- Email: [email protected]
Studying Proboscideans: knowledge, problems, and perspectives
ARTICLE IN PRESS
Quaternary International 126–128 (2005) 1–3
Preface
Studying Proboscideans: knowledge, problems, and perspectives In the course of their long history, covering approximately 55 million years, Proboscideans exhibited complex and in some ways not yet clarified evolutionary patterns. It is enough to say that in the noticeable and ponderous opera by Osborn (1933–1942), the classification system was based on the concept of adaptive radiation and more than 350 species have been recognised. The more our knowledge increases, the more windows are opened on the past and the present Proboscideans and elephant life, the more sophisticated methods of analysis we employ, more discoveries arise and more zones still remain in shadow. As generally accepted today, at least three main radiation phases have been recognised (Shoshani and Tassy, 1996) that have given rise to phenomena of convergence, to extremely specialized forms, and to taxa whose phylogenetic relationships are still debated. Moreover, Proboscideans have inhabited the most diverse environments, sometimes extreme, from semiaquatic environments, to deserts, to rain forests, to the savannah, to the tundra, often attaining a very wide geographical distribution. The elephants, the last terrestrial giant. As A.S. Romer wrote in his Vertebrate Paleontology in 1966, ‘‘one of the most spectacular stories in mammalian evolution is that of the order Proboscidea’’. Moreover for decades, Proboscideans in general have been recorded as a ‘‘model’’ of evolution. In particular, it is generally believed that some elephant lineages represent a clear example of anagenetic evolution. Moreover, rivers of ink have been spent debating Palaeolithic human and proboscideans’ relationships. Accordingly, we believed that a conference focusing on topics of interest to both palaeontologists and archaeologists might find a particularly vigorous response in the international scientific community. The city of Rome itself, and its environs, characterized by particularly significant fossiliferous localities, looked like an appropriate site to hold the 1st International Congress, ‘‘The World of Elephants’’. An extended abstract volume containing all contributions has been distributed at the meeting and subsequently mailed upon request to all research institutions (Cavarretta et al., 2001). Furthermore, we have prepared this volume as a main result of the debate raised from this conference,
collecting selected contributions discussed in our international congress, together with invited papers. Among the themes proposed for discussion at the Congress, we have chosen those that were met with a particularly vigorous response in the international scientific community. Our aim is to offer a picture about the ‘‘state-of-art’’ of our knowledge, to concentrate our attention on the unsolved problems, to suggest topics for future researches, and to provide a sound foundation for further discussions. Accordingly, the most recent hypotheses and findings on the origin and classification of Proboscideans have been highlighted in the Shoshani and Tassy paper. The authors focused their attention on ‘‘aquatic ancestry’’ of Proboscidea, interordinal relationships within Placentalia, Proboscidean taxonomy, anatomy and physiology and some ecological considerations. They then reassessed Proboscidea, classifying them in 42 genera and 10 families. Where Proboscidea from South America and Asia are concerned, Prado et al. provided new evidence to understand pattern and process of evolution of Gomphotheres as well as their palaeobiogeography, dispersal and palaeoecology. New, interesting data on Asian stegodontids provided by Saegusa et al. suggest stegodons might have originated in Asia. Moreover, even if Mammoth evolution in Eurasia represents one of the best-studied examples of evolutionary pattern and process in the terrestrial fossil record, as demonstrated by the Lister et al. paper, the gradual transformation of chronospecies did not occur in Europe but a complex evolutionary process took place, that included geographical variation across the whole of northern Eurasia and some introgression phenomena. The African Elephas lineage seems to mirror a similar situation because this lineage has been proposed as an example of anagenetic change, and chronosubspecies within it have often been used as biochronological markers. Todd’s paper suggests that Elephas recki is not a single species and highlights that a revision of the African Elephas lineage, in conjunction with reanalysis of the other elephantid lineages, will have important implications for elephantid diversity, evolution and biochronology in the African Plio–Pleistocene.
