Pleistocene Land Bridges Research Articles

Pleistocene land bridges act as semipermeable agents of avian gene flow in Wallacea

Cyclical periods of global cooling have been important drivers of biotic differentiation throughout the Quaternary. Ice age-induced sea level fluctuations can lead to changing patterns of land connections, both facilitating and disrupting gene flow. In this study, we test if species with differing life histories are differentially affected by Quaternary land connections. We used genome-wide SNPs in combination with mitochondrial gene sequences to analyse levels of divergence and gene flow between two songbird complexes across two Wallacean islands that have been repeatedly connected during glaciations. Although the two bird complexes are similar in ecological attributes, the forest and edge-inhabiting golden whistler Pachycephala pectoralis is comparatively flexible in its diet and niche requirements as compared to the henna-tailed jungle-flycatcher Cyornis colonus, which is largely restricted to the forest interior. Using population-genomic and coalescent approaches, we estimated levels of gene flow, population differentiation and divergence time between the two island populations. We observed higher levels of differentiation, an approximately two to four times deeper divergence time and near-zero levels of gene flow between the two island populations of the more forest-dependent henna-tailed jungle-flycatcher as compared to the more generalist golden whistler. Our results suggest that Quaternary land bridges act as semipermeable agents of gene flow in Wallacea, allowing only certain taxa to connect between islands while others remain isolated. Quaternary land bridges do not accommodate all terrestrial species equally, differing in suitability according to life history and species biology. More generalist species are likely to use Quaternary land connections as a conduit for gene flow between islands whereas island populations of more specialist species may continue to be reproductively isolated even during periods of Quaternary land bridges.

The effects of Pleistocene climate change on biotic differentiation in a montane songbird clade from Wallacea

The role of Pleistocene Ice Age in tropical diversification is poorly understood, especially in archipelagos, in which glaciation-induced sea level fluctuations may lead to complicated changes in land distribution. To assess how Pleistocene land bridges may have facilitated gene flow in tropical archipelagos, we investigated patterns of diversification in the rarely-collected rusty-bellied fantail Rhipidura teysmanni (Passeriformes: Rhipiduridae) complex from Wallacea using a combination of bioacoustic traits and whole-genome sequencing methods (dd-RADSeq). We report a biogeographic leapfrog pattern in the vocalizations of these birds, and uncover deep genomic divergence among island populations despite the presence of intermittent land connections between some. We demonstrate how rare instances of genetic introgression have affected the evolution of this species complex, and document the presence of double introgressive mitochondrial sweeps, highlighting the dangers of using only mitochondrial DNA in evolutionary research. By applying different tree inference approaches, we demonstrate how concatenation methods can give inaccurate results when investigating divergence in closely-related taxa. Our study highlights high levels of cryptic avian diversity in poorly-explored Wallacea, elucidates complex patterns of Pleistocene climate-mediated diversification in an elusive montane songbird, and suggests that Pleistocene land bridges may have accounted for limited connectivity among montane Wallacean biota.

Evolutionary history of the endangered shrub snapdragon (Galvezia leucantha) of the Galápagos Islands

