Anastrepha Ludens and Anastrepha Serpentina (Diptera: Tephritidae
Total Page：16
File Type：pdf, Size：1020Kb
ECOLOGY AND BEHAVIOR Anastrepha ludens and Anastrepha serpentina (Diptera: Tephritidae) Do Not Infest Psidium guajava (Myrtaceae), but Anastrepha obliqua Occasionally Shares This Resource With Anastrepha striata in Nature 1,2,3 1 ANDREA BIRKE AND MARTI´N ALUJA J. Econ. Entomol. 104(4): 1204Ð1211 (2011); DOI: 10.1603/EC11042 ABSTRACT This study examined whether economically important fruit ßy species Anastrepha ludens (Loew), Anastrepha serpentina (Wiedemann), and Anastrepha obliqua (Macquart) (Diptera: Tephritidae) may opportunistically exploit guavas, Psidium guajava L. (Myrtaceae), growing near preferred natural hosts. We collected 3,459 kg of guavas and 895 kg of other known host species [sour orange, Citrus aurantium L.; grapefruit, Citrus paradisi Macfadyen; mango, Mangifera indica L.; white sapote, Casimiroa edulis La Llave and Lex.; sapote, Pouteria sapota (Jacq.); sapodilla, Manilkara zapota L.; and wild plum, Spondias purpurea L. and Spondias mombin L.] along an altitudinal gradient over a 4-yr period (2006Ð2009). Plants were growing in sympatry in 23 localities where the guavas are usually infested in the state of Veracruz, Me´xico. The guava samples yielded 20,341 Anastrepha spp. pupae in total (overall mean, 5.88 pupae per kg of fruit). ConÞrming previous reports, Anastrepha fraterculus (Wiedemann) and Anastrepha striata (Schiner) were found heavily infesting guavas in Veracruz. Importantly, although we did not Þnd evidence that A. ludens and A. serpentina are able to attack this valuable commodity, we document for the Þrst time in the agriculturally important state of Veracruz that P. guajava is an alternative natural host plant of A. obliqua. We recovered two fruit in the mango-growing locality of la Võ´bora, Tlalixcoyan, that harbored larvae of A. striata and A. obliqua. This Þnding has important practical implications for management of A. obliqua. Over the entire altitudinal gradient, when individual fruit infestation was examined, a dynamic pattern of species dominance was unveiled with guavas growing below 800 m above sea level mainly attacked by A. striata and a progressive replacement with increasing altitude by A. fraterculus. Interestingly, most individual fruit examined (97%) harbored a single species of fruit ßy, a Þnding that may be taken as evidence of competitive displacement among sympatric species of fruit ßies. Based on this study and previously published work by us on this topic, we conclude that literature reports indicating that A. ludens and A. serpentina infest guavas under Þeld conditions should be questioned. KEY WORDS Anastrepha, Psidium guajava, competitive displacement, infestation patterns, host status The correct status of a fruit as a host of pestiferous fruit ludens (Loew); Anastrepha serpentina (Wiedemann); ßy species is critical in assessing the risk of introduc- and Anastrepha obliqua (Maquart) as species of con- tions into importing countries (Aluja and Mangan cern in guava export protocols from Me´xico to the 2008). Guava, Psidium guajava L. (Myrtaceae), is a United States (http://www.senasica.gob.mx). tropical fruit of commercial value with a long history Guavas in Veracruz are widely distributed and can of local consumption in Me´xico and a signiÞcant ex- be found all the way from sea level to localities at port potential. During the past 4 yr, exports to the altitudes of Ϸ1,600 m above sea level (Sivinski et al. United States have increased by 45% with expected 2000). There are no commercial plantations in Vera- future yearly returns ϾUS$350Ð400 million (http:// cruz, mainly because of heavy damage inßicted by A. sagarpa.gob.mx). In Me´xico, guavas are mainly at- fraterculus and A. striata. Trees grow along rural roads tacked by Anastrepha striata (Schiner) and Anastrepha and highways, in coffee (Coffea spp.) plantations, dis- fraterculus (Wiedemann) (Aluja et al. 1987, 2000, turbed patches of native vegetation, parks, home gar- 2003b,c; Sivinski et al. 2000, 2004); however, quaran- dens, backyards, or abandoned lots in urban areas tine restrictions include Mexican fruit ßy, Anastrepha often interspersed among natural hosts of economi- cally important fruit ßy species such as A. ludens, A. 1 Instituto de Ecologõ´a, A.C. Apartado Postal 63, 91000 Xalapa, serpentina, and A. obliqua. In the lowlands, ßowering Veracruz, Me´xico. begins in March and mature fruit is available between 2 Posgrado Instituto de Neuroetologõ´a, 91190 Xalapa, Veracruz, Me´xico. June and August. The latter time partially coincides 3 Corresponding author, e-mail: [email protected]. with the fruiting period of Spondias purpurea L., a 0022-0493/11/1204Ð1211$04.00/0 ᭧ 2011 Entomological Society of America August 2011 BIRKE AND ALUJA:FRUIT FLIES ASSOCIATED WITH GUAVAS IN CENTRAL ME´ XICO 1205 preferred host plant of A. obliqua, and with mango, hottlei (Standl.) Standl.