AFRICAN CENOZOIC METATHERIANS: A REVIEW

1. AFRICAN CENOZOIC
METATHERIANS: A REVIEW
V.D. Crespo1,2*, and F.J. Goin1,2
1Museo de La Plata (UNLP), La Plata, Argentina
2CONICET (Consejo Nacional de Investigaciones Científicas y Técnicas), Argentina
*vidacres@gmail.com

2. Distribution
Map of Africa and the Arabian Peninsula
showing the fossil localities mentioned in this
work. Included are localities of both
metatherians and previously alleged
metatherians; B, Stratigraphic column of the
Cenozoic Era indicating the ages of the
metatherian taxa, or alleged metatherians,
mentioned in this work; E, Equator; 1,
Zhelestidae, gen. et sp. indet; 2, ?Garatherium
todrae; 3, Garatherium mahboubii; 4,
Kasserinotherium tunisiense; 5, Ghamidtherium
dimaiensis; 6–7, Peratherium africanum; 8,
Herpetotheriidae? gen. et sp. indet.
Figure from Crespo and Goin, 2021

3. True metatherians

4. Distribution
Peratherium africanum
Kasserinotherium tunisiense
Herpetotheriidae? gen. et sp. indet.
Figure from Crespo and Goin, 2021

5. Herpetotheriidae
 Peratherium africanum Simons & Bown, 1984 from Jebel Qatrani Formation (Fayum Depression,
northern Egypt) and Taqah (Sultanate of Oman), early Oligocene.
4 molars and 3 premolars, typical metatherian trait
Figure from Hooker et al. 2008. Left: A, CGM 40236, holotype with canine, P2–M3, roots of P1. B, DPC 3820, P3–M4, alveoli of P2. Right: right maxilla with P2–M3, DPC
16946,

6. Herpetotheriidae?
 Genus and species indet. from Moroto (Uganda), early Miocene.
Figures from Crespo and Goin, 2021 and Goin et al., 2016; E–F,
Herpetotheriidae? gen et sp. indet. UM MOR II 48’04, left m4 in
occlusal and buccal views
The number of the cuspids in the talonid allow
us reject the Afrosoricids
A notch between the entoconid and the
hypoconulid is typical from herpetotherids
The herpetotheriids have hypoconulids
and entoconids of unequal height while
in peradectids entoconids and
hypoconulids are of similar height in all
lower molars, including the m4

7. ?Peradectoidea
 Kasserinotherium tunisiense Crochet, 1986 from Kasserine (Tunisia),
early Eocene
Figure from Crespo and Goin, 2021 and Goin et al., 2016; G, EY 10,
holotype, a left M3 in occlusal view; J, EY 12, right M1 (inverted drawing) in
occlusal view, after Crochet (1986, figs. 1, 2).
Presence of a line of stylar
cusps and a large stylar area
Absence of a hypocone and of a cingulum
below the protocone
Presence of a rectilinear centrocrista with a
predilambdodont morphology.

8. Other materials once
referred to the
Metatheria

9. Zhelestidae
 Zhelestidae indet. from Locality MAD93-35, Mahajanga Basin,
Northwestern Madagascar, Upper Cretaceous
Figure from Krause, 2001 and Gheerbrant and Astibia, 1994. Left: Lower-left molar (UA 8699) of the Zhelestidae indet.: a, buccal; b, mesial; c, lingual;
d, distal; and e, f, occlusal (stereopair). Scale bar, 1 mm. Right Lainodon orueetxebarriai 1-3, L1AT 14, holotype, M/1
Their belonging to the Metatherians is ruled out
because of their close resemblance to
contemporary Zelestids.
Lainodon orueetxebarriai

10. Adapisoriculidae
 Garatherium mahboubii Crochet, 1984 from El Kohol (Algeria), late early Eocene
Note the strong similarity of Garatherium mahboubii to
other African Palaeogene adapisoriculids.
Figures from Crespo and Goin, 2021 and Seiffert, 2010. Right Garatherium mahboubii, right M2. Right: Comparison of occlusal morphology of the upper second molar
(scaled to the same approximate buccolingual width) in a series of Paleogene African mammals with possible afrosoricid affinities. A. Widanelfarasia bowni (latest Eocene,
Jebel Qatrani Formation, Egypt). B. Dilambdogale gheerbranti (earliest late Eocene, Birket Qarun Formation, Egypt). C. Chambilestes foussanensis (late early or early
middle Eocene,Chambi, Tunisia). D. ?Garatherium todrae (late Paleocene, Jebel Guersif Formation, Morocco). E. Afrodon chleuhi (late Paleocene, Jbel Guersif Formation,
Morocco). F. Todralestes variabilis (late Paleocene, Jbel Guersif Formation, Morocco).

