Allometry and sexual selection of male weaponry in Wellington tree weta, Hemideina crassidens

Both male and female Wellington tree weta, Hemideina crassidens,use cavities in trees as diurnal shelters. That these galleriesare often limiting in nature offers males the opportunity toincrease their reproductive success by monopolizing galleriesand the females residing in them. Male H. crassidens, can matureat either the 8th, 9th, or 10th instar, whereas females matureat the 10th instar only, and male head (and mandible) size positivelycovaries with ultimate instar number. It has been suggestedthat males fight for control of galleries by using their enlargedmandibles as weapons, and males with larger mandibles controlgalleries with more females. In the present study, I presenta statistical examination of sexual dimorphism, showing thattraits related to head size are on average significantly largerin males, whereas traits related to body size are on averagesignificantly larger in females. I tested three predictionsaddressing the hypothesis that sexual selection is driving megacephalyin male H. crassidens. First, as predicted, traits related tohead size show a positive allometric relationship with bodysize in males but not in females. Second, adapting a novel statisticaltechnique based on maximum likelihood and bootstrapping revealedthat males, but not females, exhibit a multimodal distributionin head and body size traits. This is likely a consequence ofmales maturing at one of three instars, which results in positivecovariance between the ultimate instar number and morphologicaltraits. Third, as predicted, single adult males with largerheads reside in galleries housing larger groups of adult females.     

Endemism and sexual systems in the evergreen tree flora of the Western Ghats, India

This study reports the endemism and sexual system in the wet evergreen tree flora of the Western Ghats. A total of 656 species from 66 families and 231 genera were listed. This included a gymnosperm family (Podocarpaceae) and a monocot family (Arecaceae). No family was endemic to the Western Ghats, but 352 species (53%) from 43 families and five genera were endemic. Nearly 35% of the families had no endemics. The largest families with endemics were Dipterocarpaceae (92%), Anacardiaceae (84%), Lauraceae (72%), Fabaceae, Rubiaceae and Myrtaceae (68%). The top five contributing families in the tree flora of the Western Ghats were Euphorbiaceae, Lauraceae, Rubiaceae, Myrtaceae and Annonaceae. The 656 species were largely hermaphrodites (57%) followed by dioecious (20%), polygamous (16%) and monoecious species (5%). The rate of dioecy reported in this study (20%) is higher than reports for Puerto Rico (18%) but lower than the Malaysian rainforest (26%). Structurally, like the Neotropical forests, most evergreen forest types of the Western Ghats could be classified into four ensembles. Yet, the Western Ghats had fewer species than other tropical and Neotropical forests. The proportion of endemics in the ensembles of the Western Ghats ranged from 34% (ensemble IV) to 14% (ensemble I).     

Repeatability of extra-pair mating in tree swallows

Models of sexual selection assume that female mating preferences are heritable and, thus, repeatable for individual females across multiple mating episodes. Previous studies of the repeatability of female preference have examined individuals in captivity and focused presumably on social mate choice. However, extra-pair mating is widespread and can also influence sexual selection. We examined the repeatability of extra-pair mating in a wild population of tree swallows (Tachycineta bicolor) by experimentally inducing females to lay two clutches in rapid succession within the same season. We found that the proportion of extra-pair young and the number of extra-pair sires were highly repeatable for individual females. However, the repeatability of specific extra-pair sires was low. We suggest that this unusual pattern of mating may be due to females maximizing the heterozygosity of their offspring.     

When, how and how much: Gender-specific resource-use strategies in the dioecious tree Juniperus thurifera

BACKGROUND AND AIMS: In dioecious species male and female plants experience different selective pressures and often incur different reproductive costs. An increase in reproductive investment habitually results in a reduction of the resources available to other demands, such as vegetative growth. Tree-ring growth is an integrative measure that tracks vegetative investment through the plant's entire life span. This allows the study of gender-specific vegetative allocation strategies in dioecious tree species thoughout their life stages. METHODS: Standard dendrochronological procedures were used to measure tree-ring width. Analyses of time-series were made by means of General Mixed Models with correction of autocorrelated values by the use of an autoregressive covariance structure of order one. Bootstrapped correlation functions were used to study the relationship between climate and tree-ring width. KEY RESULTS: Male and female trees invest a similar amount of resources to ring growth during the early life stages of Juniperus thurifera. However, after reaching sexual maturity, tree-ring growth is reduced for both sexes. Furthermore, females experience a significantly stronger reduction in growth than males, which indicates a lower vegetative allocation in females. In addition, growth was positively correlated with precipitation from the current winter and spring in male trees but only to current spring precipitation in females. CONCLUSIONS: Once sexual maturity is achieved, tree rings grow proportionally more in males than in females. Differences in tree-ring growth between the genders could be a strategy to respond to different reproductive demands. Therefore, and responding to the questions of when, how and how much asked in the title, it is shown that male trees invest more resources to growth than female trees only after reaching sexual maturity, and they use these resources in a different temporal way.     

