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New carnivoran material (Creodonta, Carnivora and Incertae sedis) from the Early Miocene of Napak, Uganda
Abstract
Eight species of creodonts and carnivores have previously been reported from the Early Miocene deposits at Napak, Uganda. This paper takes into account unpublished material from old collections made by W.W. Bishop during the 1950s and 1960s, as well as new samples made by the Uganda Palaeontology Expedition between 1985 and 2005. Four carnivoran species are reported from the site for the first time, of which one is a new genus and species, and one species is removed from previous lists, making for a grand total of 12 species for Napak comprising 6 creodonts, 4 fissipeds and two incertae sedis.
... Hooijer (1966) and Guérin and Pickford (2003) dealt with the rhinocerotids from Napak, while Lavocat (1973) and Pickford et al. (2013Pickford et al. ( , 2014) described the rodents. Morales et al. (2001Morales et al. ( , 2007) described new carnivorans from Napak following up on the pioneer work by Savage (1965) and the later studies by Schmidt-Kittler (1987) and Schmidt-Kittler and Heizmann (1991). The ruminants were studied by Pickford (2002) and Sánchez et al. (2015). ...
... The upper teeth from Napak XV are close in dimensions to those in the holotype maxilla (M 19084) and show no clear morphological differences, apart from those mentioned in the discussion concerning the M1, NAP XV 382'08, which has lingual morphology which is somewhat divergent from other specimens from the same locality, and different from the molars from Napak I, in that the separation between the protocone and lingual cingulum is clearly marked. Another interesting difference in our opinion, is the stronger development of the parastyle in the P4 from Napak XV compared with the fossils from Napak I. Morales et al. (2007) described additional material of this species from Napak I including upper and lower deciduous teeth and an unworn molar. In conclusion, the forms from Napak I and Napak XV can be classified in the same species without any difficulty. ...
... This was also the opinion supported by more recent authors such as Morlo et al. (2007) and Werdelin and Peigné (2010). In contrast Morales et al. (2007), Morales and Pickford (2008) and Morales et al. (2010) have used the Hecubides for these first African amphicyonids. Hecubides euryodon is extraordinarily similar in dimensions to Cynelos lemanensis, at least as far as the dentition is concerned, and very few molars fall outside the range of variation known for the latter species in its type locality (SaintGérand-le-Puy, France), as can be observed in figures 5 and 6. ...
The Early Miocene Napak XV locality (ca 20.5 Ma), Uganda, has yielded an interesting assemblage of fossils, including the very well represented amphicyonid Hecubides euryodon. The remarkable find of a nearly complete mandible, unfortunately with poorly preserved dentition, together with new dental remains allow us to obtain a better idea about the morphology and variability of this species. Additionally, we describe a newly discovered mandible of Hecubides euryodon from the Grillental-VI locality (Sperrgebiet, Namibia), which is the most complete and diagnostic Amphicyonidae material found in this area. Comparisons with Cynelos lemanensis from Saint-Gérand-le-Puy (France), the type locality, and with an updated sample of the species of amphicyonids described in Africa leads us to validate the genus Hecubides. Hecubides would be phylogenetically related to the medium and large size species of Amphicyonidae from Africa, most of them now grouped into the genera Afrocyon and Myacyon, both endemic to this continent.
... They also refer one specimen of I. matthewi (KNM-CMF 4060) to I. andrewsi as this specimen conjoins with an I. andrewsi mandible (KNM-CMF 4023). Further material from Napak has been described as Isohyaenodon zadoki (Morales et al., 2007). The third species, I. pilgrimi (figure 26.3), is the only one of Savage's (1965) three eastern African (former) Hyaenodon species whose status has not been questioned. ...
... Perhaps this is due to the fact that it is the only species of Isohyaenodon to include material of both upper and lower dentitions. New material has recently been described from Napak, Uganda (Morales et al., 2007). As I. andrewsi is the type species of the genus, synonymizing this species with Leakitherium hiwegi or Metapterodon kaiseri will require a taxonomic revision of all members of the genus.Morales et al. (2007:74)has noted that the m2 of I. pilgrimi, M. zadoki, and a possible specimen of Leakitherium from Napak (UM NAP I 44'99) all share a " mesial valley on the anterior margin. ...
