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Home , Acytolepis, Acytolepis puspa, Badamia exclamationis, Catochrysops, Catochrysops panormus, Catopsilia

The ofPohnpei, Eastern , Micronesia1

Donald W Buden2 andJacqueline Y Miller3

Abstract: Fourteen of butterflies are recorded from , Micro­ nesia, seven for the first time. None is endemic to the island; all are widely dis­ tributed in the western Pacific, including parts of Indo-, with many extending into or beyond southeastern Asia. A long history of plant intro­ ductions and agricultural experimentation may have facilitated dispersal of but­ terflies to the island and provided a broad selection of host plants for those arriving otherwise unassisted. At least one, and possibly two or more, un­ identified species apparently confined to deep forest habitats were seen but not collected during this study. Compared with the local odonate fauna, the butter­ flies of Pohnpei differ in reaching their greatest abundance and species diversity in the lowlands, in lacking endemic species, and probably in having a higher turnover rate.

IN THE MOST RECENT and comprehensive number of species recorded from the island assessment of the distribution and status of and is based largely on collections and ob­ butterflies in , Schreiner and Na­ servations from throughout the island during fus (1997) recorded five species on Pohnpei: March 2000-September 2001. bochus, Danaus plexippus, Hypolimnas bolina, villida, and Melanitis leda. Ear­ lier reports on Pohnpei butterflies are scanty, Study Area sketchy, and scattered among several obscure and for the most part not readily available Pohnpei W 52' N, 158° 13' E) is the largest journals. A collection of butterflies assembled (355 km 2 [MacLean et al. 1986]) and highest from the field surveys of the of Mi­ island (approximately 800 m), and the capital, cronesia Project described in Gressitt's (1954) of the Federated States ofMicronesia (FSM), introduction was sent on loan to someone which also includes , Chuuk (formerly in Japan many years ago and apparently has Truk), and States (Figure 1). Warm, since been lost (S. E. Miller, pers. comm.); wet conditions persist year-round on Pohn­ there has never been a report published on pei. The average annual temperature is 27°C, butterflies in the Insects ofMicronesia publi­ and the monthly average does not vary from cation series. This lack of available informa­ that by more than 1°C (Laird 1982). The tion and paucity ofrecords provided incentive high rainfall-485 cm annually in the low­ for our further investigation into lands and up to 1015 cm estimated for the diversity on Pohnpei. Our study doubles the mountains (Merlin et al. 1992)-contributes to a lush tropical vegetation. Densely forested ridges and valleys radiate outward and down­ 1 Manuscript accepted 30 April 2002. ward from the central highlands. Mosses, 2 Corresponding author: Division of Natural Science ferns, and other epiphytes festoon the trees and Mathematics, College ofMicronesia-FSM, P.O. Box and shrubs of the cloud forest, usually oc­ 159, , Pohnpei FM 96941 (phone: 691-320-2480; fax: 691-320-2479; E-mail: [email protected]). curring above 600 m. A tropical rain forest 3 Allyn Museum of Entomology, Florida Museum of dominated by broadleaf trees reaching 30 or Natural History, 3621 Bay Shore Road, Sarasota, Florida more meters high covers much of the area 34234 (E-mail: jrniller®virtu.sar.usf.eclu). betWeen 200- ana 600 ffi, wtrh-rhe endemic kotop palm (Clinostip;ma ponapensis) forming Pacific Science (2003), vol. 57, no. 1:1-8 nearly pure stands on plateaus and gentle © 2003 by University of Hawai'i Press slopes, especially above about 450 m. The All rights reserved clearing of numerous small 1- to 2-ha plots 2 PACIFIC SCIENCE· January 2003

0 MARIANA 160 ISLANDS / __ MARSHALL ISLANDS

.i Yap 'I Pohnpei • '. I> • Chuuk .. Kosrae CAROLI NE ISLANDS NAURU,·..,...· KIRIBATI

