Feral Swine Sus Scrofa: a New Threat to the Remaining Breeding Wetlands of the Vulnerable Reticulated Flatwoods Salamander Ambystoma Bishopi

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Feral swine Sus scrofa: a new threat to the remaining breeding wetlands of the Vulnerable reticulated flatwoods salamander Ambystoma bishopi

K ELLY C. JONES,THOMAS A. GORMAN,BRANDON K. RINCON,JOHN A LLEN C AROLA A. HAAS and R ICHARD M. ENGEMAN

Abstract Feral swine Sus scrofa have been implicated as a Keywords Ambystoma bishopi, endangered species, major threat to sensitive habitats and ecosystems as well ephemeral ponds, hog control, hog damage, invasive spe- as threatened wildlife. Nevertheless, direct and indirect im- cies, rooting, Sus scrofa pacts on threatened species (especially small, fossorial species) are not well documented. The decline of the U.S. federally endangered reticulated flatwoods salamander Introduction Ambystoma bishopi, categorized as Vulnerable on the IUCN Red List, has been rapid and there are few remaining nvasive feral swine Sus scrofa cause extensive damage breeding locations for this species. The flatwoods salaman- Ito natural systems and affect many species of native wild- der depends on complex herbaceous vegetation in all life life and vegetation, both directly and indirectly (Seward stages, including eggs, larvae and adults. Historically sets et al., ). In addition, feral swine have been implicated of hog tracks have been observed only occasionally in the as a major threat to sensitive habitats and ecosystems vicinity of monitored reticulated flatwoods salamander (Engeman et al., ). The negative impact of exotic species breeding wetlands, and damage to the wetlands had never on native species and ecosystems is exceeded only by – been recorded. However, during the autumn winter breed- human-caused habitat destruction (Wilcove et al., ; – ing season of we observed a large increase in hog Parker et al., ). In the USA exotic species have played sign, including extensive rooting damage, in known flat- a role in the listing of % of the species protected by the woods salamander breeding wetlands. Our objective was Endangered Species Act (Stein & Flack, ). Worldwide, to assess the amount of hog sign and damage in these wet- invasive species are a leading contributor to extinctions of lands and to take corrective management actions to curb birds, fish and mammals (Clavero & García-Berthou,  additional impacts. Of wetlands surveyed for hog sign, ). Unlike many invasive species that may have primarily   presence was recorded at %, and damage at %. Of the direct effects on other species through predation or compe-  sites known to be occupied by flatwoods salamanders in tition, feral swine can have both direct effects as predators –   , % had presence, and % had damage. We and indirect effects by drastically altering habitats (Zengel found that regular monitoring of disturbance in wetland ha- & Conner, ; Barrios-García & Ballari, ; Rossell bitats was a valuable tool to determine when intervention et al., ). Non-native predators tend to be a greater threat was needed and to assess the effectiveness of intervention. than native predators to prey populations (Salo et al., ). Habitat damage caused by feral hogs poses a potentially ser- Feral swine have been implicated as a threat to several threa- ious threat to the salamanders, which needs to be mitigated tened wildlife species in the USA, including the lesser using methods to control and exclude hogs from this sensi- prairie-chicken Tympanuchus pallidicinctus, shorebirds tive habitat. and marine turtles (USDA, ; Engeman et al., , ), but direct and indirect impacts on small, fossorial species, including amphibians, have not been as well documented. Land managers tasked with the recovery of endangered wildlife require an increased understanding of the

KELLY C. JONES (Corresponding author), THOMAS A. GORMAN*, BRANDON existence and nature of conflicts between feral swine and K. RINCON,CAROLA A. HAAS Department of Fish and Wildlife Conservation, threatened amphibians. Virginia Tech, Blacksburg, Virginia 24061, USA. E-mail [email protected] Reticulated flatwoods salamanders Ambystoma bishopi JOHN ALLEN Animal and Plant Health Inspection Service, U.S. Department of and frosted flatwoods salamanders A. cingulatum have Agriculture, Niceville, Florida, USA been identified, along with three other amphibian species RICHARD M. ENGEMAN National Wildlife Research Center, U.S. Department of Agriculture, Fort Collins, Colorado, USA in Florida that are either federally or state listed, as being threatened by the activities of feral hogs (USDA, ). *Current address: Aquatic Resources Division, Washington State Department of Natural Resources, Chehalis, Washington, USA Although direct predation of salamanders by feral swine  Received  May . Revision requested  July . has been reported (Howe et al., ), it has been suggested Accepted  October . First published online  February . that habitat disturbance from extensive rooting has the most

