GENETIC EVIDENCE FOR MULTIPLE ORIGINS OF DIOECY IN THE HAWAIIAN SHRUB WIKSTROEMIA (THYMELAEACEAE)

Dioecy is unusually common in the Hawaiian Islands, yet little is known about the evolutionary biology of this breeding system. A native shrub, Wikstroemia, has an unusually diverse array of breeding systems: two forms of dioecy, cryptic and morphological dioecy, as well as hermaphroditism (perfect flowers). The existence of two forms of dioecy is significant for three reasons: 1) the presence of cryptic unisexuals that are functionally unisexual, but retain the appearance of hermaphroditism in both sexes, is strong evidence for the ancestral status of hermaphroditism; 2) the production of nonfunctional pollen, by female cryptic unisexuals, is a new instance of a phenomenon which has previously been reported for a few other species; 3) the two forms of dioecy are morphological markers which are useful in hybridization studies for tracing the genetic basis of their inheritance. Crosses were made between cryptically unisexual individuals (C), between morphologically unisexual individuals (M), and between the two types of unisexuality. The offspring of crosses between individuals with the same sex type usually resulted in offspring with that sex type, but most of the progeny of between-sex type crosses were, unexpectedly, perfect-flowered hermaphrodites. These results show that genetic control of sex determination is not homologous in all populations, suggesting that dioecy has evolved at least twice in Hawaiian Wikstroemia. The genetic data further suggest that males are the heterozygous sex.     

SEQUENTIAL RADIATIONS AND PATTERNS OF SPECIATION IN THE HAWAIIAN CRICKET GENUS LAUPALA INFERRED FROM DNA SEQUENCES

The tremendous diversity of endemic Hawaiian crickets is thought to have originated primarily through intraisland radiations, in contrast to an interisland mode of diversification in the native Hawaiian Drosophila. The Hawaiian cricket genus Laupala (family Gryllidae) is one of several native genera of flightless crickets found in rain-forest habitat across the Hawaiian archipelago. I examined the phylogenetic relationships among mitochondrial DNA (mtDNA) sequences sampled from 17 species of Laupala, including the 12S ribosomal RNA (rRNA), transfer RNA (RNA)^val and 16S rRNA regions. The distribution of mtDNA variants suggests that species within Laupala are endemic to single islands. The phylogenetic estimate produced from both maximum likelihood and maximum parsimony supports the hypothesis that speciation in Laupala occurred mainly within islands. The inferred biogeographical history suggests that diversification in Laupala began on Kauai, the oldest rain-forested Hawaiian island. Subsequently, colonization to younger islands in the archipelago resulted in a radiation of considerable phylogenetic diversity. Phylogenetic patterns in mtDNA are not congruent with prior systematic or taxonomic hypotheses. Hypotheses that may explain the conflict between the phylogenetic patterns of mtDNA variation and the species taxonomy are discussed.     

POLYGENIC INHERITANCE OF A BEHAVIORAL PHENOTYPE: INTERSPECIFIC GENETICS OF SONG IN THE HAWAIIAN CRICKET GENUS LAUPALA

The Hawaiian cricket genus Laupala (family Gryllidae) is one of several native genera of flightless crickets found in rain-forest habitat across the Hawaiian archipelago. Species in this genus are morphologically quite similar, but the songs produced by adult males are acoustically distinct. I examined the nature of song variation found within Laupala paranigra and between Laupala kohalensis and L. paranigra, both endemic to the island of Hawaii. Variation within and among species was most notable in the temporal structure of the song, as quantified by the pulse rate. The variation in pulse rate present in natural populations of L. paranigra bred true through the F₁ laboratory generation, suggesting that the intraspecific variation in this species has a genetic basis. Interspecific hybridizations between L. kohalensis and L. paranigra successfully produced F₁, F₂, and backcross generations. Hybrid F₁ males from reciprocal crosses sang with significantly different pulse rates, implicating an X chromosomal contribution to the phenotypic difference between these species. Interspecific patterns of inheritance are most consistent with a type-I genetic architecture. Polygenic inheritance of the interspecific pulse-rate variation was observed, and approximately eight genetic factors were estimated to underlie the difference in pulse rate between L. kohalensis and L. paranigra.     

