Laysan honeycreeper

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Laysan honeycreeper
Laysan honeycreeper photographed by Donald R. Dickey in 1923 (probably taken from his footage), a few days before the species' extinction[1]

Extinct (1923)  (IUCN 3.1)[2]
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 of Hawaii showing Laysan Island in the lower left inset box
Synonyms
List
  • Himatione fraithi
  • Himatione freethii
  • Himatione freethi
  • Himatione frethii
  • Himatione sanguinea fraithii
  • Himatione sanguinea fraithi
  • Himatione sanguinea freethii
  • Himatione sanguinea freethi

The Laysan honeycreeper (Himatione fraithii), also known as the Laysan ʻapapane or Laysan honeyeater, is an extinct species of finch that was endemic to the island of Laysan in the Northwestern Hawaiian Islands.

Taxonomy[edit]

The Laysan honeycreeper was first noticed on Laysan Island 3 April, 1828, by C. Isenbeck, surgeon of the Russian ship Moller, which was visiting the Hawaiian Islands (then called the Sandwich Islands). This report was published in an 1834 article by the German naturalist Heinrich von Kittlitz. Isenbeck referred to a "red bird" and a "humming bird" ("Colibri"), in both cases probably referring to the honeycreeper, the latter due to it feeding on nectar.[3][4][5]

In 1892, the British zoologist and banker Walter Rothschild described and named seven new bird species from Hawaii, obtained by the New Zealand collectors Henry C. Palmer and George C. Munro In 1891, including a series of ten honeycreeper specimens from Laysan. These and other collectors had been sent to the Hawaiian Islands in 1890–93 to collect specimens for Rothschild's Natural History Museum at Tring in England. Rothschild named the honeycreeper Himatione fraithii, and classified it as a Hawaiian honeycreeper of the finch family Drepanidae. He found it to resemble the ʻapapane (H. sanguinea) of the same genus which is found across the main Hawaii archipelago, while differing in various details.[6][3][7][8] The generic name is derived from the Greek word himation, a crimson cape worn by the spartans to war, in reference to the color of the ʻapapane.[9]

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

The specific name acknowledges George D. Freeth, the self-appointed US governor of Laysan, manager of the guano-mining there, and amateur naturalist, who had assisted Palmer and Munro. The misspelling of Freeth's name, fraithii, may have been due to a miscommunication or erroneous assumption, and his full name was not mentioned in the description.[10][3] From 1893–1900, Rothschild published a three-part monograph on the birds of Laysan, with further observations about the honeycreeper, which he referred to as the Laysan honey-eater. While some related green species had previously also been considered part of the genus Himatione, he agreed with the American zoologist Robert C. L. Perkins that those should be moved to Chlorodrepanis, and restricted Himatione to the red species; the ʻapapane and the Laysan honeycreeper. By this time, Rothschild had realized that he had misspelled Freeth's name, and attempted to emend the spelling to H. freethi, but also used the spellings fraithi and freethii in various sections of the work.[11][3] The German zoologist Hugo H. Schauinsland used the spelling H. frethii in 1899.[12]

After the original description, all the various spellings of the name were used by different authors, but fraithii was only used by the German ornithologist Ernst Hartert in 1920 and by the American ornithologists Edwin H. Bryn and James C. Greenway in 1944. Hartert also considered the bird a subspecies of ʻapapane, as H. sanguinea fraithii, and listed the type specimen as being an adult male. This classification was followed by most other taxonomists and the trinomial name was used throughout the 20th century.[3][10][13][14] In 1950, the American ornithologist Dean Amadon considered the spelling fraithii a lapsus calami (misspelling), and justified using freethii instead, since Rothschild later corrected the name to that.[15][9] In 2005, the American ornithologist Harold Douglas Pratt also indicated that the name had been corrected within the same publication.[9] In 2011, the American ornithologist Peter Pyle pointed out that Rothschild had not corrected the name in the same publication, but not until the first part of his monograph published in 1893, and that he also appears to have realised 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 retained.[10]

In 2015, Pratt, Pyle, and the American ornithologist Reginald E. David formally proposed to the American Ornithologists' Union (AOU) that the Laysan honeycreeper should be split into its own full species again and its specific name changed back to fraithii, based on the recent research that supported this. They also suggested that the bird should be listed in the Checklist of North American Birds under the common English name Laysan honeycreeper rather than Laysan ʻapapane, a name that had by then become popular with some writers, as this would also require a modifier for the name of the ʻapapane. They considered the name Laysan ʻapapane a modern retrofitting of Hawaiian names to species whose Hawaiian names are unknown or never had one to begin with, and noted that Hawaiians never appear to have visited Laysan Island, which has no traditional Hawaiian name either. These authors stated that they supported the use of Hawaiian neologisms for local use, but thought the AOU should continue designating English names using loan-words where appropriate, but not create new ones in other languages. They also found it appropriate that at least one species of Hawaiian honeycreeper would retain the word "honeycreeper" in its common name, for sentimental reasons.[16] In 2015 the AOU implemented these propositions in their check-list.[17] The changes were also adopted by 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 various museums across the world, but two specimens appear to have gone missing. Some museums have multiple specimens, such as 24 in the Bernice Pauahi Bishop Museum, 20 (including the type specimen) in the American Museum of Natural History, and 20 in the National Museum of Natural History. There are also at least two skeletons, three nests, and one egg preserved. The known specimens were collected between 1892 and 1913.[19][8][20]

