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A New Brachypterous Nusalala Species from Costa Rica, with Comments on the Evolution of Flightlessness in Brown Lacewings (Neuroptera: Hemerobiidae)

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A New Brachypterous Nusalala Species from Costa Rica, with Comments on the Evolution of Flightlessness in Brown Lacewings (Neuroptera: Hemerobiidae) Systematic Entomology (1996) 21, 343-352 A new brachypterous Nusalala species from Costa Rica, with comments on the evolution of flightlessness in brown lacewings (Neuroptera: Hemerobiidae) J 0 H N D . 0 S WA L D Department of Entomology, Texas A&M University, College Station, Texas, U.S.A. Abstract. A new flightless hemerobiid species, Nusalala brachyptera, collected at high elevation in Costa Rica, is described and illustrated, and a variety of data relevant to the evolution of flightlessness in the family Hemerobiidae are reviewed. Flightlessness due to brachyptery has evolved independently in at least five monophyletic [= holophyletic] lineages of the family Hemerobiidae (brown lacewings). Volant hemerobiids are primarily foliage foraging arboreal predators [presumed ancestral condition], while flightless species are predominantly associated with terricolous-type microhabitats (e.g. ground-litter, epiphytic mosses) [presumed derived condition]. These differences suggest a significant habitat shift for flightless hemerobiid species, and that the parallel evolution of flightlessness and brachyptery in hemerobiids are shared responses to the conditions of a terricolous existence. The restriction of most flightless hemerobiid species to insular andlor montanelalpine land areas may be related to the typically depauperate nature of the faunas of such areas. This faunal characteristic may facilitate ttansitions from arboreality to terricolousness by presenting ancestrally arboreal predators such as hemerobiids with novel ecological opportunities in terricolous microhabitats. Introduction Helicoconis aptera Messner and H.hirtinewis Tjeder; see Meinander, 1972), Hemerobiidae (see below), Ithonidae (an Macroptery and the ability to fly are plesiomorphic within the undescribed species from Mexico; E. G. MacLeod, personal insect superorder Neuropterida - orders Neuroptera (lacewings communication, specimen seen), and Nemopteridae and allies), Megaloptera (alderflies and fishflies) and (Stenorrhachus walkeri (McLachlan); Stange, personal Raphidioptera (snakeflies) - and flightless taxa are rare within communication, specimen seen). In the families Berothidae, this group of largely predatory insects. Only one record of a Coniopterygidae, Ithonidae and Nemopteridae flightlessness flightless megalopteran has been traced - a micropterous female has been reported only in females. In these families, species corydalid probably belonging to the genus Platychauliodes with flightless females are known or suspected to be sexually [possibly an aberration; see Barnard (1940)l. Flightlessness is dimorphic, with fully macropterous volant males. Only within not definitely known in the order Raphidioptera, although the family Hemerobiidae are species presently known that are doubts have been expressed about the flight abilities of the flightless in both sexes. females of two inocelliid species based on their relatively small This paper describes a new brachypterous species of the wings (i.e. Inocellia fulvostigmata Aspock & Aspock, see hemerobiid genus Nusalala and reviews the phenomena of Aspock et al., 1991, and Indianoinocellia mayana Aspock, wing reduction and flightlessness in the family Hemerobiidae. Aspock & Rausch, see Aspock et al., 1992:175). Within the Four topics related to the latter theme are discussed. First, the order Neuroptera, taxa inferred to be flightless on the basis of taxonomic and phylogenetic positions of flightless hemerobiids brachypterous, micropterous or apterous wing states are known are surveyed. Second, the morphological evidence for in the families Berothidae (i.e. Trichoma gracilipenne Tillyard; flightlessness in hemerobiids is examined. Third, the available see Aspock & Aspock, 1985), Coniopterygidae (e.g. data on habitat preferences of flightless hemerobiids is reviewed. Finally, factors that may have contributed to the Correspondence: Dr John D. Oswald, Department of Entomology, evolution of flightlessness in hemerobiids are considered and Texas A&M University, College Station, TX 77843-2475, U.S.A. discussed. O 1996 Blackwell Science Ltd 343 344 John D. Oswald Figs 1-7. Nusalala brachyptera, n.sp. [holotype]. 1, Left forewing (converted to right dorsal view). 2, Left hindwing (converted to right dorsal view). 3, 9th tergite and ectoproct, lateral. 4, parabaculum and laterobacula, dorsal. 5, same, lateral. 6, gonarcus complex, dorsal. 