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Evaluating Stability in Ziziphus Celata, a Highly Endangered Clonal Shrub Endemic to Lake Wales Ridge, Central Florida

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Evaluating Stability in Ziziphus Celata, a Highly Endangered Clonal Shrub Endemic to Lake Wales Ridge, Central Florida Printed October 2007 Vol. 3: 125–132, 2007 ENDANGERED SPECIES RESEARCH Endang Species Res Published online July 10, 2007 Evaluating stability in Ziziphus celata, a highly endangered clonal shrub endemic to Lake Wales Ridge, central Florida Martha M. Ellis1,*, Carl W. Weekley2, Eric S. Menges2 1Wildlife Biology Program, University of Montana, Missoula, Montana 59812, USA 2Archbold Biological Station, PO Box 2057, Lake Placid, Florida 33862, USA ABSTRACT: Florida ziziphus Ziziphus celata is a self-incompatible clonal shrub comprising fewer than a dozen remnant, uniclonal populations in south-central Florida, USA. We model the population viability of this highly endangered species based on 9 yr of data from 2 populations. The matrix pro- jection model is based on hypothetically independent ‘plants’ defined as clumps of ramets within a 25 cm radius. Seedling recruitment is unknown in the uniclonal study populations and is not mod- eled. The populations modeled here have stable demographies characterized by high survival and stasis, but variable levels of new plant production (clonal recruitment). Population growth rates sug- gest protracted long-term declines in population size, but predicted extinction rates over 50 yr are as high as 20% in one population. Changes in new plant production and survival had greater effects on population growth rates than did changes in growth rates of individual plants. Augmenting clonal plant production and protecting survival of new ‘plants’ are short-term management goals. The establishment of sexually reproducing populations through the translocation of cross-compatible genotypes is a long-term necessity for the persistence of Florida ziziphus populations. KEY WORDS: Population viability analysis · Florida ziziphus · Clonal reproduction · Endangered species · Rare plant conservation Resale or republication not permitted without written consent of the publisher INTRODUCTION (Menges 2000b). In some species, where clonal growth can be directly tied to clearly defined individuals with Population viability analysis (PVA) is a powerful tool aboveground clonal connections, a PVA can proceed for assessing the sustainability (or extinction risk) of with the addition of a ‘new clonal ramet’ life history plant populations (Menges 2000a, Morris & Doak 2002, stage (e.g. Eriksson 1988, Weppler et al. 2006). In some Reed et al. 2002). Stage-based matrix models incorpo- cases, both ramets and genets can be distinguished, rating the various sources of stochasticity (environ- and modeling can be carried out at both ramet and mental, demographic, genetic) have been increasingly genet levels, which may differ in their predictions used in recent years to aid conservationists in identify- (Eriksson 1994, Damman & Cain 1998, Colling & ing demographic threats to the persistence of rare Matthies 2006). In populations with high genetic varia- plant populations (e.g. Liu et al. 2005, Smith et al. 2005, tion, genotyping can be combined with demographic Maschinski et al. 2006, Meyer et al. 2006). modeling to provide demographic analyses (Kynci et Most PVAs for plants have not considered those with al. 2006). However, because genets cannot always be clonal growth (Menges 2000b). Clonal species con- easily identified, most clonal plant PVAs use ramet stitute a special challenge for PVAs because of the level data. difficulty of distinguishing genets (genetically distinct Ramet population viability varies among sites individuals) from ramets (modular units of a genet (Boeken & Canham 1995), among years (Huenneke & which may be physiologically independent) in the field Marks 1987), between the clone periphery and its cen- *Email: [email protected] © Inter-Research 2007 · www.int-res.com 126 Endang Species Res 3: 125–132, 2007 ter (e.g. Wikberg & Svensson 2006), and with harvest- Selection of study populations. We have conducted ing levels (Nault & Gagnon 1993). Even in species demographic research in all known Florida ziziphus without known sexual reproduction, ramet-based populations since 1997. However, small sample sizes PVAs can compare various scenarios of decline and disturbances to marked plants limit the number of (Schwartz et al. 2000). Here, we generate a PVA for a cases in the dataset. For example, mowing and dam- clonal species in which most ‘populations’ consist of a age by cattle have resulted in tags being displaced single genet. We evaluate the role of clonal reproduc- from marked plants, thereby reducing sample sizes tion in the long-term population dynamics of Florida for those populations. For this study, we chose 2 ziziphus Ziziphus celata, a federally endangered shrub populations for analysis based on many censuses and endemic to the Lake Wales Ridge of central-peninsular larger samples sizes. One population (PO3) occurs in Florida (USFWS 1999). a remnant sandhill site and one (PO4) in a pasture. Our goals in the present study are to develop stage- The PO3 