Behavioural syndromes, syndrome deviation and the within- and between-individual components of phenotypic correlations: when reality does not meet statistics

Evolutionary mechanisms leading to correlations across different behaviours, called behavioural syndromes, are hard to study, mostly because behavioural syndromes are group/population level phenomena. Recently (Herczeg and Garamszegi Behav Ecol Sociobiol 66:161–169, 2012), we introduced the concept of syndrome deviation that allows the study of behavioural syndromes at the individual level by focusing on the individual deviation from the hypothetical perfect group-level behavioural correlation. Subsequently, Dingemanse et al. (Behav Ecol Sociobiol 66:1543–1548, 2012) emphasized that behavioural syndromes refer to the between-individual component of phenotypic correlations, and only this component is relevant for syndrome deviation. They also recommended mixed models to decompose the between- and within-individual correlations. We agree that separating these components is important, but the proposed approach is impractical to apply for functionally different behaviours because (1) the assumption of constant within-individual correlations is unjustified and (2) different behaviours cannot be measured at the same time. Further, our simulations based on mixed models show that the statistical differentiation between the within- and between-individual components is inefficient when using realistic sample sizes. Until the separation of between- and within-individual correlations is resolved, we recommend alternative approaches for empirical behavioural syndrome research that consider the repeatability of the behaviours and the optimal balance between within- and between-individual sample sizes. Syndrome deviation calculated from phenotypic correlations of traits that are proven to be individual specific, or from the between-individual correlations if possible, is a meaningful metric to describe behavioural consistency and to explain its evolutionary significance.     

Individual boldness affects interspecific interactions in sticklebacks

Within populations of many species, individuals that are otherwise similar to one another in age, size or sex can differ markedly in behaviours such as resource use, risk taking and competitive ability. There has been much research into the implications of such variation for intraspecific interactions, yet little investigation into its role in influencing interspecific interactions outside of a predator–prey context. In this study, we investigated the role of individual-level behavioural variation in determining the outcomes of interactions between two ecologically similar fishes, the threespine and ninespine sticklebacks (Gasterosteus aculeatus and Pungitius pungitius). Experiment 1 asked whether individuals of both species were consistent in their expression of two behaviours: activity in novel surroundings and latency to attack prey. For each behaviour, focal individuals were assayed twice, 10 days apart. Performances were positively correlated between exposures, suggesting behavioural consistency within individuals, at least over this timescale. Experiment 2 revealed not only differences in habitat use described both by species-level variation, with ninespines spending more time in vegetated areas, but also by individual differences, with more active individuals of both species spending more time in open water than in vegetation. Experiment 3 revealed that when heterospecific pairs competed for prey, bolder individuals consumed a greater share, irrespective of species. These findings suggest that individual-level variation can facilitate overlap in habitat use between heterospecifics and also determine the outcomes of resource contests when they meet. We discuss how this might vary between populations as a function of prevailing selection pressures and suggest approaches for testing our predictions.     

Consistent crustaceans: the identification of stable behavioural syndromes in hermit crabs

Consistent individual variation in behaviour has been termed ‘animal personality’ and has been identified across a range of behavioural contexts including aggression, boldness in response to a threatening stimulus and exploration. When behaviours are correlated across multiple functional behavioural categories or ‘contexts’, ‘behavioural syndromes’ are said to be present. It is possible, however, that behavioural syndromes may also show consistencies. Here we investigated the presence of behavioural syndromes linking startle responses, exploration and aggression in hermit crabs and assessed their stability across two situations (low versus high predation risk). Correlation analyses detected behavioural syndromes between startle responses, a measure of ‘boldness’, and the latency to investigate a novel object, as well as the latency to attack an opponent in an aggressive context. The startle response–investigation and startle response–aggression syndromes were stable between situations, whilst there was a lack of relationship between investigation and aggression in each situation. Here we propose that these consistent individual differences in the expression of behavioural syndromes reveal the presence of animal personality, manifesting in not just one, but a suite of interacting traits.     

