Male Rhyothemis plutonia, from here.

Belongs within: Paneurypalpidomorpha.
Contains: Brachydiplax, Orthetrum, Libellula, Agrionoptera.

The Libellulidae, skimmers and perchers, are a diverse group of dragonflies with antenodal costal and subcostal veins in strict alignment, and a rounded base to the hindwing in both sexes (Theischinger & Hawking 2006). Members of the group vary widely in appearance; some have strongly patterned wings whereas others have entirely transparent wing membranes.

The family is divided between several subfamilies, primarily on the basis of wing venation characters. Members of the largest subfamily, the Sympetrinae, have fore wings with the last antenodal crossvein not extending beyond the subcosta, the arculus between the first and second antenodal crossveins, and distinct radial and medial planates, and hind wings with a broad base and a highly visible anal loop. The Libellulinae usually have the arculus at or distal to the second antenodal crossvein in both pairs of wings; it is always so at least in the hind wing. The Pantalinae are a subfamily of broad-winged, strong fliers (Watson & O’Farrell 1991); the pantropical Pantala flavescens may be the single most widespread odonatan species. The Old World tropical genus Rhyothemis contains broad-winged, short-bodied dragonflies with extensive dark markings on the fore and hind wings, and a typically fluttering flight (Theischinger & Hawking 2006).

Libellulidae: on the wing
Published 21 June 2018

Dragonflies of the order Odonata are unquestionably one of the more familiar groups of insects to the general public. They are large, visible and eye-catching, and may be quite colourful. Some have even taken to ‘twitching’ dragonflies in the same manner as bird species, identifying species observed on the wing and keeping a tally of how many they have seen.

And at the top of many people’s list: the wandering glider Pantala flavescens, copyright Jeevan Jose, the world’s most widespread dragonfly species.

Ecologically, in contrast, dragonflies may be called a relatively conservative group. All begin their lives as aquatic predators before emerging with adulthood as fast-moving aerial predators. All are generalists, feeding on whatever other insects may be unfortunate enough to fall into their grasp. All dragonflies conform to a fairly similar overall bauplan when compared to the diversity of forms that may be found in many other insect orders (for instance, there are no flightless dragonflies). Classification of dragonflies has often focused heavily on features of the wing venation, tracing its lines in their criss-crossing network.

Hind wing of a libellulid with the anal loop highlighted, from here.

The largest of the generally recognised families of dragonflies is the Libellulidae, containing over 1000 of the approximately 6000 known species of Odonata (Pilgrim & von Dohlen 2008). Characteristic features of the Libellulidae include the presence of the ‘anal loop’, an arrangement of veins in the hind wing forming what has been described as a boot shape. In the case of the genus Libellula, at least, the shape of the anal loop rather reminds me of one of the legs on the Manx flag. Members of the Libellulidae are commonly known as perchers or skimmers in reference to their hunting behaviours; others have similarly composed names such as darters or pondhawks. A number of members of the family have strikingly banded or coloured wings, leading to vernacular labels such as amberwings or pennants. Members of the genus Tramea are commonly known as saddlebags in reference to the dark patches at the base of their hind wings.

Common picturewing Rhyothemis variegata, copyright Tarique Sani.

Members of the Libellulidae have been divided between about a dozen subfamilies, again primarily defined on the basis of wing venation. However, distinctions between the subfamilies have always been vague with many subfamilies recognised by particular combinations of characters rather than characters unique to each subfamily alone. This vagueness has been underlined by recent molecular studies which have identified most subfamilies as polyphyletic. It seems likely that the defining features of these subfamilies are convergences related to similar ecologies. The ‘Sympetrinae’ include species with a preference for open watery habitats such as ponds and marshes where they spend a lot of time perched on exposed vegetation (Pilgrim & von Dohlen 2008). The ‘Tetrathemistinae’, with narrow wings with somewhat reduced venation, are found along forest streams (Fleck et al. 2008). The genera Tramea and Pantala, falsely united in the subfamily Trameinae by broadened bases on the hind wings, are specialised for long-distance flights spending extended periods on the wing (Pilgrim & von Dohlen 2008). Indeed, the wandering glider Pantala flavescens is the world’s most widespread dragonfly species, being found in warmer regions of the entire globe and seemingly capable of migrations between separate continents.

The slightly freakish-looking larva of Orionothemis felixorioni, from Fleck et al. (2009).

So if we’re going to have a stable classification for libellulids, we need to look past their wings. Intriguingly, larval features may prove more useful in this regard than adult characters. Fleck et al. (2008) examined a group of genera previously classified in the Tetrathemistinae but whose larvae were more similar to those found among members of the Libellulinae. A molecular phylogeny showed that, whereas the Tetrathemistinae as a whole were polyphyletic, these genera were indeed associated with the Libellulinae as their larvae indicated. With further research, we find that libellulid classification need not be all in vein.

