Chapter

Refuges of Antarctic diversity

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Abstract

Antarctic terrestrial ecosystems contribute a tiny proportion of the area of the continent and host an impoverished and often cryptic biota. In recent years it has been realized that much of this biota is unique to the continent, carrying signals of its evolutionary radiation on multi-million-year timescales, some even pre-dating the final breakup of Gondwana and the geographic isolation of Antarctica. However, for terrestrial life to have existed continuously on the continent over these timescales, appropriate ice-free land must have existed through the multiple glacial cycles that took place throughout the Miocene, Pliocene and Pleistocene eras. This challenges current glaciological reconstructions, which present a model of complete obliteration of most currently ice-free areas of ground at successive glacial maxima, with those remaining not providing viable refugia for the majority of the contemporary terrestrial biota. In this chapter, we consider the requirement for refugia across all regions of Antarctica, and the likely form that such refugia may have taken.

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... In contrast, Antarctica's paleoclimatic legacy for terrestrial communities has long been considered one of widespread extinction due to glaciation. Evidence of terrestrial species surviving in Antarctic glacial refugia (6) and discoveries of substantial endemic diversity and biogeographic structuring in some groups (7,8) is changing this narrative, indicating extended evolutionary histories on land. Yet, such evolutionary histories remain obscured by a lack of large-scale molecular phylogenetic work, with most Antarctic terrestrial research focused on small subsets of species or populations (9,10). ...
... The hypothesis that diversification has proceeded similarly in Antarctic marine and terrestrial groups has not been tested. While the extinction of a diverse continental Antarctic biota is well established (13), mounting evidence of significant and biogeographically structured Antarctic terrestrial diversity (8,14,15) with a long evolutionary history (6,16) suggests the possibility of broadly similar diversification processes across marine and terrestrial Antarctic systems. If valid for some taxa, further tests should then be sought across a wider variety of organisms. ...
... While in some respects quite different to the continental Antarctic, the islands are in other ways quite similar, providing a window into diversification processes that might be sought for continental groups, especially given their age and biogeographic structuring. Both regions share many higher taxa (e.g., ref. 25), a dynamic geo-climatic history (6,26), a profound degree of isolation, and indications that climatic events likely structured their biota (6,8,27). The terrestrial habitat on the continent and its surrounding islands is fragmented by large expanses of ice or ocean, respectively, and has been further isolated by the Antarctic Circumpolar Current for at least 34 Ma (28,29). ...
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Global cooling and glacial–interglacial cycles since Antarctica’s isolation have been responsible for the diversification of the region’s marine fauna. By contrast, these same Earth system processes are thought to have played little role terrestrially, other than driving widespread extinctions. Here, we show that on islands along the Antarctic Polar Front, paleoclimatic processes have been key to diversification of one of the world’s most geographically isolated and unique groups of herbivorous beetles—Ectemnorhinini weevils. Combining phylogenomic, phylogenetic, and phylogeographic approaches, we demonstrate that these weevils colonized the sub-Antarctic islands from Africa at least 50 Ma ago and repeatedly dispersed among them. As the climate cooled from the mid-Miocene, diversification of the beetles accelerated, resulting in two species-rich clades. One of these clades specialized to feed on cryptogams, typical of the polar habitats that came to prevail under Miocene conditions yet remarkable as a food source for any beetle. This clade’s most unusual representative is a marine weevil currently undergoing further speciation. The other clade retained the more common weevil habit of feeding on angiosperms, which likely survived glaciation in isolated refugia. Diversification of Ectemnorhinini weevils occurred in synchrony with many other Antarctic radiations, including penguins and notothenioid fishes, and coincided with major environmental changes. Our results thus indicate that geo-climatically driven diversification has progressed similarly for Antarctic marine and terrestrial organisms since the Miocene, potentially constituting a general biodiversity paradigm that should be sought broadly for the region’s taxa.
... The complex history of Antarctic colonization here recorded in B. argenteum and its persistence through glacial periods within the continent complements recent observations in multiple groups of terrestrial biota (Biersma et al., 2017;Convey et al., 2008Convey et al., , 2018Convey et al., , 2020McGaughran, Tearuds, Convey, & Fraser, 2019), including many invertebrate groups, lichens, and other mosses that currently occur in habitats and areas that are also commonly characterized by B. argenteum. These important components of the Antarctic biota must therefore have had environmentally suitable refugia available to them (Convey et al., , 2020. ...
... The complex history of Antarctic colonization here recorded in B. argenteum and its persistence through glacial periods within the continent complements recent observations in multiple groups of terrestrial biota (Biersma et al., 2017;Convey et al., 2008Convey et al., , 2018Convey et al., , 2020McGaughran, Tearuds, Convey, & Fraser, 2019), including many invertebrate groups, lichens, and other mosses that currently occur in habitats and areas that are also commonly characterized by B. argenteum. These important components of the Antarctic biota must therefore have had environmentally suitable refugia available to them (Convey et al., , 2020. However, the precise locations of such refugia remain unknown , as does the degree of connectivity or isolation between individual locations. ...
... within bank-forming mosses over a period of up to two millennia preserved in permafrost. In Victoria Land, cold-based glaciers were widespread between c. 5-7 Ma and 2.5 Ma (Di Nicola et al., 2012;Smellie et al., 2014), providing the possibility of survival during glaciations through cryptobiosis, although it has to be accepted that the duration of glacial periods (tens to hundreds of thousands of years) is considerably longer than any currently available demonstration of cryptobiotic survival other than in microorganisms (Convey et al., 2020). ...
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The dispersal routes of taxa with transoceanic disjunctions remain poorly understood, with the potential roles of Antarctica not yet demonstrated. Mosses are suitable organisms to test direct intra‐Antarctic dispersal, as major component of the extant Antarctic flora, with the cosmopolitan moss Bryum argenteum as ideal target species. We analyzed the genetic structure of B. argenteum to provide an evolutionary time frame for its radiation and shed light into its historical biogeography in the Antarctic region. We tested two alternative scenarios: (a) intra‐Antarctic panmixia and (b) intra‐Antarctic genetic differentiation. Furthermore, we tested for evidence of the existence of specific intra‐Antarctic dispersal routes. Sixty‐seven new samples (40 collected in Antarctica) were sequenced for ITS nrDNA and rps4 cpDNA regions, and phylogenetic trees of B. argenteum were constructed, with a focus on its Southern Hemisphere. Combining our new nrDNA dataset with previously published datasets, we estimated time‐calibrated phylogenies based on two different substitution rates (derived from angiosperms and bryophytes) along with ancestral area estimations. Minimum spanning network and pairwise genetic distances were also calculated. B. argenteum was potentially distributed across Africa and Antarctica soon after its origin. Its earliest intra‐Antarctic dispersal and diversification occurred during a warming period in the Pliocene. On the same timescale, a radiation took place involving a dispersal event from Antarctica to the sub‐Antarctic islands. A more recent event of dispersal and diversification within Antarctica occurred during a warm period in the Pleistocene, creating favorable conditions also for its colonization outside the Antarctic continent worldwide. We provide evidence supporting the hypothesis that contemporary populations of B. argenteum in Antarctica integrate a history of both multiple long‐range dispersal events and local persistence combined with in situ diversification. Our data support the hypothesis that B. argenteum has been characterized by strong connectivity within Antarctica, suggesting the existence of intra‐Antarctic dispersal routes.
... The evolutionary history of Antarctic springtails has been studied in increasing detail using molecular tools over the last two decades [30][31][32][33] and a growing consensus has been obtained for their long-term persistence in Antarctica and survival in local refugia during glacial maxima [34,35]. Present-day populations are the descendants of lineages that colonized suitable environments up to many millions of years ago, subsequently persisting through multiple and intense periods of glaciation [36,37]. ...
... It is also not long before the very extensive glaciations in the Miocene, around 12-14 Mya, which are associated with the final extinction of remaining "tundra" communities in the Transantarctic Mountains [37,45]. ...
... An alternative, systematic source of uncertainty may lay in the use of calibration points that are very distant, both in terms of time and taxonomy, from the group of interest. This cannot be discounted in the current analysis, as available calibration points within Collembola are limited to two Palaeozoic nodes [37,[46][47][48] for a compilation of fossil Collembola that may be deployed provided adequate taxon sampling is available). Future dating analyses specifically focused on Antarctic divergences are clearly required, although the associated difficulties (e.g., the availability of appropriate calibration points that do not themselves derive from phylogeographic interpretations, considering that the resulting dates are to be used to test phylogeographic hypotheses) at present preclude this possibility. ...
