Bultman&Uetz_oecologia_1982

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Table 1. Mean taxonomic and structural com position of leaves onthe forest floor of the study site, 6 April 1980. N =1 0 0.1 m 2 quad-ratsSpecies Percent Coefficient ofcomposition variationSpeciesBeech 98 0.02

Other 2 1.23StructureFalt 32 0.22Bent 52 0.06Tw isted 6 0.81Curled 10 0.83

c la s s e s ( s e e C .V . ' s i n Ta b le 1 ) i nd ic a t e s t he h igh un i fo rmi tyo f t h e l e a f l i t te r i n t h i s f o r e s t. N a t u r a l l i tt e r d e p t h w a s4.4 cm__0.21 (Y_+C.V.) .

Experimental Design and SamplingL e a f t r e a tm e n t s w h o s e c o m p o s i t i o n w a s s t r u c t u r a l l y le ssc o m p l e x ( 7 8 % f l a t ; 2 2 % b e n t ) , e q u a l l y c o m p l e x ( 3 2 % f l a t;5 2 % b e n t ; 6 % t w i s t ed ; 1 0 % c u r i e d ) o r m o r e c o m p l e x ( 5 0 %t w i s t e d ; 5 0 % c u r l e d ) t h a n t h o s e a t t h e s t u d y s i t e w e r e a s -s e m b l e d f o r b o t h n a t u r a l a n d a r t if i ci a l n o n - n u t r i t i v e l e av e s.A r t i f i c i a l l e a ve s w e re c o ns t ruc t e d o f v iny l p l a s ti c o f0 . 25 4 m m t h i ck n e s s . E n v i r o n m e n t a l c o n d i t i o n s w i t h i n a r t i-f i c ia l l i t t e r d id no t d i f f e r f ro m tho s e w i th in na tu ra l l e a ve s( B u l t m a n a n d U e t z , i n r e v i e w ) . O d o r s f r o m p l a s t i c s m a ya f f e c t ( a t t r a c t o r r e p e l) l i tt e r -d w e l l in g a r t h r o p o d s ; t h e p a r -t i c u l a r v i n y l u s e d w a s c h o s e n f o r i t s la c k o f o d o r . L a b o r a t o -r y r e a r e d s p i d e r s w e r e n e i t h e r r e p e l l e d b y n o r a t t r a c t e d t oi t (T . B u l t m a n , p e r s o n a l o b s . ). L e a f tr e a t m e n t s w e r e h o u s e di n 2 . 5 4 c m m e s h c h i c k e n - w i r e b a s k e ts w h o s e a r e a w a s0 .1 m 2 a nd w ho s e s ide s w e re e q ua l t o t he s tu dy s i t e me a nl i tt e r d e p t h . T h e 6 tr e a t m e n t s w e r e r e p l i c a t e d 2 4 t i m e s a n dp l a c e d o n t h e f o r e s t f l o o r 1 5 M a y 1 98 0 u si n g a r a n d o m i z e db l o c k d e s i gn . S p i d e r s c o l o n iz i n g b a s k e t s w e r e r e m o v e dm o n t h l y f o r 4 m o n t h s w i t h t h e e x c e p t i o n t h a t s a m p l i n g d i dn o t o c c u r w i t h i n 3 6 h a f t e r a r a i n . A l l o w a n c e f o r t h i s d r y i n gpe r iod w a s ne c e s s a ry be c a us e the s p ide r f a una i s no t i c e a b lyi n a c t i v e f o l l o w i n g a r a i n . S a m p l i n g w a s a c c o m p l i s h e d b ys i e v ing s p ide r s f rom l i t t e r . B u l tma n a nd U e tz ( in r e v ie w )g i v e a m o r e d e t a i le d a c c o u n t o f s a m p l i n g t e c h n i q u es .

