Benthic invertebrate fauna of high mountain streams in the Caucasus

A C T A U N I V E R S I T A T I S L O D Z I E N S I S

POL I A C A U C A S I C A 2, 1987

A n d r ze j K o w n a c ki

BENTHIC INVERTEBRATE F A U N A O F H IG H M O U N T A IN STREAMS IN THE C AU CA SU S

1. HISTORICAL OUTLINES OF FAUNI STIC INVESTIGATIONS IN THE WATERS OF THE CAUCASUS

T h e C au ca su s a n d th e T r a n sc a u c a s ia n a re a b e lo n g to th e m o st in te r e stin g te r r ito r ie s of th e USSR as for as n a tu r e in c o n c e rn e d . O n a r e l a tiv e ly sm all a r e a a g r e a t v a r i e ty of la n d sc a p e for m s a r e fo un d , s t a r -ting fro m th e sn o w -c ap p ed to p s of th e G r e a t C a u c asu s (P hoto 1) to th e su b -tro p ic al lo w lan d s of K o lch id a and L en k o ran . T h e r e a l s o o cc u r a l l

P h o t o 2. G la c ie r s tr e a m

k in d s of a q u a tic e n v iro n m e n ts: sp rin g s, o ften m in e ra l on es, tu rb id g la -cial stre a m s (Ph oto 2), n u m e ro u s ri v e r s of w h ich th e R iv e r K u ra is th e b ig g est (P hoto 3), m o u n ta in lak es, th e e n o r m o u s L a k e S ev an, s a lty la ke s, se a s id e m arsh es, an d fin a lly th e B lack an d th e C a sp ia n Seas. A p a r t from n a tu r a l a g u a tic b io to p e s a n u m b er of r e se r v o irs , ca n als, a n d fish p o n d s w e r e b u ilt th e re . T h e se w a te r s a r e in h a b ite d b y a v e r y ric h fa u n istic w o rld.

T h e fir st m en tio n s on th e C a u c a su s fre sh w a te r f a u n a c a n b e a lr e a d y fou nd in a n c ie n t w o rk s (H erod ot, Elian), th o se of A r ab ic g e o g r a p h e r s (A lIstachri, Ib nalF akich ), A rm e n ia n h isto r ia n s (K a g an k atv a ci) an d l a -te r in th e r e p o rts of R u ssia n tr a v e lle rs. In th e se p a p e r s in fo rm atio n is fo u n d m o stly on o c c u re n c e o f som e fish sp ec ie s in th e C a u c a su s w a te rs a n d th e T ra n s c a u c a s ia n a r e a . In th e 19th c e n tu r y , th e C au ca su s w as v is ite d b y n u m e ro u s n a tu r a lis ts; th e ir p a p e r s c o n ta in , am o ng o th e rs, d a ta on a q u a tic o rg an ism s a n d fish sp ec ie s of th a t re g io n .

P h o to 3. K u ra R iv e r in th e T b ili s i

S y ste m a tic h y d r o b io lo g ic al in v e stig a tio n s c a r r ie d o u t in th e C a u -c a su s b eg an in 1912 w h e n p ro f esso r A. N. D e rz h av in o rg a n ize d I-c h th y o lo - g ic al L ab o r a to ry in Baku. P ro fesso r D erz h av in w a s also th e o r g a -n ize r of th e S ev a -n H y d ro b io lo g ica l S ta tio -n (1923), o -n e of th e o r g a -n i-zers of th e Z oo log ical In stitu te of th e A z er b a ija n A ca d e m y of S ciences, an d fo r m an y y e a r s th e h e a d of th e L a b o r a to ry of H y d ro b io lo g y an d Ic h th y o lo g y in th a t In stitu te . H is scien tific ac tiv ity , lik e th a t of th e m ajo r ity of p io n e er s of h y d ro b io lo g y , w as m an y-sid ed . H e w as a n o u tsta n d in g sy ste m a tist, e x p e r t in c ru s ta c e a (C um a cea, A m p h ip o d a )

