Vulpes vulpes (Linnaeus, 1758)
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publication ID |
https://doi.org/10.26879/1510 |
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persistent identifier |
https://treatment.plazi.org/id/905687B3-FF90-FFE7-FEE0-FD1FFBC43407 |
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treatment provided by |
Felipe |
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scientific name |
Vulpes vulpes (Linnaeus, 1758) |
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Vulpes vulpes (Linnaeus, 1758) View in CoL
( Figures 9–10 View FIGURE 9 View FIGURE 10 ; Table 4; Appendix 1)
Referred material. The material ( NISP 35, MNI 5; left/right bones are given in brackets) includes one C1 (0/1), one P4 (1/0), two M1 (1/1), one M2 (0/1), eight mandibles (5/3), three c1 (3/0), three m1 (2/ 1), one scapula (1/0), one atlas, two humeri (1/1), one radius (0/1), two ulnae (1/1), two femora (2/0), one tibia (0/1), three calcanei (2/1), one mt 4 (0/1), and one phalanx 2.
Description. The elongated and robust P4 has a high and prominent paracone and a mesio-distally elongated metastyle. The moderately developed protocone is pushed more mesially than the mesial crown margin. The mesio-buccal cingulum is absent, and a small but sharply pointed parastyle is present. The disto-lingual cingulum is strongly developed. The M1 trigon is broadly expanded mesio-distally. The trigon basin is round and wide, and as deep as the talon basin, which is weakly compressed and semi-circular. There is a prominent shelf-like buccal cingulum that extends from the mesial wall of the paracone to the distal wall of the metacone. The paracone and metacone are similar in size, but the paracone is slightly higher. On the mesio-buccal side, there is a prominent parastyle. The well-developed and high protocone and the hypocone are separated from the lingual cingulum by a marked furrow. The metaconule is moderately high and prominent, the protoconule is low and small. The M2 trigon is mesio-distally less expanded than that in M1. The paracone is higher and larger than the metacone. The protocone is well developed and a small but well recognised metaconule is situated distally ( Figure 9 View FIGURE 9 ).
Length, L Breadth, B B/L ratio Tooth Locality Age M Min–Max M Min–Max M Min–Max N
C1 Silesia R 13.94 12.37-15.89
Pavlov, Předmostí 3–2 15.66 13.89-17.23
Niedźwiedzia Cave 3 16.43 14.91-17.43
Zoolithenhöhle 3 15.86 13.45-16.94
Jaurens Cave 3 13.95 11.60-15.40
Biśnik Cave, l. 7-5 3 16.24 14.86-117.18
Biśnik Cave, l. 10-9 5a 15.88 13.25-17.16
Biśnik Cave , l. 13 5e 14.28 11.39-17.14
Romain-la-Roche 6 12.44 10.30-13.80
c1 Silesia R 16.38 13.55-19.44
Niedźwiedzia Cave 3 18.68 16.22-20.78
Zoolithenhöhle 3 17.46 15.56-20.15
Jaurens Cave 3 14.22 12.00-16.10
Biśnik Cave, l. 7-5 3 17.79 14.45-20.23
Biśnik Cave, l. 10-9 5a 16.14 13.88–18.57
Dziadowa Skała, l. 3- 4 5e 13.74 12.96–14.78
Biśnik Cave , l. 13 5e 13.45 12.49–14.23
P2 Silesia R 15.38 13.69–17.14
