Spermatophore characteristics in bushcrickets vary with parasitism and remating interval

Male bushcrickets provide females with a nuptial gift, a spermatophore, which is transferred to females at mating. The spermatophore consists of a gelatinous mass, the spermatophylax, and the sperm-containing ampulla. Male spermatophore size is positively correlated with insemination rate and female refractory period and therefore with male reproductive success. In this study, we examined spermatophylax weight, ampulla weight and sperm number in males of Poecilimon mariannae parasitized by the acoustically orienting fly Therobia leonidei. We show that in parasitized males, spermatophylax weight decreases with the level of parasitism. In line with the hypothesis that parasitism is a cost to reproduction, we found that spermatophylax weight was reduced at remating. In contrast, the replenishment of the spermatophylax in unparasitized males was complete after 2 days and was increased no further after 3 days. Both sperm number and ampulla weight showed an increase over time since last mating and sperm production was estimated at a constant rate of 500,000 per day in all individuals, regardless of parasitism. The allocation of investment in components of the spermatophore varies greatly with parasitism and remating. Both factors had rather independent effects on spermatophore constitution, revealing functional constraints acting on spermatophore characteristics in bushcrickets, which are important for understanding the selection pressures working on its components. Received: 13 September 1999 / Received in revised form: 4 February 2000 / Accepted: 13 March 2000     

The cost of mating with a non-virgin male in a monandrous butterfly: experimental evidence from the speckled wood, Pararge aegeria

Contrary to vertebrates, sperm production in insects may bear considerable costs for males. This is especially true in species that donate spermatophores containing sperm and nutrient-rich accessory gland products like in butterflies. Hence, spermatophores at first and subsequent copulations can differ in a quantitative and qualitative way. Such effects have particularly been shown in polyandrous species providing large spermatophores. Here we experimentally tested the effect of male mating status (virgin male vs recently mated male) on copulation duration, spermatophore size and females’ fitness components in a monandrous butterfly Pararge aegeria that typically donates small spermatophores. Copulations with non-virgin males lasted on average five times longer than that with virgin males and resulted in a spermatophore which was on average three times smaller. Number of eggs laid and female life span were not affected by the mating status treatment, but there was a significant effect on the number of living caterpillars a female produced, as copulations with virgin males resulted in higher numbers of larval offspring. Interestingly, the difference in spermatophore mass at the first and the second copulation increased with male body size. This suggests differential spermatophore allocation decisions among males of different size. Consequences for females and potential mechanisms influencing female fitness components are discussed. Given the small absolute size of spermatophores in P. aegeria, components other than consumable nutrients (perhaps hormones) should cause the observed effects.     

Sperm economy and limitation in spiny lobsters

Sperm limitation, when female fertilisation success is constrained by the supply of sperm, is generally perceived to be an uncommon feature of reproduction in species which directly transfer gametes during copulation. Male size, previous copulations, and the balance of expected reproductive return and future mating opportunity may, however, limit the amount of sperm males transfer to females. We used laboratory experiments where mate size could be manipulated and its consequences on spermatophore size and clutch size determined, to show that in two genera of spiny lobsters (Crustacea: Palinuridae) male reproductive output limits the size of clutches brooded by females. In Panulirus argus from the Florida Keys, we show that while male size affects spermatophore area, males also vary the amount of ejaculate positively with female size. Furthermore, the area of the spermatophore has a greater influence than female size on subsequent clutch weight. In Jasus edwardsii from New Zealand, female size, male size and mate order all affect clutch weight. In both species, clutches fertilised by small males in the laboratory are significantly smaller than clutches fertilised by large males. These results suggest that to ensure they receive sufficient sperm, females should either mate several times prior to oviposition, mate as early as possible in the reproductive season, or choose large, preferably unmated males as partners and thus compete with other females for preferred males. Sperm-limited female fecundity has the potential to limit the egg production of fished populations where large males are typically rare. Received: 18 May 1998 / Received in revised form: 20 November 1998 / Accepted: 30 November 1998     

