Comparative response of Rangifer tarandus and other northern ungulates to climatic variability

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1 Cpi p f Rgif gl lii ibili Rb B. Wlji 1 *, Di R. Kli 2, Øi Hl 1 & l M 3 'Dp f il i, gill Uii f Nw, P.O. Bx 5025, N-1432 Å, Nw. 2Ii f i Bilg, Uii f lk Fibk, P.O. Bx: , Fibk, K 775, U. 3Dp f Bilg, Diii f Zlg, Uii f Ol, P.O. Bx 1050 Bli, N-0316 Ol, Nw ""pig (b.wlji@if.l.). b: T f iflig lif i i ppli i, i i iigl big gi ip f il ii. I i pp, w iw i p f kwlg ig ff f lii ii (ll glbl) ppli p f g l, wi pil pi i/ib (Rgif ). W l i l ii ff f li g b fg ilbili qli, i ii. Vi iff w ibl; wi i ppi i lig lif i i f gl, fw i w i ff lii ib T i lil b wi w ibl pl ig ifi l iflig gl ppli p. Ti b b ff f w gl pp li i lif i i l p i i iffil il lii ff f i p f. Ti fi plxi f liip bw i. W pi lii f pbili bw iffil f glizig b ff f li g bl, pi wii pi. F, i pp b k li-l f f lg f i/ib b lk i li f lii ff lg bi. I lig f i, i f f i f lg i i i ig ff f li ibili i/ib. K w: b, ib, li g, fi, i,, NO, N li Oilli,, i, x i, il, w. Rgif, 22 (1): Ii Diig f pi-pl ii i lif i i g iiil i f i gl f lg (Bg, 16). Ti qi lg- i, i f ppli iz gpi p, Rgif, 22 (1), 2002 wll iifii f ip f iflig i (Li, 1). M f lif i i ifl b g b f iii xii ii i lpl p f ppli (Cppi & Pi, 15). I ff - 33

2 ip f f iflig lif i i ppli i f gl, i i iigl big gi ip f il ii (M, 14; Ti, 15; P, 16; æ, 17). Di-ip i i i ppi i, if pi, ib bill ppli fli i gl (æ, 17). gl p g wig, f xpl, i i ff l il (Gbb, 174; æ, 17; Gill, 18). Hw ff f w ppli i i bi ill i l pfi ill (æ, 17). Cli il il bi iff l (Hll, 184; kgl, 18; Kjl, 11). F xpl, i ff f l illig ifll l i f gli (Pk & Rbbi, 185; P, 16), i ig w p i f li i w (Pk, 184). Cli l ii l gl ppli p g i ff p I, lii ii kw j ip b iil qli f pl i ilbl bi (Di, 184; V, 14). F gl, f ii i pp li p b l ilbili f i i wi (kgl, 185) (Ri, 183; Ri, 17; Hjlj & Hiøl, 1; Fi, 2000). T b iw ff f li lg bi (æ, 17) ff f li i pifi ggpi wi i fil il (U.K.; P, 16) x li (G & kgl, 17; Kli, 1). Hw, b i f w i ig lii ff gl wi pi i, i i pi æ (17) f b ig f ii bi bi lii ff lg bi. I i pp, w iw i l p f kwlg ig ff f li (ll glbl) ppli i lif i i f gl, wi pi i ib (b Rgif ). W l i gig pibl i, b i f ii ff f li il pl qi qli, i iil ip i ppli i f gl gg b P & (1). F, w i ii ff f li g i i/ib, p b lgl lk b iw i i fi I, li l iil gl, i f i ib, g ii wi i ff i ppli p (, 14; G & kgl, 17; Möl & Kli, 17; Möl, 1; Cl, 2000). g ig f i i g b glbl li g ffig (Hg, 2000). Clii ibili pl p Pl pi B f i ii i li, pl lp iff il gi p wi fbl il ii. Tiz & Zig (1) ll i «l*. li, fbl i l ii b pi (Hi, 185), pl p ii il lii, wll pi. B f gwig gll lw p ii i ppi i, lig p gg b ip f iflig pl gw lp (V, 14). B i i ip f ig wi f, lf ig wi b fi ip pi (Li, 180). El lf lp bl pil b pl iliz l gwig, pl p w b liii i lf xp i (Tiz & Zig, 1) Ll lii ii fg qli qi lg i, w, lig f gw f pl, piip i pppi ppi f pil pl gw (Ji & R, 184). Yl ii i fg qli i gl ifl b w (B0 & Hjlj, 11; Fi, 2000b). B f iff i ii i il f, B & Mi (184) f w fl iff i b g gw i pl pi q f lp xp i iff i i i i il f. w p ff lg f gwig fg qli (Hjlj & Hi0l, 1; V Wl, 2000). Tiig f wl l 34 Rgif, 22 (1), 2002

