Research Article
Environmental factors influencing the presence and abundance of a log-dwelling invertebrate, Euperipatoides rowelli (Onychophora: Peripatopsidae)
- S. Barclay, J. E. Ash, D. M. Rowell
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 425-436
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The microenvironments of decaying logs support a large invertebrate fauna, but are vulnerable to the effects of logging and land-clearing. We present an analysis of the relationship between a number of environmental variables and the occurrence and abundance of the onychophoran Euperipatoides rowelli within decomposing logs in a forest in south-eastern Australia. Aspect of slope is strongly associated with most of the environmental variables and is a good predictor of both presence and abundance of animals in logs; south-easterly slopes show higher levels of log colonization, larger populations within logs and a larger median weight of both males and females than north-westerly facing slopes. Degradation state of logs, log length, presence of termites and shrub cover are also associated with the presence of onychophorans, while log volume shows the most marked association with abundance. Euperipatoides rowelli prefers wetter logs, although there is considerable overlap between relative water content of logs with and without onychophorans. Log degradation state, an important factor in the presence of animals, is associated with years since felling, and our data indicate that logs become capable of supporting large populations of E. rowelli after a minimum of 45 years of decay. Thus future logging practices have important implications for the maintenance of suitable log environments for this species, and this result may be generalized to other log-reliant fauna.
Pheromonally mediated colonization patterns in the velvet worm Euperipatoides rowelli (Onychophora)
- S. D. Barclay, D. M. Rowell, J. E. Ash
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 437-446
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We demonstrate that the males of the peripatopsid Euperipatoides rowelli secrete a pheromone from crural papillae, which acts as an attractant to both males and females of the species. Patterns of colonization of decomposing logs and differences in sex ratio between incipient vs established populations indicate that males are the initial dispersers and colonizers, finding suitable log habitats in an exploratory fashion, while females subsequently colonize logs. This results in a disproportionately high frequency of males in newly colonized logs, followed by a gradual increase in female proportion. We argue that females use the aggregating pheromone secreted by males to target appropriate rotting log microhabitats, resulting in a clumped distribution of females compared to a more random distribution for males among recently colonized logs. This mode of colonization reduces the time that animals, especially females, spend outside the safety of suitable logs, and the risk-taking strategy of male exploration may explain the marked sex ratio bias in favour of females in the population. Rapid and non-random dispersal may also account for the contradictory evidence of unexpectedly high rates of colonization of new logs, yet a paucity of animals in leaf litter samples. The possibility that pheromone trails play a part in the dispersal process is discussed.
Reproductive biology of the onychophoran Euperipatoides rowelli
- Paul Sunnucks, Natalie C. Curach, Anthony Young, Jordan French, Ray Cameron, Dave A. Briscoe, Noel N. Tait
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 447-460
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The reproductive biology of the ovoviviparous peripatus Euperipatoides rowelli was investigated from field collections and laboratory cultures. The sexes have different demographics. The frequency distribution of individual weight is essentially L-shaped in females, but closer to normality for males: thus the sexes must exhibit different patterns of growth and/or mortality. Males are generally much smaller and rarer than females. The primary sex ratio seems to be 1:1 with equal investment in the sexes, while the tertiary ratio is highly female-biased. Logs with fewer individuals tend to be male-biased while well-populated logs tend to be female-biased. Males mature at 15–30% of the bodyweight of mature females. The weight frequency distribution of males without developed sperm in their tracts is strongly skewed to the lower weights, while that of males with sperm is more normally distributed, indicating that sperm production occurs as soon in life as possible. Males mature in their first year of life, if growth rates in culture may be extrapolated to the wild. In contrast to this rapid maturity in males, females may mature as late as their second or third years. Most mature females, and many prior to maturity, carry sperm in their spermathecae. After maturity, there is an approximately linear relationship between body mass and number of developing embryos. Reproduction in E. rowelli is significantly seasonal despite high individual variance, with a major bout of parturition in November–December (summer). A female can harbour one developed and one undeveloped batch of embryos in each uterus. Excesses of developed embryos in one uterus are counterbalanced by deficits of undeveloped ones, indicating that females can use their paired reproductive tracts independently. Individual females in culture can experience episodes of parturition approx. 6 months apart without re-mating, thus gestation may be 6 months or more. Sperm in spermathecae remain capable of vigorous swimming for at least 9.5 months.