of w h ich he d escrib ed a n u m b e r of sp ec ie s from th e T ra n sc a u c a sia n a re a ; h e w as also an em in e n t ich th y o lo g ist. H e in itia te d co m p lex in v e s -tig a tio n s of th e lo w e r c o u rse of th e R iv er K ura. A t p r e s e n t his h y d ro - b io lo g ic al a c tiv ity is c a r ri e d on b y h is pu pils, am o ng o th e r s p ro fesso r A. G. K asy m o v. T o g e th e r w ith a la rg e te am of sc ie n tists th e y c o n ti -n u e th e ic h th y o lo g ic al a -n d h y d r o b io lo g ic al i-n v estig atio -n s i-n itiate d b y p ro fe sso r D e rz h av in in th e lo w e r R iv er K ura an d in th e two dam re se r v o ir i.e. th e M in gie cha urslci an d th e V a r v ar y n s k i, b u ilt on it; th e y a lso c a rr y in v e stig a tio n s in lak es a n d stream s. R e ce n tly in v e s-tig atio n s h av e b e e n fo c u ssed on th e in flu e n ce of p e tro l p o llu tio n s on th e b io cen o ses of th e C a sp ia n Sea.

T h e b e g in n in g s of h y d r o b io lo g y in G e o rg ia d a te b a sk to th e th ir -ties' of o u r c e n tu r y a n d a r e c o n n ec te d w ith th e n am e of p ro fesso r V. N. N ik ity n . In 1931, p r o f esso r N ik ity n e sta b lish e d th e H y d ro b io lo - g ic al L a b o r a to ry a t th e B atum i V N IR O S ta tio n (A ll-U nion S cien tific I n stitu te of F is h e ry a n d O ce a n o g ra p h y ) . T h e n h e m o ved to Tbilisi w h e re h e fo u n d e d th e H y d ro b io lo g ic al-Ic h th y o lo g ica l L a b o r a to r y at th e G e o rg ian S ection of th e A c a d em y of S cie nce s a n d th e n b eca m e th e h ea d of th e D e p a rtm e n t of Z o olog y of th e U n iv e r sity of Tbilisi w h e re h e le c tu r e d a lso on h y d ro b io lo g y . T h e H y d ro b io lo g ic a l- Ich th y o lo - g ica l L a b o ra to ry (of th e I n s titu te of Z o o lo g y of t h e G e o rg ia n A c a d em y of S ciences), fo r m an y y e a r s h ea d e d b y p ro fe sso r A. S ad o w -sk i — a P o le b y o rig in — c a rr ie d o u t in v e stig a tio n s in th e u p p e r an d m id dle c o u r se of th e R iv er K u ra a n d its C a u c a sia n trib u ta r ies , in m o u n ta in lak es, in m o u n ta in a n d su b m o u n ta in d am re se rv o irs. T h e G e o r -g ia n D ivisio n V N IR O in B atum i c o n c e n tra te s its in te r e st m ain ly of fish pon ds.

In th e A rm e n ian SSR th e h y d ro b io lo g ical sta tio n on th e L ake S ev an is of g r e a t e s t im p o rta nce . T h e w o r k s of its staff o n th e u n iq u e flo ra an d fa u n a of th a t lak e a r e w o rld k n ow n .

In N o r th e r n O setia, in O rd zh o n ik id ze, th e H y d ro b io lo g ic a l S ta tion of th e A g ric u ltu ra l In s titu te , d ir e c te d b y p ro fe sso r D. A. T arn o g rad sk i, h as b e e n c a rr y in g scien tific a c tiv ity for m a n y y ea r s.

A s a r e su lt of a c tiv itie s c a r rie d o u t b y th e a b o v e m e n tio n e d h y d r o -b io lo g ic al c e n te rs an d d u e to S o v iet re se ar c h es , am o ng the m p r o f e s-so r s V. I. Z hadin, A. L. B ening, K. R. F o r tu n a to v , 2319 sp ec ies and fo rm s of fre sh w a te r fa u n a w e re re c o rd e d in th e w a te r s of the te r r :- to rie s b e y o n d th e C a u c a su s (K a s y m о v 1972).