... Until recently, H. sulzeri was only known from western Europe (Switzerland, France, Germany, and Spain)(Ginsburg, 1999). This taxon has been identified at Arrisdrift, Namibia, from a maxillary fragment and a juvenile canine and mandible (Morales et al., 2003) and has also been reported from Grillental, Namibia (Morales and Pickford, 2005), although the Grillental material was later referred to Megistotherium (Morales and Pickford, 2008).Morlo et al. (2007)suggest that all described African species of Hyainailouros may ultimately be recognized as conspecifics of the type species, given how poorly known this genus is within Africa. They suggest that all known taxa may be reducible to three: H. bugtiensis (= Megistotherium osteothlastes), H. sulzeri (all African and European specimens), and Sivapterodon lahirri (India). ...
The order Creodonta was first named by Cope (1875) and removed from its original placement in the order Carnivora. Some researchers still maintain that creodonts and carnivorans (or carnivoramorphans) are sister taxa, though not all agree. While creodonts and carnivorans share an ossified tentorium and a few basicranial and tarsal features, their possible synapomorphies are few. Like carnivorans, creodonts vary greatly in their postcranial adaptations. Unlike carnivorans, most creodonts have fissured terminal phalanges, with the exception of the European Proviverrinae, and an unfused scaphoid and lunate. All known Afro-Arabian creodonts belong to the family Hyaenodontidae and range in age from the Eocene (and possibly Paleocene) to the middle Miocene. While the earliest African members of this family are found in the north, this family eventually spread southward into eastern and southern Africa. In Africa, as elsewhere, this family is diverse both in body size and morphology, with body size ranging from some of the smallest of forms to the largest hyaenodontid known (Megistotherium). This chapter describes the systematic paleontology of Creodonta.
... The genus Leakitherium is only known from two maxillary fragments that display P 4 , M 1 and M 2 [3]. Morales et al. [41] referred a fragment of an M 2 to the genus. The taxonomic status of Leakitherium among hyainailourines is unresolved [41]. ...
... Morales et al. [41] referred a fragment of an M 2 to the genus. The taxonomic status of Leakitherium among hyainailourines is unresolved [41]. They hypothesized that Leakitherium is a synonym for Isohyaenodon andrewsi. ...
... They hypothesized that Leakitherium is a synonym for Isohyaenodon andrewsi. In this analysis Leakitherium is closely related to Miocene hyainailourines such as Megistotherium and Isohyaenodon, but the hypothesis of Morales et al. [41] is not supported here. ...
We describe a new large-sized species of hypercarnivorous hyainailourine-Kerberos langebadreae gen. & sp. nov.-from the Bartonian (MP16) locality of Montespieu (Tarn, France). These specimens consist of a skull, two hemimandibles and several hind limb elements (fibula, astragalus, calcaneum, metatarsals, and phalanges). Size estimates suggest K. langebadreae may have weighed up to 140 kg, revealing this species as the largest carnivorous mammal in Europe at that time. Besides its very large size, K. langebadreae possesses an interesting combination of primitive and derived features. The distinctive skull morphology of K. langebadreae reflects a powerful bite force. The postcranial elements, which are rarely associated with hyainailourine specimens, indicate an animal capable of a plantigrade stance and adapted for terrestrial locomotion. We performed the first phylogenetic analysis of hyainailourines to determine the systematic position of K. langebadreae and to understand the evolution of the group that includes other massive carnivores. The analysis demonstrates that Hemipsalodon, a North American taxon, is a hyainailourine and is closely related to European Paroxyaena. Based on this analysis we hypothesize the biogeographic history of the Hyainailourinae. The group appeared in Africa with a first migration to Europe during the Bartonian that likely included the ancestors of Kerberos, Paroxyaena and Hemipsalodon, which further dispersed into North America at this time. We propose that the hyainailourines dispersed into Europe also during the Priabonian. These migrants have no ecological equivalent in Europe during these intervals and likely did not conflict with the endemic hyaenodont proviverrines. The discovery of K. langebadreae shows that large body size appears early in the evolution of hyainailourines. Surprisingly, the late Miocene Hyainailouros shares a more recent common ancestor with small-bodied hyainailourines (below 15 kg). Finally, our study supports a close relationship between the Hyainailourinae and Apterodontinae and we propose the new clade: Hyainailouridae.