Oryx, 2018, 52(4), 669–676 © 2017 Fauna & Flora International doi:10.1017/S0030605316001253 Downloaded from https://www.cambridge.org/core. University Libraries - Virginia Tech, on 10 Mar 2020 at 17:52:39, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605316001253 670 K. C. Jones et al.

serious impact on amphibians with specialized habitat re- frequent drought years, smoke management risks that in- quirements (Means & Travis, ). Feral hogs create short- herently accompany prescribed fire, and slow natural dis- term disturbance of vegetation and soil, removing emergent persal by the salamanders into recently restored habitat, plants and exposing bare soil (Arrington et al., ; Doupé the species faces a long, difficult road to recovery. et al., ; Rossell et al., ). They can also affect the Feral swine are relatively ubiquitous at Eglin, where their long-term vegetation structure, as certain plants appear to damage to wetlands has been documented for many years respond positively and others, such as seedlings of the long- (Engeman et al., ). Nevertheless, for .  years only oc- leaf pine Pinus palustris, which as canopy trees provide an casional sets of hog tracks had been observed in the vicinity important source of fuel to maintain the fire-dominated of monitored flatwoods salamander breeding wetlands, and ecology of this habitat, may be consumed preferentially by damage to the sites had never been documented prior to hogs (Simberloff, ; Arrington et al., ; Means, . However, while surveying for larval flatwoods sala- ; Boughton & Boughton, ). These impacts are par- manders during the – autumn–winter breeding ticularly challenging for flatwoods salamanders because season we observed a significant increase in hog sign, in- these species rely on hydric and mesic longleaf pine–wire- cluding extensive damage from rooting in known breeding grass Aristida stricta savannah habitat for both the aquatic wetlands of the flatwoods salamander. The addition of this and terrestrial phases of their life cycles (Palis, ). new threat to an already challenging recovery poses a poten- The reticulated flatwoods salamander was listed as feder- tially serious problem, the severity of which needs to be ally endangered in  (USFWS, ) and is categorized assessed. as Vulnerable on the IUCN Red List (Palis & Hammerson, Effects of invasive species on rare species may be more ). The decline of the species has been rapid, and fewer pronounced in a location such as Florida, where rare species than  breeding wetlands across the species’ range are may be particularly vulnerable because extensive develop- known to have been occupied in recent years (USFWS, ment has depleted many of the native habitats on which ; Semlitsch et al., ). Eglin Air Force Base in they depend (Engeman et al., ). Invasive species often Florida is one of few remaining public lands where this spe- present novel control situations for managers, requiring the cies occurs, and it is the only remaining property within the acquisition of biological knowledge and the development entire range of the species to support two populations, each and testing of control technologies and strategies (e.g. occurring in wetland complexes having .  occupied wet- Engeman & Vice, ). Our objective was to assess the lands (Gorman et al., ). amount of hog sign and damage in these wetlands and to The flatwoods salamander depends on complex herb- take corrective management actions to curb additional im- aceous vegetation for all aspects of its life history. The eco- pacts from this invasive mammal. We report on our moni- tone between deeper wetland areas and uplands provides toring procedure as well as the management practices to a diverse habitat structure, and this type of shallow, well- document the benefits of regular monitoring in facilitating vegetated littoral zone is known to be important to many a rapid response. species of pond-breeding amphibians (Porej & Hethering- ton, ; Shulse et al., ). Flatwoods salamanders lay their eggs in dense herbaceous ground cover in the ecotone Study area (Gorman et al., ), and developing larvae feed and seek cover there as well (Sekerak et al., ; Gorman et al., ). Our study area comprised longleaf pine–wiregrass flat- Frequently metamorphs and adults are observed climbing woods with scattered ephemeral wetlands of various and perching above the ground in wiregrass near breeding sizes on Eglin Air Force Base, Okaloosa County, Florida. wetlands (Jones et al., ). This dense understorey herb- Within this area we selected  wetlands, of which  were aceous vegetation is maintained by the presence of an historical breeding sites for reticulated flatwoods salaman- open pine canopy, which provides fine fuels and a short ders. Eleven of the  historical sites were occupied by sala- (– years in uplands, – years in wetlands) fire return mander larvae in –. The selected wetlands typically interval, although in much of the salamander’s range fire had an overstorey consisting of a combination of slash pine suppression has facilitated the encroachment of midstorey Pinus elliottii, pond cypress Taxodium ascendens and black shrubs that shade and change this habitat (Bishop & gum Nyssa sylvatica, a midstorey consisting of some com- Haas, ; Gorman et al., ). bination of myrtle dahoon Ilex cassine var. myrtifolia and In the face of many complex challenges, habitat manage- titi Cyrilla racemosa as well as the aforementioned over- ment for reticulated flatwoods salamanders at Eglin has in- storey species, and an understorey of wiregrass, Curtiss’ creased significantly since  and includes mechanical sandgrass Calamovilfa curtissii, longleaf threeawn Aristida removal of midstorey shrubs in wetland basins (e.g. palustris, pipeworts Eriocaulon spp., shortbristle horned Gorman et al., ) and use of growing-season prescribed beaksedge Rhynchospora corniculata, and a rich diversity fire. However, with continuing challenges associated with of graminoids, forbs and small shrubs.

