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Florida Golf Courses Helpan Endangered ButterflyGolf courses in the Florida Keys offer their helpto save a colorful and rare butterfly.BY JARET C. DANIELS AND THOMAS C. EMMEL

fever. He was an amateurnaturalist who in his spare timediscovered and collectedspecimens of a newswallowtail.

When discovered, theSchaus swallowtail inhabitedtropical hardwood hammockson the south Florida mainland.This globally endangeredhabitat type, one of the mostimperiled plant communitiesin Florida, also occurs through-out the Florida Keys and iscomposed of slightly elevatedlimestone areas that supportbroad-leaved tropical decidu-ous trees. Due to poor nutrient

availability, sparse soils, lack of freshwater, and harsh growing conditions,the dense hammock canopy remainsdiminutive, rarely reaching over 40 feethigh. High, stable ground in southernFlorida was a highly sought-aftercommodity, and soon it was rapidlydwindling due to expanding urbandevelopment surrounding Miami.

Also during this period, a collectortook a specimen of the swallowtail onKey West in 1885. It is not surprisingthat the Schaus swallowtail was firstrecorded in the Lower Keys, then some200 miles from the south Florida main-land, as the only viable means of travelto the Keys was via boat, and Key Westwas a major port. In fact, Key West wasthe largest city in Florida in 1890,exceeding even Miami in population. Itwas not until some years later in 1912

ponceanus) is a large, colorful butterflyendemic to southern Florida; additionalsubspecies occur in the West Indies.12 Itis considered one of the rarest residentbutterflies in North America and islisted as an endangered species by boththe state of Florida and the federalgovernment. It is one of seven swallow-tail butterflies out of about 573 knownspecies that are listed by the Inter-national Union for Conservation ofNature and Natural Resources.1

Historically, the species was oncewidespread from the greater Miami areasouth to Lower Matecumbe Key in theFlorida Keys. It was formally named asa species, Papilio ponceanus, in 1911 byWilliam Schaus, a physician stationed inMiami during the Spanish-AmericanWar to help treat American soldiersevacuated from Cuba because of yellow

Native larval host plants and adult nectar sources were planted on golfcourses to create natural habitat suitable to maintain transient adultbutterflies, encourage adult movement and gene flow between existingcolonies, and allow for the natural establishment of new breeding colonieswithin the Florida Keys.

THE CASE OFTHESCHAUS SWALLOWTAILBy far one of the most successful andhighly publicized projects has centeredon the only endangered swallowtailbutterfly in the United States. TheSchaus swallowtail (Papilio aristodemus

T'.'"he plight of imperiledbutterfly species aroundthe world continues to

generate increased publicinterest and funding support.Recent management planscreated to conserve manycritically endangered butterfliesmirror the aggressive, creative,and cooperative nature ofthose types of plans historicallyimplemented for traditionalvertebrate conservationprograms. Laboratory andfield-based ecological researchcombined with captivepropagation, organism reintro-duction and translocation,habitat restoration or augmentation, andmovement corridor development havehelped unite university researchers,governmental agencies, non-govern-mental conservation organizations, andprivate landowners in a conunon goalof biodiversity conservation. Althoughnot yet fully realized, the outcome ofsuch ambitious recovery measures hashelped bring invertebrate conservationto the forefront and led to cautious, butoptimistic forecasts for the future of thespecies involved.

22 G R E ENS E C T ION R E COR D

Researchers at the University of Florida initiated efforts to improve and expand suitable breedinghabitat for the endangered Schaus swallowtail butterfly. Native larval host plants and adult nectarsources are being planted on golf courses in southern Florida to assist that effort.

that Henry Flagler built a new railroadthrough the Keys, linking the mainlandto Key West and opening up thenumerous small islands to tourism andcolonization. By then the remainingtropical hardwood hammock habitat onKey West had already been cleared forhousing and commercial development,and the butterfly was extirpated therewhile inhabiting the less-settled Keys tothe north.

DEVELOPMENTANDANDREWTHREATEN SURVIVALIn the ensuing decade, better roads,mosquito control, widespread electricity(making air conditioning possible), andfresh water piped in from the Floridamainland brought rapid development tomuch of the Keys. As a result, the over-all trends in the Schaus swallowtail'srange and numerical abundance con-tinued to decline. It was last recordedon the Florida mainland on May 31,1924,11and during the 1940s to 1970swas reduced in range to Key Largo andnumerous small islands to the northwithin Biscayne National Park. Up to1972, naturalists could come to KeyLargo and regularly see several hundredswallowtails along the sun-dappledhammock trails on the island's northernend.

