Temporal variation of Tubularia indivisa (Cnidaria, Tubulariidae) and associated epizoites on artificial habitat communities in the North Sea

We have analyzed the composition, diversity, density and biomass of a temporal series of samples taken in a Tubularia indivisa community, which dominates a shipwreck in the North Sea waters (N 51°23′,730–E 02°29′,790, 17 nautical miles from the coast, 30 m depth). This shipwreck has structures emerging up to 8 m above the seabed. Water temperature ranged from 4.2°C in March to 20.3°C in August. Salinity showed few variations around 33.9 psu. Bottom tidal currents followed a semi-diurnal cycle and were preferentially NE oriented with 84% of them in the range 0.25–0.75 m s⁻¹. The mean value for total suspended matter was 6.2 mg l⁻¹ with large variations on a monthly scale. The species richness of samples varied from 15 in October to 42 in August with a mean value of 33 species. Diversity indices were higher during autumn and winter because of the strong dominance of a few crustacean species during the warmer months. The total density of individuals ranged from 6,500 ind m⁻² in October to 445,800 ind m⁻² in July, most of these individuals belonging to the amphipod species Jassa herdmani. The biomass of the T. indivisa community varied from 9 g AFDW m⁻² in October to 1,106 g AFDW m⁻² in July, with T. indivisa itself constituting between 59 and 82% of the total biomass. The biomass of T. indivisa was positively correlated with species richness and with the density of 23% of the species identified on this community, suggesting that T. indivisa plays an important structural role in this habitat. This was further confirmed by the number of species associated with T. indivisa which was generally superior to 55% of the sorted species. Multivariate analysis indicated strong differences between spring/summer−autumn/winter assemblages mostly but not solely due to the abundance patterns of species. These findings support the conclusion that shipwrecks in Belgian waters allow the development of assemblages dominated by a high biomass of T. indivisa which in turn provides shelter for high densities and biomass of epizoites. These assemblages will further show large monthly variations in densities and composition due to large variation in T. indivisa biomass under an apparent repetitive annual cycle. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differences in resource utilization and behaviour between coexisting <Emphasis Type="Italic">Jassa</Emphasis> species (Crustacea,Amphipoda)

Species of the genus Jassa are an important element of marine fouling communities, several species often co-occurring on a wide range of hard substrates. At Helgoland (North Sea, German Bight), the cosmopolitan J. marmorata occurs associated with the NW European species J. falcata and J. herdmani. Field and laboratory experiments revealed some ecological differences among species which may facilitate their coexistence. Test panels in the field were instantly colonized by the complete spectrum of resident Jassa species, but J. marmorata proved clearly superior to its congeners in rapidly colonizing new habitats. Further differences among species relate to microhabitat selection (differential use of peripheral and central parts of algal thalli) and the behavioural response to mechanical disturbance.     

Testing hypotheses of exponential growth and size-dependent molting rate in two copepod species

Individual copepodids from nature of the lipidstoring, monocyclic, diapausing Calanus glacialis, and from nature and laboratory culture of the non-storing, polycyclic Eurytemora herdmani (both collected new Halifax, Nova Scotia in 1987 and 1988) were reared in excess food at ca. 3 and 10 °C. Soon after molts, prosome lengths and weights [total dry wt (TW) of E. herdmani; structural wt (SW) and estimated oil-sac wt (OSW) of C. glacialis] were measured. Stage durations were close to published temperature-dependent predictions; C. glacialis (almost all females) did not enter resting stages. Growth of body length was linear and of body weight (TW or SW) was exponential, with no sexual difference in E. herdmani. There were mixed effects of sizes on stage durations of individuals: weakly positive at 3 °C (but not significant at ca. 10°C) for length and SW of C. glacialis and generally weakly negative for E. herdmani, except for TW at 10 °C. Body condition (residuals of log SW vs log length) of C. glacialis at ca. 10°C, unlike length or SW, was negatively related to stage duration. There was no relationship between length-corrected SW and OSW in C. glacialis. The various results appear to suggest that health was more important than allometric constraints on growth rates of individuals. However, copepods reared at temperatures very different from those previously experienced may show long-term adjustments of size, whereas development rates respond immediately. If so, only the weakly positive effects on stage durations of length of C. glacialis at ca. 3 °C and of TW of E. herdmani at ca. 10 °C may illustrate expectations under stable temperatures and adequate food in nature.     

