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MARVARD
UNIVERSITY

POSTILLA

PEABODY MUSEUM
YALE UNIVERSITY

NUMBER 159 19 MARCH 1973

A REVISION OF LIZARDS OF THE GENUS
PRIONODACTYLUS, WITH A NEW GENUS
FOR P. LEUCOSTICTUS AND NOTES ON THE
GENUS EUSPONDYLUS (SAURIA, TEIIDAE)

THOMAS UZZELL

POSTILLA

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a

A REVISION OF LIZARDS OF THE GENUS PRIONODACTYLUS, WITH
A NEW GENUS FOR P. LEUCOSTICTUS AND NOTES ON THE GENUS
EUSPONDYLUS (SAURIA, TEIIDAE)

THOMAS UZZELL

Department of Biology and Peabody Museum of Natural History,
Yale University, New Haven, Connecticut*

(Received 2 May 1972)

ABSTRACT

Prionodactylus differs from Euspondylus in having a double row of widened
gular scales and keeled hexagonal scales; it is therefore removed from
synonymy with Euspondylus. Five species (P. vertebralis, P. dicrus new
species, P. manicatus, P. argulus, and P. eigenmanni) are recognized. P.
dicrus is related to P. vertebralis; in both, the loreal is separated from the
supralabials, the subdigital lamellae are not tuberculate, and the frontonasal
is not divided; P. dicrus differs in having two dorsolateral light stripes an-
teriorly that fuse to one posteriorly, rather than a single median light line
throughout. P. vertebralis is known from Pacific Ecuador and Colombia, the
Magdalena and Cauca valleys of Colombia, and from adjacent Panama and
Venezuela; Euspondylus ampuedae appears to be the same species (new
synonymy). P. dicrus occurs on the Amazonian slopes of central Ecuador.
P. manicatus has two subspecies. P. m. manicatus, with single tubercles on
the subdigital lamellae, no median collar scale, and a divided eyelid disc,
occurs in Amazonian Ecuador and northern Peru. P. m. bolivianus, with non-
tuberculate subdigital lamellae, a median collar scale, and an undivided
eyelid disc, occurs in Amazonian Bolivia and southern Peru. In both races,
the loreal touches the supralabials and the frontonasal is not divided. P.
argulus has a wide range in the Guianas and Amazonian South America, at
relatively low elevations, from northern Brazil along the slopes of the Andes
to Bolivia. It has a divided frontonasal, and the loreal touches the supra-
labials. P. eigenmanni occurs only in Amazonian Bolivia. It has a single
frontonasal, the loreal touches the supralabials, and the subdigital lamellae
may have both single and double tubercles.

*Present address: Academy of Natura! Sciences, Nineteenth and the Parkway,
Philadelphia, Pennsylvania 19103.

Prionodactylus spinalis and P. rahmi are placed in the genus Euspondylus
because they do not have a double widened row of gular scales.

Prionodactylus ocellifer is identified as Aspidolaemus affinis (new
synonymy).

Riolama, new genus, is proposed for Prionodactylus leucostictus. Riolama
differs from Prionodactylus in not having a double row of widened gular
scales, in having a superficial tympanum rather than a recessed one, in
having much depressed digits, except at the tips, and in having plicae rather
than folds on the anterior tip of the tongue. The plicae of the tongue indicate
affinities of Riolama with Ptychoglossus, Alopoglossus, and Ecpleopus. The
former two have completely plicate tongues. Riolama differs from Ecpleopus
by the much depressed toes, by the complete superciliary series, and by the
longitudinal rather than diagonal rows of ventral scales. R. leucosticta is the
only known species.

Specimens of Euspondylus maculatus (the type species of the genus) that
have been reported from Pacific coastal South America are apparently mis-
labelled. The species is reliably known from northern and central Amazonian
Peru.

Euspondylus stenolepis, on account of its completely plicate tongue and
smooth scales on the forelimbs, is referred to the genus Ptychoglossus (new
combination).

Keys to Prionodactylus and Euspondylus based on these changes are in-
cluded. The status of names proposed in or referred to Euspondylus and
Prionodactylus is summarized.

The genus Prionodactylus has as relatives Cercosaura, Pantodactylus,
Aspidolaemus, and Pholidobolus. Cercosaura is distinct in scutellation. The
other genera, particularly Aspidolaemus and Pantodactylus, may not be
distinguishable from Prionodactylus. Hemipenial features of Pantodactylus
and Cercosaura distinguish the individuals for which this organ was studied
from the individuals of the other genera for which hemipenes were
examined.

POSTILLA 159: 67p. 19 MARCH 1973

CONTENTS

ABSTRACT
I Introduction sb Bet dy arjpe arith Rae iacice Asa ee Ree eer een eRe Eon BBP 4
II A revision of the genus Nernodacivtis SB RIO peo 5
WIS SPEEICSPMEMOVEC SEROMD LT1OMODACEVIUS. cn... 0.0. vosc0s0ecencaccesdusueuadesteees 50
IV new eenus for! PrionOdactylusylCuCOStICIUS ©0000... .cccccccccerecnotecnenneenent ay
WAC OIMMIEMtS OMsthe EMS LUN PONAVIUS .o...cc.sccscyccss.0s0orvesssssuencitdeocnentors 56
Wile AmevetO) the Species Ol HUSPONRGVIUS® 2. occ. cs tesansvsesssennnssecnuneetaeteusPanne 62
VII Status of names proposed in or referred to Pronodacniue and
LEDS D IEG BOUTIN «2 el Mt sd ce I in ee ee 63

4 POSTILLA 159

I. INTRODUCTION

The genus Euspondylus, although one of the earliest named in Group II
(Boulenger, 1885) of the family Teiidae, remains one of the most ill defined,
largely because the type species of the genus, Euspondylus maculatus Tschudi
(1845), is essentially devoid of unusual external morphological features on
which a generic concept could be founded. As a result, Euspondylus became
a wastebasket into which many other taxa could be put. As the generic con-
cept gradually broadened, ever more distinctive taxa could be placed within
it, so that by the time of Burt and Burt’s (1931) study, one or more mem-
bers of at least six quite distinct genera had become included. Several of
these came into Euspondylus when the Burts merged Prionodactylus with
Euspondylus. This merger seemed justified partly because of the diversity of
taxa that had previously been included in Euspondylus and in Prionodactylus.
Gradually, however, these diverse stocks have been removed from Euspon-
dylus (Ruibal, 1952; Uzzell, 1959, 1961, 1969a, b).

The genus Prionodactylus can, I believe, be redefined to encompass five
taxa, one of which I here describe for the first time. Definition of Priono-
dactylus requires considerable reshuffling of other taxa. I have considerable
faith in some of the changes proposed here, but the collection of species
that remains in Euspondylus still seems artificial to me. The easy work, how-
ever, appears to be done.

ABBREVIATIONS
AMNH American Museum of Natural History, New York
BMNH British Museum (Natural History), London
CAS California Academy of Sciences, San Francisco
CM Carnegie Museum, Pittsburgh
FMNH Field Museum of Natural History, Chicago
IRSN Institut Royal des Sciences Naturelles de Belgique, Brussels
LACM Los Angeles County Museum
MCZ Museum of Comparative Zoology, Harvard University
MLS Museo de La Salle, Bogota
MNHN Muséum National d’Histoire Naturelle, Paris
NRM Naturhistoriska Riksmuseet, Stockholm
UIMNH University of Illinois Museum of Natural History
UKMNH University of Kansas Museum of Natural History
UMMZ University of Michigan Museum of Zoology
USNM United States National Museum
ZMB Zoologisches Museum, Berlin

ZSM Zoologische Staatssammlung, Munich

PRIONODACTYLUS AND EUSPONDYLUS 5

II. A REVISION OF THE GENUS Prionodactylus
Prionodactylus O’Shaughnessy

Prionodactylus O’Shaughnessy, 1881, Proc. Zool. Soc. London 1881: 231.
DEFINITION. Tongue with imbricate scalelike papillae. Snout moderate to
long, blunt to pointed. Head scales without striations or rugosities; single or
divided frontonasal; single frontal and interparietal; paired prefrontals and
parietals; a median and two paramedian occipitals. Nostril pierced in a single
or divided nasal; loreal and frenoocular present; first superciliary expanded
onto dorsal surface of head or not; prefrontals in contact with loreal.
Supraoculars two to four; superciliary series complete. Eyelids developed,
lower with a translucent or lightly pigmented disc, divided by vertical grooves
or not. Tympanum recessed, external ear opening small. Gular crease present,
weak. Collar fold well developed. Gular scales flat, rectangular, the two
median rows of scales, at least on the posterior part of the throat, forming
a double longitudinal series of widened scales. Limbs pentadactyl; digits
clawed. Forefoot with or without enlarged platelike scales along inner margin
of palm between thumb and wrist; if enlarged, the medial edge projecting
or not. Undersides of third and fourth toes usually with paired scales on
proximal part, the inner scale usually producing a tubercle. Dorsal scales
in diagonal and transverse rows, keeled, hexagonal. Lateral scales reduced
in size, forming a wide band between dorsal and ventral scales; upper rows
keeled. Ventral scales in transverse and longitudinal rows, smooth. Femoral
pores occasionally absent in females. Preanal scales in two rows.

REMARKS. The genus Prionodactylus was named for the tubercles under the
toes and fingers of northern populations of P. manicatus, the type species
of the genus. In these populations, the subdigital lamellae are angled on the
midline of the digit, and bear a single tubercle on the angle (Fig. 1). In

FIG. 1. Undersurface of the left forefoot of Prionodactylus m. manicatus (UKMNH
109819), showing the single medial tubercle of the basal lamellae of the third
and fourth digits. x 8.

6 POSTILLA 159

southern populations of P. manicatus and in P. vertebralis and P. dicrus the
subdigital lamellae are mostly single and are nontuberculate; in these, the
subdigital lamellae at the base of the third and fourth toes are doubled,
and the inner parts are swollen into tubercles. Prionodactylus eigenmanni
has single and double tubercles, as well as simple subdigital lamellae. In
P. argulus, most of the subdigital lamellae are paired, but not tuberculate.

The relationships of Prionodactylus appear to be with Cercosaura, Panto-
dactylus, Aspidolaemus, and Pholidobolus, with all of which it shares keeled
dorsal scales and a double longitudinal row of widened gular scales. Pri-
onodactylus may be distinguished from Cercosaura because Cercosaura has
large rectangular keeled dorsal scales in longitudinal rows. Pantodactylus,
as noted by Ruibal (1952: 520) may in fact be indistinguishable from
Prionodactylus. Aspidolaemus and Pholidobolus also may be indistinguish-
able from Prionodactylus, but at present I retain them as different genera
pending studies based on more than scalation.

The taxa that I recognize in the genus Prionodactylus may be distinguished
by the following key.

1. Frontonasal divided siagub suing essdjesdaes oe ceqeit ee eee oae IO EELELS.
Frontonasal single we elia «gaan ji ndan jeden ddqubesneahoonche ee Z
2. Loreal separated from supralabials by nasal and frenoocular.......... 3
Eoreal in contact with the supralabials 2.0.0. 4
3. A narrow middorsal light stripe from rostral to tail ....... P. vertebralis

Two narrow dorsolateral light stripes on head fusing to one at midbody
ae ae ae _ P. dicrus, new species

4. 32-35 scales around the midbody region; 13-15 subdigital lamellae
under fourth toe; some subdigital lamellae with one, others with two
tubercles .... Om a IC A a _ P. eigenmanni
33-52 scales iain che mrdbody region; ee 23 cubdigial lamellae
under fourth toe; subdigital lamellae, if tuberculate, with a single tu-

bercle: per’ lamella’ o.c...c.csscssevohteovs cesses dee 5}
5. 2 or 3 posterior preanal scales; no median collar scale; translucent disc
in lower eyelid divided into two segments _. P. m. manicatus

4 posterior preanal scales; a median collar scale; translucent disc in
lower eyelid undivided 29.35 an eee P. m. bolivianus

Although additional work may result in subdivision of some of the taxa
that I recognize, these are the basic stocks in the genus as I understand it.

Prionodactylus vertebralis (O’Shaughnessy )
(Fig. 2)

Cercosaura (Pantodactylus) vertebralis O’Shaughnessy, 1879, Ann. Mag. Nat.
Hist. Ser. 5, 4: 298.

PRIONODACTYLUS AND EUSPONDYLUS 7

Prionodactylus palmeri Boulenger, 1908, Ann. Mag. Nat. Hist. Ser. 8, 2: 518.
Prionodactylus marianus Ruthven, 1921, Occ. Pap. Mus. Zool. Univ. Michi-
gan 103: 1.
(2?) Euspondylus ampuedae Lancini, 1968, Publ. Ocas. Mus. Cienc. Nat.,
Zool. 12: 4.

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eee eee RI

FIG. 2. Side view of the head of Prionodactylus vertebralis (FMNH 43799). The
left loreal is fused to the nasal on this specimen. x 5.

HOLOTYPE. A young female (BMNH 78.1.25.13, reregistered as 1946.
8.31.35) collected by Buckley at Intac, Imbabura Province, Ecuador.

DEFINITION. A single frontonasal. Loreal separated from supralabials by con-
tact between nasal and frenoocular (98% )!. Two collar scales at midline,
usually widened. Lamellae under fingers not forming tubercles. Translucent
disc in lower eyelid divided into 3-5 parts’, often lightly pigmented. No
sexual dimorphism in femoral pore number. A conspicuous middorsal light
stripe the length of the body and along the unregenerated part of the tail.°

VARIATION. I have examined over 200 specimens of P. vertebralis. There is
considerable variation, within as well as between populations (Table 1).
Most of the specimens examined (88% ) have three supraoculars on each
side. One individual (FMNH 43806) has 4-4. Counts of 2-2 supraoculars
are relatively common (39% ) in the series from San Pedro, the type locality

1Loreal absent in the single Peruvian specimen (LACM 55881).

*Undivided in LACM 55881.

3 Absent in LACM 55881; possibly present only in male paratype of Euspon-
dylus ampuedae.

8 POSTILEA 159

of P. marianus. Eight longitudinal rows of ventrals are also relatively com-
mon in the series from San Pedro (24/43); this count occurs sporadically
elsewhere some 12 times.

There is some indication of a latitudinal gradient in whether or not the
prefrontals are in contact at midline. They more often are in contact in the
north (San Pedro, 36/41; Sonsdén, 16/27, with three additional specimens
having an azygous scale separating the prefrontals). Farther south, separated
prefrontals are more common (Pasto, 4/4 separated; Pichincha and Coto-
paxi, Ecuador, 5/9 separated). The series from “Mera” has 5/11 separated,
and the series from “Zamora” has 4/6 separated.

The two extreme southern specimens, AMNH 18312 from El Oro, Ecua-
dor, and LACM 55881 from Piura, Peru, lack prefrontal scales entirely, and
thus superficially resemble members of the genus Proctoporus. These two
specimens appear to carry to an extreme the reduction in prefrontal size as-
sociated with separation of the prefrontals.

As anomalies, occasional individuals have the median occipital missing, the
loreal fused to the nasal, frontonasal asymmetrically divided, and various
extra, minute head scales. These form no obvious geographic patterns.

