Sclerophrys Tschudi, 1838

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 : 69-78

Genus Sclerophrys Tschudi, 1838

Sclerophrys buchneri (Peters, 1882) BUCHNER’S TOAD

Bufo Buchneri Peters 1882b:147 . Holotype: ZMB 10103 (collector M. Buchner). Type locality: “Lunda (Africa occidentalis),” Angola.

Bufo buchneri : Bauer et al. (1995:40).

Bufo funereus : Tandy and Keith (1972:158), Ruas (1996:22, 2002:142).

Amietophrynus buchneri : Frétey et al. (2011:22).

Sclerophrys buchneri : Frost (2016).

Global conservation status (IUCN): Data Deficient.

Global distribution: The species is known from eastern Angola, but it may also occur in Republic of the Congo and western Democratic Republic of Congo ( Frost 2016).

Ocurrences in Angola (Map 13): Within Angola, this species is known only from the type locality “Lunda” in eastern Angola . Lunda

Sul: “Lunda” [-10.96667, 20.06667] ( Peters

1882b:146; Bauer et al. 1995:40; Ruas

1996:22, 2002:142).

Taxonomic and distributional notes:

Various authors, including Tandy and Keith

(1972), have suggested treating S. buchneri as a synonym of S. funerea (Bocage, 1866) , which was followed by Ruas (1996, 2002). Both the lack of recent data and the uncertainty regarding the taxonomic status of this species indicate that more work is needed on this species.

Boulenger (1904) recognized Bufo decorsei

Mocquard 1903, described from Brazzaville in

Republic of Congo, as a junior synonym of S.

buchneri. If true, this suggests that this species is more broadly distributed in Central Africa.

MAP 13. Distribution of Sclerophrys buchneri in Angola. We follow Ohler and Dubois (2016) in using the senior synonym Sclerophrys Tschudi, 1838 for all African bufonids recently assigned to Amietophrynus Frost et al. 2006 .

Sclerophrys funerea (Bocage, 1866) ANGOLAN TOAD

Bufo funereus Bocage 1866a:56 1866b:77. Holotype: MBL 7.5-363 (collector F. A. P. Bayão) fide Perret (1976a:17), destroyed by fire 18 March 1978. Type locality: “Duque de Bragança” [= Calandula] Malanje Province, Angola.

Bufo benguelensis Boulenger (1882:299) . Syntypes: BMNH 1872.2 .15.3. Type locality: “ Benguela ” [ Angola], “Fernando Pó” [= Bioko , Guinea] and “West Afrika.”

Bufo funereus : Bocage (1882a:304, 1895a:186, 1897a:205), Ferreira (1904:114, 1906:166), Monard (1937a:27, 1938:55, 79), Mertens (1937a:18), Themido (1941:2), Laurent (1950a:13, 1954a:71), Loveridge (1957:311), Schmidt and Inger (1959:20), Perret (1976a:17), Cei (1977:16, 17), Frost (1985:45), Ruas (1996:21, 2002:142), Channing

(2001:69).

Bufo funereus funereus : Laurent (1964a:131).

Amietophrynus funereus : Frétey et al. (2011:23),

Channing et al. (2012:129).

Sclerophrys funerea : Frost (2016), Conradie et al.

(2016:12).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from West-Central Africa from Gabon to Uganda, Burundi and Rwanda, southward to Democratic Republic of Congo and Angola.

Ocurrences in Angola (Map 14): Reports for this species range across much of Angola , extending from the southwestern to northeastern regions. Bengo: “Cabiri” [-8.91667, 13.66667] ( Ferreira 1904:114; Monard MAP 14. Distribution of Sclerophrys funerea in Angola.

