l o i i gkr de 1;i ~iiiicc iivec p'<iliciilc 1 3 - I ,SO: piiic ~iiiibiil;it~iire 1: lxisiléiiiur 0.47/0.10-0.I 2, télofdiiiiir ( j ,2~~0,?~/( /o, (~x-(~, 10. t i h i ; i ~)..~?-0:~~/~).07-0.08. h;isii;irsc 0.2R-0,24/0.06. iéloi:irsc 0 . 2 ~ - 0 , 3 3 / ~ ~ , ~ ~ ~ ; p t i c

' a i h l a t o i r c IV: fémur 0,73-(3,76/~1.17-0,I9. tibia 0,64-0.6S/0,~-0,l 1, basitarsc 0.28-0.30/0,07-0,08,

fseudohlorliriu vuicariiis n.sp. se distingue de I'csficc oroitiii n.sp.par des pattes-máchoircs plus élaiicfes, uiic taillc IégPrciiieiii plusgr;iiidc, des yciix un ~xii pliis &vclopl>cs. 1 1 projcciion ;intérolatéralc de la hanchc 1 csi plus épaissc ct ~x~urvuc dc spicules. Outrc ccla une différcncc scniblc cxistcr dans le

, 2 1 0 i ~ ~ e 0.35-0,37/0,06-0,07

13

Figs. 1 I i 13: Pscu~loi~ir>i/iric~ vuicunicc n.sp.; pédipalpc gauchc

déwcloppcincni des tarses des patics iiinbulaioircs: chcz vrilcunur les télotarses 1 ct IV sont rclativcmcnt plus longs que les basitxxs (1.26-1.37 rcsp.1,16-1.31 )que chcz oroniii (1.50-1.64 rcsp. 1,36-1.42). Ccttc diffkrcncc yourrait indiqucr titic séparation gCnétiquc bicn 6i;iblic de ccs dcux esficcs, qui soni issucs. i une époquc asscz récciiic, d'unc sotichc aiiccstrale coiiiinunc Les iles de I'archipcl des Acorcs sont toutcs d'wiginc volc;iiiiquc. Sao Jorgc ct 1Crccir;i font partic dti groupe ccnLral, qui s'cst probablcnicnt formé vcrs 1;i fin du Miocenc el qiii ;i énicrgc diiratii le Pliociinc.

Par la prksence d'uiic glande stcrnalc oroniii (ci probablcniciit aussi vulconus ) se riipproche de P.srrinarii du Jura suisse. jusqu'ici xulc espt?ce connuc avec une iclle glande abdominiilc. Ces cspiccs soni bicn distinctes par dauves caractres morphologiqucs ct niorphométriqucs.

BIBLIOGRAPIiIE

BEIEK. M. 1963. Ordnung Pscudoxorpionidca (Afierskorpione). Bcstiiiiinbüchcr Bodcnfauiia Europas 1: 313 p. -- 1969: Reliktformcn in dcr Pseudoscorpioiiidcn-~auna Europas. Mcmoric Soc.ent.ital. 48:317-

323. MAHNERT.V.sous prcs.sc.l.cs pscudoscoryioiis (Araclinida) des groiles des llcs Canarics. avcc dcscription

de dciix esp5ccs iiouvcllcs du genrc fardioc/if/ioniu.s Ucier. MCiii.Biospéol. 16 OROMI, p. J.L.MAKTIN, N.P.ASIIMOLE 1988. L i s ciividades volcanica5 en las islas Azores. Actas las.

Jornadas Atlanticai Mcio Ainbiciitc.Aiigra do Ilcroism«.Jan 1988. VACIION. M. I 954: Remarqucs morpliologiqucs ct aiiaioiniqucs sur les I'seudoscorpions (Arachnides)

iippanenani au gcnrc Psciidobioilirur (Ucicr) (Fiini. Syarinidic J.C.C.). Uull. Mus. natn. 1-list. nat. Piis. 2c sér..26 (2):212-219.

1969: Hcmanlues sur I;i Liinillc des Syariiiidac J.C.Chambcrlin (Araclinides, I'seudoscorpions) i propos de la dcscripiioii d'iiiic iiouvclle cspkcc: I'sctulobloiliricr rliieúariúi , habitaiit les cavenies de

-- SiiiSSC. I ~ C V I I C Si i isW 5í001.76i: 387-306.

Vicror*a 18: 171-1ii7 Juiiiu IYN

l

l I I I I I 1 I 1 I l 1 1

1

Arthropods of recen1 lava flows o11 Lanzarote 8

ABSTRACT: Trapping aiid searching was used 10 investigate the arthropods oí historic lava and adjaceiit islotes on Lanzarote. Recent lava away íroni the coas t is poor iii iiuinbers oí individuals and oí species. Very close LO ihe coast there are abundünt Collenibola and also a variety oí larger arthropods, especially ílightless Melyridae (Coleop- te ra ) , ílightless Gryllidae (Orthoptera) and lsopoda (Crustacea). Islotes of older rocks have richer communities in which Diptera and Coleoptera a re abundant but Collembola relatively scarce. Key words: Arthropods. lava í low, cave, aeolian ecosystenis, Tinianíaya. Lanzarote, Canary Islands.

RESUMEN: Se utilizó un muestreo por trampeo y búsqueda para investi- gar la fauna d e artrópodos d e las lavas históricas y los islotes adyacentes cn Lanzarotc. La lava rccieitte alejada de la costa es pobre tanto e n número de individuos como de cspecics; sin embargo muy cerca d e la costa hay abundantes colémbolos y una amplia variedad de artrópodos de mayor tainano, espccialiiiente Isopoda (Crustacea) y cier- tos insectos no voladores coino Melyridae (Coleopicra) y Gryllidae (Orthoptera) . Los islotes de rocas iiiás antiguas sustentan comunidades inás ricas en las que abundan coleópteros y díl~teros, niientras que los colémbolos son relativamente escasos. Palabras clave: Artrópodos, colada de lava, cueva, ccosistemas cólicos, Timaníaya. Lanzarole, Islas Canarias.

