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A Mechanistic Simulation Model of Seed Dispersal by Animals

A Mechanistic Simulation Model of Seed Dispersal by Animals

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Published by: Edgar Larrarte Rivera on Jun 19, 2012
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 Journal of Ecology
 2008,
96
 , 10111022doi: 10.1111/j.1365-2745.2007.01341.x
 © 2008 The Authors. Journal compilation © 2008 British Ecological Society
 BlackwellPublishing Ltd
 A mechanistic simulation model of seed dispersal byanimals
 Heidrun Will* and Oliver Tackenberg
 Institute of Ecology, Evolution and Diversity, University of Frankfurt, D-60323 Frankfurt (Main), Germany 
 Summary
 1.
 In order to investigate seed dispersal by animals on a landscape scale, we developed the spatiallyexplicit, individual-based mechanistic model SEED (Simulation of Epi- and Endozoochorous SeedDispersal). The purpose of the model is to predict patterns and densities of seeds dispersed byanimals (especially mammals) within a simulated landscape.
 2.
 The model was parameterized for sheep, cattle and deer as vectors but may be applied to otheranimals if data for parameterization is available. The model data base currently includes parametervalues for about 100 plant species.
 3.
 Seed attachment to and seed detachment from the fur, as well as seed excretion after passagethrough the gut, are explicitly simulated by drawing randomly from distributions that were deter-mined by standardized experiments. Animal movement is simulated as a correlated random walk,but to increase reality of the model, radio-tracking data of animals can also be used.
 4.
 A sensitivity analysis of SEED was conducted to identify the relative importance of plant andanimal traits. The analysis highlighted where the main gaps in our knowledge of seed dispersal pro-cesses lie. Even though in our study endozoochorous dispersal had the higher potential for long-distance dispersal compared to epizoochory, there is only scarce knowledge about seed productionand especially about the proportion of seeds eaten by an animal, parameters which were shown tobe of major importance for dispersal.
 5.
 A comparison of variation in plant and animal traits, respectively, showed that dispersal kernelsdepend more on changes in the animal vector than on the comparably little variation a particularplant species can exhibit. For this reason, animal movement is, from all the dispersal-relevantparameters, the one for which more exact data is most urgently needed.
 6.
 Synthesis.
 The newly developed simulation model will help to understand, quantify and predictlong-distance seed dispersal by animals. The possibility to incorporate real landscapes andmovement data from very different animals makes the model generalizable and possibly applicableto a wide range of scientific and applied questions.
 Key-words:
 animal dispersal, animal movement, long-distance dispersal, mammals, mechanisticmodel, plant traits, plant–animal interactions, seed dispersal, seed shadow, zoochory
 Introduction
 Seed dispersal is a key process in ecology, determining,among other things, colonization, local and meta-populationdynamics, and the spatial structure of plant communities(Nathan & Muller-Landau 2000). Long-distance dispersalevents, although typically rare, are especially crucial topopulation spread, to the maintenance of genetic connectivity,and hence to the regional survival of plant species (Cain
et al 
 .2000).Against the background of changing land use, alien speciesintroduction and, in particular, climate change, it is extremelyimportant to better understand dispersal processes at differentspatial scales. Accurate measures of seed dispersal are essentialto assess its importance for different plant species and theirresponse to environmental changes. Yet, long-distance dis-persal is extremely difficult to quantify empirically. Dispersalmodels have long been used to quantify dispersal processes(Levin
et al 
 . 1984). Classical diffusion models, however,generally underestimated long-distance dispersal events, aswas shown by historical records and molecular analyses (Cain
 et al 
 . 1998; Godoy & Jordano 2001). Mixed dispersal models
 *Correspondence author. E-mail: will@bio.uni-frankfurt.de
 
