Breviora

BREVIORA

Museum of Comparative Lioology

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. ZimMKus!* AND Davip C. BLACKBURN!

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) of 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:

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-

‘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@oeb.harvard.
edu.

Africa; Amphibia; Arthroleptidae; identification; Ranoidea; Phrynobatrachidae

atic research and identification in the field
and in museum collections. Within sub-
Saharan Africa, two such genera, Arthrolep-
tis Smith 1849 and Phrynobatrachus Gunther
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, 194la, b; Scott, 2005) and
the results of molecular phylogenetic analy-
ses (e.g., Bossuyt et al., 2006; Frost et al,
2006; Roelants et a/., 2007), it is now very
clear that these genera, although externally
similar, are only distantly related. In our

2 BREVIORA

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 (4rthroleptis 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 er 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 1s
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 al., 2006; Scott, 2005).
Both Scott (2005) and Frost et al. (2006)
argued for synonymizing these genera with

No. 513

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 1s
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, 1941la, 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

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

METHODOLOGY

Using the taxonomic, systematic, and
faunal literature as our guide, we examined
museum specimens for characteristics report-
edly 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
Arthroleptis 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 Arthroleptis and 246 adult
specimens representing 31 species of Phryno-
batrachus (Online Supplement: Appendix 1).

Arthroleptis and Phrynobatrachus appear
to generally exhibit different body shapes, with
Arthroleptis 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:

DIAGNOSING ARTHROLEPTIS AND PHRYNOBATRACHUS 3

Appendix 1). We used an analysis of covari-
ance to test for significant difference in the
slope of 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 « = 0.05.

RESULTS

The characters examined and evaluated in
this study are summarized in Table 1. Nearly
all characters reported to differentiate Ar-
throleptis 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, Arthroleptis have relatively
wider heads than Phrynobatrachus. The
coefficients (B) of linear regression of HdWd
on SVL are significantly different between
these two genera (Arthroleptis: B = 0.461, SE
= 0.006; Phrynobatrachus: B = 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 30mm, SVL (Pic. 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
Arthroleptis and Phrynobatrachus. The most

4 BREVIORA

No. 513

EXTERNAL MORPHOLOGICAL CHARACTERS USEFUL FOR DIFFERENTIATING BETWEEN A RTHROLEPTIS

AND PHRYNOBATRACHUS.

TABLE l.
Character Arthroleptis
Tarsal tubercle absent
Outer metatarsal tubercle absent
Heel tubercle (located on the proximal absent
end of the tarsus)
Circummarginal groove on terminal absent
phalanx
Pedal webbing absent
Median dorsal skin raphe present*
Hourglass or triple diadem pattern on variable
dorsum
Chevron-shaped glands in scapular region absent
Eyelid cornicle or spur absent
Sexually mature males
Elongate third finger variable
Dermal digital spines variable
Inguinal spines variable
Nuptial excrescence (thickened pad of absent
skin) on first finger
Femoral gland absent
Lateral vocal folds absent

Phrynobatrachus References

present Blackburn, 2005; Channing and
Boycott, 1989; Drewes and Perret,
2000: Scott, 2005

present Deckert, 1938

variable Scott, 2005

variable Blackburn, 2005; Scott, 2005

variable Schmidt and Inger, 1959; Stewart, 1967

absent Drewes and Perret, 2000; Laurent,
1957; Scott, 2005

absent Scott, 2005

variable Scott, 2005; Stewart, 1974

variable Perret, 1988; Rodel, 2000

absent Blackburn, 2005; Noble, 1931

absent Noble, 1931; Schmidt and Inger, 1959

absent Blackburn, unpublished

variable Scott, 2005; Stewart, 1967

variable Blackburn, 2005: Parker, 1935:
Stewart, 1967

variable Stewart, 1967

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

definitive characteristic that can be used to
differentiate between these two genera 1s 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
o Phrynobatrachus

E
£
Tv
=
Be}
ir
SVL (mm)
Figure 1. Head width (HdWd) plotted against

snout—vent length (SVL) for Arthroleptis (squares) and
Phrynobatrachus (circles).