... Schmidt-Kittler (1987) noted that Savage had emphasized similarities between his Hecubides Savage, 1965 and the European amphi cyonid Cynelos lemanensis (Pomel, 1846), and synonymized Hecubides under the latter generic name. Recently, Morales et al. (2007) proposed resurrecting Hecubides as distinct from Cynelos Jourdan, 1862. However, the diff erences are slight and as the material is limited and Cynelos itself in dire need of revision, we shall not here use the name Hecubides for these taxa. ...
... Finally, it may be noted that although the data are poor at best, there are some features of Bonisicyon n. gen., such as the broad lower molars and well-developed M1 protocone, that may, together with the lack of evidence of a Bonisicyon-like form outside Africa or of a migration of Amphicyonidae into Africa after the early middle Miocene, argue for a derivation of the genus from early Miocene African amphicyonids such as C. euryodon. If this is the case, there may be merit in resurrecting Hecubides for these forms, as suggested on other grounds by Morales et al. (2007). However, this line of evidence will not be truly useful until intermediate forms are recovered from the African late early and middle Miocene. ...
Amphicyonidae are a common element of carnivoran faunas in the early and middle Miocene of Eurasia and North America, but by the Messinian they had become extinct there, except possibly on the Indian subcontinent. In Africa, amphicyonids are poorly known from a scattering of records from the late Oligocene to the late Miocene. In this paper, we describe the last-surviving amphicyonid in Africa, from Messinian-age sediments (dated c. 6.5-5.3 Ma) of Ethiopia (Gona) and Kenya (Lothagam and possibly Lemudong'o). This new taxon shows unique adaptations to hypercarnivory in the lower molars and was small for an amphicyonid, dentally about the size of a coyote, Canis latrans. © Publications Scientifiques du Muséum national d'Histoire naturelle, Paris.
... Our joint work in the Lower Miocene of Eastern Africa focused on the deposits at Napak (Karamoja, Uganda). The Ferae from the Lower Miocene at Napak (Uganda) are highly diverse in size and morphology (Morales et al. 1998b, Morales et al. 2001, 2007, 2016Pickford 2017, 2018b) (Table 2). A summary of the taxa identified can be found in Morales et al. (2016). ...
This article aims to show the interest of the work of Dr Jorge Morales on the understanding of African Ferae and Ruminantia collected in Uganda, Kenya and Namibia within the framework of collaboration with our team. This is not a revision of the systematics of these groups which is beyond the scope of the paper. Eighty taxa of Ferae are identified in the Cainozoic strata, of which thirty-nine are new. Seventeen ruminant taxa are recognised in the same deposits, of which eight are new.
... Hecubides Savage, 1965 was later included in Cynelos (Ginsburg 1980), a decision followed by nearly all researchers (e.g., Schmidt-Kittler 1987; Morales et al. 1998;Pickford et al. 2003;Peigné & Heizmann 2003;Peigné et al. 2006a, b;Morlo et al. 2007;Werdelin & Simpson 2009;Werdelin & Peigné 2010;Leakey et al. 2011;Hunt & Stepleton 2015;Jiangzuo et al. 2018;Adrian et al. 2018). One exception to this is the work of Morales and colleagues (Morales & Pickford 2005;Morales et al. 2007Morales et al. , 2008Morales et al. , 2010Morales et al. , 2016, who emphasized differences between H. euryodon and the type species of Cynelos, C. lemanensis (Morales et al. 2016). However, as Jiangzuo et al. (2018 discussed, the features cited in favor of retaining the name Hecubides (Morales et al. 2016) can be regarded as intrageneric if other species of Cynelos are considered. ...
