Depth-related shift in life history strategies of a brooding and broadcasting deep-sea asteroid

Combining field and laboratory work, this study investigated the reproductive cycle, aggregative behavior, spawning periodicity, development and early growth of the sea star Henricia lisa living at bathyal depths off eastern Canada. Marked differences were found between individuals from ~1,300 and ~600 m deep. The former had a male biased sex ratio and an aperiodic reproductive cycle, whereas the latter displayed an equal sex ratio and a biannual breeding pattern. Furthermore, the maximum size was larger and female fecundity roughly five times higher in shallower compared to deeper populations. In the tanks, aggregative behavior was recorded twice a year during the summer and winter breeding periods. The onset of aggregations and spawning coincided with a temperature of 3–4°C. Males spawned first and females typically responded inside 30–60 min. Between 12 and 20 eggs were retained to be brooded under the arched arms of the female, whereas the remainder were broadcasted and developed without parental care. The fertilized eggs underwent a first cleavage after 12 h, reached the brachiolaria stage in 1 month, became juveniles within 3–4 months and reached ~ 4 mm in diameter after 14–17 months of growth. The embryos and juveniles developed at the same rate whether brooded or not, and development of winter cohorts was typically slower due to lower prevailing temperatures. This study of H. lisa provides the first evidence of lecithotrophy in a seasonally breeding deep-sea echinoderm and of brooding in a deep-sea asteroid.     

Reproductive pattern in the brooding and broadcasting sea star Pteraster militaris

Pteraster militaris (O. F. Müller, 1776) broods young in an aboral chamber and broadcasts gametes and offspring as well. The reproductive cycle of a population of P. militaris from the northwest Atlantic Ocean was examined over a 23 mo period from September 1985 to July 1987. The species is gonochoric (sex ratio 1:1), with only one incidence of hermaphroditism observed (n=261). Gonad indices were lower in females than in males, possibly an adaptation to the limited space available in the brood chamber. Body component and maturity indices, gonad histology and response to 1-methyladenine indicate that females in the population have a continuous breeding season, with a slight decrease in intensity in the spring. Males have a more pronounced decrease in the intensity of reproduction in the spring. Gonad development in males in synchronous both within and between individuals. Gonad development in females is asynchronous.     

Reproductive cycle of the subarctic brooding asteroid Leptasterias polaris

The reproductive cycle of the large brooding seastar Leptasterias polaris Müller and Troschel was examined over an 18-month period in the St. Lawrence Estuary, Québec, Canada. There is a distinct annual cycle with spawning over several months in the autumn. The female has an unusual cycle in that the ovary only slightly decreases in size during spawning, and the size structure of the oocyte population is remarkably stable throughout the year. The major annual change observed in the oocyte population is the development of a small number of 600-to 800- oocytes prior to spawning and their loss during spawning. This stability, combined with the lack of evidence of phagocytosis, suggests that oocyte development takes place over many years. By contrast, the developmental cycle of the testis is similar to that of most echinoderms. The large reservoir of oocytes probably guarantees a steady annual recruitment, and brooding during the winter probably minimizes metabolic costs for the female and ensures the release of the juveniles when conditions are favourable in the spring and summer.     

Conflicts,social economics and life history strategies in ants

Summary This paper presents a life history model for a perennial social insect colony. The model's purpose is to explore the evolutionary consequences, in terms of fitness of different parties within the colony, of alternative life history strategies. The model has been specifically developed for colonies of the slave-making ant, Harpagoxenus sublaevis, which has reproductive workers organized in dominance orders. It incorporates empirically obtained parameters, and uses computer algorithms based on numerical optimization to determine the optimum policy for a colony queen in allocating resources between workers, queens, and males. Variants of the model also consider alternative situations in which either (1) orphaned workers do not slave-raid, or (2) workers are sterile. The results correspond closely to data on colony growth and reproductive allocation obtained from the field. They suggest that a colony queen would suffer reduced fitness in the two theoretical cases as compared to the real situation. Reproduction by orphaned workers posthumously enhances the colony queen's fitness because a queen with sterile workers cannot produce enough extra sexuals in her lifetime to balance her loss in grandson production. The results also suggest that the division of labour between slave-raiding and nonraiding workers observed in H. sublaevis colonies can be explained as an outcome of worker-worker reproductive conflict: reproductively-inhibited subordinate workers can increase their inclusive fitness by slaveraiding for dominant, nonraiding egg-layers. These findings emphasize the evolutionary importance of the orphanage period and of intracolony conflict in monogynous social insect colonies.     

