Reproductive plasticity in mole crabs, Emerita brasiliensis, in sandy beaches with contrasting morphodynamics

Ovarian macroscopical analysis, histological validation and field sampling procedures were used to evaluate the variability in reproductive traits of the mole crab Emerita brasiliensis Schmitt, 1935 (Decapoda: Hippidae) in two exposed sandy beaches of Uruguay with contrasting morphodynamics. All developmental stages involved in the complex life cycle exhibited lower abundance, individual size and temporal occurrence in a harsh reflective beach, compared with a more benign dissipative environment. In addition, this population showed more compressed events of the reproductive cycle (e.g oogenesis, encounter of potential mate and female parental care) and recruitment period. However, the beginning of the vitellogenesis, ovary maturity, male sexual differentiation and ending of spermatogenesis occurred at smaller sizes under dissipative conditions. These results disagree with the recent findings of delayed sexual maturity in dissipative beaches. We postulate that, in dissipative conditions, high food availability might allow an overlapping of reproductive and moulting processes, and thus females may reach optimum size and sexual maturity with fewer moults than in reflective beaches. Hence, reproductive responses must be considered not only in relation to environmental harshness, but also in the context of life history traits and their phylogenetic and allometric constraints. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The ecology of two sandy beaches in south west India. II. Notes on Emerita holthuisi

The mole crab Emerita holthuisi Sankolli occurred on two beaches, at Shertallai and at Cochin, in southwest India. Observations on the size-frequency of samples for Shertallai suggest that there were two main periods of recruitment, one in the pre-monsoon,months of February and March, and one during the monsoon, although there may have been some recruitment throught the year. At Cochin, E. holthuisi was present in the pre-monsoon period, but disappeared from the beach during the monsoon.     

Growth,mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches

We studied some aspects of the population dynamics of the yellow clam Mesodesma mactroides (Deshayes, 1854) which inhabits the eastern sandy beaches of Uruguay. Observations were made from March 1983 through March 1985 (harvesting season), and additional data from January 1988 to December 1989 (closed season) were also included in the analyses. Parameters of growth and mortality were estimated by methods that make use of age or length information. An age/length relationship was constructed by counting growth rings. Results were consistent with those obtained with modal class progression analysis and other length/frequency methods. The growth curve obtained from the age/length relationship did not account for seasonality in growth rate, and a seasonal growth model explained growth oscillations in a better way. The instantaneous coefficient of natural mortality (M) estimated for the closed season was 1.64, whereas an increase in fishing mortality (F) was observed during 1984. Age composition did not change during the 2 yr of study, and the first 6 mm class accounted for over 50% of the population. The main contribution of recruits was observed between late summer and early fall (February to April), and the recruitment pattern suggested one main settlement period per year. Growth and recruitment success appeared to be regulated by density-dependence processes. Some implications for management are considered in this context. Methodological aspects concerning the reliability of empirical equations and programs for estimating growth and mortality are discussed.     

Larval settlement and metamorphosis in the sand crab Emerita talpoida (Crustacea: Decapoda: Anomura)

The effects of various potential cues on the survival and time to metamorphosis of larvae of the sand crab Emerita talpoida (Say) were examined. Zoeal duration, which ranged from 25 to 43 d, was not correlated with subsequent megalopal survival but had a weak, negative correlation with the duration of the megalopal stage. Neither food, sand nor water from containers with adults directly affected megalopal survival or duration. In a second experiment, the presence of sand both increased the probability of survival and shortened the duration of late zoeal stages. This is the first experimental evidence for delayed zoeal metamorphosis in a decapod. Settlement of E. talpoida appears to occur primarily during the zoeal phase, not at the megalopal stage.     

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.     

Prediction of extinction in plants: interaction of extrinsic threats and life history traits

The global extinction of species proceeds through the erosion of local populations. Using a 60-year time series of annual sighting records of plant species, we studied the correlates of local extinction risk associated with a risk of species extinction in the Park Grass Experiment where plants received long-term exposure to nutrient enrichment, soil acidification, and reductions in habitat size. We used multivariate linear models to assess how extrinsic threats and life history traits influence extinction risk. We investigated effects of four extrinsic threats (nitrogen enrichment, productivity, acidification, and plot size) as well as 11 life history traits (month of earliest flowering, flowering duration, stress tolerance, ruderalness [plant species' ability to cope with habitat disturbance], plant height, diaspore mass, seed bank, life form, dispersal mode, apomixis [the ability for a species to reproduce asexuall through seeds], and mating system). Extinction risk was not influenced by plant family. All of the 11 life history traits except life form and all threat variables influenced extinction risk but always via interactions which typically involved one threat variable and one life history trait. We detected comparatively few significant interactions between life history traits, and the interacting traits compensated for each other. These results suggest that simple predictions about extinction risk based on species' traits alone will often fail. In contrast, understanding the interactions between extrinsic threats and life history traits will allow us to make more accurate predictions of extinctions.     

