Plasma corticosterone concentrations in wild and captive juvenile tuatara (Sphenodon punctatus)

Abstract

High mortality and abnormal growth patterns commonly found in captive juvenile tuatara were hypothesised to be due in part to the effects of long‐term chronic stress of captivity. This study compared plasma concentrations of the reptilian adrenal steroid, corticosterone, in wild juvenile tuatara on Stephens Island, Cook Strait, and in captive juveniles of Stephens Island origin, held in New Zealand institutions, in February and August 1992. Seasonal variation in plasma concentration of corticosterone in wild juveniles in four seasons of the year was also examined. This is the first study of seasonal cycles in plasma corticosterone in a wild juvenile reptile. Plasma corticosterone concentrations were significantly higher in captive juvenile females (4.21 ± 0.27 ng/ml; mean ± SE) compared with wild juvenile females (2.44 ± 0.42 ng/ml) in February (P < 0.05), but not in August, and there was no difference in concentration between captive and wild juvenile males in either month. There was significant seasonal variation in plasma corticosterone in wild juvenile females (P < 0.05). However, there was no seasonal variation observed in wild juvenile males, and the magnitude of the variation in plasma corticosterone was low in both sexes (1.28 ± 0 ng/ml ‐4.65 ±3.41 ng/ml). Although mean plasma corticosterone was higher in captive juvenile females compared with wild juvenile females in February 1992, the value in captive females was within the range of mean plasma corticosterone concentrations observed in the seasonal study, and may be therefore due to asynchronicity of seasonal cycles, rather than stress. Further research is required; however, lack of correlation between plasma corticosterone concentrations and either growth rate or density indicate that captive juvenile tuatara in New Zealand are not suffering from pronounced chronic stress.     

Observations on the activity and thermoregulation of the tuatara,Sphenodon punctatus (Reptilia: Rhynchocephalia), on Stephens Island

Abstract

Observations on Stephens Island, Cook Strait, in December 1978 show that although the tuatara is generally most active at night, many animals spend much of the day at or beyond burrow entrances, apparently to increase their body temperature. During the day, tuataras tend to move further from burrows which are under shaded forest than from those in open pasture. By day, mean body temperatures (±SE) ranged from 17.2±0.5°C in forest shade to 24.6±1.1°C in full sunlight; the maximum body temperature recorded was 26.3°C. The significance of these observations is discussed.     

Pairing and insemination patterns in a giant weta (Deinacrida rugosa: Orthoptera; Anostostomatidae)

Positive size assortative mating can arise if either one or both sexes prefer bigger mates or if the success of larger males in contests for larger females leaves smaller males to mate with smaller females. Moreover, body size could not only influence pairing patterns before copulation but also the covariance between female size and size of ejaculate (number of spermatophores) transferred to a mate. In this field study, we examine the pre-copulatory mate choice, as well as insemination, patterns in the Cook Strait giant weta (Deinacrida rugosa). D. rugosa is a large orthopteran insect that exhibits strong female-biased sexual dimorphism, with females being nearly twice as heavy as males. Contrary to the general expectation of male preference for large females in insects with female-biased size dimorphism, we found only weak support for positive size assortative mating based on size (tibia length). Interestingly, although there was no correlation between male body size and the number of spermatophores transferred, we did find that males pass more spermatophores to lighter females. This pattern of sperm transfer does not appear to be a consequence of those males that mate heavier females being sperm depleted. Instead, males may provide lighter females with more spermatophores perhaps because these females pose less of a sperm competition risk to mates.     