Samples of individuals from two areas show notable variation. The speci-
mens from San Pedro, the type locality of P. marianus, fall into two groups.
Eight of the 10 paratypes of P. marianus that I have examined (UMMZ
56031, 8 specimens; CAS 54809-10) have 8 rather than 6 longitudinal rows
of ventral scales. Seven have 2-2 supraoculars, and one has 2-3. These speci-
mens also have fewer femoral pores. Nine additional specimens (MCZ
14652-57; USNM 75967, 120798; BMNH 1925.5.1.11) were collected by
Hno. Nicéforo Maria at about the same time and place as UMMZ 56031;
BMNH 1925.5.1.11 was obtained from the University of Michigan and has
the same collection data as the paratypes; it probably is a paratype. These
“topotypes” resemble the paratypes of P. marianus in having 8 longitudinal
rows of ventral scales, 2-2 supraoculars in 6 individuals and a low number
of femoral pores. In contrast, AMNH 32737-57 and MCZ 29860 and 46448
differ from the paratypes and similar specimens. Four of 23 have 2-2 supra-
oculars; 6 have 8 longitudinal rows of ventral scales; the femoral pore
number is greater (Table 2). A canonical analysis (BMDO7M; Dixon 1968)
based on total femoral pores, longitudinal rows of ventral scales, transverse
rows of ventral and of dorsal scales, scales around midbody region, and total
number of supraocular scales, groups 18 of the 19 paratypes and “‘topotypes”
together, and 18 of the 21 “non-types” together. The first two canonical axes
account for 90 per cent of the dispersion. Fig. 3 shows that the non-paratype
females are very variable; they include three of the misplaced individuals. I
conclude that the samples labelled San Pedro came from at least two dif-
ferent localities in that area.

Among the animals from near Popaydn, two quite distinct forms occur.
These can be separated by several scale counts and by total femoral pore
number (Table 1). The specimens differ also in color, with the specimens

PRIONODACTYLUS AND EUSPONDYLUS Y)

ia Paratypes
@ @ 'Topotypes"
O 9 "Non-types"

SECOND CANONICAL VARIABLE

-54 -39 -24 -09 0.6 Zal 36 5.
FIRST CANONICAL VARIABLE

FIG. 3. Distributions and means for individuals of three groups of specimens from
the type locality of Prionodactylus marianus (San Pedro, Colombia) along the
first and second canonical axes. The characters on which the analysis was based
include total femoral pores, longitudinal rows of ventral scales, transverse rows
of ventral and of dorsal scales, scales around midbody region, and total number
of supraoculars. The first two canonical axes account for 90 per cent of the
dispersion. Paratypes include UMMZ 56031 (8 specimens) and CAS 54809-10.
“Topotypes” include MCZ 14652-57, USNM 75967, 120798, BMNH 1925.5.11;
the last is probably a paratype. “Non-types” include AMNH 32737-57 and MCZ
29860 and 46448.

from Popayan (1700 m) being in general much lighter in coloration than
the specimens from El Tambo (2000 m). Part of the difference may be due
to preservation; the El Tambo specimens appear to be formalin blackened.
In the darker specimens, from El Tambo, the dark borders to the light mid-
dorsal stripe are essentially straight and parallel on the head, and separated
by the distance separating the supraocular series on the top of the head. In
the lighter colored specimens, however, the dark borders for the median
light line are much more irregular, and may run along the outer edges of
the occipitals, the parietals, the supraoculars and along the canthus rostralis.
Of the 10 light-colored specimens, 5 have the prefrontals separated; of the
12 darker-colored specimens, 2 have the prefrontals separated; the X? value
observed (2.79) indicates that this difference has a probability between 0.1
and 0.05 of occurring by chance.

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PRIONODACTYLUS AND EUSPONDYLUS 13

These two series were also compared in a canonical analysis using the same
characters used for the San Pedro specimens (Fig. 4). The first canonical
axis accounts for 91 per cent of the dispersion and for the major differences
between the samples. The second canonical axis includes largely sexual
dimorphism, and accounts for an additional 6 per cent of the dispersion.

In general, when other series are compared with those from near Popayan
by canonical analysis, the dark specimens remain distinct. Their greatest
similarity is with specimens from western Ecuador, especially in femoral pore
number and number of transverse rows of ventral scales (Table 1).

The specimen from Piura, Peru (LACM 55881) is remarkable in several
ways. It lacks prefrontals and loreals. It has no middorsal light line. Instead,
there are dorsolateral light lines with dark borders on the head and the
anterior part of the body, the lines and their borders about one dorsal scale’s
width wide; these lines disappear near midbody. The number of scales around

@@ EI! Tambo
OQ Popayan
+ Mean

Oo = us

SECOND CANONICAL VARIABLE

'

-5.l =o —2 1 -0.6 0.9 2.4 3.9 5.4
FIRST CANONICAL VARIABLE

FIG. 4. Distribution and means for individuals of two groups of Prionodactylus
vertebralis from near Popayan, Colombia, along the first and second canonical
axes. The characters on which the analysis was based include total femoral pores,
longitudinal rows of ventral scales, transverse rows of ventral and dorsal scales,
and total number of supraoculars. The second canonical axis reflects largely sexual
dimorphism. These two axes account for 97 per cent of the dispersion. The
Popayan sample includes FMNH 43794-803; the El Tambo sample includes
FMNH 43804-15.

14 POSTILLA 159

the midbody is lower than in most Ecuadorian material, especially compared
to AMNH 18312, from El Oro.

The absence of the middorsal light line gives this animal a pattern quite
unlike that of other specimens of P. vertebralis. Dorsolateral light lines are,
however, faintly suggested in AMNH 18312. The undivided disc in the
lower eyelid may reflect the common tendency for southern species and
populations of species of Group II of the Teiidae to have an undivided eyelid
disc. The absence of the loreal probably results from fusion with the nasal;
the pattern of scales on the side of the head resembles that seen in P. verte-
bralis, except that the first superciliary scale, rather than the loreal, is sepa-
rated from the supralabials by contact between the frenoocular and nasal.

Eventually, this specimen may be referred to a distinct taxon. In the
absence of additional material, and because of its similarities to AMNH
18312, which has faint dorsolateral light lines on the head and anterior part
of body and which lacks prefrontals. I consider it a specimen, although
highly unusual, of P. vertebralis.

COLORATION. Most specimens of this species appear to be badly blackened.
In life they appear to be considerably more attractive. James A. Peters kindly
described the color and pattern of two specimens from Tandayapa, Pichincha,
Ecuador, that I have not examined, as follows. Middorsal stripe bronze on
snout, darker between eyes, lightening to light bronze or light tan, and

TABLE 2. Characters of three groups of Prionodactylus vertebralis from
San Pedro, Antioquia, Colombia.

Total Ventral Dorsal Scales
femoral scale scale around
pores TOWS rows midbody region
Paratypes
626 0-6 20-21 30-34 S337
(2-7) (20.8) (31.8) (3:2)
42° 0-5 20-21 31 33-36
CEe7) (20.5) (3120) (34.2)
“Topotypes”
BIO 0-11 20-21 32-34 32-42
(2:2) (20.4) (32.8) (35.2)
42° 0-3 21-23 30-34 3137
(0-7) @iIES) (32.0) 327)
““Non-types”’
8 34 0-11 20-23 31-35 34-42
G7) (2165) (32.0) (39.0)
1699 0-9 20-23 51235 35-41
(3:33) @1ES5)) @2:9) (37.0)

Paratypes = UMMZ 56031 (8 specimens, CAS 54809-10); “topotypes” =
MCZ 14652-57, USNM 75967, 120789, BMNH 1925.5.1.11; “non-types” =
AMNH 32737-57, MCZ 29680, 46448.

PRIONODACTYLUS AND EUSPONDYLUS 15

finally almost to white over sacrum. Three to four scales on either side of
middorsal stripe jet black; rest of side olive brown. Ocelli along sides jet
black, with bluish-white centers. A reddish area around first two ocelli on
neck; another over sacrum and on lateral edge of tail. Ocelli all along side
onto tail. Forelimb dorsally has black spots with bright yellow centers; re-
mainder of limb olive brown. Hind limb olive brown with black mottling.
Dorsum of head on either side of middorsal line black, shading to dark brown
laterally. A bronze line over eye, another from nostral along labials, below
tympanum to shoulder; behind the shoulder, much lighter, a dirty yellow,
broken into spots each with an irregular black border. The chin and throat
a soft chocolate tan; entire chest, belly, anal region, tail, and lower side of
hindlimbs a vivid orangy-red, spotted and dotted with black. Ventral side of
forelimbs olive-brownish. Iris a deep glowing red.

The largest male examined is 63 mm snout to vent; the largest female, 68
mm. Twenty-five males with tails intact have tail over snout-vent length
ratios of 1.7—2.3, mean 1.99; 20 females have a range of 1.7—2.2, mean, 1.86.

SEXUAL DIMORPHISM. This species shows relatively little sexual dimorphism,
since the femoral pores number about the same in males and females. There
is probably considerable sexual difference in color in living individuals, but
most of the material examined is badly blackened after preservation.

REMARKS. The holotype of P. vertebralis came from Intac, Imbabura, Ecua-
dor. Although some of the locality data for specimens collected by Buckley
appear to be incorrect (Peters 1955), that Intac is the correct locality is
supported by recent collections of specimens from several localities close to
Intac in western Ecuador (Fig. 5).

There are records of P. vertebralis from eastern Ecuador. AMNH 60586-
97, reportedly from Mera, Tungarahua, at about 1000 m, are typical of the
species; almost certainly they are incorrectly labelled.4 Another record,
BMNH 1933.6.24.82-87 from Zamora in Chinchipe-Zamora (Parker 1934)
may be based on specimens from Loja rather than Zamora, since many of
the specimens in that collection of Carrion’s came from Loja.

The altitudinal range of P. vertebralis in Ecuador (7O0O—1600 m) suggests
that it should be confined to one side of the Andes or the other. A closely
related species, found on the eastern Andean slopes, is described below.
Most of the recent records for P. vertebralis come from the western Andean
slopes. I doubt records from the eastern slopes.

The holotype of P. palmeri Boulenger (1908), BMNH 1909.4.30.63 (re-
registered as 1946.8.31.43) is an adult female collected from San Antonio,
Valle del Cauca, Colombia by M. G. Palmer. Boulenger’s description and
figure are excellent. The holotype falls within my concept of P. vertebralis.

‘James A. Peters (personal communication) has drawn a similar conclusion
about AMNH 49943-46, 49948-50, specimens of Atelopus longirostris collected
by C. Ollala, and supposedly from Mera.

16 POSTILLA 159

LaaN

we
\ ss

—

Ne coe

@ Pvertebralis
A P dicrus

FIG. 5. Northwestern South America, showing ranges for Prionodactylus vertebralis

and for P. dicrus, new species. Certain localities discussed in the text are also
indicated.

UMMZ 56037, the holotype of P. marianus Ruthven (1921) was col-
lected by Hno. Nicéforo Maria near San Pedro, Antioquia, Colombia. I have
examined this specimen and it falls within my concept of P. vertebralis.

There is some question about the number of paratypes of P. marianus.
Ruthven gave no indication of the number of specimens that he examined.
The paratypes recorded at the University of Michigan Museum of Zoology
originally included 13 catalogued as 56031. Two of these were sent to the
California Academy of Sciences (CAS 54809-10); one was sent to the

PRIONODACTYLUS AND EUSPONDYLUS 17

Field Museum of Natural History (FMNH 109860). There are still 8
specimens catalogued as 56031 at the University of Michigan. Two of the
original paratype series are thus not accounted for; one is probably BMNH
19255-1001.

Although they are not included in Barbour and Loveridge’s (1929) list of
typical material in the Museum of Comparative Zoology, Barbour and Love-
ridge (1946) listed 10 specimens (MCZ 14651-60) as paratypes of P. mari-
anus, and indicated that they were received as an exchange from the Uni-
versity of Michigan. These specimens apparently were examined by Ruthven
before he described P. marianus (letter to T. Barbour, 10 June 1921). Ruth-
ven clearly indicated, however, that many Colombian specimens, presumably
including MCZ 1465-60, were sent by Barbour to Ruthven for identification.
Ruthven identified the MCZ Prionodactylus as conspecific with his new form.

Ruthven stated that two of the paratypes have 3-3 supraoculars. I identify
these as UMMZ 56031C and CAS 54809. Since MCZ 14653 also has 3-3
supraoculars, it seems likely that this individual, and thus the entire series,
was not considered a paratype by Ruthven. If this is true, USNM 120789,
listed by Cochran (1961) as a paratype of P. marianus, likewise is not, since
it was formerly MCZ 14660.

I have been unable to examine any specimens of Euspondylus ampuedae
Lancini (1968) and Lancini’s description does not describe several features
that I consider diagnostic of genera and species of teiid lizards of Group II.
I believe, however, that Lancini has redescribed Prionodactylus vertebralis,
and that E. ampuedae should be placed as a junior synonym of that species
(new synonymy). My main reason for thinking that E. ampuedae is con-
specific with P. vertebralis is the pattern of the light lip line, described by
Lancini as beginning on the mental, crossing the middle of the second
supralabial, the upper border of supralabials 3, 4, and 5, and continuing
to below the ear opening and above the forelimb insertion. This pattern is
duplicated in P. vertebralis (Fig. 2) and in P. dicrus, new species, but in no
other Prionodactylus. Most of the other described features of E. ampuedae
can be matched in the sample of P. vertebralis that I have examined. Few
of these features, however, are diagnostic of any species. I interpret the two
loreals to include a superior loreal separated from the supralabials by an
inferior frenoocular.

Two characteristics of E. ampuedae suggest that the present synonymy is
not correct: the number of scales around the midbody region (27-29) re-
ported by Lancini is less than I have observed in P. vertebralis (31-45 in
Colombian specimens); no middorsal light line was described for the holo-
type and female paratypes of E. ampuedae, although such a line is almost
always present in P. vertebralis. Both of these differences could easily be
accounted for, however; the former by a different convention in counting
the scales, especially the minute lower lateral scales; the latter by desquama-
tion of the specimens, which makes the middorsal light stripe almost invisible
while leaving the lip line conspicuous. The male paratype, however, does

18 POSTILLA 159

have a middorsal light stripe. Alternatively, E. ampuedae may be a distinct
taxon.

E. ampuedae was described from Villa Paez, Estado Tachira, Venezuela
(Fig. 5).

BIOLOGY. Virtually no biological data are available to me. One specimen
from Pichincha was collected in the debris of a large log near a small, fast-
flowing stream. Many of the females contain two leathery-coated eggs, one
in each oviduct.

RANGE. Specimens of Prionodactylus vertebralis are known from the Pacific
slopes of Ecuador and northern Peru at elevations of 700 to 1600 m above
sea level, from the Cauca and Magdalena valleys of Colombia at elevations
of 1500 to 2500 m, from far southern Panama, and possibly from Estado
Tachira in Venezuela.

Specimens Examined

COLOMBIA: ANTIoQuiA, Jeric6 (1967 m) USNM 84967, 92496; Las
Palmas, SE of Medellin: FMNH 63824-25; Medellin (1538 m) AMNH
32758-59, 32772-73, 32775; Rio Negrito; 15 km E of Sons6én: FMNH
63823; Rio Negro (2120 m) AMNH 38950-53; San Pedro (2560 m)
UMMZ 56031 (8 specimens), CAS 54809-10, paratypes of Prionodactylus
marianus; BMNH 1925.5.1.11, MCZ 14652-57, 29680, 46448, USNM 75967,
120789, AMNH 32737-5S7; Santa Rosa de Osos (2640 m) AMNH 32766-68;
Sons6n (2545 m) AMNH 32725-36, 35302, UMMZ 57702 (4), 89421
(10): Yarumal (2300 m) FMNH 63821-22. Ca.pas, Pueblorrico (1560 m)
BMNH 1910.7.11.19-21. Cauca, Munchique, near El Tambo (2000 m)
FMNH 43804-15; Popayan (1700 m) FMNH 43794—-803. CUNDINAMARCA,
Arracachal (200 m) UMMZ 131677-78. NaRINo, Pasto (2594 m) AMNH
91768-71. TOLIMA, no other locality: UMMZ 56441 (4), 63801 (4); Quindio
Mountains: MCZ 15953-57, 15959, USNM 75968. VALLE DEL Cauca, Rio
Riposa Virology Field Station, USNM 151620; San Antonio (1850-2200 m)
BMNH 1909.4.30.63, reregistered as 1946.8.31.43, holotype of Prionodac-
tylus palmeri.

