Unraveling the diversification and systematic puzzle of the highly polymorphic Psammobates tentorius (Bell, 1828) complex (Reptilia: Testudinidae) through phylogenetic analyses and species delimitation approaches

The high level of phenotypic diversity in southern African tent tortoises (Psammobates tentorius complex) has for decades prevented systematists from developing a stable taxonomy for the group. Here, we used a comprehensive DNA sequence dataset (mtDNA: Cytb, ND4, ND4 adjacent tRNA-His, and tRNA-Ser, 12S, 16S; and nDNA: PRLR gene) of 455 specimens, and the latest phylogenetic and species delimitation analytical procedures, to unravel the long-standing P. tentorius complex systematic puzzle. Our results for mtDNA and nDNA were incongruent, with the poorly supported nDNA phylogeny differentiating the three recognized subspecies, and showing potential hybridization in some regions. In contrast, the concatenated mtDNA phylogeny identified seven operational taxonomic units, with strong support. Clades 1, 4, 5, and 7 corresponded to tortoises identified as P. t. tentorius, clade 3 to P. t. trimeni, and clades 2 and 6 to P. t. verroxii. Our analyses showed conflicting topologies for the placement of C6 (P. t. verroxii north of the Orange River), with stronger support for it being sister to C2 + C3 than to the other clades. Clades 1, 2, and 6 had significantly higher genetic diversity than clades 3, 4, 5, and 7, perhaps because these clades inhabit substantially larger areas. The potential for future cladogenic radiations seems high in C1 and C6, particularly in C6 for which the within-clade diversification level was highest. Further research involving microsatellite DNA, phylogeographic evaluations, and morphological variation among clades is crucial for understanding the adaptive radiation of the P. tentorius complex and for modifying their taxonomy.     

Fire Management Effects on Long-Term Gopher Tortoise Population Dynamics

Long-term datasets are required to understand the response of long-lived organisms (e.g., gopher tortoises [Gopherus polyphemus]) to management actions, such as prescribed burns. Our objective was to estimate the effects of prescribed burning on gopher tortoise population dynamics over decadal time frames at Fort Stewart Army Reserve, southeastern Georgia, USA. We captured and marked adult tortoises from 1994–2020. In addition, since the early 1990s, managers at Fort Stewart collected spatial records of prescribed burns; thus, we could compare demography of the population to prescribed burning. We used a Bayesian hierarchical model (open population Jolly-Seber model) to estimate population parameters (emigration and survival, immigration and recruitment, and adult abundance) and their relationships with years since burn. We observed opposing responses to years since burn at 2 sites: abundance and the probability of staying (survival plus not emigrating) increased within 1 site when it had been more recently burned (F zones), but abundance and probability of staying in a second site increased when it had been longer since the site was burned (E zones). Some of these effects were weak but indicative of different responses to burning between the sites. Although the sites experienced similar burning regimes, they differed substantially in other habitat features: the F zones had almost twice the tree cover and lower soil sand composition, indicating that tortoise population responses to burning depend on habitat context. We inferred that the primary mechanism for demographic responses to years since burn was likely emigrating adults, which indicates the need for more detailed movement data. Our results demonstrate that gopher tortoise population responses to prescribed burning are complex, context dependent, and primarily influenced by tortoise movements. Therefore, prescribed burn plans may best accommodate spatially dynamic tortoise populations when they create spatial heterogeneity in burn ages within the range of typical tortoise movements. © 2021 The Wildlife Society.     

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.     

