4.a.1. Upper second premolar (P2)

Material. IMEDEA 90142, left (Fig. 2a).

Figure 2.

Dental material of Myotragus palomboi, n. sp. (a) left P2 IMEDEA 90142: 1 – lingual view; 2 – mesial view; 3 – labial view; 4 – distal view; 5 – occlusal view; (b) left P3 IMEDEA 90275: 1 – lingual view; 2 – mesial view; 3 – labial view; 4 – distal view; 5 – occlusal view; (c) left i3 IMEDEA 90145: 1 – occlusal view; 2 – mesial view; 3 – labial view; (d) right dc IMEDEA 90146: occlusal view; (e) left p2 IMEDEA 90143: 1 – labial view; 2 – lingual view; 3 – mesial view; 4 – distal view; 5 – occlusal view; (f) right p2 IMEDEA 90144: 1 – labial view; 2 – lingual view; 3 – mesial view; 4 – distal view; 5 – occlusal view. Scale bar 2 cm.

A very hypsodont tooth with subequal length and width in occlusal view. The tooth is slightly wider than long (L = 6.5 mm, W = 6.7 mm; see Table 1), and the labial wall is slightly concave, almost flat, with styles poorly developed or absent. No ribs on the labial wall can be observed. The parastyle is almost absent because it is extremely reduced, appearing as a small rib on the mesial side of the tooth. This weak parastyle is placed very close to the mesial extreme of the central cavity, which is mesio-distally elongated and placed on the mesial half of the tooth. This tooth is poorly developed mesially to the protocone, while distally it is large and expands lingually. The enamel is more extended on the mesial and labial walls than on the distal and lingual walls. The occlusal surface is worn and almost flat. The distal edge on the occlusal surface is sharp and divided into two lobes.

As remarked previously, the tooth of the CdR caprine is wider than long (ratio L/W < 1), as in specimens of M. pepgonellae, Ovis vignei and Capra pyrenaica (P. Bover, unpub. data). The remaining fossil and extant caprines display a P2 which is longer than wide.

A partial maxilla of M. pepgonellae (MNIB 59199) bearing P2–M3 shows that the P2 displays the same morphology as in the CdR caprine. According to the description of the P2 of Norbertia hellenica, styles are absent and the tooth is longer than broad (Köhler, Moyà-Solà & Morales, Reference Köhler, Moyà-Solà and Morales1995). This morphology is similar to that displayed by the CdR caprine, M. pepgonellae and Gallogoral, although it might be related to the fact that these remains share the same level of wear. Like Norbertia, Aragoral mudejar has a longer than broad P2 (Alcalá & Morales, Reference Alcalá and Morales1997) and, as observed in the CdR caprine, its mesial side is reduced (area placed mesially to protocone poorly developed). No P2 of Nesogoral appears described or figured in the bibliography, and no P2 from Ses Fontanelles is available for study.

Although the morphology of this premolar is conditioned by wear, the shape in occlusal view of the Caló den Rafelino P2 is more similar to Ammotragus, Ovis and Capra than to other caprines. Differences among these taxa rely on the lesser development of the area placed mesially to the protocone in the CdR caprine and Ammotragus, and the presence of a labial projection of a feeble metastyle in Capra and Ovis.

4.a.2. Upper third premolar (P3)

Material. IMEDEA 90275, left (Fig. 2b).

Although the tooth is highly worn, it is clearly hypsodont. As in P2, the P3 is wider than long (L = 8.3 mm; W = 10.8 mm; see Table 1). In occlusal view, it has a square-trapezoidal shape. The parastyle is very reduced and placed on the mesio-labial corner, and is expanded labially. The labial wall is also relatively flat and only the paracone prolongation to the root through the wall can be observed. The area placed mesially to the protocone is well developed and expanded lingually. The area placed distally to the protocone is poorly developed. The enamel is more expanded on the labial and medial walls. The central cavity is elongated and U-shaped. The medial extreme of this cavity is placed close to the parastyle, between the parastyle and the paracone. As in P2, the distal crest present on the occlusal surface is well marked and sharp. The occlusal surface is almost flat with a broad groove running in the lingual–labial direction.

