Polar Biol (1999) 2l: S4 iig i.l Springcr Verlag 1999

New records of Acari from the sub-Antarctic Prince Edward lslands

Accepledi lll Julv I998

Abstract Sixtv spccics of Acari are rccorded tiom thcsub-Antarctic Marion and Prince Edrvard lslands (thePrince Edward archipelago). Trventy of the 45 spcciescollcctcd on recent expedjtions are new and currentl).unclescribed. C)thcr neq' taxa includc a farr.rily of Meso-stismata. firur new gencra, ancl the first sub-Antarcticrecords of CilUbidae (Mesostigmatar) and Ertngioltus(Prostigmala). Fifteen ol'the 31 species prcviouslv re-ported flom thc islands are confinncd. although cight ofthe previous accoLlnts rcmitill doubtl'ul. The fauna.*hich shou,s a distinction bctwccn the shorelinc andterrestr ial componcnts. comprises endcmic. South IndianoLc.rn Pri,\in(c und suh-{nturetie mrte .pc..ies.

I 1.,

D.J. Marshall N.J.NI.B.M. OConnor ' P.J.A.E.A. IJeckermann

Gremmen L. CoetzeePugh P.I). 'Iheron

lntroduction

\larion lsland (46'54'S. 37'45'E)Eduard Island (46'60'S. ,16'97'E)

nerrbels of the South lndian Occan Provincc (sensnLcwis-Smith l9E4). Marion lsland is a mdor studycentrc lor the physiology. ecology ancl biogcography ofmarine birds and mantmals. as rvcll as terrestrial sub-Antarctic plants and invertebrates (e.g. Grcmnren l98l:Craffbrd et al. 19E6; Smith 1987: Lutjcharms 1995). Theinvcrtcbrate launa includes wcll-documentcd macro-arthropods. that is insects (Crafford et al. 1986: C--hou,nand Scholtz 1989: Chou,n 1989. 1990. 1993. 1994:Chorvn et al. 1997). although mesoarthropods. mitcs(Acari) and springtails (Collembola) have attractedrelativelv little attention.

Studies on thc Prince Edward Islands acarolaunahave concentrated on thc taxononty and phvsiology ofthc Cryptostigmilta (Or ibatida: e.g. van Plctzen and Kokl97l: Engelbrecht 1914. 1915: Marshall and Chout1995: Marshall 1996). There have been no broad-scalelield studies ol this group. As a consequence rhc rcportedacarine diversity of 20 Cr),ptostigmata. fivc Prosrigmi.rta(Actincilida). two Astigmata (Acaridida) and no fi-ee-living Mcsostigmata (Garnasida). is ve ry poor comparcdwith other sub-Antarctic islands (Block 1992: Pugh1993: Starlyl and Block 1995: Starlf et al. 1997; Stari' andBlock 1998). The current study rcports on thc Acaricollected bv rcccnt expeditions to Marion and PrinceEdward lslands and comments on thc acarofauna of theislands in light of the new findings.

Materials and methods

Free-li\ing Acari u'crc collected lion] Malion and Prince EdrvardIslands during April Mayol l996and 1997. Collections \rcrc nadefrorr a divcrse rlnge of localitics and habitat types. These includeclvcge!ated (mirc rnd non-rnire) and non-vegerarcd (l'ellliekl) habitats o\er a large alliludinal range. es lr'all rs lhe littoral and supralittoral zorrcs. Slriclly controlled access to Pdncc l:dward Islandli nitr.l .

'r.plrnc rhr'r. r,',r 'ln!lc d.r\ \ .lThc fc$ Acari obserred in the lield. especially rhose jn epilirhic

habllals. lverc collectetl r i1h an aspirator. The nra joritv of sarnples.comprising r,egelation iLnd soil cores- \rcrc hand sorted or hcirlexlractcd. Ibr 5 da1s. r'ia Tullgren or high gradicnt flnnels into

and nearbl Princcare sub-Anlarctic

D J \'larshall (X)L)iprr(menl of Zoology. lJnirersitl of Durban-Westr ille.Pn\ltc Blg X5,1001. Durban:1000. SouIh Afuica-' nail: nrarshall rr pirie.uclrr'.ac.za. Fax: +27-ll-10:11790

\.-l \1. GrcmmcntsLrcir!L Drla-Anallsc Ecologic. D$'arsdrift 2.1.-i\1 .\P Dlcver 'the Netherland\

