Why are some plant–pollinator networks more nested than others?

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Why are some plant–pollinator networks more nested than

others?

Chuliang Song

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Abstract

Ɛĺ lrbub1-Ѵv|7b;v_-;=om7|_-||_;l|-Ѵbv|b1bm|;u-1|bomv=oulbm]|_;v|u1|u; o= rѴ-m|ŋroѴѴbm-|ou m;|ouhv -u;|rb1-ѴѴ lou; m;v|;7 |_-m ;r;1|;7 0 1_-m1; -Ѵom;ĺ77b|bom-ѴѴķ|_;ou;|b1-Ѵv|7b;v_-;v_om-rovb|b;-vvo1b-|bom0;|;;m |_; m;v|;7 v|u1|u; o= l|-Ѵbv|b1 m;|ouhv -m7 1ollmb| r;uvbv|;m1;ĺ +;|ķ b| _-v 0;;m v_om |_-| vol; rѴ-m|ŋroѴѴbm-|ou m;|ouhv l- 0; lou; m;v|;7 |_-m o|_;uvķu-bvbm]|_;bm|;u;v|bm]t;v|bomo=_b1_=-1|ouv-u;u;vromvb0Ѵ;=ouv1_ ;m_-m1;7m;v|;7v|u1|u;ĺ

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Ɠĺ lrou|-m|Ѵķouo0v;u-|bomv1olrѴ;l;m||_;ouv_obm]|_-|lou;m;v|;7m;|Ŋ ouh v|u1|u;v 1-m bm1u;-v; |_; u-m]; o= ;mbuoml;m|-Ѵ 1om7b|bomv 1olr-|b0Ѵ; b|_vr;1b;v1o;bv|;m1;bml|-Ѵbv|b1vv|;lvķ-Ѵvohmom-vv|u1|u-Ѵv|-0bѴb|ĺ $_bvbm1u;-v;bmm;v|;7m;vvv_oѴ70;lou;-7-m|-];ov-m7o11ulou;o=|;mbm Ѵo1-|bomvv0f;1||ou-m7ol;mbuoml;m|-Ѵr;u|u0-|bomvķ_b1_1oѴ70;7ub;m0 _b]_Ѵ1_-m]bm]ouv;-vom-Ѵ;mbuoml;m|vĺ$_bvvm|_;vbvo=|_;ou-m7o0v;u-Ŋ |bomv1oѴ7ruo;u;Ѵ;-m|=ou-0;||;um7;uv|-m7bm]o=|_;;1oѴo]b1-Ѵruo1;vv;v 7ubbm]|_;-vv;l0Ѵ-m7r;uvbv|;m1;o=;1oѴo]b1-Ѵ1ollmb|b;vĺ  + ) ! " 1_-m]bm];mbuoml;m|vķm;v|;7m;vvķm;|ouh1olr-ubvomķv;-vom-Ѵb|ķv|u1|u-Ѵv|-0bѴb|ķ zŊv1ou;v

http://doc.rero.ch

Published in "Journal of Animal Ecology 86 (6): 1417–1424, 2017"

which should be cited to refer to this work.

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Number of species (log scale)

Nestedness (NODF) 101 ₁₀1.5 ₁₀2 ₁₀2.5 (b) r = –.71 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.0 0.2 0.4 0.6 0.8 1.0 Maximum nestedness Nestedness (NODF) (c) r = .37

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(4)

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Standard deviaiton from

randommization (a) 0.002 0.004 0.006 0.008 0.010 0.305 0.315 0.325

Maximum value of acceptance (

z = 2)

(b)

(5)

