The Odonata ofKosrae, Eastern Caroline Islands, Micronesia!

Donald W Buden 2,4 and Dennis R. Paulson 3

Abstract: A recent collection of 69 specimens together with survey counts and incidental observations during June-July 2002 provide new information on the odonate fauna of Kosrae, Micronesia. The fauna comprises one zygopteran (Isehnura aurora) and six anisopterans. It appears to have remained stable with no known extinctions or colonizations over the past half century. The fauna is nearly a subset of that of Pohnpei and the islands to the west, and it comprises six widespread weedy species and one endemic, Hemieordulia erieo. Upland aquatic habitats appear largely unexploited or underutilized by odonates, and the absence of any Teinobasis species on Kosrae is in marked contrast to the presence of six species on the nearest high island, Pohnpei.

THE ODONATE FAUNA of the Federated The specimens he examined were obtained States of Micronesia (FSM) has been largely by three collectors (T. Esaki, December unstudied since Lieftinck's (1962) compre­ 1937; R. P. Owen, July 1949; J. F. Gates hensive review of populations in greater Mi­ Clarke, January-April 1953) and from eight cronesia, including Palau, the Marianas, and localities, at least seven of which are in the the Marshall Islands. In the only other studies lowlands (highest recorded elevation 60 m), of Odonata in the FSM, Schreiner and Nafus with the elevation of Clarke's "Mt. Matante" (1989) commented briefly on species they locality unstated. There are no indications encountered during surveys of aquatic insects that any of these specimens or any records on Yap in September 1988, and Paulson that Lieftinck (1962) gleaned from the litera­ (2003) and Paulson and Buden (2003) re­ ture are from the uplands, which appear to viewed the status of populations on Pohnpei have been largely or entirely overlooked dur­ based largely on collections and observations ing previous odonate surveys. Clarke, how­ during 2001-2002. The study reported here ever, collected insects other than odonates is an extension of these more recent inves­ at numerous inland stations, including the tigations and focuses on Kosrae, the eastern­ highest summits and ridges (Clarke 1953, most island in the FSM. 1976, Gressitt 1954). Clarke (1953) reported nine (unidentified) species of odonates among Summary ofOdonate Studies on Kosrae 704 insect species in his preliminary assess­ ment of specimens collected on Kosrae for Lieftinck (1962) recorded seven species of the Insects ofMicronesia project. Whether all odonates on Kosrae, including one damselfly of the specimens of two species of odonates (Zygoptera) and six dragonflies (Anisoptera). were lost or otherwise unavailable to Lief­ tinck (1962), who recorded only seven, or I Manuscript accepted 19 February 2003. whether Clarke's original tally was in error is 2 Division of Science and Mathematics, College of unknown. Micronesia-FSM, P.O. Box 159, Kolonia, Pohnpei FM The Owen collection appears to consist of 96941 (E-mail: [email protected]). 3 Slater Museum of Natural History, University of only a few specimens from Lelu Island, and Puget Sound, Tacoma, Washington 98416 (E-mail: Esaki's specimens are from several widely [email protected]). distributed coastal or lowland localities. Most 4 Corresponding author. of the specimens collected by Esaki were deposited in the Kyusyu Imperial University Pacific Science (2003), vol. 57, no. 4:399-407 (Japan) and unavailable to Lieftinck (1962), © 2003 by University of Hawai'i Press but locality records for the Anisoptera were All rights reserved included in Asahina's (1940) study.

399 400 PACIFIC SCIENCE· October 2003

The study reported here provides new Mt. Finkol (= Mt. Crozer) near the center of information on the distribution, habitat pref­ the island, and 589 m on Mt. Mutunte (= Mt. erences, and relative abundance of odonates Bauche), whose slopes comprise much of the on Kosrae and is based largely on specimens north end. collected and observations recorded by The average annual rainfall is about 5000­ D.W.B. from throughout the island during 6000 mm/yr (Merlin et al. 1993), with esti­ 20 June-6 July 2002. mates for the mountains ranging to as high as 7500 mm (Whitesell et al. 1986). The average annual temperature is 27°C at sea level and Study Area slightly lower at higher elevations, and the Kosrae (5° 19' N, 163° E) is a densely for­ difference between the average monthly min­ ested and ruggedly mountainous, 109-km2 imum and average monthly maximum is less volcanic island (about 1.2-2.6 million yr old than 8°C throughout the year (Whitesell et al. [Keating et al. 1984]) in the eastern Caroline 1986). The abundant and evenly distributed Islands (Figure 1). It is the easternmost state rainfall feeds at least 22 perennial streams in the Federated States of Micronesia, which (T.T.P.I. Office of Planning and Statistics includes also Yap, Chuuk (formerly Truk), 1979) and, together with warm year-round and Pohnpei states. Mountains compose temperatures, contributes to the growth of about 70% of the island area (Whitesell et al. lush, tropical vegetation. 1986), with maximum elevations of 630 m on Cloud forest consisting of a luxuriant as-

