Citizen Science meets specimens in old formalin filled jars: a new species of Banded Rubber Frog, genus Phrynomantis (Anura, Phrynomeridae) from Angola

Abstract

Three species of Phrynomantis Peters, 1867, have been historically recorded for Angola: P. affinis, P. annectens and P. bifasciatus. As noted by all authors who have dealt with specimens of P. bifasciatus from the country, the Angolan population is characterized by an odd coloration pattern for the species, which led Boulenger to consider it a different variety. A revision of the extant specimens of Angolan Phrynomantis available in natural history collections, specimens collected in recent field surveys, as well as recent sightings and photographs allows the recognition of the Angolan population of P. cf. bifasciatus as a new species, endemic to the coastal lowlands of western Angola. The new taxon is described solely based on its coloration pattern and morphology, and it is separated from nominotypic P. bifasciatus by more than one thousand kilometers. The revision of these historical specimens also allowed us to confirm a second record of P. affinis in the country and to contribute to an overall better understanding of the distribution of the species of the genus on the continent.

Full text

Alytes, 2021, 38 (1–4): 18–48.
Citizen Science meets specimens
in old formalin filled jars: a new species of
Banded Rubber Frog, genus Phrynomantis
(Anura, Phrynomeridae) from Angola
Luis M. P. CERÍACO1,2, Bruna S. SANTOS3,2, Mariana P. MARQUES4,2,1,
Aaron M. BAUER5 & Arthur TIUTENKO6
1 Museu de História Natural e da Ciência da Universidade do Porto, Praça Gomes Teixeira 4099-002
Porto, Portugal
2 Departamento de Zoologia e Antropologia (Museu Bocage), Museu Nacional de História Natural e da
Ciência, Universidade de Lisboa, Rua da Escola Politécnica, 58, 1269-102 Lisboa, Portugal
3 Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, Rua do Campo Alegre,
4169-007 Porto, Portugal
4 Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO), InBIO,
Rua Padre Armando Quintas 7, Vairão, 4485-661 Porto, Portugal
5 Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University,
800 Lancaster Avenue, Villanova, Pennsylvania 19085-1699, USA
6 Friedrich-Alexander Universität, Schloßplatz 4, 91054, Erlangen, Germany
* Corresponding author <luisceriacogmail.com>.
Three species of Phrynomantis Peters, 1867, have been
historically recorded for Angola: P. affinis, P. annectens and
P. bifasciatus. As noted by all authors who have dealt with
specimens of P. bifasciatus from the country, the Angolan
population is characterized by an odd coloration pattern for the
species, which led Boulenger to consider it a different variety. A
revision of the extant specimens of Angolan Phrynomantis
available in natural history collections, specimens collected in
recent field surveys, as well as recent sightings and
photographs allows the recognition of the Angolan population
of P. cf. bifasciatus as a new species, endemic to the coastal
lowlands of western Angola. The new taxon is described solely
based on its coloration pattern and morphology, and it is
separated from nominotypic P. bifasciatus by more than one
thousand kilometers. The revision of these historical specimens
also allowed us to confirm a second record of P. affinis in the
country and to contribute to an overall better understanding of
the distribution of the species of the genus on the continent.

CERÍACO et al. 19
RESUMO
Três espécies de Phrynomantis Peters, 1867, têm sido
historicamente reportadas para Angola: P. affinis, P.
annectens e P. bifasciatus. Como referido por todos os
autores que têm lidado com espécimes de P. bifasciatus no
país, a população de Angola apresenta uma coloração atípica
para a espécie, o que levou mesmo Boulenger a considerá-la
uma variedade diferente. A revisão de todos os espécimes de
Phrynomantis de Angola existentes em coleções de história
natural, de espécimes recentemente colectados em novos
trabalhos de campo, bem como observação e fotografias
recentes, permitem o reconhecimento da população Angolana
de P. cf. bifasciatus como uma nova espécie, endémica das
zonas costeiras de baixa elevação do oeste de Angola. O novo
táxon é descrito apenas com base no seu padrão de coloração e
morfologia, e encontra-se separado da forma nominotípica de
P. bifasciatus por mais de mil quilómetros. A revisão destes
espécimes históricos permitiu também confirmar um segundo
registo de P. affinis para o país, contribuindo para uma melhor
compreensão da distribuição das espécies deste género no
continente.
urn:lsid:zoobank.org:pub:7890AFAD-EDEA-4195-BD15-72238789B721
INTRODUCTION
After being closed for research for approximately four decades due to violent
armed conflicts, Angola has in recent years experienced a renaissance of its biodiversity
studies (Marques et al. 2018; Huntley et al. 2019). This is especially true for
herpetology. In little more than a decade, dozens of papers have been published
regarding the country’s herpetofauna (see Marques et al. 2018 for a summary), with
several new species being described from several different families and genera of
amphibians and reptiles. These include the amphibian genera Hyperolius Rapp, 1842
(Conradie et al. 2012a, 2013) and Poyntonophrynus Frost et al., 2006 (Ceríaco et al.
2018), and the reptile genera Boaedon Duméril, Bibron & Duméril, 1854 (Hallermann
et al. 2020), Cordylus Laurenti, 1768 (Stanley et al. 2016; Marques et al. 2019a),
Hemidactylus Goldfuss, 1820 (Ceríaco et al. 2020a–b), Limnophis Günther, 1865
(Conradie et al. 2020), Lygodactylus Gray, 1864 (Marques et al. 2020), Nucras Gray,
1838 (Branch et al. 2019), Panaspis Cope, 1868 (Ceríaco et al. 2020c), Pedioplanis
Fitzinger, 1843 (Conradie et al. 2012b) and Trachylepis Fitzinger, 1843 (Marques et al.
2019b). Many other species from these and other genera are currently in the process of
being described, and certainly many more await detailed study.
The curious and visually striking phrynomerid frogs of the genus Phrynomantis
Peters, 1867, commonly known as Rubber Frogs due to their smooth, rubber-like dorsal
skin, are one such group. Phrynomantis is a relatively small genus comprising five
currently recognized species, all of them endemic to sub-Saharan Africa: Phrynomantis
affinis Boulenger, 1901, Phrynomantis annectens Werner, 1910, Phrynomantis
bifasciatus (Smith, 1847), Phrynomantis microps Peters, 1875 and Phrynomantis

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20
somalicus (Scortecci, 1941). Three of these have been recorded for Angola, namely P.
affinis, P. annectens and P. bifasciatus (Marques et al. 2018).
Phrynomantis affinis was originally described from “Pweto, Lake Mweru” in the
former Katanga region of the Democratic Republic of the Congo, and its known
distribution ranges from southern Democratic Republic of the Congo, southern
Tanzania, western Zambia, to northeastern Namibia and eastern Angola (Channing
2001; Channing & Rödel 2019). Parker (1940) described Phrynomantis hoeschi from
northern Namibia (type locality: “Ombujomatemba [1450 m], near the Waterberg”
[Parker 1940]), which was later considered as a synonym of P. affinis by Laurent
(1964), a decision followed by all subsequent authors (e.g. Mertens 1971; Dubois 1988;
Channing 2001; Channing & Rödel 2019; Frost 2020). In Angola P. affinis is
historically known only from a single male specimen from the Lake Calundo area
([latitude] ‒11.80º, [longitude] 20.86º; 1119 m [above sea level]) in Moxico Province,
eastern Angola (Laurent 1964). Recent reexamination of the specimens reported by
Monard (1938) as P. bifasciatus reveals that the author misidentified the species, and
these records actually refer to P. affinis. Monard’s (1938) report was based on seven
juvenile specimens collected in June 1933 in Mulondo (‒15.6479158º, 15.1955508º;
1158 m), on the high plateau of Huíla Province. Monard noted that all of the specimens
at his disposal were “dark, leaden black, decorated with very small white spots, more or
less arranged in longitudinal lines” [“sombre, d’un noir plombé, ornée de très petites
taches blanches, plus ou moins disposées en lignes longitudinales”], in which “the spots
behind the eye are the most linearly arranged” [“Les taches qui existent en arrière de
l’œil sont les mieux disposées en ligne”], and all the specimens had a “whitish
underside” [le dessous est blanchâtre”] [translation by the authors from the original in
French]. Monard stated that this coloration was “abnormal” for P. bifasciatus, which is
“normally ornamented by yellow or red lines over a brown or black background”
[“ordinairement ornée de lignes jaunes ou rouges sur fond brun ou noir”] [translation by
the authors from the original in French] (Monard 1938). Two of the seven specimens
are still present in the collections of the Musée d’Histoire Naturelle de La Chaux-de-
Fonds (MHNC 90.001–002; Fig. 1a) and are unambiguously assigned to P. affinis,
extending the known distribution of the species approximately 750 km southwest in
Angola.
Figure 1. Dorsal (a) and ventral view of the left hand (b) of Phrynomantis affinis (MHNC 90.001 [or 2])
from Mulondo, Huila Province, Angola. Photo by Nicolas Marcgraf.