1938:55). Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1950a:13; Ruas 1996:21); “Muita, Luembe E (Muíta)” [-7.80000, 21.45000] ( Laurent 1950a:13; Ruas 1996:21, 2002:142). Lunda Sul: “Alto Cuílo, rives du Cuílo, Lunda” [-10.01667, 19.55000] ( Laurent 1964a:131; Ruas 1996:22, 2002:142); “Dala” [-11.03333, 20.20000] ( Monard 1938:55, 79; Ruas 1996:21, 2002:142); “Alto Chicapa, Lunda” [-10.88333, 19.23333] ( Laurent 1964a:131; Ruas 1996:22, 2002:142). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] ( Ferreira 1904:114; Monard 1938:55; Ruas 1996:21, 2002:142); “Gumba” [-11.26667, 14.28333] ( Ferreira 1904:114; Monard 1938:55; Ruas 1996:21, 2002:142); “Lembu (Lembe)” [-12.86667, 14.11667] ( Ferreira 1904:114; Monard 1938:55; Ruas 1996:21, 2002:142). Malanje: “Duque de Bragança” [-9.10000, 15.95000] ( Bocage 1866a:56, 1866b:77, 1882a:304, 1895a:186, 1897a:205; Loveridge 1957:311; Perret 1976a:17; Frost 1985:45, 2016; Ruas 1996:21, 2002:142). Moxico: “Calombe, Luso” [-11.83333, 19.93333] ( Ruas 1996:22, 2002:142); “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:131; Ruas 1996:22). Bié: “General Machado” [-12.03333, 17.46667] ( Mertens 1937a:18). Huambo: “Serra do Moco (Luimbale) ” [-12.50000, 15.16667] ( Laurent 1954a:71; Ruas 1996:21, 2002:142). Benguela: “Benguella” [-12.58333, 13.41667] ( Boulenger 1882:299; Loveridge 1957:311). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1882a:304, 1895a:186, 1897a:205; Monard 1938:55; Themido 1941:2; Perret 1976a:17; Ruas 1996:21, 2002:142); “Mukoti” [-14.20000, 15.80000] ( Monard 1937a:27, 1938:55, 79; Ruas 1996:21, 2002:142). Cunene: “Ruisseau Mbalé (Rio Bale)” [-15.16667, 16.75000] ( Monard 1937a:27, 1938:55, 79; Ruas 1996:21, 2002:142). Cuando Cubango: “Kakindo (Caquindo)” [-15.45000, 17.05000] ( Monard 1937a:27, 1938:55, 79; Ruas 1996:21, 2002:142); “Cubango system around the villages of Kakindo [= Caiundo] and Mbale” [-12.85000, 18.29028] ( Conradie et al. 2016:12). Undetermined Locality: “without precise locality” ( Ferreira 1906:116); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: The species was noted as new by Bocage in two different papers (1866a,b), both of which appeared in the November 1866 issue of the same journal. Unlike many of the nomina in 1866a, however, that of Bufo funereus is a nomen nudum as there is no description. Boulenger (1882) described a new species, Bufo benguellensis , based on one individual from “Benguela” sent to him by Bocage as well as two other specimens from “ Fernando Po ” and “W. Africa”; Bocage (1895a) later synonymized B. benguellensis with B. funereus . The species Sclerophrys buchneri (Peters, 1882) is also sometimes treated as conspecific (Tandy and Keith 1972). Schmidt and Inger (1959) provided a map with the distribution of Bufo funereus in Central Africa, although they did not provide information about specific localities.

Sclerophrys garmani ( Meek, 1897) GARMAN’S TOAD

Bufo garmani Meek 1897:176 . Syntypes: FMNH 415, MCZ 19082 [2 specimens] (collector Field Museum

East African Expedition ). Type locality: Haili [= Haileh], southeast of Berbera, Somalia. Bufo regularis humbensis Monard 1937a:26 . Syntypes: MNHC 90.00098-99 [2 specimens] (collector

A. Monard) and MNHB 23712. Type locality: “Mulondo” ( Monard 1937a:26), Angola. Bufo regularis humbensis : Monard (1938:78), Barbour and Loveridge (1946:84), Frade (1963:254). Bufo garmani : Tandy and Keith (1972:142, 159), Frost (1985:46), Largen (2001:315). Bufo regularis sensu stricto: Ruas (1996:21). Bufo gutturalis : Ruas (2002:141). Amietophrynus garmani : Frétey et al. (2011:23). Sclerophrys garmani : Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: Sclerophrys garmani is known from disjunct populations in East Africa from Ethiopia and northern Somalia, Angola, south to northeastern parts of South Africa, and the eastern Caprivi Strip of Namibia.

Ocurrences in Angola (Map 15): The species occurs in southern and eastern Angola.