1000 km AUSTRALIA

FIGURE 1. Location map of Pohnpei and surrounding western Pacific islands. for the cultivation of sakau (= , Piper islands, the largest and nearest of which (So­ methysticum) has been steadily encroaching kehs, Takaieu, and Temwen) are connected deeper and higher into this forest; Trustrum to mainland Pohnpei by causeways. (1996) reported a decline in intact native up­ land forest from 42% of the total vegetation MATERIALS AND METHODS cover in 1975 to 15% in 1995. The lowlands (below 200 m) include Specimens were collected opportunistically mainly secondary woodland and areas other­ and intermittently throughout the island wise modified for subsistence agriculture and by D.W.B. during March 2000-September human occupation; some intact rain forest 2001. The more remote montane forest re­ occurs at the upper extreme. Approximately gions were visited by backpacking for 2-4 8000 of the nearly 35,000 islanders live in days at a time. Localities were identified using Kolonia; the others are distributed in smaller a topographic map (U.S. Geological Survey settlements along the 87-km-Iong circum­ 1983), an altimeter (Barigo) scaled in 20-ft ferential road. Spur roads extend short dis­ (6.1-m) increments, and a global positioning tances into the interior but none traverses system (GPS) navigator (Garmin III Plus) the island;- the- central highlan-ds" are unin­ wnere the canopy- aio nof 15l"ocKsatellife re:' habited. form nearly a continuous ception. All specimens were deposited in the belt around the island, covering about 55 km2 collections of the Allyn Museum of Ento­ (Petteys et al. 1986). A barrier reef encloses a mology (Sarasota, Florida), American Mu­ lagoon with numerous volcanic and seum of Natural History (New York), Bishop Butterflies of Pohnpei . Buden and Miller 3

Museum (Honolulu), and CSIRO Long and another in a seepage area with Pocket Laboratory (Indooroopilly, Australia). alata, taro, and banana on Takaieu Island on Species recorded for the first time in 1 September 2001 are the only records for Pohnpei appear in boldface type in the spe­ Pohnpei. This species is more abundant than cies accounts. The name Pale pea-blue for the scanty records indicate. It occurs syntopi­ panormus follows Braby (2000), cally with C. pomona but is less common, with whereas all other English names follow an estimated ratio of about 1: 30. Schreiner and Nafus (1997). SPECIES ACCOUNTS puspa (Horsfield). Common hedge blue. Hesperiidae Matsumura (1915) described Cyaniris exclamationis (Fabricius). Brown (= Acytolepis) puspa watasei as a new "variety" . (subspecies) from Angaur (Palau) and Po~­ Previously recorded in Micronesia from pei, but he did not indicate island localities Palau, the Marianas, and the for individual specimens. He reported that (Schreiner and Nafus 1997). The nine speci­ the new form differed from the "type" in its mens collected in Kolonia and vicinity during smaller size, narrower marginal black bands 12-16 September (5), 22 April 2000 (2), and on the wings, and smaller patches (= spoj:s?) 2-15 September 2001 (2) are the first for on the undersurface of the wings. Schreiner Pohnpei. All were on or near and Nafus (1997) reported A. puspa from catappa. Caterpillars and pupae of B. exclama­ Palau but did not mention Pohnpei in their tionis occasionally were observed in the rolled account. or folded-over leaves of Terminalia, and the many damaged leaves with ragged edges pre­ (c. Felder). Pale pea­ sumably had been eaten by these larvae. blue. Schreiner and Nafus (1997) reported Ca­ tochrysops amasea from Palau, Yap, and Chuuk, pomona (Fabricius). Lemon IIll­ and no other congeners in Micronesia. grant. Semper (1906) recorded C. strabo from the Previously recorded in Micronesia only Caroline Islands [Micronesia] and made ·no from westernmost islands, in Palau and the mention of C. amasea or any other congeqers Marianas (Schreiner and Nafus 1997). The 21 in his account. Twenty-nine specimens of specimens collected during this study are the Catochrysops were collected on Pohnpei, in the first reported from Pohnpei. vicinity ofKolonia, during our study. Twenty is one of the most common species in the of them were critically examined by j.YM. lowlands, especially along roadsides in wet and identified as C. panormus based on the areas where the candle flower, Cassia alata, is wing and genitalic characters described in present. It was abundant also in the vicinity of Corbet and Pendlebury (1978) and Seki eF .al. a tree in bloom at the Agricul­ (1991), and three others from this series were tural Experiment Station in Kolonia in June identified by D. Sands also as C. panorfnus. 2000, where 20 to 30 were in view at anyone We include the remaining six also under C. time. panormus though we did not examine them critically and the identifications by colleagues (Linnaeus). Common to whom they were sent were uncertain. migrant.- CfitochrysopspaiiorinZis is Widely Clistrlouied Previously recorded in Micronesia in Palau from to and the Philippines and Yap (Schreiner and Nafus 1997). One southward through to northern collected on a roadside adjacent to a taro and eastern Australia, and eastward to the patch on Temwen Island on 30 April 2000, , , and the Loyalty 4 PACIFIC SCIENCE· January 2003

