Elisabeth Vrba

**Elisabeth S. Vrba** (Hanburgo, 1942ko maiatzaren 17a - New Haven, 2025eko otsailaren 5a) hegoafrikar paleontologoa  izan zen, aldaketa pultsuaren hipotesia garatu zuelako eta Stephen Jay Gould lankidearekin exaptazio hitza sortu zuelako ezaguna.

## Ibilbidea 
Vrba Hanburgon jaio zen, Alemanian. 1944an, aita hil ondoren, familiarekin batera Namibiako ardi-ustiategi batera joan zen bizitzera. Vrba 1965ean Zoologia eta Estatistikan graduatu zen  eta baita ere zientzietako batxiler gisa  Lurmutur Hiriko Unibertsitatean. 1966 eta 1967 artean biologia molekularrean ikerkuntzan aritu zen. 1968an bigarren hezkuntzako irakasle gisa lan egin zuen Pretoriako San Alban Ikastetxean. 1969tik aurrera, Pretoriako Transvaal Museoko Paleontologia eta Paleoantropologia sailean lan egin zuen. Zoologian eta paleontologian doktoretza ikasketak egiteko egon zen han, doktoretza lortzeko. 1973tik 1986ra Transvaal Museoko Paleontologia eta Paleoantropologia saileko burua izan zen. Zoologia eta Paleontologia doktoregoa lortu zuen 1974an. Bobidoen familian (antilopeak, etab.) garatu zuen bere lana. 

Kromdraai guneko indusketa-zuzendaria izan zen 1976 eta 1986 artean, eta Transvaal Museoko zuzendari laguntzailea 1977 eta 1986 artean. 1986an Estatu Batuetara joan zen, Yale Unibertsitateko Geologia eta Geofisika Sailean unibertsitateko katedradun izateko. Peabody Museoko  arduraduna zan zen 1987tik. 1995ean Yale Unibertsitateko Geologia eta Geofisika Saileko lehendakaria izan zen. 1996 eta 1999 bitartean Yaleko Zientzia Bioesferikoaren Institutuko zuzendaria izan zen, eta 1992-2010 bitartean Unibertsitate bereko Ecosave Zentroko zuzendaria.

## Berrikuntzak 
Vrba eta bere lankide Stephen Jay Gould exaptazioaren teoriagatik dira ezagunak. Vrba eta Goulden ikerketak ezaugarri genetiko baten jatorri historikoak ez duela beti bere funtzio garaikidea islatzen iradoki zuen.  Egokitzapen genetikoek funtzio berriak har ditzakete, eta eboluzioan aurrerago beste helburu bat lortzeko balio diezaiokete espezie bati. Vrba eta Goulden exaptazioaren ideia ere kritikatu dute azken urteotan adituek, ezaugarri genetikoak hainbat faktoreren presiopean daudela baieztatzen baitute, eta horrek zaildu egiten du noiz dagoen egokitzapena edo exaptazioa jokoan zehaztea.

Vrbak aldaketa-pultsuaren hipotesia ere garatu zuen, teoria makroebolutiboaren gehigarri esanguratsua. 

## Hedabideetan agerpenak 
Vrbak *Planeta Milagroso* (2005) dokumental sailean eta *Ein Zufall im Paradies* (1999) dokumental austriarrean parte hartu du bere burua aurkeztuz.

