Pronozelus, a New Neotropical Harpactorine Genus and Species from Colombia (Hemiptera: Heteroptera: Reduviidae: Harpactorinae) Author(s): Dimitri Forero Source: Entomologica Americana, 118(1):278-284. 2012. Published By: The New York Entomological Society DOI: http://dx.doi.org/10.1664/12-RA-021.1 URL: http://www.bioone.org/doi/full/10.1664/12-RA-021.1

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Entomologica Americana 118(1–4):278–284, 2012

PRONOZELUS, A NEW NEOTROPICAL HARPACTORINE GENUS AND SPECIES FROM COLOMBIA (HEMIPTERA: HETEROPTERA: REDUVIIDAE: HARPACTORINAE) DIMITRI FORERO Department of Entomology, University of California Riverside, Riverside, CA 92521, USA Present address: Departamento de Biologı´a, Laboratorio de Entomologı´a (106B), Ed. 53-54, Pontificia Universidad Javeriana, Carrera 7 #43-82, Bogota´, Colombia. E-mail: [email protected] Abstract—Pronozelus schuhi gen. nov. and sp. nov. of Harpactorini is described from the eastern Colombian Andes. A dorsal habitus, photographs of male genitalia, and a key to distinguish Pronozelus from the most similar genera are provided. Key words: Andes, Harpactorinae, Harpactorini, new species, South America, taxonomy.

INTRODUCTION Harpactorinae are the largest subfamily of Reduviidae (Maldonado, 1990). Unsurprisingly, the tribal level classification of this subfamily is unsettled and still represents a challenge (Forero, 2011). Only two tribes occur in the New World: Apiomerini and Harpactorini (Davis, 1969). Recent phylogenetic analyses support the monophyly of Apiomerini (Forero et al., in press; Forero and Weirauch, in prep.) but not of Harpactorini, which is paraphyletic with respect to Rhaphidosomini (Weirauch and Munro, 2009). Among all the groups within Harpactorinae, Harpactorini is the largest with more than 2,000 species, more than 280 described genera worldwide (Maldonado, 1990), and 51 genera in the Neotropical region (Forero et al., 2008). Despite continuous taxonomic descriptive efforts in the New World (Forero et al., 2008, and references therein), seldom explored areas in the Neotropical Region, such as the Colombian Andes, provide evidence that additional fieldwork—even in relatively disturbed areas—might reveal new taxa. In this paper I describe a new genus and species of Neotropical Harpactorini from Colombia, provide documentation of its genitalia in the form of photographs, discuss its potential phylogenetic placement, and compare it with the putative closest relatives. MATERIAL AND METHODS The holotype male, the only examined specimen, is deposited in the Museo Entomolo´gico,

Facultad de Agronomı´a, Universidad Nacional de Colombia, Bogota´ (UNAB). Labels were copied verbatim. Digital photographs were taken with a Microptics-USA photomicrographic system (www. visionarydigital.com), equipped with infinity K2 lens and a CF-3 and CF-4 objectives. A Zeiss Axioscope 2 compound microscope furnished with a 53 CP-Achromat objective and a GTVision digital camera were used for the digital photographs of the genitalia. Stacks of images were digitally processed with CombineZP (Hadley, 2010) and further processed in Photoshop CS4 version 11.0. Measurements are in millimeters, except as noted. Terminology of external characters follows Forero, et al. (2008), and genitalic terms follow Forero and Weirauch (2012). The holotype was affixed with a Unique Specimen Identifier (USI) number in the form of a barcode label. KEY TO NEOTROPICAL HARPACTORINI GENERA RELATED TO PRONOZELUS GEN. NOV. This is a key to facilitate the separation of Neotropical Harpactorini genera with delicate bodies, long legs, antenniferous tubercle flat without spines, and mesepisternum without a tubercle. 1- Male without parameres . . . . . . . . . . . Atopozelus 19- Male with parameres . . . . . . . . . . . . . . . . . . . . 2 2- Abdominal margin expanded at segments 6 and 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iquitozelus 29- Abdominal margin nearly parallel, continuous, never abruptly expanded . . . . . . . . . . . . . . . . . . 3 3- Forelegs very long, length/width ratio of profemur 44 or more; 1st visible labial segment

