Lime Swallowtails in a New World
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Wayne Wehling (2) Wehling Wayne Lime Swallowtails by Wayne Wehling, Carmelo A. in a New World Núñez, and Jeffrey Glassberg Plant and animal species evolve in a particular sites throughout a large area of the Dominican region and then may naturally spread from Republic9. their place of origin to occupy a much larger In the Dominican Republic lime swallow- range. Alternatively, a species may be native tails, as would be expected, are using various to one area and be transported by humans citrus trees, especially limes and oranges, as to another part of the planet, where it then caterpillar foodplants7. However, one of us flourishes. This is the story of an animal (Núñez) has observed that the most attractive — the lime swallowtail (Papilio demoleus) caterpillar foodplant appears to be limeberry — that has now taken both routes to a global (Triphasia trifolia) a widely planted ornamen- distribution. tal, non-citrus Rutaceae that is native to Asia Lime swallowtails, part of a closely- and treated as an invasive in Florida, Puerto related group of species thought to have Rico and Texas. originated in Madagascar1, are native to Asia, In March 2005, a 19-year-old woman, ranging from Iran in the West, across India known to us only by her web alias — and southern China, to Japan and Australia phetamined — visited Rincon in far western in the East. This species has been extremely Puerto Rico. While there she photographed a successful in Asia, greatly expanding its range swallowtail on the beach and later uploaded within historic times. For example, it has been her photograph to the website Webshots. An suggested that lime swallowtails were intro- examination of the photo (page 32) clearly duced into Iran in the 10th century along with shows it to be of a lime swallowtail — the first citrus2. More recently, these strong-flying record of this species from Puerto Rico. swallowtails jumped from mainland Asia to In June 2006, lime swallowtails were many offshore islands3, 4. Throughout most found in Guanica, Puerto Rico by members of their range, lime swallowtails are pests of of the USDA’s Animal and Plant Health citrus groves5 and have been described as “the Inspection Service10. When one of the authors most destructive pests in [citrus] nurseries.”6 of this article (Glassberg) visited the Guanica Top: A lime swallowtail On March 29, 2004, a group from Harvard area of Puerto Rico on November, 17-18, in a makeshift University collected two lime swallowtails in 2006, he found lime swallowtails to be the laboratory in the the southeast portion of the Dominican Repub- most common swallowtail in an area of about Dominican Republic. lic for the first confirmed occurrence of this 25 square miles that included the Guanica Dry September, 2005. species in the New World7. DNA analysis has Forest preserve. There were no citrus groves shown that the Dominican Republic butterflies in this area, although the existence of a small are derived from the Malayan subspecies of number of citrus trees on residential properties Right: The upperside 8 lime swallowtail . could not be excluded. Most of the observed of lime swallowtails is Because of the economic implications of butterflies were nectaring at bougainvillea. quite unlike any native this finding, an international team of ento- In addition, he saw a single individual of a mologists visited the Dominican Republic in butterfly that was probably this species in the New World swallowtail. September 2004 to survey for this butterfly. Maricao Forest, in the western Puerto Rico September, 2005. The team, headed by one of the authors of highlands. Dominican Republic. this article (Wehling) found the swallowtail Although we know of no reports sub- to be common — more common than native stantiating this, it is possible that lime swal- citrus-feeding swallowtails — at numerous lowtail caterpillars are using native prickly- 30 American Butterflies,Summer/Fall 2006 31 Wayne Wehling (2) Wehling Wayne Left: The first known Right top: Fourth sighting of a lime instar lime swallowtail swallowtail on Puerto caterpillars. Sept. 2004. Rico. March 2005, Dominican Republic. Rincon, Puerto Rico. Right bottom: Second Below: Habitat for instar lime swallowtail lime swallowtails in the caterpillar. Sept. 2004. Guanica area of Puerto Dominican Republic. Rico. Note the absence of any citrus groves. Nov. 17, 2006. Given the foregoing, it seems likely that Phetamined lime swallowtails are already well-established on Cuba, and inevitable that they will soon be seen in southern Florida where the caterpillars will likely use cultivated citrus, limeberries and native citrus-family plants such as prickly- ashes. We predict that lime swallowtails, using a similar suite of caterpillar foodplants, will eventually spread to roughly the range of Giant Swallowtails, that is from southern South America to New Jersey and the upper Midwestern United States (lime swallowtails range up to 7000 ft. in the Himalayas14, 15). How did the first lime swallowtails arrive in the Dominican Republic from southeastern Asia? They could have arrived