BIOLOGY OF HALOBATES (HETEROPTERA: GERRIDAE)

8/3/2019 BIOLOGY OF HALOBATES (HETEROPTERA: GERRIDAE)

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Ann. Rev Entomol Copyrigh b Annua Reews Inc. A rgs eserved

BIOLOGY OF HOBATES

(HETEROPTERA: GERRDAE)

ChengScipps Instiutio of Oceaography, La Jolla Cafoa 92093

TDUT

Until abou10 yeas ago aloba was pobably known o only a handful ofenomologits; ouside hi egment of e enomological wold i wa geneallyunknown. Howeve dung he las 10 yeas hee has been a gea deal ofeseach on vious aspects of the biology of this unique insec genus, has

now become a well-nown oganism among maine biologists ionicallyhough it i sill no well-known amng entomologis

Insecs ae he mos common of he mezoans but i as genealy thoughtha hey ae aben fom the oceans which cove some 7% of he eah'ssuface. We now know hat insecs in at leas 4 odes wih ome 1400secies occu in vaious maine habias 26), alhough in the open seas we indonly membes o he genus aloba. These sea saes o ocean stides aein he Family Geidae of he Ode emipea and wee dscoveed some 60yeas ago by Eschscho e nclded hee species n hs fs descponof he genu bu aid nohing about thei biology Beween 8 and 188,when BuchananWhie decibed he aloba collected dung he Cha Exedon few papes wee publshe on hs genus. he 8 n 88listed species including 6 ne ones and povided a ey o all descibedspecies. Fom hen unil 961 a numbe of papes wee published many weedesciptions of ne ecies hat wee synonymied ae bu some conanedaddiional biological infomation By fa he most complee aonomic eamen ofaloba was the eview by Heing 5) who eamined all availabletypes descbed pio o

960synonymied many species and descbed

4new ones. e povided a ey o all 8 pecies map showing he type localityo disbution of each of he non-endemic species an annoaed bibliogaphyof ealie publicaion on he genu and an account of some paial life histoy

0066-70/85/00-0$0200

Annu.

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2 CN

ue ha he cre u gehe wh ne fm he leaue weehee nec emane me le gne by blg f he n ecaeunil nee wa ee by Cheng 17). Since hen alobas ha becme

much bete nwn mane blg an ha been he ubjec f a ey fbehaal blgcal n bchemcal ue N aemp mae hee eew he leaue n alobas p 9 nce can be fun in heeale eew f Heing 5) an Cheng 7). nea ecen ning n hiunue genu f inec e eewe wh anal infman fm unpublhe pena bean

AD P

hee e n nwn fl falobas he famly Geiae ae bac alea he lgcene 85) M membe f he famly e pn ae whchcmmnly cu n he uface f fehwae eam pn An ecelleneew f he w Geae wa publhe n 960 by Maua 60) an meecen ue n he phylgenec elanhp an gegaphy hae beencie u b alabee an Aneen aloba nee be ne f he me aance membe f he family. Alhugh he majiy fhe Geae ae fehwae inhaban hee ae a few genea bee alobas ha hae epeenae lng n mne ennmen e.. humao

bas Aslos obas humaomos 5). f hee nlyAslosbelng he ame ubfamily a aloba an i i he m lely anceafm 5) Heng ppe ha aloba cul hae gine fm anslos- ance in uhea Aa nce hee f he fu nwn pecefAslos e fun n ha ea 6 5) uheme a lea 50% f henwn caal aloba pecie ccu n ha ea (ee gue 1) which iepecally ich n nly n Geae bu al n he auac Heepea )

cumanal eence fm a ecen uy n he chmme falobas anAslos 6) len upp eng hyphe eample inall ehwae ge thu fa negae he chme nube angefm 2 (ee Newman Cheng un slos shrau hae 2 chmme bu n alobas pecie the numbe wa hehghe ece f any Geiae. Snce hee nec hae hlcencchmme i n uneanable pulae ha beaage f uchchmme may hae gien ie the inceae chmme numbefun in he me aance pecie 6.

MPnle fehwae pn-e whch may be ehe wnge wngle allmne alobas e pemanenly wngle Wng whch ae peumablyueful f peal mgan beween icninuu habia wul be f

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Y F HALBATES

itte use in te sea. Te body o a sea skater is typicaly sort and road, and ine adul age may reac a maxmum leng o 65 mm and may be abou mm in widt Te antennae ae long and sender and are never modiied Te

ont legs ae so and used or grasping prey wile eeding or o olding teemae durig matig tey e ot used or locomotio, wic is done pincipally y te long middle legs. e tiiae and tarsal segments o te middle legsare ined by ong ydouge airs, wic may ep in reducing riction betweente legs and water suface tereby alowing tese insects to skate moreeortlessly over te water surace () Te ind legs wic are muc sorterta te midde legs, re used more for steeng tan or propusion. Te indtbae and sa segments are also lned wit ydrouge airs e ostrum s4-segmented and reativey sort, not extending beyond te prosteum

Te nymps ae vey smlar to adus in exeal morpology, and ere isno exteal sexual dimorpism unti te last nympal insta dult malesgenerally longer and tinner tan emales, ave te last two or tree abdominalsegmets modiied to o te geitaia Otewise tere e o stucturaldiferences etween te sexes Maes o dierent species can be istinguisedy dierences in genitalia emales, owever, are vey similar and diicut todentiy Te abdominal tergum o most emales is exible lowing orexpansion during egg maturation emales tat contain matured eggs may avea markedly umped abdomen (4)

ST

Tere ave been o complete lie istor studies on any species oaloba.Some obsevations ave een made on a ew coastal and open-ocean species,and it is becoming clear tat tere re severa consistent dierences etween tetwo groups.

