Zoosyst. Evol. 98 (1) 2022, 23-42 | DOI 10.3897/zse.98.76692

Epi a aes P00 Tease eM ______________am
D> PENSOFT. aN A

NATURKUNDE
BERLIN

A new slippery frog (Amphibia, Conrauidae, Conraua Nieden, 1908)
from the Fouta Dyallon Highlands, west-central Guinea

Karla Neira-Salamea', Joseph Doumbia’, Annika Hillers?,
Laura Sandberger-Loua', N’Goran G. Kouamé’, Christian Brede?,
Marvin Schafer!, David C. Blackburn®, Michael F. Barej!, Mark-Oliver Rédel!

Museum fiir Naturkunde Berlin — Leibniz Institute for Evolution and Biodiversity Science, InvalidenstraBe 43, 10115 Berlin, Germany
ONG EnviSud Guinée, Quartier Kipé T2 Commune de Ratoma, 030 BP 558 Conakry, Guinea

Wild Chimpanzee Foundation (WCF), Liberia Office, FDA Compound, Whein Town, Mount Barclay, Montserrado County, Liberia
Université Jean Lorougnon Guédé, UFR Environnement, Laboratoire de Biodiversité et Ecologie Tropicale, Daloa, BP 150, Céte d'Ivoire
Seydlitzstrasse 12, 23564 Liibeck, Germany

Department of Natural History, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA

NnoF WN BP

http://zoobank. org/1845C392-2447-49EF-96EE-7E2ZAEOFD993D

Corresponding author: Mark-Oliver Rodel (mo.roedel@mfn.berlin)

Academic editor: Johannes Penner @# Received 18 October 2021 # Accepted 2 December 2021 Published 19 January 2022

Abstract

We describe a new species of the genus Conraua from the Fouta Djallon Highlands in Guinea. The species is recognised as distinct
from nominotypical C. alleni, based on morphological evidence and is supported by a recent species delimitation analysis, based
on DNA sequence data. The new species 1s distinguished from its congeners by the unique combination of the following characters:
medium body size, robust limbs, only one instead of two palmar tubercles, the first finger webbed to below the first subarticular tu-
bercle, presence of a lateral line system, indistinct tympanum, two subarticular tubercles on fingers III and IV, venter in adults white
with dark brown spots or dark brown with grey or whitish spots. The new species differs from all congeners by more than 6% in the
DNA sequence of mitochondrial ribosomal 16S. We discuss isolation in Pliocene and Pleistocene forest refugia as a potential driver
of speciation in the C. alleni complex. We also emphasise the importance of conserving the remaining forest fragments in the Fouta
Djallon Region for the preservation of both its unique biodiversity and its valuable water sources for local people.

Resumé

Nous décrivons une nouvelle espece du genre Conraua des hauts plateaux du Fouta Dyjallon en Guinée. L'espece est reconnue comme
distincte du C. alleni nominotypique, sur la base de preuves morphologiques et est soutenue par une analyse récente de délimitation
des especes, basée sur des données de séquence d'ADN. La nouvelle espece se distingue de ses congéneres par la combinaison
unique des caractéres suivants: taille moyenne du corps, membres robustes, un seul tubercule palmaire au lieu de deux, premier
doigt palmaire jusqu'en dessous du premier tubercule subarticulaire, présence d'un systeme de lignes latérales, tympan indistinct,
deux tubercules subarticulaires sur les doigts III et IV, ventre blanc avec des taches brun foncé ou brun foncé avec des taches gris
ou blanchatre chez les adultes. La nouvelle espéce différe de ses congénéres avec plus de 6% de sa séquence d'ADN du ribosome
mitochondrial 16S. Nous discutons de I'isolement dans les refuges forestiers du Pliocene et du Pléistocéne comme facteur potentiel
de spéciation dans le complexe C. al/leni. Nous soulignons également l'importance de conserver les fragments de forét restants dans
la région du Fouta Dyallon pour préserver a la fois sa biodiversité unique et ses sources d'eau précieuses pour les populations locales.

Key Words
Anura, conservation, forest refugia, Upper Guinea forest zone, West Africa

Copyright Neira-Salamea K et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

24

Introduction

The Fouta Dyallon is a poorly studied mountainous re-
gion in west-central Guinea (Rodel et al. 2021). It is
characterised by plateaus with irregular profiles, per-
manent rivers in deep valleys, steep slopes with water-
falls and an elevational gradient of 1000-1600 m a.s.I.
(Young and Young 1992). The proximity to the Atlantic
Ocean and south-westerly winds generate higher lev-
els of precipitation than in other parts of West Africa
(Kamara et al. 2002; Descroix et al. 2020). Together,
the topographic and climatic complexity of Fouta Dyal-
lon creates a unique ecoregion with patches of tropical
and subtropical moist broadleaf forests in valleys, sur-
rounded by tropical and subtropical grasslands, savan-
nahs and shrublands (ecoregions, sensu Dinerstein et
al. 2017).

For decades, it has been stated that the Fouta Dyallon
harbours a large and unique floral diversity and that its
conservation should be a priority (Hepper 1968; Schnell
1968). However, the habitats of this area have experi-
enced severe degradation due to agricultural expansion
and cattle ranching (Couch et al. 2019, 2020). Several
plant species, endemic to the Fouta Dyallon, have not
been seen in over 60 years and are likely to be extinct
(Couch et al. 2019). In contrast to botanical invento-
ries, the fauna of the Fouta Dyallon has been largely
neglected though there has been some recent progress.
For instance, two candidate species of barb fishes, ge-
nus Enteromius, have been recognised in different ba-
sins of the Fouta Djallon (Schmidt et al. 2019). To date,
94 amphibian species are known from Guinea (Chan-
ning and Rédel 2019), with 25 species recorded from
the Fouta Dyallon (Hillers et al. 2008). However, Hillers
et al. (2008) suggested this diversity might be higher
and this is supported by the subsequent discovery and
description of a new Arthroleptis (R6del et al. 2011)
and two Odontobatrachus species (Bare et al. 2015b).
Hillers et al. (2008) also briefly mentioned and figured
an enigmatic, likely undescribed species of Conraua.
This taxon was subsequently listed as a distinct taxon
in Channing and Rodel (2019), but it has not yet been
formally described.

Seven species of Conraua are currently recognised
as valid (Neira-Salamea et al. 2021). They are
distributed in East (C. beccarii), Central (C. goliath,
C. crassipes, C. robusta) and West Africa (C. alleni,
C. derooi, C. sagyimase). A recent phylogenetic
analysis of the genus Conraua indicated that the
frogs, known as C. alleni, actually comprise a species
complex, including lineages from the Fouta Dyallon
(Blackburn et al. 2020). Here, we take a step forward
in disentangling the taxonomy of this species complex.
Through a combination of comparative morphology
and the recent molecular phylogenetic analyses of
Blackburn et al. (2020), we describe a new species
of Conraua that is known from Konkouré Fetto,
Hore Binti and Chute de Ditinn in the Fouta Dyallon
Highlands, central Guinea.

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Neira-Salamea, K. et al.: A new Conraua from Guinea

Materials and methods

We examined four Conraua specimens from Konkouré
Fetto, five specimens from Hore Binti and five specimens
from Chute de Ditinn, all of which are from the Fouta
Djallon Region in Guinea and all are deposited in the
collection of the Museum fiir Naturkunde Berlin, Germa-
ny (ZMB). The holotype and two paratypes of the new
species were included in the study by Blackburn et al.
(2020). All individuals were preserved in 75% ethanol.