Article
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Springtails and mites are the dominant groups of terrestrial arthropods in Antarctic terrestrial ecosystems. Their Antarctic diversity includes a limited number of species, which are frequently endemic to specific regions within the continent. Advances in molecular techniques, combined with the re-evaluation of morphological characters and the availability of new samples, have recently led to the identification of a number of new springtail species within previously named, but ill-defined, species entities described in the last century. One such species, the neanurid Friesea grisea, originally described from sub-Antarctic South Georgia, was for many years considered to be the only known springtail with a pan-Antarctic distribution. With the recent availability of new morphological and molecular data, it has now been firmly established that the different representatives previously referred to this taxon from the Antarctic Peninsula and Victoria Land (continental Antarctica) should no longer be considered as representing one and the same species, and three clearly distinct taxa have been recognized: F. antarctica, F. gretae and F. propria. In this study, the relationships among these three species are further explored through the sequencing of the complete mtDNA for F. gretae and the use of complete mitogenomic as well as cytochrome c oxidase I data. The data obtained provide further support that distinct species were originally hidden within the same taxon and that, despite the difficulties in obtaining reliable diagnostic morphological characters, F. gretae is genetically differentiated from F. propria (known to be present in different locations in Northern Victoria Land), as well as from F. antarctica (distributed in the Antarctic Peninsula).
... The Antarctic terrestrial vegetation is assembled in extensive tufts over 0.3% of the permanent ice-free surface (Convey et al., 2020). The main components are lichens, which comprise 65% of the total species identified (Casanovas et al., 2015). ...
... The successful survival of this group is facilitated by their poikilohydric nature because their hydration status tends to be in equilibrium with the environment, where they are metabolically active when hydrated and dormant when dry. Dormancy constitutes an abiotic stress avoidance strategy that ensures the survival of these species . This response trait (any trait that varies in response to changes in environmental conditions according to Violle et al. (2007)) is widespread in various organisms, such as mosses, nematodes, tardigrades, rotifers, and collembolans (Convey et al., 2020). ...
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Poikilohydric autotrophs are the main colonizers of the permanent ice-free areas in the Antarctic tundra biome. Global climate warming and the small human footprint in this ecosystem make it especially vulnerable to abrupt changes. Elucidating the effects of climate change on the Antarctic ecosystem is challenging because it mainly comprises poikilohydric species, which are greatly influenced by microtopographic factors. In the present study, we investigated the potential effects of climate change on the metabolic activity and net primary photosynthesis (NPP) in the widespread lichen species Usnea aurantiaco-atra. Long-term monitoring of chlorophyll a fluorescence in the field was combined with photosynthetic performance measurements in laboratory experiments in order to establish the daily response patterns under biotic and abiotic factors at micro- and macro-scales. Our findings suggest that macroclimate is a poor predictor of NPP, thereby indicating that microclimate is the main driver due to the strong effects of microtopographic factors on cryptogams. Metabolic activity is also crucial for estimating the NPP, which is highly dependent on the type, distribution, and duration of the hydration sources available throughout the year. Under RCP 4.5 and RCP 8.5, metabolic activity will increase slightly compared with that at present due to the increased precipitation events predicted in MIROC5. Temperature is highlighted as the main driver for NPP projections, and thus climate warming will lead to an average increase in NPP of 167–171% at the end of the century. However, small changes in other drivers such as light and relative humidity may strongly modify the metabolic activity patterns of poikilohydric autotrophs, and thus their NPP. Species with similar physiological response ranges to the species investigated in the present study are expected to behave in a similar manner provided that liquid water is available.
... The Thala Hills terrestrial biota includes a high proportion of endemic species (Fig. 5), a feature that is being increasingly strongly recognized across Antarctica , Convey et al. 2020. Most of the lichens present in Thala Hills are widely represented in polar vegetation and are either Antarctic endemics or have bipolar distributions. ...
... Where such studies have been applied (e.g. Iakovenko et al. 2015), the general conclusion is that Antarctic representatives are clearly distinct from other relatives globally, as is the case in other groups of terrestrial fauna (reviewed by Convey et al. 2020). There is a similar lack of up-to-date taxonomic reconsideration of tardigrades across much of Antarctica. ...
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Knowledge of the biodiversity of the Thala Hills oasis (Enderby Land, East Antarctica) is very limited. Here, we integrate all information available since 1962, when the Russian ‘Molodyozhnaya’ station was established in the western part of the oasis. The published data on local eukaryote diversity (lichens, embryophytes, metazoans) include records of 90 species. Since 2008, Belarusian Antarctic Expedition researchers have worked in the eastern part of the oasis, accessible from the Belarusian station ‘Vechernyaya Mount'. This research revealed 95 species, including 44 species not recorded in the earlier published literature. The level of available information is uneven across major taxa. Lichens are the better-known group, with 51 species recorded in total, including 13 species recently recorded for the first time in the oasis. New records were also obtained for rotifers. Thala Hills biodiversity is consistent with wider patterns of Antarctic biogeography, with a high proportion of regionally endemic species (especially metazoans), the occurrence of both endemic and bipolar species of lichens and generally low numbers of cosmopolitan species (largely limited to aquatic rotifers, with the caveat that up-to-date taxonomic studies are required). The lack of data on marine macrobenthos, soil nematodes and terrestrial rotifers emphasizes the need for studies focusing on these groups.
... Over the last two decades, rapid advances in classical biogeographic and molecular phylogeographic research have led to a paradigm change in our understanding of the origin and antiquity of the Antarctic terrestrial biota . Studies carried out across all ice-free regions of the continent, and all the major terrestrial taxonomic groups, have confirmed that long-term presence in these regions is the norm, requiring persistence of both the organisms and terrestrial habitats through the multiple cycles of glaciation Antarctica has experienced (see Convey et al., 2020, for comprehensive review). Some phylogeographic studies have identified signals of persistence since the final breakup of Gondwana and separation of Antarctica. ...
Chapter
Antarctica's contemporary terrestrial biota have adapted to the continent's environmental challenges over many millions of years. It now faces the twin challenges of the multiple aspects of global and regional environmental change and the direct impacts of human presence and activity. Most native terrestrial biota possess wide physiological and ecological flexibility, well beyond the expected scale of environmental change over the next century and, when considered in isolation, are likely to show positive responses to environmental ameliorations, particularly the combination of increased thermal energy and liquid water availability, through increased production, populations, local distribution extent, and community complexity. However, over this timescale other, direct, human impacts on Antarctic terrestrial ecosystems and biodiversity are likely to have greater and negative consequences. Major sources of such impacts are likely to come through physical damage to the limited available area of terrestrial habitats, and the anthropogenic introduction of non-native species, a proportion of which are likely to become invasive and act as ecosystem engineers.
... Recent climate and glaciological modeling studies have highlighted greater dynamism in glacial extent than previously considered possible throughout the early Pliocene and Pleistocene (Scherer et al., 2008;Naish et al., 2009;Pollard and DeConto, 2009;De Schepper et al., 2014;DeConto and Pollard, 2016), suggesting the possibility of ice-free local refugial areas persisting throughout these periods (e.g. as suggested by Fraser et al., 2014). Molecular, phylogenetic and biogeographic studies also suggest in situ survival for many groups of terrestrial fauna in Antarctica throughout the Quaternary, Neogene and even Paleogene (see Convey et al., 2008Convey et al., , 2009Convey et al., , 2020. Recently, increased evidence has also been found of million-year persistence of the Antarctic flora, e.g. ...
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Biogeographic patterns of globally widespread species are expected to reflect regional structure, as well as connectivity caused by occasional long-distance dispersal. We assessed the level and drivers of population structure, connectivity, and timescales of population isolation in one of the most widespread and ruderal plants in the world — the common moss Ceratodon purpureus. We applied phylogenetic, population genetic, and molecular dating analyses to a global (n = 147) sampling data set, using three chloroplast loci and one nuclear locus. The plastid data revealed several distinct and geographically structured lineages, with connectivity patterns associated with worldwide, latitudinal “bands.” These imply that connectivity is strongly influenced by global atmospheric circulation patterns, with dispersal and establishment beyond these latitudinal bands less common. Biogeographic patterns were less clear within the nuclear marker, with gene duplication likely hindering the detection of these. Divergence time analyses indicated that the current matrilineal population structure in C. purpureus has developed over the past six million years, with lineages diverging during the late Miocene, Pliocene, and Quaternary. Several colonization events in the Antarctic were apparent, as well as one old and distinct Antarctic clade, possibly isolated on the continent since the Pliocene. As C. purpureus is considered a model organism, the matrilineal biogeographic structure identified here provides a useful framework for future genetic and developmental studies on bryophytes. Our general findings may also be relevant to understanding global environmental influences on the biogeography of other organisms with microscopic propagules (e.g., spores) dispersed by wind.
