Article

Pecora Incertae Sedis

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Abstract

The Ruminantia is commonly divided into two infraorders: the Tragulina and the Pecora. Pecoran monophyly is well accepted with five modern families: the Giraffidae, Bovidae, Moschidae (musk deer in Asia), Antilocapridae (pronghorns of North America), and Cervidae (deer). These are commonly placed in three superfamilies: Bovoidea, Cervoidea, and Giraffoidea. Of modern pecorans, only the Bovidae and Giraffidae are widespread in Africa (cervids dispersed into North Africa in the early late Pleistocene. In the African Miocene, there are several named taxa of pecoran ruminants that are not easily assigned to the modern families or superfamilies. Morphological analyses have often suggested a close relationship between the Bovidae and Giraffidae, and the position of the Moschidae has been particularly controversial. There is a growing consensus from molecular data and new “supertree” analyses that Bovidae and Cervidae together are the sister group of the Giraffidae. In addition to the living families of ruminants, there are numerous names given to extinct alleged families of both the infraorders Tragulina and Pecora, largely from Eurasia. This chapter describes the systematic paleontology of Pecora incertae sedis. The Ruminantia is commonly divided into two infraorders: the Tragulina and the Pecora. Pecoran monophyly is well accepted with five modern families: the Giraffidae, Bovidae, Moschidae (musk deer in Asia), Antilocapridae (pronghorns of North America), and Cervidae (deer). These are commonly placed in three superfamilies: Bovoidea, Cervoidea, and Giraffoidea. Of modern pecorans, only the Bovidae and Giraffidae are widespread in Africa (cervids dispersed into North Africa in the early late Pleistocene. In the African Miocene, there are several named taxa of pecoran ruminants that are not easily assigned to the modern families or superfamilies. Morphological analyses have often suggested a close relationship between the Bovidae and Giraffidae, and the position of the Moschidae has been particularly controversial. There is a growing consensus from molecular data and new “supertree” analyses that Bovidae and Cervidae together are the sister group of the Giraffidae. In addition to the living families of ruminants, there are numerous names given to extinct alleged families of both the infraorders Tragulina and Pecora, largely from Eurasia. This chapter describes the systematic paleontology of Pecora incertae sedis.

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... Starting with Lydekker [25], which erected the family name Palaeomerycidae, some authors diagnosed palaeomerycids on the basis of dental features and included within the group hornless forms such as Amphitragulus or Oriomeryx, which in turn were considered moschids in some other publications and some of them finally turned out to be basal pecorans [26]. On the other hand, some authors diagnosed palaeomerycids by the presence of ossicones and a single occipital appendage, and considered a more restricted group [4,7,13,19,[27][28][29]. Prolibytherium was sometimes considered part of the Palaeomerycidae despite its tremendous differences with the three-horned true palaeomerycids [10,13], however later works have assigned this taxon to Giraffoidea and Climacoceratidae [26,30]. As noted by Rössner [6] the suite of dental characters used by Janis and Scott [27] to diagnose the family Palaeomerycidae is present in other taxa such as cervids or moschids, and the only real autapomorphic features of palaeomerycids recognized so far are the presence of both a bifurcated occipital appendage and a pair of supra-orbital ossicones. ...
... The Eurasian palaeomerycids share a common ancestor with the Miocene African pecoran Propalaeoryx. Remains of this genus have been found in both South and East Africa [11,30,48,72]. It was accepted that Propalaeoryx was a member of the Giraffoidea [11,27,30,48] with some authors regarding it as a climacoceratid within giraffoids [11,48], an exclusively African family that contains forms such as Orangemeryx, included in this work. ...
... Remains of this genus have been found in both South and East Africa [11,30,48,72]. It was accepted that Propalaeoryx was a member of the Giraffoidea [11,27,30,48] with some authors regarding it as a climacoceratid within giraffoids [11,48], an exclusively African family that contains forms such as Orangemeryx, included in this work. Propalaeoryx appears with a high support as the closest sister-group to the Palaeomerycidae (PP = 0.91), and hence we reject the hypothesis of Propalaeoryx belonging to both the Climacoceratidae and the Giraffoidea. ...
