Was there a second adaptive radiation of giant tortoises in the Indian Ocean? Using mitochondrial DNA to investigate speciation and biogeography of Aldabrachelys (Reptilia,Testudinidae)

A radiation of five species of giant tortoises (Cylindraspis) existed in the southwest Indian Ocean, on the Mascarene islands, and another (of Aldabrachelys) has been postulated on small islands north of Madagascar, from where at least eight nominal species have been named and up to five have been recently recognized. Of 37 specimens of Madagascan and small-island Aldabrachelys investigated by us, 23 yielded significant portions of a 428-base-pair (bp) fragment of mitochondrial (cytochrome b and tRNA-Glu), including type material of seven nominal species (A. arnoldi, A. dussumieri, A. hololissa, A. daudinii, A. sumierei, A. ponderosa and A. gouffei). These and nearly all the remaining specimens, including 15 additional captive individuals sequenced previously, show little variation. Thirty-three exhibit no differences and the remainder diverge by only 1-4 bp (0.23-0.93%). This contrasts with more widely accepted tortoise species which show much greater inter- and intraspecific differences. The non-Madagascan material examined may therefore only represent a single species and all specimens may come from Aldabra where the common haplotype is known to occur. The present study provides no evidence against the Madagascan origin for Aldabra tortoises suggested by a previous molecular phylogenetic analysis, the direction of marine currents and phylogeography of other reptiles in the area. Ancient mitochondrial DNA from the extinct subfossil A. grandidieri of Madagascar differs at 25 sites (5.8%) from all other Aldabrachelys samples examined here.     

Persistence of distinctive morphotypes in the native range of the CITES‐listed Aldabra giant tortoise

Understanding the extent of morphological variation in the wild population of Aldabra giant tortoises is important for conservation, as morphological variation in captive populations has been interpreted as evidence for lingering genes from extinct tortoise lineages. If true, this could impact reintroduction programmes in the region. The population of giant tortoises on Aldabra Atoll is subdivided and distributed around several islands. Although pronounced morphological variation was recorded in the late 1960s, it was thought to be a temporary phenomenon. Early researchers also raised concerns over the future of the population, which was perceived to have exceeded its carrying capacity. We analyzed monthly monitoring data from 12 transects spanning a recent 15‐year period (1998–2012) during which animals from four subpopulations were counted, measured, and sexed. In addition, we analyzed survival data from individuals first tagged during the early 1970s. The population is stable with no sign of significant decline. Subpopulations differ in density, but these differences are mostly due to differences in the prevailing vegetation type. However, subpopulations differ greatly in both the size of animals and the degree of sexual dimorphism. Comparisons with historical data reveal that phenotypic differences among the subpopulations of tortoises on Aldabra have been apparent for the last 50 years with no sign of diminishing. We conclude that the giant tortoise population on Aldabra is subject to varying ecological selection pressures, giving rise to stable morphotypes in discrete subpopulations. We suggest therefore that (1) the presence of morphological differences among captive Aldabra tortoises does not alone provide convincing evidence of genes from other extinct species; and (2) Aldabra serves as an important example of how conservation and management in situ can add to the scientific value of populations and perhaps enable them to better adapt to future ecological pressures.     

The Indian Ocean giant tortoise Testudo gigantea on Aldabra

The Indian Ocean giant tortoise survives on Aldabra, an atoll north-west of Madagascar. The Bristol Seychelles Expedition estimated the total numbers as 33,000 during visits in 1964 and 1965. Growth and movements were recorded during marking experiments. The growth rate with size is plotted, the time scale being derived from the number of rings on the shields of the shell; and the growth curve is compared with that for captive tortoises.
Variation in many characters of supposed taxonomic value is explained as the influence of environmental factors on the normal growth pattern, and of non-adaptive differences expressed during adult life. The present separation of north and south Aldabra, and Seychelles forms is not supported.
An explanation is suggested for the uniformity of shape among tortoises.
Observations were made on the feeding, diurnal movements and mortality of tortoises on Aldabra.     

