The active metabolic rate predicts a male spider's proximity to females and expected fitness

Conspicuous traits, such as weaponry and body size, are often correlated with fitness. By contrast, we understand less about how inconspicuous physiological traits affect fitness. Not only is linking physiology directly to fitness a challenge, but in addition, behavioural studies most often focus on resting or basal metabolic rates, resulting in a poor understanding of how active metabolic rates affect fitness. Here we use the golden orb-web spider ( Nephila plumipes ), a species for which proximity to a female on the web predicts a male's paternity share, to examine the role of resting and active metabolic rates in fitness. Using a semi-natural experimental set-up, we show that males closer to a female have higher active metabolic rates than males further from females. This higher metabolic activity is paralleled by increased citrate synthase activity, suggesting greater mitochondrial densities. Our results link both higher active metabolic rates and increased citrate synthase activity with fitness. Coupled with the behaviour and life history of N. plumipes , these results provide insight into the evolution of physiological systems.

Male copulation frequency, sperm competition and genital damage in the golden orb-web spider (Nephila plumipes)

Copulation in many sexually cannibalistic spiders is associated with a loss of function of the male reproductive organs and, as a consequence, males that survive sexual cannibalism may nevertheless be unable to subsequently copulate successfully. Sexual cannibalism is common in the Australian golden orb-web spider (Nephila plumipes), in which the tip of the conductor typically breaks during copulation. Thus, male mating frequency may be physiologically limited to two females, irrespective of the male’s ability to avoid cannibalism or the opportunity to locate and court additional, receptive females. Laboratory experiments revealed that the likelihood of the conductor breaking depends upon the copulatory history of the female insemination duct: males were more likely to break their conductor if they inseminated a ‘virgin’ rather than ‘mated’ insemination duct. However, the choice of insemination duct did not influence the duration of copulation or quantity of sperm transferred. In field populations, the proportion of males with both conductors broken increased during the course of the mating season, but while males with broken conductors did not copulate successfully with virgin females, they were nevertheless observed on the webs of immature females. We suggest that male N. plumipes with broken conductors on the webs of females are most likely mate guarding, as this appears to be the most effective mechanism of securing paternity.

The systematics and biology of the spider genus Nephila (Araneae:Nephilidae) in the Australasian region

Five species of the nephilid genus Nephila Leach are found in the Australasian region, which for the purposes of this study was defined as Australia and its dependencies (including Lord Howe I., Norfolk I., Christmas I., Cocos (Keeling) Is), New Guinea (including Papua New Guinea and the Indonesian province of West Papua), Solomon Is, Vanuatu, New Caledonia, Fiji, Tonga, Niue, New Zealand and other parts of the south-west Pacific region. All species are redescribed and illustrated. Nephila pilipes (Fabricius) occurs in the closed forests of eastern and northern Australia, New Guinea, Solomon Is and Vanuatu (through to South-East Asia); N. plumipes (Latreille) is found in Australia (including Lord Howe I. and Norfolk I.), New Guinea, Vanuatu, Solomon Is and New Caledonia; N. tetragnathoides (Walckenaer) inhabits Fiji, Tonga and Niue; N. antipodiana (Walckenaer) occurs in northern Australia (as well as Christmas I.), New Guinea and Solomon Is (through to South-East Asia); and N. edulis (Labillardière) is found in Australia (including Cocos (Keeling) Is), New Guinea, New Zealand and New Caledonia. Epeira (Nephila) walckenaeri Doleschall, E. (N.) hasseltii Doleschall, N. maculata var. annulipes Thorell, N. maculata jalorensis Simon, N. maculata var. novae-guineae Strand, N. pictithorax Kulczyński, N. maculata var. flavornata Merian, N. pictithorax Kulczyński, N. maculata var. flavornata Merian, N. maculata piscatorum de Vis, and N. (N.) maculata var. lauterbachi Dahl are proposed as new synonyms of N. pilipes. Nephila imperialis var. novaemecklenburgiae Strand, N. ambigua Kulczyński, N. sarasinorum Merian and N. celebesiana Strand are proposed as new synonyms of N. antipodiana. Meta aerea Hogg, N. meridionalis Hogg, N. adelaidensis Hogg and N. meridionalis hermitis Hogg are proposed as new synonyms of N. edulis. Nephila picta Rainbow is removed from the synonymy of N. plumipes and treated as a synonym of N. edulis, and N. nigritarsis insulicola Pocock is removed from the synonymy of N. plumipes and treated as a synonym of N. antipodiana. Allozyme data demonstrate that N. pilipes is distinct at the 80% FD level from N. edulis, N. plumipes and N. tetragnathoides. Nephila plumipes and N. tetragnathoides, deemed to represent sister-taxa owing to the shared presence of a triangular protrusion of the male pedipalpal conductor, were found to differ at 15% FD in the genetic study. No genetic differentiation was found between 10 populations of N. edulis sampled across mainland Australia. Species of the genus Nephila have been extensively used in ecological and behavioural studies, and the biology of Nephila species in the Australasian region is extensively reviewed and compared with studies on Nephila species from other regions of the world.

Male mate choice and patterns of paternity in the polyandrous, sexually cannibalistic orb-web spider, Nephila plumipes

Studies that investigate patterns of paternity in polyandrous species typically employ double-mating trials, in which the paternity share of each male is established by either the sterile male technique or using genetic markers. However, polyandrous females may mate with more than two males and, in some species, triple-mating trials produce different patterns of paternity from double-mating trials. We investigated patterns of paternity share in triple-mating trials of the sexually cannibalistic orb-web spider Nephila plumipes. These experiments reveal little quantitative changes to paternity share when more than two males mate with the female; the third male apparently diluted the fertilisation success of the second male but not of the first male. Sexual cannibalism had little impact on the fertilisation success of the first male, but greatly increased the fertilisation success of the third male. When offered a choice, males did not prefer to mate with virgin over mated females, but males that chose virgin females were significantly heavier than those that chose mated females.

Frequency, composition and variation in external food stores constructed by orb-web spiders: Nephila edulis and Nephila plumipes (Araneae : Araneoidea)

Few examples of food hoarding have been documented in spiders, yet two Australian species of orb-web spiders, Nephila edulis and N. plumipes, typically incorporate previously captured prey into the web. The effect of prey density and prey-encounter rate on the storage behaviour of adult female N. edulis was tested in the laboratory. Prey density had a significant effect on the propensity to construct external caches: when more food was available, food caches were larger than when the supply of prey was limited. Caching behaviour also differed with the rate of prey encounter, even though the total amount of food supplied was the same. When prey were encountered at constant rates, spiders allocated more food to external storage compared with random encounter rates. Finally, we tested the quality of different prey types for external or internal storage. N. edulis were fed with blow-flies or crickets, and these prey were stored in the web, discarded or totally consumed. Crickets were typically consumed or stored, while flies were more frequently discarded. Field observations of the storage behaviour in N. edulis and N. plumipes found surprising differences in the composition of the cache. While N. plumipes incorporated only animal material, N. edulis also utilised plant material, suggesting that the storage band in N. edulis has other, non-food-storing functions. Field experiments indicated that the presence or absence of external stores in the web of N. plumipes had no influence on mortality, weight gain, or the presence of Argyrodes kleptoparasites.