INTRODLKXION

l 'he Parque Nacional de Timaníaya, oii Lanzarote. Canary Islands, Spain, wa. established priniarily becausc oí tlie spcctacuiar volcaiiic piiciiomena that i t displays 1-lowever, the lava flows and ciiidcr cones oí ihc park also Iiave considerable biulogica intercst. KUNKEL (1981) has discusscd the platit liíc oí the park, and w e rcport Iiert o n somc prelimiiiary iiivestigaiions. i n 1984 aiitl 1985, oí tlie arthropods living in t l i i

park and o n similar arcas ncarby. Wc also includc dais un arthropods lound duritig , brieí investigation oí a lava tube iii historic lava ouisidc the park.

As Fig. 1 shows. i h c eruptions bctwecn 1730 and 1736 (and sinaller ones i i 1824) covered aliiiost cxactly a quartcr of thc islaiid of 1,anzarote (and alniosi i h i whole oí the area that is now ~iiirk) w i t h lava aiid pyroclastic tiiatcrials. klost oí t11t recent lava is oí the chaotic and jagged "aa" typc. but tliere are sonie areas oí ihi snioother "pahoelioe" lava (see MacDONA 1.1). 1953). Tiic volcanic dcposits presuiiiahl!

170 171

sterilizcd the ground conipletely, bu1 soiiie old high volcanoes and some relat ively ílat areas were surrounded by ncw lava or pyroclasis without being coinpletely covered. Thcsc are pcrliaps bcst rcícrrct l to as "kipukas", íol lowii ig thc I iawaiiai i ierniinology (NiUfLLEII-UOMUOIS et al., I9Ml), I)uL c i i i Laiizarotc thcy are kiiowii as "isloies" oí oldcr rock cincrging íroiii tlic surroundiiig rccc i i i "mar dc lava".

Alihougli recci i t lava ílows I x k aiiy typc o! soil and are inliospitablc íor Iiiglicr p la i i ts , colonizatioii by iiiicr<)orgüiiisiiis, licliens, and to soiiie extent iiiosses seenis to be possiblc almosi i i t oii<:c:. ií adcqua ie iiioisturc is ava i lab le (HENII IKSSON si IIODGEHS, 1978). Oii Laiizuroic succcssion is doubtless helped by tlie accumulalion oí dust f ro i i i [he Saliara, wliich probobly arrives at a rate of several centinietres per 1000 years (SCHOTZ ct al.. l!18l). 'rl ie resultant dusi deposiis hold moisture, and are colonized by iiinsses aiid occiisioiislly by higlicr plaiits. Iiideed. KUNKEL (1981) poiiits out thai aliiiost 400 pliiiit spccics (i<:ciir iii thc piirk. Apart froiii licliens, howcvcr, tlicse plaiits are exireiiicly scattcred. ~i i id coii i r i l iuie little LO the productivity oí the area. L ic l iens niust acliicvc substaiiiial priii iary productioii where tliey are abundani, but populations oí Iicrbivorcs iii miics wi i l i licliens secm LO be vcry sparse.

I5ec~iusc priii iary l m d t i r t i o i i is so Iow oii rccci i t lava I I ~ i w s . and i n dry cliniates succcssinn is s low, tlicsc : i rvas c;iii f o r siiiiic: iitiic Ix: r<*asonol>ly iiicludcd ¡ t i ihc catcgory oí "ae~iliaii ccosystciiis" (fIXVAI<IX, 1987). whid i are thosi: wlicre animal l i le depcnds i i ia i i i ly oii tlic iiipiii oí windboriie organic nialerial (wl i icl i we reíer Lo as " h i o lo!: ic a 1 ín 1 i o u t " or " iw ii iiii"). 'l'hc rcsidciii aiiiiiials arc aliiiost al1 cornivores o r scavcngcrs. aiid t l ic i r liiod coiisists i i iaiii ly 0 1 wiiidboriic pollcii, sporcs, sccds, p l a i i t íraginents. i i u i r i c i i i pari ir lcs dcrivcd íroiii tlic S C ~ aiid morr productivc laiid, aiid a wide varieiy ol' acrial ly dispersiiig uriliropods (rcvicw iii EIIWAIWS. 1987).

Colo i i i za t ion of i l ie I i i s to r i c lava I Iows iiiay l i i ive takcii place part ly íroiii populations thai survivcd oii ~Iie islotcs (cí. KUNKEL, 1981, p. IG) , bui \vas probably inainly í ro in ihc peripl ic-ral areas u f older rocks. Soine coasial aninials inay have a r r i v e d by sea, a i i d coloii izatioii by a i r would be possiblc íor certain spiders aiid ni i tcs. I l o w c v c r . i i ios i i i ic i i ibcrs o f ilic res idei i t inlaiid comniunity are incapable of í l ight and cvidei i t ly coloiiizcd i l i c rccc i i i lava by walkiiig o n LO i t after it had cooled, or i n sonie cas<:s pcrhaps h y t r a v e l l i n g uiidergrouiid througli tlie "MSS", íroiii tl ie F rc i i c l i ter i i i "iiiili~xi souierraiii supcrficicl" (JUi>iiil'i'I IIE, 1983). which in Englisli can be rcndcred as i h e "iiics<i<:;Ivcrii<ius sliallo\v stratuiii" (el. IIOWAll'fI l. 1983; OIIOMI, LIEDINA si MARI'IN, iii prcss). l t has bccii showii tliat the fauna characteristic oí the MSS í requen i l y occurs also iii l ava tubcs ( I l O W A l I T I ~ . 1983; OROMI, MEDINA & TEJEDOII, I98ti). Ii i s worth iioliiig that tlie most practicable way oí deinonstrating the ex is tcnce oí t l i is ui idcrgr<iu i id inovci i iei i t would bc by sampliiig i n lava tubes s i tu a ted in extensive lava flows. A L tlic time oí this study i t was tliought that no substaiitial lava tubes wcrc prcsciit iii tlie park.