1040-6182/$ - see front matter r 2004 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2004.04.009
ARTICLE IN PRESS 2
Preface / Quaternary International 126–128 (2005) 1–3
Accordingly, it clearly emerges that biochronological deductions based on gradual transformation of molar morphology have to be used with great caution. Indeed, the biochronogical and paleoecological significance of Proboscideans has been highlighted in papers by Agenbroad, Vislobokova, and Palombo and Ferretti. Agenbroad concerns himself with temporal and geographic distribution for the genus Mammuthus in North America. Vislobokova concerns herself with Pliocene Proboscideans in the territory of the former Soviet Union; and Palombo and Ferretti concern themselves with elephant fossil record from Italy. In the later paper also the problems connected with the processes of reduction in size of elephants in insular environments is synthetically considered. Paleocological definition of the so-called ‘‘mammoth steppe’’ was the subject of Velichko and Zelikson’s paper. Taking into account that the identification of mammal palaeodiet can be a useful tool to reconstruct palaeoenvironment, Palombo et al. presented the results of an integrated methodological approach based on microwear scars, isotope analysis and X-ray diffraction for the late Middle Pleistocene Elephas antiquus samples from Italy. Paleoenvironmental conditions have also played a noticeable role in elephant extinctions. Nevertheless, Mammuthus primigenius and Palaeoloxodon antiquus became extinct in the last glacial–interglacial cycle, but they had very different ecologies, times of extinction and ‘last stands’ in different regions and, as pointed out by Stuart, environmental change alone appears insufficient to account for extinctions. However, the possible role of human hunters is also still unclear. Human–elephants interaction was one of the most important themes debated during the ‘‘The World of Elephants’’ congress. With archaeological evidence, this interaction can be proved for a span of time covering over two million years. The use of flakes of elephant bone, and in some cases of ivory, already appears in the Lower Palaeolithic, but is also well attested during the Upper Palaeolithic, when elephant bones were also used as building material. The paper by Gaudzinski et al. provides interesting information on the use of Proboscidean remains in every-day Palaeolithic life as an attempt to illuminate some aspects of the relationship between Proboscideans and humans from an archaeological perspective. A particular aspect was offered by Iakovleva and Djindjian’s paper, reporting preliminary results of the new excavations realised during the last ten years on the Gontsy site (Ukraine) that were focused on the different dwelling structures and the various activity around the dwellings. According to Svoboda et al., during Gravettian time the lowlands of Lower Austria–Moravia–South Poland form an important natural corridor in Central Europe,
allowing migrations of both humans and mammals. Mammoth bone accumulations from these areas can be interpreted as butchery places on the death locations and as butchery places on death/hunting sites. On the other hand, if the Lower Palaeolithic is concerned, new data regarding the hunting versus scavenging debate come from the Villa et al. paper devoted to the Spanish Middle Pleistocene Ambrona site: The site is a natural landscape regularly visited by hominids who transported some artifacts from nonlocal raw material sources, and had an organized approach to meat acquisition, but did not actively hunt. At about the same time, in a very different geographic area (Guizhou, South China), similar taphonomical perspectives came from the study of Stegodon orientalis remains from the Middle Pleistocene archaeological site of Panxian Dadong. Schepartz et al. prove that Proboscideans faunal assemblage at Dadong is probably a result of both hominid activities (either hunting or scavenging) and the natural accumulation of young stegodonts from accidental deaths. It is clear that the ongoing researches have continuously improved our knowledge and some preconceived ideas, even if deeply rooted, have now changed. In short, the Editors hope that this volume will be useful as a basis for more investigations. We hope also to solicit multidisciplinary approaches to study Proboscidea origin, phylogeny, physiology and behaviour that might explain still unsolved enigmas. This volume could also be interesting for colleagues who are curious about the history of the last terrestrial giants. Much has been achieved, but much still remains to be done. It is therefore to be hoped that a series of meetings may help in the future to further elucidate the close link between climate, environment, evolution of the taxa and structure of the palaeocommunities, and the development of the interrelation between man and elephant through the various phases of prehistory. For our colleagues and scientists interested in these subjects the date for the 2nd International Congress ‘‘The World of Elephants’’ is September 2005 in ‘‘Hot Springs’’, the South Dakota (USA) mythic Mammoth locality.
References Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), 2001. La terra degli Elefanti—The World of Elephants. Proceedings of the First International Congress. CNR, Roma, 739pp (pdf format can be downloaded at http://www.cq.rm.cnr.it/elephants2001/atti it. htm. Osborn, H.F., 1942. Proboscidea 2: Stegodontoidea, Elephantoidea, The American Museum Natural History. New York, 1676pp. Romer, A.S., 1966. Vertebrate Paleontology, 3rd edn, University of Chicago Press, 468pp. Shoshani, J., Tassy, P. (Eds.), 1996. The Proboscidea. Oxford University Press, 472pp.
ARTICLE IN PRESS Preface / Quaternary International 126–128 (2005) 1–3
M.R. Palombo Dipartimento di Scienze della Terra, CNR, Istituto di Geologia Ambientale e Geoingegneria Universita" degli Studi di Roma ‘‘La Sapienza’’, P. le Aldo Moro 5, I–00185 Roma, Italy E-mail address: [email protected] M. Mussi Dipartimento Scienze dell’Antichita" Universita" degli Studi di Roma ‘‘La Sapienza’’, P. le Aldo Moro 5, I–00185 Roma, Italy E-mail address: [email protected]
3
P. Gioia Soprintendenza Comunale ai Beni Culturali Piazza Lovatelli 35, I–00186 Roma, Italy E-mail address: [email protected] G. Cavarretta CNR, Istituto di Geologia Ambientale e Geoingegneria Via Bolognola 7, Roma, Italy E-mail address: [email protected]
Quaternary International 126–128 (2005) 1–3
Preface
Studying Proboscideans: knowledge, problems, and perspectives In the course of their long history, covering approximately 55 million years, Proboscideans exhibited complex and in some ways not yet clarified evolutionary patterns. It is enough to say that in the noticeable and ponderous opera by Osborn (1933–1942), the classification system was based on the concept of adaptive radiation and more than 350 species have been recognised. The more our knowledge increases, the more windows are opened on the past and the present Proboscideans and elephant life, the more sophisticated methods of analysis we employ, more discoveries arise and more zones still remain in shadow. As generally accepted today, at least three main radiation phases have been recognised (Shoshani and Tassy, 1996) that have given rise to phenomena of convergence, to extremely specialized forms, and to taxa whose phylogenetic relationships are still debated. Moreover, Proboscideans have inhabited the most diverse environments, sometimes extreme, from semiaquatic environments, to deserts, to rain forests, to the savannah, to the tundra, often attaining a very wide geographical distribution. The elephants, the last terrestrial giant. As A.S. Romer wrote in his Vertebrate Paleontology in 1966, ‘‘one of the most spectacular stories in mammalian evolution is that of the order Proboscidea’’. Moreover for decades, Proboscideans in general have been recorded as a ‘‘model’’ of evolution. In particular, it is generally believed that some elephant lineages represent a clear example of anagenetic evolution. Moreover, rivers of ink have been spent debating Palaeolithic human and proboscideans’ relationships. Accordingly, we believed that a conference focusing on topics of interest to both palaeontologists and archaeologists might find a particularly vigorous response in the international scientific community. The city of Rome itself, and its environs, characterized by particularly significant fossiliferous localities, looked like an appropriate site to hold the 1st International Congress, ‘‘The World of Elephants’’. An extended abstract volume containing all contributions has been distributed at the meeting and subsequently mailed upon request to all research institutions (Cavarretta et al., 2001). Furthermore, we have prepared this volume as a main result of the debate raised from this conference,
