Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on the types and type locality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and description of a new species of Eleutherodactylus from Mato Grosso, Brazil (Amphibia: Anura: Leptodactyl

18 Pages·1999·8 MB·

by  W Ronald Heyer

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(1):1-18. 1999.
Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on
the types and type locality of Telatrema heterodactylum Miranda-
Ribeiro, 1937, and description of a new species of Eleutherodactylus
from Mato Grosso, Brazil (Amphibia: Anura: Leptodactylidae)
W. Ronald Heyer and Anna M. Munoz
(WRH) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560-0109, U.S.A.;
(AMM) Department of Biology, New Mexico State University, Las Cruces, New Mexico 88003,
U.S.A.; Current address: 1306 Alvarado, Carlsbad, New Mexico 88220, U.S.A.
—
Abstract. Examination of the types of Eleutherodactylus crepitans Boker-
mann, 1965, indicates that they represent a valid species, which shoilld be
removed from the synonymy of Eleutherodactylus fenestratus (Steindachner
1864). Eleutherodactylus crepitans is only known from the type material. Cer-
tain features of the types of Telatrema heterodactylum Miranda-Ribeiro, 1937,
are described and the type locality clarified. Analysis of the advertisement call
and morphology of a series of specimens from Chapada dos Guimaraes indi-
cates that the specimens represent a species distinct from E. fenestratus (Stein-
dachner 1864), for which no name is available. We describe this new species
as Eleutherodactylus dundeei. The frog fauna of Chapada dos Guimaraes con-
tains four distinct historical units: cerrado, chaco, Amazonian hylaea, and en-
demic.
The advertisement call of what has been whether it has a distribution beyond the re-
identified as Eleutherodactylus fenestratus gion of the town of Chapada dos Guima-
from Chapada dos Guimaraes, Mato Gros- raes, we examined other specimens of
WRH
so, Brazil, was recently recorded by Eleutherodactylus from Mato Grosso, in-
PRODEAGRO. PRODEAGRO
as part of is cluding the type material of Telatrema het-
an ecological-agricultural survey of the erodactylum Miranda-Ribeiro, 1937 and
State of Mato Grosso, undertaken by the Eleutherodactylus crepitans Bokermann,
State Secretariat of Planning, funded by the 1965. The purpose of this paper is to report
World Bank, and contracted to the Sao Pau- our findings.
lo firm of consultants "Consorcio Nacional
de Engenheiros Consultores" (CENEC). As Methods and Materials
part of the National Museum of Natural
History's Research Training Program, The first recording of the Chapada dos
AMM USNM
analyzed the advertisement call of Guimaraes form is from Tape 320,
the Chapada dos Guimaraes form and com- Cut 2, Brazil, Mato Grosso, Chapada dos
pared it to the advertisement call of Eleu- Guimaraes, Estancia Monarca, recorded 30
therodactylusfenestratus from the Amazon September 1996, between 1835-1930 h, air
basin. The results indicate that the Chapada temperature 21.6°C, unvouchered, by
dos Guimaraes form is a different species WRH, one call recorded and analyzed. The
from the Amazonian E. fenestratus. To de- second recording of the Chapada dos Gui-
USNM
termine whether there is an available name maraes form is from Tape 320, Cut
for the Chapada dos Guimaraes form and 3, same data as Cut 2, except recorded at
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
USNM
1930 h, voucher 507899, one call Specimens examined are listed in Appen-
recorded and analyzed. The recordings for dix 1. Museum abbreviations follow Levi-
Eleutherodactylus fenestratus are from ton et al. (1985).
