Hawaiian Honeycreeper

Hawaiian honeycreepers are a group of small birds endemic to Hawaii. They are members of the finch family Fringillidae, closely related to the rosefinches (''Carpodacus''), but many species have evolved features unlike those present in any other finch. Their great morphological diversity is the result of adaptive radiation in an insular environment. Many have been driven to extinction since the first humans arrived in Hawaii, with extinctions increasing over the last two centuries following European discovery of the islands, with habitat destruction and especially invasive species being the main causes.

Taxonomy

Before the introduction of molecular phylogenetic techniques, the relationship of the Hawaiian honeycreepers to other bird species was controversial. The honeycreepers were sometimes categorized as a family Drepanididae,Clements, J. 2007. ''The Clements Checklist of the Birds of the World.'' 6th ed. other authorities considered them a subfamily, Drepanidinae, of Fringillidae, the finch family. The entire group was also called Drepanidini in treatments where buntings and American sparrows ( Passerellidae) were included in the finch family; this term is preferred for just one subgroup of the birds today.AO
Check-list of North American Birds
Accessed 26 December 2007 Most recently, the entire group has been subsumed into the finch subfamily Carduelinae, without a named taxon assigned to this group.

The Hawaiian honeycreepers are the sister taxon to the '' Carpodacus'' rosefinches. Their ancestors are thought to have been from Asia and diverged from ''Carpodacus'' about 7.2 million years ago, and they are thought to have first arrived and radiated on the Hawaiian Islands between 5.7–7.2 million years ago, which was roughly the same time that the islands of Ni'ihau and Kauai formed. The lineage of the recently extinct po'ouli (''Melamprosops'') was the most ancient of the Hawaiian honeycreeper lineages to survive to recent times, diverging about 5.7–5.8 million years ago. The lineage containing '' Oreomystis'' and '' Paroreomyza'' was the second to diverge, diverging about a million years after the po'ouli's lineage. Most of the other lineages with highly distinctive morphologies are thought to have originated in the mid-late Pliocene but prior to the formation of Maui. Due to this, Oahu likely played a key role in the formation of diverse morphologies among honeycreepers, allowing for cycles of colonization and speciation between Kauai and Oahu.

A phylogenetic tree of the recent Hawaiian honeycreeper lineages is shown here. Genera or clades with question marks (?) are of controversial or uncertain taxonomic placement.

The classification of '' Paroreomyza'' and '' Oreomystis'' as sister genera and forming the second most basal group is based on genetic and molecular evidence, and has been affirmed by numerous studies; however, when morphological evidence only is used, ''Paroreomyza'' is instead the second most basal genus, with ''Oreomystis'' being the third most basal genus and more closely allied with the derived Hawaiian honeycreepers, as ''Oreomystis'' shares traits with the derived honeycreepers, such as a squared-off tongue and a distinct musty odor, that ''Paroreomyza'' does not. This does not align with the genetic evidence supporting ''Paroreomyza'' and ''Oreomystis'' as sister genera, and it would be seemingly impossible for only ''Paroreomyza'' to have lost the distinctive traits but ''Oreomystis'' and all core honeycreepers to have retained or convergently evolved them, thus presenting a taxonomic conundrum.

'' Viridonia'' (containing the greater ʻamakihi) may be associated with or even synonymous with the genus '' Aidemedia'' (containing the prehistoric icterid-like and sickle-billed gapers), and has the most debated taxonomy; it was long classified within the "greater ''Hemignathus''" radiation (a now-paraphyletic grouping containing species formerly lumped within ''Hemignathus'', including ''Hemignathus'', ''Akialoa'', and '' Chlorodrepanis'') and while some sources speculate it as being sister to ''Chlorodrepanis'' (containing the lesser ʻamakihis), other sources speculate it may be a sister genus to the genus '' Loxops'' (containing the 'akepas, ʻakekeʻe and ʻalawī).

Characteristics

Nearly all species of Hawaiian honeycreepers have been noted as having a unique odor to their plumage, described by many researchers as "rather like that of old canvas tents".

Today, the flowers of the native ōhia (''Metrosideros polymorpha'') are favored by a number of nectarivorous honeycreepers. The wide range of bill shapes in this group, from thick, finch-like bills to slender, down-curved bills for probing flowers have arisen through adaptive radiation, where an ancestral finch has evolved to fill a large number of ecological niches. Some 20 species of Hawaiian honeycreeper have become extinct in the recent past, and many more in earlier times, following the arrival of humans who introduced non-native animals (ex: rats, pigs, goats, cows) and converted habitat for agriculture.