Low‐coverage reduced representation sequencing reveals subtle within‐island genetic structure in Aldabra giant tortoises

Aldabrachelys gigantea (Aldabra giant tortoise) is one of only two giant tortoise species left in the world and survives as a single wild population of over 100,000 individuals on Aldabra Atoll, Seychelles. Despite this large current population size, the species faces an uncertain future because of its extremely restricted distribution range and high vulnerability to the projected consequences of climate change. Captive‐bred A. gigantea are increasingly used in rewilding programs across the region, where they are introduced to replace extinct giant tortoises in an attempt to functionally resurrect degraded island ecosystems. However, there has been little consideration of the current levels of genetic variation and differentiation within and among the islands on Aldabra. As previous microsatellite studies were inconclusive, we combined low‐coverage and double‐digest restriction‐associated DNA (ddRAD) sequencing to analyze samples from 33 tortoises (11 from each main island). Using 5426 variant sites within the tortoise genome, we detected patterns of within‐island population structure, but no differentiation between the islands. These unexpected results highlight the importance of using genome‐wide genetic markers to capture higher‐resolution genetic structure to inform future management plans, even in a seemingly panmictic population. We show that low‐coverage ddRAD sequencing provides an affordable alternative approach to conservation genomic projects of non‐model species with large genomes.     

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 Wilmé, 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.     

Are the native giant tortoises from the Seychelles really extinct? A genetic perspective based on mtDNA and microsatellite data

The extinction of the giant tortoises of the Seychelles Archipelago has long been suspected but is not beyond doubt. A recent morphological study of the giant tortoises of the western Indian Ocean concluded that specimens of two native Seychelles species survive in captivity today alongside giant tortoises of Aldabra, which are numerous in zoos as well as in the wild. This claim has been controversial because some of the morphological characters used to identify these species, several measures of carapace morphology, are reputed to be quite sensitive to captive conditions. Nonetheless, the potential survival of giant tortoise species previously thought extinct presents an exciting scenario for conservation. We used mitochondrial DNA sequences and nuclear microsatellites to examine the validity of the rediscovered species of Seychelles giant tortoises. Our results indicate that the morphotypes suspected to represent Seychelles species do not show levels of variation and genetic structuring consistent with long periods of reproductive isolation. We found no variation in the mitochondrial control region among 55 individuals examined and no genetic structuring in eight microsatellite loci, pointing to the survival of just a single lineage of Indian Ocean tortoises.     

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.     

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.     

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.     

Seed dispersal by Galápagos tortoises

Aim Large‐bodied vertebrates often have a dramatic role in ecosystem function through herbivory, trampling, seed dispersal and nutrient cycling. The iconic Galápagos tortoises (Chelonoidis nigra) are the largest extant terrestrial ectotherms, yet their ecology is poorly known. Large body size should confer a generalist diet, benign digestive processes and long‐distance ranging ability, rendering giant tortoises adept seed dispersers. We sought to determine the extent of seed dispersal by Galápagos tortoises and their impact on seed germination for selected species, and to assess potential impacts of tortoise dispersal on the vegetation dynamics of the Galápagos. Location Galápagos, Ecuador. Methods To determine the number of seeds dispersed we identified and counted intact seeds from 120 fresh dung piles in both agricultural and national park land. To estimate the distance over which tortoises move seeds we used estimated digesta retention times from captive tortoises as a proxy for retention times of wild tortoises and tortoise movement data obtained from GPS telemetry. We conducted germination trials for five plant species to determine whether tortoise processing influenced germination success. Results In our dung sample, we found intact seeds from > 45 plant species, of which 11 were from introduced species. Tortoises defecated, on average, 464 (SE 95) seeds and 2.8 (SE 0.2) species per dung pile. Seed numbers were dominated by introduced species, particularly in agricultural land. Tortoises frequently moved seeds over long distances; during mean digesta retention times (12 days) tortoises moved an average of 394 m (SE 34) and a maximum of 4355 m over the longest recorded retention time (28 days). We did not find evidence that tortoise ingestion or the presence of dung influenced seed germination success. Main conclusions Galápagos tortoises are prodigious seed dispersers, regularly moving large quantities of seeds over long distances. This may confer important advantages to tortoise‐dispersed species, including transport of seeds away from the parent plants into sites favourable for germination. More extensive research is needed to quantify germination success, recruitment to adulthood and demography of plants under natural conditions, with and without tortoise dispersal, to determine the seed dispersal effectiveness of Galápagos tortoises.     