Te eggs o coasta aloba re laid on substrates near sore Wenbrougt to te laboratory and kept in an aquarium aloba avvnr romPalau, West Caroline Isands, aid eggs gued to te container wa, 9 to 9 mmbelow te we level, n wo dsnc dep goups one at 52 mm and teoter at 5 mm below te water suace Tere were no eggs at 9 mm or2 mm deps. 59 eggs ound, 2 wee laid singy, 6 clusers were o2 to eggs and custe ad 6 eggs, pobably a laid y a single female Gravid avvn emales dissected at Palau were ound to contain rom to uly matured, oive-green eggs (L. Ceng, unpubised data) Newly laid eggs

are also olve green, bu tey tu brigt orange in 2 to days As te embyodeveops, te eyes become crimson, and dark grey pigmentation outlining teantennae, legs, rostrum, and te ead and body segments ca be seen cearytoug te semitansparet coo Prior to atcig te eggs re d greyexcept or te eyes, wic remain rigt orange or crimson n Paau, were te


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4 NG

seawater temperature in June 1981 was29 to 30C, the duration between eggaying andhatching was 17 days

In the open ocean the eggs e laid on loating sbstates: feathes small

pieces of wod, mollsk shells, seeds, ta lmps, nsectcacae c have found two fourth-nstar nymphs of an from the Atantc Ocean, bearng eggs on the dorsa surfaces The deveopment of the embosobrnu is simiar to that avvnralthogh in a batch of

obueggs coected nea La Paz Baja Caifoia in Novemer 974 andstdied in the laboatoy, development took mch longe. In that samplesveal hndd eggs had been ld ll ove a t lup appoxmately 5 cacross; n some paces they were two or even three eggayers deep. he eggshatched (up to 50 per day) sporadcay over a period of30 days, suggesting that

embyo devlopment might take p to one month at 20C. Inbu 90%of thedeveloping embyos the vental sface faced pwads, with the cimson eyesnd the nealy folded antnnae and legs clealy visible thogh the anslcentchoron. Some 1% o the eggs were slghtly tued to one sde wth ony oneeye vsbe from the top When the embryo was ready to hatch, the abdomencontracted to times at approximately 2sec itervals A slit was then madewith the chiinized egg bse and he head emeged foowed y he abdomen The legs wee then plled ot leaving the embyonic skin behind The enti pocess took abot min

Egg hatch was folowed at hr ntevas for days Less than % hatcheddurng each of the perods of hr and12-18 hr The majority (51 %) hatchedbetween 4 and 10 hr and about20% hatched between20 and 22 hr ince newly hatched nymphs e moe ssceptible to pedation the timing of egghatch might be an ipotant facto in edcing nymphal motality

Dspite diligent seches caied ot in the eld on many occasions I havenot found ay eggs on coasta rocks or on submerged mangrove branches or roots his isnepected since neashoe aloba presumaby ovpost on

such substraeswly hatched nymphs ofaloba mease abot 1 mm in body lengthThe cticles, initially ple, become tanned within an ho o two nd theyong insects ae able to move abot in a eky fashion soon ae hatchingAttempts to eed newly hatched nymphs of obrnu nthe laboatoy have not been successfu; n one rearng study they a ded within ve days ofhatching Theyong nymphs occasionally preyed on one nother but they didnot feed on ny mine organim o on fehy edDroophla ffed to thm althogh olde nymphs fom thid insta onwads) nd alts eadily

tk sch od oganisms How st insta nymphs svive in he open ocan isstill a mystery hey may be able to utze ntents fom the oganic-ch surface m but thshas not been substanated

Deveopment and moting of third toh instar nymph have been observed


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IGY F HALBATES 115

in mnnum (24) Ahouh i a no po o oo any nnyph hough h dvopna piod y ong and xaning ho i a po o a ha ah nypha adu ad 1214 day.

hi i o h ovaion o ing (5) on . hen: 8 dayoh ai na 1012 da or ler nsrs He oo a una o oohe nyphs hrogh compee cyce

n my aoaoy popuaon o . mnnum on ourrd onyn00 and 00 h oy o 0800 Indvidua ady o ooppd dng and appd uggh. Ny od nyph vy payo o ao hi u annin gan ao diay Whn onhou mos nsecs hadtued gh rown, and whn hreehou hy hadaand noa ooaion So on o o hou a h o hy gan o

dAuin ha ah nypha adiu ak 114 day h oa dvop

men perod rom eggs oadu a a o 60 o 70 day Th opon ao o oni and nypha dvopn i poay pau a n h a o vaou ha id udid (15 77), ahouh hav no daa or lobte Oher normon on and nymphdeveopmen ws pshedy Heng (5 and evmr (74

AXM

h axono olobte ws y revewed y Herrng n hs 191 (monogrph hry-egh spees were nded n hs ey, whh s presenh ony oka ky o h gnu. Sin h puaon x n pihav n addd: tne o h By o Bna (54 . e o Buni(61 . melleu rom h dn os o he Red e (59) . mngovenromheGu o Aaa and heGu o Zuu on h Rd Sa (70) heonto Dain Auaa (66) and heng o Noh Qunand Auaia (67

e1 ss henownlobte speces ogeher wh her rod rnesodiuion or ype oes rig (53) ognzd o a goup olobte h opn-oan and h naho o oaa oup h onud vn p. Hov n ehholtz and tne renown ony rom her ype ocons hese wo spees re excded romrher onsderons o openocenlobe n hs revew

heai xonom chrersdnguhng lote p a u o h a gniaa h pi dn n oo paand odaon on any o h appnda Whn ony a aavae enghso vros ppendges my e sed xonomunan genery remn Fo h oan p hov h aaaoy okn ky vn i ony a a avaia (22). Nyph an

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116 CNG

Ls f aoba sps dsrbd pr 198 wh hr kw dsb

Sps K dsb

auaudi Brgrh Syhs; Mah I as I2 H bwn Hg Koomaaa oomo3. H. ba Hrrg Fj Mak 4. caypu Hrrg hpps Nw Ga

5 i Heng Astralia6 ehhzi Hrg" Zazbar

7 jiei Hrg F 8 avver shsz Indin Oan; a Oa9. formidbi Ds Adama a ay ga

10 H aat Hg Aaba Sa

mau Whi" Indian Ocan; Pacfic Ocan haaiiei Ug Hawa Maqa y Tah Ta

3 haan Wh Rd Sa Nw Ga Idsa MaaysaAsaa

14 aonu Esak Japa

15 kaheriae Hg Nw Cada6 keei Hn Sama

17 kudri Nasav Nw Cada

8 mauu Shadw S Nw Ga19 maraarum sak Maaas Gam Car Marsa I20 maumrai sak ap Fmsa Kra Cha

