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A species factory refers to the source that gives rise to an exceptionally large number of species. However, what is it exactly: a place, a time or a combination of places, times and environmental conditions, remains unclear. Here we search for species factories computationally, for which we develop statistical approaches to detect origination, extinction and sorting hotspots in space and time in the fossil record. Using data on European Late Cenozoic mammals, we analyse where, how and how often species factories occur, and how they potentially relate to the dynamics of environmental conditions. We find that in the Early Miocene origination hotspots tend to be located in areas with relatively low estimated net primary productivity. Our pilot study shows that species first occurring in origination hotspots tend to have a longer average longevity and a larger geographical range than other species, thus emphasizing the evolutionary importance of the species factories.

Dietary analyses of herbivorous mammals are important for paleoecological reconstruction. Several methods applicable to fossil teeth have been developed lately. The mesowear method based on wear-induced occlusal shape and relief of ungulate molars has proven to be a robust method for dietary analysis. In its original form it can only be used for selenodont, plagiolophodont, and ectolophodont ungulate molars, but the principle can be extended to other kinds of tooth morphology. We introduce a new method of dietary analysis for proboscideans similar to the mesowear method, based on angle measurements from worn dentin valleys reflecting the relief of enamel ridges. The enamel ridges should be heavily worn when the abrasiveness of diet increases, resulting in lower occlusal relief and larger angles. For testing this, we compared the mesowear angles with stable carbon isotope values from dental enamel from populations of extant and fossil species from localities from Kenya and India. This enables us to compare diet and tooth wear in proboscideans, because the stable carbon isotope ratios in tropical environments provide a reliable standard for assessing the relative amounts of C4 and C3 plants in diet, and most of the C4 plants are grasses, which should be reflected in the mesowear signal.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Bovidae indet. LOCALITY. ��� Quarry 1, Ivand district, northwest of Tabriz, Iran. MATERIAL EXAMINED. ��� Fragmentary right mandible with m2-m3 (HMNH-IV 59; Table 3), right m1 (HM- NH-IV 65; Table 3), broken right m2 (HMNH-IV 68; Table 3), left maxilla with DP2-M1 (HMNH-IV 70; Table 3), fragmentary right maxilla with M1-M2 (HMNH- IV 64; Table 3), fragmentary right maxilla with P4-M1 (HMNH-IV 58; Table 3), fragmentary left maxilla with M1-M2 (HMNH-IV 57; Table 3). DESCRIPTION The mandibles and maxillae are too fragmentary to provide any detailed character. DP2 is heavily worn and is long and narrow. The parastyle is prominent but low. There is no mesostyle, and the metastyle is weak. The paracone is the highest cusp, and the metacone is slightly lower. The protocone is large and located anteriorly to the paracone. The hypocone is larger than the protocone. There is no central fossette visible on the occlusal surface. DP3 is longer than DP2 and is submolarized. The parastyle is strong and well separated from the paracone by a wide groove. The mesostyle, however, is less developed, and the metastyle is the least developed. The anterior rib is very strong, and the posterior rib is weak. The central cavities bear a simple enamel outline. DP4 is highly molarized and, except for its smaller size, closely resembles molars. The central fossettes are open in the medium wear. P4 is subtriangular. The hypocone is relatively well developed, though smaller than protocone, thus rendering the inner wall a less angled shape. The parastyle and metastyle are well developed, while the mesostyle is weak. On M1, the parastyle is robust, the mesostyle is well developed as a vertical ridge, and the metastyle projects much less. The anterior rib is large, and the posterior rib is weak. There is no basal pillar. M2 closely resembles M 1 in tooth morphology. The hypocone triangle is relatively short and is rounder than the protocone triangle. The metastyle is slightly more developed. m1 shows a rectangular occlusal outline. The parastylid, metastylid, and the entostylid are much reduced. The lingual wall is flat with slightly convex metaconid and entoconid. The protoconid and hypoconid on m2 are triangle shaped. The parastylid is slightly more developed than the metastylid and entostylid. There is no goat fold but there is a low basal pillar. m3 resembles m2, except for its strongly labially offset hypoconulid with a central cavity. COMPARISON With no cranial or horn-cores found, discussing the systematic position of these isolated fragmentary materials in detail is difficult. Based on size and morphology, these teeth are tentatively assigned to one form herein, pending the discovery of better specimens. This form, like Urmiatherium polaki Rodler, 1889, features a developed parastyle, very strong mesostyles, a prominent anterior rib, and a concave labial metacone wall on the upper molars. It differs by having a lower tooth crown, no additional cavities near the central lingual edge of the occlusal surface on the upper molars, a convex shape of the lingual wall, and the well developed basal pillars on the lower molars. De Mecquenem (1924, 1925) reported some material of Palaeoryx pallasi (Wagner, 1857) from Maragheh. The size of the present form from Ivand locality is comparable to this Maragheh A 550 500 450 400 350 taxon. They share similar tooth morphology, such as strong parastyles and mesostyles, weak metastyles, developed anterior ribs, no basal pillars on the upper molars, a slightly convex lingual wall of the lower molars, low but distinct basal pillars, triangular labial lobes on the lower molars, and strongly labially offset hypoconulids on m3. The skull figured by de Mecquenem (1924, 1925: pl. 4, fig. 1), however, was believed to be that of a Miotragocerus Stromer, 1928 (= Tragoportax Pilgrim, 1937) by Bohlin (1936) and Gentry (1971, 2000). Gentry (1971) further stated that there was no other convincing evidence of Palaeoryx pallasi from Maragheh. Given these poor materials and pending the discovery of more complete fossils, we avoid naming them., Published as part of Ataabadi, Majid Mirzaie, Mohammadalizadeh, Jafar, Zhang, Zhaoqun, Watabe, Mahito, Kaakinen, Anu & Fortelius, Mikael, 2011, Late Miocene large mammals from Ivand (Northwestern Iran), pp. 709-728 in Geodiversitas 33 (4) on pages 721-723, DOI: 10.5252/g2011n4a7, http://zenodo.org/record/4597129, {"references":["DE MECQUENEM R. 1924. - Contribution a l'etude des fossiles de Maragha. Annales de Paleontologie 13: 135 - 160.","DE MECQUENEM R. 1925. - Contribution a l'etude des fossiles de Maragha. Annales de Paleontologie 14: 1 - 64.","ANTOINE P. O. & SARAC G. 2005. - Rhinocerotidae (Mammalia, Perissodactyla) from the Late Miocene of Akkasdagi, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkasdagi, Late Miocene of Central Anatolia. Geodiversitas 27 (4): 601 - 632.","DENG T. 2002. - Limb bones of Chilotherium wimani (Perissodactyla, Rhinocerotidae) from the Late Miocene of the Linxia Basin in Gansu, China. Vertebrata PalAsiatica 40 (4): 305 - 316.","HUTTUNEN K. & GOHLICH U. B. 2002. - A partial skeleton of Prodeinotherium bavaricum (Proboscidea, Mammalia) from the middle Miocene of Unterzolling (Upper Freshwater Molasse, Germany). Geobios 35 (4): 489 - 514.","BOHLIN B. 1936. - Bemerkungen uber einige pontischen Antilopen-Gattungen. Arkiv for Zoologi 28 A (18): 1 - 22.","GENTRY A. W. 1971. - The earliest goats and other antelopes from the Samos Hipparion fauna. Bulletin of the British Museum (Natural History), Geology 20: 231 - 296.","GENTRY A. W. 2000. - Caprinae and Hippotragini (Bovidae, Mammalia) in the Upper Miocene, in VRBA E. S. & SCHALLER G. B. (eds), Antelopes, Deer and Relatives: Fossil Record, Behavioral Ecology, Systematics and Conservation. Yale University Press, New Haven: 65 - 83."]}

