Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and demography of the Hawaiian hawk: Is delisting warranted? Journal of Wdlf Mgmt 67(1):165-176.
ABUNDANCE AND DEMOGRAPHY OFTHE HAWAIIAN HAWK: IS DELISTING WARRANTED?
JOHN L KLAVITER, Wldlfte Scrences Program, PO. Bax 352100. University of Wngtcm, Seae,WA 98195-2100, USA JOHN M MARZLUFF.' WUdllfe Sdences Progrem, P.O. Bca 362100. University of Washitun. Seattle, WA 881952100, USA MARK S. VEKASY.~Wildlife Samcws Pmgmm, PO. &ac 3521 06,University ol Washington. Seaffle. WA 86195-2106, UM
AbstmrC:'li>pix~vidr. rh~ 1J.S. Fish and Wildlifr &I vice (LrSWS) wi~hupclatrd informa~ionon the Hawaiian lra~~k ('io. Wtrlro solihrr7~r)tor rrcons~dc~auonol its currc-rlr r~rclac~gerctlslatruc, we tlewnni~rcdpopulation ~i7~.distrih ~rtion.h;tbi~a~ aviihhility, r~~niv:~l,fccl~l~dity, and finite rnrr of increase (A) ot 'io clu~ingIW8 arid 1999 can 111e island ol' I Iawaii. Wc cdnratrd tl~rtiltal ~rnpuht~o~~at 1.457 t 176.3 SE 'io. Birrlq wc-re broadly distributed arour~d the idand, but I~igt~wdr~lsiitcs were fuud in nlawce nauvt&wsr_ with agra..~u~~dc~story (0.57 r 0.12 'b/knr2). Wc- rstirnatc.d tJ;at 58.7% 01 tlrc- islil~td(6,144 kmZ) was usable habirrt for 'io. C)T the usable Jlahit;~~.3 1.8% (I .Y54 km2) currrrrrly is pro~rrtedty spate md fedcrul fi~rc-sts.parks, ar~clrefi~gm. Rased or1 an rvcrage density of 0.24 If. 0.08 'io/kln2 irr tho* XIXX?,uv ca1cnlatr.d that protected areas CCIITCIIII~ sttpport 469 'io (9S1K(:I: 244-Wl ). In all habitats combined. firstyear and ad1111strrviwl \va.- 0.50 + 0. I0 and 0.94 f 0.04, respcctiveIy, and fecurdity was 0.23 + 0.04 fco~alt:vuc~r~g/l)rceding female. Overall. )c was 1.03 k 0.04. Eladcity analyses qhowed dull survival to be the most important parnmctt-r ~qdatinp:Iiiiiw grc~wtltof this populatiori. The poplrbtion appears viable b;lsc.d on A u apprcrxirnatelv I and 11igt1adult simival. Additiortally. sukt~ntiaJareas of habitat ate protrxtcd. 'io appear rcsb tan1 do aviar~diw~wx found on thc island. no evidence mists of' rie~ativeimparts from contzuninanu. aid the hid% show rhr ability tn IIW humanaltered Isnckapes slid exotic prey. Becalm of the shon study dudon. relativeiy low population riz, variance around o~ires~in~ates (abundance, fecundity. survival, L). and envimnmental st* rhrsGcity, we believe Orat clelisti~rgto a no~ilhrentenrdstatus is ur~warnrited.Hwwr. wrr belicw that downlisting to thrr.ate~~edstatus wolrld be appropriale for this species, or ernhracing a 1ir~i11gas 'nc-ar threawncd" ur~derthr 1ntc.mational I'nion fur the G)n.wrv;l~ionol'Natum (TU<,W) Red 1,ist categories. Rqardlc-m of a change in polili- cal stat~~s.ppnlation size qntl especiall?~:irhtl~s~~~vivnnlri~ shor~ldhe ro~~tinclymonilored. JOURNAL OF WLDUFE MANAGEMENT 67(1):165-176
Key d:abuntlatirr. B~ctroscdituriur. delistin~.demopnphy, downlisting, elitiriy aiialvws. I'rcunclitv. finite ratr of incnxse. I-iawnii. Hawaiian I~awk,'io. rurvivd.
Hawaii conlaitls 349 endangered and 14 threat- size (Ominstein 1968. Berger I98 1 ): perceived ened spucics, of which 282 are plants (C:SFUrS threats to its native forest liabitat horn ae;ricul- 2000). 'This is nearly 33% oI't11r 1,107 specic..~list- (ure. logging, ca~nmercialdevelcrpmcnt (USFWS ecl ill ll~c1rSA (1lSWS 19996). Although tilany 1984): and lack of' infbnnation on the species tirrei~tenccl and cnc1angr.rt.d Han-aiialr species (Griffin 1985). Additionally, lapti>rs had been iIttrac.1 atterltiorl ol' frdetal, stare. and pm-tc. declining worldwide froln cantanli~lantssuch xi orgx~~i-~atiarrs,the future seerns ~incertilin for DDT (Newton 1979:229-262). anci the eEwrs of tnmt specics. Tct date, Iio species haw ken thcse on 'io populations wcl-c. unknown. Ho\vev- delisted or downlistccl in Hilwiii, although 4 cr, [he alirnatcd 'io pop~tlation size in 1983 Pacific island bircl specres ('Tiniarl n~o~larch (1,400-2,700: GnEn 1985, 1989) mssimilar to [ Monrirrha IIJNI/.IU~US(~~),Palall grot~nrl-dove[ (mi- thc 'io recover? plan's target af 3,000 for ciawrl- licolund)(~ ranr/n,tu]. I'alau owl ( I~lu~gInrrxlisting to "threatened- status (CJSFWS 1984). Tt~r piwhzrff't8nl. and Palau fi~~liulI IG~rf~tdurn /r/~ih] ) 1983 r-slitrlate was basetl ~IIIdislrihr~rion data cd- have becn rcmc~vedIro111 tile I 166 HAWAIIAN I-f:\M'K /\RUNDANCE.\ND UEMOCKAYHY h'hailtrrrr /I/. J. U1ildl. Manage. 67(1):2003 al. 1997). ilall et al. ( 1997) found 'io distributed eral pc~pulailonestimate. We believe that theJune throughout thr island and calculated a total pop 1998 survey best represenis the popularion of 'io ulation estimate of' 1.600 birds. They agreed with because it ( 1 ) occurred at a time of year when 'io the recornmendation to downlist, but suggested responded well to playbacks; (2) collsemtively that researchers collect long-tern1 dernopphic estimated abundance since the survey occurred data to better understand the status of the species. before younpf-theyearwere added to the popu- In 1994, rhe USFWS withdrew the proposal to lation; and (3) had the greatest number of survey downlist due to lack of public support and limit- points, providing less variability in our estimates. ed ability of the 1993 survey to ascertain the pop We followed exact survey routes and methods ulation stanu with confidence. Furthermore, sew used by Hall et al. (1997) for rhe pre-bmeding eral important facton renlainecl r~nresolvedsuch season survey, wilh 399 survey points located a total paputation size,fjreei311g pair prorfuctb minimum-&.% krrntw-d pest- ity, and fledgling surviv;ll (USFWS 1994). breeding surveys, we expanded the survey mutes. We initiated this stucly to provide the USFWS incrrd the number survey points to 677.