Distribution ofthe Chuuk Islands Giant , Acladocricus setigerus (: Rhinocricidae), and Identification of Its Defensive Compoundsl

Donald W Buden, 2,4 Athula Attygalle,3 and Xiaogang Wu3

Abstract: The spirobolidan millipede Acladocrieus setigerus (Silvestri, 1897) grows to at least 155 mm long and is so far known only from Chuuk Islands, Micronesia. It occurs mainly in well-shaded habitats, usually on the forest floor and on tree trunks. It sprays defensive secretions from paired, lateral ozopores on trunk segments; the major compounds, identified here for the first time, are benzoquinones. The secretion stains human skin a reddish brown and causes a slight burning sensation, occasionally followed by slight blistering and exfolia­ tion.

AMONG THE MORE than 600 islands com­ tations of human contact with them. The posing the Federated States of Micronesia study is based largely on specimens, field ob­ (FSM), only six of the larger islands ofChuuk servations, and anecdotal information from (formerly Truk) Lagoon are known to harbor local residents obtained during visits to a millipede exceeding 100 mm in length. Weno, Tonoas, Fefan, Uman, and Tol is­ Taxonomic and phylogenetic relationships of lands, 20 June-1 July 2003. this and other in the Pacific are incompletely studied and we follow Jeekel Previous Reports (2001) in treating the giant millipede on Chuuk as Acladocrieus setigerus (Silvestri, 1897) Silvestri (1897) described Rhinocrieus setigerus pending a systematic review of the . Al­ as a new of spirobolidan millipede though A. setigerus was described over 100 yr from "Ins. Carolinae" (= Caroline Islands) ago, almost nothing is known of its biology. based on a male measuring 150 mm long The scanty and widely scattered literature and 15 mm wide in the Dresden Museum consists largely of morphological descriptions collections. The exact provenance of the and briefly if at all annotated listings in faunal specimen is uncertain. The type was lost survey reports. This study provides new in­ apparently during World War II and there formation on the distribution and habitat is no record of it in Dresden Museum files preferences of A. setigerus together with the (K. Schniebs, pers. comm.). If the Caroline first report on the composition of its defen­ Islands locality is correct, Chuuk is the most sive compounds and the clinical manifes- likely source inasmuch as all other specimens and documented sightings are from there, and comparably sized millipedes are unknown 1 Manuscript accepted 5 February 2004. 2 Division of Natural Sciences and Mathematics, elsewhere in the FSM, which covers nearly College of Micronesia-FSM, P.O. Box 159, Kolonia, all of the Caroline Islands, and where such Pohnpei, Federated States of Micronesia 96941 (e-mail: a conspicuous organism is unlikely to have [email protected]). gone undetected. The Republic of Belau 3 Department of Chemistry and Chemical Biology, Stevens Institute of Technology, Castle Point on Hud­ (Palau) occupies the extreme, westernmost son, Hoboken, New Jersey 07030 (e-mail: aattygal@ Carolines, whence two other large rhinocri­ stevens-tech.edu). cids are recorded: Salpidobolus callosus (Karsch, 4 Corresponding author. 1881) and S. lissior (Chamberlin, 1947). The possibility that the type of A. setigerus origi­ Pacific Science (2004), vol. 58, no. 4:625-636 nated there, though slight, cannot be defi­ © 2004 by University of Hawai'i Press nitely eliminated. There is no evidence to All rights reserved suggest that the type of A. setigerus was col-

