identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
327687E4FFEAFFBDFF32886DFD27E392.text	327687E4FFEAFFBDFF32886DFD27E392.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Macromiidae Needham 1903	<div><p>Macromiidae as a family</p><p>The taxonomic history of the family  Macromiidae and its genera is quite complicated and confused. According to the principle of coordination by International Code of Zoological Nomenclature (ICZN) Art. 36.1 (International Commission on Zoological Nomenclature 1999), proposal of a family group name in either rank automatically establishes it in all other ranks of the family group. The family group name based on the genus  Macromia was first proposed as the subfamily  Macromiinae by Needham (1903), who just mentioned this name, for the first time in scientific literature, in the text as follows: “In the  Macromiinae, however, it [the antecedent trachea in the larval wing] is formed by a modification of the latter type, as shown for  Didymops transversa ” (Needham 1903: 711) . Here the family group name was mentioned as obviously and properly derived from the genus  Macromia, which hence was its type genus, and the second genus,  Didymops, was explicitly included into that subfamily. At that time both these genera were considered in the family  Corduliidae, hence  Macromiinae was proposed as belonging to it. Needham apparently intended to propose the new subfamily name in another work of the same year (Needham &amp; Hart 1903), which contained a taxonomic paragraph in its introduction where families and subfamilies were enumerated. However, another name, Synthemiinae, was used there instead of  Macromiinae . In a later work he provided under the subtitle “subfamily  MACROMIINAE ” the following footnote “The use of the name Synthemiinae for this subfamily in Bull. 111. State Lab. of Nat. Hist, VI, p. 5, was due to enforced haste in printing, whereby proof corrections made by me were not received by the printer in time for incorporation into the text.” (Needham 1904: 698).</p><p>Until quite recently (see e.g. Bridges 1994) it was erroneously considered that a family group name based on  Macromia was first introduced by Tillyard (1917), as the tribe  Macromiini of the subfamily  Corduliinae including the genera  Azuma Needham, 1904,  Didymops,  Epophthalmia,  Macromia and  Phyllomacromia (see his table inserted after page 282). Later Tillyard &amp; Fraser (1940) (this monograph was completed by the second author) proposed another name Epophthalminae Tillyard &amp; Fraser 1940, based on a different genus, for their new subfamily including  Epophthalmia,  Didymops,  Macromia,  Macromidia and  Phyllomacromia . This subfamily corresponded to the current family  Macromiidae with the only exception of the inclusion of  Macromidia . The same classification was reproduced in Fraser (1957).</p><p>Gloyd (1959) was first to suggest the family rank of  Macromiidae, to include  Didymops,  Epophthalmia,  Macromia and, tentatively,  Macromidia . The genera  Epophthalmia,  Didymops,  Macromia and  Phyllomacromia comprising the family  Macromiidae in its current sense (with  Macromidia excluded) (Dijkstra et al. 2013) were shown to form a monophyletic group based on morphological analysis by May (1997) (who still considered them as the subfamily  Macromiinae) and molecular analysis by Ware et al. (2007) (which did not involve  Epophthalmia), Dumont et al. (2010) and Carle et al. (2015).</p></div>	https://treatment.plazi.org/id/327687E4FFEAFFBDFF32886DFD27E392	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kosterin, Oleg E.;Vierstraete, Andy;Schneider, Thomas;Kompier, Tom;Hu, Fang-Shuo;Everett, Larry;Makbun, Noppadon;Onishko, Vladimir V.;Papazian, Michel;Dumont, Henri J.	Kosterin, Oleg E., Vierstraete, Andy, Schneider, Thomas, Kompier, Tom, Hu, Fang-Shuo, Everett, Larry, Makbun, Noppadon, Onishko, Vladimir V., Papazian, Michel, Dumont, Henri J. (2025): Molecular phylogenetic analysis of the family Macromiidae (Odonata) worldwide based on a mitochondrial and two nuclear markers, with a short overview of its taxonomic history. Zootaxa 5620 (4): 501-545, DOI: 10.11646/zootaxa.5620.4.1, URL: https://doi.org/10.11646/zootaxa.5620.4.1
