THE JOURNAL OF AGRICULTURE OF THE UNIVERSITY OF PUERTO RICO Issued biannually by the Agricultural Experiment Station of the University of Puerto Rico, Mayagüez Campus, for the publication of articles and research notes by staff members or others, dealing with scientific agriculture in Puerto Rico and elsewhere in the Caribbean Basin and Latin America. VOL. 100 DECEMBER 2016 Special issue No. 1 Biodiversity of Heteroptera in Puerto Rico: Part I A Conspectus of Pentatomomorpha: Pentatomoidea1,2 Alejandro E. Segarra-Carmona3, Rosa A. Franqui3 and Hariette Pérez-Martínez4 J. Agric. Univ. P.R. 100 Special issue (1): (2016) ABSTRACT Pentatomoidea fauna for Puerto Rico and its adjacent islands is documented as part of a revised account of Heteroptera on these islands. There are 71 species in six families of Pentatomoidea known from Puerto Rico: Cydnidae (11); Pentatomidae (48) including Asopinae (8), Pentatominae (37), Discocephalinae (1), and Edessinae (2); Tessaratomidae (1); Scutelleridae (8); Megarididae (1); and Corimelaenidae (2). Of this total, seven represent new state records for Puerto Rico. Taxonomic accounts include synonymies, known distribution, lists of hosts, and a listing of specimens examined. Regional taxonomical keys are provided for the identification of taxa included. Color plates for most species are also included. Key words: biodiversity, Pentatomoidea fauna, insect, Heteroptera, taxa 1Manuscript submitted to Editorial Board 4 August 2015. 2Research funded by University of Puerto Rico Agricultural Experiment Station’s In- sect Research Collection Project (C-415). 3Professor, Department of Agroenvironmental Sciences, Agricultural Experiment Station, College of Agricultural Sciences, University of Puerto Rico-Mayagüez Campus. All correspondence to [email protected]. 4Research Associate, Department of Agroenvironmental Sciences, Agricultural Ex- periment Station, College of Agricultural Sciences, University of Puerto Rico. Mayagüez. 1 2 BIODIVERSITY OF HETEROPTERA IN PUERTO RICO: PART I RESUMEN La biodiversidad de Heterópteros en Puerto Rico: Parte I Un resumen de Pentatomomorpha: Pentatomoidea La fauna de Pentatomoidea de Puerto Rico y de sus islas adyacentes se presenta como parte de una revisión de los Heterópteros de estas islas. Se presentan 71 especies en seis familias de Pentatomoidea conocidas de Puerto Rico: Cydnidae (11); Pentatomidae (48): Asopinae (8), Pentatominae (37), Discocephalinae (1); Edessinae (2); Tessaratomidae (1); Scutelleridae (8); Megarididae (1); y Corimelaenidae (2). Siete especies representan récords nuevos para Puerto Rico. Para cada récord de las especies presentadas se incluyen sus sinonimias, su distribución conocida, sus hospederos, y una lista de los especímenes examinados. También se incluyen claves taxonómicas y fotos a color para identificar la mayoría de las especies discutidas. Palabras claves: Heteroptera, Pentatomoidea, biodiversidad, taxones INTRODUCTION Geography, Geology and Biota Puerto Rico is the smallest island among the ‘Greater Antilles’ in the West Indies, which include the larger islands of Cuba, Jamaica, and Hispaniola. The area of Puerto Rico, including all associated land- masses in its archipelago, is approximately 9,100 sq. km. Puerto Rico is located in the Caribbean Basin east of Hispaniola and west of the U.S. Virgin Islands just below the Tropic of Cancer, between latitudes 17º 55’ and 18º 32’ north and longitudes 65º 12’ and 67º 58’ west (Fig- ure 1). The largest islands that form part of Puerto Rico are: Vieques (135 sq. km), Culebra (28 sq. km), Mona Island (22 sq. km), Caja de Muerto (1.5 sq. km), and Desecheo (1.5 sq. km). Puerto Rico has great physiographic variability in its terrain, which is anchored by an East- West mountain range known as the Cordillera Central, punctuated by the highest elevation at Cerro Punta (1,330 m), and bordered by more than 500 km of coastline. This variability creates a multiplicity of geo- graphic and terrain features consistent with a wide range of climates supporting diverse and contrasting life zones that simultaneously in- clude subtropical dry forests and rain forests. Ancient land connections help explain significant patterns of ende- mism and extinction of insect species observed in Puerto Rico. As with other Greater Antilles, current day Puerto Rico was formed by indepen- dent terranes that were joined until the Miocene epoch (5 to 23 million years ago). Looking beyond its insular coastal boundaries, Puerto Rico is part of a larger geological platform called the Puerto Rican Bank, created by volcanism during the Cretaceous period, emerging largely through tectonic movements in the lower Eocene between 35 and 40 J. Agric. Univ. P.R. VOL. 100, NO. 3 DECEMBER 2016 3 million years ago. Today, the