Towards a ‘Sea‐Level Sensitive’ dynamic model: impact of island ontogeny and glacio‐eustasy on global patterns of marine island biogeography

A synthetic model is presented to enlarge the evolutionary framework of the General Dynamic Model (GDM) and the Glacial Sensitive Model (GSM) of oceanic island biogeography from the terrestrial to the marine realm. The proposed ‘Sea‐Level Sensitive’ dynamic model (SLS) of marine island biogeography integrates historical and ecological biogeography with patterns of glacio‐eustasy, merging concepts from areas as diverse as taxonomy, biogeography, marine biology, volcanology, sedimentology, stratigraphy, palaeontology, geochronology and geomorphology. Fundamental to the SLS model is the dynamic variation of the littoral area of volcanic oceanic islands (defined as the area between the intertidal and the 50‐m isobath) in response to sea‐level oscillations driven by glacial–interglacial cycles. The following questions are considered by means of this revision: (i) what was the impact of (global) glacio‐eustatic sea‐level oscillations, particularly those of the Pleistocene glacial–interglacial episodes, on the littoral marine fauna and flora of volcanic oceanic islands? (ii) What are the main factors that explain the present littoral marine biodiversity on volcanic oceanic islands? (iii) How can differences in historical and ecological biogeography be reconciled, from a marine point of view? These questions are addressed by compiling the bathymetry of 11 Atlantic archipelagos/islands to obtain quantitative data regarding changes in the littoral area based on Pleistocene sea‐level oscillations, from 150 thousand years ago (ka) to the present. Within the framework of a model sensitive to changing sea levels, we discuss the principal factors affecting the geographical range of marine species; the relationships between modes of larval development, dispersal strategies and geographical range; the relationships between times of speciation, modes of larval development, ecological zonation and geographical range; the influence of sea‐surface temperatures and latitude on littoral marine species diversity; the effect of eustatic sea‐level changes and their impact on the littoral marine biota; island marine species–area relationships; and finally, the physical effects of island ontogeny and its associated submarine topography and marine substrate on littoral biota. Based on the SLS dynamic model, we offer a number of predictions for tropical, subtropical and temperate volcanic oceanic islands on how rates of immigration, colonization, in‐situ speciation, local disappearance, and extinction interact and affect the marine biodiversity around islands during glacials and interglacials, thus allowing future testing of the theory.     

Phylogenetic measures reveal eco-evolutionary drivers of biodiversity along a depth gradient

Energy and environmental stability are positively correlated with species richness along broad-scale spatial gradients in terrestrial ecosystems, so their relative importance in generating and preserving diversity cannot be readily disentangled. This study seeks to exploit the negative correlation between energy and stability along the oceanic depth gradient to better understand their relative contribution in shaping broadscale biodiversity patterns. We develop a conceptual framework by simulating speciation and extinction along energy and stability gradients to generate expected patterns of biodiversity for a suite of complementary phylogenetic diversity metrics. Using a time-calibrated molecular phylogeny for New Zealand marine ray-finned fishes and a replicated community ecological sampling design, we then modelled these metrics along large-scale depth and latitude gradients. Our results indicate that energy-rich shallow waters may be an engine of diversity for percomorphs, but also suggest that recent speciation occurs in ancient fish lineages in the deep sea, hence questioning the role of energy as a key driver of speciation. Despite potentially facing high extinction early in their evolution, ancient phylogenetic lineages specialized for the deep-sea were likely preserved by environmental stability during the Cenozoic. Furthermore, intermediate depths might be a ‘museum’ (or zone of overlap) for distinct lineages that occur predominantly in either shallow or deep-sea waters. These intermediate depths (500–900 m) may form a ‘phylogenetic diversity bank’, perhaps providing a refuge during ancient (Mesozoic) extreme anoxic events affecting the deep sea and more recent (Pliocene–Pleistocene) climatic events occurring in shallow ecosystems. Finally, the phylogenetic structures observed in fish communities at intermediate depths suggest other processes might restrict the co-occurrence of closely related species. Overall, by combining a conceptual framework with models of empirical phylogenetic diversity patterns, our study paves the way for understanding the determinants of biodiversity across the largest habitat on earth.     

