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Studies in Avian Biology No. 22:31-46, 2001.

PATTERNS OF SUCCESS AMONG INTRODUCED IN THE HAWAIIAN ISLANDS

MICHAEL I? MOULTON, KARL E. MILLER, AND ERIC A. TILLMAN

Abstract. At least 140 speciesof 14 different orders of birds have been introduced to the six main Hawaiian Islands. The introduced came from six continents and the introductions were carried out by a variety of agents including state and local governments, private citizens, and the acclimati- zation society known as the Hui Manu. The introductions mostly occurred during the early to mid- twentieth century. Most (79%) of the intentional introductions were of species from three orders: , Columbiformes, and Passeriformes. Introduction success rates were significantly greater for passeriforms than for either columbiforms or galliforms, although the reasons for this are unknown. In predicting the fate of future introductions, only the columbiforms showed an “all-or-none” pattern of introduction history. Successful species had larger native geographic ranges than did unsuccessful species, which supports the hypothesis that range size is correlated with the ability to adapt to a new environment. Finally, in a partial test of the introduction effort hypothesis we found that galliforms successfully introduced to the island of Hawai‘i were introduced in significantly larger numbers than unsuccessful species. Key Words: doves; game birds; introduced species; introduction effort; introduction success; native range size; perching birds; pigeons.

Numerous species of birds from six continents (Moulton 1993; Moulton and Pimm 1983, have been introduced to the Hawaiian Islands 1986a). Second, there is a pattern of limiting (Caum 1933, Berger 1981, Long 1981, Pratt et al. similarity among congeneric pairs of introduced 1987). These species were introduced by a variety birds: differences in bill length are significantly of groups for a variety of reasons. As noted by greater in pairs that coexist than in pairs of spe- Berger (1981), the first avian introduction came cies that were not able to coexist (Moulton with early Polynesians who brought the Red Jun- 1985). And third, successful introduced passer- glefowl (Gallus gallus) for food. Since that time, ines show a pattern of morphological overdis- a number of private citizens have brought species persion (Moulton and Pimm 1987, Moulton and to ‘ (e.g., Caum 1933). Some of these in- Lockwood 1992); i.e., successful species are troductions were made inadvertently as individual morphologically more different from each other birds escaped captivity (e.g., Melodious Laughing- than expected by chance. thrush or Hwamei, Garrulax canorus, on Oahu),‘ Although these three patterns are consistent whereas others were intentionally released for aes- with predictions from competition theory, other thetic reasons or even as an attempt at biological explanations for patterns in introduction out- control (Caum 1933). There also have been inten- comes have been advanced. These include intro- sive efforts both by private citizens (e.g., Lewin duction history of a species (Simberloff and 1971) as well as state and county agencies Boecklen 1991) and introduction effort (e.g., (Schwartz and Schwartz 1949; Walker 1966, 1967) Pimm 1991, Veltman et al. 1996). to establish populations of various game birds for The idea that introduction history can predict recreational hunting. In the early to mid-twentieth future introduction outcomes is appealing in its century, the acclimatization society known as the simplicity. The concept comes from Simberloff Hui Manu actively introduced several species to and Boecklen (1991) who argued that whenever various islands (Caum 1933, Berger 1981). and wherever a given species is introduced, it Regardless of their source, a central question tends to either always succeed or always fail. in any study of introduced birds is “Why do This leads to an “all-or-none” pattern in the dis- some species succeed and others fail?” In sev- tribution of birds introduced onto a series of is- eral papers we and our colleagues have argued lands: some species being successful on “all” that competition has played an influential role in the islands in the series and others being suc- determining the outcomes of species ’ cessful on “none” of the islands. If introduced introductions in Hawaii‘ (Moulton and Pimm birds actually follow this pattern, then predicting 1983, 1986a, 1987; Moulton 1985, 1993; Moun- the outcome of future introductions would be tainspring and Scott 1985; Moulton et al. 1990; greatly simplified. Moulton (1993) and Moulton Moulton and Lockwood 1992). These arguments and Sanderson (1997), however, argued that the are based on three main findings. First, intro- all-or-none pattern reported by Simberloff and ductions tend to be less successful when more Boecklen (1991) for passerine birds was pri- species of introduced birds are already present marily an artifact of sample size.