Islands (Tite 1959, Seki et al. 1991, Braby September 2000 are the only records for 2000). It has also been recorded in Micro­ Pohnpei. Larval host plants elsewhere include nesia on Guam (parsons 1999). The speci­ Hygrophila angustifolia and Ruellia tuberosa mens from Pohnpei are the first records for (Acanthaceae) (Braby 2000). the on the island and the first for the species in the Federated States ofMicronesia. Danaidae We have not examined examples of Gatochry­ Danaus plexippus (Linnaeus). Monarch. sops from elsewhere in Micronesia and can­ Widespread throughout Micronesia; oc­ not comment further on the identification of curring on all major islands or island groups those recorded as C. amasea and C. strabo. with the possible exception of Kosrae Larval host plants recorded for this species (Schreiner and Nafus 1997). This species was include many legumes (Fabaceae) among the apparently introduced to Pohnpei from Ha­ genera Phaseolus, Dolichos, Gajanus, Grotalaria, wai'i via a shipment of "milkweeds" in 1857 Dendrolobium, Flemingia, and Galactia (Seki (Gulick in Scudder 1875), but more likely as a et al. 1991, Parsons 1999, Braby 2000). Most stowaway adult than as eggs, larvae, or pupae of the Pohnpei specimens were collected on (Scudder 1875). Danaus plexippus is especially or near Dendrolobium sp. in coastal strand, common around settlements, usually in close and the others were in adjacent grassy, weedy proximity to crown flowers, Galatropis gigan­ areas. tea, a common ornamental.

Jamides bochus (Stoll). Cerulean. Euploea abjeeta Butler. Sicle-spotted brown Previously recorded in Micronesia in Palau crow. and Pohnpei (Schreiner and Nafus 1997). Schreiner and Nafus (1997) recorded this There is one specimen deposited at Bishop species in Micronesia only from Palau, Museum collected on Pohnpei by Z. Ono where they considered it common. Matsu­ without further data. Another specimen was mura (1915) reported it from Angaur (Palau) collected by the same collector on Palao Is­ and Pohnpei. He described coloration and land (= Palau) in the Caroline Islands on 4 pattern but gave no additional details regard­ May 1936. None was encountered on Pohn­ ing locality data for specimens collected. pei during our survey, although the host plants listed by Schreiner and Nafus (1997), including Grotolaria mucronata, Derris trifo­ liata, and Vigna marina, are common along Hypolimnas bolina (Linnaeus). Blue moon or the shore and on the lagoon islands. Common egg-fly. Recorded on all the major island groups in Zizina otis (Fabricius). Lesser grass blue. Micronesia, including Palau, the Marianas, Previously recorded in Micronesia from Yap, Chuuk, Pohnpei, Kosrae, and the Mar­ Yap, the Marianas, and Chuuk (Schreiner and shall Islands (Schreiner and Nafus 1997). It Nafus 1997). The only record for Pohnpei is is the most common species on Pohnpei, at a specimen collected in short grass and weeds least in the lowlands, especially in settled on the campus of the College of Mi­ areas and along roadsides. The variety of fe­ cronesia on 28 September 2000 and sent to male forms and different expanses of orange the Bishop Museum, where it was identified on the forewing are particularly noteworthy. to species by D. Preston. Hypolimnas pitboeka Kirsh. Dark egg-fly. Zizula bylax (Fabricius). Tiny grass blue. Previously recorded in Micronesia only on Previously "rec6iCleCl- in Micronesia froin Kosfae (Scnfemerana NafUs f9-97), wnichis Yap, the Marianas, and Chuuk (Schreiner and one of the easternmost islands and one of the Nafus 1997). Ten specimens collected in most distant from potential source popula­ short grass and weeds on the Palikir campus tions in the Indo-Australian-Melanesian re­ of the College of Micronesia during 20-28 gion. Of the six Kosraen specimens in the Butterflies of Pohnpei . Buden and Miller 5

