Ecology, Behavior, and Conservation of the Maui Parrotbill’
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The Condor 89:24-39 0 The Cooper Ornithological Society 1987 ECOLOGY, BEHAVIOR, AND CONSERVATION OF THE MAUI PARROTBILL’ STEPHEN MOUNTAINSPRING~ U.S. Fish and Wildlife Service,Patuxent Wildlife ResearchCenter, Mama Loa Field Station, P.O. Box 44, Hawaii National Park, HI 96718 Abstract. The distribution,habitat response, sexual dimorphism, foraging, breeding, and flockingbehavior of Maui Parrotbills(Pseudonestor xanthophrys) were studiedover a five year period.The speciespresent’ range is confinedto montanerainforest on easternMaui, but dry lowlandhabitats on Maui and Molokai wereoccupied before Polynesian contact. Birdsoccurred from 1,250 to 2,150 m elevation,becoming most abundantat 1,750 to 2,000 m. In Kipahulu Valley, birds moved to lower elevationsin some seasons.Maui Parrotbillsassociated with areaswhere vegetation was less disturbed from feralpig activity. Activity areashad more developedunderstories and more opencanopies than nonactivity areas. Birdstended to foragein the subcanopyand understory,with 66% of the prey captured 1 to 5 m aboveground. Plant speciesuse deviated from expectationsbased on availability. The mostfrequent means of preycapture was excavation for timber-boringinsects in dead brancheson live plants.Foraging accounted for 39% of the diurnal time budget;an average prey item appearedto accountfor 1% of the daily energyintake. Male birds had longer, more deeplyhooked bills than females,and tendedto excavateto greaterdepths. Sexes differedin tree speciesuse and foragingmaneuvers, but not in substrate use or foraging height. The principal limiting factors appeared to be habitat loss, avian disease, habitat degra- dation, predation, and competition from exotic species. Control of pig populations is a needed managementaction. Key words: Activity budget;endangered species; jlocking behavior;foraging behavior; habitat selection;Hawaiian Islands;Maui Parrotbill. INTRODUCTION natural history accounts (Perkins 1895, 1903; The Hawaiian honeycreepers(Drepanidinae) of- Henshaw 1902) were followed by a half-century fer striking examples of adaptive radiation in bill ofneglect, and the specieswas feared extinct until morphology from a single colonizing ancestor. rediscovered in 1950 (Richards and Baldwin Specialized morphological and behavioral ad- 1953). Other sightings have since been made in aptations for feeding on seeds, nectar, insects, the remote montane rainforests of East Maui fruit, bird eggs,and molluscs span the range of (Bank0 1968; Casey and Jacobi 1974; Shallen- passerine variation. The Maui Parrotbill (Pseu- berger 1974, 198 1; Scott and Sincock 1977; Co- donestorxunthophrys), a fascinating but poorly nant 1981; Carothers et al. 1983; Scott et al. known honeycreeper,is a fairly small (length 140 1986), but little more has been learned of its mm) bird distinguished by a disproportionately natural history. A major survey conducted in large parrot-like bill used to excavateborers from 1980 on the distribution and abundance of forest timber. The sexes exhibit an extreme example birds on Maui, the Hawaii Forest Bird Survey of bill dimorphism among the honeycreepers (HFBS), estimated the population of the Maui (Amadon 1950). Parrotbill to be 500 birds (95% confidence in- The Maui Parrotbill has apparently retreated terval, 270 to 730), with a geographic range of from over 95% of its original range (Scott et al. 50 km2 (Scott et al. 1986). The specieshas federal 1986). Despite its striking appearance and ap- status as an endangered species (U.S.F.W.S. proachability, the specieswas not recognized by 1983) and studies of its natural history are es- 19th century Hawaiians (Munro 1944). Early sential in determining appropriate management actions and priorities to ensure its survival (Kep- ’ ’ Received6 August1985. Final acceptance29 Sep- ler et al. 1984). tember 1986. The objectives of this report are to quantita- 2 Previouslypublished as StephenR. Sabo. tively describe aspectsof the natural history of 1241 MAUI PARROTBILL BIOLOGY 25 the Maui Parrotbill, particularly its niche, hab- extending from 1,825 to 2,175 m elevation be- itat, and behavior, and to discuss its limiting tween the forks of Hanawi Stream in the Koolau factors,and management needs.This study com- Forest Reserve, and located in an area of high bines intensive field work during 1980 to 1985 Maui Parrotbill density. Topography in the area with analyses of museum specimens, literature is ruggedand steep,with an average slope of 40% reports, and unpublished HFBS data. on ridges.During May 198 1, the daily mean tem- perature at dawn was 7°C the mean afternoon STUDY AREAS high was 18°C and the overall 24 hr mean was 11.2”C. Temperatures of -4°C occurred in Jan- THE HFBS STUDY AREA