Population dynamics, sex ratio and size at sex change in a protandric simultaneous hermaphrodite, the spiny shrimp Exhippolysmata oplophoroides

One of the main goals of sex allocation theory is understanding sex ratio evolution. However, theoretical studies predicting sex ratios in species with unusual sexual systems, such as protandric simultaneous (PS) hermaphroditism, are rare. In PS hermaphrodites, juveniles first develop into functional males that mature into simultaneous hermaphrodites later in life. Here, we report on the sex ratio (males/males + hermaphrodites) in the PS hermaphroditic shrimp Exhippolysmata oplophoroides. A 2-year study demonstrated that hermaphrodites dominated the population in two different bays. This skewed sex ratio may be explained by limited encounter rates among conspecifics. In agreement with this idea, the density of shrimps was extremely low (≤1 shrimp km⁻²) at the two study sites. Size at sex phase change and sex ratios remained relatively stable through time at the two bays. The stability of these parameters might be explained by the rather steady population structure of this species during the study period. A review of sex ratios in PS hermaphroditic shrimps (Lysmata and Exhippolysmata) revealed considerable variation; some species have male- and others hermaphrodite-skewed sex ratios. The conditions explaining inter- and intra-specific sex ratio variation in protandric simultaneous hermaphroditic species remain to be addressed.     

Reproduction of a Northumberland population of the polychaete Cirratulus cirratus

A population of Cirratulus cirratus (O. F. Müller) was sampled from September 1966 to September 1968. The population is found intertidally in crevices at Cullercoats Bay, Northumberland (UK). Measurements of the sexual maturity and size of the animals have been made. The breeding season is not restricted, spawning occurs in the population asynchronously throughout the year. Ripe worms and worms that have just spawned are invariably present, but the majority of females are always at intermediate stages of oogenesis. Adult males, however, always contain active spermatozoa. Females, but not males, are devoid of coelomic gametes after spawning; a new generation of coelomic oocytes begin to accumulate in the coelom after 30 days. The growth of the coelomic oocytes is relatively slow, the coelomic phase of oocyte development takes between 1 and 2 years for completion. The coelomoducts which serve for the emission of the gametes are described. An interpretation of the female population structure based upon length and sexual maturity data suggests that most females survive to spawn 2 or 3 times, and some to spawn 4 or 5 times. They grow in length by about 2 cm in the period between successive spawnings. The reproduction of Cirratulus cirratus is compared with that of other annelid species breeding throughout the year, and this unusual mode of reproduction is discussed.     

Reproduction and development of the polychaete Stauronereis rudolphi,including a summary of development in the superfamily Eunicea

The life history of the dorvilleid polychaete Stauronereis rudolphi (Delle Chiaje) was studied through the F₂ generation under laboratory conditions. The species released its gametes into the water where fertilization occurred. The pelagic larvae swam for one week before settling. Segments were added successively and sexual maturity was reached in one year of laboratory culture. Spawned adults recovered and began feeding a few days after spawning. The culture techniques and foods used to rear developing eggs, larvae, juveniles, and adults were described. Comparisons were made with other developmental studies in the superfamily Eunicea.This study was a portion of a thesis submitted in partial fulfillment of the requirements for the degree of Master of Arts in the Department of Biology, California State College at Long Beach. It was in part supported by research grant WP-00531-02 from the National Institute of Health, U. S. Public Health Service, under supervision of Dr. Donald J. Reish. Contribution No. 1 of the Ira C. Darling Center for Research, Teaching and Service, University of Maine, Maine 04573, U.S.A.     

Reproduction and development of Sabella media,a sabellid polychaete with extratubular brooding

Known reproductive patterns in the polychaete family Sabellidae include: (1) broadcasting of gametes, (2) depositing of benthic egg masses, (3) brooding outside the lip of the tube, and (4) brooding within the tube. There is little information for sabellids in the third category; there-fore we have studied the reproductive biology of Sabella (=Demonax) media (Bush), one of the few species known to brood its early developmental stages outside the tube. A total of 32 adults of S. media with egg cocoons were collected from the surfaces of compound ascidians (Aplidium sp. and Didemnum sp.) and from holdfasts of a brown alga (Hedophyllum sessile) at Eagle Cove, San Juan Island, Washington, USA, from June 1981 through June 1982, between tidal levels of 0.0 and 0.6 m. The donutshaped cocoons, each containing over 1 000 eggs, are deposited just outside the tube aperture from April through September. Larval development takes place within the cocoon through the 3-setiger stage, at which time the larvae emerge (about 8 d after egg deposition). The newly hatched larvae are demersal and swim near the bottom of the culture dish for 1 d before settlement. Metamorphosis takes nearly a week, and initial formation of the tube is also gradual. The reproductive patterns within the family Sabellidae are discussed, and the adaptive significance of extratubular brooding considered.     

