INSTRUCTIONS CHECKLIST - Medipac Travel Insurance

SPATIAL AND TEMPORAI, ECOLOGY OF EASTEKN SPADEFOOT TOADS ON A FLORIIIA LANDSCAPE

' [!SI>A

f i r c ~ s tSrrxicc., S o r ~ t h e nHr.rc~~r(./l ~ Stolion, fk.111Cn,ck Erpl~irr~c,r~icil I:orc.,i 1577 Br-c.oiirr1Rd., Ashi~cilli;,KC 28806, (ISA '~eorgt. IY. Tc~~~rtcr. Pr($.~.sor. Ilc~pc~rtr~zcnt of IVildliji, I.:c.~log!~nrrtl (:or~.sc,n;(~iiorr. I~~stilutc, ofFi)orl clrltl Agn'crtli~rrr~l Scic~~ccs, Nvrci~~s 7~,igli,r:303, Hor 110 f.30, Il~~icc~rc.if!y of I;/oritla, Cni~rc.st,i/k~, I T ,32611-01,30,USA

ABS.I,HA(:.I.: Efft!ctive arr1~11ibi;rncoirscwation rit~rstcor1sit1t.r i~opt~latiori ailti la~ttlsc;tpc. procc~.;sc3s,111tt information at riitlltipl(~sc;rlt.s is rare. Wc cxplorti spatial ;~rr(ltcxrnporalpattt,r~~s of 1)rcbc~tfiilg nrld i-caw-riit~ttv~rt 1)). tXastcrnsI)atic~foottoads (Scclphiopvs holhrookii), using nirit. yc3arsof c1;rt;l ti0111 coritiririolrs ino~iitoriiig\vitll tirift frriccs arrtl pitfall traps at cxiglit cpllr~rnc~ral porrtls irr It)~iglcafpin(,-wirc,grass saritlliills. Brc~cldirrgcavcxi~ts ( 2 2 5 atlr~ltsat a porrtl witlrirt a ~rronth)occlrrred 23 times on niitrs occasior~sat scvvn o f t l i ( ~c'iglit st11t1~ lxni(ls, b11t sr~l)starttialrecruitrncrtt (LlOO irletainorplts) fi)llowc2c1only five cXvcrrls.Ii(~c~riit111c~nl ~.;~r~grlt! 1*o11i0-3648 among po~itls.Orily So~lrpontls friric~tionetlas popttl;~tiorr "sorrrccs", ;inti orrly drrring sorrrt. years. 'l'hcs otlior ~ x ~ r ~ carttl l s , evc.11 "sol~rcc"pontls tluring some ycars, fnnctioned citl1c.r as "sirrks," \vlrc~c,1)rc~t~tlirrg occirrr-c~l with no resillting recnritnient, or were not tlseti at all fbr i)rcxc,tfirig.Most rc~c~r~~ittnt~nt oc.clrrrc~tlt-lurii~gSotrr years. Recrlritment was correlated with atlult breeding c,ffort, b i ~ ortly t diiri~tgso1111~ ycS;trs.Rc'~.iiptlrrt.sw'rc rare, artd ir~ter-pondexchange of adults was rtli~iinialand short-tlista~icc( < I 3 0 111; on(. was 416 r t i ) . Most (83.5%) i~idividualscaptured were rnet;rmorplrs, ant1 15.9% wr,rt, 2.51 rnrn ((,st. 2 4 yr). W11 c~onsc~rvativc4y estimated a 7-yr lifespan. Ad~llt"population" trends clearly rt~fl(~ctt~d hrc~etlingcffi)rt rat11r.r tl~arrntr~nberspelse; capture rates flirctl~atetidramatically among years, lmt slrowrtf rro ovt,rall trt~ritlstlt~ri~ig tlrcx 9-yr-strrtiy. Orrr piper is provides empirical irtforntation th;it car1 bt: ~rsc~tf to genrratc3 rc;distic rrictapoltiilatiorr ri~odr~ls for S. holhrookii as a tool in conservation planniiig.