The influence of ration level on growth and statolith increment width of the tropical squid Sepioteuthis lessoniana (Cephalopoda: Loliginidae): an experimental approach

Juvenile squids were grown in individual 2.6-l floating enclosures and were fed either a high- or a low-ration diet of fish and the crustacean Acetes. Squids were maintained for a maximum of 44 days in two experiments. The high-ration individuals reached a significantly larger size in both experiments (27, 25.5 mm mean mantle length, ML) compared to their low-ration siblings (19 mm mean ML) in both experiments. The statolith increment widths prior to the start of the experiment were significantly wider (between 3 and 4 μm) compared to the increment widths after the start of the experiment (between 2 and 3 μm) both for the low- and the high-ration squids. High-ration squids also had significantly wider increments and larger statoliths than their low-ration siblings. Even though we detected consistent trends in daily statolith increment widths for the different feeding regimes, we could not detect variation in increment widths at a daily level of resolution (i.e. as a result of differences in day-to-day food intake at an individual level). This was probably due to the relatively consistent diet experienced by each individual. These experiments revealed that ration level influences squid growth rate, statolith size and daily statolith increment width. Received: 30 March 2000 / Accepted: 30 October 2000     

Social foraging in stingless bees: how colonies of Melipona fasciata choose among nectar sources

In an experimental set-up, a colony of the stingless bee Melipona fasciata demonstrated its ability to choose the better of two nectar sources. This colony pattern was a result of the following individual behavioural decisions: continue foraging, abandon the feeder, restart foraging and initiate foraging. Only very rarely did individuals switch from one feeder to the other. With the first combination of a rich (2.7 M) and a poor (0.8 M) feeder M. fasciata behaved differently from Apis mellifera. Recruitment occurred to both feeders and the poor feeder was not abandoned completely. When the poor feeder was set to 0.4 M, M. fasciata abandoned the poor feeder rapidly and allocated more foragers to the rich feeder. These patterns were similar to those reported for A. mellifera with the first combination of feeders. Over a sequence of 4 days, experienced bees increasingly determined the colony patterns, and the major function of communication between workers became the reactivation of experienced foragers. The foragers modulated their behaviour not only according to the profitability of the feeder, but also according to previous experience with profitability switches. Thus, experience and communication together regulated colony foraging behaviour. These findings and the results of studies with honeybees suggest that M. fasciata and honeybees use similar decision-making mechanisms and only partly different tools. Received: 21 December 1998 / Accepted: 5 January 1999     

Testing for between individual correlations of personality and physiological traits in a wild bird

Recently, integration of personality traits into a ‘pace-of-life syndrome’ (POLS) context has been advocated. To be able to understand how an individual’s behavioural, physiological and life history traits may coevolve, we need to jointly quantify these traits in order to study their covariance. Few studies have established links between personality and immunity properties of an individual. We here examined covariation of a measure of skeletal size (tarsus length), three behavioural traits (activity, handling aggression and breath rate) and two immunological traits (IgG level and haematocrit), in 592 wild caught blue tits. Many individuals (201) were tested more than once, allowing quantification of individual consistency of all traits and partition of the covariances between the traits, using a multivariate mixed model, into between individual and residual covariances. We find individual consistency of all behavioural traits, indicating that these capture aspects of blue tit adult personality and also the physiological measures are repeatable. Contrary to the POLS expectation, we find no overall significant individual level correlation structure between these traits and a factor analytical approach confirmed that between individual correlations across traits were not due to a common (POLS) factor or driven by size (tarsus length). Based on a published power study, we conclude that there is no common syndrome of individual level covariance between personality and physiological traits in wild blue tits or that the effect sizes, such a syndrome generates, are too low (r?<?0.3) to detect. Future field-based work should be designed to explore low effect sizes and strive to measure specific traits whose involvement is implicated to have large effect sizes as based on, e.g. laboratory findings.     