in a shipment of citrus, but this seems very unlikely given the biology of the butterfly. Adults could not survive the long journey. Chrysalids are not normally found associated with citrus fruit and The unfortunate introduction of a pest but- establishing a new population would probably terfly into the New World illustrates the impor- require a number of unrelated individuals to tance of responsible choices in how lepidop- emerge at a similar time, mate, and produce tera livestock are handled. We will probably viable offspring. It seems much more likely never know with certainty how this butterfly Jeffrey Glassberg Jeffrey that the butterflies were intentionally imported made it to the Dominican Republic, however, ash (Zanthoxylum) species, such as West December 2006, a lime swallowtail was found into the Dominican Republic, either by a hob- we can be almost certain that negative public Indian prickly-ash (Z. flavum)(also found in in Jamaica13. byist or by an private butterfly zoo at one of relations will come. Although we doubt that the Florida Keys) and white prickly-ash (Z. Here are some approximate distances the upscale tourist resorts in the southeastern lime swallowtails will be a very serious eco- martinicense) as caterpillar foodplants since a between various West Indian islands and Dominican Republic near where the swallow- nomic pest in the United States, one concern number of Zanthoxylum species are reported Florida. Hispaniola-Jamaica,110 miles; tails were first seen. Lime swallowtails are we do have is that when lime swallowtails as hostplants in the literature11. Hispaniola-Puerto Rico, 60 miles; Hispaniola- commonly exhibited at butterfly zoos through- are found in Florida, the control/eradication In June, 2006, Dr. Jacqueline Miller Cuba, 50 miles; Cuba-Florida Keys, 90 miles; out the world. At least 19 regulated butterfly efforts may seriously harm other butterflies, spotted a butterfly on North Andros, Bahamas, North Andros, Bahamas-Florida Keys, 125 zoos in the United States have permits to especially other rutaceae-feeding swallowtails that may well have been this species12. In miles. import this species. such as Giant and Schaus’ Swallowtails. 32 American Butterflies,Summer/Fall 2006 33 1. Zakharov, E., et al. 2004. Independent gene phy- 11. http://www.nhm.ac.uk/research-curation/proj- logenies and morphology demonstrate a Malagasay ects/hostplants/#6 www.sunstreaktours.com Visit our new websites [sic] origin for a wide-ranging group of swallowtail 12. Jacqueline Miller, personal communication. www.sunstreaktours.com butterflies. Evolution 58: 2763-2782. 13. The Jamaica Observer internet edition, Dec. 20, www.sunstreakbooks.com 2. Larsen, T.B. 1977. Extension recente en Iraqu de 2006. l’aire de Papilio demoleus. Linn. Entomops 14. Winter-Blyth, M.A. 1957. Butterflies of the 42: 37-38. Indian Region. Bombay Natural History Society: New! Field Guide to 3. Matsumoto, K. 1996. Establishment of Papilio Bombay. Mexican Butterflies demoleus in Java. J. Lep. Soc. 50: 139-140. 15. Smith, Colin. 1989. Butterflies of Nepal. Tec- 4. Matsumoto, K. 2002. Papilio demoleus in press: Bangkok, Thailand. Seek Borneo and Bali. J. Lep. Soc. 56: 108-111. 5. Larsen, T.B. 1984. Butterflies of Saudi Arabia and its Neighbors. Stacey International: London. Below: Wasps are efficient predators of Vacation 6. Agribusiness Information Center website: www. lime swallowtail caterpillars. ficciagroindia.com/production-guidelines/fruits/ 2006. Dominican Republic. Elation citrus/Pests.htm Pedro G. Rodgriguez 7. Guerrero, K.A., et al. 2004. First New World New! Wyoming trip. documentation of an Old World citrus pest, the Butterfly Tours with lime swallowtail, Papilio demoleus, in the Domini- Small Groups, can Republic (Hispaniola). Am. Entomol. 50: Great Leaders Tours throughout the U.S., 227-229. Mexico and the Carribbean. 8. Eastwood, R., Boyce, S.L. and Farrell, B.D. Sensational Butterflies 2006. The provenance of Old World swallowtail Shown are views of our butterflies, Papilio demoleus, recently discovered www.sunstreaktours.com Wyoming trip. in the New World. Annals of the Entomol. Soc. of America. 99: 164-168. Call or write Sunstreak Tours for complete info: 973-285-0890, 4 Delaware Rd., Morristown, NJ 07960. 9. Wehling, W. 2004. Dominican lime swallowtail survey report to USDA. USDA Internal Report. 10. Dunkle, R.L. 2006. Lime Swallowtail (Papilio NABA Checklist & English Names demoleus) detected in the Municipality of Guanica, Puerto Rico. National Plant Board. DA-2006-55. of North American Butterflies Invaluable to everyone interested in butterflies! All 722 species of butterflies recorded in North America north of Mexico (and in Hawaii) are listed, giving both the English and scientific names. In front of each species’ name is a line for a checkmark. Following each name a space is provided to allow you to write in date and locality information. Also included is an introductory section, explain- ing the formation, deliberations, and policies of the NABA Names Committee and a commen- tary section, in which the reasons why particular names were chosen are explained and the votes of the Committee are given.