insects Review Jamaica’s Critically Endangered Butterfly: A Review of the Biology and Conservation Status of the Homerus Swallowtail (Papilio (Pterourus) homerus Fabricius) Matthew S. Lehnert 1,* , Valerie R. Kramer 1, John E. Rawlins 2, Vanessa Verdecia 2 and Jaret C. Daniels 3,4 1 Department of Biological Sciences, Kent State University at Stark, North Canton, OH 44720, USA; [email protected] 2 Carnegie Museum of Natural History, Pittsburgh, PA 15213, USA; [email protected] (J.E.R.); [email protected] (V.V.) 3 McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, Gainesville, FL 32611, USA; jdaniels@flmnh.ufl.edu 4 Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611, USA * Correspondence: [email protected]; Tel.:+1-330-244-3349; Fax: +1-330-244-3482 Academic Editor: Brian T. Forschler Received: 21 June 2017; Accepted: 7 July 2017; Published: 10 July 2017 Abstract: The Homerus swallowtail, Papilio (Pterourus) homerus Fabricius, is listed as an endangered species and is endemic to the Caribbean island of Jamaica. The largest butterfly in the Western Hemisphere, P. homerus once inhabited seven of Jamaica’s 14 parishes and consisted of at least three populations; however, now only two stronghold populations remain, a western population in the rugged Cockpit Country and an eastern population in the Blue and John Crow Mountains. Despite numerous studies of its life history, much about the population biology, including estimates of total numbers of individuals in each population, remains unknown. In addition, a breeding program is needed to establish an experimental population, which could be used to augment wild populations and ensure the continued survival of the species. Here, we present a review of the biology of P. homerus and recommendations for a conservation plan. Keywords: Homerus swallowtail; Jamaica; Papilio homerus; conservation; endangered species; flagship species 1. Introduction The Homerus swallowtail (Papilio (Pterourus) homerus Fabricius 1793) (Figure1) is the largest butterfly in the Western Hemisphere [1,2] and is critically endangered [3–5]. Endemic to the island of Jamaica [6,7], P. homerus is a flagship species that symbolizes the need for conservation efforts [8] and has been featured on postage stamps and the Jamaican $1000 bill (Figure2). Papilio homerus is listed in the IUCN Red Data Book, Threatened Swallowtail Butterflies of the World [3], and is protected as an Appendix I species by the Convention for International Trade in Endangered Species (CITES) and the Jamaican Wildlife Act of 1988. The range of this enigmatic butterfly, possibly less than 10 km2 for each of the two remaining populations, continues to dwindle, particularly due to habitat destruction [4] and recent mining initiatives [7,9]. In addition, the extant populations of P. homerus reside in areas of Jamaica famous for their difficult terrain [5,7,10,11], which further confounds the ability to perform the long-term field studies necessary for understanding the biology and population dynamics for this species. Despite these complications, lab and field studies have given insight into the ecology, Insects 2017, 8, 68; doi:10.3390/insects8030068 www.mdpi.com/journal/insects Insects 2017, 8, 68 2 of 16 Insects 2017, 8, 68 2 of 16 history, and behavior of P. homerus. Here, we provide an overview of the biology of P. homerus, history, and behavior of P. homerus. Here, we provide an overview of the biology of P. homerus, lifeincluding history, knowledge and behavior gaps of andP. homerusobjectives. Here, for future we provide conservation an overview efforts offor the this biology imperiled of P. butterfly. homerus, including knowledge gaps and objectivesobjectives forfor futurefuture conservationconservation effortsefforts forfor thisthis imperiledimperiled butterfly.butterfly. Figure 1. Photographs of representative Papilio homerus from the eastern population. (A) Female of P. Figure 1. Photographs of representative Papilio homerus from the eastern population. (A) Female of P. Figurehomerus 1. (dorsalPhotographs on left, ofventral representative on right) collectedPapilio homerus from thefrom Blue the Mountains, eastern population. Jamaica (1959). (A) Female (B) Male of homerus (dorsal on left, ventral on right) collected from the Blue Mountains, Jamaica (1959). (B) Male P.of homerusP. homerus(dorsal collected on left, at ventralCorn Puss on right) Gap, collectedJamaica (1986). from the All Blue photographs Mountains, are Jamaica used with (1959). permission(B) Male of P. homerus collected at Corn Puss Gap, Jamaica (1986). All photographs are used with permission offromP. homerus Warrencollected et al. at (2016) Corn Puss and Gap, acquired Jamaica from (1986). http://butterflie All