Conservation biogeography of the Antarctic

Aim

To present a synthesis of past biogeographic analyses and a new approach based on spatially explicit biodiversity information for the Antarctic region to identify biologically distinct areas in need of representation in a protected area network.

Location

Antarctica and the sub‐Antarctic.

Methods

We reviewed and summarized published biogeographic studies of the Antarctic. We then developed a biogeographic classification for terrestrial conservation planning in Antarctica by combining the most comprehensive source of Antarctic biodiversity data available with three spatial frameworks: (1) a 200‐km grid, (2) a set of areas based on physical parameters known as the environmental domains of Antarctica and (3) expert‐defined bioregions. We used these frameworks, or combinations thereof, together with multivariate techniques to identify biologically distinct areas.

Results

Early studies of continental Antarctica typically described broad bioregions, with the Antarctic Peninsula usually identified as biologically distinct from continental Antarctica; later studies suggested a more complex biogeography. Increasing complexity also characterizes the sub‐Antarctic and marine realms, with differences among studies often attributable to the focal taxa. Using the most comprehensive terrestrial data available and by combining the groups formed by the environmental domains and expert‐defined bioregions, we were able to identify 15 biologically distinct, ice‐free, Antarctic Conservation Biogeographic Regions (ACBRs), encompassing the continent and close lying islands.

Main conclusions

Ice‐free terrestrial Antarctica comprises several distinct bioregions that are not fully represented in the current Antarctic Specially Protected Area network. Biosecurity measures between these ACBRs should also be developed to prevent biotic homogenization in the region.     

Historical biogeography and ecology of a Continental Antarctic mite genus, Maudheimia (Acari, Oribatida): evidence for a Gondwanan origin and Pliocene-Pleistocene speciation

Of the eight genera (30 species) of extant Acari in Continental (= East) Antarctica, the genus Maudheimia Dalenius & Wilson, 1958 (Oribatida; Maudheimiidae) is uniquely endemic. A Gondwanan origin is proposed for the genus based on antiquity, inferred from endemism, a widespread distribution throughout Continental Antarctica and a limited dispersal capacity. Adaptation for a high montane epilithic existence, a necessity for the origination and long-term persistence in Antarctica, is inferred from the life history, physiology and ecology of Maudheimia. Phylogenetic analysis placed all four Maudheimia species in a single (generic) clade with the following structure (M. pelronia Wallwork, 1962 (M. tanngardenensis Coctzee, 1997 (M. manhalli Coetzee, 1997 (M. wilsoni Dalenius & Wilson, 1958)))). Geographical distributions of the Maudheimia species, in relation to their phylogenetic relationships, support the hypothesis that post-Gondwanan speciation occurred as a consequence of isolation during glaciation of Antarctica.     

Moss endemism in the Mexican flora

Although the moss flora of Mexico consists of nearly 1000 species, only 77 are endemic. The country has many poorly collected or unexplored areas, but the number of endemic mosses is not expected to undergo a substantial increase; percent endemism has in fact decreased with taxonomic revisions and monographs and better exploration in other countries. Literature and herbarium records (n?=?584) were used to obtain an updated list of endemic mosses and their state distribution in Mexico. Cluster analysis and mapping indicate that there are three main areas of endemism: Lowland areas in various states, the mountain area along 19–20°N lat., and the highlands in Oaxaca and Chiapas. Similarity by province shows that Trans-Mexican Volcanic Belt, Sierra Madre Oriental, Chihuahuan Desert, and Sierra Madre del Sur have the highest numbers of endemic species. Five monotypic genera and 76 species (including two infraspecific taxa), many of which have comparatively narrow geographical ranges, suggest that speciation is recent, that species have had little time to disperse, and were formed by Pleistocene environmental climates in the highlands; older speciation may be represented by widespread disjunct species that still are found in the highlands of Mexico.     

