Towards a model of postglacial biogeography in shallow marine species along the Patagonian Province: lessons from the limpet Nacella magellanica (Gmelin, 1791)

ABSTRACT: BACKGROUND: Patagonia extends for more than 84,000 km of irregular coasts is an area especially apt to evaluate how historic and contemporary processes influence the distribution and connectivity of shallow marine benthic organisms. The true limpet Nacella magellanica has a wide distribution in this province and represents a suitable model to infer the Quaternary glacial legacy on marine benthic organisms. This species inhabits ice-free rocky ecosystems, has a narrow bathymetric range and consequently should have been severely affected by recurrent glacial cycles during the Quaternary. We performed phylogeographic and demographic analyses of N. magellanica from 14 localities along its distribution in Pacific Patagonia, Atlantic Patagonia, and the Falkland/Malvinas Islands. RESULTS: Mitochondrial (COI) DNA analyses of 357 individuals of N. magellanica revealed an absence of genetic differentiation in the species with a single genetic unit along Pacific Patagonia. However, we detected significant genetic differences among three main groups named Pacific Patagonia, Atlantic Patagonia and Falkland/Malvinas Islands. Migration rate estimations indicated asymmetrical gene flow, primarily from Pacific Patagonia to Atlantic Patagonia (Nem=2.21) and the Falkland/Malvinas Islands (Nem=16.6). Demographic reconstruction in Pacific Patagonia suggests a recent recolonization process (< 10 ka) supported by neutrality tests, mismatch distribution and the median-joining haplotype genealogy. CONCLUSIONS: Absence of genetic structure, a single dominant haplotype, lack of correlation between geographic and genetic distance, high estimated migration rates and the signal of recent demographic growth represent a large body of evidence supporting the hypothesis of rapid postglacial expansion in this species in Pacific Patagonia. This expansion could have been sustained by larval dispersal following the main current system in this area. Lower levels of genetic diversity in inland sea areas suggest that fjords and channels represent the areas most recently colonized by the species. Hence recolonization seems to follow a west to east direction to areas that were progressively deglaciated. Significant genetic differences among Pacific, Atlantic and Falkland/Malvinas Islands populations may be also explained through disparities in their respective glaciological and geological histories. The Falkland/Malvinas Islands, more than representing a glacial refugium for the species, seems to constitute a sink area considering the strong asymmetric gene flow detected from Pacific to Atlantic sectors. These results suggest that historical and contemporary processes represent the main factors shaping the modern biogeography of most shallow marine benthic invertebrates inhabiting the Patagonian Province.     

Spiders from Beauchêne Island, Falkland Islands, South Atlantic

A collection of 55 spiders, which was made on Beauchêne Island, Falkland Islands archipelago, in December 1980, contains seven species. Three of these are known from the main body of the Falkland Islands, one is a new subspecies of a species previously known from Patagonia and Tierra del Fuego, and the remaining three are endemic to Beauchêne Island. New taxa described are Auximus fuegianus bransfieldi nssp. and Emmenomma beauchenicum n.sp.     

