A longitudinal study of age-specific reproductive output and body condition among male rhesus macaques,<Emphasis Type="Italic"> Macaca mulatta</Emphasis>

In many mammalian species, male reproductive success appears to climb sharply at young adulthood, form a brief plateau during prime ages, and decline among older animals, a pattern often attributed to reduced physical condition with ageing. However, solid evidence to either substantiate or refute this profile among nonhuman primates is lacking. Here, we combine a decade of genetic analysis of paternity among free-ranging rhesus macaques, Macaca mulatta, with information about body condition in order to evaluate how changes in morphology might govern age-specific reproduction among males. We show that age-specific reproductive success traverses the same life history profile as found in other mammals, but reductions in reproductive output with advanced age were associated with reduced chances of survivorship rather than accompanied by diminished body condition. We demonstrate that variance in male age at onset of reproduction is three times greater than variance in female age at onset of reproduction. We provide the first evidence from primates that age-specific reproductive output among males is not a consequence of age-related changes in body condition, but reflects social and demographic factors.     

Body size phenotypes are heritable and mediate fecundity but not fitness in the lepidopteran frugivore Cydia pomonella

The inheritance and functional roles of quantitative traits are central concerns of evolutionary ecology. We report two sets of experiments that investigated the heritability and reproductive consequences of body size phenotypes in a globally distributed lepidopteran frugivore, Cydia pomonella (L.). In our first set of experiments, we tested the hypotheses that (1) body size is heritable and (2) parental body size mediates egg production and offspring survival. Midparent–offspring regression analyses revealed that body mass is highly heritable for females and moderately heritable for males. The contribution of fathers to estimates of additive genetic variance was slightly greater than for mothers. Egg production increased with mean parental size, but offspring survival rates were equivalent. Based on this result, we tested two additional hypotheses in a second set of experiments: (3) male size moderates female egg production and egg fertility and (4) egg production, egg fertility, and offspring survival rate are influenced by female mating opportunities. Females paired with large males produced more eggs and a higher proportion of fertile eggs than females paired with small males. Females with multiple mating opportunities produced more fertile eggs than females paired with a single male. However, egg production and offspring survival rates were unaffected by the number of mating opportunities. Our experiments demonstrate that body mass is heritable in C. pomonella and that size phenotypes may mediate fecundity but not fitness. We conclude that male size can influence egg production and fertility, but female mate choice also plays a role in determining egg fertility.     

Social Rank, Stress, Fitness, and Life Expectancy in Wild Rabbits

 Wild rabbits of the two sexes have separate linear rank orders, which are established and maintained by intensive fights. The social rank of individuals strongly influence their fitness: males and females that gain a high social rank, at least at the outset of their second breeding season, have a much higher lifetime fitness than subordinate individuals. This is because of two separate factors: a much higher fecundity and annual reproductive success and a 50% longer reproductive life span. These results are in contrast to the view in evolutionary biology that current reproduction can be increased only at the expense of future survival and/or fecundity. These concepts entail higher physiological costs in high-ranking mammals, which is not supported by our data: In wild rabbits the physiological costs of social positions are caused predominantly by differential psychosocial stress responses that are much lower in high-ranking than in low-ranking individuals. Received: 23 February 1999 / Accepted: 17 March 1999     

Changes in Age-Related Reproductive Tactics in the Female of the Butterfly, Eurema hecabe

The occurrence of mate solicitation by virgin females was investigated in the butterfly Eurema hecabe. Young (1-day-old) virgin females rarely showed mate solicitation to male model, however, old (at least 6-day-old) virgin females frequently showed such flight. The duration of solicitation was significantly longer in older females than in younger ones. The age-related behavioral change occurs with female oogenesis (Hiroki and Kato 1996), and such behavior may thus be a result of female adaptation to maximize their fecundity. Received: 6 November 1997 / Accepted in revised form: 29 June 1998     

