Introduction

Despite lack of consistent reconstruction of phylogenetic relationships among major ensiferan lineages (Legendre et al. 2010), it is generally admitted that king crickets, raspy crickets and weta diverged from a Mesozoic paraphyletic assemblage, namely the hagloideans (Gorochov 2001; Heads & Leuzinger 2011; Sharov 1968, 1971). Because hagloidean males possess the ‘tettigonid-gryllid-like’ stridulatory apparatus in forewings (including a file, that defines the taxon Grylloptera nom. Haeckel, 1896, dis.-typ. Béthoux, 2012), some assumed that king crickets, raspy crickets and weta experienced a loss of the corresponding complex structures (Gorochov 2001; Sharov 1968, 1971; among others). However several authors recognized the wing venation of these insects as plesiotypic, compared to that of other ensiferans (Karny 1929b), and assumed a primary lack of the Grylloptera stridulatory apparatus (Zeuner 1939). The assumption that king crickets, raspy crickets and weta diverged from a group that already possessed the Grylloptera stridulatory apparatus, is primarily supported by the lack of evident Mesozoic remains.

The wing venation of extant king crickets, raspy crickets and weta is sparsely documented. Apart from Karny (1930), most papers by this author include incomplete drawings of wing venation (e.g., Karny 1928b). Several other contributions focus on species that most likely represent the most derived members of the group (e.g., Karny 1928a, Ragge 1955). Illustrations and observations by Zeuner (1939) are minimal and will prove partly inaccurate. This lack of data jeopardizes the identification of fossil stem-relatives of king crickets, raspy crickets and weta.

In order to obtain reference data for a discussion on the origin of these insects, a survey of the holdings of the Australian National Insect Collection and CSIRO Ecosystem Sciences (Canberra, Australia) was carried out, complemented by a survey of material housed at the Queensland Museum (Brisbane, Australia), and of a few other institutions. Preliminary results suggested that relatives of king crickets, raspy crickets and weta could be identified among Early Mesozoic orthopterans. This finding prompted a broadening of the scope of this contribution to include data on intra-specific wing venation variability in extant species, in order to assist delimitation of related fossil species.

The paper is organized as follows: a comparative analysis is carried out, aiming to develop wing venation topological homologies in selected species (and including data on intra-specific variability). Based on the resulting conjectures, a character-state-based nomenclatural treatment (i.e., cladotypic) is proposed, including the revision of several fossil taxa relevant for the following section; finally, conjectures and nomenclature are used to elaborate on evolutionary aspects.

Material and methods

I use the cladotypic nomenclatural procedure I elaborated (2007b, c; 2010) and Lanham's species names (Dayrat et al. 2004, Lanham 1965) [see introduction and application of this approach in JOR by Béthoux & Herd (2009)]. In order to avoid a mixing of names erected under distinct procedures, vernacular versions of taxon names erected under the Linnaean procedure will be used (e.g., gryllacrididaeans instead of Gryllacrididae). The position of the species surveyed in this contribution under the traditional nomenclatural procedure is provided in Appendix 1. In addition, traditional genera are indicated in this section, between inverted commas (except in Appendix 2). Appendix 2 provides a nomenclatural treatment at the specific and generic levels compliant with the ICZN. Appendix 3 provides provisional compositions for the newly defined taxa.

Specimens referred to as ANIC IWC OB, QM, PIN, NHM, and MHNG are housed at the Australian National Insect Collection and CSIRO Ecosystem Sciences (Canberra, Australia), the Queensland Museum (Brisbane, Australia), the Palaeontological Institute of the Russian Academy of Sciences (Moscow, Russia), the Natural History Museum (London, UK), and the Museum d'histoire Naturelle (Geneva, Switzerland), respectively. Other institutions are mentioned in the text.

Extant species and specimens used for this contribution are as follows [following Bengston (1988) for open nomenclature]:

Species cf. mexicanus de Saussure, 1859: 209 (currently assigned to the genus ‘Anabropsis’ Rehn, 1901 under the Linnaean nomenclatural procedure).

—The studied specimen was determined at the genus level. Since the type-species of the corresponding genus is mexicanus de Saussure, 1859: 209, I refer to the species as cf. mexicanus in the following.

—Specimen illustrated (ANIC pinned collection): ANIC IWC OB 26, ♀, Monte Verde, Powell Prop., ca 1580 m., Costa Rica, 23.iii.1974, determined by D.C.F. Rentz.

Species laudatum Johns, 1997 (genus “Transaevum Johns, 1997). —Determination of specimens of this species was complicated by the claim of the existence of two distinct and undescribed species in the genus by Johns (1997; balanced in Monteith & Field 2001). However, only one species has been formally described. Observation of the ‘type-series’ of the presumed second species housed at the Queensland Museum (material labeled ‘longihamatum’) revealed no distinctive features except a size comparatively smaller than material of laudatum, but nevertheless compatible with identity with this species. In addition, specimens of ‘longihamatum’ were collected from a locality where laudatum occurs, rendering the distinctiveness of ‘longihamatum’ unlikely according to CB. Monteith (pers. com. 2010). I presume that specimens housed at the ANIC and at the QM indicated as belonging to laudatum were determined by P.M. Johns; determination of two unassigned ANIC specimens by O. Béthoux was based on the fact that the set of ‘longihamatum’ specimens was composed of comparatively smaller specimens. All specimens were collected from Australia. Because of the large sample available, specimens belonging to the ANIC pinned collection with wings in resting position were excluded from the survey. The specimen of the paratype series of ‘longihamatum’ illustrated on Fig. 3H, Q was excluded from the measurement survey.

—Specimens prepared and illustrated (ANIC IWC OB 1–9 from ANIC ethanol collection; ANIC IWC OB 10 from ANIC pinned collection): ANIC IWC OB 5, ♀, 17.37S 145.34E, BS3 Massey Crk., 27.ii–26.iii.1996; ANIC IWC OB 6, ♀, 17.07S 145.38E, Mt. Haig Road, 10 kms. ENE of Tinaroo Dam Wall, Atherton Tableland, QLD, 03.ii. 1988; ANIC IWC OB 7, ♂, 17.02S 145.37E, Davies Crk., 15 km. from Kennedy Hwy., 21 km E. by S. of Mareeba, QLD, 01.iii.1988; ANIC IWC OB 8, ♀, 17.37S 145.34E, BS3 Massey Crk., QLD, 31.i–27.ii.1996; ANIC IWC OB 9, 3, 17.37S 145.34E, BS3 Massey Crk., QLD, 04.ii-06.iii.1995, determinedby O. Béthoux; ANICIWCOB 10, ♂, Kuranda, QLD, 27.ii.1981, determined by O. Béthoux; QM (unnumbered, ‘paratype series’ of ‘longihamatum’), ♂, Charmillin Ck. Xing, 950 m., Tully Falls Rd., QLD, 8.xii.l989-5.i.l990 (only left forewing prepared).

—Specimens used for measurements survey (in addition to the illustrated ones):

Females: ANIC pinned collection, 15.47S 145.17E, Moses Ck., 4 km NE of Mt. Finnigan, nr. Cooktown, QLD, 14.x. 1980 (paratype); ANIC pinned collection, Paluma, QLD, 17.x.1979 (paratype); ANIC pinned collection; Davies Ck. Rd., nr. Kuranda, QLD, 9.i.1979 (paratype); ANIC pinned collection; 16.50S 145.37E, 1.5 km S of Kuranda on Kennedy Hwy.‘Arona’, QLD, 10.xii.2003 (2 specimens); ANIC pinned collection; 17.25.43S 145.19.13E, Atherton Tableland, Mt. Hypipamee Nat. Pk., QLD, 1–12.ii.1998; ANIC ethanol collection, 17.05S 145.35E, Mt. Haig, 22 km NE by N of Atherton, QLD, 16.iii.1988 (2 specimens); 16.16S 145.02E, Forestry Camp Mt. Winsdor Tableland, NNW of Mt. Carbine, QLD, 23.1.1988; ANIC ethanol collection, 17.05S 145.35E, Mt Haig, 22 km NE by N of Atherton, QLD, 16.iii.1988; ANIC ethanol collection, 17.06S 145.36E, Mt. Haig, QLD, 4.ii–17.iii.1995; ANIC ethanol collection, 17.28S 145.29E, Longlands Gap, QLD, 5–27.ii.l996;ANICethanolcollection, 17.02S 145.37E, Davies Ck. (15 km from Kennedy Hwy.), 21 km E by S of Mareeba, QLD, 1.iii.1988 (2 specimens); QM ethanol collection, 16.26S 145.42E, Hughes Rd., Topaz, 4.xii. 1993, QLD (2 specimens, paratypes); QM ethanol collection, 16.05S 145.25E, upper roaring meg, 680m, 6 km W. of Capte tribulation, QLD, 10–22.xi.1993, (3 specimens, paratypes); QM pinned collection, Cardwell Range, Upper Broadwater Ck Valley, QLD, 17–21 .xii.1986, 700–80Om (only left forewing observed; paratype); QM pinned collection, Douglas Ck. Rd., Kirrama Range, QLD (only left forewing observed; paratype); QM pinned collection, Upper Broadwater Ck. Valley, Cardwell Range, QLD, 17–21 .xii.1986 (only right forewing observed; paratype); QM pinned collection, Head of Roots Ck, 12 km WNW Mossman, QLD, 28–29.xii.1990; QM pinned collection, Mt Lewis Rd., QLD.

