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Geobios 48 (2015) 263–270

Available online at ScienceDirect www.sciencedirect.com

Original article

Discovery of the first lacewings (: Permithonidae) from the

§

Guadalupian of the Lode`ve Basin (Southern France)

a, b c d,

Jakub Prokop *, Fla´via Rodrigues Fernandes , Jean Lapeyrie , Andre´ Nel *

a

Charles University in Prague, Faculty of Science, Department of Zoology, Vinicˇna´ 7, CZ-128 44, Praha 2, Czech Republic

b

Museu Paraense Emı´lio Goeldi, Avenida Perimetral da Cieˆncia s/n, Campus de Pesquisa, Guama´, 66040-170 Bele´m, PA, Brazil

c

Corniche de Fontbonne, 34700 Lode`ve, France

d

Muse´um national d’histoire naturelle, institut de syste´matique, e´volution, biodiversite´, ISYEB, UMR 7205 CNRS UPMC EPHE, CP50, 45, rue Buffon, 75005

Paris, France

A R T I C L E I N F O A B S T R A C T

Article history: Lacewings and their relatives (Neuroptera) are one of the ancient holometabolous order reaching

Received 5 August 2014

spectacular diversity during the and , but with a surprisingly sparse record during the

Accepted 30 March 2015

Late Palaeozoic. Here, we describe the first members of the stem-group Permithonidae from the middle

Available online 11 April 2015

Permian () of Salagou Formation in the Lode`ve Basin (Languedoc region, Southern France):

Lodevothone pectinata nov. gen., nov. sp., and Lodevosisyra reducta nov. gen., nov. sp. Both diagnoses are

Keywords:

based on unique wing venation patterns. Discovery of these two new permithonid species in the Lode`ve

Insecta

Basin fits well with the distribution of other taxa known from the of Euramerica, Siberia, and

Neuropteroidea

Gondwana. The record of Permithonidae is documented and briefly discussed in respect of

Myrmeleontida

Permian stratigraphical range and geographical distribution. A check-list of all taxa including their synonymy is

Salagou Formation provided.

France ß 2015 Elsevier Masson SAS. All rights reserved.

1. Introduction (), bearing number of plesiomorphic characters in wing

venation and supporting the close relationship between Arche-

Neuropterida together with sister-group Coleopterida repre- and Osmylidae. The latter family encompasses mem-

sent a holometabolan insect so called Neuropteroida, well bers from the Early to extant species. It should be noticed

supported by phylogenetic analyses of morphological, molecular that this hypothesis about a relationship between these two

and phylogenomic data with history dated back at least to the late families was already proposed by Riek (1976).

Pennsylvanian (Trautwein et al., 2012; Nel et al., 2013; Misof et al., The Lode`ve Basin is situated in the department of He´rault

2014). Palaeozoic lacewings (Neuroptera) are currently classified (Languedoc region, Southern France); it is an important Permian

into two families: Permithonidae recorded from the early to late insect locality from the Southwestern Europe territory. Insect

Permian deposits of Euramerica, Siberia, and (Fig. 1), are found in the Guadalupian Salagou Formation exposed in several

and Archeosmylidae known from the late Permian and outcrops close to the town of Lode`ve (Garric, 2000; Nel et al., 2009).

(Novokshonov, 1996; Makarkin et al., 2014). However, the main The depositional palaeoenvironment is interpreted as a playa with

radiation of this order took place after the PT boundary, then ephemeral pools controlled by climate oscillations (Lopez et al.,

followed by a marked peak in diversity during the Jurassic (Jepson 2008). are commonly preserved as isolated wings or body

and Penney, 2007). Novokshonov (1996) provided a critical fragments caused probably by a long decay and post mortem

systematic revision and synonymized the Permian neuropteran transportation, or due to the activity of aquatic predators like

families Sialidopsidae, Palaeomerobiidae, and Permosisyridae with crustaceans and (Gand et al., 1997). With more than fifty

Permithonidae. Makarkin et al. (2014) confirmed the position of insect species described from the Salagou Formation assigned to

Archeosmylidae based on the study of the osmylid thirteen orders or higher rank , Lode`ve demonstrates rather

Archaeosmylidia fusca Makarkin, Yang et Ren, 2014 in Daohugou diversified insect communities with some taxa directly linked to

early and late Permian ones known from assemblages in Euramerica

and Russia (e.g., Prokop and Nel, 2011; Prokop et al., in press).