1omvbv|;m| -1uovv m;|ouh vb; -m7 1omm;1|-m1;ĺ $_bv v_ov |_-| - 1ol0bm;7 v|-|bv|b1 1-m 0; - lou; u;Ѵb-0Ѵ; l;-vu; |_-m zŊv1ou;v |o 1olr-u;m;v|;7m;vv-1uovvm;|ouhvĺlrou|-m|Ѵķ|_;1omvbv|;m1o= |_bv1ol0bm;7v|-|bv|b1-ѴѴovoubm;v|b]-|bomo=|_;ro|;m|b-Ѵ-vvo1bŊ -|bom0;|;;mm;v|;7m;vv-m7|;lr;u-|u;v;-vom-Ѵb|ĺ

bm-ѴѴķ |o |;v| |_; _ro|_;vbv |_-| m;v|;7m;vv v_oѴ7 bm1u;-v; m7;u1_-m]bm];mbuoml;m|-Ѵ1om7b|bomvķ;bm;v|b]-|;7|_;-vvoŊ 1b-|bom0;|;;m|_;1ol0bm;7m;v|;7m;vvv|-|bv|b1Ő cő-m7|_; Ѵ;;Ѵo=|;lr;u-|u;v;-vom-Ѵb|Ő-ub-m1;o=7-bѴ|;lr;u-|u;őbm7b=Ŋ =;u;m|Ѵo1-|bomv-uom7|_;ouѴ7ĺ$_;Ѵ-|b|7;-m7Ѵom]b|7;o=|_; o0v;u;7rѴ-m|ŋroѴѴbm-|oum;|ouhv;u;o0|-bm;7=uol|_;r0Ѵb1u;Ŋ rovb|ouweb-of-life.esĺmbuoml;m|-Ѵ7-|-;u;o0|-bm;7=uol |_;r0Ѵb1u;rovb|ouWorldClimŐ;uvbomƐĺƓķu;Ѵ;-v;ƒķu;voѴ|bomƑĺƔ lbmőĺ&vbm]|_;v;u;rovb|oub;vķ;]-|_;u;7;mbuoml;m|-Ѵ7-|-=ouƓƒ o|o=|_;ƔƖm;|ouhvĺ);7b7mo|=bm7bm=oul-|bom-0o||_;o|_;u Ɛѵ m;|ouhv Őm;|ouh -m7 |;lr;u-|u; 7-|- -u; ruob7;7 bm "om] ;|-ѴĺķƑƏƐƕőĺ$ov|7|_;7bu;1|u;Ѵ-|bomv_br0;|;;m|_;1ol0bm;7 m;v|;7m;vvv|-|bv|b1-m7|;lr;u-|u;v;-vom-Ѵb|ķ=buv|;v;7-vblŊ rѴ;1ouu;Ѵ-|bom-m-Ѵvbvĺ;1-v;b|bvmo|;r;1|;7|_-|omѴom;=-1|ou Ő;ĺ]ĺ |;lr;u-|u; v;-vom-Ѵb|ő v_oѴ7 r;u=;1|Ѵ ;rѴ-bm |_; v|u1|u; o= ;1oѴo]b1-Ѵ m;|ouhv Ő$upf;Ѵv]--u7 ş Ѵ;v;mķ ƑƏƐѵőķ ; -Ѵvo v;7 -v|-m7-u7rubm1br-Ѵ1olrom;m|-m-ѴvbvŐĸ;];m7u;ş;];m7u;ķ ƑƏƐƑőĺ$_bv-rruo-1_-ѴѴo;7v|oruof;1||_;lѴ|b7bl;mvbom-Ѵ-ubŊ -|bomo=|_;7-|-v;|bm|o=;;u7bl;mvbomvŐrubm1br-Ѵ1olrom;m|vőĺ); v;7|_;1ol0bm;7m;v|;7m;vvv|-|bv|b1ķѴo]Ő"őķ1omm;1|-m1;-m7|;lŊ r;u-|u;v;-vom-Ѵb|-v-ub-0Ѵ;vbmouĺ$_vķ|_;bѴѴv|u-|;v |_; 1ouu;Ѵ-|bomv 0;|;;m |_;v; -ub-0Ѵ;v -m7 _o |_; ;rѴ-bm |_; -ub-m1;vbm|_;7-|-ĺ77b|bom-ѴѴķ;1olr-u;7|_;1ouu;Ѵ-|bom-m7 u;vѴ|v-]-bmv||_;u;vѴ|v=uolv|-m7-u7b;7-Ѵ;vo=m;v|;7m;vv ŐzŊv1ou;vőĺ

