Patterns in marine hydrozoan richness and biogeography around southern Africa: implications of life cycle strategy

Aim To examine patterns of marine hydrozoan richness around southern Africa and to test the hypothesis that patterns of biogeography become weaker with increasing dispersal ability. Location The coastline of southern Africa from 21° S, 14° E to 28° S, 33° E, extending from the intertidal zone seawards a distance of 200 nautical miles. Methods Published and unpublished information on the distribution of marine Hydrozoa was entered as presence/absence data onto a gridded coastline of the region. A similarity matrix between the species composition of grid squares was constructed using the Bray–Curtis index and visualized using non-metric multidimensional scaling ordinations. Separate analyses were conducted, and compared, on the three major life cycle groupings: holoplanktic, meroplanktic and benthic. Results Over 450 species of marine Hydrozoa have been reported from the region, and species richness increases eastwards, in a manner at odds with the distribution of sampling effort. There was a significant correlation between the geographic structures of the resemblance matrices generated for the three life cycle groupings. In other words, all three groups showed similar patterns of biogeography around the region, and these were broadly similar to those reported by others. However, there were differences between them that reflect the resolution at which the data were examined. At a level of 40% similarity, there was no biogeographic structure to the holoplanktic fauna, the meroplanktic taxa were simply sub-divided into cool- and warm-temperate/subtropical elements, and in the case of benthic taxa, the cool-water fauna was further split into a southern Namaqua and a depauperate northern Namib component. Even at a resolution of 70% similarity, the holopelagic taxa could be separated only into cool-temperate and warm-temperate/subtropical faunas. Main conclusions Holoplanktic taxa show comparatively less biogeographic structure than meroplanktic taxa, which in turn show less clearly defined biogeographic structure than benthic taxa. It is suggested that this is related to the interaction between oceanography and dispersive-stage duration. The role that the Agulhas Current plays in influencing the Benguela Current fauna is highlighted. This study has implications for conservation planning exercises based on protecting representative biotopes in different biogeographic regions.     

Large-scale species-richness gradients in the Atlantic Ocean

The increase in species richness from the poles to the Equator has been observed in numerous terrestrial and aquatic taxa. A number of different hypotheses have been put forward as explanations for this trend, e.g. area and energy availability. However, whether these hypotheses apply to large spatial scales in marine environments remains unclear. The present study shows a clear latitudinal gradient from high to low latitude (from 80 degrees N to 70 degrees S) in marine species richness for 6643 species (fishes and invertebrates) in 10 different taxa dwelling in benthic and pelagic habitats on both sides of the Atlantic. The patterns in benthic taxa are strongly influenced by coastal hydrographic processes, with marked peaks and troughs, and consequently the gradients are not symmetric along both Atlantic sides. Pelagic taxa show a plateau-shaped distribution and the influence from coastal events on gradients could not be demonstrated. The relationships between species richness and different environmental factors indicate that area size does not explain the latitudinal pattern in benthic species richness on a large spatial scale. Sea-surface temperature (positive relationship) is the best predictor of this pattern for benthic species, and nitrate concentration (negative relationship) is the best predictor for pelagic species. The results call into question the existence of a single primary cause that would explain the pattern in marine species richness on a large spatial scale.     

Marine, intertidal, freshwater and terrestrial biodiversity of an isolated polar archipelago

