Species identification and likely catch time period of whale bones from South Georgia

Skeletal remains of baleen whales killed during the onset of 20th century commercial whaling lie scattered across the shores and abandoned whaling stations of the subantarctic island of South Georgia. Here we report on genetic species identification of whale bones collected from South Georgia using standard historical DNA protocols. We amplified and sequenced short fragments of the mitochondrial DNA (mtDNA) control region from 281 available bone samples. Of these, 231 provided mtDNA sequences of sufficient quality and length (174–194 bp) for species identification: 158 bones were identified as humpback whale (Megaptera novaeangliae), 51 bones were identified as fin whale (Balaenoptera physalus), 18 bones were identified as blue whale (B. musculus), two bones were identified as sei whale (B. borealis), one bone was identified as a southern right whale (Eubalaena australis), and one bone was identified as a southern elephant seal (Mirounga leonina). The prominence of humpback, fin, and blue whale bones in the sample collection corresponds to the catch record of the early years of whaling on the island of South Georgia (pre‐1915), prior to the depletion of these populations.     

Summer-winter differences in copepod distribution around South Georgia

Zooplankton was sampled on a synoptic grid of stations centered on South Georgia during the austral summer of 1981/82 and winter 1983. Within the top 1000 m layer at oceanic stations, copepods averaged 48% of the total biomass in summer and winter, but outnumbered all other taxa combined by a factor of 10. In winter the mean zooplankton biomass within the top 1000 m was 68% of its summer level. Copepod biomass was 77% of its summer level. During both surveys, the large and abundant Calanoides acutus and Rhincalanus gigas dominated the copepod biomass and, with several other species, showed a marked downwards seasonal migration out of the top 250 m layer in winter. Antarctic epipelagic species predominated around the island during summer but tended to be replaced by sub-Antarctic and cosmopolitan species during the winter. Factors likely to influence our estimates of overall copepods abundance and changes in species composition include seasonality of reproduction, net mesh selection and differences in water mass distribution. The observed trends are attributed mainly to variation in the position of the Polar Front which lay north of the island during the summer survey yet lay across the survey area in winter. This resulted in a greater influence of sub-Antarctic water around South Georgia in winter and the displacement of Antarctic species.     

Multivariate statistical analysis of net diatom species distributions in the Southwestern Atlantic and Indian Ocean

Summary Vertical net haul diatom assemblages from near South Georgia, and from between Africa and Antarctica, were examined and compared. Variation among South Georgia stations was examined by principal component, cluster and canonical discriminant analyses. Diatom distributions provide evidence for at least two distinct water masses. The region north of the island is characterized by neritic, temperate diatoms and by an assemblage with low species diversity. The region south of the island is characterized by oceanic, antarctic species and relatively high species diversity. The regions are most distinct to the west of the island, intergrading east of the island. Within the north-south division, five station groupings were detected on the basis of distribution of dominant net diatoms. By comparing classical species ecological categorizations to results of principal component analysis, a neritic-oceanic factor was identified from net diatom distributions. This factor was common to both areas in spite of the fact that Biscoe and Agulhas collections were from different seasons.     

Microphytoplankton assemblages in the waters surrounding South Georgia, Antarctica during austral summer 1994

The composition and distribution of microphytoplankton assemblages in the vicinity of South Georgia were examined during a krill survey conducted during voyage 119 the RV Africana in austral summer (January/February) 1994. Microphytoplankton distribution was studied at 83 stations along a zig-zag transect in the waters surrounding South Georgia. Oceanographic and zooplankton data suggest that conditions were anomalous during the investigation, in that sub-Antarctic surface waters had probably been advected southwards and were present around the island. The two most widely distributed species were Corethron criophilum and Eucampia antarctica, which were recorded at all stations sampled. Also well-represented amongst the microphytoplankton assemblages were temperate, neritic species such as Chaetoceros atlanticus, Proboscia alata and Odontella weissflogii. This can probably be related to the intrusion of sub-Antarctic surface waters unusually far south. Cluster and ordination analysis identified three distinct groupings of stations in the waters surrounding the islands. These were found to the east, west and north of the islands. The diatom groupings identified during this investigation broadly correspond to three water masses previously described in the vicinity of South Georgia. Despite the anomalous hydrographical conditions that existed during the investigation, the differences in phytoplankton were negligible when compared to normal years. Small mesoscale features are, however, important in imposing biogeographic patterns on microphytoplankton. Received: 21 February 1996 / Accepted: 3 November 1996     