Statistical TechniquesThis s tudy w a s de s igne d s pe c i f i c a l ly fo r s t a t i s t i c a l t e s t ingb y a n a l y s is o f v a r i a n c e ( A N O V A ) . P r o b a b i l i t y l ev e l o f s ig -n i f i c a n c e a t t a c h e d t o s t a t i s t i c a l t e s t s t h r o u g h o u t w a sP < 0 . 05 . T o i n s u r e t h a t o u r s a m p l i n g t e c h n i q u e s a d e q u a t e l ys a mple d s p ide r s in the fo re s t l it t e r , w e c ons t ruc t e d s pe c ie s-s a mple s e f fo r t c u rve s . B e c a us e the s e c u rve s a ppe a r e d tor e a c h a n a s y m p t o t e a f t e r s a m p l i n g 2 4 r ep l i c a te b a s k e ts p e rt r e a tme n t pe r da t e ( i . e . , F ig . 1 ) w e c onc lude d ou r s a mpl ingm e t h o d s a d e q u a t e l y s a m p l e d t h e s p i d e r f a u n a .S p i d e r s p e ci e s d iv e r s i ty a n d i ts t w o c o m p o n e n t s o f d i v e r-s i ty w e r e c a lc u l a t e d f o r t h e 6 t r e a t m e n t t y p e s . T h e t w o c o m -pon e n t s o f s pe c ies d ive r s i ty a re s pe c ie s r i c hne s s ( s) a nd e ve n -

3530- -

5 20~6

E 1

1 1 0 0 0 0 0 0

I I I I6 I2 18 2Z~N u m b e r o f s a m p t e s

Fig. 1. Species-area curve for spiders collected in July, 1980 fromstructurally complex - natural litter

ne s s (9 ) . The Sh a nn on (1948) inde x o f d ive r s i ty t a ke s thef o r m :

sH ' = - Z P i l o g eP ii= 1

w he re P~ i s t he p rop o r t i on o f t o t a l i nd iv idua l s i n s pe c ie si , a nd s is t he num be r o f s pe c ie s . A n e s t ima te o f e ve ne s sis J (where J =H'obs / H 'max ) (P ie lou 1966) . Fo r c a l c u la -t i o n s o f d i v er s it y , d a t a w e r e s t a n d a r d i z e d t o a c c o u n t f o rm i s s in g d a t a o f d i s t u r b e d b a s k e t s u s i n g t h e f o r m u l a : S i = N i( T / T - D ) , w h e r e S i i s t h e n u m b e r o f i n d iv i d u a l s o f s pe c ie si a f t e r s t a n d a r d i z a t io n , N ~ e q u a l s t h e n u m b e r o f i n d iv i d u a l so f s pe c ie s i , D i s the nu m be r o f d i s tu rbe d ba s ke t s a n d Te q u a l s t h e t o t a l n u m b e r o f b a s k e t s ( 24 ). T h i s s t a n d a r d i z a -t i o n w a s d o n e f o r a l l t r e a t m e n t s .To fu r th e r a ss es s d i f fe re nc e s be tw e e n t r e a tm e n t s i ns p i d e r c o m m u n i t y s t r u c t u r e w e c a l c u l a t e d f r o m s t a n d a r d -i z ed d a t a a f t e r lo g t r a n s f o r m a t i o n [ l o g1 0 ( X + 1 )] ( C l i ff o r da nd S te phe ns on 1975) pe rc e n t s imi l a r i t y va lue s u s ing thef o r m u l a :

