Evolution of egg size and fertilisation efficiency in sea stars: large eggs are not fertilised more readily than small eggs in the genus<Emphasis Type="Italic"> Patiriella</Emphasis> (Echinodermata: Asteroidea)

Fertilisation kinetics theory suggests that, when sperm are limiting, the larger eggs of broadcast-spawning marine organisms ought to be fertilised more frequently than smaller eggs, because they provide a bigger target for searching sperm. Whilst this effect has been demonstrated within species, it is not known if this pattern holds among species. We tested whether a large difference in egg size between congeneric seastars with contrasting planktotrophic and lecithotrophic modes of development results in differences in the likelihood of eggs being fertilised in sperm-limiting situations. Measurement of egg sizes and sperm swimming speeds led to the prediction that the sperm–egg collision rate constant for Patiriella calcar (420-µm-diameter egg) should be nine times greater than for P. regularis (140-µm-diameter egg). Although the eggs of P. calcar should be fertilised at greater rates in low sperm concentrations, they were not. When gametes were allowed to mix for 10 s, the hypothesis that P. calcar eggs required less sperm than P. regularis to ensure 50% of eggs were fertilised was rejected. When gametes were mixed for 5 min, P. regularis eggs were more frequently fertilised, but the difference was not statistically significant. We conclude there must be a difference between these species in the likelihood that when a sperm finds a conspecific egg it can successfully fertilise. This apparent uncoupling of egg size and likelihood of fertilisation suggests that fertilisation is not a major constraint on the evolution of egg size in these seastars.Communicated by G.F. Humphrey, Sydney     

Responses to sugars and their behavioural mechanisms in the starling (Sturnus vulgaris L.)

Summary The response of the starling to sugar solutions was investigated as this species, which eats sweet fruits, does not prefer sugar solutions according to the literature.In choice tests using two identical fountains, moderate preference for higher concentrations (0.5 M and 1 M) of glucose and fructose to water was demonstrated. The same concentrations of sucrose were rejected. Lower concentrations (0.25 M and less) of all three sugars were neither preferred nor rejected. In choice tests using two differently coloured fountains the subjects did not generally respond more sensitively to sugar solutions. In learning experiments with differently coloured fountains the starlings developed very marked preferences for 0.5 M glucose and 0.5 M fructose and a corresponding rejection of 0.5 M sucrose in the course of 1–2 days. In a learning experiment with dentical fountains they also developed a pronounced preference; the preference values obtained, however, are higher if secondary cues such as the colour of the fountain or its position are available. In short time tests immediate responses to sugar solutions could not be demonstrated.The rejection of sucrose is attributed to illness-induced aversion learning, the learned preference for glucose and fructose to the preference according to gain in energy per unit of time as postulated by the optimal foraging theory. Taste sensations play, if any, only a minor role. The learning mechanisms operating in the experiments could be employed by the starling for the selection of fruits.     

Pleistocene diversification of the Pomacentrus coelestis species complex (Pisces: Pomacentridae): historical biogeography and species boundaries

Pleistocene eustatic changes in sea level are often invoked to explain genetic divergence among marine organisms. However, molecular phylogenies have revealed relatively few examples of speciation events dating to the Pleistocene. We present a species-level hypothesis of the timing of evolution for the Pomacentrus coelestis species complex (Pomacentridae), based on the nuclear S7 intron and the mitochondrial Cytb gene, and reconstruct ancestral range distributions across the timetree. Ancestral range reconstruction suggests the complex originated in the Coral Triangle and East Indian Ocean, with subsequent range expansion outward from this region. We suggest that land barriers to dispersal (e.g., Indo-Pacific barrier) may be responsible for the divergence between Indian (P. alleni, P. similis, P. caeruleopunctatus, and P. caeruleus) and Pacific (P. micronesicus, P. auriventris, and P. coelestis) species groups, and subsequent isolation by Pleistocene sea-level fluctuations in certain areas of the Coral Triangle (glacial refugia) may play an important role in the diversification of this species complex. Additionally, our analyses show cryptic lineages within P. micronesicus and highlight the need for comprehensive sampling within and among species to reveal recent speciation events.     