## Argitaratutako lanak 
* Gould, SJ eta ES Vrba. (mila bederatziehun eta laurogeita bi). "Exaptation-A Missing Term in the Science of Form" . *Paleobiologia* 8: 4-15.
* Vrba, ES eta Gould, SJ (1986). "Sailkapenaren eta hautaketaren hedapen hierarkikoa". *Paleobiologia* . 12 (2): 217-228.
* Vrba, ES (1993). "The Pulse That Produced Us". *Historia natural* **102** (5) 47-51.
* Vrba, ES, 1975.  Some evidence of chronology and palaeoecology of Sterkfontein, Swartkrans and Kromdraai from the fossil Bovidae. *Nature* 254(5498):301-304.
* Vrba, E.S. and Grine, FE, 1978. Australopithecine enamel prism patterns. Science 202:890 892.
* Vrba, E.S., 1979. A new study of the scapula of Australopithecus africanus from Sterkfontein. Am. (0 URLa). Anthrop. 51(1):117-129.
* Vrba, E.S, 1980.  The significance of bovid remains as indicators of environment and predation patterns. In Fossils in the Making, A.K. Behrensmeyer and A.P. Hill, eds., The University of Chicago Press:Chicago, pp. 247-271.
* Vrba, E.S, 1980. Evolution, species and fossils: how does life evolve? S. Sateliteak. J. Sci. 76(2):61-84.
* Grine F.E. and Vrba, ES, 1980. Prismatic enamel: a preadaptation for mammalian diphyondonty? S. Afrika J. Sci. 76(3):139-140.
* Gould SM and Vrba, ES, 1982. Exaptation - a missing term in the science of form. Paleobiology 8(1):4-15.
* Vrba, ES, 1983. Macroevolutionary trends: new perspectives on the roles of adaptation and incidentaleffect. Science 221(4608):387-389.
* Greenacre, M.J. and Vrba, E.S., 1984. A Correspondencia Analysis of biological census data. Ecology 65(3):984-997.
* Vrba, E.S. and Eldredge, N., 1984. Individual, hierarchies and processes: towards a more complete evolutionary theory. Paleobiology 10(2):123-164.
* Vrba, ES, 1985. Ecological and adaptive changes associated with early hominid evolution. In Ancestors: The Hard Evidence, E. Delson, ed., Alan R. Liss, Inc.: New York, pp. 63-71.
* Vrba, ES, 1985. Species and Speciation. Transvaal Museum Monograph 4. Transvaal Museum, Pretoria.
* Vrba, ES, 1985.  Environment and evolution: alternative causes of the temporal distribution of evolutionary events. S. Sateliteak. J. Sci. 81(5):229-236.
* Vrba, E.S. and Gould SJ, 1986. The hierarchial expansion of sorting and selection: sorting and selection cannot equated. Paleobiology 12(2)217-228.
* Vrba, E.S., 1987. Ecology in relation to speciation rates: some case histories of Miocene-Recent mamal clades. Evolutionary Ecology 1:283 300.
* Vrba, ES, 1989.  Levels of selection and sorting, with special reference to the species level. Oxford Surveys of Evolutionary Biology, Vol. 6 :111-168.Vrba, E.S. 1992. Mammals as a key to evolutionary theory. Plenary Keynote Address, 70th Congress of American Society of Mammalogists, June 1990, Frostburg, Maryland. Journal of Mammalogy 73(1):1-28.
* Gatesy, J. E., Yelon, D., DeSalle, R., and Vrba, E. S. 1992. Phylogenetic analysis based on mitochondrial DNA of the Bovidae, Mammalia. Molec. Biol. Evol. 9(3):433-446.
* Vrba, E. S.. 1993. Turnover-pulses, the Red Queen, and related topics. American Journal of Science 293-A:418 452. White, T.W., Suwa, G., Hart, W.K., Walter, R.C., WoldeGabriel, G., De Heinzelin, J., Clark, J.D., Asfaw, B.,and Vrba, E.S. 1993. New discoveries of Australopithecus at Maka in Ethiopia. Nature 366:261 265.
* Vrba, E. S., Vaisnys, J. R., Gatesy, J. E., DeSalle, R., and K-Y. Yen 1994. Analysis of paedomorphosis using allometric characters: the example of Reduncini antelopes (Bovidae, Mammalia). Systematic Biology 43 (1):92-116.