AbstractAimThe endangered Galápagos shrub snapdragon (Galvezia leucantha, Antirrhineae, Plantaginaceae) is restricted to small populations on four islands. In this study, we appraised results from taxonomy, genetics, phylogenetics, phylogeography and pollination ecology to reconstruct the evolutionary history of the genus Galvezia.LocationPeru, continental Ecuador and Galápagos.MethodsWe sequenced the nuclear ribosomal ITS and two plastid regions, ndhF and ndhF‐rpL32, to infer the origin of Galvezia and patterns of colonization to and across the Galápagos archipelago, based on Bayesian inference and statistical parsimony analyses. To investigate genetic diversity and differentiation within G. leucantha, we screened the genome of six populations and obtained 194 AFLP fingerprints. Autogamy tests and pollination network analyses were performed to evaluate the colonization potential and to investigate the structure of the pollinators’ assemblage of Galvezia.ResultsRelationships of seven nucleotide‐substitution haplotypes and 11 nucleotide‐substitution ribotypes of Galvezia revealed monophyly for the Galapagos species. Dating estimates indicated divergence of the insular Galvezia lineage in the Middle‐Upper Pleistocene (0.66–0.09 Ma). In addition, distribution of genotypes (seven haplotypes, eight ribotypes) across the three continental species showed geographical differentiation, while low differentiation and distribution of G. leucantha. AFLP genetic diversity is relatively high (HT = 0.109), but a low proportion of the total allelic variance is attributed to variation among subspecies/islands (Hb = 0.035, hierarchical AMOVA: 3.77% of total variance). The endemic bee (Xylocopa darwinii) accounted for 87.30% of the floral visits to G. leucantha.Main conclusionsWe inferred a single origin for an insular lineage that colonized the Galápagos Islands from northern Peru in the Pleistocene. Recent colonization of the archipelago, Pleistocene land bridges between islands and active gene flow promoted by X. darwinii may account for the low‐moderate genetic differentiation of G. leucantha subspecies. An unusual pollination shift from ornithophily (hummingbirds on the continent) to entomophily (Xylocopa in the Galápagos Islands) is supported.

Comparative phylogeography of the endemic Japanese weasel (Mustela itatsi) and the continental Siberian weasel (Mustela sibirica) revealed by complete mitochondrial genome sequences

We analyzed the complete mitochondrial genome for 26 individuals of the endemic Japanese weasel (Mustela itatsi) and 20 individuals of its continental sister species, the Siberian weasel (Mustela sibirica). These species diverged an estimated 1.19 Mya, in the Early Pleistocene. Mustela itatsi comprised two haplotype clades that diverged an estimated 0.93 Mya, in the Middle Pleistocene: a northern (Honshu) clade comprising geographically distinct basal, northern, and eastern subclades, and a western paraphyletic group, as well as a southern clade comprising geographically distinct subclades on Kyushu, Shikoku, and adjacent small islands. Our results indicate a single migration of an ancestral population from the Korean Peninsula to southern Japan across an Early Pleistocene land bridge, followed by allopatric speciation of M. itatsi in Japan. The southern lineage appears to have remained in place, whereas the range of the northern lineage expanded stepwise from south-western to northern Honshu between 0.31 and 0.11 Mya. Mustela sibirica also comprised two main clades that diverged an estimated 0.67 Mya; one containing haplotypes from continental Russia and Tsushima Island (Japan), and the other containing haplotypes from Korea, China, and Taiwan. The M. sibirica population on Tsushima Island is likely a relict from the continental Russian population.

Whole mitochondrial genomes illuminate ancient intercontinental dispersals of grey wolves (Canis lupus)

AbstractAimGrey wolves (Canis lupus) are widespread across the Holarctic. Here, we test the previously proposed hypothesis that extant North American wolves originate from multiple waves of colonization from Asia. We also test the hypothesis that land connections have been important in the evolutionary history of other isolated wolf populations in Japan.LocationHolarctic.MethodsWe analyse 105 previously published and newly obtained complete mitochondrial genomes from a geographically diverse sample of grey wolves and date critical branches in the phylogenetic tree. Phylogeographical hypotheses are tested in an approximate Bayesian computation approach.ResultsWe find that the mitogenomes of all living wolves in North America, including Mexican wolves, most likely derive from a single colonization event from Eurasia that expanded the grey wolf range into North America. This colonization occurred while a land bridge connected Eurasia and North America before the Cordillerian and Laurentide ice sheets fused in the Last Glacial Maximum,c. 23 ka, much more recent than predicted based on the fossil record. Pleistocene land bridges also facilitated the separate colonization of Hokkaido and the southern Japanese islands.Main conclusionsExtant wolf lineages in North America derive from wolves that migrated into North America coincident with the formation of the most recent land bridge with Eurasia. The maternal lineages from earlier Pleistocene American wolves are not represented in living American wolves, indicating that they left no descendants. The timing of colonization of North America, Hokkaido and the southern Japanese islands corresponds to the changes in land connectivity as a consequence of changing sea level.

