Tisular and population level responses to habitat harshness in sandy beaches: the reproductive strategy of <Emphasis Type="Italic">Donax hanleyanus</Emphasis>

Microscopic analysis and field sampling procedures were used to compare demographic and reproductive strategies of the intertidal wedge clam Donax hanleyanus (Bivalvia: Donacidae) in two exposed sandy beaches with contrasting morphodynamics (reflective vs. dissipative) during 13 consecutive months. Histological analysis showed that: (1) the reproductive cycle of D. hanleyanus was more extended in the dissipative beach, and this was true for all the three pre-active (beginning of gonadal activity), active (maturation) and spawning stages; and (2) males and females showed significantly smaller sizes at sexual maturity at the reflective beach. Even though successive increments in proportion and mature at size were observed, the sigmoid function was significantly steeper at the reflective Arachania for both sexes, suggesting an abrupt transition to maturity. Field sampling revealed a more extended recruitment period at the dissipative beach, where recruits were also significantly more abundant than at the reflective beach. These results give support for the habitat harshness hypothesis, which predicts that in intertidal species capable of sustaining populations across a wide spectrum of physical conditions, such as D. hanleyanus, abundance, recruitment, size at maturity and extent of reproductive and recruitment seasons increase from reflective to dissipative beaches. However, a recent hypothesis suggests that reflective beaches acting as sink populations were not sustained, because mature and spawning individuals of both sexes were found in the reflective beach throughout the study period. Thus, we suggest that post-settlement processes are critical in modulating population patterns for this bivalve.     

Habitat harshness and morphodynamics: life history traits of the mole crab Emerita brasiliensis in Uruguayan sandy beaches

The habitat harshness hypothesis (HHH) postulates that in reflective beaches the harsh environment forces organisms to divert more energy towards maintenance and they therefore have lower abundance, fecundity, growth and survival rates than in dissipative beaches. Recent investigations have tested this hypothesis through single comparisons of only two beaches, and thus the observed trends in population level variables cannot be attributed incontestably to the beach state, but only to location. Here, abundance, reproduction, recruitment, population structure and body size of the intertidal mole crab Emerita brasiliensis were compared between populations from eight microtidal exposed sandy beaches with contrasting morphodynamics, sampled bimonthly during 22 months throughout the 180 km Uruguayan Atlantic coast. Physical variables and compound indices of the beach state were used to categorize sandy beaches. The results of this bi-annual large-scale analysis were fully consistent with the predictions of the HHH: abundance (total and population components), duration of the reproduction and recruitment seasons and the individual size of megalops and females of the mole crab E. brasiliensis decreased from dissipative to reflective beaches. This was reflected by linear or, mostly, nonlinear relationships between biological and both physical variables and compound indices of beach state. In conclusion, this multi-beach sampling provides compelling evidence of a consistent response of demographic and life history traits of an intertidal beach species to morphodynamic characteristics.     

Maturity,fecundity, and reproductive cycle of the spotted ratfish, <Emphasis Type="Italic">Hydrolagus colliei</Emphasis>

Size at maturity, fecundity, and reproductive periodicity were estimated for the spotted ratfish, Hydrolagus colliei (Lay and Bennett, 1839), off the coast of California, Oregon, and Washington. Maximum body size and size at median maturity were greater for females than males. Skeletal muscle concentrations of the steroid hormones testosterone (T) and estradiol (E₂) predicted similar, but slightly smaller sizes at maturity than the morphological criterion. Stage of maturity for males was estimated identically using internal organs or external secondary sexual characters, thus allowing non-lethal maturity assessments. Size at median maturity was greater north of Point Conception for females, and north of Cape Mendocino for males. Peak parturition occurred from May to October, with increased concentrations of E₂ in skeletal muscle of females correlating with ovarian recrudescence during November to February. No significant seasonal trends in female T were apparent, but mean female 11-ketotestosterone (11KT) was 300% greater in April than any other month during the parturition season. There was a marginal evidence for increased number and size of ova with maternal size. Extrapolation of the hypothesized 6 to 8-month egg-laying season to observed mean parturition rates of captive specimens yielded an estimated annual fecundity of 19.5–28.9 egg cases. Differences in fecundity among higher taxonomic classifications of chondrichthyans were detected with chimaeriform fishes more fecund than lamniform, myliobatiform, squaliform, and rhinobatiform fishes, and less fecund than rajiform fishes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Threshold to maturity in a long-lived reptile: interactions of age, size, and growth

Thresholds to sexual maturity—either age or size—are critical life history parameters. Usually investigated in short-lived organisms, these thresholds and interactions among age, size, and growth are poorly known for long-lived species. A 34-year study of captive green turtles (Chelonia mydas) that followed individuals from hatching to beyond maturity provided an opportunity to evaluate these parameters in a long-lived species with late maturity. Age and size at maturity are best predicted by linear growth rate and mass growth rate, respectively. At maturity, resource allocation shifts from growth to reproductive output, regardless of nutrient availability or size at maturity. Although captive turtles reach maturity at younger ages than wild turtles, the extensive variation in captive turtles under similar conditions provides important insights into the variation that would exist in wild populations experiencing stochastic conditions. Variation in age/size at maturity should be incorporated into population models for conservation and management planning.     