P S = l - o . 5 z l P . - Pblw h e r e P , i s t h e p r o p o r t i o n o f a g i v e n sp e ci es i n t r e a t m e n tA a n d P b i s t h e p r o p o r t i o n o f a g i v e n sp e c ie s in t r e a t m e n tB (W hi t t a ke r 1975 ) .W e a l s o a n a l y z e d g u i ld c o m p o s i t i o n a l c h a n g e s b e t w e e nt re a tme n t t ype s , W e a s s igne d s p ide r s t o gu i ld s (R oo t 1967 )w h i c h a r e f u n c t i o n a l g r o u p s b a s e d , i n t h e c a s e o f s p i d e r s,o n f o r a g i n g m e t h o d . S p i d e r g u i ld s y s t e m s m a y r e f le c t , d e -p e n d i n g u p o n i n f o r m a t i o n a v a i l a b l e o n f o r a g i n g m e t h o d ,g ros s (B a logh a nd Loks a 1948) o r s l i gh t (Pos t a nd R ie c he r t1977 ; R ov ne r 1980) d i f fe re nc e s in s p ide r fo ra g ing be ha v io r .A g r o s s f e a t u r e o f s p i d e r f o r a g i n g i s t h e u s e o r l a c k o fu s e o f w e b s in p r e y c a p t u r e . A s i m p le d i c h o t o m o u s s y s t e mb a s e d o n w e b u s e d i s t i n g u i s h e s t h e ' m a c r o - g u i l d s ' w e b -bu i ld ing a nd hun t ing s p ide r s . W i th in th i s w e a l s o re c og -n i z e d f iv e s p id e r g u i ld s , m e m b e r s o f w h i c h c o m p r i s e d> 9 0 % o f a l l sp i d er s c o l l ec t e d . M e m b e r s o f t h e w o l f s p i d e rg u i ld a r e ' s i t a n d w a i t ' t y p e p r e d a t o r s w h i c h f r e q u e n t l yc ha nge s i t e s (Fo rd 1977) . A s e c ond gu i ld , t he runn ings p ide r s , a re a c t ive , pu rs u ing p re da to r s (G e r t s c h 1979) .M e m b e r s o f th e t h r e e r e m a i n i n g g u i l ds b u i l d w e b s : t h e s c a t -t e re d l i ne , s he e t w e b a nd va g ra n t w e b-bu i ld ing s p ide r s .


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36R e s u l t s a n d D i s c u s s i o n

Abundances of SpidersI n e a c h m o n t h b u t J u n e w e b - b u i ld i n g s p i d e rs w e r e s i g ni fi -c a n t l y m o r e a b u n d a n t i n a r t if i c ia l l it t e r t h a n i n n a t u r a l l i tt e r(Tab le 2 and F ig . 2 ) . A do m in an t w eb-bu i lde r , Neoantisteamagna ( s e e A p p e n d i x ) c o m m o n l y m a d e w e b s i n a r t i f i c i a l

T a b l e 2. F ratios o f 2-way analysis of variance on abundance ofweb-building and hunting spidersMo nth Web-builders Hunters

Source of variation F value F valueJune Structure 0.22 0.22Artificial 3,79" 8.21 *Structure * artificial 0,17 3.60 *July Structure 8.77 * 3.17 *Artificial 29.47 * 0.34Structure* artificial 4.99 * 0.08Aug Structure 5.29" 1.59

Artificial 34.23" 6.22"Structure* artificial 1.92 0.80Sept Structure 0.25 2.63Artificial 30.40" 2.68Structure* artificial 0.77 1.20* P < 0 . 0 5

l eaves (T . Bul tm an , pe rsona l obs . ) . Ar t i f i c ia l l eaves werem o r e r i g i d t h a n n a t u r a l l e a v es a n d t h e y m a y h a v e p r o v i d e dsuper io r s i te s fo r web a t t ach m en t . I f th i s i s t rue , s ign i f i can te f fec t s o f l i t t e r type (na tura l / a r t i f i c i a l ) fo r web-bu i ld ingsp ide rs (Tab le 2 ) a re m is lead ing ; w eb-bu i lde rs soug ht a rch i -t ec tu ra l , no t nu t r i t ion a l , qua l i t i es o f a r t if i c i a l l eaves.Unl ike web-bu i lde rs , hun t ing sp ide rs were s ign i f i can t lym o r e a b u n d a n t i n n a t u r a l l i t te r i n Ju n e a n d A u g u s t ( T a b l e 2and F ig . 2 ). The re a re s eve ra l pos s ib le reasons f o r th i s . Be -cause the re were m ore web-bu i ld ing sp ide rs in a r t i f i c ia ll e av e s , p r e s e n c e o f g r e a t e r a m o u n t s o f s i lk w e b s m a y h a v ede te r red hun t ing sp ide rs f rom these l eaves . Also , g rea te ra b u n d a n c e s o f p o t e n t ia l p r e y (C o l l e m b o l a a n d / o r D i p t e r a )in na tura l l eaves m ay sugges t hun t ings sp ide rs were aggre -ga t ing in a reas o f h igh prey dens i ty (Bu l tm an 1981) .Web-bui ld ing sp ide rs occur red in s ign i f i can t ly g rea te ra b u n d a n c e i n t r e a t m e n t s o f g r e a t e r st r u c t u r a l c o m p l e x i t yi n J u l y a n d A u g u s t ( F i g . 2 a n d T a b l e 2 ). O t h e r s h a v e o b -ta ined s im i la r re su l t s wi th shrub l aye r sp ide rs by p rov id ingf r a m e s m e e t i n g s t r u c t u r a l r e q u i r e m e n t s o f w e b - b u i ld i n gspec ie s (Colebourne 1974; S chae fe r 1978; Robinson 1981) .A n e x p l a n a t i o n f o r w h y s t r u c t u r e d o e s n o t e f f e c t w e b - b u i l d -i n g s p i d er a b u n d a n c e s i n J u n e a n d S e p t e m b e r i s t h a t e n v i -r o n m e n t a l p a r a m e t e r s i n f l u e n c i n g s p i d e r p o p u l a t i o n s m a yc h a n g e s e a s o n a l l y ( B u l t m a n a n d U e t z , i n r e v i e w ) . U e t z( 1 97 9 ) p r e v i o u s ly s u g g e s te d s t r u c t u r e i s o f m a j o r i m p o r -tance to l i t t e r -dwe l l ing sp ide rs dur ing m id-sum m er whi lee n v i r o n m e n t a l p a r a m e t e r s s u c h a s p r e y a b u n d a n c e , t e m p e r -a t u r e a n d h u m i d i t y c o n d i t i o n s , a re m o r e i m p o r t a n t i n e a r l y