Experimental test of socially mediated sex change in a protandric simultaneous hermaphrodite,the marine shrimp<Emphasis Type="Italic"> Lysmata wurdemanni</Emphasis> (Caridea: Hippolytidae)

In Lysmata wurdemanni, individuals begin benthic life in a male phase (MP) but later change to a female phase (FP) with female external morphology, but with both male and female reproductive capacity (protandric simultaneous hermaphroditism). Previous studies have demonstrated that the size (timing) of sex change varies considerably in natural populations. We experimentally tested for social mediation of sex change by rearing male-phase individuals (MPs) in both large and small social groups with different sexual and size composition. In the large group experiment, speed of sex change was inversely related to the abundance of female-phase individuals (FPs) in the group (sex-ratio induction). Increased allocation to female function (more rapid change to FP) may occur when male mating opportunities are lower because the simultaneous-hermaphrodite FP can immediately reproduce as a female while maintaining male mating capacity. When FPs are abundant, delayed sex change might be adaptive because the costs of female reproduction are considerable. An MP may gain reproductively by increased growth before changing to FP at a larger size (fewer but much larger broods). Size-ratio induction of sex change by small MPs was suggested but not confirmed. Experimental results from small groups (1–2 individuals) were qualitatively similar but not as conclusive as those from large groups. The number and complexity of social interactions in large groups may be necessary to stimulate labile sex change in this species. In L. wurdemanni, sex change may be influenced not only by abiotic factors related to breeding [Bauer (2002) Biol Bull 203:347–357] but also by social factors in certain demographic situations.Communicated by K. Lindström     

Population-level variation in plant secondary chemistry,and the population biology of herbivores

Summary Modern population ecology is becoming increasingly spatially-explicit. For insect hervibores, spatial variation in plant quality is a critical component of distribution and abundance. We argue that populationlevel measurements of phytochemical variation among individual plants has not kept pace with recent developments in population ecology. With examples from our own work, we demonstrate the importance of spatial variation in plant chemistry for insect herbivores, and suggest that phytochemistry should play a central role in the development of any spatially-based ecological theory.     

Behaviour that influences dispersal and connectivity in the small,young larvae of a reef fish

Determining the scale of larval dispersal and population connectivity in demersal fishes is a major challenge in marine ecology. Historically, considerations of larval dispersal have ignored the possible contributions of larval behaviour, but we show here that even young, small larvae have swimming, orientation and vertical positioning capabilities that can strongly influence dispersal outcomes. Using young (11–15 days), relatively poorly developed (8–10 mm), larvae of the pomacentrid damselfish, Amblyglyphidodon curacao (identified using mitochondrial DNA), we studied behaviour relevant to dispersal in the laboratory and sea on windward and leeward sides of Lizard Island, Great Barrier Reef. Behaviour varied little with size over the narrow size range examined. Critical speed was 27.5 ± 1.0 cm s⁻¹ (30.9 BL s⁻¹), and in situ speed was 13.6 ± 0.6 cm s⁻¹. Fastest individuals were 44.6 and 25.0 cm s⁻¹, for critical and in situ speeds, respectively. In situ speed was about 50% of critical speed and equalled mean current speed. Unfed larvae swam 172 ± 29 h at 8–10 cm s⁻¹ (52.0 ± 8.6 km), and lost 25% wet weight over that time. Vertical distribution differed between locations: modal depth was 2.5–5.0 and 10.0–12.5 m at leeward and windward sites, respectively. Over 80% of 71 larvae observed in situ had directional swimming trajectories. Larvae avoided NW bearings, with an overall mean SE swimming direction, regardless of the direction to nearest settlement habitat. Larvae made smaller changes between sequential bearings of swimming direction when swimming SE than in other directions, making it more likely they would continue to swim SE. When swimming NW, 62% of turns were left (more than in other directions), which would quickly result in swimming direction changing away from NW. This demonstrates the larvae knew the direction in which they were swimming and provides insight into how they achieved SE swimming direction. Although the cues used for orientation are unclear, some possibilities seemingly can be eliminated. Thus, A. curacao larvae near Lizard Island, on average swam into the average current at a speed equivalent to it, could do this for many hours, and chose different depths in different locations. These behaviours will strongly influence dispersal, and are similar to behaviour of other settlement-stage pomacentrid larvae that are older and larger.     