CERÍACO et al. 21
Contrasting with the relatively stable taxonomic and nomenclatural history of P.
affinis, the Namib specialist P. annectens has a more complex history. The species was
originally described by Werner (1910) from the “Aar-Rivier” in southwestern Namibia.
Several subsequent authors described three other taxa, all currently considered as junior
synonyms of P. annectens. Methuen & Hewitt (1913) described Phrynomantis nasuta
from the Great Karas mountains region in southern Namibia; Ahl (1934) described
Hoplophryne marmorata from Okahandja, central Namibia; and Ahl (1935) described
Ctenophryne marmorata (thus creating a junior secondary homonym of the latter) based
on two specimens without locality data (for a discussion on the identity of this taxon see
Zweifel 1989). This synonymy is currently accepted by all authors who have dealt with
the species (e.g. Loveridge 1936; Parker 1936; Mertens 1971; Dubois 1988; Zweifel
1989; Bauer et al. 1996; Channing 2001; Channing & Rödel 2019; Frost 2020). P.
annectens is an arid region specialist, and occurs from the xeric areas of southwestern
Angola, through western and central Namibia to the Augrabies area of the western
Northern Cape Province in South Africa (Channing 2001; Marques et al. 2018;
Channing & Rödel 2019). In Angola, P. annectens is known from a handful of records
from the coastal lowlands of western Angola (Parker 1936; Poynton & Haacke 1993;
Ceríaco et al. 2016; see Marques et al. [2018] for details on the available published data
on the distribution of the species in the country), and has recently been extensively
collected in several localities in Benguela and Namibe provinces (this paper, Fig. 2, see
Table 1). Besides the published historical records, there is an unpublished specimen
from “Benguela” (-12.5833º, 13.41667º, 15 m above sea level), Benguela Province,
collected by William John Ansorge in his 1903–1909 exploration of Angola, which is
deposited in the collections of the Royal Museum for Central Africa (RMCA) in
Tervuren (accession number RMCA B.469; Fig. 3).
Figure 2. Life photo (a) and ventral view of the left hand (b) of Phrynomantis annectens (LMPC 1149)
from Malowe, base of Serra da Neve, Namibe Province, Angola. Photos by Luis M. P. Ceríaco.

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Table 1. Specimens used for morphological and coloration comparisons.
Taxon Accession number Locality
Phrynomantis affinis MD 5726 ANGOLA: Lake Calundo region (‒11.80º,
20.86º; 1119 m), Moxico Province
Phrynomantis affinis MHNC 90.001–90.002 ANGOLA: Mulondo (‒15.6479158º,
15.1955508º; 1158 m), Huíla Province
Phrynomantis affinis NMNW 10854–10855 NAMIBIA: Farme Marne, Gobabis (‒22.41811º,
18.86524º; 1476 m)
Phrynomantis affinis NMNW NAMIBIA: Farme Marne, Gobabis (‒22.41874º,
18.854985º; 1474 m)
Phrynomantis annectens RMCA B.469 ANGOLA: Benguela (‒12.5833º, 13.41667º;
15 m), Benguela Province
Phrynomantis annectens
CAS 263085–263088;
UF 187252–187279;
INBAC/AMB 10058,
10060–10064, 10078
ANGOLA: Chimalavera Nature Reserve,
Waterhole (‒12.79168º, 13.12738º; 225 m),
Benguela Province
Phrynomantis annectens CAS 255056 ANGOLA: Omauha Lodge (‒16.19872º,
12.40008º; 338 m), Namibe Province
Phrynomantis annectens AMB 11353, 11354;
INBAC/AMB 11355
ANGOLA: Maungo Dam (‒14.57922º,
12.68057º; 549 m), Namibe Province
Phrynomantis annectens UF 187250–187251;
INBAC/AMB 10344
ANGOLA: Dolondolo (‒13.81328º, 13.13618º;
681 m) Namibe Province
Phrynomantis annectens LMPC 1149 ANGOLA: Malowe (‒13.83424º, 13.27669º;
803 m), Namibe Province
Phrynomantis annectens
BMNH 1845.5.11.131;
1858.4.25.171–172
(syntypes)
SOUTH AFRICA: “South Africa” [without
specific locality]
Phrynomantis bifasciatus
FMNH 80627–80629,
FMNH 80631–80638,
FMNH 190740,
FMNH 187346
DEMOCRATIC REPUBLIC OF THE CONGO:
Upemba National Park, Mabwe (‒8.65º,
26.516667º; 581 m)
Phrynomantis bifasciatus
FMNH 205396,
FMNH 205404,
FMNH 206462,
ESWATINI: Tshaneni (‒26.001284º,
31.759438º; 300 m)

CERÍACO et al. 23
Taxon Accession number Locality
Phrynomantis bifasciatus FMNH 2410 KENYA: Lukenya (‒1.483313º, 37.068721º;
1839 m)
Phrynomantis bifasciatus FMNH 129557 TANZANIA: Nachingwea district (‒10.5º,
38.3333º; 420 m)
Phrynomantis bifasciatus FMNH 18002–18006 BOTSWANA: Metsimaklaba road
Phrynomantis bifasciatus FMNH 18001 BOTSWANA: Molepolole (‒24.406587,
25.495079; 1138 m)
Phrynomantis bifasciatus
FMNH 205394–
205395, FMNH
205398– 205399,
FMNH 205401–
205403, FMNH
205405, FMNH
206463–206464
ESWATINI: Tshaneni (‒26.001284º,
31.759438º; 300 m)
Phrynomantis bifasciatus NMNW 25218 NAMIBIA: Tsumkwe, Otjozondjupa Region
(‒19.5833º, 20.4833º; 1115 m)
Phrynomantis bifasciatus NMNW 25624 NAMIBIA: Popa Falls (‒18.1º, 21.56667º;
1025 m), Kavango Region
Phrynomantis bifasciatus NMNW 25683 NAMIBIA: Otjikaro (‒20.61805º, 17.191074º;
1432 m), Otjozondjupa Region
Phrynomantis bifasciatus NMNW 25779 NAMIBIA: Okosomingo (‒20.623536º,
17.065656º; 1494 m), Otjozondjupa Region
Phrynomantis bifasciatus NMNW 25679 NAMIBIA: Ombujoma-temba (‒20.637249º,
17.236565º; 1420 m), Otjozondjupa Region
Phrynomantis bifasciatus NMNW 25761 NAMIBIA: Erindi, Osombaka (‒21.28333º,
17.38333º; 1410 m), Otjozondjupa Region
Phrynomantis newtoni
sp. nov.
MHNCUP/ANF-194
(holotype)
ANGOLA: Chingo (‒11.2º, 13.85º; 11 m),
Kwanza Sul Province
Phrynomantis newtoni
sp. nov. BMNH 1873.7.28.14 ANGOLA: “Angola” [without specific locality]

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Figure 3. Dorsal view of Phrynomantis annectens (RMCA A.469) from Benguela, Benguela Province,
Angola. Photo by Garin Cael.
Figure 4. Dorsal (a) and ventral view of the right hand (b) of Phrynomantis bifasciatus (FMNH 80634)
from Mabwe, Upemba National Park, Democratic Republic of the Congo. Photo by Adam Ferguson.