L Coctzce\rlionirl MuseuIn. Bloenrlontcin. Soulh Afnca

B \1. O(lonnor\ll.cum ol Zoolog). lJnivcrsilv of Michigan.\::r \rbor. NII US,{

P -l \ Pugh3:::r.h \nlarctic Survey. High Cross. C:Lmbridge CBI oET. tJK

I D I heron,7 . '.. D(f,rltnent. Pot( 1cl.tt,,,,tn I trircr,rtr.I :.::.l.rroom. Soulh Afflca

a \ [ ..k.rnann3 :...i:mirtics Di\ision. Phnl Protec!ion Ilescarch Insrilu!e.?':: :, S.ulh

^lricl

709/o cthanol (e.g. MacFaclyen 1961)- Hypersaline llotarion (Pughand Bartsch l99l) \\'as used to exlracl mites from inlerlldal algaeend thc surface of stones. The latter were scrLlbbcd into a bucket ofsea\laler and any suspcnded malerial retained by a 60 pm n],lonmesh wds extracted using hypersaline.

All Acari $ere stored jn 7070 aqueous elhanol with 59/" vrvglvcerol. macerated in 70% lactic acid and mounled for identili-cation in either lactic acid. Hoyer's medium or gll,cerine jelly {e.g.Krantz 1978). Specimens are currentlv in the iDstitutc collcclions ofthe authors and collaboralors as follows: Mesostjgmate (E.A-Ueckermann)- non-marinc Prostigmata (1,.D. Thcron). Oribatida(L. Coetzee). Astignala (B.M. OConnor) and halacarid and hva-desiid mites. L Brrlsch and P.J.A. Pugh.

Results and discussion

Acarine taxa (Tables l, 2)

The revised faunal list for the Prince Edward Islandscomprises 60 free-living acarine species and subspecies.including eight Mesostigmata, 20 Prostigmata, 23Cryptostigmata and nine Astigmata. Filteen of the 31previously reported species are confirmed, althougheight remain unsubstantiated (cf. van Pletzen and Kok1971: Bartsch 1979; Pugh 1993). A lurther 30 specics arerepofied as ncw to the Prince Edward Islands but arenot described here. Ncw taxa include one family (Me-sostigmata). four genera (Mesostigmata and Prostig-mata) and 20 species. The latter include the first sub-Antarctic records of Cillibidae (Mesostigmata) andEr ttt gi opus (Prostigmala).

85

The new genus of the l'amily Cillibidae is closely re-lated to the genus CillibaYon Heyden, known only lromthe Mediterrirnean region. and, Austrulo<illiba Athias-Binche and Bloszyk, liom Australia. lt differs from bothof these gcnera in the shape of the peritrcmes, the an-terior extension o[ thc genital shield. possession of a

claw on the tarsus of leg l. the absence of a metapodalline, submarginal setae on the opisthogaster. the lengthol the corniculi, and the shape of the hl sctae. Thelhmily Rhodacaridac is represented here by five newspecies. ol'which three appear to be closely related to thespecies of Athia.sella Lee. previously only known fron.rNew Zcaland and Australia. The new lamily ol Mcso-stignlata is currently being investigated.

Of the nine new species of Prostigmata. four are lromthe family Nanorchestidae. Nanot(he.\te.\ spccics I isclearly distinguishable from the others on account ol'itslarge size and thc characteristics of the prodorsal area.Species 2 resembles N. cupensis and N. globosas (bothdescribed from the southern Afiican subcontinent) andN. bellus. but warrants separation on the basis of the legchaetotaxy and structural diflerences in the prodosalarea and cheliceral sctae. Whereas species J shows sint-ilarities to N. antarcticu.t. it diffcrs in the detail of thegnathosoma and prodorsal area. Nunor(h(stes species 4hJS an inlcgumcntal lold in the.en.ory lrea senarllinEit lrom both N. antdrcti(us and species 3, and differsfrom the latter in setal lengths.

