Reconstruction of Brachyodus aequatorialis, copyright WillemSvdMerwe.

Belongs within: Artiodactyla.
Contains: Protoceratidae, Tragulidae, Pecora.

The Ruminantiamorpha are a clade of artiodactyls including all species closer to modern ruminants than to other living artiodactyls. Living ruminants are characterised by the possession of a multi-chambered stomach in which food is broken down by bacteria living in a large fermentation chamber, the rumen.

Horns and guts
Published 7 April 2008
Jacob breed of sheep Ovis aries, from Bide a Wee Farm.

This page’s subject will, I’m sure, be familiar to most of you. Even kids in the depths of urbania will usually learn at some stage that cows say “moo” and sheep say “baa”. Today’s subjects are the Ruminantia, the primary group of large herbivorous mammals on the planet today. Including such animals as sheep, cattle, goats and deer, ruminants are also one of the most significant groups of animals in modern human lifestyles.

Modern ruminants fall into six families. The vast majority of modern ruminant diversity is contained within two of those families, the Cervidae (deer—16 genera) and Bovidae (cattle, goats, antelope, etc.—48 genera), particularly the latter. The Tragulidae (chevrotains or mouse deer) are two or three genera of small inhabitants of Old World tropical forests. The Giraffidae includes just two genera—the giraffe Giraffa and okapi Okapia johnstoni. Finally, two genera are assigned families of their own—the pronghorn Antilocapra americana and the musk deer Moschus. Fossil-wise, there are a whole host of extinct families, mostly of small deer-like animals, and some of the smaller living families—notably the Giraffidae and Antilocapridae—were much more diverse in the past than they are now.

Banteng Bos javanicus, a species of wild cattle native to south-east Asia that has been introduced in northern Australia. Photo from here.

Ruminants are best-known for their impressive digestive systems, which are regarded as the secret of their current success. Most other mammals living mostly on low-grade plant matter (i.e. browsers and grazers) are what are called “hindgut fermenters”—the beginning of the large intestine forms a large chamber where bacteria break down the cell walls of the plants eaten, releasing the nutrients that would otherwise be locked within into the digestive system. Hindgut fermentation is a fairly simple system, but the problem is that it is relatively inefficient. By the time the plant matter has reached the large intestine, it has already passed through most of the digestive system, leaving only a relatively short distance in which the released nutrients can be absorbed. Most hindgut fermenters compensate for this inefficiency by being very large, their concordantly large guts allowing more time for digestion*. A good example of this can be seen in the gorilla, a specialised folivore (leaf-eater) that is considerably larger than its close non-hindgut-fermenting relatives.

*One exception to this pattern is the Leporidae (hares and rabbits), which have a different method of getting around the limitations of hindgut fermentation—they eat their own faeces, so passing their food through the digestive system twice.

Ruminants, in contrast, are foregut fermenters. In ruminants it is not the large intestine but the stomach that has become enlarged and subdivided to contain fermentative bacteria (the so-called “four stomachs” of cattle—Macdonald 1984). Ruminants also complement the digestive process by regurgitating and further chewing their food (“cud-chewing”) when not directly grazing. Because the plant cell walls are broken down before passing through the majority of the digestive system, ruminants are able to derive far more nutrients from their food than hindgut fermenters. They are also able to subsist on feed on lower quality, which lead to their taking over as the dominant grazers when the climate cooled and grasslands spread during the Miocene.

Barren-ground caribou Rangifer tarandus groenlandicus, a member of the Cervidae. Photo from here.

Phylogenetically, it is well-established that the Tragulidae are the basalmost of the living ruminant families, the remainder forming a clade called the Pecora. Chevrotains have a less-developed rumination system than other ruminants – while they have four stomach chambers as in other ruminants, the third chamber is poorly developed. Within Pecora, relationships are more controversial. While it seems that Cervidae, Moschus and Bovidae form a clade to the exclusion of Giraffidae and Antilocapra, it is unclear whether the latter two taxa form a sister clade to the other pecorans, or a paraphyletic series. It also has recently been suggested that Moschus may be sister to Bovidae, in contrast to its traditional position closer to the Cervidae (Hassanin & Douzery 2003). Were one to broaden the investigation to cover the various extinct taxa, the situation just degrades to a hopeless mess.

Musk deer, photographed by Ben Cooper.