... nematodes, tardigrades, mites and mosses) was proposed to feed into conservation planning and management (Terauds et al. 2012, Terauds andLee 2016). Proper conservation planning and management of the Antarctic is urgent for protecting its often endemic biota (Convey et al. 2014(Convey et al. , 2020 as most of them have narrow distributional ranges due to the long-term survival of taxa in isolated glacial refugia during ice ages , Fraser et al. 2014, Biersma et al. 2018). In the sub-Antarctic, isolation has long been thought to underpin patterns in its terrestrial biodiversity leading to three distinct provinces (Smith 1984), namely the South Atlantic, South Indian and South Pacific Provinces. ...
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Terrestrial biota in the Antarctic are more globally distinct and highly structured biogeographically than previously believed, but information on biogeographic patterns and endemism in freshwater communities is largely lacking. We studied biogeographic patterns of Antarctic freshwater diatoms based on the analysis of species occurrences in a dataset of 439 lakes spread across the Antarctic realm. Highly distinct diatom floras, both in terms of composition and richness, characterize Continental Antarctica, Maritime Antarctica and the sub‐Antarctic islands, with marked biogeographic provincialism in each region. A total of 44% of all species is estimated to be endemic to the Antarctic, and most of them are confined to a single biogeographic region. The level of endemism significantly increases with increasing latitude and geographic isolation. Our results have implications for conservation planning, and suggest that successful dispersal of freshwater diatoms to and within the Antarctic is limited, fostering the evolution of highly endemic diatom floras.
... These regions experience very different environmental conditions that strongly impact on the composition and functioning of their terrestrial ecosystems. Moreover, their microbiota and invertebrate fauna have long evolutionary histories in isolation 20,34,35 . The reduced diversity and richness of Antarctic springtail microbiomes, as well as the limited similarities observed between the four microbiomes analysed here, could be the result of strong environmental control, as well as of millions of years of evolutionary separation. ...
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Collembola are a key component of the soil biota globally, playing an important role in community and ecosystem dynamics. Equally significant are their associated microbiomes, that can contribute to key metabolic functions. In the present study, we investigated the bacterial community composition of four Antarctic springtail species to assess if and how the extreme Antarctic environment has shaped the collembolans’ microbiomes. Springtails were collected from two biogeographical regions, the maritime and the continental Antarctic. From each region, two endemic species, belonging to the genera Cryptopygus (Isotomidae, Entomobryomorpha) and Friesea (Neanuridae, Poduromorpha), were included. This experimental design allowed us to quantify the relative importance of ecological factors (different regions of occurrence) and/or phylogenetic divergence in the host (different Orders) in shaping the Collembola microbiome. The diversity and richness of springtail microbiomes was lower in the Antarctic taxa compared to published information from species from temperate regions. The microbiome composition was predominantly species-specific, with a limited core microbiome shared across the four species examined. While both geographic origin and host species influenced the associated microbiomes, the former was the prevalent driver, with closer similarity between springtails from the same bioregion than between those belonging to the same genus.
... But nunataks also have important implications for post-glacial geomorphic and environmental dynamics during the paraglacial phase, such as for vegetation colonization (Ruiz-Fern andez et al., 2019). They have been suggested to provide refugia to biodiversity during past glaciations, thus being pioneering sites for vegetation re-colonization following glacial retreat (Jørgensen et al., 2012) and key spots for species dispersal across migration routes from ice-free areas in the Maritime Antarctica to the interior of the continent (Convey et al., 2020). Therefore, a more accurate picture of the age of deglaciation and nunatak development in the Antarctic Peninsula region can be useful to better understand the pattern of colonization. ...
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The small ice caps distributed across the Antarctic Peninsula region have undergone large ice volume changes since the Last Glacial Cycle, in line with most of the Antarctic continent. While the surface extent of glacial shrinking is relatively well known, the timing of glacial oscillations and the magnitude of ice thinning remain little investigated. Cosmic-Ray Exposure (CRE) dating applied on ice-free vertical sequences can provide insights about the temporal framework of glacial oscillations. However, the potential occurrence of nuclide inheritance may overestimate the real timing of the last glacial retreat. This problem has been observed in many areas in Continental Antarctica, but similar studies have not yet been conducted in environments of the Maritime Antarctica, such as the South Shetland Islands (SSI). This research focuses on the Hurd Peninsula ice cap (HPIC, ca. 60°22′ W, 62°40’ S), located in the SW of Livingston Island, SSI. Past climate oscillations since the Last Glacial Cycle have determined the amount of ice stored in the ice cap. Today, this polythermal ice cap is surrounded by several nunataks standing out above the ice. Three of them have been selected to explore their deglaciation history and to test the potential occurrence of nuclide inheritance in deglaciated bedrocks associated with polythermal glaciers. We present a new dataset with 10 10Be exposure dates. Some of them were found to be anomalously old, evidencing that nuclide inheritance is present in bedrocks associated with polythermal ice caps and suggesting complex glacial exposure histories. We attribute this to limited erosion, given the gentle slope of the nunatak margins and the cold-based character of the surrounding ice. The remaining samples allowed to approach local surface-elevation changes of the HPIC. Our results suggest that ice thinning started during the Last Glacial Maximum (LGM) at ∼22 ka but intense glacial shrinking occurred from ∼18 to ∼13 ka, when the nunataks became exposed, being particularly intense at the end of this period (∼14–13 ka) coinciding with the time of the meltwater pulse 1a (MWP-1a) and the end of the Antarctic Cold Reversal (ACR).
... En Isla Decepción, de origen volcánico, los estudios de la diversidad y desarrollo de la vegetación se han centrado en zonas de actividad geotérmica, donde la combinación de calor, vapor, inestabilidad del sustrato y concentraciones de nutrientes a menudo insignificantes o con niveles tóxicos de gases sulfurosos y depósitos minerales, imponen severas restricciones a la biota vegetal y liquénica (Smith 2005). A su vez, estos sectores ligados a un activo vulcanismo y elevada actividad geotérmica pueden haber jugado un papel determinante en la colonización vegetal del continente antártico (Convey et al. 2020), por lo que una caracterización precisa y detallada de la distribución de especies vegetales puede ayudar en una mejor comprensión de los patrones espacio-temporales de expansión de la vegetación de la Antártica. Desde el punto de vista botánico, Isla Decepción es particularmente importante, pues alberga un gran número de especies poco comunes de la Antártica, entre ellas musgos, hepáticas, y líquenes (Smith 2005). ...
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Con el objetivo de determinar la influencia de las pingüineras sobre la diversidad de la vegetación en la Isla Decepción, se estudió la composición de briófitas de un transecto de aproximadamente 2 km entre el Lago Irízar y la pingüinera (Pygoscelis antarcticus) de Punta La Descubierta. Fueron detectados un total de 39 carpetas de vegetación formadas principalmente por briófitos, distribuidas en tres sectores principales, aledaño al Lago Irízar, en Collado Vapor y en Punta La Descubierta. Los briófitos registrados corresponden a 15 especies de musgos y sólo 2 hepáticas, con 11 familias representadas. Se detectó que dos musgos, Sanionia uncinata y Politrychastrum alpinum, con 31 y 9 registros, dominaban las carpetas, con mayor abundancia y frecuencia. Además, se encontró un nuevo registro para la isla del musgo Bryum orbiculatifolium, el cual crece directamente asociado a la pingüinera. Además, se encontró que hay 5 especies de musgos comunes a los tres sectores, aunque otras 5 especies crecen solo en el sector de la pingüinera, diferenciándose esta comunidad de las otras dos. No se detectaron plantas vasculares en todo el sitio de estudio, por lo que se discute la posibilidad de que estos sitios se encuentran en estados de colonización temprana y donde su biota está marcada por la presencia de musgos pioneros que crecen alrededor de las pingüineras, influenciadas probablemente por el aporte de nutrientes del guano depositado.
... Green algae (Viridiplantae) are noted in all studies of Antarctic lake algal communities, represented mainly by the Phylum Chlorophyta [12], with zygnematophycean green algae and other taxa phylogenetically close to higher plants (Streptophyta) being less common. De Wever et al. [8] highlighted the wide phylogenetic diversity of apparently endemic Antarctic lineages of microscopic green algae, consistent with hypotheses of strong regionalization and long-term evolutionary isolation within Antarctica, even of microbial diversity [13][14][15]. According to De Wever et al. [8], their findings, supported by molecular analyses, contrasted with most previous morphological studies, which generally concluded that Antarctic green algae were mostly represented by cosmopolitan species. ...
