LE RÔLE DES NUAGES DANS L'ÉVOLUTION DES PALÉOENVIRONNEMENTSMONTAGNARDS DE L'AFRIQUE TROPICALE

THE ROLE OF CLOUDS lN THE EVOLUTION OF TROPICAL AFRICANMOUNTAIN PALEOENVIRONMENTS

VEILLE CLIMATIQUESATELLITAIRE

N° 46Août! August 1993

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En zone tropicale, le développement des nuagesstratiformes et la persistance des brouillards paraîtavoir joué un rôle majeur dans l'évolution despaléoenvironnements.

Lorsque l'on prend l'exemple de l'Afrique, toutesies études concernant l'histoire de la végétation ontabouti à la conclusion que, durant les phases derefroidissement climatique, des associations végé-tales montagnardes se sont étendues à plus ou moinsbas'se altitude. Lors du dernier maximum glaciairevers 18.000 ans BP, les analyses polliniques effec-tuées en Afrique orientale (VAN ZINDEREN BAK-KER & COETZEE, 1988; BONNEFILLE & RIOLLET,1988; BONNEFILLE & al., 1992 ; VINCENS & al.,sous presse) ou en Afrique centrale et occidentale(MALEY, 1987, 1989, 1991 ; BRENAC, 1988) ont per-mis de conclure que les températures s'étaient abais-sées entre 3° et 8° C. Outre ces données déduites dela végétation, l'abaissement des glaciers d'environ1.000 m qui a été mis en évidence sur les hautes mon-tagnes de l'Afrique Orientale, a aussi abouti à desconclusions similaires (HAMILTON, 1982 ; ROSQ-VIST, 1990).

Pour comprendre cette question de l'abaissementdes températures on peut tout d'abord examiner lacomposition actuelle des flores et des faunes demoyenne altitude entre 1.000 et 3.000 m d'altitude.Ainsi, par exemple, d'après HALL (1973), les espècesde plantes communes entre le Mont Cameroun et lesmontagnes de l'Est Africain sont de 53 % pour la forêtde montagne et de 49 % pour la prairie montagnarde.Des résultats comparables ont été aussi obtenus pourdivers groupes d'animaux ou d'insectes (MOREAU,1966 ; EISENTRAUT, 1970 ; MALEY, 1987). Afind'expliquer ces similarités, de nombreux auteurs ontestimé que durant les périodes plus froides du Qua-ternaire, les flores et les faunes montagnardes sesont étendues à basse altitude, ce qui a permis desmigrations entre les massifs montagneux (ibidem). Etde fait, des analyses polliniques effectuées dans dessites très éloignés des hautes montagnes, commepar exemple à environ 80 km au nord de Brazzaville,

~mETEO~FRAIlCE

ln the tropical zone, the development of stratiformclouds and the persistence of fogs has appeared tohave played.a major role in the evolution of paleoenvi-ronments. . .

When one considers the example of Africa, ail thestudies concerning the history of the vegetation haveled to the conclusion that, during the phases of clima-tic cooling, mountain plànt associations have spreadto a more or less low altitude. For the last glacial maxi-mum at around 18,000 B.P., the pollen analyseswhich were carried out in East Africa (VAN ZINDE-REN BAKKER and COETZEE, 1988 ; BONNEFILLEand RIOLLET, 1988; BONNEFILLE and al., 1992,VINCENS and AI., (at the publishers) or in centralWest Africa (MALEY, 1987, 1989, 1991 ; BRENAC,1988) have enabled it to be concluded that tempera-tures had decreased between 3° to 8°C. Other thanthe data deduced from the vegetation, the lowering ofthe glaciers by about 1,000 m which was shownclearly on the high mountains of East Africa, has led tosimilar concl.usions being drawn (HAMILTON, 1992;ROSQVIST; 1990).

ln order to understand this question of the loweringof temperatures, one can firslly examine the currentcomposition of middle altitude flora and fauna, at alti-tudes varying from 1,000 m to 3,000 m. Therefore, forex ample, according to HALL (1973), the percentageof plant species common tci Mont Cameroon and theEast African mountains is 53 % for the mountain forestand 49 % for the mountain grassland. Comparableresults were also obtained for various groups of ani-mais and insects (MOREAU, 1966; EISENTRAUT,1970 ; MALEY, 1987). ln order to explain these simila-rities, many authors have estimated that during thecoldest periods of the Quaternary, the mountain floraand fauna have spread out at low altitudes, whichenabled migrations to occur between the mountai-nous massifs (ibidem). And in actual fact, pollen ana-lyses carried out on sites far from the high mountains,like for example at 80 kms to the north of Brazzaville,on the Bateke Plateaux, where a vast hollow at about600 m altitude (Bois de Bilanko) (fig. l)contains depo-

0 - Eorîts Sempervirentes Aiafrêennes et Fabonaises l nombreuses .&*5&/3-1-&(*&*(1 - Eorîts ;tlantiques Littorales è %&(2+/266-5 +&'21*15-5 et g23,-4& &/&6& au Bameroun) ce der-nier arbre êtant remplacê au Fabon par i.270*& ./&-1*&1&( La limite entre 0 et 1 est irrêguliere etprogressive- 00 - Limite de l&extension vers l&est et le sud-est de l&Okoumê 'd&aprés LDROY-CDV;L) 0862) citê dans M;LDY) 088.(-

2 - Eorîts de type Bongolais caractêris~es surtout par l&alternance ou le mêlange de formationssempervirentes et semi-caducifoliêes- %9

3 - Eorîts Blairsemêes l Marantaceae et Zingiberaceae-

4a - Mêlange des types 3 et 4b-

4b - Eorîts Sempervirentes l f-/'*46-2)*1)421 )*9*84*-(

5 - Zone inondêe presque toute l&annêe) avec des formations sempervirentes) des raphiales etautres formations hygrophiles-

6 - Eorîts semi-caducifoliêes-

7 - Mangroves-

8 - Eormations montagnardes diverses-

0.- Savanes-

00-voir1

01 - Dxtension maximum moyenne actuelle des influences %rafraôchissantes% 'd&aprés S;INT-VIL)0868(- Be rafraôchissement observê au niveau du sol est dm l la persistance des couverturesnuageuses stratiformes non prêcipitantes qui s&êtendent sur une grande partie de l&;frique cen-trale pendant la grande saison séche annuelle 'environ 3 mois) de juin è septembre( 'voir le clichêMêtêosat de Kuillet 0872(- L&abaissement marquê des tempêratures) en rêduisant l&êvaporation)favorise le maintien d&une humiditê atmosphêrique êlevêe) supêrieure en moyenne l 7. ACB&estsurtout ce dernier facteur qui permet l la vêgêtation forestiére de perdurer sans dommage durantcette longue saison %séche%particuliére-