327687E4FFEAFFADFF328F59FEBAE781.text	327687E4FFEAFFADFF328F59FEBAE781.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Macromiidae Needham 1903	<div><p>Genera of  Macromiidae</p><p>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).</p><p>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).</p><p>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).</p><p>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.</p><p>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 &amp; 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 &amp; 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.</p><p>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).</p><p>Species grouping</p><p>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.</p><p>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.</p><p>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.</p><p>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.</p><p>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.</p><p>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.</p><p>The grouping within  Phyllomacromia spp. was also repeatedly addressed (Gambles 1979; Legrand 1992; 1993; Dijkstra 2005). Eleven groups were recognised in the latest of these works, with a note that they “strictly serve convenience and (as yet) have no phylogenetic basis, conveying jizz rather than kinship” (Dijkstra 2005: 23).</p><p>Phylogenetic reconstructions</p><p>Histone H3–H4 region tree</p><p>The phylogenetic tree reconstructed from sequences of the histone H3–H4 region is shown in Fig. 2. The further text will mention branches with indication of their support as posterior probability values in brackets. All but three sequences of the histone H3–H4 region were obtained in the course of this study, so the set of species in this tree is mostly confined to our species sample.</p><p>In this tree, the  Macromiidae family is monophyletic with the perfect support (1.00). At the same time the genera  Macromidia,  Idionyx (both being well-supported monophyletic clusters) and  Oxygastra do not form a monophyletic clade, as might be expected. Together with  Macromiidae, they form a major clade with a high support (0.92), which does not include  C. aenea . In this clade,  Oxygastra curtisii (Dale, 1834) reliably clusters with  Macromidia (0.90), while association of their joint cluster with  Macromiidae is unsupported (0.53). The genus  Idionyx diverges into two clusters, one of which (0.85) includes three species from the Western Ghats of India ( Idionyx corona Fraser, 1921,  I. saffronatus Fraser, 1924,  I. travancorensis Fraser, 1931) and  I. carinatus Fraser, 1926 from Vietnam, while the other (1.00) contains three species from the eastern part of Oriental Realm ( I. montanus Karsch, 1891,  I. thailandicus Hämäläinen, 1985,  I. selysi Fraser, 1926).</p><p>The most basal branch of  Macromiidae is  Didymops, however, the opposed rest of the family is weakly supported (0.68), so it is better to speak of three polytomic clades:  Didymops,  Macromia and  Epophthalmia +  Phyllomacromia . The last cluster has the highest possible support (1.00), and inside it,  Epophthalmia and  Phyllomacromia themselves are also monophyletic with the same support. The monophyly of  Macromia is poorly supported (0.68).</p><p>Disregarding the weak node of 0.75, the genus  Macromia comprises five well supported main clusters, enumerated below from bottom to top.</p><p>- The most basal cluster of this genus (1.00) contains  M.aculeata and  Macromia katae Wilson,1993 corresponding to the  arachnomima -group by Wilson (1993) and Muraki (2010).</p><p>- One (0.89) of the three other polytomic clusters includes  M. pyramidalis and  Macromia westwoodi Selys, 1874, belonging to the  westwoodi -group by Lieftinck (Laidlaw 1922; Asahina 1987a).</p><p>- The next cluster (0.94) consists of two subclusters. One of them (1.00) contains nine species (from  M. flavocolorata to  M. daimoji), eight from the  calliope -group (Lieftinck 1929; Asahina 1987a; Muraki 2014) and  M. daimoji as a sister branch to them, thus being in line with isolation of this species into the  daimoji -group of its own by Muraki (2009).