Zonation of the interstitial polychaeteStygocapitella subterranea (Stygocapitellidae) in European sandy beaches

The zonation ofStygocapitella subterranea Knoellner (Polychaeta, Stygocapitellidae) in sandy beaches is compared for several localities along the European coasts. In tidal beaches the species occupies a zone near average high tide level (AHTL), whereas in non-tidal beaches it is found at a fairly long distance from the water line. The regularities observed inS. subterranea indicate that the interstitial fauna may prove well suited for the establishment of a general scheme of zonation in sandy beaches analogous to that for rocky shores proposed byStephenson andStephenson (1949).     

Seasonal polyphenism in life history traits: time costs of direct development in a butterfly

Insects with two or more generations per year will generally experience different selection regimes depending on the season, and accordingly show seasonal polyphenisms. In butterflies, seasonal polyphenism has been shown with respect to morphology, life history characteristics and behaviour. In temperate bivoltine species, the directly developing generation is more time-constrained than the diapause generation, and this may affect various life history traits such as mating propensity (time from eclosion to mating). Here, we test whether mating propensity differs between generations in Pieris napi, along with several physiological parameters, i.e. male sex pheromone synthesis, and female ovigeny index and fecundity. As predicted, individuals of the directly developing generation—who have shorter time for pupal development—are more immature at eclosion; males take longer to synthesise the male sex pheromone after eclosion and take longer to mate than diapause generation males. Females show the same physiological pattern; the directly developing females lay fewer eggs than diapausing females during the first days of their life. Nevertheless, the directly developing females mate faster after eclosion than diapausing females, indicating substantial adult time stress in this generation and possibly an adaptive value of shortening the pre-reproductive period. Our study highlights how time stress can be predictably different between generations, affecting both life history and behaviour. By analysing several life history traits simultaneously, we adopt a multi-trait approach to examining how adaptations and developmental constraints likely interplay to shape these seasonal polyphenisms.     

Extremes, plasticity, and invariance in vertebrate life history traits: insights from coral reef fishes

Life history theory predicts a range of directional generic responses in life history traits with increasing organism size. Among these are the relationships between size and longevity, mortality, growth rate, timing of maturity, and lifetime reproductive output. Spanning three orders of magnitude in size, coral reef fishes provide an ecologically diverse and species-rich vertebrate assemblage in which to test these generic responses. Here we examined these relationships by quantifying the life cycles of three miniature species of coral reef fish from the genus Eviota (Gobiidae) and compared their life history characteristics with other reef fish species. We found that all three species of Eviota have life spans of < 100 days, suffer high daily mortality rates of 7-8%, exhibit rapid linear growth, and matured at an earlier than expected size. Although lifetime reproductive output was low, consistent with their small body sizes, short generation times of 47-74 days help overcome low individual fecundity and appear to be a critical feature in maintaining Eviota populations. Comparisons with other coral reef fish species showed that Eviota species live on the evolutionary margins of life history possibilities for vertebrate animals. This addition of demographic information on these smallest size classes of coral reef fishes greatly extends our knowledge to encompass the full size spectrum and highlights the potential for coral reef fishes to contribute to vertebrate life history studies.     

Abnormal reproduction of the sand crab Emerita analoga in the vicinity of a nuclear generating station in Southern California

Female sand crabs (Emerita analoga) were sampled from 12 beaches along a 350 km stretch of Southern California coastline during the reproductive season in July, 1982. The percentage of mature females with egg masses and the condition of those egg masses were examined at each site. The proportion of mature females with egg masses varied only slightly between beaches. However, at sites within about 20 km of the San Onofre Nuclear Generating Station, a high percentage of females carried egg masses consisting of empty egg shells. A transplant experiment was conducted. Females with empty egg masses were taken from the 6.5 km N study site, where only 5% of the mature females carried normal egg masses, and transferred to running unfiltered seawater in Santa Barbara. After 2 wk, about a third of the transplanted females carried normal egg masses, while at the same time the proportion of mature females with normal egg masses remained at 5% on the beach of origin. Heated water discharged from the nuclear generating station is probably not the cause of the abnormal reproduction of sand crabs along the nearby coast. More likely causes are: (1) runoff of agricultural pesticides from a creek 3 km north of the nuclear generating plant; (2) release of metals from corroding cooling pipes, and/or (3) increased turbidity of the nearshore waters.     

