Canis lupus spelaeus Goldfuss, 1823

Marciszak, Adrian & Kropczyk, Aleksandra, 2025, Canidae (Carnivora, Mammalia) from Niedźwiedzia Cave (Silesia, southwest Poland), Palaeontologia Electronica (a 15) 28 (1), pp. 1-33 : 2-11

publication ID	https://doi.org/10.26879/1510
persistent identifier	https://treatment.plazi.org/id/905687B3-FF9B-FFFC-FC4A-FE90FD8F3300
treatment provided by	Felipe
scientific name	Canis lupus spelaeus Goldfuss, 1823
status

Treatment

( Figures 2–8 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 View FIGURE 7 View FIGURE 8 ; Tables 1 –3; Appendix 1)

Referred material. The material of Canis lupus spelaeus , the most abundant canid from Niedźwiedzia Cave, is represented by almost all skeletal elements ( NISP 479, MNI 18; left/right bones are given in brackets): two crania fr., nine maxillae fr. (6/3), one I2 (0/1), six I3 (3/3), 16 C1 (5/11), three P1 (1/2), nine P2 (3/6), eight P3 (5/3), 20 P4 (12/8), 15 M1 (8/7), one M2 (1/0), 32 mandibles (14/18), two i1 (1/1), nine i2 (4/5), two i3 (1/1), 18 c1 (7/11), three p1 (1/2), one p2 (0/1), 11 p3 (5/6), seven p4 (5/2), 18 m 1 (10/8), two scapulae (2/0), 20 cervical vertebrae, four thoracic vertebrae, seven lumbar vertebrae, two caudal vertebrae, ten costae, 16 humeri (7/9), eight radii (4/4), three ulnae (2/1), one scapholunar (0/1), two pisiformes (1/1), two cuboides (0/2), ten mc 2 (5/5), nine mc 3 (4/5), eight mc 4 (6/2), 16 mc 5 (9/7), 11 femora (9/ 2), 15 tibiae (7/8), three naviculares (2/1), 18 calcanei (9/9), 15 talii (7/8), 15 mt 2 (9/6), 18 mt 3 (10/8), eight mt 4 (4/4), ten mt 5 (4/6), 33 phalanges 1, 10 phalanges 2, and 10 phalanges 3.

Description. The upper tooth row is curved distally and the teeth are arranged tightly. The broad and curved incisor row is moderately extended forward ( Figure 2 View FIGURE 2 ). The simply built I1 is short mesio-distally and moderately developed bucco-lingually. The apex of the crown is oriented mesio-buccally and forms an asymmetric triangle. The mesial edge is larger than the distal one. The mesial valley between them is V-shaped, shallow, and both edges are connected to each other. The I2 has a massive and compact crown, the top of which is directed distally and slightly buccally. The buccal part of the crown is concave and separated from the convex lingual side by a thin and V-shaped cingulum, almost evenly developed mesially and distally. The large, canine-like I3 has a crown placed at an angle of 40 o to the root axis. The crown, ovoid, or oval in cross-section, is also flattened on both sides, especially on the lingual side. The top of the crown is twisted disto-lingually. Two edges extend from it, mesial and distal, which at the base of the crown connect with the moderately developed lingual cingulum, forming a thin ridge around the crown. The robust C1 are flattened laterally and elongated mesio-distally. Weak mesial and distal crests run from their apex to the base.

The oval-shaped P1 is a small, monocuspid, single-rooted tooth. Buccal and lingual margins are convex, and the lingual one is more strongly developed. The P2 is an elongated tooth, with a strong distal cingulum ( Figure 2 View FIGURE 2 ). The buccal margin is straight and the lingual margin is gently convex in its middle part. The mesial and distal margins are blunt. The protoconid is situated more mesially. Two thin edges run from the protoconid apex, the mesial one in mesio-lingual direction. The distal edge ends exactly medially and is connected with the distal cingulum. The P3 is elongated and robust ( Table 1). The crown bears an elongated distal cingular projection. The small mesio-buccal prominence of the cingulum forms a faint mesial crest to the apex of the protoconid. This protoconid is located medially and slightly mesially. Two edges running from the protoconid apex are thick and sharp. The distal one is not connected with the distal cingulum, and its end forms a small, swellinglike bulge. The cingulum is weakly developed on the lingual side. The mesial and distal margins are blunt or rounded, while the buccal margin bears a moderately developed, median concavity. At the same level, but on the lingual margin, is located a moderate convexity ( Figure 2 View FIGURE 2 ).

