Seawalls Do Not Sustain Viable Populations of Limpets
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MARINE ECOLOGY PROGRESS SERIES Vol. 322: 179–188, 2006 Published September 20 Mar Ecol Prog Ser Seawalls do not sustain viable populations of limpets J. Moreira*, M. G. Chapman, A. J. Underwood Centre for Research on Ecological Impacts of Coastal Cities, Marine Ecology Laboratories A11, University of Sydney, New South Wales 2006, Australia ABSTRACT: Increasing urban development in coastal areas is leading to reduction and fragmenta- tion of natural habitats and a proliferation of artificial structures, such as seawalls. These new struc- tures provide a novel habitat for several organisms, but recent studies have shown differences between the composition of populations in assemblages inhabiting them and those present on natural rocky shores. Preliminary observations also suggested that populations of limpets may differ in size and reproductive output between seawalls and natural rocky shores in Sydney Harbour (Australia). To test hypotheses about the influence of artificial habitat on populations of marine invertebrates, we examined the densities, size and reproductive output of the pulmonate limpet Siphonaria denticulata at midshore intertidal levels on several seawalls and natural rocky shores in Sydney Harbour. Popu- lations on seawalls were mainly composed of juveniles and small adults; large adults were generally found only on rocky shores. There were also significantly fewer and smaller egg masses on seawalls. Experimental manipulations suggested that differences in reproductive output are related to the maximal size that adults attain in each habitat. Populations on seawalls make a smaller contribution to the total reproductive output in any given area, and, thus, it can be predicted that replacement of natural habitats with seawalls may have negative effects on the maintenance of local populations. Populations of organisms living in different habitats in similar densities may show important differ- ences in biological performance. Thus, differences in reproductive output between habitats, even at the same densities, will potentially influence dynamics of populations of limpets. KEY WORDS: Urbanisation · Habitat · Seawall · Rocky shore · Intertidal · Reproduction · Limpets · Siphonaria denticulata Resale or republication not permitted without written consent of the publisher INTRODUCTION The consequences of development of new structures that replace and fragment natural habitat have not, Urban areas currently house nearly half of the however, been widely explored. For some habitats, it is world’s human population (Lim & Sodhi 2004), and it is not known whether fragmentation by building some- expected that this proportion will grow to 60% by the thing is more or less of an influence on sustainability of year 2030 (Pickett et al. 2001). This trend means a con- species than would be the case for other types of frag- tinuous increase in the expansion of cities and towns mentation. On the other hand, where there have been and, therefore, a conversion of natural landscapes to studies of artificial structures, they tend to note human-modified urban landscapes (Collinge 1996). whether abundances of animals are similar to those in Urbanisation removes and fragments natural habitats natural habitats (Dickman & Doncaster 1987, Made- and results in a number of new structures being added jczyk et al. 1998, Laakkonen et al. 2001). When sepa- to the environment (McDonnell & Pickett 1990, Glasby rated from demographic functionality, density may, & Connell 1999). To date, most research on the effects however, lead to wrong conclusions about the value of of urbanisation in terrestrial habitats has been focused artificial structures as alternative habitats, because on habitat fragmentation and performance of species they may just act as ecological sinks in which organ- in remaining patches of natural habitat (Fahrig & Mer- isms fail to breed successfully (Smallwood 2001). Con- riam 1994, Dooley & Bowers 1998, Krauss et al. 2003). sequently, survival of populations in these artificial *Email: [email protected] © Inter-Research 2006 · www.int-res.com 180 Mar Ecol Prog Ser 322: 179–188, 2006 habitats will depend on immigration from other observations suggested that, although there appeared sources (Watkinson & Sutherland 1995). There is, to be large densities on seawalls, the sizes of limpets therefore, an urgent need to assess the impact of artifi- and lengths and numbers of egg-ribbons appeared to cial structures and their value as alternative habitats be smaller than on natural rocky shores. If seawalls are that can sustain viable populations. not providing suitable habitat to sustain breeding pop- Loss of habitat is especially widespread in coastal ulations of these limpets, there are potentially serious marine systems (Moberg & Rönnbäck 2003) and is the consequences of replacing natural habitat with sea- most critical threat to marine biodiversity (Gray 1997). walls. We hypothesised that if seawalls were inade- Declines and extinctions of species can, for example, quate habitats for successful breeding by limpets, be associated with the loss of habitat needed for the there would be less fecundity per capita on seawalls completion of critical parts of the life-cycle (McDowall than in natural habitats. 