- Email: [email protected]
Saxonagrion minutus nov. gen. et sp., the oldest damselfly from the Upper Permian of France (Odonatoptera, Panodonata, Saxonagrionidae nov. fam.)
S A X O N A G R I O N M I N U T U S NOV. GEN. ET SP., THE OLDEST DAMSELFLY FROM THE U P P E R P E R M I A N OF FRANCE (ODONATOPTERA, PANODONATA, SAXONAGRIONIDAE NOV. FAM.) ANDRE NEL, GEORGESGAND, GUNTHERFLECK, OLMER BETHOUX, JEAN LAPEYRIE & JACQUES GARRIC NEL A., GAND G., FLECK G., BETHOUX O., LAPEYRIE J. & GARIC J. 1999. Saxonagrion minutus nov. gen. et sp., the oldest damselfly from the Upper Permian of France (Odonatoptera, Panodonata, Saxonagrionidae nov. faro.). [Saxonagrion minutus nov. gen. et sp., la plus ancienne libellule du Permien sup6rieur fran~ais (Odonatoptera, Panodonata, Saxonagrionidae nov. fam.)l. GEOBIOS, 32, 6: 883-888. Villeurbanne, le 31.12.1999. Manuscrit d6pos6 le 22.10.1998; accept6 d6finitivement le 25.01.1999. ABSTRACT - Saxonagrion minutus nov. gen. et sp. was found in the Saxonian (Salagou Formation) of the Lod6ve basin. It is the oldest and the first record of the modern infra-order Panodonata in the Palaeozoic (Upper Permian of France). The present discovery supports the hypothesis concerning the persistance of many groups of Odonatoptera through the Permo-Triassic boundary. KEYWORDS: INSECTA, ODONATOPTERA, SAXONAGR1ONIDAE NOV. FAM., GEN ET SP. NOV., OLDEST RECORD, PERMO-TRIASSIC BOUNDARY CRISIS. RI~SUMI~ - Saxonagrion minutus nov. gen. et sp. a 6t6 trouv6 dans le Saxonien (Formation du Salagou) du bassin de Lod~ve. C'est le premier et le plus ancien fossile se rattachant h l'infra-ordre moderne des Panodonata, dans le Pal6ozoique (Permien sup6rieur, France). Cette d6couverte conforte l'hypoth6se selon laquelle de nombreux groupes d'Odonatoptera ont surv6cu h la crise permo-triasique. MOTS-CLt~S: INSECTA, ODONATOPTERA,SAXONAGRIONIDAE NOV. FAM., GEN ET SP. NOV., PLUS ANCIEN FOSSILE, CRISE PERMO-TRIASIQUE.
INTRODUCTION
SYSTEMATIC P2d_.4~ ONTOLOGY
The discovery of fossil arthropods in the Lod6ve basin has been recorded recently by Gand et al. 1997. Insects are highly diverse with several new taxa (Nel et al. 1999a,b). Their study is going on with the present description of a new Odonatoptera: Saxonagrion minutus nov. gen. and sp.
In the following study we use the wing venation nomenclature of Riek (1976) and Riek & Kukalove-Peck (1984), e m e n d e d by Kukalov~-Peck (1991), Nel et al. (1993) and Bechly (1995, 1996), unlike t h a t of Kuka]ov~-Peck (submitted). We accept the phylogenetic classification of Anisopt era proposed by Bechly (1996), emended by Bechly (1997).
This insect remain has been found close to Salasc in the site F23 n a m ed "les Canals" of the Salagou Formation from the Saxonian Group (Odin 1986) (Figs 1, 2). Geographical and stratigraphical informations on the locality are available in this last work (Gand et al. 1997, fig 1, 2, 892-893). The channel silstone body containing Saxonagrion has been dated as Lower Kazanian. It is based on new taxa of Protozygoptera very similar to Epilestes kargalensis MARTYNOV~1937 and Permepallage angustissima MARTYNOV~1938 known from the base of this stage in the Russian Permian (Nel et al. 1999a; Gand et al. 1997).
