Mainland Relatives Research Articles

Speciation in Howea Palms Occurred in Sympatry, Was Preceded by Ancestral Admixture, and Was Associated with Edaphic and Phenological Adaptation.

Howea palms are viewed as one of the most clear-cut cases of speciation in sympatry. The sister species Howea belmoreana and H. forsteriana are endemic to the oceanic Lord Howe Island, Australia, where they have overlapping distributions and are reproductively isolated mainly by flowering time differences. However, the potential role of introgression from Australian mainland relatives had not previously been investigated, a process that has recently put other examples of sympatric speciation into question. Furthermore, the drivers of flowering time-based reproductive isolation remain unclear. We sequenced an RNA-seq data set that comprehensively sampled Howea and their closest mainland relatives (Linospadix, Laccospadix), and collected detailed soil chemistry data on Lord Howe Island to evaluate whether secondary gene flow had taken place and to examine the role of soil preference in speciation. D-statistics analyses strongly support a scenario whereby ancestral Howea hybridized frequently with its mainland relatives, but this only occurred prior to speciation. Expression analysis, population genetic and phylogenetic tests of selection, identified several flowering time genes with evidence of adaptive divergence between the Howea species. We found expression plasticity in flowering time genes in response to soil chemistry as well as adaptive expression and sequence divergence in genes pleiotropically linked to soil adaptation and flowering time. Ancestral hybridization may have provided the genetic diversity that promoted their subsequent adaptive divergence and speciation, a process that may be common for rapid ecological speciation.

Brain Changes during Phyletic Dwarfing in Elephants and Hippos

Of all known insular mammals, hippos and elephants present the extremes of body size decrease, reducing to 4 and a mere 2% of their ancestral mainland size, respectively. Despite the numerous studies on these taxa, what happens to their relative brain size during phyletic dwarfing is not well known, and results are sometimes conflicting. For example, relative brain size increase has been noted in the Sicilian dwarf elephant, Palaeoloxodon falconeri, whereas relative brain size decrease has been postulated for Malagasy dwarf hippos. Here, I perform an analysis of brain, skull, and body size of 3 insular elephants (Palaeoloxodon “mnaidriensis,” P. tiliensis, and P. falconeri) and 3 insular hippos (Hippopotamus madagascariensis, H. lemerlei, and H. minor) to address this issue and to test whether relative brain size in phyletic dwarf species can be predicted. The results presented here show that the encephalization of all insular elephants and hippos is higher than that of their continental relatives. P. falconeri in particular has an enormous encephalization increase, which has so far not been reported in any other insular mammal. Insular brain size cannot be reliably predicted using either static allometric or ontogenetic scaling models. The results of this study indicate that insular dwarf species follow brain-body allometric relationships different from the expected patterns seen for their mainland relatives.

Reduced Defenses and Increased Herbivore Preference of Island Chaparral Shrubs Compared to Mainland Relatives

The absence of herbivores from certain environments may select for plants that are less defended and more palatable compared to plants in similar environments that are exposed to herbivory. This potential loss of defenses becomes especially important when exotic herbivores are introduced to islands that historically lack such organisms, potentially resulting in negative effects on native island ecosystems. To test the hypothesis that island plants have reduced defensive traits, we measured structural defenses and palatability of 10 taxonomic pairs from contrasting island and mainland environments. Our study compares plants from Santa Catalina Island, California—which has a long history without large native herbivores followed by more recent human-caused introductions—to the adjacent southern California mainland where ungulate herbivores are native and common. Our results suggest that island plants have reduced structural defenses and are more preferred by herbivores compared to their mainland plant relatives. These patterns are likely driven by selection on plant traits in the unique and insular island environment. Reduced defenses and increased palatability of island plant species should be considered when developing management strategies for island ecosystems that historically lack native ungulate herbivores.