Desert Tortoises (Gopherus agassizii) Are Selective Herbivores that Track the Flowering Phenology of Their Preferred Food Plants

Previous studies of desert tortoise foraging ecology in the western Mojave Desert suggest that these animals are selective herbivores, which alter their diet according to the temporal availability of preferred food plants. These studies, however, did not estimate availability of potential food plants by taking into account the spatial and temporal variability in ephemeral plant abundance that occurs within the spring season. In this study, we observed 18 free-ranging adult tortoises take 35,388 bites during the spring foraging season. We also estimated the relative abundance of potential food plants by stratifying our sampling across different phenological periods of the 3-month long spring season and by different habitats and microhabitats. This methodology allowed us to conduct statistical tests comparing tortoise diet against plant abundance. Our results show that tortoises choose food plants non-randomly throughout the foraging season, a finding that corroborates the hypothesis that desert tortoises rely on key plants during different phenological periods of spring. Moreover, tortoises only consumed plants in a succulent state until the last few weeks of spring, at which time most annuals and herbaceous perennials had dried and most tortoises had ceased foraging. Many species of food plants—including several frequently eaten species—were not detected in our plant surveys, yet tortoises located these rare plants in their home ranges. Over 50% of bites consumed were in the group of undetected species. Interestingly, tortoises focused heavily on several leguminous species, which could be nutritious foods owing to their presumably high nitrogen contents. We suggest that herbaceous perennials, which were rare on our study area but represented ~30% of tortoise diet, may be important in sustaining tortoise populations during droughts when native annuals are absent. These findings highlight the vulnerability of desert tortoises to climate change if such changes alter the availability of their preferred food plants.     

The evolutionary origin of Indian Ocean tortoises (Dipsochelys)

Today, the only surviving wild population of giant tortoises in the Indian Ocean occurs on the island of Aldabra. However, giant tortoises once inhabited islands throughout the western Indian Ocean. Madagascar, Africa, and India have all been suggested as possible sources of colonization for these islands. To address the origin of Indian Ocean tortoises (Dipsochelys, formerly Geochelone gigantea), we sequenced the 12S, 16S, and cyt b genes of the mitochondrial DNA. Our phylogenetic analysis shows Dipsochelys to be embedded within the Malagasy lineage, providing evidence that Indian Ocean giant tortoises are derived from a common Malagasy ancestor. This result points to Madagascar as the source of colonization for western Indian Ocean islands by giant tortoises. Tortoises are known to survive long oceanic voyages by floating with ocean currents, and thus, currents flowing northward towards the Aldabra archipelago from the east coast of Madagascar would have provided means for the colonization of western Indian Ocean islands. Additionally, we found an accelerated rate of sequence evolution in the two Malagasy Pyxis species examined. This finding supports previous theories that shorter generation time and smaller body size are related to an increase in mitochondrial DNA substitution rate in vertebrates.     

