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.     

Hemoglobin-sulfhydryls from tortoise (Geochelone carbonaria) can reduce oxidative damage induced by organic hydroperoxide in erythrocyte membrane

Sulfhydryl groups are important to avoid oxidative damage to the cell. In RBC, tert-butyl hydroperoxide (tert-BOOH) and hydrogen peroxide (H₂O₂) are capable of oxidizing heme and promoting lipid peroxidation. H₂O₂ caused greater oxidation of heme than tert-BOOH, although the oxidation of sulfhydryl groups was similar. Geochelone carbonaria Hb, a rich sulfhydryl protein, inhibited the TBA-reactive substances formation of human erythrocytes exposed to tert-BOOH by about 30%; this decrease was smaller with Geochelone denticulata Hb. Sulfhydryl reagents diminished the number of reactive sulfhydryl groups in the G. carbonaria Hb resulting in a decrease of its antioxidant power, suggesting the involvement of sulfhydryls of Hb in the protection against lipid peroxidation.     

A role for histamine in cardiovascular regulation in late stage embryos of the red-footed tortoise, Chelonoidis carbonaria Spix, 1824

A chorioallantoic membrane artery in embryos of the red-footed tortoise, Chelonoidis carbonaria was occlusively cannulated for measurement of blood pressure and injection of drugs. Two age groups of embryos in the final 10 % of incubation were categorized by the ratio of embryonic body to yolk mass. All embryos first received cholinergic and β-adrenergic blockade. This revealed that β-adrenergic control was established in both groups whereas cholinergic control was only established in the older group immediately prior to hatching. The study then progressed as two series. Series one was conducted in a subset of embryos treated with histamine before or after injection of ranitidine, the antagonist of H₂ receptors. Injection of histamine caused an initial phasic hypertension which recovered, followed by a longer lasting hypertensive response accompanied by a tachycardia. Injection of the H₂ receptor antagonist ranitidine itself caused a hypotensive tachycardia with subsequent recovery of heart rate. Ranitidine also abolished the cardiac effects of histamine injection while leaving the initial hypertensive response intact. In series, two embryos were injected with histamine after injection of diphenhydramine, the antagonist to H₁ receptors. This abolished the whole of the pressor response to histamine injection but left the tachycardic response intact. These data indicate that histamine acts as a non-adrenergic, non-cholinergic factor, regulating the cardiovascular system of developing reptilian embryos and that its overall effects are mediated via both H₁ and H₂ receptor types.     

Flexibility of digestive responses in two generalist herbivores,the tortoises Geochelone carbonaria and Geochelone denticulata

Summary To test the prediction that digestive responses digestibility, intake and passage time-of generalist herbivores vary with different diets, feeding trials were conducted in Venezuela with two sympatric tortoise species, Geochelone carbonaria and G. denticulata. Three single-species diets (two fruit, one foliage) were fed to both species. For a given diet, digestibility, mass-specific intake and passage time did not differ between the two tortoise species, nor did they vary by sex or body mass within each species. However, the digestive parameters varied for tortoises feeding on the different diets. The responses ranged from nearly abandoning cell wall fermentation and depending entirely on extraction of cell contents to relying heavily on cell wall fermentation. Therefore, these generalist herbivores have flexible digestive responses that are influenced by diet, not fixed digestive responses that limit the diet, as previously observed in other generalist herbivores. A three-part classification of herbivores (specialist, specialized mixed feeder and opportunistic mixed feeder) is suggested as an approach to understanding flexible and inflexible digestive strategies in herbivores.     

Molecular characterization of hemoglobin alpha-D chains from Geochelone carbonaria and Geochelone denticulata land turtles

In order to help elucidate the evolution of alpha-globins, the complete cDNA and amino acid sequences of Geochelone carbonaria and Geochelone denticulata land turtles alpha-D chains have been described. In G. carbonaria, the cDNA is 539 bp with ATG start codon located at position 46, TGA stop codon at position 469 and AATAAA polyadenylation signal at position 520. In G. denticulata, the cDNA is 536 bp with ATG start codon located at position 46, TGA stop codon at position 469 and AATAAA polyadenylation signal at position 517. Both cDNAs codify 141 amino acid residues, differing from each other in only four amino acid residues. When comparing with human Hb alpha-chain, alterations in important regions can be noted: alpha110 Ala-Gly, alpha114 Pro-Gly, alpha117 Phe-Tyr and alpha122 His-Gln. There is a high homology between the amino acids of these turtles when compared with chicken alpha-D chains, progressively decreasing when compared with human, crocodile, snake, frog and fish alpha-chains. Phylogenetic analysis of alpha-D chains shows that those of turtles are closer to those of birds than to snakes and lizards.     