A wider than long P3 (ratio L/W < 1) is also displayed by M. pepgonellae (Moyà-Solà & Pons-Moyà, Reference Moyà-Solà and Pons-Moyà1982), Gallogoral (Guerin, Reference Guerin1965) and some specimens of Capra and Ovis (P. Bover, unpub. data). Although this tooth is more hypsodont in M. pepgonellae, its morphology is very similar to the CdR caprine. Norbertia hellenica and Aragoral mudejar have a weak metastyle (like the CdR caprine), but they display a flat lingual surface, while in the CdR caprine this surface is strongly convex. Gallogoral and Nesogoral have a well-developed metastyle, but Nesogoral has a weaker parastyle than Gallogoral, although it is more developed than in the CdR caprine.

Although the shape of the occlusal surface also seems to be influenced by wear, similarities in development of the protocone and parastyle between the CdR caprine and specimens of Ovis, Capra and Pseudois with similar tooth wear can be clearly noticed.

4.b.1. Third incisor (i3)

Material. IMEDEA 90145, left (Fig. 2c).

This tooth has a typical chisel shape and displays a clear distinction between root and crown. It is not a very hypsodont tooth (crown length 9.5 mm; crown maximum width 4.6 mm; root length 8.2 mm). The root is compressed mesio-distally and displays a shallow groove along the mesial side. The crown is labially curved, especially in ventral view. The wear surface has a subrectangular-oval shape. It is divided into two concavities by a medial rib and has a distinctive wear facet on the tip (labial extreme). The enamel has no longitudinal grooves and covers the entire crown region (except the wear surface).

In bovids, incisors and canines (incisiforms) usually fall from the mandible shortly after death. It is rare to get jaws retaining incisiforms in fossil specimens. When more than one species is present in the same deposit, it is difficult to assign isolated incisors and canines to one of them. Thus, for comparative purposes, only incisors and canines from recent caprines have been used, together with some teeth of Myotragus pepgonellae.

Although it depends on the wear conditions, the shape of this tooth in the CdR caprine is similar to the i3 of all caprine species studied except Myotragus. The general shape of the tooth is even more similar to Capricornis, Nemorhaedus and Rupicapra.

The i3 of M. pepgonellae displays a very different shape. It has a wear surface subtriangular in outline and is relatively hypsodont (Hypsodonty Index, HI: 2.12–2.22, calculated as crown total length/maximum crown width), but it is less hypsodont than the i3 of M. antiquus (HI: 3.1) (Moyà-Solà & Pons-Moyà, Reference Moyà-Solà and Pons-Moyà1982). The HI for the i3 of the CdR caprine is 2.06, and its shape is completely different (see above).

4.b.2. Deciduous canine or third incisor? (dc or di3?)

Material. IMEDEA 90146, right (Fig. 2d).

Crown of a small incisiform, probably a deciduous canine. Crown length 5.5 mm; crown maximum width 2.7 mm. The outline of the crown is oval.

4.b.3. Lower second premolar (p2)

Material. IMEDEA 90143, left, 90144, right (Fig. 2e, f; Table 1).

This is not a very hypsodont tooth (Table 1). Entoconid and entostylid are fused (entoconid–entostylid complex; fusion observed even in an unworn tooth, IMEDEA 90143). Paraconid and parastylid are also fused (paraconid–parastylid complex). The labial wall is almost completely flat, while there are two slightly marked ribs on the lingual wall. They correspond to the ventral expansion of the paraconid–parastylid complex and the metaconid. The paraconid–parastylid complex is small and reduced to a crest on the mesial margin of the tooth. The protoconid is just in front or placed slightly mesial to the metaconid. In the worn p2 (IMEDEA 90144) the metaconid and protoconid are the highest points of the tooth and are placed on the mesial margin of the occlusal surface. Thus, the occlusal surface is flat and obliquely inclined towards the distal part. In one of the specimens (IMEDEA 90143), two large roots can be observed.