Effects of brooding and broadcasting reproductive modes on the population genetic structure of two Antarctic gastropod molluscs

Life-history characteristics exert a profound influence upon the population structure of many marine organisms. However, relatively few genetic studies have compared direct with indirect-developing species in the same ecosystem or geographical region, and none to our knowledge within an Antarctic setting. To address this issue, we have collected novel amplified fragment length polymorphism (AFLP) data from the direct-developing top shell Margarella antarctica to form a comparison with previously published data for the broadcast-spawning Antarctic limpet Nacella concinna. We scored 270 loci in 240 M. antarctica individuals sampled from five populations spanning the full length of the Antarctic Peninsula. Profound differences were identified in the strength and pattern of population structure between the two species, consistent with gene flow being highly restricted in M. antarctica relative to N. concinna.     

Implications of dispersal and life history strategies for the persistence of Linyphiid spider populations

Linyphiid spiders have evolved the ability to disperse long distances by a process known as ballooning. It has been hypothesized that ballooning may allow populations to persist in the highly disturbed agricultural areas that the spiders prefer. In this study, I develop a stochastic population model to explore how the propensity for this type of long distance dispersal influences long term population persistence in a heterogeneous landscape where catastrophic mortality events are common. Analysis of this model indicates that although some dispersal does indeed decrease the probability of extinction of the population, the frequency of dispersal is only important in certain extremes. Instead, both the mean population birth and death rates, and the landscape composition, are much more important in determining the probability of extinction than the dispersal process. Thus, in order to develop effective conservation strategies for these spiders, better understanding of life history processes should be prioritized over an understanding of dispersal strategies.     

Colony life history and demography of a swarm-founding social wasp

Colonies of social insects are sometimes viewed as superorganisms. The birth, reproduction, and death of colonies can be studied with demographic measures analogous to those normally applied to individuals, but two additional questions arise. First, how do adaptive colony demographies arise from individual behaviors? Second, since these superorganisms are made up of genetically distinct individuals, do conflicts within the colony sometimes modify and upset optima for colonies? The interplay between individual and superindividual or colony interests appears to be particularly complex in neotropical, swarm-founding, epiponine wasps such as Parachartergus colobopterus. In a long-term study of this species, we censused 286 nests to study colony-level reproduction and survivorship and evaluated individual-level factors by assessing genetic relatedness and queen production. Colony survivorship followed a negative exponential curve very closely, indicating type II survivorship. This pattern is defined by constant mortality across ages and is more characteristic of birds and other vertebrates than of insects. Individual colonies are long-lived, lasting an average of 347 days, with a maximum of over 4.5 years. The low and constant levels of colony mortality arise in part from colony initiation by swarming, nesting on protected substrates, and an unusual expandable nest structure. The ability to requeen rapidly was also important; relatedness data suggest that colonies requeen on average once every 9–12 months. We studied whether colony optima with respect to the timing of reproduction could be upset by individual worker interests. In this species, colonies are normally polygynous but new queens are produced only after a colony reaches the monogynous state, a result which is in accord with the genetic interests of workers. Therefore colony worker interests might drive colonies to reproduce whenever queen number happens to cycled down to one rather than at the season that is otherwise optimal. However, we found reproduction to be heavily concentrated in the rainy season. The number of new colonies peaked in this season as did the percentages of males and queens. Relatedness among workers reached a seasonal low of 0.21–0.27, reflecting the higher numbers of laying queens. This seasonality was achieved in part by a modest degree of synchrony in the queen reduction cycle. Worker relatedness reached peaks of around 0.4 in the dry season, reflecting a decrease to a harmonic mean queen number of about 2.5. Thus, a significant number of colonies must be approaching monogyny entering the rainy season. Coupled with polygynous colonies rearing only males (split sex ratios), this makes it possible for a colony cycle driven by selfish worker interests to be consistent with concentrating colony reproduction during a favorable season.     