Tl ic work on Laiizaroic discusscd iii this papcr was carried out i n two seclions. In 1981 MJA aiid NPA collccted saniples at tl iree siles o11 tlie Iiistoric lava ílows, wi th the aiin oí coii ipari i ig ilic iiiiiiiiiil cniiiiiiuiiities nii tlicsc sitcs witl i those on historic f lows on T c i i c r i í r (ASI IMOLE Sr ASI- IMOIX, 19871. Tlie r csu l t s sl iowed that tl ie conipositioii o í tlir íauiiii varicd w i t h tlic distancc íroiii tl ie sea and tl iat i n Tenerife t l i c re werc st r ik i i ig dií lcrci ircs iii tlic füuiias o f kipukas antl tl ie surroundiiig recen1 lava. Thc work thai \ve rarric:tl out togethcr o n Latizarole in 1985 was planned takiiig tliese results iiito occoui~i. Wo (Jbliiiiicd series oí saiiil>lcs at t l i í í c r ~ i i t distaiiccs íroiii tl ie coast aiid sainplcd two kipuk:is (sec saiiipliiig siles).

Iii order LO obtaiii aii idea of tlic kiiirl oí subtcrraiieaii fauna that could occur i n t l i e T i n i a n í a y a lava, vic ur idcr took sa i i ip l i i ig iii 1985 i n the Cueva de los Naturalistas, probably t l i e iiinst appropiatc place fnr such a study despite i ts distaiice íroin tlie park (scc I'ig. 1 ) .

COLUCTING MEI'I IOUS

We used a trappiiig period o í íour days and Iour diríereni col leci i i ig nietliods. As we Iiave discusscd clsewlicrc (ASIIMOLE si ASIIM01.E. 1987) each of tliese methods tciids to be biasscd toward tlic capture of dif ferci i t groups oí invertebrates. l 'he use oí al1 four inctliods a t ciicli siic, I iowevcr , probably rcsu l t s iii the cap tu re oí representnLives oí' i i ios i i i i vc r i cb rn te i i ixa that play aii i i i iportani role iii the local <:liiiiiiiliiiiiy. I ~ : x < : l ~ ~ ' t l l l i i " 111 41111 s i i i i i ~ l i i l d s ~ ~ i i i p l i i i ) ; illiiliiil' iiri. i i i ~~ i l i i~ l i i~ !d ill iIi<! ilccoIIII1s of tlrc sitcs coiiccrii<id.

I

I

I

l . I'itfall traps. These t raps w c r c sc rew- top siraight-sidcd plastic jars o í 4.2 cm d i a m c i c r and 8 crn deptli, aiid coriiaincd 50 ni1 oí a 5% solution oí íorinaliti (40% íor i i i i i ldchydc) in water w i t h ii I i i tk: tlcic.rg<:iit. Tlic trrips wcrc sliadc~l íroiii direct suiiliglit. S ix wcrc uscd i it &di site iii Iioili yciirs. 2. Ik>itle vays wiih 'rurquin's liquid. l'licsc werc 250 1111 disl>osable glass becr bottlcs w i t l i 50 ni1 of Turqui i i 's l i q u i d (TtJI IQUIN, 19731, designcd to be a t t r a c t i v e to arthropods. The modilied íorni that we use consists of 10 g chloral hydrate, 5 ml íorinalin, 5 ni1 glacial acetic acid, I ni1 liquid dctergent aiid water to 1 l i t re. Three oí these traps werc used al each site. 3. Uoitle traps with c h c e s e . Thc saine type of boit le was used as with Turquin's liquid, but tl ie bait was about 3 g oí "Danish IHue" clieese. Three of these traps were used ai each site. 4. Visual scarches. Thcsc wcrc carricd out iii thc vic i i i i iy o f tlie traps i n daylight and iiornially lasied íor a total oí 1.5 Ii iii cacli siie: this time w"us always spl i t between two or more peoplc.

1 3 - 4 5 . w

I

H l S T O R l C L A V A

1 7 3 0 - 3 6 n - u

1 8 2 4

A L E G R A N Z A o M O N T A Ñ A C L A R A

U A

SAMPLING SITI-i

Sampling in 1984 was a t three s i tes (Lago d e Lava. Malpals interior and Malpaís c o s t e r o ) . I n 1985 we iiiade systenialic coilections a t eight new s i tes (including one cave) and put out a íew i raps a l a nintli (Pleitito); we also returned to Lago de Lava 10 put t raps iii deep cracks. Tlie 1984 sites a r e inentioned oiily briefly since they a r e fully described iii ASIIMOLE & ASHMOLE (1987). Some Eiiglish names íor s i tes that were iised in that papcr o r iii correspondeiice with specialists a r e given in parenthesis.

It is possible that iiie lava at one o r more o í our sampiiiig siies dates írom the 1824 eruption rather than the 1730-36 eruptions (compare Figs. I and 2); uníortunately we could not deicrmiiic thc precise bouiidaries oí the 1824 íiows (see 1-IAUSEN. 1959) and have there íore not indicatcd thcni o11 tlic iarge-scale iiiiip (Fig. 2 ) .

Lago de lava (Lava Iake) 1984 aiid 1985 UI'M reí. Fr232088. 350 ni a.s.1. More tliaii 7 kin froiii tlie sea. A sliallow basin

o í his tor ic lava oí relatively siiiooth "pahochoe" ~ y p c . but with crcvasse- iike cracks. Lichen coverage about 10% In 1985 tlie purpose oí tlie sanipling here was inainly LO see whether tlie coniposition oí tlic fauna in the cracks was diífcrent íroiii that íountl on thc su r í acc iii ihc prwious year. Wc (lid iioi use pitíall i raps or iiiakc a visual search, hut used striiigs io lowcr s i x l>ottlcs witli 'l'urquiii's liquid .aiid six witli clieesc iiiio t hc c racks (niax. depih 5.3 id.