The new genus of Halacaridae collected from thesupply rcscrvoir (van den Boogaard river) represents the

Famih Species Record Distrihulionfabl€ I Distribulions of\lcsostigmata and Prostigmatarecorded from the PrinceEdrald Islands. All new taxarte in ball. Sub-antarctic.lNtriburions are iaken fro'nBertsch (1979). Pugh (1993) andSrrr! and Block ()998). (Mar\frritimeAntarciic SorrhShetland. South Orkney andi.ruth Sandwich Islands. SouthGeorgiir. Bouveto],a. S1P Soulhlndian Ocean Province Islands

Pince Edward Islands. IslesCrozet and Kerguelen. Heardl.hnd. SPP South PacificOaean Province Islands\lecquarie, Campbell Islands,':. r ne\\ to the Prince Edwardiilands. + conlirmed prior::cord. ? doubtful record)

MesostigmataRhodrceridae

DigamasellidaeCillibidacFam. nov.Prostigmat:rNanorchestid^e

EupodidaeRhagidiidaeHalacaridae

EreynetidaeTydeidaeBdellidaeScLltlcrridaeStign1aeidaeErythraeidae

gen. nor.gen. nor.gen. nor.gen. nor.g€n. nov.

sp. nor. Isp. no\. 2sp. nov. 3sp. nov. 4sp. nov. 5

Dendrolaelaps sp. not.g€n. nov. sp, nov.gen. nov. sp. nov.

lcrrdrcreJldj sp. nov. INanorchetles sp, !no\. 2,\'an arclestas sp. nov, 3Nanorchestes sp. not, 4tulo/c.r r?in rrlr (Strandtmann)Rha!1itlia sp.gen. nor. sp, nov.lletIh?l!a !eta BarlschH alac drc I [us par il iti Bartsch11. 7?or r (Lohmann)H. rohutns LohmannLohnanne lla gauss i LohmnnnIso hac tr us m u gnus (Lohmann)Rhomhognathu.t d .r/e/ BaftschEr elne I es macqud t ie ns i s F ainTldeus ( Pertldeus) sp. not.Bdellodes sp. not.Di sparipe s an t ar c t itus RichtersEtfngiopas sp. nor ,

Balaustium sp. nor.

.l

Nerv

Neu

NcwNew+

Mdr SPPSIP

SPPMar

Mar

MarMar

SIPSIPSIPSIPSIPSIP

SIP

SIP

SPP

86

Table 2 Distribution of Crlptoslrgrnrla and Astignlrta recorded liom the Prince Edward lslands

FJ rrl il) Specics Rccord Disrribution

Cryptostigmatatsrdcht-chthonidr.

HcrnrannicllidaeGymnotlamaeidaePeloppiid:reOppiidae

Oribatulidae

ProtoribalidaeCerlltozclidac

ParakalummidaeAstigmirtaAcxridecAlgophagidae

Hyldesiidne

Wintcrsch idtiidac

Hislioslonllllidnc

Li a t h I h ot t i u ! .1lJt r//r.\ Co\ nrrubiasL. rrlli./rr:r!i,rlrr (l I.lmmcr)

JI u( quat i o tti a !t-ld.r (Wlll$ ork)Jll.rr-r]1rll.r .ro:.1.rr/l (Richlers)Al us k Lt--a t c t a t tr c t it ur nrlt rrrlr1nr.r ( Michael iA. hout(!o.tu ri.\ van Pletzen & Kok//a1r)-., ,r l?/gi.r/c (Michirel)H. ./d--.1./r.r/r (Richtcrs)H. .lrafttT.\i\ \an Pletzen & Kok11. lirhrli Englcbrechtll. rrrr'l)ll/r (Lohmrnn)H. nkxi ts l(rilli(tsi EnglebrechtIJ. t)tut iontn.\i\ EnglebrechtPodacar u s ouhu t i CrirndjcL,nl)t)nvtatinu rlrlidr./?rir ran Ple/ten & Kok7,.r'got ihd ld suhd tu,-./i( rr \'en Plezten & KokTott)bat$ nutiotrn\is van Pletzen & KokA n t u ( I o -( I t!.\ . ro:.idrrri ( Richtcrs)('!/./1r-1,1.r gdrlrii (Richtcrs)Ilagt:IIo:(t(t antut L Iiar (Michael)P ot a k o I u nltnt r.rrrrrl./ (W!ll\\,ork)

S(h\ithtt tul?u .ath!utat(ti( u FilillAlgolhagus sp. trov. IAlgophagus sp. no\.2H tv lts i 0 h d ! o I h i le F i\;11

11. lcrgat,/t,rrr,rrvir LohmannH. tuhunturtrit t FainNtntlnlia t17r.i FrinNeocaholia sp. not.,1u:;trtruoctut ktrgucIt kr?Jt.r Fain

Ne$l'l'l+++

+'l'I

'l++++

+

'l

Ne\\+

+

NcwNewNe$

I

Ncw

Mar'

