Towards a more general species–area relationship: diversity on all islands, great and small

Aim

To demonstrate a new and more general model of the species–area relationship that builds on traditional models, but includes the provision that richness may vary independently of island area on relatively small islands (the small island effect).

Location

We analysed species–area patterns for a broad diversity of insular biotas from aquatic and terrestrial archipelagoes.

Methods

We used breakpoint or piecewise regression methods by adding an additional term (the breakpoint transformation) to traditional species–area models. The resultant, more general, species–area model has three readily interpretable, biologically relevant parameters: (1) the upper limit of the small island effect (SIE), (2) an estimate of richness for relatively small islands and (3) the slope of the species–area relationship (in semi‐log or log–log space) for relatively large islands.

Results

The SIE, albeit of varying magnitude depending on the biotas in question, appeared to be a relatively common feature of the data sets we studied. The upper limit of the SIE tended to be highest for species groups with relatively high resource requirements and low dispersal abilities, and for biotas of more isolated archipelagoes.

Main conclusions

The breakpoint species–area model can be used to test for the significance, and to explore patterns of variation in small island effects, and to estimate slopes of the species–area (semi‐log or log–log) relationship after adjusting for SIE. Moreover, the breakpoint species–area model can be expanded to investigate three fundamentally different realms of the species–area relationship: (1) small islands where species richness varies independent of area, but with idiosyncratic differences among islands and with catastrophic events such as hurricanes, (2) islands beyond the upper limit of SIE where richness varies in a more deterministic and predictable manner with island area and associated, ecological factors and (3) islands large enough to provide the internal geographical isolation (large rivers, mountains and other barriers within islands) necessary for in situ speciation.

     

Novel summary metrics for insular biotic assemblages based on taxonomy and phylogeny: Biogeographical,palaeogeographical and possible conservational applications

Using the land‐bound vertebrates on the marine islands as model organisms, two metrics are presented that permit quantitative and succinct synopses of the ‘evolutionary maturity’ of the hosted faunal assemblages. In turn, these reflect the geo‐physical settings and geological developments of the substrates. The assemblage lineage‐taxonomy spectrum (ALTS) is based on the constituent lineages’ taxonomic distinctiveness and diversity. Individual lineages within assemblages can in most cases be assigned to one of six categories, LT1?LT6: LT1 is a non‐endemic taxon, whereas LT6 comprises multiple endemic genera from a family that arose elsewhere. If required, the scheme can be expanded: LT9 is an endemic order. The data can then be combined to provide an assemblage spectrum, for example, 00:08:38:30:08:15[ 13 ]. Here, the first six values denote the number of lineages assigned to each category expressed as percentages of the overall total, with the sum of the processed lineages listed as the seventh (in brackets and bold). The ALTS metric highlights efficiently the key features of a marine island's biological assemblage. Notably, the contrast between spectra for suites on geologically and geo‐physically varied island types can be striking, for instance the squamate suite on the young, proximate orogenic margin island of Taiwan is coded 78:16:05:00:00:00[ 37 ] whereas the one on the distantly located, Late Eocene composite terrane island of New Caledonia is 00:11:00:11:33:44[ 9 ]. To overcome the subjectivity that is inherent in assigning supraspecific ranks, an alternative assemblage lineage‐age spectrum (ALAS) is also introduced that makes use of the binary logarithm values of the colonization times of the island lineages (0–2, 2–4, … , 32–64, >64 Ma). It is represented using a seven‐plus‐two‐number code, for instance Madagascar's squamates are 00:06:00:00:19:62:12[ 19 ( 16 )]; most colonizations took place in the Palaeogene (66–23 Ma); there are 19 lineages, but only 16 are presently age‐dated. In addition to marine‐island biogeography studies, the ALTS–ALAS spectrum approach is potentially useful for encapsulating biotas in other sorts of insular setting (e.g. lakes, mountain tops), and for evaluating palaeogeographical models. Furthermore, it may help emphasize the conservation value of an island's faunal assemblage.     