&y tuor(1.~: Amplribian populatio~rs;Anr~ranI>rclc~ding;Eplrc~r~lcwl porrtls; Ilydroperiotl; M~~t;ipoprrletiori; S~clphiopurholbrookii; Spatiefoot toads; T(,rnporary wetlantls; \Vctlancls

AT FIRST glance, pond-breeding amphibians represent ideal rnetapopulation models using a "ponds as patches" paradigm, where pop~ilationscontint~allybecome extinct and are recolonized by ;i~npliihiansfioin r~eighboring poilds (Marsh and Trer~harn,2001). However, this paradigm may not hr realistic, either because metapopulation dynamics of a given species are not as they appear, or becalisc; assumptioris are not rnet. Assulnptions iiiclude: ( I ) extiiictions and recolor~izations commonly occtir, (2) local extinctions are tllc. result of' stochastic processes in othe~wisr suita1)le breeding llabitat, and (3) inter-pond distances influence recolonization processes clue to limited dispersal ability by amphibians (Marsh and Trenhain. 2001). In addition. perceived extinc:tions may be biased by sarrlpling prot)lems, s~icllas species that are missed in surveys, skipped breeding seasons when climate or pond cotictitioiis are unsuit-

able, or cxploslvc l)rc.cding followrtl I)y long (M,iisli ant1 Trcnperiods ultliout I)~-c~cdtng h,lns, 200 I ) Accllratc pcLrceptioris of ,implilhi,in mrtapopulation dynamics ant1 popnlatioi~ trclntls rccpir? long-tvrtr~, I,mdscC~pt~-sc'ilt. sttlcltt>\ to distingt~ishbehvrlrri local c~xt~nction ,ii~dI(.colom/attoli, n,ittll-,il popli1,ltiori fi~ictt~dtiot~\, aiitf true cfccl~nes,it ,i rc.giorla1 level (Hlali\tc.in ct ,i1 , 1994, (:lbhons ct a1 , 1997, I'cch~nann ct dl , 1991) IJong-trrit~sarnplilig at a singlp s ~ provide\ t ~ vit'il ~iiforniatio~i '1l)otit dlrtI~hit)1 ~ ~ 1poiid 1 t ~ s cant1 l i f r Illstory, l ~ u tsdcr1ficc.s ,i 1andscapc.-sc.,llc pclr\pc,ct~vcx of cotill~ni~t~ty ,illti ~nc~i~ipopnl,ttioi~ dyrl,ni~ic\ (I>odtl, 1992, S(~lr11itschet 1'1 , 1996) 0 1 1 1 - p ' i ~ 1s ~ 11it(~11(lul ~r tcvrrpor,tl 'i11(1 spdt~~il \ ~ ~ ~ - ~ ~of~ l1)rc~l111g, ) i l ~ t y to p ~ - o v ~ lc(~' ~ n p w ~ infor~rt,itlo~i cal t11,it car1 11i. rc~cr111tlrrc>1lt, l o i ~ g c