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Concept: Tortoises


The taxonomy of giant Galapagos tortoises (Chelonoidis spp.) is currently based primarily on morphological characters and island of origin. Over the last decade, compelling genetic evidence has accumulated for multiple independent evolutionary lineages, spurring the need for taxonomic revision. On the island of Santa Cruz there is currently a single named species, C. porteri. Recent genetic and morphological studies have shown that, within this taxon, there are two evolutionarily and spatially distinct lineages on the western and eastern sectors of the island, known as the Reserva and Cerro Fatal populations, respectively. Analyses of DNA from natural populations and museum specimens, including the type specimen for C. porteri, confirm the genetic distinctiveness of these two lineages and support elevation of the Cerro Fatal tortoises to the rank of species. In this paper, we identify DNA characters that define this new species, and infer evolutionary relationships relative to other species of Galapagos tortoises.

Concepts: Evolution, Biology, Charles Darwin, Galápagos tortoise, Galápagos Islands, Tortoise, Tortoises, Geochelone


In recent years red-footed tortoises have been shown to be proficient in a number of spatial cognition tasks that involve movement of the animal through space (e.g., the radial maze). The present study investigated the ability of the tortoise to learn a spatial task in which the response required was simply to touch a stimulus presented in a given position on a touchscreen. We also investigated the relation between this task and performance in a different spatial task (an arena, in which whole-body movement was required). Four red-footed tortoises learned to operate the touchscreen apparatus, and two learned the simple spatial discrimination. The side-preference trained with the touchscreen was maintained when behaviour was tested in a physical arena. When the contingencies in the arena were then reversed, the tortoises learned the reversal but in a subsequent test did not transfer it to the touchscreen. Rather they chose the side that had been rewarded originally on the touchscreen. The results show that red-footed tortoises are able to operate a touchscreen and can successfully solve a spatial two-choice task in this apparatus. There was some indication that the preference established with the touchscreen could transfer to an arena, but with subsequent training in the arena independent patterns of choice were established that could be evoked according to the test context.

Concepts: Psychology, Educational psychology, Learning, Knowledge, Tortoise, Tortoises, Geochelone, Red-footed tortoise


To recognize that a picture is a representation of a real-life object is a cognitively demanding task. It requires an organism to mentally represent the concrete object (the picture) and abstract its relation to the item that it represents. This form of representational insight has been shown in a small number of mammal and bird species. However, it has not previously been studied in reptiles. This study examined picture-object recognition in the red-footed tortoise (Chelonoidis carbonaria). In Experiment 1, five red-footed tortoises were trained to distinguish between food and non-food objects using a two-alternative forced choice procedure. After reaching criterion, they were presented with test trials in which the real objects were replaced with color photographs of those objects. There was no difference in performance between training and test trials, suggesting that the tortoises did see some correspondence between the real object and its photographic representation. Experiment 2 examined the nature of this correspondence by presenting the tortoises with a choice between the real food object and a photograph of it. The findings revealed that the tortoises confused the photograph with the real-life object. This suggests that they process real items and photographic representations of these items in the same way and, in this context, do not exhibit representational insight.

Concepts: Reptile, Ontology, Photography, Photograph, Tortoise, Tortoises, Geochelone, Red-footed tortoise


Species are being lost at an unprecedented rate due to human-driven environmental changes. The cases in which species declared extinct can be revived are rare. However, here we report that a remote volcano in the Galápagos Islands hosts many giant tortoises with high ancestry from a species previously declared as extinct: Chelonoidis elephantopus or the Floreana tortoise. Of 150 individuals with distinctive morphology sampled from the volcano, genetic analyses revealed that 65 had C. elephantopus ancestry and thirty-two were translocated from the volcano’s slopes to a captive breeding center. A genetically informed captive breeding program now being initiated will, over the next decades, return C. elephantopus tortoises to Floreana Island to serve as engineers of the island’s ecosystems. Ironically, it was the haphazard translocations by mariners killing tortoises for food centuries ago that created the unique opportunity to revive this “lost” species today.