In M. pepgonellae the presence of a second lower premolar remains controversial. In the description of the species by Moyà-Solà & Pons-Moyà (Reference Moyà-Solà and Pons-Moyà1982), the presumed presence of an alveolus for a reduced p2 was reported, but Bover & Alcover (Reference Bover and Alcover2005) mentioned that it was not observable. Thanks to a recent exhaustive cleaning of the type of M. pepgonellae (MNIB 57321) and of a second mandible (MNIB 59196) of this species, both completely covered by an opaque layer of consolidant, the root of a small tooth was revealed mesially to the p3. It is unclear if it corresponds to a p2 or to a relict dp2. The size of this premolar should be extremely reduced in relation to a ‘normal’-sized premolar (alveolar length: 1.9 mm, 2.5 mm; alveolar width: 2 mm, 2.9 mm, respectively, in the two mandibles), and its shape should have looked like a small column (like the dp3 of M. balearicus; see Bover & Alcover, Reference Bover and Alcover1999). The morphology of this tooth in the CdR caprine is more similar to the p2 of other fossil and recent caprines studied. Unfortunately, the unavailability for study of other premolars of the CdR caprine precludes the establishment of comparisons between the size of the p2 and the size of the p3 and p4, and thus of evolutionary changes or size reduction in this tooth. Nevertheless, the p2 is proportionally wider in the CdR caprine (W/L: 0.75 for IMEDEA 90143 and 0.95 for IMEDEA 90144; see Table 1) than in other caprines studied (W/L ranging between 0.55 and 0.67 except in very worn specimens, in which this ratio is higher). In other caprines studied (except Aragoral; p2 not figured in Alcalá & Morales, Reference Alcalá and Morales1997) the protoconid and metaconid are also the highest points of the tooth, but they are placed in the middle of the occlusal surface.

4.c.1. Magnum

Material. IMEDEA 90272, right, and 90273, right (Fig. 3a, b).

Figure 3.

Postcranial material of Myotragus palomboi, n. sp. (a) Right magnum IMEDEA 90272: 1 – proximal view; 2 – distal view; 3 – palmar view; (b) right magnum IMEDEA 90273: 1 – proximal view; 2 – dorsal view; (c) left pisiform IMEDEA 90138: 1 – medial view; 2 – dorsal view; (d) left astragalus IMEDEA 90131: 1 – dorsal view; 2 – plantar view; 3 – proximal view; 4 – distal view; (e) left cubonavicular IMEDEA 90141: 1 – proximal view; 2 – distal view; 3 – dorsal view; (f) right metatarsal (holotype) IMEDEA 90140: 1 – dorsal view; 2 – medial view; (g) proximal phalanx IMEDEA 90132: 1 – dorsal view; 2 – axial view; (h) proximal phalanx IMEDEA 90133: 1 – dorsal view; 2 – abaxial view; (i) proximal phalanx IMEDEA 90274: 1 – dorsal view; 2 – axial view; (j) medial phalanx IMEDEA 90134: 1 – dorsal view; 2 – abaxial view; (k) medial phalanx IMEDEA 90135: 1 – dorsal view; 2 – abaxial view; (l) distal phalanx IMEDEA 90136: 1 – proximal view; 2 – solar view; 3 – axial view; 4 – distal view; (m) distal phalanx IMEDEA 90137: axial view. Scale bar 2 cm.

Like other postcranial remains, this bone is robust. It is slightly proximo-distally compressed, and the insertion areas of ligaments and tendons are poorly preserved, as occurs with the crests present on the articular facets. The separation between the semilunar and scaphoid articular facets is relatively low in dorsal view. The distal surface is completely flat and, while in specimen IMEDEA 90272 the articular facet for the metacarpal is expanded over all the surface, in IMEDEA 90273 there is a non-articular facet on the latero-palmar part of this surface.

One of the specimens (IMEDEA 90273) displays gnawing marks on the proximo-medial and proximo-dorsal margin of the bone, probably produced by a rodent (a cricetid and a glirid have been recorded in the deposit).