Bathymetric patterns of genetic variation in a deep-sea protobranch bivalve, Deminucula atacellana

The origin of the deep-sea benthic fauna is poorly understood and represents an enormous gap in our understanding of basic evolutionary phenomena. One obstacle to studying evolutionary patterns in the deep sea has been the technical difficulty of measuring genetic variation in species that are typically minute, rare, and must be recovered from extreme depths. We used molecular genetic techniques to quantify variation in the 16S rRNA mitochondrial gene within and among populations of the common protobranch bivalve Deminucula atacellana (Schenck, 1939). We analyzed 89 individuals from nine samples collected in the 1960s along a depth gradient from 1100 to 3800 m in the western North Atlantic. Genetic variability within populations is much lower than between populations, and peak haplotype numbers occur near the center of its depth distribution. Continental slope (<2500 m) and rise (>2500 m) populations were genetically distinct despite the lack of any obvious topographic or oceanographic features that would impede gene flow. These findings indicate that the deep-sea macrofauna can have strong population structure over small (134 km) spatial scales, similar to that observed in shallow-water and terrestrial organisms. This surprisingly high biodiversity at the genetic level affords the potential for adaptation and evolutionary diversification, the ultimate historical causes of high species diversity in the deep-sea benthos. Received: 24 July 1997 / Accepted: 26 January 1998     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Feeding ecology and evolutionary survival of the living coelacanth Latimeria chalumnae

One concept of evolutionary ecology holds that a living fossil is the result of past evolutionary events, and is adapted to recent selective forces only if they are similar to the selective forces in the past. We describe the present environment of the living coelacanth Latimeria chalumnae Smith, 1939 at Grande Comore, western Indian Ocean and report depth-dependent cave distribution, temperature, salinity and oxygen values which are compared to the fish's distribution and its physiological demands. We studied the activity pattern, feeding behaviour, prey abundance and hunting success to evaluate possible links between environmental conditions, feeding ecology and evolutionary success of this ancient fish. Transmitter tracking experiments indicate nocturnal activity of the piscivorous predator which hunts between approximately 200 m below the surface to 500 m depth. Fish and prey density were measured between 200 and 400 m, both increase with depth. Feeding tracks and feeding strikes of the coelacanth at various depths were simulated with the help of video and laser techniques. Along a 9447 m video transect a total of 31 potential feeding strikes occurred. Assuming 100% hunting success, medium-sized individuals would obtain 122 g and large females 299 g of prey. Estimates of metabolic rates revealed for females 3.7 ml O₂ kg⁻¹ h⁻¹ and for males 4.5 ml O₂ kg⁻¹ h⁻¹. Today coelacanths are considered to be a specialist deep-water form and to inhabit, with their ancient morphology, a contemporary environment where they compete with advanced, modern fish. Received: 5 July 1999 / Accepted: 11 November 1999     

Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996     

Group foraging, patch exploitation time and the finder's advantage

 We use a combination of the marginal value theorem (MVT) of Charnov (1976), and a group foraging model featuring information sharing to address patch residence in an environment where food occurs in discrete patches. We shall show that among equal competitors the optimal patch time for the individual that finds the food patch is shorter than that for the non-finder among equal competitors, T _E < T _N. This is the case if the patch-finder commences food harvesting in the patch earlier and manages to monopolise a fraction of the prey items (finder's advantage) before the other individuals come to take their benefit. When individuals differ in their food-searching abilities so that some of them (producers) contribute proportionally more to food-searching than others (scroungers), and differ in ability to compete for the food found, a difference emerges between producer and scrounger individuals in the optimal patch time. Within a patch we always have the finder's advantage (T _E < T _N) regardless of phenotype. Between patches a suite of optimal patch times for encountering individuals emerges depending on the performance of producers and scroungers when changing from solitary feeding to feeding in a group. The optimal patch time for individuals that are affected more severely by competition is shorter than that for individuals of the phenotype with better competitive ability. When both phenotypes are affected similarly no difference in optimal patch times emerges. Received: 13 February 1996 / Accepted after revision: 28 September 1996     

Variability in abundances of fishes associated with seagrass habitats in relation to diets of predatory fishes