-and with demographic info~n~ationto assist in their inaeased the minimum distance between pain& w assessment of status of the 'io. 01.11- ol?jectivcs 1.6 krn for greater coverage on the island and in- were to ( I ) estimate population ahtindance by creased independence between sampling points. pointxorrnt surveys and determine whether pop Points W~L'L.mpled only once during each survey- ulation estimates were r.epeatahlc over time: (2) We surveyed each point for 10 min using play determine the amount of 'io habitat on the back recordii of adult and fledgling 'io for 2 1- island; (3) determine survival of adult and first- min periods during the first and eighth minutes. year birds in native and exoticdominated hibi- Prior to each day crf surveying, all ohservers prac- tats; (4) determillc 'io fecundity in native, mixed ticed estimating distance with a kshnell 400 nativ-xotic, and exotic habitats; and (5) deter- laser rangefinder (Bushnell Corporation. Over- mine finite rate of increase (1)in native, mixed land Park. Kansas, USA), tape measures. and nativc-exotic, and exoticdominated habitats. a~~tomobileodometers. We conducted surveys between 0900-1 700, with winds S24 km/hr, and STUDY AREA with no precipitation or light rain. At each point. Our research took place on state, federal, and we recorded {I) 'io detections, (2) the type of private lands distributed widely throughout the detection (audio or visual) and the distance at island oi Hawaii. Point counts and nest monitor- which it w~sfirst made, (3) percentage of survey- ing occurred between 10-2.150 m in elevation. or's view obstructed, and (4) the habitat sur- Detailed descriptions of the study area exist in rounding each detection according to a modified Guddihy and Stone ( 1990) and Jacobi ( 1990). habitat scheme {Table 1). This scheme modified Hall et al. (1997) habitat classerr by combining METHODS vegetation types 5 and 10 and adding 2 new class- es: urban and lava. Population Abundance and Distribution We corrected for 'io movement coward the We conducted 4 island-wide pointcount sur- ob.wrvcr prior to detection ('io were attrac3ed to veys using methods developed by Hall et al. 'io playback recordings) with methods developed (1997) and modified by Klavitter (2000) to esti- by Klavi ttcr (200). After correcting for the move- mate population density and abundance. Points ment bias, we analyzed poi~ltcoi~ntdata with pre were chosen systematically from a random start- #ran DISTANCE (hake et al. 1993. Thomas et ing point and accessible by road. Two pre-breed- al. 1998). We used a global detection function and ing surveys were done in Jaouary 1998 and 1999, allowed Qhe program to select the best model using a breeding survey in June 1998, and a post-breed- minimuin Akaike Info~matianCriterion (AIC) ing survey in September 1998. January or June ~alues(AkAke 1973, Bt~cklandet al. 1993:75-76, survep meamred the population after overwinter Hall et al. 1997). We right-truncated the largest mortality and prior to younpf-theyear entering 3% of the distances to facilitate model fitting the population (Griff~net al. 1998). therefore (Buckland et al. 1993). conservatiwly estimating total population size. We estimated 'io densities for each habitat and The September survey measored the pop~~lation calculated an island-wide popt~larion estimate after young-of-the-year en tered tlie population based on the amount of area for each major habi- and prior to significnr~tmortality. producing a lib- tat on thc island (Table I; Klavitter 2000). Mk esti- Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. J. Wildl. Manage. 67(1):2003 HAWAIIAN HAWK ABUNDANCE AND DEMOGRAPHY &Yil@el al, 167 Table 1. Scheme used to classlty he habitat sunounding each of 677suwy points on the idand of Hawad, USA. Habitat was assessed durlng 'io surveys oondwed from Jan 1998 to Jan 1999 (Tat& 2). Habltal within the area swveyed was estknated to fac~htatehabitat-speclftc projectrans of 'io densities. dassitbflon scheme was rodified fmn Hall el al. (1997). Hatntal types Areaa Codeb oeMptionc Fallow sugarcane 174 1 Sugar cane fields (fallow or Wve) with ewobc or nabve trees or shrubs at edges Exocc trees, shrubs, grasses 733 2 Short or tall exolic bees wnh emk shrubs and grasses Orchards 76 3 Macadamla nut, papaya, or ohrorchard with nab Worexdc trees or shrubs at edges Grass dominated Areas dominated by grass with few or none nathre andlor exotic trees: scattered homes Shrub domimed Nalive andlor exotic shrubs domlnab the landscape: lava occasionally visibJe Mature nahforest Nathm trees and shrubs dmmate Ihe erea Native-exotic brest Mxed exotic and native treas, melimes with mixed exotic and nathre shrubs or .grass Exurben Residenhal areas with scathed exotic and naWe vegetation Pioneer native forest Native trees and mixed exotic and native shrub vegetation on lava; a pioneer community Mature natilorest with Mature native foml wiM a gmsedominatd understory grass understory Memane-naK1 forest Mamane-na~owgebtkn with a grass andtor an exotic &rub understory Urban Dense residential and lndustrlal areas and whh Eew or no trees Lava Areas dominsted by lava wtth no trees and few or no shrubs or grasses a Total area of each habital on the island (ha). Hlgh-elevaUon pioneer native forest (237 km2) and hlgh elevation shrubs-grass- es (20.709 km2) are not shown because haMtats are OWthe normal 'io range. Vegetakn codes used by Hall el al. (1 997) Corresponding codes are listed adjacent to our habitat types. Codes 5 and 10 were combined. and 13 and 14 were added. Scientific names of plants listed: sugar cane (sacchamoilicinanrm), macadamia nut (MawWnia femiWa).papaya (Car- ca pepaya). Mamane (Sophora Chpsq~hylIa)),and nab (Myqporum sandwmsis)). mated the amount of each major habitat using foreseeable h~n~reand wouid continue to support