* Clark, J.D., De Heinzelin, J., Schick, K., Hart, W., White, T., WoldeGabriel, R. Walter, Suwa, g., Asfaw, B., Vrba, E. S. and Y. Hator-Selassie. 1994. Middle Pleistocene discoveries in the Middle Awash Valley, Ethiopia. Science 264:1907 10.
* Vrba, E.S. 1995. Species as habitat specific, complex systems. In The Recognition of Species: Speciation and the Recognition Concept [J.C. Masters, D.M. Lambert and H.G. Spencer, eds. ], pp. 3-44. Johns Hopkins University Press.
* Vrba, E. S., 1995. The fossil record of African Antelopes (Mammalia, Bovidae) in relation to human evolution and paleoclimate. Pp. 385-424. In E. S. Vrba, G. H. Denton, T. C. Partridge and L. H. Burckle (eds. ), Paleoclimate and Evolution with emphasis on Human Origins, pp.. Yale University Press, New Haven, Connecticut. 556 or.
* Vrba, E. S., 1995. On the connections between paleoclimate and evolution. Pp. 24-45. In E. S. Vrba, G. H. Denton, T. C. Partridge and L. H. Burckle (eds. ), Paleoclimate and Evolution with emphasis on Human Origins, Yale University Press, New Haven, Connecticut. 556 or.
* Vrba, E. S., G. H. Denton, T. C. Partridge and L. H. Burckle. Editors, 1995. Editors Paleoklimate and Evolution with emphasis on Human Origins, Yale University Press, New Haven, Connecticut, 556 or.
* Vrba, ES, 1996. Climate, heterochrony, and human evolution. Journal of Anthropological Research 52 (1): 1-28.
* Schaller, G.B. and Vrba, E.S. 1996. Description of the Giant Muntjac (Megamuntiacus vuquangensis) in Laos. Journal of Mammalogy 77 (3): 675-683.
* Gatesy, J., Amato, G., Vrba, E., Schaller, G. and DeSalle, R. 1997. A cladistic analysis of mitochondrial ribosomal DNA from the Bovidae. Molecular Phylogenetics and Evolution: 7(3): 303-319.Vrba, E.S. 1998. (Monografikoa) New fossils of Alcelaphini and Caprinae (Bovidae, Mammalia) from Awash, Ethiopia, and phylogenetic analysis of Alcelaphini. Palaeontologia afrikarra 34: 127-198.
* Vrba, E.S. 1998.  Multiphasic growth models and the evolution of prolonged growth exemplified by human brain evolution.  Journal of Theoretical Biology 190 : 227-239.
* de Heinzelin, J., Clark, J.D., White, T., Hart, W., Renne, P., WoldeGabriel, g., Beyene, Y. and Vrba, E.S. 1999. Environment and behavior of 2.5-million-year old Bouri hominids. Science 284 : 625-629.
* Vrba, E.S. 1999. Habitat theory in relation to the evolution in African Neogene biota and hominids. in T.G. Bromage and F. Schrenk (eds. ), African Biogeography, Climate Change, and Early Hominid Evolution, pp. 19-34. Part of New Series on Human Evolution (Series Eds Wood, B. and Ciochon, R.). Oxford University Press, Oxford, United Kingdom.
* Vrba, E.S. and Schaller, G.B. Editors 2000. Antelopes, Deer, and Relatives : Fossil Record, Behavioral Ecology, Systematics, and Conservation. Yale University Press, New Haven, Connecticut.
* Vrba, E.S. and Schaller, G.B. 2000. Phylogeny of Bovidae (Mammalia) based on behavior, glands and skull morphology. pp. 203-222 In Vrba, ES. and 2004, G.B. (arg.) Antelopes, Deer, and Relatives: Fossil Record, Behavioral Ecology, Systematics, and Conservation. Yale University Press, New Haven, Connecticut.
* Vrba, E.S. 2000. Major features of Neogene Mammalian Evolution in Africa. In T.C. Partridge and R. Maud (eds. ), Cenozoic Geology of Southern Africa, 277-304. Oxford University Press, Oxford, United Kingdom.
* Vrba, E.S. 2002. Exaptation In Pagel, M. Encyclopedia of Evolution, 123-126. Oxford University Press, Oxford, United Kingdom.
* Asfaw, B., Gilbert, H., Beyene, Y., Hart, W. Renne, P. WoldeGabriel, G., Vrba, E.S., and White, T. 2002. Remains of Homo erectus from Bouri, Middle Awash, Ethiopia. Nature 416 (6878): 317-320.
* DeGusta, D. And Vrba, E.S. 2003. A method for inferring paleohabitats from the functional morphology of bovid astragali. Journal of Archaeological Science 30(8): 1009-1022.