Bishop Museum collection, four were col­ and August 1946 (5). In addition, Hirose lected at elevations between 800 and 1200 ft (1934) reported seven males collected on 23 (244-366 m) by H. K. Townes in August December 1933, and Matsumura (1915) con­ 1946. The two others are undated and lack sidered it common on Pohnpei and elsewhere additional locality data, but were collected by in Micronesia. However, none was collected Z. Ono, all ofwhose specimens from Pohnpei nor positively identified in the field during we examined were collected in 1936. our study. Occasional sightings (one for every Eighteen H. pithoeka collected by D.W.B. 4-5 days in the field) of orange and brown during 18 October 2000-18 August 2001 are butterflies in the well-shaded understory of the first reported for Pohnpei. This species is the rain forest may have included this species. widely distributed throughout the island but Two observed briefly but at close range much more numerous in montane forest than (within about 2 m) appeared to have a wing in the lowlands, which probably contributes pattern similar to that of J. villida, but an­ to the absence of previous records. One each other more closely resembled Vanessa indica, were collected at 73 and 92 m, eight at 200­ with its distinctive orange markings on the 400 m, three at 401-600 m, and five in cloud forewings; other sightings were too brief and forest or rain forest/cloud forest transition at too great a distance for even tentative at 601-800 m. Hypolimnas pithoeka does not identifications. None of these orange and usually occur under the forest canopy but brown butterflies was observed in the low­ has a predilection for open, sun-exposed lands in the more open, grassy, weedy areas sites, including mountain ridges and sum­ representing habitat seemingly more suitable mits, and forest edges bordering grasslands, for J. villida (and V indica). fern brakes, marshes, and agricultural plots (mainly sakau patches). It tends to be territo­ Satyridae rial, returning to the same perch even when Melanitis leda (Linnaeus). Evening brown. repeatedly disturbed during unsuccessful at­ Widely distributed throughout Micro­ tempts at capture, and it is usually seen paired nesia, being unrecorded only in the Mar­ or in small groups of three or four that tend shall Islands (Schreiner and Nafus 1997). On to stay within and circuit very circumscribed Pohnpei, it is common in dense grassy, weedy areas usually covering less than 0.5 ha. Some areas, from the lowlands to the highest peaks. fresWy emerged females may appear similar Specimens are markedly reduced in size with in coloration to males with a faint indica­ occasionally a faint indication of orange on tion of the bluish white postmedian band on the proximal margin of the dorsal forewing the forewing. Hopkins (1927) recorded the ocelli. larval host plant as Cudrania sp. near java­ nensis (Urticaceae), a common epiphyte in , but on H. pithoeka DISCUSSION subspp. utilize the herbaceous weed Sida rhombifolia (Malvaceae) (Parsons 1999). Of the 14 species of butterflies recorded from Pohnpei, none is endemic; all range widely in Junonia villida (Fabricius). Meadow argus. the West-central Pacific, occurring also in at Previously recorded in Micronesia from least parts of Indo-Australia and often ex­ Palau, Chuuk, Pohnpei, and the Marshall Is­ tending well into or beyond southeastern lands (Schreiner and Nafus 1997). It probably Asia. Adler and Dudley (1994) reported that was common on Pohnpei at least until the the Pacific butterfly fauna was derived largely mid-1900s, but is now extremely scarce if still from Australia, New Guinea, and Asia, and eitant: Tlle- Bishop -Ml.iseUIfi~ na:s 13 -speci­ mat isolatToIl from putential-suurce--areas--is mens from Pohnpei obtained by three differ­ the most important geographic variable ex­ ent collectors (Z. Ono, H. K. Townes, S. plaining variation in species richness among Uchiyama) at eight different localities during these islands. S. E. Miller (1996) cited nu­ April 1927 (2), February and March 1936 (6), merous examples of comprehensive biogeo- 6 PACIFIC SCIENCE· January 2003 graphic studies involving at least six different been brought to Pohnpei from 33 different orders of insects wherein a progressive dimi­ islands or countries, both tropical and