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A partial skeleton of Deinotherium (Proboscidea, Mammalia) from the late Middle Miocene Gratkorn locality (Austria)
Abstract
A disarticulated, though still roughly associated partial Deinotherium skeleton from the late Middle Miocene (late Sarmatian sensu stricto; 12.2–12.0 Ma) Gratkorn locality (Austria) is described. Based on dimensions and morphology of the material it can be determined as a medium-sized taxon of Deinotheriidae and definitively assigned to the genus Deinotherium. This specimen from Gratkorn confirms the osteological differences in the postcrania between Prodeinotherium and Deinotherium. As the diagnostically important p/3 is missing on the specimen it can only be assigned to Deinotherium levius vel giganteum. The Gratkorn specimen is one of not many skeletons of a medium-sized taxon of Deinotheriidae and one of only a few well-dated late Middle Miocene occurrences in Central Europe with associated dental and postcranial material.
... Based on several dental, cranial and postcranial features, European deinotheres are represented by the Early-Middle Miocene Prodeinotherium and the Middle-Late Miocene Deinotherium (e.g., Huttunen 2002a; Aiglstorfer et al. 2014a;Konidaris et al. 2017). Five species are considered valid here: Prodeinotherium cuvieri (Kaup, 1832a) from the early-middle Orleanian, Prodeinotherium bavaricum (von Meyer, 1831) from the late Orleanian-early Astaracian, Deinotherium levius Jourdan, 1861 from the late Astaracian, Deinotherium giganteum Kaup, 1829 from the Vallesian and Deinotherium proavum (Eichwald, 1831) (= Deinotherium gigantissimum Stefanescu, 1892) from the latest Vallesian-Turolian. ...
... Distinctive features among the species include: a) dental dimensions, b) traits of the mandible (shape of the symphysis and the mandibular angle), and c) morphology of the p3 and the dp2/DP2. Further details on the taxonomy of European deinotheres are given in Aiglstorfer et al. (2014a), Konidaris et al. (2017), Alba et al. (2020) and Konidaris and Tsoukala (2022). ...
... The Ldp4 stands between D. levius and D. giganteum, while the Wdp4 values (CT-scan measurements) are at the upper quartile of D. levius (Fig. 8). The HAM dp4 is plotted close to the dp4 from Sopron [Hungary; the deinothere material from Sopron is attributed to D. levius by Aiglstorfer et al. (2014a)], the larger specimens from La Grive and Massenhausen, and the smaller one from Montredon (Fig. 7). ...
During the Miocene, proboscideans reached their greatest diversification, and due to their marked evolutionary changes in dental size and morphology, they comprise an important biostratigraphic/biochronological tool. In this article, we study the proboscideans from the Late Miocene hominid locality Hammerschmiede (Germany), whose fossiliferous layers HAM 6, HAM 4 and HAM 5 are dated to 11.42, 11.44 and 11.62 Ma, respectively. The studied material consists of mandibular, tusk and cheek tooth specimens, which are attributed to the deinothere Deinotherium levius and the tetralophodont gomphothere Tetralophodon longirostris. An almost complete juvenile mandible of D. levius was CT-scanned and revealed that the erupting lower tusks represent the permanent ones. The mandible is most possibly associated with a lower deciduous tusk, and therefore these specimens capture the rare, and short in duration, moment of transition between deciduous and permanent lower tusks in fossil proboscideans and represent the first such example in deinotheres. The chronologically well-constrained proboscidean fauna from Hammerschmiede and the examination of other assemblages from European localities indicate that the coexistence of D. levius and T. longirostris characterizes the late Astaracian–earliest Vallesian, while Hammerschmiede may showcase the transition from the Middle Miocene trilophodont (Gomphotherium)-dominated faunas of central Europe to the Late Miocene tetralophodont-dominated ones. Finally, in order to decipher the dietary preferences of the Hammerschmiede Tetralophodon we performed dental mesowear angle analysis, which revealed a mixed-feeding diet with an important browsing component, significantly different from the heavily browsing one of Deinotherium known from other localities. Such distinct feeding habits between the taxa indicate niche partitioning, which allowed their sympatry.
... Most small mammals (insectivores, chiroptera, rodents, lagomorphs; Prieto et al. 2014), the Perissodactyla, Artiodactyla and Proboscidea found at Gratkorn have body masses well below 150 g or above 15 kg (Aiglstorfer et al. 2014a(Aiglstorfer et al. , 2014b(Aiglstorfer et al. , 2014c(Aiglstorfer et al. and 2014dVan der Made et al. 2014; see also Merceron et al. 2012;Costeur et al. 2013, andNOW database 2023). So, it seems unlikely that they are the source of the hairs found. ...
... Presumably, like modern carnivorans with a body mass >25 kg, Albanosmilus tended to ambush prey of its own weight and beyond (>100 kg), making it an obligate hunter of large mammals (Carbone et al. 1999;Wang et al. 2018;Barret 2021). In the case of Gratkorn, this could have been the equid Anchitherium, the suids Listriodon splendens and Parachleuastochoerus steinheimensis, the even-toed ungulate 'Palaeomeryx' and, rather improbably, the much larger Chalicotherium and the rhinos Aceratherium, Brachypotherium and Lartetotherium (Aiglstorfer et al. 2014a(Aiglstorfer et al. , 2014b(Aiglstorfer et al. , 2014dVan der Made et al. 2014). Chewing marks (dentalites; Hunt et al. 2018) on a proboscidean scapula (Deinotherium; Aiglstorfer et al. 2014d) were assigned to an unknown large carnivore ) but could be from Albanosmilus. ...
... In the case of Gratkorn, this could have been the equid Anchitherium, the suids Listriodon splendens and Parachleuastochoerus steinheimensis, the even-toed ungulate 'Palaeomeryx' and, rather improbably, the much larger Chalicotherium and the rhinos Aceratherium, Brachypotherium and Lartetotherium (Aiglstorfer et al. 2014a(Aiglstorfer et al. , 2014b(Aiglstorfer et al. , 2014dVan der Made et al. 2014). Chewing marks (dentalites; Hunt et al. 2018) on a proboscidean scapula (Deinotherium; Aiglstorfer et al. 2014d) were assigned to an unknown large carnivore ) but could be from Albanosmilus. However, capturing an adolescent deinothere, weighing more than 6 t (Aiglstorfer et al. 2014d), would have required social hunting behaviour comparable to that of extant lion prides ( Van den Hoek Ostende et al. 2006). ...
An association of eighteen coprolites (specimens 01–18) and one isolated coprolite (specimen 209,210) were found in a vertebrate fossil-rich paleosol at the Gratkorn site (south-eastern Austria; late Middle Miocene). The specimens consist mostly of calcium phosphate (apatite) and a matrix formed by microglobules. Coprolites 01–18 show cylindrical and spherical morphologies and are considerably smaller than the tube-shaped specimen 209,210, in which no inclusions were observed. In contrast, coprolites 01–18 contain numerous, highly altered bone fragments (sub-mm-sized long bones and several mm-sized trabecular bone remains) as well as hair imprints, plant detritus and palynomorphs. Based on composition, morphology, size, microstructure, and inclusions, and considering the body fossil record of this site, we assume the hyaenid Protictitherium and the barbourofelid Albanosmilus, as producers of coprolites 01–18 and 209,210, respectively. The preserved bone remains in specimens 01–18 suggest that Protictitherium fed on small vertebrates, but possibly also cracked bones of medium-sized animals. The hair imprints found were either from the hyaenid itself or its prey, while the plant material was probably ingested accidentally. The lack of inclusions in specimen 209,210 is related to the presumably hypercarnivorous diet of Albanosmilus, which was certainly the apex predator in this biome.
... The number of deinotheriid taxa recorded in Europe has been debated. Some recent authors still distinguish a single genus, Deinotherium Kaup, 1829(e.g., Böhme et al., 2012Pickford and Pourabrishami, 2013), whereas most authors distinguish the smaller Prodeinotherium Éhik, 1930 (early to middle Miocene) from the larger Deinotherium sensu stricto (middle to late Miocene; Gasparik, 1993;Huttunen and Göhlich, 2002;Huttunen 2002aHuttunen , 2002bHuttunen , 2004Duranthon et al., 2007;Markov, 2008a;Vergiev and Markov 2010;Aiglstorfer et al., 2014;Konidaris et al., 2017). Although the distinction between the two genera is not well supported on dental grounds, based on purported cranial and postcranial differences (Huttunen, 2002a(Huttunen, , 2004, we prefer to keep them distinct, in agreement with most recent studies (e.g., Konidaris et al., 2017), unless an in-depth review of the family as a whole supports otherwise. ...