Deinotherium giganteum Kaup, 1829 (Fig. 5 A-R) LOCALITY. ��� Quarry 3, Ivand district, north of Tabriz (Iran). MATERIAL EXAMINED. ��� Postcranials, including proximal part of ulna (HMNH-IV 135; Fig. 5 A-C, Table 2); proximal part of radius (HMNH-IV 115; Fig. 5 P-R, Table 2); proximal part of tibia (HMNH- IV119; Fig. 5 D-F, Table 2); calcaneus (HMNH-IV 118; Fig. 5 G-I, Table 2); unciform (HMNH-IV 117; Fig. 5 J- L, Table 2) and one metapodial (HMNH-IV 114; Fig. 5 M-O, Table 2). DESCRIPTION The proximal part of an ulna is preserved (Fig. 5 A-C). The tuber olecrani is partly damaged, but shows a slight extension in the medial part. The processus coronoideus medialis and lateralis are round and of approximately the same size. The processus anconaeus is slightly higher than the tuber olecrani. The cross section of the shaft is triangular. The proximal part of a single radius is present (Fig. 5 P-R). The shape of the caput radii is sub-triangular. The articular circumference is damaged. The tibia includes only one proximal part (Fig.5 D- F). In the proximal view, only the condylus lateralis and medialis are visible. Calcaneus is an almost complete specimen with minor damage (Fig. 5 G-I). The tuber calcanei is higher than it is wide. Proximally, there are three facets, two of which are partly preserved. The lateral facet is not preserved. The two astragalus facets are separated by sulcus calcanei. The larger astragalus facet is rounded and the smaller one is on the sustentaculum tali and is partly broken. On the dorsal of the sustentaculum tali, parts of a navicular facet are preserved. The unciform is nearly complete and has a triangular outline (Fig. 5 J-L). The proximal facet is triangular and concave. The distal side has one triangular concave facet and another elongated facet. The third facet is destroyed. COMPARISON The morphologies of postcranial skeletons in deinotheriids bear notable similarities (Huttunen 2002). From the postcranial elements described here, the proximal condyle of the ulna from Ivand is very well preserved and allows us to compare it to other deinotheriid specimens from other localities. Figure 6B shows the bivariate plots of the proximal width of the ulna against the width at process olecrani in IV135 and specimens from Maragheh (MN12, NW Iran), Saloniki and Pikermi (MN12, Greece), and Unterzolling (Germany). The plot illustrates that the Ivand material, though slightly smaller in proportion, is close to both Deinotherium giganteum from Pikermi and an undetermined specimen from Maragheh. The ulna of the Ivand specimen is obviously larger than the ulna of Choerolophodon pentelici (Gaudry & Lartet, 1856) from Saloniki, Greece and of Prodeinotherium bavaricum (Meyer, 1831) from Unterzolling (Huttunen & G��hlich 2002). Therefore, based on this comparison, and since Deinotherium giganteum is the largest of the Late Miocene mammals inNW Iran, previously recorded from Maragheh, we assign the very large postcrani - als from the Ivand locality to this species. Order ARTIODACTYLA Owen, 1848, Published as part of Ataabadi, Majid Mirzaie, Mohammadalizadeh, Jafar, Zhang, Zhaoqun, Watabe, Mahito, Kaakinen, Anu & Fortelius, Mikael, 2011, Late Miocene large mammals from Ivand (Northwestern Iran), pp. 709-728 in Geodiversitas 33 (4) on page 718, DOI: 10.5252/g2011n4a7, http://zenodo.org/record/4597129, {"references":["HUTTUNEN K. & GOHLICH U. B. 2002. - A partial skeleton of Prodeinotherium bavaricum (Proboscidea, Mammalia) from the middle Miocene of Unterzolling (Upper Freshwater Molasse, Germany). Geobios 35 (4): 489 - 514."]}

Gazella sp. (Fig. 7 A-D, G, H) LOCALITY. ��� Quarry 3, Ivand district, north of Tabriz, Iran. MATERIAL EXAMINED. ��� Two left and one right broken frontlets with horn-cores (HMNH-IV 138-140; Fig. 7 A-D, G, H). DESCRIPTION Frontlet HMNH-IV138 (Fig. 7C, D) preserves an almost complete horn-core (80 mm above the pedicle), unfortunately broken in the middle and preserving little of the frontal or orbit. IV139 (Fig.7G, H) features the preserved supra-orbital fossa, but lacks the tip of the horn-core. IV140 (Fig. 7A, B) preserves a partial orbit and a small part of the parietal, but lacks the tip of the horn-core and has a broken inner side. Frontlets IV139 and IV140 bear similar sizes and horn-core morphologies. The DAP and DT at the base of IV139 are 25 mm and 21.5 mm, respectively. The horn-cores are located above the orbits, moderately curved posteriorly in lateral view, and straight in anterior view. The base of the horn-core is relatively robust and laterally compressed, and the cross section diminishes greatly upwards. The maximum width lies at the anterior side of the horn-core while the posterior side is narrow. Longitudinal grooves are also present. The supra-orbital fossa is large and oval-shaped, and is located slightly towards the middle of the horn-core mid-line. The dorsal orbital rim is wide and features a postcornual fossa. Frontlet IV138 differs from the two frontlets described above in that it is less curved in lateral view and is less robust at the base; its cross section diminishes gradually from the base upwards. COMPARISON According to the morphology of horn-cores,the three frontlets found at Ivand locality can be separated into two groups. Frontlets IV139 and IV140 show homogenous characteristics and should be classified into one form; IV138 may belong to another form. Horn-cores IV139 and IV140 show great similarities to Gazella deperdita (Gervais, 1847) in that they fea- ture a robust base, rapidly diminishing cross section, a curved outline in the side view, and a maximum width located at the anterior part. IV138 however, is much less curved in the side view, with the cross section gradually changing from the base upwards. These characteristics fit Gazella gaudryi Schlosser, 1904 (Arambourg & Piveteau 1929)., Published as part of Ataabadi, Majid Mirzaie, Mohammadalizadeh, Jafar, Zhang, Zhaoqun, Watabe, Mahito, Kaakinen, Anu & Fortelius, Mikael, 2011, Late Miocene large mammals from Ivand (Northwestern Iran), pp. 709-728 in Geodiversitas 33 (4) on page 720, DOI: 10.5252/g2011n4a7, http://zenodo.org/record/4597129, {"references":["ARAMBOURG C. & PIVETEAU J. 1929. - Les vertebres du Pontien de Salonique. Annales de Paleontologie 18: 59 - 138."]}

