The dogs of Roman Vindolanda, Part III: Quantifying juvenilization and pleiotropic effects of miniaturization
Schlagworte:
CANIS FAMILIARIS, DOMESTIC DOG, DWARF DOG, MINIATURE DOG, NEOMORPHY, PAEDOMORPHY, JUVENILIZATION, ROMANO–BRITISH, VINDOLANDAAbstract
In this report we focus on harriers, small dogs of gracile build widely represented in Roman artwork, yet not previously documented from skull or skeletal material. With them we compare remains of other types of small dogs from Vindolanda and other Roman-era sites. Accurate characterization of small dogs requires differentiation between paedomorphy and juve- nilization in skull ontogeny. We present a new method for quantifying the degree of juveniliza- tion in dog skulls which is based on understanding the differential development of endochondral and dermal centers of ossification. We enumerate functional and pleiotropic effects of skull and dental morphology and development that are characteristic of miniature dogs but rare in dwarf dogs and in large, normally-proportioned dogs. By comparing a wide array of modern dogs and wolves, we determine the degree to which small gracile dogs of the Roman era differ from prim- itive, non-juvenilized forms. We employ ratio analysis of dog skull shape utilizing parameters previously indicated by principal component analysis (PCA) as being of high diagnostic value. Although some small and medium-sized domestic dogs have highly juvenilized skulls, equally small skulls from Vindolanda are no more juvenilized than wolves or Dingoes. While small, gracile dogs existed as early as the late Iron Age, we conclude that juvenilization in dogs arose after the Roman era.
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Literaturhinweise
Ashdown, R. & Evans, C. 1981: Animal bones. Skeleton Green: A Late Iron Age and Romano–British site. Britannia Monograph 2: 205–235.
Azúa, R.V. 2000: Prehispanic dog types in Middle America. In: Crockford, S.J. (ed.): Dogs Through Time: an Archaeological Perspective: 193–204. Proceedings of the 1st ICAZ Symposium on the history of the domestic dog. B.A.R. (International Series) 889. Oxford.
Bartosiewicz, L. 2000: Metric variability in Roman period dogs in Pannonia Province and the Barbaricum (Hungary). In: Crockford, S.J. (ed.): Dogs Through Time: an Archaeological Perspective: 181-189. Proceedings of the 1st ICAZ Symposium on the history of the domestic dog. B.A.R. (International Series) 889. Oxford.
Baxter, I.L. 2002: A dwarf hound skeleton from a Romano–British grave at York Road, Leicester, England, U.K., with a discussion of other Roman small dog types and speculation regarding their respective aetiologies. In: Snyder, L.M. & Moore, E.A. (eds.): Dogs and People in Social, Working, Economic or Symbolic Interaction: 12–23. Oxbow Books, Oxford.
— 2010 a: Small Roman dogs. Alexandria Archive 901: 1–9. Available at http://alexandriaarchive.org/bonecommons/items/show/901.
— 2010 b: A re-examination of the Thistleton canids.
Alexandria Archive 885: 1–11. Available at: http://alexandriaarchive.org/bonecommons/items/show/885.
Baxter, I.L. & Nussbaumer, M. 2009: Evidence of morphometric variation in an Iron Age dog cranium from Trumpington, Cambridgeshire, U.K. Archaeofauna 18: 69–78.
Bennett, D.K. 1980: Stripes do not a zebra make, Part I: A cladistic analysis of Equus. Systematic Zoology 29: 272–287.
— 2005: Bone from the Severan ditch, Area A, 2004. In: Birley, A. & Blake, J. (eds): The Vindolanda Excavations, 2003–2004. The Vindolanda Trust, Hexham, Northumberland: 115–157.
— 2007: Ongoing mammal bone research at Vindolanda: interim report on the 2005–2006 collections. In: Birley, A. & Blake, J. (eds.): The Vindolanda excavations 2005–2006. The Vindolanda Trust, Hexham, Northumberland: 200-204.