(Ulmaceae) (Aluja et al. 1987; Mangifera indica L. ÔManilaÕ, an exotic host plant of Aluja et al. 2000, 2003c). both A. obliqua and A. ludens (Aluja and Birke 1993). There are at least 25 primary reports (sensu Aluja From Ϸ900 to 1,000 m above sea level, ßowering be- and Mangan 2008) indicating that A. obliqua is able to gins between late April and early May, and mature infest guava under Þeld conditions (information sum- fruit can be found between mid-August to late Sep- marized by Norrbom, 2004). Most credible reports tember, even though ripening of some fruit can be stem from the Caribbean Islands (Dominican Repub- delayed until December. During this time, another A. lic and Puerto Rico; Jenkins and Goenaga 2008a) and obliqua preferred host plant, Spondias mombin L., is some countries in Central and South America (Costa ripening (Lo´pez et al. 1999, Aluja et al. 2000). Also, Rica, Colombia, Venezuela, Brazil, and Ecuador; (Ma- various species of Citrus and Sapotaceae, preferred lavasi and Morgante 1980, Malavasi et al. 1980, Cara- hosts of A. ludens and A. serpentina, respectively, are ballo 1981, Jiro´n and Hedstro¨m 1988, Ohashi et al. abundant. Above 1,000 m above sea level, ßowering 1997, Katiyar et al. 2000, Zucchi 2000, Uchoˆa-Ferna´n- begins in June, and mature fruit becomes available dez et al. 2002, Araujo et al. 2005, Raga et al. 2006, between mid-August and early October. Despite this Jenkins and Goenaga 2008b, Minza˜o and Uchoˆa- altitudinal seasonality, one can Þnd off-season fruit Ferna´ndez 2008). In Me´xico, P. guajava has only been year-round in single trees, albeit in very small num- recorded as a host of A. obliqua in the southern-most bers. Soconusco region in the state of Chiapas (Aluja et al. A. ludens is one of the most economically important 1987), despite extensive fruit collecting efforts in sev- fruit pests in Me´xico. It is a polyphagous fruit ßy eral areas throughout the country (e.g., Yucata´n, species, whose geographical range spans from Central Herna´ndez-Ortõ´z et al. 2006; Veracruz, Herna´ndez- America to the southern United States (Thomas 2003, Ortõ´z and Pe´rez-Alonso 1993, Lo´pez et al. 1999, Aluja Aluja et al. 2009). Its main natural hosts are in the et al. 2000, Sivinski et al. 2000); northern Chiapas, Rutaceae and include yellow chapote, Casimiroa greg- Montes Azules, Aluja et al. 2003c; and Nayarit, Pe´rez- gii (S. Watson); white sapote, Casimiroa edulis La Staples and Aluja 2004). Llave & Lex.; and the exotic hosts bitter orange, Citrus Our null hypothesis was that in our study region aurantium L.; grapefruit, Citrus paradisi (Macfadyen); guavas would only be naturally infested by A. striata and orange, Citrus sinensis (L.) Osbeck. Other hosts and A. fraterculus, two fruit ßy species with an un- include mango, Mangifera indica L. (Anacardiaceae); equivocally proven association to this host in Me´xico. peach, Prunus persica L. (Rosaceae); and ÔManzanoÕ Alternatively, we hypothesized that A. ludens, A. ser- pepper, Capsicum pubescens L. (Solanaceae) (Aluja et pentina, and A. obliqua may opportunistically exploit al. 2000, Norrbom 2004, Thomas 2004). Guava has guavas when growing in the vicinity(within ßight been reported as a host of A. ludens (Herna´ndez-Ortiz range) of their preferred natural hosts. Hosts were and Aluja 1993, White and Elson Harris 1992), but considered to be in sympatry when found within 250 m such reports are not based on Þeld evidence but rather of each other, which represents the mean daily ßight on questionable literature. distance for Anastrepha adults (Thomas and Loera- A. serpentina is distributed from the southern Gallardo 1998). To test these hypotheses, we con- United States to Brazil and has been reported to ex- ducted a 4-yr study in 23 different Þeld sites along an ploit 22 plant species in seven families, with a marked elevation gradient in the agriculturally important state preference for fruit in the Sapotaceae family of Veracruz, Me´xico. The study was based on fruit (Robacker et al. 2009). EskaÞ and Cunningham (1987) collections in the Þeld and follow-up quantiÞcation of reported P. guajava as a host in Guatemala, but results infestation patterns. of this survey are questionable because they were reported along with erroneous recoveries of A. obliqua Materials and Methods and A. striata from Citrus sinensis (undoubtedly a mistake). Fruit Collection Sites. The study was carried out A. obliqua is distributed from Me´xico to southern during the 2006Ð2007 and 2008Ð2009 fruiting seasons. Brazil and the Caribbean Islands (Aluja 1993, Herna´n- Guavas were systematically collected along an altitu- dez-Ortiz and Aluja 1993). It does not occur in the dinal gradient (9Ð1,137 m above sea level) in central United States and is therefore considered a pest of Veracruz, Me´xico. Nineteen localities were selected quarantine signiÞcance for this country (Steck 2001). among those with the highest fruit ßy infestation rates This species predominantly attacks fruit within the and greatest guava tree abundance (see Tables 1 and Anacardiaceae family; it is considered a major pest of 2 for details; Lo´pez et al.