What is the oldest Hawaiian Island

Most emanations within the arcipelago adhere to a certain extent to a progressive control scheme, since they colonise in sequence from older isles, although the spotting often also takes place within isles. Among the multi-island types investigated, however, we find that there are a number of highly diversified samples that reflect an increased dispersion spectrum (succineids, dicranomyia). It detects Hawaiian specimens that colonise other areas (scaptomyza fly, succineids).

The taxonomy was also clarified by means of molecule experiments (agatinella slugs, drosophilia). Whilst Hawaiian wildlife has blossomed since the mid-1990s, much is still not known. But Hawaii's wildlife is in danger: more than 70 percent of Hawaii's bird population and possibly 90 percent of its snail population are dead.

Preservation is essential if this species is to be able to bring further insight into deep evolutions and biogeographical issues. Hawaii's Hawaiian Arcipelago (Figure 1) is a series of sea islets that move northwest over a fixed flag or "hot spot" in the Earth's surface that intermittently transmits magma across the surface, forming a string of volcanos, each younger than the previous one and northwest of the first.

At some point each island sinks and dies, becomes a low altitude tunnel, then a submerges sea mount and is ultimately subduced as the Pacific Ocean slab glides under the adjoining oceanic slab (Price & Clague 2002). Today's archipelago is made up of the younger "high" and the older north-western isles, which have become low atlls or small Erosted isles and battlements.

Kure Atoll (29 Ma) is the oldest island in the northwest and Kauai (5. 1 Ma) is the oldest high island, with the youngest island, Hawaii itself, being less than 0.5 Ma and still being made ((Figure 1). The report concentrates on the high islets that host spectral array of native plant and animal life (Ziegler 2002).

Hawaiian Islands Wall Chart. Above right illustration shows the location of the Hawaiian Islands in the Pacific Area. Bottom lefthand picture shows the whole Hawaiian arcipelago, which includes the Hawaiian islands in the northwest, with the Kure Atoll (29 Ma) in the extreme northwest edge. Kauai's yardstick relates to the center of Hawaii's major islands.

Hawaii's bio-diversity and evolving charms are probably more dramatic than those of the Galápagos, and one can only guess what Darwin's joy and reverence would have been if he had been to Hawaii. The Reverend John Thomas Gulick, Hawaii 1832, was intrigued by this wonderful diversity of species, especially the colourful and extremely varied Oahu slugs.

On the basis of his study of Hawaiian slugs, Gulick's work Evolution, racially and habitudinally (Gulick 1905), and some of his other papers may contain the first account of the founding effect, the initial description of genetics and of the idea of the shift of equilibrium (Carson 1987a). Hawaiian biologic radiation has been regarded since his day as some of the most important samples of evolutional experimentation (Simon 1987), in which the reproduction of clade on sequential and accurately datable isles has provided important insight into many facets of evolutional ecology, among them species (Carson 1987b), bigeography (Wagner & Funk 1995; Cowie & Holland 2006), sex-selections ( (Kaneshiro 1988, 2006; Mendelson & Shaw 2005) and eco-system dynamics (Vitousek 2002, 2004).

The emergence and increasing spread of the use of phylogeographical techniques has made it possible to address aspects of historic bio-geography and systematic such as geographical origins, monophile and clad bi-furcation, time and order. Research on the clarification of historic biogeographical models in the Hawaiian Islands is beginning to uncover some shared pattern.

The most remarkable in this context is perhaps the turning point collection by Wagner & Funk (1995), which deals with the bio-geography of a series of Hawaiian flora and fauna tataxi. There were some general designs, among them the holding of many Hawaiian lines to an island variation of the so-called "progression rule" (Hennig 1966), which identified centers of provenance with the youngest members of a monophylet ic line on the geographical outskirts, in this case the geological younger isles.

Thus, after the first settlement of the oldest island in a sequence of producing islets, a line fork accompanies the spread from older to younger ones along the series. When Wagner & Funk (1995) was compiled, it was only when biomolecular methods became routinely applied to biogeographical issues, and most of the trials in this book were primarily or completely morphologic.

In this brief retrospective, the bio-geography of Hawaiian landlife is to be taken up and updated in the wake of the large number of recent molecule surveys carried out since Wagner & Funk's groundbreaking paper (1995). Few trials can be examined in detail in the effort to be as complete as possible in the available area.

In the following, however, a large part of the work done since Wagner & Funk (1995) is summarized, which allows us to gain an insight into the large and dispersed books and to reflect on possible trends and problems. The majority of early molecule trials used a simple, mostly mesochondrial genetic markers (mtDNA). The majority of Hawaiian land based taxi trials have included non-human animals, undoubtedly because of the relatively easy sampling of a large number of specimens, generally fewer regulatory constraints and far greater numbers of mammals.

However, there are a number of interesting research on the origin and distribution of bird life, which includes fossilized taxes, and the individual indigenous mammals, a bats (there are no indigenous Hawaiian reptile or amphibian land -based animals). Previous publications dealing with the hawaiian fauna's origin in phylogenetics and biogeography often implicitly believed that every main group of Hawaiian taisa, e.g. all Hawaiian members of a particular familiy, was monophyletal and descended from a unique colonising group.