31 32 STUDIES IN AVIAN BIOLOGY NO. 22

Another factor that might influence the out- successful if they were present on an island in 1990. come of introductions is the effort invested in We considered species to be unsuccessful if there were the introduction process. Griffith et al. (1989) no recorded observations after 1990. Scientific names found that introduction effort along with of 140 species introduced in the Hawaiian Islands are provided in Appendix I. Scientific names of intro- quality were associated with introduction out- duced species not included in our statistical analyses come. Similarly, Pimm (1991) studied introduc- are provided in Appendix 2. tions of seven game species (all of which In order to determine success rates for the species in had been successfully introduced somewhere in the different orders, we considered a species to be suc- the world) in the western and cessful if it succeeded on any island, and unsuccessful found that there was a very high (3601424 = only if it failed on every island on which it was released. 85%) failure rate. Pimm’s analysis indicated that By this approach, even if a species fails on all but one the failure rate was particularly high when fewer island, we believe that environmental conditions in the than 75 individuals were released. More recent- archipelago overall were potentially suitable for establish- ment and that perhaps differences in the mechanics of the ly, studies of introduced birds in New Zealand release or interactions with other species might have oc- (Veltman et al. 1996, Duncan 1997, Green 1997) curred on islands where the species failed. We compared have concluded that introduction effort is the introduction success rates across orders with a chi-square most influential variable in determining which test of equal proportions. species succeed. In each of the three studies, the We used range maps in Long (1981) to estimate authors reported that successful species were in- native range size for all introduced species, except troduced in larger numbers and more frequently Garrulax cuerulat~s and douglasii, which than were unsuccessful species. were not included by Long. We used a grid method Several authors have reported a positive re- similar to the methods of Moulton and Pimm (1986b). We placed a small acetate grid over the native range lationship between the size of the native geo- map in Long (1981) and counted the number of graphic range of a species and its average abun- squares that were intersected. Each square represented dance (e.g., Bock and Ricklefs 1983, Brown approximately 259,000 km’. In earlier analyses of na- 1984). If widespread species tend to be ecolog- tive range size of introduced in Hawai‘i ically more generalized than species with narrow (Moulton and Pimm 1986b), Urueginthusangolensis distributions, we would predict that successful and C/. cyanocephala were omitted because of concern introduced species would tend to be those that about the potential confusion with young U. hen&us have larger native ranges. in the field. However, we included all three Uraegin- thus species in this analysis because Berger 981) re- Many analyses of avian introduction success (I ported each was seen and identified in the wild. in Hawai ‘i have focused on passerine birds (e.g., We used Mann-Whitney tests for all our range size Moulton and Pimm 1983, 1986a,b, 1987; Wil- comparisons because data were not normally distrib- liams 1987, Moulton and Lockwood 1992) yet uted. We compared native geographic range sizes of passerines represent fewer than half the total successful versus failed introductions, both within and number of birds that have been introduced to the across orders. Hawaiian Islands (Berger 1981, Long 1981). Our objectives in this paper were to examine RESULTS patterns of success for introduced species in Ha- At least 140 species of nonindigenous birds wai‘i across three taxonomic orders of birds: from 14 orders have been released in the Ha- Galliformes, Columbiformes, and Passeriformes. waiian Islands (Table I). Our results differ from Specifically, are the success rates of nonpasser- earlier totals of 162 species (Long 1981) and ine birds different from those of the passerines? 170 species (Berger 1981) for two reasons. First, Second, is an all-or-none pattern evident in the those authors followed a somewhat different tax- nonpasserine orders‘? Third, is native range size onomy. For example, Berger (1981) listed the greater for successful introduced species than for Green (Phusianus versicolor) as being unsuccessful introduced species in passerines a distinct species, whereas we followed Sibley and nonpasserines? And, fourth, does introduc- and Monroe (1990) and treated it as being con- tion effort play a role in determining the success specific with the Ring-necked Pheasant (Phasi- of introduced birds in Hawai‘i? unu.s colchicus). Second, at least among the pas- serines, we have excluded several species in- METHODS AND MATERIALS cluded by Long and Berger on grounds that sim- We used Caum (1933) Schwartz and Schwartz ply too few individuals (i.e., < 5) were released. (1949), Munro (1960), Walker (1966, 1967), Lewin Simberloff and Boecklen (1991) list 14 of these (1971) Berger (1981), Long (1981), Lever (1987), and species in their Appendix B, although based on Pratt et al. (1987) to compile lists of nonindigenous birds introduced to the Hawaiian Islands. In compiling Berger (1981) we included the two Uraeginthus our lists we attempted to ascertain not only the current species (U. angolensis and U. cyanocephala). status of each species but also the date of first intro- Although a great diversity of species has been duction. In our analyses we considered species to be released into the Hawaiian Islands, for the most INTRODUCED BIRDS-Moulton, Miller, and Tillman 33

TABLE 1. SIJECIES OF BIRDS INTRODUCED TO THE HA- troductions: Galliformes, Columbiformes, and WAIIAN ISLANDS (CAUM 1933, BERCEK 1981, LONG Passeriformes. 1981) HISTORICAL PERSPECTIVE Numbrr 01 O&r SptXitX In order to develop a historical perspective on Tinamiformes 1 the phenomenon of introductions for the galli- Pelecaniformes 1 forms, columbiforms, and passeriforms, we - Ciconiiformes 3 egorized introductions by time period (Fig. 1). Falconiformes I Historical peaks in the number of introductions Galliformes 40 were evident for each order. Turniformes 1 For galliforms, the number of species’ intro- Gruiformes I ductions increased steadily from 1901 until the 2 4 early 1960s and then declined to zero. There has Columbiformes 18 not been an introduction of a new species of Psittaciformes 14 galliform into the Hawaiian Islands since 1965 Strigiformes I (Fruncolinus udsperus). For columbiforms, the Apodiformes I peak occurred in the 1920s. Indeed, there have Passeriformes 52 been only two introductions (Zenuidu asiuticu in 1961 and Zenuidu mucrouru in 1962) of species from this order since 1960. The passeriforms part three orders accounted for the bulk of the also appear to show a decline in the number of introductions. These are the game birds (Galli- introductions after the 1960s (Fig. 1). Closer in- formes), pigeons and doves (Columbiformes), spection reveals an even sharper decline in the and perching birds (Passeriformes). These spe- frequency of introductions, with only one new cies represent 110 introductions (Appendices 3- passerine species introduced since 1980 (Estril- 5). Berger (1981) lists 14 species of a fourth du ustrild in 1981). The remaining nine species order, Psittaciformes. However, according to were all present on other islands in the archi- Berger (1981), 13 of these species were acci- pelago prior to 1975 and possibly arrived onto dental introductions. Moreover, Pratt et al. new islands via interisland colonization. (1987) considered only one species of this order SUCCESS RATES (Psittucula krumeri) to be successful in Hawai ‘i. Thus, we restricted our tests to the three orders Success rates differed significantly among or- for which there was evidence for intentional in- ders (x’ = 14.59, df = 2, P < 0.005). Among

