The fossil vertebrate locality Kossom Bougoudi,Djurab desert,Chad: A window in the distribution of the carnivoran faunas at the Mio–Pliocene boundary in Africa

The fossil vertebrate bearing locality Kossom Bougoudi (KB) is situated in the Djurab desert (Chad, Africa), 600 km north-east of N’djamena. It has yielded about 1250 specimens with many mammalian remains, principally artiodactyls. Its geologic age has been estimated to be about 5 Ma by biochronologic estimation and about 5.3 Ma by radiometric studies on cosmogenic nuclides of beryllium (authigenic ¹⁰B/⁹B). The carnivoran fauna contains few specimens which belong to five different families. All the taxa were unknown in central Africa. A large lutrine is close to Sivaonyx but different from known species of the genus. Another large lutrine is similar by its size to a species described from the Middle Pliocene of Uganda. An edentulous mandible of a small machairodont cat resembles a small species of Dinofelis, while a distal humerus indicates the presence of a larger member of the same genus. A hunting hyaenid is also much like the European species. An unidentified canid reaches the size of the recent Canis aureus and an isolated calcaneum matches that of the large extant viverrid. This small fauna allows a first look at the guild of the carnivorans at the Latest Miocene–Pliocene boundary in Central Africa and is a milestone between North African, East African and South African carnivore faunas.     

South African fossil Cercopithecoidea: A re-assessment including a description of new material from Makapansgat,Sterkfontein and Taung

The fossil cercopithecoid material from South Africa has been reviewed according to sites and species. The 722 specimens considered comprise 6 genera including 16 taxa and come from 16 sites. Aspects of taxonomic controversy and interest are discussed. In particular, the Parapapio material from Makapansgat has been re-evaluated and the taxonomy of the genus Simopithecus is reconsidered. A number of proposals are put forward. Four new specimens from Makapansgat and one from Sterkfontein are described; a previously partially described specimen from Taung is re-described in detail.     

New species of Cercopithecoides from Haasgat, North West Province, South Africa

Analyses of new cercopithecid fossil specimens from the South African site of Haasgat point to craniofacial affinities with the genus Cercopithecoides. Detailed metric and non-metric comparisons with South African Cercopithecoides williamsi, and other East African Cercopithecoides species, Cercopithecoides kimeui, Cercopithecoides meaveae, Cercopithecoides kerioensis, and Cercopithecoides alemyehui demonstrate that the Haasgat fossils have distinct craniofacial morphology and dental metrics. Specifically, material from Haasgat probably represents one of the smaller Cercopithecoides, differing from the others in its particular suite of features that vary within the genus. It is unique in its more vertical ramus, associated with a relatively lengthened mandibular body. Haasgat Cercopithecoides has a particularly narrow interorbital region between relatively larger ovoid orbits, with articulation of the maxillary bones at a suture above the triangular nasal bones. Furthermore, the maxillary arcade is more rounded than other Cercopithecoides, converging at the M² and M³. The conclusion drawn from this analysis is that the Pleistocene Haasgat fossils are colobines representing a distinct taxon of Cercopithecoides, Cercopithecoides haasgati, thus adding a second species of the genus to southern Africa.     

New observations on the Late Miocene–Early Pliocene Lutrinae (Mustelidae: Carnivora,Mammalia) from the Middle Awash,Afar Rift,Ethiopia

New observations on the Late Miocene and Earliest Pliocene mustelids from the Middle Awash of Ethiopia are presented. The Middle Awash study area samples the last six million years of African vertebrate evolutionary history. Its Latest Miocene (Asa Koma Member of the Adu-Asa Formation, 5.54–5.77 Ma) and Earliest Pliocene (Kuseralee and Gawto Members of the Sagantole Formation, 5.2 and 4.85 Ma, respectively) deposits sample a number of large and small carnivore taxa among which mustelids are numerically abundant. Among the known Late Miocene and Early Pliocene mustelid genera, the Middle Awash Late Miocene documents the earliest Mellivora in eastern Africa and its likely first appearance in Africa, a new species of Plesiogulo, and a species of Vishnuonyx. The latter possibly represents the last appearance of this genus in Africa. Torolutra ougandensis is known from both the Late Miocene and Early Pliocene deposits of the Middle Awash. The genus Sivaonyx is represented by at least two species: S. ekecaman and S. aff. S. soriae. Most of the lutrine genera documented in the Middle Awash Late Miocene/Early Pliocene are also documented in contemporaneous sites of eastern Africa. The new observations presented here show that mustelids were more diverse in the Middle Awash Late Miocene and Early Pliocene than previously documented.     