2. FAUN ISTT C C H A R A C T E R IS T IC S O F H IG H M O U N T A IN STR E AM S

A v e r y in te r e stin g fau n istic w o rld in h a b its th e m o u n tain stre am s an d riv er s of th e C au ca su s. E la b o ra tio n of th e b e n th ic fa u n a of the R iv er B a k u rian k a m ad e b y M u r v a n i d z e (1949) w a s a classica l one. T he p a p e rs b y B e n i n g and P o p o v a (1947) g iv e a h y d ro b io -lo g ic a l e la b o ra tio n of th e R iv er Z ang a from its sp rin g s to Eriv.:n. K a k a u r i d z e (1946) g iv e s a d e sc rip tio n of the b en th ic fauna of th e stre am D ab ąkh an , w h ich cr o sse s th e b o ta n ic a l g a r d e n in Tbilisi. S o m e m in o r m en tio n s o n th e b e n th ic fa u n a in so m e stre am s of th e G re a t C au c asu s can be fo u n d in p r o fe ss o r D. A. T a r n o g ra d s k i's algo- lo gical p u b lica tio n s ( T a r n o g r a d s k i , P o p o v 1932/1933). In his

stu d y d e alin g w ith fr es h w a te r fa u n a of th e C au ca su s K a s y m o v (1972) p r e se n ts th e re su lts of in v e stig a tio n s c a r r ie d ou t in 29 riv e r s an d stre a m s of th e G re a t C au c a su s a n d S m all C au ca su s in th e ir lo w er a n d m id dle co u rse. Z o s i d z e (1973), K o w n a c k i , Z o s i d z e (1980) g iv e in th e ir p a p e r s c h a r a c te r istic s of th e b en th ic fa u n a of the A d zh ar ASSR r iv e r s a n d stre am s, w h e re a s, K o w n a c k a , K o w -n a c k i (1Э71) e la b o r a t e d su c h c h a r a c te r istic s of so m e stre a m s of th e G r e at C a u c a su s in th e K azb ek a n d S v a n e ts k iy reg io n s. N e v e rth e le ss, th e C au c asu s fa u n a of stre a m s a n d r iv e r s h a s n o t b ee n fu lly d e te rm i-n ed as y e t, a i-n d r e q u ir e s f u rth e r ii-n v e stig atio i-n s.

In 1970 h y d ro b io lo g ica l in v e stig a tio n s w e re c a rr ie d o u t in som e h ig h m o u n tain stre a m s an d r iv e r s of th e G r e a t C a u casu s. In v e stig atio n s c o -v e r e d th e le ft sid e tr ib u ta r ie s of th e R i-v e r A laz an i: K ish ch ay , K ur- m u k h c h ay , T ala c h a y , K a te k h ch ay , a n d B e lak a n c h a y in th e te r r ito r y of A z e r b a ija n a n d th e u p p e r c o u r se of th e R iv er T e re k a n d its g lac ia l tr ib u ta r ie s S u atisi, M n aisid o n , C h k h e ri flo w in g d o w n fro m th e so u th -e r n a n d -e a s t-e r n slo p -e s of th -e Mt. K a zb -e k in th -e t-e r r it o r y of G -eo rg ia . T h ese in v e stig atio n s aim ed a t p r e se n tin g th e in v e r te b r a t e fa u n a co m -p osition , zo n al d istrib u tio n of fa u n istic asso c ia tio n s, an d a tte m -p te d at e lu c id a tin g th e o rig in of th is fa u n a in th e str e a m s of th e G r e at C a u -casus.

In th e in v e stig ate d stre a m s la r v a e of in sects, sp e c ia lly D iptera of th e fam ily C hiro n om id ae , a r e th e m a in c o m p o n en ts of th e in v e r te b ra ta fau na. T h e ir s h a r e u su a lly e x c e e d s 50 p e r c e n t of th e to tal fa u n a a n d e v e n in som e se c to rs of stre a m s th e y a r e th e o n ly r e p re se n ta tiv e s of in v e rte b ra te s . F ro m o t h e r g ro u p s th e l a r v a e of m ay flies, ston e-flies, an d D iptera of th e fam ily S im u liid a e (Fig. 1) a r e f a ir ly n u m e ro u s in som e strea m s. In th e se stre a m s a b o u t 100 tax o n s of in v e r te b ra te s ca n b e fo u n d w h a t is r e la t iv e ly little . In t h e b e s t re c o g n iz e d m o u n tain r iv e r s of E u ro p e i.e. th e R iv er F u ld a a n d th e R iv e r R aba, o v e r 500 sp e -cies w e re fo u n d ( M a r t e n 1984, L e h m a n n 1971, K o w n a c k i 1975). It sh o u ld be, h o w e v e r, re m e m b er e d th a t in th e C au c asu s on ly th e h ig h e st se cto r s of stre a m s w e r e in v e s tig a te d , w h e re as, in th e ab o v e m e n tio n e d r iv e r s th eir w h o le co u rse w a s d e te r m in e d .