Males: ANIC pinned collection, 32 km NW of Kennedy Hwy., QLD, 24.xii.1979 (paratype); ANIC pinnedcollection, 15.47S 145.17E, Moses Ck., 4 km NE of Mt. Finnigan, nr. Cooktown, QLD, 16.x.1980 (only left forewing observed; paratype); ANIC pinned collection; 18.59S 146.10E, Birthday Ck., 6 km. NW by W of Paluma, QLD, 25. ix.1980 (paratype); ANIC pinned collection, Kuranda, QLD, 27.ii.1981 (2 specimens; in one, only left forewing observed); ANIC pinned collection; 16.50S 145.37E, 1.5 km S of Kuranda on Kennedy Hwy. ‘Arona’, QLD, 10.xii.2003; ANIC ethanol collection, Mt. Lewis Rd., Julatten, QLD, 12.xi.1975 (paratype); ANIC ethanol collection, 17.02S 145.37E, Davies Ck. (15 km from Kennedy Hwy.), 22 km E by S of Mareeba, QLD, 24.iii.1988; ANIC ethanol collection, 17.05S 145.35E, Mt. Haig, 22 km NE by N of Atherton, QLD, 16.iii.1988; QM ethanol collection, 16.26S 145.42E, Hughes Rd., Topaz, QLD, 4.xii.1993 (paratype); QM ethanol collection, 1km S Cable Tower No7, Bellenden-Ker Range, QLD, 17–24.x.1981 (paratype); QM ethanol collection, upper roaring meg, 6 km W. Cape Tribulation, QLD, 8–9.xii.1993 (paratype); QM ethanol collection, upperlsley Ck., 9 km WSN of Edmonton, QLD, 29–30. xi.1993 (paratype); QM pinned collection, 17.16S 145.49E, Massey Range, 4 km W of Centre Bellenden Ker, 9–11.x. 1991 (holotype); QM pinned collection, 32 km NW of Kennedy, QLD, 24.ii.1979 (paratype); QM pinned collection, 17.14S 145.48E, Massey Range, 6 km NW of Centre Bellenden Ker, 11–12.xi.1991 (only right forewing observed; paratype); QM pinned collection, Charmillin Ck. Xing, Tully Falls Rd., QLD, 8.xii. 1989-5.i.1990 (2 specimens, both paratypes; only right forewings observed); QM pinned collection, head of Francis Ck, 12 km WNW Mossman, QLD, 30 xii.1989 (only right forewing observed); QM pinned collection, Mt Lewis Rd., 16km from Highway, QLD, 18.xii.1989-13.1.1990 (only left forewing observed).

Species ornata Willemse, 1963 (genus ‘Exogiyllacris’ Willemse, 1963).

—Determination of material of this species is straightforward as the species is very distinctive. All specimens were collected from Australia.

—Specimens prepared and illustrated (ANIC ethanol collection): ANIC IWC OB 15, ♂, 17.28S 145.29 E, Longlands Gap, QLD, 1–30.xi. 1995, determined by D. C.F. Rentz; ANICIWCOB 16, ♂, 17.35S 145.34E, BS3 Massey Crk., QLD, 3.x-2.xi.1995, determined by D.C.F. Rentz.

—Specimens used for measurements survey (in addition to illustrated specimens):

Females: ANIC, pinned collection, Atherton tableland, QLD (only right forewing observed); ANIC, pinned collection, no data, collected by JT Doyer (only right forewing observed); ANIC ethanol collection, 17.33S 145.32E, Mt Fisher, QLD, 4.ii–21.iii 1995; ANIC ethanol collection; 17.27S 145.29E, Hugh Nelson Rd. QLD, 1–30.xi.1995; ANIC ethanol collection, 17.15S 145.38E, Lake Barrine, Atherton Tableland, QLD, 6.xii. 1985; QM ethanol collection, 17.30S 145.36E Millaa Millaa Falls, 13–24.xi. 1994 (three specimens); QM ethanol collection, 16.26S 145.42E, Hughes Rd.,Topaz, QLD, 4.xii.1993; QM ethanol collection, 17.23S 145.28E, Upper Plath Rd., 9.xii.1995; QM ethanol collection, 17.17S 145.58E, Graham Range, 8–9.xii.1995; QM pinned collection, 17.26S 145.42E, Hughes Rd, Topaz, 6.xii.l993-25.ii.l994.

Males: ANIC pinned collection, Mt Haig, nr. Danbulla, QLD, 28.ix.1966; ANIC dry collection, Douglas Ck., lamb range, NE QLD, 12.x.1992 (only right forewing observed); ANIC ethanol collection, 17.37S 145.34E, Massey Ck., QLD, 2–20. xi.1995; QM ethanol collection, 17.28S 145.32E, Kenny Road, 25.xi.1994-10.1.1995; QM ethanol collection, 17.37S 145.46E, Palmerston NPE Margin, 10.xii.1995-7.ii.1996; QM ethanol collection, 17.27S 145.29E, Tower nr. the Crater NP, 25.xi.1994, 1230m; QM pinned collection, Bulurru, Topaz, 23.ix.1997 (only right forewing observed); QM pinned collection, 17.14S, 145.48E, Massey Range, 6 km NW of Centre Bellenden Ker, 11–12.x.1991 (only right forewing observed); QM pinned collection, 17.14S, 145.48E, Massey Range, 6 km NW of Centre Bellenden Ker, 11–12.x.1991.

Species cf. bicornis Karny, 1929a (genus ‘Gryllotaurus’ Karny, 1929a).

—An issue similar to that faced with laudatum was raised by the claim that two species, similar but distinct from bicomis exist, but are undescribed yet (Johns 1997; balanced in Monteith & Field 2001). The selected material (illustrated and measured) was labeled ‘biconulata’, presumably by P.M. Johns. Apart from a comparatively smaller size, material of ‘biconulata’ does not significantly differ from that of bicomis, therefore I consider the existence of a ‘biconulata’ dubious (specimens of intermediate size exist). In any case, the survey is based on material from a single species, either bicomis, or ‘biconulata’. At worst the intraspecific variability will have been underestimated. All specimens were collected from Australia.

—Specimens prepared and illustrated (all ANIC ethanol collection): ANIC IWC OB 11 and ANIC IWC OB 12, ♀, 17.06.S 145.37E, GS2 Mt. Edith, QLD, 4.ii–17.iii.l995; ANIC IWC OB 13, ♀, 17.33S 145.32E, BS2 Mt. Fisher, QLD, 21.iii-5.iv.1995; ANIC IWC OB 14, ♀, 17.25S 145.29E, GS3 Hugh Nelson Rd., QLD, 5.iii–4.iv.l995.

—Specimens used for measurements survey (in addition to the illustrated ones):

Females: ANIC ethanol collection, Mt Belleden-Ker, Summit, 11.iv.1979, QLD (indicated as ‘paratype’); ANIC ethanol collection, 17.33S 145.32E, Mt. Fisher, QLD, 3.x.-2.xi.1995 (fourspecimens); ANIC ethanol collection, 17.37S 145.34E, Massey Crk., QLD, 2–30.V.1996 (fourspecimens); QM ethanol collection, Belleden Ker Range, Summit TV Stn., QLD, 25– 31.x. 1981 (indicated as ‘paratype’); QM ethanol collection, paratype; Belleden Ker Range, NQ, Cable Tower 3, 1054m, 1–7.xi.1981 (only right forewing observed);

Males: ANIC ethanol collection, 17.33S 145.32E, Mt. Fisher, QLD, 3.x.-2.xi.1995; QM ethanol collection Belleden Ker Range, Summit TV Stn., 25–31 .x. 1981 (indicated as ‘holotype’).

Species pinguipes? Rentz in Morton & Rentz, 1983 (genus ‘Bothriogryllacris’ Rentz in Morton & Rentz, 1983).

—The studied specimen was collected prior to investigations by Morton & Rentz (1983). It is housed in the ANIC pinned collection, together with pinguipes specimens. However based on its forewing venation, the specimen differs from representatives of species by its higher number of RP branches. Therefore I assume it could belong to a distinct species.

—Specimen prepared and illustrated: ANIC IWC OB 24, ANIC pinned collection, ♂, Stuart Hwy., 296 km S. of Tennant Crk., NT, Australia, 29.xi.1972.

Species pinguipes Rentz in Morton & Rentz, 1983 (genus ‘Bothriogryllacris’ Rentz in Morton & Rentz, 1983).

—All specimens were collected from Australia and determined by D.C.F. Rentz. Only specimens from the ANIC ethanol collection were surveyed (i.e., pinned specimens were excluded), because this sample appeared sufficient. Two specimens were excluded from survey because of dubious determination. Collecting data of these specimens are as follows: 32.29S 124.13E, 59 km ESE of Balladonia Motel, WA, 29.1.1991.

—Specimens prepared and illustrated: ANIC IWC 17, ♂, ca 30.39S121.27E,7–11km N of Kalgoorlie„ WA, 17.xi.1978; ANIC IWC OB 18, ♀, ca 30.39S 121.27E, 7–11 km N of Kalgoorlie, WA, 17.xi.1978; ANIC IWC OB 19, ♀, 24.51S 133.HE, Stuart Hwy., 41 km N of Erlunda, NT, 4.xi.l988: ANIC IWC OB 20, ♀, 20.52S 130.16E, Tanami Desert, NT, 21 .iii.1988; ANICIWC OB 21, ♀, 23.17S 133.39E, 50 km. NNW of Alice Springs, NT, 15.xii.1989; ANIC IWC OB 22, ♂, ca 20 km E of Norseman, WA, 12–13.i.1986; ANIC IWC OB 23, ♂, 32.13S, 123.15E, 38 km. WNW of Balladonia Motel, WA, 22.xi.1980.

—Specimens used for measurements survey (in addition to the illustrated ones):

Females: 26.00S 131.25E, 26 km WSW of Milga Pk., NT, 18.i.1982 (paratype); 26.09S 130.35E, 56 km W of Amata, Musgrave Ranges, SA, 20–21.i.1982 (paratype); 25.12S 132.05E, Lasseter Hwy., 35 km ENE. of Curtin Springs, NT, 8.xi.1988; 25. HS 133.11E, Stuart Hwy., 4 km N of Erlunda, NT, 4.xi.1988; 20.52S 130.16E, Tanami Desert, NT, 28.iii.1988; 30.42S 121.28E, 4 km N by E of Kalgoorlie, WA, 17.ii.1978 (6 specimens); 25.11S 133.1E, Stuart Hwy., 4km N. of Erlunda, NT, 4.xi.1988; 25.21S 131.53E, Lasseter Hwy., 13km E by S of Curtin Springs, NT, 8.xi.1988 (2 specimens); ca. 30.39S 121.27E, 7–11 km N of Kalgoorlie, WA, 17.xi.1978 (4 specimens); 30.47S 121.27E, Kalgoorlie (airport), WA, 16.ii.1978 (4 specimens); 24.51S 133.11E, Stuart Hwy., lkm N of Erlunde, NT, 4.xi.1988; 28.47S 121.31E, 1 km S of Malcolm, WA, 19.ii.1978 (2 specimens).

