Distribution and Abundance Patterns of the Palila on Mauna Kea, Hawaii
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DISTRIBUTION AND ABUNDANCE PATTERNS OF THE PALILA ON MAUNA KEA, HAWAII CHARLESVAN RIPER, III, TMJ. MICHAEL SCOTT,2 AND DAVID M. WOODSIDE3 'Departmentof Zoology,University of Hawaii, Honolulu, Hawaii 96822 USA, 2PatuxentWildlife Research Center, U.S. Fish and WildlifeService, P.O. Box 44, Hawaii VolcanoesNational Park, Hawaii 96718 USA, and 3Hawaii Departmentof Land and Natural Resources, 1151 Punchbowl Street, Honolulu, Hawaii 96813 USA ABSTP•CT.--Censuses of the known geographicalrange of the rare and endangeredPalila were conductedin January (nonbreedingseason) and September(breeding season)1975. The habitat (mamane and naio forest of Mauna Kea, Hawaii) was divided into five major areas, with each analyzed for vegetationalcomposition, phenology of the predominanttree species,and Palila density.Using a line transectcensus technique, we determinedthat: 1) the Palila occupied5,560 ha, approximately 10% of its former range; 2) Palila populations were more restricted in the nonbreedingseason, possibly reflecting their flocking tendency;3) Palila densitieswere 38 birds per km2 in the breedingseason and 36 birdsper km2 in the nonbreedingseason; 4) Palila population movementswere small, and apparently were correlatedwith patternsof food availability; and 5) all methodsof analysisyielded a projected population of approximately 1,600 birds. These low numbers, coupled with its restricted range, make the Palila one of the most vulnerable endangered species.Received 10 May 1977, accepted16 November 1977. THE Palila (Psittirostra bailleui) is a finch-billed member of the endemic Hawaiian honeycreeperfamily Drepanididae, and is presentlyconsidered rare and endangered. We have extrapolated the pre-19th century distribution from analysis of specimens in museums and literature references(Fig. 1). Historically, its range included the mamane (Sophora chrysophylla) and mamane~naio (Myoporum sandwicense)eco- systemsof Mauna Kea, Hualalai, as well as the southwesternslope of Mauna Loa. Mamane fruit is the primary food source of the Palila, although there are no records of the bird occurring in the mamane forest on the eastern slope of Mauna Loa. It was apparently limited to the upper forest regions, as Perkins (1903) found it only from 1,220 to 1,830 m on Hualalai. Wilson and Evans (1890-1899) also reported it as being confined to the upland districts of Hualalai and Mauna Kea. Palila range has been greatly reduced since the turn of the century. Perkins noted indicationsof its declinewhen he reportedthat in Kona in August 1894he saw only two males, whereas in 1892 he had observed it "in numbers" (Munro 1960). This was apparently the last recordedsighting of a Palila in this area. The specieswas found to be locally common in 1943, 1948-49, and 1950 between 2,360 and 2,530 m on the western and northeastern slopes of Mauna Kea (Richards and Baldwin 1953). The last recorded sighting of a bird on the northern slope of Mauna Kea is that of Walker (1968), who observed three individuals above Puu Mali. Berger (1972) found the Palila on Mauna Kea common from 2,135 m to nearly tree line, rarely descendingas low as 1,980 m. Despite a much wider distribution in the past, all information available indicates that today the Palila is found only in parts of the mamane and naio ecosystemsof Mauna Kea. In order to document the distribution and abundance of the species, censusesof its entire known geographical range were conducted during January 1975 (nonbreedingseason) and again during September 1975 (breeding season). 4 Presentaddress: Cooperative National Park ResourcesStudies Unit, P.O. Box 54, Hawaii Volcanoes National Park, Hawaii 96718 USA. 518 The Auk 95: 518-527. July 1978 July 1978] Palila Distributionand Abundance 5 19 N Kailuc 0 5 10 15 20 25 I•ilo•eters 0 5 10 15 ISLAND of HAWAII Miles --- PresentPalila Range i'•...•i" '• Farmer Palila Range Fig. 1. Map of Ha•vaii indicating historicalrange and presentdistribution of the Palila. METHODS Seventeenpeople representing the Ha•vaii StateForestry and Fish and Gamedivisions, U.S. Fish and Wildlife and Forestservices, and Universityof Ha•vaii participatedin t•vo S-daycensuses. The non- breedingseason census •vas conductedfrom 13-17 January 1975, •vhile the breedingseason count •vas made from 15-19 September 1975. Five repeat censusesinvolving five observers•vere conductedfrom 22-26 Septemberto confirmthe reliability of the censustechnique. Total counts•vere made in the 80-ha enclosureat Puu Laau utilizing 16 observerson 17 and 23 September 1975. Information •vas gathered on Palila populationnumbers and density,geographical range and movements•vithin the range,as •vell as vegetationalphenology (flo•vering and fruiting densities)of mamane