Population structure,dispersal and colonization history of the garden bumblebee <Emphasis Type="Italic">Bombus hortorum</Emphasis> in the Western Isles of Scotland

New methods of analysing genetic data provide powerful tools for quantifying dispersal patterns and reconstructing population histories. Here we examine the population structure of the bumblebee Bombus hortorum in a model island system, the Western Isles of Scotland, using microsatellite markers. Following declines in other species, B. hortorum is the only remaining long-tongued bumblebee species found in much of Europe, and thus it is of particular ecological importance. Our data suggest that populations of B. hortorum in western Scotland exist as distinct genetic clusters occupying groups of nearby islands. Population structuring was higher than for other bumblebee species which have previously been studied in this same island group (F_st = 0.16). Populations showed significant isolation by distance. This relationship was greatly improved by using circuit theory to allow dispersal rates to differ over different landscape features; as we would predict, sea appears to provide far higher resistance to dispersal than land. Incorporating bathymetry data improved the fit of the model further; populations separated by shallow seas are more genetically similar than those separated by deeper seas. We argue that this probably reflects events following the last ice age when the islands were first colonized by this bee species (8,500–5,000 ybp), when the sea levels were lower and islands separated by shallow channels would have been joined. In the absence of significant gene flow these genetic clusters appear to have since diverged over the following 5,000 years and arguably may now represent locally adapted races, some occurring on single islands.     

An approach to bioassess pelagic ciliate biodiversity at different taxonomic resolutions in response to various habitats in the Amundsen Sea (Antarctica)

The rapid melting of glaciers and loss of sea ice will result in changes in habitat conditions that may drive substantial changes in biodiversity. In order to bioassess the changing polar ecosystem and evaluate biological conservation, pelagic ciliate communities at different taxonomic resolutions were studied at five habitats in the Amundsen Sea during the austral summer from December 2010 to January 2011. Distinctive spatial patterns were observed in the communities among the five habitats (oceanic areas, transitional areas, polynyas, edges of glaciers, and edges of sea ice) in response to environmental variability (e.g., temperature, salinity, chlorophyll a, and nutrients). The distributions in the numbers of different taxonomic levels and of three biodiversity indices (Shannon-Wiener H′, Pielou’s J′, and Margalef D) also revealed clear spatial variability with the maximum mean species number and indices in the polynya and maximum genus and family numbers in the transitional area. The presence/absence of data at taxonomic resolutions up to the family level provided sufficient information to evaluate the ecological patterns of pelagic ciliate communities and could accurately reflect habitat variations. The k-dominance curves illustrated clearly that maximum diversity was presented in the polynya at the species level and in the transitional area at the genus and family level. We suggest that the diversity at higher taxonomic resolutions should be considered more in future monitoring. Our findings provide basic data and an approach toward answering important questions about biological conservation, especially the biodiversity at various taxonomic resolutions in response to the increasing climate changes in polar ecosystems.     

Interaction between coastal and oceanic ecosystems of the Western and Central Pacific Ocean through predator-prey relationship studies

The Western and Central Pacific Ocean sustains the highest tuna production in the world. This province is also characterized by many islands and a complex bathymetry that induces specific current circulation patterns with the potential to create a high degree of interaction between coastal and oceanic ecosystems. Based on a large dataset of oceanic predator stomach contents, our study used generalized linear models to explore the coastal-oceanic system interaction by analyzing predator-prey relationship. We show that reef organisms are a frequent prey of oceanic predators. Predator species such as albacore (Thunnus alalunga) and yellowfin tuna (Thunnus albacares) frequently consume reef prey with higher probability of consumption closer to land and in the western part of the Pacific Ocean. For surface-caught-predators consuming reef prey, this prey type represents about one third of the diet of predators smaller than 50 cm. The proportion decreases with increasing fish size. For predators caught at depth and consuming reef prey, the proportion varies with predator species but generally represents less than 10%. The annual consumption of reef prey by the yellowfin tuna population was estimated at 0.8 ± 0.40 CV million tonnes or 2.17 × 10(12)± 0.40 CV individuals. This represents 6.1% ± 0.17 CV in weight of their diet. Our analyses identify some of the patterns of coastal-oceanic ecosystem interactions at a large scale and provides an estimate of annual consumption of reef prey by oceanic predators.     

