A new critically endangered slippery frog (Amphibia, Conrauidae, Conraua) from the Atewa Range, central Ghana

Abstract

Forty-nine years after the last description of a slippery frog, we describe a seventh species of the genus Conraua. The new Conraua is endemic to the Atewa Range Forest Reserve, central Ghana, and is described based on genetic, bioacoustics, and morphological evidence. Recent molecular phylogenetic and species delimitation analyses support this population as distinct from nominotypical C. derooi in eastern Ghana and adjacent Togo. The new species is sister to C. derooi, from which it differs ~4% in the DNA sequence for mitochondrial ribosomal 16S. Genetic divergences in 16S to other species of Conraua range from 4–12%. The new species is distinguished morphologically from its congeners, including C. derooi, by the combination of the following characters: medium body size, robust limbs, lateral dermal fringing along edges of fingers, cream ventral color with brown mottling, the presence of a lateral line system, indistinct tympanum, the presence of inner, outer, and middle palmar tubercles, and two subarticular tubercles on fingers III and IV. We compare the advertisement calls of the new species with the calls from C. derooi and find that they differ by duration, frequency modulation, and dominant frequency. We discuss two potential drivers of speciation between C. derooi and the new species, including river barriers and fragmentation of previously more widespread forests in West Africa. Finally, we highlight the importance of the Atewa Range Forest Reserve as a critical conservation area within the Upper Guinean biodiversity hotspot.

Full text

ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by M. Vences: 9 Jun. 2021; published: 29 Jun. 2021 71
Zootaxa 4995 (1): 071–095
https://www.mapress.com/j/zt/
Copyright © 2021 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4995.1.4
http://zoobank.org/urn:lsid:zoobank.org:pub:8F2BAC17-6F60-4970-8B7F-61BE4D1F2948
A new critically endangered slippery frog (Amphibia, Conrauidae, Conraua) from
the Atewa Range, central Ghana
KARLA NEIRA-SALAMEA1, CALEB OFORI-BOATENG2,8, N’GORAN G. KOUAMÉ3, DAVID C. BLACKBURN4, GABRIEL H.
SEGNIAGBETO5, ANNIKA HILLERS6, MICHAEL F. BAREJ1, ADAM D. LEACHÉ7 & MARK-OLIVER RÖDEL1,*
1 Museum für Naturkunde Berlin – Leibniz Institute for Evolution and Biodiversity Science, Invalidenstraße 43, 10115 Berlin,
German
�
karla.neira@mfn.berlin, https://orcid.org/0000-0001-7835-6574
�
mfbarej@web.de, https://orcid.org/0000-0002-8743-2185
2 CSIR-Forestry Research Institute of Ghana, P. O. Box 63, Fumesua, Kumasi, Ghana
�
calebofori@gmail.com, https://orcid.org/0000-0001-9347-5977
3 Université Jean Lorougnon Guédé, UFR Environnement, Laboratoire de Biodiversité et Ecologie Tropicale, Daloa, BP 150, Côte
d´Ivoire
�
ngoran_kouame@yahoo.fr, https://orcid.org/0000-0003-3007-605X
4 Department of Natural History, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA
�
dblackburn@flmnh.ufl.edu, https://orcid.org/0000-0002-1810-9886
5 Laboratory of Ecology and Ecotoxicology, Faculty of Sciences, University of Lomé, BP 6057 Lomé, Togo
�
h_segniagbeto@yahoo.fr, https://orcid.org/0000-0002-4697-3671
6 Wild Chimpanzee Foundation (WCF), Liberia Office, FDA Compound, Whein Town, Mount Barclay, Montserrado County, Liberia
�
hillers@wildchimps.org, https://orcid.org/0000-0003-3179-8492
7 Department of Biology & Burke Museum of Natural History and Culture, University of Washington, Seattle, WA, USA
�
leache@uw.edu, https://orcid.org/0000-0001-8929-6300
8 EDGE of Existence Programme, Zoological Society of London, Regent’s Park, London NW1 4RY, UK.
*CorrespondanCe:
�
mo.roedel@mfn.berlin; https://orcid.org/0000-0002-1666-195X
Abstract
Forty-nine years after the last description of a slippery frog, we describe a seventh species of the genus Conraua. The new
Conraua is endemic to the Atewa Range Forest Reserve, central Ghana, and is described based on genetic, bioacoustics,
and morphological evidence. Recent molecular phylogenetic and species delimitation analyses support this population
as distinct from nominotypical C. derooi in eastern Ghana and adjacent Togo. The new species is sister to C. derooi,
from which it differs ~4% in the DNA sequence for mitochondrial ribosomal 16S. Genetic divergences in 16S to other
species of Conraua range from 4–12%. The new species is distinguished morphologically from its congeners, including
C. derooi, by the combination of the following characters: medium body size, robust limbs, lateral dermal fringing along
edges of fingers, cream ventral color with brown mottling, the presence of a lateral line system, indistinct tympanum, the
presence of inner, outer, and middle palmar tubercles, and two subarticular tubercles on fingers III and IV. We compare
the advertisement calls of the new species with the calls from C. derooi and find that they differ by duration, frequency
modulation, and dominant frequency. We discuss two potential drivers of speciation between C. derooi and the new
species, including river barriers and fragmentation of previously more widespread forests in West Africa. Finally, we
highlight the importance of the Atewa Range Forest Reserve as a critical conservation area within the Upper Guinean
biodiversity hotspot.
Key Words: Anura, biodiversity hotspot, conservation, integrative taxonomy, Upper Guinean Forest, West Africa
Introduction
The genus Conraua Nieden, 1908 comprises some of the most enigmatic African frog species, including the Goliath
Frog, Conraua goliath (Boulenger, 1906), the largest living species of frog. In contrast to the previous assignment of
some species to different genera (Gigantorana, Hydrobatrachus, Paleorana, Pseudoxenopus, Rana; compare Frost
2021), as well as the placement of Conraua in different families (compare Frost et al. 2006, Blackburn & Wake

