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Influence of Host-Plant Quality on the Performance of Episimus

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Influence of Host-Plant Quality on the Performance of Episimus BioControl (2009) 54:475–484 DOI 10.1007/s10526-008-9196-3 Influence of host-plant quality on the performance of Episimus unguiculus, a candidate biological control agent of Brazilian peppertree in Florida Veronica Manrique Æ J. P. Cuda Æ W. A. Overholt Æ S. M. L. Ewe Received: 18 June 2008 / Accepted: 30 October 2008 / Published online: 15 November 2008 Ó International Organization for Biological Control (IOBC) 2008 Abstract Brazilian peppertree, Schinus terebinthifo- addition, higher survival (40%), faster development lius Raddi (Sapindales: Anacardiaceae), introduced (34 day) and higher fertility (88% eggs hatched) from South America, invades a variety of habitats in occurred in high-nutrient treatments. Based on these Florida (e.g. disturbed sites, coastal mangrove forests). results, field releases should be conducted in favorable The objective of this study was to evaluate the effect of habitats (e.g., low salinity, high fertility soils) to host-plant quality on the performance of Episimus maximize the possibility of establishment and popu- unguiculus Clarke (=E. utilis Zimmerman) (Lepidop- lation growth of E. unguiculus in Florida. tera: Tortricidae), a potential biocontrol agent of Brazilian peppertree. Experiments were conducted in Keywords Insect–plant interactions Á the laboratory using Brazilian peppertrees exposed Weed biological control Á Tortricidae Á either to different salinity levels (0, 6, 12 parts per Anacardiaceae Á Schinus terebinthifolius thousand), or to different nutrient levels (low, medium, high). Higher survival (55%) and faster development (32 day) to adulthood was observed on plants grown in fresh-water environments (0 ppt) compared to low (6 ppt) or high-salinity environments (12 ppt). In Introduction Brazilian peppertree, Schinus terebinthifolius Raddi Handling Editor: John Scott. (Sapindales: Anacardiaceae), is an introduced peren- nial plant that has become widely established & V. Manrique ( ) Á W. A. Overholt throughout central and south Florida (Cuda et al. Biological Control Research and Containment Laboratory, University of Florida, 2199 South Rock Road, 2006). This species is native to Argentina, Brazil and Fort Pierce, FL 34945, USA Paraguay (Barkley 1944, 1957), and was brought to e-mail: vero72@ufl.edu Florida as an ornamental in the 1840s (Mack 1991). Brazilian peppertree not only colonizes disturbed J. P. Cuda Department of Entomology and Nematology, University sites such as highway right-of-way, canals, fallow of Florida, Bldg. 970 Natural Area Drive, Gainesville, farmlands, and drained wetlands, but also invades FL 32611, USA natural communities, including pinelands, hardwood hammocks and mangrove forests in Florida (Myrtin- S. M. L. Ewe Ecology and Environment, Inc., 1665 Palm Beach Lakes ger and Williamson 1986; Ewe 2001; Cuda et al. Blvd., West Palm Beach, FL 33401, USA 2006). The invasion and displacement of native 123 476 V. Manrique et al. species by Brazilian peppertree poses a serious threat leaflets together to feed, whereas older instars feed to biodiversity in many ecosystems of Florida and pupate inside a cylindrical rolled leaflet. The (Morton 1978). Several attributes of this plant dimorphic adults are small, grayish brown with distinct contribute to its invasiveness, including a large wing patterns that separate the sexes (Zimmerman number of fruits produced per female plant, an 1978). Because the entire life cycle of E. unguiculus effective mechanism of dispersal by birds (Panetta occurs in the canopy of the host plant, this insect may be and McKee 1997), tolerance to shade (Ewel 1979), well adapted to areas subjected to seasonal flooding in fire (Doren et al. 1991), and drought (Nilson and south Florida. Host-specificity studies conducted in Muller 1980b), allelopathic effects on neighboring Brazil and Florida quarantine showed that larval plants (Gogue et al. 1974; Nilson and Muller 1980a; feeding damage was restricted to plants in the family Morgan and Overholt 2006), and tolerance to saline Anacardiaceae in no-choice tests, and Brazilian conditions (Ewe 2001; Ewe and Sternberg 2002). peppertree was the preferred host in choice tests Although herbicides and mechanical methods (J. P. Cuda, unpublished data). A petition for the (e.g., cutting, burning and flooding) are routinely release E. unguiculus in Florida for biological control of used for controlling existing Brazilian peppertree Brazilian peppertree is currently in preparation. stands (Gioeli and Langeland 2006; Cuda et al. 2006), Several factors influence the establishment of these methods are extremely labor intensive and can biological control agents, such as climate, natural be very expensive, especially for