Ecological guidelines for designing networks of marine reserves in the unique biophysical environment of the Gulf of California

No-take marine reserves can be powerful management tools, but only if they are well designed and effectively managed. We review how ecological guidelines for improving marine reserve design can be adapted based on an area’s unique evolutionary, oceanic, and ecological characteristics in the Gulf of California, Mexico. We provide ecological guidelines to maximize benefits for fisheries management, biodiversity conservation and climate change adaptation. These guidelines include: representing 30% of each major habitat (and multiple examples of each) in marine reserves within each of three biogeographic subregions; protecting critical areas in the life cycle of focal species (spawning and nursery areas) and sites with unique biodiversity; and establishing reserves in areas where local threats can be managed effectively. Given that strong, asymmetric oceanic currents reverse direction twice a year, to maximize connectivity on an ecological time scale, reserves should be spaced less than 50–200 km apart depending on the planktonic larval duration of target species; and reserves should be located upstream of fishing sites, taking the reproductive timing of focal species in consideration. Reserves should be established for the long term, preferably permanently, since full recovery of all fisheries species is likely to take?>?25 years. Reserve size should be based on movement patterns of focal species, although marine reserves?>?10 km long are likely to protect?~?80% of fish species. Since climate change will affect species’ geographic range, larval duration, growth, reproduction, abundance, and distribution of key recruitment habitats, these guidelines may require further modifications to maintain ecosystem function in the future.     

The developmental biogeography of hawksbill sea turtles in the North Pacific

High seas oceanic ecosystems are considered important habitat for juvenile sea turtles, yet much remains cryptic about this important life‐history period. Recent progress on climate and fishery impacts in these so‐called lost years is promising, but the developmental biogeography of hawksbill sea turtles (Eretmochelys imbricata) has not been widely described in the Pacific Ocean. This knowledge gap limits the effectiveness of conservation management for this globally endangered species. We address this with 30 years of stranding observations, 20 years of bycatch records, and recent simulations of natal dispersal trajectories in the Hawaiian Archipelago. We synthesize the analyses of these data in the context of direct empirical observations, anecdotal sightings, and historical commercial harvests from the insular Pacific. We find hawksbills 0–4 years of age, measuring 8–34 cm straight carapace length, are found predominantly in the coastal pelagic waters of Hawaii. Unlike other species, we find no direct evidence of a prolonged presence in oceanic habitats, yet satellite tracks of passive drifters (simulating natal dispersal) and our small sample sizes suggest that an oceanic phase for hawksbills cannot be dismissed. Importantly, despite over 600 million hooks deployed and nearly 6000 turtle interactions, longline fisheries have never recorded a single hawksbill take. We address whether the patterns we observe are due to population size and gear selectivity. Although most sea turtle species demonstrate clear patterns of oceanic development, hawksbills in the North Pacific may by contrast occupy a variety of ecosystems including coastal pelagic waters and shallow reefs in remote atolls. This focuses attention on hazards in these ecosystems – entanglement and ingestion of marine debris – and perhaps away from longline bycatch and decadal climate regimes that affect sea turtle development in oceanic regions.     