3 ifl iig f g f fg pl (Kli, 185; Lg, 16; P & Kli, 1). M ipl, i f p w i plg pi f wl g fg ibl i f w Fi (2000) f l ii i w ifl fg ilbili igibili g, f xpl i i i Gwig Dg D i M J. (16) p ig pi ll i bi (Bl p.) w i wi ig lw p ig f pipii i l, ig p i l. Hig il p, w, f l i ligifi i f pl ll wll, lwig igibili. i i pi qi g f i i, f i ig ppl g f pl i, i ff i p ig p ig lig ii (Di, 184). Cl b g p i f bg pl bi ii l f fg b ffig f lig i b pl (V, 14). Ti b li w pl qi ig lig i i p f pil gw lg lw li. Ri lig l, i f f fl gi, f w f, w, ip qli f lf i fg b xig l g f plg. Ti l i lw i f l piig i lii li f igi-iibi ig il i f b wi b p i x f gw f pl (B, 183). B0 & Hjlj (11) f il g ip ig l, w. T p, f iigi ip bw pi (Bl p.), wi i l ii, i i i pi: i i i f fg p ig li i, l ig pii ff fg qli b i lil pi l b pl ig (Kli, 170) lw bil p f fib i. wi w i ll, w ii ig l. Tf, ilbili f fg gl ig wi will b ff. I, l w fllwig w w i, l i l i w «iig» g f liiig f bi gi. Ti will ff pi fig il i fil l Rgif, 22 (1), 2002 i i bw, wi l b flxib Glbl lii g pl qli qi Ml f li g pi glbl p pipii will i wii x, g big p i li ig wi (Diki, 186; Mxwll, 17). Wig i pipii b i gi f i ig p, i wi li l (Cp & Wl, 13; z, 15). B & Wll (17) l i i wi wfll i Nw Tii f C. B p pj li g ff ff qli qi f pl il ilbl fg f gl g g big b i, i ilbl il i ig pl gw (Kli, 1). I f ig i xpi w lg pl gw pi wi ffii il bl i, will b i i pl bi (Kli, 1). P & (1) f N li Oilli ( f lg-l lii ibili; blw) w igifil l pl plg, pl pi bl li fllwig iigl w, w wi wi b pi big ii lii ii. Ti l ff lg f gwig wi, w x, ff f ig wi pi f pl (Kli, 1). Ti wl f l i i wi fg, b wi ibl qli (Cpi &, 16). T pj i i l ig l l i xi f pi w fg i igl igibl lil lw ll f il (Bll, 1; Kli, 1). Ti l b f g lgil ip ig li w pl gw i. Ll li gi lg bi ppli B wig B wig i f ip lif i i (Cl, 184). l p f lif i f i wi b wig; f ff b wig l ip ifl ppli i (Kli, 170b; æ, 185). 35

4 l i w liip bw b wig ii g (k & Lik, 180; Ri, 183; Lg, 16; æ, 16; Hjlj & Hiøl, 1). Lg (16) gg wig gi f (C lp) ig l gw p l b pi ig l M-J w. T i i wi plgil lp f fg pl ig pi f l w, ig lf: i igibili f pl p li lwl, ig ql i f ilbl i. Cli ibl xp g ll w ii, f, b xp j ff f g qli qi i, wi i ff gl b iz b wig. Clii ibl, b g i i ff fg qli i qii ilbili, b xp pl j l i ppli i f gl (Tbl 1). æ (185) f bii f lii ibl xpli igifi p i f l ii i wig f (l l) i. M p pipii w ip ib (16) p wi lil ig b f i w w gll i wi lw p ig f pipii i l, wi ig p lw p ipii i l l fl T l f pii li, lg igifi l f l fl, bw pi wi w b, ggig l ff f w ibl b. T ff f wi li pp l b ip, g, f xpl, ii ff f l w l fg qli w i i fg ilbili, b i ff l b ip. I wi ig wfll, ib l w lig 10 f g pbl bl gi ffii b b fll lig ( & Dl, 18). Ngi li bw b i f pig wi b p f wil i (kgl, 183). G (16) i wi ii wi li i b f pi i l fl ib. Ti w p Il, C, w ib i i ( b, l p b ) i pp b l wi i ix (Oll, 17). Hjlj & Hiøl (1) l p i b f l lig wi iig w p ig pig wi pig. F, ii f l b p fllwig wi wi lw p ig w p (æ & G, 188; Cl, 11). iill, Cl (11) f gi li bw b l w p. w p b p b l ll lii ibl igifi ff b i ø-tølg, Nw (Li & Lg, 18; Li, 1b). iil gi ifl w f blk-il Oil i (Pk, 13), wi-il O. igii (M & ig, 181). Ti iff, w, f l f i b Oll (17) ib, i fi liip bw wi i b ii. Rpi pf x i g i fi gl l b wig ii (Lik, 188; Gill, 12; Jg, 13; F- Bi, 14; Lg, 16; æ, 16; Ri, 17). Tf, lii ii i xp ff lif i p (Tbl 1), b i liip, w, iff gill iff li gi (æ, 16). Fi (2000b) p pp i f i lig lig i J w p iil l Gwig Dg D pi M J gil l b Gwig Dg D pi Jl w p wi pi bi. C (13), p pbbili f fl pg f fl ib b lgl p i ig pi big, b ii, w l ii ig l pg g ifl lig l lf i (16) f i w ig fi i f w i wi ig lil w pb w. T ppi f 2--l lig i Nw, iff i ppi lig 3--l R, l, w gil l ii i M-J g 12 li (Lg, 16). T l l ppi f i lig R li wi iig b f wi w i wi. Wi ii, l ifl p i. T pi pf f fl 36 Rgif, 22 (1), 2002

