studies of hawaiian freshwater and soil algae ii. algal colonization and succession on a dated...

I'HY 1 OPLANKTON-PRO I EIN CONTENT 171

ptiysioliiKy 'll |)i<iln/(Kt. tn HiiincT. S. M. |F<1.] Prntozna. v<il.IV AciKlfiiiit I'li'Ns, Nfw Vtuk. 11-59.

1."). SMrnkin, C. \*i7'S. Cirontli mcastirt'tnciils—(hird wcijjhi,|iiukc(l (fll vcilttmt- and opiual density. In Stoiii. J. R. IKd.Iliiinithuok oJ Phycotogirnt .\teUiiui\, C:iinbrklf{e Univcmly Press,

Itt. Siiiikl.ind. ]. 1). II. & I'aisons. T. K. ]9(>0.•'•' '' ' ' " • Binitil Ciui., Ruttetin N". 12.^. P iS-

17. MKiO. Drti*rmin:ili(m nl prntein, ht.\h. Res. BnardCan.. BuUeiin Nn. 123. 113-fi.

IS. U)68. Dficriniiiaiion of pariiiiilulf nitrngen. Ft.\H.Rf.s. Bmirii Can., Bitttetm No. lt)7. 221-2.

H). rrainiH-, V. R. l'.(71. t)cvclnpment nl Inrm in Srrttrttr.\mii.\.fn I'arkiT. B. C. & Brown, R. M.. Jr. |Ed.I Conlnbutiom tnl'liy>olngy. Allrn I'ICKS. Inc.. LiiKrcmi*. Kiinsas. Hl-il2.

-2). i'.>7S. pp. 171-I7S

STUDIES OF HAWAIIAN FRESHWATER AND SOIL ALGAE II,ALGAL COLONIZA'riON AND SUCCESSION ON A

DATED VOLCANIC SUBSTRATE'-^

Johnny L. Carson'^ and R. Malcolm Brown, Jr.nf Bnt;inv. I'tiivcrsity nf North Oirolina. Chajwl Hill, N.C, 27

ABSTRACT

Spatial studies iij colonization and succrssinn oj soil al-gae and chemical analyse.^ oj the x-arious .\oils on tbe cinderrone oj Kilaura Iki in Hawaii Volcanoes National Park,Hawaii are ontlinrd. Tbere is a j)ositivr correlation be-twrrn tbr ilii'rrsity anif quantity oj soil algar witb nutrientIciiels and organic matin accumulation in racb locale.Tbrer distinct rdnjihir-biotic zoiirs f:\isti?ig in this area arrdijjnentially rn'raied by the sod cbcmical compo.sitiori,{/uanni\ and divrrsity oj soil algar, and as tt'itlrfit vari-ations in biglier plant giowth and colonization. I'uryingColonization and .succe.'isional jibasrs oJ higber jdantgtowth around .'.tanding and jallen tree snags killed byvolcanic activity also rrjirct variations in the sod algalflora. Thrse variations ajijirar largely as a Junction oJdifjrrrntial watrr intercrjition, ab.wrjition, and retentionOS rvrll as difjrrrntial accnnnilation oj organic matter, andthe initiation of various biogeochemical cycles.

Key index word\: Bacillarioj/hyceae; biogeochemical cycles;Chloroj>hycrar; colonization; Cyanojjhyceae; ediij)hic al-gae: Hawaii: microbial ecology: soil algae: succe.'i.sion: x<ol-canic sub.\trate

The tlii\v:iii:ni Islands coinprisc a ciiaiii of sm:illislands of voUanit oiigin in ilic mi<l-l'atilK Oceani-xiendinK ' • ' " " *i>- I9''-a9*N an<l (a. inS'-lHO^W.lying i;i. :^,'2()i) km rn»ni Liny innjor land niiiss. Tliecommon cjccta of the Hnwaiian volcanoes indutica.sh. cinders, and varying texttnes ol lava. The mostahtnuUint (heniiial rompoiiLMit o( tlic Hawaiian vol-taiioes is silica which mav comniisc r>07r of the lava

'Accrfileil: It December 1977.* III ic'(»)Kiiili(iii (ll many yrais nt sdioljrly prndurtivity in phy-tlnjfy. wv Like HKMt |ilfasurr in dfdUaiinn this paj>cr m Dr.liigi I'lnva.^nli.' Adftrt'ss Inr ri'piiiil

content (9.14). Other oxides prevalent in lavas in-clude alumintmi. iron, magnesiinii, calcium andsodium (9). Prevalent volcanic gases are water vapor,carbon dioxide, sulfur dioxide and nitrogen (9).

The weathering of Lhe volcanic siibsuaies throughgeologic time has produced dilTerent types of soils.but much of che Hawaiian ccuinirysidc consists oilava flows. niotnUainotis lands, beaclies, and ash andcinder cones (1). 1 hese areas are miscellaneous landtypes and comprise appvoximately halt of ilie landarea of ihe Hawaiian Islands (1,10).

The island of Hawaii is an ideal location to inves-tigate tlie role of freshwater and soil algae in thecolonization ol" sterile land surlaccs and early eco-system development (Fig. 1). Volcanic areas havebeen the object of a number of ecological investi-gations (2.4-7,12.13). Origgs (7) noted that volcanicash fallout areas were ideal for investigations into.soil genesis and the initiation of the nitrogen cyclefrom inorganic N-free ash. He mentioned the im-portance of liverworts as pioneers on the unciis-turbed volcanic ash of Kaimai. Alaska, but notedthat they played no such role in the colonization ofKrakataii (lava). Criggs noted the absence of blue-green algae on the Katniai ash, but observed bac-teria, ftnigal liypliae. moss protonemata and thegreen alga C/j/orororfum humicolum (Naeg.) Raben. inthe liverwort complex. Brock (2) also observed li-chens, mosses, vascular plants, and chlorophytes onthe new substrate at Surtsey (Iceland) and suggestedthat temperature is responsible for the dilTerence inimportance of the C'yanophyceae and Bryophyta aspioneer forms at Krakatau. Katmai. and Surtsey.

Study .site. Kilauea Iki cinder cone in Hawaii Vol-canoes National Park (Figs. 1-4) evolved as an ashfallout area during a two month rift erui)tion ofKilauea Volcano in 1959-1960. Kilauea Iki is a pit

\TZ JOHNNY L. CARS<^N AND R. MALCOLM BROWN. JR.

