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REVIORA 



jyi niseiim oi Lomparafive Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 



24 July 2008 



Number 513 



DISTINGUISHING FEATURES OF THE SUB-SAHARAN FROG GENERA 

ARTHROLEPTIS AND PHRYNOBATRACHUS: A SHORT GUIDE FOR FIELD 

AND MUSEUM RESEARCHERS 

Breda M. Zimkus 1 * and David C. Blackburn 1 

Abstract. Typically small body size and similar external anatomy have made it difficult for researchers to 
distinguish between species of two distantly related, but often sympatric, African ranoid frog genera: Arthroleptis 
(Arthroleptidae) and Phrynobatrachus (Phrynobatrachidae). We define a suite of external morphological characters, 
extending beyond the traditionally used secondary sexual characteristics, that can be used to definitively distinguish 
between adult Arthroleptis and Phrynobatrachus. Photographs comparing absence, presence, and variation of 
morphological characters are included for clarification. Significant differences between body proportions (head 
width/snout-vent length) are also observed between these genera. Although smaller species might be more difficult to 
identify, larger species (> 30 mm snout-vent length) oi Arthroleptis can be distinguished from Phrynobatrachus 
because of their relatively wider heads. We intend this to serve as a heuristic guide for both field- and museum-based 
researchers. 

Key words: Africa; Amphibia; Arthroleptidae: identification: Ranoidea: Phrynobatrachidae 



INTRODUCTION 

Nearly every herpetologist is familiar with 
the overwhelming diversity of "little brown 
frogs." Many leaf litter anurans are small, 
generally drab in color, and morphologically 
similar, and this has hampered both system- 



1 Museum of Comparative Zoology, and Department of 
Organismic and Evolutionary Biology. Harvard Univer- 
sity, Cambridge. Massachusetts 02138. U.S.A.: 
*Corresponding author. E-mail: bzimkus<Yz oeb. harvard, 
edu. 



atic research and identification in the field 
and in museum collections. Within sub- 
Saharan Africa, two such genera, Arthrolep- 
tis Smith 1849 and Phrynobatrachus Giinther 
1862, have been confused for more than a 
century (i.e., Boulenger, 1882). On the basis 
of studies of skeletal morphology (e.g.. 
Laurent, 1940, 1941a. b; Scott, 2005) and 
the results of molecular phylogenetic analy- 
ses (e.g., Bossuyt et at.. 2006; Frost et ah. 
2006; Roelants et a/., 2007). it is now very 
clear that these genera, although externally 
similar, are only distantly related. In our 



BREVIORA 



No. 513 



experience, many researchers are unaware of 
the numerous external morphological char- 
acters that can be used to distinguish 
between each genus. Arthroleptis and 
Phrynobatrachus continue to be confused 
in the field and in museum collections, 
which can then confound further research 
such as molecular phylogenetic analyses 
(see Discussion for an example). Thus the 
external morphological differences between 
these two genera warrant a thorough treat- 
ment to improve the efficacy of identifica- 
tion. 

Squeakers (Arthroleptis sensu Frost, 2007) 
are generally small (16-54 mm snout-vent 
length), terrestrial, leaf litter anurans. and all 
of the approximately 37 species are believed 
to have lost the free-living, feeding tadpole 
stage (i.e.. direct development). The smallest 
Arthroleptis species were previously placed in 
Schoutedenella (Laurent. 1954): Frost et al. 
(2006) synonymized Schoutedenella with Ar- 
throleptis on the basis of the results of their 
molecular phylogenetic analysis. An analysis 
of the relationships within Arthroleptis and 
its putative sister taxon Cardioglossa is 
currently in preparation (Blackburn, unpub- 
lished). 

Puddle frogs (Phrynobatrachus) comprise a 
lineage of approximately 75 species found in 
diverse terrestrial habitats across sub-Sa- 
haran Africa (Frost, 2007). Phrynobatrachus 
has been a long-standing source of confusion 
to systematists because of extensive geo- 
graphic and intrapopulation variation ac- 
companied by only slight morphological 
differences among species (Largen, 2001; 
Rodel, 2000; Stewart, 1974). Molecular data 
from mitochondrial and nuclear markers 
indicate that both Dimorphognathus and 
Phrynodon might be embedded within Phry- 
nobatrachus, thus rendering the genus para- 
phyletic (Frost et a!., 2006; Scott, 2005). 
Both Scott (2005) and Frost et al. (2006) 
argued for synonymizing these genera with 



Phrynobatrachus. Whereas Dimorphognathus 
africanus is embedded deeply within Phryno- 
batrachus. supporting the synonymization of 
this monotypic genus, the placement of 
Phrynodon sandersoni within Phrynobatra- 
chus is more controversial because of its 
basal position within the lineage; the synon- 
ymy of Phrynodon with Phrynobatrachus is 
currently being examined as part of a 
phylogenetic study of the Phrynobatrachidae 
(sensu Frost, 2007; Zimkus. unpublished). 