USNM
Tape 206, Cut 2, Peru, Madre de
Dios, Tambopata, recorded 6 January 1989,
Results
1850 h, air temperature 24.0°C, voucher
USNM —
342993, by Reginald B. Cocroft III, Advertisement calls. The ideal calls to
USNM
four calls analyzed, and Tape 266, compare the Chapada dos Guimaraes form
Cut 19, Peru, Madre de Dios, Tambopata, with are those from topotypic Eleuthero-
recorded 15 November 1990, 1815 h, air dactylus fenestratus males. Eleutherodac-
temperature 27.0°C, unvouchered, by Re-
tylus fenestratus (Steindachner 1864) was
ginald B. Cocroft III, six calls analyzed. described from specimens from Rio Ma-
Call recordings were digitized using
more, Rondonia and Borba, Amazonas,
Sound Image software at a sampling fre- Brazil. The geographically most proximate
quency of 22.05 KHz and 16-bit resolution. adequate recordings we know for Amazo-
The digitized calls were converted into au-
nian E. fenestratus are from Tambopata,
dio data files using a wave converter pack- We
Peru. use these recordings to represent
age and analyzed as audiospectrograms and
E. fenestratus for our comparisons.
wave forms using Computerized Speech
Research Environment software (AVAAZ As there is some variation among the
four individual frogs recorded from Peru
Innovations Inc., Version 4.2).
and Brazil, the data are summarized by in-
Eleven parameters were analyzed for
dividual caller (Table 1).
each call following the definitions of Heyer USNM
The frog from Tambopata, Tape
et al. (1990): call duration, call rate, notes
206, Cut 2, has a mean call duration of 0.31
per call, note duration, note repetition, note
sec; 2.75 mean notes per call; and a mean
pulsation, fundamental frequency, dominant
note duration of0.07 sec. The dominant fre-
frequency, change in dominant frequency,
peak frequency, and characterization ofhar- quency is the second harmonic; additional
harmonics are present, but weak. (Fig. la).
monics.
The second frog from Tambopata,
The audiospectrograms used in Fig. 1 USNM
were produced using Canary software (Cor- Tape 266, Cut 19, has a mean call
duration of 0.18 sec; and a mean note du-
nell Laboratory of Ornithology, Version
ration of 0.07 sec. The dominant frequency
1.2).
Color pattern and external morphological is the second harmonic; additional harmon-
data were recorded using a system of ics are present, but weak.
sketches to represent all distinctive patterns The single call of the frog from Chapada
USNM
or states. These data were then summarized dos Guimaraes, Tape 320, Cut 2,
and categorized into written character has a relatively short (0.02 sec) and weak
states. Measurements were taken with dial first note, the other 4 notes are stronger and
calipers and recorded to the nearest 0.1 mm have a duration range of 0.03-0.04 sec. The
following Heyer (1984). Adult males were notes are pulsatile. The dominant frequency
determined by presence of vocal slits. Adult is the second harmonic; additional harmon-
females were determined by presence of ics are present, but weak.
mature ova visible through the body wall. The single call of the second frog from
USNM
Unclear sex determinations based on exter- Chapada dos Guimaraes, Tape 320,
nal examination were verified by dissection. Cut 3, has a relatively short (0.02 sec) and
Multivariate discriminant function analyses weak first note, all other notes are stronger
were performed using SYSTAT 7.0 for with a duration of 0.04-0.05 sec. The notes
Windows. are pulsatile. The dominant frequency is the
I
VOLUME NUMBER
112, 1
10 n
N
U
O
2 -
a
a
10 n
B
8 -
»-i
fa
^ « ^ t
6-
4-
2-
—
o 0.2 0.4 0.6 0.8 1.0
Time in Seconds
USNM
Fig. 1. A. Advertisement call of Eleutherodactylusfenestratus, Tape 206, Cut 2. B. Advertisement
call of Chapada dos Guimaraes form, USNM Tape 320, Cut 3.
second harmonic; additional harmonics are and increased note repetition rate (Schnei-
&
present, but weak. (Fig. lb). der et al. 1984; Duellman Trueb 1986).
There is some variation between individ- The three degree difference of ambient tem-
ual male vocalizations from the same lo- perature between the two recordings of the
cality. The variation observed between the Peruvian specimens is in the predicted di-
recordings from Tambopata might be ac- rection for call and note duration and note
counted for by differences of temperature repetition rate. Note, however, that the re-
of calling males, at least in part. As a gen- cordings from Chapada dos Guimeraes
eral rule for frogs, increases in temperature were recorded at lower temperatures than
result in decreased call and note duration those of Peru, yet these Brazilian calls had
4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
—
Table L Advertisement call data for four Eleutherodactylus specimens from Tambopata, Peru and Chapada
dos Guimaraes, Brazil.