Distribution of six major copepod species around South Georgia during an austral winter

Summary A net sampling survey was conducted around the island of South Georgia during July/August 1983. This study compares the age structure and vertical and horizontal distributions of the dominant copepods Calanoides acutus, Calanus simillimus, C. propinquus, Rhincalanus gigas, Metridia lucens and M. gerlachei. The chief physical and biological factors affecting the distributions of these species are assessed and the results are compared with those from a similar survey around the island carried out in early summer (1981/1982). The survey grid lay within the Polar Front during the winter survey, and horizontal changes in copepod abundance corresponded well to the temperature gradient across the front. This pattern was interrupted by the South Georgia shelf where the seasonal migrants (Calanoides acutus, Rhincalanus gigas and Calanus simillimus) occurred in high abundance. The concentration of these migrants over the shelf relative to the oceanic surface layer was attributed to the shelf having prevented their seasonal migration. Within the oceanic area the copepods occupied differing depths, with Calanoides acutus and Metridia gerlachei living deeper than Calanus simillimus, C. propinquus and M. lucens. The populations also tended to live deeper in the warmer (NW) portion of the oceanic survey area. In contrast to the summer survey the age structure of each species showed little variation throughout the survey area. This was attributed mainly to the decreased rates of copepod growth and metabolism in winter.     

Structural changes to a mycothallus along a latitudinal transect through the maritime and sub-Antarctic

Previous studies have shown the leafy liverwort Cephaloziella varians to associate consistently with fungi, typically the ericoid mycorrhizal symbiont Rhizoscyphus ericae, across a wide latitudinal gradient in the maritime and sub-Antarctic. Hitherto, however, there are no quantitative data on the intensity of colonisation of C. varians by fungal structures in the natural environment and how colonisation might vary with changing environmental conditions. A study is hence reported showing that the frequency of colonisation by fungal structures of C. varians alters along a latitudinal transect from South Georgia (54° S, 38° W) to Moutonnée Valley on Alexander Island (71° S, 68° W). The percentage of stem length colonised by dark septate (DS) hyphae increased significantly along the transect, from 30% at South Georgia to 97% at Moutonnée Valley. In contrast, the percentage of stem length colonised by hyaline hyphae decreased significantly, from 85% at South Georgia to 13% at Moutonnée Valley, and that colonised by hyphal coils similarly decreased from 71% at the former location to 15% at the latter. The frequencies of DS hyphae were negatively associated with mean annual and seasonal air temperatures, whereas those of hyaline septate hyphae and hyphal coils were positively associated with air temperatures. Coils at northerly locations were more convoluted than those at southerly locations. The data indicate that hyphal coils, usually associated with nutrient exchange between partners in ericoid mycorrhizas, do form in C. varians tissues in the maritime and sub-Antarctic, but that the frequency of these structures diminishes in colder habitats.     

Respiratory metabolism of Hydromedion sparsutum and Perimylops antarcticus (Col., Perimylopidae) from South Georgia

Summary Metabolic rates were studied in adults and larvae of the two phytophagous beetles Hydromedion sparsutum and Perimylops antarcticus (Col., Perimylopidae) indigenous to the Sub-Antarctic island of South Georgia. From the regression lines for log₁₀ metabolic rate on temperature no difference in oxygen consumption was found between adults and larvae of Hydromedion or adults of Perimylops from lower sites near sea level and upper sites at about 200 m. Elevated rates of metabolism, however, was found in larvae of Perimylops from upper sites. The slopes of the regression lines of corresponding upper and lower sites adults or larvae were similar in all cases. The metabolic rates were of the same level in adults of both species and in the larvae, but generally higher in adults than in larvae. The activation energies, calculated from Arrhenius plots, varied from 6.7 to 13.6 kcal.mol^–1 in larvae and from 10.7 to 11.3 kcal.mol^–1 in adults. The low values may be interpreted as an expression of cold adapted metabolism. Compared with published data on phytophagous beetles, the metabolic rates of the two species from South Georgia are comparable to two alpine species, but lower than the rates of an Arctic species.     

