View Image Gallery of Tribe Macariini

This tribe embraces the concept of the Macaridae of Guenée (1857), a name that should take priority (as Macariini) despite the extensive usage of Semiothisini, probably first employed by Warren (1894; as Semiothisinae).

Packard (1876) merged Guenée's Macaridae and Zerenidae with his Fidonidae as the Fidoninae (more correctly Fidoniinae). This is based on the Fidonites of Duponchel, typified by Fidonia Treitschke. Fidonia is a junior objective synonym of Eurrhanthis Hübner, having as common type species the Mediterranean plummistaria Villers. This is distinct from the concept of Semiothisini of Forbes (1948) as it backs the definitive features discussed below.

The Macariini also embrace the Fernaldellinae of Hulst (1896), as Fernaldella Hulst is placed as a semiothisine by McGuffin (1972).

Forbes referred to male genitalia with two setal spines (or horns) at the tip of the uncus and a deeply emarginate or forked valve. The chaetosemata are transversely elongate, tapering towards the centre of the head through a parting of the head scales. The antennae are bipectinate, serrate or filiform with cilia. The forewing is often foveate in a characteristic manner. There is a transverse comb of setae on sternite 3 of the male.

No single character can be used to define the tribe but a combination of the two genitalic features mentioned and that of the chaetosemata will enable members to be identified reliably. The chaetosema character is, however, present in Scardamia Guenée of the Scardamiini (See Scardamiini), a probable homoplasious occurrence given the weight of other characters. The gnathus tends to be strong, expanded deeply at the apex.

The development of the male forewing fovea requires further study across the tribe. It is strongly developed in most S.E. Asian taxa that have been attributed to Semiothisa, and is illustrated in Fig 1. The base of vein A1 is sigmoid, its anterior convexity containing a corrugate tymbal structure; this is bounded distally also by a spur from the posterior vein enclosing the cell (CuA). The vestige of A2 is strongly curved, running parallel to, but mostly well separate from, the basal curvature on Al. In Macaria and Semiothisa often only the spur from the cell is evident, but this is variable within these genera (Hua Baozheng, pers. comm.).

Normal chaetosemata are present in Lampadopteryx Warren, Hypephyra Butler and Iridoplecta Warren, but the first two have the uncus horned, and all have the valve forked, albeit weakly in Lampadopteryx. There is no fovea in these genera, hence its degree of development within the tribe will in general be apomorphic.

The apomorphic state of the chaetosemata occurs in a number of moderate to large genera such as Itame Hübner, Semiothisa Hübner, Godonela Biosduval, Tephrina Guenée and Oxymacaria Warren, as well as smaller ones such as Chiasmia Hübner, and Parosteodes Warren. However, the definition and status of these genera is far from satisfactory. Inoue (1986) indicated that the type species of Semiothisa, the South American gambaria Hübner, has features that set it apart from the Holarctic and Old World tropical taxa currently placed in the genus. It has the costal arm of the male valve slender, curved, apically acute, and lacks uncus horns. The male antennae are serrate. He placed a number of species lacking uncus horns in Heterocallia and referred Oriental taxa with horns and with filiform male antennae to Chiasmia.

In order to resolve this question a little further to determine the status of Bornean taxa, males of all relevant generic type species (as in Fletcher (1979)) were dissected. In addition, numerous N. American taxa have been featured by McGuffin (1972), and there is a review of a N. American conifer feeding complex (referable to the generic name Sciagraphia Hulst) by Ferguson (1974).

Important features appear to be the presence or absence of uncus horns, the state of the distal margin of the eighth sternite, condition of the male fovea, and, to some extent, whether male antennae are bipectinate or filiform. Secondary loss of uncus horns and the degree of pectination of the male antennae are probably somewhat homoplasious, with eighth sternite characters providing the most useful distinction.

The sternite is unmodified in Heterocallia, Oxymacaria and Parosteodes. The male antennae are filiform in the first, bipectinate in the second and serrate in the third. Only in Parosteodes are uncal horns present. In all three the saccular section of the valve is small, without ornament. Despite the antennal differences, Heterocallia and Oxymacaria show common features such as a slightly bifalcate forewing shape, a moderately strong fovea, and a row of small white marks over the anterior part of the submarginal. There are thus grounds for bringing them into synonymy . The N. American Trigrammia Herrich-Schäffer has filiform antennae and genitalic features as in Heterocallia, but is markedly different in facies characters. Mellilla Grote and Eumacaria Packard, with bipectinate antennae, also have distinct facies in this category.

