COMMUNITY AND ECOSYSTEM ECOLOGY Fruit Flies of the Genus Anastrepha (Diptera: Tephritidae) and Associated Native Parasitoids (Hymenoptera) in the Tropical Rainforest Biosphere Reserve of Montes Azules, Chiapas, Mexico MARTI´N ALUJA,1, 2 JUAN RULL,1 JOHN SIVINSKI,3 ALLEN L. NORRBOM,4 ROBERT A. WHARTON,5 ROGELIO MACI´AS-ORDO´ N˜ EZ,1 FRANCISCO DI´AZ-FLEISCHER,6 1 AND MAURILIO LO´ PEZ Environ. Entomol. 32(6): 1377Ð1385 (2003) ABSTRACT We report the results of a 2-yr survey that determined some of the host plant and parasitoid associations of Anastrepha fruit ßies (Diptera: Tephritidae) in the “Montes Azules” tropical rainforest biosphere reserve (State of Chiapas, Mexico). We collected a total of 57.38 kg of fruit representing 47 native species from 23 plant families. Of these, 13 plant species from eight plant families were found to be native hosts of 9 species of Anastrepha. The following Anastrepha host associations were observed: Bellucia pentamera Naudin (Melastomataceae) with A. coronilli Carrejo y Gonza´lez; Malmea gaumeri (Greenm.) Lundell (Annonaceae) with A. bahiensis Lima; Tabernamontana alba Mill. (Apocynaceae) with A. cordata Aldrich; Quararibea yunckeri Standl. (Bombacaceae) with A. crebra Stone; Ampelocera hottlei (Standl.) Standl. (Ulmaceae) with A. obliqua (Macquart) and A. fraterculus (Wiedemann); Zuelania guidonia Britton and Millsp. and Casearia tremula (Griseb.) Griseb. ex C. Wright (Flacourtaceae) with A. zuelaniae Stone; Psidium sartorianum (O. Berg.) Nied (Myrtaceae) with A. fraterculus; Psidium guajava L. and P. sartorianum (Myrtaceae) with A. striata Schiner; and Manilkara zapota (L.) Van Royen, Pouteria sp., Bumelia sebolana Lundell, and Calocarpum mammosum (L.) Pierre (Sapotaceae) with A. serpentina (Wiedemann). The following are new host plant records: Malmea gaumeri for A. bahiensis; Quararibea yunckeri for A. crebra; Ampelocera hottlei for A. fraterculus and A. obliqua; Bumelia sebolana for A. serpentina; and Casearia tremula for A. zuelaniae. A. coronilli is reported for the Þrst time in Mexico. Infestation levels were variable and ranged between 0 and 1.63 larvae/g of fruit depending on host species. Larvae of eight species of Anastrepha on nine plant species from six plant families were found to be parasitized by Doryctobracon areolatus Szepligeti, D. crawfordi Viereck, D. zeteki Musebeck (new report for Mexico and northernmost record for the species), Opius hirtus Fisher, Utetes anastrephae Viereck (all Hymenoptera: Braconidae), and Aganaspis pelleranoi Brethes (Hymenoptera: Figitidae). Percent parasitism ranged from 0 to 76.5%. We discuss our Þndings in light of their practical (e.g., biological control) and theoretical (e.g., species radiation) implications and highlight the importance of these types of studies given the rampant deforestation of tropical forests in Latin America and the risk of extinction of rare fruit ßy species that could shed light on the evolution of host plant and parasitoid associations within the genus Anastrepha. KEY WORDS Tephritidae, Anastrepha, hostÐplant relationships, parasitoids, conservation RAPID DEFORESTATION OF THE tropics may be causing the also threatens the associated native parasitoid species extirpation, perhaps even the extinction, of many fruit (Hymenoptera). This is important because much of ßy species (Diptera: Tephritidae) (Aluja 1999) and the information required to gain a more thorough understanding of fruit ßy biology, ecology, and evo- 1 Instituto de Ecologõ´a, A. C., Apartado Postal 63, Xalapa, Veracruz lution is to be found in areas with unperturbed native 91000, Me´xico. vegetation. Therefore, one is confronted with the se- 2 E-mail: [email protected]. 3 Center for Medical, Agricultural and Veterinary Entomology, P.O. rious challenge of Þnding the few remaining pristine Box 14565, Gainesville, FL 32604. areas in which to study topics such as ancestral fruit ßy 4 SEL-USDA, U.S. National Museum NHB-168, Washington, DC hostÐplant and parasitoid associations, patterns of re- 20560. source use, factors that regulate populations and niche 5 Department of Entomology TexasA&MUniversity, College Station, TX 77843. partitioning, and adult behavior. 