Whereas this seems sensible for the large groups native to the isles, such as the Amastridae Landsnail familiy or the Hawaiian honey tugs, the Drepanidinae, there is less reason for such an a priori adoption in a group that is not native to the isles, e.g. the Succineidae Landsnail familiy or the Drosophila aviary.

The monophyllic and thus assumed uniform origins of Hawaiian taxpayers of a more common group were affirmed for some groups. There are about 60 Hawaiian bee varieties of the global Hylaeus genera, all of which belong to the subgenera Nesoprosopis, which is otherwise mainly known from Japan, with one variety stretching west to Europe and a number of indescribable varieties in China (Magnacca & Danforth 2006).

All Hawaiian rays were detected as monophyletically using the COI, CO2 and trRNA-leucine markers of MontDNA, in combination with morphologic characteristics - the results of a unique colonisation. Under the assumption that the subgenerative classification is accurate, Hawaiian rays are likely of East Asiatic origins. The Hawaiian Havaika spiders (nine nominally occurring species), also found on the Marquesas Islands of the Southern Pacific (Gillespie et al. 2008), showed the monophilia of Hawaiian rays but did not specifically conclude their geographical origins (Arnedo & Gillespie 2006).

However, there is increasing indication of multi-populations for certain Hawaiian endemics belonging to wider, more widespread groups. In Gillespie et al (1994), with a small fraction of the 12S RNA gene, several sources of Hawaiian tetragnathide hairs were found, with two distinct colonisations resulting in independently irradiated genera and two extra colonisations that did not result in significant radiation.

Though it is a somewhat restricted overall sample collection using only mtDNA (COI and 16S) marker, Gillespie (2002) proposed that Hawaiian tetragnatha originate from an US well. Arnedo et al. (2007) in another group of Hawaiian cobwebs of the cosmopolitical species Theridion analyzed sequences of two mtDNA (COI and 16S) and three mDNA ( (18S, 28S and Histon H3) mDNAs.

Rundell et al (2004) used COI to show two different monophylet ic lines and showed at least two sources of Succineidae, a worldwide spread of the Hawaiian island snails: an older colonisation, first the island of Kauai, which is the oldest of Hawaii's major islands (Figure 1), and a younger colonisation of Hawaii, the youngest island.

Roundll et al. (2004) conjecture that this is East Asiatic source only. The Netherlands & Cowie (in the press) confirmed these results in a multi-locus survey with extended Hawaiian and world samples and proposed a South Pacific ancestry for the older Hawaiian group. There is less clarity about the roots of the well-known agatinellide terrestrial slugs, an almost Pacific indigenous species, which include the species of the Hawaiian Achatinellinae (subfamily).

It is a monophylet lineage, derived from the geographic samplings carried out to date (Holland & Hadfield 2004; Wade et al. 2006), and the individual strain taken outside Hawaii is associated with Hawaiian monophylet rays (Holland & Hadfield 2004), which suggests a unique source of Hawaiian rays.

The absence of non-Hawaiian agatinellid species in the Hawaiian genus (Cooke & Kondo 1961) indicates, however, that there may have been several Hawaiian colonisations from elsewhere in the Pacific. Agatinellidae are part of a well-funded group of' orthurethran' taxa, including the Hawaiian indigenous Amastridae genealogy, the New Caledonian indigenous Draparnaudiidae genealogy, the Pacific Island (excluding Hawaii), the indigenous Partulidae genealogy and other global distributive genealogies (Holland & Hadfield 2004; Wade et al. 2006).

However relations within the Orthurethra are not well known ( (Wade et al. 2006) and the final geographical origins of agateinellidae (and the other Pacific island taxis) remain unclear, although in Europe and North America were found to have a dubious agatinellide relationship (Solem 1976; Solem & Yochelson 1979).

Other Hawaiian groups of invertebrates that have been analyzed by genetic analysis have sometimes been able to deduce the number of colonisations independently, although, as with agateinellidae, the exclusion of a wider geographical sample of groups that are not non-harmful to Hawaii rules out fixed inferrals. While Hawaiian craneflies (Diptera, Limoniidae, Dicranomyia) can be monophyletal, indicating a unique colonisation, although with a possible second colonisation by one colonisation type (Nitta & O'Grady 2008), the nonexistence of samples of Dicranomyia from elsewhere in its widespread Pacifica dispersal range does not allow final reaffirmation.

To clarify the Hawaiian dragonfly diversity pattern and place it in a wider geographical contexts, Jordan et al. (2003) used nanodNA and mtDNA from 20 of the 23 described types of the indigenous Megalagrion family. There were eight types (in two genera), and although Megalagrion was clearly monophylet, not a single type or type appeared as a Hawaiian one.

Like many Hawaiian groups, the out groups were far away; the out group selection affected the in-group overhead, but the issue could be solved by adapting an Evolution to the in-group overhead and limiting the in-group overhead during out-grouping. Although there are two well endorsed monophylet groups of Hawaiian tettigoniids or catydides (Banza spp.), the placing of a unique specie (Banza nihoa) is ambiguous and in combination with the absence of samples of related specimens from the possible areas of origin (Banza seems to be intimately related to the New and Old World taxa) excludes the number of Hawaiian colonisations (Shapiro et al. 2006).