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Fig. 1. Pronozelus schuhi gen. nov. et sp. nov. A. Dorsal habitus. B. Detail of head and pronotum, dorsal view. C. Lateral view of the head. D. Detail of head and pronotum in lateral view.

not reaching anterior margin of eye; posterior margin of the pronotum nearly straight, humeral angles not spined . . . . . . . . Ischnoclopius 39- Forelegs not as long, length/width ratio of profemur equal or less than 30; 1st visible labial segment at least reaching anterior margin of eye; posterior margin of the pronotum angled or slightly curved, humeral angles spined or not . . . . 4 4- Lateral angles of the pronotum not greatly expanded, if slightly expanded, then posterior margin of pronotum not gently curved but angled; anterior lobe of pronotum at most half as long as posterior lobe; mesosternum nearly flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zelus 49- Lateral angles of the pronotum greatly expanded; anterior lobe of pronotum much less (0.44) than half the length of posterior lobe; mesosternum slightly enlarged . . . Pronozelus gen. nov.

PRONOZELUS GEN. NOV. Figures 1, 2 TYPE SPECIES: Pronozelus schuhi, sp. nov. DIAGNOSIS: Distinguished among all Neotropical Harpactorini by the following combination of characters: antennal tubercles unarmed (Fig. 1B, C); long and delicate legs, in particular the profemur with a length/width ratio of 26 (Fig. 1A); metafemur about as long as the profemur; mesepisternum flat, without any process or tubercle (Fig. 1D); pronotum gently curved along the posterior margin from the humeral angles, covering the base of the scutellum (Fig. 1B); humeral angles greatly expanded laterally (Fig. 1B); posterior lobe of pronotum more than two times (2.23) as long as anterior lobe; posterior margin of the

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Fig. 2. Pronozelus schuhi gen. nov. et sp. nov. A–E. Pygophore and 8th abdominal segment. A. Dorsal view. B. Lateral view. C. Caudal view. D. Lateral view of apex of abdomen. E. Caudal view of pygophore in situ. F–H. Phallus with inflated endosoma. F. Dorsal view. G. Lateral view. H. Ventral view. Abbreviations: emar 5 emargination of posterior margin of pygophore; mp 5 medial process; proc 5 triangular process of pygophore; str 5 struts; ds 5 dorsomedial sclerites of endosoma.

pygophore laterad to the medial process of the pygophore deeply emarginated (emar) (Fig. 2B, C, E); medial process (mp) of the pygophore undivided (Fig. 2A–E); parameres small (Fig. 2A); paramere socket covered laterally by a large triangular process (proc) (Fig. 2B). DESCRIPTION: Male: COLORATION (Fig. 1A): Reddish brown, with orange and yellow areas. VESTITURE: Head: Pedicel on lateral distal half with 7 trichobothria arranged in a slightly curved single row. STRUCTURE: Head (Fig. 1B, C): Anteocular region slightly shorter than postocular; anteocular gently narrowing towards the apex, postocular strongly convex to midpoint posteriorly forming a narrow collum; eye nearly hemispherical, not reaching dorsal or ventral margin of head in lateral view, slightly protruding laterally in dorsal view; postocular sulcus deep, curved, situated at level of posterior margin of eye; ocellus relatively large, equidistant to eye and other ocellus, located on large tubercle; antenniferous tubercle with cuticle flat, without spines. Second labial segment (1st visible) short, nearly

reaching anterior margin of eye; third (2nd visible) longest, straight; fourth (3rd visible) shorter than second; ratio 1: 2.24: 0.53. Scape about two times as long as head length. Thorax: Anterior lobe of pronotum about half as long as posterior lobe and about half as wide; discal ridges wide, vaguely impressed; anterolateral angles very short, blunt; longitudinal sulcus deeply impressed in posterior end; transverse sulcus impressed (Fig. 1B). Paired longitudinal, posteriorly diverging low ridges arising just anteriorly to transverse sulcus reaching into posterior lobe. Posterior lobe of pronotum densely set with pits, surface rugose, carinate on lateral margin (Fig. 1D), gently curved anteriorly, broadly curved on posterior margin, disc elevated (Fig. 1B); humeral angles produced and spined (Fig. 1B). Scutellum with disc elevated, triangular; base covered by posterior margin of pronotum. Meso- and metapleura without tubercles. Mesosternum slightly swollen as paired lobes (Fig. 1D), posterior margin with sharp, U-shaped carina. Hemelytron: Corium with veins raised, surface with dense, very shallow, darkened pits