A new large Pliocene colobine species (Mammalia: Primates) from Asa Issie, Ethiopia

The Colobinae (Mammalia: Primates) are relatively unknown from the African middle to late Miocene. When they appear in the Pliocene they are unambiguous and already fairly diverse taxonomically, geographically, and ecologically. Discoveries from Pliocene sediments in eastern and southern Africa document a radiation of large-bodied colobines very different from those known today. Paleontological research in Ethiopia has recently led to the discovery and identification of another large-bodied colobine species from the early Pliocene site of Asa Issie, discovered in 2000. This new colobine is larger than but morphologically very similar to its sister taxon Kuseracolobus aramisi, an older taxon also described from the Middle Awash. This new species has significant implications for our understanding of the Pliocene colobine adaptive radiation.     

A new genus for a rare African vespertilionid bat: insights from South Sudan

A new genus is proposed for the strikingly patterned African vespertilionid “Glauconycteris” superba Hayman, 1939 on the basis of cranial and external morphological comparisons. A review of the attributes of a newly collected specimen from South Sudan (a new country record) and other museum specimens of “Glauconycteris” superba suggests that “Glauconycteris” superba is markedly distinct ecomorphologically from other species classified in Glauconycteris and is likely the sister taxon to Glauconycteris sensu stricto. The recent capture of this rarely collected but widespread bat highlights the need for continued research in tropical sub-Saharan Africa and in particular, for more work in western South Sudan, which has received very little scientific attention. New country records for Glauconycteris cf. poensis (South Sudan) and Glauconycteris curryae (Gabon) are also reported.     

Gradual evolution in the African hunting dog lineage Systematic implications

The hunting dog, Lycaon pictus, is one of the most representative species of the extant African fauna and one of the most efficient predators in the World. This species is actually a relict within Sub-Saharan Africa, but its lineage is well recorded in Eurasia during the Pliocene, and in Eurasia and Africa during the Early Pleistocene, while its record during the Middle Pleistocene is not well documented. Though postcranial skeletal remains dating from the Early Pleistocene show a tetradactyl forelimb, a characteristic feature of the extant lycaon, unique among canids, the upper and lower dentitions show gradual evolution from a primitive morphology in the Late Pliocene specimens to the highly specialized trenchant carnassials of the extant predatory species. We propose a new systematics for the lineage, grouping all the forms within the genus Lycaon. To cite this article: B. Martínez-Navarro, L. Rook, C. R. Palevol 2 (2003).     

Les Spatangus du Miocène et du Pliocène de l’Ouest de la France

Numerous fragments of spatangoid echinoids have been discovered in the Pliocene deposits of Challans, in Vendée (western France). In spite of the fragmentary data of the samples, a reconstitution of a complete test could be realized using the different fragments and their symetrization. The general shape of the test, and its architectural and ornemental characters allow establishing the presence of the genus Spatangus in western France during the end of Neogene. It allows to precise the biogeography of the genus Spatangus and of the morphological group S. (S.) purpureus on the Atlantic coast after the Messinian crisis. The Pliocene species is compared to the Miocene Spatangus (Phymapatagus) brittanus, abundant in Anjou, Brittany and Touraine. This older species was refered to the subgenus Phymapatagus according to the presumed lack of primary tubercles on its posterior interambulacrum. The discovery of well-preserved specimens, with primary tubercles on every parts of the test, in the Middle Miocene of Brittany allows to refute this subgeneric distinction and to refer the species brittanus to the subgenus Spatangus (Spatangus). The presence of this subgenus in western France is finally confirmed from Middle Miocene to Pliocene.     

The fossil Bovidae (Artiodactyla, Mammalia) from Gesher Benot Ya'aqov, Israel: Out of Africa during the Early-Middle Pleistocene transition

We report the study of the collection of fossil bovid specimens from the Early-Middle Pleistocene Acheulian site of Gesher Benot Ya‘aqov. This locality, situated in the Levantine Corridor (the bottleneck that connects Africa and Eurasia) is a key site to explain the faunal and human dispersals out of Africa during the Matuyama/Brunhes boundary around 0.8 Ma. Two species of bovine (Bos sp., and Bovini gen. et sp. indet. cf. Bison sp.), one antelope (Gazella sp. cf. G. Gazella), and another indeterminate Bovidae gen. et sp. indet., have been recorded. The largest species, Bos sp., is an African immigrant related to the species from the Eritrean site of Buia, Bos buiaensis, which evolved from the buffalo of Olduvai Pelorovis oldowayensis, and colonized the Eurasian continent in parallel with the dispersal of the Acheulian culture into the northern continent. Numerous important species first recorded in several localities of Early-Middle Pleistocene transition from Eurasia are included in this dispersal out of Africa, including the megaherbivore, Palaeoloxodon antiquus, and the carnivores Crocuta crocuta, and later, Panthera leo and Panthera pardus. This faunal turnover is coincident with the change to colder climates that dominated the Middle Pleistocene.     

