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Transcript
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Text S1. Analytical approximation for changes in species abundances with coevolution. The
2
analyses presented in the Box in the main text can be used to both generalize the results from the
3
simulations and expose the causal pathways underlying the role of coevolution in determining
4
the changes in species abundances when climate change alters the intrinsic rate of increase of
5
one species. Consider the general model
6
N1,t+1 = N1,tF1(N1,t, 1(1,t, 2,t)N2,t, 1,t)
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N2,t+1 = N2,tF2(N2,t, 2(1,t, 2,t)N1,t, 2,t) .
(S1)
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This structure is an obvious generalization of the competition/mutualism model. It also has the
9
same structure as the predator-prey model; if N1 is the prey and N2 is the predator, then 1(1, 2)
10
= a(E, 1, 2), and 2(1, 2) = –a(E, 1, 2)c, where c is the predator conversion rate.
11
From the Implicit Function Theorem,
12
æ ¶G ö ¶G
¶X *
= -ç ÷
è ¶X ø ¶E
¶E
-1
(S2)
13
where X*=(N1*,N2*,1*,2*) is the vector of the equilibrium densities and trait values for each
14
species, and G is a vector of functions that all equal zero when population densities and traits are
15
at their equilibrium values; specifically,
16
G1(N1, 1(1, 2)N2, 1) = F1 – 1
17
G2(N2, 2(1, 2)N1, 2) = F2 – 1
18
G3(N1, 1(1, 2)N2, 1,[D11]N2) =
dF1 V1
du1 F1
19
G4(N2, 2(1, 2)N1, 2,[D22]N1) =
dF2 V2
du 2 F2
1
20
where the notation [Dif] denotes the partial derivative of f with respect to variable i; hence,
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[D11] = 1/1. Letting G/X = A, and assuming that E affects only species 1, the change in
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the sum of species abundances with respect to E is
23
 X 1 *  X 2 *
cof ( A,1,1)  cof ( A,2,1) G

E
det( A)
E
(S3)
24
where cof(A,i,j) = (–1)i+j det([A]i,j) and [A]i,j is matrix A with the ith row and jth column
25
removed.
26
The elements of A can be derived for G1 and G3 as
27
a11 = [D1G1] = –1
28
a12 = [D1G1]1
29
a13 = [D2G1][D11]N2 + [D3G1] = 0
30
a14d1 = [D2G1]N2[D21]
31
a31 = [D1G3]
32
a32 = [D2G3]1 + [D4G3][D11]
33
a33 = [D2G3]N2[D11] + [D3G3] + [D4G3][D111]N2
34
a34d1 = [D2G3]N2[D21] + [D4G4][D121]N2,
35
with symmetric equations for G2 and G4. The term d1 represents the cross derivatives [D21] and
36
[D121] that show when the interaction effect from species 2 on species 1, 1(1, 2), changes
37
with the trait value of species 2. In the specific equations we use for competition and mutualism
38
in which 1(1, 2) = 1(1 + d2), d is synonymous with d1.
39
Expanding equation (S3) in terms of aij and excluding terms of order di2 yields
40
cof(A,1,1)+ cof(A, 2,1) = (-1- a21 ) a33a44 - a23a32 a44 d2 + a23a31a44 d1
41
det(A) = (1- a12 a21 ) a33a44 + d2 ( a23a32 a44 + a23a31a12 a44 ) + d1 ( a14 a41a33 + a14 a42 a21a33 )
2
42
43
44
(S4)
Note that when d1 = d2 = 0,
¶ ( X1 *+X2 *)
(-1- a21 ) a33a44 ¶G = (1+ a21 ) ¶G
=¶E
(1- a12 a21 ) a33a44 ¶E (1- a12 a21 ) ¶E
(S5)
45
which is identical to the result in the absence of coevolution. This shows that coevolution affects
46
the response of equilibrium abundances to changes in E only when selection on the trait values of
47
one species changes its interactive effects i(1, 2) on the other species.
48
Letting C1 = (1 + a21)a33a44, C2 = (1 – a12a21)a33a44,
49
¶X1 *+X2 *
C1 + a32 a23a44 d2 - a31a23a44 d1
¶G1
=
¶E
C2 + ( a23a32 a44 + a23a31a12 a44 ) d2 + ( a14 a41a33 + a14 a42 a21a33 ) d1 ¶E (S6)
50
This expression gives a straightforward way of generalizing the simulation results for the
51
specific forms of competition, mutualism, and predator-prey equations we used. For example,
52
for competition
53
¶X1 *+X2 *
¶G1
(1- a2 ) a1a2 N1N2 + a1a22 N12 d
=
2
2
¶E
(1- a1a 2 )a1a 2 N1 N 2 + a1a 2 (a1 N 2 + a 2 N1 )d ¶E
(S7)
54
Because existence of a positive stable equilibrium requires |i| < 1, it follows that (X1*+
55
X2*)/E increases with d, implying that non-conflicting coevolution will always increase the
56
change in summed equilibrium abundances with changes in E.
57
Broader generalities can be obtained directly from equation (S6) by noting constraints on
58
the signs of the terms aij. The cases of competition and mutualism are similar, as follows. For
59
traits to have optimal values, a33 < 0 and a44 < 0. If higher trait values vi reduce the interaction
60
impact on species i from the other species (as in our models), then a23a32 > 0 and a14a41 > 0. For
61
the exponential forms of equations that we used for simulations, i.e., F1(N1, 1N2, 1) =
3
62
exp(f1(N1, 1N2, 1)), the terms a13 = a24 = 0. Assuming that these values are small, equation S6
63
simplifies to
64
¶X1 *+X2 *
C1 + a32 a23a44 d2
¶G1
=
¶E
C2 + a23a32 a44 d2 + a14 a41a33d1 ¶E
(S8)
65
For competition and mutualism, C1 > C2, which leads immediately to the conclusion that
66
increasing d = d1 = d2 will increase (X1*+ X2*)/E.
67
For predator-prey interactions, equation (S8) also applies, although the signs of the terms
68
differ from the cases of competition and mutualism. The term d1 represents the cross derivatives
69
[D21] and [D121], and similarly d2 represents [D12] and [D122]. Because there is only a single
70
function giving the predation rate, 1(1, 2) = a(1, 2), and 2(1, 2) = –a(1, 2)c, so d1 = –
71
cd2. For traits to have optimal values, a33 < 0 and a44 < 0. In contrast to competition and
72
mutualism, a23a32 ≤ 0 and a14a41 ≥ 0. Thus, positive values of d = d1 = –cd2, which will be the
73
case for predator-prey relationships, will in general decrease (X1*+ X2*)/E. Although this
74
analysis only addresses the case when the environmental change E affects the prey intrinsic rate
75
of increase, a symmetric analysis shows similar results for the case of E affecting predators.
76
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