Creodonta

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Creodonta
Temporal range: Early Paleocene to Late Miocene, 63.3–11.1Ma [1]
SarkastodonDB.jpg
Sarkastodon
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Infraclass: Eutheria
Superorder: Laurasiatheria
Order: Creodonta
(Cope, 1875)
Families

Hyaenodontidae
Oxyaenidae
Limnocyonidae (?)

Creodonta is an extinct order of archaic, carnivorous mammals that lived from the Paleocene to the Miocene epochs. Because they both possess carnassial teeth creodonts and carnivorans were once thought to have shared a common ancestor, but given that different teeth are involved in making up the carnassials (both between creodonts and carnivorans and between the main groups of creodonts), the evolutionary history is no longer clear. It is even uncertain whether the creodonts form a monophyletic group. Two distinct families are generally recognized: Oxyaenidae and Hyaenodontidae, although their relationship is not entirely clear.

Creodonts had an extensive range, both geographically and temporally. They are known from the late Paleocene through the late Oligocene in North America, the early Eocene through late Oligocene in Europe, from the early Eocene through late Miocene in Asia, and from the Paleocene to the late Miocene in Africa.[2]

Creodonts were the first large, obviously carnivorous mammals with the radiation of the oxyaenids in the late Paleocene.[3] During the Paleogene they were the most abundant form of terrestrial carnivore in the Old World.[4] In Oligocene Africa, they were the dominant predatory group. They competed with the Mesonychids and the Entelodonts and ultimately outlasted them by the start of the Oligocene and by the middle of the Miocene respectively, but lost ground to the carnivorans. The last genus became extinct 11.1 million years ago, and carnivorans now occupy their ecological niches.

Systematics[edit]

Mount of oxyaenid Patriofelis from the American Museum of Natural History.

"Creodonta" was coined by Edward Drinker Cope in 1875.[5] Cope included the oxyaenids and the viverravid Didymictis but omitted the hyaenodontids. In 1880 he expanded the term to include Miacidae, Arctocyonidae, Leptictidae (now Pseudorhyncocyonidae), Oxyaenidae, Ambloctonidae and Mesonychidae.[6] Cope originally placed creodonts within the Insectivora. In 1884, however, he regarded them as a basal group from which both carnivorans and insectivorans arose.[7] Hyaenodontidae was not included among the creodonts until 1909.[8] Over time various groups were removed, and by 1969 it contained, as it does today, only the oxyaenids and the hyaenodontids.[9]

Skull of oxyaenid Machaeroides eothen.

One view of the position of the group is that Creodonta and Carnivora are sister taxa (within a superorder Ferae).[10] Others have argued that insectivorans are more closely related to carnivorans, and creodonts, therefore, are a basal eutherian.[11] Others have suggested that Creodonta might not be monophyletic.[12] Polly has argued that the only available synapomorphy between oxyaenids and hyaenodontids is a large metastylar blade on the first molar (M1), but he believes that that feature is common for all basal eutheria.[11] Separating Oxyaenidae from Hyaenodontidae would also comport with biogeographic evidence since the first oxyaenid is known from the North American early Paleocene and the first hyaenodontids are from very late Paleocene of North Africa.[13]

Complicating this arrangement is the tentative endorsement by Gunnell[14] of the erection of a third family, Limnocyonidae.[15] The group includes taxa that were once considered oxyaenids, such as Limnocyon, Thinocyon[8] and Prolimnocyon[16] Wortman had even erected a subfamily of Limnocyoninae within the oxyaenids.[17] Van Valen nests the same subfamily (including Oxyaenodon) within Hyaenodontidae[9] Gunnell is agnostic whether Limnocyonidae is a group within Hyaenodontidae (although a sister group to the rest of hyaenodontids) or entirely separate.[18]

Lateral (A) and dorsal (B) views of the skull of the hyaenodontid Apterodon macrognathus by Henry Fairfield Osborn.

According to Gunnell, the defining features of the oxyaenids include: A small braincase low in the skull. The occiput wide at base and narrowing dorsally (to give it a triangular shape). The lacrimal bone makes a semicircular expansion on the face. The mandibles have heavy symphysis. M1 and m2 form the carnassials, while M3/m3 are absent. The manus and pes are plantigrade or subplantigrade. The fibula articulates with the calcaneum, and the astragalus articulates with the cuboid bone. The phalanges are compressed and fissured at the tip.[18]

Likewise Gunnell's list of defining features of hyaenodontids includes: Long, narrow skull with a narrow basicranium and a high narrow occiput. The frontal bones are concave between the orbital regions. M2 and m3 form the carnassials. M3 is present in most species, while m3 is always present.Manus and pes range from plantigrade to digitigrade.The fibula artiuclates with the calcaneum, while the astragalar-cuboid articulation is reduced or absent. Terminal phalanges are compressed and fissured at the tip.[19]

The limnocyonids had the following features according to Gunnell: M3/m3 were reduced or absent, other teeth were unreduced. The rostrum was elongated. The animals themselves were small to medium sized.[20]

Morphology[edit]

Sinopa fossils: (1) Right upper cheek teeth, P2-M2; (2) Left ramus of mandible (p2-m2); (3) Right ramus of mandible (c-m2).

