Nacophorini Lithinini Lithinini I. apicata Lithinini M . biplaga M . biplaga Lithinini Geometridae Scionomia Warren ( Sato 1977 ) Ennomini M . biplaga ada biplaga I. apicata Nacophorini biplaga Nacophorini Lithinini Molecular relationships of the Australian Ennominae (Lepidoptera: Geometridae) and implications for the phylogeny of the Geometridae from molecular and morphological data Young, Catherine J. Zootaxa 2006 2006-07-17 1264 1 1 147 [405,515,523,549] Insecta Geometridae Animalia Lepidoptera 68 Arthropoda tribe Lithinini  —swollen hind­tibia; U­shaped gnathos (Fig. 113); absence of discrete cornuti (Fig. 114); relatively small corpus bursae; absence of signa; inner surface of corpus bursae studded with small spicules (Fig. 115). The eggs of the two species of  Metrocampaare rather nacophorine in appearance with a broad, dorso­ventrally flattened form, irregularly arranged cells, narrow to moderately broad, recessed walls and inconspicuous aeropyles (Fig. 116) (Young, in press). They do not resemble the eggs of  I. apicata(Fig. 112) or other lithinine species ( Salkeld 1983). Shared larval features are as follows: —fern feeders; moderately stout to stout; short setae; long setae on first instar; SV1, SV3 and V 1 invertical alignment on A1; absence of extra prolegs; crochets a biordinal interrupted mesoseries although incompletely interrupted in  I. apicatain mature larvae; acuminate setae. The presence of moderately long to long setae in the first instar larvae is typical of the Australian Nacophorini(unpubl. data). Long setae were also found in  Amelora,  Capusa, Chlenia,  Fisera,  Mnesampela,  Plesanemmaand  ThalainaWalker. Howeverthis feature was also present in the asthenine  Poecilasthenaand the noctuids  ProteuxoaHampsonand  AgrotisOchsenheimer. Differences between the larval morphology of  M. biplagaand  I. apicata: —only four lateral setae present on A6; L3, SV1 and V1 aligned vertically on A3–5; anal claspers large and prominent; crochet arrangement in the first instar larva a uniordinal interrupted mesoseries, in  M. biplagaonly. Behavioural differences between the two species is the tendency of  I. apicatato drop, without the aid of silk, and rest in a coiled position when disturbed. On the other hand,  M. biplagalarvae are stout and sluggish and conceal themselves while clinging to the undersides of fronds when disturbed. Adult resting positions also vary. The adults of  Idiodesrest in typical geometrid fashion with all wings exposed in an open planiform position whereas  Metrocampaadopts the more atypical closed planiform position in which the wings are held in a horizontal plane but with the hindwings covered ( McFarland 1988). Shared pupal features are as follows: —setae very short; exposed labium; concealed meta­tibia; pro­thoracic spiracle weak in  I. apicataand absent in  M. biplaga; no A5 spiracular development; dorsal furrow welldeveloped in both species, deeply excavated in  M. biplaga; lateral furrow present in both species but weak in  M. biplaga; four pairs of hooked cremastral setae, terminal pair robust. Unlike  M. biplaga,  I. apicatahas a smooth rather than rugose cuticule and exposed fore­femora. The punctation in both species is also noticeably different. In  I. apicatathe punctation is uniformly small, shallow, dense and randomly distributed on A1–7 whereas  M. biplagahas large, deep punctures arranged in two rows on the dorsum of A1–3, and then similar punctation to  I. apicataon A4–8.  The Nacophoriniand Lithininihave been linked on the basis of the probable homology of the processes of the anellus found in both tribes and also the curved edges of the juxta that often bear cristate hairs ( Rindge 1986; Pitkin 2002). Rindge (1986)in his review of New World Lithininifound difficulty in defining apomorphies for the tribe. However Holloway (1987)defined the tribe on the following features: projected cucullus; presence of processes of the anellus, apices of processes often bearing spines; valvae directed dorsally; triangular transtilla; uncus long and slender; weak, setose socii; central signum; pleated, short ductus bursae.  I. apicatapresents all of these features (Figs 109, 111); however none, apart from possibly the modified cucullus, are unique to the Lithinini. The genitalia of  M. biplagaare not typically lithinine due to the absence of a projected cucullus and signum (Figs 113, 115). It is probable that  M. biplagahas been misplaced into the Lithininidespite being a fern­feeder. Non­lithinine fern­feeders in the Geometridaeare few, as far as I know, but one example is the genus  ScionomiaWarren ( Sato 1977), which is placed in the Ennomini. The genitalia and eggs of  M. biplagaand its congener  adalack typically lithinine features and the larvae of  biplagaare differentiated from  I. apicataon several important features. Further, the presence of a combination of all three malesecondary sexual characteristics, A3 pecten, inflated hind­tibia and hair­pencil on the hind­tibia in this species were only found in the Australian Nacophoriniin this study (unpubl. data) and is additional evidence of a link between  biplagaand the Australian nacophorines. Accordingly, both lithinine genera have been placed within the Australian Nacophoriniby the molecular analysis (28S D2) (Fig. 10) and it is possible that the two tribes could be united, as the monophyly of the Lithininiis only weakly supported on morphology.