\'OLCANOESNATIONAL PARK

Vi(.. L Map of isliiniK ol Hawaii: loc<itirin ot Hawaiio National Park and Kiktnca Vnkaiio indicatrd. 2. Miip ol Kilanra Iki stttdy area showing soil rolledion

i s : sec lexi lor explanation.

crater adjoining tho northeast ritn of Kilatica Vol-cano. The otitirc area i.s ca. 1.200 in in altitude. Theclimate oi the area is tropical inontatie. The meanannual tcmperaltirc at the park hcadtjuariers.which is at approximately ihe same altiiude ai«l 1.9km from Kilauca Iki is 15.9 (' (.5). The mean month-ly rainfall at the headqiiariers is >I00 mm extepiin Juno when it may be slightly less. A gradient dropol ca. 1,000 nitn in annual rahilall occins fVtim Ki-laiiea Iki south to the upper Kau Dcsorl (12). Theash from llie ertiption produced a large cinder coneand was also distributed from its origin (at the veniin the crater wall) southwest, by ilic trade winds overan ellipiical area ol ta. 500 ha and a dejiih ratigingfrom 15 m at the crater rim lo the suitimit of thecimier c<H\e down lo H negligible dislribiilion in iheupper Kau De.sert (12).

The volcanic activity left a peripheral porlitjn ol"the stiiTounding dimax forest ol XUtrosidrros andCibotiitm relatively unaffected. However, the ashdenuded and parlially covered a number of trees ina wide zone interior to the climax forest. Since theeruption, some of the snags have fallen with higherplant growth occurring anmnd these standing andfallen snags oi Mftrosidrros. The ash lallotit com-pletely obliierated the innermost portion of ihe af-fected area.

The edaphic and vegctational features revealthree di.stinci zones (Kig.s. 2, 5);

Zone / : Outermost region comprised of a definitesoil sub.strate lying to the peripliery of the cindercone which remained relatively unaffected by thevolcanic eruption; consists largely of climax Metro-idCibi ibrest.

Zone 11: "Btilfcr" or transitional area interior toZone I in which the pyrodaslic material either de-stroyed or considerably delbliated tlie vegetation,leaving a titiinher of tiee snags, of which some re-(oveted, but many others were killed. The lichenStrreocaulon vulcani appears commonly in hare asliareas, I'ioneer vasctilar plants and lichens are com-mon es])ecially arontid tree .snags and there is someorganic litter over the ash and cinder base (Kig. 6).

/««/• / / / : Central area interior to Zone II com-prised oi the cinder cone summit and immediaiesurroundings which xvere comj)Ietely covered byvolcatiic ash and spatter from the eruption. Thereis only nominal evidence of returning vascularplants in well dei'mcti areas, unti S. vnliani is prev-alent over the ash sub.strate. but there is liltle or noevidetice ol organic Utter over the volcanic substrate(see Kig. 5).

MATKKIAI-S AND METHODS

iininauoiK itt ihe soil al^ac were made hv Nam-ihe lop 2,5'! cm ol the voltanit snhstiaieai lemilar Intervals

along :iO.l« m iranst-ri'. at seven lolledion sites in lite Kilaiiea Ikiarea i.\-"i^. '2). The siimples were iolle(ie«l aseptimlly in ]>luMkhiiKS and shipped tiurncdiately lo the L'niversity ol North ('arn-lina. <;hapel Hill, where ten samples Irom eaih iMiirit tn the inin-.tett were ciiltiirefl liy Mispendiri^ ^^ g ali<|iiots ol the M)il in ^0 mllt(|inrl Bold's liasil Medium (BBM) (»); soniratinj! mildly; plating0.75 ml ol llie suspension over riidi ol two 100 mm petri dL-sIiesol sterile agari/ed HBM. The mltuies iimihiited at least 2 wk atta. 7.000 Ix al lil C on a I(>:H l.t) lycle helore laMinmnii ilcirr-ininalions were made.

Quantitative dvtt-rtiiinatiniis ol un\ atgiic wrrc miide l>y MIS-peruling ."> K alii|notK ol siilmiate material Irom eadi siie in .10till litjiiid HttM; sonicaiiriK inildly; plating O.I ml o( ;i 1:10 dilu-tion in liquid BU\f in eadi ol iwo 100 tntii jictii disht-K of stctilc.

sou.. AIRIU>R\t ,\U;.\K o r I1.\WAII

I AtU t I. thtiimnwlums i>[ grurru iJn'tJMt\ of Oif \otl nlgitr at Kileinteiih.

l - ' i t . . ^^. A f n . l l p l i o i t i g i . i p t i u l K i l a u c a V o U . i n o s h o u t n g j H t i -

t i o n t i l l . i i g i ' I . i t i U - i a ( I t ' l l 1. p i I i i . i l e t , i i i ( l t i i i d r t u n t r n l K i l . i u t - a

I k i s l m w n III l i n s c i T i g l i l | U M i i i ( ' / » ( « ( / ) , I K . . 1. ( . u m p . i i . i i u f i i i -

h . i i t d p l i i i t i i g i a | i l i i c v < \ i l i T i g M i l i H - i K i n t . i n l u . i t i n g n l K i L i u t M

• i l i d K i l a i i i M I k i ( i ; i t < - i I l i i o i s , i i i < l ( i m l t ' i t i i i i l c i ( i i i u ' M | K i U t i U M

I k i un I I'll h lint), t ' l i o i n g i . i p l i i c M . i K a n ; i l i s c i m ' n l ( l i l l r i t - n i i a l M il-

i a i i u i K . i l i n g w i t h l l t l i l g l i l d c t l c i [ l i . u u i a l < n l n i i i / a i i n i i a n t )

'•Ml I ( " . M O M , ( P h n i n g i . i p h s ( t i i M U ' s \ [ ' . i l i M i i ( . , I ' u l i V U , K n \ i i t i i i -

M u - i i l . i l K c v c a i i 11 I MM n i l I t ' n l M i t h i g . i i i . A i m A i l m i . )

U » \ r I h c u i l l i M c s i i u i i l M k i ! '1 w k . i l u i . ^ . f ' O t t I x m

iiN l i ^ l i i .11 I . I , 1^1 i i i l i ^ v

l a i K l n i n i m ' n t i t . t i l i o t t w n | > l . i i ( ' s t i n i n t M c h i l i ' S i g i i . i i c d M t t .