The roles of geological history and colonization abilities in genetic differentiation between mammalian populations in the Philippine archipelago

Aim To test hypotheses that: (1) late Pleistocene low sea‐level shorelines (rather than current shorelines) define patterns of genetic variation among mammals on oceanic Philippine islands; (2) species‐specific ecological attributes, especially forest fidelity and vagility, determine the extent to which common genetic patterns are exhibited among a set of species; (3) populations show reduced within‐population variation on small, isolated oceanic islands; (4) populations tend to be most highly differentiated on small, isolated islands; and (5) to assess the extent to which patterns of genetic differentiation among multiple species are determined by interactions of ecological traits and geological/geographic conditions. Location The Philippine Islands, a large group of oceanic islands in Southeast (SE) Asia with unusually high levels of endemism among mammals. Methods Starch‐gel electrophoresis of protein allozymes of six species of small fruit bats (Chiroptera, Pteropodidae) and one rodent (Rodentia, Muridae). Results Genetic distances between populations within all species are not correlated with distances between present‐day shorelines, but are positively correlated with distances between shorelines during the last Pleistocene period of low sea level; relatively little intraspecific variation was found within these ‘Pleistocene islands’. Island area and isolation of oceanic populations have only slight effects on standing genetic variation within populations, but populations on some isolated islands have heightened levels of genetic differentiation, and reduced levels of gene flow, relative to other islands. Species associated with disturbed habitat (all of which fly readily across open habitats) show more genetic variation within populations than species associated with primary rain forest (all of which avoid flying out from beneath forest canopy). Species associated with disturbed habitats, which tend to be widely distributed in SE Asia, also show higher rates of gene flow and less differentiation between populations than species associated with rain forest, which tend to be Philippine endemic species. One rain forest bat has levels of gene flow and heterozygosity similar to the forest‐living rodent in our study. Main conclusions The maximum limits of Philippine islands that were reached during Pleistocene periods of low sea level define areas of relative genetic homogeneity, whereas even narrow sea channels between adjacent but permanently isolated oceanic islands are associated with most genetic variation within the species. Moreover, the distance between ‘Pleistocene islands’ is correlated with the extent of genetic distances within species. The structure of genetic variation is strongly influenced by the ecology of the species, predominantly as a result of their varying levels of vagility and ability to tolerate open (non‐forested) habitat. Readily available information on ecology (habitat association and vagility) and geological circumstances (presence or absence of Pleistocene land‐bridges between islands, and distance between oceanic islands during periods of low sea level) are combined to produce a simple predictive model of likely patterns of genetic differentiation (and hence speciation) among these mammals, and probably among other organisms, in oceanic archipelagos.     

Understanding how biodiversity unfolds through time under neutral theory

Theoretical predictions for biodiversity patterns are typically derived under the assumption that ecological systems have reached a dynamic equilibrium. Yet, there is increasing evidence that various aspects of ecological systems, including (but not limited to) species richness, are not at equilibrium. Here, we use simulations to analyse how biodiversity patterns unfold through time. In particular, we focus on the relative time required for various biodiversity patterns (macroecological or phylogenetic) to reach equilibrium. We simulate spatially explicit metacommunities according to the Neutral Theory of Biodiversity (NTB) under three modes of speciation, which differ in how evenly a parent species is split between its two daughter species. We find that species richness stabilizes first, followed by species area relationships (SAR) and finally species abundance distributions (SAD). The difference in timing of equilibrium between these different macroecological patterns is the largest when the split of individuals between sibling species at speciation is the most uneven. Phylogenetic patterns of biodiversity take even longer to stabilize (tens to hundreds of times longer than species richness) so that equilibrium predictions from neutral theory for these patterns are unlikely to be relevant. Our results suggest that it may be unwise to assume that biodiversity patterns are at equilibrium and provide a first step in studying how these patterns unfold through time.     