The biogeographical history of Asian keelback snakes of the genusHebius(Squamata: Colubridae: Natricinae) in the Ryukyu Archipelago, Japan

The islands of the central and southern parts of the Ryukyu Archipelago and Taiwan accommodate suites of allopatric endemics for many terrestrial vertebrates. These subregions, which were isolated as three independent islands since the Late Miocene period, became connected to the continent by the Early Pleistocene land bridge (EPL). Many endemics are considered to have emerged through vicariant events accompanied with the collapse of EPL, but this is not well corroborated. We conducted divergence dating analyses for samples of four allopatric keelback snakes from 20 islands using a published timeframe for natricine snakes. Phylogenetic analyses recognised the existence of eight subclades for the four species. Four of the five subclades in the Central Ryukyus have emerged after the initial isolation of three subregions but before the EPL collapse, indicating iterative divergences within the super-island and stagnation of diverged entities. In contrast, the Southern Ryukyus was subjected to multiple incursions from both Central Ryukyus and Taiwan, irrespective of the EPL connection. Consequently, among all seven nodes, only two were consistent with the assumption of vicariance. These findings suggest that dispersals and subsequent nonvicariant divergences have contributed to the formation of the current phylogeographical pattern in this continental island chain.

Pseudamnicola Paulucci, 1878 (Caenogastropoda: Truncatelloidea) from the Aegean Islands: a long or short story?

The aims of the study were (i) to reveal the pattern of phylogeny of Pseudamnicola inhabiting the Aegean Islands, (ii) to describe and analyse the variation of the morphology in 17 populations of Pseudamnicola from the springs on the Aegean Islands not studied so far and considering also another seven populations studied earlier and (iii) to find out which model is more applicable to the island Pseudamnicola populations: either a model in which a relict fauna rich in endemics is differentiated in a way that mainly reflects the geological history of the area or a model in which a relatively young fauna is composed of more or less widely distributed taxa, with relatively high levels of gene flow among the springs they inhabit. To address the above issues, the morphology and the mitochondrial genes—cytochrome oxidase subunit I (COI) and ribosomal 16S—and nuclear genes—ribosomal 18S, 28S and histone 3 (H3)—were analysed. COI and COI+16S rRNA+18S datasets gave trees with identical topology in both ML and Bayesian inference. The 24 studied populations of Pseudamnicola form 16 clades, each of them generally having low levels of intrapopulation genetic differentiation. The generalised mixed Yule coalescent (GMYC) procedure and the Automatic Barcode Gap Discovery (ABGD) analysis for COI identified 16 Pseudamnicola entities coinciding with clades of the ML tree based on 44 haplotypes and 189 sequences. The present pattern of diversity, together with dating of divergence time, reflects a short story of colonisation/recolonisation, supported by the Late Pleistocene land bridges, rather than the consequences of earlier geological events. The principal component analysis (PCA) on the shells of the molecularly distinct clades showed differences, although variability ranges often overlap. Female reproductive organs showed no differences between the clades, and penile characters differed only in some cases.

Phylogeography of the prehensile‐tailed skinkCorucia zebrataon the Solomon Archipelago