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where setae are inserted, apex surpassing middle of membrane; membrane about as long as corium. Legs: Long and delicate. Profemur longer than mesofemur, and slightly longer than metafemur, ratio 1: 0.73: 0.93; profemur length/width ratio 26. Protibia with prominent tibial spur. Abdomen: Margin of connexival segments entire, nearly parallel, barely visible in dorsal view. 7th seternite larger than preceding segments. Genitalia: Segment 8 sclerotized only on ventral half (Fig. 2B), anterior margin gently emarginate, posterior margin straight; spiracle on membrane (Fig. 2B). Pygophore ovoid, anterior opening separated from genital opening by well-sclerotized transverse bridge (Fig. 2A); posterior margin of genital opening deeply emarginated, emargination round, surrounding parameres (Fig. 2B–E); medial process of pygophore triangular, broad basally (Fig. 2A, E), directed posteriorly at about 45u (Fig. 2B), apex of medial process with small, acute, ventrad process; paramere short, slightly surpassing posterior margin of pygophore, nearly straight, apical half slightly enlarged (Fig. 2A, B); paramere socket covered laterally by large triangular process with rounded apex, process broadly rounded in dorsal view and nearly reaching posterior margin of genital opening (Fig. 2A, B). Aedeagus ovoid when not inflated. Basal plate with arms narrow, expanded dorsally, strongly sinuous in anterior view (Fig. 2H); basal plate bridge not sclerotized. Basal plate extension very short, slightly expanded laterally and flattened dorsoventrally (Fig. 2G, H). Phallosoma with dorsal phallothecal sclerite short, transversely oblong (Fig. 2F). Endosoma (partially inflated) without protruding lobes on proximal half; distal half with medial paired dorsal sclerites and dense areas of microtrichia (Figs. F, G); endosomal struts (str) apically divergent (Fig. 2F). Female: Unknown. ETYMOLOGY: From the Greek ‘‘pronos,’’ protuberance, and the reduviid generic name Zelus, to indicate the great lateral humeral expansions and its relatedness to Zelus. The gender is masculine. Pronozelus schuhi sp. nov. Figures 1, 2 HOLOTYPE: Male: COLOMBIA: Cundinamarca: 431 m, 4u389540N, 73u199370W, 16 Oct 1999, C. Corte´s/Medina (Cund.), finca Mi Negro, 16-1099, C.Corte´s/UCR_ENT 00045992/= Pronozelus schuhi sp. nov. Forero (UNAB).