I t t u n i l i c M ' i l . i t . i . i i i . i t l K ' t n . i t u . i ) t I f t c i m i M . i l i M i t s w t - i r i n . u l c u l l l i t -

i | i i . i n i i [ . i i i ^ < - I I I I ' m i / . n i l > u p n u ' i i M . t l >>l [ h ( - ^ n i l . i l g . i c g u c i w i s o i l .

.•\ l i i n l u - i < | u a i i [ i l . i t i \ ( ' . t s M - s s i n t ' i i t 111 t h r s o i l . i l g a l l i m a a t t - a t l i

N i l e WAS i n . i i l r t i s i i i g ( l i l i > i i i ) > l i \ l l ( i i i u c i i l i i i t i n i i i i s a n i i l i t i i . i t o i i n

t h e i i t l t u i o d ( - M t i l x d t n t i | i i i i 1 i i ^ i i i v i - i l t - i c i i i i i i i . > t i i ) n > . A l a i K l o i n

I u n ' l i i i i i i t i i i w a s K - i i i i A c d I K H I I i l i t - 1 i i l i i i i t - d . i g a i M M l a t e a n t i

i - v t i a d c d I t l l t i l l - i i r H i n t ' i i i t n ( > i t t \ o l i l u - g n u v i i i g t f l U i n I I I m l

. n f t I m e a t t l i t ' ( u \ t k •»! ( i i l i i i i r g m w i l ) u a . I i n n ) . A h s o i h . t i u c w . i s

T i i » ' , i M i n - ( | .11 i ; U l I I I I I .

S l . t l i N l i i . d . m . t l v N t s i \ t ' H - d n i i c I I I d c H ' i n i m t ' I I K ' I I » I i f l . i t i n t l l > i -

t \ t t ' t - i ) M i l l n u n l e n t I t - v c l s . i n t l [ l i t - p i o d i t i t i M t \ o l l l i c s n i l i i l g . i r . l ^

•I l i i i H I I K I I n l i n l . i l m i l n t i i ' i n l g r - i K ' l . l i v > l 1 ( - i l r d , n i t l u l H ' l n l i n l n -

i n < ' \ t i l ( ' 1 i l t u i i | i l t M f . i i ' n l i s i - i v f t l i n i i i l i i i i i - . a i i < l i h t ' 1 1 i l i I I t i i . i t t ' i i . i l .

S i i i d i f s w e l l ' m n U ' i t a k t ' i i m . i ^ s f \ s t l u - n i l l i t c i u r n t t l i t - . m i i -

• M i i l . i i i i i i i o t Ml g . m i i l i t t r i . m i l M i l l I n i n i a l m n o i l t h e d n f I s i t \ . m i l

" ] i i . i i i l i l \ u l s n i l ; i | g . u - I t l \\\v M i l i . t i i i i s . t m p i i - a i t . t s .-\ T . l V J i n

t i a n s i - i i i i i n n m g N t S W w a \ < l t - h t i i ' a i t - d m / o n e I I M i t i t l l i . i l

( M i l l c i i i t w . i s - . i i i i . i l ( ' d 111 l u i f v n l i a t i H . t s l i w i i h Iii l i f t ii i i n o i g . i i i t i

l i t t f i 1 1 i e I f l i t I . l l ) m i t i n n n l t l i f i T . m M - t t p i l ^ •> l •d t i c M I n a s t i l i l . i i \

Mittii-idiU'S t u - c M L i g a i n i m t l ( \ l i i i l i p m i i t f i \ f g f l . t t n i n . | t i i i i M i i l \

tiiiddhfit iiMiitna 1 ( H I I . ( \ a \ ( ' > > t : i i i l l s l i f d a n d i l i f i c w a ^ i n i i M d i - i . i l i l f

" i g . m i i l i i t f i .111 i i i m i l . i h d n ( I i g ( i ) . I l i r d i i t - i I m n . i l . i l i g n n i f i i i u l

t i l l - M . i n s t - t t w a > ( i i i i > j s l f i i i w i i l i t h e - p i t v . i l t n t N l - t i . i i U " w i n d s

S . t i n p l t ' s n l t h e • . i i t i M i . i l f W f i f m l l f t H < 1 .11 i f g u l a l i n i c i \ . i h a l c M g

t l i c t i . i i i s r i l l i t ' g i i m i i i n a t l l i f n n i t l u M - . i e n d , I l u - s . i i n p l t ' N w c i r

t i i l i i i t i i l l i \ • . i i > i i c i i d i i i g '» g u l S ( i i l I m n i t . K l i p n m t . t | o i i | ; l l i f

t l . i i i M - i I . m I I I n t l 111 |i I III l U l M ; - . n n i t . i i i i i g n n U l K i n i :W) %. p l . i l i t i g

" . ' J . '> m l III l i l t ' M i v p c M s i n n i n ,i p c i i i i l i s l i n t . i g a t i / r d H U M . I I i f

 w t i f i T i > p f ( l ( ( ! l o i g i - i n - i n d u n s i l v

• i T i d i n n i i i i ' t l . l l i f i l n f i M i \ a n d a l i i i i u l a n t t - o t i l i t - M < I I . i l g - u '

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t;UI.()ROI'llV(:KAK(.htiini\diiiniiiia\V.hloullii(.hloiiiii'ii ittnUunnuiiiimSiinmnhlous(i<•ni^ll.^t'nlmrtliirufniyt'lUliUiHi'llSStiiliiii iirfii\l',luii\\U\

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X

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tJ...^

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/•inr l.iliiius liirrM.Biil • Aim- M.hiillri nmr

around st.mdiiig .iiul l.iilrti tKT sn.igs wvw tU-tciTiiiiic-rl hv sim-itai inrtliotls iiM-d tor ihi- i{U;tntiMtivf t(>iint> dexr ihr i l . Die "niNh n i l ) t h e s » \ t i i i l i ' s i g l l . l t f d s i t t ' S i l i i t l t t t i i n t l u - n t l u i l u l d s U l t l i i ' s

v h f i f I l i f t i i K a l K . m . i K / f d l>v > t . m d . i i d mi - l l ix i lN (."^.l t ) .