... Subsequently, a second species, Deinotherium levius Jourdan, 1861 (type locality: La Grive-Saint-Alban M, France, MN7+8), has been alternatively distinguished (Gräf, 1957) or synonymized (Bergounioux and Crouzel, 1962a;Harris, 1973;Huttunen, 2002aHuttunen, , 2002b with the former. The same applies to Deinotherium gigantissimum Stefanescu, 1892 (type locality: Gaȋceana, Romania, Turolian), frequently synonymized with the type species (Bergounioux and Crouzel, 1962a;Harris, 1973;Huttunen, 2002aHuttunen, , 2002b, but most recently considered a distinct species (Göhlich, 1999;Garevski and Markov, 2011;Böhme et al., 2012;Pickford and Pourabrishami 2013;Aiglstorfer et al., 2014;Konidaris et al., 2017;Konidaris and Koufos, 2019). The correct name for D. gigantissimum is Deinotherium proavum (Eichwald, 1831), as already noted by Kretzoi (1965) and Codrea (1994) and subsequently recognized by more recent authors with the correct year of description (Pickford and Pourabrishami, 2013;Konidaris et al., 2017;Konidaris and Koufos, 2019). ...
... The morphology of the p3 is considered to be most diagnostic for distinguishing D. giganteum (with fused protoconid and metaconid, even though their apices are still distinct; e.g., Gasparik, 2001:pl. 1, fig. 3) from D. levius, which retains a more plesiomorphic condition (with individualized protoconid and metaconid; Böhme et al., 2012:fig. 4c; see also Pickford and Pourabrishami, 2013;Aiglstorfer et al., 2014). The p3 morphology of D. proavum is more similar in this regard to that of D. giganteum. ...
Three species of Deinotherium sensu stricto (Proboscidea, Deinotheriidae), i.e., excluding Prodeinotherium, generally considered to have nonoverlapping chronostratigraphic distributions, are currently recognized from the Miocene of Europe: Deinotherium levius (late Astaracian/Aragonian, MN7+8), Deinotherium giganteum (type species; Vallesian, MN9–MN10), and Deinotherium proavum (a senior synonym of Deinotherium gigantissimum; Turolian, MN11–MN13). Here we describe a sample of 26 cheek teeth from four roughly coeval localities of Ronda Oest de Sabadell (ROS), in the Vallès-Penedès Basin, northeastern Iberian Peninsula, dated to the latest Vallesian (∼9.4–9.1 Ma, MN10) on biostratigraphic grounds. The remains from ROS-D3 represent all the permanent upper and lower dentition and can be unambiguously assigned to D. proavum, the largest deinothere from Europe, based on their large size (well above the range of D. giganteum). Remains from the other localities (ROS-D2, ROS-D5, and ROS-D8) are smaller and generally overlap in size with both D. proavum and D. giganteum. However, an assignment to the former species is more likely given that these species are not known to co-occur and that their dental size ranges were already known to overlap partially. The remains of D. proavum from ROS represent the oldest record of this species, the occurrence of which in the late Vallesian had already been recorded in the slightly younger (∼9.1 Ma) locality of Sinap 49, Turkey. Given that D. proavum is recorded before the Vallesian/Turolian boundary, and that smaller individuals of this species overlap in size with D. giganteum from the Vallesian, caution is required when making biostratigraphic correlations based on late Miocene deinothere remains from Europe.
... Young adult specimens, such as, e.g. a Deinotherium levius vel giganteum partial skeleton, with not fully fused epiphyses, are therefore not disclosed separately but included in adult specimens. A delayed fusion of the long bones and continuation of growth beyond sexual maturity has been observed in modern proboscideans (Poole 1996; in males even up to the age of 3045 years; see discussion in Aiglstorfer et al. 2014b, this issue). Specimens only documented by postcranial elements are considered adult in all cases, if fusion of long bones is completed and no signs of attrition indicate a senile age. ...
... The partial skeleton of Deinotherium levius vel giganteum comprises many bones, which are strongly weathered (weathering stage 5;after Behrensmeyer 1978) and often the bone compacta is not preserved, but only bone spongiosa. It shows clear signs of a prolonged exposure, such as fragmentation on the surface and scavenging of larger carnivores (see discussion below; Fig. 3h;and Aiglstorfer et al. 2014b, this issue). This could be explained by the size of the bones, which are much larger than in all other species from Gratkorn, and therefore the skeleton was probably not covered so soon and bones not dislocated as deep into the palaeosol as observed in other specimens. ...
... This could be explained by the size of the bones, which are much larger than in all other species from Gratkorn, and therefore the skeleton was probably not covered so soon and bones not dislocated as deep into the palaeosol as observed in other specimens. This is well in accordance with recent decomposition data in modern elephants (Coe 1978;Conybeare and Haynes 1984; see discussion in Aiglstorfer et al. 2014b, this issue). Furthermore, the deinothere remains are sticking out over the top of the palaeosol and many remains were recovered from the uppermost part of it, which led to longer exposure near or on the surface and therefore stronger weathering during early diagenesis. ...
At the Gratkorn locality (Styria, Austria), a highly diverse, late Middle Miocene (late Sarmatian sensu stricto; 12.2–12.0 Ma) faunal assemblage is preserved in a palaeosol. It represents the first systematically excavated and well-documented continental Sarmatian site in Central Europe. Taphonomical analysis of the 700 large mammal specimen excavated so far has led to the following conclusions: (1) the level of diagenetic alteration is low, as primary (aragonitic) mineralisation in gastropod shells is preserved and teeth and bones of large mammals in general show a relatively low total REE content; (2) the high degree of disarticulation and fragmentation in large mammal bones is induced by hunting, scavenging, trampling, and neotectonics; (3) there are no signs for fluviatile transportation due to the general preservation features of the bones (e.g. no record of abrasion) and the still roughly associated fragments of individual bones and skeletons; and (4) local accumulation of large mammal bones is the result of scavenging. The fossil assemblage is considered to form a more or less autochthonous taphocoenosis without any significant time averaging (or faunal mixing) in terms of geologic resolution (contemporaneously deposited).
... The number of distinct deinotheriine genera is still a matter of debate. Some recent authors (Böhme et al. 2012;Pickford and Pourabrishami 2013) still distinguish a single genus (Deinotherium Kaup, 1829), but most other authors (e.g., Harris 1973Harris , 1978Gasparik 1993Gasparik , 2001Göhlich 1999;Huttunen 2002aHuttunen ,b, 2004Huttunen and Göhlich 2002;Duranthon et al. 2007;Markov 2008;Rasmussen and Gutierrez 2009;Sanders et al. 2010;Aiglstorfer et al. 2014;Konidaris et al. 2017;Alba et al. 2020) distinguish the smaller Prodeinotherium Éhik, 1930 (late Oligocene to early middle Miocene; but see Qiu et al. 2007 for a report of Prodeinotherium from the late Miocene of China) from the larger Deinotherium s.s. (late middle to late Miocene). ...
... Some authors recognized a single valid species of Prodeinotherium in Europe (Bergounioux and Crouzel 1962a;Göhlich 1999;Huttunen 2002aHuttunen ,b, 2004Huttunen and Göhlich 2002), but most current authors (Ginsburg and Chevrier 2001;Böhme et al. 2012;Pickford and Pourabrishami 2013;Aiglstorfer et al. 2014;Konidaris et al. 2017) distinguish Prodeinotherium cuvieri (Kaup, 1832) from Prodeinotherium bavaricum (von Meyer, 1831). ...
Deinotheres (Proboscidea, Deinotheriidae) are a clade of non-elephantiform proboscideans that originated in Africa and dispersed into Eurasia by the early Miocene. In Europe, deinotheres are first recorded in Greece during MN3, although they did not become a common faunal element throughout Europe until MN4. Early Miocene (MN3–MN4) deinothere remains from Europe are generally assigned to a different species (Prodeinotherium cuvieri) than those from the early middle Miocene (Prodeinotherium bavaricum; MN5–MN6). In the Vallès-Penedès Basin (NE Iberian Peninsula), Prodeinotherium remains are very scarce and largely remain unpublished. To clarify their taxonomic assignment, we describe the available material and compare it with that from elsewhere in Europe. Based on size and a few diagnostic occlusal details, we tentatively recognize both Prodeinotherium cf. P. cuvieri and Prodeinotherium cf. P. bavaricum in the basin. Although all the studied sites had previously been correlated to MN4, the recognition of P. cf. P. bavaricum at els Casots and les Escletxes is consistent with ongoing litho- and magnetostratigraphic studies suggesting a slightly younger age for these sites. The lack of Prodeinotherium remains in older (MN3) localities from the Vallès-Penedès Basin, where Gomphotherium is already recorded, further supports the view that deinotheres dispersed into Western Europe somewhat later than gomphotheres.
... Dental terminology for proboscidean teeth follows Tassy (1996b) and Aiglstorfer et al. (2014). Measurements were taken with digital calipers (precision 0.1 mm). ...
... Based on the morphology and relatively large size of the specimens, the finds from Ravanica and Adrani most likely both represent Deinotherium giganteum. For the Ravanica material, the attribution to D. levius also can not be excluded, since the diagnostically important p3 (Gräf 1957;Aiglstorfer et al. 2014) did not represent part of the find. However, it must be noted that D. levius is not uniformly accepted as a valid taxon, and other authors (e.g. ...
