THE WILSON BULLETIN A QUARTERLY JOURNAL OF ORNITHOLOGY Published by the Wilson Ornithological Society

VOL. 110, NO. 1 MARCH 1998 PAGES 1-154

Wilson Bulletin, 110(l), 1998, pp. l-22

REFLECTIONS ON A 1975 EXPEDITION TO THE LOST WORLD OF THE ALAKAI‘ AND OTHER NOTES ON THE NATURAL HISTORY, SYSTEMATICS, AND CONSERVATION OF ‘

SHEILA CONANT1,4, H. DOUGLAS PRATT2, AND ROBERT J. SHALLENBERGER3

ABSTRACT.-We observed, tape recorded, and photographed birds of the Alakai‘ Plateau on Kauai,‘ ‘ for one week during the summer of 1975. We observed all but one of the islands’ historically known species and compared the Alakai‘ Plateau with the more accessible Kokee‘ area. Ours were the last studies before catastrophic changes in the Kauai‘ avifauna and included many observations that cannot now be repeated. This retrospective report presents our findings in the light of subsequent events. Because our Alakai‘ studies were seminal in the development of the current AOU classification of Hawaiian native , we defend that classification against recent challenges and further refine it. The controversial genus is shown to be supported by a suite of synapomorphies of plumage, bill morphology, and vocalizations. We advocate removal of the Anianiau‘ from Hemignathus and classify it as Magumma parva. Our studies of foraging behavior and vocalizations support the recent recognition of the Kauai‘ Amakihi‘ (H. kauaiensis) as a separate species and suggest that the Elepaio‘ (Chasiempis) is best split into three species (sckzteri, ibidis, and sandwichensis). Major hurricanes in 1983 and 1992 appear to have severely impacted Alakai‘ populations with the subsequent of the Kauai‘ 0‘ 0‘ (Moho braccatus) and possibly the Kamao‘ (Myadestes myadestinus), and the island population of O‘ u‘ (Psittirostrapsittacea). We report some of the last natural history observations on these species. Formerly adaptive strategies for storm survival, including taking refuge in valleys, are no longer effective because the lowlands are now infested with mosquito-borne avian diseases. The Puaiohi (M. palmen),’ a ravine specialist, suffered less from the storms although its population remains perilously low. Other forest birds, especially the Akikiki‘ (Oreomystis bairdi), show noticeable declines since 1975. We speculate that intro- duced organisms such as alien plants can have a deleterious effect on ecosystems by altering feeding methods of birds even in areas where the weeds do not occur. We caution against the overly conservative use of species- level taxa for setting conservation priorities on remote islands. Received 24 October 19%, accepted 27 June 1997.

In July 1975 we spent a week in one of fortunate to find all but one of Kauai‘ s’ his- Hawaii‘ s’ last strongholds for native birds, the torically known native bird species. At that fabled Alakai‘ Swamp on Kauai.‘ We were time, we assumed that this relatively large wilderness area, under no obvious threat, ’ ’ Department of Zoology, University of Hawaii at would remain a refuge for endangered birds Manoa, 2538 The Mall, Honolulu, HI 96822; E-mail: at least for the near future. Thus, although our [email protected] observations contributed to subsequent publi- z Museum of Natural Science, Louisiana States Uni- cations, we published no general summary of versity, Baton Rouge, LA 70803-3216 our expedition. Time has shown that our op- 3 Midway Atoll National Wildlife Refuge, c/o U. S. Fish and Wildlife Service, PO. Box 1, Midway Atoll, timism was naive; we made some of the last FPO/AP 965 16 observations and tape recordings, and took 4 Corresponding author. some of the last (or only) photographs, of sev-

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FRONTISPIECE. Upper 4 and lower left 2: Now extinct Kaua‘i ‘0‘0 (M&o braccarus) in various attitudes. Lower right 2: The nearly extinct ‘0’~ (Psittirostra psirracea) in characteristic postures. Photographed July 1975 in the Alaka‘i Swamp, Kaua‘i by Robert J. Shallenberger except lower right by H. Douglas Pratt. 2 THE WILSON BULLETIN. Vol. 110, No. 1, March I998 era1 species, and made the last natural history Endangered Species (T. Pratt, pers. comm.). In studies of several others under anything re- view of these unexpectedly rapid changes, and sembling “normal” conditions. We herein re- because the Biological Resources Division, port numerous observations that will likely U.S. Geological Survey, has initiated a new never be repeated and make comparisons with research program to study Kauai‘ s’ endan- the current status of birds on Kauai.‘ We in- gered birds as the century draws to a close (T. terpret and amplify our 1975 observations Snetsinger, pers. comm.), a report of our 1975 with subsequent observations on Kauai‘ birds work is timely. made by the authors individually and with rel- evant unpublished observations of other nat- STUDY AREA AND METHODS uralists that they have generously shared with Our primary study site was within what is now the us. Alaka‘i Wilderness Preserve. We were airlifted by hel- We also discuss the implications of our ob- icopter to headwaters of the Halehaha Stream, near one servations for ongoing systematic controver- of the late John L. Sincock’s (Harrison et al. 1992) sies, including some previously unpublished study areas we call “Sincock’s Bog” (Fig. l), late in supporting details for Pratts’ (1979) revision the afternoon of 1 July 1975. Our flight was hampered by low-hanging clouds and fog, but the next morning of the Hawaiian honeycreepers, the basis for dawned bright and clear, and such uncharacteristically the classification in widest use today (Amer- dry weather prevailed for the rest of our stay until just ican Ornithologists ’ Union 1983, Sibley and before our departure on 8 July, when clouds and fog Monroe 1990, Tarr and Fleischer 1995). We again closed in. Each night produced heavy dew. have made some refinements of Pratts’ (1979) RJS used a Nagra portable tape recorder and a Dan classification, which was “eclectic” (sensu Gibson parabola; HDP used a Sony cassette recorder and an 46 cm Sony PBR-400 parabola. All recordings Raikow 1985) but based on a cladistic anal- were later archived at the Library of Natural Sounds ysis, to make it better reflect phylogeny. (LNS), Cornell Laboratory of Ornithology, and some Only a few major scientific studies of have been published (Hardy and Parker 1985; Pratt Kauai‘ forest birds followed our expedition: 1996). SC used binoculars and a stopwatch to record the 1981 survey made as the final part of the activity patterns and behaviors of several species. All landmark U.S. Fish and Wildlife Service pop- 3 authors took numerous color photographs of birds, plants, and the habitat as 35 mm transparencies. A ulation study (Scott et al. 1986), various stud- small series of specimens collected by HDP was de- ies by the U.S. Forest Service (C. J. Ralph and posited in the Museum of Natural Science, Louisiana colleagues, unpubl. data), and several surveys State University. by other state and federal wildlife agencies The misnamed Alaka‘i Swamp is actually a wet (e.g., Engilis and Pratt 1989; Pyle 1990, 1994; montane plateau mostly above 1000 m elevation Telfer 1993). Several species declined notice- spreading northwest from Mt. Wai‘ale‘ale, often con- ably between 1975 and 1981, and Hurricanes sidered the world’s wettest spot, with mean annual rainfall of 1415 cm or 449 in. (Giambelluca et al. Iwa in 1982 and Iniki in 1992 severely re- 1986). The plateau is deeply dissected by numerous duced remaining bird populations in the forested ravines or canyons that feed into highly erod- Alakai‘ (Engilis and Pratt 1989; Pratt 1994a; ed Waimea Canyon, which drains to the south. The Pyle 1983, 1993b) apparently because the ridges between the streams can be broad and nearly montane forest birds ’ historic habit of riding flat, pocked in many places by open bogs (probably out severe storms in lowland valleys is now the origin of the “swamp” designation). maladaptive, the valleys being infested with Our study area included a small (approximately 4 ha), open bog where the helicopter landed (Fig. 1). The disease-carrying mosquitoes (Atkinson et al. surrounding dense rainforest was dominated through- 1993, Pyle 1993a, Pratt 1994a). Today, the en- out by ‘ohi‘a-lehua (Metrosideros collina). ‘Ohi‘a demic Kauai‘ 0‘ 0‘ and Kamao‘ (see species characteristically forms a dense canopy with numerous accounts) may both be extinct, and the O‘ u‘ emergent snags (Fig. 2) that provide display perches has not been certainly reported since Hurri- for several native birds. The bright red brush-like cane Iniki in 1992 despite diligent searches by ‘ohi‘a flowers, which lack petals, are the primary nec- competent biologists. Among other island en- tar source for several honeycreepers. The nectar col- lects in a cuplike calyx at the base of the red stamens demics, the Kauai‘ Nukupuu‘ barely survives so that nectarivorous birds must insert the bill deep (J. Jeffrey, pers. comm.), the Puaiohi is very into the inflorescence. The forest in the study area was rare (Pyle 1994; T. Snetsinger, pers comm.), very dense even at ground level and heavily over- and the Akikiki‘ is proposed for listing as an grown with epiphytes. The epiphytic community in- Crwurnr r, trl. * ALAKA’I BIRDS 3