population occupies a 250 m2 site character- based models of the ramet dynamics of Florida ized by plants typical of Florida sandhill (e.g. longleaf ziziphus and to use these models as the basis for a pine, wiregrass, and several sandhill endemics), but population viability analysis. Modeling the dynamics surrounded by exotic vegetation. In contrast, the PO4 of Florida ziziphus’ uniclonal populations will con- population occupies a 625 m2 site converted to pas- tribute to its recovery in 2 ways: (1) the critical life his- ture several decades ago. The PO3 and PO4 datasets tory stages of the long-lived plants that constitute these include 63 and 263 records of individual histories, ‘populations’ will be identified, and (2) it allows us to respectively. During the study period, the number of infer the fate of these remnant populations given the live plants in the PO3 population ranged from 17 to current absence of sexual reproduction. 28. At PO4 the number of live plants ranged from 29 to 143. Data collection. We censused the 2 study popula- MATERIALS AND METHODS tions annually, beginning in January 1997. We defined a ‘plant’ as a group of ramets within a 25 cm radius of Study species. Florida ziziphus Ziziphus celata Judd one another; this rule was based on observations of & D. Hall (Rhamnaceae) is a narrowly endemic and groups of stems that appeared physiologically inde- critically endangered shrub (USFWS 1999, Ward et al. pendent. While this definition was arbitrary, some way 2003, Turner et al. 2006). At the time of its description of classifying individuals at a scale between the popu- in 1984 from a 36 yr old herbarium specimen, Florida lation and the individual stem was necessary. In some ziziphus was thought to be extinct (Judd & Hall 1984). cases, stems grew too closely together to access or cen- However, between 1988 (DeLaney et al. 1989) and sus individually. Using this rule, we defined new 2002, 9 remnant populations were discovered along a ‘plants’ (clonal recruits) as new stems (i.e. root shoots) 45 km stretch of the Lake Wales Ridge in south-central occurring >25 cm from the base of an existing plant or peninsular Florida. All populations occur on yellow new stems occurring within 25 cm of a plant that had sands that historically supported longleaf pine/wire- previously died back (i.e. resprouts of a pre-existing grass Pinus palustris/Aristida stricta var. beyrichiana plant). We marked each plant with a numbered alu- sandhills, although 6 populations now occupy grazed minum tag, and, in January/February of each year (the pastures subject to frequent mowing. Of the 9 pop- time of peak flowering), recorded its height, maximum ulations, 8 are uniclonal (Godt et al. 1997, Weekley crown diameter, number of stems, flowering stage et al. 2002) and, because Florida ziziphus is self- (vegetative or reproductive), and number of flowers. incompatible (Weekley & Race 2001), most populations Data collected from 1997 to 2005 were included in this are sterile. study, resulting in 8 annual projection matrices for Extant populations have apparently persisted for each population. several decades through clonal growth. Populations Model structure and analysis. We defined a 3-stage vary in size from a few dozen to several hundred life history for Florida ziziphus: new plants (clonal ramets and comprise more or less discrete clumps of recruits), vegetative plants, and reproductive plants. ramets which may function as eco-physiologically in- Because we have never observed seedling recruitment dependent entities (‘plants’). Only one small uniclonal in in situ Florida ziziphus populations, we did not population is publicly protected. While the recovery of model this or fecundity. To identify the best model Florida ziziphus requires the creation of sexually vi- structure, we used logistic regression to model survival able populations through the translocation of geneti- (Table 1) as a function of each of the following vari- cally diverse and cross-compatible genotypes (USFWS ables: height, number of stems, number of flowers 1999, Weekley et al. 2002), management of existing (estimated in 5 log-10 categories), a binary variable uniclonal populations is a critical intermediate task. indicating whether the individual was a new ramet in Ellis et al.: Population viability of Florida ziziphus 127 Table 1. Multiple regression predicting annual survival from size and stage the percentage of individuals present variables. Survival was regressed against height, number of stems, number of in one year that was present in the fol- flowers, whether the plant first appeared in the current year (new), and stage lowing year. An individual that was (combining age and flowering). The intercepts and coefficients for each variable are shown with the standard error associated with that estimate and missed in a given year (i.e. tag not associated p-values found) was included in these calcula- tions as surviving if it was found alive Population Variable Intercept Coefficient p in subsequent years. We calculated growth transition rates as the percent- PO3 Height 1.70 ± 0.34 0.00 0.996 age of surviving individuals that tran- No. of stems 1.83 ± 0.27 0.01 ± 0.08 0.885 sitioned from stage i to stage j (g ).
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