POLISH ACADEMY OF SCIENCES
MUSEUM AND INSTITUTE OF ZOOLOGY
■A G ^M
FAUNISTICA
The su c c e ss io n o f fauna
in th e B iałow ieża prim eval p in e fo rest
FOREST BIODIVERSITY PROTECTION PROGRAM
GEF 0 5 / 2 1 6 8 5 POL
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POLI SH ACADEMY OF S C I E N C E S
MUSEUM
AND
INSTITUTE
OF
ZOOLOGY
FOREST BIODIVERSITY PROTECTION PROGRAM
PROTECTION OF FOREST ECOSYSTEMS
G EF 0 5 /2 1 6 8 5 POL
Przem ysław T r o ja n
R egina B a ń k o w sk a , E lżbieta C h u d z ic k a , Irm ina P ilipiuk
Ew a S k ib iń sk a , M aria S t e r z y ń s k a , J o la n ta W ytw er
Secondary succession of fauna
in the pine forests of Puszcza Białowieska
E dited by
PRZEMYSŁAW TROJAN
W A R S A W
1 9 9 4
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REDAKCJA
E lż b ie ta C h u d zick a , H en ry k G arb arczyk (z a s tę p c a r e d a k to r a n a c ze ln e g o ), A n n a Lian
W aldem ar M ik o ła jcz y k , R e g in a P isa r sk a (re d a k to r n a cz e ln y ), M aria S te r z y ń s k a ,
Ewa S k ib iń sk a (s e k re ta rz re d a k cji)
A d res R e d a k c ji
M u zeu m i I n s ty tu t Zoologii P o lsk iej A k ad em ii N a u k
0 0 -6 7 9 W a rsz aw a, u l. W ilcza 64
© C opyright by M uzeum i In sty tu t Zoologii PAN
W arszaw a 1994
ISBN 8 3 -8 5 1 9 2 -3 7 -9
ISSN 0015-9301
N akład 6 5 0 egz. P ap ier kl. III 80 g.
S kład kom puterow y, łam an ie i diapozytyw y - Dział W ydaw nictw MilZ PAN
D ru k - D ru k a rn ia Nr 1, W arszaw a, R akow iecka 37
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Przem ysław T r o ja n
Regina B a ń k o w sk a , Elżbieta C h u d z ic k a , Irm ina P ilipiuk
Ew a S k ib iń sk a , M aria S ter zy ń sk a , J o la n ta W ytw er
S econ d ary s u c c e s s io n o f fauna in th e pine fo rests o f P u szcza B iałow ieska
A b stract. M ultidirectional h y p o th e sis of th e seco n d ary su cc e ssio n of fa u n a is considered. It c o n ta in s
five types of su ccessio n : creative, stab ilisin g , rise-and-fall, regressive a n d resto rativ e. T he verification
o f th e se h y p o th eses d u rin g p ine forest developm ent is b ase d on ch a n g e s in th e n u m b e r of species,
th e ir a b u n d a n c e , th e s tru c tu re of th e taxocoenes, a c tu a l a n d p o ten tial species diversity. 27 tax o n s
(fam ilies or orders) w ith 1247 species o ccu rrin g in Białowieża prim eval p ine forest served a s a m aterial
for co m parisons. T hey re p re se n t all m ain tro p h ic ty p es of th e fau n a: p h y to p h ag es, p red ato rs,
p a ra s ite s an d sap ro p h a g e s am ong in v erteb rates, a n d also b ird s a n d m am m als.
A fter th e clear c u ttin g of trees, no em pty place a p p e a rs w ith reg ard to fa u n a , w hich is th e ric h e st a t
th is stage. The m u ltid irectio n a l c h a ra c te r of th e su cc essio n of fa u n a is p red o m in an t. M any indices
d ec re a se th eir v alu es in th e m a tu re forest. E cosystem stab ility is e n su re d by th e b e tte r ad a p te d
form s w hile th e p o ten tial diversity is n o t fully realised.
1. INTRODUCTION
1.1. S tu d y o b je ctiv e s
The form ation of forest ecosystem s ta k e s a long tim e. D epending on th e initial
s ta te of the environm ent, th e p ro cess m ay follow one of two possible p a tte rn s.
T he first p a tte rn is observed w hen a forest develops on a form erly u nforested
site, i.e. w hen a forest is form ed in th e a re a for th e first tim e. S u c h a pro cess is
called prim ary su ccessio n . The o th er p a tte rn is activated w hen th e forest
develops in a re a s w hich were previously occupied by a forest ecosystem , so th a t
one m ay sp e ak of forest rec o n stru c tio n in th a t case. It is u su a lly sta rte d by
p lan tin g tre e s in a clearing form ed after forest felling or a forest fire. T his process
is term ed seco n d ary succession.
An exam ple of prim ary su ccessio n is th e form ation of forest ecosystem s in
d u n e a re a s a s a re s u lt of th e drain ag e of Lake Chicago a n d the form ation of th e
shoreline of Lake M ichigan in the postglacial period. T his object re p re se n ts a
classic exam ple of forest developm ent from th e stage of a sa n d y b each to a
m aple-beech forest. The sere also in cludes a pine forest stage (C o w l e s 1899,
S u e l f o r d 1913). It h a s helped to c o n s tru c t a theory of ecological succession:
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th e f o rm in g o f t h i s e c o s y s te m to o k a b o u t 1 0 ,0 0 0 y e a rs ; in th e fin a l s ta g e , th e
v e g e ta tio n c o v e r a t t a i n e d c o n s id e r a b le s ta b ility , w h ile t h e r e a r e c o n s t a n t
c h a n g e s in t h e m in e r a l c y c lin g in t h e e c o s y s te m s (O lson 1958).
E u ro p e a n coniferous forests follow th e o th er p a tte rn - secondary succession.
C ultivated, felled an d afforested anew for h u n d re d s of y ears, they becom e
“ecologically m a tu re ” a s soon a s they reach th e felling age. For th e pine forest,
th e felling age is 8 0 -1 5 0 years. T his tim e sp a n d eterm in es th e total length of th e
process of secondary su ccessio n b e c au se forest felling re n d e rs im possible
fu rth e r tra n s fo rm a tio n s of b oth th e h a b ita t a n d th e biocenosis, a n d th e
ecosystem r e tu rn s to the initial stage w ith new c u ltu re s being planted.
E u ro p e a n a n d A m erican ecologists u su a lly consider th e p ro cesses of forest
su ccessio n from th e view point of th e clim ax theory. The theory s ta te s th a t th e
settling of th e form er forest h a b ita t is th e d o m in an t ph en o m en o n in the first
stage of su ccessio n . In th e second stage, specific su b sy ste m s a re form ed w ithin
th e biocenosis. In the final stage of su ccessio n , after th e forest ecosystem h a s
developed, th e ecosystem is a d a p te d to the h a b ita t conditions in the area, su ch
a s clim atic factors. The resu ltin g final an d stab le stage of su ccessio n is called
th e clim atic clim ax. In som e cases, however, th e final stage of su ccessio n
dep en d s on th e fertility of th e soil h a b ita t, a n d is term ed th e edaphic climax.
The pro cess of su ccessio n is accom panied by ch an g es in biocenoses a n d
h a b ita t conditions. The direction of h a b ita t tra n sfo rm atio n s is not related to th e
h a b ita t co n d itio n s occurring a t th e b eginning of the process of su ccessio n . Most
seres develop in a convergent m anner, resu ltin g in the sam e final stage. U nder
th e c irc u m sta n c e s of th e Polish Low lands, the final stage is rep resen ted by th e
Pino-Quercetum mixed forest (Ko r n a ś 1950, Pawłowski 1959). T h a n k s to th a t,
p rim ary su ccessio n , no m a tte r w h e th e r it s ta rts in a shallow ed pond, m arsh ,
s a n d d u n e or lim e s t o n e r o c k s , l e a d s , a f t e r n u m e r o u s e c o s y s te m
tra n sfo rm atio n s, to th e form ation of a m ixed forest.
The tra n sfo rm atio n of th e vegetation cover in th e co u rse of forest su ccessio n
is relatively well know n (F aliński 1991). M uch less is know n a b o u t ch an g es in
th e forest fau n a d u rin g forest s ta n d developm ent. The d a ta are incom plete, they
concern different forests, d issim ilar environm ental conditions, various tim e
sp a n s a n d , usually, a sm all n u m b e r of anim al species. The situ a tio n is th e sam e
a s reg a rd s stu d ie s of su ccessio n in E u ro p ean forests a n d forests in o th er
c o n tin e n ts.
O u r u n d e rs ta n d in g of fau n al tra n sfo rm a tio n s in th e co u rse of su ccessio n is
still m u ch b e tte r th a n th e theoretical b ack g ro u n d of su ccessio n p ro cesses
(An d e r s o n 1986). Theoretical a s su m p tio n s are usu ally m ade on th e b a sis of
diverse prem ises, w hich are u su a lly not sufficiently validated. An additional
difficulty for a zoologist is the fact th a t m ost concepts of su ccessio n are
co ncern ed w ith p la n t asso ciatio n s, so th a t th e findings, owing to specific tra its
of the b o tanical m aterial, c a n n o t be fully applied to faunal stu d ies.
B ecause of th a t the theoretical m ainstay for a zoologist is still the concept
form ulated by O dum (1969). In its term s, the process of ecological succession is:
1. u n id irectio n al a n d predicable,
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2. its p a tte rn an d rate are d eterm in ed by the physical en v iro n m en t w hich
also se ts lim its to developm ent,
3. th e process te rm in a te s in a stabilised ecosystem , w here m axim um b io m ass
is p rese n t a n d functional links betw een o rganism s e n s u re control over the
processes tak in g place a n d protection ag a in st d istu rb an c e s.
The m ec h a n ism s of ho m o eo stasis a re su p p o rte d by g ro u p s of o rg an ism s with
sim ilar functional c h a ra c te ristic s b u t different ran g es of ecological tolerance
(T r o ja n 1984). D epending on the in te rn a l a rra n g e m e n t of th e s e sy stem s
(Kikkawa 1986) they m ay be a d a p te d to m ain ta in in g sh o rt-te rm or long-term
stability.
Puszcza B iałow ieska is th e g rea test fau n al refugium in the C entral E u ro p e an
Lowland a s can be evidenced by com paring th e an im als o ccu rrin g th e re to th e
fau n a of th e C entral Polish Lowland. In the b e st know n g ro u p s of a n im a ls in
Puszcza Białow ieska, e.g. b u tterflies or Isopoda, th e n u m b e r of species recorded
in the Puszcza well exceeds 80% of th o se know n from the C en tral Polish Lowland
(T r o ja n u n p u b lish e d data). The fau n al ric h n e ss of Puszcza B iałow ieska is the
reaso n for the c o n c en tra tio n of fau n al rese a rc h in th e area. As a re su lt, it is
possible to analyze th e p h e n o m e n a occurring in the p ro cess of seco n d ary
su ccessio n of fauna. The n u m b e r of over 30 tax o n s (orders or families) stu d ied
is a rep resen tativ e sam ple allowing to tra c e reg u larities in th e p ro cess of
form ation of fau n al com plexes w ithin the pine forest biocenoses of Puszcza
Białowieska. The m aterial included in th e p re se n t p a p e r is not e x h au stiv e if one
co n sid ers the species ric h n e ss of Puszcza Białow ieska, b u t it is th e larg est fau n al
sam ple th a t h a s ever been investigated a t th e sam e place a n d stage of th e
developm ent of fa u n a of pine forests.
The d a ta on th e fa u n a of Puszcza Białow ieska a re an irreplaceable so u rc e of
com parative inform ation. The prim eval n a tu re of the forests is m u ch b e tte r
preserved, a n d th e n u m b e r of felling cycles m u ch sm aller, th a n in all oth er
E u ro p ean forests. As a resu lt, the d a ta on the fau n a of th a t forest com plex serve
as a fram e of reference for all fau n al stu d ie s con d u cted in Polish forests a s well
as in o th er co u n tries. The s tru c tu re of th e fa u n a of P uszcza B iałow ieska m ay be
used in a n exem plary fashion w hen analysing c h a n g es o ccu rrin g in fau n al
com m unities in forests s itu a te d in o th e r areas.
O bjectives o f th e stu dy. The following objectives w ere set for th e a n a ly sis of
the secondary su ccessio n of pine forest fauna:
1. D eterm ination of th e form ula for pine forest zoocenosis s tru c tu re , and
classification of specific g ro u p s of a n im a ls into functional g ro u p s w ith in the
forest biocenosis.
2. C om parison of th e species com position of th e anim al g ro u p s stu d ie d in the
various su ccessio n stag es a n d a sse s s m e n t of c h an g es in th e species com position
of the fauna.
3. E stim ation of th e n u m b e r of species in each anim al group. C om parison of
em pirical an d estim ated d a ta a n d a s se s s m e n t of th e degree to w hich th e m aterial
collected is representative.
4. D escription of the s tru c tu re of th e tax o n s in each stage of th e sere.
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5. D eterm in atio n of a c tu a l a n d potential species diversity of th e tax o n s in th e
co u rse of su ccessio n .
6. C om parison of c h an g es in th e s tru c tu re a n d diversity of each fau n al
com plex in th e c o u rse of secondary su ccessio n of the pine forests in Puszcza
Białow ieska.
7. A sse ssm e n t of th e degree of com plexity of th e zoocenosis in each stage of
su ccessio n .
T h e e n t ir e a n a l y s i s o f th e p h e n o m e n a o c c u r r in g in th e f a u n a d u r in g p in e
f o r e s t d e v e lo p m e n t is b a s e d o n t h e c o n c e p t o f f u n c tio n a l g r o u p s w ith in th e
b io c e n o s is . T h e g r o u p s a r e lin k s w h ic h fo rm t h e s u b s y s t e m s o f th e b io c e n o s is
to m a k e u p t h e h o m o e o s ta tic s y s te m (Tr o ja n 1 9 8 4 ), w ith in w h ic h th e p r o c e s s e s
o f a b u n d a n c e r e g u la tio n in fo r e s t e c o s y s te m s o c c u r.
1 .2 . R eview o f litera tu re
The c o u rse of secondary su ccessio n a n d the u n d erlying m ec h a n ism s have
b een a ttra c tin g n a tu r a lis ts ’ a tte n tio n since the tu r n of o u r century. Over the la st
15 years, n early 100 p a p e rs co ncerned w ith th is su b je c t have been m entioned
in bibliographical registers. A considerable n u m b e r of p a p e rs is concerned w ith
th e theo ry of su ccessio n (about 30 item s). Reviews of p a p e rs dealing w ith the
th eo ry of su c ce ssio n can be found in A n d e r s o n (1986) a n d F alińska (1991). The
developm ent of th e theory of su ccessio n coincides w ith ongoing reco n sid eratio n s
of th e concept of th e ecosystem , u n d e rta k e n both by h o lists a n d by red u c tio n ists.
Holistic views have been reflected in p a p e rs by Ma r g a l e f (1968), O dum (1971)
a n d W h itta k er (1975) w here th e su c ce ssio n is believed to o ccu r on ecosystem
level. T heir re d u c tio n ist o p p o n en ts, e.g. M a c m a h o n (1980), G ray et al. (1987)
a tta c h g re a te r im p o rta n c e to p o p u la tio n p ro c e s s e s. A ccording to th em ,
su c ce ssio n re s u lts from the living strateg y of species a n d th eir in te rac tio n s w ith
th e e n v iro n m en t an d w ith one another.
Views on th e theory of su ccessio n , related notions, c h a ra c te ristic fea tu re s
a n d m e c h a n ism s have been m entioned in m any ecology textbooks (Tr o ja n 1975,
S t u g r e n 1976, O dum 1977, C o l l ie r , C ox et al. 1978). A n u m b e r of p a p e rs are
con cern ed w ith vario u s a sp e c ts of th is p h en o m en o n , c o n c en tra tin g on issu e s of
stability, diversity, bioenergetics a s well a s m ec h a n ism s a n d m ath em atical
m odels of su ccessio n .
A rec e n t tre n d h a s been to tre a t b o th co n cep ts a s com plem entary (Tr o ja n
1984) so th a t the p roperties stu d ie d w hen in te rp re tin g su ccessio n p h e n o m en a
r e la te b o th to in d iv id u a ls of v a rio u s s p e c ie s a n d to p o p u la tio n s a n d
co m m u n ities. S h u g a r t (1984) a n d P r e n t ic e (1986) em phasize the hierarch ical
c h a ra c te r of th e theory of su ccessio n , according to w hich p ro cesses occurring
on higher levels stem from p ro cesses tak in g place on lower levels of org an isatio n
of th e ecological system .
The stu d ie s of su ccessio n in forest ecosystem s a re c o n d u cted on different
levels of o rg an isatio n of th e ecological system , in clu d in g b o th soil an d vegetation
a n d zoocenoses. The lite ra tu re on su ccessio n in th e se ecosystem s p e rm its to
m ake co m p ariso n s in v ario u s types of forest s ta n d a n d to trace tra n sfo rm a tio n s
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of b o th vegetation fau n a. The stu d ie s are b ased on th e theory of edaphic clim ax.
D ata on seco n d ary su ccessio n in th e pine forests in Puszcza Białow ieska c o n ta in
th e re s u lts of a n a ly se s of a n u m b e r of seres. If th e sere develops in a n su ita b le
h a b ita t, th e ra te of su ccessio n is u su a lly accelerated. C om parative a n a ly se s
sh o w th a t w h e n a fo rest is grow n in a n u n s u ita b le h a b ita t, s u b s titu te
a sso c ia tio n s em erge an d n a tu ra l su c ce ssio n is delayed.
T he s tr u c tu r e of faunal sy ste m s is a sensitive in d icato r of ch an g es occu rrin g
in th e e n v iro n m e n t. They a re m a n ife ste d a s a lte ra tio n s in th e s p e c ie s
com position, s tru c tu re an d a b u n d a n c e of fau n al com m unities. A com parative
a n a ly sis of fau n al co m m u n ities carried o u t in s ta n d s of v ario u s age m ay be u se d
for describing a n d in te rp re tin g th e p ro cesses occurring in th e su ccessio n in th e
ecosystem stu d ie d . It is u su a lly c o n d u cted in sp atial seres, b u t its re s u lts can
be extrap o lated tem porally, w hich m akes possible in te rp re ta tio n of c h a n g es
o ccu rrin g in th e co u rse of su ccessio n , while th e tim e of field observation is
co n sid erab ly sh o rten ed . A re g u la r cycle of secondary su ccessio n in th e pine
fo rests of P uszcza B iałow ieska is com pleted in 150 years.
T he lite ra tu re on th e p ro ce sse s of secondary su ccessio n in the pine fo rests of
Puszcza B iałow ieska is ra th e r poor. The bo tan ical a sp e c ts have been dealt w ith
by F aliński (1989, 1991) a n d T olkać & D a n cev ic (1974). D ata on fau n a l
su c ce ssio n in th e se ecosystem s a re provided in 1 1 item s. W ith one exception,
they are con cern ed w ith selected g ro u p s of in sects. L iterature on fa u n a is
exhaustively listed in Ka rpiń sk i a n d O k o łó w (1969) an d O kolów (1976, 1983,
1991).
2. AREA OF STUDY
P uszcza Białow ieska is s itu a te d in n o rth -e a s te rn Poland. The a re a is divided
by the sta te b o rd er betw een Poland an d B yelorussia. In its contem porary lim its,
Puszcza Białow ieska in cludes th e re m a in s of th re e forest com plexes asso ciated
in th e n o rth w ith the forest of P uszcza K nyszyńska. S itu a ted on the b o rd e r
betw een Poland, L ithuania, a n d th e historical regions of Jaćw ierz a n d R uś
W ołyńska, it w as dep o p u lated in the 13th cen tu ry (W ię c k o 1972) to rem a in
u n in h a b ite d for a b o u t 300 years. D espite th e pillage of th eir reso u rc es d u rin g
World W ars I a n d II, th e forests of Puszcza B iałow ieska have com pleted fewer
felling ro ta tio n s th a n forests s itu a te d w est of it. P articularly valuable a n d rich
a re th e d e c id u o u s forests c h a ra c te ristic of the tem p erate clim ate zone in E u ro p e
a n d u n d erg o in g “com plete se rie s of seco n d ary a n d recreative su c c e ssio n ”
(F aliński 1989). T aking into co n sid e ratio n th e n a tu ra l v alu es of th e se forests, it
w as decided th a t they w ould be p rotected a s a B iosphere Reserve.
A part from th e com m on lin d e n -o ak -h o rn b e a m forests a n d ald er forests,
Puszcza B iałow ieska also h o u se s pine forests rep re sen te d by th e u ltim a te forest
a sso c ia tio n s of Pino-Quercetum a n d Peucedano-Pinetum (su b co n tin en tal pine
forest) (Faliński 1988). The latter, re p re se n tin g th e “fre sh ” pine forest a sso ciatio n
type, grow on podzolic soils. T he m ajority of th ese forests are situ a te d in th e
s o u th -e a s te rn p a rt of P uszcza B iałow ieska, in H ajnów ka forest in sp ecto rate.
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P. T ro jan et al.
T here are a b o u t 3739 h a of pine fo rests in Puszcza Białow ieska, ac co u n tin g for
less th a n 7% of th e total forest s ta n d area. The p a tc h e s of th e pine forest are
com posed of pine (about 76%) a n d sp ru c e (15%) with a sm all add itio n of birch
(7%) a n d scan ty c o n trib u tio n s from o th e r tree species su c h a s oak or asp en .
S tu d ie s of secondary su c ce ssio n of the fau n a of th e forests of Puszcza
Białow ieska w ere carried ou t in H ajnów ka forest inspectorate, m ore specifically
in S tarzy n a forest d istrict. T hree site s w ere chosen in divisions 538, 667 an d
668. T hree even-aged coniferous forest sta n d s, several h e c ta re s each, were
selected w ithin each of th e site s for th e study. These included m a tu re s ta n d s
(tim ber sta n d s: 100-110 years), pole wood, i.e. Ill age c la ss s ta n d s (40-60 years),
young s ta n d s (15-25 years) an d c u ltu re s (3-7 years).
In order to collect cam bio- a n d xylophagous beetles, add itio n al c a tc h e s w ere
carried ou t in divisions 494 C, 519 D a n d 539 C.
T he location of stu d y sites is show n on th e m ap (Fig. 1).
B IA Ł O W IE S K I
PA R K
NARODOWY
Lutounia
I-----
138 53!
634!
Fig. 1. Location of stu d y s ta n d s in th e p in e fo rests of P uszcza Białowieska
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S econdary su c c e ssio n of fa u n a in th e p in e fo re sts of P uszcza B iałow ieska
9
Prior to collecting fau n al m aterials, s tu d ie s of p la n t asso ciatio n s (protocols)
a n d soil an a ly sis (soil profiles) w ere perform ed in th e a re a s chosen for th e stu d y
(M a t u s z k ie w ic z et al. 1993).
As re g a rd s soil a n d phytosociology, all th e a re a s included in the field stu d y
a re hom ogeneous an d re p re se n t th e su b b o re a l variety of the su b c o n tin e n ta l
(P eucedano-Pinetum ) pine forest. The variety is c h a ra c te rise d by a s u b s ta n tia l
co n trib u tio n of sp ru c e in th e s ta n d a n d u n d e rsto re y . The phytosociological
a n a ly sis c arried o u t in the s ta n d s revealed relatively insignificant differences in
floristic com position betw een various age c la sse s, even betw een stag es a s
different a s a m a tu re forest a n d a clearing w ith y o ung p lantings. Unlike
d e c id u o u s forests, th e a sso c ia tio n s form ing in th e c u ltu re an d young grow th
stag e do n o t differ phytosociologically. The a sso c ia tio n s, however, are poorer in
th e n u m b e r of species in com parison to a m a tu re pine forest, a t the sam e tim e
co m prising som e species from san d y greens.
In te rm s of soil typology, in all th e a re a s u n d e r stu d y , th e soils belong to th e
c la ss of podzols an d a re c h a ra c te rise d by acidic reactio n a n d low acid -b ase
sorption.
T he stu d y w as carried o u t in p a tc h e s of pine forest su b ject to no rm al
silv icu ltu ral practices. The a re a u n d e rg o e s clear felling w ith a cu ttin g w idth of
6 0 -8 0 m. The resto ra tio n of th e s ta n d is acco m p lish ed by p lan tin g a c u ltu re in
a specially p rep a re d felling site com posed of pine (80%), sp ru c e (10%) an d b irch
a n d oak (10%).
3. FAUNA SAMPLING METHODS AND MATERIAL
The program m e of stu d ie s en c o m p a ssed 30 g ro u p s of in v erteb rates a n d 2
g ro u p s of v e rteb rates. They re p re se n t diverse life styles an d a wide ran g e of
tro p h ic an d environm ental relatio n sh ip s. D ue to th a t, sam pling m eth o d s w ere
a d ju ste d to th e bionom ic p ecu liarities of the v ario u s g ro u p s of anim als. The
m eth o d s can be divided into two groups: qu alitativ e sam pling tech n iq u es and
s ta n d a rd q u a n tita tiv e m ethods.
T he q u a lita tiv e sam p lin g te c h n iq u e s w ere desig n ed specifically for th e
p a rtic u la r an im al g ro u p s stu d ie d . T hey a re d escrib ed in se p ara te p a p e rs
con cern ed w ith individual taxons. The m o st com m on qualitative m eth o d s
included “sta lk in g ” w ith a net, u sin g decoys to a ttra c t in se c ts an d brow sing
th ro u g h s u b s tra te s asso ciated w ith a p a rtic u la r fau n a l group.
The s ta n d a rd q u a n tita tiv e m eth o d s su ite d th e vertical s tru c tu re of th e forest
a n d activity p a tte rn s of th e a n im a ls u n d e r stu d y . W inged in sects o ccu rrin g in
th e canopy layer w ere collected u sin g M oericke's pitfall tra p s (M o e r ic k e 1950)
a tta c h e d to top b ra n c h e s of tre e s by m e a n s of special pegs. The pegs enabled
th e c o n ta in e rs to be h a u le d dow n a t re g u la r in terv als in o rder to tak e o u t th e
sp ecim en s collected. Since th e canopy layer s ta r ts to form only in grown up
p lan ta tio n s, th is sam pling tech n iq u e w as u se d in pine s ta n d s of th re e age
classes: y o u n g grow th, pole wood (age c la ss III) a n d m a tu re forest. T hree
sam pling site s were selected a n d 5 pitfall tra p s in stalled in each age class,
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10
P. T rojan et al.
resu ltin g in a total of 45 tra p s. T he sp ecim en s collected w ere ta k e n out every
fortnight, sta rtin g from th e w ithdraw al of th e snow cover u n til late November.
The m aterial collected u sin g th is m ethod in th e y e a rs 1986-1987 c o n tain ed
m ore th a n 5 00.000 of a ra c h n id a n d in se c t sp ecim en s (Tab. I). After the m aterial
h ad been sorted a n d species a n d th e ir a b u n d a n c e d eterm ined by experts in the
respective taxonom ic gro u p s, it w as su b jected to an aly ses described below.
T able I. List of m aterials collected u sin g M oericke’s pitfall tra p s in th e p ine forests of P uszcza
B iałow ieska in th e y ears 1986-87
Taxon
Psocoptera
B lattodea
Thpsanoptera
H omoptera
H eteroptera
Coleoptera
Neuroptera
Lepidoptera
H prnenoptera :
A culeata
Parasitica
S pm phijta
Diptera
N em atocera
Brachijcera
Aranei
Total
yung sta n d s
451
97
48582
2871
265
3629
1060
1308
N um bers of sp ecim en s
m a tu re s ta n d s
pole wood
695
566
32
10
83437
131262
5333
399 8
660
1243
2767
2281
794
548
889
890
Total
Average
1712
139
263281
12202
2168
867 7
2402
308 7
0.11
0.01
16.89
0 .7 8
0 .1 4
0 .5 6
0 .1 5
0 .2 0
3660
10618
615
1028
9384
581
1244
80963
653
5932
28965
1849
0 .3 8
1.86
0 .12
3002 6
63478
113
167773
29555
30581
68 9
166180
20726
20889
672
194190
80307
114948
247 4
5 2 8143
5.1 5
7.38
0 .1 6
3 3 .8 9
%
0 .3 2
0 .0 3
49 .8 4
2 .3 0
0.41
1.65
0 .4 4
0 .5 9
0 .0 0
1.12
5.49
0 .3 5
0 .0 0
15.20
2 1 .7 8
0 .4 7
1 0 0 .0 0 J
In o rder to d eterm in e th e species com position of th e h e rb layer fau n a , a
sw eeping n et w as u sed (10 sam p les of 25 sw eeps each in each area).
To collect an im als linked w ith tre e ste m s a n d b ra n c h e s, su c h a s Buprestidae.
C eram bycidae o r Scolytidae, s te m s a n d b ra n c h e s w ere s c a n n e d a n d foil
w indow -traps w ere used.
E stim atio n of th e species com position a n d a b u n d a n c e of epigeic a n im a ls w as
perform ed u sin g th e sq u a re plot m eth o d s (1 sq.m ., 10 sq.m . a n d 100 sq.m .).
This m ethod w as u se d m ostly in th e case of a n t com m unities. R odents w ere
c a u g h t into ja r s a n d tra p s placed on th e su rfa c e of soil.
Soil m eso fau n a w as a sse sse d in th e sp rin g a n d a u tu m n p eak s. A s ta n d a rd
steel corer w ith a n a re a of 20 sq. cm. w as u se d for o b tain in g sam ples. 10 sam ples
w ere ta k e n a t each site in each period. A dditional 25 sam p les w ere ta k e n a t th ree
sites in m a tu re forests in selected m ic ro h a b ita ts re p re se n te d by p a tc h e s of
bilberry, m oss, fern, litter a n d h e a th e r. The sam p les w ere th e n ex tracted in
T ullgren funnels. Special ex tractio n m eth o d s w ere em ployed for n em ato d es
(B r zesk i et al. 1976). The a n im a ls w ere th e n conserved in 4% form alin solution.
E n ch y traeid s w ere ex tracted in O’C o n n o r’s a p p a ra tu s (O’C o n n o r 19 7 1 ).
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S eco n d ary su c c e ssio n of fa u n a in th e pine fo rests of P uszcza B iałow ieska
4.
METHODS FOR PROCESSING DATA
T he d a ta collected, u su a lly quan titativ e, fulfilled th e criteria for application
in s ta tis tic a l a n a ly sis. T h is allow ed to s tu d y th e s tr u c tu r e of individual
tax o co en es a n d determ in e th eir q u a n tita tiv e ch aracteristics.
T he form ula for th e stru ctu re of v ario u s com m unities is obtained on th e
b a s is of m odels of five d istrib u tio n s, w hich a re m ost extensively u sed in faunal
s tu d ie s no w ad ay s ( T r o ja n 1992). The following m odels of s tru c tu re w ere used:
1. A geom etric series w herein a n a s su m p tio n is m ade th a t environm ental
re so u rc e s a re u sed successively in a n 1th p a rt by each species form ing th e
co m m unity, in a cco rd an ce w ith th e following series:
U ( l - I ) , (1 - I ) 2 .. • l ( 1 - 0 s-1
T h u s, th e a n ticip ated a b u n d a n c e of individual species a ssu m e s values
acco rd in g to th e form ula:
rn = r\Nl (1 - I ) 1' 1
w here 1 a n d q - eq u atio n c o n s ta n ts calcu lated from sam ple p a ra m ete rs
N
- the total n u m b e r of individuals in th e sam ple
i
- the ra n k of species in o rd er of dim inishing a b u n d a n c e .
2. B roken stick d istrib u tio n w herein it is a ssu m e d th a t environm ental
re so u rc e s a re utilised in a ran d o m m a n n e r by individual species form ing th e
com m unity; th e an ticip ated sh a re of th e species in th e com m unity is determ ined
from th e form ula:
i J =1
e Ą
=1 y
1___
(a t
w h e re i
rij
N
S
s
^
j= i
s + 1 -j
J
- th e s p e c ie s ' r a n k (fro m th e le a s t to th e m o s t a b u n d a n t)
- th e n u m b e r o f in d iv id u a ls in t h e s a m p le b e lo n g in g to th e
s p e c ie s o f j
- t h e to ta l n u m b e r o f in d iv id u a ls in th e s a m p le
- th e n u m b e r o f s p e c ie s in t h e s a m p le
3.
A lo g a r ith m ic s e r ie s w h e r e in t h e d is t r ib u tio n o f th e n u m b e r o f s p e c ie s ,
a c c o r d in g to a b u n d a n c e in c r e a s in g d is c r e te ly , a s s u m e s th e fo rm o f a s e rie s :
oo>2 a n 3
ocur
aU- ~ 2 ' ~ 3 ~ ............T
w h e re
r
- is th e h i g h e s t a b u n d a n c e o f in d iv id u a ls w ith in th e s p e c ie s ,
a n d t h e s e r ie s p a r a m e t e r s a a n d d a r e c a lc u la te d th e o r e tic a lly
r e la tiv e to
t h e n u m b e r o f s p e c ie s S a n d s a m p le a b u n d a n c e N ;
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12
P. T rojan et al.
4.
A lo g n o rm a l d is t r i b u t i o n b a s e d o n th e a s s u m p tio n t h a t s p e c ie s
a b u n d a n c e s , c o r r e s p o n d in g to th e a m o u n t s o f e n v ir o n m e n ta l r e s o u r c e s u s e d ,
p r e s e n t a n o r m a l d i s t r ib u tio n , w ith t h e g r e a te s t n u m b e r o f m o d e r a te ly a b u n d a n t
s p e c ie s , a n d s m a ll n u m b e r s o f r a r e a n d m o s t a b u n d a n t s p e c ie s ; in a lo g a r ith m ic
s c a le o f i n c r e a s in g c la s s e s o f a b u n d a n c e , c a lle d “o c ta v e s ”, in w h ic h t h e n u m b e r
o f s p e c ie s (S r ) d e te r m in e d in a c c o r d a n c e w ith t h e lo g n o r m a l d is tr ib u tio n
e q u a tio n , is d e fin e d b y t h e f u n c tio n :
Sr -
w here
S0
£
R
So e (£R)2
- n u m b e r of species in a m odal octave
- equation p a ra m ete r o btained from em pirical d istrib u tio n
- ordinal n u m b e r of a b u n d a n c e c la ss
5.
A negative binom ial d istrib u tio n b a sed on th e a ssu m p tio n th a t th e
probability of catch in g a given n u m b e r of individuals belonging to a p a rtic u la r
species is very low, corresp o n d in g to P oisson’s d istrib u tio n w ith th e X p aram eter;
if a c e rtain variability of th is p a ra m ete r is a ssu m e d , th e a n ticip ated s h a re s of
species in th e sam ple re p re sen te d by a given n u m b e r of individuals P{X = r), th e
following form is a ssu m e d for th e negative binom ial d istribution:
P ( X = r ) = r ~Y f r ~ p r qr
rl T ( v )
w here d. p, q are d istrib u tio n p a ra m e te rs determ in ed from the sam ple;
T he c o m p u ta tio n of d istrib u tio n s w as carrie d o u t in a c co rd a n ce w ith
descrip tio n s contained in a p a p e r by T r o ja n (1992) u sin g a co m p u te r program m e
of th e M useum an d In stitu te of Zoology PAS. W hen th e re s u lts w ere sim ilar to
th e m odels u sed , d a ta on q u a n tita tiv e c h a ra c te ristic s d eterm in in g th e s tru c tu re
of th e m odel w ere utilised. The sim ilarity of em pirical to theoretical d istrib u tio n s
p
w as verified on the b a sis of th e x test.
A ctual sp e c ie s d iv e rsity in individual co m m u n ities w as m e a su re d w ith
S h a n n o n a n d W eaver's diversity index:
i- 1
- th e a b u n d a n c e of th e ith species
- a b u n d a n c e of a given com m unity com posed of a total n u m b e r
of S species.