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Background Vega Island is located off the eastern tip of the Antarctic Peninsula (Maritime Antarctica), in the Weddell Sea. In this study, we used metabarcoding to investigate green algal DNA sequence diversity present in sediments from three lakes on Vega Island (Esmeralda, Copépodo, and Pan Negro Lakes). Methods and results Total DNA was extracted and the internal transcribed spacer 2 region of the nuclear ribosomal DNA was used as a DNA barcode for molecular identification. Green algae were represented by sequences representing 78 taxa belonging to Phylum Chlorophyta, of which 32% have not previously been recorded from Antarctica. Sediment from Pan Negro Lake generated the highest number of DNA reads (11,205), followed by Esmeralda (9085) and Copépodo (1595) Lakes. Esmeralda Lake was the richest in terms of number of taxa (59), with Copépodo and Pan Negro Lakes having 30 taxa each. Bray–Curtis dissimilarity among lakes was high (~ 0.80). The Order Chlamydomonadales (Chlorophyceae) gave the highest contribution in terms of numbers of taxa and DNA reads in all lakes. The most abundant taxon was Chlorococcum microstigmatum. Conclusions The study confirms the utility of DNA metabarcoding in assessing potential green algal diversity in Antarctic lakes, generating new Antarctic records.
... Only 0.3% of the land surface is currently ice-free and suitable for terrestrial life, with most ice-free areas small, fragmented and isolated from each other (Bergstrom and Chown, 1999;Convey et al., 2008;Convey and Stevens, 2007). This long-term isolation has led to evolutionary divergence and speciation, creating the high levels of endemism characterising multiple terrestrial invertebrate groups currently present in Antarctica, including mites, springtails, rotifers and nematodes (Cakil et al., 2021;Convey et al., 2020Convey et al., , 2008Pugh and Convey, 2008;Stevens et al., 2021;Velasco-Castrillón et al., 2014). ...
Article
Antarctica has been isolated and progressively glaciated for over 30 million years, with only approximately 0.3 % of its area currently ice-free and capable of supporting terrestrial ecosystems. As a result, invertebrate populations have become isolated and fragmented, in some cases leading to speciation. Terrestrial invertebrate species currently found in Antarctica often show multi-million year, and even Gondwanan, heritage, with little evidence of recent colonisation. Mesobiotus is a globally distributed tardigrade genus. It has commonly been divided into two “groups”, referred to as harmsworthi and furciger, with both groups currently considered cosmopolitan, with global reports including from both the Arctic and the Antarctic. However, some authors considered that Meb. furciger, as originally described, may represent an Antarctic-specific lineage. Using collections of tardigrades from across the Antarctic continent and publicly available sequences obtained from online databases, we use mitochondrial and nuclear ribosomal sequence data to clarify the relationships of Antarctic Mesobiotus species. Our analyses show that all Antarctic members belong to a single lineage, evolving separately from non-Antarctic representatives. Within this Antarctic lineage there are further deep divisions among geographic regions of the continent, consistent with the presence of a species complex. Based on our data confirming the deep divisions between this Antarctic lineage, which includes representatives of both groups, we recommend that the use of furciger and harmsworthi group terminology is now abandoned, as it leads to systematic and biogeographical confusion. (Bertolani et al., 2014, Itang et al., 2020, Kaczmarek et al., 2018, Kayastha et al., 2021, Mapalo et al., 2017, Mapalo et al., 2016, Roszkowska et al., 2018, Stec, 2019, Stec et al., 2018, Stec and Kristensen, 2017; Tumanov, 2020)
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The temperature experienced by micro-invertebrates in extreme environments (such as those of Antarctica) is a pivotal parameter regarding these animals' ecology and physiology. However, at present, detailed knowledge of microhabitat physical conditions in Antarctica is limited, as well as being biased towards sub-Antarctic and Maritime Antarctic regions. To better understand the temperature conditions experienced in the microhabitats of Continental Antarctica by the native microfauna, we recorded temperatures year round in ponds and soils in an area of the Victoria Land coast and compared these measurements with air temperature data from the closest automatic weather station. We identified an important difference in temperature dynamics between the air, soil and pond datasets. Ponds were the warmest sites overall, differing by up to 7.5°C in comparison with the air temperature due to their greater thermal capacity, which also drove their patterns of freeze-thaw cycles and mean daily thermal excursion.
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Aim We investigated evolutionary relationships and biogeographical patterns within the genus Boeckella to evaluate (1) whether its current widespread distribution in the Southern Hemisphere is due to recent long-distance dispersal or long-term diversification; and (2) the age and origin of sub-Antarctic and Antarctic Boeckella species, with particular focus on the most widely distributed species: Boeckella poppei. Location South America, sub-Antarctic islands, maritime Antarctica, continental Antarctica and Australasia. Methods To reconstruct phylogenetic patterns of Boeckella, we used molecular sequence data collected from 12 regions and applied Bayesian and Maximum Likelihood analyses using multiple loci. We also estimated divergence times and reconstructed ancestral ranges using two different models of species evolution. Results Phylogenetic analyses and divergence time estimates suggested that Boeckella originated on the Gondwanan supercontinent and initially split into two main clades during the late Cretaceous (ca. 80 Ma). The first clade diversified in Australasia, and the second clade is currently distributed in South America, various sub-Antarctic islands and Antarctica. Dispersal from South America to the Kerguelen and Crozet archipelagos occurred during the Eocene/Oligocene (B. vallentini) and in the late Pliocene (B. brevicaudata), while South Georgia and the maritime Antarctic were likely colonized during the late Pleistocene (B. poppei). Main conclusions Boeckella has a Gondwanan origin, with further diversifications after the physical separation of the continental landmasses. Extant populations of Boeckella from the Scotia Arc islands and Antarctic Peninsula originated from South America during the Pleistocene, suggesting that original Antarctic Gondwanan lineages did not survive repeated glacial cycles during the Quaternary ice ages. A continuous decline in the species accumulation rate is apparent within the genus as the early Eocene, suggesting that Boeckella diversification may have decreased due to progressive cooling throughout the Cenozoic era.
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In the harsh Antarctic terrestrial ecosystems, invertebrates are currently confined to sparse and restricted ice free areas, where they have survived on multi-million-year timescales in refugia. The limited dispersal abilities of these invertebrate species, their specific habitat requirements, and the presence of geographical barriers can drastically reduce gene flow between populations, resulting in high genetic differentiation. On continental Antarctica, mites are one of the most diverse invertebrate groups. Recently, two new species of the free living prostigmatid mite genus Stereotydeus Berlese, 1901 were discovered, bringing the number of Antarctic and sub-Antarctic species of this genus up to 15, of which 7 occur along the coast of Victoria Land and in the Transantarctic Mountains. To examine the biodiversity of Stereotydeus spp., the present study combines phylogenetic, morphological and population genetic data of specimens collected from nine localities in Victoria Land. Genetically distinct intraspecific groups are spatially isolated in northern Victoria Land, while, for other species, the genetic haplogroups more often occur sympatrically in southern Victoria Land. We provide a new distribution map for the Stereotydeus species of Victoria Land, which will assist future decisions in matters of the protection and conservation of the unique Antarctic terrestrial fauna.
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The invertebrate terrestrial fauna of Antarctica is being investigated with increasing interest to discover how life interacts with the extreme polar environment and how millions of years of evolution have shaped their biodiversity. Classical taxonomic approaches, complemented by molecular tools, are improving our understanding of the systematic relationships of some species, changing the nomenclature of taxa and challenging the taxonomic status of others. The springtail Friesea grisea has previously been described as the only species with a “pan-Antarctic” distribution. However, recent genetic comparisons have pointed to another scenario. The latest morphological study has confined F. grisea to the sub-Antarctic island of South Georgia, from which it was originally described, and resurrected F. antarctica as a congeneric species occurring on the continental mainland. Molecular data demonstrate that populations of this taxon, ostensibly occurring across Maritime and Continental Antarctica, as well as on some offshore islands, are evolutionarily isolated and divergent and cannot be included within a single species. The present study, combining morphological with molecular data, attempts to validate this hypothesis and challenges the taxonomic status of F. antarctica, suggesting that two additional new species, described here as Friesea gretae sp. nov. and Friesea propria sp. nov., are present in Continental Antarctica.