For comparison, we examined specimens of all other
species of Conraua, including 19 specimens of C. alleni,
three C. beccarii, four C. goliath, four C. robusta, nine
C. crassipes, 34 C. derooi and eight C. sagyimase. The
comparative material included the holotype of C. alle-
ni from the Museum of Comparative Zoology (Harvard
University, Cambridge, USA; MCZ), four paratypes of
C. derooi from the Royal Museum for Central Africa
(Tervuren, Belgium; MRAC) and the Muséum national
d’Histoire naturelle (Paris, France; MNHN), the holo-
types of C. crassipes, C. sagyimase and C. robusta from
the ZMB and the holotype of Rana Griaulei Angel, 1934
(a synonym of C. beccarii) from the MNHN. All compar-
ative material is listed in Appendices 1 & 2. MOR mea-
sured the C. alleni holotype, the four C. derooi paratypes
and the R. Griaulei holotype. All other data were collect-
ed by KNS. To account for potential individual differenc-
es when measuring museum specimens, we included only
the data collected by KNS in statistical comparisons.

We measured the following morphometric variables:
snout—vent length (SVL, from tip of snout to posterior end
of vent), head length (HL, from tip of snout to posterior end
of head protuberance), head width (HW at corners of the
mouth), snout length (SL, from anterior edge of orbit to tip
of snout), eye diameter (ED, maximum horizontal diame-
ter), interorbital distance (IOD, shortest distance between
upper eyelids), upper eyelid width (UEW, maximum width
of upper eyelid), eye to nostril distance (EN, from anterior
edge of eye to centre of nostril), eye to snout distance (ES,
from anterior edge of eye to tip of snout), internarial dis-
tance (IND, between centre of nostrils), tympanum diame-
ter (TD, maximum horizontal diameter), eye to tympanum
distance (ETD, from posterior edge of eye to anterior edge
of tympanum), crus (tibiofibula) length (TL, from the bent
knee to heel), foot length (FL, from the proximal end of
tarsus to tip of fourth toe), toe IV length (T4), hand length
(HAL, from the proximal edge of the palm to the tip of the
finger IIT), finger II length (F3) and forearm length (FLL).
All measurements were taken with a digital calliper (+ 0.1
mm) and/or a dissecting microscope and are given in milli-
metres (mm). Measurements of the type series are present-
ed in Table 1, summarised measurements for the popula-
tions of Hore Binti and Chute de Ditinn are given Table 2.

Sex and maturity were assessed by examination of
gonads through an incision in the lateral body wall. Ad-
ditional qualitative morphological characters that we
examined include: tympanum detectability (distinct/
indistinct), webbing condition (complete/incomplete),
head shape in lateral view (round/pointed/truncated),

Zoosyst. Evol. 98 (1) 2022, 23-42

25

Table 1. Measurements [mm] of the type series of Conraua kamancamarai sp. nov. (holotype in bold); m = male, f = female, s =
sub-adult; SVL = snout—vent length, HW = head width, HL = head length, SL = snout length, ED = horizontal eye diameter, EN
= eye to nostril distance, ES = eye to snout distance, IND = internarial distance, IOD = interorbital distance, UEW = upper eyelid
width, TD = tympanum diameter, ETD = eye to tympanum distance, TL = crus length, FL = foot length including toe IV, T4 = toe
IV length, HAL = hand length, F3 = finger III length, FLL = forearm length; ZMB = Museum fiir Naturkunde, Berlin.

Voucher ZMB Sex SVL HW _~ HL SL ED EN~ ES
78432 f 71.7 23.7 23.5 7.0 6.5 4.3 8.2
78429 MSE Se POISG Ord! EGO," Sees. as. ™ Gu,
78430 S466 5 lot6 3" » BA" BO ASO oS.
78431 s 494 194 168 61 50 3.7 7.5

IND
4.0
4.6
3
4.1

1OD UEW TD ETD TL FL T4 HAL F3_ FLL
5.2 3.8 4.3 4.0 33.3 44.8 29.3 16.8 9.8 12.4
45 3.2 25/32°"35.0 210 Tai2? 8:2. JOA
4.1 3.1 21.6 29.1 209 124 64 84
By FS le) 2253" 30.6 22%. 1216. 97.5. 39

Table 2. Summary measures [mm] of Conraua kamancamarai sp. nov. from Hore Binti and Chute de Ditinn (referred material; total
of four males and females, respectively). SVL = snout—vent length, HW = head width, HL = head length, SL = snout length, ED =
horizontal eye diameter, EN = eye to nostril distance, ES = eye to snout distance, IND = internarial distance, [OD = interorbital dis-
tance, VEW = upper eyelid width, TD = tympanum diameter, ETD = eye to tympanum distance, TL = crus length, FL = foot length
including toe IV, T4 = toe IV length, HAL = hand length, F3 = finger III length, FLL = forearm length.

Sex SVL HW HL SL ED EN ES IND
mean f 81.7 28.3 25.1 8.0 Vans B 4. 10:3. 6:5
sd f 5.6 2AM SS— §OS THO = O15 0.9 09
max f 866 31.5 276 87 88 61 #114 74
min f 74.3 259 203 7.2 6.5 BO -o2 5.3
mean m 78.8 26.9 26.2 8.2 Vise 5.3 9.9 6.1
sd m 9.7 3.5 2t3™ 20'6 10 04 1.3 ileal
max m 886 303 283 90 84 #57 #=11.2 7.6
min Mute ZO £23238 23:4 7/55 16.65 472 286 510

relative length of fingers (1.e. comparative lengths were
assessed by pressing fingers together and fingers were
numbered from pre-axial to postaxial I to IV), presence of
an interorbital stripe, colour and shape (curved/straight/
slightly-curved) of the supra-tympanic ridge, presence or
absence of subarticular tubercles, undivided or divided
palmar tubercles, belly and throat colouration in alcohol
and presence of a lateral line system (see Lamotte and
Perret 1968; Fritzsch et al. 1987). Concerning the nomen-
clature of these lines (infra-orbital line, supra-orbital line,
upper lateral line, lower lateral line, median lateral line,
caudal lateral line and jugular line, anterior lower lateral
line, posterior lower lateral line and mandibular lateral
line), we followed Escher (1925) and Shelton (1970). Co-
louration of the back in preservation was only specified as

1OD UEW TD ETD TL FL T4 HAL F3_ FLL
Be 54.5, 5a Oo 35. B29 632.2= 19,1 PLO 146
TOS “OS sOi4y -215% 2a 7s (G4 “Gy OS" FOi6
C7 BSG. aheST Gaye “Seb NG2e2e” SorSeerA lor ee 1S 2
44 33 48 3.1 32.1 465 243 176 10.2 13.8
BGP 85:0" 4:39 (6.3,, "Stier 42.0.8 31.7 =2L6, 013.3. 134
OF “OF =08 233- HS 7b B= 40° 34 20
69 60 45 86 35.7 494 440 269 181 164
53 45 41 #40 328 322 244 17.5 10.1 12.0

being comparatively light or dark. The variation of qual-
itative morphological characters amongst all Conraua
Species 1s summarised in Table 3.

For statistical comparisons of morphological characters
with the new taxa, we focused on populations of the C.
alleni complex that were included in the recent phylogeny
of the genus (see Blackburn et al. 2020). Our analyses in-
cluded thirteen individuals of C. alleni sensu lato (Table 4,
Appendix 2), three individuals from Soyah, Fouta Djallon
(Appendix 2) and 10 from Konkouré Fetto, Hore Binti and
Chute de Ditinn, Fouta Dyjallon. We performed a principal
component analysis (PCA) to reduce morphometric vari-
ables and explore intraspecific differences of morphomet-
ric measurements amongst individuals (Neff and Marcus
1980). A subsequent discriminant function analysis (linear

Table 3. Comparison of qualitative morphological characters of all currently recognised Conraua species. All species have com-

pletely webbed feet.

