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Title: A New Subspecies of Lizard, Cnemidophorus sacki, from Michoacan, Mexico
Author: Duellman, William E., 1930-
Language: English
As this book started as an ASCII text book there are no pictures available.

*** Start of this LibraryBlog Digital Book "A New Subspecies of Lizard, Cnemidophorus sacki, from Michoacan, Mexico" ***

                 ~University of Kansas Publications~

                     ~Museum of Natural History~

                 Vol. 10, No. 9, pp. 587-598, 2 figs.

                              May 2, 1960

                      A New Subspecies of Lizard,
              Cnemidophorus sacki, from Michoacán, México


                          WILLIAM E. DUELLMAN

                        ~University of Kansas~

    ~University of Kansas Publications, Museum of Natural History~

           Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
                           Robert W. Wilson

                 Volume 10, No. 9, pp. 587-598, 2 figs.
                        Published May 2, 1960

                        ~University of Kansas~
                           Lawrence, Kansas

                              PRINTED IN
                       THE STATE PRINTING PLANT
                            TOPEKA, KANSAS


                      A New Subspecies of Lizard,
              Cnemidophorus sacki, from Michoacán, México


                          WILLIAM E. DUELLMAN

The systematic status of the populations of lizards assignable to
_Cnemidophorus sacki_ in western México (Sonora southward to Jalisco)
has been reviewed in detail by Zweifel (1959, Bull. Amer. Mus. Nat.
Hist, 117 (2):57-116), who stated that the use of the specific name
_sacki_ for the western populations rests upon a reasonable, but as
yet unproved, assumption that intergradation occurs. Although Zweifel
examined specimens of the nominal subspecies from the upper Balsas
Basin, he did not study specimens from the intervening area--the
Tepalcatepec Valley in Michoacán.

Field studies and the examination of large series of preserved
specimens of _Cnemidophorus_ from the Tepalcatepec Valley show the
presence of _Cnemidophorus sacki_ in the region between the ranges of
_C. sacki sacki_ in the upper Balsas Basin and _C. sacki occidentalis_
in Jalisco and southern Nayarit. Furthermore, the populations of
_sacki_ inhabiting the Tepalcatepec Valley have characters of
scutellation and coloration that distinguish them from other described
subspecies of _sacki_. In recognition of the important contributions
to the systematics of the genus _Cnemidophorus_ made by Dr. Richard G.
Zweifel, I propose that the subspecies of _Cnemidophorus sacki_ in the
Tepalcatepec Valley be named as follows:

             +Cnemidophorus sacki zweifeli+ new subspecies

_Holotype._--University of Michigan Museum of Zoology No. 119542, from
Capirio, Michoacán, México (185 meters), an adult male, one of a
series collected on June 13, 1958, by William E. Duellman, Jerome B.
Tulecke, and John Wellman. Original number WED 12310.

_Paratopotypes._--UMMZ Nos. 119536-119541 and 119543-119550.

_Diagnosis._--A race of _Cnemidophorus sacki_ characterized by large
size (more than 130 mm. snout-vent length in males), approximately 106
dorsal granules around the midbody, about 41 femoral pores, and a
dorsal color-pattern in adult males consisting of lateral and
dorsolateral rows of spots, paravertebral rows fused with middorsal
light green area at least anteriorly, and pink throat having a median
light blue spot or transverse band.

_Description of Holotype._--Snout-vent length, 128 mm.; tail length,
252 mm.; tail/body ratio, 1.97; scutellation typical of _sacki_--four
supraoculars, enlarged postantebrachials, and enlarged mesoptychials;
107 granules around midbody (excluding enlarged ventrals); 45 femoral
pores; three enlarged preanal scales; supraorbital semicircle-series
extending anteriorly to posterior edge of frontal (Fig. 1).

    [Illustration: Top View Head - Cnemidophorus sacki zweifeli]

~Fig. 1.~ Top view of the head of the holotype of _Cnemidophorus sacki
    zweifeli_ (UMMZ 119542) showing scutellation. ×5.