Geographic area Peru Peru Brazil Brazil
USNM Tape and Cut 206, 2 266, 19 320, 2 320, 3
Call duration (sec) 0.21-0.36 0.16-0.20 0.29 0.53
Number of notes/call 2-3 2 5 8
Note duration (sec) 0.06-0.09 0.05-0.08 0.04 0.05
(Mean) note repetition rate per sec 7.6 9.1 14.7 13.7
Fundamental frequency range 1636-2411 1464-2497 1894-2239 1808-2497
Peak fundamental frequency 1720 2067 2067 2153
Dominant frequency range 2842^392 2842^220 3014-^220 3273-4737
Peak dominant frequency 3531 3617 3692 3875
shorter note durations and higher note rep- stronger and the audiospectrogram indicates
etition rates than the Peruvian calls, indi- it is comprised of two notes at a dominant
cating that temperature differences do not broadcast frequency of 3300 Hz. The calls
explain the differences between the Brazil- from Rondonia, as expected, match the
ian and Peruvian calls. calls from Tambopata rather than the calls
The calls from Brazil and Peru do not from Chapada dos Guimaraes.
differ in terms of carrier frequencies; they The advertisement call evidence is con-
are broadcasting in the same frequency sistent with recognizing the Chapada dos
band. The calls differ markedly in terms of Guimaraes form as a species distinct from
temporal packaging (Table 1), such as num- Eleutherodacytl—usfenestratus.
ber of notes per call, individual note dura- Morphology. There are two available
tion, and note repetition rate. Penna (1997) names for Eleutherodactylus from Mato
among others, has experimentally demon- Grosso that potentially could apply to the
strated that the kinds of temporal differenc- Chapada dos Guimaraes form. There are
es found between the Tambopata and Cha- also a number of Eleutherodactylus fenes-
pada dos Guimaraes frogs are recognized as tratus-\\k& specimens from Mato Grosso.
species differences by frogs. The purpose of this section is to compare
Unfortunately, to date, there are no re- the Chapada dos Guimaraes form with
cordings of what we consider to be Ama- these specimens to determine whether the
zonian Eleutherodactylus fenestratus from Chapada dos Guimaraes form occurs more
the hylaea of northern Mato Grosso. In fact, broadly within the State of Mato Grosso
the only other published recordings for E. and to determine whether there is an avail-
fenestratus are from Amazonian Bolivia able name for it.
(Marquez et al. 1995) and are very similar The materials for comparison comprise
to the recordings from Tambopata. Ronald the following (Fig. 2 for Mato Grosso lo-
I. Crombie made three recordings in Ron- calities; Appendix 1 for specimen lists): the
donia, Brazil in which a single call of an E. 2 types of Telatrema heterodactylum Mi-
fenestratus is in the background. In two of randa-Ribeiro, 1937; the 3 type specimens
them (USNM Tape 55, Cut 5 and USNM of Eleutherodactylus crepitans Bokermann,
Tape 56, Cut 5, both from Alto Paraiso), 1965; 11 specimens from Chapada dos Gui-
the call sounds as though it consists of two maraes; 2 specimens from Fazenda Santa
notes, but can not be confirmed on an au- Edwiges; 1 specimen from Jacubim; 2 spec-
diospectrogram due to the relative weak- imens from Barra do Tapirapes; and speci-
ness of the call and strength of the back- mens considered to be E. fenestratus', 3
USNM
ground noise. The call on Tape 56, specimens from Sao Jose do Rio Claro; 8
Cut 8 from Santa Cruz da Serra is a bit specimens from Apiacas, 48 specimens
VOLUME NUMBER
112, 1
Fig. 2. Outline map of the State of Mato Grosso. See text for key to locality numbers. Open square is the
city of Cuiaba.