Marine calanoid resting eggs in Norway: Abundance and distribution of two copepod species in the sediment of an enclosed marine basin

Abundance and distribution of calanoid resting eggs in sediment were examined in 1988–1989 in an enclosed marine basin (Svartatjønn; western Norway). The basin undergoes rotenone treatment and draining every winter. Egg densities were highest in autumn, when up to 2 × 10⁶ eggs m^–2 were found, and lowest in summer. Egg densities were also highest in the deeper parts of the basin and in the upper 1 cm of sediment. Hatching was initiated in late January by pumping warmer deep water from the outside area into the basin. Evidence is given for resting-egg formation inEurytemora affinis Poppe, 1880 andAcartia clausi Gurney, 1931 (present phenotype reclassified asA. teclae by Bradford 1976: N. Z. Jl mar. Freshwat. Res. 10: 159–202). Seasonal differences in inhibition of hatching, together with the longevity and tolerance of these eggs, strongly suggest that they are diapause eggs.     

Social network theory in the behavioural sciences: potential applications

Social network theory has made major contributions to our understanding of human social organisation but has found relatively little application in the field of animal behaviour. In this review, we identify several broad research areas where the networks approach could greatly enhance our understanding of social patterns and processes in animals. The network theory provides a quantitative framework that can be used to characterise social structure both at the level of the individual and the population. These novel quantitative variables may provide a new tool in addressing key questions in behavioural ecology particularly in relation to the evolution of social organisation and the impact of social structure on evolutionary processes. For example, network measures could be used to compare social networks of different species or populations making full use of the comparative approach. However, the networks approach can in principle go beyond identifying structural patterns and also can help with the understanding of processes within animal populations such as disease transmission and information transfer. Finally, understanding the pattern of interactions in the network (i.e. who is connected to whom) can also shed some light on the evolution of behavioural strategies.     

Calanoid copepod eggs in sea-bottom muds. II. Seasonal cycles of abundance in the populations of several species of copepods and their eggs in the Inland Sea of Japan

The seasonal cycles of abundance of populations of dominant calanoid copepods in the water column and of their eggs recovered from the bottom sediment in the central part of the Inland Sea of Japan are described. The numbers of both copepods and eggs fluctuated markedly with season in an essentially similar pattern among the 6 species studied (Tortanus forcipatus Giesbrecht, Calanopia thompsoni A. Scott, Acartia erythraea Giesbrecht, A. clausi Giesbrecht, Centropages abdominalis Sato, C. yamadai Mori). The density of eggs in the sea bottom was highest shortly before the population of adults and late copepodids disappeared from the plankton; the numbers of eggs then gradually decreased until the appearance of the next planktonic population.     

Morphological and fluorescence analysis of the Montastraea annularis species complex in Florida

The taxonomic status of the Montastraea annularis species complex is unclear. Much evidence has been accumulated to support the separation into 3 species, but the presence of intermediate morphotypes and the apparent lack of effective reproductive barriers in some areas are yet unexplained. Several authors have made a call for the introduction of new traits that can be used to resolve differences among closely related coral species. We collected skeletal and tissue samples from corals within the M. annularis species complex (15 each of M. annularis, M. faveolata, and M. franksi) and 10 morphological intermediates from several reefs in the Florida Keys. Multivariate analysis of corallite skeletal measurements supported the separation of the species complex into three taxa. We detected two main fluorescence emission peaks at 480 nm (turquoise) and 515 nm (green) that were not distributed equally among the three species. Every M. annularis colony had a major turquoise fluorescence peak. Some had a weak green secondary fluorescence peak. Colonies of M. faveolata and M. franksi had either the green or turquoise fluorescence peak, but at significantly different frequencies. The intermediate morphotypes proved to be highly heterogeneous with respect to both micromorphology and fluorescence, and their nature could not be fully explained. We were not able to separate the three species using fluorescence characters alone, however this new trait does increase our understanding of the taxonomic structure within the M. annularis species complex. Received: 21 January 2000 / Accepted: 1 September 2000     

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.     