5 ib (2, 3, 4 6 l) w ff b wi wfll wi li liig wi iig l wi w fll ig wi pi big ( & Dl, 18). I Nw æ (16) f fl l li f w wi wi l w. iill, l i w f i g (O i) i ppli liig wi ii (Jg, 13). Kk (1) f fi f fl wi ifll i p ig wi. Rl, li ibl b ff ppli x i i (Kk, 1; P, 1b; M, 2000). T ppi f l b l il f R, l, li wi wi ifll, i wi i iil i fl (Kk, 1). I Nw, ppi f l ig li wi iig w p i M (M, 2000). T l x-i w l.g. x-p li ig. Fl fw iil i wi iig i f li b R, l i Nw. T xii ifii f gl x i b li ppli l i pbbl pi (Kk, 1; P, 1b; M, 2000). Hw, pilgil i b wi x i i i i kw (Hwi & Gill, 1). Pibl i il ig i li f l ii, ifi i f x i pi / ipl i (Ri, 1; M, 2000). il, b ppli gw l liip b p bw wi w b l li i l pi f gl (Mik, 167; Cl-Bk & lb, 182; & B, 183; kgl, 185; Gill, 13;) b f l (L, 2000). Di ff f wi lii ii il l p, b l b ff f w. kgl (185), pig iff wil i, l liip bw lf il f fg ilbl ig l wi. Cib i w lk l xpi ig lf li i w wl w lil l (F & Wi, 11). l ii i ppli gw i lb i w gl gil l wi pipii. Hig gw w wi w, ff f li w g ig ii (, 2000). il f l f b x w piil l wi p gil wi wfll, wil l lig g i ff f -g lig il w i l lig pii ff f -g w l l il (Li & Lg, 18). gi ff f wi w f f R, l (Cl-Bk & lb, 182), wi lw l fl il (Tbl 1). Jil li w b i ig wi wi b ig ppli i ig Nb-J ifll (Kk, 1). i b Jg (17) i big p (Oi i), Li & Lg (18) i liip bw wi i l i T bi w iff i li b x g l wii ppli i p w ib I wpii (C i), lf il l piil wi Jl p gil wi Nb pipii wil w il w piil l wi J p, b gil l wi M p l Jl pipii ( & B, 183). I i w, ff f li i wi ppli i. I, ff f wi w il f wpii w l g ig ppli i ( & B, 183). Mli i i l ib ii. Cpig liil gi b i wi g ii i if f Nwgi ppli, æ (16) f li w ig i, w ig fw l i ig wi. T f liip bw wi f ppl lf li. bjø (2000) f l ii i il g l b lil g l, i ppli f i Nw. T ig li w f g ig i w ppli. T l li f l w iff g gi ig b wi. bjø (2000) ib p f l li lii ii i b wi. El il i (Cpl pl) fw i wi iig ifll i M Rgif, 22 (1),

6 f R 0 2 L b 0 2 i Fi 8 0 b, 0 00, l,. l l L g ^ /5 Ü C L i L i g L --^ k K 2 8 i bil i i li l l l l p i l l p p l l g l N blt 38 l L T l i b i i li Cl i p p i p i g Dp i i p k k,, l l l l N x i l fll i w l w l w i L W i i T b lf l C ib i C i b i i <-i i p w N l l, i i P w D) D (G ( J g M D ^ ig w: B i i w i PHOP i b i D D G i lp l p w I w w g g,, l l øø TT øø II I + + I + + p l f w w p /3H/: l l i i f f l l l l BCC R R l w w, NRNR III l i - w, N R l i f i J - b I li p N ig w H i Rgif, 22 (1), 2002

7 3 8, B i g i W * * VC w ö, i i G +,+ + + G-N l l C w ö, i i G ø l, j lj æj H w N w NF II l i G k k P + 3 k, l l CC, i g i M k, Y w N i L F, lp T F, P 3 g J C, lb l l i f i p p -J w w i M W l i i i i i i i l l l l l i i l 6 lf lf w w w l li l l g f l B B i g l l Y B B l C C C C C l C i i i - i i - V V f. f. R k- M i g * C M l p W Rgif, i 22 (1), 2002 Bl 3 W g w i l p l

8 J (Gill, 17). T il f i (Rpip pip) b p i w pipii w ig p w lw ig pig pig i P, wil ff w i lp, b il fllwig lw g- pig (Li, 1). I iigi f gil ff f li i, L (2000) f b f li l i Fil w gil l b w w wi w pi, pi il w w pi bi (Tbl 1). Ti i pif w li lf b w i il w ii i i Glbl lii gi lg bi ppli I N li gi, wi lii i bili i, lg x, xpli b lgl li f pi p ll N li Oilli (NO) (Rg, 184; Hll, 15). T NO f iil illi i pi wi f i Il bpil li f z Ibi Pil (Hll, 15). B ig f NO i gl gil, i i qifi ll b ipl ix b i i f g -f p bw z Il, f Db g M f wi. Pii l f NOix iz ll w, w wi i Ep ( ll l, wi i N i), wil gi l iz ll l, wi i Ep ( w, w wi i N i) (Hll, 15). B NO f f fli i wi p ipii i li (Hll, 15, 16; Hll & V L, 17) wll i i i wii p l p i b il l f li g (Hll, 16), NO l pil ff lg f pl il i N Hip (P, 1). Hw, ff f i lg-l lii ix l i iff ll w p i iff (M, 2000), ii il plx. T NO, g i ff gi lii ii, ifl l p f lif i lg f il lg - 40 li bi, ilig ppi ii, fi, x i, gpi pp li i p (P, 17, 1,b,; F, 18; P &, 18, 1; Li, 1; M, 2000). Glbl li g will likl i b x l i f p w i i, f q f i li ig li (z, 15). Iig, f, b xp i, liiig fg b ib (Kli, 1). b p f glb l g will ff bi ppli p ( G & kgl, 17; Tbl 2). B, fi x i T i li bw NO -pifi b wig kll i f l w w i (P, 17). Rl, w, ppi li bw NO b wig w p f i Nw; b ig b wig f l (Li, 1b) lw b wig f l (P, 17) fllw wi pii NO. M (2000) f ig l w b P (17) i l f g gi ff f ll i, b i NO i ig p f. ig NO ix w iii f ig pipii. wi p w f Nw i l 0 C, w p, w, ig NO (Hll 15; P, 17) l i gi li bw w p NO lw l i, w i liip i b ppxil (M, 2000). Fg i fil l i pbbl il ilbl i l lw l ig wi wi ig NO ix (Li, 1b), ig NO ix pbbl ii fbl ii f. I, b wig f i Nw i gl w piil l wi NO w kig ppli b i ig ix f ll i, lg i liip w lw NO l (. M, N.G. Yz, N.C. & R. Lg, pbli l). O R, l, P &, (1) b bi l f i fllwig pii (w, w) NO wi wil fi f fl l gil wi NO ix f wi i Rgif, 22 (1), 2002