from Juruena (measurement data taken for adequate to conclude that the variation is
10 females and 1 male), 1 specimen from meaningful (e.g., the differences in loreal
Alto Juruena, 46 specimens from Aripuana region pattern between the Mato Grosso
(measurement data taken for 10 males and populations of Eleutherodactylus fenestra-
females), and 28 specimens from Tambo- tus (Table 3, samples 7-10) and the Peru-
pata and Pakitza, Madre de Dios, Peru (the vian sample of E. fenestratus. In other cas-
specimens from the nearby locaHty of Pak- es, the differences are suggestive, but any
itza were added to those from Tambopata conclusions are compromised by sample
to bring the sample sizes up to at least 10 sizes. For example, there are two aspects of
males and 10 females). light mid-dorsal stripes that are impacted by
Patterns: The dorsal snout patterns dem- small sample size. The first is that there is
onstrate as much intra- as inter-population variation in the presence condition of the
variation and are not discussed further. The stripe. It may either be narrow or broad, and
other patterns analyzed demonstrate some it may extend from the tip of the snout to
levels of interpopulation variation (Tables the vent or from the interocular area to the
2—8). In some cases, the sample sizes are vent. Because there were relatively few in-
6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
—
Table 2. Interorbital markings. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Almost
absent; B = Outlined only, not with expansion; C = Solid, not with expansion; D = Solid with single prominent
posterior expansion; E = Solid with bifid prominent posterior expansion. Data are number of individuals dem-
onstrating the various states (question mark indicates specimen faded, state uncertain).
C. dos E.fenestratus
Taxa E. heterodactylns E. crepitans Guimaraes
Locality 1 2 3 7 8 9 9a 10 Peru
A
5
B 2 12 9 2
C 11 3 6 9 1 11 9
D 6
E 6
From illustration 2a (Miranda-Ribeiro 1937).
dividuals with light mid-dorsal stripes, the ples, if the Chapada dos Guimaraes form
information had to be collapsed for com- has mid-dorsal stripes at those same fre-
parison. Second, the sample size for the quencies of occurrence, then the sample
Chapada dos Guimaraes form is too small size is not adequate to have found individ-
to conclude that it lacks light mid-dorsal uals with that condition.
stripes, even though all 11 specimens ex- Morphological features: All but three in-
amined lack them. There are three adequate dividuals scored for tarsal fold variation
samples of E. fenestratus to compare with had discernible tarsal folds (Table 9). It is
the Chapada dos Guimaraes form: Aripua- likely that the three individuals thus scored
na, Juruena, and Peru. For Aripuana, the actually have them but are preserved in
probability of an individual having a mid- such a manner that they are not discernible.
dorsal stripe is 0.35, thus the probability of The two individuals of Eleutherodactylus
not having a stripe is 0.65. If we use that crepitans with discernible tarsal folds have
probability to ask whether the Chapada dos a very different morphology from all other
Guimaraes population could have the same specimens examined (Table 9). The varia-
occurrence of stripes, the answer is no: tion in toe fringe condition is continuous
0.65" = P < 0.001 (exact probability test). (Table 10). The fringed condition is weakly
However, when the population data from fringed, with noticeable basal toe webbing
Juruena and Peru are used, the answer is (the fringe is not as well-developed as in
yes: for Juruena, the probability of not hav- some species of Leptodactylus, for exam-
ing a stripe is 0.91, for Peru 0.92, with P ple). The variation in tubercles on the sole
=
0.35 and 0.40 respectively. Thus, based of the foot (Table 11) suggests that E. cre-
on the low frequency of occurrence of mid- pitans is distinctive (in contrast to com-
dorsal stripes in the Juruena and Peru sam- pletely distinct) from the other samples, and
—
A =
Table 3. Loreal region patterns. Locality numbers refer to those in Fig. 2 (Locality 9a not mappable).
No pattern; B = Incomplete stripe in front of eye; C = Dark stripe from eye to nostril, entire loreal region may
be dark. Data are number of individuals exhibiting various states. Pattern not discernible for specimen from
locality 6.
c dos £.fenestratus
Taxa E. heterodacnlus E. crepitans Gui.maraes 1 9 9
Locality 1 2 3 4 5 6 7 8 9 9a 10 Peru
A
8
B 1 2 4 1 9 1 1 13
C 1 7 2 3 8 12 20 7
—
VOLUME NUMBER
112, 1 7
—
Table 4. Supratympanic fold patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A =
Fold not accentuated with dark brown band; B = Fold accentuated with interrupted dark brown band; C = Fold
accentuated with solid dark brown band. Data are number of individuals demonstrating the various states (ques-
tion mark indicates state uncertain due to fading or perspective of illustration).
