Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type. An erratum to this article can be found at     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type.     

Reproductive effort in four species of intertidal limpets

Reproductive effort was measured in 4 species of intertidal limpets: Cellana tramoserica (Sowerby) Notoacmea petterdi (Tenison Woods), Patella peroni Blainville and Patelloida alticostata (Angas) from south-eastern Australia. Field studies between 1971 and 1975 enabled indices of reproductive effort to be obtained using both gonad to body weight ratios and energy budgets. The ratio of annual gonad production to somatic tissue energy content is highest for Patella peroni (2.19). The ratios for C. tramoserica and Patelloida alticostata are lower but similar (1.02), and N. petterdi has the lowest ratio (0.78). The percentage of assimilated energy allocated to reproduction is also highest in Patella peroni (26.6%), but this index of reproductive effort is similar for the other three species (10.3 to 12.9%). Consequently, these 4 species are ranked in different orders with respect to reproductive effort, depending upon which index is used. However, the difference between both indices is small, and is less than has usually been anticipated. The difference is interpreted as being the result of the low metabolic rate of N. petterdi rather than as evidence for the inadequacy of one of the indices of reproductive effort.     

Effects of contrasting modes of larval development on the genetic structures of populations of three species of prosobranch gastropods

 In south-eastern Australia, the prosobranch gastropods Morula marginalba (Blainville), Cominella lineolata (Lamarck) and Bedeva hanleyi (Angas) have similar fine-scale distributions, but appear to possess very different dispersal capabilities due to contrasting modes of larval development. M.marginalba produce planktonic larvae, whereas C. lineolata and B. hanleyi undergo direct development in benthic egg capsules and emerge as crawling juveniles. To test for possible effects of contrasting life histories on levels of genetic variation within and among populations, a survey was conducted of allozyme variation at six polymorphic loci in 8 to 9 local populations of each species. Collections of snails were made between June 1992 and November 1993. Sampling ranges spanned between 162 and 180 km of coast. Regardless of larval type, proportions of single-locus genotypes in each collection were consistent with the recruitment of offspring which had been generated through random mating. However, genotypic diversity was lower in those species that undergo direct development. Loci surveyed in C. lineolata and B. hanleyi were polymorphic (i.e. frequency of most common allele <95%) in fewer populations than those examined for M.␣marginalba (P <0.001) and, where polymorphisms occurred, also possessed significantly fewer alleles (P <0.001). Consequently, average levels of expected heterozygosity were greater in populations of M. marginalba than in those of either of the other species (P <0.001). Genetic variation among populations, expressed as the standardised variance in allele frequencies (F _ST ), was inversely related to expected larval dispersal capability. The nine collections of M. marginalba showed little overall differentiation (F _ST  = 0.017; P <0.001), reflecting the ability of planktonic larvae to interconnect local populations, and so limit divergence due to drift and natural selection. In contrast, there were high levels of allelic heterogeneity among the nine collections of C. lineolata (F _ST  = 0.523; P <0.001) and eight collections of B. hanleyi (F _ST  = 0.140; P <0.001). These data imply that for species which undergo direct development, local populations are effectively closed and evolve largely independent of one another. Received: 3 May 1996 / Accepted: 12 July 1996     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Spatial characteristics of habitat patches and population survival

The effect of the spatial arrangement of habitat patches on the survival of resident populations were considered in a stochastic model using population parameters appropriate to Peromyscus leucopus. The 34 possible arrangements of connections among five otherwise identical patches were simulated in order to determine the survival probabilities and population sizes.The main findings are that populations in completely isolated patches have lower survival probabilities than those in patches that are connected to other patches, are connected to the survival probabilities of populations in connected patches increases with the size of the largest geometric figure of which the patch is a part. The results are discussed in the context of resource management.     