625 626 PACIFIC SCIENCE· October 2004

lected anywhere other than in the Caroline grossen] Myriopoda (sitol)" as being among Islands, but it is of at least passing interest the more "striking" [auffallen] en­ to note that another millipede, Tropitrachelus countered on Truk during the German (=Anastreptus) unidentatus, described by Sil­ 1908-1910 Siidsee-Expedition; elsewhere in vestri from Dresden Museum material and the same text, he (Kramer 1932:580 [428]) labeled as from the Caroline Islands is en­ indicated that it is known locally as the "sitol demic to East Africa (R. Hoffman, pers. or sidjo1." Resident islanders interviewed comm.). during this study and asked to spell the Brolemann (1913) erected the genus Dine­ Chuukese name of this referred to it matocricus, with Acladocricus as one of three as sichon (pronounced see chon, with the ac­ subgenera, and which he (Brolemann 1914) cent on the first syllable, ch as in church, and subsequently raised to genus. Attems (1914) on as in the name John). They use this name included Acladocricus as a subgenus within his for all millipedes, however, and seemingly own newly proposed genus Polyconoceras but make no nomenclatoril distinction between without explanation. Hoffman (1974) pointed species. Goodenough and Sugita (1980) de­ out this taxonomic irregularity, and later fined sichon as a "species of black worm (Hoffman 1980) recognized Acladocricus as about six inches long." Davis (1999) defined predominately an Indo-Australian genus with it as "millipede, including several types both 11 species from the Celebes, Solomon Is­ small and large," and he included the name lands, and Philippines, and one (A. setigerus niyen as a synonym, but we did not hear by inference) from the Caroline Islands. that name used during our study. The name Kramer (1932) reported Polyconoceras setigerus sichon is applied also to one or more fern as common in Truk during the German species in the family Polypodiaceae (Good­ 1908-1910 Siidsee-Expedition. In the most enough and Sugita 1980, Merlin and Juvik recent synopses of Indo-Australian spi­ 1996). We suggest the English vernacular robolidan taxa, Jeekel (2001) recognized A. name "Chuuk Islands Giant Millipede" for A. setigerus from Truk in the Carolines, with setigerus. It is among the largest of the Pacific congeners from the Celebes (9), Philippines Islands millipedes, although on a global per­ (4), and Solomon Islands (2), and Marek et a1. spective it is more mid-sized; some of the (2003) presented a similar list of Acladocricus African spirostreptids are approximately twice species, but added two Australian species that as long as A. setigerus. Jeekel (2001) recorded under Adelobolus and transferred A. hicksoni (Pocock, 1894) of the Study Area Celebes to the genus Salpidobolus. In the preliminary report on his ento­ Chuuk (151° 22' to 150° 04' E and r 07' to mological survey of Chuuk in 1949, Potts 7° 41' N) is one offour states composing the (1949) briefly mentioned encounters with a FSM. It is located near the center of the large species of millipede that he referred Caroline Islands chain in the west-central to as Polyconoceras callosus but is almost cer­ Pacific Ocean (Figure 1). The Carolines span tainly A. setigerus; the species Polyconoceras approximately 3,200 Ian from Palau eastward (= Salpidobolus) callosus is known only from to and including all of the FSM-Yap, Palau (Jeekel 2001). A photograph (Gressitt Chuuk, Pohnpei, and Kosrae States. The ap­ 1954:83, fig. 38) taken on Mt. Winipot, Tol proximately 290 islands of Chuuk State (127 Island, in 1953 shows an aggregation of giant Ian2 total land area) include the 19 volcanic millipedes on a tree trunk, which are identi­ islands and 96 coralline islets of Chuuk La­ fied in the caption as "? Polyconoceras." goon, together with 14 widely scattered atolls and low, single coral islets. The volcanic is­ Vernacular Names lands (Figure 2) are the emergent remains of a shield volcano that originated over a hot Kramer (1932:577 [426 in the original Ger­ spot 14-8 million yr (myr) B.P. (Keating et a1. man text]) recorded the "finger-sized [finger- 1984a) and about 1,500 Ian southeast of its Chuuk Islands Giant Millipede . Buden et al. 627

GUAM ,.

10° CHUUK POHNPEI

IS LAN D S KOSRAE

SOLOMON ISLANDS

1000 kIn

FIGURE 1. Location of the Caroline Islands.

current position. The reef and its associated (18.9 km 2/373 m), Fefan (13.2 km 2/303 m), 2 2 2 islets enclose a lagoon 2,130 km in area and Tonoas (8.75 km /343 m), Udot (4.9 km / "presumably trace the former extent of the 242 m), and Uman (4.7 km 2/273 m); the volcano which may have had a height of 1,200 others are less than 2.0 km 2 and below 150 m m above present sea level" (Keating et al. (data from Bryan 1971). 1984a:350). Currently, the largest (and The climate of Chuuk is warm and humid highest) islands in decreasing order of land year-round. Average annual rainfall based 2 area are Tol (34.2 km , 443 m high), Weno on Weno weather station records for 1946- 628 PACIFIC SCIENCE· October 2004

...... "

' . .... ",

CHUUK LAGOON

N .... 1 \)<>

"

.'

.' ," " " . ,0' • ...... ' ... ' ...... ' . , .' ... ' lOKM

FIGURE 2. Location of Chuuk Lagoon Islands; dotted line indicates outer edge of the reef.