NEIRA-SALAMEA ET AL.
72 · Zootaxa 4995 (1) © 2021 Magnolia Press
2011), the number of species formally described for the genus has remained stable in contrast to many other African
anuran genera. The taxonomy of Conraua was revised by Lamotte & Perret (1968), with only one other species, C.
derooi Hulselmans, 1972, described subsequently. However, recent molecular phylogenetic and species delimitation
analyses support that there remain several undescribed species of Conraua (Blackburn et al., 2020).
Currently six Conraua species are considered valid (Channing & Rödel 2019, Frost 2021): C. alleni (Barbour
& Loveridge, 1927) known from the western part of the Upper Guinean forests, i.e. Sierra Leone, through Guinea
and Liberia to western Ivory Coast and isolated populations in western Ghana (Barbour & Loveridge 1927, 1930;
Guibé & Lamotte 1958; Lamotte 1971; Hughes 1988; Rödel & Branch 2002; Rödel 2003; Rödel & Bangoura 2004;
Rödel et al. 2004; Hillers et al. 2008); C. derooi occurring in southern-central and south-eastern Ghana and adjacent
Togo (Schiøtz 1964; Hulselmans 1972; Segniagbeto et al. 2007, 2017; Hillers et al. 2009; Channing & Rödel 2019);
C. crassipes (Buchholz & Peters, 1875) with records from eastern Nigeria through Cameroon into the Democratic
Republic of Congo and Gabon (Perret 1966, Amiet 1990, Frétey & Blanc 2000, Lötters et al. 2001, Burger et al.
2004); C. goliath from south-western Cameroon into Equatorial Guinea (Perret 1966, Gartshore 1984, Frétey &
Blanc 2000, Schäfer et al. 2019); C. robusta Nieden, 1908 confined to mountains of western Cameroon and adjacent
south-eastern Nigeria (Perret 1966, Gartshore 1984, Lawson 1993); and C. beccarii (Boulenger, 1911) known from
mountain streams in Ethiopia and Eritrea (Largen & Spawls 2010). In western Cameroon, C. goliath, C. robusta,
and C. crassipes can occur in sympatry or even syntopy (Gartshore 1984, Lawson 1993, Plath et al. 2004, Herrmann
et al. 2005), but all other species have distinct non-overlapping ranges.
Surprisingly little is known about the biology of species of Conraua. All are aquatic, inhabiting forest streams
of different size and speed, from lowlands to higher altitudes (Lamotte & Perret 1968; Amiet 1989; Rödel & Branch
2002; Rödel 2003; Hillers et al. 2008, 2009; Largen & Spawls 2010; Segniagbeto et al. 2017). The most well-
studied species is the largest species, C. goliath, including its larval biology (Sabater-Pi 1985), parental behaviour
(Schäfer et al. 2019), diet (Perret 1957, 1960; Sabater-Pi 1985), and physiology (Hutchison 1998; for a summary of
the present knowledge see AmphibiaWeb 2020). Tadpoles of several other species of Conraua have been described
(Lamotte et al. 1959, Channing et al. 2012) as well as anecdotal accounts of behaviour of some species (Knoepffler
1985, Herrmann & Edwards 2006). The latter includes the way that species typically produce advertisement calls:
whistling with an open mouth (Amiet 1990, Amiet & Goutte 2017).
Only recently has it become clear that not only is our knowledge of the biology of this genus fragmentary, but
that some species may comprise cryptic species (Rödel & Branch 2002, Rödel 2003, Rödel & Bangoura 2004, Hillers
et al. 2008, Blackburn et al. 2020, Leaché et al. 2020). For instance, Conraua derooi was unexpectedly recorded
in the Atewa Range Forest in southern-central Ghana (Kouamé et al. 2007). This area comprises a unique montane
forest vegetation, not occurring anywhere else in West Africa (McCullough et al. 2007), and is separated by more
than 120 km, including the Volta River, from populations of C. derooi in eastern Ghana. While the population in
the Atewa Range was first considered to belong to C. derooi (Kouamé et al. 2007, Channing & Rödel 2019), recent
molecular phylogenetic studies (Blackburn et al. 2020, Leaché et al. 2020) strongly support these populations as
distinct from C. derooi and representing an unconfirmed candidate species (sensu Vieites et al. 2009). Following our
study of the morphology and advertisement calls of C. derooi and the population from the Atewa Range, it is clear
that the latter represents an undescribed species, which we describe herein.
Materials and methods
Morphology. We examined eight specimens from the population of Conraua in the Atewa Range of Ghana: three
from the collection of the Museum für Naturkunde Berlin, Germany (ZMB) and five from the collection of the
Burke Museum of Natural History and Culture (University of Washington, Seattle, USA; UWBM). The holotype
of the new species was included in the study by Blackburn et al. (2019). All individuals were preserved in 75%
ethanol.
For comparison, we examined specimens of all other nominal species of Conraua, including 11 specimens of
C. alleni, three C. beccarii, four C. goliath, four C. robusta, nine C. crassipes, and 34 C. derooi. These include four
paratypes of C. derooi from the collections of the Royal Museum for Central Africa (Tervuren, Belgium; MRAC)
and the Muséum national d’Histoire naturelle (Paris, France; MNHN), as well as the holotypes of C. crassipes
and C. robusta from the ZMB, the holotype of C. alleni from the Museum of Comparative Zoology, Cambridge
(Harvard University, Cambridge, USA; MCZ), and the holotype of Rana Griaulei Angel, 1934 (a synonym of C.

A NEW CONRAUA FROM GHANA Zootaxa 4995 (1) © 2021 Magnolia Press · 73
beccarii) from the MNHN. All comparative material is listed in Appendix 1. MOR measured the four C. derooi
paratypes, the C. alleni holotype and the R. griaulei holotype. All other data were collected by KNS, and for
statistical comparisons we excluded the data collected by MOR.
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
diameter), 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 center of nostril), eye to snout
distance (ES, from anterior edge of eye to tip of snout), internarial distance (IND, between center of nostrils),
tympanum diameter (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 III), finger III length (F3), forearm length (FLL). All measurements were taken with a
digital caliper (± 0.1 mm) and/or a dissecting microscope and are given in mm. They are summarized in Table A1
for the new species and Table A2 for C. derooi. Sex and maturity were assessed by examination of gonads through
an incision in the lateral body wall. Further qualitative morphological characters examined included tympanum
detectability (distinct/indistinct), webbing condition (complete/incomplete), head shape in lateral view (round/
pointed/truncated), relative length of fingers (i.e., comparative lengths were assessed by pressing fingers together
and fingers were numbered from preaxial to postaxial I to IV), presence of the interorbital stripe, color and shape
(curved/straight/slightly-curved) of the supra-tympanic ridge, belly and throat coloration in alcohol, and presence of
a lateral line system (see Lamotte & Perret 1968, Fritzsch et al. 1987; the lateral line system of adult Conraua will
be described in a separate publication). Concerning the nomenclature of these lines (infra-orbital line, supra-orbital
line, upper lateral line, lower lateral line, median lateral line, caudal lateral line, and jugular line), we followed
Escher (1925) and Shelton (1970). Coloration in preservation was only specified as being comparatively light or
dark. The variation of qualitative morphological characters between the new species and C. derooi is summarized in
Table 1. We were able to collect all data for all specimens, except for ZMB 91138 due to its poor condition. A short
morphological summary description of C. derooi is presented in the appendix (Appendix 2).
For statistical analysis, we only used adult frogs for which we could confirm maturity by inspecting the gonads,
which included six adult specimens of the undescribed Conraua and 19 C. derooi. Because males and females
in most species of Conraua cannot be distinguished by external morphology, we ran statistical comparisons for
both sexes together. Principal component analysis (PCA) was performed to reduce morphometric variables and
determine morphometric variation between individuals (Neff & Marcus 1980); and a discriminant function analyses
(linear discriminant analysis, LDA) was performed to determine the reliability of morphological quantitative traits
in assigning individuals to the species groups identified by Blackburn et al. (2020); the two groups correspond
to “derooi 1” (Atewa Range population) and “derooi 2” (nominotypical C. derooi) of Blackburn et al. (2020).
To visualize variation independent of body size, PCA was conducted on the residuals of linear regressions of 15
measured variables and SVL; we did not, however, “size-correct” the data used in the LDA of these same 16
variables. Both sets of analyses were carried out on log-transformed data. Because the tympanum was not visible in
all individuals, we did not include tympanum diameter (TD) or eye to tympanum distance (ETD) in these analyses.
Additionally, to explore the results of the PCA and LDA analyses further, we compared ratios of measurements that
resulted on high loadings in the PCA, performing T-tests or, alternatively, Mann–Whitney U-tests (MW-U) for non-
normal distribution. Statistical analyses were performed in R version 3.5.1 (R Core Team 2018) using the prcomp
function for PCA (with values scaled and centered) and lda function for LDA from the MASS package (v7.3-
51.3; Venables & Ripley, 2002), and for PCA visualization we used the function autoplot of the ggplot2 package
(Wickham 2016).
Bioacoustics. We analyzed three recordings of two individuals of the undescribed Conraua and seven recordings
of three individuals of C. derooi. Recordings of the new species were made by Jeremy Lindsell with an AudioMoth
autonomous recorder, and recordings of C. derooi were collected by AH with a Sony WM-D6C tape recorder and a
directional microphone (Sony ECM-Z157). Air temperatures were not taken at the time of recordings.
The advertisement calls were analyzed in Raven Pro 1.6.1 (Center for Conservation Bioacoustics 2019), with a
sampling rate of 44.1 kHz, FFT of 256 points, 16-bit resolution, overlap 50%, and Hann window type. To remove the
background noise, prior to analysis we set a filter around every call, selecting an area of 0.5 kHz above and below