Does testis weight decline towards the Subarctic? A case study on the common frog, Rana temporaria

Interpopulation comparisons of variation in resource availability and in allocation patterns along altitudinal and latitudinal gradients allow insights into the mechanisms shaping the life history of animals. Patterns of between-population differences in female life history traits have been studied intensively across a wide range of taxa, but similar investigations in males have remained scarce. To study if testis weight—a measure of reproductive investment—varies on a geographical scale in anurans, we focussed on the variation in relative testis weight (RelTW) and asymmetry in 22 populations of the common frog Rana temporaria along a 1,600-km latitudinal transect across the Scandinavian peninsula. We found that RelTW decreased towards the north. Body mass and body length both had independent positive effects on testes mass. We found evidence for directional asymmetry (DA) in testis weight with the right testis being larger than the left. The level of DA in testis weight was not related to latitude, but both body mass and testes mass had independent positive effects on asymmetry. We discuss the northwards decrease in RelTW in terms of a decreased reproductive investment as a possible consequence of harsher environmental conditions, and perhaps also, weaker sexual selection in the north than in the south.     

Within-year differences in reproductive investment in laboratory zebra finches (<Emphasis Type="Italic">Taeniopygia guttata</Emphasis>), an opportunistically breeding bird

Reproduction in opportunistically breeding bird species has traditionally been considered non-seasonal with individuals taking advantage of favourable environmental conditions as they arise. However, some studies imply that this opportunistic breeding may be superimposed on an underlying seasonality, which has effects on the readiness to breed when conditions are favourable. The zebra finch (Taeniopygia guttata) is the classic opportunistic breeder and widely used as such in studies. In a series of laboratory-based breeding experiments, we found evidence to suggest that there are seasonal differences in maternal reproductive investment in the zebra finch even when photoperiod, temperature, relative humidity and diet were held constant. Females showed highly significant seasonal differences in clutch size and egg mass with laying order. Clutch size showed a spring/summer peak typical of multi-brooded species in the wild. There was also a significant increase in egg mass with laying order in all seasons except winter. This variation in breeding parameters with season may allow females to adjust investment depending on the potential fitness returns from a given reproductive attempt. These findings also raise a warning about interpreting results of multiple zebra finch breeding experiments that have been carried out in different seasons.     

Plant invasion phenomenon enhances reproduction performance in an endangered spider

Current models in evolutionary ecology predict life history alterations in response to habitat suitability to optimize fitness. Only few empirical studies have demonstrated how life history traits that are expected to trade off against each other differ among environments. In Europe, many salt marshes have been recently invaded by the grass Elymus athericus. Previous studies however showed higher densities of the endangered spider Arctosa fulvolineata (Araneae: Lycosidae) in invaded salt marshes compared to natural habitats, which suggests a lower habitat suitability in the latter. The aim of this study was to determine if this emerging habitat (1) affects the amount of resource acquisition and (2) alters the balance between life history traits that are expected to trade off against each other in this stenotopic salt marsh species. As suggested by theoretical studies, an optimization of fitness by increasing egg size at the cost of decreasing fecundity in unsuitable (i.e., natural) habitats was expected. Females presenting cocoon were then collected in close invaded and natural salt marsh areas within the Mont Saint-Michel Bay (France). By considering female mass as covariate, cocoon mass, number of eggs, and egg volume were compared between both habitats. Clutch mass was strongly determined by female mass in both habitats. Clutch mass was however significantly smaller in the natural habitat compared to the invaded habitat, indicating a higher resource acquisition in the latter. When correcting for female size, fecundity was additionally increased in the invaded habitat through a significant decrease in egg size. This phenotypic response can be explained by differences in habitat structure between invaded and natural habitats: the former offers a more complex litter favoring nocturnal wanderers like A. fulvolineata. The existence of such an adaptive reproduction strategy depending on habitat suitability constitutes an original case of an invasion that favors an endangered species.     