Males: 26.00S 131.25E, 26 km WSW of Milga Pk., NT, 18.i.1982 (paratype); 26.00S 131.25E, 26 km WSW of Milga Pk., NT, 18.1.1982 (2 specimens; both paratypes); ca. 20 km E of Norseman, WA, 12–13.i.1986 (2 specimens); 1 km W of Sangsters Bore, Tanami Desert, NT, 20.iii.1985; 25.11S 133.11E, Stuart Hwy., 4 km N of Erlunda, NT, 27.x.1988; 20.52S 130.16E, Tanami Desert, NT, 18.iii.1988 (2 specimens); ca. 30.39S 121.27E, 7–11 km N of Kalgoorlie, WA 17.11.1978; 30.47S 121.27E, Kalgoorlie (airport), WA, 16.ii.1978 (4 specimens); 28.47S 121.31E, 1 km S. of Malcolm, WA, 19.ii.1978; ca. 30.39S 121.27E, 4 km N by E of Kalgoorlie, WA 17.11.1978.

Species ‘sp. undet.’.

—The corresponding specimen belongs to a set labeled as ‘Gryllacrididae Genus 3 Sp. 1’ in the ANIC pinned collection.

—Specimen illustrated: ANIC IWC OB 25, S, Yass, NSW., Australia, 13.xii. 1932.

Species punctipennis Walker, 1869 (genus ‘Xanthogryllacris’ Karny, 1937).

—All surveyed specimens belong to the ANIC ethanol collection, were determined by D.C.F. Rentz, and were collected from Australia.

—Specimens prepared and illustrated: ANIC IWC OB 36, ♀, 11.45S, 142.35E, Heathland, QLD, 1–21.iii.1992; ANIW IWC OB 37, ♂, 15.50S, 145.20E, Gap Ck., 5 km. ESE of Mt. Finnigan, nr. Cooktown, QLD, 13–16.V.1981; ANIC IWC OB 38, ♂, 15.28S, 145.15E, 1km. w. of Cooktown, QLD, 12–13.V.1981. —Specimens used for measurements survey (in addition to the illustrated ones):

Females: 12.45S 143.17E, 8km E of Mt.Tozer, nr. Iron Range Nat. Pk., QLD, 8.vii.1986; 12.44S 143.16E, 6 km ENE of Mt. Tozer, nr. Iron Range Nat. Pk., QLD 1 .vii. 1986; Shiptons Flat, nr. Cooktown, QLD, 16–18.v.1981; 15.50S 145.20E, GapCk., 5 km ESE of Mt. Finnigan, nr. Cooktown, QLD, 13–16.V. 1981; 11.45S 142.35E, Heatlands, QLD, 18.ix.1992–21.x.1992.

Males: 12.43S 143.17E, 9 km ENE of Mt. Tozer, Iron Range Nat. Pk., QLD, 10.vii.1986; 12.44S 143.14E, 9 km ENE of Mt. Tozer, nr. Iron Range Nat. Pk., QLD, 28.vi.1986-4.vii.1986; 12.43S 143.17E, 9 km ENE of Mt. Tozer, Iron Range Nat. Pk., QLD, 10.vii.1986; 15.50S 145.20E, Gap Ck., 5 km ESE of Mt. Finnigan, nr. Cooktown, QLD., 13–16.V.1981; 15.04S 145.07E, Mt. Webb Nat. Pk., nr. Cooktown, QLD, 27–30. iv.1981; 17.00S 145.50E, Pine Ck. (nr. CSIRO Tower), 11 km SE by S of Cairns, QLD, 18.ii.1988; 17.53S 146.09E, Dunk Island (airstrip), QLD, 17.iv.1990.

Species rufovaria Kirby, 1888 (genus ‘Gryllacris’ Audinet-Serville, 1831).

—All specimens were collected from Australia, and all are housed at the ANIC (ethanol collection).

—Specimen prepared and illustrated: ANIC IWC OB 27, ♂, 10.27S, 105.40E, Christmas Island, Grants Well to Irvine, Hill Rd. track,, 27.iv. 1989, presumably determined by D.C.F Rentz. —Specimens used for measurements survey (in addition to the illustrated one; ♂ and ♀ treated together because of limited sample size):

Males: 10.27S 105.40E, Grants Well to Irvine, Hill Rd. track, Christmas Island, 27.iv.1989; 10.29S 105.38E, near Central Area, Workshop, Christmas Island, 17.iv.1989; 10.30S 105.55E, EW Parl track, Christmas Island, 28.iv.1989.

Female: 10.27S 105.40E, Grants Well to Irvine, Hill Rd. track, Christmas Island, 27.iv.1989.

Wings of illustrated extant specimens were cut off specimens preserved dry or under ethanol, softened as necessary, and mounted in white Euparal (Asco Laboratories, Manchester, UK). Left wings were mounted on their dorsal side, right wings on ventral side. As far as possible, wings were mounted unfolded.

Photographs reproduced on Figs 2–9, 11–19, 22, 23, 27, 28 were taken with a Canon EOS 450D digital camera coupled to a Canon 50 mm macro lens (and an extension tube as appropriate), and to a Canon MP-E 65 mm macro lens. Photographs reproduced on Figs 1, 21 were taken with a Nikon Coolpix C2500L. Other photographs were provided by various colleagues. For wings of extant material, transmitted light was obtained from a VisiLED ACT Basis for small wings, and from a regular light table for larger ones. Photographs were dust off manually using a stamp tool, and processed with Adobe Photoshop. Unless specified, photographs of fossil specimens were taken under dry conditions.

Except for drawings reproduced in Figs 1 and 24, drawings were prepared with the aid of a Zeiss SteREO Discovery V8 stereomicroscope equipped with a pair of W-PL 10×/23 eye pieces, a Plan Apo S 1.0x FWD objective, and a camera lucida. Due to the occurrence of structural folds in fore- and hind wings, and as a result of the compression of the three-dimensional structure that an insect wing is, creases occurred in many preparations of extant material, in particularly in hind wings. For example, ScP seems to be located posteriorly with respect to RA on Fig. 18, but is actually ‘creased’ below RA. In forewings, the membranous area posterior to AA veins was also problematic to mount unfolded, because it tends to fold even once cover glass has been put on. Drawings were corrected manually, and with the aid of Adobe Photoshop for hind wings vannuses (distortion transformation).

I use the conjecture of topological homology proposed by Béthoux & Nel (2001) for orthopterans, itself based on the serial insect wing venation pattern (Lameere 1922, 1923). Corresponding abbreviations and color-coding are as follows: CP, posterior Costa; ScA, anterior Subcosta; ScP, posteriorSubcosta; R, Radius; RA, anterior Radius; RP, posterior Radius; M, Media (green); MA, anterior Media; MP, posteriorMedia; CuA, anterior Cubitus (orange); CuP, posterior Cubitus; CuPa, anterior branch of CuP (purple); CuPb, posterior branch of CuP; CuPaα, anteriorbranch CuPa (blue); CuPaβ, posterior branch of CuPa (red); AA1, first anterior Analis. Correspondence with some other conjectures for orthopterans can be obtained from Béthoux & Nel (2002a: table 2) and on Fig. 10. The criticism expressed by Gorochov (2005) regarding Béthoux & Nel's (2001, 2002a) conjecture are addressed in Béthoux (2007a). Subsequent comments by Rasnitsyn (2007) are addressed in Béthoux (2008).

Other abbreviations are as follows: LFW, left forewing; LHW, left hind wing; RFW, right forewing; RHW, right hind wing. A vein (or vein sector) is said to be convex if located on an elevation, and concave if located in a depression. I refer to cross-veins aligned longitudinally to main veins, which are secondary structures, as ‘intercalary veins’.

Several transformation types are mentioned in the following and are better defined here. A translocation is the complete fusion of a vein (or one of its branches) with a surrounding vein (or branch), from the origin of the former, or from the wing base even. This transformation was documented in Pantcholmanvissiida nom. Béthoux 2007d, dis. Béthoux & Nel 2002b, typ. Béthoux 2007d (Béthoux 2007d) and in mantodeans (Béthoux & Wieland 2009). The term ‘abbreviation’ refers to the complete lack of forewing apex, without alteration of the basal area morphology, common among orthopterans. The term ‘pectinate fusion’ refers to a case involving the successive pectinate emergence of branches of a vein fused with another, the former lacking a distinct main stem (documented in neuropterans, see Shi et al. 2012).

Measurements of forewing length of unprepared material were made using a caliper. Although measures were made once only, a confidence interval can be estimated based on the largest difference observed between left and right forewings of a single individual. This difference amounts to 1.1 mm. This estimated error accounts for error in judging the positions of the measurement points, and the accuracy of the reading. It is considered constant (i.e., does not vary with wing size). Width of the area between M and CuPaα, opposite the free part of CuA (diverging from M + CuA, before its fusion with CuPaa), was measured under a microscope equipped with a graticule. Apositive figure represents a free CuA, while a negative one indicates (1) that a M + CuA + CuPaa fusion occurs (i.e., there is no free part of CuA: e.g., Fig. 8E, G), and (2) the length of this fusion. This approach was prompted by the fact that the series of states ranging from ‘long free part of CuA’ to ‘long M + CuA + CuPaa’ form a continuum (Fig. 13M–Q) better appreciated by a single measurement.

Fig. 1.

Species †uralica Sharov, 1968, holotype (PIN 1700/4153), drawing and photograph (right forewing, negative imprint) (see text for abbreviations and color-coding). For color version, see Plate III.