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74 · Zootaxa 4995 (1) © 2021 Magnolia Press
TABLE A1. Measurements [mm] of the type series of Conraua sagyimase 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; UWBM= Burke Museum of Natural History and Culture, Seattle; ZMB= Museum für Naturkunde, Berlin.
Voucher Sex SVL HW HL SL ED EN ES IND IOD UEW TD ETD TL FL T4 HAL F3 FLL
UWBM:Herp
05839 m 71.9 26.7 25.6 7.9 8.1 4.6 9.0 5.2 5.0 5.7 4.0 2.5 32.4 43.4 26.8 17.4 11.2 12.7
ZMB 91136 f 54.0 18.8 18.2 5.6 5.4 3.7 6.8 4.7 3.7 4.4 3.0 3.7 25.6 34.5 22.7 13.5 7.1 10.4
ZMB 91137 f 54.1 19.2 17.3 5.5 6.0 3.3 6.8 5.1 4.8 3.7 4.0 2.5 25.9 34.7 23.6 12.8 7.9 10.1
ZMB 91138 f 66.7 24.1 23.0 6.2 6.4 5.0 7.4 4.9 5.0 4.6 4.4 4.6 31.1 44.0 28.6 16.2 9.7 11.9
UWBM:Herp
05840 m 57.0 20.4 20.3 6.0 5.9 3.9 7.5 4.4 4.6 4.6 3.7 3.5 26.0 30.8 17.8 13.9 8.4 10.7
UWBM:Herp
05841 f 53.4 18.8 18.6 5.3 5.8 3.6 7.0 4.1 4.2 4.5 4.1 3.5 25.1 34.9 24.4 14.0 8.8 11.1
UWBM:Herp
05842 s 47.9 16.7 16.4 5.1 5.0 3.0 5.8 3.6 3.3 4.3 3.4 2.5 21.8 28.0 20.0 11.6 7.4 8.9
UWBM:Herp
05843 s 44.0 15.4 14.7 4.7 4.4 2.9 5.6 3.9 3.2 3.3 2.8 2.4 20.3 26.9 18.9 10.9 6.4 8.0
mean f 57.1 20.2 19.3 5.6 5.9 3.9 7.0 4.7 4.4 4.3 3.9 3.6 26.9 37.0 24.8 14.1 8.4 10.9
sd f 6.5 2.6 2.5 0.4 0.4 0.7 0.3 0.4 0.6 0.4 0.6 0.8 2.8 4.6 2.6 1.5 1.1 0.8

A NEW CONRAUA FROM GHANA Zootaxa 4995 (1) © 2021 Magnolia Press · 75
TABLE A2. Measurements [mm] of adult Conraua derooi, m= male, f= female, 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; UWBM=
Burke Museum of Natural History and Culture, Seattle; MNHN= Muséum national d’Histoire naturelle, Paris; ZMB= Museum für Naturkunde, Berlin.
Voucher Sex SVL HW HL SL ED EN ES IND IOD UEW TD ETD TL FL T4 HAL F FLL
ZMB 71293 m 73.2 30.8 25.0 5.0 8.5 4.9 8.4 6.7 6.2 6.4 NN NN 34.7 41.9 24.8 18.0 11.5 13.1
ZMB 71298 m 100.0 40.8 37.6 10.3 10.1 6.6 12.5 7.0 9.4 7.1 6.9 11.9 44.9 58.9 42.4 23.5 13.6 17.8
ZMB 71299 m 96.8 38.8 35.2 8.3 8.3 6.6 10.8 6.9 9.0 6.5 6.3 10.0 45.0 57.5 40.8 22.0 11.9 17.2
ZMB 71300 m 100.0 40.3 36.9 9.5 8.5 6.5 11.1 7.1 9.1 7.4 5.9 11.9 44.5 51.9 37.2 22.5 12.3 20.1
ZMB 71302 m 80.2 30.5 26.2 6.6 6.7 4.5 9.9 5.6 7.2 5.5 4.6 6.2 37.5 50.0 34.7 18.4 10.7 16.6
UWBM:Herp 09602 m 75.9 28.4 27.7 7.8 7.6 4.7 8.9 5.3 5.6 5.0 4.5 2.0 34.6 45.7 27.2 17.3 10.7 12.8
UWBM:Herp 09604 m 97.7 38.3 32.8 8.1 9.2 5.5 9.3 6.6 8.2 7.1 6.4 4.1 42.9 51.5 38.1 20.7 11.8 17.2
MNHN 1978.2027 m 88.7 34.3 29.0 8.5 8.3 5.4 10.7 6.4 7.4 6.1 4.6 8.6 42.7 46.2 28.3 21.6 11.6 15.2
MNHN 1978.2029 m 80.9 28.8 25.5 7.3 8.1 5.0 8.5 5.5 6.4 5.5 6.3 5.6 36.3 45.7 30.2 17.9 10.7 13.5
MNHN 1993.2628 m 78.4 27.8 23.5 7.6 7.1 4.8 8.8 4.9 5.9 5.5 5.5 5.5 35.2 48.5 29.5 18.9 11.2 13.8
MNHN 1993.2629 m 78.7 27.2 24.0 7.8 7.1 5.0 9.2 5.6 5.9 5.8 4.9 5.4 34.8 46.7 29.5 18.6 10.9 14.2
UWBM:Herp 09599 f 92.3 35.1 31.8 8.2 9.3 5.6 10.9 5.7 7.3 7.4 NN NN 43.5 50.6 34.0 21.8 12.1 16.2
ZMB 71294 f 72.6 28.2 26.2 5.9 9.3 4.3 8.1 6.5 6.0 5.7 4.5 4.9 33.6 43.4 28.6 18.7 10.4 14.0
ZMB 71301 f 98.1 36.3 27.6 8.5 8.5 5.8 10.5 6.6 7.5 7.5 4.8 9.7 43.6 57.9 41.0 21.6 12.2 15.8
UWBM:Herp 09603 f 91.8 33.7 31.2 7.8 10.6 4.8 9.7 6.3 6.7 6.7 5.5 2.5 38.4 41.4 34.5 22.0 13.0 15.9
UWBM:Herp 09600 f 95.0 35.0 32.1 8.5 9.3 5.1 10.4 5.5 7.1 6.9 5.2 2.7 41.5 52.1 36.8 20.6 12.8 15.1
UWBM:Herp 09601 f 89.0 34.4 31.0 8.3 8.7 4.9 10.0 5.8 6.7 6.3 NN NN 43.0 50.0 34.8 20.8 11.7 15.0
MNHN 1978.2030 f 86.3 32.4 29.2 7.3 8.0 5.3 9.6 6.0 6.9 5.3 5.3 8.0 39.5 49.2 30.2 20.1 10.6 16.0
MNHN 1978.2031 f 88.6 35.6 32.6 9.9 8.6 5.8 5.3 6.6 6.8 6.0 6.3 9.3 43.3 45.8 26.6 20.4 12.0 16.4
mean f 89.2 33.8 30.2 8.0 9.1 5.2 9.3 6.1 6.9 6.5 5.3 6.2 40.8 48.8 33.3 20.7 11.8 15.6
sd f 7.7 2.6 2.3 1.1 0.8 0.5 1.8 0.4 0.5 0.8 0.6 3.3 3.5 5.2 4.7 1.1 0.9 0.8
mean m 86.4 33.3 29.4 7.9 8.1 5.4 9.8 6.1 7.3 6.2 5.6 7.1 39.4 49.5 33.0 19.9 11.5 15.6
sd m 10.43 5.345 5.295 1.386 0.985 0.807 1.294 0.791 1.44 0.8 0.893 3.333 4.554 5.188 5.916 2.166 0.866 2.342