HIM I.IS

Ihc t;liUuo|>hyicaf wcir ihc piiiuiplc intupn-nt-nts of llif soil algae ilirotijrlKuit ihc Kilatica Ikisiiidv ait-a (lahU-s I, 2), Several ^cm-ia wcti- (n>-iiiopolilaii: vvlioiras. oihcrs wciv iiiorc irstriilcd inilu'if (lisit ihtiiion. Si-vi-ral m'tUMa. possil)lv liavittj;nion' riji(»rt)tis nuMitidHal rc"(|iiiicnifnEs. (KILU oiilvin ihf (litiiax iuicst atca t/.t»tu' I). The Clyanuphy-iv.w \st'H' itiainly lotttit! iti thr diinax lorcst area(/otu- n ahhou^Ii three genera vvere ie(i>\eif(iIroin the hulTer (/.one 11) and <levasiaie<l i / iuie 111)areas. Otily one genus ol (iialonis was leioxeredIrotn M)il sani]>les attd onlv troiii ihe tUniax lores!and hiilTei atea. In lernis ol intal getteta mllecled.ihe (liinax lores! and Inillei /otu-s exliibiied iliegteaicsl diversity ol soil algae while llie tonipletelvdevastated areas illtisiraiecl a more litiiiieii algalIlora.

(Jdotxtoftum sp. was ihe tnt)si lointnonK enmnn-lered geiut.s iti ihe ilitn.ix lotesi area al Siie A lol-lt)ued thiselx h\ Chlorrlla sp. ( lah le 2). Xavirula sp.was the onlv nieinhei ol ihe Uaiillaiiopluicae ot-t tit ring contmoiih in Siii- A (iliniax loresil: how-e\et, it was a pie\ak'ni totnpunent ol lite soil algallloia in lluil atea. llormulmm sp. and Stithonut its sp..ittu)ng tlie C'hloroplnteae also showed >igtiiluantrelative heqiieiuies ttl 70' ; oi more In Siie .A (1 ig.

174 JOHNNY L. CARSON AND R. MALCOLM »ROWN. JR.

TABLE 2, lirtative jifi/urncy t'A) nf soil algtil grtiera al Kihiuea th.'

/.Of IF.A

Diil-

CMLOROIMIVCKAtChtamyitomojuisCM>retlaC.hliirtHixtum

O^f^^tl•.

0 0 ().i 95 1000 0 nf)

•15 10 095 100 90

100 20 070 100 20 no fiO 904.5 90 10(t 75 100 MK)45 0 0 0 0 0

0 0 0 0 0

SU(ho(iiffH.\Jehiify\ln

CVANOI'HYCEAtAnutkimaEitUifihyyatii

OHitlaloriat'luirmultum

BACILLARIOI'UVt.E.AENovuuta

755

1550

750

20

4')35

0

000

150(1

0(I50

150

3010070

00

50150

00 10 0 0 0 0* ViybA ol !fO tiilluirWutr: jpi

ay;' A l i l i i r > u l i t i i » j i III r.ili)r I

nir iil iiiilii jiril |{rnrij in r;«ili nl

2). The only significant <K:currcncc of the Cyano-phyccac in the entire Kilaueii Iki area was in Site Awhere Sostoc sp. appeared wiih a relative IreciiieiityotTO'i.

Although widespread in Siie A, Cblorocotrum sp.is uncommon in adjaccnl Zone 11 Siie B; however,the rehuive lre{|uen(y of Hormitiium sp. intrca.sedmarkedly in Silc B. Tlie sigiiilitance ni Chlorella sp.as a [jicjiieer alga ihroughoiit tlie entire area i.s readi-ly apparent as its relative rre(]iiency was never lessthan 95'/?. The relati\e f'rec|iien(y vahie.s Cor soil al-gae of Sites B and (i in Zone II were similar, re-flecting their similar microhabitats, and edaphic andlioristic features. The high relative frequency of thealgal genera at Site D in Zone III indicales the capac-ity of this particular locale lo support a more exten-sive algal Mora than the other hare ash sites.

Quantitative assessments (Table 3) at Kilatiea Ikiindicate that the Chlorophyceac comprise the dom-inant soil algal flora for lhe entire area. The greatest

Iki.K -1. Ctiliirrifittytl ubuntwnre vfcuttureit .\tiit miiletitil fu«n Kitiium

Aitlrl'r;iiiM-<

SD

BillDcv

DcvUiil

.22

.01

.09,07.OK.20

.01-.09

.02-. 12

.01-. 12

.00(i

.009

.000

.001

.001

.001

.0L-.• Mc.iii VJIIIP •>!

numbers of Clilorophyceae occur at Site I), the sum-mit of the cinder cone, but counts approaching thismaximum wore also observed at Sites A, B. and G.Neither the Cyanophyceae nor the Bacillariopliy-ceae appeared quantitatively significant anywhere inthe Kilauea Iki area. The chlorophyll extraction olcultured soil material from Kilauea Iki (Table 4) in-dicates high chlorophyll concentrations of culturedalgae in those site.s corresponding to high genericdiversity atul high quantitative detertninations.

The chemical analy.ses of the Kilauea Iki soils(Table 5) show the soils from all siies were acidi(.with the most acidic being in the climax forest andbuffer /ones, and the least in the hare volcanic a.sh.Thegreate.st concentration of inorganic nutrients aswell as organic matter occurre<l in the climax—Me-trmidtros-dilwtinm forest of Siie A in Zotte I. "Tlteorganic maitcr content of Site A s(»ils is >-l x greaterthan in any other silc. Niiratc-N iit Silc A was alsofar in excess of thai in all other sites. The valuesobtained for the soil chetnical composition of SitesB and G in /one II were .similar. Likewise, siinilarvalues of inorganic nutrients and organic tnatter alSites C D . E. and I in Zone III suggest the groupingoi these sites into a distinct zone. Niirate-N levelswere very low throughout Zone III. Statistical anal-yses indicated a liigh level of correlation betweenthe diversity and abtindance of soil algae and thelevels of inorganic nutrients and organic matter inthe soils from which they were collected.