11. ?Eulipotyphla
 Ghamidtherium dimaiensis Sánchez-Villagra et al., 2007 from BQ-2 Quarry,
Fayum Depression (Egypt), late Eocene.
The dental morphology is typical of Eulipotyphla and Chiroptera, but
the rounded shape of the mandible is more typical of Eulipotyphla.
Figure from Sánchez-Villagra et al., 2007. Ghamidtherium dimaiensis . CGM 83699, right dentary fragment including m1-2 and alveoli
for p4 and m3.Occlusal/buccal view, scale = 2 ram; C. Buccal view, scale = 3 mm; D. Lingual view.

12. Chiroptera
 Chiroptera indet. from BQ-2 Quarry, Fayum Depression (Egypt), late Eocene.
The configuration of the talonid is typical of some
primitive bats (necromantodonty sensu Maitre, 2014),
therefore it can be ruled out that it belongs to a
marsupial.
Figure from Sánchez-Villagra et al., 2007. DPC 21498B, right m1 in occlusal view

13. Chiroptera
 ?Khonsunycteris or ?Phasmatonycteris or new genus and species from BQ-2
Quarry, Fayum Depression (Egypt), late Eocene.
This upper molars have several features that characterize
chiropteran species of the Vespertilionidae and
Myzopodidae. These are include the shape of the
protocone, absence of talon, well-developed mesial and
posterior cinguli, M2 having a small hypocone, among
other features. Notwithstanding, these materials also have
unique features, as the connection of the preprotocrista
with the paracone, which is disconnected to the
paracingulum. This combination of features does not
appear in other vespertilionid or myzopodid bats from the
Eocene. In the same site was described the myzopodid
bat Phasmatonycteris phiomensis and the nearby quarry
L-41 from latest Eocene, there was described the
vespertilionid Khonsunycteris aegypticus both taxa
described from a nearly complete mandible.
Figure from Sánchez-Villagra et al., 2007. Left: DPC 21372A, left M2. -
Right: DPC 21372B right M1.

14. Paleobiogeography

15. The origin of the African metatherians
By contrast, Kasserinotherium,
suggests a Paleocene arrival from
South America, because of the
similarities it has with South
American and Australian taxa.
This last migration probably took
place through a filter corridor such
as the Rio Grande Rise-Walvis
Ridge system, and it may also
explain the enigmatic presence of
polydolopimorphian metatherians
in the Paleogene of Turkey.
The arrival of the Herpetotheriids in
Afro-Arabia was probably through
Europe in one or more dispersal
waves since the early Oligocene.
Figure from Crespo and Goin, 2021. Position of the southern continents at 60 Ma. To the resulting palaeomap it added the contours of the Rio Grande Rise (RGR) and Walvis
Ridge (WR) in the South Atlantic.

16. The origin of the African metatherians
Another more radical hypothesis is
that all European Marsupialiformes
have an origin in South America,
and dispersed via Africa by the
Paleocene–earliest Eocene.
Figure from Crespo and Goin, 2021
Figure from Crespo and Goin, 2021. Position of the southern continents at 60 Ma. To the resulting palaeomap it added the contours of the Rio Grande Rise (RGR) and Walvis
Ridge (WR) in the South Atlantic.

17. Bibliography
 Crespo, V.D., & Goin, F.J. (2021). African Cenozoic metatherians. Spanish Journal of Palaeontology 36 (2),
2021
 Gheerbrant, E., & Astibia, H. (1994). Un nouveau mammifère du Maastrichtien de Laño (Pays Basque
espagnol). Comptes rendus de l'Académie des sciences. Série 2. Sciences de la terre et des planètes, 318(8),
1125-1131.
 Goin, F., Woodburne, M., Zimicz, A. N., Martin, G. M., & Chornogubsky, L. (2016). A brief history of South
American metatherians: evolutionary contexts and intercontinental dispersals. Springer.
 Hooker, J. J., Sánchez‐Villagra, M. R., Goin, F. J., Simons, E. L., Attia, Y., & Seiffert, E. R. (2008). The origin
of Afro‐Arabian ‘didelphimorph’ marsupials. Palaeontology, 51(3), 635-648.
 Krause, D. W. (2001). Fossil molar from a Madagascan marsupial. Nature, 412(6846), 497-498.
 Maitre, E. (2014). Western European middle Eocene to early Oligocene Chiroptera: systematics, phylogeny
and palaeoecology based on new material from the Quercy (France). Swiss journal of Palaeontology, 133(2),
141-242.
 Sánchez-Villagra, M. R., Seiffert, E. R., Martin, T., Simons, E. L., Gunnell, G. F., & Attia, Y. (2007).
Enigmatic new mammals from the late Eocene of Egypt. Paläontologische Zeitschrift, 81(4), 406-415.
 Seiffert, E. R. (2010). The oldest and youngest records of afrosoricid placentals from the Fayum Depression
of northern Egypt. Acta Palaeontologica Polonica, 55(4), 599-616.

18. Thank you for your
attention!