Control theory predictions of reproductive allocation in female dusky salamanders

A model of the reproductive ecology of female dusky salamanders was used to investigate the allocation scheme that a female might use to maximize her reproductive success. Analysis of the model with techniques of optimal control theory suggests that fecundity is maximized either by allocating food resources to reproduction for the entire time period prior to egg laying, or by growing first, then switching to reproduction later in the year. The analysis also indicates that egg mortality will be minimized if the female provides maternal care at the maximum level throughout egg brooding. These results are not specific to dusky salamanders, but can be extended to other organisms with similar reproductive characteristics.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.     

The impact of sulfur on the reproductive success of <Emphasis Type="Italic">Anagrus</Emphasis> spp. parasitoids in the field

Pesticides targeted at pest species have often been demonstrated to have strong adverse effects on the survival of biological control agents in short-term laboratory bioassays; however, studies examining the influence of pesticides on the actual reproductive success of biological control agents in the field are rare. Because natural enemy reproduction is often directly tied to biological control success, effects of pesticides on reproduction are of central importance. Here we use a new technique to examine the influence of sulfur, a fungicide widely used in grape production, on the reproductive success of Anagrus erythroneurae (Hymenoptera: Mymaridae) and Anagrus daanei (Hymenoptera: Mymaridae), egg parasitoids of the grape leafhopper, Erythroneura elegantula (Homoptera: Cicadellidae). Sulfur has previously been shown to be highly toxic to Anagrus spp. in short-term laboratory and field bioassays, creating the expectation that sulfur should also reduce Anagrus reproductive success in the field. Surprisingly, in two studies, the first comparing the oviposition success of Anagrus collected live in paired sulfur-treated versus untreated vineyards and the second comparing the lifetime reproductive success of Anagrus collected at the end of their lives in unpaired sulfur-treated versus untreated vineyards, we found no effect of sulfur on parasitoid reproductive success. In this system, traditional short-term assays of laboratory toxicity do not appear to predict effects on parasitoid reproductive success, suggesting that demographic approaches to assessing the disruptive effects of pesticides may have an important role in designing IPM programs.     

Reproductive strategies of rhesus macaques

Reproductive strategies incorporate a multitude of mechanisms that have evolved to promote the reproductive success of individuals. Evolutionary perspectives tend to emphasize the advantages of male-male competition and female choice as mediators of differential reproduction. Male rhesus macaques have not been observed to fight for access to sexually receptive females, although they suffer more wounds during the mating season. An increased likelihood of attacks appears to coincide with male troop entry. Males who spend more time in consort and mate with more females tend to sire more offspring. Genetic analysis of paternity has pinpointed age and endurance rivalry, rather than agonistic competition, as key variables associated with variation in progeny production. Female rhesus macaques often copulate with multiple males during their ovulatory period, and tend to conceive on the first cycle of the mating season. Female reproductive success is more likely to be a function of offspring survivorship than the identity of particular male partners. The role of female choice as a direct mediator of male reproductive success is unresolved, but female mate selection seems to indirectly affect male reproductive success because female preference for mating with novel males seems to foster male dispersal. Evaluating whether mating preferences for particular male phenotypes affectsfemale reproductive success is a task for the future. A common denominator to the reproductive strategies of both female and male rhesus macaques is that feeding patterns affect body condition which influences reproductive output and regulates relative reproductive success.     

Group Structure,Nest Size and Reproductive Success in the Cooperatively Breeding Cichlid Julidochromis ornatus: A Correlation Study

The effect of group size on reproductive success has long been studied in cooperatively breeding species, as it might provide an adaptive explanation for group‐living in social species. Numerous studies have shown positive effects of subordinates on reproductive success (‘helper effect’), but these studies have also revealed the importance of controlling statistically, or experimentally, for the effect of other factors that might affect reproductive success. Here, we first examine the relationships between group size, body size of group members and nest size in the cooperatively breeding cichlid Julidochromis ornatus, in which unrelated helpers frequently participate in reproduction and their breeding nests inside rock crevices may be crucial for reproduction and survival of all group members. Then, we subsequently investigate the relationship between group size and reproductive success, while controlling for these factors. The results showed that group size was significantly related to body size of group members rather than nest size; and larger breeders had larger helpers. It was found that group size significantly increased group reproductive output. More importantly, reproductive success of male breeders did not depend on the presence of mature helpers, whereas female reproductive success increased when two males assisted her and tended to decrease when two females bred cooperatively. We conclude that breeding groups of J. ornatus have size hierarchical societies that relate to group size, and group composition of genetically unrelated and co‐breeding members affects their reproductive success.     