FIG. 1. Sincock’s Bog as viewed from the air (upper)and from ground level (lower) with HDP in the foreground. Photos by RJS. THE WILSON BULLETIN l Vol. 110. No. 1, March 1998

FIG. 2. Profile of forest canopy in the study area showing emergent ‘ohi‘a snags. Photo by HDP chides abundant mosses and ferns as well as flowering lost none of its historically known bird species plants. (Richardson and Bowles 1964) and the Below the bog to the south the terrain sloped sharply Alakai‘ Plateau remained a seldom-visited to a rushing stream. In places, the banks were so steep that large ‘ohi‘a trees leaned over the stream almost wilderness. Birds in general were abundant in horizontally. In other places, the streambed widened our study area, and native species far outnum- into relatively flat amphitheater-like areas that sup- bered aliens. Most native birds appeared to ported the growth of grass as well as small trees and have just finished breeding. We saw few fledg- shrubs. Such areas had obviously been scoured by de- lings, but recently fledged immatures of sev- bris during times of high water. With the help of Dr. eral species were numerous. Except for the C. H. Lamoureux, who worked from our photographs, Kauai‘ 0‘ 0,‘ the Kamao,‘ and possibly the we were able to identify many, but not all, of the dom- inant or common plants in the study area (Table 1). O‘ u,‘ the passerines we observed did not ap- For comparative purposes, we made a short visit to pear to be defending breeding territories. Koke‘e State Park, an area just west of the Alaka‘i Rather they ranged widely through the area; swamp, immediately prior to our expedition. The park the species noted near our campsite changed area differs from the heart of the Alaka‘i in having on a daily basis. Also, few species were sing- much more koa (Acacia koa) among the canopy trees ing territorial songs; we recorded mostly call and in the presence of numerous alien plants and birds. The Koke‘e region is served by many roads and trails notes. Probably the best season for observing and we have returned to it individually many times in breeding activities in the Alakai‘ would be the ensuing decades to monitor changes in Kaua‘i’s April through June, although the peak of nest- avian community. We mostly visited areas served by ing may vary from year to year. the Pihea Trail, Camp 10 (Mohihi) Road, and the Alak- ai Swamp Trail to the edge of the Alakai Wddemess SPECIES ACCOUNTS Preserve (for maps and details see Pratt 1993). The following codes are used in this sec- THE AVIAN COMMUNITY tion: RE = resident, endemic Hawaiian spe- Through the 1960s Kauai‘ was considered cies; RK = resident, species endemic to unique among the Hawaiian Islands in having Kauai‘ Island; RI = resident indigenous Conant et al. l ALAKA‘I BIRDS 5

TABLE 1. Major plant species identified in the study area, listed alphabetically within different forest strata. (Plant names follow Wagner et al. 1990).

Scientific name Hawaiian name FlOW.% Fruit

Trees Cheirodendron sp. ‘olapa + Metrosideros collina ‘ohi‘a-lehua + Myrsine sp. kolea + + Pelea sp. ‘alani + Tetraplasandra sp. ‘ohe + Shrubs Broussaisia arguta kanawao + Cibotium sp. hapu ‘u no notes Clermontia spp. ‘oha Dubautia paleata na‘ena‘e puakea + Rubus hawaiiensis ‘akala Scaevola glabra ‘ohe naupaka + Vaccinium calycinum ‘ohelo kau la‘au Vaccinium dentatum ‘ohelo no notes Herbs and Epiphytes Adenophorus sp. unknown no notes Astelia sp. pa‘iniu + Carex sp. unknown no notes Dianella sandwichensis uki‘ uki‘ + + Drosera anglica mikinalo + ‘ie‘ie no notes Korthacella remyana var. wawrae hulumoa + + Lycopodium cernum wawae ‘iole + Lycopodium serratum unknown + Nertera granadensis unknown + Psilotum sp. moa no notes Smilax sandwichensis hoi kuahiwi + Viola kauaensis poke hiwa + +

breeding species; A = alien breeding species. centrate in lowland wetlands such as the taro Federally recognized Endangered Species are fields at Hanalei (Pratt 1993), they also nest noted as such. in the mountains (Berger 1981). White-tailed Tropicbird or Koa e-kea‘ Red Junglefowl or Moa (Gallus gallus), (Phaethon lepturus dorotheae), RI.--We saw A.-HDP flushed one bird from the ground in two individuals flying over bog at about 13:OO the low forest north of the bog on 4 July, and 8 July, as we awaited our helicopter pickup. found an abandoned large nest on the ground Tropicbirds are numerous in canyons and val- in the same area. We did not hear any dawn leys throughout Kauai‘ and nest in Hanapepe “crows” from our campsite. The unusually Valley south of our study area (HDP pers. large numbers of these birds that congregate obs.). around buildings in Kokee‘ State Park are ap- Hawaiian Duck or Koloa (Anas wyvilliana), parently lured there and maintained by food RE, endangered.-RJS flushed one individual handouts from visitors. from the stream bed just below camp. The Short-eared Owl or Pueo (Asio flammeus), bird was in a wide part (about 15 m) of the RI.-Solitary individuals were observed soar- stream where it was probably standing on a ing, possibly hunting, a few meters above the rock or in a dense growth of sedge (Carex tree tops once or twice each day, usually in sp.). The duck is also present in low numbers the afternoon or early evening. The owl is also and may nest along Kawaikoi Stream in the uncommon in the Kokee‘ region. Although Kokee‘ area. Although Hawaiian Ducks con- historically regarded as an endemic subspe- THE WILSON BULLETIN l Vol. 110, No. I, March 1998