§

Corresponding editor: Gilles Escarguel.

The present contribution deals with the formal description of two

* Corresponding authors.

new neuropteran taxa of Permithonidae, including a discussion of

E-mail addresses: [email protected] (J. Prokop), [email protected] (A. Nel).

http://dx.doi.org/10.1016/j.geobios.2015.03.001

0016-6995/ß 2015 Elsevier Masson SAS. All rights reserved.

264 J. Prokop et al. / Geobios 48 (2015) 263–270

Fig. 1. (Colour online). Palaeogeographical map of the Late Permian (260 Ma) showing the position of sites with described lacewings (Permithonidae) indicated by dots (based

on Ron Blakey’s original paleomap; http://cpgeosystems.com/). 1. Elmo (Kansas, USA); 2. Kuznetsk Basin (Russia); 3. Iva-Gora, Soyana River (Russia); 4. Tchekarda, Sylva River

(Russia); 5. Lode`ve Basin (France); 6. Russky Island (Russia); 7. Kureika River (Russia); 8. Tikhiye Gory (Republic of Tatarstan); 9. Ilimpeya River (Russia); 10. Rio Grande do Sul

(Brasil); 11. Newcastle, N.S.W. (Australia); 12. Karaungir (); 13. Kargala, Orenburg reg. (Russia); 14. Mooi River, Natal (South Africa).

their systematic position and a broader evaluation of the fossil pectinate with seven straight posterior branches (plus MA); CuP

record of this family. The checklist of all taxa attributed to family simple; wing apex with intercalary folds markedly corrugated.

Permithonidae with their geographical and stratigraphical origin is

Lodevothone pectinata nov. gen., nov. sp.

given for clear arrangement.

Figs. 2, 3

Derivation of the name: Named after the pectination of vein

MA in the forewing.

2. Material and methods

Material: Holotype specimen Ld LAP 312 A-B (print and

counterprint of a forewing), Lapeyrie coll., stored in the Muse´e

The three neuropteran specimens described below come from

Fleury, Lode`ve, France.

the Guadalupian deposits of the Salagou Formation in the Lode`ve

Age and outcrop: Middle Permian, Guadalupian, Me´rifons

Basin (He´rault, Southern France), which are well known thanks to

Member, Salagou Formation; around the Salagou lake, Lode`ve

the long-term sampling effort by one of us (JL). The material

Basin, France.

(including type specimens) resulting from these collects is housed

Measurements of the holotype: Wing: 11.7 mm long, 5.8 mm

at the Muse´e Fleury (Lapeyrie collection, prefix ‘‘Ld LAP’’) in Lode`ve

wide; width of costal area: 0.9 mm; width of area between ScP and

(He´rault, France).

RA: 0.3 mm; max. width of intraradial area: 0.5 mm.

All specimens were observed under an Olympus SZX-9

Description (Fig. 2): holotype forewing elongate, more than

stereomicroscope in a dry state. Line drawings of venations were

two times as long as wide, widest beyond mid length, narrowly

obtained directly, using a coupled camera lucida. Photographs

rounded apically, with anterior margin slightly curved, especial-

were taken using a digital camera Canon D550 with a reverse lens

ly in apical quarter; no crossvein visible in main part of the wing,

MP-E 65 mm, then processed using the image-editing software

except for the anterior branches of ScP; apex of wing corrugate

Adobe Photoshop CS.

because of the presence of intercalary folds along apical margin,

We follow the systematics of Permithonidae as proposed by

especially between RP branches; costal area regularly broad, not

Novokshonov (1996). The venational symbols used here are

expanded in its mid part, slightly wider than intraradial space in

specified as follows (symbols in capitals denote the longitudinal

proximal half of wing, with numerous simple, weakly-curved ScP

veins): ScP, subcosta posterior; RA/RP, radius anterior/posterior;

branches; trichosors and nygmata not preserved, if originally

MA/MP, media anterior/posterior; CuA/CuP, cubitus anterior/

present; RA beyond junction with ScP slightly arched along and

posterior; 1A/2A, first/second anal vein.