Characters C. kamancamarai C. alleni C. sagyimase C. derooi C. goliath C. beccarii | C. crassipes C. robusta
Webbing hands finger | webbed absent absent
Palmar tubercle single double double double single single absent
Belly colouration white with brown | white or white white or white white or white uniform uniform white or white

spots with brown with brown with brown white or grey | white or grey with brown
mottling mottling mottling mottling
Throat colouration | white with brown white white with brown | white with brown] — white or white or white or white or white
spots or brown mottling mottling white with white with white with with brown
with white or grey brown brown brown mottling
spots mottling mottling mottling
Relative length of II>IV>Hl>| II>IV>l>1 II>IV>= II>IV>l>1 IHI>IV>I1>1 HI>IV>I>I IHI>IV>I1>1 II>IV>I1>1
fingers
Snout shape rounded rounded pointed rounded rounded rounded
Interorbital stripe mostly present mostly present | mostly present mostly absent mostly mostly mostly mostly absent
Supratympanic slightly curved straight or slightly curved curved curved curved curved curved
fold slightly curved
Lateral line system present present present present absent present absent absent
Tympanum indistinct indistinct indistinct indistinct distinct distinct distinct
visibility

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26

Neira-Salamea, K. et al.: A new Conraua from Guinea

Table 4. Summary measures (in mm) of adult Conraua alleni, m= male, f = female, N = sample size; SVL = snout—vent length, HW
= head width, HL = head length, SL = snout length, ED = horizontal eye diameter, EN = eye to nostril distance, ES = eye to snout
distance, IND = internarial distance, IOD = interorbital distance, UEW = upper eyelid width, TD = tympanum diameter, ETD = eye
to tympanum distance, TL = crus length; FL = foot length including toe IV, T4 = toe IV length, HAL = hand length, F3 = finger III
length, FLL = forearm length.

Sex N SVL HW_~ HL SL ED EN’ ES __ IND
mean f 5 58.4 22.3 20.7 56 65 3.7 7.3 45.0
sd Le Se 25810 sO SS" 2533 ASI 3P res
max if "5s 63:4,"76:4 0-241 6674 87-5: 5:07 =9:0s. £670
min tT 55-509 Pe bOr5alB? | 510-0 5:24 3:08, "6:3" a2
fmean* om: *83~-55-5 21°5°204- 5:9: 33.9: 33.7" 723° 4:5
sd my 8° Ph. 2.85 "2:9" Oo: <0:8 ~O0.5, 2 "0.7
max. om 8° 71%6- 27.3526). «%.Q Wid. 4:5 TGrb5. 527
min m 8 48.3 18.9 169 4.7 50 3.2 62 3.6

1OD UEW TD ETD TL THLE FL T4 +-\$HAL F3_ FLL
49 37 40 2.9 280 27.6 384 240 149 86 10.0
201 SIRS pals! Ieb- oll: elIhy Te6y LOiL se Sal Ah3
63 44 44 40 33.4 31.9 469 28.5 17.8 10.1 12.3
44 34 34 24 236 243 324 21.8 12.8 7.2 8.0
AD) 4356" 1325 22826. GO EATS: 3614 23: Serbs “7:9?” 1003
Oris TOPs sO (OR So FS BGs? 336s w 200° S119 1-6
62 40 41 3.5 340 34.0 47.3 30.8 165 98 14.0
40. 3:3 3:0, 24 23:4. 23.7 269° 166 thd 6:5 8.8

discriminant analysis, LDA) was used to calculate the re-
liability of morphological quantitative traits in assigning
individuals to the species groups, identified by Blackburn
et al. (2020). To visualise morphometric variation indepen-
dent of SVL, PCA was run on residuals of linear regres-
sions of the measured variables. LDA was applied on raw
morphometric measurements including SVL, because the
aim of the LDA was to determine the use of all morpho-
metric measurements to distinguish the different lineag-
es (Funk et al. 2012). Both analyses were performed on
log-transformed data. In some analyses, we investigated
both sexes together because sex is difficult to determine,
based on external morphology for most Conraua species
(see Neira-Salamea et al. 2021). Tympanum diameter (TD)
and eye to tympanic distance (ETD) were excluded from
the analysis, as the tympanum was not visible in most
specimens. Statistical analyses were performed in R ver-
sion 3.5.1 (R Core Team 2018) using the prcomp function
for PCA (with values scaled and centered) and /da function
for LDA from the MASS package (v.7.3-51.3; Venables
and Ripley 2002) and for PCA visualisation, we used the
function autoplot of the ggplot2 package (Wickham 2016).

Results

Based on the multi-locus phylogeny, published by Black-
burn et al. (2020), the Conraua populations from Fouta
Djallon are distinct from C. alleni, C. derooi and C. sagy-
imase, all of which form a large clade. Uncorrected p-dis-
tances of the mitochondrial 16S rRNA fragment (569 bp)
revealed high genetic divergence between the populations
from Fouta Dyallon and all other Conraua species. Uncor-
rected p-distances between the undescribed species from
Fouta Dyallon and other species is as follows: C. derooi,
6%; C. sagyimase, 7%; C. alleni, 7%; C. goliath, 9%; C.
robusta, 9%; C. beccarii, 11%; and C. crassipes, 12%.
These values are higher than the 3% threshold, usually ap-
plied for recognising anuran candidate species (Fouquet et
al. 2007; Vieites et al. 2009). Barcoded specimens within
the type series (ZMB 78432, 78429, 78430) of the unde-
scribed species had uniform 16S sequences. The genetic
distance between these frogs and a population from Soyah
(ZMB 90175, 90176), a site well situated within the Fouta

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Djallon, was about 3% (mean + sd: 0.0295 + 0.008, range:
0.028—0.030). Therefore, we excluded the Soyah popula-
tion from the description of the new species and suggest
further bioacoustics analysis to confirm if it belongs to
the species described below. The frogs of the type series
further showed genetic differences to other populations
from the Fouta Djallon. For instance, a specimen from
Hore Binti (ZMB 90177) differed from the type series by
0.9% (this specimen differed from the Soyah population
by 0.029% + 0.0008, 0.028—0.030%). Although we be-
lieve that specimens from Hore Binti and Chute de Ditinn
represent the new species, we list and describe material
from these locations only as additional referred material.

We analysed morphological characters (Tables 1, 2
and 4) of all lineages of the C. alleni complex includ-
ed in the genus phylogeny (for details, see ‘Material and
Methods’ and Blackburn et al. 2020; for morphological
comparisons with other Conraua species, see ‘Diagno-
sis’ below). Five principal components with eigenval-
ues > 1.0 (PCI—PCS5) accounted for 72.8% of the total
character variation (Fig. 1; Table 5). PC1 accounted for
29.3% of the variation and loaded most strongly for eye
to snout distance (ES) and crus (tibiofibula) length (TL).
PC2 accounted for 14.8% of the variation and loaded
negatively most strongly for finger III length (FE) and
eye to nostril distance (EN). PC3 was mainly impacted
by upper eyelid width (UEW), eye diameter (ED) (both
negatively). PC4 was dominated by hand length (HAL,
negatively) and interorbital distance (IOD). PC5 loaded
most strongly for head length (HL, negatively) and inter-
narial distance (IND). A plot of PC1 versus PC2 showed
both large variation amongst individuals, as well as a
large amount of overlap between the four genetic lineages
included (Fig. 1). However, a subsequent linear discrim-
inant analyses (LDA) of the morphological raw data cor-
rectly assigned all individuals to their respective genetic
lineage. The characters contributing most to the species
assignment were SVL, HL and TL. Coefficients of linear
discriminant analysis (LDA) are provided in Table 6.

An unexpected diagnostic character was the number of
palmar tubercles. Whereas C. derooi, C. sagyimase and
C. alleni have a divided, double palmar tubercle (an outer
and a middle one), all Conraua populations from the
Fouta Dyjallon exhibit only an undivided palmar tubercle

Zoosyst. Evol. 98 (1) 2022, 23-42

PC2 (14.78%)
o
oO

-0.2

-0.2 0.0 0.2
PC1 (29.34%)

Figure 1. Axes I and II from the Principal Component Analyses
(PCA), based on 15 size-corrected morphometric variables of
four Conraua lineages from West Africa; only adult frogs includ-
ed. Black = Conraua kamancamarai sp. nov. types from Konk-
ouré Fetto (asterisk = female holotype), red = Conraua kamanca-
marai sp. nov. populations from Hore Binti and Chute de Ditinn,
blue = Conraua populations from Soyah, Fouta Dyallon, yellow
= C. alleni; triangle = females, circle = males; compare Table 5.