Bennett, D.K. & Timm, R.M. 2013: Microfaunal remains from the east granary, Roman Vindolanda: evidence for human–animal interactions. In: Birley, A. & Blake, J. (eds): The Vindolanda Granary Excavations. Roman Army Museum Publications, Brampton, Greenhead: 72–98.
— 2016: The dogs of Roman Vindolanda, Part II: Time-stratigraphic occurrence, ethnographic comparisons, and biotype reconstruction. Archaeofauna 26: 93–112.
Bennett, D.; Campbell, G. & Timm, R.M. 2016: The dogs of Roman Vindolanda, Part I: Morphometric techniques useful in differentiating domestic and wild canids. Archaeofauna 26: 71–92.
Birley, A. 2003: Vindolanda Research Report, 2003: The Excavations of 2002–2002, Volume I. The Vindolanda Trust, Bardon Mill. Hexham.
Blake, J. 2014: Vindolanda Research: the Excavations of 2007–2012 in the Vicus or Extramural Settlement (‘Area B’). Roman Army Museum Publications, Brampton. Greenhead.
Churcher, C.S. 1993: Dogs from Ein Tirghi cemetery, Balat, Dakleh Oasis, Western Desert of Egypt. In: Clason, A.; Payne, S. & Uerpmann, H.-P. (eds.): Skeletons in Her Cupboard: a Festschrift for Juliet Clutton-Brock: 39–59. Oxbow Monographs 34. Oxford.
Clark, K.M. 1995: The later prehistoric and protohistoric dog: the emergence of canine diversity. Archaeozoologia 2: 9–32.
— 2006: The dog assemblage. In: Fulford, M.; Clarke,
A. & Eckhardt, H. (eds.): Life and Labour in Late Roman Silchester: Excavations in Insula IX Since 1997. Britannia Monograph Series no. 22: 189–195.
— 2012: A review of the Romano–British dog. In: Fulford, M.; Clarke, A. & Eckardt, H. (eds.): Silchester and the Study of Romano–British Urbanism. Journal of Roman Archaeology, Supp. Series 90: 1651– 1684.
— In press: Tiny dog from Iron-Age Silchester. In: Fulford, M.; Clarke, A.; Durham, E. & Pankhurst, N. (eds.): Late Iron Age Calleva: the pre-conquest occupation at Silchester Insula IX, London, Society for the Promotion of Roman Studies, Britannia Monograph.
Corbett, L. 1995: The Dingo in Australia and Asia. University of New South Wales Press, Sydney.
Crockford, S.J. 2000: Dog evolution: a role for thyroid hormone physiology in domestication changes. In: Crockford, S.J. (ed.): Dogs Through Time: an Archaeological Perspective: 11–20. Proceedings of the 1stI.C.A.Z. Symposium on the history of the domestic dog. B.A.R. (International Series) 889. Oxford.
Davis, S.J.M. 1997: Animal bones from the Roman site at Redlands Farm, Stanwick, Northamptonshire, 1990 excavation. Ancient Monuments Laboratory Reports Series 106/97.
De Grossi-Mazzorin, J. & Tagliacozzo, A. 2000: Morphological and osteological changes in the dog from the Neolithic to the Roman period in Italy. In: Crockford, S.J. (ed.): Dogs Through Time: an Archaeological Perspective: 141–161. Proceedings of the 1st I.C.A.Z. Symposium on the history of the domestic dog. B.A.R. (International Series) 889. Oxford.
Drake, A.G. 2004: Evolution and development of the skull morphology of canids: an investigation of morphological integration and heterochrony. Ph.D. dissertation, University Massachusetts, Amherst.
— 2011: Dispelling dog dogma: an investigation of heterochrony in dogs using 3D geometric morphometric analysis of skull shape. Evolution and Development 13: 204–213.
Drake, A.G. & Klingenberg, C.P. 2008: The pace of morphological change: historical transformation of skull shape in St. Bernard dogs. Proceedings of the Royal Society B: Biological Sciences 275: 71–76.