Multiple colonizations of a remote oceanic archipelago by one species: how common is long‐distance dispersal?

Aim  It is well established that many groups of plants and animals have undergone long-distance dispersal, but the extent to which this continues beyond initial colonization is largely unknown. To provide further insight into the frequency of gene flow mediated by long-distance dispersal, we investigated the origins of the fern Asplenium hookerianum on the Chatham Islands, and present a review of the contribution of molecular data to elucidating the origins of this archipelago's biota.
Location  Chatham Islands and New Zealand. A. hookerianum is scarce on the Chatham Islands but common in New Zealand, some 800 km to the west.
Methods  We compared chloroplast trnL–trnF DNA sequence data from Chatham Islands' A. hookerianum with extensive phylogeographic data for this genetically variable species in mainland New Zealand.
Results  Our sequencing revealed the presence of two haplotypes in Chatham Islands' A. hookerianum . These haplotypes differed by four mutational events and were each more closely related to haplotypes found in New Zealand than to each other.
Main conclusions  Despite the rarity of A. hookerianum on the Chatham Islands, its populations there appear to derive from at least two long-distance dispersal events from New Zealand, these possibly originating from different areas. We suggest that long-distance transoceanic dispersal, and the gene flow it can mediate, may be more common than is generally appreciated.     

EVOLUTION OF UNISEXUALITY IN THE HAWAIIAN FLORA: A TEST OF MICROEVOLUTIONARY THEORY

The evolution of separate sexes as a means of avoiding self-fertilization requires the controversial coexistence of large inbreeding depression and high selfing rate in the ancestral hermaphrodite population. Fitness components of adult females and hermaphrodites in nature, of their open-pollinated progeny, and of experimental selfs and outcrosses onto hermaphrodites were compared in endemic Hawaiian Bidens sandvicensis, all of whose known populations are gynodioecious, consisting of a mixture of females and hermaphrodites. Multilocus selfing rates of hermaphrodites were also estimated, and sex morph ratio monitored over four seasons in three populations of B. sandvicensis and one population of gynodioecious B. cervicata. Total mean inbreeding depression in seed set (in the glasshouse), germination rate (in an open-air nursery on Kauai), and first year survivorship and fecundity in the field were estimated as 0.94 (SE 0.04), and occurred primarily in drought months. Lower survivorship and fecundity of selfs were partially explained by their consistently smaller size. Open-pollinated seed of females had significantly lower germination rate, proportion flowering, and fecundity than outcrossed progeny of hermaphrodites, suggesting moderate biparental inbreeding in females and a lack of any non-outcrossing advantage to progeny of females. In all fitness components, open-pollinated progeny of hermaphrodites were inferior to those of females and to outcrosses, and in most components were superior to selfs. Total performance of open-pollinated progeny of females relative to those of hermaphrodites was calculated as 2.3 (SE = 0.4), but since inflorescences of females also set 20% to 50% more seed than those of hermaphrodites, their total relative ovule success was estimated as 3.2 (SE = 0.5). If inheritance of male sterility is nuclear, this superiority is sufficient to maintain females in frequencies over 20% in populations, whose actual frequencies ranged from 14% to 33%. In four populations, selfing rates of hermaphrodites, assayed in seedlings, were 0.50, 0.45, 0.25, and 0.30, but since substantial inbreeding depression occurred prior to germination, the mean selfing rate of hermaphrodite ovules exceeded 0.57. Female frequencies were significantly higher in the two populations with higher hermaphrodite selfing rate. These results suggest that inbreeding depression can exert a profound influence on the mating system of self-compatible plants on Hawaii and perhaps other oceanic islands, and can be sufficiently strong to electively favor the elimination of the male function.     

