Tracing ancestry with chromosomal sequences

Most of the large Drosophila species of Hawaii are single-island endemics. Chromosomal sequences show that species at the new end of the archipelago have been derived stepwise from ancestral populations on older islands. The oldest high island has an endemic species with sequences that match some in the Nearctic-Palearctic robusta species group. Colonization from a continent by long-distance dispersal seems a likely origin for the Hawaiian drosophilids. Telmatogeton, a worldwide genus of marine midges, has five Hawaiian species inhabiting freshwater streams. Chromosomal sequences of a marine species in Hawaiian waters match the freshwater forms, indicating colonization from the ocean.     

Molecular phylogeny and divergence times of drosophilid species

The phylogenetic relationships and divergence times of 39 drosophilid species were studied by using the coding region of the Adh gene. Four genera--Scaptodrosophila, Zaprionus, Drosophila, and Scaptomyza (from Hawaii)--and three Drosophila subgenera--Drosophila, Engiscaptomyza, and Sophophora--were included. After conducting statistical analyses of the nucleotide sequences of the Adh, Adhr (Adh-related gene), and nuclear rRNA genes and a 905-bp segment of mitochondrial DNA, we used Scaptodrosophila as the outgroup. The phylogenetic tree obtained showed that the first major division of drosophilid species occurs between subgenus Sophophora (genus Drosophila) and the group including subgenera Drosophila and Engiscaptomyza plus the genera Zaprionus and Scaptomyza. Subgenus Sophophora is then divided into D. willistoni and the clade of D. obscura and D. melanogaster species groups. In the other major drosophilid group, Zaprionus first separates from the other species, and then D. immigrans leaves the remaining group of species. This remaining group then splits into the D. repleta group and the Hawaiian drosophilid cluster (Hawaiian Drosophila, Engiscaptomyza, and Scaptomyza). Engiscaptomyza and Scaptomyza are tightly clustered. Each of the D. repleta, D. obscura, and D. melanogaster groups is monophyletic. The splitting of subgenera Drosophila and Sophophora apparently occurred about 40 Mya, whereas the D. repleta group and the Hawaiian drosophilid cluster separated about 32 Mya. By contrast, the splitting of Engiscaptomyza and Scaptomyza occurred only about 11 Mya, suggesting that Scaptomyza experienced a rapid morphological evolution. The D. obscura and D. melanogaster groups apparently diverged about 25 Mya. Many of the D. repleta group species studied here have two functional Adh genes (Adh-1 and Adh-2), and these duplicated genes can be explained by two duplication events.      

Chromosomal sequences and interisland colonizations in hawaiian Drosophila

Of 103 picture-winged Drosophila species endemic to the high Hawaiian islands, all but three are endemic to single islands or island complexes. They are presumed to have evolved in situ on each island. The banding pattern sequences of the five major polytene chromosomes of these species have been mapped to a single set of Standard sequences. Sequential variation among these chromosomes is due to 213 paracentric inversions. An atlas of their break points is provided. Geographical, morphological and behavioral data may be used to supplement the cytological information in tracing ancestry. Starting at the newer end of the archipelago, the 26 species of the Island of Hawaii (less than 700,000 years old) are inferred to have been derived from 19 founders, 15 from the Maui complex, three from Oahu and one from Kauai. The existence of 40 Maui complex species is explicable as resulting from 12 founders, ten from Oahu and two from Kauai. The 29 Oahu species can be explained by 12 founder events, five from Kauai and seven from Maui complex (summary in Figure 5). Although the ancestry of two Kauai species can be traced to newer islands, the ten remaining ones on this island (age about 5.6 million years) are apparently ancient elements in the fauna, relating ultimately to Palearctic continental sources.     

Hawaiian biogeography and the islands' freshwater fish fauna

Aim This paper describes known patterns in the distributions and relationships of Hawaiian freshwater fishes, and compares these patterns with those exhibited by Hawaii's terrestrial biota. Location The study is based in Hawaii, and seeks patterns across the tropical and subtropical Indo‐west Pacific. Methods The study is based primarily on literature analysis. Results The Hawaiian freshwater fish fauna comprises five species of goby in five different genera (Gobiidae). Four species are Hawaiian endemics, the fifth shared with islands in the western tropical Pacific Ocean. All genera are represented widely across the Indo‐west Pacific. All five species are present on all of the major Hawaiian islands. All five species are amphidromous – their larval and early juvenile life being spent in the sea. Although there has been some local phyletic evolution to produce Hawaiian endemics, there has been no local radiation to produce single‐island endemics across the archipelago. Nor is there evidence for genetic structuring among populations in the various islands. Main conclusions In this regard, the freshwater fish fauna of Hawaii differs from the well‐known patterns of local evolution and radiation in Hawaiian Island terrestrial taxa. Amphidromy probably explains the biogeographical idiosyncrasies of the fish fauna – dispersal through the sea initially brought the fish species to Hawaii, and gene flow among populations, across the archipelago, has hitherto inhibited the evolution of local island endemics, apparently even retarding genetic structuring on individual islands.     