P o ten tia l sp e c ie s d iv ersity of v ario u s com m unities is expressed a s the
b o rd er v alu es of S h a n n o n a n d W eaver’s index ( T r o ja n 1994). B ecause of th a t,
th e degree of realisatio n of th e potential, expressed a s Pielou’s index, a s su m e s
the form:
w here
m
N
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S eco n d ary su c c essio n of fa u n a in th e p ine forests of P uszcza B iałow ieska
log 2
w here
S*
13
s
- is th e potential n u m b e r of species in the h ab itat.
A ssessm en t of sim ila rity o f s p e c ie s c o m p o sitio n of individual com m unities
w as m ade according to th e “p ercen tag e” m odification of S o eren sen 's form ula
(S0 ERENSEN 1948):
So = 2 —
100%
a+ b
w here a an d b refer to th e n u m b e r of species in the com m unities com pared
an d w to th e n u m b e r of species com m on to b oth com m unities.
5. ANALYSIS OF SECONDARY SUCCESSION OF THE PINE FORESTS
OF PUSZCZA BIAŁOWIESKA
5 .1 . A m od el o f th e stru ctu re o f p in e fo rest z o o c e n o sis
and th e co u rse o f seco n d a ry su c c e s sio n .
Anim al co m m u n ities in h ab itin g forest ecosystem s form functional system s
ch a ra c te rise d by close linkage betw een th e ir com ponents. Puszcza Białow ieska
is the first n a tu ra l forest in Poland, for w hich a m odel w as c o n stru c te d to
d eterm ine th e ecological s tru c tu re of th e ecosystem (Karpiński 1949). According
to th e m odel “th e functional gro u p s of th e florotone a n d th e zootone an d forest
h a b ita ts of c ertain types correlating w ith one a n o th e r form a forest biocenosis
of a specific type". K arpiński d istin g u ish ed 5 functional g ro u p s in th e zootone,
each w ith a different role to fulfil in the life of th e forest.
Now adays th e org an isatio n of forest ecosystem s is viewed a s a hierarch ical
system com prising 4 types of s tru c tu ra l u n its (T ro ja n 1984).
1. The su b sy ste m , w hich is c h a ra c te rise d by a specific in p u t s u b s tra te th a t
supplies th is system , a n d c o n trib u te s to reg u latin g th e whole ecosystem . T here
are five s u b sy ste m s w ithin each ecosystem : th e biotope, the pro d u cers, a n d th re e
s u b s y ste m s form ing th e zoocenosis (or zootone, acco rd in g to K arpiński):
1) sa p ro tro p h s, 2) bio tro p h s, 3) m acro tro p h s.
2. The tro p h ic (food) c h ain is the basic org an isatio n al u n it of the su b sy stem .
Individual c h a in s differ w ith regard to th e type of food (input link) consum ed
an d the m a n n e r in w hich th e food is c a p tu red . (One h o st p la n t m ay give rise to
several tro p h ic c h a in s of biotrophes). E ach c h a in co n sists of links corresponding
to trophic levels (Lindem ann 1942).
3. The com petitive asso ciatio n (guild) e n c o m p a sses species form ing a single
link of th e trophic chain. The species in h ab it identical or sim ilar ecological
niches, u n d e rsto o d a s com m on food reso u rc es utilised by th e entire guild an d
a com m on g ro u p of p redatory a n d p a ra sitic species exploiting th e guild.
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14
P. T rojan et al.
4.
T he p o p u lation is a n in te rn al org an isatio n u n it of th e guild b a s e d on
com petitive relations.
The s tru c tu re of zoocenoses b ased on th e above four types of u n its is
rep ro d u cib le a n d universal. This m ea n s th a t sim ilar o rg anisational s tru c tu re s
a re found b o th in forest ecosystem s a n d in m eadow or field ones, or even in
u rb ic e n o se s. D ifferences betw een th em c o n c ern th e s h a re s of in d iv id u al
o rg an isa tio n al u n its in a p a rtic u la r ecosystem . The differences are recognised
a n d defined a n d th e degree of u n iq u e n e ss is d eterm in ed by com parative
fau n istical analyses.
T he o rg an isatio n of forest biocenoses exhibits a high degree of com plexity
(S z u je c k i 1980). It c a n be seen b oth in conditions im posed by forest s ta n d
developm ent, its sp atial stru c tu re , se aso n a l a n d d iu rn a l rh y th m s of a n im a ls a n d
re la tio n sh ip s betw een individual com p o n en ts of th e environm ent. Any a tte m p t
to sy ste m atise th e se rela tio n sh ip s is faced w ith th e need to m ake sim plifications
a n d to a s s u m e th a t c e rtain factors are c e n tral to reg u latin g th e sy stem
o rg an isatio n . In the p re se n t p a p e r th e cen tral role is perform ed by th e tro p h ic
factor. T he s tru c tu re p resen ted is th u s equivalent to the trophic o rg an isatio n of
th e zoocenosis. The p a ra tro p h ic factor is given m inor im portance. Owing to th is,
for exam ple, p a ra sitic in se c ts have been classified after phytophages, in w hich
th e ir larvae develop. The p a ra site s play a n additional role a s m ellitophages,
feeding on honey-dew a n d n e c ta r a n d pollen of flowers, w hich a re pollinated in
th is way. T his system of feedback relatio n s h a s not been reflected, however, in
th e above outline of th e org an isatio n of th e pine forest zoocenosis of P uszcza
B iałow ieska.
The pine forest zoocenosis, like o th er forest ecosystem s of th is clim atic zone,
is com posed of th ree su b sy stem s: biotrophic, sa p ro tro p h ic an d m acrotrophic.
The p re s e n t p a p e r gives m ost c o n sid eratio n to th e biotrophic su b sy ste m b e c a u se
of th e c h a ra c te r of th e m ate ria ls collected a n d an alysed a n d the ability of
dom estic ex p erts to analyze individual anim al groups.
The biotrophic su b sy ste m com prises two g ro u p s of tro p h ic ch ain s. The c h a in s
are in itiated by first-level c o n su m e rs (C 1) i.e. chew ing phy to p h ag es a n d su c k in g
ph y to p h ag es (Fig. 2). P articularly im p o rta n t is th e chew ing p h y tophage link,
w hich com prises lep id o p teran s (Lepidoptera) an d sawflies (Sym phyta) - th e m ost
d a n g e ro u s pine foliophages w hich periodically invade som e regions of Poland
c a u sin g severe dam age to pine sta n d s . Sim ilar fu n ctio n s are also fulfilled by
weevils (Curculionidae) a n d leaf beetles (C hrysom elidae). Only th e form er pose a
significant th re a t to forests a s p e sts of pine. The next group of p h y to p h ag es is
com posed of species su c k in g leaves a n d young sp ro u ts. It in clu d es a p h id s
re p re se n te d by th e specialised family of Lachnidae asso ciated w ith coniferous
trees. Parallel c h a in s a re form ed by h o m o p te ran s (H om optera. A uchenorrhyncha )
a n d h e te ro p te ra n b u g s (Heteroptera). This group m ay be defined a s p h y to p h ag es
u n d e rm in in g th e forest s ta n d by su c k in g s u b s ta n tia l a m o u n ts of a ssim ilate s o u t
of leaves a n d sp ro u ts. A p a rtic u la r role in forest ecosystem s is played by
xylophages a n d cam biophages. T his g roup o ccu rs characteristically in forest
ecosystem s. In th e p re se n t stu d y , it is rep re sen te d by th re e beetle fam ilies:
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Secondary su c ce ssio n of fa u n a in th e p ine fo re sts of P uszcza B iałow ieska
15
BIOTROPH SUBSYSTEM
Fiuu6iiti
C2 + C3
Lathnid©
Coccinellidae
Auch©norrrhyncha
Neuropteroid©
H©t©ropt©r
S’yFphidae
others n n
V
\
Tachinidae
Lepidopter
Spheoidae
Symphyts
Ichneumonidee
Pipunculidee
Curculionida©
Chrysomelidae
Aranei
Asilidae
cotylida©
antharida©
Cerambycid
Bupr©stida©
d©ad organic matt©r
Forrnicida©
Enchytraeidae
Collem bola
N am atoda
Therevida©
others
Asilida©
D ith e rs
Soncida©
Rhagionida©
D th a rs
SAPROTROPH SUBSYSTEM
MACROTROPH SUBSYSTEM
Fig. 2. Zoocenosis o rg an isatio n in th e p ine forests of P u szcza B iałow ieska
b u p r e s tid s (B uprestidae), c e ra m b y c id s (Cerambycidae) a n d b a r k b e e tle s
(Scolytidae).
Zoophages include p a ra sitic a n d pred ato ry form s specifically a sso ciated w ith
ce rtain g ro u p s of phytophages. In th is p a p e r special a tte n tio n is given to
d istin g u is h in g p a ra s ito id s th a t red u ce the a b u n d a n c e of foliophages, i.e.
tac h in id flies (Tachinidae) an d ich n eu m o n flies (Ichneum onidae). G roups of
sp e c ie s re d u c in g th e a b u n d a n c e of s u c k in g p h y to p h ag e s, i.e. p re d a to ry
a p h id o p h ag es an d p a ra s ite s of H om opterans, a re also well rep resen ted . All th ese
anim al g ro u p s are second-level c o n su m e rs (C2). Two s u b s e q u e n t lin k s are
form ed by p redatory form s (C3) h u n tin g in flight or on th e su rface of p la n ts and
soil robber-flies (A silidae) a n d digger w asp s (Sphecidae)) a s well a s sp id ers
(Aranei) h u n tin g in th e ir w ebs.
The sap ro tro p h ic su b sy ste m in th e pine forest is well-developed b oth as
reg ard s d etrito p h ag es, polyphages a n d zoophages. Not all of th e g ro u p s of soil
fau n a ca n be an alysed a t p re se n t w ith a c cu ra cy req u ired for fau n al stu d ie s. The
re la tio n sh ip s o ccu rrin g in anim al co m m u n ities of th is su b sy ste m have been
described on the b a s is of m odel anim al g ro u p s su c h a s nem ato d es (N em atoda).
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16
P. T rojan et al.
e n c h y trae id s (E nchytraeidae) a n d s p rin g ta ils (Collembola). P olyphages a re
rep re sen te d by a n ts (Formicidae), w hich are m ost effective in controlling forest
ecosystem s. Therevidae served a s a n exam ple of soil p red a to rs.
The m acrotrophic su b sy ste m , w hich controls both the biotrophic a n d the
sap ro tro p h ic su b sy ste m s, is re p re sen te d by b ird s a n d sm all m am m als.
Ecological su ccessio n is one of th e m ore im p o rta n t p ro cesses o ccurring in
ecosystem s. It is a reflection of dynam ic p ro cesses th a t tak e place in every
ecosystem . The stu d ie s of th e forests of Puszcza B iałow ieska have provided a
b a sis for th e developm ent of th e dynam ic concept of vegetation cover (Paczo sk i
1951). The idea h a s also been developed by Ka rpiński (1949), who d istin g u ish e d
fluctuation and su ccessio n p rocesses. He also drew a tte n tio n to th e relative
stability of clim ax system s b u stlin g w ith c o n tin u o u s p ro cesses of change, w hich
are, however, no t directional. T hese co ncepts have recently been developed by
F aliński (1989). also on th e b a sis of Puszcza Białowieska. Falihski d istin g u ish e d
th re e types of p ro cesses o ccu rrin g in ecosystem s: directional, flu ctu atin g a n d
cyclical. Of th e th ree types, only th e directional p ro cesses of su ccessio n an d
regression are im p o rta n t from th e view point of th e p rese n t an aly sis. The
se p ara tio n of the two p ro cesses is, however, difficult. The ta s k is all th e m ore
com plicated b e c au se regression p ro cesses, like su ccessio n processes, consist of
m ultiple stag es an d re su lt in the system ad a p tin g to th e altered environm ental
conditions. Particularly u n c le a r is th e problem of finding m e a su re s for die
progress resu ltin g from su ccessio n . F aliński (1989: 105) lists am ong su ch
m e a su re s “a conventional d ynam ics in d ex ”, th e sta te of th e biom ass, energetic
relatio n s betw een th e co m p o n en ts, etc. All of th ese m e a su re s are increm en tal
so th a t non-grow th p ro cesses a re not regarded a s elem ents of th e notion of
succession. In zoocenology, su c h a rea so n in g h a s a n im plicit a s su m p tio n of
grow th of th e biom ass, diversity, a b u n d a n c e a n d th e n u m b e r of species. S uch
a course of su ccessio n is, however, very poorly evidenced for.
In the p rese n t paper, th e a ssu m p tio n is m ade th a t su ccessio n (prim ary as
well a s secondary) leads to relative stability of the system . The stability is
expressed, am ong o th er things, by th e fact th a t th e grow th p ro cesses in the
system are slowed down or stopped, a n d the entire energy gain is utilised in
in te rn a l regulatory p rocesses, i.e.
B « const.
AB= R
w here B - b io m ass a n d R - resp iratio n . All regulatory p ro cesses a re th u s
accom plished a t th e expense of energy, w hich, in the ecosystem 's final acco u n t,
is u sed to co m p en sate for the co sts of m ain te n a n c e an d activity, i.e. th e
m etabolism of th e co m p o n en ts form ing th e biocenosis.
It is not easy to resolve the iss u e s in q u estio n on theoretical g ro u n d s. The
n a tu re of m aterial collected for th e p u rp o se of the stu d y still p erm its verification
of the views on secondary su c ce ssio n by determ in in g w hich of th e 5 v a ria n ts of
su ccessio n p resen ted below (Fig. 3) is actu ally observed in th e co u rse of
developm ent of the pine forest of Puszcza Białow ieska.
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Secon dary su c c e ssio n of fa u n a in th e p ine fo rests of P uszcza B iałow ieska
17
b) stabilising succession
measure of succession progress (M)
a) creative succession
su cc essio n series
d) regressive succession
measure of succession progress (M)
c) rise- and- fall succession
IV
V
su ccessio n series
succ essio n series
measure of succession progress (M)
e) restorative succession
I
II
III
succ essio n series
IV
V
Fig. 3. V a ria n ts o f th e c o u rs e o f se c o n d a ry
su ccessio n
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18
P. T rojan et al.
C reative su ccessio n (a) is observed w hen th e m e a su re of su c ce ssio n progress
of a co m m u n ity in the consecutive sta g e s of th e sere is higher th a n th e preceding
one
Mo < M i < M2 . . . < M n
T he system w here su c h a series of v alu es is registered develops infinitely.
S tabilising su ccessio n (b) is seen w hen th e m e a su re of su c ce ssio n progress
in c r e a s e s in th e initial perio d to s u b s e q u e n tly sta b ilise , irre sp e c tiv e of
developm ent or grow th p ro cesses observed in o th er co m ponents
Mo < M i < M2 = M3 . . . = M n
R ise-and-fall su ccessio n (c) is c h a ra c te rise d by a n initial in cre ase in the
m e a su re of succession pro g ress followed by a fall
Mo < Ml < M2 ... < Mmax ^ . . . Mn-2 ^ Mn-1 > Mn
S u c h a series of v alu es will be c h a ra c te ristic of th o se co m m u n ities w ithin
zoocenoses w hich c o n trib u te to th e developm ent of th e system in th e initial
sta g e s of th e sere. They are often pioneer species or pioneer g ro u p s th a t set the
stag e for g ro u p s b e tte r ad ap ted to living in a n organised biocenosis.
Regressive su ccessio n (d) is observed in situ a tio n s w hen a g ro u p reach es
m ax im u m values in th e first stage of su ccessio n w here the sy stem is least
organised. The in crease in the degree of org an isatio n of th e biocenosis re su lts
in a g rad u a l red u ctio n of th e function of th is g roup a n d its elim ination or
s u b s ta n tia l decline.
R estorative su ccessio n (e) is a n a ttrib u te of com m unities w hich survived fairly
well th e c a ta stro p h e connected w ith th e d e stru c tio n of th e ecosystem , b u t
w itn e ss a d ecrease in th e m e a su re of su c ce ssio n pro g ress in the first stages. The
value in cre ase s only after th e target level of ecosystem org an isatio n is a ttain ed .
A n u m b e r of sim ple m e a su re s of th e progress of su ccessio n m ay be u sed in
a s se s s in g th e progress of su c ce ssio n of a given taxon rep re se n tin g e ith e r a
c e rta in fau n al com plex or a ce rtain guild:
1. A b undance, w hen a sse sse d w ith com parable m ethods; it m ay be referred
to a re a o r volume: or be relative in ch a ra c te r, expressing anim al activity or the
action of a h u n tin g tool. A b u n d an ce is a m e a su re of th e ecological su c c e ss of
th e tax o n stu d ied - if it in creases, or a regression - if it d ecreases.
2. The n u m b e r of species in h a b itin g th e ecosystem in a given stage of
su c ce ssio n reflects com m unity developm ent or recession. Both em pirical (S) a n d
e stim a te d (S*) n u m b e r of species m ay be used.
3. A ctual (H') a n d potential (Hmax) species diversity ind icates th e ric h n e ss of
a given taxocoene a n d th e degree to w hich it fills th e h a b ita t.
4. T he index of d om inance d eterm in in g q u a n tita tiv e re la tio n sh ip s betw een
sp ecies form ing a guild. In te rp re ta tio n of th e value of th is index is not clear,
especially w hen a sere is concerned.
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S econdary su ccessio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
19
5 .2 S u c c e ssio n in th e su b sy stem o f b iotrop h s
5 .2 .1 . P h y to p h a g es
As reg a rd s the m ode of feeding a n d the quality of th e food they ta k e in,
p h y to p h a g o u s in sects m ay be divided into th ree biotic gro u p s. The first group
is com posed of ecotrophs or chew ing p h y tophages, w hich feed on the green p a rts
of p la n ts. The second group is th a t of en d o tro p h s w ith a stin g in g -su ck in g m o u th
a p p a ra tu s designed to absorb s u b s ta n c e s produced by p la n ts w ith o u t d a m ag in g
th e a s s im ila tin g a p p a ra tu s . T he th ird g ro u p c o m p rise s x y lo p h ag es a n d
cam b io p h ag es trophically associated w ith woody p lan ts. C hew ing p h y to p h ag es
in clu d e c aterp illars of b u tterflies (Lepidoptera), larvae of sawflies (H ym en o p tera .
S y m p h yta ) a s well a s n u m e ro u s fam ilies of beetles, su c h a s leaf b eetles
(C hrysom elidae) or weevils (Curculionidae). S ucking p h y to p h ag es em brace,
above all else, h y m en o p te ra n s su c h a s a p h id s (Aphidoidea) a n d leafh o p p ers
[Auchenorrhyncha) t o g e t h e r w ith p h y t o p h a g o u s h e t e r o p t e r a n b u g s
(Heteroptera).
A nother im p o rta n t p h y to p h ag o u s group in forest ecosystem s is form ed of
xylophages a n d cam biophages biologically associated w ith trees.
S ucking ph y to p h ag es
7 species of a p h id s of the family Lachnidae, 23 species of leafhoppers a n d 39
sp e c ie s of p h y to p h a g o u s H e te ro p te ra (K o l o d z ie ja k in p r e s s , C h u d zic k a
u n p u b lis h e d d a ta , C m o lu c h o w a , L ec h o w sk i 1993) were recorded in th e can o p ies
of pine forest s ta n d s in Puszcza B iałow ieska in all the forest s ta n d age c la sse s
s tu d ie d (young grow th, pole wood, m a tu re forest). In each of th e above g ro u p s,
a sim ilar n u m b e r of species w as registered in every age c la ss of the forest s ta n d ,
w ith th e g rea test n u m b e r of ap h id species recorded in the pole wood, a n d of
leafhoppers a n d h e te ro p te ran bugs, in m a tu re s ta n d s (Fig. 4).
The species com position of all th e co m m u n ities of su c k in g p h y to p h ag es
collected in canopies of pine w as sim ilar (Soerensen’s sim ilarity indices ra n k e d
from 62% to 99%). The hig h est sim ilarity of species com position w as recorded
for co m m u n ities of a p h id s: in each age c la ss of th e fo rest s ta n d s th e se
co m m u n ities w ere solely com posed of m onophages feeding on one or a n u m b e r
of pine species. Sim ilarity indices of species com position of a p h id c o m m u n ities
o ccu rrin g in canopies of p ines in th e s ta n d s stu d ie d in Puszcza B iałow ieska
ranged betw een 80% -99% .
It sh o u ld be added th a t th e species com position of each of th e chew ing
p h y toph age co m m u n ities stu d ie d in th e canopies of tre e s in th e pine fo rests of
P uszcza B iałow ieska w as considerably sim ilar to th e species com position of
co m m u n ities in h ab itin g canopies (So) of p ines in o th er Polish forest com plexes
(K o l o d z ie ja k in p ress, C h u d zick a u n p u b lish e d d a ta , C m o lu c h o w a , L e c h o w s k i
1993).
In th e pine forest sere c h an g es in a b u n d a n c e a n d d om inance s tru c tu re o ccu r
in all th e co m m u n ities of su ck in g ph y to p h ag es u n d e r stu d y . In a p h id a n d
leafhopper co m m u n ities the indices of relative a b u n d a n c e d ecrease w ith th e
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20
P. T rojan et al.
SUCKING PHYTOPHAGES
25
20-
o
o
E
IV
—<■— Lachnidae
Auchenorrhyncha
Heteroptera
Fig. 4. C h anges in th e n u m b e r of species of su ck in g p h y to p h ag es in p in e fo rests (3 age c la sse s of
th e forest stand)
SUCKING PHYTOPHAGES
0.25
0. 2 -
ax>
£
0.15-
o
0.05-
IV
II
—
Lachni dae
Auchenorrhyncha
Heteroptera
Fig. 5. C hanges in th e a b u n d a n c e of su ck in g p h y to p h ag es in p ine fo rests (3 age c la sse s of th e forest
stan d )
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S econdary su c c e ssio n of fa u n a in th e p ine fo re sts of P uszcza B iałow ieska
21
grow ing age of the s ta n d , w hile co m m u n ities of p h y to p h ag o u s h e te ro p te ran b u g s
exhibit th e reverse (Fig. 5).
C h a n g e s in th e s t r u c tu r e of d o m in a n c e of th e p h y to p h a g o u s in se c t
c o m m u n ities u n d e r stu d y co n sisted p red o m in an tly in shifts of positions, or
r a n k s , of th e d o m in a n t sp ecies. O n th e o th e r h a n d , differences in th e
c om position of th e d o m in a n t species g ro u p w ere n o t observed in any com m unity
stu d ie d (Fig. 6-8).
A significant tra n sfo rm a tio n of th e d o m in an ce s tru c tu re in com m unities of
a p h id s took place betw een th e y o ung grow th a n d pole wood stages. C om pared
to th e y o u n g grow th com m unity, th e pole wood co m m u n ity is c h a ra c te rise d by
a d im in ish ed ra n k of Schizolachnus pineti, a species w hich form s com pact
colonies on to p s of s p ro u ts a n d feeds p red o m in an tly on y oung needles. On th e
o th e r h a n d , th ere is a n in cre ase in th e p roportion of E ulachnus agilis, a species
w hich does n o t form colonies and feeds on 2 -3 -y ear-o ld needles of pine (Fig. 6).
In the leafhopper co m m u n ities stu d ie d in th e sere, th ere is a n in crease in th e
r a n k of two pine m onophages, nam ely W agneripteryx germ ari a n d G rypotes
puncticollis. On th e o th er h a n d th e re w as a red u c tio n in b oth a b u n d a n c e an d
pro p o rtio n of S p eu dotettix su b fu sc u lu s , a polytope settling in v arious forest
h a b ita ts (Fig. 7). In th e leafhopper co m m u n ities, a considerable tra n sfo rm atio n
of the do m in an ce s tru c tu re occu rred betw een th e pole wood an d m a tu re stages.
In th e case of h e te ro p te ra n b u g s, c h an g es in th e s tru c tu re of dom inance
o ccu rrin g in th e sere c o n sisted p red o m in an tly in an in crease of the proportion
of Plesiodem a pinetellum , a m ono phage of pine, in older sta n d s. The species
Lachnidae
Fig. 6. D om inance s tru c tu re of c o m m u n ities o f a p h id s (Aphidoidea, Lachnidae) in can o p ie s of pine
in forest s ta n d s of v ario u s ages in P u szcza B iałow ieska; 1 - E ula ch n u s agilis, 2 - Schizolachnus pineti,
3 - Cinara pinea, 4 - C. pilosa, 5 - C. pini
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P. T rojan et al.
A u ch e n o rrh yn ch a
II
III
IV
Fig. 7. D om inance s tru c tu re of co m m u n ities of c ica d as (H om optera, A uchenorrhyncha) in can o p ie s
of pine in forest s ta n d s of v a rio u s ages in P u szcza B iałow ieska; 1 - S p e u d o tettix su b fu sc u lu s, 2 W agneripteryx germari, 3 - O ncopsis Jlauicollis, 4 - H esiu m domino, 5 - G rypotes puncticollis, 6 A llygus m ixtus
H eteroptera
515585^
Fig. 8. D om inance s tru c tu re of c o m m u n ities of h e te ro p te ra n s (H eteroptera ) in can o p ie s of p ine in
forest s ta n d s of vario u s ages in P uszcza Białowieska: 1 - Phytocoris intricatus, 2 - Phoenicocoris
obscurellus, 3 - C am ptozyg u m aequale, 4 - Phytocoris intricatus, 5 - A lloetom us germ anicus, 6 Phoenicocoris m odestus, 7 - P lesiodem a pinetellum
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S eco n d ary su c c e ssio n o f fa u n a in th e p in e fo rests of P uszcza B iałow ieska
23
a tta in e d p e a k a b u n d a n c e a n d proportion in the m a tu re forests of Puszcza
B iałow ieska (Fig. 8). Its a b u n d a n c e in pole wood w as nearly four tim es lower,
a n d in y oung s ta n d s , a h u n d re d tim es lower, th a n in m a tu re sta n d s. Moreover,
in older s ta n d s a d e crease w as observed in th e proportion of two species n o t so
n arro w ly specialised a s m onophages, nam ely Phytocoris intricatus a n d P. p in t
b o th of w hich a re trophically asso ciated w ith vario u s coniferous tree species.
Lachnidae
Auchenorrhyncha
H m ax
Hrnax
Heteroptera
.5 •
H m ax
3-
J
o-
74%
III
IV
Fig. 9. C h ang es in a c tu a l (H’) a n d p o ten tial (Hmax)
s p e c ie s d iv e rsity in c o m m u n itie s of s u c k in g
p h y to p h ag es a n d th e degree of realisa tio n (J) of
p o ten tial diversity of th e co m m u n ities in th e sere
of th e p in e forest
T h e s im ila rity of s p e c ie s c o m p o sitio n in all th e su c k in g p h y to p h a g e
co m m u n ities o c c u rrin g in s ta n d s of all age c lasses in Puszcza Białow ieska, a s
well a s th e d ifferences in a b u n d a n c e a n d com m unity s tru c tu re described above
su b s ta n tia lly affected species diversity m e a su re d w ith S h a n n o n a n d W eaver’s
index (Fig. 9). In th e sere u n d e r investigation, species diversity of b o th ap h id
a n d h e te r o p te r a n c o m m u n itie s d e c re a s e d w ith g re a te r sp e c ie s d iv ersity
differences n o ted in b o th co m m u n ities betw een young grow th a n d older s ta n d s
(pole wood, m a tu re forest) th a t betw een the la tte r two. C icadas exhibited a
different re sp o n se to th e p ro cess of su ccessio n , reflected in a n in crease in species
diversity w ith th e grow ing age of th e forest sta n d . In th a t case the m a tu re forest
is a stag e of g re a te r p o ten tial species diversity th a n the y ounger stages: y oung
sta n d a n d th e pole wood. In th e pine forest sere stu d ied in Puszcza B iałow ieska,
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P. T rojan et al.
th e degree to w hich species diversity approxim ated th e po ten tial of th e fau n al
system varied from co m m u n ity to com m unity. In ap h id an d h e te ro p te ra n
co m m u n ities, the degree of realisatio n of diversity (m easured w ith Pielou’s index)
is h ig h est in young s ta n d s, w hile in th e case of leafhoppers th e h ig h est v a lu e s
w ere reg istered in m a tu re s ta n d s (Fig. 9). As the forest s ta n d grows older,
differences a re also noted in th e d istrib u tio n of individual species’ s h a re s in th e ir
co m m u n ities. D ecreases in th e degree of uniform ity of individual species
d istrib u tio n w ere recorded in ap h id a n d h e te ro p te ran co m m unities, while
in c re a se s in species p ro p o rtio n s’ uniform ity w ith the grow ing age of th e s ta n d
w ere seen in com m unities of leafhoppers.
A nalysis of s tru c tu re s of th e co m m u n ities of su c k in g p h y to p h ag es show ed
th a t they do n o t fit an y of th e five d istrib u tio n m odels form ed so far i.e. th e
geom etric series, broken stick d istrib u tio n model, logarithm ic series, lognorm al
d istrib u tio n a n d negative binom ial d istrib u tio n .
T he pro cess of form ation of th e u n iq u e su ck in g phytophage fa u n a of th e
canopies of tre e s in pine forests s ta rts in young s ta n d s (or possibly even earlier,
in the c u ltu re stage), a s is in dicated by th e rem ark ab ly high sim ilarity of species
com position of co m m u n ities o ccu rrin g in different age classes of th e pine forests
of Puszcza Białow ieska. The tra n s itio n betw een consecutive su c ce ssio n stag es
of th e pine forest is m ark ed by c h an g es (although th e ir ra te a n d direction differ
from group to group) in a b u n d a n c e a n d d om inance s tru c tu re of all th e su ck in g
phy to p h ag e com m unities stu d ie d . G enerally, a s th e forest s ta n d grow s older,
both th e a b u n d a n c e a n d p ro p o rtio n s of species a sso c ia te d w ith pine ten d to
in crease, w hile th e a b u n d a n c e a n d p ro portion of poly- a n d oligotrophic species
are red u ced . The form ation of a d o m in an ce s tru c tu re c h a ra c te ristic of pine
canopies is n early com pleted in 50-year-old forest s ta n d s (aphids, h e te ro p te ra n
bugs). In leafhoppers th e p ro cess is slower, concluding in s ta n d s older th a n 50
years.
C hew ing p h y to p h ag es
115 species of sawflies (H ym enoptera, S y m p h y ta ), 23 species of leaf beetles
(Coleoptera, C hrysom elidae) 46 species of weevils (Coleoptera. Curculionidae) a n d
122 species of bu tterflies (Lepidoptera) (H u f l e jt u n p u b lish e d d a ta , W ą so w sk a
u n p u b lis h e d d a ta , W iśn ie w sk a u n p u b lis h e d d a ta , B u sz k o a n d W iniarska
u n p u b lis h e d data) w ere recorded in th e canopies of pine forest s ta n d s in P uszcza
B iałow ieska in all the forest s ta n d age c la sse s stu d ie d (young s ta n d , pole wood,
m a tu re forest). In th e m ajority of th e g ro u p s of chew ing p h y to p h ag es u n d e r
stu d y , th e n u m b e r of species form ing th e co m m u n ities ten d ed to d e crease a s
th e forest s ta n d grew older. T he rea so n for th e decrease in th e n u m b e r of species
w as th e sam e for all co m m unities. In y o u n g er s ta n d s (particularly in y o ung
sta n d s), th e re w as a significant n u m b e r of species asso ciated trophically, often
only in th e larval stage, w ith p la n ts occu rrin g in th e g ro u n d cover of th e pine
forest stu d ie d in Puszcza B iałow ieska: 30 su c h species w ere recorded in th e
co m m u n ity of sawflies in y o u n g sta n d s , 26 in the pole wood a n d 20 in m a tu re
sta n d s . The respective figures for th e C urculionid com m unity w ere 12 species
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Secondary su cc e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
25
in y o u n g sta n d s , 6 in th e pole wood a n d 7 in m a tu re s ta n d s. On th e o th er h a n d ,
th e n u m b e r of species of bu tterflies grow s steadily a s the s ta n d develops w ith
70 species recorded in young sta n d s , 91 in th e pole wood a n d 100 in m a tu re
s ta n d s (Fig. 10).
CHEW ING PHYTOPHAGES
100
9080-
3020IV
—«■— S y m p h y ta
C u rc u lio n id a
C h ryso m e lid
- e> - Le p id o p tera
Fig. 10. C h an g es in th e n u m b e r of species of chew ing p h y to p h ag es in pine forests of th ree age classes
of th e forest sta n d
T he sp ecies com position w as sim ilar in all th e co m m u n ities of biting
ph y to p h ag es studied. The h ig h est sim ilarity of species com position w as recorded
for co m m u n ities of bu tterflies an d saw flies w ith S o eren sen 's sim ilarity indices
ran g ed from 68% to 79% for individual com m unities, while the low est values
w ere ob tain ed for com m unities of C hrysom elidae (47%-58%). The values of
sim ilarity indices (So) of individual co m m u n ities in h ab itin g canopies of p ines
w ere largely d ep en d en t on local differences in the species com position of th e
g ro u n d cover an d th e u n d e rsto re y vegetation. It w as p a rticu larly evident in th e
case of in se c t species of high trophic specificity. For exam ple, H ypera nigrirostds
[Curculionidae], a species feeding on clover, w as registered in young a n d m atu re
s ta n d s (in th e la tte r it w as even a d o m in a n t species), b u t did n o t o ccur in th e
pole wood. Of th ree species asso ciated w ith cruciferous p la n ts an d recorded in
young sta n d s , only one w as found in th e pole wood a n d none in th e m a tu re
forest. In leaf beetle com m unities, virtually all th e species c a u g h t in canopies of
tre e s in th e pine forests feed on leaves of e ith e r h e rb a c e o u s p la n ts or d ecid u o u s
tre e s (only 1 o u t of 27 species found in Puszcza B iałow ieska feeds on pine). As
a resu lt, th e species com position of co m m u n ities of beetles of th is fam ily w as
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26
P. T rojan et al.
m u ch m ore d e p e n d en t on local species ric h n e ss of the g ro u n d cover an d on the
presen ce of various tree species a n d d ecid u o u s s h ru b s in th e u n d ersto rey .
C om m unities found in older s ta n d s (pole wood, m a tu re forest) exhibited
co n sp ic u o u s sim ilarity b o th in species com position (sim ilarity indices ranged
from 58% for leaf beetle co m m u n ities to 77% for sawflies) a n d th e s tru c tu re of
dom inance of individual com m unities.
On th e o th e r h a n d , a b u n d a n c e of th e saw fly, leaf b e e tle a n d weevil
c o m m u n ities stu d ied did n o t exhibit significant differences related to th e age of
th e s ta n d (Fig. 11).
C H EW IN G PHYTOPHAGES
1. 2-
x
<D
-a
c
cO
re
£
.a
re
0. 6 -
0.40. 2 -
II
—<■— S y m p h y ta
IV
C u rc u lio n id a e
C h ryso m e lid a e
Fig. 11. C h an g es in th e a b u n d a n c e of chew ing p h y to p h ag es in p ine fo rests of th re e age c la sse s of
th e forest sta n d
In th e sere stu d ied ch an g es in d om inance s tru c tu re occu rred in all the
c o m m u n it i e s of c h e w in g p h y t o p h a g e s u n d e r s tu d y . T h e y c o n s is te d
pred o m in an tly in shifts in th e ra n k of the d o m in a n t species.
In sawfly com m unities, th e p roportion of th e m onophage of pine X y e la julii
(larvae feed on m ale efflorescence) rose w ith th e growing age of th e s ta n d (from
2% in y o u n g s ta n d s to 16% in m a tu re stan d s), while the reverse p h en o m en o n
w as observed for A thalia rosae (from 33% in y o ung s ta n d s to a b o u t 10% in
m a tu re forests). A. rosae is a very com m on species o ccu rrin g in v a rio u s h a b ita ts
(Fig. 12). The proportion of Pristiphora abietina, th e larvae of w hich feed on
n eedles of sp ru ce, rem ained fairly c o n s ta n t in all th e age c la sse s of th e forest
sta n d .