collecting selected contributions discussed in our international congress, together with invited papers. Among the themes proposed for discussion at the Congress, we have chosen those that were met with a particularly vigorous response in the international scientific community. Our aim is to offer a picture about the ‘‘state-of-art’’ of our knowledge, to concentrate our attention on the unsolved problems, to suggest topics for future researches, and to provide a sound foundation for further discussions. Accordingly, the most recent hypotheses and findings on the origin and classification of Proboscideans have been highlighted in the Shoshani and Tassy paper. The authors focused their attention on ‘‘aquatic ancestry’’ of Proboscidea, interordinal relationships within Placentalia, Proboscidean taxonomy, anatomy and physiology and some ecological considerations. They then reassessed Proboscidea, classifying them in 42 genera and 10 families. Where Proboscidea from South America and Asia are concerned, Prado et al. provided new evidence to understand pattern and process of evolution of Gomphotheres as well as their palaeobiogeography, dispersal and palaeoecology. New, interesting data on Asian stegodontids provided by Saegusa et al. suggest stegodons might have originated in Asia. Moreover, even if Mammoth evolution in Eurasia represents one of the best-studied examples of evolutionary pattern and process in the terrestrial fossil record, as demonstrated by the Lister et al. paper, the gradual transformation of chronospecies did not occur in Europe but a complex evolutionary process took place, that included geographical variation across the whole of northern Eurasia and some introgression phenomena. The African Elephas lineage seems to mirror a similar situation because this lineage has been proposed as an example of anagenetic change, and chronosubspecies within it have often been used as biochronological markers. Todd’s paper suggests that Elephas recki is not a single species and highlights that a revision of the African Elephas lineage, in conjunction with reanalysis of the other elephantid lineages, will have important implications for elephantid diversity, evolution and biochronology in the African Plio–Pleistocene.
1040-6182/$ - see front matter r 2004 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2004.04.009
ARTICLE IN PRESS 2
Preface / Quaternary International 126–128 (2005) 1–3
Accordingly, it clearly emerges that biochronological deductions based on gradual transformation of molar morphology have to be used with great caution. Indeed, the biochronogical and paleoecological significance of Proboscideans has been highlighted in papers by Agenbroad, Vislobokova, and Palombo and Ferretti. Agenbroad concerns himself with temporal and geographic distribution for the genus Mammuthus in North America. Vislobokova concerns herself with Pliocene Proboscideans in the territory of the former Soviet Union; and Palombo and Ferretti concern themselves with elephant fossil record from Italy. In the later paper also the problems connected with the processes of reduction in size of elephants in insular environments is synthetically considered. Paleocological definition of the so-called ‘‘mammoth steppe’’ was the subject of Velichko and Zelikson’s paper. Taking into account that the identification of mammal palaeodiet can be a useful tool to reconstruct palaeoenvironment, Palombo et al. presented the results of an integrated methodological approach based on microwear scars, isotope analysis and X-ray diffraction for the late Middle Pleistocene Elephas antiquus samples from Italy. Paleoenvironmental conditions have also played a noticeable role in elephant extinctions. Nevertheless, Mammuthus primigenius and Palaeoloxodon antiquus became extinct in the last glacial–interglacial cycle, but they had very different ecologies, times of extinction and ‘last stands’ in different regions and, as pointed out by Stuart, environmental change alone appears insufficient to account for extinctions. However, the possible role of human hunters is also still unclear. Human–elephants interaction was one of the most important themes debated during the ‘‘The World of Elephants’’ congress. With archaeological evidence, this interaction can be proved for a span of time covering over two million years. The use of flakes of elephant bone, and in some cases of ivory, already appears in the Lower Palaeolithic, but is also well attested during the Upper Palaeolithic, when elephant bones were also used as building material. The paper by