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In intertidal communities, consumers (especially carnivorous gastropods) have historically been thought to exert strong top-down control on community composition by regulating competitively dominant mussels and barnacles. This paradigm was formulated based on wire mesh cage exclusion experiments, which have well-known artifacts such as altering hydrodynamics and excluding non-target, but potentially important, consumer species. Recent research highlights the potential importance of multiple consumers that are often subtle and transient, as well as the modifications of consumer pressure by spatial and temporal environmental variability and bottom-up processes. Manipulative experiments that target individual taxa will be essential to more clearly identify their roles in complex ecological communities. The predatory gastropod Nucella lapillus has long been considered an important consumer controlling the structure and dynamics of intertidal communities in the Gulf of Maine. To test the role of N. lapillus in shaping community structure, we manually reduced its densities for 2.33 years. Species composition, stable stage community composition (based on a Markov model), and transition probabilities (as measures of ecological pathway strength) were compared between treatments (control vs. density reductions), and among seasons (spring, summer, fall, winter). In contrast to previous studies, exclusions had no effect on community composition or long-term Markov model predictions of stable stage community structure. Reducing N. lapillus abundance increased the persistence and reduced the mortality of the barnacle Semibalanus balanoides in the model, but did not affect blue mussels, Mytilus edulis , in a similar way. Reducing N. lapillus abundance had additional indirect effects of increasing Ralfsia spp. mortality and increasing hydroid persistence. Despite differences in transition probabilities among treatments, compensatory changes in direct and indirect pathways led to communities that converged over the long-term. Our results contrast previous estimates of the role of N. lapillus in intertidal communities and might reflect low mussel recruitment, predation by alternate consumers (e.g., C. maenas , C. borealis , Tautogolabrus adspersus , birds), or spatial and/or temporal environmental variation that influenced the role of N. lapillus in structuring these communities. The role of N. lapillus may be more limited, or variable, than often assumed in the Gulf of Maine, and the methods manipulating broad functional groups such as “consumers”, though useful in developing basic conceptual models, blur the effects of individual species in community assembly and dynamics.

With anthropogenic impacts rapidly advancing into deeper waters, there is growing interest in establishing deep-sea marine protected areas (MPAs) or reserves. Reserve design depends on estimates of connectivity and scales of dispersal for the taxa of interest. Deep-sea taxa are hypothesized to disperse greater distances than shallow-water taxa, which implies that reserves would need to be larger in size and networks could be more widely spaced; however, this paradigm has not been tested. We compiled population genetic studies of deep-sea fauna and estimated dispersal distances for 51 studies using a method based on isolation-by-distance slopes. Estimates of dispersal distance ranged from 0.24 km to 2028 km with a geometric mean of 33.2 km and differed in relation to taxonomic and life-history factors as well as several study parameters. Dispersal distances were generally greater for fishes than invertebrates with the Mollusca being the least dispersive sampled phylum. Species that are pelagic as adults were more dispersive than those with sessile or sedentary lifestyles. Benthic species from soft-substrate habitats were generally less dispersive than species from hard substrate, demersal or pelagic habitats. As expected, species with pelagic and/or feeding (planktotrophic) larvae were more dispersive than other larval types. Many of these comparisons were confounded by taxonomic or other life-history differences (e.g. fishes being more dispersive than invertebrates) making any simple interpretation difficult. Our results provide the first rough estimate of the range of dispersal distances in the deep sea and allow comparisons to shallow-water assemblages. Overall, dispersal distances were greater for deeper taxa, although the differences were not large (0.3-0.6 orders of magnitude between means), and imbalanced sampling of shallow and deep taxa complicates any simple interpretation. Our analyses suggest the scales of dispersal and connectivity for reserve design in the deep sea might be comparable to or slightly larger than those in shallow water. Deep-sea reserve design will need to consider the enormous variety of taxa, life histories, hydrodynamics, spatial configuration of habitats and patterns of species distributions. The many caveats of our analyses provide a strong impetus for substantial future efforts to assess connectivity of deep-sea species from a variety of habitats, taxonomic groups and depth zones.