The long taxonomic history linking Ar- 
throleptis and Phrynobatrachus contributes 
to the present difficulties resolving the 
identifications of these distantly related frog 
genera (e.g., Boulenger, 1882). Many species 
originally described as Arthroleptis have been 
since transferred to other genera, including 
21 species that are currently considered valid 
species of Phrynobatrachus (Frost, 2007). In 
part on the basis of the work of Deckert 
(1938), Laurent (1940, 1941a. b) used oste- 
ological characters to separate species into 
two genera, Arthroleptis and Phrynobatra- 
chus. which he believed were not closely 
related. The morphological differentiation of 
these genera is further supported by recent 
molecular phylogenetic studies (e.g., Bossuyt 
et al. , 2006; Frost et al. . 2006; Roelants et al. . 
2007), which show that Arthroleptis and 
Phrynobatrachus belong to different radia- 
tions within ranoid frogs. 

The purpose of this study is to clearly 
determine those external morphological 
characters that can be used to discrimi- 
nate between Arthroleptis and Phrynobatra- 
chus. We also document characters that 
could be useful for identification but vary 
within these genera. Because we have found 
that written descriptions of these characters 
are often unclear, we provide photographs to 
illustrate the more important characters. 
Although this information might not be 
entirely novel for tenured African amphibian 
biologists, we hope that those not familiar 



2008 



DIAGNOSING \RT1I ROl.l.PIIS \\l)/7/A>) \OUATRACHUS 



wilh these genera will find this to be a useful 
guide. 

METHODOLOGY 

Using the taxonomic, systematic, and 

fauna! literature as our guide, we examined 
museum specimens for characteristics report- 
ed!) useful in differentiating between Arthro- 
leptis and Phrynobatrachus. We focused 
solely on external characteristics to make 
our observations equally useful to research- 
ers conducting field surveys or sorting 
specimens in museum collections. A total of 
15 morphological characters were investigat- 
ed; six of these are male secondary sexual 
characters. Characters are provided in Ap- 
pendix 1. and Appendix 2 comprises a data 
matrix in which all species examined are 
coded for these characters. 

The results of this work are based on our 
study of specimens from museum collections, 
as well as our respective experiences in the 
field. Museum abbreviations correspond to 
those of Leviton et al. (1985). with the 
exception of MMB (Museums of Malawi. 
Blantyre). The allocation of species to either 
Anhroleptis or Phrynobatrachus follows 
Frost (2007). High-resolution images were 
obtained with a JVC 3-CCD digital camera 
using AutoMontage Pro 5.0 (Synoptics). 
Images were saved as TIFF files, cropped, 
and contrast-adjusted with Adobe Photo- 
shop 7.0 for Macintosh. We gathered meris- 
tic data from 280 adult specimens represent- 
ing 27 species of Anhroleptis and 246 adult 
specimens representing 31 species of Phryno- 
batrachus (Online Supplement: Appendix 1). 

Anhroleptis and Phrynobatrachus appear 
to generally exhibit different body shapes, with 
Anhroleptis having relatively wider heads. To 
test whether this difference is significant, we 
took measurements (± 0.1 mm) of snout- 
vent length (SVL) and head width (HdWd) 
with digital calipers (Online Supplement: 



Appendix 1). We used an analysis of covari- 

ance to test for significant difference in the 
slope o\~ the regression lines in which HdWd 
is the dependent variable and SVL is indepen- 
dent. To take into account variation in both 
HdWd and SVL. reduced major axis regres- 
sion was used. Statistical significance was 
evaluated for oe = 0.05. 