Evolution[edit]

Taxidermied Laysan finch, Laysan honeycreeper, and Laysan rail, 1903

Schauinsland considered the Laysan honeycreeper an example of how a new species may arise through isolation in 1899.[12] 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.[21] Amadon noted in 1950 that while 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 was shorter, perhaps because it had reached Laysan earlier than the honeycreeper.[15]

In 1976, the American ornithologists Seymour O. Schlanger and George W. Gillett proposed that because Laysan been a raised coral island until 18.000 years ago whereafter erosion and tectonic subsidence reduced it in height, it could be seen as 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.[22] Olson and the American paleontologist Helen F. James considered the Laysan honeycreeper a distinct species from the ʻapapane in 1982, but without elaboration.[23][24]

The American ornithologists Storrs L. Olson and 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 have been a niche for a relative on the island of Nihoa. They also pointed out that the idea that Hawaiian honeycreepers were an upland group is an artefact of 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.[25]

Pratt and the American biologist Thane K. Pratt stated in 2001 that due to its distinct physical features, the Laysan honeycreeper was unquestionably distinct from the ʻapapane following the Phylogenetic Species Concept. They added that potential isolating mechanisms included its distinct song, feeding and nesting behaviour, and its very different habitat. They found it very unlikely that the two birds would have been able to interbreed, let alone freely, and considered it likely that future researchers would split them.[24] A 2004 phylogenetic analysis by James based on osteological features found Himatione to group in a clade similar to what Perkins suggested in 1903. Her "clade 11" is depicted in the phylogram below († denotes recent extinctions, ‡ denotes prehistoric):[26]

The 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 Drepanidae or Drepaniidae, which turned out to be preoccupied by a family of moths[15]), subfamily Drepanidinae, or tribe Drepaninini, were long recognized as constituting a natural group of finches with varying bill-shapes and plumage patterns as a result of adapting to island environments. Their relationships to other finches remained uncertain, and they were often considered a distinct lineage outside the Fringillidae. By the turn of the 21th century, genetic studies had established that drepanids were genetically close to Carduelinae, with studies from 2011 and 2012 finding them nested within that group as the sister group of the genus Carpodacus from Asia (neither study included the Laysan honeycreeper itself). This indicates the Hawaiian honeycreepers originated in Asia, and divergence was estimated to roughly coincide with when the oldest of the Hawaiian islands formed, about 5.7 million years ago, with further divergence occurring as other islands formed. It was also proposed that Drepanidinae should be treated as a junior synonym of Carduelinae.[27][28][29] Since the ʻapapane had been found to be the sister taxon of the ʻākohekohe (Palmeria dolei) by some studies, Pratt suggested in 2014 that their genera Himatione and Palmeria might be merged.[30]

Description[edit]

Illustration of an adult male Laysan honeycreeper (A), an adult female (B), and juvenile (C), and an 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).[31][20][6] The wing measured 65–69 mm (2.55–2.7 in), the tail 61 mm (2.4 in), the culmen (upper surface of the beak) 14 mm (0.55 in), and the tarsometatarsus (lower leg bone) was 23 mm (0.9 in).[11] It was bright scarlet vermilion with a faint tint of golden orange on the head, breast and upper abdomen, while the rest of its upper parts were orange scarlet. The lower abdomen was dusky gray that faded into brownish white, and the under-tail covert feathers were grayish. The wings, tail, bill, and legs were dark brown, while 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.[31] The bill was slender and downturned.[32]

The sexes were alike, though the bill, wings, and tail were slightly shorter in the female. While Rothschild stated in his 1892 description that the female was paler than the male, Fisher indicated in 1903 that such differences may have been age-related instead. Fisher also noted that the illustration of the Laysan honeycreeper published by Rothschild showed the bird as far too pale, giving an inaccurate idea of its color.[33][6] Rothschild also mentioned in his 1893–1900 work that freshly 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.[11]