gradually converging with wing margin, but not reaching wing

apex; intraradial space greatly widened basally, without visible

3. Systematic palaeontology crossveins; MA visible as a very short vein emerging from M,

reaching RP and re-appearing as its most proximal branch,

Order NEUROPTERA Linnaeus, 1758 distally pectinated with seven parallel posterior branches; RP

Family PERMITHONIDAE Tillyard, 1922 with seven simple parallel posterior branches (plus MA); MP

Included genera: see Table 1. divided into two main branches, anterior forked MP1 and

posterior three-branched MP2; CuA divided into two main

Genus Lodevothone nov.

branches, forked CuA1 and three-branched CuA2; CuP simple;

Derivation of the name: named after the Lode`ve Basin and

CuA separating from CuP 1.9 mm from base of preserved part of

Permithone, type genus of the Permithonidae. Gender feminine.

the wing; 1A forked and 2A pectinate with four short terminal

Type species: Lodevothone pectinata nov. gen., nov. sp.

branches.

Diagnosis: Forewing elongate; MA pectinate with seven long

Remarks: Lodevothone nov. gen. belongs to the family

posterior branches covering a very broad zone (apomorphy);

Permithonidae due to the vein RA distally straight close to wing

interadial space very broad near its base, broader than space

apex and CuP simple, unlike in Archeosmylidae. Lodevothone nov.

between ScP and RA and nearly as broad as costal area; RP

J. Prokop et al. / Geobios 48 (2015) 263–270 265

Table 1

Checklist of Palaeozoic and Mesozoic Permithonidae (Neuroptera).

Taxon Locality /

Elmothone Carpenter, 1976

E. martynovae Carpenter, 1976 Elmo, Kansas, USA /

Eopsychops Martynov, 1933

E. angaridensis Martynova, 1961 Kuznetsk Basin, Russia Cisuralian/Ufimian

E. sojanensis Martynov, 1933 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

Jurla Vilesov, 1995

J. bisubcostata Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/

Kunguromaritus Vilesov, 1995

K. guttatus Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

K. lacer Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

Lodevothone nov. gen.

L. pectinata nov. gen., nov. sp. Lode`ve Basin, France Kungurian/

Lodevosisyra nov. gen.

L. reducta nov. gen., nov. sp. Lode`ve Basin, France Kungurian/Roadian

Okolpania Vilesov, 1995

O. capitosa Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

O. eugeniae Shcherbakov et al., 2009 Russky Island, Russia Cisuralian/Kungurian

O. favorabilis Novokshonov and Novokschonova, 1997 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

O. observalis Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

Osmythone Ponomarenko and Shcherbakov, 2004

O. neoxenus Ponomarenko and Shcherbakov, 2004 Kureika River, Russia Tatarian

Palaemerobius Martynov, 1928

Palaemerobius proaviatus Martynov, 1928 Tikhiye Gory, Republic of Tatarstan Guadalupian/Kazanian

Palaemerobius avus Martynova, 1952 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

Palaemerobius latibasis Martynova, 1961 Kuznetsk Basin, Russia Cisuralian/Ufimian

Permantispa Ponomarenko and Shcherbakov, 2004

P. emelyanovi Ponomarenko and Shcherbakov, 2004 Ilimpeya River, Russia Tatarian - Lower Triassic

Permegalomus Martynov, 1931

P. maculipennis Martynov, 1933 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

= Permithonopsis triramosa Martynova, 1952 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

P. regularis Martynov, 1931 Tikhiye Gory, Republic of Tatarstan Guadalupian/Kazanian

P. (= Bianchia) spectabilis (Martynova, 1952) Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

Permerobius Martynova, 1961

P. latibasis Martynova, 1961 Kuznetsk Basin, Russia Cisuralian/Ufimian

Permipsythone Pinto and Pinto de Ornellas, 1980

Permipsythone panfilovi Pinto and Pinto de Ornellas, 1980 Rio Grande do Sul, Brasil