2008

African frogs, but the presence or combina-
tion of particular characters can be useful in
distinguishing between adult Phrynobatra-
chus and Arthroleptis. Expansion of the
terminal phalanx of 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. minutus, 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

DIAGNOSING ARTHROLEPTIS AND PHRYNOBATRACHUS

n

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 Phrynohatrachus 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 1s 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

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-138084) 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

2008

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 subgular 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

DIAGNOSING ARTHROLEPTIS AND PHRYNOBATRACHUS 7

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 (Rédel,
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; Rédel, 1998: Rédel
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 al, 2003; Moritz and Cicero,
2004; Vences et al., 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

froglets

illustrating extensive webbing. E, Tips of left pedal digits II 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 | mm in 2A—D, F-H;

0.1 mm in 2E.

orc

M¢

Z A-13

v1IeW

lishing between Arthroleptis and Phrynoba-

showing elongate digit III. B, Fingers I
lermal spines. C, Right hand of male P
a anus (MCZ A-136757), below.
f male A. schubotzi (CAS 201717), left, and

y dermal

Ids. E, Throat of male P. auritus (MCZ A-

ible vocal folds. Scale bar 1 mm

2008

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 by the misidentification of voucher
specimens. For instance, nucleotide BLAST
searches against GenBank sequence data for
the Arthroleptis specimens for which Vences
et al. (2003) amplified the 16S rRNA gene
reveals that these specimens are misidentified
(GenBank sequences AF215139-40). AI-
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 University) 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.

DIAGNOSING ARTHROLEPTIS AND PHRYNOBATRACHUS 3,

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

femoral glands of Phrynobatrachus are |\o-
cated on the posterior thigh. However.

BREVIORA

No. 513

15. Sexually mature males, lateral vocal
folds: (0) absent: (1) present. When pres-

glands can be situated closer to either the ent. folds run roughly parallel to the
knee or vent. These glands are most easily lower jaw, at the lateral margins of
seen in living specimens because they are the throat. and form one or multiple
usually bright yellow. creases.

APPENDIX 2.

DISTRIBUTION OF MORPHOLOGICAL CHARACTERS AMONG SPECIES OF
ARTHROLEPTIS AND PHRYNOBATRACAUS. ALL CHARACTER STATES ARE
BINARY (0. 1). MISSING DATA ARE CODED AS *?”:; 0,1 DENOTES POLYMOR-

PHISM. REFER TO APPENDIX | FOR CHARACTER DESCRIPTIONS.

| Paar. 3 4A. 5° 6 7 8 9 10. fh 2 13 14 15
Arthroleptis adelphus 0 O 0 Or at 1 0.1 0 0 1 1 1 0 0 0
A. adolfifriederici O> Oy S07 “0% to at 0 OO 1 1 1 0 0 0
A. affinis 5°00 OF tise e031 Oren O ] ] 0 0 0 0
A. brevipes OO Os 20): dinmec 0 or 0 ? ? ? 0 0 0
A. crusculum Dano 0 Te i! 1 0.1 0 0 1 1 1 0 0 0
A. francei 0 160250 7 05 ff 7 os OTTO 1 | 1 0 0 0
A_ hematogaster Ge FeO. ey O ez, si 0 O57: O72 4 ? 0 0 0
A. lameerei 0 O 0 0 1 LO 0 0 1 0 0 0 0 0
A. nikeae 0 O 0 Oe vie ed 0 0 0 ? v4 ? 0 0 0
A. nimbaensis 0 O 0 UR Fe aa 1 21 | 0 0 1 1 1 0 0 0
A. “poecilonotus™ Gc 0.540" 0) ots iy SOs OF O'S E 1 1 0 0 0
A. pyrrhoscelis O00) 0 ss 0 0 0°. 1 0 0 0 0 0
A. reichei OF05 0) On tte 6-1 0” 40 ] 1 1 0 0 0
A. schubotzi io 0" 02a tt Ot 0 O 1 1 | 0 0 0
A. stenodactylus OO: 120) Orn le 401 0 O 1 1 1 0 0 0
A. sylvaticus Ome O02 120 Certs SiO 0 O east 1 0 0 0
A. taeniatus 0 O 0 Oa yt 1 0 0 0 1 1 1 ) 0 0
A. tanneri O76) 6. OL tie 0 Gr 0 1 1 1 0 0 0
A. troglodytes OF 70a 0770 ot ie 041 0 oO ? 1 1 0 0 0
A. tuberosus O70? Cie nOa ek 1051 OP 0 1 1 ] 0 0 0
A. variabilis 0 a0e O00" te Os 0 O 1 1 1 0 0 0
A. wahlbergii O00 5:0 St 3 01 0 O 1 1 1 0 0 0
A. xenochirus 0 O 0 Oe be cit 0.1 0 0 1 1 1 0 0 0
A. xenodactyloides OO OO, Ae Ost 0 O 1 1 1 0 0 0
A. xenodactylus OOO ees et a 0 0 O 1 1 1 0 0 0
A. zimmeri 0 O 0 Oye sis kz ? 0 0 ? ? ? 0 0 0
Arthroleptis sp. nov. 0-0") 0) OG. 1 A 0 0: 0, ~2 ? ? 0 0 0
Phrynobatrachus acridoides 5 On eal ie 0 0 1 0 O 0 0 1 1 1
P. acutirostris 1 1 1 | io 0 l 0 0 0 0 1 0 ]
P. africanus PPA Oy Te 201 0 0 1 0 O 0 0 1 0 1
P. auritus 1 1 1 1 Me MG 0 1 0 0 0 0 0 1 1
P_ batesii ees | 0 1 0 O 0 ] 0 O 0 0 ? ? ?
P. bequaerti isp On 10 Oe 0 0 ] OF 20 0 0 1 0 0
P. bullans tL 4 ] Oa 0 0 1 0 O 0 0 1 1 1
P. calcaratus 1 i 0a 20020 0 1 1 0 0 0 0 0 1