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The dispersal routes of taxa with transoceanic disjunctions remain poorly understood, with the potential roles of Antarctica not yet demonstrated. Mosses are suitable organisms to test direct intra‐Antarctic dispersal, as major component of the extant Antarctic flora, with the cosmopolitan moss Bryum argenteum as ideal target species. We analyzed the genetic structure of B. argenteum to provide an evolutionary time frame for its radiation and shed light into its historical biogeography in the Antarctic region. We tested two alternative scenarios: (a) intra‐Antarctic panmixia and (b) intra‐Antarctic genetic differentiation. Furthermore, we tested for evidence of the existence of specific intra‐Antarctic dispersal routes. Sixty‐seven new samples (40 collected in Antarctica) were sequenced for ITS nrDNA and rps4 cpDNA regions, and phylogenetic trees of B. argenteum were constructed, with a focus on its Southern Hemisphere. Combining our new nrDNA dataset with previously published datasets, we estimated time‐calibrated phylogenies based on two different substitution rates (derived from angiosperms and bryophytes) along with ancestral area estimations. Minimum spanning network and pairwise genetic distances were also calculated. B. argenteum was potentially distributed across Africa and Antarctica soon after its origin. Its earliest intra‐Antarctic dispersal and diversification occurred during a warming period in the Pliocene. On the same timescale, a radiation took place involving a dispersal event from Antarctica to the sub‐Antarctic islands. A more recent event of dispersal and diversification within Antarctica occurred during a warm period in the Pleistocene, creating favorable conditions also for its colonization outside the Antarctic continent worldwide. We provide evidence supporting the hypothesis that contemporary populations of B. argenteum in Antarctica integrate a history of both multiple long‐range dispersal events and local persistence combined with in situ diversification. Our data support the hypothesis that B. argenteum has been characterized by strong connectivity within Antarctica, suggesting the existence of intra‐Antarctic dispersal routes.
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Aim Antarctica's remote and extreme terrestrial environments are inhabited by only two species of native vascular plants. We assessed genetic connectivity amongst Antarctic and South American populations of one of these species, Colobanthus quitensis, to determine its origin and age in Antarctica. Location Maritime Antarctic, sub‐Antarctic islands, South America. Taxon Antarctic pearlwort Colobanthus quitensis (Caryophyllaceae). Methods Four chloroplast markers and one nuclear marker were sequenced from 270 samples from a latitudinal transect spanning 21–68° S. Phylogeographic, population genetic and molecular dating analyses were used to assess the demographic history of C. quitensis and the age of the species in Antarctica. Results Maritime Antarctic populations consisted of two different haplotype clusters, occupying the northern and southern Maritime Antarctic. Molecular dating analyses suggested C. quitensis to be a young (<1 Ma) species, with contemporary population structure derived since the late‐Pleistocene. Main conclusions The Maritime Antarctic populations likely derived from two independent, late‐Pleistocene dispersal events. Both clusters shared haplotypes with sub‐Antarctic South Georgia, suggesting higher connectivity across the Southern Ocean than previously thought. The overall findings of multiple colonization events by a vascular plant species to Antarctica, and the recent timing of these events, are of significance with respect to future colonizations of the Antarctic Peninsula by vascular plants, particularly with predicted increases in ice‐free land in this area. This study fills a significant gap in our knowledge of the age of the contemporary Antarctic terrestrial biota. Adding to previous inferences on the other Antarctic vascular plant species (the grass Deschampsia antarctica), we suggest that both angiosperm species are likely to have arrived on a recent (late‐Pleistocene) time‐scale. While most major groups of Antarctic terrestrial biota include examples of much longer‐term Antarctic persistence, the vascular flora stands out as the first identified terrestrial group that appears to be of recent origin.
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A decrease in biodiversity and density of terrestrial organisms with increasing altitude and latitude is a well-known ecogeographical pattern. However, studies of these trends are often taxonomically-biased toward well-known organisms and especially those with relatively large bodies, and environmental variability at the local scale may perturb these general effects. Here, we focus on understudied organisms—soil invertebrates—in Antarctic deserts, which are among the driest and coldest places on Earth. We sampled two remote Antarctic sites in the Darwin Glacier area and established an altitudinal gradient running from 210 to 836 m a.s.l. We measured soil geochemistry and organic matter content and linked these parameters with the presence of soil invertebrates. We found three general outcomes, two of which are consistent with general assumptions: (a) the hostile climatic condition of the Darwin Glacier region supports an extremely low diversity of soil metazoans represented by a single nematode species—Scottnema lindsayae; (b) soil geochemistry is the main factor influencing distribution of nematodes at the local scale. Contrary to our expectations, a positive correlation was found between nematode density and altitude. This last observation could be explained by an additional effect of soil moisture as we found this increased with altitude and may be caused by orographic clouds, which are present in this region. To the best of our knowledge such effects have been described in tropical and temperate regions. Potential effect of orographic clouds on soil properties in polar deserts may be a fruitful area of ecological research on soil fauna.
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Seven species of springtail (Collembola) are present in Victoria Land, Antarctica and all have now been sequenced at the DNA barcoding region of the mitochondrial cytochrome c oxidase subunit I gene (COI). Here, we review these sequence data (n = 930) from the GenBank and Barcode of Life Datasystems (BOLD) online databases and provide additional, previously unpublished sequences (n = 392) to assess the geographic distribution of COI variants across all species. Four species (Kaylathalia klovstadi, Cryptopygus cisantarcticus, Friesea grisea, and Cryptopygus terranovus) are restricted to northern Victoria Land and three (Antarcticinella monoculata, Cryptopygus nivicolus, and Gomphiocephalus hodgsoni) are found only in southern Victoria Land, the two biogeographic zones which are separated by the vicinity of the Drygalski Ice Tongue. We found highly divergent lineages within all seven species (range 1.7–14.7%) corresponding to different geographic locations. Levels of genetic divergence for the southern Victoria Land species G. hodgsoni, the most widespread species (~27,000 km2), ranged from 5.9 to 7.3% divergence at sites located within 30 km, but separated by glaciers. We also found that the spatial patterns of genetic divergence differed between species. For example, levels of divergence were much higher for C. terranovus (>10%) than for F. grisea (<0.2%) that had been collected from the same sites in northern Victoria Land. Glaciers have been suggested to be major barriers to dispersal and two species (C. cisantarcticus and F. grisea) showed highly divergent (>5%) populations and over 87% of the total genetic variation (based on AMOVA) on either side of a single, 16 km width glacier. Collectively, these data provide evidence for limited dispersal opportunities among populations of springtails due to geological and glaciological barriers (e.g., glaciers and ice tongues). Some locations harbored highly genetically divergent populations and these areas are highlighted from a conservation perspective such as avoidance of human-mediated transport between sites. We conclude that species-specific spatial and temporal scales need to be considered when addressing ecological and physiological questions as well as conservation strategies for Antarctic Collembola.
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Construction and operation of research stations present the most pronounced human impacts on the Antarctic continent across a wide range of environmental values. Despite Antarctic Treaty Parties committing themselves to comprehensive protection of the environment, data on the spatial extent of impacts from their activities have been limited. To quantify this, we examined the area of building and ground disturbance across the entire continent using geographic information system mapping of satellite imagery. Here, we report the footprint of all buildings to be >390,000 m ² , with an additional disturbance footprint of >5,200,000 m ² just on ice-free land. These create a visual footprint similar in size to the total ice-free area of Antarctica, and impact over half of all large coastal ice-free areas. Our data demonstrate that human impacts are disproportionately concentrated in some of the most sensitive environments, with consequential implications for conservation management. This high-resolution measurement of the extent of infrastructure across the continent can be used to inform management decisions to balance sustainable scientific use and environmental protection of the Antarctic environment. © 2019, The Author(s), under exclusive licence to Springer Nature Limited.
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Snow algae are found in snowfields across cold regions of the planet, forming highly visible red and green patches below and on the snow surface. In Antarctica, they contribute significantly to terrestrial net primary productivity due to the paucity of land plants, but our knowledge of these communities is limited. Here we provide the first description of the metabolic and species diversity of green and red snow algae communities from four locations in Ryder Bay (Adelaide Island, 68°S), Antarctic Peninsula. During the 2015 austral summer season, we collected samples to measure the metabolic composition of snow algae communities and determined the species composition of these communities using metabarcoding. Green communities were protein‐rich, had a high chlorophyll content and contained many metabolites associated with nitrogen and amino acid metabolism. Red communities had a higher carotenoid content and contained more metabolites associated with carbohydrate and fatty acid metabolism. Chloromonas, Chlamydomonas and Chlorella were found in green blooms but only Chloromonas was detected in red blooms. Both communities also contained bacteria, protists and fungi. These data show the complexity and variation within snow algae communities in Antarctica and provide initial insights into the contribution they make to ecosystem functioning.