Sexually dimorphic effect of mating on the melanotic encapsulation response in the harem‐defending Wellington tree weta,Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae)

Reproductive activities are generally costly to immune responsiveness because limited resources required by reproduction are diverted away from immunity (and vice versa). Reproduction, however, is not expected to affect the immune response in males and females similarly as mating is expected to negatively affect male immunity more so than female immunity. Here, I test the phenotypic plasticity hypothesis in the Wellington tree weta (Hemideina crassidens), a sexually dimorphic orthopteran insect that is endemic to New Zealand. My laboratory experiment showed that although males had higher rates of melanotic encapsulation than females, contrary to prediction, females were the only sex significantly affected by mating and the effect was positive. In addition to immunity differing between the sexes, immune function can differ intrasexually, particularly when males are polymorphic and different investment strategies are used to maximize fitness. Male H. crassidens exhibit alternative mating strategies that are represented by three different morphotypes. I therefore explored whether the morphs differed in their melanotic encapsulation response and whether mating affected the morphs differently. I found no difference among morphs or an effect of mating on male immune response.     

Female remating and the intensity of female choice in black-horned tree crickets, Oecanthus nigricornis

Repeated mating by females of many species occurs at frequenciesin excess of those needed to acquire additional sperm for fertilizingova. I tested three alternative hypotheses for the rate of rematingby females of the courtship-feeding tree cricket, Oecanthusnigricornis Walker, by manipulating diet quality and courtshipfeeding and measuring the time to remating by the female inrelation to four aspects of male phenotype (age, condition,fluctuating asymmetry, and size). First, in courtship-feedingspecies, remating may be due to selection to increase the amountof nutritional resources provided by males, with nutrient-deprivedfemales remating more quickly. Second, remating may functionas a mechanism of postcopulatory mate choice, with females rematingquickly when the quality of a previous mate is low. Third, quicknessof remating may be the consequence of precopulatory mate choiceprior to future matings, with females remating more quicklywith high-quality males, regardless of the quality of priormates. Females on a low-quality diet remated quickly, did notvary remating speed with the phenotype of their first mate,and did not differentially reject prospective second mates withdifferent phenotypes. In contrast, both the degree of coyness(measured as the frequency of mate rejection) and the intensityof female choice (measured as the size differential betweenaccepted and rejected mates) increased with diet quality. Theseresults support both the material-benefits and the precopulatorymate-choice hypotheses for remating speed of female tree crickets.There was mixed support for the postcopulatory choice hypothesis:females on the high-quality diet remated more slowly after firstmating with relatively large males, in support of the postcopulatorychoice hypothesis; however, the remating interval of femaleson the high-quality diet decreased with the condition of thefirst mate, opposite to the prediction of the postcopulatorychoice hypothesis     

Signal diversification in Oecanthus tree crickets is shaped by energetic,morphometric, and acoustic trade‐offs

Physiology, physics, and ecological interactions can generate trade‐offs within species, but may also shape divergence among species. We tested whether signal divergence in Oecanthus tree crickets is shaped by acoustic, energetic, and behavioral trade‐offs. We found that species with faster pulse rates, produced by opening and closing wings up to twice as many times per second, did not have higher metabolic costs of calling. The relatively constant energetic cost across species is explained by trade‐offs between the duration and repetition rate of acoustic signals—species with fewer stridulatory teeth closed their wings more frequently such that the number of teeth struck per second of calling and the resulting duty cycle were relatively constant across species. Further trade‐offs were evident in relationships between signals and body size. Calling was relatively inexpensive for small males, permitting them to call for much of the night, but at low amplitude. Large males produced much louder calls, reaching up to four times more area, but the energetic costs increased substantially with increasing size and the time spent calling dropped to only 20% of the night. These trade‐offs indicate that the trait combinations that arise in these species represent a limited subset of conceivable trait combinations.     

