Discovery of new hydrothermal activity and chemosynthetic fauna on the Central Indian Ridge at 18°-20° S

Indian Ocean hydrothermal vents are believed to represent a novel biogeographic province, and are host to many novel genera and families of animals, potentially indigenous to Indian Ocean hydrothermal systems. In particular, since its discovery in 2001, much attention has been paid to a so-called 'scaly-foot' gastropod because of its unique iron-sulfide-coated dermal sclerites and the chemosynthetic symbioses in its various tissues. Despite increasing interest in the faunal assemblages at Indian Ocean hydrothermal vents, only two hydrothermal vent fields have been investigated in the Indian Ocean. Here we report two newly discovered hydrothermal vent fields, the Dodo and Solitaire fields, which are located in the Central Indian Ridge (CIR) segments 16 and 15, respectively. Chemosynthetic faunal communities at the Dodo field are emaciated in size and composition. In contrast, at the Solitaire field, we observed faunal communities that potentially contained almost all genera found at CIR hydrothermal environments to date, and even identified previously unreported taxa. Moreover, a new morphotype of 'scaly-foot' gastropod has been found at the Solitaire field. The newly discovered 'scaly-foot' gastropod has similar morphological and anatomical features to the previously reported type that inhabits the Kairei field, and both types of 'scaly-foot' gastropods genetically belong to the same species according to analyses of their COI gene and nuclear SSU rRNA gene sequences. However, the new morphotype completely lacks an iron-sulfide coating on the sclerites, which had been believed to be a novel feature restricted to 'scaly-foot' gastropods. Our new findings at the two newly discovered hydrothermal vent sites provide important insights into the biodiversity and biogeography of vent-endemic ecosystems in the Indian Ocean.     

Large scale patterns and trophic structure of southern African rocky shores: the roles of geographic variation and wave exposure

In this study we revise the biogeographic delimitation, and large-scale patterns of community structure of the intertidal rocky shores of southern Africa. We use binary (presence/absence) and per-species biomass data collected at fifteen localities and thirty-seven different rocky sites, encompassing the shores of southern Namibia, South Africa and southern Mozambique. Multivariate analyses revealed that the shores of southern Africa (south of 25°) can be divided into three main biogeographic provinces: the west coast or Namaqua province, the south coast or Agulhas province and the east coast or Natal province. The biomass structure of the intertidal rocky shores communities of southern Africa varied at a large scale, corresponding to biogeographic differences, while local-scale variation accorded with the intensity of local wave action. The average biomass of west coast communities was on average significantly greater than that of the south and east provinces. At a local scale, the community biomass on exposed shores was an order of magnitude greater than on sheltered shores, within all biogeographic provinces. Semi-exposed shores exhibited intermediate average biomass. The trophic structure of these communities varied significantly with wave action: autotrophs, filter-feeders and invertebrate predators were more prevalent on wave exposed than sheltered shores, whereas grazers were more abundant on sheltered and semi-exposed shores. Exposed shores were consistently dominated by far fewer species than semi-exposed and sheltered shores, independently of biogeographic differences. Within all biogeographic provinces semi-exposed and sheltered shores were more diverse than exposed shores. West coast intertidal communities therefore had high levels of biomass, but were consistently species-poor. Several working hypotheses that could explain these large and small-scale patterns are presented.     

New directions in biogeography?

The deduction of biogeographic process from the study of its history is complicated by the fact that history is a singular thing. This singularity allows no estimation of the degree of determinism surrounding the realization of biogeographic processes, and consequently we know almost nothing about the generality of such deductions. Here we discuss a new approach, called 'experimental biogeography', that exploits computers to model faunal build-up repeatedly against a fixed vicariant background over ecological and evolutionary time scales. This new approach enables a biogeographer to be omniscient - to know both vicariant history and actual phylogeny. Moreover, history can be replayed repeatedly to accumulate a sample of multiple phylogenies and to estimate probability density functions for biogeographic variables. Roles of stochastic, historical and ecological processes in adaptive radiations can also be assessed. Experimental biogeography allows examination of the reliability of various methods of recovering historical patterns.     