Resurrecting extinct interactions with extant substitutes

There is increasing evidence that restoration ecologists should be most concerned with restoring species interactions rather than species diversity per se [1]. Rewilding with taxon substitutes, the intentional introduction of exotic species to replace the ecosystem functions of recently extinct species, is one way to reverse ecosystem dysfunction following the loss of species interactions [2]. This is highly controversial [3], in part because of a lack of rigorous scientific studies [4]. Here we present the first empirical evidence of an in situ rewilding project undertaken as a hypothesis-driven ecosystem management option. On Ile aux Aigrettes, a 25-hectare island off Mauritius, the critically endangered large-fruited endemic ebony, Diospyros egrettarum (Ebenaceae), was seed-dispersal limited after the extinction of all native large-bodied frugivores, including giant tortoises. We introduced exotic Aldabra giant tortoises, Aldabrachelys gigantea, to disperse the ebony seeds. Not only did the tortoises ingest the large fruits and disperse substantial numbers of ebony seeds, but tortoise gut passage also improved seed germination, leading to the widespread, successful establishment of new ebony seedlings. Our results demonstrate that the introduction of these exotic frugivores is aiding the recovery of ebonies. We argue for more reversible rewilding experiments to investigate whether extinct species interactions can be restored.     

Movement patterns and habitat use of soft-released translocated spur-thighed tortoises, <Emphasis Type="Italic">Testudo graeca</Emphasis>

Wildlife translocations, the deliberate movement of animals from one part of their distribution to another, are increasingly used as a conservation method for the reestablishment of rare and endangered species. The objective of this study was to examine the movement patterns and macro- and microhabitat use of translocated and resident spur-thighed tortoises. This translocation was considered a soft-release as the tortoises were forced to be relatively inactive due to their being released at the beginning of the aestivation season. Our results suggest that forced aestivation soft-releases may succeed in reducing dispersal by forcing spur-thighed tortoises to spend time at the release site as the majority of translocated tortoises had similar activity range sizes and movement path tortuosity as resident tortoises. Spur-thighed tortoise conservation will require protecting habitat at multiple scales, with the remaining native forests in the country of Jordan being important to the spur-thighed tortoise during the activity and aestivation/hibernation seasons, as this macrohabitat was used significantly more than the human-modified habitats. Microhabitat structures such as leaf litter and availability of large stones may also be especially important in human-modified landscapes, as these microhabitats may help reduce the effects of habitat degradation.     

The Dominance of Introduced Plant Species in the Diets of Migratory Galapagos Tortoises Increases with Elevation on a Human‐Occupied Island

The distribution of resources and food selection are fundamental to the ecology, life history, physiology, population dynamics, and conservation of animals. Introduced plants are changing foraging dynamics of herbivores in many ecosystems often with unknown consequences. Galapagos tortoises, like many herbivores, undertake migrations along elevation gradients driven by variability in vegetation productivity which take them into upland areas dominated by introduced plants. We sought to characterize diet composition of two species of Galapagos tortoises, focussing on how the role of introduced forage species changes over space and the implications for tortoise conservation. We quantified the distribution of tortoises with elevation using GPS telemetry. Along the elevation gradient, we quantified the abundance of introduced and native plant species, estimated diet composition by recording foods consumed by tortoises, and assessed tortoise physical condition from body weights and blood parameter values. Tortoises ranged between 0 and 429 m in elevation over which they consumed at least 64 plant species from 26 families, 44 percent of which were introduced species. Cover of introduced species and the proportion of introduced species in tortoise diets increased with elevation. Introduced species were positively selected for by tortoises at all elevations. Tortoise physical condition was either consistent or increased with elevation at the least biologically productive season on Galapagos. Santa Cruz tortoises are generalist herbivores that have adapted their feeding behavior to consume many introduced plant species that has likely made a positive contribution to tortoise nutrition. Some transformed habitats that contain an abundance of introduced forage species are compatible with tortoise conservation.     

The rise and fall of the Aldabran giant tortoise population.

