Journal of Insect Conservation

, Volume 12, Issue 3–4, pp 383–397 | Cite as

Population studies and conservation of Jamaica’s endangered swallowtail butterfly Papilio (Pterourus) homerus

  • Eric Garraway
  • A. J. A. Bailey
  • B. E. Freeman
  • J. R. Parnell
  • T. C. Emmel
Original Paper


This is an 18-year study of the endangered Papilio (Pterourus) homerus, adding substantial information to our scanty knowledge of its ecology. The contraction of a once contiguous but narrow population on a single Caribbean island carries the serious threat of extinction. There are now two populations or probably metapopulations, effectively isolated from each other. The butterfly’s larvae feed on Hernandia catalpaefolia and H. jamaicensis, both endemic to Jamaica, and development takes ∼84 days from egg to the emerged adult. Adult numbers fluctuate rapidly, with peaks in July/August each year. Egg distribution was studied at three spatial levels: the food item (leaf cluster), the patch (tree) and the habitat (each valley). Major causes of developmental mortality were Chrysonotomyia, a eulophid parasitoid of the eggs, and bacterial infection of the larvae and pupae. Critically, the mortality from this wasp was lower in undisturbed forest than in the area disturbed by agriculture, this finding having important consequences for conservation. Although there was no evidence of a decline in numbers over the last century, we believe this is an artefact due to collectors working only at the periphery of its distribution. Even assuming that its population densities have not changed, the contraction of its usable habitat implies a similar reduction in average numbers and the small populations are susceptible to disaster. The efforts of researchers, NGOs, and Government agencies have greatly increased the level of awareness, making the people in some key areas the ‘protectors of the species’.


Egg parasitoids Physical factors Polymorphism Natural deception Population dynamics Spatial distribution Rare species Metapopulations 



Dr. George R. Proctor of the Department of Life Sciences University of the West Indies; The Jamaica Parrot Project, especially Mr. Herlitz Davis and Dr. Noel Snyder; The Butterfly Group of the Natural History Society of Jamaica, especially Catherine Murphy and Carol Miller; the Windsor Research Center, and Paula Hurlock of Dolphin Head Trust, provided useful data and information. Mr. Errol Francis and Mr. Orlando Wilson were masterful field assistants. Funding was received from the Jamaica Agricultural Research Program.