Population variation and hybridization: Comparison of finches from two archipelagos

Summary Some populations of Darwin's Finches (Emberizinae) are exceptionally variable in body size and beak traits as a result of introgressive hybridization. A study of museum specimens of honeycreeper-finches (Carduelinae) from the Hawaiian islands was undertaken to see if the same phenomenon was manifested by a different phyletic group of finches in a different archipelago. Five hundred and twenty-four specimens of the seven species with finch-like bills were measured and their coefficients of variation were compared with those of the ground finch group (six species) of Darwin's Finches. Coefficients were smaller in the Hawaiian finches. Sympatric and, hence, potentially hybridizing species on the island of Hawaii were not consistently more variable than the allopatric species on other islands in the archipelago. The one species with both sympatric and allopatric populations did not show greater variation in the sympatric population. There is little evidence from these comparisons of hybridization occurring in the last 100 years. The difference between the two finch faunas can be explained in terms of two factors. Finches have been present for a longer time in the Hawaiian archipelago than in the Galápagos archipelago and have had more time to not only diversify but to evolve pre- and post-zygotic isolating mechanisms. In the generally less seasonal and floristically richer Hawaiian islands they have evolved greater dietary specializations. Beak traits adapted to specialist feeding may have been under stronger stabilizing selection and hybrids (if formed) may have been at a strong disadvantage in the absence of an ecological niche intermediate between the niches of the two parental species. Results of published electrophoretic studies of genetic variation suggest that the early phase of differentiation, involving occasional introgressive hybridization, may last for up to 5 million years.     

Paraliparis hawaiiensis,a new species of snailfish (Scorpaeniformes: Liparidae) and the first described from the Hawaiian Archipelago

Paraliparis hawaiiensis n.sp. is described from the north‐western Hawaiian Islands from two specimens collected at 2196 and 3055 m. It differs from other North Pacific Ocean species in its chin pore arrangement, tooth pattern and body proportions. Although liparid specimens have previously been collected from Hawaii, they were undescribed and are now lost. Therefore, this is the first liparid species described from the archipelago. In situ photographs of Hawaiian snailfishes are also shown and discussed here.     

Phylogeny and age of diversification of the planitibia species group of the Hawaiian Drosophila

The Hawaiian Drosophila offer a unique opportunity to examine evolutionary questions because of the known ages of the Hawaiian Islands and the large number of species endemic to this archipelago. One of the more well studied groups of Hawaiian Drosophila is the planitibia species group, a long-standing population genetic model system. Here we present a molecular phylogenetic hypothesis of all 17 taxa in the planitibia group based on nucleotide sequences from two mitochondrial (16S and COII) and four nuclear (Adh, Gpdh, Yp1, and Yp2) loci, accounting for over 4kb of sequence per taxon. We use these data to estimate major divergence times within this group. Our results suggest that the basal diversification within this group, calculated at 6.1 +/- 0.47 MY, predates the oldest high island of Kauai. The older diversifications in this group took place on Kauai, with subsequent colonization and speciation events occurring as new islands became available to Drosophila. Understanding of the phylogenetic relationships of this important group will place the existing population genetic work in a macroevolutionary context and stimulate additional work, particularly on those taxa endemic to the Maui Nui complex of islands.     

Molecular insights into the phylogenetic structure of the spider genus Theridion (Araneae, Theridiidae) and the origin of the Hawaiian Theridion-like fauna