* Vrba, E.S. 2004. Ecology, evolution, and development : Perspectives from the fossil record. In Hall, B.K., Pearson, R.D., and Muller, G.B. (arg.) Environment, Development, and Evolution, pp. 85-105. Massachusetts Institute of Technology Press, Cambridge, Massachusetts.
* Vrba, E. S. & DeGusta, D. 2004 Do species populations really start small? New perspectives from the Late Neogene fossil record of African mammals. Phil. Trans. Reala. Lond. B 359, 285-293.
* Haile-Selassie, Y. Woldegabriel, G., White, T.D., Bernor, R.L., DeGusta, D., Renne, P.R., Hart, W.K., Vrba, E., Stanley, A., and Howell, F.H. 2004. Mio-Pliocene mammals from the Middle Awash, Ethiopia. Geobios 37(4): 536-552.
* Vrba, E.S. and Eldredge, N. (Editors) 2005. Macroevolution: Diversity and Disparity. Stephen Jay Gould-en Dedikated issue of Paleobiology in honor, Paleobiology 31(2).Hernández Fernández, M. and Vrba, E.S. 2005. Rapoport Effect and biomic specialization in African mammals: revisiting the Climatic Variability Hypothesis. Journal of Biogeography 32: 903-918.
* Hernández Fernández, M. and Vrba, E.S. 2005. A complete estimate of the phylogenetic relationships in Ruminantia: a dated species-level supertree of the extant ruminants. Biological Reviews 80: 269-302.
* Hernández Fernández, M. and Vrba, E.S. 2005. Body size, biomic secialization, and range size of African large mammals. Journal of Biogeography 32: 1243-1256.
* Vrba, E.S. 2005. Mass turnover and heterochrony events in response to physical change Paleobiology 31(2): 157-174.
* Hernández Fernández, M. and Vrba, E.S. 2005. A complete estimate of the phylogenetic relationships in Ruminantia: a dated species-level supertree of the extant ruminants. Evolutionary Ecology 19: 199-219.
* Vrba, E.S. and Hareeta-Selassie, Y. 2006. A new antelope, Zephyreduncinus oundagaissensis (Reduncini, Artiodactyla, Bovidae) from the Late Miocene of the Middle Awash, Afar Rift, Ethiopia. Journal of Structate Paleontology 26(1): 213-218. (2006ko martxoa)
* Vrba, E.S. 2007. Role of Environmental Stimuli in Hominid Origins. In Henke,W., Rothe, H., and Tattersall, I. (arg.) Handbook of palaeoanthropology, Vol. 3: Phylogeny of Hominines, pp. 1-41. Springer-Verlag: New York.
* White, Tim D., Stanley H. Ambrose, Gen Suwa, Denise F. Su, David DeGusta, Raymond L. Bernor, Jean-Renaud Boisserie, Michel Brunet, Eric Delson, Stephen Frost, Nuria Garcia, Ioannis Giaomark, Claud, Shell., Kheel, Boishin Boissheel, Bois, Kheel, Kheel, Kheel Kheat Kheat, Kheat, Kheat, Kheat, Kheat, Kheat, Kheat, Kheat, Kheat Kheat, Kheat Kheat Kheat Kheat, Kheat, Kheat, Kheat, Kheat, Kheat, Kland, Kheat Kheat Kheat, Kheat, Kheat, Kheat, Kheat, Kheat, Kheat, Kland, 2009. Makroornogabea Paleontology and the Pliocene Habitat of Ardipithecus ramidus. Science 326 (5949): 87-93.
* Bibi, F., Bukhsianidze, M., Gentry, A.W., Geraads, D., Kostopoulos, D., and Vrba, E.S. 2009. The fossil record and evolution of Bovidae: State of the field. Palaeontologia elektronikoa: 12(3):10A.
* Bibi, F. and Vrba, E.S. 2010. Unraveling bovin phylogeny: accomplishments and challenges. BMC Biology 8(50) : 1-4.

## Sariak eta aintzatespenak 
* *Elkarte Britainiarraren Domina* jaso du, berrogei urtetik beherako ikertzaile gazteei ematen zaiena.
* *Urteko Emakume Profesionala Afrikan* izendatu zuten (1982). 
* 1985ean *Hego Afrikako Errege Elkarteko kide* izateko hautatu zuten. 
* Yale Unibertsitateko Geologia eta Geofisika Saileko fakultateko kide izan zen 1980ko hamarkadaren hasieratik eta baita ere *irakasle emeritua*.

## Erreferentziak 
## Kanpo estekak