September 2002 2577
DIVERSITY–INVASIBILITY IN MARINE SYSTEMS
. 1. Experimental substrate for diversitymanipulations. See
. These species will be hereafterreferred to by genus. Although the true status of theNew England sea squirts as natives or invaders is dif-ﬁcult to resolve due to the poor fossil record of thesesoft-bodied organisms, all of these species have beenpresent in New England for as long as humans havebeen studying these animals (Couthouy 1838,
); DeKay 1843,
, cf. Van Name 1945). Thus,these species form the resident community that currentinvaders encounter upon arrival.Five species of sessile marine invertebrates havebeen recently introduced (1970s–1980s) to New Eng-land and are common on hard substrates, including twocolonial ascidians (
), two solitary ascidians (
), and one encrusting bryozoan (
). Although the
sp. has long been thought to be
, recentmorphological evidence suggests that it may insteadbe
). All of these invaders are similar in feeding modeand growth habit (they are all sessile suspension feed-ers), but they differ considerably in their ecology andlife history (Berman et al. 1992, Osman and Whitlatch1996).
brood very short-lived larvae (a few minutes to hours) that appear ca-pable of settlement almost immediately upon release(Worcester 1994; J. Stachowicz, H. Fried, R. Osman,and R. Whitlatch,
are broadcast spawners whose lar-vae generally become competent to settle within 12–24 h after fertilization (Niermann-Kerkenberg andHoffmann 1989). Thus, there may be differences in thescales over which populations of each species representopen vs. closed systems.
can be semel-parous (Whitlatch et al. 1995), with two cohorts peryear. The second cohort of
recruits in latesummer or early autumn and overwinters as small col-onies, a life history cycle remarkably similar to thenative
(Grosberg 1988). Solitary species suchas
overwinter as large juvenilesand small adults.
is a more recent invaderand exhibits extreme year-to-year and site-to-site ﬂuc-tuations in recruitment and abundance, often being ab-sent from a site for several years and then becomingnumerically dominant the following year.
are colonial and can rapidly propagateasexually, whereas
are solitary,and individual zooids can grow much larger than thoseof the colonial species, but must reproduce sexually.Thus these species, while taxonomically and ecologi-cally similar enough to facilitate direct comparisons,also provide sufﬁcient diversity in life history and eco-logical properties to allow an assessment of the gen-erality of our results.
To assess whether native community species richnessin the ﬁeld was correlated with invasion success, werecorded the presence of native and introduced sessilespecies in 25
25 cm quadrats at 12 different sites(10 quadrats per site) in eastern Long Island Sound,including the Thames, Poquonnock, and Mystic riverestuaries. Sites ranged within 1–10 m depth and in-cluded habitat provided by pilings, rocky outcrops,boulder ﬁelds, seagrass beds, and kelp forests. Sam-pling was initiated from a haphazardly chosen locationwithin each site and individual plots were located atrandomly selected distances and directions with a min-imum of 5 m between samples. An inventory of allsessile organisms present in each plot was made andthe data used to assess the correlation between the num-ber of native and nonnative species in an individualquadrat. Data were later pooled within each site to as-sess the correlation between the number of invasivespecies and the number of native species at the scaleof an entire site (
50 m). Plots of cumulativespecies totals vs. the number of quadrats sampled in-dicated that our sampling effort captured most mac-roscopic species present within a site (J. Stachowiczand R. Osman,
Manipulating species richness
We manipulated species richness and compositionusing a 10
10 cm polyvinyl chloride (PVC) substratecomposed of a ﬁve-by-ﬁve grid of 2
2 cm PVC tiles(Fig. 1), each with a single individual or colony of anative species. One side of each tile was ﬂat, and wetransplanted juveniles or allowed larvae to settle ontothis side. The other side of the tile had a T-shaped