New land tortoises (Testudines: Testudinidae) from the Miocene of Africa

The described fossil testudinids from the Miocene of Africa are reviewed. Geochelone stromeri sp. nov . is described from Lower Pliocene (Langebaanweg) and Miocene (Namib) specimens. Kinixys erosa , an extant species, is reported from Songhor Hill. Chersina sp. is reported from Arrisdrift. Impregnochelys pachytectis gen. et sp. nov . is described from Rusinga Island, Kenya, and is unique in having the anterior shell opening orientated ventrally and in having struts on the internal surface of the neurals, but shares with Kinixys a unique epiplastron shape, a high number of axillary scutes and unique orientation of the head of the femur.     

Characterization of polymorphic microsatellite markers for the endangered Malagasy radiated tortoise (Geochelone radiata)

The radiated tortoise (Geochelone radiata) is an endangered endemic species from Madagascar that inhabits the semiarid spiny forest of the southern part of the island. Habitat destruction and illegal harvesting greatly threaten this species, as attested by the significant reduction of its distribution area in the past 30 years. In order to undertake appropriate conservation actions, it is essential to acquire a better knowledge of its genetic structure. For this study, 145 blood samples were collected from three populations of radiated tortoises in southwestern Madagascar. Eight microsatellite loci were found to be polymorphic, and allelic diversity and observed heterozygosity were high for all markers.     

Seed Dispersal by Geochelone carbonaria and Geochelone denticulata in Northwestern Brazil1

The role of red‐footed tortoises (Geochelone carbonaria) and yellow‐footed tortoises (G. denticulata) as seed dispersal agents was investigated in northwestern Brazil from 5 to 26 January 2002 by analyzing fecal samples for frequency and viability of seed species and estimating daily displacement of tortoises from recaptured and thread‐trailed individuals. Fourteen of 19 fecal samples contained a total of 646 seeds represented by 11 plant species. The most abundant species was Ficus sp. (N= 400) with 100 percent of seeds viable, followed by Aechmea sp. (N = 88) with 93 percent of seeds viable, and Genipa americana (N= 59) with 91 percent of seeds viable. Mean minimum retention time of seeds was 1.6 d and mean daily displacement of tortoises based on recaptured (N= 7) and thread‐trailed tortoises (N= 2) was 57 m. Thus, the diversity and proportion of viable seeds consumed by tortoises, combined with the seed retention times and daily movements, suggest they may be effective dispersal agents. These preliminary findings warrant further investigation into the ecological role of these tortoises in Neotropical ecosystems and their contribution to the maintenance of species diversity and forest structure.     

Hesperotestudo (Testudines: Testudinidae) from the Pleistocene of Bermuda, with comments on the phylogenetic position of the genus

A recently discovered fossil land tortoise (Testudines: Testudinidae) is described from the Pleistocene of Bermuda. Its morphology is sufficiently well preserved to allow assignment to the extinct North American genus Hesperotestudo. However, several features of this tortoise are unique and it is named Hesperotestudo bermudae sp. nov. A review of the phylogenetic relationships of the better known genera of the Testudinidae suggests that the affinities of Hesperotestudo lie with other North American tortoises (Gopherus) and not with Geochelone or other testudinines; thus, Hesperotestudo is reassigned to the Xerobatinae. This is at least the fifth documentation of a testudinid dispersing over open ocean to an oceanic island (the first for Hesperotestudo) and it corroborates the hypothesis that members of this family are well suited to over-water dispersal.     

Evolutionary divergence of giant tortoises in Galapagos

The giant tortoises in the Galapagos Archipelago diverge considerably in size, and in shape and other carapace characteristics. The saddleback morphotype is known only from insular faunas lacking large terrestrial predators (i.e. Galapagos and Mauritius) and in Galapagos is associated with xeric habitats where vertical feeding range and vertical reach in agonistic encounters are adaptive. The large domed morphotype is associated with relatively cool, mesic habitats where intraspecific competition for food and other resources may be less intense than in xeric habitats. Other external characteristics that differ between tortoise populations are also correlated with ecological variation. Tortoises have radiated into a mosaic of ecological conditions in the Galapagos but critical data are lacking on the role of genetic and environmental controls on phenotypic variation. Morphological divergence in tortoises is potentially a better indicator of present ecological conditions than of evolutionary relationships.     

Microsatellite analysis of genetic divergence among populations of giant Galápagos tortoises

Giant Galápagos tortoises represent an interesting model for the study of patterns of genetic divergence and adaptive differentiation related to island colonization events. Recent mitochondrial DNA work elucidated the evolutionary history of the species and helped to clarify aspects of nomenclature. We used 10 microsatellite loci to assess levels of genetic divergence among and within island populations. In particular, we described the genetic structure of tortoises on the island of Isabela, where discrimination of different taxa is still subject of debate. Individual island populations were all genetically distinct. The island of Santa Cruz harboured two distinct populations. On Isabela, populations of Volcan Wolf, Darwin and Alcedo were significantly different from each other. On the other hand, Volcan Wolf showed allelic similarity with the island of Santiago. On Southern Isabela, lower genetic divergence was found between Northeast Sierra Negra and Volcan Alcedo, while patterns of gene flow were recorded among tortoises of Cerro Azul and Southeast Sierra Negra. These tortoises have endured heavy exploitation during the last three centuries and recently attracted much concern due to the current number of stochastic and deterministic threats to extant populations. Our study complements previous investigation based on mtDNA diversity and provides further information that may help devising tortoise management plans.     