1992). Growing urbanisation in coastal areas can lead We therefore examined reproductive output of to the reduction and fragmentation of natural habitats Siphonaria denticulata on rocky shores and seawalls in and a subsequent increase in artificial structures, like Sydney Harbour. Sampling provided quantitative data jetties, pilings, pontoons and seawalls (Glasby & Con- on densities and sizes of limpets and number and nell 1999, Wolff 1999, Davis et al. 2002, Chapman & lengths of egg-ribbons to test the hypothesis that Bulleri 2003, Bulleri & Chapman 2004). These new reproductive output would be smaller on seawalls. structures are made of a variety of materials and differ From the results, experiments were designed to test from natural habitats in terms of composition, orienta- the hypotheses that: (1) size of egg-ribbons in each tion, features of the surface and provision of microhab- habitat is related to different rates of survival (i.e. large itats (Chapman & Bulleri 2003). egg-ribbons were not observed on seawalls because Although it has been suggested that seawalls pro- they persist there for less time than on rocky shores), vide a novel habitat and might act as surrogates for (2) size of egg-ribbons depends on size of adults (i.e. rocky shores (Thompson et al. 2002), recent studies long egg-ribbons are not observed on seawalls have shown important differences between intertidal because large adult limpets are sparse or absent) and seawalls and natural rocky shores in terms of composi- (3) intrinsic characteristics of seawalls alter the way tion of biological assemblages and relative abundance eggs are laid (i.e. adults are not able to lay longer egg- of common species (Chapman 2003, Chapman & Bul- ribbons on seawalls). leri 2003, Bulleri & Chapman 2004). In addition, little attention has been paid to the study of the ecological processes of intertidal assemblages on artificial struc- MATERIALS AND METHODS tures (Bulleri et al. 2000, Bulleri 2005), and there have not been any studies on the viability of populations on Density and sizes of Siphonaria denticulata and seawalls. egg-ribbons. Eight different shores were sampled in Approximately 50% of the intertidal habitat in Syd- Sydney Harbour, 4 of them having vertical sandstone ney Harbour (New South Wales, Australia) consists of seawalls and 4 having natural horizontal rocky shores seawalls and similar artificial habitats (Chapman & Bul- (Fig. 1). Hereafter, seawalls and rocky shores are leri 2003). Such extensive creation of artificial habitats referred to as habitats and shores to as locations. Loca- allows tests of some potential consequences of frag- tions were selected because they had either a seawall mentation by unnatural surfaces. In particular, it pro- or a rocky shore extensive enough for the proposed vides a system for investigating whether maintenance sampling (at least 40 m long). They had large popula- of density of species in the artificial habitat is sufficient tions of S. denticulata (>100 ind. m–2). Locations were to sustain populations and ecological processes. separated by at least 100 m. Sampling was restricted to Grazing gastropods are among the most abundant the upper midshore (0.9 to 1.1 m above Indian Low mobile organisms on rocky shores in New South Wales Water Springs), where egg-ribbons were more abun- (Underwood 1978, Creese 1981) and play a major role dant and S. denticulata larger. The sampling design in structuring intertidal assemblages (Underwood & did not eliminate potential sources of confounding, for Jernakoff 1981, Hawkins & Hartnoll 1983, Petraitis example due to differences in exposure and slope, but 1987). Among them, the pulmonate limpet Siphonaria was a sensible scheme on the fragmented shoreline of denticulata is one of the numerically dominant grazers Sydney Harbour (Chapman & Bulleri 2003). All loca- on shores with moderate exposure to wave action tions were, however, located inside Sydney Harbour (Creese & Underwood 1982). This limpet lays benthic and apparently experienced similar wave-wash due to egg-ribbons, and their reproductive season extends intense maritime traffic. It is therefore unlikely that from November to May (Creese 1980). The limpet also differences in exposure exist between the 2 studied lives