v ~ t,nlcl > , nlt(v--porltl 1i1o\7cX- 11sc.tl to g"1l"r'ltc' rc~,illstlc lric'tgrd\sgro~trlcl eovel-, eIlri~lcc~, ri~i(fthe s ~ ~ ~ t a i ) iof' l ~ both t y I)rc.cdltig \vitlcly \paced longlcdf pine trccxs, alrd few pontls '~ntl the. s ~ ~ r r o r ~ l l d iirpl,ultIs ~ig (II(3l1s harcI\voocls Since 1976 tlic- npl'11ltl 11~,1tr1x t I , 1 ) M11cl1 of wll,it 1s l is 1e5rcq)tlolls of ~,O~~lll'ltlOll I)lllll 27 < l ( l l l l t 5 (11 < I l l \ \ l l l q l ( ~ 1 ) 0 1 1 ( 1 ~ l l 1 l l l l gtgl~t5t11tly po11d5 of 1.3 O c ' t o l > c ~1995 I)rclc~tl~ng, ((l,~tc>\) 1 ) vo5t ( $ I \ 01 I I I I I ~ , ) \ ) I - ( J C X ~ I I I ~1>(>r~o(15 rnc~t,ul~orl)li\ c ~ ~ t I'olitl c ~ l $311-o111 6 N i o \ ~ > l t t l ) ( ~( F I ~ (1111-rngS(~13tc~~nl)cr or 0ctol)cl-, I)nt 1905 tlrrot~gltM,iv I996 (1)11t1 r t o 5 t l)v Fc~l)rrl,lt~ ocac.~~rrc.tl I ) l c ~ c ~ c ! ~ l r q also occ.111-1-c~l ( ~ I I I - I I ~F~l)rlldry, ~ 1996) All ~ ~ ~ ( ~ t ' l ~ i iSI-OII~ orpl~ tl115 s l)r(w111rg b1~le~tl 11) l)itf;illt r q x \\Tt> c,ipt~n-c.tl 0.3364, RMSE =: 755 36; F I y,,= 27 88, l-' < \ ~ g ~ i ~ f i c ~I I rI O~I l- t~11\~1lc\s l~ ~ ($576%) t l ~ i ~ f(~111'11t~s r 0 0001), illtiicnting a thl-csholtl for tlrc~11111ilbr-'r (32 4% ) t111rulgIjrc,cltllltg c~ c ~ t s( I = -.3 52, of breeding athtlts, bclo\v whlcll rt~crnitlllrwt tll = 22: P < 0 005) was riegligible (Fig. 2) Whtx~i( ~ X ' ~ I ~ ~ I I Ithe III~ klost (8.33 ( k ) c , i p t ~ ~ r cS~ d1iolhn)oklz \\(.I-? years separately, lxowrvrr, wt, Iolrltd a sigllifi- ~llc)tal~rol-plts Most 'lt111lts\z7c1re(~itllc~l111 tllv cant relatlonslr~p betwc,c,n ~ d l l l t ljrc~ltl~ilg 3 ) r o1t1 (34 7% ) 01-5 \ r oltl (62 I % ) ,lgcLc l , i \ ~ \ effort and recrllit~nel~td l ~ n n g 1995 (r" (F1g -3) \'(.I-? ~ ( ' M J111(11\1(itldl\111 tllv ~111~111('\t 09325, RMSE = 13531, F , , = 8284, 1) < .iqt, cl,tsscls (1-3 !I- 2 :3% ) w c lollnci, ~ 111o\t (1111-111g ilo~iljrr~t~tlii~q 0 OOOI), 1996 (r2 = 0 9646, RhlSE = $379, of th(>scl\ \ J t s l t x c~i1>tilrc~tl tls In tllc. oltle5t colrort (6\ r F1 I = 81 81, P < 0 005), ,uld 2000 (I-' = l ~ t ~ r ~ oiirtl~\rtl~~,tl\