</p><p>The other subcluster (0.97) contains four species attributed to different groups:  M. cupricincta Fraser, 1924 (from the  cincta -group (Lieftinck 1929)),  M. irata (from the  cingulata -group (Fraser 1924)),  M. viridescens Tillyard, 1911 and  M. cydippe Laidlaw, 1922 (from  westwoodi -group in the broad sense by Lieftinck (1929; see also 1971)).</p><p>- The next cluster (0.91) again consists of two well supported subclusters. One of them (1.00) quite surprisingly includes  M. manchurica,  M. hamata and  M. annaimallaiensis Fraser, 1931 .  Macromia manchurica was not classified to any group, while  M. annaimallaiensis was attributed to the  cincta -group (Sadasivan et al. 2023). The other subcluster (0.89) includes  M. moorei,  M. splendens (Pictet, 1843),  M. malleifera Lieftinck, 1955 and  M. unca Wilson, 2004 .  Macromia splendens and  M. unca were not classified with respect to the groups of  Macromia .  Macromia moorei belongs to the  moorei -group in the sense of Lieftinck (1929). However, later Lieftinck (1955) implicitly subtracted from it his  amphigena -group, which included  M. malleifera and  M. amphigena, of which the former is found in this subcluster while the latter is found in the next cluster.</p><p>- The last cluster (1.00) includes two Palaearctic species  M. amphigena and  M. clio Ris, 1916 of the  amphigena group and all five Nearctic species involved, without Palaearctic-Nearctic divergence.</p><p>At the low level of the phylogenetic reconstruction of Fig. 2, some interesting points should be mentioned. The quantitative data on sequence differences are reported below as the number of nucleotide substitutions with the uncorrected p-distances (percentage of variable nucleotide positions) calculated on this base, and the number of indels, if any, but are not shown as alignments for the sake of space.</p><p>The sequences of the H3–H4 region of  M. flavocolorata from Cambodia and Western Ghats of India (Kerala) do not cluster together, differing in 30 substitutions (uncorrected p-distance 3.5%) and a two-nucleotide-long indel.</p><p>The sequences of  M. callisto from Cambodia and Thailand also do not cluster together and differ in 45 substitutions (5.2%) and four indels (two of one nucleotide and two of two nucleotides). At the same time, the Cambodian sequence of  M. callisto differs from that of  M. urania Ris, 1916 only in four substitutions (0.5%) while the Thai sequence of  M. callisto differs from that of  M. calliope Ris, 1916 with seven substitutions (0.8%) and a two nucleotide long indel. It is noteworthy, however, that the difference between  M. calliope and  M. flavocolorata from Cambodia is even smaller, consisting of five substitutions (0.6%) and one nucleotide indel. The situation with these three species is complicated and addressed in Discussion.</p><p>The sequences of the histone H3–H4 region of  M. manchurica and  M. hamata differs in four nucleotides only (0.5%). Such small differences would more fit specimens of the same species.</p><p>The two subspecies of  Macromidia genialis Laidlaw, 1923 (both specimens from Cambodia) appeared very close; they differ in 13 nucleotide substitutions (1.5%) and one nucleotide long indel.</p><p>In general, the tree based on the H3–H4 region (Fig. 2) corresponds rather well to the current taxonomical concept. However, in the genus  Macromia it supports monophyly of the  calliope -group and  arachnomima -group while representatives of other involved groups were found in different clusters and subclusters. The association of  M. annaimallaiensis from the Western Ghats and  M. manchurica from East Asia was not expected.</p><p>Of the two alternative treatments of  Macromidia,  Idionyx and  Oxygastra, as genera incertae sedis (Dijkstra et al. 2013; Paulson et al. 2025) or members of the family  Synthemistidae in the broad sense (Carle et al. 2015), our tree (Fig. 2) supports the former, since these genera do not cluster together.</p><p>ITS region tree</p><p>The Bayesian tree based on sequences of the other nuclear marker, the ITS region, is presented in Fig. 3. In general, it is rather similar to the H3–H4 region tree (Fig. 2), but involves more sequences, including those from GenBank.