Cobble community DNA as a tool to monitor patterns of biodiversity within kelp forest ecosystems

Kelp forest ecosystems dominate 150,000 km of global temperate coastline, rivalling the coastal occurrence of coral reefs. Despite the astounding biological diversity and productive ecological communities associated with kelp forests, patterns of species richness and composition are difficult to monitor and compare. Crustose coralline algae are a critically important substrate for propagule settlement for a range of kelp forest species. Coralline‐covered cobbles are home to hundreds of species of benthic animals and algae and form a replicable unit for ecological assays. Here, we use DNA metabarcoding of bulk DNA extracts sampled from cobbles to explore patterns of species diversity in kelp forests of the central California coast. The data from 97 cobbles within kelp forest ecosystems at three sites in Central California show the presence of 752 molecular operational taxonomic units (MOTUs) and 53 MOTUs assigned up to the species level with >95% similarity to current databases. We are able to detect spatial patterns of important management targets such as abalone recruits, and localized abundance of sea stars in 2012. Comparison of classic ecological surveys of these sites reveals large differences in species targets for these two approaches. In order to make such comparisons more quantitative, we use Presence/Absence Metabarcoding, using the fraction of replicate cobbles showing a species as a measure of its local abundance. This approach provides a fast and repeatable survey method that can be applied for biodiversity assessments across systems to shed light on the impact of different ecological disturbances and the role played by marine protected areas.     

Deglaciation explains bat extinction in the Caribbean

Ecological factors such as changing climate on land and interspecific competition have been debated as possible causes of postglacial Caribbean extinction. These hypotheses, however, have not been tested against a null model of climate‐driven postglacial area loss. Here, we use a new Quaternary mammal database and deep‐sea bathymetry to estimate species–area relationships (SARs) at present and during the Last Glacial Maximum (LGM) for bats of the Caribbean, and to model species loss as a function of area loss from rising sea level. Island area was a significant predictor of species richness in the Bahamas, Greater Antilles, and Lesser Antilles at all time periods, except for the Lesser Antilles during the LGM. Parameters of LGM and current SARs were similar in the Bahamas and Greater Antilles, but not the Lesser Antilles, which had fewer estimated species during the LGM than expected given their size. Estimated postglacial species losses in the Bahamas and Greater Antilles were largely explained by inferred area loss from rising sea level in the Holocene. However, there were more species in the Bahamas at present, and fewer species in the smaller Greater Antilles, than expected given island size and the end‐Pleistocene/early Holocene SARs. Poor fossil sampling and ecological factors may explain these departures from the null. Our analyses illustrate the importance of changes in area in explaining patterns of species richness through time and emphasize the role of the SAR as a null hypothesis in explorations of the impact of novel ecological interactions on extinction.     

Different responses of avian feeding guilds to spatial and environmental factors across an elevation gradient in the central Himalaya

Although elevational patterns of species richness have been well documented, how the drivers of richness gradients vary across ecological guilds has rarely been reported. Here, we examined the effects of spatial factors (area and mid‐domain effect; MDE) and environmental factors, including metrics of climate, productivity, and plant species richness on the richness of breeding birds across different ecological guilds defined by diet and foraging strategy. We surveyed 12 elevation bands at intervals of 300 m between 1,800 and 5,400 m a.s.l using line‐transect methods throughout the wet season in the central Himalaya, China. Multiple regression models and hierarchical partitioning were used to assess the relative importance of spatial and environmental factors on overall bird richness and guild richness (i.e., the richness of species within each guild). Our results showed that richness for all birds and most guilds displayed hump‐shaped elevational trends, which peaked at an elevation of 3,300–3,600 m, although richness of ground‐feeding birds peaked at a higher elevation band (4,200–4,500 m). The Normalized Difference Vegetation Index (NDVI)—an index of primary productivity—and habitat heterogeneity were important factors in explaining overall bird richness as well as that of insectivores and omnivores, with geometric constraints (i.e., the MDE) of secondary importance. Granivore richness was not related to primary production but rather to open habitats (granivores were negatively influenced by habitat heterogeneity), where seeds might be abundant. Our findings provide direct evidence that the richness–environment relationship is often guild‐specific. Taken together, our study highlights the importance of considering how the effects of environmental and spatial factors on patterns of species richness may differ across ecological guilds, potentially leading to a deeper understanding of elevational diversity gradients and their implications for biodiversity conservation.     