The only remarkable morphological feature of the magnum of the CdR caprine is the relatively low height difference between the semilunar and scaphoid articular facets. The ratio medial height/lateral height (ACM/ACm; see Table 1) is low in the CdR caprine (mean = 1.16; n = 2; range = 1.11–1.21). Only M. kopperi (mean = 1.22; n = 2; range = 1.17–1.27) (no magnum of M. pepgonellae or M. antiquus is available), Budorcas (mean = 1.24; n = 3; range = 1.15–1.28) and Ovibos (mean = 1.18; n = 5; range = 1.12–1.21) show similar values. This ratio is markedly higher in the other species studied (e.g. Oreamnos; mean = 1.5; n = 5; range = 1.41–1.59) and Nesogoral (1.35; n = 1, obtained from figured specimen in Van der Made, Reference Van der Made, Alcover and Bover2005).

4.c.2. Pisiform

Material. IMEDEA 90138, left (Fig. 3c).

The pisiform of the CdR caprine is robust and of globular shape. The articular facet for the unciforn and the ulna are separated by a ridge. For comparison only pisiforms of M. balearicus, Gallogoral, Oreamnos, Capra, Ovis, Rupicapra, Capricornis, Budorcas and Ovibos have been used. Although some features of the pisiform are very variable (Bover, Fornós & Alcover, Reference Bover, Fornós, Alcover, Alcover and Bover2005), the general globular shape of this bone in the CdR caprine is very similar to M. balearicus, Oreamnos and Ovibos.

4.c.3. Astragalus

Material. IMEDEA 90131, left (Fig. 3d).

The astragalus is incomplete and partially eroded, but some of its features can be described. It is short and broad. The plantar articular facet for the calcaneus is concave but not very rounded and is extended distally. The groove placed between the two proximal trochleae is broad and deep, and in dorsal view it is closed U-shaped. In this same view, and although it is slightly eroded, the lateral proximal trochlea is placed above the level of the medial one, which is expanded plantarly.

Astragali from recent caprines Budorcas, Oreamnos, Capricornis, Ovibos, Ovis and Capra and fossil Myotragus, Nesogoral and Gallogoral have been compared with the astragalus of the CdR caprine. The caprine astragali are generally short, but this bone is not significantly more robust in the CdR caprine. Values for the ratio Distal Breadth/Lateral Greatest Length (Bd/GLl × 100; measurements following von den Driesch, Reference von den Driesch1976) reach 58.5 in the CdR caprine (Table 1) and 61.8 in M. pepgonellae (n = 1) (in M. antiquus even more slender; mean = 57; n = 3; range = 55.4–58.1). In the rest of the caprines studied, this value is larger than 65 except in Nemorhaedus goral (mean = 61.2; n = 3; range = 59.2–62.4), Nesogoral (63.6) and Gallogoral (56.2).

Additionally, resemblances between the CdR caprine and Myotragus rely on the shared deep, slender and closed U-shaped outline of the groove placed between the proximal trochleae when seen in dorsal view, and the not very rounded plantar articular facet for the calcaneus. In other caprines this groove is more V-shaped (Budorcas, Capra, Ovis, Nesogoral) or open U-shaped (Capricornis, Oreamnos, Nemorhaedus, Ovibos, Gallogoral) and the facet for the calcaneus is clearly rounded (P. Bover, unpub. Ph.D. thesis, Univ. Illes Balears, 2004).

4.c.4. Cubonavicular

Material. IMEDEA 90141, left (Fig. 3e).

This bone shows the dorsal margin fragmented. It has a square outline in proximal view and is slightly compressed proximo-distally. All attachment areas of ligaments are robust. The medial and central processes are partially eroded. In plantar view, the attachment area of the plantar ligament is large and rounded. A longitudinal groove located centrally along the plantar margin can be observed. Although it is partially abraded, the postero-external metatarsal process is blunt and robust. In distal view, it displays a deep and narrow groove to accommodate the tendon of the musculus peroneus longus that continues obliquely along the lateral margin of the bone and contacts the calcaneal articular facet. The antero-external metatarsal facet and the entocuneiform facet are rounded and concave, the former being almost half the size of the latter. The posterior cuneal facet is flat and very small. The postero-external metatarsal facet is also rounded. The calcaneal articular facet is broad medio-laterally and the internal and external astragal articular facets are deep and very concave, with a sharp crest between them. There is a deep and elongated notch on the planto-lateral part of the internal astragal articular facet.