,111 lo^ 1 ) r c ~ ~ l i ndlir111g g thc' other years The ot1rc.r .tx)~l(lsfltnctlonecl e~tlier as "sinks," \ ~ l l r t ~ -l~rcltd~ng r~ o~cnrrcclrarely and In low ~it~nll,c~rs, 1)11t rt,snltc~tl In no recniitrnent (I'ontls 2, 7, ,111~1S), 01- were never used for I)rc~c~111ig (Pontl4) Arnong-pond differences tn 1)1(>(~111ig '11111 r c t . ~ ~ i ~ t ~ ~(lid l ( - ' l not i t 'Ippear I cll,itc.tl to In (11ologj, tvh~chwas similar 'imong ,Ilrrlfl-r/i~c~~N ( ~ c ~ ~ ) ) l (lit(/ rrn~ ~rrostponds ((;rtcwl~crgand Tan~ler,2004) lttf(~t"-/)olt(l 'I101 ( J l l t i , t t / (:oncc~ptu,ill), ,r " m r t a p o p l ~ l , i t ~ oIncorIlo~~" ii tot,d of 12 7% of fil-st-c,lptl~~c>tl ,itll~lts r,ltc3s ,it 1e'ist sonicJ tilspersal 'imong distinct ~ ~t111s s c'ise, ponds) (Hanskl and \\l(,r(l ~-c.captl~rc~l (-380 of 2986) Most ~-c'c,lp- I x ) p ~ I . i t ~ o(111 t11r~socct~rrc~(l (111ri11g the s ~ i ~ ?(,'ir ~ i v c~il>t~irc~(l > 2 j I- 'litc~r, 111- 1)011(1s,'intl short tl~st,inces(except once) when cl~c,itiilqtll,it tllc,> \ \ ~ c ~,it~ -lt-.,tst c t 1 1 1 ( ~ , j c1,ir5oltl ~tocc111rcc1Ycxt, tlrc intenla1 I~etweenbreed~ng ( h t ~ ~ 1 1 5nlost c ~ I ~ ~ ( ~ ~ ~ ~ I I I O\\rt,ro I - ~ I I Itoo ~ s1r1,iII c.1 tl~its,it gl\ c11 porld cuceeded the app'rrent to tor,-cl~p),h\7o ~ncli\~tltl,lls wc,~-cl~ c ~ a p t l ~ rI~ic~sn,ni t ~ l1 of ~riostirid~vid~ials JWP esttnl'ited '1 11fcsp,1u of 7 vr), too long to sustatn after fi\ > clars, dlld 0 1 1 ~~ ~ ~-c~ed~>tlt~-c~d 1 s d f t ~ ~ In,i\nnllni six ycA,irs,~ n d i c ~ i t tli'it ~ ~ i gS Ilolhrookll c,in 1 1 ~ ~ 1v~,il)lc popnlation if ponds dtd not recetve ,it least sc\ c11ycxars (T,il)lc 1 ) ~til~~iigr'i~lts fro111 sltrrolinding ponds For Most (95%))c ~ c l ~rr~c,ipt~~~-c~s ~lt occ111-1-c~l , ~tlrr t r.x,~niple,large nli~nbrrsof adults bred at Ponds pon(1wlic~rc,t1ic.y wcJrc, first c,iptl~rc~d (T,i1)1e2) 5 'ind (7 111 2000, although the last substantla1 Mln~nt,ilrxucll,irigcsof ddlllts occ~irrc'd r i ~ t l o ~ ~~-c>c~-lnt~ncnt g occt~rrc,d sorrietin~e before the Pontls 1 , 2, and (3 (n~,t\i~ril~ttr d~st,nicc~ 1 :30 In), stt~tfjbc,ga~lm 1904 (at least 7 ,5 yr earlier) 'inti o ~ i e~ n ( l i ~ ~ d lis ~ , lrcleclptl~r(~d l 'it Pond 8, T1r1s incongrn~ty1)etween ~pp,irentpond fidel,11qxo\111i.