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Prediction of Rare-Plant Occurrence: a Southern Appalachian Example

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Prediction of Rare-Plant Occurrence: a Southern Appalachian Example Friday Oct1610:17 AM ecap 8 407 Mp 909 Allen Press x DTPro File # 07sc November 1998 ECOLOGICAL CONCEPTS IN CONSERVATION BIOLOGY 909 Ecological Applications, 8(4), 1998, pp. 909±920 q 1998 by the Ecological Society of America PREDICTION OF RARE-PLANT OCCURRENCE: A SOUTHERN APPALACHIAN EXAMPLE SUSAN K. WISER,1 ROBERT K. PEET, AND PETER S. WHITE Department of Biology, CB#3280, University of North Carolina, Chapel Hill, North Carolina 27599-3280, USA Abstract. Ecologically sound efforts to manage or reintroduce populations of rare species require detailed knowledge of species habitat requirements. However, the fact that such species are rare implies that the data needed for habitat characterization are sparse and that species might well be absent from favorable sites due to chance aspects of dispersal or mortality. We use four rare plant species endemic to southern Appalachian high-elevation rock outcrops, to illustrate how nonparametric and parametric logistic regression can yield predictive models of the probability that a species will occur, given certain site conditions. Models were constructed for each species at two scales: 100-m2 plots and 1-m2 subplots. At the 100-m2 plot scale, absences beyond the current geographic range were excluded. At the 1-m2 subplot scale, absences from subplots were only included if the species occurred elsewhere on the 100-m2 plot. Six signi®cant models resulted; no signi®cant model could be constructed for Solidago spithamaea or Calamagrostis cainii on 1-m2 subplots. For 100-m2 plots, the most valuable predictors were potential solar radiation, a soils gradient related to available soil iron, boron, and copper, and coarse-scale rock surface texture, although Geum radiatum occur- rences were dif®cult to predict at this scale. For 1-m2 subplots the best predictors were available soil cations, potential solar radiation, the proportion of exposed bedrock, and vegetation height. Along individual gradients response curves were often similar, but no two species were predicted by identical sets of site parameters. Beyond current range limits, existence of suitable habitat on 100-m2 plots was demonstrated for Solidago spithamaea, supporting a view that the range limits of this species are not necessarily set by availability of suitable habitat. Habitat-based models have numerous management applications (such as to guide restoration and reintroduction efforts as well as to direct searches for additional populations) and provide a framework for future work on species-speci®c physiological requirements. Key words: Calamagrostis cainii; endemic; generalized additive models; Geum radiatum; habitat characterization; Houstonia purpurea var. montana; logistic regression; rare plants; rock outcrops; Solidago spithamaea; southern Appalachian Mountains (USA). INTRODUCTION plied to rare species at a site-by-site scale can be used Effective management of rare plant species requires by managers to (1) guide searches for unknown pop- an understanding of their ecology. Knowledge of which ulations, (2) indicate site suitability for restoration or habitat parameters most accurately predict the occur- reintroduction, (3) predict impacts of habitat degra- rence of a rare plant species, and the likelihood that dation and (4) provide a framework for further research species will occur given speci®c site conditions, is fun- on speci®c physiological requirements. damental (Simberloff 1988, Brussard 1991, Falk and In the southern Appalachian Mountains, high-ele- Olwell 1992). The development of analytical approach- vation rock outcrops constitute a rare habitat, and many es to predict the occurrence of plant species from en- of the inhabitants grow in few, if any, other habitats vironment has received considerable attention (e.g., in the region. Forty outcrop species are regionally rare, Austin et al. 1984, Nicholls 1989, Yee and Mitchell including twelve that are either southern Appalachian 1991), but has generally been applied to common spe- endemics or species disjunct from alpine, arctic, and cies at broad geographic scales. Such approaches ap- other treeless habitats far to the north. These species are noteworthy in that they are considered relicts of a Manuscript received 10 July 1995; revised 1 April 1996; Pleistocene alpine ¯ora in a region where no alpine accepted 20 April 1996; ®nal version received 20 May 1996. environment currently exists (White et al. 1984, Schaf- For reprints of this Invited Feature, see footnote 1, p. 907. 