T h e fa u n a of p a r tic u la r ca tc h m e n t a r e a s a n d e v e n p a rtic u la r s t a -tio n s w ith in th e stre am is v e r y d iffe re n tiate d . F o u r ty p e s of zo ocen o- ses w e re d istin g u ish ed . T h e m o st fr e q u e n t ty p e e n c o u n te re d in th e in v e stig a te d stre a m s is th e h ig h m o u n tain zo o c en o se in w h ich C h iro

n o m id a e, w ith la rv a e of th e g e n u s D ia m esa , c o n s titu te th e m a in e l e

-m en t. S o -m eti-m es la r v a e of S i-m u liid a e o c c u r alo n g w ith th e -m in fa ir ly la r g e ' nu m b ers, w h e re a s, p a r tic ip a tio n of m a y fly a n d sto n e-f ly la r v a e is re la tiv e ly sm all. In th e se co n d ty p e of z o o c en o se s la r v a e of

Chiro-1 Л5 O T H E R T A X A Щ E P H E M E R O P T E R A (T) T A L A C H A Y

m

I--- K t S H C H A Y --- I l --- K U R M U K H C H A Y --- 1 г © - | i--- K A T E K H C H A Y ---1 V « i 2 Э 4 5 6 7 8 9 1 0 tl 1 2 1 3 H 1 5

F ig. 1. T o ta l n u m b er o f m a c r o i n v e r t e b r a te s an d p e r c e n t a g e d is t r ib u tio n o f d iffe r e n t t a x a at s a m p lin g s i t e s in th e C a u c a s u s 'str eam s : K is h c h a y , K u r m u k h c h a y , T a la c h a y ,

K a to k c h a y , B e la k a n c h a y (A z e r b a ija n S SR)

n o m id a e — O rlh o cla d iu s (E u o rth o cla d iu s) rivico la a n d sto n eflie s d o

-m inate. M ay flies of th e g e n u s B aetis a n d E peo ru s a r e a n i-m p o rtan t e le m e n t in this zo o cen o se. In th e th ir d g ro u p of z o o c en o se s C hirono-

m ida e — E u k ie ilerie lla c y a n e a a r e th e d o m in a tin g form s. T h e le a st

com m on is th e a sso c ia tio n w h e r e m ay flies, ca dd is-flies, sto ne-flies, a n d T ricla did ae d o m inate. T h is ty p e is e n c o u n te r e d in sm all fo r est tr ib u ta r ie s of m ain strea m s.

O b se r v a tio n s of th e fa u n a d istrib u tio n alo n g th e co u rse of stre am p e r m it to d istin g u ish so m e asso ciatio n s d iffering b o th in n u m b e r of in d iv id u a ls an d sp ec ia l c o m p o sitio n as w ell a s in d o m in atin g c h a ra c te -ristic sp ecie s. In th e u p p e r c o u r se of th e R iv e r T e re k a n d its h ig h m o u n ta in tr ib u ta r ie s th e follow in g d istrib u tio n of th e fa u n a w a s fo un d (Fig. 2). In th e stre am , ju s t b e lo w th e g la cie r, n o r e p r e s e n ta tiv e s of th e f au n a w e r e re p o rte d , th o u g h n u m ero u s C h iron o m id a e of th e g e n u s

D iam esa a n d s to n e flies P ro to n em u ra a ltico la w e r e se e n fly in g ab o v e

th e stream . But a lr e a d y 50 m b e lo w th e g la c ie r in d iv id u al, v e r y y o u n g la r v a e of th e g e n u s D ia m esa w e r e e n c o u n te r e d . A t a d istan ce of 500 m from th e g lac ie r a c h a ra c te r istic a sso c iatio n , co n sistin g m ain ly of