The discovery of new deep-sea hydrothermal vent communities in the southern ocean and implications for biogeography

Since the first discovery of deep-sea hydrothermal vents along the Galápagos Rift in 1977, numerous vent sites and endemic faunal assemblages have been found along mid-ocean ridges and back-arc basins at low to mid latitudes. These discoveries have suggested the existence of separate biogeographic provinces in the Atlantic and the North West Pacific, the existence of a province including the South West Pacific and Indian Ocean, and a separation of the North East Pacific, North East Pacific Rise, and South East Pacific Rise. The Southern Ocean is known to be a region of high deep-sea species diversity and centre of origin for the global deep-sea fauna. It has also been proposed as a gateway connecting hydrothermal vents in different oceans but is little explored because of extreme conditions. Since 2009 we have explored two segments of the East Scotia Ridge (ESR) in the Southern Ocean using a remotely operated vehicle. In each segment we located deep-sea hydrothermal vents hosting high-temperature black smokers up to 382.8°C and diffuse venting. The chemosynthetic ecosystems hosted by these vents are dominated by a new yeti crab (Kiwa n. sp.), stalked barnacles, limpets, peltospiroid gastropods, anemones, and a predatory sea star. Taxa abundant in vent ecosystems in other oceans, including polychaete worms (Siboglinidae), bathymodiolid mussels, and alvinocaridid shrimps, are absent from the ESR vents. These groups, except the Siboglinidae, possess planktotrophic larvae, rare in Antarctic marine invertebrates, suggesting that the environmental conditions of the Southern Ocean may act as a dispersal filter for vent taxa. Evidence from the distinctive fauna, the unique community structure, and multivariate analyses suggest that the Antarctic vent ecosystems represent a new vent biogeographic province. However, multivariate analyses of species present at the ESR and at other deep-sea hydrothermal vents globally indicate that vent biogeography is more complex than previously recognised.     

Using directed phylogenetic networks to retrace species dispersal history

Methods designed for inferring phylogenetic trees have been widely applied to reconstruct biogeographic history. Because traditional phylogenetic methods used in biogeographic reconstruction are based on trees rather than networks, they follow the strict assumption in which dispersal among geographical units have occurred on the basis of single dispersal routes across regions and are, therefore, incapable of modelling multiple alternative dispersal scenarios. The goal of this study is to describe a new method that allows for retracing species dispersal by means of directed phylogenetic networks obtained using a horizontal gene transfer (HGT) detection method as well as to draw parallels between the processes of HGT and biogeographic reconstruction. In our case study, we reconstructed the biogeographic history of the postglacial dispersal of freshwater fishes in the Ontario province of Canada. This case study demonstrated the utility and robustness of the new method, indicating that the most important events were south-to-north dispersal patterns, as one would expect, with secondary faunal interchange among regions. Finally, we showed how our method can be used to explore additional questions regarding the commonalities in dispersal history patterns and phylogenetic similarities among species.     

When Indian crabs were not yet Asian - biogeographic evidence for Eocene proximity of India and Southeast Asia

Background  

The faunal and floral relationship of northward-drifting India with its neighboring continents is of general biogeographic interest as an important driver of regional biodiversity. However, direct biogeographic connectivity of India and Southeast Asia during the Cenozoic remains largely unexplored. We investigate timing, direction and mechanisms of faunal exchange between India and Southeast Asia, based on a molecular phylogeny, molecular clock-derived time estimates and biogeographic reconstructions of the Asian freshwater crab family Gecarcinucidae.     