Rhinocerotinae indet. (Fig. 5 S-U) LOCALITY. ��� Quarry 3, Ivand district, north of Tabriz, Iran. MATERIAL EXAMINED. ��� Femur (HMNH-IV 137; Fig.5 S- U; Table 2). DESCRIPTION An almost complete left femur is preserved (Fig. 5 S- U). In the proximal part, the femoral head (caput femoris) and trochanter major are missing. The A Hipparion sp. large Ivand H. giganteum Gromova, 1952 Grebeniki H. primigenium Meyer, 1833 Inzersdorf H. prostylum Gervais, 1849 Saloniki H. sp. Large Samos H. brachypus Hensel, 1862 Pikermi H. mediterraneum Roth & Wagner, 1855 Pikermi H. moldavicum Gromova, 1952 Tarakilia H. proboscideum Studer, 1911 Samos Hipparion sp. large Ivand H. moldavicum Gromova, 1952 H. gettyi Bernor, 1985 H. campbelli Bernor, 1985 H. prostylum Gervais, 1849 third trochanter is broken, but the basal part is preserved. The diaphysis becomes stockier distally from the third trochanter. The distal end is well preserved, and only the posterior and anterior sides are abraded. The medial and lateral trochleas are at the same level. The trochlear trough is damaged. The two condyles somehow diverge downward and the medial condyle is higher in elevation. The intercondyloid fossa is filled with sediment. COMPARISON Figure 6A shows the scatter plot of the femoral length against the distal width of well-preserved rhinocerotid femurs from several localities in Greece, Turkey, Iran, and China. The femur from Ivand locality, as illustrated, is clearly distinct from those of Chilotherium wimani Ringstr��m, 1924 and C. persiae Pohlig, 1887 which are smaller, slenderer, and less massive (Deng 2002). The femur IV137 clusters among the material assigned to Ceratotherium neumayri (Osborn, 1900) and Stephanorhinus pikermiensis (Toula, 1906) from eastern Mediterranean localities such as Pikermi, Maragheh, and Akkaşdaği (Fortelius et al. 2003; Antoine & Sara�� 2005). In general, the femur of these species is about 50 cm long and 15 cm wide distally. The total length of the Ivand femur, due to its missing proximal end, is unknown but it is estimated to be about 45 cm. Therefore, the rhinocerotid femur from Ivand locality can be assigned to Ceratotherium neumayri / Stephanorhinus pikermiensis primarily by size. Order PROBOSCIDEA Illiger, 1811, Published as part of Ataabadi, Majid Mirzaie, Mohammadalizadeh, Jafar, Zhang, Zhaoqun, Watabe, Mahito, Kaakinen, Anu & Fortelius, Mikael, 2011, Late Miocene large mammals from Ivand (Northwestern Iran), pp. 709-728 in Geodiversitas 33 (4) on pages 716-718, DOI: 10.5252/g2011n4a7, http://zenodo.org/record/4597129, {"references":["GROMOVA V. 1952. - [Gippariony (rod Hipparion) po materialam Taraklii, Pavlodara i drugim]. Trudy Paleontologicheskogo Instituta 36: 1 - 478 (in Russian).","BERNOR R. L. 1985. - Systematic and evolutionary relationships of the hipparionine horses from Maragheh, Iran (Late Miocene, Turolian age). Palaeovertebrata 15 (4): 173 - 269.","DENG T. 2002. - Limb bones of Chilotherium wimani (Perissodactyla, Rhinocerotidae) from the Late Miocene of the Linxia Basin in Gansu, China. Vertebrata PalAsiatica 40 (4): 305 - 316.","FORTELIUS M., HEISSIG K., SARAC G. & SEN S. 2003. - Rhinocerotidae (Perissodactyla), in FORTELIUS M., KAPPELMAN J., SEN S. & BERNOR R. L. (eds), Geology and Paleontology of the Miocene Sinap Formation, Turkey. Columbia University Press, New York: 282 - 307.","ANTOINE P. O. & SARAC G. 2005. - Rhinocerotidae (Mammalia, Perissodactyla) from the Late Miocene of Akkasdagi, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkasdagi, Late Miocene of Central Anatolia. Geodiversitas 27 (4): 601 - 632."]}

Oioceros atropatenes Rodler & Weithofer, 1890 (Fig. 7E, F) LOCALITY. — Quarry 1, Ivand district, north of Tabriz, Iran. MATERIAL EXAMINED. — Left horn-core (HMNH-IV 200; Fig. 7E, F), right mandible with p4-m3 (HMNH-IV 67; Table 3), right mandible with broken m2 and complete m3 (HMNH-IV 69; Table 3). DESCRIPTION The horn-core specimen is partly broken at the base and the tip. The antero-posterior diameter cannot be measured precisely due to the basal missing part on both the anterior and posterior sides. However, we estimate that it exceeds 17 mm. The medio- A M1 M2 M3 M4 M6 M7 M14 M15 M23 M25 M30 M31 M32 M33 lateral diameter is 17.4 mm. The DAP and DT at 5 cm above the base are 12.7 mm and 10.9 mm, respectively. The total preserved length is 75 mm from the pedicle. By the preserved part of the orbit and remnant of the frontal, the horn-core is located above the posterior part of the orbit, tilted slightly backwards with a weak curvature. The horn-core is slender with a roughly oval cross section. The antero-posterior and transverse diameters diminish slowly from the base upwards. There are two weak keels, one starting from the antero-lateral side, and the other from the postero-lateral side. These two keels enclose a slightly convex outer surface and a more rounded inner surface. The keels spiral clock- wise roughly one gyre from the base to the tip by estimation. There is also a postcornual fossa. p4 is long and narrow. The paraconid is not separated from the parastylid. The anterior valley is wide. The metaconid is situated posterior to the protoconid. The entoconid is close to the entostylid. A wide and shallow valley separates the protoconid from the hypoconid.m1 is well worn. The parastylid is developed, and well separated from the metaconid. There is no goat fold, thus rendering the anterior border much narrower. The lingual wall is flat with weak metastylids. The entostylid is larger. There is a large basal pillar between the protoconid and the metaconid. m2 is very similar in morphology to m1, except for its larger, more convex lingual wall, and lower basal pillar. The parastylid on m3 is more pronounced and well separated from the metaconid. The basal pillar is small and low. The hypoconulid is large and postero-labially offset. COMPARISON Based on the small size, insertion above the posterior part of orbit, the clockwise torsion on the left horn-core from the base, and two keels, the horn-core IV200 from Ivand locality can readily be assigned to the genus Oioceros (type species Antelope rothii Wagner, 1857 from Pikermi, Greece) (Gaillard 1902). Since then, numerous species were included or assigned to this genus. De Mecquenem (1924) recognized three species from Maragheh: O. rothii (Wagner, 1857), O. atropatenes and O. boulei Mecquenem, 1924. Heintz (1963) synonymized O. boulei with O. atropatenes based on his detailed description and comparison. Roussiakis (2003) recently described in detail an almost complete skull with mandibles of Oioceros rothii from Pikermi and reviewed the generic status. Besides the type species, he listed only Oioceros atropatenes in the genus. The present specimen is smaller and less laterally compressed than Oioceros rothii from Pikermi. The main keel is much weaker and the upper part of the horn-core diverges less. The size falls into the variation of Oioceros atropatenes from Maragheh, Iran (Heintz 1963). The morphology of the horn-core is also consistent with those from Maragheh. Hence, the horn-core specimen can be assigned to Oioceros atropatenes.