The enigma of guanacos in the Falkland Islands: the legacy of John Hamilton

Aim To address the biogeographical enigma of why guanacos (Lama guanicoe) are in the Falkland Islands we investigated the following questions: (1) What was the origin of the introduced guanacos? (2) What were the initial population sizes? (3) Why are they found only on one island? and (4) Who was John Hamilton and what role did he play? Location The Falkland Islands are located in the South Atlantic Ocean 600 km east of Patagonia at the southern end of South America. While dominated by East and West Falkland Islands, the archipelago is composed of some 750 islands. Sedge and Staats Islands, two small outlying islands of West Falkland, are the focus of this paper. Methods Historical information was collected from known relevant documents housed at the Falkland Islands Government Archives in Stanley, and personal interviews conducted with past and present residents of West Falklands. Research expeditions were made to Staats Island in 1999, 2002 and 2003 to assess the guanaco population size, distribution and social structure. Results Guanacos were unsuccessfully introduced in 1862 to East Falkland south of Mt Pleasant where Prince Alfred hunted them in 1871. John Hamilton, Scottish immigrant to the Falklands and Patagonia of southern Argentina and Chile, was the driving force in the introduction of guanacos from the region of Rio Gallegos, Argentina during the 1930s. The guanaco was one of several wildlife species he introduced, however, only the guanaco, Patagonia grey fox (Dusicyon griseus) and perhaps the sea otter (Lutra felina) survive. Hamilton's acting agent, Jimmy Miller, imported four shipments totalling 26 guanacos from 1934 to 1939. In 1934 the Falkland Government authorized Miller to introduce guanacos to Sedge Island, all 11 of which disappeared. Whether intentional or accidental, 15 guanacos were taken to Staats Island, an islet of 500 ha on the western edge of the archipelago. Historically, guanacos are unexpected on Staats Island because documentation authorizing their introduction is unknown. Guanaco numbers have fluctuated widely on Staats Island for 65 years primarily due to culling. In 1959 the population was dangerously close to extirpation, but today 400 thrive there. A severely reduced gene pool and genetic bottlenecking were suggested by recent field studies, revealing preliminary evidence of deleterious consequences of inbreeding. Main conclusions John Hamilton, spirited and visionary Scottish immigrant to the Falklands in the early 1880s, was responsible for the introduction of guanacos into the Falkland Islands. While there are some gaps in the historical events, the enigma of how and why guanacos were introduced to a single island in the South Atlantic Ocean is understood. Today, Staats Island, as a closed system, is a rare natural experiment in progress. It offers unique opportunities for addressing advanced questions in ungulate population, behavioural and genetic ecology. The population potentially also represents breeding stock for farming the guanaco's highly valuable wool on other islands. Thus, among his many efforts to practice land stewardship and promote economic diversity through the introduction of Patagonian wildlife, a remaining legacy of John Hamilton to the Falkland Islands is unmistakably the guanacos of Staats Island.     

Discrimination between farmed and free-living invasive salmonids in Chilean Patagonia using stable isotope analysis

In Chilean Patagonia relatively pristine aquatic environments are being modified by the introduction of exotic salmonids, initially through their deliberate release for sport fishing since the early twentieth century, and more recently via the accidental escape from fish farms. There is therefore a need to reliably distinguish between naturally reproducing and fugitive salmonids associated with the Chilean salmonid farming industry, the second largest in the world. We tested the ability of stable isotope analysis (SIA) and analysis of scale growth profiles to discriminate between farmed and free-living salmonids sampled around the Island of Chiloé. Juvenile Atlantic salmon (Salmo salar) and rainbow trout (Oncorhynchus mykiss) from aquaculture facilities were significantly more enriched in δ¹⁵N and lipid-corrected δ¹³C than river-caught individuals. Scale growth slopes during the first year in freshwater were significantly higher in farmed than in wild-caught rainbow trout, indicating faster somatic growth under hatchery conditions. Stable isotopes analysis classified 94% of juvenile Atlantic salmon and rainbow trout to their correct farm or free-living groups. Our results, therefore, can help to elucidate the origin and spread of exotic invasive salmonids in Chile, and address one of the biggest threats to native freshwater fishes in Patagonia and other temperate zones of the Southern Hemisphere.     

The galaxiid fishes of South America

Four galaxiid species in two genera are recognized and redescribed from the South American region, viz., Galaxias maculatus (Jenyns), G. platei Steindachner, G. globiceps Eigenmann, and Brachygalaxias bullocki (Regan). All of these species occur in Chile, but G. maculatus and G. platei occur also east of the Andean divide in Argentina, in Tierra del Fuego, and the Falkland Islands. The two species east of the Andes are regarded as having crossed the mountains from west to east. The galaxiids, together with Aplochiton (family Aplochitonidae) and Geotria (family Petromyzonidae), are regarded as southern circumpolar elements in the South American freshwater fish fauna, distinct from the group of species of Amazonian derivation (trichomycterids, Nematogenys, Diplomystes, Cheirodon , etc.) and distinct also from oceanic derivatives, either recent (atherinids) or ancient (serranids).     