1400 MARIANA MARSHALL ISLANDS ,: , ISLANDS . . , Yap 'Chuuk Pohnpej"': '.. ,I • l=. ' o , Palau • ..'. CAROLINE ISLANDS '\ .' Kosrae

- " I 1000 KM

AUSTRALIA

FIGURE 1. Location map of Kosrae and surrow1ding western Pacific islands. Odonata of Kosrae . Buden and Paulson 401 semblage of terrestrial and epiphytic bryo­ Other major lowland habitats assessed by phytes, ferns, and orchids and gnarled, Whitesell et al. (1986) include secondary stunted trees (5-6 m tall) and shrubs occurs vegetation (1272 ha = 11.4% of total area) on the highest peaks and ridges above about and mangrove forests (1562 ha = 14%). 480 m (Hosokowa 1952). It "represents the Freshwater swamps, with Terminalia caro­ lowest elevation in the world at which TMCF linensis as the dominant tree, occur in depres­ [tropical montane cloud forest] is known to sions or poorly drained soils, usually behind occur" (Merlin and Juvik 1993:153). Com­ beach strand. Terminalia swamps account for mon trees and shrubs in this zone include only 345 ha (3 %) of the land area, but they Astronidium kusaianum, Campnosperma brevi­ are considered "the most well developed, sci­ petiolata, Cinnamomum carolinense, Elaeocarpus entifically acclaimed swamp forests in all of carolinensis, Polyscias subcapitata, and Psychotria Micronesia today" (Merlin et al. 1993:40). sp. Tree ferns (Cyathea) are common and Other aquatic habitats widely distributed Dicranopteris ferns predominate in scattered throughout Kosrae include numerous streams open areas. Many of the tree species occur and seepage areas in the uplands, and streams, also at lower elevations where they grow to taro patches, roadside puddles, drainage greater size. Summit areas are usually open ditches, and open cisterns in the lowlands. with scattered shrubs and small trees and a Initial colonization of Kosrae by aborigi­ dense cover of ferns and grasses. The total nals, probably from Melanesia, occurred area of cloud forest is about 69 ha and com­ about 2000 B.P. (Athens 1995), and first con­ prises 0.6% of the total area of the island. tact with Europeans was in 1824 during a Rain forest, with the largest trees 25-30 m lO-day visit by the French research vessella tall, covers most ofthe slopes below the cloud Coquille (Ritter and Ritter 1982). The most forest. Campnosperma brevipetiolata, Elaeo­ recent (1994) census recorded 7317 people carpus carolinense, Horsfieldia nunu, Parinarium residing mainly in several coastal settlements laurina, and Ficus spp. are among the domi­ distributed around the entire island (Kosrae nant species; Ptychosperma ledermanniana is Branch Statistics Office 2000); the mountain­ the only indigenous palm. Tree ferns and ous interior is uninhabited. A perimeter road other ferns are common throughout. The connects nearly all the settlements; the Utwa upland forest (excluding cloud forest) covered to Walung section is still unpaved and in 5090 ha (45.5% of total area) in 1983 large part unimproved, and the Walung to (Whitesell et al. 1986), but this figure has Okat segment is still in the proposed and likely diminished in recent years following planning stages. anthropogenic changes, such as agricultural expansion and encroachment into higher ele­ MATERIALS AND METHODS vations and clearing land to otherwise ac­ commodate a growing human population All odonates encountered above 200 m dur­ (Raynor 1993, Allen et al. 2001). ing this survey were collected whenever Agroforest is widespread in the lowlands possible, though difficult terrain kept success and lower mountain slopes, where native rate low. Collections at lower elevations vegetation is partially cleared for cultivation were more selective, often bypassing the of subsistence and cash crops, including ba­ more common species to search for those less nanas (Musa spp.), breadfruit (Artocarpus alti­ common. Terms ofabundance used in species lis), betel palm (Areca catechu), coconut (Cocos accounts are appraisals based on individuals nucifera), oranges and tangerines (Citrus spp.), observed during timed counts as well as im­ papaya (Carica papaya), tapioca (Manihot escu­ pressions based on incidental observations lenta), taro (Alocasia, Colocasia, Cyrtosperma), throughout the study period: very common and yams (Dioscorea spp.). Whitesell et al. (50 or more encounters per day on most days (1986) recorded 2585 ha ofagroforest in 1983 under suitable viewing conditions and in (23.1 % of total area), compared with 1844 ha suitable habitat), common (25-50 encounters (16.5%) in 1975. per day on most days), fairly common (10-25 402 PACIFIC SCIENCE· October 2003 encounters on most days), and uncommon mann 1997). The only previous records ofA. (up to about 10 encounters per day, but un­ guttatus on Kosrae are a male collected by recorded on many days). Clarke at Mutunlik in February 1953, and Six timed survey counts were conducted one or more specimens collected by Owen on in the lowlands (below 200 m) along paved Lelu Island in July 1949 (Lieftinck 1962). and unpaved roads and recently cleared areas, During the June/July 2002 survey, A. guttatus mainly along the completed and partially was uncommon to locally fairly common in completed parts of the circumferential road, open, sparsely vegetated areas; it was most and through a wide variety of habitats. Sur­ numerous along roads, over parking lots and veys totaled 14.3 hr at different times of the fields, and in recently cleared areas. It was day and covered all areas of the island perim­ often found near standing water; some were eter with the exception of the northern