Qtiestion.s have beeti raised as lo the characteris-

Tiantm-tiir

mionir*tni '

lAHU. 3. QiuinUtatn

CHLOROPIIVCtAfc

R.inKr SI)

'e eounti t>l wil otfrnr al Kitauea 1

C VAMU'IIVCIAJ

tm' Kjiigr

'ki.'

Ml

EIA< It lAKIIH'IIVCI-AI'

" c S " * Ran r M)

CliButDfvD<vDtvDcvBiif

2.1.33t).30.7

39 52.13.2

26.4

I (Ml7-760-4

12-030-100-9

\'^^.79•ifiH.Ol

1.79219. S3

11,01H.()2

I HO. 9 3

0.0l.'i0.00.00.00.00.0

0-40.002.270.000.{)00.000.000.00

0.90.00.00.00.00.00.0

0-2 2.270.000.000.000.000.000.00

i u M iAlit>tr\jutii>n>

inirili m titmlum Mdtinjtol

S i m . . AlKBOKNK A U i A l t)I- II.WVAII /.'I

I K . , fi. \ lew l.tiitig Nl- l i i i in snininit ol Kil.Mic.i llî t i t idei(ot i f NhinMiig tlii i-c diMinit id.i( i l i i t and it-gt t.itiotml /unrs titi-ullt-i ti-(l .\Ulii'\itti-n>\-(.ihiitiinii ti i i i 'sl (/tnic 11 III li.H Lgroi i iul : Ad-iiticnc In tlim.ix |ni CM IS " h u l l f i " ^tii|> I/.I me- 11) shitwnig iTiiiindeati I n c si),igs,niil | in i inci ^cgcMticm: in In icginui id (/ luu- l i t )l i i i l t ' Milcnnt Mjltsti.itc. I'll., I). \'u'w til single |ilt>t ul piotU'fiVfg(-i.iiii<n antl teninani ttt-e snag \n /.one 11.

l i ( ; s . 7 . H. l i i l l i i e n t f o l l u r s n . i g s t m M M i o t i i i t l i n g | ) l . in i h t e .

l-l(>. 7 , s l ) t i \vs a t t i v e g i n w i h u l t e r n s Smllnui s | i . l l o i t - g r i i t i t i d 1 . i t id

Xif'hiiitif'-'i.-- s p . a i i ' i - t u d s t . t n d t i i g v t t a g s : l l < ; . S. s l t i m s \ifi/ni-lt-fi<,i.\

SI), w liii 11 . i p | i . i i r t i l K ^1 CW i i i i u i i u t a sUt i t t l i t i ^ ^>i-iK ' " K (iK'i i h;i( k

w l u i l t h e s n . i g l e l l .

l i t s o l i n s i l i i ^ n o w l l i o l l l i r s o i l a l ^ a o o n l l i f s u b -

s ( i ; i l r . p i l l i i ( i i l . i i I \ w l i f l l i n l l i r ; i l ^ ; ; i l H o i a i s i n ; t n i -

it 'Mnl in uiuhaiaiU-iiMit lu in is in (ului i i - aini iKiir t ilo iKiiun*. r iu- (v;niopl iv(c.ni Tnhfiidhrix sp . Ii.i>iKt'ii i cmj ;n i /c ( l in l o l l n l ions in.uii- c l iu 'dK l io inthe voltanit ;isli. DiiiKini (rustiilcs were iil.so cIrarKidi-i i t i lnd hv (liict t mil loMopit iii>iniu<tn ol i h r st»ilas ( o n i p o n c n i s ci l l ir soil alâl Ilitra. Oi l ier noii-(U-s( I ipi t-noiifs wliic tl \v<-if liki-l\ lr;i};im'ntN ol hoili^'irt'ii and hlnt'-giiH-n a lgae \ \ t ' i t ' o h s c n c t l . hiii tu l -niiiii^' iuul niittftMcipit i n s p n i i o n ol llu- stiil iiiiilf-lial is p n s r n i K nrcrssaiv loi l l ir ton iph- ie idcnli-litaliiMi ol i\xc soil al);al llora.

Siiuiit-.s (tl soil a lâe a long a i ransc t l in / o n e IIg sotiiliui'si Irom hitrc ;isli throuj^h a small

U- (it nig.init mallei a(einntilali<)n. |)ionfct vej;-clalioii. aiitl a i rum.nil n ct- siia^;. and liai k onl() ihe

bait* ash substrate, show a j;reater tliversitv atulabimdanif ot soil alâe to ihc iinnu-diatf It-ruartlside 111 ihr iiansect U iil)Ie {j>. AU values lov soilI hrmiial i.cim|)osition e \ t f pi lor manânrsr arr alsolnj;hri to ihr iinnu-diair Irrwaid sU\v ol the iian-srtl.

1 nvesiiâiioiiN (4 soil al^.ii- iiioinul standing; nndTalk II live .'*naj;.s in / o n e II show a more ahundaiit;iud di^r1sr tloia a round sl^nuiin^ snags thaniironntl lalleii tme.s (ligs, 7. S; lablr 7). 1 he Clhlo-lophvtrae are tlcarlv ilir most dominant (^nantila-live and <]iialiiali\e coniponent.s in boih area.s al-I hough nominal hlncgrren algae and tliatoms wrrc*ohsrrxfd. Stiil clK'initai anaKsrs around slaiuhngand lallrii Miag> inditatr grrairr levels ol organicmat i r rant! all inoig.mii compoiu lUs extejit sultale-S around standing snags.

I A«U :i./J./.7M(in«/K(H> "/ nunu-ul Ir-.rh. figuni, mulln. pit tn Kdii,,,4i Iki

27 \i

1)

V

' A l l n.i l t init liw-lh .III \\i\\.> 1-1 -'11 1 111 <lcplh

1 n '>l:v.'<III)'.I.M

:iti (I

17:11 17MShi7•>7f,

I )

I).'.I1

I'.iLIdO1 ri ' jtm11111)11L'HiOll•d-d HKI

171.1r.rj.7

.'in 7.0

l.i)

M1.N

•2-2. t

22.1Ili.O

Hl.6

NO, S

IOt.3lo.s0.0tt.O7.1O.tJ

l l l .K

Ot){4IIH.iiuiiri '1'

tl).7'i.t)o.iiO.()0,7O.ti1.5

ItMiit

••>A

t i. lfi.O5.S(i.55-5

JOHNNY L. CARSON AND R. .MALCOLM HROWS. JR.