Oioceros atropatenes Rodler & Weithofer, 1890 (Fig. 7E, F) LOCALITY. ��� Quarry 1, Ivand district, north of Tabriz, Iran. MATERIAL EXAMINED. ��� Left horn-core (HMNH-IV 200; Fig. 7E, F), right mandible with p4-m3 (HMNH-IV 67; Table 3), right mandible with broken m2 and complete m3 (HMNH-IV 69; Table 3). DESCRIPTION The horn-core specimen is partly broken at the base and the tip. The antero-posterior diameter cannot be measured precisely due to the basal missing part on both the anterior and posterior sides. However, we estimate that it exceeds 17 mm. The medio- A M1 M2 M3 M4 M6 M7 M14 M15 M23 M25 M30 M31 M32 M33 lateral diameter is 17.4 mm. The DAP and DT at 5 cm above the base are 12.7 mm and 10.9 mm, respectively. The total preserved length is 75 mm from the pedicle. By the preserved part of the orbit and remnant of the frontal, the horn-core is located above the posterior part of the orbit, tilted slightly backwards with a weak curvature. The horn-core is slender with a roughly oval cross section. The antero-posterior and transverse diameters diminish slowly from the base upwards. There are two weak keels, one starting from the antero-lateral side, and the other from the postero-lateral side. These two keels enclose a slightly convex outer surface and a more rounded inner surface. The keels spiral clock- wise roughly one gyre from the base to the tip by estimation. There is also a postcornual fossa. p4 is long and narrow. The paraconid is not separated from the parastylid. The anterior valley is wide. The metaconid is situated posterior to the protoconid. The entoconid is close to the entostylid. A wide and shallow valley separates the protoconid from the hypoconid.m1 is well worn. The parastylid is developed, and well separated from the metaconid. There is no goat fold, thus rendering the anterior border much narrower. The lingual wall is flat with weak metastylids. The entostylid is larger. There is a large basal pillar between the protoconid and the metaconid. m2 is very similar in morphology to m1, except for its larger, more convex lingual wall, and lower basal pillar. The parastylid on m3 is more pronounced and well separated from the metaconid. The basal pillar is small and low. The hypoconulid is large and postero-labially offset. COMPARISON Based on the small size, insertion above the posterior part of orbit, the clockwise torsion on the left horn-core from the base, and two keels, the horn-core IV200 from Ivand locality can readily be assigned to the genus Oioceros (type species Antelope rothii Wagner, 1857 from Pikermi, Greece) (Gaillard 1902). Since then, numerous species were included or assigned to this genus. De Mecquenem (1924) recognized three species from Maragheh: O. rothii (Wagner, 1857), O. atropatenes and O. boulei Mecquenem, 1924. Heintz (1963) synonymized O. boulei with O. atropatenes based on his detailed description and comparison. Roussiakis (2003) recently described in detail an almost complete skull with mandibles of Oioceros rothii from Pikermi and reviewed the generic status. Besides the type species, he listed only Oioceros atropatenes in the genus. The present specimen is smaller and less laterally compressed than Oioceros rothii from Pikermi. The main keel is much weaker and the upper part of the horn-core diverges less. The size falls into the variation of Oioceros atropatenes from Maragheh, Iran (Heintz 1963). The morphology of the horn-core is also consistent with those from Maragheh. Hence, the horn-core specimen can be assigned to Oioceros atropatenes., Published as part of Ataabadi, Majid Mirzaie, Mohammadalizadeh, Jafar, Zhang, Zhaoqun, Watabe, Mahito, Kaakinen, Anu & Fortelius, Mikael, 2011, Late Miocene large mammals from Ivand (Northwestern Iran), pp. 709-728 in Geodiversitas 33 (4) on pages 718-720, DOI: 10.5252/g2011n4a7, http://zenodo.org/record/4597129, {"references":["GROMOVA V. 1952. - [Gippariony (rod Hipparion) po materialam Taraklii, Pavlodara i drugim]. Trudy Paleontologicheskogo Instituta 36: 1 - 478 (in Russian).","BERNOR R. L. 1985. - Systematic and evolutionary relationships of the hipparionine horses from Maragheh, Iran (Late Miocene, Turolian age). Palaeovertebrata 15 (4): 173 - 269.","WATABE M. & NAKAYA H. 1991 b. - Cranial skeletons of Hipparion (Perissodactyla, Mammalia) from Maragheh (Turolian, Late Miocene), NW Iran. Memoirs of the Faculty of Sciences, Kyoto University 56 (1 - 2): 55 - 125.","BERNOR R. L., TOBIEN H., HAYEK L. A. & MITTMANN H. W. 1997. - The Howenegg hipparionine horses: systematic, stratigraphy, taphonomy and paleoenvironmental context. Andrias 10: 1 - 230.","GAILLARD C. 1902. - Le belier de Mendes ou le mouton domestique de l'ancienne Egypte. Bulletin de la Societe d'Anthropologie de Lyon 20: 69 - 103.","DE MECQUENEM R. 1924. - Contribution a l'etude des fossiles de Maragha. Annales de Paleontologie 13: 135 - 160.","HEINTZ E. 1963. - Complement d'etude sur Oioceros atropatenes (Rod. et Weith.), Antilope du Pontien de Maragha (Iran). Bulletin de la Societe geologique de France 5: 109 - 116.","ROUSSIAKIS S. J. 2003. - Oioceros rothii (Wagner, 1857) from the Late Miocene of Pikermi (Greece): cranial and dental morphology, comparison with related forms. Geodiversitas 25 (4): 717 - 735."]}

Hipparion sp. large (Fig. 2) LOCALITY. ��� Quarry 2, Ivand district, north of Tabriz, Iran. MATERIAL EXAMINED. ��� Skull (HMNH-IV 1) and mandible (HMNH-IV 2) (Fig. 2; Table 1). DESCRIPTION The skull is quite long and belongs to a large adult male (large canines present). The muzzle is elongated and broad, even though the application of extra material in fixing the broken snout has made the muzzle artificially longer (Fig. 2 A-C). The narial opening and nasal area have been damaged during excavation and subsequently restored, so details of this area are missing. The POF is very well preserved on the right side and partially filled with sediment on the left side (Fig.2A).The POF is far from the orbit,sub-triangular in shape, relatively large and deep, and antero-posteriorly oriented. There is no posterior pocketing. The anterior rim of the POF is well expressed and appears above the medial part of P3. The posterior border of the POF is located above the anterior part of M2. The ventral rim of the POF is not straight, features undulation, and is above the line of the lower border of the orbit.The infra-orbital foramen is at the anteroventral border of the fossa, placed slightly inferiorly. The orbit is round and its anterior border is situated close to the posterior border of M3. The facial crest is very strong and is far from the alveoli and the ventral border of the POF. The anterior border of the facial crest is situated above the medial part of P4.The palate is elongated and wide. The choanae, although filled with sediment, are