ODONATOPTERA(sensu Bechly 1996) PANODONATA Bechly, 1996 ZYGOPTERASelys, 1853 ? SAXONAGRIONIDAE nov. faro. Type genus - Saxonagrion nov.gen.
D i a g n o s i s - Within the Perm i an fauna, this family is characterized by its cubito-anal region, sharing with modern Odonata: the free basal part of CuA is a short vein perpendicular to MP and AA, exactly opposite the point of fusion between MAb
884 STRATIGRAPHY F
Marks
I
AGES
Log
Gand et al. 1997
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~
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~:Z
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FIGURE 1 - A. T h e Lod6ve b a s i n in F r a n c e . B. Location of t h e F23 Saxonagrion minutus site in t h e Lod6ve basin. Situation du gisement F23 h S a x o n a g r i o n m i n u t u s .
450
< (,9
and MP; no strong dorsal sclerotization of the aligned veins MAb and CuA (unique plesiomorphy); no concave posterior branch of MP; subnodus and Cr aligned and distinctly oblique; the branches of RP are all well distal of the nodus; only one row of cells in all the areas between the main veins.
400 Z Z<
N
G e n u s S a x o n a g r i o n nov. gen. T y p e s p e c i e s - Saxonagrion m i n u t u s sp. nov. E t y m o l o g y - A f t e r A g r i o n a n d t h e S a x o n i a n G r o u p of t h e Lod6ve basin.
220
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D i a g n o s i s - T h a t of t h e family.
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Saxonagrion minutus nov. gen. nov. sp. F i g s 3, 4 F4 H o l o t y p e - S p e c i m e n Ld L A P 173 A-B, coll. Lapeyrie, M u s 6 e Fleury, Lod6ve. P a r t a n d c o u n t e r p a r t of t h e b a s a l two t h i r d of a wing, w i t h o u t a n y t r a c e of coloration. H o r i z o n a n d L o c a l i t y - U p p e r P e r m i a n , fossiliferous site F23 located a t "les C a n a l s " to t h e s o u t h of Lod6ve. It is a c h a n n e l s i l s t o n e b o d y h a v i n g also a lot of t r i o p s i d s a n d Conchostracans. F2
D e s c r i p t i o n - W i n g a b o u t 19 m m long a n d 4 mm wide; distances from the base to the arculus, 1.9 mm, to the nodus, 6.6 mm, and to the pterostigma, 7.5 mm. Nodus in a rather basal position. Pterostigma poorly preserved, 1.5 mm long, apparently not braced. Wing well petiolated, petiole 1.6 mm 10ng and about 1.1 mm wide. A distinct vein AA basally fused with AP in the petiole and distally
-100m
F1
-0 o" 0.02 ~o O0., n . O,'t~.O
FIGURS 2 - Location of t h e F23 Saxonagrion m i n u t u s site in t h e P e r m i a n series; Leonardian ~ Kungurian from Ross & Ross (1994). Situation du gisement & Saxonagrion m i n u t u s dans la
sgrie permienne; le L#onardien est approximativement gquivalent au Koungourien d'apr~s Ross & Ross (1994).