Niche breadth and biodiversity change derived from marine Amphipoda species off Iceland

Understanding the ecological requirements and thresholds of individual species is crucial to better predict potential outcomes of climate change on species distribution. In particular, species optima and lower and upper limits along resource gradients require attention. Based on Huisman‐Olff‐Fresco (HOF) models, we determined species‐specific responses along gradients of nine environmental parameters including depth in order to estimate niche attributes of 30 deep‐sea benthic amphipods occurring around Iceland. We, furthermore, examined the relationships between niche breadth, occupancy, and geographic range assuming that species with a wider niche are spatially more widely dispersed and vice versa. Overall, our results reveal that species react very differently to environmental gradients, which is independent of the family affiliation of the respective species. We could infer a strong relationship between occupancy and geographic range and also relate this to differences in niche breadth; that is specialist species with a narrow niche had a more limited distribution and may thus be more threatened by changing environmental conditions than generalist species, which are more widespread. Given the preponderance of rare species in the deep sea, this implies that many species could be at risk. However, this must be carefully weighed against geographical data gaps in this area, given that many deep‐sea areas are severely undersampled and the true distribution of most species is unknown. After all, our results underline that an accurate taxonomic classification is of crucial importance, without which ecological niche properties cannot be determined and which is hence fundamental for the assessment and understanding of changes in biodiversity in the face of increasing human perturbations.     

Spatial patterns and scaling behaviors of Steller sea lion (Eumetopias jubatus) distributions and their environment

Fractal geometry and other multi-scale analyses have become popular tools for investigating spatial patterns of animal distributions in heterogeneous environments. In theory, changes in patterns of animal distributions with changes in scale reflect transitions between the controlling influences of one environmental factor or process over another. In an effort to find linkages between Steller sea lions (Eumetopias jubatus) and their environment, the objective of this study was to determine if the spatial distribution of Steller sea lions at sea displayed similar scaling properties to the variation of two environmental features, including bathymetry and sea surface temperature (SST). Additionally, distributions of Steller sea lion point patterns were examined with respect to measurements of bathymetric complexity. From February 2000 to May 2004, satellite transmitters were deployed on 10 groups of juvenile Steller sea lions (n=52) at eight different locations within the Aleutian Islands and Gulf of Alaska. Indices of fractal dimension were calculated for each group of sea lions using a unit square box-counting method, whereas indices of bathymetry and SST patchiness were derived by conducting a variance ratio analysis over the same scales. Distributions of Steller sea lions at sea displayed self-similar fractal patterns, suggesting that individuals were distributed in a continuous hierarchical set of clumps within clumps across scales, and foraging behavior was likely influenced by a scale invariant mechanism. Patterns of bathymetric variability also were self-similar, whereas patterns of SST variability were scale dependent and failed to retain self-similar spatial structure at larger scales. These results indicate that the distributions of Steller sea lions at sea were more influenced by bathymetry than SST at the scales examined, but scale-dependent patterns in the distribution of Steller sea lions at sea or linkages with SST may have been apparent if analyses were conducted at finer spatial scales.     