Biodiversity and biogeography of southern temperate and polar bryozoans

Aim To describe the distribution of biodiversity and endemism of bryozoans in southern temperate and polar waters. We hypothesized that we would find: (1) no strong latitudinal richness gradient; (2) striking contrasts in richness and endemism between clades and between regions; and (3) that faunal similarity of regions would cluster geographically around each southern continent. Location South Atlantic, Indian and Pacific Oceans and the Southern Ocean. Methods We constructed a data base from known literature, regional data bases and recent finds. We regionalized each southern continent, calculated levels of richness and endemism for each region and continent, and used primer 5 to perform multivariate statistical analysis. Results A third (1681) of global bryozoan species described occur south of 30° S, of which c. 87% were cheilostomes. In richness we found no latitudinal cline and change across longitude was stronger. New Zealand was richest and had the most (60%) endemic species, followed by Antarctica at 57%. There were striking contrasts in regional richness and endemism between clades but the highest levels of between‐region similarity were around Antarctica. The timing of past continent connectivity was reflected. Main conclusions Bryozoans show strong hemispherical asymmetry in richness and, like molluscs and corals, decrease away from Australasia rather than with latitude. Species endemism is much lower in Antarctic bryozoans than previously thought, and as this taxon is not particularly dispersive and is now amongst the best studied regionally, maybe Antarctic endemism in general is lower and Antarctica less cut‐off to species dispersal than previously thought. However, Antarctic generic endemism is double the level previously calculated and regional faunal similarities are much higher than around other continents – both reflecting long‐term isolation. Bryozoans, in contrast to the paradigm of Antarctic fauna, may be fairly robust to predicted climate change. Paradoxically, they may also be one of the best taxa to monitor to sensitively detect marine benthic responses.     

Diversity, endemism and evolution in the Coral Triangle

In a recent paper by D. R. Bellwood and C. P. Meyer ('Searching for heat in a marine biodiversity hotspot', Journal of Biogeography , 2009, 36 , 569–576), the authors had two evident objectives: (1) to disprove the theory that the geographical origins of reef organisms could be determined by locating concentrations of endemic species, and (2) to emphasize that the high diversity of the Coral Triangle was due to an accumulation of species from outside that area. With regard to the first point, no such theory had previously been proposed to my knowledge. Second, the accumulation theory was promoted without consideration of the facts supporting the centre of origin hypothesis, except to dismiss it by saying that it had its origin in pre-continental drift ideas. This short response outlines the properties and evidence for the operation of centres of origin in this region.     

Marine shelf habitat: biogeography and evolution

We synthesize the evolutionary implications of recent advances in the fields of phylogeography, biogeography and palaeogeography for shallow‐water marine species, focusing on marine speciation and the relationships among the biogeographic regions and provinces of the world. A recent revision of biogeographic provinces has resulted in the recognition of several new provinces and a re‐evaluation of provincial relationships. These changes, and the information that led to them, make possible a clarification of distributional dynamics and evolutionary consequences. Most of the new conclusions pertain to biodiversity hotspots in the tropical Atlantic, tropical Indo‐West Pacific, cold‐temperate North Pacific, and the cold Southern Ocean. The emphasis is on the fish fauna, although comparative information on invertebrates is utilized when possible. Although marine biogeographic provinces are characterized by endemism and thus demonstrate evolutionary innovation, dominant species appear to arise within smaller centres of high species diversity and maximum interspecies competition. Species continually disperse from such centres of origin and are readily accommodated in less diverse areas. Thus, the diversity centres increase or maintain species diversity within their areas of influence, and are part of a global system responsible for the maintenance of biodiversity over much of the marine world.     

贡嘎山东坡植物区系的垂直分布格局

为了探讨贡嘎山植物区系的垂直分化特征及其与周边地区植物区系的联系,结合样带法与样方法,对贡嘎山东坡垂直植被带进行了调查,统计得出各垂直植被带的科、属的物种数量,分析了科、属、种级区系成分的构成及其沿海拔梯度的分布格局,并对各垂直植被带区系的相似性进行了聚类分析。结果表明：1)贡嘎山植物区系在整体上具有温带性质,但在干旱河谷地带,热带和温带区系成分的比例相当：热带成分的构成和分布反映古热带和古地中海区系的残遗性影响;2)东亚(含亚型)和东亚-北美成分对贡嘎山中部森林植物区系的影响最大,这些成分以温带古老性质为主;3)北温带成分是贡嘎山植物区系的主体之一,对青藏高原隆升以来贡嘎山植物区系进化类群和特有成分的发展有主要贡献,代表区系的年轻组分;4)中国特有种类型多样,占不同垂直植被带物种数量的40％-65％,其比例随海拔上升而增大。各类型比例的垂直变化突出反映了贡嘎山及横断山脉中海拔地段的植物区系与华中地区的联系,以及高海拔地段与青藏高原及东喜马拉雅的区系之间的联系。本文还就贡嘎山在生物地理分布上的意义以及贡嘎山和横断山脉植物区系特有性的性质进行了讨论。     