Aim We ask how biodiverse is a polar archipelago; how this faunal richness is spread across marine, intertidal, freshwater, terrestrial and parasitic realms; and how fast species are accumulated with increased sampling effort. Location The South Orkney Islands (SOI), Scotia Arc, Southern Ocean. Methods We sampled mega‐ and macro‐benthos at the SOI using scuba in the shallows (0–10 m), a rough bottom otter trawl at 150–250 m and an Agassiz trawl and epibenthic sledge, both at depths of 200, 500, 1000 and 1500 m. We also collated species occurrence at the SOI in each realm from a century of literature and modern databases to investigate patterns in species accumulation, endemism, faunistic affinities and bathymetric ranges in three model taxa. Results Our 11 benthic samples showed that point biodiversity at the SOI is high, yielding 19 classes and 158 species. Nearly a third were new to the area, whilst five species and one genus were new to science. The shallowest samples were richest but had fewest new records of species. Known richness at the SOI is dominated by marine species (1026), of which 821 (83.3%) were benthic. Across all realms, 1224 species (50 classes, 24 phyla) were recorded, of which 43 were intertidal, 64 freshwater, 100 terrestrial, 60 parasitic and 40 birds. Species accumulation curves for model taxa showed new sampling yields about 0.75% per sample of known benthic richness, so by Antarctic standards we know the SOI quite well. Most species are Southern Ocean endemics, but very few occur only at the SOI. Main conclusions This first estimate of faunal biodiversity of a polar locality demonstrates both high richness and high levels of knowledge at the SOI. As suspected but never quantified, the benthos dominates polar biodiversity, at least at the SOI. Marine species there constitute 20% of those recently listed for the entire Southern Ocean, whilst > 60% of terrestrial species are known from Antarctica. The SOI, being one of the better‐studied polar locations, of known age and with a discrete shelf, represent an important source of comparison for biodiversity studies. Our data clearly show that richness and our knowledge of the polar fauna differ across environments.     

Deconstructing latitudinal species richness patterns in the ocean: does larval development hold the clue?

The consistent decrease in species richness with latitude shows several exceptions among marine organisms. We hypothesize that contrasting latitudinal diversity gradients can be explained by differences in critical life-history attributes, such as mode of larval development (MLD). We deconstructed latitudinal species richness patterns of marine benthic invertebrates according to MLD to elucidate differences in patterns of species richness and to reveal underlying processes. The patterns of species richness were remarkably similar across taxa within MLD but differed between MLD. Species richness decreased polewards in planktotrophic species and increased in direct developers. Temperature explained most of the variation in species richness. Low temperature at high latitudes may generally favour direct developing species, but, together with low chlorophyll- a concentration, limit the distribution of planktotrophic species. The contrasting influence of temperature on different MLDs might be explained by its effect on the length of planktonic life and on brooding costs.     

Increased accuracy of species lists developed for alpine lakes using morphology and cytochrome oxidase I for identification of specimens

The first step in many community ecology studies is to produce a species list from a sample of individuals. Community ecologists now have two viable ways of producing a species list: morphological and barcode identification. In this study, we compared the taxonomic resolution gained by a combined use of both methods and tested whether a change in taxonomic resolution significantly impacted richness estimates for benthic macroinvertebrates sampled from ten lakes in Sequoia National Park, USA. Across all lakes, 77 unique taxa were identified and 42% (32) were reliably identified to species using both barcode and morphological identification. Of the 32 identified to species, 63% (20) were identified solely by comparing the barcode sequence from cytochrome oxidase I to the Barcode of Life reference library. The increased resolution using a combined identification approach compared to identifications based solely on morphology resulted in a significant increase in estimated richness within a lake at the order, family, genus and species levels of taxonomy (P < 0.05). Additionally, young or damaged individuals that could not be identified using morphology were identified using their COI sequences to the genus or species level on average 75% of the time. Our results demonstrate that a combined identification approach improves accuracy of benthic macroinvertebrate species lists in alpine lakes and subsequent estimates of richness. We encourage the use of barcodes for identification purposes and specifically when morphology is insufficient, as in the case of damaged and early life stage specimens of benthic macroinvertebrates.     