Genera and species

The term "prehistoric" indicates species that became extinct between the initial human settlement of Hawaii (i.e., from the late 1st millennium AD on) and European contact in 1778.

Subfamily ''Carduelinae''

* Drepanidini
**Genus '' Aidemedia'' Olson & James, 1991 – straight thin bills, insectivores
***''Aidemedia chascax'' Olson & James, 1991 – Oahu icterid-like gaper (prehistoric)
***''Aidemedia lutetiae'' Olson & James, 1991 – Maui Nui icterid-like gaper (prehistoric)
***''Aidemedia zanclops'' Olson & James, 1991 – sickle-billed gaper (prehistoric)
**Genus '' Akialoa'' Olson & James, 1995 – pointed, long and down-curved bills, insectivorous or nectarivorous
***''Akialoa ellisiana'' Gray, 1859 – Oʻahu ʻakialoa (extinct, 1940)
***''Akialoa lanaiensis'' Rothschild, 1893 – Maui Nui ʻakialoa (extinct, 1892)
***''Akialoa stejnegeri'' Wilson, 1889 – Kauaʻi ʻakialoa (extinct, 1969)
***''Akialoa obscura'' Cabanis, 1889 – lesser ʻakialoa (extinct, 1940)
***''Akialoa upupirostris'' – hoopoe-billed ʻakialoa (prehistoric)
**Genus '' Chloridops'' Wilson, 1888 – thick-billed, hard seed (e.g. '' Myoporum sandwicense)'' specialist
***''Chloridops kona'' Wilson, 1888 – Kona grosbeak (extinct, 1894)
***''Chloridops regiskongi'' – King Kong grosbeak (prehistoric)
***''Chloridops wahi'' – wahi grosbeak (prehistoric)
**Genus '' Chlorodrepanis'' Olson & James, 1995 – pointed bills, insectivorous and nectarivorous
***''Chlorodrepanis stejnegeri'' Pratt, 1989 – Kauaʻi ʻamakihi
***''Chlorodrepanis flava'' Bloxam, 1827 – Oʻahu ʻamakihi
***''Chlorodrepanis virens'' Cabanis, 1851 – Hawaiʻi ʻamakihi
**Genus '' Ciridops'' Newton, 1892 – finch-like, fed on fruit of '' Pritchardia'' species
***''Ciridops anna'' Dole, 1879 – ʻula-ʻai-hāwane (extinct, 1892 or 1937)
***''Ciridops tenax'' Olson & James, 1991 stout-legged finch (prehistoric)
** Genus '' Drepanis'' Temminck, 1820 – down-curved bills, nectarivores
***''Drepanis funerea'' Newton, 1894 – black mamo (extinct, 1907)
***''Drepanis pacifica'' Gmelin, 1788 – Hawaiʻi mamo (extinct, 1898)
***''Drepanis coccinea'' Forster, 1780 – ʻiʻiwi
** Genus ''Dysmorodre''''panis'' Perkins, 1919 – pincer-like bill, possibly snail specialist
***''Dysmorodrepanis munroi'' Perkins, 1919 – Lanaʻi hookbill (extinct, 1918)
**Genus '' Hemignathus'' Lichtenstein, 1839 – pointed or long and down-curved bills, insectivorous
***''Hemignathus affinis'' – Maui nukupuʻu (extinct, 1995–1998)
***''Hemignathus hanapepe'' – Kauaʻi nukupuʻu (extinct, 1998)
***''Hemignathus lucidus'' – Oʻahu nukupuʻu (extinct, 1837)
***''Hemignathus vorpalis'' James & Olson, 2003 – giant nukupu'u (prehistoric)
***''Hemignathus wilsoni'' Rothschild, 1893 – ʻakiapolaʻau
**Genus ''Himatione'' – thin-billed, nectarivorous
***''Himatione sanguinea'' Gmelin, 1788 – ʻapapane
***''Himatione fraithii'' – Laysan honeycreeper (extinct, 1923)
** Genus '' Loxioides'' Oustalet, 1877 – finch-like, Fabales seed specialists
***''Loxioides bailleui'' Oustalet, 1877 – palila
***''Loxioides kikuichi'' Olson & James, 2006 – Kaua'i palila (prehistoric, possibly survived to the early 18th century)
**Genus '' Loxops'' – small pointed bills with the tips slightly crossed, insectivorous
***''Loxops caeruleirostris'' Wilson, 1890 – ‘akeke‘e
***''Loxops coccineus'' Gmelin, 1789 – Hawaiʻi ʻakepa
***''Loxops ochraceus'' Rothschild, 1893 - Maui ʻakepa (extinct, 1988)
***''Loxops wolstenholmei'' Rothschild, 1895 – Oʻahu ʻakepa (extinct, 1990s)
