Macromiidae Needham 1903

Genera of Macromiidae The genera of Macromiidae were established in 19th century according to the taxonomic concepts of that time which, in particular, overestimated importance of the venation. The chronologically first genus Epophthalmia was described by Burmeister (1839) to include many species currently considered in seven genera of three families. He did not indicate the type species, which was subsequently designated as being E. vittata by Hagen (1867: 62), who cited the personal communication by Burmeister: “Del’ Gattungsname Epophthalmia 1st auf den persönlich ausgesprochenen Wunsch Prof. Burmeister’s seiner E. vittata belassen [The genus name Epophthalmia was retained at the personal request of Prof. Burmeister’s for his E. vittata.]”. Subsequent authors used to include in Epophthalmia in the narrowed sense also some other large species, including some North American ones, all placed in Macromia since the paper by Williamson (1909). The present content of Epophthalmia as including five Asian species has been established after the comprehensive revision by Lieftinck (1931). The genera Macromia and Didymops were simultaneously established by Rambur (1842). Macromia was proposed for five new species described in the same work, of which only three remained in this genus until now, one was later moved to Epophthalmia and one to Phyllomacromia. The type species was not indicated and was later designated by Kirby (1890). He was aware of the importance of type species of genera and so made such designations for all genera known to him as missing the type species. For this purpose, he usually chose species listed first in the original description of a genus, so he designated Macromia cingulata Rambur, 1842 as the type species of the genus Macromia. This choice appeared to be not so good because this species is more peculiar than ‘typical’ among its congeners for being the smallest, having unusually short and broad hamuli in the male secondary genitalia, and very extensive yellow markings (Fraser 1936). Moreover, the holotype female (the only specimen which the original description was based on) was mislabelled, even with respect to the continent. Rambur (1842: 138) indicated its origin as follows: “Collection de M. Serville, òu elle est placée parmi les AEschna, et étiquetée sous le nom que je lui ai conservé; indiquée de l’Amérique septentrionale [Collection of M. Serville, where it is placed among Aeschna, and labeled under the name that I have kept for it; indicated from North America]”. Selys (1874) clarified that this was in error and supposed that the true type locality was ‘Bengal’. This species occurs in India, Pakistan and Nepal (Fraser 1936; Kalkman et al. 2020). The genus Didymops was proposed for the sole species Didymops servillii Rambur, 1842, described in the same work (Rambur 1842). This name was later treated as a junior synonym of Libellula transversa Say, 1840. Hence, the species under the currently valid name Didymops transversa is the type species of the genus by monotypy as to the original description (the only one mentioned therein). It is noteworthy that Selys (1871) considered Macromia as the valid name for the genus and subgenus and Didymops as its synonym, even at the subgeneric level. Thus, according to the current ICZN Art 24.2, he acted as the First Reviser who chose the valid name among simultaneously published synonyms. Although seven years later he (Selys 1878) restored the name Didymops to denote a valid subgenus, this decision persists whenever these two names are considered synonymous. The name Phyllomacromia was first proposed by Selys Longchamps (1878) for a new subgenus including two African and one Madagascarian species exhibiting foliaceous broadenings of the end of abdomen. Its type species was subsequently designated again by Kirby (1890) to be Macromia trifasciata Rambur, 1842, most probably since it was chronologically first described. It is the only species of this genus occurring in Madagascar and absent from the African continent. F.C. Fraser, who was the actual author of the text of Tillyard & Fraser (1940), wrote: “several species of Macromia have been described from Africa, but an examination of their genitalia, which I carried out recently, proves them all to belong to Phyllomacromia ” (Tillyard & Fraser 1940: 390). For some reason, in his later work, Fraser (1954) reclassified the African species under Macromia. However, May (1997) restored Phyllomacromia as a genus including all African and the only Madagascarian species (the type species of the genus) of Macromiidae on the basis of his thorough morphological analysis, first of all concerning the characters of the male vesica spermalis and cerci. This treatment is unequivocally accepted at present. The genus Azuma was proposed for the only nominal species Macromia elegans Brauer, 1865, currently Epophthalmia elegans, based on venation characters (Needham 1904). Later Williamson (1909) found