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Biology and Impacts of Pacific Island Invasive Species. 9 Biology and Impacts of Pacific Island Invasive Species. 9. Capra hircus, the Feral Goat (Mammalia: Bovidae)1 Mark W. Chynoweth,2,5 Creighton M. Litton,2 Christopher A. Lepczyk,2 Steven C. Hess,3 and Susan Cordell 4 Abstract: Domestic goats, Capra hircus, were intentionally introduced to numer- ous oceanic islands beginning in the sixteenth century. The remarkable ability of C. hircus to survive in a variety of conditions has enabled this animal to become feral and impact native ecosystems on islands throughout the world. Direct eco- logical impacts include consumption and trampling of native plants, leading to plant community modification and transformation of ecosystem structure. Al- though the negative impacts of feral goats are well known and effective manage- ment strategies have been developed to control this invasive species, large popu- lations persist on many islands. This review summarizes impacts of feral goats on Pacific island ecosystems and management strategies available to control this invasive species. Descended from the wild goat (C. aegagrus) self-perpetuating feral populations given their from central Asia (Zeder and Hesse 2000), ability to survive in a variety of habitats, on a domestic goats (Capra hircus) have been intro- wide variety of forage, and with limited water. duced to islands worldwide. The original pur- Goats have been considered by some to be pose of insular domestic goat introductions “the single most destructive herbivore” intro- was likely for sailors to populate oceanic is- duced to island ecosystems globally (King lands with a food source to access during later 1985:3). voyages (Campbell and Donlan 2005). Re- leased domesticated goats can quickly develop name Capra hircus (Linnaeus, 1758) 1 Manuscript accepted 23 August 2012. This material Synonyms: Capra hircus, Capra hircus ae- is based upon work supported by the National Science gagrus, Capra aegagrus hircus. Foundation Graduate Research Fellowship under Grant Common names: briar goat, brush goat, No. 2010094953 (to M.W.C.); USDA Forest Service, feral goat, goat, hill goat, scrub goat, Spanish Pacific Southwest Research Station, Institute of Pacific Islands Forestry (Research Joint Venture 08-JV- goat, wood goat. 11272177-074 to C.M.L.); and the W. T. Yoshimoto Foundation Endowed Fellowship in Animal Wildlife description and account of variation Conservation Biology (to M.W.C.). Support to S.C.H. was provided by the USGS Invasive Species Program. 2 Department of Natural Resources and Environmen- Goats are even-toed hoofed ungulates of the tal Management, University of Hawai‘i at Mänoa, Hono- order Artiodactyla and have been considered lulu, Hawai‘i 96822. to comprise from one to nine species (Shack- 3 U.S. Geological Survey Pacific Island Ecosystems leton and Shank 1984 and references therein). Research Center, Hawai‘i National Park, Hawai‘i 96718. 4 Feral goats on Pacific islands (Figure 1) are U.S. Department of Agriculture Forest Service, In- stitute of Pacific Islands Forestry, Hilo, Hawai‘i 96720. assumed to have been introduced by Europe- 5 Corresponding author (e-mail: chynoweth.mark@ an sailors as a food source and are, therefore, gmail.com). most likely derived from continental Europe- an domestic goat breeds. Feral goats exhibit substantial intraspecific variation and are sex- Pacific Science (2013), vol. 67, no. 2:141 – 156 doi:10.2984/67.2.1 ually dimorphic. Generally, males are 20% © 2013 by University of Hawai‘i Press larger and have larger horns than females All rights reserved (Fleming 2004). Both males and females have 141 142 PACIFIC SCIENCE · April 2013 Figure 1. Feral goats, Capra hircus, on Hawai‘i Island. (Photo: Mark Chynoweth) horns made of living bone surrounded by etary preferences. In general ungulates, in- keratin. Goats typically weigh between 25 and cluding feral goats, demonstrate preference 55 kg, stand 1–1.2 m at the shoulder, and are and avoidance at least partly based on foliage 1–1.5 m long. All males and some females chemistry (Forsyth et al. 2002). McCammon- are bearded as adults. Both sexes have 30 –32 Feldman (1980) suggested that goats actively teeth, with upper and lower teeth in the back select the highest-quality forage. Although to chew cud and a dental pad in place of upper the most palatable forage is typically sought incisors. Goats sometimes resemble sheep out and consumed first, poor-quality forage but can be distinguished by their short, up- is often used to sustain populations (Coblentz ward pointing tails. Pelage coloration is typi- 1977, Green and Newell 1982). Consequent- cally black, but individuals can be white, ly, feral goats can extirpate preferred forage gray, brown, red, black, or any combination species (Coblentz 1977). thereof. Goats are often