Expression Of Alternative Reproductive Tactics Research Articles

Kinship mediates the expression of alternative reproductive tactics in competing spider mite males

Owing to its fitness implications, the degree of genetic relatedness within and between the sexes is a highly significant determinant in mate competition, mate choice and mating success. The role of kinship in the expression of alternative reproductive tactics (ARTs) has been rarely investigated. Here we scrutinized kinship effects on male ARTs in two-spotted spider mites Tetranychus urticae.

Kin-mediated plasticity in alternative reproductive tactics.

Conditional strategies occur when the relative fitness pay-off from expressing a given phenotype is contingent upon environmental circumstances. This conditional strategy model underlies cases of alternative reproductive tactics, in which individuals of one sex employ different means to obtain reproduction. How kin structure affects the expression of alternative reproductive tactics remains unexplored. We address this using the mite Rhizoglyphus echinopus, in which large males develop into aggressive 'fighters' and small males develop into non-aggressive 'scramblers.' Because only fighters kill their rivals, they should incur a greater indirect fitness cost when competing with their relatives, and thus fighter expression could be reduced in the presence of relatives. We raised mites in full-sibling or mixed-sibship groups and found that fighters were more common at higher body weights in full-sibling groups, not less common as we predicted (small individuals were almost exclusively scramblers in both treatments). This result could be explained if relatedness and cue variability are interpreted signals of population density, since fighters are more common at low densities in this species. Alternatively, our results may indicate that males compete more intensely with relatives in this species. We provide the first evidence of kin-mediated plasticity in the expression of alternative reproductive tactics.

Behavior of "Intermediate" Males of the Dimorphic Squid Doryteuthis pleii Supports an Ontogenetic Expression of Alternative Phenotypes.

The expression of alternative reproductive tactics (ARTs) by different-sized males of loliginid squids has been extensively investigated. In loliginids, alternative phenotypes are characterized by discontinuous differences in behavior, body size, sperm deposition site, and morphology and functioning of ejaculates. Large consort males guard females, display agonistic behaviors toward rival consort males, and mate with females in the male-parallel (MP) position. Small sneaker males avoid fighting contests and instead adopt furtive behaviors to access females guarded by consort males, mating with females in the head-to-head (HH) posture. Recently, the reappraisal of preserved material from the loliginid squid Doryteuthis pleii showed that intermediate-sized males (so-called “intermediate” males) had both sneaker- and consort-like ejaculates, leading to the hypothesis of them being a transitional stage between both phenotypes. Here, we describe observations made in captivity showing that intermediate males can display agonistic behaviors toward consort males and mate with females in both mating positions, depending on the male’s current reproductive context, i.e., generally in HH, but switching to MP when the female is laying eggs. Such unusual findings of intermediate males simultaneously displaying behaviors of both sneaker and consort males comprise additional evidence corroborating the ontogenetic hypothesis for phenotypic expression of ARTs in this species. Taken together, our results indicate that (1) instead of competing with large consort males for female access and monopolization, small/young males adopt sneaker tactics to obtain mating opportunities, and (2) as they continue to grow, they gradually modify the morphology of their ejaculates and their mating behavior, going through an “intermediate” stage, before becoming large consort males.

Experimental manipulation of reproductive tactics in Seba's short-tailed bats: consequences on sperm quality and oxidative status.

To reproduce, males have to fertilize the female’s eggs, sometimes in competition with ejaculates of other males. In species where males display alternative reproductive tactics, whereby territorial males secure mating and non-territorial males have to sneak copulations, the latter might be expected to invest relatively more resources towards sperm quality compared with the territorial males. Sperm cells are especially vulnerable to oxidative stress, which reduces male fertility. Therefore, antioxidant resources are expected to modulate sperm quality, and might be allocated differently between reproductive tactics. To test the link between reproductive tactics, redox profile and sperm quality, we experimentally induced changes in the reproductive tactics of 39 captive males Seba’s short-tailed bats Carollia perspicillata. We monitored the blood and ejaculate oxidative balance, and the sperm quality before, 7 days and 21 days after the manipulation of reproductive tactic. Although ejaculates’ oxidative damage was negatively related to sperm velocity, males exhibited similar blood and ejaculates redox profiles and similar sperm quality, regardless of their reproductive tactic. Possibly, these results arise as a consequence of some constraints having been lifted during the experiment. Our results also suggest that, in Seba’s short-tailed bats, the expression of alternative reproductive tactics is not subjected to strong oxidative constraints. Furthermore, our results could reflect an absence of trade-off between pre- and post-copulatory traits in harem males, as they could be selected to invest both in female attraction and sperm quality, as a consequence of their inability to fully monopolize females.