Fecundity and reproductive strategies in deep-sea incirrate octopuses (Cephalopoda: Octopoda)

Coleoid cephalopods show flexibility in their reproductive strategies or mode of spawning, which can range from simultaneous terminal spawning over a short period at the end of the animal’s life to continuous spawning over a long period of the animal’s life. Although a simultaneous terminal spawning strategy is typical of shallow water temperate octopuses, it is not known whether deep-sea octopods would have the same reproductive strategy. The reproductive strategies and fecundity were investigated in nine species of deep-sea incirrate octopuses: Bathypolypus arcticus, Bathypolypus bairdii, Bathypolypus ergasticus, Bathypolypus sponsalis, Bathypolypus valdiviae, Benthoctopus levis, Benthoctopus normani, Benthoctopus sp., and Graneledone verrucosa (total n = 85). Egg-length frequency graphs and multivariate analysis (principal components analysis) suggest that B. sponsalis has a synchronous ovulation pattern and therefore a simultaneous terminal spawning strategy. Although a simultaneous terminal spawning strategy is most likely for B. levis and B. normani, the egg-length frequency graphs and multivariate analysis also suggest a greater variation in egg-lengths which could lead to spawning over an extended period.     

Nymphal density, behavioral development, and life history in a field cricket

Population density regulates the strength of intraspecific competition and may thereby be reflected in life-history variables, such as development time, growth rate, or investment in immune defense. However, population density may also affect the fitness payoffs of different behaviors and consequently shape the development of personality. Here we studied if population density during nymphal development (one, four, or ten individuals raised together) affects the level of boldness, measured as the latency time to recover from freezing and emerge from a shelter, aggressiveness towards conspecifics or their correlation at the adult stage in the field crickets, Gryllus integer. In addition, we tested if individuals invest more resources in immune function or speed up their development in response to a high conspecifics density during ontogeny. Nymphal density did not affect adult boldness or aggressiveness towards conspecific males per se, but these variables showed a negative association, i.e., indicated an unconventional behavioral syndrome in the highest density treatment. Supporting the effectiveness of density treatments in inducing plastic responses, individuals reached maturity sooner and invested more resources in immune function in the highest nymphal density group compared to groups consisting of one or four individuals. Our results suggest that population density may play an important role in shaping both the realized life history and development of behavioral syndromes.     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Life history correlates of alternative migratory strategies in American Dippers

Partial migration is thought to be a critical step in the evolution of avian migration, but data on the life history correlates of alternative migratory strategies are extremely limited. We have studied a partially migratory population of American Dippers since 1999. This population is composed of sedentary individuals (residents) that maintain the same territory year round and altitudinal migrants that share winter grounds with residents, but move to higher elevations to breed. We used seven years of data on individually marked birds to (1) determine if individuals consistently use the same migratory strategy, (2) determine if offspring have the same strategy as their parents, and (3) estimate reproductive and survival rates of the two migratory strategies. We evaluate hypotheses for the persistence of partial migration and discuss their implication for the evolution of migration in sedentary populations. Individual American Dippers rarely switched migratory strategy (4/169 monitored more than one year). An individual's strategy, however, was not always that of its parents, indicating that, while migratory behavior may have a genetic component, environmental or social conditions probably influence the migratory strategy that an individual adopts. Sedentary dippers consistently had higher annual productivity (approximately 1.4 more fledglings/year) than migratory dippers, but mark-recapture models suggested that migratory dippers may have slightly higher survival than residents (approximately 3.4%). Migrants were estimated to have lower lifetime reproductive success than residents because their higher survival was insufficient to offset their lower productivity. Our data suggest that alternative migratory strategies in American Dippers are unlikely to be a fixed genetic dimorphism that persists because the two strategies have equal fitness, or because the relative fitness of the two strategies fluctuates over time. Migratory strategies in American Dippers are more likely to be condition dependent, and the two strategies persist because migrants "make the best of a bad job" by moving to higher elevations to breed. Because migrants obtained no fitness benefits by moving to seasonal breeding territories, our data are consistent with the hypothesis that migration could evolve in sedentary populations if competition for limited resources forces some individuals to seek breeding opportunities outside their initial range.     