C. dos E.fenestratus
Taxa E. heterodactxlus E. crepitans Guimaraes 7 7 7
Locality 1 2 3 4 5 6 7 8 9 9a 10 Peru
A 9a 2
B 3 2 1 12
C 9 2 7 3 8 21 1 21 14
From illustration 2a (Miranda-Ribeiro 1937).
the variation in the Chapada dos Guimaraes Taxonomy
specimen tubercles is distinctive from the
Telatrema heterodactylum Miranda-Ri-
variation observed in the E. fenestratus —
beiro, 1937. Alipio Miranda-Ribeiro de-
samples.
scribed this species from "Matto-Grosso
Measurements: Male and female sizes
Caceres, 2 exemplares colhidos na gruta
vary among the samples (Table 12). Two
dita 'Fazendinha' Comm. Rondon (1937:
points are worth noting, although additional
67)." His son Paulo Miranda-Ribeiro des-
samples are necessary to verify them statis- A
ignated specimen 106 in the collection of
tically. There is no sexual size dimorphism the Museu Nacional, Rio de Janeiro, as the
in the Eleutherodactylus crepitans sample,
lectotype (1955:411). The second speci-
whereas sexual size dimorphism is pro-
men, the paralectotype, which also had the
nounced in all other samples that contain number 106 originally, was subsequently
both sexes. The Chapada dos Guimaraes
recatalogued as 5089.
form is smaller than the Eleutherodactylus Dr. R E. Vanzolini (pers. comm.) was
fenestratus samples analyzed.
able to locate A. Miranda-Ribeiro's locality.
The results of multivariate discriminant Miranda-Ribeiro was a member of the Ron-
function analyses for males and females don Commission expedition and he collect-
(analyzed separately) agree in that the ed the specimens in question himself. Cac-
Eleutherodactylus crepitans specimens are eres, as used by Miranda-Ribeiro, refers to
clearly distinct from all other individuals the municipality, which at the time of the
analyzed (Fig. 3). The rest of the popula- Rondon Commission expedition was very
tions analyzed indicate that there is differ- extensive. Vanzolini found in the Rondon
entiation among them, but the degree of Commission itinerary that Miranda-Ribeiro
variation is consistent with both intra- and travelled to "Fazendinha" and returned to
inter-specific variation based on similar Caceres in the same day and that "Fazen-
studies in other groups of frogs. dinha" was near a place called Pirizal. Van-
—
Table 5. Mid-dorsal stripe patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Absent;
B = Present. Data are number of individuals demonstrating the states.
C. dos E.fenestratus
Taxa E. heterodactylus E. crepitans Guimaraes ? ?
Locality 1 2 3 4 5 7 8 9 9a 10 Peru
A
1 11 3 8 44 1 34 26
B 4 12 2
8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
—
Table 6. Dorsal patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Indistinct or
uniform; B = Dark symmetrical spots; C = Dark chevron in medial scapular region followed by dark blotches
posteriorly; D = Dark W-shaped mark in medial scapular region followed by dark blotches posteriorly; E =
Extended dark W-shaped pattern (additional strophes to next in series from inverted V to W) in medial scapular
region followed by dark blotches posteriorly. Data are number of individuals demonstrating the various states
(question mark indicates uncertain condition due to fading).
C. dos E.fenestratus
Taxa E. heienxlactxhis E. crepitans Guimaraes
Locality I ' 2 3 7 8 9 9a 10 Peru
A
1 3 7 1 9 13
B 1 3 4
C 4 2 2
D 2 9 6 15
E
From illustration 2a, b (Miranda-Ribeiro 1937).
zolini located a "Fazendinha" that not only randa-Ribeiro's measurement of 27); head
meets these conditions, but is in an area length 8.6; head width 8.5; eye-nostril dis-
known to have sandstone caves (grutas = tance 3.5; eye-eye distance 5.1; greatest
caves) at 16°00'S, 57°36'W (Fig. 2, Local- tympanum diameter 2.4; thigh 13.9; shank
ity 1). 14.1; foot 12.0; width of 3rd finger disk 1.4;
&
Lynch Duellman (1997:225) recog- width of 4th toe disk 0.7. The lectotype has
nized Eleutherodactylus heterodactylus slits in the floor of the mouth, but they ap-
(Miranda-Ribeiro, 1937) as a distinct spe- pear to be cuts, not vocal slits. The sex of
cies and included it in the Eleutherodacty- the lectotype can not be determined with
lus binotatus species-group. As Lynch and certainty without dissection, which given
Duellman did not examine the types, we of- the poor condition of the specimen was
fer the following observations. considered inappropriate.