%

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4

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N C F ' N CArt. Nat.W e b . Bui l ders

F N C F N CArt. Nat.H u n t e r s

108

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0 F N C F N CArt. Nat.Web. Bu i l ders

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F N C F N CArt. Nat.Web. BuUders

liN C F N CArt. Nat.H u n t e r s

Fig. 2. Mean number of web-buildingand hunting spiders collected per sample.Bars represent 1 standard error o f themean. F, N, and C represent falt, naturaland complex leaf structures respectively.Art. and Nat. represent artificial andnatural leaves, respectively


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100 n n i n g S p i d e r s8 0 I l f S p i d e r sg

& 60E g r a n t W e bc ~ [ [ d e r s4 0" ~ a t t e r e d L i ne.~aversC~2 0n e e t L i n e

. ~ Q v e F s

0 F N C F N CA r t i f i c i a l N a t u r a l

F i g . 3 . R e l a t i v e a b u n d a n c e o f d o m i n a n t sp i d e r g u il d s p e r t r e a t m e n tt y p e , u s i n g n o t a t i o n i n F i g . 2

37Table 3. Duncan's multiple range test for mean values of spiderspecies richness. Nota tion as in Fig. 2

S t r u c t u r a l t y p e N u t r i t i o n a l t y p eF N C A r t . N a t .

June* 4.043 3.857 4.630 3.735 4.642July 4.957 5.830 6.043 5.814 5.400Aug ** 6.500 6.795 7.844 7.077 7.000Sept 5.348 5.553 6.136 5.986 5.353

* Significant differences in species richness between artificial/nat-ural treatments only in June (F1,~29 =6.36, P


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38Table 4 . Com mu ni ty matr ix ba sed o n percent s imi lar ity of sp iderspecies. A and N re fer to art if ical an d natu ral treatm ents respective-ly, while f , n, and c refer to flat , natural and complex structuralt rea tments

A f N f A n N n A c N cA fN f 0 .723 -A n 0.738 0.716 -N n 0.734 0.735 0.759 -Ac 0.738 0.709 0.793 0.738 -N c 0.708 0.744 0.725 0.771 0.715 -

Table 5. Araneae species diversity (H') and evenness (3) for treat-men t types. N ota t io n as in Fig . 1Tr e a t me n t sArtificial NaturalF N C F N C

