Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

Growth of the kelp Ecklonia radiata near the northern limit of its range in Western Australia

The length, growth and survivorship of mature sporophytes of Ecklonia radiata (C.Ag.) J. Agardh were measured on two patch reefs within the lagoon of a high-latitude, coralreef atoll (Easter Group, Houtman Abrolhos; 28° to 29° S; 113°35 to 114°03E) for 1 yr (1982–1983). The sites differed in their proximity to the perimeter reef, but had similar regimes of temperature, light, nutrient concentration and water movement. Kelp length, growth rate and survivorship differed significantly between sites, although plant density was similar (less than l m^-2). At the site near the lagoon perimeter, the central lamina averaged 381 mm in length, grew at an annual mean rate of 1.28 mm d^-1, and 56% of tagged individuals had died after one year. The corresponding values for the site near the centre of the lagoon were: 257 mm, 0.75 mm d^-1 and 92%. Three other sites on the coastal limestone reefs near Perth (400 km to the south) were studied for various intervals over a 5 yr period (April 1979 to August 1984). The kelp from the low-latitude sites were smaller, grew more slowly, and suffered similar or greater mortality than their southern counterparts. Latitudinal comparisons were confounded by differences in kelp density between sites, but there is no evidence for density-dependent effects on the measured parameters across the range of natural densities observed. Growth rates at all sites were negatively correlated with ambient sea temperatures when these were above 20°C. There was no evidence of adaptation to the higher sea temperatures experienced at the Abrolhos, and temperature cannot be dismissed as a factor controlling the growth of the species near the northern limit of its distribution on the Western Australian coast. Other factors however must be involved in determining the latitudinal position of that limit.     

Interactions of methylmercury,cadmium, and salinity on regeneration in the fiddler crabs Uca pugilator,U. pugnax and U. minax

After multiple autotomy in the fiddler crabs Uca pugilator, U. pugnax and U. minax, both methylmercury and cadmium retard limb regeneration and ecdysis. When crabs in sea water are exposed to a mixture of both metals, the effect is increased, indicating that the two are interacting in an additive way. In 50% sea water (=15% S), the effects of cadmium are greatly intensified so that growth of limb buds is extremely slow, if it occurs at all. When methylmercury is present in the water at the same time, the severe effects of cadmium are somewhat ameliorated, indicating an antogonistic interaction of the two metals under these conditions. Adding additional calcium to the 50% sea water also decreased the severity of the cadmium effect, thus supporting the idea of a calcium-cadmium competition.     

Effects of time of year and limb removal on rates of ecdysis of eyed and eyestalkless fiddler crabs,Uca pugilator

The incidence of ecdysis among monthly samples of female Uca pugilator, both intact and eyestalkless, was determined from May, 1974 through April, 1975. Among both the intact and eyestalkless crabs the percentages of the crabs that underwent ecdysis varied during the year. During the height of the breeding season in particular, ecdyses were rare among the eyestalkless crabs and absent among the intact ones. The effects of limb removal, either followed by or preceded by eyestalk removal, on the rate of ecdysis were also determined. Eyestalkless crabs from which 4 walking legs were removed either the day before or the day after their eyestalks had been removed underwent ecdysis at a slower rate than crabs lacking only eyestalks, but faster than crabs lacking only 4 walking legs. The high incidence of precocious ecdyses induced by removing large numbers of limbs from eyed crabs appears to be due to the number of limbs removed and not to the weight or volume of limb tissue removed.     