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DESCRIPTION: Male: MEASUREMENTS: Total length: 13.1; head length: 2.8; head width across eyes: 1.12; anteocular length: 0.85, postocular length: 1.35; scape: 5.9, pedicel: 1.52; labial segment II (1st visible): 0.75, III (2nd visible): 1.68, IV (3rd visible): 0.4; length of anterior pronotal lobe: 0.85, width: 1.6; length of posterior pronotal lobe: 1.9, width (across humeral angles including spines): 4.25; length of profemur: 6.5, width: 0.25; length of protibia: 6.65; length of mesofemur: 4.75; length of mesotibia: 5.9; length of metafemur: 6.1; length of metatibia: 6.9. COLORATION (Fig. 1): Head: Pale yellow; frons with paired longitudinal, pale brown stripes going from the antennal tubercles to transverse sulcus of head (Fig. 1B); area anterior to each ocellus pale brown; anteocular region laterally with brown stripe from antennal tubercle to middle of eye (Fig. 1C). Thorax: Pronotum and scutellum orange brown (Fig. 1B); thoracic pleura and sternum orange red (Fig. 1D). Hemelytron: Clavus and most of corium orange brown; corium on area adjacent to quadrate and at base of discal cell black, distal-most portion of corium orange; membrane dark brown (Fig. 1A). Legs: Fore, mid, and hindleg with coxae and trochanters orange red (Fig. 1A, D); femora and tibiae pale yellow; tarsi darkened (Fig. 1A). Abdomen: Orange red, becoming bright red towards the apex; mediotergite VII black. Pygophore red, parameres black. VESTITURE: Head: Very short, delicate, sparse, apically recurved setae. Thorax: Small, erect, delicate setae intermixed with sparse apically recurved setae on pleura; anterior lobe of pronotum with sparse, darkened setae located only on elevated ridges; posterior lobe of pronotum with dense, very short, apically recurved, darkened setae. Hemelytron: Clavus and corium with very short, delicate, apically recurved, simple setae. Legs: Protibia with short, stout, erect setae, meso- and metatibiae with simple, delicate, erect setae, less dense in metatibia. Abdomen: Very short, sparse, simple setae. Genitalia: Base of medial process of the pygophore laterally densely setose (Fig. 2C, E); paramere apically with sparse, long setae (Fig. 2A, B). STRUCTURE: Head, thorax and abdomen as in generic description. GENITALIA (Fig. 2): As in generic description except as follows. Dorsal phallothecal sclerite with posterior margin strongly emarginate as a pair of broad rounded lobes (Fig. 2F), directed downwards (Fig. 2G). Endosoma with a pair of dorsomedial sclerites on distal half (ds), broadly triangular, apex

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of sclerites slightly converging medially (Fig. 2F), membrane adjacent to apex of sclerites with broad, triangular heavily sclerotized denticles (Fig. 2F); membranous lobes lateral to medial sclerites with dense, not well-sclerotized, triangular spines; distal central lobe posterior to paired sclerites with central area with dense area of long, hair-like microtrichia (Fig. 2F); distal lateroventral areas with paired, broad slightly sclerotized areas (Fig. 2F, H). Female: Unknown. ETYMOLOGY: This new species is named after Randall T. Schuh in honor of his 70th birthday. This is a small token of appreciation for, and in recognition of, his productive taxonomic work on various groups of Heteroptera, particularly Miridae and Saldidae, as well as on his seminal influence on the phylogenetics of Heteroptera. DISTRIBUTION: Only known from the type locality in the eastern foothills of the eastern cordillera of the Colombian Andes. DISCUSSION: Pronozelus shares a number of characters with a group of closely related genera in the New World. The structure of the male genitalia suggest a close relationship among Pronozelus and Atopozelus Elkins (Elkins, 1954; Hart, 1972a), Ischnoclopius Sta˚l (Hart, 1975), and Zelus Fabricius (Hart, 1986, 1987). Iquitozelus Be´renger is a genus related to this group, but only females are known (see below) (Be´renger, 2003). Atopozelus, Ischnoclopius, and Zelus all share a strongly emarginate posterior border of the pygophore, as well as a general delicate appearance, long and delicate legs, long antennae, and antennal tubercles without spines. Pronozelus can be easily separated from Atopozelus because of the lack of parameres in the latter. Of the remaining genera, Ischnoclopius has very short parameres, and also very long profemora with a length/width ratio of about 44 (Hart, 1975), whereas in Pronozelus it is about 26, and in Zelus it ranges from about 13 to nearly 30 (Hart, 1972b). Ischnoclopius has the pronotum flattened and the humeral angles not expanded. Atopozelus, Zelus and Ischnoclopius all lack large prominences lateral to the paramere sockets. Pronozelus is more closely related to Zelus with regard to male genitalic characters (see below). Given that there is no phylogenetic hypothesis for Zelus, no synapomorphies are yet available that may facilitate comparisons with this new taxon. One of the outcomes of future cladistic analyses may