Assessing species richness of macrofauna associated with macroalgae in Arctic kelp forests (Hornsund,Svalbard)

Kelp beds support diverse and productive benthic coastal ecosystems and are often perceived as cold water analogs of tropical coral reefs, yet the levels of species richness in polar regions have remained largely unexplored. The present study aims to assess the magnitude of macrozoobenthic species richness associated with the macroalgae in an Arctic kelp forest. The study was conducted in Hornsund, a high latitude Arctic fjord off west Spitsbergen (76–77°N). A total of 403 samples (i.e., individual algae) were collected by scuba divers at three sites located along the fjord axis at varying depths (5–10 m). The most common invertebrate species and the sample species richness (mean 11.5 species per sample) were consistent among the most common algal species (Laminaria digitata, Saccharina latissima, Alaria esculenta, Desmarestia aculeta, Odonthalia dentata, Phycodrys rubens). Fauna associated with overstory kelps was concentrated on the holdfasts. A total of 208 species were identified; Bryozoa, Polychaeta and Hydrozoa yielded the highest numbers of species (70, 52 and 37, respectively). The non-parametric Chao2 estimator of true species richness gave an estimate of 259 species (with 95% confidence intervals from 234 to 308). Despite the high sampling effort, the total species richness was not captured. This may stem from the high level of rarity within the fauna associated with kelps; 38% of the species occurred only in one or two samples. The studied fauna was much less diverse than similar assemblages surveyed at lower latitudes. The high numbers of species and the common occurrence of colonial forms (Bryozoa, Hydrozoa) in the current study show that these taxa cannot be omitted in macrobenthic diversity surveys performed within kelp habitats.     

Polychaetes as surrogates for marine biodiversity: lower taxonomic resolution and indicator groups

Due to conservation needs, reliable rapid-assessment methods for mapping of biodiversity are needed. One approach is to use surrogates, i.e. quantities that correlate strongly with the number of species, but are easier to obtain. The purpose of this paper is to test two polychaete surrogates, one for higher taxa and one for indicator groups, that will facilitate prediction of species richness in marine soft-bottom communities. Soft sediment is an important habitat which covers most of the ocean bottom. Data on polychaetes from the North Atlantic were used since polychaetes are often numerically dominant in the benthic assemblages, both with regard to the number of species and their abundance. In the polychaete assemblages along the Norwegian coast, richness at the genus, family and order level were significantly, linearly correlated to total species richness (r 0.92). Polychaetes in the order Terebellida were found to be a good indicator of polychaete species richness and to a lesser extent also of whole benthic assemblages. The group Terebellida is potentially well suited as an indicator group, since it contains long-lived, large species that are easy to sort from the sediment and it is well defined taxonomically. Although promising as proxies for species richness in marine biodiversity studies, the use of lower taxonomic resolution and indicator groups requires further investigations in more local areas where there are conservation issues.     

Trends in the Diversity,Distribution and Life History Strategy of Arctic Hydrozoa (Cnidaria)

This is the first attempt to compile a comprehensive and updated species list for Hydrozoa in the Arctic, encompassing both hydroid and medusa stages and including Siphonophorae. We address the hypothesis that the presence of a pelagic stage (holo- or meroplanktonic) was not necessary to successfully recolonize the Arctic by Hydrozoa after the Last Glacial Maximum. Presence-absence data of Hydrozoa in the Arctic were prepared on the basis of historical and present-day literature. The Arctic was divided into ecoregions. Species were grouped into distributional categories according to their worldwide occurrences. Each species was classified according to life history strategy. The similarity of species composition among regions was calculated with the Bray-Curtis index. Average and variation in taxonomic distinctness were used to measure diversity at the taxonomic level. A total of 268 species were recorded. Arctic-boreal species were the most common and dominated each studied region. Nineteen percent of species were restricted to the Arctic. There was a predominance of benthic species over holo- and meroplanktonic species. Arctic, Arctic-Boreal and Boreal species were mostly benthic, while widely distributed species more frequently possessed a pelagic stage. Our results support hypothesis that the presence of a pelagic stage (holo- or meroplanktonic) was not necessary to successfully recolonize the Arctic. The predominance of benthic Hydrozoa suggests that the Arctic could have been colonised after the Last Glacial Maximum by hydroids rafting on floating substrata or recolonising from glacial refugia.     

Which functional processes control the short-term effect of grazing on net primary production in grasslands?