The ultimate function of nuptial feeding in the bushcricket Poecilimon veluchianus (Orthoptera : Tettigoniidae : Phaneropterinae)

Summary During mating the males of the bushcricket Poecilimon veluchianus transfer a large spermatophore of about a quarter of their body weight to the female. Such nuptial feeding is often thought to function as paternal investment by increasing the fitness of the male's offspring. According to an alternative, though not mutually exclusive, hypothesis, the size of the spermatophore is maintained because of its function as a sperm protection device. In this case the cost to the male should be classified as mating effort. To discriminate between these two hypotheses we measured the duration of sperm transfer into the female spermatheca and the time taken for spermatophore consumption. A comparison of durations revealed that spermatophore consumption interferes with the process of sperm transfer (Fig. 4). There was no significant effect of spermatophore consumption on number of eggs laid, weight of eggs or absolute weight of hatched larvae. The relative dry weight of hatched larvae, however, was increased as a result of spermatophore consumption (Table 1). Thus spermatophylax size is adjusted in accordance with a sperm protection function and the spermatophylax therefore represents mating effort. The increase in relative dry weight indicates that there may also be a paternal investment effect of the spermatophylax, if the offspring that benefit from spermatophylax materials are fathered by the donating male. Correspondence to: K. Reinhold     

Competition,fluctuating asymmetry and sperm transfer in male gryllid crickets (<Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> and <Emphasis Type="Italic">Gryllodes sigillatus</Emphasis>)

Sperm competition theory predicts that ejaculate expenditure by males should increase with the risk of competition from rivals. The tendency to modulate expenditure, however, may reflect various constraints acting on different individuals. We test this idea here by looking at the effects of competition and fluctuating asymmetry (FA) in two species of gryllid cricket. Male Gryllus bimaculatus and Gryllodes sigillatus modulated the sperm content of their spermatophores in relation to both apparent competition and FA in limb size. Males of both species increased the amount of sperm transferred when an apparent male competitor was present, sperm number increasing regardless of the species of apparent competitor in G. bimaculatus, but significantly more with a conspecific competitor in G. sigillatus. In addition, the relative size of the spermatophylax in G. sigillatus increased in the presence of apparent competition, while spermatophore transfer time decreased in G. bimaculatus. Sperm number showed opposite relationships with limb asymmetry in the two species, decreasing with increasing asymmetry in males in G. sigillatus, but increasing with asymmetry in both males and females in G. bimaculatus. G. bimaculatus, but not G. sigillatus, also transferred more sperm when paired with a larger female. Relationships between reproductive measures and limb asymmetry were significant only when competitors were present in G. bimaculatus, but were significant regardless of competition in G. sigillatus. The differences may reflect the additional burden of producing a spermatophylax in G. sigillatus.     

Male asymmetry and postcopulatory sexual selection in the fly Dryomyza anilis

The significance of male asymmetry in postcopulatory sexual selection was studied in the fly Dryomyza anilis by examining whether male asymmetry is related to fertilization success. The traits measured were wing length, tibia length and the length of small and large claspers. The male claspers are situated at the tip of the abdomen, functional pair of claspers consists of a large and a small clasper on the same side of the body. These claspers are used to tap the female abdomen after sperm transfer, which has been shown to increase fertilization success for the mating male. Fertilization success was negatively related to the fluctuating asymmetry of wing length, suggesting either female preference for more symmetrical males or a relationship between male asymmetry and intrasexual selection which was reflected in mating performance. Fertilization success was also related to the length of small claspers, decreasing with increasing length of the claspers. In addition, males with asymmetrical small claspers enjoyed higher fertilization success than symmetrical ones. This study shows that fluctuating asymmetry in wing length is an important fitness trait in postcopulatory sexual selection. Since male tapping affects sperm distribution in the female's sperm storage organs, the higher fertilization success of males with asymmetrical small claspers could have a functional relationship with the asymmetrical position of female sperm storage organs. Received: 6 March 1997 / Accepted after revision: 8 November 1997     