Speciation in Hawaiian angiosperm lineages: cause, consequence, and mode

The biota of Hawaiian Islands is derived entirely from long distance dispersal, often followed by in situ speciation. Species descended from each colonist constitute monophyletic lineages that have diverged to varying degrees under similar spatial and temporal constraints. We partitioned the Hawaiian angiosperm flora into lineages and assessed morphological, ecological, and biogeographic characteristics to examine their relationships to variation in species number (S). Lineages with external bird dispersal (through adhesion) were significantly more species-rich than those with abiotic dispersal, but only weakly more species-rich than lineages with internal bird dispersal (involving fleshy fruits). Pollination mode and growth form (woody vs. herbaceous) had no significant effect on S, in contrast to studies of angiosperm families. S relates positively to the geographic and ecological range size of whole lineages, but negatively to local abundance and mean range sizes of constituent species. Species-rich lineages represent a large proportion of major adaptive shifts, although this appears to be an artifact of having more species. Examination of 52 sister species pairs in numerous lineages provides evidence for allopatric (including peripheral isolates) and parapatric (ecological) modes, with 15 cases of each. Although postspeciational dispersal may obscure these modes in many of the remaining cases, instances of sympatric and hybrid speciation are also discussed. Because speciation is both a consequence and a cause of ecological and biogeographic traits, speciation mode may be integral to relationships between traits. We discuss the role of speciation in shaping the regional species pool.     

MULTIPLE GENETIC LINKAGES BETWEEN FEMALE PREFERENCE AND MALE SIGNAL IN RAPIDLY SPECIATING HAWAIIAN CRICKETS

Diverging sexual communication systems can lead to the evolution of new species that no longer recognize each other as potential mates. The coevolution of male and female components of sexual communication is facilitated by physical linkage between genes underlying signals and preferences. By crossing two closely related Hawaiian crickets (Laupala kohalensis and Laupala paranigra) with vastly different song pulse rates and female preferences, and assessing segregation of songs and preferences among second generation backcrosses, we show a strong genetic correlation between song and preference variation. Furthermore, multiple, but not all, quantitative trait loci underlying song variation also predict female preferences. This physical linkage or pleiotropy may have facilitated the striking diversification of pulse rates observed among Laupala species in conjunction with one of the most rapid species radiations so far recorded.     

Molecular biogeography and diversification of the endemic terrestrial fauna of the Hawaiian Islands

Oceanic islands have played a central role in biogeography and evolutionary biology. Here, we review molecular studies of the endemic terrestrial fauna of the Hawaiian archipelago. For some groups, monophyly and presumed single origin of the Hawaiian radiations have been confirmed (achatinelline tree snails, drepanidine honeycreepers, drosophilid flies, Havaika spiders, Hylaeus bees, Laupala crickets). Other radiations are derived from multiple colonizations (Tetragnatha and Theridion spiders, succineid snails, possibly Dicranomyia crane flies, Porzana rails). The geographic origins of many invertebrate groups remain obscure, largely because of inadequate sampling of possible source regions. Those of vertebrates are better known, probably because few lineages have radiated, diversity is far lower and morphological taxonomy permits identification of probable source regions. Most birds, and the bat, have New World origins. Within the archipelago, most radiations follow, to some degree, a progression rule pattern, speciating as they colonize newer from older islands sequentially, although speciation often also occurs within islands. Most invertebrates are single-island endemics. However, among multi-island species studied, complex patterns of diversification are exhibited, reflecting heightened dispersal potential (succineids, Dicranomyia). Instances of Hawaiian taxa colonizing other regions are being discovered (Scaptomyza flies, succineids). Taxonomy has also been elucidated by molecular studies (Achatinella snails, drosophilids). While molecular studies on Hawaiian fauna have burgeoned since the mid-1990s, much remains unknown. Yet the Hawaiian fauna is in peril: more than 70 per cent of the birds and possibly 90 per cent of the snails are extinct. Conservation is imperative if this unique fauna is to continue shedding light on profound evolutionary and biogeographic questions.     