_

pre-1876 1876-1900 1901-25 1926-50 1951-75 post-l 975 Year

n Galliformes Columbiformes 0 Passeriformes

FIGURE 1. Chronology of species introductions to the Hawaiian Islands. 34 STUDIES IN AVIAN BIOLOGY NO. 22

0 : Hawaii Maui Lanai Molokai Oahu Kauai Island

n Galliformes Columbiformes q Passeriformes 1

FIGURE 2. Success rates (number of successfulintroductions/total number of introductions)per order across the six main Hawaiian Islands.

passerines, 33 of 52 (64%) species have been Oahu‘ (five to Kauai‘ and two to Hawaii).‘ For successful on at least one island (Appendix 3). galliforms, only Oahu‘ and Hawaii‘ have any The success rates for galliform and columbiform unique species. species were not nearly so high. Only 12 of 40 (30%) introduced galliform species (Appendix ALL-OR-NONE PATTERNS 4) and 4 of 18 (22%) introduced columbiform The hallmark of an all-or-none distributional species (Appendix 5) have been successful on at pattern of introduced birds on islands would be least one island. presence of few, if any, mixed species. Mixed Within islands the success rates also were species are those that are successful on some variable (Fig. 2). For passerines, Molokai‘ and islands and unsuccessul on others (Simberloff Lanai‘ shared the highest rates of success at 1.00 and Boecklen 1991). In principle, species re- (13/13 for Molokai‘ and ll/ll for Lanai).‘ leased onto one island could show a mixed out- Lanai’ also had the highest success rate for gal- come if they spread to another island and then liforms (9/15, 0.60), whereas Molokai‘ had the fail on one of the two islands. In practice this is highest rate for columbiforms (3/4, 0.75; Fig. 2; very difficult to detect, because those species Table 2). Although it is tempting to compare with the ability to spread to other islands could rates among islands across the different orders, do so repeatedly giving the impression that they results of any tests would be misleading because were established on the second island even if of the high potential for nonindependence. For they were actually not able to survive there. This example, with respect to passerines, only seven would be an example of what Brown and Ko- species were introduced to islands other than dric-Brown (1977) have termed a “rescue ef- fect.” With this in mind we believe that analyses for all-or-none patterns should be limited to TABLE 2. SUCCESS RATES (NUMBER OF SUCCESSFUL those species that were physically introduced to INTRODUCTIONS/TOTAL NUMBER OF INTRODUCTIONS) PER more than one island. ORDERACROSS THE SIX MAIN HAWAIIAN ISLANDS In their analysis of introduced Hawaiian birds, Simberloff and Boecklen (1991) reported that IFland Gahformes Columbiforme3 Passeriformes among 19 introduced columbiform species, only Hawaii ‘ 0.32 0.56 0.80 one (Pterocles exustus) showed a mixed out- Maui 0.45 0.27 0.84 come, having succeeded on Hawaii,‘ and failed LZnai‘ 0.53 0.43 1.00 on Molokai‘ and Kauai.‘ However, Sibley and Molokai‘ 0.53 0.60 1.oo Monroe (1990) placed this species in the order Oahu‘ 0.26 0.33 0.60 Ciconiiformes. If this species is excluded, 18 Kauai‘ 0.40 0.375 0.75 columbiform species remain, 11 of which were