well preserved. They are wide and their anterior border is situated at the level of contact between M2 and M3. The upper tooth row is long and complete at both sides, including P2-4, M1-3. All the teeth are moderately worn and the enamel morphology is visible (Fig. 2D). The protocone is elliptical and flattened lingually, especially in the molars. The enamel plication is rich (mean plication is 20) with deep plis. The hypocone is elliptical with relatively deep distal hypoconal grooves. The plicabaline is strong, short and single or long and double/multiple. The mandible is elongated with a narrow snout and cup. The symphysis is short and narrow.The condyle and coronoid processes and the ascending ramus, though not preserved,have been restored (Fig.2 E-G). The tooth row is long and the teeth are large and wide (Fig. 2H). The parastylid is relatively well developed and closed. The metaconid is elliptical to round and the metastylid is rounded. The entoconid is elliptical to round. The ectoflexid is V- or U-shaped and moderately deep and narrow, reaching the middle of the tooth; only in m1 it reaches the linguaflexid. A plicabalinid is present in some of the teeth. COMPARASION The hipparion skull from Ivand (HMNH-IV1) has been compared to several species of hipparionine horses, from localities in the eastern Mediterranean and northern Black Sea regions. The bivariate plots of the POB width (distance between the anterior rim of the orbit and the posterior rim of the POF) against the P2-orbit distance, which serve as a useful measure of skull (face) length (Forsten 1983), appear in Figure 3. Here, Ivand skull clusters with H. giganteum Gromova, 1952 from Grebeniki, H. brachypus Hensel, 1862 from Pikermi, and Hipparion sp. large from Samos (Fig. 3A). The results of a similar plot show the distinction, based D, H on this criterion, of the Ivand skull from those of other known hipparion species in Maragheh (Fig. 3B), including ��� H ���. gettyi Bernor, 1985; ��� H. ��� aff. moldavicum (H. moldavicum Gromova, 1952 sensu Watabe & Nakaya 1991b); Hipparion prostylum Gervais, 1849; and H. campbelli Bernor, 1985 (H. urmiense Gabunia, 1959 sensu Watabe & Nakaya 1991b). The large-sized skull from Ivand (HMNH-IV1), as the logarithmic ratio diagram shows (Fig. 4A), is comparable in its basic dimensions and morphology to H. brachypus from Pikermi (Koufos 1987b), Hadjidomovo (Hristova et al. 2003), and Akkaşdaği (Koufos & Vlachou 2005). However, the Ivand skull has a longer muzzle (M1), which is indeed an artifact of incorrect restoration, and a wider snout (M15). Figure 4B shows that the large-sized skull from the Ivand locality is not comparable in size and morphology to any of the hipparionine horses from Maragheh (Bernor 1985). Hipparion brachypus is characterized by large size, an elongated skull with a relatively wide muzzle, and a deep narial opening. The preorbital fossa is oval, antero-posteriorly oriented, well-marked, deeply posteriorly pocketed, and situated far from the orbit. The upper cheek teeth also show rich enamel plication with deep plis (Koufos 1987a; Koufos & Vlachou 2005; Vlachou & Koufos 2009). Ivand skull (Fig. 4A) also partially resembles H. giganteum from Grebeniki, Moldavia (Gromova 1952). Hipparion giganteum is also a large hipparion with a single, elliptical, and deep POF with strong posterior pocketing, located far from the orbit and facial crest. Nevertheless, the large-sized skull from Ivand has a sub-triangular POF which lacks posterior pocketing. Ivand skull is also comparable to a large-sized skull from Samos (AMNH 22838), described as H. cf. proboscideum (Sondaar 1971). This specimen from Samos features more facial height than does the large skull from Ivand. Unlike the type skull of H. proboscideum Studer, 1911, AMNH 22838 bears a single and deep POF (Sondaar 1971), which makes it morphologically similar to the H. brachypus skull from Akkaşdaği (Koufos & Vlachou 2005). Based on the bivariate plots and log ratio diagrams (Figs 3, 4), the hipparion skull from the Ivand locality likely belongs to a large-sized Hipparion species of H. giganteum - H. brachypus lineage (��� Hippotherium ��� brachypus -��� Hippotherium ��� giganteum lineage, sensu Bernor et al. 1996b), previously unknown from the cranial material in NW Iran (i.e. the Maragheh area). We refrain from classifying this specimen at the species level because of some differences in its facial morphology, such as the shape of the POF and lack of posterior pocketing, and its inaccurately restored muzzle. The occurrence of large hipparions (similar to H. brachypus) in Maragheh was previously evidenced by the presence of medium to large, robust third metapodials (Watabe & Nakaya 1991a; Tobien in Bernor et al. 1996a, b). Some of these robust metapodials were assigned to H. prostylum, even though they showed greater similarity to the metapodials of H. brachypus from Pikermi (Watabe & Nakaya 1991a: fig. 15). Family RHINOCEROTIDAE Owen, 1845, Published as part of Ataabadi, Majid Mirzaie, Mohammadalizadeh, Jafar, Zhang, Zhaoqun, Watabe, Mahito, Kaakinen, Anu & Fortelius, Mikael, 2011, Late Miocene large mammals from Ivand (Northwestern Iran), pp. 709-728 in Geodiversitas 33 (4) on pages 713-716, DOI: 10.5252/g2011n4a7, http://zenodo.org/record/4597129, {"references":["BERNOR R. L., SCOTT R. S., FORTELIUS M., KAPPEL- MAN J. & SEN S. 2003. - Equidae (Perissodactyla), in FORTELIUS M., KAPPELMAN J., SEN S. & BERNOR R. L. (eds), The Geology and Paleontology of the Miocene Sinap Formation, Turkey. Columbia University Press, New York: 220 - 281.","FORSTEN A. 1983. - The preorbital fossa as a taxonomic character in some Old World Hipparion. Journal of Paleontology 57 (4): 686 - 704.","GROMOVA V. 1952. - [Gippariony (rod Hipparion) po materialam Taraklii, Pavlodara i drugim]. Trudy Paleontologicheskogo Instituta 36: 1 - 478 (in Russian).","BERNOR R. L. 1985. - Systematic and evolutionary relationships of the hipparionine horses from Maragheh, Iran (Late Miocene, Turolian age). Palaeovertebrata 15 (4): 173 - 269.","WATABE M. & NAKAYA H. 1991 b. - Cranial skeletons of Hipparion (Perissodactyla, Mammalia) from Maragheh (Turolian, Late Miocene), NW Iran. Memoirs of the Faculty of Sciences, Kyoto University 56 (1 - 2): 55 - 125.","KOUFOS G. D. 1987 b. - Study of the Pikermi hipparions. Part II: Comparisons and odontograms. Bulletin du Museum national d'Histoire naturelle, Paris, 4 e ser., 9, section C, 3: 327 - 363.