21. mica shshz' Aa a da a a a22 mbei H Ausrala

erei Hg aa Nw Ga24. ae Hg Nw Cada

25. eron Hrrg oomo ' Nw Ga hpps26 poed Hrrg Kya

27 picep Wh Cbs Maaysa Nw Ga a Masava

8 proau Wh Mas Nw Hbds ava hpps S-m Maaysa Nba

ea Cap s Sas

30 uu Bab Gapag ad

aae Hrg Tona32 rce Eshsh af a33 exuai Dsa Maasa b Wh rh an Cra a35. ede Wa Sh Amr

h rrg Mars37. ae Hg Ba f Bga

8 hieeei Sk Asaa

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(d

ps

ehrs rg4 ngrvensis mdd M

4 elles 42 eskii Mymoo43. henis m

d g

4 hingi m

a p p

IG HABATES

Ard

ow dsrbo

d d)

Dw, Ar

Qd Ar

gnrally b idniid only in assoiaion wi adls T dvlpmnalsags o nymps an b assignd by omparing sgmn lngs o varosappendags. Tabls nympal masremens ar availabl only or .mn robutu and . mrnnrum (24 3

Al lobte spcies ave been nd n marne abia ecp r heront wc wa rcnl dcbd m al R n NTrrir Asralia Howr alog spimns o is spciswr colld svral klmrs psam rom rvr m no saliniymasrmns wr akn) O 44 spis now known only 5 liv on on oan ors ar knwn slly rom yp loalis r nar oassNarsor spcis lobte sm o prr oasal aras ringd wimangrv wr ma ccr n dn reacng m and can beal cag w a pnd n a plac ranr T adl a ally nd

n e mre pn ara away m e mangre wera nympspally s irs w nsar ar mr r lss nnd o sallwwars among ros wr y mig b mr prod rom prdaors Ia en eremly dnse pacs yng nymp amng aral roo mangrovs in (. Jen am and onap (. mrnnrum and ealpagos (. robutu. In oasal aras wo mangrovs yng nympsar n nd cle erangng rances r rocks g. . multu in Solomons and nere in ala wil adls and ldr nymps ply mrpen waer

Nearsr lobte are rarly nd alng xpd ca e.g n Rranga n e Ck Iland nl mange r caal gain pn. Ir ar slands w mang r r gan m adl nscsma wandr m m d alng mr psd scn ang cass

Annu.




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18 CHENG

e.g mg he rk sds Pu. Breedg us ear be mresey ssed w veeed s bs udi bys s diver mus

Te ve ee lbte sees e usuy u by wi eus e ver e e sufe () Tey re sedm fud se sreexe er srms hve fud udreds f si iv semes f gemnu srded m wrk M sd Fij fw srewd s. Aer sees, . mn ws fud uder smr dis he Cym sds he Cbbe d he sd f Me he TukLg Care sds (wese Pf).

Augh e e ea sure ers be bi wiu ysbudares lobte eds ere hes. Phys egrh

eies my im r e uees f e ses i sues e disussed i he ex sei

Diibuin

y c f 9 c ss (udg ehholt ad tne) re sw Fure 1 Te re mry f hesees e ud he dMysi d Ausrsi reg whi srbby e eer f eui f e geus sige s sees sbee fud he A Oe Ahugh my f he s sees ekw y frm eir rg ye s (see Tbe 1) we sfeyude hey re uy edem se se skers my ur arudreme isds sedm vised by mie biiss d eve mre rey byemiss (3, 3, 3). Cei seies fud oud isds es frwh here s sderbe eg frm my deed be edem (eg. robuu he ags sds. A few he s ses re erwdey dsbued e.g vventr hynu mrnnrum bu eh s

fed me we-esbshed gegrh re where ud esy vesred frm sd isd by fw e ures. Te dsbue f. hen s s esy exed. Ths sees s beerered frm he Hwi sds sued arud N d s fm gu f isds Tii e.) ed ud 2S see Tbe 1). Seimesfrm Hw d Th re aey dsiusbe wud be eresg fd u wheer ey ud ierbreed uy r eiy sie esd rus e bu 5 k ar d here re w kw esufe ue sysems e hem. Te us f. hen

my be rems f mre wdey dsrbued sees s he se f erbe sees (W A ewm ers mmu.

The dsbus f he ve e lobe sees are w we esbsed ( 7 4) Tey e ri r subo wers werehe wier emerure des f mu bew 15C Fiur 2 Teir


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BOLOG OF HALAES 9

Figu ITy loaton o oaal

btspp ncdn

. tznd

. t).Ss ns gvn n Tb 1.

ourn nd undn on ny vn t ppnty onod yu tr tprtu (0 2) h optu tptr n or t P p 22C (2).

Hobs mns th ony oopotn p ond n thr oron (u 2). In th P nd Indn on t rng nyonnd to on tn out 20 nd 20S ut t xpnd to 0 to 0Sn th Ant On. In t On th p y tndnohd yond 20 tod Jpn, poy n rd th y thKoho rrnt th ar oth tnon n th t IndnOn ttut to th Anuh urnt ( 2 h unu dtrutono th p n th orth and South Atant pry o d yuf urnt yt. Th p o o vny dtutd n thnoth ton ut ot opty nt o th utop SouthAtnt (2) h noth xtnon poy nund y onn o u t (57)

h oth or p of on ktr a fond n h P On utth hv v dnt duton tt ( ur 2 H sus th

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N

u Rnges o dsbon o he ve mjo oen spp

mo idepread ih a bipolar diribion raher like hoe of cerain zooplankon oranim ih plankonic larvae in he aer colmn (9 68 I iraer na or a race-den eonc oranm o o c adiconino diribion The abence of seeus beeen abo and2S eablihed by an abndance of neaive daa; 42) may be de ocompeiion ih mans Alhoh pecimen of boh pecie may occroeher in ome locaion here i no overlap in heir rane in area here

he ocr in hih deniie Fire 3 2) sobnus and splendens areraher more rericed in heir diribion and he end o be more coaalhan mans or seeus