But following the 1972 flight season,the Schaus swallowtail population onKey Largo underwent a rapid anddramatic decline. In 1977, it was listed asa threatened species by the U.S. Fishand Wildlife Service (USFWS), andseveral quick studies were completed todetermine the status of the butterfly,culminating in a recovery plan writtenby the Florida Game and FreshwaterFish Commission and published in1982.

In May 1984, at the direct request ofthe USFWS office in Jacksonville, aUniversity of Florida research team wasassembled to carry out status surveys insouth Florida and make recommenda-tions for action on the existing recoveryplan. The resulting data indicated that

the observed decline in the swallowtail'shistoric range and numerical abundancefrom 1924 to 1981 had continued, withless than 70 adults recorded during the1984 flight season. The only threecolonies of any significant size werelocated, all within Biscayne NationalPark on Elliott Key, Old Rhodes Key,and Totten Key, with a fourth smallcolony in the remaining intact ham-mock on northern Key Largo beingrepresented by only a single adult.Following this report in 1984, theSchaus swallowtail was upgraded in

listing from threatened to endangeredstatus.

Field and laboratory research indi-cated that the two principal factorscontributing to the demise of theSchaus swallowtail throughout much ofits former range were habitat loss andmosquito control adulticide spraying.The final blow, starting in 1973, hadbeen the initiation that year of the useof two new organophosphate adulti-cides, Dibrom and Batex, in the Keysby the Monroe County MosquitoControl District. The resulting sprayinghad far-reaching effects on the butterfly

and contributed to it being reduced toan extremely limited geographic range.

Despite these setbacks, ensuing re-strictions on the use of these chemicalsresulted in the slow increase in popula-tion, and the Schaus remained relativelysecure from human-promoted influ-ences in the small but protected ham-mocks of Biscayne National Park. TheNongame Wildlife Section of theFlorida Game and Freshwater FishCommission subsequently funded threesuccessive two-year grants (1985-86,1987-88, 1991-92), along with addi-

tional assistance from the dePont Fundduring 1988-90, in order to continuethe status surveys of this clearlyendangered butterfly.

The additional threat to the speciesresulting from the impact of a majornatural disaster was realized on August24, 1992, when hurricane Andrewslammed into southern Florida, destroy-ing or heavily damaging all habitat areasfostering remaining butterfly popula-tions. Field surveys conducted duringthe following flight season in 1993revealed that Schaus swallowtail popu-lations in Biscayne National Park and

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northern Key Largo were extremelyreduced (17 adults on Elliott Key, 33 onAdams Key, and 7 on Key Largo).

RESEARCHERS RESPONDIn a truly serendipitous occurrence,just two months prior to HurricaneAndrew, the U. S. Fish and WildlifeService had given the University ofFlorida permission to remove 100 eggsin June 1992 as the starter nucleus of alarge-scale captive propagation program.Following the destruction wrought byHurricane Andrew, the USFWS com-mitted major funding to continue thefield surveys and captive propagationprogram, and implement experimentalreintroduction of the species withinprotected habitat areas. The bulk of thecaptive propagation work was carriedout at the University of Florida, wherethe Boender/USFWS EndangeredSpecies Laboratory became availablein June 1993, along with screenedenclosures and greenhouse supportfacilities. As a result, the captive holdingbecame the only readily available sourcefor livestock reintroductions andprompted the rapid expansion of exist-ing livestock breeding to become oneof the largest endangered invertebratecaptive propagation programs in theU.S.

This highly fortuitous timing allowedfor the first successful mating of captiveadults (via hand-pairing) in March 1993and the successful captive production of31 diapausing pupae by July 1993. Eggsproduced from these captive femaleswere increased by additional eggsbrought from Adams Key in BiscayneNational Park in June 1993 and pro-duced 49 healthy pupae early that fall,the nucleus of the 1994, 1995, and 1996captive propagation programs2,S,6,9,10,1\ forthe 1995, 1996, and 1997 reintroductionreleases.