Part of the reason for this confusion is that homoplasy seems to be rife within the ruminants. The most intriguing feature of the group for me is the repeated evolution of horns or antlers (technically, cranial appendages). Four of the five living pecoran families have some form of cranial appendage, and indications are that they have arisen independently in each family. This is reflected in the different structure of the appendages in each family (Macdonald 1984). Bovids have permanent horns of bone covered with a layer of keratin. In Cervidae, the antlers are deciduous bone, and are shed and regrown annually. Giraffids have ossicones of permanent bone covered with a layer of skin. The horns of Antilocapra are permanent bone as in Bovidae, but the outer layer of keratin is shed annually. The Tragulidae and Moschus, which lack horns, both have enlarged canines (the above photo of a musk deer shows just how enlarged). Enlarged canines are also found in the deer genera Hydropotes (the Chinese water deer) and Elaphodus (the tufted deer), which have small antlers (Elaphodus) or lack them altogether (Hydropotes). There seems to be a rough inverse relationship between enlarged canines and cranial appendages, where the development of the latter is generally (though not invariably) correlated with the loss of the former. Interestingly, recent phylogenetic analyses indicate that Hydropotes is actually nested among antlered deer, indicating that it regained enlarged canines as the antlers were lost (Gilbert et al. 2006).

Reconstruction of Hoplitomeryx, from Scontrone.

Again, when the extinct taxa are factored in, the picture becomes even more complicated. For instance, the Climacoceratidae are an extinct family that dental features indicate is related to the Giraffidae, and which had ossicones similar in structure to those of the giraffids (albeit long and branched in some forms, so similar in appearance to a deer’s antlers). However, the basal members of the Climacoceratidae lack ossicones, indicating that they were evolved independently in the two families (Morales et al. 1999). The most dramatic armament among fossil ruminants was perhaps that found on the Mediterranean endemic Hoplitomeryx, found in the Miocene on what is now Monte Gargano on the eastern coast of Italy, but was then a separate island (Hassanin & Douzery, 2003). Hoplitomeryx, as shown above, had a grand total of five horns on its head, as well as long dagger-like canines (providing an exception to the horns vs. canines rule mentioned above). It has been suggested that the overdone armation of Hoplitomeryx may have evolved as a defense against the large birds of prey that would have been its major (indeed, its only) predators on Gargano.

Systematics of Ruminantiamorpha

Synapomorphies (from Janis & Scott 1988): Tarsus with fused cuboid and navicular; upper incisors reduced or lost; incisiform lower canines.

<==Ruminantiamorpha [Tragulina]SOG09
    |--+--Mixtotherium Filhol 1880GT09, SM93 [Mixtotheriidae]
    |  |    `--M. mezi Schmidt 1913S68
    |  `--Elomeryx Marsh 1894SOG09, D07
    |       |--E. armatus (Marsh 1894)S96
    |       |--E. crispusOB13
    |       `--E. woodiD07
    `--+--Leptoreodon Wortman 1898GT09, SM93
       |    |--L. edwardsiW96
       |    |--L. leptolophus Golz 1976W96
       |    |--L. major Golz 1976W96
       |    |--L. marshi Wortman 1898W96
       |    |--L. pusillus Golz 1976W96
       |    `--L. stockiW96
       `--+--Heteromeryx disparGT09
          `--+--+--Libycosaurus petrocchii Bonarelli 1947SOG09, B78 [=Merycopotamus petrocchiiB78]
             |  `--ProtoceratidaeSOG09
             `--+--Hypertragulidae [Hypertraguloidea]JS88
                |    |--Hypertragulus Cope 1874GT09, SM93
                |    |--HypisodusJS88
                |    |--Nanotragulus loomisiJS88, TS96
                |    |--Parvitragulus Emry 1978SM93
                |    |--Miomeryx Matthew & Granger 1925SM93
                |    `--SimimeryxHD03
                |         |--S. hudsoni Stock 1934W96
                |         `--S. minutusP96
                     |  i. s.: Russa equinaM89
                     |         IndomeryxJS88
                     |         PseudomeryxJS88
                     |         Archaeomeryx Matthew & Granger 1925D07 [ArchaeomerycidaeHD03]
                     |           `--A. optatusD07
                     |         Hexameryx White 1941D07
                     |           |--H. elmoriD07
                     |           `--H. simpsoniD07
                     `--+--Bachitherium [Bachitheriidae]JS88
Ruminantiamorpha incertae sedis:
  Bothriogenys Schmidt 1913SOG09, B78
    |--*B. fraasi (Schmidt 1913) [=Brachyodus (*Bothriogenys) fraasi]B78
    |--B. africanus (Andrews 1899)B78
    |--B. andrewsi Schmidt 1913B78
    |--B. gorringei (Andrews & Beadnell 1902)B78
    |--B. parvus (Andrews 1906)B78
    `--B. rugulosus Schmidt 1913B78
  Brachyodus Deperet 1895BLB05, SM93
    |--B. aequatorialis MacInnes 1951BLB05, B78 [=Masritherium aequitorialisB78]
    |--B. andrewsi Schmidt 1913S68
    |--B. gorringei (Andrews & Beadnell 1902)S68
    |--B. parvulus (Andrews 1906)S68
    `--B. rugulosus Schmidt 1913S68