Reproduction and development of Paedoclione doliiformis,and a comparison with Clione limacina (Opisthobranchia: Gymnosomata)

Observations on breeding season, copulation, spawning, and development are described for Paedoclione doliiformis, the only gymnosomatous pteropod which consistently retains external larval features throughout its life. The number of eggs spawned per individual is dependent upon temperature and availability of Spiratella retroversa, its thecosome prey. At 17° to 19°C, planktotrophic veligers hatch from floating egg masses within 3 days, and cast their shells approximately 11 days later. The resultant shell-less veligers could complete metamorphosis to the polytrochous larval stage, in which the body is encircled by 3 ciliary bands, within 12 h. Some polytrochous larvae begin feeding on veligers of S. retroversa within 2 days. Unlike other gymnosome species, there is no further change in external appearance except for growth. A comparison of reproduction and development has been made with Clione limacina, another gymnosome which competes for food with P. doliiformis in Nova Scotian (Canada) coastal waters.     

Effects of density and simulated recruitment and mortality on sex change in a protandric simultaneous hermaphroditic shrimp, Lysmata wurdemanni

Sex change in many hermaphrodite animals has been suggested to be environmentally determined, especially socially. To investigate whether sex change in the protandric simultaneous hermaphrodite shrimp Lysmata wurdemanni (Gibbes 1850) is socially mediated, two experiments were conducted in the laboratory between September 2002 and April 2004 using laboratory-cultured shrimp that originated from Port Aransas, Texas, USA. The size at sex change (from male to simultaneous hermaphrodite) in this shrimp is variable, with the minimum around 2.4 cm in total length (TL). Large shrimp (2.4–4.5 cm TL) still in the male-phase (MP) have been found in the wild and laboratory environments. This study tested the hypothesis that large MP shrimp delay changing to the euhermaphrodite-phase (EP) due to social control. In the first experiment, ten shrimp were raised in large (110-l) and small (20-l) containers to test the effect of habitat size/density on sex change. The percentage of shrimp changing to EP was significantly higher in the large container (low density) than in the small container after 60 and 120 days. But after 570 days sex ratios were the same, 2 MP:8 EP. In the second experiment, group composition was changed over time to simulate population recruitment and mortality. MP shrimp delayed sex change when EP shrimp were present. However, if group structure is stable, some MP shrimp may not change sex during their lifetime. Under certain demographic conditions, such as when postlarvae (PL) were added to (simulating recruitment) or EP shrimp were removed from (simulating mortality) a group, all old MP (from original PL) shrimp changed to EP. The response of old MP shrimp to simulated recruitment is faster than to simulated mortality. The present study confirms that social control affects the size and timing of sex change in L. wurdemanni. However, some MP shrimp never change sex suggesting that genetics might also play a role in the sex ratios of L. wurdemanni populations.     

Reproduction,larval development and post-larval growth ofNaineris laevigata (Polychaeta,Orbiniidae) in the Mediterranean Sea

To investigate the life-history adaptations of a brackish-water population ofNaineris laevigata Grube, a population from Acquatina lagoon near Lecce (Italy) was studied. Benthos and plankton samples were collected monthly over a period of ca. 2 yr (1989–1990). Observations of gametogenesis indicate that oogenesis is intraovaric, probably with autosynthetic yolk formation. Oogenesis takes about 7 mo, from November to April–May, for both proliferative and vitellogenetic phases. Reproduction is synchronized within the population. Spawning has also been obtained under laboratory conditions where larval development and post-larval growth were studied. Eggs measured 240 µm in diameter and were spawned within a gelatinous mass. The lecithotrophic pelagic stage was very short. Reproductive effort was high, probably because of high food availability and low level of competition. One of the most interesting findings was the probable presence of a resting stage in the life-cycle. The adaptive significance of enclosing the eggs in gelatiuous masses and of reducing the duration of the pelagic stage is discussed.     