Methods were resurveyed during the same period following  December . Moreover, following the initial reports of Damage assessment damage, U.S. Department of Agriculture Wildlife Services personnel based at Eglin immediately (April ) began Following our initial observations of hog damage in several implementing hog trapping across all affected areas. Swine wetlands during the salamander breeding season in winter captured in pen traps were euthanized. The swine trapping –  , we surveyed wetlands for hog sign and dam- process required identifying the most favourable locations  –   age during April August . These included all to carry out control activities, placing baits (soured corn), wetlands that were known to be occupied by flatwoods sala- constructing pen traps around bait piles that were visited – mander larvae during the breeding season, as well consistently by swine, allowing the swine to acclimatize to  as six that have been occupied historically and potential the presence of the traps, and then setting the traps for cap- breeding sites, six of which are near historically or recently ture. The traps were custom-designed and collapsible for occupied sites, and five of which are in an area that is under- portability, durable, and large enough to capture groups of going restoration to support future recovery efforts for flat- feral swine, including the largest individuals (Engeman woods salamanders. et al., ). Sites were surveyed by walking along the approximate perimeter of the wetland within the ecotone (wetland edge, heavily used by breeding salamanders), and along per- Statistical analyses pendicular transects into the approximate centre of the wet- – land basin, at intervals of m, depending on visibility. The prevalence of damage caused by feral hogs was com- Additional surveys for hog sign were conducted along adja- pared among the three categories of wetlands, based on  cent roads and surrounding uplands within m of the wet- use by flatwoods salamanders (potential breeding sites, cur- land edges. At each site we recorded the type(s) of hog sign rently occupied breeding sites, and historical breeding sites), observed (tracks, scat, rooting damage), location of damage using Fisher’s exact test. For breeding sites with damage, the ,  (wetland basin, ecotone, uplands m of outer edge percentage of damage within the ecotone (where most of the of ecotone), approximate area damaged or disturbed by suitable vegetation associated with breeding occurs in our rooting, and percentage of overall wetland disturbed by sites) was compared to that observed in the associated wet- rooting. We also assessed whether damage was focused on land as a whole using repeated measures ANOVA. This herbaceous vegetation, woody vegetation, or a near-equal would indicate whether the salamanders’ potential breeding combination of the two. Observers had several years of ex- locations were preferred rooting habitat of feral swine. perience identifying hog damage in both upland and wetland habitats, as well as training from U.S. Department of Agriculture Wildlife Services personnel. Results    Mitigation activities Hog presence was recorded in of the ( %) wetlands surveyed, and damage caused by hogs was recorded in  of Our initial reporting of hog damage led natural resources those  (% of wetlands with hog presence, or % of total personnel at Eglin to install hog exclusion fencing in two sites). Critically, six of the  damaged sites had $ %of areas that enclosed five of the  flatwoods salamander the wetland ecotone damaged by feral hog activity, and breeding sites that were occupied in –, as well as two of those six were occupied by flatwoods salamanders some surrounding uplands. In total, , m of fencing at that time (Plate ). In all  sites where hog damage was was required, at a cost of USD . per m, including the recorded, the damage was focused exclusively on herb- cost of site preparation, equipment, labour and materials. aceous vegetation. Materials included . mmetalT-posts, gauge . × . m The prevalence of hog presence among the historical (″ × ′) hog fence panels (Behlen Manufacturing Co., breeding wetlands was %( of  sites), with %( of Columbus, USA), and wire clips. The fence panels had a ) of those having damage (% of the  historical breeding mesh width of  cm, and a mesh height ranging from sites). Of paramount concern, hogs were present and da-  cm at the bottom to  cm at the top. We surveyed these maged %( of  sites) of the breeding wetlands that fenced breeding wetlands for hog presence a minimum of were occupied by reticulated flatwoods salamanders in three times following the completion of the installation on –. Hog sign was present but there was no observed  December , to ensure that no hogs had been enclosed damage in %( of ) of current known breeding sites, and within the fence perimeters. Fifteen unfenced sites, includ- no sign or damage was observed in four other wetlands ing the remaining six breeding sites that were occupied in (%) in this category. Thus, although a little more than  and initially had been surveyed for hog presence, half of the currently occupied breeding sites were damaged,