Predictable evolution towards larger brains in birds colonizing oceanic islands

Theory and evidence suggest that some selective pressures are more common on islands than in adjacent mainland habitats, leading evolution to follow predictable trends. The existence of predictable evolutionary trends has nonetheless been difficult to demonstrate, mainly because of the challenge of separating in situ evolution from sorting processes derived from colonization events. Here we use brain size measurements of >1900 avian species to reveal the existence of one such trend: increased brain size in island dwellers. Based on sister-taxa comparisons and phylogenetic ancestral trait estimations, we show that species living on islands have relatively larger brains than their mainland relatives and that these differences mainly reflect in situ evolution rather than varying colonization success. Our findings reinforce the view that in some instances evolution may be predictable, and yield insight into why some animals evolve larger brains despite substantial energetic and developmental costs.

Convergence in reduced body size, head size, and blood glucose in three island reptiles.

Many oceanic islands harbor diverse species that differ markedly from their mainland relatives with respect to morphology, behavior, and physiology. A particularly common morphological change exhibited by a wide range of species on islands worldwide involves either a reduction in body size, termed island dwarfism, or an increase in body size, termed island gigantism. While numerous instances of dwarfism and gigantism have been well documented, documentation of other morphological changes on islands remains limited. Furthermore, we lack a basic understanding of the physiological mechanisms that underlie these changes, and whether they are convergent. A major hypothesis for the repeated evolution of dwarfism posits selection for smaller, more efficient body sizes in the context of low resource availability. Under this hypothesis, we would expect the physiological mechanisms known to be downregulated in model organisms exhibiting small body sizes due to dietary restriction or artificial selection would also be downregulated in wild species exhibiting dwarfism on islands. We measured body size, relative head size, and circulating blood glucose in three species of reptiles—two snakes and one lizard—in the California Channel Islands relative to mainland populations. Collating data from 6 years of study, we found that relative to mainland population the island populations had smaller body size (i.e., island dwarfism), smaller head sizes relative to body size, and lower levels of blood glucose, although with some variation by sex and year. These findings suggest that the island populations of these three species have independently evolved convergent physiological changes (lower glucose set point) corresponding to convergent changes in morphology that are consistent with a scenario of reduced resource availability and/or changes in prey size on the islands. This provides a powerful system to further investigate ecological, physiological, and genetic variables to elucidate the mechanisms underlying convergent changes in life history on islands.

Vomeronasal and Olfactory Structures in Bats Revealed by DiceCT Clarify Genetic Evidence of Function.

The degree to which molecular and morphological loss of function occurs synchronously during the vestigialization of traits is not well understood. The mammalian vomeronasal system, a sense critical for mediating many social and reproductive behaviors, is highly conserved across mammals. New World Leaf-nosed bats (Phyllostomidae) are under strong selection to maintain a functional vomeronasal system such that most phyllostomids possess a distinct vomeronasal organ and an intact TRPC2, a gene encoding a protein primarily involved in vomeronasal sensory neuron signal transduction. Recent genetic evidence, however, shows that TRPC2 is a pseudogene in some Caribbean nectarivorous phyllostomids. The loss-of-function mutations suggest the sensory neural tissue of the vomeronasal organ is absent in these species despite strong selection on this gene in its mainland relatives, but the anatomy was unknown in most Caribbean nectarivorous phyllostomids until this study. We used diffusible iodine-based contrast-enhanced computed tomography (diceCT) to test whether the vomeronasal and main olfactory anatomy of several phyllostomid species matched genetic evidence of function, providing insight into whether loss of a structure is linked to pseudogenization of a molecular component of the system. The vomeronasal organ is indeed rudimentary or absent in species with a disrupted TRPC2 gene. Caribbean nectar-feeders also exhibit derived olfactory turbinal morphology and a large olfactory recess that differs from closely related bats that have an intact vomeronasal organ, which may hint that the main olfactory system may compensate for loss. We emphasize non-invasive diceCT is capable of detecting the vomeronasal organ, providing a feasible approach for quantifying mammalian chemosensory anatomy across species.

