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 (Insecta: ) SHILAP Revista de Lepidopterología, vol. 32, núm. 128, diciembre, 2004, pp. 297-313 Sociedad Hispano-Luso-Americana de Lepidopterología Madrid, España

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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 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.

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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 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 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

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(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 is narrower caudal. The sternite is broader, with slightly sclerotized mediocaudal part. A large pleural membrane connects tergite and sternite of the last pregenital segment. Sternites of visceral segments, except viii, are slightly sclerotized and weakly developed; a pale yellow pigmentation is present in the ventral parts of the segments only. Three pairs of muscles extend from tergite viii to segment ix (Fig. 1): two pairs of tergal longitudi- nal intersegmental muscles (mt. 8-9) and one pair tergosternal intersegmental muscles (mt-s. 8-9). Both pairs of tergal longitudinal muscles extend paramedially from the anterior margin of segment viii (Fig. 2). The more slender mt. 8-9 (1) extends medially backwards to the anterior margin of the tegumen. Irregular muscles (m) are parallel to mt. 8-9 (1); they extend caudad from the anterior phragma of seg- ment viii to the fine intersegmental membrane. The larger and thicker mt. 8-9 (2) extend (laterally with respect to mt. 8-9 (1)) from the anterior margin of tergite viii to the anterior margin of the tegumen, in its lower part. The mt. 8-9 (2) is attached to the tight intersegmental membrane adjoining the tegumen. The third pair of muscles (tergosternal mt-s. 8-9) extends from anteromedian part of the tergite, is di- rected downwards, and inserts on the upper part of the vinculum. Dorso-ventral muscles mt-s. 8 are well developed; they are located in the caudal part of segment viii and also in other visceral segments. The mt-s. 8 extend paramedially from tergite to sternite as two bands. Dorso-ventral pleural musculature mpld-v. 8 is also well developed. Two pairs of muscles, directed towards the genitalia, are also located on sternite viii (Fig. 1, 2). Sternal intersegmental longitudinal muscles ms. 8-9 (1) extend paramedially backwards and upwards from the anterior margin of segment viii to the upper part of the vinculum inserting on it directly bene- ath mt-s. 8-9. The shorter and thicker sternal longitudinal muscles ms. 8-9 (2) extend from sternite viii

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rather caudad and medially with respect to ms. 8-9 and extend medially to the tight intersegmental membrane inserting on it behind the posterior margin of vinculum.

2. The skeleton of the genitalia (Figs 3-5) The tegumen of Oenosandra is trapezoidal, with a short sagittal reinforcing rib basally. The transi- tion between the tegumen and the hooked uncus is without any articulation. The base of the uncus late- ro-ventrally forms a pair of unarticulated sclerotized outgrowths – the socii. The comparatively long and weakly sclerotized anal cone is located directly beneath the uncus and the socii. The vinculum is narrow and sulcate, with a heavily sclerotized anterior margin. Each side of the vinculum has dorso-laterally directed apodemal plates; these plates are tightly connected to the base of the tegumen by a comparatively broad membrane. The saccus is weakly developed. The valves are rectangular, with well-developed sacculi that end in flat hooked cuillers. The scle- rotized bases of the upper margins of the valves form a transtilla of intricate shape. Each lateral lobe of the transtilla bears a short basal heavily sclerotized outgrowth that is weakly invaginated inside the ab- domen. The lobes of the transtilla are connected medially by a thin neck. The lower angles of the sac- culi approach each other. The disc-shaped juxta is located directly beneath the aedeagus between the valves. The aedeagus is thick, with a short basal outgrowth. The aedeagus and juxta are connected by the manica; this is very short and fused with the aedeagus beside its apex.

3. The musculature of the genitalia (Figs 3-5) Six pairs of muscles are present in the male genitalia of O. boisduvalii: m1, m3(2), m4, m5(6), m6(7), m7(5), m8(3); they move the copulatory appendages. One odd muscle m21 is located inside the aedeagus. m1 – musculi tergales intersegmentales longitudinales (tergal intersegmental longitudinal muscles of segments ix-x, depressors of uncus); these are very powerful and extend ventro-medially from the anterior margin of the tegumen to the base of the uncus. Muscles m2 (musculi tergales intersegmentales longitudinales of segments x-xi, retractors of the anal cone) have not been found. The valves are moved by three pairs of directly acting muscles: m3(2), m4, and m5(7) and one pair of muscles m8(3) with an indirect action. m3(2) - musculi gonopodales externi dorsolaterales, abductors of the valves. These comparatively thin muscles extend paramedially from the anterior margin of tegumen and are directly connected with the basal outgrowths of the transtilla beneath. m4 – musculi gonopodales externi dorsomediales, adductors of valves. They are shorter and much more voluminous than m3(2). Muscles m4 extend from the base of the tegumen beside the tegumino- vincular articulation medially to the upper parts of the transtilla lobes. Muscles m4 and m3(2) are anta- gonistic and cross each other. m5(7) – musculi gonopodales interni mediales, flexors of clasper. These intravalval muscles bend the valves transversely and their hard cuillers become directed medially. The cuillers immobilize the end of the female abdomen during the copulation. m8(3) – musculi laminae mediales anteriores, indirect action abductors of the valves. They con- nect the distal margin of the juxta with the anteromedial margin of the vinculum. Obviously, their con- traction produces a mechanical pressure in the region of the saccular bases and, as a result, opens the valves. m6(5) – musculi phallici externi posteriores, protractors of aedeagus. They extend from the dorso- lateral apodemal extensions of the vinculum to the basal outgrowth of the aedeagus, inserting on it late- rally and partly on top. m7(6) – musculi phallici externi anteriores, retractors of aedeagus. They extend from the anterior margin of the median area of the vinculum to the apex of the basal outgrowth of the aedeagus, inserting on it upward beside m6(5).