CERÍACO et al. 25
Regarding P. bifasciatus, the most widely distributed member of the genus in
southern Africa, its taxonomic and nomenclatural history is also complex, with the
description of two taxa that would later be sunk into synonymy, and its allocation at
different times to several genera: Brachymerus Smith, 1847, Dendrobates Wagler,
1830, Bombinator Merrem, 1820, Phrynomantis Peters, 1867 and Phrynomerus Noble,
1926 (see Dubois [1988] and Frost [2020]). This visually striking species, easily
identifiable by the two, usually continuous broad orange to red dorsolateral bands (Fig.
4‒5), was originally described by Smith (1847) based on specimens from the “country
to the east and north-east of the Cape Colony”. Poynton & Broadley (1985) noted that
although Smith did not provide many details regarding the type series and that the
author’s material in the BMNH do not correspond exactly to the original description,
the specimens BMNH 1845.5.11.131 and 1858.4.25.171–172 likely correspond to the
type material of the species. The two other nominal taxa currently in synonymy are
Dendrobates inhambanensis described by Bianconi (1849) from the Inhambane region
in Mozambique, and Phrynomerus bifasciatus nyasalandensis, described by Hoffman
(1944) from “Chitiala, Nyasaland” [Chitala River] in Malawi. While it is clear that
Bianconi did not have access to Smith’s publication at the time he wrote the description
of Dendrobates inhambanensis (the publications are separated by only a year: 1847 and
1848), the subspecies nyasalandensis was diagnosed against the nominotypical taxon on
the basis of the different shape and sizes of the snout, second finger, third toe and hind
limb (Hoffman 1944; Loveridge 1953). Both these taxa are unambiguously conspecific
with the nominotypical form and have been considered junior synonyms by all authors
that have subsequently reviewed the group (e.g. Peters 1882; Loveridge 1953; Poynton
& Broadley 1985; Channing 2001; Channing & Rödel 2019; Frost 2020). The species is
widely distributed in sub-Saharan Africa, ranging from Somalia through the Democratic
Republic of the Congo, western Angola, central and eastern Namibia, to northeastern
South Africa (Channing 2001; Channing & Rödel 2019).
In Angola, P. bifasciatus has been historically recorded from the western regions
of the country by Boulenger (1882), Bocage (1895), Ferreira (1904) and Monard
(1938). However, as noted above, Monard’s (1938) specimens represent a
misidentification of P. affinis. All historical records of P. bifasciatus in Angola present
a common characteristic that differentiates them from the majority of the specimens
from the rest of the species distribution. Boulenger (1882) notes that the single
specimen (BMNH 1873.7.28.14) from “Angola” (no specific locality provided, but
surely from Luanda region, as it was the area where the collector was based at the time;
LMPC pers. obs.) at his disposal, collected by Joachim John Monteiro, has its “upper
surfaces with large round light spots” and “no light bands”. This coloration pattern led
Boulenger (1882) to designate the Angolan specimen as “Var. C.”, contrasting with the
nominotypical “Var. A” (for which Boulenger consulted Smith’s type material), which
has “white (or red), sometimes interrupted streak on each side of the body, beginning on
the upper eyelid”, “a white subtriangular or heart-shaped marking on coccygeal region”,
and “limbs spotted with white”; and “Var. B” (Shiré Valley and Zambezi region
material), which is “similar to the preceding [Var. A], but the space between the white
bands with dark and light longitudinal lines”, and “markings on the legs arranged in
cross bars.”
This coloration difference was also noted by Bocage (1895) on the two
specimens available at the time in the Lisbon Museum collections, one specimen from

ALYTES 38 (1–4)
26
“Quissange” (‒12.43333º, 14.0500º; 870 m) and another from “Benguela”, both in
Benguela Province. According to the author the specimen from Quissange “belongs” to
Boulenger’s (1882) “var. C”, and presented a “dark dorsal coloration, with large white
or yellow spots in the dorsum and the limbs but lacking the longitudinal lines of the
typical form, the throat, the chest as well as the posterior part of the thighs and the
underside of the legs brown, speckled with whitish, the abdomen greyish white with
brown markings, and the upper side of the limbs the same color as the dorsal
coloration”. [Translation by the authors from the original in French: “Il est noir en
dessus, varié sur le dos et les membres de grandes taches blanches (jaunes ?), mais sans
les deux raies longitudinales de la forme typique; la gorge et la poitrine, ainsi que la
face postérieure des cuisses et la face inférieure des jambes, brunes tachetées de
blanchâtre; l’abdomen d’un blanc grisâtre avec des marbrures brunes; la face supérieure
des membres de la couleur du dos”]. Regarding the specimen from Benguela, Bocage
(1895) noted that the “upper parts lacking lines or spots, had a reddish-brown
background, brighter and finely dotted with brown in the head region and dorsum, paler
and uniform on the sides, with small brown spots on the upper sides of the limbs; the
undersides of grey with a fawn wash, with neither spots nor punctuations” [Translation
by the authors from the original in French: “le dos est dépourvu de raies et de taches;
sur les parties supérieures règne une teinte roux-fauve, plus vive et finement pointillée
de marron sur la tête et le dos, plus pâle et uniforme sur les flancs, variée de petites
taches marron sur la face supérieure des membres. En dessous d’un gris lavé de fauve,
sans taches ni ponctuations.”]. This latter specimen was illustrated in a plate (Bocage
1895, Fig. 6). These two specimens were lost in the fire that destroyed the Lisbon
museum in 1978.
Later, Ferreira (1904) reported on an adult specimen “of Boulenger’s Var. C”
from “Chingo (Novo Redondo)” [Chingo, in Sumbe district, formerly Novo Redondo
district], Kwanza Sul Province, Angola collected by Francisco Newton in 1903.
According to the author, the specimen had the gular region “strongly” darkly
pigmented, and all the other characters were in accordance with Boulenger’s
description. This specimen is still extant in the collections of the Museu de História
Natural e da Ciência da Universidade do Porto (MHNCUP), Porto (Portugal), under the
accession number MHNCUP/ANF-194 (Fig. 7).
More recently some photographs, putatively attributed to this species, have been
published on social media networks, namely on Facebook pages dedicated to the
dissemination of the biodiversity of Angola (“Biodiversidade de Angola” and “Angola
Selvagem” Facebook groups). One specimen was found in the Calumbo area
(‒9.1499994º, 13.3991574º; 3 m), Luanda Province, in June 2019. The specimen was
not collected but was photographed alive (Fig. 8), and it shows the same type of
coloration pattern as those described by previous authors for the Angolan population of
P. bifasciatus. On 24 April 2020, another specimen was also found in the Luanda
neighborhood of Benfica (‒8.972175º, 13.142503º; 18 m), Luanda Province, in 24 April
2020. This specimen was not collected but it was photographed (Fig. 9) and it also
presents the same coloration pattern.

CERÍACO et al. 27
Figure 5. Life photos of Phrynomantis bifasciatus from several localities across southern Africa: (a)
Mozambique; (b) Harare, Zimbabwe; (c) Muunze Forest Lodge, Zimbabwe; (d) Muunze Forest Lodge,
Zimbabwe; (e) Okonjima, Namibia; (f) Olievenhoutsrus, South Africa. Photos by Harith Farooq (a), and
François Becker (b-f).
Figure 6. Illustration of Angolan Phrynomantis bifasciatus from Bocage᾿s Herpétologie d’Angola et du
Congo (1895).

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Figure 7. Holotype of Phrynomantis newtoni sp. nov. from “Chingo (Novo Redondo)” collected by
Francisco Newton in 1903 (MHNCUP/ANF-194). Photo by Bruna Santos.
Figure 8. Live photo of Phrynomantis newtoni sp. nov. from Calumbo region, near Luanda. Specimen
photographed in June 2019, but not collected. Photo by Manuel Castanho.

CERÍACO et al. 29
Currently, there is no available phylogeny including all the members of the
genus, and a full and detailed taxonomic treatment of the genus is also lacking. The
most recent summary on the morphology and diagnostic characters of species of the
genus was provided in the field guide on African amphibians by Channing & Rödel
(2019). As noted by these authors, the main way to differentiate Phrynomantis species,
besides molecular data and probably vocalizations, is through their color patterns and
distribution ranges. Other characters, such as the shape and proportions of the body and
the presence or absence of enlarged disks on the terminal part of the finger- and toe-tips,
are also important diagnostic characters. In the light of the distinctive coloration pattern
and considerable geographic distance of the Angolan population of P. bifasciatus from
the closest records of the species in Zambia and northern Namibia, we consider the
Angolan population as a distinct taxon that we describe herein.
MATERIALS AND METHODS
For comparisons we examined 88 specimens (see Table 1) of Phrynomantis sp.
deposited in the folowing institutions: California Academy of Sciences (CAS), San
Francisco, USA; American Museum of Natural History (AMNH), New York, USA;
Florida Museum of Natural History (UF), Gainesville, USA; Field Museum of Natural
History (FMNH), Chicago, USA; Natural History Museum (BMNH), London, UK;
Royal Museum for Central Africa (RMCA), Tervuren, Belgium; Musée d’Histoire
Naturelle de La Chaux-de-Fonds (MHNC), La Chaux-de-Fonds, Switzerland; Instituto
Nacional da Biodiversidade e Áreas de Conservação (INBAC/AMB), Kilamba-Kiaxi,
Angola; Museu Regional do Dundo (MD), Dundo, Angola; National Museum of
Namibia (NMNW), Windhoek, Namibia; and Museu de História Natural e da Ciência
da Universidade do Porto (MHNCUP), Porto, Portugal. Information on morphological
characters of species and/or type material that could not be examined, as well as
supplementary data for the species of the genus was obtained from the relevant
literature (Smith 1847; Bianconi 1849; Peters 1875; Boulenger 1882, 1901; Bocage
1895; Werner 1910; Monard 1938; Scortecci 1941; Hoffman 1944; Passmore &
Carruthers 1979; Poynton & Broadley 1985; Auerbach 1987; Lambiris 1989; Channing
2001; Pickersgill 2007; Channing & Howell 2006; Spawls et al. 2006; Farook et al.
2014; Channing & Rödel 2019). In order to check the coloration patterns of the different
species across their known distributions, we also checked the research grade records
[records confirmed by a consensus of experts] of the genus Phrynomantis published on
the online citizen science platform iNaturalist (www.inaturalist.org; Anonymous 2020)
as well in relevant literature for which locality data could be retrieved (Passmore &
Carruthers 1979; Auerbach 1987; Pickersgill 2007; Du Preez & Carruthers 2009, 2017;
Farook et al. 2014). Each specimen was classified according to its coloration pattern
(Table 2).