STUDIES OF HAWAIIAN FRESHWATER AND SOIL ALGAE II. ALGAL COLONIZATION AND SUCCESSION ON A DATED VOLCANIC SUBSTRATE1,2

Spatial studies of colonization and succession of soil algae and chemical analyses of the various soils on the cinde cone of Kilauea Iki in Hawaii Volcanoes National Park, Hawaii are outlined. There is a positive correlation between the diversity and quantity of soil algae with nutrient levels and organic matter accumulation in each locale. Three distinct edaphic-biotic zones existing in this area are differentially revealed by the soil chemical composition, quantity and diversity of soil algae, and as evident variations in higher plant growth and colonization. Varying colonization and successional phases of higher plant growth around standing and fallen tree snags killed by volcanic activity also reflect variations in the soil algal flora. These variations appear largely as a function of differential water interception, absorption, and retention as well as differential accumulation of organic matter, and the initiation of various biogeochemical cycles.     

A COPROLOGICAL SURVEY OF HELMINTH PARASITES OF THE HAWAIIAN MONK SEAL FROM THE NORTHWESTERN HAWAIIAN ISLANDS

In conjunction with an Hawaiian monk seal, Monachus schauinslandi , breeding enhancement program, fecal samples were examined for the presence of helminth parasite products (eggs, larvae, and proglottids). Eighty-two samples were examined from four island groups in the northwestern Hawaiian Islands: 38 from Kure Atoll, 32 from French Frigate Shoals, 11 from Laysan Island, and one from Lisianski Island. Identified helminth products include those of Diphyllobothrium sp., D. cameroni, Contracaecum turgidum, Anisakis sp., Corynosoma rauschi , and Trematoda. In addition, eggs from Halarachne laysanae (Atari) were found in 3 samples. Eggs of C. turgidum were present in the highest percentage (>64%) of samples from all locations. Eggs of Anisakis sp. and Trematoda are reported from this host for the first time.     

东南亚生物多样性热点地区的形成与演化

东南亚地处热带,生物多样性极为丰富,可分为4个热点地区:印度-缅甸区的中南半岛、巽他区(含马来半岛、婆罗洲、苏门答腊岛)、菲律宾区(菲律宾群岛)、华莱士区(苏拉威西岛、爪哇岛、马鲁古群岛、小巽他群岛等)。中南半岛在泥盆纪便已是欧亚大陆的一部分,在印度板块撞击欧亚大陆之后受挤压而出;巽他区来自于冈瓦纳古陆和澳洲古陆;菲律宾群岛部分来自于劳亚古陆的碎片向南漂移,部分来自于太平洋西南岛弧的向北迁移;华莱士区则是劳亚古陆碎片、太平洋西南岛弧以及澳洲古陆北侧碎片的组合。巽他区地处赤道,常年温湿;菲律宾区、华莱士区、中南半岛则都受到不同程度的季风气候决定的干湿季变动。地质历史和季风气候影响程度的不同,奠定了东南亚4个生物多样性热点地区的雏形。华莱士区保存有大量的早期被子植物原始类群如睡莲目(Nymphaeales)和木兰藤目(Austrobaileyales),是现代被子植物起源地和冰期避难所之一。巽他区(婆罗洲)和中南半岛是亚洲热带植物的现代分布中心和"进化前沿",是整个东南亚地区重要的种源;而华莱士区的爪哇岛和小巽他群岛主要是物种迁入和中转的种库。这样的物种形成历史与迁移格局,塑造了东南亚4个...     

Impacts of Restored Patch Density and Distance from Natural Forests on Colonization Success

Abstract The reduction and fragmentation of forest habitats is expected to have profound effects on plant species diversity as a consequence of the decreased area and increased isolation of the remnant patches. To stop the ongoing process of forest fragmentation, much attention has been given recently to the restoration of forest habitat. The present study investigates restoration possibilities of recently established patches with respect to their geographical isolation. Because seed dispersal events over 100 m are considered to be of long distance, a threshold value of 100 m between recent and old woodland was chosen to define isolation. Total species richness, individual patch species richness, frequency distributions in species occurrences, and patch occupancy patterns of individual species were significantly different among isolated and nonisolated stands. In the short term no high species richness is to be expected in isolated stands. Establishing new forests adjacent to existing woodland ensures higher survival probabilities of existing populations. In the long term, however, the importance of long‐distance seed dispersal should not be underestimated because most species showed occasional long‐distance seed dispersal. A clear distinction should be made between populations colonizing adjacent patches and patches isolated from old woodland. The colonization of isolated stands may have important effects on the dynamics and diversity of forest networks, and more attention should be directed toward the genetic traits and viability of founding populations in isolated stands.     