Escaping the heat: range shifts of reef coral taxa in coastal Western Australia

One of the most critical challenges facing ecologists today is to understand the changing geographic distribution of species in response to current and predicted global warming. Coastal Western Australia is a natural laboratory in which to assess the effect of climate change on reef coral communities over a temporal scale unavailable to studies conducted solely on modern communities. Reef corals composing Late Pleistocene reef assemblages exposed at five distinct localities along the west Australian coast were censused and the results compared with coral occurrence data published for the modern reefs offshore of each locality. The resulting comparative data set comprises modern and Late Pleistocene reef coral communities occurring over approximately 12° of latitude. For the modern reefs this gradient includes the zone of overlap between the Dampierian and Flindersian Provinces. Modern reef coral communities show a pronounced gradient in coral composition over the latitudinal range encompassed by the study, while the gradient in community composition is not as strong for Pleistocene communities. Tropical‐adapted taxa contracted their ranges north since Late Pleistocene time, emplacing two biogeographic provinces in a region in which a single province had existed previously. Beta diversity values for adjacent communities also reflect this change. Modern reefs show a distinct peak in beta diversity in the middle of the region; the peak is not matched by Pleistocene reefs. Beta diversity is correlated with distance only for comparisons between modern reefs in the north and the fossil assemblages, further supporting change in distribution of the biogeographic provinces in the study area. Coral taxa present in modern communities clearly expanded and contracted their geographic ranges in response to climate change. Those taxa that distinguish Pleistocene from modern reefs are predicted to migrate south in response to future climate change, and potentially persist in ‘temperature refugia’ as tropical reef communities farther north decline.     

Biogeography and Potential Exchanges Among the Atlantic Equatorial Belt Cold-Seep Faunas

Like hydrothermal vents along oceanic ridges, cold seeps are patchy and isolated ecosystems along continental margins, extending from bathyal to abyssal depths. The Atlantic Equatorial Belt (AEB), from the Gulf of Mexico to the Gulf of Guinea, was one focus of the Census of Marine Life ChEss (Chemosynthetic Ecosystems) program to study biogeography of seep and vent fauna. We present a review and analysis of collections from five seep regions along the AEB: the Gulf of Mexico where extensive faunal sampling has been conducted from 400 to 3300m, the Barbados accretionary prism, the Blake ridge diapir, and in the Eastern Atlantic from the Congo and Gabon margins and the recently explored Nigeria margin. Of the 72 taxa identified at the species level, a total of 9 species or species complexes are identified as amphi-Atlantic. Similarity analyses based on both Bray Curtis and Hellinger distances among 9 faunal collections, and principal component analysis based on presence/absence of megafauna species at these sites, suggest that within the AEB seep megafauna community structure is influenced primarily by depth rather than by geographic distance. Depth segregation is observed between 1000 and 2000m, with the middle slope sites either grouped with those deeper than 2000m or with the shallower sites. The highest level of community similarity was found between the seeps of the Florida escarpment and Congo margin. In the western Atlantic, the highest degree of similarity is observed between the shallowest sites of the Barbados prism and of the Louisiana slope. The high number of amphi-atlantic cold-seep species that do not cluster according to biogeographic regions, and the importance of depth in structuring AEB cold-seep communities are the major conclusions of this study. The hydrothermal vent sites along the Mid Atlantic Ridge (MAR) did not appear as “stepping stones” for dispersal of the AEB seep fauna, however, the south MAR and off axis regions should be further explored to more fully test this hypothesis.     

Microbe-mineral biogeography from multi-year incubations in oceanic crust at North Pond,Mid-Atlantic Ridge

Subseafloor oceanic crust is a vast yet poorly sampled habitat for life. Recent studies suggest that microbial composition in crustal habitats is variable in space and time, but biogeographic patterns are difficult to determine due to a paucity of data. To address this, we deployed hundreds of mineral colonization experiments at and below the seafloor for 4–6 years at North Pond, a borehole observatory network in cool (<10°C) and oxic oceanic crust on the western flank of the Mid-Atlantic Ridge. The overall community composition of mineral incubations reveals that colonization patterns are site dependent, with no correlation to mineral type. Only a few members of the Thioalkalispiraceae and Thioprofundaceae exhibited a mineral preference pattern, with generally higher abundance on metal sulphides compared to silicates, while taxa of the Gammaproteobacteria and Deltaproteobacteria were common in the colonization experiments. In comparison to datasets from other crustal habitats, broader biogeographic patterns of crustal communities emerge based on crustal habitat type (surface-attached communities versus fluid communities), redox environment and possibly crustal age. These comparisons suggest successional biogeography patterning that might be used as an indicator of how recently permeable pathways were established within oceanic crust.     