The Hawaiian happy face spider ( Theridion grallator Simon, 1900), named for a remarkable abdominal colour pattern resembling a smiling face, has served as a model organism for understanding the generation of genetic diversity. Theridion grallator is one of 11 endemic Hawaiian species of the genus reported to date. Asserting the origin of island endemics informs on the evolutionary context of diversification, and how diversity has arisen on the islands. Studies on the genus Theridion in Hawaii, as elsewhere, have long been hampered by its large size (> 600 species) and poor definition. Here we report results of phylogenetic analyses based on DNA sequences of five genes conducted on five diverse species of Hawaiian Theridion , along with the most intensive sampling of Theridiinae analysed to date. Results indicate that the Hawaiian Islands were colonised by two independent Theridiinae lineages, one of which originated in the Americas. Both lineages have undergone local diversification in the archipelago and have convergently evolved similar bizarre morphs. Our findings confirm para- or polyphyletic status of the largest Theridiinae genera: Theridion , Achaearanea and Chrysso . Convergent simplification of the palpal organ has occurred in the Hawaiian Islands and in two continental lineages. The results confirm the convergent evolution of social behaviour and web structure, both already documented within the Theridiidae. Greater understanding of phylogenetic relationships within the Theridiinae is key to understanding of behavioural and morphological evolution in this highly diverse group.     

Out of Hawaii: the origin and biogeography of the genus Scaptomyza (Diptera: Drosophilidae)

The Hawaiian Archipelago is the most isolated island system on the planet and has been the subject of evolutionary research for over a century. The largest radiation of species in Hawaii is the Hawaiian Drosophilidae, a group of approximately 1000 species. Dispersal to isolated island systems like Hawaii is rare and the resultant flora and fauna shows high disharmony with mainland communities. The possibility that some lineages may have originated in Hawaii and subsequently 'escaped' to diversify on continental landmasses is expected to be rarer still. We present phylogenetic analysis of 134 partially sequenced mitochondrial genomes of Drosophilidae (approx. 1.3 Mb of sequence total) to address major aspects of adaptive radiation and dispersal in Hawaii. We show that the genus Scaptomyza, a group that accounts for approximately one-third of the species-level diversity of Drosophilidae in the Hawaiian Islands, originated in Hawaii, diversified there, and subsequently colonized a number of island and continental landmasses elsewhere on the globe. We propose that a combination of small body size, rapid generation time and unique ecological and physiological adaptations have allowed this genus to effectively disperse and diversify.     

Complex CIS-Acting Regulators and Locus Structure of Drosophila Tissue-Specific Adh Variants

Diverse patterns of tissue-specific expression of alcohol dehydrogenase (ADH) among species of the grimshawi subgroup of Hawaiian picture-winged Drosophila suggests control by complex or multiple, independently acting regulatory elements. These elements act by controlling Adh mRNA accumulation in individual tissue types. Restriction mapping of the Adh loci from these species reveals several insertion/deletion differences, one of which lies just outside the 5' end of the structural sequences and correlates with differences in larval patterns of ADH expression. No tissue-specific rearrangement of Adh sequences was observed.     

Using taxonomic and phylogenetic evenness to compare diversification in two Island Floras

This study compares the phylogenetic structure in the Canary Islands and Hawaii by means of the distributions of the species number for plant families (Taxonomic evenness) and lineages (Phylogenetic evenness) across archipelagos and across habitats in both archipelagos using the Gini coefficient. We then investigate phylogenies to identify particular habitats contributing to such differences using Taxonomic distinctness (AvTD) and its variation (VarTD).Our results show that the distribution of species number among Hawaiian lineages is much more uneven than the Canary Islands. In contrast, Hawaii produces a more even distribution of species number by family than the Canary Islands. This may be due to the Hawaiian Flora being derived from considerably fewer colonists than the Canarian Flora as a result of its much greater degree of isolation. At the same time, Hawaii is represented by the same number of families as the Canary Islands. This may stem from Hawaii's flora being derived from a greater range of source areas despite its isolation. Finally, there is much more diversification spread across a larger number of lineages in Hawaii. The higher degree of Hawaiian diversification may be due to a greater range of habitats, more diverse and phylogenetically distinct floristic sources, and low initial species diversity resulting from extreme isolation.Two Canarian habitats (Rock communities and Thermophilous habitats) and one Hawaiian habitat (Wet communities) contribute to the differences in phylogenetic structure between the two archipelagos. These habitats exhibit disproportionate levels of unevenness and may represent centres of diversification. We propose a combination of two habitat properties, high receptivity and low stability, to explain these results.     

Redundant cis-acting elements control expression of the Drosophila affinidisjuncta Adh gene in the larval fat body.