Characters shared by the CdR caprine and Myotragus are the robust attachment area for ligaments, the general proximo-distal compression of the bone, the shape of the distal articular facets, and the similar level at which the antero-external metatarsal facet and the entocuneiform facet are placed when seen in dorsal view.

4.c.5. Metatarsal

Material. IMEDEA 90140, right (Fig. 3f)

The metatarsal is a very short and wide bone (Fig. 4). It is more compressed dorso-plantarly than transversally. The plantar surface of the diaphysis is fairly flat and wide, and a shallow plantar longitudinal groove can be observed on the proximal and distal extremes, close to the epiphyses. The dorsal surface is also almost flat, and the dorsal longitudinal groove, wide and deep, can just be observed on the distal 1/3 of the diaphysis, especially on the area close to the intertrochlear incision.

Figure 4.

Metatarsals in dorsal view of (a) Myotragus palomboi n. sp. IMEDEA 90140 (holotype); (b) Myotragus pepgonellae MNIB 59204 and (c) Myotragus balearicus IMEDEA 85651. Dashed lines indicate the anatomical separation of metatarsal and cubonavicular (fused in the more modern species of Myotragus). Scale bar 2 cm.

In dorsal and plantar views, the transverse diameter is practically the same along all the diaphysis (that is, the lateral and medial margins of the diaphysis are parallel).

The proximal articular surface is almost flat, and the proximal metatarsal groove is deep, mainly at its central part. The posterior subfacet of the cubonavicular facet is abraded and does not allow us to conclude if it is high. The other facets and projections are also eroded. The morphology of the insertion of the lateral and medial extensor tendons cannot be fully resolved.

Although it is poorly preserved, the articulation surface of the trochlea is not as restricted as in Myotragus (Leinders, Reference Leinders1979).

The index of metatarsal robustness (TD/GL × 100; see Table 1) of the CdR caprine (22.8) is larger than in almost all fossil and extant caprines studied, except Myotragus. Among extant caprines, only Budorcas taxicolor has a similar, even higher, robustness index (mean = 23.7; n = 4; range = 22.2–24.6), but its morphology is completely different. While the tranverse diameters of the distal and proximal epiphyses in the CdR fossil are similar or slightly larger than the transverse diameter at midshaft, Budorcas displays a broad difference in transverse diameters, those of distal and proximal epiphyses being clearly wider than at midshaft (P. Bover, unpub. Ph.D. thesis, Univ. Illes Balears, 2004).

Among the fossils, only the Balearic caprines show a similar morphology and similar or larger robustness index. Two partially reconstructed metatarsals of M. pepgonellae have a robustness index of 32.6 and 36.2 (MNIB 59204 and 59205, respectively) and display almost parallel margins in dorsal view (Fig. 4). Identical morphology is shown by a metacarpal of the caprine from Ses Fontanelles, which has a robustness index of 27. Caprines show similar proportions in hind and forelimb metapodials. Thus, it is probable that the metatarsals of the caprine from Ses Fontanelles had a similar robustness index to the CdR caprine.

A similar morphology and robustness index to the CdR caprine is found in a metacarpal of a Bovidae indet. from the Baccinello-Cinigiano Basin (Tusco-Sardinian palaeoprovince, Late Miocene, Italy) that appears figured in Abbazzi et al. (Reference Abbazzi, Delfino, Gallai, Trebini and Rook2008, text-fig. 18). According to measurements taken from this figure, its robustness index falls around 22.7 and it also displays parallel margins in dorsal view. Unfortunately, no taxonomical assignment can be achieved yet regarding this bone, but the authors suggest that the metacarpal (and another one, not figured) could belong to Etruria viallii (?Antilopinae) or Turritragus casteanensis (Subfamily indet.).

4.c.6. Proximal phalanx

Material. IMEDEA 90132, 90133 and 90274 (Fig. 3g–i).

Since none of the preserved CdR caprine phalanges can be assigned to the hind or the forelimb, all phalanges of recent and fossil caprines (when available) have been used for comparative purposes.

This is a robust phalanx. No remarkable differences between the transverse diameter of the epiphysis and the diaphysis can be observed. The medial and lateral walls of the diaphysis are parallel in dorsal view.