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76 · Zootaxa 4995 (1) © 2021 Magnolia Press
the maximum and minimum frequency of the call. We measured six parameters: call duration, dominant frequency
(peak frequency), low frequency, high frequency, bandwidth 90%, and we calculated the frequency modulation by
subtracting the frequency at the beginning of the frequency at the end of the call. Temporal parameters are given
in seconds (s) and spectral parameters in kilohertz (kHz). Description and terminology of calls follow Köhler et al.
(2017).
Phylogenetic and genetic data. We relied on the recent phylogeny of the genus Conraua (Blackburn et al.
2020), which includes mitochondrial and nuclear sequences of the new species described below and all other taxa
investigated for comparisons. Phylogenetic inference methods (Bayesian and Maximum Likelihood analyses) are
detailed in Blackburn et al. (2020).
Field data. Field observations of the new species were collected by three of the co-authors (COB, NGK, ADL).
In particular, COB has studied the new species at all known sites.
TABLE 1. Comparison of qualitative morphological characters of Conraua sagyimase sp. nov. and Conraua derooi (for
morphological measurements compare Tabs. A1-A2), including characters important for diagnosing other species in the
genus Conraua.
Characters Conraua sagyimase sp. nov. Conraua derooi
Relative length of fingers III>IV>II≈I III>IV>II>I
Snout shape rounded rounded
Interorbital stripe mostly present mostly absent
Supratympanic fold slightly curved to arm curved to arm
Lateral line system present present
Belly coloration light or light with dark mottling light or light with dark mottling
Throat coloration light with dark mottling light with dark mottling
Tympanum visibility indistinct indistinct
Webbing complete complete
Results
Molecular and morphological comparisons. The multi-locus phylogeny and species delimitation analyses of
Conraua presented by Blackburn et al. (2020) strongly support that the population of Conraua from the Atewa
Range and C. derooi form a well-supported clade. Together, these two species may form a lineage that is nested
within the several lineages currently referred to as C. alleni, which also contains one or more undescribed species;
these will be dealt with in separate publications. Uncorrected p-distances of the mitochondrial 16S rRNA fragment
(569 bp) between the population from Atewa Range and all other currently recognized or candidate species of
Conraua equaled or exceeded 4% (Blackburn et al. 2020). A value of 3% difference in this barcode gene is usually
regarded as indicating candidate species in frogs (Fouquet et al. 2007, Vieites et al. 2009). Genetic distances of
the population from Atewa to C. derooi from western Ghana and Togo was 4%, 4–7% to all populations of the C.
alleni-complex, 9% to C. goliath and C. robusta, and 11–12% to C. beccarii and C. crassipes (Blackburn et al.
2020). No haplotypes were shared between the population from Atewa and nominotypical C. derooi for any of the
nuclear gene sequences generated by Blackburn et al. (2020), including for DISP2, FICD, KIAA2013, SPEN, and
SVEP (1 specimen of each population) as well as POMC and RAG1 (three specimens from Atewa and four from
C. derooi).

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FIGURE 1. Conraua derooi from the type locality Misahöhe, Togo (a – c) and Biakpa in the Volta area (d), Ghana; a: adult
male, note bulging neck and posterior part of head, b & c: adult females; d: half-grown specimen. Specimens not collected.
FIGURE 2. Principal component analyses based on 15 size-corrected morphometric variables; red = Conraua derooi, violet =
Conraua sagyimase sp. nov., triangle = females, circle = males.

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FIGURE 3. a) Spectrogram and oscillogram of advertisement call of male Conraua sagyimase sp. nov. from Atewa Range
Forest, individual not collected (call 1); b) Spectrogram and oscillogram of advertisement call of male Conraua derooi from
Biakpa, Ghana, individual not collected. FFT = 256 points.
As the molecular phylogenetic results indicated that topotypical populations of C. derooi are sister to the
populations from Atewa, we restrict our detailed morphological comparisons to these two taxa (for morphological
comparisons with other Conraua species see ‘diagnosis’ below). Measures are summarized in Tables A1–A2. A
brief summary of the morphology of C. derooi is presented in the appendix (Fig. 1; Appendix 2; compare Tab. 1).
PCA analysis revealed differences among the Atewa frogs and C. derooi (Fig. 2). Four principal components with

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eigenvalues >1.0 (PC1–PC4) accounted for the 68.9% of the total variation. An ordination of PC1 versus PC2
demonstrates that the undescribed Conraua and C. derooi are roughly separated along PC1, which accounted for
27.6% of the variation and loaded negatively and most strongly for hand length and finger III length. The highest
loadings of PC2 were head width and head length (both negative), which account for 16.4% of the variation. PC3
was dominated by eye diameter and toe IV length; PC4 was predominately impacted by snout length and eye-nostril
distance. Although separation of the two taxa on the PCA axes was not complete (Fig. 2), a linear discriminant
analyses (LDA) of the morphometric characters could successfully separate our species (as identified from molecular
results), assigning correctly all individuals to the respective taxa. Loadings, eigenvalues, and percentage of variance
explained by the PCA are summarized in Table 2.
TABLE 2. Character loading, percentage (%) and cumulative percentage of the variance of morphological measures
in a Principal Component Analyses (PCA) with eigenvalues > 1.0, (PC) I–IV; PCA was applied to the residuals from
regressions between all measurements and SVL; variables with the highest loadings are given in bold; for abbreviations
of morphological measures see ‘material and methods’ section.
Variable PC1 PC2 PC3 PC4
HW/SVL 0.12 -0.55 0.02 -0.17
HL/SVL -0.11 -0.44 -0.09 0.11
SL/SVL -0.09 0.13 -0.03 0.61
ED/SVL -0.28 -0.06 -0.45 -0.21
EN/SVL -0.27 -0.14 0.20 0.34
ES/SVL -0.23 0.25 0.17 -0.30
IND/SVL -0.37 -0.20 0.02 -0.06
IOD/SVL 0.03 -0.38 0.39 -0.28
UEW/SVL -0.28 0.02 -0.23 -0.28
TL/SVL -0.21 -0.25 0.18 0.28
FL/SVL -0.28 0.23 0.41 0.08
T4/SVL -0.11 0.25 0.42 -0.29
HAL/SVL -0.43 0.00 -0.13 0.01
F3/SVL -0.38 0.09 -0.25 -0.03
FLL/SVL -0.29 -0.19 0.24 -0.01
Eigenvalue 4.15 2.46 2.02 1.70
%27.68 16.42 13.47 11.34
Cumulative % 27.68 44.10 57.57 68.91
We additionally checked with T-test, or Mann–Whitney U-tests (MW-U) for morphological ratios which differed
between the two taxa and identified the following as significant: SL/HL (t = 2.71, d.f. = 16.17, p < 0.05), IND/IOD
(t = -4.070, df = 10.84, p < 0.01), HAL/SVL (t = 3.21, d.f. = 10.27, p < 0.01, IN/HW (W = 8, p < 0.01), ED/HW (t =
3.81, d.f. = 15.44, p < 0.01), and ES/HW (W = 103, p < 0.01). Several ratios were not significantly different between
C. derooi and frogs from the Atewa Range: IND/HW (W = 106, p > 0.01), IOD/HW (t = 0.29, d.f. = 6.05, p > 0.05),
and TL/SVL (t = 1.4766, d.f. = 14.89, p > 0.05).
Bioacoustics. The advertisement calls of the population from Atewa Range and C. derooi have a similar
structure: a whistle-like tonal call with an ascending frequency modulation. However, most parameters were
noticeably different between the two taxa: frequency modulation in the call of the Atewa population (mean = 0.92
kHz, sd ± 0.4, N = 3) was higher than in C. derooi (mean = 0.54 kHz, sd ± 0.21, N = 7), call duration was shorter
in the Atewa population (mean = 0.5 s, sd ± 0.08, N = 3) than in C. derooi (mean = 1.13 s, sd ± 0.26, N = 7), and
the dominant frequency was lower in the Atewa population (2.47 kHz, sd ± 0.2, N = 3) compared to C. derooi (3.27
kHz, sd ± 0.28, N = 7) (Fig. 3, Table 3).