Malpaís interior (Iniand nialpaís) IIW

with lichen coverage avcragiiig arouiid 50%).

Malpals cosLcro (Coiuial Malpaís) 1984 UTM re í . FT188158 ( i n c o r r e c t l y quoled in ASIIMOLE & MI-IMOLE, 1987 as

FT190150), ca. 18 m a.s.1. Aboui 200 ni írom thc sca; lava type similar to that at Malpaís interior. but witli hardly ;iiiy licliciis.

Orilla (Seaside) 1985 UTM reí. FI'189152, y 5 ni ii.s.1. Al1 the i raps werc lcss [han 20 in írom the sea

a n d wi thi i i t h e zonc aí íected by sal1 spray. 'I'lie lava reaching the sea here is oí rough "aa" type. with no iichcns and Iiardly any iiioss.

Posadero (Cullrock) 1985 UTM r e í . F1'190150, ea. I Y ni a.s.1. Aboui 200 ni inlaiid aiid about 500 m

n o r t h e a s t oí Malpals costero. No licliciis o r iiiosses seen. Wtli an onshore wind tlie taiig oí salt iii the a i r \vas vcry noticeable.

Barranco 1985 UTM ref. FT194150, ca. 20 rii a.s.1. About 500 ni iiiland. A iow-lying a r e a oí

very dissecled sliarp "aa" lava. A íew tiny patches oí nioss and some sniall lichens in dcep places. Thc only vascular plant that we saw was a single Iarge bush o í Launaea arborescens (Uau.) Murb.

Esquina (Corncr) 1985 UTM reí. FTZ16120. 180 ni a.s.1. This site is 4 kin inland on tlie park boundary

southeast oí Caldera Deriiicja. It is a very dissecied a r c a oí lava, but with no deep c r a c k s . Lichens . inainly S te reocau lo i i vesuvianuiii Pers.. a r e abundant on the nortli sides oí tlie rocks. with a total cover oí about 15%. Thcre is a Iittle m o s and along ihe side of tlic nearby trsck a r e a íew Lauiiacs arboresceiis.

lslotc I lalconcs 1985 UTM r e í . FT153090. 108 ni a.s.1. A reinote kipuka in the southwest oí the

national park. i t is a boonierang-shaped ridge oí old volcanic rock, about 0.7 km long, a r o u n d wh ich t h e 1730-36 l ava í l owed , enc los ing i t on al1 sites and isolating i t tlioroughly írom otlier vcgetated arcas. We doubled our trapping sample here, putting one set of t raps on tlie cast aiid onc oii ihe west of tlie ridge; visual searching was iiicrcascd íroiii t l i c iioriiiiil 1.5 Iir LO 2.5 lir. Tlierc a r e only scai tcrcd plants on tlie ridge, giving about 1-5 'k cover , bu1 on somc par ts oí the sides Euphorbia balsamilera Aiioii is weli cstabiishcd; a varicty oí other piants Iiave been recorded by KUNKEL

UI'M reí. 1=1'214100, 290 ni a.s.1. üii Iiistoric "aa" lava iiearly ti kiii í r m tlie sea;

(1981). Lizards a r e íairly abundant and we saw evidence oí rabbits; a pair oí ravens- (Corvus corax) apparenily nest on the islote.

F igu re 2. Map oí the Parciue Nacional de l 'imaníaya and surrounding area. 'Contours a r e a l 100 m i n i e rva l s . F o r c i a r i t y , oiily priiicipai roads and selected tracks a r e shown. S t ipp l ing i n d i c a t e s aproxi i i ia tc ly t h e a r c a s t h a t were not covered by new vo lcan ic depos i t s during tlic historie eruptions. Froni various sources. Lava ílows oí 1824 a r c not shown bccausc iheir br>tinclsrirs iirc noi cstol)lislic(i.

Abbrcviaiioiis: LL: Lago de liiv:i; Mi: M u l l ~ i í s ifli(!rior; h4C: Malpaís costero; Or: Orilla; Po: Posadero; Da: fh r r enc« ; Es: Es(ptii1;i; I I 1: lsloie I ial<:oiics; LI 1: Lava I - l a l ~ o t i ~ ~ ; IT: lslotc Tahaiilas; 1'1: I'leitilo; CN: Ciicvo I I C ic)s Nniur:ilis[iis.

I

. Lava l lalcones 1985

UI'M reí. FT154090, titi iii a.s.1. Tliis site is about 100 ni írom the northcast edge oí Islote Halcones, in tlie historic lava ílow. It is very rougli "aa" lava, wi th soiiie deep cracks. Lichen (Siercocauloii vesuviaiiuni) is patcliy but abuiidant and covers up 10 50% oí i h c rnck su r lacc i i i a íew smal l arcas. There a re sonie snial l accuniulations o í dust. in wliich iiioss 1;rows. We pul out a doiible set oí traps here. 10

provide a í a i r coinparisoii wit l i Islote Halcones, but two botile traps with Turquiri's liquid were lost dowii decp cracks; visual scarcliing was íor 2.5 Iir.

Islote Tabaibm 1985 UTM reí. FT204154. ea. 30 iii u.s.1. This sitc is about 400 m inland and iii a

large, irregular and relatively flai islote just nort l i 0 1 tlie iiiain expanse oí recent lava i n the national park; i t extcnds íroin Caldera i3lanca. 4 km inlaiid, down 10 thc shore. The site includes a low rocky ridge uiid a bowl-shapcd arca wit l i loosc rocks and ac- cumulated dust. There is about 4OX, covcr oí slirubs. doiiiiiiütcd by Eupliorbiii spp., bui therc are virtual ly no Iicrbs. I<al>bit droppiiigs are iiuiiicrous. Siinipling was carricd out here in order to provide a comperison wi ih tlie transcct on ihc historie lavii nearby; uníortunately wc did not Iiavc tiiiic to do visual searcliiiig ai th is site.