ECUADOR: Coropaxt, below Sigchos in Toachi Valley (1000 m) USNM
193944. Et Oro, El Chiral (1350 m) AMNH 18312. ImsBaBura, Intac
(1000 m) BMNH 78.1.25.13, reregistered as 1946.8.31.35, holotype of
Cercosaura vertebralis. Pastaza, Mera (1000 m) AMNH 60586-97.
PICHINCHA, Mindo (1050 m) UMMZ 119714; 3 km E of Nanegal Chico
(1600 m) USNM 193946; Pacto (1000 m) USNM 193941-43; below Pacto
(900 m) USNM 194065; Palma Real (900 m) USNM 193945; Pandayacu
USNM 193940; Rio Blanco near mouth of Rio Yambi (700 m) USNM

PRIONODACTYLUS AND EUSPONDYLUS 19
193947; Rio Caoni (200-300300 m) USNM 193939. Zamora-Chinchipe, Za-
mora (800 m) BMNH 1933.6.24.82-87.

PANAMA. Darien, Cerro Mali UKMNH 76174-75.

PERU: Piura, 16 km E of Canchaque LACM 58811.

Prionodactylus dicrus, new species
(Fig. 6)

Euspondylus festae, Burt and Myers, 1942:319.

HOLOTYPE. FMNH 36708, an adult female from Ecuador, Tungurahua, Ma-
poto; 1300 m; June 1938; William Clarke-MaclIntyre.

PARATYPES. FMNH 28043, 134152, USNM 193592: Ecuador, Tungurahua,
Banos (1800 m); BMNH 1912.11.1.34: Topo (1500 m); AMNH 24144,
CAS-SU 8253: Pastaza, Abitagua (1100-1500 m); USNM 193590-91: head-
waters of Rio Arajuno (about 600 m); USNM 194383: between Bafios and
Puyo (900 m).

FIG. 6. Side view of the head of Prionodactylus dicrus, new species (USNM
194383, paratype). <6. The loreal is separated from the supralabials by contact
between the nasal and frenoocular. The light labial line is less conspicuous than
in P. vertebralis, but follows essentially the same course.

20 POSTILLA 159

DIAGNOSIS. A member of the genus Prionodactylus closely allied to P. verte-
bralis in having an undivided frontonasal and the loreal scale separated from
the upper labial scales by contact between the nasal and frenoocular; these
two features distinguish P. vertebralis and P. dicrus from all other species of
the genus. P. dicrus differs from P. vertebralis in that it has two light lines
on the snout, superciliary scales and anterior parts of the body that fuse
posteriorly to form a median light line, rather than a median light line from
the snout to the tail; and marked sexual dimorphism in femoral pore number
(20-25 in males of P. dicrus, 7-11 females; 0-16 in males and females of
P. vertebralis. Subdigital lamellae of forefoot without tubercles (Fig. 7).

DESCRIPTION OF HOLOTYPE. Rostral broadly in contact with frontonasal, nar-
rowly in contact with nasal and first supralabial. Frontonasal wider than long.
Two prefrontals forming a short median suture. Frontal length about 1.4
times breadth, in contact with prefrontals, second supraoculars, and fronto-
parietals. Paired prefrontals, about twice as long as broad, in contact
medially. Interparietal 1.4 times as long as broad. Parietals slightly longer

‘“

FIG. 7. Underside of left forefoot of Prionodactylus dicrus, new species (USNM
4688, paratype) showing the nontuberculate subdigital lamellae. <7.

PRIONODACTYLUS AND EUSPONDYLUS 21

than wide, each in contact with interparietal, prefrontal, fourth supraocular,
a small postocular, a lateral (narrowly) and medial temporal (broadly), and
a paramedian occipital scale. A median occipital, a pair of paramedian oc-
cipitals, and a largish pair behind these that could be called postoccipitals.
Four supraoculars on each side, the anteriormost small, second largest, fourth
slightly larger than third. Superciliary series complete, 4 on right, 5 on left,
the first not expanded onto dorsal surface of head. Nasal scale low, elongate,
extending past middle of second supralabial, apparently divided on right,
entire on left. A narrow, diagonally placed loreal separated from supralabials
by contact between nasal and small, triangular frenoocular. Loreal bounded
by nasal, prefrontal, first supraocular (narrowly), first superciliary, and
frenoocular. Three (left) or 4(right) very narrow suboculars. A diagonally
placed row of 3 somewhat larger postoculars. A pigmented disc in the lower
eyelid, divided into four sections by vertical grooves. Seven supralabials on
each side, the space between the sixth and seventh and the lateral temporal
scale filled with about three longitudinal rows of elliptical scales. These and
the temporal scales weakly marked with fine wrinkles. External ear opening
round; tympanum deeply recessed, unpigmented, larger than external ear
opening. Mental followed by 1 unpaired and 3 (left) or 4 (right) paired
chin shields, all in contact with infralabials, the first (left) meeting the first
and second (right) at midline, the posterior ones separated by pregulars.
Pregulars (Ruibal, 1952) irregular, smooth, smallest on midline, in chevrons,
not forming transverse rows. Gular crease distinct, marked by small scales.
Gulars smooth, rounded posteriorly, larger in anteriormost row; a row of
seven pairs of widened gular scales on midline. Collar distinct; scales smooth,
rounded posteriorly, six in number including a large median pair.

Median anterior dorsal scales almost square, wrinkled; remainder of dorsal
body scales elongate, hexagonal, keeled, with minute ridges that converge
toward posterior tips of keel (Fig. 8); in 33 rows between occipital and
posterior margin of hind limbs. Lateral scales on neck, in axilla, and in
groin small granules; a weak fold from tympanum to lateral edge of collar.
Lateral body scales between limb insertions in two sizes; above, a zone about
6 scales high in which larger lateral rows are intercalated between ends of
transverse rows of dorsals; below, a granular series, almost a groove, three
to five scales high. Ventrals smooth, quadrangular, rounded at posterior
corners, in 8 longitudinal and 20 tranverse rows between collar and anterior
preanal scales. Two rows of smooth preanal scales, the posterior row with
four elongate medial and two minute lateral scales, the 2 median widest; the
anterior row with 2 relatively larger medial and 2 smaller lateral scales.
Femoral pore series divided; a single pore near each knee, 2 (right) and 3
(left) minute pores near groin, the innermost virtually a preanal pore. Pores
usually in center of scale.

Scales on forelimb large, very weakly keeled, weakly wrinkled on anterior
and dorsal surfaces, granular beneath. Subdigital lamellae undivided, slightly
swollen, but not forming a series of projections (Fig. 7). Scales along margin

22 POSTILLA 159

FIG. 8. Dorsal scales of Prionodactylus dicrus, new species USNM 194383, para-
type) showing keel and minute ridges converging posteriorly. x 20.

between thumb and wrist conspicuously enlarged, with a free posterior edge.
On palm scales granular. Dorsal thigh scales large, keeled and wrinkled;
anterior and ventral scales smooth; posterior scales granular. Anterior shank
scales large, keeled and wrinkled; ventral scales very large, smooth; posterior
scales granular. A row of intercalated scales on posterior side under bases of
digits one through four, the subdigital lamellae in these areas forming
tubercles. Digits weakly compressed, claws relatively robust, curved. Caudal
scales above, like dorsals, below, like ventrals, forming complete rings around
tail.

Color and pattern generally obscure; ground color olive; a pair of thin
light lines faintly discernable beginning on rostral, following canthus ros-
tralis, superciliaries, and median temporals, becoming conspicuous on body
as light bluish lines a single dorsal’s width wide, gradually converging
posteriorly, fusing at the nineteenth transverse row of dorsal scales, and con-
tinuing posteriorly to tip of unregenerated tail as a light line 2-3 scales wide.

PRIONODACTYLUS AND EUSPONDYLUS 23

Another, even thinner light line, beginning at edge of first supralabial, con-
tinuing along dorsal edge of third supralabial, across remaining supralabials
to lower edge of tympanum, along the weak neck fold, along the body above
the arm insertion and along upper edge of laterals almost to groin. Ventrally,
a uniform lead gray.

VARIATION. A median occipital is present in all individuals, the prefrontals
are in contact in all, and the loreal is separated from the infralabials in all.
Eight of the specimens examined have 4-4 supraoculars; USNM 193590 and
FMNH 28043 have 3-3; on the left in the former the location of a fourth,
posterior supraocular is indicated by grooves that do not meet in the center
of the very large third. The first supraocular is very small in all, and oc-
cupies the space on the top of the head usually filled in Prionodactylus by a
dorsal expansion of the first superciliary; as a consequence that scale ends
at the canthus rostralis. There is a pair of large median collar scales in all
specimens. The femoral pores in the females number far fewer than they do
in males (Table 3). The series in females is divided in all cases, with at least
one essentially preanal pore and at least one near the knee. There are enough
scales between these sets so that if all had pores, there would be little sexual
dimorphism in total number. One specimen (USNM 193590) has 6 rather
than 8 longitudinal rows of ventral scales. The largest male examined is 40
mm snout to vent, the largest female, 53 (the holotype). Other variational
data are included in Table 3.

RELATIONSHIPS. The relationships of P. dicrus are with P. vertebralis. This is
indicated not only by the features mentioned in the diagnosis, but also by
numerous details of structure. A striking feature shared by these two taxa is
the conspicuous wrinkling on many of the scales, especially the dorsal body
scales (Fig. 8).

Specimens of P. vertebralis have been reported from the Amazonian slopes
of the Andes at Mera in Pastaza (AMNH 60586-97) and from Zamora in
Chinchipe-Zamora (BMNH 1933.6.24.82-87; Parker 1934). While it is pos-
sible that P. vertebralis is sympatric with P. dicrus near Mera, I doubt it.
I suspect that the American Museum specimens are mislabelled. The alti-

TABLE 3. Characteristics of 10 specimens of Prionodactylus dicrus, new
species. Figures represent ranges and, in parentheses, means.

Scales
Total Dorsal Ventral around Subdigital lamellae
femoral scale scale midbody Fourth Fourth
pores rOWs rows region finger toe

536 20-25 Hg 3) 20-21 38-42 iPS =il9/ 20-235
@22) (30.4) (20.8) (40.2) (15.8) (21.0)

x5 22 tel 31-33 2OR22, 38-43 ily) DAD 2
(8.4) (126) (20.8) (40.8) (16.0) (21.3)

24 POSTILLA 159

tudinal range of P. vertebralis in Ecuador (up to about 1600 m) suggests
that it would be confined to one side or the other of the Andes. I believe it
probable that the specimens reported by Parker from Zamora actually came
from Loja in the province of Loja, some 50 kilometers to the west, whence
much of the material collected by Carrion and reported in the same paper
came; this would be more consistent with other distributional records. On
the whole, I am more inclined to believe the Zamora record than the Mera
record, since there are numerous collections from Mera, ‘and no other in-
dividuals of P. vertebralis have turned up in that area. Additional collecting
near Zamora, where the fauna is less well known, may yield new specimens
of P. vertebralis.

The derivation of P. dicrus and P. vertebralis trom a common ancestor
as a result of isolation of populations on each side of the Andes seems quite
likely. The time or area of the transgression by some ancestral population
are not known. The patterns of P. dicrus and P. vertebralis could easily be
changed one into the other. Many specimens of P. vertebralis have already
some lightening along the superciliary series.

DERIVATION OF NAME. The name is from the Greek 8:xpos, forked. It is used
as an adjective modifying Prionodactylus.

RANGE. Specimens of Prionodactylus dicrus are known from elevations of
perhaps 600 to perhaps 1800 m above sea level on the eastern Andean slopes
of central Ecuador.

Specimens Examined
These include only the holotype and paratypes.
Prionodactylus manicatus (O’Shaughnessy )

DEFINITION. A single frontonasal. Loreal almost always in contact with
frontonasal and supralabials. At least some indication of a light labial line
beginning under eye and continuing to tympanum. Subdigital lamellae either
smooth or with a single median tubercle.

VARIATION. I have examined 48 specimens that I refer to this species. These
come from localities in Amazonian Ecuador, Peru, and Bolivia, and from
the Bolivian Chaco (Fig. 9). The sample shows considerable geographic
variation (Table 4); I have grouped the populations, somewhat arbitrarily,
into two subspecies. Geographical variation in several characters is discussed
below.

1. Posterior preanal scales. Usually females of teiid lizards of Group II
have more posterior preanal scales than males have. Although there is slight

PRIONODACTYLUS AND EUSPONDYLUS 25

sexual dimorphism in this character in P. manicatus, the dominant trend is
a geographical one, independent of sex. Specimens from Ecuador usually
have 2 large lateral and 1 narrow median posterior preanal scale. A single
male (MCZ 45779) has only 2 posterior preanals. Two females have addi-
tional slivers lateral to the 3 major preanal scales. Three posterior preanals
also occur in specimens from northern and central Peru. The female from
Oxapampa and the two from Divisoria have the additional lateral slivers. All
11 specimens for which I have data from Bolivia and southern Peru have
4 posterior preanals; they are almost equally wide.

2. Enlarged collar scales. Specimens from Ecuador have 2 large collar
scales that form almost the entire collar. These 2 large scales form a con-
tinuation of the pairs of large gular scales, although they are wider than the
widened gulars. Two widened scales also occur in AMNH 90673, from
Luisiana in central Peru. The Bolivian and the remainder of the Peruvian
specimens have a median and 2 paramedian collar scales of about equal
size, except for UMMZ 96547 and FMNH 40424, in which the median

TABLE 4. Characters of 33 specimens of Prionodactylus manicatus.
Figures are ranges and, in parentheses, means.

Scales
Total Dorsal Ventral around Subdigital lamellae
femoral scale scale midbody Fourth Fourth
pores rows rows region finger toe
P. m. manicatus
Ecuador and Northern Peru
IOs 26-30 39-42 19-20 43-52 10-13 14-18
(27-9) (40.3) (19.3) (46.6) Glie3) 1(15:8)
Sao) 9 24-28 37-40 17-20 37-49 10-12 14-17
(26.6) (39.1) (18.9) (44.3) (Lica) (Gtse7)
Divisoria
ile Pil 34 20 43 12 16-17
2239 16-17 32-34 19-21 41-42 12 17-18
Oxapampa
ir? 14 34 18 38 12 --
Luisiana
je) 15 36 17 40 2 17-18

P. m. bolivianus
Southern Peru

4366 15-19 33-38 17-20 36-40 12-15 17-21
(1720) (36.3) (18.7) (38.3) (13:6) (1938)
622 5-9 35-37 17-19 35-41 13-15 17-23
(6.8) (35.7) (18.5) (37.8) (1422), 4(20:6)

Bolivia
1 2 16 36 20 36 13-14 21-22
2291 3-7 35-40 18-21 33-41 14-15 20-23

1{talicized numbers indicate holotype of P. m. bolivianus.