The spatial, diel and tidal variability in the abundance of piscivorous fishes and their teleost prey, and the dietary composition of predatory fishes were investigated in beds of Heterozostera tasmanica within Port Phillip Bay, Australia, from September 1997 to February 1998. Predatory and prey fish assemblages were sampled from beds of H. tasmanica at three locations during each combination of diel (day and night) and tidal (high and low) cycles. Pelagic and benthic crustaceans represented >60% by abundance of the diets of all predatory fishes. Seven species, 54% of all predatory fishes, were piscivorous. These piscivores consumed individuals from seven families, 36.8% of the fish families being associated with seagrass. Western Australian salmon, Arripis truttacea (Arripidae) (n = 174) and yank flathead, Platycephalus speculator (Platycephalidae) (n = 46) were the most abundant piscivores. A. truttacea consumed larval/post-larval atherinids, gobiids and sillaginids. P. speculator consumed late-juvenile/adult atherinids, clinids and gobiids. While the abundances of piscivores varied between locations (P < 0.001) and diel periods (P = 0.028), the relative differences in piscivore abundance between sites and diel periods were not consistent between tides. The abundances of A. truttacea varied in a complex way amongst sites, diel period and tidal cycle, as shown by a three-way interaction between these factors (P = 0.026). Only during diurnal periods at St. Leonards was the abundance of A. truttacea significantly higher during high than low tides (P < 0.001). During the other diel periods at each site, the abundance of A. truttacea did not vary. P. speculator was significantly more abundant nocturnally (P = 0.017). The abundance of small (prey) fishes varied significantly amongst sites (P < 0.001). During the day, the abundance of small fishes did not vary between high and low tides (P = 0.185), but their nocturnal abundance was greater during low tide (P < 0.001). Atherinids (n = 1732) and sillaginids (n = 1623) were the most abundant families of small fishes. Atherinids were significantly more abundant nocturnally (P = 0.005) and during low tides (P = 0.029), and varied significantly amongst sites (P < 0.001). Sillaginids varied significantly only amongst sites (P < 0.001). Seagrass beds provide a foraging habitat for a diverse assemblage of predatory fishes, many of which are piscivorous. Anti-predator behaviour and amongst-location variability in abundances of piscivorous fishes may explain some of the diel and tidal, and broad-scale spatial patterns in small-fish abundances. Received: 23 July 1999 / Accepted: 18 January 2000     

Seasonal vertical distribution and population dynamics of the chaetognath Parasagitta elegans in the water column and hyperbenthic zone of Conception Bay, Newfoundland

The vertical distribution and population dynamics of the chaetognath Parasagitta elegans Verrill were determined in the water column and hyperbenthic zone of Conception Bay, Newfoundland from April 1997 to June 1998. The water column depth at the study site (47°32.2′N; 53°07.9′W) was 235 m. The temperature below the thermocline was <0 °C the year round. Chaetognath samples from the water column were collected with a Tucker Trawl. Those from the hyperbenthic zone, were collected with an epibenthic sledge. Depending upon whether the hyperbenthic zone was assumed to extend either 1 m or 10 m above bottom, the grand mean, areal abundance of chaetognaths in the hyperbenthic zone ranged from 6% to 40% of the total abundance in the water column (including the hyperbenthic zone), and the grand mean, areal biomass ranged from 25% to 77%. Large, mature individuals were collected only in the hyperbenthic zone, whereas small, immature individuals were collected primarily in the water column. According to body length and ovary maturity data, three cohorts were identified in the hyperbenthic zone during the study period. Within each cohort, the length frequency of reproductively mature individuals was bimodal, with groups of mean length 33 mm and 41 mm reproducing from May to October. The recruitment period of juvenile chaetognaths extended from July to February, coinciding with the recruitment period of copepods. The estimated individual growth rate of P. elegans was 1.0 mg C year⁻¹. The approximate generation time of the two groups of individuals with mean length at maturity of 33 mm and 41 mm was 450 and 780 days, respectively. This study demonstrates that a failure to sample the large, mature P. elegans living in the hyperbenthic zone leads to serious underestimates of the total abundance and biomass of chaetognaths and an inaccurate picture of seasonal population dynamics. Received: 8 September 1999 / Accepted: 15 September 2000     

Interannual, seasonal, local and body size variations in reproduction of the prawn Penaeus (Marsupenaeus) japonicus (Crustacea: Decapoda: Penaeidae) in the Ariake Sea and Tachibana Bay, Japan