Tactic changes in dusky frillgoby Bathygobius fuscus sneaker males: effects of body size and nest availability

Field and laboratory studies were conducted to examine the effects of nest availability and body size on changes in male mating tactics from sneaking to nest‐holding in the dusky frillgoby Bathygobius fuscus. In the field, the body size of nest‐holding males decreased from early to mid‐breeding season, suggesting the possibility of a change in the tactics of sneaker males to nest‐holding. Many sneaker males did not use vacant spawning nests even when size‐matched nests were available, but they continued to reproduce as sneakers. Similarly, in aquarium experiments with available vacant nests, some sneaker males became nest‐holders irrespective of their body size, but some did not. These results showed that nest availability is not a limiting factor for changes in tactics by sneaker males in this species. Because tactic‐unchanged sneaker males were co‐housed with larger nest‐holding males in the tanks, the body size of nearby nest‐holding males may have affected the decision to change tactics for sneaker males. Moreover, smaller individuals among tactic‐changed males tended to spend more time until spawning, probably because they had relatively larger costs and smaller benefits of reproduction as nest‐holding males compared to larger males.     

Maternal investment and size-specific reproductive output in carcharhinid sharks

1.  Life-history theory predicts that organisms will provide an optimal level of parental investment for offspring survival balanced against the effects on their own survival and future reproductive potential.
2.  Optimal resource allocation models also predict an increase in reproductive output with age as expected future reproductive effort decreases. To date, maternal investment in sharks has received limited attention.
3.  We found that neonatal dusky sharks ( Carcharhinus obscurus ) are not independent from maternal resource allocation at the point of parturition but instead are provisioned with energy reserves in the form of an enlarged liver that constitutes approximately 20% of total body mass.
4.  Analysis of long-term archived data sets showed that a large proportion of this enlarged liver is utilized during the first weeks or months of life suggesting that the reported weight loss of newborn sharks signifies a natural orientation process and is not necessarily related to prey abundance and/or indicative of high mortality rates.
5.  Interrogation of near-term pup mass in two carcharhinids, the dusky and spinner shark ( Carcharhinus brevipinna ), further revealed an increase in reproductive output with maternal size, with evidence for a moderate decline in the largest mothers.
6.  For the dusky shark, there was a trade-off between increasing litter size and near-term pup mass in support of optimal offspring size theory.
7.  For both the dusky and spinner shark, there was a linear increase in near-term pup mass with month, which may indicate variable parturition strategies and/or that carcharhinids are able to adjust the length of the gestation period.
8.  The identification of optimal size-specific reproductive output has direct implications for improving the reproductive potential of exploited shark populations and for structuring future management strategies.     

Body mass and sex‐biased parasitism in wood mice Apodemus sylvaticus

Male sex‐biased parasitism (SBP) occurs across a range of mammalian taxa and two contrasting sets of hypotheses have been suggested for its establishment. The first invokes body size per se and suggests that larger individuals are either a larger target for parasites, trade off growth at the expense of immunity or cope better with parasitism than smaller individuals. The second suggests a sex‐specific handicap whereby males have reduced immunocompetence compared to females due to the immunodepressive effects of testosterone. The current study investigated whether sex‐biased parasitism is driven by host ‘body size’ or ‘sex’ using a rodent–tick (Apodemus sylvaticus–Ixodes ricinus) system. Moreover, the presence or absence of large mammals at study sites were used to control the presence of immature ticks infesting wood mice, allowing the impacts of parasitism on host body mass and female reproduction to be assessed. As expected, male mice had greater tick loads than females and analyses suggested this sex‐bias was driven by body mass as opposed to sex. It is therefore likely that larger individuals are a larger target for parasites, trade off growth at the expense of immunity or adapt behavioural responses to parasitism based on their body size. Parasite load had no effect on host body mass or female reproductive output suggesting individuals may alter behaviour or life history strategies to compensate for costs incurred through parasitism. Overall, this study lends support to the ‘body size’ hypothesis for the formation of sex‐biased parasitism.     