The eainin openocean pecie, gemanus i fond boh in hePacific and Indian ocean here like splendens and sobnus i eemo ollo he coa of he coninen and iland Fir 2). Drn hcircmnaiaional cie of "peraion Drae 3) gemau a coleced a eeral aion. I poplaion deniie ( expreed b nmber oinec cah per andard 0-min o 7) ere mch hiher in he Java Seaand Red Sea (repecie 2 and 3) han in he aciic Ocean proper

(6) b no pecmen a cah in he open rech o he ndian cean ()

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10'

o

ZO

BOLOY OF BTES 121

" . . :

Figure Aras of high population desities of hree Ha/bates spp i he ease ropcal PacifcOa (42)

mila obeatio hae bee epoed b aadach (69) ho pecthat gemnu may have diferet food referece from . mn hichcold accot for it omehat more coatal habitat

Food

ea ate ae predaceo he have tpcal hemptera c mothpahere the madiblar tylet have toothed ed ed for ctt ad themaillar tylet ae held toethe by fie ride to fom the feed tbe 0.Whe the iect ae ot feedi the tylet are retracted ito the hot otrm.he diet ofHlobte i vaied. I eeral earhore pecie feed o ladiect tapped at the ater face hich ca be rather abdat ohore9 30 bt opeocea pecie feed o vario zooplantoic oraim 0a el a o oati iect aibaim ha bee oberved both i e fieldad i the laborator amo pelaic a el a earhore ea ater. dlt

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NG

and older nmph ma feed on ouner nmph bu adul have never beeneen o feed on one anoher In he Gapao Foer & Trehee 4 fondha 5% of he pre of rbutu e pecmen of her on knd. have

never obered or been abe o nduce an lbt o dve eher or food orfor anhn ee Foer & Trehee (4 found he ame a e for H.rbutu. ea kaer probabl deec food eher b h (dead pre) or benor deecon of rpple creaed b pre trln a he aer urface, aha been obered n frehaer errd b Murphe ( Ahoh havefond ha brnu and ru prefer vn, ln fruit le ommobe one Foer & Trehee (4 repored ha coaal rbutuavoded n pre. f he pre ma enera raped b he eakaer fron e and held compleel free from conac h he aer race

(Fure 4 and 5 premabl o ha here are no ripple from a rln preo arac aenon from oher poenal predaor When he prey are andoo heav o uch a a mal h have een hree o fou ea kaerfeedn mulaneol on he ame anma Oher nfonaon on feednbehavor can be found n he paper b Foer & Trehee (4 Feedn meobvou depend on he relave ze of he pre and he conumer a honfor he frehaer Grr (5. A laboraoryreared fr rphl mlngtr can be ucked "dr b an adu lbt n le han mn In mexperience one or o ie per da ce for each adul mrnnrum

The feedn procee n lbt are premabl mar o hoe of oherfrehaer errid (14 here a deve enme ecreed and pmped nohe pre' bod o ldze he mcle and oher bod conen The qd food hen cked p hroh he food canal.

Predators

urfacefeedn eabrd are he mo mporan predaor of pea lbtpp , eman of. mn have been fond n of he brded et nthtu) n he echele (4 hen npblhed daa).Alhoh ru a fond n he reurae of 9 o of 18 brd peceded n he Norhee aaan Iand 5 th food em mporann he de of onl for pece: he blera nodd Prlt rul) heBonn perel Ptrrm hplu) he rabacked e t lunt)and Buler' pere Bulr bulr) The blera nodd and he phoenxpee Ptrrm lb) had been earler repored o ake conderabe nmber of ocean kaer (0 b n hoe ude ampe e ere oo malfor anave aemen In a more recen d, hen & arron 5

fond u n 15 ou of (4.4% rerae from he for majorbrd predao ed above For he mo mpoa redaor, Prltrul ome 81% of he amped brd had fed on lbt and he averaenmber of nec per food ample a Th dmnve brd


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BIOLOGY OF HWBATES 123

igure 4 Halobes sericeus fm dn n d-wd t y_

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124 N

Fgu Top view of living Haba iu female feeding on a de-winge it ly.

eih o abo 5 m ca a imes feed soe o H alobates ad o dobs a ver mpora predaor o sea saers he ope ocea [The eercoe ofHfavivetris ads as fod o be kca dr ( he,C S arrso ad T N e pbshed daa a vae smr o ha of hepa bPhilaaeus(Jassidae) ad ihi he ae ve fo sevea ivere

baes (6 6)] Ahoh aod he Haaa isads he coasa speciesH hawaiiesis ca ofe be fod i appecabe mbes (35) e did o fidspecmes of hs speces a of he 15 brd somach sampes e examed() e do o ko heher peac brds aso feed o coasaH abates spp he Gapaos sads pedao o H rbsts b he eo arber(Dedrica peachia), sma fsh (Sardp agax ad a speces of mead o oe occasio a ava izad (Troiurus albemarlesis) has beeobseved (46 8)

F-Chain RelationshipsHalobaes spp. hehe coasa or peac beo o he marie pesocomm (21). Ahoh aimas hs odmesioa evme arecoed o he sea srface he ieac ih oasms ha eea ive


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GY F HABAS 125

aoe o elo them. elagc oce ate fee o zooplato tappe at theea face. ea hee poplatio eitie of iect ae high (p to X1 km 42 the mt emove a coieale me of pe ogaim

oeve, ice ocea kate ae fifeee it ha ot et pove poile tot thei iet examiig thei gt cotet o to etimate the impact ofocea kate o the poplatio of a ooplakto ogaim. vetall ehol e ale to evelop immological (4 o electophoetic (51) methoitale fo ch aale Coatal albates pecie poal fee lgel ola iect that have ee lo o ahe ot to ea o have falle to theate face fom ovehagig vegetatio like thei opeoce coi,the have ot ee oeve to take mae ogaim. albates ma fall peto h o eai a thee cotte i ome a to the ecclig of

tiet i the maie eviomet, t it ill e ma e efoe e haveeogh qatitative ata fo etimatig thei tophic elatiohip i te pleo (21