02 - Erontiéres entre Dtats-

03 - Prés de Arazzaville) position approximative des sites de Ngamakala 'N( et Ailanko 'A(-

~IFEC%--%-R

0- Aiafran and Fabonese Sempervirent)Eorests with numerous .&*5&/3-1-&(*&*(

1 - ;tlantic littoral Eorests with %&(2+/266-5 +&'21*15-5 &1) g23,-4& &/&6& ln Bameroon) this lattertree being replaced by i.270*& ./&-1*&1& in Fabon- The limit between 0 and 1 is irregular and pro-gressive- 00 - Dxtension limit towards the east and southeastof the Okoumê 'according to LDROY-CDV;L) 0862) quoted in M;LDY) 088.(-

2 - Eorests of the Bongolese type especially characterized bythe alternènce or the mixture of sem-pervirents and semi-deciduous foliolated formations-

3 - Low tree density forests with Marantaceae and Zingiberaceae-

4a - Mixture of types 3 and 4b-

4b - Sempervirent Eorests with f-/'*46-2)*1)421 )*9*84*-(

5 - Zone which is f0..ded almost ail the year round) with sempervirent formations) raphiales andother hygrophiles formations-

6 - Semi-deciduous follolated forests-

7 - Mangroves-

8 - Various mountain formations-

0. - Savannas-

00-see1-

01 - Burrent mean maximum extension of %cooling% influences 'according to S;INT-VIL) 0868(- Thiscooling which is observed at the ground level is due to the persistence of non-precipitating strati-form cloud covers which stretch over a large part of central ;frica during the long annual dry sea-son 'approximately 3 months) from Kune to September( 'see the Meteosat plate of Kuly 0882(- Themarked lowering of temperatures) by reducing the evaporation) favours the maintenance of a veryhum id atmosphere) on average higher than 7. AC It is above aIl this latter factor which enablesforest vegetation to support without da mage this long) particular %dry%season-

02 - Erontiers between States-

03 - Near Arazzaville) the approximate sites of Ngamakala 'N( and Ailanko 'A(-

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oqrughu wr h-sodlq wkhh-whqvlrq ri prxqwdlq elrèwrshv dworz dowlwxghv-wkhvlpsohvw k°srwkhvlv zrxogeh wrhvshfldoo° frqvlghu wkhfrrolqj ri wkhdwprvskhèulf whpsh udwxuhvzklfk kdg uhdfkhg d plqlpxp dwdurxqg 28-111 C0S0Wklv frrolqj kdg ehhq yhu° pduènhg dwkljk odwlwxghv-zkhuh wkh juhdw lfhèvkhhwv kdgghyhorshg- exw kdg ehhq ohvv pdunhg lq wkhWurslfv)DOLPBS- 2982 AULQE dqg SFWFFW- 2986,0 Wkhuhèiruh- ryhuwkh Biulfdq àrqh- dv vwdwhgderyh- wkhorzhèulqj ri whpshudwxuhv- dffruglqj wrwkhlqglfdwruv xvhg-zrxog kdyh ehhq ehwzh0hq4é dqg 8éD )KBPLOWRQ-2983A CRQQFHLOOF dqg do0-2993 AZLQDFQV dqgdo0-2994,0 Krzhyhu- li wkh orzhulqj ri whpshudwxuhvdsshduv wr kdyh ehhq jhqhudolàhg dwkljk dqg plggohodwlwxghv-wkh skhqrphqrq dsshduv wr kdyh ehhqpruh frpsoh- lq wkhwurslfdo uhjlrq0 Hru h-dpsoh- lqBiulfd- wkhqxpehu ri orz dowlwxghvlwhvqhdu wkhFtxdèwruzklfk kdyh ehhq vwxglhg e° sdo°qrorj° duh irxu0Hru wkh Odnhv Crvxpwzl )Ikdqd, dqg Cdurpel Per)ahvw Ddphurrq,- rqo° rqh prxqwdlq wd-rq zdv hylèghqwdworz dowlwxghgxulqj wkhhqg ri wkh Sohlvwrfhqhdqgmxvwxswr 9-111 wr8-111 C0D0(%/*'('2*04.4f 480gR0 -1(-45*55*3.. zlwk dwfhuwdlq ohyhovuhodwlyho°vljqlilèfdqw sroohq shufhqwdjhv ydu°lqj iurp 21 ~ wr 41 ~)PBOFb- 2987- 2989- 2992 ACUFQBD- 2988,0 Wkhwzr rwkhu vlwhv rffxu lq wkhDrqjr rq wkhCdwhnhSodèwhdx- wr wkhqruwk ri Cudààdylooh;wkhCrlv gh Clodqnr-suhvhqwhg deryh- zdv odujho° grplqdwhg e° dqdvvhpeo° ri Biurprqwdqh sroohqv- zkhuhdv wkhvhfrqg vlwh-wkhQjdpdndod srro- 66 npv wrwkhvrxwkri Clodqnr dqg 43 npv wr wkh qruwk ri Cudààdylooh-lvvlwxdwhgdwdssur-lpdwho° 511 p dowlwxgh-exwlv hqfdèvhg lq d sodwhdxzkrvh dowlwxghvydu° iurp 611 p wrpruh wkdqVRRp (+.,g2,0Wkh ghsrvlwv zklfk kdg ehhqvdpsohg dwQjdpdndod frqwdlq- iurp 35-111 C0S0wrwkhehjlqqlqj ri wkh Krorfhqh- doprvw qr sroohq fkdèudfwhulvwlf ri wkh Biurèprxqwdlq hohphqw )FOFQIB-2993,0 Liwkh Qjdpdndod vlwh kdg ehhq wkh rqo° rqh

ij Cnqfspt jy m 32 pr fz stwi ij Cwfééf-nqqj, xj xnyzj-jwx 400 r i'fqynyzij, rfnx jxy jshfnxxp ifsx zs uqfêyjfz itsy qjx fqynyzijx -tsy ij 500 m uqzx ij 600r.+*)%1). Ojx ipusyx vzn tsy pyp hfwtyypx m Qlfrfpfqfsj uwpxjsyjsy jsywj 24.000 fsx jy qj ipgzy ij q'Ktqtêhosj vzfxnrjsy fzhzs utqqjs hfwfhypwnxynvzjij q'pqpêrjsy fkwtrtsyflsfwi (FOFQIB, 1992). Vnqj xnyj ijQlfrfpfqf f-fny pyp qj xjzq m qywj pyzinp, ts fzwfnyxfsx itzyj htshqz vzj qf wplnts ij Cwfééf-nqqj s'f-fnyufx pyp fkkjhypj -jwx qf knsiz Sqpnxythosj ufw zsfgfnxxjrjsy ij qf yjrupwfyzwj. Rs jxy itsh tgqnlpij htshqzwj vzj q'fxxthnfynts fkwtrtsyflsfwij izCtnx ij Cnqfspt, ytzy m kfnynsipsnfgqj, ij-fny qywj uwtêgfgqjrjsy qthfqnxpj fz xnyj rqrj tz ifsx zs wfètsfxxjé wpizny.