ALYTES 38 (1–4)
30
Table 2. Coloration patterns of P. bifasciatus and P. newtoni sp. nov. Color pattern description: A,
Uninterrupted orange dorsolateral bands on a homogeneous dark background; B, Uninterrupted orange
dorsolateral bands, with faint vestiges of greyish dorsal bands; C, Briefly interrupted orange dorsolateral
bands on a homogeneous dark background; D, large irregular yellowish, red or silver blotches. Catalog
and accession number abbreviations: IN, iNaturalist Record; MHNCUP, Museu de História Natural e da
Ciência da Universidade do Porto, Porto, Portugal; BMNH, Natural History Museum, London, United
Kingdom; FMNH, Field Museum of Natural History, Chicago, USA; NMNW, National Museum of
Namibia.
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
IN 66735019 P. bifasciatus BOTSWANA: Palapye ‒22.560776 27.104304 A https://www.inaturalist.org/
observations/66735019
IN 66996545 P. bifasciatus MOZAMBIQUE: Mtindiri ‒18.982363 34.353337 B https://www.inaturalist.org/
observations/66996545
IN 66884527 P. bifasciatus SOUTH AFRICA:
Mookgophong/Modimolle ‒24.319341 28.013233 A https://www.inaturalist.org/
observations/66884527
IN 66788362 P. bifasciatus SOUTH AFRICA:
Kruger National Park ‒24.395533 31.77947 C https://www.inaturalist.org/
observations/66788362
IN 66342008 P. bifasciatus ZIMBABWE: Fortunes Gate,
Bulawayo ‒20.196273 28.618005 B https://www.inaturalist.org/
observations/66342008
IN 66368557 P. bifasciatus ZIMBABWE:
Unspecificd Locality ‒20.639572 32.023923 C https://www.inaturalist.org/
observations/66368557
IN 65927954 P. bifasciatus SOUTH AFRICA: Madibeng ‒25.516886 27.801664 A https://www.inaturalist.org/
observations/65927954
IN 65201623 P. bifasciatus SOUTH AFRICa: Skukuza ‒24.993472 31.593388 A https://www.inaturalist.org/
observations/65201623
IN 65461281 P. bifasciatus SOUTH AFRICA: Waterberg ‒24.743777 28.221943 C https://www.inaturalist.org/
observations/65461281
IN 65591352 P. bifasciatus ZIMBABWE:
Granite Park, Bulawayo ‒20.209947 28.590184 C https://www.inaturalist.org/
observations/65591352
IN 65516610 P. bifasciatus SOUTH AFRICA: Tshwane
Metropolitan Municipality ‒25.706558 28.512823 A https://www.inaturalist.org/
observations/65516610
IN 64439126 P. bifasciatus SOUTH AFRICA:
Unspecificd Locality ‒25.36129 28.564498 A https://www.inaturalist.org/
observations/64439126
IN 64282565 P. bifasciatus SOUTH AFRICA:
Unspecificd Locality ‒25.292854 28.444443 A https://www.inaturalist.org/
observations/64282565
IN 62135809 P. bifasciatus SOUTH AFRICA:
Jdm Keet Plantation ‒23.781484 30.119391 A https://www.inaturalist.org/
observations/62135809
IN 60979478 P. bifasciatus SOUTH AFRICA:
Kruger National Park, Skukuza ‒24.99223 31.59261 A https://www.inaturalist.org/
observations/60979478
IN 58593708 P. bifasciatus TANZANIA: Utende ‒7.973196 39.748081 B https://www.inaturalist.org/
observations/58593708
IN 56516404 P. bifasciatus SOUTH AFRICA: Sandriv ‒22.98161 29.617859 A https://www.inaturalist.org/obs
ervations/56516404
IN 56516352 P. bifasciatus SOUTH AFRICA: Mapungubwe ‒22.184585 29.202902 C https://www.inaturalist.org/obs
ervations/56516352
IN 56594458 P. bifasciatus TANZANIA: Mafia ‒7.973248 39.7495 C https://www.inaturalist.org/
observations/56594458
IN 56516348 P. bifasciatus SOUTH AFRICA: Mapungubwe ‒22.184585 29.202902 A https://www.inaturalist.org/
observations/56516348
IN 56516392 P. bifasciatus SOUTH AFRICA: Sandriv ‒22.98161 29.617859 A https://www.inaturalist.org/
observations/56516392

CERÍACO et al. 31
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
IN 56516383 P. bifasciatus SOUTH AFRICA: Sandriv ‒22.98161 29.617859 A https://www.inaturalist.org/
observations/56516383
IN 50864304 P. bifasciatus MOZAMBIQUE:
Khodzue Gorge, Cheringoma ‒18.56025 34.87319 B https://www.inaturalist.org/
observations/50864304
IN 49221966 P. bifasciatus MOZAMBIQUE:
Unspecified Locality ‒24.101316 35.498109 B https://www.inaturalist.org/
observations/49221966
IN 44621203 P. bifasciatus SOUTH AFRICA:
North Uthungulu ‒27.580165 32.068073 A https://www.inaturalist.org/
observations/44621203
IN 42610270 P. bifasciatus SOUTH AFRICA:
Donkerhoek, South Africa ‒25.809526 28.467207 B https://www.inaturalist.org/
observations/42610270
IN 42399504 P. bifasciatus TANZANIA: Unspecified Locality ‒4.166667 38.166667 B https://www.inaturalist.org/
observations/42399504
IN 42657487 P. bifasciatus MOZAMBIQUE: Chitengo Camp,
Gorongosa National Park ‒18.98015 34.35201 A https://www.inaturalist.org/
observations/42657487
IN 41883270 P. bifasciatus TANZANIA: Ulanga ‒8.515021 36.45418 B https://www.inaturalist.org/
observations/41883270
IN 41294335 P. bifasciatus TANZANIA:
Mkomazi National Park ‒4.165637 38.166504 A https://www.inaturalist.org/
observations/41294335
IN 41589877 P. bifasciatus ZIMBABWE:
Camp Amalinda, Matobo ‒20.48142 28.43761 B https://www.inaturalist.org/
observations/41589877
IN 40180972 P. bifasciatus BOTSWANA: North-West District ‒18.440065 23.711235 C https://www.inaturalist.org/
observations/40180972
IN 39931672 P. bifasciatus SOUTH AFRICA: Thabazimbi ‒24.582633 27.402774 A https://www.inaturalist.org/
observations/39931672
IN 38323415 P. bifasciatus SOUTH AFRICA: Kruger Park ‒24.393097 31.776115 C https://www.inaturalist.org/
observations/38323415
IN 39089327 P. bifasciatus SOUTH AFRICA:
Unspecified Locality ‒25.362608 27.722178 A https://www.inaturalist.org/
observations/39089327
IN 37746911 P. bifasciatus BOTSWANA: Tutume ‒20.830012 27.075294 B https://www.inaturalist.org/
observations/37746911
IN 37503465 P. bifasciatus SOUTH AFRICA:
Kruger National Park ‒24.421383 31.785453 A https://www.inaturalist.org/
observations/37503465
IN 37482681 P. bifasciatus SOUTH AFRICA:
Unspecified Locality ‒24.150364 30.59408 A https://www.inaturalist.org/
observations/37482681
IN 36877649 P. bifasciatus SOUTH AFRICA:
Waterberg, Limpopo ‒25.30653 28.444632 C https://www.inaturalist.org/
observations/36877649
IN 36877650 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.305842 28.445615 A https://www.inaturalist.org/
observations/36877650
IN 36877648 P. bifasciatus SOUTH AFRICA:
Waterberg, Limpopo ‒25.223067 28.579498 C https://www.inaturalist.org/
observations/36877648
IN 36607649 P. bifasciatus SOUTH AFRICA: Far North ‒22.597419 29.926524 A https://www.inaturalist.org/
observations/36607649
IN 36704991 P. bifasciatus SOUTH AFRICA: Bojanala ‒25.378464 26.752327 C https://www.inaturalist.org/
observations/36704991
IN 36439932 P. bifasciatus BOTSWANA: Gaborone ‒24.673638 25.874554 A https://www.inaturalist.org/
observations/36439932
IN 36405385 P. bifasciatus KENYA: Lamu ‒2.147905 40.895092 A https://www.inaturalist.org/
observations/36405385
IN 36557268 P. bifasciatus SOUTH AFRICA:
Kruger National Park, Skukuza ‒24.992302 31.592463 A https://www.inaturalist.org/
observations/36557268
IN 36053034 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.304086 28.461982 B https://www.inaturalist.org/
observations/36053034