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The position of the protractors and retractors of the aedeagus and the presence of a short manica enable not only the retraction and protraction of the aedeagus but also allow it to pivot on the juxta. m21 – musculus phallicus internus longitudinalis, retractor of vesica. It is located inside the basal outgrowth of the aedeagus.

Description of the female terminalia

1. The skeleton of the terminalia (Figs 6-8) The end of the solid cylindrical female abdomen is formed by a big cylindrical segment vii, by a small segment viii and by setous anal papillae. The seventh segment is divided into a short proximal part with membranous pleura and a much more solid cone-shaped distal part without pleural membra- nes. The cone-shaped part of segment vii is uniformly, but weakly, sclerotized. The surface of the seg- ment is covered by a black squamose bush of hair-scales –called a corethrogyne; it hides the minute segment viii and the anal papillae. The eighth segment is heavyly sclerotized dorsolaterally and has a cover of a few strong hair-sca- les. Its ventral part is weakly sclerotized distally, has no setae and bears proximally the ostium which is a narrow transverse aperture. The sclerotization behind the ostium is fused with a dorsal sclerotization of segment viii. The anterior apophyses are attached to the anterior margin of segment viii. The anal papillae (segment ix) are weakly contracted and retracted into the interior of segment viii. They are densely covered with strong bristles. The posterior apophyses are inserted at the anterior margin of segment ix.

2. The musculature of the terminalia (Figs 8-10) Nine pairs of muscles are present in segments vii to ix: m1, m2, m3, m4, m5, m6, m7, m10, and m14. They retract and expand these segments, depress, lift, and rotate the end of the abdomen. m1 – musculi tergales intersegmentales longitudinales (tergal intersegmental longitudinal mus- cles). They extend paramedially from tergite vii to the anterior margin of tergite viii and retract segment viii. m2 - musculi apophysales anteriores 7-8 (anterior apophyseal muscles of the anterior apophyses); they connect the anterior margin of tergite vii with the tip of the anterior apophyses and retract segment viii. m4 - musculi apophysales anteriores 7-9 (anterior apophyseal muscles of the posterior apophy- ses); they connect the anterior margin of tergite vii with the tip of the posterior apophyses. Muscles m4 are attached on tergite vii together with m7 and united with each other at a distance. Muscles m4 retract the anal papillae. m5 - musculi interapophysales 8-9 (interapophyseal muscles) connect the tips of the anterior apophyses with the base of the posterior apophyses and move the anal papillae upwards. The function of these muscles significantly depends on the functional state of the other ovipositor muscles. m3 and m6 - musculi apophysales posteriores (posterior apophyseal muscles); they are antagonis- tic with the anterior apophyseal muscles. m3 - musculi apophysales posteriores 7-8 (posterior apophyseal muscles of the anterior apophy- ses) extend backwards from the tip of the anterior apophyses to the anterior margin of segment vii and protract segment viii. m6 - musculi apophysales posteriores 8-9 (posterior apophyseal muscles of the posterior apophy- ses) extend backwards from the tip of the posterior apophyses, inserting laterally at the anterior margin of segment viii; they protract the anal papillae. m7 - musculi sternales intersegmentales longitudinales (sternal intersegmental longitudinal mus- cles) extend from sternum vii to the anterior margin of the ventral part of segment viii, inserting on it paramedially. m10 - musculi apophyso-sternales 8-9 (apophyso-sternal muscles) extend from the tip of the ante-

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rior apophyses to a membrane-like fold beneath the anal papillae. Muscles m10 are depressors of the anal papillae and antagonists of the interapophyseal muscles m5. m14 - musculi pleurales dorsoventrales 9 (pleural dorsoventral muscles of segment ix). They (to- gether with the parietal and longitudinal muscles of the visceral segments) maintain the haemolymph turgor inside the abdomen.

Features of skeleton and musculature of segment viii in males of Oenosandra boisduvalii

1. Tergal longitudinal intersegmental muscles mt.8-9 (Fig. 1-2). According to FORBES (1939), these have been found in all examined Lepidoptera. These muscles have a constant position and con- nect paramedially or medially the anterior margins of tergite viii with the tegumen. O. boisduvalii has two pairs of tergal longitudinal muscles - mt.8-9 (1) and mt.8-9(2), the presence of a second pair is pos- sibly apomorphic. 2. Sternal longitudinal intersegmental muscles ms. 8-9 (Figs 1, 2). Two, three or four pairs of these muscles have been found in most examined Lepidoptera (KUZNETZOV & STEKOLNIKOV, 2001). FORBES (1939) reported 3 pairs (MS1, MS2, and MS3) in . Only two pairs of tergal longitu- dinal muscles are present in O. boisduvalii (ms.8-9 (1) and ms.8-9 (2)). 3. Tergo-sternal intersegmental muscles mt-s.8-9 (Figs 1, 2). These muscles (MT2 according to FORBES, 1939; m. tergovincularis in HANNEMANN, 1957) have been found in (HANNEMANN, 1957), , Geometridae, , Noctuidae (FORBES, 1939; KUZ- NETZOV & STEKOLNIKOV, 2001), and, recently, in Notodontidae (KUZNETZOV & STEKOLNI- KOV, 2004b, in press). Muscles mt-s.8-9 connect the anterior margin of tergite viii either with the vin- culum or the ventral part of the annulus. 4. Tergosternal intrasegmental muscles mt-s.8 (Fig. 2). These muscles have been found in O. bois- duvalii, in a species (KUZNETZOV & STEKOLNIKOV, 2001), in Lasiocampidae (KUZ- NETZOV & STEKOLNIKOV, 1985), and in Notodontidae (KUZNETZOV & STEKOLNIKOV, 2004b, in press). Muscles mt-s.8 are unknown in most other examined Lepidoptera. The origin of the muscles is obviously a result of hypertrophy in some part of the dorso-ventral pleural musculature.