  1. Aaron EM (1893) New localities for Papilio homerus. Can Ent 25:258Google Scholar
  2. Adams CD (1972) Flowering plants of Jamaica. University of the West Indies, JamaicaGoogle Scholar
  3. Andrewartha HG, Birch LC (1954) The distribution and abundance of animals. The University of Chicago Press, ChicagoGoogle Scholar
  4. Avinoff A, Shoumatoff N (1940) Jamaican summer. Carnegie Mag (Pittsburgh) 14:175–182Google Scholar
  5. Bailey AJ (2003) The biology and ecology of the endangered giant swallowtail butterfly, Papilio (Pterourus) homerus (Lepidoptera: Papilionidae) in Jamaica. PhD thesis, University of the West IndiesGoogle Scholar
  6. Beaver RA (1966) The development and expression of population tables for the bark beetle Scolytus scolytus (F.). J Anim Ecol 35:27–41CrossRefGoogle Scholar
  7. Boughton DA (1999) Empirical evidence for complex source-sink dynamics with alternative states in a butterfly metapopulation. Ecology 80:2727–2739Google Scholar
  8. Brown FM, Heineman B (1972) Jamaica and its Butterflies. E.W.Classey, LondonGoogle Scholar
  9. Butler PJ (1994) The Jamaica conservation education program. RARE Center for Tropical Conservation, Philadelphia, PennsylvaniaGoogle Scholar
  10. Chew FS, Robins RK (1984) Egg laying in butterflies. In: Vane-Wright RI, Ackery PR (eds) The biology of butterflies. Academic Press, London, pp 65–79Google Scholar
  11. Cockpit Country Stakeholders Group (2006) Bauxite mining in the Cockpit Country. Cockpit Country Stakeholders Group, Cited 12 October, 2007
  12. Collins NM, Morris MG (1985) Threatened swallowtail butterflies of the World. The IUCN Red Data Book. International Union for the Conservation of Nature and Natural Resources, CambridgeGoogle Scholar
  13. Cromartie WJ (1975) The effect of stand size and vegetational background on the colonization of cruciferous plants by herbivorous insects. J Appl Ecol 12:517–533CrossRefGoogle Scholar
  14. Dempster J (1983) The natural control of populations of butterflies and moths. Biol Rev 58:461–481CrossRefGoogle Scholar
  15. Dempster JP, Hall ML (1980) An attempt at establishing the swallowtail butterfly at Wicken Fen. Ecol Ent 5:327–334CrossRefGoogle Scholar
  16. Dolphin Head Trust (2007) Dolphin Head Trust. Cited 12 October, 2007
  17. Emmel TC (1995) Saving endangered swallowtails. The conservation biology of Papilio aristodemus ponceanus in Florida and P. homerus in Jamaica. In: Scriber JM, Tsubaki Y, Lederhouse RC (eds) Swallowtail butterflies: their ecology and evolutionary biology. Scientific Publishers, Gainesville, pp 359–369Google Scholar
  18. Emmel TC, Garraway E (1990) Ecology and conservation biology of the Homerus swallowtail in Jamaica (Lepidoptera: Papilionidae). Trop Lep 1:63–76Google Scholar
  19. Evelyn OB, Camirand R (2003) Forest cover and deforestation in Jamaica: an analysis of forest cover estimates over time. Int For Rev 5:354–363Google Scholar
  20. Eyre LA (1986) Deforestation in Jamaica: its rate and implications. Department of Geography, University of the West Indies, KingstonGoogle Scholar
  21. Eyre LA (1987) Deforestation in Jamaica. J Sci Res Counc Jamaica 6:17–24Google Scholar
  22. Feeny P(1995) Ecological opportunism and chemical constraints on the host associations of swallowtail butterflies. In: Scriber JM, Tsubaki Y, Lederhouse RC (eds) Swallowtail butterflies: their ecology and evolutionary biology. Scientific Publishers, Gainesville, pp 9–15Google Scholar
  23. Freeman BE (1976) A spatial approach to insect population dynamics. Nature 260:240–241PubMedCrossRefGoogle Scholar
  24. Freeman BE (1981) The dynamics in trinidad of the Sphecid Wasp Trypoxylon palliditarse: a Thompsonian population? J Anim Ecol 50:563–572CrossRefGoogle Scholar