26 POSTILLA 159

o—

>
BEKGalal

Pallatanga | @ Pm bolivianus
sc — a ae \ Z : (e) 200
HL Ute ; Nes a —<| Kilometers
—e@ CO ‘ io
e
Ne, Z
‘ wi
4°, a ie

Ber

16°

82° 78° 74° 70° 66° 62°

FIG. 9. Collection localities for Prionodactylus manicatus in Ecuador, Peru, and
Bolivia. Localities mentioned in the text are also indicated.

scale is fused with one of the lateral collar scales in an asymmetrical arrange-
ment.

3. Disc is lower eyelid. The translucent disc in the lower eyelid is unpig-
mented in all specimens examined. In Ecuadorian and northern and central
Peruvian specimens, the disc is divided into 2 or 3 segments by vertical
grooves, except for FMNH 134398. The Bolivian and southern Peruvian
material has the disc in the lower eyelid undivided except for FMNH 40424
and BMNH 1946.8.31.37, one of the syntypes of Prionodactylus okendeni.
This pattern of geographic variation, divided eyelid discs in the north, un-
divided ones in the south, also occurs in other widespread teiids of Group II
(Cercosaura ocellata and Neusticurus ecpleopus; Ruibal, 1952; Uzzell, 1964).

4. Longitudinal rows of ventral scales. The number recorded is the number
of longitudinal rows at midbody not interrupted by smaller scales for at least

PRIONODACTYLUS AND EUSPONDYLUS 27

four transverse rows. Most of the specimens from Ecuador have 6 longi-
tudinal rows if the criterion is strictly interpreted, but in two, the lateral
rows would be counted as rows of ventrals by most observers, and a third
specimen clearly has eight rows near midbody. The northern and central
Peruvian specimens generally have 8 longitudinal rows, except for AMNH
90673 from Luisiana in central Peru, which has 6. Bolivian and southern
Peruvian specimens have 8 longitudinal rows of ventrals except for BMNH
1946.8.31.37.

5. Coloration. In Ecuador, at one extreme of development, there is a
light line from the first supralabial along the lips, through the lower edge of
the external ear opening, above the arm, along the side, finally disappearing
near the hind limb insertion. This light line is bordered below by dark from
the third infralabial posteriorly. More usually, this band begins on the first
supralabial and terminates just posterior to the forelimb insertion; it is con-
stricted or broken where it crosses the lateral extention of the collar. The
northern and central Peruvian specimens also have this less well-developed
light line, broken at the collar, and stopping just posterior to the arm. The
specimens from Bolivia and southern Peru have a much reduced light lateral
line, beginning under the middle or posterior part of the eye, having less
regular margins, and generally broken somewhere anterior to the collar.
There is usually an ill-defined line above the arm insertion.

6. Subdigital lamellae. The generic name Prionodactylus derives from the
tubercles of the subdigital lamellae of northern population (Fig. 1). Such
tubercles are also present in the central Peruvian populations, although they
seem less well developed; this may be partly due to the condition of these
older specimens. The specimens from Bolivia and southern Peru have little
or no swelling of the subdigital lamellae.

The toes of the central and southern Peruvian specimens and of the
Bolivian specimens have many of the proximal subdigital lamellae doubled.
When tubercles are present under the toes as in the central Peruvian speci-
mens, it is the inner part that forms the tubercle.

7. Other characters. Table 4 lists ranges and means for several other char-
acters. Total femoral pore number appears to decrease from north to south.
Ecuadorian and northern Peruvian specimens show almost no sexual dimor-
phism in pore number; the dimorphism increases markedly to the south,
where females have far fewer pores than males. The number of dorsal scales
between occiput and posterior margin of hind limbs decreases from north
to south, as does the number of scales around the midbody region. Sub-
digital lamellae, in contrast, increase in number from north to south.

Because the samples of animals come from separated areas, it is im-
possible to be certain that they are connected by series of potentially inter-
breeding populations. The general pattern of variation from north to south,
without reversals, and the fact that the central Peruvian populations re-
semble the southern populations in some characters (collar scales, longi-

28 POSTIELARISS.

tudinal rows of ventral scales), the northern populations in some others
(preanal scales, disc in lower eyelid, coloration), and are intermediate in
yet others (femoral pores, scales around midbody region, subdigital lamellae)
suggest that these populations are all conspecific. Certainly, the populations
here referred to P. manicatus are more closely related to each other than
any of them is to any other species of Prionodactylus. This is attested by the
single frontonasal, the contact between the loreal and the supralabials, the
presence of a light lip line, and the generally high number of dorsal scales
and scale rows around the midbody region.

Prionodactylus manicatus manicatus (O’Shaughnessy )
(Fig. 10)

Cercosaura (Prionodactylus) manicata O’Shaughnessy, Proc. Zool. Soc. Lon-
don 1881: 231.

SYNTYPES. BMNH 80.12.8.8, reregistered as 1946.8.2.1, an adult male from
Ecuador, Chimborazo, Pallatanga, received from Buckley, and BMNH
80.12.8.5-7, reregistered as 1946.8.31.15-17, three adult females from
Ecuador, Pastaza, Canelos, also received from Buckley.

DEFINITION. A subspecies of P. manicatus distinguished by having two or
three, rather than four, large posterior preanal scales, two widened collar
scales at midline, the transparent disc in the lower eyelid divided into two or
three sections by vertical grooves, a light lip line beginning on the first

FIG. 10. Side view of the head of Prionodactylus manicatus manicatus (UKMNH
109819). 5. The loreal is in contact both with the frontonasal and the supra-

pe The light lip line is broader and straighter than in P. vertebralis and P.
aicrus.

PRIONODACTYLUS AND EUSPONDYLUS 29

supralabial, and subdigital lamellae of toes and fingers forming a serrated
series of tubercles (Fig. 1).

SEXUAL DIMORPHISM. This subspecies of P. manicatus shows remarkably
little sexual dimorphism, although females from central Peru appear to have
fewer femoral pores than the single male examined.

SIZE AND TAIL LENGTH. The largest male examined was 61 mm snout to vent;
the largest female, 73 mm. Two males with tails intact have tail over snout-
vent length ratios of 1.7 and 1.9, mean 1.84; four females with tails intact
have tail over snout-vent length ratios of 1.4 to 1.7, means 1.55.

REMARKS, Although syntypes of P. manicatus manicatus, which IJ have ex-
amined, supposedly came from both the Pacific and the Amazonian slopes of
Ecuador, the specimen from Pallatanga is undoubtedly mislabelled, as was
much Buckley material reported from both Pallatanga and eastern Euca-
dorian localities (Peters 1955: Uzzell 1961).

RANGE. Amazonian slopes of the Andes from central Peru north to Ecua-
dor (Fig. 9), at elevations of 300 to perhaps 1600 m above sea level.

Prionodactylus manicatus bolivianus Werner
(Fig. 11)

Prionodactylus bolivianus Werner, 1899, Zool. Anz. 22: 481.
Prionodactylus Okendeni Boulenger, 1907, Ann. Mag. Nat. Hist. ser. 7,
19: 486.

HOLOTYPE. MNHN 00.4, an adult female from the Chaco of Bolivia; the
specimen was received from Werner (Guibé 1954).

DEFINITION. A subspecies of P. manicatus distinguished by having 4 large
posterior preanal scales, 3 subequal collar scales at midline, the transparent
disc in the lower eyelid not divided into sections by vertical grooves, light lip
line beginning under eye, and subdigital lamellae of toes and fingers except at
bases of toes 3 and 4 not forming a serrate series of tubercles.

SEXUAL DIMORPHISM. Females have far fewer femoral pores than males, but
little other sexual dimorphism is evident. The males I have examined have
2 or 3 black ocelli with light centers, but these are ill defined. Boulenger
(1907) reported ocelli in male syntypes of P. okendeni.

SIZE AND TAIL LENGTH. The largest male examined was 56 mm snout to vent;
the largest female, 58 mm. Two females with intact tails have tail over snout-
vent length ratios of 1.7 to 2.1, mean 1.91.

REMARKS. I have examined the holotype of P. bolivianus. The description
offered by Werner is adequate. I count 4 femoral pores on the right side,

30 POSTILLA 159

FIG. 11. Side view of the head of Prionodactylus manicatus bolivianus (FMNH
40422). 6. The loreal is in contact with the supralabials, the small, oval disc
in the lower eyelid is not divided, and the light lip line is absent beneath and in
front of the eye.

3 on the left; these are weakly developed. There are 3 supraoculars on each
side; the first superciliary is expanded onto the dorsal side of the head. The
loreal is in contact with both the frontonasal and the supralabials. The small
oval translucent disc in the lower eyelid is undivided. There is an even num-
ber of collar scales at midline, and they are not particularly widened. There
is a light middorsal area 2-3 scales wide bordered by dark stripes that
end on the tail. Virtually all of these features can be found in the speci-
mens referred to P. M. bolivianus.

I have also examined a second specimen (ZSM 66/1920) identified by
Werner as P. bolivianus. It is an adult male, purchased from Werner in 1920,
and collected by Schluter in Bolivia in 1907. This individual has 8 longi-
tudinal rows of ventral scales, three large posterior preanals, 3-3 supraocu-
lars, a small undivided oval disc in the lower eyelid, the first superciliary ex-
panded onto the dorsal surface of the head, the loreal in contact with both
the frontonasal and the supralabials, and 3 subequal collar scales at midline.

The only definite statements about range in Bolivia are Werner’s locality
in Chaco, and a specimen (UMMZ 69547) from the Departamento de
Cochabamba. I suspect that the Chaco is too dry for this species, but the
fauna of Bolivia is too ill known for this to be more than conjecture.

Many specimens of Opipeuter xestus in collections from Bolivia have
been labelled Prionodactylus bolivianus; they differ from the holotype of
P. bolivianus in being totally devoid of keeling on the scales, rather than

PRIONODACTYLUS AND EUSPONDYLUS 31

strongly keeled especially on the dorsal and lateral body and caudal scales.
Werner did not mention keeling in his description of P. bolivianus but such
keeling is implied by his generic placement of the taxon. The undivided disc
in the lower eyelid of P. m. bolivianus is small, oval, and translucent, whereas
in O. xestus it is relatively enormous, nearly circular, and transparent
(Uzzell 1969).

The syntypes of Prionodactylus okendeni Boulenger (1907) are BMNH
1905.5.31.2-8, reregistered as 1946.8.31.36-42, from Puno, Oconeque, at
about 1750 m above sea level, and 1902.11.28.3, from Peru, Puno, Santo
Domingo, about 1500 m above sea level. I have examined these specimens,
but only data on 1946.8.31.37 are included in the discussion of variation and
in Table 4. The description by Boulenger places the syntypes within the
variation that I have observed in southern Peruvian specimens. I know of
no characters that will distinguish the syntypes of Prionodactylus okendeni
from the Bolivian material identified by Werner as P. bolivianus. I therefore
consider P. okendeni a junior synonym of P. bolivianus Werner (cf. Peters
and Donoso-Barros 1970).

AMNH 1705-06, recorded as from Lake Aracona, Juliaca, Peru, probably
came from near Santo Domingo in Puno, Peru (cf. Dunn 1942, Uzzell 1970).

RANGE. Specimens of P. m. bolivianus are known from the Department de
Puno in southern Peru south to the Departamento de Cochabamba or pos-
sibly the Chaco of Bolivia, at elevations of about 1200-17500 m above sea
level (Fig. 9).

Specimens Examined
Prionodactylus m. manicatus

ECUADOR: Napo: Rio Licuna USNM 194066; Rio Pindo USNM 194068;
Puerto Libre, Rio Aguarico (570 m) UKMNH 119405-7; Puerto Napo
(463 m) UIMNH 55781; Santa Cecilia (340 m) UKMNH 109817-18,
119408-09; Rio Viclano USNM 194067. Pastaza, Rio Pastaza between
Canelos and Rio Marafién MCZ 37261. MoRoNa-SANTIAGO, between Rio
Pastaza and Rio Santiago MCZ 45779.

PERU: Amazonas: Rio Cenipa Valley AMNH 56494. Cuzco, Rio Apuri-
mac, Luisiana (500 m) AMNH 90673. HuaNuco, Divisoria (1300-1600 m)
FMNH 55986-88. Pasco, Oxapampa (1800 m) FMNH 134398.

Prionodactylus m. bolivianus
BOLIVIA: no other locality: ZSM 66/1920. CHaco: MNHN 00.4, holotype

of Prionodactylus bolivianus; DEPARTAMENTO COCHABAMBA: UMMZ 69547.
PERU, PuNo (Carabaya) Santo Domingo (1862 m) FMNH 40420-25;

32 POSTILLA 159

(Sandia) Oconeque (1956 m) BMNH 1905.5.31.3, reregistered as 1946.
8.31.37, syntype of Prionodactylus okendeni; Juliaca, Lake Aracona (=Lago
de Aricoma) AMNH 1705-06.

Prionodactylus eigenmanni Griffin
Prionodactylus eigenmanni Griffin, 1917, Ann. Carnegie Museum 11: 316.

HOLOTYPE. An adult female (CM 918) collected by José Steinbach in the
Provincia de Sara, Beni, Bolivia, 400 m above sea level.

DEFINITION. A single frontonasal. Loreal in contact with frontonasal and
supralabials. Two large collar scales. Subdigital lamellae of three kinds
(Fig. 12): simple, with a median tubercle, or with two lateroventral tubercles.

FiG. 12, Undersurface of the right forefoot of Prionodactylus eigenmanni (BMNH

ia ie showing the single and double tubercles of the subdigital lamellae.
x :

PRIONODACTYLUS AND EUSPONDYLUS 33

Translucent disc in lower eyelid oval, not divided into segments by vertical
grooves. Marked sexual dimorphism in femoral pore number. No con-
spicuous dorsal light lines; no continuous light line along supralabials.

VARIATION. I have examined the holotype and six other individuals. There
is very little variation (Table 5). All specimens have a median occipital, six
transverse rows of ventral scales, 3-3 supraoculars, the first superciliary ex-
panded onto the dorsal surface of the head, an undivided translucent disc
in the lower eyelid, and two very wide collar shields at midline.

In addition to the sexual dimorphism in femoral pore number, the single
female examined has four rather than two posterior preanals. Griffin inter-
changed anterior and posterior preanals in his description.

The largest male examined is 38 mm snout-to-vent; the largest female, 40.
Three males with intact tails have tail over snout-vent length ratios of 1.2
to 1.4, mean 1.30.

REMARKS. Burt and Burt (1931) placed this species in the synonymy of
P. bolivianus Werner. I have examined the holotype of P. bolivianus;
P. eigenmanni is quite distinct.

RANGE. Amazonian slopes of Bolivia between 200 and 400 m above sea
level (Fig. 13).

Specimens Examined

BOLIVIA: BENI, Yacuma, Rurrenabaque (227 m): AMNH 22537-38;
Provincia de Sara (200-400 m): CM 981, holotype. SANTA Cruz, Ichilo,
Buena Vista (400 m): MCZ 24887, BMNH 1927.8.1.151-152.

Prionodactylus argulus (Peters)

Cercosaura (Pantodactylus) argulus Peters, 1862, Abh. Akad. Wiss. Berlin
1862: 184.

Prionodactylus oshaughnessyi Boulenger, 1885, Cat. Liz. British Mus. 2: 392.
Prionodactylus holmgreni Andersson, 1914, Ark. for Zool. 9 (3): 9.
Prionodactylus columbiensis Werner, 1916, Zool. Anz. 47: 306.