36 STUDIES IN AVIAN BIOLOGY NO. 22

TABLE 4. NUMBER OF INDIVIDUALS OF SPECIES OF for the three orders we have focused on here, ap- GAMEBIRDS,PIGEONS,AND DOVES RELEASEDON HAWAII‘ pears to be historical, with most introduction ef- (LEWIN 197 1) forts having come to a close. There have been no columbiform or galliform introductions to the Ha- Number waiian Islands in more than 30 years. Moreover, SptXE3 released StatUS no new passerine species have been introduced to virginianus” 108 F the islands since 1981. This is not to say that there Oreortyx pictus 88 F will not be future introductions from these, or oth- Callipepla squamata 14 F er, taxa. Indeed, there have been recent sightings Callipepla culifornicu 412 S of various parrot species since 1990. For example, Callipepla gambelii 546 F Pyle (1994) reported that 10 to 15 Nanday Para- Callipepla douglasii 113 F keets (Nandayus nenday) were seen on the island griseogularis 20 F montezumae 8 F of Hawaii.‘ chukar 110 S In terms of success rates, we found that pas- Alectoris barbara 104 F serine species had a significantly higher overall francolinus 226 S success rate than either of the nonpasserine or- Fruncolinus pintadeanus 10 F ders. The reasons for this are unclear, but the Francolinus pondicerianus 214 S pattern is highly significant. It is possible to ex- Francolinus adsperus 4 F plain some of this result via the propagule size Francolinus icterorhynchus 9 F hypothesis. We found a significant relationship Francolinus clappertoni 10 F between propagule size (i.e., introduction effort) Francolinus erckelii 179 S and the success rates of galliforms introduced to Francolinus leucoscepus 27 F the island of Hawaii.‘ Caum (1933) also noted chine&s 8 F Bambusicola thoracicu 12 F that several columbiform species apparently Lophuru leucomelanos 67 S were introduced in very small numbers. How- Gallus sonnerutii 14 F ever, it remains to be shown that passerines were colchicus 244 S systematically released in larger numbers. reevesii 180 F The simplest potential predictor of the out- Pavo cristatus 2 S come of species ’ introductions is introduction Meleagris gallopavo 115 S history (Simberloff and Boecklen 1991). If in- Zenaida macroura 168 S troduction history alone were an adequate pre- Zenaida asiatica 40 F dictor of introduction outcomes we should have Streptopelia risoria (= decaol20 ?) 11 F detected clear all-or-none patterns within the or- Streptopelia chinensis 8 S ders we analyzed. Moulton (1993) and Moulton Geopelia striata 18 S and Sanderson (1997) argued that the all-or-none a See Appendix1 for common names patterns reported for passerines introduced to the Hawaiian Islands and elsewhere may be due to sampling artifact. When we extended the anal- were significantly different in a Kruskal-Wallis test ysis here to include the columbiforms and gal- (H = 5.25, P = 0.02). liforms, only the columbiforms show any evi- Data for the columbiforms appear to be equal- dence for such a pattern. Thus, we found little ly compelling, although we have not tested this evidence to support the notion that introduction group since there were just seven species intro- history is an adequate predictor of future intro- duced and two of these already were established duction outcomes. on Hawaii‘ at the time of the introductions by Our analyses suggested that one consistent the Puu‘ Waawa‘ a‘ Ranch (Lewin 1971). predictor of introduction success was size of na- tive geographic range. In all three orders we ob- DISCUSSION served that successfully introduced species had The introduction process in the Hawaiian Is- larger native ranges than unsuccessful species. lands has been highly nonrandom with respect to These results are consistent with the hypothesis phylogeny. Thus 10 of the 14 orders are repre- that species with larger ranges are ecologically sented by five or fewer species. The three orders more generalized (Brown 1984) and hence better that are represented by more species are those that able to adapt to a new environment. have been the focus of intentional introductions. In a partial test of the introduction effort hy- Thus, most galliforms were likely introduced to pothesis, we found that galliforms introduced suc- enhance prospects for recreational hunting, and cessfully to Hawaii‘ were introduced in larger most columbiforms were introduced for recrea- numbers than were unsuccessfulspecies. However, tional hunting or for aesthetic reasons. Passerines it should be noted that some species were suc- were introduced for a variety of reasons, including cessful with initial releases of as few as two in- biological control and aesthetic reasons, as well as dividuals; e.g., a single pair of (Pavo cris- accidental releases of cage birds. tutus) released on the Puu‘ Waawa’ a’ Ranch in The phenomenon of avian introductions, at least 1909 led to the successful establishment of the INTRODUCED BIRDS--Moulton, Miller, and Tillman 37 species on Hawaii‘ (Lewin 1971). Also, for 6 of ACKNOWLEDGMENTS the 15 unsuccessfulspecies, >85 individuals were We thank J. M. Scott, S. Conant, R. Walker, B. Den- released (Colinus virginianus, Callipepla dougla- nis, and an anonymous reviewer for their comments sii, Callipepla gambelii, Sytmaticus reevesii, Or- on earlier versions of the manuscript. Florida Agricul- eotyx pictus, Alectoris barbara; Table 4). We do tural Experiment Station JS #R-07766. A. van Doorn not know if successfulgame birds on islands other assisted with review of the literature of psittaciform than Hawaii‘ were introduced in higher numbers introductions. MPM wishes to thank C. J. Ralph, C. I? than were unsuccessful species. Because data are Ralph, and A. C. Ziegler for their kindness and hos- lacking for passeriform and columbiform species, pitality during fieldwork in Hawai‘i. a thorough test of the introduction effort hypoth esis was not possible.

APPENDIX 1. SCIENTIFIC AND COMMON NAMES OF 142 SPECIESINTRODUCED TO THE HAWAIIAN ISLANDS(NOMEN- CLATUREFOLLOWS SIBLEY AND MONROE 1990)

Scientific name Common name

Acridotheres tristis Common Myna Agapornis roseicapillis Rosy-faced Lovebird Alauda arvensis Skylark Alectoris bat-bat-a Barbary Alectoris chukar Chukar Amandava amandava Red Avadavat Amazona ochrocephala Yellow-crowned Parrot Amazona viridigenalis Red-crowned Parrot Ammoperdix griseogularis See-see Partridge Anus discors Blue-winged Teal Anas platyrhynchos Mallard Ara macao Scarlet Macaw Bambusicola thoracica Chinese -Partridge Brotegeris jugularis Orange-chinned Parakeet Bubulcus ibis Cattle Egret Cacatua galerita Sulphur-crested Cockatoo Cacatua moluccensis Salmon-crested Cockatoo Callipepla californica Callipepla douglasii Elegant Quail Callipepla gambelii Gambel’s Quail Callipepla squamata Scaled Quail nicobarica Nicobar Pigeon cardinalis Northern Carpodacus mexicanus Cettia diphone Japanese Bush-Warbler Chalcophaps indica Emerald Dove amherstiae Lady Amherst Pheasant Chrysolophus pictus Colinus virginianus Northern Bobwhite Collocalia vanikorensis Uniform Swiftlet livia Rock Pigeon Copsychus malabaricus White-rumped Shama Copsychus saularis Oriental Magpie-Robin Coturnix chinensis Blue-breasted Quail Coturnix japonica Japanese Quail Coturnix pectoralis Stubble Quail Crux rubra Great Currasow Cyanoptila cyanomelana Blue-and-White Flycatcher Cygnus olor Mute Swan Cyrtonyx montezumae Montezuma Quail Eclectus roratus Eclectus Parrot Eolophus roseicapilla Galah Erithacus akahige Japanese Robin Erithacus komadori Ryukyu Robin Estrilda astrild Common Waxbill Estrilda caerulescens Lavendar Waxbill Estrilda melpoda Orange-cheeked Waxbill 38 STUDIES IN AVIAN BIOLOGY NO. 22

