Proboscidea from Kanapoi, Kenya.

The early Pliocene site of Kanapoi (Turkana Basin, Kenya) has a large, diverse vertebrate sample that contains the earliest representatives of the hominin genus Australopithecus. Included in this sample is an impressive assemblage of fossil proboscideans, comprised of deinotheres (Deinotherium bozasi), anancine gomphotheres (Anancus ultimus), and at least three species of elephant (Loxodonta adaurora, a primitive morph of Loxodonta exoptata, and Elephas ekorensis). A single specimen from high in the sequence could plausibly belong to a primitive stage of Elephas recki. A review of dental carbon isotope analyses indicates a range of dietary habits for these taxa, from dedicated browsing (deinotheres) to mixed feeding/grazing (elephants and gomphotheres), which in early Pliocene elephants corresponds to molars with greater crown height and more plates than in late Miocene confamilials, bringing their morphology more in phase with feeding behavior than was the case in their earlier relatives. Variation in feeding preferences among Kanapoi proboscideans corresponds to evidence for habitat heterogeny, including inferred substantial presence of grasses; the occurrence of multiple megaherbivores may have contributed to the fragmentation of ecosystems, positively affecting early hominin success and aiding diversification of other ungulate groups.

A grazing Gomphotherium in Middle Miocene Central Asia, 10 million years prior to the origin of the Elephantidae.

Feeding preference of fossil herbivorous mammals, concerning the coevolution of mammalian and floral ecosystems, has become of key research interest. In this paper, phytoliths in dental calculus from two gomphotheriid proboscideans of the middle Miocene Junggar Basin, Central Asia, have been identified, suggesting that Gomphotherium connexum was a mixed feeder, while the phytoliths from G. steinheimense indicates grazing preference. This is the earliest-known proboscidean with a predominantly grazing habit. These results are further confirmed by microwear and isotope analyses. Pollen record reveals an open steppic environment with few trees, indicating an early aridity phase in the Asian interior during the Mid-Miocene Climate Optimum, which might urge a diet remodeling of G. steinheimense. Morphological and cladistic analyses show that G. steinheimense comprises the sister taxon of tetralophodont gomphotheres, which were believed to be the general ancestral stock of derived "true elephantids"; whereas G. connexum represents a more conservative lineage in both feeding behavior and tooth morphology, which subsequently became completely extinct. Therefore, grazing by G. steinheimense may have acted as a behavior preadaptive for aridity, and allowing its lineage evolving new morphological features for surviving later in time. This study displays an interesting example of behavioral adaptation prior to morphological modification.

Dietary traits of the ungulates from the HWK EE site at Olduvai Gorge (Tanzania): Diachronic changes and seasonality.

The Oldowan site HWK EE (Olduvai Gorge, Tanzania) has yielded a large fossil and stone tool assemblage at the transition from Lower to Middle Bed II, ∼1.7 Ma. Integrated tooth wear and stable isotope analyses were performed on the three most abundant ungulate taxa from HWK EE, namely Alcelaphini, cf. Antidorcas recki (Antilopini) and Equus oldowayensis (Equini), to infer dietary traits in each taxon. Some paleodietary changes were observed for cf. A. recki and E. oldowayensis based on tooth wear at the transition from the Lemuta to the Lower Augitic Sandstone (LAS) interval within the HWK EE sequence. Stable carbon and oxygen isotope data show no significant changes in bulk diet or hydroclimate between the Lemuta and LAS intervals. The combined tooth wear and stable isotope data suggest similar paleoecological conditions across the two HWK EE intervals, but that differences in vegetation consumed among ungulates may have resulted in changes in dietary niches. Integrating tooth wear and stable isotope analyses permits the characterization of ungulate diets and habitats at HWK EE where C4 dominated and minor mixed C3 and C4 habitats were present. Our results provide a better understanding of the paleoenvironmental conditions of the Lemuta and LAS intervals. The LAS assemblage was mostly accumulated during relatively dry periods at Olduvai Gorge when grasses were not as readily available and grazing animals may have been more nutritionally-stressed than during the formation of the Lemuta assemblage. This helps to contextualize variations in hominin and carnivore feeding behavior observed from the faunal assemblages produced during the two main occupations of the site.

The ear region of earliest known elephant relatives: new light on the ancestral morphotype of proboscideans and afrotherians.