regarded as browsers. However, tendency to graze or browse is diet determined primarily by environmental con- ditions, such as seasonal and geographic vari- Feral goats are versatile generalist herbivores ation of forage. Instead, it may be more ap- capable of surviving on grasses, forbs, browse, propriate to classify goats as mixed feeding and even marine algae. Coblentz (1977) docu- opportunists (Lu 1988). As browsers, goats mented goats using almost every plant species are known to assume a bipedal stance to reach present within a study area in California. upper sections of shrubs and trees, and even However, feral goats demonstrate strong di- to climb into trees to access foliage. In the Pacific Island Invasive Species: Capra hircus, the Feral Goat · Chynoweth et al. 143 process of browsing, goats often strip bark the absence of nonnative ungulates (Merlin and girdle trees (Spatz and Mueller-Dombois and Juvik 1992). 1973). Removal of feral goats from entire islands, An important trait that enables feral goats or ungulate exclosures within islands, demon- to persist in arid island environments is their strates this transformative effect; native and remarkable ability to survive in the absence nonnative vegetation typically shows an im- of a permanent water source. Domestic goats mediate positive response to release from have a minimum water requirement of 1.0%– grazing and browsing pressure. Within fenced 1.5% body weight per day, but selective pres- units in dryland Hawaiian forests where goats sure may enable feral goats to survive in dry have been removed, native species demon- ecosystems with even less available water strate increased survival rates (Scowcroft and (Dunson 1974). Goats primarily derive pre- Hobdy 1987). On Hawai‘i Island, heavily formed water from plant foods in many sce- browsed areas demonstrate a lack of recruit- narios (Robbins 1994) but have also been ment and an older age class structure for the observed drinking salt water (Gould Burke dominant tree species, mämane (Sophora 1988). Limited water requirements have con- chrysophylla) (Scowcroft and Sakai 1983), and tributed to the success of feral goats as an in- reduced sucker growth on endemic koa (Aca- vasive mammal on numerous Pacific islands. cia koa) (Spatz and Mueller-Dombois 1973). On Guadalupe Island, Mexico, presumed ex- environmental impact and economic tinct and extirpated plant species have recov- importance ered from seed banks after goat removal (Keitt et al. 2005). Detrimental Aspects Foraging preferences of feral goats on Pa- cific islands vary greatly, depending largely on Nonnative feral goats are notorious for the composition of available plant communi- their negative impacts on island ecosystems ties. Goats are observed feeding on both na- (Coblentz 1978). Remote Pacific island plant tive and nonnative species, but native Pacific species evolved in geographic isolation from island plants are often consumed first because herbivorous mammals, losing many of the they lack defenses against herbivory and are, secondary chemical (e.g., tannins, turpenes) therefore, often more palatable. For this rea- and morphological (e.g., thorns) defense son, even low-density goat populations can mechanisms to deter herbivory (Kelsey and have negative consequences for native flora. Locken 1987, Bowen and Vuren 1997, Sheley In Hawai‘i Volcanoes National Park, D. K. and Petroff 1999). Consequently, native and Morris (unpubl. data) observed that stomach endemic plant communities are often unable contents of feral goats depended largely on to recover from persistent herbivory and density of animals present in an area. In areas trampling, resulting in their replacement by with low goat density, where native vegeta- more tolerant and resilient nonnative species tion was abundant, stomachs contained 98% (Augustine and McNaughton 1998). Intense native species. In contrast, nonnative plants browsing and grazing by goats can extirpate composed 99% of stomach contents in areas preferred species and cause the desertifica- of high goat density where native vegetation tion of entire islands. In some cases, such as was scarce. Although native species are often Santa Fe Island in the Galápagos, feral goats preferentially consumed when available, non- eliminated 100% of seedlings from large trees native plants support goat populations where (Clark and Clark 1981). Importantly, the native species do not occur. presence of nonnative ungulates can affect In addition to direct effects from browsing, competition between native and introduced grazing, and trampling, feral goats have im- plants. A comparison of Pacific islands with portant indirect effects; the alteration of plant and without introduced ungulates indicates communities through modification of plant that some island plant communities
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