Forest Bird Communities of the Hawaiian Islands: Their Dynamics, Ecology, and Conservation J MICHAEL STEPHEN SCOTT MOUNTAINSPRING U.S Fish and Wildlife Service Patuxent Wildlife Research Center Mauna Loa Field Station Hawaii National Park, Hawaii 967 18 FRED L RAMSEY Department of Statistics Oregon State University Corvallis, Oregon 9733 and CAMERON B KEPLER U.S Fish and Wildlife Service Patuxent Wildlife Research Center Maui Field Station Kula, Hawaii 96790 Drawings of native birds by H DOUGLAS PRATT Studies in Avian Biology No A PUBLICATION OFTHECOOPER ORNITHOLOGICAI SOCIEI’Y Cover Photograph: Iiwi (Vestiorio coccineo) perched on kolii (Tremotolobelio kouoiensis) on Kauai, by David Boynton STUDIES IN AVIAN BIOLOGY Edited by RALPH J RAITT with the assitance of JEAN P THOMPSON at the Department of Biology New Mexico State University Las Cruces, New Mexico 88003 EDITORIAL Joseph R Jehl, Jr ADVISORY Frank A Pitelka BOARD Dennis M Power Studies in Avian Biology, as successor to Pacific Coast Avifauna, is a series of works too long for The Condor, published at irregular intervals by the Cooper Ornithological Society Manuscripts for consideration should be submitted to the current editor, Frank A Pitelka, Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720 Style and format should follow those of previous issues Price: $26.50 including postage and handling All orders cash in advance; make checks payable to Cooper Ornithological Society Send orders to James R Northern, Assistant Treasurer, Cooper Ornithological Society, Department of Biology, University of California, Los Angeles, CA 90024 Current address of J Michael Scott: Idaho Cooperative Fish and Wildlife Research Unit, College of Forestry, University of Idaho, Moscow, Idaho 83843 Library of CongressCatalog Card Number 86-7 1720 Printed by Allen Press, Inc., Lawrence, Kansas 66044 Issued 29 August, 1986 DEDICATION We dedicate this book to all those who participated in the arduous and hazardous field work for these studies A special debt is owed to Eugene Kridler (right), first U.S Fish and Wildlife Service biologist stationed in the islands, who often went out on an administrative “limb” to support and encourage us, and to John L Sincock (left), who spent many raindrenched nights alone in the forest pioneering field techniques Without the help, encouragement, and example of these two, the Hawaii Forest Bird Survey would still be a dream 111 CONTENTS I~R~DuCTI~N THE SURVEY AND ITS OBJECTIVES THE NATURAL ENVIRONMENT Geology Climate Vegetation STUDY AREAS Kau Hamakua Puna Kipukas Kona , Mauna Kea Kohala East Maui West Maui Molokai Lanai Kauai FIELD METHODOLOGY Establishment of Transects Observer Training Bird Sampling Vegetation Sampling Insect Observations DATA ANALYSIS Estimation of Effective Area Surveyed Birds per Count Period Range Determination Population Estimates Unrecorded Species Original Ranges Analysis and Interpretation of Habitat Response Habitat variables Community variables Preliminary screening Regression models Habitat response graphs Interpreting habitat response Interspecific Competition Species-Area Relationships Comparison with Earlier Studies Survey Limitations NATIVE SPECIESACCOUNTS Hawaiian Goose (Nene) Hawaiian Hawk (10) Hawaiian Rail (Moho) iv 5 6 15 15 15 17 18 20 24 24 24 31 31 32 32 33 33 34 37 40 45 48 48 52 53 53 54 55 55 55 56 56 57 58 59 61 61 61 61 61 72 78 79 Lesser Golden-Plover (Kolea) Short-eared Owl (Pueo) Hawaiian Crow (Alala) Elepaio Kamao Olomao Omao Puaiohi (Small Kauai Thrush) Kauai 00 (Ooaa) Bishop’s00 Hawaii 00 Kioea ou Palila Lesser Koa-Finch Greater Koa-Finch Kona Grosbeak Maui Parrotbill Common Amakihi Anianiau Greater Amakihi Hawaiian Akialoa Kauai Akialoa Nukupuu Akiapolaau Kauai Creeper Hawaii Creeper Maui Creeper Molokai Creeper Akepa Ula-ai-hawane Iiwi Hawaii Mamo Black Mamo Crested Honeycreeper (Akohekohe) Apapane Poo-uli 