Morphological divergence among populations of Testudo graeca from west-central Morocco

Morphometric analyses were carried out on Testudo graeca from west-central Morocco, and compared to previously published mitochondrial DNA sequence variation. We measured 41 characters on 244 tortoises from three localities, including one population of T. g. soussensis from Souss Valley. All three populations could be statistically differentiated in a multivariate space using these characters. The population from the Jbilet mountains was the most differentiated (no overlap), those from Essaouira on the coast and from Admine showed limited overlap (5%). Sexual size dimorphism (males < females) remained constant between sites. Sexual shape dimorphism was prominent in all populations, but the degree of dimorphism differed for some characters when corrected for size. Tortoises from Jbilet living under harsher conditions (low precipitation, high thermal amplitude, low plant cover) were smaller, lighter, more flattened and less dimorphic in shape than the others. Although the Admine population (currently assigned to T. g. soussensis) was morphologically distinct, its distinctiveness was less supported than in the case of Jbilet; several other qualitative traits previously considered as defining this subspecies were also questionable. Moreover, this morphological variation contrasted with the available genetic evidence (12S rRNA mtDNA) which did not reveal significant variation between any of the populations. This implies that morphological differences have either arisen very recently or can be attributed to phenotypic plasticity. This should be taken into account when using morphological traits for taxonomic considerations and conservation management.

[1]  R. Shine,et al.  Sexual size dimorphism and sexual selection in turtles (order testudines) , 2004, Oecologia.

[2]  R. Shine,et al.  Relationships between sexual dimorphism and niche partitioning within a clade of sea-snakes (Laticaudinae) , 2002, Oecologia.

[3]  K. Nagy,et al.  Sexual dimorphism in steppe tortoises (Testudo horsfieldii): influence of the environment and sexual selection on body shape and mobility , 2001 .

[4]  R. Shine Sexual size dimorphism and male combat in snakes , 2004, Oecologia.

[5]  J. Goudsmit,et al.  Phylogenetic relationships among the species of the genus Testudo (Testudines: Testudinidae) inferred from mitochondrial 12S rRNA gene sequences. , 2002, Molecular phylogenetics and evolution.

[6]  Le Houérou,et al.  Classification écoclimatique des zones arides (s.l.) de l'Afrique du Nord , 1989 .

[7]  R. A. Anderson,et al.  Sexual selection versus alternative causes of sexual dimorphism in teiid lizards , 1990, Oecologia.

[8]  A. Hailey,et al.  Variation of adult body size of the tortoise Testudo hermanni in Greece: proximate and ultimate causes , 1999 .

[9]  D. Harris,et al.  Genetic variation in Testudo graeca from Morocco estimated using 12S rRNA sequencing , 2003 .

[10]  R. Shine,et al.  Sexual dimorphism in snakes: different reproductive roles favour different body plans , 1998, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[11]  Fred L. Bookstein,et al.  A Comment on Shearing as a Method for “Size Correction” , 1987 .

[12]  A. Boumezzough,et al.  Partage des ressources spatiales et tropiques au sein d’un peuplement de lézards insectivores des Jbilets centrales (Maroc occidental) , 2000, Revue d'Écologie (La Terre et La Vie).

[13]  J. Iverson A revised checklist with distribution maps of the turtles of the world , 1992 .

[14]  D. Germano,et al.  Growth and population structure of the Moorish tortoise (Testudo graeca graeca) in westcentral Morocco: possible effects of over-collecting for the tourist trade. , 2005 .

[15]  SEXUAL DIMORPHISM IN LIZARD BODY SHAPE: THE ROLES OF SEXUAL SELECTION AND FECUNDITY SELECTION , 2002, Evolution; international journal of organic evolution.

[16]  A. Hailey,et al.  Variation in adult survival rate of the tortoise Testudo hermanni in Greece: implications for evolution of body size , 2001 .

[17]  A. Andreu,et al.  Clutch frequency, egg and clutch characteristics, and nesting activity of spur-thighed tortoises, Testudo graeca, in southwestern Spain , 1996 .

[18]  Anthony Herrel,et al.  Sexual dimorphism of head size in Gallotia galloti: testing the niche divergence hypothesis by functional analyses , 1999 .

[19]  R. Shine,et al.  Post-hatching environment contributes greatly to phenotypic variation between two populations of the Australian garden skink, Lampropholis guichenoti , 2000 .

[20]  A. Hailey,et al.  Sexual dimorphism of body size and shell shape in European tortoises , 2003 .

[21]  D. Germano Shell Morphology of North American Tortoises , 1993 .

[22]  G. C. Packard,et al.  Environmentally induced variation in body size and condition in hatchling snapping turtles (Chelydra serpentina) , 1999 .

[23]  H. Harmon,et al.  Geographic Variation in Sexual Dichromatism in the Collared Lizard, Crotaphytus collaris (Sauria: Crotaphytidae) , 1997 .

[24]  Monteiro,et al.  Shape distances, shape spaces and the comparison of morphometric methods. , 2000, Trends in ecology & evolution.

[25]  R. Shine,et al.  Geographic variation in lizard phenotypes: importance of the incubation environment , 1998 .

[26]  Jxw,et al.  " ' ' Observations on Ecological Changes Threatening a Population of Testado graeca graeca in the Souss Valley , Southern Morocco , 2022 .

[27]  J. A. Mateo,et al.  Mitochondrial DNA haplotyping of Testudo graeca on both continental sides of the Straits of Gibraltar. , 2000, The Journal of heredity.

[28]  K. Crandall,et al.  Considering evolutionary processes in conservation biology. , 2000, Trends in ecology & evolution.

[29]  K. Nagy,et al.  Sexual size dimorphism in steppe tortoises (Testudo horsfieldi): growth, maturity, and individual variation , 2001 .

[30]  A. Hailey,et al.  BODY MASS CONDITION IN GREEK TORTOISES: REGIONAL AND INTERSPECIFIC VARIATION , 2002 .

[31]  R. Shine,et al.  THE INFLUENCE OF NATURAL INCUBATION ENVIRONMENTS ON THE PHENOTYPIC TRAITS OF HATCHLING LIZARDS , 1997 .

[32]  Brian Groombridge,et al.  1996 IUCN Red List of Threatened Animals , 1996 .

[33]  B. McArdle The structural relationship: regression in biology , 1988 .

[34]  J. Mellado S.O.S. Souss: argan forest destruction in Morocco , 1989, Oryx.

[35]  A. Hailey,et al.  Egg size and shape, clutch dynamics, and reproductive effort in European tortoises , 1988 .

[36]  S. Nylin,et al.  Adaptive plasticity and plasticity as an adaptation : a selective review of plasticity in animal morphology and life history , 1995 .

[37]  Michael LaBarbera,et al.  ANALYZING BODY SIZE AS A FACTOR IN ECOLOGY AND EVOLUTION , 1989 .