Features of skeleton and musculature of the male genitalia in Oenosandra boisduvalii

1. Muscles m1 (Fig. 3). Muscles m1 are included in the basic pattern of musculature. In , they extend from segment ix to the base of the bilobed tergite x. In , the posterior position of the attachment of these ancestral muscles has been transferred to the uncus which is formed from the bilobed tergite x. According to FORBES (1939), m1 are depressors of the uncus (they move the uncus down). The position and function of m1 in Ditrysia are constant. The position of m1 attachment is transferred to the base of the gnathos or the socii only in the case of reduction of the uncus. The Notodontidae are the only group within the Noctuiformes where a connection of m1 with the well developed uncus is lost. In the Notodontidae, m1 inserts at the socii (TIKHOMIROV, 1979, 1981). In only three studied genera (Semidonta Staudinger, 1892, Pygaera Ochsenheimer, 1810, and Fentonia Butler, 1881), has m1 an an- cestral attachment at the base of the uncus (TIKHOMIROV, 1979). In O. boisduvalii, socii are developed, but m1 inserts at the uncus; so its position is plesiomorphic.

2. Valval muscles m3(2) and m4 (Figs 3, 4). In Ditrysia, the valves are moved mainly by muscles m3(2) and m4. Muscles m4, adductors of the valves, are likewise developed in the male genitalia of all Lepidoptera (KRISTENSEN, 1984), and their reduction is very rare. Muscles m3(2) draw the valves apart and open them (abductors). These muscles are absent in most Monotrysia and are only known in and , as well as in most Ditrysia. Both pairs of muscles insert medially and paramedially on the anterior margin of the tegumen

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and are directed downwards to the apodemal structures of the upper basal margin of the valves. Mus- cles m4 have two different positions, they insert either on the vinculum or on the base of the tegumen. A comparison of these positions in different taxa indicates that the first position is primary for Ditry- sia, and the latter position is secondary and convergent (Table 2). Moreover, m4 is attached to the lo- wer part of the vinculum in archaic Lepidoptera (Psychoidea, , , Yponomeu- toidea and so on), whereas m4 inserts at the upper part of the vinculum in more advanced groups. In the most advanced taxa (, Notodontoidea, and Noctuoidea), with rare exceptions, m4 is attached to the base of the tegumen. In Noctuidae, the displacement of m4 is delayed during the evolu- tionary process owing to the development of paratergal sclerites (separate parts of the vinculum) that enable a movement of tegumen and vinculum. However, in several subfamilies of Noctuidae, the para- tergal sclerites are fused with the tegumen and, as a result, m4 inserts at the tegumen. Presumably, the displacement of m4 to the base of tegumen has also happened many times in other noctuid families. For example, in Oenosandridae, m4 is attached at the tegumen although the tegumino-vincular articu- lation is clear.

Table 2. Position of genital muscles m4 in Lepidoptera, according to KUZNETZOV & STEKOLNIKOV (2001)

Taxon Muscles attached at: Vinculum tegumen paratergal sclerites Psychoidea, Tineoidea, Gracillarioidea, + Elachistoidea: , Ethmiidae + Elachistoidea: Agonoxenidae + + Coleophoroidea: Xyloryctidae + Coleophoroidea: + + Coleophoroidea: , , , + ; ; Tortricoidea , , , Schreckensteinioidea, , + Thyridoidea, Copromorphoidea () Pterophoroidea + Calliduloidea, Uranioidea + , + + Hesperioidea, + + + Sphingoidea, Bombycoidea; Oenosandridae, Notodontidae, + Lymantriidae Arctiidae + + Aganaidae + + Noctuidae: Nolinae, Chloephorinae, Sarrothripinae + Noctuidae: Acontiinae + + Noctuidae: Acronictinae, Agaristinae, Plusiinae, Heliothinae, + Cuculliinae, Amphipyrinae, Hadeninae, Noctuinae, Herminiinae, Hypeninae Noctuidae: Pantheinae, Scoleopteryginae + Noctuidae: Rivulinae + + Noctuidae: Catocalinae + +

The valve abductors m3(2) usually connect the dorsal part of the tegumen to the valves, inser- ting on it from above. Compared with m4, muscles m3(2) are sporadically reduced in different taxa.

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There is a tendency to reduce m3(2) in the phylogenetically advanced taxa. So, m3(2) is reduced in all Hesperioidea and Papilionoidea, and also in most Lasiocampoidea and Bombycoidea. Within the Noctuiformes, m3(2) is absent in the Notodontidae, Lymantriidae, Aganaidae, Arctiidae, and in a part of the Noctuidae (Nolinae, Chloephorinae, Sarrothripinae, Herminiinae, Rivulinae, Hypeninae, Catocalinae, and Calpinae). Two pairs of antagonistic valval muscles are present in the other, the tri- fine, subfamilies of Noctuidae. There are two opinions about the homology of these muscles in the Noctuidae. According to TIKHOMIROV (1979), one pair of muscles (m4) is the adductor of valves, the second pair corresponds to the abductor m3(2). According to SPEIDEL et al. (SPEIDEL, FÄN- GER & NAUMANN, 1996), however, the abductor of valves in the trifine subfamilies is not homo- logous to the abductor muscle found outside the Noctuiformes. The trifine noctuid subfamilies are normally not considered basal within the Noctuiformes, and therefore the duplication of the valval muscles in the trifine Noctuidae is interpreted by these authors to be a result of a splitting of the mus- cles m4. Both m3(2) and m4 are well developed in the Oenosandridae. The valval abductor m3(2) of the Oenosandridae may be homologous with the same muscle found outside the Noctuiformes. This as- sumption is mainly based on the distribution of this character within the Noctuiformes. The homo- logy of these muscles is very difficult to determine, as there are slight differences in their insertion and antagonistic function making morphological evaluation complicated.

