A New Species of Lizard of the Genus Ameiva (Teiidae) from the Pacific Lowlands of Colombia

Author(s): Arthur C. Echternacht

Source: Copeia, Vol. 1977, No. 1 (Mar. 16, 1977), pp. 1-7
Published by: American Society of Ichthyologists and Herpetologists (ASIH)
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1977 No. 1 COPEIA March 16

A New Species of Lizard of the Genus Ameiva

(Teiidae) from the Pacific Lowlands of Colombia
ARTHUR C. ECHTERNACHT

Ameiva anomala is described from a series of specimens from the De-

partments of Choc6, Valle and Cauca in the Pacific lowlands of Colombia.
The new species differs from all of its congeners and from all other
"macroteiids" in the granular nature of the anterior ventral scales. A
key to the seven known or expected species of Ameiva of trans-Andean
South America is provided. These species generally replace one another
in linear fashion in the Pacific lowlands of northwestern South America.

THE bird fauna of trans-Andean northwest- Ameiva anomala sp. nov.

ern South America has been long recog-
nized for its high species diversity and extensive
endemism (Haffer, 1967, 1974). Although the
degree of sampling and analysis of other verte-
brate groups in this area lags far behind that
achieved for birds, it is apparent that the rain-
forest cul-de-sac west of the Andes in Colom-
bia and Ecuador has also been a center of evo-
lutionary activity for reptiles and amphibians.
Lizards of the genus Ameiva (Teiidae) are a
conspicuous element of the terrestrial fauna of
these wet lowlands and the more arid region
bounding it to the south. Of the five species
of Ameiva presently known from the area,
four are endemic [bridgesi (Cope), edracantha
Bocourt, orcesi Peters, septemlineata Dumeril].
The fifth, festiva Lichtenstein and Von Mar-
tens, is represented by an endemic subspecies
(eutropia Cope). In faunal works on this re-
gion, only the Ameiva of Ecuador have re-
ceived adequate taxonomic treatment (Peters,
1963). Medem (1969) wrote of the Colombian
species "Respecto a la taxonomia y nomen-
clatura, el genero Ameiva esta en un estado de
alta confusi6n y necesita urgentemente una re-
visi6n total." It is not, therefore, surprising to
find an unrecognized species of Ameiva in the
Pacific lowlands of Colombia. Because of the
unusual anterior ventral scales, which distin-
guish the new species not only from its con-
geners but from all other macroteiids, I name
it:
I
Figs. 1 and 2
Holotype.-American Museum of Natural His-
tory 109694, an adult male from Quebrado
Guangui, 0.5 km above Rio Patia (Upper Rio
Saija drainage), Depto. Cauca, Colombia, 100-
200 m, February 1973. C. W. Myers and J. W.
Daly.
Cauca: Same locality as holo-
Paratypes.-COLOMBIA:
type, AMNH 107908-9, 109685-93. Choc6: Boca de la
Raspadura, AMNH 18269-71; Condoto, UMMZ 121464;
2 km above Playa de Oro, upper Rio San Juan, AMNH
108440; Pangala and Quebrado Pangala, ca. 40 km by
river N Palestina, lower Rio San Juan, AMNH 111040-50;
Quebrado Docord6, ca. 110 km by river N Palestina,
AMNH 111054; Quebrado Taparal, ca. 12 km by river N
Palestina, lower Rio San Juan, AMNH 111051-53, MCZ
112368-69; Quebrado Vicord6, ca. 5 km by river above
Noanama, Rio San Juan, AMNH 108439; 30 km up Rio
Pune, Rio Atrato drainage, AMNH 18272-73; Rio San
Juan, AMNH 108993. Valle: Anchicaya, KU 152678-81;
Rio Zabaleta, nr. Zabeleta, AMNH 108992; Virology
Field Laboratory, Rio Raposo, USNM 151619; nr. "Ra-
poso River Station" on Rio Raposo, UMMZ 126891. See
the Acknowledgments section for a key to abbreviations of
museum names.
Diagnosis.-A moderately large (maximum ob-
served snout-vent length 110 mm), striped
Ameiva distinguished from other species of
Ameiva and from all other macroteiids (Teii-
nae: see MacLean, 1974) by the granular ven-
tral scales of the chest and the resulting low
number (19-23) of longitudinal rows of en-
larged ventral scales.
Description of the holotype.-Snout-vent length
(SVL) 97 mm; tail length 244 mm; head length

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2 COPEIA, 1977, NO. 1

f11

- I Nm

Fig. 1. Dorsal (A) and ventral (B) views of the holotype of Ameiva anomala. Note especially the granu-
lar scalesin the chest region.

