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Chapter 3 - The Miocene

Published online by Cambridge University Press:  22 April 2019

D. Margaret Avery
D. Margaret Avery
Affiliation:
Iziko Museums of South Africa

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A Fossil History of Southern African Land Mammals , pp. 12 - 38
Publisher: Cambridge University Press
Print publication year: 2019
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3.1 Miocene Mammals

During the Miocene only one extinct Order, Creodonta, is represented in southern Africa, but the total number of Orders rose to 14, with only the Soricomorpha (shrews) and Pholidota (pangolins) not present. This epoch is notable for having the largest number of families, of which slightly more than a third are extinct (Table 1.1), and for the diversity of elephants (Proboscidea), of which there are nine genera and eight species in four families. The Rodentia continued to be the most diverse Order, with 36 genera and 41 species in 12 families. Next most diverse at the family level, with six families each, are the Chiroptera (bats), Carnivora (carnivores) and Artiodactyla (even-toed ungulates), the first two of which appeared for the first time.

ORDER: AFROSORICIDA

Suborder: Tenrecomorpha

FAMILY: TENRECIDAE

†Promicrogale namibiensis Pickford, 2018. Comm. Geol. Surv. Namibia 18: 88, 89.
Type locality: Elisabethfeld.
Comments: Pickford (Reference Pickford2018d) places the Tenrecidae in Order Soricomorpha, but points out (personal communication, 2018) that there is considerable disagreement on the allocation of this family. For now, Wilson and Reeder (Reference Wilson and Reeder2005) are followed in placing Tenrecidae in Afrosoricida.

Subfamily: †Protenrecinae

†Protenrec butleri Mein and Pickford, 2003. Mem. Geol. Soc. Namibia 19: 145.
Type locality: Arrisdrift.
Additional references: Butler and Hopwood, (Reference Butler and Hopwood1957); Mein and Pickford (Reference Mein and Pickford2008a).

Suborder: Chrysochloridea

FAMILY: CHRYSOCHLORIDAE

Subfamily: †Prochrysochlorinae

†Prochrysochloris miocaenicus Butler and Hopwood, Reference Butler and Hopwood1957. Foss. Mamm. Afr. 13: 11.
Additional references: Butler (Reference Butler1984); Mein and Pickford (Reference Mein and Pickford2008a).

ORDER: MACROSCELIDEA

FAMILY: MACROSCELIDIDAE

Subfamily: Macroscelidinae

Elephantulus Thomas and Schwann, Reference Thomas and Schwann1906. Abstr. Proc. Zool. Soc. Lond. 33: 10.
Additional references: Corbet and Hanks (Reference Corbet and Hanks1968); Evans (Reference Evans1942); Holroyd (Reference Holroyd, Werdelin and Sanders2010a); Patterson (Reference Patterson1965); Van der Horst (Reference Van der Horst1944).
†Miorhynchocyon gariepensis Senut, Reference Senut2003. Geol. Soc. Namibia Mem. 19: 126.
Type locality: Arrisdrift.
†Palaeothentoides Stromer, Reference Stromer1931. Sitzungsber. Bayer. Akad. Wiss. München. Math.-Nat. Abt. 1931: 185.
Type locality: Kleinzee.
Additional references: Holroyd (Reference Holroyd, Werdelin and Sanders2010a); Patterson (Reference Patterson1965).

Subfamily: †Myohyracinae

†Myohyrax oswaldi Andrews, Reference Andrews1914. Quart. J. Geol. Soc. Lond. 70: 171.
Synonyms: doederleini; osborni.
Additional references: Holroyd (Reference Holroyd, Werdelin and Sanders2010a); Leakey (Reference Leakey1943a); Patterson (Reference Patterson1965); Stromer (Reference Stromer1923).
†Myohyrax pickfordi Senut, Reference Senut2008. Geol. Surv. Namibia Mem. 20: 189.
Type locality: Langental.
†Protypotheroides beetzi Stromer, Reference Stromer1921. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1921: 333.
Synonyms: osborni.
Type locality: Langental.
Additional references: Holroyd (Reference Holroyd, Werdelin and Sanders2010a); Patterson (Reference Patterson1965); Stromer (Reference Stromer1923).
Comments: Stromer (Reference Stromer1923: 258) refers to this taxon as Protypotheroides beetzi Stromer (1922, S. 333), but see note in Section 1.3 regarding journal volume numbers.

Subfamily: Rhinchocyoninae

†Brevirhynchocyon gariepensis Senut, Reference Senut2003. Geol. Surv. Namibia Mem. 19: 126.
Synonyms: Brachyrhynchocyon, Miorhynchocyon.
Type locality: Arrisdrift.
Additional references: Butler (Reference Butler1984); Senut and Georgalis (Reference Senut and Georgalis2014).
†Brevirhynchocyon jacobi Senut, Reference Senut2008. Geol. Surv. Namibia Mem. 20: 192.
Synonyms: Brachyrhynchocyon.
Type locality: Elisabethfeld.
Additional references: Senut and Georgalis (Reference Senut and Georgalis2014).
†Hypsorhynchocyon burrelli Senut, Reference Senut2008. Geol. Surv. Namibia Mem. 20: 194.
Type locality: Grillental 6.

ORDER: TUBULIDENTATA

FAMILY: ORYCTOPODIDAE

†Amphiorycteropus Lehmann, Reference Lehmann2009. Zool. J. Linn. Soc. 145: 665.
Synonyms: Myorycteropus minutus; Orycteropus.
Additional references: Holroyd (Reference Holroyd, Werdelin and Sanders2010b); Pickford (Reference Pickford1975, Reference Pickford1996b, Reference Pickford2003a).
Orycteropus Cuvier, Reference Cuvier1798. Tabl. Elem. Hist. Nat. Anim. 1798: 144.
Additional references: Holroyd (Reference Holroyd, Werdelin and Sanders2010b); Kitching (Reference Kitching1963); Lehmann (Reference Lehmann2007); Pickford (Reference Pickford2008a).

