Molecular data suggest long-term in Situ Antarctic persistence within Antarctica's most speciose plant genus, Schistidium

© 2018 Biersma, Jackson, Stech, Griffiths, Linse and Convey. 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.NERC STUDENTSHIP NE/K50094X/

Low genetic variation between South American and Antarctic populations of the bank-forming moss Chorisodontium aciphyllum (Dicranaceae)

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.This research was funded by a Natural Environment Research Council (NERC) Ph.D. studentship (ref. NE/K50094X/1) to E.M.B. and supported by NERC core funding to the BAS Biodiversity, Evolution and Adaptation Team

Global biogeographic patterns in bipolar moss species

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

Lichen response to ammonia deposition defines the footprint of a penguin rookery

Ammonia volatilized from penguin rookeries is a major nitrogen source in Antarctic coastal terrestrial ecosystems. However, the spatial extent of ammonia dispersion from rookeries and its impacts have not been quantified previously. We measured ammonia concentration in air and lichen ecophysiological response variables proximate to an Adèlie penguin rookery at Cape Hallett, northern Victoria Land. Ammonia emitted from the rookery was 15N-enriched (δ15N value +6.9) and concentrations in air ranged from 36–75 µg m−3 at the rookery centre to 0.05 µg m−3 at a distance of 15.3 km. δ15N values and rates of phosphomonoesterase (PME) activity in the lichens Usnea sphacelata and Umbilicaria decussata were strongly negatively related to distance from the rookery and PME activity was positively related to thallus N:P mass ratio. In contrast, the lichen Xanthomendoza borealis, which is largely restricted to within an area 0.5 km from the rookery perimeter, had high N, P and 15N concentrations but low PME activity suggesting that nutrient scavenging capacity is suppressed in highly eutrophicated sites. An ammonia dispersion model indicates that ammonia concentrations sufficient to significantly elevate PME activity and δ15N values (≥0.1 µg NH3 m−3) occurred over c. 40–300 km2 surrounding the rookery suggesting that penguin rookeries potentially can generate large spatial impact zones. In a general linear model NH3 concentration and lichen species identity were found to account for 72 % of variation in the putative proportion of lichen thallus N originating from penguin derived NH3. The results provide evidence of large scale impact of N transfer from a marine to an N-limited terrestrial ecosystem

The non-native chironomid Eretmoptera murphyi in Antarctica: erosion of the barriers to invasion

Antarctica is the continent least affected by invasive species, but climate change and increasing human activity are increasing this threat. Antarctic terrestrial ecosystems generally have low biodiversity with simple community structures and little competition for resources. Consequently, species with pre-adaptations or capabilities that allow them to tolerate polar conditions may have disproportionately large ecosystem impacts when introduced to Antarctica compared with other regions of the Earth. Here we investigate the invasion risk associated with the flightless chironomid midge, Eretmoptera murphyi, which was accidentally introduced from South Georgia (54°S) to Signy Island, South Orkney Islands (61°S), probably during plant transplantation experiments in the 1960s. Larval size class distribution analysis indicated that E. murphyi has a 2 year life cycle on Signy Island, supporting previous suggestions. Estimates of litter turnover show that recent large increases in E. murphyi population density and extent are likely to increase nutrient cycling rates on Signy Island substantially. Existing physiological adaptations may allow E. murphyi to colonise higher latitude locations. Growth rate and microhabitat climatic modelling show that temperature constraints on larval development on Anchorage Island (68°S) are theoretically similar to those on Signy Island even though it is ~750 km further south. Establishment of this non-native midge at climatically similar intervening locations along the western Antarctic Peninsula is therefore plausible. Currently, lack of effective natural dispersal mechanisms is probably limiting the spread of the midge. However, dispersal to other areas of the Antarctic Peninsula may occur via human-assisted transportation, highlighting the importance of appropriate biosecurity measures

Poa pratensis L., current status of the longest-established non-native vascular plant in the Antarctic

A single colony of the non-native grass Poa pratensis L., which was introduced inadvertently to Cierva Point, Antarctic Peninsula, during the 1954–1955 season, was still present during a survey in February 2012, making it the longest surviving non-native vascular plant colony known in Antarctica. Since 1991, the grass cover has roughly tripled in size, with an annual increase in area of approximately 0.016 m2, and an estimated maximum radial growth rate of 1.43 cm y−1. However, it remains restricted to the original site of introduction and its immediate surroundings (c. 1 m2). Annual flowering of the plants occurred during the 2010/2011 and 2011/2012 seasons; however, there has been no seed production and only incomplete development of the sexual structures. Current environmental conditions, including low temperatures, may inhibit sexual reproduction. Lack of effective vegetative dispersal may be influenced by the low level of human activity at the site, which limits opportunities for human-mediated dispersal. Although P. pratensis has existed at Cierva Point for almost 60 years, it has not yet become invasive. Scenarios for the potential future development of the species in Antarctica and the associated negative impacts upon the native vegetation from competition are discussed in the context of regional climate change. Finally, we describe the environmental risk presented by P. pratensis and argue that this non-native species should be eradicated as soon as possible in accordance with the Protocol on Environmental Protection to the Antarctic Treaty

Food choice of Antarctic soil arthropods clarified by stable isotope signatures

Antarctic soil ecosystems are amongst the most simplified on Earth and include only few soil arthropod species, generally believed to be opportunistic omnivorous feeders. Using stable isotopic analyses, we investigated the food choice of two common and widely distributed Antarctic soil arthropod species using natural abundances of 13C and 15N and an isotope labelling study. In the laboratory we fed the isotomid springtail Cryptopygus antarcticus six potential food sources (one algal species, two lichens and three mosses). Our results showed a clear preference for algae and lichens rather than mosses. These results were corroborated by field data comparing stable isotope signatures from the most dominant cryptogams and soil arthropods (C. antarcticus and the oribatid mite Alaskozetes antarcticus). Thus, for the first time in an Antarctic study, we present clear evidence that these soil arthropods show selectivity in their choice of food and have a preference for algae and lichens above mosses