and observed patrolling over freshwater swamps. northeastern sections. Counts are presented as encounters per hour and give a crude Family CORDULIIDAE approximation of the relative abundance of species in edge habitats throughout the low­ Hemicordulia erico Asahina lands. The genus Hemicordulia is widespread in All specimens are currently in the collec­ Oceania, but it is represented on many islands tion of D.R.P. but are to be transferred to the and small island groups by endemic species. International Odonata Research Institute, Asahina (1940) described H. erico endemic to Gainesville, Florida. Kosrae from a series of 11 males and one fe­ male collected by Esaki in 1937. He (Asahina SPECIES ACCOUNTS 1940) recorded all 12 in "materials examined" as collected in November, but where he listed Family COENAGRIONIDAE them as holotype, allotype, and 10 paratypes, he reported the collection date of the holo­ Ischnura aurora (Brauer) type (a male from Malem) as 22 December. This species occurs widely from India and Asahina's "materials examined" section in­ Sri Lanka to Polynesia and New Zealand cludes a male from Malem collected on 22 (Steinmann 1997), and Lieftinck (1962) listed November, but December is probably the numerous localities from throughout Micro­ correct month for this entire series, because nesia. Lieftinck (1962) recorded an unstated all the other of Esaki's Kosraean records number of1. aurora collected by Esaki, Owen, annotated as to date by Asahina (1940), and and Clarke on Kosrae in the lowlands (Lelu involving three other species, range from 1 Island, Malem, Mwot-Utwe, and Mutunlik) to 18 December. Furthermore, other locality during December-February, April, and July. records throughout Asahina's text indicate This species was encountered only occasion­ that Esaki was elsewhere in Micronesia on ally during the June/July 2002 survey, and some of the November dates in question. mainly in wet, lowland marshy areas, includ­ Consequently, the collection date of the par­ ing grassy, weedy drainage ditches and taro atype from Mwot-Utwe that Lieftinck (1962) patches. It was locally very common in some 2 reported as 8 November 1939 should be areas (3-4 m ), but scarce in similar habitats emended to 8 December 1937. elsewhere. It was found mainly at or near sea In addition to the paratype, Lieftinck level but occasionally along stream banks to (1962) examined six males and one female elevations as high as 182 m. collected by Clarke during February (5 specimens), March (1), and April (1) 1953 and Family AEscHNIDAE another male collected by Owen in July 1949. Anax guttatus (Burmeister) Clarke's material included a pair in copula at This species is widespread in the Pacific Funaunpes in February. With the possible islands through Indo-Australia to Southeast exception of Clarke's specimen from an un­ Asia, India, and the Seychelle Islands (Stein- specified elevation on Mt. Mutunte, all of Odonata of Kosrae . Buden and Paulson 403 these early records are from the lowlands. breeding habitat. The closely related H. ha­ The locality for Owen's specimen was un­ luco Asahina was similarly common at all ele­ stated, but all of his other records are from vations on Pohnpei, but its breeding habitat Lelu Island, so his specimen of H. erico prob­ was not determined certainly (Paulson and ably is from there as well. Buden 2003). During the June/July 2002 survey, H. erico Asahina (1940) was unable to describe the was encountered throughout the island from female cerci in this species. They are cylin­ sea level to the summits of the highest peaks. drical, very slightly upcurved, thickest at It was most numerous along the circum­ midlength, bluntly pointed, and 2.5 mm in ferential road passing through secondary length. Those of its congener from Pohnpei, woodland, agroforest, and swampland in the H. haluco Asahina, are similar but straight and northwestern quadrant, between the Okat just over 3 mm in length. River bridge and the causeway to the airport With the almost simultaneous collection (33 total, 19 per hour, 29 June), but it was of large series of H. erico and H. haluco (Paul­ scarce to uncommon in similar habitats along son and Buden 2003), a comparison between the road on the eastern side of the island, these two species is warranted. Hemicordulia where only 5-6 were seen throughout the erico is slightly smaller than H. haluco, with a study period. It was the most frequently en­ distinctly shorter abdomen. Mean hind wing countered odonate in the higher elevations length of males is 26.6 mm (24.5-29, n = 28) (Table 1), especially on the fern-, shrub-, and in H. erico versus 28.2 mm (27-29.5, n = 37) grass-covered summits. Four or five were ob­ in H. haluco, and mean hind wing length served together patrolling an area of several of females is 29.1 mm (28-29.5, n = 7) in square meters on the slope of Mt. Oma, just H. erico versus 29.9 (29.5-31.5, n = 11) in below the summit at about 440 m elevation H. haluco. Mean abdomen length of males is on 28 June. 28.6 mm (25-31, n = 26) in H. erico versus One female with eggs extruding from the 32.6 mm (30-35, n = 18) in H. haluco, and abdomen collected in Walung Village on the mean abdomen length offemales is 29.1 mm southwestern coast on 3 July was the only di­ (28-29.5, n = 7) in H. erico versus 36.4 mm rect evidence of breeding. However, males (36-37, n = 6) in H. haluco. were seen patrolling over both the Malem Asahina (1940:8) said of this species, "col­ and Finkol Rivers, which may have been our very dull greenish (more shining in one