Cetai ogaic compo a heav metal ae ko to e cocetateat the eaface film, t thei coetatio i the fil ae techicallifficlt to meae (52) albates pecie, hich live at the ea-ai itefacei the ope ocea, have ee fo to accmlate ome of the heav metal.Sice the pemal otai ch metal eithe thogh ioaccmlatio pthe foo chai o ikig facefilm ate, the col coceival e

e a iicato of heav-metal polltio i the ope ocea. t ma eigificat that ioaccmlatio of ome of the heav metal, a of chloiate hocao, a fo ol i pelagic albates ot i ea-hoegei.

o example, Cheg & Bema (8) fo that total DDT cocetatioi sbrius ee 51 time highe tha i pecime of the e-hoeRheumatbates aestuarius collecte fom the viciit The highe DDT cocetato albates a attite to a iet of me zooplakto ogaim, hich e themelve peato Rheumatbates o the othe ha,

feeig mail o teetial iect, ma of hich ma e phtophago,ol e le likel to cocetate DDT

The cotet of heav metal, ch a camim (C), coppe (C), io (e),lea (P) , icel (Ni), a zic (Z), have ee meae i eveal pecie ofoth pelagic a ea-hoe albates [e.g. Cheg et al (27; Cheg et al(4] geeal, the tace-elemet cotet i ea kate e ithi the ageo lightl highe tha thoe epoe fo ote mae iveeate (7oeve, the C cocetatio te to e mch highe, a facto of omoe, tha te highet vale otaie fo ooplakto ample. Sice C itoxic to hma a othe aimal (49 a ha ee exeivel ivetigate ie mae eviomet, e pe tie o the ptake a accmlatio ofti tace elemet i captive ea kate (7 7 it cocetatio i atal

Annu.




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6 NG

poplatio (4 7 4 7) ad it poible traer to the avia pedator othee mie iect ( 4) We oud that o the whole coatal lobecotaied le tha 5 ppm dr wt) o Cd i their bodie wherea opeocea

pecie ma cotai p to 00 ppm Althogh thi elemet ca be take p romther pre much o the Cd oud i ocea kater i probabl obtaid rom therace lm b drkig (4) Some o the dietar Cd ma be ecreted or toredi te cticle a ractio dotedl remai i he tie Wat eect it mahave o the biolog or phiolog o ocea kater ha ot et bee ivetigated. Cheg et (4) have oud that the bluegra odd (rocelceru[e) the mot importat avia predator o rcu ha proportioatelmore Cd i it eather tha do three other related bird rom the ame habitatggetig that thi mtal ca be aerred rom ocea kater p the ood

chai to it redatorBhavior

lobe are trogl attracted to light at ight. How thi tpe o phototacticbehavior evolved i ot ea to udertad the ope ea there are oatrall occurrg lightemittig leto orgaim that might ee a oodNeverthele i area where ea kater occr but ma be diicult to catch idatime becaue o their exreme agilit it i ote ea to capture them buig a light lure ater dark. I coatal area where iherme practice ight

ihig their light ote attract large umber o lad iect a well a varioplatoic orgaim ad ih providig a rch ood ouce or lob Seakater have well developed ee which the ue or hutig ad i avoidig orecapig rom predator. The ca ee well ot ol i daligh but alo obright moolit ight a evideced b etavoidace tdie bae o et catchdata (18 2. Sice it i almot impoible to carr out behavioral tudie opelagic lob i the ield ad we have ot et bee abe to povde uitablelaborator coditio or reag opeocea pecie iormatio o theirbehavior i limited mail to hoterm obeatio cied out at ea or toidirect dedctio baed o populatio aale o preerved ample

However i everal locatio aroud the world high deitie olobepopulatio e kow to occur ea hore where expeietal tude ca becared out with relative eae Several iteretig ield tudie have bee carriedout at the Gapago lad where robuu i oud i great abudace (7 8) ect o thi pecie oe orm dee aggregate cloe to the edge omagrove thcket or coatal lava rock. The patche o highet deit coito matig par o adlt up to 120 m) mixture o adult in o ad older

mph ted to be le dee (8) The leat dee aggregate are compoed omied populatio o matig adult igle adult ad older mph (third toith itar). Youg irt to ecod itar mph almot ever oud i uch


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BOLOY OF HALOBATS 1

aggregates nonay occur in dense patches of ther own age group aongmangrove roots, were they may avoid cannbasm by oder sbings.

Floas o rbusus anain saions in areas wh ngng angrove

alon rocky shores where hey end o anain reaivey consan posiionshoughou da cyces. Were e mangove ees re cose o shore easexposed at ow tde he otas reain aggregated but they move to poos andrean tere unti the tide retus; ten they ove bac to their orignaposons n relaon o the anrove ede. The rerable ability olbesagregatons to ainain or resue saons in hs way is probaby acheved bysoe sor o orientation by he individua nsects to topophysica feaures suchas manrove oos or rocy sore nes (8. The cohesion of such oas sprobaby atibutabe to gregarious beavior. The uncona sgnificance o

aggregatons in rbusus was studied by Foster & Treee (48 whoonsdered ha suh aggregates ay reduce susceptbity o predaors Theyfound tat te approac o a predator or ode woud cause ndvduas a tepeiphey to change ther speed and direction o oveent thereby increasingencounter rates among individuas of the otia and perhaps confusng thepredator and avoding predaton. Fueore te number o encounters wasfound to depend on roup size: avoidance behavior n a arge agregate can beacheved by na visua suaion at ama predaor approach ()Since Treee & Foster (8 observed n he ed ha the transmison o

avodance behavior though the loia greay exceeded e speed of anapproaching predaor ode hey caed i he Traalgar Eect. Ths behaviorenabes the insects to nitiate avodance behavor even beore the approachingpredator s seen. Besides adng predator avodance rbusus aggregatesaso seve as arenas or ae fndng and seecon (8 inge ales osa aggreaes whch aac eaes (and oher ales) as he size of heota increases he popoion o insecs i ul aso ncreases oser &Trehee (48 found H. rbusus pairs reanng i ul for up to 4 .This s consisent wth y own unpubshed obseatons o ieris n

Pau and rru n Guam. Presumaby such prolonged copulaonreduces copetion aong spe o copetng aes (see 65). Matefindng and copuaton may be smary actated by group foation n otherspecies olbes

In the freshwater gerid Rhgdrsus kreelii Wcox (8 84 found

that maes can atact emaes by producing surface rppes or wave signasI Athoug tis sort of system coud be expected to wor we on st surfaces of

ponds or aes reavey poteed fro wnds s dcu o vsuaze such asyste operang eceny on he open ocean ecep duing eceponaycal spes. If peac species of Hlbes depend on hs system or matefnding one woud expect to ind a coelation between teir repoduction and

Annu.