APPENDIX 1. CONTINUED.

Scientific name Common name

Estrilda troglodytes Black-rumped Waxbill F&o (rusticolus ?) Gyrfalcon? Francolinus adsperus Red-billed Francolinus clappertoni Clappertons’ Francolin Francolinus erckelii Erckels’ Francolin Francolinus francolinus Francolinus icterorhynchus Heuglins’ Francolin Francolinus leucosepus Yellow-necked Spurfowl Francolinus pintadeanus Francolinus pondicerianus Gallicolumba luzonica Luzon Bleeding-Heart Gallus gallus Red Gallus sonneratii Gurrulax albogularis White-throated Laughingthrush Garrulax caerulatus Grey-sided Laughingthrush Garrulax canorus Hwamei Garrulax chinensis Black-throated Laughingthrush Garrulax pectoralis Greater Necklaced Laughingthrush Geopelia cuneata Diamond Dove Geopelia humeralis Bar-shouldered Dove Geopelia striata Zebra Dove Geophaps lophotes Crested Pigeon Geophaps plumifera Spinifex Pigeon Geophaps smithii Partridge Pigeon Geotrygon montana Ruddy Quail-Dove Gracula religiosa Hill Myna Grallina cyanoleuca Magpie-Lark Lagonosticta senegala Red-billed Firefinch Larus novaehollandiae Silver Gull Larus occidentalis Western Gull Leiothrix lutea Red-billed Leiothrix Leptotila verreauxi White-tipped Dove Leucosarcia melanoleuca Wonga Pigeon Lonchura cantans African Silverbill Lonchura malacca Black-headed Munia Lonchura oryzivora Java Sparrow Lonchuru punctulata Scaly-breasted Munia Lophura leucomelanos Lophura nycthemera Melanocorypha mongolica Mongolian Lark Meleagris gallopavo Wild Melopsittacus undulatus Budgerigar Mimus polyglottos Myiopsitta monachus Monk Parakeet Nandayus nenday Nanday Parakeet Neochen jubatu Orinoco Goose Nothoprocta perdicaria Chilean Tinamou Numida meleagris Helmeted Oreortyx pictus Ortalis cinereiceps Grey-headed Chachalaca capitata Yellow-billed Cardinal Paroaria coronata Red-crested Cardinal Paroaria dominicana Red-cowled Cardinal Parus varius Varied Tit Passer domesticus House Sparrow Passerina ciris Painted Bunting Passerina cyanea Indigo Bunting Passerina leclancherii Orange-breasted Bunting Pave cristatus Common Peafowl Penelope purpurascens Crested perdix Grey Partridge Phalacrocorax carbo Great Cormorant Phaps chalcoptera Common Bronzewing Phasianus colchicus Ring-necked Pheasant Phoenicopterus ruber Greater Flamingo INTRODUCED BIRDS-Moulton, Miller, and Tillman 39

APPENDIX 1. CONTINUED.

Scientific name common name

Platycercus adscitus Pale-headed Rosella Porphyria porphyrio Purple Swamphen Psittacula krameri Rose-ringed Parakeet Pterocles exustus Chestnut-bellied Sandgrouse Pycnonotus cafer Red-vented Bulbul Pycnonotus jocosus Red-whiskered Bulbul Rhipidura leucophrys Willie-Wagtail Rollulus rouloul Serinus leucopygius White-rumped Seedeater Serinus mozamhicus Yellow-fronted Canary Sicalis flaveola Saffron Finch Streptopelia chinensis Spotted Dove Streptopelia decaocto Eurasian Collared-Dove Sturnella loyca Long-tailed Meadowlark Sturnella neglecta Syrmaticus reevesii Reeves’ Pheasant Syrmaticus soemmerringii Copper Pheasant Tiaris olivacea Yellow-faced Grassquit Tut-nix varia Painted Tympanuchis cupido phasianellus Sharp-tailed Tyto alba Barn Owl Uraeginthus angolensis Blue-breasted Cordonbleu Uraeginthus bengalus Red-cheeked Cordonbleu Uraeginthus cyanocephala Blue-capped Cordonbleu Urocissa erythrorhyncha Red-billed Blue Magpie Vidua macroura Pin-tailed Wydah Zenaida asiatica White-winged Dove Zenaida macroura Mourning Dove Zosterops japonicus Japanese White-Eye

