Island evolutionary biogeography: analysis of the weevils (Coleoptera: Curculionidae) of the Falkland Islands (Islas Malvinas)

Aim I analysed distributional and phylogenetic information on weevils (Coleoptera: Curculionidae) from the Falklands, and integrated it with molecular, palaeontological and geological information to infer a geobiotic scenario. Location Falkland Islands (Islas Malvinas). Methods The panbiogeographical analysis was based on data on 23 Falkland species and their related taxa from southern South America. For the cladistic biogeographical analysis I analysed six weevil taxa for which phylogenetic hypotheses are available (the generic groups Cylydrorhinus, Strangaliodes and Falklandius, and the genera Antarctobius, Germainiellus and Puranius). Results from this analysis were compared with previous regionalizations. Cenocrons (sets of taxa that share the same biogeographical history) were identified by considering temporal information provided by fossils and molecular clocks. Finally, a geobiotic scenario was proposed by integrating the available information. Results Six generalized tracks were detected: Maule–Valdivian forests, Magellanic forest, Magellanic moorland, Falkland Islands, Magellanic forest–Magellanic moorland, and Magellanic forest–Falkland Islands. A node was identified in the Magellanic forest, based on the overlap of two generalized tracks. A single general area cladogram was obtained, implying the following sequence: (Magellanic moorland (Maule–Valdivian forests (Magellanic forest, Falkland Islands))). The Falklands are classified here as a biogeographical province in the Austral realm, Andean region and Subantarctic subregion. Falkland weevils seem to belong to a single Subantarctic cenocron. The sequence of events deduced implies the following steps: development of the Subantarctic biota in southern South America, arrival of the Falkland crustal block from South Africa in the Early Cretaceous, geodispersal of the Subantarctic cenocron from southern South America to the Falklands during the Early Oligocene, vicariance of the Magellanic moorland, vicariance of the Maule–Valdivian forests, and final vicariance between the Magellanic forest and the Falkland Islands. Main conclusions The biotic components identified support the connection of the Falkland weevils with the Magellanic forest. Falkland weevils belong to a single cenocron, dated to at least the Early Oligocene, when geodispersal from southern South America may have occurred. An older African cenocron may have been replaced completely by the Subantarctic one when the proto‐Falklands made contact with the Patagonian continental shelf. A geobiotic scenario implying vicariance events related to sea‐level variations could explain the distributional patterns analysed herein.     

Divergence between passerine populations from the Malvinas – Falkland Islands and their continental counterparts: a comparative phylogeographical study

Bursts of speciation have followed colonization of remote oceanic islands by diverse taxa, a process evidenced by island endemics around the world. The present study explores whether the Malvinas – Falkland Islands (MFI), a relatively understudied archipelago off the South Atlantic coast of Patagonia, harbour endemic genetic lineages of passerine birds. Nine passerine species nest regularly in the MFI (Cinclodes antarcticus, Muscisaxicola maclovianus, Troglodytes cobbi, Cistothorus platensis, Turdus falcklandii, Anthus correndera, Melanodera melanodera, Sturnella loyca, and Carduelis barbata). Mitochondrial DNA (cytochrome c oxidase I sequences) are used to quantify and compare divergence between insular and continental populations, finding genetic patterns to vary across these nine species. Most MFI passerines do not show significant genetic differentiation from continental populations, whereas C. platensis, M. melanodera, and T. falcklandii are modestly diverged. Finally, T. cobbi differes markedly from its closest continental relative Troglodytes aedon, a result that is confirmed using nuclear and vocal data. The study also identifies broadly divergent lineages within continental populations of C. platensis and T. aedon. Taken together, these results suggest that the land bird populations of the MFI were established at different times. Troglodytes cobbi is the oldest MFI land bird, splitting from continental T. aedon during the Great Patagonian Glaciation of the Pleistocene. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, •• , ••–••.     