Mxr

Mar

Mrr

stP

MarMarMarMrr

SPPSPPSPP

SPPSPP

SPP

SPP

SIPSIPSIPSIPSIPSIP

SIPSIPSIPSIPSIPSIP

SIPstP

Mar SIPSIP SPP

SIP

SIPSIPSIPSIP

SIP

first sub-Antarctic freshwater halacarid mite (1. Bartschpersonal comnunication: cL Pugh and Dartnall 1994).All four previously reported Hllarc;rridae from the is-lands u,erc collected offshorc (Baltsch 1979). whilc allthrcc ncw records. ol Halucurallus nolas (Lohmann).IsohttLtrtr.s rlaslru/.r (Lohlran\) i\t\d RhonlhognathLts.rr.rtcl Bartsch. are intcrtidal.

The cryptostigmatid fauna contains numerous dis-crepancics between the current and earlier rccords (cf.van Pletzen and Kok l97l; Pugh 1993). Five taxa re-ported by van Pletzen and Kok (1971) remain uncon-firmed. These are Herruun iell.t sp. (Hermanniellidac).Allod0nretrs sp. (Gymnodameidae) Halo:etes uo.ete sis(Richtcrs) ard H. ed\t qrdel1.tl.! van Plctzen and Kok(Ameronothridi,re) and Afi tur( to:?te.t (r?:ctu',?sr.! (Rich-ters) (Mycobatidae). Diagnoses ol Hemrunniella andAlkttlaneus arc based on single badly preserved speci-mcns that have not subsequently bccn recordedfrom the sub-Antarctic. H. (to:etensi.\ is probablyreferable to H. .filvus (Engelbrecht I975), the onlyabundant terrestrial Hulo.et?s species (Tables 3, 5),whtle H. edts artler.ll! is probably H. ntttrinus devilliersi(Engelblecht 1914). Antorcto:e1er is a synonym ofAlt'k:orihates (Balogh and Balogh 1992). which rnaybe confused wilh Cetqto:ate,\ g.tusii (Richters)(Ceratozetidae) (Wallwork 1972). Furthermore, Pugh

(1993) wrongly placcd C. gnir.rli in the Prince EdwardIslands instead of the Iles Crozet. although a relatedspecics. Mcgcllo;cles utt{u'(i(u.\ (Michael). is a newrecord lor Princc Fdurrrd I.land.

Further uncorroborated rccords are as follows.Lioththonius finrbt'iqtis:jimus (sensu Trav6 1976a in Pugh1993) rvas probably incorrectly identified by van Pletzenand Kok (1971) and is probabll' Z. ctt.rtlalls, a species

recorded during the present study and which occurs onother sub-Antarctic islands. In contrast L. .fimb-riatissin'rus is only known from New Zealand and SouthAmerica (Hammer 1958: Luxton 1985). H. marinus(sensu Trav6 1976a. in Pugh 1993) could, like 11. erl-

vardensi.s. refer to 11. nrurirtus devilliersi. A. antarcticus(Trav6 1976a, in Pugh 1993). is unsubstantiatcd, al-though this could, like A. bouvetoruensis (sensu vanPletzen and Kok 1971; Engelbrccht 1975). be relerablelo A. qntdrcticLts intermedius (sensu Somme 1986).

The Astigmata include three new records of Hyde-sin. and new species of Algophagus and Neoculvoliu(Table 2). Both Algophagus spp. havc the small dorsalsclerites of ,4. semicollatls Fain which are not present ineithet A.u.a lur(tica.r Hughes or A.q. laticollati,\ Fa:n.Unlikc ,4. semicollaris, both have a well-devclopcd ax-illary organ extending between thc bases of legs I and II.They differ between themsclves in body size, the devel-

opment of spines on legs and axillary organ morpho-logy. The new Neotulyoliu sp. is similar to N. kergue-lenex.rls Fain" liorl which it differs in thc structure of themale genitalia and the relative positions of solcnidia I 2on tarsus L

Ecology (Tables 3 5)

There are distinct terrestrial and shoreline faunas lorMarion Island (Tables 3,4). The terrestrial species havegeneral habitat requirements. a l'cature typical of sub-Antarctic island acarofaunas (e.g. Trave 1976b: Pughand Bartsch 1993; Pugh and MacAlister 1994; Pugh1995). However, the shoreline species. particularlv lit-toral species and excluding the predacious Mesostigmatairnd Prostigmata interlopers. such as Balcurstfunt. Btlel-lodes antl Rhodacaridae spp., are more spccific withregald to habitat requirements (Tables 3,4). Some ter-restrial species occur in cdaphic, epilithic and vegetativehabitats. whereas the shoreline specics are olten re-stricted to a particular littoral zone. lichen or alga (Ta-bles 3. 4). For example. lor Halozetes, the terrestrial 11.