1NTRODUCED’SPEC;ESACCOL;;TS’:::::::::::::::::::::::::::::::::::: Black Francolin Erckel’s Francolin Gray Francolin Chukar Japanese Quail Kalij Pheasant RedJunglefowl Ring-necked Pheasant Common Peafowl Wild Turkey California Quail V , 80 81 82 86 93 96 96 101 103 106 106 107 107 111 114 114 114 115 117 128 128 130 130 131 133 139 142 146 148 149 156 157 163 167 168 170 182 183 183 199 200 206 211 211 217 218 226 226 234 Rock Dove Spotted Dove Zebra Dove Mourning Dove Common Barn-Owl Eurasian Skylark Japanese Bush-Warbler White-rumped Shama Melodious Laughing-thrush Red-billed Leiothrix Northern Mockingbird Common Myna Japanese White-eye Northern Cardinal Saffron Finch House Finch Yellow-fronted Canary House Sparrow Red-cheeked Cordonbleu Lavender Waxbill Warbling Silverbill Nutmeg Mannikin COMMUNITY ECOLOGY Species-Area Relationships Richness and Diversity General Patterns of Habitat Response Distributional Anomalies LIMITING FACTORS Habitat Modification Browsers, grazers, and rooters Introduced plants Anthropogenic habitat degradation Predation Disease Interspecific Competition Disasters CONSERVATION History of Human Disturbance Conservation Strategies Extinction Models Island Recommendations Conclusion ACKNOWLEDGMENTS SURVEY PARTICIPANTSAND HABITATS LITERATURE CITED APPENDIX TABLES vi 235 235 247 248 252 252 254 260 261 266 277 282 287 295 307 314 317 317 321 321 321 325 330 330 334 340 349 351 352 352 361 362 363 364 368 371 371 371 373 380 380 384 385 387 393 406 TABLES Table Table Table Table Table Table Table Table Table Table 10 Table 11 Table 12 Table 13 Table 14 Table Table Table Table Table Table Table 15 16 17 18 19 20 Table Table Table Table Table 22 23 24 25 26 Table 27 Table 28 Table Table Table Table Table Table 29 30 32 33 34 Table 35 Table 36 Table 37 Table 38 Table 39 Table 40 Table Table 42 Table 43 Table 44 Status and distribution of endemic Hawaiian birds Native tree and shrub genera on Kona, East Maui, and mature dry forest sites Hawaiian Forest Bird Survey study areas Number of stations sampled by elevation, habitat, and study area Adjustment factors for the effects of habitat configuration on effective area Analysis of variance for the effect of species, observer, and habitat configuration on effective detection distance Effective detection distances for Hawaiian birds Multiplicative factors for effective areas by elevation, habitat, and study area Habitat and area in assumed original range of native birds Summary statistics for native birds in the study areas on Hawaii Summary statistics for native birds in the study areas on Maui, Molokai, Lanai, and Kauai Probability of detecting at least one bird of species unrecorded during the HFBS Density of the Hawaiian Goose (Nene) and Hawaiian Crow (Alala) Regression models for habitat response of the Hawaiian Goose (Nene) and Hawaiian Crow (Alala) Incidental observations of the Hawaiian Hawk (10) Density of the Elepaio Regression models for habitat response of the Elepaio Density of the Kamao, Olomao, Omao, and Puaiohi Regression models for habitat response of the Kamao, Omao, and Puaiohi Density of the Kauai 00, Ou, Palila, Maui Parrotbill, Anianiau, and Nukupuu Regression models for habitat response of the Ott, Palila, Maui Parrotbill, Anianiau,andAkiapolaau Density of the Common Amakihi Regression models for habitat response of the Common Amakihi Density of the Akiapolaau and Poo-uli Density of the Kauai Creeper, Hawaii Creeper, and Maui Creeper Regression models for habitat response of the Kauai Creeper, Hawaii Creeper, and Maui Creeper Density of the Akepa and Crested Honeycreeper (Akohekohe) Regression models for habitat response of the Akepa and Crested Honeycreeper (Akohekohe) DensityoftheIiwi Regression models for habitat response of the Iiwi Densityofthe Apapane Regression models for habitat response of the Apapane Summary statistics for introduced birds in the study areas on Hawaii Summary