Concepts: Genetics, Evolution, Charles Darwin, Galápagos tortoise, Galápagos Islands, Tortoise, Tortoises, Aldabra Giant Tortoise


Human-mediated secondary contact of recently diverged taxa offers valuable opportunities for studying the evolutionary mechanisms involved in the establishment and maintenance of genetic boundaries between taxa. We used mitochondrial and microsatellite markers to examine a recently introduced population of the spur-thighed tortoise (Testudo graeca) of mixed origin in the Doñana National Park (SW Spain). The earliest records of tortoises in Doñana trace back to the 18th century, but several population reinforcements in the 20th century with animals from Morocco are well-documented. Consequently, different genetic lineages, which represent distinct subspecies, are thought to co-exist there. Our results confirmed the presence of distinct lineages by revealing that tortoises of the subspecies T. g. marokkensis were introduced into a local allochthonous T. g. graeca population. Unexpectedly, T. g. marokkensis haplotypes exclusively appeared in males, and admixture levels were statistically sex-biased toward males. The sex ratio of the population deviated from parity, with males being 2.36-fold more abundant than females. Our results indicated that population reinforcements had a strong effect on the genetic composition of this population and aggravated its sex ratio deviation. We predict that this sex-biased pattern of introgression is ephemeral and advocated to the near loss of T. g. marokkensis haplotypes.

Concepts: Genetics, Evolution, Sex, 20th century, 18th century, Tortoise, Tortoises, Spur-thighed Tortoise


Restoration of extirpated species via captive breeding has typically relied on population viability as the primary criterion for evaluating success. This criterion is inadequate when species reintroduction is undertaken to restore ecological functions and interactions. Herein we report on the demographic and ecological outcomes of a five-decade-long population restoration program for a critically endangered species of “ecosystem engineer”: the endemic Española giant Galapagos tortoise (Chelonoidis hoodensis). Our analysis of complementary datasets on tortoise demography and movement, tortoise-plant interactions and Española Island’s vegetation history indicated that the repatriated tortoise population is secure from a strictly demographic perspective: about half of tortoises released on the island since 1975 were still alive in 2007, in situ reproduction is now significant, and future extinction risk is low with or without continued repatriation. Declining survival rates, somatic growth rates, and body condition of repatriates suggests, however, that resources for continued population growth are increasingly limited. Soil stable carbon isotope analyses indicated a pronounced shift toward woody plants in the recent history of the island’s plant community, likely a legacy of changes in competitive relations between woody and herbaceous plants induced by now-eradicated feral goats and prolonged absence of tortoises. Woody plants are of concern because they block tortoise movement and hinder recruitment of cactus-a critical resource for tortoises. Tortoises restrict themselves to remnant cactus patches and areas of low woody plant density in the center of the island despite an apparent capacity to colonize a far greater range, likely because of a lack of cactus elsewhere on the island. We conclude that ecosystem-level criteria for success of species reintroduction efforts take much longer to achieve than population-level criteria; moreover, reinstatement of endangered species as fully functioning ecosystem engineers may often require large-scale habitat restoration efforts in concert with population restoration.

Concepts: Biodiversity, Plant, Demography, Endangered species, Extinction, Perennial plant, Galápagos tortoise, Tortoises


In the transition between Mediterranean forest and the arid subtropical shrublands of the southeastern Iberian Peninsula, humans have transformed habitat since ancient times. Understanding the role of the original mosaic landscapes in wildlife species and the effects of the current changes as pine forest plantations, performed even outside the forest ecological boundaries, are important conservation issues. We studied variation in the density of the endangered spur-thighed tortoise (Testudo graeca) in three areas that include the four most common land types within the species' range (pine forests, natural shrubs, dryland crop fields, and abandoned crop fields). Tortoise densities were estimated using a two-stage modeling approach with line transect distance sampling. Densities in dryland crop fields, abandoned crop fields and natural shrubs were higher (>6 individuals/ha) than in pine forests (1.25 individuals/ha). We also found large variation in density in the pine forests. Recent pine plantations showed higher densities than mature pine forests where shrub and herbaceous cover was taller and thicker. We hypothesize that mature pine forest might constrain tortoise activity by acting as partial barriers to movements. This issue is relevant for management purposes given that large areas in the tortoise’s range have recently been converted to pine plantations.

Concepts: Forest, Plantation, Tortoise, Tortoises, Savanna, Shrubland, Spur-thighed Tortoise, Deserts and xeric shrublands


We studied the plant resource use between and within populations of desert tortoise (Gopherus agassizii) across a precipitation gradient in the Sonoran Desert of Arizona. The carbon and nitrogen stable isotope values in animal tissues are a reflection of the carbon and nitrogen isotope values in diet, and consequently represent a powerful tool to study animal feeding ecology. We measured the δ(13)C and δ(15)N values in the growth rings on the shells of tortoises in different populations to characterize dietary specialization and track tortoise use of isotopically distinct C4/CAM versus C3 plant resources. Plants using C3 photosynthesis are generally more nutritious than C4 plants and these trait differences can have important growth and fitness consequences for consumers. We found that dietary specialization decreases in successively drier and less vegetated sites, and that broader population niche widths are accompanied by an increase in the dietary variability between individuals. Our results highlight how individual consumer plant resource use is bounded under a varying regime of precipitation and plant productivity, lending insight into how intra-individual dietary specialization varies over a spatial scale of environmental variability.

