Reproductive investment in moa: a K-selected life-history strategy?

The exaggerated K-selected life-history strategy of moa has been suggested as an important factor causing their rapid extinction. Classically, this strategy is characterized by few, large offspring and low fecundity rates. Assuming clutches with one or two eggs as derived from the fossil record, we tested if eggs of moa were larger than the average of similar-sized birds, and estimated their unknown annual breeding frequencies. Therefore, we established allometries on body mass and different reproductive traits (i.e. egg mass, clutch mass and annual clutch mass). These were derived for r-selected (r-model) and for K-selected (K-model) bird species. In agreement with our initial expectations, moa had egg to body mass relations seen in “average” extant K-selected birds. While the K-model pointed to a clutch size of one or two eggs for moa corroborating fossil data and a K-selected life-history, the r-model predicted two to three times larger sizes. Under clutch sizes between one and two eggs and an annual clutch mass as observed in other similar-sized flightless island birds (e.g. rails, ratites), the annual clutch mass allometry suggested one to three clutches per year for moa. Even when assuming less than one brood per year (K-model predicts 0.5 clutches per year); annual clutch masses were still consistent with the K-model. Further studies are needed to clarify whether or not the reproductive strategy of flightless island birds and/or of the birds underlying the K-model fits better to the moa strategy. The approach presented herein, illustrates that combining biological and paleontological data can assist in the reconstruction of species traits, which are insufficiently or not preserved in fossils, but are necessary to understand the evolution of traits.     

Sex allocation and nestling survival in a dimorphic raptor: does size matter?

Fisher's theory predicts equal sex ratios at the end of parentalcare if the costs and benefits associated with raising eachsex of offspring are equal. In raptors, which display variousdegrees of reversed sexual size dimorphism (RSD; females thelarger sex), sex ratios biased in favor of smaller males areonly infrequently reported. This suggests that offspring ofeach sex may confer different fitness advantages to parents.We examined the relative returns associated with raising eachsex of offspring of the brown falcon Falco berigora, a medium-sizedfalcon exhibiting RSD (males approximately 75% of female bodymass) and subsequent sex ratios. Female nestlings hatched eitherfirst or second did not receive more food nor did they hatchfrom larger eggs or remain dependent on parents for longer periodsthan male offspring from these hatch orders. Together with previousstudies this result indicates that even in markedly dimorphicspecies, the required investment to raise the larger sex islikely to be less than that predicted by body size differencesalone. Moreover, among last-hatched nestlings, both sexes faceda reduced food allocation and suffered a slower growth rateand thus final body size, with a concurrent increased probabilityof mortality. For last-hatched females the reduction in foodallocation was more marked, with complete mortality of all last-hatchedfemale nestlings monitored in this study. Once independent,males of any size but only larger females are likely to be recruitedinto the breeding population. The sex-biased food allocationamong last-hatched offspring favoring males thus reflects therelative returns to parents in raising a small member of eachsex.     

Reproductive effort in clonal plants: constant allocation ratios among ramets?

Summary Armstrong (1982, 1983) predicted that all ramets within a clone should have the same ratio of biomass allocation to sexual reproduction versus vegetative growth. He presented data (1984) that he interpreted as showing that Solidago altissima ramets in a clone do have the predicted constant allocation ratio. Reanalysis of his methods shows that this conclusion was an artifact of his analysis. A simulation using random numbers and Armstrong's analysis showed the same pattern as his data. Data from S. altissima ramets of a single clone grown in a greenhouse experiment, using a different analysis, illustrated that the allocation ratios within a clone can be highly variable.     

Reproductive suppression in female Damaraland mole-rats Cryptomys damarensis: dominant control or self-restraint?

Colonies of Damaraland mole-rats Cryptomys damarensis exhibit a high reproductive skew. Typically one female breeds and the others are anovulatory. Two models, the dominant control model (DCM) and the self-restraint model (SRM), have been proposed to account for this reproductive suppression. The DCM proposes that suppression is under the control of the dominant breeder and is imposed by mechanisms such as aggression, pheromones and interference with copulation, whereas the SRM does not involve aggression directed towards non-breeders and may function in order to minimize inbreeding. We investigated potential proximate mechanisms involved in the suppression of females in a series of experiments. Socially induced stress through aggression did not appear to be responsible for anovulation. Nor did breeders actively interfere with subordinate copulation. Females were physiologically suppressed when housed in intact colonies. However, as predicted by the DCM, they did not become reproductively active when removed from the presence of breeders. We found no evidence that pheromonal cues block ovulation. We suggest that the SRM is the basic model found in the Damaraland mole-rat and that self-restraint functions in order to minimize inbreeding by restricting reproduction until an unrelated male is present. This would explain the rapid onset of reproductive activation in females when paired with an unrelated male, as demonstrated in this study.     