Cueva dc los Naturalistas 1985 UTM re í . FT3010, 325 ni a.s.1. A volcanic tubc situated i i i the Malpaís de

Tizalaya, between Masdüchc aiid Lo Vegucta (k-ig. 11. Tlie lava was foriiicil during tlic eruption oí 1730-315. Tlie lava is of typical paliochoc iypc:, now covcrcd by a dcnsc g rowth oí l i che i i s niainly Stcr<wcaiiloii sp.. I'ariiiclli:i sp. and Xantlioria sp. ai id by some higher plants typical oí rocky Iiabitats sucli as Aconiuiii sp. Tlie cave, with a t o t a l length oí about lti40 ni, has two eiitraiiccs, I I I I I ~ coiiiiiiunicatiiig with a large ma in tube; a nar rower sccondary bl i i id tube branclics o l í tliis and is rathcr mure diííicult oí access (see MAI<TIN Sr UIAZ. 1985). I t is vcry near [he suríace and wit l i frequent siiiall openings wliicli allow thc penctration oí soiiic l ight írom outside, and the tcmperature is Ii igh (around 23 "C) in comparison wit l i otlicr caves. Our standard set oí traps was spl i t betwccii twci stalioiis. boili iii tlic sccoiidory tube; ilie first was i n twi l ight and the secoiid íurihcr iiito thc cave ¡ t i a tolal ly dark zone. Only a ícw niinutes searching was done iii tlie cave, at thc í i rst stiitioii.

I'leitito 1985 UTM reí. FT203157, ca. 10 ni a.s.l. Tl i is siic was un a steep slope oí old rocks

just above the intert idal zone, i iortli-iiortliwcst o[ Islote Tabaibas. We pu l out a íew traps at tliis site, but tl ierc werc varioiis problcnis iiiid wc thercíore do not present t l ie da ta syste i i ia t ica l ly . I l o w c v c r , a ícw specics íouiid here are mentioncd iii the taxonoinic section.

RESULTS

The genera l co inposi t ion oí tlie samples is sliown i n Table 1, mainly a t tlie ordinal leve1 but wi th sonie orders divided wliere tli is seemed appropriate. A summary oí the data íor eacli site is givcn i n Tablc II. Tlic sites are tnost readily compared. however. by r e í c r c n c e LO Figs. 3 and 4. whicli show the relationship between the nuniber oí individuals and tlie nuiiibcr oí species íouiid at cach site. In this analysis we have inc luded tlic saiiiples obtained by scarcliiiig, sincc tliese were important i n adding extra specics, especially oí spiders; wc have also included in Lago de Lava the ani i i ials caugh t in e x t r a t rapping i n c racks i i i 1985, since these cracks are an iniportaiit part oí thc Iiabitat that \vas not sainpled iii 1984. Fig. 3A includes al1 the data, while Fig. 311 slinws tlie picturc aí icr elii i i inatioii o f the Collembola; these were extreiiicly nuinerous at I'osadcro, Ori l la. aiid Malpaís costero, ihree sites which íoriii a clear group i n Fig. 3A, wi l l i Ii igh abuiidaiice bu1 low spccics richness. Islote Tabaibas and Is lote Iialcones, tlic two sitcs oii older rocks, are linked by their high species r ichness, and i n Fig. 3A also by Iiigli Libuiidance; Iiowevcr, iliis is caused by large niinibers oí Diptero in Islotc Tabaibas Iiut oí Collciiibola iii Islote Halcones. The Figs. 3A ai id 313 daia slinw the low divcrsity oí the rccei i i lava sites and the generally higher abundaiices closcr to tlie sea (sce below). 'flic rclat ively Iiigh number oí species a t Lago de Lava probably rcf lccts the greater structural coiiiplexity oí this habitat and perhaps also its closeiiess 10 largcr areas of older substrates (see Fig. 2); tlie cx l ra saiiiplinl: k i t t l i is silc i m y alsn hnve liad soiiic cl'írct.

Tab le l. T o t a l nuinher oí i nd iv idua ls and ( i n parenthesis) species oí arthropods oblained in al1 sampling at each site. For Lago de Lava data íor 1984 and 1985 are combined. No searching was done at Islote Tabaibas and oiily a little at Cueva de los Naturalistas (see tcxt).

t

Nos. i n c l u d i n g search

Preudorcorpionei

Opi liones

Araneae

Acari

lropoda

Diplopoda

Collembola

Thyianura (Microc.)

Thysanura (2ygenI.l

Orihoplera

Procoplera

Homoplera

Heieropiera

Coleopiera

Lepidoplera

- 1 (24 10 (24 1 (1) Diptera (Nemalocera) - 1 (1) -

Diptera (ürach./Cycl.) 7 (44) - 2 U) - 1 (1) 2 (2) 7 (24 8 (2+9 17 (4+) 300 (164) 71 (1)

ilymenoplera (Formic.) - -

Hymenoplera (oiher) - - - 10 u1 5 (39) -

- 2 W 5 (34 -

Tablc II shows tlic high dcgrec oí doniiiiaiicc o f Collciiibola in tlie receiit lava sitcs close Lo the sea. AL n o s 1 sitcs secoiid placc iii tlic ordinal rankings is held by Coleoptera or 'fliysanura, with Dipiera, Acari, Orthoptera (Gryllidae) and Isopoda also rcpresei i ted in the top tliree places. AL al1 bui t l i rcc o f tl ie siles more than íour- íiíths o í the animals collected belong to thrce orders (ir Icss.

A quan t i ta t i ve p ic ture of relationsliips aniong the siies on a broad taxonomic basis is provided by Talile 111, which gives ii i i iatrix o í villucs of Kcndall's "Tau" iridex oí rank correlatioii. CI IENf (1963) drcw attciitioii L o tlic usclulncss of this iiidex i n comniunity comparisons, and its use is discusscd by IlUllTA (1979) and ASHMOLE & ASI-IMOLE (1987). In tlie present sludy thc analysis wiis hased on al1 aiiinials obtained by t rapping aiid searching and was c a r r i c d out at tlic ordiiial lcvel (with a íew exccptioiis). Taxa absent íroni both iiicnibers oí a poir ol s i t a werc oniitted írom tiiat comparisoii. No corrcction lo r tics wíis applied siiicc iiiost tics wcrc at zero abundance

- i n une oí a pair of sitcs: ilic corrcction for tics givcs iiillated siinilarity values i r 1 this situatioii.