Concepts: Photosynthesis, Photorespiration, C4 carbon fixation, Tortoise, Tortoises, C3 carbon fixation, Desert Tortoise, Gopherus


The extinction of large herbivores, often keystone species, can dramatically modify plant communities and impose key biotic thresholds that may prevent an ecosystem returning to its previous state and threaten native biodiversity. A potentially innovative, yet controversial, landscape-based long-term restoration approach is to replace missing plant-herbivore interactions with non-native herbivores. Aldabran giant (Aldabrachelys gigantea) and Madagascan radiated (Astrochelys radiata) tortoises, taxonomically and functionally similar to the extinct Mauritian giant tortoises (Cylindraspis spp.), were introduced to Round Island, Mauritius, in 2007 to control the non-native plants that were threatening persistence of native species. We monitored the response of the plant community to tortoise grazing for 11 months in enclosures before the tortoises were released and, compared the cost of using tortoises as weeders with the cost of using manual labor. At the end of this period, plant biomass; vegetation height and cover; and adult, seedling, flower, and seed abundance were 3-136 times greater in adjacent control plots than in the tortoise enclosures. After their release, the free-roaming tortoises grazed on most non-native plants and significantly reduced vegetation cover, height, and seed production, reflecting findings from the enclosure study. The tortoises generally did not eat native species, although they consumed those native species that increased in abundance following the eradication of mammalian herbivores. Our results suggest that introduced non-native tortoises are a more cost-effective approach to control non-native vegetation than manual weeding. Numerous long-term outcomes (e.g., change in species composition and soil seed bank) are possible following tortoise releases. Monitoring and adaptive management are needed to ensure that the replacement herbivores promote the recovery of native plants. Estudiando el Potencial para Restaurar Ecosistemas Históricos de Forrajeo con Reemplazos Ecológicos de Tortugas Terrestres.

Concepts: Species, Plant, Ecology, Seed, Extinction, Herbivore, Tortoise, Tortoises


Seasonal migration has evolved in many taxa as a response to predictable spatial and temporal variation in the environment. Individual traits, physiology and social state interact with environmental factors to increase the complexity of migratory systems. Despite a huge body of research, the ultimate causes of migration remain unclear. A relatively simple, tractable system - giant tortoises on Santa Cruz Island, Galapagos, was studied to elucidate the roles of environmental variation and individual traits in a partial migratory system. Specifically, we asked: (i) do Galapagos tortoises undergo long-distance seasonal migrations? (ii) is tortoise migration ultimately driven by gradients in forage quality or temperature; and (iii) how do sex and body size influence migration patterns? We recorded the daily locations of 17 GPS-tagged tortoises and walked a monthly survey along the altitudinal gradient to characterize the movements and distribution of tortoises of different sizes and sexes. Monthly temperature and rainfall data were obtained from weather stations deployed at various altitudes, and the Normalized Difference Vegetation Index was used as a proxy for forage quality. Analyses using net displacement or daily movement characteristics did not agree on assigning individuals as either migratory or non-migratory; however, both methods suggested that some individuals were migratory. Adult tortoises of both sexes move up and down an altitudinal gradient in response to changes in vegetation dynamics, not temperature. The largest tagged individuals all moved, whereas only some mid-sized individuals moved, and the smallest individuals never left lowland areas. The timing of movements varied with body size: large individuals moved upward (as lowland forage quality declined) earlier in the year than did mid-sized individuals, while the timing of downward movements was unrelated to body size and occurred as lowland vegetation productivity peaked. Giant tortoises are unlikely candidates for forage-driven migration as they are well buffered against environmental fluctuations by large body size and a slow metabolism. Notably the largest, and presumably most dominant, individuals were most likely to migrate. This characteristic and the lack of sex-based differences in movement behaviour distinguish Galapagos tortoise movement from previously described partial migratory systems.

Concepts: Human migration, Charles Darwin, Galápagos tortoise, Galápagos Islands, Tortoise, Tortoises, Geochelone, Darwin's finches