885
FIGURE 3
-
Saxonagrion rninutus
nov. gen. et sp., holotypus, Fleury Museum of Lod~ve (H~rault); Saxonian Group, Salagou Formation, Upper Permian; fig. 3.1 = counterpart n~ LdLAP173B; fig. 3.2 = part n~ LdLAP173A;fig. 3.3 = the wing venation with abbreviations corresponding to the names of the veins; specimens kept in the Fleury Museum of Lod~ve.Saxonagrionminutus nov. gen. et sp. ; holotype conservd au Musde Fleury de Lod~ve (Hdrault); Groupe Saxonien, Formation du Salagou, Permien supdrieu~ Les fig. 3.1 et 3.2 sont successivement la contre-empreinte et l'empreinte de l'aile, La fig. 3.3 reprdsente les nervures nommdes par leur abrdviations
RAq N"
3
MAa
fused with CuA. The organization of AA and CuA is identical to t h a t of a modern Zygoptera. CuP not preserved but it was only present as a CuP crossing (transverse vein) between M + Cu and AA, without any distal free part. CuA separated from MP exactly opposite posterior branch MAb of MA. Free part of CuA very short, 0.2 mm long before its fusion with AA. Distally, CuA well-defined, parallel with MP and weakly zigzagged. MP with no concave posterior branch. RP separating from MA midway between RA and angle of MA in arculus. Angle between RA and RP + MA in arculus very open. MAb (sensu Nel et al. 1993) very strong. Discoidal space basally open. A strong and oblique vein Cr between ScP and RA in nodus and a oblique subnodus Sn well aligned with Cr. No preserved antenodal crossveins but only the two primary antenodal crossveins were probably present. No crossvein in area between RA and RP and only one crossvein in area between RP and MA, basal of subnodus. Three preserved postnodal crossveins but they were probably six in the postnodal area. Base of IR2 is 1.5 mm (two cells) distal of subno-
dus, that of RP2 4.6 mm (five cells) distal of subnodus and base of IR1 is 4.6 mm (five cells) distal of base of RP2. Only one row of cells between all main veins. No oblique crossvein "O" between IR2 and RP2.
PHYLOGENETIC AND BIOSTRATIGRAPHIC CONSIDERATIONS PHYLOGENY r n i n u t u s nov. gen. et sp. has several synapomorphies of the Panodonata justifying its present attribution to this "modern" group: nodal Cr and Sn present, aligned and oblique (synapomorphies of the Nodialata); a sclerotized pterostigma between RA and C (synapomorphy of the Stigmoptera); MA with a strong posterior branch MAb (synapomorphy of the Discoidalia); CuA separating from MP opposite the contact of MAb with MP; free part of CuA very short before its fusion with AA; arculus, MAb and free part of CuA nearly aligned; arculus making a very open angle Saxonagrion
886
2
3
4
F~GURE 4 - Saxonagrion minutus nov. gem et sp.; fig. 4.1-2 = counterpart n ~ LdLAP173B and fig. 4.3-4 = part n ~ LdLAP173A lighted differently to show veins and cells. The fig. 4.4 is an enlargment of the posterior p a r t of the wing seen from the fig. 4.3. Saxonagrion minutus nov. gen. et sp.; empreinte (Fig. 4.3, 4.4) et contre empreinte (Fig. 4.1, 4.2) gclair~es de mani~re diffgrente afin de souligner les cellules.
887 with RA (synapomorphies of the Panodonata). After the original description of the U p p e r Permian Protozygoptera Permagrion falklandicus proposed by Tillyard (1928), the organization of the veins AA, CuA and MP would be very similar to that of Saxonagrion minutus nov. gen. et sp. Nevertheless, we reexamined the original material of Permagrion (specimen n ~ Ar 2168, collection Naturh. Riksmuseum Paleozoo. Adv. Stockholm) (Bethoux 1998, Bethoux & Nel, in prep.). The interpretation of Tillyard is erroneous and the wing venation of Permagrion is completely different to that of Saxonagrion in the following points: CuA separated from MP well before MAb, presence of a strong posterior branch of MP. Permagrion is closely related to the Permolestidae sensu Bechly (1996). The character "MP with no strong posterior branch" is a plesiomorphic condition present in Panodonata, but not in Protozygoptera Permolestidae and Permagrionidae. Saxonagrion has no strict panodonatan "discal brace" sensu Carle (1982), i.e. no strong dorsal sclerotization of the aligned veins MAb and CuA. This is a plesiomorphic condition suggesting that Saxonagrion has a basal position within the Panodonata.
