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Results

We found 40 Least Concern and Data Deficient species that possess a high combined score of λM and ED (see Table 1). In total, 42 of the island mammal species we assessed were listed by the IUCN as Data Deficient, 47 as Least Concern, with the remainder as threatened species. Those species already listed as threatened were potentially suffering from other threats (e.g. non-native species as predators/competitors). Focusing on those species that are Data Deficient or Least Concern and have higher λM-ED scores would be most beneficial, as their rarity indicate them to be at risk and a high λM value represents an important patch, and one that would pay off greatly to conserve.

The five islands with the highest average λM-ED scores, taken by adding all the scores and dividing by our (island-restricted mammals) species richness per island were Jamaica, Guadalcanal, Isle of Pines, Madagascar, and Nggela Sule (see Table 2, Fig. 3 for map). Interestingly, Madagascar held 39 of the highest λM-ED species, and ranked fourth in our λM-ED islands list.

We found that combining evolutionary distinctness with λM revealed species that may be of concern that were not otherwise noticed. Because quantifying fragmentation effects on species takes into account spatial configuration, this can help to improve threat status assessments. The EDGE programme has already sought to visualize regions in the world with the most rare species and moved to prioritize those particular species. This adds a spatial understanding of the species distribution to that prioritization.

Table 1 Top 10 species in order of decreasing λM-ED score

Table 2 Top 10 islands, in order of decreasing λM-ED score

Fig. 3 Map highlighting the top five islands, coloured from warm to cool (i.e. red to blue), in decreasing λM-ED score (see also Table 2)

Discussion

Summary

We found Least Concern and Data Deficient island-restricted mammals that possess a high combined score of λM and ED. This method can be the start to finding species with a combination of phylogenetic rarity and long-term extinction risk due to island isolation. Further analyses are needed, as global prioritizations risk overgeneralizing among distinct animals, and yet suitable datasets, spatial and otherwise, are difficult to come by.

Island Studies

Islands represent less than 5 % of the earth's land area, harbour 80 % of known species extinctions since 1500 (Ricketts et al. 2005), and make up 39 % of today's IUCN Critically Endangered species (TIB 2012). Endangered island species, such as those targeted and listed in the Threatened Island Biodiversity (TIB) database, are currently of major concern due to invasive species. However, we can still examine the effects of isolation and area from an island point of view. On a global scale, this method aims to show which islands or species are most important for conservation, based on the spatial properties of the islands and the phylogenetic rarity of the species themselves.

Islands are a natural laboratory for evolutionary specialization and adaptation, because such an environment greatly shapes the select set of species living there in such isolation (Losos and Ricklefs 2009). From a conservation perspective, islands are unique because with less spatial area to begin with, they can only support smaller populations to evolve on them (Diamond 1975; Frankham 1998). Furthermore, recolonisation, the process responsible for maintaining population size from a larger source population, decreases because of spatial isolation and size (MacArthur and Wilson 1963, 1967; Simberloff and Wilson 1970), and dispersal amongst islands can be far more limited than on terrestrial “islands”. We expect that islands suffer more from stochastic extinction processes, in addition to anthropogenic effects such as introduced species, so they are on the whole in much greater need of immediate conservation action. In fact, islands have previously been the focus of research on prioritisation schemes for conservation planning (TIB 2012).

However, much complexity remains in studying islands. Most threatened species have small geographic distributions, and the distributions of island species are inevitably smaller than the distributions of continental species (Manne et al. 1999). Yet, some island populations can “show greater persistence than mainland populations of the same species, notwithstanding their smaller range sizes” (Channell and Lomolino 2000), perhaps reflecting the advantages of living in sheltered isolation. Another study found that island endemics are not relatively more threatened than continental ones, considering their distribution size, “suggesting that evolutionary isolation is not the reason for their vulnerability” (Purvis et al. 2000). Perhaps unravelling isolation and evolutionary factors can lead to a greater understanding of the unique state that island animals seem to occupy.

Small distribution area and island endemicity were the most important predictors of mammal extinction risk found through literature survey (Purvis et al. 2000). Because of such isolation, we would expect evolutionary history to reflect the spatial fragmentation. Moreover, there is a certain importance to the isolation of islands, given the limits of animal dispersal (Diamond 1974). For instance, the number of threatened endemic bird species has been found to correlate with deforestation on islands, and single-island endemics are considerably more at risk than more widespread species (Brooks et al. 1997), hence examining spatial aspects of islands is a sensible route.

Islands, particularly larger ones, are likely to contain multiple landscape types, and our islands borders, although defined at high resolution by GSHHS, can likely overestimate the amount of suitable habitat for a species. For instance, we found Madagascar ranked fourth in our list, but including additional information would scale down the habitat size from islands to the actual size of primary habitat. Then Madagascar might very well outrank all the other islands, due to unique species that possess ranges limited to parts of the island. With species records from GBIF and publicly available environmental layers, we could perhaps improve on this by creating approximate species distribution “maps” that we might be able to prune down the current IUCN extent of occurrence maps to a more realistically “fragmented” habitat extent. Calculating the λM of such maps would be an improved and more realistic estimate as to long-term species persistence.

It might be that island species have some adaptation for having historically small isolated populations, such that the little area available has shaped the species' phylogeny (Cardillo et al. 2008). On the other hand, age of the islands (equivalently, patches) might have a significant influence on metapopulation persistence (Hastings 2010). It could also be that the most sensitive species were previously driven to extinction and modern day survivors have already been selected for (Manne et al. 1999). Human impact cannot be overestimated, because despite exceptional habitat loss on all terrestrial land types, “the human impact index” was considerably greater on islands (Kier et al. 2009). It is still a puzzle to be teased apart, how the interaction of intrinsic factors, e.g. innate biological susceptibility, and extrinsic factors, i.e. those mostly due to human impact, affect the outcome that ultimately leads to extinction (Bennett and Owens 1997).

Already there are numerous efforts underway to stave off the extinction of island species, such as the previously mentioned Threatened Island Biodiversity (TIB) database (tib.islandconservation.org/), whose primary focus is on eradicating threatening non-natives. The high levels of endemic richness already warrant special conservation protection (Kier et al. 2009). Species on continents can experience island effects, e.g. mountains or islands within lakes, which would still make island conservation studies, such as this, applicable to them.

 
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