The proximal articular surface is broad, and the incision for the metapodial verticillus is shallow (as in Myotragus). In lateral view, it is concave with a rounded articular area (suggesting that joint movement was slightly restricted, but not as in Myotragus).

The dorsal surface of the diaphysis is convex, and the insertion area for the interdigital ligament is prominent and located slightly distal to the midshaft. There is no groove for the interosseum tendon.

The plantar/palmar surface is slightly concave in transverse section. The distal articular facets do not extend onto the plantar/palmar surface and their margins are flat-V shaped in this view. Sesamoidean facets are small and the insertion areas for the sesamoidean ligaments are also small and are oriented proximo-plantarly/palmarly.

No proximal phalanges of M. pepgonellae have been recovered as yet. Thus, for comparative analysis, the proximal phalanges of the more recent species M. antiquus have been used (Fig. 5). No proximal phalanges of the caprine from Ses Fontanelles or of Aragoral were available for study.

Figure 5.

Comparison between phalanges of Myotragus palomboi n. sp. (a–c) and Myotragus antiquus (d–f). (a) Proximal phalanx IMEDEA 90132: 1 – axial view; 2 – dorsal view; (b) medial phalanx IMEDEA 90134: 1 – abaxial view; 2 – palmar/plantar view; 3 – axial view; (c) distal phalanx IMEDEA 90136: 1 – axial view; 2 – solar view; (d) proximal phalanx: 1 – MNIB 57902: medial view; 2 – MNIB 57901: reversed, dorsal view; (e) medial phalanx: 1 – MNIB 57935, reversed, abaxial view; 2 – MNIB 57922, reversed, palmar/plantar view; 3 – MNIB 57928, axial view; (f) distal phalanx: 1 – MNIB 57950, axial view; 2 – MNIB 57951, reversed, solar view. Scale bar 2 cm.

When compared to other caprines, the most relevant feature of the Caló den Rafelino phalanges is their great robustness (Mean Robustness Index, RI (TD/GL × 100; measurements following von den Driesch, Reference von den Driesch1976) = 39.8; n = 2 (one of the three phalanges, IMEDEA 90274, is fragmented); range = 38.2–41.4). Among recent caprines, only Budorcas and Ovibos display larger or similar RI (48.7 (n = 1) and mean = 39.4 (n = 5; range = 36.7–44.9), respectively), while among the fossils, only Myotragus antiquus shows larger values (even larger than 50; mean = 48; n = 5; range = 43.9–51.8).

In addition to their robustness, the phalanges of the CdR caprine and Myotragus share other morphological features (Fig. 5). The articular facets for the sesamoids are small. The protuberances where the ligaments between sesamoids and the proximal part of the phalanges are attached (sesamoidean ligaments, following Köhler, Reference Köhler1993) are also small and oriented in the proximo-plantar/palmar direction. In other caprines studied, these areas are rough surfaces oriented plantarly/palmarly and extend distalwards to approximately one-third of the diaphysis. Even in species with very robust proximal phalanges (Budorcas) this area is extended.

A characteristic feature of Myotragus (Leinders, Reference Leinders1979; Spoor, Reference Spoor1988) is the reduction of the articular movement between the proximal and medial phalanges, manifested in the flat morphology of the distal articular facet of the proximal phalanx (in lateral view). The caprine from Caló den Rafelino is similar to Norbertia, Gallogoral, Budorcas or Nesogoral in that respect, showing a rounded facet, but without an expansion of these facets on the dorsal and plantar/palmar surface of the diaphysis, as displayed by other caprine taxa (e.g. Capra and Ovis). In addition, the proximal articular surface is slightly reduced in the CdR caprine, Myotragus, Norbertia and Nesogoral (where it has a less concave morphology in lateral view), while in the other caprines it has a rounded outline.

4.c.7. Medial phalanx

Material. IMEDEA 90134 and 90135 (Fig. 3j, k).

Like the proximal phalanx, the second phalanx is short and robust. The post-articular plateau and the dorsal extensor process are very weak or almost absent. The articular fovea is concave.

No articular notches (in the sense of Köhler & Moyà-Solà, Reference Köhler and Moyà-Solà2001) can be observed on the distal and proximal articular surfaces. The distal articular lateral facet has a rounded outline, while the medial facet has a subtriangular outline in medial view (it is divided by a crest).