L'origine géologique des iles des Indes Occidentales en relation avec la dispersion de quelques Malacostracés stygobiontes

The relation between the geological theories andthe origin of the biotas of the West Indian islands is analyzed, mainly based upon observations on the stygobiont fauna of an important number of islands.Two models for the geological origin of theislands have been postulated, viz. an active model (ab novo formation by submarine volcanic action) and a passive model (fragmentation and drift of continental material originating from the protocentral-american region).The same models are encountered in biogeographic considerations on the origin of the biotas of the archipelago. According to the active model (or dispersion model), the biotas arrived from nearby terrestrial regions through dispersion. No doubt this route is, and has been, very important in the West Indies.The passive model (or vicariance model) supposes a passive transport of the biotas «on the back of floating continental fragments. Certain arguments in favour of this model are weighed by the author and are not considered parsimonious. at least for the islands in question.A modification of the passive model is the evolution according to the regission model («stranding of populations of marine origin during periods of marine regression or tectonic uplift); this phenomenon has played an extremely important role in the evolution of limnic biotas, chiefly stygobiont ones, of the Antilles, an island chain that is in a quasi-permanent stage of uplifting.The Bahamas, in the contrary, are in a continuoussubsidence stage. Their limnic faunas do not comprise regression elements, but show clear traces of invasions, probably fairly recent ones, by marine animals.     

Of mice and mammoths: generality and antiquity of the island rule

Aim

We assessed the generality of the island rule in a database comprising 1593 populations of insular mammals (439 species, including 63 species of fossil mammals), and tested whether observed patterns differed among taxonomic and functional groups.

Location

Islands world‐wide.

Methods

We measured museum specimens (fossil mammals) and reviewed the literature to compile a database of insular animal body size (S_i = mean mass of individuals from an insular population divided by that of individuals from an ancestral or mainland population, M). We used linear regressions to investigate the relationship between S_i and M, and ANCOVA to compare trends among taxonomic and functional groups.

Results

S_i was significantly and negatively related to the mass of the ancestral or mainland population across all mammals and within all orders of extant mammals analysed, and across palaeo‐insular (considered separately) mammals as well. Insular body size was significantly smaller for bats and insectivores than for the other orders studied here, but significantly larger for mammals that utilized aquatic prey than for those restricted to terrestrial prey.

Main conclusions

The island rule appears to be a pervasive pattern, exhibited by mammals from a broad range of orders, functional groups and time periods. There remains, however, much scatter about the general trend; this residual variation may be highly informative as it appears consistent with differences among species, islands and environmental characteristics hypothesized to influence body size evolution in general. The more pronounced gigantism and dwarfism of palaeo‐insular mammals, in particular, is consistent with a hypothesis that emphasizes the importance of ecological interactions (time in isolation from mammalian predators and competitors was 0.1 to > 1.0 Myr for palaeo‐insular mammals, but < 0.01 Myr for extant populations of insular mammals). While ecological displacement may be a major force driving diversification in body size in high‐diversity biotas, ecological release in species‐poor biotas often results in the convergence of insular mammals on the size of intermediate but absent species.     

‘On being the right size’ – Do aliens follow the rules?

Aim

To assess whether mammalian species introduced onto islands across the globe have evolved to exhibit body size patterns consistent with the ‘island rule,’, and to test an ecological explanation for body size evolution of insular mammals.

Location

Islands worldwide.

Methods

We assembled data on body mass, geographical characteristics (latitude, maximum elevation) and ecological communities (number of mammalian competitors, predators and prey) for 385 introduced populations across 285 islands, comprising 56 species of extant, non‐volant mammals. We used linear regression, ANCOVA and regression tree analyses to test whether introduced populations of mammals exhibit the island rule pattern, whether the degree of body size change increased with time in isolation and whether residual variation about the general trend can be attributed to the geographical and ecological characteristics of the islands.