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be to have a paraphyletic Zelus with respect to Pronozelus. Until those analyses are produced, I present evidence (based on the available data) that supports the proposal of this taxon as a new genus when compared with several species of Zelus. Furthermore, there is no combination of characters that unambiguously allow diagnosing Zelus, being identified by its lack of autapomorphies (Hart, 1972a, b, 1986, 1987). The endosoma is routinely not inflated in Reduviidae species descriptions, although it has useful characters for taxa delimitation (e.g., Ishikawa and Okajima, 2006; Ishikawa et al., 2006; Redei and Ishikawa, 2007), including certain harpactorine groups (Forero and Weirauch, 2012; Wygodzinsky, 1947a, 1948; Forero et al., 2008; Berniker et al., 2011). Pronozelus can be differentiated from Zelus because the dorsal phallothecal sclerite is shorter than any of the known species of Zelus, and in those species with an emarginate posterior margin, the resulting lobes are not as broad as in Pronozelus. Some described species of Zelus, like Z. ambulans Sta˚l and Z. luridus Sta˚l, have the medial process of the pygophore wide at base, but only in a few undescribed species of Zelus, the base of the medial process of the pygophore is relatively wide and has an emarginate posterior border of the dorsal phallothecal sclerite (Hart, 1972b), similar to the structure found in Pronozelus. Nonetheless, the expansion of the humeral angles in Pronozelus is much greater than in any of the known species of Zelus. In these undescribed species, the humeral angles are barely produced beyond the margin of the thorax. Only in Z. armillatus (Lepeletier et Serville) are the humeral angles slightly more expanded than other species and spined, but the posterior margin of the pronotum is angled lateral to the scutellum, not gently curved as in Pronozelus. In all these species, the anterior lobe of the pronotum is about half as long as the posterior lobe, whereas in Pronozelus is less (0.44). Furthermore, Pronozelus has a pair of dorsomedial sclerites on the posterior half of the endosoma, in addition to dense areas of microtrichia (Figs. 2F, G). Other Neotropical Harpactorini have large paired endosomal sclerites, such as Aristathlus Bergroth (Forero et al., 2008), although in Pronozelus these sclerites are not as large. The endosomal structure has remained unknown in Atopozelus and Ischnoclopius, and virtually undocumented for Zelus. Hart (1972b, 1986, 1987)

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never documented any endosomal sclerites in any Zelus species because the phallus was never inflated. The endosoma has been illustrated only for Z. illotus Berg (as Z. carvalhoi Wygodzinsky) and Z. mattogrossensis Wygodzinsky (Wygodzinsky, 1947b). In these two species the endosoma seems to be either completely membranous (Z. illotus) or with weakly sclerotized processes (Z. mattogrossensis). Thus, the particular structure of the endosoma might reveal additional characters to clarify the relationships among Atopozelus, Ischnoclopius, Zelus, and Pronozelus. Iquitozelus, a genus described from the Amazon Region in Peru, is only known from females (Be´renger, 2003; Forero, 2006). Iquitozelus is obviously related to Atopozelus, Ischnoclopius, and Zelus (Be´renger, 2003). Iquitozelus shares with Pronozelus a similar head structure, slightly rounded posterior margin of the pronotum, and delicate legs. The only known specimens of Iquitozelus (I. couturieri holotype total length 5 16 mm) are much larger than the male of Pronozelus (13 mm), a difference too large to be attributed to sexual dimorphism. In addition, the expanded connexivum on segments 6 and 7 of Iquitozelus readily separates this taxon from Pronozelus. Other differences include the blunt anterolateral angles of the pronotum in Pronozelus, whereas Iquitozelus has small conical tubercles on top of them (Be´renger, 2003). The length/width ratio of the profemur is also different, which in Iquitozelus is about 13, whereas in Pronozelus it is 26. The postocular region in dorsal view is slightly wider in Iquitozelus when compared with Pronozelus. Finally, the humeral angles are much more developed laterally and are armed with stouter spines in Pronozelus, whereas in Iquitozelus the humeral angles barely project beyond the pleural region and the spines are more delicate (Be´renger, 2003). Zhang and Weirauch (in press) recently documented the presence of sticky glands on a number of Neotropical Harpactorini taxa. Given that the holotype is the only specimen of Pronozelus, no attempt was done to study them. Nonetheless, because the forelegs are covered in a resinous substance (Fig. 1A), and given that the protibiae have erect setae that can be interpreted as sundew setae, which are always associated with sticky glands (Zhang and Weirauch, in press), it is likely that Pronozelus also possesses sticky glands. Furthermore Atopozelus, Ischnoclopius and Zelus,