RESULTS 

The characters examined and evaluated in 
this study are summarized in Table 1. Nearly 
all characters reported to differentiate Ar- 
thro/eptis and Phrynobatrachus were found 
to vary both within and between species in 
each genus (Table 1). Only two external 
morphological characters definitively differ- 
entiate these genera: 

1 . the presence of a tarsal tubercle and 

2. the presence of an outer metatarsal 
tubercle. 

However, among African anurans. only the 
presence of a tarsal tubercle is unique to 
Phrynobatrachus. 

In general. Anhroleptis have relatively 
wider heads than Phrynobatrachus. The 
coefficients (P) of linear regression of HdWd 
on SVL are significantly different between 
these two genera {Anhroleptis: P = 0.461. SE 
= 0.006: Phrynobatrachus: p = 0.348. SE = 
0.005: P < 0.0001, F = 64.39). Although 
highly significant, in practice it is very 
difficult to discriminate between these genera 
on the basis only of these data if specimens 
are less than 30 mm SVL (Fig. 1). The 
difference in relative head width is only 
obvious for specimens more than 30 mm 
SVL (HdWd/SVL: 40% vs. 34%). 

DISCUSSION 

In this study, we examined a suite of 
morphological characters believed to be 
useful in distinguishing between adults of 
Anhroleptis and Phrynobatrachus. The most 



BREVIORA 



No. 513 



Table 1. External morphological characters useful for differentiating between Arthroleptis 

AND PhRYSOBATRACHCS. 



Character 



Arthroleptis Phrynobatrachus 



References 



Tarsal tubercle 



absent 



Hourglass or triple diadem pattern on 

dorsum 
Chevron-shaped glands in scapular region 
Eyelid cornicle or spur 
Sexually mature males 

Elongate third finger 

Dermal digital spines 

Inguinal spines 

Nuptial excrescence (thickened pad of 
skin) on first finger 

Femoral gland 

Lateral vocal folds 



variable 



present 



Outer metatarsal tubercle 


absent 


present 


Heel tubercle (located on the proximal 


absent 


variable 


end of the tarsus) 






Circummarginal groove on terminal 


absent 


variable 


phalanx 






Pedal webbing 


absent 


variable 


Median dorsal skin raphe 


present* 


absent 



absent 



absent 


variable 


absent 


variable 


variable 


absent 


variable 


absent 


variable 


absent 


absent 


variable 


absent 


variable 


absent 


variable 



Blackburn. 2005; Channing and 
Boycott. 1989; Drewes and Perret. 
2000; Scott. 2005 

Deckert. 1938 

Scott. 2005 

Blackburn. 2005; Scott. 2005 

Schmidt and Inger. 1959; Stewart. 1967 
Drewes and Perret, 2000; Laurent, 

1957; Scott. 2005 
Scott. 2005 

Scott, 2005; Stewart. 1974 
Perret, 1988; Rodel. 2000 

Blackburn, 2005; Noble. 1931 
Noble. 1931; Schmidt and Inger. 1959 
Blackburn, unpublished 
Scott. 2005; Stewart, 1967 

Blackburn, 2005; Parker. 1935: 

Stewart. 1967 
Stewart, 1967 



*Variation is the result of preservation, desiccation, or both. 



definitive characteristic that can be used to 
differentiate between these two genera is the 
presence of both an outer metatarsal tubercle 
and a tarsal tubercle in Phrynobatrachus 
(including the previously synonymized Di- 
morphognathus and Phrynodon) (Fig. 2B). 
Arthroleptis (including Schoutedenella) only 
exhibits an inner metatarsal tubercle, which 
is also found in Phrynobatrachus. Although 
Drewes and Perret (2000) found Phrynoba- 
trachus (Dimorphognathus) africanus and 
Phrynobatrachus (Phrynodon) sandersoni to 
lack a tarsal tubercle, we found the tarsal 
tubercle present in both taxa, albeit rather 
reduced in size in the latter species. 

Numerous morphological characters ex- 
amined in this study are present in other 



° Arthroleptis 

° Phrynobatrachus 




SVL(mm) 

Figure 1. Head width (HdWd) plotted against 
snout-vent length (SVL) for Arthroleptis (squares) and 
Phrynobatrachus (circles). 