Results

Introduced populations follow the predicted island rule trend, with body size shifts more pronounced for populations with greater residence times on the islands. Small mammals evolved to larger body sizes in lower latitudes and on islands with limited topographic relief. Consistent with our hypothesis on the ecology of evolution, body size of insular introduced populations was influenced by co‐occurring species of mammalian competitors, predators and prey.

Conclusion

The island rule is a pervasive pattern, exhibited across a broad span of geographical regions, taxa, time periods and, as evidenced here, for introduced as well as native mammals. Time in isolation impacts body size evolution profoundly. Body size shift of introduced mammals was much more pronounced with increasing residence times, yet far less than that exhibited by native, palaeo‐insular mammals (residence times > 10,000 years). Given the antiquity of many species introductions, it appears that much of what we view as the natural character and ecological dynamics of recent insular communities may have been rendered artefacts of ancient colonizations by humans and commensals.     

Insularity and the evolution of melanism,sexual dichromatism and body size in the worldwide‐distributed barn owl

Island biogeography has provided fundamental hypotheses in population genetics, ecology and evolutionary biology. Insular populations usually face different feeding conditions, predation pressure, intraspecific and interspecific competition than continental populations. This so‐called island syndrome can promote the evolution of specific phenotypes like a small (or large) body size and a light (or dark) colouration as well as influence the evolution of sexual dimorphism. To examine whether insularity leads to phenotypic differentiation in a consistent way in a worldwide‐distributed nonmigratory species, we compared body size, body shape and colouration between insular and continental barn owl (Tyto alba) populations by controlling indirectly for phylogeny. This species is suitable because it varies in pheomelanin‐based colouration from reddish‐brown to white, and it displays eumelanic black spots for which the number and size vary between individuals, populations and species. Females are on average darker pheomelanic and display more and larger eumelanic spots than males. Our results show that on islands barn owls exhibited smaller and fewer eumelanic spots and lighter pheomelanic colouration, and shorter wings than on continents. Sexual dimorphism in pheomelanin‐based colouration was less pronounced on islands than continents (i.e. on islands males tended to be as pheomelanic as females), and on small islands owls were redder pheomelanic and smaller in size than owls living on larger islands. Sexual dimorphism in the size of eumelanic spots was more pronounced (i.e. females displayed much larger spots than males) in barn owls living on islands located further away from a continent. Our study indicates that insular conditions drive the evolution towards a lower degree of eumelanism, smaller body size and affects the evolution of sexual dichromatism in melanin‐based colour traits. The effect of insularity was more pronounced on body size and shape than on melanic traits.     

Narrow water barriers prevent multiple colonizations and limit gene flow among California Channel Island wild buckwheats (Eriogonum: Polygonaceae)

The relative roles of chance colonization and subsequent gene flow in the development of insular endemic biotas have been extensively studied in remote oceanic archipelagos, but are less well characterized on nearshore island systems. The current study investigated patterns of colonization and divergence between and within two wild buckwheat species (Polygonaceae), Eriogonum arborescens and E. giganteum, endemic to the California Channel Islands to determine whether geographical isolation is driving diversification. Using plastid and nuclear sequence data and microsatellite allele frequencies, we determined that gene flow in these Eriogonum spp. is restricted by isolation. The data suggest that successful colonization of and gene flow among the islands are infrequent. Colonization appears to have followed a stepping‐stone model that is consistent with a north‐to‐south pattern across the islands. This colonization pattern coupled with relatively little post‐colonization inter‐island gene flow, particularly among southern islands, has generated a pattern of more divergent lineages on the isolated southern islands. These results run counter to the general expectation that all islands close to a continental source should receive a high level of gene flow. Finally, management recommendations focused on protecting the lineages from loss of private alleles and the erosion of the remaining genetic diversity are offered.     