been thoroughly investigated, but the reper- toire is qualitatively distinguishable from those of the ‘ and Hawaii‘ forms (Pratt 1996). E. VanderWerf (pers. comm.) recently found that Oahu‘ Elepaio‘ respond differen- tially to playback of their own vocalizations and those of Hawaii‘ birds. Similar studies with the Kauai‘ Elepaio‘ are ongoing. All ele-‘ paio are bold and inquisitive, but the Kauai‘ birds are particularly so, the juveniles more than adults. Juvenile Kauai‘ Elepaio‘ ap- proach humans closer, sometimes perching within arms’ reach, and follow them farther FIG. 3. Juvenile Kaua‘i ‘Elepaio. Photo by HDP than do elepaio‘ on other islands. Both age categories respond readily to imitations of their whistled songs and calls as well as ties, this cosmopolitan species may be a post- “spishing.” We saw several fledglings includ- Polynesian colonizer of the Hawaiian Islands ing one whose rectrices were only l-2 cm (Olson and James 1991). long. This suggests that the breeding season Kaua i‘ Elepaio‘ (Chasiempis sclateri), ends about the same time as that of the Oahu‘ RI.-In 1975, only one species of elepaio‘ Elepaio‘ (Conant 1977, Sherwood 1995), but was recognized, with subspecies on Kauai‘ earlier than that of the Hawaii‘ representative (C. sandwichensis sclateri), Oahu‘ (C. s. gayi, (Bollinger and Bollinger 1987). later emended to C. s. ibidis by Olson 1989a), Ecologically, all three elepaio‘ species are and Hawaii‘ (C. s. sandwichensis). Subse- forest birds, but as their islands differ, so do quently, the Hawaii‘ form was shown to ex- the birds ’ respective habitat preferences. We hibit inn-a-island variation at the subspecies observed both adult and immature Kauai‘ level, with at least three forms recognizable Elepaio‘ frequently every day in our study (Pratt 1980). The striking plumage differences area, and found them also common at Kokee.‘ of the Kauai‘ and Oahu‘ forms clearly qualify They are restricted to the higher and wetter them as “megasubspecies” (sensu Amadon parts of the island (Scott et al. 1986) and have and Short 1976), and they are unquestionably not undergone any obvious decline since 1975 “phylogenetic” species (sensu Cracraft 1983). (Engilis and Pratt 1989, Walther 1995). In Olson and James (1991; see footnote to Table contrast, the Oahu‘ Elepaio‘ has declined 1) recognized three species of Chasiempis, but strikingly during the same period (Williams whether they consider them biological or phy- 1987, Pratt 1994a, Conant 1995) and is now logenetic species is unclear. rare, with a fragmented distribution restricted Our work on Kauai‘ combined with obser- primarily to relatively mesic valleys at middle vations on other islands (Conant 1977, Pratt elevations (VanderWerf et al. 1997). It has ap- 1980) supports the recognition of three bio- parently always been more common in drier, logical species of Chasiempis. The adult mid-elevation forests (Bryan 1905). On Ha- Kauai‘ Elepaio‘ is the plainest and the only waii,‘ the intra-island variation in plumage one with a gray dorsum. Juveniles are mostly color is correlated with rainfall (Pratt 1980). rufous, without the white rump and wing-bars Apparently the three Chasiempis were first of the adults, and closely resemble juveniles “lumped” by Bryan and Greenway (1944), of the Oahu‘ form in color but seem subjec- who gave no data to support the change in tively to be subtly different (more rotund, less status; and subsequent authors, until Olson linear) in shape (Fig. 3). Plumage dimorphism and James (1991), accepted that by is lacking in adults on Kauai‘ and Oahu,‘ but default. We believe a return to recognition of Hawaii‘ adults are strongly dichromatic (Pratt three species, with C. sandwichensis having 1980, Pratt et al. 1987). three intra-island subspecies, provides a more The variations in tempo and pitch of the meaningful and balanced taxonomic treatment vocalizations of the Kauai‘ Elepaio‘ have not that better reflects the observed geographic Conant et al. l ALAKAI‘ BIRDS 7 variation. Because the three forms are allo- able” sightings, but no birds were heard (Pyle patric, biological species limits must be as- 1989). That report noted that on a 1985 DFW sessed by inference. Our observations, as well survey many Kamao‘ were heard. DFW again as those of VanderWerf (in press), show that surveyed the Alakai‘ Swamp in 1993, and two the striking plumage differences are backed “brief but inadequate” possible sightings of up by potential behavioral and ecological iso- Kamao‘ were reported (Pyle 1993b). lating mechanisms. We can discern no reason, Puaiohi (Myadestes palmeri), RK, endan- other than the general similarities reflected at gered.-Our first sighting of this species oc- the generic level, to consider them conspecif- curred at dusk on 3 July. A single bird flew ic. from the forest underbrush and perched si- Kama o‘ (Myadestes myadestinus), RK, en- lently above our camp site. In the fading light dangered, possibly extinct.-This larger of the we could see the bird was a heavily spotted two Kauai‘ solitaires (Pratt 1982), then called immature, with a few down feathers still the Large Kauai‘ Thrush, was not uncommon clinging to the head. The pink feet were evi- in the study area during our visit. As many as dent as the bird perched upright facing us. seven individuals could be seen simultaneous- Later, HDP saw two other individuals, both ly perched on emergent ohi‘ a‘ snags in the adults. One was located by slowly searching forest surrounding the bog above our camp in out the source of a dry, toneless hiss that the early morning and at dusk. They perched proved to be the call note. The bird was upright with the tail down, From inside the perched almost motionless on a mossy stump forest, Kamao‘ were difficult to see, but we under a concealing clump of vines and shrubs heard their songs and calls throughout the day at the upper rim of a small ravine. In posture, in the vicinity of the campsite. They gave two this individual resembled the bird seen pre- different calls, one a cat-like rasp and the oth- viously, as it sat bolt upright rather in the er resembling the sound of a police whistle. manner of a tyrannid flycatcher. It remained These calls are similar to, but distinguishable for some minutes in this position, uttering its from, calls of the Oma‘ o‘ (M. obscurus) of calls at about 10-s intervals. It then flew to Hawaii.‘ The song of the Kamao‘ is entirely the end of a small stump that extended from different from that of the Hawaii‘ bird and a high bank over the stream. Again, the bird resembles closely the song of Townsends’ Sol- remained motionless for some minutes and itaire (Myadestes townsendi) of western North then flew away into the forest. The third in- America, with some of the tonal quality of the dividual was glimpsed briefly as it flushed song of the Slate-colored Solitaire (M. occi- from concealment in a small (ca. 5 m) olapa‘ dentalis) of Mexico. These song differences (Cheirodendron trigynum) tree that grew in a were among early indications that the Kamao‘ wide, flat area of the streambed (Fig. 4). These and Oma‘ o‘ were separate species, later con- observations demonstrate that this species is firmed by Pratts’ (1982) playback experiments extremely difficult to detect where it occurs, on Hawaii‘ using recordings made during our and thus may have been more common in the Alakai‘ visit. The song of the Kamao‘ is long Alakai‘ than the few published observations and complex with an ethereal, echoing, and would indicate. All of our sightings were in cascading quality. A typical singing perfor- or near a deep, protected stream valley, as mance ended with the bird rising on its wings, have been other observations (Ashman et al. hovering over the forest while singing con- 1984, Kepler and Kepler 1983, Pyle 1984), stantly, and then abruptly diving into the trees. whereas most of the Kamao‘ we observed Pratt (1982) presented sonagrams of the two were near the more exposed ridge-top bog. call types and the song, and recordings of all The smaller solitaire appears to be a ravine three have now been published (Pratt 1996). specialist. This may partly explain why the What was once Kauai‘ s’ most abundant forest Puaiohi, although historically much the rarer bird (Munro 1944) may well be extinct; it has of the two Kauai‘ thrushes (Perkins 1903), has not been seen since Hurricane Iniki (Walther weathered the two recent hurricanes much 1995; T. Snetsinger, pers. comm.). A February better than its larger relative. A very small 1989 survey by the Hawaii Division of For- population of Puaiohi persists in the Alakai‘ estry and Wildlife (DFW) reported two “prob- (T Snetsinger, unpubl. data) and is the subject THE WILSON BULLETIN * Vol. 110, No. I, Much 1998

FIG. 4. An open area of stream bed on Halehaha Stream where HDP flushed a Puaiohi. Photo by HDF?

of intense observation and possible recovery This has become the most actions by federal agencies and The Peregrine abundant forest bird on Kaua‘i (HDP, pers. Fund. These ongoing studies have already dis- obs.; Scott et al. 1986). covered a larger than expected population Kaua i‘ 0‘ o‘ or 0‘ o‘ a‘ a‘ (Moho braccatus), (Pyle 1996) in the Koai‘e Stream area (almost RK, endangered, possibly extinct.-On 2 July directly between Koke‘e and our 1975 study we awoke to the song of the ‘0‘0 (06: 15). site) where two nests were found about a de- Shortly after the dawn chorus ceased and the cade ago (Ashman et al. 1984, Kepler and Ke- sun was up, a single ‘0’0 appeared near our pler 1983). campsite and provided our first look. Interest- Melodious Laughing-thrush (Garrulax can- ingly, the bird did not visit us on subsequent orus), A.-RJS briefly saw one bird in low, mornings, indicating that it may have deviated dense vegetation above the campsite. On from its usual feeding route to “check us out” Kaua‘i, this bird is found in dense mesic and that first day. The song (for recordings see wet forests of both native and introduced Pratt 1996) is a series of loud, clear melodious plants. It is very secretive and almost always whistles with a “tune” reminiscent of the seen near the ground. Its loud song is the best song of the Western Meadowlark (Sturnella indication of its presence. Although we have neglecta). Its tone quality is distinctly meli- no record of vocalizations in our field notes, phagine, however, with a haunting, echoing HDP later noticed the song in the background quality, and is surprisingly similar to portions of one of his tapes. of the song of the Tui (Prosthemadera novae- Japanese White-eye (Zosterops japonicus), seelandiaee) of New Zealand (for an example A.-White-eyes were present in the area but of the latter, see Gunn and Gulledge 1977). were greatly outnumbered by native birds. We Other portions of the Tui song sound remark- saw only a few small groups. In contrast, they ably like the unrelated drepanidine ‘I‘iwi (dis- were common to abundant in the Koke‘e area. cussed below). The Kaua‘i ‘0‘0’s song is Conant et al. - ALAKAI‘ BIRDS 9

TABLE 2. Percentage of time devoted to 7 activ- that they patrolled systematically because they ities of 2 Kauai‘ 0‘ 0‘ during 61 minutes and 36 sec- would leave this area for 20-40 min at a time, onds of observation. usually flying off to the northeast and retum- ing northwest. This foraging pattern is consis- Activity Percentage of time tent with observations of “trap-line” feeding Feeding at flowers 55 in East . There, both the I‘ iwi‘ and the Inactive perching 29 Akohekohe‘ (Palmeria dolei) revisit blooming Preening 9 Flying or hopping 3 ohi‘ a‘ trees with a regular cyclicity of about Singing 3 20-35 min (HDP pers. obs.). Probably the cy- Insect gleaning”

TABLE 3. Foraging behavior of two Kauai‘ 0‘ ‘ during 47 minutes and 11 seconds of observation (data recorded only while birds were feeding).