Comparative analysis

The rationale of such analysis is clarified in Béthoux (2012: 44). In summary, conjectures of topological homology (=THCs) are established so that the amount of transformation that has to be assumed to explain differences between patterns is minimized. Ideally, sets of THCs (=STHCs) can be compared based on the total amount of transformation assumed by each STHC However the methodology for this step is still little developed. In the following competing STHCs (see Fig. 10; ‘classical’ STHC including Fig. 10EH; alternative STHC including Fig. 10I–N) will be compared based on the occurrence (or lack thereof) of predicted intermediate patterns, and their consistence with respect to observed variants. This approach is grounded in that any given STHC can be challenged by a new, more performing one.

The wing venation of cf. mexicanus de Saussure, 1859: 209, laudatum Johns, 1997, ornata Willemse, 1963, cf. bicomis Karny, 1929a, pinguipes? Rentz in Morton & Rentz, 1983, pinguipes Rentz in Morton & Rentz, 1983, sp. undet., punctipennis Walker, 1869, and rufovaria Kirby, 1888, will be compared successively in the following. These species were selected based on their position as putative intermediates (i.e., exhibiting a unique pattern), reliable determination at the species level, and/or abundance of specimens available to the author. Variations observed in several aspects of wing venation are summarized in Appendix 4.

The fossil species †uralica Sharov, 1968 (Fig. 1) was selected to represent the presumed plesiotypic ‘oedischioid’ forewing morphology. As with most other Permian stem-orthopterans and Carboniferous relatives (Béthoux et al. 2012), †uralica possesses a common stem composed of M (green) and CuA (orange), a branched M (into MA and MP), a distinct free part of CuA, and CuP (purple) giving rise to CuPa (purple) and CuPb, a CuPa giving rise to CuPaα (blue) and CuPaβ (red), and a fusion of CuA with CuPaα resulting into a composite stem. Béthoux (2007d) suggested that CuA is simple in oedischoids and groups that derived from them. A simple CuA is conjectured for †uralica and in the following.

Species ornata

This species proved to have forewings often exhibiting rare and unusual traits, to the point that a usual condition was impossible to delimit based on the available material. The most noticeable aspect regards the organization of RP. As observed on forewing represented on Fig. 6A,B,D,E,F,H, the stem of R emits several posterior stems earlier to the usual RA anterior branches, and sometimes additional posterior stem(s) distal to the first anterior ones. These successive posterior stems are interpreted as RP branches. In other words, it is assumed that a pectinate fusion of RP with RA occurs. An alternative conjecture would be that the first posterior branch is MA, but this option is easily discarded: (1) as in †uralica, cf. mexicanus, and laudatum, the stem of M near the origin of CuA is forked into two simple branches, therefore the anterior stem of M observed in ornata is likely the genuine MA, and (2) the left forewing of the specimen ANICIWCOB 16 (Fig. 6C,G) has a MA actually translocated onto R (demonstrated by the basal origin of this stem, and the simple free part of M, hence composed of MP alone), yet it possesses ‘successive posterior branches of R [indeed RP]’. The organization resulting from the pectinate fusion is variable: points of divergence of the successive RP branches, and the number of their own branches, vary widely. As a consequence apical branches are not always evident to assign either to RA or RP, and the range of RP branches is difficult to delimit (ca 4–6).

In this species it is nearly equally common that CuA diverges distal to the MA / MP fork (Fig. 6A,E; i.e., CuA is fused with MP for a short distance before diverging), basal to the MA / MP fork (as in species surveyed above), or that CuA is not distinguishable from surrounding cross-veins (Fig. 6D,H). The branching pattern of CuA + CuPaα is also variable: usually its two components diverge distally (Fig. 6A, E), but can keep fused (Fig. 6C,G; Appendix4). Intermediate conditions occur, either with CuA (Fig. 6D,H) or CuPaα vanishing. A rare and informative case was observed in the right forewing of the specimen ANIC IWC OB 15 (Fig. 6B,F). At the first glance, a ‘Vein-like portion’ (indicated as ° on Fig. 6B) could have been considered as homologous to the basal section of CuPaα, as usual, and as observed in the other wing of the same individual (°on Fig. 6A). However, this conjecture must be discarded: in this particular forewing the portion of CuPaα distal to the origin of CuA (indicated as * on Fig. 6B) diverges from CuPa, i.e., is not fused with CuA. As a consequence the structure indicated as °on Fig. 6B is a ‘phantom CuPaα’, i.e., a remnant of the usual course of CuPaα. This is possibly a case of ‘tracheal un-capture’ (see Béthoux 2012 for this transformation type).

Fig. 2.

Species cf. mexicanus de Saussure, 1859: 209, specimen ANIC IWC OB 26 (♀; A—F at the same scale; G—H at the same scale; A, * indicates CuPaα); A, D, drawing and photograph of the left forewing; B, E, drawing and photograph of the right forewing; C, F, drawing and photograph of the left hind wing; G, detail of the left forewing, as located on D; H, detail of the left forewing, as located on E; I, J, detail of the left forewing, as located on D.

Fig. 3.

Species laudatum Johns, 1997 (all at the same scale); A—B, J—K, specimen ANICIWC OB 7 (♂); A, J, drawing and photograph of the right forewing; B, K, drawing and photograph of the left forewing; C, L, specimen ANIC IWC OB 5(♀), drawing and photograph of the left forewing; D, M, specimen ANIC IWC OB 6 (♀), drawing and photograph of the left forewing; E, N, specimen ANIC IWC OB 10 (♂), drawing and photograph of the left forewing.

F—G, O—P, specimen ANIC IWC OB 8 (♀); F, O, drawing and photograph of the right forewing; G, P, drawing and photograph of the left forewing; H, Q, specimen QM (♂), drawing and photograph of the left forewing; I, R, specimen ANIC IWC OB 9 (♂), drawing and photograph of the left forewing.

Fig. 4.

Species laudatum Johns, 1997, details as located on Fig. 2; A–B, specimen ANIC IWC OB 7 (♂), detail of costal area; A, right forewing, arrows indicate the first ScP branch (reaching ScA); B, left forewing, arrow indicates the origin of the first ScP branch (reaching anterior wing margin); C—G, detail of medio-cubital area; C, specimen ANIC IWC OB 5(♀), left forewing; D, specimen ANIC IWC OB 6(♀), left forewing; E, specimen ANIC IWC OB 10 (♂), left forewing; F, specimen ANIC OWC OB 8 (♀), left forewing; G, specimen ANIC IWC OB 9 (♂), left forewing.

Hind wings of ornata lack the pectinate fusion of RP with RA, and lack the RP + MA fusion observed in laudatum. The number of branches of RP varies between 3 and 4 (based on two hind wing pairs). A single intercalary vein occurs between MP and CuA + CuPaα + CuPaβ. A noticeable trait is the very basal origin of RP.

Species cf. bicornis

Illustrations of parts of the wing venation of bicornis are available from Karny (1929a: fig. 29). A more comprehensive treatment will prove necessary. Owing to their comparatively long ScA and short ScP, forewings of cf. bicornis might be abbreviated.

The usual organization of the connection of CuA with CuPaα is represented on Fig. 8C,I (and see Appendix 4). It involves a short CuA diverging from M near the point of divergence of MA and MP, and fusing with CuPaα. However, a M + CuA + CuPaα fusion (therefore without free part of CuA) was observed in 6 cases (4 of them composing 2 forewing pairs of single individuals; Fig. 8D–F,J–L). The standard deviation of the width of the ‘CuA area / length of M + CuA + CuPaα’ is higher than in other species (Table 1), indicating an unusual variability of this character in cf. bicornis. Notice that standard deviation falls down to 0.44 if forewings of the specimen ANIC IWC OB 12 (the one with the two longest MP + CuA + CuPaα fusions observed; Fig. 8E–F, K–L) is removed. Yet it is higher than in the other surveyed species.

In addition, the free part of CuA was observed to diverge distal to the MA / MP fork (i.e., CuA is briefly fused with MP) in 6 cases (Fig. 8A,G). Also, MP was observed to diverge from CuA + CuPaα (is re-routed along the free part of CuA) in 2 cases (Fig. 8B,H).

Except from the area involving the free part of CuA (or the lack thereof), the forewing venation of this species is comparatively stable. Veinlets of ScP rarely reach ScA (Appendix 4), RP is usually simple (20 cases), sometimes forked (13 cases; including 3 presumed cases of ‘RA + RP pectinate fusion’).

The hind wing morphology of cf. bicornis is unusual in that M + CuA and CuPa are very inconspicuous, and fuse near the wing base (Fig. 9). Although inconspicuous, a CuPaβ distinct from CuA + CuPaα was observed, as in species surveyed above (both veins fuse before reaching the wing margin). Another noticeable aspect is the occurrence of strong ScP branches. Such organization is common in forewings of orthopterans, but its occurrence in hind wings seems unique (at least among species considered herein).

Fig. 5.

Species laudatum Johns, 1997 (A—B, D—E at the same scale); A—D, specimen ANIC IWC OB 10 (♂); A—B, left hind wing, drawing and photograph; C, detail as located on B; D, portion of right hind wing, photograph; E, specimen ANIC IWC OB 7 (♂), portion of left hind wine. photograph.

Species pinguipes?

The corresponding specimen is essential to the establishment of forewing venation homologies of the species surveyed thereafter. Two possible transformation series are summarized on Fig. 10. They concern topological homologies of the medio-cubital area in forewings [according to Béthoux & Nel's (2001) conjecture and nomenclature for the whole orthopterans; correspondence with the Zeuner's (1939), Sharov's (1968, 1971), and Ragge's (1955) conjectures, under which the structure indicated as * is MP, is indicated by annotations in gray; annotations in light gray follow Karny's (1928b) conjecture, under which the structure indicated as ^* is a cross-vein].