G, Colnniiahnu piitmtial ami MIH tnMly.\es around a solitary tter\nag u-tth fnimrn vff^rtotion ami iiif^anit ilfbti.\ at Kilaiiea Ih.

distance (cm)Niimlxrr genera

colleclctjTi>t:il ('olonic!i

ChlorophyceacPhosphorus"I'utassiumCaltitim.MugneMumM;ingjnt'%cStillatr-SAmmoniiim-NNiiraic-NOrg.iiii( matter %pi 1 ill u';itcr

Nor

-366

1

7531.5

244.015200.'.51.0

13.66.4

84.20.00.75.9

ihcjii Sn tg

-1K3 0

4

65H76.5

431.72,1600.'.13.2

2.914.4H9.440.S

1.05.(>

.Viuiliwrti

1H3

6

14 OH

157.5753.K

404007.SH.7

I.K24.0«J7.(>

135.H2.65.7

366

2

2.14

45.0

253.412100413.1

3.214.4.16.70.0O.H5.7

' Klf l lJ II ' 20 tm itr|M)i.

DISCUSSION

Site A in Zone I is comprised primarily of undis-Itirbed climax forest o(Xfelrosidrros and Cibolium andhas developed a mature and diverse algal floi a. SitesB and G, lying on opposite sides of the cinder conein Zone 11 show edaphic and floristic .similarities andhave the second greatest diversity of soil algae. InZone II. moisture from precipitation and fog ismore readily intercepted and retained by the tteesnags; and. the initial layer of organic matter andincreased nutrients may enhance microbial growthover that on the bare ash. Site B may be regardedas the windward component, while Site G is tlie lee-ward component of Zone II. The relatively greateravailahiliiy of moisture to the windward si<le mayacccnint for the greater generic diversity of the soilalgae there. The close proximity ofthe climax Ibresito Site B may also result in an accelerated rate ofmaturation ofthe soil algal flora. The tree snags andother vegetational barriers to the wind flow in thiszc>ne may also increase the friction a I element of thewind flow resulting in a greater deposition of air-borne propagula. Site D illustrates significant diver-sity in the soil algae and conversely a marked paucityof inorganic nutrients and organic tnatter. The rea-sons for this are twofold. The collection points atSite D were at the summit of the cinder cone wherethe wind speed at times reaches 05-75 kph and themicrometeorological turbulence is considerablygreater than lot the area in general. 1 hese featuresallow for: i) greater inotulaiion of ait borne propag-ula; and. ii) greater influx of moisture. These fac-tors in concert result in greater colonization of soilalgae near the sumtnit of the dnder cone. Sites Cand V lie interior to Sites B and G respectively, butin Zone III where there is little edaphic or floiisiicsimilarity (Fig. 2). The lestilting generic diversity ofthe soil algae in Sites C and I' is substantially lower.The generic diversity of Site t is low\ comparableto that of Sites C and F. This may be due to several

TABLF 7. Iiit>ti< anil rilaplui /'raliim nf (hr mirrohahilah anmnil staml-ni^ ami laltfn Itff snugrs al Kilaiua Iki.

t'|)tiKlii •nuR l-rfllm fiuic

t. Algal gcnciii divcisity Chhttlla(Moriiroiium/I mm lilt urnPrfitnrnfCUiXa'i'iorhliin.i\ii.\torNavtriiln

Onrystis

11. AlgjI qiiatiti(ati(jt)"

<caf (tit*

I I I . Soil chctnii-at romposi-

roi:is.siiitn('iilriiim

Atnmnnivttn-N

(' tnattt-r '.r

t.4

0.0

27.0

3OH0O-195.7

.1.2I2 .M0.1.0lO .S

2..1

0.0

0.0

9.0137.6

140002IS.7

2,620. H

<t.6I.I

Mri in irl [;)Tiil<irTi s nyt iil 10 iK / l < : / l l l l

Mlll;<[r.

factors. Kirst. the substrate is basically bare volcaniciish with little organic matter accumulation, pioneervegetation, or tree snags. This allows le.ss letentioiiof tnoisture and considerable desiccation by the con-si.stent ttade winds: and. there ate indications thatthe turbulence characteristic of the cinder conesummil has subsided at this point with a correspond-ingly lower intKulation potential (unpubl.). Theubiqtiitous genera of ;ilgae (i.e. tho.se ofctirritig a(every site) are very iinportaiu as primary colonizerso( Sites C. E. and K whete envitonmentui conditionsare severe. However. CJiIoroeoirtim sp. and Protocoe-nis sp. were collected at only fotu" sites, clesct ibed ashaving an acctnnulation of litter, greater water re-tention in the substrate, or stibject to high wind,turbulence, and a greater influx of moisture, sug-gesting that their re(|uiretnents are more rigid andthat they may represent threshold pioneer form.s.

Kr<»m the <|uantitative data obtained, it is estimat-ed that a minimum viable colotiization potential ofca. 5.5(H) cohmi/ing uniis/g soil exist. A potentialmaximum viable colonization potential of ca.310.000 colonizing luiits/g of soil material may existwithin these particular volcanic substrates. Ihe vari-ation in the counts of soil algae in the Kilauea Ikiarea is greatest in those siles wbete avetage couniswere also high. This implies ihat areas with higherlevels of nutrients and organic matter are in a con-stant flux of metabolic activity in a variety of micro-habitats with a resulting departure from the normin (juantitative cotints as opposed to tho.se unlavor-

SOIL. AIRBORNE ALGAE OF HAWAII 177

at)Ic or less iciiilc areas witli (Vwcr niclics. whereaverage (DUIUS and slaiulard ctevialions were lower.The low levels ol oiganic mailer, lack ol iiilrogenas well as other hiorganic ninricnis. and relativelylov\' counts of soil algae on the bare ash sites eni-phasi/e the irni)ortance ol" niicrobial and pioneerhigher plant metabolism in the initiation ol'the ni-li'ogen cycle as well as olher biogeochemical cyclesin the.se locales.