Beijing's coordinated development of population,resources, environment,and economy

Since the Olympic Games will be held in Beijing at 2008, the urban development plan of Beijing deserves special attention. The essence of regional sustainable development is that population (P), resources (R), environment (E) and economy (E) could develop with sustenance, order and coordination during a certain period, which forms a complex system called the PREE complex system. For this system, a multi-objective integrated model is formulated and solved by a nonlinear goal programming technique. Beijing's PREE complex system is then used as an empirical study. Twelve interesting goals are carefully selected, the parameters of each goal function were estimated by using Beijing's 50-year (1949–1999) statistical data, and their ideal goal limits for 2000 and 2005 are chosen according to the 9th and 10th Five-Year Economic Development Plans of Beijing. The results showed that population size is the most sensitive element in Beijing's PREE complex system. Furthermore, some suggestions related to public policies about the urban development of Beijing are proposed.     

Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata

Dipolydora armata (Langerhans, 1880) is a small (4 to 5 mm) spionid polychaete found burrowing in the calcareous hydrozoan Millepora complanata Lamarck, 1816, on coral reefs at Barbados, West Indies. It excavates complex networks of interconnecting burrows and forms aggregations of worms in cavities within branches of the coral. Adult worms have a mixed feeding mode (suspension feeding and deposit feeding). Size–frequency distributions of worms in branch samples suggest that they mature in a single year and that reproduction occurs throughout the year. Burrow openings on the surface of the coral develop distinctive, erect spines caused by combined growth of worm tubes and host tissue. Millepore zooids were absent in the vicinity of tube openings and on spines, and thus the potential feeding surface of the coral will be reduced in heavily colonized branches. Burrows and openings were densest at the bases of millepore branches where weakening of the skeleton would be expected to occur. The absence of openings near the branch tips suggests difficulty in larval settlement there, amongst stinging zooids. Reproduction␣and larval development of the worms were examined, and a sequence of larval stages from one to 20 segments and a juvenile stage of 22 segments are described. Eggs are deposited in brood sacs attached to the burrow wall, and the larvae feed upon nurse eggs (adelphophagy). The presence of larvae and juveniles occurring free in the burrows suggests that larval development may be completed within the host coral as an alternative or in addition to a planktonic larval phase. Lack of provisional larval setae, early development of adult capillary setae, production of special spermatophores and a protracted breeding cycle in D. armata are all traits which would favour complete development within the host skeleton. Received: 6 March 1997 / Accepted: 25 October 1997     

Reproduction and development of the blue coralHeliopora coerulea (Alcyonaria: Coenothecalia)

Up to the present time, the reproductive cycle and larval development ofHeliopora coerulea, living fossil and sole member of the alcyonarian order Coenothecalia, have been undescribed. Populations ofH. coerulea studied on the Great Barrier Reef between 1984 and 1989 were found to have separate sexes and an annual gametogenic cycle. Mature eggs were large (>800µm diam), and their development was initiated internally. There appeared to be a lunar or semi-lunar periodicity in reproductive behaviour, with brooding of larvae commencing around either full or new moon. Development proceeded via superficial cleavage and a solid stereoblastula. Subsequent development took place on the surface of the colony, within a temporary brood-space created by protracted distension of the polyps. Larvae lacked zooxanthellae and were brooded for between 6 and 14 d prior to leaving the colony. Reproduction and development reflected the taxonomic affinities ofH. coerulea, but brooding and embryological development also appeared to be constrained by its morphology. Reproductive and larval characteristics ofH. coerulea may offer a partial explanation of the current biogeographic range ofH. coerulea.     