Taxonomic notes on the genus Otoptera (Phaseoleae, Fabaceae) in southern Africa

A taxonomic study of the genus Otoptera (tribe Phaseoleae) is presented. This distinct genus consists of two species, O. burchellii in southern Africa (Namibia, Botswana, South Africa and Zimbabwe) and O. madagascariensis endemic to Madagascar. The two species differ in the shape and size of leaflets, size and orientation of the spur located at the base of the wing petal, and also by size of the pod. A key to the two species is provided. This study focuses mainly on the southern African O. burchellii DC., which is described and illustrated. The species name is lectotypified and the known geographical distribution is recorded for the first time.     

Integrative taxonomy of a poorly known Sahelian rodent, Gerbillus nancillus (Muridae,Gerbillinae)

Aridification processes that affected the Sahelian area of West Africa during the last decades have induced significant changes in plant and animal communities of this region. In rodents, the genus Gerbillus characteristic of North African and Asian arid habitats has been affected by this climatic and environmental trends. Several species of this genus showed a southward range expansion in recent years into the Sahelian bioclimatic zone. Recent sampling in several localities of West Africa (Mali, Niger and Senegal) enabled us to collect numerous specimens of small gerbils. An integrative study of these samples using molecular, morphological and cytogenetical methods revealed that many of them were attributable to Gerbillus nancillus, a secretive and poorly known species. Gerbillus nancillus appears characterized by a well differentiated karyotype with 2n = 56 chromosomes, and to represent a unique genetic lineage within this genus. Body and skull measurements of G. nancillus were compared with those of the morphologically similar Gerbillus henleyi, which provided diagnostic clues between them. These new data significantly expand the distribution area of G. nancillus, which now ranges from Sudan in the East to Senegal in the West. G. nancillus is here reported from numerous new localities in Niger and Mali, and for the first time in Senegal, which raises questions about the origin of its presence and the colonization routes it followed to get there. We also show that G. henleyi and G. nancillus are sympatric and apparently often syntopic in the sub-Saharan part of the distribution of G. henleyi.     

Speciation slowing down in widespread and long-living tree taxa: insights from the tropical timber tree genus Milicia (Moraceae)

The long generation time and large effective size of widespread forest tree species can result in slow evolutionary rate and incomplete lineage sorting, complicating species delimitation. We addressed this issue with the African timber tree genus Milicia that comprises two morphologically similar and often confounded species: M. excelsa, widespread from West to East Africa, and M. regia, endemic to West Africa. We combined information from nuclear microsatellites (nSSRs), nuclear and plastid DNA sequences, and morphological systematics to identify significant evolutionary units and infer their evolutionary and biogeographical history. We detected five geographically coherent genetic clusters using nSSRs and three levels of genetic differentiation. First, one West African cluster matched perfectly with the morphospecies M. regia that formed a monophyletic clade at both DNA sequences. Second, a West African M. excelsa cluster formed a monophyletic group at plastid DNA and was more related to M. regia than to Central African M. excelsa, but shared many haplotypes with the latter at nuclear DNA. Third, three Central African clusters appeared little differentiated and shared most of their haplotypes. Although gene tree paraphyly could suggest a single species in Milicia following the phylogenetic species concept, the existence of mutual haplotypic exclusivity and nonadmixed genetic clusters in the contact area of the two taxa indicate strong reproductive isolation and, thus, two species following the biological species concept. Molecular dating of the first divergence events showed that speciation in Milicia is ancient (Tertiary), indicating that long-living tree taxa exhibiting genetic speciation may remain similar morphologically.     

The cushion–star Parvulastra exigua in South Africa: one species or more?