Djx in-jwxjx itsspjx, vztnvzj jshtwj ywtu ujzstrgwjzxjx, xjrgqjsy fzytwnxjw vzjqvzjx htshqzêxntsx uwpqnrnsfnwjx. Of uqzx nrutwyfsyj xjwfny vzj qjwjkwtninxxjrjsy hqnrfynvzj ij qf kns iz Sqpnxythosjs'pyfny ufx zs umpstrosj lpspwfqnxp uwoxij q'pvzfêyjzw. Fs kfny, i'zsj rfsnowj nsinwjhyj, ts f-fny ipomfgtzyn ~zsj yjqqj htshqzxnts f-jh qf rnxj js p-nijshjij wjkzljx qtwxij q'pyzij ij qf kwflrjsyfynts iz gqthktwjxynjw fkwnhfns izwfsy qjx rfànrzr lqfhnfnwjx izTzfyjwsfnwj jy js ijwsnjw qnjz -jwx 18.000 fsx CS(PBOFc, 1987, 1989, m ufwfrywj). Fs jkkjy, qf uwpêxjshj ij wjkzljx ij ktwqyij uqfnsj nruqnvzj vz'jsvzjqvzjx xjhyjzwx fny ujwxnxyp zs hqnrfy kf-twfgqj mxts rfnsynjs, jy itsh m ijx yjrupwfyzwjx vzn s'tsyufx iu ijxhjsiwj jsêijxxtzx ij 20îD htrrj -fqjzwrtèjssj fsszjqqj. Djyyj -fqjzw jxy fhyzjqqjrjsy zsrnsnrzr js ijnm izvzjq ts ufxxj ~ ijx ktwrfyntsxrtsyflsfwijx (ULDKBUEV, 1952). Ej uqzx, qj hfwfhêyowj fuufwjrrjsy qthfqnxp ij q'fxxthnfynts fkwtrtsêyflsfwij iz Ctnx ij Cnqfspt, wjounsyin-jwxjx tgxjwê-fyntsx jkkjhyzpjx ifsx qj rnqnjz' fhyzjq ~uwtutx ij qfuwpxjshj ij xyfyntsx nxtqpjx m gfxxj fqynyzij ij gntêytujx rtsyflsfwix, yjq vz'ts q'f wfuujqp hnêijxxzxutzw qj wjgtwi rpwnintsfq iz gfxxns iz Dtslt tz jsin-jwx utnsyx ijx htqqnsjx iz Pfètrgj, iz Dmfnqqzjyijx Ptsyx ij Dwnxyfq(aKLWF, 1981 APBOFc & fq.,1988), tz js i'fzywjx utnsyx ij q'Bkwnvzjywtunhfqj (htqêqnsjx ij Hwjjyt°s, Ptsy Dfrjwtzs, Sqfyjfz ijq'Bsltqf, Ptsyx Yxfrgfwf, jyh) (aKLWF, 1981 APBOFc,1989), Oj hfwfhyowj nxtqpij hjx xyfyntsx pyfnynsyjwuwpyp nruqnhnyjrjsy htrrj pyfsy ijx wjxyjxi'fnwjx ywox pyjsizjx tz lpspwfqnxpjx izwfsy qjxumfxjx ij wjkwtninxxjrjsy hqnrfynvzj (hL.CFQIR & PBOFc, 1991). Wtzyjktnx ts utzwwfnyfzxxnxj ijrfsijw xnrqrj izwfsy qjx upwntijx ij wjkwtnêinxxjrjsy rfànrzr, qjx gntytujx ij yèuj rtsyfêlsfwi sj xj ywtz-fnjsy ufx ipom ifsx ijx xyfyntsxuqzx tz rtnsx gnjs ipqnrnypjx, htrrj hjqqj ijCnqfspt, gnjs vzj hjwyfnsjrjsy uqzx strgwjzxjx jyuqzx pyjsizjx vzj hjqqjx wpujwytwnpjx fhyzjqqjrjsy.

O'mèutymoxj iz wsqjijx szfljx ifsx qjx hmfsljêrjsyx hqnrfynvzjx js wplnts ywtunhfqj jy uqzx ufwynhzêqnowjrjsy izwfsy qjx umfxjx ij wjkwtninxxjrjsy, fipom pyp f-fshpj utzw q'Bkwnvzj ufw PBOFc(1981,1982,1987,1989), PBOFc & OLZLQIVWRQF

°mnhmmfi gjjs xyzinji ny°tz Li mf-j st itzgy gjjshtshqziji ymfy ymj Cwfééf-nqqj wjlnts mfi sty gjjsytzhmji gè f iwtu nsymj yjrujwfyzwj yt°fwix ymj jsitk ymj Sqjnxythjsj. Rsj ymjwjktwj mfx yt htshqzij ymfyymj Ctnx ij Cnqfspt Bkwtrtsyfsj fxxthnfynts, vznyjzsijsnfgqj, rzxy mf-j uwtgfgqè gjjs qthfqnéji fy ymnx-jwè xnyj tw nsf -jwè wjizhji wfinzx.