The production of sexual and asexual larvae within single broods of the scleractinian coral, Pocillopora damicornis

The genotypic compositions of two populations of the brooding coral, Pocillopora damicornis, were studied by exhaustive sampling of adult colonies on a fine scale in southern Taiwan. In addition, 100 larvae were randomly selected among more than 1,000 brooded larvae collected from a single broodparent within each population. Using 7 polymorphic microsatellite markers, both populations were found to be highly clonal, and 7 clonal lineages were characterized. One clonal lineage (C1) dominated both study areas and comprises 54.9% of all colonies sampled, while any of the other 6 clonal lineages represented no more than 5%. Among the 100 larvae randomly selected for genotyping from each broodparent, the extent of clonal reproduction was high, and only 29 and 6 larvae, respectively, were found to be genotypically different from their broodparent. Among the 35 genotypically distinct larvae, 33 were thought to be derived from sexual reproduction, and 2 were assumed to be clonal propagules that had undergone somatic mutations. Two genotypically identical larvae were also found in one of the 2 sexually derived larva arrays, indicating the possible existence of polyembryony. The high proportions of clonality in both adult colonies and brooded larvae suggest that asexually produced larvae might significantly contribute to the recruitment of local populations. The dense clonal population of P. damicornis in the study area favors the quick recolonization view of clonal propagules after disturbances.     

Population biology and larval development of the terebellid polychaete Nicolea zostericola

A 3-year study was conducted on a population of Nicolea zostericola (Oersted: Grube, 1860) at Nahant, Massachusetts (USA), which indicated that this species has a life span of about 1 year. Spawing occurred from mid-April to early July, after a rise in sea-water temperature to about 6°C. Pair formation occurred between sexually mature adults prior to and during spawning. Eggs were deposited in cocoons, where development occurred in 14 to 22 days. Larval develomment is described along with juvenile and adult substrate preferences and behavior. The entire intertidal population disappears following the spawning season in late June, and reappears in September as a result of migration of worms from the sub-littoral zone. Oogenesis consists of 3 phases: a short gonadal and follicular phase, followed by a longer vitellogenic phase during which oocytes are free-floating in the coelomic fluid.Contribution no. 31 from the Marine Science Institute, Northeastern University, Nahant, Massachusetts, USA.     

Population structure and foraging biology of the predaceous chilean asteroid Meyenaster gelatinosus and the escape biology of its prey

Observations of the feeding biology of Meyenaster gelatinosus (Meyen) were made between Horcón and the southern Golfo de Penas, Chile. Of 811 sea stars examined, 436 were feeding on individuals representing 30 prey species. M. gelatinosus preys upon almost all the echinoderms and molluscs in its habitat, yet most of the prey species have extremely effective running escape behaviour in which they eventually release their attachment to the substratum, usually assuring that they will be swept to safety. Many of the molluscs exaggerate this by dorsoventral flattening of their mantles, so that they glide even farther. The echinoid Loxechinus albus has a very effective pedicillariae defense. Even at a distance, prey species usually discern foraging M. gelatinosus from non-foraging individuals, and on several occasions were observed touching M. gelatinosus which were eating conspecifics. Comparisons of the sizes of individual M. gelatinosus and their L. albus prey items showed no correlation; none of the prey species except possibly Concholepas choncholepas and M. gelatinosus itself has a refuge in size from attacking M. gelatinosus. The density of M. gelatinosus in 3 of 4 widely separated study areas where such data were collected was 0.04/m². The mean radius of M. gelatinosus ranged from 150 to 210 mm in five study areas.     

Movement patterns of young Caribbean reef sharks, Carcharhinus perezi, at Fernando de Noronha Archipelago, Brazil: the potential of marine protected areas for conservation of a nursery ground

The movement patterns and long-term site-fidelity of primarily juvenile Caribbean reef sharks, Carcharhinus perezi, were investigated using tag-recapture and automated telemetry at an insular nursery area, the Fernando de Noronha Archipelago, Brazil. Of the 143 externally tagged juvenile sharks (<110 cm), 22 (15.3%) were recaptured between 0 and 5 km from the site of tagging after 5–800 days at liberty, suggesting some site-fidelity in young individuals of this species. Site-fidelity and movement patterns of ten juvenile sharks ranging from 78 to 110 cm total length (TL) and one opportunistically captured adult female (224 cm TL) were also investigated for periods of up to 2 years with an array of automated telemetry receivers. Tagging and telemetry data from both inside and outside a marine protected area (MPA) show that shark abundance and activity is greatest along the part of the archipelago’s coastline least disturbed by human activity. Telemetry tracking also showed that juvenile reef sharks demonstrated a high degree of site-fidelity and occupied specific locations along the coast throughout the year, with some evidence of an increase in activity space with ontogeny. Sharks appeared to range more widely at night and there were no seasonal variations in habitat use. Our results suggest that MPAs may be a useful conservation tool to protect young C. perezi and potentially other reef-dwelling carcharhinid sharks during their early life history.     