Pavlov, Předmostí 3–2 16.14 14.31–17.79
Niedźwiedzia Cave 3 16.63 14.69–18.64
Zoolithenhöhle 3 16.27 14.45–18.04
Jaurens Cave 3 15.75 15.00–16.20
Biśnik Cave, l. 7-5 3 16.59 14.78–18.84
Biśnik Cave, l. 10-9 5a 15.43 13.82–17.98
Biśnik Cave , l. 13 5e 15.04 13.38–16.18
Romain-la-Roche 6 14.01 11.8–16.50
p2 Silesia R 13.41 11.55–16.21
Niedźwiedzia Cave 3 14.92 13.78–15.88
Pavlov, Předmostí 3–2 14.66 12.16–15.65
Zoolithenhöhle 3 14.42 12.56–15.84
Jaurens Cave 3 12.66 10.80–14.00
Biśnik Cave, l. 7-5 3 14.88 13.65–16.04
Biśnik Cave, l. 10-9 5a 13.97 12.72–15.14
Banwell Bone Cave 5a 13.02 11.22–14.00
Biśnik Cave , l. 13 5e 13.34 11.47–15.66
P3 Silesia R 16.99 15.74–19.29
Pavlov, Předmostí 3–2 17.54 15.82–19.97
Niedźwiedzia Cave 3 18.76 17.37–20.27
Zoolithenhöhle 3 18.04 14.97–20.09
Jaurens Cave 3 17.03 16.30–17.50
Biśnik Cave, l. 7-5 3 17.98 15.66–19.41
Biśnik Cave, l. 10-9 5a 17.84 15.97–18.93
7.91 7.56-9.59 63.3 58.8-69.1 40
9.48 8.14-12.04 67.1 60.9-72.1 22
9.97 8.29-12.21 59.8 52.8-68.8 17
9.52 7.97-11.94 66.9 56.7-71.8 18
8.96 7.70-9.90 66.3 59.2-79.2 8
9.57 8.59-11.97 63.4 56.7-68.8 8
9.22 8.22-11.84 66.1 62.2-72.4 12
8.79 6.96-10.66 61.8 58.3-66.9 9
7.72 6.40-8.90 62.1 59.7-64.5 43
10.44 8.56-11.89 65.6 57.7-76.9 38
11.27 9.09-13.45 60.2 53.1-67.9 16
10.97 8.89-13.16 66.9 57.9-72.6 10
9.17 8.40-9.90 64.6 55.5-71.7 12
11.14 8.97-13.26 66.4 58.7-69.7 7
10.21 8.73–11.74 63.2 59.6–66.8 7
9.38 8.84–10.15 61.9 63.6–68.7 5
8.51 7.86–9.14 63.2 58.2–68.6 8
6.41 5.37–7.39 41.7 36.4–47.4 40
6.69 5.64–8.44 42.2 37.7–48.6 23
7.31 5.75–8.72 43.8 38.4–49.4 11
7.04 5.48–8.29 43.1 36.6–48.7 19
6.88 6.60–7.30 43.3 41.0–45.6 4
6.97 5.69–8.66 42.7 38.4–46.9 9
7.15 6.17–8.71 46.5 41.4–56.7 13
6.34 5.62–6.94 42.1 38.7–49.5 8
6.17 5.40–7.00 44.1 42.4–45.8 31
6.45 5.48–7.78 48.2 41.9–53.1 40
7.39 6.55–7.97 49.6 47.3–51.9 11
7.43 6.63–8.04 52.9 44.8–56.8 34
7.19 6.38–7.96 49.8 45.7–54.3 18
6.23 5.60–6.90 49.7 44.4–59.3 10
7.44 6.42–8.06 52.4 47.9–56.2 11
7.15 6.62–7.67 51.6 49.7–55.6 13
6.75 5.93–8.00 51.4 44.3–59.3 7
6.89 5.78–7.91 50.9 45.6–55.2 8
7.09 6.16–8.17 41.8 37.1–47.3 40
8.03 7.36–8.78 46.2 40.8–51.6 14
8.79 7.75–9.66 46.8 43.2–48.8 12
8.31 6.54–9.43 42.9 37.1–52.1 23
7.49 6.80–8.00 43.9 41.7–45.7 6
8.64 7.44–9.21 46.4 43.5–50.5 11
8.56 7.53–9.76 47.6 43.5–51.9 13
The mandibular corpus is deep and stout, becoming medio-laterally thicker ventrally. Its height gradually increases distally. Two mental foramina are located below the p1 (mesial, larger) and below the mesial root of the p3 (smaller, distal) that emerges more dorsally than the former ( Figure 9 View FIGURE 9 ). The rounded mesial edge of the deep masseteric fossa reaches the m3. 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, with the strongest curvature under m1. The symphysis part is elongated and moderately robust. The cheek teeth row is almost straight, and the premolars are located more buccally in relation to the molars. The premolars are loosely arranged and separated by small diastemas.