3. Tegumino-vincular articulation (Fig. 3) The articulation of tegumen and vinculum evolved independently in several ditrysian Lepidop- tera (e. g. in Tineiformes, Yponomeutiformes, Gelechiiformes and in other Lepidoptera). This articu- lation has developed in an unusual way in the Noctuidae, Aganaidae, and Arctiidae. These groups are characterized by a convergent evolution of the paratergal sclerites instead of a direct tegumino-vin- cular articulation. The paratergal sclerite is occasionally fused with the vinculum in some Noctuidae. Within the Noctuoidea s. str., paratergal sclerites are absent and the tegumen and vinculum are fused in the Lymantriidae. In the Oenosandridae, the tegumino-vincular articulation is present, but parater- gal sclerites are absent corresponding to the plesiomorphic condition.

4. Muscles m8 (3) (Figs 3, 4, 5) The presence of muscles m8(3) is included in the ground plan of the lepidopterous male genita- lia in one hypothesis (KRISTENSEN, 1984). In the other hypothesis (KUZNETZOV & STEKOLNI- KOV, 2001), m8(3) evolved repeatedly in the Lepidoptera by a splitting of the phallic muscles. In any case, the presence of these muscles in ditrysian Lepidoptera is a symplesiomorphic feature. The position and function of m8(3) undergo a well-defined evolutionary trend (KUZNETZOV & STEKOLNIKOV, 2001, Table 3). In Tineoidea and Yponomeutoidea, these muscles extend from the vinculum to the valves or to the annellus and retract the valves. In the phylogenetically advanced Lepidoptera, the position of the m8(3) attachment is transferred from the annellus to the juxta. So, the juxto-vincular position of these muscles has become the most widespread one in Lepidoptera. This is characteristic for Coleophoroidea, Tortricoidea, Cossoidea, Zygaenoidea, Sesioidea, Pyraloi- dea, , Uranioidea, Geometroidea, Drepanoidea, Papilionoidea and for some Sphingoidea and Bombycoidea. In this case the muscles m8(3) are still responsible for the abduction of the val- ves. But, in some superfamilies and at the same time, muscles m8(3) changed their position from a vinculo-valval to a juxto-valval one without modification of their function (abductors of the valves). This transfer of the insertion evolved in the morphologically most specialised taxa, such as Crambi- dae (Pyraloidea), Geometrinae (Geometridae), Hesperiidae, a part of the Papilionoidea (, partim, and most ), Lemoniidae (Lasiocampoidea), and Bombycoidea. Mus- cles m8(3) have a secondary juxto-valval position in the genitalia of all examined Notodontoidea and Noctuoidea. O. boisduvalii (Oenosandridae) is the only species of Noctuiformes with a plesiomorphic juxta- vincular position of muscles m8(3).

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Table 3. Position of muscles m8 (3). Abbreviations accepted in the table: vnc-vlv - from vinculum to valvae; vnc- anl - from vinculum to anellus; vnc-jxt - from vinculum to juxta; jxt-vlv - from juxta to valvae

Taxon Position of muscles m8 (3) vnc-vlv vnc-anl vnc-jxt jxt-vlv, vnc-jxt jxt-vlv without m8(3) Tineoidea + + + Psychoidea + + Gracillarioidea + Epermenioidea + Yponomeutoidea + + + Elachistoidea + + + + Coleophoroidea + + + + Gelechioidea ++ Tortricoidea + + Cossoidea + + Zygaenoidea + + Sesioidea, Schreckensteinioidea + Pyraloidea + + Thyridoidea + Copromorphoidea: Carposinidae + Alucitoidea, Pterophoroidea, Calliduloidea, Uranioidea, + Geometroidea + + Drepanoidea + Hesperioidea, Papilionoidea, Lasiocampoidea, Sphingoidea + + + Bombycoidea + + + Oenosandridae + Notodontidae, Lymantriidae, Aganaidae, Arctiidae, Noctuidae +

5. Intravalval muscles m5(7) (Figs 3, 4). Intravalval muscles have been found in ditrysian Lepidoptera only. These muscles help to link the sexes in copulation. They bend the valves in order to fix the female abdomen with the help of the harpe, cuiller and other sclerites. However, these muscles are occasionally reduced in many families. In Noc- tuiformes and related taxa, m5(7) is usually well developed. It is reduced only in the Notodontidae or it remains in an underdeveloped state in some notodont species (e.g Spatalia argentina ([Denis et Schif- fermüller], 1775), Spatalia doerriesi Graeser, 1888, Semidonta biloba (Oberthür, 1880), Peridea gigan- tea Butler, 1877 (= monetaria Oberthür, 1879) and others (TIKHOMIROV, 1981)). The absence of m5(7) in other Noctuiformes is rare. This character clearly distinguishes the Notodontidae from Oeno- sandra boisduvalii which has well developed intravalval muscles as in most Noctuiformes.