Effect of salinity on hypoxia tolerance of resting green crabs, Carcinus maenas, after feeding

Mechanisms that can influence the tolerance of hypoxia in brackish waters were studied in resting and fed crabs, Carcinus maenas, at 15?°C. Mortality, blood oxygenation, acid-base status and lactate concentration were analysed in fed crabs held in full-strength normoxic seawater (32.5‰?S) and then transferred for 24?h to a partial pressure of oxygen (P_o2) of 3?kPa (1.4?mg?l^?1) and various salinities (17, 12.5, 10, 8‰?S). At salinity levels >10‰, fed crabs tolerated P_o2 values as low as 3?kPa in the ambient water and 0.5?kPa in their arterial blood for 24?h without switching to anaerobic metabolism. Only below 10‰?S did their blood-lactate content rise, leading to their death despite the fact that their blood O₂-content was twice the control value measured in full-strength normoxic seawater and their blood P_o2 did not decrease below values recorded at higher salinity levels. Addition of CO₂ to 8‰?S water (CO₂ partial pressure increasing from 0.1 to 0.3?kPa) decreased blood-lactate production and mortality, suggesting that at 10‰?S impairment of the O₂ supply is limited by an excessive blood O₂-affinity. The results are discussed in terms of the distribution (?10‰?S) of C. maenas along salinity gradients in estuaries and bays.     

Body size,morphological constraints,and mated pair formation in four populations of horseshoe crabs (Limulus polyphemus) along a geographic cline

The relationship of body size to mating success was studied in four populations of horseshoe crabs, Limulus polyphemus L., along the east coast of the U.S.A. in the spring and summer from 1986 to 1989. Crabs of both sexes from Great Bay, New Hampshire, were significantly smaller than crabs from three middle Atlantic coast populations: Sandy Hook Bay and Delaware Bay, New Jersey, and Chincoteague Bay, Virginia. The formation of mated pairs was independent of body size in each population; there were no significant size differences between mated and single individuals, and size assortative mating did not occur. A comparision of male clasper dimensions with the corresponding point of attachment on the female indicates that there are no morphological constraints limiting amplexus between any male and any female within a population. The ratios of male to female prosoma width within amplexed mated pairs averaged from 0.78 to 0.80 in each population, despite the large difference in absolute size between southern and nothern populations. This may suggest a role for natural selection in regulating the relative sizes of each sex.     

Aspects of population ecology in two populations of fiddler crabs, <Emphasis Type="Italic">Uca pugnax</Emphasis>

Fiddler crabs, Uca pugnax, were collected from a highly contaminated site and a relatively clean site, both in New Jersey to determine if and how environments with varying levels of pollutants may impact aspects of population biology including individual size, morphology (major cheliped size), population density, fecundity, recruitment and survivorship of early benthic phases. Crabs from the highly contaminated site were significantly larger in size, but had lower population density, lower recruitment, reduced reproductive season and lower survivorship of early benthic phases. Our study suggests that contamination may play a role in population ecology of U. pugnax. This study also determined that the reproductive season for U. pugnax in New Jersey is much longer than reported in the literature and could potentially be impacted by global climate change.     

An investigation of genetic population structure in blue crabs,Callinectes sapidus,using nuclear gene sequences

Spatial and temporal genetic heterogeneity in the sequences of nuclear protein-coding genes were investigated in populations of the blue crab, Callinectes sapidus. Haplotype frequencies of these novel markers were determined for adult crabs collected along 300 km in the northern Gulf of Mexico in two different years (2010 and 2011), as well as for megalopal recruits collected in 2010. Tests of genetic differentiation among all locations and between locations spanning known genetic breaks in other species were conducted. In addition, samples from distant locations within the range of C. sapidus were used to assess genetic divergence on a broader geographic scale. Significant between-year differences were found for adults at one location and near significant spatial differentiation was found across northern Gulf of Mexico locations in 2010. These results suggest that although the large population sizes and meroplanktonic life history of blue crabs promote widespread gene flow on a regional scale, genetic composition can change over just one year. Substantial divergence between the northern Gulf of Mexico and Venezuela suggests the possibility that temporal shifts in haplotype frequencies could result from variation in the rate of immigration from genetically distinct source populations. The possible effects of the Deepwater Horizon Oil Spill and attendant fisheries closures during the sampling phase of this study are also considered.     