The c1 is long and robust, with a proportionally elongated and hook-shaped crown ( Figure 9 View FIGURE 9 ). Two longitudinal grooves run on the buccal and lingual sides of the crown. The relatively large p1 is an elongated, oval, small, and one-rooted tooth. The two-rooted p2 is high-crowned, with the protoconid strongly displaced mesially. The crown is elongated and narrow, gently convex buccally and with weak median convexity. It has an elongated distal cingular projection. After the protoconid is located slightly disto-buccally a minute cuspid, which in some specimens is weakly marked only as a ridge. The elongated and narrow p3 gently widens distally. It has straight buccal and lingual margins, while the mesial and distal margins are blunt. The protoconid is also displaced mesio-medially, however less than in p2. An elongated distal cingular projection is oriented more disto-buccally. On the distal edge, a small tubercular convexity is present just behind the top. The mesial and distal cingulum is weakly developed. The high-crowned p4 has the protoconid strongly displaced mesio-medially. A prominent cuspid is present after the protoconid. It is associated with the distal crest, running distally from the protoconid apex. The crown is slightly broadened in the distal direction, with straight buccal and lingual margins. The mesial and distal margins are blunt. The distal cinminimal gular projection is less elongated in comparison to the p2–p3 and collared by a thick cingulum. The inner surface of this projection is crescent-shaped and shallow.
The elongated and robust m1 has a massive and high trigonid and proportionally long and low talonid, slightly narrower than the trigonid. The paraconid is low and short. The large and trapezoidal metaconid is moderately distinct from the high and prominent protoconid. The mesial margin is rounded to triangular, and the distal one is blunt. The lingual margin is straight, while the buccal one holds a strong concavity on the transition between the trigonid and talonid. The cingulum is moderately developed. The hypoconid is large and high, while the entoconid is lower and smaller. The m1 is bucco-lingually stout and high, especially at the level of the trigonid, whereas the protoconid is stout and large. The talonid basin is round, smooth, and wide, partially enclosed lingually by the entoconulid. Distally, there is a prominent cingulid ( Figure 9 View FIGURE 9 ).
The broad and large m2 has a slightly irregular, rounded occlusal outline. On the trigonid are located mesio-buccaly a larger and higher protoconid and medio-lingually a lower and smaller metaconid. The talonid is narrower, with a conical and low hypoconid and a distinctly lower and smaller entoconid. A small but well-recognised mesoconid is also present. Before the entoconid, which developed an inner edge, is also present a small proentoconid. The moderately developed cingulum is stronger only on the distal margin. The relatively weakly reduced m3 has oval or rounded outline and bears two equally sized cusps, the protoconid and the metaconid. The talonid is preserved as a semi-circular ridge ( Figure 9 View FIGURE 9 ).
Comparison. The analysed material of Vulpes vulpes from Niedźwiedzia Cave is a good example of great metrical and morphological variability of this species (Lucenti and Madurell-Malapeira, 2020). The single P4 (JN.2.5) has a narrow mesial margin of the paracone, typical for the species. However, the morphology of this structure is highly variable, and this part can be reduced or, on the contrary, expanded. The enlarged and separated protocone dominated in V. vulpes . Nevertheless, in some P4, it is reduced or fairly prominent and partially associated to the paracone, like in JN.2.5. The mesial embayment of P4 (JN.2.5) is moderately developed, and strongly vary in V. vulpes , where it can be narrow, wide, or even absent. Vulpes vulpes generally does not possess cingulum on the mesio-buccal side of the P4, but specimens with fairly to well-developed cingula are not uncommon, like that in JN.2.5. The occurrence of a distinct parastyle is rare in V. vulpes and V. corsac and more common in V. lagopus (Gimranov, 2017; Lucenti and Madurell-Malapeira, 2020). It is also absent in P4 from Niedźwiedzia Cave. Some P4 of V. vulpes possesses an accessory cuspule on the preparacrista, located just above the mesial embayment. Such structure is also present in JN. 2.5 in a form of minute, but well-recognised ridge.
Characteristic for Vulpes vulpes is that the medial protocone crest is fused to the preparacrista, but in some specimens it is incomplete or curved distally not reaching it. Generally, the morphology of the medial protocone crest and its relation to the preparacrista is highly variable in this canid (Lucenti and Madurell-Malapeira, 2020). In JN.2.5, this structure is straight, projecting medially toward the lower third of the preparacrista, and this type of morphology dominated in V. vulpes . Much less commonly occurs an arched medial protocone crest, with parabola-like shape, which extends linguo-medially and then ventrally toward the apex of the crown. Typical for V. vulpes , which is also observed in JN.2.5 tooth, is the protocone collared by a prominent cingulum, which is sometimes incomplete and rarely absent. The P4 paracone is longer than the metastyle, like in JN.2.5, although in some specimens they are comparable in length. The disto-lingual cingulum in JN.2.5 is moderately marked. The stage of development of this feature is highly variable, from nearly absent to particularly strong.