The paper provides an overview of the Neogene fossil mammals recorded in the vicinity of Kraljevo in Central Serbia. Large mammals are represented exclusively by the accidental finds of proboscidean dental specimens, deposited both in the Čačak-Kraljevo and Gruža basins. We provide re-descriptions and comparative analyses of the proboscidean material. The following taxa are recorded: Deinotherium giganteum at Ravanica and Adrani; Gomphotherium angustidens at Bogutovac; Tetralophodon longirostris at Godačica and at an unidentified sand pit (majdan) near Kraljevo; Anancus sp. at Miločaj; Proboscidea indet. at Oplanići. Small fossil mammals are recorded at three sites. The Progorelica assemblage is currently attributed to the MN6 zone of the Middle Miocene (Langhian), based on the finds of Eulipotyphla sp., Eomyops sp., Cricetodon sp., Megacricetodon sp. and Alloptox sp. – one of the first finds of this Asian genus in the Balkan Peninsula. At Tavnik, more than 120 remains of small mammals were attributed to the MN9 zone of the Late Miocene (Tortonian); the site yielded remains of Eulipotyphla indet., Prolagus sp., Megacricetodon similis and Miodyromys sp. Two fossil teeth of Talpidae sp. were also recorded at Oplanići.
... Dental terminology for proboscidean teeth follows Tassy (1996b) and Aiglstorfer et al. (2014). Measurements were taken with digital calipers (precision 0.1 mm). ...
... Based on the morphology and relatively large size of the specimens, the finds from Ravanica and Adrani most likely both represent Deinotherium giganteum. For the Ravanica material, the attribution to D. levius also can not be excluded, since the diagnostically important p3 (Gräf 1957;Aiglstorfer et al. 2014) did not represent part of the find. However, it must be noted that D. levius is not uniformly accepted as a valid taxon, and other authors (e.g. ...
... In particular, the Suomusjärvi specimen is close in size to the Deinotherium giganteum specimen from Munich described by Stromer (1938). Another example of a comparable medium-sized Deinotherium is the partial skeleton from the late Middle Miocene locality of Gratkorn, Austria, identified as Deinotherium levius or early D. giganteum, although this skeleton lacks humeri almost entirely (Aiglstorfer et al., 2014). Erosion of the Suomusjärvi specimen could have had some (but probably not major) effect on its measurements. ...
... However, the comparably small size of the specimen for the genus Deinotherium could even point at a late Middle Miocene (ca. 16-11 Ma) age, because Aiglstorfer et al., 2014). This is, however, highly speculative, especially because the age or growth stage of the individual animal cannot be reliably estimated. ...
Abstract
We discuss a proboscidean bone fragment discovered in southern Finland, including the morphological analysis of the bone, as well as pollen and diatom analyses from sediment contained in the marrow cavity. Preliminary analysis of the bone suggested petrification and thus an apparently old age, while the microfossil assemblages include numerous unequivocally pre-Quaternary pollen, spore, and diatom types. A Miocene age for the bone is determined based on the presence of the diatom genus Alveolophora, indicating a minimum age of 5 Ma, and based on the earliest appearance of proboscideans outside Africa, setting a maximum age of 19 Ma. Based on morphology, the bone is determined as a partial humerus of the left foreleg of a large proboscidean. The bone is tentatively assigned to cf. Deinotherium sp., which is consistent with the diatom-based minimum age. The pollen assemblage is rich in spores of shoreline pteridophytes, while the diatom assemblage is also consistent with a shoreline freshwater environment, suggesting that the bone was deposited post-mortem near the shore of a lake or a stream. Miocene sediments do not currently exist in southern Finland or in the near vicinity. This implies that the bone has been transported over a considerable distance. Due to the discovery of the bone in early-Holocene Baltic Sea clay, the final transport phase and deposition must have taken place via iceberg rafting. This was likely preceded by one or more phases of glacial and/or glacio-fluvial transport. While we are unable to conclusive ascertain the region of origin, the alkaline composition of the contained sediment and diatoms point towards the Russian Plain region in the east. This specimen represents the oldest mammalian bone discovered in Finland and the northernmost discovery of a Miocene proboscidean bone in Europe.
... The largest forms of Deinotheriidae are within the last genus Deinotherium. The taxonomy of this genus has been somewhat chaotic for many decades (Gräf 1957;Bergounioux and Crouzel 1962;Harris 1973Harris , 1978Antoine 1994;Huttunen 2000;Markov 2008;Vergiev and Markov 2010;Böhme et al. 2012;Pickford and Pourabrishami 2013;Aiglstorfer et al. 2014). For this paper the morphospecies concept defended by Aiglstorfer et al. (2014) was followed, along with the African species (Arambourg 1934), which included the next species within the genus: D. levius, D. giganteum, D. proavum (= D. gigantissimum and D. thraceiensis) and D. bozasi. ...
... The taxonomy of this genus has been somewhat chaotic for many decades (Gräf 1957;Bergounioux and Crouzel 1962;Harris 1973Harris , 1978Antoine 1994;Huttunen 2000;Markov 2008;Vergiev and Markov 2010;Böhme et al. 2012;Pickford and Pourabrishami 2013;Aiglstorfer et al. 2014). For this paper the morphospecies concept defended by Aiglstorfer et al. (2014) was followed, along with the African species (Arambourg 1934), which included the next species within the genus: D. levius, D. giganteum, D. proavum (= D. gigantissimum and D. thraceiensis) and D. bozasi. Moreover, the gigantic, nearly-complete specimen from Ezerovo, described by Kovachev and Nikolov (2006) as D. thraceiensis, is very problematic. ...
In recent decades there has been a growing interest in proboscideans’ body size, given that mass is highly correlated with biological functions. Different allometric equations have been proposed in the recent decades to estimate their body masses, based on a large number of living examples. However, the results obtained by these formulae are not accurate because extinct animals often had different body proportions and some were outside the size range of extant samples. Here the body mass of a large number of extinct proboscideans has been calculated by the Graphic Double Integration volumetric method which is based on technical restorations from graphical reconstructions of fossils employing photos, measurements and comparative anatomy of extant forms. The method has been tested on extant elephants with highly accurate results. The reconstructions necessary to apply this method give important information such as body proportions. On the other hand, equations to calculate the skeletal shoulder height have been developed, with a large number of published shoulder heights being recalculated. From the shoulder heights, several equations were created to find out the body mass of a series of extant and extinct species. A few of the largest proboscideans, namely Mammut borsoni and Palaeoloxodon namadicus, were found out to have reached and surpassed the body size of the largest indricotheres. Bearing this in mind, the largest land mammal that ever existed seems to be within the order of Proboscidea, contrary to previous understanding.
... Distinguishing features among these species include (a) dental dimensions, (b) traits of the mandibular symphysis and angle, and (c) morphology of the p3 and the dp2/DP2. Further details on the taxonomy of European deinotheres are given in Aiglstorfer et al. (2014), Konidaris et al. (2017Konidaris et al. ( , 2019Konidaris et al. ( , 2023a, Alba et al. (2020) and Konidaris and Tsoukala (2022). ...
In this article, we describe so far unpublished proboscidean specimens from several Late Miocene localities of Romania. A partial mandible and the complete upper/lower cheek tooth rows of a deinothere individual from the site of Gherghești 1 belong to Deinotherium proavum and comprise one of the few examples of entire cheek tooth rows of the same individual of this species. Gherghești 1 is geographically close to Mânzaţi from where the celebrated skeleton of “Deinotherium gigantissimum” was discovered at the end of the nineteenth century, and thus further highlights the importance of Romania in the study of this emblematic deinothere. Deinotherium proavum represents the last deinothere species in Europe and corresponds to the terminal stage of the size increase characterizing the evolution of European deinotheres. Two zygodont molars are attributed to the rare “Mammut” cf. obliquelophus and add to the scarce record of “Mammut” in the Miocene of Eurasia. They document the secure presence of “Mammut” in the Miocene of Romania. The small size of the studied molars compared to known specimens of the Pliocene “Mammut” borsoni and the weak development of the distal cingulum in the lower third molars may have taxonomic and biostratigraphic importance. Furthermore, the presence of an amebelodontid is documented by a large-sized and dorsoventrally flattened lower tusk fragment that shows tubular dentine in its inner part and is attributed to the tetralophodont shovel-tusker Konobelodon. This specimen marks the first record of the genus in Romania. Finally, the biostratigraphic distribution of the taxa is discussed.
... The most recent investigations of European deinotheres approve five different valid morphospecies (Böhme et al. 2012) or chronospecies (Pickford and Pourabrishami 2013), whereas previous studies were in favour of four species (Gasparik 1993(Gasparik , 2001Markov 2008a;Vergiev and Markov 2010) or even only two species (Huttunen 2002). In this present investigation, I follow the five species concept, but also the two genera concept as proposed by Éhik (1930), which refers to dental and postcranial differences in the genera Prodeinotherium and Deinotherium (see also Aiglstorfer et al. 2014). Therefore, in this work, I consider the following European taxa (from oldest to youngest) to be valid: Prodeinotherium cuvieri (Kaup, 1832), P. bavaricum (von Meyer, 1831, Deinotherium levius Jourdan, 1861, D. giganteum Kaup, 1829 andD. ...