Table 5. Principal component analysis on morphometric mea-
surements for Conraua kamancamarai sp. nov., C. alleni and
Conraua populations from Soyah. Character loading, percent-
age (%) and cumulative percentage of explained variance for
principal components with eigenvalues > 1.0, (PC) I-V; vari-
ables with the highest loadings are given in bold; compare Fig.
1; for abbreviations of morphological measures, see ‘Materials
and Methods’ section or Tables 1, 2 and 4.

Variable PC1 PC2 PC3 PC4 PC5
HW 0.33 0.08 -0.03 0.16 -0.07
HL 0.26 0.06 -0.19 G25 -0.62
SL 0.23 -0.31 -0.04 Qa 3. 0.31
ED 0.10 0.30 -0.44 0.10 -0.13
EN 0.13 -0.45 0.19 0.34 0.15
ES 0.37 -0.07 0.04 0.25 0.10
IND 0.26 0.05 -0.35 OxL1 0.45
1OD 0.30 -0.04 0.10 0.40 -0.22
UEW -0.20 -0.29 -0.49 0.06 -0.09
Te 0.35 0.08 -0.04 -0.33 -0.17
FL 0.31 0.20 0.19 -0.39 0.24
T4 0.30 0.32 0.19 0.02 0.09
HAL 0.26 -0.25 -0.33 -0.45 0.01
FS Only, -0.49 -0.04 -0.28 -0.13
FLL 0.07 -0.23 0.41 -0.16 -0.31
Eigenvalue 4.40 222. 75 1.35 1.20
% 29.34 14.78 11.64 8.98 8.02
Cumulative % 29.34 44.12 SDaLO 64.74 $2.75

(Fig. 2). In C. derooi, the divided palmar tubercles are
nearly connected and the thenar tubercle is clearly sepa-
rated from the middle palmar tubercle. In C. alleni, the di-
vided palmar tubercles (outer and middle) are very close

27

Table 6. Coefficients of linear discriminant analysis (LDA)
based on the 16 morphometric variables of three Conraua pop-
ulations from West Africa: Conraua kamancamarai sp. nov.,
Conraua populations from Soyah, Fouta Djallon and C. alle-
ni, those characters that contribute most are given in bold; for
abbreviations of morphological measures, see ‘Materials and
Methods’ section or Tables 1, 2 and 4.

LD1 LD2
SVL -33.06 -6.11
HW 5.43 6.73
HL 0.67 31.97
ol -13.74 -9.47
ED 4.93 ae aw as
EN 573 1Sso7
ES 19.41 -0.79
IND 1.26 3:27
lOD 10.45 -18.10
UEW 3123 -2.18
TL 23.24 -14.45
FL Srli2 are
T4 -12.27 6.19
HAL 8.53 5.04
ES Dome te) -2.96
FIL -1.84 -9.82

to each other, but not connected and the thenar tubercle
is separated from the middle palmar tubercle. In C. sagy-
imase, the divided palmar tubercles (outer and middle)
are close but separated and the middle palmar and the the-
nar tubercle are distinctly separated (Fig. 2).

The most conspicuous morphological character of di-
agnostic value is the ventral pattern. Whereas most West
African Conraua species have a predominantly white
or light grey venter, often with dark mottling, but never
with distinct spots or blotches, the new species exhib-
its a white ventral colour with well-defined dark brown
spots and blotches (or dark colour with scattered grey or
whitish spots) (Figs 3, 5 and 6). This ventral colour pat-
terning is most distinct in adults, but already developed
in some subadults.

Based on the genetic differences and the above diag-
nostic characters, we describe the Conraua populations
from Konkouré Fetto, Hore Binti and Chute de Ditinn as
a species new to science. Due to their genetic divergence
from other Fouta Dyallon populations, we exclude the
population from Soyah until larger sample size and call
recordings are available (e.g. compare Neira-Salamea et
al. 2021). This population might represent another cryptic
species that would be described separately.

Description of the new species

Conraua kamancamarai sp. nov.
http://zoobank.org/625E8A78-5BB3-4EB1-8405-159C93 135699
Figs 2-6

Holotype. ZMB 78432 (field and tissue #: GN11-140;
GenBank # for 16S: MT669400) adult female, Guinea,
Fouta Dyjallon Region, Konkouré Fetto, 10°20'28.21"N,

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28

Neira-Salamea, K. et al.: A new Conraua from Guinea

Figure 2. Ventral view of hands of four Conraua species showing palmar tubercles differences. Whereas three species possess a
divided (outer and middle) palmar and a thenar tubercle, Conraua kamancamarai sp. nov. has an undivided palmar and a thenar tu-
bercle. a. Conraua kamancamarai sp. nov. holotype (ZMB 78432); b. C. derooi (MNHN-RA 1993 4087); ¢. C. sagyimase (UWBM
Herp 05841, paratype); d. C. alleni (ZMB 90390); Scale bars: 5 mm.

12°10'16.82"W, 650 m a.s.1., 20 June 2011, collected by
Laura Sandberger-Loua & Joseph Doumbia.

Paratypes. ZMB 78429 (field and tissue #: GN11-130;
GenBank # for 16S: MT669399) adult male; ZMB 78430,
ZMB 78433 (field and tissue #: GN11-133, GN11-136;
GenBank # for 16S: KF693389) subadults, all other data
as holotype.

Additional referred material. For the description of the
new species, we restrict the type series to the population
from Konkouré Fetto. We do so because there may be ad-
ditional undescribed diversity within the group identified
as “C. alleni 1b” by Blackburn et al. (2020). The vouch-
ers from Hore Binti, Pita and Chute de Ditinn, Dalaba
(all from Fouta Dyallon, Guinea) are, therefore, listed as
additional referred material (see Table 2).

Hore Binti, Pita. ZMB 90301, ZMB 90304 (field and
tissue #: CB2010-055, CB2010-061), adult females; ZMB
90302 (field and tissue #: CB2010-056), adult male; ZMB
90177 (field and tissue #: CB2010-057; GenBank # for
16S: MT669401), subadult; ZMB 90303 (field and tissue
#: CB2010-059), subadult, 10°51'04.8"N, 12°31'14.1"W,
657 ma.s.l.; 22 July 2010, collected by Christian Brede
& Joseph Doumbia.

Chute de Ditinn, Dalaba. ZMB 90305, ZMB 90307,
ZMB 90309 (field and tissue #: CB2010 082, CB2010
089, CB2010 091), adult males; ZMB 90306, ZMB 90308
(field and tissue #: CB2010 088, CB2010 090), adult fe-
males, 10°49'13.1"N, 12°11'30.7"W, 760 m a.s.1.; 24 July
2010, collected by Christian Brede & Joseph Doumbia.

Diagnosis. The new species resembles other members of
the genus Conraua Nieden, 1908. Conraua kamancama-
rai Sp. nov. is an aquatic frog with the following traits:
smooth dorsal skin, covered with scattered small, round-
ed warts on back and longitudinal ridges on dorsal part of
hind legs; venter skin smooth; three odontoid projections
on lower jaw, one at symphysis and one to each side on
dentary; vocal sacs absent; fully webbed feet, 1.e. to end
of last phalanx of toe. Conraua kamancamardi sp. nov. 1s

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closely related to a clade including C. alleni sensu stricto,
C. derooi and C. sagyimase (see Blackburn et al. 2020).
Genetic distances between the new species and all other

Figure 3. Conraua kamancamarai sp. nov., female holotype
(ZMB 78432) in lateral, dorsal and ventral view. Scale bar: 20 mm.

Zoosyst. Evol. 98 (1) 2022, 23-42

29

Figure 4. Conraua kamancamarai sp. nov., female holotype (ZMB 78432) lateral view of head and ventral view of left foot. Scale

bar: 10 mm.

Conraua species were higher than 6% in the examined
part of the 16S gene.

