Xenopus sp.

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 : 60-65

Xenopus sp.

Xenopus fraseri : Laurent (1950a:13, 1954a:70), Cei (1977:16), Kobel (1981:120), Ruas (1996:20), Channing (2001:240), Channing et al. (2012:294), Wagner et al. (2013:206), Ernst et al. (2015:147).

Xenopus (Xenopus) fraseri : Frétey et al. (2011:22).

Global distribution: The taxon is presently known from northeastern Angola, although it likely represents a species that is more widespread and ranging from northern Angola to Cameroon and Central African Republic.

Ocurrences in Angola (Map 3): Occurs in the extreme northeast of the country. Lunda

Norte: “Muita, Luembe E (Muita, Luachimo)”

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

1996:20); “Dundo” [-7.36667, 20.83333] ( Laurent 1954a:70; Ruas 1996:20); “ Lunda Norte

(probably Dundo or Muita)” ( Kobel 1981:120).

Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ”

( Cei 1977:16).

Taxonomy and natural history notes:

The Xenopus amieti subgroup (sensu Evans et al., 2015) contains several morphologically similar species that mainly occur in Central

Africa ( Wagner et al. 2013; Ernst et al. 2015).

Previously this group contained X. fraseri , but

Evans et al. (2015) demonstrated that Central

African species referred to this taxon instead MAP 3. Distribution of Xenopus sp. in Angola. represent either X. allofraseri or X. parafraseri . The identity of the Angolan populations of Xenopus sp. remains uncertain, but they most likely represent a species within the amieti subgroup. Only two species, X. pygmaeus and X. fraseri , have previously been reported from south of the Congo Basin ( Laurent 1950a, 1954a; Ruas 1996; Wagner et al. 2013). The species X. fraseri has frequently been confused with X. andrei , a closely related species previously believed to be restrict- ed to Cameroon, Gabon, and the Central African Republic, but which was recently found in Angola ( Ernst et al. 2015).

Laurent (1950, 1954a) reported many specimens (~133) from Angola, collected in Lunda Norte Province, in “Dundo” and “Muita” regions in northeastern Angola, an area bordering the Democratic Republic of Congo. Ruas (1996) referred these previously published records from the northeastern Angola to X. fraseri although she also provided a complementary map ( Ruas 1996) with other records for X. fraseri in central and southern Angola, with no further detail, on the basis of these records. It is likely that all previous records of X. fraseri in Angola actually refer to other species, including X. andrei or recently described species ( Evans et al. 2015). Because of the lack of corresponding vouchers and tissue samples for genetic analyses, it cannot at this time be determined to which species these records might be best assigned.

Xenopus muelleri ( Peters, 1844) MÜLLER’S CLAWED FROG

Dactylethra Muelleri Peters 1844:37 . Syntypes: ZMB 3556 [3 tadpoles], 3557 [3 specimens], 6164 [2 specimens] and MZUT An264 fide Bauer et al. (1995:48) (collector W.C.H. Peters). Type locality: “ Mozambique ” ( Peters 1844:37), restricted to “ Tete, Zambezi River” [ Mozambique] by Loveridege (1953:308).

Xenopus muelleri : Conradie et al. (2016:17) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known to occur along the East African coastal belt from extreme southern Kenya through Tanzania to the northwestern border of South Africa. There is an apparently allopatric population from Burkina Faso eastward across Sudan-Guinea to northeastern Democratic Republic of Congo and Angola.

Ocurrences in Angola ( Map 4 ): Occurs in the eastern regions of Angola. Cuando Cubango: “Cubango basin (29)” [-17.8729, 19.83333] ( Conradie et al. 2016:17); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9 -10,17).

Taxonomic and distributional notes: Given the known distribution for X. muelleri and the fresh material collected by Conradie et al. (2016), it is expected that the species is more widespread in the eastern regions of Angola, and probably occurs in sympatry with X. poweri .