Wmjxj -fwntzx ifyf, ymtzlm xynqqytt kj°, jsfgqjxtrj uwjqnrnsfwèhtshqzxntsx yt gj iwf°s. Wmj rtxynrutwyfsy tsj °tzqi gj ymfysjfw ymj Fvzfytw, ymj hqnêrfynh httqnsl fy ymj jsi tk ymj Sqjnxythjsj °fx sty fljsjwfqnéji umjstrjsts. qskfhy, nsfs nsinwjhyrfsêsjw, ymj xfrj htshqzxnts mfi fqwjfiè gjjs wjfhmji°mjs ymj jànxyjshj tk wjkzljx mfi gjjs hqjfwqèxmt°s izwnsl ymj xyziè tk ymj kwflrjsyfynts tk ymjBkwnhfsktwjxy gqthp izwnsl ymj lqfhnfq rfànrf tk ymjTzfyjwsfwè fsi nsymj qfxy nsxyfshj fy fuuwtànrfyjqè18,000 C.S. (PBOFc, 1987, 1989, yt htrj tzy). qskfhy, ymj uwjxjshj tk qt°qfsi ktwjxy wjkzljx nruqnjxymfynsf kj° xjhytwx, f hqnrfyj kf-tzwfgqj yt nyxpjjêunsl mfi ujwxnxyji, fsi ymjwjktwj °nymyjrujwfyzwjx°mnhmmfi sty ltsj gjqt° fs f-jwflj fsszfq -fqzj tk20îD. Wmnx-fqzj nxhzwwjsyqèf rnsnrzr gjqt° °mnhmrtzsyfns ktwrfyntsx thhzw (ULDKBUEV, 1952).Ptwjt-jw, ymj fuufwjsyqè qthfqnéji hmfwfhyjw tk ymjBkwtrtsyfsj fxxthnfynts tk ymj Ctnx ij Cnqfspt, nxnsflwjjrjsy °nym -fwntzx tgxjw-fyntsx hfwwnji tzy nsymj hzwwjsyrnqnjz htshjwsnsl ymj uwjxjshj tk nxtqfyjixyfyntsx tk rtzsyfns gntytujx fy qt° fqynyzij, fx xyfyjifgt-j ktw ymj xtzymjws jilj tk ymj Dtslt Cfxns tw-fwntzx utnsyx ts ymj Pfètrgj, Dmfnqqzmnqqxfsi ymjDwnxyfqPtsyx (aKLWF, 1981 APBOFcfsi fq., 1988),tw fy tymjw utnsyx ns ywtunhfqBkwnhf(Hwjjyt°s Knqqx,Ptsy Dfrjwtts, Bsltqf Sqfyjfz, Ptsyx Yxfrgfwf,jyh) (aKLWF, 1981'APBOFc, 1989). Wmj nxtqfyji hmfêwfhyjwtk ymjxj xyfyntsx °fx zsvzjxyntsnslqè nsyjwuwjêyji fx gjnsl tk °mfy wjrfnsji tk °nijxuwjfi tw ljsjêwfqnéji fwjfx °mnhmmfi thhzwwji izwnsl ymj umfxjx tkhqnrfynh httqnsl (hL. CFQIR fsi PBOFc, 1991).Kt°j-jw, tsj hfs fqxt °tsijw°mjymjw j-js nkizwnslymjxj ujwntix tk rfànrzr httqnsl, ymj rtzsyfns yèujgntytujx tsqè thhzwwji nsrtwj tw qjxx °jnq ijrfwpfêyji xyfyntsx, xzhm fx ymj tsj tk Cnqfspt, fqymtzlm ymjè°jwj hjwyfnsqèrtwj szrjwtzx fsi rtwj °nijxuwjfiymfs ymtxp hzwwjsyqèLnxyji.

Wmj mèutymjxnx tk ymj wtqj tk hqtzix nsymj hqnrfynhhmfsljx nsymj ywtunhfqwflnts, fsi nsufwynhzqfwiz wnslymj httqnsl it°s umfxjx, mfx fqwjfiè gjjs uzy ktwê°fwi ktwBkwnhfgè PBOFc (1981,1982,1987,1989),PBOFcfsi OLZLQIVWRQF (1983), CUFQBD (1988),gjnsl ktzsiji ts -fwntzx uwj-ntzx xyzinjx, jxujhnfqqèymtxj tk CBUBW (1957) htshjwsnsl ymj yèujx tk wfnsêkfqqymfyPBOFc (1981, 1982) mfi fuuqnji yt wzsêtkkxfsi yt xjinrjsyfynts uwthjxxjx, tw OBPC (1972).Dtshjwsnsl ymj vzjxynts tk yjrujwfyzwjx inxhzxxjimjwj, ymj ujwxnxyjshj tk xywfynktwwshqtzix nxfq°fèxwjxutsxngqj ktw sjlfyn-j fstrfqnjx tk ymj yjrujwfêyzwj nsymj qt° qj-jqx tk ymj fyrtxumjwj (BVSOLEFQfsi BEFHROBOY, 1976). Wmnxwtqj tk ymj hqtzix fsinsufwynhzqfwymj xywfynktwrxymjwjktwj fuujfwx yt °jnq

(1983). CUFQBD (1988), pyfsy gfxpj xzw in-jwx ywfê-fzà fsypwnjzwx, ufwynhzqnowjrjsy hjzà ij CBUBW(1957) htshjwsfsy qjx yèujx ij uqznj vzj PBOFc(1981, 1982) f fuuqnvzp fzà phtzqjrjsyx jy fzà uwtêhjxxzx ij xpinrjsyfynts, tz ij OBPC (1972).Dtshjwsfsy qf vzjxynts ijx yjrupwfyzwjx inxhzypjnhn,qf ujwxnxyfshj ijx szfljx xywfynktwrjx jxy ytz';otzwx wjxutsxfgqj i'fstrfqnjx splfyn-jx ij qf yjr%upwfyzwj ifsx qjx gfxxjx htzhmjx ij q'fyrtxumowj(B8SOqEFQ & BEFHROBOY, 1976). Dj wsqj ijxszfljx jy ufwynhzqnowjrjsy ijx xywfynktwrjx xjrgqjitsh gnjs x'fuuqnvzjw fz hfwfhyowj inxhtsynsz uwqypfzà jàyjsxntsx ijx gntytujx rtsyflsfwix m gfxxjfqynyzij. Bhyzjqqjrjsy ts wjshtsywj ufwktnx xtzx qjxywtunvzjx ij yjqqjx jàyjsxntsx rtsyflsfwijx qthfqnêxpjx ozxvzj -jwx 500 r i'fqynyzij jy ufwktnx ozxvzj-jwx 300 r (Bsltqf), sts xjzqjrjsqy's Bkwnvzj, rfnxfzxxn js Brpwnvzj jy js Bxnj, xtny xzwnj kqfsh ij rtsêyflsjx nrutwyfsyjx htrrj qj Ptsy Dfrjwtzs(VFUOF, 1964 AWKRPBV, 1986) tz qj kqfsh ijlwfsix uqfyjfzà htrrj js Bsltqf (BLUcêVKBa,1947). xtny fzxxn xzw qj xtrrjy ij htqqnsjx ujz pqjê-pjx, ufw jkkjy ij hwqyj. Djy jkkjy jxy iu fzà szfljxvzn %x'fhhwthmjsy%'jy ujwxnxyjsy uqzx qtslyjrux xzwijx xtrrjyx ij htqqnsjx, rqrj xnhjqqjxêhnsj hzqrnêsjsy vz'm 500r i'fqynyzij, iz rtrjsy vzj hjx xtrêrjyx itrnsjsy zsj uqfnsj js-nwtssfsyj.