25.9 mm. Tongue sheathed posteriorly. Ros-
tral entire; nasal divided, nostril in prenasal-
postnasal suture; frontonasal entire but deeply
grooved longitudinally; postnasals contacting
prefrontals; prefrontals paired, strongly keeled,
separated medially by a pair of smaller keeled
scales; ca. 11 scales in the area normally occu-
pied by the frontal, a series of 3 strongly keeled
scales on either side, the anteriormost separated
medially by 2 smaller keeled scales, other scales
in the area smooth or weakly keeled; many
small, irregularly arranged and usually keeled
scales in the area normally occupied by fronto-
parietal and parietal scales with a single keeled,
moderately large, heart-shaped scale in the posi-
tion of the interparietal; 2 strongly keeled, sub-
equal supraoculars; 5 superciliaries, the second
5-10 times larger than the others; 3 or 4 rows of
small (granular) circumorbital scales separating
the posterior half of the second supraocular
from the median head scales, becoming 2 rows
to the middle of the first supraocular and a sin-
gle row forward of that point; 3 rows of small
(granular) scales separating the superciliaries
from the posterior half of the second supraocu-
lar, 2 rows separating the rest of the supraocu-
lar series from the superciliaries; 2 loreals, the
smaller posteroventral to the larger and 5-6
times smaller; one strongly keeled preocular; 3
left, 4 right strongly keeled suboculars, the

