Phrynobatrachus cryptotis, Schmidt and Inger, 1959

Marques, Mariana P., Ceríaco, Luis M. P., Blackburn, David C. & Bauer, Aaron M., 2018, Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola Atlas of Historical and Bibliographic Records (1840 - 2017), Proceedings of the California Academy of Sciences 65, pp. 1-501 : 144-148

Phrynobatrachus cryptotis group

Given the considerable nomenclatural and taxonomic confusion regarding P. cryptotis , P. mababiensis , P. minutus and P. parvulus , we have opted to combine all of their distributional and taxonomic notes into the same account, although, specific accounts for each one, with the list of chresonyms and type specimen data still follow below. Records attributed to each species are provided below separately, indicated by the name and noting the different symbols used to plot the records on the combined map.

Ocurrences in Angola (Map 94):

Phrynobatrachus cryptotis (plotted as circles). Lunda Sul: “Alto Cuílo” [-10.01667,

19.55000] ( Laurent 1964a:144; Ruas 1996:27);

“Alto Chicapa” [-10.88333, 19.23333] ( Laurent 1964a:144; Ruas 1996:27). Huíla: “ Boca de Humpata ” [-14.93333, 13.51667] ( Laurent

1964a:144; Ruas 1996:27). Undetermined

Locality: “arid territories along the coast” ( Cei

1977:18). Phrynobatrachus mababiensis

(plotted as triangles). Bié: “Cuando basin

(11)” [-13.69413, 17.06177] ( Conradie et al.

2016:8-9,16); “Cuando basin (12a)”

[-13.59333, 16.87986] ( Conradie et al. 2016:8 -

9,16). Huambo: “Cuando basin (18)”

[-12.67105, 16.11111] ( Conradie et al. 2016:8 -

9,16). Huíla: “Nuntechite lagoon” [-15.13333,

13.41667] ( Poynton and Haacke 1993:14; Ruas MAP 94. Distribution of Phrynobatrachus cryptotis group in Angola.

1996:27). Cuando Cubango: “Cuando basin

(3)” [-14.94277, 17.71863] ( Conradie et al. 2016:8 -9,16); “Cuando basin (6b)” [-14.67458,

17.73544] ( Conradie et al. 2016:8 -9,16); “Cuando basin (19)” [-14.70213, 17.37772] ( Conradie et al. 2016:8 -9,16); “Cuando basin (21)” [-14.66586, 17.07661] ( Conradie et al. 2016:8 -9,16); “Cuando basin (22c)” [-14.65386, 16.93547] ( Conradie et al. 2016:8 -9,16); “Cuando basin (22d)” [-14.64991, 16.90739] ( Conradie et al. 2016:8 -9,16); “Cuando basin (29)” [-17.87291, 19.833333] ( Conradie et al. 2016:9 -10,16); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016: 9 -10,16); “Cuando basin (38)” [-17.58830, 22.65694] ( Conradie et al. 2016:9 -10,16); “Cuando basin (39)” [-17.46333, 22.86638] ( Conradie et al. 2016:9 -10,16); “Cuando basin (40)” [-17.45786, 22.91191] ( Conradie et al. 2016:16); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9 -10,16); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9 -10,16). Phrynobatrachus minutus (plotted as pentagons). Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1954a:74; Ruas 1996:27); “galerie forestière de la Luachimo (Luachimo)” [-7.38333, 20.85000] ( Laurent 1950a:15; Ruas 1996:27); “Marrura rive droite de la Tshikapa, 50 km sud-oest de Dundo (Marrora)” [- 7.60000, 20.51667] ( Laurent 1954a:74; Ruas 1996:27). Cunene: “Donguena (Cunene)” [-17.01667, 14.71667] ( Laurent 1954a:74; Ruas 1996:27). Undetermined Locality: “Station Quanza (Estação do Quanza)” ( Laurent 1954a:74; Ruas 1996:27); with no precise locality ( Laurent 1954a:74); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “plateaus regions” ( Cei 1977:17). Phrynobatrachus parvulus (plotted as diamonds). Lunda Sul: “Alto Cuílo, Poste de Cacolo, ruisseau cascade Ná-Ipanha” [-10.00000, 19.58333] ( Laurent 1964a:144); “Alto Cuílo, rives du Cuílo (Cuílo)” [-10.01667, 19.55000] ( Laurent 1964a:144); “Alto Chicapa, humidiherbosa des sources de la Kamutongola” [-10.88333, 19.25000] ( Laurent 1964a:144). Moxico: “Teixeira de Sousa” [-10.70000, 22.23333] Mertens (1937:20). Malanje: “Bange N’golla (Dange)” [-8.43333, 16.56667] ( Boulenger 1905:109; Loveridge 1933:386; Marx 1958:425; Schmidt and Inger 1959:160; Poynton and Broadley 1985b:169; Frost 1985:449, 2016; Ruas 1996:27). Kwanza Sul: “Calaongo below Congulu (Carlaongo)” [-10.78333, 14.26667] ( Parker 1936:142; Ruas 1996:27). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1936:131; Ruas 1996:27); “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:131, Ruas 1996:27); “General Machado” [-12.03333, 17.46667] Mertens (1937:20). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937a:58, 1938:118, Ruas 1996:27). Benguela: “Cubal” [-13.03333, 14.25000] Mertens (1937:20). Cuando Cubango: “Cuando basin (43)” [-17.53500, 23.189916] ( Conradie et al. 2016:9 -10,17). Undetermined Locality: “without precise locality” ( Schmidt 1936:131).