Presented here is a middle Miocene proboscidean fauna—deinotheres and gomphotheres—from lignites of the coal mine in Gračanica (Bugojno Basin) in Bosnia–Herzegovina. The material, which exclusively comprises teeth (n = 42), is described and
systematically discussed. The majority of the studied fossils represent Prodeinotherium bavaricum, but gomphotheres are also a common faunal element and are identified as Gomphotherium angustidens and tentatively as cf. Gomphotherium subtapiroideum. Noteworthy is the occurrence of an amebelodontine, identified as cf. Protanancus, and represented in the present material only by a single milk tooth. Protanancus is generally known from Africa and Asia and extremely rarely documented in southeastern Europe so far. The Gračanica fauna is supposed to correlate with the Mammalian Neogene unit
late MN5 or early MN6.
... Five species are considered valid here: P. cuvieri (Kaup, 1832) from the early-middle Orleanian, P. bavaricum (von Meyer, 1831) from the late Orleanian-early Astaracian, D. levius Jourdan, 1861 from the late Astaracian, D. giganteum Kaup, 1829 from the Vallesian, and D. proavum (von Eichwald, 1831) (= D. gigantissimum Stefanescu, 1892) from the Turolian. Distinctive features among the species include (a) dental dimensions, (b) traits of the mandible (shape of the symphysis and the mandibular angle), and (c) the morphology of the p3 and the dp2/DP2 [see below; for further details on the taxonomy of European deinotheres, see Aiglstorfer et al. (2014), Konidaris et al. (2017) and references cited in both]. ...
In this article, we present new proboscidean remains from the late Miocene (Turolian) of Samos Island (Greece), which are stored in the old Samos collections of Darmstadt, Frankfurt a.M. (Germany), Lausanne (Switzerland), and Vienna (Austria), and originate from the excavations or fossil collections that took place on the island at the end of the nineteenth century and the beginning of the twentieth century. The specimens belong to juvenile individuals of deinotheres, choerolophodonts and amebelodonts. The deinothere material is attributed to the last European huge-sized deinothere, Deinotherium proavum. The described skull from Samos is currently the most complete specimen of all known Miocene juvenile deinotheres from Eurasia and Africa. The majority of the Samos choerolophodont specimens belong to the advanced morph of Choerolophodon pentelici, whereas one shows more archaic features and belongs to the primitive evolutionary stage of this species. This more primitive morph could originate from the lower fossiliferous horizons of Samos, which are dated to the early Turolian. The third proboscidean is attributed to the tetralophodont shovel-tusker Konobelodon atticus, a rare taxon in the Samos fauna. Together with the previously described zygodont Mammut from Samos, these four proboscideans are typical of the Turolian proboscidean fauna of southeastern Europe. We discuss the biostratigraphy of the Samos proboscideans with the aim of unraveling some aspects of the chronological range of the late Miocene proboscideans, focusing in particular on the Southern Balkans and Turkey.
... She identified it as Deinotherium levius Jourdan, 1861 and suggested a late Sarmatian age for the specimen. Nevertheless, Mottl (1958) mentioned that Deinotherium species identifications, solely based on tusks, are difficult and are not conclusive for stratigraphic purposes (see Aiglstorfer et al. 2014). ...
This paper describes the section and fossil content of a former gravel pit in the Eastern Styrian Basin (SE Austria), which exposes sediments of a fluvial system, ranging from within channel to overbank environments. A predominately terrestrial gastropod fauna of 15 species so far, was recovered from a palaeosol formed in a moist and vegetated, floodplain or abandoned channel. Up-section, a shallow freshwater pond/lake developed within the floodplain, settled by fishes, molluscs and ostracods. By integrating regional geological and biostratigraphical data derived from the terrestrial gastropod fauna as well as from the other recovered biota, these strata are of late middle Miocene (late Sarmatian s.str.) age. Hence, this fossil site provides a rare insight into the terrestrial habitats in the hinterland of the Sarmatian Sea and their biota, which are otherwise barely known in Central Europe.
We have described here proboscideans fossil remnants which belong to six species: Prodeinotherium pentapotamiae, Gomphotherium browni, Choerolophodon corrugatus, Protanancus chinjiensis, and Elephas planifrons. The described specimens are collected from six different localities of the Siwaliks. These localities are Bhilomar, Chinji type section, Lawa, Dhok Bun Ameer Khatoon (Chinji Formation), Ochri (Dhok Pathan Formation) and Rathian (Pinjor Formation), belongs to the Lower to Upper Siwaliks, district Chakwal and Jhelum, Punjab, Pakistan. The specimens were collected from these localities after extensive fieldwork from 2015 to 2020, which confirms the presence of large mammalian fauna during the Middle Miocene to Late Pleistocene. Prodeinotherium pentapotamiae is rare taxa in the Middle Siwaliks of Pakistan, first time discovered from new fossil section Satroma wali dhal in the vicinity of the Dhok Bun Ameer Khatoon locality. Bhilomar is a less explored fossil locality belongs to the Chinji Formation, Gomphotherium browni and Choerolophodon corrugatus is reported from the Chinji type section. A beautiful juvenile mandibular ramus of Protanancus chinjiensis has been recovered first time from Lawa and Elephas planifrons were recovered from Rathian which belongs to Pinjor Formation dated as Plio-Pleistocene age of Upper subgroup of Siwaliks. Based on palaeoenvironment and stratigraphy the age of these localities is suggested as Miocene to Pleistocene. The recovered proboscideans fossil fauna indicates about the climatic changes and confirms that more seasonal extensive grasslands and savannah like ecosystem were present at that time.
Proboscideans (Mammalia: Proboscidea) originated during the Eocene (perhaps already during the Paleocene) in Africa. Their fossil record narrates an amazing evolutionary history, ranging from the Paleogene to the Quaternary. Proboscideans experienced in the past a great diversification and wide distribution in Africa, Europe, Asia, and the Americas. They persist until today with only two genera, Loxodonta and Elephas, geographically confined in regions of Africa and Asia, respectively. The review of the fossil record of the Neogene proboscideans (excluding the members of Elephantidae that are treated elsewhere) in Greece revealed the presence of deinotheres (Deinotheriidae), mammutids (Mammutidae), choerolophodonts (Choerolophodontidae), amebelodonts (Amebelodontidae), tetralophodont gomphotheres (Gomphotheriidae), and stegodonts (Stegodontidae) in more than fifty localities, ranging from the early Miocene to the Early Pleistocene. Fourteen taxa are here considered valid, three of them (Choerolophodon chioticus, C. pentelici, and Konobelodon atticus) erected from type localities in Greece. The most diverse localities are Pikermi and Samos, where at least four proboscidean species have been recorded. The peak in taxonomic diversity occurred during the Turolian (late Miocene). The Greek proboscidean fossil record contains several highlights. The earliest appearance of the family Deinotheriidae in Europe is documented in Gavathas of Lesvos Island, and it is the proboscidean family with the widest temporal distribution in Greece. A deinotheriid skull from Samos Island is so far the most complete juvenile one known from Eurasia and Africa. Choerolophodon presents the widest temporal distribution among the genera in Greece, and where present, it is the dominant genus in terms of abundance. The rich choerolophodont sample allows the distinction into evolutionary stages and renders the genus as biostatigraphically important for Southeastern Europe. The late Miocene Anancus from Chomateri represents the first appearance of the genus in Greece and one of the earliest occurences in Europe. The sample of the late Pliocene Mammut borsoni from Milia, Grevena, is the richest one of this species, including partial skeletons, the longest upper tusks ever recorded in the world and the most complete mandible in Europe. During the Pliocene–Early Pleistocene, the most frequent and widespread proboscidean is the last European gomphothere Anancus arvernensis. Finally, the Siatista Stegodon is the first evidence of the presence of stegodontids in Europe.
The Late Miocene fossiliferous locality of Kerassia is located in northwestern Euboea, near the homonymous village. In the current undergraduate thesis material from all the excavations that have taken place in the area was studied. The material consists of bones from the most common mammal families that are represented in the Turolian faunas of the Balkan-Iranian Province. Perissodactyla (Equidae, Rhinocerotidae, Chalicotheriidae) and artiodactyla (Suidae, Giraffidae, Bovidae) are the most common taxa in the fauna of Kerassia. Other important taxa that were identified in the area are Aves, Proboscidea and Carnivora, which are considered rare findings. In the locality of Kerassia, two fossiliferous horizons have been recognized, as mentioned in previous works. The lower one includes the sites K2, K3 and K4, while the upper one includes the sites K1 and K6. The palaeoenvironment is characterized as mosaic savanna-woodland, but the species distribution in the two horizons indicates a transition into more forested conditions from the lower to the upper horizon.