these characters unreliable. Ris (1916) assumed the names Azuma and Epophthalmia as synonyms but some subsequent authors used the former as valid since they considered the latter to be a junior synonym of Cordulia Leach, 1815, because of its above mentioned too broad sense when it was proposed by Burmeister (1839). At last, the name Azuma was claimed to be a junior synonym of Epophthalmia (as defined by its type species) by Lieftinck (1931). Additionally, the name Azuma turned out to be a junior homonym of a genus of fishes (Lieftinck 1931). Species grouping The large number of species in the genera Macromia and Phyllomacromia and their obvious morphological diversity in their current senses inspired repeated attempts of their grouping. To avoid confusion of the informal species groups with species, further in the text we will designate the species groups with a specific name only, e.g. “ calliope - group” rather than “ M. calliope group”, unless in verbatim citations. It should be noted that all attempts of grouping in Macromia concerned numerous Asian species but did not involve American species. Laidlaw (1922), based on the marking of the face and mesepisternum and the shape of S10, subdivided the Asian species into three groups: westwoodi -, cincta - and calliope - groups. Fraser (1924) added the fourth, cingulata -group. However, the latter looked unnatural as being too heterogeneous with respect to the male secondary genitalia, as followed from illustrations provided by Fraser (1924; 1936) himself. Lieftinck (1929) added the moorei -group, isolating it from Laidlaw’s calliope -group. The grouping by Lieftinck (1929) was most broadly accepted for a long time. Thus, up to 1930s, two grouping systems of Macromia had been proposed, of which Fraser’s concerned the Indian species and Lieftinck’s concerned mostly Malesian/Australasian species. Both were broadly accepted for the respective regions but underwent further independent splitting. It should be noted that Lieftinck considered his groups very informally, not as quasi-taxonomic entities of some more or less definite rank. For instance, in his later paper he mentioned that “ urania clearly belongs to the gerstaeckeri group” (Lieftinck 1950: 702), without defining that group, and then wrote that “the groups defined earlier were admittedly artificial”. Then he (Lieftinck 1955) provisionally attributed five North Asiatic species to the amphigena -group (isolating it from his moorei -group), which included Macromia amphigena, but did not provide the diagnosis of this group. At last, Lieftinck (1971) considered “the Papuasian group of Macromia ”, composed of species “very large alike and obviously closely interrelated”, which he further split into three groups: “Group I of M. terpsichore Foerster ”, “Group II of M. melpomene Ris ”, and “Group III of M. chalciope Lieftinck ” (Lieftinck 1971: 30–32). However, according to his earlier seminal work (Lieftinck 1929), Macromia terpsichore Förster, 1900 and M. melpomene Ris, 1913 were members of the “I Group of M. westwoodi Selys ”, hence all three of his later Papuasian ‘groups’ were actually subdivisions of his primary westwoodi -group. However, later authors assumed Lieftinck’s groups rather seriously. The arachnomima -group was independently proposed by Wilson (1993) and Muraki (2010). However, both authors did not consider the fact that Macromia arachnomima Lieftinck, 1953 is very similar to M. aculeata Fraser, 1927 (Kosterin 2015; Muraki, 2021), which Fraser (1927) had classified under his cingulata -group. Muraki (2004) isolated the urania -group from the calliope - group, although this could be considered the gerstaeckeri -group in the sense by Lieftinck (1950). Muraki (2009) mentioned additional groups, namely daimoji -group and vangviengensis -group, and an unnamed group for an undescribed species, with a reservation that this grouping had not been published yet. Muraki (2014) provided short diagnoses and lists of included species for seven groups, which were the four groups by Lieftinck (1929) plus urania -group, arachnomima -group, and vangviengensis -group by Muraki, but for some reason did not mention his daimoji -group in that paper (Muraki 2014). Such over-splitting of the SE Asian representatives of the genus in groups seems to be more misleading than helpful. The Fraser’s grouping system was recently reconsidered by Sadasivan et al. (2023) who, based on the same ‘Fraser’s’ characters of the male cerci and secondary genitalia structures, isolated M. ellisoni from Fraser’s cingulata - group into the ellisoni -group of its own. They also split the Western Ghats species of cingulata -group into two subgroups, flavicincta-bellicosa-irata - and cingulata - subgroups. Isolation of the latter species, peculiar for its small size and short, broad hamuli, looks reasonable. The grouping within Phyllomacromia spp. was also repeatedly addressed (Gambles 1979; Legrand 1992; 1993; Dijkstra 2005). Eleven groups were recognise