</p><p>Again, the family  Macromiidae is monophyletic with the highest support (1.00) while  Macromidia,  Idionyx (both again supported at 1.00) and  Oxygastra do not form a monophyletic clade.  Oxygastra is recovered as a branch of its own. Curiously,  Idionyx,  Oxygastra,  Cordulia and  Macromiidae appeared to comprise a monophyletic clade with the highest support of 1.00 (with  Idionyx being a sister clade to the rest), while  Macromidia is the most basal clade of the tree. The inner structure of  Idionyx is reshuffled as compared to the H3–H4 region tree (Fig. 2).</p><p>The ITS tree (Fig. 3) corresponds less to the current taxonomy than the H3–H4 region tree (Fig. 2). First,  Didymops is no longer the most basal clade of  Macromiidae but is found deep inside  Macromia, clustering with  M. moorei,  M. splendens and  M. malleifera . The most basal clade of  Macromiidae in the ITS tree (0.97) is now formed by  M. arachnomima,  M. aculeata,  M. katae and  M. westwoodi . This clade is opposed to the rest of the family, which is poorly supported (0.76). Hence the genus  Macromia has lost its monophyly.</p><p>Epophthalmia +  Phyllomacromia (1.00) form a sister clade to the large clade (0.97) including the rest of  Macromia, without the three above mentioned species but with  Didymops . Monophyly of  Epophthalmia and  Phyllomacromia has the maximum support.</p><p>In the remaining  Macromia spp., ten well supported clusters (0.96–1.00) and solitary lineages are resolved:</p><p>-  Macromia pyramidalis .</p><p>-  Macromia unca (one sequence).</p><p>-  Macromia viridescens (one sequence).</p><p>-  Macromia cupricincta,  Macromia cincta Rambur, 1842 and  M. irata .</p><p>-  Macromia cydippe (one sequence).</p><p>- The  calliope -group:  M. calliope,  M. callisto,  M. flavocolorata,  Macromia septima Martin, 1904,  M. urania,  Macromia gerstaeckeri Krüger, 1899 (but see ‘Discussion’ concerning this sequence),  Macromia chaiyaphumensis Hämäläinen, 1986,  Macromia sp3 .; again together with  M. daimoji forming a separate subcluster.</p><p>-  Macromia moorei,  M. splendens,  M. malleifera and  D. transversa .</p><p>-  Macromia manchurica and  M. annalmallaiensis .</p><p>- Five American species of  Macromia .</p><p>-  Macromia amphigena,  M. kubokaiya Asahina, 1964,  M. clio .</p><p>We see these clusters to be mostly the same as in the H3–H4 tree (Fig. 2), with the following differences:</p><p>- The  amphigena -group is now only very weakly coupled (0.65) with the American branch.</p><p>-  Macromia unca gets decoupled from the  moorei -group, which is, however, updated with  D. transversa .</p><p>-  Macromia westwoodi is decoupled from  M. pyramidalis and moved to the common clade with  M. arachnomima,  M. aculeata and  M. katae .</p><p>-  Macromia cupricincta and  M. cydippe are decoupled from  M. cingulata and  M. irata and are separate lineages.</p><p>In general, the clusters inside  Macromia remained mostly the same as in Fig. 2, while the basal topology of  Macromiidae changed substantially.</p><p>At the low level, the following notes are necessary.</p><p>The sequence of  M. flavocolorata from the Western Ghats (Kerala) again does not cluster with those from Vietnam and Cambodia at all, differing from them in a 36 nucleotide long insertion and 24 nucleotide substitutions (4.2%).</p><p>FIGURE 3. (Continued).</p><p>FIGURE 3. (Continued).</p><p>The sequences identified as  M. callisto are now found in three different places of the  calliope -group cluster. That from Cambodia is again found among  M. urania . Its sequence is identical to those of  M. urania from Japan and Vietnam, except for a small indel region where it has two nucleotides while the two latter have six and none, respectively.  M. callisto from Thailand again clusters with  M. flavocolorata and  M. calliope and has only one nucleotide different (0.1%) (plus one heterozygous position) from the sequence of  M. calliope from Vietnam. The GenBank sequence  M. callisto from Malaysia lies separately from other sequences identified as that species.</p><p>In the same  calliope -group cluster, there are two more pairs of species raising suspicions of their identity or misidentifications.