 The reproductive biology of female Penaeus japonicus Bate was investigated in the Ariake Sea and Tachibana Bay (located outside the Ariake Sea), Japan from 1994 to 1996. Interannual, seasonal, individual female body size and spatial influences on the incidence of spawning were examined. The proportion of inseminated females, increased with increasing body size up to 170 mm body length (BL) and decreased thereafter. The minimum size at maturity was similar between years; 130 to 140 mm BL. The minimum size of individuals with developing ovaries or spermatophores differed from that of ripe females by 20 to 25 mm. Spawning occurred in the central part of the Ariake Sea and Tachibana Bay but rarely in the inner part. P. japonicus had a clear reproductive cycle. Spawning started earlier and ended later, occurring from mid-May to mid-October, in the eastern central part of and outside of the Ariake Sea compared with the western central part, where spawning occurred from mid-June to mid-September. Spawning dynamics differed across seasons, body sizes and areas but not across years. The seasonal peak in the proportion of ripe females varied with body size. In small individuals (130 to 169 mm BL), no peak was observed, whereas in large individuals (>170 mm BL) the proportion of ripe individuals peaked in June. The proportion of ripe individuals increased with increasing body size and was high outside the Ariake Sea. Factors causing the variation in spawning dynamics are discussed. Received: 30 November 1998 / Accepted: 12 October 1999     

Mitochondrial DNA analysis of the genetic relationships among populations of scad mackerel (Decapterus macarellus, D. macrosoma, and D. russelli) in South-East Asia

The genetic relationships among South-East Asian populations of the scad mackerels Decapterus macarellus, D. macrosoma and D. russelli (Pisces: Carangidae) were investigated. In 1995 and 1996, 216 fish were sampled in seven localities spanning the seas of Indonesia and were examined for restriction-site polymorphisms using ten restriction enzymes for the mitochondrial (mt) DNA control region, amplified by the polymerase chain-reaction. The inferred phylogeny of haplotypes led to the recognition of three distinct mitochondrial lineages or phylads consistent with the distinctions of current taxonomy. All 15 mtDNA haplotypes found in D. macarellus and all 9 haplotypes found in D. macrosoma were arranged as star-like clusters, suggesting recent evolutionary history. In contrast, the phylad formed by 6 haplotypes in D. russelli from the Sulawesi Sea exhibited diffuse topology, suggesting that ancestral lineages of this species have been retained to the present. Average nucleotide-divergence estimates between haplotypes of different phylads were between 0.042 and 0.135, suggesting ancient separation, in consistency with published allozyme data. High levels of haplotype diversity, but no geographical heterogeneity, was detected within D. macarellus from the Molucca Sea and the Banda Sea. Populations of D. macrosoma exhibited both significant differences between adjacent regions (Sunda Strait and Java Sea), and broadscale genetic homogeneity from the South China Sea to the Sulawesi Sea via the Java Sea and Makassar Strait. The geographic isolation of the D. macrosoma population sampled in the Sunda Strait suggests that this region constitute a sharp transition zone between the Indian Ocean and the Sunda Shelf. Near-monomorphism of haplotypes and low nucleotide diversity (d _X) were observed in the samples of D. macrosoma from the continental shelf (haplotype-diversity estimates, h, = 0.00 to 0.25 ± 0.08 and d _X = 0.000 to 0.002). This was in contrast to the comparatively high haplotype and nucleotide diversities observed in other pelagic fish species including D. macarellus (h = 0.82 ± 0.05, d _X = 0.012 to 0.015) and D. russelli (h = 0.63 ± 0.12, d _X = 0.016), and in the oceanic D. macrosoma population sampled in the Sunda Strait (h = 0.67 ± 0.31, d _X = 0.005). We hypothesise that this may be the consequence of recent and perhaps repeated bottleneck events that have affected the D. macrosoma population sampled on the continental shelf. Received: 29 September 1997 / Accepted: 3 September 1999     

Distribution and diversity of mtDNA lineages among southern right whales (Eubalaena australis) from Australia and New Zealand

