Canis lupus variabilis

Zhoukoudian wolf
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: Variabilis
Subspecies: C. l. variabilis
Trinomial name
Canis lupus variabilis
Pei, 1934[1]

Canis lupus variabilis (the Zhoukoudian wolf) is an extinct small wolf that once inhabited parts of what is now present-day China and Yakutia. The small wolf was later recognised as a variant of Canis variabilis (Pei 1934) that was also discovered and named by Pei in the same year.[2]

Discovery

The wolf's fossils were found at the Zhoukoudian (once spelt Choukoutien) cave system and archaeological site in 1934 and named by its discoverer, Pei Wenzhong.

"Although no sharp line can be traced between the above described Canis and a true C. lupus, the marked differences found in size, and in cranial characters, seem to be sufficient for creating, at least, a new variety, Canis lupus variabilis, for the Zhoukoudian Locality 1 small wolf.[3]:17

Pei stated that the Nihewan wolves[4] attributed to Canis chihliens should also be included in this new category.[3]:18 Canis lupus variabilis was also known from Lantian County in Shaanxi Province,[5] so it had a wide range in time and space.

At the site, the small wolf's remains were in close proximity to Homo erectus pekinensis or Peking man, in layers dating back to 500,000-200,000 years before present (YBP).

Distribution

Although discovered in China, fossil remains of Canis variabilis (Pei 1934) have been discovered in central Yakutia in Siberia on the Alaseya River and the Aldan River.[6] They are the oldest recorded samples of Olesky fauna found in Yakutia.[7] Canis cf. variabilis is thought to have been widespread in Eurasia until around 300,000 YBP and does not appear to overlap with the earliest occurrence of the morphologically distinctive gray wolf.[8]

Canis lupus variabilis - relationship to the domestic dog

Pei describes this small wolf as exhibiting variation in size and tooth adaptations, stating that it's skull differs from the typical wolf in much smaller size (about 175.0 mm total length for a large Canis lupus variabilis specimen), with a more slender muzzle and noticeably reduced or absent sagittal crest. In addition, the lower border of some Canis lupus variabilis mandibles is "strongly convex as in the dog".[3]:15 The one trait aligning Canis lupus variabilis with wolves is relatively large carnassial teeth (P1 20.4 – 23.0 mm; M1 22.0 – 24 mm). A later researcher has confirmed Pei's measurements, and describes the wolf's skull as having "heavy, wolf-like proportions although smaller than any extant Canis lupus.[9] More recent researchers have revisited Pei's view that the ancestor of the dog is a now extinct Canis lupus, and proposed that Canis lupus variabilis might be an ancestor of the dog lineage.[10][11]:7

The small wolf was later recognised as a variant of Canis variabilis (Pei 1934), also discovered and named by Pei in the same year.[2]

Canis variabilis - relationship to the dog and wolf

In 2010, a study found that the diversity of the Canis group decreased by the end of the Early Pleistocene to Middle Pleistocene and was limited to the small wolves of the Canis mosbachensis–Canis variabilis group and the large hypercarnivorous Canis (Xenocyon) lycaonoides in Eurasia. The true gray wolves made their appearance at the end of the Middle Pleistocene at about 0.5–0.3 million YBP.[12] Based on morphology, Canis mosbachensis has been proposed as the ancestor of the Canis lupus lineage.[13]:239–245 Based on genetics, Canis variabilis has been proposed as contributing to the wolf/dog lineage.[8] The two small wolves, Canis variabilis and Canis mosbachensis, are contemporaries in mid-latitude Eurasia and are similar in morphology, with phylogenetic analysis showing them to group together in a common region on a cladogram and they could represent one geographically-widespread mid-Pleistocene wolf.[14]:181

In 2015, a mitochondrial DNA analysis was conducted on 14 ancient canid remains from Arctic Siberia and suggests a genetic contribution from regional sources of wolves, including possibly Canis cf. variabilis, to the dog and modern wolf lineages. This was the first study to extract DNA material from Canis variabilis.[8]

References

  1. ↑ Wang, Xiao Ming & Tedford, R. H. (2008), Dogs: their fossil relatives and evolutionary history, Columbia University Press
  2. 1 2 Teilhard de Chardin, P., and W.-C. Pei. 1941. The fossil mammals from locality 13 of Choukoutien. Palaeontologica Sinica Series C 11: 1–106.
  3. 1 2 3 Pei, W.C. (1934). The carnivora from locality 1 of Choukoutien. Palaeontologia Sinica, Series C, vol. 8, Fascicle 1. Geological Survey of China, Beijing. pp. 1–45.
  4. ↑ Teilhard de Chardin, P and Pivetean, J. (1930). Les Mammiferes Fossiles De Nihowan (China). Annales de Paleontology vol 19, Paris. pp. 88–89.
  5. ↑ Hu, C. & Qi, T. (1978). Gongwangling Pleistocene mammalian fauna of Lantian, Shaanxi. Paleontologia Sinica, Whole Series no. 155, New Series C no. 21. Geological Survey of China, Beijing. pp. 1–64. Cited in S. Olsen 1985
  6. ↑ Vangengeim, E.A. (1961). "Paleontological study of anthropogenic sediment stratigraphy north of Eastern Siberia (on the fauna of mammals)". Proc. Geological Sciences of the USSR 48: 183.
  7. ↑ Sher, Andrei (1971). Pleistocene Mammals and stratigraphy of the Far Northeast USSR and North America. Science, Moscow. pp. 1–310.
  8. 1 2 3 Lee, E. (2015). "Ancient DNA analysis of the oldest canid species from the Siberian Arctic and genetic contribution to the domestic dog". PLoS ONE 10 (5): e0125759. doi:10.1371/journal.pone.0125759. PMC 4446326. PMID 26018528.
  9. ↑ Olsen, S. J. (1985). Origins of the domestic dog: the fossil record. Univ. of Arizona Press, Tucson, USA. pp. 88–89.
  10. ↑ Lawrence, B. (1967). "Early domestic dogs". Zeitschrift für Säugetierkunde 32: 44–59.
  11. ↑ Koler-Matznick, Janice (2002). "The Origin of the Dog Revisited" (PDF). Anthrozoös 15 (2): 98–118. doi:10.2752/089279302786992595.
  12. ↑ Sotnikova, M (2010). "Dispersal of the Canini (Mammalia, Canidae: Caninae) across Eurasia during the Late Miocene to Early Pleistocene". Quaternary International 212: 86–97. doi:10.1016/j.quaint.2009.06.008.
  13. ↑ Mech, L. David; Boitani, Luigi (2003). Wolves: Behaviour, Ecology and Conservation. University of Chicago Press. ISBN 0-226-51696-2.
  14. ↑ Tedford, R (2009). "Phylogenetic Systematics of the North American Fossil Caninae (Carnivora: Canidae)". Bulletin of the American Museum of Natural History 325: 1–218. doi:10.1206/574.1.
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