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About 400 subglacial lakes are known from Antarctica. The question of whether life unique of subglacial lakes exists has been paramount since their discovery. Despite frequent evidence of microbial life mostly from accretion ice, subglacial lakes are characterized by physiologically hostile conditions to metazoan life, as we know it. Pure water (salinity ≤0.4–1.2%), extreme cold (−3°C), high hydrostatic pressure, areas of limited or no oxygen availability and permanent darkness altogether require physiological adaptations to these harsh conditions. The record of gene sequences including some associated with hydrothermal vents does foster the idea of metazoan life in Lake Vostok. Here, we synthesize the physico-chemical environment surrounding sub-glacial lakes and potential sites of hydrothermal activity and advocate that the physico-chemical stability found at these sites may be the most likely sites for metazoan life to exist. The unique conditions presented by Lake Vostok may also offer an outlook on life to be expected in extra-terrestrial subglacial environments, such as on Jupiter's moon Europa or Saturn's moon Enceladus.
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We have obtained the first data demonstrating the capability of multicellular organisms for longterm cryobiosis in permafrost deposits of the Arctic. The viable soil nematodes Panagrolaimus aff. detritophagus (Rhabditida) and Plectus aff. parvus (Plectida) were isolated from the samples of Pleistocene permafrost deposits of the Kolyma River Lowland. The duration of natural cryopreservation of the nematodes corresponds to the age of the deposits, 30 000-40 000 years.
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From glacial reconstructions it is clear that Antarctic terrestrial life must have been extremely limited throughout Quaternary glacial periods. In contrast, recent biological studies provide clear evidence for long-term in situ persistence throughout glacial times within most extant Antarctic faunal and several microbial groups. However, even now, the evolutionary history of the Antarctic flora—despite playing major role in Antarctic ecosystems—remains poorly studied. We assessed the diversity, richness and relative age divergences within Schistidium (Grimmiaceae, Bryophyta), the most species-rich plant genus in the Antarctic, as well as the plant genus containing most Antarctic endemic species. We applied phylogenetic and molecular dating methods based on nuclear ribosomal Internal Transcribed Spacer sequences, including all known Antarctic Schistidium species with available sample material. We additionally investigated the continent-wide genetic diversity within the most common Antarctic representative of the genus—the endemic species Schistidium antarctici—and performed preliminary phylogeographic analyses of the bipolar species Schistidium rivulare. Most previously described Antarctic Schistidium species were genetically distinct, confirming their specific status. Interspecific divergences of all species took place at least ~1 Mya, suggesting a likely in situ persistence in Antarctica for (at least) all endemic Schistidium species. The widespread endemic species, Schistidium antarctici, diverged from other Antarctic congeners in the late Miocene, thereby revealing the oldest extant plant species currently known in Antarctica, and providing increasing support for the hypothesis of vegetation survival through multiple glacial periods. Within S. antarctici we identified several distinct clades dividing the eastern Antarctic Peninsula and Scotia Arc islands from the western Antarctic Peninsula and all continental locations. This suggests that the mountainous spine on the Antarctic Peninsula forms a strong barrier to gene flow in this species, while increased genetic diversity in the northern Maritime Antarctic indicates likely glacial refugia in this area. This study provides an important first step toward assessing the diversity and evolutionary history of the most speciose moss genus in the Antarctic. The multi-million year presence of several endemic species contributes to studies on their adaptive potential to survive climate change over both historical and contemporary timescales.
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Deschampsia antarctica is the only grass naturally occurring in Antarctica, and it is also indigenous to southern South America. We aimed to evaluate patterns of within-population genetic diversity and between the focal areas Patagonia and Antarctica by using 144 sequences of nuclear internal transcribed spacer and non-coding plastid regions. We analysed phylogenetic relationships between these two main areas and performed demographic and landscape analysis. To test the divergence time between Antarctic and Patagonian populations we used approximate Bayesian computation. We found 17 nuclear and eight plastid haplotypes. For both molecular markers, Patagonia was the most genetically variable area in the range of D. antarctica. The divergence time between populations from Antarctica and Patagonia was dated to the mid to late Pleistocene. The large number of private haplotypes found in Patagonia and the great genetic variability support the hypothesis of a South American origin of the Antarctic populations of D. antarctica. Finally, we suggest that D. antarctica probably survived the Last Glacial Maximum and possibly earlier glaciations in ice-free refugia in Patagonia and Antarctica. Dispersal to Antarctica possibly occurred in the mid to late Pleistocene through bird-aided long-distance transport from South America.
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In the present paper, we compare how the kelp gull, Larus dominicanus, utilizes various nest building materials, particularly vascular plants, bryophytes, lichens and other components, in the Fildes Peninsula area (King George Island) and on the Argentine Islands area. In both areas, nest material primarily consisted of the Antarctic hairgrass (Deschampsia antarctica), bryophytes, lichens, feathers, limpets, and algae. Our study reveals area-specific differences in the utilization of plants for nest building related to local conditions during the nesting season. In the Fildes area, vegetation emerges from under the winter snow cover earlier in the spring, giving the gulls greater choice locally, meaning that the gulls need not resort to long distance material transfer. Here, mosses and lichens dominate in the nest material, likely collected from the nearby vegetation formations. The Antarctic hairgrass in these conditions is mostly found in nests located directly within hairgrass formations. However, on the more southern Argentine Islands, kelp gulls routinely use D. antarctica and some mosses, transferring them from coastal hill tops where snow generally disappears earlier. Here, the gulls appear to be selective still, as they rarely use some mosses, such as Polytrichum strictum, that are abundant near the nesting locations. In the Argentine Islands area, we documented long-range transfer of the Antarctic hairgrass and some other vegetation materials from places of abundance to bare rocks of low islands lacking developed vegetation. This demonstrates the potential of the gulls to serve as dispersal and gene pool exchange agents for the local terrestrial biota in the maritime Antarctic, especially between highly isolated populations from small islands and ice-free areas.
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The Antarctic–South American bank-forming moss Chorisodontium aciphyllum is known for having the oldest sub-fossils of any extant plant in Antarctica as well as extreme survival abilities, making it a candidate species for possible long-term survival in Antarctica. Applying phylogeographic and population genetic methods using the plastid markers trnL-F and rps4 and the nuclear internal transcribed spacer, we investigated the genetic diversity within C. aciphyllum throughout its range. Low genetic variation was found in all loci, both between and within Antarctic and southern South American populations, suggesting a relatively recent (likely within the last million years) colonization of this moss to the Antarctic, as well as a likely severe bottleneck during Pleistocene glaciations in southern South America. We also performed a simple atmospheric transfer modeling approach to study potential colonization rates of small (microscopic/microbial) or spore-dispersed organisms (such as many mosses and lichens). These suggested that the northern Antarctic Peninsula shows potentially regular connectivity from southern South America, with air masses transferring, particularly southbound, between the two regions. We found elevated genetic variation of C. aciphyllum in Elephant Island, also the location of the oldest known moss banks (> 5500 years), suggesting this location to be a genetic hotspot for this species in the Antarctic.
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A bipolar disjunction is an extreme, yet common, biogeographic pattern in non-vascular plants, yet its underlyingmechanisms (vicariance or long-distance dispersal), origin and timing remain poorly understood. Here, combining a large-scale population dataset and multiple dating analyses, we examine the biogeography of four bipolar Polytrichales mosses, common to the Holarctic (temperate and polar Northern Hemisphere regions) and the Antarctic region (Antarctic, sub-Antarctic, southern South America) and other Southern Hemisphere (SH) regions. Our data reveal contrasting patterns, for three species were of Holarctic origin, with subsequent dispersal to the SH, while one, currently a particularly common species in the Holarctic (Polytrichum juniperinum), diversified in the Antarctic region and from here colonized both the Holarctic and other SH regions. Our findings suggest long-distance dispersal as the driver of bipolar disjunctions. We find such inter-hemispheric dispersals are rare, occurring on multi-million-year timescales. High-altitude tropical populations did not act as trans-equatorial ‘steppingstones’, but rather were derived from later dispersal events. All arrivals to the Antarctic region occurred well before the Last Glacial Maximum and previous glaciations, suggesting that, despite the harsh climate during these past glacial maxima, plants have had a much longer presence in this southern region than previously thought.