Australia's marine biogeography revisited: Back to the future?

The recognition of broad biogeographic provinces provides an important framework for ecological and conservation biological research. Marine biologists have long recognized distinct biogeographic provinces in southern Australia, primarily on the basis of qualitative differences in intertidal species assemblages. Here we provide an a priori test for these traditional eastern (Peronian), western (Flindersian) and south‐eastern (Maugean) provinces. Specifically, we analyse distributional data for approximately 1500 algal species using the newly available Australian Virtual Herbarium, an online database of herbarium specimens. Our quantitative algal analyses across southern Australia identify three distinct biogeographic assemblages, consistent with traditional qualitative provinces. We argue that these broad provinces provide a highly effective framework for understanding and managing Australia's marine biodiversity. In particular, biogeographic provinces provide a regional framework for integrating the ongoing discovery of biological variation at finer scales. More broadly therefore we recommend that biologists undertake quantitative analyses to test provincial biogeographic boundaries around the globe.     

Microdistribution of Faunal Assemblages at Deep-Sea Hydrothermal Vents in the Southern Ocean

Chemosynthetic primary production by microbes supports abundant faunal assemblages at deep-sea hydrothermal vents, with zonation of invertebrate species typically occurring along physico-chemical gradients. Recently discovered vent fields on the East Scotia Ridge (ESR) in the Southern Ocean represent a new province of vent biogeography, but the spatial dynamics of their distinct fauna have yet to be elucidated. This study determines patterns of faunal zonation, species associations, and relationships between faunal microdistribution and hydrothermal activity in a vent field at a depth of 2,400 m on the ESR. Remotely operated vehicle (ROV) dives obtained high-definition imagery of three chimney structures with varying levels of hydrothermal activity, and a mosaic image of >250 m² of seafloor co-registered with temperature measurements. Analysis of faunal microdistribution within the mosaiced seafloor reveals a consistent pattern of faunal zonation with increasing distance from vent sources and peak temperatures. Assemblages closest to vent sources are visibly dominated by a new species of anomuran crab, Kiwa n. sp. (abundance >700 individuals m⁻²), followed by a peltospiroid gastropod (>1,500 individuals m⁻²), eolepadid barnacle (>1,500 individuals m⁻²), and carnivorous actinostolid anemone (>30 individuals m⁻²). Peripheral fauna are not dominated by a single taxon, but include predatory and scavenger taxa such as stichasterid seastars, pycnogonids and octopus. Variation in faunal microdistribution on chimneys with differing levels of activity suggests a possible successional sequence for vent fauna in this new biogeographic province. An increase in δ³⁴S values of primary consumers with distance from vent sources, and variation in their δ¹³C values also indicate possible zonation of nutritional modes of the vent fauna. By using ROV videography to obtain a high-resolution representation of a vent environment over a greater extent than previous studies, these results provide a baseline for determining temporal change and investigations of processes structuring faunal assemblages at Southern Ocean vents.     