The reproductive cycle of the Thecosomatous pteropod Limacina retroversa in the western South Atlantic

The reproductive cycle of Limacina retroversa in the Argentine Sea was studied by gonadal analysis. Samples were collected in 1978–1979. The relation between gonadal maturity and size is dependent on the season and the geographical area. Evidence of in situ reproduction was found at the beginning of spring and at the end of summer, indicating that the life cycle in the area consists of two generations per year. In accordance with the season of birth, both generations develop different strategies. Individuals born in spring mature early and reproduce before the end of summer. The offspring born in summer survive the cold season without reaching sexual maturity and reproduce in the following spring.     

Climate-induced reaction norms for life-history traits in pythons

Climate change modelers predict increasingly frequent "extreme events," so it is critical to quantify whether organismal responses (such as reproductive output) measured over the range of usual climatic conditions can predict responses under more extreme conditions. In a 20-year field study on water pythons (Liasis fuscus), we quantified the effects of climatically driven annual variation in food supply on demographic traits of female pythons (feeding rate, body size, body mass, and reproductive output). Reaction norms linking food supply to feeding rates and residual body mass were broadly linear, whereas norms linking food supply to female body size became curvilinear when a dramatic (flooding-induced) famine reduced the mean body size at sexual maturity. Thus, the reaction norms recorded over 16 years of mal" (albeit highly variable) climatic conditions gave little insight into the population's response to a more extreme nutritional crisis.     

Food or sex—males and females in a sex role reversed pipefish have different interests

In a sex role reversed pipefish, Syngnathus typhle, we found that basic life history allocations were directly influenced by sexual selection. We investigated time allocation to foraging and mating, respectively, in a choice experiment, giving males and females, of small or large body size, a choice between food and a potential partner. We found that males were more interested in foraging than mating, i.e., were more frequently observed in front of the food than in front of the partner, whereas females were more interested in the potential partner. This reflects sexual selection operating differently on the two sexes, as males and females are relatively similar in other life history traits, such as growth, mortality, age of maturity, dispersal, and parental expenditure. Moreover, large individuals allocated more time to mating activities, small to feeding. Individuals more interested in mating compared to food were subsequently more critical when given a choice between a large (high-quality) and a small (low-quality) partner, whereas individuals more interested in food were not selective. These findings are consistent with our predictions: sex-role reversed males can be relatively sure of achieving one or more matings, and should allocate more time to feeding and, hence, to parental investment, growth and/or future reproduction. Females, on the other hand, have more uncertain mating prospects and should allocate time to imminent reproductive activities, thereby foregoing other life history traits such as growth and future egg production. By this, they also sacrifice future fecundity and attractiveness.     

Morphodynamic responses of nourished beaches in SW Spain

Coastal erosion in SW Spain is affecting man-made structures and beaches that represent an important economic resource in the area. In the last decade the Spanish government carried out several nourishment works that have limited durability. Most of the artificial beach fills consist of a spill of natural dredged sand on the visible beach, leading to a flat, artificial berm with an important seaward slope and narrow foreshore. As a result, the initial dissipative profile was transformed into a fully reflective one. A beach monitoring program was carried out to record morphological evolution after the nourishment works. Several field assessments of disturbance depth were also made to characterize beach morphodynamics of a nourished beach (Rota) and a natural dissipative one (Tres Piedras), whose slope was similar to the pre-nourishment gradient of Rota beach. Natural dissipative beaches were characterized by spilling breakers that did not significantly affect bottom sand. The severe erosion recorded in the nourished zones was related to the new morphodynamic regime acting on these beaches, which was controlled by high erosive plunging breakers associated with high foreshore slopes. In conclusion, other nourishment practices should be used, better adapted to the natural beach morphodynamics of the zone, taking also into account the original grain size and density of the beach sands, in order to obtain more durable artificial beaches.     