Jacohi's (1990) vegetation map and analysis of hawks. We then determined the overlap of pro- SPOT mtellitc- imagery (SPOT 1994). Most of the tected lands and the mas supporting a minimum habitat rnappcd I>y Jacobi ( 1990) consistc.d of density of 0.07 'io/km2 to calculate the amount native vegetation communi tics between 500 and 'io habitat currently protected on the island. We 2,500 m elevation. All tnapped areas were field- did not include habitats with densities 4.07 checked along transecn located Ihrongl1out the 'io/km2 in our measurement because (1) we want- island of Hawaii (Scor t et al. 1981 ) with 7,863 total ed a conservative measure and (2) the,% habitats sampling points. We converted the level-l clascs stlppon little or no productivity (Rlavitter 2000). from Jacohi's ( 1990) map to the 13 habitat types in Table I. We used satellite imagery (SPOT 1994) Estimating Survival to visually map the remaining 5.5 16 km2 of habi- From January to September 1998, we used bal- vat on the island to complete our habitat map cllatri tl-aps (Berger and Mueller 1959) with live (Klavitter 2000). We used 1,076 'io survey poinh black rata (Rnltw mnus) to trap adult and recent- and 75 'io nest sites to ground- ruth classifications. ly fledged (68-89 days of age) male and female We used habitatqxcific density estinlates from 'io in native- and exotic-~io~tedhabitats to the June 1998 survey and the habitat map determine survival for a I-year period. Son~e described previously to nlap 'io ab\m&ance and nestlings (3845 days of age) were captured by distribution otl the island. We used the June 1998 hand in the nest. We weighed, measured, and sunfey because we iwlicved that ir provided the handed 'io with a unique combination of plastic IIIOSLaccurate, vrt conservative estimates. color bands and a USFWS aluminuln band. We We used program ARCMEW (Envtronmental scxed 'io (all ages) by body mass, wing chord. Systems Research Insti~ute1998) to calculate the and tarsus mensuremerm and aged birds (1.2, 3, amoun[ of 'io hal>itat protcctcd by state iorcst 4+ year) by plumage. molt, and color of iris, cere, and natutal area reserves and fcderal parks and tarsus, and feet (Griffin 1985, Clarkson and hni- refitges ((oral protected land on island = 3.241 awe 2000). We attached a backpack radiotra~~s- km2). We axsumed thal protected lands tha~cur- tnitter (Advanced Telemetry Systems, Isan ti, Min- rently srlppol-I 'io wolild not changc. greatly in [be nesota, LISA) to each bird using a break-.way Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. 168 HAWAlL4iV HAM7K ABUNDANCE AND DEMOGRAPHY Kluvifkr rt 01. J. Wildl. Manage. 67(1):2003 harness dcsign modified from Buehler et al. cally lay 1 egg, we used simple binomial propor- (1995) and Vekasy et &l,(19%). We added a tions (Zar 1996: 5 13-568) to calculate success. medium-weight cotton thread *weak link" LO the Radiotagged and known bl-ceding-aged females harness on the ventral surface of the bird tha~ thal did not attempt ro breed were recorded to connected dl 4 straps. We placed a small leather de~ernnnethe portion of the population not patch under the weak link that prevented 'io breeding in a given year. We determined fecun- from rising their hill to sever the cotton thread. dity (j)as the number of female offspring prtr Radiotransmitters (9 g for n~ales. 13 g fbr duced per breeding-age I'emale per year. Vaxi- Females) weighed 5.3% of the bird's Mymars. ance was calcl~latedar Transmitters werc rated to last 210 and 24 rnorrths LOr males and females, respectively, and included a mortality switch. We located 'io a min- imum of once/month from the ground. except for October-December 1998. We searched for missing birds fro111tlelicopters. where 71 = wcil number of females used to calcu- Wc dctel~ninedadult and tint-year siirvi\al for late fcc~lndity. a I-year period (Apr 1998-Apr 1999) usirig simple binomial proportions (Zar 1996:5 13-568) and Estimating Finite Rate of Increase asrrumcd 100% detection rales for rddiomarkcd We used program RAMA!3 GIs (Akcakayd 1999) birds. Variance was calcula~edas and Leslie matrices (1,eslie 1945, 1948) to esti- Inate the finite rite of increase (A),the long-term stable age distribution (SAD), arid the average ageclass I-esidence. We set up our stage matrix usirig 4 age classes: I. 2. 3. and 4c. Age of first- reproduction was age claw 4 (3 years of age; f Clarlrson and Laniawe 2000). We used our esti- where S = number of lirds surviving after 1 year, mate of firstyear survival for age class I, atxd our if = number of mortali$eq _after I year, and X = adult survival for 2.3, and 4+ age classes. We did total number of 1md.s (S + M). Wc chose April as riot niodel demographic or enviro~lmentalst* the beginning of'the survivdl period based on our chastici ty or density dependence. We used elas- 1998 egg-laying date (mean + SD) of 23 April i ticity .analyses (Cawell 1989,2000; de Kroon et al. 26.2 days. 2000; Heppell et a]. 