1990 is 3,632 mm, with the driest months were counted during the 2000 census, in­ (January-March) averaging 150-250 mm/ cluding 8,930 on Tol, the largest island month (Merlin and Juvik 1996). The average (treated as three islands [Tol, Polle, and annual temperature is approximately noc, Patta] separated by mangrove channels in with a daily range of about 6°C (Merlin and some references), and 13,808 on Weno, the Juvik 1996), and the humidity ranges from capital (Division of Statistics 2002). Centuries an average of 78% during the day to 87% at of human occupation have altered the vege­ night (Department of Planning and Statistics tation greatly, which currently consists of 1996). weedy shrubs and herbs, grasslands, second­ Chuuk is the most populous of the FSM ary woodland, agroforests, and other areas of states, with about half of the nation's popula­ cultivation. Remnants of primary forest in tion on less than one-fifth its land area Chuuk are now confined to the less-accessible (Merlin and Juvik 1996); 53,595 residents central peaks on the higher islands (Merlin Chuuk Islands Giant Millipede . Buden et at. 629 and Juvik 1996, Mueller-Dombois and Fos­ Meristics and Mensuration berg 1998). Acladoericus setigerus is a large, robust milli­ pede (Figure 3A,B,C,E): maximum length 155.0 mm, width 15.7 mm, weight 25.0 g; MATERIALS AND METHODS number of trunk segments 50-55, modally 52 Logistical problems involving interisland (10 of 26 = 39%). No sexual dimorphism in transport from Weno limited the survey to size was evident in 24 males (Table 1) and five islands. With the exception of Weno, three females-two from Tonoas (102 and no more than 6 hr was spent on each island 113 mm long) and one from Uman (148 mm). during each visit; Tonoas was visited twice, No hatchlings or very young were observed; the others once each. Millipedes were col­ the smallest individual encountered was 80 lected by hand using forceps 23 em long to mm long. Aggregations of millipedes ap­ coax them gently into cardboard boxes con­ peared to consist largely of individuals of taining leaf litter and duff, though rarely similar size. The six males collected within an without their ejeoting defensive secretions; area of about 1.5 m 2 on the ground beneath a they were brought to Weno for processing large abo tree (Dysoxylon abo) on a mountain later the same day. The collection was some­ slope in Fefan were uniformly large and what biased toward obtaining large speci­ nearly completely separated from all others in mens. Defensive compounds were obtained total length (Table 1). Ozopores through by placing a millipede in a glass jar and shak­ which defensive compounds are released are ing the jar to stimulate release of the secre­ present bilaterally on nearly all trunk seg­ tions, and by striking the millipede directly. ments, beginning with the sixth. They are Secretions collected on filter paper and in occasionally absent or very much reduced in glass microcapillary tubes were sealed in glass size on the terminal segment and less fre­ ampules using a microtorch and shipped to quently so among some of the midbody seg­ laboratory facilities in Hoboken, NewJersey, ments. Scobinae (paired, dorsal paramedian where they were kept frozen until chemical pits present on the trunk segments in some analyses could be performed. rhinocricids) are absent in A. setigerus.

Analytical Procedures Distribution For gas-chromatographic analyses, the tubes Acladocricus setigerus is known only from were opened and each sample was extracted Weno [=Moen], Tonoas [= Dublon], Fefan with 50 J..lL of dichloromethane. A 1.0-J..lI ali­ [= Fefen], Uman, Udot, and Tol, the six quot of each sample was analyzed by GC-MS largest islands in Chuuk Lagoon. We include using a gas chromatograph (HP 5890 II) Udot in the list of locality records based on linked to a mass selective detector (HP 5970); personal communications from a resident of a 25 m by 0.20 mm fused-silica column the island and from a former resident now coated with 0.33 Jlm DB-5 was used. The living on Pohnpei, both ofwhom we consider oven temperature was kept at 40°C for 3 min reliable sources; all other locality records and raised 6°C/min to 270°C. are from Potts (1949) and this study. Potts (1949:10) stated "the large julid milliped of the Truk area was quite common at the OBSERVATIONS AND RESULTS higher altitudes on Moen, Dublon, and Tol." Coloration During our study, A. setigerus was encoun­ tered on Tonoas, Fefan, Uman, and Tol Is­ Body color in A. setigerus is dark brown lands, but not on Weno despite more time (nearly black in most specimens), occasionally spent in the field there than elsewhere; other paler (medium to light brown), and rarely islands were not visited. Residents of Weno with a grayish cast. variously indicated that it was common in the FIGURE 3. Acladocricus setige'r'lts (Silvestri). A, whole animal; B, side view of midbody segments showing ozopores near top of view; C, part of an aggregation of six on trunk of coconut tree; D, fingers of collector stained with defensive compounds ofA. setigel"llS; E, coiled, defensive posture. Chuuk Islands Giant Millipede . Buden et at. 631

