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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 301 1, 43 pp., 85 figures, 4 tables June 26, 1991 Cladistics and the Biogeography of Two Trans-Caribbean Hairstreak Butterfly Genera: Nesiostrymon and Terra (Lepidoptera, Lycaenidae) KURT JOHNSON' ABSTRACT Two sister genera composed of notably non- T. hispaniola Johnson and Matusik (southern His- vagile members distributed both in the Greater paniola); on the mainland, type species T. tera Antilles and mainland Neotropics are revised. (Hewitson) (central Mexico to Colombia); three Cladistic classification is employed from numer- species transferred from Thecla, T. calchinia ical analysis (PAUP, Swofford) ofthe ingroup and (Hewitson) (upper Amazon), T. cana (Hayward) three outgroups restricted to the mainland. Ne- (northwest Argentina), T. chilica (Schaus) (south- siostrymon includes five species: in the Antilles, east Brazil, Paraguay); and one new species, T. type species N. celida (Lucas) (with three interis- andevaga (Ecuador). Comparison of area clado- land subspecies-nominate [Cuba] N. c. aibonito grams of Nesiostrymon and Terra shows congru- (Comstock and Huntington) [Puerto Rico and ence of plesiotypic Antillean elements and apo- northern Hispaniola], N. c. baorucoensis, new sub- typic members occurring in Central America and species [southern Hispaniola]), and N. shoumatoffi South America, respectively. These are construed (Comstock and Huntington) (Jamaica); on the as more compatible with late-Cretaceous-Tertiary mainland, pan-Neotropical N. celona (Hewitson) vicariance of a proto-Greater Antilles from the (transferred from Thecla) and two new insular spe- Central and South American regions than a com- cies, N. milleri (cloud forests, Aragua, Venezuela) plex dispersal explanation requiring numerous and N. australivaga (scrub-steppe, Mendoza, Ar- concordant dispersals. gentina). Terra includes six species: in the Antilles, ' Research Associate, Department of Entomology, American Museum of Natural History. Copyright © American Museum of Natural History 1991 ISSN 0003-0082 / Price $5.00 2 AMERICAN MUSEUM NOVITATES NO. 3011 INTRODUCTION Nesiostrymon Clench (1964) and Terra tory (FMNH); Hope Entomological Collec- Johnson and Matusik (1988) are sister genera tion, Oxford University (HEC); Instituto of notably nonvagile "hairstreak butterflies" Miquel Lillo (Tucuman, Argentina) (IML); (Theclinae, Eumaeini) occurring both in the Los Angeles County Museum (LACM); Mil- Antilles and the mainland Neotropics. Nu- waukee Public Museum (MPM); Museum merical cladistic analyses associated with the National d'Histoire Naturelle (Paris, France) description of Terra indicated that it and Ne- (MNHN); Museo Nacional de Historia Nat- siostrymon are the sister group of three eu- ural (Santo Domingo, Dominican Republic) maeine assemblages restricted to the main- (MNDR); Zoologisches Museum der Hum- land (Johnson and Matusik, 1988). Observing boldt Universitiit zu Berlin (ZMH). Speci- the local habitat restriction and widely dis- mens were also examined from three private junct distributions ofNesiostrymon and Ter- collections with extensive Antillean or aus- ra taxa, Johnson and Matusik (1988) sug- tral South American holdings: Robert C. Ei- gested that Antillean species of these genera sele Collection (Jujuy, Argentina) (REC); Da- (along with two other recently discovered vid Matusik Collection (Skokie, Illinois) south Hispaniolan endemics) most likely "re- (DM); and Albert Schwartz Collection (Mi- flect a tectonic relationship between ... ami, Florida) (ASC). regions . on the northern edge of the Ca- ribbean plate." SYSTEMATICS Recent biogeographical studies (Shields and Phylogenetic Analyses ofNesiostrymon and Dvorak, 1979; Buskirk, 1985; Johnson and Terra: Johnson and Matusik (1988) used nu- Descimon, 1989; Miller and Miller, 1989) merical cladistic analysis (PAUP; Swofford, recognized a component of Antillean Rho- 1985) to establish the monophyly of Nesios- palocera more suggestive of an ancient vi- trymon, Terra, and certain outgroup taxa of cariant origin than the prominent waif-dis- the lycaenid grade "Thecla" (sensu Bridges, persal origin attributed to the fauna by early 1988) (fig. 1; tables 1, 2). Focusing primarily workers (Comstock and Huntington, 1949; on the Caribbean region, Johnson and Ma- Scott, 1972; Riley, 1975; Brown, 1978). Mil- tusik (1988) distinguished Antillean mem- ler and Miller (1989), summarizing a prob- bers of Nesiostrymon and Terra and listed able vicariance and dispersal model for mainland relatives from the "Thecla" grade. Antillean butterfly origins, included Nesio- Subsequently, Johnson (1991) published a strymon and Terra as one of nine butterfly nomenclature for outgroup taxa (figs. 84, 85). groups most probably arising from late Cre- The cladistic study of Johnson and Ma- taceous-Eocene vicariance. tusik (1988) indicated that Terra and Ne- This study reports a cladistic taxonomy of Neotropical Nesiostrymon and siostrymon form a monophyletic group Terra and distinguished from outgroups by six synapo- presents the ecological and ev- biogeographic morphies of the wings, genitalia, and termi- idence supporting a vicariant origin for An- tillean members of two nal abdominal morphology (table 1: charac- these hairstreak but- ters 2, 4-6, 12, and Terra terfly genera. 