Spatial patterns and evolutionary processes in southern South America: A study of dental morphometric variation
Corresponding Author
Valeria Bernal
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
División Antropología, Facultad de Ciencias Naturales y Museo, Museo de La Plata, Paseo del Bosque s/n, La Plata (1900), ArgentinaSearch for more papers by this authorS. Ivan Perez
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorPaula N. Gonzalez
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorMarina L. Sardi
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorHéctor M. Pucciarelli
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorCorresponding Author
Valeria Bernal
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
División Antropología, Facultad de Ciencias Naturales y Museo, Museo de La Plata, Paseo del Bosque s/n, La Plata (1900), ArgentinaSearch for more papers by this authorS. Ivan Perez
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorPaula N. Gonzalez
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorMarina L. Sardi
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorHéctor M. Pucciarelli
División Antropología, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, CONICET, Buenos Aires, Argentina
Search for more papers by this authorAbstract
The purpose of this article is to examine the patterns of evolutionary relationships between human populations from the later Late Holocene (1,500–100 years BP) of southern South America on the basis of dental morphometric data. We tested the hypotheses that the variation observed in this region would be explained by the existence of populations with different phylogenetic origin or differential action of gene flow and genetic drift. In this study, we analyzed permanent teeth from 17 samples of male and female adult individuals from throughout southern South America. We measured mesiodistal and buccolingual diameters at the base of the crown, along the cement–enamel junction. The results of multiple regression analysis and a mantel correlogram indicate the existence of spatial structure in dental shape variation, as the D2 Mahalanobis distance between samples increases with increasing geographical distance between them. In addition, the correlation test results show a trend toward reduction of the internal variation of samples with increasing latitude. The detected pattern of dental variation agrees with the one expected as an outcome of founder serial effects related to an expansion of range during the initial occupation of southern South America. Am J Phys Anthropol, 2010. © 2009 Wiley-Liss, Inc.
Supporting Information
Additional Supporting Information may be found in the online version of this article.
Filename | Description |
---|---|
AJPA_21206_sm_suppinfoappendix1.doc38.5 KB | Supporting Information Appendix 1 |
AJPA_21206_sm_suppinfoappendix2.doc302 KB | Supporting Information Appendix 2 |
Please note: The publisher is not responsible for the content or functionality of any supporting information supplied by the authors. Any queries (other than missing content) should be directed to the corresponding author for the article.
LITERATURE CITED
- Alvesalo L,Tigerstedt PMA. 1974. Heritabilities of human tooth dimensions. Hereditas 77: 311–318.
- Aschero CA,Goñi RA,Civalero MT,Molinari RL,Espinosa SL,Guraieb AG,Bellelli CT. 2005. Holocenic park: Arqueología del Parque Nacional Perito Moreno. Anales de Parques Nacionales 17: 71–119.
- Barbujani G. 2000. Geographical patterns: how to identify them, and why. Hum Biol 72: 133–153.
- Barrientos G,Perez SI. 2004. La expansión y dispersión de poblaciones del norte de Patagonia durante el Holoceno tardío: evidencia arqueológica y modelo explicativo. In: MT Civalero, PM Fernandez, AG Guraieb, editors. Contra viento y marea. Arqueología de Patagonia. Buenos Aires: Instituto Nacional de Antropología y Pensamiento Latinoamericano. p 179–195.
- Barrientos G,Perez SI,Bernal V,González P,Béguelin M,Del Papa M. 2005. Changing views about the local evolution of human populations in the southeastern pampas of Argentina during the Holocene. In: S Zakrzewski, M Clegg, editors. Proceedings of the 5th Annual BABAO Conference, BAR International Series 1383. Oxford: Archeopress. p 93–104.
- Béguelin M. 2009. Variación geográfica en la morfología del esqueleto postcraneal de las poblaciones humanas de Pampa y Patagonia durante el Holoceno tardío: una aproximación morfométrica, Unpublished Ph.D. thesis, Universidad Nacional de La Plata, La Plata.