Evaluating Potential Effects of Exotic Freshwater Fish From Incomplete Species Presence–absence Data

Many freshwater ecosystems and biotas around the world are threatened with extinction. Freshwater fishes, for example, are the most endangered vertebrates after amphibians. Exotic fish are widely recognized as a major disturbance agent for native fish. Evaluating the ecological effects of invaders presents many challenges and the problem is greatly augmented in parts of the world where the native fauna is poorly known and where exotic species are commonplace. We use the fish community of Patagonia, a small and distinct native biota dominated by exotic salmonids, as a case study to ask: what can we learn about the effects of exotic fish species from fragmentary or partial data and how do such data point the way to what needs to be learned? We review the available data and literature on the distribution and status of native and introduced fish. We compile a novel regional presence/absence species database, build fish distribution maps, describe distribution patterns of native and exotic species, and identify critical information voids. A comparative review of literature from Patagonia and Australasia, where a similar native and exotic fish fauna is found, helps us to identify research priorities and promising management strategies for the conservation of native fish fauna. We conclude that the main challenge for fish conservation in Patagonia is to identify management strategies that could preserve native species while maintaining the quality of salmonid fisheries.     

Physiological responses to temperature in Merizodus soledadinus (Col., Carabidae), a subpolar carabid beetle invading sub-Antarctic islands

Recent human activities and rising air temperature have increased the vulnerability of sub-Antarctic islands to alien species. At the Kerguelen Islands, the predaceous ground beetle Merizodus soledadinus is the only invasive insect originating from the southern cold temperate area (native from Patagonia, Tierra del Fuego and Falkland Islands). This austral origin raises the question of the limits of its physiological tolerance and capability to withstand (1) global change and (2) warmer climates of archipelagos where its accidental transportation could be facilitated from the Kerguelen Islands (namely Amsterdam and Saint Paul). Using gas chromatography/mass spectrometry metabolomics, we compared metabotypes of adults exposed to different temperatures (0, 4, 8, 12, 16, 20 °C). All individuals survived after 2 weeks regardless of the temperature they were exposed to. The physiological changes observed were consistent with increased metabolic rate at increased temperatures, without extreme metabotypes that are characteristic of acute stress. First cues of sublethal stress were observed after prolonged exposure to 20 °C, a warm regime unrealistic for such duration in sub-Antarctic Islands. Overall, M. soledadinus’ thermal tolerance exceeded temperatures currently experienced in nature, suggesting that climate warming may boost its invasion by eliciting its activity and broadening habitat suitability in both invaded and still pristine islands. This thermal tolerance may allow survival aboard ships and development in sub-Antarctic islands with conditions warmer than the Kerguelen Islands, such as Amsterdam and St Paul. Stringent biosecurity measures are thus needed to prevent transfer from Kerguelen to these islands. The native range of this predaceous beetle limited to the Falkland Islands and southernmost South America may be partly constrained by factors other than temperature, such as desiccation, predation or competition.     

Crying wolf, crying foul, or crying shame: alien salmonids and a biodiversity crisis in the southern cool-temperate galaxioid fishes?

The galaxioid fishes are the dominant, most speciose group of freshwater fishes (with >50 species) in the lands of the cool southern hemisphere, with representatives in western and eastern Australia, Tasmania, New Caledonia, Lord Howe Island, New Zealand, the Chatham, Auckland and Campbell Islands, Patagonian South America (Chile, Argentina), the Falkland Islands and South Africa. The group is most diverse in Australia and New Zealand. Lepidogalaxiidae is found only in Australia, Retropinnidae in Australia and New Zealand, and Galaxiidae across the entire range of the group. Many species are in serious conservation crisis for a diversity of reasons, including habitat deterioration and possibly fisheries exploitation, but there is enduring and pervasive information that shows that the group has been seriously impacted by the acclimatisation of salmonid fishes originating in the cool-temperate northern hemisphere, particularly brown and rainbow trout. With few exceptions, where these trout have been introduced there has been major decline in the galaxioids, especially Galaxiidae, as a result of a complexly interacting series of adverse impacts from these introduced fishes. In some places, centrarchids and cichlids may also have adverse impacts. In addition, there appear to have been adverse impacts from the translocation of galaxioids into communities where they do not naturally occur. In many instances it appears that displacement of the galaxioids has led to a situation where galaxioids and salmonids no longer co-occur, owing either to displacement or predation, leading to fish communities in which there is no explicit evidence for displacement. These effects are resulting in the galaxioid fishes being amongst the most seriously threatened fishes known.     