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8 CHENG

doldrm codiions. Howeer analyses of feld collecons so hae noshown ny cler indicion o sesonliy 19 4

Spcial Aapain

s enioned erlier alobates is e only insec ens liin on he openocean; i is also he only marne ineebrae aculy feeding he seaairinerace. ha speci dpaons enable i o occpy such speced ndofen hosie enronmen? nd how does i nerac wh rous bologcaphysic and cheic cors peculir o he seair inerfce? These are someo e probles we e o sole i we re o ndersnd he biooy o hisnusu insec o fr we hae only lmied inormion on soe of eadapaions of pelaic aloba o is special habia snce here are obious

diculies inoled in condcing research on he open ocean. O necessiymos o he relen inormaion hs been obned fro specily preseredspeciens nd ro sudies on cosl species

ie e seir inerce is in soe wys siple, b in ohers i s beremarably compliced. In his essenially wodimensiona eninmen hesur o e nil s predced largely on is aby o mnime accidensubergence and o oid drowning. aloba does his by enclosing isel inwha s anlogous o a waerrepelen enelope or expe he body andes o pelic alobates re coered by lyer of hydroe crorichi in

ddon o he cohir co 6) n H rmas e ooshpedmicrcha re abou 5 m high 0 m wde across he op 05 wide he base nd spaced m par while he aerage nercap disnce isabou Ther densy ws ound o be n he range o67 >< 0 mm

for javii nd x 10 m for H ga n such nrrneen yer o ir rpped on he hirs srroundin he insec iesi buoyncy nd enbles i o pop p o he se surce ie n ir bubble i i isaccdenaly submerged In he freshwaer geid Vi he cha aepeglie 6 ccordin o horpe Crisp ) hooshped microrichiaprde a ch ore effcen plasrn hn pegle ones.

To es he eficiency of he irreanng pasron oaloba Icaed ousoe eperiens he rine lboroy in l Wes Croline Islnds in9 nd 92 usin he cos species . javivetris peciens wereforced o rein sbered in boied sewer (conining 35 pp O) nd innboiled sewer (coninin7.4 pp O). Wiin 5 inues e speciens inhe boiled wer exhbed signs of paralysis wh heir mdde nd hind egsexended bcwrds (ire ) ie alobates hod heir les exed in

crcersic sin posiion (see iure 5) In nboiled sewer prysisse n ony er 30 min ( Cheng npublished d). I speciens wereremoed ro he experimen esses shorly er pryss ws obseved(whn ns) hen dried and reued o he surce o seawer hey soon


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BIOLOG OF HALOBAES 12

rcovrd and wthn a fw mnt bgan to kat normally If howvrnct n bold awat w lft to down thy ank wthn 4 mn f thntkn ot and dd vn thogh thy ht tll b alv thy nv and

no ovnt, a t l had vdntly ffrd prmannt pant.On th othr hand atr bng hld ndr nbold awatr vn aftr a 2hrbmrgnc mot ( ot of 12 pcmn) rmand aoat and appard to balv. Th th mcotcha layr on avveris hld cnt ar to allowth nct to rvv ndr watr for at lat 2 h at 315C whra n boldawatr th oxygn wa qckly dpltd to lthal lvl Indvdal of thocanc seres whch occr n mch coolr watr (23C) wr abl torman bmrgd for p to 1 hr wthot ffrng any apparnt ll ffct (L.Chng npblhd data)

Aprt from bng abl to avod drownng a katr mt b capabl ofmantanng thr body d at an omolty condably blow that of thawat (zng-pont dpron a = 1C). Th omotc pr ofth hmolmh of r wa ond to b 2 m omol -97 a (E Edny and hng npblhd data) whch not vry dfrnt

Fgure 6 view of dead alobate erceu fmale.


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130 EN

ro tat o terestria insects (-0839 0c) (75) Wen speciens werepaced in water containing Y3 or2 tie te osoarity o seawater (0.33 X or x SW or4 , eoyp osotic pressure decreased (to 4358 oso

1) or increased (to 366 oso 1) and te anias beca suggis.Tose pced on res wter or2 X SW died win 48 r Speciens pced onO5x or15x SW owever beaved noray and survived or up to 4 dayse coasta avv was abe to ive on 05 X SW and even on res wateror up to 7 and 5 days, respectivey (L Ceng unpubised data t ay beore adaptabe tan te peagic species since it occasionay as t cope witeavy freswater unoff after stors

bs are agie insects and can sate at speeds o about 50100 csec (20) ey can aso jup to eigts of 112 c (24) Te ydofuge

airs coverng teir egs presuby ep to reduce rctiona drag (1) Teirabiity to jup is a ery eective eans of escaping fro predators coingup fr beow or ying above te. Since ere are no seters of any kind onte open ocean suc agiity is o considerabe suviva vaue

t is reasonabe to assue tat ood or ese oceanic insects is usuay essabundant tan it woud be at te suraces o aes streas or ponds and tattey woud need considerabe energy reseves to tide te over periods ofstavation. Lee & Ceng (58) ound tat two species of ocean saters mau and . u store rigycerides in aounts of up to 92% of teir

tota ipids but in teir bracis-ater(Rhubs) and resater(G) reatives te corresponding vaues were uc ower in te range of4672% Since trigycerides are easiy etaboized tey are presuabyuseu energy reseves or tese insects Atoug tese two ocean saters storeany o te sae stuctura faty acids as are coon in insect ipids tey asocontain te poyunsaturated aty acids 20:4, 20:5 and 22:6, noray oundony in arine zoopanton Tus tese bs store not ony pospoipidsand structura ipids coon in oter insects but aso ipids pecuir to arinezoopanton. is as been sown aso or te nearsore sea sater. fby Hoand et a (55) Soe poyunsaturated atty acids suc as 05 and 6ay be essentia for ebrane foration in arine anias and since insectsare typicy unabe to syntesize ongcain aty cids (above 18 crboncains or eongate te oecuar cains o unsaurated ay acids (50) teywoud ave to obtain te o teir diet as suggested by Hoand et a (55)

n reswater as we as nearsore arine environents gerrs are otenound under sade duing sunny periods a beavior tat presuay eps toprtect te insects ro possibe daage fro soar UV igt. n te openocean owever tere is no sade and ocean skaters ave ad to evove soeind o adaptation orUV potection. n ceain insects soe o te energy ote incident UV ay be refected or reeitted by uorescence but tesepenoena ave not been observed in bs. e fate oUV energy in te

Annu.