Dia-SUATISI and TEREK MNAI5IDON CZHERI DIAMESA S P (JU V ) 12 29 12+ - - - -DIAMESA GR LATITARSIS 22 16« + + - -ORTHOCLADIUS RIVICOLA О - 23<b + + -EUKIEFFERIELLA OA NEA - 20 DIAMESA GR INSIGNIPES 23 - 15+ - - -DIAMESA GR CINERELLA PROTONEMOURA s p 15 18 - + + - -BAETIS OR RHODANI + - - -Ml ... * - + -tor-2D + -+ -+ - + + +

-I

atH

P ig . 2. D is tr ib u tio n of b o tto m fa u n a c o m m u n itie s in t h e C a u c a s u s s tr e a m s : T erelc, S u a tis i, M n a is id o n a n d C h k h e ri ( = C z h e r i) (G e o r g ia n SS R ). T h e c o m m u n i tie s a re c o m p o s e d a c c o r d in g to th e i n d e x of d o m in a n c e ; n u m b e r in c i r c l e = fir s t d o m in a n t,

+ = su b d o m inan L , — = a d o m in a n t, s . = s p r i n g s tr e a m s

m esa lar v a e , w as found. T h ese w e re h ar d to id e n tify b u t on th e b asis

of sp ec im en s c o lle cte d a t le a st t h re e sp e cie s w e r e fo u n d to o ccu r: Dia-

m esa ca ucasica, D. tsk h o m e lid ze i, D. sa k a r tv e lla . T h is a sso c ia tio n d e

-v e lo p e d in th e stre a m s M n a is id o n a n d C h k h e ri alo n g th e ir w h o le co u rse . In th e str ea m S u a tisi fo rm e d b y m erg in g of so m e g la c ia l a n d sp rin g stre a m s th e ab o v e m en tio n e d a sso c ia tio n d e v e lo p e d o n ly in g la c ia l stre am s. In th e sp rin g stre a m th e first d o m in a n t w e r e la rv a e of O rth o cla d iu s rivico la. T h is sp ec ies d o m in a ted also alo n g th e w h o le len g th of th e stre a m S u atisi fro m th e p la ce w h e r e all th e stre a m s join. T his p r o v e s a m ark e d in f lu e n ce of sp rin g str ea m s a n d th e n u m er o u s sp rin g s situ a te d on th e riv e r b a n k on th e c h a ra c te r of b io cen o ses. W h e -reas, in th e in v e stig a te d se c to r of th e R iv e r T e re k (at th e a ltitu d e of 1700 to 2200 m) m ain ly a n a ss o c ia tio n co n sistin g of la rv a e of E u kief-

fe iie lla c ya n e a d ev e lo p s. T h is sp ec ie s d o m in ates also in th e trib u ta rie s

3. Z O O G G O G R A PH 1C R E M A RK S O N T H E F A U N A O F H I G H M O U N T A I N S T R E A M S

Z o o g eo g r ap h ie in v e stig atio n s of fr esh w a te r fa u n a of th e C au c a su s w e re s t a r te d b y p ro fe sso r D e r zh a v in w ith a s er ie s of p a p e rs on d is trib u tio n a n d o rig in of fish a n d c ru sta ce a . T h e se in v es tig atio n s w e re s u trib -se q u e n tly co n tin u e d by K a s y m o v (1972).

Z o o g eo g ra p hie c o n sid e ra tio n s p r e s e n te d in th is p a p e r w e re c a r -ried o u t e x c lu siv e ly o n th e b a sis of th e g e n u s D iam esa . T his g e n u s w as ch o se n sin c e its la r v a e an d p u p a e a r e ty p ic al in h ab ita n ts of m o-u n ta in strea m s a n d th ese a r e th e in v e stig atio n o bject.

In th e G re at an d S m all C au c a su s 13 sp ec ies of th e g e n u s D iam esa w e r e foun d to o cc u r ( K o w n a c k i , K o w n a c k a 1973a, 1973b, 1974, 1980, S h i l o v a 1968). D iam esa lin d ro th i, r e g a rd e d by som e a u th o rs as a b o real form , h a d a lr e a d y b e e n sy g n alize d fro m o th e r m o u n ta in s of E u ro p e an d sh o u ld b e r e g a r d e d as a n b o re o -a lp in e elem en t. T h e fo l-lo w in g a r e b o re o - alp in e sp ecies of la r g e ra n g e : D iam es aberrata, D.