/irlyas is found in 20 habitatisites on the islands. whereasits marine counterparts, H. belgicue- H. nurionensi.s andH. narinus deyilliersi, show restrictive distributions(Tablcs 3 5). These differences in niche specificity maybe explained by historical climatic events. The ice-cap-ping of Marion Island that occurrcd during glaciationmust have obliterated most of the tcrrestrial faunatChown 19U9. 1990) while having a lesser effect on the

117

shoreline fauna. Recolonization of the terrestrial cnvi-ronmcnt under low interspecific competition would havelacilitated niche expansion.

Biogeography

Tables I and 2 would suggest that all Mesostigmata.5ll0/o ol'Prostigmata. l3ol, of Clryptostigmata and 33%of Astigmata are endemic. Endcmism of these taxa mayrelatc to differences in their dispersal capacities in thesub-Antarctic which. in turn. may be associated withdiet. energetics and lile histories. The least endemicCryptostigmata have low-energy dicts (fungivorous, al-givorous andior detritivorous) and low metabolic rates.A number ol thc Mcsostigmata. Prostigmata and As-tigmata taxa are predacious or parasitic (more correctlyinvertebratc haematophages). or have phoretic lifestages. The phoretic representatives of these groups aremore likcly the products ol huma.n-mediated introduc-tion (Pugh 1997). Thcse would include the new Cillib-idae. a member of thc phoretic Uropodina andDendrolaelaps sp. (Digamascllid.re). Buluustium sp.(Erythraeidae) is a probable insect parasite (as juve-niles), while most Winterschmidtiidae (though not yl;tconfirmed for Naotulwlia) and possibly Austrqnoetuskerguelenetsi.s Fain (Histiostomatidac) are phoretic.

A number ol species, particularly of the Cryptostig-mata and Astigmlrta. occur on other Southern Oceanislands (Tables l. 2: cf. Pugh 1993: Stary and Block1995: Stari et al. 1997). Of thesc nrite groups. the nu-

Ordcrrspccies SitersubstralaTable 3 Terrestrial Acari of\laiorl lsland: sampling sitcsJnd substrata. 1,.{ D AzorellaKop {400 m): A Poa tor*iirussock. -8 l--orella plant. aBlxck lrvr rocks. l) Ditti.hunlmoss)cushioni t l KattedraalKrans (800 m): -E rocksrlichcn.I DittiLhunl (moss) cushiotl:6 l1 Stony Ridge: G Dlr,n,rxrTrrmoss) cushion. H L):tolodiuDl:/ J Skua Ridgc (100 tn): ,a soil.l. rl.,ra drainage linc,./ soil..lqtotis stuloniJard; K Ttypotr.)il and plant. Poa rcokii; L\rchs'ay Bay: soil and plant.(.)rrla: M N Log Beach: ,I1-t.,JJr lrrr (moss) cushion. NSrereatauhn (lichen): O Ju-nior's Kop: Dir-i(rr, (moss):ushion: P Van den BoogaardRi\ cr D{m: vcgctation at dam:dge]

MesostigmataRhodacaridae sp. nov. 2

sp. nov- 5

Dcndrolaclaps sp. nov.Cillibidae sp. nov.Prostigmatar\d,loral€,ri.,J sp. nov. I

sp. no\'. 2sp. nov. lsp. nov. 4

Eupotles ninunsRhugilia sp.Halacaridae sp. nov.EreJ trcles nwcq ariensi.tn d?r.r (Perlr'/eu.r) sp. nov.Bdellotles sp. nov.Balaus!iun sp. no\.CryptostigmataLioc h I h oiliue austa Ii sMacquat iopllia striutaA sttappia trcaetcnsi.t

Podacarus auhettiDonrcIo, ina nturio (.nsi\AstigmataSc hf i e h?a I a [pa tu b.n tar (t ical/grp,lragrrs sp. nor'. I