APPENDIX 2. LIST OF 3 1 SPECIES FROM 11 ORDERS NOT INCLUDED IN STATISTICAL ANALYSES. WITHIN EACH CELL, THE FIRST LINE INDICATES DAT!Z OF “RST INTRODUCTION (OR FIRST REFERENCE TO INTRODUCTION) AND STATUS (s = SUCCESSFUL; F = FAILED); THE SECOND LINE INDICATES MODE OF INTRODUCTION (1 = PRIVATE; 2 = STATE OR COUNTY AGENCY; 3 = UNKNOWN, INCLUDES ESCAPE FROM CAPTIVITY; 4 = POLYNESIANS; 5 = HUI MANU); AND THE THIRD LINE INDICATES REFERENCE

Species OahU‘ KaW.3 Maui Hawaii’ Molokai’ Lzna'i

Nothoprocta perdicaria 1966 F

Phalacrocorax carbo 1890s F 1

Phoenicopterus ruber 1929 F 1 1 Bubulcus ibis 1959 s 1959 s 1959 s 1959 s 1959 s 1959 s 1 1 1 1 1 1 7 7 7 I 7 7 Pterocles exustus 1961 F 1961 S 1961 F 2 2 2 5,ll 5,ll 5,ll Falco (rusticolus?)a 1929 F 40 STUDIES IN AVIAN BIOLOGY NO. 22

APPENDIX 2. CONTINUED.

Species Oahu’ Kauai’ Maui Hawai’ Molokai’ L%Ell‘

Turnix varia 1922 F 1922 F 2 2 1 1 Poyphyrio porphyrio 1933 F 1928 F 3 2 1 Larus novaehollandiae 1924 F 3 1 Larus occidentalis 1933 F 1933 F 3 3 1 Cygnus olor 1920 F 1

Neochen juhata 1922 F 2 1 Anas platyrhynchosb 1955 s

6 Anus disco& 1932 F 2 1 Tyto alba 1959 s 1959 s 1959 s 1958 S 1959 s 2 2 2 2 2 7 7 7 7 7,ll Collocalia vanikorensis 1962 S 2 7 Brotegeris jugularis 1933 F 3 1 Cacatua galerita 1933 F 3 1 Cacatua roseicapilla 1933 F 3 1 Cacatua moluccensis 1981 F 3 7 Ara macao 1933 F 3 1 Melopsittachus undulutus 1933 F 3 1 Psittacula krameri 1933 s 1981 S 1981 S 3 3 3 I,7 7,lO 7,lO Nandayus nenday 1981 F 1981 U 3 3 7 13 Myiopsitta monachus 1970 F 3 7 Amazona viridigenalis 1971 u 3 7 INTRODUCED BIRDS-Moulton, Miller, and Tillman 41

APPENDIX 2. CONTINUED.

Species Oahu’ Kauai‘ Malli Hawaii’ Molokai’ Amazonu ochrocephala 1969 F 3 7 Eclectus roratus 1981 F 3 7 Agapornis roseicapillis 1973 F 3 7 Platycercus adscitus 1877 F I 1 Urocissa erythrorhyncha 1966 F 1 7

Rqfwences: I = Caum 1933; 2 = Schwartz and Schwartz 1949; 3 = Munro 1960; 4 = Walker 1966; 5 = Walker 1967; 6 = Lewin 1971; 7 = Berger 1981: 8 = Moulton and Pimm 1983; 9 = Scott et al. 1986; 10 = Pratt et al. 1987; II = simherloffand Boecklen 1991; 12 = Moulton 1993; 13 = Pyle 1994; 14 = Wunz 1992. BCaum (1933) listed F rusricolus only a\ a tentative identification. ’ ’ May have interbred with natural migrants, as well as feral mdiwduals. ’ ’ SpCCiCs identity uncertain. Caum (1933) stated the spews IS Qurrquedulrr d~.scor.$(Blue-winged Teal, AU., dbcor,~); however, he also reponed that the individuals came from where the Blue-winged Teal does not occur.

APPENDIX 3. INTRODUCED PASSERINES ON SIX MAIN HAWAIIAN ISLANDS (SEE APPENDIX 2 FOR EXPLANATION OF TERMS)

Spec,es Oahu‘ Kauai‘ Maui Hawaii’ Molokai‘ LHnai‘

Acridotheres tristis 1872 S 1883 S 1883 S 1883 S 1883 s 1883 s 1 3 3 3 3 3 12 8 8 8 8 8 Alauda arvensis 1867 S 1870 F 1886 s 1902 S 1917 s 1917 s 3 1 3 3 3 3 12 1,8 8 8 8 8 Amandava amandava 1900 s 1987 S 1987 s 3 3 3 12 11 11 Cardinalis cardinalis 1929 S 1929 S 1949 s 1929 S 1951 s 1957 s 3s 1 3 2 3 3 I,8 1,8 8 1 8 8 Carpodacus mexicanus 1870 S 1886 s 1886 s 1886 S 1886 S 1886 s 3 3 3 3 3 3 12 8 8 8 8 8 Cettia diphone 1929 S 1988 S 1980 S 1979 s 1980 s 1,2 3 3 3 3 1 11 11 11 11 Copsychus mnlabaricus 1939 s 1931 s 5 1 11 I Copsychus saularis 1932 F 1922 S 5 1 1 1,12 Cyanoptila cyanomelanu 1929 F 1937 F 2,5 5 1,8 8 Erithacus akahige 1929 F 2 1 Erithacus komadori 1931 F 3 8 Estrilda astrild 1981 s 3 12 42 STUDIES IN AVIAN BIOLOGY NO. 22

APPENDIX 3. CONTINUED.

