Corresponding author: Davide Badano ( firstname.lastname@example.org )
Academic editor: Susanne Randolf
© 2017 Davide Badano, Horst Aspöck, Ulrike Aspöck.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Badano D, Aspöck H, Aspöck U (2017) Taxonomy and phylogeny of the genera Gymnocnemia Schneider, 1845, and Megistopus Rambur, 1842, with remarks on the systematization of the tribe Nemoleontini (Neuroptera, Myrmeleontidae). Deutsche Entomologische Zeitschrift 64(1): 43-60. https://doi.org/10.3897/dez.64.11704
The delineation of antlion genera has often been based on morphological characters not tested in a phylogenetic context, thus seriously impairing the study of systematics of the family Myrmeleontidae. Nebulous generic limits also impede the taxonomy and study of the affinities of closely related species. As a case study, the generic placement of Megistopus mirabilis Hölzel, 1980, was based on a single leg character. To test the position of this species, the reciprocal relationships of the members of the genera Gymnocnemia Schneider, 1845, and Megistopus Rambur, 1842 were investigated, using a morphology-based phylogenetic analysis. This approach demonstrated that M. mirabilis should actually be assigned to the genus Gymnocnemia, as G. mirabilis comb. n. This analysis also supports the subdivision of the tribe Nemoleontini in two subclades based on morphology of male and female genitalia. A new characterisation of these genera is provided, as well as a redescription of the very rare G. mirabilis and the poorly investigated Megistopus lucasi (Navás, 1912). An updated identification key to the members of the genera Gymnocnemia and Megistopus is presented.
Systematics, Myrmeleontinae, Nemoleontini, antlion, Western Palaearctic
Myrmeleontidae, commonly known as antlions, are the most species-rich family of the order Neuroptera, including 1657 described species (
Examined specimens are deposited in the following collections: collection of Davide Badano, Taggia, Italy (DB); collection of Horst and Ulrike Aspöck, Vienna, Austria (HUAC); collection of Herbert Hölzel, Natural History Museum Vienna, Austria (NHMW) (Suppl. material
Specimens were examined and measured with a Leica® MZ 9.5 stereomicroscope equipped with an optical micrometre and photographed with a Canon® EOS 600D digital camera with Canon® lens MP-E 65 mm; obtained images were stacked with the software Zerene® STACKER. Specimens were measured following the protocol of
The distribution map was made with Natural Earth, free vector and raster map data, available at www.naturalearthdata.com.
Abbreviations used in the text: Biogeogr, biogeography; Biol, biology; Cat, catalogue; Com, comment; Distr, distribution; Key, identification key; List, checklist; ODescr, original description; Syn, synonymy.
Characters and states were coded for a set of antlion taxa illustrative of morphological variability across the family. Palpares libelluloides Linnaeus (Palparinae) was selected as the outgroup. A data matrix including 56 characters and 121 states was prepared in MESQUITE version 3.03 (
1. Labial palpus: (0) short and stout; (1) extremely long and thin. The labial palpus is extremely elongated, much longer than the head width in Palpares Rambur.
2. Labial palpus, sensory area (palpimacula): (0) rounded; (1) slit-like. The sensory area of the apical palpomere is slit-like in Palpares libelluloides (see also
3. Pronotum: (0) wider than long; (1) as long as wide, (2) noticeably longer than wide. The proportions of pronotum widely vary across the family. In Palpares it is much wider than long, while in most antlions it is subquadrate and as long as wide. In Dendroleontini (Dendroleon Brauer, Tricholeon Esben-Petersen) and in G. mirabilis the pronotum is characteristically much longer than wide (see also
4. Thorax, outstanding setae: (0) absent; (1) present. In members of the genus Gymnocnemia, the thorax is covered with large and robust setae (Fig.
5. Legs: (0) short and stout; (1) long and slender. Most myrmeleontids have relatively short and robust legs. Several species of Dendroleontini, such as Dendroleon and Tricholeon, and Nemoleontini, such as Gymnocnemia, Megistopus and Nedroledon Navás, have very long and slender legs, with the femur at least 7 times longer than wide (Fig.