Experimental analysis of the contribution of swimming and drifting to the displacement of reef fish larvae

The extent to which behaviour affects the dispersal of pelagic larvae in reef fishes has been a topic of major discussion among marine ecologists. Here, we experimentally quantified the extent to which the displacement of late-stage larvae of Abudefduf saxatilis is due to active movement (i.e. swimming) and drifting. We consider drifting as the component of larval displacement accounted for by the current. Drifting was quantified by comparing larval displacement to the displacement of passive particles in an extended flow chamber that gave larvae the free choice of swimming (i.e. swim with or against the current or not swim at all). We also determine whether drifting results from currents exceeding larval swimming capabilities or from the behavioural choice of larvae of not to swim against adverse currents. To do this, we compare the speeds of larval swimming in the extended flow chamber to those obtained in a smaller chamber in which larvae are behaviourally forced to swim due to space constraints and a retaining fence (most available data on larval swimming is based on this sort of chamber). Within the extended chamber, larvae tended to face the current and swim slower than it. This resulted in a net displacement increasingly determined by drifting. We also found that in the extended chamber, larvae swam at speeds between one and six times slower than the speeds they achieved in the “behaviourally modifying” smaller chamber. This suggests that the net displacement in the extended chamber was in part due to the behavioural choice of the larvae of not to swim. The importance of this “behavioural drifting” is discussed in terms of energy savings required for successful completion of the larval period and post-settlement survival. The idea that larvae may modulate their swimming behaviour raises caution for the use of published data regarding swimming capabilities of reef fish larvae when assessing the extent to which these fish actively affect their dispersal.     

Predation on egg capsules of <Emphasis Type="Italic">Zidona</Emphasis><Emphasis Type="Italic">dufresnei</Emphasis> (Volutidae): ecological implications

Among the diverse patterns of energy allocation to the offspring of gastropods, the presence of egg capsules to protect embryos is common. Females of the edible snail Zidona dufresnei attach egg capsules to hard substrates in shallow Argentine Patagonian waters (40°45′S, 64°56′W) during spring-summer. Embryonic development takes about 30 days at 22°C. In this study, three likely capsule predator species and the marks left by each on egg capsule walls were identified in laboratory experiments in February 2010. Abundances of predators and egg capsules with evidence of predation were assessed in the field in the summers of 2010 and 2011. Under laboratory conditions (N = 10 replicates per treatment and control), the predation rate by the chiton Chaetopleura isabellei was the highest (up to 90%), followed by the gastropod Tegula patagonica and the crab Neohelice granulata (~20% each). Nearly 60% of 41 capsules found in the field showed signs of predation. According to the marks identified in the laboratory, C. isabellei was responsible for 79% of this predation, and T. patagonica for the rest. Predation appears to be important during the encapsulated early life and could be an agent for selecting for resistant capsule walls and a relatively shorter development time.     

Geographic structure in Alaskan Pacific ocean perch (<Emphasis Type="Italic">Sebastes alutus</Emphasis>) indicates limited lifetime dispersal

Prevailing oceanographic processes, pelagic larvae, adult mobility, and large populations of many marine species often leads to the assumption of wide-ranging populations. Applying this assumption to more localized populations can lead to inappropriate conservation measures. The Pacific ocean perch (Sebastes alutus, POP) is economically and ecologically valuable, but little is known about its population structure and life history in Alaskan waters. Fourteen microsatellite loci were used to characterize geographic structure and connectivity of POP collections (1999–2005) sampled along the continental shelf break from Dixon Entrance to the Bering Sea. Despite opportunities for dispersal, there was significant, geographically related genetic structure (F _ST = 0.0123, P < 10⁻⁵). Adults appear to belong to neighborhoods at geographic scales less than 400 km, and possibly as small as 70 km, which indicates limited dispersal throughout their lives. The population structure observed has a finer geographic scale than current management, which suggests that measures for POP fisheries conservation should be revisited.     

