Giant Tortoises Research Articles

Genetic Pedigree Analysis of the Pilot Breeding Program for the Rediscovered Galapagos Giant Tortoise from Floreana Island.

An aim of many captive breeding programs is to increase population sizes for reintroduction and establishment of self-sustaining wild populations. Genetic analyses play a critical role in these programs: monitoring genetic variation, identifying the origin of individuals, and assigning parentage to track family sizes. Here, we use genetic pedigree analyses to examine 3 seasons of a pilot breeding program for the Floreana island Galapagos giant tortoise, C. niger, that had been declared extinct for ~150 years until individuals with mixed ancestry were recently discovered. We determined that 8 of 9 founding individuals were assigned parentage to at least 1 of 130 offspring produced, though there was considerable reproductive skew. In addition, we observed that genetic diversity of the progeny was lower than that of the founders. Despite the observed reproductive skew, we did not see evidence for assortative mating based on relatedness, but there was a trend toward reduced fitness when more related individuals bred. Finally, we found that the majority of progeny had ancestry assigned to the Floreana species (mean ± SE = 0.51 ± 0.02), though individual estimates varied. The success of these pilot seasons bodes well for a larger breeding program to help restore the previously extinct tortoise from Floreana island. Future efforts should continue to monitor for reproductive skew and assortative mating to maintain allelic diversity. We would also recommend forming smaller breeding groups and rotating individuals among them to prevent long-term reproductive skew among pairs.

Net displacement and temporal scaling: Model fitting, interpretation and implementation

Abstract Net displacement is an integral component of numerous ecological processes and is critically dependent on the tortuosity of a movement trajectory and hence on the temporal scale of observation. Numerous attempts have been made to quantitatively describe net displacement while accommodating tortuosity, typically evoking a power law, but scale‐dependency in tortuosity limits the utility of approaches based on power law relationships that must assume scale‐invariant tortuosity. We describe a phenomenological model of net displacement that permits both scale‐variant and scale‐invariant movement. Movement trajectories are divided into pairs of relocations specifying start‐ and end‐points, and net displacements between points are calculated across a vector of time intervals. A bootstrap is implemented to create new datasets that are independent both across and within time intervals, and the model is fitted to the bootstrapped dataset using log–log regression. We apply this model to simulated trajectories and both fine‐grain and coarse‐grain trajectories obtained from an Aldabra giant tortoise Aldabrachelys gigantea, African elephants Loxodonta africana, black‐backed jackals Canis mesomelas and Northern elephant seals Mirounga angustirostris. The model was able to quantify the characteristics of net displacement from simulated movement trajectories corresponding to both scale‐variant (e.g. correlated random walks) and scale‐invariant (e.g. random walk) movement models. Furthermore, the model produced identical outputs across time vectors corresponding to different intervals and absolute ranges of time for scale‐invariant models. The model characterized the tortoise as generally exhibiting long scale‐invariant steps, which was corroborated by visual comparison of model outputs to observed trajectories. Elephants, jackals and seals exhibited movement parameters consistent with their known movement behaviours (nomadism, territoriality and widely ranging searching). We describe how the model may be used to compare movements within and between species, for example by partitioning movement into scale‐variant and scale‐invariant components, and by calculating a unitless net displacement scaled to the basal movement capacities of an animal. We also identify several useful derived quantities and realistic parameter ranges and discuss how the model may be implemented in a variety of ecological studies.

Patterns of activity and body temperature of Aldabra giant tortoises in relation to environmental temperature.

We studied the temperature relations of wild and zoo Aldabra giant tortoises (Aldabrachelys gigantea) focusing on (1) the relationship between environmental temperature and tortoise activity patterns (n = 8 wild individuals) and (2) on tortoise body temperature fluctuations, including how their core and external body temperatures vary in relation to different environmental temperature ranges (seasons; n = 4 wild and n = 5 zoo individuals). In addition, we surveyed the literature to review the effect of body mass on core body temperature range in relation to environmental temperature in the Testudinidae. Diurnal activity of tortoises was bimodally distributed and influenced by environmental temperature and season. The mean air temperature at which activity is maximized was 27.9°C, with a range of 25.8–31.7°C. Furthermore, air temperature explained changes in the core body temperature better than did mass, and only during the coldest trial, did tortoises with higher mass show more stable temperatures. Our results, together with the overall Testudinidae overview, suggest that, once variation in environmental temperature has been taken into account, there is little effect of mass on the temperature stability of tortoises. Moreover, the presence of thermal inertia in an individual tortoise depends on the environmental temperatures, and we found no evidence for inertial homeothermy. Finally, patterns of core and external body temperatures in comparison with environmental temperatures suggest that Aldabra giant tortoises act as mixed conformer–regulators. Our study provides a baseline to manage the thermal environment of wild and rewilded populations of an important island ecosystem engineer species in an era of climate change.