b ertra m i, an d D. th ien e m a n n i. A m o ng E u ro -C au ca sia n sp ec ie s D. m ar- ta e an d D. m od e sta h a v e a n A lp in e -C a u c a sia n ty p e of d is trib u tio n

D. la v ille i P y r en e sia n -C a u c a sia n ty p e of d istr ib u tio n , w h e re as , D. va il-

la n li an d D. th o m a si a m o u n ta in ty p e of la rg e ra n g e b o th in th e g e o

-g ra p h ic al an d v e r tic a l sense. S u b p o n to - M ed iterr a n e a n sp ec ies a re :

Dia-m esa sa k a rtv e lla a n d D. k a s y Dia-m o v i fo u n d in th e C au c a su s an d in th e

m o u n tain s of S m all A sia (the m o u n ta in of L eb an o n — M o u bay ed , La- v ille 1983) as w ell a s sp ec ies found , so far, in t h e C au c asu s i.e.

Dia-m esa cau casica a n d D. tsk h o Dia-m e lid ze i (Tab. 1).

It ca n b e se e n from th e a b o v e p r e s e n te d d a t a th a t in fa u n a of

Diam esa of th e C au c a su s th e p a r tic ip a tio n of b o re o lp in e sp ec ies in re la

-tiv e ly sm all w h e n c o m p a re d w ith o th e r m o u n tain s of E urope. Let u s ta k e for e x a m p le th e T a tr a M ts w h e r e for 14 sp ec ie s of th e g e n u s

D iam esa 7 sh o w a b o re o - m o u n ta in d is trib u tio n ( K o w n a c k i 1986).

In th e fa u n a of th e C a u ca su s, on th e o th e r h and , E u ro - C a u ca sia n and su b p o n to -M e d ite rr a n e a n sp ec ies p re v a il. It sho u ld, th u s, b e assu m ed th a t th e p e rio d of th e Q a r t e r n a r y g la ciatio n s, w h ic h m ar k e d ly in flu e n ce d th e fa u n a of D iam esa of th e T a tr a M ts did n o t p la y su ch an im -p o r ta n t ro le th ere . In s-p ite of it th e r e is e v id e n t c o n c u r re n c e b e tw e e n th e m o u n tain fa u n a of E u ro p e a n d th a t of th e C a u ca su s. F o r 13 spe cie s of D iam esa from th e C au c a su s o n ly 4 h a v e n o t y e t b ee n fo u nd in th e m o u n tain s of E urop e. H e n ce, th e r e w a s a p e r m a n e n t e x c h an g e of sp e -cies b e tw e e n th e C au c a su s an d th e m o u n tain s of E uro pa, w h e re a s co n n ec tio n s b e tw e e n th e C au c a su s an d th e o th e r m o u n tain s of A sia

T a b l e 1 Zoog<4>grtip h ic a l e le m e n ts in th e fa u n a of th e g e n u s D i a m e s a in C a u c a s u s Z o o g e o g r a p h ic a l e l e m e n ts S p e c i e s D, a b e r r at a D. b e r t r am i B o r e o -A lp in e D. t h i e n e m a n n i D. l i n d r o t h i D. m a r t a e D. m o d e s t a E u r o -C a u c a s ia n D . t h o m a s i D . l a v i U e i D. s a k a r l v e l l a S u b p o n to -M e d it e r r a n o a n D. k a s y m o v i C a u c a s ia n D. c a u c a s i c a D . ts k h o m e li d z e i

a r e c o n sid e ra b ly lo oser. A s a m a tte r of fac t D ia m esa of th e m o u n tain s of A sia a r e n o t w e ll reco g n ize d h ith er to , n e v e r th e le ss som e co n c lu

-sio n s c a n a lr e a d y b e d ra w n o n th e b a sis of th e e x is tin g d ata. F o r ex a m p le in th e w h o le H im alay as, 11 sp ecies of D iam esa w e re foun d b u t on ly 1 sp ecies, i.e. D iam esa a b eria ta , is a sp ecies co m m o n to th e H im alay as, th e C au ca su s, a n d th e m o u n tain s of E urop e.