Reproduction and population dynamics of the fissiparous brittle star Ophiactis savignyi (Echinodermata: Ophiuroidea)

Reproductive cycle, asexual reproduction, and population dynamics of the fissiparous brittle star, Ophiactis savignyi, which inhabits the exhalant passages of the sponge Haliclona sp. were examined monthly from February 1991 to January 1992 at Wanlitung, southern Taiwan (22°N; 120°E). Mature gametes were found from March to December, but release was mainly in May and June. Sexual recruits were found from May to December, with the highest frequency (14.1% of population) in June. Sexual recruits composed 2.4% of the 1-yr sample. Sex ratio of male to female was 24:1. Fission occurred throughout the year, although the frequency of recently split individuals was lower from January to June (6 to 31%), and higher from July to December (42 to 52%). The occurrence of fission was highest after spawning. Regenerating individuals composed 48.2% of the 1-yr sample. Population density fluctuated greatly during summer due to recruitment by fission and mortality or dispersal due to the stressful environmental conditions. Both sexual and asexual reproduction of O. savignyi were successful at this site.     

Gonadal development and infrequent sex change in a population of the humbug damselfish,Dascyllus aruanus,in continuous coral-cover habitat

The expression of protogyny often differs among populations of a single species. The humbug damselfish, Dascyllus aruanus (Linnaeus), typically lives in spatially discrete groups of several individuals around live coral colonies and has been reported to be protogynous. The size and sexual composition of groups of this species, however, vary with the size of the coral patch or the degree of coral cover. The present study examined gonadal development and sexual pattern in a population of D. aruanus living in continuous coral-cover habitat. Fish were collected in Tumon Bay, Guam (13°31′N; 144°47′E), during a 2-week period in July and August 1996. Reflecting their distributions, small fish (n=54) were collected at depths of 1–2 m over continuous Porites spp. patches, and large fish (n=56) were collected at depths of 2–3 m over a continuous Acropora spp. patch. Gonadal development was examined histologically. Gonads of D. aruanus first developed an ovarian lumen and primary-growth stage oocytes. From this ovarian state or from more developed ovaries with cortical-alveolus stage oocytes, some gonads developed into testes through degeneration of oocytes and development of spermatogenic tissue. Developing spermatogenic tissue only occurred in gonads having pre-cortical alveolus stage oocytes, and the size of individuals with gonads of this type overlapped mainly with the size range of individuals with gonads containing only pre-vitellogenic oocytes (i.e. immature females). This suggested that functional female-to-male sex change was rare or absent in this population of D. aruanus. Moreover, the different habitat distributions of small and large fish suggest that sex determination in immature fish occurs in the absence of adults.     

Female reproductive biology and oocyte development of the red frog crab,Ranina ranina,off Hachijojima,Izu Islands,Japan

Speciments of the red frog crab, Ranina ranina, examined in this study were collected off Hachijojima, Japan from 1987–1988 and 1990–1991. Ovigerous females occurred from May to September, and their carapace length ranged from 61.8 to 102.1 mm. Minimum size at the maturity was estimated to be 50 to 60 mm carapace length. The lobule-type ovary had a longitudinal ovarian cavity at the center. Germinal zones were located along the sides of the cavity. Oogenesis was divided into five main and ten detailed stages: proliferation (oogonium); previtellogenesis (bouquet, early chromatin nucleolus, late chromatin nucleolus); primary vitellogenesis (oil globule and yolk granule); secondary vitellogenesis (primary yolk platelet, secondary yolk platelet and prematuration); and maturation (maturation) stages. Vitellogenesis started at the oil globule stage when weakly positive periodic acid-Schiff granules appear in the cytoplasm. Based on the gonadosomatic index and histological changes in ovary, the reproductive cycle was divided into five stages: multiplication, from December to January; developing, from February to March; ripe in April and early May; spawning, from late May to August; and recovery, from August to November. Each crab spawned more than once during the spawning season.