Laysan honeycreeper

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Laysan honeycreeper
Black-and-white photo of a bird with open jaws on a rock
Male Laysan honeycreeper photographed by
Donald R. Dickey in 1923, a few days before the extinction
of the species

Extinct (1923)  (IUCN 3.1)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Family: Fringillidae
Subfamily: Carduelinae
Genus: Himatione
Species:
†H. fraithii
Binomial name
†Himatione fraithii
Map showing the Hawaaiian Islands with inset close ups of some areas
Map of the
Laysan Island
in the lower left inset box
Synonyms[2]
List
  • H. fraithi
    Rothschild, 1892
  • H. freethii
    Rothschild, 1893–1900
  • H. freethi
    Rothschild, 1893–1900
  • H. frethii
    Schauinsland, 1899
  • H. sanguinea fraithii
    Hartert
    , 1919
  • H. sanguinea fraithi
    Delacour
  • H. sanguinea freethii
    Amadon
    , 1950
  • H. sanguinea freethi
    Pratt
    et al., 1987

The Laysan honeycreeper (Himatione fraithii), also known as the Laysan ʻapapane or Laysan honeyeater, is an

ʻapapane. The specific name, fraithii, refers to George D. Freeth, the self-appointed governor of Laysan, but was misspelled. Rothschild attempted to emend it to freethi in a later publication. This was accepted by most subsequent authors throughout the 20th century, and the bird was also considered a subspecies of the ʻapapane, as H. sanguinea freethii, for most of this time. By the 21st century, after further research, the original name was reinstated and it was considered a full species again. As a Hawaiian honeycreeper, a grouping within the finch subfamily Carduelinae
, its ancestors are thought to have come from Asia.

The Laysan honeycreeper was 13–15 cm (5–6 in) long and its wing measured 64–69 mm (2.5–2.7 in). It was bright scarlet

covert feathers were grayish. The wings, tail, bill, and legs were dark brown, and the iris was black with a brown outline. Immature birds were brown with paler lower parts, and had green edges to their wing-covert feathers. The bill was slender and downturned. The sexes were alike, though the bill, wings, and tail were slightly shorter in the female. The ʻapapane differs from the Laysan honeycreeper in features such as being blood-red overall and having a longer bill. The song of the Laysan honeycreeper was described as low and sweet, consisting of several notes. Laysan is a remote coral island with an area of 3.6 km2 (1.4 sq mi). The honeycreeper lived throughout, but was most abundant in the interior among tall grass and low bushes near the open plain that bordered the island's lagoon
.

This bird was very active and, though less trusting than other birds, sometimes entered buildings to hunt

sandstorm
hit the island, and the last birds perished due to lack of cover. The destruction of Laysan's vegetation led to the extinction of three out of five of its endemic land birds.

Taxonomy

The Laysan honeycreeper was first recorded on

humming bird" ("Colibri" in German), in both cases probably referring to the honeycreeper, the latter due to it feeding on nectar.[2][3][4][5]

In 1892, the British zoologist and banker

Spartans to war, in reference to the color of the ʻapapane.[10]

Frederick W. Frohawk
, 1893–1900; 23 (lower left) is the Laysan honeycreeper, with the variant spelling H. freethii

The

Hugo H. Schauinsland used the spelling frethii in 1899.[13]

After the original description, all the spellings of the name were used by different authors, but the original fraithii was only used a few times.

Harold D. Pratt also indicated that the name had been corrected within the original description.[10] In 2011, the American ornithologist Peter Pyle pointed out that Rothschild had not corrected the name until the first part of his monograph was published in 1893, and that he also appears to have realized his emendation was inappropriate, since he reverted to the original spelling fraithi in the third part. Pyle therefore concluded that according to the International Code of Zoological Nomenclature, the name should not be emended and the original spelling should be reinstated.[11]

In 2015, Pratt, Pyle, and the American ornithologist Reginald E. David formally proposed to the

International Ornithological Committee in their world list of birds the same year.[18]

There are at least 105 known specimens (six of them mounted) of the Laysan honeycreeper in museums across the world, but two specimens appear to have gone missing. Some museums have multiple specimens, including 24 in the