Vicariant versus dispersal processes in the settlement of Pseudamnicola (Caenogastropoda,Hydrobiidae) in the Mediterranean Balearic Islands

The genus Pseudamnicola comprises a group of tiny springsnails inhabiting several continental and insular regions of the Mediterranean basin. Given the limited dispersal capabilities of these animals, it is difficult to explain the wide distribution range of the genus and, more specifically, its presence in isolated habitats, such as on islands. Thus, to investigate the process(es) that may explain these distribution patterns, we morphologically re‐described and genetically analysed the six Pseudamnicola (Pseudamnicola) species occurring in the Iberian Peninsula and the nearby Balearic Islands. Genetic relationships were explored by sequencing two mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) and one nuclear (28S rRNA) gene in 19 populations. Our morphological study confirmed the presence of previously described species, whereas our phylogenetic results revealed three lineages within the subgenus: one clade grouping the species from Minorca Island with an Iberian Peninsula species, a second clade grouping the three species from Majorca Island, and a third clade that consists of a single species, which occurs in both the Iberian Peninsula and Ibiza Island. Calculated speciation times show that the cladogenetic events involving the insular species seem to have occurred after the current conformation of the Balearic Islands (c. 20 Mya). Therefore, the speciation process may have been related to subsequent transmarine colonizations, probably during the Messinian Salinity Crisis, and the Pleistocene glaciations when landmass corridors connected the islands with the continent. © 2014 The Linnean Society of London     

Diversification in the tropical pacific: comparisons between marine and terrestrial systems and the importance of founder speciation

Patterns of distribution and processes of differentiation haveoften been contrasted between terrestrial and marine biotas.The islands of Oceania offer an excellent setting to explorethis contrast, because the geographic setting for terrestrialand shallow-water, benthic, marine organisms are the same: themyriad islands strewn across the vast Pacific. The size of speciesranges and the geographic distribution of endemism are two biogeographicattributes that are thought to differ markedly between terrestrialand marine biotas in the Pacific. While terrestrial speciesare frequently confined to single islands or archipelagoes throughoutOceania, marine species tend to have wide to very wide distributions,and are rarely restricted to single island groups except forthe most isolated archipelagoes. We explore the conditions underwhich species can reach an island by dispersal and differentiate.Genetic differentiation can occur either through founder speciationor vicariance; these processes are requisite ends of a continuum.We show that founder speciation is most likely when few propagulesenter the dispersal medium and survive well while they travelfar. We argue that conditions favorable to founder speciationare common in marine as well as terrestrial systems, and thatterrestrial-type, archipelagic-level endemism is likely commonin marine taxa. We give examples of marine groups that showarchipelagic level endemism on most Pacific island groups aswell as of terrestrial species that are widespread. Thus boththe patterns and processes of insular diversification are variable,and overlap more between land and sea than previously considered.     

Why birds with deferred sexual maturity are sedentary on islands: a systematic review

Background

Island faunas have played central roles in the development of evolutionary biology and ecology. Birds are among the most studied organisms on islands, in part because of their dispersal powers linked to migration. Even so, we lack of information about differences in the movement ecology of island versus mainland populations of birds.

Methodology/Principal Findings

Here we present a new general pattern indicating that large birds with deferred sexual maturity are sedentary on islands, and that they become so even when they are migratory on the mainland. Density-dependent variation in the age at first breeding affects the survivorship of insular populations and this, in turn, affects the movement ecology of large birds. Because density-dependent variation in the age of first breeding is critical to the long-term survival of small isolated populations of long-lived species, migratory forms can successfully colonize islands only if they become sedentary once there. Analyses of the movement ecology of continental and insular populations of 314 species of raptors, 113 species of Ciconiiformes and 136 species of passerines, along with individual-based population simulations confirm this prediction.

Conclusions

This finding has several consequences for speciation, colonization and survival of small isolated population of species with deferred sexual maturity.     

If hippopotamuses cannot swim,how did they colonize islands?