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Cryptobiosis is a reversible ametabolic state of life characterized by the ceasing of all metabolic processes, allowing survival of periods of intense adverse conditions. Here we show that 1) entire moss individuals, dated by ¹⁴C, survived through cryptobiosis during six centuries of cold-based glacier burial in Antarctica, 2) after re-exposure due to glacier retreat, instead of dying (due to high rates of respiration supporting repair processes), at least some of these mosses were able to return to a metabolically active state and remain alive. Moss survival was assessed through growth experiments and, for the first time, through vitality measurements. Future investigations on the genetic pathways involved in cryptobiosis and the subsequent recovery mechanisms will provide key information on their applicability to other systematic groups, with implications for fields as divergent as medicine, biodiversity conservation, agriculture and space exploration.
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Subglacial environments are known to harbour metabolically diverse microbial communities. These microbial communities drive chemical weathering of underlying bedrock and influence the geochemistry of glacial meltwater. Despite its importance in weathering reactions, the microbial cycling of iron in subglacial environments, in particular the role of microbial iron reduction, is poorly understood. In this study we address the prevalence of viable iron-reducing microorganisms in subglacial sediments from five geographically isolated glaciers. Iron-reducing enrichment cultures were established with sediment from beneath Engabreen (Norway), Finsterwalderbreen (Svalbard), Leverett and Russell glaciers (Greenland), and Lower Wright Glacier (Antarctica). Rates of iron reduction were higher at 4 °C compared with 15 °C in all but one duplicated second-generation enrichment culture, indicative of cold-tolerant and perhaps cold-adapted iron reducers. Analysis of bacterial 16S rRNA genes indicates Desulfosporosinus were the dominant iron-reducing microorganisms in low-temperature Engabreen, Finsterwalderbreen and Lower Wright Glacier enrichments, and Geobacter dominated in Russell and Leverett enrichments. Results from this study suggest microbial iron reduction is widespread in subglacial environments and may have important implications for global biogeochemical iron cycling and export to marine ecosystems.
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Historical views have characterized Antarctica as a frozen desert with low diversity, although recent studies suggest that this may not be true for microscopic organisms. For microbes, assessing endemism in the Antarctic region has been particularly important, especially against a backdrop of debate regarding their presumed cosmopolitan nature. To contribute to this conversation, we highlight the observed endemism of the freshwater diatom genus Luticola in Antarctica by synthesizing the results of a modern high-resolution taxonomy from the Continental, Maritime, and sub-Antarctic regions. We report that Luticola has one of the highest endemic rates of any diatom genus in Antarctica, in terms of total number of species (taxon endemism) and percentage of the entire genus (phylogenetic endemism). Of the over 200 species of Luticola globally, nearly 20% (43) occur in the Antarctic, with 42 of these being endemic. Within regions, Maritime Antarctica has the largest number of Luticola species and endemics (28 and 23, respectively), followed by Continental Antarctica (14, 9) and sub-Antarctic islands (8, 6). Thus, 38 of the 42 endemics are found in a single region only. While the timing of Luticola diversification has not been established, fossil evidence suggests recent invasions and/or diversification over a relatively short geologic timescale. Understanding the origin and evolution of endemic diatom species in Antarctica will help us better understand microbial biogeography, as well as assess and interpret impacts of large-scale environmental change taking place at southern latitudes.
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Previous work focused on allozymes and mitochondrial haplotypes has detected high levels of genetic variability between Cryptopygus terranovus populations, a springtail species endemic to Antarctica, until recently named Gressittacantha terranova . This study expands these biogeographical surveys using additional analytical techniques, providing a denser haplotype dataset and a wider sampling of localities. Specimens were collected from 11 sites across Victoria Land and sequenced for the cytochrome c oxidase subunit I mitochondrial gene ( cox1 ). Haplotypes were used for population genetics, demographic, molecular clock and Bayesian phylogenetic analyses. Landscape distribution and clustering of haplotypes were also examined for the first time in this species. Only three (out of 67) haplotypes are shared among populations, suggesting high genetic structure and limited gene flow between sites. As in previous studies, the population of Apostrophe Island has a closer genetic similarity with those of the central sites, rather than with its neighbours. Molecular clock estimates point to early differentiation of haplotypes in the late/mid-Miocene, also supporting the view that C. terranovus is a relict species that survived on the Antarctic continent during the Last Glacial Maximum. The present genetic composition of populations represents a mixture of ancient and more recent haplotypes, sometimes occurring in the same localities.
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We present Bedmap2, a new suite of gridded products describing surface elevation, ice-thickness and the seafloor and subglacial bed elevation of the Antarctic south of 60° S. We derived these products using data from a variety of sources, including many substantial surveys completed since the original Bedmap compilation (Bedmap1) in 2001. In particular, the Bedmap2 ice thickness grid is made from 25 million measurements, over two orders of magnitude more than were used in Bedmap1. In most parts of Antarctica the subglacial landscape is visible in much greater detail than was previously available and the improved data-coverage has in many areas revealed the full scale of mountain ranges, valleys, basins and troughs, only fragments of which were previously indicated in local surveys. The derived statistics for Bedmap2 show that the volume of ice contained in the Antarctic ice sheet (27 million km<sup>3</sup>) and its potential contribution to sea-level rise (58 m) are similar to those of Bedmap1, but the mean thickness of the ice sheet is 4.6% greater, the mean depth of the bed beneath the grounded ice sheet is 72 m lower and the area of ice sheet grounded on bed below sea level is increased by 10%. The Bedmap2 compilation highlights several areas beneath the ice sheet where the bed elevation is substantially lower than the deepest bed indicated by Bedmap1. These products, along with grids of data coverage and uncertainty, provide new opportunities for detailed modelling of the past and future evolution of the Antarctic ice sheets.
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As the accuracy and sensitivity of remote-sensing satellites improve, there is an increasing demand for more accurate and updated base datasets for surveying and monitoring. However, differentiating rock outcrop from snow and ice is a particular problem in Antarctica, where extensive cloud cover and widespread shaded regions lead to classification errors. The existing rock outcrop dataset has significant georeferencing issues as well as overestimation and generalisation of rock exposure areas. The most commonly used method for automated rock and snow differentiation, the normalised difference snow index (NDSI), has difficulty differentiating rock and snow in Antarctica due to misclassification of shaded pixels and is not able to differentiate illuminated rock from clouds. This study presents a new method for identifying rock exposures using Landsat 8 data. This is the first automated methodology for snow and rock differentiation that excludes areas of snow (both illuminated and shaded), clouds and liquid water whilst identifying both sunlit and shaded rock, achieving higher and more consistent accuracies than alternative data and methods such as the NDSI. The new methodology has been applied to the whole Antarctic continent (north of 82°40′ S) using Landsat 8 data to produce a new rock outcrop dataset for Antarctica. The new data (merged with existing data where Landsat 8 tiles are unavailable; most extensively south of 82°40′ S) reveal that exposed rock forms 0.18 % (21 745 km2) of the total land area of Antarctica: half of previous estimates.
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Although most models suggest continental Antarctica was covered by ice during the Last Glacial Maximum (LGM) it has been speculated that endemic species of soil invertebrates could have survived the Pleistocene at high elevation habitats protruding above the ice sheets. We analyzed a series of soil samples from different elevations at three locations along the Beardmore Glacier in the Transantarctic Mountains (in order of increasing elevation): Ebony Ridge (ER), Cloudmaker (CM), and Meyer Desert (MD). Geochemical analyses show the MD soils, which were exposed during the LGM, were the least weathered compared to lower elevations, and also had the highest total dissolved solids (TDS). MD soils are dominated by nitrate salts (NO3/Cl ratios >10) that can be observed in SEM images. High δ17O and δ18O values of the nitrate indicate that its source is solely of atmospheric origin. It is suggested that nitrate concentrations in the soil may be utilized to determine a relative “wetting age” to better assess invertebrate habitat suitability. The highest elevation sites at MD have been exposed and accumulating salts for the longest times, and because of the salt accumulations, they were not suitable as invertebrate refugia during the LGM.
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Climate changes are likely to have major influences on the distribution and abundance of Antarctic terrestrial biota. To assess arthropod distribution and diversity within the Ross Sea region, we examined mitochondrial DNA (COI) sequences for three currently recognized species of springtail (Collembola) collected from sites in the vicinity, and to the north of, the Mackay Glacier (77°S). This area acts as a transition between two biogeographic regions (northern and southern Victoria Land). We found populations of highly divergent individuals (5%-11.3% intraspecific sequence divergence) for each of the three putative springtail species, suggesting the possibility of cryptic diversity. Based on molecular clock estimates, these divergent lineages are likely to have been isolated for 3-5 million years. It was during this time that the Western Antarctic Ice Sheet (WAIS) was likely to have completely collapsed, potentially facilitating springtail dispersal via rafting on running waters and open seaways. The reformation of the WAIS would have isolated newly established populations, with subsequent dispersal restricted by glaciers and ice-covered areas. Given the currently limited distributions for these genetically divergent populations, any future changes in species' distributions can be easily tracked through the DNA barcoding of springtails from within the Mackay Glacier ecotone.