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Palaeomerycids were strange three-horned Eurasian Miocene ruminants known through fossils from Spain to China. We here study their systematics, offering the first cladistic phylogeny of the best-known species of the group, and also reassess their phylogenetic position among ruminants, which is currently disputed. The beautifully preserved remains of a new palaeomerycid from middle Miocene deposits of Spain, Xenokeryx amidalae gen. et sp. nov., helps us to better understand palaeomerycid anatomy, especially that of the nuchal region in the skull, significantly improving our current knowledge on these enigmatic ruminants. Our results show two main lineages of palaeomerycids, one containing the genus Ampelomeryx diagnosed by a characteristic type of cranium / cranial appendages and some dental derived traits, and another one that clusters those forms more closely related to Triceromeryx than to Ampelomeryx, characterized by a more derived dentition and a set of apomorphic cranial features. Xenokeryx branches as a basal offshoot of this clade. Also, we find that Eurasian palaeomerycids are not closely related to North American dromomerycids, thus rejecting the currently more accepted view of palaeomerycids as the Eurasian part of the dromomerycid lineage. Instead of this, palaeomerycids are nested with the African Miocene pecoran Propalaeoryx and with giraffoids. On the other hand, dromomerycids are closely related to cervids. We define a clade Giraffomorpha that includes palaeomerycids and giraffids, and propose an emended diagnosis of the Palaeomerycidae based on cranial and postcranial characters, including several features of the cranium not described so far. We also define the Palaeomerycidae as the least inclusive clade of pecorans containing Triceromeryx and Ampelomeryx. Finally, we reassess the taxonomy of several palaeomerycid taxa.
... c Lists other Miocene sites where the taxon is found. Data from published faunal lists (Pickford, 1986a;Drake et al., 1988;Pickford and Mein, 2006;Cote, 2008Cote, , 2010Leakey et al., 2011;Grossman et al., 2014) with updates from Werdelin and Sanders (2010). Unpublished data are included for Moroto (S. ...
... Kenyasus has been reported from Kalodirr and Moruorot, and Rusinga is the type locality for K. rusingensis (Pickford, 1986a;Leakey et al., 2011). The pecoran Propalaeoryx is known from Rusinga and Elisabethfeld (Whitworth, 1958;Drake et al., 1988;Cote, 2010), while Canthumeryx has been reported from Kalodirr, Moruorot, and Rusinga, as well as at Jebel Zelten in North Africa (Hamilton, 1978;Drake et al., 1988;Harris et al., 2010). ...
Article
Field expeditions to Bukwa in the late 1960s and early 1970s established that the site had a small but diverse early Miocene fauna, including the catarrhine primate Limnopithecus legetet. Initial potassium-argon radiometric dating indicated that Bukwa was 22 Ma, making it the oldest of the East African early Miocene fossil localities known at the time. In contrast, the fauna collected from Bukwa was similar to other fossil localities in the region that were several million years younger. This discrepancy was never resolved, and due to the paucity of primate remains at the site, little subsequent research took place. We have collected new fossils at Bukwa, reanalyzed the existing fossil collections, and provided new radiometric dating. ⁴⁰Ar/³⁹Ar incremental heating ages on lavas bracketing the site indicate that the Bukwa fossils were deposited ∼19 Ma, roughly 3 Ma younger than the original radiometric age. Our radiometric dating results are corroborated by a thorough reanalysis of the faunal assemblage. Bukwa shares taxa with both stratigraphically older localities (Tinderet, Napak) and with stratigraphically younger localities (Kisingiri, Turkana Basin) perfectly corresponding to our revised radiometric age. This revised age for Bukwa is important because it indicates that significant faunal turnover may have occurred in East Africa between 20 and 19 Ma. Bukwa samples immigrant taxa such as large suids, large ruminants, and ochotonids that are absent from stratigraphically older but well-sampled localities in the region, such as Tinderet (∼20 Ma) and Napak (20 Ma). Further age refinements for Bukwa and the entire East African early Miocene sequence will help to constrain the timing of this faunal turnover event, of particular importance in paleoanthropology since this temporal sequence also provides us with what is currently our best window into the early evolution of cercopithecoid and hominoid primates.
