Tomopterna Duméril and Bibron, 1841

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 : 153-156

Genus Tomopterna Duméril and Bibron, 1841

Tomopterna cryptotis (Boulenger, 1907) TREMELO SAND FROG

Rana cryptotis Boulenger 1907b:109 . Syntypes: BMNH 1947.2.1.73–78, 1947.2.28.48–5, and MCZ A-19268, exchanged from BMNH fide Barbour and Loveridge (1946:182) (collector Dr. W.J. Ansorge), formerly BMNH 1907.6.29.82–96. Type locality: “Catequero,” “Ponang Kuma (Dongwenna)” and “Kafitu swamps” ( Boulenger 1907b:109) Namibe Province, Angola.

Rana cryptotis : Barbour and Loveridge (1946:182).

Tomopterna cryptotis : Poynton and Broadley (1985b:125), Poynton and Haacke (1993:13), Frost (1985:528, 2016), Ruas (1996:23), Channing (2001:365), Largen (2001:330), Channing and Howell (2006:329), Pickersgill (2007a:118).

Tomopterna (Tomopterna) cryptotis : Frétey et al. (2011:43).

Tomopterna cryptotis : Conradie et al. (2016:19) .

Global conservation status (IUCN): Least Concern.

Global distribution: This widespread species is found in xeric regions across much of sub-Saharan Africa.

Ocurrences in Angola (Map 100): The species occurs in southwestern Angola. Namibe: “ 25 km W of Virei” [-15.66667, 12.76667] (Poynton and Haacke 1993:13; Ruas 1996:23); “Miranda” ( Boulenger 1907b:109; Poynton and Haacke 1993:13; Ruas 1996:23). Cunene: “ 2 km NW of Calequero (Cahama)” [-16.28333, 14.30000] (Poynton and Haacke 1993:13); “Catequero” [-16.56667, 14.90000] ( Boulenger 1907b:109; Poynton and Broadley 1985b:125; Frost 2016, 1985:528; Ruas 1996:23; Largen 2001:330); “Ponang Kuma (Dongwenna)” [-17.05000, 14.65000] ( Boulenger 1907b:109; Barbour and Loveridge 1946:182; Poynton and Broadley 1985b:125; Frost 2016, 1985:528; Ruas 1996:23); “Kafitu swamps” ( Boulenger 1907b:109; Poynton and Broadley 1985b:125; Frost 2016, 1985:528; Largen 2001:330). Cuando Cubango: “Cubango basin (22c)” [-14.65386, 16.93547] ( Conradie et al. 2016:8 -9,19); “Cubango basin (29)” [-17.87291, 19.83333] ( Conradie et al. 2016:9 -10,19).

Taxonomic and distributional notes:

Originally described based on specimens from

Angola, many subsequent authors have identified populations across much of Africa as

T. cryptotis . Recent molecular genetic analyses suggest at least some populations found far from Angola, including Somalia and Kenya,

represent other distinct species (Zimkus and

Larson 2011; Wasonga and Channing 2013).

Because of the difficulty in identifying some species of Tomopterna based on morphology alone, the extent of the distribution of this species remains uncertain ( Channing 2001;

Channing and Howell 2006). Some records of

T. cryptotis might refer instead to T. tandyi .

Channing (2001) provided a map for T. tandyi with records from southern Angola ( Pickersgill

2007a; Channing et al. 2012), between Namibe MAP 100. Distribution of Tomopterna cryptotis in Angola.

and Cunene provinces.

Tomopterna damarensis Dawood and Channing, 2002 DAMARALAND SAND FROG

Tomopterna damarensis Dawood and Channing 2002:130 , figs 1, 2, 3. Holotype: TM 83913 (collector

A. Channing). Type locality: “Khorixas” (Dawood and Channing 2002:133), Kunene Region, Namibia. Tomopterna damarensis : Frost (2016), Ceríaco et al. (2016a:20) .

Global conservation status (IUCN):

Data Deficient.

Global distribution: The species is known from southwestern Angola and northern

Namibia.

Ocurrences in Angola (Map 101): The species occurs in southwestern Angola .

Namibe: “Pediva Hot springs” [-16.28462,

12.56329] (Ceríaco et al. 2016a:20).

Taxonomic and distributional notes:

Dawood and Channing (2002) suggested that this species might occur more broadly. Recently Ceríaco et al. (2016a) reported the first record for Angola from Iona Natinoal Park.