Using biological traits to explain ladybird distribution patterns

Aim  Determining to what extent differing distribution patterns are governed by species’ life‐history and resource‐use traits may lead to an improved understanding of the impacts of environmental change on biodiversity. We investigated the extent to which traits can explain distribution patterns in the ladybird fauna (Coleoptera: Coccinellidae) of Great Britain. Location  The British mainland and inshore islands (Anglesey, the Isle of Wight and the Inner Hebrides). Methods  The distributions of 26 ladybird species resident in Britain were characterized in terms of their range size (from 2661 10‐km grid squares across Britain) and proportional range fill (at 10‐ and 50‐km scales). These were assessed relative to five traits (body length, elytral colour pattern polymorphism, voltinism, habitat specificity and diet breadth). The role of phylogenetic autocorrelation was examined by comparing the results of phylogenetic and generalized least‐squares regressions. Results  Diet breadth was the only trait correlated with range size: species with broad diets had larger range sizes than dietary specialists. Range fill was sensitive to recording intensity (a per‐species measure of the mean number of records across occupied squares); models including both recording intensity and range size provided more explanatory power than models incorporating ecological traits alone. Main conclusions  Habitat specificity is often invoked to explain the distribution patterns of species, but here we found diet breadth to be the only ecological correlate of both range fill and range size. This highlights the importance of understanding predator–prey interactions when attempting to explain the distribution patterns of predatory species. Our results suggest that the diet breadth of predatory species is a better correlate of range size and fill than other measures, such as habitat specificity.     

Niche divergence by deep‐sea octocorals in the genus Callogorgia across the continental slope of the Gulf of Mexico

Environmental variables that are correlated with depth have been suggested to be among the major forces underlying speciation in the deep sea. This study incorporated phylogenetics and ecological niche models (ENM) to examine whether congeneric species of Callogorgia (Octocorallia: Primnoidae) occupy different ecological niches across the continental slope of the Gulf of Mexico (GoM) and whether this niche divergence could be important in the evolution of these closely related species. Callogorgia americana americana, Callogorgia americana delta and Callogorgia gracilis were documented at 13 sites in the GoM (250–1000 m) from specimen collections and extensive video observations. On a first order, these species were separated by depth, with C. gracilis occurring at the shallowest sites, C. a. americana at mid‐depths and C. a. delta at the deepest sites. Callogorgia a. delta was associated with areas of increased seep activity, whereas C. gracilis and C. a. americana were associated with narrow, yet warmer, temperature ranges and did not occur near cold seeps. ENM background and identity tests revealed little to no overlap in ecological niches between species. Temporal calibration of the phylogeny revealed the formation of the Isthmus of Panama was a vicariance event that may explain some of the patterns of speciation within this genus. These results elucidate the potential mechanisms for speciation in the deep sea, emphasizing both bathymetric speciation and vicariance events in the evolution of a genus across multiple regions.     

Patterns and controlling factors of species diversity in the Arctic Ocean

Aim The Arctic Ocean is one of the last near‐pristine regions on Earth, and, although human activities are expected to impact on Arctic ecosystems, we know very little about baseline patterns of Arctic Ocean biodiversity. This paper aims to describe Arctic Ocean‐wide patterns of benthic biodiversity and to explore factors related to the large‐scale species diversity patterns. Location Arctic Ocean. Methods We used large ostracode and foraminiferal datasets to describe the biodiversity patterns and applied comprehensive ecological modelling to test the degree to which these patterns are potentially governed by environmental factors, such as temperature, productivity, seasonality, ice cover and others. To test environmental control of the observed diversity patterns, subsets of samples for which all environmental parameters were available were analysed with multiple regression and model averaging. Results Well‐known negative latitudinal species diversity gradients (LSDGs) were found in metazoan Ostracoda, but the LSDGs were unimodal with an intermediate maximum with respect to latitude in protozoan foraminifera. Depth species diversity gradients were unimodal, with peaks in diversity shallower than those in other oceans. Our modelling results showed that several factors are significant predictors of diversity, but the significant predictors were different among shallow marine ostracodes, deep‐sea ostracodes and deep‐sea foraminifera. Main conclusions On the basis of these Arctic Ocean‐wide comprehensive datasets, we document large‐scale diversity patterns with respect to latitude and depth. Our modelling results suggest that the underlying mechanisms causing these species diversity patterns are unexpectedly complex. The environmental parameters of temperature, surface productivity, seasonality of productivity, salinity and ice cover can all play a role in shaping large‐scale diversity patterns, but their relative importance may depend on the ecological preferences of taxa and the oceanographic context of regions. These results suggest that a multiplicity of variables appear to be related to community structure in this system.     