- Béguelin M,Bernal V,Del Papa M,Novellino P,Barrientos G. 2006. El poblamiento humano tardío del sur de Mendoza y su relación con el Norte de Patagonia: una discusión bioarqueológica. In: V Durán, V Cortegoso, editors. Arqueología y ambiente de áreas naturales protegidas de la provincia de Mendoza. Mendoza: Anales de Arqueología y Etnología 61. p 5–25.
- Bernal V. 2008. Procesos de diferenciación biológica entre poblaciones humanas del Holoceno tardío de Patagonia. Una aproximación desde la variación morfométrica dental, Unpublished Ph.D. thesis, Universidad Nacional de La Plata, La Plata.
- Bernal V,Gonzalez PN,Perez SI,Pucciarelli HM. 2008a. Entierros humanos del noreste de Patagonia: nuevos fechados radiocarbónicos. Magallania 36: 175–184.
- Bernal V,Perez SI,Gonzalez PN. 2006. Variation and causal factors of craniofacial robusticity in Patagonian hunter-gatherers from late Holocene. Am J Hum Biol 18: 748–765.
- Bernal V,Sardi ML,Pucciarelli HP. 2008b. Evaluación de la señal filogenética en datos dentales del sur americano. X Congreso de la Asociación Latinoamericana de Antropología Biológica. La Plata ALAB.
- Borrero LA. 1999. The prehistoric exploration and colonization of Fuego-Patagonia. J World Prehist 13: 321–355.
- Borrero LA. 2001a. El poblamiento de la Patagonia. Toldos, milodones y volcanes. Buenos Aires: Emecé.
-
Borrero LA.
2001b.
Cambios, continuidades, discontinuidades: discusiones sobre arqueología Fuego-Patagónica. In:
EE Berberián, AE Nielsen, editors.
Historia Argentina prehispánica, Tomo 2.
Córdoba:
Brujas. p
815–838.
10.1007/s007050170150 Google Scholar
- Buikstra J,Ubelaker D. 1994. Standards for data collection from human skeletal remains. Archaeological Survey Research series 44. Fayetteville, AR: Archaeological Survey Research.
- Cavalli-Sforza LL,Menozzi P,Piazza A. 1994. The history and geography of human genes. Princeton: Princeton University Press.
- Cheverud JM,Routman EJ,Irschick DK. 1997. Pleiotropic effects of individual gene loci on mandibular morphology. Evolution 51: 2004–2014.
- Clapperton CM. 1992. La última glaciación y deglaciación en el Estrecho de Magallanes, implicaciones para el poblamiento de Tierra del Fuego. An Inst Patagon 21: 113–128.
- Cocilovo JA. 1981. Estudio sobre discriminación y clasificación de poblaciones prehispánicas del NO Argentino. Publicación Ocasional No. 36. Santiago: Museo Nacional de Historia Natural.
- Cocilovo JA,Di Rienzo J. 1984–1985. Un modelo biológico para el estudio del poblamiento prehispánico del territorio argentino. Correlación fenético-espacial. Relac Soc Argent Antropol 16: 119–135.
- Cocilovo JA,Guichón RA. 1985–1986. Propuesta para el Estudio de las Poblaciones Aborígenes del Extremo Austral de Patagonia. An Inst Patagon/Soc 16: 111–123.
- Darroch JN,Mosimann JE. 1985. Canonical and principal component of shape. Biométrika 72: 241–252.
- Dejean CB,Lanata JL,Martino L,Carnese FR,Osella A. 2007. Demografía y distribución de haplogrupos mitocondriales durante la dispersión inicial en las Américas. Rev Argent Antropol Biol 9: 51.
- Dempsey PJ,Townsend GC. 2001. Genetic and environmental contributions to variation in human tooth size. Heredity 86: 685–693.
- Dillehay TD. 2000. The settlement of the Americas. New York: Basic Books.
- Excoffier L. 2004. Patterns of DNA sequence diversity and genetic structure after a range expansion: lessons from the infinite-island model. Mol Ecol 13: 853–864.
- Fabra M,Laguens AG,Demarchi DA. 2007. Human colonization of the central territory of Argentina: design matrix models and craniometric evidence. Am J Phys Anthropol 133: 1060–1066.