Identifying a causal agent of sexual selection on weaponry in an insect

In many animal species, males do not seek females directly butinstead locate and defend sites that contain spatially or temporallylimited resources essential to female survival and reproduction.Resident males that successfully repel conspecific rivals canmate with females attracted to these resources. In theory, increasingresource value increases harem size and thus increases the opportunity(I_mates) for and strength of sexual selection on traits crucialto male resource-holding potential and mating success. I experimentallytested this hypothesis in the field using the Wellington treeweta, Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae),a sexually dimorphic insect in which males use their enlargedmandibles as weapons in male–male contests over accessto females sheltering in tree cavities (galleries). By manipulatinggallery size, I showed that, compared with smaller galleries,larger galleries housed larger harems. Variation in gallerysize was an important determinant of I_mates, but contrary toexpectation, greater opportunity existed in small galleriescompared with large galleries. As predicted, male weapon sizewas under stronger directional selection in large galleriesbecause the fitness benefits were greater under these conditionscompared with small galleries. My results help explain the positiveassociation between average weapon size and average gallerysize observed within and among tree weta populations in NewZealand.     

Historical biogeography of the New Zealand freshwater crayfishes (Parastacidae,Paranephrops spp.): Restoration of a refugial survivor?

Abstract

New Zealand's two species of freshwater parastacid crayfishes have allopatric distributions, with one species in the North Island and northwestern South Island and the other in the eastern and southern South Island and Stewart Island. This gives the appearance of a vicariance event driven by uplift of the Southern Alps beginning in the Pliocene, and of former land connections across both Cook Strait and Foveaux Strait. However, separation of the two species may date from before the Southern Alps were formed. A diverse series of historical geological events is invoked to explain details of the distributions of these two species. Absence of Paranephrops from intermontane valleys of eastern flanks of the Southern Alps is notably different from patterns seen in freshwater fish species.     

The Relationship Between Resource Control, Association with Females and Male Weapon Size in a Male Dominance Insect

In species with a resource‐defence (male dominance) mating system, males are expected to maximize fitness by controlling resources deemed more valuable by sexually receptive females because these sites attract more mates. Furthermore, males, which control more valuable resources should themselves be of high quality. I experimentally tested these predictions in the laboratory using the sexually dimorphic Wellington tree weta, Hemideina crassidens (Blanchard) (Orthoptera: Tettigonioidea: Anostostomatidae). Male H. crassidens use their mandibular weaponry to fight for control of harems (groups of adult females) that seek shelter in trees cavities (galleries). As predicted, larger galleries housed significantly larger groups of females and males with larger weaponry controlled large galleries significantly more often. Therefore, galleries with a larger volume are likely considered more valuable by males because they house larger harems. However, contrary to prediction, males with larger weaponry did not reside with significantly more females overall because females did not always form the largest possible groups in galleries and males with smaller weaponry were able to reside with single females in small galleries. The latter observation suggests a possible alternative mating strategy by disadvantaged males.     

Variation in biomass and carbon storage by stand age in pine (Pinus tabulaeformis) planted ecosystem in Mt. Taiyue,Shanxi, China

Forest ecosystems play dominant roles in global carbon budget because of the large quantities stored in live biomass, detritus, and soil organic matter. Researchers in various countries have investigated regional and continental scale patterns of carbon (C) stocks in forest ecosystems; however, the relationship between stand age in different components (vegetation, forest floor detritus, and mineral soil) and C storage and sequestration remains poorly understood. In this paper, we assessed an age sequence of 18-, 20-, 25-, 38-, and 42-year-old Pinus tabulaeformis planted by analyzing the vertical distribution of different components biomass with similar site conditions on Mt. Taiyue, Shanxi, China. The results showed that biomass of P. tabulaeformis planted stands was ranged from 88.59 Mg ha^?1 for the 25-year-old stand to 231.05 Mg ha^?1 for the 42-year-old stand and the major biomass was in the stems. Biomass of the ground vegetation varied from 0.51 to 1.35 Mg C ha^?1 between the five stands. The forest floor biomass increased with increasing stand age. The mean C concentration of total tree was 49.94%, which was higher than C concentrations of ground vegetation and forest floor. Different organs of trees C concentration were between 54.14% and 47.74%. C concentrations stored in the mineral soil for each stand experienced decline with increasing soil depth, but were age-independent. Total C storage of five planted forests ranged from 122.15 to 229.85 Mg C ha^?1, of which 51.44–68.38% of C storage was in the soil and 28.46–45.21% in vegetation. The study provided not only with an estimation biomass of P. tabulaeformis planted forest in Mt. Taiyue, Shanxi, China, but also with accurately estimating forest C storage at ecosystem scale.     