</p><p>The sequences of  M. calliope differ from those of  M. flavocolorata from Vietnam and Cambodia in as few as three substitutions (0.5%) (plus one heterozygous position), the latter two differing from each other in two indels, one and two nucleotides long.</p><p>The sequence of  M. gerstaeckeri from GenBank and the two sequences of  M. chaiyaphumensis are identical.</p><p>The sequences of  M. daimoji of different origin tightly cluster together without geographic regularity. Their alignment (not shown) exhibits some substitutions and six sites of indels from one- to six-nucleotide-long: five of them are in the sequence from Taiwan and one in that from Russia; one of the Japanese sequences also differs from the two others in a two-nucleotide-long indel in one of the same sites.</p><p>The four sequences of  M. cupricincta are identical, while the two sequences of  M. berlandi Lieftinck, 1941 differ from them only in one substitution (0.2%) and a one-nucleotide-long deletion.</p><p>In all three cases where subspecies of the same species were involved, their sequences appeared to differ rather significantly, as follows.</p><p>Macromia amphigena amphigena,  M. amphigena fraenata,  M. clio and  M. kubokaiya are very close in the phylogenetic reconstructions of Fig. 3 but nevertheless formed four different and very well supported (0.94-1.00) clusters, of which the two presumed subspecies of  M. amphigena are most distant. Their sequences appeared to differ more in indels than in substitutions. Only 19 nucleotide positions are variable, but seven regions were affected by indels (two of them one-nucleotide-long). There are six (0.9%) diagnostic (found only in this taxon) substitutions and a diagnostic eight-nucleotide-long insertion of  M. kubokaiya, four (0.6%) diagnostic substitutions and a diagnostic eight-nucleotide-long insertion of  M. amphigena amphigena, one (0.2%) diagnostic substitution and two diagnostic two-nucleotide-long deletions of  M. amphigena fraenata, and just one (0.2%) diagnostic substitution of  M. clio, the latter species being variable for longer indels. The three sequences of  M. amphigena fraenata contain six (0.9%) variable positions and two indels (one- and two-nucleotide long) without geographic regularity.</p><p>Macromia moorei moorei from Himachal Pradesh, India, is found inside the tight cluster formed by  M. moorei malayana . The sequence of the former has a four-nucleotide-long insertion and four nucleotide substitutions (0.6%) not found in those of the latter, which in turn differ from each other in few substitutions.</p><p>The difference between the two subspecies of  Macromidia genialis appeared substantial. The sequence of  M. genialis buusraaensis Kosterin, 2018 differs from that of  M. genialis shanensis Fraser, 1927 in two eight-nucleotide-long insertions, a one-nucleotide-long insertion, three one-nucleotide-long deletions, and 12 nucleotide substitutions (1.5%).</p><p>All the five sequences of  E. vittata regardless of their origin (India, Cambodia, Laos and Vietnam) are identical.</p><p>COI tree</p><p>The phylogenetic tree reconstructed on the base of the mitochondrial COI barcoding fragment is shown in Fig. 4.</p><p>In addition to  Macromidia,  Idionyx and  Oxygastra, the genera of the ‘GSI clade’ in the sense by Ware et al. (2007) are now updated with  Synthemis . They again do not form a monophyletic clade. As in the ITS-tree (Fig. 3) and unlike the H3–H4-tree (Fig. 2),  Macromidia occupies the most basal position while  Idionyx clusters with  Macromiidae . However,  Macromidia donaldi (Fraser, 1924) appeared outside the main  Macromidia clade and comprises an independent branch, like  Cordulia aenea and  Synthemis eustalacta (Fig. 4).  Oxygastra now shows a tight clustering with  Idionyx (Fig. 4), although it clustered with  Macromidia in the H3–H4 tree (Fig. 2) and was independent in the ITS-tree (Fig. 3).</p><p>As in the two other trees,  Macromiidae are again monophyletic with the maximum support (1.00) in the COI tree (Fig. 4). Inside it, the most basal clade (1.00), opposed to the rest of the family, is represented by  M. arachnomima,  M. aculeata and  M. katae . The rest of the family (0.98) then diverges into four clades with moderate support:</p><p>-  Phyllomacromia (0.84).