ORDER: HYRACOIDEA

FAMILY: †PLIOHYRACIDAE

†Parapliohyrax Lavocat, Reference Lavocat1961. Notes Mém. Serv. Géol. Maroc 155: 87.
Additional references: Pickford (Reference Pickford1996a).
†Parapliohyrax ngororaensis Pickford and Fischer, Reference Pickford and Fischer1987. N. Jhb. Geol. Palaeontol. Abh. 175: 212.
Additional references: Pickford (Reference Pickford2003b); Rasmussen and Gutiérrez (Reference Rasmussen, Gutiérrez, Werdelin and Sanders2010).
†Prohyrax hendeyi Pickford, Reference Pickford1994. Geol. Surv. Namibia Comm. 9: 45.
Type locality: Arrisdrift.
Additional references: Pickford (Reference Pickford2003b); Rasmussen and Gutiérrez (Reference Rasmussen, Gutiérrez, Werdelin and Sanders2010).
†Prohyrax tertiarius Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 256.
Type locality: Langental.
Additional references: Rasmussen and Gutiérrez (Reference Rasmussen, Gutiérrez, Werdelin and Sanders2010).

FAMILY: PROCAVIIDAE

†Heterohyrax auricampensis Rasmussen, Pickford, Mein, Senut and Conroy, Reference Rasmussen, Pickford, Mein, Senut and Conroy1996. J. Mammal. 77(3): 746.
Type locality: Berg Aukas 1.
Additional references: Rasmussen and Gutiérrez (Reference Rasmussen, Gutiérrez, Werdelin and Sanders2010).

FAMILY: †TITANOHYRACIDAE

†Afrohyrax namibiensis Pickford, 2008. Mem. Geol. Surv. Namibia 20: 314.
Type locality: Grillental 6.
Additional references: Pickford and Senut (Reference Pickford and Senut2018).

ORDER: PROBOSCIDEA

FAMILY: †DEINOTHERIIDAE

Subfamily: †Deinotheriinae

†Prodeinotherium hobleyi Andrews, Reference Andrews1911. Proc. Zool. Soc. Lond. 1911: 944.
Synonyms: Deinotherium; Dinotherium.
Additional references: Éhik (Reference Éhik1930); Harris (Reference Harris1977); Sanders et al. (Reference Sanders, Gheerbrandt, Harris, Saegusa, Delmer, Werdelin and Sanders2010a).

FAMILY: ELEPHANTIDAE

Subfamily: Elephantinae

Loxodonta Anonymous, 1827. Zool. J. 3: 140.
Synonyms: Tetralophodon?.
†Loxodonta cookei Sanders, Reference Sanders2007. Trans. R. Soc. S. Afr. 62(1): 7.
Synonyms: Lukeino Stage.
Type locality: Langebaanweg.
Additional references: Pickford and Senut (Reference Pickford and Senut1997); Sanders et al. (Reference Sanders, Gheerbrandt, Harris, Saegusa, Delmer, Werdelin and Sanders2010a); Todd (Reference Todd2010).

FAMILY: †GOMPHOTHERIIDAE

Subfamily: †Amebelodontinae

†Afromastodon coppensi Pickford, 2003. Geol. Surv. Namibia Mem. 19: 219.
Type locality: Arrisdrift.
Additional references: Sanders et al. (Reference Sanders, Gheerbrandt, Harris, Saegusa, Delmer, Werdelin and Sanders2010a).
†Progomphotherium maraisi Pickford, Reference Pickford2003c. Geol. Surv. Namibia Mem. 19: 211.
Type locality: Auchas.
Additional references: Osborn (Reference Osborn1934); Sanders et al. (Reference Sanders, Rasmussen, Kappelman, Werdelin and Sanders2010b).

Subfamily: †Anancinae

†Anancus kenyensis MacInnes, Reference MacInnes1942 . Trans. Zool. Soc. Lond. 25(2): 33–106.
Additional references: Cooke (Reference Cooke1993b); Hautier et al. (Reference Hautier, Mackaye, Lihoreau and Tassy2009); Pickford and Senut (Reference Pickford and Senut1997); Sanders et al. (Reference Sanders, Rasmussen, Kappelman, Werdelin and Sanders2010b); Tassy (Reference Tassy1986).

Subfamily: †Choerolophodontinae

†Afrochoerodon kisumuensis MacInnes, Reference MacInnes1942. Trans. Zool. Soc. Lond. 25(2): 51.
Additional references: Pickford (Reference Pickford2001a, Reference Pickford2005a); Tassy (Reference Tassy1986).
†Gomphotherium pygmaeus Depéret, Reference Depéret1897. Bull. Soc. Géol. France, Series 3, 25: 520.
Synonyms: Choerolophodon, Mastodon; angustidens.
Additional references: Beck (Reference Beck1906); Fraas (Reference Fraas1907); Hamilton (Reference Hamilton1973); Pickford (Reference Pickford2005a); Sanders et al. (Reference Sanders, Gheerbrandt, Harris, Saegusa, Delmer, Werdelin and Sanders2010a).

FAMILY: †MAMMUTHIDAE

†Eozygodon morotoensis Pickford and Tassy, Reference Pickford and Tassy1980. Neues Jahrb. Geol. Palaontol., Monatsch. 4: 242.
Synonyms: Zygolophodon aegyptensis cf.
Additional references: Pickford (Reference Pickford2007, Reference Pickford2008c).

ORDER: PRIMATES

Suborder: Haplorrhini

FAMILY: CERCOPITHECIDAE

Subfamily: Colobinae

†Microcolobus Benefit and Pickford, Reference Benefit and Pickford1986. Amer. Phys. Anthrop. 69: 446.