2008 DIAGNOSING ARTHROLEPTIS AND PHRYNOBATRACHUS 11
APPENDIX 2. Continued.

iC 2 3, 4.57 46 7 S. Serie iy] 2 13 14 15
P. cornutus 1 ] 0 i 0 6 0 ] 1 0 0 0 ? 2 ?
P. cricogaster ] ] ] ] 1 O 0 1 0 0 0 0 ] 0 0
P. cryptotis Pei (Os Os 0 0 0 ] 0 O 0 0 0 0 1
P. dendrobates ] l 0 L076 0 1 0 0 0 0 | 0 1
P. francisci 1 1 OF 2:0) Ail 20 0 ] 0 O 0 0 ? ? ?
P. inexpectatus heel OF 0 OL 30 0 G7 09 56 0 0 0 ] ]
P. keniensis 1 1 0 OF~ 04.20 0 1 0 0 0 0 1 0 1
P. kinangopensis 19 oh 0:16 SO aes 0 0 1 0 O 0 0 ] 0 ]
P. krefftii ee | 0 Pst 2D 0 1 0 O 0 0 1 1 1
P. mababiensis fe i O81 0; 0" 50 0 ] > san0 0 0 ? ? ?
P. manengoubensis | 0 1200 0 ] VucO 0 0 0 ] 1
P. minutus ie 00" 7.0'470 0 | Of>.0 0 0 ? ? ?
P. natalensis ] Por) Sil =p 0 0.1 O 0 0 0 1 1 1
P. pakenhami Os ae, eee att 0) 038" 20 0 0 0 1 ]
P. parvulus Pad ee OS 0S a 0 is=_ 05 =70 0 0 ] ] 1
P. plicatus bee te (0-2 Lr SO 0 ft 0 20 0 0 0 0 ]
P. rungwensis i as | 1 10: <0 0 1 0 O 0 0 1 1 1
P. sandersoni 1 1 1 1 0 0 1 0 0 0 0 1 1 1
P. scheffleri i, ot 1 0% 202740 0 1 0 O 0 0 ? ? ?
P. steindachneri i. AO Pe tes SO 0 ] 0 0 0 0 1 1 0
P. stewartae 1s 5 a4 | Gr 0 ] 0 0 0 0 1 1 1
P. ukingensis ee els OES Os SD 0 0.1 0 O 0 0 1 1 1
P. versicolor 1 1 0 ] tea0 0 ] 0 0 0 0 1 0 1

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