TABLE 1 Measurements (Range, Mean, Sample Size in Parentheses) for Male AcladocricZlS setigenlS Collected on Five Chuuk Lagoon Islands during June and July 2003

Character Tonoas Fefan Uman Tol

Body length (mm) 80.0-135.0 134.0-155.0 114.0-135.0 135.0 (1) 112.4 (14) 144.0 (6) 121.7 (3) Body width (mm) 9.6-14.0 14.6-15.5 13.7-14.7 15.7 (1) 12.5 (14) 15.1 (6) 14.2 (3) Weight" (g) 19.0-20.0" 23.5-25.0 22.0 (1) 19.7 (3) 24.6 (6)

n Measured on the three largest specimens; 126, 130, and 135 mm long.

mountains, that it was so in the past but not (1949) mentioned only the three large islands now, or that they were completely unfamiliar where he found it common, though he visited with it. One resident provided directions to at least five other lagoon islands (Gressitt a specific site in central Weno where she 1954). A student at the College ofMicronesia believed the sichon could be found, but a reported (pers. comm.) that it did not occur half-day search of that area and beyond to an on her home island, Eot, which is just off elevation of about 274 m with two local as­ Udot, where it does occur, and we received sistants was unsuccessful. Harvey Segal, a an unconfirmed report from a member of the long-term local naturalist and instructor in Chuuk Public Health Immunization Survey the Education Department at the College of Team that it does not occur on Romanum, Micronesia, recalled seeing giant millipedes following his inquiry on our behalf during a in the vicinity of the Japanese gun emplace­ brief visit to the island. ments above the Chuuk Hospital grounds when he first arrived in Micronesia in August Identification ofDefensive Secretions 1964, but two visits during the recent survey revealed none. IfA. setigerus is still present on The major defensive compounds of A. Weno, it appears to be very scarce or ex­ setigerus are toluquinone (2-methyl-l,4­ tremely localized or both. On other islands benzoquinone) and 2 methoxy-3-methyl-l,4­ visited during 2003, it was encountered benzoquinone. widely but discontinuously from sea level to high on the forested slopes, being abundant Effects ofDefensive Secretions on Humans in some areas but apparently absent in inter­ vening ones of similar habitat. Some were The clinical manifestations of human contact exposed on tree trunks (mainly coconut, Cocos with the secretions of A. setigerus were first nucifera), but most were on the ground com­ reported by Potts (1949: 10), who wrote ofhis pletely or partially covered by leaves, dead encounters with "Polyconoceras callosus" on branches, clumps of grass, and procumbent Chuuk as follows [typographical errors re­ vines. Acladocricus setigerus prefers shaded tained as in original]: areas usually among the sprawling root bases "On one occasion, an average sized speci­ of large trees, and it was found in the settle­ men (about 4-5 inches) was disturbed when I ments as well as (and more frequently) in put my hand within an inch of it while more remote and less-disturbed parts of the scrambling up a moist slope. A considerable forest. blob of the brown liquid emmited by the To what extent A. setigerus occurs on the beast was sprayed onto the back of my hand smaller islands, if at all, is uncertain. Potts while small drops of it were thrown up my 632 PACIFIC SCIENCE· October 2004