</p><p>-  Epophthalmia (0.98).</p><p>- The  calliope -group without  M. daimoji (0.85).</p><p>- The rest of  Macromia with  D. transversa (0.90).</p><p>The following smaller clusters and separate lineages were revealed with supports above 0.8:</p><p>-  Macromia daimoji (1.00).</p><p>-  Macromia westwoodi and  M. cydippe (0.95).</p><p>-  Macromia cupricincta,  M. irata,  M. cincta and  M. viridescens (0.86).</p><p>-  Macromia pyramidalis and  M. unca (0.85).</p><p>-  Macromia annaimallaiensis .</p><p>-  Macromia moorei,  M. malleifera,  M. splendens (0.99).</p><p>-  Macromia manchurica and  M. hamata (1.00) (just four variable positions, 0.6%, without any regularity of substitutions as to the three sequences).</p><p>-  Disymops transversa (1.00).</p><p>- The American species of  Macromia (0.93).</p><p>-  Macromia amphigena sspp and  M. clio (1.00).</p><p>We see that</p><p>- clustering of  Phyllomacromia with  Epophthalmia is no longer supported (0.53);</p><p>-  Didymops is found inside  Macromia, as in the ITS tree (Fig. 3);</p><p>-  Macromia daimoji is now decoupled from the  calliope -group;</p><p>FIGURE 4. (Continued).</p><p>FIGURE 4. (Continued).</p><p>-  Macromia annaimallaiensis is decoupled from  M. manchurica, which it clustered with in other trees (Figs 2–3).</p><p>In general, the COI tree (Fig. 4) is more or less similar to the H3–H4 (Fig. 2) and ITS (Fig. 3) trees as having similar clusters, but they are less supported and the position of some species differed strongly, especially of  M. donaldi (Fraser, 1924),  O. curtisii and  M. annaimallaiensis .</p><p>At the species level the following details are the same as in the ITS tree:</p><p>-  Macromia moorei moorei (India) is again found inside the branch of  M. moorei malayana . The haplotypic network constructed for their COI sequences (Fig. 5) did not reveal the two subspecies but shows that the sequence from India, Vietnam and the two Thai sequences taken together are equidistant.</p><p>-  Macromia hamata is inside the branch of  M. manchurica .</p><p>-  Maccromia amphigena fraenata clusters with  M. clio rather than with  M. amphigena amphigena . To better illustrate the latter case, we constructed the haplotypic network for the COI sequences of  M. amphigena sspp. and  M. clio (Fig. 6). The clusters of the three taxa included are separated by significant distances (21 and 44 mutation steps; 3.2 and 6.3%), so best corresponding to closely related but distinct species. The sequences of  M. amphigena fraenata from West Siberia and Primorye differ in six (0.9%) substitutions only.</p><p>The situation in the  calliope -group in the COI tree (Fig. 1) is as complicated as in the two previous trees (Figs 2– 3) (unfortunately, we did not manage to get the ITS sequence from the specimen of  M. calliope itself), as follows:</p><p>- The sequences of  M. daimoji from Korea, Russia and Taiwan (the latter woud be identified as  M. chui in recent past) are very similar: just four (0.6%) substitutions in total, three of which are in the sequence from Taiwan.</p><p>-  Macromia flavocolorata from the Western Ghats (Kerala) again does not cluster with those from Vietnam and Cambodia but lies separately.</p><p>- The sequences of  M. callisto are found in two different places, again near  M. flavocolorata from Vietnam and Cambodia and near  M. urania, to which  M. chaiyaphumensis is now added.</p><p>To illustrate the situation in the  calliope -group, we constructed its COI haplotypic network (Fig. 7). The sequence of  M. chaiyaphumensis differs from the closest ones of  M. urania and  M. callisto from Cambodia in four mutation steps only (0.6%).  Macromia sp1,  Macromia sp3 and  M. flavocolorata from Kerala are species most distant from the rest.</p><p>Two sequences of  Macromidia genialis shanensis from Thailand and Cambodia and the sequence of  M. genialis buusraaensis from Cambodia appeared equally distant from each other (Fig. 8), having in total 101 (as much as 14.5%) variable positions.</p><p>The sequences of  E. vittata contained 18 variable positions (2.5%) showing no geographical regularity, but ten substitutions were confined to the beginning of the sequence of a specimen from Cambodia.