云南蕨类植物的物种多样性和区系组成

在一定实地调查和分类的基础上,采用统计分析及区系地理分析法,对云南地区蕨类植物进行研究。结果表明:云南地区共有蕨类植物60科193属1 530种(包括变种和变型),科、属、种分别占中国蕨类植物科总数的58.8%,属的83.9%和种的95.2%;在这60个科中含30种以上的有13个,含5个属以上的有11个,分别占该蕨类植物区系属、种数的56.5%和79.2%,特别是鳞毛蕨科、蹄盖蕨科、水龙骨科和金星蕨科,这4个科共拥有70属761种,分别占总属数、总种数的36.3%、49.7%;193属中含30种以上的有11个,分别占总属、总种数的5.7%和41.7%,最具优势的是鳞毛蕨属和耳蕨属、蹄盖蕨属、铁角蕨属、卷柏属,这5个属共有403种,占总种数的26.3%;该区系中无云南特有科,但具有4个亚洲特有科:雨蕨科、稀子蕨科、柄盖蕨科和骨碎补科;该区系以热带、亚热带性质为主,科的区系成分中热带亚热带分布的科占66.6%,热带成分的属有112个,占68.7%(除世界分布类型);该区系是东亚地区蕨类植物区系的重要组成部分,东亚分布31属和中国特有分布6属,分别占总属数(除世界分布属外)的19.0%和3.7%;该地区蕨类植物区系与西藏、台湾具有共同的区系起源和物种分化形成的背景,属的相似性系数约为70%,种的相似性系数约为30%,均起源于热带亚热带地区;属的热带区系成分与温带区系成分所占比例分别为68.7%和27.6%,存在一定的区系过渡性;该区系科的分化强度为3.2,属的分化强度为7.9,在科、属水平上均表现出较强的区系分化特征。     

Diversity, distribution and dispersal of Antarctic terrestrial algae

Terrestrial algae have been studied at widespread Antarctic localities. However, their diversity is not fully known as often collections have not been made from all habitats and techniques have been inadequate for recognition of the total flora. Identifications can be unreliable and are often left at generic level. Despite this it seems that they largely comprise cosmopolitan species but at a reduced diversity relative to other regions. There is a small element of endemic species which, in general, differ only slightly from related species elsewhere. Lack of base-line taxonomic knowledge, detailed characterization of environmental factors, and application of multivariate analysis restricts our ability to define communities and interpret their distribution patterns. Examples are provided where understanding would be greatly improved by more detailed analyses. Dispersal barriers could be operating both to and within Antarctica. However, local dispersal by wind seems readily accomplished. Overcoming dispersal barriers to Antarctica, and between ice-free localities within Antarctica, could present a greater problem. Investigation of long-distance transport of propagules by wind, birds and humans is needed.     

Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia

Recent data revealed that metazoans such as mites and springtails have persisted in Antarctica throughout several glacial–interglacial cycles, which contradicts the existing paradigm that terrestrial life was wiped out by successive glacial events and that the current inhabitants are recent colonizers. We used molecular phylogenetic techniques to study Antarctic microchlorophyte strains isolated from lacustrine habitats from maritime and continental Antarctica. The 14 distinct chlorophycean and trebouxiophycean lineages observed point to a wide phylogenetic diversity of apparently endemic Antarctic lineages at different taxonomic levels. This supports the hypothesis that long-term survival took place in glacial refugia, resulting in a specific Antarctic flora. The majority of the lineages have estimated ages between 17 and 84 Ma and probably diverged from their closest relatives around the time of the opening of Drake Passage (30–45 Ma), while some lineages with longer branch lengths have estimated ages that precede the break-up of Gondwana. The variation in branch length and estimated age points to several independent but rare colonization events.     