Higher testicular activity in laboratory gerbils compared to wild Mongolian gerbils (Meriones unguiculatus)
- S. Blottner, C. Franz, M. Rohleder, O. Zinke, I. W. Stuermer
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 461-466
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The Mongolian gerbil has been used as laboratory animal since 1935. Breeding gerbils as an isolated laboratory population for decades may have led to a domestication process whose effects include changes in brain size. Quantitative changes in testicular activity could be assumed. Comparative intraspecific measurements were performed in 34 adult males of the laboratory strain (LAB) and in males raised as offspring of wild Mongolian gerbils (WILD) caught in central Mongolia (F1, n = 16; F2, n = 17). LAB and WILD were examined in January. Testicular spermatozoa were counted, proportions of different cell types were analysed using DNA flow cytometry, and mitotic and meiotic activity was calculated from DNA histograms. Intratesticular testosterone concentrations were measured with an enzyme immunoassay. In the WILD, testicular activity was lower and varied more. The overall weight, the efficiency of spermatogenesis (sperm/g testis) and resulting total sperm per testis were significantly less in offspring of wild gerbils. This corresponded with lower levels of haploid cells, total germ cell transformation of diploid cells to spermatids and meiotic transformation of spermatocytes to spermatids. The most profound difference was found in testicular testosterone concentration: the mean level was 405.7 ± 41.2 ng/g testis in LAB vs 6.4 ± 2.0 ng/g in WILD F1. All parameters changed in WILD F2 generation compared with F1 and diminished the differences with LAB. Differences between F1 and F2 were significant for testis mass, testis/body weight ratio, percentages of haploid cells and cells in G2/M phase, both germ cell transformations and testosterone concentration. The results suggest rapid, adaptive changes of male reproductive physiology in the early offspring generations from wild populations under laboratory breeding conditions. The breeding of Mongolian gerbils in the laboratory has influenced the testicular function resulting in increased spermatogenic activity and highly stimulated testosterone production.
Roosts used by the golden-tipped bat Kerivoula papuensis (Chiroptera: Vespertilionidae)
- M. Schulz
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 467-478
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Use of roosts by the golden-tipped bat Kerivoula papuensis was investigated principally in the Richmond Range National Park, north-eastern New South Wales and Mt Baldy State Forest in north-eastern Queensland, Australia. Roosts were located primarily using radiotelemetry; although others were found by light tagging, visual location and nest searching techniques. In the main study site in the Richmond Range a total of 54 K. papuensis diurnal roosts were located, 96.5% occurring in the suspended nests of the yellow-throated scrubwren Sericornis citreogularis (Pardalotidae) (93%) and the brown gerygone Gerygone mouki (Pardalotidae) (3.5%). At Mt Baldy, K. papuensis displayed greater variation in day roost selection, although 73% were in S. citreogularis nests. In both sites, day roosts were confined to the lower stratum of rainforest with a small proportion in eucalypt tall open forest with a rainforest subcanopy. Diurnal roosts were located at distances varying from 20 to 1200 m to the nearest major vegetation ecotone, ranging from along watercourses upslope to ridgelines. Diurnal roosts were predominantly occupied by single bats, with a maximum group size of eight individuals. All roosts in hanging nests were modified by the presence of a basal hole. Strong circumstantial evidence was provided that K. papuensis modified these roost structures in a manner allied to tent-making bats. A single maternity roost was located in the hollow of a rainforest canopy tree, Flindersia australis (Rutaceae).