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o1vbm] om |_; vblrѴ; 1ouu;Ѵ-|bomv 0;|;;m |_; m;v|;7m;vv v|-|bvŊ |b1v-m7|;lr;u-|u;v;-vom-Ѵb|ķ b]u;Ɠ-ķ0v_ov|_-||_;zŊv1ou;v 7;ub;7=uol|_;ruo0-0bѴbv|b1-m7|_;=b;7lo7;Ѵv_-;m;-uѴ-mѴѴ 1ouu;Ѵ-|bom ŐrƷĺƏƑ -m7 ƴĺƏƕķ ;-uvom 1ouu;Ѵ-|bomőĺ m 1om|u-v|ķ |_; =b]u; Ő b]u;Ɠ1ő v_ov |_-| |_; 1ol0bm;7 m;v|;7m;vv v|-|bv|b1 _-v - rovb|b; 1ouu;Ѵ-|bom b|_ |;lr;u-|u; v;-vom-Ѵb| ŐrƷĺƑƕķ ;-uvom 1ouu;Ѵ-|bomőĺo|;|_-|0o|_|_;o|Ѵb;uv-m7|_;Ѵo1ouu;Ѵ-|bom-um 1-|bom om bm|;uru;|bm] |_bv u;vѴ|ĺ ou ;-lrѴ;ķ u;lobm] robm|v Ѵ-u];u|_-m|oSDv--=uol|_;l;-mom0o|_-ub-0Ѵ;vl-h;v|_bv rovb|b;1ouu;Ѵ-|bommomŊvb]mb=b1-m|ĺ