6. Legs, colour pattern: (0) similar in all legs; (1) differing in each leg pair. In members of the genus Gymnocnemia, pro-, meso- and metathoracic legs have a strikingly different colour pattern.
7. Tarsus, segments 1-4: (0) T1-T4 short, of the same size; (1) T1 much longer than the others, T2-T4 short; (2) T1-T3 elongated, of the same size. In Palpares, tarsomeres 1-4 are of similar length and equally short. In most genera of myrmeleontids, the basal tarsomere (T1) is much longer than tarsomeres 2-4, which are short; the basal tarsomere is usually as long as tarsomeres 2 and 3 together (T2+T3). In Megistopus, tarsomeres 1-3 are subequal in size, equally elongated while T4 is short (Fig.
8. Tibial spurs: (0) absent; (1) present. Tibial spurs are absent in G. variegata and G. editaerevayae.
9. Tibial spurs (if present), length: (0) long, at least 1/3 of tarsus length; (1) short, 1/4-1/5 of tarsus length; (2) minute, 1/6 of tarsus length. Tibial spurs are relatively long (at least one third of tarsus length) in most analysed taxa; however, they are relatively short, less than one fourth of tarsus length in Cueta lineosa (Rambur) and Megistopus (Fig.
10. Tarsal claws: (0) not opposable; (1) opposable. In some antlions, the tarsal claws can be folded against a brush of thickened setae on the distal tarsomere. Of the analysed species, this condition is found in members of the genera Megistopus, Gymnocnemia and Nedroledon (Fig.
11. Prothoracic leg: (0) similar in length to meso- and metathoracic legs; (1) longer than meso- and metathoracic legs. In Gymnocnemia the prothoracic leg is much longer and thinner than the following pairs (Fig.
12. Prothoracic leg femur, hair-like sensillum (femoral sense hair sensu
13. Forewing, origin of Rs: (0) 1/4 of wing length or less; (1) 1/3 of wing length; (2) 1/2 of wing length. Rs arises near wing base (1/4 of wing length) in Dendroleontini (Dendroleon and Tricholeon), while it originates distally (around 1/3 of wing length) in all the other analysed taxa. In G. mirabilis, Rs arises at half wing length (Fig.
14. Forewing, vein CuP: (0) long vein, running independently from 1A for its entire length; (1) short vein, parallel to 1A for a short distance and then merging with it, (2) crossvein-like. In Palpares and other Palparini, CuP is not fused with 1A, rather it is a distinct vein. In most other myrmeleontids, CuP is a very short vein fusing with 1A (Fig.
15. Forewing, vein CuP+1A: (0) convergent with wing margin; (1) parallel to wing margin. The vein CuP+1A runs toward wing margin in almost all the analysed taxa, while it is parallel to wing margin in Dendroleon, Tricholeon and Creoleon Tillyard.
16. Forewing, vein 2A: (0) gently curved; (1) close to 1A, then strongly bent toward to 3A. In Palpares, Dendroleon, Tricholeon and Solter Navás, vein 2A runs as a smooth curve. In Cueta Navás, Myrmeleon Linnaeus and in the members of Nemoleontini 2A is characteristically angled (Fig.
17. Hind wing, presectoral area: (0) 1 crossvein; (1) more than 4 crossveins. The hind wing presectoral area is crossed by one crossvein in Dendroleontini (Dendroleon, Tricholeon) and Nemoleontini (Nemoleon Navás, Distoleon Banks, Neuroleon, Macronemurus, Creoleon) (Fig.
18. Pilula axillaris: (0) absent; (1) present. Among the analysed taxa, the pilula axillaris is absent in Cueta and Nemoleontini (see Discussion).
19. Male, abdomen length: (0) as long as the wings; (1) noticeably longer than the wings. The males of several antlion genera, such as Cueta, Distoleon, Macronemurus, Creoleon and Nedroledon, are characterized by elongated abdomens, much longer than wings.
20. Male gonocoxites 9 and 11: (0) not fused; (1) fused into a tube-like structure. In Palpares and Cueta gonocoxites 9 and 11 are amalgamated into a tube-like structure.