2000) to determine the rela- tive contributio~lof each demographic parame- Productivity ter lo A. Elasticity values ringe From zero to I. b'e ~nor~stolednest5 to deterlriirle success from with 1 being the most infliielltial on population M~J-chd~rough Septcmhcr during 1998 ant1 regulation. Paranietels with highest elasticity =I- 1999. We checked historical 11estitlg territo~ies tics were used to idenlib potential management and followed radiomarked birds LO find rlests targeu (de Kroon el al. 2000, Saethcr and Bakke containing epgs or young. We checked empty 2000. Wisdom et al. 2000). We used PI-ow ncsts containiug new rnaterial (stick, etc.) a mill- MATHEMATICA (Wolfram Research IW) and imum of once every 2 weeks to see if the pair Iaicl the Delta method (Seber 1973) to lind the vari- eggs. We determined egg laying by observing ance of' li given the variances of the survival and eggs or an incubaung 'io on the nest. To avoid fecundity rates. We used I-tailed &tests and u = disturb;incc. and posqible abandonment, nest 0.05 to test H,: A s I. Uees were not cl~nlbeclduring incubatiori or the litst 5 weeks of' chick realirig. Mre avoided pro- Other Potential Limiting Factors longed aest observations dilrillg this time. We collected 'io eggs from ilests 2-6 weeks after We deter~ilir~edllcstitlg success for pairs found nest failitre ro delemine levels of envir-onmental during the incubation phase (Steenhof and co~iri~minaiitssuch as organochlorines and heavy Kocllert 1982, Steenliol' 1987). Pairs fbuncl after metals. We weighed and meas~~redeggs, extlact- inc~~badonwere not usctl in the analyses. Suc- cd contents. and froze contents in sterile glass C~SS~LIIpain raised at least I othpring to llerlging containe~~.The U.S. (kological Survey (USCS) qe(59-63 days). Failed pain were il~isuccevvful Patuxent Analytical Control Facility in Laurel, in fledgling at least 1 offspring. Because 'io iypi- Maryland, analyzed sarilples (detection limiu: Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. 1. Wiltll Manage. 67( IM?O3 HAWMIAN HAWK ABC'NDANCE AND DF,MmRAPHY 9 K1mrirtr~d 169 native Hawaiia11 forest bids (van Riper III et al. 1986. Work et al. 2000). We sent samples to the USGS Biological Research Division in Volcano, Fiawaii, to check for avian malaria, and to the U.S. Department of Agriculture in Beitsville, Mqland, to check for 7: gondii. We visually checked all captured 'io for avian pox (Poxiviru~aviuin) lesions and also checked birds and nests fbr ectoparasites. Nests were checked after nest failure or during the late chick phase. Population Abundance We courltrd 78 'io in-january 199R. 178 in June Fig. 1. Estimates of the total popuhtton of 'oon Hawaii, USA. 1998. 144 in September 1998, and 72 in January (mean i 95% CI) obta~nedduring 4 pointcount surveys aa 1999 surveys. Altllough population estimates part of this study (Jan 1998, Jun 1998, Sep 1998, Jan 1996) were similar among all surveys (Fig. I), derlsities and an estimate by Hall el al (1997,Dec 1993). Eslimetes are slansticalty similar bemuse all 85% Cis overlap. Jan 1998. varied among habitas and among surveys (Table Jun 1888, and Dec 1993 all ldlowed the same sumy routes 2). Dcnsitit.~were lowest in January 1999. Areas and consisted of 399 pomts. Jun 1998 and Sep 1998 bolh Id- lowed the same survey route consisting of 677 points. ~300m elevation on the west and southwesr por- tions of the island supported few or no 'io. 'Io were absent fmm areas >2,100 m elevation. Based on the analyes of the June 1998 survey, densities 0.01 -0.76 ppm) for PCBs, 20 organochlorines, differed significantly among habitats and ranged and 18 heavy metals (Klaviuer 2000). rmnl0.01 to 0.57 'io/km2 (Table2; Fig. 2), For all We obtained 'io plasma samples (n = 7) from surveys combined, native forat wilh pas. fallow the USFWS to test for avian malaria (Phnou!ium sugarcane fields, and orchards had the highest wlict?1.mtapidrnnnne) and toxoplasmosis ( Tox@krr- average density rank (Table 2). Mamanenaio, m ~wdii),known to cause mortality iu other urban, and lava had the lowest densities. Table 2. 'lo hab~tar-specificdensities ('idkm2) tor Jan 1998-Jan 1999 surveys and the average rank of each habitat (based on density) on the island of Hawail. USA. Densnies were derived trom analysis of point counts at 399 (prebreeding Jan surveys) and 677 (breeding and postbreeding Jun and Sep surveys) pdnts. 'lo calls were broadcast during counts to facilitate detection; bhs was measured by observing radlotagged 'io responses to playbacks and accounted for in density estimates. We test& for dlf- ferences between habitat density estimates for the Jun 1998 survey because this was our most accurate survey. Jan l9Q& Jan 1998 Jun 1898 SW 1998 Jan 1999 Jan 1QSEl No Avo. Habitat Denshy SE Density SE diffe.renc# Density SE Denshy SE rank Native torest with grass 0.62 0.40 0.57 0.12 1 0.45 0.12 0.10 0.10 2.25 Fallow sugarcane 024 0.11 0.40 0.11 1.2 0.38 0.10 0.10 0.06 3.75 Orchards 0.35 0.12 0.28 0.09 2. 3 0.38 0.12 0.22 0.10 2.25 Mature native forest 0.33 0.11 0.27 0.07 2. 3 0.12 0.08 0.22 0.09 4.50 Exok trees, shrubs, grasses 0.24 0.1 1 0.24 0.10 2. 3.4 0.19 0.08 0.29 0.10 4.00 Nativeexotic forest 0.15 0.11 0.15 0.05 3. 4. 5 0.25 0.08 0.15 0.06 5.75 Grass dominated 0.29 0.09 0.13 0.04 3, 4.5 0.17 0.06 0.22 0.10 4.75 Pioneer natlve bred 0.08 0.08 0.09 0.04 4. 5 0.14 0.07 0.04 0.03 8.75 Shrub dominated 0.19 0.09 0.06 0.04 5 0.03 0.03 0.07 0.07 8.50 Exurban 0.04 0.03 0.06 0.03 5 0.17 0.06 0.05 0.05 8.50 Mamane-Naio forest 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 11.00 Urban 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 11.00 Lava 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 11.00 a Habltats dth the same numben Indite no diffsrence in dmiiy estimates (Z-test,n = 0.10). Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. ,1,3O ~~~~DEMOGRAP'HYr Klardt~ttd J. Wildl. h4anage. 67(1):2003 Nest-tree sites were distributed widely through- our the island (Fig. 2). Most (81%) nests occor-red in native 'ohi'a (1MehAdem~po~morpha) trees, hui some occurred in exotic trees. We found 50 nests with eggs, young, or fresh nest lin- ing in 1098 and 63 in 1999. Of these, we found 39 (1998) and 49 (1 999) nests with incubating par- ents. In all habitats combined, 59% (1998) ancl 49% ( 1999) of the pairs found during incubation Iledged young (Fig. 3). In 1998, success in native habitat (71%) tended to be slightly higher than succty in exotic- habitat [44%: &= 2.94, P = 0.086), In 1999, less difference in nesting success occurred between native (56%) and exotic habi- tat (35%: X{ = 1.97, P= 0.160). In all s~iccessfulnesh. only 1 young fledged. Of the 113 nests monitored during the study. females laid 2 eggs on only 2 occasions. Fledgling sex ratio was 51% male, 49% female (n = 43). Cambining all habitats, overall fecundity was 0.23 f 0.04 female young/adult female (Table 3). We found no difference in fecundity between native (0.28) and mixed (0.17; % = 1.27. P= 0.20). native and exotic (0.10; Z= 1 -35, P= 0.18), or mixed and Fig. 2. 