NATIVE TERRESTRIAL INVERTEBRATES

Wayne C. Gagn6

The fint collections of terrestrial invertebrates from the Hawaiian Islands were made during Captain James Cook's voyages in 1778 and 1779. Little of this material was examined scientifically other than by Fabricius (1792-1794), who described the large ichneumon wasp Eclltllromorphufuscator and the vespid hornet Odynem radulu from these collections. The ultimate disposition of specimens, and which ones subsequently survived, became immenselv comolicated (Whitehead 1978).

The &llect/ons of t&restrial mollusk;, including the genus Acltarinellrr (large tree snail?, endemic to O'ahu) and other groups of tree snails, assembled th r~whout the 1XOOs hv earlv ex~lorer-naturalists. curious mkionaries. and their i~ffrpring gained .&on5id&ably more attention than other invertehratec (Ka! 1472). prewmahly hecause of comparative ease of recervatlon in addi- lion to \~~ect;rcuIar color patterns. Toward the close of t i e 19th ccnt~lrv and for many decades following, professional malacologists (mollusk reseaichers), including C.M. Cooke, Jr. and Y. Kondo at B.P. Bishop Museum in Honolulu, published several large systematic treatments of these groups, documenting several hundred endemic soecies. The recoenition of intra-soecific varia- tion, speciation, and endenh t ) of ~awaiia; landthells hy ~ ; h n Gulick helped hring the .mplicationr of Darwinian evolution before the glohal srient~fic hodv lGul~ck 1932). The imoortance of hasine taxonomic rclation- ships and the%ore classification of mbllusks on the en&e animal ra the~ than only the shell was also recognized.

Apait from some largely incydental collections made during various expeditions from 1778 to 1877 (see Illingworth 1923), other Hawaiian arthropods (insects, spiders, sowbu s, and so forth) remained largely B unknown. Exce tin a collection o true bu s (Hemiptera) made during the *sit of the ~ w e g s h !ripe Eugrni in 1853 8.1 IXSY), it was not until the arrival of the Rev. T. lackburn in 1877 (resident in Hawai'i for about a decade) that the first major systematic collections of insects were assembled (Blackburn 1877, 1881). Thus, a century of Western exploration passed while the terrestrial invertebrates, other than large land snails, remained largely unknown.

The most important collections of terrestrial arthropods were assembled by the extraordinary naturalist R.C.L. Perkins from 1893 to 1903. He w a sponsored by the Royal Society of London and the British Association for the Advancement of Science, which, in conjunction with the B.P. Bishop Museum,

ublished the results of Perkins' labors in three monumental volumes of the ! auna Hawaiiensis series (Sharp 1899.1913). Duplicate material was depos- ited at the British Museum of Natural History and the B.P. Bishop Museum (Manning 1986).

For a number of major groups, especially the Coleoptera (beetles) and many Hymenoptera (bees, wasps, and ants),Fauna Hawaiiencir, though now serious1 dated, is still the "state of the art." Revisions of several genera and liig i er taxonomic groups (see natural classification in Glossa ) and 2' biolo 1cal studies of aquatic insects (Williams 1936-1944) appeare In the decacfes following publication of Fauna. In 1948, E.C. Zimmerman began the immense task of updating the Hawaiian arthro ods in his Insects of Hawaii series. Fourteen volumes have appeared thus Par, with the Diptera or flies authored in five volumes by D.E. Hardy (1960-1981).

Swezey (1954) summarized the host associations of all insects known to be associated with native trees. Examination of his Forcsr Entomolog~ and Zimmerman's volumes shows that the life histories of the bulk of the de- scribed native species are still unknown.

Work on the Hvmenootera for the Insects of Hawaii volumes bv C.Y. Yoshirnoto and J.\\'. l<c.ar:lile~ i.; airrently inp;ogress. Nothing ofthis magnitude is yet in the \ r w h ror the Coleo tera G A S~muelsox and G.M. Nishida of the I:ntomol<~p lkpnrrment at f ; . ~ . Ekshbiicseurn are cornput- erizing the card file o i I!;~rr..~~ian rcrrestrial invertebrates, hoth nat.ve a r J alien, developed hy the lare ;\m!. Suehiro. Guslw Pa.lley of the Lni\,ercity of Washington has prcp:~rcd a computer listing or 11: known endernlc terrrr- trial invertchrates. Whcn these projects are complemi, we will knou how many species have been described. as well as the~r b~ogeographic sr;itus. Recent cstimates of the number of endemic arthropod species lie hetaccn 7.000 and 10,000 (Simon 1987: F.G. Howarth, personal communicnr~~~rj. For the mites, Gnff (19x7) has provided an up-to-date carafog containi.~g 4h11 species. I3ut the endemicity of acarineo mites and reldrlve~) i c etcc,pr~~~ri;~lly tlifficult to determine, and Goff does no! attempt to do so. Surn:~n ( I ~ ? J pruvided a cdtalog of Hawaiian spiders totalling 149 specier, of a.111cli 3': are endemic. 170 are indigenous (naturally occurring in llswsi i 1v.r naturallv found elsewherealso), and the rest are alien.