Saxonagrion gen. nov. shares with the Zygoptera the long petiolated wing, not present in the most basal group of the Panodonata, i.e. the Tarsophlebioptera (Upper Jurassic family Tarsophlebiidae), sister group of the "true" Odonata. Nevertheless, this character is highly homoplasic and not sufficient for its attribution to the Zygoptera. Furthermore, Saxonagrion has no known autapomorphy in its wing venation. The exact affinities of Saxonagrionidae fam. nov. within Panodonata remain uncertain. BIOSTRATIGRAPHY The Odonatoptera were highly diverse in the Upper Permian and had already a long history since the Middle Carboniferous. Upper Permian and Lower to Middle Triassic Odonatoptera belong to numerous suborders, Protanisoptera, Triadophlebiomorpha and Protozygoptera (Bechty 1996). A true Odonatoptera Discoidalia has been recently discovered in the Westphalian of England (Jarzembowski & Nel in prep.), but the previously oldest known Panodonata (Tarsophlebioptera + Zygoptera + Epiproctophora) are from the Middle and Upper Triassic (Nel et al. 1993); Protozygoptera [Permagrionoidea + Archizygoptera (= Kennedyomorpha sensu Pritykina, 1981) sensu Bechly 1996)] being the sister group of the Panodonata (Bechly 1996). From this, it appears clearly that the present fossil wing from the Upper Permian of the Lod6ve basin is the oldest record of the Panodonata.
CONCLUSION The present discovery of a putative Panodonata in the Upper Permian supports the hypothesis of Nel et al. (1999a,b) concerning the persistance of many groups of Odonatoptera through the PermoTriassic boundary and the greater antiquity of more advanced groups than was previously supposed. Nevertheless, as the Protozygoptera and Panodonata are considered as sister-groups (Bechly 1996), the presence of the former in the Lower Permian implies that of the latter during the same period. Thus, the" discovery of this new family Saxonagrionidae is not so surprising than what could be supposed. Nevertheless, both the phylogenetic analysis and the present find contradict the results of Labandeira & Seposki (1993) and Labandeira (1994), strictly based on the fossil record, indicating that the Panodonata were absent in the Upper Permian. It confirms that phylogenetic analyses are necessary in order to successfully study the historical variations of biodiversity. A c k n o w l e g e m e n t s - This paper is a contribution to the project "Pal~ontologie et Biostratigraphie du Permien" of the UMR 5561 of CNRS: "Biog~osciences" and to the 'UPRES' project "Syst~matique et crises de la biodiversit~ chez les insectes" in the Laboratoire d'Entomologie du Museum National d'Histoire Naturelle, Paris. This work was supported by the UMR 5561 of the CNRS.
REFERENCES BECHLY G. 1995 - Morphologische Untersuchungen am Flfigelgeader der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Beriicksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, BSblingen, sp. vol. 1, 341 p. BECHLY G. 1996 - Morphologische Untersuchungen am Fliigelge~ider der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Ber~icksichtigung der Phytogenetischen Systematik und des Grundplanes der Odonata. Petalura, BSblingen, sp. vol. 2, 402 p. (revised edition of the previous publication, with an english appendix including a new phylogenetic systern of fossil and Recent odonates). BECHLYG. 1997 - Phylogenetic Systematics of Odonata. - homepage on the Internet: http://members.aol.com/odonatadat/ phylogeny/bechl3;htm. BETHOUX O. 1998 - Les Protozygoptera (Odonatoptera) permotriaslques. Essai d'analyse phylog~n6tique. Impact du choix du groupe externe sur les phylogenies. M~moire de DiplSme d'Etudes Approfondies, Syst~matique animale et v~g~tale, Musdum National d'Histoire naturelle de Paris [in6dit]. CARLE F.L. 1982 - The wing vein homologies and phylogeny of the Odonata: A continuing debate. Societas Internationalis Odonatologica, Rapid Communications, 4:10 + 66 p. GAND G,. LAPEYRIEJ., GARRICJ., NEL A., SCHNEIDERJ. & WALTER H. 1997 - D~couverte d'Arthropodes et de Bivalves in6dits dans le Permien continental (Lod~vois, France). Comptes