Byssus drifting and the drifting threads of the young post-larval mussel Mytilus edulis

The long, drifting threads secreted by young post-larval mussels (Mytilus edulis L.) are simple monofilaments, distinct in form and function from the attachment byssus threads. The diameters of both thread types are in the micron or sub-micron range but, whereas the attachment threads are of restricted length and terminate in an attachment plaque, drifting threads exceed the post-larva in length by more than two orders in magnitude and are without plaques or any other structures. Transmission electron micrographs of drifting threads show no evidence of internal sub-structure. In contrast, attachment threads appear to be made up of filaments. These studies confirm that the drifting threads are highly effective in enhancing the dispersal of young mussels. The terminal sinking velocity of young drifters is typically ca. 1 mm s^-1. At this velocity the suspension range above the sea bed, assuming given values of vertical diffusivity, is estimated to be 0.5 to 5 m. Calculations of the fluid drag experienced by post-larvae in the water column show that the theoretical viscous drag force on the thread is sufficient to account for the reduced sinking rate of drifters. The calculated contribution of the thread to the total drag is approximately one order of magnitude greater than that of the post-larval body. A rapid thread-deployment strategy, shown by post-larvae which are brought into suspension, may prolong each drifting excursion and thus further enhance dispersal in turbulent marine environments.     

In situ ontogeny of behaviour in pelagic larvae of three temperate, marine, demersal fishes

The ontogeny of behaviour relevant to dispersal was studied in situ with reared pelagic larvae of three warm temperate, marine, demersal fishes: Argyrosomus japonicus (Sciaenidae), Acanthopagrus australis and Pagrus auratus (both Sparidae). Larvae of 5–14 mm SL were released in the sea, and their swimming speed, depth and direction were observed by divers. Behaviour differed among species, and to some extent, among locations. Swimming speed increased linearly at 0.4–2.0 cm s⁻¹ per mm size, depending on species. The sciaenid was slower than the sparids by 2–6 cm s⁻¹ at any size, but uniquely, it swam faster in a sheltered bay than in the ocean. Mean speeds were 4–10 body lengths s⁻¹. At settlement size, mean speed was 5–10 cm s⁻¹, and the best performing individuals swam up to twice the mean speed. In situ swimming speed was linearly correlated (R ²=0.72) with a laboratory measure of swimming speed (critical speed): the slope of the relationship was 0.32, but due to a non-zero intercept, overall, in situ speed was 25% of critical speed. Ontogenetic vertical migrations of several metres were found in all three species: the sciaenid and one sparid descended, whereas the other sparid ascended to the surface. Overall, 74–84% of individual larvae swam in a non-random way, and the frequency of directional individuals did not change ontogenetically. Indications of ontogenetic change in orientated swimming (i.e. the direction of non-random swimming) were found in all three species, with orientated swimming having developed in the sparids by about 8 mm. One sparid swam W (towards shore) when <10 mm, and changed direction towards NE (parallel to shore) when >10 mm. These results are consistent with limited in situ observations of settlement-stage wild larvae of the two sparids. In situ, larvae of these three species have swimming, depth determination and orientation behaviour sufficiently well developed to substantially influence dispersal trajectories for most of their pelagic period.     

Phylogeographic structure of false clownfish, Amphiprion ocellaris, explained by sea level changes on the Sunda shelf

The phylogenetic structure of the anemonefish Amphiprion ocellaris was established based on sequence data generated for the 5′ end of the cytochrome b (cytb) gene at 11 sites throughout Southeast Asia. Considerable genetic structuring was observed throughout the range of A. ocellaris. The region has a complex geography, with the Sunda shelf having been exposed during the lower sea levels associated with periods of glaciation. The direction of subsequent flooding of the Sunda shelf, as a direct consequence of the retreating glacial sheets, can be predicted based on the major river drainage systems in the region. Much of the phylogeographic structure, including levels of intra-site genetic variation, can be explained in terms of the “seeding” of sites on the Sunda shelf, by those on the shelf rim, as sea levels rose. We often found surface ocean currents in the region, which should influence larval dispersal, to be poorly correlated with phylogeographic structure. Several geographically close sites, which appear to be connected by surface currents, showed significant genetic stratification. We hypothesise that the phylogeographic structure of A. ocellaris is more reminiscent of the Pleistocene sea level changes than surface currents. The high contribution of sea level changes to the phylogeographic structure was also supported by several missing haplotypes in the generated spanning network. Cytb DNA sequences generated for recently recruited A. ocellaris individuals sampled from Singapore indicate that, on a local scale, there is a directional inflow of recruits, which is dependent on the monsoon seasons. The nature of dispersal and genetic structure of reef fish species found on the Sunda shelf is clearly complex, and should take into consideration past phylogeographic events. Received: 13 November 1999 / Accepted: 12 July 2000     