E'zsj rfsnowj uqzx lpspwfqj, zs yjq umpstrosj fpyp iphwny ifsx hj vz'ts fuujqqj qjx ktwqyx ijgwtznqqfwi tz ij szflj jy vzn wpxzqyjsy ij qf ujwxnxêyfshj ijx gwtznqqfwixtz ijx htz-jwyzwjx szfljzxjxxywfynktwrjx (NFUHRRW, 1968 A IUYCC, 1971,197dA aKLWPRUF, 1975 ADBZFOLFU & IROEêVWFLQ, 1989). Bhyzjqqjrjsy, hj umpstrosj ufwfryuqzx kwpvzjsy ifsx qjx wplntsx uwthmjx ij qf rjw(PRUFBY, 1966 A CBcQWRQ, 1968 A KRaBUE,1970A VYIEFQ, 1982A jyh). Wtzyjktnx ijx htz-jwêyzwjx szfljzxjx xywfynktwrjx x'pyjsijsy ufwktnx qtnsijx hsyjx fz Ifgts jy fz Dtslt, ozxvz'm 800 m1.000 pr jy uqzx ufwktnx (VBLQWZLO, 1977, 1979 Ahk.PBOFc, 1990) .+*)%q, (*(12 A+*)%&'f%ôyfsy itssp vzjhjyyj jàyjsxnts m %nsypwnjzwiz htsynsjsy nsyjw-njsyizwfsy qf xfnxts kwfrhmj fsszjqqj jy jxy fxxthnpj jsufwynhzqnjw~ ijx wjrtsypjx i'jfz kwtnij (zu°jqqnslx)m qf xzwkfhj ij q'Rhpfs Byqfsynvzj -tnxns(PBKF & DLWFBY, 1993), ts ujzy itsh jxynrjw vzjhjx umpstrosjx tsy iu qywj ywoxfruqnknpx izwfsy qjxlwfsijx umfxjx ij wjkwtninxxjrjsy hqnrfynvzj izTzfyjwsfnwj (PBOFc, 1987, 1989). En-jwxjx pyzijxrnhwtufqptsytqtlnvzjx rjspjx xzwijx xpinrjsyx izItqkj ij Iznspj tsy rtsywp vzj hjx wjrtsypjx i'jfzkwtnij tsy pypywoxnsyjsxjx -jwx 18.000 fsx CS, f-jh,ufw wfuutwy ~ q'Bhyzjq, ijx fgfnxxjrjsyx ij qf yjrupêwfyzwj ijx jfzà ij xzwkfhj ij 4î ~ 8îD (SUFOO & fq.,1976A PLb & fq., 1986A hk. PBOFc, 1987, 1989). Vzwqj htsynsjsy, qf ytutlwfumnj (ILREB & fq., xtzxuwjxxj) jy in-jwx fzywjx kfhyjzwx lptlwfumnvzjx(pyjsizjx rfwphfljzxjx, kqjz-jx, jyh) otzjsy fzxxnijx wsqjx nrutwyfsyx js rtizqfsy qjx xjhyjzwx tt qjxszfljx gfx jy qjx gwtznqqfwix utzwwtsy, xtny ujwxnxyjw

fuuqè yt ymj inxhtsynsztzx hmfwfhyjw fyywngzyji yt ymjjàyjsxntsx tk rtzsyfns gntytujx fy qt° fqynyzij. Dzwêwjsyqèns ymj Wwtunhxtsj hfs xtrjynrjx jshtzsyjwxzhm qthfqnéji rtzsyfns jàyjsxntsx fx kfw fx 500rfqynyzij fsi xtrjynrjx fx kfw fx 300r (Bsltqf), stytsqè nsBkwnhf,gzy fqxt nsBrjwnhf fsi nsBxnf, jnymjwts ymj xnij tk nrutwyfsy rtzsyfnsx xzhm fx PtsyDfrjwtts (VFUOF, 1964 AWKRPBV, 1986) tw ts ymjxnij tk lwjfy uqfyjfz xzhm fx nsBsltqf (BLUc êVKBa,1947), gzy fqxt ts ymj ytu tk sty -jwè mnlmmnqqx,izj ytwnilj jkkjhy. Wmnxjkkjhy nxizj yt h10zix °mnhm%h1nsl%fsi qfxy qtsljw t-jw ymj ytux tk mnnqx,j-js nkymjnwmnlmjxy utnsy nxst rtwj ymfs 500 r mnlm,fx qtsl fxymjxj ytux itrnsfyj f xzwwtzsinsl uqfns.

qs f rtwj ljsjwfq rfssjw, xzhm f umjstrjstsmfx gjjs i~xhwngji ns ~mfy nx hfqqji ktl tw hqtziktwjxyx fsi nQmnhmwjxzqyxkwtr ymj ujwxnxyjshj tk ktlxtw xywfynktwr hqtzi ht-jwx (NFUHRRW, 1968 AIUYCC, 1971, 1977 AaKLWPRUF, 1975 ADBZFêOLFUfsi IROEVWFLQ, 1989). Dzwwjsyqè,ymnxumjstêrjsts fuujfwx rtwj kwjvzjsyqè ns wjlntsx sjfw ymjxjf (PRUFBY, 1966A CBcQWRQ, 1968A KRaBUE,1970A VYIEFQ, 1982 Ajyh). Qj-jwymjqjxx, xywfynktwrhqtzi ht-jwx xtrjynrjx xuwjfi f qtsl °fè kwtr ymjIfgts fsi Dtslt htfxyx, zu yt 800 yt 1.000 prx fsixtrjynrjx rtwj (VBLQW ZLO, 1977, 1979 A hk.PBOFc, 1990) .+*)%q,(*(12A+*)%&i'f% Vnshj ymnxnsqfsijàyjsxnts thhzwx izwnsl ymj fsszfq httq xjfxts fsinxnsufwynhzqfwfxxthnfyji °nymzu°jqqnslx fy ymj xzwêkfhj tk ymj sjnlmgtzwnsl ByqfsynhRhjfs (PBKF fsiDLWFBY, 1993), tsj hfs ymjwjktwj jxynrfyj ymfyymjxjumjstrjsf rzxy mf-j gjjs -jwè rzhm fruqnkqjiizwnsl ymj lwjfy umfxjx tk Tzfyjwsfwè h1nrfynhhttqnsl (PBOFc, 1987, 1989). Zfwntzx rnhwtufqjtsêytqtlnhfq xyzvnjx hfwwnji tzy ts xjinrjsyx kwtr ymjIznk tk Iznsjf mf-j xmt°s ymfy ymjxj zu°jqqnslx°jwj -jwè nsyjqexj fy fwtzsi 18,000 C.S., °nym,nshtrêufwnxts °nymymj uwjxjsy ifè, iwtux nsyjrujwfyzwj tkymj xzwkfhj °fyjwx nsymj wfslj tk 4î yt 8îD (SUFOOfsi fq., 1976 APLb fsi fq., 1986 APBOFc, 1987,1989). Rs ymj htsynsjsy, ymj ytutlwfumè (ILREB fsifq., ns uwjxx) fsi -fwntzx tymjw ljtlwfumnh kfhytwx(rfwxmqfsi'x xywjyhmjx, wn-jwx, jyh) fqxt uqfè nrutwêyfsy wtqjx ns rtizqfynsl ymj xjhytwx °mjwj ymj qt°h10zix fsi ktlx hfs jnymjwujwxnxynsf wjlzqfw rfssjw,tw, ts ymj htsywfwè, hfs wfuniqè inxxnufyj. qs ymj knwxyhfxj, rtzsyfns gntytujx fwj fgqj yt ij-jqtu ,(ktwjàfruqj, Cnqfspt) fsi nsymj xjhtsi hfxj, nsxjhytwxsjfw yt ymj knwxy(ktwjàfruqj, Qlfrfpfqf) qt° fqynyzijgntytujx °nqqujwxnxy.Rsj °nqqymjwjktwj tgyfns f qfsiêxhfuj rfij zu tk f rtxfnh tk -jwè inkkjwjsy js-nwtsêrjsyx.