~t(,lv 41 6 In f I - ~ I I'o11t1 ~ I 7 W ~ I ~ ~ ~t I - ~C ~ 119 5 vt.1 s11s w~tllln-poncl recrt~ttnlent~nterv'ils o r ~ g i ~ i ~ ~11idrk(~1 lly (Tcibl(22 ) 13c>e~11s(> nlost t l ~ slloltltl t leati to loccl-1extinct~on,coultf be nic~t'lniorl-~lis \ v c ~ vnot ~ n ~ i r k\vv t ~ ico111(1not I)cc,t~lsc> ( I ) atlt~ltS holhrookzz are phtlopatric dc~tc~t-lnlnc \vl~ctllc~~t1ic.y 11sc~1 ~ronn,ltalponds Imt ] I \ ( ' Ionq(-'r th,in we estimated, (2) adltlts lor I)rc~c~tl~~ig ,is ,idrtIts nio\cJ ,lnlolig 1x)ncIs morcJ than we detectecl, or (*3)~ric~t,i~no~-phs d~spcrsc.and breed In nonnatdl 1 ) 0 ~ ~ 1'155 ,ldlllt5 13cc,i11sclwc dltl not nark cmlgratlng niet'i()Ill" d'lfd l)rO\ ldc' ( ' 1 1 1 ~ ) 1 1 ~ 1 1~ ldl ~f 0 l llldtlOll tll'lt tnornlis \vcl c ~ ) l ~ lnot ( l assess their d~snersal I L c,tn I)(, 1 i \ c ~ 1111 tic\ c~lop~ng I c'.~l~st~c ~nc~t~ll>ol)l~Most ~il(-'t~iniorpIis entering ponds were capI ~ t ~ o~rlotl(>ls n for S Ilol1)rool~lliI'(1 fo1111tlthat tlitc~ltl111-trig ~rt~lss clxodlls events frorrl those ,111 t x ) ~ ~ (wc,rcx l s not "crtl,itt.cl c ~ c ~ ~ c~~a l~"t l,isi t ~ lx)~i(ls,\~tggt>sting that they were part of the 1)rc~c~ctirtg 51tc.s or ,is sot~rcc~s of j11\c1111(~ rc- s'i111~r(~crltitnlent('vc'nt IIowever. oilr data t~lllltl~1(~llt .s llo/l)~-ool,ll l)l-(Yl I11 \ < l l - l < l l ~ l c ~t11d s r ~ g g c ~t11,it t solric. short-distance Inter1 1 1 1 1 1 1 ~ ) ( ' JlllOl1g ~~ 50111(', 1)llt 110t 'ill 5t'('ll1111~~> pontl ~rio\vnicnt occlirrecl (Greenberg and st~~t,il)lc 11oncls il~ttonqtllc. c,~qlrtpo~ltlswcx I dn~ic~r 2004) (:c)nct~c ,rnalvscs would shed studlcYi, Olll\, 11'ilf (Po11tls 1 . .3, 5, 'llltl (5) \\/(xr(x 1110r(' ligllt on tlicl ~rlc~tnl>opl~l,ltio~~ connectivity llsc~clfl(Yjtl(~lltl?, tllollgll lrl c~glll l)r(~(~(llllq I-(,(,\ (,lit ,it ,11i\ gncn 1x)11cl, stiggcsting that there 5llltcYl 111 I 1 0 l-t~c.lllltlll(~llt) or \\('I.(' not 11\e'(1 'it 111'1) 1)c' ~1crit~c'iltlirc.sliold of I~reediogeffort,