Resource availability, mating opportunity and sexual selection intensity influence the expression of male alternative reproductive tactics.

The expression of alternative reproductive tactics can be plastic and occur simultaneously depending on cues that vary spatially or temporally. For example, variation in resources and sexual selection intensity is expected to influence the pay-off of each tactic and shape the decision of which tactic to employ. Males of the nuptial gift-giving spider Pisaura mirabilis can adopt three tactics: offering a genuine prey gift, a 'worthless' non-nutritious gift or no gift. We hypothesized that resources and/or male body condition, and mating opportunity and sexual selection intensity, vary over the course of the mating season to shape the co-existence of alternative traits. We measured these variables in the field over two seasons, to investigate the predictions that as the mating season progresses, (i) males become more likely to employ a gift-giving tactic, and (ii) the likelihood of switching from worthless to genuine gifts increases. Prey availability increased over the season and co-varied with the propensity of males to employ the gift-giving tactic, but we found no support for condition-dependent gift giving. Males responded to an increase in female availability by increasing their mating effort (gift production). Furthermore, the frequency of genuine gift use increased with sexual selection intensity, consistent with the assumption that sperm competition intensity increases with time. Our results suggest that the frequency of alternative tactics is shaped by seasonal changes in ecological factors and sexual selection. This leads to relaxed selection for the gift-giving tactic early in the season when females are less choosy and resources more scarce, and increased selection for genuine gifts later in the season driven by mating opportunity and risk of sperm competition.

Evidence for ontogenetically and morphologically distinct alternative reproductive tactics in the invasive Round Goby Neogobius melanostomus.

Alternative reproductive tactics are characterized by the occurrence of discrete alternative morphs that differ in behavioural, morphological and physiological traits within the same sex. Although much effort has been made to describe the behaviour, morphology and physiology of such alternative morphs, less effort has been invested investigating how much overlap there is in the characteristics of such morphs in natural populations. We studied random population samples of the invasive Round Goby Neogobius melanostomus from five different localities in the river Rhine system in the Netherlands. We found two morphologically and physiologically distinct male morphs which likely represent alternative reproductive tactics. Almost all mature males under 9.35 cm total length had a gonadosomatic index > 3%, suggestive of a sneaker tactic, while nearly all males above 9.35 cm has a gonadosomatic index of < 3%, suggestive of a parental tactic. Cheek size and eye diameter alone were sufficient to distinguish the two morphs. Gonads had a different relationship with size in the two morphs, indicating separate growth trajectories. The gonad mass of sneaker morphs would be ca. 7.5 times as high as the gonad mass of parental morphs of the same total length after extrapolation. Few (9%) intermediates were found, suggesting that the expression of alternative reproductive tactics is determined before the first breeding season. This contrasts with studies on other goby species, which show evidence of plastic tactics that can be affected by social circumstances. We conclude that it is possible to distinguish two alternative male morphs in the Dutch Round Goby population using morphological measurements alone. Although behavioural observations are needed to provide conclusive evidence, the difference in GSI between these morphs indicates that these morphs reflect alternative reproductive tactics.

Birth date predicts alternative life‐history pathways in a fish with sequential reproductive tactics

Summary In species with plastic expression of alternative reproductive tactics (ARTs), individuals of the same sex, usually males, can adopt different reproductive tactics depending on factors such as body size. The ‘birth date hypothesis’ proposes that condition‐dependent expression of ARTs may ultimately depend on birth date, because individuals born at different times of the year may achieve different sizes and express different reproductive tactics accordingly. However, this has rarely been tested. Here, we tested this hypothesis in a fish with ARTs, the peacock blenny (Salaria pavo). A long‐term (6 years) mark–recapture study demonstrated that ARTs in the peacock blenny were sequential and that males may follow at least two alternative life‐history pathways: a nest‐holder pathway, in which males express the nest‐holder tactic from their first breeding season onwards, and a parasitic pathway, where males reproduce on their first breeding season as sneaker males and subsequently as nest‐holders. We have found evidence of a birth date effect on the expression of ARTs in the peacock blenny. Males following the nest‐holder pathway are born earlier and are larger at the first breeding season than males following the parasitic pathway, but they have similar growth curves. The mechanisms underlying a birth date effect are far from clear and might be diverse. We have not found support for a mechanism of body size threshold triggering sexual maturation and subsequent ARTs. A mechanism of tactic determination that is strictly based on timing of first maturation is also unlikely. A proxy of lifetime reproductive success shows crossing (body size associated) fitness curves for the two main life‐history pathways.