21. Male, gonocoxites 9, relative position to gonocoxites 11 (if not fused): (0) ventro-caudal to gonocoxites 11; (1) partly encircling gonocoxites 11. In G. variegata and G. editaerevayae, the proximal part of gonocoxites 9 characteristically curves upward, partly surrounding gonocoxites 11 in lateral view (Fig.
22. Male, gonocoxites 9, shape: (0) plate-like; (1) fused in a Y-shaped structure; (2) fused in an upward hook-shaped structure. In ventro-caudal view, the apex of gonocoxites 9 appears as paired sclerotized plate-like structures in most myrmeleontid taxa. In several genera of Nemoleontini, including Nemoleon, Distoleon, Neuroleon, Macronemurus, Creoleon, gonocoxites 9 are fused into a structure resembling a “Y”, with a ventral projection and two processes curved upward (Fig.
23. Male, gonocoxites 9 (if plate-like), relative position: (0) parallel; (1) oblique, converging apically. In ventro-caudal view, the plate-like apexes of gonocoxites 9 are parallel to each other in Palpares, Solter and Gymnocnemia (Fig.
24. Male, ectoproct, ventrocaudal projections: (0) absent; (1) present. In the males of several antlion genera, including Palpares, Cueta, Macronemurus, Nedroledon, and Nemoleon, the ectoproct is equipped with a posterior “clasper-like” processus (Fig.
25. Male ectoproct, length of ventrocaudal projection (if present): (0) short, as long or slightly longer than ectoproct; (1) extremely long, much longer than ectoproct. The posterior projections of ectoproct are short in Cueta, while they are extremely long in Palpares, Macronemurus and Nemoleon (Fig.
26. Female, process on segment 8: (0) absent; (1) present. In the female of several myrmeleontid genera, on the segment 8, proximal to gonocoxites 8 or at their base a pair of setiferous processes is present, usually termed “anterior gonapophyses” (e.g.
27. Female, gonocoxite 7: (0) absent; (1) present. The gonocoxite 7 (praegenitale of authors, see
28. Female, gonocoxites 8: (0) not prominent; (1) short, 3-4 times longer than wide; (2) long, digitiform, over 5 times longer than wide. The gonocoxites 8 (posterior gonapophyses sensu
29. Female, processes on segment 9: (0) absent; (1) present. Setiferous processes at base of gonocoxites 9 are present in the analysed species of Cueta, Dendroleon and Tricholeon.
30. Female, gonocoxites 9, chaetotaxy: (0) only thin setae; (1) stout setae present. Gonocoxites 9 are only covered with thin setae in Dendroleon and Tricholeon, while the remaining analysed genera are characterised by the presence of stout digging setae. (Figs
31. Female, ectoproct, chaetotaxy: (0) only thin setae; (1) stout setae present. The female ectoprocts are only covered with thin setae in Cueta, Solter, Dendroleon and Tricholeon, while all other analysed genera are provided with stout digging setae.
32. Larva, chaetotaxy of the dorsal side of the head capsule: (0) sensilla mainly bristle-like; (1) sensilla mainly dolichasters; (2) sensilla scale-like. See
33. Larva, ocular tubercle, size: (0) large; (1) small. See
34. Larva, upward bending of mandible: (0) not bent, i.e. straight; (1) bent upward. See
35. Larva, fringe of extremely long setae on lateral side of the mandible: (0) absent; (1) present. See
36. Larva, mesothoracic spiracle: (0) not raised; (1) raised. See
37. Larva, metanotum tuft of setae: (0) absent; (1) present. See
38. Larva, anterior mesothoracic setiferous process, type: (0) scolus-like; (1) small protuberance. See
39. Larva, posterior mesothoracic setiferous process: (0) scolus-like; (1) tubercle-like. See
40. Larva, anterior metathoracic setiferous process, type: (0) scolus-like; (1) tubercle-like; (2) small protuberance. See
41. Larva, fringe of elongated setae on meso- and metathoracic legs: (0) absent; (1) present. See
42. Larva, abdominal segment 9, shape: (0) longer than wide; (1) wider than long. See
43. Larva, abdominal sternite 9, specialized digging setae on ventral surface: (0) absent; (1) present. See
44. Larva, abdominal sternite 9, specialized digging setae (if present), number and pattern: (0) one line of 2-4 narrow digging setae at the base of rastra; (1) six stout digging setae arranged into a X-shaped pattern; (2) one or two transversal rows of stout digging setae. See