Taxonomic and distributional notes: Opinions in the literature vary as to whether three commonly reported species from Angola are conspecific: P. mababiensis FitzSimons, 1932 , P. parvulus ( Boulenger, 1905) and P. minutus (Boulenger, 1895) . Loveridge (1953) assigned a specimen previously identified as P. parvulus to P. mababiensis , suggesting that P. parvulus was a western relative of P. mababiensis or perhaps a synonym of Phrynobatrachus minutus . Poynton and Broadley (1985b:165, 170) discussed difficulties in separating P. parvulus and P. mababiensis based on external morphology. Zimkus et al. (2010) recently used molecular phylogenetic analyses to reveal that there are probably three different species currently recognized as P. mababiensis . If P. minutus is known only from Ethiopia, it is highly likely that the Angolan records assigned to this species are misidentified and instead belong to one of the other three species. With some reservations, Laurent (1964a) cited P. cryptotis for Angola, though he recognized that these might represent P. parvulus instead, which is a similar species (Drewes and Vindum 1994). It is likely that the Angolan specimens mapped here represent a combination of P. mababiensis and P. parvulus .

Phrynobatrachus cryptotis Schmidt and Inger, 1959 CRYPTIC RIVER FROG

Phrynobatrachus cryptotis Schmidt and Inger 1959:143 , fig. 60, pl. 5, fig. 5. Holotype: IRSNB 1.493 (collector G.F. de Witte), formerly Institut des Parcs Nationaux du Congo Belge n. 1926. Type locality: “upper Bwalo River (an affluent from the left in on the left of the Muye, which is an affluent from the right of the Lufira)” (Schmidt and Inger 1959:143), [= Parc National de l’Upemba, upper Katanga], Democratic of Republic of Congo.

Phrynobatrachus cryptotis : Laurent (1964a:144), Cei (1977:18), Frost (1985:445, 2016), Ruas (1996:27), Frétey et al. (2011:39).

Global conservation status (IUCN): Data Deficient.

Global distribution: It is known with certainty only from the Democratic Republic of Congo.

Phrynobatrachus mababiensis FitzSimons, 1932 MABABE PUDDLE FROG

Phrynobatrachus mababiensis FitzSimons 1932:40 . Holotype: TM 14838, formerly Vernay-Lang Khalari Expedition no. 989 (collector V.F. FitzSimons). Type locality: “Tsotsoroga Pan, Mababe Flats” ( FitzSimons 1932:40), South Africa.

Phrynobatrachus mababiensis : Poynton and Broadley (1985b:165), Forst (1985:447, 2016), Poynton (1992:67), Poynton and Haacke (1993:14), Ruas (1996:27), Channing (2001:305), Channing and Howell (2006:284), Zimkus et al. (2010:42), Frétey et al. (2011:39), Channing et al. (2012:278), Conradie et al. (2016:16) .

Global conservation status (IUCN): Least Concern.

Global distribution: This species occurs in savannas across southern and central Africa.

Phrynobatrachus minutus (Boulenger, 1895) ETHIOPIAN DWARF PUDDLE FROG

Arthroleptis minutus Boulenger 1895a:539 , pl. 30, fig. 4. Holotype: BMNH 1947.2.30.51 (collector A.D. Smith), formerly 95.6.11.8 fide Grandison in Frost (1985:448). Type locality: “Durro” ( Boulenger 1895a:539), [= Duro], Ethiopia.

Arthroleptis minutus : Monard (1938:118).

Phrynobatrachus minutus : Laurent (1950a:15, 1954a:74), Loveridge (1957:349), Inger (1959:540), Cei (1977:16, 17), Ruas (1996:27), Frétey et al. (2011:39), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: This species is likely limited to Ethiopia.