Vertical canopy occupation Horizontal canopy occupation

Height (m) Percent time ZOX” Percent time

2-4 18 Inner 14 4-7 11 Center 7 71 Outer 86 Movement Patterns During Feeding Percent time moving 57 Percent time stopped (feeding pause) 43 Rate of feeding pauses 3.9 pauseslmin Rate of movement (not including feeding pauses) 2.1 m/min Rate of movement (including feeding pauses) 1.2 m/min Average distance between feeding pauses 0.3 m

a Canopy zones: Inner = within I mof the central axis of tree; Outer = within I mof outermost crown foliage; Center = between inner and outer. 10 THE WILSON BULLETIN l Vol. 110, No. I, March I998

FIG. 5. ‘0% (Psitrirostra psirtacea) male (1.) and female (r.). Photos by RJS. See also Frontispiece.

Krakow&i, who heard an ‘0‘0 calling on 28 their wings clumsily. The bill was open during and 29 April 1987 (Pyle 1987). A survey by singing and the body quivered slightly. state agency biologists in February of 1989 All the birds we saw sat quietly or moved found no ‘0‘0, and the report speculated that slowly and heavily in upper portions of dead the birds were “now probably gone” (Pyle snags or on high exposed perches. As a result, 1989). They have not been reported since and most of our photographs, which appear to be are undoubtedly extinct, their loud vocaliza- the only ones ever taken of this species, are tions being difficult to overlook. strongly back-lighted and fail to show the Northern Cardinal (Cardinalis cardinalis), birds’ colors to good effect. Their perching A.-SC heard one bird singing the morning of posture was distinctively horizontal, with the 8 July for our only observation. Northern tail held slightly up but not cocked. They ap- Cardinals were and remain common to abun- peared rather pot-bellied (Fig. 5, Frontis- dant at Koke‘e, where the forest is more dis- piece). Although we spent several hours turbed. In general, they are one of the more watching various individuals, HDP saw only widespread introduced birds throughout the one bird actually feed, apparently taking nec- Hawaiian Islands, but are rare in primary for- tar from an ‘ohi‘a flower. est (Pratt et al. 1987). A Kaua‘i ‘O‘u population in the low hun- ‘0 ‘u (Psittirostra psittacea), RE, possibly dreds in the late 1970s was apparently deci- extirpated.-We located several pairs of ‘O‘u mated by Hurricane Iwa in 1982 (Pyle 1983, near our campsite (we estimated about one Engilis and Pratt 1989). The last published pair per 4 ha). They were usually heard before Kaua‘i sightings were made on 16 and 17 they were seen, giving their characteristic, February 1989 (Pyle 1989). The population on loud, mellow, upslurred call-note, but were Hawai‘i has likewise dwindled, and there have difficult to locate because the call is ventri- been no recent reports of ‘O‘u on that island. loquial. SC saw males singing on several oc- The ‘O‘u is on the brink of extinction, if any casions, but was unable to record the song, a yet survive. complex canary-like performance, starting Kaua ‘i ‘Amakihi (Hemignathus kauaiensis), with two downwardly inflected notes, fol- RK.-This was the scarcest of the “common” lowed by a short, upslurred note, and a de- honeycreepers in our study area, and was ob- scending warbling trill and ending with a low served on only three occasions. HDP saw an downwardly inflected note of about the same individual near the camp on 6 July, and above frequency as the second note. HDP also noted it on 8 July. The first bird was clinging to a a single short song, comprising whistles and vertical ‘ohi‘a trunk and picking over the a trill, that he was unable to record but trans- bark, very much the way the ‘Akikiki forages. literated as feee-tooo-ter-wheet-wheet-wheet- SC and RJS observed a bird feeding on ‘ohe wheet-wheet-feee-tooo. When the males sang, naupaka (Scaevola glabra) flowers, which are they fluffed all the body plumage and flapped 4-6 cm long. To obtain nectar, the bird in- Conant ef al. l ALAKAI‘ BIRDS 11 serted its head into the bright yellow tubular CULMEN LENGTH corolla, rather than piercing a hole at the flow- 9ibngpthw fem.5 vripns er base as we have seen other Kauai‘ Ama-‘ 45+ kihi do at morphologically similar flowers. It 8 Au. doni (Maui) + also took insects from around flower bases. 15 HZL t&&mu' (Molokai)’ At Kokee‘ with more koa the Kauai‘ + Amakihi‘ was common in 1975 and remains 20 tiu. doni (Lanai)’ so today (HDP, pers. obs.). We attribute its -t rarity in the study area to the absence of koa. 19-H d&ti In 1975, the Kauai‘ Amakihi‘ was regarded as YL&&?~,+r5 one of four subspecies of Amakihi‘ (H. vi- rem), although Bock (1970) considered its bill r, I ! I I I I I I I i2 IA 16 16 20 22 size differences sufficient to separate it spe- cifically from other amakihis.‘ The bill is much larger than those of other amakihis,‘ with little or no overlap in measurements (Fig. BILL DEPTH 6). Interestingly, the recently split Oahu‘ Amakihi‘ (H. chbris), the Kauai‘ birds’ ap- parent closest relative (Tarr and Fleischer 1994), shows the least approach or overlap of all. Birders inexperienced on Kauai‘ may fail to appreciate the degree of difference and con- sequently misidentify a Kauai‘ Amakihi‘ as a Nukupuu‘ (discussed below). Therefore, re- ports of Nukupuu‘ on Kauai‘ should be re- garded with some skepticism unless accom- panied by unequivocal details. ! 9 I I 1 I 1 I I II Our 1975 observations and subsequent 3 4 5 6 7 6