- Franco NV,Borrero LA. 1996. El stress temporal y los artefactos líticos. La cuenca superior del Río Santa Cruz. In: J Gómez Otero, editor. Arqueología. Sólo Patagonia. Puerto Madryn: Centro Nacional Patagónico. p 341–348.
- García-Bour J,Pérez-Pérez A,Alvarez S,Fernandez A,Lopez-Parra AM,Arroyo-Pardo E,Turbon D. 2004. Early population differentiation in extinct aborigines from Tierra del Fuego-Patagonia: ancient mtDNA sequences and Y chromosome STR characterization. Am J Phys Anthropol 123: 361–370.
- Goñi R,Barrientos G,Cassiodoro G. 2000–2002. Las condiciones previas a la extinción de las poblaciones humanas del sur de Patagonia: una discusión a partir del análisis de la estructura del registro arqueológico de la cuenca del Lago Salitroso. Cuad Antropol Pensam Latinoam 19: 249–266.
- González-José R. 2003. El poblamiento de la Patagonia. Análisis de la variación craneofacial en el contexto del poblamiento Americano, Unpublished Ph.D. thesis, Universidad de Barcelona, Barcelona.
- González-José R,Dahinten S,Hernandez M. 2001a. The settlement of Patagonia: a matrix correlation study. Hum Biol 73: 233–248.
- González-José R,Dahinten SL,Luis MA,Hernández M,Pucciarelli HM. 2001b. Craniometric variation and the settlement of the Americas: testing hypotheses by means of R-matrix and matrix correlation analyzes. Am J Phys Anthropol 116: 154–165.
- González-José R,Martínez Abadías N,Van Der Molen S,García Moro C,Dahinten S,Hernández M. 2004. Hipótesis acerca del poblamiento de Tierra del Fuego-Patagonia a partir del análisis genético-poblacional de la variación craneofacial. Magallania 32: 79–78.
- Graw M. 2001. Significance of the classical morphological criteria for identifying gender using recent skulls. Forensic Sci Commun 2001: 1.
- Hair JF,Anderson RE,Tatham RL,Black WC. 2005. Multivariate data analysis. Englewood Cliffs, NJ: Prentice Hall.
- Hamilton MJ,Milne BT,Walker RS,Burger O,Brown JH. 2007. The complex structure of hunter-gatherer social networks. Proc R Soc Lond B Biol Sci 274: 2195–2202.
- Harzer W. 1995. Size relationships and interactions between crown diameters in twins and family members. In: RJ Radlanski, H Renz, editors. Proceedings of the 10th international symposium on dental morphology. Berlin: “M” Marketing Services. p 124–135.
-
Hernández M,Lalueza C,García-Moro C.
1997.
Fueguian cranial morphology: the adaptation to a cold, harsh environment.
Am J Phys Anthropol
103:
103–117.
10.1002/(SICI)1096-8644(199705)103:1<103::AID-AJPA7>3.0.CO;2-X PubMed Web of Science® Google Scholar
- Heusser CJ. 1993. Late Quaternary forest-steppe contact zone. Isla Grande de Tierras del Fuego, Subantarctic South America. Quat Sci Rev 12: 169–178.
- Hillson SW,Fitzgerald CM,Flinn HM. 2005. Alternative dental measurements: proposals and relationships with other measurements. Am J Phys Anthropol 126: 413–426.
- Hutchison DW,Templeton AR. 1999. Correlation of pairwise genetic and geographic distance measures: inferring the relative influences of gene flow and drift on the distribution of genetic variability. Evolution 53: 1898–1914.
- Imbelloni J. 1937. Fuéguidos y Láguidos. Posición actual de la raza paleo-americana o de Lagoa Santa. An Mus Argent Cien Nat 39: 6–103.
- Johnson GA. 1982. Organizational structure and scalar stress. In: C Renfrew, M Rowlands, BA Reagraves, editors. Theory and explanation in archaeology. New York: Academic Press. p 389–421.
- Johnson JB,Omland KS. 2004. Model selection in ecology and evolution. Trends Ecol Evol 19: 101–108.
- Jungers WL,Falsetti A,Wall CE. 1995. Shape, relative size and size-adjustments in morphometrics. Yearb Phys Anthropol 38: 137–161.