6. Transtilla (Fig. 4). The transtilla is often present in Lepidoptera. This is likely to be due to convergence, as the medial accretion of the hemitranstillan lobes is a simple process. Of all noctuid Lepidoptera, the Noctuidae ha- ve the best developed transtilla, but the presence of basal outgrowths of the transtilla is undoubtedly an

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autapomorphic feature of the Oenosandridae. A very similar transtilla with well developed basal out- growths, as in Oenosandra boisduvalii, is present only in the Sphingidae (KUZNETZOV & STEKOL- NIKOV, 2001); and is considered a result of homoplasy.

7. The morphology of the aedeagus with special reference to its function (Figs 3, 5). A significant contraction of the manica and a displacement of m7(6) on the basal outgrowth of the aedeagus beside m6(5) occur, in addition to Oenosandra boisduvalii, in the Notodontidae, subfamily Cerurinae (TIKHOMIROV, 1979) and in Germar, 1812 (KUZNETZOV & STEKOLNI- KOV, 2004, in press). Moreover, the same morphological constructions with identical functions are also known in the Lasiocampidae.

8. Bulbus ejaculatorius The morphology of the bulbus ejaculatorius was first considered in the phylogeny of the noctuid Lepidoptera by SPEIDEL et al. (SPEIDEL, FÄNGER & NAUMANN, 1996). It has been shown that the Oenosandridae have a shorter bulbus ejaculatorius than the Arctiidae+Aganaidae+Noctuidae. The long bulbus ejaculatorius is considered as a synapomorphic feature of these families. The form of the bulbus seems to be correlated with the form of the spermatophore.

Features of the skeleton and musculature of the female genitalia of O. boisduvalii

1. Anal tufts (corethrogyne) In various noctuid Lepidoptera, the scales of the anal tufts (corethrogyne) are concentrated in dif- ferent regions at the end of abdomen; in Thaumetopoea on the tergite of segment vii and in Lymantrii- dae, Panthea, and Diloba over the whole of segment viii. In Oenosandra boisduvalii, the scales cover the complete cone-shaped segment vii. The different location of the corethrogyne testifies to its inde- pendent origin in the different noctuid Lepidoptera. Sometimes, Lepidoptera with a corethrogyne also have a similar internal structure of the genitalia by convergence. So, in Thaumetopoea and Oenosan- dra, irregular muscle-bundles extend from the base of the apophyses of segment viii to the dorsal wall of segment vii; their contraction probably promotes the deposition of the scales of the corethrogyne.

2. Muscles of the ground plan (Figs 8-10) Muscles m1, m2, m3, m4, m5, m6, m7, m10 are included into the ground plan of the Noctuifor- mes (KUZNETZOV & STEKOLNIKOV, 2004a); their position in Oenosandra is indistinguishable from the earlier described scheme.

3. Posterior apophyseal muscles of the apophyses of segment ix – m6 (Figs 8, 10) Only one pair of posterior apophyseal muscles (m6) has been found in Oenosandra boisduvalii. In most Noctuiformes examined, 2 pairs of these muscles (m6a and m6b) are present. The reduction to one pair occurs in only two species of Notodontidae ( dodonaea ([Denis & Schiffermüller], 1775) and (Clerck, 1759)) and happened independently in Oenosandra.

4. Oviduct dilators m12 Among Noctuiformes, muscles m12 are present in Lymantriidae and Noctuidae, but absent in No- todontidae (except in Pterostoma palpina), Arctiidae, and, as shown here, in Oenosandridae. These muscles, probably, arose convergently in various groups of ditrysian Lepidoptera and were not included in the ground plan of the Noctuiformes (KUZNETZOV & STEKOLNIKOV, 2004, in press). The presence of m12 in the most specialised species of Notodontidae, Pterostoma palpina, strengthens this assumption.

5. Tergo-sternal dorso-ventral muscles m14 in segment vii (Fig. 8) These muscles have evolved from the parietal pleural musculature in different segments, both in

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the genital segments as well as in the pregenital ones. As shown in Notodontidae, Lymantriidae (Ly- mantria dispar (Linnaeus, 1758)), and in several Arctiidae (KUZNETZOV & STEKOLNIKOV, 2004, in press), these muscles are developed in segment viii (m13). Only in Xyleutes persona Le Guillou, 1841 (Cossidae; KUZNETZOV & STEKOLNIKOV, 2004a), are these muscles well developed in seg- ment vii, as in Oenosandra boisduvalii.