'lo dmtribution and 4enslly 0dtetmined from exotic habitats (Z= 0.69, I-'= 0.49; Table 3). the Jun 1998 survey, and the tocations of 75 'io pairs found during the 1998-1888 held seasons (only Lhe first nest found Estimating Finite Rate of for each pa11 is shown) on the island of Hawaii, USA. From the Increase 75 total .pa~rs.50 nests with eggs. young. or hesh nesting We used survhal and fecundity results to set up material were located In 1998 and 62 in 1999. State and fed- eral lands (resemalrefuges) mat overlap suitaWe 'io habitat the following habitar-specific Leslie matrices: (areas with 'io dendies d 0.07 'ldkm2 or greater) are oul- lined. 'lo occurred widcly throughout the island on protected and unprotected areas (Fig. 2). We esti- mated that 58.7% of the island (614,405 ha) was 0 0 usable 'io habitat (supporting densities 20.07 (2) Native 'io/km2). The total anount of usable habitat that 0.27 0 0 0 is protected amounted to 1,954 km2 (32% of all usable habitat, Fig. 2). with an average density of 0.24 f 0.08 'io/km2. We esli~natedthat protected lands suppc~rted469 'io (95% GI: 244-901 ). Very (3) Mixed 0 0 0.17 few high-density areas (0.40-0.57 'io/km2) were protected, while some of Lhe moderatedensity (0.154.39 'io/km2) were protected. 0 0.94 0.94 Survival and Productivity 0 0 0 0.10 Sittvival rates (br both age class I and 4+ were (4) Exotic high (Table 3). Age cks-l sil~valwas signtfi- 0.82 0 0 0 1 caatly greater in exotic forat than rlativc forest 0 0.94 0 0 . = = I (x: 772, I-' 0.005; Table 3). Age class-4+ sur- ( 0 0 0.94 0.94) viva1 was slightly higher in exotic foresr thati native forest, altliough rhc difference was not sig- Overall finite raw of incwxrc wm 1.03 f 0.04 nificant (x: = 2.90, P = 0.089; Table 3 ) . ('L'able 3). Wt- failed to reject Ho: ;l S I (Z= 0.76, Klavitter, J.L.,1. J.M. Wildl. Marzluff, Manage. and 67(M.S. 1 ):ZOO3Vekasy. 2003.HAWAIIAN Abundance HAWK and ABUNDARGEDemography of ANDthe Hawaiian DEMOGRAPHY Hawk: Is Delisting Kfdt&sl Warranted? d Journal171 of Wildlife Mgmt 67(1):165-176. Table 3. Survival, fecundity. and finite rate of increase (5) estimates among all 'io age dasses In native, mixed m-otlc (mixed), exotic, and all habitats combined (overall) beween 1998-1999 on Me Island of Hawaii, USA:Nathte hWIs dominet- ed by natlve vegetallon. Mixed nalive-exotic habttat Is composed of both native and exotlc vegetation, and ex& is domlneted by exotk vegetation. Age dass 1 and age dasses 2.3,and 4+ survival and fecundtty estimates and sample sizeb are shown as %,, ns2, and q,, rnspeMvely. Age 1 Age 2,3.4+ Age 4+ Habtlt suml ng, SE survival nt2 SE fecundity , SE 5 Sf overarl 0.50 28 0.10 0.94 34 0.04 0.23 102 (73) 0.04 1.03 0.04 Nathe 0.27 15 0.1 1 0.88 17 0.08 0.28 SS (44) 0.06 0.95 0.08 Mixed 0.82 11 0.12 0.94 17 0.04 0.17 43(28) 0.06 1.W 0.05 Exotic 0.W8 11 0 12 0.94 17 0.04 0.10 16 (lo) 0.07 1.01 0.06 EsWnale from m~xednalive-exotk was used for the exotk esttmate. Age ciass 4e surwal esbmates were used for age dasses 2. 3, and 4+. No 'b deaths oaxwred m mixed hahitat. We used the 'overall' survival estlmate &r mixed. - Numbers In parentheses are the number of different 'io pain. P= 0.22. 1 - P = 0.20). We found no difference in viva1 held constant, adult sunival needs to exceed h. between native (0.95) and mixed (1.04; % = 0.91% for the finite rate of increase to remain sta- 1.02, P= 0.31). native and exouc (1.01; %=0.24, P ble (h= I) or to increase (1 > I ). Our estimated = 0.81), and mixed and exotic habitats (%= 0.15, adult survi-ml was not statistically different from P = 0.88: Table 2). The long-term stable agr dis 0.91 (H,: S S 0.91, I-tailed, n = 0.05, Z= 0.74, P= tribution of the popr~lation is expected to be 0.229, 1- P = 0.190). 15.6% in age class I. 7.5% in age class 2, 6.9% in age class 3, and 70.0% of the population in the 4+ Other Potential Limiting Factors age cla~s.The avenge time of residence in the 4+ We found no evidence that environmental can- age class is cxpected to be 16.7 years. taminants, avian pox, avian malaria, or I: gondzi Adult su~vivalwa5 the most important parame- limited this popuhdon. Eggs (n = 5) contained ter regulating the popula~ion (elasticity = 0.72). none or trace amounts of organochlorines, PCBs. First-year survival and fea~ndityhad elasticity val- or heavy metals. Plasnra samples (n = 7) tested ues of only 0.87. With fecundity arrd firs~yearsur- negative fimalaria, and only 1 .parnple showed a positive result lor 'I: gmdii (tiler = 25, borderline positive). We tbund 1 adult 'io with avian pox (n = 155), which had healed_lesionson 2 digits of both the left zllld right tiusomelatamus, causing slight deformitits in each. The defolmities did not appear to affect the bird's perching ability. We found low incidence of ectoparasites. Five adult 'io (n = 155) had minor HippoMdae infestatioas. but they did not appear t~ be adversely affecting the birds. We okd4 'io nests and 1 'io chick infested with trcrpical EawI mites (Omicfwnysus ba). When the chick was first observed in the nest (approx 28 days of age). its eyes were nearly swollen shut and it was extremely lethargic. Over the riext 2 weeks, the swellit~gdecreased dnmaucalty and lethargy ceased. The chick successfx~llyfledged, although 1 week early. Fate after fledging was unknown. Fig. 3. 