Urban marine ecology in southern California: the ability of riprap structures to serve as rocky intertidal habitat

Increasing human populations along marine coastlines has lead to increasing urbanization of the marine environment. Despite decades of investigations on terrestrial ecosystems, the effect of urbanization on marine life is not well understood. Riprap is the rocky rubble used to build jetties, breakwaters, and armored shorelines. Roughly 30% of the southern California shoreline supports some form of riprap, while 29% of the shoreline is natural rocky substrate. Astonishingly few studies have investigated this anthropogenic rocky habitat even though it rivals a natural habitat in area on a regional scale along a coastline that has been extensively studied. In this study, I compared the diversity and community structure of exposed rocky intertidal communities on four riprap and four natural sites in southern California. I ask the following questions: (1) does diversity or community composition differ between intertidal communities on riprap and natural rocky habitats in southern California, (2) if so, which organisms contribute to those differences, (3) which physical factors are contributing to these differences, and (4) do riprap habitats support higher abundances of invasive species than natural habitats? On average, riprap and natural rocky habitats in wave exposed environments in southern California did not differ from each other in diversity or community composition when considering the entire assemblage. However, when only mobile species were considered, they occurred in greater diversity on natural shores. These differences appear to be driven by wave exposure. The presence of invasive species was negligible in both natural and riprap habitats.     

Climate change and population genetic structure of marine species

Climate change influences populations by reducing or extirpating local populations, by disrupting patterns of migration and by shifting geographical distributions. These events can affect genetic population structure in several ways. Molecular markers have been used in numerous population genetic and phylogeographical studies of marine species and have detected population responses to climate change in the last few decades, such as range expansions, adaptative shifts and declines or increases in abundance. Little is known, however, about the molecular and physiological basis of adaptive responses to climate change in marine Mediterranean species. The Mediterranean Sea ecosystem is a ‘living laboratory’ with native species that are challenged by environmental change and by invasive species and a ‘gene-climate’ approach should be adopted as a way of focusing on the relationship between climate warming and genetic diversity.     

Contrasting population genetic structures of sympatric, mass-spawning Caribbean corals

Coral reef conservation management policy often focuses on larval retention and recruitment of marine fish with scant data available on important, less motile reef-building species such as corals. To evaluate the concept of population connectivity in corals, we tested whether broadcast spawning reproduction per se confers the same degree of dispersal to two sister species, Montastraea annularis (Anthozoa: Scleractinia; Ellis and Solander 1786) and M. faveolata (Ellis and Solander 1786), both dominant taxa in reefs of the northern Caribbean. Genetic analyses of ten nuclear DNA loci (seven microsatellite and three single-copy RFLP) reveal strikingly different patterns of population genetic subdivision for these closely related, sympatric species, in spite of likely identical dispersal abilities. Strong population genetic structure typified the architecture of M. annularis, whereas M. faveolata populations were principally genetically well mixed. A higher level of clonality was observed in M. annularis potentially because of a susceptibility to physical fragmentation. Clonality did not, however, significantly contribute to population genetic structure or low-level Hardy–Weinberg and linkage disequilibria observed in some populations. The lack of consistent association between reproductive mode and dispersal reinforces the perspective that population connectivity is not so much a function of predictable marine population source and sink relationships as is due to a more complex interface of oceanic currents interacting with and amplifying stochastic fluctuations in larval supply and settlement success. Our results support others promoting an overall ecosystem approach in marine protected area design.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Morphological and habitat divergence in the intertidal limpet Patelloida pygmaea

In the intertidal limpet Patelloida pygmaea, two distinctive morphs, the forms pygmaea and conulus, have been recorded. The former possesses a flat elongated shell, and the latter has an extremely high round shell. It has been observed in the field that pygmaea is found on oyster shells Crassostrea gigas. The form conulus uses an unusual substrate for attachment. It is found on the living shells of the intertidal gastropod Batillaria cumingi. Although conulus is normally found only on shells of Batillaria, it can also be found on oyster shells when pygmaea and Batillaria shells are not present in nature. An electrophoretic analysis of allozymes showed that these two forms are reproductively isolated from each other and coexist without gene exchange on the same mudflat. Laboratory experiments showed that pygmaea prefers oyster shells and conulus prefers Batillaria shells as substrates for attachment when both oyster and Batillaria shells are present. The form pygmaea did not attach to Batillaria shells, even when only Batillaria shells were available. However, conulus also attached to oyster shells when Batillaria shells were not available. The proportion of individuals of conulus that attached to oyster shells decreased significantly when pygmaea was attaching to the oyster shells. These results suggest that pygmaea is ecologically more specialized to living on oyster shells than conulus.     