The biogeography of islands is often strongly influenced by prior geological events. Corucia zebrata (Squamata: Scincidae) is endemic to the geologically complex Solomon Archipelago in Northern Melanesia. We examined the level of divergence for different island populations of C. zebrata and discussed these patterns in light of Pleistocene land bridges, island isolation, and island age. Corucia zebrata was sampled from 14 locations across the Solomon Archipelago and sequenced at two mitochondrial genes (ND2 and ND4; 1697 bp in total) and four nuclear loci (rhodopsin, an unknown intron, AKAP9, and PTPN12). Measures of genetic divergence, analyses of genetic variation, and Bayesian phylogenetic inference were used and the data assessed in light of geological information. Populations of C. zebrata on separate islands were found to be genetically different from each other, with reciprocal monophyly on mitochondrial DNA. Populations on islands previously connected by Pleistocene land bridges were marginally less divergent from each other than from populations on other nearby but isolated islands. There are indications that C. zebrata has radiated across the eastern islands of the archipelago within the last 1–4 million years. Nuclear loci were not sufficiently informative to yield further information about the phylogeography of C. zebrata on the Solomon Archipelago. Analyses of the mitochondrial data suggest that dispersal between islands has been very limited and that there are barriers to gene flow within the major islands. Islands that have been isolated during the Pleistocene glacial cycles are somewhat divergent in their mitochondrial genotypes, however, isolation by distance (IBD) and recent colonization of isolated but geologically younger islands appear to have had stronger effects on the phylogeography of C. zebrata than the Pleistocene glacial cycles. This contrasts with patterns reported for avian taxa, and highlights the fact that biogeographic regions for island species cannot be directly extrapolated among taxa of differing dispersal ability.

Biogeography of tenebrionid beetles (Coleoptera: Tenebrionidae) in the circum-Sicilian islands (Italy, Sicily): Multiple biogeographical patterns require multiple explanations

The tenebrionid beetles on 25 circum-Sicilian islands were studied to determine the influence of island geographical and landscape features on three main intercorrelated biogeographical patterns: (1) species richness, studied using species-area and species environment relationships, (2) species assemblage composition, investigated using Canonical Correspondence Analysis (CCA), and (3) inter-site faunal similarity, investigated using Canonical Correlation Analysis (CANCOR) applied to multidimensional scaling of inter-island faunal dissimilarities. Species richness was mostly influenced by island area and landscape heterogeneity (expressed using various indices of diversity based on land cover categories). When species identities were considered in the CCA, no substantial effect of landscape was detected. Current island isolation did not have a strong influence on species richness, but has a distinct effect in determining species assortments on the remotest islands. Historical influences of Pleistocene landbridge connections were not detectable in species richness relationships using geographical variables in species richness analyses or in assemblage gradients in the CCA, but emerged distinctly from inter-island similarities in the CANCOR.

Phylogeography and historical demography of the Italian treefrog,Hyla intermedia,reveals multiple refugia, population expansions and secondary contacts within peninsular Italy

We investigated the geographical patterns of genetic diversity in the Italian treefrog through sequence analysis of a mitochondrial cytochrome b gene fragment. Three main mitochondrial lineages were identified, distributed in northern, central and southern Italy, respectively. Their divergence appears indicative of a split time largely predating Late Pleistocene climatic oscillations, and syntopy between them was only observed in the geographically intermediate populations. The historical demographic reconstructions suggest that in both northern and central Italy, an expansion occurred during the last major glacial phase, when a vast widening of the lowland habitats followed the glaciation-induced fall of the sea level. Instead, in southern Italy an expansion event likely followed the end of the last glaciation, although the inference of expansion appears less reliable for the southern clade than for the others. Within this geographical area, a sharp phylogeographic discontinuity separated peninsular from Sicilian populations, and the overall pattern of diversity suggests that the latter derived from a recent colonization of the island, probably through a Late Pleistocene land bridge. Phylogenetic, phylogeographic and historical demographic analyses thus concur in delineating a scenario of multiple refugia, with four groups of populations which survived the last glacial-interglacial cycles in at least three distinct refugia arranged along peninsular Italy, and have recently come into contact following range expansions. Therefore, these results support the hypothesis that a plethora of microevolutionary processes, rather than the prolonged stability of populations, were mainly responsible for shaping the patterns of diversity within this major biodiversity hotspot.

