Reproductive biology of harvestmen(Arachnida:Opiliones):a review of a rapidly evolving research field

Harvestmen are a major arachnid order that has experienced a dramatic increase in biological knowledge in the 21st century.The publication of the book Harvestmen:The Biology of Opiliones in 2007 stimulated the development of many behavioral studies.Although the book is relatively recent,our understanding of the reproductive biology of harvestmen is already outdated due to the fast accumulation of new data.Our goal is to provide an updated review of the subject to serve as a benchmark for the following years.In the pre-copulatory phase,we explore the evolu-tion of facultative parthenogenesis,the factors that may affect the types of mating system,and the role of nuptial gifts in courtship.Regarding the copulatory phase,harvestmen are unique arachnids because they have aflagellate spermatozoa and a penis with complex morphology.We discuss the implications of these two features for sperm competition and cryptic female choice.In the post-copulatory phase,we connect ovi-position site selection and climate conditions to the widespread occurrence of resource defense polygyny,alternative reproductive tactics,and sexual dimorphism in several clades of tropical harvestmen.Finally,we present the different forms of parental care in the order,and discuss the benefits and costs of this behavior,which can be performed either by females or males.Throughout the review,we indicate gaps in our knowl-edge and subjects that deserve further studies.Hopefully,the information synthesized here will stimulate researchers worldwide to embrace harvestmen as a study system and to improve our effort to unravel the mysteries of their reproductive biology.

Male nutritional status does not impact the reproductive potential of female Cnaphalocrocis medinalis moths under conditions of nutrient shortage

In addition to sperm,some accessory substances transferred to females during copulation act as nuptial gifts by passing on valuable nutrients in many insect species.The nutritional status of the males can thus have a great effect on the mating behavior,fecundity and even the longevity of females.However,little is known about the effect of male nutritional status on the female reproductive traits in migratory insect species,particularly when females experience nutrient shortage and have to choose between reproduction and migration.Here,Cnaphalocrocis medinalis,a migratory rice pest in Asia,was studied to explore this issue.Our results showed that in male moths fed with honey solution,their gonads had higher energy content than gonads of starved males,resulting in increased energy content of the bursa copulatrix of females after mating with fed males.Such females showed increased mating frequency,fecundity and longevity compared to females mating with starved males,indicating that male moths deliver nuptial gifts to females and improve their reproductive performance.However,when females were starved,only about 45%mated,with just a single copulation,regardless of male nutritional status.Starved females showed lower fecundity,and a longer pre-oviposition period(indicating a greater propensity to migrate),compared to fed females.However,copulation still significantly extended their longevity.These results suggest that starved females invest in migration to escape deteriorating habitats,rather than investing the nuptial gift to increased fecundity.Our results further our understanding of the reproductive adaptability of migratory insects under conditions of food stress.

Evolution of deceit by worthless donations in a nuptial gift-giving spider

Males of the nursery web spider Pisaura mirabil usually offer an insect prey wrapped in white silk as a nuptial gift to facilitate copulation. Males exploit female foraging preferences in a sexual context as females feed on the gift during copula- tion. It is possible for males to copulate without a gift, however strong female preference for the gift leads to dramatically higher mating success for gift-giving males. Females are polyandrous, and gift-giving males achieve higher mating success, longer copulations, and increased sperm transfer that confer advantages in sperm competition. Intriguingly, field studies show that ap- proximately one third of males carry a worthless gift consisting of dry and empty insect exoskeletons or plant fragments wrapped in white silk. Silk wrapping disguises gift content and females are able to disclose gift content only after accepting and feeding on the gift, meanwhile males succeed in transferring sperm. The evolution of deceit by worthless gift donation may be favoured by strong intra-sexual competition and costs of gift-construction including prey capture, lost foraging opportunities and investment in silk wrapping. Females that receive empty worthless gifts terminate copulation sooner, which reduces sperm transfer and likely disadvantages males in sperm competition. The gift-giving trait may thus become a target of sexually antagonistic co-evolution, where deceit by worthless gifts leads to female resistance to the trait. We discuss factors such as female mating rate and intensity of sperm competition that may shape the evolution of male deception, and how ecological factors may influence the evolution and maintenance of worthless gifts as an evolutionarily stable alternative mating strategy by frequency dependent selection

The evolution of multi-component visual signals in darters （genus Etheostoma）

As complex traits evolve, each component of the trait may be under different selection pressures and could respond independently to distinct evolutionary forces. We used comparative methods to examine patterns of evolution in multiple components of a complex courtship signal in darters, specifically addressing the question of how nuptial coloration evolves across different areas of the body. Using spectral reflectance, we defined 4 broad color classes present on the body and fins of 17 species of freshwater fishes （genus Etheostoma） and quantified differences in hue within each color class. Ancestral state reconstruction suggests that most color traits were expressed in the most recent common ancestor of sampled species and that differences among species are mostly due to losses in coloration. The evolutionary lability of coloration varied across body regions; we found sig- nificant phylogenetic signal for orange color on the body but not for most colors on fins. Finally, patterns of color evolution and hue Of the colors were correlated among the two dorsal fins and between the anterior dorsal and anal fins, but not between any of the fins and the body. The observed patterns support the hypothesis that different components of complex signals may be subject to distinct evolutionary pressures, and suggests that the combination of behavioral displays and morphology in communication may have a strong influence on patterns of signal evolution .

Patterns of color phase indicate spawn timing at a Nassau grouper Epinephelus striatus spawning aggregation

Nassau grouper Epinephelus striatus are a large bodied, top level predator that is ecologically important throughout the Caribbean. Although typically solitary, Nassau grouper form large annual spawning aggregations at predictable times in spe- cific locations. In 2003, The Cayman Islands Marine Conservation Board established protection for a newly rediscovered Nassau grouper spawning aggregation on Little Cayman, British West Indies. The large size of this aggregation provides a unique oppor- tunity to study the behavior of Nassau grouper on a relatively intact spawning aggregation. During non-spawning periods Nassau grouper display a reddish-brown-and-white barred coloration. However, while aggregating they exhibit three additional color phases： ＂bicolor＂, ＂dark＂, and ＂white belly＂. We video sampled the population on multiple days leading up to spawning across five spawning years. Divers focused a laser caliper equipped video camera on individual fish at the aggregation. We later ana- lyzed the video to determine the length of the fish and record the color phase. Our observations show that the relative proportion of fish in the bicolor color phase increases significantly on the day leading up to the primary night of spawning. The increase in the proportion of the bicolor color phase from 0.05 early in the aggregation to 0.40 on the day of spawning suggests that this color phase conveys that a fish is behaviorally and physiologically prepared to spawn. Additionally, 82.7% of fish exhibiting dark or white belly coloration early in the aggregation period suggests that these color phases are not only shown by female fish as was previously posited [Current Zoology 58 （1）： 73-83, 2012].