P. (= Permithonopsis) perantiqua (Martynova, 1961) Kuznetsk Basin, Russia Cisuralian/Ufimian

Permithone Tillyard, 1922

P. belmontensis Tillyard, 1922 Newcastle, N.S.W., Australia Tatarian

P. neonexus Riek, 1953 Newcastle, N.S.W., Australia Tatarian

P. oliarcoides Tillyard, 1926 Belmont, Australia Tatarian

P. venosa Davis, 1943 New South Wales, Australia Tatarian

Permithonopsis Martynov, 1933

P. cellulosa Martynova, 1952 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

P. dubia Vilesov and Novokshonov, 1994 Karaungir, East Kazakhstan Tatarian

P. enormis Martynova, 1952 Letopala, Soyana River, Russia Guadalupian/Kazanian

P. ivensis Martynov, 1933 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

= P. proxima Martynov, 1933 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

P. kaltanensis Martynova, 1961 Kuznetsk Basin, Russia Cisuralian/Ufimian

P. longifurcata Martynova, 1952 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

P. obscura Martynova, 1952 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

P. sharovi Martynova, 1961 Kuznetsk Basin, Russia Cisuralian/Ufimian

Permopsychops Tillyard, 1926

P. altajopanorpa Novokshonov, 1997 Kuznetsk Basin, Russia Cisuralian/Ufimian

P. belmontensis Tillyard, 1926 Newcastle, N.S.W., Australia Tatarian

P. karaungirensis Vilesov and Novokshonov, 1994 Karaungir, East Kazakhstan Tatarian

P. saurensis Vilesov and Novokshonov, 1994 Karaungir, East Kazakhstan Tatarian

Permorapisma Tillyard, 1926

P. biserialis Tillyard, 1926 Newcastle, N.S.W., Australia Tatarian

P. fragmentata Vilesov and Novokshonov, 1994 Karaungir, East Kazakhstan Tatarian

P. gori Ponomarenko and Shcherbakov, 2004 Kureika, Tunguska Basin, Russia Lopingian - Lower Triassic

P. trisetalis Tillyard, 1926 Newcastle, N.S.W., Australia Tatarian

Permosisyra Martynov, 1933

P. latipennis Martynov, 1933 Sheimo-Gora, Soyana R., Russia Guadalupian/Kazanian

P. punctata Martynov, 1933 Iva-Gora, Soyana River, Russia Guadalupian/Kazanian

P. (= Permithonopsis) paurovenosa (Martynova, 1952) Letopala, Soyana River, Russia Guadalupian/Kazanian

P. (= Sialidopsis) sojanensis (Martynova, 1952) Letopala, Soyana River, Russia Guadalupian/Kazanian

Permosmylus Tillyard, 1926

P. pincombeae Tillyard, 1926 Belmont, Australia Tatarian

Sauropsychops Novokshonov, 1996

S. (= Permithone) kazakhstanensis (Vilesov and Novokshonov, 1994) Karaungir, East Kazakhstan Tatarian

S. (= Permithone) tillyardi (Vilesov and Novokshonov, 1994) Karaungir, East Kazakhstan Tatarian

Sialidopsis Zalessky, 1926

S. kargalensis Zalessky, 1926 Kargala, Orenburg reg., Russia Tatarian

S. similis Martynov, 1937 Kargala, Orenburg reg., Russia Tatarian

266 J. Prokop et al. / Geobios 48 (2015) 263–270

Table 1 (Continued )

Taxon Locality Epoch/Stage

Sismerobius Riek, 1976

S. pusillus Riek, 1976 Mooi River, Natal, South Africa Lopingian

Sylvamarita Vilesov, 1995

S. minor Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

Sylvasenex Vilesov, 1995

S. lacrimabunda Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

Tshekardithonopsis Vilesov, 1995

T. oblivius Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

T. pictus Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

T. zalesskyi Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

Tychtobius Martynova, 1958

T. brevicostatus Martynova, 1958 Kuznetsk Basin, Russia Cisuralian/Ufimian

Uralisyra Vilesov, 1995

U. angusta Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

U. prolubnikovi Vilesov, 1995 Tchekarda, Sylva River, Russia Cisuralian/Kungurian

gen. differs from Sauropsychops Novokshonov, 1996, Sylvasenex Diagnosis: Forewing shape oval; ScP ending in costal margin;