Bernice Pauahi Bishop Museum in Honolulu, 20 (including the type specimen) in the American Museum of Natural History in New York City, and 20 in the National Museum of Natural History in Washington. There are also at least two skeletons, three nests, and one egg preserved. The known specimens were collected between 1892 and 1913.[9][19][20] The ratio of males to females between adult specimens in museum collections is 1.7:1.[21]

Evolution

Photo of three taxidermied bird
Taxidermied Laysan finch (upper left), Laysan honeycreeper (upper middle), and Laysan rail (below), 1903; the latter two are extinct

In 1899, Schauinsland considered the Laysan honeycreeper an example of how a new species may arise through isolation and noted its resemblance to the ʻapapane.[13] Perkins stated in 1903 that the Laysan honeycreeper was descended from the ʻapapane colonizing the island, and he divided the Hawaiian honeycreepers into two main groups.[22] Amadon noted in 1950 that although the Laysan finch (Telespiza cantans) and the Laysan honeycreeper spent more time on the ground than their relatives, their power of flight was seemingly not reduced. He pointed out that the length of their wings was rather short, which perhaps indicated a beginning tendency in such a direction. The wing of the finch is shorter, perhaps because it had reached Laysan earlier than the honeycreeper.[15]

In 1976, the American geologist Seymour O. Schlanger and botanist George W. Gillett proposed that because Laysan had been a raised coral island until 18,000 years ago whereafter erosion and tectonic subsidence reduced its height, it could have been a refugium for upland and montane species that had adapted to the drastic changes in habitat. They pointed to the Laysan honeycreeper and Laysan finch as evidence for this, being the only Hawaiian honeycreepers living close to beaches.[23] The American ornithologists Storrs L. Olson and Helen F. James considered the Laysan honeycreeper a distinct species from the ʻapapane in 1982, but without elaboration.[24][25]

Olson and the American ornithologist Alan C. Ziegler stated in 1995 that while the Laysan honeycreeper was often considered a subspecies of ʻapapane, its skull features indicate it was distinct and probably more primitive, and therefore perhaps a remnant of an earlier evolutionary stage rather than being particularly specialized for the conditions on Laysan. They speculated that if this bird could survive on Laysan, there could also be a niche for a relative on the nearby island of Nihoa. They also argued that Hawaiian honeycreepers are not actually an upland group, but that this perception of them as such comes from them having been wiped out from lowland areas of Hawaii in prehistoric times by human-made habitat destruction, and that many fossils of the group (including of Himatione) have been found in areas just above sea level. They therefore disagreed with the idea that the species found there were a remnant of upland populations or necessarily ancient occupants.[5]

The American biologist Mark J. Rauzon speculated in 2001 that the Laysan honeycreeper could have descended from ʻapapanes that flew there from the rainforests of

osteological features found Himatione to group in a clade similar to what Perkins suggested in 1903. Her "clade 11" is depicted in the cladogram below († denotes recent extinctions, ‡ denotes prehistoric):[27]

ʻapapane
, the closest relative of the Laysan honeycreeper

Vestiaria coccinea
(ʻiʻiwi)

†

Drepanis funerea
(black mamo)

†

Drepanis pacifica
(Hawaii mamo)

Himatione sanguinea
(ʻapapane)

†Himatione freethii (Laysan honeycreeper, now fraithii)

Palmeria dolei
(ʻākohekohe)

‡

Ciridops tenax
(stout-legged finch)

‡Ciridops sp. (from Oahu)

†

Ciridops anna
(ʻula-ʻai-hāwane)

The Hawaiian honeycreepers, variously considered to constitute the family Drepanididae (formerly spelled "

ʻākohekohe (Palmeria dolei) by some studies, Pratt suggested in 2014 that their genera Himatione and Palmeria might be merged.[31]

Description

Illustration of four birds, three light and one blood-red red
Adult male Laysan honeycreeper (A), adult female (B), juvenile (C), and ʻapapane (D), by John Gerrard Keulemans, 1893–1900

The Laysan honeycreeper was a small bird, with published length measurements ranging from 13–15 cm (5–6 in).