Planktotrophic larval development is associated with a restricted geographic range in Lasaea,a genus of brooding,hermaphroditic bivalves

Members of the intertidal, near-cosmopolitan mollusc genus Lasaea brood their young either to a planktotrophic veliger or crawl-away juvenile stage of development. Developmental mode can be reliably inferred from brood masses and from prodissoconch structure. I have conducted a global developmental survey of this genus based mainly on examination of hundreds of museum lots. With one exception, Lasaea species with a planktotrophic larval development were restricted to the western Pacific. Congeners that lack planktotrophic larvae were found on all continents apart from Antarctica, and also on a large number of oceanic islands. These results indicate that (1) Lasaea species releasing crawl-away juveniles have a markedly greater collective geographic range than congeners with planktotrophic larvae; (2) pelagic larvae are not necessary for long-distance dispersal in this genus; (3) rafting has played a key role in the evolutionary success of the genus Lasaea; (4) cross-fertilizing Lasaea species with feeding larvae are less successful in utilizing chance rafting events to colonize new areas than are congeners lacking these traits.     

Temperature tolerance and latitudinal range of brown algae from temperate Pacific South America

Lower and upper survival temperatures of microthalli of 25 species of South American Phaeophyceae isolated from central Peru (14°S) to the Canal Beagle (55°S) were determined using 2-wk exposure for the upper and 4-wk exposure for the lower limit. All species survive 4 wks at -2°C. With respect to the upper limit, species reported only from southernmost South America tolerate 19.9 to 24.5°C (n=8), and species occurring from Cape Horn to central Chile 24.6 to 27.4°C (n=7). Three species that occurred as far north as northern Chile and Peru before the 1982–1983 El Niño event, and whose northern limit was dramatically shifted southwards in 1983, tolerate 20.8 to 25.3°C, whereas five species that have survived in Peru tolerate 25.6 to 28.5°C. Tinocladia falklandica which tolerates 27.8 to 28.1°C but lives only in southernmost South America and Striaria attenuata, which tolerates 31.6 to 31.9°C but occurs at ca. 42°S, are exceptional. Their high temperature tolerance may have no adaptive value in South America. They are restricted to the cold-temperate region due to low temperature requirements for reproduction or for reasons yet unknown. In general, the northern distributional limits of the Phaeophyceae studied along the temperate Pacific coast of South America are reproduction boundaries, except in El Niño years when they are redefined according to the species' upper suvival limits. Temperature tolerance of isolates from northern Chile and Peru agrees well with maximum temperatures reached during the 1983 El Niño.     

Genetic structuring and gene flow in Cerastoderma glaucum (Bivalvia: Cardiidae): evidence from allozyme variation at different geographic scales

Genetic variation at 16 enzyme loci was analysed in 20 Mediterranean and 1 Baltic population of Cerastoderma glaucum. Spatial genetic variation at different geographic scales was investigated. In general, this species was fairly genetically structured (over all loci Š=0.088), according to the fragmentary nature of its habitat. Spatial structuring of genetic diversity was shown to follow different models, depending on the geographic scale considered: a stepping-stone model provided a good fit at a wide scale, with gene flow inversely related to geographic distance, whereas at small scale, genetic relationships among samples could not be interpreted as simply the effect of physical distance among populations. Results are discussed taking into account the intrinsic and extrinsic factors which are most likely to affect the patterns of genetic structuring in coastal marine animals. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-001-0753-x     

Recruitment to adult habitats following marine planktonic development in the fiddler crabs, Uca pugilator, U. pugnax, and U. minax