14). taxa share a salient apomorphy: the prominent sclero- tized bulb protruding ventrally between the MATERIALS AND METHODS cephalic and caudal elements of the ductus COLLECrIONS bursae (table 1, character 11). Nesiostrymon taxa share a specialized prominently pronged Specimens examined included samples tergite 8 in males (table 1, characters 1, 9). from the Allyn Museum of Entomology, High consistency values (fig. 1) for clado- Florida Museum of Natural History (AME); grams result from the various very distinctive American Museum of Natural History morphological characters which separate (AMNH); British Museum (Natural History) these taxa (e.g., the incised posterior cavity (BMNH); Carnegie Museum of Natural His- ofmale tergite 8, distribution ofmicrotrichia, tory (CMNH); Field Museum ofNatural His- etc.). "Outgroup" and "Lundberg outstate" 1991 JOHNSON: HAIRSTREAK BUTTERFLY GENERA 3 AB C D A B CD E F 5-' - 1 4 12 ' 2,4,5,6,12,1 4 1 , 9 T' caption, A caption, B caption, C (Tb. 1) 2,4,5,6,12,1l4 Fig. 1. Cladogram ofNesiostrymon (D), Terra (C), and relatives (A, B) (Johnson and Matusik, Fig. 2. Ingroup cladogram ofNesiostrymon (A, 1988). Four-taxon statement derived from parsi- B, C) and Terra (D, E, F). Six-taxon statement monious distribution ofunweighted characters and derived from parsimonious distribution of un- rooted using (1) outgroup[s] described in text (con- weighted characters and rooted using Lundberg sistency index = 0.923) and (2) Lundberg method outstates based on monophyly of Nesiostrymon (Swofford, 1985) based on presumed primitive and Terra, figure 1. Consistency index = 0.909. states listed at bottom oftable 2 (consistency index Apomorphies are specified by horizontal bars and = 0.889). Apomorphies for Nesiostrymon and Ter- represent characters enumerated in tables 3 and ra specified by horizontal bars enumerated from 4. Synapomorphies noted for Nesiostrymon/Terra 1 tables and 2. Outgroups and their apomorphies: are from figure 1, tables 1 and 2. Terminal groups A, B. Outgroup rooting preferred " Thecla uzza include A, N. celona/milleri/australivaga; B, N. complex" (fig. 84) as sister group ofNesiostrymon! celida; C, N. shoumatoffi; D, T. hispaniola; E, T. Terra based on characters 9, 10 with " Thecla cel- tera; F, T. andevaga/chilica/calchinia/cana. mus complex" (fig. 84) as the outgroup; Lundberg rooting preferred the opposite based on characters, 1, 3, 7, 13. C. "Thecla" outgroup (fig. 85), char- acters enumerated by Johnson and Matusik, 1988. demic T. hispaniola being the most primitive in character. Based on these results, species- level taxonomies for Nesiostrymon and Terra (Swofford, 1985) rootings (fig. 1) give iden- are enumerated below. tical results concerning the monophyly of Terra and Nesiostrymon. TAXONOMY OF NESIOSTRYMON and To determine a cladistic classification of TERRA species in Nesiostrymon and Terra, parsi- GENUS NESIOSTR YMON CLENCH monious distributions of characters were constructed for shared characters of species Figures 3-32, 57, 59-68 in each genus (fig. 2, tables 3, 4). Monophyly Nesiostrymon Clench, 1964: 248, 251. Brown and of the ingroup was assumed from Johnson Heineman, 1971: 4; 1972: 232. - Scott, 1986: and Matusik's (1988) results; species criteria 21. - Riley, 1975: 101. - Bridges, 1988: 1.74, were based on standard taxonomic proce- II.77. - Johnson and Matusik, 1988: 236. - dures involving consistent in Schwartz, 1989: 240. - Miller and Miller, 1989: differences 245. characters of the wings and genitalic and ter- gal morphology. Table 3 lists the apomorphic DIAGNOSIS: Compared to all Eumaeini: and plesiomorphic states ofcharacters delim- male tergite 8 modified to a "subcordate in- ited from the final rooted tree (fig. 2). This cised posterior cavity" (the "sipc" sensu Field, analysis indicates the following: Nesiostry- 1967a, 1967b; Johnson, 1988a, 1989a, 1989b; mon is a monophyletic group with two nested Johnson and Matusik, 1988) terminating in sets of taxa, the Jamaican endemic, N. shou- a dorsally produced lobe or prong, genital matoffi, being the most primitive in charac- valvae multiplanar with protruding, winglike ter; Terra is a monophyletic
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