A significant tra n sfo rm atio n of th e d om inance s tru c tu re in co m m u n ities of
beetles of th e family Curculionidae could be seen betw een th e y o u n g grow th and
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S econdary su c c e ssio n of fa u n a in th e p ine forests of P uszcza B iałow ieska
27
Fig. 12. D om inance s tru c tu re of c o m m u n ities of saw flies {Hymenoptera, S ym p h yta ) in can o p ies of
p ine in forest s ta n d s of v ario u s ages in P uszcza B iałow ieska; 1 - A thalia rosae, 2 - Pristiphora abietina,
3 - X yelajulii, 4 - R hogogaster picta. 5 - A thalia bicolor. 6 - Pristiphora pseudodecipiens, 7 - N em atus
dispar. 8 - Dineara viridorsata. 9 - Shaliphora parva, 10 - F en u sa pusilla, 11 - A thalia circularis
C urculionidae
Fig. 13. D om inance s tru c tu r e of co m m u n ities of xxx (Coleoptera, Curculionidae ) in can o p ie s o f pine
in forest s ta n d s of v ario u s ages in P uszcza Białow ieska; 1 - B rachonyx pineti, 2 - R hinom acer
attelaboides, 3 - A nthonom u s phyllocola. 4 - A ttela b u s nitens, 5 - M agdalis linearis. 6 - Straphosom a
capitatum , 7 - D ayd rh yn ch u s austriacus. 8 - H ypera nigrirostris
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P. T ro jan e t al.
pole wood stages. It c o n sisted in a decrease in th e ra n k of B rachonyx pineti, a
species feeding on pine needles, for th e benefit of Rhinom acer attelahoides, w hich
develops in m ale efflorescence of coniferous trees. In y oung s ta n d s , B. pineti w as
th e e u d o m in a n t w ith a p ro p o rtio n of 47.4% , an d w as followed by R. attelaboid.es
(13%) (Fig. 13). In older s ta n d s (pole wood, m a tu re sta n d s) b o th species w ere
co -d o m in an t.
In th e C hrysom elidae beetle co m m u n ities th e proportion of Galerucella lineola
in cre ase d a s th e s ta n d grew older. T he im agines of th is sp ecies feed on d eciduous
tre e s a n d s h ru b s . On th e o th e r h a n d , th ere w as a re d u c tio n in th e sh a re of
A ph to n a euphorbiae, a species a sso c ia te d w ith th e g ro u n d cover vegetation
(Fig. 14).
C hrysom elidae
5(H
45
4035
3025
20
10IV
Fig. 14. D om inance s tru c tu re of c o m m u n itie s of leaf beetles (Coleoptera , C h rysom elidae ) in canopies
of pine in forest s ta n d s of vario u s a g es in P u szcza Białowieska; 1 - A p h to n a euphorbiae, 2 - Galerucella
lineola, 3 - Longitarsus parvulus, 4 - C ryptocephalus parvulus, 5 - C ryptocephalus Jlavipes, 6 Longitarsus longiseta, 7 - Phyllotreta nem orum , 8 - Phratora laticollis, 9 - C ryptocephalus saliceti.
In Lepidoptera co m m u n ities th e pine m onophage, O lethreutes bifasciana is
th e d o m in a n t species in all su c c e ssio n stages. T here is also a h igh proportion
of species a sso ciated w ith th e g ro u n d cover or even d e c id u o u s tre e s (Rhopobota
naevana, A potom is betuletana, Alcis repandata). A. betu leta n a is only found in
m a tu re s ta n d s , while R. n a e v a n a a n d A. rep cm d atab elo n g to th e d o m in a n t group
in all age classes. C h a ra c te ristic is a n extrem ely low p ro p o rtio n of species
recognised a s p rim a iy pine foliophages (L ym antria m onacha a n d B npalus
piniariits).
T he v a lu e s of S h a n n o n a n d W eaver’s index co rresp o n d in g to total species
diversity of individual c o m m u n ities of chew ing p h y to p h ag es d isplay relatively
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S eco n d ary successio n of fa u n a in th e p in e fo re sts of P uszcza B iałow ieska
29
insig n ifican t differences in relation to th e age of th e sta n d (Fig. 15). D espite th e
sm all differences, the highest species diversity in all the com m unities of chew ing
p h y to p h ag e s w as recorded in yo u n g er sta n d s : y o ung s ta n d s (Curculionidae) or
th e pole wood (S y m p h y ta , C hrysom elidae)
In th e se re of th e pine forest of Puszcza B iałow ieska potential species diversity
(m easu red w ith Pielou's index) ranged from 7 2 -8 2 % for sawfly com m unities, a n d
76-85% for leaf beetle com m unities. S om ew hat low er values of p o tential species
diversity w ere o btained for Curculionidae co m m u n ities, b u t even in th is case,
th e v a lu e s of Pielou’s index w ere sim ilar in all s ta n d age cla sse s (Fig. 15).
Symphyta
Curculionidae
H m ax
H m ax
0j------II
III
IV
Symphyta
II
III
IV
Fig. 15. C h a n g es in a c tu a l ( H ) a n d p o ten tial ( H m a x )
s p e c ie s d iv e rsity in c o m m u n itie s o f c h ew in g
p h y to p h a g e s a n d th e degree of realisa tio n (J) of
p o ten tial diversity of th e co m m u n ities in th e sere
of th e p in e forest
A nalysis of s tru c tu re s of th e co m m u n ities of chew ing p h y tophages show ed
th a t they do no t fit an y of th e five m odels of a b u n d a n c e d istrib u tio n am ong
individual sp ecies i.e. the geom etric series b ro k en stick d istrib u tio n se rie s,
logarithm ic, lognorm al d istrib u tio n a n d negative binom ial d istrib u tio n .
T he p ro cess of form ation of th e chew ing p h y to p h ag e fa u n a specific to th e pine
forests of P uszcza Białow ieska ta k e s place in th e pole wood stage, a s is in d icated
by th e c o n sid erab le sim ilarity of species com position a n d dom inance s tr u c tu r e s
in co m m u n ities o ccurring in the pole wood a n d m a tu re s ta n d s.
In all th e chew ing p h y to p h ag esr c o m m u n ities stu d ied tra n sitio n s betw een
consecutive sta g e s of th e pine forest sere a re m ark ed by ch an g es in a b u n d a n c e
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P. T rojan et al.
30
an d d o m in an ce s tru c tu re . The c h a n g es in th e dom inance s tr u c tu r e c o n sist
p red o m in an tly in shifts in th e position, or ran k , of species p red o m in atin g in a
given com m unity. In weevil an d sawfly com m unities older s ta n d s w itness an
in crease in th e prop o rtio n s of species w hose larvae feed on m ale efflorescence
of pine; in the leaf beetles com m unity th ere is a d ecrease in th e proportion of
th e species feeding on d ecid u o u s tre e leaves. As th e s ta n d grows older, a decrease
is observed in th e n u m b e r of species trophically asso ciated (often only in th e
larval stage) w ith h erb a c e o u s p la n ts th a t only p e n e tra te canopies of pines. T his
holds tru e b oth for m any species of sawflies, th e m ajority of a d u lt form s of w hich
are m ellitophagous, a n d m an y species of beetles, w hich are p h y to p h ag o u s b o th
in th e larval an d a d u lt stages.
X ylophages a n d cam biophages
The consecutive stag es of the pine forest sere cre a te different conditions for
the developm ent of xylophages a n d cam biophages, w hich develop in or u n d e r
th e b ark , or in th e wood of various species of tre e s an d s h ru b s . Individual species
of th ese beetles are asso ciated w ith specific p la n t species, in h a b itin g different
layers from roots, stem s, b ra n c h e s to th in twigs. They also settle in decaying
stu m p s, dam aged d u rin g forest felling a n d clearance, a n d in wood rem aining
after th e felling. V arious species of th ese in sects show preference for certain
p h a se s of b a rk decom position. In ju ries to th e foot of a tree, w hich w eaken the
e n tire o rg an ism , in cre ase in severity a s the s ta n d grows, pro m o tin g the
developm ent of xylo- a n d cam biophages.
XYLO- and CAM BIO PHAG ES
30
25-
o
a>
10-
III
—■ — S co lytid ae
C e ra m b y c id a e
IV
B up restid ae
Fig. 16. C h an g es in th e n u m b e r of sp ecies of xylophages a n d cam b io p h ag es in p ine fo rests of three
age c la s se s of th e forest sta n d
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S eco n d ary su ccessio n of fa u n a in th e p ine forests of P uszcza B iałow ieska
31
XYLO- and CAM BIOPHAGES
1200
1000 x
<i>
800-
3
600400200-
II
—■ — S colytid ae
IV
C e ra m b y c id a e
B up restid ae
Fig. 17. C h an g es in th e a b u n d a n c e of xylophages a n d cam biophages in pine forests of th re e age
c la s se s of th e forest sta n d
B ark beetles (Scolytidae)
B ark beetle com m unities are c h a ra c te rise d by a stead y rise in th e n u m b e r of
species th ro u g h o u t the sere (Fig. 16). T he lowest n u m b e r of species - 10 - w as
recorded in th e c u ltu re stage. T hose species show a n affinity for h a b ita ts form ing
im m ediately after felling a n d a re m ore a p t to in h a b it fresh s tu m p s (Hylurgos
p a llia tu s, H yla stes ater,; H. opacus a n d H ylurgos lignipera). Som e of th em prefer
th e u n d e rg ro u n d p a rt of th e s tu m p d u e to higher m o istu re (H ylastes a ter a n d
H. opacus), o th ers in h ab it th e whole of th e stu m p (Mokrzycki, oral inform ation).
The species dam age th e b a rk an d m ay o ccu r in co nsiderable a b u n d a n c e in
3 -5 -y ear-o ld c u ltu res. In th e y oung s ta n d stage, th e species com position of th e
b a rk beetle com m unity is even richer: all the species o ccurring in th e c u ltu re
are still p rese n t, th ere a p p e a r species settling on tru n k s , s u c h a s P ityogenes
quadridens, or c h a ra c te ristic of older sta n d s , e.g. Ips se xd e n ta tu s. In older
s ta n d s th e species com position of th e co m m u n ities becom es ric h e r still, owing
to, a m o n g o th e r factors, a n influx of species se ttlin g on b ra n c h e s . The
co m m u n ities m ost sim ilar in species com position are th o se in h ab itin g pole wood
an d m a tu re s ta n d s (S oerensen's sim ilarity index ranged am ong 81 to 100%).
The low est a b u n d a n c e is noted for co m m u n ities in young sta n d s . In older sta n d s ,
the a b u n d a n c e in creases steadily, achieving a m axim um in th e m a tu re stage
(Fig. 17). A nalysis of th e s tru c tu re of dom inance reveals th a t th e s tru c tu re of
the co m m u n ity is tran sfo rm ed in th e co u rse of su ccessio n since different species
are d o m in a n t in th e co n secu tiv e age c la sse s (Fig. 18). T he sim ilarity of
dom inance s tru c tu re s of Scolytidae co m m u n ities in h ab itin g older s ta n d s (pole
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32
P. T rojan et al.
wood, m a tu re stands) is high. In yo u n g er s ta n d s (culture, y o ung stan d s) on th e
o th er h a n d , com m unity s tru c tu re s differ b oth from one a n o th e r a n d from
co m m u n ities of older sta n d s . It is therefore suggested th a t th e co m m u n ities
becom e s ta b le a fte r th e s ta n d re a c h e s th e pole w ood sta g e ( M o k r z y c k i
u n p u b lish e d data).
S colytidae
Fig. 18. D om inance s tru c tu re of c o m m u n ities of b a rk b eetles (Scolytidae) in forest s ta n d s of v a rio u s
ages in P uszcza Białowieska; 1 - C rypturgus hispidulus, 2 - C rypturgus cinereus, 3 - Trypodendron
lineatum , 4 - Tom icus piniperda, 5 - Ips a cum inalus, 6 - Ips se x d e n ta tu s , 7 - P ityogenes b id en ta tu s,
8 - O rthom icus suturalis, 9 - P ityogenes quadridens, 10 - H yla stes opacns, 11 - O rthom icus laricis,
12 - H ylurgops palliatus, 13 - H yla stes a ter , 14 - H ylurgus lignipedra
Longicorns (C eram bycidae)
L ongicorns co m m u n ities are c h a ra c te rise d by a n in crease in th e n u m b e r of
species in th e pole wood a n d m a tu re forest stage a s com pared to y o u n g s ta n d s
(Fig. 16). A sim ilar n u m b e r of species in every age c la ss of pine forest (young
sta n d , pole wood, m a tu re stand) w as recorded only in th e case of tree canopies.
The c u ltu re stag e is ch a ra c te rise d by a relatively rich C eram bycidae taxocoene,
w hich is a sso ciated w ith th e taxon settling s tu m p s a n d w h a t h a s rem ain ed after
clearfelling. The species com position in th e young s ta n d stage is th e p oorest a n d
m ost different in com parison to co m m u n ities o ccurring in older s ta n d s . In the
pole wood, th e Longicorn beetle co m m u n ity is sim ilar in species com position to
th e co m m u n ity in h ab itin g n a tu ra l m a tu re s ta n d s (S oerensen’s index is 67%)
( G u t o w s k i u n p u b lish e d data). In th e sere u n d e r stu d y , th e a b u n d a n c e of
C eram bycidae re a c h e s th e low est v alu es in young sta n d s , an d in c re a se s in the
pole wood a n d m a tu re s ta n d s (Fig. 17). In th e C eram bycidae c o m m u n ities u n d e r
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S econdary su cc e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
33
investigation, A se m u m striatum is d o m in a n t in the c u ltu re stage (16.5%),
Pogonocherus decoratus in young s ta n d s (32.8%), Corym bia rubra in th e pole
w ood (15.0%) an d R hagium inquisitor in th e m a tu re s ta n d (17.7%) (Fig. 19).
C e ra m b ycid ae
/zzzzm
Fig. 19. D om inance s tru c tu re of co m m u n ities of C eram bycidae in forest s ta n d s o f v a rio u s ag es in
P uszcza B iałowieska; 1 - R hagium inquisitor, 2 - Stenurella m elanura, 3 - M olorchus minor, 4 A rhopalus rusticus, 5 - Tetropium ca sta n eu m , 6 - Pogonecherus fa scic u la tu s, 7 - A canthocinus
griseus, 8 - A se m u m striataum , 9 - Callidium coriaceum, 10 - Corym bia rubra, 11 - Pogonecherus
decoratus, 12 - Spondylis buprestoides, 13 - Callidium violaceum
B u p re stid s (B uprestidae)
T he h ig h est n u m b e r of species is recorded in b u p re stid c o m m u n ities from
pine c u ltu re s a n d significantly th in n e d m a tu re forests (Fig. 16). The h ig h est
sim ilarity of species com position w as noted for co m m u n ities of c u ltu re s, pole
wood a n d m a tu re s ta n d s (S oerensen’s sim ilarity index ranged em ong 67 to 70%).
The a b u n d a n c e of B uprestidae co m m u n ity is th e h ig h est in th e c u ltu re stage
(Fig. 17). T he sam e species, nam ely A n th a xia quadripunctata , is d o m in a n t in all
th e age c la s s e s of th e pine forest se re a n d th e c o m m u n itie s in h a b itin g
consecutive stag es of forest s ta n d developm ent are c h a ra c te rise d by relatively
high sim ilarity of do m in an ce s tru c tu re s (Fig. 20). The m ost sim ilar in th is resp ect
are co m m u n ities occupying c u ltu re s a n d y oung s ta n d s, w hile th e g re a te st
differences in dom in an ce s tru c tu re s w ere recorded for th e c u ltu re a n d m a tu re
sta n d co m m u n ities (G utow ski u n p u b lish e d data).
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34
P. T rojan et al.
Buprestidae
869
67^40882
Fig. 20. D om inance s tru c tu re of co m m u n ities of b u p re stid s (B u p restid a e ) in forest s ta n d s of v ario u s
ages in P uszcza Białowieska; 1 - A n th a xia quadripunctata, 2 - Chalcophora m ariana, 3 - B uprestis
rustica, 4 - A n th a xia subm ontana, 5 - B uprestis haem horroidalis, 6 - Agrilus sulcicollis, 7 - T rachys
m inuta, 8 - P haenops cyanea, 9 - B uprestis octoguttata, 10 - Agrilus a n g u stu lu s
A nalysis of c h a n g es in xylo- a n d cam biophage co m m u n ities
in th e co u rse of su c ce ssio n
T ra n sfo rm a tio n s of the com m unities of th e xylo- an d cam biophage groups
stu d ie d a re a little different in each group. T he g re a te st n u m b e r of species is
noted for th e xylo- an d cam biophage com m unities o ccurring in m a tu re s ta n d s
(Fig. 16). B uprestidae an d C eram bycidae a re c h a ra c te rise d by relatively high
species ric h n e ss in th e c u ltu re stage a s well, b u t Scolytidae a re re p re sen te d by
th e low est n u m b e r of species a t th is stage. Young s ta n d s a re hom e to th e low est
n u m b e rs of species of B uprestidae a n d C eram bycidae. T he a b u n d a n c e of
Scolytidae an d C eram bycidae in cre ase s tow ard s m ore m a tu re sta n d s , a n d it is
only th e m arkedly x erophilous B uprestidae th a t a re m ost a b u n d a n t in the
c u ltu re stage. In th e sere u n d e r investigation, th e d o m in a n t species group of
Scolytidae an d C eram bycidae u n d erg o es s u b s ta n tia l tra n sfo rm atio n . O ther
species are d o m in an t in yo u n g er s ta n d s th a n prevail in older ones, a lth o u g h the
form er d o m in a n t group is not com pletely su p p la n te d . B uprestidae a re th e only
tax o n w here one species (Anthaxia quadripunctata) is d o m in a n t in all the
co m m u n ities studied. In Scolytidae, C eram bycidae a n d B u p restid a e , th e group
of d o m in a n ts is com posed of a n u m b e r of species in every age class.
A nalysis of sim ilarity of em pirical xylo- a n d cam biophage co m m u n ities w ith
th e five theoretical sta tistic a l d istrib u tio n s show ed th a t th e d istrib u tio n of
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S eco n d ary su ccessio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
sp ecies in th e co m m u n ities exam ined is closest to th e negative binom ial
d istrib u tio n (Scolytidae a n d Ceram bycidae). In th e case of C eram bycidae it is
also sim ilar to th e logarithm ic series (com m unities of c u ltu re s, pole wood an d
m a tu re forest) a n d to the geom etric series (young stan d s). In som e in sta n c e s,
th e e stim ated n u m b e r of species is h ig h er th a n th e one a c tu a lly recorded
(C eram bycidae) w hich su g g ests the p resen ce of h ig h er species cap acity of th e
h a b ita t. In Scolytidae the estim ated n u m b e r of species co rre sp o n d s to th e a c tu a l
one.
Species diversity, m ea su red w ith S h a n n o n an d W eaver’s index, is h ig h est in
m a tu re sta g e s (Scolytidae - pole wood, B uprestidae a n d C eram bycidae - m a tu re
stan d s) (Fig. 21). The degree of realisatio n of potential species diversity (Hmax)
in c re a se s in generalfor th e co u rse of of th e g ro u p s stu d ied in each su c ce ssio n
stage.
Scolytidae
Cerambycidae
H m ax
3.5-1Hmax
IV
Buprestidae
H m ax
97%
i! 7 6 %
53%
Fig. 21. C han g es in a c tu a l ( H ’) a n d p o ten tial ( H m a x )
sp ecies diversity in c o m m u n ities of xylophages
a n d cam b io p h ag es a n d th e degree of realisatio n
(J) of p o ten tial diversity of th e c o m m u n itie s in th e
sere of th e pine forest
A nalysis of s tru c tu re s of th e co m m u n ities of selected xylo- a n d cam b io p h ag es
p erm its a fuller d escription of th e c h a n g es tak in g place in th e su c c e ssio n sta g e s
stu d ied in th e p in e forest sere.
T he c u ltu re stag e c re a te s m ost favourable conditions for B u p restid a e to reach
a high a b u n d a n c e a n d a n u m b e r of species sim ilar to th a t of th e m a tu re forest.
C eram bycidae are also c h a ra c te rise d by relatively rich fa u n a a t th is stage. T his
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P. T rojan et al.
is related to th e possibility of settlin g in stu m p s, clearfelling rem a in s a n d to high
insolation w hich, is p a rticu larly cru cial for the a p p e a ra n c e of B uprestidae.
The y o u n g s ta n d s h a rb o u r th e low est n u m b e rs of C eram bycidae a n d
B uprestidae species. The decaying s tu m p s in young s ta n d s a re no longer a
su ita b le h a b ita t for m ost species of C eram bycidae a n d B u p restid a e, a n d the th in
ste m s a n d b ra n c h e s com bined w ith low exposure to su n lig h t do n o t provide a
good feeding b a sis for a w ider ran g e of species. The situ a tio n m ay change if
deadw ood a p p e a rs in younger age classes. Cam bio- a n d xylophagous species of
b oth g ro u p s m ay th e n in crease th e ir n u m b ers. B ark beetle co m m u n ities have
rich er species com position a t th is stage. The species com position of b a rk beetle
c o m m u n itie s b ecom es ste a d ily ric h e r a s th e fo rest grow s older. S pecies
c h a ra c te ristic of the s u b s e q u e n t stag es s ta rt to a p p e a r while th e species
occurring in younger age c lasses do n o t retrea t, b u t only change th eir positions
in the s tru c tu re of dom inance.
The pole wood stage is a p re p a ra to ry p h a se for th e form ation of com m unities
ch a ra c te ristic of n a tu ra l m a tu re pine forests. Especially im p o rta n t for the
developm ent of xylophages a n d cam b io p h ag es is th e a p p e a ra n c e of litter, dead
trees, developm ent of th e h erb layer a n d a n in crease in in so latio n a t th is stage.
The a n a ly sis of the selected g ro u p s of xylo- a n d cam biophages revealed th a t
th e beetle g ro u p s stu d ied becom e stab ilised in older s ta n d s (pole wood). The
xylo- a n d cam biophage fa u n a o ccu rrin g a t th is stage m ost closely resem b les the
fa u n a of a n a tu ra l m a tu re forest.
5 .2 .2 . E p ip h y tic p redators
P red ato rs preying in th e g ro u n d cover, s h ru b an d tree canopy layers m ay be
divided into two g ro u p s playing different ecological roles of specialised a n d
n o n -specialised pred ato rs. Prim e exam ples of no n -sp ecialised p re d a to rs (or
form s w ith hard ly -m ark ed trophic specialisation) a b u n d a n tly re p re sen te d in the
c a n o p y lay e r of p in e fo re st s ta n d s a re s p id e rs (Aranei), so ld ie r b e e tles
(C antharidae. Coleoptera), robber-flies (Asilidae, Diptera) a n d digger w asp s
(S p h ecid a e, H ym enoptera). In specialised p red a to rs, one can find g ro u p s of
a p h id o p h a g o u s species belonging to sy rp h id s (S y rp h id a e , Diptera), digger w asp s
(Sphecidae. H ym enoptera), n e u ro p te ra .n s (Neuropteroidea) a n d la d y b ir d s
(Coccinellidae, Coleoptera).
In pine forests th ese g ro u p s of p re d a to rs exhibit high species ric h n e ss, a t the
sam e tim e o ccurring in high a b u n d a n c e s to play a d oubtlessly significant role
in the functioning of th e biocenosis. From am ong no n -sp ecialised p red ato rs,
large-sized form s m ay even a tta c k larger anim als, a n d th e m arkedly sclerotised
protective layer of th eir victim s’ bodies poses no o bstacle for them . A dult
robber-flies a n d digger w asps, w hich are good flyers, p e n e tra te into all vegetation
layers w hile h u n tin g .
Web sp id e rs of pine s ta n d s have been stu d ie d by Ł u c z a k (1963), sp id ers of
m iddle-aged s ta n d s have been investigated by D z i a b a s z e w s k i (1976), a n d sp id ers
living in forest s ta n d s of v ario u s ages have been an aly sed by S t e r z y ń s k a a n d
Ś l e p o W r o ń s k i (in press).
Pine forest robber-flies have n o t been dealt w ith in a se p a ra te p a p e r to date.
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S econdary su ccessio n of fa u n a in th e p ine forests of P uszcza B iałow ieska
37
T h e s p e c ie s c o m p o s itio n a n d d o m in a n c e s t r u c t u r e of d ig g e r w a s p
c o m m u n ities of th ese ecosystem s have been an aly sed in p a p e rs devoted to
W arsaw en v iro n s ( S k i b i ń s k a 1981) a n d older pine s ta n d s in B yelo ru ssia
( S h l y a k h t e n o k in p ress).
T he species com position a n d do m in an ce s tru c tu re of syrphid co m m u n ities
have been p rese n ted in a p a p e r on th e pine forest of Puszcza K am pinoska and
B iałołęka D w orska ( B a ń k o w s k a 1982). To date, th ere h a s been only one p a p e r
w h ich co m p ares the course of secondary successio n of syrphid co m m u n ities in
th re e age c lasses of pine forest s ta n d s in th re e different pine forests ( B a ń k o w s k a
in press).
L adybirds of older s ta n d s of pine have been an alysed by H o r e g o t t (1960),
K l a u s n i t z e r (1968), K lo m p an d T e e r i n k (1973) a n d C z e c h o w s k a (1981).
Few p a p e rs have been devoted to n e u ro p te ra n s of pine forests ( C z e c h o w s k a
1985) a n d n u ie of th em is concerned w ith succession.
S p id ers (Aranei)
75 sp id er species occur in th e pine forest s ta n d s stu d ied in all th e age classes,
w ith 51 species in young sta n d s , 31 in th e pole wood a n d 3 9 in m a tu re s ta n d s
(Fig. 22) (S terzy ń sk a u n p u b lish e d data). 15 species w ere found to occur in every
su c ce ssio n stage in th e pine forest, a c co u n tin g for a b o u t 20% of the sp id e r fa u n a
collected w ith M oericke’s pitfall tra p s in th e stu d y a reas. The species com position
of sp id ers in young pine s ta n d s is c h a ra c te rise d by a m u ch g reater u n iq u e n e s s
t h a t th e fa u n a registered in older sta n d s . The degree of species com position
sim ilarity (So) of th e young s ta n d a n d pole wood co m m u n ities w as 46% , a n d of
ZO O PH AG ES
55
504540-
a.
35302520 -
15IV
—'■— A silidae
- e> -
A ranei
S ph ecid ae
C o c cin e llid a e - * * - C an th a rid ae
Fig. 22. C hanges in th e n u m b e r of species of p re d a to rs in pine forests of th ree age c la s se s of th e
forest sta n d
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38
P. T rojan et al.
sp id er com m unities of young a n d m a tu re s ta n d s - 44%, while th e figure for th e
pole wood an d m a tu re s ta n d s w as 60%.
The differences in a b u n d a n c e of sp id er co m m u n ities in pine s ta n d s of various
ages in Puszcza B iałow ieska were n o t a s significant a s differences in th e n u m b er
of species. The m ost a b u n d a n t com m unity lived in young sta n d s , while in pole
wood th e com m unity a b u n d a n c e w as lower by a b o u t 40% a n d in m a tu re forests
- by 30% (Fig. 23).
ZO O PH AG ES
60
50x
|
40-
c
30-
E
20
T03CJ
03
10IV
—
Asi l i dae
-■+■■■
C o ccinellidae —
A ranei
S p h eo id ae
Cant har i dae
Fig. 23. C hanges in th e a b u n d a n c e o f p re d a to rs in p ine fo rests of th re e age c la sse s of th e forest sta n d
The s tru c tu re s of d om inance of sp id er co m m u n ities w ere sim ilar in all the
su c ce ssio n stag es of pine forests (Fig. 24). H eliophanus d u b iu s w as d o m in a n t in
y oung s ta n d s an d th e pole wood, an d su b d o m in a n t in m a tu re s ta n d s , w here
Diaea dorsata w as th e d o m in a n t species. From am ong th e 10 m o st a b u n d a n t
species c a u g h t in canopies of pines, a s m any a s 7 belonged to the g roup of species
prevailing in all th e forest s ta n d s stu d ied . The species included: H eliophanus
dubius. A n y p h a e n a accentuata, Trem atocephalus cd sia tu s. Theridion tinctum.
Diaea dorsata. X y stic u s cristatus a n d A te a sturm i. The s h a re of th is g ro u p w as
significant in each com m unity exceeding 45% in young sta n d s , rea c h in g nearly
65% in pole wood a n d a b o u t 50% in m a tu re sta n d s . All of th e above species are
forest polytopes, asso ciated m ostly w ith coniferous forests. O nly T. cristatus is
considered to o ccu r m ore often in d ecid u o u s forests ( S t e r z y ń s k a u n p u b lish e d
data).
As th e s ta n d s grew older, th e a b u n d a n c e of H. dubius, A. accentuata. T.
cristatus an d T. tinctum decreased while th e a b u n d a n c e of D. d o rsa ta a n d A.
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S eco n d ary su ccessio n of fa u n a in th e p ine fo rests of P uszcza B iałow ieska
39
Aranei
Fig. 24. D om inance s tru c tu re of co m m u n ities of sp id ers (A ranei ) in forest s ta n d s of v a rio u s ages in
P uszcza Białow ieska; 1 - H eliophanus diibius, 2 - Euryclubiona s u b s u lta n s , 3 - D en d ryp h a n tes ru d is ,
4 - A n y p h a e n a a ccentuata , 5 - Trem atocephalus c rista tu s , 6 - Entelecara congenerci, 7 - Theridion
tinctum , 8 - A tea sturm i, 9 - Diaea dorsata, 10 - D ictyna arundinacea
sturm i rose to g eth er w ith th eir ra n k s in th e com m unity a s they occupied higher
positions in th e s tru c tu re of d om inance in older s ta n d s. In the case of X. cristatus
th e a b u n d a n c e rem ained th e sam e in all the pine s ta n d s stu d ied , while its sh a re
in older s ta n d s ’ co m m u n ities in creased since th e total a b u n d a n c e of th e se
c o m m u n itie s w as low er th a n th e a b u n d a n c e of th e y o u n g s ta n d sp id e r
com m unity.
Soldier beetles (C antharidae)
In the pine forest s ta n d s stu d ie d in P uszcza Białow ieska soldier b eetles w ere
rep re sen te d by 16 species ( C h o b o t o w 1993), all of w hich w ere registered in
m edium age s ta n d s . The com m unity poorest in regard to species com position
(11 species), b u t a t th e sam e tim e th e m o st a b u n d a n t, w as found in y o u n g s ta n d s
(Fig. 22). 11 species w ere found to be com m on to all th e com m unities stu d ied ,
so th a t soldier beetle com m unities w ere c h a ra c te rise d by a high degree of species
com position sim ilarity. The v alu es of S o e ren se n ’s index ranged from 81% to 92%.
The com position of th e d o m in a n t species group in th e beetle co m m u n ities w as
quite sim ilar in th e s ta n d s of v ario u s ages, b u t th e young s ta n d soldier beetle
com m unity w as a little m ore u n iq u e th a n co m m u n ities of older s ta n d s (Fig. 25).
The two co -d o m in an t species from young s ta n d s, Cantharis nigricans a n d C.
pellucida fu n ctioned a s su b d o m in a n ts in older sta n d s; in th e pole wood th e
c o -d o m in a n ts w ere C. o b scu ra (a s u b d o m in a n t in y o u n g s ta n d s ) a n d
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P. T ro jan et al.
R hagonycha lignosa, w hich b ecam e a stro n g d o m in a n t in m a tu re forest. A
c h a ra c te ristic finding is th a t polytopic species w ere d o m in a n t in y oung sta n d s,
w hile o th e r form s p red o m in ated in older s ta n d s - a forest polytope a n d
coniferous forest oligotope in th e pole wood, an d a coniferous forest oligotope in
th e m a tu re forest.
C antharidae
^
Fig. 25. D om inance s tru c tu re of c o m m u n ities of c a n th a rid e s (C antharidae ) in forest s ta n d s of various
ages in Puszcza Białowieska; 1 - C antharis nigricans, 2 - C antharis pellucida, 3 - C antharis obscura,
4 - R hagonycha limbata, 5. R hagonycha lignosa, 6 - R h agonycha elongala
J u s t like in th e case of spiders, robber-flies (A silidae) ( T r o j a n u n p u b lish e d
data) a n d digger w asp s (Sphecidae) ( S k i b i ń s k a u n p u b lish e d data) w ere also
re p re sen te d by a m u ch h igher n u m b e r of species in y o ung s ta n d s (Fig. 22). Those
p red ato ry in se c ts v ere also m ore a b u n d a n t in young s ta n d s.
R obber-flies (A silidae)
A total of 22 species of A silidae w ere found in the pine forest s ta n d s stu d ied
in Puszcza Białow ieska, a s m any a s 20 of w hich w ere registered in y oung s ta n d s
com pared to 10 species c a u g h t in th e pole wood a n d 13, in m a tu re sta n d s . Most
of th e m w ere r a th e r com m on sp e cie s, w ith th e exception of Lasiopogon
im m aculatus, a species only recently found in Poland a n d c h a ra c te ristic of
Puszcza B iałow ieska. The species w as am ong the m ost a b u n d a n t species in
m a tu re forests. 9 species w ere found to be com m on to all th e s ta n d s stu d ied . A
high degree of species com position sim ilarity w as n oted for A silidae com m unities
of older s ta n d s , w ith S o e ren se n ’s index equalling 78% for pole wood a n d m a tu re
forest com m unities.
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Secondary su c c e ssio n of fa u n a in th e p ine fo rests of P uszcza B iałow ieska
41
T he a b u n d a n c e of A silidae co m m u n ities in pole wood a n d m atu re fo rests w as
sim ilar a n d nearly th re e tim es lower th a n th e a b u n d a n c e of the d ip te ra n
c o m m u n ity in young s ta n d s (Fig. 23).
From th e view point of th e com position of the m ost a b u n d a n t species group,
th e s tr u c tu r e of dom inance w as sim ilar in all th e su ccessio n sta g e s of pine forests
(Fig. 26) w ith N eoitam us cyanurus fu nctioning a s a d o m in a n t an d N. so c iu s, a s
a su b d o m in a n t. The a b u n d a n c e of each of th e two species w as m u ch lower in
m a tu re th a n in young forests. T he group of th e m ost a b u n d a n t species in all th e
c o m m u n ities stu d ied included also Tolm erus atricapillus. The co n trib u tio n of
th e se species to robber-flie co m m u n ities w as significant an d a m o u n te d to over
80% in yo u n g s ta n d s, over 90% in pole wood a n d alm ost 70% in m a tu re sta n d s .
Asilidae
Fig. 26. D om inance s tru c tu re of co m m u n ities of A silid a e in forest s ta n d s of v ario u s ages in P uszcza
B iałowieska; 1 - N eoitam us cya n u ru s , 2 - N eoitam us socius, 3 -N eo ita m u s cothurnatus, 4 - Dioctria
hyalipennis. 5 -T olm erus atricapillus, 6 - Lasiopogon im m aculatus, 7 -E pitriptus arthiticus
Digger w asp s (Sphecidae)
In th e pine forest sere of Puszcza B iałow ieska 77 species of digger w a sp s w ere
found, a c co u n tin g for V3 of the total n u m b e r of digger w asp s ever found in
Poland. T his group of p re d a to rs w as also ric h e st in yo u n g s ta n d s b o th in th e
n u m b e r of species a n d in a b u n d a n c e . 53 species were found in yo u n g sta n d s ,
39 in th e pole wood a n d 44 in m a tu re s ta n d s.