45. Larva, rastra: (0) absent; (1) present. See
46. Larva, rastra (if present), size: (0) very small, not prominent; (1) prominent and sclerotized.
47. Larva, rastra (if present), digging setae: (0) fused; (1) unfused. See
48. Larva, rastra (if present), relative dimensions of digging setae: (0) subequal in size; (1) medial pair distinctly smaller than lateral ones. See
The cladistic analysis yielded two most parsimonious trees with a tree length of 83 steps, a consistency index (CI) of 0.687 and a retention index (RI) of 0.752. One of the two most parsimonious trees was selected to map inferred character changes and is shown in Fig.
Clade G was supported by one nonhomoplasious apomorphy (10:1, tarsal claws opposable) and one homoplasious apomorphy (5:1) with a Bremer support value of 1. Within the latter group, the genus Megistopus broke up as polyphyletic with “Megistopus” mirabilis forming a clade (clade H) with the members of the genus Gymnocnemia. The monophyly of this genus (“M.” mirabilis + (G. variegata + G. editaerevayae)), relied on three nonhomoplasious apomorphies (4:1, thorax with outstanding setae; 6:1, leg colour pattern differing in each leg pair; 11:1, prothoracic leg longer than meso- and metathoracic legs) and obtained a Bremer support value of 3. A sister relationship between G. variegata and G. editaerevayae was retrieved based on one nonhomoplasious apomorphy (8:0, tibial spurs absent). Clade I, composed by the species of Megistopus (exclusive of “M.” mirabilis) and Nedroledon was supported by just one homoplasious apomorphy (23:1) with a Bremer support value of 1. A clade composed by M. flavicornis and M. lucasi (clade J) was reconstructed as monophyletic based on one nonhomoplasious apomorphy (7:2, tarsomeres T1-T4 elongated, of the same size) and one homoplasious apomorphy (9:1) and garnered a Bremer support value of 2. The genus Nedroledon (clade K), whose monophyly was supported by one homoplasious apomorphy (24:1), received a Bremer support value of 1. N. anatolicus and N. iranensis were recovered as sister taxa based on one homoplasious apomorphy (25:0).
The genus Gymnocnemia was recently revised by
Gymnocnemia Schneider, 1845: 343.
Type species Megistopus variegatus Schneider, 1845.
Aplectrocnemus Costa, 1855: 18.
Type species Aplectrocnemus multipunctatus Costa, 1855 (Syn).
G. variegata, G. editaerevayae, and G. mirabilis.
Thorax covered with robust, outstanding setae. Legs extremely long and thin. Prothoracic leg much longer and slender than meso- and metathoracic legs. Femora differing in pattern: prothoracic femur with small dark dots, mesothoracic femur with two parallel dark stripes, metathoracic femur with an apical dark marking. Tibial spurs absent (G. variegata, G. editaerevayae) or present (G. mirabilis). Tarsomeres 1-4 of prothoracic leg differentiated in size, with the first (basal) tarsomere at least as long as one and a half times the second tarsomere. Tarsal claws opposable, closing against a brush of thickened setae on the fifth tarsomere. Male (based on G. variegata and G. editaerevayae): ectoproct rounded, gonocoxites 11 arch-like, gonocoxites 9 partly encircling gonocoxites, 11 plate-like in shape. Female: gonocoxite 7 tooth-like, gonocoxites 8 digitiform, gonocoxites 9 covered with digging setae, ectoproct provided with digging setae.