0 I-Yk ) ,ilso wc.1-c, I ,)I-(,, tlirl 1,lrgc.st 111(1i\ 1(111'11s c a p t ~ ~ r\vc,rt> c ~ l SO Inn1 S171, (]:IS $3) I\'(XI1o(1(Fig .3)

7

J

i )

26

I l ~ l ~ l ~ l ~ , ~ l ~ ~ l , ~ ~ ~ , l ~ ~ ~ \

1 1'01

6I

1\'0

1

1 1 c,\t~~ict 01ircl,lt,i sJro\v t l l ~ tharcd , clggs, or t,ltlpol(hs 101- sll1)\tri~it~dl 51lr\~i\dlto ( ~ ( ~ locally i at I)l-c~c~tl~rig po~lds,pc~rcc>l)t~o~t\ of m c t ~ o r to o occrtr 1)odtl (1992) re- o ~ tr.ipp111g .ircl Il(lri\~ly1)liiscd 1)y pol-tc~lonly ortc, 1,irgc. S I~olhr-ookll1)rc~c~tllrlqS 1rolh1-ookzz,il,t~tld,u~cc~ cvrrit In five. y(,,i~-s,it '1 polld 111 1rortlr-c(,1ltr,ll \\icx,itllcv-, po~ltl Itytll-olok~ ((;rc~c~lil)c~~-g ,111d Flol-~tl,t,1)nt 1 1 0 rc~c1-n1tlrlc~11t S(~111llt5cll c%t'11 'I',illlic.r, 2004), ,ulcl I)rc,c~tl~~rg 1>Ile1101ogy 111 (1096) also rrxpol-tcld otlc. sriccvsslt~l rc~c1-111t-0111- stttcl~,~il)l)clr(x~~t I ( Y ~ I I ( , ~ Y !l ~ o l > r ~ ~ 01 ~it~o~ ~ n c ~ of n t S I~olI)~-ookli tlc.sIxt(l ~-cy)cx,ltc~l 1 ) r ( ~ ~ lr-i ( l ~ ~Sl t 11o/l)r-ooX11 (1111-111g \o111('1(lri1 s cI(~~LI-I\ 111gc v ( > 1 l t 5 o\7c1r 16 y(,~-ot(~l~t~Lil ~iI)r(~vdl~ig follo\v~~~g o~llyfi\c of 2:3 ~ ) I - ( Y Y I I I(I ~ ~ e ~ l t'i11c1 s , (b\c,~it 1111-tli(>1 1)1;i\(,s l)op11I~it1011 e~t1111;itvs \vcis 11ot \K)~it~c~lly 01- t ( ~ ~ r ~ p o rs~J~ ~I~l yc l ~ ~ - o ~((;os~~cx~ ~ i / ( ~ ( 'i11(1 l 13lcic*l, I I I ( I I \ ~ I ( I I I ~18-50 ~ I ~ 1r111r \ ~ r r x( e~l j c'ip,lcross tlrcx stt~dyponds I i o ~ ' i - l ~ ~ t occ11r1c~cl ~il(~it \vcXro ,it only o~lc.of ciglrt potrcls tlrlrl~lgt111c.c. >c,,t~s tl~rc'cl,it po~itls,rrrost c,ll>trlrcd11rc11\1dt1,ils (1994, 1995, arrtf 2001 ), ,111tl ,it hi7oI)o~~(isS 1-72 trill1 SVI,, or ,tl)oi~t4-Fj )I-old (\Yi-tght, t l ~ ~I I - ~ tli~ring2000 0tlrc.r strt(l~c>\ I I ~ I L ~sI10\\711 , t11,lt 2002) rI'li~s11i(11c~ltc~\ lt rcaltlcllll111 \i~rvlvdlof d ~ l l l r ~I,in~,lr~ l ~ l to ~ric~t,i~rlorpllos~s 1s ] I I \ ( > I ~ Irollsf,tctors ~ r ~ c l r t t i w,~tc~r ~ ~ i g 11~>1~i1icls 5111 ro1111c111ig ]301i(15 lo1 - 3 4 c,:i1-5 at l x ) ~ ~ lfor s tllc,~r first t c ~ ~ n p ( ~ r ~((:osnt.r i t i ~ r c ~ ,uld Hl,ick, I955), prcl- I>c~fotc.,y)l)c,,n.