T he degree of species com position sim ilarity w as 57% for th e y oung s ta n d
a n d pole wood digger w asp co m m unities, 54% for th e y oung a n d m a tu re s ta n d
com m unities an d 70% for th e pole wood a n d m a tu re s ta n d c o m m u n ities of
Sphecidae. S tu d ie s of Sphecidae in o th e r h a b ita ts show th a t a v a lu e of
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P. T rojan et al.
S o e ren se n ’s index equalling 70% is extrem ely high for th is taxon. It m ay be th u s
c o n ten ted th a t in older s ta n d s of pine Sphecidae a re rep re sen te d by a taxocoene
of a ra th e r uniform species com position.
C onsiderable differences were, however, noted in com m unity a b u n d a n c e . The
digger w a sp s of young s ta n d s w ere nearly five tim es as a b u n d a n t a s in pole wood
a n d alm o st th re e tim es a s n u m e ro u s a s in m a tu re forests (Fig. 23).
The s tru c tu re of dom inance of pine forest digger w asp co m m u n ities in
P uszcza Białow ieska is c h a ra c te rise d by a strongly m ark ed position of th e
all-stage d o m in a n t - M ellinus a w e n s is (Fig. 27). Two species w ith a lower ra n k
- Crabro cribrarius an d Pem phredon balticus - a re in the group of 10 m ost
a b u n d a n t species th a t are p re s e n t in each age c la ss of th e forest sta n d . In total,
th e th re e species acco u n ted for a b o u t 87% of th e total a b u n d a n c e of the young
s ta n d com m unity, com pared to a b o u t 60% of th e com m unities in h ab itin g pole
wood a n d m a tu re forests.
S ph e cid ae
Fig. 27. D om inance s tru c tu re of c o m m u n ities of digger w a sp s (Sphecidae ) in forest s ta n d s of v ario u s
ages in P u szcza Białowieska; 1 - M ellinus a rv e n sis, 2 - Crabro cribrarius, 3 - P a ssa lo ecu s insignis,
4 - Crabro scutellatus, 5 - N y sso n sp in o su s, 6 - A m m ophila p u b escen s, 7 - Pem phredon balticus, 8
- P em phredon lugubris, 9 - Trypoxylon Jigulus, 10 - Trypoxylon m inus
The c h a ra c te r of th e s tru c tu re of dom in an ce of digger w asp com m unities
confirm s th e close rela tio n sh ip of co m m u n ities occurring in older s ta n d s a n d
th e m ore u n iq u e c h a ra c te r of th e yo u n g s ta n d com m unity. Am ong th e 10 m ost
a b u n d a n t species in the co m m u n ities in h a b itin g the pole wood a n d young
s ta n d s, th e re w ere 4 com m on species: M ellinus arvensis, Crabro cribrarius,
Pem phredon balticus a n d P assaloecus turionunx, a sim ilar co m parison for pole
wood a n d m a tu re sta n d co m m u n ities yielded 7 com m on species - M ellinus
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S eco n d ary su cc e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
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a r v e n s is . Crabro cribrarius, P em phredon balticus, P assaloecas insignis. P.
erem ita, R hopalum clavipes a n d E ntom ognathus brevis.
O ne m ay sp eak of trophic sp ecialisation am ong Sphecidae, a t lea st som e
species of th e taxon. The g rea test n u m b e r of species a n d th e h ig h est a b u n d a n c e
c h a ra c te riz e s th e group of digger w asps, w hose diet c o n sists p red o m in an tly of
v ario u s d ip te ran s. In the pine forests stu d ie d in Puszcza B iałow ieska th is tro p h ic
g roup w as re p re sen te d by a total 27 species. In y oung sta n d s , the a b u n d a n c e
of 22 of th e se species acco u n ted for 94% of th e total com m unity a b u n d a n c e . In
older s ta n d s , the group w as m u ch poorer b oth in the n u m b e r of species an d
a b u n d a n c e , w ith 12 species recorded in th e pole wood a n d 14 in m a tu re s ta n d s
a n d th e a b u n d a n c e being several tim es lower th a n in y oung sta n d s . The degree
of sim ilarity of th e Sphecidae com m unities w hich feed th eir offspring w ith
d ip te ra n s w as 65% for young s ta n d s a n d th e pole wood, 56% for y o u n g as
com pared to m a tu re s ta n d s an d 70% for th e pole wood an d m a tu re forest.
Sim ilar tre n d s in c h an g es in th e n u m b e r of species a n d a b u n d a n c e c a n be
observed in th e g roup of Sphecidae w hich feed th e ir offspring w ith larvae of
b u tte rflie s a n d sawflies (8 species altogether): in older s ta n d s th is g roup is n early
15 tim es less a b u n d a n t th a n in young s ta n d s.
Reverse tre n d s are observed in th e g ro u p s of digger w asp s w hich feed th eir
offspring w ith a p h id s or spiders. The ra n k of th is g roup rises co n sid erab ly in
older s ta n d s , w here only a slight in crease in th e n u m b e r of species ta k e s place,
b u t the a b u n d a n c e su rg e s su b sta n tia lly . The group feeding th eir offspring w ith
sp id er is no t n u m e ro u s (5 species) and 20 tim es m ore a b u n d a n t in m a tu re th a n
in young sta n d s .
APHIDO PHAG ES
28
26
24
CO
.2
o<x>
22
w
20
o
z
18
14
12
10
Coccinellidae
II
III
Syrphidae
IV
Neuropteroidea -a -- Sphecidae
Fig. 28. C h an g es in th e n u m b e r of species of ap h id o p h ag e s in p ine forests of th ree age c la s se s of th e
forest s ta n d
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P. T rojan et al.
The ap h id o p h a g o u s digger w asp group is rep re sen te d in th e pine s ta n d s
stu d ie d in Puszcza Białow ieska by 22 species w ith 14 species in young s ta n d s,
16 in th e pole wood an d 17 in m a tu re s ta n d s (Fig. 28). T he degree of species
com position sim ilarity (So) w as a s high a s 90% for th e pole wood an d m a tu re
sta n d s, a n d 64-66% for young a s com pared to older s ta n d s . In th e pole wood,
th e a p h id o p h a g e s w ere th re e tim es a s a b u n d a n t a s in young s ta n d s (four tim es
m ore a b u n d a n t in m a tu re s ta n d s th a n in young ones) (Fig. 29).
APH ID O PHAG ES
50
4540-
10-
III
Coccinellidae
Syrphidae
Neuropteroidea
IV
-b -
Sphecidae
Fig. 29. C h an g es in th e a b u n d a n c e of ap h id o p h ag e s in p ine fo rests of th ree age c la s se s of th e forest
sta n d
N eu ro p teran s (Neuropteroidea)
Like ap h id o p h a g o u s Sphecidae, ap h id o p h a g o u s N europteroidea also in crease
th e n u m b e r of species a n d a b u n d a n c e a s th e s ta n d grow s older (Fig. 28-29)
( C z e c h o w s k a u n p u b lish e d data). T he total n u m b e r of species recorded w as 23,
w ith 14 registered in young sta n d s , 22 in th e pole wood an d 2 0 in m a tu re s ta n d s.
A bout 58% of the N europteroidea fa u n a (13 species) w ere found in every
su ccessio n stage of th e pine forest stu d ie d in Puszcza Białow ieska. C om m unity
a b u n d a n c e in th e pole wood a n d m a tu re forest w as th re e a n d four tim es higher
th a n in y oung s ta n d s respectively (Fig. 29). The m ajority of th e species registered
w ere m ore a b u n d a n t in older th a n y o u n g er sta n d s . In the s ta n d s of v ario u s age
u n d e r investigation th e com position of th e m ost a b u n d a n t species g roup in
n e u ro p te ra n com m unities w as sim ilar (Fig. 30). Am ong th e 10 m o st a b u n d a n t
species in each age class th e re w ere five com m on species. In y o u n g s ta n d s th eir
s h a re s a m o u n te d to 70% of to tal co m m u n ity a b u n d a n ce , w hile in older s ta n d s
th e figure w as a b o u t 85%. The c h a ra c te r of th e s tru c tu re of d o m in an ce testifies
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S eco n d ary su c c e ssio n of fa u n a in th e p ine fo re sts of P uszcza B iałow ieska
45
to th e close re la tio n sh ip s betw een pole wood a n d m a tu re s ta n d Neuropteroidea
c o m m u n ities.
N e u ro p te ro id e a
9999990224
Fig. 30. D om inance s tru c tu re of c o m m u n ities of n e u ro p te ra n s (Neuropteroidea) in forest sta n d s of
v ario u s ages in P uszcza B iałow ieska; 1 - M allada p ra sin u s, 2 - W esm aelius concinnus, 3 - R aphidia
xanthostigm a, 4 - R aphidia nota ta , 5 - H em erobius stigm a, 6 - C oniopteryx parthenia, 7 - C hnjsopa
se p te m p im cta ta
S y rp h id s (Syrphidae)
C o m m u n ities of a p h id o p h a g o u s d ip te ra n s of th e family Syrphidae revealed a
reverse tendency: a s th e sta n d grew older, th ere w as a decrease in both
a b u n d a n c e a n d th e n u m b e r of species (Fig. 28-29) ( B a ń k o w s k a in press). This
p h e n o m e n o n is d u e to th e elim ination, in older sta n d s , of species associated
w ith o th e r h a b ita ts (meadows) w hich can o c c u r in considerable a b u n d a n c e in
yo u n g s ta n d s (e.g. Pipizella uaripes). O n th e o th e r h a n d , older s ta n d s see a rise
in th e a b u n d a n c e of sp ecies closely a sso c ia te d w ith pine forest (e.g. D a sy sy rp h u s
v e n u stu s, D idea interm edia). The total n u m b e r of a p h id o p h a g o u s syrphid species
registered in th e pine fo rests stu d ied in P uszcza B iałow ieska w as 41, 13 of w hich
w ere recorded in every age class of the forest s ta n d s . The species com position
of a p h id o p h a g o u s sy rp h id com m unities in h a b itin g s ta n d s of various age w as
q u ite sim ilar, b u t th e com m unity living in y o u n g s ta n d s w as m ore u n iq u e w hen
co m p ared to co m m u n ities in h ab itin g older s ta n d s . The v alues of S o e ren se n ’s
index w ere 60% for co m m u n ities from y oung s ta n d s a n d th e pole wood, a n d 65%
for co m m u n ities from pole wood a n d m a tu re sta n d s . E pisyrphus balteatus,
M eta syrp h u s corollae a n d D a sy sy rp h u s v e n u stu s w ere do m in an t in all th e
co m m u n ities stu d ie d (Fig. 31). E pisyrphus b a ltea tu s w as a little less a b u n d a n t
in older s ta n d s th a n in young sta n d s , b u t its p roportion in the com m unity
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P. T rojan et al.
in cre ase d d u e to the lower a b u n d a n c e of th e whole com m unity of aphidophago u s
S y rp h id a e in older s ta n d s. A nalogous tre n d s can be detected in M eta syrp h u s
corollae, w hich undergoes a n evident d ecrease in a b u n d a n c e (nearly 2 timers),
w hile th e th ird d o m in a n t species of S yrphidae - D a sy sy rp h u s v e n u stu s doublles
its a b u n d a n c e in older s ta n d s.
S yrphidae
Fig. 31. D om inance s tru c tu re of c o m m u n ities of sy rp h id s (Syrphidae) in forest s ta n d s of various a g es
in P uszcza B iałow ieska; 1 - E p isyrp h u s balteatus, 2 - M eta syrp h a s corollae, 3 - D a sy sy rp h u s
v e n u s tu s , 4 - M elanostom a m ellinum, 5 - Sphaerohoria m enthastri, 6 - D idea interm edia, 7 - S y rp h u s
torvus
A p h id o p h ag o u s lad y -b ird s (Coccirtellidae) a re re p re s e n te d by th e sa m e
n u m b e r of 11 species in each sta n d stu d ie d (Fig. 28). ( C z e c h o w s k a u n p u b lish e d
data). The species com position of th is in sect group in s ta n d s of v ario u s aged
s itu a te d in th e pine forest of Puszcza Białow ieska w as sim ilar. A total of 13
sp ecies w as registered including 9 com m on to each su ccessio n stage stu d ied .
S o e re n se n ’s index ranged from 83% -87% . W hile no significant differences
o ccu rred in regard to th e species com position of th e com m unities, th e re w ere
d iffe re n c e s in a b u n d a n c e (Fig. 29). As th e s ta n d s grow older, lad y b ird
c o m m u n ities d ecrease in a b u n d a n ce . In th e pole wood th e a b u n d a n c e of th e
c o m m u n ity w as twice lower a n d in m a tu re s ta n d s th ree tim es lower, th a n in
yo u n g sta n d s .
T he s tru c tu re of d om inance w as sim ilar in th e co m m u n ities a s far a s th e m o st
a b u n d a n t species group w as concerned (Fig. 32). In all th e co m m u n ities u n d e r
investigation, th e m ost n u m e ro u s species included Scym rtus suturalis a n d
M yrrha octodecim guttata, w hich collectively acco u n ted for a b o u t 87% of total
c o m m u n ity a b u n d a n c e in young s ta n d s a n d th e pole wood, com pared w ith 73%
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S econdary su c c e ssio n of fa u n a in th e p ine fo rests of P uszcza B iałow ieska
4 /
in m a tu re forests. Coccinella sep te m p u n c ta ta w as a n o th e r ra th e r a b u n d a n t
species in y o u n g sta n d s . In older s ta n d s th e species red u ced its a b u n d a n c e j u s t
like S c y m n u s suturalis.
Coccinellidae
II
mi
f
IV
Fig. 32. D om inance s tru c tu re of co m m u n ities of ap h id o p h a g o u s coccinelids (Coccinelidae) in forest
s ta n d s of v ario u s ages in P uszcza B iałowieska; 1 - S c y m n u s suturalis, 2 - M yrrha octodecim punctata,
3 - Coccinella septem puncta ta , 4 - H arm onia quadripunctata, 5 - Calvia quatuordecim punctata, 6 M ysia oblongoguttata, 1 - A n a tis ocellata
8 species of a p h id o p h a g o u s Coccinellidae develop on coniferous trees. T heir
total proportion in th e com m unity w as 91% in y o ung s ta n d s a n d a b o u t 96% in
older s ta n d s (pole wood, m a tu re stan d s). As for species closely asso ciated w ith
pine, nam ely S c y m n u s suturalis, S. nigrinus. H arm onia quadripunctata a n d
Myrrha octodecim guttata, th e ir total sh a re in th e Coccinellidae co m m u n ity w as
a b o u t 86% in y oung s ta n d s a n d th e pole wood a n d 82% in older sta n d s .
T he r e m a rk a b ly h ig h q u a lita tiv e a n d s tr u c tu r a l s im ila rity b e tw e e n
com m unities of ap h id o p h a g o u s Coccinellidae in th e canopy layer of pine s ta n d s
of various age re s u lts from th is h a b ita t being colonized by species w ith definite
h a b ita t a n d tro p h ic req u irem en ts. 80% of a co m m u n ity w as com posed of
stenotopic species, feeding only on pine a p h id s. A ttention should be p aid to th e
fa c t t h a t th e e u ry to p ic s p e c ie s (Coccinella se p te m p u n c ta ta a n d Calvia
quatuordecim punctata) were m ore a b u n d a n t in th e com m unity in h ab itin g yo u n g
s ta n d s (7%) th a n in older s ta n d s (3.2% -3.9% ).
C om m unities of p red ato ry a rth ro p o d s o ccu rrin g in pine s ta n d s of v a rio u s
ages in Puszcza Białow ieska can be classified into two groups; one com prising
p red ato rs from y o u n g s ta n d s, th e o th e r com posed of form s w hich in h a b it older
sta n d s. Young s ta n d a rth ro p o d co m m u n ities a re u su a lly m ore u n iq u e th a t th o se
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P. T rojan et al.
reg istered in pole wood an d m a tu re sta n d s . C om m unities from older s ta n d s w ere
sim ilar to one a n o th e r in species com position, dom inance s tru c tu re , uniform ity
of d istrib u tio n a n d a b u n d a n c e . T his regularity w as observed in co m m u n ities of
b o th n on-specialised p red a to rs an d of a phidophages.
The degree of sim ilarity of species com position of p re d a to rs is h ig h er for
co m m u n ities from older s ta n d s (pole wood a n d m a tu re sta n d s). The value of
S o e re n se n 's index for all th e co m m u n ities stu d ied in older s ta n d s ran g e s from
65% for a p h id o p h a g o u s S yrp h id a e to 90% for co m m u n ities of C antharidae.
N europteroidea a n d a p h i d o p h a g o u s Sphecidae. T h e d e g r e e o f s p e c i e s
com position sim ilarity betw een co m m u n ities occurring in y o u n g s ta n d s an d
older s ta n d s is lower ranging from 54% for Sphecidae co m m u n ities of y o u n g a n d
m a tu re s ta n d s to 97% for co m m u n ities of Coccinellidae from y o u n g a n d m a tu re
s ta n d s . C antharidae com m unities in h ab itin g young a n d m a tu re s ta n d s a re the
only p a ir a s sim ilar to each o th er a s co m m u n ities from older sta n d s .
T he species com position of co m m u n ities of su c h p red a to rs a s A silidae. A ranei,
Sph ecid a e a n d Syrphidae w as m u ch rich er in young s ta n d s th a n in older sta n d s .
O n th e o th e r h a n d , Neuropteroidea. C antharidae a n d a p h id o p h a g o u s Sphecidae
w ere c o m p o se d of a h ig h e r n u m b e r of s p e c ie s in o ld e r s ta n d s , w hile
a p h id o p h a g o u s Coccinellidae co m m u n ities consisted of th e sa m e n u m b e r of
species in each age class.
In th e m ajority of co m m u n ities registered in y oung sta n d s , th ere w ere higher
p ro p o rtio n s of bo th species from o th er h a b ita ts , n o t associated w ith pine forests,
a n d of eurytopic species, th a n in co m m u n ities stu d ied in older s ta n d s . In older
s ta n d s , species of open a re a s (meadows) are elim inated or th e ir a b u n d a n c e
d e c re a ses considerably, while forest polytopes a n d coniferous forest oligotopes
rem ain , often in creasin g th eir a b u n d a n c e . S u ch a re su lt of re se a rc h c o n d u cted
in P uszcza B iałow ieska h a s been confirm ed by stu d ie s co ncerned w ith c e rtain
g r o u p s o f p r e d a t o r s (A ra n e i, S y r p h id a e , Coccinellidae, N europteroidea.
Sphecidae) in pine forests situ a te d in o th e r regions of Poland (Bory T ucholskie,
P uszcza Biała).
S ignificant ch an g es w ere observed in a b u n d a n c e of individual g ro u p s of
p red a to ry a rth ro p o d s. A decreasin g p a tte rn of a b u n d a n c e is c h a ra c te ristic of
co m m u n ities of Asilidae. Cantharidae, A ranei a n d Sphecidae. S u c h a direction
of a b u n d a n c e ch an g es w as also registered for co m m u n ities of p olyphagous
d ip te ra n s of th e family Dolichopodidae, w hich are not, however, thoro u g h ly
an a ly se d in th e paper, b u t, n ev erth eless, occur n u m ero u sly in c a n o p ie s of pine
fo rests in Puszcza Białow ieska. A nother group of non-specialized p red a to rs,
nam ely sp id e r w asp s (E m pididae), w as also relatively a b u n d a n t in th e canopy
layer. The d ip te ra n s o ccurred in approxim ately th e sam e a b u n d a n c e in y oung
s ta n d s an d m a tu re s ta n d s, w hereas th ey w ere a little m ore a b u n d a n t in th e pole
wood, b u t the differences w ere n o t significant (the pole wood co m m u n ity
a b u n d a n c e w as 25% h igher th a n co m m u n ity a b u n d a n c e in y o u n g sta n d s).
Am ong aph id o p h ag es, all th e m ost a b u n d a n t tax o n s are th o ro u g h ly an a ly se d
in th e p re se n t paper. A lthough th is group h a s th e sam e tro p h ic preferences,
individual tax o n s form ing th e guild exhibited diverse p a tte rn s of c h a n g e s in the
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S eco n d ary su c c e ssio n of fa u n a in th e pine fo rests of P uszcza B iałow ieska
Asilidae
Aranei
H m ax
H m a x py
J
I
83%
III
IV
Sphecidae
IV
Coccinellidae
H m ax
J
J
45%
126%
III
IV
Cantharidae
H m ax
Fig. 3 3 . C h ang es in a c tu a l ( H ‘) a n d p o ten tial (H m a x )
s p e c ie s
d iv e r s it y
in
c o m m u n it ie s
of
n o n -s p e c ia lis e d p re d a to rs a n d th e d e g re e of
r e a l is a tio n ( J ) o f p o te n tia l d iv e r s ity o f th e
c o m m u n ities in th e sere of th e pine forest
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P. T rojan et al.
pine forest sere. The changes, b o th in resp ect to th e n u m b e r of species a n d
a b u n d a n c e , are balanced, resu ltin g in sm all total alte ra tio n s.
C o m m u n ities of p red a to rs in th e pine s ta n d s of v ario u s age investigated in
P uszcza Białow ieska displayed sim ilar stru c tu re s . In all age c lasses of th e
s ta n d s , th e g roup of d o m in a n ts h a s sim ilar species com position. The differences
c o n sist in c h an g es of p a rtic u la r sp ecies’ positions in th e s tru c tu re of dom inance
i.e. c h a n g e s of th eir ra n k s in th e com m unities.
A nalysis of sim ilarity of a b u n d a n c e d istrib u tio n in th e a c tu a l (empirical)
co m m u n ities of p red a to rs to th e five th eoretical sta tistic a l d istrib u tio n s revealed
th a t m o st of th e a c tu a l d istrib u tio n s a re related to th e negative binom ial
d istrib u tio n a n d th e logarithm ic series. T he sim ilarity is significant for v ario u s
g r o u p s in v a rio u s p in e fo re s t s u c c e s s io n s ta g e s : for ro b b e r-flie s a n d
a p h id o p h a g o u s digger w asp s in th e pole wood, for ap h id o p h ag o u s syrp h id s in
y o u n g a n d m a tu re forests. The d istrib u tio n of a b u n d a n c e in soldier beetle
c o m m u n ities is also closest to th e negative binom ial d istrib u tio n (in young a n d
m a tu re stan d s) being also th e only one to b e a r sim ilarity to th e lognorm al
d istrib u tio n (in young stan d s). The d istrib u tio n of sp id er com m unity a b u n d a n c e
is to som e e x ten t sim ilar to th e geom etric series type of d istrib u tio n (in th e pole
wood). The e stim ated n u m b e r of species for all g ro u p s of p red a to rs (except th e
y o u n g s ta n d com m unity of soldier beetles) w as hig h er th a n th e em pirical one,
w hich su g g e sts g rea ter potential cap acity of th ese h a b ita ts.
Q u a n tita tiv e s t r u c t u r e s of d ig g e r w a sp , la d y b ird a n d n e u r o p te r a n
co m m u n ities did not b e a r any sim ilarity to any of the theoretical d istrib u tio n
m odels.
S pecies diversity of n on-specialised p red a to rs (Fig. 33) d ep en d s on the g roup
of a n im a ls stu d ied . The h ig h est a c tu a l diversity w as recorded for spiders, w ith
a b sen c e of significant differences betw een th e v ario u s stag es of su ccessio n . The
m ax im u m diversity, w hich is a m e a su re of th e potential h a b ita t capacity, is also
h igh in th e case of spiders, w hile th e degree of approxim ation of th e h a b ita t
p o ten tial is th e h ig h est in th is g ro u p (J > 80%).
Sim ilar p a tte rn s of changes, b u t o ccurring on a lower level, were observed for
a c tu a l a n d potential species diversity in Asilidae. Here, a c tu a l diversity also
d e c re a se s in th e pole wood, w hilst m axim um diversity is a tta in e d in th e oldest
s ta n d s . T he degree of ap p ro x im atio n of the a c tu a l diversity to th e h a b ita t
p o ten tial in c re a se s visibly in th e consecutive stag es of th e sere (Fig. 33).
T he low est values of a c tu a l species diversity w ere noted for digger w asp s
(S p h ecidae) in y oung s ta n d s of pine. In older s ta n d s the species diversity rises
nearly two tim es. The a c tu a l species diversity of digger w asp s is m ost d istin ct
from th e po ten tial in the pine forest, p a rticu larly in young s ta n d s w here J= 26% .
A ste a d y in crease is noted for species diversity of ladybirds (Coccinellidae) in
th e pine forest sere. The tre n d is observed a s reg ard s b o th th e po ten tial an d
a c tu a l species diversity a s well a s w ith resp ect to the ap p ro x im atio n of the
p o ten tial diversity in th e h a b ita t (Fig. 33).
Inverse ten d e n c ie s a re noted only in soldier beetles (Cantharidae), w h ere the
a c tu a l species diversity in y oung s ta n d s is quite high, reach in g a m ax im u m in
th e pole wood a n d visibly d e c re a sin g th ere after (i.e. in m a tu re stan d s). This
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S eco n d ary su c c e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
51
g ro u p is also c h a ra c te rise d by a d ecrease in Pielou’s index (J) in th e consecutive
sta g e s of th e pine forest sere (Fig. 33).
S pecies diversity in ap h id o p h ag es m ay behave according to one of th re e
p a tte rn s in th e pine forest sere. In ladybirds {Coccinellidae), th ere is a n in cre ase
(Fig. 34) b o th in a c tu a l diversity a n d th e degree of approxim ation of th e h a b ita t
p o te n tia l. A rev e rse tre n d is se en in n e u ro p te ra n s (Neuropteroidea) a n d
a p h id o p h a g o u s d ig g e r w a s p s (Sphecidae). In a p h id o p h a g o u s s y r p h id s
(Syrphidae), however, bo th a c tu a l an d p o tential species diversity re a c h e s a
m ax im u m in th e pole wood stage, b u t th e degree of ap p roxim ation of th e
po ten tial diversity is th e low est here.
Coccinellidae
Syrphidae
3.5Hm ax
3-
H m ax
2.52-
1
0.5-
o-
III
Neuropteroidea
Sphecidae
H m ax
66% t:
Fig. 3 4 . C h an g es in a c tu a l ( H ') a n d p o ten tial ( H m a x ) species diversity in co m m u n ities of a p h id o p h a g e s
a n d th e degree of realisatio n (J) of po ten tial diversity of th e c o m m u n ities in th e sere of th e p ine forest
5 .2 .3 . P a rasitoid s o f p h y to p h a g es
A p a rt from p re d a to rs , p a ra s ito id s w h o se la rv a e p re s e n t p a ra s itic or
se m ip arasitic m ode of life play a n im p o rta n t role in red u cin g th e a b u n d a n c e of
p h y to p h a g e s . T h re e g ro u p s of p a ra s ito id s a re p a rtic u la rly im p o rta n t in
ecosystem s: tac h in id flies (Tachinidae), ich n eu m o n flies (Ichneum onidae) a n d
Pipunculidae. They will be described successively.
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P. T rojan et al.
PARASITOIDS
100
9080-
2
o
S.
7060-
ift
o
s
50-
|
40-
=
3020-
10IV
—<■— T a c h in id a e
P ip u n c u lid a e
Ich n e u m o n in ae
Fig. 35. C h an g es in th e n u m b e r of species of p a ra sito id s in p ine forests of th ree age cla sse s of th e
forest sta n d
T achinid flies (Tachinidae)
In th e pine fo rests of Puszcza B iałow ieska tach in id flies are rep re sen te d by
1 14 species (D r a b e r -M oń k o u n p u b lish e d data). Most of th em (82 species) a re
p a ra s ite s of lep id o p te ra n s (Lepidoptera). O th er g ro u p s given in th e o rder of
d e c re a s in g a b u n d a n c e in c lu d e p a r a s ite s of b e e tle s {Coleoptera), m o stly
p h y to p h ag o u s form s (12 species), p a ra s ite s of saw llies (S y m p h y ta ) (7 species).
G ro u p s of ta c h in id flies p a ra sitic on o rth o p te ra n s [Orthoptera), d ip te ra n s
(Diptera). h e te ro p te ra n b u g s [Heteroptera), centipedes (Chilopoda) a n d earw igs
(Dermaptera) a re less a b u n d a n t, including no m ore th a n 5 species each.
The h ig h est n u m b e r of tach in id species is recorded in young s ta n d s (Fig. 35).
T his is d u e to th e h igher proportion of flowering p la n ts th a n in older s ta n d s.
T heir n e c ta r is e sse n tia l for th e developm ent of go n ad s of tachinid flies. Young
s ta n d s also h a rb o u r th e ir ho sts: n u m e ro u s p h y to p h ag o u s species not asso ciated
w ith older s ta n d s . The lowest n u m b e r of tach in id species w as recorded for the
pole wood, w hile it w as a little h igher in m a tu re forests. T achinid a b u n d a n c e
follows a sim ilar p a tte rn w ith th e low est v alu es in the pole wood (Fig. 36). It is
in te restin g to n o te th a t th e n u m b e r of species of Tachinidae p a ra sitic on
bu tterflies, so hig h in y oung s ta n d s (65), becom es stabilised in older s ta n d s (42
species). F lu c tu a tio n s of a b u n d a n c e in th e c o u rse of th e sere are observed in
o th er h o st g ro u p s of tac h in id flies. The n u m b e r of species estim ated on th e b a sis
of th e lognorm al d istrib u tio n in d icates com plete convergence with em pirical d a ta
in each of th e th re e age c la sse s of forest s ta n d s.
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S econdary su cc e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
53
PARASITOIDS
20
16x
14-
£ 12a>
£
03
10-
-C
03
IV
—
Ta chi ni dae
P ip u n c u lid a e
Ic h n e u m o n in a e
Fig. 36. C h an g es in th e a b u n d a n c e of p a ra sito id s in p in e forests of th ree age c la sse s o f th e forest
s ta n d
Tachinidae
1
;i
Fig. 37. D om inance s tru c tu re of c o m m u n ities of ta c h in id flies (Tachinidae) in forest s ta n d s of v ario u s
ages in Puszcza Białowieska; 1 - Medina separata, 2 - Carcelia bombylans, 3 - Lypha dubia, 4 Ernestia rudis, 5 - Campylochaeta inepta. 6 - Eloceria delecta, 7 - Ceranthia abdominalis, 8 - Peribaea
fissicornis, 9 - Triarthria septipennis, 10 - Blepharomyia angustifrons
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P. T rojan et al.
The s tru c tu re of d o m in an ce of tach in id fly co m m u n ities (Fig. 37) is of th e
d o m in an t-ty p e w ith a co n sid erab le overlap betw een th e com m unity of the y o u n g
a n d m a tu re sta n d . M elina se p a ra ta . Coralia bom bylans, L yp h a dubia a n d
C am pylochaeta inepta are d o m in a n t in b oth stages. In th e pole wood, the to p
position in th e dom in an ce s tru c tu re is occupied by E rnestia ru d is, while th e
o th e r d o m in a n ts occupy th e second a n d farth er positions.
T he species diversity of tac h in id flies in th e th re e sta g e s of secondary
su c ce ssio n of the pine forest is nearly identical a n d q u ite high (Fig. 38) - a little
m ore th a n 4.5. The degree of approxim ation of p o tential species diversity of
ta c h in id flies by th e a c tu a l v alu es is also sim ilar in all th e age c la sse s of th e
forest sta n d (the h ig h est in pole wood, th e low est in young stan d s).
Tachinidae
Ichneumoninae
H m ax
H m ax
II
III
IV
I
II
III
IV
Pipunculidae
H m ax
Fig. 3 8 . C h an g es in a c tu a l ( H j a n d p o ten tial ( H m a x )
species diversity in co m m u n itie s of p a ra s ito id s an d
th e degree of realisa tio n (J) of p o ten tia l diversity of
th e c o m m u n ities in th e sere of th e p in e forest
Ich n eu m o n flies (Ichneum onidae)
In th e p re se n t p a p e r ich n e u m o n flies a re re p re sen te d only by p a ra s ite s of
lep id o p teran s [Lepidoptera), th u s in clu d in g form s biologically a sso c ia te d w ith
th e m o st d an g ero u s foliophagous p e s ts of Polish forests.
T he n u m b er of species in the pin e forest sere (Fig. 35) h a s a m ax im u m in
y o u n g s ta n d s a n d a m in im u m in m a tu re sta n d s . The c u ltu re stag e is in h ab ite d
m ostly by open a re a ich n e u m o n flies, while th e com m unity from y oung s ta n d s
h a s featu res related to th e ecotone n a tu re of th is h a b ita t. T here are n u m e ro u s
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S econdary su c ce ssio n of fa u n a in th e p ine fo rests of P u szcza B iałow ieska
55
sp ecies p re s e n t in th e c u ltu re stage a n d also typical forest species. In older
s ta n d s , open a re a s species recede to be replaced by a poorer com plex of
p a ra s ito id s of M icrolepidoptera, m a in ly Rhyaciortia buoliana, a n d sp e c ie s
p a ra sitic on th e p u p a e living in th e litter ( S a w o n i e w i c z u n p u b lish e d data).
T he a b u n d a n c e of ich n eu m o n flies p re s e n ts a sim ilar p a tte rn (Fig. 36). It is
th e low est in pine c u ltu res, rea c h e s a m axim um in young s ta n d s to becom e
sta b ilised in older s ta n d s a s a com m unity c h a ra c te ristic of th e pine forest.
T he s tru c tu re of Ichneum onidae co m m u n ities o ccu rrin g in th e consecutive
sta g e s of th e sere in Puszcza Białow ieska is sim ilar to th a t registered in
c o rre sp o n d in g forest s ta n d s in o th er regions of Poland (Fig. 39). The core of th e
co m m u n ity is form ed of 14 species w ith Vulgichneum on deceptor being the one
sp ecies to a ssu m e a m arkedly d o m in a n t position b u t only in th e c u ltu re stage.
T he com m unity of ich n e u m o n flies of th e c u ltu re stag e is quite unique. Of th e
six d o m in a n ts, th ere is only one species (C tenichenum on funereus) w hich o ccu rs
in th e y oung sta n d a s well. It is only in th e la tte r stag e th a t th re e typically forest
species (A eth eru s n itid u s. E. saw oniew iczi a n d Tycherus dodecellae) a p p e a r and
occupy top positions in th e s tru c tu re of dom inance. In older sta n d s , th e
d o m in a n t species g ro u p is su p p lem en ted by th re e o th e r species to form a stab le
core of th e com m unity.
Ichneum oninae
I
II
III
IV
Fig. 39. D om inance s tru c tu r e of co m m u n ities of ic h n e u m o n s (Ichneumonidae) in forest s ta n d s of
v ario u s ages in P uszcza Białow ieska; 1 - Platylabops virginalis, 2 - Cratichneumon sicarius, 3 Cratichneumon viator, 4 - Vulgichneumon deceptor, 5 - Virgichneumon diagrammus, 6 - Barichneumon
plcigiarius, 7 - Ichneumon minutorius, 8 - Ctenichneumon funereus, 9 - Eriplatys ardeicollis, 10 Eriplatys sawoniewiczi, 11 - Aetherus nitidus, 12 - Tycheurus dodecellae, 13 - Tycherus vagus. 14
- Tycherus verecundus
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P. T rojan et al.
Ichneum onidae species diversity in pine c u ltu re s is quite high (Fig. 38),
reach in g a m axim um in young s ta n d s w hich h a rb o u r th e g re a te st n um ber of
ich n e u m o n fly species. The diversity d ecreases in older s ta n d s to reach a
m in im u m in th e m a tu re forest. A sim ilar p a tte rn is rep o rted for potential
diversity w ith a m axim um in young s ta n d s a n d a m inium in m a tu re sta n d s. The
degree of ap p ro x im atio n of th e potential diversity by th e a c tu a l diversity
d e c re a ses in th e consecutive stag es of th e sere. The v alu es of Pielou’s index fall
from 86% in c u ltu re s to 72% in m a tu re forest. T his is suggestive of the
developm ent of in te rn a l rela tio n s w ithin the com m unity co n sistin g in an
in cre ase in th e degree of in te rn a l org an isatio n a t th e expense of decreasing
species diversity, an d th e developm ent of a s tru c tu re of d o m in an ce characterised
by a m ore d isp ersed distrib u tio n .
