Canis lupus Linnaeus, 1758

TEDFORD R. H., WANG X. & TAYLOR B. E., 2009, Phylogenetic Systematics Of The North American Fossil Caninae (Carnivora: Canidae), Bulletin of the American Museum of Natural History 2009 (325), pp. 1-218 : 148-150

Canis lupus Linnaeus, 1758 Appendix 3

Canis lupus Linnaeus, 1758 .

Canis lupus lunellensis Bonifay, 1971 .

Medial Pleistocene North American Material: From Cripple Creek Sump (Olyorian, late Irvingtonian), near Fairbanks, Alaska: F:AM 67186, left ramus with i1–i3 alveoli, i2 root, c–m2, and m3 alveolus; F:AM 67183, right ramus with i1–p3 alveoli and p4 broken, m1–m2, and m3 alveolus; F:AM 67181, left ramus with i1–i3 alveoli, c, p1 alveolus, p2–m2, m3 alveolus; F:AM 67173, left maxillary fragment with part C alveolus, p1–p3 alveoli, P4–M2; F:AM 97107, fragment of right ramus with i1–i3, c, p1– p3 alveoli, p4, m1 alveolus; F:AM 97109, left ramus with i1–i3, c, p1–p4, m1 alveolus; F:AM 97091 and 97091A, two left partial humeri; F:AM 97093, 97093A, and 97093B, left proximal and two distal ends of right radii; F:AM 97095, 97092, 97092A, and 97092B, three right partial ulnae; F:AM 68008C, right partial ulna; F:AM 97095 and 97095A, left partial ulna and partial femur; F:AM 90974A and 90974B, right partial femur, right and left partial tibiae; F:AM 68010A, right incomplete tibia; F:AM 68010D, right tibia; F:AM 68012O, right metacarpal V; F:AM 68012J, left metacarpal V; and F:AM 68013F, left calcaneum.

From Old Crow River, northern Yukon, northwestern Canada, CRH locality 47, unit 2, beneath Surprise Creek Tephra in rocks of normal polarity attributed to the Jaramillo Subchron, Matuyama Chron, medial Pleistocene (1.11 Ma, see Repenning, 1992, for discussion): CMN 39490, right M1.

Distribution: Earliest North American record is medial Pleistocene (Olyorian, equivalent in part to late Irvingtonian) in the Arctic at Old Crow locality CRH 47 in Yukon, Canada. More material is available in slightly younger beds at the Cripple Creek Sump locality near Fairbanks, Alaska. Midcontinent records begin in the late Rancholabrean.

Description and Comparison: The unworn right M1 from the Yukon (CMN 39490) is the oldest (Jaramillo) material attributed to C. lupus from North America. This early C. lupus is presently represented only by a single upper cheek tooth (M1, length, 17.4 mm, width, 20.2 mm). The available specimen shows a number of dental features that are synapomorphic with C. lupus . Notable among these are the hypertrophy of the paracone with respect to the metacone [table 1, character 54(1)], enlargement of hypocone [1(1)], preparacrista di- rected anteriorly, lingual to the parastyle [3(2)], and reduction of the metaconule [59(1)].

The gold-bearing alluvial deposits of the region west of Fairbanks, Alaska, have produced large quantities of Quaternary mammalian fossils. The largest samples reside in the Frick Collection at AMNH and at the University of Alaska. The exigencies of collection of this material during hydraulic mining of the deposits provided few opportunities for stratigraphic allocation of the remains. Moreover, most of this work was carried out in the 1930s and early 1940s before a consistent stratigraphy had been developed for the Quaternary deposits. Troy Péwé’s monumental studies (1975a, 1975b, 1989) were the first synthesis of the stratigraphy of the Quaternary deposits in the Fairbanks area. This was later refined with the application of tephrochronology and magnetostratigraphy ( Westgate et al., 1990) so that it is now possible to see that the gold-bearing deposits extend, with hiatuses, from the medial Pliocene through the Pleistocene.

Unfortunately, very few of the large collections of mammals can be clearly related to this stratigraphic scheme and their ages are consequently uncertain. However, there is one locality, on Cripple Creek, at the dredging site call the Sump, that yielded a suite of mammalian remains from a more circumscribed lower part of the local column. At the Cripple Creek Sump, faulting has preserved a section of the Fox Gravels and the lower part of the overlying Gold Hill Formation. Closely associated occurrences of the Ester Ash Bed in the lower Gold Hill Formation yielded an isothermal plateau fission track age of 0.81 ± 0.07 Ma ( Westgate et al., 1990). At nearby Gold Hill this part of the sequence lies in a reversed magnetozone correlated with the latest phase of the Matuyama Chron.

This chronology corresponds with the late Irvingtonian of mid-latitude North America, although the Alaskan fauna of this time contains many taxa that do not appear south of the continental glacial barrier until the Rancholabrean (e.g., Bison , Ovibos , Ovis , and Panthera leo ). The fauna at the Cripple Creek Sump (verified with F:AM records and examination of specimens) includes: Lepus sp. , Ochotona sp. , Castor sp. , Xenocyon lycaonoides , Canis cf. lupus , Panthera leo , Ursus arctos , Taxidea taxus , Mammut sp. , Mammuthus sp. , Cervus cf. elaphus , Rangifer sp. , Alces cf. latifrons, Bison cf. priscus , Bootherium bombifrons ( McDonald and Ray, 1989), Praeovibos recticornis and P. priscus ( McDonald et al., 1991), Ovibos sp. , Ovis sp. , Saiga tartarica ( Harington, 1981) , Camelops sp. , and Equus sp. In its composition it more closely corresponds with the Olyor faunas of adjacent northeastern Siberia ( Sher, 1986), especially in the co-occurrence of Xenocyon , Canis cf. lupus , lion, Saiga , Bison , Praeovibos , and Alces cf. latrifrons. North American elements in this New World Olyorian fauna include the badger Taxidea , the lamine camel Camelops , and the endemic ovibovine Bootherium .