Dome-headed, small-brained island mammal from the Late Cretaceous of Romania

The island effect is a well-known evolutionary phenomenon, in which island-dwelling species isolated in a resource-limited environment often modify their size, anatomy, and behaviors compared with mainland relatives. This has been well documented in modern and Cenozoic mammals, but it remains unclear whether older, more primitive Mesozoic mammals responded in similar ways to island habitats. We describe a reasonably complete and well-preserved skeleton of a kogaionid, an enigmatic radiation of Cretaceous island-dwelling multituberculate mammals previously represented by fragmentary fossils. This skeleton, from the latest Cretaceous of Romania, belongs to a previously unreported genus and species that possesses several aberrant features, including an autapomorphically domed skull and one of the smallest brains relative to body size of any advanced mammaliaform, which nonetheless retains enlarged olfactory bulbs and paraflocculi for sensory processing. Drawing on parallels with more recent island mammals, we interpret these unusual neurosensory features as related to the island effect. This indicates that the ability to adapt to insular environments developed early in mammalian history, before the advent of therian mammals, and mammals with insular-related modifications were key components of well-known dwarfed dinosaur faunas. Furthermore, the specimen suggests that brain size reduction, in association with heightened sensory acuity but without marked body size change, is a novel expression of the island effect in mammals.

Wildlife-vehicle collisions in Lanzarote Biosphere Reserve, Canary Islands.

Insular wildlife is more prone to extinction than their mainland relatives. Thus, a basic understanding of non-natural mortality sources is the first step in the development of conservation management plans. The Canary Islands are an important tourist destination due to their unique climate and rich scenery and biodiversity. During the last few decades, there has been significant development of urban areas and busy road networks. However, there have been no studies describing the effects of road mortality on wildlife in this archipelago. We describe the temporal and spatial patterns of wildlife roadkill in Lanzarote (UNESCO Biosphere Reserve), using counts from cars for an entire annual cycle. A total of 666 roadkills were recorded (monthly average of 0.09 birds/km and 0.14 mammals/km) comprising at least 37 species including native birds and introduced mammals. Seasonal abundance, richness and diversity of roadkills showed a high peak during summer months for both mammals and birds. GLMs indicated that accidents (including birds and mammals) have a higher probability of occurrence close to houses and on roads with high speed limits. When analysed separately, mammal kills occurred in sectors with high speed limits, close to houses and in areas surrounded by exotic bushes, while bird roadkills appeared in road sectors with high speed limits, close to houses and low traffic volume. Our findings highlight that roads are a potential threat to native birds in the eastern Canary Islands. Detailed studies on the local population dynamics of highly affected species, such as the Houbara Bustard, Eurasian Stone Curlew, Barn Owl or Southern Shrike, are urgently needed to determine whether these levels of road mortality are sustainable.

Dietary niche variation and its relationship to lizard population density.

Insular species are predicted to broaden their niches, in response to having fewer competitors. They can thus exploit a greater proportion of the resource spectrum. In turn, broader niches are hypothesized to facilitate (or be a consequence of) increased population densities. We tested whether insular lizards have broader dietary niches than mainland species, how it relates to competitor and predator richness, and the nature of the relationship between population density and dietary niche breadth. We collected population density and dietary niche breadth data for 36 insular and 59 mainland lizard species, and estimated competitor and predator richness at the localities where diet data were collected. We estimated dietary niche shift by comparing island species to their mainland relatives. We controlled for phylogenetic relatedness, body mass and the size of the plots over which densities were estimated. We found that island and mainland species had similar niche breadths. Dietary niche breadth was unrelated to competitor and predator richness, on both islands and the mainland. Population density was unrelated to dietary niche breadth across island and mainland populations. Our results indicate that dietary generalism is not an effective way of increasing population density nor is it result of lower competitive pressure. A lower variety of resources on islands may prevent insular animals from increasing their niche breadths even in the face of few competitors.

Testing the adaptive radiation hypothesis for the lemurs of Madagascar.

Lemurs, the diverse, endemic primates of Madagascar, are thought to represent a classic example of adaptive radiation. Based on the most complete phylogeny of living and extinct lemurs yet assembled, I tested predictions of adaptive radiation theory by estimating rates of speciation, extinction and adaptive phenotypic evolution. As predicted, lemur speciation rate exceeded that of their sister clade by nearly twofold, indicating the diversification dynamics of lemurs and mainland relatives may have been decoupled. Lemur diversification rates did not decline over time, however, as predicted by adaptive radiation theory. Optimal body masses diverged among dietary and activity pattern niches as lineages diversified into unique multidimensional ecospace. Based on these results, lemurs only partially fulfil the predictions of adaptive radiation theory, with phenotypic evolution corresponding to an ‘early burst’ of adaptive differentiation. The results must be interpreted with caution, however, because over the long evolutionary history of lemurs (approx. 50 million years), the ‘early burst’ signal of adaptive radiation may have been eroded by extinction.