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21/25

BIOLOGY OF HALOBATES 11

biologically damaging range (26020 nm) absorbed by geid cuticles wasstue by Cheng et al (1) We foun that, although the UV-absorbespet of the oean skater sus was qualtatvely sml to that of

gerrs ro braksh water (Rhuatobats astuaus) an fresh water G g) at a waelength o 280 nm only 00002% of the raiation isansmie throgh the cuticle oH. u compared to 04% in Rhuatobat an 50% in G. g t is pobable that the relatie opacity oalobats cutcle is attbutable to a hgh content of aomatc amno acs(notably tyrsne, typtophan, an phenylalanne) (6) but ths subjet neesfurther nvestgaton

UD MAS

The uniqueness of alobat presents us with a vaiety of intellectual hallenges here is, irst ts vey exstene nmore oetly, onthe sea nspte of the common isconception that marine nsects o not exist (to uge bytextbooks in entomology and marne biology) there are in act seeral hundredinsect genera assoiated with mane littoral enironments Howeer alobats is the only genus with species that are wholly oceanic Why are there notmore? How does alobats succee where others have failed? How an twithstand aily tropical insolaton wthout a lea r a so to hie uner n the

heat of noon? How can it stalk about on a % solution o salts an neverthelessmaintain a hypotonic blood? It may see rain too rarely; dew an seamlets arequte outse its ken. How does it walk on water? Without any possible shelter,how oes it suvive in stors? How es t n ates? These ae some of thequestions that we hae staed to answer, but there ae others.

The world ofHalobat is the most strctly twodimensional habitat o anynown animal. This habita is a way neace ha setches or thousands okilometers noth and south east and west. However any deviation more than aouple of mllmeters above or below the water sufae must be outse thsnset' s mmeate wol. et vtually all of ts prey must oat up f belowas plankton or a o above as a sparse rain of aerial insets or the habitat ofHalobat is quite without its own pimary sources o prouctiity. And all oits predators likewise except or its cannibalistic cousins, ie down upon it(as do birds like noddies) or swim up o snatch it rom below (as do ceainshes, f they ae quck enough)

A futher question ones subets suh as gene flow, speaton, anevoluton The eology of alobats has zoogeography wthout topology, for

the ocean surace is eatureless uniied by mountain riges, rier alleys, ordese plains How in such circumstances hae ierent species eole, andhow do they maintain their ierences? emperature gradients? Perhaps, butcould they suice? It seems unlikely. ould cuents an gyres be suciently

Annu.




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12 N

isinc an cnsan eng eimi spcies We n nw a as dn') Wa bservans sd be mae an wa expermens devis am p w w p

prbms nmiss an marne bigiss an i i pmes researca may answr sme es qesins in i wi a sr i pps

ACKNOWLEDGMENTS

M sea abae w ave be psb wi agmn, abran, a p rm nmrs rins a a,many wm ar cars pbicains cie in e revew, an gae ei cpeain n paica ms menin e ae PressJn D saacs my sband Rap A Lewin and he S pgraper

Larr Fr Te ancies a ave pe in nin vas researc prjecs ob n Amran Asscain Unvry Wmen, eNaina Grapic Sciy Naina Sence Fnain Fnan Ocean Researc San Deg) Oc Nava Rseac eNaina Ocanic an Amsperic Aminsain e Nana Sea GranPrgram e Scey Sigma Xi and e Scripps nsin Orpy

Leraure Ced

Andrn, N M 76 A compatitudy of locomotion on th watr urfacin miaquatc bug Vdnk dd.Dan. Nauh. Fn. 7-6

Andrn N M 77 Fin ucur ofth body air layr and morphology ofth pirac of miauatic bug (ncta Hmptra Gomopha) in rationto lif o th wat ufac Vdnk.dd Dan Nauh Fn 07-7

Andrn N. M 7 Phylogntic infrnc a applid to th tudy of olu

tiony dirication of miauaticbug (Hmipta: omorpha S00 8:5578

Andrn N M Th miauaticbug (Hmiptra omorpha Phylogny adaptation, biogography andclaifcation Enmngaph Vol Klmpnborg Dnmak Scd Sci pp

Andrn N M Polhmu J T 76Watrtidr (Hmiptra idaVlida tc In an nc d Chng pp 7- Amtrdam

NorthHoland pp6 Ahmol N P Ahmol M J 67Compaat fding cology of a birdof a opial ic ild u Peabd u Na H Ya Un N 24pp

7 Bb C W 6 Th uu Adnu Nw Yor/London P Putnam'Son pp

Birch M C Chng, L Trh J .7 Ditribution and nionmntalynchronization of th mrn inctHaba buu in th alpago and P S ndn S 06

Brnton 7 Paatr rlating toth diibution of platonic organim pcially Euphauiid in th at

opical Pacific Pg Oang

10 Brwn, M hng L 98 New ntfor ampling th oca urfac aE Pg. S -7

Bull K R Muon R K Obo DWard P Chng L 77 High ll ofcadmium in Atantic abird and aatr Nau 607-

Calabr D M 0 Zoogographyand claditic analyi of th ida(Hmpta Htoptra c PubEnm S. Am l -

Calabr D M Wng polymophim and cladogn in th ida (Hmiptra Htroptra n ThEu Sgncance nc Pmphm d M W Stoc A C Bartlt pp


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4 Chng, 966 Studis n th bilgy th Grrda (Hm. , Htroptra). :Obsrvans n h dng Lmnogonusfossau (F) Entomol Mon. Mag

102:21-285 Chng, . 966 Sudis n th bilgy t Grrda (Hm. , Htrptra) IIh li hisr Meocois tenuicoisEski. Entomol. Mon. Mag 122738