The long and robust P4 has straight buccal and lingual margins of the talon ( Figure 2 View FIGURE 2 ). The mesial and distal margins are blunt or rounded. The moderately high paracone bears a thin crest across the mesial border from the apex to the base of the crown. It is separated from the moderately long and low protocone, whose mesial margin is aligned with that of the paracone. Its length is on average smaller than the distal breadth of the crown ( Table 2). The metacone is separated from the paracone by a deep valley. The cingulum is more strongly developed on the lingual margin of the metacone. The M1 is large; its breadth is smaller than the P4 length (Table 3; Figure 2 View FIGURE 2 ). The trigon is moderately wide and short, with a moderate and abrupt concavity of the buccal margin. The paracone and the metacone are elongated and well developed, where paracone is larger. They are well separated by a deep, narrow, V-shaped valley. Apexes of both main cusps are connected by a thin, long crest. The talon is shorter than the trigon and separated by a deep and broad depression running through the middle part of the crown. The protocone is low and long. It is divided by a shallow and wide valley into two parts of similar length and height. The reduced metaconule is not connected with any other cusp or crest, and is a low, elongated cuspid situated in the middle part of the crown. A long and thin crest corresponding to the buccal cingulum margin runs in parallel along the whole talon length. The well-developed lingual cingulum forms a thick crest ( Figure 2 View FIGURE 2 ).

The mandibular body is long and high, and its height measured behind the m1 is larger than the m1 length ( Figure 3 View FIGURE 3 ). The height of the mandibular body gradually increases distally. Two rounded mental foramens are moderately spaced and similar in size. The mesial one is situated under the p2, while the distal mental foramen is located under the distal root of p3, slightly lower than the mesial one ( Figure 3 View FIGURE 3 ). The masseteric fossa is moderately deep and its rounded, mesial edge reaches the m1/m2 boundary. The mesial part narrows dorso-ventrally and its ventral margin only slightly exceeds the midline of the mandibular body in dorso-ventral direction. The lower mandibular body margin forms a gently curved arch, uniformly domed mesially and distally, with the strongest curvature under m1. The symphysis part is massive and elongated.

The series of cheek teeth is almost straight and the premolars are located more buccally in relation to the molars. The premolars are tightly arranged and all teeth are situated at a similar level. The triangular crown of the i3 is double cusped, with the main protoconid occupying the larger surface. It is slightly asymmetrical, being more developed on the lingual side. Its vertical top is massive and rather blunt. The c1 is long and robust, with a proportionally elongated and hook-shaped crown ( Figure 3 View FIGURE 3 ). Two longitudinal grooves run on the buccal and lingual sides of the crown.

The relatively large p1 is an oval, small, and one-rooted tooth. It is tightly squeezed between c1 and p2. The two-rooted p2 is low-crowned, with the protoconid strongly displaced mesially. Its occlusal outline is almost rectangular, with an elongated distal part. Two thin crests run in the mesial and distal direction from the protoconid top. The distal cingulum forms a thin ridge, collaring the smooth area in the distal part. The mesial part of the crown bears an elongated distal cingular projection ( Figure 3 View FIGURE 3 ). The larger p3 has a similar outline in occlusal view, with straight buccal and lingual margins. The mesial and distal margins are blunt. The protoconid is also displaced mesio-medially, but less so than in p2. An elongated distal, cingular projection is oriented slightly disto-buccally. Two thin edges running from the protoconid apex. On the distal edge, a small tubercular convexity is present just behind the top. The mesial and distal cingulum are relatively strongly developed. The two-rooted p4 is relatively high-crowned and has the protoconid placed almost exactly centrally and pushed slightly mesially. A relatively large hypoconid is present after the protoconid. It is associated with the distal crest, running distally from the protoconid apex. The mesial ridge is thinner than the distal one. The mesial and distal halves of the tooth are equal in length. The mesial part is narrower than the distal one. The crown is slightly broadened in the distal direction. A gentle lingual convexity occurs in the middle part of the crown. The lingual margin is straight. The mesial margin is blunt, while the distal one is rounded. The mesial and distal cingulum are relatively strongly developed. The distal, cingular projection is less elongated compared to the rest of

MARCISZAK & KROPCZYK: CANIDS FROM NIEDŹWIEDZIA CAVE the crown. Collared by a thick cingulum, the inner surface of this projection is crescent-shaped and shallow ( Figure 3 View FIGURE 3 ).

The large m1 has a massive and high trigonid ( Figure 4 View FIGURE 4 ). The short and low talonid is narrower than the trigonid. The small metaconid is connected with the protoconid. A thin, longitudinal ridge, surrounding the talonid field, ending on the metaconid base, runs from the elongated and low hypoconid. The paraconid edge is weakly developed. The mesial margin is rounded, while the distal one is blunt. The buccal margin is almost straight, with gentle concavity on the transition between the trigonid and talonid. The lingual margin of the paraconid and half of the protoconid is straight, while more distally a moderate convexity is present. The cingulum is moderately developed. The m2 is moderately reduced. The crown has a slightly irregular, rounded occlusal outline. The large and low paraconid and protoconid are located on the trigonid. The larger paraconid is located mesio-buccaly, while the smaller, but not lower protoconid is situated medially and lingually. The talonid is narrower, with a conical and low hypoconid. The moderately developed cingulum is stronger only on the distal margin ( Figures 4–5 View FIGURE 4 View FIGURE 5 ).