Ntoralvtliu trayei

tJHI J

ABDE F

F

AB

oo

GGGGG K

KI J KL

FDF

J KLJ

JKMJ K L MN PJ KLJ MN P

FGGH

F GHB

AB

c

D

DE

IIII

I

I JKKL

(; H

Order species Sitetsubstrata

rJ8

Table 4 Littoral rnd suprrlittoral Acari of N,larion Island: sam-pling siles dnd subslrala. (.1 F Macaroni Br): .! L/d.r/.)./Lr. su-pralittoral lichen. B Cuhplu(u. supralitloral )rchen. C I'errucuriu.upper-shore ltchen. D Porph.rnt. mid-shorc alga. t red alga fronllow-shorc. I pool in supralittoral zone: G H Transvaal CoVe: GPorph ru. md shorc al€!:r. l/ rcd xlgd liom lor sho|e; 1 J Ship s

Core: / beach debris near pcngLlin ncst. .J Mastulict- supftlllttorallichen)

merous species shareil with iles Crozet (ll). iles Ker-guclcn (14) and Heard Island (5) in particular, supportthe ploposal of a South lndian Ocean Province (sensuLewis-Snith l9lJ4). The l5 spccics which also occur onthe Maritimc Antarctic lslands (including 7 on SouthGeorgia). and the 11 on the South Pacific Islands (in-cluding 7 on Macquarie) corroborate a sub-Antarcticelement of the Prince Edward lslands acarofauna (sensue.g. Gressitt I967).

The Prince Edward Islands are geologicallv young atczr 2 milljon ycars old. and so were most likely colonised,during the Holocene, by lrans-occanic waifs (sensuPeake l98l; Kuschcl l99l). Most of these waifs origi-nated on other sub-Antarctic islands and the numbers ofcommon spccies suggcst that the Mesozoic continentalioceanic microplates of the Iles Crozet and Kcrguclcn.both of which contained Pleistocene glacial refugia(Chown 1994). are the most likell''. sources of the currentPrincc Edrvard .[slands acaroftruna.

Ackno$ledgements I B:rrtscb (Biologrsche Anstalt Helgoland.Halnburg. Germany). GCI Loots (Zoolog! Dcpxrtmenl. Universityol Polchefslroom. South Aftica) and L Croblcr (National Muse-um. Bloemlbntein. South Africa) are lhanked for conlirmingi<lentillcations. R Mcrcer helped with the lield r,"ork and collectedAlatko:tttt antur(titu\ int.ruk,dius. SL Chorvn (Uni\'ersit], ol'Pretoria. South Africe) commcntcd on.r draf! nanuscl-ipt. TheSouth Alrican l)eprrtrnent of EDvironmenl Affairs and Tourismprovided logistical and financial support through the Soulh AlricanComrnittee for Anlirrclic Research. I'JA Pugh \\rs supportedthrough Ihc British Antirrctic Surre\.

References

, P.(,g1. J. B.rl.'gl. P ' l'rr:' lhe.'nb:rtrd rnl(\ pc.lcr,r oi rhr \\,,rld.Ilrrnirr:rrr \ut:,.rr:.J \4u.(u r Pr(-. BuJ.rf(.1

Bartsch I (1979) Halacariclae (Acnri) eus der Subantarktis. CahBiol Mar 20:325 319

Block W (1992) An annotrted bibliographl of Antarctic inverre,brales (terrcstrirl and licsh\\rtcr). British Antarctic Surve!.Cambridge

Chown SL (19E9) Ilabitat use and diet as biogeogrdphic indicatorslor subantarctic Ectemnorhinini (Coleoptera: Curculionidae).Antdrcl Sci l:2J 30

Chown SL ( 1990) Possible efi'ccts of Quatcrnary climatc change onthe conposition of insect comnunilies of the South lidianOcean Province Islands. S Afr J Sci 86:186 19l

Chown SL (1993) Desiccation resisrance in six sub-Antarctic wee-vils (Coleopterar Curculionidae): humidity as an abjotic factotinlluencing assemblage struciure. Funct Ecol 7:318 125

Chown SL (199.1) Historical ecology of the sub-Antarctic weevils(Coleopterar Curculionidae): patterns and processes on isolatedislands. J Nat Hist 28:4ll ,131

Chown SL. Scholtz CH (1989) Biology and ecology ofruyrrre.e/e.tJcanncl (Colcoptcra. Curculionidac) spccics complcx on Mari-on Island. Oecologia 80:93 99

Chorvn SL. \.ln de. Merwe M. Snlilh VR (1997) The influence ofhabitat and altitude on oxygen uptake in sub-Antarctic weevils.Physiol Zool 70:ll6 124