Reduced population size does not affect the mating strategy of a vulnerable and endemic seabird

Bottleneck episodes may occur in small and isolated animal populations, which may result in decreased genetic diversity and increased inbreeding, but also in mating strategy adjustment. This was evaluated in the vulnerable and socially monogamous Monteiro’s Storm-petrel Hydrobates monteiroi, a seabird endemic to the Azores archipelago which has suffered a dramatic population decline since the XVth century. To do this, we conducted a genetic study (18 microsatellite markers) in the population from Praia islet, which has been monitored over 16 years. We found no evidence that a genetic bottleneck was associated with this demographic decline. Monteiro’s Storm-petrels paired randomly with respect to genetic relatedness and body measurements. Pair fecundity was unrelated to genetic relatedness between partners. We detected only two cases of extra-pair parentage associated with an extra-pair copulation (out of 71 offspring). Unsuccessful pairs were most likely to divorce the next year, but genetic relatedness between pair mates and pair breeding experience did not influence divorce. Divorce enabled individuals to improve their reproductive performances after re-mating only when the new partner was experienced. Re-pairing with an experienced partner occurred more frequently when divorcees changed nest than when they retained their nest. This study shows that even in strongly reduced populations, genetic diversity can be maintained, inbreeding does not necessarily occur, and random pairing is not risky in terms of pair lifetime reproductive success. Given, however, that we found no clear phenotypic mate choice criteria, the part played by non-morphological traits should be assessed more accurately in order to better understand seabird mating strategies.     

Intrinsic worker mortality depends on behavioral caste and the queens’ presence in a social insect

According to the classic life history theory, selection for longevity depends on age-dependant extrinsic mortality and fecundity. In social insects, the common life history trade-off between fecundity and longevity appears to be reversed, as the most fecund individual, the queen, often exceeds workers in lifespan several fold. But does fecundity directly affect intrinsic mortality also in social insect workers? And what is the effect of task on worker mortality? Here, we studied how social environment and behavioral caste affect intrinsic mortality of ant workers. We compared worker survival between queenless and queenright Temnothorax longispinosus nests and demonstrate that workers survive longer under the queens’ absence. Temnothorax ant workers fight over reproduction when the queen is absent and dominant workers lay eggs. Worker fertility might therefore increase lifespan, possibly due to a positive physiological link between fecundity and longevity, or better care for fertile workers. In social insects, division of labor among workers is age-dependant with young workers caring for the brood and old ones going out to forage. We therefore expected nurses to survive longer than foragers, which is what we found. Surprisingly, inactive inside workers showed a lower survival than nurses but comparable to that of foragers. The reduced longevity of inactive workers could be due to them being older than the nurses, or due to a positive effect of activity on lifespan. Overall, our study points to behavioral caste-dependent intrinsic mortality rates and a positive association between fertility and longevity not only in queens but also in ant workers.     

Parental conflict and blue egg coloration in a seabird

When both parents provide offspring care, equal sharing of costly parental duties may enhance reproductive success. This is crucial for longlived species, where increased parental effort in current reproduction profoundly affects future reproduction. Indication of reproductive value or willingness to invest in reproduction may promote matching responses by mates, thus reducing the conflict over care. In birds with biparental care, blue-green eggshell color may function as a signal of reproductive value that affects parental effort, as predicted by the signaling hypothesis of blue-green eggshell coloration. However, this hypothesis has not been explored during incubation, when the potential stimulus of egg color is present, and has been little studied in longlived birds. We experimentally studied if egg color affected incubation patterns in the blue-footed booby, a longlived species with biparental care and blue eggs. We exchanged fresh eggs between nests of the same laying date and recorded parental incubation effort on the following 4 days. Although egg color did not affect male effort, original eggshell color was correlated with pair matching in incubation. Exchanged eggshell color did not affect incubation patterns. This suggests that biliverdin-based egg coloration reflects female quality features that are associated with pair incubation effort or that blue-footed boobies mate assortatively high-quality pairs incubating more colorful clutches. An intriguing possibility is that egg coloration facilitates an equal sharing of incubation, the signal being functional only during a short period close to laying. Results also suggest that indication of reproductive value reduces the conflict over care.     