Mendelson & Shaw (2005), on the basis of an AFLP (amplified fragment length polymorphism) of 25 types of Hawaiian cricket endemics (Laupala), have proposed that the genera are monophyletal, although they have not tried far beyond Hawaii and have therefore not addressed their geographical ancestry. In spite of the relatively large amount of research on Hawaiian drosophiles, and although they were progressively acknowledged as originating from a singular colonisation (O'Grady 2002; Remsen & O'Grady 2002; O'Grady & DeSalle 2008), their geographical origins were unclear, although the opportunity to mention Eastern Asia existed from times to times (Remsen & DeSalle 1998; Davis 2000).

Hawaiian mammals, although far less varied than terrestrial gastropods or terrestrial gastropods, have nevertheless triggered a number of genetic engineering trials on their origin, although much is still not known. Butischer & MacIntosh (2001) studied bird molecules, mostly using the use of metDNA biomarkers, largely from the point of view of how they understand their phosphogenetic relations and origin.

Only other indigenous vertebrates are bats. Hawaiian honey veils (Drepanidinae) with over 50 different bird varieties are the most visible. Others have emitted far less spectral images and since over 70 percent of all avian fauna is now endangered (Boyer 2008), the challenge of molecule analysis is a challenge. However, a combined analysis of existing taxa and significant achievements in recovering bone sequence from dead specimens has provided significant insights into the physiogenetic and geographical origin of the various groups and non-radiating specimens.

Hawaiian air-impaired tracks, Porzana spp (perhaps more than 12 species), may have originated from two separate colonisations from Asia or the West Pacific by volanteas ("Slikas et al. 2002"). Hawaii's mockingbird, Myadestes obscurus, the only surviving member of five types of ray, originates from the New World (Caribbean or the west of North America).

Hawaiian crows, Corvus hairawaiiensis, the only one of four raven types, seems to be more close related to the Pacific Island ravenses than to other Pacific Islandraven. They are however related to the New World, perhaps to South America's Anas or Anas relations, although they separated from them a long while ago, before the island of Hawaii, from which they are away, emerged and became airless and giant (see also Sorenson et al. 1999).

The existing geese, the Hawaiian canard, Anas vyvilliana, and the Laysan canard, Anas LAYSANENENSIS, appear to be colonies in their own right, however, with the former most likely being related to Northern mallard or spotted Ducks and the latter to the South Pacific teal. Eudocimus and Eudocimus are the two most related to the New World' s Eudocimus alb.

Hawaiian shelduck ( "Branta sandvicensis"), together with fossilized rays of Hawaiian branchta is embedded in the widely spread Nearctic Canada shelduck ( "Branta canadensis"), which makes the latter para-phyletic, but confirms the northern US origins of Hawaiian brancha (see also Paxinos et al. 2002). Hawaiian falcon, Buteo soloitarius, is grouped with the New World and not the Old World types.

Unlike most of these species, which originate in the New World or in some cases in the Southern Ocean, the Hawaiian Adler, a relatively young but pre-human species arriving on the island, is of Palearctic descent and has differed little from the sea elder Haliaeetus albicilla (see also Fleischer et al. 2000).

The use of molecule approach has not been used to investigate the origin of other Hawaiian birds whose understanding is due to morphologic taxonomies. Hawaii's only indigenous land animal, the Hawaiian bats, Lasiurus conereus semiotus, has shown that the mtDNA restricted site map is more intimately related to the species of the bats in Northern America than to the species of the bats in Southern America (Morales & Bickham 1995), indicating a relatively new colonisation of the Hawaiian islands from Northern America.

Whereas few rather few molecular investigations have dealt with the origin of Hawaiian wildlife in an explicit and satisfactory manner, perhaps with the exceptions of the bird, a number of research projects have dealt with the evolutive emanations and the pattern of diversity within the island chains after first colonisation. Most of these designs more or less closely follow the progressive rules, although there may be resettlement from younger to older isles, larger emanations within and not between isles, and significant random spreads, especially in more vaguely or active scattering species.

The results of Thacker & Hadfield (2000) with 16S and Holland & Hadfield (2004) with COI dealt with the diversity of the agateinelline linear slugs, whereby the results between the two trials were substantially constant. Philogenetic reconstruction proposed colonizing Oahu as the first island, with ensuing settlement and diversity, which generally followed the principle of progressive settlement, but with a number of cases of back settlement (Figure 2).

The mysterious find (Gage 1996) of a solitary fossilized kind of the agateinellin type Newcombia on Kauai (no other agateinellins are known from Kauai and Newcombia is otherwise only known from Maui and Molokai) probably is a recolonisation from Maui/Molokai, the skip from Oahu (Figure 2), instead of a Kauai source of agatinellin rays, since Newcombia is a more derivative than basic taxion within the agatinellinae (Holland & Hadfield 2004).

Intra-island colonisation route for the agateinelline gastropods from the Holland & Hadfield MDNA fylogeny (2004). They are the knots and blades in the trees, except for the one from Molokai to Kauai, which is the only agatinelline on the island, representing the sub-fossil Newcombia sp. on Kauai.