Computer animations stimulate contagious yawning in chimpanzees

People empathize with fictional displays of behaviour, including those of cartoons and computer animations, even though the stimuli are obviously artificial. However, the extent to which other animals also may respond empathetically to animations has yet to be determined. Animations provide a potentially useful tool for exploring non-human behaviour, cognition and empathy because computer-generated stimuli offer complete control over variables and the ability to program stimuli that could not be captured on video. Establishing computer animations as a viable tool requires that non-human subjects identify with and respond to animations in a way similar to the way they do to images of actual conspecifics. Contagious yawning has been linked to empathy and poses a good test of involuntary identification and motor mimicry. We presented 24 chimpanzees with three-dimensional computer-animated chimpanzees yawning or displaying control mouth movements. The apes yawned significantly more in response to the yawn animations than to the controls, implying identification with the animations. These results support the phenomenon of contagious yawning in chimpanzees and suggest an empathic response to animations. Understanding how chimpanzees connect with animations, to both empathize and imitate, may help us to understand how humans do the same.     

Pyrogens fail to produce fever in the leopard tortoise Geochelone pardalis

1. The body temperature of seven tortoises, Geochelone pardalis, was measured in a thermal gradient chamber, by indwelling thermocouples, after injection of various pyrogens. 2. The tortoises regulated their body temperature by moving in the chamber. 3. The tortoises did not develop fever in response to any of the pyrogens we tested. 4. The results support the contention that fever in reptiles is not ubiquitous.     

Absence of contagious yawning in children with autism spectrum disorder

This study is the first to report the disturbance of contagious yawning in individuals with autism spectrum disorder (ASD). Twenty-four children with ASD as well as 25 age-matched typically developing (TD) children observed video clips of either yawning or control mouth movements. Yawning video clips elicited more yawns in TD children than in children with ASD, but the frequency of yawns did not differ between groups when they observed control video clips. Moreover, TD children yawned more during or after the yawn video clips than the control video clips, but the type of video clips did not affect the amount of yawning in children with ASD. Current results suggest that contagious yawning is impaired in ASD, which may relate to their impairment in empathy. It supports the claim that contagious yawning is based on the capacity for empathy.     

Ingroup-outgroup bias in contagious yawning by chimpanzees supports link to empathy

Humans favor others seen as similar to themselves (ingroup) over people seen as different (outgroup), even without explicitly stated bias. Ingroup-outgroup bias extends to involuntary responses, such as empathy for pain. However, empathy biases have not been tested in our close primate relatives. Contagious yawning has been theoretically and empirically linked to empathy. If empathy underlies contagious yawning, we predict that subjects should show an ingroup-outgroup bias by yawning more in response to watching ingroup members yawn than outgroup. Twenty-three chimpanzees (Pan troglodytes) from two separate groups watched videos of familiar and unfamiliar individuals yawning or at rest (control). The chimpanzees yawned more when watching the familiar yawns than the familiar control or the unfamiliar yawns, demonstrating an ingroup-outgroup bias in contagious yawning. These results provide further empirical support that contagious yawning is a measure of empathy, which may be useful for evolutionary biology and mental health.     

Plant dispersal by the Aldabran giant tortoise,Geochelone gigantea (Schweigger)

Summary A total of 28 species of grasses, herbs, and woody plants germinated from faeces of Aldabran giant tortoises. It seems likely that all members of the land flora can be ingested by tortoises and so may be dispersed by them over short distances.The tortoises are thought to have derived from a Madagascan stock and reached Aldabra from time to time by transmarine migration until the extermination of the Madagascan tortoises about 1100 years ago. The time taken by a tortoise to void its last meal is much longer than that required to float, at present current speeds, from Madagascar to Aldabra. If neither of these parameters have changed since the tortoise migrations to Aldabra, the tortoises may have been responsible for the introduction of plant species to Aldabra.     

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.     