Vilesov, 1995, Jurla Vilesov, 1995, Sylvamarita Vilesov, 1995, RA with numerous simple anterior branches; RP with three simple

Kunguromaritus Vilesov, 1995, Olkopania Vilesov, 1995, Tshekar- branches; convex MA simple with its base independent of radius;

dithinopsis Vilesov, 1995, Bianchia Martynova, 1952, Palaemerobius MA shortly coalesced to RP; CuA branched.

Martynov, 1928, Permosisyra Martynov, 1933, Permantispa Pono-

Lodevosisyra reducta nov. gen., nov. sp.

marenko and Shcherbakov, 2004, Permegalomus Martynov, 1931,

Figs. 4, 5

Permerobius Novokshonov, 1996, and Permithone Tillyard, 1922 in

Derivation of the name: Named after the reduction of cubital

the following characters: more branches of RP and MA with

veins CuA and CuP.

numerous posterior branches. The genera Uralisyra Vilesov, 1995,

Material: Holotype specimen Ld LAP 347 (forewing), Lapeyrie

and Permithonopsis Martynov, 1933 are more similar to Lode-

coll., stored in the Muse´e Fleury, Lode`ve, France.

vothone nov. gen. in having relatively numerous posterior branches

Age and outcrop: Middle Permian, Guadalupian, outcrop of

of RP, but they have less branches of MA (four instead of seven).

‘‘Les Vignasses’’, Me´rifons Member, Salagou Formation; around the

Also Permithonopsis has not its MA shortly fused with RP just distal

Salagou lake, Lode`ve Basin, France.

to its base. Elmothone Carpenter, 1976 has numerous branches of

Measurements of the holotype: wing fragment: 5 mm long,

RP similarly to Lodevothone nov.gen., but its MA is not posteriorly

2.5 mm wide, estimated length 6 mm; width of costal area

but anteriorly branched. Permosmylus Tillyard, 1926, Permopsy-

0.35 mm; width of area between ScP and RA 0.2 mm; max width of

chops Tillyard, 1926, Permithonopsis Martynov, 1933, Eopsychops

interadial area 0.3 mm.

Martynov, 1933, Tychtobius Martynova, 1958, and Permorapisma

Description: A small wing with a distally rounded shape,

Tillyard, 1926 have a wing shape distinctly triangular, and MA not

part missing, probably forewing; strongly concave ScP ending on

pectinate but with complex branching. Permipsythone Pinto and

costal margin 4.6 mm from wing apex, convex RA simple, slightly

Pinto de Ornellas, 1980 differs from Lodevothone in a completely

distally curved bearing numerous anterior branches; division of RA

different relative position of apex of ScP and a free MA.

and RP on same level as MA and MP from the wing base, concave RP

Due to and limited state of preservation of the

running parallel to RA towards wing apex with three simple

holotype of Lodevothone pectinata nov. gen., nov. sp., in the basal

branches ending close to apex; simple MA shortly coalesced to RP,

part of the wing, we can admit tiny modifications of the original

ending in wing apex; MP deeply forked, anterior branch forked

course of veins MA and MP as well as possibly missing crossveins.

again and resulting into five apical branches, posterior deeply

Any doubts resulted from the weak preservation of the holotype

forked; CuA shortly forked distally.

cannot be solved until another specimens of this species will be

Remarks: Novokshonov (1996: p. 40) proposed the synonymy

discovered.

of the Permian families Palaemerobiidae, Permosisyridae, and

Genus Lodevosisyra nov. Sialidopsidae with Permithonidae, which currently comprises

Derivation of the name: Named after the Lode`ve Basin and 25 genera (Table 1). Permithonidae differs from the Permian-

Sisyra, genus of Sisyridae. Gender feminine. Triassic family Archeosmylidae by the presence of clearly straight

Type species: Lodevosisyra reducta nov. gen., nov. sp. apex of RA and simple or terminally twigged CuP, which corresponds

to the venation of our fossil. Lodevosisyra nov. gen. differs from all

previously described permithonid genera by the following combi-

nation of venation characters: RP with three branches, MA simple,

MP with three branches, and reduced CuA and CuP. Lodevosisyra nov.

gen. is probably related to Permosisyra Martynov, 1933 based on the

reduced venations with only few branches of RP, but differs in MA

probably simple instead of having distal branches and base of MA

still clearly visible. Therefore, the new genus name introduced here

is justified by the extremely reduced pattern of venation of the new

fossil from the Lode`ve Basin.