Tlie highest value o f thc indcx is that hetwccn Miilpais costero and Posadero, two si tcs niily a fc-w liuiidrcd III~~I":~ iipiiri CII ihc s i ~ i i i ~ <list;liic(: írolii tlic s r a . biit

sampled a t d i í l c rcn i seiisons iind i n d i í lercni years. l'hesc two sites are also l inked by h ig l i index values to Uarranco ( i i si lc i n tlie saine area bu1 300 m lurther inland), Lava Halcones (also on rcccnt aa lava fair ly closc to the sea), and especially to Lago de Lava (oii a ratlier dií ícrci i t typc o f lava aiid niucli íurther inland). Malpaís interior also shows inoderatc siii i i larity wit l i ihe incnibers oí tliis group. The only othcr high i n d e x value links ihc two oldcr lava sites, Islote 1-lalcones and Islote Tabaibas. The l a t t e r site i i a tu ra l l y sliows vcry l i t i l c similari ty LO thc rccent lava sites, espccially tliose close to tlie sea. Orilla, riglit o11 tlie sliore, sliows strikiiigly low index values i n coniparisons wi t l i al1 thc otlicr sites, ii icluding I'osadcro only 200 m inland lrom it. Esquina also stands OUI by i ts rat l icr low similarity to most other sites: i ts closest re la t ionsh ip is LO Malpais interior, anotlier site lar from the sea where we caught very few aninials.

u> W - 0 4 0 - W P u> u 3 0 - O

Table I I . Suiiiniary of data o11 divcrsity aiid dominant taxa lor tlie samples írorn cach site. Sites as i n Table 1, abbrcviatcd as in Figs. I and 2.

SITE LL MI MC Or Po Ea Es LH 111 IT CN

No. of orderr 12 6 5

No. oí individuals (iolail -111 -55 413

No. or species (ioial) 2 3 i 10 6

No. lndivs. (minus Coll.) 51 14 84

No. of spp. (rninus Coll.) 19+ 7 5

Dominance by ordcr:

Firsl C d k m C d h C d h

Second nv- cob4L Thvur

Third C d a P Thyul cCr0;p

7

994

I I

123

9

C d l e n

C d a P

c*uip

6 9 8 8 1 1

1603 77 -59 cal17 -371

10 I I+ 15+ 14* 38*

37 62 29 13 103

6 101 12+ 9+ 31+

13

459

51+

415

so+

D p l

CdaP C d l e n

6

105

8

92

7

D i n

Cdlen

c*uip

% domin. oí 1st rank ca.52 ca.56 80 88 98 30 ca.50 ca.72 ca.73 80 68

?6 dom. oí 1st-3rd rank ca.75 ca.95 97 96 >99 (70) (ca.75) ca.88 ca.86 86 89

Some íurther iiisiglits can be obtained by looking at the taxonomic composition oí the sainplcs in i i iorc detail (Table I and the Appendix). It is clear that the community at Oril la is distiiict íroiii al1 the otliers. I t is r i ch in individuals but not in taxa. and altliougli i t sl iarcs abundance of Colleinbola with Posadero and Malpaís costero, whicli i i rc 200 ni iiiland, i t dií lers íroni theiii in the abscnce oí lepismatid 'I'hysanura (but thc prcsr i icc o í soiiic niachilids), thc prcsence oí isopods m d i n thc abundance oí c r i c k e t s anrl t w o kiiids oí unusual íl iglii lcss melyrid beetles, Gictel la ínrtun;i ta and I í i i idit is- p<-t r ico la. I lar ra i ico, wl i ic l i i s a l i t t l c íu r the r inland than I'osadcro aild h.l;ilpaís costcro ltui si111 i i i the ~ o i i e that 1s virtuolly Iackiiig in liclicns, dil íered íroni them inaiiily iii the draniatically lower abundance of Collembola.

l

l

I

I l I i i i I l 1 i I I 1 I 1 I I 1 I I

i I i

i I

I 1 i l i

Tab le I I I . M a t r i n of similarity valucs lo r coniparisoiis iiiiiong the suríace sites usink Kendall's "tau" rank correlat ion coeíficient. Ihscd on data iii .iablc l.

:;I'rl LL M I MC Or Po Ba E s LII 111 IT

t

Lava Lakc . .- .50 .65 . 2 U .70 .61 .27 .SU .42 .31

Malpais iiileriur .SO - .48 .11 .46 .28 .44 .19 .33 .15

Malpais cusleru .65 .4U - .25 .U0 .53 .16 .57 .20 . I 2

Orilla .20 .13 .25 . J 6 .22 .24 .O5 .O5 -.O2

Posadero .70 .46 .80 .J6 -- .56 .18 .42 .24 .O9

Barranw . b l .28 .53 .22 .56 --- - . O 2 .28 .17 .O3

Esquina .27 A 4 .16 .24 .18 -.o2 - .13 .29 .16

Lava Halconcs .58 .39 .57 .O5 .42 .28 .13 - .21 .15

lslole Halcones .42 .33 .20 .O5 2 4 .17 .29 . i l - .60

Islote Tabaibas .11 .15 .12 -.O2 .O9 .O3 .16 . i 5 .60 -

.I '"1 1 1 ,

1 l H

0 1 , 1

1 0 0 1 0 0 4 0 0 ' O 0 0 1 5 5 0

N U M B E R O F I N D I V I D U A L C

Figure 3. A: lielatioiisliip Itci\v<:eii tlic nuiuhcr oí iiidivitluals ai id ihc iiuiiibcr of specic> arthropotls (iiicludiiic <'«llciiibola) recortlcd Ironi cach sitc. Upward-pointing arrow>

iiidicate thai nunil>crs 01 spccics at ilirsc si tes wcrc iiiiniina, sincc not al1 taxa coulc Iw workcd oiil I(J ihi. XIII:<'I<'S I iw~l. A l i l i r i ~ v ~ ~ i t i i ~ i i x -1% in Fip.1 iiiid 2. 1): l 'hc siitti<\ ;I:

I'ig. 3A, but w i ih Collciiiboia oiiiitivtl.