The plesiotypic condition observed in †uralica (see Fig. 1) is schematized on Fig. 10A, and two patterns observed in pinguipes and closely related species are schematized on Fig. 10B,C. The aim of this section is to interpret these two patterns according to the conjecture of topological homologies posited for †uralica. Fig. 10F represents a conjecture proposed by Sharov (1968, 1971: fig. 27.B; see also Karny 1928b: figs 2, 9), assuming a simple M (Sharov's ‘MA’), and a 3-branched CuA + CuPaa (Sharov's ‘MP + CuA1’). According to this author, his ‘MA’ is branched in †uralica, so the implied transformation is M becoming simple. Another needed transformation involves the branching pattern of his ‘MP + CuA1’ (here CuA + CuPaα), splitting into a simple ‘MP’ (here CuA) and a forked ‘CuA1’ (here CuPaα).

Fig. 10G represents the conjecture for the pattern reproduced on Fig. 10C according to Zeuner (1939: pl. 2, upper figure). This conjecture assumes a simple M (Zeuner's ‘MA’) and a 2-branched CuA + CuPaα (Zeuner's ‘MP + CuA’). Provided that Zeuner assumes a forewing venation ground plan of orthopterans with a branched ‘MA’ (Zeuner 1939: pi. 9, upper figure), this author assumes the same transformation as Sharov (1968, 1971), viz. M (or ‘MA’) becoming simple in pinguipes and closely related species.

Two patterns can bridge those represented on Fig. 10A and Fig. 10F. One of them is represented on Fig. 10E, and assumes that the branching pattern of CuA + CuPaα was modified first. The other one (not represented) assumes that M becomes simple first. The most critical aspects of the argument being made here is that neither of these two patterns has ever been observed in actual specimens.

To date, another transformation series has never been considered. Starting from Fig. 10A, it assumes that CuPaα becomes simple (Fig. 101). The corresponding pattern fits that observed in cf. mexicanus, laudatum, ornata, cf. bicornis, and was observed in the left forewing of the specimen ANIC IWC OB 24 (Fig. 11A,C,E,G). In the last case, this morphology is unusual for the species.

Then (Fig. 10J) MP is assumed to be re-routed together with the free part of CuA; the composite stem fuses with CuPaα; MP and CuA + CuPaa diverge distally; CuA and CuPaα diverge distally. Such a re-routing of MP was documented by Sharov (1968, 1971) in the Gigantitanidae (and see Béthoux 2007d: fig. 1D). This transformation was also documented in cf. bicomis (rare occurrences), and is therefore highly plausible.

Fig. 6.

Species ornata Willemse, 1963 (all ♂; all at the same scale; * indicates the portion of CuPaα distal to its divergence from CuA; ° indicates the portion of CuPaα basal to its fusion with CuA, or its ‘phantom’; × indicates RA + RP partim stems); A—B, E—F, specimen ANICIWC OB 15; A, E, left forewing, drawing and photograph; B, F, right forewing, drawing and photograph; C—D, G—H, specimen ANIC IWC OB 16; C, G, left forewing, drawing and photograph; D, H, right forewing, drawing and photograph.

Continued.

This alternative transformation series supposes the occurrence of a common stem MP + CuA. Direct evidence for this common stem is provided by the right forewing of the specimen ANICIWC OB 24 (Fig. 11B,D,F,H). Exceptionally enough, this wing lacks the fusion of CuA (or MP + CuA) with CuPaα, present in all orthopterans since the Late Carboniferous (at least). Homologies in the right forewing are readily established based on the left one (compare Fig. 11E and F): the distal portions match, with a two-branched M, and simple CuA, CuPaα, and CuPaβ. Tracing MP and CuA backwards in the right forewing reveals that they form the common stem MP + CuA, as predicted.

This specimen indicates that the most plausible interpretation of the pattern represented on Fig. 10B is provided on Fig. 10J. From this step, interpretation of the pattern represented on Fig. 10C is straightforward: CuA and CuPaα keep being fused up to the posterior wing margin (Fig. 10 K).

Species pinguipes

Forewing THCs in this species are mainly established based on the previous case, but an additional specimen belonging to pinguipes supports the ‘MP + CuA’ conjecture (schematized on Fig. 10I–K). Forewings of this specimen are represented on Fig. 12. In both forewings M + CuA (presumably) separates into MA and MP + CuA, but MP diverges from MP + CuA and re-unites with MA, ‘re-forming’ the M stem observed in previous species. Another interesting aspect of this specimen regards CuA + CuPaα. This stem is easily identified in both wings as being located posterior to MP, and anterior to CuPaα (itself easily located based on the location of its point of origin, similar in both wings). The point is that CuA + CuPaα is forked in the left forewing (Fig. 12A,C) and simple in the right one (Fig. 12B,D). The former case in uncommon in this species and was observed in females only (Table 1). This specimen validates the transformation assumed to have occurred between patterns represented on Fig. 10B and Fig. 10C (namely, CuA + CuPaα becoming simple): apart from the brief MA — MP reunification, its left and right forewings are schematized in Fig. 10J and Fig. 10K, respectively.

The species could be surveyed based on a large sample (Appendix 4), a selection of which is represented in Figs 13–14. A trait that significantly varies is the connection of CuA with CuPaα. The usual organization is represented on Fig. 13G,J, with a short free part of CuA. A complete series ranging from a long free CuA (Fig. 13A,D,M) to a long M + CuA + CuPaα fusion (longest fusion represented on Fig. 131,L,Q) was documented in both males and females (Fig. 13; Appendix 4).

Unusual morphologies are documented in Fig. 14. Although the right forewing of the specimen ANIC IWC OB 20 possesses the usual CuPaα stem fusing with CuA (emerging from M + CuA; Fig. 13A,D,M),its left forewing has no such free CuPaα stem (Fig. 14A–B, K). Therefore it is assumed that CuPaα is translocated onto M + CuA. The left forewing of the specimen ANIC IWC OB 17 (Fig. 14D,F,L) has a MA fused with R opposite the fusion of MP and CuA with CuPaα. The fusion of MA with the radial system continues on along RP; at some point MA diverges from RP, shortly fuses with MP, and recovers its usual route. The right forewing of the same specimen (Fig. 14C, E) has the usual MA distinct from R and RP. Finally, the specimen ANICIWC OB 21 has both forewings with a branched CuA + CuPaα (Fig. 14G–J; already observed in the specimen ANIC IWC OB 19, Fig. 12A,C), and the left forewing has a short RP + MA fusion (Fig. 14H,J).

Instead of having a stem of R straight, as in species surveyed above, the hind wings of pinguipes possess a R stem posteriorly bent basal to the point of divergence of RP (Fig. 15). Another peculiar aspect regards M. A fusion of a ‘median element’ with RP is evident (Fig. 15C). Posterior to this ‘median element’ occurs the genuine CuA + CuPaα, identified based on its distal fusion with CuPaβ. Whether the ‘median element’ is MA, or M, will be demonstrated based on the next case. Suffices to notice that (1) the free part of the ‘median element’ is simple (genuine RP branches are located very distally from its origin), and that (2) two intercalary veins occur between the free part of the ‘median element’ and CuA + CuPaα + CuPaβ (unlike in previous species, in which a single intercalary vein occurs). Therefore two plausible options are to be considered: (a) a fusion of M with RP occurs (i.e., MA and MP stay fused); or (b) a fusion of RP with MA occurs, and MP runs together with CuA + CuPaα, as in forewings (but does not diverge distally).

Fig. 7.

Species ornata Willemse, 1963, specimen ANIC IWC OB 15 (♂), left hind wing; A, B, drawing and photograph (same scale); C, detail as located on B.

With respect to previous species, a peculiar trait is the organization of RA branches: in both fore- and hind wings, they arise basal to the first RP fork, and run parallel and close to the main RA stem for some distance.

Species punctipennis

The usual organization of M, CuA, and of branches of CuPa in forewings of punctipennis is represented on Fig. 17A,E, and is schematized on Fig. 10D. As aptly recognized by Karny (1910), Zeuner (1939), and Ragge (1955), MA (‘M’ of previous authors) is fused with R at the wing base (Fig. 10H,L–N). It is the most plausible conjecture for the occurrence of a single stem basal to the RA / RP fork. Although tracheae were uneasy to observe in material of punctipennis, the stem of ‘R’ is provided with two evident tracheae in forewings of rufovaria (Fig. 19E), viz. R and MA.

Several conjectures are possible forthe remaining medio-cubital elements. According to Ragge (1955: fig. 54), M is simple and no fusion of ‘MP’ (here CuA) occurs, and ‘Cu_1a’ (here CuPaα) and ‘Cu_1b’ (here CuPaβ) diverge distally. However, this option is balanced in the text (pp. 76–77), in which this author suggested that ‘MP’ (CuA) could be fused with ‘Cu₁’ (CuPa). This option is schematized on Fig. 10H. An issue with this interpretation is that it can hardly account for the occurrence of the pattern observed in the left forewing of the specimen ANIC IWC OB 37 (Fig. 17B,F; schematized on Fig. 10L): it would necessitate a branched ‘Cu^1b’ (here CuPaβ) (but this vein is simple in all orthopterans), or a translocation of ‘Cu1a’ (here CuPaα) onto ‘Cu^1b’ (here CuPaβ). A more plausible option is based on one of the THC proposed for pinguipes and schematized on Fig. 10K), implying the occurrence of a common stem MP + CuA + CuPaα which splits into MP and a simple CuA + CuPaα. Applying this pattern to the left forewing of the specimen ANIC IWC OB 37 (Fig. 17B, F) leads to the interpretation schematized on Fig. 10L: in both cases (Fig. 10K, L), posterior to MA successively occur the simple MP, then the composite (and simple) CuA + CuPaα stem, and then CuPaβ. On Fig. 10L, if traced backwards, the course of these elements reveals a composite stem CuA + CuPaα + CuPaβ, and a more basal composite stem MP + CuA + CuPa (CuPa or CuPaα + CuPaβ). No evidence was found for a ‘composite anterior stem’ [i.e., composed of MP and other ‘elements’, as suggested by Ragge (1955)]. Therefore it is assumed that in the usual forewing morphology of punctipennis MP + CuA + CuPa / CuPaα + CuPaβ divides into MP and a simple CuA + CuPa / CuPaα + CuPaβ (Fig. 10M). Among specimens of punctipennis, a single forewing was observed with an undivided composite MP + CuA + CuPa / CuPaα + CuPaβ stem (Fig. 17D,H), but it is not uncommon to observe the first fork of MP + CuA + CuPa / CuPaα + CuPaβ in a distal position (Fig. 17C,G). One specimen was observed with MP diverging from MA (hence M; Appendix 4). Very probably CuPa does not actually forks into CuPaα and CuPaβ in the usual morphology.