Visual perception of light organ patterns in deep‐sea shrimps and implications for conspecific recognition

Darkness and low biomass make it challenging for animals to find and identify one another in the deep sea. While spatiotemporal variation in bioluminescence is thought to underlie mate recognition for some species, its role in conspecific recognition remains unclear. The deep‐sea shrimp genus, Sergestes sensu lato (s.l.), is one group that is characterized by species‐specific variation in light organ arrangement, providing us the opportunity to test whether organ variation permits recognition to the species level. To test this, we analyzed the visual capabilities of three species of Sergestes s.l. in order to (a) test for sexual dimorphism in eye‐to‐body size scaling relationships, (b) model the visual ranges (i.e., sighting distances) over which these shrimps can detect intraspecific bioluminescence, and (c) assess the maximum possible spatial resolution of the eyes of these shrimps to estimate their capacity to distinguish the light organs of each species. Our results showed that relative eye size scaled negatively with body length across species and without sexual dimorphism. Though the three species appear capable of detecting one another's bioluminescence over distances ranging from < 1 to ~6 m, their limited spatial resolution suggests they cannot resolve light organ variation for the purpose of conspecific recognition. Our findings point to factors other than conspecific recognition (e.g., neutral drift, phenotypic constraint) that have led to the extensive diversification of light organs in Sergestes s.l and impart caution about interpreting ecological significance of visual characters based on the resolution of human vision. This work provides new insight into deep‐sea animal interaction, supporting the idea that—at least for these mesopelagic shrimps—nonvisual signals may be required for conspecific recognition.     

The absence of sharks from abyssal regions of the world's oceans

The oceanic abyss (depths greater than 3000 m), one of the largest environments on the planet, is characterized by absence of solar light, high pressures and remoteness from surface food supply necessitating special molecular, physiological, behavioural and ecological adaptations of organisms that live there. Sampling by trawl, baited hooks and cameras we show that the Chondrichthyes (sharks, rays and chimaeras) are absent from, or very rare in this region. Analysis of a global data set shows a trend of rapid disappearance of chondrichthyan species with depth when compared with bony fishes. Sharks, apparently well adapted to life at high pressures are conspicuous on slopes down to 2000 m including scavenging at food falls such as dead whales. We propose that they are excluded from the abyss by high-energy demand, including an oil-rich liver for buoyancy, which cannot be sustained in extreme oligotrophic conditions. Sharks are apparently confined to ca 30% of the total ocean and distribution of many species is fragmented around sea mounts, ocean ridges and ocean margins. All populations are therefore within reach of human fisheries, and there is no hidden reserve of chondrichthyan biomass or biodiversity in the deep sea. Sharks may be more vulnerable to over-exploitation than previously thought.     

物种丰富度垂直分布格局及影响机制

物种丰富度分布格局是一定地域内物种丰富度沿三维空间的立体分布,包括物种丰富度在经度、纬度和垂直梯度(海拔高度和海水深度)三个维度上的空间分异。近年来物种多样性的垂直分布格局与机制研究得到了生物地理学家和生态学家的重视。物种丰富度的垂直分布格局存在多种类型,但随海拔增加而物种数减少的单调递减模型和中海拔物种丰富度最高的单峰模型较为常见。目前在机制研究中验证较多的是气候稳定性、生物因子(种间相互作用)、能量、生境异质性、干扰、进化时间、物种分化速率、面积、中域效应(mid-domain effect)、生态位保守性(niche conservatism)等假说和机制。物种丰富度的分布格局是多方面因素综合作用的结果;由于地理、地形、气候、地质演化历史、物种库和进化历史、物种分化速率、干扰等差异,在不同地区存在着特别的物种丰富度空间分布格局和机制;处于同一地区的不同类群的物种也因进化扩散历史和生态适应能力不同而呈现多样化的分布格局。因此,对不同地区和类群的物种丰富度格局和机制进行研究应具体分析后才能得到可信结论。