物种分化速率

Three

Assumptions

in

Neutral

Community

EcologyNeutrality

assumptionZero-Sum

assumptionPoint

mutuation

assumptionHubbell,

S.

P.

2001.

The

unified

neutral

theory

of

biodiversityand

biogeography.

-

Princeton

University

Press.第二页，共十九页，2022年，8月28日Relax

point

mutuation

assumption---

random

fissionspeciation•Hubbell

(2001)

stated

that

the

equilibriummetacommunity

SAD

resulting

from

random

fissionspeciation

is

a

zero-sum

multinomial.•Ricklefs

(2003)

provided

some

approximate

formulas

for

thetotal

species

richness

in

the

metacommunity

under

randomfission

speciation.•Etienne

and

Haegeman

(2010)

derive

analyticalexpressions

for

the

distribution

of

abundancesaccording

to

the

neutral

model

with

recruitment

(i.e.,dispersal

and

establishment)

limitation

and

randomfission

speciation.第四页，共十九页，2022年，8月28日Relax

point

mutuation

assumption---

random

fission

speciationEtienne,

R.S.

and

Bart,

Haegeman,

2010.

The

neutral

theory

biodiversity

with

random

fission

speciation.Theor

EcolFigure

5

shows

the

abundance

distributions

with

the

fitted

models.

Clearly

the

model

with

random

fission

speciation

never

performs

significantlybetter

than

the

point

mutationmodel.第五页，共十九页，2022年，8月28日Relax

point

mutuation

assumptionThe

metacommunity

abundance

distribution

under

random

fission

is

identical

to

thebroken-stick

abundance

distribution,

and

thus

provides

a

dynamical

explanation

for

thgrand

old

lady

of

abundance

distributions.---

random

fission

speciationThe

metacommunity

abundance

distribution

under

random

fission

:第六页，共十九页，2022年，8月28日Relax

point

mutuation

assumption---

speciation

rate

proportional

to

abundanceEtienne

RS,

Apol

MEF,

Olff

H,

Weissing

FJ

(2007b)

Modes

of

speciation

and

the

neutral

theory

of

biodiversity.

Oikos116:241–258.第七页，共十九页，2022年，8月28日Relax

point

mutuation

assumptiontion

rate:---

Protracted

speciation••

PPrroottrraacctteeddspsepceicaitaitoino:n:bybmyomdoedleilnignsgpsepceicaitaitoinoans

asgargardaudaula,lp,rportortarcatcetde,dp,rporcoecses

rsartahtehretrtanah

instana

intsatnaenotuasneventsuo

e.vent.••

SSpeciationaicep

rate:,

whe,rewheu

irsegueisren

gaelnsepreaclisapteicoina-tiinoint-iiantiitoinartaitoen.rate.第八页，共十九页，2022年，8月28日Relax

point

mutuation

assumption---

Protracted

speciationRosindell

et

al.,2010.

Protracted

speciation

revitalizes

the

neutral

theory

of

biodiversity.

Ecol.

Lett.

13:716-727.第九页，共十九页，2022年，8月28日Relax

point

mutuation

assumption---

ProtractedspeciationRosindell

et

al.,2010.

Protracted

speciation

revitalizes

the

neutral

theory

of

biodiversity.

Ecol.

Lett.

13:716-727.第十页，共十九页，2022年，8月28日Three

Assumptions

in

Neutral

Community

EcologyWhile

the

neutrality

assumption

is

the

heart

of

the

neutraltheory,

the

other

two

assumptions

are

assumptions

ofparticular

model

implementations

ofthe

theory,allowing

foranalytical

expressions

for

diversity

measures,

rather

than

thefundamental

assumption

of

the

theory

itself.

A

mismatchbetween

observes

and

theoretical

predictions

can,

be

due

tothese

additional

assumptions

and

therefore

cannot

beimmediately

interpreted

as

calling

for

a

rejection

of

the

neutraltheory

as

awhole.Rampal

EtienneEtienne,

R.S.

and

Bart,

Haegeman,

2010.

The

neutral

theory

biodiversity

with

random

fission

speciation.Theor

Ecol第十一页，共十九页，2022年，8月28日Further

study

–

niche-ovlappedHowever,

the

degree

to

which

real

ecological

communites

are

acttually

neutral

is

a

more

openquestion

(Gotelli

and

McGill,

2006).

That

is,

what

is

a

distribution

of

ecological

communitiesalong

a

continuum

form

purely

neutral,

througth

weekly

nich-structured,

to

strongly

niche-structured

(Adler,

HilleRisLambers

&

Levine

2007)?---

Different

but

equal:

the

implausible

assumption

at

the

heart

of neutral

theory,

2011In

the

present

case,

our

question

is:

when

is

the

limited

complexity

of

the

neutral

model

sufficient

to

provide

auseful

approximation

to

a

model

of

biodiversity

with

simple

niche

structure?---Theory

predicts

a

rapid

transition

from

niche-structured

to

neutralbiodiversity

pattera

speciation-rate

gradient,

2011.第十二页，共十九页，2022年，8月28日Further

study

–

niche-ovlappedChishom

and

Palaca

(2010)

present

an

analytical

proof

that,

in

the

limit

as

diversitybecomes

large,

a

strong

niche

model

give

rises

to

exactly

the

same

asymptotic

form

of

SAD

as

the

neutral

model.Haegeman

and

Etienne

(2011)

argue

that

their

result

is

an

artefact

because

theimodel

for

high

diversity

is

equivalent

to

an

independent-species

model.

theyanalyse

this

independent-species

model,

and

show

that

its

SADs

are

identical

toneutral

predictions

for

all

levels

of

diversity.第十三页，共十九页，2022年，8月28日Further

study

–

niche-ovlappedChishom

and

Palaca

(2011)

:

As

the

speciation

rate

in

the

hybrid

model

increases,

we

observe

a

surprisingly

rapid

transitionfrom

an

ecosystem

in

which

diversity

is

almost

entirely

governed

by

niche

structure

to

one

in

which

diversity

is

statisticallyindistinguishable

from

that

of

the

neutral

model.Posterior

probability

that

an

observed

species

abundance

distribution

from

the

hybrid

model

comes

from

a

neutral

model,

given

an

uninformativedichotomous

prior

that

specifies

equal

probabilities

for

the

neutral

model

and

the

pure

(broken-stick)

niche

model

(P

(neutral)

=

P(brokenstick)=0.5),

for

different

values

of

the

speciation

rate.

Other

parameters

are

m=0.1,

J=20,000,

JM=1012,

K=16.

See

text

for

model

details.

Dashedlines

show

95%

confidence

intervals

(estimated

from

20

simulations)第十四页，共十九页，2022年，8月28日Further

study

–

Relax

neutrality

assumptionZhou

and

Zhang

(2008):

The

neutral

theory

can

be

extended

to

explain

theseobservations

by

allowing

species

to

differ

slightly

in

their

competitive

ability(fitness).第十五页，共十九页，2022年，8月28日Further

study

–

Relax

neutrality

assumptionZhang

et

al.(2011)

demonstrate

how

such

a

weaker

form

of

symmetry

can

generatesignificant

macroecological

diversity

patterns,

leading

to

a

very

differentinterpretation

of

the

relation

between

Fisher’s

a

and

Hubbell’s

fundamentalbiodiversity

number.,

where第十六页，共十九页，2022年，8月28日Further

st