Phrynobatrachus parvulus ( Boulenger, 1905) DWARF RIVER FROG

Arthroleptis parvulus Boulenger 1905:109 , pl. 4, figs; 3, 3b. Syntypes: BMNH 1947.2.6.93-95 [3 specimens], formerly 1904.5.2.97-99 and FMNH 75381 (collector W.J. Ansorge) fide Grandison in Frost (1985:449). Type locality: “Bange N’golla” ( Boulenger 1905:119), [= Dange] Malanje Province, Angola.

Arthroleptis parvulus : Loveridge (1933:386), Schmidt (1936:131), Parker (1936:142), Mertens (1937:20).

Arthroleptis minutus : Monard (1937a:58, 1938:118).

Phrynobatrachus parvulus : Marx (1958:425), Schmidt and Inger (1959:160), Laurent (1964a:144), Poynton and Broadley (1985b:169), Frost (1985:449, 2016), Poynton (1992:67), Ruas (1996:27), Channing (2001:308), Channing and Howell (2006:289), Frétey et al. (2011:40).

Phrynobatrachus cf. parvulus : Conradie et al. (2016:17) .

Global conservation status (IUCN): Least Concern.

Global distribution: This species occurs in savannas across southern and central Africa.

Phrynobatrachus natalensis (Smith, 1849) NATAL DWARF PUDDLE FROG

Stenhorhynchus natalensis Smith 1849b:24 . Holotype: BMNH 1947.2.5.13 (formerly BMNH 62.3.14.20) (collector A. Smith) fide Grandison in Frost (1985:448). Type locality: “the country around Port Natal” [= Durban, KwaZulu-Natal], South Africa.

Phrynobatrachus natalensis : Günther (1865a:481), Bocage (1866a:54, 1870:68, 1879c:89, 1895a:162, 1897b:211), Boulenger (1882:112, 1905:108), Ferreira (1904:113, 1906:166), de Witte (1919:223), Schmidt (1936:130), Monard (1937a:57, 1938:117), Mertens (1937a:20), Parker (1939:142), Laurent (1950a:15, 1954a:74, 1964a:143), Loveridge (1957:348), Hellmich (1957a:24), Schmidt and Inger (1959:155), Inger (1959:540), Cei (1977:16, 17, 18), Poynton and Haacke (1993:14), Ruas (1996:27, 2002:145), Largen (2001:348), Channing (2001:307), Zimkus et al. (2010:886), Frétey et al. (2011:39), Frost (2016), Ceríaco et al. (2016b:39), Conradie et al. (2016:17) .

Phrynobatrachus cf. natalensis : Ceríaco et al. (2014b:669).

Global conservation status (IUCN): Least Concern.

Global distribution: Populations referred to P. natalensis occur across sub-Saharan Africa, including in Angola, though this taxon likely contains multiple undescribed species.

Ocurrences in Angola (Map 95): The species is widespread across nearly all of

Angola. Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1950a:15, 1954a:74; Ruas

1996:27); “Andrada (Luembe O)” [-7.70000,

21.38333] ( Laurent 1954a:74; Ruas 1996:27);

“ Muita (Luembe E)” [-7.80000, 21.45000]

( Laurent 1950a:15, 1954a:74; Ruas 1996:27).

Lunda Sul: “Dala” [-11.03333, 20.20000]

( Monard 1937a:57, 1938:117; Ruas 1996:27);

“Alto Cuílo, rives du Cuílo (rio Cuílo)”

[-10.01667, 19.55000] ( Laurent 1964a:143;

Ruas 1996:27). Malanje: “Bange N’golla

(Dange)” [-8.43333, 16.56667] ( Boulenger

1905:108; Ruas 1996:27); “Duque de Bragança ” [-9.10000, 15.95000] ( Bocage 1866a:54,

1895a:162; Günther 1865a:481; Boulenger

1882:112; de Witte 1919:223; Poynton and MAP 95. Distribution of Phrynobatrachus natalensis in Angola.

Haacke 1993:14; Ruas 1996:27); “ 16 km SE of

Quissol - Garibo” [-9.63333, 16.41667] (Poynton and Haacke 1993:14; Ruas 1996:27); “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:39); “Capanda” [-11.58333, 17.61667] (Ceríaco et al. 2014b:669); “ Reserva da Palanca Preta (nascentes do rio Cuanza)” [-11.11667, 17.46667] ( Ruas 2002:145); “ Rio Caluando (nascente)” [-11.11667, 17.46667] ( Ruas 1996:27, 2002:145). Moxico: “Teixeira de Sousa” [-10.70000, 22.23333] (Mertens 1937:20); “Posto Bussaco” [-11.43333, 20.16667] ( Ruas 2002:145; Ruas 1996:27); “Rives du lac Calundo (Lago Calundo)” [-11.80000, 20.86667] ( Laurent 1964a:143; Ruas 1996:27); “ Rio Calombe (Reserva da Palanca Preta)” [-11.83333, 19.93333] ( Ruas 1996:27, 2002:145); “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:143; Ruas 1996:27). Kwanza Norte: “Mucoso” [-9.53333, 14.65000] ( Hellmich 1957a:24). Bengo: “Catete” [-9.11667, 13.70000] ( Ferreira 1904:113; Ruas 1996:27). Kwanza Sul: “Congulu (Congulo)” [-10.86667, 14.28333] ( Parker 1936:142; Ruas 1996:27). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1936:130; Ruas 1996:27); “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:130; Ruas 1996:27); “General