and naio in September. All the high mamane and naio forest on Mauna Kea was studied, •vith most of the transects being •vithin the fence-lineof the Mauna Kea Game Management District. Natural featuressuch as roads, gulches, and areas lacking trees •vere used to divide the mountain into five survey sections(Fig. 2). Section One •vas further divided into t•vo smaller units for the first census and into three on the second (Puu Nanaha to Kemole •vascovered only on the first count). Within this sectionthe Palila •vascensused from tree line at 2,815 m to 2,195 m on Parker Ranch. SectionsThree, Four, and Five •vere all covered on a single day in each survey, as each containedonly a small relict of mamane forest. Sections•vere divided into transectsalong either 60 or 30 m contours, except for the eight transects from Puu Laau to Puu O Kauha, •vhich •vere run from high to low elevationsduring the first census. Elevations •vere initially measured in feet and the numbers later converted to meters and rounded. Observersused altimeters, topographic maps, and compassesto maintain correctcontours. Each section 520 VANRIPER, SCOTT, AND WOODSIDE [Auk,Vol. 95 z000 % 15LAND of Hawaii Fig. 2. Map of Mauna Kea, Hawaii indicating sectionscensused. was walkedfrom oneend to the other,except Section Two duringthe secondcensus. During this count, observersstarted simultaneously from a mid-linepoint on eachtransect, and walked both directionsto respectiveedges of the forest.The changein techniquewas to reducepossible bias in the data, as during the first countobservations extended well into the afternoon.All othercensusing was done during the first4 h after sunrise,when birds were mostconspicuous. The initial day of eachcensus period was spent refamiliarizing participants with the Palila,and the remainderof the weekcensusing different forested areas. Each observerwalked at slowspeeds and recordedage, sex,and socialgrouping. To minimizedouble registrations, time of the observationalong with directionof flightwas recorded. Besides observations of Palila,all endangeredspecies were noted. Observationsmade by eachobserver were recordedas audio when a bird was heard but not seen, audio-visualwhen it washeard first then seen, or visualif observedfirst. No "squeaking"or othersounds were usedto lure birds. The right angle distanceof the sightingfrom the transectline was determined with a rangefinderor by pacing.Occurrences of the threeobservational types were plotted against the 276 distanceobservations for whichdetection type was noted.This indicatedthat Palila were detected by audio(or audio-visual)and visualstimuli in approximatelyequal numbers at distancesup to 25 m. However,as the distancefrom observerincreased beyond 25 m therewas an increasingreliance on audio stimuli;observations beyond 37 m werealmost entirely the resultof auraldetection. Screening effect of the mamanecanopy was probablyresponsible for the sharpdecline in visualdetections at greaterdis- tances. The effectivearea surveyed on eachside of the observerwas determined by poolingall sightingsand findingthat distancewhere the numberof sightingsfirst decreased dramatically (Emlen 1971). It was 15 m for the firstcount and 6 m for the second.This gavea coefficientof detectability(Emlen 1971)for the Palilaof 0.21 duringthe nonbreedingseason (January) and 0.11 duringthe breedingseason (September). Thesedetectability values are relativelylow, and reflectthe quietnessof the speciesand its habit of remainingwithin the canopy.Palila appeared to vocalizeand fly lessfrequently during the breeding season, and thus we found the difference in distribution of observational distances between the two censuseshighly significant (X 2 = 38.7; df = 10;P < 0.005). July 1978] Palila Distributionand Abundance 5 2 1 Density and total numbers of Palila were determined for each sectionusing the following formulas: D = P/A T• =D'O where D = density of Palila; Tj - total number of Pallia; P = number of Pallia tallied within effective detection area; A = hectares surveyed within effective detection area; and O = number of hectares bounded by Palila observations (effective habitat). The effective detection area was calculated for each section using the formula: A = M'L'C where L = total length of all transectsbounded by outermost Palila sightings;C = subjectively derived correction factor of either 1.1 or 1.2 for deviation from transect due to roughnessof terrain; and M = maximum probable detectability distance from the transect (January = 15 m, September = 6 m) mul- tiplied by 2. Population and densityestimates were developedfor each survey using: r=rj j=0 D= T/O where T = total number of Palila; D - density of Palila; O - number of hectares bounded by Palila observations; andj = section number. In order to measurediscontinuity of Palila distribution,a clear acetatesheet