Ultrastructure of the neck membrane in dragonflies (Insecta, Odonata)
- Stanislav N. Gorb
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 479-494
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This study describes and quantifies the microsculpture and ultrastructural design of the neck membrane of adult Odonata using scanning and transmission electron microscopy. The membranous cuticle has a complex pattern of microfolds, which appear to have a specialized mechanical function. The membrane has two layers of cuticle: the epicuticle and exocuticle. The outermost layer is the electron-opaque epicuticle, which repeats the shapes of the microfolds. The epicuticle has no fibrillar elements. The exocuticle is electron-lucent and rather thin (0.5–1.5 μm) compared with the sclerite cuticle. The cuticle microfibrils in successive lamellae are at angles to each other. The epidermal cells underlying the membrane have an electron-lucent matrix filled with electron-opaque spherical vesicles of 0.1–0.4 μm in diameter. The behaviour of the membrane folds under loading was studied by shock-freezing experiments; these showed that the shape of the folds changed in response to head movements, and stretched under loading. A comparison of the surface patterns of the membrane in 10 odonate species from seven families (Zygoptera: Coenagrionidae, Lestidae, Megapodagrionidae; Anisoptera: Cordulegastridae, Aeshnidae, Corduliidae, Libellulidae) had not revealed any correlation of the measured parameters with the size, sex or systematic position of species studied.
Diet composition of badgers (Meles meles) in a pristine forest and rural habitats of Poland compared to other European populations
- Jacek Goszczyński, Bogumiła Jedrzejewska, Włodzimierz Jedrzejewski
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 495-505
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Diets of the Eurasian badger Meles meles were studied by scat analysis in three localities of eastern and central Poland, representing pristine forests and rural landscapes with little wood cover. In spring, earthworms constituted 82–89% of biomass consumed by badgers in all localities. In summer and autumn, the proportion declined to 56% in the pristine forest, and to 24% in the mosaic of forests, fields, and orchards. Supplementary resources at this time were amphibians (in forests) or garden fruits (in a rural landscape). Literature on badger diet composition in Europe showed that earthworms and vegetable matter were the dominant food types of badgers, but their roles changed with latitude. The share of earthworms grew from nil at 37–40°N to 40–70% at 55–63°N; the opposite trend was observed for vegetable food. Also, of two major supplementary resources, vertebrates were taken by badgers more often at northern latitudes, and insects in the south. In consequence, the food niche of badgers was broadest at 45–55°N and became narrow at both lower and higher latitudes. Moreover, in the temperate zone of Europe, the degree of habitat transformation by humans significantly affected badger feeding habits. In forests, badgers relied predominantly on earthworms (on average, 62% in diets). In farmlands and pastures, earthworms and plant material (usually garden fruit and cereals) played equally important roles (34% each). This biogeographical pattern of feeding habits can be explained by variation in abundance and availability of earthworms.
Interspecific scaling of the hindlimb skeleton in lizards, crocodilians, felids and canids: does limb bone shape correlate with limb posture?
- Richard W. Blob
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 507-531
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During locomotion, lizards and crocodilians generally use a more sprawling limb posture than most mammals and experience substantial axial rotation of the femur. Consequently, the limb bones of most mammals are loaded predominantly in bending, but the limb bones of lizards and crocodilians are loaded primarily in torsion. As body size increases, torsional shear stress in limb bones is expected to increase more than bending stress; therefore, limb bone diameters of lizards and crocodilians might be expected to scale with relatively greater positive allometry than limb bone diameters of mammals that use upright posture. To test this hypothesis, scaling patterns of the femur and tibia in lizards (iguanians and varanids) and crocodilians were compared with patterns in felid and canid mammals, using both non-phylogenetic statistical methods and phylogenetically independent contrasts. Comparisons with theoretical models indicate that size-related changes in limb bone geometry do not completely compensate for size-related increases in limb bone stress in the lizard or crocodilian lineages examined. Unless lizards and crocodilians compensate for size-related increases in limb bone stress through other mechanisms (e.g. changes in limb kinematics or the mechanical properties of limb bones), limb bone stresses are predicted to be relatively greater among larger species of these lineages. However, limb bone diameters appear to scale with greater positive allometry (relative to body mass) in varanids than in iguanians, suggesting that larger lizard lineages might compensate for increased stress through changes in bone geometry to a greater degree than smaller lineages. Allometric scaling patterns for many limb bone diameters among iguanians are more similar to those of felids and canids than to those of varanids; thus, sprawling locomotor habits do not correlate clearly with a particular pattern of limb bone scaling. This suggests that similarity of interspecific scaling patterns of limb bone lengths and diameters is not sufficient to justify inferences of similar locomotor function.
