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Recruitment of Desert Tortoises (Gopherus Agassizii and G

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Recruitment of Desert Tortoises (Gopherus Agassizii and G Herpetological Conservation and Biology 10(2):583–591. Submitted: 29 September 2014; Accepted: 7 April 2015; Published: 31 August 2015. RECRUITMENT OF DESERT TORTOISES (GOPHERUS AGASSIZII AND G. MORAFKAI): A SYNTHESIS OF REPRODUCTION AND FIRST-YEAR SURVIVAL STEVEN P. CAMPBELL1,2,3, ROBERT J. STEIDL1, AND ERIN R. ZYLSTRA1 1School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona 85721, USA 2Albany Pine Bush Preserve Commission, Albany, New York 12205, USA 3Corresponding author, email: [email protected] Abstract.—Recruitment is integral to population persistence, therefore characterizing this process is essential for evaluating recovery actions for species in decline. We gathered all data available and used Bayesian analyses to quantify annual recruitment of Mojave Desert (Gopherus agassizii) and Sonoran Desert (G. morafkai) tortoises as the product of four components: proportion of females that reproduced, number of eggs produced per reproducing female, hatching success, and hatchling survival. For Mojave Desert Tortoises, the estimated proportion of females that reproduced (0.81 [95% Credible Interval: 0.52–0.99]) and number of eggs produced per year (6.90 [5.51–8.16]) were higher than for Sonoran Desert Tortoises (0.52 [0.07–0.94] and 5.17 [3.05–7.60], respectively). For Mojave Desert Tortoises, hatching success averaged 0.61 (0.25–0.90). Data on hatching success for Sonoran Desert Tortoises and hatchling survival for both species were sparse, therefore we represented these components with a range of plausible values. When we combined components, average recruitment for Mojave Desert Tortoises ranged from 0.51 females/female/y assuming that hatchling survival was 0.30 to 1.18 females/female/y with hatchling survival assumed to be 0.70. For Sonoran Desert Tortoises, average recruitment ranged from 0.25 to 0.57 females/female/y for the same values of hatchling survival. Differences in recruitment between species likely reflect the evolution of different life-history strategies for tortoises in Mojave and Sonoran Deserts, perhaps in response to variation in precipitation regimes. To better inform conservation and recovery of desert tortoises, more information is needed for all recruitment components, but especially for hatching success and hatchling survival. Key Words.—Bayesian analysis; conservation; demography; Mojave Desert; Sonoran Desert species, and disease (USFWS 1990, 2010, 2011; Darst et INTRODUCTION al. 2013; Gray and Steidl 2015), and they are likely vulnerable to a suite of emerging threats that includes Demography provides a foundation for exploring climate change (Zylstra et al. 2013; Lovich et al. 2014). processes that govern population dynamics and for These declines have led the U. S. Fish and Wildlife developing reliable strategies to recover species in Service to classify the Mojave Desert Tortoise as a decline. For many vulnerable species, however, threatened species and the Sonoran Desert Tortoise as a demographic information is limited and often imprecise, candidate for listing (USFWS 1990, 2010). Developing which can make it difficult to determine an appropriate effective conservation strategies for these species will course of action for recovery (Tear et al. 1995; require understanding the nature and extent to which Beissinger and Westphal 1998; Morris et al. 2002). these threats affect different life stages and how life Nevertheless, delaying recovery actions in anticipation stages interact to affect demography and population of additional information can prove detrimental for dynamics. Although progress has been made toward this species whose populations are declining (Grantham et al. goal (Bury and Germano 1994; Van Devender 2002; 2009). Therefore, it is often better to make weaker USFWS 2011; Rostal et al. 2014), information is still inference based on available data than to make no lacking for key aspects of their life histories. inference at all (Link and Barker 2010). Information on recruitment, which we define as the The need to develop and implement recovery actions number of offspring produced that survive their first based on limited demographic information is especially year, is especially limited. There have been few relevant for Mojave and Sonoran Desert Tortoises attempts to estimate recruitment for Mojave Desert (Gopherus agassizii and G. morafkai, respectively). Tortoises (Turner et al. 1987; Doak et al. 1994; Karl Across large portions of their geographic ranges, both 1998) and none for Sonoran Desert Tortoises, probably species are thought to be declining in response to an because different life-history events or components that array of threats that includes habitat loss, invasive comprise recruitment are challenging to study (Morafka Copyright. © 2015. Steven P. Campbell 583 All Rights Reserved. Campbell et al.