In spring 1995, the frrst reintroductionefforts were initiated. A total of 764pupae were released at 7 sites, from theDeering Estate in south Miami on themainland to Key Largo. Despite heavypredation by migrating warblers, suc-

24 GREEN SECTION RECORD

cessful adult emergence and subsequentreproduction were identified at all sites,representing the first time since 1924that the Schaus swallowtail was foundon the south Florida mainland. Thesubsequent 1996 and 1997 releases of500 and 209 adult butterflies enhancedthe previous year's offSpring in theexisting population and established,directly or indirectly (via local move-n1ents),6 additional colonies in theUpper to Middle Keys.

Following these three years of intro-ductions, as of June 1997, the butterflyoccupied sites stretching from the southMiami area in Dade County to LowerMatecumbe Key in the Middle Keys ofMonroe County, across a graphic rangeof 57 miles. Thus the reintroductionshave resulted in the quadrupling of thespecies' geographic range from what itwas in the 20 years prior to the destruc-tion by Hurricane Andrew. Addition-ally, the total annual wild adult Schausswallowtail population rose to morethan 1,200 butterflies as of the 1997flight season. Still, the celebration ofthe project's success was tempered bylingering concerns regarding existinghabitat quality and long-term manage-ment as well as efficient gene flowbetween populations.

In early 1998, under direct fundingsupport from the National Fish andWildlife Foundation (NFWF), habitatimprovement was initiated by theplanting of hundreds of wild lime trees(Zanthoxylum fagara), one of two nativelarval host plants, within several selectedKey Largo colonies. The ultimate goalwas to improve and expand the suitablebreeding habitat available to the butter-fly within already existing protectedcolony sites and allow for the naturalincrease of the wild population to suffi-cient and stable levels. While habitatimprovement was currently beingaddressed, efficient gene flow betweenthe numerous existing and newly estab-lished colonies remained a criticalconcern.

Historically, the Schaus swallowtailenjoyed an intact range of pristine

tropical hardwood hammock habitatthroughout much of the Florida Keys,broken only by periodic but negotiablewater barriers. Individuals from neigh-boring colonies regularly infiltratedeach other, allowing for more or lessconstant gene flow between popula-tions. Additionally, wild populationnumbers annually waxed and waned,creating periodic localized extinctionsthat could be overcome by founderindividuals wandering in from nearbycolonies.

Today, the remaining Schaus swallow-tail populations no longer have thatsimple luxury. Adult butterflies nowhave to deal with urban developmentthat has left the Florida Keys with apatchwork of isolated and often distantpockets of suitable habitat, making con-tact between colonies an ever increas-ingly difficult task. Since all newlyestablished colonies were derived froma relatively small initial nucleus ofmaterial obtained from a single colony,all clearly face the continued threat of anarrowing genetic base, as well asunforeseen future natural disasters.

USGAAND FLORIDA GOLFCOURSES GET INVOLVEDThe opportunity to develop a viablecorridor system to encourage adultbutterfly movement and regular geneflow between colonies presented itselfin the spring of 1999 through grantfunding from the USGA's WildlifeLinks Program and the NFWF, andwith the direct cooperation of the U.S.Fish and Wildlife Service, the Universityof Florida, and two private golf clubs.The project, funded for three years,involved improving and restoring re-maining tropical hardwood hammockhabitat on the golf course property ofSombrero Country Club in Marathonand Cheeca Lodge on Islamorada.

Native larval host plants and adultnectar sources are being planted tocreate sufficient natural habitat suitableto maintain transient adult butterflies,encourage adult movement and geneflow between existing colonies, and

allow for the natural establishment ofnew breeding colonies within the Keys.Central to the project's success is thecooperation achieved between thegovernment agencies, private organiza-tions, and private landowners involved,including the current development oftwo Safe Harbor agreements.

If the current proj ect proceeds asexpected and additional funding be-comes available for additional years ofcaptive propagation, reintroduction, andmonitoring, the Schaus swallowtail willoccur over a broad enough geographicrange in protected habitat areas and insufficient areas and in sufficient numbersoutside of the Biscayne National Parkpopulation that a major catastrophicevent such as a hurricane, fire, or otherfocused environmental event no longerthreatens extinction or major depletionof the species. At such a point, it seemsevident that reclassification of thebutterfly's status from endangered tothreatened can occur, making it the firstinvertebrate successfully removed fromthe U.S. endangered species list.