No cost of male mating experience on female reproductive success in the almond moth, <Emphasis Type="BoldItalic">Cadra cautella</Emphasis> (Lepidoptera; Pyralidae)

Male copulation experience may have a profound impact on female reproductive success if male reproductive investment declines over consecutive copulations and if females are unlikely to re-mate. Male reproductive investment is particularly interesting in lepidopterans because males produce dimorphic sperm: a fertilizing (eupyrene) and a non-fertilising (apyrene) sperm. In two experiments, we explored the lifetime reproductive investment of male almond moths, Cadra cautella (also known as Ephestia cautella) and examined its influence on female reproductive success. In the almond moth, females re-mate infrequently and males transfer sperm in a spermatophore. Attached to the spermatophore is a large chitinous process, the function of which is unknown. One group of males were permitted consecutive copulations with virgin females and the amount of sperm and size of the spermatophore transferred were compared for all females. We found that the number of both eupyrene and apyrene sperm per ejaculate decreased with his increased mating frequency, while the size of the spermatophore process decreased dramatically after the male’s first copulation. In a second experiment, we allowed males to mate with females throughout their lives and then compared female fecundity and fertilisation success. We found no obvious decrease in female fecundity and fertilisation success with increased male copulation experience, despite the likely reduction in male gametic investment. We discuss potential explanations for the development of this enlarged and elaborate first spermatophore of male almond moths given that it confers no clear fitness advantage to females.     

Multiple mating in the ant Acromyrmex versicolor: a case of female control

In eusocial insects, polyandrous mating has the potential to reduce genetic relatedness of individuals within a colony, which may have a profound effect on the stability and social structure of the colony. Here we present evidence that multiple mating is common in both males and females of the desert leaf-cutter ant Acromyrmex versicolor. Females seem to have complete control over the number of matings, and mate on average with three males, even though the sperm transferred in a single copulation is sufficient to fill the spermatheca. To determine whether there is a bias in the representation of sperm from different mates in the spermatheca, females were mated to three or four males in controlled mating experiments and were subsequently allowed to found colonies in the laboratory. Paternity analysis of the offspring by random amplified polymorphic DNA analysis showed that all males that have been mated to a female successfully contributed sperm to the production of her offspring. No significant asymmetry in sperm use was detected, suggesting complete sperm mixing. Different hypotheses to explain polyandrous mating are discussed, and it is argued that the best hypothesis to explain polyandrous mating and complete sperm mixing in A. versicolor is that utilizing genetically diverse sperm confers a selective advantage on females. Received: 14 June 1995/Accepted after revision: 26 November 1995     

Different ejaculate delivery strategies in first versus subsequent matings in the swallowtail butterfly Papilio machaon L.

Summary Evidence suggests that males incur costs in producing ejaculates. Therefore some discrimination should be expected with respect to their pattern of allocating ejaculates. In a laboratory study, males of the European swallowtail butterfly Papilio machaon were mated with conspecific virgin females up to six times. The weight of the spermatophore and of the accessory substances delivered by males, the duration of mating, and the number of sperm in ejaculates were measured. The weight of the first spermatophore was related to male weight but that of accessory substances was not. The weight of the spermatophore delivered by males upon remating was always less than that of the first spermatophore. Second spermatophores Produced by males were smallest when delivered on the day after the first mating and increased asymptotically to about half the initial size with time elapsed since the first mating. Copula duration was highest for males remating on the day after the first mating but decreased to the duration of the first mating for males that were remated on the third day after the first mating or later. The weight of accessory substances delivered by males and the number of sperm in ejaculates were also higher in first matings compared to all subsequent matings. Both of two males that were handpaired on three consecutive days after the first mating produced a cumulative ejaculate mass equalling that transferred at the first mating. Thus the fact that males delivered smaller ejaculates upon remating on the third day after the first mating or later requires explanation, especially in view of the fact that copulations were not prolonged. We argue that the male ejaculate delivery strategy in mildly polyandrous species should be to mate for the first time with virgin females, when a maximum size ejaculate should be delivered, and then to maximize the number of matings. The rationale behind the male shift towards smaller ejaculate sizes delivered at subsequent matings is that: (1) the size of the ejaculate that can be produced is dependent on the time elapsed since the last mating; (2) the abundance of virgin females decreases as the flight season proceeds; and (3) there is an inverse relationship between the size and number of ejaculates that a male can produce in his life-time.     