June H ' 3.377 3.429 3.724 4.022 3.650 4.2 21J 0.482 0.505 0.539 0.594 0.513 0.572July H ' 2 .933 2 .802 2 .947 3 .999 3 .3 8 8 3 .325J 0 .385 0 . 3 3 1 0 . 3 4 1 0 .447 0 .424 0 .417Au g H ' 3 .405 3 .307 3 .2 7 1 3 .852 3 .573 3 .615J 0 .413 0 .390 0 .376 0 .485 0 .4 5 1 0 .433Sept H ' 3 .596 3 .662 3 .894 3 .720 3 .9 2 1 3 .902J 0 .462 0 .477 0 .498 0 .5 1 1 0 .506 0 .512

a t t he s pe c i e s l e ve l ( s e e Appe nd i x f o r s pe c i e s l i s t ) . A l l s i mi -l a r it y v al u e s a r e > 7 0 % . M a t r i c e s o f i n d i v i d u a l s a m p l i n gm o n t h s a l s o s h o w e d n o d i f f e r e n c e s b e t w e e n t r e a t m e n tt y p e s . T h i s s u p p o r t s o u r p r e v i o u s c o n t e n t i o n t h a t l i t t e rs t r u c t u r a l h e t e r o g e n e i t y w e a k l y a f f e c t s s p i d e r c o m m u n i t yo r g a n i z a t i o n .S p i d e r s p e c i e s d i v e r s i t y ( H ' ) s h o w s n o t r e n d w i t h s t r u c -t u r a l c o m p l e x i t y w i t h in n a t u r a l a n d a r ti f ic i a l tr e a t m e n t s( T a b l e 5 ) . M e a n s p e c i e s d i v e r s i t y f r o m n a t u r a l l e a v e s ( 3 . 7 6 )h o w e v e r , i s c o n s i d e r a b l y h i g h e r t h a n t h a t f r o m a r t i f i c i a ll e a v e s ( 3 . 3 6 ) . D i f f e r e n c e s i n d i v e r s i t y b e t w e e n n a t u r a l a n da r t i fi c i a l t r e a t m e n t s a r e s i g n i f i c a n t l y d i f f e r e n t ( M a n n - W h i t -n e y U t e s t, P < 0 . 0 5 ). A s w i t h s p e c i e s d i v e rs i t y , e v e n n e s si s a l s o s i g n i f i c a n t l y h i g h e r i n n a t u r a l t h a n i n a r t i fi c i a l t r e a t -m e n t s . T h e r e i s n o d i f f e r e nc e h o w e v e r , b e t w e e n s p i d e r s p e -c i e s r i c h n e s s i n n a t u r a l a n d a r t if i c ia l l i t te r .

S p e c i e s d i v e r s i t y i n a r t if i c i a l le a v e s w a s l e s s t h a n i n n a t u -

r a l l e av e s b e c a u s e d o m i n a n t s p e c ie s b e c a m e m o r e a b u n d a n ti n a r t i f i c i a l l i t t e r ( s e e A p p e n d i x ) , a n d t h e r e f o r e e v e n n e s sa n d t o t a l d i v e r s i t y d e c l i n e d . D o m i n a n t s p e c i e s i n b o t h a r t i f i -c i a l a n d n a t u r a l l i t t e r w e r e p r i m a r i l y w e b - b u i l d e r s . W e f e e ld i f f e r e n c e s in d i v e r s i t y b e t w e e n t h e t w o l e a f ty p e s ( n a t u r a l /a r t if i c ia l ) a r e a r e s u l t o f g r e a t e r r i g i d i t y o r t h e a r t i fi c i a lm a t e r i a l a s c o m p a r e d t o n a t u r a l l e a v es a n d a r e u n r e l a t e dt o d i f f e r e n c e s i n n u t r i t i o n a l c o n t e n t b e t w e e n l e a f t y p e s .

Conc lus ionsI n f l u e n c e o f th e l i tt e r h a b i t a t o n s p i d e r c o m m u n i t i e s isc l e a r l y c o m p l e x , i n v o l v i n g a c o n c e r t o f p o t e n t i a l l y i n t e r a c t -i n g v a r i a b le s . L i t t e r - d w e l l i n g s p i d e r s a r e i n f l u e n c e d b y v a r i -a t i o n i n l i t t e r h a b i t a t s ( U e t z 1 9 7 9 ) . V a r i a t i o n i n l i t t e r d e p t hi n f l u e n c e s s e v e r a l v a r i a b l e s : l i t t e r c o m p l e x i t y , p r e y a b u n -d a n c e , t e m p e r a t u r e a n d h u m i d i t y . P r o b a b l e p a t h w a y s b e -t w e e n i m p o r t a n t v a r i a b l e s a s s o c i a t e d w i t h l i t t e r d e p t h a r ed i a g r a m m e d i n F i g . 4 .