History, ocean channels, and distance determine phylogeographic patterns in three widespread Philippine fruit bats (Pteropodidae)

The comparative phylogeography of widespread, codistributed species provides unique insights into regional biodiversity and diversification patterns. I used partial DNA sequences of the mitochondrial genes ND2 and cyt b to investigate phylogeographic structure in three widespread Philippine fruit bats. Ptenochirus jagori is endemic to the oceanic region of the Philippines and is most abundant in lowland primary forest. Macroglossus minimus and Cynopterus brachyotis are most common in disturbed and open habitats and are not endemic. In all three, genetic differentiation is present at multiple spatial scales and is associated to some degree with Pleistocene landbridge island groups. In P. jagori and C. brachyotis, genetic distance is correlated with geographic distance; in C. brachyotis and M. minimus, it is correlated with the sea-crossing distance between islands. P. jagori has the least overall genetic structure of these three species, whereas C. brachyotis and M. minimus have more geographic association among haplotypes, suggesting that phylogeographic patterns are linked to ecology and habitat preference. However, contrary to expectation, the two widespread, disturbed habitat species have more structure than the endemic species. Mismatch distributions suggest rapid changes in effective population size in C. brachyotis and P. jagori, whereas M. minimus appears to be demographically more stable. Geologic and geographic history are important in structuring variation, and phylogeographic patterns are the result of dynamic long-term processes rather than simply reflecting current conditions.

ASSESSING THE TAXONOMIC STATUS OF THE PALAWAN PANGOLIN MANIS CULIONENSIS (PHOLIDOTA) USING DISCRETE MORPHOLOGICAL CHARACTERS

Abstract We use discrete morphological characters in a statistical framework to reassess the taxonomic status of the Palawan pangolin Manis culionensis relative to the Sunda pangolin M. javanica. We recommend that the 15 species-level traits previously proposed in the literature to distinguish the 2 pangolins be replaced by 5 newly defined diagnostic characters related to skull and external scales. Our study supports species-level partition between the Palawan and Sunda pangolins at a frequency of expected polymorphism threshold fixed to 0.10. Isolation through sea level rising (approximately 800,000–500,000 years ago) of proto-Palawan pangolins coming from Borneo through Early Pleistocene land bridges might have promoted the speciation of M. culionensis, a Palawan endemic species to be considered of high conservation concern.

Phylogeographic patterns of HawaiianMegalagriondamselflies (Odonata: Coenagrionidae) correlate with Pleistocene island boundaries

The Pleistocene geological history of the Hawaiian Islands is becoming well understood. Numerous predictions about the influence of this history on the genetic diversity of Hawaiian organisms have been made, including the idea that changing sea levels would lead to the genetic differentiation of populations isolated on individual volcanoes during high sea stands. Here, we analyse DNA sequence data from two closely related, endemic Hawaiian damselfly species in order to test these predictions, and generate novel insights into the effects of Pleistocene glaciation and climate change on island organisms. Megalagrion xanthomelas and Megalagrion pacificum are currently restricted to five islands, including three islands of the Maui Nui super-island complex (Molokai, Lanai, and Maui) that were connected during periods of Pleistocene glaciation, and Hawaii island, which has never been subdivided. Maui Nui and Hawaii are effectively a controlled, natural experiment on the genetic effects of Pleistocene sea level change. We confirm well-defined morphological species boundaries using data from the nuclear EF-1alpha gene and show that the species are reciprocally monophyletic. We perform phylogeographic analyses of 663 base pairs (bp) of cytochrome oxidase subunit II (COII) gene sequence data from 157 individuals representing 25 populations. Our results point to the importance of Pleistocene land bridges and historical island habitat availability in maintaining inter-island gene flow. We also propose that repeated bottlenecks on Maui Nui caused by sea level change and restricted habitat availability are likely responsible for low genetic diversity there. An island analogue to northern genetic purity and southern diversity is proposed, whereby islands with little suitable habitat exhibit genetic purity while islands with more exhibit genetic diversity.