Sperm precedence, mating interval, and a novel mechanism of paternity bias in a beetle (Tenebrio molitor L.)

When females mate with more than one male, the ensuing sperm competition leads to the evolution of male mechanisms that skew paternity. Males of the yellow mealworm beetle (Tenebrio molitor) transfer a spermatophore to females during copulation, but sperm release and storage occur later. We investigated how the interval between two matings with different males affects sperm precedence by varying the interval between the copulations so that the second mating was either: (1) before sperm release from the first spermatophore (<5 min); (2) after sperm release but before spermatophore ejection (15–20 min); (3) after spermatophore ejection but before sperm storage (4 h), or (4) after complete sperm storage (24 h). We collected offspring over a period of 2 weeks and determined paternity by protein electrophoresis. There was second-male sperm precedence in all treatments, but when the interval was <5 min, the second male usually (86% of cases) had complete sperm precedence (i.e., P ₂=1). Investigations into the mechanism of second-male sperm precedence during <5-min mating intervals indicate that sperm release from the first spermatophore is inhibited, a phenomenon which has not been previously documented. Received: 31 January 2000 / Revised: 9 June 2000 / Accepted: 26 August 2000     

Proximal traits and mechanisms for biasing paternity in the red flour beetle Tribolium castaneum (Coleoptera: Tenebrionidae)

A comprehensive understanding of sexual selection requires knowledge of the traits and mechanisms responsible for increasing a male’s paternity share (proportion of progeny sired) relative to that of other males mating with the same female. In this study we manipulated by starvation the expression of traits that might influence male paternity share in Tribolium castaneum. We then conducted experiments to examine how male starvation affects male performance during sequential episodes of sexual selection from mating to progeny production, and investigated female control over specific stages by using live vs dead females. Comparison of starved vs fed males revealed that T. castaneum females have control over spermatophore transfer during mating, as live females rejected inseminations by starved (“low quality”) males. None of the measured male copulatory behaviors (leg-rubbing frequency, asymmetry, and percent of time spent rubbing) affected the probability of successful insemination, but the last two were positively associated with male paternity share. Spermatophore positioning within the female reproductive tract was not affected by male treatment (starved/fed), by female treatment (live/dead), or by male copulatory behaviors. Starvation, however, had a dramatic effect on male reproductive physiology, decreasing both accessory gland size and total number of sperms transferred (but not sperm viability in seminal vesicles). In addition, females who mated to starved males stored fewer sperms in their spermathecae, which, together with decreased ejaculate size, may explain the reduced paternity share of starved males compared to fed males. This study elucidates some cryptic mechanisms influencing male reproductive success and aids our understanding of trait evolution through sexual selection.     