A hierarchical classification of freshwater mussel diversity in North America

Aim North America harbours the most diverse freshwater mussel fauna on Earth. This fauna has high endemism at the continental scale and within individual river systems. Previous faunal classifications for North America were based on intuitive, subjective assessments of species distributions, primarily the occurrence of endemic species, and do not portray continent‐wide patterns of faunal similarity. The aim of this study is to provide an analytical portrayal of patterns of mussel diversity in a hierarchical framework that informs the biogeographical history of the fauna. Location The study considered the mussel fauna of North America from the Rio Grande system northwards. Methods Patterns of mussel faunal similarity in 126 river systems or lake watersheds across North America were examined. The dataset was developed from the literature and consisted of recent species presence/absence (282 species) in each drainage unit; subspecies were not included. Patterns of mussel diversity were examined with hierarchical cluster analysis, based on a pairwise distance matrix between all drainage units. Results Cluster analysis revealed 17 faunal provinces within four major faunal regions: Mississippian, Atlantic, Eastern Gulf and Pacific. The Mississippian Region dominates the North American fauna with 11 provinces, including five not recognized by previous classifications: Mississippi Embayment, Upper Mississippi, Great Plains, Ohioan and Pontchartrain–Pearl–Pascagoula. Within the Eastern Gulf Region (containing three provinces), the Escambia–Choctawhatchee Province is distinctive from the Apalachicolan Province, under which it was previously subsumed. Patterns of diversity in the Atlantic Region (two provinces) and Pacific Region (one province) were similar to previous classifications. Main conclusions The classification proposed in this study largely corroborates earlier schemes based on the occurrence of endemic species but identifies additional heterogeneity that reflects unique assemblages of widely distributed species. The study proposes a hierarchical structure that illustrates relationships among these provinces. Although some provinces in the Mississippian Region have high endemism, all Mississippian provinces share a group of widely distributed species. The Atlantic and Eastern Gulf regions have distinctive, endemic faunas suggesting limited past connectivity with the Mississippian Region. The Pacific Region is the most distinct fauna in North America and bears close affinity to the Eurasian mussel fauna.     

土壤动物群落空间格局和构建机制研究进展

群落空间格局和构建机制一直是生态学研究的核心内容。在生物多样性严重丧失的背景下, 揭示群落空间格局及其构建机制, 有助于深刻理解生物多样性丧失的原因, 更有助于应对生物多样性保护等重大生态环境问题。然而, 陆地生态系统的研究多集中于地上生物群落, 对地下生态系统, 尤其是土壤动物空间格局和构建机制的研究尚不充分。事实上, 土壤动物多样性是全球生物多样性的关键组成之一, 是地下生态系统结构和功能维持的重要部分。对土壤动物空间格局和构建机制的研究, 能明确不同空间尺度条件下土壤动物多样性的维持机制。土壤动物群落常在多种空间尺度形成复杂的空间分布格局, 因此, 本文首先介绍了不同空间尺度主要土壤动物群落的空间自相关性特征, 阐述了土壤动物群落斑块和孔隙镶嵌分布的复杂空间格局。继而阐明这种空间格局主要受生物间作用、环境过滤和随机扩散的调控, 并说明这三个过程对土壤动物群落的调控能力和作用方式。作者提出, 这三个过程仍是今后土壤动物群落空间格局和构建机制研究的重点内容, 需要进一步加强以土壤动物为研究对象的群落构建理论的验证和发展。我国土壤动物群落空间格局和构建机制起步较晚, 希望本文能够促进我国土壤动物生态学相关领域的研究。     

Biogeography of the phoracanthine beetles (Coleoptera: Cerambycidae)

Six biogeographic subregions of the phoracanthine beetles are proposed on the basis of 136 valid species in all eleven genera of this Australasian group of insects: the Kosciuskan, Western, Eyrean, Torresian, Timorian and New Guinean. Both phenetic and cladistic approaches are employed in the biogeographic study of these beetles. Phenetically, the Kosciuskan and Torresian fauna are the most similar and the least faunal similarity occurs between the New Guinean subregion and all others. Cladistically, two distinct peripheral faunas are divided: the southern Western+Kosciuskan and the northern New Guinean+Torresian+Timorian. The Eyrean fauna in the central Australian continent is more distantly related to the remainder than the latter are to each other. Biogeographic history of the phoracanthine beetles is discussed extensively.     