Top of head and nape dusty brown; tip of rostral and lateral edges of
superciliaries dark cream-color; upper labials and sides of head
anterior to eyes cream-color, mottled with blue; lower labials and
postocular region pale blue; mental, postmental, and sublabials
cream-color. Upper surfaces of forelimbs dull bluish gray, spotted
with pale greenish yellow; dorsal surfaces of proximal one-fourth of
tail light brownish gray turning to pale orange-brown posteriorly;
lateral surfaces of tail bluish gray anteriorly and creamy brown
posteriorly. Nuchal region light bluish gray; flanks dark gray; dorsal
ground-color dark brown, somewhat paler posteriorly. Body having a row
of cream-colored spots in place of a lateral stripe, and another row
in place of dorsolateral stripe; dorsally, large diffuse tan or light
green spots partially fused and tending to form irregular transverse
markings. Chin to posterior end of sublabials pale pink, bordered
posteriorly by bluish white area, and then pink. Mesoptychials, under
surfaces of hind limbs, and belly cream-colored; anterior edges of
belly scales dark blue; lateral two rows of ventrals on posterior
two-thirds of body dark blue having light blue or cream-colored spots.
Under surfaces of forelimbs bluish cream; ventral surface of tail

_Variation in Size and Scutellation._--The largest male has a
snout-vent length of 132 mm., the largest female, 114 mm., and the
smallest juvenile, 34 mm. The number of dorsal granules at the midbody
varies from 91 to 117 (106.2 ± 0.43); the ratio of the number of
granules between the paravertebral stripes to the number of granules
around the body (PV/GAB) varies from 0.064 to 0.157 (0.097 ±0.007);
the number of femoral pores varies from 32 to 49 (41.1 ± 0.20).
Usually there are only three enlarged preanals, but 18 specimens have
a somewhat enlarged scale anterior to the normal complement of three.
In 15 specimens the supraorbital semicircle-series terminate short of
the posterior edge of the frontal; in the others the series reach the

_Variation in Coloration._--The coloration of juveniles and subadults
varies little; large adults vary considerably especially in the amount
of diffusion of the light green middorsal area. In some individuals
the vertebral pale area does not include the paravertebral spots; in
other individuals the pale area includes not only the paravertebral
rows, but, at least anteriorly, the dorsolateral rows. In large males
of about equal size (and collected at the same time) there is
considerable variation in the amount of blue on the belly. In a few of
the males the belly is white with only the anterior edge of each scale
blue; in some only the lateral rows of ventrals on the posterior
two-thirds of the body are blue; in others all of the posterior
two-thirds of the belly is blue.

_Ontogenetic Change in Color Pattern._--The metamorphosis of color
pattern in _Cnemidophorus sacki zweifeli_ results in the dorsal
ground-color becoming paler with age, the replacement of the stripes
by spots, and finally in large males the suffusion of these spots.

A single hatchling (UMMZ 114732) is available; this specimen has a
prominent umbilical scar and a snout-vent length of 34 mm. The top of
the head is olive brown; the dorsal surfaces of the limbs are dark
brown with cream mottling; the dorsal ground-color is brownish black;
this is paler on the lower flanks. The lateral and dorsolateral
stripes are cream-colored; the paravertebral stripes are white. There
is a faint, diffuse vertebral stripe anteriorly (Fig. 2 A). The throat
and undersides of the limbs and tail are cream-colored; the belly is
bluish white. In life the stripes were pale yellowish green, and the
tip of the tail was pink.

In larger individuals the dorsal ground-color is dark brown; the
lower flanks are grayish tan. Light brown diffuse spots are present
in the lateral and dorsolateral dark fields. The tan vertebral stripe
is diffuse and nearly fills the paravertebral dark fields; the
paravertebral stripes are faint posteriorly; throughout their length
they are scalloped--the beginning of their fragmentation into spots
(Fig. 2 B).

In subadults (± 80 mm. snout-vent length) the paravertebral stripes
are fragmented into spots posteriorly. Also, the dorsolateral stripes
in some individuals are fragmented posteriorly. The dorsolateral dark
fields are somewhat paler than the lateral dark fields. Cream-colored
spots are present on the flanks. The mottling on the thighs tends
towards the formation of light spots (Fig. 2 C).

In small adults (± 100 mm. snout-vent length) the paravertebral
stripes are entirely fragmented into spots. The lateral and
dorsolateral stripes are broken into spots posteriorly. The middorsal
pale area (formed by the suffusion of the vertebral stripe) and
paravertebral and dorsolateral rows of spots are pale green. The
cream-colored spots on the flanks are expanded to form vertical bars
(Fig. 2 D).

