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Huxel Displacement Natives by Invasives Hybridization

Huxel Displacement Natives by Invasives Hybridization

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07/20/2013

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Rapid displacement of native species by invasive species: eects of hybridization
Gary R. Huxel*
Department of Environmental Science and Policy and Institute of Theoretical Dynamics, University of California, Davis, CA 95616, USA
Received 23 March 1998; received in revised form 24 October 1998; accepted 4 November 1998
Abstract
The introduction of non-native populations can lead to the competitive exclusion (displacement) of native populations. This hasbeen hypothesized to be further exacerbated by the potential of hybridization, which can dilute or genetically assimilate the nativegenotype leaving no ``pure'' natives. With relatively moderate to high rates of immigration, the loss of the native species can berapid with or without hybridization. Using single-locus, two-allele models, I ®nd that species replacement can occur very rapidlyand the time to displacement decreases rapidly with increasing immigration and selection dierential. Immigration and selection actin two dierent ways: increasing immigration results in displacement by overwhelming the native; whereas increasing the selectiondierential in favor of the invader leads to displacement via genetic assimilation. The implications of these results are the need formore empirical studies on the immigration patterns of invasive species and their potential for interbreeding with natives.
#
1999Elsevier Science Ltd. All rights reserved.
Keywords:
Introgression; Species displacement; Hybridization; Genetic assimilation; Invading species
1. Introduction
Hybridization is a common phenomenon in plants,birds, ®sh, and many other taxa (Mayr, 1942; Grant,1981; Harrison, 1990; Grant and Grant, 1992, 1996;Dowling and DeMarais, 1993; Levin et al., 1996; Rhy-mer and Simberlo, 1996; Williamson, 1996; Arnold,1997). Yet in studying the displacement of native speciesby invading species the potential for interbreedingbetween an invading species and a native species is oftenignored or understated. It has been suggested thatinterbreeding increases the threat of extinction for anumber of species due to hybridization introgression(Levin et al., 1996; Rhymer and Simberlo, 1996). Forexample, ``pure'' native Pecos pup®sh may no longerexist due to introgression with an introduced bait ®sh,the sheepshead minnow (Echelle and Connor, 1989).Conditions under which hybridization is expected toincrease or decrease the rate of species displacement hasnot been explicitly examined. In cases where relatedtaxa are able to interbreed, introductions may lead tothe introduced taxa dying out, coexistence, new hybridtaxa or extinction of the native taxa. Although theore-tical studies have addressed the introduction of newalleles into populations (i.e. the third phase of Wright'sshifting balance theory), few have focused on the dis-placement of native taxa (alleles). Here, I use singlelocus, two-allele (one native and one non-native) toexamine the in¯uence of interbreeding with invadingtaxa on native taxa.Historically, geographical isolation has been a majorfactor in limiting the impacts of hybridization andintrogression. Today, however, taxa are being broughtinto contact with related taxa from which they havebeen isolated through many anthropogenic pathways(Carlton, 1979, 1989; Carlton and Geller, 1993; Wil-liamson, 1996). For example, the movement of ballastwater throughout the world is increasing the opportu-nities for closely related taxa to interbreed leading tohomogenization of near-shore marine communitiesworldwide and the widespread loss of species (Carlton,1979, 1989, 1996; Carlton and Geller, 1993). Similarlarge distance (inter-regional), short time scale immi-gration occurs with the introduction of nurse stocks of plants and the incidental movement of herbivores andparasitoids (Heywood, 1989). Furthermore, introduc-tion of bird species has been extremely widespread
0006-3207/99/$ - see front matter
#
1999 Elsevier Science Ltd. All rights reserved.PII: S0006-3207(98)00153-0Biological Conservation 89 (1999) 143±152* Tel.: +1-530-752-5162; fax: +1-530-752-3350; e-mail: grhuxel@ucdavis.edu.
 