$o =u|_;u bm;v|b]-|; |_;v; u;Ѵ-|bomv_brv -0o;ķ ; v_b=| ou =o1v |o |_; ĺ b]u;Ɣ v_ov |_; |o rubm1br-Ѵ 1olrom;m|v 7;ub;7 =uol |_; =ou-ub-0Ѵ;vĹ m;v|;7m;vv v|-|bv|b1 ŐzŊv1ou;v -m7 1ol0bm;7 m;v|;7m;vvőķ ml0;u o= vr;1b;v ŐѴo] o= |_; ];ol;|ub1 l;-mo=rѴ-m|v-m7roѴѴbm-|ouvőķ1omm;1|-m1;-m7|;lr;u-|u;v;-Ŋ vom-Ѵb|ĺ$_;|o1olrom;m|v-11om|;7=oucĺƕƔѷo=|_;-ub-m1; bm|_;7-|-ĺ$_;=b]u;Ő b]u;Ɣ-ķ0őv_ov|_-|vbm]zŊv1ou;vķ|_; m;v|;7m;vvv|-|bv|b1bv_b]_Ѵ1ouu;Ѵ-|;7b|_vb;-m71omm;1|-m1; =ou|_;ruo0-0bѴbv|b1-m7=b;7lo7;Ѵvu;vr;1|b;Ѵĺ$_bv-vvo1b-|bom 1-m0;o0v;u;70mo|b1bm]|_;v-l;7bu;1|bomo=|_;bu1ouu;vrom7Ŋ bm];1|ouvĺ o| vurubvbm]Ѵķ 0o|_ zŊv1ou;v;u; roouѴ 1ouu;Ѵ-|;7 b|_|;lr;u-|u;v;-vom-Ѵb|ķbĺ;ĺ|_;;1|ouvo=|_;m;v|;7m;vvv|-Ŋ |bv|b1-m7|;lr;u-|u;v;-vom-Ѵb|;u;rovb|b;Ѵ1ouu;Ѵ-|;7omom; 1olrom;m|0|m;]-|b;Ѵom|_;o|_;uĺ$_;v;u;vѴ|v1om=bulru;Ŋ bovouhv_obm]|_-|zŊv1ou;v7;ub;7=uol7b==;u;m|mѴѴlo7;Ѵv Őruo0-0bѴbv|b1-m7=b;7lo7;Ѵvő1-m];m;u-|;7b==;u;m|r-||;umv-m7 mo mb=b;7 -mv;uv Ő";0-v|b࢙mŊom࢙Ѵ; ;|-Ѵĺķ ƑƏƐƔĸ $-h;lo|o ş -fb_-u-ķƑƏƐѵőĺ & ! ƒ Պ olr-ubm]m;v|;7m;vv-1uovvb7;m|b1-ѴѴ];m;u-|;7m;|ouhvĺ ou;mv;l0Ѵ;vo=ƐķƏƏƏu-m7olѴ];m;u-|;7m;|ouhvb|_|_; v-l;mv|u1|u;7l;1_-mbvlŐvbm]|_;;tbruo0-0Ѵ;lo7;Ѵőķ|_;=b]u;v_ov|_;-;u-];m;v|;7m;vvv|-|bv|b1Őom-Ѵo]v1-Ѵ;=oubv-Ѵb-|bom rurov;vő-v=m1|bomo=7b==;u;m|-Ѵ;vo=1omm;1|-m1;ŒƏĺƐĸƏĺƓœŐxŊ-bvő-m71ollmb|vb;ŐƑƏķƓƏ-m7ѵƏvr;1b;vőĺ$_;0Ѵ;vt-u;vķou-m]; 1bu1Ѵ;v-m7]u;;m|ub-m]Ѵ;v1ouu;vrom7|o|_;m;v|;7m;vvv|-|bv|b1vbm]|_;zŊv1ou;b|_ruo0-0bѴbv|b1lo7;ѴķzŊv1ou;b|_=b;7lo7;Ѵ-m7 |_;1ol0bm;7m;v|;7m;vvu;vr;1|b;Ѵĺo|;|_-|omѴ|_;1ol0bm;7m;v|;7m;vvv|-|bv|b17bvrѴ-v-lou;1omvbv|;m|l;-vu;-1uovvm;|ouh 7bl;mvbomvĺѴѴ;uuou0-uv-uom7|_;-;u-];-Ѵ;;u;m;]Ѵb]b0Ѵ;ĺ;1-v;|_;m;v|;7m;vvv|-|bv|b1=ou|_;=b;7lo7;Ѵbvm;]-|b;ķ;v;7 Ѵo]ŐzŊv1ou;ƳƐƏő=oub|vbv-Ѵb-|bomŒoѴou=b]u;1-m0;b;;7-|bѴ;omѴbm;Ѵb0u-uĺ1olœ

20 Species 40 Species 60 Species

0.1 0.2 0.3 0.4 0.1 0.2 0.3 0.4 0.1 0.2 0.3 0.4 102.5 102 101.5 101 100 100.5

Connectance

Nestedness statistic (log scale) Probabilistic Fixed Combined

Connectance

http://doc.rero.ch

(6)