Moxico: “ Rio Calombe ( Reserva da Palanca

Negra)” [-11.83333, 19.93333] ( Ruas

2002:141). As Sclerophrys garmani humbensis

(plotted as triangles): Huíla: “Mulondo”

[-15.63333, 15.20000] ( Monard 1937a:26,

1938:78; Barbour and Loveridge 1946:84;

Ruas 1996:21). Cunene: “Mupa” [-16.18333,

15.75000] ( Monard 1937a:26, 1938:78; Ruas

1996:21); “Humbi (Humbe)” [-16.68333,

14.90000] ( Monard 1937a:26, 1938:78; Ruas

1996:21). Undetermined Locality: “western subregion and Angolan highlands ( Angola coastal or watersheds)” ( Frade 1963:254).

Taxonomic and distributional notes:

Monard (1937a) originally described a new subspecies of Bufo regularis Reuss, 1833 based MAP 15. Distribution of Sclerophrys garmani in Angola. on specimens from “Molundo,” giving it the name Bufo regularis humbensis Monard, 1937 . These specimens were deposited in the Musée d’histoire naturelle de La Chaux-de-Fonds, Switzerland, though Barbour and Loveridge (1946) also reported one cotype (MCZ A-23712) from “Molundo.” Ruas (1996) identified B. r. humbensis as Bufo regularis sensu stricto, however she noted that some records identified as B. regularis may instead correspond to other Bufo (= Sclerophrys ) species (following Tandy and Keith, 1972). Ruas (1996) recognized S. humbensis as a synonym of S. garmani rather than S. regularis . If S. garmani is restricted to northeastern Africa ( Largen 2001), then Angolan specimens would either represent a relatively isolated population located near the Namibian border (Tandy and Keith 1972), correspond to another species such as S. poweri , or represent an undescribed species. Channing (2001) did not include A. garmani among the species of Angola, in contrast to both Frétey et al. (2011) and Frost (2016).

Sclerophrys gutturalis ( Power, 1927) GUTTURAL TOAD

Bufo spinosus Bocage 1867a:845 . Holotype: MBL 324 (collector J.A. d’Anchieta), not located by Perret

(1976a:18), destroyed by fire 18 March 1978. Type locality: “Benguella,” later corrected to “Dombe,

Benguella, Angola,” by Perret (1976a:18). Preoccupied by Bufo spinosus Daudin, 1803 . Bufo regularis gutturalis Power 1927:416 , Pl. 21, fig. 2. Syntypes: MCZ A-15403 and MMK [originally

4 specimens but now presumably 3] (collector J.H. Power). Type locality: “Lobatsi” and “Kuruman”

Botswana-Northern Cape Province border, South Africa. Bufo spinosus : Bocage (1867b:227), Loveridge (1936a:82). Bufo regularis : Boulenger (1882:299), Bocage (1895a:185), Barbour (1911:135). Bufo regularis gutturalis : Power (1927:416). Bufo regularis regularis : Loveridge (1957:310). Bufo gutturalis : Tandy and Keith (1972:159), Perret (1976a:18), Poynton and Broadley (1988:452), Poynton and Haacke (1993:13), Ruas (1996:21, 2002:141), Channing (2001:74). Amietophrynus gutturalis : Frétey et al. (2011:23), Channing et al. (2012:130). Sclerophrys gutturalis : Frost (2016), Conradie et al. (2016:12).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from southeastern Uganda eastward to coastal Kenya and Tanzania, south through Mozambique to KwaZulu-Natal. Also present in Swaziland, Botswana, northern Namibia, and Angola.

Ocurrences in Angola (Map 16): The species is distributed across the middle of

Angola, from the coast to the country’s eastern border with Zambia and the Democratic

Republic of Congo. Kwanza Norte: “Dondo,

bank of Quanza River ” [-9.68333, 14.43333]

(Poynton and Haacke 1993:13; Ruas 1996:21,

Ruas 2002:142). Malanje: “ Reserva da Palanca Preta (Rio Caluando )” [-11.46667,

17.70000] ( Ruas 1996:21, 2002:141). Moxico:

“ Lago Dilolo ” [-11.50000, 22.01667] ( Ruas

1996:21, 2002:141); “Luso” [-11.78333,

19.91667] ( Ruas 2002:141); “Reserva da

Palanca Preta (Rio Calombe )” [-11.83333,

19.93333] ( Ruas 1996:21, 2002:141);

“Luvuei” [-13.06667, 21.16667] ( Ruas

1996:21, 2002:141). Benguela: “Benguella

(Benguela)” [-12.58333, 13.41667] ( Bocage MAP 16. Distribution of Sclerophrys gutturalis in Angola.