For unicellular organisms, a lack of effects of local species richness on ecosystem function has been proposed due to their locally high species richness and their ubiquitous distribution. High dispersal ability and high individual numbers may enable unicellular taxa to occur everywhere. Using our own and published data sets on uni- and multicellular organisms, we conducted thorough statistical analyses to test whether (1) unicellular taxa show higher relative local species richness compared to multicellular taxa, (2) unicellular taxa show lower slopes of the species:area relationships and species:individuals relationships, and (3) the species composition of unicellular taxa is less influenced by geographic distance compared to multicellular taxa. We found higher local species richness compared to the global species pool for unicellular organisms than for metazoan taxa. The difference was significant if global species richness was conservatively estimated but not if extrapolated, and therefore higher richness estimates were used. Both microalgae and protozoans showed lower slopes between species richness and sample size (area or individuals) compared to macrozoobenthos, also indicating higher local species richness for unicellular taxa. The similarity of species composition of both benthic diatoms and ciliates decreased with increasing geographic distance. This indicated restricted dispersal ability of protists and the absence of ubiquity. However, a steeper slope between similarity and distance was found for polychaetes and corals, suggesting a stronger effect of distance on the dispersal of metazoans compared to unicellular taxa. In conclusion, we found partly different species richness patterns among uni- and multicellular eukaryotes, but no strict ubiquity of unicellular taxa. Therefore, the effect of local unicellular species richness on ecosystem function has to be reanalyzed. Macroecological patterns suggested for multicellular organisms may differ in unicellular communities.     

Geographical distribution and extinction risk: lessons from Triassic–Jurassic marine benthic organisms

Aim To evaluate the influence of geographical distribution on the extinction risk of benthic marine invertebrates using data from the fossil record, both during times of background extinction and across a mass‐extinction episode. Total geographical range is contrasted with proxies of global abundance to assess the relationships between the two essential components of geographical distribution and extinction risk. Location A global occurrence data base of fossil benthic macro‐organisms from the Triassic and Jurassic periods was used for this study. Methods Geographical distributions and biodiversity dynamics were assessed for each genus (all taxa) or species (bivalves) based on a sample‐standardized data set and palaeogeographical reconstructions. Geographical ranges were measured by the maximum great circle distance of a taxon within a stratigraphic interval. Global abundance was assessed by the number of localities at which a taxon was recorded. Widespread and rare taxa were separated using median and percentile values of the frequency distributions of occurrences. Results The frequency distribution of geographical ranges is very similar to that for modern taxa. Although no significant correlation could be established between local abundance and geographical range, proxies of global abundance are strongly correlated with geographical range. Taxon longevities are correlated with both mean geographical range and mean global abundance, but range size appears to be more critical than abundance in determining extinction risk. These results are valid when geographical distribution is treated as a trait of taxa and when assessed for individual geological stages. Main conclusions Geographical distribution is a key predictor of extinction risk of Triassic and Jurassic benthic marine invertebrates. An important exception is in the end‐Triassic mass extinction, which equally affected geographically restricted and widespread genera, as well as common and rare genera. This suggests that global diversity crises may curtail the role of geographical distribution in determining extinction risk.     

The Relationship between Genus Richness and Geographic Area in Late Cretaceous Marine Biotas: Epicontinental Sea versus Open-Ocean-Facing Settings

For present-day biotas, close relationships have been documented between the number of species in a given region and the area of the region. To date, however, there have been only limited studies of these relationships in the geologic record, particularly for ancient marine biotas. The recent development of large-scale marine paleontological databases, in conjunction with enhanced geographical mapping tools, now allow for their investigation. At the same time, there has been renewed interest in comparing the environmental and paleobiological properties of two broad-scale marine settings: epicontinental seas, broad expanses of shallow water covering continental areas, and open-ocean-facing settings, shallow shelves and coastlines that rim ocean basins. Recent studies indicate that spatial distributions of taxa and the kinetics of taxon origination and extinction may have differed in these two settings. Against this backdrop, we analyze regional Genus-Area Relationships (GARs) of Late Cretaceous marine invertebrates in epicontinental sea and open-ocean settings using data from the Paleobiology Database. We present a new method for assessing GARs that is particularly appropriate for fossil data when the geographic distribution of these data is patchy and uneven. Results demonstrate clear relationships between genus richness and area for regions worldwide, but indicate that as area increases, genus richness increases more per unit area in epicontinental seas than in open-ocean settings. This difference implies a greater degree of compositional heterogeneity as a function of geographic area in epicontinental sea settings, a finding that is consistent with the emerging understanding of physical differences in the nature of water masses between the two marine settings.     