Increasing trend in the number of Southern Rockhopper Penguins (Eudyptes c. chrysocome) breeding at the Falkland Islands

The Falkland Islands currently supports one of the largest Southern Rockhopper Penguin (Eudyptes c. chrysocome) populations. Archipelago-wide censuses conducted in 2000 and 2005 revealed that the number of breeding pairs had declined by 30 % during this period. To establish whether the breeding population continued to decline, an archipelago-wide census was conducted in 2010. We report a conservative estimate of 319,163 ±SD 24,820 pairs breeding at the Falkland Islands in 2010. This represents a 51 % increase when compared with the number counted in 2005. A simple stochastic population model was developed to investigate the extent to which changes in demographic parameters between 2005 and 2010 could account for the increase in breeding pairs. The population model predicted a 38 % increase in the number of breeding pairs over a 5-year period (289,431 ±SD 24,615). The increase in the number of breeding pairs was therefore probably attributed to improved vital rates in the period between the 2005 and 2010 archipelago-wide censuses in combination with other factors such as a reduction in the proportion of adult birds that abstained from breeding. Based on the 2010 Falkland Islands estimate, the global population of the subspecies E. c. chrysocome is now closer to 870,000 breeding pairs of which the Falkland Islands accounts for approximately 36 %, the second largest proportion after Chile. We conclude that despite fluctuations, the number of Southern Rockhopper Penguins breeding at the Falkland Islands has increased over the last 15 years and suggest that the ‘Vulnerable’ conservation status of the species be re-assessed.     

Minimal evidence of interspecific hybridisation between the Yellow-billed Duck and introduced Mallard in central and northwestern South Africa

Hybridisation is the interbreeding of genetically distinct groups that can lead to introgression – an exchange of genetic material between species. Hybridisation is of conservation concern when an alien invasive species is involved, as it can lead to a loss of local genetic adaptations and genetic diversity. Hybridisation is a significant threat for many dabbling ducks where interbreeding with the closely related invasive Mallard Anas platyrhynchos is extremely common. Phenotypic evidence suggests that Mallard populations in South Africa hybridise with the indigenous Yellow-billed Duck Anas undulata. The aim of this study was to determine the incidence of hybridisation between Yellow-billed Ducks, occurring in central and northwestern South Africa, and introduced Mallards. Genetic variation between Mallards, Yellow-billed Ducks, and their inferred hybrids was assessed using mitochondrial and microsatellite DNA markers. All samples inferred to be hybrids based on the phenotype were found to have Yellowbilled Duck mitochondrial DNA and showed minimal evidence of admixture across the microsatellite markers. Thus, these results do not support the notion that hybridisation between Mallards and Yellow-billed Ducks is prevalent in central and northwestern South Africa. However, hybridisation could be occurring where Mallards are found in higher abundance, such as in the Western Cape Province. Therefore, continued monitoring of this potential hybridisation should be performed frequently and throughout South Africa.     