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 ECHTERNAGHT-NEW
jr
P:
Fig. 2. TopoparatypicAmeiva anomala, AMNH 109688,female, snout-ventlength 52 mm. Collectedand
photographedby CharlesW. Myers. 14 February 1973.
second 2 (right) or 3 (left) times the length of
the others, which are subequal in size; 7 supra-
labials; 5 infralabials; a single postmental fol-
lowed by 4 pairs of chin shields, none in con-
tact medially, the posterior 3 chin shields
separated by smaller scales from the infra-
labials; no enlarged gular scales; mesoptychial
scales small with fields of slightly larger, ir-
regularly arranged scales on either side of the
midline; 258 dorsal granules at midbody (GAB);
64 granules between (and including) the dor-
solateral light stripes (PV); 5 granules across
the vertebral stripe at midbody (VG); 307 gran-
ules occiput to rump (GOR); 6 longitudinal
rows of enlarged ventral scales at midbody, the
outermost a third the width of the others;
scales in the chest region granular and irregu-
lar in arrangement; 21 transverse rows of en-
larged ventral scutes; 17 preanal scales in the
midline, the 16th much enlarged and circular,
the 17th scales (terminal preanals) paired and
each about a fourth the size of the 16th; no
preanal spurs; caudal scales keeled, 16 scales
in the 15th verticel; prebrachial scales granular
except a few moderately enlarged, keeled
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AMEIVA 3
scales near the forearm on the left side only; a
single row of much enlarged preantebrachial
scales; postbrachials much enlarged, irregular
in arrangement; postantebrachials not en-
larged; 2 or 3 rows of much enlarged prefe-
moral scales; 2 rows of enlarged infratibial
scales; 27 left, 25 right femoral pores; 21 left,
21 right subdigital lamellae on the fourth hind
toes.
Color and pattern in alcohol: Dorsal ground
color black with indistinct, irregular gray
blotches; middorsal field divided by a narrow,
grayish-white vertebral stripe (pale brown on
the head) beginning on the rostral scale and
continuing onto the tail; narrow grayish-white
dorsolateral light stripes bordering the middor-
sal field, beginning above and just behind the
eyes and continuing onto the tail; velvet black
dorsolateral dark field (stripe) extending from
behind the eye to base of the tail; narrow gray-
ish-white ventrolateral stripe beginning below
eye and continuing over the ear to the hind
leg, thence from behind the femur onto the
tail; ventrolateral field gray, containing a nar-
row, wavy, occasionally interrupted grayish-
4 COPEIA, 1977, NO. 1
white stripe originating below the ear and ter-
minating at the anterior point of insertion of
the hind limb; posterior half to two-thirds of
the tail brown; limbs dark gray to black above
with irregularly arranged grayish-white lines
and spots on lower hind leg; all ventral sur-
faces blue-gray except femoral pore scales
white; head brown above, gray on sides except
for a black stripe interrupted by the eye ex-
tending from the nares across the upper half
of the loreal and to a point above the ear.
Variation.-Twenty-two males and 23 females
were available for study. Sexual dichromatism
was not evident, nor was sexual dimorphism
apparent for any of the morphometric or scale
characters summarized below. Four samples
were sufficiently large for preliminary statisti-
cal analysis: Vicinity of Pangala (N = 11) and
Quebrado Taparal (N = 5), Depto. Choc6;
Anchicaya and localities on the Rio Raposo,
Depto. Valle (N = 6); and Quebrado Guangui,
Depto. Cauca (N = 12). No geographic trends
were noted and the samples did not differ in a
statistically significant manner with respect to
any of the characters examined. A summary of
variation for the total sample is given below.
For each character, the range is given followed
by (in parentheses) the mean, estimated stan-
dard deviation of the mean, and sample size.
Characters and abbreviations are as defined by
Echternacht (1971).
Maximum observed SVL 109 mm for males,
110 mm for females; maximum observed tail
length 244 mm for males, 249 mm for females;
tail length/total length ratio 0.68-0.72 (0.70 ?
0.01, 21); supralabials 12-14 (12.5 ? 0.79, 45);
infralabials 10-12 (10.3 ? 0.58, 45); supaocu-
lars 4-6 (4.1 + 0.44, 45); COP 4-8 (7.1 ? 1.08,
45); SO-SC 5-10 (8.4 ? 1.46, 45); loreals 2-7
(3.2 ? 1.3, 45); scales between eyes 3-6 (4.2 ?
0.68, 45); GAB 228-295 (253.8 ? 14.49, 40); VG
4-9 (6.3 ? 1.35, 36); PV 46-76 (59.1 ? 5.24, 40);
GOR 270-370 (312.9 ? 20.26, 44); VG/GAB
0.015-0.038 (0.025 ? 0.006, 35), PV/GAB 0.20-
0.27 (0.232 ? 0.012, 38); GAB/GOR 0.71-0.91
(0.81 - 0.05, 39); SAT 12-16 (15.4 ? 0.94, 45);
total preanal scales 10-20 (14.4 ? 2.93, 45);
terminal scales 1-4 (2.3 ?
preanal 0.67, 45);
total femoral pores 41-59 (49.9 ? 4.49, 43); sub-
digital lamellae on 4th toe of left hind foot
21-26 (23.6 ? 1.22, 43); longitudinal rows of
ventral scales 19-23 (21.1 ? 0.87, 45); transverse
rows of ventral scales 4-6 (6.0 ? 0.30, 45).
Small specimens may have 7 distinct stripes,
but some large individuals lose the ventral-
most stripe on either side leaving only the ver-
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tebral stripe and those bordering the dorso-
lateral dark field. In very large specimens, the
vertebral stripe may be lost or quite faded.
Paired blotches may occur on either side of
the vertebral stripe in larger specimens (e.g.,
the holotype and UMMZ 126891), but these are
absent in hatchlings and subadults and not
present in all adults.
Color and pattern in life.-The following is ex-
cerpted from my field notes of 21 May 1971 de-
scribing a series of 4 Ameiva anomala (3 juve-
niles, SVL 42-59 mm, KU 152678-80; 1 adult
female, SVL 82 mm, KU 152681) collected at
Anchicaya, Depto. Valle: Juveniles "Dorsum
velvet black with a golden-orange vertebral
stripe beginning on the rostrum and extending
onto the tail where it becomes irridescent blue-
green, then irridescent blue. Dorsolateral stripes
. . . golden-orange turning blue-green poste-
riorly and on the tail paralleling the color
change of the vertebral stripe. Sides velvet
black with an orange ventrolateral stripe be-
ginning beneath and behind the eye and ex-
tending to the axilla of the hind leg. A con-
tinuation of this stripe, but blue, runs along
the side of the tail. The hind legs are mot-
tled with blue and green on black; the fore-
limbs are mottled with orange as is the side
of the head below the eye. The venter is
coppery to the hind legs, then blue-green onto
the ventral surface of the tail. The adult is
as described above except the stripes on the
tail are more distinct and less irridescent. The
ventral surface of the tail is white and black."
Color notes provided by C. W. Myers on a
series of anomala from several localities are
in general agreement with my own with the
exception that the stripes are variously de-
scribed by Myers as "vivid yellow" (AMNH
108439, 72 mm SVL) and "yellow" (AMNH
109688-90, 52-84 mm) or "orange-yellow"
(AMNH 108440, 59 mm), the vertebral stripe
becoming "greenish-blue" or "blue" posteriorly
on smaller individuals.
Comparisons.-Of the seven species of Ameiva
known or expected in trans-Andean South
America, only festiva occurs sympatrically with
anomala, both species having been collected at
Pangala and Condoto (Depto. Choc6) in Co-
lombia. Ameiva festiva is found from Estado
Tabasco, Mexico through Central America to
Pacific coastal and inter-Andean valleys of
northern Colombia (Echternacht, 1971). Ameiva
leptophrys ranges from southeastern Costa
Rica through the Pacific lowlands of Panama
 ECHTERNACHT-NEW AMEIVA 5