The life history of Rhodophysema georgii in laboratory culture

Rhodophysema georgii Batt., a rare epiphyte on Zostera spp., was collected from lowtide-level lagoons at Bembridge (Isle of Wight), and its life history examined in laboratory culture. The tetraspores, the only method of reproduction known, gave rise to small crusts which again developed tetrasporangial sori on the surface, similar to those of the parental material. It is suggested that this direct type of life history is probably due to the repression of meiosis in the tetrasporangial mother cell.     

Reproductive strategies and diet in deep-sea nuculanid protobranchs (Bivalvia: Nuculoidea) from the Rockall Trough

Although the taxonomy of deep-sea protobranch bivalves is becoming better known, relatively little information is available on their reproductive biology and whether or not populations show reproductive periodicities. We have examined the reproductive biology of three common sympatric species as part of a long-term time-series of samples taken from 2900 m in the Rockall Trough from 1973 to 1983. Malletia cuneata Jeffreys, 1876 produces a maximum of 30 oocytes at any one time and these grow to a maximum size of 240 m. Maximum fecundity of Ledella pustulosa (Jeffreys, 1876) and Yoldiella jeffreysi (Hidalgo, 1877) is 174 and 360, respectively, and both species produce an egg of 120 m in diameter. These data indicate lecithotrophic early development in L. pustulosa and Y. jeffreysi, but direct development in M. cuneata; however, evidence from the prodissoconch length of M. cuneata suggests lecithotrophic development. L. pustulosa and Y. jeffreysi also differ from M. cuneata in having a distinct reproductive cycle with spawnout in the winter months. Both the continuously breeding M. cuneata and the seasonally breeding L. pustulosa ingest diatoms, coccoliths and foraminiferans, but whereas the diet of M. cuneata appears to be constant throughout the year there is an apparent reduction in the feeding activity of L. pustulosa concomitant with the deposition of phytodetritus on the deep-sea bed.     

Linking life history to landscape for threatened species conservation in a multiuse region

Landscape-scale conservation that considers metapopulation dynamics will be essential for preventing declines of species facing multiple threats to their survival. Toward this end, we developed a novel approach that combines occurrence records, spatial–environmental data, and genetic information to model habitat, connectivity, and patterns of genetic structure and link spatial attributes to underlying ecological mechanisms. Using the threatened northern quoll (Dasyurus hallucatus) as a case study, we applied this approach to address the need for conservation decision-making tools that promote resilient metapopulations of this threatened species in the Pilbara, Western Australia, a multiuse landscape that is a hotspot for biodiversity and mining. Habitat and connectivity were predicted by different landscape characteristics. Whereas habitat suitability was overwhelmingly driven by terrain ruggedness, dispersal was facilitated by proximity to watercourses. Although there is limited evidence for major physical barriers in the Pilbara, areas with high silt and clay content (i.e., alluvial and hardpan plains) showed high resistance to dispersal. Climate subtlety shaped distributions and patterns of genetic turnover, suggesting the potential for local adaptation. By understanding these spatial–environmental associations and linking them to life-history and metapopulation dynamics, we highlight opportunities to provide targeted species management. To support this, we have created habitat, connectivity, and genetic uniqueness maps for conservation decision-making in the region. These tools have the potential to provide a more holistic approach to conservation in multiuse landscapes globally.     

A model for a species with two life history stages and added mortality

A differential equation model of a species with two life stages is constructed, and the behavior of the equilibria investigated when density-independent mortality is continuously imposed. With density dependence largely in recruitment, an increase in either life stage is possible even though mortality is being imposed on both life stages. The equilibrium of a given life stage can increase if density dependence is mainly in that stage.The most effective stage to attack is that which is numerically dominant. With species having a high reproductive potential the most effective way of reducing either stage is to attack the stage with the lesser natural density-independent mortality.Certain parameter values allow multiple positive steady states. Data from Lucilia cuprina appear to conform to some of the predictions of the model.     