throughout the world (Moulton and Pimm, 1986;Moulton, 1993; Case, 1996). Human-mediated intro-ductions can be frequent and re-occurring, greatlyincreasing immigration rates (Carlton and Geller, 1993).In the absence of hybridization the interactionbetween the native and introduced species is essentiallya competitive one between species that are similar inmany aspects of their ecology and life histories. Assum-ing a homogeneous environment, without interbreeding,the species with the greater ®tness (e.g. better compe-titor or higher reproductive rate) will dominate at equi-librium. Switching from discussing individuals, I nowfocus on interactions between native and non-nativegenotypes. If immigration rates are very low and givenno (or a weak) selection advantage to the invader, thenone would expect that Allee or stochastic eects maylead quickly to elimination of the invader as its popula-tion density remains low. At very high rates of immi-gration, Kondrashov (1992) has shown thatimmigration eects alone in the absence of interbreed-ing can cause the loss of the native species. Alter-natively, large ®tness dierentials, alone, lead to the®xation of a favored genotype, therefore the relativelyslow process of selection acting without immigrationcan be quickened by increasing the ®tness dierential.For example, intermediate genotypes can be made lethalfor a population with a large, but ®nite, population sizeso that the inferior parental genotype is rapidly lost(Kondrashov, 1992).In the presence of hybridization, one would expectcosts associated with genetic assimilation and/or bene-®ts of infusion of new genes into one or both parentaltaxa. However, hybrids can be fertile yet reproductivelyisolated from adults for various reasons including ferti-lity selection (allopolyploidy, translocation hetero-zygosity, recombination speciation, species-speci®cdierences in mitochrondial DNA) and pre-mating fac-tors (behavioural changes, reproductive phenology,niche separation). Interbreeding may also allow raregenotypes to become established and then increase asbackcrosses with the non-native parental taxa or inter-breeding between hybrids occurs. This of course, issimple in the single locus models used here, but is morecomplicated in real situations involving multiloci (seebelow). Large selection advantages for non-nativealleles should result in more rapid displacement of native alleles with hybridization than without.While studies have shown that lowering the ®tness of hybrids can result in displacement (Kondrashov, 1992),the question remains as to whether the rate of speciesdisplacement increases or decreases with increasedinterbreeding between an invading genotype and anative genotype. Genotypic displacement occurs when anative species becomes extinct and is replaced by aninvading species. By ®tness, I mean the relative compe-titive ability expressed as the number of viable ospringproduced compared with the number produced by othergenotypes. Thus, the interesting question in terms of theconservation of species, and the main question of thispaper, is whether hybridization enhances or impedes the
rate
of species displacement and whether this phenom-enon is aected by introgression. I address this questionusing single locus, two-allele models with varyingdegrees of interbreeding. Additionally, I examined thein¯uence of varying the ®tness of the heterozygote fromunderdominance to overdominance in a system thatexhibits introgression.
2. Model descriptions and analyses
Let us consider two populations with non-over-lapping generations. One population is native and has a®xed allele at a given locus (BB) and the other is non-native with a dierent ®xed allele at the same locus(AA). Non-native individuals can migrate into thenative habitat but the opposite in not allowed. This mayrepresent stocking programs or other human-mediateddispersal so that immigration rates can be much greaterthan otherwise found in natural systems. The popula-tion is assumed to follow Hardy±Weinberg proportionswith selection immediately following immigration. Thisassumes no mutation, large populations and randommating. Furthermore, any competitive dierences areaccounted for in ®tness dierentials. In the model, thepopulation size is in®nite and ®xation of AA is said tooccur when its frequency is greater than 0.99. Fourcases of post mating isolation are modeled: (1) lethalheterozygotes; (2) sterile hybrids; (3) hybrids withbackcross sterility and (4) introgression. A ®fth case wasexamined which also involves introgression, but the ®t-ness of the heterozygote is varied from underdominanceto overdominance. The frequency of AA,
p
AA
, (theintroduced genotype) at time (
t
1):
 p
H
ee
p
ee
1
À
m
 
m
Y
I
where
m
is the migration rate. This assumes thatmigrating individuals each replace one resident indivi-dual proportionally (no preference for displacement of any genotype by immigrants). Fitness of the native spe-cies was assumed to be 1 in all models for mathematicaland interpretive ease.Most theoretical studies have focussed on equilibriumanalyses of introduced alleles, but I was most interestedin the time to displacement. The time frame of dis-placement has important conservation consequencesdue to human-mediated immigration rates and the lackof taxonomic work in many systems (Carleton andGeller, 1993; Geller et al., 1997). Thus I allowed thesystems to run assuming a large population of constantsize from one generation to the next until displacement
144
G.R. Huxel/Biological Conservation 89 (1999) 143±152
 