Male reproductive success in harem troops of hanuman langurs (Presbytis entellus)

Based on cross-sectional and longitudinal data collected in 1967–1988 by various observers, male reproductive success was studied in the Hanuman langurs of Jodhpur, India. The harem-structured social organization ensures a high degree of paternity certainty. Births occur throughout the year, with significant peaks and minima in March and November, respectively (n =398).The interbirth interval averages 16.7 months (n = 114).The duration of harem residencies varies between 3 days and ≥ 74.0 months, with a mean of 26.5 (n = 64). Harem holder replacements occur during all months of the year. No male achieves residency in more than one troop, suggesting that residency is associated with a distinct peak in the resource holding potential of a given male. Reproductive success among males varies considerably. Male mortality is high due to migration and intrasexual competition, leading to an adult sex ratio of 1:4.9. It is estimated that one-quarter of all adult males will never gain harem residency. Conceptions achieved outside harem residencies are so rare (4.7%) that a viable low-risk strategy, opting for longevity instead of harem residency, is unlikely. Tenure length has a stronger influence on male reproductive success than harem size because interbirth intervals are significantly shorter in small harems than in larger ones. It is assumed that females in one-male breeding structures compete for sperm and that such competition is more intense in larger harems.     

Habitat use by three rat species (

Ship rats (Rattus rattus) were removed from sites on Pearl Island, southern Stewart Island, in 2004 and 2005, to test whether they excluded Pacific rats (R. exulans) or Norway rats (R. norvegicus) or both from podocarp-broadleaf forest. As predators can influence habitat use in rodents, Pearl Island was selected because no mammalian predators of rodents are present. Rats were trapped in two other habitats to clarify rat distribution on the island and to obtain samples for stable isotope investigation of food partitioning within habitats. The experimental removal of ship rats failed, as Pacific rats were found to share forest and shrubland with ship rats. This result contrasted with the restricted distribution of Pacific rats on Stewart Island. Ship rats were ubiquitous, and appear to have been the dominant species in podocarp-broadleaf forest on Pearl Island. The largest species, the Norway rat, was trapped only on the foreshore of Pearl Island, but on Stewart Island it is more widespread. Ship rats and Norway rats were partitioning the coastal habitat by exploiting different food sources. Stable isotopic ratios (δ¹⁵N and δ¹³C) in muscle samples from Norway rats revealed a strong marine signature, suggesting intensive foraging in the intertidal zone. Ship rats trapped in the same habitat exhibited mixed terrestrial and marine sources in their diet. There was little obvious partitioning between ship rats and Pacific rats in forest, except a possible delay in breeding in Pacific rats relative to ship rats. Whether Norway rats select the intertidal zone to forage, or were excluded from forest by ship rats is unknown, but competitive exclusion is likely. Estimated densities of rats were low (2.1–5.1 rats ha^-1 in forest, 1.42 rats ha^-1 in shrubland) and similar to other New Zealand sites with low soil fertility. Further research will be required to elucidate the roles of food quality, habitat structure and predation in facilitating habitat selection in these species.     

Low survival rate and high predation in the African hingeback tortoise Kinixys spekii

Live and dead Kinixys spekii were collected in the Sengwa Wildlife Research Area, Zimbabwe over a 12‐year period. Live tortoises were sexable at a midline plastron length of 100 mm; females were considered sexually mature at 140 mm (reached by age 9 years) and males at 120 mm (at age 7 years). Adult females were significantly larger than males, on average by 14 mm in length and by 1.43 times in mass. Mark–recapture analysis in a 2 km² area showed a population density of sexable tortoises of 0.16 ha^?1. The survival rate was estimated by recaptures, by the frequency distribution of age at death, and by the mean age of live tortoises, and averaged 0.74 year^?1. Seventy‐seven to 89% of dead tortoises showed evidence of predation, depending on the criteria used. Damage occurred in characteristic forms, loss of the front or rear of the plastron, or holes in the carapace and plastron, which were attributed to predation by mammals and ground hornbills, respectively. K. spekii had similar body size and sexual size dimorphism to Mediterranean tortoises (Testudo), but population density was much lower and the mortality rate was twice as high, probably due to the abundant African predators. High mortality was offset by a rate of juvenile growth twice that of Testudo.     