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TABLE 3. Advertisement call parameters of Conraua sagyimase sp. nov. (Cs) and C. derooi (Cd; several calls per
individual available; compare Fig. 3). Given are: call number (call#); individual, minimum frequency (minFr) in Hz;
maximum frequency (maxFr) in Hz; bandwidth 90% (FrBW 90%) in Hz; call duration 90% (CD 90%) in seconds,
dominate frequency (DoFr) in Hz; frequency modulation (FrMod) in Hz. For all call parameters, minimum, maximum,
mean, and standard deviation values are provided.
Call# Individual minFr (Hz) maxFr (Hz)
FrBW 90%
(Hz) CD 90% (s)
DoFr
(Hz) FrMod (Hz)
1 Cs1 1.12 3.16 0.86 0.42 2.24 1.38
2 Cs2 1.63 3.42 0.69 0.49 2.58 0.69
3 Cs2 1.66 3.48 0.69 0.58 2.58 0.69
Conraua
sagyimase
sp. nov.
mean 1.47 3.36 0.75 0.5 2.47 0.92
max 1.66 3.48 0.86 0.58 2.58 1.38
min 1.12 3.16 0.69 0.42 2.24 0.69
sd 0.31 0.17 0.1 0.08 0.2 0.4
1 Cd1 2.38 3.57 0.52 1.43 3.1 0.34
2 Cd1 2.37 3.37 0.52 0.92 2.93 0.17
3 Cd1 2.49 3.58 0.52 1.46 3.1 0.52
4 Cd2 2.16 3.78 0.69 1.26 3.1 0.69
5 Cd3 2.65 4.11 0.34 0.79 3.62 0.69
6 Cd3 2.03 4.14 0.52 1.06 3.62 0.69
7 Cd3 2.04 4.09 0.34 1 3.45 0.69
Conraua
derooi
mean 2.3 3.81 0.49 1.13 3.27 0.54
max 2.65 4.14 0.69 1.46 3.62 0.69
min 2.03 3.37 0.34 0.79 2.93 0.17
sd 0.23 0.31 0.12 0.26 0.28 0.21
Based on the phylogenetic and species delimitation analyses presented by Blackburn et al. (2020) and the
morphological and acoustic differences presented here, we recognize the populations from the Atewa Range Forest
Reserve as a species new to science. This new species is separated by more than 120 km from C. derooi, including
a well-known biogeographic barrier, the Volta River.
Description of the new species
Conraua sagyimase sp. nov.
Figs. 4-7
Holotype. UWBM:Herp 5839 (field #: ADL 3858, GenBank #: 16S: MT669421, DISP2: MT669442, FICD:
MT669459, KIAA2013: MT669477, POMC: MT669511, SPEN: MT669573, SVEP: MT669590, TYR: MT669624),
adult male, Ghana, Eastern Region, Atewa Range Forest Reserve, 06º13’57.79” N, 0º33’07.08” W, 633 m asl; 11
May 2011, leg. Adam D. Leaché. & Caleb Ofori-Boateng.
Paratypes. Four females, one male, two subadults; all from Atewa Range Forest, Ghana; ZMB 91136 (field #:
JP 0041.1), adult female, Asiakwa South, 06º15’44.3” N, 0º33’18.8” W, 783 m asl, 11–16 June 2006, leg. N’goran
Germain Kouamé & Caleb Ofori-Boateng; ZMB 91137–91138 (field #: JP 0041.2–3), adult females, Asiakwa North,

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06º16’16.1” N, 0º33’52.7” W, 814 m asl, 17–22 June 2006, leg. N’goran Germain Kouamé & Caleb Ofori-Boateng;
UWBM:Herp 5840 (field #: ADL 3859), adult male, UWBM:Herp 5841 (field #: ADL 3860), adult female, UWBM:
Herp 5842 (field #: ADL 3861), subadult, UWBM:Herp 5843 (field #: ADL 3862), subadult, 25 May 2011, other
data as holotype.
FIGURE 4. Conraua sagyimase sp. nov., male holotype (UWBM:Herp 5839) in lateral, dorsal, and ventral view. Scale =
distance between two small strokes: 1 mm.

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FIGURE 5. Conraua sagyimase sp. nov. male holotype (UWBM:Herp 5839) ventral views of throat and arms, right hand; and
right foot. Scale = distance between two small strokes: 1 mm.
Diagnosis. The new species generally resembles other members of the genus Conraua Nieden, 1908. Conraua
sagyimase sp. nov. is the smallest species of its genus and a mid-sized (SVL of adults: 53–89 mm) aquatic frog,
relative to other frog species (see Womack & Bell 2020). It has smooth dorsal skin, covered with scattered small,
rounded warts; skin on belly smooth; large and protruding eyes, positioned latero-dorsally; three odontoid projections
on lower jaw, one on the symphysis and one on each side of the central one on the dentary; fully webbed feet, i.e.,
webbing extends to the end of the last phalange of toe, disc remaining free of web.

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FIGURE 6. Four individuals of Conraua sagyimase sp. nov. from the Atewa Range Forest Reserve, southern Ghana (a: photo
by Piotr Naskrecki); specimens not collected.
Conraua sagyimase sp. nov. differs in the 16S sequences at least by 4% from other species in the genus, and
its closest relative is C. derooi (see above and Blackburn et al. 2020). Adult body size varies substantially across
species of Conraua (numbers reported are maximum values): Conraua sagyimase sp. nov. (females SVL = 66.7