'lo nesllng success on the island of Hawaii, USA, dur- ing the 1998 and 1999 breed~ngseasons. Success Is stratified into 5 habitats that cofredpond to habitat types in Table 1. DISCUSSION 'Native' comb~nesnative forest, nathre torest with grass, and ndxed nativdexotic bresl when dominated by natlve compo- Population Status nents. Txotii include exotic forest. wu-n, and mixed na(rve/exotic forest that were dominated by exotic vegetation. 'io were abundant throughout their entire Numbabm ban are sample sizes. range, and their population appeared stable Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. 1 72 HAWAIIAN HAWK ABL.'NDAN<;E AN11 DEMOGR4PHY Klavdrrcr PI 01. J. Wildl. Manage. 67(1):2003 (Figs. 1, 2). Our current populatjorl estimates al. 1997) 'lo appcar to have saturated mature were similar to those obtained in 1993 (Hall et al. native forest with grass understory. lowe~leva- 1997) and only slightly less than tl~oseestimated Go11 forests, gr.wslands, and plantations with for- in 1983 (GrifEn 1985). Methods used by Griff~n est edges. These habitats have numerous and (1985) to estimate the population were likely pos- accessible perch sites, an operi undelstory, sel- itively biawd, because he assumed 'io were dis- dom-wed dirt roads, and many intrrduced birds ~ributedisland wide at the same high density as in and small mammals (Tomich 1986:37-50. Hill his small study area ( remalrl viable with expected land-use clia~igemay future anthropogcnic changes to the island. include (1) the species' ability to adapt to non- Human population growth and comme~cbl native habitat and prey (Griffin 1985), (2) a lar.ge development on the island will increase over time population size. and (3) protection of substantial (Hawaii State Department of Business, Econmnk portions of the species' natural habitat Development, and Tourism 1998). As of 1998, Ilespite our estimate falling below the downlist- 143.1 35 people resided on the island, an &ate ing target of 2,000 birds (USFWS 19841, our data expected to increase by approxinlawly 2,85Q/year suppon dowr~listirlgthis species to threatened. (U.S. Burcall of the Census 1998). In 199%.773 'Io are clearly not at immediate risk orextinction. new family homes were built on the island We caution again.5~removing 'io from the list of (Hawaii State Department of Busi~wss.Economic species covered by the Entlangered Species Act Development and Tourism 1998). These figures (ESA) of 1973 because theirsrnall popularion size show modes1 growth, but future growth is diflti- (relalive to mainhnd species] m?&a 'io-vuinera- cult to predict FFowth devTopmi3t con- ble to future e~ivironmentalstochasticity (Spiller tinued to a point where all lands (except those et al. 1998) and a~~tl~ropogenicchanges to the protected by reserves, parks. and refuges) were island (Cuddilly and Sro~ltr1990). The idand of converted to urban, we caladated a minimum of Hawaii is comprisecl of 3 active volcanoes any of 244 'io wot~ldstill remain. Thus, the 'io popda- which could quickly change habitat suitability for tion can persist into the foreseeable kture but 'io by ~vplacingforest with law. may always be threatened with extinction. It is Most islands are relatively small in terms of land uncertain if a population at 244 'io would be area. resulting in species with naturally small pop viable, being mostly dependen1 on habitat type, ulation sizes. Based on small populatioti sizes and productivity, and survival (Table 3). We found lirnited di~tributions,by definition, tiearly every that some human-altered hahitat contained 'io islancl species could be considered thr-eatened densities higher than those supported by native under the ESA. One possible altenrative for habitats. However, urban and largescale agricul- islancl species is for federal agencies to adopt spe- turc without forest edge and windrows of large cies classification categoties similar to tt~oseused trees are illcompatible with 'io. by the IUCN, whose designations allow for finer ckt.wifi~lionof species. Recovered and/or sruble and Management island species col~ldbe listed as 'vul~terable" (not critically endangcmd or endangered bu~facing a Although 'io appear viable on Hawaii. their high risk of extincdon in die wild in the medium- abundance and adult survival should be moni- term future). or tifired as 'near ~hrealened" tored periodically. We recommend monitoring (close to qualifying for w Inelable; In ternatiorlal survival wih a mdio~ggingstudy at Syear inter- Union for C~~iservationof Nature 1994). Our vals, tagging a minimum of 30 birds (probably analysis suggests that 'io arc near threatened. r 100 lor sufficient statistical power) From ~ous We have some concenis with basing a reclassif% habitats around the island. To minimize costs, ration on only 2 years of data. Brook and Kikkawa continuous tracking would not be needed. Birds ( 1998) advised cauuo~iwhen using relatively would only need to be tracked at the end of tllc short (4year) periods of dau collection to pre- I-year monitoring period to confirm swvi~lor dirt population stability, especialiv of eridangered mortality. To avoid biasing Stlr~i~lestimates, it species on islands. Beissinger and Westphal would be important to incorporate a measwe of (1 908) recommended a conservative approach trarumitter failure in he study. Mortality may be whrn using deterministic models (finite popula- overestimated. due to birds that are alive but uo11 growth models) to appriise the status of whose tra~ismittershave ceased to transmit due to eaclangerecl species because of demographic esti- malfunction or loss. Using color bands in place of' rnate variability. U'e acknoivledge that in some tadiotag would not be adequate because survival species. a year effect on snmval coultl exist that may be underestimated. We observed I adult might not he detected during short-term srxdies. female change territories and would hme assumed A slight decrease (20.04) in 'iu adult s11114val the bird dead had it not been radiotagged. from our estimate of 0.94 wo~ddcause a down- 'lo abundance should be monitored island ward population tt-djectory. wide using survey methods dewloped by Klavitter Because 'io are found only on the island of' (2000). However, the 'io survey technique prtr Hamii. we are concenied alumt the possiblr duces rather wide confidence itlklvals and cur- Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. 