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Causes of Landscape-Scale Rarity in Cobble Beach Plant Communities

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Causes of Landscape-Scale Rarity in Cobble Beach Plant Communities Ecology, 83(8), 2002, pp. 2304±2314 q 2002 by the Ecological Society of America CAUSES OF LANDSCAPE-SCALE RARITY IN COBBLE BEACH PLANT COMMUNITIES JOHN F. B RUNO1 Department of Ecology and Evolutionary Biology, Brown University, Providence, Rhode Island 02912 USA Abstract. In most communities, a considerable number of the constituent species will only occupy a small percentage of sites or habitat patches. However, the factors that limit the distribution and abundance of rare species are seldom examined and are poorly un- derstood. I tested several explanations of landscape-scale rarity in the New England cobble beach plant community, an assemblage of halophytic forbs associated with fringing beds of the marine intertidal grass Spartina alterni¯ora. Spartina reduces ¯ow velocity and stabilizes the cobble substrate, thereby reducing the burial of seeds and seedlings of other species. Frequencies of the presence of forb species behind 387 sampled Spartina beds distributed along 120 km of shoreline in Narragansett Bay, Rhode Island, USA, ranged from 0.005 to 0.382. Both the whole forb assemblage and individual species were strongly nested with respect to patch size and species richness. Species that were rare on a landscape scale were generally only found behind the longest beds, and bed length was positively related to species diversity. Experimental seed additions to 40 beds ranging in length from 8 to 688 m indicated that intermediate-sized beds often occupied by common species are not habitable by three rare forbs. When present behind the longest beds, these rare species are always associated with a microhabitat characterized by a relatively ®ne-grained and highly stable substrate. A series of substrate manipulations combined with experimental seed additions indicated that the rare annuals Spergularia marina, Salicornia biglovii, and Atriplex arenaria are restricted to the longest Spartina beds because smaller beds lack suitable substrate. These experiments also suggest that seed supply, competitors, and her- bivores are not the proximate causes of the absence of these species from smaller beds, although dispersal limitation may prevent the colonization of many larger habitable patches. The microhabitat required by the rare forbs appears to be generated by the deposition of ®ne-grained particles behind the center of the longest Spartina beds. Consequently, bed length is causally related to the distribution of populations at a landscape scale and to species richness and composition at the scale of a whole habitat patch. Key words: cobble beach; diversity; facilitation; halophytic forbs; nestedness; rarity; species richness. INTRODUCTION 1994, Skaggs and Boecklen 1996). Rare species could A nearly universal attribute of terrestrial and aquatic be poor dispersers that tend to arrive preferentially at communities is a nested distributional pattern in which larger habitat patches due to the large target they pre- the biota at species-poor sites represent a subset of sent (Rabinowitz 1978, Rabinowitz and Rapp 1981, those inhabiting richer sites (Simberloff and Martin Skaggs and Boecklen 1996). Likewise, they can be the 1991, Boecklen 1997). Nestedness is caused by the last to colonize a recently formed, denuded, or highly restriction of rare species to the largest, oldest, and isolated island (Kadmon and Pulliam 1993, Kadmon most species-rich locations (Cutler 1991, Rosenzweig 1995, Hubbell et al. 1999). Rare species can also rep- 1995). Rare species are generally de®ned as those that resent specialists that are unable to colonize small hab- have low abundances and/or small ranges (Gaston itat patches. Required resources, such as food or refuge 1994). Rarity can occur at several spatial scales, as rare from predators, may simply be absent on species-poor species can be absent from most locations within a site islands (Ward and Lakhani 1977, Gaston 1994), a con- or habitat patch or from most sites across a landscape. dition called ``habitat nesting'' (Simberloff and Levin Rare species might also be restricted to a small geo- 1985, Quinn and Harrison 1988, Calme and Desrochers graphic region or even a single landscape or site. 1999). Alternatively, resources might be in short sup- There are numerous potential causes of rarity (Ra- ply, and less common species could be restricted from binowitz 1978, Simberloff and Martin 1991, Gaston small islands by superior competitors. Rare species might be low-disturbance specialists that, unlike hab- Manuscript received 29 January 2001; revised 21 October itat generalists, cannot occupy frequently or severely 2001; accepted 20 November 2001. disturbed sites (Simberloff and Levin 1985). Finally, 1 Present address: Department of Marine Sciences, Uni- versity of North Carolina, Chapel Hill, North Carolina 27599- rare species might require more area or resources to 3300 USA. E-mail: [email protected] maintain viable populations than common species (Pat- 2304 August 