Female resistance and male preference in a stream-dwelling isopod: effects of female molt characteristics

In the stream-dwelling isopod, Lirceus fontinalis, mating contests between males and females occur prior to pair formation. We examined the relative contribution of male preference and female resistance to contest outcomes. We first quantified male and female behavior during typical mating interactions and examined the relationship between time until molt (TTM) and mating outcomes. We then examined the role of male preference and female resistance in determining mating outcomes when females differed in molt type (growth, egg deposition) and appeared to differ in TTM (due to hormone applications). Both male preference and female resistance contributed to different components of the mating sequence but female resistance ultimately determined whether or not pair formation occurred. Males expressed a preference for females that appeared to be close to molt, using variation in levels of molt hormone as a cue. However, males did not discriminate between females based on molt type. Received: 5 March 1999 / Received in revised form: 10 August 1999 / Accepted: 16 October 1999     

Male control of sperm transfer dynamics in a spermatophore-donating bushcricket

In many animal species, male and female interests often differ when it comes to decisions over mating and fertilization. However, it is intrinsically difficult to determine the degree to which males and females exert control over the various processes that determine the outcome of reproductive interactions, and thus to predict how such conflicts will be resolved. For example, in species where sperm are transferred to females via a spermatophore or other external sperm packaging device, it is unclear which sex determines subsequent sperm transfer dynamics to the female reproductive tract. To address this question, we used a reciprocal cross experimental design in a bushcricket species (Poecilimon veluchianus) comprising two subspecies differing in the dynamics of sperm transfer. The reciprocal crosses show that in these bushcrickets the timing of sperm transfer in inter-subspecies crosses closely resemble those typical of the subspecies of the male partner, indicating that it is the properties of the spermatophore rather than its handling by the female that determine sperm transfer dynamics. There was neither a significant female influence nor any indication of an interaction between males and females with regard to the number of sperm transferred after a set interval. Our study suggests that males rather than females appear to control the timing of the insemination process in this species.     

Asymmetry in a visual signaling character and sexual selection in a wolf spider

Fluctuating asymmetry (FA) can indicate developmental instability in bilaterally symmetric organisms, and studies have shown that the degree of asymmetry in male secondary sexual characters influences female mate choice in a number of taxa. In male Schizocosa ocreata wolf spiders, conspicuous tufts of bristles on the forelegs are a critical component of visual courtship signals, which appear to play a role in female mate choice. Previous studies have shown that females exhibit reduced receptivity to males with regenerative asymmetry, a consequence of leg loss and regeneration that causes males to be grossly asymmetric with respect to this important signaling character. We provide data on the occurrence of FA in the tufts of S. ocreata, and examine further the influence of asymmetry on female mate choice. The distribution of tuft area asymmetry values from a sample of field-collected males was normal, with a mean value of zero, indicating true FA. For a subset of males measured directly after field collection and prior to feeding, tuft asymmetry was significantly negatively correlated with measures of body size (body length) and condition (abdomen volume/cephalothorax width). Receptivity responses of females to visual signals from live males of similar size varied with the degree of asymmetry in male tufts. Since FA covaries with male body size and condition, which may also influence behavioral vigor, we used video image manipulation to alter the degree of asymmetry in tufts of a courting male while holding size and condition constant. Asymmetry treatments represented values within the range of natural FA variation as well as more extreme values characteristic of regenerative asymmetry. With the confounding effects of male size, condition, and behavior held constant, female spiders exhibited reduced receptivity responses to all experimental asymmetric video images relative to a control video stimulus. There were no differences in the frequency of female receptivity among the various asymmetry treatments, suggesting that discrimination against asymmetry in conspecific male signal characters is not simply a rejection of extreme phenotypes. Results suggest that asymmetry in a key male secondary character used in visual signaling, independent of any concomitant behavioral or size factor, is an important criterion in mate choice. Received: 26 February 1998 / Accepted after revision: 12 September 1998     

Testes mass and reproductive mode of minnows

Most comparative analyses of relative testes mass find that testes are larger in species in which more sperm competition is predicted (multiple males mate with individual females). I tested for differences in adjusted testes mass (for body mass) by spawning mode and by sexual size dimorphism in a comparative analysis of 37 minnow species. No significant differences were found for testes mass by spawning mode or sexual size dimorphism. These results imply a lack of response to selection on testes size from sperm competition in minnow species. Possible explanations for the lack of the expected relationship between testes mass and mating systems in minnows are presented. Received: 8 November 1999 / Received in revised form: 27 January 2000 / Accepted: 13 February 2000     