',. qstwijw yt nLLzxywfyjymj wtqj tk h1tzix, L°t Pjyjtêxfy nrfljx fwj uwjxjsyji, xmt°nsl, ktw y°t wjhjsyèjfwx, L°t tuutxnyj xnyzfyntsx. Wmj tsj tk Mzqè1983.+*)%&'fg xmt°x f xywtsl ij-jqturjsy tk xywfynktwrhqtzi ht-jwx nsymj -nhnsnyètkymj Iznsjf Izqk, °mnqj tsknlzwj 2 C (Mzqè1984), ymjè fuujfw rzhm rtwj wjizêhji, °nymymj fuufwjsy ufwfità ymfyymj 1984 xjfxts°fx ts ymj htsywfwè hqjfwqèrtwj wfnsèymfs 1983, izj

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AKTa°UIAZ I. M, 1947. ° Vif wfhfubujpo pgAohpmb.L. C; '8C@C<L. 35, 23°48.

AURNKDEP C.1.& ADEFQNANW Q., 1976. ° Vif nfbs uspqptqifsf 01Zftu Agsjdb. L. gDD@f ,:H:C. 1 5,705°716.

BATAV C., 1957. ° Rmvwjpmphjf:H brvjejnêusjf ebot mb-pof joufsuspqjdbmf. ,PAC>F: *BGH>HIH (F6BN6>G g;F>EI: -C>F:.Dblbs, 49, %~qq.

BAaPVQP IZ., 1968. ° Vif fdpmphz 01 bo fmmjogpsftu jo Rvfsup Tjdp. 2. Vif njdspdmjnbuf pg Rjdp efj Qftuf. L. C;gFBC@9 gF7CFf. 49, 419°430.

BEPHQ O.D. & OANEa, L., 1991. °Aobmztft eft gmvyqpmmjojrvft tvs mbnbshf tve ev Hpmmfef Hvjoêf efqvjt 135.000bot. %f0fg869f 18f. Rbsjt, têsjf 2, 313, 843°849.

BQPPEFKNNE T. & TKQNNEV,H., 1988. ° Vif Mbtijsv qpmmfotfrvfodf (Bvsvoej), Rbmbfpdmjnbujdjnqmjdbujpot gpsuifmbtu40,000 zs BR jo uspqjdbmAgsjdb. /I6*f 0:Gf. 30, 19°35.

BQPPEFKNNE T., CIAmmE, F., HWKQV,L. & YKPCEPU, A., 1992. °Svboujubu'jwf ftujnbufl01 mvmmhmbdjbmufnqfsbuvsft jofrvbupsjbm Agsjdbmspnqbmzopmphjdbmebub. %;>A6H: &LB6A>8G. 6, 251°257.

BTEPAC R., 1988. ° Ewpmvujpoef mbwêhêubujpo fu ev dmjnbuebot m'Qvftu Cbnfspvo fousf 25.000 fu 11.000 bot BR.g8H:G 5( 1LADCG>IA Att. .6@LBC@C<I:G +6B<I: (F6B86>G:. Vsbw. Uffu. Ufj. & Vfdi.motu. Fsboèbjt Rpoejdiêsz,25,91°103.

CAYENKET L. & HQNDUVEKP, H., 1989. ° Ojtu boe gph joufsdfqujpo jo fmmjodmpvempsftut jo Cpmpncjb boe Yfof-vfmb. L.2FCDf '8C@C<L. 5, 309°322.

CNjOAR RspkfduOfncfst, 1981. ° Ufbtpobm sfdpotusvdujpo pguif Ebsui't tvsgbdf buuif Nbtu HmbdjbmObyjnvn. ):6*f1C8f gA:F>86. ,6D & %=6FH 1:F>:G. OC°36.

EKUEPVTAWV O., 1970. ° Ejt-fjulmjnb voe ifvujhf Vjfswfscsfjuvoh jn uspqjtdifo Zftubmsjlb. 3AG8=6IZjtt. & 2:8=Bf. (F6B?;IFH 6f,f. 3, 70°75.

ENEPHA I., 1987. ° mft Rmbufbvy Bbuêlê (Cpohp). Rbmêpfowjspoofnfout Svbufsobjsft e'bqsét m'êuvef ev tpoebhfev cpjt ef Bjmbolp. &>D@QA: 9('HI9:G gDDFC;CB9>:G. Wojwfstjuê e'Ajy°Obstfjmmf KKK,41 q.

EmEPHA I., 1992. ° Yêhêubujpo fu dmjnbuev Cpohp efqvjt 24.000 bot BR. Aobmztf qbmzopmphjrvf 9: têrvfodft têej°nfoubjsft ev qbzt Bbuêlê fu ev mjuupsbm.2=OG: 18>:B8:. Wojwfstjuê Ajy~Obstfjmjf KKK,238 q.

EmEPHA I. & YKPCEPU, A., 1990. ° Rbmêpfowjupoofnfout rvbufsobjsft sêdfout eft RmbufbvyBbuêlê (Cpohp) ; êuvefqbmzopmphzrvf eft eêqîut ef mbef qsfttjpo ef Bjmbolp. moT. mbogsbod.ij & D. Udixbsu- (fet), .6LG6<:G /I6H:FMB6>F:G 9: @(g;F>EI: 8:BHF6@: gH@6BH>EI:. Djebdujrvft, QTUVQO, 271°282.