Winter ecology of house mice and the prospects for their eradication from Steeple Jason (Falkland Islands)

Invasive house mice Mus musculus are known to impact on seabird, invertebrate and plant communities on temperate and subantarctic islands, particularly where they are the sole rodent species. Steeple Jason, in the Falkland Islands, is an island which supports globally important seabird populations as well as introduced mice. To evaluate the prospects for mouse eradication, we investigated mouse ecology and undertook bait uptake trials on Steeple Jason in late winter. Mice were present in all habitats but were most abundant in tussac Poa flabellata where they occurred at 20–35 mice ha^?1. From 58 mature perforate females, 16 % were pregnant, with litters of 4–8 pups. The first lactating female was caught at the end of August, suggesting that breeding had recently begun. Bait trials replicating an aerial eradication were undertaken on two trapping grids of 7.7 and 6.8 ha, with non-toxic pellets containing the biomarker pyranine spread at 7.5–7.7 kg ha^?1. All 447 mice captured after baiting had consumed bait. The relatively low winter density, distribution and biology of house mice on Steeple Jason are similar to those observed before other successful mice eradications, and the study indicated 100 % bait acceptance. Before an eradication attempt, we suggest investigating whether breeding ceases completely earlier in the winter and urge careful consideration of non-target species.     

Going native,going local: revegetating eroded soils on the Falkland Islands using native seeds and farmland waste

Remote island ecosystems are vulnerable to human disturbance and habitat destruction, yet they often have limited capacity to revegetate degraded habitats, especially with native species. To revegetate degraded island habitats, practitioners often rely on importing non‐native species, thereby increasing the number of introduced species on islands. In this study, we investigated the effectiveness of sowing wild collected native seeds and locally sourced treatments for revegetating different eroded soil types (clay, peat, and sand) across the Falkland Islands. A seed mixture of 15 native species was sown with different supportive treatments (sheep dung, sheep dags [woolly off‐cuts], and geotextile matting [coir]) and their combinations. After 1 year, native seeds provided up to 70% plant cover and accrued 1.98 kg/m² in biomass. Three key native species Elymus magellanicus, Poa flabellata, and Poa alopecurus occurred in 64, 50, and 50% of all sown plots. However, supportive treatments equally facilitated the colonization and establishment of non‐native species. At the same time, there was no difference in native plant cover and biomass across different treatments or soil types, although in the absence of supportive treatments there was little to no revegetation. Thus, locally sourced treatments (i.e. sheep dung and dags) may provide an equally effective but low‐cost alternative to imported treatments (i.e. geotextiles). We further discuss challenges of integrating revegetation using native seeds and livestock grazing on the Falkland Islands. Our study demonstrates that native species and local treatments can provide a rapid approach to revegetating degraded island habitats.     

Photosynthesis and water‐use efficiency: A comparison between invasive (exotic) and non‐invasive (native) species

Abstract Invasive species have been hypothesized to out‐compete natives though either a Jack‐of‐all‐trades strategy, where they are able to utilize resources effectively in unfavourable environments, a master‐of‐some, where resource utilization is greater than its competitors in favourable environments, or a combination of the two (Jack‐and‐master). We examined the invasive strategy of Berberis darwinii in New Zealand compared with four co‐occurring native species by examining germination, seedling survival, photosynthetic characteristics and water‐use efficiency of adult plants, in sun and shade environments. Berberis darwinii seeds germinated more in shady sites than the other natives, but survival was low. In contrast, while germination of B. darwinii was the same as the native species in sunny sites, seedling survival after 18 months was nearly twice that of the all native species. The maximum photosynthetic rate of B. darwinii was nearly double that of all native species in the sun, but was similar among all species in the shade. Other photosynthetic traits (quantum yield and stomatal conductance) did not generally differ between B. darwinii and the native species, regardless of light environment. Berberis darwinii had more positive values of δ¹³C than the four native species, suggesting that it gains more carbon per unit water transpired than the competing native species. These results suggest that the invasion success of B. darwinii may be partially explained by combination of a Jack‐of‐all‐trades scenario of widespread germination with a master‐of‐some scenario through its ability to photosynthesize at higher rates in the sun and, hence, gain a rapid height and biomass advantage over native species in favourable environments.     