lrou|-m|Ѵķ b]u;Ɣ1v_ov|_-||_;1ol0bm;7m;v|;7m;vvv|-|bvŊ |b1_-v-v|uom]1ouu;Ѵ-|bomb|_|;lr;u-|u;v;-vom-Ѵb|ĺ$_-|bvķ|_; ;1|ouvo=m;v|;7m;vv-m7v;-vom-Ѵb|v_ovblbѴ-u7bu;1|bomvom0o|_ 1olrom;m|vĺ o|; |_-| 1ol0bm;7 m;v|;7m;vv v|-|bv|b1 -m7 |;lr;u-Ŋ |u;v;-vom-Ѵb|-u;m;-uѴou|_o]om-Ѵ|o1omm;1|-m1;-m7vb;ĺ$_;v; t-Ѵb|-|b; u;vѴ|v 7b7 mo| 1_-m]; 0 u;lobm] |_; o|Ѵb;uv 7;=bm;7 -0o;=uol b]u;Ɠ1Őv;; b]u;"Ƒőĺ$_;v;u;vѴ|v|_;mvrrou||_; _ro|_;vbv|_-|lou;m;v|;7v|u1|u;vo=rѴ-m|ŋroѴѴbm-|oum;|ouhv Ő_;m -11om|bm] =ou |_; bm|ubmvb1 ;==;1|v o= 1omm;1|-m1; -m7 1olŊ lmb|vb;őv_oѴ70;=om7bmlou;v;-vom-Ѵou;mbuoml;m|-ѴѴѴ;vv ru;7b1|-0Ѵ;Ѵo1-|bomvĺ

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1;m|u-Ѵ Ѵbm; o= u;v;-u1_ om |_; v|7 o= l|-Ѵbv|b1 1ollmb|b;v _-v0;;m|om7;uv|-m7|_;;|;m||o_b1_|_;v|u1|u;o=rѴ-m|ŋ roѴѴbm-|ou m;|ouhv 1-m lo7Ѵ-|; |_; Ѵbh;Ѵb_oo7 o= vr;1b;v r;uvbvŊ |;m1;Ő-v1olr|;şou7-moķƑƏƐƒőĺ$_;ou;|b1-ѴѴķb|_-v0;;mv_om |_-||_;m;v|;7v|u1|u;o=l|-Ѵbv|b1m;|ouhv1-m_-;blrou|-m| 1omv;t;m1;v =ou 0bo7b;uvb| l-bm|;m-m1; Ő-v|oѴѴ- ;|-Ѵĺķ ƑƏƏƖĸ !o_u ;|-Ѵĺķ ƑƏƐƓĸ "--;7u- ;|-Ѵĺķ ƑƏƐƒĸ "--;7u-ķ !o_uķ Ѵ;v;mķ ;|-Ѵĺķ ƑƏƐѵőĺ $_vķ v|7b;v _-; 0;;m bm;v|b]-|bm] _b1_ ;1oѴo]b1-Ѵ -m7 ;mbuoml;m|-Ѵ 1om7b|bomv 1oѴ7 0; -vvo1b-|;7 b|_ -m bm1u;-v;