Calanoid resting eggs in the Baltic Sea: implications for the population dynamics of Acartia bifilosa (Copepoda)

Resting eggs of Acartia spp. (presumably A. bifilosa Giesbr.) at densities between 65 and 125 eggs cm^-2 were found in sediment off the southern coast of Finland and represent the first observations of dormant calanoid eggs in the Baltic Sea. The highest egg densities were found at sediment depths between 4 and 8 cm. The hatching percentage varied between 0 and 90% in the different sediment layers and at different experimental temperatures (13, 16 and 19°C). Some nauplii emerged from the eggs extracted from sediment layers 9 to 10 cm beneath the surface layer and other nauplii emerged from eggs after 82 d of dark storage at 3°C. Light was not needed to trigger hatching; the nauplii were able to hatch in darkness when the eggs were resuspended in filtered seawater. Indirect evidence suggests that the marked spring peak of Acartia spp. nauplii seen in a monitoring data from the years 1973 to 1984 was derived from benthic eggs and not from spawning. The possible mechanisms for induction of hatching and the adaptive benefits of resting egg production for A. bifilosa are discussed.     

Model selection and model averaging in behavioural ecology: the utility of the IT-AIC framework

Behavioural ecologists often study complex systems in which multiple hypotheses could be proposed to explain observed phenomena. For some systems, simple controlled experiments can be employed to reveal part of the complexity; often, however, observational studies that incorporate a multitude of causal factors may be the only (or preferred) avenue of study. We assess the value of recently advocated approaches to inference in both contexts. Specifically, we examine the use of information theoretic (IT) model selection using Akaike’s information criterion (AIC). We find that, for simple analyses, the advantages of switching to an IT-AIC approach are likely to be slight, especially given recent emphasis on biological rather than statistical significance. By contrast, the model selection approach embodied by IT approaches offers significant advantages when applied to problems of more complex causality. Model averaging is an intuitively appealing extension to model selection. However, we were unable to demonstrate consistent improvements in prediction accuracy when using model averaging with IT-AIC; our equivocal results suggest that more research is needed on its utility. We illustrate our arguments with worked examples from behavioural experiments.     

Mitochondrial DNA polymorphisms reveal additional genetic heterogeneity within the Leptasterias hexactis (Echinodermata: Asteroidea) species complex

Sea stars of the Leptasterias hexactis complex are important members of the rocky intertidal community of the temperate and subarctic North American Pacific coasts. Recent attempts to determine the genetic structure and taxonomic relationships of these sea stars have not been completely successful, due to the combination of extreme morphological variability and relative scarcity of taxonomically-informative genetic (allozyme) polymorphisms. The present study used mitochondrial DNA restriction-site polymorphisms for PCR (polymerase chain-reaction)-amplified products to provide additional information about this complex. Five restriction-site haplotypes were identified in southern Alaska and in the Puget Sound region that were allozymically distinct. Because the allozymic differences (Nei's unbiased genetic differences >0.08) are maintained in sympatry, the haplotypes behave evolutionarily as conventional species. Although most haplotypes show deficiencies of heterozygotes for polymorphic allozyme loci, there is no evidence for the existence of cryptic species within any of the haplotypes. Rather than consisting of a few polytypic and cosmopolitan species, the L. hexactis complex is apparently composed of numerous species with more restricted ranges. Determining the exact ranges and taxonomic relationships of the various species will require additional study.     

