Re‐approaching the small island effect

Aim To propose a new approach to the small island effect (SIE) and a simple mathematical procedure for the estimation of its upper limit. The main feature of the SIE is that below an upper size threshold an increase of species number with increase of area in small islands is not observed. Location Species richness patterns from different taxa and insular systems are analysed. Methods Sixteen different data sets from 12 studies are analysed. Path analysis was used for the estimation of the upper limit of the SIE. We studied each data set in order to detect whether there was a certain island size under which the direct effects of area were eliminated. This detection was carried out through the sequential exclusion of islands from the largest to the smallest. For the cases where an SIE was detected, a log‐log plot of species number against area is presented. The relationships between habitat diversity, species number and area are studied within the limits of the SIE. In previous studies only area was used for the detection of the SIE, whereas we also encompass habitat diversity, a parameter with well documented influence on species richness, especially at small scales. Results An SIE was detected in six out of the 16 studied cases. The upper limit of the SIE varies, depending on the characteristics of the taxon and the archipelago under study. In general, the values of the upper limit of the SIE calculated according to the approach undertaken in our study differ from the values calculated in previous studies. Main conclusions Although the classical species–area models have been used to estimate the upper limit of the SIE, we propose that the detection of this phenomenon should be undertaken independently from the species–area relationship, so that the net effects of area are calculated excluding the surrogate action of area on other variables, such as environmental heterogeneity. The SIE appears when and where area ceases to influence species richness directly. There are two distinct SIE patterns: (1) the classical SIE where both the direct and indirect effects of area are eliminated and (2) the cryptic SIE where area affects species richness indirectly. Our approach offers the opportunity of studying the different factors influencing biodiversity on small scales more accurately. The SIE cannot be considered a general pattern with fixed behaviour that can be described by the same model for different island groups and taxa. The SIE should be recognized as a genuine but idiosyncratic phenomenon.     

Partitioning the sources of demographic variation reveals density‐dependent nest predation in an island bird population

Ecological factors often shape demography through multiple mechanisms, making it difficult to identify the sources of demographic variation. In particular, conspecific density can influence both the strength of competition and the predation rate, but density‐dependent competition has received more attention, particularly among terrestrial vertebrates and in island populations. A better understanding of how both competition and predation contribute to density‐dependent variation in fecundity can be gained by partitioning the effects of density on offspring number from its effects on reproductive failure, while also evaluating how biotic and abiotic factors jointly shape demography. We examined the effects of population density and precipitation on fecundity, nest survival, and adult survival in an insular population of orange‐crowned warblers (Oreothlypis celata) that breeds at high densities and exhibits a suite of traits suggesting strong intraspecific competition. Breeding density had a negative influence on fecundity, but it acted by increasing the probability of reproductive failure through nest predation, rather than through competition, which was predicted to reduce the number of offspring produced by successful individuals. Our results demonstrate that density‐dependent nest predation can underlie the relationship between population density and fecundity even in a high‐density, insular population where intraspecific competition should be strong.     

Long‐term non‐equilibrium dynamics of insular floras: a 17‐year record

Aim To document long‐term rates of immigration, extinction and turnover in insular floras and evaluate the relative impacts of recent hurricane activity and climate change. Location Three archipelagos of small islands, in the Exuma Cays, Andros and Abacos, Bahamas. Methods I surveyed the floras of 194 vegetated islands in three archipelagos over several multi‐year periods, spanning up to 17 years. Changes in abundance (foliar cover) of persistent populations were measured on a subset of 14 islands in the Exuma Cays over a 9‐year period. Results Rates of plant turnover were generally low compared with other organisms, but varied among archipelagos and time periods. Turnover rates were usually higher in the second decade of this study, and extinction rates were often dramatically higher than immigration rates in the second decade, resulting in overall decreases in species richness. Turnover did not differ significantly among island types based on generalized location and surrounding water depths, and extinctions were not more likely to occur on more exposed islands. The abundance (foliar cover) of populations that did not go extinct decreased steadily over the second decade of this study, indicating, along with higher extinction rates, a generalized decline in these insular floras. Main conclusions Although some islands may have been at or near a state of dynamic equilibrium in the first decade of this study, average species richness declined in all three archipelagos during the second decade, when extinctions greatly outnumbered immigrations. Four major hurricanes affected the study archipelagos in the second decade of this study, although the available evidence suggests that the hurricanes were not directly responsible for the declines. Indirect effects of hurricanes such as increased herbivory and possible decreased nutrient availability, along with a long‐term (25 years) increase in temperature and decline in rainfall are likely contributing factors.     