The cushion–star Parvulastra exigua (Lamarck, 1816) is a widely distributed member of the temperate intertidal fauna in the southern hemisphere. In South Africa, it occurs in sympatry with the endemic Parvulastra dyscrita (Clark, 1923), the two species being differentiated predominantly by gonopore placement. Several recent studies have suggested that there may be additional cryptic species within the Parvulastra exigua complex in South Africa, based variously on color morphology, genetic evidence and the differential placement of the gonopores. This paper attempts to resolve whether one or more species are represented within Parvulastra exigua. A total of 346 Parvulastra exigua and 8 Parvulastra dyscrita were collected from sites on the west and south–west coasts of South Africa; morphological, anatomical and genetic analyses were performed to determine whether cryptic species and/or Parvulastra exigua specimens with aboral gonopores were present. Results show that neither cryptic species nor Parvulastra exigua specimens with aboral gonopores occur at these sites. This study thus refutes previous claims of the existence of aboral gonopores in South African Parvulastra exigua, and suggests that a single species is represented. The distinction between Parvulastra exigua and Parvulastra dyscrita is also confirmed, and features separating these two species are clarified and documented.     

Comparative pollen morphology of Brachylena, Tarchananthus and two species of Tubulifloridites (Asteraceae) from the Eocene, Knysna Lignite of South Africa

Two fossil taxa Tubulifloridites antipodica and T. viteauensis recovered from the Eocene Knysna Lignite of South Africa were examined with scanning (SEM) and transmission electron microscopy (TEM). The details of their sculpturing and wall structure are similar to the same species of fossil dispersed pollen taxa recovered from southwestern Africa and South America. Fifteen species of the woody South African taxa, Brachylaena (9 species) and Tarchonanthus (6 species) were investigated with SEM and TEM. All of the taxa are tricolporate, spherical to slightly prolate, microechinate to echinate and have a bilayered columellate infrastructure, except B.ilicifolia, which has a single columellate infrastructural level with the “granularization” of the outer portion of the infrastructural layer or the inner layer of the tectum. There is a similar distribution of plesiomorphic and derived pollen characters in a number of aster subfamilies and tribes suggesting a similar evolutionary progression of pollen, and pollen wall character evolution was occurring synchronously in a variety of aster subfamilies during the middle Tertiary and that these unique pollen features may be important to the evolution and diversification of the Asteraceae.     

Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa

Drimolen is one of the newest and most productive hominin sites in South Africa, and is dated on faunal grounds between 2.0 Ma to 1.5 Ma. This paper provides the first overview of the Carnivora from Drimolen, updating the previously published preliminary faunal list, and describing all currently prepared craniodental and postcranial material. The Drimolen specimens are described in comparison with other modern and fossil South African carnivore material. The carnivores cover a range of taxa including hyaenids, felids, canids and herpestids. Most notable amongst these are the sabretooth Dinofelis aff. piveteaui craniodental and postcranial remains, which are described in detail, and a Chasmaporthetes nitidula cranium. The genus Chasmaporthetes is found at three other sites in the area - Sterkfontein, Swartkrans and Coopers D. There are two models for the geographic origin of Dinofelis piveteaui, in that it may have arisen in either eastern or southern Africa. These possibilities are discussed in the light of the new South African Dinofelis material, as the Drimolen material appears to represent a more primitive form with affinities with D. piveteaui. Fossil leopard material from Kromdraai B and Drimolen is also discussed, as the metapodia assigned to P. pardus from these two sites are very small, but lie within the variation of modern leopards. Such size differences in fossil postcrania may have implications for the niches that these animals may have occupied in the past.     

The genus Macroteleia Westwood in Middle Miocene amber from Peru (Hymenoptera,Platygastridae s.l., Scelioninae)

A new species of the scelionine genus Macroteleia Westwood (Platygastridae s.l., Scelioninae) is described and figured from a female beautifully preserved in Middle Miocene amber from Peru. Macroteleia yaguarum Perrichot & Engel, sp. n., shows a unique combination of characters otherwise seen independently within its congeners. It is most similar to the modern M. surfacei Brues, but differs from it by the non-foveolate notauli, the contiguous punctures of the vertex, and the continuous propodeum. The new species is the first New World fossil of the genus, suggesting a Cretaceous origin for the group and a relatively old age of the South American, tropical African, and Australian faunas, and a younger age of the modern Holarctic faunas.     

Plectranthus: A plant for the future?

The genus Plectranthus (Lamiaceae) is a significant, prolific and extensively used genus in southern Africa. It plays a dominant role in both horticulture and traditional medicine. Some 12 species are documented for their use in treating ailments by various indigenous peoples of southern Africa. It is a firm favourite in gardens and Plectranthus has been bred to further utilise the remarkable diversity of indigenous South African wildflowers with amenity horticultural potential. Although previously subjected to both horticultural (Van Jaarsveld, 2006) and ethnobotanical (Lukhoba et al., 2006) review, Plectranthus is a genus with economic potential in various sectors, and this article aims to review this potential of southern African species.     

