Macariini, GUENEE, [1858]

Scoble, Malcolm J. & Krüger, Martin, 2002, A review of the genera of Macariini with a revised classification of the tribe (Geometridae: Ennominae), Zoological Journal of the Linnean Society 134 (3), pp. 257-315 : 262-263

TRIBE MACARIINI GUENÉE, [1858]

Fernaldellini Hulst, 1896

Semiothisini Warren, 1894

The tribe is recognizable by the presence of one or more of three characters. These are: chaetosemata extending across the head, either completely or partly; uncus with a pair of enlarged setae (horns) at or near the apex; and valva divided typically into two fairly broad lobes (unlike the condition in Cassymini and Eutoeini ), although modifications to the lobes do occur (e.g. as in Semiothisa and Oxymacaria ). In many supraspecific groupings, abdominal sternum A8 of the male is modified – cleft, excavated, or emarginated. It has been suggested by Holloway ([1994]: 155; fig. 1) that the form of the fovea, where present, might be distinctive for the tribe, although this requires confirmation based on further examiniation across the Ennominae .

Within the tribe are three main groupings, not necessarily monophyletic. Indeed, so loose are these grouping that we have not made headings in the descriptive section of the text.

The Platypepla-group of genera

( Platypepla to Narraga )

This group is probably monophyletic. The moths are small to medium, ochreous to brown, and often have a falcate forewing. The uncus is weakly to strongly setose in most species, but it bears strong spines in two species of a new genus ( Krüger, 2001). The valva is deeply, almost entirely, divided and frequently has spine-like setae on the costa. The saccus is usually elongated. In the female, the corpus bursae appears short in relation to the length of the apophyses and the ductus. Usually the corpus is round and covered densely with spines, but sometimes spines are absent entirely. Most species occur in the Afrotropics, including Madagascar, but there are Holarctic representatives. Two Australian species listed under Paramelora by McQuillan & Edwards (1996) have a valva divided as deeply as that in Platypepla and its relatives. However, the bursa copulatrix is not densely spinose and the affinity with Platypepla is not established.

Genera lacking horns on the uncus

( Heliomata to Semiothisa )

This assemblage is not monophyletic. Indeed, some doubt is expressed below as to whether Heliomata belongs to the Macariini . Among the grade of genera lacking uncus horns are several groups of species distinguishable on the basis of male genitalia and wing shape, colour or both. We have applied existing generic names to these groups, despite the fact that the male genitalia are broadly similar in valva shape and, to a lesser extent, the shape of the uncus and of the gnathos.

A particular difficulty was encountered with the genera Isturgia , Bichroma , Enconista and Tephrina . The type species of all four differ in wing colour and pattern. In wing shape, however, they are all much more similar to each other than they are to Mellilla , even though Mellilla has very similar genitalia. The shapes of the uncus and gnathos are so similar in Isturgia , Bichroma and Enconista that they appear to form a monophyletic group. These genera are synonymized in this work. Further similarities between Isturgia and Bichroma in habitat, foodplant and larval appearance were noted by Rödel & Trusch (1997). The uncus and, particularly, the gnathos of Tephrina differ in some species, but it seems likely that these structures are modifications of the Isturgia pattern. If that is the case, then Isturgia would be rendered paraphyletic without Tephrina . After much agonizing we have therefore also synonymized Tephrina with Isturgia . The recorded foodplants of all these genera (including Tephrina ) belong almost exclusively to the Papilionoidea (Leguminosae ( Fabaceae )).

Itame , which includes just vincularia and teknaria , is retained as a valid genus since the male genitalia are sufficiently distinctive from the genera mentioned above. I. vincularia is recorded from Rhamnaceae not, as is the case in Isturgia s. str., Fabaceae .

The other genera lacking uncus horns, and which are somewhat more remote from Isturgia but with broadly similar male genitalia, are Trigrammia , Mellilla (externally distinctive), Dissomorphia and Paramelora . The phylogenetic relationships of these genera, individually, to Isturgia (expanded sense) remain unclear, and we have not synonymized them with Isturgia .

Genera with horns on the uncus ( Macaria to Hypephyra )

Most species of Macariini have uncus horns, for these structures occur in the two species-rich genera Chiasmia and Macaria . It is unclear if horns are basal to the macariine clade and lost on one or more occasions, or if they appeared later in the evolution of the group. It is unlikely, however, either that all genera with horns form a monophyletic group. For example, although Semiothisa lacks horns, and differs from Macaria in yet other features of the male genitalia, the two genera both share a cleft condition of sternum A 8 in the male and a similar wing shape and pattern in many species. Further discussion of macariine phylogeny is given by Krüger (2001).