Xenopus petersii Bocage, 1895 PETERS’ CLAWED FROG

X enopus Petersii Bocage 1895a:187 . Syntypes: MBL 3.476 (S. Salvador do Congo, collector António Barroso), 3.473 (Quimbundo), 3.474-477 (Caconda), 3.478 (Dombe), 3.479 (Quibula), 3.481-483 (Huíla), 3.485 (Cassange), 3.486-488 (Dondo) (collector J.A. d’Anchieta) [15 specimens] fide Perret (1976a:16). All destroyed by fire 18 March 1978. Bauer et al. (1996:271) reported one syntype from “Catumbella” ZMB 5831, as a surviving specimen from exchanged from the MBL. Type locality: “S. Salvador do Congo,” “Dondo,” “Caconda,” “Dombe,” “Quibula,” “Huíla,” “Cassange” and “Quindumbo” ( Bocage 1895a:187), [= M’Banza Congo, Dondo, Caconda, Dombe, Huíla, Cassange and Quindumbo], Angola.

Dactylethra laevis : Günther (1865a:480).

Dactylethra mülleri : Bocage (1867b:227).

Dactylethra Multeri : Bocage (1879b:89, 1879c:96).

Xenopus muelleri : Boulenger (1882:457), Loveridge (1957:308).

Xenopus petersii : Bocage (1896a:113, 1897a:206), Ferreira (1906:166), Schmidt (1936:128), Bauer et al. (1996:271), Channing (2001:248), Measey and Channing (2003:325), Furman et al. (2015:910), Frost (2016), Ceríaco et al. (2016b:41).

Xenopus laevis : Boulenger (1905:107), Monard (1937a:25, 1938:55, 76), Hellmich (1957a:22), Inger (1959:540), Kobel (1981:120), Gavetti and Andreone (1993:41).

Xenopus laevis poweri : Laurent (1964a:129), Cei (1977:17).

Xenopus laevis petersi : Loveridge (1957:308), Schmidt and Inger (1959:8), Laurent (1964a:130), Perret (1976a:17), Cei (1977:17), Poynton and Broadley (1985a:508), Poynton and Haacke (1993:13), Ruas (1996:20, 2002:141).

Xenopus (Xenopus) laevis : Frétey et al. (2011:22).

Xenopus cf. petersii : Ceríaco et al. (2014b:669).

Xenopus petersii : Furman et al. (2015:910), Frost (2016), Conradie et al. (2016:17) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from southern Gabon, southwards to the western Democratic Republic of Congo, Angola, and northern Namibia.

Ocurrences in Angola (Map 5): The species likely is widely distributed across western and central Angola. Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] ( Bocage 1895a:187, 1897a:206; Loveridge 1957:308; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20). Kwanza

Norte: “Cambondo” [-9.15963, 14.65771]

( Ferreira 1906:166; Monard 1938:55); “ Rio

Luinha” [-9.26667, 14.53333] ( Ferreira

1906:166); “Ambaca” [-9.26667, 15.18333]

( Bocage 1895a:187, 1897a:206; Monard

1938:55; Loveridge 1957:308; Bauer et al.

1996:271; Ruas 1996:20, 2002:141); “Dondo”

[-9.68333, 14.43333] ( Bocage 1895a:187,

1897a:206; Monard 1938:55; Loveridge

1957:308; Perret 1976a:17; Loumont

1984:728; Bauer et al. 1996:271; Ruas

1996:20). Malanje: “Duque de Bragança ”

[-9.13333, 16.06667] (Schmidt and Inger

1959:8; Loumont 1984:728; Ruas 1996:20,

2002:141); “Capanda” [-9.72841, 15.34585]

(Ceríaco et al. 2014b:669); “Reserva da Palanca Preta (margens do Rio Cuanza)”