TABLE 5. Characters of seven specimens of Prionodactylus eigenmanni.
Figures represent ranges and, in parentheses, means.

Scales
Total Dorsal Ventral around Subdigital lamellae
femoral scale scale midbody Fourth Fourth
pores rows TOWS region finger toe
626 12-15 32-35 17-19 29-31 9-11 13-15

(1320) (33.0) (17.8) (2958) (10:0) (14:2)
ee 0 32 18 30 10-10 13-14

34 POSTILLA 159

FIG. 13. Bolivia, showing localities for Prionodactylus eigenmanni.

HOLOTYPE. ZMB 4555, an adult male from the mountainous regions around
Santa Fe de Bogota, Cundinamarca, Colombia collected by Liedig.

DEFINITION. Frontonasal divided. Loreal in contact with frontonasal and
supralabials. Two or three large collar scales at midline. Subdigital lamellae
mostly divided but not tuberculate except at bases of toes 1-4. Translucent
disc in lower eyelid divided into two pieces by a vertical groove. Marked
sexual dimorphism in femoral pore number. Usually a pair of dorsolateral
light line; supralabials light, the light area continued posteriorly as a lateral
light line through external ear opening and above forelimb.

VARIATION. P. argulus is a wide-ranging species, but I have seen relatively
few specimens from the northeastern and southwestern parts of its range
(Fig. 14). For the characters that I have investigated, there seems to be
little geographic variation (Table 6). Virtually all specimens have a median
occipital, although it is irregular in several specimens. In UMMZ 68087, it
is fused to one of the paramedian occipitals. The loreal is in contact with

PRIONODACTYLUS AND EUSPONDYLUS 35

: ; =e

e P arguilus

8°

Santa Rosa de Osos pe

; \

SE2A40 8

a a \ i Cees ae
eto 4 ~ “~y KILOMETERS

Riobamba

Pallatanga \é
ht
{

Wy
Ke
eh

ZZ

Lp,

io

8°

16°)

FIG. 14. Northern South America showing savannah and grassland formations, and
localities for Prionodactylus argulus. Open symbols are literature records. Certain
localities discussed in the text are indicated. Abbreviations: DG, broadleaf decidu-
ous trees and grass; Di, broadleaf deciduous trees, plants sufficiently far apart
so that they frequently do not touch; DsG, broadleaf deciduous shrub forms,
minimum height 3 feet, with grass; GBp, grass with broadleaf evergreen trees
growing singly or in groups or patches; GSp, grass with semideciduous trees
growing singly or in groups or patches.

the supralabials in all specimens. The supraoculars mostly number 3-3;
occasional specimens have 4 or 2 on one side, 3 on the other. The first
superciliary is expanded onto the dorsal surface of the head in all speci-
mens, but the area on the dorsal surface varies considerably. The translucent
disc in the lower eyelid is divided into two parts by vertical grooves. Some
14 specimens from various localities lack a median collar scale and thus
have an even rather than an odd number of collar scales.

Most of the specimens examined have 6 longitudinal rows of ventral
scales. In the small sample from Bolivia, including the holotype of P. holm-
greni, all specimens have 8 longitudinal rows of ventral scales. These far-
southern specimens also possibly have a slightly larger number of dorsal and
ventral scale rows, and a lower number of scales around the midbody region.
The samples are, however, very small, and similar counts also occur in the
far northeast. Because of the general absence of distinction in scale counts,
I see no present merit in recognizing a southern race of P. argulus.

The samples from Colombia seem distinctive in some counts, particularly
in their lower numbers of dorsal scale rows. The interpretation of these data

26 POSTILLA 159

TABLE 6. Characters of 73 specimens of Prionodactylus argulus.
Figures represent ranges and, in parentheses, means.

Scales
Total Dorsal Ventral around Subdigital lamellae
femoral scale scale midbody Fourth Fourth
pores TOWS rows region finger toe
Brazil
ne 20 39 19 34 16-17 21-22
L-9 0 42 20 37 17-18 22
Guyana
eG: 16 38 20 35 15-16 20-21
2.29 4-5 37-40 (18-20 32-33 15-16 19-21
(4.5) (38.5) (19.0) (32:5) (Gey 7) (20.0)
Colombia
Meta
466 14-16 31-36 18-20 29-33 14-16 18-21
15:5) (33.5) 2" (19:0); (Gied) (GISe3))) (19:5)
Bag? 7-11 33-35 18-19 30-32 15-16 19-22
(9:0). (340) (18-3) (31.0) (152) (20.7)
Putumayo
1 3 14 32 18 30 16 18-19
Miscellaneous
4é6¢ 15-16 31-34 17-20 29-30 14-16 18-20
GIS.3) 8" 432.7) (LSE) 197) (15.0) dor)
ies i 33 20 30 13 16
Ecuador!
14 $¢ 12-23 38-42 16-21 30-39 13-18 18-22
(18:8) (39.6) Csi6y) (3643) (GS) (19.4)
13 Oe2 4-13 38-43 17-21 32-39 14-18 17-24
ClOn)~ (4) (18.0) (35:5) (15.9) (1929)
Peru
Loreto and San Martin
gf Steines 13-25 37-44 18-23 31-41 13-18 18-23
C17-0)) (39:9) (19:9) G36) (153) (19.9)
70 9 4-15 36-45 19-20 31-41 14-17 18-21
(9.8) (41.2) (19.2). (36:8) C1527) (19.6)
Huanuco
ZO6 16-20 37-43 19-21 36-37 16-18 20-22
12 ‘ 43 19 36 16-17 20-22
Madre de Dios
er: 12 38 22 32 14-15 18-20
Bolivia
DOG. = 13-15 42 22 32, 16 20-22
PAR) 2 41-44 Pn Sil 15-18 18-21

1Based on AMNH 32724, 60630, 89831; BMNH 80.12.8.17, 1946.-8.31.18,
1956.1.15.91; MLS 372 (Colombia) UIMNH 65701-2, 04-15; UKMNH 98604-05;
UMMZ 84745, 84846 (1), 90771, USNM 193949-50.

“Holotype of P. holmgreni italicized.

PRIONODACTYLUS AND EUSPONDYLUS Si7/

is confused by some doubts about the origin of some of the specimens. The
specimens that supposedly came from west of the eastern range of the Andes
seem no more distinct from other populations than those that certainly
come from Amazonian Colombia. Thus, even though the dorsal counts
given by Werner for his P. columbiensis are lower than those Werner gave
for conspecific nominal taxa that have Amazonian ranges, the data do not
permit recognition of a Magdalena Valley race, both because the western
specimens are not distinct from the Amazonian specimens from Colombia,
and because the total Colombian sample is only very weakly distinguished
from the remainder of the populations.

SEXUAL DIMORPHISM. The greatest sexual dimorphism is in color. Mature
males have one to several light-centered, dark-bordered ocelli along the sides
of the body beginning in the shoulder region. Such ocelli are absent or at
most faint in the females and young. The number of femoral pores also shows
sexual dimorphism; where appreciable samples have been examined, the
highest total counts for the females overlap the lowest total counts for
males (Table 6).

REMARKS. The holotype of Cercosaura argulus (ZMB 4555) is an adult
male. I have examined the specimen. Peters’ (1862) description is very good.
The holotype is unusual in having the prefrontals separated from each by an
anterior extension of the frontal. I have not seen this condition elsewhere in
the specimens I refer to this taxon, but the holotype is so similar to the
material examined in so many other features that I consider this distinction
an anomaly.

Prionodactylus oshaughnessyi was distinguished from P. argulus by Boul-
enger (1885) primarily, I believe, because of the anomalous separation of
the prefrontals in Peters’ holotype. Boulenger also listed other differences
in scale dimension and number. I have examined the syntypes of P. oshaugh-
nessyi (BMNH 80.12.8.1 from Pallatanga, and 80.12.8.14-16, recatalogued
as 1946.8.31.18-20), a male and two females from Canelos. These speci-
mens fall within my concept of P. argulus. All the syntypes were collected
by Buckley; the specimen labelled Pallatanga, like much other Buckley ma-
terial from Pallatanga, is undoubtedly mislabelled (Peters, 1955; Uzzell,
1961).

Through the courtesy of Greta Vestergren, I have examined the holotype
of Prionodactylus holmgreni, an adult male (NRM 3226) collected by
N. Holmgren at San Fermin, in La Paz (Caupolican), about 850 m above
sea level. I consider P. holmgreni a junior synonym of P. argulus (cf. Peters
and Donoso-Barros, 1970). Although this and the other Bolivian material
examined show slight differences from the rest of the material referred to
P. argulus, 1 do not presently believe that any purpose is served by recog-
nition of a distinct southern subspecies of P. argulus, although the name
holmgreni remains available should that course later prove useful.

38 POSTILLA 159

Burt and Burt (1931) considered P. holmgreni a subspecies of P. oken-
deni. P. okendeni, however, differs in having a single frontonasal, rather
than a divided one, and belongs with P. manicatus. The specimens identified
by the Burts as P. o. holmgreni (AMNH 22740-41, 38955-62) are Opipeuter
xestus.

Prionodactylus columbiensis Werner (1916) was described from a speci-
men purportedly from the Canon de Tolima. I have been unable to locate the
holotype. Werner described the specimen in enough detail so that it clearly
may be placed with the rest of the material that I identify as Prionodactylus
argulus. Especially, the divided frontonasal and the presence of a lateral
light line are characteristic of P. argulus rather than any of the other species
of Prionodactylus.

The number of scales around the midbody region in P. columbiensis, ac-
cording to Werner’s key, varies from 25 to 29; the holotype had 25. I have
not seen specimens with this few scales (Table 6). All of the other data
given by Werner can, however, be matched within the specimens I refer
to P. argulus.

Werner gave no indication of the sex of the holotype. The presence of two
large posterior preanal scales suggests that it was a male; most females have
more than two posterior preanals. Most males, especially individuals as
long as Werner’s holotype (SO mm snout-to-vent) have at least one well-
marked ocellus on the side of the body, but Werner mentioned none.

Prionodactylus argulus is basically an Amazonian drainage species
(Fig. 14). In Colombia, most of the recent records are from the lower
Amazonian slopes. I have examined 9 specimens from these areas. In addi-
tion, however, I have examined the following specimens from west of the
easternmost Andes in Colombia: MLS 373 (Humbo near Muzo, Boyaca),
MCZ 42190 and 61157 (Muzo, Boyacé), MCZ 22014 (Bogota, Cundina-
marca), AMNH 32769 (Santa Rosa de Osos, Antioquia), and UMNZ
131628 (Fusagasuga, Cundinamarca). The holotype of P. argulus reportedly
came from the mountainous surroundings of Bogota. The holotype of
P. columbiensis reportedly came from the Cafién de Tolima.

Dunn (1944) was almost certainly correct in his surmise that P. argulus
is not part of the herpetofauna of the Bogota region. AMNH 32769 had
been identified (Burt and Burt, 1931) as one of a series of P. vertebralis
from Santa Rosa de Osos, and almost surely some error in record keeping
accounts for the locality data of this specimen. Since P. argulus is basically
a lowland Amazonian form, I doubt that it crosses the easternmost Andes
either near Andalucia or at the Paso de los Cruces, both of which are at
about 1800 m. The records are, however, too numerous to reject out of
hand, and I have indicated these western localities on the range map for
P. argulus (Fig. 14). Verification that P. argulus does not occur in the
Magdalena valley is needed. Regardless, there is nothing distinctive about
the specimens from these western localities, which are listed in Table 6
as miscellaneous Colombian localities.

PRIONODACTYLUS AND EUSPONDYLUS 39

AMNH 14561-62, 23328, and 32724, supposedly from Riomamba, Chim-
borazo Province, Ecuador, are probably mislabelled.

RANGE. Specimens of Prionodactylus argulus are known from elevations
from about 100 m above sea level to perhaps 1600 m above sea level at
the periphery of the Amazon basin from Bolivia north and east to Amapa in
Brazil.

Specimens Examined

BOLIVIA. BENI (Ichillo), Buena Vista (400 m) UMMZ 60600. CocHa-
BAMBA: UMMZ 68087-88 (Cundinamarca), Monteredondo (1420 m)
ZSM 63/1959 (2 specimens). LA Paz (Caupolican), San Fermin (700-
100 m) NMNH 3226, holotype of Prionodactylus holmgreni.

BRAZIL. AmaPA, upper Rio Lunier, Tumuc-Humac (Tumucumaque; 100-
200 m) MNHN 1899.73. PardA, Utinga, near Belém (100-200 m) ZSM
203/1911.

COLOMBIA. Amazonas, Puerto Narifio (280 m) MLS 372. ANTIOQUIA,
Santa Rosa de Osos: AMNH 32769. Boyaca, Garagoa (1634 m) MCZ
31860; Humbo, near Muzo: MLS 373; Muzo: MCZ 42190, 61157. ?CuN-
DINAMARCA, Bogota: ZMB 4555, holotype of Cercosaura argulus, MCZ
22014; Fusagasuga (1800 m) UMMZ 131680; Meta, Acacias MLS 375;
Cano Guayapa, tributary of Rio Guejar (400 m) FMNH 81316, UMMZ,
131682; upper Rio Guejar near San Juan de Arama (400 m) UMMZ
131679; Villavicencio (450 m) MCZ 31606-07, 31859; MLS 376. PuTUu-
MAYO, Puerto Asis MLS 374. VAUPES, upper Rio Apaporis, tributary of Rio
Caqueta: UMMZ 131681.

ECUADOR. No other locality: AMNH 38954. Eastern Ecuador: AMNH
89831. CHIMBORAZO, Pallatanga: BMNH 80.12.8.17, syntype of Priono-
dactylus oshaughnessyi; Riobamba (2250 m) AMNH 14561-62, 23328,
32724. MorONA-SANTIAGO, between Rio Pastaza and Rio Santiago: MCZ
45780. Napo, Cuyabeno (227 m): UIMNH 65707; Limén Cocha (200-
500 m) UIMNH 65701-02, 65704-05, 65708-13, 65715, UKMNH 98604-
05, 119429; 2.5 km S of Ongota: UMMZ 123893; Puerto Libre, Rio Agua-
rico (570 m) UKMNH 119425-28; San Francisco (1200 m) UMMZ 84746
(4 specimens), 84747 (7), UIMNH 65714; Santa Cecilia (340 m) UKMNH
105384-85, 109820-22, 112227, 119410-24; near Tena (SOO m) UMMZ
84745. Pastaza, Alpayacu UMMZ 90771; Anga Cocha, Rio Bobonaza
AMNH 60630; 2.5 km downstream from Cabaceras, Rio Bobonaza (550 m)
USNM_ 193949; Canelos (700 m) BMNH 80.12.8.14, reregistered as
1946.8.31.18, syntype of Prionodactylus oshaughnessyi; 6 km W of Cane-
los (700 m) UKMNH 119432; Rio Conambo (300 m) USNM 193953;

40 POSTIELAVIS2.

Mera (1140 m) UKMNH 119430-31; upper Rio Pastaza (800 m) USNM
193952; 2.5 km SE of Puyo (800 m) USNM 193948; Sarayacu (400 m)
MCZ 37710; BMNH 1956.1.15.91; Veracruz, 10 km E of Puyo (900 m)
USNM 193950-51, UKMNH 119433.

GUYANA. Essiquibo, Bartica District (10 m) AMNH 21265; Marudi
(250 m) AMNH 61386; Shudikar-wau (300 m) AMNH 61434.