LITERATURE CITED1. Collins, N. M., and M. G. Morris. 1985.Threatened swallowtail butterflies of the world:

The IUCN Red Data Book. Cambridge:IUCN.

2. Emmel, T. C. 1985. Status survey of the Schausswallowtail in Florida in 1984. Technical ReportNo. 14. Florida Cooperative Fish and WildlifeResearch Unit, University of Florida,Gainesville. 39 pp.

3. Emmel, T. C. 1986. Status survey and habitatrequirements of Florida's endemic Schausswallowtail butterfly. Florida Game and Fresh-water Fish Commission, Nongame WildlifeSection, Gainesville.

4. Emmel, T. C. 1988. Habitat requirements andstatus of the endemic Schaus swallowtail in theFlorida Keys. Florida Game and Freshwater FishCommission, Nongame Wildlife Section,Tallahassee. 202 pp.

5. Emmel, T. c., P.J. Eliazar,J. C. Daniels, S. D.Larson, andJ.A. Sarvis. 1993. Status monitoringand experimental reintroduction of the endan-gered Schaus swallowtail. Florida CooperativeFish and Wildlife Research Unit Annual Report,January to December 1992. Gainesville, Florida.Pp.19-20.

6. Emmel, T. c., P.J. Eliazar,]. C. Daniels, M. C.Minno, S. D. Larson, L. L. Groce,J. A. Fletcher,andJ. L. Nation,Jr. 1994. Captive propagationand habitat reintroduction for the Schausswallowtail following Hurricane Andrew (RWO113). Pp. 20-21. In Florida Cooperative Fish andWildlife Research Unit Annual Report,Januaryto December 1993. Gainesville, Florida. 54 pp.

7. Emmel, T. c., P.J. Eliazar,J. C. Daniels, M. C.Minno, S. D. Larson, and J. A. Sarvis. 1994. Statusmonitoring and experimental reintroduction ofthe endangered Schaus swallowtail (RWO 84). P.20, in Florida Cooperative Fish and WildlifeResearch Unit Annual Report,January toDecember 1993. Gainesville, Florida.

8. Emmel, T. C. 1995. Habitat requirements andstatus of the endemic Schaus swallowtail in theFlorida Keys. Florida Game and Freshwater FishCommission, Nongame Wildlife Section,Tallahassee. 202 pp.

9. Emmel, T. c., A. Sourakov, P.J. Eliazar,J. C.Daniels, V. Kroutov,J. Hall, K. Willmott, S. D.ScWachta,J. B. ScWachta, S. Sanchez, R. Worth,and K. A. Schwarz. 1998. Captive propagationand experimental reintroduction of Florida'sSchaus swallowtail (RWO 151). FloridaCooperative Fish and Wildlife Research UnitAnnual Report,January to December 1997.Gainesville, Florida.

10. Emmel, T. c., A. Sourakov, P.J. Eliazar,J. C.Daniels, V. Kroutov,J. Hall, K. Willmott, S. D.ScWachta,J. B. ScWachta, S. Sanchez, N. Eliazar,I. D. Kincade, and R. Morarnz. 1998. Breedingand reintroduction of the endangered Schausswallowtail (RWO 179). Florida CooperativeFish and Wildlife Research Unit Annual Report,January to December 1997. Gainesville, Florida.

11. Kimball, C. P. 1965. The Lepidoptera ofFlorida. An annotated checklist. Arthropods ofFlorida and neighboring land areas. Volume 1.Gainesville: Division of Plant Industry, FloridaDepartment of Agriculture.

12. Smith, D. S., L. D. Miller, andJ.Y Miller.1994. The butterflies of the West Indies andsouth Florida. Oxford University Press, NewYork.

JARET C. DANIELS, PH.D., and THOMAS

C. EMMEL, PH.D., McGuire Center forLepidoptera and Biodiversit}/JFloridaMuseum if Natural Histor}/JUniversity ifFlorida) Gainesville) Florida.

The Schaus swallowtail isconsidered one of therarest resident butterfliesin North America and islisted as an endangeredspecies by both the state ofFlorida and the federalgovernment.Wild limeplants serve as host sitesfor swallowtail eggs.

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