The ʻapapane differs from the Laysan honeycreeper in being blood-red overall, with black wings and tail, whiter under-tail covert feathers, and a longer bill.[31] The two also differ in that the Laysan species had a shorter, stouter bill, and in that its primary feathers did not have oblique truncation (in the ʻapapane this truncation produces a "wing note" sounding like vocalization).[9] 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 difference in plumage of the Laysan honeycreeper was due to fading caused by the intense sunlight, but pointed out it had been found to be distinct in osteological features.[25][34] Pratt countered in 2005 that white undertail feathers cannot fade to brown since they lack pigmentation to begin with.[9]

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 breeding-season,and was in "full song" during January and February. While catching and skinning birds in 1891, Palmer caught a Laysan honeycreeper in his net, which proceeded to sing in his hand; he answered it with a whistle, which it returned, continuing for some minutes without seeming frightened.[11][19] The American ornithologist Alexander Wetmore noted that their "charming song is out of proportion to their size" in 1923.[1]

Habitat[edit]

1912 map of Laysan (left) and 2010 aerial photo

The Laysan honeycreeper was endemic to Laysan, a remote island which has a total land area of 3.6 km2 (1.4 sq mi), and is the largest of the northwestern Hawaiian Islands inn the central Pacific Ocean. Laysan is the eroded remnant of a once high island, built up by volcanic activity, perhaps the flattened top of a volcano that formed in the Miocene. The island is roughly triangular, and rises into up to 12 m (40 ft) high crest elevations. Its subsurface substratum is coralline rock, and its topography suggests it was once part of an atoll with a lagoon which 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 species, though the extent of the vegetation had almost recovered by 1973.[19][4][31]

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 land-birds appeared to congregate. This was also its favored nesting area, with its broad patches of the succulent Portulaca that these birds fed from. Munro added that it also frequented grass tops 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.[33][35][36] It was the only nectar-feeding finch of the northwestern Hawaiian Islands.[3]

Behavior and ecology[edit]

The flower maiapilo which the Laysan honeycreeper fed nectar from

Few naturalists personally encountered the bird, and few accounts were left of its life history. It was very active, like the Laysan millerbird, always present in vegetation around buildings, and while perhaps less confiding than the millerbird, they were reported to sometimes enter buildings for moths and for roosting at night. Unlike the ʻapapane, it also foraged on the ground.[32][19][37] Palmer stated he generally saw them in pairs.[11] Munro described its flight as "weak and low". He stated they would gather around houses and drink rain water from leaks in barrels, indicating that they "missed water" more than the other birds of the island. He speculated they had perhaps not adapted to the meager supply of water there; apart from rain and dew, the only natural moisture was a seep of brackish water.[36]

The Laysan honeycreeper was nectarivorous and insectivorous, and insects were probably a more important part of their diet than nectar in some seasons. The bird originally fed on nectar from the native flower maiapilo (Capparis sandwichiana), but when that species disappeared, it switched to the ʻākulikuli (Sesuvium portulacastrum) and ʻihi (Portulaca lutea).[19] It was also observed visiting nohu (Tribulus cistoides) and pōhuehue (Ipomoea pes-caprae).[34][36] The Laysan honeycreeper spent the day foraging while walking like pipits after small insects or drinking from flowers with its brush-like tongue. The way it rapidly went from flower to flower and precisely inserted its bill between their petals reminded Schauinsland and Fisher of hummingbirds, though it did so by walking rather than hovering in front of them.[33][35][12][19] The Laysan honeycreeper gathered insects from flowers, such as small, green caterpillars, and like the other insect-eating birds were fond of large, brownish moths called millers, which were abundant on the island. Fisher observed honeycreepers extracting moths from between boards, grasping them with one foot while eating the soft parts.[33][35][36]

Reproduction[edit]

Photo of a nest, by Walter K. Fisher, 1902

Fisher noted the nest of the Laysan honeycreeper was more difficult to find than that of the Laysan millerbird, and only one that contained a single egg was found, in the middle of a grass tuft about 61 cm (2 ft) above ground. Schauinsland noted it also nested in thick aweoweo (Chenopodium oahuense) shrubs. The nest was made of fine grass (identified as the species kawelu (Eragrostis variabilis) by Pratt based on Fisher's photo of the nest[9]) and rootlets with some dry grass (variously described as loosely or well built), and its bowl measured about 5.7 cm (2.25 in) across and 4.13 cm (1.625 in) in depth. It was lined with fine rootlets, petrel feathers, and brown down from Laysan 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, and the nests of different land birds of the island may have been similar due to the limited building materials.[33][35][11][12]