Fungal community demonstrates stronger dispersal limitation and less network connectivity than bacterial community in sediments along a large river

Despite the essential functions of sedimentary bacterial and fungal communities in biogeochemical cycling, little is known about their biogeographic patterns and driving processes in large rivers. Here we investigated the biogeographic assemblies and co-occurrence patterns of sedimentary bacterial and fungal communities in the Jinsha River, one of the largest rivers in southwestern China. The mainstream of river was divided into upstream, midstream and downstream. The results showed that both bacterial and fungal communities differed significantly among three sections. For both communities, their composition variations in all sites or each river section were controlled by the combination of dispersal limitation and environmental selection, and dispersal limitation was the dominant factor. Compared with bacteria, fungi had stronger dispersal limitation. Co-occurrence network analyses revealed higher network connectivity but a lower proportion of positive interaction in the bacterial than fungal network at all sites. In particular, the keystone species belonging to bacterial phyla Proteobacteria and Firmicutes and fungal phyla Ascomycota and Chytridiomycota may play critical roles in maintaining community function. Together, these observations indicate that fungi have a stronger dispersal limitation influence and less network connectivity than bacteria, implying different community assembly mechanisms and ecological functions between bacteria and fungi in large rivers.     

Deconstructing the mammal species richness pattern in Europe – towards an understanding of the relative importance of climate,biogeographic history,habitat heterogeneity and humans

Aim We deconstructed the mammal species richness pattern in Europe to assess the importance of large‐scale gradients in current macroclimate relative to biogeographic history, habitat heterogeneity and human influence (HHH variables) as richness determinants for total species, and for widespread and endemic species separately. Location Europe, west of 30° E. Methods We deconstructed total species richness (50‐km resolution) into its widespread and endemic species richness components. We used simultaneous autoregressive modelling (SAR) with information‐theoretic model selection and variation partitioning to assess the importance of macroclimate and HHH variables. The HHH variables included two historical factors, estimated by novel methodologies: (1) ice‐age‐driven dynamics, represented by accessibility to recolonization from hindcasting‐estimated glacial refugia, and (2) biogeographic peninsular dynamics, represented by distance to the entry region for the main European faunal source in western Asia. Results A large fraction of explained variation was shared between macroclimate and HHH in the SAR models. For total species richness, more variation could be uniquely attributed to macroclimate than to HHH, whereas for the deconstructed patterns (widespread and endemic species) the opposite was the case. Considering the individual factors, there was a strong peninsula effect on both widespread and endemic species richness but not on total richness. Main conclusions Both macroclimate and HHH variables (history, habitat heterogeneity and human influence) proved important predictors of species richness, but also difficult to disentangle. Notably, biogeographic history, in particular peninsular dynamics, is an important determinant of widespread and endemic species richness.     

The macroecology of reef fish agonistic behaviour

Understanding the interplay between processes operating at large and small spatiotemporal scales in shaping biotic interactions remains challenging. Recent studies illustrate how phenotypic specialization, species life-history traits and/or resource partitioning recurrently underlie the structure of mutualistic interactions in terrestrial ecosystems along large latitudinal gradients of biodiversity. However, we know considerably less about how local processes interact with large-scale patterns of biodiversity in modulating biotic interactions in the marine realm. Considering agonistic behaviour as a proxy for contest competition, we empirically investigate whether the structure of reef fish agonistic interactions is conserved across a 34 000-km longitudinal gradient of biodiversity. By sampling coral reefs using standardized remote underwater video, we found recurrent patterns of fish agonistic behaviour in disparate communities distributed across five biogeographic provinces of the Pacific and Atlantic oceans. While the sheer number of species increases with regional richness, the number of aggressive disputes at the habitat scale is similar across communities. We then combined generalized linear models and network theory to reveal that, the emergent structure of local agonistic networks is not modular but instead recurrently display a nested structure, with a core of highly interactive site-attached herbivores of the Pomacentridae family. Therefore, despite the increase in the number of species involved in agonistic interactions toward speciose communities, the network structure is conserved along the longitudinal richness gradient because local disputes are mostly driven by closely-related, functionally-similar species. These findings suggest that evolutionary and local processes interact in modulating reef fish agonistic behaviour and that fine-scale niche-partitioning can structure the ecological networks in marine ecosystems.