Survival and movements of head-started Mojave desert tortoises

Head-starting is a conservation strategy in which young animals are protected in captivity temporarily before their release into the wild at a larger size, when their survival is presumably increased. The Mojave desert tortoise (Gopherus agassizii) is in decline, and head-starting has been identified as one of several conservation measures to assist in recovery. To evaluate the efficacy of indoor head-starting, we released and radio-tracked 68 juvenile tortoises from a 2015 cohort in the Mojave National Preserve, California, USA. We released 20 tortoises at hatching (control) in September 2015, and reared 28 indoors and 20 outdoors in predator-proof enclosures for 7 months before releasing them in April 2016. We monitored tortoises at least weekly after release until 27 October 2016, and documented survivorship, movement, and surface activity. We estimated survivorship by treatment and evaluated effects of treatment, proximity to a raven (Corvus corax) nest (predator) coincidentally established after release, distance moved between monitoring events, surface activity, and release size on individual fate in a generalized linear model. Although indoor head-start tortoises reached the size of 5–6-year-old wild tortoises by release at 7 months of age, survival did not differ significantly among the 3 treatment groups. Combined annual survival was 0.44 (95% CI = 0.34–0.58). Tortoises that were closer to an active raven nest were significantly more likely to die, as were those seen more often outside their burrows and active aboveground. Predicted estimates for short-term probability of survival approached 1.0 as distance from a raven nest exceeded approximately 1.6 km. Rearing treatment, movement distance, and body size were not significant predictors of fate over the 1-year monitoring period. Head-started tortoises released ≥1.6 km from areas of raven activity will likely have higher short-term survival. Population recovery through head-starting alone is unlikely to be successful if systemic ecosystem-level issues, such as habitat degradation and conditions that promote human-subsidized predators, are not ameliorated. © 2019 The Wildlife Society.     

Enriching Tortoises: Assessing Color Preference

Environmental enrichment is a principle that is used to enhance the quality of care for nonhuman animals in captivity. To achieve this, it is necessary to understand the animal's needs. This study focused on color preference to provide food stimuli as a source of environmental enrichment for the tortoise, Chelonoidis denticulata. During this study, the stimuli green-, blue-, yellow-, and red-colored bananas and plaster blocks were randomly offered to the tortoises. Analysis of the data showed that the tortoises had a preference for the stimuli dyed with colors red and yellow over the other presented colors. It was possible to conclude that presenting food in different colors stimulated the animals to evaluate their environment and make choices in relation to their color preference. Thus, this experiment introduced an element of choice into their lives, beyond identifying color food preferences for the tortoises. The element of choice is known to be important to animal welfare.     

Selection of large host plants for oviposition by a monophagous leaf beetle: nutritional quality or enemy‐free space?

Abstract.  1. Oviposition site selection is crucial for the reproductive success of herbivorous insects. According to the preference–performance hypothesis, females should oviposit on host plants that enhance the performance of their offspring. More specifically, the plant vigour hypothesis predicts that females should prefer large and vigorously growing host plants for oviposition and that larvae should perform best on these plants.
2. The present study examined whether females of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) prefer to oviposit on large host plant individuals of the meadow clary and whether large host plants are of higher nutritional quality than small host plants. Subsequently, it was tested whether the female preference correlates with offspring performance and survival.
3. In the field, females preferred large host plant individuals for oviposition and host plant quality, i.e. leaf nitrogen content, was significantly higher in leaves of large than of small host plants.
4. In the laboratory, larval development time was shorter on leaves of large host plant individuals than on small host plant individuals, but this could not be shown in the field.
5. However, a predator-exclusion experiment in the field resulted in a higher survival of larvae on large host plants than on small host plants when all predators had free access to the plants. On caged host plants there was no difference in survival of larvae between plant size categories.
6. It is concluded that females of C. canaliculata select oviposition sites that enhance both performance and survival of their offspring, which meets the predictions of the plant vigour hypothesis.     

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.     

Polymorphic microsatellite markers for the Mojave desert tortoise, Gopherus agassizii

We describe primers and polymerase chain reaction (PCR) conditions to amplify 14 tri- and tetranucleotide microsatellite loci for the Mojave desert tortoise (Gopherus agassizii). Across three populations (87 individuals) located in the Mojave Desert, USA, the markers yielded a range of four to 33 alleles and an average observed heterozygosity of 0.733 (range 0.433 to 0.933). We neither detected linkage disequilibrium between any pair of loci nor did we find a consistent pattern of deviation from Hardy-Weinberg equilibrium. These microsatellites are designed for PCR multiplexing, and provide higher throughput capacity to aid in conservation genetics studies for this threatened species.