Research Article
Grouping behaviour, tail-biting behaviour and sexual dimorphism in the armadillo lizard (Cordylus cataphractus) from South Africa
- P. le F. N. Mouton, A. F. Flemming, E. M. Kanga
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- Published online by Cambridge University Press:
- 01 September 1999, pp. 1-10
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Grouping behaviour, tail-biting behaviour and sexual dimorphism were investigated in Cordylus cataphractus, a rock-dwelling lizard from South Africa. Eighty-five per cent of individuals collected during different seasons occurred in groups of two or more, which shows that grouping behaviour is a year-round phenomenon in this species. Group size ranged from one to approximately 30. Groups smaller than nine individuals normally included only one adult male, but larger groups often had more than one. Sixty-eight per cent of lone individuals were adult males. All individuals (n = 134) attempted to bite their tails and roll into a ball on collection. Cordylus cataphractus prefers horizontal crevices with an accompanying ledge on which animals can perch in close proximity to the crevice. Males apparently reach larger asymptotic body sizes than females and also have larger heads and longer tails. The higher incidence of scars among males suggests that sexual dimorphism could result from sexual selection, but sexual differences in scaling of head and tail dimensions, however, suggest that the dimorphism may be the result of differential energy allocation by females. Unlike most other Cordylus species which have female-biased sex ratios, the ratio is close to one in C. cataphractus.
Field body temperatures, activity levels and opportunities for thermoregulation in an extreme microhabitat specialist, the girdled lizard (Cordylus macropholis)
- Dirk Bauwens, Aurora M. Castilla, P. le Fras N. Mouton
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- 01 September 1999, pp. 11-18
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The girdled lizard Cordylus macropholis inhabits semi-desert areas along the west coast of South Africa. Unlike many other small diurnal lizards living in open habitats, it exhibits extreme low levels of above-ground activity and does not show overt thermoregulatory behaviours, such as basking or shuttling. It has an extreme low movement rate and confines routine activity to the microhabitat of the succulent Euphorbia caput-medusae. We document this restriction of activity in time and space in relation to aspects of this lizard's thermal biology. Throughout the day, a majority of the lizards was found sheltering (‘covered’) among the branches of E. caput-medusae plants, and surface-active (‘exposed’) individuals were seen during only a restricted time period (i.e. 1100–1700 h). Mean body temperatures (Tbs) of both ‘exposed’ (29.4 °C) and ‘covered’ (28.4 °C) individuals were surprisingly low for a lizard inhabiting a hot and dry climate zone. This similarity of the Tbs of both groups, as well as the correspondence between the ambient temperatures in the plant microhabitat and the Tbs of ‘exposed’ lizards indicate that, for a considerable part of day, thermal conditions under plant cover allow achievement of Tbs at a similar level as that maintained by the ‘exposed’ lizards. Moreover, the absence of sightings of overtly basking lizards supports the view that C. macropholis warm up by adopting the ambient temperature in their shelters thereby reducing the time spent in surface activity. All lizards observed were within the periphery of E. caput-medusae plants, indicating that C. macropholis is an extreme microhabitat specialist. The succulent provides more and safer hiding places than other common shrubs in the environment. In addition, estimates of ambient temperatures indicate that these plants offer superior opportunities for thermoregulation compared with other available shrub microhabitats.
Morphology, evolution and function of feeding in the giant anteater (Myrmecophaga tridactyla)
- Virginia L. Naples
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- 01 September 1999, pp. 19-41
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The edentulous giant anteater (Myrmecophaga tridactyla) ingests food using a slender, elongated sticky tongue which can project to a distance greater than the cranial length. A large and elongated hyoid apparatus, including a long stylohyal, epihyal, reduced ceratohyal and fused basihyal-thyrohyal fused to a partially ossified thyroid cartilage, supports the tongue. The fusion pattern and relative hyoid element sizes in adult Myrmecophaga differ from those in other xenarthran anteaters, tree and ground sloths, and armadillos. The hyoid bones have synovial joints with articular surfaces permitting great freedom of movement. A unique hyoid muscle arrangement enables Myrmecophaga to project the tongue with great speed and precise positional control. This muscle arrangement combined with an elongated secondary palate, accommodates the retracted tongue within the oropharynx without compromising the animal's ability to breathe. Maximum gape is reached at a few degrees of mandibular depression, but the oral opening is increased to 2 cm by the extreme length of the anterior facial region. Gape is further increased by medial rotation and depression of the unfused mandibular rami at the mental symphysis. This movement, even without mandibular depression, permits protraction and retraction of the elongated tongue. Mandibular rotation in lieu of depression and elevation simplifies jaw movements made by a smaller uniquely modified muscle mass over shorter distances, therefore increasing the speed with which anteaters can ingest food, and complements the extremely rapid tongue protrusion–retraction cycle.