PERU. No other locality: AMNH 56293. AMaAzonas, mouth of Rio Santiago
(100-200 m) AMNH 56285, 56291; HuANuco, Divisoria (1300-1600 m)
FMNH 55983-85; Hacienda Pampayacu (750 m) MCZ 43767. LoreETo,
Cerros de Contaya, E of Contamana (100-200 m) AMNH 56297; Cerro
Azul (100-200 m) FMNH 55982; Iquitos (100 m) AMNH 56299; Isla
Cedro (100-200 m) AMNH 56292; mouth of Rio Aquaytia (100-200 m)
AMNH 56286; Rio Bombo, upper Rio Tapiche Valley (100-200 m) AMNH
56221; mouth of Rio Napo, Lago Mirano region (100-200 m) AMNH
56295-96; mouth of Rio Tigre (100-200 m) AMNH 56493; upper Rio
Utoquinia (200-500 m) AMNH 56280; Pucallpa (100-200 m) MCZ 45878;
San Regis (100-200 m) AMNH 56294; Utoquinia-Tapiche region: AMNH
$6281-83; Yarinacocha (100-200 m) FMNH 55981. MapbRrE DE DyIos,
Avispa (1000 m) FMNH_ 154832. SAN MarTIN, Juan del Monte,
AMNH 56394.

Hemipenes of Species of Prionodactylus

I have examined hemipenes of all five species of Prionodactylus. Members
of this genus, like most species of Boulenger’s (1885) Group II of the
family Teiidae, have calcareous spinules in the flounces of the hemipenis. I
have developed a technique for examining these (Figs. 14-21). The hemi-
penis is dissected free from the tail; for consistency, I have used the left organ
if possible. The organ is washed in several changes of distilled water, soaked
briefly (several hours) in 0.5% KOH, stained in a solution of Alizarin Red
S in 1% KOH for 12 to 24 hours, destained in several rinses of distilled
water (by the time unbound dye is removed, excess KOH is also removed)
and stored in 70% alcohol. Staining is more uniform and more rapid if the
organ is slit before being stained. For uniformity, I have slit the organ along
the length of the sulcus spermaticus from its attachment at the cloacal wall
to the end of the undivided section of the hemipenis. Additional slitting
beyond this point, both medially to separate the two lobes, and laterally, to
open up each lobe, allows the internal pattern to be examined. The organ
loses little of its integrity in the KOH solution, and may be manipulated with
forceps so that the details, especially those hidden beneath the median welt,
may be examined.

The feature most objectively viewed is the pattern of the calcareous
spinules. At the tip of the organ, the two lobes form a series of complex folds,

PRIONODACTYLUS AND EUSPONDYLUS 41

but these are difficult to examine, and easily destroyed or damaged while
slitting the lobes. I have placed little emphasis on variation in this part of
the organ.

Because the illustrations are based on slit and spread preparations, it
should be pointed out that in the intact organ, the spinous areas on either
side of the median welt form two distinct pouches. Before dissection, the
sulcus is in contact with the median welt, and in cross section, the undivided
part of the organ has a C-shaped space, with the median welt inside the C.

There are numerous slight differences in the hemipenes examined. Since
I have examined only one from each species, it is not possible at this time
to be certain that the differences do not reflect individual or seasonal dif-
ferences rather than species-specific differences.

The hemipenis of P. vertebralis (Fig. 15A) has a series of chevron-shaped
flounces with calcareous spinules in both the lateral (left) and medial (right)
pockets. The apices of the chevrons are basal and lie beneath the free
edge of the median welt when the organ is slit along the sulcus spermaticus.
There are 22 complete chevrons in the lateral pocket, plus two incomplete
ones at the base of the organ; there are 21 complete and one incomplete
chevron in the medial pocket. Teeth are present across the apices of the
chevrons, but in the distal 4 or 5 chevrons, the apex is occupied by an en-
larged tooth, usually with lateral spurs (Fig. 15B, C). These spurs have
not been seen in other species of Group II. The spinules in the flounces are
relatively uniform in size (Fig. 15D). The base of the median welt has a
series of five tooth-bearing flounces that continue across it (Fig. 15E). The
teeth in these vary considerably in size, being smallest in the basal rows,
and at the lateral edges of each row; the teeth in the distalmost of these five
rows are slightly smaller than the teeth in the row basal to it.

The hemipenis of P. dicrus (Fig. 16A) is generally similar to that of
P. vertebralis. There are 14 toothed flounces in the lateral pocket, and 15
in the medial pocket. The chevron-shaped flounces lack teeth at the apices
(Fig. 16B); there are five short rows of teeth across the base of the median
welt (Fig. 16C); the teeth are larger medially, smaller at the ends of the
rows. The teeth of the third and fourth rows from the base are larger than
those of the remaining rows.

In the hemipenis of Prionodactylus m. manicatus (Fig. 17A), the toothed
flounces form chevron-shaped structures; teeth are continuous across the
apex. There is, however, no enlarged tooth at the apex of the distal chevrons.
The chevrons number 23 in the lateral pocket, 24 in the medial. There are
three complete and one incomplete rows of teeth across the base of the
median welt (Fig. 17B). The teeth are larger medially, and larger in the
distal rows.

In the hemipenis of Prionodactylus argulus (Fig. 18A) the flounces do
not form complete chevrons; this is conspicuously true of the basal flounces
(Fig. 18B). These toothed flounces number 18 in the lateral pocket, 17 in the
medial pocket. A single complete row of teeth crosses the base of the median

POSTILEAVWISS

PRIONODACTYLUS AND EUSPONDYLUS 43

A

"Dy

ee ih, Wabi, a

yy

FIG. 16 (above). Structure of the left hemispenis of Prionodactylus dicrus, new
species (FMNH 28043). The inverted organ has been slit along the sulcus sperma-
ticus. A) The entire organ showing general arrangement of the flounces in the
lateral (left) and medial (right) pockets. x 10. B) The median welt pulled back to
reveal the apices of the chevron-shaped flounces; the teeth are not continuous
across the apex, and there are no enlarged teeth at the apex. x22. C) The short
rows of teeth in flounces that cross the base of the median welt. x 33.

FIG. 15 (left). Structure of the left hemipenis of Prionodactylus vertebralis (USNM
_ 193939). The inverted organ has been slit along the sulcus spermaticus. A) The
_ entire organ showing general arrangement of the flounces in the lateral (left) and
_ medial (right) pockets. x7. B) The median welt pulled back to reveal the apices
of the chevron-shaped flounces. «16. C) Details of the enlarged teeth at the
apices of the distal chevrons of the medial pocket. x43. D) Details of the teeth
_ in the arms of the chevrons. x 43. E) The short rows of teeth in flounces crossing
_ the base of the median welt. x 16.

\

44 POSTILLA 159

welt (Fig. 18C), but there is also a partial row, and there are one or two
isolated teeth.

In Prionodactylus eigenmanni, the hemipenis has 14 toothed flounces in
the lateral pocket, 15 in the medial pocket. These flounces are chevron
shaped, with teeth complete across the apices. There is no enlarged tooth
at the apex of the chevron. There are 4 short rows across the base of the
median welt, and part of a fifth. The teeth in these are largest medially,
smallest at the ends of the rows. The teeth of the middle rows are larger.

In general, the hemipenial structures in Prionodactylus thus seem to be
rather similar. There are differences in the few organs examined. Some of
these, such as number of rows of flounces, are associated with body size;
there are more flounces with teeth in P. vertebralis and P. m. manicatus, the
two largest taxa. The enlarged teeth at the apices of the chevrons of P.
vertebralis are the most distinctive feature observed in the series.

FIG. 17. Structure of the hemipenis of Prionodactylus manicatus manicatus (FMNH
45779). The inverted organ has been slit along the sulcus spermaticus. A) The
entire organ showing general arrangement of the flounces in the lateral (left)
and medial (right) pockets. x6. B) The short rows of teeth in flounces that
cross the base of the median welt. x 17.

PRIONODACTYLUS AND EUSPONDYLUS 45

FIG. 18. Structure of the left hemipenis of Prionodactylus argulus (FMNH 81316).
The inverted organ has been slit along the sulcus spermaticus. A) The entire
organ showing general arrangement of the flounces in the lateral (left) and
medial (right) pockets. 6.5. B) Detail of the flounces in the basal part of the
medial pocket, showing the absence of teeth at the apices of the chevrons. x 34.
C) The short row of teeth in the flounces crossing the base of the median welt.
x 34.

46 POSTILEA 159

Figures 20-22 show hemipenial structures of individuals of three related
genera. The hemipenis of Pantodactylus schreibersii is very distinctive in
comparison to the hemipenes of species of Prionodactylus. There are 15
tooth-bearing flounces in both the lateral and medial pockets (Fig. 20A).
Teeth are absent across the apices of the chevrons, especially basally; in this,
there is some similarity to the hemipenis of P. argulus (Fig. 18B). In marked
distinction, however, the teeth of the basal five flounces of, the left pocket
are much larger than those of the distal flounces (Fig. 20B). The teeth in
the short rows crossing the base of the medial welt also seem to be relatively
much larger than the corresponding teeth in species of Prionodactylus
(Fig. 20C).

The hemipenis of Cercosaura ocellata (Fig. 21) also shows some dis-
tinctive features. There are 12 chevron-shaped tooth bearing flounces in the
lateral pocket, and 17 in the medial pocket. Three rows of teeth that cross
the base of the median welt are continuous with 3 of the more basal chevrons
in each pocket (Fig. 21B). This condition has not been seen in Prionodactylus
or the other genera discussed here.

The hemipenis of Aspidolaemus affinis (Fig. 22) seems rather like that of
several forms of Prionodactylus, especially P. vertebralis and P. m. manicatus.
There are 27 tooth-bearing flounces in the lateral and 28 in the medial
pocket. Most of these form chevrons that are toothed across the apex; there
is no enlarged tooth at the apex. There are 4 short rows of teeth across the
base of the median welt. In addition, the ends of two of the incomplete basal
chevrons are essentially joined on the median welt by an enlarged tooth
(Fig. 22B). The general similarity of the hemipenis of P. vertebralis, P. m.
manicatus, and Aspidolaemus affinis can be observed in comparing Figures
15; tang 22.

Relationships within the Genus Prionodactylus

Within Prionodactylus, the closest pair of species clearly seems to be P.
vertebralis and P. dicrus. These share nontuberculate subdigital lamellae on
the fingers, an undivided frontonasal, a loreal separated from the supralabials,
minutely ridged dorsal scales, a distinctive light lip line, and parts of the
dorsal pattern: the posterior middorsal light line of P. dicrus resembles that
of P. vertebralis. The anterior pair of dorsolateral stripes on the head and
anterior part of the body of P. dicrus is paralleled in some P. vertebralis by
light lines on the superciliary scales. This pair seems clearly to represent two
daughter species derived from a single ancestral species.

P. manicatus seems in many ways related to P. vertebralis and P. dicrus.
after formaldehyde preservation, some degree of minute ridging on the dorsal
scales, the nontuberculate subdigital lamellae on the forelimbs of P. m.
bolivianus, and the light lip line, especially of P. m. manicatus. P. manicatus
has a much higher number of scales around the midbody region, and in
P. manicatus, the loreal is in contact with the supralabials.

PRIONODACTYLUS AND EUSPONDYLUS 47

A

FIG. 19. Structure of the hemipenis of Prionodactylus eigenmanni (BMNH 1927.
8.1.152). The inverted organ has been slit along the sulcus spermaticus. A) The
entire organ showing the general arrangement of the flounces in the lateral (left)
and medial (right) pockets. x12. B) The median welt pulled back to show the
teeth continuing across the apices of the flounces in the lateral pocket, and details
of the spinules. x 27.

Prionodactylus argulus and P. eigenmanni seem rather alike, but this
general impression results from their both being brownish rather than
blackish when preserved; the venter in both is light. This perhaps reflects
altitude, and possibly is related to insolation, rather than closeness of com-
mon ancestry. Indeed, P. manicatus in general occurs at altitudes ranging
between those of P. argulus and P. eigenmanni on the one hand and those of
P. vertebralis and P. dicrus on the other, and is considerably less uniformly
black beneath than the two higher elevation forms, although blacker than
the lower elevation forms.

P. argulus seems to be closer to P. vertebralis, P. dicrus, and particularly

48 POSTIELAMIS9.

P. manicatus in that it has a light lip; this forms a line only posteriorly where
it is bordered beneath by a dark line. P. argulus has the loreal in contact
with the supralabials, but is particularly distinctive in its longer snout and
divided frontonasal.

Prionodactylus eigenmanni also has the loreal in contact with the supra-
labials, but it has an undivided frontonasal. The single and double tubercles
on the subdigital lamellae set this species apart from the others in the genus.

The arrangement of the loreal and of the frontonasal both appear to be
related to snout length. P. dicrus and P. vertebralis, in which the loreal is
separated from the supralabials, have, relative to body size, the shortest
snouts in the genus. The three species in which the loreal touches the
supralabial series have relatively longer snouts. The longest snouted species,
P. argulus, also has the frontonasal divided. Divided frontonasals are unusual
in Group II of the Teiidae, and do not occur in all long-snouted forms, for
example, Anadia and Placosoma. A divided frontonasal does occur, however,
in the longer-snouted forms of Echinosaura.

Relationships to Other Genera

Ruibal (1952: 520) pointed out that Pantodactylus and Prionodactyius are
closely related, in fact, perhaps not distinguishable. Among the species of
Prionodactylus, P. argulus seems closest to Pantodactylus. They share,
among other features, the general brownish color in preservation, a lined
pattern, at least in some forms of Pantodactylus; and a light venter. More
importantly, both Pantodactylus schreibersii and Prionodactylus argulus have
all but the distal subdigital lamellae of the toes and fingers divided into two
parts, although not tuberculate. There is possibly some confirmation of a
relationship in the structure of the hemipenis, in the absence of teeth at the
apices of the basal chevrons (Figs. 18B, 20A). The enlarged teeth in the
basal chevrons of Pantodactylus is, however, quite distinctive.

Cercosaura also seems to be closer to P. argulus or P. eigenmanni than to
the other species of Prionodactylus, but is a very distinctive animal, and
apparently not particularly close to any Prionodactylus.

Aspidolaemus affinis appears in its general features to be closer to the
high-elevation species of Prionodactylus, P. vertebralis and P. dicrus, than
to other members of the genus. This similarity, however, in such features as
the generally dark coloration, and slightly pigmented disc in the lower eyelid,
may well reflect adaptations to high elevations. Some suggestion of affinity
occurs in the hemipenial structure, in the numerous flounces with teeth in
them and the larger number of short, tooth-bearing folds across the base of
the median welt. There is some similarity to P. manicatus as well: the loreal
in A. affinis touches the supralabials, and there are dorsolateral light lines,
ill marked, in both.

These notes are very tentative. A study of skeletal and muscular features
would shed much light on the affinities of this complex group of lizards.

PRIONODACTYLUS AND EUSPONDYLUS 49

C

FIG. 20. Structure of the left hemipenis of Pantodactylus schreibersii (AMNH
101604). The inverted organ has been slit along the sulcus spermaticus. A) The
entire organ showing the general arrangement of the flounces in the lateral (left)
and median (right) pockets. The chevron-shaped flounces, especially the basal
ones, lack teeth at the apices. «9. B) Detail of the enlarged teeth on the lateral
arms of the chevrons of the lateral pockets. x44. C) The large teeth in the
short rows in the flounces that cross the base of the median welt. « 44.

50 POSTILLA 159

II]. SPECIES REMOVED FROM THE GENUS Prionodactylus

Four species described as Prionodactylus do not belong in the genus as I
define it. Three of these can appropriately be transferred to other genera.