studies (Pratt 1979) found potential isolating FIG. 6. Bill measurements of amakihis‘ from var- mechanisms of the Kauai‘ Amakihi‘ among ious islands, adapted from Pratt (1979). Means are in- ecological and behavioral characters. The bird dicated by long vertical lines, ranges by horizontal spends much more time picking prey from the bars. All measurements in millimeters. Number of bark of trees and less time feeding among specimens examined at ends of bars. leaves and flowers than other amakihis.‘ Rai- kow (1974) found such distinctive foraging behavior to be innate. The typical call of both present in amakihi‘ songs from other islands the Common Amakihi‘ and the Oahu‘ Ama-‘ (Pratt 1996). Individual trills of H. virens typ- kihi is a short, buzzy or mewing note rather ically last about two seconds, but those of H. similar to the call of the Blue-gray Gnatcatch- kauaiensis are about a half-second shorter. er (Polioptila caerulea) of eastern North Further, although the songs of all species are America. The Kauai‘ Amakihi‘ also utters variable, the Kauai‘ bird seems to have a wid- such a note, but only occasionally. Its typical er range of variation that includes trills on a call is a loud upslurred chirp (Pratt 1996), nearly level pitch. This observation may ex- sometimes indistinguishable from calls of plain why birds on Hawaii‘ that responded to Anianiau,‘ Akeke‘ e,‘ and Akikiki,‘ all of songs of conspecifics from Maui failed to re- which join mixed flocks on Kauai. All ama-‘ spond to playback of Kauai‘ songs, whereas kihis sing short trills (Pratt et al. 1987, Pratt Kauai‘ birds responded equally to either tape 1996), but the loudest notes of the Kauai‘ (Pratt 1979). Kauai‘ birds may have recog- birds’ trill usually drop in pitch whereas those nized the level-pitched trills as within their of the other forms remain level. The repeated range of variation, but Hawaii‘ birds did not elements of the Kauai‘ birds’ trill appear sim- recognize the descending trills from Kauai.‘ pler in sonagrams, and the song also usually Based on the large suite of potential isolat- has a distinctive introductory note that is not ing mechanisms and a level of morphological 12 THE WILSON BULLETIN * Vol. 110, No. I, March 1998 differentiation comparable to species-level well as firsthand investigations of plumage rank in other passerines, Pratt (1979) and Pratt coloration and ontogeny, sexual dimorphism, et al. (1987) recognized the Kauai‘ Amakihi‘ vocalizations, foraging behavior, and ecology. as a species, as did James and Olson (1991), Qualitative characters such as bill shape and on the basis of osteological studies and John- plumage pattern were the most useful in de- son et al. (1989) and Tarr and Fleischer (1994) limiting genera, whereas such characters as based on biochemical studies. The American tongue morphology and, surprisingly, vocali- Ornithologists ’ Union (1995) has now recog- zations proved more useful as indicators of nized the split. The Kauai‘ Amakihi‘ s’ tax- higher relationships. Numerous recent studies onomy is rather convoluted because its gener- have shown that the traditional prejudice ic placement has been controversial. The cor- against the use of plumage color in alpha tax- rect specific epithet depends on whether one onomy (as stated by Amadon 1950:166 and places the amakihis‘ in Hemignathus or in clearly evident in many other works of that some other genus. The specific epithet of the era) has no justification. In the two other most Kauai‘ Amakihi‘ is stejnegeri in any genus frequently cited avian examples of adaptive other than Hemignathus, where that name is radiation, the Vangidae (Langrand 1990) and preoccupied (Olson and James 1988). Pratt Geospizinae (Lack 1968, Grant 1986) plum- (1989a) proposed kauaiensis as a replacement age pattern is as good a predictor of relation- in Hemignathus. ships as any external character, and is much Historically, Hemignathus was used either more useful in delimiting genera than bill size for the akialoas,‘ with the bill greatly elon- or length, a character that Amadon (1950) ap- gated, and the nukupuus,‘ with a short man- parently considered pre-eminent and reiterated dible and elongated maxilla, together (Ama- in his critique of Pratts’ (1979) revision (Ama- don 1950), or for the akialoas‘ alone with the don 1986). Members of Pratts’ Hemignathus nukupu us‘ segregated as Heterorhynchus have sharply pointed downcurved bills that (e.g., Perkins 1903). Olson and James (1995) are dark colored with a pale base to the man- pointed out that both names are based on the dible. Adults are dull green above and more same type species; they restricted Hemigna- or less yellow below, with narrow dark lores. thus to the nukupuus‘ and named the genus Sexual dimorphism is pronounced, with males based on the Hawaiian name for those larger, longer billed, and always yellower than birds. Pratt (1979) was the first to place the females. Juveniles resemble adult females but similarly plumaged but short-billed amakihis‘ have pale wing-bars (retained in adult females in Hemignathus, although R. C. L. Perkins (in of H. chloris). The bill shape and most of the Wilson and Evans 1890-99) also considered suite of color characters differ from the an- the amakihis‘ closer to the akialoas‘ and nu- cestral condition in Drepanidinae (Raikow kupuus‘ than to the akepas‘ () with 1977), are not found among the which Amadon (1950) grouped them. Olson (for illustrations of all species see Clements and James (1995) denigrated those (e.g., 1993), and can only be regarded as synapo- American Ornithologists ’ Union 1983, Sibley morphies. Except for the Anianiau‘ (discussed and Monroe 1990, Tarr and Fleischer 1995) below), which we no longer consider to be- who followed Pratts’ (1979) classification, long to the genus, the various Hemignathus stating wrongly that it was adopted “without are so similar in plumage at any age and in any consideration having been given to its either sex that the bill would have to be seen merits” (Olson and James 1995:374). Because to confirm identification in the field (Pratt et the observations we began in 1975 in the al. 1987). Some other avian genera (Calidris, Alaka‘ i‘ Swamp were seminal in the devel- Nectarinia,Toxostoma, and Geospiza come to opment of Pratts’ (1979) taxonomy, we here- mind) exhibit a similar range of variation in with reconsider its merits, as compared to the bill length and shape. classification of the Olson/James team (see James and Olson (1991) based their alter- also Pratt 1994b). native classification primarily on cranial os- Pratts’ (1979) classification was based on teology, particularly the feeding apparatus, be- the literature dealing with tongue morphology, cause the postcranial anatomy of Hawaiian breeding biology, myology, and osteology as honeycreepers is so uniform as to provide lit- Conant et al. l ALAKA‘I BIRDS 13 tle phylogenetic information (Raikow 1977). biochemical studies do not always resolve Because they were classifying numerous re- systematic disputes. cently found subfossils, their focus on osteo- A classification compatible with both views logical characters was understandable, and would recognize the genus Viridonia for the they stated forthrightly (James and Olson amakihis‘ with Loxops, Akialoa, and Hemig- 1991:23) that they had not attempted a phy- nathus as separate genera. We consider this a logenetic analysis. In their most recent revi- counsel of despair, however, because it would sion, Olson and James (1995) recognized the turn back the taxonomic clock a century, genera Akialoa and Hemignathus (nukupuus‘ when nearly every species of Hawaiian hon- only) with the amakihis‘ being placed in Lox- eycreeper had its own genus. Such a classifi- ops along with the akepas‘ and the Hawaii‘ cation is devoid of phylogenetic information Creeper (“L. ” mana). In so doing they recon- and ignores demonstrated synapomorphies stituted Amadons’ (1950) Loxops, minus the that cluster the three curve-billed green dre- three species of and the Akikiki‘ panidine groups. The only way to taxonomi- (discussed below). tally recognize that cluster is to consider it a Amadons’ (1950) Loxops is probably the genus with Viridonia, Hemignathus, and Aki- most extreme example of alpha-level taxo- aloa (but not Loxops) as subgenera because nomic over-lumping in the annals of avian the entire drepanidine taxon, once considered systematics. It included one species of cross- a family, may be only a tribe (Sibley and billed akepa,‘ now shown to be at least two Monroe 1990; James and Olson 199 1). biological species (Pratt 1989b); four subspe- Anianiau‘ (Magumma parva), RK.-This cies of curve-billed amakihi‘ now regarded as bird was common, and we observed it fre- three species (Tarr and Fleischer 1994); the quently every day taking nectar and probably nearly straight-billed Greater Amakihi‘ (Viri- insects from ohi‘ a‘ flowers. Birds foraged in donia sensu stricto) which may be related to the tops of flowering trees along with Ake-‘ the newly discovered fossil genus Aidemedia kee‘ and Apapane,‘ but we saw no interspe- (James and Olson 1991); the Anianiau‘ (Ma- cific aggression. Occasionally the Anianiau‘ gumma, discussed below) which may not be foraged much lower, particularly among kan- closely related to the amakihis‘ after all (Tan: awao (Broussaisia arguta) plants. On 8 July and Fleischer 1995); and a “species” Amadon SC observed a single individual taking nectar called “the Creeper” now regarded by all au- by inserting its bill into the tubular corollas of thors as five species in two (Pratt 1992b) or ohe‘ naupaka (Scaevola glabra) flowers. three (Olson and James 1982, James and Ol- When ohi‘ a‘ is not in full bloom, the Ani-‘ son 1991) possibly unrelated genera! The only aniau is an active and agile insectivore, seek- conceivable character these birds have in ing its prey among smaller branches and common is a short bill (Pratt 1979). The re- leaves (Raikow 1974, HDP pers. obs.). In its moval of four of Amadons’ “subspecies” of movements and postures it resembles a wood- “creeper” from his Loxops does not make the (Parulinae). genus any more tenable. It has no defining HDP saw an adult feeding a fledgling bare- synapomorphies or even any general similar- ly able to fly on 8 July. July may be the last ities to unite its disparate members. Olson and month of the breeding season. Hart (1976) James (1995:375) stated misleadingly that un- saw a male giving courtship displays before a published genetic studies by R. Fleischer sup- female on 26 May 1976. Berger et al (1969) port their treatment “. .with the proviso that suggested that the nesting season of this spe- the akialoas‘ have not yet been analyzed and cies extends from mid-February to late June. that the amakihis may need to be further We heard no singing by this species in our split.” They fail to mention that Tarr and study area, but had learned the song, a trill of Fleischers’ (1995) recently published findings doublets or triplets, during our prior visit to lend equal support to Pratts’ treatment, de- Kokee.