Little is known about how larvae disperse in deep ocean currents despite how critical estimates of population connectivity are for ecology, evolution and conservation. Estimates of connectivity can provide important insights about the mechanisms that shape patterns of genetic variation. Strong population genetic divergence above and below about 3000 m has been documented for multiple protobranch bivalves in the western North Atlantic. One possible explanation for this congruent divergence is that the Deep Western Boundary Current (DWBC), which flows southwestward along the slope in this region, entrains larvae and impedes dispersal between the upper/middle slope and the lower slope or abyss. We used Lagrangian particle trajectories based on an eddy-resolving ocean general circulation model (specifically FLAME – Family of Linked Atlantic Model Experiments) to estimate the nature and scale of dispersal of passive larvae released near the sea floor at 4 depths across the continental slope (1500, 2000, 2500 and 3200 m) in the western North Atlantic and to test the potential role of the DWBC in explaining patterns of genetic variation on the continental margin. Passive particles released into the model DWBC followed highly complex trajectories that led to both onshore and offshore transport. Transport averaged about 1 km d−1 with dispersal kernels skewed strongly right indicating that some larvae dispersed much greater distances. Offshore transport was more likely than onshore and, despite a prevailing southwestward flow, some particles drifted north and east. Dispersal trajectories and estimates of population connectivity suggested that the DWBC is unlikely to prevent dispersal among depths, in part because of strong cross-slope forces induced by interactions between the DWBC and the deeper flows of the Gulf Stream. The strong genetic divergence we find in this region of the Northwest Atlantic is therefore likely driven by larval behaviors and/or mortality that limit dispersal, or local selective processes (both pre and post-settlement) that limit recruitment of immigrants from some depths.

The deep ocean supports a highly diverse and mostly endemic fauna, yet little is known about how or where new species form in this remote ecosystem. How speciation occurs is especially intriguing in the deep sea because few obvious barriers exist that would disrupt gene flow. Geographic and bathymetric patterns of genetic variation can provide key insights into how and where new species form. We quantified the population genetic structure of a protobranch bivalve, Neilonella salicensis, along a depth gradient (2200‐3800 m) in the western North Atlantic using both nuclear (28S and calmodulin intron) and mitochondrial (cytochrome c oxidase subunit I) loci. A sharp genetic break occurred for each locus between populations above 2800 m and below 3200 m, defining two distinct clades with no nuclear or mitochondrial haplotypes shared between depth regimes. Bayesian phylogenetic analyses provided strong support for two clades, separated by depth, within N. salicensis. Although no morphological divergence was apparent, we suggest that the depth-related population genetic and phylogenetic divergence is indicative of a cryptic species. The frequent occurrence of various stages of divergence associated with species formation along bathymetric gradients suggests that depth, and the environmental gradients that attend changes in depth, probably play a fundamental role in the diversification of marine organisms, especially in deep water. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, ••, ••‐••.

The Pleistocene and post-Pleistocene evolutionary history of many North Atlantic intertidal invertebrate species is well known, but the evolutionary history of the deep North Atlantic fauna is poorly understood, specifically whether colonization of the deep North Atlantic paralleled the patterns observed in shallow water. Contemporary pan-Atlantic species distributions could result from several colonization pathways that connected different regions of the Atlantic at different times (e.g. Arctic, Antarctic or Panamanian pathways). To test potential colonization pathways we quantified geographic variation in nuclear and mitochondrial markers from Atlantic samples ofNucula atacellana, a pan-Atlantic deep-sea protobranch bivalve, usingN. profundorumin the eastern central Pacific as an outgroup. We combined existing 16S data from North and South Atlantic populations ofN. atacellanawith new sequences of 16S, COI, and an intron of calmodulin from those populations, and newly sampled populations near Iceland. Population genetic analyses indicated a subtropical expansion via the Central American Seaway. We found no evidence for Transarctic migration to the Atlantic inN. atacellana, which suggests that colonization pathways may differ significantly between shallow- and deep-water fauna.