Surviving out in the cold: Antarctic endemic invertebrates and their refugia

Aim To identify Antarctic palaeoendemic taxa and their probable glacial refugia from regional groups of endemic species records. Location Antarctica. Methods We compiled a list of Antarctic non‐marine invertebrates from published literature, and then deleted all records relating to non‐endemic, zoochoric (phoretic and parasitic), marine and partially identified species to leave only the elements endemic to Antarctica. We then used cluster analysis and principal components analysis to identify regional groupings within this endemic fauna. Results Some 170+ of the reported 520+ Antarctic invertebrates are free‐living and endemic, but only nine of these are pan‐Antarctic, with the majority having either ‘continental’/eastern or ‘maritime’/western distributions. Main conclusions All invertebrates endemic to continental Antarctica are confined to, or found adjacent to, ice‐free palaeorefugial mountains, nunataks and coastal exposures. By contrast, only one maritime Antarctic palaeorefugium has been identified, and most endemic taxa are currently associated with coastal lowland neorefugia. We suggest which regions of Antarctica (1) are likely to be refugial, and (2) simply require more data in order that the nature and origin of their fauna can be elucidated.     

The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

Diversity and genomics of Antarctic marine micro-organisms

Marine bacterioplanktons are thought to play a vital role in Southern Ocean ecology and ecosystem function, as they do in other ocean systems. However, our understanding of phylogenetic diversity, genome-enabled capabilities and specific adaptations to this persistently cold environment is limited. Bacterioplankton community composition shifts significantly over the annual cycle as sea ice melts and phytoplankton bloom. Microbial diversity in sea ice is better known than that of the plankton, where culture collections do not appear to represent organisms detected with molecular surveys. Broad phylogenetic groupings of Antarctic bacterioplankton such as the marine group I Crenarchaeota, alpha-Proteobacteria (Roseobacter-related and SAR-11 clusters), gamma-Proteobacteria (both cultivated and uncultivated groups) and Bacteriodetes-affiliated organisms in Southern Ocean waters are in common with other ocean systems. Antarctic SSU rRNA gene phylotypes are typically affiliated with other polar sequences. Some species such as Polaribacter irgensii and currently uncultivated gamma-Proteobacteria (Ant4D3 and Ant10A4) may flourish in Antarctic waters, though further studies are needed to address diversity on a larger scale. Insights from initial genomics studies on both cultivated organisms and genomes accessed through shotgun cloning of environmental samples suggest that there are many unique features of these organisms that facilitate survival in high-latitude, persistently cold environments.     

A critical review of Antarctic Conoidea (Neogastropoda)

ABSTRACT

The Antarctic Conoidean fauna is critically reviewed based on published data and specimens in the collections of the USNM, IORAS and MNHN. Forty-two species and subspecies of the superfamily Conoidea are recorded as occurring within the Antarctic Convergence (excluding the fauna of the Kerguelen Islands) and are attributed to 14 genera and seven families. These include the new taxa: Antarctospira n. gen. (type species—Leucosyrinx badenpowelli Dell, 1990); Drilliola antarctica n. sp.; Pleurotomella (Pleutoromella) tippetti n. sp.; Pleurotomella (Anomalotomella) petiti n. sp.; Xanthodaphne pastorinoi n. sp. Aforia watsoni is introduced as a new name for Pleurotoma (Surcula) lepta Watson, 1881, non Pleurotoma lepta Edwards, 1861. A lectotype is designated for Conorbella antarctica (Strebel, 1908). New combinations are also proposed. Antarctospira badenpowelli (Dell, 1990), n. comb. (previously assigned to Leucosyrinx); Antarctospira principalis (Thiele, 1912), n. comb. (previously assigned to Typhlomangelia); Antarctospira mawsoni (Powell, 1958), n. comb. (previously assigned to Leucosyrinx); Typhlodaphne paratenoceras (Powell, 1951), n. comb. (previously assigned to Leucosyrinx); Belalora weirichi (Engl, 2008), n. comb. (previously assigned to Oenopota); Pleurotomella (Anomalotomella) innocentia (Dell, 1990), n. comb. (previously assigned to Typhlodaphne); Pleurotomella (Anomalotomella) nipri (Numanami, 1996), n. comb. (previously assigned to Typhlodaphne); Xanthodaphne raineri (Engl, 2008), n. comb. (previously assigned to Pleurotomella); Aforia hedleyi (Dell, 1990), n. comb. (previously assigned to Pontiothauma). The majority of Antarctic conoidean taxa have hypodermic marginal teeth. Although there is a similar relative abundance of conoideans in Antarctic waters to that seen in other well-studied faunas, the low number of conoideans is indicative of the general impoverishment of the gastropod fauna in the region. Fourteen percent (2 of 14) of conoidean genera that occur within the Antarctic Convergence are endemic to Antarctic waters, as are 82% (34 of 42) of the species. Most taxa have very broad bathymetric ranges, some extending from bathyal to hadal depths. The greatest species diversity was at bathyal depths.     