& ! Ɣ Պ 7;m|b=bm]|_;7bu;1|bomo=1_-m];0;|;;mm;v|;7m;vv-m7|;lr;u-|u;v;-vom-Ѵb|ĺ&vbm]-rubm1br-Ѵ1olrom;m|-m-Ѵvbv Ő;];m7u;ş;];m7u;ķƑƏƐƑőķ|_;=b]u;v_ov|_;|orubm1br-Ѵ1olrom;m|v=ou|_;=ou-ub-0Ѵ;vbm;v|b]-|;7Ĺml0;uo=vr;1b;vŐѴo]o= |_;];ol;|ub1l;-mo=rѴ-m|v-m7roѴѴbm-|ouvőķ1omm;1|-m1;ķm;v|;7m;vvv|-|bv|b1-m7|;lr;u-|u;v;-vom-Ѵb|ĺ$_;-uuov1ouu;vrom7|o|_; =ou-vvo1b-|;7;b];m;1|ouvķ-m7;-1_vl-ѴѴ7o|1ouu;vrom7v|oom;o=|_;Ɠƒo0v;u;7rѴ-m|ŋroѴѴbm-|oum;|ouhvŐb|_;mbuoml;m|-Ѵ7-|-őĺ o|;|_-||_;0o||ol-m7Ѵ;=|-;v1ouu;vrom7|o|_;v1-Ѵ;7-Ѵ;vķ_;u;-v|_;|or-m7ub]_|-;v1ouu;vrom7|o|_;;1|ouѴo-7bm]vĺŐ-ŋ1ő |1ouu;vrom7|o|_;m;v|;7m;vvv|-|bv|b1vbm]|_;zŊv1ou;b|_ruo0-0bѴbv|b1lo7;ѴķzŊv1ou;b|_=b;7lo7;Ѵķ-m7|_;1ol0bm;7m;v|;7m;vv u;vr;1|b;Ѵĺo|;|_-|omѴ|_;1ol0bm;7m;v|;7m;vvv|-|bv|b1Ő1ő7bvrѴ-v-v|uom]o;uѴ-rb|_|;lr;u-|u;v;-vom-Ѵb|ķ-m70o|_-u;-Ѵlov| ou|_o]om-Ѵ|o1omm;1|-m1;-m7ml0;uo=vr;1b;vŒoѴou=b]u;1-m0;b;;7-|bѴ;omѴbm;Ѵb0u-uĺ1olœ −0.2 −0.1 0.0 0.1 0.2 0.3 Comp.1 Comp.2 −5 0 5 Nestedness (probabilistic) Connectance Species T_Seasonality (a) Comp.1 Comp.2 −5 0 5 Nestedness (fixed) Connectance Species T_Seasonality (b) −0.4 −0.2 0.0 0.2 0.4 −4 −2 0 2 4 6 −0.4 −0.2 0.0 0.2 0.4 − 4 − 2 0246 −0.4 −0.2 0.0 0.2 0.4 −0.2 −0.1 0.0 0.1 0.2 0.3 −0.2 −0.1 0.0 0.1 0.2 0.3 Comp.1 Comp.2 −5 0 5 −4 −2 0 2 4 6 Nestedness (combined) Connectance Species T_Seasonality (c) & ! Ɠ Պ vvo1b-|bomo=|;lr;u-|u;v;-vom-Ѵb|b|_m;v|;7m;vvv|-|bv|b1vĺ ouƓƒo0v;u;7rѴ-m|ŋroѴѴbm-|oum;|ouhvŐb|_ ;mbuoml;m|-Ѵ7-|-őķŐ-ŋ1őbѴѴv|u-|;|_;-vvo1b-|bomo=|_;v1-Ѵ;7|;lr;u-|u;v;-vom-Ѵb|ŐxŊ-bvőb|_|_;m;v|;7m;vvv|-|bv|b17;ub;7=uol |_;ruo0-0bѴbv|b1lo7;Ѵķ|_;=b;7lo7;Ѵ-m7|_;1ol0bm;7m;v|;7m;vvŐv1-Ѵ;7-Ѵ;vőu;vr;1|b;Ѵĺ-1_robm|1ouu;vrom7v|oom;o=|_;Ɠƒ m;|ouhvĺ$_;voѴb7Ѵbm;v1ouu;vrom7|o|_;Ѵbm;-uu;]u;vvbomŐbm|;m7;7|o]b7;|_;;;őķ-m7r1ouu;vrom7v|o|_;;-uvom1ouu;Ѵ-|bomŐomѴ|_; 1ouu;Ѵ-|bomb|_1ol0bm;7m;v|;7m;vvv|-|bv|b1-vvb]mb=b1-m|-||_;Ɣѷ1om=b7;m1;Ѵ;;ѴőŒoѴou=b]u;1-m0;b;;7-|bѴ;omѴbm;Ѵb0u-uĺ 1olœ −1 0 1 2 3 0 5 10 15 Temperature seasonality (scaled values)

Nestedness statistic from

probabilistic model (a) r = .02 −1 0 1 2 3 −6 −4 −2 0 2 Temperature seasonality (scaled values) Nestedness statistic from fixed model

(b) r = –.07 −1 0 1 2 3 −2 −1 0 1 2 Temperature seasonality (scaled values)

Nestedness statistic from

combined metric (scaled values)

(c)

r = .27

(7)

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