- Kelly RL. 2003. Colonization of new land by hunter-gatherers. Expectations and implications based on ethnographic data. In: M Rockman, J Steele, editors. Colonization of unfamiliar landscapes. Londres: Routledge. p 44–58.
- Lahr MM. 1995. Patterns of modern human diversification: implications for Amerindian origins. Am J Phys Anthropol 38: 163–198.
-
Lalueza Fox C,Hernández M,García Moro C.
1996.
Craniometric analysis in groups from Tierra del Fuego/Patagonia and the peopling of the south extreme of the Americas.
Hum Evol
11:
217–224.
10.1007/BF02436625 Google Scholar
- Lanata JL,Garcia A. 2005. Environmental corridors and early human dispersal in South America. Presented at the 70th Annual Meeting of the Society for American Archaeology, Salt Lake City, UT, March 30–April 3,2005.
- Lanata JL,Martino L,Osella A,Garcia-Herbst A. 2008. Demographic conditions necessary to colonize new spaces: the case for early human dispersal in the Americas. World Archaeol 40: 520–537.
- Lapointe FJ,Legendre P. 1992. Statistical significance of the matrix correlation coefficient for comparing independent phylogenetic trees. Syst Biol 41: 378–384.
- Leamy LJ,Routman EJ,Cheverud JM. 1999. Quantitative trait loci for early- and late developing skull characters in mice: a test of the genetic independence model of morphological integration. Am Nat 153: 201–214.
- Legendre P. 1993. Spatial autocorrelation: trouble or new paradigm? Ecology 74: 1659–1673.
- Legendre P. 2002. Program for multiple linear regression (ordinary or through the origin) with permutation test-user's notes, Département de Sciences Biologiques, Université de Montréal, Montreal.
- Legendre P,Fortin MJ. 1989. Spatial pattern and ecological analysis. Vegetatio 80: 107–138.
- Legendre P,Lapointe F-J,Casgrain P. 1994. Modelling brain evolution from behavior: a permutational regression approach. Evolution 48: 1478–1499.
- Legendre P,Legendre L. 2003. Numerical ecology, 2nd ed. Amsterdam: Elsevier.
- Lieberman DE. 1999. Homology and hominid phylogeny: problems and potential solutions. Evol Anthropol 7: 142–151.
- Mahalanobis PC. 1936. On the generalised distance in statistics. Proc Natl Inst Sci India 2: 49–55.
- Merriwether DA,Rothhammer F,Ferrell RE. 1995. Distribution of the four founding lineage haplotypes in Native Americans suggests a single wave of migration for the New World. Am J Phys Anthropol 98: 411–430.
- Miotti LL,Salemme MC. 2004. Poblamiento, movilidad y territorios entre las sociedades cazadoras recolectoras de Patagonia. Complutum 15: 177–200.
- Moraga M,Rocco P,Miquel JF,Nervi F,Llop E,Chakraborty R,Rothhammer F,Carvallo P. 2000. Mitochondrial DNA polymorphisms in Chilean aboriginal populations: implications for the peopling of the southern cone of the continent. Am J Phys Anthropol 113: 19–29.
- Neme G,Gil A,Duran V. 2005. Late Holocene in southern Mendoza (northwestern Patagonia): radiocarbon pattern and human occupation. Before Farming 2:Article 5.
- Neves WA,Powell JF,Ozolins EG. 1999. Extra-continental morphological affinities of Palli-Aike, Southern Chile. Interciencia 24: 258–263.
- Oden NL,Sokal RR. 1986. Directional autocorrelation: an extension of spatial correlograms to two dimensions. Syst Zool 35: 608–617.
-
Pearson OM,Millones M.
2005.
Rasgos esqueletales de adaptación al clima y a la actividad entre los habitantes aborígenes de tierra del fuego.
Magallania
33:
37–50.
10.4067/S0718-22442005000100003 Google Scholar
- Perez SI. 2006. El poblamiento holocénico del Sudeste de la Región Pampeana: un estudio de morfometría geométrica craneofacial, Unpublished Ph.D. thesis, Universidad Nacional de La Plata, La Plata.