and has been collected in extreme eastern 2. Chest scales small, sharply differentiated
Panama. It is to be expected in adjacent north- from posterior ventral scales, and ir-
western Colombia, and it is sympatric with regular in arrangement anomala
festiva over its entire range although differ- Ventral scales enlarged overall, arranged
ences in habitat preference have been noted in well-defined rows between mesop-
(Echternacht, 1971). In Colombia, bridgesi tychium and preanal region 3
has been found in the vicinity of the Rio Ma- 3. Mesoptychial scales enlarged, in a trans-
taje at the border between Colombia and Ecua- verse series; prebrachial scales much en-
dor and at Tumaco (Depto. Nariio), and on larged, in one or two rows __septemlineata
Isla Gorgona (Depto. Cauca). Records of Mesoptychial scales not or slightly en-
Ameiva from between Quebrado Guangul and larged, if enlarged usually not in con-
Tumaco are lacking (Fig. 3), so the exact tinuous series; prebrachial scales not
northern range limit of bridgesi and the south- enlarged or, if enlarged, slightly to mod-
ern limit for anomala are unknown. Ameiva erately with a distinct keel --___---bridgesi
septemlineata replaces bridgesi in northwestern 4. Midgular scales much enlarged, the largest
Ecuador and ranges south to at least Zaruma larger than the rostral scale festiva
(El Oro Prov.) where it is itself replaced by Midgular scales not or moderately en-
edracantha. Ameiva edracantha occupies arid larged, the largest always smaller than
and semiarid habitats in southwestern Ecuador the rostral scale 5
and northwestern Peru. Ameiva orcesi is en- 5. Midgular scales subequal; postnasal con-
demic to the arid valley of the Rio Jubones in tacts prefrontal orcesi
southwestern Ecuador, east of the ranges of Posterior gular scales smaller than ante-
both septemlineata and edracantha. rior gular scales; postnasal does not
Ameiva anomala is most closely related to contact prefrontal 6
bridgesi and septemlineata but both of these 6. Five parietal scales, only the lateral-most
lack the irregularly arranged, granular scales separated from the frontoparietal; total
of the chest region of anomala. Ameiva festiva femoral pores 35 or fewer; males with
has eight transverse rows of ventral scales at preanal spurs edracantha
midbody, a transverse band of enlarged mesop- Three parietal scales (when not irregu-
tychial scales, one or more very large midgular larly subdivided), the lateral parietal
scales, and the anterior scales of the chest are scales separated from the frontoparietal
not reduced. Both festiva and edracantha gen- by one or more rows of smaller scales;
erally have the "normal" pattern of dorsal total femoral pores 40 or more; males
head scales for macroteiids in that the frontal without preanal spurs leptophrys
is undivided. All of the known or expected
trans-Andean Ameiva except leptophrys have Range.-Ameiva anomala is presently known
only from the Pacific lowlands of Colombia
prominent vertebral stripes although the stripes from the Rio Atrato southward
upper
may fade in larger (older) individuals. Details to central Cauca at the region
of color and pattern are, however, often lost in type locality. This
comprises a latitudinal range of little more
preservative and I have found the frontal (at
to be divided in some of than 3? (ca. 350 km airline) and the species
least) specimens every
has nowhere been found more than ca. 80 km
species of Ameiva that I have examined (Ech-
ternacht 1970, 1971). For these reasons, the from the coast (Fig. 3).
following key to the identification of trans- Remarks.-Ameiva anomala has been found in
Andean species of Ameiva relies on other both
primary and secondary forests as well as
characters. Those characters which distinguish in
edge situations along roads and riverbanks.
bridgesi and septemlineata, the species most Individuals were most often encountered sun-
likely to be confused, are clearly illustrated by ning or prowling in patches of sun. I never
Peters (1963; Figs. 1 and 2). found them in grass or in cut-over areas. None
were seen on overcast days, nor were they
A Key to Known or Expected Trans- found active in even very light rain. The four
Andean Species of Ameiva
specimens collected at Anchicaya were all ob-
1. Six transverse rows of ventral scales at tained during a one-half hour sunny period in
midbody 2 the midst of a three day span of almost con-
Eight transverse rows of ventral scales at stant overcast during which no other Ameiva
midbody 4 were seen.