2008 



DIAGNOSING ARTHROLEPTIS AND PHRYNOBATRACHUS 



African frogs, but the presence or combina- 
tion oi' particular characters can be useful in 
distinguishing between adult Phrynobatra- 
chus and Arthroleptis, Expansion o\~ the 
terminal phalanx o\' manual and pedal digits 
is found in both genera, but circummarginal 
grooves are found only in some species of 
Phrynobatrachus (Blackburn. 2005; Scott, 
2005; Fig. 2E). Phrynobatrachus differs from 
Arthroleptis by the presence of pedal web- 
bing in many species of the former (Schmidt 
and Inger. 1959). Webbing ranges from 
absent or rudimentary in numerous smaller 
species such as P. miliums, P. parvulus, and 
P. ukingensis to extensive in the majority of 
larger, rheophilic species such as P. acutiros- 
tris, P. krefftii, and P. versicolor (Fig. 2C, 
D). Five species of Phrynobatrachus, includ- 
ing P. annulatus, P. calcaratus, P. cornutus, 
P. taiensis, and P. villiersi, possess a single 
spinelike dermal tubercle on the upper eyelid 
(Rodel, 2000). a character that can be used to 
distinguish Phrynobatrachus from Arthrolep- 
tis (Perret. 1988), as well as other African 
anurans. 

A number of external morphological 
characters vary both within and between 
species in each genus, and this variation is 
due to either natural variation within or 
between populations or the condition of 
preserved specimens. Chevron-shaped glands 
in the scapular region of Phrynobatrachus 
and a dorsal triple diadem pattern in 
Arthroleptis can both be used to identify 
species of these genera (Scott, 2005; Stewart, 
1974). However, these are not always prom- 
inent in all members of a species or may be 
completely lacking in some species. The size 
and shape of the chevron-shaped glands of 
Phrynobatrachus are quite variable and can 
originate and terminate in the scapular 
region or instead extend almost the entire 
length of the body (Fig. 2H). The median 
dorsal skin raphe of Arthroleptis can be a 
difficult character to visualize, especially in 



poorly preserved specimens (Fig. 2G); how- 
ever, the presence of this character can be 
used to accurately identify Arthroleptis. 
Finally, some species of Phrynobatrachus 
exhibit circummarginal grooves on the man- 
ual or pedal digit tips; in some species, these 
furrows are found only on the longest digits. 
In addition, the presence of circummarginal 
grooves could be difficult to determine 
because of desiccation of preserved speci- 
mens. However, if circummarginal grooves 
are present, this character can be used to 
differentiate Phrynobatrachus from Arthro- 
leptis, because it is lacking in the latter, but 
not necessarily from other African frog 
genera (Blackburn, 2005; Scott, 2005; 
Fig. 2E). 

The presence of male secondary sexual 
characters can lead to the most straightfor- 
ward identifications of specimens as Arthro- 
leptis or Phrynobatrachus. An elongate third 
finger is found in many male Arthroleptis, as 
well as most species of Cardioglossa, the 
putative sister genus of Arthroleptis (Black- 
burn, in press; Fig. 3A, B). This sexual 
dimorphism is not found in Phrynobatrachus 
or any other anurans. Males of nearly all 
Arthroleptis species have dermal spines lining 
the medial surface of the elongate third 
finger and are also sometimes found on the 
lateral, medial, or both surfaces of the 
second finger (Fig. 3B). In some species, 
the presence of spines can vary seasonally 
(e.g., Schmidt and Inger, 1959). Males of 
some species of Phrynobatrachus possess a 
nuptial excrescence or thickened pad of skin 
on the medial and dorsal surface of the first 
finger, which was hypothesized by Parker 
(1940) to be an adaptation to aquatic 
amplexus. In P. africanus, this pad is greatly 
hypertrophied and covers much of the dorsal 
surface of the hand (Fig. 3C). The presence 
of an elongate ovoid femoral gland, which is 
most evident in life because it often can be 
hardened and yellow in color, allows the 



BREVIORA 



No. 513 




&';'- ? "J' 


- 


c — 


D 








Figure 2. Morphological characters useful in distinguishing between Arthroleptis and Phrynobatrachus. A, Left 
foot of A. stenodactylus (MCZ A-137060) in ventral view illustrating inner metatarsal tubercle and lack of webbing. 
B, Left foot of P. natalensis (MCZ A-1380S4) in ventral view illustrating inner metatarsal tubercle (a), outer 
metatarsal tubercle (b), and tarsal tubercle (c). C, Left foot of P. parvulus (MCZ A-137121) in ventral view 
illustrating absent or rudimentary webbing. D, Right foot of P. steindachneri (MCZ A- 136907) in ventral view 