Conraua kamancamarai sp. nov. can be distinguished
from C. goliath by a rounded snout (pointed in C. go-
liath), the absence of short dorsal skin ridges, a white
venter with dark brown botches (yellow venter in C. go-
liath), the presence of a lateral line system, an indistinct
tympanum, a wide tarsal fold and by having more than
one subarticular tubercle on fingers (one in C. goliath).
Conraua kamancamarai sp. nov. differs from C. crassipes
by a white venter with dark brown blotches (uniform

white or cream in C. crassipes), an indistinct tympanum,
the presence of a lateral line system, by a conspicuous out-
er metatarsal tubercle (less conspicuous in C. crassipes)
and by lacking a dermal fold near the elbow. Conraua
kamancamarai sp. nov. differs from C. beccarii by the
absence of a transverse fold behind the eyes and across
the interorbital region, by lacking a swollen post-occipital
and suprascapular region in adult males, by a white-co-
loured venter with dark brown blotches (no spots in C.
beccarii) and by having a head that is as wide as long
(wider than long in C. beccarii). Conraua kamancamarai

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30 Neira-Salamea, K. et al.: A new Conraua from Guinea

Figure 5. Conraua kamancamarai sp. nov., female holotype (ZMB 78432) in life.

sp. nov. differs from C. robusta by having ahead thatisas_ —formly white or with dark mottling in C. robusta) and the
wide as long (wider than long in C. robusta), by havinga presence of a lateral line system. Conraua kamancamarai
U-shaped notched tongue-tip (tip of tongue rounded inC. _ sp. nov. differs from C. alleni sensu stricto by having an
robusta), by a white venter with dark brown blotches (uni- —_— undivided palmar tubercle, by having a white-coloured

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Zoosyst. Evol. 98 (1) 2022, 23-42

eal

Figure 6. Dorsal and ventral views of paratypes of Conraua kamancamarai sp. nov., male (ZMB 78429), subadults (ZMB 78430,

ZMB 78431). Scale bars: 20 mm.

venter with dark brown blotches (uniform light or light
with dark mottling in C. alleni), by a larger inner meta-
tarsal tubercle, a wider tarsal fold and by the presence of
webbing between fingers I and II. Conraua kamancama-
rai sp. nov. differs from C. derooi by having a more slen-
der body and limbs, a slightly curved supratympanic fold
(distinctly curved in C. derooi), two subarticular tubercles
on finger III (one in C. derooi), by lacking a swollen pos-
toccipital and suprascapular region in adult males, by the
absence of a divided palmar tubercle, by a white venter
with dark brown blotches (uniform whitish or with dark
mottling in C. derooi) and by the presence of webbing
between fingers I and I. Conraua kamancamardi sp. nov.
differs from C. sagyimase by having narrower fingertips,
a wider tarsal fold, by the absence of a divided palmar

tubercle, by a white venter with dark brown blotches (uni-
form pale or with dark mottling in C. sagyimase) and by
the presence of webbing between fingers I and II.

Description of the holotype (Figs 2—5; measurements
in mm). Adult female; slightly dorsoventrally flattened,
short and rounded body; snout rounded in dorsal and lat-
eral view, with upper lip slightly projecting forward; SVL
71.7; head width 23.7, approximately equal to head length
23.5; head length 33% of SVL; snout length 7.0, 30% of
head length; eye—-nostril distance 4.25; eye—snout distance
8.1; internarial distance 4.0, slightly larger than interorbit-
al distance 5.2; nostrils protuberant, directed dorsolateral-
ly, visible in lateral and dorsal view; large eyes, project-
ing laterally beyond margins of head in dorsal view; eyes

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OZ

projecting slightly above dorsal margin of head in lateral
view; eye diameter 6.5, horizontal diameter of tympanum
4.3; upper eyelid width 3.8, 73% of interorbital distance;
eye-tympanum distance 3.9; tympanum indistinct; can-
thus rostralis distinct and rounded; loreal region concave;
slightly curved supratympanic fold extending from poste-
rior edge of eye to shoulder, joining the lateral fold; upper
lip slightly protruding; premaxillary and maxillary teeth
slender and pointed, three odontoid projections on lower
jaw, one on at symphysis and one to each side on dentary;
vomerine teeth pointed; about half of anterior tongue at-
tached to floor of mouth, tongue-tip with U-shaped notch.

Forelimbs robust; forearm length 12.4, 74% of hand
length 16.8; thenar and palmar tubercle oval and protrud-
ing, palmar tubercle larger than thenar tubercle; shape of
fingers conical, wider at bases and narrower towards tips;
finger tips rounded, non-expanded; one subarticular tuber-
cle on fingers I and I; two subarticular tubercles on fingers
II and IV; subarticular tubercles absent on base of fingers;
relative length of fingers: III > ITV > I ~ I, length of finger
III 9.8; fingers I and IT webbed to first subarticular tubercle.

Hind limbs moderately robust; crus length 33.3, 46%
of the SVL; foot including longest toe 44.8, 62% of
SVL; elongated, prominent oval inner metatarsal tuber-
cle, more than twice as long (4.4) as wide (1.7); outer
metatarsal tubercle absent; supernumerary plantar tu-
bercles absent; subarticular basal tubercles absent; one
subarticular tubercle on toes I and II; two subarticular
tubercles on toes III and V, three subarticular tubercles
on toe IV; toe tips rounded, forming small discs, as broad
as subarticular tubercles; relative lengths of toes: VI > HI
>V> II >I; length of toe IV 29.3; webbing complete,
i.e. to end of last phalanx of toe; dermal fringing on outer
surfaces of toes I and V, forming lateral skin folds; wide
tarsal fold.

Skin texture on dorsal parts of head, body, flanks and
limbs smooth with scattered, small, rounded warts; upper
eyelid skin with many warts; inner surface of upper arm
smooth; dorsal surface of crus with 12 rows of longitudi-
nal ridges; ventral skin smooth, throat with longitudinal
folds; a post-gular (thoracic) fold extending to level of
forelimbs insertion; lateral line system with jugular line,
upper lateral line, lower lateral line, median lateral line,
caudal lateral line, infra-orbital line, supra-orbital line,
mandibular lateral line and anterior lower lateral line (see
Shelton 1970).

Colouration in preservative (after 10 years in 75%
ethanol; Figs 2—4). Dorsum brown with scattered small
cream warts, warts more abundant on posterior surface;
light interorbital stripe; upper eyelids brown with abun-
dant lighter spots; lips brown or brown with lighter mot-
tling; supratympanic fold brown; lateral fold light; upper
arm fold light; dorsal surface of flanks brown; ventral sur-
face of flanks white or light grey with brown spots; dorsal
surface of legs brown with dark scattered spots; dorsal
surface of toes I, I, III and IV with brown mottling; dor-
sal surface of toe V brown; dorsal surface of arms brown

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Neira-Salamea, K. et al.: A new Conraua from Guinea

with scattered light brown warts; dorsal surface of fin-
gers I and II light brown with dark mottling, fingers III
and IV brown; ventral surface of throat and belly whitish
with dark brown blotches; ventral surface of crus and feet
light brown with dark mottling; ventral surface of hands
brown.

Colouration in life (Fig. 5). Dorsum dark brown with
scattered lighter dots and dark spots; lips same colour as
dorsum, paler on lateral surfaces; flanks dark; iris gold;
ventral surfaces white with dark brown spots; ventral sur-
face of hands dark, tubercles lighter; ventral surfaces of
legs with scattered reddish dots; lateral fold light brown.

Variation (Figs 6—7; Tables 2 and 3). Overall, the para-
types are similar to the holotype in external appearance
and colouration. Dorsal colouration ranges from uniform
dark brown (ZMB 78432) to predominantly brown with
dark mottling (ZMB 78430, ZMB 78431) or predomi-
nantly brown with dark spots. In the male paratype (ZMB
78429), the jugular lateral line is more conspicuous than
in the holotype and the upper lip is light brown. Ventral
colour pattern of all specimens similar: whitish with dis-
tinct brown blotches, however, these blotches are lighter
in the subadult paratypes (ZMB 78430, ZMB 78431).