A molecular phylogeny of Protobranchia, the subclass of bivalve mollusks sister to the remaining Bivalvia, has long proven elusive, because many constituent lineages are deep-sea endemics, which creates methodological challenges for collecting and preserving genetic material. We obtained 74 representatives of all 12 extant protobranch families and investigated the internal phylogeny of this group using sequence data from five molecular loci (16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase subunit I, and histone H3). Model-based and dynamic homology parsimony approaches to phylogenetic reconstruction unanimously supported four major clades of Protobranchia, irrespective of treatment of hypervariable regions in the nuclear ribosomal genes 18S rRNA and 28S rRNA. These four clades correspond to the superfamilies Nuculoidea (excluding Sareptidae), Nuculanoidea (including Sareptidae), Solemyoidea, and Manzanelloidea. Salient aspects of the phylogeny include (1) support for the placement of the family Sareptidae with Nuculanoidea; (2) the non-monophyly of the order Solemyida (Solemyidae + Nucinellidae); (3) and the non-monophyly of most nuculoid and nuculanoid genera and families. In light of this first family-level phylogeny of Protobranchia, we present a revised classification of the group. Estimation of divergence times in concert with analyses of diversification rates demonstrate the signature of the end-Permian mass extinction in the phylogeny of extant protobranchs.

We describe PCR primers and amplification protocols developed to obtain introns from conserved nuclear genes in deep-sea protobranch bivalves. Because almost no sequence data for protobranchs are publically available, mollusk and other protostome sequences from GenBank were used to design degenerate primers, making these loci potentially useful in other invertebrate taxa. Amplification and sequencing success varied across the test group of 30 species, and we present five loci spanning this range of outcomes. Intron presence in the targeted regions also varied across genes and species, often within single genera; for instance, the calmodulin and β-tubulin loci contained introns with high frequency, whereas the triose phosphate isomerase locus never contained an intron. In introns for which we were able to obtain preliminary estimates of polymorphism levels in single species, polymorphism was greater than traditional mitochondrial loci. These markers will greatly increase the ability to assess population structure in the ecologically important protobranchs, and may prove useful in other taxa as well.

The physical orientation of rocky substrate profoundly affects subtidal marine commu- nities of sessile organisms. Anecdotal descriptions of dramatic differences between communities on rock walls and adjacent horizontal rocky bottoms abound in the literature and are common knowl- edge among scuba divers, yet these differences have rarely been quantified by ecologists. We sam- pled rock walls and adjacent horizontal rock bottoms at 8 subtidal sites across the Gulf of Maine. Spe- cies richness and abundance, in percent cover, of sessile invertebrates on vertical walls averaged 4 times higher than that on horizontal platforms, whereas abundance of macroalgae on horizontal rock was about 3 times that on vertical walls. Both macroalgae and sessile invertebrates were less abundant, particularly on horizontal surfaces, at sites with high sea urchin densities. The consistency of sessile invertebrate domination of vertical walls versus macroalgal domination of horizontal rock, combined with previous experimental results, suggests that competition for space between autotro- phic and heterotrophic organisms drives this pattern. The partitioning of autotrophs (macroalgae) and heterotrophs (sessile invertebrates) between horizontal and vertical surfaces respectively implies that topographic heterogeneity plays an important role in the structure, composition and function of rocky subtidal ecosystems.