Using a biopsy dart system, samples of skin tissue were collected from southern right whales (Eubalaena australis) in 1995 on two wintering grounds, southwest Australia (n = 20) and the Auckland Islands of New Zealand (n = 20); and on offshore feeding grounds at Latitudes 40 to 43°, south of Western Australia (n = 5). A variable section of the mitochondrial DNA control-region (289 nucleotides) was amplified and sequenced from these 45 individuals (21 males, 20 females and 4 of unknown sex), distinguishing a total of seven unique sequences (i.e. mtDNA haplotypes). Two haplotypes were found on both wintering grounds (including a common type representing 45% of each sample), and five types were unique to only one wintering ground. An analysis of variance adapted for molecular information revealed significant genetic differentiation between the two wintering grounds (p = 0.017). The feeding-ground sample was too small for statistical comparison with the wintering grounds, but included two haplotypes found only in the Auckland Islands as well as the common haplotype found on both wintering grounds. The nucleotide diversity and differentiation of mtDNA among the right whales was similar to that among humpback whales (Megaptera novaeangliae) from the same regions (Baker et al. 1998), but haplotype diversity was significantly reduced, perhaps as a result of more intensive hunting during the last century and continued illegal hunting during this century. Received: 16 March 1998 / Accepted: 18 December 1998     

Genetic variation and clonal structure in the scleractinian coral Pocillopora damicornis in the Ryukyu Archipelago, southern Japan

Samples of the scleractinian coral Pocillopora damicornis were collected from six sites located around four islands in the Ryukyu Archipelago, southern Japan, and subjected to allozyme electrophoresis. Seven polymorphic loci were examined for their allelic patterns. The ratio of observed to expected genotypic diversity (0.30 < G _o :G _e  < 0.64), the ratio of the observed number of genotypes to the number of individuals (0.47 < N _g :N _i  < 0.75), and deviations from Hardy–Weinberg equilibrium indicated that asexual reproduction plays a major role in the maintenance of established populations. However, populations were not completely dominated by a single or a few clones, and most clones were represented by only a few individual samples. The high frequency of typhoons in the region suggests that, in P. damicornis, fragmentation caused through occasional exposure to powerful waves is a major mode of asexual reproduction, but asexual production of planulae may also be contributing to the maintenance of populations. A significant genetic differentiation (F ST) was found between the six populations examined (0.027 < F ST < 0.092, average F ST = 0.056). The moderate gene flow is discussed according to characteristics of the larval stage of the species, and to circulation patterns in the region. Received: 7 August 1998 / Accepted: 18 May 1999     

Responses of ovaries and testes of Lytechinus variegatus (Echinodermata: Echinoidea) to dietary administration of estradiol, progesterone and testosterone

 In many vertebrates, environmental factors influence gamete differentiation and growth of the mature gonad through alteration of sex steroid production or action; however, it is unclear how gamete differentiation and gonadal growth are regulated in echinoids. The purpose of this study was to examine the influence of dietary administration of estradiol (E2), progesterone (P4), testosterone (T) and finasteride (F, a 5α-reductase inhibitor) on the ovaries and testes of mature Lytechinus variegatus (Lamarck) during gonadal growth. Echinoids were fed a formulated diet supplemented with steroids or steroids in combination with finasteride for 36 d. The effects of dietary administration of steroids on L. variegatus were both steroid- and sex-specific. The mean ovary index was 54% greater in individuals fed E2 than from individuals fed the control (C) diet (10.0 ± 1.1 vs 6.5 ± 0.7, respectively; P < 0.05). Individuals fed E2, P4, E2/P4, and P4 in combination with F had significantly smaller oocytes (P < 0.005) than individuals fed C. The volume fraction occupied by nutritive phagocytes in ovarian tubules from individuals fed E2/P4 or P4 were significantly larger than the volume fraction occupied by nutritive phagocytes from individuals fed C, indicating that E2 and 5α-reduced progestins may promote nutrient allocation to nutritive phagocytes. Although oocytes from the individuals fed T alone were significantly smaller than those fed T in combination with F, oocytes from individuals in both treatments were significantly larger compared to oocytes from individuals fed C. These data suggest that upon removal of 5α-reduced androgens, T is able to promote an increase in oocyte diameters. In contrast, the mean testis index was 56% greater in individuals fed P4 than in individuals fed C (8.9 ± 0.6 vs 5.7 ± 0.9, respectively; P < 0.05); the testis index did not increase in individuals fed P4/F, suggesting that 5α-reduced progestins stimulate testicular growth. Testes growth in the presence of 5α-reduced progestins was accomplished by significant increases in the volume fraction occupied by nutritive phagocytes and by a significant reduction in the volume fraction occupied by spermatogenic columns in testicular tubules. These data further indicate that 5α-reduced progestins (or 5α-reduced androgens) may inhibit spermatogenic column formation. In conclusion, E2 stimulated ovarian growth but inhibited oocyte growth, whereas T had no affect on ovarian growth but promoted oocyte growth in L. variegatus. We hypothesize that the E2 (or E2 metabolites) and/or 5α-reduced androgens in combination with T regulate oocyte growth in the echinoid L. variegatus. In addition, 5α-reduced progestins promoted nutrient accumulation in nutritive phagocytes within the ovaries and the testes. Furthermore, 5α-reduced progestins stimulated growth of the testes and inhibited spermatogenic column formation, suggesting that 5α-reduced progestins regulate nutrient accumulation into nutritive phagocytes and spermatogenic column formation in L. variegatus. The differences in estrogen effects between echinoids and asteroids may be related to differences in gonad morphology and, ultimately, the differences in cellular signalling pathways (paracrine vs endocrine). Received: 22 May 1999 / Accepted: 24 May 2000     