Gender- and sequence-dependent predation within group colonizers of defended plants: a constraint on cheating among bark beetles?

Bark beetles engage in one of the most pronounced examples of group procurement of defended plants. Their aggregation pheromones attract both sexes and are essential to overcome constitutive and rapidly inducible lethal defenses. The relative benefits to senders versus receivers of these signals are only partly understood. Because the initial stage of host entry can be hazardous, there may be benefit to a cheating strategy, whose practitioners respond to pheromones but do not engage in host searching. Several disadvantages to cheating have been proposed, but the role of predators has not been considered. Predators exploit bark beetle pheromones to locate prey, accumulate at the breeding site, and consume adult bark beetles before they enter the tree. Preliminary experiments quantified arrival patterns in the field. We used a laboratory assay to investigate relative predation on pioneers (those that initially select and enter hosts) and responders (those that arrive at a host in response to pheromones) during host colonization. Our model system utilized the pine engraver, Ips pini, which exhibits male harem polygamy. We allowed male I. pini to colonize host tissue and added females 1 day later. Also 1 day later, we variably added additional males and predacious checkered beetles, Thanasimus dubius. These treatments included two densities of males and three densities of predators that were selected to emulate field conditions. Responding males experienced higher predation than pioneers. T. dubius ate more males than females, independent of the presence or absence of responding males. T. dubius affected the distribution of females per male, although the number of females that survived to construct ovipositional galleries was constant. We discuss the viability of cheating, implications for biological control, and predator-prey coevolution in this cooperative, group-colonizing herbivore.     

Intracolonial social distance, signaling modality and association choice in the greater sac-winged bat (Saccopteryx bilineata)

Understanding patterns and motivations for social spacing greatly illuminates the structure and internal dynamics of given groups or social systems. The highly social, polygynous greater sac-winged bat, Saccopteryx bilineata, represents an excellent model for social spacing analysis, since the choice of individual roost-sites within a day-roost constitutes an enduring, often repeated decision about relative proximity to conspecifics. Day-roost colonies consist of one or multiple harem groups, each with several females and a single male. Additionally, non-harem males without females may be present. A social-distance–time-budget metric revealed that harem males, females, and non-harem males differed significantly in their respective spatial associations while roosting. Harem males and females were most closely associated, with harem males located at the center of harems instead of at the borders. Non-harem males associated significantly closer with one another than with harem females that they were trying to access. The signaling modality mediating social interactions depended on the respective social distances between communicating bats. Our results suggest a concentric social organization based around harem males, which may select roost sites in closest possible proximity to females to enhance courtship signal strength. This constitutes an interesting deviation from the normative form of harem maintenance, patrolling borders, in mammals.     

Survival,density and population composition of bellbirds (Anthornis melanura) on the Poor Knights Islands,New Zealand

Abstract

Annual survival by age, sex and population size and structure were estimated for bellbirds (Anthornis melanura) from a banding study undertaken on 66 ha Aorangi Island, the Poor Knights Islands, during the period 1978–86. The most parsimonious model showed that survival varied little with age and between years, and survival of males was significantly greater than that of females. Recapture probabilities of males and females were similar, but did vary with age and among years. There was a greater proportion of adult males than adult females and sub‐adult males and females in the population, and among adults the male bias in the population was greater during the breeding season than at other times of the year. The high density of bellbirds on the Poor Knights and larger size of males may explain the reduced survival of females relative to that of males.     