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three of the putative closest relatives of Pronozelus, all possess sticky glands (Zhang and Weirauch, in press). Additional specimens of Pronozelus—in particular females—together with additional documentation on the endosoma of Atopozelus, Ischnoclopius, and Zelus, and males of Iquitozelus, will provide further characters to elucidate their relationships. ACKNOWLEDGEMENTS I am grateful to Francisco Serna and Erika Vergara (UNAB) for arranging the loan of the specimen. Guanyang Zhang (University of California Riverside) and an anonymous reviewer kindly provided suggestions to improve the manuscript. Thomas J. Henry provided editorial recommendations. LITERATURE CITED Be´renger, J. M. 2003. Iquitozelus couturieri, nouveau genre et nouvelle espe`ce d’Harpactorinae du Pe´rou (Heteroptera, Reduviidae). Nouvelle Revue d’Entomologie (N. S.) 20: 23–27. Berniker, L., S. Szerlip, D. Forero and C. Weirauch. 2011. Revision of the crassipes and pictipes species groups of Apiomerus Hahn (Hemiptera: Reduviidae: Harpactorinae). Zootaxa 2949: 1–113. Davis, N. T. 1969. Contribution to the morphology and phylogeny of the Reduvioidea. Part IV. The Harpactoroid complex. Annals of the Entomological Society of America 62: 74–94. Elkins, J. C. 1954. A new American harpactorine genus (Hemiptera, Reduviidae). Texas Reports on Biology and Medicine 12: 39–48. Forero, D. 2006. New records of Reduviidae (Hemiptera: Heteroptera) from Colombia and other Neotropical countries. Zootaxa 1107: 1–47. Forero, D. 2011. Classification of Harpactorinae assassin bugs (Hemiptera: Heteroptera: Reduviidae). Boletı´n del Museo Entomolo´gico Francisco Luis Gallego 3: 9–24. Forero, D., L. Berniker and C. Weirauch. In press. Phylogeny and character evolution in the beeassassins (Insecta: Heteroptera: Reduviidae). Molecular Phylogenetics and Evolution. Forero, D., H. R. Gil-Santana and P. H. van Doesburg. 2008. Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae), pp. 85–103 in S. Grozeva and N. Simov (eds.), Advances in Heteroptera Research: Festschrift in Honor of 80th Anniversary of Michail Josifov. Pensoft, Sofia-Moscow. Forero, D. and C. Weirauch. 2012. Comparative genitalic morphology in the New World resin bugs Apiomerini (Hemiptera, Heteroptera, Reduviidae,

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Harpactorinae). Deutsche Entomologische Zeitschrift 59: 5–41. Hadley, A. 2010. CombineZP. Available at: http://www. hadleyweb.pwp.blueyonder.co.uk Hart, E. R. 1972a. A new species of the genus Atopozelus Elkins, with a review of the genus. Journal of the Kansas Entomological Society 45: 392–396. Hart, E. R. 1972b. A systematic revision of the genus Zelus Fabricius (Hemiptera: Reduviidae). Ph.D. dissertation (Entomology), Texas A&M University, College Station. Hart, E. R. 1975. A new species of Ischnoclopius Sta˚l, with notes on the systematic position of the genus (Hemiptera: Reduviidae). Proceedings of the Entomological Society of Washington 77: 419–425. Hart, E. R. 1986. Genus Zelus Fabricius in the United States, Canada, and Northern Mexico (Hemiptera: Reduviidae). Annals of the Entomological Society of America 79: 535–548. Hart, E. R. 1987. The genus Zelus Fabricius in the West Indies (Hemiptera: Reduviidae). Annals of the Entomological Society of America 80: 293–305. Ishikawa, T., W. Cai and M. Tomokuni. 2006. A new species of Oncocephalus (Heteroptera, Reduviidae, Stenopodainae) from Japan. Denisia 19: 475–481. Ishikawa, T. and S. Okajima. 2006. The assassin bug genus Emesopsis (Heteroptera, Reduviidae, Emesinae) in Thailand. Denisia 19: 457–474.