Hydrology influences population genetic structure and connectivity of the intertidal amphipod Corophium volutator in the northwest Atlantic

The mechanisms driving genetic structure in marine systems are elusive due to the difficulty of identifying temporal and spatial barriers to dispersal. By studying marine invertebrate species with limited dispersal potential, genetic structure can be directly related to physical and biological factors restricting connectivity. In the northwest Atlantic, the benthic brood-rearing amphipod Corophium volutator is distributed across basins with distinct circulation patterns and has the potential to disperse passively during its adult stage. We analyzed spatial genetic variation and migration rates across C. volutator’s North American range with sequence data for mitochondrial DNA and three novel nuclear markers using frequency and coalescent-based methods. We found low genetic differentiation within basins, but strong subdivision within the Bay of Fundy and a striking biogeographic break between the Bay of Fundy and Gulf of Maine, suggesting that genetic drift may act on populations in which connectivity is restricted due to the limitation of passive dispersal by hydrological patterns.     

Revealing life‐history traits by contrasting genetic estimations with predictions of effective population size

Effective population size, a central concept in conservation biology, is now routinely estimated from genetic surveys and can also be theoretically predicted from demographic, life‐history, and mating‐system data. By evaluating the consistency of theoretical predictions with empirically estimated effective size, insights can be gained regarding life‐history characteristics and the relative impact of different life‐history traits on genetic drift. These insights can be used to design and inform management strategies aimed at increasing effective population size. We demonstrated this approach by addressing the conservation of a reintroduced population of Asiatic wild ass (Equus hemionus). We estimated the variance effective size (N_ev) from genetic data () and formulated predictions for the impacts on N_ev of demography, polygyny, female variance in lifetime reproductive success (RS), and heritability of female RS. By contrasting the genetic estimation with theoretical predictions, we found that polygyny was the strongest factor affecting genetic drift because only when accounting for polygyny were predictions consistent with the genetically measured N_ev. The comparison of effective‐size estimation and predictions indicated that 10.6% of the males mated per generation when heritability of female RS was unaccounted for (polygyny responsible for 81% decrease in N_ev) and 19.5% mated when female RS was accounted for (polygyny responsible for 67% decrease in N_ev). Heritability of female RS also affected N_ev; (heritability responsible for 41% decrease in N_ev). The low effective size is of concern, and we suggest that management actions focus on factors identified as strongly affecting , namely, increasing the availability of artificial water sources to increase number of dominant males contributing to the gene pool. This approach, evaluating life‐history hypotheses in light of their impact on effective population size, and contrasting predictions with genetic measurements, is a general, applicable strategy that can be used to inform conservation practice.     

The genetic effects of larval dispersal depend on spatial scale and habitat characteristics

The intertidal gastropods Bembicium vittatum and Austrocochlea constricta, which have direct and planktonic larval development, respectively, occur sympatrically at sites across a number of islands at the Houtman Abrolhos archipelago and two harbours at Albany in Western Australia. Their distribution provide an opportunity to examine the effect of dispersal ability on levels of genetic subdivision at a number of spatial scales. F _ST (standardised variance in allelic frequencies) values in the range 0.361 to 0.396, determined from allozyme frequencies at 12 to 13 polymorphic loci, confirm isolation of Abrolhos and Albany populations, which are separated by 900 km of coastline, in both species. Within the Abrolhos and Albany, levels of subdivision in B. vittatum were high, but similar, as indicated by F _ST values of 0.091 and 0.090, respectively. In A. constricta, a mean value of 0.160 at the Abrolhos suggests severe restrictions to gene flow, while 0.021 at Albany indicates much stronger connections among populations. F _ST values at the Abrolhos support previous suggestions that this archipelago favours genetic subdivision in both direct and planktonic-developing species. The Albany harbours favoured subdivision only in B. vittatum, the low values of F _ST in A. constricta being attributed to strong mixing between the harbours, thus facilitating gene flow via planktonic larvae. The isolation of A. constricta populations at the Abrolhos can be explained in terms of highly localised recruitment, the result of limited water movement in complex intertidal habitats. The study illustrates the value of examining sympatric direct and planktonic developers in assessing the role of larval dispersal in patterns of genetic subdivision, and concludes that planktonic larvae may not promote gene flow over broad or even some fine spatial scales.     