7, same, lateral (arrow points into compressed space separating neo- and extrahemigonarcus). Abbreviations: 9t, 9th tergite; ect, ectoproct; egpp, extragonopontal process; ehgp, extrahemigonarcal process; ehgs, extrahemigonarcus; igs, intragonarcus; ltb, laterobaculum; med, mediuncus; ngps, neogonopons; nhgs, neohemigonarcus; pa, parabacular apophysis; tl, terminal lobe. Nusalala brachyptera sp.n. (Figs 1-7) Forewing (Fig. 1). Brachypterous, 3.54.5 mm long (tegula to apex), oval and coriaceous; coloration variable, mostly Diagnosis. Differentiated from other Nusalala species by testaceous with scattered darker markings, e.g. some crossveins the following combination of characters: (i) Forewings and along wing margin; venation reduced and slightly irregular; coriaceous and brachypterous (Fig. 1); (ii) hindwings costal space narrow, with or without a few (1-3) crossveins micropterous and scale-like, venation vestigial (Fig. 2); (iii) (not veinlets) present, pterostigma absent; subcostal space with male ectoproct with a prominent posteroventral spine (Fig. 3). crossveins irregular in number and position; radius with 4-6 The first two characters will differentiate brachyptera from all 'prestigmal' oblique radial branches; radial space with 34 described species of Nusalala except andina Penny & Sturm. irregular gradate series; bases of media and radius fused; These two brachypterous species are easily separated by the M3 + 4 not always (e.g. paratypes) fused to Cu as in other presence of a distinct posteroventral male ectoproct spine in Nusalala species. Hindwing (Fig. 2): micropterous, apex brachyptera (absent in andina). The detailed conformations of truncate, venation vestigial. the extragonarcal processes (extrahemigonarcal, extragono- pontal and mediuncus) and laterobacula of brachyptera are also diagnostic (Figs 4-7). Male terminalia (holotype). 9th tergite (Fig. 3, 9t): divided sagittally into a pair of hemitergites; apodemes of Description. (Holotype and paratype males; morphological proximoventral hemitergite angles prominent. Ectoprocts terminology after Oswald, 1993a.) (Fig. 3, ect): indistinguishably fused to ipsilateral 9th O 1996 Blackwell Science Ltd, Systematic Entomology, 21, 343-352 Flightlessness in brown lacewings 345 hemitergites ventrally, but free dorsally; posteroventral spine on Cerro de la Muerte, 7.vi.1992, 2750 m [9000'], ex. litter, short but prominent. Gonarcus (Figs 6-7): intragonarcus (igs) M. L. Jameson (CAS, in alcohol). extensive, narrowed dorsally and extending on to neogonarcus, ventral margin with a rounded indentation, no clear Other Material (1 9). COSTA RICA: Cartago Prov.: 1 9 3 differentiation between intragonopons and intrahemigonarcus; mi. [5 km] S. Empalme, 10.vii.1974, 2620 m [86007],O'Brien, extragonarcus highly modified; each extrahemigonarcus (ehgs) O'Brien & Marshall (CAS, pinned). Because this specimen produced posterodorsally as a stout glabrous processes (ehgp) was not collected in association with an identifiable male it is and posteroventrally as a rounded lobe, margins of specifically excluded from the type series and is only extrahemigonarcus inwardly revolute below posterodorsal provisionally placed with this species. processes; extragonopons composed of a pair of slender acuminate processes (egpp) laterally and a median mediuncus Etymology. From Greek brachys, short, and pteron, feather (med); bases of extragonopontal processes connected medially or wing, in reference to the species' abbreviated wings. by a tenuous sclerotized strap; mediuncus attached to distal margin of strap by a transverse membranous articulation; Discussion. The genus Nusalala (Navis, 1913) currently neogonarcus present; neogonopontal region (ngps) narrowed contains ;= 20 valid species and is endemic to South and and articulated sagittally in dorsal view, projectigg Central America and the West Indies (Monserrat, 1990). The conspicuously over base of extragonopons in lateral view; genus is in need of a comprehensive revision, and the discovery neohemigonarcus (nhgs) present but inconspicuous, adpressed of additional new species is expected. Flightlessness and to proximodorsal surfaces of extrahemigonarcus. Parabaculum brachyptery was first noted in Nusalala by Penny & Sturm (Figs 4-5): parabacular apophysis (pa) a well-developed, (1984) in the Colombian species andina. Nusalala brachyptera arcuate, vertical plate; terminal lobe (tl) apices splayed, is the second flightless species to be described in this genus. principal sclerotization of lobes on medial faces, lateral faces The single female specimen cited above was collected ;= 20- largely membranous, lobe surfaces adjacent to laterobacula 30 km WNW of the two male specimens. Its tentative forming a longitudinal v-shaped channel. Laterobacula (Figs 4- identification as brachyptera is supported by its similar wing 5; ltb): glabrous, apices revolute, each with a small process or modifications, high elevation collection site in the Cordillera lobe at distal end of articulation with parabaculum. Hypandrium de Talamanca, geographical proximity to the two known males internurn: present. Variation: One of
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