174 t.LAWtUlAN HAWK ARIJNDANCE AND DEMOC; rendy only has the power to detect major changes has increased recently because anthropogenic in the popularion. hmingthe same amount of alteration ol' native forest (increasing the pps variance around the June 1998 density estimate. understory) has allowed 'io densities to douhlc in the population would need to drop 48% LO 700 these habitats (Table 2) where native forest 'io (95% CI: 427-1,147) to detect a significant passerines, including endangered crows. reside. decrease in the population with 95% confidence. Maintaining a stable, rather than increasing, 'io We recommend excluding poor habitats and population is the pnldent course and may allow sampling higherdensity areas w decrease vari- for a greater overall amount of' Hawaiian biodi- ance. Excluded hahitan could hc surveyed pel<- versity LO be conserved. odically (i.e. survey every third suwq period). Fu1u1-e survq should be performed every 5 yearn ACKNOWLEDGMENTS and w~etauonchanges tracked at s~rxypoints fhsmdy msfumied p+mmily+y the USFWS and on a larldscape level using satellite imagery under a cooperative agreement ([ISFWS (hn- and geographic information systems. tract # 1448-12200-98-JO 12). Acldi tional sllpport Althollgh not the prirnary regulating parame- ancl f1111dingwas provided by the Biologicd Re- ter for this popnlarion. reprociuction sho~ddbe search Division of the U.S. Geological Survey moni tc~red.We recomnlerld monitoring a mini- (Kilauea Field Station'), ancl the Geraldine 8. mum of 30 nests in a mixtt~reof habitats dis- Dtxige Foundation. Special thanks are reserved pemd around the island every 10 years. If pm for the 'Ada Recovery Team. C. Atkinson. D. ductivity stl~diesocrurred simultaneously with Ball, M. kny, H. Rrown,J. Rurgett, N. Carroll, T. survival studies, adult f'emales could be mcked Casey, M. Chstillo, K. (2arkson. C. Collins, C. directly to their nests. Since fecundity is currenr- Cunningham, J. Dnbey, J. Enderson. J. Giiffin, P. ly low, it may be one of the few parameters thai Harrity. D. Hu. 'lo Recovery Working Group, K. can be enhanced by mar~agerncnt. Jacobson, J. Jeffrey, S. Johnston, Kau Agribusiness, We recommend ~nanagingfor 'ohi'a trees. Kealia Ranch, P. Kealotra, G. Klir~gler,M. Kochcrr. implementing public educatiorr programs, and 1. Laakc. L. Laniawe. D. Ledig. T. Lum. D. protecting additional habitat. kind managers Mantnval, S. Martin. C. Manllrff. McCandlas need to ensure long-term persistence and contin- Kanrh, H. McDermond, M. Morrison, J. Osen- ued recruitment of native 'o1hi'a trees beca~rsct kowski, M. Reynolds, K Rosa. T. Rubenstein, D. hey are the principle nest tree Tor 'io. Invasive Short,J. Skalski, K. Smith, B. Van Home. M. Wal- exotic vegetation, tire, and intensive pazing cur- ten. I). Was, and T. Work. We thank slate and rently seem to limit 'ohi'a recruitment (Cuddihy federal land managers nc well as the many private and Stone 1990). Pt~bliceducation programs need landowners who allowed us access to their lands for to be continued and improved to protect habitat Lhe study. This project was approved by the Univer- and nest trees, prevent harassment at nest sitcs. sity of Washington (UM7) Animal Care Committee and discourage the purposefill shooting of birds. (UW#3077-04). M. Rochert, I). Manuwal.J. Sk& A large portion of"io habitat currently is protcct- ki, and 2 anonymous reviewers provided excel- eti. hut fcw ol'tlie liighest-densi~varca. Fall within Icnt comments tllat imprc~vedthe rrraniscript. resrlve. ~.cfi~gc.or park boundalies (Fig. 2). Incentives need ro he implementecl tliar encour- LITERATURE CITED age landowners to manage their lands for 'io. AMIu., H. 1973. Il~forrnatiuntheory ancl tlxlc~lsionof Perhapb management activities could be initial- th~-rmaxirnun~ likelihotxl principle. Pages 267-281 irc H. N. l'ctran ~ndI;. (:saki, r.tlitcm. Secol~dintrrna- ed to increase rhc popillation above 2,000 'io. liollal syli~posi~~mon ililbrlnaticltl tlicory. hhderniwi Mangers could alter rni1tu1.c native forest to Kizrdh. Buchpclst. Ht~npry. native forest with grass understory or reinrrodl~ce AK~AK~YA.M. K. 1999. RAMA. CIS: [.inking lmdscapc 'io to other islands where they previously existcd. rhta wit11 pr>puIa~ionviability analysis. Applied Bie based on stlbfossil records (Olson and Jamcs n~athernatirs,Sc~tl~kct. New York USA. KA~WJ.W. E. 1980. Abian history reporl 6A. liislon of 1982, 199 1, 19%). However. serious conse- endcmic Hawaiian birds. Pan I. Pc~p~llationhistb quences to other native communities and birds, ries-species eccou~~ts.<'aope~ative National Park especially endangered ones, could resulr fro~n Resoulre Srt~rlira1Jni1, University of Hawaii. Honolu- such actions. 1)uring the last 4 yea~s.'io preyed lu, USA. BEISSINGER.9. K., MI) 34. 1. WVSI'IIIAI.. 1998. On rhc: LISC upon endangered Hawaiian crows ((lmtuIrnwrn- or de~nograpliicnloclels of population viahiliv ill msu; D. I.. Ball. USFWS, unp~iblisl~edclam). Pre- endangered species rnanagemcn~..joumalof-Wildlife dation nray have occurred historically. but likely manage men^ 62:R:! 