TABLE 1 Distribution of Odonates Collected on Kosrae during 20 June-6 July 2002, with Encounter Rates" during Lowland Surveys

Specimens Collected"

Species Below 200 m Above 200 m Sightings/hr

Zygoptera Ischnura aurora 3M, IF 1.0 Anisoptera Anax guttatus 1M 1.0 He11licordulia e"ico 14M, IF 14M,6F 4.0 Diplacodes bipunctata SM, IF 2.6 Pamala fiavescens 6M,2F 3.8 Tholy11lis tillarga 7M,2F IF 7.0 Tramea translnm'ina 3M, IF IF 0.8

" Calculated as the average number ofindividuals encountered on six timed roadside counts totaling 14.3 hr at different times of the day from 0720 to 1520 hours during 20 June-6 July 2002: eastern section (between Tofol and Malem Village, 6.0 hr), southwestern section (between Utwa and Walung Villages, 6.5 hr), northwestern section (between Okat River bridge and airport causeway, 1.8 hr). /, M, males; F, females. 404 PACIFIC SCIENCE· October 2003 specimen)." In fact, H. erico is not at all dull low, open, sparsely vegetated areas during the but has a glittering metallic thorax much like June/July 2002 survey. It was most numerous that of H. haluco, but it is an overall darker in the vicinity of rain-filled ruts on dirt and species. The thorax in mature male H. erico is gravel roads and road construction sites. One entirely metallic green on top and sides; that pair was collected in tandem ovipositing in a of male H. haluco is the same color but with roadside puddle at Saulung at the southern the nonmetallic orange brown of the ventral end of the island on 1July 2002. surface penetrating diffusely up the sides. Fe­ male H. erico are colored as males, but female Pantala flavescens (Fabricius) H. haluco are even less brightly colored than This nearly cosmopolitan species ranges the males, with the metallic green on the throughout the Tropics and subtropics thorax restricted to the upper parts of the worldwide (Steinmann 1997). Lieftinck mesepisternum and isolated stripes the length (1962:85) indicated that Clarke encountered of the mesepimeron and metepimeron. Addi­ P. flavescens at Mutunlik, Kosrae, in February tional differences are seen in the hind femora 1953 but did not list specimens and did not (black in H. erico, mostly orange brown in H. annotate records for any Micronesian islands, haluco) and abdominal segment 10 and its stating only that there were "Many Micro­ cerci (black in H. erico, brown in H. haluco). nesian specimens, collected on almost all is­ In both species, the wings of immature land groups; data not all recorded." This males are completely hyaline, and those of species was uncommon to locally common immature females have an orange wash over throughout the lowlands on Kosrae during the base of both wings not quite reaching the the June/July 2002 survey. It was most nu­ subtriangle in the forewings and the triangle merous over roadways and along roadsides. in the hind wings. In H. erico, the wings of One pair oviposited in tandem on the rain­ most mature males become washed with wet cement surface of a basketball court, and dusky brown, but in H. haluco relatively few another was in copula over a roadside puddle; males are so colored (difficult to quantify). both pairs were collected in Utwa Village on In mature females, the wings are similarly 1 July 2002, and both of the females have almost entirely dusky in H. erico, but in H. eggs extruding from the abdomen. haluco the dusky washes are restricted to the postnodal areas of all wings and another dif­ Tholymis tillarga (Fabricius) fuse area from the distal end of the anal loop This species ranges from Africa eastward back to the wing margin; the base remains across southern Asia and to Australia and orange in all females. Oceania (Lieftinck 1962). The only previous records from Kosrae are the eight males and four females collected by Esaki during 3-18 Family LIBELLULIDAE December 1937 (Asahina 1940). Tholymis till­ Diplacodes bipunctata (Brauer) arga was the most common odonate encoun­ This species ranges widely in Australasia tered on Kosrae during the June/July 2002 and East Asia (Steinmann 1997), and Lief­ survey. It was most numerous in coastal low­ tinck (1962) listed numerous locality records lands and was observed mainly at dusk. The from throughout Micronesia. Asahina (1940) greatest concentration was in the Tenwak/ recorded five males and three females col­ Mutunlik area along the shoreline and adja­ lected by Esaki