:ik)n 



DIAGNOSING [RTHROLEPTIS AND PHRYNOBATRACHl S 



identification of breeding Phrynobatrachus 
males (Blackburn, 2005; Stewart. 1967). 
However, in most species, this gland might 
be difficult to identify after preservation. 
Breeding Phrynobatrachus males have a 
single SLibgular vocal sac, which, when not 
distended, forms one or multiple folds, 
roughly parallel to the lower jaw, on the 
lateral margins of the throat (Stewart, 1967; 
Fig. 3E). Although the gular region of 
Arthroleptis can be distended and wrinkled, 
prominent gular folds are not present 
(Fig. 3D). 

For many field biologists that regularly 
collect Arthroleptis and Phrynobatrachus, it is 
obvious that the body proportions of these 
frogs generally differ. Indeed, we find that 
Arthroleptis have heads that are relatively 
wider than Phrynobatrachus. Meristic data 
collected in this study illustrate that the 
linear regression coefficients differ signifi- 
cantly between these two genera. However, 
in general, the difference in body proportions 
is only obvious in large specimens (> 30 mm 
SVL). Thus, at least for larger specimens, the 
relative width of the head could be a useful 
quick diagnostic feature, especially if used in 
concert with other morphological characters 
discussed herein (Table 1). 

It is also important to note that numerous 
nonmorphological characters such as call 
structure, breeding biology, and habitat 
preference can also be used in the field to 
assist in the identification of Arthroleptis and 
Phrynobatrachus. All Arthroleptis are be- 
lieved to have direct development, in which 



froglets hatch from a small clutch of 
terrestrially deposited eggs. In contrast, most 
Phrynobatrachus species deposit hundreds to 
thousands of eggs in ponds, streams, or 
pools, and a small number of species deposit 
small clutches of eggs in stagnant water 
found in tree holes, in empty fruit capsules, 
within snail shells, or terrestrially (Rodel, 
1998; Rodel and Ernst, 2002). Species 
exhibiting these alternative reproductive 
modes include P. dendrobates, P. guineensis. 
P. krefftii, P. phyllophilus, P. sandersoni, and 
P. tokba, although all have free-living 
tadpoles (Amiet, 1981; Rodel, 1998; Rodel 
and Ernst, 2002). Although few advertise- 
ment calls of these genera are published, 
those currently available will undoubtedly 
assist in distinguishing Phrynobatrachus from 
Arthroleptis (e.g., Drewes and Perret, 2000; 
Rodel, 2000; Schiotz, 1964). 

Although we demonstrate in this study 
that it is possible to distinguish adult 
Phrynobatrachus and Arthroleptis with the 
use of morphological characters, the identi- 
fication of juvenile and subadult Arthroleptis 
and Phrynobatrachus continues to be chal- 
lenging, even occasionally for the authors of 
this study. Fortunately, the identification of 
numerous taxa has been greatly facilitated by 
the recent use of DNA barcoding (e.g., 
Hebert et a/., 2003; Moritz and Cicero, 
2004; Vences et a!., 2005). The use of the 
mitochondrial 16S ribosomal RNA (rRNA) 
gene has proven to be particularly successful 
in amplification of amphibian DNA (Vences 
et al, 2005). We have found that the 



illustrating extensive webbing. E, Tips of left pedal digits III and IV of P. tokba (MCZ A-26905) illustrating 
expansion of digit tip and circummarginal grooves. F, Dorsal surface of A. stenodactylus (MCZ A- 137060) 
illustrating the hourglass (i.e.. triple diadem) pattern. G. Dorsal surface of A. sp. nov. (MCZ A-137978), in life, 
illustrating the median dorsal skin raphe. H, Dorsal surface of P. steindachneri (MCZ A- 136907) on left and P. 
auritus (MCZ A-138095) on right illustrating variability in chevron-shaped glands. Scale bar 1 mm in 2A-D, F-H; 
0.1 mm in 2E. 



