Indirect Effects of Ant Eradication Efforts on Offshore Islets in the Hawaiian Archipelago
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Biol Invasions (2011) 13:545–557 DOI 10.1007/s10530-010-9848-y ORIGINAL PAPER Indirect effects of ant eradication efforts on offshore islets in the Hawaiian Archipelago Sheldon Plentovich • Jakob Eijzenga • Heather Eijzenga • David Smith Received: 15 February 2010 / Accepted: 2 August 2010 / Published online: 18 August 2010 Ó Springer Science+Business Media B.V. 2010 Abstract Invasive species eradication and control geminata numbers declined, but the species remained are considered vital components of the conservation, present. Target ant densities remained high on restoration, and management of many native ecosys- untreated islets. Application of hydramethylnon tems. Invasive ants, which are notoriously difficult to reduced numbers of alien cockroaches (Order: Blat- eradicate, can cause catastrophic changes in ecosys- taria), but we did not detect effects on other non- tems and are aggressive colonists. Here we report the target arthropods. The eradication of P. megacephala eradication and control of two widely distributed was followed by dynamic compositional changes in invasive ants and subsequent unanticipated effects on the ant community, including the apparent coloniza- arthropod and avian communities. We used a paired tion by three species (S. geminata, Tetramorium experimental design that included 1 year of baseline bicarinatum and Anoplolepis gracilipes) previously data collection, to test the effects of the formicide undetected on the islet. One of these, A. gracilipes, hydramethylnon on abundances of two ant species on underwent a rapid range expansion during 2006–2008 two pairs of offshore islets. Pheidole megacephala which corresponded with reduced seabird nesting was eradicated from the treated islet in pair 1 and was success. We conclude that hydramethylnon can be not detected during 2003–2008. On pair 2 Solenopsis used effectively to eradicate P. megacephala. How- ever, ant eradications can have detrimental effects on ecosystems and the potential for subsequent coloni- S. Plentovich (&) Department of Zoology, University of Hawaii, zation of sites by other ant species that may be more Edmondson Hall 152, 2538 McCarthy Mall, harmful and more difficult to eradicate needs to be Honolulu, HI 96822-2279, USA considered. e-mail: [email protected] Ò J. Eijzenga Keywords AMDRO Á Island Á Non-target effects Á SWCA Environmental Consultants, 201 Merchant Street, Pheidole megacephala Á Seabirds Á Solenopsis Suite 2310, Honolulu, HI 96813, USA H. Eijzenga Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817, USA Introduction D. Smith Eradication of invasive species is a vital part of the Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife, 2135 Makiki Heights conservation and management of ecosystems (Myers Drive, Honolulu, HI 96822, USA et al. 2000; Ruiz and Carlton 2003). There have been 123 546 S. Plentovich et al. advances in the eradication of some groups (e.g., rats) in all of these cases there was at best only limited from increasingly large islands (Howald et al. 2007), monitoring of the effects of reduced ant densities and but for the majority of species, eradication remains the eradication process itself on non-target organisms more challenging (Clout and Veitch 2002), and for with the exception of other ant species. some species, such as invasive ants, it is notoriously Attempts to eradicate or control ants often involve difficult and requires ongoing treatment (Holway the broadcast of chemical ant baits or use of bait et al. 2002). This has global significance as hundreds stations (Reimer and Beardsley 1990; Zerhusen and of invasive ant species are undergoing human-med- Rashid 1992; Green and O’Dowd 2010). Ants are iated range expansions (McGlynn 1999; Holway known for their efficient exploitation of available et al. 2002). These invasions can alter the structure food sources (Holldolbler and Wilson 1990) and can and functioning of natural communities (Christian quickly transport bait underground before it is 2001; Holway et al. 2002; Hill et al. 2003; O’Dowd ingested by other arthropods. This preemption of et al. 2003; Krushelnycky and Gillespie 2008) and bait is thought to minimize effects to non-target can be catastrophic to native biota by reducing, arthropods and therefore non-target effects may be extirpating and possibly causing the extinction of more pronounced if baits are broadcast in areas with arthropod species (Perkins 1913; Risch and Carroll low densities of ants. However, this hypothesis has 1982; Cole et al. 1992; Gillespie and Reimer 1993; not been well studied and some evidence suggests LaPolla et al. 2000; Hill et al. 2003; O’Dowd et al. that the broadcast of formicides can negatively affect 