Identification of newts (Triturus sp.) and their hybrids using molecular methods
- T. J. C. Beebee, G. Rowe, J. W. Arntzen
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- 01 September 1999, pp. 43-47
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Identification of hybrids is important for the study of introgression between related species, but can be difficult when morphological differences between parental species are small. In the case of the newts Triturus vulgaris and T. helveticus, males are readily identifiable during the breeding season when secondary sexual characters are highly developed, but female dimorphism is always much less. Male hybrids with intermediate characters are therefore more likely to be detected than female ones. RAPD (random amplified polymorphic DNA) analysis of three putative newt hybrids and examples of the two parent species T. vulgaris and T. helveticus confirmed hybrid status in all three cases. The maternal parents of each hybrid were identified by amplification of mitochondrial cytochrome b and ATPase genes, including partial sequencing of the latter. One hybrid had a T. vulgaris mother while the other two had T. helveticus mothers. The PCR-based methods therefore provided a quick and simple approach for identifying both hybrid status and parentage in crosses between these two urodele species. The RAPD technique should also prove useful for the identification of larvae, which are morphologically indistinguishable in T. vulgaris and T. helveticus.
A phylogeny of the European lizard genus Algyroides (Reptilia: Lacertidae) based on DNA sequences, with comments on the evolution of the group
- D. James Harris, E. Nicholas Arnold, Richard H. Thomas
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- 01 September 1999, pp. 49-60
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The four species of Algyroides Bibron & Bory, 1833 form part of the relatively plesiomorphic Palaearctic clade of lacertids comprising Lacerta and its allies. An estimate of phylogeny based on DNA sequence from parts of the 12S and 16S rRNA mitochondrial genes confirms the monophyly of the genus already suggested by several morphological features. The molecular data also indicates that relationships within the clade are: (A. nigropunctatus (A. moreoticus (A. fitzingeri, A. marchi))); this agrees with an estimate of phylogeny based on morphology that assumes the taxon ancestral to Alygroides was relatively robust in body form, and not strongly adapted to using crevices. Initial morphological evolution within Algyroides appears to involve adaptation to crypsis in woodland habitats. The most plesiomorphic form (A. nigropunctatus) is likely to have originally climbed extensively on tree boles and branches and there may have been two subsequent independent shifts to increased use of litter and vegetation matrices with related anatomical changes (A. moreoticus, A. fitzingeri), and one to increased use of crevices (A. marchi). Some members of Algyroides are strikingly similar in superficial morphology to particular species of the equatorial African genus Adolfus. This resemblance results from a combination of many shared primitive features plus a few independently acquired derived ones that are likely to give performance advantage in the relatively similar structural niches that these forms occupy. This study provides evidence that: (1) the use of a combination of molecular and morphological data may sometimes allow the estimation of ancestral anatomical features when these are otherwise unknown; (2) process considerations may permit a choice to be made in cases of character evolution where tree topology means that equally parsimonious alternatives exist; such decisions about character evolution may allow ecological shifts to be similarly assessed; (3) parallel evolution in ecological analogues may involve relatively few characters.
Variations in life-history traits in the common frog Rana temporaria (Amphibia: Anura): a literature review and new data from the French Alps
- Claude Miaud, Robert Guyétant, Johan Elmberg
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- 01 September 1999, pp. 61-73
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Life-history traits of Rana temporaria were studied in an alpine French population and in the literature. In the living frogs, mean adult body length was greater in females than in males. Sexual dimorphism in body length was 0.109 using Lovich & Gibbon's (1992) formula, but tended to decrease with age. Age of adult frogs was assessed by skeletochronology, and age distribution was not significantly different between the sexes (range 4–15 years in males, 5–12 in females). Adult survival rate was about 0.80 in both sexes. Once maturity was reached, the total expected longevity was 6.1 years in males and 5.5 years in females. Age and body length were positively correlated in both sexes. The growth coefficient (K) was 0.47 in males, and 0.55 in females, mainly reflected as faster female growth between metamorphosis and maturation. Growth rate generally decreased before sexual maturity was reached. On average, females matured 1 year later than males. Newly metamorphosed froglets averaged 16.1 mm. When combined with published data from 12 European populations of R. temporaria, the following general patterns emerge. Mean adult body length is significantly greater in females than in males, and mean body length at maturity shows the same trend. Variation in mean age at maturity and in longevity are considerable among populations, but there is no consistent trend of difference between the sexes. Body length and age are correlated between males and females, i.e. populations with long and old males also have long and old females. Mean adult body length, mean body length at maturity, age at maturity, and longevity all increase with decreasing activity period. Adults exposed to a short activity period grow slower but attain a greater final length. Sexual dimorphism in body length generally increases as activity period gets shorter. Polygons describing norms of reaction for maturation in an age–body length space are similarly oriented in both sexes, but with a wider range in age for females. This is due to an older age at maturity for females in populations with a short activity season. Mean age and length at maturity are significantly correlated in females, but not in males, partly supporting the hypothesis that this species has a flexible pattern of development. Observed patterns are compared with predictions from life-history theory, paying attention to all life stages and environmental variation.