Male reproductive investment relative to age and flight behaviour in the monandrous butterfly Pararge aegeria

Male reproductive investment may signify a considerable cost to male insects that produce sperm packages or spermatophores. Male butterflies allocate much of their active time budget to mate location, and they may adopt different behavioural strategies to do so. In the speckled wood butterfly (Pararge aegeria L.), males adopt either a territorial wait-and-fight strategy (territorial perching) or a fly-and-search strategy in wider areas (patrolling). In this study, we analysed the impact of male age, male size and male behaviour (i.e. behavioural strategies and levels of activity) on spermatophore investment (i.e. spermatophore mass, number of eupyrene sperm bundles). As predicted, reproductive investment increased with male age and size. Nevertheless, the increase of spermatophore mass with age and the number of eupyrene sperm bundles (i.e. fertile sperm) was stronger in low-activity males compared to active flying males. This suggests that flight activity has a negative impact on male reproductive investment. However, males that were forced to fly in the laboratory produced more eupyrene sperm bundles than resting males. We discuss the potential effects of male–male competition and predation risk on current versus future male reproduction. Males adopting different mate-locating strategies (perching and patrolling) in outdoor cages did not differ in spermatophore traits as was predicted from their very different flight performances. Copulations of territorial perching males took somewhat longer than copulations with non-perching males. There was a significant family effect of spermatophore size and of the expression of male mate-locating strategies suggesting heritable variation. Female traits (i.e. age and size) did not strongly affect spermatophore production. We discuss the results relative to both ultimate and proximate explanations of the complex relationships between butterfly activity, behavioural strategies, age and spermatophore production.     

Mating effort or paternal investment? Incorporation rate and cost of male donations in the wartbiter

Summary Provision of nuptial gifts and incorporation of male-derived substances occurs in several insects. There has been some discussion as to whether these gifts should be regarded as a form of paternal investment or part of mating effort. One assumption of the paternal investment hypothesis is that the donating male is expected to father all, or the majority of, the offspring. The rate of incorporation into developing eggs is therefore assumed to be rapid enough to assure the donating male of paternity of the offspring in which he is investing. Spermatophore production in a paternal investment system is also expected to be costly, since males are providing females with highly nutritious gifts during copulation, thereby increasing female fecundity. Therefore, under conditions of food limitation males are expected to alter either mating frequency or spermatophore size. Experiments on the wartbiter (Decticus verrucivorus), using radioactive isotopes, show that the incorporation rate of male-derived substances is so slow that females become sexually receptive before any material received from the last male have been incorporated into eggs laid by the female. Since sperm mixing occurs in this species and since the label is present in eggs laid up to more than 30 days after mating, males might potentially benefit another male's offspring. The nuptial gift (the spermatophylax) was found, regardless of mating frequency and diet, to have a low protein content. These findings coincide with previous results showing that consumption of the gift had no effect on female fecundity. Instead, wartbiter males keep mating frequency high, are capable of mating every day regardless of food availability, invest the same proportion of the body weight in spermatophore production regardless of size, and produce a spermatophylax that correlates with ampulla size. These results indicate that the nuptial gift mainly functions to increase fertilization success in the wartbiter.     

Effects of mating history on ejaculate size, fecundity, longevity, and copulation duration in the ant-tended lycaenid butterfly, Jalmenus evagoras