The numbers on the isles are the numbers of indigenous agatinelline types on each island (Cowie et al. 1995). In the two cochlear colonizations by Rundell et al. (2004) and Holland & Cowie (in the press) with succinide terrestrial spiral irradiations using radioactive nuclei the older irradiation followed approximately the progressive regulation, with the first colonisation site being Kauai.

The recent settlement of Hawaii, the youngest island, has so far only led to the presence of succinea on the island, with the sole exceptions (Holland & Cowie 2007) of Succinea comaduca, which is found on all the major Hawaiian islands and is a derivative of this type of Hawaiian ray (Figure 3, see below).

Interisland colonisation pattern for Hawaiian native Hawaiian succin gastropods from human genetics information (Holland & Cowie in the press). The numbers on the isles are the numbers of the indigenous succinide types on each island (Cowie et al. 1995). Each of the two arrowheads, one of each of the two lines pointing south, one from Hawaii called "Tahiti", the other from Kauai called "Samoa", represents a South Pacific type, indicating a historic settlement of these places from the Hawaiian isles.

Shadowed squares emphasize the two well-supported Hawaiian lines, each of which retains its integrality in overall phylogeny, with several lines between them. Succinea is the only indigenous succinea of the B archipelago found on non-Hawaiian isles ("Holland & Cowie 2007").

Hylaeus bee irradiation (Magnacca & Danforth 2006) also appears to be due to early settlement on the youngest island of Hawaii, followed in this case by spread and spotting northwest of the entire island group. Most of the types were found between isles ( "sister species" that occur on different islands) and not within isles ( "sister species" that occur on the same island), with the exemption of the large rays on the island of Hawaii, the site of the island's primordial population.

In Jordan et al. (2003), various methyl DNA biomarkers and the EF1? atomic nucleus were analyzed in 20 of the 23 genera of the Hawaiian Megalagrion type end-emic monophylet seed. It is not a uniform spectrum, but a series of well established subklades with different pattern and levels of sophistication.

A group may have colonised in the opposite sense, from the youngest to the older isles. Every group forms approximately one group of types with similar ecologies (breeding habitat). Within Kauai, however, a group has spread out to cover every available hatchery. Tetragnatha clads have a pattern among the spider which approximately corresponds to the progressive rules, since the most basic specimens are found on Kauai and Oahu, with more derivative specimens on younger isles ((Gillespie et al. 1997; Pons & Gillespie 2004).

The Tetragnatha blade of the "prickly legs", however, is the most related of all the types on a particular island, although there is a general trend towards the phaeolith. That is in contrast to an earlier morphologic study (Gillespie & Croom 1995), which showed that there was no nurse axa on the same island.

1997 ) is that the pathology, as well as the color, was converging between several islets. Orsonwelles, the Hawaiian monophylet species of amber, also has a patterns of first settlement of Kauai, followed by a somewhat ambivalent patterns which at least do not conflict with the principle of progressive development, although the patterns are predominantly within the island species (Hormiga et al. 2003).

It seems that the initial colonisation took place after the formation of Kauai, Oahu and Maui Nui, and after the colonisation of one of these isles, the spread to other isles in all direction was fast. It seems that there has been more inter-island than intra-island diversity. Of the Hawaiian drosophiles, the Drosophila planitibia group's genetic analyses of two mirochondrial and four atomic labels (Bonacum et al. 2005) proposed an ety. 6.

Ma on an Ma on an island older than Kauai, but with a great diversity that begins on Kauai and goes hand in hand with colonisation and diversity as younger isles. As the overall scheme follows the principle of evolution, the species has appeared both within and between the isles, some resettlements, and the Maui Nui Group's diversion schemes (the Molokai, Maui, Lanai and Kahoolawe merged isles, e.g. Price & Elliott Fisk 2004) are particularly complicated and probably include both variance and dispersion.

A remarkable group of basals containing only a few from each of the Kauai and Hawaii isles, with the suspicion that the lack of members of this group of islets is most likely a consequence of disappearance. However, no progressive control patterns were found in the Drosophila halela haleakalia group ("O'Grady & Zilversmit 2004").

Hyposmocoma (Cosmopterigidae), the Hawaiian moths' indigenous family, competes with drosophiles in the number of genera with more than 350 recognised and much more unknown (Rubinoff 2008). The investigation of a small fraction of this variety, the acquatic, so-called "conical" taxpahs, showed that they followed a progressive control patterns with a small emission of three taxpahs on Kauai, resulting in a unique type on Oahu, Molokai and Maui.

Hawaii's varied cosmopterigidae are ready for further research. In Shaw (2002), phylogeny from nearly 40 Hawaiian Laupala endemics (crickets) was analysed using phylogeny from DNA and mtDNA files. They both showed a patterns of colonisation from older to younger isles, but in detail contradictory. However, the mtDNA trees generally indicated that the strains on each island had emerged from a population.

An ensuing study (Mendelson et al. 2004) with AFLPs continued to support an overall colonisation patterns from older to younger isles, but disagreed with the existing nDNA-based Laupala cerasina on Hawaii. There is a clear message from these contradictory research that all of them have value and that a better way is to evaluate the congruency of partsitions in a formal way, to mix them if congruency testing considers it appropriate, and to look for an explanation if they do not (Rubinoff & Holland 2005).

