Published December 31, 2003
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Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata
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Petrulevičius, J. F., Nel, A. (2003): Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata. Journal of Natural History 37 (24): 2909-2917, DOI: 10.1080/0022293021000007543, URL: http://dx.doi.org/10.1080/0022293021000007543
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- urn:lsid:plazi.org:pub:4A6B2C7BFF9860623A39FFACFB69FFE5
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- http://publication.plazi.org/id/4A6B2C7BFF9860623A39FFACFB69FFE5
References
- Bechly (1996) proposed several potential wing venation synapomorphies of the Zygoptera. (1) 'Both wing pair distinctly stalked with a petiolus that is at least somewhat longer than broad'. Fleck et al. (submitted a) noted that this character, present in Frenguelliidae, is also present in some Epiproctophora: Stenophlebiidae and that its polarity is uncertain because the Protozygoptera also have long petioles. (2) 'Both wing pair of identical shape and venation'. This character is also present in the Triadophlebiomorpha, the Protozygoptera and the Epiproctophora: Epiophlebiidae. Thus it is probably plesiomorphic for the [Zygoptera+ Epiproctophora] and subject to several homoplasies within this group. It is probable but unknown in Frenguelliidae. (3) 'On the ventral wing surface the posterior part of the basal brace (Ax0) is obliterated or covered by a rather extensive sclerotization of the wing base'. The polarization of this character is uncertain (Fleck et al., submitted a) and it is unknown in Frenguelliidae. (4) 'Reduction of the long spines of the dorsal surface of the RP and MP'. This character cannot be observed in Frenguelliidae. (5) 'A significant increase of spine-density at the apical costal margin'. This character is unknown in the Triadophlebiomorpha and Protozygoptera, thus, its exact polarity remains unknown. Furthermore, Frenguellia has no such increase. The body characters of the Zygoptera are not available in Frenguellia.
- Bechly (1996) added a possible synapomorphy of the Zygoptera, apparently present in Frenguelliidae, i.e. 'maybe the reduction of the terminal kink of the CP at the nodus is also an autapomorphy of Zygoptera, correlated with a reduction of the nodal furrow since this character is very distinct in the most basal Panodonata († Tarsophlebiidae) and most Epiproctophora, including all extant Anisoptera, while it seems to be convergently reduced in Epiophlebia and † Aeschnidiidae'. The problem is that the terminal kink of CP and the nodal furrow are also reduced in the Protozygoptera. Thus, the polarity of this character could be contrary to that proposed by Bechly (1996).
- The alignment of the second antenodal cross-vein Ax2 with the arculus, shared by the Zygoptera and Frenguelliidae, is of little value for its potential attribution to the Zygoptera. In the Protozygoptera, the arculus is in a rather variable position but generally aligned with Ax2 or in a distal position. Also, in some Epiproctophora: Stenophlebiidae, the arculus is aligned with Ax2 ( Fleck et al., submitted b). Heliocharis amazona Selys, 1853 (Calopterygoidea: Dicteriastidae) has its arculus closer to Ax1 than to Ax2. Inversely, the arculus of the Anisoptera Synthemis montaguei Campion, 1921 is shifted well distally from Ax2.
- Lastly, Frenguellia shares with the Zygoptera some characters absent in the more advanced Epiproctophora (Stenophlebioidea, Anisoptera), i.e. 'discoidal cell not divided into a triangle and a hypertriangle' (symplesiomorphy); 'vein RP2 distally recessed far from subnodus' (character of uncertain polarity as its situation varies among the Protozygoptera). Frenguellia shares with the Caloptera sensu Bechly (1996) the presence of a midfork recessed basally to a position between 12 and 26% of wing length (also present in Sieblosiidae and convergent to Lestinoidea and Hypolestinae; Bechly, 1996); and a pterostigmal brace vein obsolete (also present in Sieblosiidae and some Heterophlebioidea, Stenophlebiidae, Aeschnidiidae among the Epiproctophora).
- The phylogenetic relationships of the Sieblosiidae are still controversial. Bechly (1996) placed this family in a very basal position within the Zygoptera, as sister group of the Eucaloptera Bechly, 1996 (modern and fossil 'calopterygoid' families). Fleck et al. (submitted a) discussed this position on the basis of their nodal structures, among other characters. The Sieblosiidae and the Epiproctophora share some potential synapomorphies in the presence of a strong kink of CP aligned with the nodal Cr, a strong nodal furrow and ScP ventrally visible along costal margin, correlated with a sclerotized thickening of C distal to nodal furrow (contra Nel and Paicheler, 1994). On the contrary, Frenguellia has several plesiomorphic structures in its nodus, present in Zygoptera and absent in the Sieblosiidae, the Epiproctophora and the Tarsophlebiidae, i.e. 'no strong nodal furrow', 'costal margin basal of the nodus curved on it' and 'no thickening of C in the nodus'. Fleck et al. (submitted a) consider these last character states as plesiomorphic, contra Bechly (1996), because they are present in the basal clades Protanisoptera, Triadophlebiomorpha and Protozygoptera and in Zygoptera (Nel et al., in press; Huguet et al., submitted ).
- Frenguellia shares with the Epiproctophora, the Tarsophlebiidae, the Sieblosiidae but also the Zygoptera: Lestinoidea sensu Bechly (1996), the presence of the lestine oblique vein 'O'. Thus, this character is subject to convergencies or more.
- The hypothesis of basal positions in Epiproctophora for both Sieblosiidae and Frenguelliidae is not congruent with their rather recent ages (Oligo-Miocene and Eocene). The oldest record of Epiproctophora is Upper Triassic (Bechly, 1996, 1997). Thus both the lineages of the Sieblosiidae and Frenguelliidae should have been cryptic between the Triassic and the Eocene. A similar situation occurs with the modern epiproctophoran family Epiophlebiidae. Its sister group is at least Lower Jurassic (at least older than the oldest known Anisoptera) but it is unknown in the fossil record. These contradictions can be explained because of the fragmentary knowledge of the history of Odonata, especially for the Triassic. Only the discovery of new fossils shall help to clarify this situation. Outcrops potentially important but still understudied are the Triassic of the UK, Italy, southern Africa or Australia, among others.