The mating system of the Australian lycaenid butterfly, Jalmenus evagoras, is highly unusual compared to most other Lepidoptera. Characteristics of this system, which has been termed an ’explosive mating strategy,’ include the formation of an intensely competitive mating aggregation of males, a highly male biased operational sex ratio, a lack of discrimination and mate choice by both sexes, a high variance in male mating success, and female monogamy. In this study, we tested the hypothesis that multiple mating by males imposes physiological costs resulting in smaller spermatophores, and that this results in a fitness cost to females. We found that male J. evagoras transferred only 2.2% of their eclosion weight during their first mating, consistent with the hypothesis that males of monandrous species produce a relatively small investment. The wet weight of the ejaculate declined by an average of 27% at the second mating and the dry weight by 29%, and an intermating interval of 5–9 days was needed for the ejaculate to return to the size at the first mating, regardless of male size or age. Wet ejaculate mass increased proportionally with male size, though dry mass was proportionally larger in smaller males. Ejaculate mass tended to increase with male age at both first and second matings. Female characteristics, in general, did not affect ejaculate mass, although the wet weight of the ejaculate was positively associated with female weight at the second mating. Copulation duration increased from 2.4 h to approximately 3 h at the second mating, and to over 4 h at the third and fourth matings. Fecundity was positively correlated with female size but not with mating history, copulation duration, or any other characteristics measured for either males or females. Female longevity declined significantly as the number of times the male partner had previously mated increased. We conclude that despite the small male investment in ejaculate, the costs of multiple mating may nonetheless be significant, as indicated by the reduction in ejaculate mass, an increase in copulation duration, and reduction in female lifespan with increasing mating number. Received: 22 January 1999 / Received in revised form: 28 July 1999 / Accepted: 18 September 1999     

Mating system and alternative male mating tactics in the grass goby Zosterisessor ophiocephalus (Teleostei: Gobiidae)

 Peculiar fertilization dynamics, with males releasing sperm in mucous trails lasting several hours, characterize some demersal spawning fish. The mating system was investigated in a natural population of one of these species: the grass goby Zosterisessor ophiocephalus (Pallas, 1814), a large coastal goby inhabiting seagrass meadows in shallow brackish water. Adult males ranged in size from 7.4 to 23 cm total length, but only larger ones were observed to dig and defend a burrow, where they performed parental care on eggs laid by one to several females. Field observations together with analyses of age, sperm production, trail sperm content and sperm competition tests indicated the occurrence of alternative male mating tactics, likely the expression of an ontogenetic gradient. Larger males are older than smaller ones, and while the former are territorial, the latter “sneak” territorial male spawns. The ejaculate characteristics indicate that grass goby males have functionally polymorphic spawns: in fact sperm trails of larger males last longer and release fewer sperm than those of smaller males. Sperm production over several days is more constant in larger than in smaller males, but the total number of sperm released is higher in the latter. The influence of seminal fluid in the functional intraspecific variability in sperm release in this species is discussed. Received: 30 December 1999 / Accepted: 31 July 2000     

Paternal investment in Poecilimon veluchianus bushcrickets: beneficial effects of nuptial feeding on offspring viability

In most bushcrickets, males transfer a large spermatophore during copulation that is afterwards consumed by the female. In some species this nuptial gift enhances offspring fitness and is therefore believed to function as paternal investment. To determine whether this is the case, I examined whether a male's own offspring benefit from spermatophore consumption in the bushcricket Poecilimon veluchianus. Females that consumed a spermatophore produced offspring with increased residuals of dry weight compared to females that were prevented from feeding on the spermatophore. This beneficial effect of spermatophore consumption occurred within the first 4 days after copulation. An increased dry weight indicates higher energy reserves because offspring dry weight correlates significantly with the lifespan of starved larvae and because spermatophore consumption increased the lifespan of starved offspring. During egg-laying, females apply a liquid substance to the soil that hardens and probably serves as protection for the egg clutch. The amount of this substance correlated with the number of eggs laid but did not differ between spermatophore treatments. In P. veluchianus, females mate frequently and there is last-male sperm precedence. The spermatophore thus only constitutes paternal investment when offspring produced before female remating benefit from spermatophore consumption. The dry weight of offspring increased during the first 4 days after spermatophore consumption and thus within the natural remating interval. This shows that the spermatophore has a paternal investment function in addition to its already known sperm protection function. Received: 15 May 1998 / Accepted after revision: 11 October 1998