Energetics of gestation relative to lactation in a precocial rodent, the guinea pig (Cavia porcellus)
- Joachim Künkele
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 533-539
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Caviomorph rodents stand out by showing long gestation periods and by producing few, extremely precocial young. This study compares the energy cost of gestation vs lactation in the guinea pig Cavia porcellus. It tests whether the long gestation period (68 days) relative to the total period of maternal care (88 days) of this species results in a high overall energetic efficiency of reproduction. Although maternal net production during gestation was as high as during lactation, the net energy cost of gestation was much lower than the energy cost of lactation. Mothers' average increase in energy intake (above the non-reproductive level) was only 16% during gestation, but 92% during lactation. Similarly, the peak in maternal energy intake was low during gestation (2.4 times basal metabolic rate) compared to the peak during lactation (3.7 times basal metabolic rate). Consequently, the efficiency of energy conversion into offspring tissue during gestation (62%) was almost twice as high as the efficiency during lactation (35%). The results agree with the hypothesis that a prolonged gestation period increases the overall energetic efficiency of reproduction.
Use of a seaweed habitat by red deer (Cervus elaphus L.)
- L. Conradt
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 541-549
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The use of a seaweed habitat by red deer Cervus elaphus L. on the Isle of Rum, Scotland, was examined in detail. New information is provided on diet selection, timing of seaweed use relative to tides, inter-individual differences in seaweed use, and sex differences in site use within the seaweed habitat (‘site segregation’). Interestingly, seaweed use by adult males and females was closely correlated to that of their mothers. This implies that deer ‘learn’ early in life to include seaweed into their diet. Formerly, it has been suggested that male inferiority in indirect competitive ability relative to females causes site segregation in dimorphic ungulates (‘indirect competition hypothesis’). The observed pattern of site segregation within the seaweed habitat was used to test the hypothesis, which predicts that males should be found at sites where they can achieve higher intake rates, but where forage quality is lower than at female sites. With respect to seaweed use, the hypothesis further predicts that segregation should be lower within the seaweed habitat than within terrestrial vegetation communities, and that males should time seaweed use earlier (relative to the tide) than females. This is because seaweed availability is more subject to tidal rhythm than to indirect competition in comparison to terrestrial habitats. Males and females used different bays, and within bays they used different fractions of seaweed. However, male-preferred sites did not yield higher intake rates and were not of lower forage quality than sites preferred by females. Moreover, segregation was not lower within the seaweed habitat than within terrestrial vegetation communities, and males did not time their seaweed use earlier relative to the tide than did females. The indirect competition hypothesis could not explain the observed pattern of site segregation. Other factors, such as sex differences in sheltering or anti-predator behaviour, or social harassment, could be responsible instead.
Testes size, body size and male–male competition in acanthocephalan parasites
- Robert Poulin, Serge Morand
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- Published online by Cambridge University Press:
- 01 April 2000, pp. 551-558
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In most invertebrate taxa, males are smaller than females and large male sizes are only favoured when male–male competition for access to females is intense. In addition, in species where sperm competition is important, relatively large investments in testis growth and ejaculate size will also be selected. The use of copulatory plugs by males of some taxa, however, can prevent sperm competition. We hypothesized that, across related species in which males use plugs, relative testis size would decrease with increases in the severity of male–male competition. We tested this prediction across 112 species of acanthocephalan parasites, worms that use copulatory plugs and in which male–male competition occurs. As a measure of the intensity of male–male competition in a given species, we used sexual size dimorphism. Male and female body volume covaried allometrically, suggesting that sexual size dimorphism in acanthocephalans is the product of sexual selection. Our main finding is that relative testis volume, corrected for male body volume, decreases significantly as male body volume relative to female volume increases, i.e. as the sexual size dimorphism becomes less female-biased. All our results remained unchanged after we controlled for potential phylogenetic effects. The relationship indicates that investment in testis growth beyond the minimum size required for efficient fertilization becomes increasingly less important in species where males appear to compete intensely for mating opportunities.