Little is known about the breeding cycle of the Laysan honeycreeper, and most observers did not record when nests and young were found.[19][11] Freeth told Palmer that the bird was in "full song" in January and February, when there was also a golden gloss over the red plumage. This indicates that the breeding season was between that time and June, when Palmer saw full-grown juveniles.[11] Fisher collected a nest with an egg in mid-may, while the American zoologist William Alanson Bryan collected an egg on May 10. Bailey stated the clutch size was four or five eggs, while sets of three were taken by various collectors.[19] The ovate eggs were glossless white, with grayish blotches and spots at the larger end, and reddish brown spots above these, these markings often forming circles. The eggs varied in size, but a typical egg measured 18 by 13.75 mm (0.709 by 0.541 in). The eggs were similar to those of the short-toed treecreeper and the barn swallow, but much less glossy.[33][35][11]

Extinction[edit]

The collector Henry C. Palmer among frigatebirds on Laysan in the early 1890s, by Frohawk, 1893–1900

The few observations of the Laysan honeycreeper indicate it was not abundant on Laysan to begin with, and only three gave population estimates. Isenbeck already considered the bird uncommon when visiting in 1828, while Palmer considered it the rarest of the island's birds in 1890, though finding them in fair numbers, and Palmer gave a similar status in 1903.[19] The American zoologist Charles C. Nutting stated in 1903 that the species of Laysan were abundant, and that there were excellent conditions for collecting and studying birds. He suggested that when guano supplies ran out, Laysan should become a government preserve for bird life, protected by human-made destruction.[38] Laysan was exploited for the guano produced by its large sea bird colonies from 1890, but this became unprofitable by 1904. The German superintendent of the guano operation, Max Schlemmer, then introduced domestic rabbits and European hares to the island to start a meat-canning business that would provide food for guano miners. This did not succeed, but the rabbits instead proceeded to destroy the island's vegetation.[31][34][37]

In 1909, the US president Theodore Roosevelt issued an executive order that made several Hawaiian islets and reefs (including Laysan) part of the Hawaiian Islands Reservation. During a 1911 expedition to assess the condition of bird life on Laysan, the American ornithologist Homer R. Dill and Bryan found rabbits everywhere, and foresaw that there would be no vegetation left if drastic measures were not taken. They found thousands of bird skeletons left over by feather hunters, as well as several Laysan honeycreeper skins, but noted the birds were still fearless towards humans after this wholesale slaughter. They estimated that 300 Laysan honeycreepers remained, and that it and other birds there were "doomed to extermination" if their food supply was not preserved.[37] In 1915, the American First Leutenant William H. Munter reported that the Laysan honeyeter was fairly common, and that they were judged to number a 1000.[39] 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 next year.[19]

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

The bird was filmed in 1923 during the Tanager Expedition. Shortly after, Laysan was battered by a strong storm, and later attempts at finding any remaining Laysan honeycreeper failed.[40] Other birds also inhabited the island, including the Laysan millerbird, the Laysan rail, the Laysan duck, and the Laysan finch. Of these, only the finch and the duck remain extant.[41]

References[edit]

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  32. ^ a b Bailey, Alfred M. (1956). "Birds of Midway and Laysan Islands". Denver Museum of Natural History Museum Pictorial. 12: 119–122.
  33. ^ a b c d e f Fisher, Walter K. (1903). "Birds of Laysan and the Leeward Islands, Hawaiian group". Bulletin of the United States Fish Commission. 23 (3): 803–804.
  34. ^ a b c Fancy, Steven G.; C. John Ralph (1997). "ʻApapane". In A. Poole; F. Gill (eds.). Birds of North America. Vol. 296. Academy of Natural Sciences. doi:10.2173/bna.296.
  35. ^ a b c d e Fisher, Walter K. (1903). "Notes on the Birds Peculiar to Laysan Island, Hawaiian Group". The Auk. 20 (4): 384–397. doi:10.2307/4069753. JSTOR 4069753.
  36. ^ a b c d Munro, George C. (1971). Birds of Hawaii. Rutland: Tuttle. pp. 84–85. ISBN 978-0804800631.
  37. ^ a b c Bryan, William Alanson; Dill, Homer R. (1912). "Report of an expedition to Laysan Island in 1911: under the joint auspices of the United States Department of Agriculture and University of Iowa". U.S. Department of Agriculture, Biological Survey. 42: 3–30. doi:10.5962/bhl.title.62940.
  38. ^ Nutting, Charles Cleveland (1903). "The Bird Rookeries on the Island of Laysan". The Popular Science Monthly. 63: 321–332.
  39. ^ Munter, William H. (1915). "Report of destruction of bird life on Laysan Island, Pacific Ocean". Annual Report of the United States Coast Guard: 130–140.
  40. ^ "1923 USS Tanager Expedition". Video. Northwestern Hawaiian Islands Mult-Agency Education Project. Archived from the original on 2008-09-29. Retrieved 2009-03-16.
  41. ^ Wetmore, Alexander (1925). "Bird life among lava rock and coral sand". National Geographic Magazine. 48: 77–108.
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