FAMILY: HOMINIDAE

Subfamily: Homininae

†Otavipithecus namibiensis Conroy, Pickford, Senut, Van Couvering and Mein Reference Conroy, Pickford, Senut, Van Couvering and Mein1992. Nature 356: 144.
Type locality: Berg Aukas I.
Additional references: Gommery (Reference Gommery2000); Harrison (Reference Harrison, Werdelin and Sanders2010); Pickford et al. (Reference Pickford, Senut and Mein1992 Reference Pickford1997); Singleton (Reference Singleton2000).

Subfamily: †Kenyapithecinae

†Kenyapithecus Leakey, Reference Leakey1961. Ann. Mag. Nat. Hist., Series 13, 4: 690.
Additional references: McCrossin and Benefit (Reference McCrossin, Benefit, Begun, Ward and Rose1997); Pickford and Senut (Reference Pickford and Senut1997).
Comments: specimen listed as Hominoidea by Senut et al. (Reference Senut, Pickford and Wessels1997) and as a proconsulid (Nyanzapithecinae indet.) by Harrison (Reference Harrison, Werdelin and Sanders2010).

ORDER: RODENTIA

Suborder: Sciuromorpha

FAMILY: SCIURIDAE

Subfamily: Xerinae

†Heteroxerus karsticus Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia 12: 381.
Synonyms: Vulcanisciurus.
Type locality: Harasib 3a.
Additional references: Viriot et al. (Reference Viriot, Pelaez-Campomanes and Vignaud2011).
Comments: the version of this publication available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.
†Vulcanisciurus Lavocat, Reference Lavocat1973. Mem. Trav. Inst. Montpellier Ecole Prat. Hautes Etudes 1: 1–284.
Comments: Lavocat (Reference Lavocat1973) not seen.
†Vulcanisciurus africanus Lavocat, Reference Lavocat1973. Mem. Trav. Inst. Montpellier Ecole Prat. Hautes Etudes 1: 1–284.
Additional references: Viriot et al. (Reference Viriot, Pelaez-Campomanes and Vignaud2011).
Comments: Lavocat (Reference Lavocat1973) not seen.

Suborder: Myomorpha

FAMILY: GLIRIDAE

Subfamily: Graphiurinae

†Otaviglis daamsi Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia 12: 383.
Type locality: Harasib 3a.
Comments: the version of this publication available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.

FAMILY: SPALACIDAE

Subfamily: Rhizomyinae

†Harasibomys Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia, 12: 387.
Synonyms: Brachyuromys cf.
Type locality: Harasib 3a.
Comments: the version of this publication available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.
†Harasibomys petteri Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia, 12: 387.
Synonyms: Brachyuromys cf.
Type locality: Harasib 3a.
Comments: the version of this publication available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.
†Nakalimys lavocati Flynn and Sabatier, Reference Flynn and Sabatier1984. J. Paleontol. 3: 161.

FAMILY: NESOMYIDAE

Subfamily: Afrocricetodontinae

†Notocricetodon Lavocat, Reference Lavocat1973. Mem. Trav. Inst. Montpellier Ecole Prat. Hautes Etudes 1: 1–284.
Comments: Lavocat (Reference Lavocat1973) not seen.
†Protarsomys Lavocat, Reference Lavocat1973. Mem. Trav. Inst. Montpellier Ecole Prat. Hautes Etudes 1: 1–284.
Comments: Lavocat (Reference Lavocat1973) not seen.
†Protarsomys lavocati Mein and Pickford, Reference Mein and Pickford2003. Geol. Soc. Namibia Mem. 19: 148.
Type locality: Arrisdrift.
†Protarsomys macinnesi Lavocat, Reference Lavocat1973. Mem. Trav. Inst. Montpellier Ecole Prat. Hautes Etudes 1: 1–284.
Comments: Lavocat (Reference Lavocat1973) not seen.

Subfamily: Cricetomyinae

Saccostomus Peters, Reference Peters1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 11: 258.
†Saccostomus geraadsi Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Soc. Namibia 13: 43.
Synonyms: major; “K.”majus Geraads, Reference Geraads2001.

Subfamily: Dendromurinae

†Dendromus denysae Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Surv. Namibia 13: 49.
Type locality: Harasib 3a.
Steatomys Peters, Reference Peters1846. Bericht Verhandl. K. Preuss. Akad. Wiss. Berlin 11: 258.
†Steatomys harasibensis Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Surv. Namibia 13: 45.
Type locality: Harasib 3a.
†Steatomys jaegeri Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Surv. Namibia 13: 48.
Type locality: Harasib 3a.

Subfamily: †Otavimyinae

†Otavimys senegasi Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Surv. Namibia 13: 51.
Type locality: Harasib 3a.

Subfamily: Petromyscinae

†Harimyscus hoali Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia, 12: 395.
Type locality: Harasib 3a.
Petromyscus Thomas, Reference Thomas1926. Ann. Mag. Nat. Hist., Series 9, 17: 179.
Additional references: Petter (Reference Petter1967).

FAMILY: MURIDAE

Subfamily: Cricetodontinae

†Afaromys guillemoti Geraads, Reference Geraads1998. Palaeovertebrata 27(3–4): 205.
Additional references: Mein et al. (Reference Mein, Pickford and Senut2004).

Subfamily: Deomyinae

†Preacomys griffini Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Surv. Namibia 13: 57.
Synonyms: Karnimata.
Type locality: Harasib 3a.
†Preacomys karsticus Mein, Pickford and Senut, Reference Mein, Pickford and Senut2004. Comm. Geol. Surv. Namibia 13: 57.
Type locality: Harasib 3a.
†Preacomys kikiae Geraads, Reference Geraads2001. Palaeovertebrata 30: 91.
Additional references: Mein et al. (Reference Mein, Pickford and Senut2004).