Gaudzinski et al. provides interesting information on the use of Proboscidean remains in every-day Palaeolithic life as an attempt to illuminate some aspects of the relationship between Proboscideans and humans from an archaeological perspective. A particular aspect was offered by Iakovleva and Djindjian’s paper, reporting preliminary results of the new excavations realised during the last ten years on the Gontsy site (Ukraine) that were focused on the different dwelling structures and the various activity around the dwellings. According to Svoboda et al., during Gravettian time the lowlands of Lower Austria–Moravia–South Poland form an important natural corridor in Central Europe,
allowing migrations of both humans and mammals. Mammoth bone accumulations from these areas can be interpreted as butchery places on the death locations and as butchery places on death/hunting sites. On the other hand, if the Lower Palaeolithic is concerned, new data regarding the hunting versus scavenging debate come from the Villa et al. paper devoted to the Spanish Middle Pleistocene Ambrona site: The site is a natural landscape regularly visited by hominids who transported some artifacts from nonlocal raw material sources, and had an organized approach to meat acquisition, but did not actively hunt. At about the same time, in a very different geographic area (Guizhou, South China), similar taphonomical perspectives came from the study of Stegodon orientalis remains from the Middle Pleistocene archaeological site of Panxian Dadong. Schepartz et al. prove that Proboscideans faunal assemblage at Dadong is probably a result of both hominid activities (either hunting or scavenging) and the natural accumulation of young stegodonts from accidental deaths. It is clear that the ongoing researches have continuously improved our knowledge and some preconceived ideas, even if deeply rooted, have now changed. In short, the Editors hope that this volume will be useful as a basis for more investigations. We hope also to solicit multidisciplinary approaches to study Proboscidea origin, phylogeny, physiology and behaviour that might explain still unsolved enigmas. This volume could also be interesting for colleagues who are curious about the history of the last terrestrial giants. Much has been achieved, but much still remains to be done. It is therefore to be hoped that a series of meetings may help in the future to further elucidate the close link between climate, environment, evolution of the taxa and structure of the palaeocommunities, and the development of the interrelation between man and elephant through the various phases of prehistory. For our colleagues and scientists interested in these subjects the date for the 2nd International Congress ‘‘The World of Elephants’’ is September 2005 in ‘‘Hot Springs’’, the South Dakota (USA) mythic Mammoth locality.
References Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), 2001. La terra degli Elefanti—The World of Elephants. Proceedings of the First International Congress. CNR, Roma, 739pp (pdf format can be downloaded at http://www.cq.rm.cnr.it/elephants2001/atti it. htm. Osborn, H.F., 1942. Proboscidea 2: Stegodontoidea, Elephantoidea, The American Museum Natural History. New York, 1676pp. Romer, A.S., 1966. Vertebrate Paleontology, 3rd edn, University of Chicago Press, 468pp. Shoshani, J., Tassy, P. (Eds.), 1996. The Proboscidea. Oxford University Press, 472pp.
ARTICLE IN PRESS Preface / Quaternary International 126–128 (2005) 1–3
M.R. Palombo Dipartimento di Scienze della Terra, CNR, Istituto di Geologia Ambientale e Geoingegneria Universita" degli Studi di Roma ‘‘La Sapienza’’, P. le Aldo Moro 5, I–00185 Roma, Italy E-mail address: [email protected] M. Mussi Dipartimento Scienze dell’Antichita" Universita" degli Studi di Roma ‘‘La Sapienza’’, P. le Aldo Moro 5, I–00185 Roma, Italy E-mail address: [email protected]
3
P. Gioia Soprintendenza Comunale ai Beni Culturali Piazza Lovatelli 35, I–00186 Roma, Italy E-mail address: [email protected] G. Cavarretta CNR, Istituto di Geologia Ambientale e Geoingegneria Via Bolognola 7, Roma, Italy E-mail address: [email protected]