Sex ratios, size distributions, and sexual dimorphism in the dioecious tree Ilex aquifolium (Aquifoliaceae)

Sex ratio and sexual dimorphism in physiology and growth were studied in the dioecious tree Ilex aquifolium at two localities in northern Spain. Genet sex ratio was significantly male biased in one locality but not in the other. However, ramet and flowering ramet sex ratios were male biased at both study sites. Males had significantly thicker main trunks than females in one locality and produced more ramets in the other. Growth rate, estimated from mean width of annual rings, did not differ between localities, but males produced wider rings than females at both sites. Mean annual growth rates over the last 10, 20, and 30 yr were significantly higher for males. Measurements of chlorophyll fluorescence indicated that the efficiency of photosynthesis of leaves on nonfruiting branches of females was higher than for leaves on branches of male plants under low-light conditions, though not under saturating-light conditions. Efficiency of photosynthesis was significantly lower on fruiting branches of female plants than on nonfruiting branches. We discuss whether the observed between-sex differences are attributable to the higher cost of reproduction in females and/or to pollen competition.     

Geographic parthenogenesis and the common tea‐tree stick insect of New Zealand

Worldwide, parthenogenetic reproduction has evolved many times in the stick insects (Phasmatidae). Many parthenogenetic stick insects show the distribution pattern known as geographic parthenogenesis, in that they occupy habitats that are at higher altitude or latitude compared with their sexual relatives. Although it is often assumed that, in the short term, parthenogenetic populations will have a reproductive advantage over sexual populations; this is not necessarily the case. We present data on the distribution and evolutionary relationships of sexual and asexual populations of the New Zealand stick insect, Clitarchus hookeri. Males are common in the northern half of the species’ range but rare or absent elsewhere, and we found that most C. hookeri from putative‐parthenogenetic populations share a common ancestor. Female stick insects from bisexual populations of Clitarchus hookeri are capable of parthenogenetic reproduction, but those insects from putative‐parthenogenetic populations produced few offspring via sexual reproduction when males were available. We found similar fertility (hatching success) in mated and virgin females. Mated females produce equal numbers of male and female offspring, with most hatching about 9–16 weeks after laying. In contrast, most eggs from unmated females took longer to hatch (21–23 weeks), and most offspring were female. It appears that all C. hookeri females are capable of parthenogenetic reproduction, and thus could benefit from the numerical advantage this yields. Nevertheless, our phylogeographic evidence shows that the majority of all‐female populations over a wide geographic area originate from a single loss of sexual reproduction.     

Floral development in the androdioecious tree Tapiscia sinensis: Implications for the evolution to androdioecy

The evolutionary pathway between hermaphroditism and dioecy (females and males in a single population) draws widespread interests, and androdioecy (bisexuals and males in a single population) is rarely achieved as an intermediate state between the two breeding systems. Flower bud differentiations in the pistils of hermaphrodites and the pistillodes of males in androdioecious Tapiscia sinensis Oliv. are investigated by routine paraffin section technology, light microscopy, and scanning electron microscopy. A phylogenetic approach is used to analyze the origin of androdioecy. In T. sinensis, hermaphroditic flowers (HF) and male flowers (MF) experienced a similar development pattern in early flower bud differentiation, including the initiation of tepals and stamens. However, the carpel differentiation of MF and HF proceed in different patterns. In HF, the central zone bulges out and produces a ring meristem on which two to three carpel primordia emerge, which eventually developed into a normal pistil with a stigma, a style, and an ovary. However, in most MF, vestigial pistils are stem‐like (type I), and very few have an empty ovary (type II) or a sterile ovule (type III). Moreover, the evolution of sexual systems within the Huerteales indicates that hermaphroditism is the primitive character of T. sinensis. Tapiscia sinensis shows different degrees of reduction between male flowers and bisexual ones in the evolution to dioecy. Functional androdioecy originated from a hermaphroditic ancestor in T. sinensis and, as an intermediate sexual system, involves evolution from hermaphrodites to dioecy.     