`","HRISTOVA L., KOVACHEV D. & SPASSOV N. 2003. - Hipparion brachypus Hensel, 1862 from the Late Miocene of Hadjidimovo, Southwestern Bulgaria. Comptes Rendus de l'Academie bulgare des Sciences 56 (2): 77 - 84.","KOUFOS G. D. & VLACHOU T. D. 2005. - Equidae (Mammalia, Perissodactyla) from the Late Miocene of Akkasdagi, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkasdagi, Late Miocene of Central Anatolia. Geodiversitas 27 (4): 633 - 705.","KOUFOS G. D. 1987 a. - Study of the Pikermi hipparions. Part I: Generalities and taxonomy. Bulletin du Museum national d'Histoire naturelle, Paris, 4 e ser., 9, section C, 2: 197 - 252.","SONDAAR P. Y. 1971. - The Samos Hipparion. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen B 74: 417 - 441.","BERNOR R. L., KOUFOS G. D., WOODBURNE M. & FORTELIUS M. 1996 b - The evolutionary history and biochronology of European and southeastern Asian Late Miocene and Pliocene hipparionine horses, in BERNOR R. L., FAHLBUSCH V. & MITTMANN H. W. (eds), The Evolution of Western Eurasian Neogene Mammal Faunas. Columbia University Press, New York: 307 - 338.","WATABE M. & NAKAYA H. 1991 a. - Phylogenetic significance of the postcranial skeletons of the hipparions from Maragheh (Late Miocene), NW Iran. Memoirs of the Faculty of Sciences, Kyoto University 56 (1 - 2): 11 - 53.","BERNOR R. L., SOLOUNIAS N., SWISHER III C. C. & VAN COUVERING J. 1996 a - The correlation of three classical \" Pikermian \" mammal faunas (Maragheh, Samos, and Pikermi) with the European MN unit system, in BERNOR R. L., FAHLBUSCH V. & MITT- MANN H. W. (eds), The Evolution of Western Eurasian Neogene Mammal Faunas. Columbia University Press, New York: 137 - 154."]}

Liu, Liping, Kostopoulos, Dimitris S., Fortelius, Mikael (2005): Suidae (Mammalia, Artiodactyla) from the late Miocene of Akkaşdağı, Turkey. Geodiversitas 27 (4): 715-733, DOI: http://doi.org/10.5281/zenodo.4665351, {"references":["GAUDRY A. 1862-1867. - Animaux fossiles et geologie de l'Attique. F. Savy, Paris, 2 vols, 472 p.","GERVAIS P. 1848-1852. - Zoologie et paleontologie francaises. A. Bertrand, Paris, 2 vols, 3 vols.","HELLMUND M. 1995. - The vertebrate locality Maramena (Macedonia, Greece) at the Turolian- Ruscinian boundary (Neogene) 13. Suidae (Artiodactyla, Mammalia). Munchener Geowissenschaftliche Abhandlungen, series A (28): 143-156.","HUNERMANN K. A. 1968. - Die Suidae (Mammalia, Artiodactyla) aus den Dinotheriensanden (Unterpliozan+pont) Rheinhessens (Sudwestdeutschland). Schweizerische Palaontologische Abhandlung 86: 1-96.","KARADENIZLI L., SEYITOG LU G., SEN S., ARNAUD N., KAZANCi N., SARAC G. F ALCICEK C. 2005. - Mammal bearing late Miocene tuffs of the Akkasdagwi region; distribution, age, petrographical and geochemical characteristics, in SEN S. (ed.), Geology, mammals and environments at Akkasdagwi, late Miocene of Central Anatolia. Geodiversitas 27 (4): 553-566 (this volume).","KAZANCi N., SEN S., SEYITOG LU G., BONIS L. DE, BOUVRAIN G., ARAZ H., VAROL B. F KARADENIZLI L. 1999. - Geology of a new late Miocene mammal locality in Central Anatolia, Turkey. Comptes Rendus de l'Academie des Sciences, Paris, Sciences de la Terre et des Planetes 329: 503-510.","KOSTOPOULOS D. S. 1994. - Microstonyx major (Suidae, Artiodactyla) from the late Miocene locality of \"Nikiti-1\", Macedonia, Greece; some remarks about the species. Bulletin of the Geological Society of Greece 30 (1): 341-355.","KOSTOPOULOS D. S. 2005.- The Bovidae (Mammalia, Artiodactyla) from the late Miocene of Akkasdagwi, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkasdagwi, late Miocene of Central Anatolia. Geodiversitas 27 (4): 747-791 (this volume).","KOSTOPOULOS D. S. F SARAC G. 2005. - Giraffidae (Mammalia, Artiodactyla) from the late Miocene of Akkasdagwi, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkasdagwi, late Miocene of Central Anatolia. Geodiversitas 27 (4): 735-745 (this volume).","KOSTOPOULOS D. S., SPASSOV N. F KOVACHEV D. 2001. - Contribution to the study of Microstonyx: evidence from Bulgaria and the SE European populations. Geodiversitas 23 (3): 411-437.","KOUFOS G. D. F VLACHOU T. D. 2005. - Equidae (Mammalia, Perissodactyla) from the late Miocene of Akkasdagwi, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkasdagwi, late Miocene of Central Anatolia. Geodiversitas 27 (4): 633-705 (this volume).","LIU D. S., LI C. K. F ZHAI R. J. 1978. - [Pliocene vertebrates of Lantian, Shensis. Professional Papers on Stratigraphy and Paleontology 7: 149-200 (in Chinese).","LIU L. P. 2003. - Chinese Fossil Suidea: Systematics, Evolution and Paleoecology. University Printing House, Helsinki, 41 p.","LIU L. P., KOSTOPOULOS D. S. F FORTELIUS M. 2004 . - Late Miocene Microstonyx remains (Suidae, Artiodactyla) from Hezheng areas. Geobios 37 (1): 49-64.","NOW DATABASE 2003. - Neogene of the Old World. Database of fossil mammals: www.helsinki. fi/science/now","OZANSOY F. 1965. - Etude des gisements continentaux et de mammiferes du Cenozoiquede Turquie. Memoires de la Societe geologique de France N. S. 102: 5-92.","PEARSON H. 1928. - Chinese fossil Suidae. Palaeontologia Sinica C (5): 1-75.","PICKFORD M. 1988. - Revision of the Miocene Suidae of the Indian Subcontinent. Munchener Geowissenschaftliche Abhandlungen, series A, 12 (1): 1-95.","PICKFORD M. 1993. - Old world suoid systematics, phylogeny, biogeography and biostratigraphy. Paleontologia i Evolucio 26-27: 237-269.","PICKFORD M. F ERTURK C. 1979. - Suidae and Tayassuidae from Turkey. Bulletin of the Geological Society of Turkey 22: 141-154.","SEN S. 1997. - Magnetostratigraphic calibration of the European Neogene mammal chronology. Palaeogeography Palaeoclimatology Palaeoecology 133: 181-20.","SENYUREK M. S. 1952. - A study of the Pontian fauna of Gokdere (Elmadagi), southeast of Ankara. Turk Tarih Kurumu Belleten 16: 449-492.","SICKENBERG O. 1975. - Die Gliederung des hoheren Jungtertiars und Altquartars in der Turkei nach Vertebraten und ihre Bedeutung fur die interna- tionale Neogen-Stratigraphie. Geologisches Jahrbuch B, 15: 1-167.","STEININGER F. F. 1999. - Chronostratigraphy, geochronology and biochronology of the Miocene \"European Land Mammal Mega-Zones\" (ELMMZ) and the Miocene \"Mammal-Zones (MN-Zones)\", in ROSSNER G. E. F HEISSIG K. (eds), The Miocene Land Mammals of Europe. Verlag Dr. Friedrich Pfeil, Munchen: 9-24.","STEININGER F. F., BERGGREN W. A., KENT D. V., BERNOR R. L., SEN S. F AGUSTIJ. 1996. - Circum- Mediterranean Neogene (Miocene-Pliocene) marine-continental chronologic correlations of European mammal units, in BERNOR R. L., FAHLBUSCH V. F MITTMANN H. W. (eds), The Evolution of Western Eurasian Neogene Mammal Faunas. Columbia University Press, New York: 7-46.","TANG Y. J., LIU Z. Q., CHEN D. F CHEN L. Q. 1985. - The fossil Suidae from late Miocene of Binxian, Shaanxi. Vertebrata PalAsiatica 23 (1): 60-68.","TROFIMOV B. A. 1954. - The fossil suids of the genus Microstonyx, in Tertiary Mammals, part 2: on the Mammalia of the southern SSSR and Mongolia. Doklady Akademii Nauk SSSR 47: 61-99.","VAN DER MADE J. 1997. - The fossil pig from the late Miocene of Dorn-Durkheim 1 in Germany. Courier Forschungs-Institut Senckenberg 197: 205- 230.","VAN DER MADE J. F HUSSAIN S. T. 1989 . - \"Microstonyx\" major (Suidae, Artiodactyla) from Nagri. Estudios Geologicos 45: 409-416.","VAN DER MADE J. F MOYA- SOLA S. 1989 . - European Suinae (Artiodactyla) from the late Miocene onwards . Bollettino della Societa Paleontologica Italiana 28 (2-3): 329-339.","VAN DER MADE J., MONTOYA P. F ALCALA L. 1992. - Microstonyx (Suidae, Mammalia) from the upper Miocene of Spain. Geobios 25 (3): 395-413."]}