Crrllord JE. Schohz CH, Chonn SL (i986) The insects ofsub-Antarctic Marion and Princc Edward Islands: with a

biblioglaphy oI entornology of the Kerguelen BiogeographicalProvince. S Afr J Antarct Res 22:51 58

1,E{elbre.hr ( \'l {lq'4) Thc genus Hal"--are.' rOribdtei Acdrilon' Marion lsl.nd N.rror' NJ' Mu. tBlocrnfontrinr l.l 25

l\{€sostigmataRhodacrridac sp. no\'. I

sp. nov. 3

sp. nov 4sp. no\" 5

ProstigmataHtltarrellu,s novttsLtohattru.s ntagnutRhonthogmrhus uu.:tcrBrlc//odcs sp. nov.t-rrr.{iryrlr sp. no\'.Bdfurslirrr? sp. no\'.CryptostigmataA Ia.\ko:tIcs dnIar(Iit: .\

Htlo:ates helgiLut,IL nturinus loilitrsi

Podacarus aubcrtiTa I o ba I L,s n t u t i otk,n si \Porokuhrntntu ratunduAstigmataAlgophagus sp. nor. 2

Htadtsiu htrlophilaH. kt rgutlcncnsi.;

,4 u s t r u nl us k t r gue I e ntn t i :

AB IJ

H

(iH

(; tl

IJ

CHGG

II

EC

FBBts

B

AB

I.IJ

'I able 5 Terresllal Acal.l ofPrince Edlr'ird Island: san1pling silesand subslrata. (.,1 FCa\c Ba):,,1 P,d!ir[r mat on old Pod sIool. BCrrlo2|rrr'a on black lenal rocks- C A:otcllu ar:tl Sd!<ino. D A(ucndclrainage line. Z'open /i/r,rlurrrar'br'lophyte \egelalion. F dr') urirevegetalion: 6 / (;oldcn (;rtc: 6 lichcns ol shehered rcrticxl fllce.l/ lichens and rocks- I Blethnrnt Plagi.rll.? mire; J Kraterkoppie:J uplancl ileldmalk ol scoriae)

Order specics

NlesostigmataLtolroluelaps sp. novCillibidae sp. no\.Fam. no\'. sp. nov.Prostigmatafr.futlr ( P{,rtr ../.r/.\ ) sp. no!.Brlc//orle.r sp. nor'.1ir-r',rglorrr.r sp. nov.Baldustiutn sp. tlo\.CryptostigmataMacqttutia )ia stti dAusttullia (to:(Iensis

Poda(atus tufuttiD on1? I a I i nd nktr i one n si s

ToIob(Iet ntat iatk tisM a gt I lo:e t e.s an tar( I i( usAstigmataS(hfiehea t.r[pd suhantdt.Iic4.1 [ gophugu r sp. (unidentifi cd)NcocaArlia sp. nov.

DDF F

B EFDF

BR

CAC

CAC

FGHI J

EFG IEFGHI J

FHEFG I J

GCH

E

FG

lzrlitlbrccht ( M ( lq-5, \($ anreronolhrokl rOrihaler' A(arii la\d' Ir.'m lhe Reptrblre .rt Soulh Africh antl rhc r.hnd' ol Gouglr

.nd Mrrion- Navors Nas Mus (Bloemfbntein) 3153 88

Gremmcn NJM (19131) The vegetalion of the Subantarctic islands

Marion and Prince Edward. Dr w Junk. The Hague

Gressitt JL (1967) Introduction to entomology of ,Anlarctica

-Anlnrc Re\ Ser l0:t 1.1

(rmnrer \4 tlq58, In\(\tl!allun\ on thc orrhatid laLrnJ of th(" And.s Mountains. L The Argcntine end Bolivia Biol Skr Vid

Selsk l0: l 129

Kranlz GW (l97ti),4 manual of Acarology, 2nd edn. C)regon Statc

University Bookstorcs. CorvallisKuschel C (1991) Biogeographic aspccls of the subantarclic is-

lands. Atti Accad Na7 Lincei Rc 85:578 591

Lewis-Smith RI (1984) Terrestrial plant biology of the sub-Ant-

arctic and Antarctic ln: Lalvs RM (ed) Antarctjc ecology

Academic. London. PP 61 162

Lutieharms JRII (1995) bibliography ofthc South Aliican Journal,)l AntarcLrc Re'(,rrch. S Alr J AnlJril Re. 25:51 84