~~~g t r I i it I ~ i t r s p c ~ i Ic) I ( T ~ I I'itI ~l,~ge\ (Alforti, 1989; l1(>trank,i i~rrprc>s\to~l oL ,~~ltorlg-pori(l c>xt~~ic~ttons ,tntl ,ind Kcnrrc~tly,1999, Milll)ltr, 1983, 1987) \I/'(, r c ~ c o l o ~ ~ ~ / , ltll,it t ~ o ~II,I\C> l s ~~rlportarit ~~riphcasuggest th;tt p r ( ' ( l a t ~ )on ~ ~ cggs a~rcl !;ir\,icx t1o11\for ~ ~ ~ c ~ t ' ~ p o ~I >I ~ I ~~( !~~~i >t ~ ~ o I Ir Ii ~ S ~ ~ I ~ I ~111pl11lg ~ ~ L I - Iri~iy ~ , slll)\('t of o11r c~igllt had a ~niijor c 4 l i ~ ton ~-ccrrt~t~lic>~lt 51iccc,\s l)cca~~sia ( I ) l)repdi~lgcvclrlts \vr>1-castro~rglv \tiid! polids 1111glltI t ' l ~ v lcxlt 115 ~ l t 1 1 \'PI-> IIII~>~~Y\Io OfI ~S /~ol/)~*ookzi l)r(~c~(l~~ig ,~s\ociatetlwith po~ldst11,it Il,id rccctitly tlnc~tl (l~lf(>~cvit I I - ~ I I ~ I I I ~I I TI115 ~ rc~s~tlt 11111sa11tl rc~fill(~ti-otic~wciy to , ~ \ o ~ l11g11 c l I)r(~(frito~I ofIo~rg-tc,r~rr, lci~~dscs~p(~dc1is1tic.s tll,it n1,ly Incrc,lscl wit11 lotrqc~~ 11: (11-0- tl-,itcxstliv ~~~ll)ort,iilc.e periotl5, (2) c,icli ~ ~ i l ~ s t ~ rcc~-lt~t~rlc.~lt i~~t~al Ic\c,l st11t11c.sto c I ~ \ t ~ ~ ~ g 11~it11rdl ~lisl~ '1111p1111)1;111 occr~rrc,tlo111)111 pontls tll,lt hati rc~c~c~rltly tli-~c~lK ) o l ~ ~ ~fl11et11~it1o115, l ~ l t ~ o ~ ~or l)(>r(q)tio11\ ofI>ol>fro111t1-11c'tIc~c1111c~s (Pclclr,nit1 rclfillrd 131-lorto I)rc.c~1111g c ~ c w t \((:I-~Y.I~- rtl,ltlo~rIl~~ctti,itiorls, I \ I I - I ) 1.:\c.11 tilt, rc~l,iti\cxl\ l>el-g,lncl 'T,lrlnc.r, 111 prcl\s), ,ind (-3) tlic,~c,\\l,is I spdt1~1I< i ~ i ctc>1111)or~i/ I sco~)(> 01 0111- st11(1) ,111 ,1pp,ircxnt" I ~ ~ I I I I I I ~ ~ Itlrl-c>sl~oltl" III 01 I)rc~c~hiig1'11-g(>1 I t - I 1111tg I 1 1 tI q 1 1s 111\(~ly~ ~ i ~ i ( l (to~ c t ~ t1-11c~ ' t111i(>1,irvacl th,it \vc.~c. pot('rit~.ill) lr~glic>~iol~glt to I,irrtl~c,il~c~ e o r ~ ~ ~ ~ l 01e ~ \t11(>1r ~ t \ 1)0~)11I'itto1l " s 1 1 i ~t o 1 - - t c 1 i l l o i ~ gI - tl\rr,uri~c.