Owing to their aquatic lifestyle, hippopotamuses are normally believed to have reached islands by swimming. Yet, some studies suggest they cannot swim due to their relatively high density. If so, this raises the question of how hippopotamuses would have reached some islands. Their immigration into the British Isles, Sicily, Malta, Zanzibar and Mafia can be accounted for, because these islands sit on continental shelves and were often linked to the mainland during the Pleistocene glacio‐eustatic sea‐level falls. In contrast, their occurrence in Crete, Cyprus and Madagascar would be more difficult to explain. Available geological evidence does not seem to rule out that the latter islands might have been connected with the nearest mainland areas in very recent times. This study intends to consider possibilities about how hippopotamuses reached islands and to show that more effective collaboration is required among specialists involved with the study of insular evolution, colonization and speciation.     

The extent of population genetic subdivision differs among four co-distributed shark species in the Indo-Australian archipelago

Background  

The territorial fishing zones of Australia and Indonesia are contiguous to the north of Australia in the Timor and Arafura Seas and in the Indian Ocean to the north of Christmas Island. The area surrounding the shared boundary consists of a variety of bio-diverse marine habitats including shallow continental shelf waters, oceanic trenches and numerous offshore islands. Both countries exploit a variety of fisheries species, including whaler (Carcharhinus spp.) and hammerhead sharks (Sphyrna spp.). Despite their differences in social and financial arrangements, the two countries are motivated to develop complementary co-management practices to achieve resource sustainability. An essential starting point is knowledge of the degree of population subdivision, and hence fisheries stock status, in exploited species.     

Disentangling ancient interactions: a new extinct passerine provides insights on character displacement among extinct and extant island finches

Background

Evolutionary studies of insular biotas are based mainly on extant taxa, although such biotas represent artificial subsets of original faunas because of human-caused extinctions of indigenous species augmented by introduced exotic taxa. This makes it difficult to obtain a full understanding of the history of ecological interactions between extant sympatric species. Morphological bill variation of Fringilla coelebs and F. teydea (common and blue chaffinches) has been previously studied in the North Atlantic Macaronesian archipelagos. Character displacement between both species has been argued to explain bill sizes in sympatry. However, this explanation is incomplete, as similar patterns of bill size have been recorded in F. coelebs populations from islands with and without F. teydea.

Methodology/Principal Findings

The discovery of a new extinct species in Tenerife (Canary Islands), here named Carduelis aurelioi n. sp. (slender-billed greenfinch), provides the opportunity to study ancient ecological interactions among Macaronesian finches. To help understand the evolutionary histories of forest granivores in space and time, we have performed a multidisciplinary study combining: (1) morphological analyses and radiocarbon dating (11,460±60 yr BP) of the new taxon and, (2) molecular divergence among the extant finch species and populations in order to infer colonization times (1.99 and 1.09 My for F. teydea and F. coelebs respectively).

Conclusion/Significance

C. aurelioi, F. coelebs and F. teydea co-habited in Tenerife for at least one million years. The unique anatomical trends of the new species, namely chaffinch-like beak and modified hind and forelimbs, reveal that there was a process of divergence of resource competition traits among the three sympatric finches. The results of our study, combined with the presence of more extinct greenfinches in other Macaronesian islands with significant variation in their beak sizes, suggests that the character displacement has influenced patterns of divergence in bill size and shape on other Macaronesian islands as well.     

Conservation biogeography of the Antarctic

Aim

To present a synthesis of past biogeographic analyses and a new approach based on spatially explicit biodiversity information for the Antarctic region to identify biologically distinct areas in need of representation in a protected area network.

Location

Antarctica and the sub‐Antarctic.

Methods

We reviewed and summarized published biogeographic studies of the Antarctic. We then developed a biogeographic classification for terrestrial conservation planning in Antarctica by combining the most comprehensive source of Antarctic biodiversity data available with three spatial frameworks: (1) a 200‐km grid, (2) a set of areas based on physical parameters known as the environmental domains of Antarctica and (3) expert‐defined bioregions. We used these frameworks, or combinations thereof, together with multivariate techniques to identify biologically distinct areas.