  25. Freeman BE, Smith DC (1990) Variation of density-dependence with spatial scale in the leaf-mining fly Liriomyza commelinae (Diptera, Agromyzidae). Ecl Ent 15:265–274CrossRefGoogle Scholar
  26. Garraway E, Freeman BE (1981) Studies on the population dynamics of the Juniper bark beetle (Phloeosinus neotropicus) in Jamaica. Oikos 36:363–368CrossRefGoogle Scholar
  27. Garraway E, Bailey AJA (1992) Parasitoid induced mortality in the eggs of the endangered giant swallowtail butterfly Papilio homerus (Papilionidae). J Lepidopt Soc 46:233–234Google Scholar
  28. Garraway E, Parnell J (1993) Notes on the osmeteria of Papilio homerus (Lepidoptera: Papilionidae). Trop Lep 4:29–30Google Scholar
  29. Garraway E, Bailey AJA, Emmel TC (1993) Contribution to the ecology and conservation biology of the endangered Papilio homerus (Lepidoptera: Papilionidae). Trop Lep 4:83–91Google Scholar
  30. Gilbert LE, Singer MC (1973) Dispersal and gene flow in a butterfly species. Am Nat 107:58–72CrossRefGoogle Scholar
  31. Gosse PH (1879) On Papilio homerus, its ovum and larvae. Proc Ent Soc: 1v–lviiiGoogle Scholar
  32. Government of Jamaica (1945) Jamaica Wild Life Protection Act. Govt. JamaicaGoogle Scholar
  33. Hanski I, Gyllenberg M (1993) Two general metapopulation models and the core-satellite species hypothesis. Am Nat 142:17–41CrossRefGoogle Scholar
  34. Hanski I, Kuussaari M, Nieminen M (1994) Metapopulation structure and migration in the butterfly Melitaea cinxia. Ecology 75:747–762CrossRefGoogle Scholar
  35. Hassell MP, Southwood TRE (1978) Foraging strategies in insects. Ann Rev Ecol Syst 9:75–98CrossRefGoogle Scholar
  36. Hazel WN (1995) The causes and evolution of phenotypic plasticity in pupal colour in swallowtail butterflies. In: Scriber JM, Tsubaki Y, Lederhouse RC (eds) Swallowtail butterflies: their ecology and evolutionary biology. Scientific Publishers, Gainesville, pp 205–210Google Scholar
  37. Headley M, Evelyn OB (2000) Jamaica country report. Proceedings of the sub-regional workshop on data collection and outlook effort for forestry in the CaribbeanGoogle Scholar
  38. Higman BW (1988) Jamaica surveyed: plantation maps and plans of the eighteenth and nineteenth centuries. Institute of Jamaica Publications Ltd., KingstonGoogle Scholar
  39. Hinton HE (1973) Natural deception. In: Gregory RL, Gombrich EH (eds) Illusion in nature and art. Scribners, New YorkGoogle Scholar
  40. Hirose Y, Takagi M (1995) Population dynamics and life history of selected Papilionidae. In: Scriber JM, Tsubaki Y, Lederhouse RC (eds) Swallowtail butterflies: their ecology and evolutionary biology. Scientific Publishers, Gainesville, pp 107–113Google Scholar
  41. Hirose Y, Suzuki I, Tagaki M, Hichata K, Yamasaki M, Kimoto H, Yamanaka M, Iga M, Yamaguchi K (1980) Population dynamics of the citrus swallowtail Papilio xuthus Linné (Lepidoptera: Papilionidae). Mechanisms stabilizing its numbers. Res Popul Ecol 21:260–285CrossRefGoogle Scholar
  42. Hooper EDM (1886) Report upon the forests of Jamaica. Waterlow and Sons Ltd., LondonGoogle Scholar
  43. Ittyeipe K, Taffe CA (1982) The biology and population dynamics of Monobia mochii Soika, a rare, solitary eumenid in Jamaica. Caribbean J Sci 17:45–58Google Scholar
  44. Janzen DH (1984) Two ways to be a tropical big moth: Santa Rosa Saturniids and Sphingids. Oxf Surv Evol Biol 1:85–140Google Scholar
  45. Jayasingh DB, Freeman BE (1980) The comparative population dynamics of eight solitary bees and wasps (Aculeata; Apocrita; Hymenoptera) trap-nested in Jamaica. Biotropica 12:214–219CrossRefGoogle Scholar
  46. Jones CG, Lawton JH (1994) Linking species and ecosystems. Chapman and Hall, New YorkGoogle Scholar
  47. Kaye WJ (1926) The butterflies of Jamaica. Trans R Ent Soc 1925:455–504Google Scholar