Dietary analyses of herbivorous mammals are important for paleoecological reconstruction. Several methods applicable to fossil teeth have been developed lately. The mesowear method based on wear-induced occlusal shape and relief of ungulate molars has proven to be a robust method for dietary analysis. In its original form it can only be used for selenodont, plagiolophodont, and ectolophodont ungulate molars, but the principle can be extended to other kinds of tooth morphology. We introduce a new method of dietary analysis for proboscideans similar to the mesowear method, based on angle measurements from worn dentin valleys reflecting the relief of enamel ridges. The enamel ridges should be heavily worn when the abrasiveness of diet increases, resulting in lower occlusal relief and larger angles. For testing this, we compared the mesowear angles with stable carbon isotope values from dental enamel from populations of extant and fossil species from localities from Kenya and India. This enables us to compare diet and tooth wear in proboscideans, because the stable carbon isotope ratios in tropical environments provide a reliable standard for assessing the relative amounts of C4 and C3 plants in diet, and most of the C4 plants are grasses, which should be reflected in the mesowear signal.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Dietary analyses of herbivorous mammals are important for paleoecological reconstruction. Several methods applicable to fossil teeth have been developed lately. The mesowear method based on wear-induced occlusal shape and relief of ungulate molars has proven to be a robust method for dietary analysis. In its original form it can only be used for selenodont, plagiolophodont, and ectolophodont ungulate molars, but the principle can be extended to other kinds of tooth morphology. We introduce a new method of dietary analysis for proboscideans similar to the mesowear method, based on angle measurements from worn dentin valleys reflecting the relief of enamel ridges. The enamel ridges should be heavily worn when the abrasiveness of diet increases, resulting in lower occlusal relief and larger angles. For testing this, we compared the mesowear angles with stable carbon isotope values from dental enamel from populations of extant and fossil species from localities from Kenya and India. This enables us to compare diet and tooth wear in proboscideans, because the stable carbon isotope ratios in tropical environments provide a reliable standard for assessing the relative amounts of C4 and C3 plants in diet, and most of the C4 plants are grasses, which should be reflected in the mesowear signal.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Dietary analyses of herbivorous mammals are important for paleoecological reconstruction. Several methods applicable to fossil teeth have been developed lately. The mesowear method based on wear-induced occlusal shape and relief of ungulate molars has proven to be a robust method for dietary analysis. In its original form it can only be used for selenodont, plagiolophodont, and ectolophodont ungulate molars, but the principle can be extended to other kinds of tooth morphology. We introduce a new method of dietary analysis for proboscideans similar to the mesowear method, based on angle measurements from worn dentin valleys reflecting the relief of enamel ridges. The enamel ridges should be heavily worn when the abrasiveness of diet increases, resulting in lower occlusal relief and larger angles. For testing this, we compared the mesowear angles with stable carbon isotope values from dental enamel from populations of extant and fossil species from localities from Kenya and India. This enables us to compare diet and tooth wear in proboscideans, because the stable carbon isotope ratios in tropical environments provide a reliable standard for assessing the relative amounts of C4 and C3 plants in diet, and most of the C4 plants are grasses, which should be reflected in the mesowear signal.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Fil: Raia, Pasquale. Università degli Studi di Napoli Federico II; Italia Fil: Passaro, Federico. Università degli Studi di Napoli Federico II; Italia Fil: Carotenuto, Francesco. Università degli Studi di Napoli Federico II; Italia Fil: Maiorino, Leonardo. Università degli Studi Roma Tre; Italia Fil: Piras, Paolo. Università degli Studi Roma Tre; Italia Fil: Teresi, Luciano. Università degli Studi Roma Tre; Italia Fil: Meiri, Shai. Tel Aviv University; Israel Fil: Itescu, Yuval. Tel Aviv University; Israel Fil: Novosolov, Maria. Tel Aviv University; Israel Fil: Baiano, Mattia A. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Baiano, Mattia A. Instituto Catalán de Paleontología Miquel Crusafont; España Fil: Baiano, Mattia A. Universidad Nacional de Río Negro; Argentina Fil: Martínez, Ricard. Universitat Autònoma de Barcelona; España Fil: Fortelius, Mikael. University Of Helsinski; Finlandia Luxuriant, bushy antlers, bizarre crests, and huge, twisting horns and tusks are conventionally understood as products of sexual selection. This view stems from both direct observation and from the empirical finding that the size of these structures grows faster than body size (i.e., ornament size shows positive allometry).We contend that the familiar evolutionary increase in the complexity of ornaments over time in many animal clades is decoupled from ornament size evolution. Increased body size comes with extended growth. Since growth scales to the quarter power of body size, we predicted that ornament complexity should scale according to the quarter power law as well, irrespective of the role of sexual selection in the evolution and function of the ornament. To test this hypothesis, we selected three clades (ammonites, deer, and ceratopsian dinosaurs) whose species bore ornaments that differ in terms of the importance of sexual selection to their evolution.We found that the exponent of the regression of ornament complexity to body size is the same for the three groups and is statistically indistinguishable from0.25.We suggest that the evolution of ornament complexity is a by-product of Cope’s rule. We argue that although sexual selection may control size in most ornaments, it does not influence their shape.