Selective harvesting and habitat loss produce long-term life history changes in a mouflon population.

We examined the long-term effects (28 years) of habitat loss and phenotype-based selective harvest on body mass, horn size, and horn shape of mouflon (Ovis gmelini musimon) in southern France. This population has experienced habitat deterioration (loss of 50.8% of open area) since its introduction in 1956 and unrestricted selective hunting of the largest horned males since 1973. Both processes are predicted to lead to a decrease in phenotype quality by decreasing habitat quality and by reducing the reproductive contribution of individuals carrying traits that are targeted by hunters. Body mass and body size of both sexes and horn measurements of males markedly decreased (by 3.4-38.3%) in all age classes from the 1970s. Lamb body mass varied in relation to the spatiotemporal variation of habitat closure within the hunting-free reserve, suggesting that habitat closure explains part of these changes. However, the fact that there was no significant spatial variation in body mass in the early part of the study, when a decline in phenotypic quality already had occurred, provided support for the influence of selective harvesting. We also found that the allometric relationship between horn breadth and horn length changed over the study period. For a given horn length, horn breadth was lower during the second part of the study. This result, as well as changes in horn curve diameter, supports the interpretation that selective harvesting of males based on their horn configuration had evolutionary consequences for horn shape, since this phenotypic trait is less likely to be affected by changes in habitat characteristics. Moreover, males required more time (approximately four years) to develop a desirable trophy, suggesting that trophy hunting favors the reproductive contribution of animals with slow-growing horns. Managers should exploit hunters' desire for trophy males to finance management strategies which ensure a balance between the population and its environment. However, for long-term sustainable exploitation, harvest strategy should also ensure that selectively targeted males are allowed to contribute genetically to the next generations.     

Depth-related enzymic activities in muscle,brain and heart of deep-living pelagic marine teleosts

The activities [units^-1 wet weight tissue] of lactate dehydrogenase (LDH), pyruvate kinase (PK), malate dehydrogenase (MDH) and isocitrate dehydrogenase (IDH) in white skeletal muscle, brain and heart of 24 pelagic teleost fishes were determined. In addition, for several of the foregoing species, citrate synthase (CS) activities were examined in white skeletal muscle. In muscle, the activities of all these enzymes decrease exponentially with increasing minimum depth of occurrence of the species; this decrease closely parallels the decrease in respiratory rate found previously for these same species. The decline in enzymic activity with increasing minimum depth of occurrence is much greater than the decline in body protein content of the whole fishes, suggesting a disproportionately rapid fall in enzyme concentration in comparison to contractile and structural protein concentrations. The similar reductions in activities of both glycolytic (LDH and PK) and citric acid cycle (CS, MDH and IDH) enzymes with depth indicate that both standard and active metabolisms of deeper-dwelling species are reduced relative to shallower-dwelling forms. There is no suggestion of increased anaerobic capacity with depth or in relation to species, occurrence in the oxygen minimum layer. In brain and heart, there is no significant decrease in enzymic activity with increasing minimum depth of occurrence. These two tissues have similar capacities for metabolism in most fishes, when comparisons are based on enzymic activity per gram wet weight of tissue.     

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.     

Life history of Gonatus onyx (Cephalopoda: Teuthoidea): deep-sea spawning and post-spawning egg care

 A reproductive strategy consisting of deep- water spawning and egg-care was inferred for the midwater squid Gonatus onyx Young, 1972. Brooding females and associated eggs and hatchlings, captured between 1250 and 1750 m depth off southern California, are described. Brooding females appear to be senescent and lack tentacles. Large eggs of this species (3 mm) at cold temperatures (3 °C at capture depth) may require as long as 9 mo to develop. The high lipid content of the digestive gland in adult females of this species may provide fuel to support such an extended “brooding” period. Received: 22 February 1999 / Accepted: 25 May 2000