Siaiisiital analyses indicate tbat there is a highpositive linear correlation of generic diversity andchlorophyll cottcentration with the levels ol' phos-phorus, potassium, calcium, magnesium, man-ganese, NH4'-N, and NO.r-N in the soils. A positivecorrelation ol geiiei ic diversity and clilorc»phyll con-centration with levels orSO<^*-S in the soils was neg-ligible. This is likely due to the inconsistent dept)-sition ofsulfur-coniaining pyroclastics over the area(luring tbe eru[>tion at Kilauea Iki. Correlatii)ii anal-yses of the mean number of colonies of Clilorophy-ceae with the various nutrient levels was less defin-itive, althongh positive correlations were observedin all cases except against levels ofSO^'^-S.

Qualitative and quantitative determinations of tliesoil algal flora along the transect running from bareash thrtnigb a /one of organic matter acciunnlationand back onto the bare ash strongly suggest a moreextensive algal flora tti tfie leeward side of tfie tran-sect. We suggest that this may be due to the actionof ifie snag and nearfjy vegetation as singular bar-rieis to ibr generally unidirectioniil wind (low. Thisbarrier metliatcs nol only the dei>o.sitinn of debrisand litter from vcgctatioti in the plol, but also thedeposition ol airborne propagiila toward the lee-ward side of ifie snag, resulting in a tnore rapid rateof cr)loni/ation (unpubt.). Increased numbers anddiversity of soil algae in addition to higher levels ofinorganic and organic nutrients have been noted ontfic lee\vard side of tree snags. Tfiis suggests ifiepreferential tadiation of microbial colonization willitrade wind direciion and empfiasizes the importantrole of tho trade winds in the unidirectional distri-bution of viable microorganisms, organic debris,atid the initial colonization of these isolated locales.These observations imply that tlie accumulation oforganic litter atid accompanying increased moisturelevels are re(|uisite for tbe growth, succession, andmaturation of the soil algal flora.

The studies oi algal growth around selectedstanding and fallen Metro.\i<Iero.\ snags reflectedgreater algal generic diveisity and (juantity aioiindtbe standing snags as opposed to the fallen ones(Figs. 7. 8). I'urtberinore. ferns appear lo be grf>w-ing in profusion around standing snags, but theyfreqtiently wither and die when the snags fall over(Fig. H). I'pon inspeclitni. standing snags wereIbtiiul lo contain significant amounts of water intbeir trunks, apparently ac(|uired by interception ofprecipitation, or fjy capillary action from the sub-strate, or both; however, it appears tbat the fallen

snags, lacking this moistme, inhifjit successful colo-nization of the soil algal Hora in that microhabitat.Soils analyses indicate greatly increased nutrient Icv-els except for SO4'*-S in the soil near single standingsnags as opposed to fallen tree snags. The highSO^^^-S content of tbe substrate around lhe fallensnag is probably a remnant of volcanic activity ratherthan a result of niicrobial or plant metabolic activity,rhe content of organic matter around the standingsnag was over two times as great as that for the fallensnag. Such data accentuate tbe imiiortance oi or-ganic debris in the ac<|uisi(ion of suitable nutrientand moisture levels to establish viable microhabitats,the initiation of liiogeochemical cycles, and ultimatecolonization by higber plants.

Many investigators bave studied tfie processes ofcolonization and succession of plant life on new sur-faces. Tlie examples provided by the Kilauea Ikimodel have added to our knowletlge of early plantcolonization and succession because: i) the originalsubstrate being completely sterile, colonization wasnecessarily effected by propagnla arising from otliermature substrates. Complete initial sterility is an es-sential feature for allowing the study of colonizingforms and mechanistns withont interference, con-tamination, or deflection from previously existingbiota, ii) Specific sites of residual organic matter onthe substrate (i.e. tree snags) am fje readily delin-eated and correlated with the productivity of'the soilmicroflora. iii) i he N-free, inorganic ash providessotiie insight into the initiation oi vaiious biogeo-chemical cycles necessary for the establishment oflhe algal Hora.

Tfif atiihors express tlu-ir sinu-ic apprcriatitin to Di. MiixwcllDoty. Dcparliiient ol Botanv. t'liivcrsiiy "i Hawaii, lor dis intcr-csi and assisiaiuc dining this ]>roitit. Wt- also thank, iht* Apro-noinii DivLsioiiol the Norili C^-iniltiu Di-p:iMnu'iit of AgriciiliureInr help in conchKliiig the soit ttieniital analyses, and parlitiilarlvMr. Ray Ttukei wlui uiis mtisi lielphil in (he inti-ipietation olthe data. This work was supporietl in part bv a f>rani ty (he llaivaii Internaiional Biologital ProgramEtology I'rojcd. This nianiisirii>t is a iMiriion ol ihc dixioraldissertaiitm ot" j . L. QiiM

t. Armslrong, R- W. \h.tl.] 1973. Attm of Hau-aii. lhe fnivcr-sily Press. Hoiiohilti, Hawaii. 222 pp.

2. t^iock. T. n. t*,t7S. I'liinaiy loloni/atmn ol Siivt-Sfv. wtihspmal relfieneo lo ihe hlue-grcen uigue, fMoi 24:'JS9-4S.

:l, Brown. R. M.. j r . &• Bold. H. t:. I»tl>4. Hno.logital MudicsV. (Comparative stndies ol the iitj;at genera reliary\li\ andChtttrttrofcum. L'niversilv ol Texas. Austin, i'lihtitation No.(1417. 213 pp.

4. Campbell, a H. UKMt. The new flora of Krakatau. .-Jw. .Wjf.43;49-tiO.

5. Holy. M. S. & MiiclIt-i-Dombois. D. l<Hi(i. Alias lor Biiveroltigy Studies in Hawaii \'olran(K-v National l'ark. lla^saiiBoianieal S«it-ruf Papri No. 2, l'ni\eisity ol Hawaii. Mun-ohilti. 307 |))>.

fi. FRglei, W. .A. lyfi t, I'Linf Hleor Pariiuiin \okano. Mexiioei}<ht years alter atuvity teased, Am. Mul. Sal. 6*t:3S-(iS.