I n t h e p r e s e n t s t u d y , r o l e s o f li t te r a s a s t r u c t u r a l l y h e t -e r o g e n e o us h a b i t a t a n d a s a t r o p h i c b a se w e r e d e c o u p l e df r o m e a c h o t h e r t h r o u g h f ie ld e x p e r i m e n t s c o n t r o l l i n g l i tt e rc o m p l e x i t y a n d n u t r i t i o n a l c o n t e n t . L i t t e r c o m p l e x i t y s i g -n i f ic a n t l y af f ec t s a b u n d a n c e s o f w e b - b u i l d in g b u t n o th u n t i n g s p i d er s ( T a b l e 2 ) ; th i s m a y b e r e l a t e d t o t h e f o r m e rg r o u p ' s n e e d f o r w e b - a t t a c h m e n t s i te s. U n l i k e a b u n d a n c e ,c o m m u n i t y s t r u c t u r e o f li t te r - d w e ll i n g s p i d e r s i s o n l ys l i g h t l y a f f e c t e d b y t h e c o m p l e x i t y a n d n u t r i t i o n a l c o n t e n to f li t te r . I n l i g h t o f p r e v i o u s w o r k , i t a p p e a r s t h a t d e p t ho f l i tt e r, r a t h e r t h a n i t s l e a f h e t e r o g e n i t y o r n u t r i t i o n a l c o n -t e n t m o r e s t r o n g l y a ff e c ts c o m m u n i t y p a r a m e t e r s o f sp e c ie sd i v e r s i t y a n d c o m p o s i t i o n . I n r e f e r e n c e t o F i g . 4 , w e f e e lp a t h w a y ' A ' i s m o r e i m p o r t a n t t h a n p a t h w a y s ' B ' o r ' C 'i n d e t e r m i n i n g p a r a m e t e r s o f s p i d er c o m m u n i t y o r g a n i z a -t i o n . T h e m e a n s b y w h i c h t h i s o c c u r s m a y b e t h r o u g h v e r t i-c a l la y e r i n g o f l it te r o r b y t h e c o n t r i b u t i o n o f d e p t h t ot o t a l l i t t e r v o l u m e , a n d i t s i n f l u e n c e o n p o p u l a t i o n s i z e sa n d e x t i n c t i o n r a t e s .Acknowledgements. A num ber o f people were extemely helpful inthe i r ass is tance dur ing the f ie ld work. They are : M. Bruggeman,J . Bul tman, J . Chapman, N. Fol ino , B. Hol l i s , W. Hopple , D.Jackson, T. Jones, C. Meininger, G. Stratton and B. Sweeney.We wish to thank H. Levi for kindly identifying the difficultsp ider species . We are gra teful to the author i t ies of Cam p K ernfor allowing the use of their land for this study. We also extendthanks to S. Faeth, T. Kane, M. Miller and S. Rissing for helpfulcriticisms of earlier drafts o f the ma nusc ript.This research was suppo r ted by- Sigma Xi , the Univers i ty ofC i nc i nna t i Summe r Fe l l ows h i p P r og r a m a nd t he De pa r t me n t o fBiological Sciences Research Council at the University of Cincin-nati.

L i t t e rD e p t h

V o l u m e o f A = S p i d e r S p e c i e sH a b i t a t R i c h n e s s

" e : r s , t yT r o p h i c _ Prey + _ S p i d e rB a s e - A b u n d a n c e - A b u n d a n c e

I Fig. 4. Probable in terac t ion pathway s o f severa l l it te r p arameterswi th popula t ions of spiders and the i r prey


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A p p e n d i xSp e c i e s l i s t