We describe two large amphicyonid (Amphicyonidae, Carnivora) mandibles from Moghra, Early Miocene, Egypt. One of these represents a new species of Cynelos Jourdan, 1862, which is in the same size range as C. macrodon (Savage, 1965) and C. ginsburgi n. comb., but exhibits a relatively longer m1 paraconid blade. The other is allocated to Amphicyon giganteus (Schinz, 1825). Based on this new material the differences between Cynelos, Amphicyon Lartet in Michelin, 1836, and Afro-cyon Arambourg, 1961 are clarified. We also reassign three (P4, M1, M2) of four isolated and unas-732 GEODIVERSITAS • 2019 • 41 (21) Morlo M. et al.
... Unfortunately, phylogenetic relationships among Miocene hyainailourines have been difficult to unravel given the fragmentary nature of much of the hyainailourine fossil record (Morales et al., 2003(Morales et al., , 2007Lewis and Morlo, 2010). The genus Hyainailouros, named by Biedermann (1863), has served as somewhat of a wastebasket taxon for many large but often fragmentary dental and postcranial remains recovered from early to middle Miocene sites across Eurasia and Africa (Pilgrim, 1932;Ginsburg, 1980;Lewis and Morlo, 2010). ...
Hyainailourine hyaenodonts are among the largest terrestrial carnivorous mammals known. The clade is widely dispersed, found in Eurasia, North America, and Afro-Arabia in the Paleogene and early Neogene. In this study, we describe dental and postcranial material from Simbakubwa kutokaafrika, gen. et sp. nov., the most complete hyainailourine known from sub-Saharan Africa. The material is from a relatively young adult from the early Miocene locality of Meswa Bridge, Kenya. Simbakubwa differs from Hyainailouros in exhibiting lingually oriented molar protocones, gracile metastyles, and buccolingually compressed, shearing canines. Like other large Miocene hyainailourines, Simbakubwa has deep carnassial notches on the molars and tall paracones fused to shorter metacones forming single piercing cusps. A Bayesian phylogenetic analysis recovers Simbakubwa as the sister taxon of a clade of large-bodied Miocene hyainailourines that includes Hyainailouros and Megistotherium. Bayesian ancestral state reconstruction supports an Afro-Arabian origin for Hyainailourinae with subsequent dispersal to Eurasia during the early Miocene. Regression analysis based on carnassial size is applied to Simbakubwa and closely related hyainailourines, recovering a body mass up to 1,500 kg for the new taxon. The evolution and extinction of Hyainailourinae offers important insights for interpreting ecological transitions from Paleogene to Neogene faunas in Afro-Arabia and Eurasia. http://zoobank.org/urn:lsid:zoobank.
... Amphicyonids assigned to Cynelos in Africa are somewhat controversial. Hecubides euryodon was described by Savage (1965) based on a partial maxilla with P3-M2 and several isolated teeth from Napak, Uganda (20-18.5 Ma, Morales et al. 2007). This species is comparable to C. lemanensis in size. ...
Amphicyonidae is one of the most widely distributed families of Carnivora in the Early and Middle
Miocene, yet the fossil record of this family is extremely poor in Eastern Asia. In this study, we report an
unanticipated diversity of Amphicyonidae from the Middle Miocene Halamagai Formation in Ulungur
River area, Xinjiang, Northwestern China. At least five species of very different size: Amphicyon ulungurensis (including Amphicyon cf. ulungurensis), Gobicyon zhegalloi, Cynelos cf. bohemicus, Cynelos aff.
helbingi and cf. Cynelos sp. based on rare teeth and postcranials were found in the Halamagai
Formation. Such diversity of amphicyonids from East Asia is previously unrecorded and indicates the
probable presence of a representative Mid-Miocene amphicyonid guild. The geographic location of
these species supports the existence of a Palaearctic Neogene faunal exchange of amphicyonids
between Europe and East Asia during the early Middle Miocene interval.