The deep sea is a vast and essentially continuous environment with few obvious barriers to gene flow. How populations diverge and new species form in this remote ecosystem is poorly understood. Phylogeographical analyses have begun to provide some insight into evolutionary processes at bathyal depths ( 3000 m). Here, we quantify geographical and bathymetric patterns of genetic variation (16S rRNA mitochondrial gene) in the protobranch bivalve Ledella ultima, which is one of the most abundant abyssal protobranchs in the Atlantic with a broad bathymetric and geographical distribution. We found virtually no genetic divergence within basins and only modest divergence among eight Atlantic basins. Levels of population divergence among basins were related to geographical distance and were greater in the South Atlantic than in the North Atlantic. Ocean-wide patterns of genetic variation indicate basin-wide divergence that exceeds what others have found for abyssal organisms, but considerably less than bathyal protobranchs across similar geographical scales. Populations on either side of the Mid-Atlantic Ridge in the North Atlantic differed, suggesting the Ridge might impede gene flow at abyssal depths. Our results indicate that abyssal populations might be quite large (cosmopolitan), exhibit only modest genetic structure and probably provide little potential for the formation of new species.

Dramatic shifts in community composition occur between vertical and horizontal rocky surfaces in subtidal environments worldwide, yet the forces mediating this transition are poorly understood. Vertical rock walls are often covered by lush, diverse communities of sessile suspension-feeding invertebrates, while adjacent horizontal substrates are dominated by algae, or corals in the tropics. Multiple factors, including light, sedimentation, water flow, and predation have been proposed to explain this pattern, but experimental tests of these hypotheses are lacking. We manipulated light level and predation to test whether variation in these mechanisms could be responsible for the shift in composition of sessile communities between vertical and horizontal surfaces in the rocky subtidal Gulf of Maine. Shaded horizontally oriented granite plots were dominated by invertebrates (e.g., ascidians, barnacles, bryozoans) after 25 months. Unshaded plots were dominated by macroalgae, which was virtually absent in shaded plots. Exclusion of grazers with cages had no effect on percent cover of invertebrates or algae. Preferential settlement of invertebrate larvae to shaded plots, due to larval behaviors such as negative phototaxis, did not seem to play a role. Shading likely affects post-settlement mortality of invertebrates by alleviating competition for space with algae, although greater abundance of micropredators in algal- dominated communities could also be important. Communities on shaded plots lacked many taxa present on natural wall communities, likely due to greater disturbance on horizontal substrates and/or lack of sufficient time for colonization of these taxa. These results suggest that light plays a key role in controlling the structure, composition, and function of shallow subtidal communities.

The relationship between surface production and benthic standing stock is fundamental to understanding biogeography in the deep sea. While much has been learned about the complex oceanographic processes involved in energy transfer to the benthos on local scales, the correspondence of overhead production to benthic community structure on regional scales remains poorly characterized. We compiled a database on the biomass and abundance of deep-sea macrobenthos in the western North Atlantic collected from 1961 to 1985. Using SeaWiFS satellite color imagery, we calculated POC from surface chlorophyll a concentrations (from 1997 to 2001), and estimated POC flux to the seafloor by using the empirically derived Pace et al. [1987. Primary production, new production and vertical flux in the eastern Pacific Ocean. Nature 325, 803–804] algorithm. The standing stock and surface production data are not concurrent, but their basic geographic trends at these very large spatial scales appear to be relatively stable over the time scales of measurement. Estimated POC flux at depth accounts for 62–67% of the variance ðPo0:0001Þ in benthic standing stock, suggesting that macroecological studies of the relationship between satellite-derived surface production and deep-sea community structure may be possible. r 2007 Elsevier Ltd. All rights reserved.

The deep-sea soft-sediment environment hosts a diverse and highly endemic fauna of uncertain origin. We know little about how this fauna evolved because geographic patterns of genetic variation, the essential information for inferring patterns of population differentiation and speciation are poorly understood. Using formalin-fixed specimens from archival collections, we quantify patterns of genetic variation in the protobranch bivalve Deminucula atacellana, a species widespread throughout the Atlantic Ocean at bathyal and abyssal depths. Samples were taken from 18 localities in the North American, West European and Argentine basins. A hypervariable region of mitochondrial 16S rDNA was amplified by polymerase chain reaction (PCR) and sequenced from 130 individuals revealing 21 haplotypes. Except for several important exceptions, haplotypes are unique to each basin. Overall gene diversity is high ( h = 0.73) with pronounced population structure (Φ ST = 0.877) and highly significant geographic associations ( P < 0.0001). Sequences cluster into four major clades corresponding to differences in geography and depth. Genetic divergence was much greater among populations at different depths within the same basin, than among those at similar depths but separated by thousands of kilometres. Isolation by distance probably explains much of the interbasin variation. Depth-related divergence may reflect historical patterns of colonization or strong environmental selective gradients. Broadly distributed deep-sea organisms can possess highly genetically divergent populations, despite the lack of any morphological divergence.