which eight had recent hog damage during initial surveys (%), six (%) had recent hog rooting damage (December –May ) and were the only sites with any observed hog sign. Trapping efforts began in early April  across the three areas where all of our monitoring sites were located, and a total of  feral hogs had been trapped by  May  in two major areas of concern ( in one and  in the other). In one of these areas, trapping efforts were made both inside and outside the fence perimeter but hogs were trapped only outside the fenced area, and to date there has been no observed evidence of hogs within the fenced areas. No hogs have yet been trapped in the third area, largely because of frequent tampering of the traps by people.

Discussion

Our assessments indicate that feral swine are damaging habitat that is critical to egg laying (Gorman et al., ), larval development (Gorman et al., ), and above-ground activity of metamorph and adult flatwoods salamanders (Jones et al., ). Our findings show that whether a site is currently used for breeding, known to have been used for breeding previously, or could potentially be used for breeding, if hogs are present there is a high likelihood they will damage the site. These habitats are characterized by

PLATE 1 (a) Intact flatwoods wetland ecotone, and (b) flatwoods dense coverage of herbaceous vegetation, including wetland ecotone with recent rooting damage to herbaceous Aristida spp., Eriocaulon spp. and Xyris spp. among others vegetation caused by feral swine Sus scrofa, on Eglin Air Force (Gorman et al., ), and we found that it was exactly these Base, Florida, USA. complex herbaceous plant communities that were the focus of hog rooting in these valuable wetlands. There are few remaining known breeding sites for the federally endangered % of sites where hog presence was recorded were da- reticulated flatwoods salamander, and the damage caused by maged (Fig. ). feral swine in Eglin Air Force Base flatwoods ponds was ef- No differences in prevalence of damage were detected fectively reversing wetland restoration efforts aimed at in- (Fisher’s exact test, P = .) among potential breeding creasing the coverage of complex herbaceous vegetation sites (%), current breeding sites (%), and historical (e.g. Gorman et al., ). Given the importance of the breeding sites (not including sites occupied in –; breeding sites at Eglin for the species, persistent damage %). Of concern for salamander breeding success, the to these sites by feral swine could jeopardize the long-term mean percentage of area damaged by rooting within wetland survival of the species and increase the cost of future recov-  ±       ecotones ( SE . %) was higher (F, = . ,P= . ) ery actions. than for the associated wetlands as a whole (. ± SE .%), We knew from an ongoing study that, for several weeks indicating that areas most suitable for salamander breeding leading up to these observations, dozens of recently meta- may also be attractive to rooting swine. morphosed reticulated flatwoods salamanders had been During follow-up surveys (December –May ) moving out of the breeding pond basin into areas damaged for hog presence within fenced areas, no hog sign was de- by hogs. We have also observed metamorphs sheltering tected inside the perimeter of any of the fenced areas. Hog under clumps of wiregrass near the surface while still in sign and damage were detected both in the general vicinity the vicinity of the breeding site (Jones et al., ). Threats and immediately along the outside of the fenced areas, indi- to metamorphs sheltering in this way were indicated on  cating that hogs were still present, although they were being May , when we observed the partial remains of an ap- excluded from these areas. Furthermore, of the  unfenced parently swine-predated eastern glass lizard Ophisaurus sites that were resurveyed during this same time period, of ventralis in a freshly rooted patch of vegetation in the