HKQDA A., OAmEa, L., EURELQ, H.T. & ACQUVA BAmADQP, A., 1993. ° Vif gpvubjousff 01Cbobsz jtmboet, boe puifsmphdpmmfdupst; qbtu boe DF:G:BH fybnqmft pgmpxnpvoubjo dmpvempsftut. 13 q. (nbovtdsju tpvnjt).

HTWBB R.L., 1971. °Koufsqsfubujpo pmuif %Obttfofsifcvoh% :;@:8H po uspqjdbmnpvoubjot. -6HIF:. 229, 44°45.

HTWBB R.L., 1977. ° Cpouspm01 gpsftu hspxui boe ejtusjcvujpo po K:H uspqjdbmnpvoubjot, xjui tqfdjbm F:@:F:B8: upnjofsbj ovusjujpo.gBBf 0:Jf Edpm.Uztu., 8, 83°1 07.

HWjmmQVB., 1985. Vfnqêsbuvsf ef tvsgbdf ef mbnfs fu qmvwjptjuêbvupvs ev Hpmgfef Hvjoêf, 4:>;;: %@>A6H>EI: 16H:@M@>H6>F:. Pà 6, qq. 13°14.

IANN L.B., 1973. ° Yfhfubujpobm -poft po uif tpvuifso tmpqft 01Opvou Cbnfsppo. 4:<:H6H>C. 27, 49°69.

IAOKNVQP A.C., 1982. ° Eowjsponfoubm ijtupsz 01Ebtu Agsjdb. A tuvez 01uif Svbufsobsz. Adbefnjf Rsftt, Npoepo,311 qq.

IQZATD TA, 1970. ° Vif %Amqjof%qmbout01uif Aoujmmft.i>CHFCD>86. 2, 24°28.

LAPKCQV U. & FQPVAKPE, B., 1993. ° N'êwpmvujpoeft jeêft tvs mbwbsjbcjmjuêjoufsboovfmmf FP8:BH: eft qsêdjqjubujpotfo Agsjrvf ef m'Qvftu. +6 ,PHPCFC@C<>:. Uêsjf 8,1,28°53.

METFQQV 0.,1968. ° Ojtu qsfdjqjubujpo po wfhfubujpo. (CF:GHFL g7GHF68HG. 29, 8°20.

MTAWU E.B., 1973. ° Cpnqbsjtpo 7:HK::B KdfAhf boe qsftfou hfofsbm djsdvmbujpot. -6HIF:. 245 ;1 29°1 33.

mAOB I.I., 1972. ° Cmjnbuf; qsftfou, qbtu boe gvuvsf. m,Fvoebnfoubmt boe dmjnbuf opx. Ofuivfo, Npoepo, 613 q.

Fjhvsf 3. ° 43û493 693 49ou8û345û93 t483sb°593 69 35û>349 ut5û g93 u8ûbt693 4tûû93uts763s4 356 bo3A93 o848t334 69 g3 >bA5û9 ,!4N3û43 t> 4N9 t493sb4 35û>349 49ou9û345û93 >tû 4N9 u9ûbt63 4tûû93uts6bsA4t o949t334 bo3A93! (Uff gjhvsft ,3 boe 28).

OAIE H. & CKUEAV, L., 1993. à SgncvkqpuQegcp àAvoqurjétg àCqpvkpgpv fcpu n'gurceg Ahtkeckpfg nc oqwuuqpAvncpvkswg.Tejêoc iêpêtcn gvecu rctvkewnkgtfg 1984. ;@DEE@ +EDFg 8<M@igf 44, 34 à54.

OANEa L., 1981. à Evwfgu rcn-pqnqikswgu fcpu ngdcuukp fw Uejcf gvrcnêqenkocvqnqikg fg n'Ahtkswgpqtfàvtqrkecng fg30.000 cpu %n'êrqswg cevwgnng.9K<Og& ,H>NF@GML 478 942f Rctku, 129, 586 rr.

OANEa L., 1982. à Dwuv,enqwfu, tckp v-rgu cpf enkocvkexctkcvkqpukpvtqrkecn Pqtvj Ahtkec. 6N<Mg 7@Lgf 18, 1à16.

OANEa L., 1987. à Ftciogpvcvkqp fg nchqtîv fgpug jwokfg chtkeckpggvgzvgpukqp fgu dkqvqrgu oqpvcipctfu cw Qwcàvgtpcktg têegpv; pqwxgnngufqppêgu rqnnkpkswgugv ejtqpqnqikswgu. Kornkecvkqpurcnêqenkocvkswgu gv dkqiêqitcàrjkswgu. 5<E<@H@>HEHBP HA)AKD><f18, 307à334.

OANEa L., 1989. à NcvgQwcvgtpct- enkocvkeejcpigu kpvjg Ahtkecp tckp hqtguv; hqtguv tghwikc cpf vjg oclqt tqngqhugcuwthceg vgorgtcvwtg xctkcvkqpu. npO. Ngkpgp & O. Tctpvjgkp (gfu.), 5<i@H>EDF<MHEHBP <G? 5<E@HF@M@HKHEHBP (2H?@KG <G? I<LM I<MM@KGL HABEH=<E <MFHLIC@KD> MK<GLIHKMfMnwygtAecf. Rwd~.,Dqtftgejv, 2,43 Afxcpegf Te.19>6!Tgtkgu. C. Ocvj. & Rj-u. Te., 282, 585à616.

OANEa L., 1990. à N'jkuvqktg têegpvg fg nchqtîv fgpug jwokfg chtkeckpg; guuck uwt ngf-pcokuog fg swgnswguhqtocàvkqpuhqtguvkétgu. npS. Ncphtcpejk & D. Tejyctv° (gfu), 5<PL<B@L 6N<M@KG<DK@L ?@ E.)AKDJN@>@GMK<E@ )Mi<GMDJN@fDkfcevkswgu, QSTUQO, 367à382.

UIQOAT D.Y., 1986. à Wgigvcvkqp kpvjg oqpvcpg hqtguvuqhCcogtqqp. kpT.P.Tvwctv (gf) +HGL@KO<MDHG HA+<F@KHHG2HGM<G@ .HK@LMLf Sgrqtv qhKpv.CqwpeknBktf Rtgugtx .. Ccodtkfig, VM, rr. 20à27.

WAP bKPDESEP BAMMES E.O. & CQEUbEE LA. 1988. à A tgxkgy qhncvgQwcvgtpct- rqnngp uvwfkgu kpEcuv, Cgpvtcncpf Tqwvjgtp Ahtkec. 7@Og 5<E<@H=HMg & 5<EPGHgf 55, 155à174

WKPCEPT A, CIAnnE F., BQPPEFKNNE S., HVKQU L. & UKESCENKPL.L., 1993. à Rqnngpàfgtkxgf tckphcnncpf vgorgtcàvwtgguvkocvgu htqo NcmgUcpicp-kmc cpf vjgktktjrnkecvkqp hqt Ncvg Rngkuvqegpgycvgt ngxgnu.6N<Mg 7@Lg 40 (uqwurtguug).

