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Boggs Et Al 2006.Pdf Journal of Animal Blackwell Publishing Ltd Ecology 2006 Delayed population explosion of an introduced butterfly 75, 466–475 CAROL L. BOGGS*†, CHERYL E. HOLDREN*‡, IPEK G. KULAHCI*†§, TIMOTHY C. BONEBRAKE*†, BRIAN D. INOUYE*¶, JOHN P. FAY†**, ANN MCMILLAN†, ERNEST H. WILLIAMS†† and PAUL R. EHRLICH*† *Rocky Mountain Biological Laboratory, Crested Butte, CO, USA; †Center for Conservation Biology, Department of Biological Sciences, Stanford University, Stanford, CA, USA; **Nicholas School of the Environment and Earth Sciences, Duke University, Durham, NC, USA; and ††Department of Biology, Hamilton College, Clinton, NY, USA Summary 1. The causes of lagged population and geographical range expansions after species introductions are poorly understood, and there are relatively few detailed case studies. 2. We document the 29-year history of population dynamics and structure for a population of Euphydryas gillettii Barnes that was introduced to the Colorado Rocky Mountains, USA in 1977. 3. The population size remained low (< 200 individuals) and confined to a single habitat patch (∼2·25 ha) to 1998. These values are similar to those of many other populations within the natural geographical range of the species. 4. However, by 2002 the population increased dramatically to > 3000 individuals and covered ∼70 ha, nearly all to the south of the original site. The direction of population expansion was the same as that of predominant winds. 5. By 2004, the butterfly’s local distribution had retracted mainly to three habitat patches. It thus exhibited a ‘surge/contraction’ form of population growth. Searches within 15 km of the original site yielded no other new populations. 6. In 2005, butterfly numbers crashed, but all three habitat patches remained occupied. The populations within each patch did not decrease in the same proportions, suggesting independent dynamics that are characteristic of metapopulations. 7. We postulate that this behaviour results, in this species, in establishment of satellite populations and, given appropriate habitat structure, may result in lagged or punctuated expansions of introduced populations. Key-words: dispersal, Euphydryas, invasion dynamics, Nymphalidae, population structure. Journal of Animal Ecology (2006) 75, 466–475 doi: 10.1111/j.1365-2656.2006.01067.x poorly understood (e.g. Ehrlich 1989; Sakai et al. Introduction 2001). Stasis/expansion presents the largest problem Introduced species suffer one of several fates: extinc- for identification of invasion status, and is attributed tion; immediate population and geographical expan- generally to lag effects in the population dynamics sion; or stasis followed possibly by expansion (reviewed (reviewed in Crooks & Soulé 1999). These can be due to in Sakai et al. 2001). The factors determining what a need for multiple infusions of propagules (e.g. Wing occurs and what allows successful invasion remain 1943), slow growth or Allee effects at low population sizes (e.g. Lewis & Kareiva 1993; Grevstad 1999; Mem- mott et al. 2005), changes in habitat structure allowing Correspondence: Carol L. Boggs, Department of Biological expansion (e.g. Lonsdale 1993; Essink & Dekker 2002; Sciences, 371 Serra Mall, Stanford University, Stanford, CA Rilov, Benayahu & Gasith 2004), invasion of a critical 94305-5020, USA. E-mail: [email protected] mutualist (e.g. Parker 2001) or hybridization (reviewed ‡Current address: 11 Old Colony Place, Falmouth, MA, in Ellstrand & Schierenbeck 2000) or other genetic USA. © 2006 The Authors. §Current address: Department of Ecology and Evolutionary change (Thomas et al. 2001). However, stasis/expan- Journal compilation Biology, University of Arizona, Tucson, AZ, USA. sion patterns could also occur in species that normally © 2006 British ¶Current address: Biological Sciences, Florida State Univer- exhibit episodic outbreaks, such as ungulates (Caughley Ecological Society sity, Tallahassee, FL, USA. 1970) or forest pest insects (Myers 1998). 467 The traditionally envisioned pattern of geographical more than an order of magnitude and then contracting. Population range expansion is a travelling wave converging on a Placed in the context of our current understanding of explosion of an constant rate of expansion across space, defined by a invasion biology, this relatively detailed case study illu- introduced combination of the dispersal parameters and the minates both invasion dynamics over an intermediate butterfly population growth rate of the organism (e.g. Neubert time-scale of decades and patterns associated with & Parker 2004). Modelled expansion speeds differ range expansion from suitable habitat. between scenarios with constant environments and with periodic or stochastic environments affecting the Materials and methods