[- 11.11667, 17.46667] ( Ruas 2002:141), MAP 5. Distribution of Xenopus petersii in Angola. “Pungo Andongo” [-9.66667, 15.58333] ( Boulenger 1905:107; Monard 1938:55; Ruas 1996:20); “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:41). Kwanza Sul: “Cassongue” [-11.85000, 15.05000] ( Loumont 1984:728; Ruas 1996:20). Lunda Norte: “Cassange” [-9.58333, 17.86667] ( Bocage 1895a:187, 1897a:206; Monard 1938:55; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20, 2002:141). Lunda Sul: “mare Tchifuka, Alto Cuílo, Lunda (= Chifuca)” [-10.00000, 19.58333] ( Laurent 1964a:130; Loumont 1984:728; Ruas 1996:20, 2002:141); “Alto Chicapa, Lunda” [-10.88333, 19.23333] ( Laurent 1964a:130; Ruas 1996:20, 2002:141); “sources du Cuílo, Alto Chicapa, rives sans forêt, Lunda (= Rio Cuílo)” [-10.86667, 19.40000] ( Laurent 1964a:130; Loumont 1984:728; Ruas 1996:20, 2002:141); “Dala” [-11.03333, 20.20000] ( Monard 1937a:25, 1938:76; Ruas 1996:20, 2002:141); “mare des rives du Kutele, affl. droit du Cuango, Alto Chicapa, Lunda (= Rio Cutele)” [-11.06667, 18.86667] ( Laurent 1964a:130; Ruas 1996:20, 2002:141). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1936:128; Monard 1938:55; Schmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:128; Monard 1938:55; Schmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Nequilo” [-12.58500, 17.07000] ( Furman et al. 2015 - see Table S1); “Cubango basin (10)” [-13.71616, 17.09661] ( Conradie et al. 2016:8-9, 17). Benguela: “Quibula” [-12.28333, 14.68333] ( Bocage 1895a:187; Monard 1938:55; Loveridge 1957:308; Perret 1976a:17; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Catumbella (= Catumbela)” [-12.43333, 13.55000] ( Bocage 1895a:187; Monard 1938:55; Loveridge 1957:308; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:187; Monard 1938:55; Loveridge 1957:308; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Benguella (= Benguela)” [-12.58333, 13.41667] ( Bocage 1895a:187, 1897a:206; Boulenger 1882:457, 1905:107; Monard 1938:55; Loveridge 1957:308; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Dombe” [-12.95000, 13.10000] ( Bocage 1879b:89; 1895a:187, 1897a:206; Monard 1938:55; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Caimbambo” [-13.01667, 14.01667] (Loumont 1894:728; Ruas 2002:141); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:22); “Catengue” [-13.03333, 13.73333] (Schmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Cubal da Ganda (Marco de Canavezes)” [-13.08333, 14.33333] ( Laurent 1964a:129; Ruas 1996:20, 2002:141);

“Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113; Monard 1938:55; Ruas 1996:20, 2002:141). “Rio Coporolo (= Coporola)” [-13.93333, 12.96667] (Loumont 1894:728; Ruas 2002:141). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:187; Monard 1938:55; Loveridge 1957:308; Perret 1976a:17; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Kalukembé (= Caluquembe)” [-13.78333, 14.68333] ( Monard 1937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Kuvangu (= Cubango)” [-14.46667, 16.30000] ( Monard 1937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Huilla (= Huíla)” [-15.05000, 13.55000] ( Günther 1865a:480; Bocage 1895a:187, 1897a:206; Monard 1938:55; Schmidt and Inger 1959:9; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Osi (= Osse)” [-15.08333, 15.41667] ( Monard 1937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Lagoa Nuntechite” [-15.13333, 13.41667] (Poynton and Haacke 1993:13); “Kampulu (= Campulu-Cambissa)” [-15.21667, 16.11667] ( Monard 1937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Kuluï (= Cului)” [-15.41667, 15.73333] ( Monard 1937a:25, 1938:55, 76; Ruas 1996:20, 2002:141). Namibe: “Môssamedes” [-15.20000, 12.15000] (Bocage 1967b:227; Loumont 1984:728; Ruas 1996:20, 2002:141). Cuando Cubango: “Kandingu” [-14.66667, 17.70000] ( Monard 1937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Cubango basin (3)” [-14.94277, 17.71863] ( Conradie et al. 2016:8-9, 17); “Cubango basin (5)” [-14.74628, 17.66844] ( Conradie et al. 2016:8-9, 17); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8-9, 17); “Cubango basin (21)” [-14.66586, 17.07661] ( Conradie et al. 2016:8-9, 17); “Cuito basin (24)” [-14.60622, 18.46722] ( Conradie et al. 2016:8-9, 17). Undetermined Locality: without precise locality ( Bocage 1879b:95); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:17); “plateaus regions” ( Cei 1977:17); “arid territories along the coast” ( Cei 1977:17).

Taxonomic and distributional notes: See notes below for Xenopus poweri .

Xenopus poweri Hewitt, 1927 POWER’S CLAWED FROG

Xenopus poweri Hewit 1927 a:413, pl. 24. fig. 3. Holotype: MMK/F/898 (collector J.H. Power). Type locality: “ Victoria Falls ,” Zambia and Zimbabwe border.

Xenopus laevis poweri : Schmidt and Inger (1959:8), Laurent (1964a:129).

