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Coleoptera, Curculionidae, Entiminae) Machado_Pruebas 27/12/10 12:51 Página 233 Graellsia, 66(2): 233-280 julio-diciembre 2010 ISSN: 0367-5041 doi:10.3989/graellsia.2010.v66.025 LA MORFOLOGÍA DE LAPAROCERUS UNDATUS WOLLASTON, 1864 Y CONSIDERACIONES SOBRE LA TRIBU LAPAROCERINI LACORDAIRE, 1863 (COLEOPTERA, CURCULIONIDAE, ENTIMINAE) A. Machado* RESUMEN A. Machado. 2010. La morfología de Laparocerus undatus Wollaston, 1864 y consideracio- nes sobre la tribu Laparocerini Lacordaire, 1863 (Coleoptera, Curculionidae, Entiminae). Graellsia, 66(2): 233-280. Se presenta un estudio morfológico de Laparocerus undatus Wollaston, 1864 que com- prende la anatomía externa e interna del adulto, incluido el canal alimentario, sistema ner- vioso central y sistema reproductor. También se describen las anatomías de la larva y de la pupa, hasta ahora desconocidas en el género. Los Laparocerus son curculiónidos entimi- nos que muestran algunos caracteres primitivos infrecuentes en el grupo. Se plantea si se justifica o no mantener la tribu Laparocerini, pendiente de revisión desde que fuera esta- blecida por Lacordaire en 1863. Se discuten las semejanzas halladas en los caracteres ima- ginales y preimaginales con otros géneros y tribus, y se buscan las relaciones genéticas más cercanas empleando secuencias parciales del gen 16SrRNA de una veintena de espe- cies, obtenidas de GenBank. Laparocerus presenta importantes caracteres singulares o combinación de ellos, y se separa claramente de los demás Entiminae estudiados, ocupan- do una posición posiblemente basal dentro del grupo. Se proponen algunos caracteres diagnósticos para Laparocerini: presencia de mandíbulas con prolongación dehiscente (o su cicatriz), piezas bucales fanerognatas, escapo antenal largo, ojos y escrobas no dorsa- les, sutura entre el primer y segundo ventrito sinuosa al medio, éste tan largo como el ter- cero y cuarto reunidos; fémures sin dientes, tibias mucronadas con cestilla abierta, uñas soldadas y esternito VIII masculino completamente membranoso y provisto de un pequeño espículo relicto; en la larva coinciden las antenas de base oval con sensilo en forma de almohadilla y una mala con cinco setas en su cara ventral; la pupa carece de setas mandi- bulares. En función de estos y otros caracteres, se discuten los demás géneros atribuidos a Laparocerini, corroborando las exclusiones hechas por otros autores recientemente, y pro- poniendo la exclusión de Merimnetes, Neomerimnetes y Cyrtozemia. Quedan pendientes de confirmar si son Laparocerini dos géneros africanos: Asmaratrox y Straticus. Finalmente se discute en contexto biogeográfico la conveniencia de desglosar Laparocerus en una quincena de géneros independientes, o de mantenerlo unificado como un único género con varios subgéneros. Palabras clave: Anatomía; Curculionidae; Entiminae; Laparocerus; canal alimentario; 16SrRNA; sistema nervioso; larva; pupa; islas Canarias. * Calle Chopin 1, 38208 La Laguna, Tenerife, islas Canarias, España. [email protected] Machado_Pruebas 27/12/10 12:51 Página 234 234 Machado ABSTRACT A. Machado. 2010. The morphology of Laparocerus undatus Wollaston, 1864 and conside- rations on the tribe Laparocerini Lacordaire, 1863 (Coleoptera, Curculionidae, Entiminae). Graellsia, 66(2): 233-280 (in Spanish). This morphological study of Laparocerus undatus Wollaston, 1864 covers the external and internal anatomy of the imago, including the alimentary canal, the central nervous system and the reproductive system. The anatomy of the larva and pupa, unknown in this genus, is also described. Laparocerus are Entimin weevils that bear some primitive characters, not frequent in the group. The justification of a tribe Laparocerini, not revised since it was established by Lacordaire in 1863, is considered. Similarities of adult and preimaginal characters with those of other genera and tribes are discussed, and genetic relationships are researched using par- tial sequences of the 16SrRNA gen obtained from GenBank for some twenty tribe represen- tative species. Laparocerus has important singular characters or a combination of them and is clearly separate from the other Entiminae studied. It has possibly a basal phylogenetic position within the group. Some combined diagnostic characters for Laparocerini are proposed, in par- ticular: presence of a dehiscent mandible process (or its scar), phanerognatus mouth parts, long scape, eyes and antennal scroba not dorsal, suture of ventrites 1-2 arcuate at middle, ven- trite 2 as long as 3 and 4 together, femora unarmed, tibiae mucronate with open corbels, claws connate, and VIII sternite of male membranaceous bearing a short spiculum relictum; in the larvae, antennae are cushion shaped with an oval base and the mala has five ventral setae; the pupa lacks mandibular