Distribution,diet and habitat selection by four sympatrically breeding gull species in the south-eastern North Sea

Food choice, feeding habitat use and spatial distribution of black-headed gulls ( Larus ridibundus), mew gulls ( L. canus), herring gulls ( L. argentatus) and lesser black-backed gulls ( L. fuscus) were studied in the south-eastern North Sea. At-sea distribution during the breeding period was assessed by transect counts from ships. Clear differences could be established between the four species, with lesser black-backed gulls being widespread and numerous far out at sea and mew and black-headed gulls being restricted to the coastal area; herring gulls showed an intermediate pattern. The diet of these sympatrically breeding gulls was analysed from pellet and faecal samples collected at two colonies. Considering biases in this method, the following components appeared to comprise the principal food for each species: black-headed gulls: bivalves, crustaceans; mew gulls: bivalves, polychaetes; herring gulls: bivalves, crustaceans; and lesser black-backed gulls: fish and crustaceans. Black-headed and mew gulls had much broader dietary niches than herring and lesser black-backed gulls. Based on spatial distribution, food choice and feeding habitat use, three types of foraging strategies are apparent: (1) black-headed and mew gulls forage both intertidally and on land, (2) herring gulls feed most frequently in the intertidal zone and (3) lesser black-backed gulls feed at sea. The foraging behaviour of these four species may well have influenced their current population trends in the study area.     

Comparisons between the diets of four abundant species of elasmobranchs in a subtropical embayment: implications for resource partitioning

The diets of one ray species (Rhinobatus typus) and three shark species (Carcharhinus cautus, Negaprion acutidens, Rhizoprionodon acutus) undergo size-related changes and differ among these species in the nearshore waters of a large subtropical embayment (Shark Bay) in which these elasmobranchs are abundant, thereby reducing the potential for competition for food within and among these four species. R. typus fed almost exclusively on penaeid prawns and portunid crabs, which is reflected in its narrow dietary breadth, whereas different species of teleosts constituted a major component of the diets of each size class of the three shark species. The prey consumed by the three shark species was diverse, with representatives of 15 teleost families being consumed by C. cautus and substantial volumes of cephalopods being ingested by that species and R. acutus. The pronounced differences in the diet of the single ray species and three shark species reflect differences between a bottom-dwelling and more pelagic life, and between modes of feeding and relative mouth sizes. The relative contributions of the different species of teleost to the diets of the three shark species varied. Thus, although each of these species fed on atherinids, labrids and sillaginids, C. cautus also consumed substantial amounts of platycephalids and terapontids and R. acutus and N. acutidens also ingested considerable amounts of clupeids. Furthermore, R. acutus, which is the only one of the four species that typically occurs over seagrass, was the only species that fed on the centropomid Psammoperca waigensis, which is very abundant in seagrass meadows. However, the sparid Rhabdosargus sarba, which lives in unvegetated areas, was never ingested by R. acutus, but was consumed by C. cautus and N. acutidens. As the individuals of R. typus increased in size, they progressively consumed proportionately smaller volumes of the penaeid prawns Penaeus merguiensis and Melicertus latisulcatus and relatively greater volumes of the portunid crab Portunus pelagicus, which is slightly larger and has a harder exoskeleton. In addition to teleosts, large C. cautus ingested substantial volumes of portunid crabs and ophidian reptiles, presumably sea snakes, while large N. acutidens also fed on the ray R. typus.Communicated by G.F. Humphrey, Sydney     

A comparative analysis of sperm storage and mating strategies in five species of brachyuran crabs

Variation in female sperm storage is explained, in part, by the amount of sperm transferred at mating. Laboratory mating experiments were conducted on Eurypanopeus depressus and Rhithropanopeus harrisii from the Chesapeake Bay and Pachygrapsus transversus from Florida, while mated pairs of Uca beebei and U. terpsichores were collected from mudflats in Panama. All experiments and collections were conducted during the summer of 2006 and 2007. More sperm was transferred to larger than smaller females, and by species with long copulation durations (R. harrisii and E. depressus). These two species live in cryptic habitats, have high sperm/egg ratios, and likely store sperm across multiple broods. In contrast, P. transversus and U. beebei mate conspicuously, have short copulations, transfer fewer sperm, and have low sperm/egg ratios. Comparisons of sperm transfer across different mating strategies and habitats provide a better understanding of female sperm storage in the Brachyura.     