photographssofamerica.com/L/t/Pterourus_ are used with permission from from Warren et al. (2016) and acquired from http://butterfliesofamerica.com/L/t/Pterourus_ Warrenhomerus_a.htm. et al. (2016) and acquired from http://butterfliesofamerica.com/L/t/Pterourus_homerus_a.htm. homerus_a.htm. Figure 2. Printings and postage stamps of Papilio homerus that contribute to its status as a flagship Figure 2. Printings and postage stamps of Papilio homerus thatthat contribute to to its status as a flagshipflagship species. (A) The Jamaican $1000 bill features P. homerus in the upper-right corner. Representative species. ( (AA)) The The Jamaican Jamaican $1000 $1000 bill bill features P. homerushomerus inin thethe upper-rightupper-right corner.corner. RepresentativeRepresentative stamps issued in 1964 (B), 1975 (C), 1989 (D), and 1994 (E,F) that feature P. homerus. stamps issued in 1964 (B), 1975 (C), 1989 ((D),), andand 19941994 ((E,,F)) thatthat featurefeature P.P. homerushomerus.. Insects 2017, 8, 68 3 of 16 2. Papilio homerus Historical Records Insects 2017, 8, 68 3 of 16 The grandeur of the Homerus swallowtail has inspired artists since its discovery by Western culture.2. PapilioPapilio homerus homerus Historicalwas first Records described by Johan Christian Fabricius [12] (not from a specimen, but fromThe a painting grandeur by of William the Homerus Jones, swallowtail Esq. of Chelsea, has inspired Figure artists3C) [ 13since], who, its discovery awestruck by byWestern its beauty, namedculture. thebutterfly Papilio homerusPapilio was homerus first describedin honor by of Johan the GreekChristian poet, Fabricius Homer, [12] the (not author from a of specimen,The Iliad and The Odysseybut from. Evena painting before by William Jones’ paintings, Jones, Esq. P.of homerusChelsea, Figurewas the 3C) subject [13], who, of paintings awestruck byby Henryits beauty, Seymer (1768,named Figure the3A) butterfly and collaborative Papilio homerus paintings in honor withof the his Greek son, poet, Henry Homer, Seymer the author Jr. (1773, of The Figure Iliad 3andB) [ 13]. ThreeThe years Odyssey after. Even the descriptionbefore Jones’ by paintings, Fabricius, P. homerus the German was the entomologist subject of paintings Eugenius by Henry Johann Seymer Christoph (1768, Figure 3A) and collaborative paintings with his son, Henry Seymer Jr. (1773, Figure 3B) [13]. Esper illustrated P. homerus (Figure3D) in his series of booklets, Die Schmetterlinge in Abbildungen nach Three years after the description by Fabricius, the German entomologist Eugenius Johann Christoph der Natur mit Beschreibungen [14]. Much of the biology of P. homerus, however, remained unreported Esper illustrated P. homerus (Figure 3D) in his series of booklets, Die Schmetterlinge in Abbildungen nach untilder Rutherford Natur mit [Beschreibungen15] and Gosse [14]. [16] Much provided of the information biology of P. of homerus the immature, however, stages remained and generalunreported habits of the butterfly.until Rutherford The morphology [15] and Gosse of the[16] immature provided information stages of P. of homerus the immaturewas also stages discussed and general by Pantonhabits [17] and Taylorof the butterfly. [18]. Many The morphology of these early of the reports immature were stages anecdotal of P. homerus and itwas was also not discussed until descriptions by Panton by Andr[17]é Avinoff and Taylor and [18]. his nephew,Many of these Nicholas early reports Shoumatoff, were anecdotal that details and wereit was providednot until descriptions about the biologyby and preferredAndré Avinoff habitats and his of thenephew, adult Nicholas life stage Shoumatoff, ([19], discussed that details in [ 20were]). Despiteprovided recent about the efforts biology to study the biologyand preferred of P. homerushabitats of[4 –the7,10 adult,11, 21life– 24stage], there ([19], are discussed still gaps in [20]). in our Despit knowledgee recent efforts about to the study species, particularlythe biology about of theP. homerus biology [4–7,10,11,21–24], of the adults, the there total are number still gaps of individualsin our knowledge remaining about in the the species, stronghold particularly about the biology of the adults, the total number of individuals remaining in the populations, and population dynamics. In addition, information about the genetic architecture of stronghold populations, and population dynamics. In addition, information about the genetic P. homerus populations is completely lacking because no genetic studies have been performed, which is architecture of P. homerus populations is completely lacking because no genetic studies have been necessaryperformed, for the which development is necessary of for an the effective development conservation of an effective