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models predict that species richness should peak at intermediate levels of both factors. such
unimodal responses have been documented in many field and laboratory studies and have usually
been attributed to differences among species in competitive ability and / or trade-offs betwen
competitive ability and tolerance to disturbance. here we show that most documented patterns of
disturbance-richness and productivity richness relationships, as well as the observed interactions
between the two factors, can be explained by a smile neutral model where all species are
ecologically identical and lack trade-offs in species characteristics. this finding suggests that
current neutral theories cam be extended to explain patterns of species responses to productivity
and disturbance.
a variety of ecological models predict that species richness should peak at intermediate levels of
disturbance and productivity. empirical evidence consistent with such unimodal responses has
been reported in a wide range of field studies and laboratory experiments . however, other
una variedad de modelos ecológicos predicen que la riqueza de especies debe pico en los niveles
intermedios de disturbio y productividad. evidencia empírica consistente con estas respuestas
unimodal se ha divulgado en una amplia gama de estudios de campo y experimentos de
laboratorio. Sin embargo, otros
forms of responses, such as positive monotonic, negative monotonic, and nonsignificant patterns,
have also been reported frequently. explaining these variable responses and the mechanisms by
which disturbance and productivity affect patterns of species richness has been a continuing
challenge for ecologists during the last decade.
the unimodal response of species richness to disturbance has been attributed to a trade-off
between competitive ability and tolerance to disturbance . this trade-off is the essence of the
intermediate disturbance hypothesis . the decline of species richness at high levels of productivity
has usually been attributed to an increase in competition intensity, which leads to competitive
exclusion of inferior competitors from the community.
the interactions between the effects of disturbance and productivity on species richness have also
been attributed to trade-offs between competitive ability and life-history traits that facilitate
tolerance to disturbance. thus, most existing theories attribute the observed relationships
between species richness, disturbance, and productivity to ecological trade-offs ans / or
differences among species in competitive ability.
las interacciones entre los efectos de la perturbación y la productividad en la riqueza de especies
también se han atribuido a las compensaciones entre rasgos de historia de vida que facilitan a la
alteración de la tolerancia y la capacidad competitiva. por lo tanto, la mayoría de las teorías
existentes atribuyen las relaciones observadas entre la riqueza de especies, disturbio y la
productividad ecológica compensaciones ans / o las diferencias entre especies en capacidad
competitiva.
in this study, we provide an alternative explanation for the observed relationships between
species richness, disturbance, and productivity. specifically, we show that most documented
effects of disturbance and productivity on species richness, including their interactions, can be
generated by a simple neutral model of community dynamics. the basic premise of neutral models
is that all species are ecologically equivalent and posses the sam prospects of reproduction,
mortality, and migration. since all species in our model are assumed to be identical, our results
contrast with the widely accepted notion that ecological trade-offs and / or differences in
competitive ability are essential for explaining the observed relationships between species
richness, disturbance, and productivity. these findings provide further support for the view that
neutral processes can explain fundamental patterns in community ecology.
en este estudio, le ofrecemos una explicación alternativa para las relaciones observadas entre la
riqueza de especies, disturbio y la productividad. específicamente, mostramos que más efectos
documentados de perturbación y la productividad en la riqueza de especies, incluyendo sus
interacciones, pueden ser generados por un simple modelo neutral de la dinámica comunitaria.