As far as dental measurements are concerned, the Cripple Creek Sump wolves are clearly nested within a suite (n 5 12) of C. lupus lycaon from Minnesota in the AMNH(M) collection. They lie below most values of C. l. occidentalis (n 5 11) from Alberta and Montana, but fall close to the two females of C. lupus from northeastern Siberia in the AMNH(M) collection. The Sump specimens have dentitions that lie above the range of samples of the medial Pleistocene wolf C. mosbachensis from Europe (see appendix 1): length of m1: Untermassfeld, Germany (n 5 16, 23.3– 25.8 mm; Sotnikova, 2001); Stránska Skála, Czechoslovakia (n 5 18, 20.5–24.1 mm, Musil, 1972); Mosbach, Germany (n 5 5, 22.4–24.7 mm, Adam, 1959: table 4); Hundesheim, Austria (n 5 3, 23.9–24.0 mm, Thenius, 1954: table 4); Heppenloch, Germany (n 5 2, 24.0–26.0 mm, Adam, 1959: table 4), and Westbury-sub-Mendip, England (n 5 7, 21.5–25.7 mm, Bishop, 1982: table 28).

Only a few ramal fragments and a single maxillary are known from the Cripple Creek Sump. The rami provide most of the characters: none of the incisors are known and the canines are so worn that they provide no basis for description. The lower premolar row curves medially as is typical of C. lupus . The premolars are separated by diastemata and are closely similar in individual length, forming a series that increases posteriorly at about 1 mm per tooth. The p2 may have a posterior cusp, and the p3 consistently does so. The p4 has a tiny second posterior cusp lying in front of, or discernible from, the posterior cingulum in three of the four rami in which it can be observed. This is a primitive feature in Canis sp. , more consistently present in C. mosbachensis than in C. lupus , where this cusp cannot be differentiat- ed from the posterior cingulum. The tip of p4 lies below the paraconid of m1; p3 lies below p4 and p2; and the p4 lacks a strong posterolingual shelf as in C. mosbachensis . The m1 and m2 show no peculiarities except for the better developed anterolabial cingulum on m2 of the Sump fossils. The largest and most anterior mental foramen lies below the anterior root of p2, with the smaller one below p3. The horizontal ramus gently tapers forward, and there is a shallow inflection in the ventral border behind the symphysis. The masseteric fossa is deep, especially anteriorly, and extends beneath m3; ventrally it is marked by a strong masseteric crest. The angular process is typical of Canis .

The left maxillary fragment, F:AM 67173, bears P4, M1–M2, and parts of the palate and alveoli for P1–P3. From this specimen the following cranial measurements, in millimeters, can be approximated following the craniometry adopted herein: LPM 79.0, MW about 76, PW P1 about 30, P4L 23.2, WM2 11.5. These dimensions fall within the Minnesota sample of C. lupus used for this work. As in C. lupus , the infraorbital foramen lies above the posterior root of P3 and the largest posterior palatine foramen lies on the maxillary-palatine suture opposite the paracone of P4; a smaller second foramen lies within the edge of the palatine opposite M1. The most anterior position of the palatine notch lies opposite the rear of M2. The P4 has a stout protocone that extends slightly anterolingually but does not extend beyond the anterior margin of the tooth. The M1 has a conspicuously enlarged paracone and a low parastyle connected to the paracone by the preparacrista; the labial cingulum is subdued but largely present. The talon is narrow, but it includes a differentiated weak metaconule, a trace of the paraconule, and a low protocone. The hypocone is also sub- dued, and the lingual cingulum joins the metaconule and extends around the protocone to the parastyle. The M2 is relatively large compared with M1, and the paracone is considerably larger than the metacone. No conules are developed from the protocristae. The lingual and labial cingula nearly completely encircle the tooth except across the metacone. The hypocone is not a differentiated cusp.

Wolves close in size and morphology to the Canis sample from the Cripple Creek Sump appear in Europe near the close of the medial Pleistocene (‘‘Mindel-Riss’’ interglacial, sensu Bonifay, 1971, ca. 0.4 Ma), where a sample from the Lunel-Viel Cave of southern France (Herault) was designated Canis lupus lunellensis by Bonifay (1971). This taxon was differentiated from living wolves solely on its smaller size. The length of m1 of 10 individuals ranged from 23.8 to 27.4 mm, the m2 (n 5 9) from 9.6 to 10.8 mm. The m1 and m2 fall mostly below the range of Cripple Creek Sump. However, the single upper dentition from the Sump lies within the range of the dental measurements of C. l. lunellensis (taking into account differences in the measurement of width due to choices of reference points). Morphologically the Sump wolves closely resemble living North American wolves of comparable size as well as C. l. lunellensis.

Thus, the Alaskan material of medial Pleistocene age indicates the presence of a population that clearly belongs to Canis lupus . If the geological attribution of this material is correct, and its age is ca. 1.0 Ma, wolves similar to the living species were part of a high-latitude Arctic fauna (Olyorian) that existed contemporaneously with other species of Canis in the Irvingtonian of the middle latitudes of North America.