Histological and biochemical bases of defense mechanisms in four species of Polybranchioidea ascoglossan molluscs

The multiple defensive strategies of four Mediterranean ascoglossan molluscan species, belonging to two different Polybranchioidea families, Cyerce cristallina (Trinchese, 1881) and Caliphylla mediterranea (Costa A., 1869) (Polybranchiidae), and Ercolania funerea (Costa A., 1867) and Placida dendritica (Alder and Hancock, 1843) (Stiligeridae), were studied. E. funerea, P. dendritica and C. mediterranea were collected from the lake Fusaro (Arco Felice, Naples) in 1992. C. cristallina was collected from Capo Miseno (Bay of Naples) in 1991. C. cristallina and E. funerea easily undergo autotomy of dorso-lateral appendages (cerata) followed by an extraordinarily quick (8 to 10 d) regeneration of the latter. Histological analyses showed the presence, at the basis of both normal and regenerated cerata of these species, of a muscular sphincter which facilitates the autotomic process. C. mediterranea and P. dendritica, which do not undergo autotomy, lack this anatomical feature but feed upon algae belonging to the Chaetomorpha and Bryopsis genera, and, as shown by electron microscopy studies, retain large quantities of chloroplasts which they use as camouflage amidst algae and to escape predation. E. funerea also exploits this behavioral defense mechanism. Histological investigations also revealed in the cerata of all four species several multi-cellular mucous glands responsible for the secretion of the slime typical for these molluscs. C. cristallina, E. funerea and P. dendritica secrete large amounts of slime, whose extracts displayed ichthyotoxic activity when assayed by the Gambusia affinis ichthyotoxicity assay, while extracts of C. mediterranea slime were not toxic. A chemical analysis of the slime, mantle and cerata led to the isolation of polypropionate - and -pyrones from all species except C. mediterranea. These secondary metabolites possess structures that differ only by the degree of methylation and the geometry of double bonds of the side chain and are specifically distributed in cerata and slime of C. cristallina and E. funerea. The pyrones also display different activities in the Hydra vulgaris regeneration assay and in the G. affinis ichthyotoxicity test and, depending on their structural features and tissue distribution, are likely to play a role either as defense allomones or as supportive inducers of cerata regeneration. In conclusion, combined biological observations and histological and chemical techniques generated valuable information on the defensive behavior of these four ascoglossan species which exploit various combinations of the same behavioral and/or chemical defensive strategies and thus successfully avoid predation.     

The behavioural basis of a species replacement: differential aggresssion and predation between the introduced Gammarus pulex and the native G. duebeni celticus (Amphipoda)

Previous studies have shown that differential predation by males on moulted female congenerics may be largely responsible for the elimination and replacement of the native Irish freshwater amphipod Gammarus duebeni celticus by the introduced G. pulex. Predation of moulted females occurs both shortly after their release from precopulatory mate-guarding and whilst they are being guarded by their mates. In the present study, two hypotheses concerning the underlying cause(s) of the differential predation pattern are tested. Firstly, female G. d. celticus may be more vulnerable to predation than female G. pulex due to the former being released from precopula guarding with the new exoskeleton in a less hardened state. Secondly, G. pulex may be an inherently more aggressive species than G. d. celticus during predatory interactions over guarded females. The first experiment indicated that differential predation was not mediated by species differences in the state of the female exoskeleton at the time of release from precopula by guarding males. The second experiment, however, showed that male G. pulex were significantly more aggressive than male G. d. celticus in attacking both guarding male congenerics and guarded moulted female congenerics. In addition, in defence against predatory attacks, paired male and female G. pulex were significantly more aggressive than paired male and female G. d. celticus. These differences in aggressive behaviour led to a significantly higher frequency of predation on G. d. celticus females than on G. pulex females, and also explains this finding in previous studies. It is concluded that differential predation due to differences in aggressive behaviour may explain the pattern of replacement between these species.     