Fig. 8.

Species cf. bicornis Karny, 1929a (all ♀; all at the same scale); A, G, specimen ANlC IWC OB 13, right forewing, drawing and photograph; B, H, specimen ANIC IWC OB 11, left forewing, drawing and photograph; C—D, I—J, specimen ANIC IWC OB 14; C, I, left forewing, drawing and photograph; D, J, right forewing, drawing and photograph; E—F, K—L, specimen ANIC IWC OB 12; E, K, left forewing, drawing and photograph; F, L, right forewing, drawing and photograph.

Fig. 9.

Species cf. bicornis Karny, 1929a, specimen ANIC IWC OB 11 (♀), right hind wing; A, B, drawing and photograph (same scale); C, detail as located on B.

Limited variation was observed in the number of RP branches (♂, 11 forewings with 2-branched RP, 7 with 3-branched RP; ♀, 4 forewings with 2-branched RP, 8 with 3-branched RP). The organization of ScA is peculiar in that branches of the main stem originate at the wing base, resulting in a seemingly simple ScA complemented by branches of another vein / vein sector anterior to it. However, all these branches clearly belong to ScA.

As for hind wings, a fusion of M with R (instead of with RP as in pinguipes) occurs near the base in punctipennis (Figs 171, 18). Then M diverges from R but usually re-fuses with RP, near the origin of the latter, for some distance (Fig. 18B–D). This second fusion is very variable, ranging from ‘fusion absent’ (Fig. 18A) to a full fusion of M with R and RP, with a distal free part of M only (Fig. 18E–F). The length of the M + RP fusion itself varies widely (Fig. 18B–D).

Summary and comment

The comparative analysis demonstrates that the ‘classical’ STHC (Fig. 10E–H), which basically assumes a simple M (starting from the pattern represented on Fig. 10F), does not fit the data as well as the alternative one (Fig. 10I–N). In particular, the pattern represented on Fig. 10E, which is needed to complete the ‘classical’ series, has never been actually observed. In addition the alternative STHC accommodates more satisfactorily the observed variants, such as those represented on Figs 11, 12, 17B,F. In particular the wing venation pattern represented on Fig. 17B,F (and schematized on Fig. 10L) cannot be satisfactorily explained if one assumes a simple M. In order to facilitate comparison with actual material, the favored THCs are summarized on Fig. 20 (including interpretation for †perfecta Sharov, 1968; see below).

Fig. 10.

Schematic conjectures of topological homology, forewing venation, medio-cubital area, according to Béthoux & Nel's (2001) conjecture and venation nomenclature for the orthopterans [annotations in gray follow Sharov's (1968) conjecture and nomenclature for orthopterans, under which the structure indicated as * is MP, in which respect it is similar to Zeuner's (1939) and Ragge's (1955) conjectures; annotations in light gray follow Karny's (1928b) conjecture, under which the structure indicated as * is a cross-vein]; A, plesiotypic condition, as observed in †uralica Sharov, 1968 (see Fig. 1), with M branched (Sharov's ‘MA’); B, C, D, patterns observed in various gryllacrididaeans (see Fig. 12G,J, Fig. 13G,I, and Figs 17–18, respectively), to be interpreted; E–H: ‘classical’ set of conjectures according to various previous authors; I–N: alternative set of conjectures favored herein; E, hypothetical pattern, and conjecture, bridging A with F; F–H, conjectures by various authors, adapted under Béthoux & Nel's (2001) coniecture and nomenclature: F, pattern B conjectured according to Sharov (1968: fig. 27.B; see also Karny 1928b: figs 2, 9), assuming a simple M (Sharov's ‘MA’) and a 3-branched CuA + CuPaα (Sharov's ‘MP + CuA1’); G, pattern C conjectured according to Zeuner (1939: pl. 2, upper figure), assuming a simple M (Zeuner's ‘MA’) and a 2-branched CuA + CuPaα (Zeuner's ‘MP + CuA’); H, pattern D conjectured according to Ragge (1955: figs 54, 55; see also Karny 1928a: fig. 209), assuming a simple M (Ragge's ‘M’; but see Ragge 1955: 76–77) fused with R, and an isolated CuPaβ (Ragge's ‘Cu2’); I–N, conjectures endorsed herein; I, pattern observed in laudatum Johns 1997 (Fig. 3) and †perfecta Sharov 1968 (Figs 27, 28), among others, with both M and CuA + CuPaα forked; J, conjecture for pattern B, assuming a fusion of MP with the free part of CuA (* on A); K, conjecture for C, assuming a full fusion of CuA and CuPaα (i.e., CuA + CuPaα simple); L–N, conjectures for patterns observed in punctipennis Walker, 1869 (Fig. 17), all assuming a fusion of MA with R, and a basal composite stem MP + CuA + CuPa (/CuPaα + CuPaβ); L, conjecture for the condition represented on Fig. 17B,F, assuming a CuPaβ diverging distally; M, conjecture for D (and condition represented on Fig. 17A,E), assuming a full fusion of CuPaβ with CuA + CuPaα; N, conjecture for the condition represented on Fig. 17D,H, assuming a full fusion of MP with CuA + CuPaα + CuPaβ. For color version, see Plate IV.

Fig. 11.

Species pinguipes? Rentz in Morton & Rentz, 1983, specimen ANIC IWC OB 24 (♂; A–D at the same scale; E–H at the same scale); A, C, left forewing, drawing and photograph; B, D, right forewing drawing and photograph; E–H, details of medio-cubital area (see text for color-coding); E, G, interpretation and photograph, as located on C; F, H, interpretation and photograph, as located on D. For color version, see Plate V.

An oblique and comparatively strong cross-vein-like structure was observed between R and M + CuA in various specimens (* on Figs 12G, 14K, 19E). It has never been observed in Palaeozoic orthopterans and therefore is supernumerary. Its nature is unknown.

Taxonomic and nomenclatural implications

Taxonomy and nomenclature of several fossil species and taxa only remotely related to extant king crickets, raspy crickets & weta are considered first, It can be mentioned here that material yielded by the Dzaylyaucho locality [Madygen Formation, Kyrgyzstan; Ladinian/ Carnian, late Middle to early Late Triassic, according to Dobruskina (1995)] was deformed during fossilization (Rasnitsyn 1982; Sharov 1968, 1971; Figs 21, 27C,I). Based on preliminary investigations on material from this locality (in prep.), a plastic elongation of 150% best explains the observed aspect ratios. Species involved by such deformation are †abscissa Gorochov, 1987a, †triassica Sharov, 1968: 168, †perfecta, and †orientalis Sharov, 1968 (and species considered as junior synonyms of these in the following).

Fig. 12.

Species pinguipes Rentz in Morton & Rentz, 1983, specimen ANIC IWC OB 19(♀, A–D at the same scale; E–H at the same scale); A, C, left forewing, drawing and photograph; B, D, right forewing drawing and photograph; E–H, details of medio-cubital area (see text for color-coding); E, G, interpretation and photograph, as located on C; F, H, interpretation and photograph, as located on D. For color version, see Plate VI.

Fig. 13.

(Preceding page) Species pinguipes Rentz in Morton & Rentz, 1983 (A–F, M–O, ♀; G–L, P–Q, ♂; A–L at the same scale; M–Q at the same scale); A, D, M, specimen ANIC IWC OB 20, right forewing, drawing, photograph, and detail of medio-cubital area as located on D; B–C, E–F, N–O, specimen ANIC IWC OB 18; B, E, N, right forewing, drawing, photograph, and detail of medio-cubital area as located on E; C, F, O, left forewing, drawing, photograph, and detail of medio-cubital area as located on F; G, J, specimen ANIC IWC OB 23, right forewing, drawing and photograph; H–I, K–L, P–Q, specimens ANIC IWC OB 22; H, K, P, left forewing, drawing, photograph, and detail of medio-cubital area as located on P; I, L, Q, left forewing, drawing, photograph, and detail of medio-cubital area as located on Q.

Fig. 14.

(Preceding page) Species pinguipes Rentz in Morton & Rentz, 1983 (A–J at the same scale; K–L at the same scale); A–B, K, specimen ANIC IWC OB 20 (♀), left forewing, drawing, photograph, and detail of medio-cubital area as located on B; C–F, L, specimen ANIC IWC OB 17 (♂); C E, right forewing, drawing and photograph; D, F, L, left forewing, drawing, photograph, and detail of medio-cubital area; G–J, specimen ANIC IWC OB 21 (♀); G, I, right forewing; H, J, left forewing.

Fig. 15.

Species pinguipes Rentz in Morton & Rentz, 1983, specimen ANIC IWC OB 18 (♀), left hind wing; A, B, drawing and photograph (same scale); C, detail as located on B.

Fig. 16.

(Preceding page) Species undetermined, specimen ANIC IWC OB 25 (♂S; A–F at the same scale; G–H at the same scale); A, D, G, right forewing, drawing, photograph, and detail of medio-cubital area as located on D; B, E, H, left forewing, drawing, photograph, and detail of medio-cubital area as located on E; C, F, I, left hind wing drawing, photograph, and detail as located on F.

Fig. 17.

Species punctipennis Walker, 1869 (all same scale); A–B, E–F, specimen ANIC IWC OB 37 (♂); A, E, right forewing, drawing and photograph; B, F, left forewing, drawing and photograph; C, G, specimen ANIC IWC OB 36 (♀), right forewing, drawing and photograph; D, H, specimen ANIC IWC OB 38 (♂), left forewing, drawing and photograph; I–J, specimen ANIC IWC OB 37 (♂), left hind wing, drawing and photograph.

Fig. 18.