Island shape,size, and isolation affect nest‐site selection by Little Terns

ABSTRACT Populations of many seabirds and other species that nest along coasts are declining due to habitat degradation and loss. An improved understanding of the species‐specific factors that determine nest density across a landscape is therefore critical for conservation efforts. We examined factors that affected the density (number per hectare) and abundance (number at a sampling site) of nests of Little Terns (Sternula albifrons) on the Sinai Peninsula, Egypt. Terns preferred to nest on islands rather than the mainland, with islands constituting 64% of the area surveyed, but containing 99% of the 439 tern nests we found. Nest densities were highest on islands that were small, located at moderate distances from the mainland, and irregularly shaped or elongated. Most nests (69%) were on islands with areas < 3 ha, although these islands represented < 5% of total island area, and islands with the highest nest densities were 80–300 m from the mainland. Terrestrial predators were more likely to occur on larger islands, visiting three of the largest four islands. Most tern nests were within 1 m of shorelines, causing island perimeter to be a strong influence on nest density. Island shape was the only factor that significantly affected nest abundance, with more nests on islands with relatively long perimeters for their size. Our results suggest that protection or creation of relatively small, slender islands at moderate distances from shore may be an effective means of increasing the number of breeding sites for Little Terns. Although not generally considered a potential determinant of nest site preferences for seabirds, island shape is likely to be important for species that prefer sites adjacent to water, including species that nest on beaches and seaside cliffs.     

Evolutionary size changes in plants of the south‐west Pacific

Aim To investigate evolutionary changes in the size of leaves, stems and seeds of plants inhabiting isolated islands surrounding New Zealand. Location Antipodes, Auckland, Campbell, Chatham, Kermadec, Three Kings and Poor Knights Islands. Methods First, we compared the size of leaves and stems produced by 14 pairs of plant taxa between offshore islands and the New Zealand mainland, which were grown in a common garden to control for environmental effects. Similar comparisons of seed sizes were made between eight additional pairs of taxa. Second, we used herbarium specimens from 13 species pairs to investigate scaling relationships between leaf and stem sizes in an attempt to pinpoint which trait might be under selection. Third, we used herbarium specimens from 20 species to test whether changes in leaf size vary among islands located at different latitudes. Lastly, we compiled published records of plant heights to test whether insular species in the genus Hebe differed in size from their respective subgenera on the mainland. Results Although some evidence of dwarfism was observed, most insular taxa were larger than their mainland relatives. Leaf sizes scaled positively with stem diameters, with island taxa consistently producing larger leaves for any given stem size than mainland species. Leaf sizes also increased similarly among islands located at different latitudes. Size changes in insular Hebe species were unrelated to the average size of the respective subgenera on the mainland. Main conclusions Consistent evidence of gigantism was observed, suggesting that plants do not obey the island rule. Because our analyses were restricted to woody plants, results are also inconsistent with the ‘weeds‐to‐trees’ hypothesis. Disproportionate increases in leaf size relative to other plant traits suggest that selection may favour the evolution of larger leaves on islands, perhaps due to release from predation or increased intra‐specific competition.     