9 wi w i. T l p g i liip f b i lf fl fi wi NO i Fil Nw pil (Tbl 2). O, Li (1b) f b f w piil l NO, b lgg. l, wil lig i Nw w i if b fllwig pii NO wi, b f i w, w wi w w il, li fllwig w, w wi (P &, 1). Ti gg ff f NO ppli p i gill. P (1) l f NO ig fl lp f i i ifl f i, b g. Wi wi ig NO ix l l iigl l bi x i f ffpig, ip f g i g f i (P, 1b; M, 2000; Tbl 2; l ii b). T w, w, il ff f NO ff f w p ( lw li) w ll f lbi wi iig w p (M, 2000). b il I, F (18) b, w, i wi (ig NO) w i wi b (g wi li fllwig ), w l w- ff f w, i wi pii ff b (g i fi f b fllwig ig NO), wi pii ff big lil lg. T gg NO ff w x-p ifi, pig piil g fl g f ppli. I l b w l fl l p iffl li i ibili (P, 1). P (1) p i b 0-T0lg (Nw) R (l), l b f i w f iigl w wi (Tbl 2). W i Nw, i i w ib f pifi fi f 2 -l i, R i w ibbl i i l fi f 3 -l i. I, l l b w gil l pi NO R (P &, 1). B b (P &, 1) lb il (Mil, 1) f p (Oi i) l b p fllwig ig N O. Li (1) f l i b f pii NO (iii f ig p lw pipii i wi) xibi lw wi il i P, wil i lp, il fllwig p ii NO. igl, gg i b ff f wi w il f bi pli pil. O w i f li, ib,, kx (Oib ) wi-il b i fllwig ig NO (Tbl 2) i W Gl, Il Rl, E Gl Mi pil (P &, 1). T f i f b wi-il i N i w ifl b glbl lii fli 2 3 lg, wll l i-p fbk wlf pi (P &, 18). Cli, i ii i iig ff f lii ibili gl ppli gll l i ff f i. Hw, i i pill b ip. Kli (11) gg b ki wbl (Hp i, Oi) l b fli (Cpi p, Oi), i Gl i 150, ifi b i l b j f iflig li i b ii ppi f ppli f W Gl ib. kkii (15) i ib i l l p 2 kg f bl qi ig i Ti Pil, Ri. Cl (2000) p lw b i ppli f i i Nw f wi i f l (Tbl 3). I pp i ig pi f gig p wig w ig i wi w (pb ), g gig p w f b qli, i ff i/ib b ii ipl f g qli. Cibl bil i pp b i ip l lik bw w p lw b ii f i. I, gggi i f i ib ig w w i wi i ( & Nil, 16; Nl, 18), wi l i i i p ig / ig, i p fig, wi il iil - Rgif, 22 (1),

10 f R l L T, I P l l,w i i i i 8,,,,, I I I I I PP PPP l w w w w i w w F i N N N N I III + i 8 F P w N g l - l l l l l l l III, I P ø T,,,, w,, ø RRRR ø N RR II I 7 P w N 00 b 00 b 2 2 1,,,..,,.,, l l l l.. l. 1 l....,. l... II i i i l i L P MM L L P P M i l w w H NN III Z F, lp T,, + I I I 88,, II PP BB,, l R i MI l l i b i i li Cl i p p i g p pi D p i i p i i i p w 1 OO NN i i R O O O O O O O O O N N N N N N N N N li g l Y M b R b l T b l l 6 l ) i i g g O OOO O N N N N N l l ) f x i p ( P l g l l l l i l i l bb b L b i C O i P N p x k M > ' gg 3 2 O O NN i i f f ^ï W 42 Rgif, 22 (1), 2002

11 q (Hll, 12; Tpi, 16; Möl & Kli, 17; Cl, 2000). Uig ii g p i ii f, Möl & Kli (17) p ig f ii g g ib i l lk w i w p. Tp wi p igifi ff f i i g f ib (Tbl 3), flig gli f i ii b bii f. Bil p i i l b. I, p b Pll (16), ib, ii bi, bi w igifi l ifl b f pii i qi i, ig p-lig pi. Epk & Lg (17) p lig w w g p wi fl i i Nw. Ti ii pibl il ff f i l pi f. T j pii i f pi f i: qi (pill p., Clii), wbl fli l b fli (G & kgl, 17; Möl & Kli, 17). B, & Nil (18) p i f pil ibi i b blk fli (ilii) fli (Tbi) wl Rb (178) b ig p g f ig ig lw pg f lig i fl iil ll f qi. Möl & Kli (17) f p f i fli lg ifl ii bg f ib i qi b bill iig f i p ig ig f i p fig. I i ig, wi i pk pi i g i ik qi f Rgif, 22 (1), 2002 q l i bil i i li l p I 3. l blt f R l L I l i b i i li Cl 6, N i - < w N, U w k, N l j > i g g I 0 2, l l C ,, i i i i,. 0 2, l l l H i p l & & l ö C TM U U,, w k k i l l l i g i g i p g i i f I Q i i 6.l l l P, k l l b b b b b i i i i i q qq q i i i l p p i i i l i i l p i p i V