Prionodactylus spinalis Boulenger
Prionodactylus spinalis Boulenger, 1911, Ann. Mag. Nat. Hist. ser. 8, 7: 23.

This species was described (Boulenger 1911) from BMNH 1911.12.13.33-
46, reregistered as 1946.8.31.44-57. The type locality, Huancabamba, above
750 m, is certainly the central Peruvian town of that name in Pasco rather
than the northern town in Piura. Although Boulenger indicated that the
specimens may have come from Oxapampa (Barbour and Noble, 1921), I
see no reason to doubt the original information; at most it is a matter of a
few kilometers distance and a few meters in elevation.

I have examined the syntypes of P. spinalis. The absence of a double
row of widened gular scales indicates that this species is not a Prionodactylus.
P. spinalis shares most of the features by which I discriminate genera of
teiid lizards in Group II with the type of the genus Euspondylus, E. macu-
latus. For this reason it seems appropriate to recognize this taxon as
Euspondylus spinalis (cf. Peters and Donoso-Barros, 1970).

Prionodactylus rahmi Grijs
Prionodactylus rahmi Grijs, 1936, Zool. Anz. 116: 27.

This species was described from two specimens, a male and a female,
numbered 5221 in the museum in Hamburg. The two syntypes appear to have
been lost during World War II. The locality given by Grijs is Cuzco, Peru,
3000 m.

The quadrilateral rather than hexagonal dorsal scales and the absence of
two median rows of conspicuously widened gular scales indicate that rahmi
is not a member of the genus Prionodactylus as I define that genus. Presently,
it seems best to recognize this taxon as Euspondylus rahmi (cf. Peters and
Donoso-Barros, 1970). It is probably close to Euspondylus spinalis, as sug-
gested by Grijs.

Prionodactylus ocellifer Werner

Ecpleopus (Aspidolaemus) affinis Peters 1862, Abh. Akad. Wiss. Berlin: 199.
Prionodactylus ocellifer Werner, 1901, Verh. Zool.-Bot. Ges. Wien, 51: 596.

Prionodactylus ocellifer was described (Werner, 1901) from a single
specimen collected by Richard Haensch in Ecuador. Many of the labels for

PRIONODACTYLUS AND EUSPONDYLUS 51

Haensch’s collection were illegible, however, and no specific locality is
associated with the holotype. Other material in this collection came from
both high and low elevations on both sides of the Andes, as well as from
the inter-Andean valleys. No other examples of this taxon have been reported.
I have examined the holotype of P. ocellifer, ZMB 16593. It is an adult
female in poor condition. Werner’s description is adequate, but the following
points are worthy of note.

1. The loreal is large, in contact with the frontonasal, prefrontal, first
superciliary, frenoocular, nasal, supralabial two (broadly) and three (nar-
rowly; Fig. 23A, B.).

2. The first superciliary is large and expanded onto the dorsal surface of
the head; the superciliary series is complete.

3. The prefrontals are ever so slightly separated by contact between the
frontonasal and frontal.

4. Contact between supraoculars one and three at the superciliary series
excludes supraocular two from the superciliaries (Fig. 23C, D).

5. The disc in the lower eyelid is divided into three pieces by vertical
grooves.

6. There is a light-colored ring around the external ear opening; two or
three larger tubercles project into the opening from the anterodorsal margin
of the external ear opening (Fig. 23C, D).

The dorsal scales and the chin and throat scales are illustrated in Fig.
23E and 23F.

Most of these features (and many others) can be matched in the taxon
usually called Ecpleopus affinis (= Aspidolaemus affinis; Uzzell, 1969b). I
have compared the holotype of P. ocellifer with some of these (MCZ 45666-
67; FMNH 36714-22) and consider them to be conspecific, both on the
basis of the characters described by Werner as well as those mentioned above;
their general appearance also supports this conclusion, although the color-
ation is slightly different.5 Much of the color difference may be due to the
formalin-b\ackened condition of the specimens of A. affinis that I have
examined, and to the poor condition and desquamation of the holotype of
P. ocellifer.

I have examined the holotype of Aspidolaemus affinis (ZSM 644/0O). It
was carefully figured by Peters (1862); the specimen was labelled as type.
Despite Peters’ statement to the contrary, where epidermis persists, the
dorsal scales have a distinct but weak keel. There is no wrinkling on the
dorsals, however. I thus know of no trenchant differences between the holo-
types of P. ocellifer and A. affinis.

Boulenger (1885) reported specimens of E. (A.) affinis in the British
Museum from “western Ecuador” collected by Fraser, and from Intac
(Imbabura Province), Ecuador, collected by Buckley. Others of Fraser’s

5Benjamin Shreve made this identification without benefit of seeing the holo-
type.

Bye POSTILEA YW59

specimens, for example, specimens of Neusticurus ecpleopus and Euspondylus
maculatus, have erroneous locality data. Many specimens collected by Buckley
reportedly both at Pallatanga and eastern Andean localities were apparently
mislabelled (Peters 1955; Uzzell 1961). It is not unlikely that some other
Buckley locality data are also incorrect. There is at present no reason to
believe that Aspidolaemus affinis® occurs on the Pacific Andean slopes of
Ecuador. ;

In view of the similarities of Prionodactylus ocellifer and Aspidolaemus
affinis, | consider P. ocellifer to be a junior synonym of A. affinis (new
synonymy)’. The name P. ocellifer is available, however, to designate a
subpopulation within A. affinis should that prove useful.

IV. A NEw GENUS FOR Prionodactylus leucostictus

The only known specimen of Prionodactylus leucostictus Boulenger (1900)
was collected by F. V. McConnell and J. J. Quelch on the summit of
Mount Roraima (2150 m) in Guyana. I have examined the holotype (BMNH
99,3.25.4, reregistered as 1946.8.2.8). It is an adult female in relatively
good condition. While it bears a superficial resemblance to other species of
Prionodactylus because of its keeled, hexagonal dorsal scales, my observations
convince me that it is unrelated to Prionodactylus, but instead belongs with
Leposoma and its relatives (Ecpleopus, Alopoglossus, Ptychoglossus, Arthro-
saura, and possibly Amapasaurus; Ruibal, 1952; Uzzell, 1969b; da Cunha,
1970). Because P. leucostictus differs from each of these genera in a variety
of ways, I here erect a new genus for the taxon. The only known specimen
shows some anomalies in scalation. I believe, however, that acquisition of
additional material will reveal that most of the differences are real, and that
leucostictus cannot readily be placed into any of the already established
genera.

Riolama, new genus

TYPE SPECIES: Prionodactylus leucostictus Boulenger.

DESCRIPTION. Tongue with chevron-shaped plicae on anterior and posterior
thirds, with imbricate scalelike papillae on middle third. Snout moderately
long, blunt. Head scales without striations or rugosities. Single frontonasal
and frontal; a relatively broad interparietal extending posterior to parietals;
no median occipital (Fig. 24A). Paired prefrontals, frontoparietals, and

SA label with ZSM 644, 0 indicates that it was collected in Pichincha by Wagler.
The label is relatively recent, however, and Peters (1862) gave no such informa-
tion.

7Richard R. Montanucci, who is revising the systematics of Aspidolaemus and
Pholidobolus, concurs in this identification.

PRIONODACTYLUS AND EUSPONDYLUS 53

parietals; a pair of occipitals and postoccipitals behind interparietal. An
enlarged temporal scale lateral to parietal and occipital. Nostril pierced in
middle of nasal; no evidence of division above or below nostril although
there is an indistinct groove between the nostril and the first upper labial.
Nasals not in contact; on right side, nasal extended posteriorly almost to
anterior angle of eye; a loreal incompletely divided from the nasal by a
groove beginning near the posterior angle of the nasal. On left side, a loreal
is present, separated from labials. A large frenoocular and a series of
moderate infraoculars. Superciliary series complete; first superciliary short
on right side; fused with first supraocular on left. Four supraoculars on each
side, the first in contact with the prefrontals, separated from frontonasal.
Eyelids weli developed, the lower with a heavily pigmented central disc
divided into three pieces by vertical grooves. Ear opening large; tympanum
heavily pigmented, very shallowly recessed. A mental followed by one un-
paired and four paired chin shields (Fig. 24B); the two anterior pairs form
a median suture; the two posterior pairs are separated by small scales; the
posteriormost pair is separated from the infralabials by small scales. Pregular
(Ruibal, 1952) and gular scales are not clearly separated; collar fold very

FIG. 21. Structure of the left hemipenis of Cercosaura ocellata (BMNH 1927.-
8.1.150). The inverted organ has been slit along the sulcus spermaticus. A) The
entire organ showing the general arrangement of the flounces in the lateral (left)
and medial (right) pockets. x10. B) Detail showing the teeth in the arms of
the chevrons that join across the base of the median welt. x 30.

54 POSTILEA 159

well developed. Gulars flat, imbricate, irregular, large along midline, not
forming pairs of large scales, rounded behind; collar scales flat, six in
number including a large median scale, slightly rounded posteriorly. Limbs
pentadactyl; digits much depressed except at tip; distalmost subdigital
lamellae swollen (Fig. 25); digits clawed. Scales along inner margin of palm
between thumb and wrist enlarged, with a free medial edge; 13 lamellae
under fourth finger, the basal divided; 14 lamellae under fourth toe, the
four basal ones divided, but not tuberculate. Dorsal scales keeled, bluntly
hexagonal, in transverse rows. A zone of lateral scales four high, about half
as large as dorsals, about % size of ventrals; one or two upper rows keeled.
Ventral scales in transverse and longitudinal rows, smooth, rounded pos-
teriorly. Anterior and posterior median preanal scales; each lateral preanal
scale adjacent to both of these (Fig. 24C). Femoral pores well developed,
7 right, 6 left, none preanal.

DIAGNOSIS. The widely separated nasal scales, each surrounding a nostril,
and the pentadactyl limbs with all digits clawed place Riolama in Boulenger’s
(1885) Group II of the Teiidae. The combination of uniformly keeled,
hexagonal dorsal scales, slightly recessed tympanum, oblique plicae rather
than papillae on the anterior part of the tongue, and depressed digits with
swollen tips distinguish Riolama from all other genera in Group II.

RELATIONSHIPS. Several of the features of /eucostictus distinguish it from the
species I place in the genus Prionodactylus. In particular, the presence of
oblique folds rather than papillae on the anterior part of the tongue, the
large, nearly superficial heavily pigmented tympanum, and the markedly
depressed toes are features not seen in Prionodactylus. The last pair of
chin shields are widely separated from the infralabials, a condition that I
have not noted in Prionodactylus, and the arrangement of the preanal scales
with large anterior and posterior median scales, and large lateral scales that
are neither anterior nor posterior adjacent to these also does not occur in
Prionodactylus. Clearly, leuctostictus is not a Prionodactylus.

The presence of plicae rather than papillae on the anterior as well as the
posterior part of the tongue suggests that Riolama is close to Alopoglossus,
Ptychoglossus, and Ecpleopus, other genera of Group II that have plicae
rather than papillae on the anterior part of the tongue. In fact, the arrange-
ment of the plicae in Ecpleopus (present on posterior part and usually on
anterior part, but not the middle; Uzzell, 1969b) is an exact parallel for
the condition seen in Riolama leucosticta.

Other characters also support this arrangement, albeit weakly. The pos-
terior pair of chin shields are separated from the labials in several forms of
Leposoma, Alopoglossus, and Ptychoglossus (Uzzell, 1969b). This condition
is unusual in other genera of Group II. The arrangement of the prefrontals
with respect to the supraoculars is reminiscent of the condition in Ecpleopus
(cf. Uzzell, 1969b, fig. 1; Boulenger, 1899, fig. 3a) except that in Riolama,

PRIONODACTYLUS AND EUSPONDYLUS 55

the prefrontals are relatively much farther forward. The digits in Ecpleopus
are also slightly depressed, but not markedly so as in Riolama. The arrange-
ment of the preanal scales in Riolama is reminiscent of that seen in some
species of Arthrosaura (cf. da Cunha, 1967, fig. 1). The location of the
tympanum close to the surface of the head is also reminiscent of the con-
dition in Ecpleopus, although seen as well in such diverse genera as Neu-
sticurus, Placosoma, Anadia, and Echinosaura. None of these similarities
except the condition of the tongue, however, is more than suggestive.

On the other hand, it is easy to distinguish Riolama from Leposoma and
its relatives. I made a detailed comparison of those genera except for Amapa-
saurus (Uzzell 1969b, table 2). On the basis of data provided in the generic
diagnosis, I am not prepared to suggest that Riolama is closer to any one
of those genera than to the others.

The only known species of the genus is Riolama leucosticta (Boulenger),
new combination.

DERIVATION OF NAME. Riolama was the name and home of Rima, who in-
habited another isolated mountain of northern South America in W. H.
Hudson’s Green Mansions. It is dedicated to a friend akin to Rima in spirit.

B

FIG. 22. Structure of the hemipenis of Aspidolaemus affinis (USNM 194382).
The inverted organ has been slit along the sulcus spermaticus. A) The entire
organ showing general arrangement of the fiounces in the lateral (left) and
medial (right) pockets. x 6.5. B) Detail of teeth in the short rows in the flounces
that cross the base of the median welt and of the large tooth where two basal
chevrons of the two pockets meet at the middle of the median welt. x 18.

56 POSTILEAMI5S9

FIG. 23, Comparisons of the holotype of Prionodactylus ocellifer Werner (ZMB
16593) with recent specimens of Aspidolaemus affinis (FMNH 36720 and 36722)
A-F, <5. A) Side of the head of Aspidolaemus affinis (FMNH 36720). B) Side
of the head of Prionodactylus ocellifer «5. C) Top of head of Aspidolaemus
affinis (FMNH 36722). x5. D) Top of head of Prionodactylus ocellifer x5.
E) Dorsal scales of Prionodactylus ocellifer. F) Chin and throat scales of
Prionodactylus ocellifer.

V. COMMENTS ON SPECIES OF THE GENUS Euspondylus

My work on Prionodactylus necessitated consideration of the forms of
Euspondylus with which Prionodactylus has been long united (Burt and Burt,
1931). I record here observations of forms described in or referred to
Euspondylus other than those discussed as Prionodactylus. These comments,

PRIONODACTYLUS AND EUSPONDYLUS = |

together with those on Prionodactylus rahmi and P. spinalis, provide a
synopsis of the genus Euspondylus.

Euspondylus maculatus Tschudi

Euspondylus maculatus Tschudi, 1845, Arch. fiir Naturg. 11: 161.
Ecpleopus (Proctoporus) fraseri O’Shaughnessy, 1881, Ann. Mag. Nat.
Hist. (5) 4: 296.

This species, the type of the genus Euspondylus, was described by
Tschudi in 1845; additional information was provided in 1846; the type
locality was specified as Moyobamba in the front range of the Andes. This
locality is at about 750 m above sea level in the valley of the Rio Mayo, a
tributary of the Rio Huallaga in San Martin, Peru. There are two syntypes
in the Musée d’Histoire Naturelle in Neuchatel, Switzerland. Peters (1862)
examined these, described them carefully, and figured the male. I have
examined them also. Peters’ description is excellent; the second syntype is
a female. The male has about 42 scales around the midbody region, the
female, 44. The female appears to have a single femoral pore on each side.
The disc in the lower eyelid is divided into 2-3 parts by vertical grooves in
the male, into 3-4 in the female. There are 5 posterior preanal scales in the
female. According to Peters, the dorsal scales were slightly keeled in the
sacral region.