Intraclutch egg‐size variation in acanthosomatid bugs: adaptive allocation of maternal investment?

If there are differences in predation risk among the offspring within a clutch, parents may allocate less resources to the offspring facing higher risk. I examined parental investment in terms of egg size within clutches in five species of stink bugs (Heteroptera, Acanthosomatidae). In subsocial Elasmucha and Sastragala species, the female guards her eggs and first-instar nymphs against invertebrate predators by covering her clutch with her body. Large differences in survival from predation between offspring at the centre and offspring at the periphery of the clutch have been reported in such subsocial insects. I found that Elasmucha and Sastragala females laid significantly smaller eggs in the peripheral (and thus more vulnerable) part of the clutch. Phenotypic trade-offs between egg size and clutch size were detected in these subsocial species. Egg size was positively correlated with hatched first-instar nymph size: smaller nymphs hatched from smaller peripheral eggs. In asocial Elasmostethus humeralis , however, no significant difference in size was detected between the eggs at the centre of and those at the periphery of the clutch. Thus, in subsocial acanthosomatid bugs, females seem to allocate their resources according to the different predation risks faced by the offspring within the clutch.     

Reproductive skew in female common marmosets: what can proximate mechanisms tell us about ultimate causes?

Common marmosets are cooperatively breeding monkeys that exhibit high reproductive skew: most subordinate females fail to reproduce, while others attempt to breed but produce very few surviving infants. An extensive dataset on the mechanisms limiting reproduction in laboratory-housed and free living subordinate females provides unique insights into the causes of reproductive skew. Non-breeding adult females undergo suppression of ovulation and inhibition of sexual behaviour; however, they receive little or no aggression or mating interference by dominants and do not exhibit behavioural or physiological signs of stress. Breeding subordinate females receive comparable amounts of aggression to non-breeding females but are able to conceive, gestate and lactate normally. In groups containing two breeding females,however, both dominant and subordinate breeders kill one another's infants. These findings suggest that preconception reproductive suppression is not imposed on subordinate females by dominants, at a proximate level, but is instead self-imposed by most subordinates, consistent with restraint models of reproductive skew. In contrast to restraint models, however, this self-suppression probably evolved not in response to the threat of eviction by dominant females but in response to the threat of infanticide. Thus,reproductive skew in this species appears to be generated predominantly by subordinate self-restraint, in a proximate sense, but ultimately by dominant control over subordinates' reproductive attempts.     

Resource allocation: a conflict in the fig/fig wasp mutualism?

Conflicts of interest are omnipresent between mutualist species. In the monoecious fig/pollinator wasp mutualism, each female flower produces either a seed or a pollinator offspring (which has fed on a single seed). Pollen from a syconium (i.e. fig, a closed urn-shaped inflorescence) is only dispersed by female pollinator offspring born in this syconium. Thus the fig tree is selected to produce both seed and pollinator offspring whereas for the pollinator there is no short term advantage in seed production. Using controlled pollination experiments (pollen injection, and foundress introduction), we show that 1) The relative proportion of seeds and pollinator offspring produced (i.e., the effective allocation between female and male function) depends mainly on the number of foundresses that entered the syconium. 2) Many female flowers within every syconium mature neither a seed nor a wasp (from 25% to 33%). 3) All the female flowers within a syconium that are not vacant at maturity have the potential to produce a seed, and at least 80% of them can produce a pollinator. Several hypotheses concerning mechanisms that govern the partitioning between seed and wasp production are discussed, and their evolutionary consequences are considered.     

Does predation result in adult sex ratio skew in a sexually dimorphic insect genus?