continuity of the original mountain axis of the Puerto Rican Bank is interrupted on the eastern end of Puerto Rico and reaches lower elevations through the islands of Culebra, Vie- ques, St. Thomas, St. John, Tortola, Virgin Gorda, Guana, Jost Van Dike, Saba, and Anegada. Furthermore, from the Miocene through the Pliocene epochs, Cuba, the Puerto Rican Bank, and eastern and central Hispaniola were part of the same microplate, and thus share many animal and plant taxa. According to Acevedo-Rodríguez and Strong (2012) the flora of the Puerto Rican Bank includes 3,243 plant spe- cies of which an estimated 13.6% appear to be endemic. Maldonado- Capriles (1996) found 5,066 insect species reported from Puerto Rico, and according to this author, the insect fauna of Puerto Rico is much better known than that of other Antillean islands. A General Account of Heteroptera Among insects with incomplete metamorphosis, true bugs (He- miptera: Heteroptera) are the largest and most diverse group. Heter- opterans show a remarkable ability to exploit food sources including plants, animals, and fungi, often in direct competition with humans. Heteropterans differ from other hemipteran suborders (i.e., Auchen- orrhyncha, Coleorrhyncha, and Sternorrhyncha) because their front wings are partly leathery and partly membranous (i.e., hence the name of “hemi-elytron”), and because their hind wings are wholly membranous. (Schaefer and Panizzi, 2000). These insects also differ from other sister suborders because of their well-developed scutellum, which is a shield-shaped modification to the thoracic dorsum, and for having abdominal and thoracic scent glands. Finally, although all other non-heteropteran groups are herbivorous, several heteropteran lines have evolved to feed on animal sources, even on warm-blooded vertebrates. Heteropterans are characterized by their elongate, straw-like mouthparts designed for piercing plant and/or animal tissues, and for sucking up their fluids for nourishment. While most species appear to be plant feeders, large numbers are animal feeders, some even feeding on humans (e.g., Reduviidae and Cimicidae). Another important dif- ference with other suborders is the way plant-feeding heteropterans obtain their food. While members of other suborders feed either within plant cells, or within phloem or xylem vessels, heteropterans (especially Pentatomomorpha) prefer to feed on reproductive plant parts: flowers, ovules, ovaries, and seeds. Other heteropterans, like the Cimicomor- pha (i.e., Tingidae and the Miridae ), damage non-reproductive plant parts. Thus, their feeding causes the plant to mobilize large quantities of nitrogen to make repairs (Schaefer and Panizzi, 2000). Both heter- 4 BIODIVERSITY OF HETEROPTERA IN PUERTO RICO: PART I opteran feeding-site preferences result in lower crop yields, and almost inevitably make many species important human competitors. Seven infraorders are recognized within the Heteroptera: Enico- cephalomorpha, Gerromorpha, Nepomorpha, Dipsocoromorpha, Lep- topodomorpha, Cimicomorpha, and Pentatomomorpha. They are all represented in the Puerto Rican fauna (See Schuh and Slater, 1995 for thorough descriptions of these taxa). Infraorder Pentatomomorpha Insects belonging to infraorder Pentatomomorpha are all terres- trial and the majority are plant feeders, consuming the fluids of many plant parts, especially flowers, seeds, and fruit. Except for the Ara- doidea, all other Pentatomomorpha are characterized by the presence of trichobothria: specialized slender sensory setae arising from spots and tubercles or pits on the abdominal venter (segments III-VII). Five superfamilies are currently recognized inside Pentatomomorpha (after Schuh and Slater, 1995), and all occur in Puerto Rico: Aradoidea (e.g., flat bugs), Pentatomoidea (e.g., stink bugs), Lygaeoidea (e.g., stilt, and seed bugs), Pyrrhocoridea (e.g., cotton stainers, and bordered plant bugs), and Coreoidea (e.g., leaf-footed bugs). Pentatomoidea According to an important study by Grazia et al. (2008), the Pentato- moidea is most likely a monophyletic group within Pentatomomorpha. Four key morphological characters distinguish this group of insects: (1) their scutellum reaches or surpasses an imaginary line crossing the connexivum at apical angles of 3rd abdominal segment; (2) the claval commissure is obsolete, with apices close together but not contiguous; (3) their abdominal trichobothria positioned laterally on urosternites II-VII; and (4) tergite VIII covering tergite IX in females. Sixteen fam- ilies are recognized within Pentatomoidea (sensu Schuh and Slater, 1995; and Grazia et al., 2008): Urostylidae, Saileriolidae, Plataspidae, Phoeidae, Parastrachiidae, Thaumastellidae, Dinidoridae, Canopi- dae, Lestoniidae, Acanthosomatidae,