</p><p>Joint analysis</p><p>In order to summarise and ‘average’ the phylogenetic signals from the three markers involved, we undertook their joint analysis using StarBeast. This software (i) is able of utilising any number of sequences and markers available for each species and (ii) takes into account that sequences evolve not alone but as incorporated in some species, which actually evolve, and reconstructs the most plausible phylogenetic tree of species rather than sequences. To make this possible, it takes into account the species identifications of each particular input sequence and hence is influenced by expert opinion of identifiers. Because of this, we removed from this analysis all sequences identified as  M. callisto, since this identification appeared confusing in view of the phylogenetic reconstructions of particular markers (Figs 2–4). We also removed the only sequence of  M. gerstaeckeri (of ITS region), which was obviously misidentified  M. chaiyaphumensis (see ‘Discussion’). Also, we separated  M. flavocolorata from India, Kerala as a species different from  M. flavocolorata from Vietnam and Cambodia, as was obvious from the above analysis. The resulting tree is shown in Fig. 9.</p><p>In this tree, the family  Macromiidae is monophyletic with a high support (0.99), as in all other trees. Curiously, the genera  Idionyx,  Oxygastra,  Macromidia and  Cordulia cluster with it to form a major clade (0.96), but do not reliably cluster with each other, while  Synthemis is left outside, but this could be an artifact due to representation of the latter with just one COI sequence. The genus  Macromia becomes a monophyletic but rather not supported (0.65) major clade and only if  Didymops is included in it.</p><p>As could be expected from a tree based on three rather contradicting markers at once, most of its clades are weakly supported. However, the remaining highly supported clades should correspond to actually existing monophyletic entities and worth being enumerated. These are:</p><p>- The genus  Idionyx (0.98), divided into three inner branches, two of which the have maximum support (1.00):  I. saffronatus with  I. travancorensis and four species ( I. montanus,  Idionyx murcia Lieftinck, 1971,  I. selysi,  I. thailandicus) referring to the yolanda -group.</p><p>- The genera  Phyllomacromia and  Epophthalmia (both supported at 1.00), together also forming a very well supported (0.96) clade.</p><p>- The  calliope -group of  Macromia (0.99), including  M. daimoji as its most basal branch.</p><p>-  Macromia arachnomima,  M. aculeata and  M. katae (1.00).</p><p>-  Macromia cupricincta,  M. berlandi,  M. cincta and  M. irata (1.00).</p><p>-  Macromia splendens,  M. moorei,  M. malleifera together with  D. transversa (0.93)!</p><p>-A large clade with the support of 0.99 with three inner branches of the highest support of 1.00: (i)  M.manchurica,  M. hamata and  M. annaimallaiensis; (ii) the North American species, and (iii) the  amphigena -group.</p><p>We also reconstructed the StarBeast tree only for our two nuclear markers, the histone H3–H4 and ITS regions. It showed the same, locally a bit less supported results for  Macromiidae, so we do not show it. The pattern of the ‘GSI clade’ genera was somewhat different, that could be expected from the particular trees of the nuclear markers (Figs 2–3).</p></div>	https://treatment.plazi.org/id/327687E4FFEAFFADFF328F59FEBAE781	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kosterin, Oleg E.;Vierstraete, Andy;Schneider, Thomas;Kompier, Tom;Hu, Fang-Shuo;Everett, Larry;Makbun, Noppadon;Onishko, Vladimir V.;Papazian, Michel;Dumont, Henri J.	Kosterin, Oleg E., Vierstraete, Andy, Schneider, Thomas, Kompier, Tom, Hu, Fang-Shuo, Everett, Larry, Makbun, Noppadon, Onishko, Vladimir V., Papazian, Michel, Dumont, Henri J. (2025): Molecular phylogenetic analysis of the family Macromiidae (Odonata) worldwide based on a mitochondrial and two nuclear markers, with a short overview of its taxonomic history. Zootaxa 5620 (4): 501-545, DOI: 10.11646/zootaxa.5620.4.1, URL: https://doi.org/10.11646/zootaxa.5620.4.1