arm to a distance of at least 15 inches. There DISCUSSION was an almost immediate slight burning sen­ sation, particularly on the tenderer skin of the Defensive Secretions arm. The sensation presisted for some five or Millipedes secrete a variety of defensive ten minutes. The caustic action, however ap­ compounds (Eisner et al. 1978, Attygalle et al. peared very minor although there was a per­ 1993, Dernl and Huth 2000, Huth 2000). sistent red-brown stain. At another time I Acladocricus setigerus secretes quinones, as is inadvertently crushed a specimen under my typical of species in the orders Spirobolida, light tennis shoe. No discomfort was noted, Spirostreptida, and Julida, and it can spray but the stain penetrating the canvas lasted for them at least 38 cm (Potts 1949). Maximum around two weeks despite repeated purpose­ ejection distances for millipede secretions ful scrubbing with every shower." range to about 84 cm in Rhinocricus latespargor During the 2003 survey, all millipedes from Haiti (Loomis 1941:192) and "more were collected and handled by D.W.B. using than 60-70 cm" in Polyconoceras (= Salpidobolus metal forceps 23 cm long, bringing the hand ?) sp. in New Guinea (Jolivet 1970:51). Qui­ to within about 15 cm of the specimens. nones are repellent to many and Many sprayed their secretions when touched other potential millipede predators (Eisner et even gently. The spray was extremely fine or al. 1978). In a possible zoopharmacological was colorless when released, because it was application of these compounds, monkeys not observed until after it was in contact with in gallery forest of Venezuela were observed a solid surface. No irritation from the secre­ rubbing the quinine-producing, spirostreptid tion was felt on the thumb and forefinger, millipede Orthoporus dorsovittatus on their which were the most extensively stained areas bodies, with as many as four monkeys sharing (Figure 3D), but a slight burning sensation a single millipede (Valderrama et al. 2000). lasting for several minutes to about half an Although the protective, repugnatorial effects hour was felt on the back of the hand, which have not been confirmed in that case, a posi­ was only slightly spotted with secretion each tive correlation exists between anointment time. None of the secretion apparently frequency and periods of high rainfall when reached farther up the arm. The stain per­ mosquitoes and possibly other biting insects sisted on the skin for about a week, and there are more abundant (Valderrama et al. 2000). was no blistering or exfoliation. In addition to Weldon et al. (2003) reported similar behav­ staining and irritating the skin, the secretions ior involving self-anointment with millipedes ofA. setigerus have a disagreeable, acrid, pun­ in wild and captive capuchin monkeys, and gent odor, somewhat reminiscent of an elec­ they experimentally demonstrated possible trical wiring fire. repellency of quinone compounds found in Resident islanders queried throughout the the millipedes to Aedes aegypti mosquitoes. study reported only minor discomfort from All millipedes that are known to spray contact with the secretion, the most frequent their defensive compounds produce quinones comment being that millipedes stained the (Eisner et al. 1978). These compounds stain feet red when stepped on. Three of them reddish brown or purple and cause various stated that children often played with the degrees of irritation to skin and mucous millipedes. One former teacher on Tol re­ membranes of eyes and nose in humans (Eis­ called that one of his students was sprayed in ner et al. 1978, Schpall and Frieden 1991, the eyes during a field trip to Mt. Winipot Hopkin and Read 1992, Mason et al. 1994, many years ago and though temporarily Hudson and Parsons 1997, Huth 2000). blinded suffered no long-lasting injury; she McNamara (in Musgrave 1943) reported that was able to descend the mountain without his dog was permanently blinded in one eye assistance later in the day. None of several following an encounter with a large millipede Chuuk Hospital workers questioned during in New Guinea, but Hudson and Parsons this study knew of any incidents of millipede (1997) found no documented evidence to burns being treated at the hospital. support the claim by many New Guinea is- Chuuk Islands Giant Millipede . Buden et at. 633