Rendus de l'Acaddmie des Sciences de Paris, Sciences de la terre et des plan~tes, 325: 891-898. KUKALOvA-PECKJ. 1991 - Chapter 6: Fossil history and the evolution of Hexapod structures. In NAUMANN I.D. (ed.), The
888 insects of Australia, A textbook for students and research workers (2nd ed.), 1. Melbourne University Press: 141-179. LABANDEmA C.C. 1994 - A compendium of fossil insect. Milwaukee Public Museum Contributions in Biology and Geology, 88, 71 p. LABANDEmAC.C. & SEPOSKIJ.J.Jr. 1993 - Insect diversity in the fossil record. Science, 261: 310-314. MARTYNOVA.B. 1938 - l~tude sur l'histoire g~ologique et de phylog~nie des ordres des insectes (Pterygota). Premiere partie. Palaeoptera et Neoptera-Polyneoptera. Travaux de
fossils from Argentina (Insecta: Odonatoidea) and basic characters states in pterygote wings. Canadian Journal of Zoology, 62: 1150-1166. Ross C.A & Ross J.R.R 1994 - Permian sequence stratigraphy and fossil zonation. In BEAUCHAMPB., EMBRYA. & GLASS D. (eds), Pangea: Global Environments and Resources, Canadian Society Petroleum Geology, 17: 219-231. TILLYARDR.J. 1928 - A Permian fossil damselfly wing from the Falkland islands. Transactions of the Royal Entomological Society of London, 76: 55-63.
l'Institut Pal6ontologique, Acaddmie des Sciences de I'URSS, 7, 149 p. [in Russian, with a French summary]. NEL A., GAND G. & GARRICJ. 1999a - A new family of Odonatoptera from the continental Upper Permian: the Lapeyriidae (Lod~ve Basin, France). Geobios, 32, 1: 63-72. NEL A., GANDG., GARRICJ. & LAPEYRIEJ. 1999b - The first recorded protozygopteran insects from the Upper Permian of France. Palaeontology, 42, 1: 83-97. NEL A., MARTINEZ-DELCLOSX., PAICHELEa J.-C. & HENROTaY M. 1993 - Les "Anisozygoptera" fossiles. Phylog6nie et classification. (Odonata). Martinia, H.S., 3,311 p. ODIN B. 1986 - Les formations permiennes, Autunien sup~rieur Thuringien, du "bassin" de Lod~ve (H~rault, France): stratigraphie, min~ralogie, pal6oenvironnement, corr61ations. University d'Aix-Marseille, 375 p. [in6dit]. PRnS~dNA L.N. 1981 - New Triassic Odonata of middle Asia. In VISHNIAKOVAV.N., DLUSSKYG.M. & PRITYKINAL.N. (eds), New insects from the territory of the U.S.S.R. Trudy Paleontologiceskogo Instituta Akademie S.S.S.R., 183:5-42 [in Russian]. RIEK E.F. 1976 - A new collection of insects from the Upper Triassic of south Africa. Annals of the Natal Museum, 22 (3): 791-820. RIEK E.F. & KUKALOVA-PECKJ. 1984 - A new interpretation of dragonfly wing venation based upon early Carboniferous
A. NEL, G. F L E C K & O. B E T H O U X Laboratoire d'Entomologie Mus6um National d'Histoire Naturelle 45 rue de Buffon F-75005 Paris e-mail: [email protected]. G. GAND UMR 5561 du CNRS, Universit~ de Bourgogne Centre des Sciences de la Terre 6, boulevard Gabriel F-21000 Dijon e-mail: umr [email protected] ou [email protected]. J. L A P E Y R I E Corniche de Fontbonne F-34700 Lod~ve J. GARRIC 16 rue des Azal~es, La Chamberte F-34070 Montpellier
INTRODUCTION
SYSTEMATIC P2d_.4~ ONTOLOGY
The discovery of fossil arthropods in the Lod6ve basin has been recorded recently by Gand et al. 1997. Insects are highly diverse with several new taxa (Nel et al. 1999a,b). Their study is going on with the present description of a new Odonatoptera: Saxonagrion minutus nov. gen. and sp.