Variation in the diet of Genypterus blacodes(Ophidiidae) around the Falkland Islands

The diet change with size, season and area was investigated using the stomachs of 496 kingclip Genypterus blacodes collected around the Falkland Islands (south‐west Atlantic) between August 2001 and September 2002. The key prey species were rockcod Patagonotothen spp., benthic isopods and Patagonian grenadier Macruronus magellanicus . Kingclip <50 cm total length ( L _T) fed mainly on crustaceans and small fishes. With size the diet shifted away from crustaceans towards Patagonotothen spp. in kingclip 50–100 cm L _T, and finally towards larger fishes such as M. magellanicus and Micromesistius australis australis in kingclip >100 cm L _T. The niche breadth was highest in fish >100 cm L _T and the lowest in fish <50 cm L _T. The larger kingclip generally selected larger individuals of the same prey species, with the exception of the Patagonian squid Loligo gahi , where all ingested squid were of similar size, regardless of the predator length. The importance of the main prey species varied substantially between five consequent seasons studied, and appeared to follow the seasonal abundance and availability of prey. The spatial variability in the diet was found in kingclip caught in regions occupied by transformed temperate and sub‐Antarctic waters. The rockcod, which is available throughout the year around the Falkland Islands, was the most important prey in the kingclip diet. Kingclip takes advantage of other seasonally abundant prey species during their seasonal migrations ( e.g . L. gahi ) and also scavenge on discards from fishing vessels when available.     

Effects of desiccation on native and non‐native molluscs in rivers

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Biocontrol: the response of native oystercatchers to a non‐native clam invasion

Global change, including invasive species introduction, has already had observable effects on migrant bird species, from northern breeding areas to wintering grounds. In this study we analyze the response of the Eurasian oystercatcher abundance to the density of an invasive clam species (Corbicula fluminea) and its potential role as biological control. As a case study, the oystercatcher population fluctuations over a 30‐yr time period, coupled with video‐recorded estimates of its feeding behavior on C. fluminea, and results from an exclosure experiment, were analyzed in the NW of the Iberian Peninsula. Results showed that oystercatchers exert a top‐down control over C. fluminea density. In addition, oystercatchers doubled its wintering numbers in a C. fluminea invaded estuary where they actively feed upon this invasive clam. Given that, the facilitative interaction between the invasive C. fluminea and the migratory Eurasian oystercatcher seems to respond to bottom‐up forces. Altogether, our results suggest that control measures applied to long term biological invasions must be carefully analyzed since non‐native species may be sustaining dependent native communities.     

Self‐incompatibilty in gamete recognition: Single self‐recognizing determinants and multiple,non‐self‐recognizing ones function in the same individual

The frameworks (key mechanisms) of the self/non‐self‐discrimination systems that are found in various organisms have not been actively selected for, but have evolved by genetic drift such that the genetic frequency of random, advantageous mutations has increased within the genomes of these species by natural selection. The passive nature of this process leads to an important conclusion: in the self/non‐self‐discrimination system, the number of self‐recognizing determinants becomes one compared to multiple non‐self‐recognizing determinants. Thus, the number of determinants is defined not by the character of the determinant, but by the system framework. Mol. Reprod. Dev. 80: 2–7, 2013. © 2012 Wiley Periodicals, Inc.     

Echoes of the whispering land: interacting roles of vicariance and selection in shaping the evolutionary divergence of two <Emphasis Type="Italic">Calceolaria</Emphasis> (Calceolariaceae) species from Patagonia and Malvinas/Falkland Islands

A key to understanding the origin and identity of young species lays on the knowledge of the Quaternary climatic oscillations’ effect on gene flow and vicariance. Even though the effect of climatic fluctuations is relatively well understood for southern hemisphere plant species, little is known about their effect on the evolutionary histories of species from mainland and islands. Thus, we investigated whether Quaternary climate-driven fluctuations translated into lineage divergence and speciation, followed or not by climatic niche differentiation, in two allopatric plant species, Calceolaria uniflora and C. fothergillii from Patagonia and Malvinas/Falkland islands, respectively. We sampled the range of both species, and sequenced two chloroplastic (cpDNA; trnS–trnG and trnH–psbA), and one single copy “anonymous” non-coding nuclear region (nDNA). We performed phylogeographic and dating analyses, and adjusted spatio-temporal diffusion models. We complemented molecular evidence with climatic niche differentiation analyses and species paleo-distribution projections. A species coalescent reconstruction based on multi-locus data retrieved both species as monophyletic. Estimates from cpDNA indicated the species diverged during the Great Patagonian Glaciation. Chloroplast and nuclear DNA showed east–west distribution of the main genetic groups but with contrasting spatial genetic diversity. The spatio-temporal diffusion analyses showed that between 1–0.8 Mya and 570 Kya the lineage leading to C. fothergillii diverged from C. uniflora and arrived to the islands. Climatic niche projections hindcasted range expansions during glaciations, and contractions during the interglacial periods. Comparisons of climatic niches between the two study species indicated that temperature variables show evidence of niche conservatism while precipitation regimes supported niche divergence, even when considering the background environmental divergence. Our study indicates that glacial fluctuations affected the mainland/islands connections favouring speciation mediated not only by isolation, but also by climatic niche differentiation.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Reproductive biology of Patagonotothen ramsayi (Regan, 1913) (Pisces: Nototheniidae) around the Falkland Islands