Responses of fish assemblage structures to sandy beach types in Kyushu Island, southern Japan

To determine differences in fish assemblage structures between beach morphodynamic types on an exposed sandy shore, three beach types (reflective, intermediate, and dissipative) were sampled at Fukiagehama Beach, Kyushu Island, southern Japan, in May, August, and November 2006 and 2007. Distinct differences in physical conditions and the amount of major prey invertebrates, such as copepods, mysids, and amphipods, were recognized between the three beach types. The reflective beach was characterized mainly by turbulent swashes, coarse sediment, and a low abundance of major prey invertebrates. In contrast, the dissipative beach was characterized mostly by benign swashes, fine sediment, and a high abundance of prey invertebrates. The intermediate beach tended toward an intermediate condition between the reflective and dissipative beaches. The fish assemblage structures also differed clearly between the three beach types, with species and individual numbers being greatest on the dissipative beach and lowest on the reflective beach. A similarity index indicated differences in species composition of the assemblages between the beach types. Such differences arose primarily from the differential distributions of two dominant trophic groups, zooplankton (mostly copepods) and epibenthic crustacean (mainly mysids and amphipods) feeders, between the beach types, both groups tending to increase in species and individual numbers from the reflective beach to the dissipative beach. In addition, the mean standard length of all fish collected in each month tended to decrease toward the dissipative beach. These results suggested that beach types strongly influence sandy beach fish assemblage structures.     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Reproductive biology of the holothurians from the major communities of the New Caledonian Lagoon

The reproductive biology of nine species of aspidochirote holothurians from the major communities of the New Caledonian lagoon has been analysed. The data were obtained from monthly sampling from 1978 to 1982 of their populations on reef slopes and passes for Holothuria nobilis, H. fuscogilva, H. fuscopunctata and Thelenota ananas, in the inner lagoon for Stichopus variegatus, H. scabra and H. scabra versicolor and on reef flats for Actinopyga mauritiana, A. echinites and H. atra. The morphology and anatomy of the gonads and their annual cycle are described. The reproductive cycles, composed of the five phases of gonad growth, maturing, spawning, post-spawning and resting, were determined from the changes in the sexual stages, the gonad indices and the percentage of indeterminate sex. Three main reproductive patterns are recognized, annual cycles with spawning occurring during the warm season or during the cool season and a semiannual cycle. Other population parameters, such as the size of the mature oocytes, the mean size of the individuals at first sexual maturity, and absolute and relative fecundities were calculated and related to the mean size of the species, the taxonomy and the ecology. The reproductive strategies of the species show a gradient that is probably related to the stability of the biotopes, with smaller species living in the more intrable outer reef flats and larger species in deeper lagoon floors.     

Live fast, die young: flexibility of life history traits in the fat-tailed dwarf lemur (Cheirogaleus medius)

The fat-tailed dwarf lemur, Cheirogaleus medius, occurs in ecologically very different habitat types (rainforest and dry forest) across Madagascar. Its extraordinary biological characteristics, such as monogamy and long-term hibernation, allow us to investigate behavioral, ecological, and physiological flexibility of this species in populations across different ecological environments. This study aims to determine whether different life history and physiological traits show variation in adaptation to the differing ambient conditions or are conservative and influenced more by the organism's evolutionary history. We compared body masses, life history traits, social organization, and hibernation duration of two populations of C. medius, one from a littoral rainforest and one from the dry deciduous forest. We revealed clear geographical differences in the length of hibernation duration as well as, more surprisingly, in life history traits. We found that this species' reproductive strategies seem to be highly flexible. Animals in the rainforest can spend more time in the active state due to a shorter hibernation period, but have, in general, a shorter life expectancy due to higher mortality rates. Hence, they seem to maximize their total reproductive output with higher reproductive rates (larger litter sizes, greater number of litters). Home ranges and social organization, on the other hand, did not vary between habitats, suggesting that the general requirements of this species are independent of environmental conditions. In conclusion, some life history traits, formerly assumed to be genetically fixed parameters of primate species, prove in fact to be highly flexible. Different populations of the same species show distinct adaptations according to the prevailing conditions in order to maximize individual reproductive output.     