la premisa básica de modelos neutrales es que todas las especies son ecológicamente equivalente
y poseen las perspectivas sam de reproducción, mortalidad y migración. puesto que todas las
especies en nuestro modelo se asumieron que son idénticos, nuestros resultados contrastan con
la idea ampliamente aceptada que compensaciones ecológicas y / o las diferencias en capacidad
competitiva son esenciales para explicar las relaciones observadas entre la riqueza de especies,
disturbio y la productividad. Estos resultados proporcionan la ayuda adicional para la vista que
los procesos neutrales pueden explicar patrones fundamentales en ecología de comunidades
Methods
model description
the model we used is a modification of the neutral model developed by bell. the model describes
the dynamics of a local community embedded within a regional species pool from which the local
community is colonized. the local community has a carrying capacity of J individuals, and the total
number of species in the regional species pool is S. each iteration of the model involves four
successive processes : mortality - each individual in the community dies with probability d ( resulting
in D1 deaths for the i the species) reproduction - each individual that survives gives rise to a single
offspring with probability r (totaling R1) immigration - a fixed number I of individuals are drawn at
random from the regional species pool such that each individual has a probability of 1/S of belonging
to each of the S species ( I1 individuals of the ith species) and recruitment - if the number of potencial
colonizers ( inmigrants plus locally produced offspring) are added to the community.
Métodos
Modelo Descripción
el modelo que utilizamos es una modificación del modelo neutro desarrollado por bell. el modelo
describe la dinámica de una comunidad local incrustada dentro de un grupo de especies
regionales de los cuales la comunidad local es colonizada. la comunidad local tiene una capacidad
de carga de individuos J, y el número total de especies en la piscina de especies regionales es S.
cada iteración del modelo involucra cuatro procesos sucesivos: mortalidad - muere cada individuo
en la comunidad con la reproducción de probabilidad d (resultando en muertes D1 para la i las
especies) - cada individuo que sobrevive da lugar a una descendencia sola con la inmigración de
probabilidad r (totalizando R1) - un número fijo de individuos son extraídos al azar de la especies
regionales piscina tal que cada individuo tiene una probabilidad de 1/S de pertenencia a cada una
de las especies de S (I1 individuos de la especie de ith) y contratación - si el número de
colonizadores potenciales (inmigrantes más localmente producción descendencia) se agregan a la
comunidad.
alternatively, if the number of potencial colonizers exceeds the number of vacant sites , individuals
are draen at random from the potencial colonizers and addes to the community until it reaches a
size of exactly J individuals. in this case, the number of added individuals from species 1, X1 , has a
hypergeometric distribution with three parameters : the number of vacant sites *** , the number
of individuals of all species in the pool of potential colonizers * and the number of individuals of the
ith species in the pool of potencial colonizers * . disturbance and productivity are introduced to the
model as modifiers of mortality and reproduction probabilities . specifically , an increase in
disturbance increases the death rate of all individuals by a constant ** that is , * and an increase in
productivity multiplies the birth rate of all individuals by a constant pi that is , *.
in contrast to previous neutral models that were limited to satured communities , our model applies
to both satured and unsatured.
our model can be considered a demographic formulation of the classical macarthurd and wilson
model of island biogrography. as in the original theory , the focus of the model is on an island ( local
community in our terminology) that receives immigrants fro an external mainland that functions as
a regional species pool. there is no speciation in the system, and the steady state number of species
on the island is determined by the balance between local colonization and extinction rates.
however, in constrast to the original formulation by macarthur and wilson , where the basic
processes are colonization and extincion, in our model, colonization and extincion rates are derived
from individual-level processes of reproduction, mortality, and immigration.
en contraste con modelos anteriores neutrales que se limitaban a las comunidades saturadas,
nuestro modelo se aplica a ambos saturadas e insaturados. nuestro modelo puede ser
considerado una formulación demográfica de la macarthurd clásica y wilson modelo de
biogrography de la isla. al igual que en la teoría original, el enfoque del modelo está en una isla
(comunidad local en nuestra terminología) que recibe a inmigrantes afro un continente externo
que funciona como un grupo de especies regionales. No hay ningún especiación en el sistema, y
el número de estado estacionario de especies en la isla se determina por el equilibrio entre la
colonización local y las tasas de extinción. Sin embargo, en contraste con la formulación original
de macarthur y wilson, donde los procesos básicos son la colonización y extincion, en nuestro
modelo, las tasas de colonización y extincion derivan de nivel individual procesos de
reproducción, mortalidad e inmigración.
it should be emphasized that the term neutral model as used here , means that all individuals of all
species have the same per capita demographic rates. this definition is similar to that used in most
previous studies but it differs from that in models incorporating colonization-extincion trade-offs in
which neutrality was defined as ecological equivalence in the ability to persist at the absence of
interspecific competition.
