Indo-Malayan extension of the continent which almost certainly existed when
the peculiar Malayan fauna was developed; or that, if it has been so
united, it has been subsequently submerged and broken up to such an extent,
as to cause the extinction of many of the absent types.
It appears from Lord Walden's careful analysis, that 31 of the Philippine
species occur in the Papuan sub-region, and 47 in Celebes; 69 occur also in
India, and 75 in Java. This last fact is curious, since Java is the most
remote of the Malayan islands, but it is found to arise almost wholly from
the birds of that island being better known, since only one species,
_Xantholæma rosea_, is confined to the Philippine Islands and Java.
The wading and swimming birds are mostly of wide-spread forms, only 6 out
of the 60 species being peculiar to the Philippine archipelago. Confining
ourselves to the land-birds, and combining several of the minutely
subdivided genera of Lord Walden's paper so as to agree with the
arrangement adopted in this work, we find that there are 112 genera of
land-birds represented in the islands. Of these, 50 are either
cosmopolitan, of wide range, or common to the Oriental and Australian
regions, and may be put aside as affording few indications of geographical
affinity. Of the remaining 62 no less than 40 are exclusively {348}or
mainly Oriental, and most of them are genera which range widely over the
region, only two (_Philentoma_ and _Rollulus_) being exclusively Malayan,
and two others (_Megalurus_ and _Malacocircus_) more especially Indian or
continental. Five other genera, though having a wide range, are typically
Palæarctic, and have reached the islands through North China. They are,
_Monticola_, _Acrocephalus_, _Phylloscopus_, _Calliope_, and _Passer_; the
two first having extended their range southward into the Moluccas. The
peculiarly Australian genera are only 12, the majority being characteristic
Papuan and Moluccan forms; such as--_Campephaga_, _Alcyone_, _Cacatua_,
_Tanygnathus_, _Ptilopus_, _Janthænas_, _
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