Large adult males (± 120 mm. snout-vent length) have all of the
stripes fragmented into spots. The diffuse middorsal area is expanded
and encloses the paravertebral rows of spots. The pale spots present
in the dark fields in smaller individuals are either absent or fused
with spots resulting from the fragmentation of the stripes (Fig. 2 E).

_Sexual dimorphism._--Males attain a larger size (known maximum
snout-vent length of 132 mm., as compared with 114 mm. in females).
Males have larger but not more numerous, femoral pores, blue bellies,
and pink and blue throats, whereas females are unicolor creamy white
ventrally. The more nearly complete metamorphosis of color pattern
exhibited by adult males probably is correlated with their large adult
size. Large females retain complete lateral and dorsolateral stripes.
The jowls of breeding males are swollen.

           [Illustration: Color Pattern Change Diagram]

~Fig. 2.~ Diagrammatic representation of  ontogenetic change in color
    pattern in _Cnemidophorus sacki zweifeli_: A--hatchling, 34 mm.
    snout-vent length; B--juvenile,  55 mm. snout-vent length;
    C--subadult male, 80 mm. snout-vent length; D--small adult male,
    100 mm. snout-vent length; E--large adult male, 120 mm. snout-vent

_Geographic variation._--No noticeable geographic variation in this
subspecies is evident in the series from the Tepalcatepec Valley.
However, lizards from eastern Michoacán (Chinapa, 6 km. N of Tafetan,
6 km. S of Tzitzio, and 19 km. S of Tzitzio) differ slightly from
those from the Tepalcatepec Valley; the eastern specimens have fewer
dorsal granules and femoral pores, and a higher ratio of dorsal
granules between the paravertebral stripes to the number of granules
around the body (see Tables 1-3). No large adult males are present in
the eastern series; the subadults and small adult males have color
patterns like lizards of similar size from the Tepalcatepec Valley.
The largest male from the east has a snout-vent length of 110 mm.,
rows of pale spots, and no trace of brown and tan cross-bars.
Specimens of _Cnemidophorus sacki sacki_ of equal size from Guerrero,
Morelos, and Puebla in the upper Balsas Basin have a tan dorsum with
dark brown cross-bars. The localities in eastern Michoacán are
intermediate geographically between the Tepalcatepec Valley and the
known range of the nominal subspecies in the upper Balsas Basin. In
characters of scutellation specimens from the east are intermediate
between _C. sacki sacki_ and _C. sacki zweifeli_ in the Tepalcatepec
Valley. However, in coloration the lizards from the east are like
those from the Tepalcatepec Valley, but differ distinctly from the
nominal subspecies. Therefore, the eastern series is referred to the
subspecies _C. sacki zweifeli_.

_Comparisons._--Four other species of _Cnemidophorus_ occur in the
Tepalcatepec Valley with _Cnemidophorus sacki zweifeli_. Of these, _C.
calidipes_ has a maximum snout-vent length of 79 mm., 66 to 86 dorsal
granules, and a light brown dorsum with pale blue spots and vertical
bars; _C. communis communis_ has a maximum snout-vent length of 135
mm., 105 to 144 dorsal granules, and a greenish tan dorsum with yellow
spots; _C. deppei infernalis_ has a maximum snout-vent length of 84
mm., 91 to 120 dorsal granules, and a striped pattern throughout life;
and _C. lineatissimus exoristus_ has a maximum snout-vent length of 98
mm., 108 to 135 dorsal granules, and a middorsal yellow stripe and
vertical bars on the flanks. Both _calidipes_ and _communis_ are like
_sacki_ in possessing four enlarged supraoculars and enlarged
postantebrachials, whereas _deppei_ and _lineatissimus_ have three
enlarged supraoculars and granular postantebrachials. Juveniles of
_calidipes_ and _sacki_ are alike in coloration but different in the
extent of the supraorbital semicircle-series. In _calidipes_ the
supraorbital semicircle-series usually are complete, whereas in
_sacki_ the series never extended anterior to the posterior edge of
the frontal.