ALYTES 38 (1–4)
32
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
IN 36088381 P. bifasciatus SOUTH AFRICA:
Ngaka Modiri Molema ‒26.763176 25.722552 A https://www.inaturalist.org/
observations/36088381
IN 35942350 P. bifasciatus SOUTH AFRICA:
Waterberg, Limpopo ‒25.260875 28.439055 A https://www.inaturalist.org/
observations/35942350
IN 36285967 P. bifasciatus MOZAMBIQUE: Mecula ‒12.177257 38.084187 C https://www.inaturalist.org/
observations/36285967
IN 35779955 P. bifasciatus SOUTH AFRICA:
North Uthungulu ‒27.973542 32.329152 B https://www.inaturalist.org/
observations/35779955
IN 9809840 P. bifasciatus ZIMBABWE:
Dambari Field Station ‒20.267638 28.815321 B https://www.inaturalist.org/
observations/9809840
IN 30533908 P. bifasciatus SOUTH AFRICA:
Mjejaqne Gr, Ehlanzeni ‒25.407292 31.679869 A https://www.inaturalist.org/
observations/30533908
IN 26246732 P. bifasciatus
TANZANIA:
University of Dar es Salaam
Mwalimu Nyerere Main
Campus, Dar es Salaam
‒6.782012 39.20498 C https://www.inaturalist.org/
observations/26246732
IN 25205915 P. bifasciatus KENYA: Kilifi North ‒3.340291 39.947621 A https://www.inaturalist.org/
observations/25205915
IN 22292373 P. bifasciatus TANZANIA: Arumeru ‒3.335476 36.88807 C https://www.inaturalist.org/
observations/22292373
IN 20975060 P. bifasciatus MOZAMBIQUE: Mecula ‒12.176764 38.091802 B https://www.inaturalist.org/
observations/20975060
IN 19672290 P. bifasciatus MOZAMBIQUE: Mecula ‒12.168727 38.091888 B https://www.inaturalist.org/
observations/19672290
IN 19371383 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.261945 28.442643 C https://www.inaturalist.org/
observations/19371383
IN 19312853 P. bifasciatus SOUTH AFRICA:
Waterberg, Limpopo ‒24.69725 28.599787 A https://www.inaturalist.org/
observations/19312853
IN 19311550 P. bifasciatus NAMIBIA: Otjozondjupa Region ‒20.573325 16.891549 B https://www.inaturalist.org/
observations/19311550
IN 19312850 P. bifasciatus SOUTH AFRICA:
Waterberg, Limpopo ‒24.715207 28.482347 A https://www.inaturalist.org/
observations/19312850
IN 19311553 P. bifasciatus NAMIBIA: Otjozondjupa Region ‒20.573255 16.891581 A https://www.inaturalist.org/
observations/19311553
IN 19102682 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.223307 28.579777 A https://www.inaturalist.org/
observations/19102682
IN 18918404 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.223267 28.579603 C https://www.inaturalist.org/
observations/18918404
IN 18918405 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.223267 28.579612 A https://www.inaturalist.org/
observations/18918405
IN 18918403 P. bifasciatus SOUTH AFRICA:
Waterberg, Limpopo ‒25.223255 28.579918 A https://www.inaturalist.org/
observations/18918403
IN 18875696 P. bifasciatus SOUTH AFRICA:
Ukuwela Nature Reserve ‒27.902686 32.309613 B https://www.inaturalist.org/
observations/18875696
IN 14934278 P. bifasciatus ZAMBIA: Lusaka ‒15.359975 28.445497 B https://www.inaturalist.org/
observations/14934278
IN 640808 P. bifasciatus MOZAMBIQUE: Moatize ‒16.026366 33.701748 C https://www.inaturalist.org/
observations/640808
IN 13766881 P. bifasciatus TANZANIA: Unnamed Road ‒4.536827 39.05308 A https://www.inaturalist.org/
observations/13766881
IN 13206401 P. bifasciatus TANZANIA:
Kinondoni, Dar es Salaam ‒6.782103 39.03737 A https://www.inaturalist.org/
observations/13206401

CERÍACO et al. 33
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
IN 13206403 P. bifasciatus TANZANIA:
Kinondoni, Dar es Salaam ‒6.781938 39.037228 B https://www.inaturalist.org/
observations/13206403
IN 13138048 P. bifasciatus TANZANIA: Kilosa, Morogoro ‒7.396207 37.00168 A https://www.inaturalist.org/
observations/13138048
IN 11228065 P. bifasciatus SOUTH AFRICA:
Itibane Game Lodge ‒27.854429 32.080342 B https://www.inaturalist.org/
observations/11228065
IN 13006271 P. bifasciatus TANZANIA: Bagamoyo ‒6.433013 38.904557 B https://www.inaturalist.org/
observations/13006271
IN 11225670 P. bifasciatus SOUTH AFRICA:
Dinokeng Central ‒ Pan ‒25.347036 28.373247 A https://www.inaturalist.org/
observations/11225670
IN 11216311 P. bifasciatus SOUTH AFRICA:
Stockpoort Lodge ‒23.411281 27.354544 A https://www.inaturalist.org/
observations/11216311
IN 11014425 P. bifasciatus SOUTH AFRICA: Satara ‒24.391589 31.779882 A https://www.inaturalist.org/
observations/11014425
IN 10960486 P. bifasciatus SOUTH AFRICA: Koedoeskop ‒25.37165 29.54609 A https://www.inaturalist.org/
observations/10960486
IN 10916294 P. bifasciatus SOUTH AFRICA:
Masebe Nature Reserve ‒23.6409 28.5565 A https://www.inaturalist.org/
observations/10916294
IN 10707046 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.261415 28.45675 B https://www.inaturalist.org/
observations/10707046
IN 9654333 P. bifasciatus SOUTH AFRICA: Bosveld ‒25.275208 28.47221 A https://www.inaturalist.org/
observations/9654333
IN 9087919 P. bifasciatus SOUTH AFRICA: Waterberg ‒24.529772 27.502291 B https://www.inaturalist.org/
observations/9087919
IN 8634331 P. bifasciatus TANZANIA:
Kinondoni, Dar es Salaam ‒6.781945 39.036755 A https://www.inaturalist.org/
observations/8634331
IN 8455099 P. bifasciatus ZIMBABWE: Unnamed Road ‒20.347688 32.027631 C https://www.inaturalist.org/
observations/8455099
IN 8455577 P. bifasciatus TANZANIA:
Kibamba, Dar es Salaam ‒6.782042 39.037298 B https://www.inaturalist.org/
observations/8455577
IN 8412237 P. bifasciatus ESWATINI: Lubombo Region ‒26.134463 32.00433 A https://www.inaturalist.org/
observations/8412237
IN 7107167 P. bifasciatus TANZANIA:
Kinondoni, Dar es Salaam ‒6.781823 39.036932 B https://www.inaturalist.org/
observations/7107167
IN 7107175 P. bifasciatus TANZANIA:
Kibamba, Dar es Salaam ‒6.781645 39.036743 B https://www.inaturalist.org/
observations/7107175
IN 7107050 P. bifasciatus TANZANIA:
Kinondoni, Dar es Salaam ‒6.781645 39.036743 B https://www.inaturalist.org/
observations/7107050
IN 491088 P. bifasciatus ZIMBABWE: Hwange (Wankie) ‒18.865 26.433 B https://www.inaturalist.org/
observations/491088
IN 917967 P. bifasciatus TANZANIA: Haidom, Manyara ‒4.17685 35.02098 A https://www.inaturalist.org/
observations/917967
IN 917966 P. bifasciatus TANZANIA: Haidom, Manyara ‒4.17685 35.02098 B https://www.inaturalist.org/
observations/917966
IN 2706580 P. bifasciatus TANZANIA: Haidom ‒4.192396 35.013535 A https://www.inaturalist.org/
observations/2706580
IN 3585579 P. bifasciatus TANZANIA: Pangani ‒5.38343 38.971081 A https://www.inaturalist.org/
observations/3585579
IN 1158557 P. bifasciatus SOUTH AFRICA:
Iphiva Camp, St Lucia ‒28.352007 32.434076 A https://www.inaturalist.org/
observations/1158557