Annual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus

Interannual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus (De Haan) was studied for a 7-yr period (1981 to 1987) in an intertidal mudflat in the estuary of the Tatara-Umi River, Fukuoka, Japan. Density of juveniles fluctuated greatly, and maximum annual density was low in 1981 to 1984 (22.0 to 44.0 m^-2) and very high in 1985 to 1987 (98.5 to 188.6 m^-2). In 1980, recruits may have been very few, though there is no data on or before 1980, because no crab of the 1980 cohort was found during the study period. Density of adult crabs was almost stable (11.0 to 32.6 m^-2), perhaps because of density-dependent mortality. Under the high density conditions prevailing from 1985 to 1987 crabs grew slowly and longevity was shorter. Male chela and female abdomen were dwarfish, and females produced few and small broods. The changes of life-history traits under high density conditions were thought to result from strong intraspecific competition for space.     

The effects of size-dependent predation risk on the interaction between behavioral and life history traits in a stream-dwelling isopod

In field surveys, laboratory observations and field-based assays of behavior, I examined the effects of size-dependent predation risk on the interaction between size at reproductive maturity and maternal care behavior in the stream-dwelling isopod, Lirceus fontinalis. L. fontinalis exhibit population-specific sizes at reproductive maturity which result in population differences in predation risk during the adult phase. Females from streams containing salamander larvae (that prefer small prey) mature at large sizes and then become relatively safe from predation. Females from streams containing fish (that consume all size classes of prey equally) mature at small sizes and remain at risk. I tested whether these differences in expected survival were reflected in the behavior of females during the maternal phase (i.e., the period during which females exhibit maternal care). Female L. fontinalis carry developing juveniles inside a brood pouch. I simulated predatory attacks on gravid female L. fontinalis from the different population types and found that female behavior correlated with population differences in risk. When “attacked”, females from streams with predatory fish (that experience high risk to adult females) released juveniles from the brood pouch, whereas females from populations with predatory salamander larvae (that pose relatively little risk to adult females) did not release juveniles. I discuss the results with reference to the joint evolution of behavioral and life history traits. Received: 6 March 1996 /Accepted after revision: 12 August 1996     

Embryogenesis of decapod crustaceans with different life history traits,feeding ecologies and habitats: a fatty acid approach

Variations in embryo size and fatty acid (FA) dynamics during embryogenesis were evaluated in deep-sea pandalids and portunid swimming crabs from the Portuguese continental margin and Madeira Island slope and compared with previous data on neritic and deep-sea lobsters and shrimps (collected between February 2001 and March 2004). Inter-specific variations in embryo size seem to be dictated primarily by phylogeny rather than by differences in reproductive or early life history traits. FA reserves were significantly correlated with embryo size (P < 0.001). Principal component analysis revealed differences among three groups (1—neritic caridean shrimps, 2—deep-sea pandalids of the genus Plesionika, and lobsters, 3—portunid crabs and the deep-sea pandalid Chlorotocus crassicornis, Costa 1871). Group 1 was clearly separated by PC1 mainly due to the higher percentage of essential C₁₈ (linoleic and linolenic acids) and C₂₀ (namely eicosapentaenoic) polyunsaturated FA (specific markers of primary producers). PC2 separated Group 2 from Group 3 due to differences in the percentage of several saturated FA (including odd-numbered FA—bacterial markers) and C₁₈ monounsaturated FA (namely 18:1n − 9, a general marker of carnivory). Therefore, these differences among groups seem to result from distinctions in diet and ecological niche. Intra-specific differences in FA composition between western and southern Plesionika martia martia (A. Milne-Edwards, 1883) populations may reflect higher water temperatures on the south sub-tropical coast. Lobster embryonic development was more demanding of lipid energy than that of the other decapod species, which may reflect an evolutionary trend in decapod taxa related to an increasing degree of lecithotrophy. However, a lower FA catabolism can be interpreted as an enhanced independence of the newly hatched larvae from external energy sources. Higher FA content at hatching and, as a consequence, a greater independence from the external environment should increase the chances of larval survival. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Effect of intraspecific density on life history traits and population growth rate of Neanthes arenaceodentata (Polychaeta: Nereidae) in the laboratory

The effects of intraspecific density on life history traits and population dynamics of the nereid polychaete Neanthes arenaceodentata Moore were assessed in a laboratory experiment. Survival, growth, and fecundity were measured for one generation of worms at densities of 40, 80 and 160 worms per 840 cm² (1x, 2x, and 4x treatments, respectively). Density did not affect size (prior to pairing), percentage of worms paired, time to pairing, or size of mature paired males. Density did have a significant negative effect on survival, size of mature paired females, time to spawning, percentage of females that reproduced, and number of eggs per reproducing female. As density increased, mean survival was 90.0, 80.8 and 74.0%; mean size of mature females was 52.2, 49.2 and 48.1 segments; mean time to spawning was 100.6, 102.4 and 109.4 d; and mean fecundity was 881, 622, and 598 eggs per female, for 1x, 2x and 4x treatments, respectively. Increased density reduced the potential population growth rate, ; for a given rate of larval survivorship, was lower in the 2x and 4x treatments than the 1x treatment. Analysis of sensitivity of to changes in survivorship indicated that population growth rate at the highest density was sensitive to both changes in larval survivorship and the probability of producing a successful brood, although at low densities, was sensitive only to changes in larval survivorship. We attribute these density effects to aggressive bahavioral interactions between the worms, primarily the adults. This experiment identifies key life history traits that could be measured in future experiments to test population level responses of N. arenaceodentata populations to pollutants, both in the laboratory and in the field.Contribution No. 820 of the U.S. Environmental Protection Agency, Environmental Research Laboratory and Contribution No. 189 of the Marine Sciences Institute, University of Connecticut     

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.     

