Pecora

Pecora; Temporal range: 45–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Eocene - recent
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Suborder:	Ruminantia
Infraorder:	Pecora; Flower, 1883
Subgroups
	†Dromomerycidae; Antilocapridae; Bovimorpha Cervoidea Cervidae; ; Bovoidea Bovidae; Moschidae; ; ; Giraffomorpha †Palaeomerycidae; Giraffoidea Giraffidae; †Climacoceratidae; ; ;

Pecora is an infraorder of even-toed hoofed mammals with ruminant digestion. Most members of Pecora have cranial appendages projecting from their frontal bones; only two extant genera lack them, Hydropotes and Moschus.^[2] The name "Pecora" comes from the Latin word pecus, which means "cattle".^[3] Although most pecorans have cranial appendages, only some of these are properly called "horns", and many scientists agree that these appendages did not arise from a common ancestor, but instead evolved independently on at least two occasions.^[2]^[3]^[4]^[5] Likewise, while Pecora as a group is supported by both molecular and morphological studies, morphological support for interrelationships between pecoran families is disputed.^[2]

Evolutionary history

The first fossil ruminants appeared in the Early Eocene and were small, likely omnivorous, forest-dwellers.^[6] Molecular dating studies estimate that Ruminantia split into the two sister clades Pecora and Tragulina around 45 million years ago, during the Eocene.^[2] However, it was not until 15 million years later, at around 30 million years ago during the Oligocene, that the evolutionary radiation of Pecora began and the five families appeared (Bovidae, Cervidae, Moschidae, Giraffidae, and Antilocapridae).^[2]

The appearance of many pecoran fossils during the Miocene suggests that its rapid diversification may correspond to the climate change events of that epoch,^[6]^[7] as this time period was marked by much of Earth's forest habitats being replaced by grasslands due to widespread cooling and drying.^[2]

It is likely that the antelopes, giraffids, and pronghorns evolved in an open environment while the cervids, including the caribou, evolved in a woodland habitat.^[8] The type of gallop in pecoran species is shown to be closely related to their environment and anatomy, where light pecoran species use both flexed and extended suspensions in their fast gallops.^[8] The white-tail and mule-deer have been observed to primarily use the extended suspension, since in this phase of their gallop they leap over bushes and logs that are present in their brush environment.^[8] However, heavy pecoran species do not use extended suspensions as most have backs that slope downward with shorter hind legs.^[8]

Taxonomy and classification

Pecora is an infraorder within the larger suborder Ruminantia, and is the sister clade to the infraorder Tragulina (of which Tragulidae is the only surviving family).

Pecora's placement within Artiodactyla can be represented in the following cladogram:^[9]^[10]^[11]^[12]^[13]

Artiodactyla

Tylopoda (camels)

Artiofabula

Suina (pigs)

Cetruminantia

Ruminantia (ruminants)

	Tragulidae (mouse deer)

	Pecora (horn bearers)

Cetancodonta/Whippomorpha

	Hippopotamidae (hippopotamuses)

	Cetacea (whales)

Current attempts to determine the relationships among pecoran families (as well as all artiodactyls) rely on molecular studies, as little consensus exists in morphological studies.^[2] Different families within Pecora are recognized as valid by different groups of scientists.^[6]^{and sources therein, pp. 4–5}

Until the beginning of the 21st century it was understood that the family Moschidae (musk deer) was sister to Cervidae. However, a 2003 phylogenetic study by Alexandre Hassanin (of National Museum of Natural History, France) and colleagues, based on mitochondrial and nuclear analyses, revealed that Moschidae and Bovidae form a clade sister to Cervidae. According to the study, Cervidae diverged from the Bovidae-Moschidae clade 27 to 28 million years ago.^[14] The following cladogram is based on the 2003 study.^[14]

Pecora

	Bovidae

	Moschidae

Infraorder Pecora ("horned ruminants", "higher ruminants")

Incertae sedis
- Family †Dromomerycidae
- Family Antilocapridae (pronghorn)
Parvorder Bovimorpha
- Superfamily Cervoidea
  - Family Cervidae (deer)
- Superfamily Bovoidea
  - Family Moschidae (musk deer)
  - Family Bovidae (cattle, goats, sheep, and antelopes)
Parvorder Giraffomorpha
- Family †Palaeomerycidae
- Superfamily Giraffoidea
  - Family Giraffidae (giraffes and okapi)
  - Family †Climacoceratidae

Anatomy

Pecorans share characteristics with other artiodactyls, including a four-chambered stomach, and a paraxonic foot, meaning that it supports weight on the third and fourth digits. Several characteristics distinguish Pecora from its sister taxon, Tragulina: an astragalus with parallel sides, a loss of the trapezium, and differences in parts of the skull such as the petrosal bone.^[4]

The distinguishing features of most pecoran families are cranial appendages. Most modern pecorans (with the exception of the Moschidae) have one of four types of cranial appendages: horns, antlers, ossicones, or pronghorns.^[6]

True horns have a bone core that is covered in a permanent sheath of keratin. They are indicative of Bovidae. Horns develop in the periosteum over the frontal bone, and can be curved or straight.^[4] Surface features on the keratin sheath (e.g., ridges or twists) are thought to be caused by differential rates of growth around the bone core.^[4]
Antlers are bony structures that are shed and replaced each year in members of the family Cervidae. They grow from a permanent outgrowth of the frontal bone called the pedicle.^[4] Antlers can be branched, as in the white-tailed deer (Odocoileus virginianus), or palmate, as in the moose (Alces alces).
Ossicones are permanent bone structures that fuse to the frontal or parietal bones during the lifetime of an animal.^[4] They are found only in the Giraffidae and closely related extinct clades,^[4] represented in modern animals by the giraffe (Giraffa camelopardalis) and the okapi (Okapia johnstoni).
Pronghorns are similar to horns in that they have keratinous sheaths covering permanent bone cores; however, these sheaths are deciduous and can be shed like antlers.^[4] Very little is known about the development of pronghorns, but they are generally presumed to have evolved independently.^[4] The only extant animal with pronghorns is the pronghorn antelope (Antilocapra americana).