Cabe destacar que el término neutro modelo como utilizado aquí, significa que todos los
individuos de todas las especies tienen las mismas tasas demográficas per cápita. Esta definición
es similar a la utilizada en la mayoría de los estudios anterior pero difiere en los modelos
incorporan la colonización-extincion compensaciones en el cual neutralidad fue definido como
equivalencia ecológica en la capacidad de persistir en la ausencia de competencia interespecífica.
simulations
the effects of disturbance and productivity on species richness were analyzed for disturbance levels
* ranging from 0.0 to 0.4 (intervals of 0.02) and productivity levels pi ranging from 1 to 7 . these
ranges were chosen to obtain both saturated and unsaturated steady state communities. each
combination of disturbance and productivity was simulated using carrying capacity values of J= ###
individuals and immigration rates i=## nd 32 individuals. values of d, r and S were constant in all
simulations (d=### r=# s=200) . for each of the # sets of parameters , we ran 10 simulations. each
simulation was initiaded by # startuo steps to allow sufficient time for the community ro reach
steady state. all simulations were started with satured communities where each species was
represented by J/S individuals (we also ran simulations starting with monodominance and verified
that they give similar results). species richness of steady state communities was determined as the
mean value of 100 sampling points ( every 20steps) after the # startup steps. extincion rates
(number of species extinct per time step) , colonization rates (number of new species arriving per
time step) , mean number of individuals per species , and the relative frequency of immigrants in
the pool of potential colonizers were determined as the mean values of * sampling points (every
five steps) after the # startuo steps. all attributes were averaged across the 1o replicates of each
parameter st. the results presented here are for J = # and i =8 but the observed responses of species
richness to disturbance and productivity were consistent in all simulations.
Results
model simulations show that disturbance and productivity may have positive, negative, or
unimodal effects on species richness . at intermediate levels of productivity, disturbance has a
unimodal effect on species richness, as predicted by the intermediate disturbance hypothesis .
similarly, at intermediate levels of disturbance, species richness shows a unimodal response to
productivity. thus , in contrast to existing ecological theory, unimodal responses of species richness
to disturbance and productivity can be obtained in the absence of any trade-offs or differences
among species in competitive ability.
the results further show that the effect of disturbance depends on the level of productivity and
vice versa. specifically, the productivity level that maximizes species richness increases with
increasing disturbance, while the level of disturbance that maximizes species richness increases with
increasing productivity. at the extremes, species richness is negatively correlated with productivity
when disturbance is very low but positively correlated with productivity when disturbance is very
high. similarly, species richness only decreases with disturbance when productivity is low and only
increases with disturbance when productivity is high these patterns are similar to those predicted
by kondoh , but in contrast to this model, our simulations do not incorporate any differences among
species in competitive ability, and all patterns are generated by chance events of reproduction,
mortality, and inmigration.
how can the unimodal patterns generated by the neutral model be explained ? detailed examination
of the results indicates that different mechanisms are responsible for the increasing and decreasing
phases ofeach response. at relatively low levels of productivity, the steady state communities are
unsaturated ( total number individuals < J) are increasing productivity is associated with a
corresponding increase in the mean population size leads to a decrease in the probability of
stochastic extinctions and therefore to an increase in species richness. this mechanism was termed
the more individuals hypothesis in previous studies.