covert feathers were grayish. The wings, tail, bill, and legs were dark brown, and the iris was black with a brown outline. Immature birds were brown, with paler lower parts, and had green edges to their wing-covert feathers.[32] The bill was slender and downturned.[33]

The sexes were alike, though the bill, wings, and tail were slightly shorter in the female. Although Rothschild stated in his 1892 description that the female was paler than the male, the American zoologist

molted Laysan honeycreepers were a deeper red and not as easy to distinguish from the ʻapapane, while the latter did not fade to as pale a red.[12]

The ʻapapane differs from the Laysan honeycreeper in being blood-red overall, with black wings and tail, whiter undertail covert feathers, and a longer bill.

primary feathers did not have oblique truncation (in the ʻapapane this truncation produces a "wing note" sounding like vocalization).[10] Amadon suggested in 1950 that the fading and bleaching of Laysan landbirds was in part due to the exposed nature of the island.[15] Olson and Ziegler also suggested in 1995 that the intense sunlight of Lysan had caused the honeycreeper's plumage to fade, accounting for the difference from the ʻapapane, but noted it had been found to be distinct in osteological features.[5][21] Pratt countered in 2005 that white undertail feathers cannot fade to brown since they lack pigmentation to begin with.[10]

Palmer reported the song of the Laysan honeycreeper as low and "sweet", consisting of several notes. He noted it was usually silent, except during the

Donald R. Dickey stated that their "charming song is out of proportion to their size" in 1923.[36]

Habitat

aerial photograph

The Laysan honeycreeper was

subsurface substratum is coralline rock, and its topography suggests it was once part of an atoll with a lagoon that occupies about one-fifth of the island's center, and is now almost filled with sand and coral fragments. The island is ringed by sand dunes, but is otherwise well-vegetated. The island's original flora was the most varied of the northwestern Hawaiian islands, but much of it was destroyed by human activities by 1923, leaving near-desert-like conditions and several extinct plant species, though the extent of the vegetation had almost recovered by 1973.[19][3][32]

In 1903, Fisher stated that the Laysan honeycreeper was found all over Laysan Island, but was most abundant in the interior among tall grass and low bushes near the open plain that bordered the lagoon, an area where all the landbirds appeared to congregate. This was also the favored nesting area, with its broad patches of the succulent Portulaca that these birds fed from. Munro added that they also frequented grasstops and other plants on the fringes of the lagoon. Their bright, scarlet plumage made them conspicuous as they fluttered among the soft green Chenopodium bushes.[34][37][35] This species was the only nectar-feeding finch of the northwestern Hawaiian Islands.[2]

Behavior and ecology

ʻākulikuli, some of the flowers that the Laysan honeycreeper fed nectar
from

Few naturalists encountered the Laysan honeycreeper and few accounts were left of its life history. It was very active, like the

brackish water.[35]

The Laysan honeycreeper was

petals reminded Schauinsland and Fisher of hummingbirds, though it did so by walking rather than hovering in front of them.[34][37][13][19] The Laysan honeycreeper gathered insects from flowers, such as small, green caterpillars, and were fond of the large, brownish moths called millers (including species of Agrotis and others that have since disappeared[39]), which were abundant on the island, and were also fed on by other insect-eating birds. The honeycreepers were observed extracting moths from between boards, grasping them with one foot (always the left according to Freeth) while eating the soft parts, leaving the wings and other hard parts.[34][37][35]

Reproduction

Walter K. Fisher
, 1902

Fisher noted that the nest of the Laysan honeycreeper was more difficult to find than that of the Laysan millerbird, and found only one, in the middle of a

albatrosses, but there were no large, white feathers, which made the nest indistinguishable from that of the Laysan millerbird, which built nests in nearby tufts. It differed from that of the millerbird in being tighter in construction and having a shallower cup. The nest was also likened to that of the Laysan finch; the nests of different land birds of the island may have been similar due to the limited selection of building materials.[34][37][12][13]

Little is known about the breeding cycle of the Laysan honeycreeper, and most observers did not record when nests and young were found.