Chng, L 973 Mn nd shatskats dincs in suac in strucurs. Natu 242 3233

17 Chng, L 1973 Halobats Oanog.Ma. Bio Ann v 13-35

18 Chng, 973 Can alobates ddgnts? B dalght? imol Oceanog8663-65

9 Chng, 973 Th can-sdalobates (Hoptr Grida) in thAlanic Ocan Oanology 3570(Oiginall publishd in Russan in 973Okanologia 3683-90)

20 Chng, . 974 Nts n h clg th canic insc Halobats Ma Fish 357

2 Chng, n plutnAnimals at th saai intacOanog Ma. Bio. Ann v. 1318-22

22 Chng, 975 nscta Hmpra

Hrotra Grrda gnus Halobatesih Ident Zooplanton 147 Cns Intxpl ), 4 pp

23 Chng, . 980 alobates and thr lttlkn marn inscs h SuhPacic gin n Po Int Symp onMain Biogogaphy and Evolution inthe outh misphe Aucklad,NZ. pp. 5839

24 Chng, . 981 alobates rom icnsia, ith nts n kping maiannaum ali in th labatyMionsia 79706

25 Chng, 98 lobtes om thcasts Cha Oaol imol i13:788 (in Chins)

26 Chng 1982 ach and castal inscts n cclopedia of each adCoasal nonmens d. M .Sch, pp 489 PhiladlphiaHutchnsn Rss 960 pp

27 Chng, . Aladr, G V. rac, I 97 Cadmium and thr havy mtals in saskatrs (Grrda: Halobateshumatobats) Wat Ai Soil Pollut.633-38

8Chng, , idlman, .

1977Chlrinatd hycarbns in marn inscts.

Estuain oastal Ma. Si 5289919 Chng . ch M C 1977 st

ial nscts at sa J Ma Bio AssoU 5799-97

30 Chng , Bih C 1978 sct

BOLOGY O 133

ltsm: an unstudid mrin rsurcEcol Entoml. 38797

3 1 . Chng . Duk , Grng, D. A. 1978 UV absptin b grd cuticls

imnol Oanog3555

32 Chng, . , nrgh, I 1973 Canalobats ddg nts? y mnlight?imol. Oceaog 186669

33 Chng, , ad, C. H 199 Ataxnmic sudy th Malayan Grrda(Hmptra Htrpra) th nts nthi bilgy and dsibutin Oiet st 397-

34 Chng, L, Fac P I Schulzalds, 979 av mtals n a saskatralobates obustus m th Galpagsslands: Cncntrans in naur and up

a pmnts, it spcial nct cadmium Ma. io 420635 Chng Harrisn C S 983 Sabid

prdatn n th saskatr Halobats(Htptra: Grrida) Ma Bio 723039

36 Chng, ., Hill, D. S. 1982 in inscts Hng Kng Po 1st Int Mao. Wokhop: Mae loa adauna ofHong Kong and South hina Hong Kong 1980 pp 7383 HngKng ng Kng Univ rss

37 Chng, , Hlda, 983 alo

bates clctd durng th circuavigatinl pditin Opratn Dk Zool.J in So. 78 8798

38 Chng aild 1980 mphs t saskatrs, Halobats obustusand H mians (Hpa Gda).Syst Entomol 5:437

39 Chng Schmitt D 982 ainnscs th gnra Halbats ademabats (Hrptra) m nustnts arund id Islnd, Grat BairR Aust Ma shwat s33109-2

40 Chng, , Schulals, 98rqunc and populatn cmpositinHalobats mians (Htpta Grda) m t cnal and sut AtlancOca "Meteo oshugsegeb.he D o pp -

4 Chng, L , Schulalds, M. , rsn, C S 984 Cadum in cskatrs, alobats seiceus (Insca),and in thir sabird prdats. Ma Bio75 n prss

42 Chng, , Shulnbrgr E 1980 Distibuin and abundnc Halobats

spcis (nscta: Htptra) in th ast pica aciic ish ull 785799

43 Diamnd A W. 1976 Subannual bding and mult ccls in th brndld tSta anathtus in t Sychlls bis 8449

Annu.R

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134 EN

Eschscholz I. F. 822 nomogapin 1:16- Beln: Remer 186pp (n Gen)

45 Feller, R I. , Taghon, G L GallagerE D. Kenny G E. Iums P A.979 nologcl mehds for fdweb analyss n a soboom benhconiy. Ma. o. 546-74

46 Foser W A Treee I 80Feedng edaon and aggegaon behavor n a mare nsc Haloao Brber (Hempra Gedae) he Gapagos slnds o. R. So.ondon S B 20953953

47 Foser W A Trehee E. 8Evdence for he dluon effec n hesesh hed fm sh edaon on amrne nsec Na 293467

48 Foser W. A. Trhee E. 82eproducve havour of he ceanskaer Haloae o (HemeraGedae) n he Glapagos slnds.Oologa 55:202-7

49 Frberg L. Pscaor M. Nordberg G.97 admm n h nvonmnClevelnd CRC Press. 66 pp

0 Gler, L O'onnor . 9 Lpd nso n ahopods n emal Zoology ed. M Florkn B TScheer 5253 New York Academ 460 .

5 Gller P S 82 The naurl desof waerbugs (HemteraHeeroera):elecrohoress as a otenal mehod ofanalyss. col nomol 723337

52 Hrdy J T. 182 The sea suace mclayer bology emsy nd nhoogenc ehmen og eanog 37-28

53 Herrng L 6 The genus Haloa (emer Gerdae) a ne (2)22

4 Heng, L 64 A new seces ofHaloa m he ay of Bengal onomol So. Wa. 668-86

55 Hollnd L rs D as 983 hanges n he fatty acd omoson of he oean-srder Haloa jen eeoea; Gedae) aesrvaon Ma. Bio. L. 425-65

6 Jmeson G. S. Scudder G G. E.977 Food onsumon n Geemera elga 23-4

57 ohn, H 82 Dsbuon of Haloae mian (eteera Gerrdae nhe nohe Ala dung sng comments on s cahably n neuson

nes "Mo Fohngeg ehe No 34 336

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BIOLOGY OF HALOBATES (HETEROPTERA: GERRIDAE)