Trophic interactions between larger crocodylians and giant tortoises on Aldabra Atoll, Western Indian Ocean, during the Late Pleistocene.

Today, the UNESCO World Heritage Site of Aldabra Atoll is home to about 100 000 giant tortoises, Aldabrachelys gigantea, whose fossil record goes back to the Late Pleistocene. New Late Pleistocene fossils (age ca. 90–125 000 years) from the atoll revealed some appendicular bones and numerous shell fragments of giant tortoises and cranial and postcranial elements of crocodylians. Several tortoise bones show circular holes, pits and scratch marks that are interpreted as bite marks of crocodylians. The presence of a Late Pleistocene crocodylian species, Aldabrachampsus dilophus, has been known for some time, but the recently found crocodylian remains presented herein are distinctly larger than those previously described. This indicates the presence of at least some larger crocodylians, either of the same or of a different species, on the atoll. These larger crocodylians, likely the apex predators in the Aldabra ecosystem at the time, were well capable of inflicting damage on even very large giant tortoises. We thus propose an extinct predator–prey interaction between crocodylians and giant tortoises during the Late Pleistocene, when both groups were living sympatrically on Aldabra, and we discuss scenarios for the crocodylians directly attacking the tortoises or scavenging on recently deceased animals.

Do Species Really Matter?

Many environmentalists hold that the loss of a species is intrinsically bad, and many also think that we have moral obligations to species as such. In an attempt to capture these thoughts, some philosophers have suggested that species are bearers of intrinsic value. This approach works well in paradigmatic cases. However, it begins to break down in more difficult scenarios, such as when species boundaries are unclear or when resources are scarce. The case study of the Galápagos giant tortoises in this essay illustrates the limitations of traditional accounts of the non-instrumental value of species. Careful attention to this case indicates how species-centric accounts diverge from the evaluative attitudes of environmentalists, and suggests new directions for theoretical replacements.

When Earth was planet of the giant tortoises

Tortoises evolved into giants on at least seven occasions and on four continents. The finding undermines the long-standing idea that tortoises become enormous only if they are stranded on remote islands. There are more than 40 species of tortoise, the most spectacular being the giant tortoises. On the Galapagos islands in the Pacific and Seychelles islands in the Indian Ocean, these animals can have shells more than 120 centimeters long. Evangelos Vlachos at the Museum of Paleontology Egidio Feruglio in Trelew Argentina, and Marton Rabi at the University of Halle-Wittenberg Germany, wanted to clarify the history of gigantism. They drew the tortoise family tree using data from extinct and living species.

Self-righting potential and the evolution of shell shape in Gal\xe1pagos tortoises

Self-righting, the capacity of an animal to self-turn after falling on its back, is a fitness-related trait. Delayed self-righting can result in loss of mating opportunities or death. Traits involved in self-righting may therefore be under selection. Galápagos giant tortoises have two main shell morphologies - saddleback and domed – that have been proposed to be adaptive. The more sloped shape on the sides of the shell and the longer extension of neck and legs of the saddlebacks could have evolved to optimize self-righting. The drier environments with more uneven surfaces where the saddleback tortoises occur increases their risk to fall on their back while walking. The ability to fast overturn could reduce the danger of dying. To test this hypothesis, we used 3D shell reconstructions of 89 Galápagos giant tortoises from three domed and two saddleback species to compare self-righting potential of the two shell morphotypes. Our results indicate that saddleback shells require higher energy input to self-right than domed ones. This suggests that several traits associated with the saddleback shell morphology could have evolved to facilitate self-righting. Studying the functional performances of fitness-related traits, as in this work, could provide important insight into the adaptive value of traits.

Theory, practice, and conservation in the age of genomics: The Galápagos giant tortoise as a case study.