~Table 1.--Variation in the Number of Dorsal Granules
           in Three Subspecies of Cnemidophorus sacki~

Key for Table:
   No. = Number of Specimens  SD = Standard Deviation
   SE  = Standard Error   C/V = Coefficient of Variation

|   ~Population~           | No. |  Range  | Mean |  SD  |  SE  | C/V  |
|_sacki sacki_             |     |         |      |      |      |      |
|   Entire Sample          | 106 |  88-105 | 96.3 | 4.16 | 0.40 | 3.92 |
|   Puebla: Tehuitzingo    |  22 |  88-103 | 95.1 | 4.18 | 0.89 | 4.39 |
|   Guerrero: Chilpancingo |  23 |  90-105 | 95.8 | 3.86 | 0.80 | 4.02 |
|   Guerrero: Mexcala      |  22 |  90-102 | 96.5 | 3.56 | 0.76 | 3.69 |
|   Morelos                |  39 |  89-105 | 97.2 | 4.56 | 0.73 | 4.69 |
|                          |     |         |      |      |      |      |
|_sacki zweifeli_          |     |         |      |      |      |      |
|   Entire Sample          | 191 |  91-117 | 106.2| 5.98 | 0.43 | 3.13 |
|   Michoacán: Tafetan     |  21 |  91-116 | 101.4| 8.04 | 1.75 | 7.92 |
|   Michoacán: Apatzingán  | 170 |  95-117 | 106.8| 5.42 | 1.32 | 3.19 |
|                          |     |         |      |      |      |      |
|_sacki occidentalis_[A]   |  62 |  97-118 | 106.3| 4.72 | 0.60 | 7.61 |

[Footnote A: Data for _C. sacki occidentalis_ in Tables 1-3 are from
Zweifel (1959, Bull. Amer. Mus. Nat. Hist., 117: tables 1-3).]

Table 2.--Variation in the Number of Femoral Pores in Three
Subspecies of Cnemidophorus sacki

Key for Table:
   No. = Number of Specimens  SD = Standard Deviation
   SE  = Standard Error   C/V = Coefficient of Variation

|   ~Population~           | No. |  Range  | Mean |  SD  |  SE  | C/V  |
|_sacki sacki_             |     |         |      |      |      |      |
|   Entire Sample          | 106 |  32-44  | 36.2 | 2.42 | 0.25 | 2.28 |
|   Puebla: Tehuitzingo    |  22 |  33-41  | 36.7 | 2.36 | 0.50 | 6.43 |
|   Guerrero: Chilpancingo |  23 |  32-39  | 35.7 | 2.15 | 0.45 | 6.02 |
|   Guerrero: Mexcala      |  22 |  33-44  | 37.5 | 2.59 | 0.55 | 6.91 |
|   Morelos                |  39 |  32-40  | 35.4 | 2.19 | 0.35 | 6.18 |
|                          |     |         |      |      |      |      |
|_sacki zweifeli_          |     |         |      |      |      |      |
|   Entire Sample          | 189 |  32-49  | 41.1 | 2.77 | 0.20 | 1.46 |
|   Michoacán: Tafetan     |  19 |  33-43  | 38.1 | 2.61 | 0.60 | 6.85 |
|   Michoacán: Apatzingán  | 170 |   2-49  | 41.4 | 2.58 | 0.62 | 1.52 |
|                          |     |         |      |      |      |      |
|_sacki occidentalis_      |  67 |  32-45  | 38.8 | 2.46 | 0.30 | 3.67 |

~Table 3.--Ratio of Number of Granules Separating Paravertebral
           Stripes to Granules Abound Midbody (PV/GAB) in Three
           Subspecies of Cnemidophorus sacki~

Key for Table:
   No. = Number of Specimens  SD = Standard Deviation
   SE  = Standard Error   C/V = Coefficient of Variation