In the northern Swiss Plateau and Jura Mountains, non marine Paleogene and Neogene deposits of the Swiss Molasse Basin or linked to the Upper Rhine Graben are examined in detail. The Late Eocene (Middle–Late Priabonian) is characterized by charophytes of the Vasiformis–Tuberculata Zone (Diegten Süsswasserkalk) and Vectensis Zone (lower Tuberculata Superzone, Oberdorf Süsswasserkalk, Terre jaune). The Rupelian and Early Chattian appear as little developed and extended. The Rupelian is characterized by charophytes of the Pinguis Zone (upper Tuberculata Superzone, basal Conglomérat de Porrentruy) and Major Zone (Conglomérat de Porrentruy, lower “Molasse alsacienne”, Marnes rouges). In the Neuchâtel region of central Jura, a paleokarst marine filling of UMM (Montmollin) yielded nannofossils of Early Rupelian (NP 21) age. Early Chattian sediments are for the first time precisely dated by small mammals (MP 25–26a) and typical charophytes of the Microcera Zone (Calcaire d’eau douce de Trois-Rods). The biostratigraphy of charophytes for the Early Miocene to early Late Miocene (MN 1–9) can also be defined more precisely ranging from Nitida Zone to Etrusca Zone. The biozonation of charophytes for the Swiss Paleogene and Neogene (SPN) is revised and completed by the creation of 11 assemblage zones SPN-EC 1–2 (Eocene charophytes, Late Eocene), SPN-OC 1–2 (Oligocene charophytes, Rupelian), SPN-OC 3–5 (Chattian), SPN-MC 1–2 (Miocene charophytes, Aquitanian), SPN-MC 3 (Burdigalian) and SPN-MC 4 (Langhian–Early Tortonian).
Presented here are the deinotheriid dental and postcranial remains from the Late Miocene localities Pikermi and Halmyropotamos (Greece). The study and comparison of the available juvenile dental material from Pikermi with other relevant specimens from Europe showed that it belongs to the huge-sized Deinotherium proavum (= D. gigantissimum). Additionally, several postcranial specimens from Pikermi, as well as from Halmyropotamos, present deinotheriid features, which distinguish them from elephantoids, and permit their attribution also to D. proavum. This species is known from the Turolian and represents the terminal evolutionary stage of the European deinotheres. Its presence in Pikermi and Halmyropotamos is compatible with the middle Turolian (MN 12) age of the localities. Additionally, the taxonomy of European deinotheres is discussed, as well as the biostratigraphical and geographical distribution of D. proavum.
Presented here are the deinotheriid dental and postcranial remains from the Late Miocene localities Pikermi and Halmyropotamos (Greece). The study and comparison of the available juvenile dental material from Pikermi with other relevant specimens from Europe showed that it belongs to the huge-sized Deinotherium proavum (= D. gigantissimum). Additionally, several postcranial specimens from Pikermi, as well as from Halmyropotamos, present deinotheriid features, which distinguish them from elephantoids, and permit their attribution also to D. proavum. This species is known from the Turolian and represents the terminal evolutionary stage of the European deinotheres. Its presence in Pikermi and Halmyropotamos is compatible with the middle Turolian (MN 12) age of the localities. Additionally, the taxonomy of European deinotheres is discussed, as well as the biostratigraphical and geographical distribution of D. proavum.
The data on an isolated upper tooth (P4) of Deinotherium sp. from the Late Miocene beds of the Maikop 1 locality (Maikop, Republic of Adygea) are reported. This is the first record of Deinotherium from the Upper Miocene of Russia. The tooth crown of P4 is similar in size to D. proavum Eichwald, 1831 (= D. gigantissimum Stefanescu, 1892).
This article summarises the history of research, the geological background and the stratigraphy of the Gratkorn locality (SE Austria). Since its discovery in 2005, 65 vertebrate taxa, comprising fishes, amphibians, reptiles, birds, and small and large mammals have been documented, as well as a variety of plant and invertebrate fossils. Due to its origin from a rapidly accumulated floodplain paleosol, time-averaging is low and the taphocoenose reflects well the original vertebrate community. The Gratkorn site is dated by integrated stratigra-phy, but independent from vertebrate biochronology, to about 12.2–12.0 Ma (late Middle Miocene). Thus, it probably yields the most diverse, systematically excavated vertebrate fauna of that age in Europe and is an extremely important benchmark for a vertebrate-based, continental biostratigraphy of the Central Paratethyan realm and beyond.
δ18OCO3, δ13C and 87Sr/86Sr measurements were performed on tooth enamel of several species to gain information on the diet and mobility of herbivorous large mammals from Gratkorn (Austria; late Sarmatian sensu stricto; 12.2–12.0 Ma). Except for the tragulid Dorcatherium naui, which was most likely frugivorous to a certain degree, the mean values and the total ranges of δ13C and δ18O of the large mammal taxa are typical for an exclusively C3 vegetation diet and point to predominantly browsing in mesic/woodland environments. Occupation of different ecological niches is indicated by variation in δ18O and δ13C among the taxa, and could be shown to be typical for the species by comparison with other Miocene localities from different areas and ages. The small moschid Micromeryx flourensianus might have occasionally fed on fruits. The cervid Euprox furcatus represents a typical subcanopy browsing taxon. The proboscidean Deinotherium levius vel giganteum browsed on canopy plants in the higher parts of an exclusively C3 vegetation as did the bovid Tethytragus sp.. Generally higher values for δ18O and δ13C of Lartetotherium sansaniense indicate feeding in a more open environment. Different ecological niches can be reconstructed for the two suids. While Listriodon splendens was a browsing taxon with a considerable input of fruits and maybe some grass in its diet, Parachleuastochoerus steinheimensis might have included roots. Distinct differences in 87Sr/86Sr values indicate that most of the larger mammals (Deinotherium levius vel giganteum, Parachleuastochoerus steinheimensis, Euprox furcatus, Lartetotherium sansaniense and to a minor degree maybe Listriodon splendens) were not permanent residents of the area around Gratkorn but rather inhabited a wider area, most likely including the Styrian Basin and the higher altitudes of the Eastern Alps’ palaeozoic basement.
One of the rare records of a rich ruminant fauna of late Middle Miocene age (Sarmatian sensu stricto; 12.2–12.0 Ma) was discovered at the Gratkorn locality (Styria, Austria). It comprises, besides Micromeryx flourensianus, ?Hispanomeryx sp., Euprox furcatus, Palaeomerycidae gen. et sp. indet., and Tethytragus sp., one of the oldest records of Dorcatherium naui. Gratkorn specimens of the latter species are in metric and morphologic accordance (e.g. selenodont teeth, bicuspid p2, non-fusion of malleolus lateralis and tibia) with type material from Eppelsheim (Germany) and conspecific material from Atzelsdorf (Austria), and do not show an intermediate morphology between Late Miocene Dorcatherium naui and Middle Miocene Dorcatherium crassum, thus enforcing the clear separation of the two species. It furthermore confirms the assignation of Dorcatherium naui to a selenodont lineage (together with Dorcatherium guntianum) distinct from a bunoselenodont lineage (including Dorcatherium crassum). The record of ?Hispanomeryx sp. is the first of this genus in Central Europe. While Tethytragus sp. could also be a new bovid representative for the Sarmatian of Central Europe, Micromeryx flourensianus and Euprox furcatus are well-known taxa in the Middle Miocene of Central Europe, but comprise their first records from Styria. Morphological data from this work in combination with isotopic measurements (δ18OCO3, δ13C; Aiglstorfer et al. 2014a, this issue) indicate a niche partitioning for the ruminants from Gratkorn with subcanopy browsing (Euprox furcatus), top canopy browsing (Tethytragus sp.) and even a certain amount of frugivory (Dorcatherium naui and Micromeryx flourensianus).
This article summarises the history of research, the geological background and the stratigraphy of the Gratkorn locality (SE Austria). Since its discovery in 2005, 65 vertebrate taxa, comprising fishes, amphibians, reptiles, birds, and small and large mammals have been documented, as well as a variety of plant and invertebrate fossils. Due to its origin from a rapidly accumulated floodplain paleosol, time-averaging is low and the taphocoenose reflects well the original vertebrate community. The Gratkorn site is dated by integrated stratigra-phy, but independent from vertebrate biochronology, to about 12.2–12.0 Ma (late Middle Miocene). Thus, it probably yields the most diverse, systematically excavated vertebrate fauna of that age in Europe and is an extremely important benchmark for a vertebrate-based, continental biostratigraphy of the Central Paratethyan realm and beyond.