Extinction and recolonization of maritime Antarctica in the limpet Nacella concinna (Strebel, 1908) during the last glacial cycle: toward a model of Quaternary biogeography in shallow Antarctic invertebrates

Quaternary glaciations in Antarctica drastically modified geographical ranges and population sizes of marine benthic invertebrates and thus affected the amount and distribution of intraspecific genetic variation. Here, we present new genetic information in the Antarctic limpet Nacella concinna, a dominant Antarctic benthic species along shallow ice‐free rocky ecosystems. We examined the patterns of genetic diversity and structure in this broadcast spawner along maritime Antarctica and from the peri‐Antarctic island of South Georgia. Genetic analyses showed that N. concinna represents a single panmictic unit in maritime Antarctic. Low levels of genetic diversity characterized this population; its median‐joining haplotype network revealed a typical star‐like topology with a short genealogy and a dominant haplotype broadly distributed. As previously reported with nuclear markers, we detected significant genetic differentiation between South Georgia Island and maritime Antarctica populations. Higher levels of genetic diversity, a more expanded genealogy and the presence of more private haplotypes support the hypothesis of glacial persistence in this peri‐Antarctic island. Bayesian Skyline plot and mismatch distribution analyses recognized an older demographic history in South Georgia. Approximate Bayesian computations did not support the persistence of N. concinna along maritime Antarctica during the last glacial period, but indicated the resilience of the species in peri‐Antarctic refugia (South Georgia Island). We proposed a model of Quaternary Biogeography for Antarctic marine benthic invertebrates with shallow and narrow bathymetric ranges including (i) extinction of maritime Antarctic populations during glacial periods; (ii) persistence of populations in peri‐Antarctic refugia; and (iii) recolonization of maritime Antarctica following the deglaciation process.     

Historical biogeography and post-glacial recolonization of South American temperate rain forest by the relictual marsupial Dromiciops gliroides

Aim Long‐term climatic variation has generated historical expansions and contractions of species ranges, with accompanying fragmentation and population bottlenecks, which are evidenced by spatial variation in genetic structure of populations. We examine here hypotheses concerning dispersal and vicariance in response to historical geoclimatic change and potential isolation produced by mountains and water barriers. Location The temperate rain forest of southern South America, which is distributed from coastal Chile, including the large continental island of Chiloé, across the Andes into Argentina. Methods We investigated our hypotheses in the phylogenetically and biogeographically relictual marsupial Dromiciops gliroides. We examined 56 specimens, which resulted from field samples and museum study skins from 21 localities. We evaluated the influence of two major barriers, the Andean cordillera and the waterway between the mainland and the large island of Chiloé, by performing Bayesian and maximum‐likelihood phylogenetic analyses on sequences of 877 base pairs of mitochondrial DNA. We further tested the contribution of the proposed geographical barriers using analysis of molecular variance (amova ). We also evaluated the responses of populations to historical north–south shifts of habitat associated with glacial history and sea‐level change. Results Our analyses revealed a phylogeny with three clades, two of which are widespread and contain nearly all the haplotypes: a northern clade (36–39° S) and a southern clade (40–43° S). These two clades contain forms from both sides of the Andes. Within the southern clade, island and mainland forms were not significantly differentiated. Tests of recent demographic change revealed that southern populations have experienced recent expansion, whereas northern populations exhibit long‐term stability. The direction of recent gene flow and range expansion is predominantly from Chile to Argentina, with a modest reciprocal exchange across the Andes. Recent gene flow from the island of Chiloé to the mainland is also supported. Main conclusions The genetic structure of contemporary D. gliroides populations suggests recent gene flow across the Andes and between the mainland and the island of Chiloé. Differences in demographic history that we detected between northern and southern populations have resulted from historical southward shifts of habitat associated with glacial recession in South America. Our results add to a growing literature that demonstrates the value of genetic data to illuminate how environmental history shapes species range and population structure.     