Permosisyra has been previously placed in Sialidopsidae

(Martynova, 1962). It is currently known by the type species P.

latipennis Martynov, 1933, plus two more dubious taxa: ?P.

Fig. 2. Forewing venation of Lodevothone pectinata nov. gen., nov. sp., holotype punctata Martynov, 1933, and ?P. paurovenosa (Martynova, 1952)

specimen, N8 Ld LAP 312 A-B, line drawing. Scale bar: 2 mm. (= Sialidopsis sojanensis Martynova, 1952), both from the late

J. Prokop et al. / Geobios 48 (2015) 263–270 267

Fig. 3. (Colour online). Forewing venation of Lodevothone pectinata nov. gen., nov. sp., holotype specimen, N8 Ld LAP 312 A-B, photographs of print and counterprint. Scale bar:

2 mm.

Permian (early Kazanian) of the Republic of Tatarstan, Russia oldest family record to date (Kukalova´, 1964: 40; JP, pers. obs.).

(Novokshonov, 1996). Therefore, it seems reasonable to postulate an origin of the order

Neuroptera at least in the late , as indicated by the

calibrated phylogeny proposed by Misof et al. (2014).

4. Remarks on palaeobiogeography and palaeobiodiversity

The of the Permithonidae and included genera is

still not demonstrated due to the lack of phylogenetic analysis of

Current record of the 25 genera assigned to Permithonidae

these Permian Neuroptera. Even the exact phylogenetic relation-

corresponds to a broad, if not cosmopolitan distribution encom-

ships of the Permithonidae with other, younger neuropteran

passing data from Euramerica, Siberia, and Gondwana (Fig. 1; Table

groups remain obscure. Nevertheless it seems that two phases of

1). However, the majority of these fossils actually comes from the

radiation during the Permian period occurred, distinguishable in

Russian localities dominated by Tchekarda (Syva River, Ural) and

the time chart of plotted genera (Fig. 6): a first one during the late

others, an obvious byproduct of the large sampling of these sites.

Number of fossils still remains undescribed in the private and

institutional collections, as for instance specimens from Obora in

the Boskovice Basin (Czech Republic), actually representing the

Fig. 5. (Colour online). Forewing venation of Lodevosisyra reducta nov. gen., nov. sp.,

holotype specimen, N8 Ld LAP 347, photograph of print and counterprint. Scale bar:

Fig. 4. Forewing venation of Lodevosisyra reducta nov. gen., nov. sp., holotype

2 mm. specimen, N8 Ld LAP 347, line drawing. Scale bar: 1 mm.

268 J. Prokop et al. / Geobios 48 (2015) 263–270

Fig. 6. Stratigraphic range of genera currently assigned to Permithonidae (Neuroptera).

Cisuralian (Kungurian), and a second one starting close to the (Chumakov and Zharkov, 2002, 2003). Only two genera, namely

Guadalupian/Lopingian boundary. Thus, a rapid Permian radiation Permantisma and Permorapisma, reached and possibly survived the

of permithonids was possibly affected by marked climatic oscilla- Permian/Triassic boundary.

tions like two prominent warming periods in the late Sakhmarian However, the record of all the genera attributed to Permitho-

and the end of the Lopigian; nevertheless the fossil record may also nidae is based entirely on isolated wings (Fig. 7), which determines

remain simply rather patchy during the middle-late Guadalupian the limits for and closer comparisons with lacewing taxa

Fig. 7. (Colour online). Photographs of selected permithonid wing venation: A. Elmothone martynovae Carpenter, 1976, holotype, MCZ 5585, Elmo, Kansas, USA.