1341. Klavitter, J.L., J.M. Marzluff, and M.S. Vekasy. 2003. Abundance and Demography of the Hawaiian Hawk: Is Delisting Warranted? Journal of Wildlife Mgmt 67(1):165-176. 1.Wilrll. Managc. 67(1 ).ZOO3 HAWAILAN H.9\.VK ABUNL>ANCl?AND DEMOGRAPHY Kkrviltwd d 175 B~GER,A. J. 1981. Hawaiian birdlifc. Uniwliity of ca, lnc., Philadelphia, Pennsylvania, UStl. Hawaii Prcu~,Ifot~oIrrlu, USA. I~'ITRK.~~NA~.UNION rn~ CX)NS~?R\'ATION OF NATURE. BER(;Y.R.D. D.,ANI) H. C. ML:ELKZR.1959. Thc Ilakhatri: 1994. Red list utcgories. Species Survival Cornmi* a Lrap for bird9 of prey. Bird Banding 30: 1 8-26. sicir, Gland, Switzerland. BROOK, B. W.. axnJ. RIKMWA.1998. Examining threats h AM. K. 1999. F~ckel'sfrancolin (Arwrcdinw a&&), faced by island birds: a ppuhtion viability anal~is black francolin (fi~~dntcrfimcdinur) , and gray on the Capricorn silvereye using long-tern1 data. francolin (liancdinrcs ptdirminntrr). Pages 1-24 ink Journal of Applied Ecology 35:491-503. Pcmle and F. Gill. ediron. The birds of Nonh Ameri- SUC:KIANI).S. T., U. R. MKKSON.K. A%T)ER%)N, K. P. ca, no. 394-396. The Birds of NohAmerica, lnc., BURNI~U.mn J. I-. IA~KL. 1993. Diutance sampling Philadelphia, Pennsylvania. USA. estimating abundance of biological poptdations. JAWRI. J. D. 1990. Dislribudon maps. ecological rda Chapman and Hall. L~ndon.L'aited Kingclom. tionships, and status of native plant communities on BIIEWER,D, A, 1. D. FRASER,M. K. Rrl~m.1, S. McA, the inland of Hawaii. Dissertation, Universiq of Hawaii ~.yr~,AND J. D. SF&(+. 1995. Captive and fieEd tat- at h4anoa, Honolulu, USA. ed radio attachment techniques f& bad eagleKJ011~- mNtfm,P.-L. I**- goshawk- rlal of Field Orniltlology 66: 173-1 80. piilis): is there evidence of a popdaulan decline? CILFHCII., H. 1989. Matrix population mdcls: coiwtnlc- Jodoiltaptor Research 31:95-106. " rion. analpis, and interpretation. Sinauer Asmiates, Kuwmx,J. L. 2000. Swvcy methodology, abundaace, S~lnderlanci.Masrachiwetu. LISA. and demography of the endangered Ha-iian hawk -. 2000. Prospecrive and retrospective' pertur-ha- is delishg warranted? Thesis, University of Washing- tion analyses: ~tieirmles in ronse~~-ationbiology. ton. Seattle. USA. Ecolow 8 1:61%627. ~A%M,J.l... S. T. BBUCXIAND,D. R. ANDERSIN, AND K. P. CI-.\\KKUSON,K. AND I.. LMIAHT. 200. Hawaiian hawk BLTRNIIAM.1Y93. Dis~anceuser's guide, version 2.0, (Bum solitnnuc). Pa~cs1-1 6 irr A. Poole and E Gill, @e.p9prdatSirm,mrate. bbgy 8 1 :#42453. -. 1994. Endangemi and threatelled wildlife and &., AND S. A. TMPLE.1988. Breeding biolrk plants; withdrawal oi proposed nrle lo reclassify the gp and dlet d &iJed hawks in Pilertrb Kicr). Hawiliian hawk (ih:Im sdikrfi:rcs) frcjn~endangered to Biotropica 20: 15 1-1 60. threatend. 1J.S. Ash and Wildlife Selvice, Honolulu. %XT,J. M..J. D. JACC)BI. AND I? I.. R\MsE~,.1981. Avian llawaii. USA surveys of large gewaphi~i~lareas: a sy?ilcmatic . 1999a. U.S. Fish and Wildlife Service proposes approach. Wildlife Society Bulktill 9:190-200. to remow Tinian monarch fror~iEndangered Species , S. MOUNTAINSPKINC;,E R~MS~Y,*D C'. KLPIEK. Act protection. U.S. Fish and Wildlife Semce. Hoti- 1986. Forest bird communities of the I-lawaiian ol~~lu.Hawaii, USA. islmds: their dyna~nics. ecolcbgy. and ronscwitio~i. . 1999b. Listinp and recovery plans. Studies in Avian Biology No. 9. hltp://enclangerrcl.hus.g~~/bc~~.scorr.html.Accessed SEHBKG. A. F. 1973. The rstimaiion of animal abun- 27 October 1999. dance. Griffin, Inc., London, United fi~fiom. . 2000. US. Ks11 and Wildlife Service. Pacific SFIU!ZR.D. A., J. B. IDSOS,AND T. W.SCJ-IDENEK. 1998 Islands Ecoreion homepgei endangered species. Impact of a catmtrophic hurricane on island popula- h~~p://mrw.r.l.fws.gov/pacific/wesa/en~indm.ittrn tions. Science 28 1:695697. I. Accessed 2 May 2000. SPOT. 1994. Systelnc pour I'observadon de la Lrnp VAH WR111, C., S. G, VAN RLPPFJ~, M. L. Wm, ~WI)M. (SPOT).Hawaii statewide XS coverage. CD-ROM. IAIIU).1986. The epizootiology and ecological signifi- order 3467006003. SPOT lma~cCi)~poratio~i, Rmto~i. cance ol' malaria in Hawaiiau lalid birdq. Z~~lokficid Virginia, USA. Monog-nphs 56:327-344. Smutr~~.K. 1987. thesing rapror rrp~~ucuvcstlrcm VFX~~Y.M. S.. J. M. M~~zl.llW,M. N. KW~~LKI,R. N. and productivi~y.Pages 151-170 ~trB. A. Giro11 I'r~~dle- 1,nru~~.ANI) K !~TENHOF.1996. Inflr~enceof r~dio ron. B. A. Millsap, K W. Cline. slid D. M. Rird, edilors, transmirto~on prairie falcons. Journal of Ficid Kapcor management techniques manual. National Ornithology 67:680-690. Wildlife Federation. Waslii~~gton,D.C., USA. WISUOM.M.J.. 1.. S. MILL\AND D. F. DOAK.2000. lifestagc . AND M. N. KOCHERI. 1982. An evaluatioii of .uimulaljonanalysis: estimating vital-mtc vffccts on pop methocis used to estimate raptor ~lclstingsuccess. ulation gnnvrh tar ronservatio~~.Ecology 8 1 :628-641. Journal of Wildlife Managrlnent 46385-93. W( 'OLJRAM RF~FAUW. I W9. MKIWENWTICA. wolf ran^ TI~OMW.L.. J. LAAXE. J. ti. DEKRY,S. UUCUAND, I). Kmearch, Chatnl>aip~.Illinuis, USA. R~RCW.ILS.D. ANDER~~)N. K IJL~KNHAM.S. SIRINDREKC;. WCIW T. M., J. C. Mwscn, B. k b~>r.cxrr,C. H. GAR- S. 1. H~nl~s,M. 1.. 611~1,F. MARQUES,.^. H. P(>I.IMI). I)INER. D. B. IEIII(;.<). c. H. KwOR, AND I. ?. UUREE ANI) K. M. FF.ws?w. 1998. Distance version 3.5, relrasct 2000. ~amltoxopJasalr~is in freeranging &r~ln~eI'ed 5. Kcsea~cl~Unit Ibr Wilclliii Popularion Assesr~ten~, 'alala from Hawaii (Chms hnwrtiisnsts). Journal of Unive~rityof St. Antlrews, L'IIIL~~Kingdom. Wildlife 13isews 36:20.S-2 12. T~MIc:?~.F? Q 1986. Mammals it1 Hawair. Bisl~opM~ar- ~AR,J. H. 1%. Bn9statistjcal atrtl&s. Prenrice-H111, tun Press, Ho~iolulu,Hawaii. USA. 1:pprr .Saddle Kjvcr, NewJc~sqUSA. 1I.S. BUWLI