Power law relationships among hierarchical taxonomic categories in algae reveal a new paradox of the plankton

Aim In this continental‐scale study, the biodiversity of benthic and planktonic algal communities was explored. A recent analysis of extinct and extant tree communities by Enquist et al. (2002) showed that richness of higher taxa was a power function of species richness, invariant across temporal and spatial scales. Here we examined whether the relationships between algal richness at hierarchical taxonomic levels conform to power laws as seen for trees, and if these relationships differ between benthic and planktonic habitats. Location Streams from more than 50 major watersheds in the United States. Method A total of 3698 samples were collected from 1277 locations by the National Water‐Quality Assessment Program. Three types of stream habitat were sampled: richest targeted habitats, depositional targeted habitats, and phytoplankton. The relationships between taxonomic richness at the species level vs. all higher categories from genus to phylum across the three habitats were examined by ordinary least squares (OLS) regressions after ln‐transformation of all variables. The slopes, b, of these regressions represent the exponents of the power functions that scaled the richness of higher taxonomic levels (T) to species richness (S) in the form: T∝S^b. Results Algal richness at hierarchical taxonomic categories (genus to phylum) is a power function of species richness. The scaling exponent of this function, which captures the diversification of higher taxa, i.e. the rate of increase of their richness with the increase of species richness, is significantly different across environments. Main conclusions The differential algal diversification in the three studied habitats emphasizes the fundamental role of the environment in structuring the communities of simple organisms such as algae. The finding that the diversification of higher taxa is greater in the seemingly homogeneous planktonic environment, when compared to benthic habitats, encompassing an array of ecological niches, poses a new paradox of the plankton.     

Regional‐scale patterns and predictors of species richness and abundance across twelve major tropical inter‐reef taxa

Species richness and abundance are biodiversity metrics widely used to describe and estimate changes in biodiversity. Studies of marine species richness and abundance typically focus on one, or just a few, taxa. Consequently, it is currently not possible to understand the performance of predictors of species richness and abundance across marine taxa. Using a taxonomically comprehensive dataset of twelve major taxa of flora and fauna from eight phyla sampled from the inter‐reef seabed region of the Great Barrier Reef, Australia, we used boosted regression trees to test the performance of fourteen environmental and spatial predictors of species richness and abundance. Sediment composition predicted richness best for all taxa: gravel contributed up to 39% relative influence for one group and all taxa had low richness in muddy habitats. Sea surface temperature, seabed current shear stress, depth and latitude were also influential predictors for species richness for eight groups. Sediment was frequently an influential predictor for abundance also, while distance to domain (reef/coast) and longitude were relatively influential for six taxa. Within‐site richness was correlated between nearly all pairs of taxa, as was within‐site abundance, however ρ values were low. Overall, model performance was high, explaining up to 62% deviance of species richness, and 38% of abundance. Typically, deviance explained was greater for richness than abundance and may indicate that some drivers of species richness operate independently of any effects on species richness mediated by their effect on abundance. Deviance explained differed most between richness and abundance for bryozoans (23.3% difference) and soft corals (15.2% difference). While sediments were consistently the best predictors across all taxa, the inconsistent influence of all other predictors across taxonomic groups, as well as the low correlation of richness and abundance across taxonomic groups, cautions against predicting regional patterns of species richness and abundance from few taxa.