The reproductive biology of one of the most abundant notothenioids, Patagonotothen ramsayi, was investigated between February 2003 and November 2004 on the Falkland and Patagonian Shelves (Southwest Atlantic). Male and female P. ramsayi were mature at 27.6 and 24.8 cm L _T, respectively. P. ramsayi is a total spawner, with a total fecundity ranging between 24,300 and 76,700 eggs. Spawning occurs on the shelf breaks between June and August with the peak in gonado-somatic indices in June. Analysis of length frequency distributions over the year and sex ratios of mature fish during the spawning season may indicate the presence of a nesting and nest guarding behaviour in male fish, similar to other rockcods. Features of its reproductive strategy, which enabled P. ramsayi to dominate the medium-sized demersal fishes on the Patagonian shelf are discussed and compared with those of other nototheniids and cottid sculpins from the Northern Hemisphere.     

Parasite fauna of native and non‐native populations of Neogobius melanostomus (Pallas, 1814) (Gobiidae) in the longitudinal profile of the Danube River

Parasite fauna of round goby Neogobius melanostomus (Pallas, 1814) in the Danube River was investigated in both its native range (two sites in the Bulgarian stretch of the Danube) and non‐native range of distribution (Croatian, Slovak and Austrian stretches) during 2005 and 2006. The aim was to identify possible changes in parasite communities associated with the introduction of a host into the new environment. A total of 29 metazoan parasite species were found to parasitize round goby in the Danube River; twelve of these parasite species were found in both the native and non‐native range of distribution. Introduction of a novel parasite species to the non‐native range via the round goby was not found. Eight parasite species occurred only in the native range and nine species only in the non‐native range of the round goby distribution. Losses of native parasite species in non‐native round goby populations and/or acquiring of novel parasite species in a new environment were not significant. Thirteen parasite taxa were recorded for the first time in round gobies. Three parasite taxa (Diplostomum spp., Pomphorhynchus laevis and Raphidascaris acus) were found in high prevalence and abundance at each sampling site in both the native and non‐native range. Parasite species diversity was assessed for each sampling site and season using three diversity indices (the Shannon, Simpson and Equitability indices), with the highest same‐season values found in a non‐native site in Slovakia (1.38, 0.69 and 0.60, respectively) and the lowest in a native site in Bulgaria (0.28, 0.12 and 0.14, respectively). Species diversity was higher in both non‐native round goby populations (Slovak and Austrian) compared to native Bulgarian populations. However, diversity indices values varied among almost all sampling sites.     

The good with the bad: when ecological restoration facilitates native and non‐native species

Organisms interact with each other along a spectrum ranging from competition to facilitation. A theme in restoration ecology is tipping the balance of these interactions to favor desired species and site conditions, exemplified by restoring fertile islands and their nurse plant effects to encourage plant recruitment. We tested the effectiveness of outplanting nursery‐grown native perennials and vertical mulching (placing dead plant material upright in soil) for stimulating annual plant recruitment in a disturbed Mojave Desert shrubland in Joshua Tree National Park, California, U.S.A. Over 9 years, differences in annual species richness and cover between interspaces and below outplants and vertical mulch varied among years, potentially via inter‐annual fluctuations in precipitation or maturation of restoration sites. In the ninth year, which was the wettest, both native and non‐native cover averaged 3× higher below outplants than in interspaces. Overall among years at the microsite scale, non‐native annual plants more consistently exploited environments provided by outplants and vertical mulch structures than did native annuals. However, these restoration structures were important for native annual diversity. At the 40‐m² plot scale, disturbed plots that received outplanting supported greater richness of native annual species than disturbed unrestored plots. By facilitating both non‐native and native plants, reestablishing fertile islands to restore dryland ecosystems is a conundrum for restoration. Treatments reducing non‐native plants may need to accompany fertile island restoration to tip the balance of facilitative plant interactions in favor of native species.     