Escape responses of four Chilean intertidal limpets to seastars

The Chilean intertidal Lottiidae limpet fauna consists of at least nine species, grouped into the monophyletic taxon Scurria and a single taxon preliminarily designated as `Lottia' orbignyi (Dall). Within the Scurria clade, the most basal species are Scurria variabilis (Sowerby), S. viridula (Lamarck), and S.zebrina (Lesson), and among the most crown groups are S. ceciliana (Orbigny) and S. araucana (Orbigny). This phylogenetic information, obtained by the analysis of molecular characters (16S mtDNA), provides a framework for a comparative study of behavioral characters. In this study, we analyze the escape responses of two basal limpet species (`L.' orbignyi and S. viridula) and two derived limpet species (S. araucana and S. ceciliana) in the presence of the limpet predatory seastar Heliaster helianthus (Lamarck) and the limpet non-predatory seastars Stichaster striatus Muller and Troschel and Patiria chilensis Lutken. Neither P.chilensis nor Stichaster striatus induced escape responses such as those observed with the predatory H. helianthus. Moreover, in the presence of H. helianthus, basal and derived limpet species differed significantly in the percentage of individuals responding, reaction time, and duration of the response. The basal species `L.' orbignyi and Scurriaviridula exhibited instantaneous and vigorous locomotor responses, whereas the derived species S. araucana and S. ceciliana displayed no locomotor responses. The same pattern was found in responses to H. helianthus homogenates. The results strongly suggest a correlation between phylogeny and anti-predator escape responses to the seastar H. helianthus. Alternative escape responses have evolved for basal and derived species within the Chilean clade. Received: 27 April 2000 / Accepted: 16 August 2000     

Effects of settlement density on spatial arrangement in four intertidal barnacles

Balanus amphitrite was studied in an estuary (Halifax River) in central Florida from October to November 1990, while three other barnacle species were studied on the central coast (Monterey Bay) of California from April 1988 to July 1989. Mean nearest-neighbor distances indicated that in the majority of cases the spatial arrangement of settlers was random for three different balanomorph barnacles-B. amphitrite, B. glandula, and Chthamalus dalli. In Pollicipes polymerus, a lepidomorph species, strong aggregation among settlers was almost always observed. In the three balanomorph species there was no apparent relationship between settler density and the degree of aggregation. In P. polymerus there was a significant positive correlation between settler density and the degree of aggregation. Morphological differences between balanomorph and lepidomorph barnacles may influence the evolution of settlement behavior.     

The role of habitat heterogeneity in structuring the community of intertidal free-living marine nematodes

The role of habitat complexity has been widely neglected in the study of meiofaunal community patterns. We studied the intertidal nematode community of a structurally complex macrotidal beach exhibiting contrasting microhabitats (sandbars and runnels) to understand the influence of environmental gradients and habitat heterogeneity in the community structure. We tested whether topographical complexity affected (1) the zonation pattern in terms of abundance and diversity, and (2) local diversity by promoting compartmentalization into distinct faunal groups. Our analyses revealed three major faunal assemblages along the exposure gradient associated to differences in mean grain size and chlorophyll a. Diversity patterns involved a mid-intertidal peak, consistent with the intermediate disturbance hypothesis, and another peak at the limit with the subtidal region, consistent with the transition zone. These results highlight the predominance of environmental gradients in establishing intertidal zonation. However, microhabitats differed in environmental conditions and possessed significantly distinct nematofaunal communities. Runnels featured higher levels of taxonomic and functional diversity, many unique genera, and the community differed from the assemblage at the limit to the subtidal, stressing their role as distinct microhabitats. The nematofauna of the structurally complex beach was more diverse than the one from a homogeneous beach nearby, supporting the hypothesis that structural heterogeneity promotes diversity by compartmentalization and highlighting the importance of microhabitats in the assessment of biodiversity. Contrary to previous predictions, our results indicate potentially high regional marine nematode diversity in the Upper Gulf of California.