REPRODUCTIVE PATTERNS OF THE HAWAIIAN MONK SEAL

We evaluated reproductive patterns of the Hawaiian monk seal ( Monachus schauinslandi ) using a combination of fitted age-specific reproductive curves and analysis of reproductive patterns of individual females. We review the difficulties inherent in the acquisition and modeling of reproductive data with emphasis on the significance of reproductive senescence to populations with dissimilar age/sex compositions. Validation of the fitted reproductive parameters was accomplished by Monte Carlo sampling of parameter distributions to compare the expected number of pups with the observed production. Although the fitted reproductive functions appear to provide an acceptable fit to the raw reproductive data, we found that the fitted curves did a poor job of predicting the actual pup production in individual years because of high variability among years. To further verify, and elaborate on, the patterns in the pooled (multi-seal, and multi-year) rates, we examined attributes of the reproductive performance of individual seals. The attributes included age of primiparity, reproductive rates computed over several age ranges, and the relationship between reproductive performance and seal longevity. Analysis of individual seal patterns reinforced the conclusion that reproductive senescence is operative in monk seal populations.     

ISLAND BIOGEOGRAPHY OF GALÁPAGOS LAVA LIZARDS (TROPIDURIDAE: MICROLOPHUS): SPECIES DIVERSITY AND COLONIZATION OF THE ARCHIPELAGO

The "lava lizards" ( Microlophus ) are distributed throughout the Galápagos Archipelago, and consist of radiations derived from two independent colonizations. The "Eastern Radiation" includes M. bivittatus and M. habeli endemic to San Cristobal and Marchena Islands. The "Western Radiation" includes five to seven historically recognized species distributed across almost the entire Archipelago. We combine dense geographic sampling and multilocus sequence data to estimate a phylogenetic hypothesis for the Western Radiation, to delimit species boundaries in this radiation, and to estimate a time frame for colonization events. Our phylogenetic hypothesis rejects two earlier topologies for the Western Radiation and paraphyly of M. albemarlensis , while providing strong support for single colonizations on each island. The colonization history implied by our phylogeny is consistent with general expectations of an east-to-west route predicted by the putative age of island groups, and prevailing ocean currents in the Archipelago. Additionally, combined evidence suggests that M. indefatigabilis from Santa Fe should be recognized as a full species. Finally, molecular divergence estimates suggest that the two colonization events likely occurred on the oldest existing islands, and the Western Radiation represents a recent radiation that, in most cases, has produced species that are considerably younger than the islands they inhabit.     

Ocean barriers and glaciation: evidence for explosive radiation of mitochondrial lineages in the Antarctic sea slug Doris kerguelenensis (Mollusca, Nudibranchia)

Strong currents and deep passages of water can be barriers for larval dispersal of continental marine animals, but potential effects on direct developers are under-investigated. We examined the genetic structure of Doris kerguelenensis , a directly developing sea slug that occurs across the Drake Passage, the body of water separating Antarctica from South America. We found deep mitochondrial divergences within populations on both sides of the Drake Passage, and South American animals formed multiple sister-group relationships with Antarctic animals. A generalised molecular clock suggested these trans-Drake pairs diverged during the Pliocene–Pleistocene, after the formation of the Drake Passage. Statistical parsimony methods recovered 29 separate haplotype networks (many sympatric) that likely correlate with allopatric events caused by repeated glacial cycles. Data from 16S were congruent but more conserved than COI, and the estimated ancestral 16S haplotype was widespread. The marked difference in the substitution rates between these two mitochondrial genes results in different estimates of connectivity. Demographic analyses on networks revealed some evidence for selection and expanding populations. Contrasting with the Northern Hemisphere, glaciation in Antarctica appears to have increased rather than reduced genetic diversity. This suggests orbitally forced range dynamics based on Northern Hemisphere phylogeography do not hold for Antarctica. The diverse lineages found in D. kerguelenensis point towards a recent, explosive radiation, likely reflecting multiple refuges during glaciation events, combined with limited subsequent dispersal. Whether recognised as cryptic species or not, genetic diversity in Antarctic marine invertebrates appears higher than expected from morphological analyses, and supports the Antarctic biodiversity pump phenomenon.     