Morphological variation in the flowers of Jacaratia mexicana A. DC. (Caricaceae), a subdioecious tree

The Caricaceae is a small family of tropical trees and herbs in which most species are dioecious. In the present study, we extend our previous work on dioecy in the Caricaceae, characterising the morphological variation in sexual expression in flowers of the dioecious tree Jacaratia mexicana . We found that, in J. mexicana , female plants produce only pistillate flowers, while male plants are sexually variable and can bear three different types of flowers: staminate, pistillate and perfect. To characterise the distinct types of flowers, we measured 26 morphological variables. Our results indicate that: (i) pistillate flowers from male trees carry healthy-looking ovules and are morphologically similar, although smaller than, pistillate flowers on female plants; (ii) staminate flowers have a rudimentary, non-functional pistil and are the only flowers capable of producing nectar; and (iii) perfect flowers produce healthy-looking ovules and pollen, but have smaller ovaries than pistillate flowers and fewer anthers than staminate flowers, and do not produce nectar. The restriction of sexual variation to male trees is consistent with the evolutionary path of dioecy from hermaphrodite ancestors through the initial invasion of male-sterile plants and a subsequent gradual reduction in female fertility in cosexual individuals (gynodioecy pathway), but further work is needed to confirm this hypothesis.     

Baffling: a condition-dependent alternative mate attraction strategy using self-made tools in tree crickets

Intense sexual selection in the form of mate choice can facilitate the evolution of different alternative reproductive strategies, which can be condition-dependent. Tree cricket males produce long-distance acoustic signals which are used by conspecific females for mate localization and mate choice. Our study shows that baffling, an acoustic call amplification strategy employed by male tree crickets using self-made tools, is a classic example of a condition-dependent alternative strategy. We show that though most males can baffle, less preferred males, such as smaller and lower-amplitude callers, predominantly use this alternative strategy. Baffling allows these males to increase their call amplitude and advertisement range, which attracts a higher number of females. Baffling also gives these males a mating benefit because females mate for longer durations with them. Our results suggest that the advantage of baffling in terms of gain in the number of sperm cells transferred while mating is primarily limited to less preferred males, thus maintaining the polymorphism of calling strategies in the population. We summarize that baffling is a condition-dependent strategy used by less preferred tree cricket males to obtain mating benefits.     

Sexually dimorphic immune response in the harem polygynous Wellington tree weta Hemideina crassidens

Abstract Adult males are often less immunocompetent than females. One explanation for this is that intense sexual selection causes males to trade‐off investment in immunity with traits that increase mating success. This hypothesis is tested in the Wellington tree weta (Hemideina crassidens), a large, sexually dimorphic orthopteran insect in which males possess enormous mandibular weaponry used during fights for access to female mates. Field‐collected males have a significantly greater immune response (greater melanotic encapsulation) than females, suggesting that body condition, longevity or an allied trait is important to male fitness, or that females require materials for egg production that would otherwise be used to boost immunity. Although immunity is expected to trade‐off against reproductive traits in both sexes, there is no significant relationship between immune response and weapon or testes size in males, nor fecundity in females.     

SEX-LINKAGE OF SEXUALLY ANTAGONISTIC GENES IS PREDICTED BY FEMALE, BUT NOT MALE, EFFECTS IN BIRDS

Evolutionary theory predicts that sexually antagonistic loci will be preferentially sex-linked, and this association can be empirically testes with data on sex-biased gene expression with the assumption that sex-biased gene expression represents the resolution of past sexual antagonism. However, incomplete dosage compensating mechanisms and meiotic sex chromosome inactivation have hampered efforts to connect expression data to theoretical predictions regarding the genomic distribution of sexually antagonistic loci in a variety of animals. Here we use data on the underlying regulatory mechanism that produce expression sex-bias to test the genomic distribution of sexually antagonistic genes in chicken. Using this approach, which is free from problems associated with the lack of dosage compensation in birds, we show that female-detriment genes are significantly overrepresented on the Z chromosome, and female-benefit genes underrepresented. By contrast, male-effect genes show no over- or underrepresentation on the Z chromosome. These data are consistent with a dominant mode of inheritance for sexually antagonistic genes, in which male-benefit coding mutations are more likely to be fixed on the Z due to stronger male-specific selective pressures. After fixation of male-benefit alleles, regulatory changes in females evolve to minimize antagonism by reducing female expression.     