Referred specimens from Hore Binti: Dorsal coloura-
tion varies from similar to the type series (ZMB 90177,
ZMB 90303) to darker (ZMB 90301, ZMB 90302, ZMB
90304) or a dark dorsum with light brown spots (ZMB
90302). Ventral colouration of some individuals deviates
from that of the type series by a greyish belly with dark
mottling and a dark throat with lighter spots (ZMB 90304,
ZMB 90301). Ventral colouration of ZMB 90302 is diffi-
cult to define due to its preservation condition. Ventral co-
louration of juveniles is showing fewer dark spots than in
adults (ZMB 90177, ZMB 90303). Lateral line system in
ZMB 90304 is more conspicuous than in the type series.

Referred specimens from Chute de Ditinn: Dorsal
colouration of all individuals darker than that of type
series, some individuals with light brown spots (ZMB
90307, ZMB 90308) that are absent in the type series.
Some individuals present a different colouration pattern
than type series on the throat with a dark base colour and
lighter spots (ZMB 90306, ZMB 90307, ZMB 90308).
Belly of ZMB 90305 greyish with dark mottling, differ-
ent to the colour pattern of the type series, probably due
to preservation. The lateral line system in all individuals
is more conspicuous than in the type series. A posterior
lower lateral line (see Shelton 1970) is present in ZMB
90307 and ZMB 90309. This part of the lateral line sys-
tem was not detected in the type series.

For variation in life colouration of Conraua specimens
from various localities in the Fouta Dyallon Region, see
Figure 7.

Distribution (Fig. 8). So far, the known range of Conraua
kamancamardi sp. nov. is restricted to the type locality at
Konkouré Fetto and to two other sites, Hore Binti and

Zoosyst. Evol. 98 (1) 2022, 23-42 oie)

Figure 7. Colouration of life Conraua kamancamarai sp. nov. from the Fouta Djallon and surrounding region, Guinea, illustrating
variation in colour pattern and skin texture. a. From Dubreka, River Bindinbandan (10°22'21.9"N, 13°9'16.8"W, 199 m as.1.);
b. From Dalaba, Chute de Ditinn; c. From Hore Binti; d. From Dubreka, River Bindinbandan (10°22'21.9"N, 13°9'16.8"W, 199 m
a.s.l.); e. From Dalaba, Chute de Ditinn; f. From Télimeélé, locality Kourakoto, river Didounpourigue (10°55'30.4"N, 13°47'39.4"W,
238 ma.s.l.); frogs in lower row in typical calling position, sitting in shallow water; specimens either not collected or not assignable
to a voucher specimen, whereas the frogs from Hore Binti and Chute de Ditinn can be assigned to Conraua kamancamardi sp. nov.
without doubt; the other frogs may represent an undescribed Conraua.

Chute de Ditinn. Although the species probably occurs Biology and habitat (Figs 9 and 10). The new species
in a few more rivers that have not yet been surveyed, the occupies fast-flowing rocky streams with waterfalls within
entire range almost certainly will not extend beyond the riverine forest in mountainous areas in the Fouta Dyallon,
Fouta Dyjallon, where it 1s very likely patchily distributed. | Guinea (Figs 9 and 10). Like other frogs of the genus, they

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34

Neira-Salamea, K. et al.: A new Conraua from Guinea

- Sierra Leone

Figure 8. Map of known localities of Conraua kamancamarai sp. nov. in the Fouta Djallon Highlands, Guinea. Inset (upper right)
shows a map of West Africa indicating in red, the area of occurrence of the new species in the Fouta Dyallon and in blue, the assumed
type locality of C. alleni. Known localities of Conraua kamancamarai sp. nov. are shown in red (Konkouré Fetto, type locality: star;
Hore Binti: circle; Chute de Ditinn: square), the population from Soyah, potentially representing another undescribed Conraua, is
given in yellow. Altitudinal range is indicated with light shading from lowlands (112 m a.s.1.) to dark shading highlands (1089 m
a.s.l.). Sources: OpenStreetMap (2020), U.S. Geological Survey (2020).

are predominately nocturnal and aquatic. Despite their oc-
currence in fast flowing streams, adults show a preference
for calmer river sections, where turbulent water is absent.
Usually, frogs are encountered at least partly submerged in
shallow water, facing the riverbanks. When outside of the
water, they remain within jumping distance to water. Dis-
turbed frogs seek shelter on the ground of pools, sometimes
trying to burrow deeper into them and cover themselves
with gravel or substrate (Fig. 11). This behaviour is similar
to what Knoepffler (1985) described for Conraua crassipes.
They call (whistle) with an open mouth, sitting in shallow
water (Fig. 7; compare Amiet 1990). Mating has never been
observed by the authors; however, single observations of
clutches and jelly remnants of spawn indicate that oviposi-
tion sites are small puddles or depressions on the riverbanks
near the spray water zone of cascades and waterfalls. Con-
raua tadpoles usually were observed in silted calm ponds
where up to 50 tadpoles of about the same size have been
encountered. If this and other species of the genus show
breeding behaviour comparable to Conraua goliath (com-
pare Schafer et al. 2019), remains to be researched.

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The surroundings of the forest fragments where the spe-
cies occurs are generally degraded by anthropogenic distur-
bance, particularly peanut and rice crops and cattle grazing.
The type locality (Fig. 10) is located between Konkouré and
the largest city within the Fouta Djallon, Mamou, within a
relatively short distance to the connection road and was sur-
veyed on 20 June 2011. Along the national route one (N1),
one of the largest roads connecting the East with the West of
Guinea, houses are numerous, but already within a relatively
short distance to the road, human presence may be consider-
ably scarcer. Slopes are either covered by an open, short, dry
forest with signs of cattle grazing and used for charcoal pro-
duction (Fig. 10) or comparatively large fields for peanuts or
rice crops. Only steep slopes surrounding rivers had some-
times larger trees and denser vegetation with higher humidi-
ty levels than the surroundings. The type locality is at a river
within denser forest, with large boulders and some cascades,
allowing for a diverse river site with fast and slow flowing
parts and comparatively clear water (Fig. 10). These forests
are not protected and were in the past burned by the popu-
lation as protest against government decisions in Conakry.

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35

Figure 9. The Fouta Djallon, Guinea and habitats of Conraua kamancamarai sp. noy. Hore Binti landscape (upper photo) and
typical river habitats.

The classified forest (partly protected areas allowing
forestry) of Hore Binti is located within a mountainous
area containing several freshwater sources. It was sur-
veyed from 22—23 July 2010. Many fast-flowing streams
with cascades have its source on the mountain. The habi-
tat degradation due to anthropogenic alterations was dra-
matic and only very small forest fragments remained. The
anthropogenic pressure consisted of cultivations/fields
(mainly peanut and rice) and grazing cattle. Only streams
were surrounded by some remaining larger trees. The Di-

tinn / Dalaba site was within a small fragment of gallery
forest with a stream, next to the waterfall of Ditinn. It
was surveyed from 24—25 July 2010. Although there is a
small village next to the forest, only minor anthropogenic
alterations were detectable.

Threat status. Conraua kamancamarai sp. nov. should
be considered Data Deficient (DD) because more infor-
mation is required to make an adequate assessment of the
species’ extinction risk. However, if the species range is

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36

i

ACE

nich LE ‘cat ine

Neira-Salamea, K. et al.: A new Conraua from Guinea

Figure 10. The type locality of Conraua kamancamarai sp. nov. near Konkouré Fetto, Fouta Djallon, Guinea (10°20'28.21"N,
12°10'16.82"W, 650 ma.s.1.). The frogs live in clear, fast flowing streams, with riverine forest. The surroundings are heavily degrad-
ed by agriculture, cattle grazing and charcoal production (inset figure).

indeed restricted to the sites of Konkouré Fetto, Hore Binti
and Chute de Ditinn, the species should be categorised as
Endangered (EN) following the IUCN (2012) criteria B2bi
(continuing decline, observed, inferred or projected in extent
of occurrence) and B2bii1 (continuing decline, observed, in-
ferred or projected, in area, extent and/or quality of habitat).