Theory proposes that sexually dimorphic, polygynous species are at particularly high risk of sex-biased predation, because conspicuous males are more often preyed upon compared to females. We tested the effects of predation on population sex ratio in a highly sexually dimorphic insect genus (Hemideina). In addition, introduction of a suite of novel mammalian predators to New Zealand during the last 800 years is likely to have modified selection pressures on native tree weta. We predicted that the balance between natural and sexual selection would be disrupted by the new predator species. We expected to see a sex ratio skew resulting from higher mortality in males with expensive secondary sexual weaponry; combat occurs outside refuge cavities between male tree weta. We took a meta-analytic approach using generalized linear mixed models to compare sex ratio variation in 58 populations for six of the seven species in Hemideina. We investigated adult sex ratio across these populations to determine how much variation in sex ratio can be attributed to sex-biased predation in populations with either low or high number of invasive mammalian predators. Surprisingly, we did not detect any significant deviation from 1 : 1 parity for adult sex ratio and found little difference between populations or species. We conclude that there is little evidence of sex-biased predation by either native or mammalian predators and observed sex ratio skew in individual populations of tree weta is probably an artefact of sampling error. We argue that sex-biased predation may be less prevalent in sexually dimorphic species than previously suspected and emphasize the usefulness of a meta-analytic approach to robustly analyse disparate and heterogeneous data.     

Nuptial gift consumption influences female remating in a scorpionfly: male or female control of mating rate?

In the scorpionfly Panorpa cognata, males provide females with saliva secretions as nuptial food gifts. Consequently, females derive material benefits and possibly also genetic benefits from multiple matings. Females therefore generally should have a high motivation to remate. Males, on the other hand, do not share this interest, which will generate a sexual conflict over remating interval, possibly leading to male adaptations that prevent females from remating with other males. In this study, I found that mated females were less prone to copulate than virgin females, despite female benefits of multiple matings. Further, I found that the remating interval was significantly longer if the first copulation was long compared to shorter matings. This effect does not entirely depend on copulation duration per se, but on the amount of saliva, that a female is consuming during copulation. These results suggest a mating-induced refractory period and can be interpreted as male manipulation of female remating behaviour mediated through substances in the nuptial gift. Alternatively, receiving large nuptial gifts may decrease the prospective direct fitness benefits from further copulations, and thus change optimal female remating rate. Furthermore, gift size has been shown to correlate with male nutritional condition, which may be an indicator of male genetic quality. Females may therefore benefit indirectly by not remating following copulations involving large saliva gifts. In this scenario, female remating interval would be an effect of cryptic female choice.     

When should a female avoid adding eggs to the clutch of another female? A simultaneous oviposition and sex allocation game

Summary Avoidance of double oviposition (ADO) is the strategy not to oviposit on food patches where another female has oviposited before. If two females oviposit on the same patch, competitive and mating interactions within and between broods may lead to both a clutch size game and a sex allocation game between the two visitors. Though the two games interact, they are usually considered separately. Here, the ESS conditions for ADO are investigated in an analysis that combines the two games into one. The analysis strengthens the notion that it is really ADO that needs to be explained, because role-dependent net pay-off from an additional egg is most likely to favour double oviposition (DO). To a first female, the net payoff includes the effect on the eggs already present, whereas to a second female only the egg's gross pay-off matters. ADO is the evolutionary stable strategy (ESS) if there are enough patches still without eggs and either (1) the fitness of an additional egg is so low that the first female would not lay it even in the absence of detrimental effects on earlier offspring, so neither would a second female, or (2) differences in either the survival probability of the offspring or their reproductive success are sufficient to counterbalance the differential interest in the eggs already present. The first condition requires that eggs are relatively large, because then the decrease in pay-off between two successive eggs can be large. The second condition may be met when there is a time interval between ovipositions of subsequent females. The resulting developmental lag of the second clutch will (1) diminish its ability to compete for food and (2) lower its reproductive success when there is local mate competition and sons are too late to mate with daughters of the first female. If sons of first and second females compete on equal terms, however, ADO is unlikely. Male migration between patches reduces the influence of sex allocation strategies on clutch size decisions; the same holds for small clutch sizes. To illustrate the importance of considering sex allocation and clutch size decisions in an integrated way, oviposition strategies of plant-inhabiting predatory mites (Acari: Phytoseiidae) are discussed.     

Parasite transmission by insects: a female affair?