Reproductive development of the barnacle Chthamalus malayensis in Hong Kong: implications for the life-history patterns of barnacles on seasonal, tropical shores

Hong Kong, lying just below the Tropic of Cancer (22°17′N, 114°09′E), experiences a strongly seasonal environment, with a cool almost temperate winter and a hot, tropical, summer. Histological sectioning of the gonads of the high-shore barnacle, Chthamalus malayensis Pilsbry, showed a seasonal trend in the development of its reproductive organs. Four stages of female gonad development were identified according to the cell types present: post-spawning, resting, growth and mature stages. The female gonad was mature from April to November, which was related to seawater temperatures, and entered a resting phase from December to March. Although the male gonad showed a seasonal developmental trend and reached maximum maturity in summer, the seminal vesicles were full of spermatozoa and functional throughout the year. The reproductive season of this species is therefore solely dependant upon the maturity of the female gonad. The estimated maximum number of broods per year was up to 10 and the maximum number of eggs produced per brood can reach 3,000 eggs. The minimum size for female gonad maturity was 6 mm rostro-carinal diameter (RCD) at which size, the barnacles were ~6-month old. Sperm production occurred at a smaller size (2 mm=2-month old). Compared with Chthamalus montagui and Chthamalus stellatus from temperate regions, C. malayensis produced a greater number of broods per year, had a longer reproductive period and faster gonad development. Chthamaloid barnacles in tropical regions may, therefore, invest more energy per year in reproduction during their life span. Contrary to the seasonal gonad developmental pattern of C. malayensis in the present study, however, C. malayensis in Singapore (which experiences only slight seasonal variation) had mature female and male gonads throughout the year, further supporting the strong role of climatic conditions effecting the reproductive biology of barnacles.     

Female egg investment in relation to male sexual traits and the potential for transgenerational effects in sexual selection

Life-history theory predicts that individuals should increase their reproductive effort when the fitness return from reproduction is high. Females mated with high-quality males are therefore expected to have higher investment than females mated with low-quality males, which could bias estimates of paternal effects. Investigating the traits females use in their allocation decisions and the aspects of reproduction that are altered is essential for understanding how sexual selection is affected. We studied the potential for differential female allocation in a captive population of a precocial bird, the Chinese quail, Coturnix chinensis. Females paired with males with large sexual ornaments laid larger, but not more, eggs than females paired with males with small sexual ornaments. Furthermore, female egg mass was also significantly positively affected by male testis size, probably via some unknown effect of testis size on male phenotype. Testis size and ornament size were not correlated. Thus, both primary and secondary male sexual traits could be important components of female allocation decisions. Experimental manipulation of hormone levels during embryonic development showed that both male and female traits influencing female egg size were sensitive to early hormone exposure. Differences in prenatal hormone exposure as a result of maternal steroid allocation to eggs may explain some of the variation in reproductive success among individuals, with important implications for non-genetic transgenerational effects in sexual selection.Communicated by C. Brown     

Sexual reproduction of the scyphomedusa Aurelia aurita in relation to temperature and variable food supply

The effects of food availability and temperature on sexual maturation and female reproductive output of the scyphomedusa Aurelia aurita was examined in two populations from the contrasting environments of Southampton Water and Horsea Lake, England. Trends in oogenesis and subsequent reproductive output differed markedly between the two populations. In Southampton Water, the onset of sexual maturation occurred earliest in the larger medusae, but eventually all females became ripe, the smallest being 45 mm bell diameter (BD). The decrease in minimum size at maturity was correlated with increasing temperature. In A. aurita from Horsea Lake, size at maturity varied on a seasonal basis, with the smallest ripe female being only 19 to 20 mm BD. There were spring and autumn periods of sexual maturation in this population. During the autumn period, it is likely that food limitation was playing a more critical role in determining medusa size, with decreasing temperature indirectly affecting A. aurita by limiting primary and secondary production. In similar-sized ripe medusae, fecundity was greater in Southampton Water, but the planula larvae produced were significantly smaller than those in Horsea Lake. It is suggested that in Horsea Lake, the quality of the larvae are greater in terms of biochemical content to ensure survival of the few gametes produced (i.e. K-strategy). Comparison of the reproductive effort of the two A. aurita populations revealed that medusae from Southampton Water, which experience greater food availability, are able to direct more energy to reproduction than Horsea Lake medusae. In the latter, A. aurita medusae appear to partition the available food resources into either somatic growth (and therefore increased future fecundity) when food is abundant, or reproductive growth when food is scarce. Received: 24 June 1997 / Accepted: 23 March 1998     

Reproductive biology of the antipatharian black coral Antipathes fiordensis in Doubtful Sound, Fiordland, New Zealand