BREVIORA 



So. 513 




Figure 3. Secondary sexual characters of males useful in distinguishing between Arthroleptis and Phrynoba- 
trachus. A. Right hand of A. stenodactylus (MCZ A-137060) in ventral view showing elongate digit III. B. Fingers II 
and III of male A. schubotzi (CAS 201717) in ventral view illustrating dermal spines. C. Right hand of male P. 
natalensis (MCZ A-138084). above, and left hand of male P. (Dimorphognathas) afrkanus (MCZ A-136757). below. 
in dorsal view illustrating nuptial excrescence (pad) on digit I. D. Throat of male A. schuborzHCAS 201717). left, and 
A. stenodaaylus (MCZ A-137061), right, illustrating lack of vocal folds. E. Throat of male P. auritus (MCZ A- 
138095) and P. (Phrynodon) sanderso?iH\\CZ A- 136790) illustrating visible vocal folds. Scale bar 1 mm. 



2IMIS 



DI.\ti\()M\ii IRTIIROLEPIIS AND PHRYNOBATRACHl V 



amplification of this gene is effective in 
identifying tadpoles, juveniles, and subadults 

of numerous African ranoid frogs, including 
Phrynobatrachus and Arthroleptis. However, 

molecular data and analyses can be con- 
founded b\ the misidentificatioD of voucher 
specimens. For instance, nucleotide BLAST 
searches against GenBank sequence data for 
the Arthroleptis specimens for which Yences 
et al (2003) amplified the 16S rRNA gene 
reveals that these specimens are misidentified 
(GenBank sequences AF21 5 139^10). Al- 
though Vences et al. (2003) attribute the 
unusual paraphyly of Arthroleptis 16S genes 
to uncertainties in their sequence data, this 
result is easily explained: These sequences 
have high similarity to Phrynobatrachus and 
thus are misidentified at the genus level. We 
advocate DNA identification of specimens 
only to the level of genus because of the 
difficulty in accurately identifying species of 
either genus. Thorough taxonomic study in 
combination with molecular data will be 
necessary before DNA identification can be 
used confidently to identify species of these 
two genera. 

ACKNOWLEDGMENTS 

This study was supported by the Depart- 
ment of Organismic and Evolutionary Biol- 
ogy and Museum of Comparative Zoology 
(Harvard Lniversity) and NSF grant EF- 
0334939 (AmphibiaTree) to J. Hanken. 
Many thanks to the following individuals 
who made specimens under their care avail- 
able for study: M. Burger. J. Campbell. D. 
Cannatella. B. Clarke. R. Drewes. R. Gun- 
ther. V. Gvodzik. T. Harstell. T. LaDuc. L. 
Lawson. S. Loader. P. Malonza. L. Mazi- 
buko. J. McGuire. J. Measey. A. Ohler. A. 
Resetar. J. Rosado. A. Schmitz. E. Scott. J. 
Simmons. L. Trueb. J. Vindum, D. Wake. A. 
Wynn. J. Hanken provided useful comments 
on a draft of the manuscript. 



APPENDIX 1. MORPHOLOGICAL 
CHARACTERS EXAMINED. 

External Morphology 

1. Tarsal tubercle: (0) absent: ( 1 ) present. 

2. Outer metatarsal tubercle: (0) absent: ( 1 ) 
present. 

3. Heel tubercle (located at the proximal 
end of the tarsus): (0) absent: ( 1 ) present. 

4. Circummarginal groove at manual and/ 
or pedal digit tips: (0) absent: (1) present. In 
some species, this groove is present only on 
the tips of the longest digits. Variation in 
museum specimens might be the result of 
preservation, desiccation, or both. 

5. Pedal webbing: (0) absent or rudimen- 
tary with more than two distalmost phalan- 
ges of digit IV unwebbed: ( 1 ) moderate to 
extensive with no more than the two distal- 
most phalanges of digit IV unwebbed. 

6. Median dorsal skin raphe: (0) absent: 
( 1 ) present. 

7. Hourglass or triple diadem pattern on 
dorsum: (0) absent: (1) present. 

8. Chevron-shaped glands in scapular 
region: (0) absent: (1) present. These ridges 
of skin can be short or can extend the entire 
length of the body when present. 

9. Eyelid cornicle or spur: (0) absent: (1) 
present. 

Secondary Sexual Characteristics 

10. Sexually mature males, third finger 
relatively longer than in females: (0) absent: 
(1) present. 

11. Sexually mature males, dermal spines 
on fingers: (0) absent: ( 1 ) present. 

12. Sexually mature males, inguinal spines: 
(0) absent: ( 1 ) present. 

13. Sexually mature males, nuptial excres- 
cence (thickened pad of skin that can appear 
velvety) on first finger: (0) absent: ( 1 ) present. 