At the Gratkorn locality (Styria, Austria), a highly diverse, late Middle Miocene (late Sarmatian sensu stricto; 12.2–12.0 Ma) faunal assemblage is preserved in a palaeosol. It represents the first systematically excavated and well-documented continental Sarmatian site in Central Europe. Taphonomical analysis of the 700 large mammal specimen excavated so far has led to the following conclusions: (1) the level of diagenetic alteration is low, as primary (aragonitic) mineralisation in gastropod shells is preserved and teeth and bones of large mammals in general show a relatively low total REE content; (2) the high degree of disarticulation and fragmentation in large mammal bones is induced by hunting, scavenging, trampling, and neotectonics; (3) there are no signs for fluviatile transportation due to the general preservation features of the bones (e.g. no record of abrasion) and the still roughly associated fragments of individual bones and skeletons; and (4) local accumulation of large mammal bones is the result of scavenging. The fossil assemblage is considered to form a more or less autochthonous taphocoenosis without any significant time averaging (or faunal mixing) in terms of geologic resolution (contemporaneously deposited).
The paper describes a mandible from Aksakovo near Varna, NE Bulgaria, referred to Deinotherium giganteum on the base of dental size, since morphology of p3 is not directly observable due to poor preservation. Also from Aksakovo, Prodeinotherium bavaricum is known with a molar recovered and published in the 1960s. The two deinotheriid specimens are the only fossil finds from Aksakovo so far and, while not associated, indicate a pre-Turolian, most probably middle Miocene age for the locality. Pre-Turolian land vertebrates are rare in Bulgaria, coming mostly from the northeast part of the country, mainly from the vicinities of Varna on the Black Sea coast. This is the first Deinotherium giganteum mandible from Bulgaria, with most of the deinotheriid finds from the country belonging to the Turolian species Deinotherium gigantissimum.
Here, a partial skeleton of Prodeinotherium bavaricum from Unterzolling (Southern Germany) is documented. The following elements are preserved and described for the first time: cervical vertebrae 1–2 and 5–7, the first thoracic vertebra, one lumbar vertebra, trapezium, metacarpals 1–5, tibia, calcaneus, endo- and mesocuneiform, cuboid, the fourth metatarsal, and some phalanges. Comparisons with the skeletons of P. bavaricum from Franzensbad (Czech Republic) and Deinotherium giganteum from Eserovo (Bulgaria) show osteological differences that are described and discussed.
Hundreds of non-cultural elephant bone sites have been studied in southern Africa, starting at or before the actual moments of death and continuing through bone burial or destruction. In some sites, dozens of elephants died en masse due to drought. These sites contain spirally fractured limb bones in proportions as high as 62% of counted limb elements. Many naturally broken tusk fragments are similar to specimens that have been interpreted as artifacts in fossil proboscidean collections. Trampling marks on bones closely mimic cut marks made by stone tools. As reported here, numerous attributes of non-cultural assemblages are virtually indistinguishable from attributes that archaeologists have believed to be created by human behaviour alone.
The fossil record richly illustrates the origin of morphological adaptation through time. However, our understanding of the selective forces responsible in a given case, and the role of behaviour in the process, is hindered by assumptions of synchrony between environmental change, behavioural innovation and morphological response. Here I show, from independent proxy data through a 20-million-year sequence of fossil proboscideans in East Africa, that changes in environment, diet and morphology are often significantly offset chronologically, allowing dissection of the roles of behaviour and different selective drivers. These findings point the way to hypothesis-driven testing of the interplay between habitat change, behaviour and morphological adaptation with the use of independent proxies in the fossil record.
In August of 1982 some bones of large Deinotherium were excavated in the Obuhovka sand pit (Fig. 1). That locality is situated on the high terrace of the Paleo-Don in 1.5 km to the west from Yanov khutor (Rostov region, Russia) on the right bank of the Grushevka River (the confluent of the Tuzlov River). The excavations were realized by collaborators of Azov local museum. Most part of bones was laid in anatomical sequence (Fig. 2). Skeleton was destroyed partly during its extracting. This find is important because of well preserved teeth were discovered jointly with the most part of the postcranial elements. There were some short publications about this finding (BAJGUSHEVA & TISHKOV, 1998; BAJGUSHEVA, 1998). Nowadays all remains were restored and the specimen is exhibiting in Azov local museum (Northeast Sea of Azov Region, Russia).
This paper presents one of the richest and most complete vertebrate faunas of the late Middle Miocene (~12Ma) of Central
Europe. Up to now, sixty-two vertebrate taxa, comprising all major groups (fishes, amphibians, reptiles, birds, mammals),
have been recorded. Based on sedimentological and palaeobiological evidences, this Fossillagerstätte is assumed to originate from a floodplain paleosol formed on top of a braided river sequence. The fauna points to a highly
structured, somewhat vegetated landscape with a wide array of habitats (e.g., fluvial channels, sporadically moist floodplains,
short-lived ponds, savannah-like open areas and screes). It was preserved due to a rapid drowning and the switch to a freshwater
lake environment. Palaeoclimatological data, derived from pedogenic features as well as from biota, indicate an overall semi-arid,
subtropical climate with distinct seasonality (mean annual precipitation 486±252mm, mean annual temperature ~15°C). This
underlines the late Middle/early Late Miocene dry-spell in Central Europe. From taphonomical point of view, the irregularly
distributed but roughly associated larger vertebrate remains refer to an in situ accumulation of the bone bed. Splintered
bones, gnawing marks as well as rhizoconcretions and root corrosion structures record some pre- and post-burial modification
of the taphocoenose. However, the findings of pellet remains argue for a very fast burial and thus to a low degree of time-averaging.
For this reason, the fossil fauna reflects the original vertebrate community rather well and is a cornerstone for the understanding
of late Middle Miocene terrestrial ecosystems in this region. Certainly, Gratkorn will be one of the key faunas for a high-resolution
continental biostratigraphy and the comprehension of Europe’s faunal interchanges near the Middle/Late Miocene transition.
KeywordsVertebrate fauna–Late Middle Miocene–Sarmatian s.str.–Central Paratethys–Styria/Austria–Fossillagerstätte
Mammalian fossils from the Eppelsheim Formation (Dinotheriensande) have been a benchmark for Neogene vertebrate palaeontology since 200 years. Worldwide famous sites like Eppelsheim serve as key localities for biochronologic, palaeobiologic, environmental, and mammal community studies. So far the formation is considered to be of early Late Miocene age (~9.5 Ma, Vallesian), representing the oldest sediments of the Rhine River. The stratigraphic unity of the formation and of its fossil content was disputed at times, but persists unresolved.
Here we investigate a new fossil sample from Sprendlingen, composed by over 300 mammalian specimens and silicified wood. The mammals comprise entirely Middle Miocene species, like cervids Dicrocerus elegans, Paradicrocerus elegantulus, and deinotheres Deinotherium bavaricum and D. levius. A stratigraphic evaluation of Miocene Central European deer and deinothere species proof the stratigraphic inhomogenity of the sample, and suggest late Middle Miocene (~12.5 Ma) reworking of early Middle Miocene (~15 Ma) sediments. This results agree with taxonomic and palaeoclimatic analysis of plant fossils from above and within the mammalian assemblage. Based on the new fossil sample and published data three biochronologic levels within the Dinotheriensand fauna can be differentiated, corresponding to early Middle Miocene (late Orleanian to early Astaracian), late Middle Miocene (late Astaracian), and early Late Miocene (Vallesian) ages.
This study documents complex faunal mixing of classical Dinotheriensand fauna, covering at least six million years, during a time of low subsidence in the Mainz Basin and shifts back the origination of the Rhine River by some five million years. Our results have severe implications for biostratigraphy and palaeobiology of the Middle to Late Miocene. They suggest that turnover events may be obliterated and challenge the proposed 'supersaturated' biodiversity, caused by Middle Miocene superstites, of Vallesian ecosystems in Central Europe.
An unusually severe drought in 1982 led to a temporary die-off of elephants at a natural water source in Hwange National Park, Zimbabwe. Compared to living populations, the age structure of the animals killed by drought is strongly biased toward 2-to 8-year-old animals. However, the fresh carcasses of these young elephants were commingled with weathered remains of adults that had died earlier, creating a mixed skeletal sample whose age structure was much closer to that of living populations. Observations of elephant bones that have accumulated due to natural mortality at water holes might provide analogs for paleoecological interpretations of fossil proboscidean assemblages.
An important area of biology involves investigating the origins in animals of traits that are thought of as uniquely human. One way that humans appear unique is in the importance they attach to the dead bodies of other humans, particularly those of their close kin, and the rituals that they have developed for burying them. In contrast, most animals appear to show only limited interest in the carcasses or associated remains of dead individuals of their own species. African elephants (Loxodonta africana) are unusual in that they not only give dramatic reactions to the dead bodies of other elephants, but are also reported to systematically investigate elephant bones and tusks that they encounter, and it has sometimes been suggested that they visit the bones of relatives. Here, we use systematic presentations of object arrays to demonstrate that African elephants show higher levels of interest in elephant skulls and ivory than in natural objects or the skulls of other large terrestrial mammals. However, they do not appear to specifically select the skulls of their own relatives for investigation so that visits to dead relatives probably result from a more general attraction to elephant remains.