2002 CAUSES OF LANDSCAPE-SCALE RARITY 2305 terson and Atmar 1986, Wright et al. 1998). In this recruitment (Bruno and Kennedy 2000). Thus, the scenario, rare species tend to go extinct more frequently smallest beds, which are much more common than large than common species on smaller or species-poor habitat beds, do not represent habitable patches for any plant islands (Quinn and Harrison 1988, Cutler 1991). species. The dispersal-limitation explanation of rarity can be tested by measuring propagule arrival and by experi- METHODS mentally adding propagules, which can determine Pattern description whether a vacant site is indeed habitable by a given species. If the species is unable to colonize the site or Landscape-scale patterns of plant-species abundance habitat patch, a series of habitat manipulations (e.g., and richness were quanti®ed by sampling 387 Spartina predator exclusions or competitor removals) can be beds in Narragansett Bay, Rhode Island, USA, in Au- performed to determine the proximate cause of exclu- gust 2000. I sampled all Spartina beds along 120 km sion. Unfortunately, such ®eld manipulations are sel- of shoreline within a 20 3 30 km study area (see map dom performed at appropriate spatial scales (Gaston in Bruno 2000b:175). I searched the substrate behind 1994). As a result, ecologists generally do not know each bed for the presence of cobble beach plants by why rare species are absent from most locations. This slowly walking along the entire length of each bed two has signi®cantly limited our overall understanding of times. Because cobble beach plants are only found the forces structuring communities and has hampered within a narrow band (1±3 m in width; Kennedy and conservation efforts in many habitat types (Rey and Bruno 2000), and since this is a relatively simple sys- Strong 1983, Simberloff and Martin 1991, Caughley tem with only 12 species of forbs that are all relatively 1994). large and easily distinguished, this technique can be The purpose of this study was to determine the caus- used to accurately quantify species composition. I used es of landscape-scale rarity in the New England cobble the nested temperature calculator to determine the de- beach plant community. The rare species in this system gree of nestedness displayed by the forb assemblage can form dense populations that can be found across a as a whole. This program calculates the system ``tem- large geographic range, but are only present at a small perature'' (T), a measure of the degree of deviation percentage of sites within a given landscape. Prag- from perfect nestedness (Kelt 1997), by comparing the matically, it is often necessary to consider rarity a cat- table of species presence/absence data (with species egorical variable and to de®ne a cutoff point, distin- ranked by abundance and sites ranked by size or rich- guishing rare from common species. As recommended ness; see Appendix), to a randomly generated table in by Gaston (1994), I de®ned rare species as those that which both the number of species at a given site and occupied #25% of the number of habitat patches oc- the number of sites occupied by each species is con- cupied by the most abundant species (i.e., the quartile served. T can range from 0±1008: a system with a T of de®nition). The rare species also occupied #10% of 08 is ``maximally cold'' and is perfectly nested, while all patches. Initial observations suggested that the dis- a system that is structured randomly has a T of 1008 tribution of plant species in this system was highly (Atmar and Patterson 1993). The probability that the nested as rare species were only found in the largest, observed distributional pattern was randomly produced most species-rich patches. Speci®cally, I asked, why was determined by comparing the observed measure are rare species restricted from medium sized patches of T to a distribution of 1000 expected T values cal- that are frequently occupied by common species? culated with a Monte Carlo simulation routine (Atmar The cobble beach plant community is dominated by and Patterson 1993). I also tested the nestedness of halophytic forbs (salt-loving, nonwoody, dicot plants) each individual species in respect to both forb richness and is found below the mean high water mark on es- and Spartina bed length using Wilcoxon two-sample tuarine cobble beaches behind the upper border of rank-sum tests. This simple method can be used to fringing Spartina alterni¯ora beds (Bruno 2000a). assess to what degree (if at all) the distribution of each Spartina acts as a foundation species (sensu Dayton species conforms to a nested pattern by comparing the 1972, Bruno and Bertness 2001) and facilitates this observed pattern of site occupancy (with sites ranked community by reducing ¯ow velocity and stabilizing by size or richness) to a randomly generated pattern in the cobble substrate. This habitat modi®cation reduces which the species occupies the same number of sites the burial of seeds and seedlings of other species, which (Schoener and Schoener 1983, Simberloff and Levin are unable to emerge and survive on portions of the 1985, Simberloff and Martin 1991).
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