In Shapiro et al. (2006), about 2 kb of Banza (one to three types per island) cricket irradiation (catydides) analyzed 2 kb of 2?kb of Banza cricket and nbDNA sequence, but could not make fixed biogeographical inferences, except that the irradiation may have occurred in Oahu and diversified into two classes, of which no clear progressive control patter.

With four mtDNA biomarkers, Nitta & O'Grady (2008) examined the bio-geography of the 13 types of Hawaiian dicranomyia (crane flies), most of which are not endemic on an island, and found no proof of the progressive ischemia. They were not able to dissolve the ancestor island for the plurality of knots in their phylogenetics.

This shortage of patterns was declared as a result of the relatively lightness with which cranes, although large bugs, are passive from island to island. However, you have found some indications of a differentiation of the rules of evolution within the different types (see below). City of Rivera et al (2002) tried to identify the origin of troglobic (obligatory cave-dwelling) terrestric and epigetic ancestors.

In their COI analyses, two groups of cavern Isoopods originating from Hawaiian Islands colonisations were upheld. Littorophiloscia (one of these groups) derives the individual type of caves from the individual caves. In the other group (Hawaiioscia), however, there are four cave-adapted types, one each on Kauai, Oahu, Molokai and Maui.

of a previously common (and now possibly extinct) surfacing type or group of close related types instead of spreading to the isles. However, due to the lack of surfacing ancestors, this problem is still unsolved. Only the only remaining large Hawaiian vertebrates are the honey veils (Drepanidinae).

One Kauai is the most basic and generally the progressive rules are broadly followed within the various small groups and, as is to be anticipated for such vagil animals, there was no specialization within the island (Fleischer et al. 1998, 2001). These patterns provide a pre-dictive theoretic frame against which the parentage stage, the order of the bifurcations and the colonisation time can be better comprehended, especially when used in a comparing method.

Overall, many of the groups studied so far at least broadly adhere to a progressive control model of diversifications, with the most basic species on the oldest island occurring in the line-spread. However some biogeographical models in Hawaiian radiation are complicated (Funk & Wagner 1995; Holland & Hadfield 2004).

Whereas some endemics groups have clearly emerged on Kauai, others seem to have first colonised either Oahu (agatinelline slugs), Maui Nui (Laupala cerasina group, crickets) or the island of Hawaii (Hylaeus honeybees and a line of mountain slugs: Class A, Fig. 2). For a few groups (Havaika spin, bana grill, Dicranomyia craneflies ) neither the island's initial colonization nor the later patterns of differentiation are clear.

First colonisation and the vagibility of the organism are decisive factors in the determination of the actual allocation samples, in particular compliance with the progressive rules of diversity. So if the descent or colonisation period is approximately the same as the old island and the vagilance is enough to allow them to arrive at islets that have emerged later but are scarce enough to preserve biodiversity, there is the possibility of diversity to comply with the progressive rules.

If, for example, the original colonisation took place before the Oahu suberaerial ascent, then a progressive control patterns from Kauai or an older island would be possible with appropriate vagilance. But if the first colonisation took place after the foundation of e.g. Oahu and Maui Nui, there is no other cause than the coincidence that Kauai is the first colonisation of the island.

If it were even younger, Hawaii could be the place of the first colonisation, in which case a progressional control patterns of diversity from the oldest to the youngest island would not be foreseen. Propagation capability also seems to be an important factor in the extent to which monophylet and islet distribution monophylet radiation follows the progressive principle.

In the case of radiations with a powerful proliferation mechanisms, whether powered or not ( "passive" as recommended for dicranomyia, bird, fly insects), a clear progressive control pathway is not foreseen, regardless of the originally colonised island. The identification of sustainable, correspondingly varied and widely dispersed lines for which extensive Phylogenia can be developped and the determination of the time of first settlement (e.g. Price & Clague 2002) is therefore of central importance for our comprehension of these diversity-paths.

Most of the types of most groups of terrestrial Hawaiian organism are individual islands (Simon 1987), e.g. Landschnecken, about 90 percent (Cowie 1995; Cowie et al. 1995), and Drosophilie, more than 90 percent (Nitta & O'Grady 2008). Less vagil animals are more endemic and some Hawaiian cabbages have decreased their ability to spread actively in comparison to their continentals.

Nevertheless, some types are found on more than one island, others on all major isles and certain groups (e.g. ³cranic flies) show a significantly lower proportion of individual island endemic. Most recent work on a number of different species has discovered intra- and inter-island phylogeographical structures specific to intras. The Drosophila gramshawi is one of a kind among the Drosophila, both in its spread on the major Hawaiian isles and in its seeming patterns of diversity, which do not reflect the predictions that inter-island colonisation will lead to speciesation.

A tribe was formed on Kauai and through the spread two lines were formed, one in Oahu and one in Hawaii (the derivative D. pullipes). While the second group only exists on the Maui Nui Isles, where the genetic flux has been improved by connecting the two, the recent increase in ocean levels has led to their present character.

Out of the 13 Hawaiian Dicranomyia fly varieties, 10 are found on more than one island and 7 on all major high islets. The Nitta & O'Grady (2008) found a sophisticated set of phylogeographical samples within these types, some with a populations structure that reflects an origins on Kauai and ensuing diversity with recolonisation and island skiing, while others appear to have emerged on younger and later colonised older isles.