uary 1985. Annual rainfall at 2,000 m elevation Native bird populations and general habitat in the area averages 500 cm, well distributed structure were surveyed in all areas of native throughout the year, except for irregular 6 to 10 upland forest on Maui during the HFBS. Four week periods of low rainfall (Blumenstock and major habitat types occurred within the range of Price 1967). the Maui Parrotbill. A mesic to wet forest type, At the upper limit of the study area, small dominated by koa (Acacia koa) and ohia (Metro- patches of alpine grassland dominated by Des- siderospolymorpha), occurred along the lateral champsia australis and Holcus lanatus lie margins of the main ohia rainforest of northeast imbedded in mesic subalpine scrub composedof Haleakala. This wet ohia forest had an upper Vaccinium reticulatum, V. berbertfolium, pu- strip adjacent to the treeline of mesic ohia forest kiawe (Styphelia tameiameiae), kukae-nene (Co- that was shorter in stature, somewhat drier, and prosma ernodeoidesvar. mauiensis), pilo (C. poorer in tree speciesthan areas below. Above montana), and the fern Sadleria cyatheoides.The the treeline a mesic subalpine scrub community subalpine scrubgradually gives way to mesic ohia marked the lower boundary of a thermal inver- forest at 2,070 m elevation, which grades into sion layer. wet ohia forest at 2,000 m elevation. The forest For this paper, the range of the species was canopy is dominated by ohia; the major subcan- divided into four contiguous units (Fig. 1). Ele- opy trees are olapa (Cheirodendron trigynum), vational boundaries of each unit were deter- kolea (Myrsine lessertiana), pilo (Coprosma mined by the highestand lowest Maui Parrotbill ochracea),alani (Pelea clusiaefoliaand Pelea un- recordsin the area. The Waikamoi unit extended describedspecies), kawau (Rex anomala), oheohe from the pasture-forestboundary north of Hos- (Tetraplasandra kavaiensis)at lower elevations, mer Grove to the east edge of Koolau Gap on and hoawa (Pittosporumconfertiforum) near the the north slopesof Haleakala, and contained me- treeline. The major shrubsin the forest are ohelo sic subalpine scrub and mesic ohia, wet ohia, and (Vaccinium calycinum), naenae (Dubautia plan- koa-ohia forest. The Wailua unit which extended tagineaand Dubautia undescribed species),akala eastfrom Koolau Gap to the west fork of Hanawi (Rubus hawaiiensis), kanawao (Broussaisia ar- Stream, and the Hana unit which extended from guta), and pukiawe. Henrickson (197 1) de- Hanawi Stream to the north headwall of Kipa- scribed the flora of the area in detail. Feral pigs hulu Valley contained subalpine scrub and mesic (Sus scrofa) have caused moderate to severe and wet ohia forest. The Kipahulu unit was de- damage to the dense understory. fined by the headwalls of Kipahulu Valley and contained mesic ohia, wet ohia, and at lower METHODS elevations koa-ohia forest. Additional areas out- LARGE-SCALE SURVEY side the Maui Parrotbill range were surveyed on windward East Maui at elevations below the The sampling design, field methodology, statis- known range, and over a wide variety of habitats tical analysis, and general results of the HFBS at Waihoi Valley, Kaumakani, Manawainui Pla- were described in Scott et al. (1986). Transects neze, Haleakala Crater, Kaupo Gap, Kahikinui, were systematically positioned at 1.6 to 3.2 km Auwahi, Kula, and the West Maui Mountains. intervals on Maui in native forest bird habitat. Stations were placed 135 m apart along the tran- THE HANAWI STUDY AREA sectsand sampled once in May to August 1980. Studieswere conductedintermittently from 1980 At each station observers conducted 8-min to 1985 on the 50-ha Hanawi study area (Fig. l), counts, recording distancesto all birds detected, 26 STEPHEN MOUNTAINSPRING Pacific Ocean @ Hanawi study area l fossil record m Maui Parrotbill range FIGURE 1. Location of Hanawi study area, four range units used in analysis, fossil records, and upper elevational limits of mosquitoes(from Scott et al. 1986). and botanists recorded habitat structure and pig tion, but for the elevation, habitat type, and pig damage to the vegetation. Incidental observa- damagedistributions, the recordsused were those tions of Maui Parrotbills and other endangered where the birds were within 50 m of the station speciesalong the transects, outside the system- or where the habitat of the bird was unequivocal. atic sampling periods, were also noted. Pig dam- Records of distant calling birds were excluded agewas classifiedinto five generalcategories based from the habitat analysis becausethey may have on a variety of native and exotic indicator plants occupied different habitats than at the station. that differed in sensitivity and response to pig This method of analysis differs from that in Scott damage. The categoriesconstituted a monotonic et al. (1986)