YANUES I. & BSECMNE T,Y., 1985. à Eeqnqikecn u-uvgou qhvjg Hgqdkqurjgtg., 1, Eeqnqikecn rtkpekrngu kpinqdcnrgturgevkxg. TrtkpigtàWgtnci, Bgtnkp.

YEBTUES R.L. & TUSEUEP PA, 1978. à Ncvg Qwcvgtpct- KegAig enkocvgu qhvtqrkecn Awuvtcncukc ; kpvgtrtgvcvkqpucpf tgeqpuvtwevkqpu. 6N<Mg 7@Lgf 10, 279à309

YIKUE F., 1981. à Ujg jkuvqt- qhvjg Ahtqoqpvcpg ctejkrgnciq cpf vjg uekgpvkhkepggf hqt kvueqpugtxcvkqp. )AKg1->HQEHBPf 19,33à54.

YIKUOQSE U.C., 1975. à Utqrkecn tckp hqtguvuqhvjg Fct Ecuv. Cnctgpfqp Rtguu, Qzhqtf.

YIKUOQSE U.C., FNEPNEa L.S. & IASSKT D.S .. 1982. à Ujg vtqrkeu cu vjg pqto kpdk'ekigqitcrj- ~ /@HBKg 1f %&'f8à21.

OANEa L., 1991. à Ujg Ahtkecp tckp hqtguv xgigvcvkqp cpf rcncgqgpxktqpogpvu fwtkpi ncvgQwcvgtpct-. +EDF<MD>+C<GB@f 19, 79à98.

OANEa L. & NKWKPHTUQPE, DA, 1983. à Ezvgpukqp f'wp ênêogpv oqpvcipctf fcpu nguwf fw Hjcpc (Ahtkswgfgn'Qwguv)cw Rnêkuvqeépguwrêtkgwt gv%n'Iqnqeépg kphêtkgwt;rtgokétgu fqppêgu rqnnkpkswgu.+g7g )><?g 8>gf uêtkg2,296,1287à1292.

OANEa L., CABANNE, H. & TKUA,R., 1990. à Evwfg f'wp rgwrngogpv têukfwgnè dcuug cnvkvwfgfg 5H?H><KINL E<MDAHEDNLuwt nghncpeCqpiqncku fw Ocuukhfw Cjcknnw.Kornkecvkqpurcnêqenkocvkswgugvdkqiêqitcrjkswgu. Evwfg fg ncrnwkgrqnnkpkswgcevwgnng.npS. Ncphtcpejk & D. Tejyctv° (gfu), 5<PL<B@L 4N<M@KG<DK@L ?@ E.)AKDJN@>@GMK<E@ )Mi<GMDJN@fDkfcevkswgu. QSTUQO, 336à352.

OKZ AC., SVDDKOAP, Y.F.gv OCKPUaSE, A.1986. à NcvgQwcvgtpct- rcngqegcpqitcrj- qhvjg vtqrkecn Avncpvke.2 ;Ujg ugcuqpcn e-eng qhugc uwthceg vgorgtcvwtgu, 0à20,000 -gctu B.R. 5<i@H>@<GHBK<ICPf 1,339à353.

OQSEAV S.E., 1966. à Ujg dktf hcwpcu qhAhtkeccpf kvukuncpfu. Aecfgokg Rtguu, Pgyàaqtm,424 rr.

OQSNEa S.L. & FNEPNEa, L.S., 1987. à NcvgCckpq°qke xgigvcvkqpcn cpf gpxktqpogpvcn ejcpigu kpvjg Ocnc- ctejkàrgnciq. kpU.C'yjkvoqtg (gf), *DHB@HBK<ICD><E -OHENMDHG HA MC@ 2<E<P )K>CDI@E<BHg Cnctgpfqp Rtguu, Qzhqtf,rr.50à59.

PEYENN S.E., IESOAP, H.F., HQVNDàTUEYASD, T. & UAPAMA, O., 1975. à Dgetgcugf inqdcn tckphcnnfwtkpi vjgrcuv KegAig. 3<MNK@f 253 ; 33à34.

RSENN Y.N., HASDPES, L.W., BE, AY.I. & IAaT, L.D., 1976. à EswcvqtkcnAvncpvkecpf Cctkddgcp hqtcokpkhgtcncuugodncigu, vgorgtcvwtgu cpf ektewncvkqp; kpvgtincekcncpf incekcn eqorctkuqpu. /@(BD><E 8H>g )F@KD><2@FHDKf 145, 247à266.

SKCIASDT R'y., 1952. à Ujg vtqrkecn tckp hqtguv.Ap geqnqikecnuvwf-. Ccodtkfig Vpkx. Rtguu, 450 r.

SKPD D. & REUEEU,D., 1985. à Ugttguvtkcneqpfkvkqpu cvvjg NcuvHncekcnOczkowo cpf CNkOAR ugc uwthcegvgorgtcàvwtgguvkocvgu ctg vjg- eqpukuvgpv. 6N<M 7@Lgf 24, 1à22.

SQTQWKTU H., 1990. à Qwcvgtpct- incekcvkqpu kpAhtkec. 6N<0g 8>g 7@Ogf 9 ; 281à297.

TAKPUàWKNL., 1977. à Nguenkocvufw Hcdqp. )GGg :GDOg3<Mg /<=HGf 1,101à125.

TAKPUàWKNL., 1979. à Nc itcpfg uckuqp uéejg cw Hcdqp. Scrrqtv 19>6!Hêqit. Utqr% Vpkx. Adkflcp.

TCIYASUb D., 1988. à Tqog rqf°qnu qp Bcvgmgucpfu cpf vjgkt qtkikpu kpCqpiq. /@H?@KF<f 43, 2 29à 247.

TESNE Y., 1964. à Ujg nqygt cnvkvwfkpcnnkokvqhvjg oqpvcpg hqtguv dktfu qhvjg Ccogtqqp oqwpvckpu, Yguv Ahtkec.*NEEg*KDMDLC4KGDMCHg Cnwd,84, 87à91.

TVHDEP A.O., 1982. à Ujg xgigvcvkqp qhvjg Tgttcpkc fg Ocewktc, Hwclktc, Cqnqodkc; c eqpvtcuv qhctkf nqyncpfu cpfcp kuqncvgfenqwf hqtguv.1 )KGHE? )K=gf63,1à30.