This volume honours Professor Jorge Morales – one of the world´s leading experts in fossil carnivorans, a great scholar, and inspiration of many to follow him in the discipline of palaeontology – to mark his recent retirement. In July 2021, Prof. Morales retired after 50 years working and building up a reputation not only as a leading scientist but also as a teacher, facilitator of science, and mentor. This festschrift celebrates his lifelong, outstanding scientific and cultural achievements to mammal palaeontology. It contains contributions by colleagues, former students, and friends that come from far and wide. Articles pay homage to the jubilarian´s research interests, covering the systematics, palaeoecology, biochronology, palaeobiogeography, taphonomy, and geoheritage. On the basis of the written acknowledgements, it is easy to recognise the affection and deep esteem Prof. Morales induces in everyone. If we think about him, our professional and
personal lives would not be the same without his support, mentorship, friendship, and good humour. It is a proud privilege to thank him for his career in terms of production and excellence which without doubt placed Spanish palaeontology among the global elite. Prof. Morales is retired, but he remains active in research, and as inspiration to all of us.
Early Miocene carnivorous mammals from Buluk, Kenya, are described and discussed. Four taxa belonging to Hyaenodonta and four belonging to Carnivora are identified. Members of Hyaenodonta include Hyainailouros sulzeri, Hyainailouros cf. napakensis, a third taxon about the size of Leakitherium, represented only by postcranial material, and a fourth taxon represented by an edentulous jaw, in the size range of Sectisodon. Members of Carnivora include a new species of Cynelos jitu, which represents the largest species of Cynelos known. The first m2 of Cynelos macrodon is described, and the differentiation of this species from Cynelos ginsburgi and Cynelos peignei is confirmed. A third carnivoran species is represented by a mandibular fragment attributed to a viverrid similar to Mioprionodon, and a fourth taxon is represented by a feliform distal humerus, the size of that of a small cat. An ecomorphological guild structure analysis reveals that the Buluk carnivore have estimated body sizes spanning from <1 kg to >100 kg. Three very large species (>100 kg), and another two in the 30–100 kg range are present, while only two taxa are present in the 3–10 kg category. Carnivores in the 1–3 kg and the 10–30 kg categories are absent. Locomotor pattern could be obtained for only four taxa, and all are characterized by terrestrial locomotion. A minimum of three dietary classes (insectivorous, carnivorous, scavenging) are represented. The co-occurrence of multiple very large carnivores is not uncommon in early Miocene faunas, but the taphonomy of Buluk may also contribute to the favored preservation of larger and terrestrially adapted animals.
In this beautifully illustrated natural history that links extinct larger feline species with those still in existence, collaborators Alan Turner and Mauricio Anton weave together the evidence of modern feline behavior with that of the fossil record. Turner's clear, insightful prose and Anton's masterly illustrations combine to offer specialists and newcomers alike an accurate and accessible guide to the evolution of cats.
The recovery of a new species of small creodont from the Early Miocene of coastal Namibia including associated upper and lower dental elements reveals that the group is in need of revision. Previously, East African samples assigned to Metapterodon consisted only of maxillae while Isohyaenodon was only represented by lower jaws. It is now clear that the East African Metapterodon specimens represent the upper jaws of Isohyaenodon, while the Namibian material belongs to a separate genus Metapterodon which seems to be absent from East Africa.
Since the late 1950's, many ruminant specimens have been recovered from the Early Miocene volcano-sedimentary succession at Napak, Uganda. Beginning in 1958 when the first fossils were recognised by J.G. Wilson, and the subsequent surveys of Bishop and colleagues, more than 340 specimens have been found. Preliminary publications in the 1960's identified three species, Dorcatherium songhorensis, Dorcatherium ? chappuisi, and Palaeomeryx africanus. Napak material hitherto identified as Dorcatherium ? chappuisi represents a new species of tragulid smaller than D. chappuisi, and is here identified as Dorcatherium iririensis nov. sp., while material previously assigned to Palaeomeryx is transferred to Walangania. A new finding for the site is the tiny tragulid, Dorcatherium parvum. There are thus four species of ruminants at Napak.