Larval morphology (based on G. variegata). Mandible long and slender, provided with 3 teeth of which the apical and median are subequal in size. Ocular tubercle prominent. Mesothoracic spiracle raised on tubercle. Meso- and metathoracic setiferous processes scolus-like. Odontoid processes atrophied. Rastra with the internal pair of digging setae smaller than the others (
Medium-sized antlion with a variegated habitus, with an alternating ochre and dark brown pattern. Forewing Rs at half the wing length with at least 8 crossveins. Thorax covered with outstanding black and white setae. Tibia without spurs. Tarsus shorter than tibia.
Gymnocnemia and Megistopus, habitus and details of thorax, lateral view. A Megistopus lucasi [Italy, Sardinia]; B Gymnocnemia variegata [Greece, Corfu]; C–D Gymnocnemia mirabilis [Oman, Jabal Shams], C thorax and prothoracic leg, D tarsus. Numbers indicate morphological characters of the data matrix and their state. Abbreviation: ta – tarsus.
Head and pronotum of Gymnocnemia and Megistopus, dorsal view. A–C Gymnocnemia variegata, variability, A Italy, Lazio, Rome [type locality], B Slovenia, C Italy, Sardinia; D Gymnocnemia editaerevayae, holotype [Morocco, High Atlas] (Photo: B. Michel, CBGP); E Gymnocnemia mirabilis [Oman, Jabal Shams]; F Megistopus flavicornis [France, Gard]; G Megistopus lucasi [Italy, Sardinia].
(Based on examination of 7 males and 8 females, see Suppl. material
G. variegata is a widely distributed but exceedingly localized species ranging from the Mediterranean to Central Asia (H.
Hölzel, 1980 (ODescr): H.
Small-sized antlion with a variegated habitus, with an alternating ochre and dark brown pattern. Wing venation sparse. Forewing Rs at half the wing length, with few crossveins. Thorax and prothoracic leg covered with prominent white setae. Tarsus exceptionally elongated, longer than tibia.
Head of Gymnocnemia and Megistopus, frontal view. A Gymnocnemia variegata [Italy, Sardinia]; B Gymnocnemia mirabilis [Oman, Jabal Shams]; C Megistopus flavicornis [France, Gard]; D Megistopus lucasi [Italy, Sardinia].
Wings of Gymnocnemia and Megistopus. A Gymnocnemia mirabilis [Oman, Jabal Shams]; B Gymnocnemia variegata [Italy, Lazio, Rome, type locality]; C Megistopus lucasi [Italy, Sardinia]; D Megistopus flavicornis [France, Gard]. Abbreviations: C – Costa, Sc – Subcosta, R – Radius, Rs – Radius sector, MA – Media anterior, MP – Media posterior, CuA – Cubitus anterior, CuP – Cubitus posterior, A – Anal vein.
Oman 18.VIII.1988 / Wadi Bani Kharus / Al Misra / M.J. Ebejer, 1 ♀ (NHMW). Oman: JABAL SHAMS / 23°14’N, 57°14’E 1900M / 3Jul95 BSKule, 4 ♀, 1 ex. (NHMW); OMAN: Jabal Shams / Wadi Dar Sawda, 1865 m / 23°14’N, 57°12’E, 20.X.95 / to light, MD Gallagher, 1 ♀ (NHMW).
Colouration. Vertex ochre dorsally and darker anteriorly. Frons dark brown. Clypeus dark brown with the ventral margin ochre. Labrum brown. Genae light brown. Maxillary and labial palpi ochre (Fig.
Dimensions (based on 5 specimens). Average body length 14.62 mm (min-max 14.02-15.23); forewing length 18.06 mm (16.66-18.25), ratio width/length 0.23; hind wing length 17.05 mm (15.39-19.04), ratio width/length 0.21.
Head. Vertex with a slightly raised transversal ridge (Fig.
Thorax. Pronotum slightly longer than wide. Pronotum and mesonotum covered with white outstanding setae (Fig.
Wings. Relatively narrow with an acute apex (Figs
Female genitalia. As for genus. Gonocoxite 7 relatively large and sclerotized, tooth-like in shape. Gonocoxites 8 covered with thin black setae. Gonapophyses 8 narrow, ribbon-like. Ectoproct with a short ventrocaudal projection, ventral side covered with stout setae.