Geometric constraints represent a class of null models that describe how species diversity may vary between hard boundaries that limit geographic distributions. Recent studies have suggested that a number of large scale biogeographic patterns of diversity (e.g. latitude, altitude, depth) may reflect boundary constraints. However, few studies have rigorously tested the degree to which mid-domain null predictions match empirical patterns or how sensitive the null models are to various assumptions. We explore how variation in the assumptions of these models alter null depth ranges and consequently bathymetric variation in diversity, and test the extent to which bathymetric patterns of species diversity in deep sea gastropods, bivalves, and polychaetes match null predictions based on geometric constraints. Range–size distributions and geographic patterns of diversity produced by these null models are sensitive to the relative position of the hard boundaries, the specific algorithms used to generate range sizes, and whether species are continuously or patchily distributed between range end points. How well empirical patterns support null expectations is highly dependent on these assumptions. Bathymetric patterns of species diversity for gastropods, bivalves and polychaetes differ substantially from null expectations suggesting that geometric constraints do not account for diversity–depth patterns in the deep sea benthos.

With the development of new methods to sequence DNA from preserved organisms, existing archival collections can be used to document the population genetic structure of deep-sea species. This has made possible the first direct inferences about patterns of evolutionary diversification in the soft-sediment macrofauna. Here we report protocols and success rates for amplifying and sequencing regions of the mitochondrial 16S rDNA, Cytochrome oxidase I (COI), and Cytochrome b (cytb) genes from formalin-fixed protobranch bivalves and gastropods, major components of the deep-sea benthos. DNA was extracted from 1532 individuals of 12 common bathyal and abyssal species that had been fixed in formalin and preserved in alcohol for up to 36 years. DNA was also extracted from 53 individuals that were dried upon collection, some of which were collected more than 100 years ago. The overall success rate for amplification by PCR was 44%, but this varied considerably among species, stations, and cruises. When DNA amplified, sequencing success was generally high, averaging 85% and ranging from 19% to 100%. The reliability of amplification and sequencing depend strongly on how samples are treated during collection and storage. Amplification success was similar among samples collected from the same station and samples collected on the same cruise. We provide recommendations on strategies for primer design, PCR, and sample selection to improve success rates for genetic analysis of preserved deep-sea organisms. The success rates from different collections, sampling stations, and cruises provide important guidance for selecting material for future genetic work on deep-sea collections examined here.

▪ Abstract Most of our knowledge of biodiversity and its causes in the deep-sea benthos derives from regional-scale sampling studies of the macrofauna. Improved sampling methods and the expansion of investigations into a wide variety of habitats have revolutionized our understanding of the deep sea. Local species diversity shows clear geographic variation on spatial scales of 100–1000 km. Recent sampling programs have revealed unexpected complexity in community structure at the landscape level that is associated with large-scale oceanographic processes and their environmental consequences. We review the relationships between variation in local species diversity and the regional-scale phenomena of boundary constraints, gradients of productivity, sediment heterogeneity, oxygen availability, hydrodynamic regimes, and catastrophic physical disturbance. We present a conceptual model of how these interdependent environmental factors shape regional-scale variation in local diversity. Local communities in the deep sea may be composed of species that exist as metapopulations whose regional distribution depends on a balance among global-scale, landscape-scale, and small-scale dynamics. Environmental gradients may form geographic patterns of diversity by influencing local processes such as predation, resource partitioning, competitive exclusion, and facilitation that determine species coexistence. The measurement of deep-sea species diversity remains a vital issue in comparing geographic patterns and evaluating their potential causes. Recent assessments of diversity using species accumulation curves with randomly pooled samples confirm the often-disputed claim that the deep sea supports higher diversity than the continental shelf. However, more intensive quantitative sampling is required to fully characterize the diversity of deep-sea sediments, the most extensive habitat on Earth. Once considered to be constant, spatially uniform, and isolated, deep-sea sediments are now recognized as a dynamic, richly textured environment that is inextricably linked to the global biosphere. Regional studies of the last two decades provide the empirical background necessary to formulate and test specific hypotheses of causality by controlled sampling designs and experimental approaches.