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     

Spatial and temporal genetic homogeneity in the Arctic surfclam (<Emphasis Type="Italic">Mactromeris polynyma</Emphasis>)

Commercially harvested marine bivalve populations show a broad range of population-genetic patterns that may be driven by planktonic larval dispersal (gene flow) or by historical (genetic drift) and ecological processes (selection). We characterized microsatellite genetic variation among populations and year classes of the commercially harvested Arctic surfclam, Mactromeris polynyma, in order to test the relative significance of gene flow and drift on three spatial scales: within commercially harvested populations in the northwest Atlantic; among Atlantic populations; and between the Atlantic and Pacific oceans. We found small nonsignificant genetic subdivision among eight populations from the northwest Atlantic (F _ST = 0.002). All of these Atlantic populations were highly significantly differentiated from a northeast Pacific population (F _ST = 0.087); all populations showed high inbreeding coefficients (F _IS = 0.432). We tested one likely source of heterozygote deficits by aging individual clams and exploring genetic variation among age classes within populations (a temporal Wahlund effect). Populations showed strikingly different patterns of age structure, but we found little differentiation among age classes. In one case, we were able to analyze genetic diversity between age classes older or younger than the advent of intensive commercial harvesting. The results generally suggest spatially broad and temporally persistent genetic homogeneity of these bivalves. We discuss the implications of the results for the biology and management of surfclam populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

The effect of micrograzers on algal community structure in a coral reef microcosm

The effect of amphipod grazing on algal community structure was studied within a 75 l refuge tank connected to a 6500 l closed-system, coral reef microcosm. When amphipods (Ampithoe ramondi) were absent or present in low numbers, a high biomass of mostly filamentous algal species resulted, including Bryopsis hypnoides, Centroceras clavulatum, Ceramium flaccidum, Derbesia vaucheriaeformis, Enteromorpha prolifera, Giffordia rallsiae, and Polysiphonia havanensis. These microalgae disappeared when amphipod density increase beyond approximately 1 individual cm^-2 of tank surface. The macroalga Hypnea spinella germinated in the system in association with amphipod tube sites. H. spinella plants remained rare until filamentous species were eliminated by amphipod grazing. Feeding trials confirmed that H. spinella was protected from grazing by its size rather than a chemical defense strategy. The H. spinella community we observed is similar to the flora described on algal ridges where physical conditions exclude fish grazing. We suggest that amphipods and similar micrograzers are responsible for the algal community structure of these ridges. Caging experiments may be subject to similar effects from increased amphipod grazing on the algae. Introduction of fish that are amphipod predators into the refuge tank caused an increase in algal species diversity but total H. spinella growth rates fell from 25 g dry wt month^-1 to less than 8 g dry wt month^-1. We describe amphipod behavior in relation to changes in population density and food supply, and we stress the potential for increasing the productivity of commercial seaweeds through maintenance of appropriate amphipod species in mariculture facilities.     

Embryogenesis and larval biology of the ahermatypic scleractinian<Emphasis Type="Italic"> Oculina varicosa</Emphasis>

The ivory tree coral Oculina varicosa (Leseur, 1820) is an ahermatypic branching scleractinian that colonizes limestone ledges at depths of 6–100 m along the Atlantic coast of Florida. This paper describes the development of embryos and larvae from shallow-water O. varicosa, collected at 6–8 m depth in July 1999 off Fort Pierce, Florida (27°32.542 N; 79°58.732 W). The effect of temperature on embryogenesis, larval survival, and larval swimming speed were examined in the laboratory. Ontogenetic changes in geotaxis and phototaxis were also investigated. Embryos developed via spiral cleavage from small (100 µm), negatively buoyant eggs. Ciliated larvae developed after 6–9 h at 25°C. Embryogenesis ceased at 10°C, was inhibited at 17°C, and progressed normally at 25°C and 30°C. Larval survival, however, was high across the full range of experimental temperatures (11–31°C), although mortality increased in the warmest treatments (26°C and 31°C). Larval swimming speed was highest at 25°C, and lower at the temperature extremes (5°C and 35°C). An ontogenetic change in geotaxis was observed; newly ciliated larvae swam to the water surface and remained there for approximately 18 h, after which they swam briefly throughout the water column, then became demersal. Early larvae showed no response to light stimulation, but at 14 and 23 days larvae appeared to exhibit negatively phototactic behavior. Although low temperatures inhibited the development of O. varicosa embryos, the larvae survived temperature extremes for extended periods of time. Ontogenetic changes in larval behavior may ensure that competent larvae are close to the benthos to facilitate settlement. Previous experiments on survival, swimming speeds, and observations on behavior of O. varicosa larvae from deep-water adults indicate that there is no difference between larvae of the deep and shallow populations.Communicated by J.P. Grassle, New Brunswick     