Pipunculidae
T his is a group of sm all-sized d ip te ra n s functioning a s specialised parasites
of leafh o p p ers (Homoptera. Auchenorrhyncha). 9 Pipunculidae species occur in
th e pine forest of Puszcza B iałow ieska ( B a ń k o w s k a u n p u b lish e d data) with the
g re a te st n u m b e r recorded in y oung s ta n d s (Fig. 35) a n d only four species
o ccu rrin g in older s ta n d s. The a b u n d a n c e of th is taxon is low est in young stan d s
a n d h ig h est in the pole wood (Fig. 36), decreasing twice in m a tu re forest. With
th e species com position growing poorer a s th e sta n d grow s older, species start
to a p p e a r th a t are restricted to older s ta n d s (Pipunculus sp in ip es a n d Chalarus
fim briatus). However, th eir a sso ciatio n w ith older s ta n d s n e e d s to be confirmed
Pipunculidae
70-i
60
504030
20
10IV
Fig. 40. D om inance s tru c tu re o f c o m m u n ities of p a ra sitic Pipunculidae in forest s ta n d s o f various
ag es in P u szcza B iałow ieska; 1 - Nephrocerus scutellatus, 2 - Eudorylas subfascipes, 3 Tomosvariella sylvatica. 4 - Cephalops semifumosus, 5 - Verrallia villosa, 6 - Cephalops aenus, 7Pipunculus spinipes, 8 - Chalarus fimbriatus
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S econdary su ccessio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
57
on th e b a s is of a m ore sizeable m aterial. T he d o m in a n t role in th e co m m u n ity
is p la y e d by N ephrocerus scu tella tu s a n d Verralia villosa, w h ic h o c c u r
th ro u g h o u t th e sere a n d m ay be considered d o m in a n t species (Fig. 40).
Species diversity te n d s to d ecrease th ro u g h o u t th e sere (Fig. 38), from a
m ax im u m in young s ta n d s to a m in im u m in m a tu re forest. T he values of Pielou
in d ex also exhibit th a t tren d . T his su g g e sts a n even a b u n d a n c e d istrib u tio n of
sp ecies in h ab itin g young s ta n d s of pine. In th e s u b s e q u e n t su ccessio n stag es,
a tra n sfo rm a tio n ta k e s place in th e c o m m u n ity of Pipunciilidae to re s u lt in a
do m in an ce-ty p e com m unity, th e n u m b e r of species d ecreases a n d differences
in th eir p ro p o rtio n s becom e sh arp er.
5 .3 . Secon d ary su c c e s s io n in th e sa p ro tro p h ic su b sy stem
The s u b s y ste m of s a p ro tro p h s is a s so c ia te d w ith dead organic m a tte r
p ro d u ced w ithin the ecosystem . D eposited a s organic se d im e n ts on th e su rfa c e
of soil, it becom es a source of energy a n d m in e rals for th e com plex su b sy ste m
of sa p ro tro p h s. Owing to th is su b sy ste m , organic m a tte r is b ro k en dow n, a n d
th e organic s u b s ta n c e s it c o n ta in s are recycled in th e ecosystem . A n o th er
im p o rta n t p ro d u ct of sa p ro tro p h activity is soil. Its su rface layers are com posed
largely of m aterials p rocessed by th e s u b sy ste m of sa p ro tro p h s. T h a n k s to them ,
th e soil h a s a definite s tru c tu re a n d type of h u m u s. The su b sy ste m is com posed
of sap ro p h ag es, om nivorous an im a ls (pantophages) an d pred ato rs.
S ta rtin g a pine c u ltu re on a pine clearing a c celerates pine forest su ccessio n .
In the first period, th e soil is considerably degraded, w hich is accom panied by
com plete c h an g es in th e living co n d itio n s of a n im a ls living there. T his is c a u se d
by th e rem oval of th e su rface cover of th e soil, stro n g insolation, in c re a se s in
te m p e r a tu r e a m p litu d e a n d c h a n g e s in m o is tu re re la tio n s . T h e ty p ic a l
c ru m b -an d -p o re s tru c tu re d isa p p e a rs a n d n u trie n ts are leached leading to
d ra s tic im p o v e rish m e n t of th e h a b ita t. As th e c u ltu re grow s, s e c o n d a ry
su c ce ssio n of p la n ts b rin g s a b o u t significant ch a n g es in soil conditions a n d in
th e rate of decom position. Insolation d e c re a ses c a u sin g ch an g es in m o istu re a n d
th e rm a l conditions, an d soil receives m ore a n d m ore n u trie n ts from falling
needles, leaves, p la n ts of th e g ro u n d cover a n d dead roots. All th e se factors
p ro m o te th e developm ent of m icroflora, m ain ly fungi so th a t fav o u ra b le
co n d itio n s are c reated for th e form ation of h u m u s a n d developm ent of soil
an im als. As the s ta n d grows, it becom es th in n e d , the u n d e rsto re y develops a n d
m icroclim ate ch an g es, also affecting soil co n d itio n s a n d th e fau n a in h a b itin g
th a t p a rt of th e ecosystem . As a re s u lt of su ccessio n , m esotrophic pine forest
develops w ith m o d e r/m o r type of h u m u s p o ssessin g fea tu re s of ed aphic clim ax.
5 .3 .1 . S o il sa p ro p h a g es
S ap ro p h ag es p a rticip a tin g in organic m a tte r decom position are re p re se n te d
by a large group of a n im a ls of diverse tro p h ism . An im p o rta n t role in soil
form ation in pine forests is played by sm all soil a n im a ls su c h a s e n c h y tra e id s
[Enchytraeidae), m it e s (Acari), s p r i n g t a i l s (Collembola) a n d n e m a t o d e s
(N em atoda). The fa u n a is classified a s soil sa p ro p h a g e s feeding on d e tritu s
to g eth er w ith b a c te ria an d fungi. It stim u la te s th e m icroflora.
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P. T rojan et al.
SAPROPHAGES
30
25-
w
o
.£
20-
Q.
co
0
15-
n3
1 io-
c
III
II
—'■— E n c h y tra e id a e
C o llem b o la
IV
N e m a to d a
Fig. 41. C h an g es in th e n u m b e r of species of soil sa p ro p h a g e s in p ine fo rests of th re e age c la sses of
th e forest s ta n d
SAPROPHAGES
25
20-
•o
c
15-
3O
£
10-
MI
—«■— E n ch ytrae id ae
C o llem b o la
IV
N e m a to d a
Fig. 42. C h an g es in th e a b u n d a n c e of soil sa p ro p h a g e s in p ine forests of th ree ag e c la sse s of th e
forest s ta n d
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S econdary su c ce ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
59
D ue to c h an g es in soil conditions observed in th e co u rse of su c ce ssio n a n d
th e s ta n d grow ing older, ch an g es occur also in th e com m unities of a n im a ls
in h a b itin g th e s ta n d (species co m p o sitio n , a b u n d a n c e a n d s tr u c tu r e of
dom inance). The an aly sis of tra n sfo rm atio n of co m m u n ities of selected soil
s a p ro p h a g e s (N em atoda . E n chytraeidae, Collembola) in pine forests of v ario u s
age allows to tra c k the p ro g ress of su ccessio n .
S tu d ie s of the effect of seco n d ary su c ce ssio n on m esofaunal co m m unities,
in clu d in g co m m u n ities of soil sa p ro p h a g e s, a re m ore frequently con cern ed open
a re a s (m eadows, field) th a n w ith fo rests ( K o e h l e r , B o r n , 1989; P e t e r s e n ,
K r o g h , 1987). In pine forests, investigations of co m m u n ities of soil m eso fa u n a
in forest p la n ta tio n s of v ario u s age have b e e n carried ou t by L o h m et al. (1977).
N em atodes (N em atoda)
N em atodes occupy a special position am ong o rg an ism s p articip atin g in th e
seco n d ary su ccessio n of pine forest. The n u m b e r of species (Fig. 41) is not
su b ject to c h a n g es d u rin g ecosystem tra n sfo rm atio n s, w ith 22 species p rese n t.
More co nsiderable c h an g es a re observed in a b u n d a n c e of nem atodes. T he initial
stage of su c ce ssio n is a sso ciated w ith a n a b u n d a n c e m inim um , while in y o u n g
s ta n d s N em atoda achieve a m axim um of a b u n d a n c e , exceeding th e m in im u m
n early 3 tim es. In older s ta n d s th e a b u n d a n c e becom es stable. However, in
co m p ariso n to a b u n d a n c e ch an g es in sp rin g tails a n d E nchytraeidae (Fig. 42),
flu c tu a tio n s in the a b u n d a n c e of n em a to d es m ay be considered insignificant.
The p a tte rn of c h an g es in a b u n d a n c e is related to a d ecrease in inso latio n a n d
N em atoda
Fig. 43. D om inance s tru c tu re o f c o m m u n ities of n e m ato d e s (Nematoda) in forest s ta n d s of v a rio u s
ages in P uszcza B iałow ieska; 1 - Filenchus ditissimus, 2 - Acrobeloides nanus, 3 - Aphelenchoides
spp. , 4 - Plectus pusteri, 5 - Wilsonema otophorum, 6 - Teratocephalus dadayi, 7 - Ypsylonellus
insubdcus, 8 - Eudonjlaimus silvaticus, 9 - Rhabditis s. I
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P. T rojan et al.
60
d e n se r forest floor cover asso ciated with su ccessio n , w hich re s u lts in sm aller
v a ria tio n s in soil m oisture.
The s tru c tu re of d om inance in soil N em atoda co m m u n ities is sim ilar in th e
four su c c e ssio n sta g e s (Fig. 43). T he core of the com m unity does not change,
th ere are sm all c h an g es in th e d o m in a n t species group. G reater tra n sfo rm a tio n s
a re no ted in th e s tru c tu re of less n u m e ro u s species. As th e s ta n d grows older,
t h e c o n t r i b u t i o n of f u n g iv o r o u s n e m a to d e s i n c r e a s e s w h ile t h a t of
b a c te rio p h a g o u s N em atoda decreases, w hich is a sso ciated w ith a n increase of
th e s h a re of fungi in th e forest floor m icroflora ( B r z e s k i u n p u b lis h e d data).
T he species diversity of n e m a to d es u n d erg o es sim ilar stab ilisatio n , a s th e
p a ra m e te rs d isc u sse d above (Fig. 44). The value of th e diversity index(H ’)
flu c tu a te s a ro u n d 3.0. A dditionally, p o tential species diversity Hmax in th e
consecutive stag es n e a rs 4.5 so th a t th e degree of ap p roxim ation of p o tential
diversity does not change considerably.
Collembola
Enchytraeidae
4 \H m ax
r'
H m ax
J
66%
Nematoda
H m ax
J
64%
Fig. 44. C h an g es in a c tu a l (H’) an d p o ten tial (Hmax)
s p e c i e s d iv e r s it y in c o m m u n it ie s o f s o il
sa p ro p h a g e s a n d th e degree of rea lisa tio n (J) of
p o ten tial diversity of th e c o m m u n ities in th e sere
of th e pine forest
E n c h y tra e id s (E nchytraeidae)
E n ch y traeid co m m u n ities in th e soil of pine forests of P uszcza Białow ieska
are poor in th e n u m b e r of species, w ith one d o m in an t species, nam ely C ognettia
sphagnetorum . In th e initial stage of su c ce ssio n (culture), w here the soil is m o st
degraded, only one species w as recorded. The g rea test n u m b e r of species w as
registered in th e young sta n d . In older s ta n d s th eir n u m b e r d e c re a ses (Fig. 41),
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S econdary su ccessio n of fa u n a in th e pine fo rests of Puszcza B iałow ieska
61
a p p ro a c h in g th e n u m b e r c h a ra c te ris tic of m a tu re forests in o th e r forest
com plexes in Poland ( P i li p iu k 1993). C om m unity a b u n d a n c e in c u ltu re s an d
y o u n g s ta n d s is lower th a n in the pole wood stage an d m a tu re forests, w here it
re a c h e s a m axim um (Fig. 42).
T he s tru c tu re of the enchytraeid com m unity (Fig. 45) is c h a ra c te rise d by a
m ark e d dom inance of one species in every stage of succession. Cognettia
spha g n eto ru m a c c o u n ts for m ore th a n 90% of th e com m unity th ro u g h o u t th e
sere, rea c h in g a m axim um in the pole wood to u n dergo a slight fall in th e m a tu re
s ta n d . If we ignore the restrictin g c h a ra c te r of th e c u ltu re stage w here only one
e n c h y trae id species is p rese n t, the young s ta n d is capable of accom m odating
seven o th e r accessory species. In th e s u b s e q u e n t stag es the s tru c tu re of th e
co m m u n ity becom es poorer due to elim ination of accessory species ( P i l i p i u k
u n p u b lis h e d data).
E nchytraeidae
100
90
80
70
60
X/A 1
50
□ 2
40
ip
30
20 H
10
0
IV
Fig. 45. D om inance s tru c tu re of co m m u n ities of Enchytraeidae in forest s ta n d s of v a rio u s ages in
P uszcza B iałow ieska: 1 - Cognettia sphagnetorum, 2 - Achaeta sp. , 3 - Mesenchytraeus sp. , 4 Marionina sp. , 5 - Buchholzia simplex
The species diversity of en ch y traeid s is profoundly influenced by s tru c tu ra l
rela tio n s (Fig. 44). A ctual diversity (H’), equalling 0 in th e first sta g e of
su ccessio n , never exceeds 0.5 in the su b s e q u e n t stages. G reater c h a n g es o ccu r
w ith resp e c t to potential diversity (Hmax). It rea c h e s a peak in th e y oung s ta n d
to fall considerably in older s ta n d s. The u tilisatio n of th e p o tential h a b ita t
capacity to in crease th e diversity of th e com m un ity of e n ch y traeid s is low, never
reach in g 20% in the entire sere.
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62
P. T rojan et al.
S p ringtails (Collembola)
The n u m b e r of species of th e se a n im a ls in the pine forest sere te n d s to
in crease (Fig. 41). It re a c h e s a m axim um in th e pole wood to fall slightly in the
m a tu re sta n d . A sim ilar p a tte rn is observed in a b u n d a n c e of sp rin g tails (Fig. 42),
w hich is m u ch lower th a n th a t of enchytraeids, while being slightly hig h er th a n
th e a b u n d a n c e of nem atodes.
T he species com position of Collembola differs considerably in th e consecutive
sta g e s of the pine forest sere in Puszcza B iałow ieska (Fig. 46). C onvergence of
species com position is only observed w hen com paring co m m u n ities in h ab itin g
older s ta n d s (pole wood an d m a tu re forest). T here are also differences in the
s tru c tu re of dom inance of the springtail com m unities. In yo u n g er stages, th ere
is a decrease in th e n u m b e r of n o n -a b u n d a n t species com bined w ith a m arked
d o m in an ce of one species (80%). In older age classes (pole wood, m a tu re forest)
th e core of the com m unity is form ed of a n u m b e r of co -d o m in an t species an d
th e low -share species g roup is m u ch m ore n u m ero u s. In th e consecutive stages
of th e pine forest sere, species form ing th e d o m in an t g roup a re replaced by o th ers
( S t e r z y ń s k a u n p u b lish e d data).
Species diversity of sp rin g tails (Fig. 44) is on th e sam e level a s in nem ato d es.
However, a m axim um is reach ed in the young s ta n d stage, while in older s ta n d s
a c tu a l species diversity (H') oscillates a ro u n d 2.8. P otential diversity re a c h e s a
p e a k in th e two age classes. T he hig h est degree of approxim ation of th e potential
C ollem bola
69
Fig. 46. D om inance s tru c tu re of co m m u n ities of sp rin g tails (Collembola) in forest s ta n d s of v ario u s
ages in Puszcza Białowieska; 1 - Pseudosinella zygophora, 2 - Isotomiella minor, 3 - Mesaphorura
macrochaeta, 4 - Isotoma notabilis, 5 - Schoetella iinunguiculata, 6 - Onychiurus subuliginatus, 7 Isotoma sp.j. . 8 - Folsomia quad.riocula.ta, 9 - Anurophorus septentrionalis, 10 - Entomobryidae sp.
j . , 11 - Willemia aspinata, 12 - Pogonognathellus flavescens, 13 - Anurida pygmaea, 14 - Proisotoma
minima
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S econdary su ccessio n of fa u n a in th e pine fo rests of P uszcza B iałow ieska
63
is seen in y o u n g s ta n d s, while th e difference is th e g re a te st in m atu re forests.
T he form ation of com m unity s tru c tu re in Collembola is accom plished a t th e
expense of species diversity.
Two m odels of do m inance s tru c tu re occur in th e sere studied. The first of
th em h a s th e sam e species of th e d o m in a n t g roup form ing the core of th e
c o m m u n ity th ro u g h o u t th e sere. In th is m odel ch an g es are seen the group of
species w ith lower sh a re s in the com m unity. T his m odel is conform ed to by two
g ro u p s - N em atoda an d E nchytraeidae (Fig. 43 an d 45). The o th er m odel of
d o m in an ce s tru c tu re is observed in Collembola (Fig. 46). In th is taxon th ere are
co n sid erab le tra n sfo rm atio n s in th e d o m in a n t species group w ith different
species prevailing in young s ta n d s a n d older ones, a lth o u g h one m u s t n o t sp eak
a b o u t com plete su b stitu tio n of species com position. W hat is m ore, Collembola
c o m m u n ities from older s ta n d s have several co-dom inants, while only one
m ark ed d o m in a n t a p p e ars in y ounger s ta n d s. A nalysis of sim ilarity of th e a c tu a l
co m m u n ities of soil sap ro p h ag es to th e five theoretical sta tistic a l d istrib u tio n s
show ed th a t th e d istrib u tio n of species in th e com m unities u n d e r stu d y is closest
to th e negative binom ial distrib u tio n . T he sim ilarity is statistically significant
for v ario u s g ro u p s in v arious su ccessio n stages. In som e ca se s th e estim ated
n u m b e r of species is higher th a n th e a c tu a l one (Collembola) indicating a h ig h er
species cap acity of th e h a b ita t. In o th ers (E nchytraeidae), th e e stim atio n
c o rre sp o n d s to em pirical findings.
A nalysis of selected groups of soil sa p ro p h a g e s show ed th a t the co m m u n ities
of th e g ro u p s u n d e r stu d y becom e stab ilised in older s ta n d s of pine. T he
co m m u n ities of th ese an im als in h ab itin g m iddle age class s ta n d s p re se n t a
“p rep a ra to ry stag e” for the form ation of com m unities c h a ra c te ristic of n a tu ra l
m a tu re pine forests.
In the sere u n d e r investigation, th e pole wood is a stage of g reater potential
for realizing species diversity in relation to co m m u n ities from m a tu re sta n d s . In
m a tu re s ta n d s , w here h a b ita t conditions becom e stabilized to a c ertain extent,
species diversity of soil sa p ro p h a g e s is lower. G i l l e r (1984) claim s a link betw een
th e degree of species diversity an d h a b ita t heterogeneity. In th e sere stu d ied , a n
in crease in h a b ita t heterogeneity is stim u la te d by th e developm ent of th e layer
s tru c tu re of the s ta n d a n d c h an g es in soil conditions. S tratification, b o th
horizontal a n d vertical, provides m an y new niches. P articularly im p o rta n t for
sa p ro p h a g e s are litter an d h u m u s. T he relatio n s betw een soil sa p ro p h a g e
co m m u n ities an d th e type of h u m u s form ed are described by P o n g e (1993).
Species diversity m ay also rise w ith in cre asin g p ro d u ctio n of food su c h a s fungi
an d m icroorganism s a n d organic m a tte r c o n te n t in the soil ( P i a n k a 1966). In
m a tu re clim ax ecosystem s, productivity often decreases or rem ain s c o n sta n t.
Species diversity also m ay fall in su c h ecosystem s d u e to th e rising n u m b e r of
in terd ep en d en cies betw een o rg an ism s even th o u g h it is still stim u lated by th e
growing n u m b e r of niches. The m a tu re pine forest is a m a tu re ecosystem w here
h a b ita t co n d itio n s becom e stabilized. T he decrease in th e species diversity of
soil sa p ro p h a g e s, w hich can be observed in th e m a tu re forest is connected above
all else to th e stabilization of the ra te of h u m u s fixation a n d of h u m u s type.
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P. T ro jan et al.
5 .3 .2 . P a n to p h a g es and so il predators
An im p o rta n t role in reg u latin g th e a b u n d a n c e of th e sa p ro p h a g e su b sy ste m
is played by om nivorous a n d p red a to ry an im als. The form er a re rep re sen te d in
th e pine fo rests by a n ts , w hich control th e w hole ecosystem feeding on a wide
ran g e of anim al a n d p la n t foods. Two fam ilies of d ip te ra n s (Rhagionidae a n d
Therevidae) serve a s exam ples of soil p red a to rs. The lim bless larvae of th e se
d ip te ra n s p e n e tra te into the soil h a b ita t h u n tin g sm all in v erteb rates.
Soil p a n to p h a g e s
A nts (Formicidae)
A nts play a p a rtic u la r role am o n g th e m an y g ro u p s of p a n to p h a g o u s soil
in v erteb rates. D ue to th eir high a b u n d a n c e a n d exceptional biological activity,
th ey a re a fu n ctio n al g ro u p vital to m a in ta in in g h o m o e o sta sis in fo rest
ecosystem s, p a rticu larly in coniferous forests ( K a r p i ń s k i 1956, K a c z m a r e k
1963). More or less closely a sso ciated w ith th is h a b ita t are a b o u t 30 species of
a n ts of th e nearly 80 species form ing th e Polish a n t fa u n a (PISARSKI 1975,
C z e c h o w s k i u n p u b lis h e d d a ta ). T h e ir role in soil fo rm a tio n c o n s is ts in
su p p lem en tin g the soil w ith organic co m p o u n d s an d sh a p in g th e com position
of the m icroflora ( P ę t a ł 1977). It is also closely linked w ith th e ch a n g es o ccu rrin g
d u rin g su ccession.
PANTO PHAG ES
o
E
IV
—«■— F o rm icid ae
Fig. 47. C h an g es in th e n u m b e r of sp ecies of a n ts in th e seco n d ary sere of p in e forests
Only 19 species of a n ts a re a sso c ia te d w ith th e pine forest in Puszcza
Białow ieska. 12 species in h a b it th e c u ltu re stag e (Fig. 47). In th e y o u n g sta n d
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S eco n d ary su ccessio n of fa u n a in th e p ine forests of P uszcza B iałow ieska
65
a n d pole wood th eir n u m b e r decreases slightly to rea c h a m axim um in th e
m a tu re forest. D uring th e secondary su ccessio n of th e pine forests of Puszcza
B iałow ieska s u b s ta n tia l ch an g es take place in th e com position an d s tru c tu re of
th e a n t co m m u n ity ( C e c h o w s k i u n p u b lish e d data). The su ccessio n in th e
Formicidae taxocoene co n sists of two p h a ses. The first of them is observed in
y o u n g s ta n d s an d the pole wood. The two com m unities b e a r a 70% species
com position sim ilarity (according to S o e ren se n ’s index) a n d identical s tru c tu re s
of d o m in an ce (M orisita's index = 1). The m ost a b u n d a n t species in th ese
c o m m u n itie s a re p o ly to p ic s p e c ie s of d ry h a b it a ts (Fig. 48), a n d th e
x e ro th e rm o p h ilo u s Tetram orium c a e sp itu m w a s a d o m i n a n t in b o th
c o m m u n ities (64% on the average), while L asius niger, a ub iq u ito u s, typically
pioneering species, w as a s u b d o m in a n t (25%). C om m unities from the m a tu rin g
an d m a tu re s ta n d s form ed th e second p h a se of su ccessio n w ith a sim ilar degree
of species com position sim ilarity (70%) a n d sim ilar s tru c tu re s of dom inance
(0.95). T he m o st a b u n d a n t species in th a t p h a se included polytopes of wet
h a b ita ts w ith th e do m in an t M yrmica ruginodis (59%). In te rm s of th e n u m b e r of
a n t n e sts, th e second position w as occupied by th e forest oligotope Leptothorax
acervorw n (24%). Formicidae com m unities of the two d istin ct su ccessio n p h a se s
differed n o t so m u ch in species com position (average sim ilarity 66%) a s in th e
s tru c tu re of do m in an ce (average sim ilarity a s low a s 0.07).
Form icidae
I
II
III
IV
Fig. 48. D om inance s tru c tu re of co m m u n ities of a n ts (Formicidae) in forest sta n d s of v a rio u s ages
in Puszcza Białow ieska; 1 - Myrmica ruginodis, 2 - Leptothorax acervorum. 3 - Lasius niger. 4 - Lasius
umhratus, 5 - Leptothorax muscorum, 6 - Lasius flavus, 7 - Myrmica lobicornis, 8 - Tetramorium
caespitum, 9 - Myrmica schencki, 10 - FormicaJ'usca
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P. T ro jan et al.
C h a n g e s in total d ensity of a n t n e s ts w ere observed in th e Formicidae
co m m u n ities of the pine forest sere (Fig. 49). T his p a ra m e te r is subject to
s u b s ta n tia l variatio n s in consecutive sta g e s to finally rea c h a value (in m a tu re
forests) sim ilar to th a t recorded for th e earliest stage stu d ied (pine culture).
A ctual species diversity (Hj m e a s u re d w ith S hannon-W eaver’s index exhibits
a gentle b u t steady in crease w ith th e grow ing age of th e forest sta n d : from 1.52
to 1.77 (Fig. 50). Potential diversity (Hmax) is also th e h ig h est in th e m a tu re s ta n d
stage (3.91) b u t the tra n sitio n a l sta g e s a re c h a ra c te rise d by slightly lower values
(3.32-3.58). The degree of ap p ro x im atio n of th e p o tential does n o t exhibit
c o n sid erab le ch an g es e ith e r (Fig. 50), being, however, th e low est in the initial
sta g e s of su c ce ssio n (42% a n d 45% in th e c u ltu re a n d y oung s ta n d stage
respectively) to grow slightly in th e older sta g e s (53% a n d 74% in th e m atu rin g
s ta n d a n d m a tu re forest respectively).
PANTO PHAG ES
7500
70006500® 6000a> 5500o
c
-§
5000-
■§ 4500400035003000
IV
—1■— F o rm icid ae
Fig. 49. C h an g es in th e a b u n d a n c e of a n t n e s ts in relatio n to th e age o f pine forest s ta n d s
T he specific fea tu re s of th e se co n d a ry su ccessio n tak in g place in th e plots of
pine forest d istrib u ted in a relatively “p a tc h y ” m a n n e r a n d going th ro u g h v arious
sta g e s of m atu rity , include th e p re se n c e of the forest a n t species of th e su b g e n u s
Formica s. str. in every su c ce ssio n stage, especially in th e c u ltu re s, th e species
itself being only typical of th e clim ax sta g e s in forests. They re p re s e n t th e K-type
ecological strateg y i.e. co n q u er th e ir en v iro n m en t slowly b u t rem a in th ere forever
( P i s a r s k i , C z e c h o w s k i 1994). U nder th e c irc u m sta n c e s of th e large forest
com plex of Puszcza B iałow ieska, large colonies of th ese a n ts a re able to survive
forest felling a n d silvicultural p ra c tic e s leading to th e e s ta b lis h m e n t of a
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Secondary su c ce ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
67
p la n ta tio n , a n d to m a in ta in th eir n e s ts by feeding in a d ja c e n t old s ta n d s . A
s p e c ta c u la r exam ple w as th e com plete a b sen c e of an th ills of Formica p o lyctena
in one of the m a tu re forest a re a s stu d ied , alth o u g h th e species h ad a hig h s h a re
in th e m aterial obtained u sin g B arb er’s pitfall tra p s ( C z e c h o w s k i u n p u b lis h e d
data).
Form icidae
3.5-
H m ax
2.5-
1.553%
42%
45%
0.5-
III
IV
Fig. 50. C hanges in a c tu a l (H') a n d po ten tial (Hmax) sp ecies diversity in co m m u n ities of a n ts a n d th e
d egree of realisation (J) of p otential diversity of th e c o m m u n ities in th e sere of th e p ine forest
Soil p re d a to rs
O ut of a n u m b e r of g ro u p s of d ip te ra n s w hose larvae develop in th e soil leading
a p redatory m ode of life, the fam ilies Therevidae a n d R hagionidae are a n aly sed
in o u r study. The d a ta on th eir o ccu rren ce in Poland in to -d ate lite ra tu re a re
quite scarce ( T r o j a n 1970, M i k o ł a j c z y k 1991), n e ith e r h a s the s tru c tu re of
com m unities b een stu d ied .
Therevidae
A total of 14 T herevidae species w ere found in pine forest s ta n d s in P uszcza
B iałow ieska b a s e d on m aterial collected w ith M oericke’s pitfall tra p s i.e.
im agines (Fig. 51). The a b u n d a n c e of Therevidae a n d th e n u m b e r of sp ecies in
th re e different age c la sse s of th e pine forest: y oung s ta n d , pole wood a n d m a tu re
sta n d , do no t vary significantly (Fig. 52). T he co m m u n ities are quite sim ilar w ith
regard to th eir species com position (the v a lu e s of S o e ren se n ’s index a re 78%,
82% a n d 76%). Thereva handlirschiis the d o m in a n t species a t all sta g e s (Fig. 53).
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P. T rojan et al.
SO IL PREDATORS
10 -
o
o
E
III
—<■— T h e re v id a e
IV
R ha g io n id a e
Fig. 51. C h an g es in th e n u m b e r of sp ecies of soil p re d a to rs in th e seco n d ary sere of p in e forests
SO IL PREDATORS
1
IV
—■ — T h erevid ae
••**■••• R h agio nidae
Fig. 52. C h an g es in th e a b u n d a n c e o f soil p re d a to rs in relation to th e age of p in e fo rest s ta n d s
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S econdary su cc e ssio n of fa u n a in th e p ine forests of P u szcza B iałow ieska
69
T h e sp ecies o c c u rs com m only th ro u g h o u t th e co u n try . The sim ilarity of
co m m u n ity s tru c tu re s is also high in Therevidae, achieving v alu es from 0.80 to
0 .9 7 according to M orisita’s coefficient. The only difference we have observed
betw een co m m u n ities of Therevidae in different age c lasses of th e pine forest is
th a t in th e y o ung s ta n d th e d istrib u tio n of a b u n d a n c e per species parallels, in
all likelihood, th re e d istrib u tio n s a t the sam e tim e, nam ely th e geom etric series,
th e logarithm ic series an d th e binom ial negative d istrib u tio n . The degree of
sim ilarity to th ese d istrib u tio n s d rops considerably in co m m u n ities o ccurring
in th e pole wood, w hile Therevidae a b u n d a n c e d istrib u tio n in the m a tu re forest
does not b e a r any sim ilarity to any of th e theoretical d istrib u tio n s considered.
The sim ilarity to th e geom etric series, w hich indicates, a s th e th eo retical
principle h a s it, th a t th e a b u n d a n c e of each species is p roportional to th e h a b ita t
reso u rc es it utilises, su g g e sts th a t in th e case of Therevidae su c h a s itu a tio n
m ay actually tak e place in y ounger age c lasses of pine forest. As th e s ta n d grows
older, th e s tru c tu re of the com m unity probably becom es less regular, w hich
seem s easy to u n d e rs ta n d considering th e m ethod of sam ple collection. In th e
consecutive stag es of the pine forest sere, M oericke’s pitfall tra p s w ere u se d to
collect in se c ts flying a t increasingly high altitu d es, possibly resu ltin g in a m ore
ra n d o m selection of th e Therevidae fau n a , w hich m ig h t be m ore closely
asso ciated w ith the g ro u n d cover w here it is often registered. The slight fall in
th e a c tu a l a n d potential diversity indices observed in th e consecutive stag es of
th e pine forest sere ste m s from slight v ariatio n s in th e n u m b e r of species. The
Therevidae
Fig. 53. D om inance s tru c tu r e of co m m u n ities of Therevidae in forest s ta n d s of v ario u s ag es in Puszcza
Białowieska; 1 - Thereva handlirschi. 2 - Thereva nobilitata. 3 - Thereva lanata. 4 - Thereva plebeja.
5 - Thereva circumscripta, 6 - Cliorismia ardea
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P. T rojan et al.
Therevidae
3.5-
H m ax
2.5-
1.5-
85%
83%
§§I80%
0.5-
IV
Fig. 54. C h an g es in ac tu a l (Hj a n d p o ten tial (Hmax) species diversity in c o m m u n ities of Therevidae
a n d th e degree of realisation (J) of poten tial diversity of th e co m m u n ities in th e sere of th e p ine forest
c o n s ta n t n a tu re of th is co m m u n ity p a ra m e te r is confirm ed by th e degree of
ap p ro x im atio n of potential species diversity, w hich rem a in s a t a m ore or less the
sam e high level ranging from 8 0.4 to 8 4.7 (Fig. 54).
R hagionidae
C o m m u n ities of R hagionidae, d ip te ra n s exhibiting, in the larval a n d imago
stages, a p red ato ry m ode of life, sim ilar to Therevidae, w ere also stu d ie d in three
sta g e s of pine forest in Puszcza Białow ieska. T hese co m m u n ities exhibited
sim ilarly high co n stan cy of th e species com position a n d s tru c tu re . They are
com posed of only th ree species, th e com m on Rhagio lineola being th e d o m in a n t
w ith a t least a 95% sh a re in each of th e co m m u n ities u n d e r stu d y .
5.4.
Vertebrates
V erteb rates are considerably diversified in species com position, size a n d the
role th a t individual species play in n a tu re . In ecological term s, they are regarded
a s belonging to two su b sy ste m s. P hytophages, w hich include hoofed m am m als
a n d m o st ro d en ts, are b io tro p h s. The o th e r su b sy stem , m ac ro tro p h s, em b races
p re d a to rs co n tro llin g th e a b u n d a n c e of th e b io tro p h ic a n d sa p ro tro p h ic
su b sy ste m s. Special m eth o d s a re em ployed for a sse ssin g species com position
a n d a b u n d a n c e of verteb rates, dep en d in g on the size a n d m ode of life so th a t
v e rte b ra te s are seldom m entioned in com prehensive fau n al p a p e rs, u su a lly
dealing only w ith invertebrates. T his a n a ly sis is b ased on a p a p e r by W o l k and
G u t o w s k i (1984) devoted to sm all m am m als a n d b ird s of pine fo rests of Puszcza
B ia ło w ie sk a a n d on s tu d ie s of s m a ll m a m m a ls of p in e fo re s ts ( P u c e k
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S eco n d ary su c c e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
71
u n p u b lish e d data). N either of th ese p a p e rs in clu d e s large m am m als, w h e th e r
hoofed or predatory, or b ird s of prey. T hese groups, however, do n o t c o n ta in
g ro u p s c h a ra c te ristic of pine forests.
M am m als (M am m alia)
S m a ll m a m m a ls (Micromammalia) e n c o m p a s s tw o g ro u p s p e rfo rm in g
different biological roles: p hytophages, w hich com prise ro d en ts, a n d insectivores
(som e species m ay be considered polyphagous). 15 species of sm all m am m a ls
w ere found in th e pine forests of Puszcza Białow ieska. The n u m b e r of species in
individual sta g e s of seco n d ary su ccessio n , th e n u m b e r of species revolves
a ro u n d 10 (Fig. 55). A noticeable d ecrease in th e n u m b e r of species - dow n to 7
( P u c e k u n p u b lish e d data) or 4 ( W o l k a n d G u t o w s k i 1984) - is seen in th e m o st
m a tu re sta n d s . T hese differences tu rn o u t to be ra th e r insignificant co n sid erin g
th e period w hen th e m ate ria ls w ere collected an d th e species re p re se n te d by
single ind iv id u als collected d u rin g th e five y e a rs of catc h e s. T hese species (see
inventory) a re regarded a s fau n al rarities in th e a re a of Puszcza B iałow ieska.