Results

Early studies of continental Antarctica typically described broad bioregions, with the Antarctic Peninsula usually identified as biologically distinct from continental Antarctica; later studies suggested a more complex biogeography. Increasing complexity also characterizes the sub‐Antarctic and marine realms, with differences among studies often attributable to the focal taxa. Using the most comprehensive terrestrial data available and by combining the groups formed by the environmental domains and expert‐defined bioregions, we were able to identify 15 biologically distinct, ice‐free, Antarctic Conservation Biogeographic Regions (ACBRs), encompassing the continent and close lying islands.

Main conclusions

Ice‐free terrestrial Antarctica comprises several distinct bioregions that are not fully represented in the current Antarctic Specially Protected Area network. Biosecurity measures between these ACBRs should also be developed to prevent biotic homogenization in the region.     

Origins of endemic island tortoises in the western Indian Ocean: a critique of the human‐translocation hypothesis

How do organisms arrive on isolated islands, and how do insular evolutionary radiations arise? In a recent paper, Wilmé et al. ( 2016a ) argue that early Austronesians that colonized Madagascar from Southeast Asia translocated giant tortoises to islands in the western Indian Ocean. In the Mascarene Islands, moreover, the human‐translocated tortoises then evolved and radiated in an endemic genus (Cylindraspis). Their proposal ignores the broad, established understanding of the processes leading to the formation of native island biotas, including endemic radiations. We find Wilmé et al.'s suggestion poorly conceived, using a flawed methodology and missing two critical pieces of information: the timing and the specifics of proposed translocations. In response, we here summarize the arguments that could be used to defend the natural origin not only of Indian Ocean giant tortoises but also of scores of insular endemic radiations world‐wide. Reinforcing a generalist's objection, the phylogenetic and ecological data on giant tortoises, and current knowledge of environmental and palaeogeographical history of the Indian Ocean, make Wilmé et al.'s argument even more unlikely.     

Pliocene climatic change in insular Southeast Asia as an engine of diversification in Ficedula flycatchers

Aims Insular Southeast Asia and adjacent regions are geographically complex, and were dramatically affected by both Pliocene and Pleistocene changes in climate, sea level and geology. These circumstances allow the testing of several biogeographical hypotheses regarding species distribution patterns and phylogeny. Avian species in this area present a challenge to biogeographers, as many are less hindered by barriers that may block the movements of other species. Widely distributed Southeast Asian avian lineages, of which there are many, have been generally neglected. Ficedula flycatchers are distributed across Eurasia, but are most diverse within southern Asia and Southeast Asian and Indo‐Australian islands. We tested the roles of vicariance, dispersal and the evolution of migratory behaviours as mechanisms of speciation within the Ficedula flycatchers, with a focus on species distributed in insular Southeast Asia. Methods Using a published molecular phylogeny of Ficedula flycatchers, we reconstructed ancestral geographical areas using dispersal vicariance analysis, weighted ancestral area analysis, and a maximum likelihood method. We evaluated the evolution of migratory behaviours using maximum likelihood ancestral character state reconstruction. Speciation timing estimates were calculated via local molecular clock methods. Results Ficedula originated in southern mainland Asia, c. 6.5 Ma. Our analyses indicate that two lineages within Ficedula independently and contemporaneously colonized insular Southeast Asia and Indo‐Australia, c. 5 Ma. The potential impact of vicariance due to rising sea levels is difficult to assess in these early colonization events because the ancestral areas to these clades are reconstructed as oceanic islands. Within each of these clades, inter‐island dispersal was critical to species’ diversification across oceanic and continental islands. Furthermore, Pliocene and Pleistocene climatic change may have caused the disjunct island distributions between several pairs of sister taxa. Both vicariance and dispersal shaped the distributions of continental species. Main conclusions This study presents the first evaluation, for Ficedula, of the importance of vicariance and dispersal in shaping distributions, particularly across insular Southeast Asia and Indo‐Australia. Although vicariant speciation may have initially separated the island clades from mainland ancestors, speciation within these clades was driven primarily by dispersal. Our results contribute to the emerging body of literature concluding that dynamic geological processes and climatic change throughout the Pliocene and Pleistocene have been important factors in faunal diversification across continental and oceanic islands.     