Both types are faded such that most fea- The following support recognition of Te-
tures of any color patterns are no longer latrema heterodactylum as a distinct and
discernible. The lectotype is in poor con- valid species ofEleutherodactylus, confirm-
&
dition, the paralectotype is in worse condi- ing Lynch Duellmans' (1997) assess-
tion. The paralectotype is very brittle and ment.
fragile and disintegrates more each time it The disks on fingers 3 and 4 are very
is handled. The lectotype is the (noticeably) large, much larger than the disk on the
larger of the two, and the only specimen fourth toe, and the shape is ovate (Fig. 4a).
WRH
measured for purposes of this paper. The disks on fingers 3 and 4 of the other
recorded the measurements (in mm) of the Eleutherodactylus examined are expanded,
lectotype as: SVL 24.6 (contrasted to Mi- but either the same size or just larger than
—
A =
Table 7. Posterior face of thigh patterns. Locality numbers refer to those in Fig. 2 (9a not mappable).
Uniform; B = Finely mottled around vent, rest uniform; C = Finely mottled; D = Mottled. Data are number
of individuals demonstrating the various states.
C. dos Guima- E.fenestratus
Taxa E. heterodactylus E. crepitans raes
Locality 1 ' 2 3 7 8 9 9a 10 Peru
A
3 7 18 20 5
B 1 3 1 1 8
C 15
D
VOLUME NUMBER
112, 1 9
—
Table 8. Belly patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Almost immaculate;
B = Noticeably flecked or mottled. Data are number of individuals demonstrating the states. Specimen from
locality 6 too faded to score.
C. dos E.fenestratus
Taxa E. heterodactxlus E. crepitans Guimaraes I •7
Locality 1 2 3 4 5 6 7 8 9 9a 10 Peru
A 1 3 10 2 1 6 10 11 28
B 1 3 2 11 1 10
the disks on toe 4 and they are broadly tri- Eleutherodactylus heterodactylus (Miran-
angular in shape (Fig. 4b, c). da-Ribeiro 1937) is thus far known only
The type illustrations (most likely that of from the original two specimens collected
the lectotype, Miranda-Ribeiro 1937, Fig. by Miranda-Ribeiro from "Fazendinha."
2a-b) show a pair of ill-defined light lon- All other members of the Eleutherodactylus
&
gitudinal stripes from behind the eye to the hinotatus group, to which Lynch Duell-
sacrum. This feature is not seen in any of man (1997) assigned E. heterodactylus, oc-
the other Eleutherodactylus examined for cur in eastern Brazil, primarily in the At-
this study. lantic Forest Morphoclimatic Domain. In-
The type locality is characterized by a clusion of E. heterodactylus in this group
very open vegetation formation lacking gal- does not make zoogeographic sense.
lery forests. Eleutherodactylus fenestratus Status of Eleut—herodactylus crepitans
is a forest denizen and the Chapada dos Bokermann, 1965. Werner C. A. Boker-
Guimaraes form inhabits well-developed mann described the species based on three
gallery forests. Eleutherodactylus crepitans specimens from Sao Vicente, Mato Grosso.
also occurs in open habitats, but is morpho- The holotype and allotype are now in the
logically very distinct from E. heterodac- collections of the Museu de Zoologia,
tylus. Universidade de Sao Paulo and the other
One other morphological feature will re- paratype is in the Museu Nacional, Rio de
quire fresh material to evaluate. There is no Janeiro.