1 Present address: Landcare Research, P.O. Box 69, Lin- ale and Weakley 1990). Beyond the observation that coln, New Zealand. E-mail: [email protected] they are restricted to outcrops above 1200 m, little 909 Friday Oct1610:17 AM ecap 8 407 Mp 910 Allen Press x DTPro File # 07sc 910 INVITED FEATURE Ecological Applications Vol. 8, No. 4 about their distribution relative to environment has 1987, Murdock 1993, Saunders 1996). All four species been published. Implicit in earlier publications is the are perennials. The limited data on seed dispersal sug- assumption that these species persist because condi- gest that most dispersal is passive by wind, gravity, or tions peculiar to certain high-elevation outcrops, such overland ¯ow. Birds and rodents are frequent on the as moist north-facing slopes, closely resemble past al- outcrops, but their role in seed dispersal is unknown pine conditions. Within these macrosites, the species and probably minimal. Propagation by rhizomes has may be expected to follow the general pattern observed been observed in Geum and Houstonia. Only Cala- for other outcrop endemics of the Southeastern United magrostis has been observed to colonize disturbed States and occur on shallow-soiled microsites where sites, and this was adjacent to a pre-existing population competition and shading from other species is low (S. K. Wiser, personal observation). Herbarium records (Baskin and Baskin 1988). suggest that these species have been present at most of In this paper, nonparametric and parametric logistic their current sites for at least a century. Field obser- regression are used to determine the utility of a range vations suggest recent population declines in Solidago, of physical site parameters in predicting four of the Geum, and Houstonia, whereas insuf®cient data are twelve rare, Pleistocene-relict outcrop species at two available to assess population trends in Calamagrostis spatial scales. The following questions are addressed (N. Murdock [U.S. Fish and Wildlife Service, Ashe- for these four species. (1) At the scale of the 100-m2 ville, North Carolina, USA], personal communication). plot, which environmental factors best predict species occurrence? Are these relicts predicted to occur at the Study area highest elevations and coolest (i.e., lowest potential All populations of the four study species occur ex- solar radiation) sites? (2) At the scale of the 1-m2 sub- clusively on outcrop sites above 1200 m and south of plots, which environmental factors best predict species the Virginia±North Carolina border (Fig. 1). These out- occurrence? Are soil depth and shading important? (3) crops are widely scattered and primarily occur in seven Do the models predict species to occur on mountain mountain ranges, each range 25±60 km from the nearest ranges where they are absent, which would suggest non-environmental control of distribution across moun- range. Compositional differences in vegetation among tain ranges? We conclude by examining the strengths outcrops relate to differences in elevation, geology, po- and weaknesses of habitat-based models for conser- tential solar radiation, and soil chemistry (Wiser et al. vation applications. 1996). Wiser et al. (1996) provide a detailed description of the study area and dominant compositional and en- METHODS vironmental gradients. Study species Data collection Predictive models were constructed for four rare, high-elevation rock-outcrop endemics: Geum radiatum As part of a larger study on outcrop vegetation, pres- Michx. (Rosaceae), Solidago spithamaea M. A. Curtis ence/absence of the four study species was recorded in (Asteraceae), Calamagrostis cainii A. S. Hitchcock 154 10 3 10 m (100-m2) plots distributed among 42 (Poaceae), and Houstonia purpurea L. var. montana peaks in western North Carolina and eastern Tennessee. (Small) Terrell (Rubiaceae) (hereafter Geum, Solidago, Plots were located to include all possible combinations Calamagrostis and Houstonia, respectively; botanical of major site parameters such as elevation, slope, as- nomenclature follows Kartesz 1994). All four are pect, geology, and topographic position. All known ranked as critically imperiled globally (G1) by the Na- populations of the four target species were included. ture Conservancy and are listed, or proposed for listing, Within each 100-m2 plot, two to seven 1 3 1 m (1-m2) as either threatened or endangered by the U.S. Fish and subplots were sampled, depending on the internal het- Wildlife Service (Murdock and Sutter 1987, Murdock erogeneity of the larger plot. For 100-m2 plots, the 1993, Saunders 1996). These species were chosen for following site parameters were determined: latitude, study because they are globally rare, and also because elevation, slope percentage, aspect, bedrock type, pres- they occurred with suf®cient frequency in the study ence of perennial seepage, percentage of exposed bed- area to allow adequate sampling. In the typology of rock, and degree of fracturing (using an index such that Rabinowitz (1981:210), these species have small geo- nearly smooth outcrops containing ledges and cracks graphic ranges and narrow habitat requirements (rel- ,0.3 m wide were classi®ed as ``1,'' and highly frac- ative to the landscape as a whole), and are ``classic tured and angular outcrops containing ledges and rarities in the sense of restricted endemics.'' Local pop- cracks .0.3 m wide were classi®ed as ``3''). The av- ulation sizes vary from a few dozen individuals to hun- erage topographic position of each plot was recorded dreds. Life-history characteristics (which are mostly as an index from 0 to 10 with 1 being the base of the poorly understood) and population data have been as- outcrop, and 10 being the top.
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