The long and massive humeri have a flat and robust head ( Figure 6 View FIGURE 6 ). The greater tubercle extends distally along the side of the articulation surface. The distally directed neck and the smaller tubercle are well-developed. The lateral epicondylar crest is very prominent towards the lateral epicondyle from the last distal fourth. The medial epicondylar crest is prominent and has a narrow supracondylar foramen. The prominent rugose area occurs on the inner surface of the shaft for the attachment of the inner humeral head of the triceps muscle. The strong medial condyle is located slightly back, while the lateral condyle is pushed deeper in mesio-distal direction. A rounded and deep coronoid fossa does not connect with the wide and deep olecranon pit ( Figure 6 View FIGURE 6 ).

The radius is robust, flattened dorso-ventrally, and has a slightly concave ventral surface ( Figure 6 View FIGURE 6 ). The large head has a shallow and rounded articular surface. It is mesially collared by a prominent coronoid process. The articulation surface for the lesser sigmoid cavity is large. The distal end is widened laterally, with a well-marked elliptical glenoid cavity, framed by a rough condyle on ventral and medial sides. The facet for the ulna and the styloid process are well developed. The grooves for extensor muscle tendons, extensors carpi radialis longior and brevior muscles are broad and deep. The scapholunar articulation area is relatively short and broad ( Figure 6 View FIGURE 6 ).

The ulna is elongated and narrow, with a strong but short proximal end ( Figure 6 View FIGURE 6 ). The surface for attachment of the scapular head of the triceps muscle is broad and strongly rugose. The olecranon has a quadrilateral shape with its apex hollowed out in part of a small cavity bordered by two fairly prominent tubercles. The shaft is curved in the lateral and medial view and massive in its longitudinal extent. The articulating surface for the radius is large, while the articulating surface for the pisiform is quite narrow. The distal end is characterised by a significant protrusion of the ulnar styloid process ( Figure 6 View FIGURE 6 ).

Long, cylindrical femora have a spherical head with a prominent neck ( Figure 6 View FIGURE 6 ). The trochanteric fossa is broad and deep. The lesser trochanter is a slight conical prominence. The shaft is slightly bowed in its longitudinal extent. The greater trochanter rises distinctly above the head level and is obliquely truncated on its lateral side. The surface of the trochanter extends further down the proximal extremity of the femur. The tibial articulations are nearly equal in size. At the distal end, the condyles are separated by a broad and deep groove. The lateral condyle is slightly more developed as the medial one ( Figure 6 View FIGURE 6 ).

The body of long and massive tibiae are triangular in its proximal third, slightly convex medially, with a large tibial fossa from the lateral side ( Figure 6 View FIGURE 6 ). The distal surface below the head is narrow and shallow. The tibial crest is well marked but relatively short. At the distal end, the articular surface consists of two separate grooves. The medial throat is deeper than the lateral one. The grooves for tendons of the flexor longus digitorum and tibialis posticus muscles are well defined at the distal epiphysis. The notch incising the mesial border of the distal surface is quite small ( Figure 6 View FIGURE 6 ).

The calcanei are long and robust, with strongly developed tuber calcanei. The coracoid process is prominent and distinct from the sustentaculum tali. The medial part of the bone is flat and wide. Its proximal articular surface is concave whereas the distal articular surface is flat. The distal end of a tali has a well-marked neck and head. The body is flattened in the dorsoplantar direction. The trochlear margins are slightly asymmetrical. The lateral lip is slightly more prominent but shorter than the medial one.

Comparison. The comparison of various European populations, dated to the last 130 kya, showed considerable size variation within this period. Some significant differences were found in tooth proportions regarding the proportions of the length and width of the trigonid, but those from cold periods have more massive m1 overall ( Figures 1– 3 View FIGURE 1 View FIGURE 2 View FIGURE 3 ). From the Eemian, the length of m1 rarely exceeds 30 mm, and the specimens are characterised by moderate dimensions and stature. Since these wolves are metrically and morphologically indistinguishable from the Central European populations, they were classified as Canis lupus lupus . A sharp increasing of the size is observed since MIS 5d, with the first cold period. The period between MIS 5d–5c is poorly represented in Central European sites, but few specimens from dated on MIS 5d layer 12 of Biśnik Cave showed presence of robust form. Interesting similarity was found between British site Banwell Bone Cave and layers 10–9 of Biśnik Cave, dated on MIS 5b–5a. In this cold period, large wolves were found in both localities. The largest specimens were documented from the MIS 3, which is the best represented by the whole analysed time period. Later in the Holocene (MIS 1), a decrease in size is observed, and wolves from that period are comparable to that of MIS 5e ( Figure 8 View FIGURE 8 ). The numerous French or Italian sites with abundant wolf material

PALAEO- ELECTRONICA.ORG were not used for comparison, because wolves from contemporary Central European localities are significantly larger than those from the southern or western part of the continent.