Tragoportax

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Tragoportax
Temporal range: Turolian–Ruscinian
9.0–5.3  Ma
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Tragoceros amaltheus.jpg
Horns of Tragoportax amalthea
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Bovidae
Subfamily: Bovinae
Tribe:† Tragoportacini
Genus:† Tragoportax
Pilgrim, 1937
Type species
Tragoportax salmontanus
Pilgrim, 1937
Other species

See text

Synonyms
Synonyms
  • Capra amalthea Roth & Wagner, 1854 [1]
  • TragocerusGaudry, 1861 [2] [3]
  • PontoportaxKretzoi, 1941 [3]
  • GazelloportaxKretzoi, 1941 [3]
  •  ?MirabilocerusHadjiev, 1961 [3]
  • Tragoceridus Kretzoi, 1968 [3]
  • MesembriportaxGentry, 1974 [3]
  • MesotragocerusKorotkevich, 1982 [3]

Tragoportax is an extinct genus of bovid ungulate. It lived from the upper Miocene to the earliest Pliocene, and its fossils have been found in southeastern Europe, parts of Africa, and the Indian subcontinent. Tragoportax was formerly considered a close relative of the extant nilgai, though more recent studies suggest that it, and several other Miocene "boselaphins", formed a tribe of their own. The number of Tragoportax species has varied over the years, and some have been reassigned to the related genus Miotragocerus . The first Tragoportax specimens were discovered in Greece, and were originally assigned to Capra . Subsequently, they were reassigned to Tragocerus. This genus name was preoccupied by a beetle, and thus Tragoportax is the correct name.

Contents

Like many modern antelopes, Tragoportax exhibited sexual dimorphism. Both sexes were horned, with those of males being longer and slenderer, often curving backwards. The postcrania of the genus resemble those of large deer, such as red deer, and it may have been similar ecologically. Some species, like T. amaltheae, inhabited lightly wooded environments, and fed predominantly on leaves. Others, like T. rugosifrons, were probably mixed feeders, feeding on both leaves and grasses. Tragoportax might have been cursorial.

Taxonomy

Early history

The first of the specimens currently assigned to Tragoportax, three horn cores with parts of their skulls still attached, were discovered in Pikermi, Greece. In 1854, Johannes Roth and Johann Andreas Wagner assigned the fossils to the goat genus, Capra . The new taxon was named Capra amalthea, after Amalthea, a nymph that nursed the infant Zeus on goat milk in Greek mythology. [1] A few years later, in 1861 Jean Albert Gaudry recognised that the Pikermi fossils were distinct enough from Capra to warrant a genus of their own. He assigned them to the genus Tragocerus; however, Tragocerus was preoccupied by a longhorn beetle. [4] In 1937, Guy Ellcock Pilgrim coined the generic name Tragoportax, to include fossils recovered from the Sivalik Hills in Pakistan. For the time being, the two genera were kept separate, though Pilgrim noted strong similarities between Tragocerus and the new genus. [5] In 1971, Alan William Gentry rendered Tragocerus invalid, and assigned some of its species to Miotragocerus . [6] By 2009, Tragocerus amalthea had been reassigned to Tragoportax, [7] though Tragocerus was apparently regarded as valid by some authorities well into the 1980s. [8]

In 1904, Max Schlosser named a second species of Tragoportax (then Tragocerus), T. rugosifrons, based on material recovered from Samos, Greece. [9] Specimens from Bulgaria, Moldova, Pakistan and Ukraine have been subsequently assigned to T. rugosifrons, as well as a possible specimen from Iran. [3]

Tragoportax curvicornis, also originally found in Samos, was originally assigned to the genus Mirabilocerus, though that genus may be a synonym of Tragocerus. [3] In 1941, Miklós Kretzoi described Gazelloportax and Pontoportax to accommodate fossils now assigned to Tragoportax. [10] In 1974, Gentry named Mesembriportax acrae based on remains from the Early Pliocene of South Africa. It has since been determined that Mesembriportax exhibits all of the cranial characteristics of Tragoportax, and as such, it was reassigned to that genus in 2004. [3] In 1968, Krezoi attempted to revive Tragocerus as a subgenus, Tragoceridus; [11] this was subsequently synonymised. [3]

Classification

Tragoportax has often been assigned to the tribe Boselaphini or subfamily Boselaphinae, alongside the modern nilgai (Boselaphus tragocamelus). [3] [5] However, Bibi et al. (2009) suggested that Boselaphinae as defined was probably non-monophyletic, and that Tragoportax and its relatives should form a separate tribe, Tragoportacini. [7] This tribe would additionally include Austroportax , Kipsigicerus , Miotragocerus, Protragocerus , and Strepsiportax . [7] [12]