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6 COPEIA, 1977, NO. 1

Fig. 3. Locality recordsof Ameiva anomala (solid symbols) and Colombian records of Ameiva bridgesi
(hollow symbols). The off-shore locality for bridgesiis Isla Gorgona,Depto. Cauca. The dashed line rep-
resents the approximateposition of the western cordilleraof the Andes.

Etymology.-The name anomala is derived from
the Latin adjective anomalus, in reference to
condition of the anterior ventral scales.
ACKNOWLEDGMENTS
I am indebted to the following individuals
for allowing me to examine specimens in their
care and for relevant information: Charles W.
Myers and Richard G. Zweifel of the American
Museum of Natural History (AMNH), Ernest
E. Williams and Patricia Haneline of the Mu-
seum of Comparative Zoology (MCZ), the late
James A. Peters of the National Museum of
Natural History (USNM), William E. Duellman
and Alan H. Savitzky of the University of Kan-
sas Museum of Natural History (KU), and Ar-
nold G. Kluge and Scott Moody of the Univer-
sity of Michigan Museum of Zoology (UMMZ).
Federico Medem very kindly supplied informa-
tion on some Colombian localities, and the late
F. Carlos Lehman made possible my visit to
Cali. Capable field assistance was provided by
Sr. Humberto Salazar.
I am especially grateful to Charles W. Myers
for his efforts in obtaining specimens of
Ameiva incidental to his own field studies in
Colombia, for his patience as a source of in-
formation, and for the use of the photograph
of a living Ameiva anomala.
My studies of the Ameiva of northwestern
South America have been supported in part by
the Boston University Graduate School and the
Penrose Fund (Grant No. 5980) of the Ameri-

