SHILAP Revista de Lepidopterología ISSN: 0300-5267 [email protected] Sociedad Hispano-Luso-Americana de Lepidopterología España Kuznetzov, V. I.; Naumann, C. M.; Speidel, W.; Stekolnikov, A. A. The skeleton and musculature of male and female terminalia in Oenosandra boisduvalii Newman, 1856 and the phylogenetic position of the family Oenosandridae (Insecta: Lepidoptera) SHILAP Revista de Lepidopterología, vol. 32, núm. 128, diciembre, 2004, pp. 297-313 Sociedad Hispano-Luso-Americana de Lepidopterología Madrid, España Available in: http://www.redalyc.org/articulo.oa?id=45512810 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative 297 Kuznetzov (4-10-04) 3/1/77 18:27 Página 297 SHILAP Revta. lepid., 32 (128), 2004: 297-313 SRLPEF ISSN:0300-5267 The skeleton and musculature of male and female terminalia in Oenosandra boisduvalii Newman, 1856 and the phylogenetic position of the family Oenosandridae (Insecta: Lepidoptera) V. I. Kuznetzov, C. M. Naumann (†), W. Speidel & A. A. Stekolnikov Abstract The skeleton and musculature of the male and female terminalia have been studied in Oenosandra boisduvalii Newman, 1856, type species of the family Oenosandridae. It has been discovered that a comparatively high number of muscles of the male and female terminalia are included in the ground plan of ditrysian Lepidoptera. These are muscles m1, m3(2), m4, m5(7), m6(5), m7(6), m8(3), and m21. In the male genitalia only m2 (retractors of the anal cone) and m1-m7 and m10 in female terminalia have not been found. The position of muscles’ insertion is plesio- morphic in most cases. In Oenosandra, muscles m3(2) are retained and muscle m8(5) is in a plesiomorphic juxto- vincular position which is remarkably different to all other examined Noctuiformes. These two plesiomorphic featu- res and the general plesiomorphic nature of the male and female terminalia of the Oenosandridae are evidence of their basal position in the Noctuiformes. Reliable synapomorphic features of the Oenosandridae and other families of the Noctuiformes have not been found. Presumably, the similarities between the Oenosandridae and the Notodon- toidea or the Noctuoidea are a result of parallel evolution. KEY WORDS: Insecta, Lepidoptera, Oenosandridae, Notodontoidea, Noctuoidea, phylogeny. El esqueleto y la musculatura terminal del macho y de la hembra en Oenosandra boisduvalii Newman, 1856 y la posición filogenética de la familia Oenosandridae (Insecta: Lepidoptera) Resumen Se ha estudiado el esqueleto y la musculatura terminal en Oenosandra boisduvalii Newman, 1856, especie tipo de la familia Oenosandridae. Comparativamente se ha descubierto un elevado números de músculos terminales en el macho y la hembra que los incluye dentro del plan de los Lepidoptera ditrisianos. Son los músculos m1, m3(2), m4, m5(7), m6(5), m7(6), m8(3) y m21. En la genitalia del macho sólo el m2 (retractor del cono anal) y m1-m7 y m10 en la zona terminal de la hembra, no han sido encontrados. La posición de la inserción de los músculos es pleiso- mórfico en la mayor parte de los casos. En Oenosandra, se conserva el músculo m3(2) y el músculo m8(5) está en una posición pleisomórfica juxto-vincular lo que está en una posición destacadamente diferente a todos los otros Noctuiformes examinados. Estas dos características pleisomórficas y la naturaleza general pleisomórfica de los ter- minales de los machos y de las hembras de los Oenosandridae son una evidencia de su posición básica en los Noc- tuiformes. Las características sinapomórficas encontradas en los Oenosandridae y en otras familias de los Noctuifor- mes no han sido encontradas. Presumiblemente, las semejanzas entre los Oenosandridae y los Notodontoidea o los Noctuoidea son un resultado de evolución paralela. PALABRAS CLAVE: Insecta, Lepidoptera, Oenosandridae, Notodontoidea, Noctuoidea, filogenia. 297 297 Kuznetzov (4-10-04) 3/1/77 18:27 Página 298 V. I. KUZNETZOV, C. M. NAUMANN (†), W. SPEIDEL & A. A. STEKOLNIKOV Introduction Recent investigations of Lepidoptera morphology (KRISTENSEN, 1999) have discovered small groups within speciose taxa whose systematic position is questionable. Some of these groups have a nodal position in the lepidopterous system (Agathiphagidae, Heterobathmiidae, Hedylidae and so on). One of these groups is the small Australian family Oenosandridae. The genera Oenosandra Newman, 1856 and Discophlebia Felder, 1874 were considered by taxonomists as Thaumetopoeinae or Notodon- tinae (TURNER, 1903, 1922; KIRIAKOFF, 1970; COMMON, 1970, 1990). MILLER (1991) extracted these genera from the family Notodontidae and erected the family Oenosandridae. He placed the Oeno- sandridae at the base of the