We have developed cellular automaton models for two species competing in a patchy environment. We have modeled three common types of competition: facilitation (in which the winning species can colonize only after the losing species has arrived) inhibition (in which either species is able to prevent the other from colonizing) and tolerance (in which the species most tolerant of reduced resource levels wins). The state of a patch is defined by the presence or absence of each species. State transition probabilities are determined by rates of disturbance, competitive exclusion, and colonization. Colonization is restricted to neighboring patches. In all three models, disturbance permits regional persistence of species that are excluded by competition locally. Persistence, and hence diversity, is maximized at intermediate disturbance frequencies. If disturbance and dispersal rates are sufficiently high, the inferior competitor need not have a dispersal advantage to persist. Using a new method for measuring the spatial patterns of nominal data, we show that none of these competition models generates patchiness at equilibrium. In the inhibition model, however, transient patchiness decays very slowly. We compare the cellular automaton models to the corresponding mean-field patch-occupancy models, in which colonization is not restricted to neighboring patches and depends on spatially averaged species frequencies. The patch-occupancy model does an excellent job of predicting the equilibrium frequencies of the species and the conditions required for coexistence, but not of predicting transient behavior.

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) 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.

The deep-sea soft-sediment environment hosts a diverse and highly endemic fauna of uncertain origin. We know little about how this fauna evolved because geographic patterns of genetic variation, the essential information for inferring patterns of population differentiation and speciation are poorly understood. Using formalin-fixed specimens from archival collections, we quantify patterns of genetic variation in the protobranch bivalve Deminucula atacellana, a species widespread throughout the Atlantic Ocean at bathyal and abyssal depths. Samples were taken from 18 localities in the North American, West European and Argentine basins. A hypervariable region of mitochondrial 16S rDNA was amplified by polymerase chain reaction (PCR) and sequenced from 130 individuals revealing 21 haplotypes. Except for several important exceptions, haplotypes are unique to each basin. Overall gene diversity is high (h = 0.73) with pronounced population structure (Phi(ST) = 0.877) and highly significant geographic associations (P0.0001). Sequences cluster into four major clades corresponding to differences in geography and depth. Genetic divergence was much greater among populations at different depths within the same basin, than among those at similar depths but separated by thousands of kilometres. Isolation by distance probably explains much of the interbasin variation. Depth-related divergence may reflect historical patterns of colonization or strong environmental selective gradients. Broadly distributed deep-sea organisms can possess highly genetically divergent populations, despite the lack of any morphological divergence.