Inner systematics

Tragoportax fossils have been recovered from southeastern Europe, Libya, South Africa, and the Indian subcontinent. [3] [13] Most specimens are from the Turolian European Land Mammal Age (equivalent to the Tortonian and Messinian stages) of the Miocene, though there are some specimens that may date to the earliest Pliocene (the Ruscinian, corresponding to the Zanclean and early Piacenzian). The question of how many species are actually valid is a matter of debate: T. rugosifrons alone has five junior synonyms. Denis Geraads and Nikolai Spassov, in 2004, recognised only eight, or nine if T. maius is not a synonym of T. eldaricus. A list of Tragoportax species, per that study, is as follows: [3]

TaxonCountry of originAuthor(s) of taxonTaxon publication year
T. acraeSouth AfricaGentry1974
T. amaltheaGreece Roth & Wagner 1854
T. curvicornisPakistanAndree1926
T. cyrenaicusLibyaThomas1979
T. eldaricusAzerbaijanGabashvili1956
T. macedoniensisGreeceBouvrain1988
?T. maiusGeorgiaMeladze1967
T. rugosifronsBulgaria, Greece, ?Iran, Moldova, Pakistan, & UkraineSchlosser1904
T. salmonatusPakistan Pilgrim 1937

Multiple taxonomic revisions have taken place in the intervening years. T. macedoniensis is treated in some papers as a species of Miotragocerus. [12] In 2009, a new Tragoportax species, T. abyssinicus, was described from the Late Miocene of Afar, Ethiopia. [14] In 2023, another new species, T. perses, was described from Iran. [15]

Description

Upper jaw and mandibles of Tragoportax amalthea from Pikermi Tragoportax fossils.jpg
Upper jaw and mandibles of Tragoportax amalthea from Pikermi

Tragoportax species varied in size. Most species were approximately the size of a red deer. [3] In 1993, Meike Köhler estimated the body mass of T. amalthea at around 80 kg (180 lb). [16]

Skull and dentition

The snout of Tragoportax was short, though the basicranium was fairly long. [3] As with many modern antelope species, the horns are fairly diagnostic, and can be used to differentiate species. [17] In some species, like T. amalthea, the horns were twisted, whereas in others they were relatively straight. The horns of adult males were longer and slenderer than those of females and younger individuals, and often curved backwards. They were triangular or sub-triangular in cross-section, and were less laterally compressed than those of the related Miotragocerus. Some specimens of T. curvicornis are completely hornless. [3] Tragoportax's teeth were brachydont (low-crowned) to mesodont (having mid-sized crowns). The premolars are fairly long and are not molarised, though show the first signs of it. [18] They are shorter than those of Miotragocerus. The second premolar is shorter than the third, particularly the anterior (front) portion, with a parastyle that curved backwards. [3]

Postcrania

The postcranial elements of Tragoportax were similar in many respects to cervids. The limbs overall were fairly long, and it may have been at least somewhat cursorial. [16] The metapodials were large, slender and transversally compressed, overall resembling those of cervids. They were robust compared to those of Miotragocerus. [3]

Palaeoecology

Tragoportax amalthea likely inhabited lightly wooded environments. [16] On the other hand, T. macedoniensis (if it is Tragoportax), however, likely inhabited more forested environments, as it possesses traits characteristic of forest-dwelling bovids, such as reductions in body and horn size. [3] Based on morphological similarities with cervids like the red deer, at least some species probably were folivorous (had a leaf-heavy diet), [16] while others were graminivorous (had a grass-heavy diet). [19] On the other hand, analysis of dental isotopes and microwear suggest that T. rugosifrons was a mixed feeder, eating both leaves and grasses. [20] The evolution of browsing habits in Tragoportax may have been driven by increased competition from other grazing mammal clades, such as rhinocerotids and equids. [19]