Three congeneric species of fiddler crabs, Uca pugilator (Bosc, 1802), U. pugnax (Smith, 1870), and U. minax (LeConte, 1855), co-occur in estuaries along the east coast of North America, from Cape Cod to northern Florida. Although U. minax adults are generally found at lower salinities than the other two species, the distributions of all three species overlap to some degree. The distribution of megalopae and juvenile fiddler crabs (from first crab stage to those with a carapace width of 3.0 mm) was examined at four sites along a salinity gradient (from 35.0±2.0 to 3.0±3.0; ) in the North Inlet Estuary, South Carolina, USA, in August 2002. A PCR-RFLP technique was developed to identify individuals from the genus Uca to species from first zoea through the early crab stages. An examination of the distribution of early life stages showed that U. minax reinvades low-salinity adult habitats at settlement, following planktonic larval development in the coastal ocean. Also, juveniles of U. pugilator were found to occupy Spartina alterniflora stands, where adult conspecifics rarely occur. Species frequencies were different for adults compared to early life stages in low-salinity areas of the marsh, where populations overlap. Settlement and survival dynamics of early life-history stages in wet and dry years likely determine the distribution of adult Uca spp. populations along salinity gradients in estuarine ecosystems.Communicated by J.P. Grassle, New Brunswick     

Effects of salinity gradients on the tolerance and bioenergetics of juvenile blue crabs (Callinectes sapidus) from waters of different environmental salinities

Juvenile Callinectes sapidus Rathbun were collected from brackish and hypersaline coastal environments in August 1986 and July 1987, respectively. The brackish collection site was a salt-marsh near Grand Isle, Louisiana (USA), and the hypersaline site was in the barrier island system on the north end of the Laguna Madre near Corpus Christi, Texas (USA). On the dates of collection, salinities fluctuated daily between 20 and 30 S and between 30 and 45 S at the brackish and hypersaline collection sites, respectively. The high-salinity 21 d LC₅₀ (50% mortality) was 56.0 for brackish-water individuals and 66.5 S for hypersaline individuals. The brackish-water individuals survived 0 S. The lowsalinity 21 d LC₅₀ was 0.5 S for the hypersaline individuals. Respiration and excretion comprised a small portion of the energy budget and did not vary with salinity for individuals from brackish water. However, both respiration and excretion increased with decreasing salinity in individuals from the hypersaline environment. Respiration accounted for more energy than excretion. As energetic expenditure (due to respiration and excretion) was relatively small, scope for growth usually paralleled energy absorption. Scope for growth responses to salinity differed significantly between crabs from the two environments. Peaks in scope for growth for both the brackish-water and hypersaline individuals corresponded to salinities normally encountered by these crabs in their natural habitats. Individuals from the brackish-water population had maximal energy absorption and scope for growth at 10 and 25 S. Individuals from the hypersaline population displayed maximal energy absorption at 35 S and maximal scope for growth at 35 and 50 S.     

Environmental preferences of yellowfin tuna (Thunnus albacares) at the northern extent of its range

We used acoustic telemetry to examine the small-scale movement patterns of yellowfin tuna (Thunnus albacares) in the California Bight at the northern extent of their range. Oceanographic profiles of temperature, oxygen, currents and fluorometry were used to determine the relationship between movements and environmental features. Three yellowfin tuna (8 to 16 kg) were tracked for 2 to 3 d. All three fish spent the majority of their time above the thermocline (18 to 45 m in depth) in water temperatures >17.5 °C. In the California Bight, yellowfin tuna have a limited vertical distribution due to the restriction imposed by temperature. The three fish made periodic short dives below the thermocline (60 to 80 m), encountering cooler temperatures (>11 °C). When swimming in northern latitudes, the depth of the mixed layer largely defines the spatial distribution of yellowfin tuna within the water column. Yellowfin prefer to spend most of their time just above the top of the thermocline. Oxygen profiles indicated that the tunas encountered oceanic water masses that ranged most often from 6.8 to 8.6 mg O₂ l⁻¹, indicating no limitation due to oxygen concentrations. The yellowfin tuna traveled at speeds ranging from 0.46 to 0.90 m s⁻¹ (0.9 to 1.8 knots h⁻¹) and frequently exhibited an oscillatory diving pattern previously suggested to be a possible strategy for conserving energy during swimming. Received: 14 February 1997 / Accepted: 14 April 1997     