A GIS‐based decision‐making approach for prioritizing seabird management following predator eradication

Given that 29% of seabird species are threatened with extinction, protecting seabird colonies on offshore islands is a global conservation priority. Seabirds are vulnerable to non‐native predator invasions, which reduce or eliminate colonies. Accordingly, conservation efforts have focused on predator eradication. However, affected populations are often left to passively recover following eradications. Although seabirds are highly mobile, their life history traits such as philopatry can limit passive recolonization of newly predator‐free habitat. In such cases, seabird colonies can potentially be re‐instated with active restoration via chick translocations or social attraction methods, which can be risky and expensive. We used biogeographic and species‐specific behavioral data in the Hauraki Gulf, New Zealand, a global hotspot of seabird diversity and predator eradications, to illustrate the use of geographic information systems multi‐criteria decision analysis to prioritize islands for active seabird restoration. We identified nine islands with low observed passive recovery of seabirds posteradication over a 50‐year timeframe, and classified these as sites where active seabird management could be prioritized. Such spatially explicit tools are flexible, allowing for managers to choose case‐specific criteria such as time, funding, and goals constrained for their conservation needs. Furthermore, this flexibility can also be applied to threatened species management by customizing the decision criteria for individual species' capacity to passively recolonize islands. On islands with complex restoration challenges, decision tools that help island restoration practitioners decide whether active seabird management should be paired with eradication can optimize restoration outcomes and ecosystem recovery.     

Environmentally‐ and human‐induced body‐size responses in Macropus robustus and Macropus rufus,two widespread kangaroo species with largely overlapping distributions

Progressive body‐size dwarfing of animal populations is predicted under chronic mortality stress, such as that inflicted by human harvesting. However, empirical support for such declines in body size due to elevated mortality is lacking. In fact, the size of three macropodid species ─ the two grey kangaroo species, Macropus fuliginosus and M. giganteus, and the Red‐necked Wallaby, M. rufogriseus ─ appears to have increased since European settlement in Australia, despite these species being subjected to size‐selective harvesting over this period. To test whether this unexpected trend also characterises other species, we sought evidence of human‐induced body‐size changes in the two most widely distributed kangaroo species, the Euro Macropus robustus and Red Kangaroo M. rufus, from the late 19th Century onwards. Spatial autoregressive models controlling for age, sex and island effects were first used to identify environmental predictors of body size and to evaluate multi‐causal explanations for spatial body‐size patterns. Primary productivity emerged as the key driver of body size in both species, while heat conservation was supported as a further mechanism explaining the large body size of M. robustus in cold climatic regions. After controlling for these environmental factors, we find that the size of M. rufus has been stable over time and limited support for a small increase in the size of M. robustus. Hence, there is no empirical evidence that contemporary size‐selective harvesting has reduced body size in these species. Rather, the latter result supports the possibility that pasture improvement and/or dingo control (and associated reduction in predation pressure) facilitated body‐size increases following European settlement in Australia.     

Trapped in the web of water: Groundwater‐fed springs are island‐like ecosystems for the meiofauna

We investigated whether the equilibrium theory of island biogeography (ETIB) can be applied to the meiofauna of groundwater‐fed springs. We tested whether copepod species richness was related with spring area, discharge, and elevation. Additionally, five hypotheses are tested based on species distribution patterns, dispersal ability, and life‐history characteristics of several guilds (stygobiotic, nonstygobiotic, cold stenotherm, and noncold stenotherm species). Thirty springs in the central Apennines (Italy) were considered. A multimodel selection procedure was applied to select best‐fit models using both ordinary least‐squares regressions and autoregressive models. Mantel tests were used to investigate the impact of spatial autocorrelation in determining interspring similarity (ßsor), pure turnover (ßsim), intersite nestedness (ßnest = ßsor ? ßsim), and matrix nestedness (measured using NODF and other metrics). Explicit consideration of spatial correlations reduced the importance of predictors of overall species richness, noncold stenotherm species (both negatively affected by elevation), cold stenotherm species, and nonstygobiotic species, but increased the importance of area for the stygobiotic species. We detected nested patterns in all cases, except for the stygobites. Interspring distances were positively correlated with ßsor and ßnest (but not with ßsim) for the entire data set and for nonstygobiotic, cold stenotherm, and noncold stenotherm species. In the case of stygobites, interspring geographical distances were marginally correlated with ßsor and no correlation was found for ßsim and ßnest. We found support for ETIB predictions about species richness, which was positively influenced by area and negatively by elevation (which expresses the size of source of immigrants). Low turnover and high nestedness are consistent with an equilibrium scenario mainly regulated by immigration and extinction. Stygobites, which include many distributional and evolutionary relicts, have a low capability to disperse through the aquifers and tend to be mainly confined to the springs where they drifted out and were trapped by springbed sediments.     