Body Size Evolution in Insular Speckled Rattlesnakes (Viperidae: Crotalus mitchellii)

Background

Speckled rattlesnakes (Crotalus mitchellii) inhabit multiple islands off the coast of Baja California, Mexico. Two of the 14 known insular populations have been recognized as subspecies based primarily on body size divergence from putative mainland ancestral populations; however, a survey of body size variation from other islands occupied by these snakes has not been previously reported. We examined body size variation between island and mainland speckled rattlesnakes, and the relationship between body size and various island physical variables among 12 island populations. We also examined relative head size among giant, dwarfed, and mainland speckled rattlesnakes to determine whether allometric differences conformed to predictions of gape size (and indirectly body size) evolving in response to shifts in prey size.

Methodology/Principal Findings

Insular speckled rattlesnakes show considerable variation in body size when compared to mainland source subspecies. In addition to previously known instances of gigantism on Ángel de la Guarda and dwarfism on El Muerto, various degrees of body size decrease have occurred frequently in this taxon, with dwarfed rattlesnakes occurring mostly on small, recently isolated, land-bridge islands. Regression models using the Akaike information criterion (AIC) showed that mean SVL of insular populations was most strongly correlated with island area, suggesting the influence of selection for different body size optima for islands of different size. Allometric differences in head size of giant and dwarf rattlesnakes revealed patterns consistent with shifts to larger and smaller prey, respectively.

Conclusions/Significance

Our data provide the first example of a clear relationship between body size and island area in a squamate reptile species; among vertebrates this pattern has been previously documented in few insular mammals. This finding suggests that selection for body size is influenced by changes in community dynamics that are related to graded differences in area over what are otherwise similar bioclimatic conditions. We hypothesize that in this system shifts to larger prey, episodic saturation and depression of primary prey density, and predator release may have led to insular gigantism, and that shifts to smaller prey and increased reproductive efficiency in the presence of intense intraspecific competition may have led to insular dwarfism.     

Two decades of interaction between the MacArthur-Wilson model and the complexities of mammalian distributions

More than two decades after its publication, MacArthur and Wilson's equilibrium model of insular biogeography continues to provide the conceptual foundation for investigating the distribution of species on islands and the composition of insular biotas. During this period, studies of the distributions of mammals among insular habitats have tested, modified, and extended MacArthur and Wilson's simple formalism to enhance greatly our understanding of the complexities of biogeographic patterns and processes. The papers in this symposium summarize many of the past contributions of mammalian biogeographers and introduce important new data and ideas. The diversity of biological characteristics and associated distributional patterns exhibited by mammals has facilitated this endeavour. Some insular mammalian faunas appear to represent approximate equilibria between opposing rates of contemporary colonization and extinction. Other faunas are currently decreasing in diversity because of extinctions, owing either to natural habitat fragmentation that has occurred since the Pleistocene or to human activities within the last few centuries. Still other faunas have been increasing in diversity (at least until recent human impacts) because limiting rates of origination, both colonization and speciation, have been extremely low. The questions and analyses of island biogeography can also be applied to continents with comparable overall results: the distributions of continental faunas reflect the consequences of similar processes of colonization, speciation and extinction. Analyses of insular distributions show unequivocally that probabilities of extinction, colonization and speciation are highly deterministic and vary in predictable ways among different taxa and archipelagos. These findings have important implications for applying the theory and data of insular biogeography to the pressing practical problems of designing natural reserves to preserve native species.     

Do freshwater ecoregions and continental shelf width predict patterns of historical gene flow in the freshwater fish Poecilia butleri?