landers that the secretions of giant millipedes ingly without close relationships." Lieftinck sprayed into the eyes can cause permanent (1962:3) declared that Teinobasis carolinensis, a loss of sight in humans. damselfly endemic to Chuuk, "is a highly in­ teresting and aberrant zygopter, unlike any­ Biogeography thing known in this diversified genus ... ," and Craig and Joy (2000) and Craig et al. Millipedes have been largely understudied, (2001) reported that among the 40 species of and the Spirobolida of Oceania are among black flies in the Pacific subgenus lnseliellum those that are poorly known (Jeekel 2001); that they studied, molecular evidence in­ uncertainty surrounds the of many dicated that Simulium trukense, endemic to species currently assigned to the genus Acla­ Chuuk, is the most distantly related lineage docricus and allied Rhinocricidae (c. A. W. and the only species of the subgenus in the Jeekel, pers. comm.). Whether A. setigerus is Caroline Islands. Among terrestrial verte­ indigenous and possibly endemic to Chuuk, brates, three of 12 resident breeding land­ or whether it was introduced by the earliest bird species are endemic (Pratt et al. 1987), aboriginal colonists or their followers, or the most distinctive being the Chuuk Mon­ brought to the islands later by European and arch, Metabolus rugensis, belonging to a American explorers, traders, and missionaries monotypic genus and apparently most closely during travels in the Pacific cannot be stated related to a species endemic to the Solomon with certainty. Islands (Baker 1951). The taxonomic relationships and distribu­ If A. setigerus is not endemic to Chuuk it tion of A. setigerus (as currently understood), may still be indigenous to these islands, hav­ along with the geological history of Chuuk ing arrived from an as yet unknown source, and the occurrence of well-differentiated probably by rafting. Alternatively, it may have forms among taxa other than millipedes, sup­ been introduced by the first aboriginal set­ port, or at least are compatible with, the view tlers. Humans appear to have dispersed into that A. setigerus is endemic to these islands. central Micronesia, including Chuuk, from Chuuk islands are among the oldest in the western Micronesia and from Melanesia to Carolines (14-8 myr B.P. [Keating et al. the south about 2,000 yr ago (Intoh 1997, 1984a,b]). They are the emergent peaks of a 1999). East-to-west movements within Mi­ once-massive island that would have provided cronesia occurred about the same time, as did a large target for colonists and opportunities dispersal northwestward from Polynesia to at for establishment and differentiation. Al­ least as far as Nukuoro and Kapingamarangi though described well over 100 yr ago, A. se­ atolls in Pohnpei State (Intoh 1997). Inas­ tigerus remains known definitely only from much as "the cultures of Micronesia were Chuuk. Its absence elsewhere in the Caro­ formed by complex movements of human lines, at least in the FSM, is real because such populations" and from different directions a large and conspicuous animal that occurs in (Intoh 1997:25), our current knowledge of aggregations and exposed on tree trunks in these movements does not by itself point to a daylight would not have gone unnoticed; most likely source ofA. setigerus on Chuuk. there are no records of large millipedes on A more recent introduction by post­ Yap, Pohnpei, and Kosrae. Chuuk hosts well­ sixteenth-century European and American differentiated endemic species in a variety explorations is another possible explanation of taxa, some without any close represen­ for the occurrence of A. setigerus on Chuuk, tatives elsewhere in the FSM, and A. setigerus but those visits were relatively infrequent. possibly falls into this category. Gressitt Chuuk "remained relatively unknown to the (1954:139) stated that the insect fauna of outside world throughout most of the 19th Truk [Chuuk] "is puzzling in its represen­ century ... and enjoys the distinction ofbeing tation and affinities ... [being] exceedingly the last major island group in Micronesia to depauperate in some groups, whereas in have been opened to intensive contact with others it has peculiar endemic forms, seem- the colonial powers ofEast and West" (Hezel 634 PACIFIC SCIENCE· October 2004