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mm, males SVL = 71.9 mm) is smaller than C. derooi (females 98 mm, males 100 mm), C. robusta (females 122
mm, males 140 mm), C. goliath (females 220 mm, males 340 mm), and C. beccarii (155 mm). The new species is
similar in size to populations of the C. alleni (females 86.6 mm, males 91.9 mm) and C. crassipes (females 76 mm,
males 81 mm) (see Channing & Rödel 2019). Conraua sagyimase sp. nov. can be distinguished from C. goliath by a
more rounded snout (pointed in C. goliath), the absence of short dorsal skin ridges (present in C. goliath), a creamy
venter with brown mottling (yellow venter in C. goliath), by relatively larger odontoid projections on the lower
jaw, the presence of a lateral line system (absent in C. goliath), and an indistinct tympanum (distinct in C. goliath).
Conraua sagyimase sp. nov. differs from C. crassipes by a cream-colored belly with brown mottling (white or cream
in C. crassipes), an indistinct tympanum (distinct in C. crassipes), the presence of a lateral line system (absent in C.
crassipes) and lacking a dermal fold near the elbow (present in C. crassipes). Conraua sagyimase sp. nov. differs
from C. beccarii by the absence of a transverse fold behind the eyes and across the interorbital region (present in
C. beccarii), and in having a head that is as wide as long (wider than long in C. beccarii; adult C. beccarii males
develop an extremely wide and flat skull, see Lamotte & Perret 1968, Largen & Spawls 2010, Paluh et al. 2020).
Conraua sagyimase sp. nov. differs from C. robusta by having a head that is as wide as long (wider than long in C.
robusta; see Lamotte & Perret 1968), having non-expanded finger tips (slightly expanded in C. robusta), and the
presence of a lateral line system (absent in C. robusta). Conraua sagyimase sp. nov. differs from individuals of C.
alleni sensu lato by having a wider lateral dermal fringing along edges of fingers (narrower fringes in C. alleni sensu
lato), and large and oval toe discs (small and rounded in C. alleni sensu lato). Finally, Conraua sagyimase sp. nov.
differs from C. derooi by having a slimmer body and limbs (more robust body and limbs in C. derooi), a slightly
curved supratympanic fold (distinctly curved in C. derooi), two subarticular tubercles on finger III (one in C. derooi)
and lacking a swollen postoccipital and suprascapular region in adult males (swollen in C. derooi; Fig. 1a).
Description of the holotype (Figs. 4–5; measurements in mm). Adult male; slightly dorsoventrally flattened,
short and rounded body; snout rounded in dorsal view, rounded in lateral view, with upper lip slightly projecting
forward; SVL 71.9; head width 26.7, approximately equal to head length 25.6; head length 36% of SVL; snout
length 7.9, 31% of head length; eye–nostril distance 4.6; eye–snout distance 9.0; internarial distance 5.2, slightly
larger than interorbital distance; nostrils protuberant, directed dorsolaterally, visible in lateral and dorsal view; large
protruding eyes, projecting laterally beyond margins of head in dorsal view; eyes projecting above dorsal margin of
head in lateral view; eye diameter 8.1, twice as large as horizontal diameter of tympanum 3.9; upper eyelid width
5.7, 114% of interorbital distance; eye–tympanum distance 2.5; tympanum indistinct; interorbital distance 5.0, 62%
of eye diameter; canthus rostralis distinct and rounded; loreal region concave; slightly curved supratympanic fold
extending from posterior edge of eye to shoulder; upper lip slightly protruding; vocal sacs absent; premaxillary and
maxillary teeth are slender and pointed, teeth are long and short in premaxilla and short in maxilla, three odontoid
projections on lower jaw, one at symphysis and one to each side on dentaries.
Forelimbs moderately robust; forearm length 12.7, 73% of hand length, 17.4; outer and middle palmar tubercles
barely visible, smaller than inner palmar tubercle; supernumerary tubercles absent; one subarticular tubercle on
fingers I, II, two subarticular tubercles on fingers III and IV; subarticular tubercles absent on the base of fingers, tips
of fingers non-expanded, rounded; lateral dermal fringing along edges of fingers, wider on finger III; relative length
of fingers: III>IV>II≈I, , finger III length 11.1; no webbing between fingers.
Hind limbs moderately robust; crus length 32.4, 45% of SVL; foot including longest toe 43.4, 60% of SVL;
elongated, prominent oval inner metatarsal tubercle; outer metatarsal tubercle absent; supernumerary plantar
tubercles absent; subarticular basal tubercles absent; subarticular penultimate tubercles prominent, ovoid in dorsal
view; subarticular distal tubercles present in toes III, IV and V; toe tips with large oval discs; relative lengths of toes:
VI>III>V>II>I, toe IV length 26.7; webbing complete, i.e., to the end of the last phalanx of toe; dermal fringing on
outer surfaces of toe I and V forming lateral skin folds.
Skin texture on dorsal parts of head, body, flanks, and limbs smooth with scattered, small, rounded warts; inner
surface of upper arm smooth; dorsal surface of crus with 17 rows of longitudinal ridges; ventral skin smooth, throat
with longitudinal folds; a post-gular (thoracic) fold extending to level of forelimbs insertion; tarsal fold distinct.
The lateral line system comprises a jugular line, upper lateral line, lower lateral line, median lateral line, and caudal
lateral line, all distinct; an infra-orbital line, and a supra-orbital line are present but indistinct.
Coloration in preservative (after nine years in 75% ethanol). Dorsum brown with dark-brown spots and
scattered small cream warts; abundant cream spots on dorsolateral surface, flanks, and shoulders; lateral surface of
lips cream with brown spots, tip of snout mottled cream with brown; cream spots on interorbital region, shaping

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diffuse interorbital stripe; nostrils cream. Dorsal surface of arms and legs cream with brown spots; longitudinal
ridges on legs cream; dorsal surface of fingers I and II cream; fingers III and IV mottled cream with brown; throat
cream mottled in brown; belly cream with brown mottling on anterior part, cream on posterior part; ventral surface
of hands brown, edges of fingers cream; ventral surface of legs, feet and toes cream with almost imperceptible
mottled brown; toe IV and V with brown spots; discs brown; area surrounding cloaca brown with many cream
warts.
FIGURE 7. Conraua sagyimase sp. nov. from the Atewa Range Forest Reserve, southern Ghana (photos by Alan Channing);
specimen not collected.

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Variation. Overall the paratypes are similar to the holotype in external appearance and coloration (see Table
A1). Size (SVL) of adult females ranges from 53.4–66.7 mm, thus being smaller than male SVL, ranging from 57.0–
71.9 mm. Additional field measures of 49 individuals, with unidentified age and sex, revealed a maximum SVL of
89 mm (range: 52–89 mm; x ± sd: 65.9 ± 10.5 mm). The color of dorsal surfaces varied from predominantly brown
(UWBM:Herp 5839) to dark brown (UWBM:Herp 5841), with many dark spots (UWBM:Herp 5840, 5841; ZMB
91136). All paratypes exhibited more dark spots and fewer cream dots on dorsum than the holotype. Some specimens
exhibit a continuous and straight interorbital stripe (UWBM:Herp 5840–5843; ZMB 91136), in other specimens the
interorbital line was composed of many light spots (UWBM:Herp 5839; ZMB 91137) and not perfectly straight. In
some specimens the tympanum was distinct (UWBM:Herp 5841, 5842), in others it was hidden and only visible
under particular illumination (UWBM:Herp 5839, 5840, 5843; ZMB 91136, 91137). This variation is probably due
to preservation differences. Throat pigmentation varies among specimens, some (ZMB 91136, 91137; UWBM:Herp
5841) have dark pigmentation and others are lighter; however, all specimens exhibited some throat pigmentation.
The belly was generally uniform cream (UWBM:Herp 5840–5843); however, two specimens (ZMB 91136, 91137)
exhibit darker pigmentation, and the holotype (UWBM:Herp 5839) showed a slightly mottled brown.
FIGURE 8. Inset shows a map of West Africa showing the location of the Atewa Range in Ghana (upper left), known localities
of Conraua sagyimase sp. nov. are shown in red and know localities of Conraua derooi in yellow. Stars indicate type localities.
Altitudinal range is indicated with shading from lowlands (light) to high elevation (dark). Sources: OpenStreetMap (2020), U.S.
Geological Survey (2020).
The presence of vomerine teeth and shape of tongue was not possible to examine in the holotype without breaking
the mandibles. However, we could check for these characters in two paratypes. ZMB 91136 and ZMB 91137 possess
pointed vomerine teeth, and a tongue that bifurcates into rounding lobes, separated by a wide U-shaped indentation.
The tongue is attached to the floor of the mouth along its anterior third. Compared to the holotype the lateral line
system was more distinct in some paratypes (UWBM:Herp 5841, ZMB 91136, ZMB 91137).

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FIGURE 9. Habitat from Conraua sagyimase sp. nov. in the Atewa Range Forest Reserve (a–b), southern Ghana (photos:
courtesy of Piotr Naskrecki); and type locality of Conraua derooi, Misahöhe, Togo (c–d).