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Maldonado, J. 1990. Systematic catalogue of the Reduviidae of the World. Caribbean Journal of Science, Special publication No. 1, University of Puerto Rico, Mayagu¨ez. Redei, D. and T. Ishikawa. 2007. The generic placement of Endochus stalianus Horvath, 1879 (Heteroptera, Reduviidae, Harpactorinae), with proposal of a new synonymy. Biogeography 9: 1–5. Weirauch, C. and J. B. Munro. 2009. Molecular phylogeny of the assassin bugs (Hemiptera: Reduviidae), based on mitochondrial and nuclear ribosomal genes. Molecular Phylogenetics and Evolution 53: 287–299. Wygodzinsky, P. 1947a. Contribuic¸a˜o ao conhecimento do geˆnero Heniartes Spinola, 1837 (Apiomerinae, Reduviidae, Hemiptera). Arquivos do Museu Nacional 4: 3–65. Wygodzinsky, P. 1947b. Soˆbre alguns ‘‘Reduviidae’’ do Brasil central (Hemiptera). Revista Brasileira de Biolgia 7: 423–434. Wygodzinsky, P. 1948. Sobre alguns Reduviidae da Regia˜o Amazoˆnica (Hemiptera). Revista de Entomologia 10: 557–564. Zhang, G. and C. Weirauch. In press. Sticky predators: a comparative study of sticky glands in harpactorine assassin bugs (Insecta: Hemiptera: Reduviidae). Acta Zoologica. DOI: 10.1111/j.14636395.2011.00522.x

Pronozelus, a New Neotropical Harpactorine Genus ...

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and ... Department of Entomology, University of California Riverside, Riverside, CA 92521, USA ..... sity, College Station. Hart, E. R. 1975 ...

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Feb 28, 2011 - Ono-H. Paratypes: same data as holotype (1 ♂, 2♀♀, 1 sex?); Nemoto, Shirahama-machi, Minamibôsô-shi,. Chiba-ken, 7 XII 2007, Ono-H. (1 ...

New Record of the Seashore Genus Salinamexus ...
Feb 28, 2011 - The seashore rove beetle genus Salinamexus is recorded from Japan for the first time, and the following species are ... Key words: taxonomy, Coleoptera, Staphylinidae, new record, coarse sand beach, gravel beach,. Salinamexus browni ..

genus Eothenomys - Semantic Scholar
... 49.69 50.37 50.38 49.39 42.34 39.07 39.17 40.21 39.58 38.71 40.52 37.48 40.61 40.53 42.45 39.92 47.08 55.48 43.62 55.62 40.22 35.47 35.61 36.54. 360. J.

Two New Species of Centrolenella, with a Brief Review of the Genus ...
Mar 20, 2008 - http://www.jstor.org/journals/herpetologists.html. Each copy of ... Two species are named as new--C. pluuialis and C. phenax, both from cloud.

Neotropical region: a shapefile of Morrone's (2014 ... - Biotaxa
May 26, 2014 - Environmental Systems Resource Institute (1998) Shapefile Technical Description. Redlands, CA, 30p. Environmental Systems Resource ...

genus Eothenomys - Semantic Scholar
summarized all available karyotype data and discussed ..... Clethrionomyini and 9 outgroup taxa are shown in (A,B); analysis involving ingroup taxa sequences only are shown in (C,D). ..... (2001CB711103), and the National Natural Science.

Chirleja buckii, a new genus and species of lichenized ...
conducted a 500-replicate rapid bootstrap ana- lysis (the selection of the number ..... independent, community-supported software for describing and comparing ...

A KEY TO THE NEOTROPICAL GENERA OF ...
Central XI, San Lorenzo, PARAGUAY. e-mail: [email protected]. Abstract.- A key to .... más las islas del Caribe, esto es, la misma área cubierta por el ...

of Neotropical Montane Forests
Jun 21, 1993 - pasturelands for the growing industry of cattle ranching. The current ..... tion in elfin cloud forests in Colombia and Venezuela. J. Trop. Ecol.