Host genotype and age have no effect on rejection of parasitic eggs

Egg rejection belongs to a widely used host tactic to prevent the costs incurred by avian brood parasitism. However, the genetic basis of this behaviour and the effect of host age on the probability of rejecting the parasitic egg remain largely unknown. Here, we used a set of 15 polymorphic microsatellite loci, including a previously detected candidate locus (Ase64), to link genotypes of female great reed warblers (Acrocephalus arundinaceus), a known rejecter, with their egg rejection responses in two host populations. We also tested whether host female age, as a measure of the experience with own eggs, plays a role in rejection of common cuckoo (Cuculus canorus) eggs. We failed to find any consistent association of egg rejection responses with host female genotypes or age. It seems that host decisions on egg rejection show high levels of phenotypic plasticity and are likely to depend on the spatiotemporal variation in the parasitism pressure. Future studies exploring the repeatability of host responses towards parasitic eggs and the role of host individual experience with parasitic eggs would greatly improve our understanding of the variations in host behaviours considering the persistence of brood parasitism in host populations with rejecter phenotypes.     

Eggshell colour does not predict measures of maternal investment in eggs of <Emphasis Type="Italic">Turdus</Emphasis> thrushes

The striking diversity of avian eggshell colour has long fascinated biologists. Recently, it has been proposed that the blue-green colour of some eggs may function as a post-mating sexually selected signal of female phenotypic quality to their mates to induce higher allocation of paternal care. It has been suggested that maternally deposited yolk carotenoids may be the specific aspect of reproductive quality that the female is signalling via eggshell colour. We use the known properties of the thrush visual system (Turdus sp.) to calculate photon capture for the four single cone photoreceptors, and the principal member of the double cone class for eggs in clutches of two introduced European thrush species (Turdus merula and Turdus philomelos) in New Zealand. We show that differences in the avian-perceived colours of individual eggs are not consistently correlated with different measures of maternal investment in the egg. Given the growing extent of the knowledge between maternal quality, parental investment and eggshell pigmentation across avian taxa, we encourage the use of avian perceptual modelling for testing alternative non-signalling explanations for the structural and physiological basis of these relationships.     

黑岱沟露天煤矿复垦土地野生植物侵入规律及对生态系统的影响

以黄土高原区露天煤矿复垦地为对象,于2005年8月对准格尔旗黑岱沟露天煤矿1992-2005年复垦的364hm2排土场野生植物入侵的种类进行了调查与研究,分析了42个样地上不同野生植物侵入的多度、频度、侵入规律及对生态系统的影响.结果表明:1992-2005年的近15年中,共有115种野生植物自然侵入到排土场内,群落中以1年生植物占优势(＞50%),但多年生植物已比复垦初期(＜5%)增加了很多,在组成上约占总种数的40%～50%,多度高的植物已有多年生植物,且分布于植物群落中.影响复垦植被生长的主要侵入种是披碱草和拂子茅.复垦地野生杂草侵入,表现为①平台侵入种数＞边坡侵入种数,阴坡侵入种数＞阳坡侵入种数;②纯林侵入种数＞混交林侵入种数;③沙棘灌丛侵入种数＞油松侵入种数＞杨树林侵入种数;④黄土覆盖区侵入种数＞土石混排区侵入种数.      