H a c k l e d b a n d w e a v e r sD i c t y n i i d a eDictyna muraria E m e r t o nA m a u r o b i i d a eCallobius sp.S h e e t l i n e w e a v e r sL i n y p h i i d a eBathyphantes pallida ( B a n k s )Centromerus cornupalpis ( O . P . - C a m b r i d g e )C. latidens ( E m e r t o n )Ceraticelusfissiceps ( O . P . - C a m b r i d g e )Ceratinopsis atolma C h a m b e r l i nCorniculari sp.Eperigone maculata E m e r t o nE. tridenta E m e r t o nErigone antumnalis E m e r t o nHelophora insignis ( B l a c k w a l l )

Islandia sp.Lepthyphantes zebra ( E m e r t o n )Meioneta unimaculata (Banks)Microneta viaria ( Blackwall)Neriene variablis ( B a n k s )Prolinyphia marginata ( C . L . K o c h )A A A

N u m b e r p e r t r e a t m e n t t y p e xA f N f A n N n A c N c

B B BC C CD D DE E EF F F

1 4 25 1 2

S c a t t e r e d l i n e w e a v e r sM y s m e n i d a e

3 130 172 2

4 2112 6

15 0

133391 4

2

19

751342163

21

2 4

233228

363 9

13917

222

4 2

14 32

113

7 21 4

22 718

312

14 6

51932

1218

14 036563 6

32

4 4

Mysmena guttata ( B a n k s )T h e r i d i i d a eEuryopis limbata ( W a l c k e n a e r )Pholcomma hirsutum E m e r t o nSteatoda albomaculata ( D e G e e r )Theridion frondeum H e n t zA A AB B BC C C

1431516

11 24 17

223

1118

V a g r a n t w e b b u i l d e r sH a h n i i d a eHahnia flaviceps E m e r t o nNeoantistea magna ( K e y s e r l i n g )A g e l e n i d a e

Agelenopsis pennsylvanica ( C . L . K o c h )Circurina robusta S i m o nWadotes calcaratus ( K e y s e r l i n g )Tegenaria sp.

34 0

82191

7168 439

1 0 88 722 13

62 1 3

8823

24 8 8

17116

W o l f S p i d e rsP i s a u r i d a eDolomedes sp.Pisaurina mira ( W a l c k e n a e r )P. brevipes ( E m e r t o n )L y c o s i d a eLycosa helluo W a l c k e n a e rPirata minutus E m e r t o nSchizocosa ocreata ( H e n t z )

110 33 4

16 5

9 7 1 3 6 1 1 842 41 61

21 1 94 9

1 N o t a t i o n a s i n T a b l e 4

1 1 12 1 2 6 28 21 18 35 208 29 7 46 5

1 1 11

1 i 1 1

43

62 3 4

6142 7

1 1 65 0

39

7112 42

8 22 5411


7/8

40Species list (continued)Species list Number per treatment type 1

Af Nf An Nn Ac NcRunn ing spidersGnaphosidaeDrassyllus virginianus Chamberlin 2 1 1 2 8

Litopyl lus temporarius Chamberlin 2 6 4 2 2 5ClubionidaeCastianeira cingulata (C.L. Koch) 10 5 8 13 7 9Clubiona sp. 1 4 2 5 3Phrurotimpus alarius (Hentz) 25 47 23 81 23 88P. borealis (Emerton) 5 11 7 13 10 19AnyphaenidaeAnyphaena celer (Hentz) 8 2 12 13 14 7PhilodromidaePhi lodromus marxi Keyserling 3 1 2 1Crab spidersThomisidaeXys t i cus fra t ernu s BanksX . f e r o x (Hentz)

Tamarus angulatus (Walckenaer)10 274 4 22 24 5 344

1 3Jumping spidersSalticidaeEris m arginata (Walckenaer)Marpissa l ineata (C.L. Koch)Zygobal lus be t t i n i G. & E. Peckham

