The highly diverse tropical freshwater fish family Cichlidae is sparsely represented in the fossil record. Here we describe the new cichlid yTugenchromis pickfordi, gen. et sp. nov., from the Upper Miocene (9-10 Ma) of central Kenya. The new taxon possesses a unique combination of characters, including six lateral line foramina on the lacrimal, three lateral line segments, cycloid scales, and a low number of vertebrae (29), dorsal fin spines (13), and dorsal soft rays (9). Its lacrimal morphology and tripartite lateral line suggest an affinity with the present-day Lake Tanganyika tribes Ectodini and Limnochromini, and thus with members of the 'East African Radiation' among the African cichlids. To further elucidate the relationships of yT. pickfordi, we used a comprehensive comparative data set comprising meristic data from all present-day tribes of the 'East African Radiation.' Principal coordinates analyses support links between the fossil and Ectodini C Limnochromini, and additionally with modern Haplochromini. We conclude that yT. pickfordi could be an extinct lineage within the 'most ancient Tanganyika tribes,' or a stem lineage of the 'ancient Tanganyika mouthbrooders.' A direct relationship to the Haplochromini is unlikely because its members do not exhibit the derived characteristics of the lacrimal as seen in yT. pickfordi. Because Lake Tanganyika is located in the western branch of the East African Rift System, yT. pickfordi from the eastern branch supports the 'melting-pot Tanganyika hypothesis,' which posits that the cichlids of modern Lake Tanganyika are derived from riverine lineages that had already diversified prior to the lake formation.
The Miocene epoch was a time of major change in the East African Rift System (EARS) as forest habitats were transformed into grasslands and hominids appeared in the landscape. Here we provide new sedimentological and palynological data on the middle–upper Miocene Ngorora Formation (Tugen Hills, Central Kenya Rift, EARS), together with clay mineral characterizations, mammal finds and a description of the Ngorora fish Lagerstätte. Furthermore, we introduce a revised age of c. 13.3 Ma for the onset of the Ngorora Formation. The older part of the Ngorora Formation (c. 13.3–12 Ma) records low-energy settings of lakes, floodplains and palaeosols, and evidence of analcime indicates that lakes were alkaline. The palynomorph spectrum consists of tree pollen (Juniperus, Podocarpus), Euphorbiaceae pollen (Acalypha, Croton) and herbaceous pollen of Poaceae and Asteraceae, suggestive of wooded grasslands or grassy woodlands. Alkaline lakes, floodplains and palaeosols continue upsection (c. 12–9 Ma), but environmental fluctuations become more dynamic. Paucity of palynomorphs and the presence of an equid may point to progressively drier conditions. A total of about 500 articulated fish fossils were recovered from distinctive layers of almost all sections studied and represent different lineages of the Haplotilapiines (Pseudocrenilabrinae, Cichlidae). Some of the fish kills may be attributable to rapid water acidification and/or asphyxiation by episodic ash falls. Repeated instances of abrupt change in water depth in many sections are more likely to be due to synsedimentary tectonic activity of the Central Kenya Rift than to climatic variation. Overall, the preservation of the Ngorora fish Lagerstätte resulted from the interplay of tectonics, formation of alkaline lakes and explosive volcanism. As records of grasslands that pre-date late Miocene time are rare, our finding of middle Miocene (12–13 Ma) grassy savannah in the Central Kenya Rift is also relevant to models of human evolution in East Africa.
A new genus and species of fossil cichlid fishes of middle Miocene age (12.5 Ma) is described from the Ngorora fish Lagerstätte (Tugen Hills, Kenya) in the East African Rift Valley. Parsimony analysis of morphological characters using published phylogenetic frameworks for extant cichlids combined with the application of a comprehensive best-fit approach based on morphology was employed to place the new fossil taxon in the phylogenetic context of the African cichlids. The data reveal that the fossil specimens can be assigned to the tribe Oreochromini within the haplotilapiines. † Oreochromimos kabchorensis gen. et sp. nov. shows a mosaic set of characters bearing many similarities to the almost pan-African Oreochromis and the East African lake-endemic Alcolapia . As the striking diversity of present-day African cichlids, with 1100 recognised species, has remained largely invisible in the fossil record, the material described here adds significantly to our knowledge of the Miocene diversity of the group. It effectively doubles the age of a fossil calibration point, which has hitherto been used to calibrate divergence times of the East African cichlids in molecular phylogenetic investigations. Furthermore, the comparative dataset derived from extant cichlids presented here will greatly facilitate the classification of fossil cichlids in future studies.
The African Cichlidae Oreochromis (Alcolapia) and Oreochromis amphimelas can survive in extremely alkaline environments and represent the only known modern alkaliphilic cichlid fish found in Africa. The presence of fossil cichlids from the Miocene of central Kenya (Tugen Hills) that are morphologically similar to Oreochromis (Alcolapia) has been noted in previous works, but the conclusions remained tentative. The purpose of this study is to examine newly discovered fossil cichlids from the Tugen Hills and to compare their osteology with that of extant Oreochromis (Alcolapia). This is performed based on a comprehensive collection of comparative material, using microscopy and computed microtomography (μCT). We provide a revised diagnosis for the genus †Rebekkachromis, and revise its systematic relationships by assigning it to the Oreochromini (rather than to the Etiini). Two new species of †Rebekkachromis are described, i.e., †R. valyricus, sp. nov., and †R. vancouveringae, sp. nov., and a morphologically diverse assemblage of cooccurring †Rebekkachromis specimens is documented. Moreover, we found that †Rebekkachromis had three sensory canal pores (instead of four) on the lower arm of the preopercle, a feature that distinguishes both the modern Oreochromis (Alcolapia) and our fossil specimens from almost all other modern African cichlid fish. Our new data indicate that alkaliphile cichlids similar to Oreochromis (Alcolapia) were present in Central Kenya about 10-13 Ma ago and that the ability of African cichlid fishes to thrive in highly alkaline waters had already developed by that time.
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