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 ECHTERNACHT-NEW AMEIVA 7

can Philosophical Society. Fred Maynard pro-
vided photographic assistance.
LITERATURECITED
ECHTERNACHT, A. C. 1970. Taxonomic and eco-
logical notes on some Middle and South American
lizards of the genus Ameiva (Teiidae). Breviora
354:1-9.
. 1971. Middle American lizards of the
genus Ameiva (Teiidae) with emphasis on geo-
graphic variation. Univ. Kansas Mus. Nat. Hist.
Misc. Publ. 55:1-86.
HAFFER, J. 1967. Speciation in Colombian forest
birds west of the Andes. Amer. Mus. Novit. 2294:
1-57.
. 1974. Avian speciation in tropical South
America with a systematic survey of the Toucans
(Ramphestidae) and Jacamars (Galbulidae). Publ.
14 Nuttall Ornith. Club.
MACLEAN,W. P. 1974. Feeding and locomotor
mechanismsof teiid lizards: functional morphol-
ogy and evolution. Paplis Avulsos Zool. 27:179-
213.
MEDEM,F. 1969. El desarrollo de la herpetologia
en Colombia. Rev. Acad. Colombiana de Cs.
Exact., Fis., Nat. 13:149-199.
J. A. 1963. The lizard genus Ameiva in
PETERS,
Ecuador. Bull. So. California Acad. Sci. 63:113-
127.
DEPARTMENT OF ZOOLOGY AND GRADUATE PRO-
GRAM IN ECOLOGY, THE UNIVERSITY OF TEN-
Ac-
NESSEE, KNOXVILLE, TENNESSEE 37916.
cepted 3 March 1976.

A New Western Atlantic Wormfish

(Pisces: Microdesmidae)
C. E. DAWSON
Microdesmus luscus n. sp., closely related to M. lanceolatus (Gulf of
Mexico), M. hildebrandi and M. multiradiatus (Eastern Pacific), is de-
scribed from Caribbean waters. A key is provided to Western Atlantic
species of Microdesmus.

SIX species of wormfishes (Microdesmidae) Paratypes.-GCRL 14237, 26.5 mm (cleared and

have been described from western Atlantic
waters. These include Cerdale fasciata and C.
floridana (see Dawson, 1974), and four species
of Microdesmus (M. carri, M. bahianus, M. lan-
ceolatus and M. longipinnis). Since there will
be some delay in completing my review of the
genus, I here describe an unusual new species
of Microdesmus from the Caribbean Sea.
Methods follow Dawson (1974). Measure-
ments are in millimeters (mm), proportions are
in % standard length (SL) or head length (HL),
collection depth is reported in meters (m), lati-
tude and longitude are approximations.
Type material has been deposited in collec-
tions of the Academy of Natural Sciences of
Philadelphia (ANSP), Gulf Coast Research Lab-
oratory Museum (GCRL) and National Museum
of Natural History, Smithsonian Institution
(USNM).
Microdesmus luscus n. sp.
Fig. 1
Holotype.-USNM 198261, 45.9 mm, Dominica,
B.W.I., W coast, rock and coral area just S of
Mahaut, 15? 21' 28" N, 61? 24' 39" W, 0-7.6 m,
29 Oct. 1964, VGS 64-10, V. G. Springer and R.
H. Reckweg.
stained), Puerto Rico, La Parquera, Laurel reef,
17? 56' 40" N, 67? 03' 30" W, 1 m. ANSP
133205, Panami, Canal Zone, San Lorenzo, 09?
19' 30" N, 80? 02' 00" W.
Diagnosis.-Eye inconspicuous; caudal fin lan-
ceolate; dorsal fin origin in advance of vertical
from tip of adpressed pectoral fin; pectoral fin
13-14; gill opening broad, not tubiform; dor-
sal fin elements total 50-51; anal rays 35;
caudal vertebrae more numerous than abdomi-
nal vertebrae; 1st proximal dorsal pterygio-
phore inserted between neural spines 3 and 4.
Description.-Dorsal fin XIII, 37-38, total ele-
ments 50-51; anal fin 35; pelvic fin I, 3; seg-
mented caudal rays 17; vertebrae 21 + 29 = 50.
Measurements (mm) of holotype are followed
in parentheses by range, in % SL or HL*, of
holotype and paratypes: caudal fin length 6.0
(13.1-15.6), caudal peduncle depth 1.3 (2.8-
3.8), body depth 2.7 (5.4-5.9), predorsal length
8.2 (17.9-18.9), preanal length 22.6 (49.2-51.0),
pectoral fin length 3.3 (7.2-9.2), pelvic fin
length 3.2 (7.2-9.1), pelvic-anal length 16.7
(35.8-36.4), head length 6.2 (13.5-15.8). Eye
diameter 0.4 (5.9-7.6*), preorbital length 1.1

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