MULTIPLE MODES OF SPECIATION INVOLVED IN THE PARALLEL EVOLUTION OF SYMPATRIC MORPHOTYPES OF LAKE WHITEFISH (COREGONUS CLUPEAFORMIS,SALMONIDAE)

We performed a phylogenetic analysis of mtDNA variation among seven sympatric pairs of dwarf and normal morphotypes of whitefish from northern Québec and the St. John River drainage to address three questions relevant to understanding their radiation. Are all sympatric pairs reproductively isolated? Do phylogenetic analyses confirm that sympatric whitefish morphotypes found in eastern North America represent the outcome of polyphyletic evolutionary events? If so, did all sympatric pairs from the St. John River drainage originate from the same scenario of allopatric divergence and secondary contact? The hypothesis of genetic differentiation was supported for all sympatric pairs from the St. John River drainage, whereas lack of mtDNA diversity precluded any test of reproductive isolation for northern Québec populations. Patterns of mtDNA variation confirmed that dwarf and normal morphotypes evolved in parallel among independent, yet closely related, lineages, thus providing indirect evidence for the role of natural selection in promoting phenotypic radiation in whitefish. Patterns of mtDNA diversity among sympatric pairs of the St. John River indicated a complex picture of whitefish evolution that implied sympatric divergence and multiple allopatric divergence/secondary contact events on a small geographic scale. These results suggests that ecological opportunities, namely trophic niche availability, may promote population divergence in whitefish.     

ENDOLITHIC MICROFLORA ARE MAJOR PRIMARY PRODUCERS IN DEAD CARBONATE SUBSTRATES OF HAWAIIAN CORAL REEFS1

To quantify the contribution of endolithic phototrophs to primary production of dead carbonate substrates, experimental blocks of cleaned Porites lobata Dana skeleton were placed at three different sites in Kaneohe Bay, Hawaii: inshore, lagoonal, and oceanic. After 6 months of exposure, experimental blocks were colonized by communities characteristic of their estuarine (inshore, lagoonal) and oceanic (ocean) environments. Blocks were sub-sampled; net photosynthesis (NP) and chl a concentrations of the whole blocks (epi- and endoliths) and scrapped blocks (only endoliths) were quantified. Green turf algae colonized predominantly inshore and lagoonal blocks, while encrusting corallines were the dominant epiliths colonizing oceanic blocks. Four main species of endolithic phototrophs were identified in all blocks: Mastigocoleus testarum Lagerheim, Plectonema terebrans Bornet and Flahault (cyanobacteria), Phaeophila dendroides Crouan and Crouan, and Ostreobium quekettii Bornet and Flahault (Chlorophytes). While epiliths were very different between sites, NP rates and chl a concentration of endoliths did not vary significantly and were positively correlated (191±25 mmol C·m⁻²·day⁻¹ and 590±150 mg chl a·m⁻² of reef, respectively). Assimilation numbers for whole communities, including both epilithic and endolithic communities, were similar to those measured for endolithic communities alone (average of 0.3 g C·g chl a·h⁻¹). Under experimental conditions, the contribution of endolithic phototrophs to community NP rates of blocks ranged from 56% to 81%, and under natural conditions, we estimated that this contribution ranged between 32% and 46%. Thus, we showed that the endolithic phototrophs are one of the major primary producers in dead coral substrates in a wide range of coral reef environments.     