Effects of body size and timing of reproduction on reproductive success in female natterjack toads (Bufo calamita)

This study estimated variability in components of reproductive success in female natterjack toads ( Bufo calamita , Laurenti, 1768) as a function of their body size and timing of reproduction. Two components of reproductive success were analysed: fecundity and metamorphic success. Fecundity, as a function of body size, tended to increase exponentially. Age had a negligible effect once body size was statistically controlled. Egg size was also related to female body size but showed greater variation. Metamorphic success depended on the timing of reproduction. Later breeders suffered an exponential decline in metamorphic success because females spawned at recently-filled ponds of ephemeral duration. Since female body size decreased as the breeding season progressed, it may be inferred that larger, early-breeding females would have a differential reproductive success. The inverse relationship between the timing of reproduction and body size may be based on the energetic physiology of vitellogenesis, and on the potential trade-offs between the allocation of energy to growth—higher in smaller females—and reproduction.     

Capital energy drives production of multiple clutches whereas income energy fuels growth in female collared lizards Crotaphytus collaris

Animals invest energy in reproduction that is obtained at two distinct times relative to the reproductive cycle. Energy obtained during egg production is referred to as income energy whereas stored energy acquired prior to reproduction is capital energy. Similar to most ectotherms, squamate reptiles are generally hypothesized to be capital breeders. Nearly all squamates in which income/capital energy investment has been examined thus far produce only one clutch per reproductive season. Although it is likely that squamates producing multiple seasonal clutches fuel first clutches with capital energy, either capital or income energy may be used to produce later clutches. We first monitored female eastern collared lizards over 14 reproductive seasons to confirm that the number of clutches females produce seasonally is a plastic response to variable environmental parameters, and to examine the effects of female body condition at the beginning of the reproductive season on clutch production. Clutch production varied annually and both the size and number of clutches were positively correlated with body condition. We then tested the competing predictions of the income and capital hypotheses experimentally by supplementing the diets of female collared lizards in situ for one season. Diet‐supplementation had no effect on the number of clutches produced but increased growth rates of gravid females. We further tested the competing predictions of these two hypotheses by examining variation in maternal energy investment per clutch using preserved specimens collected near our primary field site. Clutch size was highly correlated with female body size. Together, our results suggest that variation in reproductive output by female collared lizards is linked to stored capital energy rather than income energy, similar to most ectotherms.     

Ejaculate economics: an experimental test in a moth

Increasing evidence shows that spermatogenesis is costly. As a consequence, males should optimize the use of their sperm to maximize their reproductive outputs in their lifetime. However, experimental evidence on this prediction is largely lacking. Here, we examine how a male moth Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) responds to the presence of rivals or additional mates and how such response influences his lifetime reproductive fitness. We show that when rival males are present around a copulating pair, the male ejaculates more sperm to win a sperm competition battle but in such an environment he inseminates fewer females, sires fewer offspring and lives shorter. The opposite is the case when additional females are present during copulation. These findings reveal that elevated reproductive expenditure owing to sperm competition intensity is made at the expense of longevity and future reproduction.     

Folliculogenesis in the domestic cat (Felis catus)

The dynamic regulation of mammalian folliculogenesis is a key component of the reproductive process. Traditionally, the rodent had been used as a model to study ovarian function and reproductive physiology due to the availability of animals, their relatively short cycle length, high rate of fecundity and short generation interval. We maintain that much basic information can be determined using domestic cat ovaries retrieved from local veterinary clinics following routine spaying, without having the expense of maintaining a colony of laboratory cats. Studies of normal feline reproductive physiology and advances in reproductive technology may be extrapolated for use in endangered non-domestic felids. Increased understanding of feline reproduction will be beneficial to veterinary medicine, and to groups working to control feral cat populations. It is important to examine reproductive mechanisms in alternative animal models as there are a vast number of threatened and endangered species in which we lack the critical reproductive information needed to assist in preserving their long-term survival.     