Discussion

At least two different classifications exist of the with thoracic tympanal organs. They re- flect no substantial difference, but only a formal one. One classification distinguishes two superfami- lies; Notodontoidea for the group with basically trifid forewing venation and Noctuoidea for the group characterized by a quadrifid forewing venation, whereas the other classification recognizes only a sin- gle superfamily Noctuoidea (s. l.). In the first classification another name, Noctuiformes, is used to uni- te the two superfamilies, whereas in the second classification one of the branches from the primary di- chotomy of the Noctuoidea is without a name. Both classifications have deficiencies: in the first system, there is no traditional category (rank) available for the name Noctuiformes, if we accept Ditry- sia as a subordinal name. In the other system, accepting the Noctuoidea in the wide sense, there is no traditional rank available to subdivide the Noctuoidea into the two main-groups and yet retain the tradi- tional families. The wide concept of Noctuoidea has a small advantage, as the difference in the tympanal organs between Notodontoidea and Noctuoidea seems to be less fundamental than originally indicated by Ki- riakoff when establishing the two superfamilies. He thought that the tympanal organs of both groups were at least diphyletic and did not believe in the monophyly of the Noctuiformes. However, the posi- tion of some groups (e.g. Platychasmatinae, Dilobidae) has recently been changed, which seems to in- dicate that the tympanal organs are not so fundamentally different, so there is no reason in our opinion to doubt their monophyly. However, the functional organisation of the male genitalia of the Notodonti- dae demonstrates a particular trend in the evolution of this group and another trend can be seen in the other Noctuiformes. Therefore, there is evidence for a primary dichotomy of the Noctuoidea s. l. Thus, the problem of noctuid Lepidoptera macroevolution will continue to be a subject of future investigation and evaluation. Unfortunately, this study of male and female genitalia in Oenosandra did not solve this taxonomic dilemma and provided no clear primary division of the Noctuiformes. Many muscles of the male and female terminalia of Oenosandra boisduvalii are included in the basic structural pattern of ditrysian Lepidoptera. These are m1, m3(2), m4, m5(7), m6(5), m7(6), m8(3), and m21 in male genitalia. Only m2, retractors of the anal cone, have not been found in O. bois- duvalii. However, the reduction of these muscles sporadically occurs in many groups of Lepidoptera and this feature cannot be considered in an assessment of the phylogenetic position of the group. In the female terminalia, almost all muscles are also included in the basic structural pattern of the ditrysian Lepidoptera. These are muscles m1-m7 and m10. In other Noctuiformes, all muscles described above are also present. However, a tendency to redu- ce m3(2) is found in the male genitalia. In the Notodontoidea, the same tendency occurs in m5(7). The positions of attachment of many muscles of O. boisduvalii are also ancestral. In the male ge- nitalia muscle m1 is inserted at the uncus (in Notodontidae at the socii); m3(2) is attached to the dorso- median section of the anterior margin of the tegumen; m3(8) has a juxto-vincular position (in other Noctuiformes it has a secondary juxto-valval position); m5(7) is situated inside the sacculi. At the same time, the position of the insertion of m4 is transferred towards the base of the tegumen in Oenosandra boisduvalii (as in most Noctuiformes). All muscles, described in the female terminalia, also have a position that is ancestral for ditrysian Lepidoptera. So, male and female terminalia of Oenosandra boisduvalii include a large complex of plesiomorp- hic muscular features that denote a basal position of the group in the noctuiform phylogenetic branch.

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At the same time, the functional organisation of the abdominal terminalia in O. boisduvalii is similar to both the Notodontoidea and the Noctuoidea. Some primitive characters were previously unknown in the Noctuiformes and change our concepts of the ground-plan of this group. Characters found only in the male genitalia of the Oenosandridae in- clude:

1.- mt-s.8 is present. The interpretation of the presence of this muscle is difficult, but it could indicate a relationship to the Lasiocampidae where it is also found, provided that the homology is accepted. It is a clue that the Lasiocampidae or the group to which they belong (Bombycoidea or Lasiocampoidea) could be the adelphotaxon of the Noctuoidea. Too few studies of the muscles of the pregenital segments in Lepidop- tera have been published to investigate this further.

2.- m3(2) is well developed in Oenosandridae. It is doubtfully present in the trifine Noctuidae and this may be due to convergence or parallelism.

Shared features of the Oenosandridae and Notodontoidea:

1.- Contraction of the manica and parallel displacement of m7(6) on the basal outgrowth of the aedeagus. As a result, the aedeagus moves vertically forward in an arch. A similar situation is known in some Noto- dontidae; in Cerurinae (TIKHOMIROV, 1979), in Pterostoma (KUZNETZOV & STEKOLNIKOV, 2004b, in press), in the Lasiocampidae (ROZSHKOV, 1965; STEKOLNIKOV, 1965) and in (KUZ- NETZOV & STEKOLNIKOV, 1973). This similarity can be considered as an example of homoplasy.

2.- Oenosandridae have a shorter bulbus ejaculatorius compared to the Arctiidae, Aganaidae, and Noctui- dae. The long bulbus ejaculatorius found in the latter families is considered an apomorphic feature (SPEI- DEL, FÄNGER & NAUMANN, 1996). All examined species of Notodontidae and Lymantriidae have a very short bulbus, and compared with them, the bulbus ejaculatorius of Oenosandra is rather long. So the intermediate length found in Oenosandra provides no evidence of a relationship to any of these families.

3.- Among Noctuiformes with anal tufts, the corethrogyne scales are situated on segment vii only in Oenosandridae and Notodontidae (Thaumetopoea). Moreover, both groups have irregular muscular bundles, extending from the anterior apophyses to tergite vii. However, the distribution of the scales on the segment and the structure of segment vii itself are different in these groups. In O. boisduvalii the scales cover the whole segment whereas in Thaumetopoea only the tergite. In O. boisduvalii, tergite and sternite of segment vii are fused into a cone-shaped annulus whereas in Thaumetopoea segment vii retains large membranous pleural regions. It is possible, that the same segmental location of the coreth- rogyne in Oenosandridae and Notodontidae is a parallelism.

4.- In females of the Noctuiformes, as in other ditrysian Lepidoptera, there are two pairs of muscles of the posterior apophyses in segment vii (m6a and m6b). However, in Oenosandra boisduvalii and in two species of Notodontidae (Drymonia dodonaea and Pterostoma palpina) the muscles, attached to the ventral area of tergite viii have been lost. This similarity is a result of homoplasy since the reduction of muscles is recorded in derived notodontid-genera.

5.- O. boisduvalii as well as the Notodontidae have no oviduct dilators m12 and this similarity is a sym- plesiomorphic feature.