Subfamily: Gerbillinae

†Dakkamyoides Lindsay, Reference Lindsay1988. Palaeovertebrata 18(2): 123.

Subfamily: †Myocricetodontinae

†Myocricetodon Lavocat, Reference Lavocat1952. C. R. Acad. Sci. Paris 235: 190.
†Mioharimys milleri Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia 12: 391.
Type locality: Harasib 3a.
†Mioharimys schneideri Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia 12: 394.
Type locality: Harasib 3a.

Subfamily: †Namibimyinae

†Namibimys angustidens Mein, Pickford and Senut, 2000. Comm. Geol. Soc. Namibia 12: 398.
Type locality: Berg Aukas I.

Suborder: Anomaluromorpha

FAMILY: †PARAPEDETIDAE

†Parapedetes Stromer, Reference Stromer1923. Sitz. Math.-Physik. Abt. Bayer. Akad. Wiss. München 1923 (II): 261.
Type locality: Elisabethfeld.
†Parapedetes namaquensis Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 261.
Type locality: Elisabethfeld.
Additional references: Pickford and Mein (Reference Pickford and Mein2011).

FAMILY: PEDETIDAE

†Megapedetes gariepensis Mein and Senut, Reference Mein and Senut2003. Geol. Soc. Namibia Mem. 19: 162.
Type locality: Arrisdrift.
Additional references: Pickford and Mein (Reference Pickford and Mein2011).
†Oldrichpedetes brigittae Pickford and Mein, Reference Pickford and Mein2011. Estud. Geol. 67(2): 465.
Type locality: Zebra Hill.
†Oldrichpedetes pickfordi Mein and Senut, Reference Mein and Senut2003. Geol. Soc. Namibia Mem. 19: 166.
Synonyms: Megapedetes.
Type locality: Arrisdrift.
Additional references: Pickford and Mein (Reference Pickford and Mein2011).
†Propedetes Mein and Pickford, 2008. Geol. Surv. Namibia Mem. 20: 242.
Type locality: Elisabethfeld.
†Propedetes efeldensis Mein and Pickford, 2008. Geol. Surv. Namibia Mem. 20: 242.
Type locality: Elisabethfeld.
†Propedetes laetoliensis Davies, Reference Davies, Leakey and Harris1987. In: Leakey and Harris. The Pliocene Site of Laetoli, Northern Tanzania: 172.
Synonyms: Pedetes.
Additional references: Pickford and Mein (Reference Pickford and Mein2011).

Suborder: Hystricomorpha

FAMILY: †DIAMANTOMYIDAE

Subfamily: †Diamantomyinae

†Diamantomys luederitzi Stromer, Reference Stromer1921. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1921: 334.
Type locality: Sperrgebiet.
Comments: Stromer (Reference Stromer1923: 265) refers to this taxon as Diamantomys luederitzi Stromer (1922, S. 334), but see note in Section 1.3 regarding journal volume numbers.
†Pomonomys dubius Stromer, Reference Stromer1921. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1921: 334.
Type locality: Grillental.
Comments: Stromer (Reference Stromer1923: 265) refers to this taxon as Pomonomys dubius Stromer (1922, S. 334/5), but see note in Section 1.3 regarding journal volume numbers.

FAMILY: †BATHYERGOIDIDAE

†Bathyergoides Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 263.
Type locality: Sperrgebiet.
†Bathyergoides neotertiarius Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 263.
Type locality: Sperrgebiet.
†Geofossor corvinusae Mein and Pickford, 2003. Geol. Soc. Namibia Mem. 19: 144.
Type locality: Arrisdrift.
†Geofossor moralesi Mein and Pickford, 2008. Geol. Surv. Namibia Mem. 20: 262.
Type locality: Elisabethfeld.
†Microfossor biradiculatus Mein and Pickford, 2008. Geol. Surv. Namibia Mem. 20: 262.
Type locality: Elisabethfeld.

FAMILY: BATHYERGIDAE

Subfamily: Bathyerginae

†Proheliophobius Lavocat, Reference Lavocat1973. Mem. Trav. Inst. Montpellier Ecole Prat. Hautes Etudes 1: 1–284.
Synonyms: Paracryptomys mackennae (see comment below).
Comments: Lavocat (Reference Lavocat1973) not seen. Mein et al. (Reference Mein, Pickford and Senut2000a: 386) state: ‘Denys and Jaeger (Reference Denys and Jaeger1992) provide measurements of a bathyergid (AD 1638) from Arrisdrift, Namibia which they identify as Paracryptomys mackennae, citing Hamilton and Van Couvering (Reference Hamilton and Van Couvering1977) as the source of these measurements. There are two points that should be made, firstly, the Arrisdrift species is far too small to belong to Paracryptomys mackennae, which is a giant bathyergid, possibly synonymous with Bathyergoides neotertiarius Stromer, and secondly, the site of Arrisdrift was unknown to Hamilton and Van Couvering as it was discovered in 1976 after they had submitted their manuscript for publication. In our opinion the Arrisdrift species is close to Proheliophobius but is slightly larger than the type species P. leakeyi Lavocat, Reference Lavocat1973. The version of Mein et al. (Reference Mein, Pickford and Senut2000a) available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.

FAMILY: †MYOPHIOMYIDAE

Subfamily: †Myophiomyinae

†Phiomyoides humilis Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 264.
Type locality: Sperrgebiet.