High‐throughput DNA sequencing allows efficient discovery of thousands of single nucleotide polymorphisms (SNPs) in nonmodel species. Population genetic theory predicts that this large number of independent markers should provide detailed insights into population structure, even when only a few individuals are sampled. Still, sampling design can have a strong impact on such inferences. Here, we use simulations and empirical SNP data to investigate the impacts of sampling design on estimating genetic differentiation among populations that represent three species of Galápagos giant tortoises (Chelonoidis spp.). Though microsatellite and mitochondrial DNA analyses have supported the distinctiveness of these species, a recent study called into question how well these markers matched with data from genomic SNPs, thereby questioning decades of studies in nonmodel organisms. Using >20,000 genomewide SNPs from 30 individuals from three Galápagos giant tortoise species, we find distinct structure that matches the relationships described by the traditional genetic markers. Furthermore, we confirm that accurate estimates of genetic differentiation in highly structured natural populations can be obtained using thousands of SNPs and 2–5 individuals, or hundreds of SNPs and 10 individuals, but only if the units of analysis are delineated in a way that is consistent with evolutionary history. We show that the lack of structure in the recent SNP‐based study was likely due to unnatural grouping of individuals and erroneous genotype filtering. Our study demonstrates that genomic data enable patterns of genetic differentiation among populations to be elucidated even with few samples per population, and underscores the importance of sampling design. These results have specific implications for studies of population structure in endangered species and subsequent management decisions.

Assessing colour preference in Aldabra giant tortoises (Geochelone gigantea)

Tortoises perceive different colours and rely on the visual system to find food. However, few studies have considered colour preference in tortoises, especially in land species. The aim of this study is to investigate whether Aldabra giant tortoises (Geochelone gigantea) housed in Parco Natura Viva (VR), an Italian zoological garden, show a specific colour preference in their social context. The study was divided into two different periods in which red and yellow balls of the same size were scattered around in the outdoor enclosure. In the first period, pieces of carrots were placed behind each ball whereas in the second period, carrots were replaced with apples. Data on the frequencies of interaction with red and yellow balls were collected. First, tortoises interacted more with the balls when pieces of apples rather than carrots were hidden behind them. No significant group-level colour preference was found; however, individual-level variation in colour preference was reported. In addition, tortoises interacted significantly more with the yellow balls in the second than in the first period. Food typology seems therefore to play an important role in determining colour preference in chelonians. Research aimed at identifying individual differences in animal preferences might be valuable to improve captive animal husbandry (e.g.: development of enrichment programmes, diets and rewards).

The early scientific history of Galapagos tortoises

Specimens of Galapagos tortoises (Testudo, now Chelonoidis) obtained by scientific institutions up to and including the voyage of the Beagle (1835) are reviewed, along with the scientific literature of that period. The idea that all giant tortoises (Galapagos and Indian Ocean) were a single species (Testudo indica) was challenged as early as 1817 by Amasa Delano and refuted scientifically by Harlan in 1826, although his publication was widely ignored; some authors continued to use T. indica for Galapagos tortoises at least until 1855. That Galapagos tortoises varied from island to island was reported first by David Porter in 1815, but such differences continued to be in the realm of hearsay until well after the publication of the second edition of Darwin's Journal of researches in 1845. Contrary to what has been written by modern authors, Darwin did not think that tortoises had been introduced to the Galapagos from elsewhere. Also contrary to popular mythology, tortoises did not play an important role in the development of Darwin's evolutionary theories.

Rewilding defaunated Atlantic Forests with tortoises to restore lost seed dispersal functions

The extinction of frugivores has been considered one of the main drivers of the disruption of important ecological processes, such as seed dispersal. Many defaunated forests are too small to restore function by reintroducing large frugivores, such as tapirs or Ateline monkeys, and the long-term fate of large-seeded plants in these areas is uncertain. However, such small fragments still host many species and play relevant ecosystem services. Here, we explore the use of two tortoise species, the red-footed tortoise (Chelonoidis carbonarius) and the yellow-footed tortoise (Chelonoidis denticulatus), as ecological substitutes for locally extinct large seed dispersers in small forest patches in the Brazilian Atlantic Forest. We employed prior knowledge on the known occurrences of Chelonoidis species and used ecological niche modeling (ENM) to identify forest patches for tortoise rewilding. Based on habitat suitability, food availability and conservation co-benefits, we further refined our analysis and identified that the more suitable areas for tortoise reintroduction are forest patches of northern Atlantic Forest, areas with high defaunation intensity. Giant tortoises have been used to restore lost ecological services in island ecosystems. We argue that reintroducing relatively smaller tortoises is an easy-to-use/control conservation measure that could be employed to partially substitute the seed dispersal services of extinct large disperser species, mitigating the negative cascading effects of defaunation on reducing plant diversity.