1867a:845; Loveridge 1936a:82, 1957:310;

Ruas 1996:21, 2002:142); “Dombe” [-12.95000, 13.10000] ( Bocage 1895a:185; Perret 1976a:18). Huíla: “Sá da Bandeira” [-14.91667, 13.50000] ( Ruas 1996:21, 2002:141); “Humpata” [-15.03333, 13.40000] (Poynton and Haacke 1993:13; Ruas 1996:21, 2002:142). Namibe: “Mossamedes” [-15.20000, 12.15000] ( Bocage 1867b:277); “Cima/Saco do Giraul” [-15.06667, 12.15000] (Poynton and Haacke 1993:13; Ruas 1996:21; 2002:142); “ Rio Coroca” [-15.78333, 12.06667] (Poynton and Haacke 1993:13; Ruas 1996:21, 2002:142). Cuando Cubango: “Cubango basin (2)” [-15.70452, 17.45897] (Conradie et al. 2017:8-9, 12); “Cubango basin (29)” [-17.87291, 19.83333] (Conradie et al. 2017:9-10, 12); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2017:9-10, 12); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2017:9- 10, 12). “Cuito basin (32)” [-17.04880, 19.05333] (Conradie et al. 2017:9-10, 12); “Cuando basin (44)” (Conradie et al. 2017:12); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2017:9-10, 12).

Taxonomic and distributional notes: The nomen Bufo spinosus Bocage, 1867 was preoccupied by the Iberian Spiny-toad (Daudin 1803) (Loveridge 1936). Prior to this recognition, it had been considered a synonym of Bufo regularis Reuss, 1833 by Boulenger (1882) and Bocage (1895a). Power (1927) described this taxon as the subspecies Bufo regularis gutturalis based on material from the border of Botswana and the Northen Cape. This taxon was later elevated to full species by Tandy and Keith (1972), and is currently uniformly accepted as a valid species ( Channing 2001; du Preez and Carruthers 2009; Frost 2016; Conradie et al. 2016).

Sclerophrys lemairii ( Boulenger, 1901) LEMAIRE’S TOAD

Bufo lemairii : Boulenger 1901:1, Pl. 1, fig. 1. Holotype: not stated, but presumably originally in MRAC (collector C. Lemaire). Type locality: “Pweto, sur le lac Moero” [= Pweto, Moero Lake ], Democratic Republic of Congo.

Bufo lemairei : Laurent (1950a:13, 1964a:131), Cei (1977:16).

Bufo lemairii : Poynton and Broadley (1988:464), Frost (1985:51), Haacke (1982a:11), Ruas (1996:21), Channing (2001:82).

Amietophrynus lemairii : du Preez and Carruthers (2009:144), Frétey et al. (2011:23).

Sclerophrys lemairii : Frost (2016), Conradie et al. (2016:12) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from the Okavango Delta of northern Botswana, western Zambia and adjacent areas of eastern and northeastern Angola. It also occurs in the Caprivi Strip of Namibia. The Democratic Republic of Congo and Republic of Congo appear to be the northern limits of the species.

Ocurrences in Angola (Map 17): The species occurs in northeastern Angola. Lunda

Norte: “Muita, Luembe E (Muíta)”

[-7.80000, 21.45000] ( Laurent 1950a:13; Ruas

1996:21). Lunda Sul: “Alto Chicapa, humidiherbosa des sources du Tchimboma affl. Rive gauche du Cuango-Muqué, Lunda (Rio Chimboma)” [-10.76667, 19.20000] ( Laurent

1964a:131; Ruas 1996:21); “Alto Chicapa,

sources du Cuílo, patie sans forêt riveraine,

Lunda (Rio Cuílo )” [-10.88333, 19.23333]

( Laurent 1964a:131; Ruas 1996:21). Moxico:

“Rive du lac Calundo (Lago Calundo)”

[-11.80000, 20.86667] ( Laurent 1964a:131;

Ruas 1996:21); “Cazombo” [-11.88333,

22.91667] ( Laurent 1964a:131; Haacke

1982a:11; Ruas 1996:21). Cuando Cubango:

MAP 17. Distribution of Sclerophrys lemairii in Angola. “Cubango basin (7a)” [-14.42966, 17.82658]

( Conradie et al. 2016:8-9, 12); “ Cubango basin (7b)” [-14.43377, 17.82957] ( Conradie et al. 2016:8-9, 12); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9-10, 12); “Cuito basin (30d)” [-17.51327, 20.06111] ( Conradie et al. 2016:9-10, 12); “Cuito basin (31)” [-17.46777, 20.03333] ( Conradie et al. 2016:9-10, 12); “Cuíto basin (63)” [-17.93611, 21.10269] ( Conradie et al. 2016:9, 12). Undetermined Locality: “areas of forest and savanna in north and northeast Angola ” ( Cei 1977:169).