ALYTES 38 (1–4)
34
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
IN 1089785 P. bifasciatus SOUTH AFRICA: Hluhluwe ‒28.074789 32.270483 A https://www.inaturalist.org/
observations/1089785
IN 1041061 P. bifasciatus TANZANIA: Arusha ‒3.383354 36.712138 A https://www.inaturalist.org/
observations/1041061
n/a P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A Passmore & Carruthers (1979)
n/a P. bifasciatus SOUTH AFRICA: Naboomspruit ‒24.481563 28.715058 A Passmore & Carruthers (1979)
n/a P. bifasciatus BOTSWANA:
Unspecified Locality n/a n/a A Auerbach (1987)
n/a P. bifasciatus SOUTH AFRICA: Britts ‒25.60999 27.795971 A Pickersgill (2007)
n/a P. bifasciatus ESWATINI: Manzini ‒26.5082 31.371316 A Du Preez & Carruthers
(2009, 2017)
n/a P. bifasciatus MOZAMBIQUE: Pemba n/a n/a B Farook et al. (2014)
n/a P. bifasciatus ZIMBABWE: Harare ‒17.821629 31.049226 B This paper
n/a P. bifasciatus SOUTH AFRICA: Olievenhoutsrus ‒24.157348 28.079843 B This paper
n/a P. bifasciatus ZIMBABWE:
Muunze Forest Lodge ‒20.139996 31.069837 B This paper
n/a P. bifasciatus ZIMBABWE:
Muunze Forest Lodge ‒20.139996 31.069837 B This paper
n/a P. bifasciatus NAMIBIA: Okonjima ‒20.695690 16.525283 A/C This paper
FMNH 80628 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80629 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80627 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80635 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80632 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80631 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80638 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80634 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 C This paper
FMNH 80633 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80636 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 80637 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper

CERÍACO et al. 35
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
FMNH 190740 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 187346 P. bifasciatus
DEMOCRATIC REPUBLIC OF THE
CONGO: Upemba National Park,
Mabwe
‒8.65 26.516667 A This paper
FMNH 206462 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 205404 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 205396 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 2410 P. bifasciatus KENYA: Lukenya ‒1.483313 37.068721 A This paper
FMNH 129557 P. bifasciatus TANZANIA: Nachingwea District ‒10.5 38.3333 A This paper
FMNH 18006 P. bifasciatus BOTSWANA: Metsimaklaba Road n/a n/a A This paper
FMNH 18005 P. bifasciatus BOTSWANA: Metsimaklaba Road n/a n/a A This paper
FMNH 18004 P. bifasciatus BOTSWANA: Metsimaklaba Road n/a n/a A This paper
FMNH 18003 P. bifasciatus BOTSWANA: Metsimaklaba Road n/a n/a A This paper
FMNH 18002 P. bifasciatus BOTSWANA: Metsimaklaba Road n/a n/a A This paper
FMNH 18001 P. bifasciatus BOTSWANA: Molepolole ‒24.406587 25.495079 A This paper
FMNH 205399 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 206463 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 C This paper
FMNH 205403 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 205395 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 205394 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 205402 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 206464 P. bifasciatus SOUTH AFRICA: Pongola ‒27.383147 31.619753 A This paper
FMNH 205401 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 C This paper
FMNH 205398 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
FMNH 205405 P. bifasciatus ESWATINI: Tshaneni ‒26.001284 31.759438 A This paper
NMNW 25218 P. bifasciatus NAMIBIA:
Tsumkwe, Bushmanland ‒19.5833 20.4833 A This paper
NMNW 25624 P. bifasciatus NAMIBIA: Popa Falls, Kavango ‒18.1 21.56667 A This paper
NMNW 25683 P. bifasciatus NAMIBIA: Otjikaro, Otjiwarongo ‒20.61805 17.191074 A This paper
NMNW 25779 P. bifasciatus NAMIBIA:
Okosomingo, Otjiwarongo ‒20.623536 17.065656 A This paper
NMNW 25679 P. bifasciatus NAMIBIA:
Ombujoma-Temba, Otjiwarongo ‒20.637249 17.236565 A This paper
NMNW 25761 P. bifasciatus NAMIBIA:
Erindi, Osombaka, Otjiwarongo ‒21.28333 17.38333 A This paper
MHNCUP/AN
F 194 P. newtoni
sp. nov. ANGOLA: Chingo ‒11.2 13.85 D Ferreira (1904); this paper
BMHN
1873.7.28.14 P. newtoni
sp. nov. ANGOLA: Unspecified Locality n/a n/a D Boulenger (1882);
this paper
n/a P. newtoni
sp. nov. ANGOLA: Benfica, Luanda ‒8.972175 13.142503 D
https://www.facebook.com/
groups/BiodiversidadeAngola/
permalink/1263494427338968
this paper

ALYTES 38 (1–4)
36
Catalog nº /
Accession nº Taxon Locality Latitude Longitude Color
pattern
Source
All internet links consulted on
21 January 2021
n/a P. newtoni
sp. nov. ANGOLA: Calumbo ‒9.1499994 13.3991574 D
https://www.facebook.com/
groups/1045499302182009/
permalink/2326357597429500
this paper
n/a P. newtoni
sp. nov. ANGOLA: Longa ‒10.197147 13.519337 D
https://www.facebook.com/
groups/BiodiversidadeAngola/
permalink/1106083439746735
n/a P. newtoni
sp. nov. ANGOLA: Hote ‒11.319767 13.91653 D
https://biodiversidadeangola.
com/amphibians/scientific/
9441
Relevant specimens were measured with a digital caliper and coloration patterns
were noted. The following measurements (following Watters et al. 2016) were taken (to
the nearest 0.1 mm): snout-vent length (SVL); head width (HW) at the widest
point/angle of the jaws; head length (HL) from the posterior angle of the jaws to the tip
of the snout; snout length (SL) from the tip of the snout to the anterior corner of the eye;
interorbital distance (IOD) as the shortest distance between the anterior corners of the
orbits; internarial distance (IND) as the shortest distance between the inner margins of
the nostrils; eye-nostril distance (EN) from the anterior corner of the eye to the posterior
margin of the nostril; eye diameter (ED) horizontally from the anterior to posterior
corner of the eye; upper eyelid width (UEW) as the greatest width of the upper eyelid
margins; forearm length (FLL) from the flexed elbow to the base of the outer palmar
tubercle; hand length (HAL) from the base of the outer palmar tubercle to the tip Finger
IV; Finger IV disk width (Fin4DW) as the widest horizontal diameter of Finger IV; tibia
length (TL) as the distance from the outer surface of the flexed knee to the heel; thigh
length (THL) as the distance from the vent to the knee; and foot length (FL) from the
base of the inner metatarsal tubercle to the tip of Toe IV. Locality data are reported in
the form of decimal degrees and use the WGS 84 map datum. Older (non-GPS) records
are mostly derived from Marques et al. (2018) and have been georeferenced using
GEOLocate web application (http://www.geo-locate.org). Elevations are all reported as
meters above sea level.
RESULTS
Angolan Phrynomantis belong to three separate species: P. affinis, P. annectens
and a new species, historically referred as P. bifasciatus. The three taxa of
Phrynomantis occurring in Angola differ from each other and from other African
species, in several consistent morphological and coloration characters, and they are
identified through the key presented below. Our results support the observations of all
the other authors who previously dealt with Angolan population of P. bifasciatus
(Boulenger 1882; Bocage 1895; Ferreira 1904). All Angolan specimens present a
distinct coloration pattern that allows it to be differentiated from the nominotypical
form (Table 2), as well as from most of its congeners. This pattern consists of large
irregular yellowish, red or silver blotches, sometimes arranged in the appearance of an
interrupted dorsolateral band, whereas, despite some variation, all the specimens of P.
bifasciatus from across its known distribution in Northern Namibia, Botswana, Zambia,

CERÍACO et al. 37
Zimbabwe, South Africa, Mozambique, Tanzania and Kenya, either exhibit (A) two
well marked and uninterrupted orangish to reddish dorsolateral bands on a
homogeneous dark background (n = 95), (B) two well marked and uninterrupted
orangish to reddish dorsolateral bands, with faint vestiges of greyish dorsal bands
(n = 34), or (C) briefly interrupted orange dorsolateral bands on a homogeneous dark
background (n = 22) (Anonymous 2020; Table 2; Fig. 9). Even in the very few cases
where these bands are slightly interrupted, the band is always quite conspicuous,
contrary to the specimens from the Angolan population, where there is never a complete
band. The Angolan population is, therefore, here described as a new species (see
below). Consequently, P. bifasciatus is excluded from the list of currently known
species for the country. A detailed differential diagnosis of the newly described species
and its congeners is presented in the systematic account below. Our reexamination of
historical specimens, namely those cited by Monard (1938), resulted in a second record
and considerable range extension of P. affinis in the country. Newly collected
specimens of P. annectens contribute to knowledge of the species’ distribution range
with additional records in the southwestern areas of the country.
SYSTEMATICS
AMPHIBIA, ANURA, PHRYNOMERIDAE
Phrynomantis newtoni sp. nov.
(Fig. 7–10)
lsid:zoobank.org:act:33B59511-9D68-4DD3-A9B2-322ABCD63275
Phrynomantis bifasciata [part]: Boulenger (1882: 173), Bocage (1895: 181, Plate XVIII figure 3).
Phrynomantis bifasciatus: Ferreira (1904: 113), Marques et al. (2018: 80), Baptista et al. (2019: 273)
Figure 9. Live photo of Phrynomantis newtoni sp. nov. from Benfica neighborhood, city of Luanda.
Specimen photographed on 24 April 2020, but not collected. Photo by Eduardo Catumbela.