M ICRO M AM M ALIA
10-
o
a>
.e
E
3
9
20
30
120
1 5 0 years
age of fo re st
Fig. 55. C h anges in th e n u m b e r of species of sm all m am m als in relatio n to th e age of p in e forest
sta n d s
The p a tte rn of c h a n g es in a b u n d a n c e is c h a ra c te rise d by a high d e n sity of
sm all m am m als in pine c u ltu re s (Fig. 56). T here is a 30% fall in young s ta n d s
a n d th e pole wood, a n d in m a tu re forest th e a b u n d a n c e of M icromam malia
re tu rn s to th e original values.
The s tru c tu re of th e sm all m am m al co m m u n ity show s a stab le g roup of
d o m in an ts th ro u g h o u t th e secondary sere (Fig. 57). It is form ed of four species.
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72
P. T rojan et al.
MICRO M AM M ALIA
300
280-
g
260-
c
o
ć
240-
«
220 -
200-
180
9
20
30
age of forest
120
150 years
Fig. 56. C h an g es in the a b u n d a n c e of sm all m am m als in relation to th e age of pin e forest s ta n d s
th e ro d en ts: b a n k vole (Clethrionom ys glareolus) an d field yellow -necked m ouse
(A podem us Jlavicollis) a n d th e insectivore shrew s: Sorex a raneus a n d Sorex
m inutiis. It is in terestin g to note th a t th e b a n k vole in creases its sh a re in the
co u rse of th e sere, a c co u n tin g for a s m any a s 67% of the total n u m b e r of sm all
m am m a ls cau g h t, while the peak a b u n d a n c e of yellow -necked field m ouse is
registered in th e pole wood. Reverse tre n d s a re observed in the zoohagous group.
S. araneiis is m ost a b u n d a n t in pine c u ltu res, his a b u n d a n c e d ecreasin g w ith
th e grow ing age of th e s ta n d . S. m inutiis exhibits larger v ariatio n s in a b u n d a n c e
th a t a re n o t associated w ith the age of th e forest sta n d . T here is a considerable
ag e-related d ecrease in th e sh a re of m eadow a n d field species. C om m on vole
(Microtus arvalis), quite com m on in th e c u ltu re stage, d isa p p e a rs in th e pole
wood no t to em erge again in m a tu re forests. S h ort-tailed vole (Microtus agrestis)
h a s a sh a re of several percen t in co m m u n ities in each age class, b u t it w as not
registered in th e pole wood. H arvest m o u se (Micromys minutus) a n d n o rth e rn
b irch m o u se (Sicista betulina) are ch a ra c te rise d by sim ilar sh a re s. O th er species
o ccu r in th e pine forest only sporadically.
S pecies diversity of sm all m am m als is h ig h est in c u ltu re s an d y o ung s ta n d s
of pine (Fig. 58) d ecreasing in older s ta n d s to reach a m inim um in th e final link
of th e seco n d ary sere. In th is case also, th e in crease of com m unity o rg an isatio n
is accom panied by the elim ination of accessory species a n d th e stre n g th e n in g
of th e d o m in a n t position of species form ing the core of th e com m unity. T his is
obviously reflected in th e value of Pielou’s index (J), reflecting the degree of
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S econdary su cc e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
73
M ICRO M AM M ALIA
141^362018
H 3
□ 4
I*
□ 6
30
1 1 0 -1 2 0 1 5 0 years
Fig. 57. D om inance s tru c tu re of co m m u n ities of sm all m a m m als in forest s ta n d s of v a rio u s ages in
P u szcza B iałowieska; 1 - Clethrionomys glareolus, 2 - Sorex araneus, 3 - Apodemus JlavicoUis. 4 Sorex minutus, 5 - Microtus agrestis, 6 - Sicista betulina, 7 - Apodemus agrarius, 8 - Micromys
minutus, 9 - Microtus arvalis
M ICRO M AM M ALIA
3.5
H m ax
2 .5 -
1.5-
77% :
H
71%
64%
58%
0.5 -
9
20
30
120
1 5 0 years
aqe of fo re s t
Fig. 58. C h an g es in a c tu a l (FI') a n d p otential (Hmax) species diversity in c o m m u n ities of sm all
m am m als a n d th e degree of realisa tio n (J) of p o ten tial diversity of th e co m m u n ities in th e sere of th e
pine forest
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P. T rojan et al.
74
realisatio n of th e potential for species diversity. The value of th is index falls
steadily th ro u g h o u t th e sere.
B irds (Aues)
N esting b ird s are re p re se n te d in th e pine forests of Puszcza Białow ieska by
23 species. None of th em o ccu rs a t every stage of th e seco n d ary su ccessio n
( W o ł k , G u t o w s k i 1984). D uring th e first y ears after a p la n ta tio n of pine is
sta rte d , th e afforested clearings are settled by th re e field species. T heir n u m b e r
in cre ase s u p to eight species in a 9 - 17-year-old s ta n d (Fig. 59), u p to 11 after
22 years, to rea c h 16 in th e oldest s ta n d s . In the la tte r stage, only typically forest
species occur. T his set of species, however, is only c h a ra c te ristic of larger a n d
older pine forest com plexes. In enclaves of m a tu re forest s u rro u n d e d by younger
s ta n d s, th e n u m b e r of species of n e stin g b ird s falls by alm o st a V3 i.e. dow n to
9. D istrib u tio n -b a se d estim atio n of th e n u m b e r of species w as only possible for
2 2 -2 4 -y e a r-o ld s ta n d s. A value of S =11.07 in d icates th a t em pirical d a ta
co rrespond to theoretical estim ates.
AVES
1
1
o
©
E
3-5
9-17
22-24
> 1 0 0 years
ag e of fo rest
Fig. 59. C h an g es in th e n u m b e r of sp ecies of b ird s in relation to th e age of p ine fo rest s ta n d s
The n u m b e r of couples of n e stin g b ird s p er one h e c ta re of forest is low est in
th e c u ltu re stage a n d grow s th ro u g h o u t the sere (Fig. 60). In c u ltu re s several
y e a rs old th e figure is 13, co m p ared to 33 c o u p le s /h a in large m a tu re com plexes.
In sm all enclaves of m a tu re forest c h a ra c te rise d by poorer species com position,
th e n e s t d en sity is even h igher (39 c o u p le s/h a ). T his effect is m ostly d u e to the
high p roportion of chaffinch (Fringilla coelebs) a n d goldcrest (Regulus regulus)
w hich n e s t in m a tu re enclaves.
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S econdary su ccessio n of fa u n a in th e p ine forests of P uszcza B iałow ieska
75
AVES
35
30xo
c
25-
20 -
03
15-
3-5
9-17
22-24
> 1 0 0 years
ag e of fo rest
Fig. 60. C h an g es in th e a b u n d a n c e of b ird s in relatio n to th e age of p in e forest sta n d s
The s tru c tu re of th e com m unity of b ird s is a do m in an t-ty p e one (Fig. 61).
T ree pipit (A n th u s trivialis) is d o m in a n t in 3-5 -y ear-o ld p lan ta tio n s, followed by
wood lark (Lullula arborea), a n d yellow ham m er (E m beriza citrinella) h a s th e
low est sh a re . The la tte r is d o m in a n t in 9 - 17-year-old sta n d s , w here tree pipit
m oves to th e second position. The th ird position is occupied by hedge sparrow
(Prunella m odularis), a new species. In s ta n d s older th a n 20 years, th e top
position is occupied by robin (Erithacus rubecula), followed by two species:
chiffchaf (Phylloscopus collybita) a n d wood w arbler (P. sibilatrvc\). Hedge sp arro w
re ta in s its high proportion. Sim ilarly high is th e sh a re of a new species black b ird (Turdus merula). In s ta n d s older th a n 100 y e a rs the s tru c tu re of th e
avian com m unity is m u ch m ore uniform , w ith chaffinch, goldcrest a n d wood
w arbler functioning a s d o m in an ts. C rested tit (Parus cristatus) also h a s a
significant sh a re . Since older s ta n d s c o n ta in trees with hollows in th e stem s, a
g roup of hollow-dwelling b ird s also a p p e ars.
A ctual species diversity (IT) of th e avian com m unity of pine forest in cre ase s
th ro u g h o u t the secondary sere from 1.42 to 3.43 in the oldest s ta n d s (Fig. 62).
A sim ilar tre n d is noted in potential species diversity (Hmaxk w hich eq u a ls 1.58
in th e c u ltu re stage an d 4 .00 in older sta n d s . The tre n d is p re se n t th ro u g h o u t
th e sere. The degree of ap p roxim ation of th e potential diversity, m ea su red w ith
Pielou’s index, is very high in y ounger (< 25 years) s ta n d s (J=90.0% ). It falls in
older s ta n d s to 85.8% , suggesting th a t in th e case of b ird s, too, the process of
form ation of com m unities in m a tu re ecosystem s is accom panied by th e increase
of th e degree of dom inancy.
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76
P. T rojan et al.
AVES
1
2
3
4
5
6
7
8
9
10
11
12
13
3-5
9-17
22-24
> 100 years
Fig. 61. D om inance stru c tu re of co m m u n ities of n estin g b ird s (Aues) in forest s ta n d s of v ario u s ages
in Puszcza Białowieska; 1 - Fringilla coelebs, 2 - Phylloscopus sibilatrix, 3 - Regulus regulus, 4 Pams cristatus. 5 - Erithacus mbecula, 6 - Turdus philomelos, 7 - Phylloscopus colybila, 8 - Turdus
memla. 9 - Pmnella modularis. 10 - Phylloscopus trochilus, 11 - Emberiza citńnella, 12 - Anthus
tdvialis. 13 - Lullula arborea
4
3.5
3
2.5
2
1.5
1
0.5
0
aqe of forest
Fig. 62. C hanges in ac tu a l (Hj a n d p otential (Hmax) species diversity in co m m u n ities of b ird s a n d
th e degree of realisation (J) of p o ten tial diversity of th e co m m u n ities in th e sere of th e p ine forest
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S econdary su ccessio n of fa u n a in th e p in e fo rests of Puszcza B iałow ieska
6. SUMMARY
B ecause of th e econom ic value of pine tim ber, Polish pine forests have been
exploited a n d cultivated for h u n d re d s of years. Most of th e forests co n sist of
even-aged s ta n d s , w hich are u su a lly form ed a s a re su lt of artificial resto ra tio n
of com plete c u t down of trees. S ta rtin g a pine p lan ta tio n in a clearing, u su a lly
w ith a n ad dition of com panion or sh e lte r trees, initiates th e pro cess of secondary
su ccessio n . In the course of seco n d ary su ccessio n , th e forest goes th ro u g h
several stages: c u ltu re, young grow th, thicket, pole wood a n d m atu rin g sta n d
un til it re a c h e s th e m atu re stage. In Polish pine forests, th e process tak e s
12 0 -1 5 0 y e a rs ( S z u j e c k i 1980).
E ach of th e se stag es is ch aracterized by slightly different forest m icroclim atic
conditions, w hich depend on th e com position of the s ta n d an d h a b ita t pro p erties
(O b m iń sk j 1978). The ch an g es influence th e rec o n stru c tio n of the p la n t cover.
S econdary su c ce ssio n p ro cesses in th e pine forests of Puszcza Białow ieska are
rem ark ab ly creative in th is resp ect a s they lead to th e form ation of the sp atial
s tru c tu re typical of the coniferous forest w ith c o n sp ic u o u s vertical alignm ent
a n d to filling th e living space of th e forest interior w ith species a d a p te d to the
h a b ita t conditions w hich have developed in the co u rse of secondary su ccessio n .
It is a repetitive p ro cess th a t h a s been observed an d described in coniferous
forest h a b ita ts in various p a rts of th e country.
As reg a rd s fau n a, th ese p ro cesses have been only partially u n d e rsto o d and
d a ta on the secondary su ccessio n of fa u n a p e rta in only to individual g ro u p s of
a n im a ls so th a t it is im possible to fully p o rtray th e c h an g es in com position and
org an isatio n of fau n al com plexes. On th e b ack g ro u n d of the bibliography
concerned w ith ecological succession, w hich is quite a b u n d a n t, b u t often poorly
related to “th e real thing", th e a u th o rs of th is p a p e r have a d d re sse d several
q u e stio n s th a t take into a c co u n t the c h a ra c te r of fau n istical d ata, e n su rin g th a t
a w ell-docum ented an sw er can be obtained.
The problem w hich a p p e a rs cen tral to o u r u n d e rs ta n d in g of the topic is th e
c h a ra c te r of th e secondary su ccessio n of fa u n a - w h a t a n d in w h at way does
th is p ro cess lead to. It is by no m ea n s obvious, u nlike the case of forest p lan t
cover, th a t in th e pro cess of secondary su c ce ssio n of fa u n a th e b io m ass in cre ase s
considerably an d sp atial s tru c tu re s are form ed. Even if th ese p h en o m en a
actu ally tak e place, they still need to be evidenced for. The m ost im p o rta n t
q u e s ti o n is w h e t h e r th e s e c o n d a r y s u c c e s s io n o f f a u n a is u n i- or
m ultidirectional. The qu estio n c o n c ern s th e very m odel of the p ro cess - w h e th e r
it is entirely creative, an d w h eth er its sta g e s co n sist in po p u latin g th e ecological
vacu u m , form ing ecosystem s a n d a d a p tin g th em to the ecological s tru c tu re of
th e m a tu rin g forest. T his clear pictu re ra ise s a n u m b e r of d o u b ts. Is th e re an
a c tu a l ecological v acu u m d u rin g som e tim e after clear-cu ttin g , or is th e open
a re a im m ediately occupied? Do th e v ario u s su b sy ste m s of th e zoocenosis have
to com e into being or do they alread y exist in n a tu re ? And, finally, w h a t does
th e a d a p ta tio n c o n sist in? More exactly, w h a t do the v ario u s fau n al g ro u p s
actu ally a d a p t to?
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78
P. T rojan et al.
In o rd er to o b tain a n unequivocal a n sw er to th e above q u estio n s, two w orking
h y p o th ese s were p u t forw ard, w hich w ere th e n verified w ith th e m a te ria ls
collected a n d analysed.
T he first h y p o th esis s ta te s th a t th e seco n d ary successio n of fa u n a can develop
in v a rio u s w ays a n d som etim es lead to d issim ilar final sy stem s. Five ty p es of
su c ce ssio n have been described: creative, stabilising, rise-and-fall, regressive
a n d resto rativ e (Fig. 3).
T he second hyp o th esis in te rp re ts th e in te rd ep e n d e n c ie s w ith in th e fa u n a a s
a h iera rc h ica l system of tro p h ic relatio n s. T he core of th e s tru c tu re of th e fa u n a
is accordingly form ed of sy ste m s related to one a n o th e r by m e a n s of feeding
relatio n s, su c h a s su b sy ste m , food ch ain , com petitive guild a n d po p u latio n of a
given a n im al species. T hese in te rre la tio n s have been p rese n ted on the b a sis of
30 fau n a l g ro u p s included in th e s tu d y of th e final stage of seco n d ary su c ce ssio n
- th e fa u n a of a m a tu re pine forest.
So form ulated, th e w orking h y p o th ese s have m ade it possible to com pare
fau n al d a ta w ith th e ideas a b o u t th e c o u rse of secondary su ccessio n . T he
following d o c u m e n tatio n m ate ria ls for each of the age c la sse s provided a b a s is
for verifying th e hypotheses:
1) th e n u m b e r of species form ing th e tax o cen es or com petitive guilds:
2) th e a b u n d a n c e of th e se u n its, u su a lly expressed in th e form of indices in
o rd er to illu stra te tre n d s in a b u n d a n c e changes:
3) th e s tru c tu re of th e taxocenes for illu stra tin g ch an g es in th e o rg an isatio n
of zoocenotic system s:
4) a c tu a l a n d p o tential species diversity a n d th eir ratio m e a su re d w ith Pielou’s
index.
T he n u m b e r of species (Tab. II) ex h ib its six types of c h an g es in the c o u rse of
th e se co n d a ry su ccessio n of pine forests in P uszcza B iałow ieska. T he d o m in a n t
tre n d is resto rativ e su ccessio n w here a high n u m b e r of species o c c u rs in the
pine s ta n d in the c u ltu re stage, d ecreasin g in the pole wood stag e to rise again
in th e m a tu re forest. T his tre n d w as observed in h a lf of the com petitive guilds
a n d 8 taxocenes stu d ied . The seco n d position is occupied by two o th e r m odes
of seco n d ary su ccessio n . Regressive su ccessio n , in th e co u rse of w hich th e
n u m b e r of species c o n sta n tly decreases, is found in 4 com petitive g ro u p s a n d 5
taxocenes. Sim ilarly, th e rise-and-fall su ccessio n , w here a n initial in cre ase in
th e n u m b e r of species is followed by a fall, w as observed in 5 com petitive groups.
C reative su ccessio n , ch aracterized by a c o n s ta n t grow th in th e su c ce ssio n
series, w as only found in 4 com petitive g ro u p s. S tab ilisatio n of th e n u m b e r of
species following a n initial in cre ase o ccu rred only in 2 g ro u p s of an im als. A part
from th e se 5 types of ch an g es in th e n u m b e r of species, a b se n c e of su c h c h a n g es
in th e c o u rse of forest developm ent w as observed in th e following 5 in sta n c es:
t h e l e a f b e e t l e s [C hrysom elidae), l a d y - b i r d s {Coccinellidae), p a r a s i ti c
Pipunculidae, n e m a to d es (N em atoda) a n d R hagionidae (Diptera).
A sim ilar diversification of re sp o n se s of individual an im al g ro u p s to the
reg en eratio n of a pine forest c a n be seen w h e n an aly sin g th e com petitive guilds
se p ara te ly (Tab. II). W here a g ro u p is com posed of a n u m b e r of tax o n s, th e u su a l
finding is th a t each of th e tax o n s follows a different p a tte rn of c h a n g e s in the
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79
S eco n d ary su cc e ssio n of fa u n a in th e p in e fo rests of Puszcza B iałow ieska
n u m b e r of species. Z oophages a s well a s xylo- an d cam biophages a re exceptional
in th is resp ect, rep re sen tin g m ostly the restorative type of c h an g es in th e n u m b e r
of sp e cie s, w hile th e d e c re a s in g tr e n d s prevail in th e c a se of ch ew in g
ph y to p h ag es.
T able II. C h an g es in th e n u m b e r of sp ecies of an im al co m m u n ities in th e co u rse of th e seco n d ary
su c c e ssio n of th e pine forests in P uszcza B iałow ieska
S u b sy stem
no ch an g es
-
1
-
1
-
1
-
-
2
-
B iotrophes
1
xylo- a n d cam b io p h ag es
-
1
-
-
2
-
zoophages
-
-
1
-
3
-
ap h id o p h ag es
1
-
1
1
-
1
p arasito id s
-
-
1
-
1
1
sap ro p h a g e s
-
1
1
-
-
1
soil p a n to p h a g e s a n d p re d ato rs
-
-
-
1
1
1
birds
1
m am m als
Total
regressive
1
stab ilisin g
su ck in g p h y to p h ag es
chew ing p h y to p h ag es
creativ
resto rativ e
C om petitive guild
rise-and-fall
S u ccessio n type
-
-
-
1
-
-
4
2
5
5
8
5
The re s u lts of the an aly sis of c h a n g es in th e basic co m p o n en ts of forest
ecosystem o rganisation, a s one can call anim al species, indicate th a t th e
co nstructive p ro cesses of expansion or reg en eratio n of system s, re p re se n te d by
creative, stabilising a n d restorative su ccessio n , a re a s m arked a s th e d e stru c tiv e
processes reflected in th e final outcom e a s a fall in th e n u m b e r of species. At
th e sam e tim e it is sym ptom atic th a t one o u t of every six g ro u p s of a n im a ls does
not respond to the p ro cesses operating in th e developing forest by ch an g in g th e
n u m b e r of species.
The c h a n g es in th e a b u n d a n c e of an im al com m unities (Tab. Ill) are a s diverse
a s the tre n d s in the n u m b e r of species. However, two p a tte rn s prevail: reg ressio n
a n d resto ra tio n . A bout 1/3 of th e anim al com m unities are m ore a b u n d a n t a t th e
beginning of su c ce ssio n th a n in m a tu re s ta n d s , so th a t th e decreasin g tre n d
p redom inates. The second m ost freq u en t type of su c ce ssio n is seen in th o se
com m unities w herein in th e course of su c ce ssio n th e decrease is a rre s te d a n d
a b u n d a n c e in c r e a s e s in o ld er s ta n d s . A s te a d y in c re a s e in a b u n d a n c e
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P. T rojan e t al.
th ro u g h o u t the su ccessio n series is reported only in one ou t of every 7 taxocenes.
Also in every seventh group no a b u n d a n c e c h an g es are recorded resu ltin g in a
c o n s ta n t d en sity th ro u g h o u t th e series. A d ecrease following a n initial rise is
observed in every fifteenth taxocene, while in one case only, nam ely in sp rin g tails
(Collembola), the c h an g es in a b u n d a n c e indicate stab ilisatio n in older s ta n d s.
T able III. Types of ch an g e s in species a b u n d a n c e in an im al c o m m u n ities in th e co u rse of th e
secondary su ccessio n of th e p in e forests in P uszcza Białow ieska
S u cce ssio n type
regressive
resto rativ e
no c h a n g e s
M acrotrophes
rise-and-fall
S ap ro tro p h es
stab ilisin g
B iotrophes
C om petitive guild
creativ
S u b sy stem
suck in g ph y to p hag es
1
-
-
2
-
-
chew ing p h y to p h ag es
-
-
1
-
-
2
xylo- a n d cam b io p h ag es
-
-
-
1
2
-
zoophages
-
-
-
3
1
-
ap h id o p h ag e s
2
-
-
1
1
-
p arasito id s
-
-
-
1
1
-
sap ro p h a g es
-
1
-
-
1
1
soil p an to p h a g e s an d p re d ato rs
-
-
1
1
1
-
birds
1
-
-
-
-
-
m am m als
-
-
-
-
1
1
4
1
2
9
8
4
Total
W hen th e p h e n o m e n a a re a n a ly se d from th e view point of in d iv id u al
com petitive groups, it can be seen th a t th e resp o n se s to su ccessio n w ithin each
group are visibly diversified. In each g roup of in v erteb ra te s one can find
taxocenes w hose a b u n d a n c e in creases steadily, a n d o th ers, w hose a b u n d a n c e
falls. N on-specialised zoophages are an exception in th is resp ect since th e
prevailing ten d en cy in th a t group is a decrease in a b u n d a n c e .
A nalysis of c h an g es in a b u n d a n c e of th e fa u n a provides a n a n sw er to two
im p o rtan t q u e stio n s related to secondary su ccessio n of pine forests. T he notion
of a “fau n al v a c u u m ” is not com patible w ith the facts. The forest clearing is
im m ediately occupied by anim al p o p u latio n s of high a b u n d a n ce ; alm o st % of
anim al co m m u n ities decrease th eir a b u n d a n c e in the co u rse of succession.
In creasin g a b u n d a n c e c a n be observed only in Va of the anim al g ro u p s
(Tab. Ill): su ck in g phytophages, ap h id o p h ag es a n d birds.
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S eco n d ary su ccessio n of fa u n a in th e p ine fo rests of P u szcza B iałow ieska
81
A b u n d an ce is a m ea su re of the ecological su c c e ss of b o th th e species a n d th e
a n im a l com m unity. In th e co u rse of seco n d ary su ccessio n of a pine forest,
c h a n g e s are seen in th e m a n n e r of u tilisa tio n of th e m ass p ro d u ced by th e green
p la n ts. As th e forest develops, p h y tophages w ith a biting m o u th a p p a ra tu s w hich
e a t conifers of pine are replaced by th e less d a n g e ro u s su ck in g ph y to p h ag es
w hich su c k ou t the p ro d u cts of assim ilation. The p h en o m en o n can be seen at
le a st in Puszcza Białow ieska, w here th e pine forests a re n o t su b ject to d isa s tro u s
in v asio n s of th e chew ing phytophages. An in crease in th e a b u n d a n c e of th e
s u c k in g p h y to p h a g e s is a c c o m p a n ie d by th e d ev elo p m en t of g ro u p s of
a p h id o p h a g e s w hich regulate th e a b u n d a n c e of th e form er. Very im p o rta n t is
th e in crease in a b u n d a n c e of th e com petitive group of birds, m ostly insectivores,
w hich act a s a n elem ent controlling o th er elem ents, p a rticu larly th e su b sy ste m
of biotrophes.
C h a n g e s in a c t u a l s p e c i e s d i v e r s it y (T ab . IV), m e a s u r e d w ith
S hannon-W eaver’s index (Hj, are m ost uniform ly d istrib u ted am ong th e v arious
m odels of succession. The m ost frequent p ro cesses are red u ctio n of diversity in
co m p ariso n w ith th e initial situ a tio n , b u t, all in all, the in creasin g tre n d in th e
diversity of anim al com m unities is en c o u n te re d a s frequently a s th e reverse
process. A nother frequently observed ph en o m en o n is a fall in diversity after a
period of rise. The p ictu re c h a n g es a little w hen one considers it from th e
view point of realising th e p o tential for species diversity in th e ecosystem . Here,
th e prevailing p ro cess is red u ctio n of diversity, especially in m a tu re s ta n d s.
R eduction w as observed in 17 com m unities o u t of 28 investigated in th is respect.
Increase, resto ra tio n or stab ilisatio n of diversity after a n in crease w ere registered
in only 1/3 of th e cases. A bsence of ch an g es of diversity th ro u g h o u t th e
su c ce ssio n series is very rare.
T he re s u lts of th e above a n a ly sis indicate clearly th a t M acA rthur's (1958)
th e sis “diversity m akes stab ility ” is not confirm ed by em pirical d a ta . On th e
co n trary , th e decrease in species diversity observed in th e m a tu re s ta n d s of th e
pine forest im plies th a t the in crease in th e degree of org an isatio n of a system
ta k e s place a t th e expense of decreasing its in te rn al diversification. T his is
related to com m unity stru c tu re . In m a tu re sta n d s , th ere is u su a lly one or a few
d o m in a n t species in a com m unity a n d th e “tail” of accessory species becom es
n otably shorter. T he org an isatio n of a com m unity is u su a lly b u ilt on a few
w ell-adapted species form ing com petitive groups.
In conclusion it sh o u ld be s tre sse d th a t th e m ost o u tsta n d in g fea tu re of
seco n d ary su ccessio n of th e fa u n a in th e pine forests in Puszcza B iałow ieska is
th e m ultidirectional c h a ra c te r of p ro cesses tak in g place in a growing pine forest
w hich is developing its spatial stru c tu re . If one co n sid ers the p a ra m e te rs
studied: th e n u m b e r of species, a b u n d a n c e , species diversity an d s tru c tu re , it
is e a sy to n o tic e t h a t th e d e c re a sin g tre n d s in th e p a ra m e te rs stu d ie d
pred o m in ate - e ith e r th ro u g h o u t th e su ccessio n series or in its final stag e (44%
of c a se s). S tab ilisatio n of th ese p a ra m e te rs th ro u g h o u t th e series or a fte r a period
of grow th can only be seen in 22% of cases. An increase, eith er ste a d y or following
a period of reduction, is only observed in 34% of th e cases.
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82
P. T rojan et al.
T able IV. T ypes of ch an g es in a c tu a l sp ecies diversity (Hj in an im al co m m u n ities in th e co u rse of
th e se c o n d a iy su ccessio n of th e p in e fo rests in P uszcza B iałow ieska
S u ccessio n type
regressive
resto rativ e
no ch a n g e s
su ck in g p h y to p h ag es
1
-
-
2
-
-
chew ing p h y to p h ag es
-
-
1
-
-
2
xylo- a n d cam b io p h ag es
1
1
-
-
1
-
creativ
rise-an d -fall
Com petitive guild
stab ilisin g
S u b sy ste m
B iotrophes
Sapr oti o p h cs
zoophages
1
1
1
-
2
-
ap h id o p h ag e s
1
-
1
2
-
-
p arasito id s
-
1
1
1
-
-
sap ro p h a g e s
-
1
1
-
-
1
soil p a n to p h ag e s a n d p re d a to rs
-
1
-
1
-
-
b ird s
1
-
-
-
1
-
-
5
5
5
7
3
3
m am m als
Total
T h ese d a ta confirm th e h y p o th esis of m ultid irectio n al c h a ra c te r of the
p ro c e s s e s of se c o n d a ry s u c c e s s io n in a n im a l c o m m u n itie s. P a rtic u la rly
im p o rta n t is th e decrease in th e value of m any indices c h aracterizin g th e fau n a
in th e final stage of su ccessio n , w hich ca n be term ed th e stage of com m unity
o rg an isa tio n . It tak e s place a t th e cost of reducing th e n u m b e r of species an d
diversity for th e sa k e of sta b ilisatio n of d om inance s tru c tu re s . Ecosystem
sta b ility is th u s e n su re d by th e b e tte r a d a p te d form s w hile th e diversity potential
is n o t fully utilised.
All elem en ts of th e zoocenotic sy stem are form ed sim u lta n e o u sly in th e early
stag e of p lan tin g th e s ta n d after clear-cu ttin g . At th is stag e th e sy stem is the
ric h e st. In th e s u b s e q u e n t sta g e s its o rg an isatio n is only re s tru c tu re d in
a c co rd a n ce w ith th e developm ent of th e forest a s a n ecosystem .
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8. AN INVENTORY OF SPECIES OCCURRING IN THE PINE FORESTS
OF PUSZCZA BIAŁOWIESKA
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LACHNIDAE {APHIDOIDEA, HOMOPTERA)
Cinara pinea ( M o r d r . )
Cinara pilosa ( Z e t t .)
Cinara pini (L.)
Cinara pinihalbitans ( M o r d r . )
Schizolachnus pineti (F.)
Eulachnus agilis ( K a lt.)
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AUCHENORRHYNCHA (HOMOPTERA)
Delphacidae
Criomorphus albomarginatus C u r t .
Javesella pellucida (F.)
Paradelphacodes paludosus ( F t.)
Cicadidae
Cicadetta montana (S c o p .)
Cercopidae
Aphrophora corticea (G e rm .)
Membracidae
Centrotus cornutus (L.)
Cicadellidae
Oncopsis Jlauicollis (L.)
Empoasca vitis ( G o e th e )
Eupteryx atropunctata ( G o e z e )
Eurhadina pulchella ( F a l l .)
Wagneripteryx germari ( Z e t t .)
Arboridia parvula (Bon.)
ZyginaJlammigera ( F o u r c r . )
Allygus mixtus (F.)
Colobotettix morbillosus (M e l.)
Grypotes puncticollis (H.-S.)
Hesium domino ( R e u t.)
Jassargus Jlori (F ie b .)
Macrosteles laevis (R ib.)
Psammotettix alienus ( D h lb .)
Speudotettix subfusculus ( F a l l .)
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HETEROPTERA
Sehirus Inctuosus M u l s . e t R .
Adomerus biguttatus (L.)
Tritomegas bicolor (L.)
Aelia acuminata (L.)
Aelia klugi H a h n
Eysarcoris aeneus (S c o p .)
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Stagonomus pusillus (H .-S .)
Holcostethus vernalis ( W o l f f .)
Palomena prasina (L.)
Palomeria viridissima (P o d a )
Pitediajuniperina (L.)
Pitedia pinicola ( M u ls . e t R .)
Carpocoris purpureipennis (D e G e e r )
Carpocoris fuscispinus (B o h .)
Carpocoris pudicus (P o d a )
Dolycoris baccarum (L.)
Eurydema oleraceum (L.)
Piezodorus lituratus (F.)
Picromerus bidens (L.)
Troilus luridus (F.)
Rhacognathus punctatus (L.)
Acanthosoma haemorrhoidale (L.)
Elastotethus interstinctus (L.)
Elasmuchaferrugata (F.)
ElasmuchaJieberi (J a k .)
Elasmucha grisea (L.)
Coreus marginatus (L.)
Corizus hyoscyami (L.)
Rhopalus parumpunctaius ( S c h i l l .)
Rhopalus maculatus (F ie b .)
Myrmus miriformis ( F a l l .)
Stictopleurus pictus (F ie b .)
Dicranocephalus agalis (S c o p .)
Nithecusjacobeae ( S c h i l l . )
Nysius thymi ( W o l f f )
Nysius ericae ( S c h i l l .)
Nysius helveticus (H.-S.)
Kleidocerys resedae (P a n z .)
Cymus obliguus ( H o r v .)
Cymus claviculus ( F a l l .)
Cymus melanocephalus (F ie b .)
Geocoris grylloides (L.)
Geocoris ater (F.)
Stygnocoris pygmaeus (F. S a h l b . )
Stygnocoris pedestris ( F a l l .)
Megalonotus chiragra (F.)
Megalonotus sabulicola (T h o m s.)
Pterotmetus staphyliformis ( S c h i l l .)
Rhyparochromus pini (L.)
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S econdary su ccessio n of fa u n a in th e p in e fo rests of Puszcza B iałow ieska
Ischnocoris angustulus (B o h .)
Drymiis ryei D o u g l . e t Sc.
Drymus brunneus (F. S a h l b . )
Eremocoris plebejus ( F a l l .)
Eremmocoris abietis (L.)
Scolopostethus decoratus (H a h n )
Pachybrachius fracticollis ( S c h i l l .)
Neides tipularius L.
Saldula uestita ( D o u g l.)
Acalypta ccuinata (P a n z .)
Acalypta nigrina ( F a l l .)
Acalypta marginata ( W o lf f )
Acalypta gracilis (F ie b .)
Himacerus apterus (F.)
Nabicula limbata ( D a h lb .)
Nabis Jerus (L.)
66 Nabis pseudoferus R em .
67 Nabis punctatus A . C o s t a
68 Nabis ericetorum S c h o l t z
69 Nabis rugosus (L.)
70 Myrmedobia exilis ( F a l l .)
71 Myrmedobia distinguenda R e u t .
72 Anthocoris nemomm (L.)
73 Anthocoris confusus R e u t .
74 Acompocoris pygmaeus ( F a l l .)
75 Orius niger W o l f f
76 Orius minutus (L.)
77 Orius agilis ( F l o r )
78 Monalocoris Jilicis (L.)
79 Bothynotus pilosus (B o h .)
80 Deraeocoris ruber (L.)
81 Alloeotomus gothicus ( F a l l .)
82 Myrmecoris gracilis ( J . S a h l b . )
83 Leptopternaferrugata ( F a l l .)
84 Leptoplerna dolobrata (L.)
85 Stenoderma calcaratum ( F a l l .)
86 Stenoderma uirens (L.)
87 Stenoderma laevigatum (L.)
88 Stenoderma holsatum (F.)
89 Notosistra erratica (L.)
90 Trigonotylus coelestialium (K irk .)
91 Trigonotylus ruficornis ( G e o f f r . )
92 Phytocoris longipennis F l o r
93 Phytocoris pini K i r s c h b .
94 Adelphocoris reicheli (F ie b .)
95 Caloris biclavatus (H.-S.)
96 Caloris ajfinis (H.-S.)
97 Caloris noruegicus (G m e l.)
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Stenotus binotatus (F.)
Lygocoris pabulinus (L.)
Lygus rugulipennis (P opp.)
Lygus pratensis (L.)
Lygus punctatus ( Z e t t .)
Camptozygum aequale (V ill.)