Taxonomic and distributional notes: None.

Sclerophrys poweri ( Hewitt, 1935) POWER’S TOAD

Bufo regularis poweri Hewitt 1935:293 . Syntypes: AMG? [5 specimens] (collector J.H. Power), probably lost,

fide Conradie et al. (2015:66). Type locality: “Kimberley,” Northern Cape Province, Republic of South

Africa, fide Conradie et al. (2015:66). Bufo garmani : Tandy and Keith (1972:159). Bufo poweri : Channing (2001:91), Pickersgill (2007a:521). Amietophrynus poweri : Frétey et al. (2011:23), Channing et al. (2012:135), Channing et al. (2013a). Sclerophrys poweri : Frost (2016), Conradie et al. (2016:14).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is found in arid southwestern Africa, from extreme southern

Angola, through northwestern southern Africa, including Namibia, Botswana and western Zimbabwe. Ocurrences in Angola (Map 18): The species occurs in the extreme southern Angola. Cuando Cubango: “Angola side of the Cubango River near Calai (29)” [-17.87291, 19.83333] ( Conradie et al. 2016:9-10, 14). Taxonomic and distributional notes: Tandy and Keith (1972) considered this species to be a member of Bufo regularis group and a synonym of what is now recognized as Sclerophrys garmani ( Meek, 1897) . Due to its similarities with S. garmani, Poynton and Broadley (1988) suggested that poweri might be conspecific with Bufo pseudogarmani Hulselmans, 1969 , although they chose to retain it as a synonym of garmani . Pickergill (2007a) considered S. pseudogarmani a synonym of S. poweri and suggested that southern

MAP 18. Distribution of Sclerophrys poweri in Angola. African populations should be referred to poweri . According to M. Largen, J. Poynton, L. Mazibuko and M. Tandy (pers. comm. in Channing et al. 2013a), the eastern limit of the distribution of S. poweri is uncertain due to remaining confusion with S. garmani . Whereas Channing (2001) did not consider S. garmani to occur in Angola, he did recognize populations from southernmost Angola as S. poweri . Recently, Conradie et al. (2016) cited a single specimen collected in Cuando Cubango Province, which represents the first record for Angola, although it is expected to be more widely distributed in southern and eastern Angola.

Sclerophrys pusilla (Mertens, 1937) MERTENS’ STRIPED TOAD

Bufo cinereus Hallowell 1844:169 . Name preoccupied by Bufo cinereus Schneider, 1799 = syn. Bufo bufo ( Linnaeus, 1758) .

Bufo maculatus Hallowell 1854:101 . Neotype: BMNH 1984.163 fide Poynton et al. (2016:83). Replacement name for Bufo cinereus Hallowell, 1844 . Type locality: “Monrovia, Liberia ” fide Poynton et al. (2016:83).

Bufo regularis pusillus Mertens 1937:17 . Holotype: SMF 22247 (collected by F. Haas). Type locality: “Letaba Camp,” Kruger National Park , South Africa fide Poynton et al. (2016:84).

Bufo maculatus : Tandy and Keith (1972:159), Poynton and Broadley (1988:460), Channing (1989:1, 2001:84), Poynton and Haacke (1993:13), Pickersgill (2007a:541).

Bufo funereus : Monard (1937a:26, 1938:55, 77, 78).

Bufo pusilus : Cei (1977:17), Lambiris (1988:51).

Bufo maculatus : Ruas (1996:21, 2002:141).

Bufo funereus : Ruas (1996:21, 2002:142).

Amietophrynus maculatus : Frétey et al. (2011:23),

Sclerophrys maculata : Frost (2016), Ceríaco et al. (2016a:19).

Sclerophrys pusilla : Poynton et al. (2016:84), Conradie et al. (2016:14) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from eastern, central and southern Africa from southern and western Cameroon, eastern through the Central African Republic and Sudan south to Mozambique and northern regions of South Africa and Swaziland. The species also occurs in western Zimbabwe, Zambia, and northern Namibia, with a possible wide range in Angola.