6 Campan˜ a Nacional Contra Moscas de la Fruta, Desarrollo de The types of information that can be gathered Me´todos, 2a. Av. Sur No. 5 Altos, 30700 Tapachula, Chiapas, Me´xico. through surveys in areas with unperturbed native veg- 0046-225X/03/1377Ð1385$04.00/0 ᭧ 2003 Entomological Society of America 1378 ENVIRONMENTAL ENTOMOLOGY Vol. 32, no. 6 etation and the potential uses to which it may be put [44%] described species, but even those species may can be illustrated by the work of Herna´ndez-Ortõ´z and have additional wild hosts). There is mounting evi- collaborators (Herna´ndez-Ortõ´z and Pe´rez-Alonso dence that one reason the host plants for many species 1993, Herna´ndez-Ortõ´z et al. 1994). Their collections are still undiscovered is that most efforts have been in a protected tropical evergreen rainforest (Estacio´n directed at surveys of commercial plants or those with de Biologõ´a Tropical “Los Tuxtlas,” Veracruz, Mexico) similar types of fruit with the expectation of Þnding discovered rare hymenopterous parasitoids such as larvae in the ripening pulp of fruit. Too little attention Microcrasis sp., Nealiolus sp. (both Braconidae) and has been paid to wild native plants, especially those Lopheucoila sp. (Cynipidae), associated with various with small fruits, and to examining immature fruits, Anastrepha species. They also determined new fruit seeds, and other plant parts (Aluja 1999). Recent sur- ßyÐparasitoid associations (e.g., Anastrepha cordata veys carried out in Mexico and Brazil have shown that AldrichÐOpius hirtus Fischer) and fruit ßyÐhost there is a large suite of native hymenopterous para- plant associations (e.g., A. bahiensis LimaÐPseudolme- sitoids whose potential as biological control agents has dia oxyphyllaria J.D. Smith [Moraceae]). Similarly, been inadequately assessed because of overemphasis Lo´pez et al. (1999) and Aluja et al. (2000a), collecting on exotic parasitoids (Ovruski et al. 2000). In the case in isolated patches of tropical deciduous and sub- of Anastrepha, 43 native parasitoid species have been deciduous forests, discovered new ßy hostÐplant identiÞed, 58.1% of which belong to the family Bra- and parasitoid associations such as those between conidae, and 18.6, 11.6, 6.9, and 2.3% to the families A. alveata Stone, Ximenia americana L. (Olacaceae), Figitidae, Diapriidae, Pteromalidae, and Eulophidae, and Doryctobracon areolatus Sze´pligeti (Braconidae); respectively (Ovruski et al. 2000). By far, the most between A. spatulata Stone, Schoepfia schreberi J.F. widespread (and perhaps abundant) native parasi- Gmel. (Olacaceae), and D. areolatus and D. toxotrypa- toids of Anastrepha, based on existing reports, are the nae Muesebeck (Braconidae); and between A. ludens braconids D. areolatus, D. crawfordi Viereck, and (Loew) and A. striata Schiner and the pupal parasitoid Utetes anastrephae Viereck, and the eucoiline Þgitid Coptera haywardi Ogloblin (Diapriidae). These Þnd- Aganaspis pelleranoi Brethes. These range from either ings can aid the “biorational” control of pestiferous southern Florida or equivalent latitudes in northern fruit ßy species by improving the conservation of Mexico through northern Argentina (Ovruski et al. biological control agents (Aluja 1999, Sivinski et al. 2000). Finally, many wild plants serve as parasitoid 2000). They also provide the samples needed for com- reservoirs and should therefore be protected or cul- parative studies aimed at understanding the evolution tivated (Aluja 1999). of Anastrepha behavior (Aluja and Norrbom 1999). In an effort to further contribute to our scant knowl- For example, thanks to the efforts of Herna´ndez- edge of Anastrepha diversity and hostÐplant and para- Ortõ´z and collaborators (Herna´ndez-Ortõ´z and Pe´rez- sitoid associations in unperturbed habitats, we report Alonso 1993, Herna´ndez-Ortõ´z et al. 1994), the habits Þndings of a 2-yr fruit collection carried out in the of primitive Anastrepha species such as A. cordata were “Montes Azules” biosphere reserve (State of Chiapas, discovered, and this has allowed hypotheses about the Mexico). Montes Azules is a 331,220 ha protected area evolution of oviposititon behavior of ßies in the genus within the Lacandon jungle (selva Lacandona) bor- Anastrepha (Aluja et al. 1999, Dõ´az-Fleischer et al. dering Guatemala and a Mesoamerican biodiversity 1999). “hotspot” (Myers et al. 2000). This tropical rain forest Most published information on Anastrepha hostÐ holds 15% of MexicoÕs plant species, 50% of MexicoÕs plant relationships and parasitoid associations has 800 butterßy species, and the greatest diversity of bats been recently summarized by Ovruski et al. (2000), in the country (Arriaga et al. 2000). Aluja et al. (2000b), Zucchi (2000), Uramoto et al. (2001), and Norrbom (2002, 2003). Several patterns Materials and Methods arise from these reviews. Most species of Anastrepha, particularly those of no economic importance and Study Area. We worked in the “Estacio´n Chajul,” even a few that are (i.e., A. grandis Macquart and located in Chajul, Chiapas (SE Mexico) and in several A. striata), are either monophagous or stenophagous sampling points along the Lacantu´ n river. Chajul is and have long adult lifespans (Aluja et al. 1999). The located between 16Ð17Њ N, between 90Њ30ЈÐ91Њ31Ј W Þve most economically important Anastrepha species (Fig. 1), and at an elevation