***''Loxops mana'' Wilson, 1891 – Hawaiʻi creeper
**Genus ''Magumma -'' small pointed bills, insectivorous and nectarivorous
***''Magumma parva'' Stejneger, 1887 - ʻanianiau
**Genus ''Melamprosops'' Casey & Jacobi, 1974 – short pointed bill, insectivorous and snail specialist
***''Melamprosops phaeosoma'' Casey & Jacobi, 1974 – poʻouli (extinct, 2004)
** Genus ''Oreomystis'' Wilson, 1891 – short pointed bills, insectivorous
***''Oreomystis bairdi'' Stejneger, 1887 – ʻakikiki
**Genus ''Orthiospiza'' – large weak bill, possibly soft seed or fruit specialist?
***''Orthiospiza howarthi'' James & Olson, 1991 - highland finch (prehistoric)
**Genus ''Palmeria'' Rothschild, 1893 – thin-billed, nectarivorous, favors ''Metrosideros polymorpha''
***''Palmeria dolei'' Wilson, 1891 – ʻakohekohe
**Genus '' Paroreomyza'' – short pointed bills, insectivorous
***''Paroreomyza maculata'' Cabanis, 1850 – Oʻahu ʻalauahio (possibly extinct, early 1990s?)
***''Paroreomyza flammea'' ( Wilson, 1889) – kākāwahie (extinct, 1963)
***''Paroreomyza ''
****''Paroreomyza '' Wilson, 1890 – Lana'i 'alauahio (extinct, 1937)
****''Paroreomyza newtoni'' (Rothschild, 1893) – Maui ‘alauahio
**Genus ''Pseudonestor'' – parrot-like bill, probes wood for insect larvae
***''Pseudonestor xanthophrys'' Rothschild, 1893 – Maui parrotbill or kiwikiu
**Genus ''Psittirostra'' – slightly hooked bill, '' Freycinetia arborea'' fruit specialist
***''Psittirostra psittacea'' Gmelin, 1789 – ʻōʻū (probably extinct, 1998?)
**Genus '' Rhodacanthis'' – large-billed, granivorous, legume specialists
***''Rhodacanthis flaviceps'' Rothschild, 1892 – lesser koa-finch (extinct, 1891)
***''Rhodacanthis forfex'' James & Olson, 2005 – scissor-billed koa-finch (prehistoric)
***''Rhodacanthis litotes'' James & Olson, 2005 – primitive koa-finch (prehistoric)
***''Rhodacanthis palmeri'' Rothschild, 1892 – greater koa-finch (extinct, 1896)
**Genus '' Telespiza'' Wilson, 1890 – finch-like, granivorous, opportunistic scavengers
***''Telespiza cantans'' Wilson, 1890 – Laysan finch
***''Telespiza persecutrix'' James & Olson, 1991 – Kauaʻi finch (prehistoric)
***''Telespiza ultima'' Bryan, 1917 – Nihoa finch
***''Telespiza ypsilon'' James & Olson, 1991 – Maui Nui finch (prehistoric)
**Genus '' Vangulifer'' – flat rounded bills, possibly caught flying insects
***''Vangulifer mirandus'' – strange-billed finch (prehistoric)
***''Vangulifer neophasis'' – thin-billed finch (prehistoric)
**Genus ''Viridonia''
***''Viridonia sagittirostris'' Rothschild, 1892 – greater ʻamakihi (extinct, 1901)
** Genus '' Xestospiza'' James & Oslon, 1991 – cone-shaped bills, possibly insectivorous
***''Xestospiza conica'' James & Olson, 1991 – cone-billed finch (prehistoric)
***''Xestospiza fastigialis'' James & Olson, 1991 – ridge-billed finch (prehistoric)

Hawaiian honeycreepers were formerly classified into three tribes – Hemignathini, Psittirostrini, and Drepanidini – but they are not currently classified as such.

Conservation

See also

* Hawaiian honeycreeper conservation
* List of adaptive radiated Hawaiian honeycreepers by form

Cited references

Other references

*Groth, J. G. 1998. Molecular phylogeny of the cardueline finches and Hawaiian honeycreepers. Ostrich, 69: 401.

External links

Hawaiian Honeycreepers (Drepanididae)
information, including 4 species with videos and 11 with photographs at th
Internet Bird Collection

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Bird common names
Carduelinae