The utility of the 16S gene in investigating cryptic speciation within the Brachionus plicatilis species complex

Recent reports indicate an extensive amount of molecular evolution separating cryptic taxa as well as significant population structure at a microgeographical scale. Appropriate molecular markers are particularly suitable for distinguishing cryptic biological species. In this study, we examine the phylogenetic utility of 16S rRNA in elucidating the evolutionary relationships within the recently described euryhaline Brachionus plicatilis species complex. In addition, we assess the applicability of this marker in the genetic identification and monitoring of rotifer populations. We have sequenced a 378-bp fragment of the mitochondrial 16S rRNA gene in laboratory reference strains, hatchery clones as well as collections from a wild population of the subsaline Lake Koroneia (Northern Greece). Also, restriction fragment length polymorphism (RFLP) analysis was performed with eight restriction endonucleases. Rotifer samples are distinguished into six genetically divergent lineages. Average sequence divergence between lineages is 0.1038. The evolutionary relationships and divergence time-scales revealed with the 16S sequence data are in agreement with previous analyses using different mitochondrial and nuclear markers. The 16S region appears to have several advantages over other regions of the genome regarding use of species-specific primers, ease of amplification from single specimens and undiluted informational content over both recent and more ancient separations. It has also exhibited maximum discriminatory power (100% success) between lineages during RFLP analysis. The 16S assayed region has proven especially informative and consistent in detecting, supporting and establishing the lineage status within the B. plicatilis species complex both from a phylogenetic perspective and as an identification tool.     

Precise sex ratio in a parasitic wasp: the result of counting eggs

Summary The parasitic wasp Nesolynx albiclavus maintains a very skewed offspring sex ratio (ca. 4% males). We studied proximate, causal factors of the son-daughter choice mechanism to obtain insight into the precision of the procedures and the underlying rules. We analyzed the number of males in clutches produced during sessions of 12 or 24 h. The results show that the timing of the next male egg depends on the time elapsed and the number of eggs laid since the previous one. The influence of the passage of time per se during non-laying periods was reduced in an experiment with 4-h periods. The data show that the first male egg is produced by a fairly rigid process: about the sixth egg in a run will be left unfertilized. No further male egg will appear in the next 10–20 eggs if they are laid in rapid succession.     

Genetic differentiation in two sibling species of the brackish-water polychaete Hediste japonica complex (Nereididae)

Genetic divergence among ten populations of small- and large-egg forms of the brackish-water polychaete Hediste japonica complex was investigated on 14 isozyme loci by electrophoretic analysis. The two forms were distinguishable by complete allele substitutions at five loci, resulting in high genetic differentiation (Nei's D: 0.533 to 0.662). No genetic evidence of hybridization between the two forms was detected in sympatric populations in three rivers. These results indicate that the two forms are reproductively isolated, clearly showing that the two forms are distinct species. The genetic differentiation among populations was higher in the large-egg form (D: 0.005 to 0.111, G _ST: 0.435) than that in the small-egg form (D: 0.000 to 0.001, G _ST: 0.020). This genetic difference between the two forms seems to be attributable to a difference in their life histories. The average expected heterozygosity was low in populations of both the large-egg form (0.005 to 0.068) and the small-egg form (0.014 to 0.038) in comparison with other marine invertebrates. Received: 11 April 1997 / Accepted: 8 September 1997     

Hatching with the enemy: Daphnia diapausing eggs hatch in the presence of fish kairomones

Summary. Infochemicals are known to play a key role in mediating predator-prey interactions, both in aquatic and terrestrial communities. However, state-dependent variation may exist in how effectively individuals can use this information, depending on genotype, life stage and experience. For our study, we used the predator-prey model system fish-waterflea Daphnia magna Straus (Cladocera, Daphniidae). Adult Daphnia use fish-derived infochemicals, so-called kairomones, as indicators of predation risk, and exhibit a spectrum of morphological, behavioural and life-history responses to the presence of fish kairomones. Here, we investigate whether diapausing eggs, an embryonic resting stage in the life cycle of D. magna, also use fish kairomones and tune their hatching to the risk of fish predation, as reported for diapausing stages of dinoflagellates. In two laboratory experiments, we studied hatching proportion and time until hatching of D. magna diapausing eggs in the absence and presence of fish kairomones. D. magna families differed significantly in their response to the presence of fish kairomones; some families reduced hatching proportion, whereas others increased it. Our results imply genotype-dependent differences in the hatching reactions to fish kairomones as observed for other traits in adult Daphnia.