At the end of the 19th century, after prolonged and extensive harvesting, indigenous giant tortoises had been eliminated from all islands in the Indian Ocean, except Aldabra atoll, where only a few survived. With greatly reduced levels of exploitation during the 20th century, the population recovered to a revised estimated total of 129,000 in 1973-1974, when the first sample census was conducted. A repeat census in 1997 revealed a highly significant reduction in numbers over the past 24 years to an estimated total of 100,000. The great majority of tortoises are still found at relatively high density in south-eastern Grande Terre, where the number of animals has declined by more than one-third. In contrast, low-density subpopulations on Malabar and Picard have almost doubled in size, but they represent less than 5% of the total population. Corroborative evidence for the crash in the Grande Terre subpopulation comes from two independent observations: a significant increase in tortoise mortality; and a significant decline in tortoise counts on long-term population monitoring transects. These population changes are attributed to natural population regulatory mechanisms, exacerbated by low rainfall years in the period 1980-1997, including two consecutive years of below average rainfall in 1995-1996 and 1996-1997.     

Tick prevalence and species diversity on Aldabran giant tortoises (Dipsochelys dussumieri) in relation to host range and host size in a restored ecosystem, Kenya

Tick species density and diversity on Aldabran tortoises was investigated in relation to the habitat range and size of each tortoise. Identification of tick infestation patterns forms an important aspect of effective tick control. Ten Aldabran tortoises were de‐ticked and monitored over the course of 2 months. Tick species found were Amblyomma sparsum, Amblyomma nuttalli, Amblyomma hebraeum and Boophilus decoloratus, with the most prevalent species found being A. sparsum. Tick loads varied considerably from 20 to 214 ticks per tortoise, with most ticks collected from the head/neck region. Tortoises ranging outside Haller Park had higher tick loads (70–214) compared with tortoises ranging within Haller Park (20–99). Tick load was not correlated with tortoise size. Results indicate that tick loads are related to the habitat range of the tortoises and may indirectly also be related to food preference and host food availability. Implications of the findings and appropriate tick control measures are discussed.     

Digesta retention time in the Galápagos tortoise (Chelonoidis nigra)

The retention time of food in the digestive tract of animals has important implications for digestive physiology. Retention time impacts digestive efficiency and among herbivores affects plant–animal interactions including herbivory and seed dispersal. Poorly studied yet iconic Galápagos tortoises are large-bodied generalist herbivores and ecosystem engineers which migrate seasonally. Potentially variable digesta retention times due to strong seasonal and altitudinal temperature gradients may influence tortoise seed dispersal abilities and rates of herbivory. We fed captive adult tortoises living in semi-natural conditions on Galápagos with inert particles and seeds from locally available fruits to determine whether seed size and ambient temperature influenced retention time. Median retention time varied from 6 to 28 days, with a mode of 12 days. Seed size had no effect on any of our measures of retention time, but ambient temperature was inversely correlated with retention times. Long retention time facilitates long distance seed dispersal by Galápagos tortoises, which may improve effectiveness. The effect of temperature, which may double from hot lowlands to cold highlands through the seasonal cycle, on tortoise digesta retention time will strongly influence seed dispersal efficiency and may influence patterns of food selection and migration in this species.     

Using motion-sensor camera technology to infer seasonal activity and thermal niche of the desert tortoise (Gopherus agassizii)

Understanding the relationships between environmental variables and wildlife activity is an important part of effective management. The desert tortoise (Gopherus agassizii), an imperiled species of arid environments in the southwest US, may have increasingly restricted windows for activity due to current warming trends. In summer 2013, we deployed 48 motion sensor cameras at the entrances of tortoise burrows to investigate the effects of temperature, sex, and day of the year on the activity of desert tortoises. Using generalized estimating equations, we found that the relative probability of activity was associated with temperature (linear and quadratic), sex, and day of the year. Sex effects showed that male tortoises are generally more active than female tortoises. Temperature had a quadratic effect, indicating that tortoise activity was heightened at a range of temperatures. In addition, we found significant support for interactions between sex and day of the year, and sex and temperature as predictors of the probability of activity. Using our models, we were able to estimate air temperatures and times (days and hours) that were associated with maximum activity during the study. Because tortoise activity is constrained by environmental conditions such as temperature, it is increasingly vital to conduct studies on how tortoises vary their activity throughout the Sonoran Desert to better understand the effects of a changing climate.     

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.