Machado” [-12.03333, 17.46667] ( Mertens 1937a:20); “Bihé” [-12.38333, 16.95000] ( Bocage 1879a:89, 1895a:162; de Witte 1919:223; Ruas 1996:27); “Cubango basin (10)” [-13.71616, 17.09661] ( Conradie et al. 2016:8 -9,17); “Cubango basin (12a)” [-13.59333, 16.87986] ( Conradie et al. 2016:17); “Cubango basin (17)” [-12.57008, 16.49111] ( Conradie et al. 2016:8 -9,17). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937a:57, 1938:117; Ruas 1996:27). Benguela: “Quissange” [-12.43333, 14.05000] ( Bocage 1895a:162; de Witte 1919:223; Ruas 1996:27); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:162; de Witte 1919:223; Ruas 1996:27); “Ebanga” [-12.73333, 14.73333] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:24); “Hanha” [-13.30000, 14.20000] ( Bocage 1897b:211; Ruas 1996:27). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:162; de Witte 1919:223; Ruas 1996:27); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Sangevé” [-13.88333, 15.83333] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Mukoti” [-14.20000, 15.80000] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Boca de Humpata” [-14.93333, 13.51667] ( Laurent 1964a:143; Ruas 1996:27); “Kangela (Kului)” [-15.41667, 15.73333] ( Monard 1937a:57; 1938:117; Ruas 1996:27). Namibe: “ Cunene mouth (Foz do Cunene)” [-17.28333, 11.80000] (Poynton and Haacke 1993:14; Ruas 1996:27). Cunene: “ruisseau Mbalé (Bale)” [-15.16667, 16.75000] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Kuvelaï” [-15.65000, 15.80000] ( Monard 1937a:57, 1938:117; Ruas 1996:27); “Donguena” [-17.01667, 14.71667] ( Laurent 1954a:74; Ruas 1996:27). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] ( Monard 1938:117; Ruas 1996:27); “Cubango basin (3)” [-14.94277, 17.71863] ( Conradie et al. 2016:8 -9,17); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8 -9,17); “Cubango basin (19)” [-14.70213, 17.37772] ( Conradie et al. 2016:8 -9,17); “Cubango basin (21)” [-14.66586, 17.07661] ( Conradie et al. 2016:17); “Cubango basin (22d)” [-14.66278, 16.96081] ( Conradie et al. 2016:17). Undetermined Locality: without precise locality ( Bocage 1870:68; Ferreira 1906:166; Schmidt 1936:130); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “plateaus regions” ( Cei 1977:17); “arid territories along the coast” ( Cei 1977:18).

Taxonomic and distributional notes: Schmidt and Inger (1959) provided a distribution map for Phrynobatrachus natalensis (Smith, 1849) with some records in Angola, though without additional information. The molecular phylogenetic analysis by Zimkus et al. (2010) revealed Phrynobatrachus natalensis to be a species complex, with Angolan records likely corresponding to their lineage “ natalensis A.”

Phrynobatrachus plicatus (Günther, 1858) COAST RIVER FROG

Hyperolius plicatus Günther 1858a:326 . Holotype: BMNH 1947.2.29.97 [wrongly given as 1947.2.29.47 by Grandison in Frost (1985)], formerly BMNH 49.10.9.1 fide Grandison in Frost (1985:449) (collector unknown). Holotype is depicted in Günther “1858b” 1859:8, pl. 7, fig. C. Type locality: “ Guinea ” ( Günther 1858a:326), later modified to “Coast of Guinea ” (Günther “1858b” 1859:88).

Arthroleptis plicatus : Peters (1877a:618).

Phrynobatrachus plicatus : Zimkus et al. (2010:896), IUCN SSC Amphibian Specialist Group (2013c), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: This is a widespread species from western Africa.

Ocurrences in Angola (Map 96): The only record of this species in Angola is from the Cabinda enclave . Cabinda: “Chinchoxo” [-5.10000, 12.10000] ( Peters 1877a:618) .

Taxonomic and distributional notes: The r ecord of this species in Angola is almost certain-