tem­ nution of species occurred west to east across perate, mainly during the Japanese adminis­ the Pacific Basin. The number of species tration, 1914-1945. of butterflies recorded for Micronesia by The absence of butterflies endemic to Schreiner and Nafus (1997) demonstrates this Pohnpei or to any of the other remote Caro­ attenuation from west to east (away from pri­ line Islands fits a pattern described by Adler mary source areas), with at least 41 species on and Dudley (1994) for the Pacific islands as a Palau, 22 on Yap, 20 on Guam, 9 on Chuuk, whole, wherein the largest and nearest to po­ 5 on Pohnpei, 4 on Kosrae, and 4 in the tential source areas have the greatest number Marshalls. The seven additional species re­ of endemics. They (Adler and Dudley 1994) corded on Pohnpei during this study, to­ suggested that constraints of host plant spe­ gether with earlier records of cificity and the mechanics of -plant co­ and Euploea abjecta that were not included in evolution may impede diversification among Schreiner and Nafus (1997), increase the these more remote populations. It is of at Pohnpei list by nearly threefold to 14 and is least passing interest that compared with a perturbation in this otherwise smooth se­ butterflies, 8 (53%) of the 15 species of Odo­ quence; Pohnpei now has five more species nata (dragonflies and damselflies) recorded on than does Chuuk. Possibly Chuuk is incom­ Pohnpei are endemic to the island (Paulson pletely surveyed. Also, some species recorded and Buden in press). Odonates, like butter­ earlier in Pohnpei but not recently may no flies, are relatively large, visually conspicuous, longer be extant. On the other hand, Pohn­ diurnal fliers, but they have very different pei, being markedly larger than Chuuk (355 life history patterns, including larvae that are km 2 versus 117 km 2 divided among numerous aquatic carnivores as opposed to terrestrial islets), higher (approximately 800 m versus herbivores, and the absence of host plant 435 m), and with a more diverse flora (at least specificity. Compared with butterflies, the 438 native vascular plant species recorded odonate population of Pohnpei also appears versus 298 [Fosberg et al. 1979, 1982, 1987 more stable over time. All 14 species recorded in Falanruw, unpublished report submitted during the 1940s and 1950s (Lieftinck 1962), to FSM Advisory Panel for National Bio­ along with one undescribed presumed auc­ diversity Strategy and Action Plan]), might be tochthon (Paulson in press), but no other expected to support a larger butterfly fauna species, were collected by one of us (D.W.B.) despite its more remote location 710 km far­ during the course of this study. Six of them ther east. Furthermore, the dispersal of but­ represent a radiation within the damselfly terflies to Pohnpei may have been facilitated genus Teinobasis. On the other hand, the cur­ by the introduction of plants possibly bearing rent list of butterflies differs from earlier eggs or larval stages and serving also as hosts versions in both the apparent absence of pre­ for species that arrived otherwise unassisted. viously recorded species (Acytolepis puspa, The probable introduction of Danaus plex­ Jamides bochus, Euploea abjecta, and possibly ippus to Pohnpei via a shipment of milkweeds Junonia villida) as well as the addition ofthose (Asclepias sp.) from Hawai'i was reported by recorded for the first time. Scudder (1875). Ragone et al. (2001:290) The absence of Eurema hecabe from Pohn­ stated that "Pohnpei has been a center for pei lists is somewhat surprising in view of its plant introductions in Micronesia for more widespread occurrence elsewhere in the Pa­ than 100 years," alld that "the Pohnpei Agri­ cific (Schreiner and Nafus 1997) and the culture Station ... [was] once one of the availability of many of its leguminous host foremostcentersin~the-~worh:l~~for~ me- study p-Iant- species on The isla:nd.~Possioly it was of tropical agriculture...." They (Ragone present but undetected in the past. The et al. 2001: appendix 1) listed 395 species of Bishop Museum has one specimen recorded plants (along with additional hybrids, culti­ from Chuuk and another from Kosrae (thus vars, and varieties) documented as having bracketing Pohnpei), both collected by Z. Butterflies of Pohnpei . Buden and Miller 7