The white-chinned petrel (<Emphasis Type="Italic">Procellaria aequinoctialis</Emphasis>) on South Georgia: population size,distribution and global significance

More white-chinned petrels (Procellaria aequinoctialis) are accidentally killed in fisheries than probably any other seabird in the world, but the population impact of this mortality is poorly understood, partly because there have been no recent estimates of the species’ abundance. The breeding aggregation on the sub-Antarctic island of South Georgia is believed to be larger than all others combined. We estimated the size of this population by calculating the area of suitable habitat and the density of occupied burrows within it. Some 670,000 occupied nests were estimated for the island at mid-incubation, representing 0.9 million pairs of breeding-age birds associated with South Georgia in the survey seasons (2005/06 and 06/07). This is 40–45% of the previous estimate, but still represents well over half of the global population. If the population is declining due to fishery bycatch, as is likely, the scale of annual mortality in this population alone is at least in the high tens of thousands, and plausibly hundreds of thousands.     

Collembola of sub-Antarctic South Georgia

Qualitative samples of Collembola were obtained from a range of substrates near Husvik, Stromness Bay, South Georgia, between January and March 1996. Collections made at Hope Point near Grytviken (Cumberland East Bay) in 1980/1982 and 1997 were also examined. Fifteen species of Collembola were recorded around Husvik; most were widely distributed. Two of these, Friesia sp. nov. and Cryptopygus badasa, represent additions to the previously recognised fauna of 17 species. A new record of an introduced species, Hypogastrura purpurescens, was identified in collections from Hope Point in 1980/1982, bringing the total South Georgian fauna to 20 species. A key to South Georgian Collembola is included. H. purpurescens and the congeneric Hypogastrura viatica, both cosmopolitan invasive species, have also been recognised on other sub-Antarctic islands and have displaced resident species from some habitats. Their presence on South Georgia, and the dominance of H. viatica in some habitats, highlight the importance of strict quarantine measures to avoid the introduction of further alien invertebrates. Accepted: 30 November 1998     

Low population density and biology of an island population of house mice <Emphasis Type="Italic">Mus musculus</Emphasis> on South Georgia

South Georgia has been the southernmost location where populations of invasive house mice Mus musculus are found. The distribution of mice was investigated at Cape Rosa and the Nuñez Peninsula, the two areas of South Georgia where the species is known to occur. Live-trapping and kill-trapping took place during March 2012, at the end of the austral summer, with traps in four grids (3.2–4.0 ha in area) and on 34 trap lines distributed across a range of habitats from the shoreline to 250 m above sea level. Mice were scarce, with just 68 captures in about 1750 trap nights. Mouse densities in tussock habitat were estimated at 2.1–2.8 mice/ha, with higher densities of 5.3–6.4 mice/ha along the coastline. Mice were found in all habitats apart from higher-altitude fellfield, but were relatively less abundant in tussock habitat with large numbers of seals. Mice were breeding at both sites with 58 % of mature females pregnant or lactating. Litter size (7.1 ± 2.3 embryos) and adult body mass (21.4 ± 4.6 g) were typical of most other island mice populations. Population densities of mice on South Georgia are two orders of magnitude lower than mouse densities measured at other sub-Antarctic islands. The very low population density and its restricted distribution, with most captures close to the shore, and the presence of large numbers of burrowing petrels and South Georgia pipits Anthus antarcticus at both sites, suggest that mice have a relatively limited impact on South Georgia’s vertebrate biodiversity.     

Recent lepidopteran records from sub-Antarctic South Georgia

No Lepidoptera (butterflies and moths) are known to be residents of South Georgia. This paper presents new records of three lepidopterans on the island. Two, Agrotis ipsilon (Noctuidae) and Plutella xylostella (Yponomeutidae), are well-known migrants. The third, Plodia interpunctella (Pyralidae), is closely associated with human habitation. In the context of regional trends of climate warming P. xylostella may already possess the ecophysiological capacity to permit establishment on South Georgia.     