Dentition[edit]

Among primitive creodonts the dental formula is 3.1.4.33.1.4.3, but later forms often had reduced numbers of incisors, premolars and/or molars.[21] The canines are always large and pointed. The lateral incisors are large, while the medial incisors are usually small.[22] Premolars are primitive, with one primary cusp and various secondary cusps.[23]

Creodonts have two or three pairs of carnassial teeth, but only one pair performed the cutting function (either M1/m2 or M2/m3).[22] This arrangement is unlike modern carnivorans, which use P4 and m1 for carnassials,[24] and this suggests a separate evolutionary history and an order-level distinction.[25]

Cranium[edit]

Dorsal view of the skull of the limnocyonid (?) Limnocyon verus.

Creodonts had long, narrow skulls with small brains. The skull narrowed considerably behind the eyes, producing a distinct splanchnocranium and neurocranium segments of the cranium. They had large sagittal crests and usually broad mastoids (which were probably derived features for the group).[22]

Many creodonts had (porportionately) large heads.[26] In primitive forms the auditory bullae was not ossified. Generally the temporal fossae were very broad.[22]

Postcranial skeleton[edit]

Mounted skeleton of the hyaenodontid Sinopa rapax from Bridger Basin.

Creodonts had generalized postcranial skeletons. Their limbs were mesaxonic (with the axis of the foot provided by the middle of their five digits). Their method of locomotion ranged from plantigrade to digitigrade. The terminal phalanges were fused claws.[27]

Size[edit]

Lateral outline and front view of skull of Sarkastodon mongoliensis.

Creodonts ranged in size from the size of a small cat to hyaena-sized.[28] The larger sized animals, however, were not known until late in the Paleocene with the radiation of the oxyaenids,[29] such as the puma-sized Dipsalidictis and the probably bone-crushing scavenger Dipsalodon.[30]

Certain creodonts (Arfia, Prolimnocyon and Palaeonictis) seem to have experienced the dwarfing phenomenon during the Paleocene-Eocene Thermal Maximum seen in other mammal genera. A proposed explanation for this phenomenon is that the increased carbon dioxide levels in the atmosphere directly affected carnivores through increased temperature and aridity and also indirectly affected them by reducing the size of their herbivorous prey through the same selective pressures.[31]

The largest North American creodont is the Patriofelis. A specimen of P. ferox collected in the Bridger Basin of southern Wyoming was the size of a full-grown black bear with a head almost the size of an adult male lion.[32]

During the Central Asia Expedition of 1930 by the American Museum of Natural History the largest creodont ever discovered was collected: Sarkastodon mongoliensis. Its dimensions were described as 50% greater than the Patriofelis to which it bore many similarities.[33] It has been estimated that Sarkastodon attained the body mass of twice the largest American lion.[34] It is perhaps noteworthy that the Mongolian Eocene and Oligocene epochs produced two other massive mammals: the rhinocertid Baluchitherium and the entelodont Andrewsarchus.[33]

Upper teeth of Creodonts from Middle Eocene Bridger Basin, Wyoming

Biology[edit]

Diet and feeding[edit]

Early creodonts (both oxyaenids and hyaenodontids) displayed the tribosphenic molars common for basal therians. Small forms had somewhat strong postmetacrista-metastellar crests[35] suggesting they were probably opportunistic feeders, eating such things as eggs, birds, small mammals, insects and possibly plant matter as well,[36] possibly like extant viverrids.[21] Larger forms had greater shearing capacity and the capacity increased over time. Arfia, one of the most common carnivourous mammals in early Eocene North America, developed a more open trigonid on M3 over the course of the Early Eocene, increasing the shearing ability of the carnassials.[37] A similar development can be seen by comparing Oxaeyna, Protomus and Lymnocyon with the smaller, more generalized feeders among the creodonts.[36]

Evolution[edit]

Creodonts were traditionally considered ancestors to Carnivora, but are now considered to have shared a common ancestor. They share with the Carnivora the carnassial shear, a modification of teeth that evolved to slice meat in a manner like scissors and gave both orders the tools to dominate the niche. Some researchers argue that the creodonts represent a group of mammals of diverse biological ancestry that resemble one another via convergent evolution, rather than being the descendants of a single common ancestor. Their origins lie at least as far back as the late Cretaceous, though they did not radiate much until the Cenozoic. Creodonts were the dominant carnivorous mammals from 55 to 35 million years ago, peaking in diversity and prevalence during the Eocene.[38] By the mid Oligocene, Creodonts eventually supplanted the Mesonychids entirely in North America, and Eurasia, underwent a diversification in Africa, and in turn, competed with their own relatives, the Carnivorans. The last genus, Dissopsalis, became extinct about 11.1 million years ago.