7. Grig^s, R. F. 1*)33. The ioloni/atio)i ol Katmai ash. a newand inorganic 'm\\:Am.J. Boi, 20:92-113.

178 JOHNNY I.. CARSON AND R. MALCOLM BROWN.JR.

8. Jacksiin. M. L. 195S. Soil Chnnunl Anatyîs. Prcnli(c-llall.Inc., FnglfW(H)d Clilis. New Jprsry. 19K pp.

9. Matdonald, G. A. He Ifuhbiird. U. M. l<)73. rotranon of OirXaltotuit I'arki in Ituti-aii. Tong); l*ub[tsliiii){ Co.. lnr. lloito-lulii. Iliiwuii. 56 pji.

10. MtCall. W. W. 1973. SW CMiMifuaiion nt Hau-aii. Cwjpera-ti%'r Extrn&ion SctA-itt;, Univrrsity u( I1iiw;iii. lloiioltilu. 31pji.

11. Mt'liliih. A. I'J.'iS. Dflfi'ininatinii of |ihosphoiiis. pnuis-magnesium. «Kliiini. und amin«mi(im. Noiili

Soil Tmting Division. Nfiinco^rîplicd p;ipcr. Rii-trigh. N'orili Ciirolina.

12. SinatlitTs. (;. A, it Miiclk*r-l>oinlH)is. I). li>7:V Invasion iinilrcrovi'iy ol vf(4ciaii<in iilit-r u voUatiit fiuphon in H;iwaii.I>i)ii)i{iil Report No. 10. 2ti<I issue. I'.S. InttTiKitionjI Bi-oloîtal Pin^rain. Inland KcosyMeiiis l^t(•Îill(•d RL• ear(llI'lojet't. I'niversily ol Miiwaii, Hon<iliil(i. 172 pp.

13. Iifnl). M, 1HH8. Noli(cstii la notivclk- flora dt- Kiakaloa.Ann. Jttrd. Hut. HuitrinuTf^. ~i.'l\i-2'.\.

\'\. Ufhaia. C , Ikawa, H. &: Sato, II. 1071. C.uidf toSoih. College oi •ri-oi»i<al Afjriciiltinr. Hawaii

Siaiion. Hnnohilii. MistelUinc-oiiN I'liblitaiioii

J Phyct.. 14(2), 1978. pp. 178-lB'i

TRANSLOCATION OF '*C. IN MACHOCVSTIS INTEGRIFOUA (PHAEOrilVCKAE)1.2

Christof)hrr S. Lohhnn'^

Bumlield Marine Staiinn, Bamlieltl. British Colnmtiia. Canada VtlR I BO und

Depariment «f Biologiral Sciences. Simon Frascr University,Burnaby. British Columbia. Ciinada V.'>A lSG

ABSTRACT

Transloration f/allenui in the giant kelfi, MaiTot-ystisintegrifolia Bory, were itwestigaled in .sit7i usinf^ '^Ctracer; souTcr.\ and .sijiks werv itfrritified. Export ii'a.\ firstdetertrd after 4 h of labeling; exfjnimenLs were routinrh24 h continiwii.hylh and frond initials at thebases of thejronds. Blades <0.3 mjrom the apex imfwrtedand fiid not fxfwrt; tfiis distaruf did not change seasonally.In spring export jrom blades 0.3—1.25 m from the aftexvm, exclusively up-wards; oldn blades also exfmrted down-wardly. In fait downward export began 0.5 m from theapex, arid blades >2 m from ihe aftex exftorted exclusivelydown-wards. Carbon itnftorted by frond initials, yoitngfronds, and sftorophylls in fall may fjurtly be stored forgrowth in early spring. Xo trawilocation was seen in veryyoung fjlanb until one blade (secondary frond initial) hatlbeen freed from the apical blade; this blade exfrnrted to theaf)ical blade for a time, hut imfiorted when it began todei'elop into a frond. The .second- and third-formed bladeson the primary fronds (sfwrofthylU) also exfmrted when<0J mfrom thf apex, and later stol)f;ed. Frond initialsand sf)urof)hyli\ un later-formed fronds did not exfunt atall. The translocation fmttern in M. integrirolia differsfrom that f}rei'ioiisly reftorted in M. pyrifcra in setLwnalchange and in distances from the aftex at which the changestake place.

Key index words: kelp;translocation

Macrocysli.s; Phaeofihyceae;

f 16 Drrfmber /977.' Deditntrd to Luigi l*rovaM>h with sincere wishes for rnntin-

ued Kholarly produciivity.' I'rcNcnt addreu and address lor reprint rcf)iic»t.i; Division of

Science, University of New Brunswick, Saint John, Newwick, Canada E2L 4L5.

Ciiatii k e l p s o i ' t h e g c i i i i s Macrocy.sti.s, itiwith other Laininariales, have long bt'cn known toli;ivc sieve elements wiili the pciforiited ctid wallstypical oi vascular plant jihlocni sieve elements(Esau 1909). Cnifis (19M9) showed exiKhiiion Ironicut sti|)e.s oi' Macrocysti.s fyyrifvra (L.) C-. A. Agai(ih,and Sargent and Uinlrip (1952) showed that growthol young blades at the tips ol ironds ol this speciesconki not be accounted ("or by their own photosyn-thesis, and must have been supported by olderblades which fixed excess carlion. Direct observationot translocaiion was reported by Parker (I9t)3,I9()5) who showed movement of" ' C and fluorescein<lye upwards and downwards from s(turce blades.in particular, he showed that '*C. could move fromthe 60th blade to the apical blade, supporting Sar-gent and ljntrip's conclusion. I have recentiy ex-tended Parkei's work f»n this species (LolibanI97Kb), showing that there is also downward ttans-location into sporophylls and young fronds, andslu)witig the changes in imjôrt aiui exjxirt i>atternas ',\ blade matures, in the present paper 1 (iescribemy experiments on tbe smaller, mot e boreal species.M. integrifolia Bory. 'I'he transltKation pattern wasinvestigated in relaiion to seasonal changes in tfiegrowth of the plant.s, and to their early deveiop-ment.

MATP:KIAI..S AND MFTHODSt'Tan\lofaliim fwllrrm iii miutt fitnnts. rxpniiiK-nts wcir tariird

oiil in Bat kify .Sound. Vanrouvrr hiand, Brili^h Cohniibia, |irin-ripally im platits in a Itelp t>pd at the mouth ol" Batnlield Intet,

studies of hawaiian freshwater and soil algae ii. algal colonization and succession on a dated...

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