Previous records of some other species of

Tomopterna , including T. tandyi , might refer to this species (Heinicke et al. 2017).

MAP 101. Distribution of Tomopterna damarensis in Angola.

Tomopterna krugerensis Passmore and Carruthers, 1975 KNOCKING SAND FROG

T omopterna krugerensis Passmore and Carruthers 1975:32 , figs. 4,5,6,7, 8. Holotype: TM 44670 (collectors N.I. Passmore and V.C. Carruthers). Type locality: “Machayipan, Kruger National Park ,” South Africa.

Tomopterna krugerensis : Poynton and Broadley (1985b:127), Channing (2001:368).

Tomopterna (Tomopterna) krugerensis : Frétey et al. (2011:43).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is occurs in savannas extending from Angola and Namibia, east through Botswana into northern South African and southern Mozambique.

Ocurrences in Angola: The species occurs in southwestern Angola. Undetermined Locality: without precise locality ( Channing 2001:369).

Taxonomic and distributional notes: This species is similar morphologically to T. cryptotis but has a distinct call (Poynton and Broadley 1985b) and is phylogenetically distinct (Zimkus and Larson 2011). Because of the morphological similarity, some previous reports from Angola for T. cryptotis might be referable to this species or T. tandyi . Channing (2001) presented the only records in Angola for T. krugerensis with mapped points in Namibe and Cunene provinces, though without specific locality details.

Tomopterna tandyi Channing and Bogart, 1996 TANDY’S SAND FROG

Tomopterna tandyi Channing and Bogart 1996:80 , figs. 2, 3. Holotype: PEM A-2283 (collector M. Snyman). Type locality: “Bedford in the Eastern Cape ” (Channing and Bogart 1996:80), South Africa.

Tomopterna tandyi : Channing (2001:372), Pickersgill (2007a:117), Frétey et al. (2011:43), Channing et al. (2012:360), Frost (2016).

Global conservation status (IUCN): Least Concern.

Global distribution: The species occurs in South Africa, Namibia, and southwestern Angola.

Ocurrences in Angola: The species occurs in southwestern Angola. Undetermined Locality: “without precise locality” ( Channing 2001:369).

Taxonomic and distributional notes: A tetraploid species, likely of hybrid origin between Tomopterna cryptotis (Boulenger, 1907) and Tomopterna delalandii ( Tschudi, 1838) (Channing and Bogart 1996, Frost 2016). This species is morphologically indistinguishable from T. cryptotis and T. delalandii and many records of Tomopterna cryptotis , including in Angola, may refer to T. tandyi . Channing (2001) provided a map for T. tandyi with records from southwestern Angola between Namibe and Cunene provinces (followed by Pickersgill 2007a, Channing et al. 2012, and Frost 2016), though without specific information.

Tomopterna tuberculosa ( Boulenger, 1882) ROUGH SAND FROG

Pyxicephalus rugosus Günther 1865a:479 , pl. 33, fig. 1. Syntypes: BMNH 1947.2.29.10–11 (collector F.M.J.

Welwitsch), formerly BMNH 64.7.13.5–6. Type locality: “Pungo Andongo” and “ W. Africa ” ( Günther

1865a:479), Angola. Preoccupied by Rana rugosa Temminck and Schlegel, 1838 . Rana tuberculosa Boulenger 1882:30 . Replacement name for Pyxicephalus rugosus Günther, 1865 . Rana (Tomopterna) signata Ahl “1923” 1925:43. Syntypes: ZMB 6458, 7765 [2 specimens] (donor J.V.B.

Bocage). Type locality: “Huilla” [= Huíla], Malanje Province, Angola. Synonymy by Laurent (1954a:72). Rana (Tomopterna) cacondana Ahl “1923” 1925:43. Holotype: ZMB 15477 (purchased Linnea). Type locality: “Caconda,” Huíla Province, Angola. Synonymy by Laurent (1954a:72). Pyxicephalus rugosus : Bocage (1870:68, 1873b:226, 1887c:211). Rana tuberculosa Bocage (1895a:156 , 1897b:211), Ferreira (1904:111), Schmidt (1936:130), Mertens

(1937a:19), Schmidt and Inger (1959:36). Rana (Tomopterna) tuberculosa : Monard (1938:102). Pyxicephalus tuberculosus : Parker (1939:142), Haacke (1970:276). Tomopterna signata : Cei (1997:17). Tomopterna rugosa : Laurent (1954a:72). Tomopterna tuberculosa : Loveridge (1957:345), Laurent (1964a:133), Cei (1997:17), Poynton and Broadley

(1985b:130), Poynton and Haacke (1993:13), (1996:23), Channing (2001:372), Frost (2016).