setae. These and other characters are used to evaluate the other gen- era traditionally assigned to Laparocerini. Recent exclusion of some of these genera by other authors is corroborated and the removal of Merimnetes, Neomerimnetes and Cyrtozemia is here proposed. Asmaratrox and Straticus from Africa remain as Laparocerini, pending confir- mation. Finally, the convenience of splitting Laparocerus into several genera or keeping it as a single genus with many subgenera is discussed in a biogeographical context. Keywords: Anatomy; Curculionidae; Entiminae; Alimentary canal; 16SrRNA; Nervous sys- tem; Larva; Pupa; Laparocerus; Canary Islands. Introducción Los “Laparocérides” (“Laparoceridi” in Bargagli, 1883) han sido posteriormente tratados como tribu Este estudio está inspirado en el homólogo de Laparocerini (e.g. Heyne & Taschenberg, 1907; Van den Berg (1972) dedicado a la morfología de Pierce, 1913; Lona, 1938; Alonso-Zarazaga & Lyal, Eremnus cerealis (Marshall, 1921), Curculionidae, 1999), como familia Laparoceridae por Heller Entiminiae, Tanyrhynchini. La ausencia de trabajos (1909) o como superfamilia Laparoceroidea por equivalentes en español justifica en parte este ensa- Ienistea (1986), sin que falten autores que no reco- yo, pero su objetivo principal es profundizar más en nocen el grupo, como Seidlitz (1868) que los reúne el conocimiento del género Laparocerus Schoenherr, con los “Otiorhynchides vrais”, o Winkler (1932) 1834, sobre el que el autor viene trabajando en los que asigna Laparocerus a los Mylacini. últimos años, y contribuir a esclarecer el concepto de Hoy sabemos que los caracteres que justificaron la tribu Laparocerini, a todas luces insatisfactorio. este agrupamiento inicial son homoplásicos. El pro- Lacordaire (1863), en su Histoire Naturelle des pio género Laparocerus, del que en aquellas fechas Insectes. Genera des Coléoptéres, establece los se conocía solo un par de especies, cuenta en la “Laparocérides” como grupo VII de los Otiorrhyn- actualidad con casi dos centenas, mostrando una chides, agrupando siete géneros: Elytrodon Bohe- plasticidad morfológica extraordinaria que confir- man, 1843; Laparocerus Schoenherr, 1834; Aomus ma la inutilidad de la mayoría de los caracteres Schoenherr, 1834; Aprepes Schoenherr, 1847; empleados por Lacordaire para establecer relacio- Pholicodes Schoenherr, 1826; Epiphaneus Schoen- nes de parentesco. herr, 1842 y Merimnetes Schoenherr, 1842. Poste- Lona (1938) excluye de Laparocerini a Elytrodon riormente, Faust (1890) añade a esta lista (Omiini), Aprepes (Phyllobiini), Pholicodes y Mitophorus Gerstaecker, 1855 y Edmundia Faust, Epiphaneus (Brachyderini), Edmundia (Sciaphilini) 1891 (=Reitteria Faust, 1890). y Mitophorus (Peritelini), a la vez que añade nuevos Graellsia, 66(2), Diciembre 2010, pp. 233-280 — ISSN: 0367-5041 doi:10.3989/graellsia.2010.v66.025 Machado_Pruebas 27/12/10 12:51 Página 235 Morfología de Laparocerus undatus 235 géneros a la tribu, quedando el grupo como sigue: ción real de Aphyonotus, Asmaratrox, Cyrtozemia, Aomus (Irán, Arabia Saudita y Siria) Laparocerus Merimnetes, Neomerimnetes y Straticus es inexis- (Macaronesia) y Merimnetes (Australia), más los tente o difícil de localizar. Suelen aparecer registra- añadidos Aphyonotus Faust 1895 (Etiopía), Asmara- dos como Laparocerini por defecto, o se omite su trox Heller, 1909 (Eritrea), Cyrtozemia Pascoe, 1872 adscripción tribal, seguramente a la espera de que (India), Lichenophagus, Wollaston 1854 (Macarone- alguien aclare el concepto de esta tribu. A diferen- sia), Neomerimnetes Blackburn, 1900 (Australia) y cia de Aomus, el aspecto externo de estos insectos, Straticus, Pascoe 1886 (Zaire); nueve géneros en sobre todo Merimnetes y Neomerimnetes, recuerda total. muy poco a los Laparocerus, pero ante la inusitada La acepción de Laparocerini más generalizada variación morfotípica que se da dentro de este en la actualidad es ésta de Lona, recogida directa- género, dicho criterio no es nada concluyente. No mente en el catálogo mundial de familias y géneros disponemos de material para poder abordar un estu- de Curculionoidea de Alonso-Zarazaga & Lyal dio comparativo; sin embargo, al ser Laparocerus (1999), donde la tribu Laparocerini aparece bajo la el genero tipo de los Laparocerini, un estudio mor- subfamilia Entiminae ¡junto a otras 54 más! Bien es fológico en profundidad como el que se presenta sabido que la sistemática de los curculiónidos en aquí puede ayudar a detectar caracteres singulares general, y la de los entiminos en particular (con del grupo o aquéllos otros que pudieran aportar más de 12.000 especies), dista mucho de reflejar un algún indicio sobre las relaciones de Laparocerus esquema natural (Oberprieler et al., 2007). con otros géneros conocidos. También se ensaya Los estudios detallados
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