ALYTES 38 (1–4)
38
Holotype
MHNCUP/ANF-194 (old catalogue number MHNFCUP 127327; Fig. 7), male
adult, from Chingo (‒11.2º, 13.85º; 11 m), Kwanza Sul Province, Angola, collected by
Francisco Newton around May to July 1903.
Additional specimen
One specimen: BMNH 1873.7.28.14, unsexed adult without precise locality
(“Angola”), Angola, collected by Joachim John Monteiro on an unknown date.
Diagnosis
Phrynomantis newtoni sp. nov. has a small, short, blunt head and an elongated,
slightly pear-shaped body. Like all members of the genus, it has a smooth, rubber-like
dorsal skin. It can be distinguished from other members of the genus by the following
combinations of characters: (1) dorsal pattern consisting of large irregular yellowish, red
or silver blotches, sometimes arranged in the appearance of an interrupted dorsolateral
band; (2) tips of fingers expanded into large disks; (3) ventral coloration mottled with
dark markings; (4) throat dark in males.
Figure 10. Distribution map of the different color patterns (A, B, C) of Phrynomantis bifasciatus and of
Phrynomantis newtoni sp. nov. (D) in Africa. Purple circles: P. newtoni sp. nov.; Green diamonds: P.
bifasciatus color pattern A; Green triangles: P. bifasciatus color pattern B; Green circles: P. bifasciatus
color pattern C. Locality data derived from Table 2.

CERÍACO et al. 39
Description of the holotype
Small (SVL 41.1 mm), elongated and pear-shaped male specimen, with slender
limbs (Fig. 7); head blunt, wider (HW 13.5 mm) than long (HL 10.4 mm); snout
projecting beyond upper jaw (SL 4.5 mm); eyes projecting laterally beyond the eyelids
and approximately flush with margins of head in dorsal view; eye not projecting about
above dorsal margin of head in lateral view (UEW 1.6 mm); interorbital distance 1.2
times eye diameter; pupil large and round; loreal region concave; naris small; round,
directed dorsolaterally; canthus rostralis short; eye diameter (ED 3.7 mm) 1.15 times
eye-narial distance (EN 3.2 mm); eye diameter (ED 3.7 mm) 0.8 times naris to rostral
tip; internarial region bumpy; interorbital distance (IOD 4.3 mm) approximately two
times internarial distance (IND 2.3 mm), tympanum extremely small. Skin of limbs,
dorsal and dorsolateral surface of head and body smooth, rubber-like. Skin of venter
leathery; skin of gular region smooth.
Limbs (FLL 11.2 mm; HAL 11.7 mm; TL 12.9 mm; THL 12.9 mm; FL 13.6
mm) and digits well developed but relatively short when compared to the SVL; digits of
both manus and pes stout; relative length of fingers: III > II > IV = I; finger tips largely
expanded forming disks; fingers with rounded, prominent subarticular tubercles, two
palmar tubercles distinct and well separated from one another, one at ventromedial
surface of first finger and other at proximal plantar surface, latter being about two times
larger than first; webbing between manual digits absent; relative length of toes
IV > V = III > II > I; toe tips expanded forming disks; toes with prominent, single
subarticular tubercles; webbing between toes absent; prominent inner metatarsal
tubercle, length 40 % of first toe length.
Dorsal coloration consisting of large irregular cream blotches arranged in the
appearance of an interrupted dorsolateral band, on a dark-brown background; a large
irregular blotch above vent; smaller cream blotches on the upper and lateral sides of the
limbs; venter and ventral side of the limbs marbled with dark marking on a cream
background, throat dark; palm of hands and feet homogenous gray.
Photos of live specimens (see below) agree entirely with the holotype, but
blotches sometimes shows a more reddish to orangish coloration.
Comparison with other species of the genus
Comparing P. newtoni sp. nov. with P. bifasciatus, the newly described species
can be easily distinguished by having a dorsal pattern consisting of large irregular
yellowish, red or silver blotches, sometimes arranged in the appearance of an
interrupted dorsolateral band (versus dorsal pattern consisting either of (A), two well
marked and uninterrupted orangish to reddish dorsolateral bands on a homogeneous
dark background, (B), two well marked and uninterrupted orangish to reddish
dorsolateral bands, with faint vestiges of greyish dorsal bands, or (C), briefly interrupted
orange dorsolateral bands on a homogeneous dark background (see Passmore &
Carruthers 1979; Poynton & Broadley 1985; Auerbach 1987; Lambiris 1989; Channing
2001; Pickersgill 2007; Channing & Howell 2006; Spawls et al. 2006; Farook et al.
2014; Channing & Rödel 2019; see Table 2; see Fig. 4, 5, 10, 11). Comparing P.
newtoni sp. nov. with P. affinis, the newly described species can be easily distinguished
by having a dorsal pattern consisting of large irregular blotches (versus small orange or

ALYTES 38 (1–4)
40
red spots in P. affinis) and the tips of fingers expanded into disks (versus tips enlarged
but not forming disks in P. affinis; see Fig. 1b). Comparing it with P. annectens, the
newly described species can be easily distinguished by its larger blotches, having the
tips of fingers expanded into round disks (versus tips of fingers expanded forming small
truncated disks in P. annectens, see Fig. 2b) and by having the ventral coloration
mottled with dark markings (versus venter homogeneous pinkish-brown without spots
in P. annectens). Comparing it with P. microps, the newly described species can be
easily distinguished by having a dorsal pattern consisting of large irregular blotches
(versus a dorsal pattern entirely red, sometimes with a fine median black line in P.
microps). Phrynomantis newtoni sp. nov. can be distinguished from P. somalicus by
having the tips of fingers expanded into large disks (versus tips of fingers expanded
forming small truncated disks in P. somalicus).
Distribution
The species is known from low elevation (below 20 m) in the coastal areas of
southern and central Angola, from Benguela to Luanda Province (Fig. 12). We
tentatively accept the identification of the Quissange specimen cited by Bocage as
representative of Boulenger’s “var. C” as a representative of the new species, but refrain
from assigning the Benguela specimen to any form. As both of these specimens were
lost in the fire that destroyed the Lisbon collection in 1978, these identifications cannot
be confirmed without newly collected material from the same areas.
Habitat and natural history notes
The dominant habitats where the specimens have been collected is a typical
western Angolan savannah, with sandy soils dominated by Adansonia digitata,
Euphorbia conspicua, Acacia welwitschi and Combretum spp., together with a good
grass coverage (Grandvaux-Barbosa 1970).
Etymology
The specific epithet newtoni is formed in the genitive masculine singular and
honors the Portuguese naturalist and explorer Francisco Newton (1864–1909), who
collected the holotype. We propose the Portuguese common name of “Rã de Borracha
de Newton”, and the English common name of “Newton’s Rubber Frog”.

CERÍACO et al. 41
Figure 11. Species of the genus Phrynomantis Peters, 1867. 1, P. affinis; 2, P. annectens; 3, P. microps;
4, P. newtoni sp. nov.; 5, P. bifasciatus; 6, P. somalicus. All depicted specimens are males.