I l

1 1 1 1 l

1 1

1

i

1 l l

1 I 1

I

z c ooe- r m n 0 O01

o

n o O

I 00,

1-

111

![ .. 00,

-3- s3

se6L

OOOI OOOI O05 OOL O

r- in

n

n

4 S

O m O O v)

.o* ..

several other possibilities. A t present t l iere is Iiardly any iníormation on the microorganisms involved in

successional processes on historic lava oí Lanzarote. Studies on the nitrogeii cycie oí the recent vo lcanic islaiid oí Surtsey, olí lceland (e.& HENRIKSSON & RODGEIIS. 1978). deiiionstratc substancial nitrogeii f ixat ion by blue-green algae, but this process is unl ike ly to be SO iiriportaiii oii Lanzarote since i t depends on high hutnidity. We Iiave, however, dciiionstrated bacierial ac l iv i iy on the Lanzarole lava: cellulose sticky tape iiiipressioiis, trai iskrred io "iiiariiie iigar" (UIFCO Laboratories, Detroit, Michigan) and subsequently iiicubaied at 20 "C by Dr. W.U; Grani, sliowed that Posadero and Uarranco had signiíicam populations of i ieterotrophic bacteria.

ASHMOLE & ASHM0I.E (1987) suggested that the coastal parts of tlie historic lava on Lanzarotc iniglii rcccive sigiiiíicant quaiitities oí organic material in the íorni oí niarine bacteria concentrütcd in bursting bubbles at sea and carried over the larid in the "sea-salt aerosol" ( I>LANCiIAI<D, 1983). Iii 1985 wc aitenipted Lo í ind out whetlier input oí t l i is ki i id was grcatcr close Lo the coast. by exposing marine agar plates on sniall posts a i tlie íour sitcs (III our sampling traiisect íroiii Ori l la inland io Esquina. T W o scrics oí cxixri i i ienis wcrc spoiled respectively by rain aiid iiiteiisc sun. but single plates cxposcd for aI>out iiii hour oii ilic liist day o f our íield work al Ori- lla. I'osadero aiid I larrai iro cocli ptotliicrtl u varicty o í bacteria1 colonics on iricubatiori (W.D. GIIANT. pcrs.coiiiiii.). I t i l icrcforv secins possiblc that input oí iiiariiie burteria plays a signiíicant role iii ilic tiiiiiiitciiaiicc o f tlic lava ecosysieni close lo tlic sea.

As indicatcd uiir l icr. wc supi>osc tiiat in inwt barren lava ílows o i i thc Caiiaries ilie niain resource oí ilic ariliropod coiiiinuiiiiics is biological íallout, inainly coiiiprisiiig aerisl ly dispcrsing artlirol>otls. I t i 1984 we uscd water iraps oii Latizarote to confirii i that some íallout occurrcd i i t Lago de Lava (ASIIMOLE & ASIIMOLE, unpublished). I L seeiiis uiilikely, Iiowcver, iliui ilic rccci i t lava iiear tlie coast rcccives tnucli arthropod fallout, sincc i t is aii arca w i i l i prevaleiit oiisliore i iort l icr ly winds, although tiierc are sonietiincs pcriods with castcrly wiiids Iroiii ilic Satiara.

Arioiher possibility is siiiiply thai Iiigli biological ac l iv i iy i n the i i i tcrt idai zone, couplcd with ilie i i i o l~ i l i i y (11 iiidivicliial aiiiiiials, leads LO traiisfer oí organic nialerial up to a íew Iiuntlrcd iiiclres iiiliiiid: tl i is input could i i i priticiple suppori populotions of microorgaiiisnis and a coiiiiiiuiiity oí rirtliropods.

A L p resc i i i wc I iavc iio I m s i s [<ir judging wliicli oí thcse processes is iiiost i inportaiit on tlie historie lava oii Laiirarotc, and i t is clear tliat inuch iiiore work is needed before the i iutritional hasis oí t l i is unusual ccosystem is íul ly understood.

The sainpling oii Lanzarote iii 1984 provided the basis íor a comparison between the fauna oí historie lava ílows on Teneriíe and Lanzarote (ASI-IMOLE & ASHMOLE, 1987), aiid the ncw daia do iioi cliaiige that picture significantly. In a subsequent study of a r e c c i i t l ava í low iicar tlie coast on Hierro Island, MARTIN, OROMl 8; IZQUIEI iDO (1987) foutid i l i o t iii t l ic iriipping slation closest to the sea ihe most abundaiit arthropod was thc i i ic lyr id I>cctle Gictcll;i íortunnta, wliich also occurs o11

r e c e n i l ava i i ca r LIIC sca oii 1,:iii~arotc. Oii the Hierro lava ílow Collenibola were relat ivcly scürce. thc coiiiiiiuiiity sliglii ly íurthcr froin tlic sea beiiig domiiiated by itie dcrmapteraii Aiiai;icliri Iaviciila Martín & Oroiiií; aiiothcr specics oí Anaiaelia is typical oí coasta l Iiabitais oii T ~ i i c r i í c (GANGWEIIIS c i al.. 1972) aiid is nlso known íroin Goniera. but ihc gciius lias iiot bceii recordcd I r c i i i i t i ran Canaria, Fuerteventura or Lanzarote.

