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Chersobius signatus (GMELIN, 1789)

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Higher TaxaTestudinidae, Testudinoidea, Testudines (turtles) 
Subspecies 
Common NamesSpeckled Cape Tortoise, Speckled Padloper 
SynonymTestudo signata WALBAUM 1782 (nomen illegitimum)
Testudo signata GMELIN in LINNAEUS 1789
Testudo signata SCHOEPFF 1801
Testudo cafra DAUDIN 1802
Testudo juvencella DAUDIN 1831 (in GRAY)
Homopus signatus — DUMÉRIL & BIBRON 1835: 152
Testudo (Chersobius) signata — FITZINGER 1836: 122
Testudo (Chersobius) cafra — FITZINGER 1836: 122
Testudo signata — BELL 1836-1842
Homopus Signatus — DUMÉRIL & BIBRON 1854: 222
Homopus signatus — GRAY 1873: 320
Pseudomopus signatus — HEWITT 1931: 496
Pseudomopus signatus peersi HEWITT 1935
Chersobius signatus — HEWITT 1937
Homopus signatus — BOUR 1980
Homopus signatus cafer — IVERSON 1992: 267
Homopus signatus — VALVERDE 2005
Homopus signatus — TTWG 2014
Chersobius signatus — HOFMEYR et al. 2016 
DistributionRepublic of South Africa (Little Namaqualand, from Kleinzee and Springbok southward to Graafwater and Clanwilliam)

Type locality: unknown (fide WERMUTH & MERTENS 1977); designated as "vicinity of Springbok. Cape Province, South Africa" by Bour (1988:3)

cafer: near Clanwilliam and southwestward, Republic of South Africa]; Type locality: "Cafrerie" [= Kaffraria, Republic of South Africa], restricted to "Drainage of the Olifants River, Cape Province, South Africa" by Bour (1988a:3)  
Reproductionoviparous. 
TypesHolotype: nondesignated; lecotype: the Plate labelled "Testud. Sign. var." in Walbaum (1782), designated by Bour (1988a: 1).
Holotype: MNHN 7924 [cafer]
Syntype: DNMNH (= TM) 14878 [Pseudomopus signatus peersi] 
DiagnosisDiagnosis (genus): All Chersobius species have a single, large inguinal scute, whereas the number is variable in Homopus. The midline pectoral scute of Chersobius is consistently longer, and the nuchal scute shorter, than in Homopus. The fourth to fifth vertebral suture is narrow in Homopus and broad in Chersobius. Forelimb scales also differ with 3–4 large rows and 5–6 smaller rows, respectively, in Homopus and Chersobius. Chersobius males have a plastral concavity and Homopus males not. Two Chersobius species (C. signatus and C. boulengeri) are distinguishable from Homopus by having 12 or more marginal scutes, whereas Homopus species have 11 or fewer. The glans penes are respectively V-shaped and heart-shaped in Homopus and Chersobius. Ecological differences include that Chersobius species are rupicolous and inhabit arid regions, whereas Homopus species inhabit more mesic habitats. Homopus females produce multi-egg clutches and tend to be bigger compared to single-egg clutching Chersobius females. Females of all species are larger than conspecific males, and all males have larger shell openings than females have (Hofmeyr & Branch 2018). 
CommentConservation: Homopus signatus cafer is one of the most endangered turtle species according to a 2003 assessment by the IUCN.