Although low in diversity, megaherbivores (mammals weighting over 10(3) kg) and especially proboscideans have a powerful impact on the structure and dynamics of present-day ecosystems. During the Neogene (23 to 2.6 Ma) of Europe, the diversity and geographic distribution of these megaherbivores was much greater. Nonetheless, their role in past ecosystems is unclear. Nutrition is one of the main bonds between organisms and their environment. Therefore, the ecology of organisms can be inferred from their dietary habits. The present study is aimed at characterizing the feeding habits of diverse megaherbivores through dental microwear analyses. This method was applied on cheek teeth of three sympatric species of proboscideans from the middle/late Miocene of the Molasse Basin in Southern Germany: Gomphotherium subtapiroideum, Gomphotherium steinheimense, and Deinotherium giganteum. The microwear signatures are significantly different between these taxa, suggesting differences in feeding habits and ecological niches within a woodland environment. D. giganteum probably browsed on dicotyledonous foliages whereas the two species of gomphotheres were neither strict grazers nor strict browsers and instead probably fed on a large spectrum of vegetal resources. The differences of occlusal molar morphology between the two gomphotheres are supported by the dental microwear pattern. Indeed, G. subtapiroideum probably ingested more abrasive material than G. steinheimense. Thus, our results suggest that these proboscideans did not compete for food resources.
The Proboscidea, of which only two species of elephant survive today, were one of the great mammalian orders of the Cenozoic. Their success over evolutionary time is reflected by their morphological and taxonomic diversity, their nearly worldwide distribution on every continent except Australia and Antarctica, and their persistence through nearly fifty million years. Their great past ability to migrate and to adapt to changing climatic conditions and interspecific competition provides a unique laboratory for the testing of evolutionary theories and development of new concepts. This is the first complete treatise on the evolution and palaeoecology of this group for half a century. It reviews their classification and phylogeny, the early differentiation of proboscideans, the major adaptive radiations and their evolutionary patterns, and the origins and current status of extant elephant species. Written by leading international experts, this is a major study documenting the record of terrestrial biodiversity.
The paper presents to the international palaeontological community a description of the full skeleton of a deinothere found and excavated near the village of Ezerovo by D. Kovachev in 1965, and described, restored and mounted by the late Ivan Nikolov. The fossil remains are found in sediments of Maeotian age. Former excavations have yielded fossil bones of Trilophodon angustidens Cuvier, Hipparion microtaton Nicolov and parts of mandible of Mastodon sp. When the whole skeleton was excavated only a few bones were missing, and namely, the whole rear left thighbone, the right rear fibula, most of the ribs as well as some caudal vertebrae. They are restored on the basis of symmetrical analogues existing. The measurements on the bones found and the comparisons with other deinotheres described in the literature showed certain differences. Some of the differences relative to the skull have certainly a taxonomic value. Therefore, the deinothere described is referred to a new species, Deinotherium thraceiensis sp. n.
δ18OCO3, δ13C and 87Sr/86Sr measurements were performed on tooth enamel of several species to gain information on the diet and mobility of herbivorous large mammals from Gratkorn (Austria; late Sarmatian sensu stricto; 12.2–12.0 Ma). Except for the tragulid Dorcatherium naui, which was most likely frugivorous to a certain degree, the mean values and the total ranges of δ13C and δ18O of the large mammal taxa are typical for an exclusively C3 vegetation diet and point to predominantly browsing in mesic/woodland environments. Occupation of different ecological niches is indicated by variation in δ18O and δ13C among the taxa, and could be shown to be typical for the species by comparison with other Miocene localities from different areas and ages. The small moschid Micromeryx flourensianus might have occasionally fed on fruits. The cervid Euprox furcatus represents a typical subcanopy browsing taxon. The proboscidean Deinotherium levius vel giganteum browsed on canopy plants in the higher parts of an exclusively C3 vegetation as did the bovid Tethytragus sp.. Generally higher values for δ18O and δ13C of Lartetotherium sansaniense indicate feeding in a more open environment. Different ecological niches can be reconstructed for the two suids. While Listriodon splendens was a browsing taxon with a considerable input of fruits and maybe some grass in its diet, Parachleuastochoerus steinheimensis might have included roots. Distinct differences in 87Sr/86Sr values indicate that most of the larger mammals (Deinotherium levius vel giganteum, Parachleuastochoerus steinheimensis, Euprox furcatus, Lartetotherium sansaniense and to a minor degree maybe Listriodon splendens) were not permanent residents of the area around Gratkorn but rather inhabited a wider area, most likely including the Styrian Basin and the higher altitudes of the Eastern Alps’ palaeozoic basement.
Interpretation of the Neogene fluvial deposits of the Ur-Rhein—the Dinotheriensande—has a long history in which fossil mammals have played an ever present role. Most of the collections made prior to the 1930s consist of fossils from more than one biostratigraphic horizon, and this has obscured proper assessment of the age of the fossils from the deposits. As is usual in fluviatile deposits, there is evidence of reworking of many fossils, but most of them have probably not moved far from their original depositional locus. For many years, the Dinotheriensande deposits were correlated to the Early Pliocene (in the old sense of the term when “Pontian” was thought to be Early Pliocene), but for the past half century or so they have been correlated to the basal part of the late Miocene—the Vallesian. There has always been the suggestion that earlier deposits are present in the system, but because most of the sites have yielded remains of the equid, Hipparion (today Hippotherium), most authors have opted for a late Miocene age for the Eppelsheim Formation, which is the modern term for the Dinotheriensande. Even though Dorn-Dürkheim 1 is not strictly speaking part of the Dinotheriensande, it is geographically close to the deposits traditionally included in the deposits. This contribution examines the deinothere fossils from the Mainz Basin, and compares them to rich control samples excavated from horizons or sites which span a short time interval, including Langenau (MN 4, 62 teeth), Massenhausen (MN 8, 84 teeth) and Montredon (MN 10, 190 teeth). The results indicate that there are substantial quantities of fossils of the middle Miocene age in the Mainz Basin, especially well represented at Sprendlingen (MN 6, 113 teeth) and Gau-Weinheim (Wissberg) (MN 7/8 and MN 9–10, 84 teeth). The type area of the formation at Eppelsheim, which yielded 174 teeth, includes fossil specimens that correlate to MN 4–5, while most of the fossils correlate to MN 9, and some correlate to MN 10, whereas there are a few specimens suggesting an age equivalent to MN 11. Dorn-Dürkheim 1 (35 teeth) correlates to MN 11, although the presence of Anancus at the site pleads for a younger correlation (MN 12 or MN 13) (Gaziry 1997; Metz-Muller 2000). The paper probes the taxonomy of the Dorn-Dürkheim 1 deinotheres and other “huge” deinotheres from the Turolian of Europe, Asia and Africa, with the aim of determining the taxonomic identification of the material from Dorn-Dürkheim 1, which was previously interpreted as a form intermediate between D. giganteum and D. gigantissimum. It is concluded that it belongs to the latter species which is a junior synonym of two species names which have priority—D. proavum (Eichwald Nova Acta Phys-Med Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, 17:677–760, 1831) and D. indicum Falconer (Q J Geol Soc Lond 1:356–372, 1845). A final aim of this article is to describe and interpret huge deinothere dental remains from Iran which help fill the geographic gap that used to separate the Eastern European Deinotherium proavum (ex D. gigantissimum) from equally large Deinotherium indicum. This discovery suggests that the huge Indian and European deinotheres belong to a single widespread species.
The study of the original species descriptions showed that many European species were based on a small number of specimens and limited comparisons. Many type specimens mentioned in the original descripti-ons are no longer available. Currently only two European species are considered valid. The Early to Middle Miocene species Prodeinotherium bavaricum and the Middle to Late Miocene Deinotherium giganteum can be identified mainly by the cranial and postcranial characters. The interrelationships between the European, African and Asian Prodeinotherium and Deinotherium species are not known in detail. Zusammenfassung Viele europäische Deinotheriidae Arten wurden auf Grund von beschränktem Material oder ohne Vergleiche beschrieben. Viele von den beschriebenen Typusmaterialien existieren nicht mehr. Zwei europäische Arten sind zur Zeit bekannt. Die Früh-bis Mittelmiozäne Art Prodeinotherium bavaricum und die Mittel-bis Spätmiozäne Art Deinotherium giganteum sind unterschiedlich insbesondere in den crania-len und postcranialen Merkmalen. Die taxonomische Zusammenhänge zwischen den europäischen, afrika-nischen und asiatischen Prodeinotherium und Deinotherium Arten sind noch nicht bekannt.