3 11 21 1

6 4 1 0 133 2 2

R e f e r e n c e s

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Gertsch WJ (1979) Amercian Spiders. 2nd ed. New York: VanNos-trand Reinhold CoGibson WW (1947) An ecological study of the spiders of a riverterrace forest in western Tennessee. Ohio J Sci 47:38-44Gorman OT, Karr KR (1978) Habitat structure and stream fishcommunities. Ecology 59: 507-515Huh ta V (1971) Succession in the spider communities of the forestfloor after clear-cutting and prescribed burning. Ann Zool Fenn8:483-542Jocqu6 R (1973) The spider fauna of adjacent woodland areaswith different humus types. Biol Jaarb 41:153-179Karr JR (1971) Structure of avian communities in selected Panamaand Illinois habitats. Ecol Monogr 41:207-229Karr JR, Roth RR (1971) Vegetation structure and avian diversityin several new world areas. A m Nat 105:423-435Lowrie DC (1948) The ecological succession of spiders of theChicago area dunes. Ecology 29:334-351Luczak J (1963) Differences in the structure of communities ofweb-building spiders in one type of env ironment (young pineforest). Ekol Pol Ser A 11 : 59-221MacArthur RH, MacArthur JW (1961) On bird species diversity.

Ecology 42: 594-598MacArthur R, Recher H, Cody M (1966) On the relationship be-tween habit at selection and species diversity. Am Nat100:319-332MacArthur RH, Wilson EO (1967) The theory of island biogeogra-phy. Princeton: Princeton Univer PressM'Closkey RT (1976) Community structure in sympatric rodents.Ecology 57 : 728-739Moulder BC, Reichle DE (1972) Significance of spider predationin the energy dynamics of forest floor arthropod communities.Ecol Monogr 42: 473-498Murdoch WW, Evans EC, Peterson CH (1972) Diversity and pat-terns in plants and insects. Ecology 53:819-829Penniman AB (1975) The ecology of the spiders of the old fieldsuccession in central Ohio. Unpubl. MS Thesis, Ohio St Univer,Columbus, Ohio


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4:[P i a n k a E R ( 19 6 7 ) O n l i z a r d sp e c i es d i v e r s i t y : N o r t h A m e r i c a nf l a t l and deser t s . E colog y 48 : 333-351P i e l o u E C ( 19 6 6) S p e c i e s - d i v e r si t y a n d p a t t e r n - d i v e r s i t y i n t h es tudy of ecolog ica l succes s ion . J The or Biol : [0 :371~383P o s t W M , R e i c h e r t S E ( 1 9 77 ) I n i t i a l i n v e s t i g a t io n i n t o t h e s t r u c t u r eof sp ider comm uni t i es . I . Com pet i t i ve eff ect s. J A nim Ecol46 : 72%749Rec her H F ( :[969) Bi rd species d iver s i t y and h abi t a t d iver s i t y i nA u s t r a l ia a n d N o r t h A m e r i c a . A m N a t : [0 3 :7 5 - 8 0Riech er t SE ( :[974) Thoug ht s o n the e cologica l s igni f i cance ofs p i d e rs . B i o S c i e n ce 2 4 : 3 5 2 - 3 5 6R o b i n s o n J V ( 1 9 8 1) T h e e f f e c t o f a r c h i t e c t u r a l v a r i a t i o n i n h a b i t a to n a s p i d e r c o m m u n i t y : a n e x p e r i m e n t a l f i e ld s t u d y . E c o l o g y62: 73-80R o o t R B ( :[ 96 7 ) T h e n i c h e e x p l o i t a t io n p a t t e r n o f t h e b l u e - g r a yg a n t c a t c h e r . E c o l M o n o g r 3 7 : 3 : [ 7 -3 5 0R o s e n z w e i g M L , W i n a k u r J ( :[ 96 9 ) P o p u l a t i o n e c o l o g y o f d e s e r tr o d e n t c o m m u n i t i e s : h a b i t a t a n d e n v i r o n m e n t a l c o m p l e x i t y .E c o l o g y 5 0 : 5 5 8 -5 7 2R o t h R R ( :[ 97 6 ) S p a t i a l h e r e r o g e n e i t y a n d b i r d s p e c i e s d i v e r s it y .E c o l o g y 5 7 : 7 7 3 - 7 8 2R o v n e r J S ( : [9 8 0) M o r p h o l o g i c a l a n d e t h o l o g i c a l a d a p t a t i o n s f o rp r e y c a p t u r e i n w o l f s p i d e r s ( A r a n e a e , L y c o s i d a e ) . J A r a c h n8:20:[ 2:[5

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