Harem size and oviposition behaviour in a polygynous bark beetle

Abstract. 1. Harem polygyny can have fitness benefits and costs on females. In bark beetles of the genus Ips the latter may include within‐harem competition between larvae. However, earlier competition between females for male care and mating opportunities may also influence oviposition behaviour. There has been relatively little investigation into the relationship between harem size and initial egg output. The present study investigated this relationship in the bark beetle Ips grandicollis. 2. The measure of egg output used was the number of eggs in the gallery with the most eggs in each harem. Mean (±SE) harem size of 242 observed harems was 3.25 ± 0.10. A curvilinear relationship was found between egg output and harem size, with females in smaller harems (one to four females) laying more eggs with increased harem size. However, females in larger harems (five to seven females) laid fewer eggs as harem size increased. The optimal harem size (in terms of number of eggs laid) was close to four females. 3. We found no evidence from a behavioural assay that females could preferentially choose unmated males over mated males with harems of two females. Additionally, the distribution of harem sizes suggests that females distribute themselves among males randomly. 4. The results suggest that harem size has effects on female reproduction that extend beyond larval competition and influence patterns of oviposition. The mechanism that determines why egg laying is greatest at intermediate levels is unknown. There is no evidence that smaller harems belong to lower quality males, but females may adjust egg‐laying behaviour in large harems as a result of reduced male attendance or anticipated larval competition.     

Social Organisation of the Bat Tadarida (Chaerephon) pumila (Chiroptera: Molossidae) in Ghana,West Africa

The social organisation of the polyoestrous bat, Tadarida pumila, was studied in northern Ghana (W. Africa). Although this small (8.5 g) free-tailed bat is not sexually dimorphic in size, adult males develop an odorous inter-aural crest of long hairs. A social system based on female defense polygyny was found which, however, also involved elements of resource defense. Harems, each generally established in the roof space of separate dwelling houses with restricted access, consisted of an adult male and up to 21 females, with their young, and the larger harems were held by heavier males. Harem composition was stable and both harem males and females showed high site fidelity over the study span of 16 months. Some female young were recruited to their natal harems, at a sufficient rate to replace the annual loss of harem females. However, most young, which were born in three successive cohorts during the rainy season, apparently dispersed over the dry season, following early sexual maturation. The potential causes and benefits of female associations are discussed.     

Population genetic subdivision in the New Zealand greenshell mussel (Perna canaliculus) inferred from single-strand conformation polymorphism analysis of mitochondrial DNA

Single-strand conformation polymorphism (SSCP) analysis of the NADH IV region of the mitochondrial DNA (mtDNA) molecule in greenshell mussels (Perna canaliculus) indicated strong population genetic structuring in this endemic New Zealand species. A northern and a southern group were differentiated by frequency shifts in common haplotypes and by the occurrence of a unique southern haplotype at approximately 20% frequency. This split occurred south of Cook Strait (the body of water between the North and the South Island) at approximately 42 degrees S latitude. Northern populations were less genetically diverse than southern populations and mussels from the west coast of the South Island were most distinct from northern mussels. We hypothesize that the unique haplotype VIII originated in the lower South Island, and that its spread northwards was obstructed by the opening of Cook Strait approximately 15 000-16 000 years ago and the subsequent establishment of present-day surface water circulation patterns in Greater Cook Strait. We suggest that present-day strong tidal flows and turbulent mixing of water masses in Cook Strait, and intense up-welling on the east and west coasts in this region, represent a barrier to gene flow between mussels located in the North Island and northern South Island vs. mussels in most of the South Island and Stewart Island.     