J . d h H. Bryan, Jr.. the first to popularize Hawaiim inverrehr:~rc~. [~ubli,hed a series of nature notes in the Honolulu Yar-Bullc,tin nca.'pap~,r r!mw the 1920s (Brvan 1935). He also ~uhlishccl a short. elemcn!clr\ ;I:CIIIIII: of co&non. laree'arihronnds'for use in the schools (Brva" 19401. ~ ~~

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It was "ot Cntil the 1970s that focus was brought upon the &ervation status of terrestrial invertebrates (Cam6 1975: Hart 1978: Hadfield 1986). Followine Hart's article. the U.S. DeGrtmentof the Interior (USDI) olaced the entir;genus~cl,ar;iaa on t h e ~;tdangered Species list. ~ e v e r a i ' hundred other terrestr~al inwrrehrutcz were placed under rcview hy USDI, but Drosoects for timelv official listing of anvof these now aooear dim.

iven thk prevahng p$i:icdl clini:~tc.'~ot hhclcred by such ;;~nstrair~t;, the hernational L'fiion for the Conwm~tion of Nature and Natural Resnllrces (IUCN) has listed several Hawaiian arthropod groups in itsIUCNRed Dara Bookon Invertebrates (Wells pt al. 19x3). The cnnservatinn stat~ls nf endemic ~ ~- ~ -.

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terrestrial invertebrates i!i :r.nc.r~l na, di,c.~rr&l hy GagnC and Christensen (1985): that of L e ~ i d o m r i I hut!e:rlle; snJ mrlthr) bv GaenC and Howarth (1985); that of a vblne;able damselfly by Moore and GagG (1982); and that of Hawai'i's cave invertebrates by Howarth (1983).

In recent years, major strides have been made in elucidatine the rvstematics and evolutibn of a few maior eronns: the nrnsonhil:dae (see

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&son and Kaneshiro 1976), cave-ad$& and aeolian (dependent & wind-distributed food) arthropods (Howarth 1983, 1987). and ~redatory inch worms (Montgomerv 1983). The ecolow and altitudin&dktri'hution or these

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and several ofi~cr grbupr xrr. di.;cus\ed hy various authors in Mueller- Domhoic et <I/. (10x1 J. I\ rcvuv of thc. +milicmce uf all these studies to entomolon wsr m:~J r I>\ Slrnm~ uI. ( l4h4 ) and Smon (1987 I. Several

MAJOR PROBLEMS

We arc f ~ r from ~.t~mpletic~n of a full systematic treatment of all terrestri:~l inverlehrdtes in Hawai'~. particularly groups with many s ecies, such as t h t small mollusks, ncmotodcs. spiders, mites, and insects. ? he b~olow, ecolopv. and evolution of th&e and the remainine invertebrates isiareelv a vast ~Gknown. Nor do u e know much about the hportance of invcr6b;atcc to the rest of the n:ltlve liotn. For example, pollinat~on hiolop I < ntarlv ;I . . -. virgin field.

With support of the U.S. Fish and Wildlife Service, I intend to sample tree canopy arthropods to see if there are any correlations with distributions of certain Endangered forest birds. What is the nature of other interdepen- dencies of native arthropods and the remainder of the biota? What is the role of native invertebrates in nutrient cycling? These and other equally critical research questions should he answered.

In regard to popularized treatments of Hawaiian invertebrates, there are but two (Williams 1931; Fullaway and Krauss 1945), which are very dated, long out of print, and not readily available. Frank Howarth, William Mull, and I are preparing a copiously illustrated, popularized account of native insects.