statistics for introduced birds in the study areas on Maui, Molokai, Lanai,and Kauai Density of the Black Francolin and Gray Fancolin Regression models for habitat response of the Black Francolin, Erckel’s Francolin, and Gray Francolin Density of the Erckel’s Francolin Density of the Chukar and Red Junglefowl Regression models for habitat response of the Chukar, Kalij Pheasant, and Red Junglefowl Density of the Japanese Quail and Kalij Pheasant Density of the Ring-necked Pheasant Regression models for habitat response of the Ring-necked Pheasant and Common Peafowl Density of the Common Peafowl and Wild Turkey Regression models for habitat response of the Wild Turkey and California Quail vii 11 37 38 48 51 52 53 54 62 65 69 73 77 78 87 93 95 97 105 113 126 127 134 139 141 154 155 158 167 171 181 184 191 196 199 200 206 211 212 219 220 227 230 Table Table Table Table Table Table 45 46 47 48 49 50 Table 51 Table Table Table Table Table Table 52 53 54 55 56 57 Table Table Table Table Table 58 59 60 62 Table 63 Table 64 Table 65 Table Table Table Table 66 67 68 69 Table Table Table Table 70 71 72 73 Table 74 Table 75 Density of the California Quail Density of the Spotted Dove Regression models for habitat response of the Spotted Dove and Zebra Dove Density of the Zebra Dove and Mourning Dove Density of the Eurasian Skylark Regression models for habitat response of the Eurasian Skylark, Japanese BushWarbler, and Northern Mockingbird Density of the Japanese Bush-Warbler, White-rumped Shama, and Northern Mockingbird Density of the Melodious Laughing-thrush Regression models for habitat response of the Melodious Laughing-thrush Density of the Red-billed Leiothrix Regression models for habitat response of the Red-billed Leiothrix Density of the Common Myna Regression models for habitat response of the Common Myna, Saffron Finch, and Yellow-fronted Canary Density of the Japanese White-eye Regression models for habitat response of the Japanese White-eye Density of the Northern Cardinal Regression models for habitat response of the Northern Cardinal Density of the Saffron Finch, Yellow-fronted Canary, Red-cheeked Cordonbleu, Lavender Waxbill, and Warbling Silverbill Density of the House Finch Regression models for habitat response of the House Finch Regression models for habitat response of the Warbling Silverbill and Nutmeg Mann&in Density of the Nutmeg Mann&in Regression models for habitat response of native species richness Regression models for habitat response of introduced species richness Regression models for habitat response of bird species diversity (Simpson’s Index) Relative importance of habitat variables Elevational and lateral distributional anomalies Response of native birds to mosquito presence Distribution of negative and positive partial correlations across study areas by native or introduced status of the members of each species pair Percentages of negative partial correlations among primary and secondary potential competitors in native/introduced species pairs Status and management recommendations for native Hawaiian forest birds Vlll 235 246 247 248 255 260 261 262 269 270 277 281 287 288 296 305 306 307 308 319 323 326 336 337 338 341 349 367 369 370 374 FIGURES Figure Figure Figure Figures Figures Figures Figures 4-8 9-11 12-15 16-18 Figures 19-21 Figures 22-24 Figures 25-27 Figures 28-30 Figures l-33 Figures 3436 Figures 37-39 Figures 40-42 Figures Figure 43-44 45 Figures Figure Figure 46-60 61 62 Figure Figures 63 64-68 Figure Figures Figures Figures Figure Figures Figures Figures Figures Figures Figures Figures 69 70-71 72-79 80-81 82 83-86 87-88 89-90 l-94 95-96 97-98 99-109 Figures Figures Figures Figures Figures Figures Figures Figures 110-111 112-113 114-118 119-120 121-124 125-126 127-133 134-144 Map of the Hawaiian Archipelago Field crew for the Kau forest bird survey of 1976 The main Hawaiian Islands Vegetation zones Study area