Ocurrences in Angola (Map 19): The species occurs in western Angola. Malanje:

“Duque de Bragança falls (Duque de Bragança)” [-9.13333, 16.06667] (Poynton and Haacke 1993:13; Ruas 1996:21, 2002:141). Bié:

“ Caccuchi River ” [-13.593333, 16.879861]

( Poynton et al. 2016:89). Huambo: “Santo

Amaro” [-12.70000, 15.85000] ( Monard

1937a:26, 1938:77); “ Huambo ” [-12.737167,

15.81825] ( Poynton et al. 2016:89); “Cubango basin (12a)” [-13.59333, 16.87986] ( Conradie et al. 2016:14). Benguela: “ 31 km NE of Sousa

Lara-Chila (Bocoio)” [-12.46667, 14.13333]

(Poynton and Haacke 1993:13; Ruas 1996:21);

“Ebanga” [-12.73333, 14.73333] ( Monard

1937a:26, 1938:78); “Monguavalo Farm”

[-13.45000, 14.61667] (Poynton and Haacke

MAP 19. Distribution of Sclerophrys pusilla in Angola. 1993:13; Ruas 1996:21). Huíla: “Kalukembé

(Caluquembe)” [-13.78333, 14.68333] ( Monard 1938:55, 76; Ruas 1996:21, 2002:142); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937a:26, 1938:77). Namibe: “Leba pass, between river and highway” [-15.07033, 13.24381] (Ceríaco et al. 2016a:19); “Cainde” [-15.48333, 13.36667] (Poynton and Haacke 1993:13; Ruas 1996:21; Ceríaco et al. 2016a:19); “ 16 km W of Vila Nova” (Poynton and Haacke 1993:13; Ruas 1996:21; Ceríaco et al. 2016a:19). Cuando Cubango: “Cubango basin (3)” [-14.94277, 17.71863] ( Conradie et al. 2016:14); “Cubango basin (6a)” [-14.67155, 17.73525] ( Conradie et al. 2016:14); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:14); “Cubango basin (20)” [-14.67175, 17.15331] ( Conradie et al. 2016:14); “Cubango basin (22d)” [-14.64991, 16.90739] ( Conradie et al. 2016:14); “Cuito basin (24)” [-14.60622, 18.46722] ( Conradie et al. 2016:14); “Cubango basin (29)” [-17.87291, 19.83333] ( Conradie et al. 2016:14); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:14); “Cuito basin (32)” [-17.04880, 19.53333] ( Conradie et al. 2016:14); “Cuando basin (41)” ( Conradie et al. 2016:14); “Cubango basin (47)” [-14.70214, 17.37806] ( Conradie et al. 2016:14); “Cuito basin (62)” [-17.50875, 20.06608] ( Conradie et al. 2016:14).

Taxonomic and distributional notes: Tandy and Keith (1972) placed Bufo pusillus (sensu Pienaar, 1963) in the synonymy of Sclerophrys maculata (Hallowell, 1854) due the similar morphology and mating calls. While Hulselmans (1969) questioned this synonymy, immunological evidence ( Maxson 1981; Pickergill 2007a) suggested that it is not distinct from S. maculata . A recent study conducted by Poynton et al. (2016) established that S. maculata is now restricted to West Africa from the Gulf of Guinea eastwards to Cameroon, while S. pusilla is found in eastern and southern Africa, including Angola. Conradie et al. (2016) collected some new material from Cuando Cubango Province identified as S. pusilla . Monard (1937 a, 1938) identified specimens from “Kalukembé” as Bufo funereus (Bocage) [= Sclerophrys funera (Bocage, 1866) ] but we recently determined these to be S. maculata (Ceríaco et al. 2016b) , which now should be refered to S. pusilla .