Type species: Testudo signata WALBAUM 1782 is the type species of the genus Pseudomopus HEWITT 1931 and the (sub-) genus Chersobius FITZINGER 1835. HEWITT 1937 realized that Pseudomopus is a synonym of Chersobius though. Loveridge & Williams 1957 synonymized both with Homopus. See also discussion in Lindholm 1929: 284. However, Hofmeyr et al. 2016 resurrected Chersobius. 
Etymology 
References
  • Bates, M.F.; Branch, W.R., Bauer, A.M.; Burger, M., Marais, J.; Alexander, G.J. & de Villliers, M.S. (eds.) 2014. Atlas and Red List of the Reptiles of South Africa, Lesotho, and Swaziland. Suricata 1. South African National Biodiversity Institute, Pretoria, 512 pp.
  • Bauer, A.M., and Branch, W.R. 2003. The herpetofauna of the Richtersveld National Park, Northern Cape Province, Republic of South Africa. Herpetological Natural History 8: 111-160 [2001]
  • Bayoff, Nicolas 1995. Observations and morphometric data on the Namaqualand speckled tortoise, Homopus signatus signatus (Gmelin, 1789), in South Africa. Chelonian Conserv. Biol. 1 (3): 215-220
  • Bell, T. 1836. A monograph of the Testudinata. London, 1836-1842 (sometimes given as 1832-1836, not date given, unnumbered pages) - get paper here
  • Bonin, F., Devaux, B. & Dupré, A. 2006. Turtles of the World. English translation by P.C.H. Pritchard. Johns Hopkins University Press, 416 pp.
  • Boulenger,G.A. 1888. On new or little known South African reptiles. Ann. Mag. nat. Hist. (6) 2: 136-141 - get paper here
  • Bour, R. H. 1988. Taxonomic and Nomenclatural Status of Homopus signatus. J. Herp. Assoc. Africa (35): 1-6 - get paper here
  • Bour,R. 1980. Essai sur la taxinomie des Testudinidae actuels (Reptilia, Chelonii). Bull. Mus. natl. Hist. nat. Paris (4) 2 (2): 541-546
  • Boycott, R. 1987. A review of Homopus signata (Schoepff) with notes on related species (Cryptodira: Testudinidae). J. Herp. Assoc. Africa 32:10-16 1986] - get paper here
  • Branch, William R. 1993. A Photographic Guide to Snakes and Other Reptiles of Southern Africa. Cape Town: Struik Publishers, 144 S.
  • Dobiey, M. 2006. Gesägte Flachschildkröte - Homopus signatus signatus (GMELIN 1789). Reptilia (Münster) 11 (61): 51-54 - get paper here
  • Duerden, J.E. 1906. South African tortoises of the genus Homopus, with description of a new species. Records of the Albany Museum 1: 405-411 - get paper here
  • Duméril, A. M.C., G. BIBRON & A. DUMÉRIL 1854. Erpétologie générale ou Histoire Naturelle complète des Reptiles. Vol. 9. Paris, XX + 440 S. - get paper here
  • Duméril, A.M.C., and G. Bibron. 1835. Erpétologie Générale ou Histoire Naturelle Complète des Reptiles, Vol. 2. Librairie Encyclopédique de Roret, Paris, iv + 680 p. - get paper here
  • Ernst,C.H. and Barbour,R.W. 1989. Turtles of the World. Smithsonian Institution Press, Washington D.C. - London
  • Fitzinger, Leopold 1836. Entwurf einer Systematischen Anordnung der Schildkröten nach den Grundsätzen der Natürlichen methode. Annalen des Wiener Museums der Naturgeschichte 1 (1): 103-128 [1835][Facsimile Reprint, SSAR 1997] - get paper here
  • Gmelin, J.F. 1789. Caroli a Linné Systema naturae. 13. ed., Tom 1 Pars 3. G. E. Beer, Lipsiae. 1033-1516 pp. - get paper here
  • Gray,J.E. 1873. Additional notes on the form of the bones in the sternum of very young tortoises, and their development. Ann. Mag. Nat. Hist. (4) 12: 319-323 - get paper here
  • Greig, J.C., and P.D. Burdett. 1976. Patterns in the distributions of Southern African terrestrial tortoises (Cryptodira: Testudinidae). Zool. Africana 11(2): 250-267.
  • Hewitt, J. 1931. Descriptions of some African tortoises. Annals of the Natal Museum, Pietermaritzburg. 6: 461-506.
  • Hewitt, J. 1935. Some new forms of batrachians and reptiles from South Africa. Rec. Albany Mus. 4: 283-357
  • Hewitt, J. 1937. S. Afr. J. Sci. 33: 791