locations Place names Transect locations, habitat types, and canopy cover in the Kau study area Transect locations, habitat types, and canopy cover in the windward Hawaii study areas Transect locations, habitat types, and canopy cover in the Kona study area Transect locations, habitat types, and canopy cover in the Mauna Kea study area Transect locations, habitat types, and canopy cover in the Kohala study area Transect locations, habitat types, and canopy cover in the East Mauistudyarea Transect locations, habitat types, and canopy cover in the West Mauistudyarea Transect locations, habitat types, and canopy cover in the Molokai study area Transect locations, habitat types, and canopy cover in the Lanai study area Transect locations and canopy cover in the Kauai study area Observer at top of transect prepared for lo-day bout in the rainforest Photographs of typical habitat in the study areas The cumulative detection curve and its envelope Relative abundance of dominant tree species in forest and woodland habitat types on Hawaii and Maui Sample sizes for cells on the habitat response graphs Distribution, abundance, and habitat response of Hawaiian Goose Distribution of the Hawaiian Hawk on the island of Hawaii Distribution, abundance, and habitat response of Hawaiian Crow Distribution, abundance, and habitat response of Elepaio Distribution and abundance of Kamao Distribution and abundance of Olomao Distribution, abundance, and habitat response of Omao Distribution and abundance of Puaiohi Distribution and abundance of Kauai 00 (Ooaa) Distribution, abundance, and habitat response of Ou Distribution, abundance, and habitat response of Palila Distribution, abundance, and habitat response of Maui Parrotbill Distribution, abundance, and habitat response of Common Amakihi Distribution, abundance, and habitat response of Anianiau Distribution, abundance, and habitat response of Nukupuu Distribution, abundance, and habitat response of Akiapolaau Distribution, abundance, and habitat response of Kauai Creeper Distribution, abundance, and habitat response of Hawaii Creeper Distribution, abundance, and habitat response of Maui Creeper Distribution, abundance, and habitat response of Akepa Distribution, abundance, and habitat response of Iiwi ix xii 8-10 15-16 17-19 19-20 21-23 24-26 27-28 28-29 30-3 32-34 35-36 36,38 39 40 41-50 51 58 59 73-76 79 83, 84 87-92 94 95 97-100 102 104 108-l 10 112 116 118-125 129 132 134-138 140 142-145 147-148 149-153 159-166 420 STUDIES IN AVIAN BIOLOGY hhh : : : : N-40 : - .Rxs ::: 0000;;;; j; : : : : : : : : : : : : hhh T‘ ,,,d-43+ : -,,,,o: am~vr+ +++ ovvvvvv i,,,,: : : : 000000 CC? c_~ “” : : : : : : : : : : ooooooooof j; i NO HAWAIIAN hhh :::::oclom 427 FOREST BIRDS r‘-4t-40’;::::: wvv =‘ w: : : : : :g 4’ “ “C-4 w : : : : :ooo .cT v WV -+ -hhhhhh +dd4c.Jmm ::.: : -‘ ’ : -36wmoo-W’ -n-n ~oooLC~2~:::f -mmT; :: VI’ : ’ s,:: ;: : ” : ” : sci -1 r‘ ncr :: :o: W”“’ : : :oo;;; :::::::::oooooo c Gi _ w::::: b’ hhhh ; ++++ ovvwvovvvo ++-+ :ooooo -o~ ‘ ?Tc +mm T?G ++m : : ‘.’ f: : : ‘ zz,: : ; “f f ” t‘-4 00 ’ z,: : : : :z + + 430 STUDIES IN AVIAN BIOLOGY NO 2: :s :::: + _ : : : : :ooo s: : : ;o-:7 + + + w +’ : “ + T? VW : T 00-o + Y 05 + +’ : 10 : : : 00 j i ?T ooooo- ? ++ + hhh +++ omo _ + - + o^ -’ : : _ : : : : hh ++ ++ w-0: hh ++ ’ ++’ : : O ;0 + + -0 : .’ :o ... the landing of Captain Cook in 1778, including an entire group of large, flightless geese, at least eight rails, and a constellation of lowland dry habitat passerines In the 200 years since Western... EX NE EX EX STUDIES IN AVIAN NO BIOLOGY TABLE CONTINUED Taxa Hawaii Maui Molokai Lanai Oahu Kauai NWHI Apapane Himatione Himatione sanguinea sanguinea sanguinea freethii NE NE NE... first reliable listings of the birds of the Hawaiian Islands were by Dole (1869, 1879) Ornithological interest in the islands increased dramatically in the second phase, beginning with the last