They have been associated with the relatively lightness with which craneflies are flown from island to island and not with their own proliferation. Dragonflies Megalagrion xanthomelas and Megalagrion philicifica are currently found on five of the Hawaiian islands, all three of them in the Maui Nui area.

2005 affirmed the mutual monophilia of both types using atomic and mitochondrial labels and investigated their physiogeography using the carbonyl mitochondrial and 157 specimens from 25 population. It underlines the importance of the Pleistocene landbridges for the historic transgenic river in Megalagrion and indicates that the recurrent shortages that arose when Maui Nui shifted from almost double the height of the low seaside stocks to the much smaller area we see today, led to the low levels of biodiversity that have been seen on these archipelagos in comparison to the island of Hawaii.

S. coaduca is the only one of 42 Hawaiian succinide varieties to be found on all of Hawaii's major isles ('Cowie et al. 1995). The Netherlands & Cowie (2007) showed that the spread and genetic flux between the isles was sufficiently common to avoid the formation of specimens. However, older isles and volcanos such as Kauai and West Oahu have a tendency to host more divergent population groups, while the population of younger isles and volcanos such as East Oahu, Molokai, Lanai and Maui is structurally more homogenous.

These patterns became manifest in fractured haplotypes for West Oahu and Kauai and a unique continual web of gastropods from East Oahu and Maui Nui (Molokai, Maui and Lanai), once forest landbridge links (Carson & Clague 1995). The patterns indicate that beginning species may occur on older islets and show the importance of the Pleistocene landbridges for improving the genetic diversity between East Oahu and Maui Nui (Figure 3).

A multilocal hawaiian succineid pharmacogenetic study embedded S. coaduca from six Hawaiian Isles within the island of Hawaii Cowade. Intra-specific divergences in the S. coaduca line indicate that the strain is older than the island of Hawaii (0. 43 Ma), which is home to the strains that make up the basalt blade.

The general overviews show, however, that all Hawaiian isles are the sisters of S. Çaduca. However, this sample indicates that a number of succinide strains have emerged on an older island, but that either the older line constituents have become extinct on other isles or that older population groups have not been studied during this work.

Of the Hawaiian honey tractors that can be found on more than one island, some do not show an island distinction of Mt DNA and others, as is to be anticipated for volatile spp. (Tarr & Fleischer 1995). In particular the Amakihi (Loxops stjnegeri from Kauai, Loxops viruses from Oahu, Maui and Hawaii) follow a very clear progressive regulation on Cytb (Fleischer et al. 1998).

For certain endemics on an island, there is some indication of a profound phylogeographical patterning. On Hawaii, for example, three types of Tetragnatha spider show intra-specific pattern of gene structures that reflect lavast flow fractionation due to relatively recent and sustained vulcanic activities (Vandergast et al. 2004). The Oahu's Oahu's endemic and threatened slugs, Achatinella mustelina, showed profound hereditary fractures corresponding to the extinct terrain of the 3. 7 Ma Waianae Mountain Ranges (Holland & Hadfield 2002).

More than 5 percent of the population's molecule divergences indicate that this strain is several million years old and that sexually diverging population groups are reproductive and may be subject to initial spotting. Phylogeographical proofs suggest that this type once had a wide panmic spread, which was restricted to the fragmentation of woodland along hills.

However, within L. ceresina on the island of Hawaii there is a clear phylogeographical rift between the north and south population. The Maui is the source of these population, but it is not clear whether they originated from one (Mendelson et al. 2004) or two colonisations (Shaw 2002). If Hawaiian sorts are endemic or more common on an island depends basically on their vagibility.

The level of phylogeographical structure both between and within isles is also related to vagilance. In order to truly comprehend such a structure, we must grasp the fundamental ecological nature of the relevant type in a comparable context that incorporates other types into the group. Why, for example, is S. Çaduca the only mountain gastropod on all the archipelago?

So what is unique about his ecological background, his biography or his behavior? How is it different from an island's native amber? In general it is assumed that isles are used as basins and cul-de-sacs, with uni-directional movements of types from continent to continent over temporal scale (e.g. Wilson 1961).

Often regarded as some of the most isolated islets in the worid, furthest from a continent's landmass, the Hawaiian isles are often seen as the end of the biogeographical divide. In fact, many Hawaiian once arriving, colonised and developed were regarded as cul-de-sacs - poultry and invertebrates, crops that have been deprived of the propagation mechanism of bird and air seeds (Ziegler 2002) - that did not lead to further spread.

However, more and more cases of spread from the Hawaiian islands are being detected, leading to the colonisation of remote archipelago and evencontinent. Roundll et al (2004) showed that a succinea strain of Tahiti cluster within the island of Hawaii is a group of mountain gastropods (see above) originating in Hawaii.

That has been corroborated by Holland & Cowie (in the press), who also showed that a Samoa succineide in the other large group of Hawaiian island varieties, particularly on the island of Kauai, indicates the origins of Samoan varieties on Kauai. The most spectacular thing is that O'Grady & DeSalle (2008), who have partly resolved the long-standing dispute over the Hawaiian drosophiles' generics classification (see below), have shown that the Hawaiian scaptomyza originates in the Hawaiian isles, is broadly spread there and has then colonised other isles and continents all over the world.