Sclerophrys regularis ( Reuss, 1833) AFRICAN COMMON TOAD

Bufo regularis Reuss 1833:60 . Syntypes: SMF 3429, formerly 1298.1c and SMF 1298.d [2 specimens] (collector probably E. Rüppell), the specimens represented by two figures of Grenouille Ponctuée from SaintHilaire 1809: p. 4, fig. 1, 2 are also syntypes. Type locality: “Aegypten” [= Egypt]. Bufo guineensis : Peters (1877a:618), Bocage (1879b:89). Bufo pantherinus : Bocage (1866a:56). Bufo regularis : Boulenger (1882:298, 1905:107), Bocage (1887a:192, 1887b:208, 1895a:185, 1896a:113),

Ferreira (1903:114), Monard (1937a:26, 1938:55, 77), Parker (1939:145), Themido (1941:2), Laurent

(1954a:70), Inger (1959:540), Cei (1977:16, 17), Largen (2001:322). Bufo regularis regularis : Schmidt (1936:128), Loveridge (1936a:81), Monard (1938:78), Mertens (1937a:17,

1938a:429), Laurent (1950a:13), Hellmich (1957a:23), Laurent (1964a:130). Bufo regularis sensu lato: Ruas (1996:21). Amietophrynus regularis : Frétey et al. (2011:23), Channing et al. (2012:137), Ceríaco et al. (2014b:669),

IUCN SSC Amphibian Specialist Group (2016), Ceríaco et al. (2016b:27). Sclerophrys regularis : Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: This species has among the broadest distributions of all African amphibian species. It is known from northern Africa, western Eritrea and Ethiopia (also included Uganda and southern Kenya), southwards along the western coast from Cameroon to the Demoratic Republic of Congo to Angola.

Ocurrences in Angola (Map 20): The species is widespread in Angola, except in the southeast . Cabinda: “ Chinchoxo ” [-5.10000,

12.10000] (Peters 1877:618). Zaire: “ S. Salvador do Congo ” [-6.26667, 14.23333] ( Bocage

1887a:192; Ruas 1996:21). Lunda Norte:

“Dundo” [-7.36667, 20.83333] ( Laurent

1950a:13, 1954a:70, 1964a:130; Ruas

1996:21); “Muita, Luembe E (Muíta)”

[-7.80000, 21.45000] ( Laurent 1950a:13,

1954a:70; Ruas 1996:21). Lunda Sul: “Poste de Luangue, humidiherbosa du ruisseau

Katcheleke affl. W. du Luangue, entre le Lunguena et la Tchá-Pemba, Lunda (Posto do

Luangue)” [-9.33333, 19.23333] ( Laurent

1964a:130; Ruas 1996:21); “Riv. Cuvuemba

Alto Cuílo, Lunda (Rio Cavemba)”

[-10.01667, 19.55000] ( Laurent 1964a:130; MAP 20. Distribution of Sclerophrys regularis in Angola. Ruas 1996:21); “Lunda” [-10.96667, 20.06667] ( Monard 1938:78); “Alto Chicapa, humidiherbosa des sources du Tchimboma affl. Rive gauche du Cuango-Muqué, Lunda (Rio Chimboma)” [-10.76667, 19.20000] ( Laurent 1964a:130; Ruas 1996:21); “Dala” [-11.03333, 20.20000] ( Monard 1937a:26, 1938:77; Ruas 1996:21). Kwanza Norte: “N’dalla Tando” [-9.30000, 14.91667] (Ferrreira 1904:113; Monard 1938:55; Ruas 1996:21); “Cazengo” [-9.33333, 14.76667] (Ferrreira 1904:113; Monard 1938:55; Ruas 1996:21); Kwanza Sul: “Congulu (Congulo)” [-10.86667, 14.28333] ( Parker 1936:145; Ruas 1996:21); “Novo Redondo” [-11.20000, 13.85000] (Ferrreira 1904:113; Monard 1938:55; Ruas 1996:21); “Furna s/riv. N’Gunza, Novo Redondo (N’Gunza)” [-11.20000, 13.93333] ( Laurent 1954a:70; Ruas 1996:21). Malanje: “Duque de Bragança” [-9.10000, 15.95000] ( Boulenger 1882:298, 1905:107; Bocage 1866a:56; 1895a:185;