Distribution and ecology of <Emphasis Type="Italic">Chaenocephalus aceratus</Emphasis> (Channichthyidae) around South Georgia and Shag Rocks (Southern Ocean)

Chaenocephalus aceratus (Family Channicthyidae) is one of the dominant species of demersal fish living on the South Georgia shelf where it is caught in low numbers as by-catch in the mackerel icefish and Antarctic krill commercial fisheries. Data collected during 14 demersal fish surveys, from 1986 to 2006, are analysed to investigate biomass, distribution, growth and diet. Biomass estimates from a swept area method ranged from 4,462 to 28,740 tonnes on the South Georgia and Shag Rock shelves although few fish were caught at Shag Rocks. Analysis of length frequency data indicated that growth was fast in the first five years with males and females attaining lengths at first spawning of 440 mm TL and 520 mm TL. The diet was comprised of fish and crustaceans, with an ontogenetic shift in diet from Euphausia superba and mysids to benthic fish and decapods observed to begin at 250 mm TL. In larger fish (>500 mm TL) the diet was dominated by fish. C. aceratus diet is sufficiently different from the other species of channichthyids around South Georgia to suggest that these species have undergone resource partitioning.     

Morphometric and genetic evidence for four species of gentoo penguin

Gentoo penguins (Pygoscelis papua) are found across the Southern Ocean with a circumpolar distribution and notable genetic and morphological variation across their geographic range. Whether this geographic variation represents species‐level diversity has yet to be investigated in an integrative taxonomic framework. Here, we show that four distinct populations of gentoo penguins (Iles Kerguelen, Falkland Islands, South Georgia, and South Shetlands/Western Antarctic Peninsula) are genetically and morphologically distinct from one another. We present here a revised taxonomic treatment including formal nomenclatural changes. We suggest the designation of four species of gentoo penguin: P. papua in the Falkland Islands, P. ellsworthi in the South Shetland Islands/Western Antarctic Peninsula, P. taeniata in Iles Kerguelen, and a new gentoo species P. poncetii, described herein, in South Georgia. These findings of cryptic diversity add to many other such findings across the avian tree of life in recent years. Our results further highlight the importance of reassessing species boundaries as methodological advances are made, particularly for taxa of conservation concern. We recommend reassessment by the IUCN of each species, particularly P. taeniata and P. poncetii, which both show evidence of decline.     

Current distribution and potential extent of the most invasive alien plant species on La Réunion (Indian Ocean,Mascarene islands)

Abstract La Réunion Island has the largest area of intact vegetation of the islands in the Mascarene archipelago. Biological invasions are the primary threat to biodiversity in the intact habitats of the island (those not already transformed by agriculture and urbanization). Our study aimed to identify areas to prioritize in managing invasive alien plants for biodiversity conservation. We used extensive surveys of 238 distinct untransformed areas on La Réunion to define the current distribution patterns of all invasive species. Using expert knowledge, we compiled maps of the current distribution of the 46 most widespread/important invasive plants at the habitat scale (identified according to vegetation structure). Data from 440 botanical relevés for the 20 most threatening invasive alien plant species across the island and climatic envelope models were used to derive climatic suitability surfaces; these were used to map potential distributions for these species. More than 10 species invade 16.7% of the remaining habitat. Five habitat types are invaded by 25 or more species, and eight have fewer than 10 invasive alien plant species. Cluster analysis based on presence/absence of species in the 18 habitat types produced eight groups of species that invade particular habitats. Potential distribution models show that some species have invaded large parts of their potential range (e.g. Fuchsia magellanica, Furcraea foetida, Hiptage benghalensis), whereas others have the potential to increase their range substantially (e.g. Clidemia hirta, Strobilanthes hamiltonianus, Ulex europaeus). Management implications are identified for both groups. Three broad groups of habitats were identified: (i) intact habitats with a low level of invasion (e.g. subalpine shrubland); (ii) moderately invaded habitats with varying levels of intactness (ranging from windward submountain rainforest to the Acacia heterophylla forest); and (iii) habitats with little remaining intact area and high levels of invasion (e.g. lowland rainforest). Different management interventions are appropriate for these three groups.     

Dietary composition of juvenile Dissostichus eleginoides (Pisces, Nototheniidae) around Shag Rocks and South Georgia, Antarctica

A diet analysis of the Patagonian toothfish Dissostichus eleginoides, trawled in the South Georgia Islands area in March–April 1996, was carried out by frequency of occurrence (F%) and coefficient Q (%) methods. The samples consisted chiefly of immature specimens, with predominant length ranges of 30–70 cm (TL). Fish was by far the main food on the shelves of Shag Rocks and South Georgia, accounting for about 70% of prey. Krill appeared as secondary food, although its importance was overestimated by the frequency of occurrence method. Cephalopods and mysids were infrequent in the stomachs, and only at Shag Rocks and South Georgia, respectively. Lepidonotothen kempi, Champsocephalus gunnari and Chaenocephalus aceratus constituted the main fish prey and their variability between Shag Rocks and South Georgia depended on their local abundance. The large proportion of fish exhibiting stomachs full or close to fullness (together 62%) suggests that feeding intensity of the species was high.     