Ono in 1936. Schreiner and Nafus (1997) did in German and to Ms. K. Kodama for trans­ not encounter this species during more recent lations ofarticles inJapanese, and we thank S. visits to these islands and suggested that the E. Miller and M. S. Moulds for their helpful species is rare or that the two specimens (the comments on an early draft of the manu­ only records) may be mislabeled. script. The recent additions of Catopsilia pomona, C. and pyranthe, Catochrysops panormus, Mela­ Literature Cited nitis leda to the Pohnpei list probably represent very recent colonizations or re­ Adler, G. H., and R Dudley. 1994. Butterfly colonizations, or at least recent population biogeography and endemism on tropical expansions because these species are com­ Pacific islands. BioI. J. Linn. Soc. 51: 151­ mon, conspicuous, and not likely to have es­ 162. caped detection in their current numbers Braby, M. F. 2000. Butterflies of Australia: during earlier surveys. Badamia exclamationis, Their identification, biology and distribu­ Zizina otis, and Zizula hylax are less conspic­ tion. CSIRO Entomology, Canberra. uous and possibly were overlooked by previ­ Corbet, A. S., and H. M. Pendlebury. 1978. ous collectors. Whether Hypolimnas pithoeka is The butterflies of the Malay Peninsula. a recent arrival or a long-term resident that 3rd ed. (J. N. Eliot, rev.). Malayan Nature has avoided detection in part because of its Society, Kuala Lumpur. scarcity in the lowlands is less certain. Of Fosberg, F. R, M.-H. Sachet, and R the 10 species of butterflies identified in the Oliver. 1979. Geographical checklist of the field during this study, only H. pithoeka was Micronesian Dicotyledonae. Micronesica more abundant in the upland forest than in 15:41-295. the lowlands; Melanitis leda was common in ---. 1982. Geographical checklist of the grassy areas throughout. The eight others Micronesian Pteridophyta and Gymno­ were encountered only in the lowlands, spermae. Micronesica 18:23-82. chiefly settled areas, roadsides, and coastal ---. 1987. Geographical checklist of strand. the Micronesian Monocotyledonae. Mi­ There remains the issue of the identity cronesica 20:19-129. of probably two different species of small to Gressitt, J. L. 1954. Insects ofMicronesia, in­ medium-sized, extremely wary orange and troduction. Insects ofMicronesia 1:1-257. brown butterflies that are seldom encoun­ Hirose, T. 1934. Butterflies from the Mar­ tered and only in the forest understory. shall and Caroline islands. Trans. Kansai Whether they are species previously reported Entomol. Soc. 5:29-31. from the island (at least two sighted resem­ Hopkins, G. H. E. 192 7. Butterflies of Samoa bled ]unonia villida) now occurring atypically and some neighboring island-groups. In­ in deep forest, or whether they represent sects of Samoa and other Samoan terres­ range extensions for a species as yet unre­ trial Arthropoda. Part III. corded on Pohnpei, or possibly examples of 1:1-64. as yet undescribed species, or some combina­ Laird, W. E. 1982. Soil survey of island of tion thereof is unknown and high priority for Pohnpei, Federated States of Micronesia. further investigation. U.S. Department of Agriculture, Soil Conservation Service. U.S. Government Printing Office, Washington, D.C. ACKNOWLEDGMENTS Lieftinck, M. A. 1962. Insects of Micronesia, For their assistance in identifying specimens Odonata. Insects ofMicronesia 5:1-95. ana pioviaing material for sfiidy,we iliiiIlk MaCLean, c.n.,-l.G. Cole;C. D. Wrute­ P. Ackery (BMNH), L. Miller (AME/FLMNH), D. sell, M. C. Falanruw, and A. H. Ambacher. Preston, F. Rindge (AMNH), Scott E. Miller 1986. Vegetation surveys of Pohnpei, (NMNH), and D. Sands. We are also grateful Federated States of Micronesia. U. S. For. to Rev. E. Kalau for translations of passages Servo Resour. Bull. PSW-18. 8 PACIFIC SCIENCE· January 2003