12 gl b i. Rll (13) p wbl l b fl l f f J il l g, wil qi w piil pbl i Jl. I Nw i l f 10 Jl 20 g (Nil & Hg, 14). lg i i bl g i w bii f w i bl ff ll f i, i i l p, wi li ii pi iig f f i ib, li ip f ig wi i f, li (li li ), gi b p, il gp iz bi. Wl (12) bi p «g---ql» 13 C wi «l-» 6 / pi i wil B & Wll (17) gg ii p f 6 C f i fl ii. B, lg piz li w pibl f iff b g i ll ifi ll, Nil (17) w l w i ff i ppli gl. Pll (16) f qi b w piil l wi p gil l wi wi li li ii i lk. Cl (2000) p i f wil i i Nw b piil l iig p gil l wi i ig l (Tbl 3). Dii l i b i iigig ifl f ll glbl w ii pl, w i i ff ppli p f gl (Tbl 1). T ii ff f li g i ip gi i fi g fil i w ig i l ii p l i ll f p i pl i, pl i, wil lwig pi: i f p fg pl (J, 186; Li & H, 187; B0 & Hjlj, 11). T i g ff ilbili f f p pi, il ii pi pi i pii f g Tf, il ii w b igifi ifl f igl pi gl ig lw ii ( & Dl, 18). V Wl (2000) gg li f 44 pl qi ig b i b lw ll f ilbl pl bi ig fg qli ig gwig i ig i, w i p gi wi ig fg bi, li f pl qli b f g ip f bi. I lb i, f, li f ig pl bi i likl l fbl ig g li f ig pl qli (V Wl, 2000). lg xi f ff f w p (ll glbl) gl ppl i i i wil p, i lil ii (pii gi) f i ff wi ibl ip. f ig l pbbl i f lgil iff, fil f igifi i, i (.g. M, 2000 i i b). F xpl, M (187) li gi ff f pi wi' w f ppli i i wi-il. Mi (11) lig i f p ff f w li ppl i i. P & (18) l b igil i f wi-il ig f il li ig NO (wi li wi wfll) f f i f wi-il i b igil w ifl b glbl lii fli 2-3- lg. Hw, Li (1) p ppi ff f i i pig p i i lp P, wig iff ll p glbl lii ibil i i-pifi ll ig lg. Ti gg ig p li b ll xpli b lgil pb T i ibl bilgil ii i p li bw gi,, llii, bw pi wii pi. l, i ig NO ll w i liip wi ll w ibl f i ll i. T i i l lil g g liz li ip f pifi lii ibl i iflig gl ppli p. T i ibl gil ll ibili i w gl p pifi li i ibl (Kli, 1). Ti plxi f il/ liip l gill, lll Rgif, 22 (1), 2002

13 b pi ppli (x, g iz f xpl). Ti gg w l q i f giig i f pifi w ibl b i i fil, iill i i f pibl fi li g ibl f l il (.g. Liö, 16). i/ib ppi I iw f i b, i i-ll il ip, i f i f lg i i i ig ff f li i ibili i/ib. I, iig ppl, ilig Gwi'i f lk w C, Ii f C, lk Gl, i Ki f Fi w Ri, ppl f ibi Ri F E p i/ib f i lili. Ri i Nw xpi lg l lii ii wi ip pi p, b li wig f l l l i i wi lii ii (Riifflig, 18; R.B. Wlji & 0. Hl, pb l). F, pbbl g g gl, i i pii, wi i l w ii, i f pi l ip i i lg (Möl & Kli, 17; Kli, 1; Cl, 2000). T l fgig i i i b i i f fig i Rgif (Ri, 180). I i pp i ig w il ff b ii pii f i ib g l fgig i, i g xpi i pi b. Ml iigig ff f li g i pi i i i i, i i g l p (G & kgl, 17; B & Wll, 17). Ti lll, w, b ll f p ig wi li, i i wiwp (Wl, 12). O f iflig i ll i il l (Cl, 2000), li ii «i lif» bi (Pll, 16; Möl & Kli, 17). T ff f qi, biig fli, wbl fli l b fli i ib iffill i li w ii. Ti i l iff iig f g Rgif, 22 (1), 2002 f l p f Dip pi, i lif p i ii flig pii iff p, ii wi ii ( Rb, 178; Rl, 13; Möl & Kli, 17 b). w, g i ff i ii pl il l i/ib ii ig, p big k f. I, i b ii g l ff f i i ib, p il pp pl ip l i iig ll p f i bi f i ib, i ii b f i (Möl & Kli, 17). T xp i p wi gig glbl li g l, f, b fl f i g W pi lii f pbili bw iffil f glizig b ff f li g bl, pi wii pi. T pj i p i ii p f Ei i N l f l i il ff i ib g i ff i ii w p, wi li l i ii pppi. T plx l f i i lg f i ib, i ifl f w p lg f i, f li ibili ppli i f i ib pill iffi E l i kw gig pibl ff f pii gp. Hl (1) p i f ii f pii i lb i ig i wi. igl, glbl g pi ff wi li, pii il ii i wi i wi p. T liip bw li pi i gl l f b f T ffilii bw i/ib li i g (Bj, 10; Ci, 2000). Pipii i f j p li i li (Ci, 2000). T, i pipii ig wi pi b li g l (Diki, 186; Mxwll, 17), l ip i/ib g ig wi. B i p f pl p ip wi ii, pii g f li b, lig li ikg i f -fig 45