  48. Larsson S, Ekbom B (1995) Oviposition mistakes in herbivorous insects: confusion or a step towards a new host plant? Oikos 72:155–160CrossRefGoogle Scholar
  49. Lehnert MS (2008) The population biology and ecology of the homerus swallowtail, Papilio (Pterourus) homerus, in the Cockpit Country, Jamaica. J Insect Conserv 12:179–188CrossRefGoogle Scholar
  50. Levins R (1969) Some demographic and genetic consequences of environmental heterogeneity for biological control. Bull Entomol Soc Am 15:237–240Google Scholar
  51. Levins R (1970) Extinctions. In: Some mathematical questions in biology: lectures on mathematics in the life sciences. Am Math Soc, Providence, Rhode Island, pp 77–107Google Scholar
  52. Lewis CB (1944a) Butterfly notes. Nat Hist Notes, Nat Hist Soc Jamaica (Kingston) (old issue) 17:78Google Scholar
  53. Lewis CB (1944b) Butterfly notes and records. Nat Hist Notes, Nat Hist Soc Jamaica (Kingston) (old issue) 19:118Google Scholar
  54. Lewis CB (1948) Butterfly notes. Nat Hist Notes, Nat His Soc Jamaica (Kingston) (old issue) 3:204Google Scholar
  55. Lewis CB (1949) Butterfly notes. Nat Hist Notes, Nat Hist Soc Jamaica (Kingston) (old issue) 4:48Google Scholar
  56. Matsumoto K (1985) Population dynamics of the Japanese coloured Apollo Parnassius glacialis Butler (Lepidoptera: Papilionidae). I. Changes in population size and related population parameters for three successive generations. Res Popul Ecol 27:301–312CrossRefGoogle Scholar
  57. McPeek MA, Holt RD (1992) The evolution of dispersal in spatially and temporally varying environments. Am Nat 140:1010–1027CrossRefGoogle Scholar
  58. National Environmental Protection Agency (NEPA) (2002) The Giant Swallowtail Butterfly Recovery Plan. NEPA. Kingston, Jamaica. Cited 15 October, 2007
  59. National Environmental Protection Agency (NEPA) (2003) Jamaica’s commitment to the conservation and management of natural resources, ten years in retrospect. Unpublished Paper 2003, NEPA, Kingston, JamaicaGoogle Scholar
  60. Neufville Z (2001) Environment-Jamaica: Bauxite mining blamed for deforestation. Inter Press Service. Cited 11 November 2007
  61. Panton ES (1893) A description of the larva of Papilio homerus. J Inst Jamaica 1:375–376Google Scholar
  62. Parnell J (1984) Papilio homerus, the vanishing swallowtail. Documentary Video. University of the West Indies, JamaicaGoogle Scholar
  63. Parsons P (1992) Book Publishing at University Presses. In: Encyclopedia of Library and information Science, 49(Suppl 12)Google Scholar
  64. Proctor GR (1986) Cockpit Country and its vegetation. In: Thompson DA, Bretting PK, Humphreys M (eds) Forest of Jamaica. Institute of Jamaica Publications LtdGoogle Scholar
  65. Proctor GR (2001) Interim checklist of plants found in Dolphin Head project Area. Dolphin Head Trust. Cited 11 November, 2007
  66. Pulliam HR (1988) Sources, sinks, and population regulation. Am Nat 132:653–661CrossRefGoogle Scholar
  67. Renwick JAA, Chew FS (1994) Oviposition behavior in Lepidoptera. Ann Rev Entomol 39:377–400CrossRefGoogle Scholar
  68. Roland J, Taylor PD (1997) Insect parasitoid species respond to forest structure on several spatial scales. Nature 386:710–713CrossRefGoogle Scholar
  69. Rutherford DG (1878) Notes regarding some rare Papiliones. Ent Monthly Mag 15:28–31Google Scholar
  70. Satchell VM (1990) From plots to plantations: land transactions in Jamaica, 1866–1900. Inst Social and Econ Res, University of the West Indies, KingstonGoogle Scholar
  71. Scriber MS, Tsubaki Y, Lederhouse RC (eds) (1995) Swallowtail butterflies: their ecology and evolutionary biology. Scientific Publishers, GainesvilleGoogle Scholar