Xenopus laevis : Kobel (1981:120), Channing (2001:243).

Xenopus laevis petersii : Ruas (1996:20, 2002:141).

Xenopus (Xenopus) laevis : Frétey et al. (2011:22).

Xenopus poweri : Furman et al. (2015:910), Frost (2016), Conradie et al. (2016:17) .

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from central and eastern Nigeria to eastern Cameroon and western Central Africa Republic, south to the Democratic Republic of Congo, eastern and southeastern Angola, the Okavango Region of Namibia, Zambia, and western Zimbabwe to northern Botswana.

Ocurrences in Angola (Map 6): The species occurs in central and western Angola, while X. poweri is more restricted to the east and southeastern regions. Moxico: “Reserva da Palanca Preta (Rio Calombe)” [-11.83333, 19.93333] ( Ruas 1996:20; Ruas 2002:141); “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:130; Ruas 1996:20, 2002:141); “Lucusse” [-12.51667, 20.81667] ( Ruas 1996:20, 2002:141). Cuando Cubango: “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9-10, 17); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9-10, 17).

Taxonomic and distributional notes: The African clawed frog Xenopus laevis (Daudin, 1903) has a large native distribution over much of sub-Saharan Africa, but despite its important role in biological research, its phylogeography and evolutionary history remains poorly known. The historic data for Xenopus laevis in Angola begins with Günther (1865a), who cited the species for “Huíla,” followed by several authors including Boulenger (1905), Monard (1937 a, 1938), and Hellmich (1957a). After Günther’s citation, Bocage (1895a) described a new species, Xenopus petersii , based on numerous specimens deposited in Museu Bocage, Lisboa (see above), all considered syntypes by Perret (1976a). Unfortunately, the syntypes were destroyed in the fire in the Museu Bocage in 1978. Bauer et al. (1996) reported a surviving syntype from “Catumbella” deposited in Museum für Naturkunde, Berlin.

Parker (1936) described four races or groups of X. laevis , reducing X. petersii to a subspecies of X. laevis and suggesting that MAP 6. Distribution of Xenopus poweri in Angola. Xenopus poweri Hewitt, 1927 is a synonym of X. petersii (Loveridge 1957) . When describing X. petersii, Bocage (1895a) listed three “varieties” based on differences in ventral markings. He gave no indication that this variation was geographical, but Schmidt and Inger (1959) reported that 12 FMNH specimens from northern Angola represent Bocage’s “var. A” pattern, whereas five southern Angolan specimens exhibit the “var. B” pattern. They refered X. laevis petersii to “var. A,” which represents the northern form distributed from the Upper Cuanza River northward. They assigned Xenopus laevis poweri to “var. B” reaching from the Lower Cuanza River across all of southern Angola to Zambia (Schmidt and Inger 1959; Poynton and Broadley 1985a). Mertens (1971), however, rejected this view. Channing (2001) considered X. petersii as a full species, removing it from synonymy with Xenopus laevis . Measey and Channing (2003) provided molecular evidence for the distinctiveness of X. petersii from X. laevis and considered X. poweri to be a synonym of X. petersii , although other authors, including Frétey et al. (2011) nonetheless later included X. petersii and X. poweri as synonyms of Xenopus laevis .

Recently, Furman et al. (2015) provided an analysis of molecular variation in the X. laevis group, which clarified the distribution of these lineages and supported the recognition of X. laevis sensu stricto, X. petersii Bocage, 1895 , and X. victorianus Ahl, 1924 and revalidated X. poweri Hewitt, 1927 as a separate species. These authors restricted X. laevis to South Africa and the population from Western Central Africa to X. petersii , which indicates that the central and western Angolan population should be referred to the latter. They also proposed that portions of the currently recognized distributions of X. laevis north of the Congo Basin and X. petersii south of the Congo Basin should be referred to X. poweri . This suggests that populations from eastern and southeastern Angola should be referred to X. poweri ( Furman et al. 2015) .

Based on the available studies and in some new material recently collected by Conradie et al. (2016), we refer records from Moxico Province to X. poweri , including those from “Lucusse,” “Reserva da Palanca Preta,” “ Rio Calombe” ( Ruas 1996, 2002), and “Cazombo” ( Laurent 1964a).

Figures

MAP 4. Distribution of Xenopus muelleri in Angola.