The Mouse-colored Tyrannulet (Phaeomyias murina) is a species complex that includes the Cocos Flycatcher (Nesotriccus ridgwayi), an island form that underwent a population bottleneck

Simultaneous examination of evolutionary history in island forms and closely related mainland relatives can provide reciprocal insight into the evolution of island and mainland faunas. The Cocos Flycatcher (Nesotriccus ridgwayi) is a small tyrant flycatcher (Tyrannidae) endemic to Cocos Island, an oceanic island in the eastern Pacific Ocean. We first established its close relationship to the mainland species Mouse-colored Tyrannulet (Phaeomyias murina) using a phylogeny from genome-wide ultraconserved elements and exons. We then used mitochondrial DNA to explore the relationships between Nesotriccus and Phaeomyias populations from across its distribution in Central and South America. We found that Nesotriccus is nested within the Phaeomyias evolutionary tree, and that Phaeomyias represents a complex of at least four evolutionarily distinct species that differ in plumage, voice, and habitat association. Nesotriccus underwent a population bottleneck subsequent to its divergence from Central American and northern South American Phaeomyias populations in the middle Pleistocene. The 46 UCE loci containing alleles that are fixed between the two species are widely distributed across the genome, which suggests that selective or neutral processes responsible for divergence have occurred genome-wide. Overall, our simultaneous examination of Phaeomyias and Nesotriccus revealed divergent levels of genetic diversity and evolutionary histories between island and mainland forms.

Worldwide patterns of bird colouration on islands.

Island environments share distinctive characteristics that offer unique opportunities to investigate parallel evolution. Previous research has produced evidence of an island syndrome for morphological traits, life-history strategies and ecological niches, but little is known about the response to insularity of other important traits such as animal signals. Here, we tested whether birds' plumage colouration is part of the island syndrome. We analysed with spectrophotometry the colouration of 116 species endemic to islands and their 116 closest mainland relatives. We found a pattern of reduced brightness and colour intensity for both sexes on islands. In addition, we found a decrease in the number of colour patches on islands that, in males, was associated with a decrease in the number of same-family sympatric species. These results demonstrate a worldwide pattern of parallel colour changes on islands and suggest that a relaxation of selection on species recognition may be one of the mechanisms involved.

Origins of the Endemic Scaly Tree Ferns on the Galápagos and Cocos Islands

Premise of research. Successful long-distance dispersal is rarely observed in scaly tree ferns (Cyatheaceae). Nevertheless, recent molecular evidence has suggested that the four endemic scaly tree ferns on the Galápagos Archipelago (Cyathea weatherbyana) and Cocos Island (Cyathea alfonsiana, Cyathea nesiotica, and Cyathea notabilis), two oceanic island groups west of Central and northern South America, probably each originated from different mainland America ancestors. However, the phylogenetic relationships inferred among these endemics and their mainland relatives have been unclear. This study is aimed at better resolving the relationships and tracing the origins of these island endemics.Methodology. Five plastid regions from 35 Cyathea species were analyzed to reconstruct phylogenetic relationships using parsimony, likelihood, and Bayesian approaches. We also estimated divergence times of these species, and our chronogram was used to reconstruct their biogeographical range history.Pivotal results. Our well-resolved phylogenetic tree of Cyathea, which is in agreement with previous studies, shows that when the four Galápagos and Cocos endemics are included, they each belong to separate subclades. Our biogeographical study suggests that the four endemics originated from independent colonization events from mainland America and that there was no dispersal of Cyathea between the island groups. We reveal more detailed relationships among the endemics and their respective close mainland relatives; some of these relationships differ from previous studies. Our findings are corroborated by new morphological data from ongoing stem anatomy studies.Conclusions. The four scaly tree ferns endemic to the Galápagos and Cocos Islands each did indeed originate as independent colonization events from separate sources in mainland America, and their closest relatives are identified here.