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FIGURE 10. Habitat from Conraua sagyimase sp. nov. in the Atewa Range Forest Reserve, southern Ghana.
Life coloration (based on photos of uncollected individuals, Figs. 6–7). Dorsum light to dark brown with light
spots and scattered cream warts; lips with same coloration as dorsum or lighter; flanks light to dark brown with
lighter mottling; dorsal surfaces of arms and legs with scattered cream warts; rows of parallel longitudinal ridges on
legs cream, lighter than body; iris gold; throat cream, mottled in brown; belly cream with brown mottling; ventral
surfaces of legs and feet brown with cream mottling; ventral surface of hands brown; edges of fingers cream; ventral
surface of toes brown.
Acoustics. The advertisement of Conraua sagyimase sp. nov. consist of a whistle-like tonal call with ascending
frequency modulation 0.69 – 1.38 kHz (mean ± sd, 0.92 ± 0.4 kHz, N = 3; Fig. 3). The call has a duration of 0.42
– 0.58 seconds (mean ± sd, 0.5 ± 0.08 s, N = 3). Dominant frequency ranges from 2.24 – 2.58 kHz (mean ± sd, 2.47
± 0.2 kHz, N = 3), minimum frequency 1.12 – 1.66 kHz (mean ± sd, 1.47 ± 0.31 kHz, N = 3) kHz, and maximum
frequency 3.16 – 3.48 kHz (mean ± sd, 3.36 ± 0.17 kHz, N = 3). The bandwidth 90% ranges from 0.69 – 0.86 kHz,
(mean ± sd, 0.75 ± 0.1 kHz, N = 3) (Fig. 3). Call parameters are summarized in Table 3.
Range. Conraua sagyimase sp. nov. is so far known from only five streams in the northern part of the Atewa
Range Forest Reserve in eastern Ghana (Fig. 8). However, we suppose that populations may also exist in the
southern part of the Atewa Range Forest Reserve. Lowland forest areas in Ghana west of Atewa are inhabited by
Conraua alleni senso lato, the region east of the Volta River is inhabited by C. derooi. We thus assume that the new
species is endemic to the Atewa Range.
Life history. The new species occupies relatively pristine upland evergreen forest habitats within an elevation
range of ~ 500–750 m asl. The frogs occur in rocky, clear, generally fast-flowing streams and waterfalls, although
some individuals have been recorded in slow-flowing streams. Stream widths range from one meter to over six
meters (Figs. 9–10).
Conraua sagyimase sp. nov. is highly aquatic, spending over 95% of the time in water. Some individuals however,
have been found at the banks of streams both day and night. One individual was recorded nine meters away from the
nearest stream. Along the streams, individuals perch on logs, rocks, sand, and in some cases on artificial drainage
systems such as culverts. Although usually solitary and patchily distributed, adult frogs occasionally congregate

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in small but deep pools (> 2 m) within streams, probably for feeding or mating. Habitat preferences are probably
related to prey availability (as prey may also be specialized to these habitats) or microclimatic conditions, but
this requires further investigation. Tadpoles have usually been found in shallow pools within streams, co-existing
sometimes with tiny fishes. We have found higher number of frogs in streams bordered by large trees (diameter at
breast height > 20 cm), and dense canopy coverage (> 70%), but we have also recorded individuals in streams with
overgrown bushes, suggesting the species could thrive in moderately degraded habitats.
In a recent population assessment in two streams (COB, unpubl. data), 433 individuals were marked over
approximately 5 km river lengths. Most streams did not have enough individuals to make estimates of population
feasible based on mark-recapture techniques. Based on the field counts we assume that more than 1000 individuals
could occur throughout the Atewa Hills Forest Reserve.
The female paratypes contain large ovarian eggs with a diameter of 1.5 mm (ZMB 91136), 3.2 mm (ZMB 91137)
and 2.5 mm (ZMB 91136). The eggs have a yellowish and dark pole. We assume deposition of aquatic eggs.
Threat status. The new species, although not yet officially assessed for the IUCN Red List, should be regarded
as Critically Endangered (CR), compare discussion and IUCN (2012).
Etymology. The name of the new species has been chosen in order to honor the people of the Sagyimase
community. This small community has supported the research of COB and the anti-mining campaigns during the
early 2006–2007. We hope that the naming of this endemic species will further encourage this community in their
fight for an intact Atewa Range. The species epithet is used as an invariable noun in apposition to the generic name.
As English common name, we suggest Atewa Slippery Frog, and as Akan common name we suggest kwaeɛ mu
nsutene apɔnkyerɛne, meaning the frog of the forest streams.
Discussion
With the description of Conraua sagyimase sp. nov., we confirm a presumed candidate species (Blackburn et al.
2020, Leaché et al. 2020) that is endemic to the Atewa Range Forest Reserve in southeastern Ghana. Morphological,
acoustic, and molecular data support the distinctiveness of Conraua sagyimase sp. nov. from other congeners,
becoming the seventh species currently regarded as valid in the genus.
The 16S barcode gene, usually used for comparisons in anurans, revealed uncorrected p-distances between
Conraua sagyimase sp. nov. and all other species of the genus exceeding 4%. These values are comparable to
species-level distances in other West and Central African frog genera (e.g., Barej et al. 2010; Rödel et al. 2011,
2012). Although the sample size of call recordings was small and we had no temperature values available for these
recordings, the differences were large enough to justify the recognition of two distinct species. In particular frequency
modulation of the advertisement call of C. sagyimase sp. nov. was higher, the call was shorter, and the dominant
frequency was lower than in calls from C. derooi. Finally and although Conraua species are morphologically very
similar (Lamotte & Perret 1968, Channing & Rödel 2019), the new species can also be distinguished morphologically
from other congeners (see diagnosis; Lamotte & Perret 1968; Channing & Rödel 2019). Conraua sagyimase sp.
nov. is a small species compared to congeners. For instance, maximum female body size (SVL) is 76 and 98 mm in
C. crassipes and C. derooi, respectively (sensu Channing & Rödel 2019). In contrast two female paratypes (ZMB
91136 and UWBM:Herp 05841) of C. sagyimase sp. nov. were gravid but only had body sizes of 54.0 mm and 53.4
mm, respectively, which indicates that these relatively small-sized individuals were already adult.
A previously published phylogeny of the genus already confirmed C. derooi as the closest relative of the new
species (Blackburn et al. 2020). The non-overlapping ranges of both species suggest that allopatric speciation may
have played a role in the evolutionary divergence of these two frog species. The estimated divergence between C.
derooi and C. sagyimase sp. nov. likely occurred during the late Miocene and Pleistocene (3–8 Myr; Blackburn
et al. 2020). Two possible mechanisms that may have both promoted the splitting include fragmentation of an
ancestral widespread forest-restricted species, associated with a reduction of forest cover during alternating cycles
of climatic change (Couvreur et al. 2021), and / or the lack of gene flow due to the large Volta River separating the
Atewa Range from the Togo-Volta Highlands (Wallace 1852, Penner et al. 2011). Speciation related to reduction
and fragmentation in forest cover has been suggested in many West and Central African taxa (e.g., frogs: Jongsma
et al. 2018, fruit bats: Hassanin et al. 2015, legumes: Duminil et al. 2013). Similarly, divergence across Lake Volta
has already been found in other pairs of sister species, such as Hyperolius picturatus and H. baumanni (Portik et