The shaping of mitochondrial DNA phylogeographic patterns of the brown hare ( Lepus europaeus) under the combined influence of Late Pleistocene climatic fluctuations and anthropogenic translocations

The phylogeographic structure of the brown hare ( Lepus europaeus) was studied by analysing mtDNA control region sequences of 98 individuals from continental and insular Greece, Bulgaria, Cyprus and northern Israel, together with 44 published sequences from Italy and central Europe. We found two distinct clades separated by an average nucleotide divergence of 6.6%, which may correspond to a Balkan and to an Asia Minor refugium. The estimated time of separation of the two clades was dated back to 105,000– 490,000 years ago. These two clades coexist in the area of northeastern Greece and Bulgaria, most likely as a result of a post-glacial northward expansion. Within the southern Balkan refugium, network analyses showed geographical structuring, which supports the hypothesis of several isolated Late Pleistocene populations. The central European and Italian populations appear to have originated from a non-detected northern Balkan population that was genetically closely related to some northern Greek populations, as a result of postglacial expansion, translocations or a combination of both. Moreover, several cases of ancient and recent translocations by humans were detected, especially for some island populations, while the eastern Aegean islands off the Asia Minor coast were most likely colonized naturally through Late Pleistocene land bridge connection. The genetic analysis presented here provides a framework for designing proper conservation and management guidelines for this species.

Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys whiteheadi)

The Sunda region of south-east Asia comprises the Malay Peninsula and the islands of Java, Sumatra and Borneo, all of which lie on a shallow continental shelf projecting from Indochina. Pleistocene glacial cycles caused sea levels to drop repeatedly, exposing vast areas of the Sunda shelf and creating land bridges among the islands and mainland. These land bridges, the latest of which connected all three of the major Sunda islands to the Malay Peninsula as recently as 9500 years ago, may have enabled mammalian migrations across the Sunda shelf. Pleistocene land bridges on the Sunda shelf have been invoked to explain the current distributions of mammalian taxa occupying ranges corresponding with the Pleistocene limits of land and the appearance of new mammal species in the Pleistocene fossil record. The ability of mammals to move throughout the exposed shelf during periods of low sea level would, however, have been influenced by topographic and ecological features, which have been variously described as savanna-like or as moist tropical rain forest. Using a phylogeographical approach, we test the hypothesis that Pleistocene land bridges enabled widespread movements in three rain-forest-restricted murine rodents of the Sunda shelf: Maxomys surifer, Leopoldamys sabanus and Maxomys whiteheadi. Our results do not support the hypothesis of broad Pleistocene migrations in these taxa, but instead suggest a deep history of vicariant evolution that may correspond with the Pliocene fragmentation of the Sunda block. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81, 91–109.

Biogeography of mammals in SE Asia: estimates of rates of colonization, extinction and speciation

Four categories of islands in SE Asia may be identified on the basis of their histories of landbridge connections. Those islands on the shallow, continental Sunda Shelf were joined to the Asian mainland by a broad landbridge during the late Pleistocene; other islands were connected to the Sunda Shelf by a middle Pleistocene landbridge; some were parts of larger oceanic islands; and others remained as isolated oceanic islands. The limits of late Pleistocene islands, defined by the 120 m bathymetric line, are highly concordant with the limits of faunal regions. Faunal variation among non-volant mammals is high between faunal regions and low within the faunal regions; endemism of faunal regions characteristically exceeds 70%. Small and geologically young oceanic islands are depauperate; larger and older islands are more species-rich. The number of endemic species is correlated with island area; however, continental shelf islands less than 125 000 km2 do not have endemic species, whereas isolated oceanic islands as small as 47 km2 often have endemic species. Geologically old oceanic islands have many endemic species, whereas young oceanic islands have few endemic species. Colonization across sea channels that were 5–25 km wide during the Pleistocene has been low, with a rate of about 1–2/500000 years. Comparison of species-area curves for mainland areas, late Pleistocene islands, and middle Pleistocene islands indicates that extinction occurs rapidly when landbridge islands are first isolated, with the extent of extinction dependent upon island size; extinction then slows to an average rate of 1–2%/10 000 years. The great majority of the non-volant Philippine mammals arrived from the Sunda Shelf, the geographically closest of the possible source areas. Speciation within the Philippines has contributed substantially to species richness, perhaps exceeding colonization by a factor of two or more as a contributor to species number. Colonization, extinction and speciation rates differ among taxonomic groups, with murid rodents being most successful and carnivores least successful. In order for any model of island biogeography to be widely applicable to insular faunas, the model must include speciation as a major variable. It is suggested that insular mammalian faunas typically are not in equilibrium, because geological and climatic changes can occur as rapidly as colonization and speciation.