14 li, wi ig b il f i ib. Pii b pibl ff f gig glbl wig b wi i, b will likl b bl, lg i bl bw iff ggpi. kwlg W gfll kwlg fiil pp f Nwgi Ri Hb Dlp F R.B. Wlji Ø. H T xp gi w Cipl PD Nwk i i Eil i w llw R.B. Wlji piip i i «Ri K pi i N» i pb 2000, w i wk w i i i i. W l gfl iw O. Hjlj f kig g gi ip ip. Rf, R., æ, B. E. & Øil, N Fli f i ppli f lb i: ff f i p lii ii. Egp 23: k, E. & Lik, D Wig lp i g i. I: Ri, E., G, E., & kjbg,. (.). Pig 2 I. Ri Cib p. Rø, Nw, 17. Dik f il g fkfik, Ti, pp , L. G. & Dl, B. W. 18. Rpi pf f fl lk Cib. J. Wil Mg. 62: , J. R. & Nil,. C. 16. Tppig i pi f i: T p f Cpi p Hp i bi p. Mil Vi Elg 10: , J. R. & Nil,. C. 18. D i ggg w p b pii fli gl? Rgif 18: 3 17., J. R., Nil,. C. & Fl, I. 14. Mig bi gli f i wbl fl, Hp i L. (Dip: Oi). Jl f I Bi 7: Bll, J. P., Ei, G. & Wlli, K. 1. Cli g, i p. - Elgil Blli 47: Bg, M., Hp, J. L. & Tw, C. R. 16. Elg. 3. Blkwll i. B, O. L. & Mi, E. L Eff f lp ili il fg qli f pil g lg. I: Ril, H. & kjlg,.o. (.). T ip f li g pi qli. Pig f 10 Gl Mig f Ep Gl Fi., Nw, J, pp Bø,. & Hjlj, O. 11. D il g ip ig l? C. J. Z 6: Bj, R. D. 10. Di qli ik qi f l fl ib f Dli, lk. Jl f ppli Elg 27: B, J. & Wll, G. 17. Cli g B Cib i Nw Tii, C. Clii Cg 35: B, J. P., Cpi, F.., III & Kli, D. R Cb/i bl f bl pl i li b bi. Oik 40: Cl, W iz, fi, lif i. H Uii P, Cbig, M L, Eg C, R. D., i, W. T., F,. G., G, K. L. & Wi, R. G. 13. Clig f fl ib i li b-wig. C. J. Z 71: Cppi, N. & Pi, P. W. 15. Ppli i, w pp i. L: i P. Cl, G. N.,, H. G. & P,. 11. B i f l i li wi i. J. Wil Mg. 55: Cpi, F.., III &, G. R. 16. Pilgil gw p f i pl fil f xpi ilig lii g. Elg 77: Cp, W. L. & Wl, J. E. 13. R ii f i i-p i ig-li. Blli f i Mlgil i 74: Cl-Bk, T. H. & lb,. D Wi li i (C lp). J. Z L 18: Cl, J. E Bi p f wil i i li p pii fli. PD i. Uii f Ol, Ol, Nw. Ci, P. D p f lg f fig li. Rgif 20: Di, B Cil pii ii l f bg i li li. I: Ril, H & kjlg,.o. (.). T ip f li g pi qli. Pig f 10 Gl Mig f Ep Gl Fi., Nw, J, pp Diki, R T li llig f f li. I: Bli, B., D, B., Jg, J. & Wik, R.. (.). T g ff, li g,. Wil, Ci, UK, pp Rgif, 22 (1), 2002