indication of any tarsal fold or other tarsal John Lynch (1980:8) synonymized
decoration on either the lectotype or para- Eleutherodactylus crepitans Bokermann,
lectotype, suggesting that, if in fact the tar- 1965 with E. fenestratus (Steindachner
sus lacks a fold, that would be another fea- 1864). Lynch did not examine the speci-
ture distinguishing the species from all oth- mens of E. crepitans (1980:6). Lynch ar-
er Mato Grosso Eleutherodactylus. gued that because Bokermann was unaware
—
Table 9. Tarsal fold states. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Absent; B =
Short, oblique, lying more than length of inner metatarsal tubercle from inner metatarsal tubercle; C = Mod-
erately short, less than Vi length oftarsus, lying less than length ofinner metatarsal tubercle from innermetatarsal
tubercle; D = Long, greater than Vi length of tarsus, lying less than length of inner metatarsal tubercle from
inner metatarsal tubercle. Data are number of individuals demonstrating the various states. Data not taken for
one desiccated individual from locality 10.
C. dos E.fenestratus
Taxa £. heterodactxlus E. crepitans Guimaraes
Legality 1 2 3 7 8 9 9a 10 Peru
A
1 1 1
B 2
C 10 3 8 21 1 19 28
D
1
10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
—
Table 10. Toe fringe states. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Toes free;
B = Toes with lateral ridges; C = Toes with modest lateral fringes. Data are numberofindividuals demonstrating
the states.
c dos E fenestratus
E. hetenxiactxlus E. crepitans Gui.maraes 7 7 .
Locality 1 2 3 4 5 6 7 8 9 9a 10 Peru
A
1 3 I 5 5
B 3 3 8 6 17
C 8 2 1 2 3 5 12 1 10 6
that E. fenestratus was a valid species, Bok- of E. fenestratus and the Chapada dos Gui-
ermann compared his new species with in- maraes form (Fig. 3).
appropriate species from Ecuador, from Bokermann (1965:264) made a point of
which they were very distinct. However, stating that the specimens of Eleutherodac-
had Lynch examined the type material ofE. tylus crepitans were obtained in grass-shrub
crepitans, he would have found that they vegetation in the middle of large blocks of
differ from E. fenestratus as well. granite from such a dry habitat that he
The following argue for recognizing would never have expected to find frogs
Eleutherodactylus crepitans as a valid spe- there. The immediate vicinity of Sao Vi-
cies. cente does have a granitic intrusion of rath-
Eleutherodactylus crepitans has short, er small extent and is the only such granitic
oblique tarsal folds, separated from the in- habitat in Mato Grosso (Brasil 1982). It
ner metatarsal tubercle by a distance greater would not be surprising if E. crepitans is
than the length of the fold. Eleutherodac- limited to this distinctive habitat.
tylus heterodactylus apparently lacks tarsal Bokermann (1965:264) indicated that
folds. Eleutherodactylusfenestratus and the males called at night from sandy hiding
Chapada dos Guimaraes form have longer places among rocks. He described the call
(but still relatively short) tarsal folds, par- as consisting of two notes in sequence, like
allel to the long axis ofthe tarsus, separated a short and coarse click, repeated many
by a short distance (less than length of the times but irregularly (translation by P. E.
fold) from the inner metatarsal tubercle. VanzoUni). The two note call of Eleuther-
Eleutherodactylus crepitans does not odactylus crepitans is more similar to
have marked sexual dimorphism in size. Eleutherodactylusfenestratus than the Cha-
Both Eleutherodactylusfenestratus and the pada dos Guimaraes form, but the frequent
Chapada dos Guimaraes form show marked calling of E. crepitans is different from the
sexual size dimorphism. very infrequent calls of both E. fenestratus
The overall body shape of Eleutheroda- and the Chapada dos Guimaraes form.
—
cytlus crepitans differs markedly from that Questionable status specimens. The
—
Table 11. Maximum number of tubercles on sole of foot. Locality numbers refer to those in Fig. 2 (9a not
mappable). Data are number of individuals demonstrating the states.
C. dos E.fenestratus
Taxa E. heterodactylus E. crepitans Guimaraes
Locality 1 2 3 7 8 9 9a 10 Peru
1
5 9 9 23
1 2 9 9 5
2 1 3 1 2