The occlusal outline of the M1 is highly variable, especially the proportion and stage of development between the trigon and talon. In JN.2.5, the M1 has a particularly broad and expanded trigon, T-shaped in the occlusal view, with proportionally narrow and short talon. The M1 embayment on the buccal cingulum is strongly marked, while in Vulpes vulpes varies from reduced to strongly developed. The JN.2.5 has an expanded buccal cingulum on the metacone, where the hypocone lobe connects mesially to the mesial cingulum. The bean-shaped M2 has a moderately expanded trigon and possesses a distinctly marked notch, situated lingually to the metacone. A strong median notch is also located in the buccal margin.
All p4 from Niedźwiedzia Cave have the distal portion slightly elongated and enlarged distally, with a rounded outline, typical for the species. Of the 10 m 1 from Niedźwiedzia Cave, eight are stout and enlarged in bucco-lingual direction. Two other teeth are bucco-lingually compressed, narrow, and smaller. The morphology of the m1 from Niedźwiedzia Cave is highly variable, similar to that of the extant Vulpes vulpes . Of the 10 m 1 from Niedźwiedzia Cave, seven have reduced or weakly developed inflexion on its lingual side of the protoconid and the area is nearly convex. In another three teeth, this structure is stronger developed. The metaconid is well developed, prominent, and well separated from the protoconid in nine m1 from Niedźwiedzia Cave, while only a single specimen possesses a more reduced and less separated metaconid. The presence of a transverse cristid is common for the extant V. vulpes , as in the material from Niedźwiedzia Cave, where this structure is present in all 10 teeth analysed. In six m1, this feature is strongly marked, while in four others is weakly to moderately developed. Very variable is also a constellation of accessory lingual cuspulids, located mesially to the m1 entoconid on the m1 talonid. In eight teeth, the presence of a reduced (n = 5) or an enlarged (n = 3) entoconulid was found, while in two others the entoconulid was present with the addition of a mesial accessory cuspulid. No m1 from Niedźwiedzia Cave with the absence of cuspulids mesial to the entoconid have been found. Another highly variable feature is the morphology of the distal margin of the m1. In three specimens, the margin is a simple cristid bounding the distal margin, while in three others distal accessory cuspulids are located in place of a cingulid. In single specimens this structure is developed into a form of a distal cristid arising from an evident hypoconulid, as an enlarged hypoconulid with no distal cristid, a strongly reduced one, or a reduced distal cristid or no cristid.
In three m1 from Niedźwiedzia Cave, the m2 has a more ovoid shape, with moderately to strongly marked hypoflexid and buccal median concavity. This structure is not present in the other two teeth, and their occlusal outline is more bean-
Vulpes vulpes Vulpes lagopus Parameter Sex M Min–Max N M Min–Max N
humerus
GL ♂♂ 154.6 124.8–164.6 ♀♀ 126.7 115.6–134.8
pL ♂♂ 27.4 22.8–32.7
♀♀ 23.3 16.9–26.2
dB ♂♂ 24.6 18.9–29.7
♀♀ 20.4 19.6–21.1
GL ♂♂ 144.6 119.7–163.9
♀♀ 116.9 108.1–143.7 pL ♂♂ 13.4 10.7–14.5 ♀♀ 11.2 8.7–12.9
dB ♂♂ 17.4 14.6–19.1 ♀♀ 14.4 13.7–16.6
GL ♂♂ 148.88 124.84–166.97 ♀♀ 126.66 117.84–134.67 pL ♂♂ 28.97 26.56–34.38
♀♀ 24.66 23.19–26.64 dB ♂♂ 23.79 21.45–25.56 ♀♀ 20.32 19.34–21.87
GL ♂♂ 167.97 146.79–175.74 ♀♀ 141.45 126.78–156.56
pL ♂♂ 25.67 22.78–32.34
♀♀ 21.84 19.97–25.39
dB ♂♂ 18.74 16.45–25.27
♀♀ 14.88 12.59–20.37
GL ♂♂ 35.54 30.29–40.44
♀♀ 31.53 27.54–35.56
GL ♂♂ 21.94 18.73–24.45
♀♀ 19.56 17.24–21.56
like. Similarly variable is the morphology of a buccal cingulid situated on the mesio-buccal side of the protoconid. In four m2 from Niedźwiedzia Cave, it is reduced to moderately developed, limited to the mesial part of the protoconid. Only a sole specimen possesses an enlarged buccal cingulid that prominently extends distally on the buccal side of the hypoconid. In single specimens, the
37 114.4 103.6–124.7 34
31 101.6 90.4–111.8 28
37 20.4 16.7–24.1 34
31 17.4 14.9–19.4 28
37 18.4 16.6–20.6 34
31 17.1 14.8–18.8 28
radius