Farello, P. 1995: I cani tardoantichi rinvenuti in un condoto fognario de Classe (RA). Estratto da Padusa Quaderni 1: 295–308.
Fillios, M.A. & Taçon, P.S.C. 2016: Who let the dogs in? A review of recent genetic evidence for the introduction of the dingo to Australia and implications for the movement of people. Journal of Archaeological Science: Reports http://dx.doi.org/10.1016/j.jasrep.2016.03.001, pp. 1–9.
Goodrich, E.S. 1958: Studies on the Structure & Development of Vertebrates. Vol. I: 1–485. Dover Publications, New York.
Grant, A. 1984: Animal husbandry, in Danebury: An Iron Age Hillfort in Hampshire, Vol. 2: The Excavations 1969–1978. Council for British Archaeology (CAB) Research Reports 52: 533–563.
Hall, S. 2000: Indigenous domestic dogs of southern Africa: an introduction. In: Blench, R.M. & MacDonald,
K.C. (eds.): The Origins and Development of African Livestock: Archaeology, Genetics, Linguistics and Ethnography: 302-311. University College of London Press, London.
Hammer, Ø.; Harper, D.A.T. & Ryan, P.D. 2001: PAST: Paleontological statistics software package for education and data analysis. Palaeontologica Electronica 4: 1–9. Available at http://palaeo-electronica. org/2001_1/past/issue1_01.htm.
Harcourt, R.A. 1974: The dog in prehistoric and early historic Britain. Journal of Archaeological Science 1: 151–175.
Huber, W. 1974: Biometrische analyse der Brachycephalie beim Haushund. L’Année Biologique 13(3–4): 135–141.
Kohler-Matznick, J.; Brisbin, I.L.jr.; Feinstein, M. & Bulmer, S. 2003: An updated description of the New Guinea singing dog (Canis hallstromi, Troughton 1957). Journal of Zoology (London) 261: 109–118.
Kruska, D. 1988: Mammalian domestication and its effect on brain structure and behavior. In: Jerison, H.J. & Jerison, I. (eds.): Intelligence and Evolutionary Biology, NATO ASI Series vol. G17: 211–250.
Larson, G.; Karlsson, E.K.; Perri, A.; Webster, M.T.; Ho, S.Y.W.; Peters, J.; Stahl, P.W.; Piper, P.J.; Lingaas, F.; Fredholm, M.; Comstock, K.E.; Modiano, J.F.; Schelling, C.; Agoulnik, A.L.; Leegwater, P.A.; Dobney, K.; Vigne, J-D.; Vila, C.; Andersson, L. & Lindblad-Toh, K. 2012: Rethinking dog domestication by integrating genetics, archeology, and biogeography. Proceedings of the National Academy of Sciences, USA 109: 1–6.
Lawrence, B. & Reed, C.A. 1983: The dogs of Jarmo. In: Braidwood, L.S.; Braidwood, R.J.; Howe, B.; Reed,
C.A. & Watson, P. (eds.): Prehistoric Archaeology along the Zagros Flanks: 485-494. University of Chicago Oriental Institute Publication 1.
Lüps, P. 1974: Biometrische Untersuchungen an der Schadelbasis des Haushundes. Zoologischer Anzeiger 192: 383–413.
Luttschwager, J. 1965: Ein zwerghund aus römischen Graberfeld and der Berliner Strasse zu Heidelberg-Neuenheim. Zeitschrift für Saügeteirkunde 30: 24–30.
Mackinnon, M. & Belanger, K. 2002: In sickness and health: care for an arthritic Maltese dog from the Roman cemetery of Yasmina, Carthage, Tunisia. In: Snyder, L.M. & Moore, E.A. (eds.): Dogs and People in Social, Working, Economic or Symbolic Interaction: 38–43. Proceedings of the 9th ICAZ Conference, Durham. Oxbow Books 85. Oxford.
Maltby, M. 1987: Animal bones from the excavations at Owslebury, Hampshire: an Iron Age and early Romano–British settlement. English Heritage AML Report, 6/87. London.