Ilion \4 1tol51 Crlpl$lr!mriJ iArrchnrdJ: A(arrr a cnnLI\e- ,.r,.*. Fauna NZ 7:l l0o

\+ir6T-acjven A (lq6l, lrnpro\eLl lunnel t)fc e\lraclnr' for \oll/ ,rrhron,,d'. J Anirn fc.'l .l0.l7l 184

, )+dfi'r ll i).1 (luq6t Cornpornll\e r\aler rclation\ ol .uh-Anlarclrc/ ,nd continenlal ArrLaritrc oribatrJ nrile' Polal Biol lo::R_ 2ql

\lashall DJ. Choun SL (lqq5J Temnerrlur( effecl' un locomolor.- - ,,.ri!it\ rilc\ ot rub-Anlar(ltc oribalrd mil(. Polar Brol

l5:'17 49Peakc JF (19u1) Thc land snails of islands a dispersalist s view-

o.rinr ln CrccnuU,'d PH. Foft! PL {eds' ( hrnce chrnge 3nd

.h.rllenpc. rhc <vnlring brosphtre Brili5h \4u-cum rNirlur3l

}li.tor], irnJ Crmbridgc I nirer.rt) ftes'. pp l'1- 261

e/"n lL+ 1toot) ^

-\nonlmic cirlalogLrc ol the Acrri from Anl-!' _rrctica.

the sub-Antarctic islands and the Soulhern Ocean

J Nal Hist 2?:321-'121Pu.eh PJA (1995) Air-breathing litlortrl mites of sub-Antarctic

South Gcorgia. J Zool Lond 236:649 666

l{9

Pugh PJA (1997) Acarine colonisation of Antarctica and thc is-

lands of the Southern Ocean: thc role ol zoohoria. Polar Rec

3l:113 122

Pugh PJA, Bartsch I (1993) Ecology of the littoral Halacaridae(Acari: Prostigmata) of South Georgia in the sub-Antarctic.I Nat Hist 28:75-85

Pugh I'JA. Dartnall HJG ( 1994) Thc Acari of fresh- and brackishwater habilats in the Antarctic and sub-Antarclic regions PolarBiol 14:401 404

. Ptrch PJA. MrcAli'lcr H 1loa41 Ac.rri of lhc supr.llirlurrl /one 'n- -ruh-{nt.rrcttc Suulh Ceorgid. Pedobr.rl.-'gia .t8:552 )05

Smith VR (198?) The environmcnt and biota of Marion IslandS Ali J Sci 8l:2ll 220

Somme L (19{36) Terrcstrial arthropods of Bouvetoya Polar Res(N S.) 4:l l5 l2q

r,Sf.rrj J. Blo.k \ (lq')5,Oribarid mrrcs rAcarr: Oribatidar olS.ruth' C.orp',. South Arlanlic. J \Hl Hi.l 29:146q l4ql

Stari J. tlock W (1998) Distribution and biogcographv of oribatidmites (Acari: Oribatida) in Antarctica. the sub-Antarctic islandsrnJ neirrb\ land rrr<.r. J \dl Hitt ll:8ol 894

, stlrv J. Block w. (,reen\lade P tlqq'l onb,rtid mrles iAcirri:r o''hrr'd.,r of rhe sub-AnlJrcli( Heard l.land. J Nrr Hi'r

3l:545 553Travi (1976a) Les oribaies (acariens) des archipel de Kerguelen ct

Crozet: essai biogcographique. CNFRA 4l:61 70

. trr\ i ll,r-bbj Rccherche'.ur le. Microiillhropnde- lerrcllre- J(r I rrchipel dc. Kcrguelen donnec. quantrtaltre' anal)'e d( deu\group;s d Acarierls Oribatida et Acaridida Rev Ecol Biol Sol

ll:5s -67

' V,ad Pl.,r.n R. Kok DJ tlgTll orrhdlei In ran Zinderen Bakkcr- f\4, Drle' RA. Wrnlerbottom J\4 (eJ'r \4Jrion .rnd Princ(

Edward Islands: Report on the South African biological andgeological expedition. 1965 l966 Balkema. Cape To\\'n.

nf ll+126. Mit"",L JA (lq-2, Sonre crlJ'to\ligmJlrd mrle\ rAecr,: Crlplo-- .lignrrtr) from Crozet l.land. Pac In'e(lq l4::? 1-