FAMILY: THRYONOMYIDAE

†Apodecter Hopwood, Reference Hopwood1929. Amer. Mus. Novit. 344: 3.
Type locality: Lüderitz Bay (south of).
†Apodecter stromeri Hopwood, Reference Hopwood1929. Amer. Mus. Novit. 344: 3.
Type locality: Lüderitz Bay (south of) (?Langental: Mein and Pickford, Reference Mein and Pickford2008c).
†Neosciuromys africanus Stromer, Reference Stromer1921. Sitz. Math.-Physik. Klass. Bayer. Akad. Wiss. München 1921 (II): 333.
Synonyms: fractus.
Type locality: Sperrgebiet.
Additional references: Stromer (Reference Stromer1923).
†Paraphiomys australis Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia 12: 378.
Type locality: Harasib 3a.
Comments: the version of this publication available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.
†Paraphiomys orangeus Mein and Pickford, 2003. Geol. Soc. Namibia Mem. 19: 143.
Type locality: Arrisdrift.
†Paraphiomys roessneri Mein, Pickford and Senut, 2000. Comm. Geol. Surv. Namibia 12: 375.
Synonyms: Apodecter cf.
Type locality: Harasib 3a.
Comments: the version of this publication available on the Geological Survey of Namibia website at www.mme.gov.na/files/publications/5dd_Mein%20et%20al_Late%20Miocene%20micromammals_Harasib.pdf appears to be wrongly paginated.
†Paraulacodus johanesi Jaeger, Michaux and Sabatier, Reference Jaeger, Michaux and Sabatier1980. Palaeovertebrata 9 ext. Mem. JubiI. R. Lavocat: 367.
†Phthinylla fracta Hopwood, Reference Hopwood1929. Amer. Mus. Novit. 344: 4.
Synonyms: Neosciuromys.
Type locality: Lüderitz Bay (south of) (?Langental: Mein and Pickford, Reference Mein and Pickford2008c).

FAMILY: TUFAMYIDAE

†Efeldomys loliae Mein and Pickford, 2008. Geol. Surv. Namibia Mem. 20: 259.
Type locality: Elisabethfeld.

ORDER: LAGOMORPHA

FAMILY: OCHOTONIDAE

†Austrolagomys hendeyi Mein and Pickford, 2003. Geol. Soc. Namibia Mem. 19: 172.
Synonyms: Kenyalagomys.
Type locality: Arrisdrift.
†Austrolagomys inexpectatus Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 261.
Type locality: Sperrgebiet.
Synonyms: Simpsoni.
Additional references: Hopwood (Reference Hopwood1929); Mein and Pickford (Reference Mein and Pickford2003d, Reference Mein and Pickford2008b).

ORDER: ERINACEOMORPHA

FAMILY: ERINACEIDAE

Subfamily: Erinaceinae

†Amphechinus rusingensis Butler, Reference Butler1956. Foss. Mamm. Africa 11: 54.
Additional references: Butler (Reference Butler1984, Reference Butler, Werdelin and Sanders2010); Mein and Pickford (Reference Mein and Pickford2008a).
†Gymnurechinus leakeyi Butler, Reference Butler1956. Foss. Mamm. Afr. 11: 3.
Additional references: Mein and Pickford (Reference Mein and Pickford2008a).

Subfamily: †Galericinae

†Galerix Pomel, Reference Pomel1848. Arch. Sci. Phys. Nat. Genève 9: 164.

ORDER: CHIROPTERA

Suborder: Microchiroptera

FAMILY: RHINOLOPHIDAE

Rhinolophus Lacépède, Reference Lacépède1799. Tableau des Divisions, Sous-divisions, Ordres, et Genres des Mammifères: 15.
Additional references: Andersen Reference Andersen1905); Taylor et al. (Reference Taylor, Stoffberg and Monadjem2012).
Comments: Lacépède, Reference Lacépède1799 not seen. Citation according to Wilson and Reeder (Reference Wilson and Reeder2005).
†Rhinolophus contrarius Mein and Pickford, Reference Mein and Pickford2003. Geol. Surv. Namibia Mem. 19: 115.
Type locality: Arrisdrift.

FAMILY: HIPPOSIDERIDAE

Asellia Gray, Reference Gray1838. Mag. Zool. Bot. 2: 493.
Additional references: Thomas (Reference Thomas1904a).
Hipposideros gigas Wagner, Reference Wagner1845. Arch. Naturgesch. 11(1): 148. Giant leaf-nosed bat.

FAMILY: MEGADERMATIDAE

Megaderma Geoffroy Saint-Hilaire, Reference Geoffroy Saint-Hilaire1810. Ann. Mus. Hist. Nat. Paris 14: 197.

FAMILY: EMBALLONURIDAE

†Taphozous incognita Butler and Hopwood, Reference Butler and Hopwood1957. Foss. Mamm. Africa 13: 29.
Synonyms: Saccolaimus.
Additional references: Butler (Reference Butler1984).

FAMILY: MOLOSSIDAE

Subfamily: Molossinae

Tadarida Rafinesque, Reference Rafinesque1814. Précis des découvertes et travaux somiologiques: 55.

FAMILY: VESPERTILIONIDAE

Subfamily: Myotinae

Myotis Kaup, Reference Kaup1829. Skizz. Entwickel.-Gesch. Nat. Syst. Europ. Thierwelt 1: 106.
Additional references: Gray (Reference Gray1842).

ORDER: †CREODONTA

FAMILY: †HYAENODONTIDAE

Subfamily: †Hyaenodontinae

†Metapterodon kaiseri Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 254.
Synonyms: Isohyaenodon.
Type locality: Elisabethfeld.
Additional references: Lewis and Morlo (Reference Lewis, Morlo, Werdelin and Sanders2010); Morales et al. (Reference Morales, Pickford and Salesa2008a); Savage (Reference Savage1965).
†Metapterodon stromeri Morales, Pickford and Soria, Reference Morales, Pickford and Soria1998. C.R. Acad. Sci. Paris, Sci. Terre Plan. 327: 634.
Synonyms: Pterodon.
Type locality: Langental.
Additional references: Lewis and Morlo (Reference Lewis, Morlo, Werdelin and Sanders2010); Morales et al. (Reference Morales, Pickford and Salesa2008a).