- Perez SI,Béguelin M,Del Papa M. 2004. Evaluación de relaciones biológicas regionales y extra regionales de muestras del N.O. de Santa Cruz. In: MT Civalero, PM Fernández, AG Guráieb, editors. Contra viento y marea. Arqueología de Patagonia. Buenos Aires: Instituto Nacional de Antropología y Pensamiento Latinoamericano y Sociedad Argentina de Antropología. p 437–359.
- Perez SI,Bernal V,Gonzalez P. 2007. Morphological differentiation of aboriginal human populations from Tierra del Fuego (Patagonia): implications for South American peopling. Am J Phys Anthropol 133: 1067–1079.
- Perez SI,Monteiro LM. 2009. Nonrandom factors in modern human morphological diversification: a study of craniofacial variation in southern South American populations. Evolution 63: 978–993.
- Peterson MA,Denno RF. 1998. The influence of dispersal and diet breadth on patterns of genetic isolation by distance in phytophagous insects. Am Nat 152: 428–446.
- Phenice TW. 1969. A newly developed visual methods of sexing os pubis. Am J Phys Anthropol 30: 297–301.
- Powell JF,Neves WA. 1999. Craniofacial morphology of the first Americans: pattern and process in the peopling of the New World. Yearb Phys Anthrpol 42: 153–188.
- Pucciarelli H,Neves W,González-José R,Sardi M,Ramírez Rozzi F,Struck A,Bonilla MY. 2006. East–west cranial differentiation in pre-Columbian human populations of South America. Homo 57: 133–150.
-
R Development Core Team.
2008.
R: a language and environment for statistical computing.
Vienna:
R Foundation for Statistical Computing.
10.1890/0012-9658(2002)083[3097:CFHIWS]2.0.CO;2 Google Scholar
- Ramachandran S,Deshpande O,Roseman CC,Rosenberg NA,Feldman MW,Cavalli-Sforza LL. 2005. Support from the relationship of genetic and geographic distance in human populations for a serial founder effect originating in Africa. Proc Natl Acad Sci USA 102: 15942–15947.
- Relethford JH. 2004. Global patterns of isolation by distance based on genetic and morphological data. Hum Biol 76: 499–513.
- Relethford JH. 2008. Rmet 5.0. Computer software for Windows. http://konig.la.utk.edu/relethsoft.html.
- Relethford JH,Blangero J. 1990. Detection of differential gene flow from patterns of quantitative variation. Hum Biol 62: 5–25.
- Relethford JH,Harpending HC. 1994. Craniometric variation, genetic theory and modern human origins. Am J Phys Anthropol 95: 249–270.
-
Relethford JH,Lees FC.
1982.
The use of quantitative traits in the study of human population structure.
Yearb Phys Anthropol
25:
113–132.
10.1002/ajpa.1330250507 Google Scholar
- Ross AH,Ubelaker DH,Guillén S. 2008. Craniometric patterning within ancient Peru. Lat Am Antiq 19: 158–166.
- Rothhammer F,Silva C. 1990. Craniometrical variation among South American prehistoric populations: climatic, altitudinal, chronological and geographic contributions. Am J Phys Anthropol 82: 9–17.
- Sardi ML. 2002. Diferenciación craneofacial en aborígenes de la Patagonia y su relación con grupos americanos y extra-americanos, Unpublished Ph.D. thesis, Universidad Nacional de La Plata, La Plata.
- Sardi ML,Novellino PS,Pucciarelli HM. 2006. Craniofacial morphology in the Argentine center-west: consequences of the transition to food production. Am J Phys Anthropol 130: 333–343.
- Sardi ML,Ramírez Rozzi F,González-José R,Pucciarelli HM. 2005. South Amerindian craniofacial morphology: diversity and implications for Amerindian evolution. Am J Phys Anthropol 128: 747–756.
- Schafer JL. 1999a. Multiple imputation: a primer. Stat Methods Med Res 8: 3–15.
- Schafer JL. 1999b. NORM 2.03. http://www.stat.psu.edu/∼jls/misoftwa.html.
- Schafer JL,Olsen MK. 1998. Multiple imputation for multivariate missing-data problems: a data analyst's perspective. Multivariate Behav Res 33: 545–571.