Tomopterna (Tomopterna) tuberculosa : Frétey et al. (2011:43).

Global conservation status (IUCN): Least Concern.

Global distribution: This is a widespread species occurring in savannas extending from Angola and northwestern Namibia across southern Democratic Republic of Congo to Tanzania, Zambia, and Zimbabwe.

Ocurrences in Angola (Map 102): The species occurs throughout much of Angola.

Malanje: “Pungo-Andongo” [-9.66667,

15.58333] ( Günther 1865a:480, 1869:479;

Boulenger 1882:30; Bocage 1895a:156;

Loveridge 1957:345; Schmidt and Inger

1959:36; Poynton and Broadley 1985b:130;

Ruas 1996:23). Kwanza Sul: “Quirimbo”

[-10.68333, 14.26667] ( Parker 1936:142; Ruas

1996:23); “Novo Redondo” [-11.20000,

13.85000] ( Ferreira 1904:111; Schmidt and

Inger 1959:37; Ruas 1996:23). Moxico:

“ Réserve de Cameia (Cameia)” [-11.83333,

21.00000] ( Laurent 1964a:133; Ruas 1996:23).

Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt

1936:130; Ruas 1996:23); “General Machado”

[-12.03333, 17.46667] ( Mertens 1937a:19).

Huambo: “Bimbi” [-11.81667, 15.83333] MAP 102. Distribution of Tomopterna tuberculosa in Angola .

( Monard 1938:102; Ruas 1996:23); “Galanga”

[-12.06667, 15.15000] ( Bocage 1895a:156; Ruas 1996:23); “ Mt. Moco ” [-12.41667, 15.18333] ( Parker 1936:142; Ruas 1996:23). Benguela: “Quissange” [-12.43333, 14.05000] ( Bocage 1887c:211, 1895a:156; Ruas 1996:23); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:156; Ruas 1996:23); “Ebanga” [-12.73333, 14.73333] ( Monard 1938:102; Ruas 1996:23); “Hanha” [-13.30000, 14.20000] ( Bocage 1897b:211; Ruas 1996:23). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:156; Ahl “1923” 1925:45; Ruas 1996:23); “Kalukembé (Caluquembe)” [-13.78333, 14.68333] ( Monard 1938:102; Ruas 1996:23); “ 13 km N of Cutenda (Cutenda 13 km N)” [-14.25000, 15.05000] (Poynton and Haacke 1993:13; Ruas 1996:23); “ 7 km N of Cutenda (Cutenda 7 km N)” [-14.36667, 15.08333] (Poynton and Haacke 1993:13; Ruas 1996:23); “Kuvangu” [-14.46667, 16.30000] ( Monard 1938:102; Ruas 1996:23); “Huilla” [-15.05000, 13.55000] ( Bocage 1873b:226, 1895a:156; Ahl “1923” 1925:45; Ruas 1996:23); “ 7 km SE of Jau (Jau 7 km)” [-15.21667, 13.51667] (Poynton and Haacke 1993:13); “ Cascade de Ongueria (Chibia)” [-15.30000, 13.51667] ( Laurent 1954a:72; Ruas 1996:23); “Christo Rei (Cristo Rei)” [-14.93333, 13.51667] (Poynton and Haacke 1993:13; Ruas 1996:23). Cunene: “riusseau Mbalé (Bale)” [-15.16667, 16.75000] ( Monard 1938:102; Ruas 1996:23). Namibe: “Biballa (Bibala)” [-14.76667, 13.36667] ( Bocage 1895a:156; Ruas 1996:23); “Bottom of Leba Pass (Leba)” [-15.06667, 13.23333] (Poynton and Haacke 1993:13; Ruas 1996:23). Undetermined Locality: without precise locality ( Bocage 1870:68).

Taxonomic and distributional notes: The nomen Pyxicephalus rugosus was preoccupied and was replaced by Rana tuberculosa Boulenger, 1882 . Ahl (1925 “1923”) described two new species of Tomopterna based on Angolan types, which Laurent (1954a) subsequently treated as junior synonyms of what is now T. tuberculosa .