\ Eff('ctl\or~il (l l ) o l > ~ ~ l (~I i\ tI ~ ~ ~oI~I I~Ic I~~ ~\ l ( I ,i~ttocori-c~l,~t~o~l 111,iyli,nc, t ~ l ~ l ) o ~ -1111p11c~it~ii~t I~lc,-lr~stor\ tl,iit\ that cilf(~ct tions fi)~-tic~c>lol)~~tg p ~ - o l t , ~ l ) ~ ltllotl(.ls ~ s t ~ c of spc~cie~\-spc~c~fic dt d l~i~i(ls(~,q>(~ 1(,\(,1 0 1 1 1 st11c1) p,itcll occt~p,i~icj 1)) & I I ~ P I I I I ) I ~ I( IK~ I I ~ I ~('t) ~'11> , 11(~1-\1stt~11c(l 11111\t1-~~t(~s ill(, pot(iilt1d1 c o ~ r t r ~ l ) r ol ~ t (>deb ~o~~ 200.'3; T I - ~ ~ I c'It ~,u-s, to otlrcsr\ '1s ~ c l ( ~ ~ tI)\c c\ ~ t11P 'ithllt ~~0~111;III(,> ili . t i o ~ ~ 17: l20:(. (:., \\', 1). KIII,,\I(;,hl. J. hloi\\11\, S. 1,. t-(,g~~l;ktio~r. !\t~~(>ric;t~r Katl~r;~Iist 1:32:f I I I r o :~II(I ( l ~ t~c~t~i ~ ( ~~t~: t~~ r r oirl l~ ~ I ~ o s i sc11ro11>. 1col11gic;1l!\l~l>lic;~tio~~\ 13: 1.722-15'12. i::~~-olin;~ gopl~(srliog (I{/III/I~/IJJ;IO I Y I ~ ~ I OI I)I S O I I ~ I I \ \ ' I I . II{,~ ~ 11. \I. IW7, li(~,g11I;iti1111 I I ~ ' \ ~ ~ I I ( , ~ I I iI lYl , ~ I I I I I ~ ) I C ~ \ (;;~rolil~u. ]ot11-11;11 01' I Ierl~c~tolo#v 2H:fi12-lil-i. \ > S ~ ( > I I I \ : c~\lx".i111(~11t;t1 ~ ( ' I I I ~ ) O I W > I ) o ~ ~( (. ~l I I I I I I I I I I ~ ~ ~ ( ' \ S I . : \ ~ I . I ~ I13. ~ ~I)., ~ : I 1). I , 1.S( I Y I - ~ ,j.. 11. K. I > P ; ( : I I \ ~ \ \\ \\ I. ) I(iS. I 4:37- 1.452. 1. If'. (;1131io\\. 19$K;. S ~ I - I I C ;~~ I II ( I! ~O)II:IIII~(,~ I~ of' :III \\'11.1311%. 11. hl., 1). j. \ I O l i l \ . \ \ I ) 11. N 1 1 \ 1 i I ~ l \ . l>)S:3, ;~~rrpIril~i;~r~ ( ~ o ~ ~ t ~ ~ rc~vitlt~r~cr~ i i t r i t ~ :fiotlr ;I I(i->.c,;kt-\tlldyof' S:I~:IIII;~II(~(~Il)~-(~(l:~tiotr ;111el t11c \ ~ ~ I I ( , ~ I I II -~ ( ~~ ' ( ~ \ ~ ~ I ~ ~ ~ I I I ( ~ I I ~ ~ I ~ ;I I I ; I ~ I I ~ , Ll /> o ~ ~1'1(1.i . 21 7- 24S. 111 11. I.. (;11(iy ; L I I ~J . !\. ~ , ( J I I I I I I I I ;ILII I~I I~~ : ~I I III T' s~ C: I ) O ~ I C ( ' L1(.010q (%I: 142:1-I.42!). S~rr;tll\~c~c~tl (I':tlc.), 1.111rg-tc>r-tr~ St~rtlic,s oi' \ ' c ~ t c ~ l ~ ~ ~\\.NI(;II.I'. tc. I\. 11. 2002. 1,1f2~-11ictori(~\ of' t 1 1 ~1:rogs oi' ~ I I I , ( ; I ~ I I I I I I I ~ I;\c;~tI(,~~iic I ~ ~ ~ ( > \l'r~xss. . S ~ Il)i(,go, I ( ; i ~ l i f i ~ r ~ ~ i : ~Okc4i1rolic~c~ , S\v;itrll~.i;cv~rgi;~. N ~ r i l r!\t~~c.i-ic.nrr S:~li(x~~ti;~ 0.S.A. iiZ1111r;ONo. 7. (:or111~11I j ~ ~ i \ ( l r \ iI)I-(~c\, t) I t l ~ i t c . ; ~\. i ( ~ SKP;I.I,I,1). K . , b:, 1 . \\'I.:I~\I,:I%. ~ L I )S. [I, ( : O I V I . \ ~ I ~ I ( ; I I T . Yot-k, [l,S,j\. 1