References

^ Flower, W. On the Arrangement of the Orders and Families of Existing Mammalia. Proceedings of the Zoological Society of London. 178 -186. 1883.
^ ^a ^b ^c ^d ^e ^f ^g Hassanin, A.; Douzery, E. (2003). "Molecular and Morphological Phylogenies of Ruminantia and the Alternative Position of the Moschidae". Systematic Biology. 52 (2): 206–228. doi:10.1080/10635150390192726. PMID 12746147.
^ ^a ^b Bubenik, A. Epigenetical, Morphological, Physiological, and Behavioral Aspects of Evolution of Horns, Pronghorns, and Antlers. in Horns, Pronghorns, and Antlers. G. Bubenik and A. Bubenik eds. Springer-Verlag. New York. 1990
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Janis, C., K. Scott. The Interrelationships of Higher Ruminant Families with Special Emphasis on the Members of the Cervoidea. American Museum Novitates. 2893: 1-85. 1987. http://digitallibrary.amnh.org/dspace/handle/2246/5180
^ Hassanin, A.; Delsuc, F.; Ropiquet, A.; Hammer, C.; Matthee, C.; Ruiz-Garcia, M.; Catzeflis, F.; Areskoug, V.; Nguyen, T. T.; Couloux, A. (2012). "Pattern and Timing of Diversification of Cetartiodactyla (Mammalia, Laurasiatheria), as Revealed by a Comprehensive Analysis of Mitochondrial Genomes". Comptes Rendus Biologies. 335 (1): 32–50. doi:10.1016/j.crvi.2011.11.002. PMID 22226162.
^ ^a ^b ^c ^d DeMiguel, D.; Azanza, B.; Morales, J. (2014). "Key Innovations in Ruminant Evolution: A Paleontological Perspective". Integrative Zoology. 9 (4): 412–433. doi:10.1111/1749-4877.12080. PMID 24148672.
^ Morales, J.; Pickford, M.; Soria, D.; Pachyostosis (1993). "Lorancameryx pachyostoticus Nov. Gen. Nov. Sp. and Its Bearing on the Evolution of Bony Appendages in Artiodactyls". Geobios. 26 (2): 207–230. doi:10.1016/S0016-6995(93)80016-K.
^ ^a ^b ^c ^d Dagg, Anne Innis (1967). Gaits and Their Development in the Infraorder Pecora (Thesis) – via UWSpace.
^ Beck, N.R. (2006). "A higher-level MRP supertree of placental mammals". BMC Evol Biol. 6: 93. doi:10.1186/1471-2148-6-93. PMC 1654192. PMID 17101039.
^ O'Leary, M.A.; Bloch, J.I.; Flynn, J.J.; Gaudin, T.J.; Giallombardo, A.; Giannini, N.P.; Goldberg, S.L.; Kraatz, B.P.; Luo, Z.-X.; Meng, J.; Ni, X.; Novacek, M.J.; Perini, F.A.; Randall, Z.S.; Rougier, G.W.; Sargis, E.J.; Silcox, M.T.; Simmons, N.B.; Spaulding, M.; Velazco, P.M.; Weksler, M.; Wible, J.R.; Cirranello, A.L. (2013). "The Placental Mammal Ancestor and the Post-K-Pg Radiation of Placentals". Science. 339 (6120): 662–667. doi:10.1126/science.1229237. hdl:11336/7302. PMID 23393258. S2CID 206544776.
^ Song, S.; Liu, L.; Edwards, S.V.; Wu, S. (2012). "Resolving conflict in eutherian mammal phylogeny using phylogenomics and the multispecies coalescent model". Proceedings of the National Academy of Sciences. 109 (37): 14942–14947. doi:10.1073/pnas.1211733109. PMC 3443116. PMID 22930817.
^ dos Reis, M.; Inoue, J.; Hasegawa, M.; Asher, R.J.; Donoghue, P.C.J.; Yang, Z. (2012). "Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny". Proceedings of the Royal Society B: Biological Sciences. 279 (1742): 3491–3500. doi:10.1098/rspb.2012.0683. PMC 3396900. PMID 22628470.
^ Upham, N.S.; Esselstyn, J.A.; Jetz, W. (2019). "Inferring the mammal tree: Species-level sets of phylogenies for questions in ecology, evolution, and conservation". PLOS Biology. 17 (12): e3000494. doi:10.1371/journal.pbio.3000494. PMC 6892540. PMID 31800571.(see e.g. Fig S10)
^ ^a ^b Hassanin, A.; Douzery, E. J. P. (2003). "Molecular and morphological phylogenies of Ruminantia and the alternative position of the Moschidae". Systematic Biology. 52 (2): 206–28. doi:10.1080/10635150390192726. PMID 12746147.