Etymology. This species is dedicated to Kaman Camara

(Fig. 12), our long-term field assistant and friend, who
started working with MOR in 2002 on a survey to the

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Simandou Range that was organised by Conservation In-
ternational (Rodel and Bangoura 2004). From 2007 until
his recent death, Kaman was a member of our Guinean
team, investigating the amphibians of the Nimba Moun-
tains and other Guinean areas. Kaman had outstanding
skills in detecting and catching frogs, and, more impor-
tantly, an unswerving positive attitude. A day could be
completely exhausting and frustrating, but with a simple
joke from Kaman all was good again! Kaman was born
and lived in a remote village at the western foothills of

Zoosyst. Evol. 98 (1) 2022, 23-42 37

Figure 12. Kaman Camara in June 2007 on Mount Nimba, Guinea. Inset figure taken on a Rapid Assessment to south-eastern Guin-
ea, organised by Conservation International and Kaman’s first experience with frog work, from left to right: Mark-Oliver Rédel,
Mohamed Alhassane Bangoura and Kaman Camara.

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38

the Simandou Range. He never received any formal ed-
ucation. Still, he repeatedly rejected other better paying
job offers from mining companies, preferring instead to
work with his frog team whenever it was possible. Ka-
man died in June 2020 after a short severe disease. These
frogs shall be a permanent memory to an outstanding per-
son! We suggest ‘Kaman Camara’s Slippery Frog’ as the
English common name, ‘la grenouille glissante de Kaman
Camara’ in French and in the local language Poular: ‘T6ti
bhowroundi de Kaman Camara’.

Discussion

Surveys in West Africa over the past 20 years have re-
vealed previously unknown populations of the genus
Conraua and prompted the need for re-evaluating the
taxonomy of this genus, including exploring the poten-
tial for undescribed and morphologically-cryptic species
(Rodel and Branch 2002; Rodel and Bangoura 2004;
Hillers et al. 2009; Leaché et al. 2020). Based on com-
parative morphology and molecular phylogenetic data,
we describe a new species, Conraua kamancamarai
from three sites in the Fouta Djallon, Guinea, thus con-
firming the existence of an Unconfirmed Candidate Spe-
cies (UCS) (sensu Vieites et al. 2009), as suggested by
Blackburn et al. (2020).

The uncorrected p-distances of mitochondrial 16S be-
tween the new species and all other Conraua species was
greater than 6% and, thus, well above the usual thresh-
old (~ 3%) for potential anuran candidate species (Fou-
quet et al. 2007; Vieites et al. 2009). Although we had
no bio-acoustic data available and individuals overlap
in their body shape with those of C. alleni, these pop-
ulations are diagnosable as a distinct species by various
characters, including the absence of a divided palmar
tubercle, having a first finger webbed to below the first
subarticular tubercle and the unique ventral colour pat-
tern in adults.

Conraua kamancamarai is the most westerly distrib-
uted Conraua species. Its closest congeners are found to
the east, including the C. alleni complex from Guinea
(east of the Fouta Dyallon to lowland forests in western
Ghana, west of the Atewa Forest Range), C. sagyvimase in
the Atewa Range Forest in Central Ghana and C. derooi
along the escarpment of the montane border in western
Ghana and eastern Togo. Similar patterns of distribution
are also found in other pairs or groups of closely related
vertebrates; for instance, toothed frogs (Odontobatra-
chus; Bare] et al. 2015b), reed frogs (Hyperolius, Rodel
et al. 2010; Channing and Rodel 2019), horseshoe bats
(Rhinolophus, Fahr et al. 2002) and suckermouth catfish-
es (Chiloganis, Schmidt et al. 2017).

Although the fauna of the Fouta Dyallon has been poor-
ly studied compared to other forests in West Africa (R6del
et al. 2021), studies of its flora have identified the region
as a centre of endemism and species richness (Porembski
et al. 1994; Poorter et al. 2004). These two metrics are
commonly used to recognise areas that may have been

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Neira-Salamea, K. et al.: A new Conraua from Guinea

forest refugia (Hillers 2008; Tchouto et al. 2009), ie.
forests that persisted during cold and dry climatic peri-
ods (Haffer 1969; Couvreur et al. 2021). By investigat-
ing the phylogeography of several puddle frog species,
Phrynobatrachus, Hillers (2008) confirmed the presence
of macro- and micro-forest refugia in the highlands of
Fouta Dyallon during the Pleistocene. This finding, to-
gether with the results of Blackburn et al. (2020), which
found the divergence of C. a/leni into various lineages
during the Pliocene and Pleistocene, suggests allopatric
speciation, associated with isolation in Pleistocene forest
refugia, led to the evolution of one or more endemic Con-
raua species in the Fouta Djallon. Pliocene—Pleistocene
speciation events are well documented for other African
forest taxa (e.g. rodents: Nicolas et al. 2010; Bohoussou
et al. 2015; legumes: Duminil et al. 2013; bats: Hassanin
et al. 2015).

Today the ecosystems of the Fouta Dyallon are heavily
degraded (Wilson 1992; Couch et al. 2019), threatening
the area’s unique biodiversity (Schmidt 2014; Bare] et
al. 2015a) including this newly-described Conraua spe-
cies. A preliminary assessment of the conservation status
of vascular plants identified Guinea as amongst the top
countries in tropical Africa with the highest proportion
of threatened species (Stévart et al. 2019). In Fouta Dyal-
lon, there are still patches of forest (Wilson 1992) that
seem sufficient to facilitate the survival of endemic frog
species, such as the new Conraua or the endemic Odon-
tobatrachus species (Bare et al. 2015b; this study). The
conservation of preserved small areas can be key to the
survival of regional biodiversity and these isolated forests
that served as refugia in the past may also be more resis-
tant to future climatic changes (Hillers 2008). Conraua
kamancamarai occurs in classified forests (Konkouré
Fetto and Binti classified forests), i.e. areas designated
for sustainable management of forest resources (Bru-
giere and Kormos 2008). However, these forests have
historically been given low priority, have been illegally
degraded (Brugiere and Kormos 2008), being ineffective
in mitigating the effects of anthropogenic intervention on
vegetation change (Liu et al. 2017). A more rigorous pro-
tection of these forests and potentially upgrading of the
conservation status of these forests would be important
steps towards the preservation of the region’s biodiver-
sity, as well as for the long-term maintenance of this 1m-
portant water source for local people. The Fouta Djallon
is known as the “Water Tower” of West Africa, being the
principal water supply for several countries in the region.
For instance, Mauritania and Niger rely on this source
for 96% and 70% of their water, respectively (Descroix
et al. 2020).

With the description of Conraua kamancamarai, we
continue to refine the taxonomy of the West African slip-
pery frogs. However, there still remains work to be done.
Further data and analyses are required to re-evaluate the
larger C. alleni complex and to identify whether other
populations in the Fouta Dyallon, such as at Soyah (sub-
clade la; see Blackburn et al. 2020), might represent ad-
ditional undescribed species.