Red Iron-Pigmented Tooth Enamel in a Multituberculate Mammal from the Late Cretaceous Transylvanian "Haţeg Island".

Mammals that inhabit islands are characterized by peculiar morphologies in comparison to their mainland relatives. Here we report the discovery of a partial skull associated with the lower jaws of a Late Cretaceous (≈70 Ma) multituberculate mammal from the Carpathian “Haţeg Island” of Transylvania, Romania. The mammal belongs to the Kogaionidae, one of the rare families that survived the Cretaceous—Paleogene mass extinction in Europe. The excellent preservation of this specimen allows for the first time description of the complete dentition of a kogaionid and demonstration that the enigmatic Barbatodon transylvanicus presents a mosaic of primitive and derived characters, and that it is phylogenetically basal among the Cimolodonta. Another peculiarity is the presence of red pigmentation in its tooth enamel. The red coloration is present on the anterior side of the incisors and on the cusps of most of the teeth. Energy-dispersive X-ray spectrometry (EDS) analysis reveals that the pigmented enamel contains iron, as in living placentals. Such a red pigmentation is known in living soricine shrews and many families of rodents, where it is thought to increase the resistance of the enamel to the abrasion that occurs during “grinding” mastication. The extended pattern of red pigment distribution in Barbatodon is more similar to that in eulipotyplan insectivores than to that in rodents and suggests a very hard diet and, importantly, demonstrates that its grasping incisors were not ever-growing. As inferred for other endemic Transylvanian vertebrates such as dwarf herbivorous dinosaurs and unusual theropod dinosaurs, insularity was probably the main factor of survival of such a primitive mammalian lineage relative to other mainland contemporaries of the Northern hemisphere.

Evolutionary Relationships, Gynodioecy, and Polyploidy in the Galápagos EndemicLycium minimum(Solanaceae)

Premise of research. Whereas most Lycium species are diploid and hermaphroditic, an association across Lycium species has been demonstrated between higher ploidy levels and gender dimorphism. Previous monographic work, in combination with a single tetraploid chromosome count, suggested that the Galápagos endemic Lycium minimum may be dimorphic. If substantiated, this would be the only documented Lycium species from South America to be dimorphic in gender expression, which is especially notable given that species diversity is highest in South America. The goals of this study are to (1) test the monophyly, characterize population-level genetic diversity, and determine the closest mainland relatives of L. minimum; (2) infer the sexual system of L. minimum; and (3) assess/verify ploidy using DNA content measurements.Methodology. Phylogenetic, morphological, and cytometric approaches were used to document evolutionary affinities, gynodioecy, and polyploidy. Phylogenetic analyses were conducted with DNA sequence data from both plastid and nuclear regions. Plastid data were also subject to network analysis, and genetic diversity within and among islands was examined using nuclear data. To determine whether populations were dimorphic in gender expression, we made floral measurements and documented pollen presence/absence. DNA content was determined using flow cytometry for multiple individuals from four islands.Pivotal results. Our findings support a South American origin for the Galápagos endemic L. minimum, with the Argentine L. athium as the closest living relative. Island populations exhibit significant population structure, with geographic distance positively correlated with genetic distance. Populations of L. minimum are gynodioecious; plants vary in the presence of pollen, and those without pollen have reduced anther length, width, and height.Conclusions. Morphological evidence and DNA content data confirm gender dimorphism and polyploidy in L. minimum, the first documented instance of a dimorphic Lycium species in South America.

The repeated evolution of large seeds on islands.

Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous species often evolve to become woody and species frequently evolve larger leaves, regardless of growth form. However, our understanding of how seed sizes might evolve on islands lags far behind other plant traits. Here, we conduct the first test for macroevolutionary patterns of seed size on islands. We tested for differences in seed size between 40 island-mainland taxonomic pairings from four island groups surrounding New Zealand. Seed size data were collected in the field and then augmented by published seed descriptions to produce a more comprehensive dataset. Seed sizes of insular plants were consistently larger than mainland relatives, even after accounting for differences in growth form, dispersal mode and evolutionary history. Selection may favour seed size increases on islands to reduce dispersibility, as long-distance dispersal may result in propagule mortality at sea. Alternatively, larger seeds tend to generate larger seedlings, which are more likely to establish and outcompete neighbours. Our results indicate there is a general tendency for the evolution of large seeds on islands, but the mechanisms responsible for this evolutionary pathway have yet to be fully resolved.