notodontoid branch after a cladistic analysis and suggested that this taxon was the sister group of the clade Doa + Notodontidae. At the same time, MILLER (loc. cit.) noted a conflicting set of characters in the type species Oenosandra boisduvalii. Therefore, MILLER’S hypot- hesis of the phylogenetic position of the family is problematic. MILLER (loc. cit.) reported that the me- tathoracic tympanum is similar to that in Lymantriidae and Arctiidae, but not to the Notodontidae. La- ter, the hypothesis concerning the relationships between the Oenosandridae and the Noctuoidea s. l. (Noctuiformes according to KUZNETZOV & STEKOLNIKOV, 2001) was discussed and the Oenosan- dridae were tentatively given a basal position within that superfamily (SPEIDEL, FÄNGER & NAU- MANN, 1996). KITCHING & RAWLINS (1999) also considered the Oenosandridae to be the sister group of the remaining Noctuoidea s. l. (Noctuoidea + Notodontoidea). In a recent catalogue, ED- WARDS (in NIELSEN et al., 1996) included the genera Diceratucha Swinhoe, 1904 (1 species), Dis- cophlebia (5 species), Nycteropa Turner, 1941 (1 species), and Oenosandra (1 species) within the fa- mily Oenosandridae. So, there are two hypotheses on the position of the Oenosandridae in the classification of the Lepi- doptera: The family is opposed to the notodontoid branch only (MILLER, loc. cit.) or to all other Noc- tuoidea s. l. (Noctuiformes) (SPEIDEL, FÄNGER & NAUMANN, 1996; KITCHING & RAWLINS, 1999). For the solution of this phylogenetic problem, the functional organisation of the terminalia of both sexes of O. boisduvalii Newman, 1856 has been examined in details. In the previous literature the- re is only a sketchy drawing of the male and female genitalia of O. boisduvalii (KIRIAKOFF, 1970) and fragmentary data on the male musculature of the same species (SPEIDEL, FÄNGER & NAU- MANN, 1996). Materials and methods The insects were collected by Dr. E. D. Edwards in Australia. The terminalia were fixed in ethanol and prepared in a Petri dish to study the position of the muscles. The end of the abdomen was dissected sagittally with microsurgical scissors. Each half of the abdomen was attached to a wax layer in the Petri dish bottom. Internals, adipose tissues, and tracheas were removed with forceps. Pointed dissecting ne- edles and microsurgical scissors were used for manipulation and dissection. In total, three males and one female were investigated. Nomenclature Two systems of numerical nomenclature are used in morphological publications on the musculatu- re of lepidopterous male genitalia. In American and Western European papers, Forbes’ nomenclature (FORBES, 1939) is employed. At the same time, scientists from Eastern European countries and Russia number the muscles according to KUZNETZOV & STEKOLNIKOV (1973). The two parallel nomen- clatures makes it difficult to compare data reported by different authors. Replacement of one nomencla- ture by the other one does not help understand previous publications. Here, we make use of a recently introduced compromise nomenclature (SPEIDEL, FÄNGER & NAUMANN, 1996). Muscles are given two numbers: the first number is according to FORBES (1939), and the number in brackets is accor- ding to KUZNETZOV & STEKOLNIKOV (1973). Only one number is used if the muscle’s number is the same in both nomenclatures. The two nomenclatures are compared in Table 1. Recently, IVANOV 298 SHILAP Revta. lepid., 32 (128), 2004 297 Kuznetzov (4-10-04) 3/1/77 18:27 Página 299 THE SKELETON AND MUSCULATURE OF MALE AND FEMALE TERMINALIA IN OENOSANDRA BOISDUVALII NEWMAN, 1856 (2003) reconstructed the ground plan of the male terminalia of the Amphiesmenoptera (Trichoptera + Lepidoptera) and proposed a nomenclature using the abbreviations of KUZNETZOV & STEKOLNI- KOV (1987, 2001) for both orders. Table 1. The nomenclature of male genital muscles in the publications of the different authors. Forbes, 1939 Kuznetzov & Speidel, Fänger & Function of muscles Stekolnikov (1973) Naumann (1996) m1 m1 m1 Depressors of uncus m2 m10 Retractors of anal tube m3 m2 m3(2) Abductors of the valves m4 m4 m4 Adductors of the valves m5 m7 m5(7) Flexors of valves - intervalval muscles (flexors of clasper according to FORBES, 1939) m6 m5 m6(5) Retractors of aedeagus m7 m6 m7(6) Protractors of aedeagus m8 m3 m8(3) Ventral adductors of the valves - - m21 Retractors of vesica Muscles of the female terminalia are numbered according to the system proposed in Zygaena trifolii (Esper, 1783) (FÄNGER & NAUMANN, 1997). Results Description of the male terminalia 1. Pregenital segment (viii) (Figs 1, 2) The male genitalia are retracted into segment viii; tergite viii is a small plate, which