37 104.9 97.8–116.6 34
31 93.2 85.6–104.2 28
37 10.9 7.4–12.2 34
31 9.9 6.6–10.9 28
37 14.2 13.4–16.8 34
31 12.9 11.66–14.78 28
femur
37 111.9 102.6–122.4 34
31 100.2 93.7–108.7 28
37 23.41 21.56–24.48 34
31 20.44 18.89–21.79 28
37 19.77 17.24–20.44 34
31 17.66 15.52–18.89 28
tibia
37 129.76 115.64–145.81 34
31 116.78 104.69–126.45 28
37 20.88 17.87–24.45 34
31 19.64 15.86–21.44 28
37 14.97 13.11–21.22 34
31 13.92 12.68–18.64 28
calcaneus
37 28.97 26.32–32.34 34
31 26.91 25.54–29.56 28
talus
37 17.97 15.93–19.97 34
31 16.04 15.06–17.56 28
m2 possess the entoconid as a single cuspid or a very reduced one, while the rest of the specimens show a large entoconid and a mesial accessory cuspulid ( Figure 9 View FIGURE 9 ). All three m3 from Niedźwiedzia Cave are small and rounded in occlusal view, while in the extant V. vulpes this tooth is often large and rounded or large and oval (Lucenti and Madurell-Malapeira, 2020). The buccal protoconid and the lingual metaconid are generally subequal in size, while the protoconid can be larger than the metaconid. In some specimens of the extant V. vulpes the metaconid is absent. Instead, those teeth have a single buccal cuspid and a lingual cristid or a single central cuspid (Lucenti and Madurell-Malapeira, 2020).
Despite the great variability of Vulpes vulpes from Niedźwiedzia Cave, we did not find any differences between the fossil material from Niedźwiedzia Cave and extant Polish populations. While no particular morphological features distinguishing the Niedźwiedzia Cave material and the extant V. vulpes have been found, there are some interesting metric patterns. Firstly, the red fox from this locality is characterised by considerable size, exceeding the average size of the extant Silesian population, even if their ranges of variation overlap. It is well documented, when compared the L m1 of the material from Niedźwiedzia Cave (17.19 mm, 16.31–17.88 mm, n = 6 in males and 14.75 mm, 14.22–15.22 mm, n = 4 in females) with the extant Silesian V. vulpes (16.32 mm, 14.78–18.16 mm, n = 114 in males and 14.48 mm, 12.66–15.56 mm, n = 108 in females). Documented sexual dimorphism is the second factor, typical for the carnivores (Dayan et al., 1989). The result of this pattern is a separation of food niches of particular species and a consistent trend for females to consume smaller preys than males (Pimm and Gittleman, 1990). Across carnivores there is a general tendency for carnassials to be less dimorphic than canines. Canids display less specialised killing behaviour than, for example, mustelids, the canines are 8– 10% larger in males, while the m1 is larger by 5– 6% (Gittleman and Van Valkenburgh, 1997; Szuma, 2000, 2004).
Such great metric and morphological variability is characteristic for Vulpes vulpes (Szuma, 2000, 2003, 2004, 2007, 2008a, 2008b), while is less pronounced in other foxes (Szuma, 2008c, 2011; Gimranov et al., 2015; Gimranov, 2017; Lucenti and Madurell-Malapeira, 2020). This variability results from the combination of biological, climatic, and geographic factors. Among them latitude, habitat productivity, differential food availability, intraguild, intrafamily or intraspecific competition, character displacement, genetic diversity, and population density are the most important (Szuma, 2008b; Szuma and Germonpré, 2020). Different Vulpes species are susceptible to different combinations of these proxies, with resulting different variability and features (Szuma, 2011). For V. vulpes , latitude, mean annual temperature, and longitude significantly affect the distribution of morphotypes (Szuma, 2007). Instead, geographic and climatic factors are less important for V. lagopus , which is more influenced by its interspecific competition with V. vulpes and food source variability and accessibility (Szuma, 2011; Szuma and Germonpré, 2020). Additionally, the geographic range of the extant V. vulpes is enormously larger and more diverse in terms of habitat conditions and types, compared to that of V. lagopus , which affects the importance of the above-mentioned factors (Szuma 2007, 2008b, 2011; Lucenti and Madurell-Malapeira, 2020).
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