14. Sexually mature males, femoral glands: 
(0) absent: (1) present. Most often the 



10 



BREVIOR-4 



\o. 513 



femoral glands of Phrynobatrachus are lo- 

. on the posterior thigh. However. 

glands can be situated closer to either the 

_ s or vent. These glands are most easily 

seen in living specimens because they are 

usually bright yellow. 



15. Sexually mature males, lateral vocal 
folds: (0) absent: (1) present. When pres- 
ent, folds run roughly parallel to the 
lower jaw. at the lateral margins of 
the throat, and form one or multiple 
crease>. 



APPENDIX 2. 
DISTRIBUTION OF MORPHOLOGICAL CHARACTERS AMONG SPECIES OF 
ARTHROLEPTIS AND PHRY\OBATH4CHlS. ALL CHARACTER STATES ARE 
BINARY (0, 1). MISSING DATA ARE CODED AS "?"; 0.1 DENOTES POLYMOR- 
PHISM. REFER TO APPENDIX 1 FOR CHARACTER DESCRIPTIONS. 





1 


2 


3 


4 


5 6 


- 


v 


9 


: 


11 


12 


13 


14 


15 


Arthroleptis adelphus 
















0.1 






















A. adolfifriederici 








































:>iis 
















0.1 
























A. brevipes 








































A. crusculwn 
















0.1 























:cei 
















0.1 























A. hematogaster 








































A. Iameerei 
















0.1 
























A. nikeae 








































A. nimbaensis 
















0.1 























A. "poecilonotus" 
















0.1 























A. pyrrhoscelis 










































A. reichei 
















0.1 























A. schubotzi 
















0.1 























wdactylus 
















0.1 























A. sylvaricus 
















0.1 










0.1 













A. taeniatus 








































A. icmneri 








































A. Troglodytes 
















0.1 























A. tuberosus 
















0.1 























A. variabilis 
















0.1 























A. wahlbergii 
















0.1 























A. xenochirus 
















0.1 























A. xenodactyloides 
















0.1 























A. xenodaciylus 








































A. zimmeri 














1 ° 


9 








9 


■1 


9 











Anhroleptis sp. now 














1 1 











9 


9 


9 











Phrynobatrachus acridoides 






0.1 




1 



















1 


1 


1 


P. acutirostris 






1 




1 



















1 





1 


P. africanus 






0.1 
























1 





1 


P. auritus 






1 




1 






















1 


1 


P. batesii 





































9 


P. bequaerti 
































1 








P. bullans 






1 





1 



















1 


1 


1 


P. calcaratus 






0.1 


1 










1 

















1 



2008 



DIAGNOSING ART/IROLEPTIS AND PHRYSOBATRACHIS 



APPENDIX 2. Continued. 



1 


"> ^ 


4 


5 


6 




g 


9 


10 


11 


12 


1? 


14 15 


P. comutus 1 


! 


1 


• 1 










1 











9 


? ? 


P. cricogaster 1 


1 1 


1 


1 






















1 





P. cryptotis 1 


1 0.1 































1 


P. dendrobates 1 


I 


1 

























1 


1 


P. francisci 1 


1 





1 






















9 


9 


P. inexpectatus 1 


1 














0.1 

















1 1 


P. keniensis 1 


1 




























1 


1 


P. kinangopensis 1 







1 






















1 


1 


P. Art://?// 1 


1 


1 


1 






















1 


1 1 


P. nnibabiensis 1 


1 0.1 




























9 


o 9 


P. manengoubensis 1 


1 


1 




























1 1 


P. minimis 1 


1 




























1 


9 9 


P. natalensis 1 


1 0.1 





1 








0.1 














1 


1 1 


P. pakenhami 1 


1 


1 


1 








0.1 

















1 1 


P. parvulus I 


1 




























1 


1 1 


P. plicatus 1 


1 


1 


1 

























1 


P. rungwensis 1 


1 1 


1 



























1 1 


P. sandersoni 1 


1 1 


1 


1 
























1 1 


P. scheffleri 1 


1 1 




























9 


o 9 


P. sieindachneri I 


1 0.1 


1 


1 
























1 


P. stewartae 1 


1 ' 





1 
























1 1 


P. ukingensis 1 


1 0.1 


1 











0.1 
















1 1 


P. versicolor 1 


1 


1 


1 


o 


o 


1 











o 




1 



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