Subfamily: †Hyainailourinae

†Hyainailouros sulzeri Biedermann, Reference Biedermann1863. Petrefakten aus der Umgegend von Winterthur. Zweites Heft. 20.
Additional references: Helbing (Reference Helbing1924); Morales et al. (Reference Morales, Pickford, Soria and Fraile1998b, Reference Morales, Soria, Sánchez, Quiralte and Pickford2003d); Morlo et al. (Reference Morlo, Miller and El-Barkooky2007); Sach and Heizmann (Reference Sach and Heizmann2001).
Comments: Biedermann (Reference Biedermann1863) not seen.

FAMILY: †PRIONOGALIDAE

†Namasector soriae Morales, Pickford and Salesa, 2008. Geol. Surv. Namibia Mem. 20: 295.
Type locality: Elisabethfeld (Tortoise Site).
Comments: Werdelin and Cote (Reference Werdelin, Cote, Werdelin and Sanders2010) are doubtful that this family should be placed in Creodonta and prefer to retain Prionogalidae as Mammalia incertae sedis.

ORDER: CARNIVORA

Suborder: Feliformia

FAMILY: †BARBOUROFELIDAE

†Afrosmilus africanus Andrews, Reference Andrews1914. Quart. J. Geol. Soc. Lond. 70: 179.
Synonyms: Pseudaelurus; Metailurus.
Additional references: Savage (Reference Savage1965); Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010).

Subfamily: †Barbourofelinae

†Ginsburgsmilus napakensis Morales, Salesa, Pickford and Soria, Reference Morales, Salesa, Pickford and Soria2001. Trans. R. Soc. Edinburgh 92: 98.
Additional references: Morales and Pickford (Reference Morales and Pickford2018).

FAMILY: FELIDAE

†Diamantofelis ferox Morales, Pickford, Soria and Fraile, 1998. Eclogae Geol. Helv. 91(1): 36.
Type locality: Arrisdrift.
Additional references: Morales et al. (Reference Morales, Soria, Sánchez, Quiralte and Pickford2003d).
†Namafelis minor Morales, Pickford, Fraile, Salesa and Soria, Reference Morales, Pickford, Fraile, Salesa and Soria2003. Mem. Geol. Surv. Namibia 19: 184.
Type locality: Arrisdrift.

Subfamily: Felinae

†Metailurus Zdansky, Reference Zdansky1924. Palaeontol. Sinica, Series C, 2(1): 123.
Comments: publication not seen but citation confirmed by L. Werdelin (personal communication, 12 December 2016).

Subfamily: Pantherinae

Panthera leo Linnaeus, Reference Linnaeus1758. Systema Naturae Regnum Animale, 10th edition, 1, Lion.
Additional references: Haas et al. (Reference Haas, Hatssen and Krausman2005); Lacruz (Reference Lacruz2009).

FAMILY: †STENOPLESICTIDAE

†Orangictis gariepensis Morales, Pickford, Soria and Fraile, 2001. Palaeontol. Afr. 37: 99.
Type locality: Arrisdrift.
Additional references: Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010).

Subfamily: †Stenoplesictinae

†Africanictis hyaenoides Morales, Pickford, Fraile, Salesa and Soria, Reference Morales, Pickford, Fraile, Salesa and Soria2003. Geol. Soc. Namibia Mem. 19: 183.
Type locality: Arrisdrift.
†Africanictis meini Morales, Pickford, Soria and Fraile, 1998. Eclogae Geol. Helv. 91: 34.
Type locality: Arrisdrift.
†Africanictis schmidtkittleri Morales, Pickford, Soria and Fraile, 1998. Eclogae Geol. Helv. 91: 35.
Type locality: Arrisdrift.
Additional references: Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010).

FAMILY: VIVERRIDAE

†Leptoplesictis namibiensis Morales, Pickford and Salesa, 2008. Geol. Surv. Namibia Mem. 20: 304.
Type locality: Langental.
Comments: according to Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010), Leptoplesictis is generally considered to be a herpestid rather than a viverrid, although it was placed in the latter Family by Morales et al. (Reference Morales, Pickford and Salesa2008a).
†Leptoplesictis senutae Morales, Pickford and Salesa, 2008. Geol. Surv. Namibia Mem. 20: 303.
Type locality: Grillental.
Comments: see under Leptoplesictis namibiensis.

Suborder: Caniformia

FAMILY: †AMPHICYONIDAE

†Agnotherium Kaup, Reference Kaup1833. Déscription d’Ossements Fossiles de Mammifères 2: 28.
Additional references: Kurtén (Reference Kurtén1976); Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010).

Subfamily †Amphicyoninae

†Amphicyon giganteus Schinz, Reference Schinz1825. In: Cuvier, Thierreich IV: 342.
Synonyms: Pseudocyon; steinheimensis.
Additional references: Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010).
†Hecubides euryodon Savage, Reference Savage1965. Foss. Mamm. Africa 19: 189.
Additional references: Morales et al. (Reference Morales, Pickford and Valenciano2016).
†Ysengrinia Ginsburg, Reference Ginsburg1965. Bull. Mus. Natl d’Hist. Nat., Series 2, 37(4): 727.
Additional references: Werdelin and Peigné (Reference Werdelin, Peigné, Werdelin and Sanders2010).
†Ysengrinia ginsburgi Morales, Pickford, Soria, and Fraile, 1998. Eclogae Geol. Helv. 91(1): 30.
Type locality: Arrisdrift.

FAMILY: MUSTELIDAE

†Namibictis senuti Morales, Pickford, Soria, and Fraile, 1998. Eclogae Geol. Helv. 91(1): 32.
Type locality: Arrisdrift.