The distal articular facets are proximally expanded on the dorsal part and they tend to converge near the midshaft. On the plantar/palmar surface these facets extend to one-third of the distal extreme. While in one specimen (IMEDEA 90134) this surface is flat, in the other specimen (IMEDEA 90135) a sagittal groove along the whole diaphysis can be clearly observed. The lateral and medial extremes of the distal articular facets are approximately at the same level.

No medial phalanges of M. pepgonellae have been recovered as yet. Thus, for comparative analysis, medial phalanges of the derived species M. antiquus have been used (Fig. 5). No data for Budorcas, Aragoral or the caprine from Ses Fontanelles were available for study.

As in the proximal phalanx, the medial phalanx of the CdR caprine is very robust (mean RI = 55; n = 2; range = 54–56.1; calculated as in proximal phalange; Table 1). Among the available living caprines, only Ovibos has a similar RI (mean = 55.2; n = 5; range = 48.9–61.5). Again, Myotragus is the taxon with the greatest RI, with values close to 60 (mean = 58.6; n = 5; range = 50.3–70.3). The CdR caprine and Myotragus also share characteristic features not observed in the other caprines studied, such as the very weak morphology of the postarticular plateau and the dorsal extensor process, and an important reduction in the articular movements (as derived from the flat proximal and distal articular facets in lateral view) (Fig. 5). This morphology of the articular facet is also displayed by Ovibos. In both fossil taxa, the lateral and medial extremes of the distal articular facets are approximately at the same level, as in Nesogoral, Capricornis and Gallogoral.

4.c.8. Distal phalanx

Material. IMEDEA 90136 and 90137 (Fig. 3l, m).

Like other phalanges, the distal phalanx is short and robust. Both phalanges available are broken in the processus extensorius zone. The dorsal ridge (on the dorsal border) is rounded. On the wedge/interdigital margin of the plantar or palmar surface of the phalanx there is a prominent protuberance extended axially (clearly visible in IMEDEA 90137, but IMEDEA 90136 is abraded and this protuberance cannot be observed). The solar (distal) border is flat or slightly concave. The border of the plantar wedge (where the flexor digitorum profundus inserts) is poorly developed. In the coronary (proximal) border, the articular surface is broad (in accordance with the size of the articular surface in the medial phalanx) and the crest that separates the two facets is axially curved. The articular surface is open (that is, not very rounded). The apex is blunt. As IMEDEA 90136 is abraded, the protuberance for the extensor insertion is not visible and the dorsal border is also eroded. In this specimen a central notch can be observed on the articular facets, while in IMEDEA 90137 it is absent.

In lateral view the dorsal border is strongly convex, almost rounded. The axial surface is concave and has a small crest on the anterior part, running in the dorso-plantar direction, and the axial foramen is at the same level as the dorsal extreme of the crest on the articular facet.

No distal phalanges of M. pepgonellae have been recovered as yet. Thus, for comparative analysis, distal phalanges of the derived species M. antiquus have been used (Fig. 5), in addition to distal phalanges of Norbertia (fragmented), Nesogoral, Gallogoral, Capra, Ovis, Oreamnos, Nemorhaedus and Ovibos.

The robustness of the distal phalanx in the CdR caprine is 33.6 (n = 1; phalanx IMEDEA 90137 is fragmented); measured as the ratio middle breadth of the sole/diagonal length of the sole (MBS/DLS × 100; Table 1), following von den Driesch, Reference von den Driesch1976). Only Ovibos (mean = 33.2; n = 5; range = 30.6–35.2) and Myotragus (mean = 39.7; n = 5; range = 35.2–42) display similar or greater robustness. The CdR caprine shares with the other studied caprines an almost rounded shape of the distal part of the dorsal ridge. In Myotragus, a prominent protuberance extended axially on the wedge of the phalanx can also be observed, although it is not as developed as in the CdR caprine (Fig. 5). In the other caprines this protuberance is a weak and elongated rough area (except in Ovibos, where this protuberance projects axially from the broad articular facet). Broad articular facets and practically flat soles are shared by Ovibos, Myotragus and the CdR caprine.