15 Epk, Y. & Lg, R. 17. Lig w i glbl wig. Elgil i f ig fl i (C 124: lp). Bil Elg ibilg 5: Hl, O., i,., Ii, J., Lg, R. & F,. G. & Wi, K. R. 11. li f l ig g b Ppi ib. C. J. Z 6: F-Bi, M., Uq, M. & i, K. G. 14. Mi g i: ki pi il big g. C. J. Z 72: Fi, G. L., Bg, M., Kill, K. & Pi,. K Clii ifl fg qli, gw pi f i w Pil I: li fg qli. I: Pig f 8 N i Cib Wkp, Wi, Yk, C. Rgif pil I N. 12: 144. Fi, G. L., Bg, M., Kill, K. & Pi,. K. 2000b. Clii ifl fg qli, gw pi f i w Pil II: Ri Gw Rpi. I: Pig f 8 N i Cib Wkp, Wi, Yk, C. Rgif pil I N. 12: 144. F, M. C.,, N. C., P, E. & Lg, R. 18. Ppli Di Of Nwgi R D: Di-Dp Clii Vii. Pig Of T Rl i Of L i B-Bilgil i 265: Gill, J. M., F-Bi, M. & Yz, N. G. 18. Ppli i f lg bi: i bl i wi l i T i Elg Eli 13: Gill, J. M., p,. J., Bi, J. M., V L, G. & Bib, B. 12. Eff f g b-wig ppi f fl big i ppli f (Cpl pl). C. J. Z 70: Gill, J. M., Bi, J. M., Dl, D., V L, G., D, P. & Lb, J. D. 17. El il i : q f ii i w ppli. Olgi 112: Gill, J. M., DL, D., Bi, T. H., V L, G., Bib, G. & Pl, R. 13. R il p: pi li f ig ppli. J. i. Elg 62: G, K. L., Wi, R. G., C, R. D. & Rll, D. E. 16. B pii i f i ib. C. J. Z 74: Gbb, P Ppli i f p. I: Jwll, P.., Mil, C. & M, B.J. (.). Il i: lg f p f. Ki l P, L, UK. G,. & kgl, T. 17. Rp f ib lb,. 1. Ei f i ii f pii i i ig i wi. Iil Jl f Pilg 2: Hi, O. M Pilgil p f lii pi i pl wi pil f igli i. I: Ki, Å., Jil, O. & Nil, J. (.). Pl pi i. Pig f «Pl pi wkp». Tø, Nw, pb 4-. Nwgi Uii P, Ol, pp Hll, T Fgig bi f i- i i (Rgif L.) i li w i Fii Lp K bi R i Rp 1, pp Hll, T., pi, J., Lp, K. & Tki, E. 12. gi i biig fli b i: fil xpi wi il p. l Zlgii Fii: 2: Hwi,. J. M. & Gill, J. M. 1. fl g g? T Ti-Will l x-bi l i i g T i Elg Eli 14: Hjlj, O. & Hiøl, T. 1. Rg-b i i f gl f pi. Olgi 11: Hg, L Bilgil q f glbl wig: i igl l pp? T i Elg Eli 15: Hll, J. W. 15. Dl i N li Oilli: gil p pipii. i 26: Hll, J. W. 16. Ifl f ii i x pil wii li i p p. Gp. R. L. 23: Hll, J. W. & V L, H. 17. Dl i i i li i wi N li illi. Clii Cg 36: J, O. W., B, J. P., Cpi, F.., III &, M Pl pl i i li ii i li gzig. i Nli 128: Jg, J. T., F-Bi, M., Gill, J. M. & Wi, W. D. 17. Eff f g, x, i, i il f big p. Elg 78: Jg, J. T., F-Bi, M., Lii, M. & Wi, W. D. 13. Eff f b-iz, ppl i-i, l ii g fi pi i big w. C. J. Z 71: Ji', T. & R, L Ifl f lii Rgif, 22 (1),

16 ii il qli f gl i Ei. I: Ril, H. & kjlg,.o. (.). T ip f li g pi qli. Pig f 10 Gl Mig f Ep Gl Fi., Nw, J. Kli, D. R T g f bl f. Jl f Rg Mg 23: Kli, D. R. 170b. F li b N i i p -ilizi f pf pl pi. I: W,. (.). il ppl i i li i f. Blkwll i Pb, Oxf, UK, pp Kli, D. R Ppli lg: ii bw i f ppl. I: F, P.F. & Dw, K.R. (.). Bilg f pi. T Rl i f Nw Zl, Wllig, Nw Zl, pp Kli, D. R. 11. Cib i gig N. ppli il Bi i 2: Kli, D. R. 1. T l f li ili i bli pi f Rgif ppli i ig i. Elgil Blli 47: Kjl, I. 11. Rpi iffil p kig i i fl i. ppli il Bi i 32: 1-3. Kk, L. E. B., Cl-Bk, T. H., lb,. D., Pb, J. M. & Gi, F. E. 1. Ppli i ff x i ii i. N 3: Li, K. M. & H, H Pli/ig i i blb Vii ill i l i p ii i i Fii Lp Oik 50: Lg, R., lb,. D., Bk, T. & Cl- Bk, T. H. 16. Cli, pl plg ii i g fi pi i p bi. J. i. Elg 65: L,. E., P, M. C., L, J.., Igl, T. & Kil, T Rgil ff f li g i: f Mki gi i Fii Lp Pl R 1: Lik, D., B0, E. & Fjlli, Rliip bw wig f i l i i ' g wig i pi pig. Rgif 8: Li, J Rp f pl il. V 1. i P, Nw Yk. Li, J. 16. W g ppli i. Willif Bilg 2: 3-. Li,. & Lg, R. 18. lg ff f wi pig w gw il f i Nw. Olgi 116: Li,., Jli J. M. & M, P. 1. Rliip bw i i il ii i glbl ll li gi: ig xpl f lp P. Elgil Blli 47: Li,., Lg, R. & lbg, E. J. 1b. B wi li i l: i glig x li ff. Egp 22: Mik, C. J Mli f M pg i wi. J. Wil Mg. 31: Mxwll, B. 17. R li p i i. I: Ol, W.C., Cllg, T., Gil, T., Hl, J.I., Mxwll, B., Ml, U. & ibj, B. (.). Glbl g i il. pig, Nw Yk. M, L. D., MRb, R. E., P, R. O. & Pg, R. E Rliip f ppli pi wi' w. J. i. Elg 56: Mi, F. 11. T igifi f liiig g lig f gp f wiil. J. i. Elg 60: Mil, J. M., El, D.. & lb, D. 1. Eiig ibi f ppli i lii fli il ii i il f p. J. i. Elg 68: M,. & ig, C. W T l wig l il f wil-il i Nw Yk. Nw Yk Fi G J. 28: Möl, F. M. 1. U f lii l p f i fli i ib. J. Wil Mg. 63: Möl, F. M. & Kli, D. R. 17 Eff f w pii i bi gp i f ib f Dl H, lk. C. J. Z 75: M, W. W. 14. Ppli gli i pi. Elg 75: M,., Yz, N. G.,, N. C. & Lg, R Rliip bw x i, li i i : ip f pil J. i. Elg 6: Nil,. C. 17. Eff f p ppl lp li i i i Hp i Cpi p (Dip: Oi). Ei. E 26: Nil,. C. & Hg, R. E. 14. T iig p f l f wbl fl Hp (= Og) i (L.) b fl Cpi p (M) (Dip: Oi) f i. Rgif 14: Nl, L. E., Pll, R. H., Bll, W. B. & Ci, M ii f i gl p 48 Rgif, 22 (1), 2002