Cost of reproduction in Callosobruchus maculatus: effects of mating on male longevity and the effect of male mating status on female longevity

One of the most studied life-history trade-offs is that resulting from the cost of reproduction: a trade-off arises when reproduction diverts limited resources from other life-history traits. We examine the cost of reproduction in male, and the effect of male mating status on female Callosobruchus maculatus seed beetles. Cost of reproduction for male C. maculatus was manifested as reduced longevity. There was also a positive relationship between male body size and male longevity. Females mated to males that had already copulated twice did not live as long as females mated to males that had copulated once or not at all. The third copulation of males also lasted longer than the two previous ones. We conclude that even though the cost of reproduction for males has been studied much less than that in females, there is growing evidence that male reproductive effort is more complex than has traditionally been thought.     

Effects of Timber Management on Pond-Breeding Salamanders

Abstract: Pond-breeding salamanders spend most of their lives in forested habitat surrounding the vernal pools where they breed. Timber harvesting has been demonstrated to have negative impacts on salamander populations due to changes in soil temperature, soil compaction, and general degradation of habitat. However, little is known about how long it takes for harvested forest habitat to once again become suitable for salamanders. Questions also remain as to whether salamanders will use an area that has been harvested in recent years if an older intact forest area is available. We used drift fences and pitfall traps to capture adult spotted salamanders (Ambystoma maculatum) and marbled salamanders (A. opacum) migrating to 3 vernal ponds during their breeding seasons. The study area contained tracts of forest that were clear-cut 11–12 years prior to the study. All 3 ponds were surrounded by areas of clear-cut and intact forest and drift fences were placed in both habitat types. Similar numbers of spotted salamanders entered the ponds from clear-cut and intact forest areas. The number of marbled salamanders migrating to the ponds did not differ between areas of clear-cut and intact forest. These results suggest that clear-cut habitats may become suitable for adult pond-breeding salamanders after a relatively short regeneration period.     

Reproductive suppression in female cooperatively breeding cichlids

Suppression by dominants of female subordinate reproduction has been found in many vertebrate social groups, but has rarely been shown experimentally. Here experimental evidence is provided for reproductive suppression in the group-living Lake Tanganyika cichlid Neolamprologus pulcher. Within groups of three unrelated females, suppression was due to medium- and small-sized females laying less frequently compared with large females, and compared with medium females in control pairs. Clutch size and average egg mass of all females depended on body size, but not on rank. In a second step, a large female was removed from the group and a very small female was added to keep the group size constant. The medium females immediately seized the dominant breeding position in the group and started to reproduce as frequently as control pairs, whereas clutch size and egg mass did not change. These results show that female subordinate cichlids are reproductively capable, but apparently suppressed with respect to egg laying. Nevertheless, some reproduction is tolerated, possibly to ensure continued alloparental care by subordinate females.     

Empirical evidence of a trade-off between reproductive effort and expectation of future reproduction in female three-spined sticklebacks

Optimal life-history models generally predict that the reproductive effort of iteroparous organisms may increase with age, as their expectation of future reproduction decreases. The population of three-spined sticklebacks (Gasterosteus aculeatus) in the Camargue (Rhone River Delta, France) is annual, all adults dying after their first breeding season. As the three-spined stickleback is a multiple spawner, we tested the hypothesis that reproductive effort may increase during the breeding season on field data. From 1987 to 1998, 653 female sticklebacks were collected in the field during the breeding seasons. The body size, body weight and weights of the liver, gonads and carcass were measured for these individuals. Only gravid females with mature eggs (176 fish) were included in the analysis. Considering the female three-spined stickleback as a capital breeder, the energetic resources available for allocation between soma and gonads were estimated by its body weight. Somatic condition decreased during the breeding season and reproductive effort (gonad weight relative to body weight) increased. These patterns did not vary significantly between years. These observed variations in reproductive effort during the breeding season can be interpreted as empirical evidence of a trade-off between reproductive effort and expectation of future reproduction.     