on Kosrae during 1-14 De­ cent road bank, where 300-400 were esti­ cember 1937. At least one other specimen mated in an area covering about 100 by 20 m. recorded by Lieftinck (1962) as collected at Most were flying close to the ground or just Mutunlik in February 1952 probably is a part above the surface of the water along the of Clarke's 1953 collection and should be in­ shoreline. Others were in small groups flying cluded with his January and February (1953) in tight circles near the tops of scattered records; the number of specimens was left roadside trees. Tholymis tillarga was the only unstated. This species was fairly common in odonate regularly encountered along rain- Odonata of Kosrae . Buden and Paulson 405 forest trails, where individuals would dislodge would provide seemingly suitable resources from understory vegetation when disturbed for one or more Teinobasis species or an and fly higher into the canopy. It occurred ecological equivalent. Although dragonflies from sea level to the mountain summits, but and damselflies are occasionally encountered was less numerous at the higher elevations along Kosrae forest streams, this habitat ap­ where water was less common. pears largely unexploited or underutilized by odonates. Small island size and distance from Tramea transmarina Brauer potential sources almost certainly contribute This species occurs widely in the Indo­ to the depauperate fauna on Kosrae, as ex­ Australian region and the islands of Oce­ pected according to the MacArthur and Wil­ ania (Lieftinck 1962, Steinmann 1997). son (1967) model of island biogeography. Asahina (1940) recorded three T. limbata Kosrae is about one-third the size ofPohnpei, (=transmarina) collected by Esaki at Mwot­ and, excluding a scattering of low-lying atolls, Utwe on 8 December 1937, and Lieftinck it is the easternmost island in Micronesia and (1962) recorded five collected by Clarke at the most distant from potential source pop­ Mutunlik during February-March 1953 and ulations on the larger Indo-Australian land another by Owen at Lelu Island in July 1949; masses. Area/distance effects notwithstanding, all nine were males. Lieftinck (1962) included the ecological breadth ofKosrae suggests that Kosrae and other eastern Micronesian islands it could support more than the seven species within the range of the subspecies T. t. pro­ of odonates it now has (Paulson and Buden pinqua Lieftinck. During the survey reported 2003). The Kosrae odonate fauna appears to here, this species was scarce or uncommon have remained stable with no known extinc­ (uplands) to occasionally fairly common tions or colonizations during the past half (lowlands), mainly over roads and other open century; all the species recorded by Lieftinck areas. Two or three were observed on the (1962), but no others, were observed during open summits ofMt. Mutunte on 5-6 July. this study. With the exception of the Teinobasis spe­ cies complex, the odonate faunas of Pohnpei DISCUSSION and Kosrae are very similar in species com­ Of the seven species of Odonata known from position. Both islands have Isehnura aurora, Kosrae, Isehnura aurora is the only zygopteran Anax guttatus, Diplaeodes bipunetata, Tholymis and Hemieordulia erieo the only endemic spe­ tillarga, Pantala flaveseens, and Tramea trans­ cies. By contrast, the adjacent island Pohnpei marina, all of which are widespread, weedy hosts nearly twice as many species (15), of species common among the islands of the which nearly half (7) are endemic, and 1. au­ western Pacific and, in many cases, well be­ rora is only one of seven Zygoptera (Paulson yond. Except for Hemieordulia erieo, which and Buden 2003). The greatest disparity in appears closely related to the Pohnpeian en­ the two faunas results from the presence ofsix demic H. halueo, Kosrae has no species that species of the zygopteran genus Teinobasis on does not also occur on Pohnpei. However, Pohnpei and none on Kosrae. The radiation Pohnpei has Paeifieothemis esakii, endemic to of Teinobasis on Pohnpei is a unique situation the island at the genus level, and Agrionoptera in the Caroline Islands; Palau and Chuuk sanguinolenta, which is also absent from (=Truk) each have only one species and Yap, Kosrae. The absence of A. sanguinolenta is like Kosrae, has none. But Chuuk and Yap somewhat unexpected because it is common are relatively low islands with few