Species punctipennis Walker, 1869; A–D, F, details of the connection between RP and M, hind wing (all at the same scale; * indicates the cell delimited by R, RP, and M); A–B, specimen ANIC IWC OB 36 (♀); A, right hind wing; B, left hind wing; C–D, specimen ANIC IWC OB 37 (♂); C, left hind wing, detail as located on Fig. 17J; D, right hind wing; E–F, specimen ANIC IWC OB 38 (♂), right hind wing; E, photograph; F, detail as located on E.

Fig. 19.

Species rufovaria Kirby, 1888, specimen ANIC IWC OB 27 (♂; A–D at the same scale); A, C, right forewing, drawing and photograph; B, D, right hind wing, drawing and photograph; E, detail of the R + MA fusion, as located on C (* indicates a strong crossvein free of trachea).

Fig. 20.

Summary of topological homologies in Tagryllacris nom.-dis.-typ. nov. (compare to Fig. 1); A–B, species laudatum Johns, 1997; C–D, species †perfecta Sharov, 1968; E–H, species pinguipes Rentz in Morton & Rentz 1983; I–J, species punctipennis Walker, 1869. For color version, see Plate VII.

Fig. 21.

Species †reducta Sharov, 1968, specimen PIN 2555/1445 (holotype; Madygen Formation, Kyrgyzstan; Ladinian/Carnian, late Middle to early Late Triassic), photograph (negative imprint, light-mirrored), witnessing deformation experienced by material from the Dzaylyaucho locality.

Fig. 22.

Species †abscissa Gorochov, 1987a, specimen PIN 2069/2240 (holotype; Madygen Formation, Kyrgyzstan; Ladinian/ Carnian, late Middle to early Late Triassic), drawing (* indicate presumed intercalary veins) and photograph (left forewing, positive imprint, flipped horizontally).

Fig. 23.

Species †triassica Sharov, 1968: 168 (all the same scale; all Madygen Formation, Kyrgyzstan; Ladinian/Carnian, late Middle to early Late Triassic); A–B, specimen PIN 2069/2320 (holotype; right fore- and hind wing), drawing and photograph (positive imprint); C–D, specimen PIN 2240/4312 (holotype of primaria Sharov, 1968; right forewing), drawing and photograph (negative imprint; flipped horizontally).

Fig. 24.

Species †brodiei Handlirsch, 1938, forewings (all the same scale; except for NHM I. 6790 (K–M), Binton, Warwickshire, UK; all Lower Jurassic); A–B, presumed ♀; C–P presumed ♂; A–B, specimen NHM I.10463 (holotype; left forewing), drawing and photograph (positive imprint, flipped horizontally); C–E, specimen NHM I.6784 (right forewing), drawing and photographs (positive imprint; light mirrored, and under ethanol); F–G, specimen NHM I.10464 (right forewing), drawing and photograph (negative imprint; flipped horizontally); H–I, specimen NHM I.3383 (right forewing), drawing and photograph (positive imprint); K–M, specimen NHM I.6790 (right forewing; ‘climbers’, Warwickshire?, UK), drawing and photographs (positive imprint; light mirrored, and under ethanol); N–P, specimen NHM I.6656 (right forewing), drawing and photographs (positive imprint; dry, and under ethanol).

Fig. 25.

Species †brodiei Handlirsch, 1938 (all the same scale; all Binton, Warwickshire, UK; Lower Jurassic); A–B, presumed ♀; C–G, hind wings, presumed ♂) A–B, specimen NHM I. 10668 (right forewing), drawing and photograph (negative imprint, flipped horizontally); C–E, specimen NHM I. 10536 (left hind wing), drawing and photographs (negative imprint, light-mirrored; dry, and under ethanol); F–G, specimen NHM I. 6779 (left hind wing), drawing and photograph (positive imprint, flipped horizontally; under ethanol).

Fig. 26.

Selected cladotypes housed at the MHNG (photographs courtesy of P. Schwendinger); A—B, species carli Griffini, 1911a, holotype; A, habitus; B, detail of left forewing venation, as located on A (arrows without labels indicate the endings of CuA and CuPaα); C-D, species magnifica Brunner von Wattenwyl, 1888, holotype; C, habitus (arrow indicates the end of CuA + CuPaa); D, detail of right forewing venation, as located on C.

Fig. 27.

(Following 2 pages) Species †perfecta Sharov, 1968, presumed ♂ (all at the same scale; all Madygen Formation, Kyrgyzstan; Ladinian/Carnian, late Middle to early Late Triassic); A, G, specimen PIN 2240/4132 (holotype; left forewing), drawing and photograph (positive imprint, flipped horizontally); B, H, specimen PIN 2555/1237 (right and left forewings), drawing and photograph (positive imprint); C, I, specimen PIN 2240/4113 (right and left forewings), drawing and photograph (positive imprint, flipped horizontally); D, J, specimen PIN 2240/4159 (right forewing), drawing and photograph (negative imprint, flipped horizontally); E, K, specimen PIN 2240/4124 (right forewing), drawing and photograph (positive imprint); F, L, specimen PIN 2240/4120 (left forewing), drawing and photograph (positive imprint, flipped horizontally); G, M, specimen PIN 2240/4123 (right forewing), drawing and photograph (positive imprint).

Label

Fig. 28.

Species †perfecta Sharov, 1968, presumed ♀ (all at the same scale; all Madygen Formation, Kyrgyzstan; Ladinian/Carnian, late Middle to early Late Triassic); A, D, specimen PIN 2555/1200 (right forewing), drawing and photograph (positive imprint); B, E, specimen PIN 2240/4112 (left forewing), drawing and photograph (negative imprint); C, F, specimen PIN 2240/1865 (right forewing), drawing and photograph (positive imprint).

Fig. 29.

Species †perfecta Sharov, 1968, specimen PIN 2240/4112 (Madygen Formation, Kyrgyzstan; Ladinian/Carnian, late Middle to early Late Triassic), detail of the tarsus (photograph courtesy of A.P. Rasnitsyn); arrows and numbers indicate tarsus segmentation; A, Sharov's (1968) interpretation; B, alternative interpretation.

Fig. 30.

Hypothetical phylogenetic relationships (based on the favored conjectures, and summarizing the presumed sequence of character state acquisitions), and nomenclatural treatment (with relevant character states).

Discussion

Origin and evolution.—According to Dobruskina (1995; followed by Shcherbakov 2008), the Dzaylyaucho locality, from which the material of †perfecta and †orientalis (and ‘†orientalis-like’ species) has been unearthed, is Ladinian-Carnian in age (late Middle to early Late Triassic). Therefore the assignment of these species to the taxon Agryllacris implies that this lineage diverged ca 235 million years ago (mya) at least. This is in sharp contrast with the prevalent view (Heads & Leuzinger 2011), predicting a divergence of king crickets, raspy crickets & weta (forming a monophyletic group, or not) ca 65 mya (at best).

The new proposition has direct implications on scenarios regarding the lack of the Grylloptera stridulatory apparatus in extant king crickets, raspy crickets & weta. The earliest species showing such a stridulatory apparatus were collected from the same locality as †perfecta and †orientalis (Béthoux 2012; Gorochov, 1986; Sharov 1968, 1971). According to Béthoux (2012) the file, typical of this apparatus, was acquired once. The point is to determine whether Agryllacris belongs to Grylloptera (i.e., if Agryllacris derived from a lineage possessing the Grylloptera file), or not (if Agryllacris does not derive from a lineage possessing the Grylloptera file).

Different degrees of complexity in stridulatory apparatuses are documented in Grylloptera from Dzaylyaucho (Béthoux 2012; Gorochov 1986; Sharov 1968, 1971). In other words, by the time of existence of †perfecta and †orientalis, the Grylloptera stridulatory apparatus was undergoing its early evolution. Therefore it is unparsimonious to assume that †perfecta and †orientalis could have already gained and lost the corresponding organization. Indeed Gorochov (1995a, b) roots †perfecta and †orientalis in the ‘oedischioid wastebasket’, composed of stem-orthopterans that never gained the Grylloptera stridulatory apparatus. I concur with this view: indeed the ‘oedischioid’ †buttsi Béthoux & Beckemeyer, 2007 and †noblensis Béthoux & Beckemeyer, 2007, possessing forewings with a low number of MA and CuA + CuPaα branches, and lacking the Grylloptera file, could well form the Early Permian ‘oedischioid’ stock from which Agryllacris derived.

Particular aspects of wing venation evolution.—In forewing, species of the taxon Metagryllacris exhibit a ScA with a long posterior stem, often simple, and separated from other ScA branches by broad area filled with strong cross-veins (Figs 17, 19). This single stem cannot be mistaken with ScP because it is strongly convex (additionally the genuine ScP is evident). In contrast, in other species of the taxon Agryllacris, ScA is composed of a single main stem regularly emitting anterior branches, more or less all similar. Without proper phylogenetic scheme, the anterior area in forewing of Metagryllacris species could have been interpreted as composed of a simple ScA, and of a CP area anterior to it. The occurrence of a Costal system to which CP would belong is hypothesized by Kukalová-Peck ( 1991; and references therein) in her insect ground plan. However observations by this author have been questioned. In particular the presumably plesiomorphic occurrence of a CP area in stem-Amphiesmenoptera, observed by Kukalová-Peck & Willmann (1990), was later discarded by Willmann (1997). Elcanid orthopterans provide the only convincing support for a presumed CP area: in the most recent representatives (see Sharov 1968, 1971: fig. 14H), three veins occur anterior to RA, which could be interpreted as ScP (Sharov's ‘Sc’), ScA (Sharov's C'), and CP (not labelled by Sharov) (notice that Carpenter dealt with this case by hypothesizing a bifid ScP; Carpenter 1992: fig. 99.7). However observations in Metagryllacris match the interpretation by Sharov (1968, 1971) of the elcanid pattern, with the anterior area being filled with a ScA provided with a simple and long posterior stem, and with another anterior stem. In other words no hypothetical CP occurs. These observations suggest that the occurrence of a Costal system in insect wings might have to be reconsidered.

Fig. 31.

Cases exemplifying the concepts of atavism, pronatism, interonatism, parallelism, reversal and cryptoparallelism. For color version, see Plate VII.