- Scherer AK. 2007. Population structure of the classic period Maya. Am J Phys Anthropol 132: 367–380.
- Schurr TG. 2004. The peopling of the New World: perspectives from molecular anthropology. Ann Rev Anthropol 33: 551–583.
- Sharma K,Corruccini RS,Henderson AM. 1985. Genetic variance in dental dimensions of Punjabi twins. J Dent Res 64: 1389–1391.
-
Shrout PE,Fleiss JL.
1979.
Intraclass correlations: uses in assessing rater reliability.
Psychol Bull
2:
420–428.
10.1037/0033-2909.86.2.420 Google Scholar
- Slatkin M. 1993. Isolation by distance in equilibrium and non-equilibrium populations. Evolution 47: 264–279.
- Smouse PE,Long JC,Sokal RR. 1986. Multiple regression and correlation extensions of the Mantel test of matrix correspondence. Syst Zool 35: 627–632.
- Sokal RR. 1984. Comment to “Beals KL, Smith CL, Dodd SM. 1984. Brain size, cranial morphology, climate, and time machines”. Curr Anthropol 25: 322–323.
- Sokal RR. 1986. Spatial data analysis and historical processes. In: E Diday, L Escoufier, J Lebart, J Pages, Y Schektman, R Tomassone, editors. Data analysis and informatics, IV. Proceedings of the Fourth Internacional Symposium on Data Analysis and Informatics, Versailles, France. p 29–43.
- Sokal RR,Geoffrey MJ,Wooten MC. 1989a. Spatial autocorrelation analysis of migration and selection. Genetics 121: 845–855.
- Sokal RR,Harding RM,Oden NL. 1989b. Spatial patterns of human gene frequencies in Europe. Am J Phys Anthropol 80: 267–294.
- Sparks CS,Jantz RL. 2002. A reassessment of human cranial plasticity: Boas revisited. PNAS 99: 14636–14639.
- Sparks CS,Jantz RL. 2003. Changing times, changing faces: Franz Boas' Immigrant study in modern perspective. Am Anthropol 105: 333–337.
- Sperber GH. 2004. The genetics of odontogenesis: implications in dental anthropology and palaeoodontology. Dent Anthropol 17: 1–7.
- Steele J,Politis G. 2008. AMS 14C dating of early human occupation of southern South America. J Archaeol Sci 36: 419–429.
- Stefan VH. 2004. Assessing intrasample variation: analysis of Rapa Nui (Easter Island) museum cranial collections example. Am J Phys Anthropol 124: 45–58.
- Stojanowski CM. 2004. Population history of native groups in pre- and postcontact Spanish Florida: aggregation, gene flow, and genetic drift on the Southeastern U.S. Atlantic Coast. Am J Phys Anthropol 123: 316–332.
- Sturges H. 1926. The choice of a class-interval. J Am Statist Assoc 21: 65–66.
- Templeton AR. 1998. Nested clade analyses of phylogeographic data: testing hypotheses about gene flow and population history. Mol Ecol 7: 381–397.
- Templeton AR. 2007. Genetics and recent human evolution. Evolution 61: 1507–1519.
- Templeton AR,Routman E,Phillips CA. 1995. Separating population structure from population history: a cladistic analysis of the geographical distribution of mitochondrial DNA haplotypes in the tiger salamander, Ambystoma tigrinum. Genetics 140: 767–782.
- Weber DC,Skillings JH. 2000. A first course in the design of experiments. A linear models approach. Boca Raton: CRC Press.
- Wegmann D,Currat M,Excoffier L. 2006. Molecular diversity after a range expansion in heterogeneous environments. Genetics 174: 2009–2020.
- Workman MS,Leamy LJ,Routman EJ,Cheverud JM. 2002. Analysis of QTL effects on the size and shape of mandibular molars in mice. Genetics 160: 1573–1586.
- Wright S. 1943. Isolated by distance. Genetics 28: 114–138.
-
Zar JH.
1999.
Biostatistical analysis.
New York:
Prentice Hall.
10.1111/j.1461-0248.2004.00674.x Google Scholar