1973:51). Furthermore, A. setigerus does Team and arranged by Jean-Paul Chaine and not appear to be especially predisposed to Kidsen Iohp. We thank R Hoffman, R Me­ human-assisted introduction. It has been sibov, and an anonymous reviewer for their known from Chuuk for about 100 yr, but it helpful comments on a preliminary draft of has not become established (or at least is not the manuscript, and Doug Kelly for prepar­ now present) on any of the other islands in ing the composite illustration ofA. setigerus. the FSM, where seemingly suitable habitat and comparable environmental conditions are Literature Cited available and where interisland traffic has be­ come increasingly more frequent. Attems, C. G. 1914. Die indo-Australischen In summary, A. setigerus has been recorded Myriopoden. Arch. Naturgesch., Abt. A definitely only in Chuuk during the approxi­ 80 (4): 1-398. mately 100 yr since it was first described and Attygalle, A B., S.-c. Xu, J. Meinwald, and could justifiably be considered endemic to T. Eisner. 1993. Defensive secretion ofthe these islands; its absence elsewhere in the millipede Floridobolus penneri. J. Nat. Prod. FSM almost certainly is real. But in view of 56:1700-1706. the lack of comprehensive systematic studies Baker, R H. 1951. The avifauna of Micro­ of millipedes in the Pacific area and the pos­ nesia, its origin, evolution, and distribu­ sibility of it occurring elsewhere outside the tion. Univ. Kans. Mus. Nat. Rist. Publ. FSM, we suggest that it be treated as a cryp­ 3:1-359. togenic species (sensu Carlton 1996), possibly Br6lemann, H. W. 1913. The in being indigenous and endemic to Chuuk but the Australian Museum. Part II. Dip­ of unknown origin and of incompletely re­ lopoda. Rec. Aust. Mus. 10:77-158. solved phylogeny. ---. 1914. Etude sur les Spirobolides (Myriapodes). Ann. Soc. Entomol. Fr. 83:1-38. ACKNOWLEDGMENTS Bryan, E. H., Jr. 1971. Guide to place names in the Trust Territory of the Pacific Is­ For responding to our numerous queries, we lands. Pacific Science Information Center, thank Sergie Golovatch, Richard Hoffman, B. P. Bishop Museum, Honolulu. C. A. W. Jeekel, Robert Mesibov, Rowland Carlton, J. T. 1996. Biological invasions Shelley, Petra Sierwald, and Tsutomu Ta­ and cryptogenic species. Ecology 77:1653­ nabe, and we are especially grateful to R 1655. Hoffman for providing the initial identi­ Craig, D. A, and D. A Joy. 2000. New fication of specimens. For their mu,ch­ species and rediscriptions in the central­ appreciated assistance with the literature sur­ western Pacific subgenus Inseliellum (Dip­ vey, we thank the librarians at the University tera: Simuliidae). Ann. Entomol. Soc. Am. ofGuam, University ofHawai'i, University of 93:1236-1262. Massachusetts Medical School, the College Craig, D. A., D. C. Currie, and D. A Joy. of Micronesia, and Micronesia Seminar, and 2001. Geographical history of the central­ we extend a special thanks to Mary Sears, western Pacific black fly subgenus In­ Ernst Mayr Library, Museum of Comparative seliellum (Diptera: Simuliidae: Simulium) Zoology (MCZ) for her efforts in providing based on reconstructed phylogeny of the obscure reference materials; Van Wallach species, hot-spot archipelagoes and hy­ (MCZ) kindly examined additional references drological considerations. J. Biogeogr. for us. We also thank Katrin Schniebs for 28:1101-1127. her efforts in trying to locate information on Davis, AD. 1999. A preliminary list of ani­ the type material deposited in the Dresden mal names in the Chuuk District, with Museum. Transportation to Tol and Tonoas some notes on plant names. Micronesica Islands from Weno was provided courtesy 32:1-245. of the Public Health Immunization Survey Deml, R, and A Huth. 2000. Benzoquinones Chuuk Islands Giant Millipede . Buden et at. 635