All other genera examined have a pair of processes from the eighth sternite. These are large, rounded lobes in Semiothisa and in some N. American taxa referable to Digrammia Gumppenberg (McGuffin, 1972: californiaria and hebetata groups). In the latter the uncus is apically narrowed, with the horns vestigial or lost, and there appear to be diagnostic features of the female genitalia: a bilobed sterigma and loss of the signum typical of most Macariini. The valves tend to have the saccus portion produced, angular, but the sternal lobes tend in some taxa towards the status described below. The male antennae are filiform.

The genera Tephrina, Gnopharmia, Isturgia Hübner and typical Itame (vincularia Hübner) lack uncus horns and have strongly bipectinate male antennae. The eighth sternite has a pair of processes, usually apically sclerotised, separated from each other by only a narrow, shallow cleft except in Isturgia where it is deeper, rounded, and Gnopharmia where it is broad. The N. American Itame taxa illustrated by McGuffin (1974) have small uncus horns, a strong valvula (a central lobe between the arms of the valve) and rather broad sternal processes separated by a strong but narrow cleft. Their status needs further investigation.

The bulk of taxa currently in Semiothisa should probably be reassigned to other genera as suggested by Inoue, e.g. Macaria Curtis, as suggested by Ferguson (in litt.). The extent of the true Semiothisa and of Macaria in the Neotropics is reviewed by Hua & Scoble (in preparation).

Macaria (embracing also Philobia Duponchel, Eutropa Hübner and Sciagraphia Hulst) includes species with sternal processes rather acute, separated by a deep, basally narrow cleft. The uncus horns are small. There is no valvula, and the saccular portion of the valve is rounded, with submarginal ornamentation usually lobes and flanges. The concept embraces many Holarctic (e.g. the ulsterata and signaria (Sciagraphia) groups of McGuffin) taxa and a number of Neotropical ones. Male antennae can be serrate or ciliate. The fovea is not well developed.

Inoue's (1986) concept of Chiasmia includes species with short, heavily sclerotised sternal processes, separated by a shallow, moderate to broad cleft (but much reduced in Automolodes and Iulocera Warren). The male antennae are filiform. The uncus horns are mostly strongly developed. The saccular process of the valve is diversely developed: it is variably acute, often ornamented and accompanied by a valvula in the robust species of the Evarzia Walker (= Gubaria Moore) group, well represented in Borneo. This concept embraces the generic names Godonela Boisduval, Acadra Herrich-Schäffer, Evarzia, Gubaria, Iulocera, Automolodes, Allochrosis Strand and Thyridesia Wehrli. A facies feature that recurs through the group is an angled forewing postmedial with a pale mark set just anteriorly and distally to the angle (seen clearly here in albipuncta Warren). Typical Chiasmia has a ventral angular process to the costal arm of the valve: a similar feature is seen in the type species of Allochrosis and in albipuncta).

However, a major problem with this concept of Chiasmia is that typical Chiasmia lack a fovea on the male forewing, whereas it is very strongly developed in most Indo-Australian taxa. The facies of typical Chiasmia is highly modified into a striking reticulate pattern. Hence a more appropriate generic combination must be sought. Godonela Boisduval is the oldest name where the type species has a strong fovea, hence this should be applied to foveate taxa within Inoue's concept of Chiasmia. All the generic names mentioned above fall within this concept apart from Chiasmia.

Given the paucity of strong apomorphic features, the decisions made here on placement of species within genera may be subject to revision in future when a more rigorous, global, morphological survey can be undertaken. Current work on African macariines by M. Kruger is likely to throw further light on the situation.

Azata Walker (type species idriasaria Walker, the type a female without locality data) was associated by Fletcher (1979) with Indian material related to the Sundanian species mutabilis Warren and allies , but the facies of the type specimen is atypical, resembling more S. American taxa in true Semiothisa, particularly an unnamed series from Jamaica in the NHM (Hua Baozheng, pers. comm.).

Chiasmiodes Warren Gen. rev. is not a macamine. The male genitalia of the Himalayan type species variolinea Warren have features more akin to those of Lithinini.

Eggs and larvae of N. American taxa are described by McGuffin (1974). Arboreal feeding is general. Pupation is at the surface of the soil in a slight cocoon. Host-plants are diverse in temperate regions, including several conifer-feeding taxa or groups. In the tropics there is some preference for Leguminosae, particularly Acacia, Albizia and allies.

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