FIG. 1 Proportion of wetlands (n = ; grouped according to presence of hog Sus scrofa sign and damage) on Eglin Air Force Base, Florida, USA, used by reticulated flatwoods salamanders Ambystoma bishopi, based on surveys conducted during  April– August . Known (n = ) represents all sites where the occurrence of A. bishopi was ever recorded, including sites detected during –; Detected – (n = ) represents sites occupied during the – breeding season; and Potential (n = ) represents sites where the species was not detected but which have potential breeding habitat.

wiregrass-dominated ecotone of one of the few remaining local residents, reduces the harvest pressure on native wild- breeding wetlands occupied by reticulated flatwoods sala- life, and limits growth of the hog population (Desbiez et al., manders (Jones et al., ). If swine can depredate a glass ), in the USA and Australia hunting of feral swine seems lizard while rooting, it seems likely that they could also de- to be self-reinforcing, with hunters (often illegally) trans- predate a flatwoods salamander. For many ephemeral pond- porting and releasing hogs to new areas for sport-hunting breeding species successful reproduction occurs sporadical- (Spencer & Hampton, ; Barrios-García & Ballari, ly, with most metamorphosis concentrated in time. In many ; Bevins et al., ). The rapid expansion of the range breeding wetlands used by flatwoods salamanders, includ- of feral swine in the USA is believed to be primarily a result ing the one where this observation occurred, successful re- of people transplanting them to increase recreational hunt- cruitment into the adult population occurs in , %of ing opportunities (USDA, ), and thus spreading their years (Gorman & Haas, ; Chandler et al., ). Given negative impacts through disease and damage to the envir- the concentration of new cohorts of amphibians in a re- onment, agriculture, wildlife (especially rare species), do- stricted space and time, there is potential for feral hogs to mestic animals, and other human interests (Seward et al., consume entire cohorts, posing a serious threat to the con- ; USDA, ). At Eglin Air Force Base recreational tinued existence of the species. hunting had been taking place for many years and was de- Since we recognized and reported the increase in hog ac- monstrated to have some impact on reducing hog numbers tivity in and around the breeding wetlands of flatwoods sal- and their damage to seepage slopes, but the beneficial im- amanders, focused trapping efforts have been implemented pacts from operational hog trapping were much greater in these areas. These efforts are ongoing and have reduced (Engeman et al., ). hog activity from  to % of resurveyed sites. Other mea- There have been reports of feral hogs consuming amphi- sures to control hogs may need to be considered in these vul- bians at other locations, and their ability to consume large nerable habitats of endangered species, as there is evidence numbers of amphibians during mass migrations to or from that people continue to release hogs into these sensitive breeding sites is a concern, especially for species limited to areas. At the five occupied breeding wetlands where fencing only a few remaining breeding sites (Schley & Roper, ; was installed in an attempt to exclude hogs, this appears to Jolley et al., ; Wishart et al., ). Feral hogs may also have been successful. The hog fencing has required some re- degrade the quality of breeding habitats by disturbing vege- pairs following damage by a falling tree or by adult black tation, spreading invasive plants, altering invertebrate com- bears Ursus americanus climbing over it but most checks munities (which are both important prey and predators of yielded no observations of damage. The potential breeding salamanders), reducing dissolved oxygen, and altering the area where human interference rendered trapping efforts in- microtopography (e.g. Kaller & Kelso, ; Engeman effective has now been closed to motorized vehicles. et al., ; Bankovich et al., ). Such destructive effects Whereas in the Brazilian Pantanal, for example, hunting have been observed elsewhere, including at breeding sites of of feral hogs provides an important source of protein to the Houston toad Bufo houstonensis, another federally