This thesis study examines how different species traits are related to the evolutionary survivorship of land mammal taxa and how the trait structure of mammalian communities is related to the climate, habitats and biogeography at the continental level. The study uses large datasets of both fossil and present land mammals from different continents. The results of the evolutionary survivorship of land mammal taxa demonstrate a recurring pattern whereby large mammal genera and species have higher origination and extinction rates and therefore shorter durations. This observed pattern can be explained because of behaviours such as hibernation, torpor and burrowing, collectively termed ‘‘sleep-or-hide’’ (SLOH). SLOH behaviour is more common in small mammals, and they are therefore better protected from environmental extremes than are large mammals, resulting in higher average survivorship and lower origination probabilities. The results of the relationship between species traits and climate provide a reasonably accurate estimate of precipitation values for today’s world, having the strongest correlations between annual precipitation and absolute and relative numbers of species, diet, tooth crown height, and diet and tooth crown height combined. The structures of tropical mammalian communities indicate that the community structure significantly differs between all of the studied vegetation categories, being the highest at major and minor ecosystem levels. The results demonstrate that community structure defined by both dietary and locomotor adaptations is a powerful discriminator of tropical ecosystems and habitats across the continents, but body mass should be interpreted with caution when the research question pertains to multiple continents. In North America climatic differentiation exists at the scale where mammals are divided into 11 distinct faunas and, in Europe, at the scale where there are five faunas. Functional trait differentiation in body mass occurs at a larger spatial scale in North America (eight distinct faunas), but locomotor differentiation is present at all spatial scales, and dietary differentiation is not present at any scale. No significant differentiation in any functional trait at any scale is present in Europe. Since the functional traits (body mass, locomotion, diet) are observable in the fossil record, community-level functional trait analysis has the potential to be used to reconstruct past climatic gradients. Tämä väitöskirja tutkii kuinka erilaiset lajien ominaisuudet liittyvät maanisäkkäiden evolutiiviseen selviytymiseen ja kuinka lajien ominaisuuksiin perustuva nisäkäsyhteisöjen rakenne liittyy ilmastoon, elinympäristöihin ja eliömaantieteeseen. Tutkimuksessa käytettiin laajoja tietokantoja sekä fossiilisista että nykyisistä maanisäkkäistä eri mantereilla. Tulokset maanisäkkäiden evolutiivisesta selviytymisestä osoittavat säännöllisen ilmiön, jossa isommilla nisäkässuvuilla ja -lajeilla on suurempi lajiutumisen ja sukupuuttoon kuolemisen tahti ja siten lyhyempi ajallinen kesto. Tulosta voidaan selittää niiden lajien avulla, jotka viettävät talviunta, horrostavat tai kaivautuvat maahan, ‘‘sleep-or-hide’’ (SLOH) -lajeilla. Koska SLOH-käyttäytyminen on yleisempää pienillä nisäkkäillä, ovat ne siten keskimäärin enemmän suojattuja ympäristön ääriolosuhteilta kuin isommat nisäkkäät ja näin ollen kokonaisuudessaan vaikuttavat pienempien lajien korkeampaan keskimääräiseen selviytymiseen ja matalampaan lajiutumistodennököisyyteen. Tulokset lajien ominaisuuksien suhteesta ilmastoon tuottavat kohtuullisen tarkat arviot nykyajan sadannasta, suurimpien korrelaatioiden ollessa vuotuisen sadannan ja lajien absoluuttisen ja suhteellisen lukumäärän, ruokavalion, hampaan kruunun korkeuden sekä ruokavalion ja hampaan kruunun korkeuden yhdistelmän välillä. Trooppisten lajien yhteisörakenteet taasen olivat tilastollisesti merkitsevästi erilaisia tutkittujen kasvillisuusluokkien välillä, ollen suurin ekosysteemitasolla. Etenkin nisäkäsyhteisöt jotka on jaettu ruokavalio- tai liikkumismuotoryhmiin, erottelevat hyvin mantereiden trooppiset ekosysteemit sekä elinympäristöt, mutta ruumiinkokoon perustuvaa ryhmittelyä kannattaa tulkita varovaisemmin, mikäli useita eri mantereita käsitellään tutkimuksessa. Pohjois-Amerikassa lajistot ovat ilmastoltaan erilaisia mittakaavassa, jossa nisäkkäät on jaettu 11 erilliseen lajistoon kun taasen Euroopassa ne jakautuvat alueellisesti viiteen erilaiseen lajistoon. Ruumiinpainon erojen perusteella Pohjois-Amerikan lajisto jakautuu kahdeksaan kokonaisuuteen, kun taasen liikkumismuodon perusteella lajistot eroavat kaikissa jakoluokissa. Ruokavalion perusteella ei lajistojen välille muodostunut tilastollisia eroja. Euroopassa yksikään lajisto ei eronnut tilastollisesti toisistaan minkään lajiominaisuuden perusteella millään jaolla. Tutkimuksessa saatujen tulosten perusteella voidaan todeta, että niitä havaittavia lajiominaisuuksia, joita voidaan erottaa myös fossiiliaineistosta (ruumiinpaino, liikkumismuoto, ruokavalio) voidaan käyttää nisäkäsyhteisöjen analysointiin ja muinaisten ilmasto-olosuhteiden mallintamiseen.