and hydroquinones in defensive secre­ Jeekel, c. A. W. 2001. A bibliographic cata­ tions of tropical millipedes. Naturwissen­ logue of the Spirobolida of the Orien­ schaften 87:80-82. tal and Australian regions (Diplopoda). Department of Planning and Statistics. 1996. Myriapod Memo. 4:5-103 [privately Chuuk State census report, 1994 FSM published by the author, Oisterwijk, The Census of Population and Housing. Netherlands]. Chuuk State Government. Weno, Chuuk. Jolivet, P. 1970. A propos de l'extreme tox­ Division of Statistics. 2002. FSM national icite d'une espece geant de Myriapode detailed tables, 2000 FSM Census of Diplopode de la Nouvelle-Guinee aus­ Population and Housing. Department of tralienne appartenant au genre Poly­ Economic Affairs. FSM National Govern­ conoceras Attems. Entomologiste (Paris) ment, Palikir, Pohnpei. 26 (1-3): 31-40,47-54. Eisner, T., D. Alsop, K. Hicks, and J. Mein­ Keating, B. H., D. P. Mattey, J. Naughton, wald. 1978. Defensive secretions of mil­ and C. K Helsley. 1984a. Age and origin lipedes. Pages 41-72 in S. Bettini, ed. of Truk Atoll, eastern Caroline Islands: Arthropod venoms. Springer-Verlag, Ber­ Geochemical, radiometric-age, and paleo­ lin. magnetic evidence. Geol. Soc. Am. Bull. Goodenough, W. H., and H. Sugita. 1980. 95:350-356. Truk-English dictionary. American Philo­ Keating, B. H., D. P. Mattey, C. E. Helsley, sophical Society, Philadelphia. J.J. Naughton, D. Epp, A. Lazarewicz, and Gressitt, J. L. 1954. Insects of Micronesia, D. Schwank. 1984b. Evidence for a hot­ introduction. Insects Micronesia 1:1-257. spot origin of the Caroline Islands. J. Hezel, F. X. 1973. The beginnings offoreign Geophys. Res. 89:9937-9948. contact with Truk. J. Pac. Hist. 8:51-73. Kramer, A. 1932. Truk. Ergibnisse der Hoffman, R. L. 1974. Studies on spiroboloid Siidsee-Expedition 1908-1910. II. Ethno­ millipeds. X. Commentary on the status of graphie: B. Mikronesien, Vol. 5. Friede­ Salpidobolus and some related rhinocricid richsen, de Gruyter and Co. Hamburg genera. Rev. Suisse Zool. 81:189-203. [translated from the German in 1942 for --. 1980 [1979]. Classification of the the Yale Cross-Cultural Survey in con­ Diplopoda. Museum d'Histoire Naturelle, nection with the Navy Pacific Islands Geneva. Handbook Project. Human Relations Area Hopkin, S. P., and H. J. Read. 1992. The Files 1968]. biology of millipedes. Oxford University Lieftinck, M. A. 1962. Insects of Micronesia. Press, Oxford. Odonata. Insects Micronesia 5:1-95. Hudson, B.]., and G. A. Parsons. 1997. Giant Loomis, H. F. 1941. New genera and species millipede 'burns' and the eye. Trans. R. of millipeds from the southern peninsula Soc. Trop. Med. Hyg. 91:183-185. of Haiti. J. Wash. Acad. Sci. 31:188-195. Huth, A. 2000. Defensive secretions of milli­ Marek, P. K, J. K Bond, and P. Sierwald. pedes: More than just a product ofmelting 2003. Rhinocricidae systematics. II: A spe­ point decrease? Pages 191-200 in J. cies catalog of the Rhinocricidae (Dip­ Wytwer and S. Golovatch, eds. Progress in lopoda: Spirobolida) with synonymies. studies on Myriapoda and Onychophora. Zootaxa 308:1-108. Fragm. Faun. (Warsaw) 43 (Supplement). Mason, G. H., H. D. P. Thomson, P. Fergin, Intoh, M. 1997. Human dispersals into Mi­ and R. Anderson. 1994. Mysterious le­ cronesia. Anthropol. Sci. 105:15-28. sions. Med. J. Aust. 160:718, 726. ---. 1999. Cultural contacts between Mi­ Merlin, M., and J. Juvik. 1996. Plants and cronesia and Melanesia. Pages 407-422 in their environments in Chuuk. East-West Jean-Christophe Galipaud and 1. Lilley, Center, Honolulu. eds. The Pacific from 5000 to 2000 B. P. Mueller-Dombois, D., and F. R. Fosberg. Institut de Recherche pour Ie Developpe­ 1998. Vegetation of the tropical Pacific Is­ ment, Paris. lands. Springer-Verlag, New York. 636 PACIFIC SCIENCE· October 2004

Musgrave, A. 1943. Some arachnids and mil­ 1987. A field guide to the birds of Hawaii lipedes from New Guinea. Aust. Mus. and the tropical Pacific. Princeton Uni­ Mag. 8:132-135. versity Press, Princeton, NewJersey. Potts, R. W. L. 1949. Preliminary report on Schpall, S., and 1. Frieden. 1991. Mahogany an insect survey of Truk. [mimeograph.] discoloration of the skin due to the defen­ Pacific Science Board of the National Re­ sive secretion of a millipede. Pediatr. Der­ search Council. [At least two versions of matol. 8:25-27. this article may exist. The copy made Valderrama, x.,]. G. Robinson, A. Attygalle, available through the University of Ha­ and T. Eisner. 2000. Seasonal anoint­ wai'i is undated by the author, but 1954 is ment with millipedes in a wild primate: A part of the "call number" handwritten in chemical defense against insects? ]. Chern. the upper left corner of the title page; the Ecol. 26:2781-2790. article is 23 pages long (19 numbered Weldon, P. J.,]. R. Aldrich,]. A. Klun,]. E. pages plus four appendices, A-D). Gressitt Oliver, and M. Debboun. 2003. Benzo­ (1954:213) indicated a publication date of quinones from millipedes deter mosqui­ 1949 and 25 pages for presumably the toes and elicit self-anointing in capuchin same report.] monkeys (Cebus sp.). Naturwissenschaften Pratt, H. D., P. L. Bruner, and D.]. Berrett. 90:301-304.