B. Palaemerobius latibasis Martynova, 1961, holotype, PIN 600-471, Kaltan, Russia. C. Permegalomus maculipennis Martynov, 1933, holotype, PIN 2334-68, Sojana-Iva gora,

Russia. D. Permomegalopus spectabilis (Martynova, 1952), holotype, PIN 117-2513, Sojana-Iva gora, Russia. E. Permithonopsis ivensis Martynov, 1933, holotype, PIN 2455-37,

Sojana-Iva gora, Russia. F. Permipsythone perantiqua (Martynova, 1961), holotype, PIN 1197-419, Kaltan – Kuzbas, Russia. G. Sialidopsis similis Martynov, 1937, hindwing,

holotype, PIN 100-11, Kargala, Russia. H. Permosisyra punctata Martynov, 1933, holotype, PIN 2455-9, Sojana-Iva gora, Russia. I. Tychtobius brevicostatus Martynova, 1958,

holotype, PIN 506-119, Kuzbas, Russia. Scale bars: 2 mm.

J. Prokop et al. / Geobios 48 (2015) 263–270 269

Fig. 8. (Colour online). Hindwing venation of Permithonidae gen. et sp. indet, specimen No. Ld LAP 507, photograph of print. Scale bar: 2 mm.

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reducta nov. gen., nov. sp., are the first described neuropterans d’Autun 161, 21–38.

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from the middle Permian (Guadalupian) of the Lode`ve Basin

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(southern France). In addition, the fragmentary specimen of

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another neuropteran wing (N8 Ld LAP 507: print of an anterior tions for the fauna of the Wealden. Palaeogeography, Palaeoclima-

part of a wing; Fig. 8) was discovered. The presence of a nearly tology, Palaeoecology 248, 109–118.

Kukalova´, J., 1964. Permian insects of Moravia. Part 2. Liomopteridae. Sbornı´k

straight RA in wing apex supports the attribution to the family

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Permithonidae, but the precise generic placement cannot be

Lopez, M., Gand, G., Garric, J., Ko¨rner, F., Schneider, J., 2008. The playa environments

established. Therefore, we consider this specimen as a Permitho- of the Lode`ve Permian basin (Languedoc-France). Journal of Iberian Geology 34,

29–56.

nidae gen. et sp. indet.

Makarkin, V.V., Yang, Q., Ren, D., 2014. A new basal osmylid neuropteran insect from

On the basis of the wing venation, these two newly described

the Middle Jurassic of China linking Osmylidae to the Permian-Triassic Arche-

permithonid species can be compared to other taxa previously osmylidae. Acta Palaeontologica Polonica 59, 209–214.

Martynov, A.V., 1928. Permian fossil insects of North-East Europe. Geologicheskij

known from early to late Permian and early Triassic deposits,

Muzej Akademii nauk SSSR [Travaux du Muse´e Ge´ologique pre`s l’Acade´mie des

supporting broad faunal exchanges across Euramerica at this time.

Sciences de l’URSS] Trudy 4, 1–118.

Nevertheless, besides wings the common lack of other body Martynov, A.V., 1931. [New fossil insects from Tikhie Gory. 2. (excluding

Miomoptera)]. Akademia nauk SSSR, Paleontologicheskii Institut. Trudy 8,

structures remains an important limitation for further assessment

149–212.

of diversity and broader picture of the morphological disparity of

Martynov, A.V., 1933. Permian fossil insects from the Arkhangelsk district. Part

these tiny insects. 2. Neuroptera, and Coleoptera, with the description of two new

from Tikhiye Gory. Akademia nauk SSSR. Paleontologicheskii Institut,

Trudy 2, 63–96.

Acknowledgements

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Authors are indebted to Vladimir N. Makarkin (Institute of

mia nauk SSSR, Paleontologicheskii Institut. Trudy 40, 197–237 (in Russian).

Biology and Soil Sciences, Far Eastern Branch of the Russian Academy

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sincerely thank the curators of the Muse´e Fleury in Lode`ve for the

komye kuznetskogo bassejna [Palaeozoic insects of the Kuznetsk basin], 85Aka-

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at Harvard University, Cambridge, USA) who allowed studying and

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