Small mammal communities and fire in the Brazilian Cerrado
- Emerson M. Vieira
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- 01 September 1999, pp. 75-81
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The Cerrado biome is a complex of open forests, gallery forests and savanna vegetation that harbours about 60 species of small mammals. Despite the frequent occurrence of fire in this habitat, few studies have analysed its effects on the small mammal communities in the Cerrado. The purpose of this study was to compare small mammal communities of two Cerrado areas with different fire frequencies (2–3 years or more than 18 years since the last fire). Species diversity, species composition, and relative abundance of the most common species of these communities were analysed in both areas before and after fire. One trapping grid was established in each of the two areas. These areas were sampled monthly or bi-monthly during 1 year before and 1 year after prescribed fires. Both areas had communities with similar species composition, but the protected site had more species, before and after fire (seven and eight, respectively), than the frequently burned site (five and six species, respectively). Both sites showed significant post-fire changes for a short time period (3 months), but only the area that had been protected against fire for more time showed significant post-fire changes in relative abundance of the most common species (the murid rodents Bolomys lasiurus, Calomys callosus, C. tener, Oryzomys subflavus, and Thalpomys cerradensis) for the 1 year following fire. Overall diversity tended to be greater in the protected site both before and after fire. Species composition and diversity indices of the small mammal communities were relatively unaffected by the fires. Thalpomys cerradensis, however, appeared to use only areas that burn frequently and C. callosus showed an immediate post-fire increase in population size in both areas, contrasting with a simultaneous decrease in number of B. lasiurus.
Habitat correlates of European rabbit (Oryctolagus cuniculus) distribution after the spread of RVHD in Cadiz Province, Spain
- John E. Fa, Colin M. Sharples, Diana J. Bell
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- 01 September 1999, pp. 83-96
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European rabbit Oryctolagus cuniculus abundance was evaluated relative to habitat variables within two zones in Cadiz Province (south-east Spain), 2 years after the spread of rabbit viral haemorrhagic disease (RVHD). The study areas were low-lying regions of mixed pasture/agriculture, Mediterranean forest and matorral, west (Zone A) and east (Zone B) of the Sierra de Cadiz mountain chain. A total of 111 sites was selected to sample all existing habitat types. Relative rabbit density in each site was estimated using a pellet count method in one 50×2 m transect. Mean pellet densities were 21.05 pellet m-2 in Zone B and significantly lower, 5.85 pellets m-2 in Zone A. Physical variables, such as vegetation characteristics (density, height and patchiness of cover-layer vegetation, etc.), topographical characteristics and human influences, cover-layer vegetation species (woody stemmed shrub species or trees forming dense stands), and ground-layer vegetation species (herbaceous species forming a low-lying layer) were determined within transects. In addition to these variables, soil types, rock type and land form, as well as the presence of aquifers, irrigation, and the proximity of rivers, were determined for each site. Significantly higher pellet densities were found in mixed grassland/matorral and cork oak/matorral than in all other habitats. Pellet density was negatively correlated with density of ground-layer vegetation, while high pellet densities were associated with medium-height ground-layer vegetation (50–100 cm). The presence of Entisols was negatively associated with pellet density. The cover-layer species Pistacia lentiscus, Quercus coccifera, Quercus suber, Opuntia ficus-indica and the ground-layer grasses all showed positive associations with pellet density, while the ground-layer species Echium gaditana and Scorpiurus vermiculatus were negatively associated with pellet density. Multivariate analyses identified the mixed grassland/matorral and cork oak forest/matorral habitats as explaining most of the variation in pellet density. The height of cover-layer vegetation was negatively associated, while Opuntia and table-and/calcareous sandstone were positively associated, with mixed grassland/matorral. Distance from habitation and human impact were negatively associated and Quercus suber was positively associated with cork oak forest/matorral. Comparison with previous studies suggest that the low pellet counts in this study represent very low rabbit densities and the population as a whole appears depleted and fragmented.