Habitat[edit]

The creodonts ranged across North America, Eurasia and Africa, in forms that resemble those of some modern carnivores. Amongst their number was Sarkastodon, one of the largest mammalian land predators of all time, weighing an estimated 800 kg.[39] Their dominance over the early Carnivora, known as miacids, began to wane after 35 million years ago. The creodonts survived until 8 million years ago; the last form, Dissopsalis, died out in Pakistan. Bears, cats, mustelids, hyenas, canids, such as wolves, and other Carnivora now occupy the former creodont niches.

Extinction[edit]

Skull of Hyaenodon

It is not known exactly why the creodonts were replaced by Carnivora. It may be because of their smaller brains and their locomotion, which was somewhat less energy-efficient (especially while running).[40] Their limb structure limited leg movement to a vertical plane, as in horses; they were unable to turn their wrists and forearms inward to trip, slash, or grab prey as modern carnivores can. Creodonts had to depend entirely on their jaws to capture prey, which may be why creodonts generally had a larger head size in relation to their bodies than carnivores of similar stature. The creodont lumbosacral spine was not arranged as efficiently for running as in Carnivora. The arrangement of the teeth was also somewhat different. In the miacids (as with the modern Carnivora), the last upper premolar and the first lower molar are the carnassials, allowing grinding teeth to be retained behind for feeding on non-meat foods (the Canidae are the closest modern analog to miacid dentition). In creodonts, the carnassials were further back—either the first upper and second lower molars, or the second upper and third lower molars. This committed them to eating meat almost exclusively. These limits may have created important disadvantages over millions of years.

In the most hypercarnivorous family of modern Carnivora, the Felidae, the second and third molars have disappeared completely, and the first upper molars behind the carnassials have become vestigial. Modern cats thus eat plant food only incidentally.

References[edit]