SELECTION FOR ALTERNATIVE MALE REPRODUCTIVE TACTICS ALTERS INTRALOCUS SEXUAL CONFLICT

Intralocus sexual conflict (IASC) arises when fitness optima for a shared trait differ between the sexes; such conflict may help maintain genetic variation within populations. Sex-limited expression of sexually antagonistic traits may help resolve the conflict, but the extent of this resolution remains a subject of debate. In species with alternative male reproductive tactics, unresolved conflict should manifest more in a more sexually dimorphic male phenotype. We tested this prediction in the bulb mite (Rhizoglyphus robini), a species in which aggressive fighters coexist with benign scramblers. To do this, we established replicated lines in which we increased the proportion of each of the alternative male morphs using artificial selection. After approximately 40 generations, the proportion of fighters and scramblers stabilized at >0.9 in fighter- and scrambler-selected lines, respectively. We then measured several female fitness components. As predicted by IASC theory, female fecundity and longevity were lower in lines selected for fighters and higher in lines selected for scramblers. This finding indicates that sexually selected phenotypes are associated with an ontogenetic conflict that is not easily resolved. Furthermore, we suggest that IASC may be an important mechanism contributing to the maintenance of genetic variation in the expression of alternative reproductive tactics.

Experimental increase of testosterone levels in free-ranging juvenile male African striped mice (Rhabdomys pumilio) induces physiological, morphological, and behavioral changes

Testosterone influences sexual differentiation in early development, and activates sexual maturation and sex-related behavior in males during puberty. Testosterone can also influence the expression of male alternative reproductive tactics, by either organizational effects (fixed tactics) or by activational effects (plastic tactics). However, the roles of testosterone in sexual maturation and at the same time the expression of alternative reproductive tactics have been little investigated experimentally, and studies of free-ranging mammals are lacking. We conducted a field experiment in free-ranging juvenile African striped mice (Rhabdomys pumilio), a species with alternative reproductive tactics. Juvenile male striped mice reaching puberty can remain in their family as philopatric group-living males with low testosterone levels, or they can disperse and become solitary living roamers with much higher testosterone levels. We tested whether experimentally increased testosterone levels in non-scrotal juvenile males induces puberty and leads to the expression of the roaming tactic. Testosterone-treated males received the hormone for 15days by silastic implants which were empty in control-treated males. When compared to control-treated males, testosterone-treated males had higher testosterone levels, lower corticosterone levels, and became scrotal with descended testes. Testosterone-treated males also had larger testes, larger epididymides, and showed indication of spermatogenesis. Testosterone-treated males did not become solitary-living roamers, but had larger home ranges than control males. We conclude that testosterone can induce sexual maturation and causes juvenile males to increase their home ranges, maybe to search for dispersal opportunities.

Temporal variation in cuckoldry and paternity in two sunfish species (Lepomis spp.) with alternative reproductive tactics

Male alternative reproductive tactics have been described in many mating systems. In fishes, these tactics typically involve a territorial male that defends a spawning site or nest and a parasitic male that uses sneaking or female mimicry to steal fertilizations from the territorial male. In this paper, we use molecular genetic markers to examine the success of males that adopt alternative reproductive tactics in two sunfishes, comprising the bluegill ( Lepomis macrochirus Rafinesque, 1819) and the pumpkinseed ( Lepomis gibbosus (L., 1758)). In sunfishes, the tactics are referred to as parental (territorial male) and cuckolder (parasitic male). We show that cuckoldry rates peak in the second trimester of the breeding season in bluegill, whereas cuckoldry rates are lowest during this period in pumpkinseed. We also show that paternity of parental male bluegill is positively correlated with body condition, but not body length or mass. No relationship between these phenotypic variables and paternity in pumpkinseed was found. We discuss the patterns of cuckoldry in relation to differences between the species in mating opportunities, parental male defence ability, and cuckolder density. Finally, we discuss how the paternity data can be used to differentiate between two mechanisms underlying the expression of alternative reproductive tactics, comprising the condition strategy and alternative strategies.