In the following study we use the wing venation nomenclature of Riek (1976) and Riek & Kukalove-Peck (1984), e m e n d e d by Kukalov~-Peck (1991), Nel et al. (1993) and Bechly (1995, 1996), unlike t h a t of Kuka]ov~-Peck (submitted). We accept the phylogenetic classification of Anisopt era proposed by Bechly (1996), emended by Bechly (1997).
This insect remain has been found close to Salasc in the site F23 n a m ed "les Canals" of the Salagou Formation from the Saxonian Group (Odin 1986) (Figs 1, 2). Geographical and stratigraphical informations on the locality are available in this last work (Gand et al. 1997, fig 1, 2, 892-893). The channel silstone body containing Saxonagrion has been dated as Lower Kazanian. It is based on new taxa of Protozygoptera very similar to Epilestes kargalensis MARTYNOV~1937 and Permepallage angustissima MARTYNOV~1938 known from the base of this stage in the Russian Permian (Nel et al. 1999a; Gand et al. 1997).
ODONATOPTERA(sensu Bechly 1996) PANODONATA Bechly, 1996 ZYGOPTERASelys, 1853 ? SAXONAGRIONIDAE nov. faro. Type genus - Saxonagrion nov.gen.
D i a g n o s i s - Within the Perm i an fauna, this family is characterized by its cubito-anal region, sharing with modern Odonata: the free basal part of CuA is a short vein perpendicular to MP and AA, exactly opposite the point of fusion between MAb
884 STRATIGRAPHY F
Marks
I
AGES
Log
Gand et al. 1997
.',"':
~
I000
"---Z
~-
.....
,:s
---
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~(
800 ?
Z ~
%-2
__.
~:Z
k-C-ZCanals ~
o F23
1----!
"~
~/
---[
soo ~
Z <
)~
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FIGURE 1 - A. T h e Lod6ve b a s i n in F r a n c e . B. Location of t h e F23 Saxonagrion minutus site in t h e Lod6ve basin. Situation du gisement F23 h S a x o n a g r i o n m i n u t u s .
450
< (,9
and MP; no strong dorsal sclerotization of the aligned veins MAb and CuA (unique plesiomorphy); no concave posterior branch of MP; subnodus and Cr aligned and distinctly oblique; the branches of RP are all well distal of the nodus; only one row of cells in all the areas between the main veins.
400 Z Z<
N
G e n u s S a x o n a g r i o n nov. gen. T y p e s p e c i e s - Saxonagrion m i n u t u s sp. nov. E t y m o l o g y - A f t e r A g r i o n a n d t h e S a x o n i a n G r o u p of t h e Lod6ve basin.
220
rr'(.9
,,o~ u
._--._-J
tl
F4
loo
D i a g n o s i s - T h a t of t h e family.
~r~#, l,
Saxonagrion minutus nov. gen. nov. sp. F i g s 3, 4 F4 H o l o t y p e - S p e c i m e n Ld L A P 173 A-B, coll. Lapeyrie, M u s 6 e Fleury, Lod6ve. P a r t a n d c o u n t e r p a r t of t h e b a s a l two t h i r d of a wing, w i t h o u t a n y t r a c e of coloration. H o r i z o n a n d L o c a l i t y - U p p e r P e r m i a n , fossiliferous site F23 located a t "les C a n a l s " to t h e s o u t h of Lod6ve. It is a c h a n n e l s i l s t o n e b o d y h a v i n g also a lot of t r i o p s i d s a n d Conchostracans. F2
D e s c r i p t i o n - W i n g a b o u t 19 m m long a n d 4 mm wide; distances from the base to the arculus, 1.9 mm, to the nodus, 6.6 mm, and to the pterostigma, 7.5 mm. Nodus in a rather basal position. Pterostigma poorly preserved, 1.5 mm long, apparently not braced. Wing well petiolated, petiole 1.6 mm 10ng and about 1.1 mm wide. A distinct vein AA basally fused with AP in the petiole and distally
-100m
F1
-0 o" 0.02 ~o O0., n . O,'t~.O
FIGURS 2 - Location of t h e F23 Saxonagrion m i n u t u s site in t h e P e r m i a n series; Leonardian ~ Kungurian from Ross & Ross (1994). Situation du gisement & Saxonagrion m i n u t u s dans la
sgrie permienne; le L#onardien est approximativement gquivalent au Koungourien d'apr~s Ross & Ross (1994).