Genetic benefits of a female mating preference in gray tree frogs are context-dependent

Abstract "Good genes" models of sexual selection predict that male courtship displays can advertise genetic quality and that, by mating with males with extreme displays, females can obtain genetic benefits for their offspring. However, because the relative performance of different genotypes can vary across environments, these genetic benefits may depend on the environmental context; in which case, static mating preferences may not be adaptive. To better understand how selection acts on the preference that female gray tree frogs ( Hyla versicolor ) express for long advertisement calls, I tested for genetic benefits in two realistic natural environments, by comparing the performance of half-sibling offspring sired by males with long versus short calls. Tadpoles from twelve such maternal half-sibships were raised in enclosures in their natal pond at two densities. In the low-density treatment, offspring of long-call males were larger at metamorphosis than were offspring of short-call males, whereas in the high-density treatment, offspring of males with long calls tended to metamorphose later than offspring of males with short calls. Thus, although the genes indicated by long calls were advantageous under low-density conditions, they were not beneficial under all conditions, suggesting that a static preference for long calls may not be adaptive in all environments. Such a genotype-by-environment interaction in the genetic consequences of mate choice predicts that when the environment is variable, selection may favor plasticity in female preferences or female selectivity among environments to control the conditions experienced by the offspring.     

The geography of sex‐specific selection,local adaptation,and sexual dimorphism

Local adaptation and sexual dimorphism are iconic evolutionary scenarios of intraspecific adaptive differentiation in the face of gene flow. Although theory has traditionally considered local adaptation and sexual dimorphism as conceptually distinct processes, emerging data suggest that they often act concurrently during evolutionary diversification. Here, I merge theories of local adaptation in space and sex‐specific adaptation over time, and show that their confluence yields several new predictions about the roles of context‐specific selection, migration, and genetic correlations, in adaptive diversification. I specifically revisit two influential predictions from classical studies of clinal adaptation and sexual dimorphism: (1) that local adaptation should decrease with distance from the species’ range center and (2) that opposing directional selection between the sexes (sexual antagonism) should inevitably accompany the evolution of sexual dimorphism. I show that both predictions can break down under clinally varying selection. First, the geography of local adaptation can be sexually dimorphic, with locations of relatively high local adaptation differing profoundly between the sexes. Second, the intensity of sexual antagonism varies across the species’ range, with subpopulations near the range center representing hotspots for antagonistic selection. The results highlight the context‐dependent roles of migration versus sexual conflict as primary constraints to adaptive diversification.     

Enhanced expression of tubulin-specific chaperone protein A, mitochondrial ribosomal protein S27, and the DNA excision repair protein XPACCH in the song system of juvenile male zebra finches

Recent evidence suggests that sexual dimorphisms in the zebra finch song system and behavior arise due to factors intrinsic to the brain, rather than being solely organized by circulating steroid hormones. The present study examined expression of 10 sex chromosome genes in the song system of 25-day-old zebra finches in an attempt to further elucidate these factors. Increased expression in males was confirmed for nine of the genes by real-time qPCR using cDNA from individual whole telecephalons. In situ hybridization at the same age revealed specific, male-enhanced mRNA for three of the nine genes in one or more song control nuclei. These genes encode tubulin-specific chaperone A, mitochondrial ribosomal protein S27, and a DNA repair protein XPACCH. Based on what is currently known about these proteins' functions and their localization to particular components of the song circuit, we hypothesize that they each may be involved in specific aspects of masculinization.     

Perspective: female remating,operational sex ratio,and the arena of sexual selection in Drosophila species

Abstract.— As commonly observed among closely related species within a variety of taxa, Drosophila species differ considerably in whether they exhibit sexual dimorphism in coloration or morphology. Those Drosophila species in which male external sexual characters are minimal or absent tend, instead, to have exaggerated ejaculate traits such as sperm gigantism or seminal nutrient donations. Underlying explanations for the interspecific differences in the presence of external morphological sexual dimorphism versus exaggerated ejaculate traits are addressed here by examining the opportunity for sexual selection on males to occur before versus after mating in 21 species of Drosophila . Female remating frequency, an important component of the operational sex ratio, differs widely among Drosophila species and appears to dictate whether the arena of sexual selection is prior to, as opposed to after, copulation. Infrequent female mating results in fewer mating opportunities for males and thus stronger competition for receptive females that favors the evolution of male characters that maximize mating success. On the other hand, rapid female remating results in overlapping ejaculates in the female reproductive tract, such that ejaculate traits which enhance fertilization success are favored. The strong association between female remating frequency in a given species and the presence of sexually selected external versus internal male characters indicates that the relationship be examined in other taxa as well.