large infestations. In enemies present in the introduced range and host-plant addition, chemical and mechanical controls are quality (Sutherst and Maywald 1985; Wheeler and unsuitable for some natural areas (e.g., mangrove Center 1996; Hunter 2001; Wheeler 2001; Byrne et al. forests) because they may have negative effects on 2003; Senaratne et al. 2006). Furthermore, the nutri- non-target species (Doren and Jones 1997) and tional status of many weeds has played an important may increase water pollution. Therefore, alternative role in successful biological control programs (Myers methods for long-term control of Brazilian peppertree 1987; Julien and Griffiths 1998). For example, initial are urgently needed. A classical biological control applications of nitrogen fertilizer helped in the program was initiated in the 1980s, and the leaflet- establishment of Cactoblastis cactorum (Berg) roller moth Episimus unguiculus Clarke (Lepidoptera: (Lepidoptera: Pyralidae) on prickly pear (Opuntia Tortricidae) was selected as a potential biocontrol spp.) cactus (Dodd 1940)andtheCyrtobagous salvinia agent of Brazilian peppertree because it clearly Calder and Sands (Coleoptera: Curculionidae) on damaged the plant and appeared to be host specific Salvinia molesta Mitchell (Salviniaceae) (Thomas and in its native range (Martin et al. 2004;Cudaetal. Room 1986). In addition, plant palatability varies 2006). Previous studies on this insect were published across salinity gradients (Levine et al. 1998;Moonand under the junior synonym Episimus utilis Zimmerman Stiling 2000;GoransonandHo2004), and high salinity (Razowski and Brown 2008). In the 1950s, E. ungui- levels may be detrimental to some insect herbivores culus was released and established in Hawaii, but (Hemminga and van Soelen 1988; Moon and Stiling successful control of Brazilian peppertree populations 2002; Schile and Mopper 2006). was not achieved (Goeden 1977; Yoshioka and Markin Because Brazilian peppertree invades different 1991; Julien and Griffiths 1998). However, its inability habitats in Florida (e.g., mangrove forests, pinelands, to control the plant in Hawaii should not preclude its abandoned farms), the environmental conditions release into other areas infested with Brazilian pepper- encountered in different habitats may influence the tree. Unfavorable abiotic or biotic factors may have survival and effectiveness of potential biocontrol contributed to its failure in Hawaii. For example, high agents. Therefore, the objective of this study was to larval mortality of E. unguiculus by introduced and evaluate the performance of E. unguiculus reared on native parasitoids and predators of agricultural pests Brazilian peppertree exposed to different salinity and was documented following its release (Davis 1959; nutrient levels in the laboratory. Results from this Krauss 1963). study will provide a better understanding of the The larvae of E. unguiculus feed on Brazilian interaction between Brazilian peppertree and peppertree leaflets and can completely defoliate small E. unguiculus, one of its principal herbivores. This plants (Martin et al. 2004). Early instars tie young information will improve the selection of release sites 123 Influence of host-plant quality on Episimus unguiculus performance 477 favorable for establishing the insect in Florida (ppt)), (2) low saline environment (6 ppt salinity), following its release from quarantine. and (3) high saline environment (12 ppt salinity). Brazilian peppertree plants were either irrigated with tap water (fresh water environment) or with water Materials and methods supplemented with seawater (36 ppt) to obtain 6 or 12 ppt salinity. In order to prevent plant stress, a Insect rearing stepwise increase in salt concentration (1 ppt every three days) was used. Plants were irrigated with each Laboratory experiments were conducted at the Bio- salinity level for the duration of the experiment logical Control Research and Containment Laboratory (*two months). Brazilian peppertree invades (BCRCL) located at the Indian River Research different habitats in Florida, including transitional and Education Center (IRREC) of the University of mangrove forests where salinity levels range between Florida, Fort Pierce, FL, USA. A colony of 0 and 25 ppt depending on the time of the year (rainy E. unguiculus was initiated in August of 2006 at the vs. dry season) (Ewe 2001). Thus, the salinity levels BCRCL, and insects were reared on individual used in this study (6 and 12 ppt) are within the range Brazilian peppertree plants (Florida genotypes) grown of values encountered in the field. in nursery pots (18 cm height, 17 cm diameter) inside Several plant parameters were recorded at the environmental growth chambers (25 ± 2°C, 60–70% beginning of the experiment (four to six replicates RH, 14L: 10D photoperiod) (see Martin et al. (2004) per treatment):
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