Areas of endemism and patterns of diversity for aphids of the Qinghai-Tibetan Plateau and the Himalayas

Aim  The study aimed to identify areas of endemism for aphids in the Qinghai-Tibetan Plateau and the Himalayas (QTPH), and to test congruence between patterns of endemism and patterns of overall species richness identified in a previous study.
Location  The QTPH.
Methods  A distribution data base of 326 endemic aphids in the QTPH was compiled. The study area was divided into a grid of 2°× 2° operative geographical units. Parsimony analysis of endemicity (PAE) was used to identify areas of endemism, and the diversity patterns of endemic species were then mapped using GIS.
Results  We identified 326 endemic species belonging to 138 genera within Adelgidae and 14 subfamilies of Aphididae. Five areas of endemism were found using PAE analysis: the eastern Himalayas, the western Himalayas, north-western Yunnan, southern Tibet and the eastern QTPH. Maps of patterns of endemism identified four major centres for endemic aphids, namely the western Himalayas, the eastern Himalayas (or Sikkim-Assam Himalayas), north-western Hengduan Mountains and the mountains of southern Gansu Province, and three minor centres, southern Tibet, south-eastern Tibet and the eastern Qinghai Province in the north-eastern QTPH.
Main conclusions  Our study identifies major centres of aphid endemism. Furthermore, there is a noticeable congruence between patterns of endemism and patterns of species richness. The patterns of endemism were most likely influenced by the recent uplift of the QTPH.     

The influence of past and present climate on the biogeography of modern mammal diversity

Within most terrestrial groups of animals, including mammals, species richness varies along two axes of environmental variation, representing energy availability and plant productivity. This relationship has led to a search for mechanistic links between climate and diversity. Explanations have traditionally focused on single mechanisms, such as variation in environmental carrying capacity or evolutionary rates. Consensus, though, has proved difficult to achieve and there is growing appreciation that geographical patterns of species richness are a product of many interacting factors including biogeographic history and biological traits. Here, we review some current hypotheses on the causes of gradients in mammal richness and range sizes since the two quantities are intimately linked. We then present novel analyses using recent datasets to explore the structure of the environment-richness relationship for mammals. Specifically, we consider the impact of glaciation on present day mammalian diversity gradients. We conclude that not only are multiple processes important in structuring diversity gradients, but also that different processes predominate in different places.     

The biogeography of the Anura of sub-equatorial Africa and the prioritisation of areas for their conservation

There is an increasing need for protected areas to conserve biodiversity efficiently. The Anura of sub-equatorial Africa have received little attention, but we quantitatively analyse a database containing presence-only data for anurans of sub-equatorial Africa to determine patterns of distribution and species richness, and discuss the roles of present and past environmental conditions in shaping these patterns. We consider the distribution of areas rich in endemic, range-restricted and Red Data Book (RDB) species to identify areas of significance to conservation. The Eastern Highlands of Zimbabwe and adjacent area in Mozambique, southeastern Malawi and the northern coast of KwaZulu/Natal are particularly species rich, whereas the southwestern Cape of South Africa and northwestern Zambia exhibit high degrees of endemism. Four major biogeographical sub-regions are identified, which can be further subdivided into provinces. All statistically significant, current environmental factors together account for 52.6% of species richness. Annual maximum rainfall, soil type variation, minimum temperature and range of elevation were all positively correlated with species richness. Thus, both habitat influences and history appear to have influenced patterns of anuran richness in the region. Generally, areas of high species richness coincide with those high in range-restricted, endemic and RDB species. In South Africa, the northeastern coast and southwestern Cape are hypothesised to have been both refugia and centres of speciation. Results suggest that the current reserve system in sub-equatorial Africa is inadequate for the conservation of the full complement of anuran species in the region.