A genomic perspective timely needed for re-evaluating the species delimitations, evolutionary trajectories, and conservation strategies of the Galapagos giant tortoises

Conservation policy in the giant Galápagos tortoise, an iconic endangered animal, has been assisted by genetic markers for ~15 years: a dozen loci have been used to delineate thirteen (sub)species, between which hybridization is prevented. Here, comparative reanalysis of a previously published NGS data set reveals a conflict with traditional markers. Genetic diversity and population substructure in the giant Galápagos tortoise are found to be particularly low, questioning the genetic relevance of current conservation practices. Further examination of giant Galápagos tortoise population genomics is critically needed.

Identification of Genetically Important Individuals of the Rediscovered Floreana Gal\xe1pagos Giant Tortoise (Chelonoidis elephantopus) Provides Founders for Species Restoration Program

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.

Los primeros viajes audiovisuales (1936) de Rolf Blomberg en Ecuador

El objetivo del presente artículo es indagar, a partir del análisis de los filmes “En Canoa a la Tierra de los Reductores de Cabezas” y “Vikingos en las Islas de las Tortugas Gigantes” de Rolf David Blomberg (Suecia, 1912- Ecuador, 1996), la producción y reproducción de imágenes de exploradores europeos en Ecuador durante la época de entre-guerras (1919-1939). Con el mencionado objetivo pretende poner en discusión la construcción de los “viajes audiovisuales” a partir del cruce teórico entre las narrativas de las crónicas de exploradores y las miradas del cine etnográfico. Aventureros como Blomberg, que transitaban ambos campos, nos hacen dar cuenta de una fragilidad en los límites de los mismos. Metodológicamente se propone en el presente artículo un encuentro entre el análisis del modo de representación de los filmes con los libros publicados por el mismo autor. Las preguntas que van a guiar este trabajo son ¿Qué tipo de narrativas e imaginarios construyeron los “viajes audiovisuales” de los exploradores/realizadores europeos en Ecuador? ¿Cómo circulaban esas ideas? ¿Cuáles elementos incorporan de las crónicas de exploradores y cuáles del cine etnográfico de la época? ¿Cuál es la importancia de estudiar este tipo de exploradores/realizadores desde y para la Antropología Audiovisual?

Contribution by vertebrates to seed dispersal effectiveness in the Galápagos Islands: a community-wide approach.

Seed dispersal and seedling recruitment are crucial phases in the life cycle of all spermatophyte plants. The net contribution of seed dispersers to plant establishment is known as seed dispersal effectiveness (SDE) and is defined as the product of a quantitative (number of seeds dispersed) and a qualitative (probability of recruitment) component. In Galápagos, we studied the direct contribution to SDE (number of seeds dispersed and effect on seedling emergence) provided by the five island groups of frugivores (giant tortoises, lizards, medium-sized passerine birds, small non-finch passerine birds, and finches) in the two main habitats in this archipelago: the lowland and the highland zones, and found 16 vertebrate species dispersing 58 plant species. Data on frequency of occurrence of seeds in droppings and number of seeds dispersed per unit area produced contrasting patterns of seed dispersal. Based on the former, giant tortoises and medium-sized passerines were the most important seed dispersers. However, based on the latter, small non-finch passerines were the most important dispersers, followed by finches and medium-sized passerines. The effect of disperser gut passage on seedling emergence varied greatly depending on both the disperser and the plant species. Although the contribution to SDE provided by different disperser guilds changed across plant species, medium-sized passerines (e.g., mockingbirds) provided a higher contribution to SDE than lava lizards in 10 out of 16 plant species analysed, whereas lava lizards provided a higher contribution to SDE than birds in five plant species. While both the quantitative and qualitative components addressed are important, our data suggests that the former is a better predictor of SDE in the Galápagos archipelago.

Human-mediated dispersals do not explain tortoise distribution on the Indian Ocean's islands

Origins of giant tortoises on the Indian Ocean's islands have been debated, and most recently attributed to human translocation (see Wilme, Patrick, & Ganzhorn, 2017). To resolve the issue, we analyse all available molecular sequences from extinct and extant Indian Ocean's giant tortoises, along with major clades of the family Testudinidae using phylogenetic methods, Bayesian inference and maximum likelihood, and a relaxed time calibration approach. Our results most strongly support giant tortoises of the genus Cylindraspis evolving in situ in the Mascarene Islands since the early mid-Miocence, and Aldabra tortoises diverging from a Madagascan lineage in the early Oligocene. The geologic time-scale of these speciation events and the resulting island endemism for each lineage do not support human translocation, but rather demonstrate the impressive ability of giant tortoises to disperse long distances across oceans.

A new species of extinct Late Quaternary giant tortoise from Hispaniola.