Species Oahu‘ Kauai’ Maui Hawai‘ Molokai‘ LaWa?

Estrilda caerulescens 1965 S 1978 s 3 3 12 11 Estrilda melpoda 1965 S 1989 s 3 3 12 MPM Estrilda troglodytes 1965 F 1975 s 3 3 12 11 Garrulax albogularis 1919 F 1 1 Garrulax caerulatus 1947 s 3 8 Garrulax canorus 1900 s 1918 s 1902 S 1909 s 1909 s 3 1 1 1 1 1,s 1,8 1,s 1,s 1.8 Garrulax chine&s 1931 F 1 1 Garrulax pectoralis 1962 S 3 11 Gracula religiosa 1960 S 3 11 Grallina cyanoleuca 1922 F 1922 F 2 2 1 1 Lagonosticta senegala 1965 F 3 11 Leiothrix lutea 1928 S 1918 s 1928 S 1928 S 1928 S 2 1 2 2 1 1 1 1 Lonchura cantans 1984 S 1984 s 1978 s 1972 S 1981 s 1979 s 3 3 3 3 3 3 11 11 11 11 11 11 Lonchura malacca 1936 S 1976 S 3 3 8 11 Lonchura oryzivora 1964 S 1983 s 1986 S 1981 S 3 3 3 3 12 11 11 11 Lonchura punctulata 1883 s 1883 s 1883 s 1883 s 1883 s 1883 S 3 3 3 3 3 3 8,12 8 8 8 8 8 Melanocotypha mongolica 1914 F 1 8 Mimus polyglottos 1931 s 1946 S 1933 s 1959 s 1951 s 1970 s 5 3 5 3 3 3 1,8 8 138 8 8 11 Paroaria capitata 1973 s 3 11 Paroaria coronata 1928 S 1928 S 1960 S 1976 S 1963 S 1976 S 1,5 3 3 3 3 3 1,ll 8,ll 11 11 11 11 INTRODUCED BIRDS-Moulton, Miller, and Tillmcm 43

APPENDIX 3. CONTINUED.

Species O‘ahu Kaua‘i Maui Hawai’i Moloka’l LBna‘i

Paroaria dominicana 1931 F 5 1 Parus varius 1928 F 1890 F 1928 F 1928 F 2 1 2 2 138 1,8 138 1,8 Passer domesticus 1871 S 1917 s 1917 s 1917 s 1917 s 1917 s 3 3 3 3 3 3 1,8 8 8 8 8 8 Passerina ciris 1937 F 5 8 Passerina cyanea 1934 F 1937 F 3 5 8 8 Passerina leclancherii 1941 F 1941 F 5 5 8 8,ll Pycnonotus cafer 1966 S 3 11 Pycnonotus jocosus 1965 S 3 11 Rhipidura leucophrys 1926 F 2 1,8 Serinus leucopygius 1965 F 3 11 Serinus mozambicus 1964 S 1977 s 3 1 11 11 Sicalis jlaveola 1965 S 1966 S 3 3 11 11 Sturnella loyca 1931 F 1 1 Sturnella neglecta 1931 F 1931 s 1934 F 2 1 3 8 1,ll 3 Tiaris olivacea 1974 s 3 11 Uraeginthus angolensis 1965 F 3 7 Uraeginthus bengalus 1965 F 1973 s 3 3 11 11 Uraeginthus cyanocephala 1969 F 3 12 Vidua macroura 1962 F 3 12 Zosterops japonicus 1929 S 1929 S 1938 S 1937 s 1938 S 1938 S 2,5 5 3 5 3 3 I,1 1 I,11 8 8 8 8 44 STUDIES IN AVIAN BIOLOGY NO. 22

APPENDIX 4. INTRODUCED GAME BIRDS ON THE SIX MAIN HAWAIIAN ISLANDS (SEE APPENDIX 2 FOR EXPLAINATION OF TERMS)

Species O'ahU Kaua‘i Mall1 Hawai'l Moloka‘i LZna‘i

Crm rubra 1928 F 2 1 Penelope purpurascens 1928 F 2 1 Ortalis cinereiceps 1928 F 2 1 Numida meleagris 1928 F 1874 F 1928 F 1928 F 1908 F 1914 F 1 1 1 1 1 1 1,lO 1,lO I,10 1,lO 1,lO 1,lO Colinus virginianus 1906 F 1906 F 1906 F 1906 F 1906 F 1906 F 2 2 2 1 2 2 1,4 1,4 1,4 4 1,4 1,4 Oreortyx pictus 1929 F 1929 F 2 2 1 1 Callipepla squamata 1961 F 2 6 Callipepla californica 18.55 F 1855 s 1855 s 1855 s 1855 s 1855 s 3 3 3 3 3 3 1,lO 1,lO 1,lO 1,lO 1,lO 1,lO Callipepla gambelii 1958 F 1958 F 1958 F 1958 S 1958 s 2 2 2 1,2 2 4,lO 4,lO 4,lO 6,lO 4,lO Callipepla douglasii 1959 F 1 6 Tympanuchus cupido 1895a F 1933a,b F 1 1 1 1 Tympanuchus phasianellus 1932 F 2 1 Cyrtonyx montezumae 1961 F 1 6 Ammoperdix griseogularis 1959 F 1 6 Alectoris chukar 1923 F 1957 s 1957 s 1949 s 1923 S 1923 S 2 2 2 2 3 3 1,lO 4,lO 4,lO 5,lO 4,lO 4,lO Alectoris barbara 1961 F 1959 F 1961 F 1959 F 2 1,2 2 2 4,lO 4,6 4,lO 4,lO Francolinus francolinus 1959 s 1959 s 1959 s 1959 s 2 2 1,2 2 9 9 6 9 Francolinus pintadeanus 1962 F 1 6 Francolinus pondicerianus 1958 S 1958 s 1958 s 1959 s 1958 s 1958 s 2 2 2 1 2 2 4,5,10 4,5,10 4,5,10 6 4,5,10 5 Francolinus adsperus 1965 F 1 6 Francolinus icterorhyn- 1961 F thus 1 6 INTRODUCED BIRDS-MouZton, Miller, and Tillman 45