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The Antarctic Conservation Biogeographic Regions (ACBRs), originally proposed in 2012, are now established as an important tool in Antarctic science, conservation, management and policy. Here, we provide a revised version of the ACBRs, reflecting updates in underlying spatial layers, together with the results of new analyses justifying the inclusion of a 16th bioregion. This updated version now covers all ice-free areas of Antarctica and is publicly available through the Australian Antarctic Data Centre. In light of the interest in the ACBRs across a variety of research fields, we also provide a new set of summary statistics for the updated spatial layer, including landscape metrics, climate data, protected area coverage and an overview of human activity. The updated ACBRs represent a contemporary, practical and evidence-based foundation for understanding, conserving and managing Antarctic biodiversity at a continental scale.
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We review the geomorphological, sedimentological and chronological evidence for palaeo-ice streams on the continental shelf of Antarctica and use this information to investigate basal conditions and processes, and to identify factors controlling grounding-line retreat. A comprehensive circum-Antarctic inventory of known palaeo-ice streams, their basal characteristics and minimum ages for their retreat following the Last Glacial Maximum (LGM) is also provided. Antarctic palaeo-ice streams are identified by a set of diagnostic landforms that, nonetheless, display considerable spatial variability due to the influence of substrate, flow velocity and subglacial processes. During the LGM, palaeo-ice streams extended, via bathymetric troughs, to the shelf edge of the Antarctic Peninsula and West Antarctica, and typically, to the mid-outer shelf of East Antarctica. The retreat history of the Antarctic Ice Sheet since the LGM is characterised by considerable asynchroneity, with individual ice streams exhibiting different retreat histories. This variability allows Antarctic palaeo-ice streams to be classified into discrete retreat styles and the controls on grounding-line retreat to be investigated. Such analysis highlights the important impact of internal factors on ice stream dynamics, such as bed characteristics and slope, and drainage basin size. Whilst grounding-line retreat may be triggered, and to some extent paced, by external (atmospheric and oceanic) forcing, the individual characteristics of each ice stream will modulate the precise timing and rate of retreat through time.
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Polar temperatures over the last several million years have, at times, been slightly warmer than today, yet global mean sea level has been 6-9 metres higher as recently as the Last Interglacial (130,000 to 115,000 years ago) and possibly higher during the Pliocene epoch (about three million years ago). In both cases the Antarctic ice sheet has been implicated as the primary contributor, hinting at its future vulnerability. Here we use a model coupling ice sheet and climate dynamics - including previously underappreciated processes linking atmospheric warming with hydrofracturing of buttressing ice shelves and structural collapse of marine-terminating ice cliffs - that is calibrated against Pliocene and Last Interglacial sea-level estimates and applied to future greenhouse gas emission scenarios. Antarctica has the potential to contribute more than a metre of sea-level rise by 2100 and more than 15 metres by 2500, if emissions continue unabated. In this case atmospheric warming will soon become the dominant driver of ice loss, but prolonged ocean warming will delay its recovery for thousands of years.
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The Antarctic region comprises the continent, the Maritime Antarctic, the sub-Antarctic islands, and the southern cold temperate islands. Continental Antarctica is devoid of insects, but elsewhere diversity varies from 2 to more than 200 species, of which flies and beetles constitute the majority. Much is known about the drivers of this diversity at local and regional scales; current climate and glacial history play important roles. Investigations of responses to low temperatures, dry conditions, and varying salinity have spanned the ecological to the genomic, revealing new insights into how insects respond to stressful conditions. Biological invasions are common across much of the region and are expected to increase as climates become warmer. The drivers of invasion are reasonably well understood, although less is known about the impacts of invasion. Antarctic entomology has advanced considerably over the past 50 years, but key areas, such as interspecific interactions, remain underexplored.
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Bryophytes were mentioned for the first time in the Svalhard literature in 1675. The early bryological history up to 1875 is thoroughly reviewed, and the important later papers are presented. All known papers and books with reports of bryophytes from Svalbard have been scanned for taxonomic and geographical information. In all, 563 bryophyte species (current names excluding synonyms) have been considered. Of these, 190 have been shown or critically considered to be erroneous, leaving a total of 373 accepted bryophytes (85 hepatics and 288 mosses in 137 genera). One hundred and fifty (26 hepatics and 124 mosses) are accepted from Bjørnøya, and five of these are not known from the rest of Svalbard. Short or sometimes more extensive comments are made on 315 accepted and all rejected species; the comments usually include all reported localities with literature references, as well as numerous unreported localities based on own herbarium material. Six species are reported from Svalbard for the first time, and Plagiothecium svalbardense is described as new. One hundred and one valid bryophyte names (29 species, 3 subspecies, 58 varieties and 11 forms) have been based on type material from Svalbard, and 18 of these are basionyms of accepted species (Appendix 1). Two exsiccates have appeared, viz. Musci Spetsbergenses Exsiccati (Berggren 1875, with 220 numbered specimens) and Bryophyta Svalbardensia exsiccata (Bednarek-Ochyra et al. 1987, with 80 numbers). Numerical and alphabetical lists, including some revisions, are given to both (Appendix 2). Many of the considered Svalbard papers describe bryophyte vegetation, and a survey of all communities whose names are based on or include bryophytes, are provided (Appendix 3).
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In contrast to northern polar areas, the dipteran fauna of Antarctica is depauperate, with only two naturally occurring species of Chironomidae. Surprisingly little is known of the biology of these species. One, Parochlus steinenii, reaches the southern limit of a distribution covering the high Andes. Tierra del Fuego, South Georgia and the South Shetland Islands in the maritime Antarctic. The other. Belgica antarctica, is endemic to the maritime Antarctic. What factors influence the distribution and past colonisation of these two species in the Antarctic? Distributional data and evidence from an accidental introduction of a third chironomid (Eretmoptera murphuyi) suggest that difficulty of colonisation is the major factor limiting the number of dipteran species present in the maritime Antarctic. Other sub-Antarctic species are likely to be preadapted to more rigorous conditions, should natural or man-induced clonisation opportunities occur. Evidence from physiological and ecological studies identify adaptations which allow these two species to survive in the harsh terrestrial environment of Antarctica. Life history characteristics of all three species include flexibility in development rates and size achieved, and the ability to continue activity at low positive temperatures. The distribution of the endemic B. antarctica is probably limited by the availability of suitable moist, vegetated, habitats and its brachyptery, rather than biological constraints directly, whereas the southern distribution of P. steinenii may be temperature-limited. Competition does not appear to be an important factor in the biology and ecology of either species, or in Antarctic terrestrial ecosystems generally.
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Discusses the ecology of the tall turf-forming mosses Chorisodontium aciphyllum and Polytrichum alpestre, and the development and dynamics of the deep moss banks formed solely by these species within the maritime Antarctic botanical zone where these banks are a unique feature of the vegetation. On Signy Island a radiocarbon date of 4801 yr BP, has been obtained for a basal peat sample taken from a bank. This suggests that this area of Signy Island has been ice-free and available for plant colonization and moss-bank development for at least 4500-5000 years. -from Authors
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A population of 172 moss balls of Grimmia longirostris Hook. was studied in the Páramo de Piedras Blancas (Venezuelan Andes) at 4380 m. Moss ball transport in the paramo is caused primarily by needle-ice activity, with wind being a secondary agent. Moss ball size and shape varied widely, but were closely correlated. Larger globoids were significantly less spheroidal than smaller ones. This occurs because heavier moss balls rest for increasingly long periods of time on one side, thus becoming flattened. Several moss forms are described, including a type (pedestal mosses) attached to pebbles. Globular mosses lacked pebbles at their core, which was formed by silty soil and organic matter. Moss balls had a high content of organic matter (19%) and fine mineral grains (69.3%); the last is nearly four times higher than in site soils. This concentration of silt and clay results from moss balls trapping fine eolian dust and soil material as globoids roll over the ground. Due to their composition, globular mosses have a superior ability to store water compared to paramo soils. Larger moss balls had a greater water-holding capacity than smaller ones; they also lost water at considerably slower rates. This may be ecologically significant for moss ball growth and survival during dry periods.