Comment on ‘Island biogeography: patterns of marine shallow‐water organisms’ by Hachich et al., Journal of Biogeography (2015)

In a recent article, Hachich et al. (2015, Journal of Biogeography, 42 , 1871–1882) studied the large‐scale biogeographical patterns of the species–area, species–island age and species–isolation relationships associated with marine shallow‐water groups (reef fish, gastropods and seaweeds) from 11 Atlantic archipelagos. We here express our concerns regarding the data accuracy used to compute the different models that tested the null hypothesis of species richness being independent of the selected variables. In our commentary, we focus mainly on the use of out‐of‐date checklists of gastropod and seaweed species from different archipelagos, but we also point out inaccuracies in some island age estimates and explain our disagreement with the use of the 200 m depth limit for the shallow‐water gastropods and seaweeds.     

Genomic and morphological data help uncover extinction‐in‐progress of an unsustainably traded hill myna radiation

The Asian songbird crisis which is currently unfolding in Southeast Asia has seen multiple bird taxa go extinct in the wild and even more slip into regional or local extinction over the span of only a few years. The hill mynas Gracula spp. are among its main victims, encompassing the Critically Endangered Nias Hill Myna Gracula [religiosa] robusta and other endangered populations across the West Sumatran Archipelago. Hill mynas are known to be present throughout this island chain but the taxonomic relationships of West Sumatran Gracula populations remain poorly understood. We hypothesized that the unique history of this island chain may have given rise to multiple distinct insular forms. Here we use genome‐wide DNA data in concert with morphological analyses to investigate the evolutionary distinctness of these taxa. Our results identify one taxon that is surprisingly distinct despite lacking recognition in most classifications, the ‘Simeulue Hill Myna’ (taxon miotera), and a range extension of the Nias Hill Myna. Despite their lack of recognition, Simeulue Hill Mynas are genomically and morphologically as unique as their Nias counterpart, in accordance with the lack of glacial land bridges between the island of Simeulue and mainland Sumatra. Simeulue Hill Mynas went extinct in the wild sometime within the last 2–3 years, and the rescue of the last captive individuals should now be the highest priority.     

Genetic structure of the critically endangered Red‐headed Wood Pigeon Columba janthina nitens and its implications for the management of threatened island populations

The Red‐headed Wood Pigeon Columba janthina nitens is endemic to the Ogasawara Islands, an oceanic island chain located 1000 km south of the main islands of Japan. The subspecies is at high risk of extinction because of its small population size and restricted habitat range. We undertook genetic analyses of this pigeon using sequences of a portion of the mitochondrial control region and five microsatellite markers to estimate the genetic characteristics of two wild populations from the Bonin and Volcano Islands, as well as one captive breeding population. The genetic diversity of the wild individuals was exceptionally low in both the mitochondria (nucleotide diversity = 0.00105) and at the microsatellite (3.2 alleles per locus and H_E = 0.12) loci. Higher numbers of microsatellite genotypes were observed in the Volcano Islands population than in the Bonin Islands population, which may be because of the relatively low impact of human disturbance. The most common mitochondrial haplotypes and microsatellite alleles observed in the two wild populations were completely fixed in the captive population. Our results suggest that the genetic diversity of the captive population needs to be increased. However, introduction of a wild individual into a captive population can lead to a decreased genetic diversity in the wild population and therefore should be done with caution. The genetic differentiation between the Bonin and the Volcano island groups was low, and the populations of the two island groups should be regarded as a single evolutionarily significant unit. However, special consideration is required for habitat conservation in the Volcano Islands, which may be functioning as a sanctuary for the Red‐headed Wood Pigeon. For the long‐term conservation of threatened bird species that live on remote oceanic islands, determination of management units considering gene flow caused by their flying capacity and maintenance of genetically suitable wild and captive populations are essential.     