The actual course and organization of tracheae within wing veins have long been recognized as a potential source of inference for establishing conjectures of topological homologies (Comstock & Needham 1898,1899).Typically, theobservation of a pair of tracheae running in a single ‘vein’ is indicative of a composite stem, such as R and MA in rufovaria, each represented by a trachea within the ‘R vein’ (Fig. 19E). However, the same material demonstrates that the corollary, namely that a single trachea is indicative a single vein, is not supported: the eminently composite stem MP + CuA + CuPa / CuPaα + CuPaβ is provided with a single trachea (Fig. 19E) in the same species. It implies that vein fusion is probably ‘completed’ by tracheal fusion at some point.

Variation, variability, and iterative evolution.—Variation in the variability of wing venation has been seldom investigated (but see Ross 2012; and references therein). The available data on Agryllacris demonstrate that the range of variation varies among the surveyed species (Tab. 1). For example, although the number of branches of CuA + CuPaα was found to be very stable in laudatum and bicomis, it varies within a limited range in pinquipes and punctipennis, and more widely in ornata. The character ‘width of the area with the free part of CuA / length of the M + CuA + CuPaα common stem’ is comparatively stable, and variable, in laudatum and pinquipes, respectively (Appendix 4).

The connection (or the lack thereof) of MP with CuA and CuA + CuPaα is also a relevant case. A connection is consistently absent in laudatum and ornata, while species of the taxon Tagryllacris usually exhibit a fusion of MP with CuA and CuA + CuPaα (as a consequence, there is not free stem of M). However, the left forewing of the specimen ANICIWC OB 24 (Fig. 11 A,C,E,G), itself belonging to the taxon Tagryllacris, exhibits the condition observed in laudatum and ornata. In addition a peculiar intermediate condition was observed in a single pinguipes specimen: the actual fusion of MP with CuA and CuA + CuPaα occurs, but MP further re-unites with MA (Fig. 12), reforming the stem of M observed in the earlier lineages. The fusion of MP with CuA and CuA + CuPaα occurs more consistently in species of the taxon Metagryllacris, without intermediate condition being observed. These examples indicate that variability possibly varies according to the phylogenetic position of a given species. Note that a similar variation in variability has been documented in the fusion of a branch of RP with RA in mantodeans (Béthoux & Wieland 2009), with occurrence of intermediate conditions in variable species.

Based on these examples, variation in character states distribution can be encompasses by three cases (Fig. 31). First, within a species exhibiting an apomorphic condition, rare variants exhibit the condition usually present in successive stem-lineages. This is the case of CuA + CuPaα being forked in pinguipes and punctipennis (Appendix 4), for example. These are atavisms, viz. the rare occurrence of a plesiomorphic condition. Although this concept usually applies to species only remotely related to the corresponding stem-lineages, it seems appropriate to use it even in species closely related to these (Fig. 31, sp. 8 in case 1): it can be argued that atavism borders on intra-specific variability indeed. The connection between atavism and reversal (or taxic atavism) is evident (Hall 2003; Stiassny 1992): both are the consequence of the persistence of functional genetic modules responsible for plesiomorphic states. The term ‘atavism’ refers to low frequency of occurrence of the plesiomorphic state in a given species (Fig. 31, sp. 10 in case 2), while reversal is characterized by a high frequency of occurrence (Fig. 31, sp. 11 in case 2).

The complementary configuration, viz. the rare occurrence of an apomorphic condition in a species usually exhibiting the plesiomorphic one, is equally plausible. Examples were documented above, such as the pectiante fusion of RP with RA, usually observed in ornata, and found to occur rarelyin laudatum (Fig. 3H,Q; and see Béthoux & Wieland 2009: fig. 13C–D). I failed to find the proper counterpart to the term ‘atavism’ for such case, therefore I propose to coin the term ‘pronatism’ (‘pro-’ = before; ‘natus’ = born). It is proposed to apply this concept only to lineages which derived directly from the corresponding stem-lineage (such as spp. 2 and 6 in case 1 on Fig. 31; see below for sp. 3; referred to as direct relative in the following). As for atavism, it is assumed that pronatism is based on a genetic organization similar to that underlying the high-frequency condition (but see below for a broader sense). And as for atavism and reversal, an evident connection exists between pronatism and parallelism (sensu Hall 2003): assuming similar genetic ground, the former is characterized by low frequency, the latter high frequency. And as for atavism, pronatism borders on intra-specific variability.

At this step confusion between parallelism on one hand, and atavism, reversal, and pronatism on the other, is possible. Hall (2003: tab. 2) defines parallelism as follows: “a feature present in closely related organisms but not present continuously in all members of the lineage” and which development is based on “normally similar developmental pathways”. However, this definition applies to atavism and reversal as well, because they are the re-expression of a genetic organization inherited from the same stem-lineage (therefore arguably similar), and of course to pronatism as defined above. I propose to give a narrower sense to parallelism, restricted to the repeated acquisition of a derived condition at high frequency, due to similar genetic ground. Atavism and reversal apply to expression of a plesiomorphic conditions.

We are left with a final case involving (1) a derived condition occurring at high frequency in a lineage, and (2) the same derived condition occurring at low frequency within a lineage which is not a direct relative of the former (Fig. 31, case 3, spp. 3 and 4 on one hand, and 7 and 11 on the other; and sp. 3 as opposed to spp. 7–11 in case 1). The translocation of CuPaα onto M + CuA fits this case: it occurs at low frequency in laudatum (Fig. 31, R), pinguipes (Fig. 14A–B, K), and in one wing of the specimen ANICIWC OB 25 (Fig. 16B,E,H), but at high frequency in †brodiei (Fig. 24), which is only remotely related to species of the taxon Agryllacris. Assuming similar genetic ground, the observed rare occurrences do not qualify for pronatism: they do not predate the expression of the corresponding derived condition at high frequency. This case better fits with parallelism as outlined above, except for the occurrence frequency. I propose to refer to the low frequency occurrences as ‘cryptoparallelisms’.

At this stage I propose to coin the term ‘iteronastism’ (‘intero-’= repeated) to encompass the concepts of atavism, reversal, cryptoparallelism, pronatism, and parallelism altogether. All have in common the repeated occurrence of a condition (either plesiomorphic or apomorphic) due to a similar underlying genetic organization. Defined on this ground, an iteronatism differs from a convergence in that the latter is due to a different underlying genetic organization (Hall 2003; and references therein).

Finally, I propose to extend the concept of iteronatism beyond repeated appearances due to similar underlying genetic organization. Given an initial pattern (i.e., the orthopteran forewing venation pattern) and a transformation type commonly observed (i.e. vein translocation), one can predict iterative appearances of the very same derived condition (i.e. CuPaα diverging from M + CuA as a consequence of a translocation) in various lineages, regardless of the adaptative significance of the transformation. In other words, provided an initial morphological context, exhaustion of morphological character states (sensu Wagner 2000), due to a limited number of transformation types, is likely to occur. A similar genetic organization is not necessarily underlying these iterative appearances, but a propensity of the condition to appear can be advocated. This broader perspective applies to all cases of iteronatism detailed above, and to the actual cases investigated herein. The concepts elaborated above are summarized as follows:

Iteronatism: repeated occurrence of a condition of any polarity, and at any species-level frequency, due to a similar underlying genetic organization (s.str.) / to a shared propensity of the condition to appear, provided the ancestral context (s.l.).

Atavism: occurrence of a plesiomorphic condition at low species-level frequency, due to the persistence and re-expression of an ancestral genetic organization; can repeat.

Reversal: occurrence of a plesiomorphic condition at high species-level frequency, due to the persistence and re-expression of an ancestral genetic organization; can repeat.

Cryptoparallelism: repeated occurrence of an apomorphic condition at low species-level frequency, due to a similar underlying genetic organization (s.str.) / to a shared propensity of the condition to appear, provided the ancestral context (s.l.).

Pronatism: appearance of an apomorphic condition at low species-- level frequency, predating its expression at high frequency in a direct relative, due to a similar underlying genetic organization (s.sfr.) / to a shared propensity of the condition to appear, provided the ancestral context (s.l.); can repeat.

Parallelism: repeated occurrence of an apomorphic condition at high species-level frequency, due to a similar underlying genetic organization (s.sir.) / a shared propensity of the condition to appear, provided the ancestral context (s.l).

Conclusion

This contribution exemplifies the importance of giving full consideration to intra-specific variation for establishing conjectures of topological homology. Indeed within-species variants can serve as Remane's (1952) intermediates and fill a gap in a transformation series otherwise incomplete. In that respect within-specimen variation taking place in symmetrical and/or serially arranged organs, such as wings, is a pinnacle. The most illuminating encountered case regards the specimen ANIC IWC OB 24 (Fig. 11), which right forewing exhibits an extremely rare configuration providing logical evidence in favor of a new STHC.

There is an unfortunate tendency of palaeoentomologists to erect new species without giving proper consideration to intra-specific variation (Béthoux 2009b). The current contribution exemplifies this issue, with six species, erected by a single author, demonstrated to be synonyms of a junior one, thanks to data on extant material. Where this trend is dramatic is that demonstrating junior synonymy is much more labor-intensive than erection of doubtful species, and can be achieved only when appropriate extant material is available (e.g., Schneider 1977, 1978), and/or on the basis of exceptional sample of fossil material (e.g., Cui et al. 2011). The routine erection of new species based on any observed difference is fundamentally flawed in that it neglects the fact that biological evolution could not take place without intra-specific variability.

A consequence of the assignment of †perfecta to Agryllacris is that this group stems in the Triassic, at least, in other words ca 170 my earlier than previously assumed. A loss of the complex gryllopteran stridulatory apparatus is no longer needed to account for the morphology of king crickets, raspy crickets and weta forewing.

Only a subset of recognized transformations was used to develop a new nomenclatural scheme (Fig. 30). The surveyed material was insufficient to allow relative timing of acquisition of the corresponding character states, and to define corresponding lineages. It is anticipated that this comparative analysis will allow wing venation homologies to be assessed in additional species, and the longest branches in the proposed transformation series to be split according to the sequence of character states acquisition.