  • Hofmeyr, Margaretha D. & William R. Branch 2018. The padloper’s tortuous path (Chelonia: Testudinidae): Two genera, not one. African Journal of Herpetology 67 (2): 99-112 - get paper here
  • Hofmeyr, Margaretha D.; Melita Vamberger, William Branch, Alfred Schleicher and Savel R. Daniels 2016. Tortoise (Reptilia, Testudinidae) radiations in Southern Africa from the Eocene to the present. Zoologica Scripta 46: 389–400, DOI: 10.1111/zsc.12223 [Record online: 18 NOV 2016] - get paper here
  • Kooijman, Martijn. 2015. Constipation in a specimen of the Speckled Cape Tortoise, Homopus signatus (Gmelin, 1789). Radiata (English edition) 24 (4): 16-24
  • Loehr, V. J. T., B. T. Henen and M. D. Hofmeyer 2004. Reproduction of the smalles tortoise, the Namaqualand Speckled Padloper, Homopus signatus signatus. Herpetologica 60 (4):444-454 - get paper here
  • LOEHR, V.J .T. 2008. Homopus s. signatus (Gmelin, 1789). Natural oviposition and incubation. African Herp News (44): 9 - get paper here
  • Loehr, Victor J. T. 2006. Natural diet of the Namaqualand Speckled Padloper Homopus signatus signatus. Chelonian Conserv. Biol. 5 (1): 149-152
  • Loehr, Victor J. T. 2015. Small Vernal Home Ranges in the Namaqualand Speckled Tortoise, Homopus signatus Journal of Herpetology Sep 2015, Vol. 49, No. 3: 447-451. - get paper here
  • Loehr, Victor J. T., Brian T. Henen and Margaretha D. Hofmeyr 2006. Shell characteristics and sexual dimorphism in the Namaqualand Speckled Padloper, Homopus signatus signatus. African Journal of Herpetology 55(1):1-11. - get paper here
  • Loehr, Victor J. T.; Brian T. Henen, and Margaretha D. Hofmeyr 2011. Reproductive Responses to Rainfall in the Namaqualand Speckled Tortoise. Copeia 2011 (2): 278-284. - get paper here
  • Loehr, Victor J.T. 2016. Wide variation in carapacial scute patterns in a natural population of speckled tortoises, Homopus signatus. African Journal of Herpetology 65 (1): 47-54; DOI: 10.1080/21564574.2016.1146168 - get paper here
  • Loehr, Victor J.T.; Carlos G.P. Voogdt, Declan Nolan 2019. Ultrastructure of Eggshells from Wild and Captive Speckled Dwarf Tortoises, Chersobius signatus. Herpetologica 75 (1): - get paper here
  • Loehr, Victor J.T.; Tariq Stark, Martijn Weterings and Henry Kuipers. 2015. Overcoming low environmental temperatures in the primary feeding season: low-level activity and long basking in the tortoise Homopus signatus. Amphibia-Reptilia 36 (3): 207–214 - get paper here
  • LOON, F. H.A. VAN 2018. Homopus signatus - Gmelin, 1789. Two-egg clutching. African Herp News (67): 25 - get paper here
  • Loveridge, Arthur & Williams, Ernest E. 1957. Revision of the African Tortoises and Turtles of the Suborder Cryptodira. Bull. Mus. Comp. Zool. Harvard 115 (6): 163-557 - get paper here
  • Mashinini, P.L.; Mahlangu, L.M. 2014. An annotated catalogue of the types of chelonians (Reptilia: Testudines) in the Herpetology collection of the Ditsong National Museum of Natural History, South Africa. Annals of the Ditsong National Museum of Natural History 4: - get paper here
  • Schoepff, J.D. 1792. Historia Testudinum Iconibus Illustrata. [part 1]. J.J. Palm, Erlangae [=Erlangen]. xii + 136 pp. [pp. 1-32] - get paper here
  • STARK, T. 2013. Homopus signatus (Gmelin, 1789) Diet. African Herp News (59): 19-21 - get paper here
  • TTWG [Peter Paul van Dijk, John B. Iverson, Anders G.J. Rhodin, H. Bradley Shaffer, and Roger Bour] 2014. Turtles of the World, 7th Edition: Annotated Checklist of Taxonomy, Synonymy, Distribution with Maps, and Conservation Status. 000.v7. Chelonian Research Monographs (ISSN 1088-7105) No. 5, doi:10.3854/crm.5.000.checklist.v7.2014 - get paper here
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  • Valverde, J. 2005. African Tortoises. Reptilia (GB) (43): 12-20 - get paper here
  • Valverde, J. 2005. Afrikanische Landschildkröten. Reptilia (Münster) 10 (56): 18-25 - get paper here
 
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