Phenological mismatch and the effectiveness of assisted gene flow

The persistence of narrowly adapted species under climate change will depend on their ability to migrate apace with their historical climatic envelope or to adapt in place to maintain fitness. This second path to persistence can only occur if there is sufficient genetic variance for response to new selection regimes. Inadequate levels of genetic variation can be remedied through assisted gene flow (AGF), that is the intentional introduction of individuals genetically adapted to localities with historic climates similar to the current or future climate experienced by the resident population. However, the timing of reproduction is frequently adapted to local conditions. Phenological mismatch between residents and migrants can reduce resident × migrant mating frequencies, slowing the introgression of migrant alleles into the resident genetic background and impeding evolutionary rescue efforts. Focusing on plants, we devised a method to estimate the frequency of resident × migrant matings based on flowering schedules and applied it in an experiment that mimicked the first generation of an AGF program with Chamaecrista fasciculata, a prairie annual, under current and expected future temperature regimes. Phenological mismatch reduced the potential for resident × migrant matings by 40–90%, regardless of thermal treatment. The most successful migrant sires were the most resident like in their flowering time, further biasing the genetic admixture between resident and migrant populations. Other loci contributing to local adaptation—heat‐tolerance genes, for instance—may be in linkage disequilibrium with phenology when residents and migrants are combined into a single mating pool. Thus, introgression of potentially adaptive migrant alleles into the resident genetic background is slowed when selection acts against migrant phenology. Successful AGF programs may require sustained high immigration rates or preliminary breeding programs when phenologically matched migrant source populations are unavailable.     

Richness,abundance and shell use of subarctic and arctic hermit crabs

Hermit crabs are an obvious, common and abundant feature of global shallow water environments but almost nothing is known of their ecology at their extreme latitudinal ranges, the Arctic and southern South America. In this study, we report the first investigation on hermit crabs and their use of a key resource, gastropod shells, in a subpolar and a polar environment—amongst the most rapidly changing places on earth. Hypothesised low levels of richness were confirmed by surveys—only Pagurus pubescens was found in western Spitsbergen and only three pagurids were found in northern Norway. At the northern-most of their extent, hermit crabs were fairly common but Arctic abundances (1–5 m⁻²) were an order of magnitude lower than in many warm temperate or tropical localities. Along the open coast of Spitsbergen, the occurrence of P. pubescens was infrequent and very patchy, but it was more abundant in the fjords. In Isfjorden, the largest studied fjord, the population of P. pubescens was mainly represented by small individuals. Spitsbergen P. pubescens used few shell types and >87% just occurred in Margarites or Buccinum shells. The proportion of the gastropod shells, used by P. pubescens, which were damaged, was high and increased with shell size. These hermit crabs are at the edge of the range for both their species and Anomura. The extremes of their location are reflected by: They probably represent the least rich assemblage, with the lowest and most patchy typical abundances reported to date and are amongst the smallest individuals using the least diverse and most damaged gastropod shells.     

Temperature and pressure tolerance of larvae of Crepidula fornicata suggest thermal limitation of bathymetric range

The extant deep-sea fauna is thought to result from recolonisation of this environment by shallow-water organisms following climate-driven mass extinctions. Planktonic larval tolerance to high pressure is considered an important preadaptation for successful deep-sea invasion. In this study, the pressure and temperature tolerance of a species without any known confamilial deep-sea relative were assessed for the first time. Early- and late-veliger larvae of the shallow-water species Crepidula fornicata were subjected to a temperature/hydrostatic pressure regime from 5 to 25 °C and from 0.1 to 40 MPa. Although early and late veliger survived pressures equivalent to 2,000 m water depth or greater at all temperatures, decreased larval activity indicated significant sublethal temperature and pressure effects. Reduced larval activity of early veliger at low temperatures suggests that the bathymetric range of this species may be thermally constrained. A mechanistic model is proposed to explain the emerging pattern of ontogenetic shifts in pressure tolerance of shallow-water benthic invertebrates.