Ruas 1996:21); “Pungo-Andongo (Pungo Andongo)” [-9.66667, 15.58333] ( Bocage 1895a:185; Boulenger 1905:107; Ruas 1996:21); “Capanda” [-9.72841, 15.34585] (Ceríaco et al. 2014b:669); “Marimba” [-8.36667, 17.03333] ( Boulenger 1905:107; Ruas 1996:21) “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:27). Moxico: “Teixeira de Sousa, Distr. Villa Luso” [-10.70000, 22.23333] ( Mertens 1937a:17); “environs du lac Calundo, rives de la Lumeje, Moxico (Rio Lumege)” [-11.51667, 20.76667] ( Laurent 1964a:130; Ruas 1996:21); “Rives du lac Calundo Moxico” [-11.80000, 20.86667] ( Laurent 1964a:130; Ruas 1996:21); “Reserve de Chasse de Cameia, 120 km à l’est de Luso (Cameia)” [-11.83333, 21.00000] ( Laurent 1964a:130; Ruas 1996:21); “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:130; Ruas 1996:21). Bié: “Bihé (Bié)” [-12.38333, 16.95000] ( Bocage 1879b:78, 1895a:185; Ruas 1996:21); “Gauca” [-11.18333, 17.45000] ( Schmidt 1936:128; Monard 1938:55; Ruas 1996:21); “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:128; Monard 1938:55; Ruas 1996:21); “General Machado” [-12.03333, 17.46667] ( Mertens 1937a:17). Huambo: “Bimbi (Bimbe)” [-11.81667, 15.83333] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Cuito, Mt. Moco ” [-12.45000, 15.26667] ( Parker 1936:145; Ruas 1996:21); “Santo Amaro” [-12.70000, 15.85000] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Cuma” [-12.86667, 15.06667] ( Loveridge 1936a:81). Benguela: “Benguella (Benguela)” [-12.58333, 13.41667] ( Bocage 1895a:185; Ruas 1996:21); “Dombe” [-12.95000, 13.1000] ( Bocage 1895a:185); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:23); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:426; Ruas 1996:21); “Cubal da Ganda, Marco de Canavezes (Marco de Canavezes)” [-13.08333, 14.33333] ( Laurent 1964a:130; Ruas 1996:21); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:185); “Kalukembé (Caluquembé)” [-13.78333, 14.68333] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Sangevé (Sangueve)” [-13.88333, 15.83333] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Quilengues” [-14.10000, 14.08333] ( Laurent 1954a:70; Ruas 1996:21); “Kuvangu (Cubango)” [-14.46667, 16.30000] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Kapelongo (Capelongo)” [-14.88333, 15.08333] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Osi (Osse)” [-15.08333, 15.41667] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Kangela (Cangela)” [-15.41667, 15.73333] ( Monard 1937a:26, 1938:77; Ruas 1996:21). Nambie: “Mossamedes” [-15.20000, 12.15000] ( Bocage 1895a:185; Ruas 1996:21). Cunene: “Kuvelaï (Cuvelai)” [-15.65000, 15.80000] ( Monard 1937a:26, 1938:77; Ruas 1996:21); “Mupa” [-16.18333, 15.75000] ( Monard 1937a:26, 1938:77; Ruas 1996:21). Cuando Cubango: “Kakindo (Caquindo)” [-15.45000, 17.05000] ( Monard 1937a:26, 1938:77; Ruas 1996:21). Undetermined Locality: “from almost all localities that he [J. d’Anchieta] visited” ( Bocage 1895a:185); “Locomi” ( Boulenger 1905:107; Ruas 1996:21); without precise locality (Laurent 1954:70); “Carangigo” ( Boulenger 1882:298); “Between Benguela and Bihé” ( Boulenger 1905:107). “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “arid territories along the coast” ( Cei 1977:17).

Taxonomic and distributional notes: Sclerophrys regularis ( Reuss, 1833) remains a taxonomic challenge. While many specimens from Angola have been identified as S. regularis, Ruas (1996) noted that the Angolan records are not representative of Bufo regularis sensu stricto and may represent Bufo maculatus (Hallowell, 1854) (currently Sclerophrys pusilla (Mertens, 1937) , see S. pusilla account) and/or Bufo guturallis ( Power, 1927) . Sclerophrys regularis is a widely distributed in Africa ((IUCN SSC Amphibian Specialist Group 2016), but the boundaries between S. regularis and S. gutturalis in Angola, the Democratic Republic of Congo, Kenya, Uganda, and Tanzania remain poorly understood. Further studies are needed to clarify the distribution and the known range should be regarded as provisional. Ruas (1996) cited several localities (e.g., “Huíla,” “Bibala,” “Quibula,” “Quindumbo,” “Cahata,” “Dondo,” “Ambaca,” “Catumbela,” “Galanga,” “Humbe,” “Rio Quando,” “Gambos,” “Capagombe,” and “Massabi”) referring to Bocage (1895a) as a source, although these localities are not cited in Bocage’s work.