17 b Cib, Rgif gi, wi i P B il fil, lk. Ci Fil-Nli 112: Oll, J. P., H, D. C., Bi,. & Ml, R. 17. pi f b ii p i f ib w p-f i i C. J. Z 71: Pk, K. L. & Rbbi, C. T Tgli i g I: H, R.J. & Wi, R.G. (.). Bigi f wil bi. CRC P, pp ii ppi. Rgif 17: Pk, K. L., Rbbi, C. T. & Hl, T Ri, E. 1. Fl x i i wil i Eg xpi f li b l Rgif i li l ii lk. J. Wil Mg. 48: g. Wil Bi 5: Pk, K. L., Gillig, M. P., Hl, T.. & Ri, E., Kli, D. R. & øgå, R Rbbi, C. T. 13. l p i b Clig i, gw b iz f pii w f f f gig Nwgi i iff g. i pi blk-il (Oil i lpi R 15: iki) i lk. C. J. Z 71: Pll, R. H., Bll, W. B., Nl, L. E. & Ci, M Pii i b ili f gl p wii P B il fil, lk, i li b Cib, Rgif gi. Ci Fil-Nli 110: P, E. & Kli, D. R. 1. Cib lf pi l g qli ig ppli li. J. Wil Mg. 63: P, E. &, N. C. 18. Lg-l lii fli ppli i f wi-il. J. i. Elg 67: P, E. &, N. C. 1. Clii ibili, pl plg, g Elg 80: P, E., F, M. C. &, N. C. 1. Ppli lg N li Oilli (NO). Elgil Blli 47: P, E., F, M. C.,, N. C. & Lg, R. 1b. Exii ifii f b x i b lii g. i Nli 154: P, E., Lg, R., F, M. C. &, N. C. 1. Eil ii p xl ipi i. Pig f P, R. J., Lgbi, J., Hwi,. J. M. &,. K. 16. Rli l f ip i-ip f i pp li i f Bii. Ml Riw 26: Rgif, 22 (1), 2002 Ri, E ii p; j i f gw fig i Rgif? I: Ri, E., G, E., & kjbg,. (.). Pig 2 I. Ri Cib p. Rø, Nw, 17. Dik f il g fkfik, Ti, pp Ri, E Gw b iz iff i Rgif, f ff. Rgif 3: Ri, E. 17. Rgif ppli lg: Riifflig. 18. Rgkp f iifæig. 181 pp. Rb, D. D Bil p f b-g ib f Cl i H j T lk il pipli. M.. i, Uii f lk, Fibk. Rg, J. C T ii bw N li Oilli Oilli i ip. Ml W Riw 112: Rll, D. E. 13. Eff f glbl wig bilg g f Ppi Cib H. I: Wll, G. (.). Ip f li g g i N. Dp f Ggp, Uii f Wl, pp æ, B. E l ii i wig f Nwgi i li li lg l iil gi. J. Wil Mg. 4: æ, B. E. 17. Eil ii ppli i f lg bi: f i. T i Elg Eli 12: æ, B. E. & G,. J l ii i wi b ii f Nwgi l. J. Wil Mg. 52: æ, B. E.,, R., Hjlj, O. & Hi, M. Nil f i, U 6: Elgil l f gil ii i P, E.,, N. C., Lg, R. & Fi, lif i f lg bi, l J. M. 17. Glbl li g ppi Elg 77: ii g. Pig Of T, H., P, Å. & Cl, G. 16. l Rl i Of L i B-Bilgil i 264: ii i b pi i i (l l). - T ff f w ii f qli. I:, H., 16. Lif ii i (l l): ggpil pl ii i b gw pi. PD i, wi Uii f gill i, pp , J. R. & B, M Di p 4

18 il f lk i w Wig. J. Wil Mg. 47: z, M. C., R, M. C., B, R. G., Wl, J. E. & Rbi, D Vii i lgill i i pipii wfl. Gli 21: kgl, T T ff f i p liii iz f wil i. Olgi 60: kgl, T T ff f i-p liii gp f wil i. J. i. Elg 54: kgl, T. 18. Cpi il gizi f i i li f, p i. i Elg 2. bjø, T., æ, B. E., lbg, E. J., Hi, M. & Rl, C. M M (l l) il i ppli i Nw. C. J. Z 78: kkii, E. E. 15. Wil i. ii Iii Lib, Wig, DC. Tiz, L. & Zig, E. 1- Pl pilg. 2 ii. i i, I., Pbli. Tpi, B., H, J. & M, M. 16. Eff f i bi f Riiè Gg ib. i 4: Ti, P. 15. Ppli gli: l g w i. I: Cppi, N. & Pi, P.W.(.). Ppli i. i P, pp V Wl, R., M, N., Li,., D, C., Lg, R. & lb,. D Tig fg qli f qi? Pl plg p i b lb i. Olgi 123: V, P. J. 14. Niil Elg f Ri. ii. Wl, N. E., F,. G., MCb, T. R. & Pk, L. F. 12. Hbi b ppi ib ig pi i. J. Wi Mg. 56: Mip i 22 Db, 2000 p 25 pb, Rgif, 22 (1), 2002

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