  1. ^ "Creodonta". Paleobiology Database. Retrieved May 22, 2015. 
  2. ^ Gunnell, Gregg F. (1998). "Creodonta." pp. 91-109 at p. 91 in C.M. Janis, K.M. Scott, and L.L. Jacobs (eds.) Evolution of Tertiary Mammals of North America. Volume 1: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals. Cambridge University Press, Cambridge. ISBN 0-521-35519-2. (Hereafter "Gunnell.")
  3. ^ Janis, Christen M.; Baskin, Jon A.; Berta, Annalisa, et al. (1998). "Carnivorous mammals." pp. 73-90 at p. 73 in C.M. Janis, K.M. Scott, and L.L. Jacobs (eds.) Evolution of Tertiary Mammals of North America. Volume 1: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals. Cambridge University Press, Cambridge. ISBN 0-521-35519-2. (Hereafter "Janis, et al.")
  4. ^ Rose, Kenneth David; Archibald, J. David (2005). The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades. Baltimore: Johns Hopkiins University Press. p. 185.  (Hereafter "Rose.")
  5. ^ Cope, E.D. (1875). "On the Supposed Carnivora of the Eocene of the Rocky Mountains". Proceedings of the Academy of Natural Sciences, Philadelphia 27. pp. 444–449. 
  6. ^ Cope, E.D. (1880). "On the Genera of the Creodonta". Proceedings of the American Philosophical Society 19. pp. 76–82. 
  7. ^ Cope, E.D. (1884). The Vertebrata of the Tertiary Formations of the West. Washington, D.C.: U.S. Government Printing Office. 
  8. ^ a b Matthew, William Diller (August 1909). "The Carnivora and Insectivora of the Bridger Basin, Middle Eocene". Memoirs of the American Museum of Natural History 9. pp. 289–576. 
  9. ^ a b Van Valen, Leigh M. (1966). "Deltatheridia, a New Order of Mammals". Bulletin of the American Museum of Natural History 132. 
  10. ^ McKenna, M.C. (1975). "Toward a phylogenetic classification of the Mammalia." pp. 21-46 in W.P. Luckett and F.S. Szalay (eds.) Phylogeny of the Primates. Plenum, New York.
  11. ^ a b Polly, P.D. (1994). "What, if anything, is a creodont?" Journal of Vertebrate Paleontology 4. p. 42A.
  12. ^ Rose, p. 176
  13. ^ Janis, et al., p. 74.
  14. ^ Gunnell, pp. 91, 98-99
  15. ^ Gazin, Charles Lewis (January 17, 1962). "A further study of the lower Eocene mammalian faunas of southwestern Wyoming" (PDF). Smithsonian Miscellaneous Collections 144. Washington, D.C.: Smithsonian Museum. pp. 1–98. 
  16. ^ Matthew, William Diller; Granger, Walter (1915). "A revision of the Lower Eocene Wasatch and Wind River faunas". Bulletin of the American Museum of Natural History 34. pp. 4–103.  In this paper the authors rename Marsh's Limnocyon protenus as Didymictis protenus and include it among the myacids.
  17. ^ Wortman, J. Lewis (July 1902). "Studies of Eocene Mammalia in the Marsh Collection, Peabody Museum". American Journal of Science 164. pp. 17–23. 
  18. ^ a b Gunnell, p. 94.
  19. ^ Gunnell, p. 96
  20. ^ Gunnell, p. 98
  21. ^ a b Denison, Robert Howland (October 1937). "The Broad-Skulled Pseudocreodi". Annals of the New York Academy of Sciences 37. pp. 163–255.  (Subscription or payment required.)
  22. ^ a b c d Gunnell, p. 92.
  23. ^ Rose, p. 177
  24. ^ Feldhamer, George A.; Drickamer, Lee C.; Vessey, Stephen H.; Merritt, Joseph F.; Krajewski, Carey (2015). Mammalogy: Adaptation, Diversity, Ecology. Baltimore: Johns Hopkins University Press. p. 356. ISBN 978-0801886959. 
  25. ^ Turner, Alan; Antón, Mauricio (2004). Evolving Eden: An Illustrated Guide to the Evolution of the African Large-Mammal Fauna. New York: Columbia University Press. p. 77. ISBN 0-231-11944-5. 
  26. ^ Rose, p. 178
  27. ^ Gunnell, p. 93.
  28. ^ Gunnell, p. 91
  29. ^ Janis, p. 73
  30. ^ Gunnell, Gregg F.; Gingerich, Philip D. (September 30, 1991). "Systematics and evolution of late Paleocene and early Eocene Oxyaenidae (Mammalia, Creodonta) in the Clarks Fork Basin, Wyoming" (PDF). Contributions from the Museum of Paleontology, University of Michigan 28. Ann Arbor: Museum of Paleontology, University of Michigan. pp. 141–180. 
  31. ^ Chester, Stephen G.B.; Bloch, Jonathan I.; Secord, Ross; Boyer, Doug M. (2010). "A New Small-Bodied Species of Palaeonictis (Creodonta, Oxyaenidae) from the Paleocene-Eocene Thermal Maximum". Journal of Mammalian Evolution 17. pp. 227–243. 
  32. ^ Wortman, Jacob L. (1894). "Osteology of Patriofelis, a Middle Eocene Creodont" (PDF). Bulletin American Museum of Natural History 6. pp. 129–164. 
  33. ^ a b Granger, Walter (April 21, 1938). "A Giant oxyaenid from the Upper Eocene of Mongolia" (PDF). American Museum Novitates (969). pp. 1–5. 
  34. ^ Sorkin, Boris (2008). "A biomechanical constraint on body mass in terrestrial mammalian predators" (PDF). Lethaia 41. pp. 333–347. 
  35. ^ Gingerich, Philip D. (1989). "New Earliest Wasatchian Mammalian Fauna from the Eocene of Northwestern Wyoming: Composition and Diversity in a Rarely Sampled High-Floodplain Assemblage". Papers on Paleontology (28). Ann Arbor: Museum of Paleontology, University of Michigan. p. 37.  (Hereafter “Gingerich.”)
  36. ^ a b Gunnell, p. 100
  37. ^ Gingerich, p. 34, Fig. 20 on p. 35 & Fig. 22 on p. 37.
  38. ^ Lambert, 162.
  39. ^ Sorkin, B. 2008: A biomechanical constraint on body mass in terrestrial mammalian predators. Lethaia, Vol. 41, pp. 333–347
  40. ^ "The Elements of Geology". Globusz. Retrieved March 11, 2008. 

Additional Reading[edit]

  • The Velvet Claw: A Natural History of the Carnivores, David Macdonald, BBC Books, ISBN 0-563-20844-9
  • David Lambert and the Diagram Group. The Field Guide to Prehistoric Life. New York: Facts on File Publications, 1985. ISBN 0-8160-1125-7