Herbivorous amphipods inhabit protective microhabitats within thalli of giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Many small marine herbivores utilize specific algal hosts, but the ultimate factors that shape host selection are not well understood. For example, the use of particular microhabitats within algal hosts and the functional role of these microhabitats have received little attention, especially in large algae such as kelps. We studied microhabitat use of the herbivorous amphipod Peramphithoe femorata that inhabits nest-like domiciles on the blades of giant kelp Macrocystis pyrifera. The vertical position of nest-bearing blades along the stipe of the algal thallus and the position of the nests within the lateral blades of M. pyrifera were surveyed in two kelp forests in northern-central Chile. Additionally, we conducted laboratory and field experiments to unravel the mechanisms driving the observed distributions. Peramphithoe femorata nests were predominantly built on the distal blade tips in apical sections of the stipes. Within-blade and within-stipe feeding preferences of P. femorata did not explain the amphipod distribution. Amphipods did not consistently select distal over proximal blade sections in habitat choice experiments. Mortality of tethered amphipods without nests was higher at the seafloor than at the sea surface in the field. Nests mitigated mortality of tethered amphipods, especially at the seafloor. Thus, protective microhabitats within thalli of large kelp species can substantially enhance survival of small marine herbivores. Our results suggest that differential survival from predation might be more important than food preferences in determining the microhabitat distribution of these herbivores.     

Phylogeography and Limited Genetic Connectivity in the Endangered Boring Giant Clam across the Coral Triangle

Abstract: The Coral Triangle is the global center of marine biodiversity; however, its coral reefs are critically threatened. Because of the bipartite life history of many marine species with sedentary adults and dispersive pelagic larvae, designing effective marine protected areas requires an understanding of patterns of larval dispersal and connectivity among geographically discrete populations. We used mtDNA sequence data to examine patterns of genetic connectivity in the boring giant clam (Tridacna crocea) in an effort to guide conservation efforts within the Coral Triangle. We collected an approximately 485 base pair fragment of mtDNA cytochrome c oxidase 1 (CO1) from 414 individuals at 26 sites across Indonesia. Genetic structure was strong between regions (φ_ST=0.549, p < 0.00001) with 3 strongly supported clades: one restricted to western Sumatra, another distributed across central Indonesia, and a third limited to eastern Indonesia and Papua. Even within the single largest clade, small but significant genetic structure was documented (φ_ST=0.069, p < 0.00001), which indicates limited gene flow within and among phylogeographic regions. Significant patterns of isolation by distance indicated an average dispersal distance of only 25–50 km, which is far below dispersal predictions of 406–708 km derived from estimates of passive dispersal over 10 days via surface currents. The strong regional genetic structure we found indicates potent limits to genetic and demographic connectivity for this species throughout the Coral Triangle and provides a regional context for conservation planning. The recovery of 3 distinct evolutionarily significant units within a well‐studied taxonomic group suggests that biodiversity in this region may be significantly underestimated and that Tridacna taxa may be more endangered than currently recognized.     

Population dynamics and growth of <Emphasis Type="Italic">Nassarius reticulatus </Emphasis>(Gastropoda: Nassariidae) in Rhosneigr (Anglesey,UK)