... The monophyly of the infraorder Pecora is well accepted and includes the following modern families: Giraffidae, Bovidae, Moschidae, Antilocapridae, and Cervidae; however, the African Early Miocene fossil record includes several taxa that cannot be clearly attributed to one of the modern families (Cote 2010). Morphological analyses have suggested a relationship between the Bovidae and Giraffidae (Morales et al. 1986;Gentry 2000), as well as a relationship between the Giraffidae and Antilocapridae (Hassanin and Douzery 2003). ...
... The upper molars are brachydont with large metaconules on M1 and M2 (Barry et al., 2005). Cote (2008Cote ( , 2010 notes the presence of "Gelocus" whitworthi in the Songhor collections but I was unable to find the 6 specimens she refers to in her dissertation. ...
... Arambourg (1933) and Whitworth (1958) described its dentition as cervoid, but some other authors considered Propalaeoryx to be a giraffoid (Janis & Scott, 1987;Gentry, 1994) or more specifically a climacoceratid giraffoid (Morales et al., 1999. Subsequently, Cote (2010) classified Propalaeoryx as a pecoran incertae sedis, although she admitted that the giraffoid hypothesis was probably the most likely to be valid. However, none of these authors included Propalaeoryx in a phylogenetic analysis. ...
... The first ruminant diversification event occurred in Asia during the middle Eocene (Archaeomerycidae and Lophiomerycidae) and then in North America (Leptomerycidae and Hypertragulidae) (Webb, 1998;Métais and Vislobokova, 2007). In contrast, recent studies confirm that ruminants reached Western Europe during the Oligocene Grande-Coupure dispersal event (Mennecart, 2012;Mennecart and Métais, 2015) and Africa during the early Miocene ( Van der Made, 1999;Cote, 2010). Only scarce remains of Eocene large mammals are known from Eastern Europe and Western Asia, in contrast to the very abundant fossil record from Central Asia (mainly China and Mongolia) and Western Europe. ...
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A fragmented right branch of a ruminant mandible from Rusce (Pčinja basin, Serbia) was originally published in the first half the twentieth century as Micromeryx flourensianus, a small ruminant common in the middle Miocene of Europe. Based on this determination, sedimentary filling of the Pčinja basin was considered to be of late Miocene age. However, later paleobotanical and micromammalian studies pointed to a late Eocene age for these deposits. The redescription and discussion of the ruminant fossil mandible from Rusce led to the conclusion that the specimen may belong to a small species of Bachitheriidae, probably to Bachitherium thraciensis. This ruminant was originally only known from late Eocene strata in Bulgaria. The peculiar late Eocene faunal composition from the Balkans (e.g., rodents, perissodactyls, and ruminants) confirms that the “Balkanian High” was a distinct paleobiogeographical province from that of Western Europe until the Bachitherium dispersal event, which occurred during the early Oligocene ca. 31 Mya.
... The relationships among these stem Pecora and modern pecoran families remain obscure. These forms are generally poorly known and distributed across Eurasia during the late Oligocene-early Miocene, and later in Africa during the early Miocene (Cote, 2010). The best records of these stem Pecora come from Western Europe and Central Asia, but very few data are currently available from Southern Asia. ...
... Compared to Climacoceras, UCMP 40461 has shorter, more robust tines and a shorter beam. In O. hendeyi the tines are found at the distal terminus of the beam, very different from the arrange-which may belong in the Giraffoidea, but then again may not (Cote 2010; Harris et al. 2010). The Giraffoidea community of the Kalodirr Member of the Lothidok Formation includes Climacoceras sp., the taxon represented by UCMP 40461, and two giraffids: C. sirtensis and Giraffidae gen. ...
Article
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Excavations at Kalodirr and Moruorot from the Lothidok Formation (ca. 17 mya) in the West Turkana Region of Kenya have yielded several cranial appendages, dentitions and postcranial fossils that can be attributed to either Climacoceratidae or the Giraffidae. An additional unusual and unique fossil, we describe in this paper for the first time, is, in our opinion, a novel stem-giraffoid cranial appen-dage. The Climacoceras from Kalodirr is the oldest representative of the genus, extending Climacoceratidae into the Early Miocene. We establish that by the end of the Early Miocene in Africa the Giraffoidea included at least two families, Climacoceratidae and Giraffidae, distinguished by different types of cranial appendages. Furthermore, the Giraffidae include at least two distinct lineages, represented by distinctive ossicones found at Kalodirr and Moruorot. Thus, we recognize that unlike at older sites where only Canthumeryx is recognized, by the later part of the early Miocene, Giraffoidea communities in East Africa include as many as three genera, and perhaps even four. This pattern of diversity in giraffoid communities persisted into the Middle Miocene and beyond.