|   ~Population~           | No. |   Range   | Mean |  SD  |  SE  | C/V  |
|_sacki sacki_             |     |           |      |      |      |      |
|   Entire Sample          |  72 |0.101-0.205| 0.154| 0.052| 0.006| 0.073|
|   Puebla: Tehuitzingo    |  12 |0.140-0.205| 0.169| 0.052| 0.015| 0.433|
|   Guerrero: Chilpancingo |  16 |0.120-0.192| 0.157| 0.021| 0.005| 0.131|
|   Guerrero: Mexcala      |  21 |0.142-0.180| 0.159| 0.031| 0.007| 0.147|
|   Morelos                |  23 |0.101-0.180| 0.137| 0.023| 0.005| 0.100|
|                          |     |           |      |      |      |      |
|_sacki zweifeli_          |     |           |      |      |      |      |
|   Entire Sample          | 105 |0.064-0.157| 0.097| 0.070| 0.007| 0.067|
|   Michoacán: Tafetan     |  21 |0.103-0.157| 0.128| 0.039| 0.009| 0.186|
|   Michoacán: Apatzingán  |  84 |0.064-0.126| 0.089| 0.051| 0.005| 0.060|
|                          |     |           |      |      |      |      |
|_sacki occidentalis_      |  50 |0.086-0.183| 0.130| 0.021| 0.003| 0.042|

From the geographically adjacent populations of _sacki_ (_sacki_ and
_occidentalis_), _zweifeli_ differs in coloration and scutellation
(see Tables I-III). _Cnemidophorus sacki sacki_ has a dorsal pattern
in adult males of dark brown cross-bars on a tan ground-color. Both
_occidentalis_ and _zweifeli_ have variable, diffuse color patterns
in large adults, but _zweifeli_ differs from _occidentalis_ in having
a blue spot on the pink throat.

_Ecology._--In the arid Tepalcatepec Valley _Cnemidophorus sacki
zweifeli_ lives at elevations of 160 to 1300 meters. In the lower
parts of the valley the lizards live primarily in open scrub forests,
characterized by deciduous trees offering only partial shade from the
sun, especially during the prolonged dry season. Common trees in
this scrub forest are _Acacia cymbispina_, _Cercidium
plurifoliolatum_, _Mimosa distachya_, and _Prosopis juliflora_.

During the dry season (November through May) adult males apparently
aestivate; several large series collected in the winter include only
subadults and females. This absence of males is corroborated by
personal observations in the Tepalcatepec Valley in April and May. In
the summer rainy season the lizards are active in the morning and
again in the late afternoon; only _Cnemidophorus calidipes_ is active
during the heat of the midday. In some areas of the scrub forest
_Cnemidophorus sacki zweifeli_ is found in association with
_Cnemidophorus communis communis_. Throughout the scrub forest _C.
sacki zweifeli_ occurs with _C. deppei infernalis_. In some of the
more dense scrub forest, where _C. sacki zweifeli_ is not so abundant
as in the more open forest, it has been taken with _C. lineatissimus
exoristus_. In the open _Acacia-Cercidium_ associations on the valley
floor _C. sacki zweifeli_ occurs with _C. calidipes_.

This subspecies is not restricted to the scrub forest. On the lower
slopes of the Cordillera Volcanica in Michoacán _C. sacki zweifeli_
has been collected in open pine-oak forest near Zirimicuaro and

_Distribution._--_Cnemidophorus sacki zweifeli_ inhabits the valley
of the Río Tepalcatepec in Michoacán and probably extreme southwestern
Jalisco, and the western part of the Balsas Basin in Michoacán. No
specimens have been seen from extreme western Guerrero, but _C. s.
zweifeli_ may occur there.

_Specimens examined._--Catalogue numbers are preceded by abbreviations
of the name of the institution as listed in the acknowledgements.

_Cnemidophorus sacki occidentalis_, 22 specimens, as follows:
_Jalisco_: 8 km. E of Ameca, UMMZ 102045; 7 km. SE of Ameca, UMMZ
102046; Autlán, UMMZ 102044, 102219-21; 7 km. ESE of El Arenal, UMMZ
114736 (2); San Gabriel, UMMZ 102040, 102042-3. _Michoacán_: 2 km. ESE
of Jiquilpan, UMMZ 117557 (3). _Nayarit_: Ixtlán del Río, UMMZ 104747;
San José de la Conde, UMMZ 102047 (4); 5 km. N of Santa Isabel, UMMZ

_Cnemidophorus sacki sacki_, 108 specimens, as follows: _Guerrero_:
Chilpancingo, UMMZ 72426 (8), 73937 (7), 88422 (4); 8 km. W of
Chilpancingo, UMMZ 119144 (4); 5 km. N of Iguala, UMMZ 114712 (11);
15 km. N of Iguala, UMMZ 99039; Mexcala, UMMZ 114711 (10); 8 km. S of
Taxco, UMMZ 114709 (2). _Morelos_: Amacuzac, UMMZ 114716; 3 km. S of
Cuautla, UMMZ 99031 (9), 99917 (11): Río Cuautla, UMMZ 99038 (6); 5 km.
S of Temixco, UMMZ 114718 (12); _Puebla_: 5 km. SE of Izúcar de
Matamoros, UMMZ 112650 (4); 13 km. SE of Izúcar de Matamoros, UMMZ
117497 (2); 3 km. NW of Tehuitzingo, UMMZ 114714 (10); 1 km. N of
Teyuca, UMMZ 114713 (6).