Attachment sites of the tick Amblyomma marmoreum on its tortoise host,Geochelone pardalis

Leopard tortoises (n=162) were collected over a period of 19 months at the National Zoological gardens, Pretoria, South Africa and the attachment sites of larvae, nymphs and adults of A. marmoreum monitored. Most of the larvae (80%) and nymphs (87%) were found on the anterior parts of the tortoise while adults (90%) were found mainly on the posterior regions of the host body. The immature stages were predominant in soft skinned areas protected by the carapace especially the neck and upper leg areas. Male and female ticks were most commonly found on the posterior armpit. posterior legs and the areas surrounding the base of the tail. Interstadial competition as a major factor influencing attachment site segregation is doubtful. Anterior attachment site selection of the immature stages is believed to result largely from ticks lying passively in wait and then climbing onto the host as it moves through the vegetation and the requirement for soft skinned and protected areas to allow for blood feeding. Adult females usually attach posteriorly in areas that allow them to fully engorge without being dislodged by mechanical disturbance, while males prefer to attach at sites where they are protected but still remain in close proximity to female pheromone signals.     

The primary structure of hemoglobin D from the Aldabra giant tortoise,Geochelone gigantea

The complete primary structures of alpha D-2- and beta-globin of hemoglobin D (Hb D) from the Aldabra giant tortoise, Geochelone gigantea, have been constructed by amino acid sequencing analysis in assistance with nucleotide sequencing analysis of PCR fragments amplified using degenerate oligonucleotide primers. Using computer-assisted sequence comparisons, the alpha D-2-globin shared a 92.0% sequence identity versus alpha D-globin of Geochelone carbonaria, a 75.2% versus alpha D-globin of Aves (Rhea americana) and a 62.4% versus alpha A-globin of Hb A expressed in adult red blood cells of Geochelone gigantea. Additionally, judging from their primary structures, an identical beta-globin was common to the two hemoglobin components, Hb A and Hb D. The alpha D-2- and beta-globin genes contained the three-exon and two-intron configurations and showed the characteristic of all functional vertebrate hemoglobin genes except an abnormal GC dinucleotide instead of the invariant GT at the 5' end of the second intron sequence. The introns of alpha D-2-globin gene were both small (224-bp/first intron, 227-bp/second intron) such that they were quite similar to those of adult alpha-type globins; the beta-globin gene has one small intron (approximately 130-bp) and one large intron (approximately 1590-bp). A phylogenetic tree constructed on primary structures of 7 alpha D-globins from Reptilia (4 species of turtles, 2 species of squamates, and 1 species of sphenodontids) and two embryonic alpha-like globins from Aves (Gullus gullus) and Mammals (Homo sapiens) showed the following results: (1) alpha D-globins except those of squamates were clustered, in which Sphenodon punctatus was a closer species to birds than turtles; (2) separation of the alpha A- and alpha D-globin genes occurred approximately 250 million years ago after the embryonic alpha-type globin-genes (pi' and zeta) first split off from the ancestor of alpha-type globin gene family.     

The daily time course of contagious and spontaneous yawning among humans

Yawning, besides being a spontaneous behavior, can also be evoked by observing others yawn. However, contagious yawning does not always occur, depending possibly on several factors, such as one’s propensity to spontaneously yawn and a heightened level of sleepiness. The aim of this study is to investigate in young adults whether contagious yawning frequency varies throughout the day, and if it is related to the daily time course of spontaneous yawning frequency and level of sleepiness. For the study, 22 subjects were instructed to log hourly, throughout wakefulness, the number of spontaneous yawns and sleepiness level. Subjects were required to continue this procedure for 3 consecutive days, after which they underwent five experimental sessions occurring at regular intervals throughout the same day. During each experimental session, subjects observed others yawn (stimulus condition) and smile (control condition). Our findings show that the contagious effect of yawning is always present throughout the daytime. Both contagious and spontaneous yawning peaked in the early morning and in the late evening, according to the sleepiness time course. However, the frequency of spontaneous yawns was remarkably lower than the frequency of contagious yawns around 19:00. This difference suggests that different mechanisms control spontaneous and contagious yawning.     

Seasonal reproductive cycle of the Galápagos tortoise (Geochelone nigra) in captivity

The reproductive physiology of nine Galápagos tortoises (Geochelone nigra) was studied from February 1988 to May 1989. The study encompassed the annual reproductive cycle to include complete mating and nesting sequences. Male (n = 4) and female (n = 5) seasonal reproductive changes were determined throughout the study with endocrine analysis and ultrasonographic examinations. Males displayed a prenuptial rise in serum testosterone (x― ±SE = 6.62 ± 0.92 ng/ml in August) during which gonadal maturation and spermatogenesis are thought to occur. The male reproductive cycle appears consistent with the prenuptial spermatogenic pattern exhibited by other tropical turtles. In the females, testosterone rose during the mating period (x― ± SE = 499.3 ± 124.6 pg/ml in October) prior to ovulation and is probably related to receptivity in the females. Progesterone was more variable, but also peaked during the mating period (x― ± SE = 1,017.2 ± 220.6 pg/ml in October) and appears related to ovulation. Estradiol rose several months prior to mating (x― ± SE = 75.5 ± 11.9 pg/ml in July) and was correlated with increased serum calcium levels. This increase in estradiol is thought to stimulate vitellogenesis several months prior to mating. Nesting occurred from November 1988 to April 1989, during which six clutches were laid. Clutch size ranged from eight to 17 eggs. Both male and female Galápagos tortoises display seasonal physiological changes that function to regulate annual reproductive patterns. Zoo Biol 17:505–517, 1998. © 1998 Wiley-Liss, Inc.     