Early life history traits of the gold-eye rockfish, Sebastes thompsoni, in relation to successful utilization of drifting seaweed

The gold-eye rockfish Sebastes thompsoni lives in association with drifting seaweed during the early developmental period. Early life history traits of the gold-eye rockfish were studied by means of otolith analysis of larvae and juveniles associated and unassociated with drifting seaweed in the waters off southern Sanriku, Tohoku, northern Japan, from 1993 to 1995.␣Daily growth increments and the existence of an extrusion check were confirmed by both a series of tetracycline marking experiments and an analysis of the relation between the number of increments and the number of days after extrusion. Larvae and juveniles associated with drifting seaweed occurred from late May to late July, with a peak in late June, showing the same seasonal fluctuation in abundance as drifting seaweed. Larvae seemed to associate with drifting seaweed after aggregation to surface slick areas. Body size at the commencement and the end of the seaweed-associated period was estimated at 18 and 50 mm total length, respectively. Growth rates were low until the beginning of the association with seaweed; thereafter they became high but declined towards the end of seaweed-associated life. Larvae and juveniles born later grew faster than those born earlier. This tendency was detected both before and during the association with drifting seaweed. Although the season of abundant drifting seaweed was about 2␣months shorter, and occurred 2 months later, than the extrusion season of the gold-eye rockfish from early February to early June, this seemed to be compensated by the growth variation dependent on birth month. Early life history traits of the gold-eye rockfish are discussed in relation to the successful utilization of drifting seaweed. Received: 3 August 1997 / Accepted: 24 March 1998     

The ecology of two sandy beaches in south west India. I. Seasonal changes in physical and chemical factors,and in the macrofauna

Changes during one year in the fauna of two beaches in south India are described in relation to the nutrient content of the water and sand, and the physical changes brought about by the south-west monsoon. At both beaches, the most stable conditions were reached in the pre-monsoon months when the greatest number of species occurred and when there was marked zonation. During the monsoon, erosion took place and only actively migrating species remained to form a permanent element of the beach fauna. Data on population density and biomass indicate markedly seasonal recruitment and rapid growth of individual species, most of which are suspension feeders.     

Effects of Cd2+ exposure on key life history traits and activities of four metabolic enzymes in Helicoverpa armigera (Lepidopteran: Noctuidae)

Cadmium (Cd) is a toxic non-essential element accumulated easily in living organisms due to its non-biodegradable nature. A slight increase in Cd levels will threaten the health of terrestrial and aquatic organisms. Therefore, Cd contamination and its effects on the ecosystems and organisms have become a major topic of eco-toxicological research. The present study investigated the eco-toxicological effects of Cd-supplemented diet on some life history traits and specific enzyme activities of carboxylesterase (CarE), glutathione S-transferase (GST), acetylcholinesterase (AChE) and cytochrome P450 monooxygenases (P450s) in the cotton bollworm, Helicoverpa armigera. The results showed that Cd significantly decreased larval survival, pupation and emergence, and reduced body mass of the sixth instar larvae, pupae, adult moths and female fecundity. Cd-supplemented diet also delayed the development of larvae and pupae and shortened the life span of the cotton bollworm moth. In addition, dietary Cd inhibited the activities of GST, CarE and P450, but increased AChE activity of the sixth instar larvae. We concluded that life history traits and metabolic enzymes are sensitive to Cd stress in the cotton bollworm. These parameters are potential biomarkers for monitoring environmental stress in Lepidopteran insect pests.     

The ecology of two sandy beaches in south west India. III. Observations on the population of Donax incarnatus and D. spiculum

The biology of two species of Donax, D. incarnatus Gmelin and D. spiculum Reeve on two beaches in south west India is described. Two year groups of D. incarnatus were present on both beaches, from settlements during the monsoon period in 1967 and 1968. At Shertallai, both groups were studied through the year, and data on growth, mortality and production are presented. At Cochin, the rate of growth was slower and the maximum size attained smaller, but mortality during the early monsoon precluded study of D. incarnatus through a full year. D. spiculum occurred at Cochin mainly during the pre-monsoon period, and at Shertallai during the post-monsoon. The species has a shorter life-span than D. incarnatus, but its irregular occurrence did not allow detailed production estimates.