  72. South R (1939) The Months of the British Isles. Frederick Warne, LondonGoogle Scholar
  73. Southwood TRE (1978) Ecological methods, 2nd. edn. Chapman and Hall, LondonGoogle Scholar
  74. Swabey C (1945) Forestry in Jamaica. Forestry bulletin No. 1, forest department, KingstonGoogle Scholar
  75. Swainson EM (1901) Notes on lepidopterous larvae from Jamaica. J New York Ent Soc 9:77–82Google Scholar
  76. Taylor CB (1894) The description of the larva and pupa of Papilio homerus Fabricius. Trans Ent Soc 42:409–410Google Scholar
  77. Taylor L (1961) Aggregation, variance and the mean. Nature 189:732–735CrossRefGoogle Scholar
  78. Trivers RL (1985) Social evolution. Benjamin/Cummings, Menlo Park, CAGoogle Scholar
  79. Tsubaki Y (1973) The natural mortality and its factors of the immature stages of a population of the swallowtail butterfly Papilio xuthus. Jap J Ecol 23:210–217Google Scholar
  80. Turner TW (1991) Papilio homerus (Papilionidae) in Jamaica, West Indies: field observations and description of immature stages. J Lepidopt Soc 45:259–271Google Scholar
  81. Tyler HA, Brown KS, Wilson KH (1994) Swallowtail butterflies of the Americas. Scientific Publishers, GainesvilleGoogle Scholar
  82. Van Alphen JJM, Vet LEM (1986) An evolutionary approach to host finding and selection. In: Waage J, Greathead D (eds) Insect parasitoids. Academic Press, London, pp 23–61Google Scholar
  83. Vane-Wright RI, Ackery PR (eds) (1984) The biology of butterflies. Academic Press, LondonGoogle Scholar
  84. Varley GC (1947) The natural control of population balance in the knapweed gall fly (Urophora jaceana). J Anim Ecol 16:139–187CrossRefGoogle Scholar
  85. Vinson SB (1976) Host selection by insect parasitoids. Ann Rev Entomol 21:109–133CrossRefGoogle Scholar
  86. Walker DJR (1945) Papilio homerus. Nat Hist Notes, Nat Hist Soc Jamaica (Kingston) (old issue) 1:161–163Google Scholar
  87. Watanabe M (1976) A preliminary study on population dynamics of the swallowtail butterfly Papilio xuthus L., in a deforested area. Res Popul Ecol 17:200–210CrossRefGoogle Scholar
  88. Watanabe M (1981) Population dynamics of the swallowtail butterfly Papilio xuthus L., in a deforested area. Res Pop Ecol 23:74–93CrossRefGoogle Scholar
  89. Watanabe M (1983) Radial growth patterns in a pioneer tree, Zanthoxylum ailanthoides Sieb, et Zucc. (Rutales: Rutaceae) related to the population dynamics of a swallowtail butterfly, Papilio xuthus L. (Lpidoptera: Papilionidae). Jap J Ecol 33:253–261Google Scholar
  90. Watanabe M (1995) Population dynamics of Papilio xuthus Larvae in relation to life history of the host tree. In: Scriber JM, Tsubaki Y, Lederhouse RC (eds) Swallowtail Butterflies: their ecology and evolutionary biology. Scientific Publishers, Gainesville, pp 101–106Google Scholar
  91. Watkinson AR, Sutherland WJ (1995) Sources, sinks and pseudo-sinks. J Anim Ecol 64:126–130CrossRefGoogle Scholar
  92. Watmough RH (1983) Mortality, sex-ratio and fecundity in natural populations of large carpenter bees (Xylocopa spp.). J Anim Ecol 52:111–125CrossRefGoogle Scholar
  93. Wiklund C (1977) Oviposition, feeding and spatial separation of breeding and foraging habitats in a population of Leptidea sinapis. Oikos 28:56–68CrossRefGoogle Scholar
  94. Williams C (1964) Patterns in the balance of Nature. Academic Press, LondonGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2008

Authors and Affiliations

  • Eric Garraway
    • 1
  • A. J. A. Bailey
    • 1
  • B. E. Freeman
    • 1
  • J. R. Parnell
    • 2
  • T. C. Emmel
    • 3
  1. 1.Department of Life SciencesThe University of the West IndiesMonaJamaica
  2. 2.Safety HarborUSA
  3. 3.McGuire Center for Lepidoptera and BiodiversityFlorida Museum of Natural HistoryGainesvilleUSA

Personalised recommendations