APPENDIX 4. CONTINUED.

Species O‘ahu Kaua’i Maui Hawu’, Moloka‘i LBIXl‘i

Francolinus clappertoni 1961 F 1 6 Francolinus erckelii 1957 s 1957 s 1957 s 1958 s 1957 s 1957 s 2 2 2 1,2 2 2 5,lO 5,lO 5,lO 6 5,lO 5,lO Francolinus leucosepus 1959 F 2 6 Perdix perdix 1910 F 1926 F 1929 F 1 1 2 1 1 1 Coturnix chinensis 1922 F 1910 F 1922 F 1922 F 1922 F 2 1 2 2 2 1 1 1 1 1 Coturnix pectoralis 1922 F 1922 F 2 2 3 3 Coturnix japonica 1921 F 1921 S 1921 S 1921 S 1921 S 1921 S 3 3 2 3 3 2 2,lO 2,lO 1,lO 2,10 2,lO 1,lO Rollulus rouloul 1924 F 2 1 Bambusicola thoracica 1959 F 1961 F 2 1 4,5,10 6 Lophura leucomelanos 1962 S 1 6,lO Lophura nycthemera 1932 F 1870 F 2 1 1 1 Gallus gallus PHC S PH S PH F PH F PH F PH F 4 4 4 4 4 4 2,lO 2,lO 2,lO 2,lO 2,lO 2,lO Gallus sonnerati 1962 F 1 6 Phasianus colchicus 1865 S 1865 S 1865 S 1865 S 1865 S 1865 S 1 1 1 1 1 1 1,lO 1,lO I,10 1,lO 1,lO 1,lO Syrmaticus reevesii 1960 F 1960 F 1960 F 1959 F 1960 F 1960 F 2 2 2 1 2 2 4,lO 4,lO 4,lO 6 4,lO 4,lO Syrmaticus soemmerringii 1907 F 1907 F 1907 F 2 2 2 1 1 1 Chrysolophus pictus 1932 F 1870 F 2 1 1 1 Chrysolophus amherstiae 1932 F 2 1 Pave cristatus 1860 s 1860 F 1860 S 1928 S 1860 F 1860 F 1 1 1 1 1 1 1,lO 1,lO 1,lO 1,lO 1,lO 1,lO Meleagris gallopavo 1815 F 1815 F 1815 s 1815 s 1815 S 1815 S 1 1 1 1 1 1 1,lO 1,lO 1,lO 1,lO I,14 1,lO

a May have been Tympanuchus phasianellur (Caum 1931). h Based on “indefinite reports” (Caum 1913). ‘ Prehistoric introduction. 46 STUDIES IN AVIAN BIOLOGY NO. 22

APPENDIX 5. INTRODUCED COLUMBIDS ON SIX MAIN HAWAIIAN ISLANDS (SEE APPENDIX 2 FOR EXPLANATION OF TERMS)

Species O‘ahu Kaua'i Moloka‘i Hawai‘i Moloka‘l LHna'i

Caloenas nicobarica 1928 F 1922 F 2 2 1 1 Chalcophaps indica 1924 F 2 1 Columba livia 1796 S 1796 S 1796 S 1796 S 1796 S 1796 S 3 3 3 3 3 3 1 1 1 1 1 1 Gallicolumba luzonica 1929 F 1 1 Geopelia cuneata 1928 F 1929 F 2 2 1 1 Geopelia humeralis 1992 F 1922 F 1928 F 2 1 2 1 1 1 Geopelia striata 1922 S 1922 S 1922 S 1922 S 1922 S 1922 S 2 2 2 2 2 2 1 1 1 1 1 1 Geophaps lophotes 1922 F 1922 F 1922 F 1922 F 2 2 2 2 1 1 1 1 Geophaps plumifera 1922 F 1922 F 2 2 1 1 Geophaps smithii 1992 F 1922 F 2 2 1 1 Geotrygon montana 1933 F 3 3 Leptotila verreauni 1933 F 3 3 Leucosarcia melanoleuca 1922 F 1922 F 2 \, 2 \ 1 1 Phaps chalcoptera 1922 F 2 1 Streptopelia chinensis 1879 S 1890 s 1890 S 1890 S 1890 s 1890 S 3 3 3 3 3 3 1 8 8 8 8 8 Streptopelia decaocto 1928 F 1920 F 1928 F 1 1 2 1 1 1 Zenaida asiatica 1961 F 2 6 Zenaida macrowa 1962 S 1 9