... Exceptions are taxa listed in NOW as Giraffoidea (Walangania, "Gelocus" whitworthi, Propalaeoryx, Prolibytherium, Sperrgebietomeryx, Orangemeryx, Namibiomeryx, Canthumeryx), which have been revised as Pecora indet. (Gentry 1994, Cote 2010, and taxa listed in NOW as Moschidae (Amphitragulus, Dremotherium, Pomelomeryx, Friburgomeryx), which were excluded (Sánchez et al. 2010) and compiled as Pecora indet. here. ...
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The omasum of pecoran ruminants (which is absent in tragulids) and shorter gestation periods in non-giraffid crown pecorans (as opposed to giraffids) could represent cases of key innovations that caused disparity in species diversity in extant ruminants. Literature suggests that the different ruminant groups inhabited similar niche spectra at different times, supporting the ‘increased fitness’ interpretation where a key innovation does not mainly open new niches, but allows more efficient use of existing ones. In this respect, we explored data on fossil species diversity of Afro-Eurasian ruminants from the Neogene and Quaternary. Tragulid and giraffid diversity first increased during the Early/Middle Miocene with subsequent declines, whereas bovid and cervid diversity increased distinctively. Our resulting narrative, combining digestive physiology, life history and the fossil record, thus provides an explanation for the sequence of diversity patterns in Old-World ruminants.
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The ecological preferences of ruminant artiodactyls are commonly used to reconstruct the paleoenvironment of Neogene fossil localities throughout Africa. However, comparatively little research has focused on the ecology of ruminant artiodactyls from the Miocene of Africa. Here, we contribute new molar mesowear and hypsodonty data for the ruminant artiodactyls from the early and middle Miocene of Kenya and Uganda. Macroscopic dental characteristics of 608 tragulids, stem pecorans, giraffoids, and bovids dated to between 20 and 13.7 Ma were analyzed. Our hypsodonty results reveal that, whereas tragulids remain brachydont throughout the early and middle Miocene, pecoran ruminants experience an increase in hypsodonty due to the appearance of high-crowned bovids and climacoceratids that migrate into eastern Africa in the middle Miocene. Results from dental mesowear analysis suggest that all tragulids and pecorans were likely browsers, with only one taxon showing mesowear values that overlap with both browsers and mixed feeders in both the upper and lower molars (Canthumeryx sirtensis). None of the taxa analyzed had mesowear scores indicative of a grazing diet. Surprisingly, middle Miocene bovids and climacoceratids, despite possessing gross tooth morphologies adapted to abrasive diets, were largely utilizing a browsing diet. Although the early and middle Miocene habitats of eastern Africa were likely very heterogenous, none of the ruminant artiodactyls present in these habitats is interpreted as having incorporated grasses into their diet in significant quantities.
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The description of new material and the reassessment of specimens previously assigned to Gelocus quercyi lead us to propose a new genus for this species, Mosaicomeryx gen. nov. Moreover, the description of a juvenile skull and other specimens of Prodremotherium elongatum, and comparison with Dremotherium, provide evidence that these two genera are not closely related as previously thought. A phylogenetic analysis based on 40 dental, cranial and postcranial features highlights the misidentification of Gelocus quercyi and suggests that Mosaicomeryx gen. nov. is closely related to Prodremotherium elongatum. Mosaicomeryx quercyi and Prodremotherium elongatum form a monophyletic group of stem Pecora that first appeared in Western Europe by the late Early Oligocene (MP25–26), and Prodremotherium elongatum persisted up to MP28; following this time both taxa appear to be replaced by Dremotherium and Amphitragulus. This time interval covers two major Oligocene faunal and climate changes: Extinction 1 (MP24), associated with regression of the inner European sea, and Extinction 3/Migrations 3 (MP28), associated with Late Oligocene Warming.
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