The reproductive biology of Antipathes fiordensis Grange, a species endemic to south-western New Zealand, was followed from April 1994 to May 1995. Ten colonies were individually tagged in Doubtful Sound and sampled on a monthly basis in order to determine their reproductive activity. The fecundity of each of the five tagged female colonies was determined by estimating the total number of polyps per colony from photographs of each colony and by planimetry, the proportion of gravid polyps per colony, and the mean number of oocytes per gravid polyp. In addition, 56 colonies were sampled in March 1995 to estimate the sex ratio, height at sexual maturity, and mean sizes of females and males. A. fiordensis was found to be a dioecious species which, in the absence of gonads in the polyps, has no obvious external morphological differences between the sexes. Broadcast spawning of gametes is the likely mode of reproduction. Gametogenesis began in November 1994 and was highly synchronous within and between colonies, with spawning occurring in March 1995. The sex ratio in adults was 1:1. Colonies reached sexual maturity between the heights of 70 and 105 cm which, based on existing estimates of growth rate, corresponds to a minimum age for sexual maturity of about 31 yr. The largest oocytes measured ranged from 100 to 140 μm in size. Female colonies produced between 1.3 and 16.9 million oocytes, with the larger colonies dominating the reproductive output of the population. Received: 23 June 1997 / Accepted: 1 August 1997     

Egg testosterone affects wattle color and trait covariation in the ring-necked pheasant

Transfer of maternal hormones to the eggs is a major source of offspring phenotypic variation. The developmental and organizational effects of egg hormones can extend into adulthood and affect behavioral and morphological traits involved in sexual and reproductive behavior, with important consequences for offspring fitness. In this study, we injected testosterone (T) in egg albumen of captive ring-necked pheasant (Phasianus colchicus) eggs. We then assessed the consequences for chick growth, cell-mediated immunity, and multiple male secondary sexual traits at maturity by comparison with a control group. We also compared the covariation between traits in the two experimental groups. We found that control males had redder wattles than males from T-injected eggs, suggesting that attractiveness and reproductive success of the offspring might vary depending on maternal transfer of T to the eggs. T treatment also modified the covariation between cell-mediated immunity and wattle coloration and between the area of the wattle and the expression of another secondary sexual trait, the ear tufts. These effects are likely to translate into fitness differences among the offspring if mate acquisition depends on the simultaneous expression of several traits that are differentially affected by the same maternal contribution. Maternal effects mediated by egg hormones might affect the fitness of the offspring not only by directional modification of phenotypic traits, but also by facilitating or inhibiting their covariation. This suggests the possibility that female choice based on the relative expression of multiple secondary sexual traits exerts a pressure on how maternal transfer of androgens contributes to developmental programs.     

Growth and population dynamic model for the non-zooxanthellate temperate solitary coral Leptopsammia pruvoti (Scleractinia, Dendrophylliidae)

In corals where complex life history processes decoupling age from size (e.g., fragmentation, fusion, partial colony mortality) are rare or clearly detectable, individual age may be determined from size, and age-based growth and population dynamic models may be applied. One example is the solitary Mediterranean coral Leptopsammia pruvoti Lacaze-Duthiers 1897, whose population size and structure, and growth rates were determined at Calafuria (43°28′N and 10°20′E), Ligurian Sea, from December 2007 to June 2009. Growth rate decreased with increasing size. The growth curve derived from field measurements confirmed the one obtained by growth band analysis. The frequency of individuals decreased exponentially with age, indicating a steady state population. Turnover time was 2.3 years. Maximum life span was 13 years. Most reproductive output was from intermediate age classes (6 years), while older individuals (>7 years), although having higher fecundity, were rare and accounted for a minority of population reproductive output. In comparison with other solitary dendrophylliids, L. pruvoti life strategy was characterized by a reproduction with r-strategy correlates (high fecundity, short embryo incubation, small planula size, fast achievement of sexual maturity), and a rate of demographic renewal occurring halfway along the r–K continuum (intermediate turnover time and maximum longevity).     

Analysis of the relationship between growth and sexual maturation in Phacosoma japonicum (Bivalvia: Veneridae)

The relationship between shell growth and sexual maturation was studied in the venerid bivalve Phacosoma japonicum (Reeve) based on specimens from six populations around the Japanese coast in 1991 and 1992. A distinct latitudinal variation in the patterns of shell growth and gonad development was detected. Speciments from northern populations are characterized by slower rates of gonad development, later offset of interval of shell growth, and larger shell size at a given age than those from southern populations, excluding the population from the Ariake Bay, Kyushu. These data indicate the presence of a tradeoff between reproductive effort and continued growth in this species. However, in all populations bivalves attain sexual maturity before ending shell growth. First sexual maturity occurs at a shell size of about 60% of the maximum asymptotic shell height. Maximum reproductive effort appears to start when the energy available for shell growth (i.e., the yearly growth rate of shell weight) attains a maximum. These relationships between shell growth and sexual maturation were also confirmed in some other bivalve species.