Alfred M. Bailey, who visited Laysan in 1912, stated that the clutch size was four or five eggs; sets of three were taken by collectors.[19] The ovate eggs were glossless white, with grayish blotches and spots at the larger end, and reddish brown spots above them, these markings often forming circles. The eggs varied in size, but a typical egg measured 18 by 13.7 mm (0.71 by 0.54 in). The eggs were similar to those of the short-toed treecreeper (Certhia brachydactyla) and the barn swallow (Hirundo rustica), but much less glossy.[34][37][12]

Extinction

sandstorm
that killed the last three Laysan honeycreepers (lower right)

The few observations of the Laysan honeycreeper indicate it was not abundant on Laysan to begin with, and only three population estimates were made. Isenbeck already considered the bird uncommon when visiting in 1828 in the first report of the species; Palmer considered it the rarest of the island's birds in 1890, though finding them in fair numbers, and Fisher gave a similar assessment in 1903.

meat-canning business that would provide food for guano miners and to amuse his children. The venture did not succeed, but the rabbits proceeded to destroy the island's vegetation.[32][21][38][41]

In 1909, the US president

feather hunters, as well as several Laysan honeycreeper skins, but noted the birds were still fearless towards humans. They estimated that 300 Laysan honeycreepers remained and that they and other birds there were "doomed to extermination" if their food supply was not preserved.[38]

Bailey recalled in 1956 that a singing honeycreeper perched on a dead

wild tobacco, the few remaining Scaevola plants, and grass tufts.[33] In 1915, the American naval officer William H. Munter reported that the Laysan honeycreeper was fairly common, and that they were judged to number 1000.[42] The American ornithologists Charles A. Ely and Roger B. Clapp pointed out in 1973 that Munter's estimates were perhaps too generous, as he did not consider them numerous the following year.[19]

Male filmed by Dickey in 1923, a few days before the species' extinction

In 1923, the

field notes
on the Laysan honeycreeper from April 11, 1923, read as follows:

Reno's report of finding 3 specimens of Himatione alive and thriving today in the tobacco patch SW of the lagoon makes me all the more certain that the species could all have adapted themselves to the changed conditions and gained a livelihood had they only been granted nesting cover to guarantee new recruits to take the place of the older birds as they died off. This tiny Honey Eater was probably the most specialized in its feeding habits of all the endemic land birds on the island. God knows when the last flower bloomed on this barren waste, yet here are at least three individuals of this specialized form persisting as a sort of heritage from the last nest of the species that was built in sufficient cover to survive. But as it seems to me, old age and death now inevitably stalk this childless remnant of a vanishing species.[36][26]

The destruction of Laysan's vegetation led to the extinction of three out of five of its endemic land birds, and many other bird species have been driven to extinction by human activities across Hawaii (both in prehistoric and modern times).[21] Wetmore reported that the Laysan millerbird had disappeared entirely by his 1923 visit (probably during another sandstorm only months earlier[36]) and only two Laysan rails (Zapornia palmeri) remained, while about 20 Laysan ducks (Anas laysanensis) had survived. Laysan finches were still singing and hopping about, and the large seabird colonies were not as affected. The rabbits were eradicated from Laysan by the Tanager Expedition during 1923, leading to a remarkable recovery of the vegetation, but too late for the Laysan honeycreeper.[43][33][32][41] Though many searches for the bird were subsequently attempted, such as the Vanderbilt Pacific Equatorial Expedition of 1951, none could be found.[20] While the Laysan rail probably did not survive on the island for long after the storm, it had been introduced to Midway Atoll, but went extinct there too by 1945 because rats (which preyed on eggs, chicks, and perhaps adults) were introduced accidentally during US military construction beginning in 1940.[41]

References

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  2. ^ a b c d e f g h Pyle, Robert L.; Pyle, Peter (2017). "The Birds of the Hawaiian Islands: occurrence, history, distribution, and status" (PDF). Bishop Museum. Archived (PDF) from the original on 27 April 2024. Retrieved 4 June 2024.
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  16. ^ "Proposal 2015-A-10: Split Laysan Honeycreeper from Apapane Himatione sanguinea and change its specific epithet to fraithii" (PDF). American Ornithologists' Union. Archived from the original (PDF) on 24 July 2015. Retrieved 24 July 2015.
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