A reconstruction of Palaeo‐Macaronesia,with particular reference to the long‐term biogeography of the Atlantic island laurel forests

Macaronesia is a biogeographical region comprising five Atlantic Oceanic archipelagos: the Azores, Madeira, Selvagen (Savage Islands), Canaries and Cape Verde. It has strong affinities with the Atlantic coast of the Iberian Peninsula and the north‐western fringes of Africa. This paper re‐evaluates the biogeographical history and relationships of Macaronesia in the light of geological evidence, which suggests that large and high islands may have been continuously available in the region for very much longer than is indicated by the maximum surface area of the oldest current island (27 Ma) – possibly for as long as 60 million years. We review this literature, attempting a sequential reconstruction of Palaeo‐Macaronesia from 60 Ma to the present. We consider the implications of these geological dynamics for our understanding of the history of colonization of the present islands of Macaronesia. We also evaluate the role of these archipelagos as stepping stones and as both repositories of palaeo‐endemic forms and crucibles of neo‐endemic radiations of plant and animal groups. Our principal focus is on the laurel forest communities, long considered impoverished relicts of the Palaeotropical Tethyan flora. This account is therefore contextualized by reference to the long‐term climatic and biogeographical history of Southern Europe and North Africa and by consideration of the implications of changes in land–sea configuration, climate and ocean circulation for Macaronesian biogeography. We go on to provide a synthesis of the more recent history of Macaronesian forests, which has involved a process of impoverishment of the native elements of the biota that has accelerated since human conquest of the islands. We comment briefly on these processes and on the contemporary status and varied conservation opportunities and threats facing these forests across the Macaronesian biogeographical region.     

Predation risk of eggs and nestlings relative to nest‐site characteristics of the Bull‐headed Shrike Lanius bucephalus

Appropriate nest‐site selection is one of the most important ways to minimize loss of reproductive investment due to predation. We determined the environmental characteristics associated with nest predation during the incubation and nestling periods of arboreal nesting Bull‐headed Shrikes on the oceanic Minami‐Daito Island where the predator community has low species diversity and includes only three introduced mammals: Ship Rat Rattus rattus, Japanese Weasel Mustela itatsi and Feral Cat Felis catus. Egg predation declined with increasing grassland cover around nests, whereas nestling predation declined with increasing nest concealment and nest height. Our results suggest that effective nest‐site characteristics for avoiding nest predation differ during the incubation and nestling periods and are dependent on the predator species and their search strategies, at least in habitats with low predator species diversity.     

Colonization of the Philippines from Taiwan: a multi‐locus test of the biogeographic and phylogenetic relationships of isolated populations of shrews

Aim Colonization of the Philippines from Taiwan or neighbouring areas of the Asian mainland has been proposed as an important source of diversity for some plant and animal groups in the northern Philippines. Previous inferences, however, were based on taxonomic groupings, which sometimes fail to reflect phylogenetic history. Here, we test for colonization of the Philippines from the north in a group of shrews (Soricomorpha: Crocidura) using explicit inferences of evolutionary history. Location Southeast Asia. Methods We estimate the phylogenetic relationships of populations of shrews from Batan and Sabtang islands in the northern Philippines using DNA sequences from two mitochondrial genes and three nuclear loci. We employ topology tests to evaluate the possible relationships of these shrews to species from throughout Southeast Asia. Results We find conclusive evidence that shrews from Batan and Sabtang are closely related to Crocidura tanakae from Taiwan and additional specimens from the Asian mainland. Bayesian and frequentist topology tests using alignments of individual loci strongly reject any notion that shrews from Batan and Sabtang are part of the main Philippine radiation of Crocidura, indicating that the northernmost Philippine islands were almost certainly colonized by shrews from Taiwan or mainland Asia. Main conclusions Our results provide the first compelling evidence for colonization of the Philippine archipelago by a terrestrial vertebrate via a northern route. Invasion of the northern Philippines by shrews, however, did not lead to further range expansion to more southerly parts of the Philippines. This study, combined with previous results, documents that Crocidura colonized the Philippines at least three times. However, only one of these invasions led to in situ speciation and ubiquity across the archipelago. Our findings are part of a growing body of literature suggesting that oceanic archipelagos are often colonized multiple times by groups of closely related species, and occasionally from multiple sources.