The species was originally described from a single specimen found in the Sinai Peninsula (Egypt). The examined specimens are the first records of this species from Oman and the Arabian Peninsula, which significantly expands the known area of distribution (Fig.
Base of fore- and hindwing of Megistopus, showing diagnostic wing characters of Nemoleontini. A forewing of Megistopus flavicornis [Italy, Liguria]; B hindwing of same specimen. Abbreviations: C – Costa, Sc – Subcosta, R – Radius, Rs – Radius sector, MA – Media anterior, MP – Media posterior, CuA – Cubitus anterior, CuP – Cubitus posterior, A – Anal vein, cr – presectoral crossvein.
Megistopus Rambur, 1842: 343. Type species Megistopus bisignatus Rambur, 1842, by monotypy.
Legs long and slender, not abruptly differentiated in size. Tibial spurs as long as the first tarsomere. Tarsomeres 1-3 of prothoracic leg of similar size, tarsomere 4 shorter than the others. Tarsal claws opposable. Male: ectoproct rounded, gonocoxites 11 arch-like, gonocoxites 9 plate-like, converging apically. Female: gonocoxite 7 tooth-like, gonocoxites 8 digitiform, gonocoxites 9 provided with digging setae, ectoproct equipped with digging setae.
Mandible relatively long, armed with 3 teeth. Ocular tubercle prominent. Mesothoracic spiracle raised on tubercle. Mesothoracic setiferous processes scolus-like, metathoracic processes tubercle-like. Odontoid processes atrophied or absent. Rastra with the internal pair of digging setae less than 1/4 the length of the others (
Navás, 1912 (ODescr):
(Navás, 1912): H.
Hölzel, 1980 (misidentification
Medium-sized antlion with a mottled light and dark brown body pattern. Wings relatively narrow. Forewing Rs arising at the height of Cubital fork. Forewing gradates shaded with brown, cubital mark brown.
Megistopus lucasi: male and female terminalia, lateral view. A male [Italy, Sardinia]; B female [Italy, Sardinia]. Abbreviations: ect – ectoproct, S9 – sternite 9, gx9 – gonocoxite 9, S8 – sternite 8, gx8 – gonocoxite 8, S7 – sternite 7.
Male genitalia of Nemoleontini, complex of gonocoxites 9 + gonocoxites 11 sensu
Italy: Sardinia, Torre dei Corsari, 39°41,017’N 8°27,044’E, 28.V.2003, 50 m, H. and U. Aspöck and R.A. Pantaleoni leg, 3 ♀ (HUAC); Italy: Sardinia, Arbus (Oristano) / Torre dei Corsari, 39°41’23.09”N 8°27’14.50”E, coastal sand dunes /V.2010 D. Badano leg, 1 ♀ ex larva (DB); Italy: Sardinia, Arbus (Oristano)/ Torre dei Corsari, 39°41’23.09”N 8°27’14.50”E, coastal sand dunes /V.2011 D. Badano leg, 1 ♂ ex larva; Italy: Sardinia/ Chia (Cagliari), beach, XI.2011 (D. Badano), 1 ♂ ex larva (DB).
Colouration. Vertex ochre with paired dark markings. Paired blackish markings extend from the anterior margin of the vertex to the lateral margins of the frons, surrounding the base of the antenna. Frons light brown, except the lateral margins. Clypeus light brown with suffused dark markings. Labrum brown. Genae, maxillary and labial palpi testaceous. Antennae light brown, scape with dark brown suffusions, antennomeres darker basally (Fig.
Dimensions (based on 2 specimens). Body length 20.31, 26.66 mm; forewing length 21.43, 26.61 mm, ratio width/length 0.24; hind wing length 20.8, 25.87 mm, ratio width/length 0.20.
Head. Vertex slightly inflated. Distal palpomere of labial palpi fusiform, relatively thin, palpimacula elliptical. Antennae clavate.
Thorax. Pronotum longer than wide (Fig.
