Hypsidia Rothschild: a review and a reassessment (: , )

MALCOLM J. SCOBLE and E. D. EDWARDS

The Australian genus Hypsidia Rothschild, currently assignedto the , is reassigned to the Drepanoidea (Drepanidae sensu Minet (1983), 'Cymatophoridengruppe' of Kennel & Eggers (1933)). Baryphanes Turner, a genus previously mIsplaced in the is synonymizedwith Hypsidia. A new (Hypsidia grisea) is described. EggersopsSick, a genus with a single species, is correctly assignedto the Drepanoidea, and is synonymizedwith Hypsidia. Hypsidia is divided into the monophyletic erythropsalisgroup and, for convenience, the niphosema 'group', which may or may not be monophyletic.The distribution of the expanded genus is disjunct, with the erythropsalisgroup confined to rainforests of northeastern Queens- land and the niphosema 'group' to southern Western Australia. The reassignmentof Hypsidia(in the revised sense) to the Drepanoidea is based largely on the form of the tympanal organs. The structure of these organs is summarized. The position of Hypsidiawithin the Drepanoidea remains uncertain, but we briefly explore the question taking particular account of the presence of ocelli in the speciesof the niphosema 'group', the arrangement of the veinsin the hindwing,and the clubbed frenulum of the male. Key words: Drepanoidea. Drepanidae. Pyralidae, Noctuidae, Hypsidia, Baryphanes, Egger- sia, Eggersops, Australia. tympanal organs. M. J. Scoble. British Museum (Natural History), CromwellRoad, London SW75BD, UK. E. D. Edwards. Division of Entomology. CSIRO. GPO Box 1700,Canberra A.C.T. 2601, Australia.

The Australian genus Hypsidia Rothschild was as- scaled at the base, a condition unlike that in any signed originally to the Pyralidae (Rothschild other member of the family. The exceptional na- 1896). Recently, the genus was redescribed by ture of Hypsidia was also noted by Hacobian Hacobian (1986) who provided an illustrated ac- ( 1986) . count of the two included species H. erythropsalis Since the publication of Hacobian (1986) some Rothschild and H. robinsoni Hacobian. Both additional findings have come to light. Two species are brightly coloured , and have species of the genus Baryphanes Turner, until this been collected only from rainforest in northeast- study assigned to the Noctuidae, arc here found to ern Queensland (Fig. 33). be closely related to erythropsalis and robinsoni. Although Hypsidia has been retained in the Another species, Eggersops australica (Sick), Pyralidae (e.g., Fletcher & Nye 1984), its position originally placed in the Cymatophoridae, has also there has long been recognized as anomalous been found to be closely related. The species of (I.F.B. Common pers. comtn.). Munroe (1972) Hypsidia, Baryphanes, and Eggersops exhibit a did not refer to Hypsidiu by name, but presumably marked similarity in genitalia, and below we had this genus in mind when he stated that in one synonymize the genera with Hypsidia as the senior Australian pyralid genus the proboscis was not synonym. 334

The purpose of the present paper is to reassess Tympanal organs of Hypsidia and the taxonomic position of Hypsidia and to rede- Drepanidae (Drepanoidea) fine the genus to include the additional specics. The structure of the abdominal hearing organs of Lepidoptera was described in detail by Kennel & Eggers (1933), and summarized by Abbreviations Bourgogne (1951) and Minet (1983). Minet made extensive ANIC Australian National Collection, Divi- use of these structures in his work on the higher sion of Entomology, CSIRO, Canberra classification of the The of EDE E. D. Edwards Lepidoptera. summary ESN E. S. Nielsen the structure of the drepanid tympanal organs pro- IFBC I. F. B.Common vided here, refers particularly to those of Hypsidia MSLJ M. S. Upton ?Vf/!A'0/7.M//.S. SAMA South Australia. Museum, Adelaide In drcpanids, the tympanal organs include com- ponents from plcural and sternal regions at the base of the abdomen. At the base of the first visi- Taxonomic history ble sternum (sternum II) there is a pair of small, oval structures (Fig. 11). Each is composed of two The genus Baryphanes was proposed by Turner interconnecting chambers (Fig. 12), one small and (1933) to include Mortoctenia niphosema Lower, more anterior (`kleine Sternalblase' of Kennel & described earlier (Lower 1908) in the Geomet- Eggers 1933), the other larger and more caudal ridac. Turner placed the species in the Noctuidae ('grosse Stcrnalblasc' of Kennel & Eggers). The and subsequently described an additional species, tympanum is unique in being situated within the B. microspila Turner (Turner 1942). Howcver, sternal component of the tympanal organ between Common ( 1969), in his study of the locking mech- the small and large chambers (Kennel & Eggers anism of the metathorax and the forewing, recog- 1933: fig. 17). nized that Baryphanes was misplaced in the Noc- The pleural component of the tympanal organ is tuidae. In thc absence of an alternativc position, formed from a prominent pleural fold, supported the genus was retained in that family by Nye by a large, three-armed sclerite. Anterodorsally ( 1975 ) . this distinctive sclerite meets I at its an- Sick described australica Sick tergum ( 1938) Eggersia terolateral and divides into and, as a result of a of its margin, prespiracular study tympanal organs, and it to the postspiracular components (Fig. 13). This assigned (as Cymatophori- sclerite and the fold can be seen on Sick is a plcural clearly nae) [Drepanidae]. (Egger.sia junior dried when the hair-like of a of bectles and was specimens, particularly homonym genus replaced scales are removed or The scierites Fletcher Indeed, it turns parted. pro- by Eggersops (1979).) vide a framework across which is stretched an an- out that the of this entire systematic position terior counter and a of is revealed the structure of the tympanum posterior tym- group species by These membranes enclose the Like those of the panum. pleural tympanal organs (Figs. 11-13). chamber of each these arc found at the base of tympanal organ. Pyralidae, organs The form of the sternal chambers is characteris- the abdomen, but close examination shows them tic of the as is that to be different in form. In fact are iden- Drepanidae including Hypsidia, quite they of the chamber with its tical to the characteristic of pleural three-armed, very tympanal organs arched sclerite. the Drepanidae (sensu Minet 1983, i.e., including , Thyatirinae, and Cyclidiinae) and it is to this family that the species under discussion Monophyly of Hypsidia belong. The drepanid (as 'Cymatophoridengruppe') The monophyly of Hypsidia is supported by the hearing organ was well described by Kennel & Eg- presence of four, prominent, specialized struc- gers (1933) in their unsurpassed study of the ab- tures. As far as we know these are unique within dominal tympanal organs of Lepidoptera, namely the Drepanidae: 1 ) a pair of characteristic arms, those of gcometrids, pyralids, cymatophorids arising from between the uncus and the tegumen, (thyatirines) + drepanids, and uraniids + epi- each arm with a pectiniferous, spatulate head; 2) a plemids. long, narrow sclerite in the dorsal part of the dia-