*Type species of generic name indicated


[B78] Black, C. C. 1978. Anthracotheriidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 423–434. Harvard University Press: Cambridge (Massachusetts).

[BLB05] Boisserie, J.-R., F. Lihoreau & M. Brunet. 2005. The position of Hippopotamidae within Cetartiodactyla. Proceedings of the National Academy of Sciences of the USA 102 (5): 1537–1541.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GT09] Geisler, J. H., & J. M. Theodor. 2009. Hippopotamus and whale phylogeny. Nature 458: E1–E5.

Gilbert, C., A. Ropiquet & A. Hassanin. 2006. Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): systematics, morphology, and biogeography. Molecular Phylogenetics and Evolution 40 (1): 101–117.

[HD03] Hassanin, A., & E. J. P. Douzery. 2003. Molecular and morphological phylogenies of Ruminantia and the alternative position of the Moschidae. Systematic Biology 52 (2): 206–228.

[JS88] Janis, C. M., & K. M. Scott. 1988. The phylogeny of the Ruminantia (Artiodactyla, Mammalia). In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 273–282. Clarendon Press: Oxford.

Macdonald, D. 1984. All the World’s Animals: Hoofed Mammals. Torstar Books: New York.

[M89] Modigliani, E. 1889. Appunti intorno ai mammiferi dell’isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 238–245.

Morales, J., D. Soria & M. Pickford. 1999. New stem giraffoid ruminants from the early and middle Miocene of Namibia. Geodiversitas 21 (2): 229–253.

[OB13] O’Leary, M. A., J. I. Bloch, J. J. Flynn, T. J. Gaudin, A. Giallombardo, N. P. Giannini, S. L. Goldberg, B. P. Kraatz, Z.-X. Luo, J. Meng, X. Ni, M. J. Novacek, F. A. Perini, Z. S. Randall, G. W. Rougier, E. J. Sargis, M. T. Silcox, N. B. Simmons, M. Spaulding, P. M. Velazco, M. Weksler, J. R. Wible & A. L. Cirranello. 2013. The placental mammal ancestor and the post-K–Pg radiation of placentals. Science 339: 662–667.

[P96] Prothero, D. R. 1996. Magnetic stratigraphy and biostratigraphy of the Middle Eocene Uinta Formation, Uinta Basin, Utah. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 3–24. Cambridge University Press.

[S68] Simons, E. L. 1968. African Oligocene mammals: introduction, history of study, and faunal succession. Peabody Museum of Natural History, Yale University, Bulletin 28: 1–21.

[SOG09] Spaulding, M., M. A. O’Leary & J. Gatesy. 2009. Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretation of key fossils and character evolution. PLoS One 4 (9): e7062.

[S96] Storer, J. E. 1996. Eocene-Oligocene faunas of the Cypress Hills Formation, Saskatchewan. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 240–261. Cambridge University Press.

[SM93] Stucky, R. K., & M. C. McKenna. 1993. Mammalia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 739–771. Chapman & Hall: London.

[TS96] Tedford, R. H., J. B. Swinehart, C. C. Swisher III, D. R. Prothero, S. A. King & T. E. Tierney. 1996. The Whitneyan-Arikareean transition in the High Plains. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 312–334. Cambridge University Press.

[W96] Walsh, S. L. 1996. S. Middle Eocene mammal faunas of San Diego County, California. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 75–119. Cambridge University Press.

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