Sperm competition and sex change: a comparative analysis across fishes

Current theory to explain the adaptive significance of sex change over gonochorism predicts that female-first sex change could be adaptive when relative reproductive success increases at a faster rate with body size for males than for females. A faster rate of reproductive gain with body size can occur if larger males are more effective in controlling females and excluding competitors from fertilizations. The most simple consequence of this theoretical scenario, based on sexual allocation theory, is that natural breeding sex ratios are expected to be female biased in female-first sex changers, because average male fecundity will exceed that of females. A second prediction is that the intensity of sperm competition is expected to be lower in female-first sex-changing species because larger males should be able to more completely monopolize females and therefore reduce male-male competition during spawning. Relative testis size has been shown to be an indicator of the level of sperm competition, so we use this metric to examine evolutionary responses to selection from postcopulatory male-male competition. We used data from 116 comparable female-first sex-changing and nonhermaphroditic (gonochoristic) fish species to test these two predictions. In addition to cross-species analyses we also controlled for potential phylogenetic nonindependence by analyzing independent contrasts. As expected, breeding sex ratios were significantly more female biased in female-first sex-changing than nonhermaphroditic taxa. In addition, males in female-first sex changers had significantly smaller relative testis sizes that were one-fifth the size of those of nonhermaphroditic species, revealing a new evolutionary correlate of female-first sex change. These results, which are based on data from a wide range of taxa and across the same body-size range for either mode of reproduction, provide direct empirical support for current evolutionary theories regarding the benefits of female-first sex change.     

Size and sex allocation in monoecious woody plants

The female size advantage hypothesis predicts that the allocation ratio of female: male reproductive effort should increase with plant size (total reproductive effort). A male height advantage hypothesis has also been proposed, based on the supposed greater advantage of height to male reproductive success in wind-pollinated plants. These ideas were tested with data for wind-pollinated, monoecious trees and shrubs which exhibit a suitably large range of sizes. Number of male inflorescences increased faster with size than did number of female inflorescences in 2 of 9 species; in the remaining 7 species there was no significant difference. The male:female ratio of inflorescence numbers increased with height in 4 of 7 species and did not change significantly in the remaining 3 species, as shown by regression. Height and size are highly correlated and so their effects could not be distinguished. The fact that many conifers place the female cones uppermost in the crown suggests that size and not height favors increased allocation to male function, as does well-established theory connecting the existence of male versus female size advantage to pollen and seed dispersal chacteristics. Regression analysis of the relation between male and female reproductive effort should be done by reduced major axis regression; ordinary least squares regression underestimates slopes; in this study opposite conclusions could be drawn from ordinary least squares and reduced major axis regressions.     

Reproductive strategies in snakes

Snakes of both sexes display remarkable flexibility and diversity in their reproductive tactics. Many features of reproduction in female snakes (such as reproductive mode and frequency, seasonality and multiple mating) allow flexible maternal control. For example, females can manipulate not only the genotypes of their offspring (through mate choice or enhanced sperm competition) but also the phenotypes of their offspring (through allocation 'decisions', behavioural and physiological thermoregulation, and nest-site selection). Reliance on stored energy ('capital') to fuel breeding results in low frequencies of female reproduction and, in extreme cases, semelparity. A sophisticated vomeronasal system not only allows male snakes to locate reproductive females by following scent trails, but also facilitates pheromonally mediated mate choice by males. Male-male rivalry takes diverse forms, including female mimicry and mate guarding; combat bouts impose strong selection for large body size in males of some species. Intraspecific (geographical) variation and phenotypic plasticity in a wide array of reproductive traits (offspring size and number; reproductive frequency; incidence of multiple mating; male tactics such as mate guarding and combat; mate choice criteria) provide exceptional opportunities for future studies.     

Mating does not influence reproductive investment, in a viviparous lizard

Mating is crucial for females that reproduce exclusively sexually and should influence their investment into reproduction. Although reproductive adjustments in response to mate quality have been tested in a wide range of species, the effect of exposure to males and mating per se has seldom been studied. Compensatory mechanisms against the absence of mating may evolve more frequently in viviparous females, which pay higher direct costs of reproduction, due to gestation, than oviparous females. To test the existence of such mechanisms in a viviparous species, we experimentally manipulated the mating opportunity of viviparous female lizard, Lacerta (Zootoca) vivipara. We assessed the effect of mating on ovulation, postpartum body condition and parturition date, as well as on changes in locomotor performances and body temperatures during the breeding cycle. Female lizards ovulated spontaneously and mating had no influence on litter size, locomotor impairment or on selected body temperature. However, offspring production induced a more pronounced locomotor impairment and physical burden than the production of undeveloped eggs. Postpartum body condition and parturition dates were not different among females. This result suggests that gestation length is not determined by an embryonic signal. In the common lizard, viviparity is not associated with facultative ovulation and a control of litter size after ovulation, in response to the absence of mating.