Differential temporal beta‐diversity patterns of native and non‐native arthropod species in a fragmented native forest landscape

An important factor that hinders the management of non‐native species is a general lack of information regarding the biogeography of non‐natives, and, in particular, their rates of turnover. Here, we address this research gap by analysing differences in temporal beta‐diversity (using both pairwise and multiple‐time dissimilarity metrics) between native and non‐native species, using a novel time‐series dataset of arthropods sampled in native forest fragments in the Azores. We use a null model approach to determine whether temporal beta‐diversity was due to deterministic processes or stochastic colonisation and extinction events, and linear modelling selection to assess the factors driving variation in temporal beta‐diversity between plots. In accordance with our predictions, we found that the temporal beta‐diversity was much greater for non‐native species than for native species, and the null model analyses indicated that the turnover of non‐native species was due to stochastic events. No predictor variables were found to explain the turnover of native or non‐native species. We attribute the greater turnover of non‐native species to source‐sink processes and the close proximity of anthropogenic habitats to the fragmented native forest plots sampled in our study. Thus, our findings point to ways in which the study of turnover can be adapted for future applications in habitat island systems. The implications of this for biodiversity conservation and management are significant. The high rate of stochastic turnover of non‐native species indicates that attempts to simply reduce the populations of non‐native species in situ within native habitats may not be successful. A more efficient management strategy would be to interrupt source‐sink dynamics by improving the harsh boundaries between native and adjacent anthropogenic habitats.     

Management intensity affects the relationship between non‐native and native species in subtropical wetlands

Question: Does management intensity affect the association between non‐native and native species and between non‐native species and soil nutrients in wetlands? Location: MacArthur Agro‐Ecology Research Center, Florida, USA. Methods: We evaluated native and non‐native plant richness and relative frequency in 15 1‐m² plots in 40 wetlands across two types of pastures, highly managed (fertilized, ditched, planted, heavily grazed by cattle) and semi‐natural (unfertilized, lightly seasonally grazed). Plant biomass was collected in five 0.25‐m² plots per wetland and sorted to species. Soil cores were collected to analyse soil total nitrogen (N) and phosphorus (P). An information‐theoretic approach was used to compare mixed effects models considering the association of non‐native richness, relative frequency, and biomass with native richness, relative frequency, biomass, C₃ grass relative frequency (a dominant native group), N, P and wetland‐type. Results: Non‐native richness was negatively correlated with native richness in semi‐natural wetlands, but there was no evidence of an association between these variables in highly managed wetlands. Non‐native richness increased with increasing soil N in semi‐natural wetlands, but not in the highly managed wetlands. Soil P was positively related to non‐native frequency in semi‐natural wetlands but negatively related in highly managed wetlands. Non‐native frequency and biomass were negatively related to relative frequency of C₃ grasses in both management types. Conclusions: Our results indicate that management intensity influences relationships between native and non‐native richness. Management intensity interacts with abiotic or biotic factors, such as soil nutrients and composition, in predicting where non‐native species will most likely need control.     

Two spiders in subfamily Mynogleninae (Araneae: Linyphiidae) from the Falkland Islands, South Atlantic

Two new taxa, Falklandoglenes spinosa gen. nov. and sp. nov. and Beauchenia striata gen. nov. and sp. nov. , are described from Beauchêne Island, the most remote island in the Falkland Islands archipelago. Both species are in subfamily Mynogleninae, previously thought to be confined to Central Africa and to New Zealand and its neighbouring subantarctic islands. Both species show intermediate characters between the Mynogleninae and the remainder of the Linyphiidae, and hence they throw some light on the phylogeny of this large, complex, world–wide family of spiders.