‘ The typical call note is a 2-syllable pending on which of two algorithms one fol- tew-weet, the first note low, the second rising. lows, and even then the genetic studies say Other calls resemble those of Kauai‘ Ama-‘ nothing about where generic limits should be kihi, Akeke‘ e,‘ and Akikiki.‘ The Anianiau‘ set among closely related groups. Obviously, remains as common today as it was in 1975, 14 THE WILSON BULLETIN - Vol. 110, No. I, March 1998 despite an apparently temporary reduction in Kokee‘ area by isolated observers (Pyle numbers following Hurricane Iniki (HDP, 1985b, 1988, 1992) have lacked sufficient de- pers. obs.). It is by far the most abundant of tails for unequivocal elimination of Kauai‘ the “green” honeycreepers in the Kokee‘ Amakihi‘ (discussed above). The Nukupuu‘ area. differs from the amakihi‘ in having the head When first described, the Anianiau‘ was paler (bright yellow in males, duller in fe- placed in the genus (which, at the males) without a contrasting darker crown and time, included the amakihis‘ and the “creep- a darker, thinner-based bill (Pratt et al. 1987) ers”) but was thought to be closer to the and it virtually never feeds in flowers (Perkins creeper group than to the amakihis‘ (Wilson 1903). and Evans 1890-99:xxi). As the species be- Because searches immediately following came better known it was placed with the Hurricane Iniki failed to find any Nukupuu‘ amakihis‘ in (Perkins 1903), (Pyle 1993c), many feared that it shared the and some authors even used the English name fate of the Kamao.‘ However, on 10 May “Lesser Amakihi” for it. In his eclectic clas- 1995, while conducting bird surveys along the sification, Pratt (1979) uncritically accepted Mohihi-Waialae Trail near Koaie‘ Stream, T the amakihi‘ relationship and placed the An-‘ L. C. Casey (pers. comm.) heard a short whis- ianiau in Hemignathus. Although it is a yel- tle, resembling one call of the low-green bird that resembles the amakihis‘ (Pseudonestor xanthophrys), that turned out to behaviorally, close examination reveals that be given by a Nukupuu.‘ Her notes include a the Anianiau‘ lacks the defining synapomor- detailed description of the bill as “. . .longer phies of Hemignathus. Sexual dimorphism is and slimmer than [that ofj a Kauai‘ Amakihi,‘ much less pronounced than in amakihis,‘ nu- particularly at the base where the [bill of the] kupuus,’ and akialoas,‘ and the dark lores are Kauai‘ Amakihi‘ is quite heavy.” She identi- totally lacking. The bill is short, only slightly fied the bird as a female on the basis of its downcurved, and flesh-colored with a dusky dull yellow head and noted that it “. . .spent culmen. A cladistic reinterpretation of Pratts’ several minutes foraging through branches of (1979) classification would place the Ani-‘ ohi‘ a‘ and olapa,‘ working each branch up- aniau in its own monotypic genus between wards from side to side.” Casey watched the L.oxops and Hemignathus. The phylogeny of bird for about three minutes as it foraged with those three genera cannot be resolved on the a mixed-species flock of small birds that in- basis of present knowledge. This taxonomy is eluded Anianiau‘ and Kauai‘ Amakihi.‘ This consistent with Tarr and Fleischers’ (1995) ge- observation is consistent with Perkins ’ (1903) netic studies (although not predicted by them), report that Nukupuu‘ often accompany such whichever algorithm one follows, as well as hocks. with those of Johnson and coworkers (1989). In the same general area, Casey saw a male Mathews (1925) proposed the generic name Nukupuu‘ on the morning of 22 June 1995. It Magumma as a replacement for the preoccu- gave a call similar to that of the previous bird pied Rothschildia so that a name would be but showed a much brighter yellow head. Lat- available if the Anianiau‘ was ever placed in er the same day at the same locality, J. Jeffrey its own genus. (pers. comm.) observed a female Nukupuu‘ Kaua ‘i Nukupu’u (Hemignathus lucidus for approximately 30 seconds. His descrip- hanapepee), RK, endangered.-A single bird, tions, particularly of the birds’ behavior are observed by SC, perched on the top of an definitive. He noted that when foraging, “the olapa‘ tree for about 30 seconds at 19:00 on bird used its bill to rip open the thin bark and 4 July. This subspecies is extremely rare, and, wood and probe with the upper bill [showing] at the time, had been seen on only two occa- that the maxilla was longer than the mandible sions (Sincock in Haley 1975) since its redis- and that the mandible was also curved down- covery in 1960 by Richardson and Bowles ward. To feed, the bird appeared to push the (1964). The USFWS forest bird survey in maxilla into or under the bark or soft wood 1981 failed to find any Nukupuu,‘ but 1985 then pull back on the maxilla while using the saw a flurry of credible sightings in several mandible as a fulcrum, tearing the bark or localities (Pyle 1985a, b). Reports in the wood.” Although these sightings seem to in- Conant et al. l ALAKA‘I BIRDS 15 dicate the presence of at least one pair at this ber two years later which was which! As not- locality, other observers (D. Kuhn, J. Lepson, ed earlier, we heard many O‘ u‘ whistles that pers. comms.) subsequently failed to find any were upslurred. Thus, we were intrigued by a Nukupuu‘ there. This species ’ long term sur- downslurred whistle, recorded by both RJS vival at very low numbers is quite remarkable. and HDP, we heard late in the afternoon of 2 Kaua i‘ Akialoa‘ (Hemignathus stejnegeri), July 1975 northwest of Sincocks’ Bog. The RK, endangered, extinct.-This was the only author of this call was too far away in dense historically known Kauai‘ forest bird that we forest for us to see it (the sun was setting at failed to see during our 1975 expedition. Fol- the time), and we never heard the downslurred lowing the 1960 rediscovery of this species call again. Did we, in fact, hear the “last” (Richardson and Bowles 1964), the only de- Kauai‘ Akialoa?‘ The question remained open tailed published report of it was that of Huber until 1989, when A. Engilis (pers. con-m.) (1966) of a sighting in 1964 in the same gen- heard a similar call and saw that it was uttered eral area as those of 1960. His description of by an 0‘ ~.‘ On that basis, and because Huber the birds’ awkward use of its extremely long may well have heard the call of an unseen bill as it fed among ohi‘ a‘ flowers is convinc- O‘ u‘ and attributed it to his akialoa,‘ Pratt ing. The Kauai‘ Akialoa‘ was both an insec- (1996) presented our downslurred call as tivore that probed among epiphytic mosses, “O‘ u,‘ Call 2.” Noteworthy is that Perkins lichens, and ferns and a nectarivore that fed (1903) described no similar note for the on deep lobelioid flowers as well as the more Kauai‘ Akialoa‘ but instead described its calls open flowers of ohi‘ a‘ (Perkins 1903, Rich- as distinct but intermediate between those of ardson and Bowles 1964). Although details its congeners, the Kauai‘ Amakihi‘ and the have not been previously published, l? L. Bru- Nukupuu.’ Nevertheless, we will probably al- ner (pers. comm., Pratt et al. 1987) saw a ways be haunted by our unseen whistler. Kauai‘ Akialoa‘ in 1969. An undergraduate Akikiki‘ (Oreomystis bairdi), RK, proposed and beginning ornithologist at the time, he endangered.-Although we observed Akiki-‘ was taken to a spot by local hunters and ki (formerly called Kauai‘ Creeper) every day, shown the bird, but could not reconstruct the they were not as common as Elepaio,‘ Ake-‘ exact locality. Apparently it was on land kee‘ or Apapane.‘ The Akikiki‘ forages by owned by Gay and Robinson, because his creeping along larger branches and takes prey guides were former employees of that corpo- from the bark surfaces, crevices, lichens, and ration, and thus it was not the same place as mosses. One bird spent three minutes taking the previous sightings. In conversations with insects from the bases of ohi‘ a‘ inflorescences HDP in 1971, Bruner described the birds’ (SC, pers. obs.). HDP noted several family head-down probing of the undersides of large groups consisting of immatures and a pair of branches and reported being surprised by the adults. The young were fully feathered and birds’ very large size, an attribute that would seemingly adept at foraging but still begged. not have been apparent to him from the pop- Immatures are distinguished by white “spec- ular literature of the time (e.g., Peterson 1961, tacles” (Fig. 7). We heard no songs from the which gives a deceptively small rendering). Akikiki,‘ only call notes, which resemble Only later, upon examining specimens, did those of the Hawaii‘ Creeper (Oreomystis HDP come to realize how much bigger the mana) but are a bit louder (Pratt 1992b). Since akialoa‘ was than other green honeycreepers 1975, the Akikiki‘ has declined catastrophi- on Kauai.‘ We are convinced that Bruner was cally. A 1990 expedition to Sincocks’ Bog the last person to see a Kauai‘ Akialoa.‘ failed to find any (Pyle 1990), although sub- Hubers’ (1966) report contains an enigmatic sequent searches have found them in the area description of the akialoa‘ s’ call. He saw the in low numbers, even after Hurricane Iniki bird in the same tree as an O‘ u‘ that was char- (Pyle 1993b). None were found by Walther acteristically sitting so motionless that he (1995) in his 1994 surveys west of the Alakai‘ failed to notice it at first. He stated that both Wilderness Preserve, although D. Kuhn (pers. birds were calling and that the notes were comm.) and tour groups led by HDP have identical except that one was upslurred, the consistently found them through 1995 at the other downslurred, and he could not remem- Kawaikoi Stream crossing on the Alakai‘ 16 THE WILSON BULLETIN l Vol. 110, No. I, March 1998