ORDER: PERISSODACTYLA

FAMILY: EQUIDAE

†Eurygnathohippus cornelianus Van Hoepen, Reference Van Hoepen1930. Paleontol. Nav. Nas. Mus. Bloemfontein 2(2): 23.
Synonyms: Hipparion; Stylohipparion; hipkini; libycum; sitifense in part?; steytleri.
Type locality: Cornelia.
Additional references: Bernor et al. (Reference Bernor, Armour-Chelu, Gilbert, Kaiser, Schulz, Werdelin and Sanders2010); Churcher (Reference Churcher1970, Reference Churcher2000); Churcher and Watson (Reference Churcher, Watson and Brain1993); Franz-Odendaal et al. (Reference Franz-Odendaal, Kaiser and Bernor2003); Pomel (Reference Pomel1897); Van Hoepen (Reference Van Hoepen1932a).
Comments: the material from Hondeklip Bay was doubtfully ascribed by Pickford and Senut (Reference Pickford and Senut1997) to Hipparion sitifense, which Bernor et al. (Reference Bernor, Armour-Chelu, Gilbert, Kaiser, Schulz, Werdelin and Sanders2010) consider a nomen dubium. See also the comment under Hipparion below.

FAMILY: RHINOCEROTIDAE

Subfamily: Rhinocerotinae

†Brachypotherium Roger, Reference Roger1902. Ber. Naturwiss. Ver. Schwaben und Neuburg 36: 12.
†Brachypotherium heinzelini Hooijer, Reference Hooijer1963. Ann. Mus. Roy. Afr. Cent., Tervuren Sci. Géol. 46: 45.
Synonyms: lewisi?.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010a); Guérin (Reference Guérin2008); Heissig (Reference Heissig1971); Hooijer (Reference Hooijer1973).
†Chilotheridium pattersoni Hooijer, Reference Hooijer1971. Bull. Mus. Comp. Zool. 144: 342.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010a); Guérin (Reference Guérin2003, Reference Guérin2008); Hooijer (Reference Hooijer1973).
†Diceros australis Guérin, Reference Guérin2000. Palaeontol. Afr. 36: 122.
Synonyms: Dicerorhinus.
Type locality: Arrisdrift.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010a); Guérin (Reference Guérin2003).
†Diceros praecox Hooijer and Patterson, Reference Hooijer and Patterson1972. Bull. Mus. Comp. Zool. 144: 19.
Synonyms: Ceratotherium.
Additional references: Geraads (Reference Geraads2005, Reference Geraads, Werdelin and Sanders2010a); Guérin (Reference Guérin, Leakey and Harris1987); Hooijer (Reference Hooijer1972 Reference Hooijer1973); Pickford and Senut (Reference Pickford and Senut1997).
†Turkanatherium acutirostrum Deraniyagala, Reference Deraniyagala1951. Spol. Zeyl. 26(2): 134.
Synonyms: Aceratatherium.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010a); Guérin (Reference Guérin2008); Hooijer (Reference Hooijer1973).

ORDER: ARTIODACTYLA

†Namibiomeryx senuti Morales, Soria and Pickford, Reference Morales, Soria and Pickford1995. C. R. Acad. Sci. Paris, Série lla, 321: 1211.
Type locality: Elisabethfeld.
Additional references: Cote (Reference Cote, Werdelin and Sanders2010).
†Namibiomeryx spaggiarii Morales, Soria and Pickford, 2008. Mem. Geol. Surv. 20: 442.
Type locality: Langental.
†Orangemeryx hendeyi Morales, Soria and Pickford, Reference Morales, Soria and Pickford1999. Geodiversitas, Series 21, 2: 241.
Synonyms: Climacoceras.
Type locality: Arrisdrift.
Additional references: Azanza et al. (Reference Azanza, Morales and Pickford2003); Cote (Reference Cote, Werdelin and Sanders2010); Hamilton (Reference Hamilton1978); MacInnes (Reference MacInnes1936); Morales et al. (Reference Morales, Soria, Sánchez, Quiralte and Pickford2003d).

Subfamily: †Sperrgebietomerycinae

†Sperrgebietomeryx wardi Morales, Soria and Pickford, Reference Morales, Soria and Pickford1999. Geodiversitas, Series 21, 2: 232.
Type locality: Elisabethfeld.
Additional references: Cote (Reference Cote, Werdelin and Sanders2010).

FAMILY: †CLIMACOCERATIDAE

Subfamily: †Propalaeorycinae

†Propalaeoryx austroafricanus Stromer, Reference Stromer1923. Sitz. Math.-Physik. Klasse Bayer. Akad. Wiss. München 1923(II): 256.
Type locality: Elisabethfeld.
Additional references: Cote (Reference Cote, Werdelin and Sanders2010).
†Propalaeoryx stromeri Morales, Soria and Pickford, 2008. Geol. Surv. Namibia Mem. 20: 410.
Type locality: Langental.

FAMILY: SUIDAE

Subfamily: †Kubanochoerinae

†Kenyasus namaquensis Pickford and Senut, Reference Pickford and Senut1997. Palaeontol. Afr. 34: 208.
Type locality: Ryskop.
Additional references: Bishop (Reference Bishop, Werdelin and Sanders2010); Pickford (Reference Pickford1986).
†Kenyasus rusingensis Pickford, Reference Pickford1986. Tert. Res. Spec. Pap. 7: 24.
Additional references: Bishop (Reference Bishop, Werdelin and Sanders2010); Pickford and Senut (Reference Pickford and Senut1997).
†Nguruwe namibensis Pickford, Reference Pickford1986. Tert. Res. Spec. Pap. 7: 25.
Synonyms: Hyotherium; Kenyasus; kijivium.
Type locality: Langental.
Additional references: Bishop (Reference Bishop, Werdelin and Sanders2010); Pickford (Reference Pickford1997, Reference Pickford2003e, Reference Pickford2008e).