Hunting and social behaviour of leopard seals (Hydrurga leptonyx) at Seal Island, South Shetland Islands, Antarctica
- Lisa M. Hiruki, Michael K. Schwartz, Peter L. Boveng
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- Published online by Cambridge University Press:
- 01 September 1999, pp. 97-109
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The hunting behaviour of leopard seals Hydrurga leptonyx was monitored opportunistically at Seal Island, South Shetland Islands, during the austral summers from 1986/87 to 1994/95. Leopard seals used several methods to catch Antarctic fur seal pups Arctocephalus gazella and chinstrap penguins Pygoscelis antarctica, and individuals showed different hunting styles and hunting success. One to two leopard seals per year were responsible for an average of 60% of observed captures of fur seal pups. Leopard seals preyed on penguins throughout the summer, but preyed on fur seal pups only between late December and mid-February. Hunting behaviour differed significantly between different locations on the island; fur seals were hunted only at one colony, and penguins were hunted in several areas. The relative abundance of prey types, size of prey in relation to predator, and specialization of individual leopard seals to hunt fur seal prey probably influence individual prey preferences among leopard seals. On five occasions, two leopard seals were seen together on Seal Island. Possible interpretations of the relationship between the inter-acting leopard seals included a mother–offspring relationship, a consorting male–female pair, and an adult leopard seal followed by an unrelated juvenile. In two incidents at Seal Island, two leopard seals were observed interacting while hunting: one seal captured fur seal pups and appeared to release them to the other seal. Observations of leopard seals interacting during hunting sessions were difficult to confirm as co-operative hunting, but they strongly implied that the two seals were not agonistic toward one another. The hunting success of individual leopard seals pursuing penguins or fur seals is probably high enough for co-operative hunting not to become a common hunting strategy; however, it may occur infrequently when it increases the hunting productivity of the seals.
Testicular and ovarian development in the harbour porpoise (Phocoena phocoena)
- C. V. Karakosta, P. D. Jepson, H. Ohira, A. Moore, P. M. Bennett, W. V. Holt
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- 01 September 1999, pp. 111-121
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Testicular and ovarian development in neonatal and juvenile harbour porpoises Phocoena phocoena was examined, using tissue from animals (males n = 65 and females n = 10) stranded or caught off the coast of England and Wales. Classification of the animals according to their stage of sexual development was undertaken using characteristics of gonadal morphology. Developmental correlates of the immature testis were increased incidence of prespermatogonia in seminiferous tubules, and the increasing proportion of testicular volume occupied by the seminiferous tubules. Using a low magnification (×200) assessment of testis sections, immature specimens could be grouped into three developmentally distinct classes, based on the relative amounts of interstitial and seminiferous tubule tissue and the frequency of prespermatogonia. Adult testes, showing either active spermatogenesis or seasonal quiescence, could be clearly distinguished from immature testes using histological criteria such as the presence of spermatocytes and spermatids. This classification system was used to determine relationships between testicular development and body size: males were classified as immature when they had body lengths < 135 cm and body weights < 30 kg; mature males always exceeded 140 cm in length and 40 kg in weight; and a transitional group, which was regarded as juvenile, had body lengths between 110 and 140 cm and body weights between 20 and 40 kg. Ovarian morphology was used to distinguish neonatal and juvenile females. Neonates displayed characteristic cords of naked ova which dispersed as development progressed. Neonatal animals had body lengths < 100 cm and body weights < 18 kg; within this group (n = 5) there were significant (P < 0.01) left-right asymmetries in the number of naked ova present, the left ovary containing nearly twice as many ova as the right. Although this difference was not apparent in females with body lengths > 100 cm and body weights between 19 and 30 kg (juveniles) it could be related to the almost total asymmetry of ovulation from the left ovary in this species (Harrison, Brownwell & Boice, 1972).
Occurrence of the genus Chiasmocleis (Anura: Microhylidae) in the State of Alagoas, north-eastern Brazil, with a description of a new species
- C. A. G. Cruz, U. Caramaschi, E. M. X. Freire
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- 01 September 1999, pp. 123-126
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The range of the genus Chiasmocleis found in the south-eastern and southern parts of the Atlantic rainforest of eastern Brazil, is expanded to the north-eastern region. Chiasmocleis alagoanus sp. nov. is described from two localities in the State of Alagoas. It is characterized by: medium size (snout–vent length 25.5–27.8 mm in females, 22.7–23.4 mm in males); snout short, tip truncate in dorsal and rounded in lateral profile; hand and foot not webbed; fingers and toes without lateral dermal spines; dorsal and ventral surfaces of body and limbs smooth, without dermal spines; colour on dorsum uniformly dark brown; venter roughly marbled in dark brown and pale cream.