occurred or 500 generations had elapsed. I used therange of values for ®tness (
3
ee
 Ð®tness of AA in the®rst four cases or
3
ef
 Ð®tness of AB in the last case)from 0 to 4.0 with step increases of 0.1 and m from 0 to1.0 with step increases of 0.1. The initial community iscomprised only of natives (
 p
ff
1).
2.1. Lethal heterozygotes
I simpli®ed the analyses by only following the intro-duced species in this case. If 
p
AA
is the frequency of theintroduced genotype in the generation after selectionand if no interbreeding can occur then
 p
H
ee
3
ee
 p
2
ee
a
3
ee
 p
2
ee
m
 
3
ef

1
À
p
ee
2
À
m

P
where
3
ee
is the ®tness of the ``pure'' invader genotype,
 p
AA
is the frequency of the ``pure'' invader genotype,
m
is the migration rate and
3
ff
is the ®tness of the nativegenotype. This is the baseline case for the discussion,which is essentially a competitive situation where thespecies have remained reproductively isolated.Fixation of the invader genotype (
 p
ee
1) alwaysoccurred when both immigration and
3
ee
were atmoderate levels [Fig. 1(a)]. Fixation also occurred whenthe invader had a disadvantage in ®tness (
3
ee
`
1). Forexample, when
3
ee
b
0
X
20, ®xation occurred wheneverimmigration was greater than 0.30. Under the conditionof no immigration (
m
0
X
0) the invader genotype isalways lost. An interesting dynamic happens at
m
0
X
1which is below the critical immigration rate for which aspecies cannot successfully replace another withoutselection when
3
ee
1 (Crow et al., 1990; Kondra-shov, 1992). As predicted with
m
at this level, at andbelow
3
ee
of 2.0 the
p
AA
slowly approaches 0.07 so that®xation does not occur in 500 generations, but increas-ing
3
ee
to 2.1 leads to ®xation in just 33 generations.This is typical of the dynamics with no hybridizationwhere there exists a sharp boundary between rapid andno ®xation with 500 generations (eventually however®xation will always occur with one way migration and
3
ee
b
1). The frequency of AA after 500 generationswas always less than 0.25 when ®xation did not occurand was usually less than 0.10 [Fig. 1(b)]. When ®xationoccurred it always did so in less than 33 generations andas both
3
ee
and
m
increased, this time to ®xation wasreduced to less than 10 generations.The case of no interbreeding demonstrates that dis-placement is greatly in¯uenced by immigration and dif-ferences in relative ®tness. Time to displacement, whenit occurred, was extremely rapid and decreased as
m
and
3
ee
increased. Immigration rates may be increased byhuman activities; examples of this are found in stockingprograms, in which the introduced species may havegreater population sizes than the native species and inthe absence of introgression may displace the native.This seems to be occurring in the introduction of brooktrout (
Salvelinus fontinalis
) into the habitat of nativebull trout (
S. con¯uentus
) in the Columbia and KlamathRiver drainages (Leary et al., 1993).
2.2. Hybrid sterility
Next I examine the extreme case of hybridizationwithout introgression in which the hybrids are sterile(
3
ef
0), but are part of the
t
1 generation. Theequation for the frequency of AA (
 p
AA
) is the similar toEq. (2), however to maintain a constant population size,
 p
AA
at time
t
1 one must account for the loss of hybrid individuals each generation. Therefore I assumedthat the two parental lineages produce sucient num-bers of ospring so that the population size is constantfrom one generation to the next and do so at theirrespective proportions after selection. For example, if 
 p
ee
0
X
40 and
p
ff
0
X
60 (®tness of all genotypes is set
Fig. 1. The no interbreeding case; (a) time to extinction and (b) fre-quency of AA. These patterns hold for the sterile hybrid and back-cross sterility cases as well.
G.R. Huxel/Biological Conservation 89 (1999) 143±152
145

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