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References

  1. 1 2 Roth, Johannes; Wagner, Johann Andreas (1854). "Die fossilen Knochenüberreste von Pikermi in Griechenland: Gemeinschaftlich bestimmt u. beschrieben nach d. Materialien, welche durch die von dem Erstgenannten im Winter 1852/3 dortselbst vorgenommenen Ausgrabungen erlangt wurden (in German)". Abhandlungen der Bayerische Akademie der Wissenschaften . 7: 371–464.
  2. ↑ Gaudry A. 1861. - Résultats des fouilles exécutées en Grèce under the auspices of the Académie. Comptes Rendus de l'Académie des Sciences de Paris 52: 297-300.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Spassov N. & Geraads D. 2004. - Tragoportax Pilgrim, 1937 and Miotragocerus Stromer, 1928 (Mammalia, Bovidae) from the Turolian of Hadjidimovo, Bulgaria, and a revision of the late Miocene Mediterranean Boselaphini. Geodiversitas26 (2): 339-370.
  4. ↑ Mindat.org: Tragocerus. mindat.org. Retrieved 2021-02-10.
  5. 1 2 Pilgrim, Guy Ellcock (1937). "Siwalik antelopes and oxen in the American Museum of Natural History". Bulletin of the American Museum of Natural History.
  6. ↑ Gentry, Alan William (1971). "The earliest goats and other antelopes from the Samos Hipparion fauna". Bulletin of the British Museum (Natural History) Geology. 20: 231––296.
  7. 1 2 3 Bibi, Faysal, Bukhsianidze, Maia, Gentry, Alan W., Geraads, Denis, Kostopoulos, Dimitris S., and Vrba, Elisabeth S., 2009. The Fossil Record and Evolution of Bovidae: State of the Field. Palaeontologia Electronica Vol. 12, Issue 3; 10A: 11p; http://palaeo-electronica.org/2009_3/169/index.html
  8. ↑ Carroll, Robert L. Vertebrate Paleontology and Evolution. WH Freeman and Company, New York. ISBN   0-7167-1822-7.
  9. ↑ Schlosser, Max (1904). "Die fossilen Cavicornia von Samos". Beitrage Paläontologie und Geologie 17. 17: 118.
  10. ↑ Kretzoi, Miklós (1941). "Neue antilopen-form aus dem Soproner Sarmat". Foldtani Közlöny. 71: 7–11.
  11. ↑ Kretzoi, Miklós (1968). "New generic names for homonyms" (PDF). Vertebrata Hungarica. 10 (1–2): 163–166.
  12. 1 2 Qin-Qin Shi & Zhao-Qun Zhang (2023) New material of Miotragocerus (Bovidae, Artiodactyla) from northern China and its systematic implications, Journal of Systematic Palaeontology, 21:1, doi : 10.1080/14772019.2023.2194891
  13. ↑ Solounias N. 1981. - The Turolian fauna from the Island of Samos, Greece with special emphasis on the Hyaenids and the Bovids. Contribution to Vertebrate Evolution 6: 1-232.
  14. ↑ "Bovidae | Ardipithecus kadabba: Late Miocene Evidence from the Middle Awash, Ethiopia | California Scholarship Online | Oxford Academic". academic.oup.com. Archived from the original on 2024-07-05. Retrieved 2024-12-12.
  15. ↑ Orak, Zahra; Kostopoulos, Dimitri S.; Ataabadi, Majid M. (May 22, 2023). "Late Miocene large-sized Bovidae (Mammalia) from Dimeh, SW Iran: contribution to depositional diachrony and palaeobiogeography". Geobios.
  16. 1 2 3 4 Köhler, Meike (1993). "Skeleton and habitat of recent and fossil ruminants" (PDF). Münchner Geowissenschaftliche Abhandlungen.
  17. ↑ Khan, Muhammad Akbar; Akhtar, Muhammad; Iqbal, Mehboob (2010). "The Late Miocene artiodactyls in the Dhok Pathan type locality of the Dhok Pathan Formation, the Middle Siwaliks, Pakistan". Pakistan Journal of Zoology Supplementary Series. 10: 1–87.
  18. ↑ Köhler, Meike (1987). "Boviden des turkischen Miozäns (Känozoikum und Braunkohlen der Türkei)" (PDF). Paleontologia y Evolucion. 21: 133–246.
  19. 1 2 Merceron, Gildas; Zazzo, Antoine; Spassov, Nikolaï; Geraads, Denis; Kovachev, Dimitar (2006-11-14). "Bovid paleoecology and paleoenvironments from the Late Miocene of Bulgaria: Evidence from dental microwear and stable isotopes". Palaeogeography, Palaeoclimatology, Palaeoecology. 241 (3): 637–654. doi:10.1016/j.palaeo.2006.05.005. ISSN   0031-0182.
  20. ↑ Merceron, Gildas; Zazzo, Antoine; Spassov, Nikolaï; Geraads, Denis; Kovachev, Dimitar (14 November 2006). "Bovid paleoecology and paleoenvironments from the Late Miocene of Bulgaria: Evidence from dental microwear and stable isotopes". Palaeogeography, Palaeoclimatology, Palaeoecology . 241 (3–4): 637–654. doi:10.1016/j.palaeo.2006.05.005 . Retrieved 6 September 2024– via Elsevier Science Direct.
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