Understanding the relationship between the gender of insects and their ability to act as vectors of insect-borne diseases (IBDs) could provide clues as to the origin of the intimate interplay among insect, pathogen and vertebrate hosts. The vector activity of several species of blood-feeding insects is linked to adult females. Interestingly, the only known exception is the transmission of canine and human thelaziosis by a male dipteran fly. This biological difference raises the question as to whether the parasitic behaviour of male and female insects transmitting IBDs is an expression of a co-evolution of vectors and pathogens.     

A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic

Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8 % (vs. 18.1 % of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male’s territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.     

Reproductive ecology of Ameroglossum pernambucense (Scrophulariaceae): is this ornithophilous and threatened shrub highly adapted to a naturally fragmented habitat?

Many rare plants exist in a naturally fragmented distribution and are expected to exhibit reproductive adaptations to isolation. Thus, understanding the reproduction of these plants might be important in predicting the future of artificially fragmented species. Ameroglossum pernambucense is a threatened ornithophilous shrub with naturally fragmented populations on granite outcrops in north-eastern Brazil. The current research studied two populations of A. pernambucense and determined if three reproductive features (mating system, flowering phenology and nectar dynamics) facilitate the colonisation of new areas and the gene flow among the outcrops. To verify the species’ reproductive efficiency, pollinator efficiency and pollen limitation were tested. Ameroglossum pernambucense has a mixed mating system, and pollinators increased seed production. The pollinator efficiency test revealed that hummingbirds can supply the entire pollen demand. No pollen limitation was detected. Variation in sunset time was the main factor to explain the flowering pattern of A. pernambucense. Nectar was abundantly produced after successive removals. The reproductive traits observed in A. pernambucense were interpreted as highly adapted to its fragmented condition. Phenological photoperiodic regulation may increase the flowering synchrony and chances of gene flow among outcrops in environmentally distinct microsites. The mixed mating system favours both the colonisation of isolated outcrops and allogamy. High flower production and its high energetic value may have an important role in pollinator attraction and may increase long-distance outcrossing. Our data suggest that artificially fragmented plants with reproductive features similar to those observed here might be reproductively less susceptible to fragmentation.     

Sexually dimorphic legs in a neotropical harvestman (Arachnida,Opiliones): Ornament or weapon?

The evolution of sexually dimorphic traits has been the focus of much theoretical work, but empirical approaches to this topic have not been equally prolific. Males of the neotropical family Gonyleptidae usually present a strong fourth pair of legs armed with spines, but their functional significance is unknown. We investigated the putative functions of the leg armature in the harvestman Neosadocus maximus. Being a non-visual species, the spines on male legs can only be perceived by females through physical contact. Thus, we could expect females to touch the armature on the legs of their mates if they were to evaluate it. However, we found no support for this hypothesis. We did show that (1) leg armature is used as a weapon in contests between males and (2) spines and associated sensilla are sexually dimorphic structures involved in "nipping behavior", during which a winner emerged in most fights. Finally, we demonstrate that five body structures directly involved in male-male fights show positive allometry in males, presenting slopes higher than 1, whereas the same structures show either no or negative allometry in the case of females. In conclusion, leg armature in male harvestmen is clearly used as a device in intrasexual contests.     

Modeling intra-sexual competition in a sex pheromone system: how much can female movement affect female mating success?

Mating disruption theory predicts that high concentrations of female pheromone, and/or large numbers of release sites, should confuse males orienting to "calling" females, reduce the number of successful matings, and decrease the reproductive potential of the population. In this scenario, females are regarded as stationary point sources of pheromone. Past behavioral observations, however, have shown virgin female grape root borers, Vitacea polistiformis Harris, significantly alter their behavior in mating disruption treatments. Treated females call at different heights, move less before call initiation, and move more after call initiation than control females. Pheromone gland dragging and wing fanning also increase significantly during pheromone treatments. These behavioral differences are significant only if they alter the mating success of females. Because long-term field studies are impractical, we used known behavior of male and female GRB to build a Fortran language time step model, adding the effects of female movement to past models of male pheromone plume following. Females were distributed randomly, and then assigned a conditional movement strategy. If females were within the competitive portion of another female's plume, the downwind female moved. Except in the lowest population density tested, females moving upwind and crosswind when in a competing female's pheromone plume mated significantly more often than females remaining stationary. In all population simulations, mating success was significantly reduced when females moved downwind. These field and simulation studies provide strong evidence for female movement as a previously overlooked potential mechanism for resistance to mating disruption treatments, as well as a shaping behavior in the evolution of pheromone communication systems.