One of the last major clades of placental mammals recognized was the Afrotheria, which comprises all main endemic African mammals. This group includes the ungulate-like paenungulates, and among them the elephant order Proboscidea. Among afrotherians, the petrosal anatomy remains especially poorly known in Proboscidea. We provide here the first comparative CT scan study of the ear region of the two earliest known proboscideans (and paenungulates), Eritherium and Phosphatherium, from the mid Palaeocene and early Eocene of Morocco. It is helpful to characterize the ancestral morphotype of Proboscidea to understand petrosal evolution within proboscideans and afrotherians. The petrosal structure of these two taxa shows several differences. Eritherium is more primitive than Phosphatherium and closer to the basal paenungulate Ocepeia in several traits (inflated tegmen tympani, very deep fossa subarcuata and ossified canal for ramus superior of stapedial artery). Phosphatherium, however, retains plesiomorphies such as a true crus commune secundaria. A cladistic analysis of petrosal traits of Eritherium and Phosphatherium among Proboscidea results in a single tree with a low level of homoplasy in which Eritherium, Phosphatherium and Numidotherium are basal. This contrasts with previous phylogenetic studies showing homoplasy in petrosal evolution among Tethytheria. It suggests that evolutionary modalities of petrosal characters differ with the taxonomic level among Afrotheria: noticeable convergences occurred among the paenungulate orders, whereas little homoplasy seems to have occurred at intra-ordinal level in orders such as Proboscidea. Most petrosal features of both Eritherium and Phosphatherium are primitive. The ancestral petrosal morphotype of Proboscidea was not specialized but was close to the generalized condition of paenungulates, afrotherians, and even eutherians. This is consistent with cranial and dental characters of Eritherium, suggesting that the ancestral morphotypes of the different paenungulate orders were close to each other. Specializations occurred rapidly after the ordinal radiation of Paenungulata.

The first record of Deinotherium (Proboscidea, Mammalia) in the Miocene of Adygea.

The data on an isolated upper tooth (P4) of Deinotherium sp. from the Late Miocene beds of the Maikop 1 locality (Maikop, Republic of Adygea) are reported. This is the first record of Deinotherium from the Upper Miocene of Russia. The tooth crown of P4 is similar in size to D. proavum Eichwald, 1831 (= D. gigantissimum Stefanescu, 1892).

The role of behaviour in adaptive morphological evolution of African proboscideans.

The fossil record richly illustrates the origin of morphological adaptation through time. However, our understanding of the selective forces responsible in a given case, and the role of behaviour in the process, is hindered by assumptions of synchrony between environmental change, behavioural innovation and morphological response. Here I show, from independent proxy data through a 20-million-year sequence of fossil proboscideans in East Africa, that changes in environment, diet and morphology are often significantly offset chronologically, allowing dissection of the roles of behaviour and different selective drivers. These findings point the way to hypothesis-driven testing of the interplay between habitat change, behaviour and morphological adaptation with the use of independent proxies in the fossil record.

Diversity in the later Paleogene proboscidean radiation: a small barytheriid from the Oligocene of Dhofar Governorate, Sultanate of Oman.

Despite significant recent improvements to our understanding of the early evolution of the Order Proboscidea (elephants and their extinct relatives), geographic sampling of the group's Paleogene fossil record remains strongly biased, with the first ~30 million years of proboscidean evolution documented solely in near-coastal deposits of northern Africa. The considerable morphological disparity that is observable among the late Eocene and early Oligocene proboscideans of northern Africa suggests that other, as yet unsampled, parts of Afro-Arabia might have served as important centers for the early diversification of major proboscidean clades. Here we describe the oldest taxonomically diagnostic remains of a fossil proboscidean from the Arabian Peninsula, a partial mandible of Omanitherium dhofarensis (new genus and species), from near the base of the early Oligocene Shizar Member of the Ashawq Formation, in the Dhofar Governorate of the Sultanate of Oman. The molars and premolars of Omanitherium are morphologically intermediate between those of Arcanotherium and Barytherium from northern Africa, but its specialized lower incisors are unlike those of other known Paleogene proboscideans in being greatly enlarged, high-crowned, conical, and tusk-like. Omanitherium is consistently placed close to late Eocene Barytherium in our phylogenetic analyses, and we place the new genus in the Family Barytheriidae. Some features of Omanitherium, such as tusk-like lower second incisors, the possible loss of the lower central incisors, an enlarged anterior mental foramen, and inferred elongate mandibular symphysis and diminutive P(2), suggest a possible phylogenetic link with Deinotheriidae, an extinct family of proboscideans whose origins have long been mysterious.

Morphological variation in the ear region of pleistocene elephantimorpha (Mammalia, Proboscidea) from central Texas.

A large sample of isolated elephantimorph petrosal bones was recovered from Pleistocene deposits in Friesenhahn Cave, Bexar County, Texas. Morphology of the middle and inner ear of the elephantimorphs is described and variation within the sample is identified. Observed variations occur in the stapedial ratio, morphology of the aquaeductus Fallopii, and connection of the crista interfenestralis to the tympanohyal on the posterior portion of the petrosal to form a foramen for passage of the stapedius muscle. The morphology of the aquaeductus Fallopii supports an ontogenetic explanation for some variation, and a sequence of ossification surrounding the aquaeductus Fallopii, from the anterior end of the canal to the posterior, is hypothesized. The stapedial ratio varies to a high degree across the sample, and such variation should be considered when the ratio is used in phylogenetic analyses. Within the inner ear, the absence of the secondary lamina suggests evolution of low-frequency hearing in extinct proboscideans, which is known for extant elephants. The morphology of the petrosals from Friesenhahn Cave is compared to published descriptions of the ear regions of other extinct proboscideans, and the distribution and evolution of morphologic characters are discussed. J. Morphol., 2011. © 2011 Wiley-Liss, Inc.