The dental remains of the species Prodeinotherium bavaricum and Deinotherium giganteum can be identified by size and p3 morphology. In Lower Austria P. bavaricum is present in localities of Middle Miocene and early Late Miocene age. D. giganteum is present in the Late Miocene localities only. The species have a simi-lar distribution in all European countries. P. bavaricum is known from the biozones MN4 to MN9, whereas D. giganteum is present from MN7/8 to MN13. The supposed presence of both species in the early Late Miocene of Austria is either due to real coexistence of both species or uncertain stratigraphic determinations. Zusammenfassung Zahnfossilien von Prodeinotherium bavaricum und Deinotherium giganteum identifiziet man durch Größe und p3 Morphologie. In Niederösterreich ist P. bavaricum bekannt aus Mittel-bis früh Spätmiozänen Fund-stellen. D. giganteum ist bekannt nur aus den spätmiozänen Fundstellen. Die Arten haben eine ähnliche strati-graphische Verteilung europaweit. P. bavaricum ist bekannt aus den Biozonen MN4 bis MN9. D. giganteum ist bekannt von MN7/8 bis MN13. Das Vorhandensein von beiden Arten in frühen Spätmiozänen Fundstellen in Österreich deutet entweder auf tatsächliche Koexistenz, oder die stratigraphische Bestimmung der Fund-stellen ist nicht sicher.
Zusammenfassung Säugerfaunen stehen häufig nicht in einem erkennbaren regionalen Schichtzusammenhang. In solchen Fällen kann die Frage ihrer Gleichaltrigkeit vermittels der regionalen Verteilung möglichst zahlreicher Einzelfundpunkte geklärt werden. Unter diesem Gesichtspunkt werden hier zwei besondere Vorkommensgruppen untersucht: die Faunen der jungtertiären Oberen Süßwassermolasse im süddeutschen Alpenvorland und die weit verstreuten obereocänen Faunen Ostasiens. Im Molasse-Gebiet lassen sich die Faunenunterschiede auf Altersverschiedenheit und Wechsel des Klimas zurückführen, in Ostasien aber auf Klimaunterschiede bei Gleichaltrigkeit.
MARKOV G. N. 2008. The Turolian proboscideans (Mammalia) of Europe: preliminary observations. - Historia naturalis bulgarica, 19: 153-178. Abstract. The paper deals with six proboscidean species from the Turolian of Europe. Problems of their taxonomy, phylogeny, assumed geographic range and time span are discussed; an attempt is made to outline the most important localities and material for each of the six taxa. With the exception of Anancus, a later immigrant, the other five proboscideans seem to be part of the large- scale mammal migrations from West Asia into Europe around the Vallesian/Turolian transition.
The co-occurrence of DeinotheriumgiganteumKaup, 1829 and Prodeinotheriumbavaricum (Meyer, 1831) in Castelnau-Magnoac is demonstrated by the discovery of new remains and the revision of old collections. It confirms the coexistence of these two species, whose dental evolutionary tendencies are specified, in Astaracian (MN6 and MN7) of the Aquitaine basin.
As migratory animals, sustainable management of African elephant populations, both within and around protected areas, is a major challenge in the conservation policy of many African countries. We captured seven female elephants, representative members of family groups, in different parts of Tarangire National Park (TNP), Tanzania, and used GPS satellite radio-tracking (November 1997-June 2000) to monitor their space and habitat use and seasonal migrations throughout wet and dry seasons. Patterns of home range overlap revealed the existence of two Large clans that occupied the north-central and southern parts of TNP, respectively. At the end of the dry season, elephants from the southern clan migrated about 100 km southeast of the park boundary, those from the northern clan remained mostly inside the park, or used periodically wet-season core areas in the nearby Game Controlled Areas. No natural mortality occurred during the study, but two elephants were poached outside the park. Human disturbance also affected activity patterns, and elephants were Less active at day outside than inside the park. Home range size varied from 477 to 1078 km(2) for the northern elephants, and from 1630 to 5060 km(2) for the southern elephants. Migration routes were characterised by higher cover (open and closed forest) than core areas. Our results indicate that elephant management must be considered across park boundaries and that migration corridors must be protected against human disturbance and land cultivation. Society problems Linked to elephant conservation can be solved by creating alternative, sustainable, use of natural resources that enhance the livelihood of Local communities. (c) 2005 Deutsche Gesellschaft fur Saugetierkunde. Published by Elsevier GmbH. All rights reserved.
The fossil record of the extinct family Deinotheriidae is restricted to Neogene and early Quaternary faunas of the Old World. The family is assigned to the Order Proboscidae but the nature of its relationship to other proboscidean families is still uncertain. Deinotheres are rendered unique by the retention of a single pair of lower tusks and by the morphology of the cheekteeth.
The earliest recorded deinotheres exhibiteo morphological adaptations for a proboscis. In terms of length and prehensibility, the proboscises of the early deinotheres were more tapir-like than elephantine. Later representatives had larger and more powerful but not necessarily longer proboscises.
It is unlikely that the tusks were used for digging and the most probable functions were for stripping vegetation, for providing a source of purchase for the proboscis, and in providing a means of recognition of the individual.
The cheek teeth were deployed in two distinct functional batteries: an anterior crushing battery and a posterior shearing battery. The anterior molar acted initially as part of the shearing battery but, on becoming worn, subscribed to the crushing function of the anterior battery.
Deinotheres were “shearing browsers” and replaced barytheres in the African Paleogene faunas. Evolutionary trends in the known representatives afford some indication of the nature of pre-Miocene adaptations.
Limb-bone allometry was investigated for 19 species of proboscideans, spanning almost the entire phylogenetic spectrum. More archaic proboscideans (‘gompthotheres’) have substantially thicker long-bone diaphyses relative to length than elephantids, as has been suggested previously, but contrary to previous suggestions it could not be confirmed that Mammuthus had more massive long-bone diaphyses on average than extant Elephas and Loxodonta. When correcting for phylogeny, the circumference of the limb bones to their length in proboscideans as a group generally scale with negative allometry, becoming stouter with increased length, as would be expected from limb mechanics. Few slopes were, however, statistically significantly negatively allometric. Correcting for phylogeny produced better correlations than traditional regression analyses, in contrast to most other studies where the reverse is the case. Intraspecific analyses of extant Elephas and Loxodonta, in addition to Mammuthus primigenius, Mammut americanum, and Gomphotherium productum, also resulted in negatively allometric regression slopes, frequently conforming to the theory of elastic similarity, as could be expected from the columnar posture of proboscideans. At present the reasons for the more massive limbs of gomphotheres s.l. are not fully understood. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 423–436.
A partial skeleton of Prodeinotherium bavaricum on exhibit at the Museum of Natural History in Vienna is described. The skeleton originates from the Middle Miocene (MN5) locality Franzensbad, Czech Repub-lic and is one of the best-preserved Prodeinotherium skeletons available. The identification and taxonomic determination was carried out based on a comparative study with other European Deinotheres. The present report is the first comprehensive investigation on the variability of the postcranial morphology of Pro-deinotherium in Europe. The results support the contemporary Deinotheriidae taxonomy, which recognizes only one Early to Middle Miocene Prodeinotherium species in Europe. An expanded diagnosis of the genus is provided. Zusammenfassung Teile des Skeletts von Prodeinotherium bavaricum des Naturhistorischen Museums in Wien werden be-schrieben. Das Skelett stammt aus der mittelmiozänen (MN5) Fundstelle Franzensbad, Tschechische Repu-blik und gehört zu den besterhaltenen Prodeinotherium Skeletten der Welt. Die Identifizierung und taxono-mische Bestimmung wurde nach einer Vergleichsstudie durchgeführt. Diese Studie ist die erste umfangre-iche Studie über die Variabilität der postcranialen Morphologie des Prodeinotherium in Europa. Die Ergeb-nisse unterstützen der Ansicht der modernen Deinotheriidae-Taxonomie, die nur eine Früh-bis Mittel-miozäne Prodeinotherium Art in Europa kennt. Als Ergebnis wird eine erweiterte Gattungsdiagnose erstellt.
MarkoV G. N. 2008. Fossil proboscideans (Mammalia) from the vicinities of Varna: a rare indication of middle Miocene vertebrate fauna in Bulgaria – Historia naturalis bulgarica, 19: 137-152. Abstract. Proboscideans from the area of Varna (NE Bulgaria) are discussed, with emphasis on two finds from Galata: an elephantoid molar mistakenly identified as Tetralophodon longirostris in earlier Bulgarian literature but actually belonging to an amebelodontid, and a previously unpublished premolar of Deinotherium giganteum. The proboscideans from the region of Varna are a rare example of pre-Turolian vertebrates from Bulgaria and might indicate middle Miocene fossiliferous outcrops in the area.
Mass estimates for a number of fossil proboscideans were computed using regression analyses on appendicular bones to body mass, for seven specimens of modern elephants, for which body masses had been recorded prior to death. The marked differences in physical proportions between extant Loxodonta and Elephas, implying substantial differences in body mass at any given shoulder height, were not present in their long bone parameters. Length and least circumferences proved to be the best parameters for prediction of body mass. Some extinct proboscideans, notably certain Mammuthus and Deinotherium, were much larger than extant elephants. Both the basal and the field metabolic rates of extant elephants are lower than predicted for a hypothetical mammal, in accordance with their body size and subsistence on low-quality foods. The feeding quantities often ascribed to extant wild elephants are exaggerated, and would in fact have sufficed to nourish much larger species. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 523–549.
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