freshwater throughout the lowlands ofPohnpei (Paulson streams, and Palau, though with only one and Buden 2003), occurs on several disparate Teinobasis, has a total of at least 18 odonate Micronesian atolls (D.W.B., unpub!. data), species. Kosrae, like Pohnpei, is a densely and has been recorded in the Bonin Islands, forested, ruggedly mountainous volcanic is­ Japan, about 2600 km north of the eastern land with high annual rainfall and numerous Carolines (Karube and Sugimura 1997). mountain streams and other wetlands that Odonates are more common in the low- 406 PACIFIC SCIENCE· October 2003 lands than in the mountains on both Pohnpei Pacific Science Board. 7 pp. (mimeo­ and Kosrae, but some species found on both graphed). Available at Hamilton Library, islands tend to extend higher up on the slopes University of Hawai'i at Manoa, Hono­ of Kosrae. Tramea transmarina and Tholymis lulu. tillarga were encountered on the highest ---. 1976. Insects of Micronesia, Micro­ peaks on Kosrae, and T. tilla1'ga was fre­ lepidoptera: Tortricoidea. Insects Micro­ quently flushed from trailside vegetation in nesia 9 (1): 1-144. low- to moderate-elevation rain forest. Nei­ Gressitt, ]. L. 1954. Insects of Micronesia, ther species was observed in cloud forest on Introduction. Insects Micronesia 1:1-257. Pohnpei, and no anisopterans were regularly Hosokowa, T. 1952. A plant sociological encountered in the forest understory there. In study in the mossy forests of Micronesian addition, no Ischnura aurora was observed islands. Mem. Fac. Sci. Kyushu University above 30 m on Pohnpei, but the species was Ser. E (BioI) 1:65-82. encountered at elevations as high as 182 m on Karube, H., and M. Sugimura. 1997. A new Kosrae. Whether such a distribution is a form record of Agrionoptera sanguinolenta Lief­ of ecological release in the presence of fewer tinck from Japan. Aeschna 34:1-4. species on Kosrae requires additional study. Keating, B. H., D. P. Mattey, ]. Naughton, C. E. Helsley, D. Epp, A. Larawicz, and D. Schwank. 1984. Evidence for a hot spot ACKNOWLEDGMENTS origin of the Caroline Islands.]. Geophys. We thank the many residents of Kosrae for Res. 89:9937-9948. their numerous kindnesses and courtesies ex­ Kosrae Branch Statistics Office. 2000. 1999 tended to D.W.B. during the course of field Kosrae State statistical bulletin. FSM De­ investigation, and we are especially grateful partment of Economic Affairs, FSM Na­ to Joseph Methukulathel, biologist at College tional Government, Kosrae State. of Micronesia, Kosrae campus, for providing Lieftinck, M. A. 1962. Insects of Micronesia, housing accommodations in his personal res­ Odonata. Insects Micronesia 5:1-95. idence for the duration of the study. Madison MacArthur, R H., and E. O. Wilson. 1967. Nena and Katrina Adams of the Kosrae Vil­ The theory of island biogeography. lage Hotel and Simpson Abraham and staff at Princeton University Press, Princeton, Kosrae Island Development Review Com­ New Jersey. mission were very helpful in arranging con­ Merlin, M. D., and]. O. Juvik. 1993. Mon­ tacts with local guides. tane cloud forest in the tropical Pacific: Some aspects of their floristics, biogeogra­ Literature Cited phy, ecology, and conservation. Pages 149-162 in L. S. Hamilton, ]. O. Juvik, Allen, ]. A., K. C. Ewel, and]. Jack. 2001. and F. N. Scatena, eds. Tropical montane Patterns of natural and anthropogenic dis­ cloud forests. East-West Center, Hono­ turbance of the mangroves on the Pacific lulu, Hawai'i. Island of Kosrae. Wetlands Ecol. Manage. Merlin, M. D., R Taulung, and]. Juvik. 9:279-289. 1993. Plants and environments of Kosrae Asahina, S. 1940. Odonata-Anisoptera ofMi­ [Sahk kap ac kain in acn Kosrae]. East­ cronesia. Tenthredo 3:1-23. West Center, Honolulu, Hawai'i. Athens, S.]. 1995. Landscape archaeology: Paulson, D. R 2003. Teinobasis budeni sp. nov. Prehistoric settlement, subsistence, and from Pohnpei, eastern Caroline Islands, environment of Kosrae, Eastern Caroline Micronesia (Odonata: Coenagrionidae). Islands, Micronesia. International Archae­ Int.]. Odonatol. (in press). ological Research Institute, Inc., Hono­ Paulson, D. R, and D. W. Buden. 2003. The lulu. Odonata of Pohnpei, eastern Caroline Is­ Clarke, ]. F. Gates. 1953. Report on insect lands, Micronesia. Int. ]. Odonatol. (in collecting trip to Kusaie, Caroline Islands. press). Odonata of Kosrae . Buden and Paulson 407