Speciation via floral heterochrony and presumed mycorrhizal host switching of endemic butterfly orchids on the Azorean archipelago

• Premise of the study: Most orchid species native to the Macaronesian islands reflect immigration from western Europe or North Africa followed by anagenesis. The only putative exception is the butterfly orchids (Platanthera) of the Azores, where three species apparently reflect at least one cladogenetic speciation event. This multidisciplinary study explores the origin, speciation, phenotypic, and genotypic cohesion of these Azorean species and their mainland relatives.• Methods: Plants of Platanthera from 30 localities spanning all nine Azorean islands were compared with those of four continental European relatives for 38 morphometric characters; substantial subsets were also analyzed for plastid microsatellites, and for nrITS of both the orchids and their mycorrhizae.• Key results: Although the three Azorean and four mainland species are all readily distinguished morphometrically using several floral characters, and hybridization appears rare, divergence in ITS and especially plastid sequences is small. Despite occupying similar laurisilva habitats, the Azorean species differ radically in the identities and diversity of their mycorrhizal partners; specialism apparently increases rarity.• Conclusions: Although morphological evidence suggests two invasions of the islands from NW Africa and/or SW Europe, ITS data imply only one. As the molecular data are unable to distinguish among the potential mainland ancestors, two scenarios of relationship are explored that imply different ancestors. Both scenarios require both anagenetic and cladogenetic speciation events, involving homoplastic shifts in overall flower size and (often substantial) changes in the relative dimensions of individual floral organs. Limited genotypic divergence among the three species compared with greater phenotypic divergence suggests comparatively recent speciation. Mycorrhizae may be the most critical factor dictating the respective ecological tolerances, and thus the relative frequencies, of these species. The recent IUCN Red-List amalgamation of Azorean Platanthera taxa into a single species urgently requires reappraisal, as P. micrantha is an excellent indicator species of seminatural laurisilva forest and P. azorica is arguably Europe's rarest orchid.

Parallel patterns and trends in functional structures in extinct island mammals

Endemic mammalian species on islands are generally known to have followed a different evolutionary pathway than their mainland relatives. General patterns, such as body size trends, have been described regularly. However, most island mammal species are unique and each of them is adapted to a specific local niche as part of an equally specific ecological assemblage. Therefore, comparing island species across taxa, islands and time is inherently dangerous without understanding the adaptational value of the studied feature in the compared taxa and without taking the ecological setting of the taxa into account. In this contribution, general and recurring patterns are described per taxon. Some features, like body mass change and sturdy limbs, are relatively general, whereas most features, like bone fusions and change of orbital axis, occur only in a very few taxa. Some features are even contradictory, such as brain size and degree of hypsodonty, with each taxon having its own particular design. In conclusion, general patterns are more often than not just trends and need to be applied with caution.

Island tameness: living on islands reduces flight initiation distance

One of Darwin's most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator-prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.

The Implications of Increased Body Size in the Song Sparrows of the California Islands

We investigated morphological divergence between mainland and California Island populations of the Song Sparrow (Melospiza melodia). Body size in small terrestrial vertebrates has been hypothesized to be larger in island forms than in their mainland relatives. Previously, we established that although bill size in island populations was similar to that in mainland populations with similar climate, bill-size dimorphism averaged much greater on the islands. In this paper, we compare various measures of structural body size and body mass between the California Islands and the mainland. We found that average body mass is over 10% higher in island Song Sparrows than in comparable mainland populations. However, a lack of commensurate structural body size suggests that body condition might be better in island Song Sparrows. This higher size-corrected mass may be explained by the dominance hypothesis, greater food availability and/or reduced competition, or adaptive mass regulation in the face of reduced predation pressure on adults. Both greater sexual bill dimorphism and greater body mass suggest reduced competition for food for the island Song Sparrow: a possibility that invites further investigation.