Similar features between Oenosandridae and Noctuoidea (s. str. or quadrifid Noctuoidea):

1.- Oenosandridae and Noctuoidea share the presence of a transtilla in the male genitalia. However, this

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structure evolved independently in various groups of ditrysian Lepidoptera, including some small taxa. The muscles of the transtilla of Oenosandra are most similar to that of hawk moths (Sphingidae). This similarity of Oenosandridae and Noctuoidea is possibly convergent.

2.- Oenosandridae and Noctuoidea are similar in having a well-defined tegumino-vincular articulation in the male genitalia. But the structure of this articulation is different in various noctuid taxa. In con- trast to the Oenosandridae, the Noctuoidea have well developed paratergal sclerites in the tegumino- vincular articulation which are considered a result of secondary separation of the upper-lateral parts of the vinculum (KUZNETZOV & STEKOLNIKOV, 1989). Moreover, different forms of paratergal scle- rites are present in various Aganaidae. So, an independent origin of the articulation between tegumen and vinculum seems to exist in Oenosandridae and Noctuoidea. The similarity between the Oenosandridae and Noctuoidea in the presence of well developed in- travalval muscles m5(7) is a symplesiomorphic feature. So, the similarities of the Oenosandridae with both superfamilies of the Noctuiformes have evol- ved independently in most cases or constitute symplesiomorphies. Only the corethrogyne on segment vii can possibly be interpreted as a synapomorphy of the Oenosandridae and Notodontidae (Thaumeto- poeinae). However, in Oenosandridae a juxto-vincular position of muscles m8(5) occurs and muscles m3(2) are present which are remarkable differences to all other examined Noctuiformes. Muscles m3(2) are certainly reduced in most other Noctuiformes, excepting possibly the trifine Noctuidae, if these muscles are homologous in these groups. These two plesiomorphic features and the general ple- siomorphic character of the male and female terminalia indicate a basal position of the Oenosandridae in the Noctuiformes. On the other hand, convincing synapomorphies to link the Oenosandridae with certainty with any superfamily of the Noctuiformes have not been found.

Acknowledgments

The authors are grateful to Mr. E. D. Edwards (Canberra, Australia) for providing the males and females of Oenosandra boisduvalii. Moreover Ted Edwards kindly agreed to read the manuscript and has made many useful notes about the English text. The work was financially supported by the Russian Foundation for Basic Research, projects no. 02-04-49138 and 02-04-04002; the program “Universities of Russia – Fundamental Science”, project no. 07. 01. 052; the program “Leader Scientific Schools”, project no. 2232.2003.4. Furthermore, we thank the German Research Council (Deutsche Forschungs- gemeinschaft, DFG) for financial support (Na 90/17-1).

BIBLIOGRAPHY

COMMON, I. F. B., 1970.– Lepidoptera. In D. F. Waterhouse ed. The Insects of Australia: 765-866. Melbourne Uni- versity Press, Carlton, Victoria. COMMON, I. F. B. 1990.– Moths of Australia: 535 pp. E. J. Brill, Leiden, New York, Kobenhavn, Köln. EDWARDS, E. D., 1996.– Oenosandridae (page 271). In E. S. NIELSEN, E. D. EDWARDS & T. V. RANGSI. Checklist of the Lepidoptera of Australia: XIV + 529 pp. CSIRO. Australia. FÄNGER, H. & NAUMANN, C. M., 1997.– Genital morphology and copulatory mechanism in Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera, ).– Acta Zoologica, 79(1): 9-24. FORBES, W. T. M., 1939.– The muscles of the lepidopterous male genitalia.– Ann. ent. Soc. Amer., 39(1): 1-10. IVANOV, V. D., 2003.– Ground plan and nomenclature of muscles in the male terminalia of the Amphiesmenoptera (Insecta: Trichoptera and Lepidoptera).– Ent. obozr., 82(1): 17-32. KIRIAKOFF, S. G., 1970.– Lepidoptera. Familia Thaumetopoeidae. In P. WYTSMAN. Genera Insectorum. Lepi- doptera. Fasc. 219e: 54 pp., 3 pls. Kraainem. KITCHING, I. J. & RAWLINS, J. E., 1999.– 19. Noctuoidea. In N. P. KRISTENSEN (ed.). Handbook of Zoology. Vol. IV. Arthropoda: Insecta. Pt. 35. Lepidoptera, moths and . 1: Evolution, systematics and biogeo- graphy: 356-401. W. de Gruyter GmbH & Co. Berlin, New York.

SHILAP Revta. lepid., 32 (128), 2004 309 297 Kuznetzov (4-10-04) 3/1/77 18:28 Página 310