_Cnemidophorus sacki zweifeli_, 207 specimens, as follows: _Michoacán_:
Apatzingán, CNHM 36966-8, 38969, 38971 (18), 38972 (50), UIMNH
36772-7, USNM 135967-8, 135970, 135972-3; 4 km. E of Apatzingán, UMMZ
85412; 6.5 km. E of Apatzingán, UMMZ 114731 (5); 5 km. W of
Apatzingán, KU 29289-90, 29293-7; 10 km. W of Apatzingán, UMMZ 114730
(4); 12.3 km. S of Apatzingán, UMMZ 112647; 16 km. S of Apatzingán, KU
29298; 14 km. SSW of Apatzingán, KU 29735, 29746, 29750-2; 10 km. W of
Buenavista, UMMZ 114719 (3); Capirio, UMMZ 112643, 114722 (2), 114733,
119536-50; 4 km. N of Capirio, UMMZ 112644 (3), 112645; 5.6 km. N of
Capirio, UMMZ 114732; 2 km. S of Charapendo, UMMZ 112639 (12);
Chinapa, UMMZ 119556 (2); Jazmin, UMMZ 114725 (2); between La Playa
and Volcán Jorullo, UMMZ 104748 (2); Limoncito, UMMZ 119552 (3);
14 km. S of Lombardia, KU 29299-301, 29303, 29305-11; 6 km. SW of
Nueva Italia, UMMZ 112640 (2); 2.7 km. S of Nueva Italia, UMMZ 112641
(4); 5 km. N of Nueva Italia, UMMZ 114721; Río Marquez, 10 km. S of
Lombardia, UMMZ 112642, 112646; Río Marquez, 13 km. SE of Nueva
Italia, UMMZ 114726; 6 km. N of Tafetan, UMMZ 119555 (18); 14.5 km.
E of Tepalcatepec, UMMZ 114720 (2); 6 km. S of Tzitzio, UMMZ 99199,
99200 (2); 19 km. S of Tzitzio, UMMZ 99154; Volcán Jorullo, UMMZ
104449 (4), 104750; Ziracuaretiro, UMMZ 114724; 3 km. NW of
Zirimicuaro, UMMZ 114723.

_Acknowledgments._--For the loan of specimens under their care I am
indebted to Doris M. Cochran, United States National Museum (USNM);
Norman Hartweg, University of Michigan Museum of Zoology (UMMZ);
Robert F. Inger, Chicago Natural History Museum (CNHM); and Hobart M.
Smith, University of Illinois Museum of Natural History (UIMNH). I
thank Ann S. Duellman, Richard E. Etheridge, Fred G. Thompson, Jerome
B. Tulecke, and John Wellman for their assistance in the field, Lorna
Cordonnier for the drawing reproduced as Figure 1, and Richard G.
Zweifel for helpful suggestions and criticism. Field work in México
was made possible by grants from the Penrose Fund of the American
Philosophical Society and the Bache Fund of the National Academy of
Sciences in co-operation with the Museum of Zoology of the University
of Michigan.

_Transmitted February 2, 1960._


   *   *   *   *

Transcriber's Notes.

This file was derived from scanned images. With the exception of two
typographical errors that were corrected, the original text is
presented. Figure 1 has the notation ×5 after the caption to let the
reader know that the image was enlarged by a factor of five.

Typographical Errors Corrected:

      Page 589, Paragraph 2: Tepalcatapec => Tepalcatepec

      Page 592, Paragraph 2: ground color => ground-color
         the hypented versionappearsn Page 596 in the paragraph
         following the table.

Emphasis Notation:

      _text_  -  italicized
      +text+  -  bold
      ~text~  -  small caps

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