We examined historical patterns of gene flow in the freshwater fish Poecilia butleri in western Mexico. We tested the hypothesis that the boundaries between four freshwater ecological communities (ecoregions) might have limited the movement of P. butleri because changes in species compositions might restrict establishment between adjacent ecoregions, even in situations where a physical barrier is absent. Hence, we predicted that boundaries between ecoregions should correspond to phylogeographical breaks in P. butleri. We also tested the hypothesis that the width of the continental shelf affected historical gene flow in P. butleri because a broad continental shelf provides a greater opportunity for rivers to coalesce during historical episodes of low sea levels as opposed to a narrow continental shelf that should restrict the potential for gene flow among adjacent rivers. Hence, we predicted greater amounts of historical gene flow among neighbouring river basins in the region of western Mexico where the continental shelf is wider, whereas, in the region where the continental shelf is narrower, we expected to detect limited levels of historical gene flow. We analyzed mitochondrial DNA sequence data (cytochrome b) taken from 264 individuals of P. butleri collected from 34 locations distributed across four different ecoregions in western Mexico. To examine patterns of phylogenetic diversification and historical gene flow in P. butleri, we employed several analytical approaches, including traditional tree‐based phylogenetic analyses (likelihood and parsimony), haplotype network reconstruction, analyses of molecular variance, and spatial analysis of molecular variance. We found genetic breaks coinciding with two out of three different ecoregion boundaries, suggesting limited historical gene flow. In addition to different species compositions between these adjacent ecoregions, geological features such as the Trans‐Mexican Volcanic Belt and the mountainous topography in south‐western Mexico, likely contributed to these observed genetic breaks. By contrast, no genetic break was evident between two other ecoregions, a result that partially rejects our first hypothesis. Several results were consistent with our second hypothesis. Changes in the width of the continental shelf in western Mexico are associated with the observed patterns of historical gene flow. Our results indicate that the interactions among multiple geological and biological factors affect the spatial patterns of genetic diversity of widespread freshwater species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 399–416.     

Complex inter‐island colonization and peripatric founder speciation promote diversification of flightless Pachyrhynchus weevils in the Taiwan–Luzon volcanic belt

Aim

We investigated the spatial and temporal patterns of diversification among colourful and flightless weevils, the Pachyrhynchus orbifer complex, to test the stepping‐stone hypothesis of colonization across the Taiwan–Luzon volcanic belt.

Location

Southeast Asia.

Methods

The phylogeny of the P. orbifer complex was reconstructed from a multi‐locus data set of mitochondrial and nuclear genes using maximum likelihood in RAxML and Bayesian inference in MRBAYES. Likelihood‐based tests in CONSEL were used to evaluate alternative tree topologies. Divergence times were estimated in beast based on a range of mutation rates. Ancestral range and biogeographical history were reconstructed using Bayesian binary MCMC (BBM) methods in RASP and in BioGeoBEARS. Demographic histories were inferred using the extended Bayesian skyline plot (EBSP). Species boundaries were tested using BPP.

Results

The phylogeny of the P. orbifer complex indicated strong support for seven reciprocally monophyletic lineages grouped by current island boundaries (Camiguin, Fuga, Dalupiri, Calayan, Babuyan, Orchid and Yaeyama Islands), except for a sister Green + Itbayat lineage. Complex and stochastic colonization of P. orbifer was inferred to have involved both northward and southward directions with short‐ and long‐distance dispersal events, which are strongly inconsistent with the strict stepping‐stone hypothesis. Divergence time estimates for all extant island lineages (<1 Myr of Middle Pleistocene) are much more recent than the geological ages (22.4–1.7 Myr) and subaerial existence (c. 3 Myr) of the islands. The statistically delimited seven cryptic species imply that the diversity of Pachyrhynchus from small peripheral islands continues to be largely under‐estimated.

Main conclusions

The non‐linear, more complex spatial and temporal settings of the archipelago and stochastic dispersal were probable key factors shaping the colonization history of the P. orbifer complex. Speciation of the P. orbifer complex may have occurred only between islands, indicating that peripatric speciation through the founders of stochastic dispersals was the major evolutionary driver.