RESEARCH AND MANAGEMENT NEEDS

Much ecological work involving terrestrial invertebrates k potentially hamstrune hv the rudimentarv knowledee of their svstematics and a lack of ~ ~~~~ ,~ ~ ~~~

trained an% paid s)stemati~ts'to car? nzt this work: There arc less than a half-dozen Invertebrate s\stemat~~t.; in I Inwai'i. who are paid to study 7.000-10.000 soecies. There are no Hawai'i-based work& to studv the systemahcs of'terrestrial arachnids, myriapods, crustaceans, annehds, mollusks, and a number of insect groups.

This near Impasse at the .;?&tnatic le\cl makes intelligent management of the immense invertehrntc rcwurse LI iflwlr. ti not impnsihlc. 'lhc recent work of Howarth, h o p e . I \ l cdc~r~~*. ;11111 ( ; : l ~ ~ ~ l > ~ n ~ on the impcts of intro- duced invertebrate$ upon the cndcmlc hi or:^ p l r l t . 111 a senrc of urgency (see Howarth and 'rqedcirm, thl\ \t~l.llnc,. 'I hc col~~crv;~t;on hiulu:y of native Species will proceed with \ t ~ l l more hl~nd >pot\ and poter~tid prtfalls unless further reseirch is supported

RECOMMENDATIONS

A number of actions a r e needed t o remedy the problems cited above:

1. Secure sufficient long-term funding to complete a thorough survey of na- tive terrestrial invertebrates of the Hawaiian Islands. Thls endeavor might take a full-time team of 12 systematists at least a decade.

2. Hi re systematists and house them a t the B.P. Bishop Museum and the Universitv of Hawaii to carrv out Recommendation 1.

3. F.nco.lraie and iorter grad~t:lte students to wt rk on the systematics and ccologv of 11;t\i3i'i's i n w r t c h r ~ t e resource\.

1. C u n i ~ l e r e :In : l w x m w n t of lhe con,crvatlon status of native terrestrial . .. . . . . . .. . -. . . .

5. Accelerate research o n conservation biology and interdependencies of terrestrial invertebrates with the remainder of the native btota.

, \ l t l t~~ugh exiitin* d ~ w w c . r i e s and research of the past have hrou ht ~ ~ ~ t e r n ~ t i o n : t l attention to the remarkable evolutionafy a n d ecologlca f! impr~rt :~ncc oi our i n d u invertehrates, understandings of the basic biology. cmlogv. and espcci;~lly ystematics of the Hawaiian arthropods, a r e just hcpinning. 'l'hcrc i s :I cri~icnl need t o support much expanded work In all these areah :ind 111 intetpret the importance of t h ~ s work through educators to decision-nukers a n d the general public.

Important References

Blackhurn, T. 1877. Insect-noles from the Sandwich Isles. Enlonmiogical Monrlfly Magazine 13:227-228.

Blackhurn, T. 1881. Hawaiian entomology. Hawoiion Annuol for 1882: 58-61. Bryan, E.H. 1935. Huwuiian Nahm Noles. Star-Bulletin Publ. Co., Honolulu. Bryan, E.H. 1940. Insecls We See in Hawaii. Tongg Publ. Co., Honolulu. Carson, H.L. 1987. Thc pruecss whcrcby spccies originate. Bioscicncr 37(10):715-720. Carson. H.L.. and K.Y. Kaneshiro. 1976. Dm.~ophila of Hawaii: syslematics and . .

ecolonieal eenctics. A?vtuolRoiew of ~co;on, and ~vsremorici 7511-345. - ~ - ~ - ~~ -~ ~ ~ - Fabridus, J.C. l m - l m 4 . Ctfontologico Sysfemarico . . . . Hafniac (Copenhagen). 4 vols.

(References to Hawaiian insaclr in vol. 2, p. 269 and vol. 3, p. 463.) FullamyY D.T., and N.L.H. Krauss. 1945. Common l&ecf.r of Hawaii. Tongg Publ. Co.,

Honoltdu. GagnC, W.C. 1975. HawaiS's tragic dismemberment. Defendem 50961-469. Game. W.C.. and C.C. Christensen. 1985. Conservation status of nalive lcrrcstrial .. .

invrnchralcs in Hawai'i. Pp. 105.126 IN C.P. Stone and J . M . Scott (ed5 ), llmm r'r Ternrial Ecorys~em! Presm~ofion ondMonopna,r. Univ Hawaii P r m fw i.'c.i Hawaii Cooperative National Park Rcsourccs Studics Unil. Hono!th

GagnC. W.C.. and F.G. Howarlh. 1985 Conscwation status olendcmic i lan:<~! .~n I .ptdc,pl< r a