Prospective attempts to explore the Pacific far and more comprehensively will allow us to determine how often such'out-of-Hawaii' colonies have been and whether the Hawaiian islands, and perhaps by way of comparison with other ocean archipels, are not bio-geographically the isolated cul-de-sacs that have often been adopted. The" DNA barcoding", the use of a fraction of a lone biomtDNA label (usually COI) as a means of identification and categorization of the entire biological diversity, was developed in this radical attempt (Hebert et al. 2003).

Hawaiian Islands, with their wonderful variety in many groups, are without a doubt an outstanding place to carry out such work, at least as a precursor to a more extensive systematic work. In Hawaiian landlife, the use of molecule information has provided important insights into the fundamental taxonomies and systems of a number of groups.

Hawaiian drosophiles' genetic diversity has led to significant controversies (O'Grady 2002; van der Linde et al. 2007; O'Grady et al. 2008; Thompson et al. 2008). O'Grady et al (2003) reworked the Hawaiian drosophil species and subspecies on the basis of the morphologic and genetic reasons, all of which were combined within the two genus Drosophila and Scaptomyza, the latter also containing Engiscaptomyza as subspecies.

Drosophila and Scaptomyza are both global, so if the Hawaiian drosophiles actually come from a unique colonisation, these species become para-phyletic, unless Hawaii was the origin of the non-Hawaiian members of one of these groups, Scaptomyza (Russo et al. 1995), which is still leaving Drosophila paraphyletically.

One option that has not been mentioned in formal terms is the option of a separated introduction of Drosophila and Scaptomyza in Hawaii, as suggested by Thomas & Hunt (1991), followed by a hybridisation that makes Hawaiian tacias appear mono-phyletic in human genetic analysis (K. Y. Kaneshiro 2008, Person-to-person communications; see also O'Grady & Zilversmit 2004).

Whereas hybridisation in drosophils was not officially proven by means of genetic proof, it was suggested to elucidate phylogeographical and systemic disparities in Laupala grilles (Shaw 2002) and megalagrion virgins (Jordan et al. 2003). Recent molecule information, however, confirms that Hawaiian drosophilides (Drosophila and Scaptomyza) are monophylet and embedded in the global species Drosophila (O'Grady 2002; O'Grady & DeSalle 2008).

This makes Drosophila indeed para-phyletic with respect to scaptomyza (including non-Hawaiian species) nesting therein. In particular for migratory animals, genetic analyses were used to assist the fundamental system, which included the differentiation between the Laysan and Nihoa millerbreds ( (Fleischer et al. 2007), the phosphogenetic status of the recently deceased Po'o-uli (Fleischer et al. 2001) and the consspecificity of the now extinct Hawaiian Sea Hawk with the Palearctic Sea Eagle ("Fleischer et al. 2000").

Thus, the division of the Havaika spiders into nine nominally oriented morphological specimens turned out to be incompatible with the Arnedo & Gillespie (2006) molecule datasets, which showed four monophylet ic lines on several isles, each line showing a degree of diversity mainly between isles. However, in general, despite the common detection of cryptically modified specimens as a consequence of genetic analyses of invertebrate organisms (e.g. Holland et al. 2004; Bickford et al. 2007), few cases seem to have been found in systematic Hawaiian faunal surveys, as suggested by Holland & Hadfield (2002) and Shaw (2002) (see Schmitz et al. 2008).

One big issue in some groups is that many Hawaiian strains, when initially described, have been classified into poorly differential, overall fishing genus, e.g. Theridion in the spider (Arnedo et al. 2007) and Succinea in the snail (Rundell et al. 2004; Holland & Cowie in the press), and their proper location in these or other genus (including new genera) will depend on the research still to be carried out.

The application of global analytics to control this fundamental system and to solve other challenging taxonomical issues is becoming more and more important in the investigation of the complexity of Hawaiian sediment. With the emergence and growing user-friendliness of human models in the fields of pharmogenetics, bio-geography and photogeography, there has been an increase in the number of trials on the Hawaiian biotas that have expanded our understanding beyond the groundbreaking Wagner & Funk (1995) collection.

Others are beginning to concentrate on improving global cycle estimates and have started to provide a better understanding of the temporal course of important colonisation and cladogenesis ("Price & Clague 2002"). While we know more about the geographical and phyllogenetic origin of Hawaiian vertebrate animals than invertebrate animals, there is much more research into the bio-geography and phyllogeography of invertebrate animals within the islands.

Presumably this is a reflection of the interaction of the generally greater vertebrate sizes, mainly those of migratory animals, in comparison to invertebrate animals, the greater vagueness of migratory animals and the far greater biodiversity of invertebrate animals, which allows rugged analysis of biogeographical samples that are not possible with only three or four of them. Since many of the Hawaiian endemics come from individual, non-radiating colonies, the origin of these colonies is questionable.

Some examples from Asia and North and South America, led by the most well-known morphologic taxionomy, will probably give the answers - and that's why we have the answers for most bird lines in Hawaii. With e.g. 1000 types of drosophiles, these origin issues are not so easy to be answered, and the immediate issues that come to one' s minds are more local, e.g. the pattern and process of diversity within and between the isles.