The alcohol dehydrogenase (Adh) gene in the Hawaiian species of fruit fly, Drosophila affinidisjuncta, like the Adh genes from all Drosophila species analyzed, is expressed at high levels in the larval fat body via a larval-specific promoter. To identify the cis-acting elements involved in this highly conserved aspect of Adh gene expression, deleted D. affinidisjuncta genes were introduced into D. melanogaster by somatic transformation. Unlike previously described methods, this transformation system allows analysis of Adh gene expression specifically in the larval fat body. The arrangement of sequences influencing expression of the proximal promoter of this gene in the larval fat body differs markedly from that described for the Adh gene from the distant relative, D. melanogaster. Multiple redundant elements dispersed 5' and 3' to the gene, only some of which map to regions carrying evolutionarily conserved sequences, affect expression in the fat body. D. affinidisjuncta employs a novel mode of Adh gene regulation in which the proximal promoter is influenced by sequences having roles in expression of the distal promoter. This gene is also unique in that far upstream sequences can compensate for loss of sequences within 200 bp of the proximal RNA start site. Furthermore, expression is influenced in an unusual, context-dependent manner by a naturally-occurring 3' duplication of the proximal promoter--a feature found only in Hawaiian species.     

Ecology, biogeography and evolution of corals in Hawaii

Recent studies in Hawaii have contributed much to the understanding of the ecology and evolution of Hawaiian corals and are forcing a reevaluation of our basic concepts concerning the zoogeography, ecology, taxonomy and population biology of these important reef-forming organisms. Geographic isolation rather than physical adversity of the environment seems largely to have determined the number of coral species that are found in Hawaii, but the physical environment controls growth of Hawaiian species with increasing latitude along the archipelago. Annual broadcast spawning has recently been shown to be the dominant mode of sexual reproduction, rather than brooding of larvae on a lunar cycle as previously believed. Asexual reproduction through colony fragmentation or by production of asexually produced larvae is now known to result in extensive representation of a single genotype in some coral populations.     

MULTIPLE COLONIZATIONS OF ASPLENIUM ADIANTUM-NIGRUM ONTO THE HAWAIIAN ARCHIPELAGO

The extreme isolation and mid-Pacific origin of the Hawaiian archipelago has ensured that all indigenous organisms have arrived via long-distance dispersal or have evolved from successfully colonizing species. Although this isolation has also produced high rates of species endemism in angiosperms (89% or more), that rate in pteridophytes is considerably less (76%). The ratio of native species to the estimated number of original successful colonizing species in angiosperms (3.4) is more than double that for pteridophytes (1.6). One possible explanation for the lower speciation rate in pteridophytes is that populations of these species are more likely to experience interpopulational gene flow because of the great vagility of their wind-dispersed spores. We conducted isozymic surveys of populations from the island of Hawaii of the indigenous allotetraploid species Asplenium adiantum-nigrum, putatively derived from two strictly European diploid taxa. Our data support multiple hybrid origins for the populations surveyed, with a minimum of 3, and possibly as many as 17, discrete hybridization events having produced the genetic diversity observed. Since the parental taxa are not found in Hawaii, each hybrid lineage must have arrived in the archipelago independently of the others. Similar long-distance, repeated dispersal events may be occurring between insular and noninsular populations of other native pteridophytes in Hawaii and in other insular regions of the world, thus contributing to the relatively low rates of speciation and insular endemism in this ancient group of plants.     

Rates of DNA change and phylogeny from the DNA sequences of the alcohol dehydrogenase gene for five closely related species of Hawaiian Drosophila

The sequence of 1.6 kb of DNA surrounding the alcohol dehydrogenase (Adh) gene from five species of the Planitibia subgroup of the Hawaiian picture-winged Drosophila, with estimated divergence times of 0.4-5.1 Myr, has been determined. The gene trees which were found by using the sequence divergence from different regions of the sequences are generally in accord with the phylogeny proposed for these species when chromosomal inversions and island of origin are used. One of the species (D. picticornis) appears to be more distant from the other species in this group than they are from a member of the Grimshawi group (D. affinidisjuncta) which is chromosomally more distant. Two of the species (D. differens and D. plantibia) show heterogeneity in the nucleotide changes in the Adh coding region, heterogeneity which is interpreted to be due to a gene conversion or recombination after hybridization between the two species. The minimal rate of nucleotide substitution of synonymous nucleotides and of nontranscribed nucleotides downstream from the coding region is estimated as 1.5 x 10(-8) and 1.1 x 10(-8) substitutions/nucleotide/year, respectively. This rate is two to three times the maximal rate estimated for mammalian synonymous substitutions.     

Molecular phylogeny of butterflies Parnassius glacialis and P. stubbendorfii at various localities in East Asia.