ACKNOWUUGEMEMS

Tli is work was carricd out wi ih grants froin thc Nuíí ield Foundation, tlie Uritish Counc i l aiid Edinburgli Uiiivcrsity, whose Iielp is grateíully acknowledged. N.P.A. aiid M.J.A. wish 10 thank tlie Director and staíí oí tlie Departamento de Biología Animal (Zoología), Univcrs idad d c La I ~ g i i i i a , for i l ic i r wcIc«iiie and for tlic provisioii oí l abora to ry racilities. Sr. L u i s i'ascual, who \vas ü i r e c t o r Coiiservador del i'arque Nacioiial de 'fiinaiifay;i, kiiidly gsve perinission íor work iii ihe park, ivhilc Sr. Ai i toi i io Fernández de Tejad;i. currci i t Uirector-Conservador oí tl ic park, prtivided us with up- to-date details o f the park bouiidary. Sr. Esicbaii Robayria gavc niucli practica1 help aiid also assisted with ilie íield work. W.U. Graiit kindly gave advice and hclp with microbiological tcchniqucs.

We a r e niucli iiidebted tu tlie followiiig people íor identification of specimeiis: M.A. Alonso Zarazaga (Curculionidae); K I I . Askew ( 1 lyiiienoptera); C. Uacli (lhysanura);

J. Uarquín (Formic idae) ; K. D land (Lepidoptera); R. Constantin (Melyrldae); R.H.L. Disney (Phoridae); M.M. da Gama (Col lembola) ; G. G i l l e r í o r s (Coleoptera); J.M. GonzSlez (Dryophyta) ; D. Hoese (Isopoda); M. Ibáher (Gastropoda); D.K.McE. Kevan (Orthoptera); D. MacFariane (Acari); A. Machado (isopoda, Coleoptere); M. Meinander ( I'soc o p t e r a ; J. J. Ranib I a (Opi l iones) ; J. liibes (tieieropiera); L. Shncliez-l'iiiio (Liclieris); G. Schmidt (Araneae); J. Wünder l ich (Arancae). t ic íerence inaterial is being kept in the Departamento de Biología Aiiinial (Zoología), Universidad dc La Laguna; type spccimens wil l be depos- ited i n ihc Museo Insular dc Ciciiciah Naiuroics <Ir Siiiiia Cruz d r Tciierif(:.

M e n i qr ( M e 1 y r i dae 1 ; J .A. Mur ph y (G tia p hos idac) ; M .

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OKOMI, l'., A.L. MEDINA & J.L. MARTIN. in press. Tlie g e n u s Licinopsis Bedel (Col., C a r a b o i d e a ) in thc C a n a r y l s l a n d s a n d its d i s t r i b u t i o n i n . t h e u n d e r g r o u n d envi ronnien t . irl(.tii.l3iospCoI.

OKOMI, P., A.L. MEDINA Sr M.L. 'I'EJEDOH. 1986. Oii ilie existence of a super í ic ia l u i i d e r g r o u n d c o i i i p a r t i n e n t iii t h c C a n a r y Islands. A c t . l X Congr . ln t .Espe leo1 . I3arcclona, 2: 147-151.

PLATA-NEGHACI-IE. 1'. & A.M.J. EVERS. 1987. Kevisión d e l g é n e r o línidius E s c a l e r a

261 -270.

' 158.

." S -

1

GASTROPOOA n ieba pisana íMüller) Canariella sp .

Geogarpus canaricnsis í Tullgrenl

P5EUOOSCORPIONES GARYPIDAE

OPILIONES PHALANGIIOAE

üunochelis spinifera (Lucas) ARANEAE

PIIOLCIDAE Sperinophora fuertevenlurae

SwriiiolJioru sp . OECOBIIOAE

Oecobius sp. L lNYPHl lDAE

Indei. D YSDE RlDAE

Dysdera sp. GNAPHOSIDAE

Scolagnapha ? canvesq (Simon) SALTICIDAE

Phlegm lucasi (Roewerl Pelienes sp. Chalcascyrlus sp.

SCYTODIDAE ScyladeS lenerifensis Wunderlich

THERlDllDAE Stealada gmssa (C.L.Koch) Sleatada sp.

Wunderlich

ACARI ANYSTIDAE

Fam. indei.

PORCELLIONIDAE

Indei.

ISDPODA

Parcell io laevis Laire i l le Parcellionides sexfasciatus

ARMADILLIDAE Armadil lo sp.

(Budde-Lund)

2 - - - - _ _ _ c o n d e s c r i p c i ó n de u n a especie n u e v a Ifiiidius p e t r i c o l a n.sp. de las Islas C a n a r i a s . Eiit.lMi¡iier, 83 (213): 160- 165.

SCiimZ, L.. K. JAENICKE & 11. I'IE'I'KEK. 1981. S a h a r a n dust t r a n s p o r t e d o v e r t h e . .

- + l 1 Noril i At la i i i i c Occan . Geol.Soc.Amer.Spec.Pap., 186: 87- 100.

Kappe. c . K . 9 6 ~ Congr.aut.Soc.sav., Toulouse 197 1 , S c i e n c e s , 3: 235-256. 'I'Ui<QUlN, M.-j. 1973. Une bioccnose c a v c r n i c o l e or ig tna le pour le Ilugcy: le pui t s d e

- -+2 _ _ _ - - - - _

2 - 1 - 7+1

- - - _ _ - - - _ _

2 -

t t - - - - _ - _

- - - _ _

2 -

- - 4 - -

- - - -

- + I -

- - + l - - + 1 - -+u

- - - _ _ _ - - _ _

2 2 3 22+2 13 - - - - - - _ _ - - _

2 - 1 1

- - 1 -

1 -

- -

HALOPHILOSCIIOAE fialophiloscia couchi íK inahan)

Tylos la l re i l ie i Audouin TYLIDAE

Fam. indei. DIPLOPODA

POLYXENIDAE Indet.

COLLEMBOLA H Y POGASTRU RID AE

Xenylia b. brevisimilis Slach Xenylla sp. l lalosenylla arfiniforniis (Siachl

ISOTOMIOAE Fblsoinides angiiloris (hxelron)