And with so much indigenous variety, as opposed to just one or a few per line of vertebrates, such issues can indeed only be pertinent in these different groups of invertebrates. The geographical origin of many invertebrates is not known ( (e.g. Drosophiliden, Megalagrion, Laupala and others, for which only rudimentary speculations are possible) and this mirrors the groups' worldwide ignorance and thus necessarily inadequate sample testing of the huge prospective sources (the whole Pacific Rim and the Pacific Islands).

The geographical origin and routes of colonisation, in many cases probably in the form of a springboard, island jumping over the Pacific, remains a broad area of study. With increasing number and dissolution of phylogeographical molecule trials, our capacity to identify common and singular characteristics of the radiation diversity processes increases.

If we look, for example, at the relationship between colonisation processes, colonisation processes and the number of colonisation processes, we can ask the question: Do lines that have colonised the Hawaiian islands several species more than those that originate from a specific process? We are therefore beginning to appreciate the importance of determinants such as the number of long haul versus island populations for determining biological density in Hawaiian lines.

It may not be a particularly determinist topic in itself whether a particular type of irradiation originates from one or more colonisations when it comes to the production of biological diversity. The agateinelline terrestrial slugs ( "a subfamily"; the porcine subsfamily Auriculellinae may have evolved from the same unique colonization), for example, are probably of a common ancestry, but the Hawaiian succinides (one family) are two or more.

So on the sub-family plane there was only one emission per line, as in the agatinelles. However, while agatinelline rays more than doubled the biodiversity of the two succineiden rays (Holland & Hadfield 2004; Holland & Cowie in the press), succinides are much more ecological.

The Hawaiian bird (many orders and families) originated from various colonisations, but Drepanidine from only one. Why, however, agatinelline subfamily radiations comprise far more types (99) than succineide subfamily radiation/colonization (42 in total) (Cowie et al. 1995) is indeed a matter that deserves to be addressed, as it refers to the basic causes why some groups are more likely than others.

Why, for example, are there only two Hawaiian endemics, but well over a thousand moth ( "Ziegler 2002"), 13 craneflies, but probably a thousand drosophilides ( "Kaneshiro 2006"; "Nitta & O'Grady 2008"), and so on? These are just some of the interesting new fields of research we expect in island biological research, especially in the fields of comparing photogeography and co-evolutionary research.

Compare the two types of phylogeographical research has started to uncover penetrating and unpredictable samples that suggest that Cryptic biologically diversified has been more important in the evolution of marine island assemblies than traditionally thought, and that these phenomena are not so readily discernible in the analysis of individual lines. Compare and compare phenomena of geologic and biogeographical origin and the fate of island lines.

Koevolutionary research attempts to comprehend how inter-specific interaction is characterized by mutual nature selectivity, is subjected to concentrated evolving changes and persists through place and place (Thompson 2005). Co-evolution on the Oceania is a largely uncharted boundary that involves several interactive communities spread across multifaceted and multi-island areas, dependent on incidents such as climatic changes, El Niño South Oscillation, volcano eruption and invasions by colonization of speciation.

Even if the Hawaiian islands still lack the necessary genetic information, there is still room for interesting new findings. A great challange for a complete comprehension of phylogenetic, systematic and evolutive aspects of Hawaiian tacias is the pure variety of many invertebrates. More than 5000 recognised indigenous insects exist (Eldredge & Evenhuis 2003), but this number could slightly be doubled as many groups comprise a large number of unrecorded insects (e.g. Kaneshiro 2006; Rubinoff 2008).

Another confusion in the system or hawaiian geological system or hawaiian geography is the recent acceleration in human extinction. 2. Hawaiian islands are known for the extremely extinct populations: over 70 percent of the bird population (Boyer 2008), up to 90 percent of the snail (Cowie 2001; Lydeard et al. 2004) and Undocumented numbers of arthropod (Wagner & Funk 1995, p. 421), which are usually understaffed and whose disappearance is particularly hard to record (Dunn 2005).

In addition, most of the remaining types are restricted to heavily contracting areas in relatively remote, high-altitude areas, making collecting harder. The lack of specimens in the analysis of phylogenetics can result in misinterpretations of the tree, so the extent of its disappearance in Hawaii is of particular importance in the field of phosphogenetic restoration. The issue has already been mentioned, for example in the explanation of the spread of the D. plantitibia group, the ages of the island of Hawaii cloud of mountain slugs and the development of troglobic Isopoda.

However, great progress has been made in developing the ability to isolate genetic material from samples in museums, especially bird samples, and hopefully in the FFH. Since the mid-1990s, Hawaiian wildlife has undergone a number of research projects. Hawaiian wildlife much more than we did when Wagner & Funk (1995) released their turning point album.

Thanx to Rob Fleischer and Sheila Conant for their help with bird literary, Neal Evenhuis, Ken Kaneshiro and Patrick O'Grady for drosophiliden and drosophiliden literary discussions and Chris Simon and Rob DeSalle for their commentaries on the work. An article of 15 on a topic "Evolution on Pacific Islands":

Landsnail modells in island biogeography: a story of two slugs.