endangered amphibian (Brown et al., ). Our results sup- analyses. KJ guided and oversaw fence installation. JA pro- port the conclusions of others that fencing (supported by vided training in hog sign identification, conducted swine trapping) can mitigate the negative effects of feral hogs on trapping, and recorded trapping success. KJ, TG, RE and wetland-breeding amphibians and reptiles (Doupé et al., CH wrote the article, and all authors contributed to editing ; Brown et al., ). Although there may be negative the article. consequences of using hog fencing around wetlands, such as excluding alligators and large turtles, other control measures cannot fully protect threatened amphibians and sensi- References tive habitats. Trapping at Eglin has been demonstrated to   ARRINGTON, D.A., TOTH, L.A. & KOEBEL, J.W. ( ) Effects of substantially reduce hog numbers (Engeman et al., ). rooting by feral hogs Sus scrofa L. on the structure of a floodplain However, eradication of hogs from Eglin is not feasible be- vegetation assemblage. Wetlands, , –. cause of a lack of complete access to an extensive area of BANKOVICH, B., BOUGHTON, E., BOUGHTON, R., AVERY, M.L. & challenging terrain, boundaries with rivers and terrain WISELY, S.M. () Plant community shifts caused by feral swine that cannot be maintained to exclude hogs, and illegal trans- rooting devalue Florida rangeland. Agriculture, Ecosystems & Environment, , –. location of hogs to the area by local people for recreational BARRIOS-GARCÍA, M.N. & BALLARI, S.A. () Impact of wild boar hunting purposes. For an extremely restricted habitat of a (Sus scrofa) in its introduced and native range: a review. Biological rare species such as the reticulated flatwoods salamander, Invasions, , –. a single hog can have a significant negative impact. Thus, BEVINS, S.N., PEDERSEN, K., LUTMAN, M.W., GIDLEWSKI,T.&  fencing the breeding sites serves as a localized eradication DELIBERTO, T.J. ( ) Consequences associated with the recent range expansion of nonnative feral swine. BioScience, , –. of hogs from around the breeding sites. During the study   BISHOP, D.C. & HAAS, C.A. ( ) Burning trends and potential period, fencing reduced damage by %. negative effects of suppressing wetland fires on flatwoods Our ability to identify the disturbance quickly through salamanders. Natural Areas Journal, , –. monitoring efforts, in concert with the presence of trained BOUGHTON, E.H. & BOUGHTON, R.K. () Modification by an wildlife control professionals, facilitated a rapid response invasive ecosystem engineer shifts a wet prairie to a monotypic  – to conserve the reticulated flatwoods salamander. We en- stand. Biological Invasions, , . BROWN, D.J., JONES, M.C., BELL,J.&FORSTNER, M.R.J. () Feral courage others who manage land that supports populations hog damage to endangered Houston toad (Bufo houstonensis) of rare or threatened species to institute pre-emptive prac- habitat in the Lost Pines of Texas. Texas Journal of Science, , tices. A first step is to make sure that personnel on the –. ground know how to identify likely invasive species and CHANDLER, H.C., RYPEL, A.L., JIAO, Y., HAAS, C.A. & GORMAN, T.A.  their signs, and to liaise with damage management profes- ( ) Hindcasting historical breeding conditions for an endangered salamander in ephemeral wetlands of the southeastern USA: sionals for advice on how to survey and respond. Regular implications of climate change. PLoS ONE, (), e. site visits should then be implemented to monitor sensitive CLAVERO,M.&GARCÍA-BERTHOU,E.() Invasive species are a areas, using survey techniques such as those described here leading cause of animal extinctions. Trends in Ecology and or by Zengel & Conner (). We recommend that land Evolution, , . managers work with invasive species control specialists to DESBIEZ, A.L.J., KEUROGHLIAN, A., PIOVEZAN,U.&BODMER, R.E. () Invasive species and bushmeat hunting contributing to implement techniques that will reduce or prevent damage wildlife conservation: the case of feral pigs in a Neotropical wetland. to sensitive habitats and species. Oryx, , –. DOUPÉ, R.G., SCHAFFER, J., KNOTT, M.J. & DICKY,P.W.()A description of freshwater turtle habitat destruction by feral pigs in Acknowledgements tropical north-eastern Australia. Herpetological Conservation and Biology, , –. We thank the Natural Resources Branch (Jackson Guard) of ENGEMAN, R.M., ADDISON,D.&GRIFFIN, J.C. () Defending Eglin Air Force Base and the Department of Fisheries and against disparate marine turtle nest predators: nesting success Wildlife Sciences at Virginia Tech for their financial and lo- benefits from eradicating invasive feral swine and caging nests from raccoons. Oryx, , –. gistical support. We specifically thank the leadership and ENGEMAN, R.M., CONSTANTIN, B.U., GRUVER, K.S. & ROSSI,C. supporting staff at Jackson Guard for their rapid response, () Managing predators to protect endangered species and Sandy Pizzolato for coordinating fence installation, and the promote their successful reproduction. In Endangered Species: New U.S. Fish and Wildlife Service Panama City Ecological Research (eds A.M. Columbus & L. Kuznetsov), pp. –. Nova Services team for permitting support. Science Publishers, Hauppauge, USA. ENGEMAN, R.M., DUFFINEY, A., BRAEM, S., OLSEN, C., CONSTANTIN, B.U., SMALL,P.etal.() Dramatic and immediate improvements in insular nesting success for threatened sea turtles and shorebirds Author contributions following predator management. Journal of Experimental Marine Biology and Ecology, , –. KJ and TG developed the survey methodology, and KJ and ENGEMAN, R.M., STEVENS, A., ALLEN, J., DUNLAP, J., DANIEL, M., BR conducted the damage surveys. RE conducted statistical TEAGUE,D.&CONSTANTIN,B.() Feral swine management for