885
FIGURE 3
-
Saxonagrion rninutus
nov. gen. et sp., holotypus, Fleury Museum of Lod~ve (H~rault); Saxonian Group, Salagou Formation, Upper Permian; fig. 3.1 = counterpart n~ LdLAP173B; fig. 3.2 = part n~ LdLAP173A;fig. 3.3 = the wing venation with abbreviations corresponding to the names of the veins; specimens kept in the Fleury Museum of Lod~ve.Saxonagrionminutus nov. gen. et sp. ; holotype conservd au Musde Fleury de Lod~ve (Hdrault); Groupe Saxonien, Formation du Salagou, Permien supdrieu~ Les fig. 3.1 et 3.2 sont successivement la contre-empreinte et l'empreinte de l'aile, La fig. 3.3 reprdsente les nervures nommdes par leur abrdviations
RAq N"
3
MAa
fused with CuA. The organization of AA and CuA is identical to t h a t of a modern Zygoptera. CuP not preserved but it was only present as a CuP crossing (transverse vein) between M + Cu and AA, without any distal free part. CuA separated from MP exactly opposite posterior branch MAb of MA. Free part of CuA very short, 0.2 mm long before its fusion with AA. Distally, CuA well-defined, parallel with MP and weakly zigzagged. MP with no concave posterior branch. RP separating from MA midway between RA and angle of MA in arculus. Angle between RA and RP + MA in arculus very open. MAb (sensu Nel et al. 1993) very strong. Discoidal space basally open. A strong and oblique vein Cr between ScP and RA in nodus and a oblique subnodus Sn well aligned with Cr. No preserved antenodal crossveins but only the two primary antenodal crossveins were probably present. No crossvein in area between RA and RP and only one crossvein in area between RP and MA, basal of subnodus. Three preserved postnodal crossveins but they were probably six in the postnodal area. Base of IR2 is 1.5 mm (two cells) distal of subno-
dus, that of RP2 4.6 mm (five cells) distal of subnodus and base of IR1 is 4.6 mm (five cells) distal of base of RP2. Only one row of cells between all main veins. No oblique crossvein "O" between IR2 and RP2.
PHYLOGENETIC AND BIOSTRATIGRAPHIC CONSIDERATIONS PHYLOGENY r n i n u t u s nov. gen. et sp. has several synapomorphies of the Panodonata justifying its present attribution to this "modern" group: nodal Cr and Sn present, aligned and oblique (synapomorphies of the Nodialata); a sclerotized pterostigma between RA and C (synapomorphy of the Stigmoptera); MA with a strong posterior branch MAb (synapomorphy of the Discoidalia); CuA separating from MP opposite the contact of MAb with MP; free part of CuA very short before its fusion with AA; arculus, MAb and free part of CuA nearly aligned; arculus making a very open angle Saxonagrion
886
2
3
4
F~GURE 4 - Saxonagrion minutus nov. gem et sp.; fig. 4.1-2 = counterpart n ~ LdLAP173B and fig. 4.3-4 = part n ~ LdLAP173A lighted differently to show veins and cells. The fig. 4.4 is an enlargment of the posterior p a r t of the wing seen from the fig. 4.3. Saxonagrion minutus nov. gen. et sp.; empreinte (Fig. 4.3, 4.4) et contre empreinte (Fig. 4.1, 4.2) gclair~es de mani~re diffgrente afin de souligner les cellules.