dispersal and/or population growth parameters (Neu- bert, Kot & Lewis 2000b; Neubert & Parker 2004). In periodically fluctuating environments spread may not be constant, but rather the organism may invade new The study population is located at Gothic, Gunnison areas in one generation and retreat the next generation County, Colorado, USA [38°57·5′ N, 106°59·6′ W, in a ‘surge/contraction’ pattern, depending on model 2912 m above sea level (asl)] (Fig. 1). The introduction parameters (Neubert et al. 2000b). Further, in both site, referred to hereafter as the origin patch, is an east– constant and variable environments, the shape of the north-east sloping area of ∼2·25 ha at the base of an dispersal kernel (distribution of movement distances) avalanche run-out zone on Gothic mountain, with influences whether the population’s expansion rate small streams and active beaver ponds (Holdren & converges to a constant rate. If there is an excess of Ehrlich 1981). Beaver and avalanche activity are dis- long-distance dispersers, the expansion rate may accel- turbances that help maintain the habitat, which reverts erate with time (Kot, Lewis & Van den Driessche 1996; otherwise to tall willows and/or spruce forest with time. Neubert & Caswell 2000a; Neubert et al. 2000b). In the The origin patch is a meadow with clumped spruce extreme, populations experiencing episodic outbreaks [Picea engelmannii Parry ex Engelm. (Pinaceae)], have large fluctuations in both their demographic and willow [Salix spp. (Salicaceae)], bog birch [Betula dispersal parameters and potentially also in the shape glandulosa Michx. (Corylaceae)], abundant larval host of the dispersal kernel. Such populations may exhibit plants [primarily Lonicera involucrata Banks ex Spreng. outward migration from what is, in effect, a temporary (Caprifoliaceae), but also Valeriana occidentalis Heller source population, resulting in temporally and/or geo- (Valerianaceae)] and adult nectar plants [e.g. Senecio graphically staggered pulses of expansion surges across triangularis Hook., Erigeron spp. (Compositae) and a landscape. Depending on the details of variation in Heracleum lanatum Michx. (Umbelliferae)]. Casual demographic and dispersal parameters, these expan- observations in this patch indicate increases in height sion surges may result in the establishment of new of spruce and height and coverage of willows, and viable populations and permanent range expansion. decreases in channelization of some streams over the Alternatively, the surges may be followed by total or course of the 29-year study. partial contraction of populations to the original geo- Two other habitat patches, Barclay (∼0·6 ha) and graphical distribution in a pulsed surge/contraction avalanche (0·3 ha), lie 400 m and 565 m, respectively, pattern. This alternative scenario may also be respon- to the south-east of the centre of the origin patch, sible for the establishment of satellite populations downstream along the East River. These patches were around an original core population. initially unoccupied. Both are on east–south-east slop- These alternative scenarios will determine the rate ing areas at the base of avalanche run-out zones, and and pattern of range expansions. Understanding these contain small streams and active beaver ponds. Barclay patterns is a precursor to insight into the processes is the steepest site of the three and contains abundant associated with successful catastrophic invasions. Such willows and old avalanche debris, along with larval patterns may also elucidate the dynamics of range host plants and adult nectar plants. Avalanche is the expansions unaided by humans, via establishment of most open patch with most vegetation under 0·5 m, new populations. except for scattered spruce and a boundary of willows Here we document the 29-year history of the popu- > 1 m tall. lation size and structure of an introduced population of Euphydryas gillettii is univoltine, flying for 3– the butterfly Euphydryas gillettii (Barnes) (Nymphalidae) 4 weeks in late June–late July. Males locate female in Colorado, USA. This population, located in the mates either by searching or by perching on tall spruce southern Rocky Mountains, lies to the south of the or willow (Kulahci & Boggs in preparation). Females butterfly’s natural range, which extends from Wyoming lay eggs in clusters on the underside of the host plant north to southern Alberta. A large number of prop- leaves near the top of the plant, exposed to morning © 2006 The Authors. agules was introduced intentionally into habitat that sun (Williams 1981; Carrillo & Boggs in preparation). Journal compilation matched the habitat of the donor population as closely Pre-diapause larvae spin webs that incorporate sequen- © 2006 British as possible (Holdren & Ehrlich
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