Increase in membrane thickness during development compensates for eggshell thinning due to calcium uptake by the embryo in falcons

We compared membrane thickness of fully developed eggs with those of non-developed eggs in different endangered falcon taxa. To our knowledge, membrane thickness variation during development has never been examined before in falcons or any other wild bird. Yet, the egg membrane constitutes an important protective barrier for the developing embryo. Because eggshell thinning is a general process that occurs during bird development, caused by calcium uptake by the embryo, eggs are expected to be less protected and vulnerable to breakage near the end of development. Thus, egg membranes could play an important protective role in the later stages of development by getting relatively thicker. We used linear mixed models to explore the variation in membrane thickness (n = 378 eggs) in relation to developmental stage, taxon, female age, mass and identity (73 females), egg-laying sequence (105 clutches) and the study zone. Our results are consistent with the prediction that egg membranes are thicker in fully developed eggs than in non-developed eggs, suggesting that the increase in membrane thickness during development may compensate for eggshell thinning. In addition, our data shown that thicker membranes are associated with larger, heavier and relatively wider eggs, as well as with eggs that had thinner eggshells. Egg-laying sequence, female age and the study zone did not explain the observed variation of membrane thickness in the falcon taxa studied. As we provide quantitative data on membrane thickness variation during development in falcons not subjected to contamination or food limitation (i.e. bred under captive conditions), our data may be used as a reference for studies on eggs from natural populations. Considering the large variation in membrane thickness and the multiple factors affecting on it and its importance in the protection of the embryo, we encourage other researchers to include measurements on membranes in studies exploring eggshell thickness variation.     

Sex-biased investment in yolk androgens depends on female quality and laying order in zebra finches (Taeniopygia guttata)

The Trivers–Willard hypothesis predicts sex biases in parental investment according to parental condition. In addition, parents may need to sex bias their investment if there is an asymmetry between the sexes in offspring fitness under different conditions. For studying maternal differential investment, egg resources are ideal subjects because they are self contained and allocated unequivocally by the female. Recent studies show that yolk androgens can be beneficial to offspring, so here we test for sex-biased investment with maternal investment of yolk testosterone (T) in zebra finch (Taeniopygia guttata) eggs. From the Trivers–Willard hypothesis, we predicted females to invest more in male eggs in optimum circumstances (e.g. good-condition mother, early-laid egg), and more in female eggs under suboptimal conditions (e.g. poor-condition mother, late-laid egg). This latter prediction is also because in this species there is a female nestling disadvantage in poor conditions and we expected mothers to help compensate for this in female eggs. Indeed, we found more yolk T in female than male eggs. Moreover, in accordance with our predictions, yolk T in male eggs increased with maternal quality relative to female eggs, and decreased with laying order relative to female eggs. This supports our predictions for the different needs and value of male and female offspring in zebra finches. Our results support the idea that females may use yolk androgens as a tool to adaptively manipulate the inequalities between different nestlings.     

The symbiotic lifestyle and its evolutionary consequences: social monogamy and sex allocation in the hermaphroditic shrimp Lysmata pederseni

Sex allocation theory predicts female-biased sex allocation for simultaneous hermaphrodites with a monogamous mating system. Mating systems theory predicts that monogamy is advantageous in environments where refuges are discrete, scarce, relatively small, and when predation risk is high outside of these refuges. These predictions were tested with the Caribbean shrimp Lysmata pederseni, a simultaneous hermaphrodite which has an early male phase and lives inside tubes of the sponge Callyspongia vaginalis. This host sponge is a scarce resource that, together with the high predation risk typical of tropical environments, should favor monogamy in the shrimp. Field observations demonstrated that shrimps were frequently encountered as pairs within these tube sponges. Pairs were equally likely to comprise two hermaphrodites or one hermaphrodite and one male. Several of these pairs were observed for long periods of time in the field. Experiments demonstrated that hermaphrodites tolerated other hermaphrodites but not males in their host sponge. These results suggest that pairs of hermaphroditic L. pederseni are socially monogamous; they share the same host individual and might reproduce exclusively with their host partners for long periods of time. Nevertheless, males appeared less likely to establish long-term associations with hermaphrodites as indicated by the rate of their disappearance from their hosts (greater than that of hermaphrodites). Sex allocation was female biased in monogamous hermaphrodites. On average, hermaphrodites invested 34 times more to female than to male reproductive structures. Monogamy and female-biased sex allocation seem to be evolutionary consequences of adopting a symbiotic lifestyle in simultaneous hermaphrodites.     