¿Cómo pueden explicarse los patrones unimodal generados por el modelo neutral? examen
detallado de los resultados indica que diferentes mecanismos son responsables por el aumento y
la disminución de cada respuesta de fases. a niveles relativamente bajos de productividad, las
comunidades de estado estacionario son insaturadas (total individuos números < J) están
aumentando la productividad está asociada con un aumento correspondiente en la población
media tamaño conduce a una disminución en la probabilidad de extinción estocástico y por lo
tanto a un aumento en la riqueza de especies. Este mecanismo se denominó la hipótesis de los
individuos más en estudios previos
the negative effect of disturbance on species richness at relatively high levels of disturbance can
also be attributed to the MIH mechanism. here an increasein disturbance reduces the mean number
of individuals per species in the community . as a result , the probability of sthochastic extincion
increases and the steady state numer of species decreases. note that the MIH mechanism does not
require any trade-offs or ecological differences among species. it is therefore not surprising that it
influenced the patterns generated by our neutral model. the decrease in species richness with
increasing productivity at relatively high levels of productivity is more interesting because all
previous theories have attributed this pattern to differences among species in competitive ability.
we relate the decreasing phase of the productivity-richness relationships observed in our
simulations to changes in the balance between reproduction and immigration along the productivity
gradient. since reproduction rates are positively correlated with productivity, while immigration
rates are indenpendent of productivity , an increase in productivity reduces the relative frequency
of immigrants in the pool of potencial colonizers
el efecto negativo del disturbio en riqueza de especies en niveles relativamente altos de disturbio
también puede atribuirse al mecanismo de MIH. aquí un disturbio incremento reduce el número
de individuos por especie en la comunidad. como resultado, la probabilidad de extincion
estocásticos aumenta y disminuye el número de estado estacionario de especies. tenga en cuenta
que el mecanismo MIH no exige compensaciones ni las diferencias ecológicas entre las especies.
por lo tanto no es sorprendente que influyó a los patrones generados por nuestro modelo neutral.
la disminución de la riqueza de especies con el aumento de la productividad en niveles
relativamente altos de productividad es más interesante porque todas las teorías anteriores han
atribuido este patrón a las diferencias entre especies en capacidad competitiva. relacionamos la
fase decreciente de las relaciones de productividad-riqueza observada en nuestras simulaciones
a cambios en el equilibrio entre la reproducción y la inmigración a lo largo del gradiente de
productividad. desde tasas de reproducción están correlacionadas positivamente con la
productividad, mientras que las tasas de inmigración son independientes de la productividad, un
aumento de la productividad reduce la frecuencia relativa de los inmigrantes en la piscina de los
colonizadores potenciales
since only new immigrants have the potencial to increase local species richness, increasing
productivity is associated with a decrease in the rate by which new species are added to the
community and therefore reduces the steady state numer of species. this mechanism is limited to
relatively high levels of productivity because only then do locally produced individuals compete with
immigrating individuals for vacant sites.
we term this mechanism the dilution effect because the increase in the number of locally produced
individuals dilutes the concentration of new immigrants in the poolof potencial colonizers and thus
decreases the likelihood that new species will be added to the community after stochastic
extinctions.
puesto que sólo los inmigrantes tienen el potencial para aumentar la riqueza de especies locales,
aumento de la productividad se asocia con una disminución en la tasa que se agregan a la
comunidad y por lo tanto reduce el número de estado estacionario de especies nuevas especies.
Este mecanismo se limita a niveles relativamente altos de productividad porque sólo entonces
individuos producidos localmente que compiten con las personas inmigrantes para sitios
vacantes. plazo este mecanismo el efecto de dilución porque el aumento en el número de
individuos producidos localmente diluye la concentración de inmigrantes en los piscona de
colonizadores potencial y por lo tanto disminuye la probabilidad que se añadirán nuevas especies
a la comunidad después de las extinciones estocásticas.
the dilution effect also explains the increase in species richness along the increasing part of the
disturbance gradient. however in this case it operates in an opposite direction : as disturbance
increases, more individuals die at each time step , and the total number of offspring produced by
the remaining individuals decreases. since immigration rates are fixed, the relative frequency of
inmigrants in the pool of potential colonizers increases, which leads to a corresponding increase in
the rate by which new species are added to the community. the result is an increase in the steady
state number of species.