Insular giant tortoise diversity has been depleted by Late Quaternary extinctions, but the taxonomic status of many extinct populations remains poorly understood due to limited available fossil or subfossil material, hindering our ability to reconstruct Quaternary island biotas and environments. Giant tortoises are absent from current-day insular Caribbean ecosystems, but tortoise remains from Quaternary deposits indicate the former widespread occurrence of these animals across the northern Caribbean. We report new Quaternary giant tortoise material from several cave sites in Pedernales Province, southern Dominican Republic, Hispaniola, representing at least seven individuals, which we describe as Chelonoidis marcanoi sp. nov. Although giant tortoise material was first reported from the Quaternary record of Hispaniola almost 35 years ago, tortoises are absent from most Quaternary deposits on the island, which has been studied extensively over the past century. The surprising abundance of giant tortoise remains in both vertical and horizontal caves in Hispaniola's semi-arid ecoregion may indicate that this species was adapted to open dry habitats and became restricted to a habitat refugium in southeastern Hispaniola following climatic-driven environmental change at the Pleistocene-Holocene boundary. Hispaniola's dry forest ecosystem may therefore have been shaped by giant tortoises for much of its evolutionary history.

Giant tortoise habitats under increasing drought conditions on Aldabra Atoll—Ecological indicators to monitor rainfall anomalies and related vegetation activity

Aldabra Atoll has the largest population of giant tortoises (Aldabrachelys gigantea) in the world. As such an important biological resource, it is necessary to understand how the effects of climate change will impact this keystone species; in particular the frequency of drought, which is likely to affect tortoise habitat. To assess whether drought frequency has changed over the last 50 years on Aldabra, we calculated the standardized precipitation index (SPI) to identify drought periods using monthly rainfall data collected during 1969–2013. We found that drought frequency has increased to more than six drought months per year today compared with about two months per year in the 1970s (t=2.884, p=0.006). We used MODIS normalized difference vegetation index (NDVI) as a proxy for vegetation activity, to determine how vegetation has responded to the changing drought frequency between 2000 and 2013. We found that Aldabra’s vegetation is highly responsive to changes in rainfall: anomalies in long-term mean monthly NDVI across Aldabra were found to decrease below the mean during most drought periods and increase above the mean during most non-drought periods. To investigate the response of tortoise habitat to rainfall, we extracted mean NDVI anomalies for three key habitat types. Open mixed scrub and grasslands, the preferred habitat of tortoises, showed the greatest decrease in vegetation activity during drought periods, and the greatest increase in average greenness during non-drought periods. Recent analysis has shown vegetation changes on Aldabra in recent decades. If these changes are caused by decreased precipitation, then the increased frequency of drought could impact the tortoise population, in both the short and long term, by limiting the quality and quantity of forage and/or shade availability within favoured habitats, and by changing the habitat composition across the atoll.

A Conference on the Origin (and Evolution) of Baryonic Galaxy Halos

A conference was held in March 2017 in the Galapagos Islands on the topic of The Origin (and Evolution) of Baryonic Galaxy Halos. It attracted some 120 researchers from around the world. They presented 68 talks (nine of which were invited) and 30 posters over five days. A novel element of the talk schedule was that participants were asked which talks they wanted to hear and the schedule was made up based on their votes and those of the Scientific Organizing Committee SOC . The final talk schedule had 34% of the talks given by women. An emphasis was given to discussion time directly after each talk. Combined with limited/no access to the internet, this resulted in high level of engagement and lively discussions. A prize was given to the poster voted the best by participants. A free afternoon included organized excursions to see the local scenery and wildlife of the Galapagos (e.g., the giant tortoises). Four public talks were given, in Spanish, for the local residents of the town. A post-conference survey was conducted, with most participants agreeing that the conference met their scientific needs and helped to initiate new research directions. Although it was challenging to organize such a large international meeting in such an isolated location as the Galapagos Islands (and much credit goes to the Local Organizing Committee LOC and staff of Quito Astronomical Observatory for their logistical efforts, organizing the meeting for over a year), it was very much a successful conference. We hope it will play a small part in further developing astronomy in Ecuador.

Comparison of the Effect of Three Food Diets Provided to Giant Tortoises (Chelonoidis guntheri and Chelonoidis vicina) Bred in Captivity at Isabela Island – Galápagos

Comparison of the Effect of Three Food Diets Provided to Giant Tortoises (Chelonoidis guntheri and Chelonoidis vicina) Bred in Captivity at Isabela Island – Galápagos