I have seen other specimens that I believe are this species. They come
from localities in southern and central Peru, on the Amazonian side of the
Andes; elevations that are believable vary from about 500-1500 m above
sea level.

Ecpleopus (Proctoporus) fraseri, described by O’Shaughnessey on the
basis of BMNH 58.7.25.14 (= 1946.8.2.48), a female, appears to be the
same taxon, but lacks prefrontals. This synonymy, suggested by Boulenger
(1885), appears to be correct; the type locality for E. fraseri (Guayaquil,
Ecuador) almost certainly is incorrect. An alternative possibility is Guala-
quiza, Morona-Santiago, Ecuador (600 m), where Fraser spent considerable
time in 1857 and early 1858. Most of Fraser’s collections of birds from
Ecuador in 1857 and 1858 were reported by Sclater (Chapman, 1926).
The localities for 1857 are all in eastern Ecuador. Fraser collected birds in
western Ecuador in late 1858 (August-December) and in 1859. The holo-
type of E. fraseri was accessioned 25 July 1858 (A. F. Stimson, personal
communication).

The large male in Brussels that Boulenger (1885) reported is MRHNB
951, I. G. 9422. Its locality is Chile. I believe that this is incorrect.

AMNH 1704, reportedly from Lake Aracona (= Lago de Aricoma),
Juliaca, above 4000 m, certainly is incorrectly labelled. Dunn (1942) re-
ported that Harvey Bassler suggested that American Museum material
labelled Juliaca was sent there by a member of the Inca Mining Company

58 POSTILLA 159

(apparently H. H. Keays), but was probably collected near the mine at
Santo Domingo, at about 1200 m above sea level in Puno (cf. Uzzell
1970: 26).

Other specimens of Euspondylus maculatus that have satisfactory locality
data include BMNH 1911.12.20.5-6 (Peru, Puno, Chaquimayo, 600-750 m)
and FMNH 40584, 45474 and AMNH 56268 (Peru, Junin, Chanchamayo,
1200-1300 m). The valid locality records suggest an eastern Andean dis-
tribution at 500-1500 m throughout Peru and southern Ecuador, rather than
the coastal areas of northern Peru and southern Ecuador suggested by
Peters and Donoso-Barros (1970).

Euspondylus simonsii Boulenger
Euspondylus simonsii Boulenger, 1901, Ann. Mag. Nat. Hist. (7) 7: 549.

This species was described by Boulenger (1901) from a single female
(BMNH 1900.11.27.34; reregistered as 1946. 8. 2. 3) collected at Punto-
yacu, Rio Perene, Peru, about 1250 m above sea level. No other specimens
have been reported.

Euspondylus simonsii can be distinguished from E. maculatus, which it
resembles, by the undivided disc in the lower eyelid and by the light line
between the angle of the mouth and the forelimb insertion. It may also
have a reduced number of scales around the midbody region. The palm of
E. simonsii has no enlarged scales between the thumb and wrist. Enlarged
scales with a projecting medial edge are present in E. maculatus.

Euspondylus guentheri

Ecpleopus (Euspondylus) guentheri O'Shaughnessy, 1881, Proc. Zool. Soc.
London 1881: 235.

This very distinct species, based on BMNH 80.12.8.18, reregistered as
1946.8.8.99, an adult female collected by Buckley near Sarayacu, Pastaza,
Ecuador, has been reported from other localities in eastern Ecuador by
Peters (1959).

Euspondylus brevifrontalis Boulenger

Euspondylus brevifrontalis Boulenger, 1903, Ann. Mag. Nat. Hist. (7)
12:°431.

The syntypes of E. brevifrontalis Boulenger (1903) include BMNH 1903.-
6.30.34, reregistered as 1946.8.31.62, an adult female from Rio Albireggas,
Venezuela (about 2900 m), and BMNH 1903.10.30.1-2, reregistered as
1946.8.32-33, two adult males from about 2500 m at Escorial, Venezuela,

PRIONODACTYLUS AND EUSPONDYLUS 59

FIG. 24. Scale features of the holotype of Riolama leucosticta (BMNH 1946.-
8.2.8.) A) Top of head 3.5. B) Chin and throat scales. «3.5. C) Scales of
cloacal region. x3. Based on Boulenger, 1900.

collected by Bricefio. As noted by Boulenger, these specimens are very
similar to Anadia bitaeniata. Although this species perhaps should be trans-
ferred to the genus Anadia (Olav Oftedal and E. E. Williams, personal com-
munication), minute differences between the taxa in arching of the distal
phalanx of the toes, division of throat scales into two groups by a gular
crease, and slightly smaller lateral scales may justify placing these two
taxa on different sides of a generic boundary. An examination of features in
addition to the scalation is needed to help place these two taxa correctly.
At present I retain this form in Euspondylus but with no real conviction.

60 POSTILLA 159

FIG. 25. Lateral view of the digits of the right forefoot of the holotype of
Riolama leucosticta (BMNH 1946.8.2.8), showing the general depressed shape of
the digit and the swollen digit tip. «25.

Euspondylus acutirostris (Peters)

Ecpleopus (Euspondylus) acutirostris Peters, 1862 Abh. Akad. Wiss. Berlin
1862: 209.

The holotype of Ecpleopus (Euspondylus acutirostris) Peters (1862) is
ZMB 4597, an adult male from Venezuela. I have examined both the holo-
type and the small female (UMMZ 125769) reported by Test et al. (1966)
from Rancho Grande, Aragua, Venezuela. I have compared the specimens
directly, and I see no difficulty in their being conspecific. The arrangement
of the subocular scales appears to be the same in both specimens.

The subdigital lamellae of the basal phalanges of the fore- and hind feet
of E. acutirostris are somewhat widened and flattened; the subdigital la-
mellae of the distal phalanx on each toe are compressed. Although the
distal phalanx is not arched, the arrangement is otherwise similar to that on
the toes of arboreal species of Anadia, and supports the hypothesis that this
may be an arboreal species (Test et al., 1966).

Other specimens of this taxon have been reported by Lancini (1968) from
the Cordillera de la Costa in Venezuela.

Euspondylus phelpsi Lancini

Euspondylus phelpsi Lancini, 1968, Pub. Ocas. Mus. Cienc. Nat., Zool.
1222:

I have seen no material of this recently described taxon (Lancini, (1968).
I am unable to determine whether it belongs in Euspondylus, Prionodactylus,

PRIONODACTYLUS AND EUSPONDYLUS 61

or some third genus. It is unusual among taxa referred to Euspondylus in
having a light line beginning in the supralabial region behind the eye and
passing above the forelimb insertion; such a light line is also present in
E. simonsii and E. acutirostris.

Euspondylus stenolepis Boulenger

Euspondylus stenolepis Boulenger, 1908, Ann. Mag. Nat. Hist. Ser. 8,
Dersg:

Euspondylus stenolepis was described from an adult male (BMNH
1909.4.30.64, reregistered as 1946.8.2.39) collected by M. G. Palmer, who
took it from the stomach of a bird shot at San Antonio, Valle del Cauca,
Colombia at about 2200 m above sea level. Boulenger’s (1908) description
and figure are very good; most of the characters of E. stenolepis that argue
for its generic placement can be seen in Boulenger’s figure, which is re-
produced here as Figure 26. There are four supraoculars, the anteriormost
elongate; all are separated from the palpebrals by a2 complete row of super-
ciliaries (Fig. 26A). The narrow interparietal and the parietals are almost
coequal in posterior extension (Fig. 26A). The loreal is a diagonally placed
scale, not reaching the supralabiais, and extending dorsal to a line con-
necting the dorsal edge of the first superciliary and the nasal (Fig. 26B).

FIG. 26. Scale features of the holotype of Euspondylus stenolepis (BMNH 1946.-
8.2.39). x3. A) Top of head. B) Side of head. C) Chin and throat. D)
Cloacal region. After Boulenger, 1908.

62 POSTIELAMID9

The first superciliary is elongate (Fig. 26B). The fourth, posteriormost pair
of chin shields is separated from the infralabials by a narrow, elongate
scale (Fig. 26C). None of the median gular scales is widened; all are some-
what elongate rectangles (Fig. 26C). There are four elongate posterior
preanal scales (Fig. 26D). There are no preanal pores (Fig. 26D).

Additional characters indicating the generic placement of E. stenolepis
include the very narrow, obtusely keeled dorsal scales, the smooth gular,
ventral, and preanal scales, and the elongate, rectangular shape of the
ventrals.

I have examined the holotype of E. stenolepis. The most important fea-
ture to be added to Boulenger’s description is the structure of the tongue,
which has chevron-shaped plicae with the apices of the chevrons anterior.
The scales on the upper and lower parts of the forelimb are smooth. In
addition, the scales on the underside of the palm, between the base of the
thumb and the wrist, are enlarged and have free medial edges. The subdigital
lamellae at the bases of the third and fourth toes are divided, and the
anterior member of each pair is swollen into a tubercle.

This collection of characters, especially the structure of the tongue, the
elongate first superciliary and first supraocular, the coequal extension of
the interparietal and parietals, the narrow, elongate dorsals and ventrals,
the elongate posterior preanal scales, and the enlarged thenar scales on the
palm of the forefeet clearly identify stenolepis as a member of the genus
Ptychoglossus Boulenger (1890) and distinguish it from the type species
of Euspondylus. This form must therefore stand as Ptychoglossus stenolepis
(Boulenger), new combination.

VI. A KEY TO THE SPECIES OF Euspondylus

The following key is modified from Peters and Donoso-Barros’ (1970) key.
It has not been tried on all species, and only on limited samples.

1. Either all dorsal scales smooth or wrinkled, or most smooth with some

weakly keeled on posterior part of body ae ae me
All dorsal scales weakly to strongly keeled WOE 5s)
2. 40-50 scales from occiput to posterior edge of hind limbs; ventral sur-
face uniform lead colored E. brevifrontalis
39 or fewer dorsal scale rows: ventral surface often heavily marked
with gray, but not uniform lead colored .......... oth oct eS
3. Infraorbitals smaller than upper labials DoS a ee
Infraorbitals subequal to upper labials _. . _ E. guentheri

4. Transparent disc in lower eyelid divided; 40 scales around midbody
region: no light line between tympanum and forelimb E. maculatus
Transparent disc in lower eyelid not divided; 35 or fewer scales

PRIONODACTYLUS AND EUSPONDYLUS 63

around midbody region; a light line between tympanum and _ fore-
ITED ks once ey een _ E. simonsii

5. Fewer than 25 transverse rows of ventral scales between collar and
preamal scales .2..........5.:. . E. acutirostris
More than 25 transverse rows of eeatrall ecales between collar and

CAM AUISCALCS: 8s by-B evs ostissctveesenes a an eres cee Meee fae)
6. Four supraoculars, first of series smallest E. rahmi
MATES SSUPHAOCUIAES ooo e nei cv sesncs cnet. =a CPE ee ae al
7. No light lateral line on lip or above limb insertion E. spinalis

A light lateral line on posterior supralabials, below ear opening, and
above limb insertion E. phelpsi

VII. STATUS OF TAXA PROPOSED IN OR REFERRED TO THE GENERA
Prionodactylus AND Euspondylus

Some 19 taxa have been named in or referred to the genus Prionodactylus.
Names preceded by asterisks have not been discussed in this paper.

*albostrigatus Griffin 1917

argulus Peters 1862
bolivianus Werner 1899

*champsonotus Werner 1910

columbiensis Werner 1916
dicrus, new species
eigenmanni Griffin 1917
holmgreni Andersson 1914
leucostictus Boulenger 1900
manicatus O’Shaughnessy 1881

marianus Ruthven 1921
ocellifer Werner 1910
okendeni Boulenger 1907

oshaughnessyi Boulenger 1885
*quadrilineatus Boettger 1876

palmeri Boulenger 1908

rahmi Grijs 1936

spinalis Boulenger 1911

vertebralis O'Shaughnessy 1879

Pantodactylus schreibersii
albostrigatus
Prionodactylus argulus
Prionodactylus manicatus
bolivianus
Placosoma cordylinum
champsonotus
Prionodactylus argulus
Prionodactylus dicrus
Prionodactylus eigenmanni
Prionodactylus argulus
Riolama leucosticta
Prionodactylus manicatus
manicatus
Prionodactylus vertebralis
Aspidolaemus affinis
Prionodactylus manicatus
bolivianus
Prionodactylus argulus
Pantodactylus quadrilineatus
Prionodactylus vertebralis
Euspondylus rahmi
Euspondylus spinalis
Prionodactylus vertebralis

64 POSTILLA U59

Some 14 forms have been named in or referred to Euspondylus in addi-
tion to the names above that were explicitly or implicitly treated as
Euspondylus by Burt and Burt (1931). Names preceded by asterisks have
not been discussed in this paper.

acutirostris Peters 1862 Euspondylus acutirostris
ampuedae Lancini 1968 (?)Prionodactylys vertebralis
brevifrontalis Boulenger 1903 Euspondylus brevifrontalis
*cupreus Andersson 1916 Placosoma cordylinum
champsonotus
*festae Peracca 1897 Neusticurus strangulatus
fraseri O'Shaughnessy 1879 Euspondylus maculatus
guentheri O’Shaughnessy 1881 Euspondylus guentheri
maculatus Tschudi 1845 Euspondylus maculatus
*ocellatus Gray 1845 Anadia ocellata
Dhelpsi Lancini 1868 Euspondylus phelpsi
*rhombifer Gunther 1859 Anadia rhombifera
simonsii Boulenger 1901 Euspondylus simonsii
stenolepis Boulenger 1908 Ptychoglossus stenolepis
*strangulatus Cope 1868 Neusticurus strangulatus
ACKNOWLEDGMENTS

Many people have aided me in this study. For use of material in their
charge, I thank William E. Duellman, University of Kansas Museum of
Natural History; Alice G. C. Grandison, British Museum (Natural History) ;
Jean Guibé, Muséum National d’Histoire Naturelle; Walter Hellmich, Zoo-
logische Staatssammlung, Munich; Alan Leviton, California Academy of
Sciences; Hymen Marx, Field Museum of Natural History; George S. Myers,
formerly of Stanford University Natural History Museum; Hno. Nicéforo
Maria, Instituto de La Salle, Bogota; Giinther Peters, Zoologisches Museum,
Berlin; James A. Peters, U. S. National Museum; Neil Richmond, Carnegie
Museum; Hobart Smith, formerly of the University of Illinois Museum of
Natural History; Greta Vestergren, Naturhistoriska Riksmuseet, Stockholm;
Ernest E. Williams, Museum of Comparative Zoology, Harvard University;
John Wright, Los Angeles County Museum; Charles F. Walker, University
of Michigan Museum of Zoology; and Richard G. Zweifel, American
Museum of Natural History. For hospitality while I visited their institu-
tions, I particularly thank Dieter Fuchs (Zoologische Staatssammlung,
Munich), Miss Grandison and A. F. Stimson (British Museum), Jean
Guibé, Hymen Marx and Robert F. Inger (Field Museum), Giinther Peters,
James A. Peters, Charles F. Walker, Ernest E. Williams, Richard Zweifel
and Charles Myers (American Museum). Most of the illustrations are the
work of A. H. Coleman, photographer, and Rosanne Rowen, draftsman; the

PRIONODACTYLUS AND EUSPONDYLUS 65

photograph for Figure 25 was presented by the British Museum (Natural
History). Computer time was available through National Science Founda-
tion Grant 29114. Travel in Europe was made possible by Yale University
and a gift from Evan Commager.

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PRIONODACTYLUS AND EUSPONDYLUS 67

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