ALYTES 38 (1–4)
42
Key to the genus Phrynomantis
Asterisks (*) denote the presence of the species in Angola.
(1a) Dorsal pattern generally consisting on two continuous broad orange to red dorsolateral bands from
the snout to the hind limbs ............................................................................................... P. bifasciatus
(1b) Dorsal pattern without continuous broad dorsolateral bands .............................................................. 2
(2a) Dorsal pattern entirely red, sometimes with a fine median black line ................................. P. microps
(2b) Dorsal pattern consisting of spots or blotches ..................................................................................... 3
(3a) Tips of fingers not expanded, dorsal pattern consisting of small spots ................................. P. affinis*
(3b) Tips of fingers expanded, dorsal pattern consisting of irregular blotches ........................................... 4
(4a) Tips of the fingers expanded forming large rounded disks .................................. P. newtoni sp. nov.*
(4b) Tips of fingers expanded forming small truncated disks .................................................................... 5
(5a) Dorsal coloration marbled with large, irregular-shaped silver, golden, yellow, orange, pink, or red
spots on dark background ................................................................................................ P. annectens*
(5b) Dorsal coloration consisting in discontinuous dorsolateral bands, composed of irregular blotches,
starting on the snout and extending to the hind limbs ........................................................ P. somalicus
Figure 12. Distribution map of the three species of Phrynomantis in Angola and general distribution of
the species in Africa. Blue circles: confirmed records of P. affinis; open red triangles: confirmed records
of P. annectens; dotted red triangles: historical records of P. annectens; open purple pentagons:
confirmed records of P. newtoni sp. nov.; dotted purple pentagon: historical records of P. newtoni sp.
nov.; purple star: type locality of P. newtoni sp. nov. Historical records from Marques et al. (2018),
confirmed records from this paper (Table 1). In the general map, blue: P. affinis; red: P. annectens; bright
green: P. bifasciatus; orange: P. microps; purple: P. newtoni sp. nov.; yellow: P. somalicus.

CERÍACO et al. 43
DISCUSSION
Although modern species descriptions without the use of molecular supporting
data are becoming less common due to the uncertainty surrounding the value of some
morphological characters, for the case of Phrynomantis species, and especially for
Phrynomantis newtoni sp. nov., the available morphological data, but especially
coloration patterns, are sufficiently convincing and stable to support its recognition as a
separate species. This is also supported when considering the biogeographic context of
its known distribution. As noted by Marques et al. (2018), the coastal lowlands of
Angola represent an overlooked hotspot of herpetological endemism. Several species of
reptiles are endemic from this region, as for example Hemidactylus longicephalus
Bocage, 1873, Hemidactylyus bayonii Bocage, 1893, Sepsina bayoni (Bocage, 1866),
Sepsina copei Bocage, 1873, Agama mucosoensis Hellmich, 1957, and Boaedon
bocagei Hallermann et al. 2020 (see Marques et al. 2018; Ceríaco et al. 2020a;
Hallermann et al. 2020). All these species have their known distributions limited by the
Atlantic Ocean in the West and the Angolan Escarpment in the East, being endemic to
the thin strip of coastal lowlands that begins north of Luanda and extends south to the
Namibe Province. Many of these species have their closest relatives in the plateau
regions (e.g. Hemidactylus paivae Ceríaco et al. 2020, Sepsina angolensis Bocage,
1866) making them interesting examples of the role of the Angolan Escarpment in
promoting the isolation and diversification of herpetological taxa. In this area, the rapid
transition from a low elevation area towards a more elevated one, is accompanied by an
abrupt habitat change, which may act as a barrier. Due to the lack of molecular data, it is
currently impossible to evaluate the phylogenetic relationship of the newly described
species to the remaining members of the genus. Although, on the basis of molecular
data, Dubois et al. (2021: 416) retrieved the species P. annectens as sister to the well-
supported pair of species P. bifasciatus and P. microps, there is currently no available
phylogeny of the whole genus, and therefore the intra-generic relationships are still
partially unknown. Further surveys in Angola are needed in order to secure fresh
material that will allow placing the newly described species in its phylogenetic context.
The arguments that support the recognition of newly described species (its
coloration pattern and geographic distribution) were retrieved from a combination of
two completely different and contrasting, data sources: (1) modern online citizen
science platforms (iNaturalist) and social media webpages (Facebook), and (2)
historical specimens housed in museum collections. This is not the first species to be
described after someone had shared a photo on a social media platform. Very recently a
new species of millipede fungus was described after a photo was shared on the social
media network Twitter (Santamaria et al. 2020). As a tool used by millions of people
around the world, some of the shared photos by amateurs and citizen scientists can
provide important hints and serve as a source of novel data for researchers. Although
these photos by themselves can never substitute for the actual specimens (Ceríaco et al.
2016), they can provide further evidence to augment specimen-based research.
The fact that the newly described species is based on the study of historical
specimens also highlights the importance of historical, and sometimes overlooked,
natural history collections for modern biodiversity studies. As noted by Fontaine et al.
(2014), many new species wait for decades on museum shelves until they are properly
studied by competent taxonomists and subsequently described. For the case of

ALYTES 38 (1–4)
44
Phrynomantis newtoni sp. nov., the specimens were available for researchers for more
than a century in the BMNH and the MHNCUP. Although the BMNH is amongst the
largest natural history collections in the world, that is certainly not the case of the
MHNCUP. Despite their size, small collections can have disproportionate importance
(Winker & Withrow 2013). This is certainly the case of the MHNCUP collections,
whose number of type specimens is considerable for its size (see Ceríaco et al. 2014),
but also harbour interesting and rare specimens, such as the one used as the holotype of
the newly described species. Within the MHNCUP collections, the specimens collected
by Francisco Newton during his 1903–1905 expedition to Angola are of special
relevance for the study of the herpetofauna of the whole region. This collection was
studied in the past by Ferreira (1904, 1906) but was then subsequently abandoned and
forgotten by the scientific community until Ceríaco et al. (2014) published its type
catalog. The collection has recently been reviewed and a detailed and commented list of
its specimens published (Santos et al. in press). Besides the description of Phrynomantis
newtoni sp. nov., other specimens are currently being used as type material for the
description of new reptile taxa (Hallermann et al. 2020). But the study of historical
material results not only on the description of new species but also in the critical
revision of previously misidentified records. These revisions have serious implications
for the understanding of species distributions, which by themselves have different
consequences for a vast array of topics and areas, such as conservation, national
checklists, species distribution model-based studies, etc. The revision of historical
material allowed us to rectify the most recent available checklists for Angolan
amphibians (Marques et al. 2018, Baptista et al. 2019), by adding a new endemic
species for the country and taking P. bifasciatus from the confirmed species list, with
far reaching consequences for the understanding of the species, distribution throughout
the continent. These new data contradict the common belief that the species was likely
distributed through Angola, as assumed by Ruas (1996) and subsequently accepted by
Channing (2001), Marques et al. (2018) and Channing & Rödel (2019). However, we
tentatively suggest that the species may still be found in the country, especially in the
far-eastern regions of Moxico and Cuando-Cubango provinces, as there are several
confirmed records of P. bifasciatus for Zambia and northeastern Namibia (Anonymous
2020).
ACKNOWLEDGMENTS
The present work is a result of the ongoing collaboration between the Instituto
Nacional da Biodiversidade e Áreas de Conservação (INBAC) from the Ministry of
Environment of Angola and its international partners. INBAC provided institutional and
logistical support, as well as the necessary permits for carrying out the field surveys that
led to the collection of some specimens used on this research. We thank António
“Muloge” Lopes, Arianna L. Kuhn, Jens Vindum, Joyce Janota, Suzana Bandeira,
Hilária Valério and Álvaro “Varito” Baptista for their support and camaraderie during
field work. We want to thank to the “Angola Ambiente” Facebook group, in the person
of its administrators Sara Fernandes, David Elizalde and Luis Querido for providing an
important platform for sharing data and photos about Angolan biodiversity, and to
Manuel Castanho and Eduardo Catumbela for allowing us to use his photo, previously
published in the Facebook group. François Becker and Harith Farook kindly provided

CERÍACO et al. 45
photos of P. bifasciatus. The following collection managers and curators are thanked for
providing data and permission to visit and use the collections their care: Lauren
Scheinberg and Erica Ely from the California Academy of Sciences (CAS), USA; Alan
Resetar, Joshua Mata and Adam Fergunson from the Field Museum of Natural History
(FMNH), USA; Jeff Streicher from the Natural History Museum of London (BMNH),
United Kingdom; Garin Cael from the Royal Museum for Central Africa (RMCA),
Belgium; Arnaud Maeder and Nicolas Margraf from the the Musée d’Histoire Naturelle
de La Chaux-de-Fonds (MHNC), Switzerland; François Becker and Bertha Buiswalelo
from the National Museum of Namibia (NMNW), Namibia; and Ilunga André from the
Museu Regional do Dundo (MD), Angola. This work was funded by grant from the JRS
Biodiversity Foundation to AMB and David C. Blackburn. MPM is currently supported
by FCT, contract SFRH/BD/129924/2017. Annemarie Ohler, Alan Channing and Mark-
Oliver Rödel provided important insights and comments to the first version of this
manuscript.
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Submitted: 26 August 2020.
Accepted: 7 April 2021.
Published: 21 July 2021.
Corresponding editor: Annemarie Ohler.