Zoosyst. Evol. 98 (1) 2022, 23-42

Acknowledgements

We thank Joseph Martinez (MCZ), Annemarie Ohler
(MNHN), Danny Meirte (MRAC), Peter S. Miller
(UWBM), as well as Claudia Koch, Dennis Rédder and
Wolfgang Bohme (ZFMK) for providing access to vouch-
er specimens under their care. Frank Tillack assisted with
access to specimens and data at ZMB and took the pho-
tos of the vouchers of the new species. Amanda Quezada
(Tropical Herping) helped with photo editing. Guillaume
Demare assisted with the map. We particularly thank our
assistants, Kaman Camara, Francois Gbémou, Blaise Pivi
and Ouo Ouo Kolié for their invaluable support in the
field. We gratefully acknowledge the Guinean ‘Ministere
en charge de |’ Agriculture Elevage Environnement Eaux
et Foréts, the ‘Ministere de |’ Environnement et du Dével-
oppement Durable’ and the ‘Ministere de |’ Enseignement
Supérieur et de la Recherche Scientifique’ for research, col-
lection and export permits (Certificat d’Origine N° 00136,
00139, 00602, 0000189, 0002707, 0002712, 0003227,
0003255; 0003312, 0003314; n° 061, 072, 177/MERS/
DNRST). Belén Ceollar and Virgile Lescure helped with
the translation of the abstract into French. We thank Alan
Channing, Vaclav Gvozdik and Werner Conradie for con-
structive feedback in their reviews of this manuscript.

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Appendix |

Additional Conraua specimens examined for qual-
itative comparisons. MCZ = Museum of Comparative
Zoology at Harvard University, Cambridge; MNHN =
Muséum national d‘Histoire naturelle, Paris; MRAC =
Musée Royal de |‘Afrique Centrale, Tervuren, Belgium;
UWBM = Burke Museum of Natural History and Cul-
ture, Seattle; ZFMK = Zoologisches Forschungsinstitut
und Museum Alexander Koenig, Bonn, ZMB = Museum
fur Naturkunde, Berlin.

Conraua alleni (sensu lato = all Conraua from the
western Upper Guinea forest zone, morphologically as-
signable to C. alleni). MCZ A-11991, subadult, holotype,
Liberia, Firestone Plantation No. 3, Du River, Liberia;
ZMB 90351, adult male, Guinea, Mont Béro Classified
Forest, 08°08'30.9"N, 08°34'09.6"W; ZMB 90443, adult
female, Sierra Leone, Tingi Hills, 08°53.4'N, 10°47.4'W;
ZMB 90426, adult female, Sierra Leone, Loma Moun-
tains Forest Reserve, 9°12.752'N, 11°08.623'W; ZMB
90178, adult female, Sierra Leone, Nimini Forest Reserve,
8°30.035'N, 11°08.800'W; ZMB 90388, ZMB 90389,
adult males; ZMB 90179, adult female, Liberia, Putu
Range, 05°39'44.6"N, 08°09'39.2"W, 306 m a.s.l.; ZMB
90304, adult male, Guinea, Fouta Dyallon, 10°49'13.1"N,
12°11'30.7"W, 513 m a.s.l.; ZMB 90357, adult male and
ZMB 90358, adult female, Ivory Coast, Haute Dodo,
04°59'14"N, 07°19'39"W; ZMB 90310, ZMB 90313, adult
females, ZMB 90311, ZMB 90312, adult males, Guinea,
Forét Classée de Ziama, 8°21'02.7"N, 9°25'01.8"W, 467 m
a.s.1.; ZMB 90325, adult female, ZMB 90326, adult male,
Guinea, Nimba Range, 07°38'56.3"N, 8°25'25.56"W, 680
m a.s.l.; ZMB 90388, ZMB 90389, adult males, Liberia,
Slabbert’s Ville, 05°39'44.6"N, 8°09'39.2"W, 316 ma.s.1.;
ZMB 90342, adult male, Liberia, Gola National Forest,
07°27'17"N, 10°41'52"W, 299 maz.s.l.

Conraua beccarii. ZFMK 15749-15750, Ethiopia, II-
lubator; MNHN 1933. 21, adult male, holotype of Rana
Griaulei Angel, 1934, Ethiopia, Gondar, 2,200 ma.s.1.

Conraua crassipes. ZMB 8360, holotype, “Abo”,
north of Douala, Cameroon; ZFMK 73216, Gabon,

41

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76588-9_1

Barrage de Kinguélé, Tchimbeélé; ZFMK 69351, 69353,
69354, Cameroon, Mt. Nlonako, Nguengue, Campsite;
ZMB 90400-90403, Gabon, Moukalaba-Doudou Na-
tional Park.

Conraua derooi. MRAC 112077—112078, paratypes,
Togo, Missahohe; MRAC 112079-112080, paratypes,
Togo, Missahohe; ZMB 71293, adult male and ZMB
71294, adult female, Ghana, Biakpa, 06°50.652'N,
00°25.280'E; ZMB 71298-71300, ZMB 71302 adult
males and ZMB 71301, adult female, Togo, Muissa-
hohe, 6°57.094'N, 0°33.878'E; UWBM:Herp 09599-—
09603, adult females and UWBM:Herp 09604, adult
male, Ghana, Volta Region, Adaklu-Anyigbe; MNHN
1978.2027, 1978.2029, adult males and MNHN
1978.2030, 1978.2031, adult females and MNHN
1978.2028, adult with unknown sex, Togo, Dangi Ati-
ba; MNHN 1993.2627, 1993.2629, adult males and
MNHN_ 1993.2630, 1993.2631, adult females and
MNHN 1993.2628 adult with unknown sex, Togo, Klo-
to; MNHN 1993.4084—1993.4087, Togo, Missahohe;
MNHN 1995.5726, 1995.5727, Togo, Kluto; MNHN
1987.2026, Togo, Dangi Atigba.

Conraua goliath. ZFMK 77927, 77928, 77930, 77932,
Cameroon, Mt. Nlonako, Ekomtolo, 500 m a.s.1.

Conraua robusta. ZMB 20085, holotype, Camer-
oon; ZMB 78427, Cameroon; ZMB 90174, Cameroon,
Manengouba Village, Mt. Manengouba; ZFMK 67288,
Cameroon, Bakossi Mts., Kodmin.

Conraua sagyimase. UWBM: Herp 5839, holotype,
adult male, Ghana, Eastern Region, Atewa Range
Forest Reserve, 06°13'57.79"N, 0°33'07.08"W, 633
m a.s.l.; ZMB 91136, paratype, adult female, Ghana,
Asiakwa South, 06°15'44.3"N, 0°33'18.8"W, 783 m
a.s.l.; ZMB 91137-91138, paratypes, adult females,
Asiakwa North, 06°16'16.1"N, 0°33'52.7"W, 814 m
a.s.1.; UWBM:Herp 5840-5843, paratypes, adult male
and female and two subadults, Ghana, Eastern Re-
gion, Atewa Range Forest Reserve, 06°13'57.79"N,
0°33'07.08"W, 633 ma.s.l.

zse.pensoft.net

42
Appendix 2

Conraua specimens examined for quantitative com-
parisons. ZMB = Museum fir Naturkunde, Berlin.

Conraua sp. from Soya, Fouta Djallon. ZMB 90341,
ZMB 90175, ZMB 90176, adult males, Guinea, Soya,
10°17'50.9"N, 11°56'32.4"W, 515 ma.s.l.

Conraua alleni. ZMB 90310, ZMB 90313, adult fe-
males, ZMB90311,ZMB 90312, adult males, Guinea, Forét
Classée de Ziama, 8°21'02.7"N, 9°25'01.8"W, 467 ma.s.L.;
ZMB 90325, adult female, ZMB 90326, adult male, Guin-

zse.pensoft.net

Neira-Salamea, K. et al.: A new Conraua from Guinea

ea, Nimba Range, 07°38'56.3"N, 8°25'25.56"W, 680 m
a.s.1.; ZMB 90388, ZMB 90389, adult males, ZMB 90390,
adult female, Liberia, Slabbert’s Ville, 05°39'44.6"N,
8°09'39.2"W, 316 ma.s.l.; ZMB 90427, ZMB 90428, adult
males, Guinea, Nimba Range, 07°42'82"N, 8°21'70"W,
848 m a.s.l.; ZMB 90342; adult male, Liberia, Gola Na-
tional Forest, 07°27'17"N, 10°41'52"W, 299 m a.s.1.; ZMB
90350, adult female, Guinea, Mont Béro Classified Forest,
08°08'30.9"N, 08°34'09.6"W, 622 maz.s.l.