Matsumura, S. 1915. Micronesian butterflies. Pohnpei, Micronesia and the role of the Entomol. Mag. Kyoto 1:63-68. Pohnpei Agriculture Station. Econ. Bot. Merlin, M., D. ]ano, W. Raynor, T. Keene, 55:290-324. J. ]uvik, and B. Sebastion. 1992. Tuke en Schreiner, I. H., and D. M. Nafus. 1997. Pohnpei [Plants ofPohnpei]. Environment Butterflies of Micronesia. Agricultural Ex­ and Policy Institute of the East-West periment Station, College of Agriculture Center, Honolulu. and Life Sciences, University of Guam, Miller, S. E. 1996. Biogeography of Pacific Mangilao. insects and other terrestrial invertebrates: Scudder, S. H. 1875. The introduction of A status report. Pages 463-475 in A. Keast Danaida plexippus into the Pacific islands. and S. E. Miller, eds. The origin and Psyche (Camb.) 1:81-84. evolution of Pacific island biotas, New Seki, Y., Y. Takanami, and K. Otsuka. 1991. Guinea to eastern : Patterns and Butterflies of . Vol. 2, No. 1. Ly­ processes. SPB Academic Publishing, Am­ caenidae (K. Otsuka, ed). Tobishima Cor­ sterdam. poration, Tokyo. Parsons, M. 1999. The butterflies of Papua Semper, G. 1906. Beitrag zur lepidopteren­ New Guinea: Their systematics and biol­ fauna des Karolinen-Archipels. Deutche ogy. Academic Press, San Diego. Entomol. Z. Iris 18:245-267. Paulson, D. R in press. Teinobasis budeni Tite, G. E. 1959. The genus Catochrysops sp. nov. from Pohnpei, Eastern Caroline Lepidoptera: Lycaenidae. Entomologist Islands, Micronesia (Odonata: Coena­ (Lond.) 92:201-212. grionidae). Int. J. Odonatol. Trustrum, N. A. 1996. Pohnpei's watershed Paulson, D. R, and D. W. Buden. in press. spatial plan and management guidelines. The Odonata of Pohnpei, Eastern Caro­ Landcare Research, , Ltd. line Islands, Micronesia. Int. ]. Odonatol. Palmerston North, New Zealand. Avail­ Petteys, E., S. Peter, R Rugg, and T. Cole. able at The Nature Conservancy, Kolonia, 1986. Timber volumes in the Pohnpei. forests of Pohnpei, Federated States of U.S. Geological Survey. 1983. Topographic Micronesia. U.S. For. Servo Resour. Bull. map of the island of Ponape [2 sheets]. PSW-19. Scale 1: 25,000. U.S. Geological Survey, Ragone, D., D. H. Lorence, and T. Flynn. Reston, Virginia. 2001. History of plant introductions to