The phylogeny of butterflies, Parnassius stubbendorfii and P. glacialis, collected at various localities in the Japan archipelago and the eastern part of the Asian continent was analyzed using mitochondrial DNA sequences coding for NADH dehydrogenase subunit 5 (805 bp). The molecular phylogenetic trees revealed that P. glacialis and P. stubbendorfii diverged from a common ancestor, and then the populations inhabiting the Japan archipelago and the Asian continent diverged in each species. The reliability of these divergences was supported by high bootstrap values. The divergences within the Japan archipelago and within the Asian continent in each species were unclear because of low bootstrap values. The genetic distance and a rough time-estimation in the UPGMA tree suggest that the both populations of P. glacialis and P. stubbendorfii may have been isolated in the Japan archipelago at the early time (about 1.7-2.0 Mya) of the glacial period in the Pleistocene. The genetic distance between the Japanese and the continental subspecies may be large enough that they can be classified as different species, in comparison with the genetic distances among some other parnassian species.     

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.     

Tissue-Specific Expression Phenotypes of Hawaiian Drosophila Adh Genes in Drosophila Melanogaster Transformants

Interspecific differences in the tissue-specific patterns of expression displayed by the alcohol dehydrogenase (Adh) genes within the Hawaiian picture-winged Drosophila represent a rich source of evolutionary variation in gene regulation. Study of the cis-acting elements responsible for regulatory differences between Adh genes from various species is greatly facilitated by analyzing the behavior of the different Adh genes in a homogeneous background. Accordingly, the Adh gene from Drosophila grimshawi was introduced into the germ line of Drosophila melanogaster by means of P element-mediated transformation, and transformants carrying this gene were compared to transformants carrying the Adh genes from Drosophila affinidisjuncta and Drosophila hawaiiensis. The results indicate that the D. affinidisjuncta and D. grimshawi genes have relatively higher levels of expression and broader tissue distribution of expression than the D. hawaiiensis gene in larvae. All three genes are expressed at similar overall levels in adults, with differences in tissue distribution of enzyme activity corresponding to the pattern in the donor species. However, certain systematic differences between Adh gene expression in transformants and in the Hawaiian Drosophila are noted along with tissue-specific position effects in some cases. The implications of these findings for the understanding of evolved regulatory variation are discussed.     

Origin of the endemic fern genus Diellia coincides with the renewal of Hawaiian terrestrial life in the Miocene

The enigmatic fern genus Diellia, endemic to the Hawaiian archipelago, consists of five extant and one recently extinct species. Diellia is morphologically highly variable, and a unique combination of characters has led to several contrasting hypotheses regarding the relationship of Diellia to other ferns. A phylogenetic analysis of four chloroplast loci places Diellia within 'black-stemmed' rock spleenworts of the species-rich genus Asplenium, as previously suggested by W. H. Wagner. Using an external calibration point, we estimate the divergence of the Diellia lineage from its nearest relatives to have occurred at ca. 24.3 Myr ago matching an independent estimate for the renewal of Hawaiian terrestrial life (ca. 23 Myr ago). We therefore suggest that the ancestor of the Diellia lineage may have been among the first successful colonists of the newly emerging islands in the archipelago. Disparity between morphological and nucleotide sequence variation within Diellia is consistent with a recent rapid radiation. Our estimated time of the Diellia radiation (ca. 2 Myr ago) is younger than the oldest island of Kaua'i (ca. 5.1 Myr ago) but older than the younger major islands of Maui (ca. 1.3 Myr ago), Lana'i (ca. 1.3 Myr ago) and Hawaii (ca. 0.43 Myr ago).     

Seasonal surveillance confirms the range expansion of Aedes japonicus japonicus (Theobald) (Diptera: Culicidae) to the Hawaiian Islands of Oahu and Kauai

The Asian bush mosquito, Aedes japonicus japonicus (Theobald) was not known to occur in the Hawaii archipelago until it was identified on the island of Hawaii in 2003. This mosquito species remained undetected on the neighboring islands for 8?years before it was discovered at the Honolulu International Airport on Oahu in 2012. By 2015, four Ae. j. japonicus mosquitoes were collected in the western mountains of Oahu and one was collected in the central mountains of Kauai. The collection of this invasive mosquito species across the neighboring Hawaiian Islands of Oahu and Kauai indicated the need for increased seasonal surveillance on these islands. Following nearly four years of surveillance, Ae. j. japonicus was also confirmed to occur in the eastern mountains of Oahu and in the central mountainous region of Kauai. To expand the knowledge of the spread of invasive mosquitoes species further surveillance is necessary to identify all possible areas where populations of Ae. j. japonicus and other invasive mosquito species occur in Hawaiian archipelago.