Charagochilus gyllenhali ( F a l l .)
Capsus piliferREM.
Capsodes gothicus (L.)
Halticus pusillus (H.-S.)
Heterotoma meriopterum (S c o p .)
Heterocordylus genistae [S c o p .)
Orthotylus ericetorum ( F a l l .)
GlobicepsJlavomaculatus (F.)
Globiceps cruciatus R e u t .
Blepharidoptems breuicoornis (W a g n .)
Phoenicocoris modestus (M e y .-D .)
Phoenicocoris obscurellus ( F a l l .)
Atractotomus mali (M e y .-D .)
Psallus varians (H.-S.)
Plesiodema pinetellum ( Z e t t .)
Lopus decolor ( F a l l .)
SYMPHYTA
Xyelajulii ( B r e b .)
Acantholyda posticalis ( M a ts .)
Cephalcia arvensis (P a n z .)
Pamphilius hortomm (K lu g )
Pamphilius marginatus (L ep.)
Pamphilius sylvaticus (L.)
Pamphilius oarius (L ep.)
Janus Jemoratus ( C u r t . )
Janus luteipes (L ep.)
Arge dimidiata ( F a l l .)
Arge ustulata (L.)
Microdiprion pakkipes ( F a l l .)
Macrophya duodecimpunctata (L.)
Eriocampa ouata (L.)
Athalia bicolor L ep.
Athalia circularis (K lu g )
Athalia cordata L ep .
Athalia cordatoides P r i e s n .
Athalia glabricollis T h o m s o n
Athalia lugens (K lu g )
Athalia rosae (L.)
Monosoma pulueratum (R e tz .)
Empria longicornis (T h o m so n )
Ametastegia equseti ( F a l l .)
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Ametastegia glabrata ( F a l l .)
Ametastegia tener (S p in o la )
Allantiis cingillum ( K lu g )
Allantus togatus (P a n z .)
Harpiphoms lepidus (K lu g )
Monophadnus pallescens (G m e l.)
Stethomostus fulginosus ( S c h r k .)
Eutomostethus luteiventris (K lu g )
Periclista albida (K lu g )
Claremontia alternipes (K lu g )
Claremontia confusa (K o n .)
Diprion similis ( H a r t . )
Gilpinia polytoma ( H a r t . )
Gilpinia virens (K lu g )
Gilpiniafrutetorum ( F a b r .)
Strongylogaster lineala ( C h r i s t )
Strongylogaster macula (K lu g )
Strongylogaster mixta (K lu g )
Aneugmenus padi (L.)
Birka cinereipes (K lu g )
Nesoselandria morio ( F a b r .)
Selandria serva ( F a b r .)
Dolerus gessneri A n d r e
Aglaostigmafulvipes (S c o p .)
Tenthredopsis excisa (T h o m so n )
Tenthredopsis fresei (K o n .)
Tenthredopsis litterata ( G e o f f . )
Tenthredopsis nassata (L.)
Tenthredopsis scutellaris ( F a b r .)
Rhogogaster picta ( K lu g )
Rhogogaster viridis (L.)
Tenthredo atra (L.)
Tenthredo livida (L.)
Tenthredo maculata ( G e o f f . )
Halidamia affinis ( F a l l .)
Caliroa cinxia (K lu g )
Hetererthrns nemoratus ( F a l l .)
Hetererthrus ochropodus (K lu g )
Hetererthrns uagans ( F a l l .)
Scolioneura betuleti (K lu g )
Messa glaucopis (K o n .)
Messa nana (K lu g )
Projenusa pygmaea (K lu g )
Profenusa thomsoni (K o n .)
Fenusa dohrnii (T is c h .)
Fenusa pusilla (L ep.)
Cladius pectinicornis ( G e o f f .)
Priophorus morio (L ep.)
Priophoms pallipes (L ep.)
Hoplocampa flava (L.)
Hoplocampa testudinea (K lu g )
Mesoneura opaca ( F a b r .)
Dineura uiridorsata ( R e tz .)
Hemichroa australis (L ep.)
Hemichroa crocea ( G e o f f . )
Nematinus lutens (P a n z .)
Pristiphora cincta (N e w m a n )
Pristiphora conjugata ( D h lb .)
Pristiphora melanocarpa ( H a r t . )
Pristiphora pallidiuentris ( F a l l .)
Pristiphora ruficornis (O liv .)
Pristiphora subbifida (T h o m .)
Pristiphora testacea ( J u r i n e )
88 Pristiphora alpestris (K o n .)
89 Pristiphora mollis ( H a r t . )
90 Pristiphora abietina ( C h r i s t )
91 Pristiphora compressa ( H a r t . )
92 Pristiphora decipiens ( E n s l.)
93 Pristiphora gerula (K o n .)
94 Pristiphora leucopodia ( H a r t . )
95 Pristiphora pallida (K o n .)
96 Pristiphora pseudodecipiens (B & K)
97 Pristiphora sexesenii ( H a r t . )
98 Pristiphora tenuicornis ( L in d q u is t)
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P ristiphora te n u ise rra (L in d q u is t)
S harliphora amphibola ( F o e r s t e r )
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Sharliphora nigella ( F o e r s t e r )
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102 Sharliphora parva ( H a r t . )
Pikonema insigne ( H a r t . )
Pikonema montanum (Z a d d .)
Pikonema nigriceps ( H a r t . )
Pikonema pallescens ( H a r t . )
Pikonema scutellatum ( H a r t . )
Croesus septentrionalis (L.)
Nematus princeps (Z a d d .)
1 10 Nematus cadderensis C am .
111 Nematus dispar B r i s c h .
1 12 Nematus umbratus T h o m .
113 Nematus viridis S t e p h .
114 Euura mucronata ( H a r t . )
1 15 Potania coriacea (B e n s .)
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CURCULIONIDAE
1 Rhinomacer attelaboides F .
2 Doydirhynchus austriacus (O liv .)
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Pselaphorhynchites longiceps (T h o m s.)
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Pselaphorhynchites nanus (P ay k .)
Deporaus betulae (L.)
Deporaus mannerheimi (H um m .)
Attelabus nitens (S c o p .)
Apoderus coryli (L.)
Apion curtirostre G e r m .
Apion simile K ir b y
Apion virens H e r b s t
ApionJulvipes ( G e o f f r . )
Otiorhynchus ovatus (L.)
Phyllobius arborator ( H e r b s t )
Polydrusus pallidus ( G y ll.)
Polydrusus pilosus G r e d l .
Brachyderes incanus (L.)
Strophosoma capitatum (D e g .)
Sitona lineatus (L.)
Hypera zoilus (S c o p .)
Hypera nigrirostris (F.)
Anthonomus phyllocola ( H e r b s t )
Brachonyx pineti (P ay k .)
Curculio rubidus G y l l .
Curculio pyrrhoceras M a r s h .
Magdalis exarata (B r is .)
Magdalis phlegmatica ( H e r b s t )
Magdalis nitida ( G y ll.)
Magdalis linearis ( G y ll.)
Magdalis violacea (L.)
Magdalis duplicata G e rm .
Hylobius abietis (L.)
Pelenomus quadricorniger ( C o lo n n .)
Rhinoncus bruchoides ( H e r b s t )
Rhinoncus castor (F.)
Coeliodes ruhicundus ( H e r b s t )
Ceutorhynchus pleurostigma ( M r s h .)
Ceutorhynchus assimilis (P ay k .)
Ceutorhynchus erysimi (F.)
Ceutorhynchus punctiger ( G y ll.)
Ceutorhynchus Jloralis (P ay k .)
Anoplus plantaris (N a e z .)
Rhynchaenus avellanae (D o n o v .)
Rhynchaenus rusici ( H e r b s t )
Rhynchaenus stigma (G e rm .)
Rhamphus pulicarius ( H e r b s t )
CHRYSOMELIDAE
1 Cryptocephalus Jlavipes F a b r .
2 Cryptocephalus parvulus M u e l l .
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Cryptocephalus pini (L in n .)
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Cryptocephalus punctiger P a y k .
Cryptocephalus pusillus F a b r .
Cryptocephalus saliceti Z e b e
Plagiodera versicolora (L a ic h .)
Chrysomela lapponica L.
Phratora atrovirens ( C o r n e l . )
Phratora laticollis ( S u f f r . )
Phratora vitellinae (L.)
Phratora vulgatissima (L.)
Galerucella lineola ( F a b r .)
Galerucella calmariensis (L.)
Phyllotreta vittula {R e d t .)
Phyllotreta nemorum (L.)
Phyllotreta undulata K u t s c h . )
PhyllotretaJlexuosa (III.)
Phyllotreta vittata (F.)
Aphthona euphorbiae ( S c h r a n k )
Longitarsus longiseta W e i s e
Longitarsus nasturtii (F.)
Longitarsus parvulus (P ay k .)
Altica quercetorum F o u d r .
Asiorestia ferruginea (S c o p .)
Chaetocnema heikertingeri L j u b .
Chaetocnema hortensis ( G e o f f r . )
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LEPIDOPTERA
Tortricidae
Acleris laterana (F.)
Acleris sparsana (D e n . e t S c h i f f . )
Acleris maccana ( T r e it.)
Eana incanana ( S te p h .)
Cnephasia communana (H.-S.)
Cnephasia stephensia ( D o u b l.)
Cnephasia asseclana (D e n . et S c h i f f . )
Cnephasia genitalana ( P i e r c e et M e c t .)
Capua vulgana F r o l .
Ramapesia gnomana ( C l e r c k )
Argyrotaenia ljugiana (T h n b g .)
Archips oporanus (L.)
Archips xylosteanus (L.)
Archips rosanus (L.)
Choristoneura diversana (H b n .)
Ptycholoma lecheana (L.)
Pandemis cerasana (H b n .)
Syndemis musculana (H b n .)
Adoxophys orana (F. v. R.)
Hedya atropunctana ( Z e t t .)
Cymolomia hartigiana (S a x .)
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22 Orthotaenia undulana (D e n .
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e t S c h if f .)
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Pseudohermenias abietana ( F a b r .)
Apotomis turbidana (H b n .)
Apotomis betuletana (H aw .)
Olethreutes bifasciana (H aw .)
Olethreutes lacunana (D e n . e t S c h i f f . )
Ancylis laetana (F.)
Ancylis upupana ( T r e it.)
Ancylis badiana (D e n . e t S c h i f f . )
Ancylis unculana (H aw .)
Ancylis apicella (D e n . e t S c h i f f . )
Epinotia solandriana (L.)
Epinotia tenerana ( D e n . e t S c h i f f . )
Epinotia tedella ( C l e r c k )
Rhopobota naevana (H b n .)
Spilonota ocellana (D e n . e t S c h i f f .)
Blastesthia tnrionella (L.)
Rhyacionia buoliana (D e n . e t S c h i f f .)
Rhyacionia pinicolana ( D o u b l.)
Rhyacionia pinivorana (L ie n , e t Z e l l . )
Rhyacionia duplana (H b n .)
Retinia resinella (L.)
Pammene obscurana ( S te p h .)
Pammene argyrana (H b n .)
Cydia cosmophorana ( T r e it.)
Cydia coniferana (S a x .)
Choreutidae
AnthophilaJabriciana (L.)
Pyralidae
Assara terebrella (Z in c k .)
Crambus lathoniellus (Z in c k .)
Crambus ericellus (H b n .)
Crambus heringiellus (H .-S .)
Agriphila inquinatella (D e n . e t S c h i f f .)
Catoptria margaritella (D e n . e t S c h i f f . )
Scoparia basistrigalis K n a g g s
Eudonia truncicolella ( S t t . )
Eurrhypis pollinalis (D e n . e t S c h i f f .)
Udea lutealis (H b n .)
Zygenidae
Adscita statices (L.)
Pieridae
Pieris napi (L.)
Pieris brassicae (L.)
Gonepteryx rhamni (L.)
Lycaenidae
Quercusia quercus (L.)
Callophrys rubi (L.)
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Lycaena virgaureae (L.)
Lycaena tityrus (P o d a )
Plebejus argus (L.)
Nymphalidae
Nymhalis polychloros (L.)
Inachis io (L.)
Aglais urticae (L.)
Araschnia levana (L.)
Issoria lathonia (L.)
Boloria dia (L.)
Boloria euphrosyne (L.)
Melitaea athalia ( R o t t . )
Satyridae
Aphantopus hyperantus (L.)
Coenonympha pamphilus (L.)
Coenonympha arcania (L.)
Pararge aegeria (L.)
Thyatiridae
Thyatira batis (L.)
Tethea or (D e n . e t S c h i f f . )
TetheellaJluctuosa (H b n .)
Geometridae
Jodis putata (L.)
Cyclophora punctaria (L.)
Scopula nigropiuictata ( H u f n .)
Scopulafloslactata (H a w .)
Idaea straminata ( B o r k h . )
Xanthorrhoe spadiceraria (D e n . e t S c h if f .)
Xanthorrhoe quadri/asciata ( C l e r c k )
Xanthorrhoe montanata (D e n . e t S c h i f f . )
Chloroclysta citrata (L.)
Chloroclysta truncata ( H u f n .)
Thara obeliscata (H b n .)
Thera uariata (D e n . e t S c h i f f . )
Therajuniperata (L.)
Electrophaes corylata (T h n b g .)
Hydriomena furcata (T h n b g .)
Rheumaptera ui~idulata (L.)
Euphyia unangulata (H a w .)
Epirrita autumnata ( B o r k h .)
Perizoma alchemillata (L.)
Perizoma hydrata ( T r e i t .)
Eupithecia plumbeolata (H a w .)
Eupithecia venosata ( F a b r .)
Eupithecia intricata ( Z e t t .)
Eupithecia subfuscata (H a w .)
Eupithecia indigata (H b n .)
Eupithecia pusillata ( D e n . e t S c h i f f .)
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S econdary su ccessio n of fa u n a in th e p ine fo rests of P u szcza B iałow ieska
Eupithecia tantillaria (B s d v .)
Chloroclystis debiliata (H b n .)
111 Semiothisa liturata ( C l e r c k )
1 1 2 Alcis repandata (L.)
1 1 3 Ematurga atomaria (L.)
1 1 4 Bupalus piniarius (L.)
1 1 5 Hylaea Jasciaria (L.)
1 1 6 Gnophos obscurata (D e n . e t S c h i f f .)
Notodontidae
1 17 Drymonia obliterata (E sp .)
1 18 Ptilodon capucina (L.)
Lymantriidae
1 19 Lymantria monacha (L.)
Arctiidae
120 Eilema deplana (E sp .)
Herminiidae
121 Herminia strigillata (L.)
Noctuidae
122 Rivula sericealis (S c o p .)
1 2 3 Hypena crassalis (F.)
1 2 4 Hypena proboscidalis (L.)
1 2 5 Catocala fraxini (L.)
1 2 6 Catocala sponsa (L.)
1 2 7 LaspeyriaJlexula (D e n . e t S c h i f f . )
1 2 8 Protodeltote pygarga (H u f n .)
1 2 9 Autographa gamma (L.)
1 3 0 Acronicta aceds (L.)
1 3 1 Amphipyra pyramidea (L.)
1 3 2 Dypterygia scabriuscula (L.)
1 3 3 Trachea atriplicis (L.)
1 3 4 Euplexia lucipara (L.)
1 3 5 Enargia paleacea (E sp .)
1 3 6 Apamea crenata ( H u f n .)
1 3 7 Apamea remissa (H b n .)
1 3 8 Oligia strigilis (L.)
1 3 9 Oligia latruncula (D e n . e t S c h i f f . )
1 4 0 Mesoligiafuruncula (D e n . e t S c h i f f . )
1 4 1 Mesapamea secalis (L.)
1 4 2 Celaena leucostigma (H b n .)
1 4 3 Archanara geminipuncta (H aw .)
1 4 4 Lithophanefurcifera (H u fn .)
1 4 5 Conistra vaccinii (L.)
1 4 6 Agrochola helvola (L.)
1 4 7 Polia bombycina ( H u f n .)
1 4 8 Polia tincta (B ra h m )
1 4 9 Polia nebulosa ( H u f n .)
1 5 0 Mamestra brassicae (L.)
1 5 1 Melanchra persicariae (L.)
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Lacanobia thalassina ( H u f n .)
Lacanobia suasa (D e n . e t S c h i f f . )
Orthosia incerta (H u f n .)
Mythimna albipunctata (D e n . e t S c h i f f . )
Mythimna pallens (L.)
Euxoa nigricans (L.)
Agrostis ipsilon (H u f n .)
Eugraphe subrosea ( S te p h .)
Diarsia brunnea (D e n . e t S c h i f f . )
Xestia c-nigmm (L.)
Xestia baja (D e n . e t S c h i f f . )
Eurois occulta (L.)
Anaplectoides prasina (D e n . e t S c h i f f . )
SCOLYTIDAE
1 Hylurgops palliatus (G y l l .)
2 Hylastes ater (P ayk.)
Hylastes opacus E r .
Hylurgus lignipedra ( F a b r .)
5 Tomicus minor ( H a r t . )
6 Tomicus piniperda (L.)
7 Pityogenes bidentatus ( H e r b s t )
8 Pityogenes chalcographus (L.)
9 Pityogenes quadridens ( H a r t . )
10 Pityogenes trepanatus ( N o r d l .)
11 Orthotomicus laricis (F.)
12 Orthotomicus proximus ( E ic h h .)
1 3 Orthotomicus suturalis ( G y ll.)
1 4 Ips acuminatus ( E ic h h .)
1 5 Ips sexdentatus (B o r n .)
1 6 Dryocoetes autographus (R a tz .)
1 7 Crypturgus cinereus ( H e r b s t )
1 8 Crypturgus hispidulus (T h o m s.)
1 9 Crypturgus pusillus ( G y ll.)
20 Trypodendron lineatum (O liv .)
21 Pityophthorus lichtensteini ( R a tz .)
22 Pityophthorus pityographus ( R a tz .)
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CERAMBYCIDAE
1 Prionus coriarius (L.)
2 Tetropium castaneum (L.)
Tetropiumfuscum (F.)
Spondylis buprestoides (L.)
5 Arhopalus rusticus (L.)
6 Asemum striatum (L.)
7 Molorchus minor (L.)
8 Callidium aeneum (D e G e e r )
9 Callidium violaceum (L.)
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Callidium coriacewn P a y k .
Rhagium mordax (D e G e e r )
12 Rhagium inquisitor (L.)
1 3 Pachyta quadrimaculata (L.)
1 4 Cortoderafemorata ( F a b r .)
1 5 Pedostrangalia pubescens ( F a b r .)
1 6 Leptura quadrifasciata (L.)
1 7 Anastrangalia sanguinolenta (L.)
1 8 Corymbia rubra (L.)
1 9 Brachyleptura maculicornis (D e G e e r )
20 Judolia sexmaculata (L.)
21 Alosterna tabacicolor (D e G e e r )
22 Strangalia attenuata (L.)
2 3 Stenurella melanura (L.)
2 4 Stenurella nigra (L.)
2 5 Monochamus galloprovincialis (O liv .)
2 6 Monochamus sutor (L.)
2 7 Monochamus urussouii ( F is c h .)
2 8 Pogonocherus Jasciculatus (D e G e e r )
2 9 Pogonocherus decoratus (F a irm .)
3 0 Acanthocinus griseus ( F a b r .)
3 1 Acanthocinus aedilis (L.)
3 2 Leiopus nebulosus (L.)
3 3 Saperda populnea (L.)
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Epitriptus cingulatus (F.)
Neoitamus cothumatus (Mg.)
15 Neoitamus cyanurus (Lw.)
1 6 Neoitamus socius (Lw.)
17 Neomochtherus pallipes (Mg.)
18 Pamponerus germanicus (L.)
19 Paritamus geniculatus (Mg.)
20 Tolmerus atricapillus ( F l l .)
21 Tolmerus atripes Lw.
22 Tolmerus pyragra ( Z l l .)
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ARANEl
1 Anyphaena accentuata ( W a lc k .)
2 Dictyna arundinacea (L.)
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BUPRESTIDAE
Chalcophora mariana (L.)
Buprestis haemhorroidalis H e r b s t
Buprestis novemmaculata (L.)
Buprestis octoguttata (L.)
Buprestis rustica (L.)
Phaenops cyanea F a b r .
Anthaxia quadripunctata (L.)
Anthaxia similis ( S a u n d )
Anthaxia submontana ( O b e n b .)
Agrilus angustulus (III.)
Agrilus sulcicollis ( L a c o r d .)
Trachys minuta (L.)
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ASIUDAE
Choerades fimbriata (Mg.)
Dioctria humeralis ( Z e l l .)
Dioctria hyalipennis (F.)
Dioctria oelandica (L.)
Lasiopogon cinctus (F.)
Lasiopogon immaculatus S t r .
Cyrtopogon lateralis ( F l l .)
Leptogaster pubicornis (Lw.)
Dysmachus cochleatus (Lw.)
10 Dysmachus picipes (Mg.)
11 Dysmachus trigonus (Mg.)
12 Epitriptus arthriticus ( Z ll.)
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Dictyna pusilla T i l
Cicurina cicurea Mg.
Arctosa cinerea (F.)
Xerolycosa nemoralis ( W e s t r .)
Drassodes pubescens ( T il)
Micaria silesiaca L. K.
Phrurolithusfestiuus (C.L.K.)
Micrommata roseum (C l.)
Clubiona lutescens W e s t r .
Euryclubiona stagnatilis (K u l.)
Euryclubiona subsultans (T h.)
Ballus chalybeius (W lk .)
Dendryphanthes hastatus (C.L.)
Dendryphanthes rudis (S u n d .)
Heliophanus aeneus (H a h n )
Heliophanus dubius (C.L.K.)
Heliophanusjlavipes (H a h n )
Marpissa muscosa C l .
Neon reticulatus ( B la c k .)
Phlegra v-incignita (C l.)
Pseudicius encarpatus (W a lc k .)
Salticus oleari (S c o p .)
Coriarachne depressa (C.L.K.)
Diaea dorsata (F.)
Xysticus cristatus ( C l.)
Xysticus lanio C.L.K.
Artanes dispar ( W a lc k .)
HorodromoidesJuscomarginatxis (D e G e e r )
Philodromus aureolus (C l.)
Philodromus collinus (C.L.K.)
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S eco n d ary su c ce ssio n of fa u n a in th e p in e fo re sts of P uszcza B iałow ieska
Araneus angulatus (C l .)
Araneus diadematus (C l .)
Araniella cucurbitina (C l .)
Araniella incospicua (Sim .)
Atea sturmi (H a h n )
Gibbaranea bituberculata ( W a lc k .)
Mangora acalypha ( W a lc k .)
Nuctenea umbratica ( C l.)
Pachygnatha clercki S u n d .
Tetragnatha pinicola K.
Agyneta rurestris (C.L.K.)
Centromerus sylvaticus (B l.)
Drapetisca socialis (S u n d .)
Kaestneda dorsalis (W id .)
Leptyphanthes Jlavipes (B l.)
Leptyphanthes obscurus (B l.)
Porrhomma convexum ( W e s t r .)
Porrhomma pygmaeum ( B la c k )
Ceratinella brevis (W id e r.)
Diplocephalus hiemalis F . D a h l .
Dismodicus cornutus (B l.)
Dismodicus elevatus P e t r , n e c C.L.K.
Entelecara congenera (O.P-C.)
Erigone atra ( B l.)
Erigone dentipalpis ( W id e r.)
Gongylidiellum compar ( W e s t r .)
Metopobactrus prominulis (O.P-C.)
Moebelia penicillata ( W e s t r .)
Oedothorax gibbosus (B l.)
Peponocranium orbiculatnm (O.P-C.)
Savignyafrontata B l .
Tapinocyba pallens (O.P-C.)
Trematocephalus cristatus (W id .)
66 Crustulina guttata (W id .)
67 Dipoena tristis (H a h n )
68 Enoplognatha ovata (C l .)
69 Neottiura bimaculata (L.)
70 Theridion mystaceum (L.K.)
71 Theridion pictum (W a lc k .)
72 Theridion pinastri (L.K.)
73 Theridion sisyphium (C l.)
74 Theridion tinclum (W a lc k .)
75 Theridion varians H a h n .
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SPHECIDAE
(* g a tu n k i afidofagiczne)
1 Podalonia ajjinis K ir b y
2 Ammophila campestris L a t r .
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Ammophila lujfi S a u n d .
Ammophila pubescens C u r t .
Ammophila sabulosa (L.)
Mimumesa unicolor L in d .
Psenulus concolor ( D a h lb .)
Psenulus fuscipennis ( D a h lb .) *
Psenulus laevigatus (S c h c k .)
Psenulus pallipes (P a n z .) *
Pemphredon balticus ( M e r is u o ) *
Pemphredon lethijer ( S c h u c k .) *
Pemphredon lugubris (F.) *
Pemphredon morio L in d . *
Pemphredon shuckardi (A. M o r .) *
Passaloecus borealis D a h l b . *
Passaloecus brevilabris *
Passaloecus clypealis F a e s t . *
Passaloecus corniger S h u c k . *
Passaloecus eremita K o h l. *
Passaloecus gracilis ( C u r t . ) *
Passaloecus insignis (L in d .) *
Passaloecus monilicornis D a h l b . *
Passaloecus singularis D a h l b . *
Passaloecus turionum D a h l b . *
Spilomena beata B l u t h g .
Spilomena difjerens B l u t h g .
Spilomena punctatissima B l u t h g .
Spilomena troglodytes (L ind.)
Spilomena vagans B l u t h g .
Tachysphex obscuripennis ( S c h e n c .)
Tachysphex pompiliformis (P a n z .)
Tachysphex psammobius (K o h l)
Nitela spinolae L a t r . *
Trypoxylon attenuatum S m ith
Trypoxylon clavicerum L e p . et S e r v .
Trypoxylonfigulus (L.)
Trypoxylon medium B e a u m o n t
Trypoxylon minus B e a u m o n t
Entomognatus brevis (L ind.)
Lindenius albilabris (F.)
Lindenius panzeri (L in d .) *
Rhopalum clavipes (L.) *
Rhopalum coarctatum (S c o p .)
Crossocerus annulipes (L ep. et B r u l .
Crossocerus barbipes ( D a h lb .)
Crossocerus cetratus S h u c k . *
Crossocerus dimidiatus (F.)
Crossocerus inermis t h o m s .
Crossocerus megacephalus (Ros.)
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Crossocems nigritus (L ep. et B r u l . )
Crossocerus ovalis (L ep. et B r u l . )
Crossocerus palmipes (L.)
Crossocems pusillus L ep . et B r u l .
Crossocems quadrimaculatus (F.)
Crossocems styrius ( K o h l)
Crossocems walkeri ( S h u c k .)
Crossocems wesmaeli (L in d .) *
Crabro cribrarius (L.)
Crabro peltarius ( S c h r e b . )
Crabro scutellatus ( S c h e v .)
Ectemnius cavifrons (T h o m s.)
Ebtemnius cephalotes (O liv .)
Ectemnius continuus (F.)
Ectemnius lapidarius (P a n z .)
66 Ectemnius mjicornis ( Z e t t .)
67 Ectemnius sexcinctus (F.)
68 Lestica clypeata ( S c h r e b . )
69 Lestica subterranea (F.)
70 Mellinus amensis (L.)
71 Nysson niger C h e v r .
72 Nysson spinosus (J . F o r s t . )
73 Nysson trimaculatus (Rossi)
74 Alyssonfuscatus (P a n z .)
75 Argogorytes mystaceas (L.)
76 Cerceris arenaria (L.)
77 Cerceris quadrifasciata (P a n z .)
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COCCINELLIDAE
Scymnus haemorrhoidalis H e r b s t
Scymnus suturalis T h u n b .
Scymnus nigrinus K u g e l .
Ebcochomus quadripustnlatus (L.)
Aphidecta obliterata (L.)
Adalia conglomerata (L.)
Adalia decempunctata (L.)
Coccinella septempunctat (L.)
Harmonia quadripunctata ( P o n t.)
Myrrha octodecimpunctata (L.)
Calvia decemguttata (L.)
Calvia quatuordecimpunctata (L.)
Mysia oblogoguttata (L.)
Anatis ocellata (L.)
CANTHARIDAE
1 Cantharis decipiens B a u d i
2 Cantharis jlavilabris F a l l .
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Cantharis livida mfipes H e r b s t
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Cantharis nigricans (O . F . M u l l .)
Cantharis obscura L.
Cantharis pellucida F.
Cantharis mfa L.
Cantharis rustica F a l l .
Rhagonycha atra (L.)
Rhagonycha elongata ( F a l l .)
Rhagonychafulva (S c o p .)
Rhagonycha lignosa (O.F. M u l l .)
Rhagonycha limbata T h o m s .
Silis nitidula (F.)
Malthinus biguttatus (L.)
Malthinus facialis T h o m s .
Malthinus frontalis ( M a r s h .)
Malthinus punctatus ( F o u r c r . )
Malthodes breuicollis (P ayk.)
Malthodes crassicornis (M a e k l.)
Malthodes guttifer K ie s e n w .
Malthodes mysticus K ie s e n w .
Malthodes pumilus ( B r e b .)
SYRPHIDAE
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(* g a tu n k i zoofagiczne)
Eristalis arbustomm (L.)
Eristalis tenax (L.)
Eristalis mpium F.
Eristalis nemomm (L.)
Eristalis horticola (D e c .)
Eristalinus sepulchralis (L.)
Myathropajlorea (L.)
Helophilus pendulus (L.)
Helophilus hybridus L o e w
Helophilus parallelus H a r r .
Helophilus affinis W h a l b .
Helophilus lineatus (F.)
Parhelophilus versicolor (F.)
Helophilus lunulatus M e ig .
Parhelophilus Jmtetomm (F.)
Lathyrophtalmus aeneus (S c o p .)
Eumems strigatus ( F a l l .)
Temnostoma bombylans (F.)
Temnostoma vespiforme (L.)
Temnostoma apiforme (F.)
Xylota segnis (L.)
Xylota tarda M e ig .
Chalcosyrphus piger (F.)
Xylotafemorata (L.)
Xylota sylvamm (L.)
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Xylota abiens M e ig .
Xylotaforum (F.)
Chalcosyrphus nemorum (F.)
Xylota ignava (P a n z .)
Syritta pipiens (L.)
Lejota nificornis ( Z e t t .)
Triglyphus primus L o e w *
Pipizella varipes (M e ig .) *
Neocnemodonfulvimanus ( Z e t t .) *
Neocnemodon uitripennis (M e ig .) *
Pipiza quadrimaculata (P a n z .) *
Cheilosia pagana (M e ig .)
Cheiosia ruralis (M e ig .)
Cheilosia vernalis ( F a l l .)
Cheilosia chloris (M e ig .)
Orthoneura nobilis ( F a l l .)
Orthoneura geniculata M e ig .
Orthoneura intermedia L u n d b .
Orthoneurafrontalis L o e w
Chrysogaster viduata (L.)
Didea intermedia L o e w *
DideaJasciata M a c q . *
Didea alneti ( F a l l .) *
Scaeua pyrastri (L.) *
Scaeua selenitica (M e ig .) *
Dasysyrphus albostriatus ( F a l l .) *
Dasysyrphus venustus (M e ig .) *
Syrphus torvus O.-S. *
Episyrphus balteatus (D e g .) *
Metasyrphus corollae (F.) *
Syrphus vitripennis M e ig . *
Syrphus ribesii (L.) *
Parasyrphus lineolus ( Z e t t .) *
Meligramma cincta ( F a l l .) *
Meliscaeva cinctella ( Z e t t .) *
Metasyrphus latifasciatus ( M a c q .) *
Meligramma guttata ( F a l l .) *
Dasysyrphus tricinctus ( F a l l .) *
Epistrophe eligans H a r r . *
Melanostoma mellinum (L.) *
Melanostoma scalare (F.) *
Sphaerophoria scripta (L.) *
Sphaerophoria menthastri (L.) *
Sphaerophoria philanthus M e ig . *
Xanthogramma citrofasciatum (D e g .)
Platycheirus albimanus (F.) *
Platycheirus peltatus (M e ig .) *
Platycheirus clypeatus (M e ig .) *
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Xanthandrus comtus ( H a r r . ) *
Melangyna quadrimaculata ( V e r r . )
Baccha elongata (F.) *
Baccha obscuripennis M e ig . *
Neoascia dispar (M e ig .)
Neoascia aenea (M e ig .)
NeoasciaJloralis (M e ig .)
Neoascia geniculata (M e ig .)
Chrysotoxum vernale L o e w *
Chrysotoxumfestiuum (L.) *
Chrysotoxum elegans L o e w *
Chrysotoxum octomaculatum C u r t .
Brachyopa parizeń G o f f e
Brachyopa scutellaris R o b . - D e s v .
Microdon devius (L.)
BleraJallax (L.)
Sericomyia lappona (L.)
Sericomyia silentis H a r r .
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NEUROPTEROIDEA
Raphidia ophiopsis ophiopsis L.
Raphidia notata F.
Raphidia xanthostigma S c h u m m .
Inocellia crassicornis (S c h u m m .)
Coniopteryx parthenia (N av. et M a r c ,
Conventzia pineticola E n d .
Wesmaelius concinnus ( S te p h .)
Wesmaelius nervosus (F.)
Wesmaelius mortoni (Me L a c h l.)
Hemerobius humulinus L.
Hemerobius stigma S t e p h .
Hemerobius pini S t e p h .
Hemerobius Jenestratus T j e d .
Hemerobius nitidulus F.
Hemerobius micans O liv .
Sympherobius fuscescens ( W a ll.)
Nineta vittata (W e sm .)
Chrysopa perla (L.)
Chrysopa septempunctata W e s m .
Mallada prasinus (B u rm .)
Mallada ventralis ( C u r t . )
Chrysoperla carnea ( S te p h .)
Cunctochrysa albplineata (K ill.)
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TACHINIDAE
Acemyia ruftibia (V o n R o s e r )
Actia crassicornis (M e ig .)
Actia nudibasis (S te in )
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Actia pilipennis ( F a l l .)
Admontia blanda ( F a l l .)
Admontia podomyia (B. & B.)
Aplomyia conjinis ( F a l l .)
Appendicia truncata ( Z e t t .)
Athrycia curvinervis ( Z e t t .)
Athrycia trepida (M e ig .)
Bactromyia aurulenta (M e ig .)
Bessa parallela (M e ig .)
Bessa selecta (M e ig .)
Blepharomyia angustifrons H e r t .
Blepharomyia pagana (M e ig .)
Blepharomyia piliceps ( Z e t t .)
Blondelia nigripes ( F a l l .)
Bothria subalpina V i l l .
Brachychaeta stigata (H e ig .)
Campylochaeta inepta (M e ig .)
Carcelia (C.) bombylans R.-D.
Carcelia (C .) gnaua (M e ig .)
Carcelia (C .) rasa (M a c q .)
Carcelia (Euryclea) tibialis (R.-D.)
Caracelina stackelbergi (M e s n .)
Ceranthia abdominalis (R.-D.)
Ceranthia lichtwardtiana (V ill.)
Ceranthia pallida H e r t .
Ceromyia monstrosicornis (S te in )
Ceromyia silacea (M e ig .)
Cylindromyia brassicaria ( F a b r .)
Cylindromyia pilipes (L o ew )
Cyzenis albicans ( F a l l .)
Cyzenis jucunda (M e ig .)
Chaetoptilia puella (R o n d .)
Compsilura concinnata (M e ig .)
Dexiosoma canina ( F a b r .)
Dineraferina ( F a l l .)
Drino bohemica M e s n .
Drino inconspicua (M e ig .)
Elfia cingulata (R.-D.)
Eljia nigroaenea H e r t .
Eloceria delecta (M e ig .)
Entomophaga nigrohalterata ( V ill.)
Eriothrix accolus K o l.
Eriothrix monochaeta W a in w r .
Eriothrix rufomaculata (D e G e e r )
Ernestia rudis ( F a l l .)