Extreme intraspecific mitochondrial DNA sequence divergence in Galaxias maculatus (Osteichthys: Galaxiidae), one of the world's most widespread freshwater fish

Biogeographic controversies surrounding the widespread freshwater fish, Galaxias maculatus, were addressed with DNA sequence data. Mitochondrial cytochrome b and 16S rRNA sequences were obtained from representatives of six populations of this species. Substantial levels of cytochrome b (maximum 14.6%) and 16S rRNA sequence divergence (maximum 6.0%) were detected between western Pacific (Tasmania-New Zealand) and South American (Chile-Falkland Islands) haplotypes. A considerable level of divergence was also detected between Tasmanian and New Zealand haplotypes (maximum 5.1%) and within and among Chilean and Falkland Island G. maculatus (maximum 3. 8%). The phylogenetic structure of haplotypes conflicts with the accepted pattern of continental fragmentation. Molecular clock calibrations suggest that haplotype divergences postdate the fragmentation of Gondwana. These findings point to marine dispersal rather than ancient vicariance as an explanation for the wide distribution. The phylogenetic structure of South American haplotypes was not consistent with their geographic distribution. We consider factors such as population divergence, population size, dispersal, secondary contact, and philopatry as potential causes of the high level of mtDNA nucleotide diversity in this species.     

Biology and host range testing of Triarthria setipennis and Ocytata pallipes (Diptera: Tachinidae) for the control of the European earwig (Forficula auricularia)

The European earwig (Forficula auricularia) is an invasive species in the Falkland Islands, causing considerable problems for local horticulture, as well threatening indigenous ecosystems. To assess the potential of a classical biological control introduction two parasitoid fly species, Triarthria setipennis and Ocytata pallipes (Diptera: Tachinidae), were collected from sites in southern and northern England and then tested for their suitability as earwig biological controls at Egham, UK. Both species had previously been introduced into North America for earwig control however little is known of their long-term efficacy and host specificity. Host range tests including both target and non-target species were done. As there are no native Dermaptera on the Falkland Islands, tests were restricted to the field cricket Gryllus assimilis and the Dubia cockroach Blaptica dubia, as representatives of insect orders phylogenetically closely related to earwigs. A second cricket species (Gryllus bimaculatus) was included in an egg-depositing experiment for O. pallipes. Both tachinid species successfully parasitised and emerged from earwigs under laboratory conditions but no signs of parasitisation and development were observed in either the cricket or cockroach.     

Trophic interactions among native and introduced fishes in a northern Patagonian oligotrophic lake

In summer the littoral zone of Lake Rivadavia contained almost all the juveniles of the native fish species present in the lake together with juveniles of the introduced salmonid Oncorhynchus mykiss . In particular, a high degree of foraging interaction was found among juveniles of the native species Aplochiton zebra and Percichthys trucha and juveniles O. mykiss. Differences in the juvenile assemblage composition, both in the presence and in the relative proportion of the species were associated with areas with different densities of vegetation. The adults of nearly all species were generalized benthic invertebrate predators. Only adults of A. zebra were absent in the littoral. Juvenile and adult O. mykiss were the only fish that preyed on planktonic organisms. Oncorhynchus mykiss was the only species that performed an ontogenetic diet shift between juveniles and adults, changing main prey categories from benthic to planktonic species.     

Species relative abundance and direction of introgression in oaks

Successful hybridisation and subsequent introgression lead to the transfer of genetic material across species boundaries. In this process, species relative abundance can play a significant role. If one species is less abundant than the other, its females will receive many heterospecific gametes, increasing mate-recognition errors and thus hybridisation rate. Moreover, first-generation hybrids will also more likely mate with the more abundant species, leading to asymmetric introgression. These predictions have important fundamental consequences, especially during biological invasions or when a rare species threatened by extinction is surrounded by individuals from a related species. However, experimental tests in nature of the importance of the relative abundance of each species on hybridisation dynamics remain scarce. We assess here the impact of species relative abundance on hybridisation dynamics among four species from the European white oak species complex. A total of 2107 oak trees were genotyped at 10 microsatellite markers and Bayesian clustering methods were used to identify reference trees of each species. We then used these reference trees to simulate purebred and hybrid genotypes to determine optimal threshold for genetic assignment. With this approach, we found widespread evidence of hybridisation between all studied oak species, with high occurrence of hybrids, varying from 11% to 31% according to stand and sampling strategies. This finding suggests that hybridisation is a common phenomenon that plays a significant role in evolution of this oak species complex. In addition, we demonstrate a strong impact of species abundance on both hybridisation rate and introgression directionality.     