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90 · Zootaxa 4995 (1) © 2021 Magnolia Press
al. 2019) [although there exists a potential photographic record of the later species from the Atewa Range], found
from Sierra Leone to central Ghana, and from eastern Lake Volta in Ghana to western Togo, respectively (Channing
& Rödel 2019). Phrynobatrachus afiabirago, described from the Atewa Range and another southern Ghanaian
locality, has its closest relatives in the western part of the Upper Guinean forest zone (Ofori-Boateng et al. 2018).
These patterns are also found more generally across western Africa in other frogs such as within West African
Astylosternus (Rödel et al. 2012) or Acanthixalus (Rödel et al. 2003, Portik et al. 2019), as well as in sister pairs
of other vertebrate species (e.g., birds: Fuchs & Bowie 2015, shrews: Jacquet et al, 2014, rodents: Nicolas et al.
2010).
All other populations of Conraua recorded in lowland forests of western and eastern Ghana correspond to C.
alleni sensu lato and C. derooi, respectively (Hillers et al. 2009; this paper; G.B. Adum & M.-O. Rödel, unpubl.
data). It is very likely that the new species described here is endemic to the Atewa Range. Its distribution seems
bound to clear streams at higher altitudes within the Atewa Range Forest Reserve. This site is threatened by many
human activities such as farming, mining, logging (Ghartey-Tagoe et al. 2021) and wildfires that have well-
documented negative effects on wildlife in this area (e.g., dragonflies; Seidu et al. 2017; butterflies; Addo-Fordjour
et al. 2015) that were recently summarized by Lindsell et al. (2019). As such, Conraua sagyimase sp. nov. qualifies
to be classified as Critically Endangered, following the IUCN (2012) criteria B2bi (continuing decline, observed,
inferred, or projected in extent of occurrence) and B2biii (continuing decline, observed, inferred, or projected, in
area, extent and/or quality of habitat; see Kusimi 2015). The discovery and description of Conraua sagyimase sp.
nov. underlines the importance of the Atewa Range Forest Reserve as a critical conservation area within the Upper
Guinea biodiversity hotspot (Myers et al. 2000, Mittermeier et al. 2004). However, the potential expansion of bauxite
mining concessions in this area is likely the greatest threat to the Atewa Range Forest Reserve (Lindsell et al. 2019).
The protection of this forest is fundamental for the survival of many endemic and threatened species such as various
arthropods (Naskrecki 2008a–c), various plants (Siaw & Dabo 2007, Hodgetts et al. 2016) and now also Conraua
sagyimase sp. nov. Future research and continuous monitoring are needed to uncover details of the life history,
population size and population demography of Conraua sagyimase sp. nov. Lastly, with the official description of
Conraua sagyimase sp. nov. as a species distinct from C. derooi, the range of the latter species—which is already
considered to be Critically Endangered—is now further restricted to the Togo-Volta Highlands of eastern Ghana and
western Togo (Hulselmans 1972). As remaining C. derooi populations are very small and patchily distributed, these
sites in the Togo-Volta Highlands also deserve urgent protection (Hillers et al. 2009, Segniagbeto et al. 2017).
Acknowledgments
The Ghana Forestry Commission and the Ghana Wildlife Division supported our work, provided permission to work
in the different forests in southern Ghana, and issued export permits. In Togo, the Ministère de l’Environnement et
des Ressources Forestières issued export permits. We are particularly indebted to all village chiefs in Ghana and Togo
who allowed us to work in their forests, and to all other villagers who helped us with their hospitality. Our assistants
and guides were of invaluable help at all investigated sites. Piotr Naskrecki and Alan Channing gave permission
to use their photos and Jeremy Lindsell provided the call recordings of the new species. We thank Peter S. Miller
(UWBM), Danny Meirte (MRAC), Wolfgang Böhme (ZFMK) and Annemarie Ohler (MNHN) for providing access
to voucher specimens under their care. Frank Tillack assisted with access to specimens and data at ZMB and prepared
the photos of the holotype of the new species. Amanda Quezada (MZUA) helped with photo editing. Mike Emmrich
introduced KNS into the analyses of call recordings, and Guillaume Demare (both ZMB) helped KNS with the
map. COB and NGK are indebted to Conservation International for organizing Rapid Assessment expeditions to the
Atewa range. COB additional work on the species has been supported by the ZSL EDGE of existence programme,
the Conservation Leadership Program (CLP) and the Rufford Small Grants Programme (RSG). Funding for the
molecular phylogenetic component of this work presented in Blackburn et al. (2020) came from the US National
Science Foundation to DCB (NSF DEB-1202609, 1560667 and 1556559).

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A NEW CONRAUA FROM GHANA Zootaxa 4995 (1) © 2021 Magnolia Press · 95
Appendix 1.
Additional specimens examined. MCZ= Museum of Comparative Zoology at Harvard University, Cambridge;
MNHN-RA= Muséum national d’Histoire naturelle, Paris; MRAC= Musée Royal de l’Afrique Centrale, Tervuren,
Belgium; UWBM= Burke Museum of Natural History and Culture, Seattle; ZFMK= Zoologisches Forschungsinstitut
und Museum Alexander Koenig, Bonn, ZMB= Museum für Naturkunde, Berlin.
Conraua alleni sensu lato. MCZ A-11991, semiadult, 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 Mountains 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 asl; ZMB 90304, adult male, Guinea, Fouta Djalon,
10°49’13.1”N, 12°11’30.7”W, 513 m asl; ZMB 90357, adult male and ZMB 90358, adult female, Ivory Coast,
Haute Dodo, 04°59’14’’N, 07°19’39’’W.
Conraua beccarii. ZFMK 15749–15750, Ethiopia, Illubator; MNHNP 1933. 21, adult male, holotype of Rana
Griaulei Angel, 1934, Ethiopia, Gondar, 2,200 m asl.
Conraua crassipes. ZMB 8360, holotype, “Abo”, north of Douala, Cameroon; ZFMK 73216, Gabon, Barrage
de Kinguélé, Tchimbélé; ZFMK 69351, 69353, 69354, Cameroon, Mt. Nlonako, Nguengue, Campsite; ZMB
90400–90403, Gabon, Moukalaba-Doudou National 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, Missahohe, 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-RA 1978.2027, 1978.2029, adult males and MNHN-RA 1978.2030, 1978.2031, adult
females and MNHN-RA 1978.2028, adult with unknown sex, Togo, Dangi Atiba; MNHN-RA 1993.2627, 1993.2629,
adult males and MNHN-RA 1993.2630, 1993.2631, adult females and MNHN-RA 1993.2628 adult with unknown
sex, Togo, Kloto; MNHN-RA 1993.4084–1993.4087, Togo, Missahohe; MNHN-RA 1995.5726, 1995.5727, Togo,
Kluto; MNHN-RA 1987.2026, Togo, Dangi Atigba.
Conraua goliath. ZFMK 77927, 77928, 77930, 77932, Cameroon, Mt. Nlonako, Ekomtolo, 500 m asl.
Conraua robusta. ZMB 20085, holotype, 1908, Cameroon; ZMB 78427, Cameroon; ZMB 90174, Cameroon,
Manengouba Village, Mt. Manengouba; ZFMK 67288, Cameroon, Bakossi Mts.: Kodmin.
Appendix 2.
Brief morphological description of Conraua derooi (for measures and comparisons with Conraua
sagyimase sp. nov. see Tables A1-A2).
Robust and rounded body shape; snout rounded in dorsal lateral view; dorsal skin smooth with small, rounded
scattered warts, skin on belly and throat smooth, tympanum hidden or indistinct, protruding eyes, positioned latero-
dorsally; vocal sacs absent; post occipital and suprascapular regions swollen in males; curved supratympanic fold,
extending from posterior edge of eye to shoulder; canthus rostralis distinct and rounded; loreal region concave;
legs with 15–21 rows of parallel longitudinal ridges; tarsal fold distinct; feet fully webbed; toes with well-defined,
big, rounded discs, dermal fringing on outer surfaces of toes I and V forming lateral skin folds; inner metatarsal
tubercle conspicuous; finger tips not expanded, rounded; lateral dermal fringing along edges of finger III; inner,
outer and middle palmar tubercles present; maxillary and premaxillary tooth present; three odontoid projections on
lower jaw, one at symphysis and one to each side on dentaries, vomerine teeth present; lateral line system comprises
infra-orbital line, supra-orbital line, upper lateral line, lower lateral line, median lateral line, jugular line and caudal
lateral line.