Feeding response of the Aldabra giant tortoise (Geochelone gigantea) to island plants showing heterophylly

Aim Heterophylly is present in many plant species on oceanic islands. Almost all of these plants are island endemics, and heterophylly may have evolved as a response to feeding from large insular browsers such as giant tortoises and flightless birds. We tested this anti‐browser hypothesis by feeding Aldabra giant tortoises (Geochelone gigantea) with leaves of native Mauritian plants to see if they distinguished between juvenile and adult leaves and between heteophyllous and homophyllous species. Location Mauritius. Methods In a choice experiment we recorded feeding response of four captive Aldabra giant tortoises to 10 species of Mauritian plants, of which seven were heterophyllous and three homophyllous. Results In general, juvenile leaves of heterophyllous species showed convergence in shape and midrib coloration. Homophyllous foliage was preferred to heterophyllous, and among heterophyllous species adult foliage was preferred to juvenile. Main conclusions Several Mascarene heterophyllous plants show convergence in morphology of juvenile leaves and these are avoided by giant tortoises. This indicates a strong selection history from large browsers such as the giant tortoises. The Mascarene example is in accordance with several other comparable cases of plant‐large browser interactions from other archipelagos.     

The Role of Endangered Species Reintroduction in Ecosystem Restoration: Tortoise–Cactus Interactions on Española Island, Galápagos

We evaluated the role that endangered species reintroduction efforts can play in the larger context of ecosystem restoration. To do so, we examined interactions between endangered giant tortoises (Geochelone nigra hoodensis), currently being reintroduced to Isla Española, Galápagos, and an arboreal cactus (Opuntia megasperma var. megasperma), which is itself endangered and a keystone resource for many animals on the island. We collected information on spatial patterns of occurrence of cacti, tortoises, and woody vegetation and compared recruitment of juvenile cacti in areas occupied versus unoccupied by tortoises. Reintroduced tortoises appeared to suppress cactus recruitment near the few remaining adult cacti at the study site, but facilitate it at longer distances, with tortoise–cactus interactions mediated by the presence of woody vegetation, which likely alters tortoise movements and thereby patterns of cactus seed dispersal. The net effect of tortoises on cacti appeared to be positive insofar as tortoise presence was associated with greater recruitment of juveniles into cactus populations. Our study provides support for reintroducing endangered reptiles and other animals to aid ecosystem restoration in areas where they might once have played an important role in grazing upon and dispersing plants.     

Observations on the reproductive biology of the land tortoises Geochelone emys and Geochelone yniphora in the Honolulu Zoo

Captive reproduction of the Burmese brown tortoise (Geochelone emys) has occurred repeatedly over a five-year period in the Honolulu Zoo. Maximum clutch size (51 eggs) exceeds that reported for other members of the Testudinidae. Nesting behavior in G emys includes construction of a large leaf-litter mound. Collection of nest-building material is accomplished by “back-sweeping” of ground litter within a distance of up to 4 m from the nest site. The nesting female assumes a “nest-guarding” behavior for a period of 2–3 days following egg laying. Eggs incubated artificially hatched in 63–84 days, with egg fertility at 78–84%. One hundred and three G emys were successfully hatched and reared during the period 1978–82. A colony of endangered Madagascar angulated tortoises (Geochelone yniphora) have been maintained at the Honolulu Zoo since 1971. Courtship and mating behavior is similar to that described for the consubgeneric G radiata. Distinctive male “hooking” behavior during courtship in G yniphora is related to the exaggerated epiplastron development in this species. Egg laying has occurred repeatedly in the G yniphora colony since 1979 (mean clutch size = 4.2; N = 6) although no eggs have yet proved fertile.     

Body size development of captive and free‐ranging Leopard tortoises (Geochelone pardalis)

The growth and weight development of Leopard tortoise hatchings (Geochelone pardalis) kept at the Al Wabra Wildlife Preservation (AWWP), Qatar, was observed for more than four years, and compared to data in literature for free‐ranging animals on body weight or carapace measurements. The results document a distinctively faster growth in the captive animals. Indications for the same phenomenon in other tortoise species (Galapagos giant tortoises, G. nigra; Spur‐thighed tortoises, Testudo graeca; Desert tortoises, Gopherus agassizi) were found in the literature. The cause of the high growth rate most likely is the constant provision with highly digestible food of low fiber content. Increased growth rates are suspected to have negative consequences such as obesity, high mortality, gastrointestinal illnesses, renal diseases, “pyramiding,” fibrous osteodystrophy or metabolic bone disease. The apparently widespread occurrence of high growth rates in intensively managed tortoises underlines how easily ectothermic animals can be oversupplemented with nutrients. Zoo Biol 29:517–525, 2010. © 2009 Wiley‐Liss, Inc.