1992b) will be discussed elsewhere (Pratt in press). ‘Akeke ‘e (Loxops caeruleirostris), RK.- This species, formerly called Kaua‘i ‘Akepa, was common in the study area and provided daily observations, but none was singing at the time. They kept to the outer canopy where feeding movements were difficult to discern in detail. ‘Akeke‘e often appeared among in- florescences of ‘ohi‘a, but we could not con- firm whether they actually took nectar. Sub- FIG. 7. Juvenile ‘Akikiki (Oreomystis bairdi) sequent research has revealed that the ‘Ake- showing distinctive white “spectacles,” Alaka‘i ke‘e forages almost exclusively in ‘ohi‘a Swamp, 1975. Photos by FCJS. (Lepson and Pratt in press), unlike its sister species, the ‘Akepa (15.coccineus), which for- ages in a wide variety of trees and shrubs (Pratt 1989b, Conant 1981, Lepson in press). Swamp Trail, the most reliable locality outside The bills of the two L~xops, with their the preserve (Pratt 1993). Recently, ‘Akikiki slightly crossed tips, have been the subject of have again been found in encouraging num- much speculation as to the manner of their bers along the Mohihi-Waialae Trail (Pyle use. Based solely on their study of the birds’ 1996). Appropriately, the ‘Akikiki is being strongly asymmetrical jaw musculature and considered for listing as an Endangered Spe- skeleton, Richards and Bock (1973) presented cies (T Pratt, pers. comm.). a lengthy and highly detailed scenario of how As with other insectivorous Hawaiian hon- the bill might be used. Both methods they de- eycreepers, the classification of the ‘Akikiki scribed involve a 90” rotation of the head dur- has been somewhat controversial. Pratt (1979) ing feeding, a twisting motion that should be placed Oreomystis with the other thin-billed obvious to an observer, yet none of our ob- insectivores, excluding the enigmatic and pos- servations in 1975 or subsequent ones of both sibly nondrepanidine (Pratt 1992a, Tar-r and species involved any such movement, nor has F’leischer 1995) Puroreomyza, in the tribe it been reported by any other observers. When Hemignathini, but a cladistic analysis sup- foraging among leaf buds, both the ‘Akeke‘e ported its independent derivation from a - and the ‘Akepa look just like birds feeding at like ancestor (Pratt 1992b). Johnson and co- inflorescences, i.e., the head is held still and workers (1989) whose analysis of drepanidi- all action is accomplished by the bill and ne phylogeny substantially and irreconcilably tongue. Bet&man (1989) described the feed- disagreed with those of other biochemical sys- ing of the ‘Akeke’e at Pu‘u o Kila in the tematists on many points, recommended that Koke‘e area in some detail: no head-twisting Oreornystis and Paroreomyza be placed in one occurs; the bird parts the scales of ‘ohi‘a leaf tribe and all other drepanidines in another. buds by gaping and laterally abducting the Tarr and Pleischer (1995) found Oreomystis to lower mandible after the closed bill has been be allied with the finchlike Psittirostrini and inserted. HDP confirmed Benkman’s obser- not closely related to Paroreomyza. We be- vations at the same locality in October 1992 lieve transfer of Oreomystis from the Hem- and also observed the action of the tongue, ignathini to the Psittirostrini is justified, even which Benkman (1989) did not report. After though it would become the only insectivo- the leaf scales were pushed apart, the tongue rous genus in that assemblage, assuming was extended vertically down into the crevice, Pratt’s (1979) placement of the Maui - then withdrawn into the bill with a small lin- bill (Pseudonestor xanthophrys) with the ear object (presumably an insect larva) entan- Hemignathini (contra AOU 1983) is correct. gled at a 90” angle in the brush-like tip. On The question of whether Oreomystis is mono- 24 September 1995 HDP observed identical typic (Olson and James 1982, 1991) or in- feeding movements by several ‘Akeke‘e in the cludes the Hawai‘i Creeper as 0. mana (Pratt upper Kawaikoi Stream valley along the Conantet al. - ALAKAI‘ BIRDS 17