 1012
 H. Will & O. Tackenberg 
 © 2008 The Authors. Journal compilation © 2008 British Ecological Society,
Journal of Ecology
 ,
96
 , 1011–1022
 with fat tails are also of limited use. They are indeed able topredict long-distance dispersal as inferred from historicalrecords (Clark 1998), but disregard the fact that seed dispersalby animals can basically not be modelled as a decreasingfunction of distance. Instead, it requires the inclusion of habitat preferences, behaviour and movement patterns of theanimals (Vellend
et al 
 . 2003; Russo
et al 
 . 2006).Spatially explicit mechanistic models are a promising toolfor the study of long-distance dispersal and have achievedconsiderable progress in quantifying wind dispersal processes(Nathan
et al 
 . 2001; Tackenberg 2003; Soons
et al 
 . 2004;Katul
et al 
 . 2005; Kuparinen 2006). Zoochorous dispersal,however, is influenced by a large variety of – sometimes inter-related – plant and animal traits, which represent a seriousobstacle in developing dispersal models for zoochory. Empiricaldata for many of the relevant factors are scarce and generalrules often not known. Consequently, few attempts have beenmade to model seed dispersal by animals (Pakeman 2001;Westcott
et al 
 . 2005; Morales & Carlo 2006; Russo
et al 
 . 2006;Couvreur
et al 
 . 2007). With the exception of Pakeman (2001),all of these studies simulated either only endo- or onlyepizoochory by one specific animal vector within one specificlandscape. Pakeman (2001) took data for several animals intoaccount, but made very simplified assumptions about animalmovement.To our knowledge, no study has hereto combined anexplicit simulation of animal movement with empiricallybased routines for epi- and endozoochorous seed dispersal.Recent studies of attachment and detachment of plant seedsto and from animal fur (Couvreur
et al 
 . 2005; Tackenberg
 et al 
 . 2006; Will
et al 
 . 2007) filled an important knowledgegap and made it possible to develop a model for epi- as well asendozoochory which operates at the landscape scale and isbased on parameters empirically determined for differentplant and animal species.Here, we present the spatially explicit, individual-basedmechanistic simulation model SEED (Simulation of Epi- andEndozoochorous Seed Dispersal). The focus of our model lieson dispersal via large mammals. Due to their comparativelylarge home ranges and relatively long food retention times inthe gut (Warner 1981), they are potentially very effectivelong-distance dispersal agents. We show that the mechanisticmodel presented is a valuable tool not only to characterizeseed rain patterns, but also to better understand the processesassociated with seed dispersal and their relative importance.Our model is the first to simulate both epi- and endozoo-chorous dispersal within the same modelling framework andbased on empirical trials. Using wild as well as domesticanimals as vectors, it is possible to run simulations for variousreal and artificial landscapes within time scales relevant for long-distance dispersal. Suitable for studying many aspects of seeddispersal, the model will be particularly useful for ecologistsand conservationists looking at plant species responses tohabitat fragmentation, climate change, invasive species andrestoration issues. Evolutionary ecologists will also gain fromthe model as trade-offs between certain plant traits (e.g. thosethat facilitate either epi- or endozoochory) can be studied.
 Modelling approach
 The model predicts patterns and densities of animal-dispersedseeds within a simulated landscape. It is designed to show howcertain plant and animal traits relate to the shape and scale of animal-generated dispersal kernels. It was parameterized forsheep, cattle and deer as vectors, but may be applied to otheranimals so long as data for parameterization are available.The model data base currently includes parameter values forabout 100 plant species (see Table 1 and Table S1 in Supple-mentary Material).Here, we first outline the basic model concepts, discuss theposition of the model within the context of current knowledgeon seed dispersal processes and highlight how it advances ourunderstanding of these processes. We next introduce the indi-vidual components of the model and show how the resultingdispersal kernels and spatial patterns change when morefactors enter incrementally into the model (Fig. 1). We willthen proceed with a short overview of the most importantmodel processes. A detailed model description is included inAppendix S1 in the Supplementary Material.
 (1)
 
DETACHMENT
 
 AND
 
DEFECATION
 Plant seed retention in animal fur is probably one of thedispersal processes that has been studied longest (e.g. Agnew& Flux 1970 and Bullock & Primack 1977; more recent studiesinclude Fischer
et al 
 . 1996; Couvreur
et al 
 . 2005; Mouissie
 et al 
 . 2005). Several studies have shown that seed retention inthe fur or seed detachment from the fur, respectively, dependson fur type, seed mass and seed morphology (Couvreur
 et al 
 . 2004; Mouissie
et al 
 . 2005; Römermann
et al 
 . 2005b;Tackenberg
et al 
 . 2006). Until now, however, our knowledgeof seed retention capabilities was restricted to relatively fewplant species. Besides, data from different studies were oftennot comparable due to differing study designs or, in the caseof field experiments, due to experimental conditions that werehard to control or to repeat. To parameterize our model, weexperimentally determined detachment curves for more than100 plant species according to the protocol in Tackenberg
 et al 
 . (2006). To our knowledge, no previous study has compiledsuch curves for as many species following one standardizedprotocol. The parameters of the observed detachment curveswere determined for each measured plant species by maximumlikelihood fit to a gamma distribution. These parameters areshape, scale and the proportion of retained seeds, i.e. thoseseeds that did not detach from the fur during the time spanof the experiment and that are hereafter referred to as ‘stuck’(F. Schurr, pers. comm.).Excretion curves for endozoochory were derived fromfeeding experiments with sheep and cattle (Bonn 2005). Theywere assumed to be identical for different plant species,whereas survival rates (see next section) were determinedseparately for each species. Excretion curves are mathemati-cally similar to the detachment curves from the fur and werealso derived by maximum likelihood fit to a gamma distribution.The main difference between detachment (fur) and excretion
 