Pmceedings, 3rd Conpen on Ettropeon Lrpidoplcr(~. 1982, pp. 74.85. (.arnI,riJy;. I- n h ~ l tioff, M.L. 1987. A CorologofAcon o/rlte Hawuiton lsland~. llniv. Ilawaii C'ull~c~: r I

Tropical Agriculture Human Rcsowcv Extension Senice No 075, ilon~dult,. Gulick, A. 1932. Evolrtrio~mf and Mtsrronary John Tltomar Qd;ck. Ilniv. (:hiclf:o Prrs Hadlicld, M.G. 19% Exiinclion tn Hawaiian achalincllinc mails. Molocolupio 2767-XI. - Hardy, D.E. 1%0-1981. Insem ofHowaii. Vols. 10-14 Wiptera). Univ. Hawaii Press,

Honolulu.

Native Temslrial h~wrIebru1eer\GngnC 81

Har1,A.L. 1578. The onslaught against Hawaii's true snails. Nontra1Hi.m~ 67(10):46-57.

Trends in Ecolow and Evoh,Iinn 2(7):220-223 .- . . Horvarlh, F.G., and A C Medriros. [Thk vnl~mwl Non-nalivc inrrr1chr;~lr5. Illinpmrth. J.F. 1923. Ea* Rpfmnrer lo llawaiio~~ btlomololy. Hi\hc,p \ l u w m

Bullelin 2. Bishop Muscurn Press, Ilonululu. Kay. EA 1972 Hawaiian natural hktory I77X-IWI. Pp.flWfiS3 IN FA. Kay ( t J.)..4

Naluml Hdslor). oflhe Hawiian Idon&: Selerrrrrl R~udinm. Univ. Press H a ~ 1 1 . Honolulu.

Manning. A. 19%. The Sandwich Islands Committee, Bishop Museum, and R.C.L. Perkins: Cooperative zoological exploration and publication. OccnsionolPopen of 11te emi ice Pmrohi Bisl8op Mueunz 26:l-46

Monlgorncry. S L. 1M3 Carnivorous calcrpillm the br.haviur, hin~mgraphy and convrvalion of Et:llpirhecio (Lrptdoptern (irurnctridnr) in the llswaiiitn Island\ Geojowzd 7549556.

Moore, N.W., and W.C. Gagnt. 1982. fifegoI~1@m poci/ic~un (McLachlan) - a preliminary study of the coosewation requirements of an endangered species. Repm ofthe Odorran Specialist Gmtp 3. International Union for the Conservation of Nature and Natural Resources Gland. Switzerland.

Mueller-Dombois, D., K.W. Bridgeq and H.L. Carson (eds.). 1981. IslondEco~s~e~ns: Biologic111 0,pmiurrion in Selecred Hmvoiian Communities. Hutchinsan Ross PuM. GI. Stmudsbur& Pennsyhnia

Sharp, D. (ed.). 18991913. Forrao Howoiiensiq Beirq lhe Land Fauna of IhcHmuaiian Is10,tds. 3 vols. The Uni\.ersity Press, Cambridge, England.

Simon, C.M. 1987. Hawaiian evolutionary biology: an intraduaion. Trm& in Ecology and Evohrtion 2(7):175-178.

Simon, C.M., W.C. Gagni, F.G. Howarlh, and FJ. Radavsky. 1984. Hawai'i: a natural entomological laboratorj. Bull&, E~lomdogic~11 Socier). ofAmerica 30(3):&17.

Stal, C. 1859. Hemiptcra. INCA. Virgin, Voyopa~lorvdu MondeswlnFregoe Suedoise I%ugmie . . . lhl(1-1853. Zoolo,qk. PA. Nontedt et Fils, Stockholm.

Suman, T.W. 1964. Spiders of the Hawaiian Islands: catalog and bibliography. Pacific Insect.? 6:6L5-687.

Publ. 44. Bishoo Museum Press. ~&olulu.

Switzerland. Whilehead, P.J. 1978. A guide to the dispersal of zoological material from Captain Cook's

voyages. Pacific Snldirs 253.93.

Association, Honolulu. Williams, F.X. 1936-1944. Biological sludies in Hawaiian water-loving insects. Part 1.

Coleoptera or bcetles. Parl 11. Odonata or dragonflies. Part 111. Diplera or flies Part IV. Le~ ido~te ra or moths and butterflies. Parl V. Hemiptera or bugs.