Sighting frequency and food habits of the leopard tortoise, Geochelone pardalis, in northern Tanzania

Sighting frequency and food habits of the leopard tortoise (Geochelone pardalis) were studied in northern Tanzania from October 1993 to June 1996. Sighting frequency varied significantly between protected (0.22 mhr^?1) and unprotected sites (0.59 mhr^?1), and between Arusha (0.27 mhr^?1) and Serengeti sites (0.56 mhr^?1). The tortoise diet comprised mostly plants (97.8%) and rarely inorganic matter (2.2%). A total of 47 plant species from 21 families was eaten. Forbs made up 74.5% of the plant items and monocots the remainder. Succulents contributed 51.0% of the plant items eaten while grasses and legumes accounted for 16.8% and 13.5%, respectively. Some plant items were eaten more frequently than their occurrence in the habitats.     

Growth and carapacial colour variation of the leopard tortoise, Geochelone pardalis babcocki, in northern Tanzania

Observations were made of growth rate and variation in colour pattern of the leopard tortoise ( Geochelone pardalis babcocki ) in northern Tanzania between October 1993 and June 1996. Growth rate differed significantly between tortoises, with immature animals (6.9 ± 1.4 mm month⁻¹) and females (5.7 ± 1.3 mm month⁻¹) exhibiting higher rates of growth than adult animals (2.9 ± 0.8 mm month⁻¹) and males (3.2 ± 1.1 mm month⁻¹), respectively. The faster growth rate in immature animals suggested that growth rate declined with age. The difference in growth rates between the sexes was ascribed to sexual dimorphism, adult females being larger and therefore growing faster than males.
The carapacial colour pattern observed in the northern Tanzanian tortoises was generally similar to that described elsewhere in the species' range. The colour pattern varied between Arusha and Serengeti District tortoises. Animals at the Arusha sites tended to be sparsely coloured (55.6%) compared to the Serengeti tortoises, which had a buff colouration with radiating spots (89.3%).; The sparse colouration provides effective camouflage in the drier Arusha habitats, as does the buff colouration with black spots in the wetter, darker habitats in the Serengeti.     

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.     

When genes meet nomenclature: tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone

We used a five-gene data set (mtDNA: 12S rRNA, 16S rRNA, cyt-b; nDNA: Cmos, Rag2) comprising approximately two-thirds of all extant testudinid species and, for the first time, including all five Testudo species to investigate the question of whether all western Palaearctic testudinids are monophyletic. Further, we examined whether the recently suggested allocation of the African Geochelone pardalis in the otherwise exclusively South African genus Psammobates and of the Malagasy G. yniphora in the monotypic genus Angonoka is justified in the face of considerable morphological evidence against such placements. Our phylogenetic analyses do not support the paraphyly and generic break-up of Testudo, as suggested by previous papers using a smaller taxon sampling and mtDNA data only. We propose a continued usage of the generic name Testudo for all five western Palaearctic tortoise species. Within Testudo, two monophyletic subclades are present, one containing T. hermanni+T. horsfieldii, and the other comprising (T. kleinmanni+T. marginata)+T. graeca. Nomenclaturally, we demonstrate that Eurotestudo Lapparent de Broin et al., 2006, which was recently erected with the type species T. hermanni, is an objective junior synonym of Chersine Merrem, 1820 and Medaestia Wussow, 1916. Recognition of a monotypic genus Angonoka for G. yniphora is unwarranted according to both our re-analysis of sequence data and morphological data. Acknowledging the strong morphological similarity between G. yniphora and G. radiata, we suggest placing both species into the genus Astrochelys. Although sequence data for only one of the three Psammobates species was available for analysis, there is currently no cause to challenge the monophyly of this genus as established on the basis of morphological evidence. Thus, we hypothesize that G. pardalis is sister to a monophyletic Psammobates. In light of the clear morphological gap between G. pardalis and Psammobates species, the recognition of a distinct genus Stigmochelys for the former seems justified.