Parker��s sneak-guard model revisited: why do reproductively parasitic males heavily invest in testes?

Alternative reproductive tactics are widespread in males and may cause intraspecific differences in testes investment. Parker's sneak-guard model predicts that sneaker males, who mate under sperm competition risk, invest in testes relatively more than bourgeois conspecifics that have lower risk. Given that sneakers are much smaller than bourgeois males, sneakers may increase testes investment to overcome their limited sperm productivity because of their small body sizes. In this study, we examined the mechanism that mediates differential testes investment across tactics in the Lake Tanganyika cichlid fish Lamprologus callipterus. In the Rumonge population of Burundi, bourgeois males are small compared with those in other populations and have a body size close to sneaky dwarf males. Therefore, if differences in relative testis investment depend on sperm competition, the rank order of relative testis investment should be dwarf males?>?bourgeois males in Rumonge?=?bourgeois males in the other populations. If differences in relative testis investment depend on body size, the rank order of relative testes investment should be dwarf males?>?bourgeois males in Rumonge?>?bourgeois males in the other populations. Comparisons of relative testis investment among the three male groups supported the role of sperm competition, as predicted by the sneak-guard model. Nevertheless, the effects of absolute body size on testes investment should be considered to understand the mechanisms underlying intraspecific variation in testes investment caused by alternative reproductive tactics.     

Cross-species correlation between queen mating numbers and worker ovary sizes suggests kin conflict may influence ovary size evolution in honeybees

During social evolution, the ovary size of reproductively specialized honey bee queens has dramatically increased while their workers have evolved much smaller ovaries. However, worker division of labor and reproductive competition under queenless conditions are influenced by worker ovary size. Little comparative information on ovary size exists in the different honey bee species. Here, we report ovariole numbers of freshly dissected workers from six Apis species from two locations in Southeast Asia. The average number of worker ovarioles differs significantly among species. It is strongly correlated with the average mating number of queens, irrespective of body size. Apis dorsata, in particular, is characterized by numerous matings and very large worker ovaries. The relation between queen mating number and ovary size across the six species suggests that individual selection via reproductive competition plays a role in worker ovary size evolution. This indicates that genetic diversity, generated by multiple mating, may bear a fitness cost at the colony level.     

Loss of legs: is it or not a handicap for an orb-weaving spider?

Leg loss is a common phenomenon in spiders, and according to the species 5% to 40% of the adults can present at least one missing leg. There is no possibility of regeneration after adult moult and the animal must manage with its missing appendages until its death. With the loss of one or more legs, female orb-weaving spiders can be penalized twice: firstly, because the legs are necessary for web construction and secondly, the legs are essential for the control of the prey after its interception by the web. During development, spiders may be also penalized because regeneration has energetic costs that take away resources for survival, growth and reproduction. All these consequences should influence negatively the development of the spider and thus its fitness. We investigated the impact of leg loss in the orb-weaving spider, Zygiella x-notata by studying its frequency in a natural population and web building and prey capture behaviours in laboratory. In field populations, 9.5% to 13%, of the adult females presented the loss of one or more legs; the majority of individuals had lost only one leg (in 48% of cases, a first one). Leg loss seems to affect all the adult spiders, as there is no difference of mass between intact spiders and those with missing leg. Data obtained with laboratory-reared spiders, showed that the loss of legs due to the moult is rare (less than 1%). Considering changes in web design, spiders with missing legs decreased their silk investment, increased the distance between spiral turns but did not change the capture surface of the web. Under our laboratory experimental conditions, spiders with one or two lost legs did not present any difference in prey capture efficiency. In laboratory conditions, spiders with lost leg(s) did not show any difference in egg sac production or in longevity (adult lifespan) compared to intact spiders.