Male genitalia. As for genus. Gonocoxites 9 plate-like, converging apically (Fig.
Female genitalia. As for genus. Gonocoxites 8 dark brown, covered with thin black setae. Gonocoxites 9 and ectoproct covered with sparse stout setae (Fig.
The species is only known from few coastal localities of Algeria, Tunisia and Italy (Lazio and Sardinia) (
M. lucasi is an extremely rare species and is exclusively known from well preserved coastal sand dunes and beaches with junipers. The unmistakable larva of this species (see
|1||Forewing vein Rs originating at or distal to cubital fork; forewing vein 2A strongly bent before merging with 3A (Fig.
|–||Other combination of characters||other Myrmeleontidae|
|2||Thorax covered with outstanding, robust setae (Fig.
|–||Thorax without outstanding setae; prothoracic leg as long as the other legs||5|
|3||Frons dark brown (Fig.
|–||Frons light brown (Fig.
|4||Wings relatively broad; forewing with cubital and gradate markings faint (Fig.
|–||Wings relatively narrow; forewing with cubital and gradate markings dark, well evident (Fig.
|5||Tarsomere 1 as long as T2 or slightly shorter (Fig.
|–||Tarsomere 1 as long as T2+T3; male: abdomen longer than wings, ectoproct with ventrocaudal projections (Fig.
|6||Body pattern: blackish with pale areas in the abdomen; frons yellow (Fig.
|–||Body pattern: ochre with dark brown markings; frons ochre with brown markings below the base of the antennae (Fig.
Female genitalia of Gymnocnemia. A–B Gymnocnemia variegata [Greece, Corfu], A lateral view, B ventral view; C–D Gymnocnemia mirabilis [Oman, Jabal Shams], C lateral view, D ventral view. Abbreviations: ect – ectoproct, gp8 – gonapophysis 8, gx7 – gonocoxites 7, gx8 – gonocoxite 8, gx9 – gonocoxite 9.
Preferred tree of the two most parsimonious trees (83 steps, C.I. 0.687) obtained in the analysis. Black squares indicate nonhomoplasious apomorphies, white squares homoplasious ones. Letters indicate clades discussed in the text.
Strict consensus tree obtained from the two most-parsimonious trees with Bremer support values indicated above branches.
The phylogenetic analysis was conducted mainly to investigate the relationships among the species included in the genera Gymnocnemia and Megistopus and to test their respective monophyly. In addition, it is possible to draw some interesting, even if preliminary, observations on affinities at the tribal level. In agreement with
Our analysis consistently recovered Gymnocnemia mirabilis, originally described as a member of Megistopus, within the genus Gymnocnemia. The monophyly of this genus was well supported by elongated prothoracic legs, strikingly different pattern of leg pairs and the presence of robust, prominent thoracic setae. The characteristic shape of male gonocoxites 9, partly encircling gonocoxites 11, characterizes both G. variegata and G. editaerevayae (see also
The present analysis confirms the importance of a phylogenetic approach (i) to circumscribe genera as monophyletic groups in a family characterized by an overall homogeneous morphology, such as Myrmeleontidae, and (ii) to test the actual systematic value of commonly used diagnostic characters. A more inclusive taxon sampling and the integration of molecular data is advisable to delimit suprageneric taxa and to reconstruct the relationships within the family.
The present study was supported by the project SYNTHESYS (www.synthesys.info), which is financed by the European Community Research Infrastructure Action under FP7 programme. Special thanks to Susanne Randolf and Harald Bruckner (NHMW, Vienna) for their support during the visit of D.B. in Vienna. We also thank Bruno Michel (CIRAD CBGP, Montpellier) for the photos of the specimen in his care, and the careful review of the manuscript. Grateful thanks to Dusan Devetak (University of Maribor, Maribor), Alexi Popov (National Museum of Natural History, Sofia), and to Susanne Randolf for the critical review of the article and for valuable suggestions. Great thanks to Dr. John Plant (Guilford, Connecticut) for critically reading the manuscript and checking the English. We also acknowledge the Museum für Naturkunde, Berlin, for the support during the publication of this article.