Subfamily: †Namachoerinae

†Namachoerus moruoroti Wilkinson, Reference Wilkinson1976. Foss. Vert. Afr. 4: 173–282.
Synonyms: Lopholistriodon.
Additional references: Pickford (Reference Pickford1987, Reference Pickford1995, Reference Pickford2003f).

Subfamily: †Tetraconodontinae

†Nyanzachoerus kanamensis Leakey, Reference Leakey1958. Foss. Mamm. Afr. 14: 10.
Additional references: Bishop (Reference Bishop, Werdelin and Sanders2010); Harris and White (Reference Harris and White1979).

FAMILY: †ANTHRACOTHERIIDAE

†Brachyodus aequatorialis MacInnes, Reference MacInnes1951. Foss. Mamm. Afr. 4: 3.
Synonyms: Masritherium.
Additional references: Holroyd et al. (Reference Holroyd, Lihoreau, Gunnell, Miller, Werdelin and Sanders2010); Madden et al. (Reference Madden, Schmidt and Whitmore1983); Pickford (Reference Pickford1991a, Reference Pickford1996b, Reference Pickford2015f).
†Brachyodus depereti Fourtau, Reference Fourtau1918. Contribution à l’étude des vertébrés miocènes de l’Egypte: 64.
Synonyms: Masritherium.
Additional references: Holroyd et al. (Reference Holroyd, Lihoreau, Gunnell, Miller, Werdelin and Sanders2010); Miller et al. (Reference Miller, Gunnell and Gawad2014); Pickford (Reference Pickford2003d, Reference Pickford2008d).
Comments: Fourtau (Reference Fourtau1918) not seen. M. Pickford (personal communication 1 December 2016) believes that the earliest evidence for the transfer of this species to Brachyodus is that given by Dineur (Reference Dineur1982).

FAMILY: †SANITHERIIDAE

†Diamantohyus africanus Stromer, Reference Stromer1921. Sitz. Math.-Physik. Klass. Bayer. Akad. Wiss. München 1921 (II): 332.
Type locality: Sperrgebiet?
Additional references: Bishop (Reference Bishop, Werdelin and Sanders2010); Pickford (Reference Pickford1984, Reference Pickford1997, Reference Pickford2004, Reference Pickford2008f).
Comments: this taxon has been ascribed to Stromer (Reference Stromer and Kaiser1926) by Bishop (Reference Bishop, Werdelin and Sanders2010) and others.

FAMILY: TRAGULIDAE

†Dorcatherium moruorotensis Pickford, Reference Pickford2001. Geobios 34(4): 437.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010b); Quiralte et al. (Reference Quiralte, Sánchez, Morales and Pickford2008).
†Dorcatherium parvum Whitworth, Reference Whitworth1958. Foss. Mamm. Afr. 15: 11.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010b); Quiralte et al. (Reference Quiralte, Sánchez, Morales and Pickford2008).
†Dorcatherium pigotti Whitworth, Reference Whitworth1958. Foss. Mamm. Afr. 15: 9.
Additional references: Geraads (Reference Geraads, Werdelin and Sanders2010b); Morales et al. (Reference Morales, Soria, Sánchez, Quiralte and Pickford2003d); Pickford et al. (Reference Pickford, Senut and Mein1996); Quiralte et al. (Reference Quiralte, Sánchez, Morales and Pickford2008).
†Dorcatherium songhorensis Whitworth, Reference Whitworth1958. Foss. Mamm. Afr. 15: 14.
Additional references: Quiralte et al. (Reference Quiralte, Sánchez, Morales and Pickford2008).

FAMILY: BOVIDAE

†Namacerus gariepensis Morales, Soria, Pickford and Nieto, Reference Morales, Soria, Pickford and Nieto2003. Geol. Surv. Namibia Mem. 19: 372.
Type locality: Arrisdrift.
Additional references: Gentry (Reference Gentry, Werdelin and Sanders2010).

Subfamily: Alcelaphinae

†Damalacra acalla Gentry, Reference Gentry1980. Ann. S. Afr. Mus. 79(8): 272.
Type locality: Langebaanweg.
Additional references: Brink and Stynder (Reference Brink and Stynder2009).

Subfamily: Antilopinae

†Homoiodorcas Thomas, Reference Thomas1981. Proc. Kon. Nederl. Akad. Weten. Ser. B. Phys. Sci. 84(3): 364.

Subfamily: Bovinae

†Ugandax demissum Gentry, Reference Gentry1980. Ann. S. Afr. Mus. 79(8): 233.
Synonyms: Simatherium.
Type locality: Langebaanweg.
Additional references: Gentry (Reference Gentry2006, Reference Gentry, Werdelin and Sanders2010); Geraads (Reference Geraads1992); Pickford and Senut (Reference Pickford and Senut1997).

3.2 Miocene Sites

The 25 Miocene sites are restricted to west of 20°E and north of about 30°S (Figure 3.1). Particularly rich sites are Harasib 3a in the north, and Arrisdrift, Langental and Grillental in the south. Exploration of the southern African Miocene began fairly early, notably with the work of Stromer (Reference Stromer1921), but Corvinus (Reference Corvinus1978) and Hendey (Reference Hendey1978c) started what might be described as the modern era. However, the discovery of Otavipithecus namibiensis at Berg Aukas (Conroy et al. Reference Conroy, Senut, Gommery, Pickford and Mein1996) provided the impetus for the subsequent work undertaken by Martin Pickford and colleagues (see references), which greatly expanded coverage of the Miocene in the region.

Figure 3.1 Location of Miocene sites.

Figure 0

Figure 3.1 Location of Miocene sites.

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