Evidence for rapid faunal changes on islands in a man-made lake

Summary Few studies of island biogeography have been made on islands in which the time of insularization is precisely known. We tested the effects of island formation on ant species diversity in a man-made lake in South Africa, to determine whether island effects are detectable after only 16 years of insularization. The number of ant species observed at trap-line censuses on islands was significantly correlated with island size (r=0.608; P<0.05) and ant species diversity was generally low compared with similar mainland habitats. Mean species number for all islands, including landbridge islands, was 5.5±3.3 species, and on mainland sites was 7.9±2.85 species. Island effects were more marked on islands <20 ha, which had a mean of 3.3±2.5 species per island. Species number on islands was inversely related to densities of the aggressive Anoplolepis custodiens and A. steingroeveri. These two species were only patchily distributed on mainlands, but these ants were nearly ubiquitous on small islands. Several lines of evidence suggest that this single species domination may be responsible for island effects. Island sites also differed in the number of ant species in different trophic groupings, tending to have fewer granivorous species than the mainland sites, but species in other diet groups were similar in both island and mainland habitats. We conclude that there have been marked changes in the ant faunas on islands smaller than 20 ha apparently due to changes in abundance of the dominant ant species. However, the causes of these changes are unknown.     

Distribution of six major copepod species around South Georgia in early summer

Summary An intensive net sampling survey was conducted around the island of South Georgia during November/December 1981. The distribution and copepodite stage structure of the dominant copepods Calanoides acutus, Calanus simillimus, C. propinquus, Rhincalanus gigas, Metridia lucens and Metridia gerlachei were compared. The herbivorous species had completed their spring vertical migration and their summer generations were developing during the survey. At every station, Calanoides acutus was noticeably more advanced than Rhincalanus gigas in its reproductive cycle. The species were also more advanced in their development in the SE oceanic part of the survey area than in the NW. However, copepod development rates are rapid at this time of year, and this apparent regional difference may be due mainly to temporal variation during the four weeks of the survey. A truly regional variation in timing of reproduction was found when development was compared between shelf and oceanic waters. Spawning of Rhincalanus gigas and particularly of Calanoides acutus was later over the shelf. The age structure of the epi-mesopelagic metridinids also differed between shelf and oceanic waters, but no age differences were found for either Calanus simillimus or C. propinquus, both of which live and spawn higher in the water column than the other species. The presence of the South Georgia shelf also limited the overall abundance of the two deeper living metridinids. However, the length of time between spawning (and production of large numbers of early copepodids) and sampling had the largest influence on observed species abundance within the survey area.     

A summer-winter comparison of zooplankton in the oceanic area Around South Georgia

Summary Zooplankton was sampled with RMT (1+8) gear on a synoptic grid of stations centred on South Georgia during the austral summer (November/December 1981) and winter (July/August 1983). This initial paper compares zooplankton biomass, vertical distribution and species composition from RMT 1 catches in the oceanic portion of the grid (water depth greater than 2000 m) during the two surveys. In the winter survey, mean zooplankton biomass within the top 1000 m of the water column was 68% of its summer level. This drop was largely due to a decrease in abundance of krill (Euphausia superba), although biomass of copepods and remaining zooplankton also decreased. Copepods averaged 48% of total biomass in summer and winter, but outnumbered all other taxa put together by a factor of 10. Antarctic epipelagic species predominated around the island in the summer survey but tended to be replaced by sub-Antarctic or cosmopolitan species during the winter survey. The majority of zooplankton also showed a downwards seasonal migration out of the top 250 m layer in winter. However, several epipelagic species, including E. superba, did not migrate, and these tended to have the largest summer-winter differences in overall abundance. These trends were attributed to variation in the position of the Polar Front, which lay north of the island during the summer survey but lay across the survey area in winter, resulting in a greater influence of sub-Antarctic water and the displacement of Antarctic species.