A reassessment of the taxonomic status and phyletic relationships of Papio baringensis and Papio quadratirostris (Primates: Cercopithecidae)

Comparison of the skulls of Papio baringensis Leakey 1969 from the Chemeron Formation and P. quadratirostris Iwamoto 1982 from the Usno Formation with those of Theropithecus gelada, T. brumpti, T. darti, T. oswaldi, and several species of Papio indicate that the species from Chemeron and Usno exhibit all, or most, respectively, of the shared derived characters of Theropithecus. We propose that they be removed from Papio and placed in Theropithecus as T. baringensis and T. quadratirostris. Comparison of the specimens of T. baringensis and T. quadratirostris with those of T. brumpti indicate that the former two species have some of the derived characters of T. brumpti but are primitive in others. We propose that the three species form the following phyletic lineage: T. baringensis-T. quadratirostris-T. brumpti. With these referrals, there are now six species of the genus Theropithecus. Based on geologic grounds, the specimens from Chemeron are about 4.0 million years (m.y.), that from Usno between 3.3 and 3.4 m.y., and those of T. brumpti between 2.0 and 2.8 m.y. in age. We also show that the most complete specimen of Papio sp. nov. from Olduval Gorge belongs to T. oswaldi. With the removal of these specimens from Papio, the East African fossil record of this genus, apart from isolated teeth, comprises only 21 specimens, while that of Theropithecus comprises at least 300 specimens.     

Hominin diversity in the Middle Pliocene of eastern Africa: the maxilla of KNM-WT 40000

The 3.5-Myr-old hominin cranium KNM-WT 40000 from Lomekwi, west of Lake Turkana, has been assigned to a new hominin genus and species, Kenyanthropus platyops, on the basis of a unique combination of derived facial and primitive neurocranial features. Central to the diagnosis of K. platyops is the morphology of the maxilla, characterized by a flat and relatively orthognathic subnasal region, anteriorly placed zygomatic processes and small molars. To study this morphology in more detail, we compare the maxillae of African Plio-Pleistocene hominin fossils and samples of modern humans, chimpanzees and gorillas, using conventional and geometric morphometric methods. Computed tomography scans and detailed preparation of the KNM-WT 40000 maxilla enable comprehensive assessment of post-mortem changes, so that landmark data characterizing the morphology can be corrected for distortion. Based on a substantially larger comparative sample than previously available, the results of statistical analyses show that KNM-WT 40000 is indeed significantly different from and falls outside the known range of variation of species of Australopithecus and Paranthropus, contemporary Australopithecus afarensis in particular. These results support the attribution of KNM-WT 40000 to a separate species and the notion that hominin taxonomic diversity in Africa extends back well into the Middle Pliocene.     

Why Africa matters: evolution of Old World Salvia (Lamiaceae) in Africa

Background and Aims

Salvia is the largest genus in Lamiaceae and it has recently been found to be non-monophyletic. Molecular data on Old World Salvia are largely lacking. In this study, we present data concerning Salvia in Africa. The focus is on the colonization of the continent, character evolution and the switch of pollination systems in the genus.

Methods

Maximum likelihood and Bayesian inference were used for phylogenetic reconstruction. Analyses were based on two nuclear markers [internal transcribed spacer (ITS) and external transcribed spacer (ETS)] and one plastid marker (rpl32-trnL). Sequence data were generated for 41 of the 62 African taxa (66 %). Mesquite was used to reconstruct ancestral character states for distribution, life form, calyx shape, stamen type and pollination syndrome.

Key Results

Salvia in Africa is non-monophyletic. Each of the five major regions in Africa, except Madagascar, was colonized at least twice, and floristic links between North African, south-west Asian and European species are strongly supported. The large radiation in Sub-Saharan Africa (23 species) can be traced back to dispersal from North Africa via East Africa to the Cape Region. Adaptation to bird pollination in southern Africa and Madagascar reflects parallel evolution.

Conclusions

The phenotypic diversity in African Salvia is associated with repeated introductions to the continent. Many important evolutionary processes, such as colonization, adaptation, parallelism and character transformation, are reflected in this comparatively small group. The data presented in this study can help to understand the evolution of Salvia sensu lato and other large genera.