Alakai‘ Swamp Trail. To date, no aspect of tionary relationship between the bills of Ha- Richards and Bocks’ (1973) scenario has been waiian honeycreepers and the flowers they confirmed by field observation. feed upon (Givnish et al. 199.5, Lammers In February 1993, HDP made an enigmatic 1995, Patterson 1995). It clearly amounts to observation of Akeke‘ e‘ feeding behavior at nectar-robbing because the bird avoids contact the northern edge of the Alakai Wilderness with the flower pistil. This anomaly leads us Preserve: a male Akeke‘ e‘ twice bit the edge to speculate that this habit may be a recently of a mature ohi‘ a‘ leaf, pinking the edge with learned feeding behavior influenced by intro- two v-shaped indentations about 1 cm apart. duced flowers with flower structures that The leaf could not be examined closely but evolved with different pollinators. Similar be- through 10 X 40 binoculars appeared healthy havioral modification has been reported and without any insect damage. Whether the among Australian nectarivores (McCulloch bird ate the leaf sections could not be deter- 1977). Henshaw (1902) observed that I‘ iwi‘ mined. Among HDPs’ 1995 observations were on Hawaii‘ had learned to pierce the “spur” several “nibbles” at leaf edges, but no sec- of nasturtiums to reach the nectar, and Perkins tions were removed. Folivory in general is (1903) reported Hawaii‘ Amakihi‘ feeding on very rare in birds (Munson and Robinson long lobelioid flowers that “they have learnt 1992) and has previously been reported in to pierce at the base, at least in certain local- only one , the Laysan ities.” The flowers of the introduced South Finch ( cantans, Conant 1988). American passionflower PassiJlora mollissima (Himatione sanguinea), Apapane‘ RE.- known locally as banana poka, are too long This was the most common bird in our study and too straight for the I‘ iwi‘ s’ curved bill, so area. Numerous adults and immatures were the birds must pierce the corolla at its base to taking nectar from ohi‘ a‘ flowers in the tree reach the base, as we have frequently seen tops. Songs and call notes were given virtually them do. When in heavy bloom, these choking all day. This was one of the two species that vines attract large assemblages of birds with the Kauai‘ 0‘ 0‘ chased from a grove of heavi- the I‘ iwi‘ predominating. Other species that ly flowering ohi‘ a‘ trees (probably the 0‘ 0‘ s’ frequent banana poka include Kauai‘ Ama-‘ feeding territory). Apapane‘ were abundant at kihi, Anianiau,‘ Apapane,‘ and Japanese Kokee‘ in 1975 and remain common today but White-eye. Whether the smaller-billed species were noticeably reduced by Hurricane In&i. are capable of piercing the large and relatively One seen in a ravine in Kalaheo, a lowland tough banana poka corollas or simply take ad- locality directly south of Sincocks’ Bog, by vantage of holes made by I‘ iwi‘ is not known. members of a tour group led by HDP six Such feeding assemblages were conspicuous weeks after the storm provided modern con- at Kokee‘ in 1975, but the alien plant had not firmation of historical accounts of displace- yet penetrated the heart of the Alakai.‘ Nev- ment of montane birds by storms (Pratt ertheless, banana poka may have had an in- 1994a). Z‘ iwi‘ (Vestiaria coccinea), RE.We saw sidious impact even in areas remote from in- this bird daily, but not frequently. Solitary in- festations. Many large flowers native to Ha- dividuals took nectar from ohi‘ a,‘ Clermontia waiian forests, including representatives of spp., and Scaevola glabra. We did not see it such disparate families as gleaning for insects amongst stems and fo- (Givnish et al. 1995, Lammers 1995), Mal- liage, as it may do when nectar is scarce. vaceae (Funk and Wagner 1995), and Good- I‘ iwi,‘ as well as Apapane‘ were the target of eniaceae (Patterson 1995) in which flowers are feeding territory aggression by Kauai‘ 0‘ 0.‘ not usually curved, have curved corollas and When taking nectar from flowers of Clermon- other structural modifications for feeding, and tia and Scaevolu, I‘ iwi‘ invariably pierced a presumably pollination, by native birds. By hole in the base of the long tubular corollas “teaching” birds to rob nectar without polli- characteristic of these plants. This feeding nating flowers, banana poka and other alien method contrasts with earlier observations plants may have broken the evolutionary link such as that of Spieth (1966) and contravenes between Hawaiian plants and their bird pol- the seemingly obvious and often-cited evolu- linators. Thus, alien plants could be damaging 18 THE WILSON BULLETIN - Vol. 110, No. I, March 1998 native ecosystems in a particularly devious Although major preserves were established by way. the federal government and The Nature Con- The voice of the I‘ iwi‘ is highly distinctive servancy of Hawaii in forests on Maui, Mo- compared to those of other drepanidines. It lokai,‘ and Hawaii,‘ Kauai‘ saw only the be- includes metallic screeches, dissonant reedy lated designation of the Alakai‘ Wilderness notes, bell-like notes, clicks, and humanlike Preserve by the State of Hawaii. We cannot whistles delivered in a measured, seemingly fault authorities for such actions; we, too, con- random cadence quite unlike the canarylike sidered Kauai‘ “safe” when compared to oth- songs of drepanidine and the simple er islands that appeared to have more pressing warbles and trills of the Hemignathini. Other concerns. The sudden collapse of Kauai‘ s’ members of the , though they avifauna was a surprise. During the same pe- share some of the I‘ iwi‘ s’ tonal qualities, have riod, Guam experienced an equally sudden much more “conventional” songs. Remark- ecological catastrophe that was also unex- ably, the I‘ iwi‘ s’ song bears a strikingly close pected and rather mysterious (Pratt et al. 1979, resemblance to portions of the song of the Savidge 1987, Jaffe 1994). Neither island was aforementioned New Zealand Tui (see Kauai‘ perceived to be in imminent danger in the ear- 0‘ 0‘ account). In fact, one could produce a ly 1970s and both experiences show that is- reasonable facsimile of the Tui song by splic- land avifaunas can quite literally be here to- ing together pieces of the songs of the I‘ iwi‘ day, gone tomorrow. We now know that island and the Kauai‘ O‘ o!‘ None of these songs are avifaunas worldwide are mere remnants of much like the songs of meliphagids of central what was present before the influence of hu- Polynesia (Foulehaio carunculata, Gymno- mans (Olson 1989b; Steadman 1995), but myza spp.). Why the songs of two unrelated these geologically rapid were slow Hawaiian birds should resemble so closely compared to recent insular avian disasters. that of a bird that lives thousands of kilome- Any small-island endemic might well be con- ters away is fertile ground for speculation. sidered inherently endangered. Although in One possible explanation is that the song of the past HDP argued against listing locally the I‘ iwi‘ developed as a response to food ter- common island endemics as Endangered Spe- ritoriality by as yet unknown Hawaiian meli- cies (Pratt et al. 1979), we now believe that phagids. those who set conservation priorities should Nutmeg Mannikin (Lonchura punctulataa), use different criteria for island versus conti- A.-HDP saw three birds in Sincocks’ Bog on nental species. That insularity, in and of itself, 8 July, having heard one calling earlier in the greatly increases any species ’ vulnerability week. Although primarily found in lowland can no longer be disputed. open habitats, Nutmeg Mann&ins are com- We further believe that conservationists monly seen along trails in the Kokee‘ region should revise their view of nominal island and are the most likely of Kauai‘ s’ introduced subspecies. Some conservationists (e.g., Haz- estrildids to be found in forest openings. evoet 1996) have suggested that the new “phylogenetic” species concept (Cracraft CONCLUSION 1983), in which all distinctive island forms Our 1975 experience in the Alakai‘ wilder- would be considered species, would better ness, viewed in two-decade hindsight, teaches serve their purposes. However, such practical several important lessons for biologists and considerations are largely irrelevant to the on- environmentalists. Because it retained a rela- going philosophical debate in ornithology tively intact avifauna into the 1970s biolo- over species concepts (see Zink and Mc- gists of the time were comparatively compla- Kitrick 1995 for a review). In our view, con- cent with regard to both research and conser- servationists should work to save all distinc- vation on Kauai.‘ Whereas Maui and Hawaii‘ tive island populations whatever species con- were the subject of major research and con- cept ornithologists ultimately adopt. We pre- servation efforts in the 1970s Kauai‘ s’ forests dict that most such populations will ultimately held the long-term attention only of John Sin- be shown to be good biological as well as cock, and it was the last of the islands to be phylogenetic species. In many cases, the req- surveyed systematically (Scott et al. 1986). uisite data necessary to shift the burden of Conanr et al. l ALAKA‘I BIRDS 19 proof to the lumpers is available even in the ACKNOWLEDGMENTS original taxonomic revisions, which only need We owe an obvious debt to the late John Sincock to be reinterpreted in the light of modern in- for allowing us the use of his base camp, playing of a sight. In other cases, recent observations of tape of the Kaua‘i ‘0‘0 to aid our search, and being vocalizations and ecology can help in the reas- in general as helpful as one could be. We are further sessment of species limits as we have done indebted to other biologists who shared observations with us and allowed us to quote from their unpublished herein for several Kauai‘ forms. notes. These include Phillip L. Bruner, Tonnie L. C. The “species only” policy of Collar et al. Casey, Andrew Engilis, Jr., Jack Jeffrey, David Kuhn, (1994), although understandable given the Tom Snetsinger, and Eric VanderWerf. Robert L. Pyle, size of the problem, is unfortunate. The regional editor for National Audubon Society Field Kamao,‘ which was considered a subspecies Notes (formerly American Birds) kept us up to date on recent sightings. Other colleagues, including Robert of the Oma‘ o‘ (Myudestes obscurus) until C. Fleischer, Jaan K. Lepson, C. John Ralph, J. Mi- Pratt (1982) showed it to be a full biological chael Scott, J. V. Remsen, and Charles van Riper, III, species, is a case in point. By the time the new helped us formulate our ideas. Some of Pratt’s field classification was officially adopted (AOU equipment was provided by the Museum of Natural 1985), the population was so low that little Science, Louisiana State University. Assistance in ar- could be done to save it. In 1975, biologists chiving and analyzing tape recordings was provided by James L. Gulledge, Gregory L. Budney, and An- had far more concern for the Puaiohi, yet to- drea Priori of the Library of Natural Sounds, Cornell day the historically rarer thrush survives while Laboratory of Ornithology. Assistance with specimens the Kamao‘ apparently does not. A captive was provided by Alan Ziegler, Robert L. Pyle, and breeding program begun in 1975 might have Karla Kishinami of the Bernice l? Bishop Museum; gotten the species through the recent hurri- Robert Dickerman, the late John Farrand, Franqois canes, but none was even contemplated for a Vuilleumier, and Mary LeCroy at the American Mu- seum of Natural History; Richard C. Banks and Robert bird that was “only” a remote island subspe- Clapp at the National Museum of Natural History cies. (Smithsonian Insititution); and J. V. Remsen and Steve Other conservation lessons from our obser- Cardiff at the LSU Museum of Natural Science. vations include the findings that alien species on islands may have deleterious effects in pre- LITERATURE CITED viously unforeseen ways and that survival strategies that worked well in a pristine en- AMADON, D. 1950. The Hawaiian honeycreepers. Bull. Am. Mus. Nat. Hist. 95:155-262, plates 9- vironment can become maladaptive in dam- 15. aged ecosystems. Our studies also show that AMADON, D. 1986. The Hawaiian honeycreepers re- even seemingly superficial observations of is- visited. ‘Elepaio 46:83-84. land birds are important and should be pub- AMADON, D., AND L. L. SHORT. 1976. Treatment of lished; we can never know when a given study subspecies approaching species status. Syst. Zool. may be the last. Most oceanic islands are 25:161-167. AMERICAN ORNITHOLOGISTS’ UNION (AOU). 1983. much less frequently visited by ornithologists Check-list of North American birds, 6th. Edition. than Kauai.‘ Gaps in the ornithological liter- American Ornithologists’ Union, Washington, ature of 30 to 50 years are not uncommon for D.C. remote islands. Even a short interlude on a AMERICAN ORNITHOLOGISTS’ UNION. 1985. Thirty-fifth seldom-visited island can provide important supplement to the American Ornithologists’ data, and both scientists and recreational bird- Union check-list of North American birds. Auk 102:680-686. ers should neither overlook opportunities to AMERICAN ORNITHOLOGISTS’ UNION. 1995. Fortieth make such observations nor denigrate the re- supplement to the American Ornithologists’ sults as too superficial to be of value. Simply Union check-list of North American birds. Auk knowing whether a given species was present 112:819%830. on a certain date may prove critical. Because ASHMAN, P R., P PYLE, AND J. JEFFREY. 1984. A sec- islands are extremely useful natural laborato- ond nest of the Small Kauai Thrush. ‘Elepaio 45: 33-34. ries for evolutionary and ecological studies, ATKINSON, C., W. HANSEN, J. PRICE, L. SILEO, D. preservation of their avifaunas is particularly LAPOINTE, M. L. GOFF, C. LOCHER, AND L. TAM. important and must be addressed immediately. 1993. Role of disease in limiting the distribution Procrastination means extinction. and abundance of Hawaiian forest birds. Annu. 20 THE WILSON BULLETIN l Vol. 110, No. I, March 1998

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LAMMERS, T. G. 1995. Patterns of speciation and bio- PRAY, H. D. 1979. A systematic analysis of the en- geography in Clermontia (Campanulaceae, Lobe- demic avifauna of the Hawaiian Islands. Unpubl. liodeae). Pp. 338-362 in Hawaiian biogeography: Ph.D. diss., Louisiana State Univ. University Mi- evolution on a hot spot archipelago (W. L. Wagner crofilms 79-28440. and V. A. Funk, Eds.). Smithsonian Institution PRATT, H. D. 1980. Intra-island variation in the Ele-‘ Press, Washington. paio on the Island of Hawaii. Condor 82:449-458. LANGRAND, 0. 1990. Guide to the birds of Madagas- PRAY, H. D. 1982. Relationships and speciation of car. Yale Univ. Press, New Haven, Connecticut. the Hawaiian thrushes. Living Bird 19:73-90. LEPSON,J. K. In press. Akepa. in The Birds of North PRATr, H. D. 1989a. A new name for the Kauai Ama- America (A. Poole, F? Stettenheim, and E Gill, kihi (Drepanidinae: Hemignathus).Elepaio‘ 49: Eds.). 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