SUMMER DISTRIBUTION OF PELAGIC BIRDS OFF THE COAST OF ARGENTINA

Eighteen species of sea–birds were observed from CSS 'Hudson' in the South Atlantic between the Rio de la Plata and Tierra del Fuego in late January and late February 1970. Major feeding areas were noted off Mar del Plata, the Valdes Peninsula and Tierra del Fuego, Argentina, the first two in rich modified or unmodified water of the subantarctic Falkland Current, associated with upwelling. The southernmost area had most birds, the northernmost fewest. A temperate–latitude assemblage including Yellow–nosed Albatross and Cory's Shearwater occurred off Mar del Plata in warmed water of subantarctic (Falkland Current) origin. South of 39°S these two species were replaced by Black–browed Albatross and Great Shearwater in cold offshore water of the Falkland Current. Off the Valdes Peninsula Black–browed Albatross, Great and Sooty Shearwaters, White–chinned Petrels and family groups of Magellanic Penguins occurred near the edge of highly–productive water of the Falkland Current. The largest feeding area, east of Tierra del Fuego, had Magellanic and Rockhopper Penguins, Great Shearwaters, King Cormorants, South American Terns and jaegers, but no oceanographic information is available. At 50°S prions and diving petrels appeared for the first time, marking a possible oceanographic discontinuity or proximity to colonies on the Falkland Islands. Several common species of the South Atlantic were not seen on the transect including Cape Pigeon, Southern Fulmar, Soft–plumaged Petrel, Schlegel's Petrel and Pediunker. Kelp Gull, South American Tern, Manx Shearwater and jaegers were rare or absent far from land. New southern limits for the western South Atlantic Ocean were established for Cory's and Manx Shearwaters, and the Great Shearwater was observed for the first time in Chilean waters.     

A New Species of Kinorhynchus (Homalorhagida, Pycnophyidae) from Australia with a Redescription and Range Extension of Other Kinorhyncha from the South Pacific

Kinorhynchus phyllotropis sp.n., from Sydney Harbour, is the first species of the phylum Kinorhyncha to be described from Australian coasts. It appears to be most closely related to K. anomalus (Lang, 1953) from the Chilean coast and possibly, but to a lesser extent, to K. spinosus (Lang, 1949) from the Falkland Islands in the South Atlantic. Kinorhynchus anomalus is rede-scribed and, along with the new species, is compared with the South Atlantic species, the two species known from the Northeast Pacific and the remaining four members of this genus from the North Atlantic, Mediterranean Sea and Black Sea. Cateria styx Gerlach, 1956, is reported from beaches north of Valparaiso, Chile, the first report of this mesopsarnmic cryptorhagid kinorhynch from the Pacific Ocean.     

Multiple losses of flight and recent speciation in steamer ducks

Steamer ducks (Tachyeres) comprise four species, three of which are flightless. The flightless species are believed to have diverged from a flying common ancestor during the Late Pleistocene; however, their taxonomy remains contentious. Of particular interest is the previously unstudied population of flying steamer ducks in the Falkland Islands. We present the first genetic data from this insular population, and illustrate that the flying and flightless steamer ducks on the Falkland Islands are genetically indistinguishable, in contrast to their traditional classification as separate species. The three species that reside in continental South America form a genetically distinct lineage from the Falkland Island ducks. The Falkland steamer ducks diverged from their continental relatives 2.2-0.6 million years ago, coincident with a probable land bridge connecting the Falkland Islands to the mainland. The three continental species share a common ancestor approximately 15 000 years ago, possibly owing to isolation during a recent glacial advance. The continental steamer duck species are not reciprocally monophyletic, but show some amount of genetic differentiation between them. Each lineage of Tachyeres represents a different stage between flight and flightlessness. Their phylogenetic relationships suggest multiple losses of flight and/or long-term persistence of mixed-flight capability. As such, steamer ducks may provide a model system to study the evolution of flightlessness.