Fleshy-fruitedness in the New Zealand flora

Aim It has been claimed that the New Zealand flora has an unusually high frequency of fleshy-fruitedness. This paper tests whether fleshy-fruitedness is indeed more common in New Zealand than in other temperate floras, then examines the distribution of fleshy-fruitedness among taxa and floristic elements to determine whether the flora conforms to predictions for a continental island with a relictual floristic element. Lastly, I test the extent to which fleshy-fruitedness has influenced colonization success and subsequent speciation within New Zealand. Methods Information on fruit characteristics for all indigenous seed plants was extracted from the Flora of New Zealand series and analysed with χ² tests. Results Contrary to previous claims fleshy-fruitedness was not unusually common in the New Zealand flora as a whole, when compared with other temperate floras. It is only more common in alpine communities and among trees. I also found no evidence for selective immigration; fleshy-fruited New Zealand genera were not more likely, than dry-fruited genera, to also occur in Australia. Furthermore there is no evidence that the New Zealand environment has favoured fleshy-fruited taxa; there has been no autochthonous evolution of fleshy-fruitedness in New Zealand, fleshy-fruitedness has had no significant effect on speciation within New Zealand, and endemic genera are no more likely to be fleshy-fruited than nonendemic genera. Fleshy-fruitedness in New Zealand is, however, strongly related to floristic elements of the flora. New Zealand is a continental island and therefore, theoretically, those elements of the flora dating from a time when the landmass was less isolated, should show a more balanced representation of dispersal modes. Contrary to this, fleshy-fruitedness is more common among species in Gondwanan taxa or in taxa with pollen records dating to before the Miocene. Main conclusions Fleshy-fruitedness in New Zealand conforms to neither the expectations for an isolated landmasses, namely a disharmonic range of dispersal modes, nor expectations for a continental island. I suggest that this pattern may be a product of selective survival of highly vagile taxa in the low-lying archipelago that was New Zealand during the late Cretaceous to mid-Cenozoic, followed by an invasion by taxa with a broader range of dispersal modes facilitated by the establishment of the circumpolar current.     

SPECIATION BY POLYPLOIDY IN TREEFROGS: MULTIPLE ORIGINS OF THE TETRAPLOID,HYLA VERSICOLOR

Speciation by polyploidy is rare in animals, yet, in vertebrates, there is a disproportionate concentration of polyploid species in anuran amphibians. Sequences from the cytochrome b gene of the mitochondrial DNA (mtDNA) were used to determine phylogenetic relationships among 37 populations of the diploid-tetraploid species pair of gray treefrogs, Hyla chrysoscelis and Hyla versicolor. The diploid species, H. chrysoscelis, consists of an eastern and a western lineage that have 2.3% sequence divergence between them. The tetraploid species, H. versicolor, had at least three separate, independent origins. Two of the tetraploid lineages are more closely related to one or the other of the diploid lineages (0.18%–1.4% sequence divergence) than they are to each other (1.9%–3.4% sequence divergence). The maternal ancestor of the third tetraploid lineage is unknown. The phylogenetic relationships between the two species and among lineages within each species support the hypothesis of multiple origins of the tetraploid lineages.     

NORTH PACIFIC HUMPBACK WHALE SONGS: A COMPARISON OF SOUTHEAST ALASKAN FEEDING GROUND SONGS WITH HAWAIIAN WINTERING GROUND SONGS

Humpback whales sing long, complex songs on their wintering grounds. On 25 August 1979 and 3 September 1981, we made recordings of humpback whale songs in southeastern Alaska, showing that humpback whales also sing on the summer feeding grounds. Both these Alaskan samples are songs in that they are repeating cyclical sound patterns and follow the known structure for humpback whale song. The Alaskan songs contain all the same material sung in the same order as that heard off Mexico and Hawaii in the surrounding wintering seasons. However, song, theme and some phrase durations are abbreviated in the Alaskan songs. The recording of these two songs represents the full sample of song recorded from 155 days over five years of attempting to record humpback whale song in Alaskan waters.