Seasonal changes in catch rate, growth and mortality of Nassarius reticulatus from an intertidal lagoon and a wave-exposed beach at Rhosneigr (Anglesey, North Wales, UK) are described. The number of N. reticulatus caught in baited traps from the lagoon was significantly higher (>125 individuals trap⁻¹) during the summer (>18°C), than at <12°C (<65 individuals trap⁻¹), and the numbers caught in the lagoon were an order of magnitude greater than on the beach, >13 individuals trap⁻¹ in July (>16°C), and <5 individuals trap⁻¹ between December and April (<9.5°C). Predictions of shell growth attained by N. reticulatus annually in the lagoon using graphical modal progression analysis (MPA) of length frequency data, were similar to the growth of marked and recaptured lagoon N. reticulatus. Predictions of shell growth using computerised length frequency distribution analysis (LFDA), however, did not reflect the growth as accurately as MPA. Modal progression analysis demonstrated that N. reticulatus from the lagoon achieved a higher asymptotic maximum shell length (L _∞) and a lower growth constant (K) than animals from the beach. Shell growth was seasonal with growth of the lagoon individuals slowing down towards the end of September and resuming in early April, about a month later than the beach individuals. Mortality of N. reticulatus was greater during the summer, and survival was lower in the lagoon than on the beach. Recruitment patterns were similar in the lagoon and on the beach, and MPA and LFDA predicted that larval N. reticulatus settled between late summer and early autumn, with juveniles (7–8.9 mm) appearing in the population the following year, between February and April. Growth of male and female N. reticulatus in the laboratory was similar and was temperature and size dependent. The different growth patterns between N. reticulatus from the two habitats, predicted using MPA, were maintained when individuals were reared under laboratory conditions for ∼6 months; N. reticulatus <21 mm from the beach grew faster than individuals from the lagoon, although N. reticulatus >21 mm from the lagoon grew faster and attained a larger length (26 mm) than individuals from the beach (24 mm). Low food availability did not affect N. reticulatus survival in the laboratory but significantly suppressed shell growth.     

Energetic costs of swarming behavior for the copepod Dioithona oculata

The cyclopoid copepod Dioithona oculata forms dense swarms within shafts of sunlight that penetrate the mangrove prop-root habitat of islands off the coast of Belize. Previous studies, based on in situ video recordings and laboratory studies, have shown that D. oculata is capable of maintaining fixed-position swarms in spite of currents of up to 2 cm s⁻¹. The purpose of this study was to examine the energetic costs of maintaining these swarms, in terms of increased metabolic costs of maintaining position in currents and in terms of reduced feeding rates in densely packed swarms during the day. Using a sealed, variable-speed flow-through chamber, the respiration rates of D. oculata were measured while swarms maintained position in different current speeds. The results indicate that active metabolism (swimming at maximum speed to maintain the swarm in a current) is approximately three times greater than routine metabolism (normal swimming speeds in the absence of currents), indicating a significant metabolic cost of maintaining swarms in the presence of currents. In addition, gut-pigment analysis indicated that feeding rates of these copepods were often reduced in swarms during the day compared to when the copepods were dispersed at night. Given the high “cost” of swarming, the adaptive value of swarming in terms of reduced predation, increased opportunities for mating, and reduced dispersal, must be substantial. Received: 4 June 1997 / Accepted: 18 September 1997     

Sociality reduces individual direct fitness in a communally breeding rodent,the colonial tuco-tuco (<Emphasis Type="Italic">Ctenomys sociabilis</Emphasis>)

In many social vertebrates, remaining in the natal group leads to at least short-term reductions in the direct fitness of philopatric animals. Among communally breeding rodents, the direct fitness costs of philopatry appear to increase as the frequency of successful natal dispersal decreases, suggesting a functional link between constraints on natal dispersal and the reproductive consequences of sociality. To explore this relationship empirically, I documented patterns of direct fitness among female colonial tuco-tucos (Ctenomys sociabilis), which are group-living subterranean rodents from southwestern Argentina. Demographic data suggest that successful natal dispersal is rare in this species, leading to the prediction that natal philopatry in C. sociabilis is associated with significant reductions in individual direct fitness. Using data obtained during 1996–2001, I compared the direct fitness of females that dispersed from their natal group and bred alone as yearlings to that of females that lived and bred in their natal group as yearlings. Philopatric yearlings reared significantly fewer young to weaning than did disperser (lone) yearlings. Although neither survival to a second breeding season nor the estimated lifetime number of pups reared to weaning differed between dispersal strategies, the annual direct fitness of group-living females was 23–40% less than expected, suggesting that philopatric animals experienced a substantial direct fitness cost by remaining in their natal group. These data yield important insights into the adaptive bases for group living in C. sociabilis and suggest that constraints on natal dispersal are an important factor favoring group living in this species.Communicated by J. Wilkinson