P o lis h A c a d e m y o f S c ie n c e s L a b o r a tor y of W a te r B io lo g y K r a k ó w B I B L IO G R A PH Y B c h n i n g A. L., P o p o v a A . N „ M a t e r i a ł y p o g i d r o b i o l o g i i г. Z an g i ot i s t o k a d o g. E r y v a n a , T r u d y s e v a n . g id r o b io l. S tan c ., 7, 1947. K a k a u r i d z e T. G., R a s p r e d e l e n i e b e n t i ć e s k o i f a u n y v g o r n o i r e k e D a b a k h a n a , T r u d y In st. Z oo l., T b il is i, 12, 1946. K a s y m o v A . G., P r e s n o v o d n a j a l a u n a K a v k a z a , Izd. E. L. M ., 1972. K o w n a c k a M. , K o w n a c k i A ., V e r t i c a l d i s t r i b u t i o n o l z o o c e n o s e s i n t h e s t r e a m s o l t h e T a tr a , C a u c a s u s a n d B a l k a n s M t s „ V e r b . I n te r n at. V e r e in . L im - n o l., 18, 1972. K o w n a c k i A ., R i v e r R a b a a n d i t s b i o c e n o s e s ( S c i e n t i f i c g u i d e ) , II S ym p . n a tem a t E p h e m e ro p te ra , 1975. K o w n a c k i A ., E c o l o g y a n d b i o g e o g r a p h y of D i a m e s i n a e ( C h i r o n o m i d a e , D i p t e r a ) f r o m P o l a n d , A c t a U n iv . L od z., F o li a lim n o l., 2, 1986.

K o w n a c k i A. , K o w n a c k a M ., C h i r o n o m i d a e ( D i p t e r a ) f r o m t h e C a u c a s u s . D i a m e s a W a l l ! g r o u p s l e i n b o e c k i , B u ll. A c a d . p o l. S e i., C l. II, S er. S ei. b iol., 21, 1973 a.

K o w n a c k i A. , K o w n a c k a M ., C h i r o n o m i d a e ( D i p t e r a ) I r om t h e C a u c a s u s . II. D i a m e s a W a l t 1 g r o u p l a t i t a r s i s , B u ll. A c a d . p o l. S ei., C l. 11, S er. S e i. b iol., 21, 1973b. K o w n a c k i A. , K o w n a c k a M ., R e l a t i o n o l C h i r o n o m i d a e I r o m T a t r a a n d C a u -c a s u s M t s . , Ent. T id s k r ., S u p p l. 95, 1974. K o w n a c k i A. , K o w n a c k a M ., D i a m e s a m a r t a e s p . n. (D i p t e r a : C h i r o n o m i -d a e ) a n -d g e o g r a p h i c a l -d i s t r i b u t i o n о I t h e s p e c i e s o l D i a m e s a l a t i t a r s i s gr ou p , A c t a H y d r o b io l., 22, 1980a. K o w n a c k i A ., Z o s i d z и R., T a x o c e n s o l C h i r o n o m i d a e ( D i p t e r a ) in s o m e r i -v e r s a n d s t r e a m s o l t h e A d z h a r A S S R (L i t t l e C a u c a s u s M t s ) , A c t a H y d r o b io l ., 22, 1980b. L e h m a n n J., D i e C h i r o n o m i d e n d e r F u lda. S y s t e m a t i s c h e , ö k o l o g i s c h e u n d l au n i - s t i s c h e U n t e r s u c h u n g e n , A r c h . H y d r o b io l. , S u p p l., 37, 1971. M a r t e n M „ T h e m a c r o z o o b e n l h o s o l t h e r i v e r F u ld a: l a u n i s t i c s , l o n g i t u d i n a l z o o - n a t i o n a n d c h a n g e s in t h e l as t 30 y e a r s , V e r h . I n ter n at. V e r e in . L im n o l., 22, 1984. M u r v a n i d z e D. I„ M a t e r i a ł y к g y d r o b i o l o g i i r. B a k u r i a n k i , T r u d y In st. Z o ol A k a d . N a u k G ru z. S SR , 8, 1949. S h i l o v a A . I., N o v a n a k h o d k a D i a m e s a l a v i l l e i ( D i p t e r a , C h i r o n o m i d a e ) v l a u n e S o v e l s k o g o S o y u z a , Z o ol. Z h., 57, 1978. T a r n о g r a d s к у D. A ., P о p о v K. K., К b i o l o g i i i r a s p r o s t r a n e n i j u p e r e d a t c h i k a l a s c i o l e z a L i m n e a I r u n k u l a t a M ü l l , na S e v . K a v k a z e , P ab . S e v .-K a v k . g id r o b io l. st., 1, 4, 1932/1933. Z o s i d z e R. Sh „ B e n t o t a u n a r e k A d z h a r s k o i A S S R , A v t o r e f. D is s , Izd. T b ilis k , U n iv ., T b il is i 1973. A n d r z e j K o w n a c k i F A U N A B E Z K R Ę G O W C Ó W W W Y S O K O G Ó R S K IC H P O T O K A C H K A U K A Z U W p o to k a c h K a u k az u g ł ó w n y m s k ł a d n i k ie m fa u n y b e z k r ę g o w c ó w są la r w y o w a d ó w , a z w ł a s z c z a m u c h ó w k i z r o d z in y C h i r o n o m i d a e (ry s. 1). O b s e r w u ją c r o z -m i e s z c z e n ie f a u n y w z d łu ż b ie g u p o to k ó w w y r ó ż n i o n o k i lk a z e s p o ł ó w r ó ż n ią c y c h s i ę z a r ó w n o i lo ś c ią o s o b n ik ó w , s k ła d e m g a tu n k o w y m , ja k i c h a r a k te r y s t y c z n y m i g a tu n k a m i d o m in u j ą c y m i (rys. 2). W p o to k a c h l o d o w c o w y c h u w r ó t lo d o w c a n ie z n a le z i o n o p r z e d s ta w i c i e l i fa u n y . A le ju ż p o n iż e j lo d o w c a w o d l e g ło ś c i o k o ło 50 m s p o t y k a n o p o j e d y n c z e m ło d e l a r w y C h i r o n o m i d a e z ro d za ju D i a m e s a . W o d le g ł o ś c i 500 m o d io d o w c a r o z w ija s i ę c h a r a k t e r y s t y c z n y z e s p ó ł z ło ż o n y g ł ó w n i e z la rw k ilk u g a t u n k ó w D iam es a . N a to m ia s t w p o to k a c h b io r ą c y c h p o c z ą t e k z e ź r ó d e ł d o -m in o w a ły la r w y O r t h o c l a d i u s (E.) r i v i c o l a . Po p o łą c z e n iu s ię w s z y s t k i c h w ię k s z y c h p o t o k ó w p o w s t a je d u ż a w y s o k o g ó r s k a r z e k a , w k tó r e j r o z w ij a s i ę g ł ó w n ie z e s p ó ł złoólon y z la r w E u k i e i l e r i e l l a c y a n e a .