Systematics of Libellulidae

Characters (from Theischinger & Hawking 2006): Wings with antenodal costal and subcostal veins in strict alignment, primary antenodals not discernible; sectors of arculus generally stalked at base; anal loop generally well developed; hindwing base rounded in both sexes. Lavae with labium ladle-shaped, bearing premental and palpal setae; labial palps smooth to strongly toothed; abdomen generally laterally and (more rarely) dorsally armed.

    |    |--Austrothemis nigrescens (Martin 1901)WO91, TH06
    |    `--TrithemisSL89
    |         |--T. adelpha Selys 1878SL89
    |         |--T. festiva [=Libellula festiva; incl. Dythemis infernalis]SL89
    |         |--T. fraternaSL89
    |         `--T. tirbyiIm92
    |    |--Tholymis tillarga (Fabricius 1798)WO91, TH06
    |    `--ZyxommaTH06
    |         |--*Z. petiolatum Rambur 1842SL89, TH06
    |         |--Z. atlanticumSL89
    |         |--Z. elgneri Ris 1913TH06
    |         |--Z. multinervorum Carpenter 1897TH06
    |         `--Z. obtusumSL89
    |    |--BrachydiplaxTH06
    |    |--Raphismia bispina (Hagen 1867)WO91, TH06
    |    `--Nannophya Rambur 1842WO91, T08a
    |         |--N. australis Brauer 1865TH06
    |         |--N. dalei (Tillyard 1908)TH06
    |         |--N. occidentalis (Tillyard 1908)TH06 [=N. dalei occidentalisC70]
    |         |--N. paulsoni Theischinger 2003TH06
    |         `--N. pygmaea [incl. N. exigua]SL89
    |    |--TetrathemisTH06
    |    |    |--T. flavescensT08c
    |    |    |--T. irregularisTH06
    |    |    |    |--T. i. irregularisTH06
    |    |    |    `--T. i. cladophila Tillyard 1908TH06
    |    |    `--T. platypteraTH06
    |    `--NannophlebiaWO91
    |         |--N. eludens Tillyard 1908TH06
    |         |--N. imitansT08c
    |         |--N. injibandi Watson 1969TH06
    |         |--N. lorquiniT08c
    |         |--N. mudginberri Watson & Theischinger 1991TH06
    |         `--N. risi Tillyard 1913TH06
    |    |--Pantala flavescens (Fabricius 1798)TH06 [incl. Libellula terminalisSL89, L. viridulaSL89]
    |    |--Hydrobasileus brevistylus (Brauer 1865)WO91, TH06
    |    |--TrapezostigmaWO91
    |    |    |--T. loewi Brauer 1866T13
    |    |    `--T. stenolobaPH10
    |    |--CamaciniaWO91
    |    |    |--C. giganteaTH06
    |    |    `--C. othello Tillyard 1908TH06
    |    `--TrameaWO91
    |         |--T. euryaleSL89
    |         |--T. eurybia Selys 1878TH06
    |         |--T. loewii Kaup 1866TH06
    |         |--T. propinqua Lieftinck 1942TH06
    |         `--T. stenoloba (Watson 1962)TH06
    |--RhyothemisTH06 [RhyothemistinaeWO91]
    |    |--R. alcestisT08b
    |    |--R. braganza Karsch 1890TH06
    |    |--R. chloeT08b
    |    |--R. curiosa Selys Longchamps 1889 (see below for synonymy)SL89
    |    |--R. fulgens Selys Longchamps 1889SL89
    |    |--R. fuliginosaSL89
    |    |--R. graphiptera (Rambur 1842)TH06
    |    |--R. phyllis (Sulzer 1776)TH06
    |    |    |--R. p. phyllisTH06
    |    |    |--R. p. beatricisTH06
    |    |    `--R. p. chloeTH06
    |    |--R. plutoniaSL89
    |    |--R. princeps Kirby 1894TH06
    |    |--R. resplendens Selys 1878TH06
    |    `--R. viduaSL89
    |    |--OrthetrumWO91
    |    |--Notolibellula bicolor Theischinger & Watson 1977WO91, TH06
    |    |--LibellulaRS10
    |    |--AgrionopteraWO91
    |    |--Potamarcha congener (Rambur 1842)O70, TH06
    |    |--Nannodythemis Brauer 1868T08a
    |    |    |--*N. australis Brauer 1868T08a
    |    |    |--N. dalei Tillyard 1908T08a
    |    |    `--N. occidentalis Tillyard 1908T08a
    |    |--LathrecistaWO91
    |    |    |--L. asiaticaTH06
    |    |    |    |--L. a. asiaticaTH06
    |    |    |    `--L. a. festa (Selys 1879)TH06 [=Agrionoptera festaSL89, Orthemis festaSL89]
    |    |    |--L. difficilis [=Agrionoptera difficilis, Orthemis difficilis]SL89
    |    |    |--L. pectoralis [=Agrionoptera pectoralis, Orthemis pectoralis]SL89
    |    |    `--L. simulans [=Agrionoptera simulans, Orthemis simulans]SL89