Ernestia vagans (M e ig .)
Erycillaferruginea (M e ig .)
Eurithia connivens ( Z e t t .)
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Eurithia consobrina (M e ig .)
Eurithiajuncta (Zim.)
Eumea linnearicornis ( Z e t t .)
Eurysthaea scutellaris (R.-D.)
HebiaJlavicornis R.-D.
Hemyda vittata (M e ig .)
Huebneria ajjinis ( F a l l .)
Linnaemyia tessellans (R.-D.)
Loewiafoeda (M e ig .)
Loewia phaeoptera (M e ig .)
Lydina aenea (M e ig .)
Lydella stabulans (M e ig .)
Lypha dubia ( F a l l .)
Macquartia grisea ( F a l l .)
Macquartia tenebricosa (M e ig .)
Masicera silvatica ( F a l l .)
Medina collaris ( F a l l .)
Medina luctuosa (M e ig .)
Medina melania (M e ig .)
Medina multispina H e r t .
Medina separata (M e ig .)
Meigenia incana ( F a l l .)
Myxexoristops blondelli (R.-D.)
Ocytata pallipes ( F a l l .)
Oswaldia eggeri (B. & B.)
Oswaldia muscaria ( F a l l .)
Parasetigena silvestris (R.-D.)
Parathryphera barbatula (R o n d .)
Peletachina tibialis ( F a l l .)
PeribaeaJissicornis ( S t r o b l )
Peribaea apicalis R.-D.
Phorinia aurifrons R.-D.
Phryxe erythrostoma ( H a r t . )
Phryxe magnicornis ( Z e t t .)
Phryxe nemea (M e ig .)
PlatymyiaJimbriata (M e ig .)
Platymyia mitis (M e ig .)
Prosopea nigricans ( E g g .)
Pseudopachystylum gonioides ( Z e t t .)
Pseudoperichaeta nigrolineata (W a lk .)
Pales pumicata (M e ig .)
Phebellia glauca (M e ig .)
Phorocera obscura ( F a l l .)
Rondaniafasciata ( M a c q .)
Senometopia excisa ( F a l l .)
SiphonaJlavifrons S t a e g .
Siphona confusa M e s n .
Siphona cristata ( F a b r .)
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Siphona rossica M e s n .
102 Siphona setosa M e s n .
1 0 3 Smidtia conspersa (M e ig .)
1 0 4 Tachina magnicornis ( Z e t t .)
1 0 5 Thelaira nigripes ( F a b r .)
1 0 6 Timavia amoena (M e ig .)
1 0 7 Tdarthria setipennis (F a l l .)
1 0 8 Vibdssina turrita (M e ig .)
1 0 9 V b ria ruralis ( F a l l .)
110 Winthemia cruentata (R o n d .)
111 Winthemia variegata (M e ig .)
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PIPUNCUUDAE
1 Nephrocerus scutellatus (M a c q .)
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Chalarus fimbdatus C o e
Verrallia villosa (V. R o s e r )
Eudorylas subjascipes C o l l .
Tomosvariella sylvatica (Mg.)
Pipunculus zugmmeyede Kow.
Pipnnculus spinipes Mg.
Cephalops semifumosus (Kow.)
Cephalops aeneus F a l l .
ICHNEUMONIDAE
Protichneumonini
1 Coelichneumon deliratodus (L.)
2 Coelichneumon desinatodus (T h u m b .)
3 CoelichneumonJcdsificus (W e sm .)
4 Coelichneumonfasciatus (G m e l.)
5 Coelichneumon sugillatodus (L.)
Heresiarchini
6 Heresiarches eudoxius (W e s m .)
Listrodonini
7 Anisobas platystylus (T h o m s.)
Acanthojoppini
8 Pseudoplatybalus uniguttatus (G r a v .)
9 Pseudoplatybalus violentus (G ra v .)
Ichneumonini
10 Stenichneumon culpator ( S c h r a n k )
11 Syspasis alboguttatus ( G ra v .)
12 Syspasis eburnifrons (W e sm .)
1 3 Syspasis scutellador (G ra v .)
1 4 Stenoaopius pictus (G ra v .)
1 5 Aoplus defraudator (W esm .)
1 6 Aoplus ochropis (G m e l.)
1 7 Platylabops virginalis (W esm .)
18 Stenobarichneumon bascdis ( P e r k .)
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Lymantdchneumon dispar (P o d a )
Cratichneumon culex ( M u ll.)
CratichneumonJ'abdcator (F.)
Cratichneumon palliditarsus (T h o m s.)
Cratichneumon punctijrons (G ra v .)
Cratichneumon rujifrons (G ra v .)
Cratichneumon sicadus (G ra v .)
Cratichneumon versator (T h u m b .)
Cratichneumon viator (S c o p .)
Eupalamus lacteator (G ra v .)
Eupalamus oscillator (W esm .)
Eupalamus wesmaeli (T h o m s.)
Cryptejfigies lanius (G ra v .)
Homotherus locutor (T h u m b .)
Homotherus varipes (G ra v .)
Edsticus clarigator (W esm .)
Vulgichneumon bimaculatus ( S c h r a n k . )
Vulgichneumon deceptor (S c o p .)
Vulgichneumon saturatodus (L.)
Virgichneumon ? albosignatus (G r a v .)
Virgichneumon diagrammus ( G ra v .)
Virgichneumon dumeticola (G ra v .)
Virgichneumon extremator (T h u m b .)
Virgichneumonfaunus (G ra v .)
Virgichneumon maculicauda (P e r k .)
Virgichneumon monostagon (G ra v .)
Badchneumon praeceptor (T h u m b .)
Badchneumon ? “perversus K r j e c h b . ”
Badclmeumon ? plagiadus (W esm .)
Chasmias motatodus (F.)
Ichneumon albicollis W e s m .
Ichneumon cessator M u l l .
Ichneumon extensodus L.
Ichneumon grasilentus W e s m .
Ichneumon insidiosus W e s m .
Ichneumon latrator F.
Ichneumon lugens G r a v .
Ichneumon melanotis H o m g r .
Ichneumon memorator W e s m .
Ichneumon minutodus D e s v .
Ichneumon nereni T h o m s .
Ichneumon sarcitorius L.
Ichneumon subquadratus T h o m s .
Limerodops elongatus ( B r is c h k e )
CtenichneumonJunereus ( G e o f f r . )
Ctenichneumon divisodus (G ra v .)
Diphyus amatodus ( M u ll.)
Diphyus luctatodus (L.)
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Spilothyrateles punctiis (G ra v .)
Spilichneumon ammonius (G ra v .)
Platylabini
69 Platylabus nigrocyaneus ( G ra v .)
70 Poecilosticus cothurnatus ( G ra v .)
71 Cyclolabus nigricollis (W e sm .)
72 Lincyus exhortator (F.)
Phaeogenini
73 Eriplatys ardeicollis (W e sm .)
74 Eriplatys sawoniewiczi ( D i l l e r )
75 Hemichneumon elongatus (R a tz .)
76 Trachyarus corvinus T h o m s .
77 Stenodontus marginellus (G ra v .)
78 Dicaelotus crassifemur T h o m s .
79 Dicaelotus inflexus T h o m s .
80 Dicaelotus pumilus (G ra v .)
81 Dicaelotus rujilimbatus (G ra v .)
82 Deloglyptus pictus S c h m ie d .
83 Diadromus subtilicornis (G r a v .)
84 Diadromus troglodytes (G ra v .)
85 Colpognathus celerator (G ra v .)
86 Aetheurus discolor W e s m .
87 Aetheurus nitidus W esm .
88 Phaeogenes melanogonos (G m e l.)
89 Phaeogenes spiniger (G ra v .)
90 Dirophanes callopus (W esm .)
91 Dirophanes maculicornis ( S te p h .)
92 Dirophanes fulvitarsis (W e sm .)
93 Tycherus amoenus (W e sm .)
94 Tycherus clypearis ( B r is c h k e )
95 Tycherus coriaceus (P e r k .)
96 Tycherus dodecellae R a n in
97 Tycherus cf. jucundus (W e sm .)
98 Tycherus ophtalmicus (W esm .)
99 Tycherus osculator (T h u m b .)
00 Tycherus vagus ( B e r t h . )
01 Tycherus verecundus R a n in
02 Epitomus infuscatus (G ra v .)
03 Misetus oculatus (W e sm .)
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1 Mesenchytraeus pelicensis I s s e l .
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Mesenchytraeus s p . Eis.
Buchholiza simplex N ie l. e t C h r i s t .
Buchholiza s p . M ic h .
Buchholiza ehlersi U d e
Cognettia sphagnetorum (V e jd .)
Marionina s p . M ic h .
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Fridericia ratzeli Eis.
Fridericia sp. M ic h .
Achaeta sp. V e j d .
COLLEMBOLA
Hypogastruridae
1 Schoetella ununguiculata ( T u llb .)
2 Xenylla brevicauda T u l l b .
3 Xenylla boerneri A x e l .
4 Willemia anophthalma B o r n e r
5 Willemia aspinata S t a c h
6 Hypogastrura sp. j.
Neanuridae
7 Pseudachorutes parvulus B o r n e r
8 Microanurida pygmaea B o r n e r
9 Neanura muscorum (T e m p l.)
Onychiuridae
1 0 Oligapphorura absoloni ( B o r n e r )
11 Propaphorura armata ( T u l l b .)
1 2 Propaphorura subuliginatus G is in
1 3 Propaphorurafurcifera ( B o r n e r )
1 4 Mesaphorura macrochaeta R u s e k
1 5 Mesaphorura yossi R u s e k
Isotomidae
1 6 Anurophorus laricis S t a c h
1 7 Anurophorus septentrionalis P a l i s s a
1 8 Folsomia quadrioculata ( T u llb .)
1 9 Proisotoma minima (A b s.)
2 0 Isotomiella minor ( S c h a f f .)
2 1 Isotoma notabilis S c h a f f .
2 2 Isotoma hiemalis S c h o t t
2 3 Isotoma tigrina (Nic.)
2 4 Isotoma sp. j.
2 5 Vertagopus cinereus (Nic.)
2 6 Pachyotoma sp .n .
Entomobryidae
T7 OrchesellaJlavescens ( B o u r l . )
2 8 Entomobrya nivalis (L.)
2 9 Entomobrya sp. j.
3 0 Lepidocyrtus cyaneus T u l l b .
3 1 Lepidocyrtus lignorum ( F a b r .)
3 2 Lepidocyrtus sp. j.
3 3 Pseudosinella zygophora ( S c h i l l e )
3 4 Cyphoderus albinus Nic.
3 5 Pogonognathellus Jlavescens ( T u l l b .)
3 6 Tomocerus sp. j
Sminthuridae
3 7 Lipothrix lubbocki ( T u llb .)
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Allacmafusca (L.)
Bourletiellidae
Bourletiella hortensis (F itc h )
NEMATODA
1 Rhabditis s.
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L.
Bunonema reticulatum R i c h t .
Heterocephalobus elongatus (D e M an )
Acrobeloides nanus (D e M an )
Ypsylonellus insubricus ( S te i n .)
Teratocephalus dadayi A n d r .
Teratocephalus tenuis A n d r .
Metateratocephalus crassidens (D e M an )
Filenchus ditissimus ( B rz .)
Malenchus bryophilus ( S te in .)
Ditylenchus anchilisposomus T a r i a n
Ditylenchus equalis H e i n s
Ditylenchus longimatricalia (K az.)
Geocenamus longus (W u)
Helicotylenchus pseudorobustus S t e i n .
Paratylenchus nanus C o b b .
Aphelenchoides spp.
Seinura winchesi G o o d e y
Paraphelenchus sp.
Eymonhystera sp.
Geomonchystera sp.
Plectus geophilus D e M a n
Plectus longicaudatus (B u t.)
Plectus pusteri F u c h s
Wilsonema otophorum (D e M an )
Prodesmodora terricola M i c o l e t z k y
Alaimus mucronatus A l t h .
Prionchulus punctatus (C o b b .)
Mesodorylaimus sp.
Eudorylaimus siluaticus B r z .
Aporcelaimellus sp.
Tylencholaimus mirabilis B u t .
Tylencholaimus stecki S t e i n
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THEREVIDAE
Acrosathe annulata (F.)
Cliorismia ardea (F.)
Dialineura anilis (L.)
Spiriverpa lunulata ( Z e t t .)
Thereva cinifera (M e jg .)
TherevaJ'ulva (M e ig .)
Thereva circumscripta Lw.
Thereva handlirschi K r o e b e r
Thereva lanata Z e t t .
Thereva microcephala Lw .
Thereva inornata V e r r .
Thereva nobilitata (F.)
Thereva strigata (F.)
Thereva plebeja (L.)
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RHAGIONIDAE
Rlw.gio lineola (F.)
Rhagio scolopaceus (L.)
Rhagio tringarius (L.)
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FORMICIDAE
1 Myrmica ruginodis N y l.
2 Myrmica scabrinodis N y l.
3 Myrmica sabuleti M e in .
4 Myrmica lobicornis N y l.
5 Myrmica schencki Em.
6 Leptothorax acervorum ( F a b r .)
7 Leptothorax muscorum (N y l.)
8 Tetramorium caespitum (L.)
Strongylognathus testaceus ( S c h e n c k )
Camponotus herculeanus (L.)
Lasius niger (L.)
Lasius Jlavus (F a b r .)
Lasius umbratus umbratus (N y l.)
Lasius fuliginosus ( L a t r .)
Formicafusca L.
Formica truncorum F.
Formica rufa L.
Formica polyctena F o e r s t .
Formica sanguinea ( L a t r .)
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MICROMAMMALIA
Rodentia
Arvicolidae
Clethrionomys glareolus ( S c h r e b . )
Pitymys subterraneus (S.-L.)
Microtus agrestis (L.)
Microtus arval is ( P a l l a s )
Microtus oeconomus ( P a l l a s )
Muridae
Micromys minutus ( P a l l a s )
Apodemus agrarius ( P a l l a s )
Apodemus flavicollis ( M e lc h .)
Apodemus sylvaticus (L.)
Zapodidae
Sicista betulina ( P a l l a s )
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Insectivora
Soricidae
Sorex araneus (L.)
Sorex caecutiens L a x m a n n
Sorex minutus (L.)
Neomys anomalus ( C a b r .)
Neomys Jodiens ( S c h r e b . )
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12
13
14
AVES
1
2
3
4
5
6
7
8
Passeriformes
Alaudidae
Lullula arborea (L.)
Motacillidae
Anthus trivialis (L.)
Troglodytidae
Troglodytes troglodytes (L.)
Prunellidae
Prunella modularis (L.)
Turdidae
Erithacus rubecula (L.)
Phoenicurus phoenicurus (L.)
Turdus merula (L.)
Turdus philomelos (C . L. Brehm)
15
16
17
18
19
20
21
22
23
Sylvidae
Sylvia atricapilla (L.)
Sylvia borin (B o d d .)
Phylloscopus collybita (Vieill.)
Phylloscopus sibilatrix ( B e c h s t .
Phylloscopus trochilus (L.)
Regulus regulus (L.)
Aegithalidae
Aegithalos caudatus (L.)
Paridae
Parus ater (L.)
Parus cristatus (L.)
Certhiidae
CerthiaJamiliaris (L.)
Fringillidae
Fringilla coelebs (L.)
Carduelis spinus (L.)
Pyrrhula pyrrhula (L.)
Emberizidae
Emberiza citrinella (L.)
Picidae
Dendrocopos major (L.) ,
STRESZCZENIE
[Tytuł: S u k c e sja w tó rn a fauny w b o ra c h świeżych Puszczy Białowieskiej]
Bory sosnow e Polski ze względu n a w artości użytkow e drew n a sosnow ego
e ksploatow ane są i u p raw ia n e od se te k lat. Na w iększość z n ich sk ła d a ją się
d rz e w o s ta n y je d n o w ie k o w e , p o w s ta łe n a jc z ę śc ie j w w y n ik u s z tu c z n e g o
odnow ienia zrębów zupełnych. Założenie n a porębie upraw y sosny, zwykle z
dom ieszką drzew tow arzyszących b ąd ź osłonow ych, u ru c h a m ia p ro ces su k cesji
w tórnej. W jej trak cie b ó r przechodzi przez kilka etapów : upraw y, m łodnika,
tyczkow iny, drągow iny, drzew o stan u dojrzew ającego aż do s ta d iu m dojrzałości.
Proces ten, w p rzypadku borów sosnow ych świeżych, trw a w naszy ch w a ru n k a c h
10-150 lat ( S z u j e c k i 1980).
Każdy z tych etapów c h a ra k te ry z u je się nieco odm iennym i w a ru n k a m i
m ik ro k lim atu leśnego, zależnie od s k ła d u d rzew ostanu i w łaściw ości siedliska
( O b m iń s k i 1978). Zm iany te z n a jd u ją swe odzw ierciedlenie w przebudow ie szaty
roślinnej. Tu procesy su k cesji w tórnej w b o rach świeżych Puszczy Białowieskiej
m a ją c h a r a k te r w y b itn ie k rea ty w n y . P row adzą bow iem do u fo rm o w a n ia
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s tru k tu ry przestrzennej b oru o w yraźnej piętrow ości, a przestrzeń życiowa
w n ętrza la su je s t zapełniona przez g a tu n k i dopasow ane do uform ow anych w
to k u su k c esji w tórnej w aru n k ó w środow iska. J e s t to proces pow tarzalny,
obserw ow any i opisany dla siedlisk borow ych z różnych regionów k raju .
Procesy te w odniesieniu do fauny rozpoznane są fragm entarycznie, a
d o k u m e n ta c ja su k c esji w tórnej fau n y obejm uje tylko poszczególne g rupy
zw ierząt i nie pozw ala n a stw orzenie pełnego obrazu przem ian zachodzących w
składzie i organizacji kom pleksów faunistycznych. Na tle dość bogatego, lecz
c z ę s to s ła b o p o w ią z a n e g o z r e a lia m i p r z y ro d n ic z y m i, p iś m ie n n ic tw a
pośw ięconego tem atyce sukcesji ekologicznej autorzy niniejszego opracow ania
p o s ta w il i s z e r e g p y t a ń d o s to s o w a n y c h do c h a r a k t e r u m a t e r i a łó w
faunistycznych, więc takich, które d a ją m ożliwość u z y sk an ia u d o k u m en to w an ej
odpowiedzi.
Podstaw ow ą w naszym rozum ieniu te m a tu kw estią je s t c h a ra k te r su k cesji
w tórnej fauny, czyli ja k i do czego ona prow adzi. Nie je s t bowiem oczywiste, ta k
ja k w p rzy p ad k u szaty roślinnej boru, że w procesie sukcesji w tórnej m a m iejsce
w yraźny rozwój biom asy zw ierząt i uform ow anie się s tr u k tu r przestrzen n y ch .
Z jaw iska takie, jeśli n aw et m ają m iejsce w ym agają u d o k u m en to w an ia. Na czoło
w ysuw a się kw estia jednotorow ości czy w ielotorowości p rocesu su k cesji w tórnej
fauny. Chodziło tu o sa m m odel procesu. Czy je s t on w pełni kreatyw ny, a jego
etapy polegają n a zasied lan iu p u stk i ekologicznej, form ow aniu podsystem ów i
d o p a so w y w a n iu ich do ekologicznej s tr u k tu r y dojrzew ającego la s u . T en
klarow ny obraz budzi szereg w ątpliw ości. Czy istnieje bowiem przez ja k iś o kres
p u s tk a fa u n is ty c z n a po w ycięciu la s u , czy też o tw a rta p rz e s trz e ń j e s t
n a ty c h m ia st z asied lan a? Czy poszczególne podsystem y zoocenozy dopiero się
form ują, czy są ju ż gotowe w przyrodzie? I w końcu, n a czym m a polegać owo
dopasow yw anie, lub dokładniej, do czego w rzeczyw istości d o p aso w u ją się
poszczególne grupy fauny?
Dla u z y sk an ia jednoznaczej odpowiedzi n a postaw ione p y tan ia zbudow ano
dwie hipotezy robocze, które stanow iły n a stę p n ie podstaw ę do w eryfikacji w
op arciu o zeb ran e i opracow ane m ateriały.
H ipoteza pierw sza określa, że su k c e sja w tó rn a fauny może m ieć rozm aity
przebieg i prow adzić do odm iennych niekiedy układów końcow ych. O pisano pięć
typów przebiegu sukcesji: kreatyw ny, stabilizujący, załam any, regresyw ny i
re sta u ra c y jn y (Fig. 3).
H ipoteza d ru g a o k reśla pow iązania fau n y w system zależności troficznych o
c h a ra k te rz e hierarchicznym . Zgodnie z n ią zrąb s tru k tu ry fauny stan o w ią
pow iązane zależnościam i pokarm ow ym i u k ład y takie ja k podsystem , ła ń c u c h
pokarm ow y, zespół k o n k u ren cy jn y i p o pulacja określonego g a tu n k u zw ierząt.
Zależności te przedstaw iono w o d niesieniu do 30 g ru p fau n y objętej pro g ram em
b a d a ń d la s ta d iu m końcow ego su k c esji w tórnej, fau n y sta ro d rz e w u b o ru
sosnow ego świeżego.
T ak sform ułow ane hipotezy robocze pozwoliły n a konfrontację d a n y c h o
faunie z w yobrażeniam i o przebiegu su k cesji w tórnej. J a k o m a te ria łu do
weryfikacji hipotez użyto m ateriałów do k u m en tacy jn y ch d la poszczególnych k las
w ieku lasu : 1) liczbę g a tu n k ó w tw orzących poszczególne tak so cen y lu b zespoły
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P. T rojan et al.
k onkurencyjne; 2) liczebność tych jed n o ste k , zwykle ujm ow aną wskaźnikow o
dla zobrazow ania tendencji zm ian liczebności; 3) s tru k tu rę taksocenów dla
zobrazow ania zm ian w organizacji układów zoocenotycznych; 4) różnorodność
g atu n k o w ą a k tu a ln ą i poten cjaln ą oraz zachodzący między nim i sto su n ek
m ierzony w skaźnikiem Pielou.
Liczba g atu n k ó w (Tab. II) w to k u sukcesji w tórnej borów sosnow ych świeżych
w Puszczy Białowieskiej w ykazuje zm iany idące w sześciu k ieru n k a ch . Dom .nuje
typ su k cesji restau racy jn ej c h a rak tery zu jący się w ysoką liczbą gatunków w
u p raw a ch sosny, spadkiem w fazie drągow iny i odbudow ą w starodrzew ie Ten
typ przebiegu został stw ierdzoy w połowie zbad an y ch zespołów konkurencyjnych
i 8 tak so c e n ac h . Na drugim pod względem u d ziału m iejscu znalazły się dw a
dalsze typy przebiegu su k cesji w tórnej. S u k c e sja regresyw na, polegająca n a
sp a d k u liczby g atu n k ó w w całym szeregu sukcesji w ystępuje w obrębie 4
zespołów i 5 taksocenów zwierząt. Podobnie su k c e sja załam ana, w której po
początkow ym w zroście m a m iejsce sp a d ek liczby g a tu n k ó w została stw ierdzona
w 5 zespołach k o n k urencyjnych. S u k cesja k reaty w n a ch arak tery zu jąca się
stałym w zrostem liczby g atu n k ó w w szeregu sukcesyjnym w ystąpiła ty k o w
o b rę b ie 4 ze sp o łó w k o n k u re n c y jn y c h . S ta b iliz a c ja liczby g a tu n k ó w po
początkow ym ich w zroście m iała m iejsce tylko w 2 zespołach zwierząt. Poza tymi
5 typam i zm ian liczby g a tu n k ó w należy wym ienić 5 przypadków b ra k u zm ian
liczby g a tu n k ó w w to k u procesów rozwoju boru. Taki s ta n rzeczy stw ierdzono
dla stonkow atych (C hrysom elidae), afidofagicznych biedronek [Coccinelliiae),
pasożytniczych m uchów ek Pipunculidae, n icieni (Nematoda) i R hagioridae
(Diptera).
Podobny obraz zróżnicow ania reakcji poszczególnych gru p zwierząt n a proces
odbudow y b o ru świeżego m a m iejsce przy ro zp a try w a n iu poszczególnych
zespołów ko n k u ren cy jn y ch (Tab. II). Tam , gdzie m am y do czynienia z kilkom a
tak so n a m i w chodzącym i w skład zespołu, zazwyczaj każdy z nich prezentuje
odm ienny k ieru n e k zm ian liczby gatunków . W yjątek pod tym względem stanow ią
zoofagi oraz ksylo- i kam biofagi, w śród których dom inuje resta u ra c y jn y kierunek
zm ian liczby gatunków . N atom iast w śród fitofagów zgryzających przew ażają
ten d en cje spadkow e w odniesieniu do liczby g a tu n k ó w w toku su kcesji wtórnej.
W ynik analizy zm ian w tworzywie organizacji ekosystem u leśnego, ja k i
stanow ią g a tu n k i zw ierząt, w skazuje n a to, że procesy twórcze rozbudowy lub
o d b u d o w y u k ła d ó w , r e p r e z e n to w a n e p rz e z ty p s u k c e s ji k re a ty w n e j,
s ta b iliz u ją c e j i re s ta u ra c y jn e j s ą rów nie siln e j a k p ro cesy d e stru k c y jn e
w yrażające się w ostatecznym ra c h u n k u spadkiem liczby gatunków . Z nam ienne
je s t przy tym , że co 6 g ru p a zwierząt nie reaguje z m ian ą liczby gatunków n a
procesy zachodzące w rozw ijającym się borze. Liczebność zgrupow ań zwierząt
(Tab. III) przed staw ia rów nie zróżnicow any obraz przebiegu zm ian ja k liczba
g atunków , je d n a k dw a kieru n k i procesów są dom inujące: regresja i restau racja.
O koło 1 /3 z g ru p o w a ń zw ierząt s ta r tu je do p ro c e s u su k c e sji z w iększą
liczebnością populacji niż w drzew o stan ach dojrzałych, dom inuje więc spadek
liczebności. Na drugim m iejscu zn a jd u ją się te zgrupow ania, których spadek
zostaje w to k u su k cesji zatrzym any i w d rzew o stan ach starszy ch m a miejsce
w zrost liczebności populacji. Tylko co 7 tak so cen w ykazuje w zrost liczebności
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S econdary su c c e ssio n of fa u n a in th e p in e fo rests of P uszcza B iałow ieska
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populacji w całym szeregu su k cesji i ta k a sa m a liczba g ru p nie w ykazuje zm ian
liczebności u trzy m u jąc się w całym szeregu sukcesji n a tym sam ym poziom ie
zagęszczenia. Co 15 tak so cen po początkow ym w zroście liczebności w chodzi w
fazę sp a d k u , zaś tylko 1 tak so cen w śród saprofagów , m ianow icie skoczogonki
{Collembola) m a przebieg liczebności w skazujący n a stabilizację zgrupow ania w
d rze w o sta n a ch starszy ch .
Z ja w is k a te r o z p a t r y w a n e od s t r o n y p o s z c z e g ó ln y c h z e s p o łó w
k o n k u ren cy jn y ch w ykazują w yraźne rozstrzelenie reakcji liczebności w ich
obrębie. W każdym zespole bezkręgowców spotykam y obok siebie tak so cen y
w ykazujące, je d n e w zrost, drugie sp a d ek liczebości. W yjątek pod tym względem
s ta n o w ią niew yspecjalizow ane zoofagi, w śród których d o m in u ją ten d e n c je
sp a d k u liczebności.
A naliza zm ian liczebności fauny pozw ala n a uzy sk an ie odpowiedzi od n o śn ie
dw óch w ażnych kw estii zw iązanych z su k c e sją w tó rn ą borów sosnow ych
św ie ż y c h . P ojęcie “p u s tk i f a u n is ty c z n e j” n ie o d p o w ia d a rz e c z y w isto śc i
przyrodniczej. Poręba leśn a zostaje n a ty c h m ia st zasiedlona przez populacje
zw ierząt o wysokiej liczebności, niem al 2 / 3 zgrupow ań zw ierząt w to k u su k cesji
zan iża sw ą liczebność w s to s u n k u do jej s ta n u w u p raw a ch sosny.
W zrost liczebności w ykazuje tylko 1 /4 zespołów zwierzęcych (Tab. III), s ą to
fitofagi ssące, afidofagi i ptaki. Liczebność stanow i m iarę su k c e su ekologicznego
zarów no g a tu n k u ja k i zgrupow ania zwierzęcego. W toku sukcesji w tórnej bo ru
so s n o w e g o św ie ż e g o m a m ie js c e z a s tę p o w a n ie s y s te m u e k s p lo a ta c ji
produkow anej przez rośliny zielonej m asy. Z jadanie igieł sosny przez fitofagi o
zgryzającym a p a ra c ie gębowym, typowe dla szkodników pierw otnych sosny,
w raz z rozwojem lasu u stę p u je n a rzecz w ysysania p roduktów asym ilacji przez
m niej groźne dla la su fitofagi ssące. Zjaw isko to w ystępuje, w każdym razie w
Puszczy B iałow ieskiej, której bory so sn o w e n ie s ą d o ty k an e klęskow ym i
pojaw am i fitofagów zgryzających. W zrostow i liczebności fitofagów ssący ch
tow arzyszy rozwój zespołów afidofagów reg u lu jący ch ich liczebność. D uże
znaczenie m a w zrost liczebności zespołów ptaków (w w iększości ow adożernych),
któ re stanow ią ogniwo kontrolujące szczególnie podsystem biotrofów.
R óżnorodność gatu n k o w a a k tu a ln a (Tab. IV) o cen ian a m iarą S h a n n o n a i
W eavera (Hj w to k u sukcesji w tórnej b o ru świeżego w ykazuje najbardziej
ró w n o m ie rn y ro z k ła d typów re a k c ji. N ajczęściej m a ją m ie jsc e p ro c e sy
z m n ie jsz a n ia się ró żn o ro d n o śc i w zględem jej s ta n u w yjściow ego, je d n a k
sum arycznie, procesy w zrostu różnorodności są rów nie częste w zgrupow aniach
zwierzęcych ja k procesy jej sp a d k u . Z naczny udział m ają rów nież procesy
załam yw ania się różnorodności po okresie w zrostu. Nieco inaczej przedstaw ia
się ta kw estia o g ląd an a od stro n y w ypełniania możliwości potencjalnych rozwoju
ró ż n o ro d n o śc i g a tu n k o w e j w e k o sy ste m ie . T u p ro c e sy z m n ie js z a n ia się
ró ż n o ro d n o śc i, szczególnie w d rz e w o s ta n a c h d o jrz a ły c h m a ją c h a r a k te r
d o m in u jący , stw ierd zo n o je w obrębie 17 zg ru p o w ań zw ierzęcych n a 28
przeanalizow anych pod tym w zględem . W zrost, odbudow a lub stabilizacja
różnorodności po w zroście dotyczyły tylko 1 /3 obserw ow anych przypadków .
Stabilizacja różnorodności n a tym sam ym poziom ie w całym szeregu su k cesji
należy do wyjątków.
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P. T rojan et al.
W ynik przedstaw ionej analizy w skazuje w yraźnie, że teza Mac A rth u ra
“diversity m ak es stab ility ” nie znajd u je żadnego odbicia w d a n y ch em pirycznych
( M a c A r t h u r 1955). W ręcz p rzeciw nie, sp a d e k ró żn o ro d n o śc i g atu n k o w ej
zgrupow ań zw ierząt w staro d rzew ach b o ru sosnow ego w skazuje n a to, że w zrost
sto p n ia organizacji u k ła d u odbyw a się kosztem zm niejszenia jego różnorodności
w e w n ę trz n e j. M a to ś c is ły z w ią ze k ze z ja w is k a m i s tr u k t u r a l n y m i . W
d rzew o stan ach stary ch , zazwyczaj w każdym zgrupow aniu zwierzęcym je s t je d e n
lub p arę g atu n k ó w w yraźnie dom inujących, “ogon” g atu n k ó w akcesorycznych
ulega zwykle skróceniu. O rganizacja zgrupow ania o p a rta je s t zwykle n a kilku
dobrze dostosow anych g a tu n k a c h b u d u jąc y c h zespoły zwierzęce.
W zakończeniu należy podkreślić, że cechą najbardziej c h a ra k te ry sty c z n ą
su k cesji w tórnej fau n y w b o rac h sosnow ych świeżych Puszczy Białowieskiej je s t
w ielokierunkow ość procesów zachodzących n a tle rosnącego, rozw ijającego sw ą
s tr u k tu rę p rz e strz e n n ą b o ru świeżego. J e śli wziąć pod uw agę przeanalizow ane
param etry: liczbę g atunków , liczebność, różn orodność g atu n k o w ą i s tru k tu rę ,
to dom inujące znaczenie m ają procesy sp a d k u w artości w skaźników , w całym
szeregu su kcesji lub w jej końcowej fazie (44% przypadków ). Stabilizacja tych
w a rto ś c i w całym sz e re g u lu b po o k re s ie w z ro stu o b e jm u je tylko 22%
przypadków . Tylko w 34% przy p ad k ó w obserw ujem y rozwój od p o c z ątk u
su k cesji do sta d iu m dojrzałego lub po okresie sp a d k u w środkow ym sta d iu m .
D ane te potw ierdzają hipotezę o w ielokierunkow ości przebiegu procesów
su k cesji w tórnej w obrębie zgrupow ań fauny. Szczególne znaczenie m a sp a d ek
w arto ści w ielu w skaźników c h a ra k te ry z u ją c y ch fau n ę w końcow ym etapie
sukcesji, który m ożna określić ja k o fazę organizacji zgrupow ań, k tó ra odbyw a
się kosztem redukcji liczby g a tu n k ó w i różnorodności n a rzecz u stab ilizo w an ia
s t r u k t u r d o m in acy jn y ch . T a k więc sta b iln o ś ć ekosystem ów tw orzą lepiej
przystosow ani, a różnorodność je s t jej b u d u lcem nie w ykorzystanym w pełni.
Cały u k ład zoocenotyczny pow staje rów nocześnie ju ż w fazie n a s a d z a n ia
d rzew o stan u po wyrębie i tu je s t najbogatszy. Dalej toczy sie jed y n ie przebudow a
jego organizacji w raz z rozwojem bo ru ja k o ekosystem u.
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TABLE OF CONTENTS
1. In tro d u c tio n ...............................................................................................................................................
1.1. S tu d y objectives ..............................................................................................................................
1.2. Review of l i t e r a t u r e ........................................................................................................................
3
3
6
2. A rea of s t u d y ...........................................................................................................................................
7
3. F a u n a sam p lin g m eth o d s a n d m a t e r i a l ..........................................................................................
9
4. M ethods for processing d a ta
.............................................................................................................
11
5. A nalysis of seco n d ary su c c essio n of th e p ine fo rests of P uszcza B i a ł o w i e s k a ...................
5.1. A m odel of th e s tru c tu re of p ine forest zoocenosis a n d th e c o u rse
of seco n d ary su ccessio n .............................................................................................................
5.2. S u ccessio n in th e su b sy ste m of b i o t r o p h s ...........................................................................
5.2.1. P h y to p h a g e s ........................................................................................................................
5.2.2. E piphytic p r e d a t o r s .........................................................................................................
5.2.3. P arasito id s of p h y to p h a g e s ..............................................................................................
5.3. S econdary su ccessio n in th e sa p ro tro p h ic su b sy ste m .....................................................
5.3 .1. Soil s a p r o p h a g e s .................................................................................................................
5.3.2. P an to p h ag es a n d soil p r e d a t o r s ...................................................................................
5.4. V e r te b r a te s .......................................................................................................................................
13
13
19
19
36
51
57
57
64
70
6. S u m m a r y ..................................................................................................................................................
77
7. B ibliography
...........................................................................................................................................
82
8. An inventory of species o ccu rrin g in th e pine forests of P uszcza B ia ł o w i e s k a ...................
86
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