P o n ie w a ż g a tu n k i z r o d z a ju D i a m e s a s ą t y p o w y m i m i e s z k a ń c a m i w y s o k o g ó r -s k ic h p o to k ó w W i e l k i e g o K au k a zu , -s ta r a n o -s ię w y j a ś n ić -s p r a w ę ic h p o c h o d z e n i a S tw ie r d z o n o w y s t ę p o w a n i e 13 g a tu n k ó w z ro d z aju D i a m e s a (tab . 1). T y l k o 7 g a

-t u n k ó w m a r o z m i e s z c z e n ie b o r e o - a lp e js ld e . P o z o s t a ł o to g a t u n k i e u r o -k a tik a s k i e i s u b p o n to -m e d y to r r a n e ń s k ie or a z d w a u z n a n e c h w il o w o za e n d e m i ty k a u k a s k i e .

N a 13 g a t u n k ó w D i a m e s a z K a u k a zu t y lk o 4 n ie z o s t a ły d o t y c h c z a s s t w i e r d z o n e w g ó r a c h E u ro p y. I s tn ia ła z a t e m s t a ł a w y m ia n a g a t u n k ó w p o m ią d z y K a u k az e m a g ó r a m i e u r o p e j s k im i . N a t o m ia s t z n a c z n ie m n i e js z e s ą p o w ią z a n i a p o m i ę d z y K a u -k a ze m a p o z o s ta ły m i gó r am i A z ji. P r z y -k ł a d o w o w c a ły c h H im a la ja c h s t w ie r d z o n o 11 g a t u n k ó w D i a m e s a , a le t y lk o D i a m e s a a b e r r a t a j e s t w s p ó ln a d la H im a la jó w , K a u k a z u i g ó r E u rop y .