2003), directly and indirectly harming vertebrates non-target organisms (Green and O’Dowd 2010). (Meek 2000; Holway et al. 2002; Davis et al. 2008; Limited data on the impacts of hydramethylnon, a Plentovich et al. 2009) and altering plant communi- commonly used formicide, on non-target species ties (Bach 1991; Green et al. 1997; Christian 2001; indicate no changes in populations of non-target Hill et al. 2003; O’Dowd et al. 2003; Handler et al. species in pine forests in the southeastern US (Colby 2007). Endemic biota, especially those in areas such 2002) and no effects when applied topically to Apis as the Hawaiian Islands where ants are believed to be mellifera (European honeybee) (NPIC 2002). How- absent from the native fauna, seem to experience the ever, hand broadcast of hydramethylnon (tradename: most significant losses (Perkins 1913; Zimmerman MaxForce) on 6 ha Spit Island, Midway Atoll 1970; Howarth 1985; Cole et al. 1992; Gillespie and resulted in reduced numbers of alien cockroaches Reimer 1993; LaPolla et al. 2000; Krushelnycky and (Blattaria) and alien crickets (Orthoptera: Gryllidae, Gillespie 2010). Plentovich et al. 2010). Historically ant eradication efforts met limited The Hawaiian Islands, including islets offshore, success, sometimes resulting in a temporary reduction provide opportunities to study the feasibility, effec- in ant numbers followed by a sharp rebound (Williams tiveness, and ecological cost of ant control method- et al. 2001; Tschinkel 2006). Recent successes include ology. Their isolation and small size increase chances the eradication of Wasmannia. auropunctata (little fire of successful eradications and make recolonization ant) from 3 ha on Santa Fe Island (Abedrabbo 1994) more difficult. Because ants are believed to be absent and 21 ha on Marchena Island (Causton et al. 2005) from the native fauna (Wilson 1996), there is no risk both in the Galapagos archipelago, eradication of P. for nontarget effects on native ant species. At least megacephala (big-headed ant) from treated pineapple fifty species of invasive ants are established in Hawaii plantations in Hawaii (Reimer and Beardsley 1990), (http://www.antweb.org/hawaii.jsp). Pheidole mega- and eradication of multiple infestations of Pheidole cephala and other species such as Anoplolepis gra- megacephala and Solenopsis geminata (tropical fire cilipes (yellow crazy ant) and S. geminata dominate ant) from approximately 30 and 3 ha, respectively lowland areas (Nishida 1992; Gillespie and Reimer around human settlements in Kakadu National Park, 1993; Wilson 1996; Krushelnycky et al. 2005). Here Australia (Hoffmann and O’Connor 2004). These we attempted to eradicate two of these species while successes indicate that eradication is possible for carefully monitoring the non-target effects of the certain species and circumstances given early detec- formicide application, as well as the non-target tion of small, well-delineated infestations followed by effects of reduced ant densities following treatment. rapid response and intensive management. However, Identification and quantification of non-target effects 123 Indirect effects of ant eradication 547 will help managers determine ecological costs of steep, rocky terrain, soil disturbance by burrowing control efforts and assess whether such efforts are seabirds and approximately 5 years of drought warranted. Our specific objectives were to (1) eradi- (ending in 2004) left the Mokulua sparsely vegetated. cate two dominant lowland ant species, big-headed The most abundant plant species on the islets were ant (P. megacephala) and tropical fire ant the alien grasses Cenchrus echinatus and Chloris (S. geminata) from small (\5 ha) islets offshore of barbata, along with the following native species; Oahu, Hawaii, (2) assess the effects of the formicide Jacquemontia ovalifolia subsp. sandwicensis, Sida AMDROÒ (active ingredient hydramethylnon) on fallax, Scaevola taccada, Sesuvium portulacastrum, non-target arthropods, and (3) quantify subsequent and Chenopodium oahuense. The arthropod commu- potential indirect effects of reduced target ant num- nity on both islets was dominated by two species: the bers on other ant species and on number of nesting tropical fire ant (S. geminata) and the crazy ant seabirds. (Paratrechina longicornis). Solenopsis geminata and P. megacephala were targeted in this study due to anecdotal observations of these two species attacking Methods seabirds in the Hawaiian archipelago (A. Hebshi and C. Vanderlip pers obs.) Study sites Ant control Based on the results of preliminary surveys we selected two pairs of offshore islets (Pair 1: Popoia/Mokuauia We attempted to eradicate targeted ant species on one and Pair 2: Moku Iki/Moku Nui, Table 1, Fig. 1). Each randomly chosen islet in each pair (i.e., Mokuauia pair was similar in location,