V. I. KUZNETZOV, C. M. NAUMANN (†), W. SPEIDEL & A. A. STEKOLNIKOV

KRISTENSEN, N. P., 1984.– The male genitalia of (Lepidoptera: Agathiphagidae) and the lepidopte- ran ground plan.– Ent. scand., 15(2): 151-178. KUZNETZOV, V. I. & STEKOLNIKOV, A. A., 1973.– Phylogenetic relationships in the family Tortricidae (Lepi- doptera) treated on the base of study of functional morphology of genital apparatus.– Trudy vses. ent. obshch., 56: 18-43. KUZNETZOV, V. I. & STEKOLNIKOV, A. A., 1985.– Comparative and functional morphology of the male genita- lia of the bombycoid moths (Lepidoptera, Papilionomorpha: Lasiocampoidea, Sphingoidea, Bombycoidea) and their systematic position.– Trudy zool. Inst. Leningr., 134: 3-48. KUZNETZOV, V. I. & STEKOLNIKOV, A. A., 1989.– Position of the tropical family Aganaidae in the system of the Noctuoidea (Lepidoptera) according to the results of a study of functional morphology of the male genita- lia.– Trudy zool. Inst. Leningr., 202: 200-210. KUZNETZOV, V. I. & STEKOLNIKOV, A. A., 2001.– New approaches to the system of Lepidoptera of world fau- na (on the base of the functional morphology of abdomen).– Trudy zool. Inst. St. Peterburg, 282: 1-462. KUZNETZOV, V. I. & STEKOLNIKOV, A. A., 2004a.– The musculature of female terminal segments and its im- plication for the systematics of the superfamilies Notodontoidea and Noctuoidea (Lepidoptera: Noctuifor- mes).– Ent. obozr., 83(2): 286-307. KUZNETZOV, V. I. & STEKOLNIKOV, A. A., 2004b.– New tribus Pterostomini Kuznetzov et Stekolnikov (Lepi- doptera, ) and the characters of terminalia muscles in both sexes.– Ent. obozr., 83 (in prep.). MILLER, J. S. 1991.– Cladistics and classification of the Notodontidae (Lepidoptera, Noctuoidea) based on larval and adult morphology.– Bull. Amer. Mus. Nat. Hist., 204: 1-230. ROZSHKOV, A. S., 1963.– Sibirskij shelkopryad. Moskow. Izd. AN USSR: 176 pp. SPEIDEL, W., FÄNGER H. & NAUMANN C. M., 1996.– The phylogeny of the Noctuidae (Lepidoptera).– Syst. Ent. 21 (3): 219-251. SPEIDEL, W. & NAUMANN C. M., 1995.– Further morphological characters for a phylogenetic classification of the Noctuidae (Lepidoptera).– Beitr. Ent., 45(1): 119-135. STEKOLNIKOV, A. A., 1965.– Functional morphology of the copulatory apparatus in some Lepidoptera.– Ent. obozr. 44: 143-149. TIKHOMIROV, A. M., 1979.– The functional morphology of the male genital apparatus and taxonomic division of the European Notodontidae (Lepidoptera).– Trudy zool. Inst. Leningr., 83: 104-119. TIKHOMIROV, A. M., 1981.– Taxonomic structure of the family Notodontidae and its position in the system of Le- pidoptera with regard of functional morphology of genitalia of species from the Far East.– Trudy zool. Inst. Le- ningr., 103: 62-72. TURNER, A. J., 1903.– Revision of Australian Lepidoptera.– Proc. Linn. Soc. New South Wales, 18: 42-92. TURNER, A. J., 1922.– Revision of Australian Lepidoptera. , , , Notodonti- dae.– Proc. Linn. Soc. New South Wales, 47: 348-390.

V. I. K. C. M. N. (†), W. S. Zoological Institute of Russian Academy Sciences Alexander Koenig Research Institute Universitetskaya emd. 1 and Museum of Zoology RUS-199034 Saint Petersburg Adenauerallee, 160 RUSIA / RUSSIA D-53113 Bonn ALEMANIA / GERMANY E-mail: [email protected]

A. A. S. Department of Entomology Saint Petersburg State University Universitetskaya emb., 7/9 RUS-199034 Saint Petersburg RUSIA / RUSSIA E-mail: [email protected]

(Recibido para publicación / Received for publication 6-IX-2004) (Revisado y aceptado / Revised and accepted 1-X-2004)

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mt-s. 8-9

mt. 8-9(1) mt. 8-9(1) mt-s. 8

mt. 8-9(2) mt. 8-9(2)

m3(2) mt-s. 8-9 mp ms. 8-9(1)

1 ms. 8-9(2) ms. 8-9(2) ms. 8-9(1) 2

M1 M3(2) M4

M4 M3(2)

M6(5) M5(7) M8(3)

M5(7) M8(3) 4 M7(6) 3

Figs. 1-4.– 1. Male genitalia of Oenosandra boisduvalii and muscles of the eighth segment, lateral view. 2. Muscles of the eighth segment of the male abdomen of Oenosandra boisduvalii, inner view. 3. Male genitalia and genital muscles of Oenosandra boisduvalii, lateral view. 4. Ventral complex of genitalia appendages of male Oenosandra boisduvalii: valvae, transtilla, juxta and muscles, posterior view.

SHILAP Revta. lepid., 32 (128), 2004 311 297 Kuznetzov (4-10-04) 3/1/77 18:29 Página 312

V. I. KUZNETZOV, C. M. NAUMANN (†), W. SPEIDEL & A. A. STEKOLNIKOV

VI

VII

VIII an.p

M6(5) 6

M8(3) 5 M7(6)

VIII

VII

7 ost.b

Figs. 5-7.– 5. Aedeagus of Oenosandra boisduvalii with juxta and muscles, lateral view. 6. Female abdominal end of Oenosandra boisduvalii, lateral view. 7. dto., ventral view.

312 SHILAP Revta. lepid., 32 (128), 2004 297 Kuznetzov (4-10-04) 3/1/77 18:29 Página 313

THE SKELETON AND MUSCULATURE OF MALE AND FEMALE TERMINALIA IN OENOSANDRA BOISDUVALII NEWMAN, 1856

m6 m4

m5

m10 10 m1 m6

m2

m5 m10 m3 VII

m7 9

m4 m1 m5 m2

mp

m2

m14 m10 an.p

8

m7

Figs. 8-10.– 8. Section of the abdominal end of Oenosandra boisduvalii female showing the positions of the attachment of the muscles. 9. Section of the eighth abdominal segment of Oenosandra boisduvalii female showing the positions of the attachment of the muscles. 10. Section of the ninth abdominal segment of Oenosandra boisduvalii female showing the positions where the muscles are attached.

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