conservation of an imperiled wetland habitat: Florida’s vanishing SALO, P., KORPIMÄKI, E., BANKS, P.B., NORDSTRÖM,M.&DICKMAN, seepage slopes. Biological Conservation, , –. C.R. () Alien predators are more dangerous than native ENGEMAN, R.M. & VICE, D.S. () Objectives and integrated predators to prey populations. Proceedings of the Royal Society of approaches for the control of brown tree snakes. Integrated Pest London B, , –. Management Reviews, , –. SCHLEY,L.&ROPER, T.J. () Diet of wild boar Sus scrofa in western GORMAN, T.A. & HAAS, C.A. () Evaluation of Mechanical Europe, with particular reference to consumption of agricultural Mid-Story Removal as a Tool to Restore Endangered Species Habitat. crops. Mammal Review, , –. Final report. DOD Legacy Resource Management Program. Virginia SEKERAK, C.M., TANNER, G.W. & PALIS, J.G. () Ecology of Tech, Blacksburg, USA. flatwoods salamander larvae in breeding ponds in Apalachicola GORMAN, T.A., HAAS, C.A. & BISHOP, D.C. () Factors related to National Forest. Proceedings of the Annual Conference of the occupancy of breeding wetlands by flatwoods salamander larvae. Southeastern Association of Fish and Wildlife Agencies, , Wetlands, , –. –. GORMAN, T.A., HAAS, C.A. & HIMES, J.G. () Evaluating methods SEMLITSCH, R.D., WALLS, S.C., BARICHIVICH, W.J. & O’DONNELL, to restore amphibian habitat in fire-suppressed pine flatwoods K.M. () Extinction debt as a driver of amphibian declines: an wetlands. Fire Ecology, , –. example with imperiled flatwoods salamanders. Journal of GORMAN, T.A., POWELL, S.D., JONES, K.C. & HAAS, C.A. () Herpetology, , –. Microhabitat characteristics of egg deposition sites used by SEWARD, N.W., VERCAUTEREN, K.C., WITMER, G.W. & ENGEMAN, reticulated flatwoods salamanders. 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Biographical sketches longleaf pine system. JOHN ALLEN has been the lead technician in the implementation of the Eglin feral hog management pro- KELLY JONES coordinates field operations for Virginia Tech’swork gramme since it began in . He has also conducted beaver con- on threatened species at Eglin Air Force Base. His interests include trol, threatened and endangered species protection, and bird/ natural history, habitat management and conservation of rare spe- animal aircraft strike hazard-related control work. CAROLA HAAS cies and ecosystems. THOMAS GORMAN manages aquatic resources conducts research on wildlife populations and behaviour in mana- – and conducts research on wildlife habitat relationships, habitat ged habitats. RICHARD ENGEMAN carries out research to develop management, and approaches to conserving amphibians and rep- wildlife indexing and ecological sampling methods, primarily fo- tiles. BRANDON RINCON is a wildlife biologist whose interests in- cusing on applications for management of invasive species and clude ecology and ecosystem management, especially within the protection of threatened species.