    |    |--Zygonidia Kirby 1900K00a
    |    |    `--*Z. insignis Kirby 1900K00a
    |    `--AcisomaK00a
    |         |--A. ascalaphoidesSL89
    |         `--A. panorpoides (Ramb. 1842) [=Libellula panorpoides]K00a
         |--Nannodiplax rubra Brauer 1868WO91, TH06
         |--Rhodothemis lieftincki Fraser 1954WO91, TH06
         |    |--C. nigrifrons (Kirby 1894)TH06
         |    |--C. servilia (Drury 1773) [=Libellula servilia]K00a
         |    `--C. soror [=Trithemis soror]SL89
         |    |--S. corruptum [=Tarnetrum corruptum]A72
         |    |--S. flaveolumK01
         |    |--S. internumA07
         |    |--S. obtrusumA07
         |    |--S. pedemontanumK01
         |    |--S. rubicundulumA07
         |    |--S. sanguineumK01
         |    |--S. scoticumK01
         |    `--S. striolatumRD77
         |--Diplacodes Kirby 1889WO91, FB03
         |    |--*D. tetraK00a [=Libellula tetraSL89, Diplacina tetraSL89]
         |    |--D. bipunctata (Brauer 1865)TH06
         |    |--D. concinna [=Libellula concinna, Diplacina concinna]SL89
         |    |--D. flavistyla [=Libellula flavistyla, Diplacina flavistyla]SL89
         |    |--D. haematodes (Burmeister 1839)TH06
         |    |--D. melanopsis (Martin 1901)TH06
         |    |--D. nebulosa (Fabricius 1793)TH06 [=Libellula nebulosaSL89, Diplacina nebulosaSL89]
         |    `--D. trivialis (Rambur 1842)TH06 [=Libellula trivialisK00a, Trithemis trivialisK00a]
         `--NeurothemisWO91 [incl. PolynevraSL89]
              |--N. basalisSL89
              |--N. degener Selys Longchamps 1879 [incl. Libellula intermedia]SL89
              |--N. feralis [incl. Libellula equestris]SL89
              |--N. fluctuans Selys 1879SL89
              |    |--N. f. fluctuansSL89
              |    `--N. f. disparilis Kirby 1889SL89
              |--N. oligoneura Brauer 1867TH06
              |--N. stigmatizans (Fabricius 1775)TH06
              `--N. tullia (Drury 1773) [=Libellula tullia]K00a
Libellulidae incertae sedis:
  Nannothemis Brauer 1868FB03
  Palaeothemis tillyardi Fraser 1923FB03
  Neodythemis Karsch 1889FB03
  Diplax trivialisSL89
  Tyriobapta torridaSL89
  Lepthemis vesiculosaSL89
  Huonia melvillensis Brown & Theischinger 1998TH06
  Celithemis eponinaA07
  Erythemis rufaA07, T08c
  Orchithemis [incl. Calothemis]SL89
    |--O. pruinans (Selys 1878) [=Calothemis pruinans]SL89
    `--O. pulcherrima Brauer 1878 [incl. Calothemis exsudans Selys 1878]SL89
  Nesciothemis Longfield 1955L55
    |--*N. farinosum (Förster 1898) (see below for synonymy)L55
    `--N. fitzgeraldi (Pinhey 1955) [=Orthetrum fitzgeraldi]L55
    |--O. congenerSL89
    `--O. lineataSL89
    |--L. cleliaSL89
    |--L. glauca Brauer 1865SL89
    |--L. luzonica Brauer 1868SL89
    `--L. pruinosa [=Libellula pruinosa]SL89
    |--*D. nanaSL89
    |--D. braueriSL89
    |--D. militarisSL89
    |--D. nigrilabrisSL89
    |--D. obscuraSL89
    `--D. smaragdinaSL89
    |--L. frigidaMG06
    |--L. hudsonicaB-PE07
    |--L. intactaMG06
    |--L. intermediaIw92
    |    |--L. i. intermediaIw92
    |    `--L. i. ijimai Asahina 1961Iw92
    |--L. pectoralisK01
    |--L. proximaMG06
    `--L. rubicundaK01
  Condalia woottoni Whalley & Jarzembowski 1985RJ93
  Zygonyx Hagen 1867K00a, K00b
    `--*Z. irisK00b
  Neurocena Kirby 1900K00b
    `--*N. ida [=Zygonyx ida; incl. Pseudomacromia luxuriosa Karsch 1893]K00b
  Schizonyx Karsch 1890K00b
    `--S. luctifera (De Selys 1869) [=Zygonyx luctifera]K00b
  Cratilla Kirby 1900K00b
    `--*C. metallica (Brauer 1878) [=Orthemis metallica, Protorthemis metallica]K00b

*Nesciothemis farinosum (Förster 1898) [=Orthetrum farinosum; incl. Hadrothemis hyalina Fraser 1927 (n. n.), O. pollinosum Karsch 1899]L55

Rhyothemis curiosa Selys Longchamps 1889 [incl. R. curiosa var. apicalis Selys Longchamps 1889, R. curiosa var. transversa Selys Longchamps 1889]SL89

*Type species of generic name indicated


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