Raynor, B. 1993. Montane cloud forests in aquatic habitats of Yap. Uillv. Guam Mar. Micronesia: Status and future manage­ Lab. Rep. 92. ment. Pages 176-182 in L. S. Hamilton,]' Steinmann, H. 1997. World catalogue of O. Juvik, and F. N. Scatena, eds. Tropical Odonata. Vols. 1 and 2. Walter de Gruyer, montane cloud forests. East-West Center, New York. Honolulu, Hawai'i. TTP.I. [Trust Territory of the Pacific Is­ Ritter, L. T, and P. L. Ritter (transls. and lands] Office of Planning and Statistics. eds.). 1982. The European discovery of 1979. Kosrae land use guide. TTP.I. Of­ Kosrae Island. Micronesian Archaeological fice of Planning and Statistics, Saipan. Survey Report No. 13. Historic Preserva­ Whitesell, C. D., C. D. MacLean, M. C. Fa­ tion Office, Trust Territory of the Pacific lanruw, T. G. Cole, and A. H. Ambacher. Islands, Saipan. 1986. Vegetation survey of Kosrae, Fed­ Schreiner, I., and D. Nafus. 1989. Insects in erated States of Micronesia. U.S. Dep. the freshwater environments ofYap. Pages Agric., For. Serv., Pac. Southwest For. 21-32 in S. G. Nelson, ed. The inland Range Exp. Stn. Resour. Bull. PSW-17.