887 with RA (synapomorphies of the Panodonata). After the original description of the U p p e r Permian Protozygoptera Permagrion falklandicus proposed by Tillyard (1928), the organization of the veins AA, CuA and MP would be very similar to that of Saxonagrion minutus nov. gen. et sp. Nevertheless, we reexamined the original material of Permagrion (specimen n ~ Ar 2168, collection Naturh. Riksmuseum Paleozoo. Adv. Stockholm) (Bethoux 1998, Bethoux & Nel, in prep.). The interpretation of Tillyard is erroneous and the wing venation of Permagrion is completely different to that of Saxonagrion in the following points: CuA separated from MP well before MAb, presence of a strong posterior branch of MP. Permagrion is closely related to the Permolestidae sensu Bechly (1996). The character "MP with no strong posterior branch" is a plesiomorphic condition present in Panodonata, but not in Protozygoptera Permolestidae and Permagrionidae. Saxonagrion has no strict panodonatan "discal brace" sensu Carle (1982), i.e. no strong dorsal sclerotization of the aligned veins MAb and CuA. This is a plesiomorphic condition suggesting that Saxonagrion has a basal position within the Panodonata.
Saxonagrion gen. nov. shares with the Zygoptera the long petiolated wing, not present in the most basal group of the Panodonata, i.e. the Tarsophlebioptera (Upper Jurassic family Tarsophlebiidae), sister group of the "true" Odonata. Nevertheless, this character is highly homoplasic and not sufficient for its attribution to the Zygoptera. Furthermore, Saxonagrion has no known autapomorphy in its wing venation. The exact affinities of Saxonagrionidae fam. nov. within Panodonata remain uncertain. BIOSTRATIGRAPHY The Odonatoptera were highly diverse in the Upper Permian and had already a long history since the Middle Carboniferous. Upper Permian and Lower to Middle Triassic Odonatoptera belong to numerous suborders, Protanisoptera, Triadophlebiomorpha and Protozygoptera (Bechty 1996). A true Odonatoptera Discoidalia has been recently discovered in the Westphalian of England (Jarzembowski & Nel in prep.), but the previously oldest known Panodonata (Tarsophlebioptera + Zygoptera + Epiproctophora) are from the Middle and Upper Triassic (Nel et al. 1993); Protozygoptera [Permagrionoidea + Archizygoptera (= Kennedyomorpha sensu Pritykina, 1981) sensu Bechly 1996)] being the sister group of the Panodonata (Bechly 1996). From this, it appears clearly that the present fossil wing from the Upper Permian of the Lod6ve basin is the oldest record of the Panodonata.
CONCLUSION The present discovery of a putative Panodonata in the Upper Permian supports the hypothesis of Nel et al. (1999a,b) concerning the persistance of many groups of Odonatoptera through the PermoTriassic boundary and the greater antiquity of more advanced groups than was previously supposed. Nevertheless, as the Protozygoptera and Panodonata are considered as sister-groups (Bechly 1996), the presence of the former in the Lower Permian implies that of the latter during the same period. Thus, the" discovery of this new family Saxonagrionidae is not so surprising than what could be supposed. Nevertheless, both the phylogenetic analysis and the present find contradict the results of Labandeira & Seposki (1993) and Labandeira (1994), strictly based on the fossil record, indicating that the Panodonata were absent in the Upper Permian. It confirms that phylogenetic analyses are necessary in order to successfully study the historical variations of biodiversity. A c k n o w l e g e m e n t s - This paper is a contribution to the project "Pal~ontologie et Biostratigraphie du Permien" of the UMR 5561 of CNRS: "Biog~osciences" and to the 'UPRES' project "Syst~matique et crises de la biodiversit~ chez les insectes" in the Laboratoire d'Entomologie du Museum National d'Histoire Naturelle, Paris. This work was supported by the UMR 5561 of the CNRS.
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A. NEL, G. F L E C K & O. B E T H O U X Laboratoire d'Entomologie Mus6um National d'Histoire Naturelle 45 rue de Buffon F-75005 Paris e-mail: [email protected]. G. GAND UMR 5561 du CNRS, Universit~ de Bourgogne Centre des Sciences de la Terre 6, boulevard Gabriel F-21000 Dijon e-mail: umr [email protected] ou [email protected]. J. L A P E Y R I E Corniche de Fontbonne F-34700 Lod~ve J. GARRIC 16 rue des Azal~es, La Chamberte F-34070 Montpellier