Spawning behavior of the collared knifefish, Cymolutes torquatus (Labridae) in Papua New Guinea

A colony of about 10 males and nearly 50 females of Cymolutes torquatus was found living over a sandy substrate on the northwest coast of Normanby Island, Papua New Guinea at depths 14.6 to 18.3 m. Each male appeared to maintain a harem territory, approximately 10 to 14 m², with 4 or 5 females living in sub-units of this territory. Eleven matings of 4 males with 2 to 4 females each were observed on 9 and 10 April 1993. The broadcast spawning of C. torquatus is similar to that of razorfishes in the genus Xyrichtys, but in contrast to most tropical sand-dwelling fishes which mate at dawn or dusk, C. torquatus mated mid-morning between 9:38 to 10:50 h. Sexual dichromatism, color changes during mating, and cleaning of C. torquatus by juvenile Halichoeres zeylonicus are documented with color photographs.     

Distribution and movements of Buller's albatross (Diomedea bulleri) in Australasian seas

Abstract

The distribution and movements of Buller's albatross in Australasian seas are analysed using results of shipborne surveys (13 238 10‐min counts), counts from trawlers, banding data, recoveries on beaches and fishing vessels, and records from the literature. Patterns of marine distribution are documented by monthly accounts and maps. During the breeding season, highest abundances are recorded over shelves and slopes off southern New Zealand (The Snares shelf to 41–43°S off the South Island, D. b. bulleri), around the Chatham Islands and over oceanic subtropical waters east of New Zealand (probably D. b. platei), with marked seasonal variations observed off southern New Zealand. Both subspecies disperse mostly outside Australasian waters during the non‐breeding season. Birds banded on The Snares were recovered off south‐eastern New Zealand (Stewart Island to Cook Strait) and in the eastern tropical Pacific. Immatures accounted for only 0.25% of birds censused during the ship‐borne surveys; they are recorded around the New Zealand mainland in August‐October and February‐May, off south‐eastern Australia and in the Tasman Sea in November‐December, February, and June‐July. Around New Zealand, males predominate among birds recovered along the eastern seaboard, whereas the sex ratio in south‐western waters tends to vary according to water depth and season. Distribution patterns and movements in New Zealand and Australian seas are discussed in relation to breeding events and breeding status.     

The Powelliphanta gilliesi — traversi — hochstetteri — rossiana — lignaria — superba ring species (Mollusca: Pulmonata)

Abstract

An alternative classification of the New Zealand rhytidid genus Powelliphanta O'Connor, 1945 is presented. P. gilliesi (Smith, 1880) and P. hochstetteri (Pfeiffer, 1862) are recognised as species, and the following subspecies are recognised within hochstetteri: h. superba (Powell, 1930); h. lignaria (Hutton, 1880); h. rossiana (Powell, 1930); h. fiordlandica (Climo, 1971). A model is explained which suggests that all described species and subspecies of Powelliphanta are part of a complex ring species. This developed as a result of glacial disjunction of South Island populations and hybridisation across a Cook Strait land bridge formed during the last Pleistocene glaciation.     

Reduced MHC variation in a threatened tuatara species

The relationship between neutral and adaptive genetic diversity is important to understand in assessing the implications of a population bottleneck. Fitness-related genes, such as those of the major histocompatibility complex (MHC), may be influenced by selection, and so retain diversity even when it is lost at neutral markers. We measured MHC class I variation in an archaic reptile species Sphenodon guntheri [North Brother Island (NBI) tuatara], which naturally occurs on one 4 ha island in Cook Strait, New Zealand, and has low levels of microsatellite diversity. MHC variation in S. guntheri was compared with microsatellite DNA variation, and with MHC variation in a large population of Sphenodon punctatus (Cook Strait tuatara) on Stephens Island. The NBI population shows significantly decreased levels of genetic diversity compared with the Stephens Island population. Only three different MHC sequences and three genotypes were found on NBI, compared with 15 sequences and 21 genotypes in a similar sample size from Stephens Island. Two sequences appear to be unique to the NBI population. The assortment of sequence variants into genotypes suggests strong gametic disequilibrium between two MHC class I loci in S. guntheri , and only two haplotypes that were present in Hardy–Weinberg proportions were identified. MHC diversity in NBI tuatara appears to be largely influenced by genetic drift, consistent with a recent population bottleneck. This may compromise the ability of this population to respond to novel disease threats.