Nussbaumer, M. 1978: Biometrischer vergleich der Topogenesemuster an der Schädelbasis kleiner und mittelgrosser Hunde. Sonderdruck aus Zeitschrift für Tierzüchtung und Züchtungsbiologie 95: 1–14.
— 1982: Über die Variabilität der dorso-basalen Schädelknickungen bei Haushunden [On the variabil
ity of dorso–basal curvatures in skulls of domestic dogs]. Zoologischer Anzeiger 209: 1–32.
Onar, V.; Pazvant, G.; Gezer Ince, N.; Alpak, H.; Janeczek, M. & Kiziltan, Z. 2013: Morphometric analysis of the foramen magnum of Byzantine dogs excavated in Istanbul Yenikapi at the side of Theodosius harbour. Mediterranean Archaeology and Archaeometry 13: 135–142.
Parker, H.G. et al. 2009: An expressed Fgf4 retrogene is associated with breed-defining chondrodysplasia in domestic dogs. Science 325: 995–998.
Phillips, C.; Baxter, I.L. & Nussbaumer, M. 2009: The application of discriminant function analysis to archaeological dog remains as an aid to the elucidation of possible affinities with modern breeds. Archaeofauna 18: 51–64.
Rackham, J. 1979: Rattus rattus: the introduction of the black rat into Britain. Antiquity 53: 112–120.
Reed, C.A. 1961: Osteological evidences for prehistoric domestication in southwestern Asia. Zeitschrift für Tierzüchtung und Züchstungsbiologie 76(1): 31–38.
— 1969: The pattern of animal domestication in the prehistoric Near East. In: Ucko, P.J. & Dimbleby, G.W. (eds.): The Domestication and Exploitation of Plants and Animals: 361–380. Aldine Press, Chicago.
— 1983: Archaeozoological studies in the Near East: a short history (1960–1980). In: Braidwood, L.S. et al. (eds.): Prehistoric Archaeology along the Zagros Flanks: 511–536. Oriental Institute Publications no.
University of Chicago.
Röhrs, V.M. 1985: Cephalization, neocorticalization and the effects of domestication on brains in mammals. In: Duncker, H.R. & Fleischer, G. (eds.): Functional Morphology in Vertebrates: 544–547. G. Fischer, Stuttgart.
Simoens, P.; Poels, P. & Lauwers, H. 1994: Morphometric analysis of the foramen magnum in Pekingese dogs. American Journal of Veterinary Research 55: 34–39.
Smith, B.P. & Litchfield, C.A. 2009: A review of the relationship between indigenous Australians, dingoes (Canis dingo) and domestic dogs (Canis familiaris). Anthrozöos 22(2): 111–128.
Thalmann, O. et al. 2013: Complete mitochondrial genomes of ancient canids suggest a European origin of domestic dogs. Science 342: 871–874.
Turnbull, P.F. & Reed, C.A. 1974: The fauna from the terminal Pleistocene of Palegawra Cave, a Zarzian occupation site in Northeastern Iraq. Fieldiana: Anthroplogy 63(3): 1–36.
Watson, A.G.; De Lahunta, A. & Evans, H.E. 1989: Dorsal notch of foramen magnum due to incomplete ossification of occipital bone in dogs. Journal of Small Animal Practice 30: 666–673.
Yalden, D. 1999: The History of British Mammals. T. & A. D. Poyser, London.
Zeder, M.A. 2006: A critical assessment of markers of initial domestication in goats (Capra hircus). In: Zeder, M.A.; Emschwiller, E.; Smith, B. & Bradley, D. (eds.): Documenting Domestication: New Genetic and Archaeological Paradigms: 181-206. The University of California Press, Berkeley.
— 2012: The domestication of animals. Journal of Anthropological Research 68(2): 161–190.
— 2015: Core questions in domestication research. Proceedings of the National Academy of Sciences USA 112(11): 3191–3198.