Intraspecific physiological variability of the gastropod Littorina saxatilis related to the vertical shore gradient in the White and North Seas

 Physiological responses to desiccation and temperature stress as well as behavioural responses to fast and abrupt environmental changes were investigated in high- and low-shore Littorina saxatilis (Olivi) from several populations from the White and North Seas. Variations in evaporation rates, resistance to air exposure and to acute and chronic temperature stress between animals from different shore levels were similar in White and North Sea periwinkles, consistent with the adaptive nature of these variations. High-shore snails were found to be able to conserve body water reserves better, to resist higher temperatures and to survive longer under conditions of combined temperature and desiccation stress than their low-shore counterparts. In a temperature range of 25 to 35 °C, the rate of evaporative water loss was positively correlated with temperature in low-shore snails while being largely temperature-independent in high-shore snails. Median lethal time during air exposure in L. saxatilis was negatively but not linearly related to the temperature of exposure. In a temperature range of 30 to 38 °C, the resistance to heat exposure in air was only slightly dependent on the temperature, with Q ₁₀ = 1.4 for the median lethal time; the heat resistance dropped drastically at temperatures above 38 °C, with Q ₁₀ = 593.8. This suggests different mechanisms of temperature resistance in different parts of the studied temperature range. In contrast, behavioural response to extreme salinity fluctuations was not uniform in the high- and low-shore periwinkles from the White and North Seas, which may reflect specific environmental conditions at different shore levels in the two areas studied. Observed physiological and behavioural variations are discussed from the viewpoint of different adaptive strategies employed by eulittoral and eulittoral-fringe animals within populations of a single species. Received: 13 December 1999 / Accepted: 11 April 2000     

Effect of arm regeneration on energy storage and gonad production in Ophiocoma echinata (Echinodermata: Ophiuroidea)

Two laboratory experiments were conducted on Ophiocoma echinata (Lamarck, 1816) collected from Missouri Key, Florida, USA, to ascertain the effects of arm regeneration on energy storage and gonad production. In each experiment individuals were divided into non-regenerating (all arms intact) and regenerating (three arms autotomized) groups and maintained together in a re-circulating seawater system for 2 mo. In Experiment 1, individuals were fed once a week, and in Experiment 2 once every 2 wk. The amount of food energy required for maintenance was estimated to be 0.172 kJ d⁻¹. The lower feeding level (Experiment 2) was below maintenance ration and both groups lost material relative to an initial group. At the higher feeding level (Experiment 1), regeneration of three arms resulted in less storage of organic material, mainly lipid in the stomach, and less gonad production. Thus, regeneration may reduce energy reserves and reproductive output. Received: 23 April 1998 / Accepted: 11 January 1999