—Recruitment of Desert Tortoises. 1994). For example, nests can be difficult to find where PR is the proportion of females in the population because they are usually located inside burrows and are 퐸 that reproduced, ⁄2 is the number of female eggs abandoned by female tortoises soon after they deposit produced per reproducing female per year, PH is the eggs (Averill-Murray et al. 2014). Likewise, hatchlings proportion of eggs that hatched successfully, and PS is can be difficult to find, mark, and track because they are the proportion of hatchlings that survived to the end of cryptic and small. In addition, studies of recruitment their first year. In our analyses, we assumed that the sex may be considered lower priority because rates of ratio was 50:50 at laying, that rates of hatching success population change for desert tortoises are thought to be and hatchling survival were equal for males and females, governed primarily by rates of adult survival (Congdon and that tortoises were surveyed annually immediately et al. 1993; Doak et al. 1994; Heppell et al. 1996; prior to reproduction, such that recruitment equates to Heppell 1998; but see Wisdom et al. 2000). egg production offset by hatchling survival. Although The lack of reliable information on recruitment has recruitment could be subdivided into more components made it difficult to evaluate the importance of this life- (e.g., PH could be divided into the proportion of eggs history stage on population dynamics. In particular, a that were fertile and the proportion of fertile eggs that lack of estimates of recruitment inhibits the use of stage- hatched successfully), we used components that were or age-structured population models, which integrate studied most commonly. information from multiple life-history stages to evaluate rates of population change, gauge the relative Data.—We assembled data on recruitment importance of different life-history stages to population components from journal articles, theses, and dynamics, predict population-level responses to threats government reports (Table 1). In our analyses, we used or management, assess long-term population viability, point estimates of components from each site and year and develop conservation strategies (Crouse et al. 1987; combination where data were available. Where raw data Caswell 2001; Morris et al. 2002; Morris and Doak were available, we estimated recruitment components 2002). The utility of single demographic rates for these ourselves (73% of estimates), but otherwise we used purposes is limited to the extent that a specific rate estimates as reported (27%). Although there is reflects other demographic processes in a population uncertainty associated with all estimates, we did not (Radchuk et al. 2013). For example, recovery actions to include variance of estimates in our analyses because increase adult survival might be misdirected if they were not provided for some reported estimates. recruitment or juvenile survival is limiting population We considered the proportion of females reproducing growth. Thus, recruitment estimates are necessary to (PR) and annual egg production (E) as the reproductive understand population dynamics, and the lack of portion of recruitment. For both components, we recruitment estimates currently impedes our ability to included only data from studies that used radiography to evaluate the relative efficacy of alternative recovery classify reproductive status and to quantify egg actions (USFWS 2010, 2011). production because other methods are unreliable (Turner Given the importance of understanding the role of et al. 1984). We excluded estimates of egg production recruitment in recovery of Sonoran and Mojave desert that were based on fewer than four tortoises (Table 1). tortoises, our primary objective was to estimate Hatching success (PH) describes reductions in recruitment of tortoises by assembling and synthesizing recruitment between egg laying and hatchling emergence data from studies across their geographic ranges in the from the nest due to egg breakage, infertility, incomplete United States. Despite limitations in the data available, development, and nest predation. We estimated this we estimate recruitment because of its central component as the number of hatchlings that emerged importance in assessing the viability of these vulnerable from all nests monitored at a site divided by the total species. Our secondary objectives were to summarize number of eggs in all the nests. Although fates of eggs research on recruitment of desert tortoises, identify in the same clutch are probably not independent, we deficiencies in available data, and recommend directions used eggs as sample
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