The deep sea is the largest ecosystem on Earth. Recent exploration has revealed that it supports a highly diverse and endemic benthic invertebrate fauna, yet the evolutionary processes that generate this remarkable species richness are virtually unknown. Environmental heterogeneity, topographic complexity, and morphological divergence all tend to decrease with depth, suggesting that the potential for population differentiation may decrease with depth. To test this hypothesis, we use mitochondrial DNA (16S rRNA gene) to examine patterns of population differentiation in four species of protobranch bivalves (Nuculoma similis, Deminucula atacellana, Malletia abyssorum, and Ledella ultima) distributed along a depth gradient in the western North Atlantic. We sequenced 268 individuals from formalin-fixed samples and found 45 haplotypes. The level of sequence divergence among haplotypes within species was similar, but shifted from between populations at bathyal depths to within populations at abyssal depths. Levels of population structure as measured by phiST were considerably greater in the upper bathyal species (N. similis = 0.755 and D. atacellana = 0.931; 530-3834 m) than in the lower bathyal/abyssal species (M. abyssorum = 0.071 and L. ultima = 0.045; 2864-4970 m). Pairwise genetic distances among the samples within each species also decreased with depth. Population trees (UPGMA) based on modified coancestry coefficients and nested clade analysis both indicated strong population-level divergence in the two upper bathyal species but little for the deeper species. The population genetic structure in these protobranch bivalves parallels depth-related morphological divergence observed in deep-sea gastropods. The higher level of genetic and morphological divergence, coupled with the strong biotic and abiotic heterogeneity at bathyal depths, suggests this region may be an active area of species formation. We suggest that the steep, topographically complex, and dynamic bathyal zone, which stretches as a narrow band along continental margins, plays a more important role in the evolutionary radiation of the deep-sea fauna than the much more extensive abyss.

The deep sea is the largest ecosystem on Earth. Recent exploration has revealed that it supports a highly diverse and endemic benthic invertebrate fauna, yet the evolutionary processes that generate this remarkable species richness are virtually unknown. Environmental heterogeneity, topographic complexity, and morphological divergence all tend to decrease with depth, suggesting that the potential for population differentiation may decrease with depth. To test this hypothesis, we use mitochondrial DNA (16S rRNA gene) to examine patterns of population differentiation in four species of protobranch bivalves (Nuculoma similis, Deminucula atacellana, Malletia abyssorum, and Ledella ultima) distributed along a depth gradient in the western North Atlantic. We sequenced 268 individuals from formalin- fixed samples and found 45 haplotypes. The level of sequence divergence among haplotypes within species was similar, but shifted from between populations at bathyal depths to within populations at abyssal depths. Levels of population structure as measured by FST were considerably greater in the upper bathyal species (N. similis 5 0.755 and D. atacellana 5 0.931; 530-3834 m) than in the lower bathyal/abyssal species (M. abyssorum 5 0.071 and L. ultima 5 0.045; 2864-4970 m). Pairwise genetic distances among the samples within each species also decreased with depth. Population trees (UPGMA) based on modified coancestry coefficients and nested clade analysis both indicated strong population-level divergence in the two upper bathyal species but little for the deeper species. The population genetic structure in these protobranch bivalves parallels depth-related morphological divergence observed in deep-sea gas- tropods. The higher level of genetic and morphological divergence, coupled with the strong biotic and abiotic het- erogeneity at bathyal depths, suggests this region may be an active area of species formation. We suggest that the steep, topographically complex, and dynamic bathyal zone, which stretches as a narrow band along continental margins, plays a more important role in the evolutionary radiation of the deep-sea fauna than the much more extensive abyss.

Demographic and life history characteristics of the intertidal whelk Nucella lapillus were measured across a wave—exposure gradient to quantify the variation in, and identify the ecological forces shaping, each trait. Growth rates, survivorship, size and age at maturity, fecundity, and per—offspring parental investment were estimated from marked snails during a 3—yr period. Growth rates as indicated by changes in shell length, total mass, shell mass, and body mass varied among populations from different exposure regimes. Relative to more protected sites, snails from exposed shores grew more slowly and terminated growth at a smaller size. Age at maturity did not differ between whelks from high— and low—wave—energy populations, but those on the exposed shore matured at a smaller size. Mortality rates increased with wave energy. Size—specific mortality rates indicated that the higher mortality on wave—swept shores reflected decreased survivorship of large (@> 15 mm) adults relative to similar sized individuals on more protected shores. Exposed shore snails deposited twice as many egg capsules with twice as many hatchlings emerging from each capsule. Although the hatchlings were °25% smaller, four times as many were produced, suggesting that reproductive effort was considerably greater on exposed coasts, offsetting the higher mortality rates. The ecological forces (energetic, physiological, abiotic, etc.) potentially responsible for the life history variation among populations from different wave—exposure regimes are discussed.