Plate tectonics and the evolution of intertidal and shallow-water benthic biotic distribution patterns of the southwest Pacific

A brief summary is given of recent information on the changing palaeogeography of the southwest Pacific region following the break-up of Gondwanaland and the parallel palaeoceanographic changes that occurred. Present intertidal and shallow-water benthic marine provinces in Australasia can be grouped into tropical—subtropical, warm-temperate, transitional warm-temperate, cold-temperate, transitional subantarctic and subantarctic regions. A notable feature is the marked faunistic attenuation that occurs with increasing latitude in most groups. Historical explanations of biotic distribution patterns fall into two classes: dispersal explanations and vicariance explanations. The problems involved in evaluating the relative roles of dispersal and vicariance in the evolution of the biota of a region are outlined and the conditions which must be met before larval dispersal across wide stretches of ocean can be successful are reviewed. It is concluded that rafting of adults is restricted to a few groups of animals and is of limited importance as a dispersal mechanism. An examination of the external relationships anddfossil record of Australasian intertidal and shallow water biota has identified a number of elements of which the predominant one is a Malayo-Pacific one comprising plants and animals of tropical affinities. The evolutionary history of the biota is re-interpreted in the light of palaeogeographic and palaeoceanographic changes with reference to a series of palaeogeographic reconstructions showing successive positions of Antarctica, Australia and New Zealand which have been used as a geographic base upon which speculative patterns of oceanic circulation and positions of oceanographic fronts have been depicted. Tectonic events, coupled with changes in circulation patterns and sea temperatures and the establishment of the Antarctic and Subtropical Convergences with a series of water masses forming belts from the poles to the tropics have been controlling influences of the distribution and evolution of the marine biota of the region. A major factor has been the migrations that have occurred in the position of the warm-water/cold-water boundary, the Subtropical Convergence. This migration coupled with changing island archipelago nature of the New Zealand land area has played a major role in speciation processes in the New Zealand region. It also led to the extraction of many warm-water genera and species as it moved north. In Australia the larger land mass extending further to the north provided opportunities for species to migrate up both the east and west coasts. This, coupled with the Pleistocene land bridges which connected Tasmania and New Guinea to Australia, led to isolation and subsequent allopatric speciation, followed by sympatry when the derived species migrated along the coast as the Subtropical Convergence retreated south to its present position and the Torres and Bass Straits barriers disappeared with rises in sea level.

Density Compensation in Island Faunas

This paper analyzes factors determining the extent of density compensation on islands: i.e., is the summed population density of individuals of all species on islands equal to the summed mainland density as a result of niche expansions and higher abundances of island species compensating for the absence of many mainland species? In addition, a method is described for estimating bird population densities based on analysis of the time dependence or mist—netting yields. Puercos Island in the Pearl Archipelago off Panama has less than one—third as many resident birds species as comparable mainland habitats. Analysis of the Pearl avifauna suggests that about one—quarter of the island species may be relicts of the Pleistocene land bridge and that the remainder are subsequent over—water colonists. The successful colonists are a highly nonrandom sample of the mainland avifauna in such respects as family composition, social structure, and second—growth habitat origin. Song—based censuses and analysis of mist—netting show that Puercos has a slightly density of individuals than the mainland. Niche shifts between islands and mainland, or among different islands, include habitat expansions, wider ranges of vertical foraging strata, abundance increases checkerboard distribution patterns, and decreased morphological variability. Comparison of the present study with other studies shows that summed population densities on islands may be higher than, comparable to, or less than mainland levels, depending upon the particular island, habitat, and group of animals studied. Among factors affecting the extent of density colonists less appropriate to the vacated habitat, tending to lower island densities; and underrepresentation of large species on islands, tending to increase island population densities for a given biomass.