 Simulation model of seed dispersal by animals
 1013
 © 2008 The Authors. Journal compilation © 2008 British Ecological Society,
Journal of Ecology
 ,
96
 , 1011–1022
 (gut) curves is that the latter start with a lag phase (with noseeds being excreted), followed by a steady increase in thenumber of excreted seeds. A peak of seed numbers is reachedsome time after ingestion and following his peak the curvesteadily declines to zero (Fig. 1a). Although gut passage rateshave already been investigated for a relatively long time(e.g. Warner 1981; Jones & Simao Neto 1987; Gardener
et al 
 .1993; Ramos
et al 
 . 2006; Varela & Bucher 2006), only a fewstudies have simulated endozoochorous seed dispersal basedon empirically determined gut passage rates. The mostprominent exceptions (Westcott
et al 
 . 2005; Russo
et al 
 . 2006)designed their models so as to reflect the specific environmentand dispersing animal under study (tropical forest, a largebird and a monkey species, respectively), whereas our modelaims at a more general simulation of seed dispersal – includingdifferent dispersal vectors in various kinds of landscapes.
 (2)
 
 ATTACHMENT
 ,
INGESTION
 
 AND
 
SURVIVAL
 Unlike detachment and excretion curves, the proportion of seeds that actually attach to the fur of a passing animal wasnot known until recently. Will
et al 
 . (2007) developed ageneral linear model of seed attachment to sheep wool. Thismodel predicts the proportion of attaching seeds (from allseeds in the infructescence) after one passing event. Accord-ing to this study, attachment depends on seed exposure andseed surface structure and may differ markedly between plantspecies. For the current study, we also developed mathematicalmodels to predict attachment to cattle and deer fur (seeAppendix S1). By combining seed attachment with seeddetachment, we were able to calculate dispersal kernels inrelation to all seeds passed by the animal (Fig. 1b).Unfortunately, exact data on the amount of seeds of a givenplant species eaten by a certain animal were not available.To substitute missing data, the proportion of seeds eaten bythe animal is determined through a constant uptake ratedepending on the distance travelled, i.e. 10% of all seeds theanimal passed while taking one step.The number of endozoochorously dispersed seeds, however,is not only reduced because only a certain proportion of seedsare eaten. During gut passage, there might be an additionalloss of seeds due to digestion (in the study described by Bonn,2005, only 3.6% of all viable seeds survived gut passage, i.e.germinated from the dung). To account for this fact, theexcretion curve, like the detachment curve, also contains athird plant species-specific parameter for all the seeds that arenot deposited due to chewing or digestion-related destruction.Based on our data and the assumptions of the proportionof seeds eaten, it is possible to assess the relative proportionsof epi- and endozoochory (Fig. 1b). Previous studies usually
Table 1.
Overview of model parameters. For each parameter, a typical value is given. Plant species parameters are known for approximately 100species; animal species parameters for sheep, cattle and deer (partial). Table S1 in Supplementary Material lists the values known for each plantspeciesParameterUnitCommon valueRangeReference for method
Seed attachment fur
Seed exposureEnclosedEnclosed to fully exposedWill
et al 
. (2007)Seed surface structure0.801Will
et al 
. (2007)Height of infructescencem0.50.091.5Bonn
et al 
. (2000)Animal height (min., max.)m0.30.80.31.5Grzimek (1988)Seed productionm
 –2
1 000 00027618 780 889Sera & Sery (2004)
Seed detachment fur
Retention after one hour (determiningthe parameters of the retention curve)%500100Römermann
et al 
. (2005b)
Seed ingestion
Proportion of seeds eaten%100100EstimationSeed productionm
 –2
1 000 00027618 780 889Sera & Sery (2004)
Gut passage
Parameters of excretion curve:Shape51.87Bonn (2005);Scale8621Römermann
et al.
(2005a)Prop. of seeds digested%960100
Animal movement
Speedm min
 –1
5413EstimationTurning angle
°
1800360Estimation
Landscape
Plant cover %251100Landscape sizekm
2
10
×
10 km1100Arbitrarily determined orbased on real landscapes (maps)Number of attractive sites150-anySize of attractive sitesm
2
20 0001-anyPosition of attractive sitesrandom
567

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