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ISSN: 1989-6581
Petrulevičius & Gutiérrez (2016)
ARQUIVOS ENTOMOLÓXICOS, 16: 341-358
341
ARTIGO / ARTÍCULO / ARTICLE
New basal Odonatoptera (Insecta) from the lower Carboniferous
(Serpukhovian) of Argentina.
Julián F. Petrulevičius 1, 2 & Pedro R. Gutiérrez 1, 3
1 Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET).
2 Museo de La Plata, División Paleozoología Invertebrados, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La
Plata, Paseo del Bosque s/n, La Plata (1900), ARGENTINA. e-mail: levicius@museo.fcnym.unlp.edu.ar
3 División Paleobotánica y Paleopalinología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Av. Ángel Gallardo 470,
Ciudad de Buenos Aires (C1405DJR), ARGENTINA. e-mail: pedroraulgutierrez@gmail.com
Abstract: Three new basal species of Odonatoptera from the upper Serpukhovian (325-324 Ma) of Guandacol 1 locality,
Quebrada de las Libélulas, Guandacol Formation, La Rioja province, central West Argentina, are described. Two known
species also from the Serpukhovian, Eugeropteron lunatum Riek, 1983 and Geropteron arcuatum Riek, 1983, from
Cuestita de La Herradura, Malanzán Formation, La Rioja province, are discussed. Several higher taxa are nominated to
include these species, resulting in a new classification: 1 Superorder Odonatoptera, 1.1 Eugeroptera ord. nov., 1.1.1
Eugeropteridae, 1.1.1.1 Eugeropteron, 1.1.1.1.1 Eugeropteron lunatum, 1.1.1.1.2 Tupacsala niunamenos gen. nov. et sp. nov.,
1.2 Palaeodonatoptera taxon nov., 1.2.1 Kukaloptera ord. nov., 1.2.1.1 Kirchneralidae fam. nov., 1.2.1.1.1 Kirchnerala
treintamil gen. nov. et sp. nov., 1.2.2 Plesiodonatoptera taxon nov., 1.2.2.1 Argentinoptera ord. nov., 1.2.2.1.1
Argentinalidae fam. nov., 1.2.2.1.1.1 Argentinala cristinae gen. nov. et sp. nov., 1.2.2.2 Apodonatoptera taxon nov.,
1.2.2.2.1 Order Geroptera, 1.2.2.2.1.1 Geropteridae fam. nov., 1.2.2.2.1.1.1 Geropteron, 1.2.2.2.1.1.1.1 Geropteron
arcuatum, 1.2.2.2.2 Neodonatoptera.
Key words: Eugeroptera ord. nov., Tupacsala niunamenos gen. nov. et sp. nov., Kukaloptera ord. nov., Kirchneralidae
fam. nov., Kirchnerala treintamil gen. nov. et sp. nov., Argentinoptera ord. nov., Argentinalidae fam. nov., Argentinala
cristinae gen. nov. et sp. nov., Geropteridae fam. nov.
Resumen: Nuevos Odonatoptera (Insecta) basales del Carbonífero inferior (Serpukhoviano) de la Argentina. Nuevos
Odonatoptera basales del Serpukhoviano superior (325-324 Ma) son descriptos de la localidad Guandacol 1, Quebrada de
las Libélulas, Formación Guandacol, provincia de La Rioja, centro oeste de la Argentina. Otras dos especies conocidas del
Serpukhoviano, Eugeropteron lunatum Riek, 1983 y Geropteron arcuatum Riek, 1983, de Cuestita de La Herradura,
Formación Malanzán, provincia de La Rioja, son discutidas. Varios taxones de orden superior nuevos son nominados para
incluir estas especies, resultando en una nueva clasificación: 1 Superorden Odonatoptera, 1.1 Eugeroptera ord. nov., 1.1.1
Eugeropteridae, 1.1.1.1 Eugeropteron, 1.1.1.1.1 Eugeropteron lunatum, 1.1.1.1.2 Tupacsala niunamenos gen. nov. et sp. nov.,
1.2 Palaeodonatoptera taxon nov., 1.2.1 Kukaloptera ord. nov., 1.2.1.1 Kirchneralidae fam. nov., 1.2.1.1.1 Kirchnerala
treintamil gen. nov. et sp. nov., 1.2.2 Plesiodonatoptera taxon nov., 1.2.2.1 Argentinoptera ord. nov., 1.2.2.1.1
Argentinalidae fam. nov., 1.2.2.1.1.1 Argentinala cristinae gen. nov. et sp. nov., 1.2.2.2 Apodonatoptera taxon nov.,
1.2.2.2.1 Orden Geroptera, 1.2.2.2.1.1 Geropteridae fam. nov., 1.2.2.2.1.1.1 Geropteron, 1.2.2.2.1.1.1.1 Geropteron
arcuatum, 1.2.2.2.2 Neodonatoptera.
Palabras clave: Eugeroptera ord. nov., Tupacsala niunamenos gen. et sp. nov., Kukaloptera ord. nov., Kirchneralidae
fam. nov., Kirchnerala treintamil gen. nov. et sp. nov., Argentinoptera ord. nov., Argentinalidae fam. nov., Argentinala
cristinae gen. nov. et sp. nov., Geropteridae fam. nov.
Recibido: 29 de noviembre de 2016
Publicado on-line: 20 de diciembre de 2016
Aceptado: 5 de diciembre de 2016
urn:lsid:zoobank.org:pub:3F60AC3C-F343-4460-8B43-C9C4242F794B
Petrulevičius & Gutiérrez (2016): New basal Odonatoptera from the lower Carboniferous (Serpukhovian) of Argentina.
342
Introduction
Odonatoptera basal to the Neodonatoptera Bechly, 2001 are extremely rare in the fossil record.
Only two previous genera were described from the Argentinean locality Cuestita de la Herradura, i.e.,
Eugeropteron Riek, 1983 and Geropteron Riek, 1983 (Riek & Kukalová-Peck, 1984). These two insects and
Xenoptera riojaensis Pinto, 1986 were recovered from the Malanzán Formation (Pinto, 1986), considered
a lateral equivalent of Guandacol Formation (Césari & Gutiérrez, 2001) (see below). Eugeropteron and
Geropteron were considered the most basal of the Odonatoptera (Kukalová-Peck, 1991, 2009;
Brauckmann et al., 1996; Bechly, 2007). These taxa and those described here are coincidently treated
as basal to Neodonatoptera in the present contribution.
The new species described here were exhumed from the base of the Guandacol Section of the
Guandacol Formation with an age of circa 325-324 Ma (Césari et al., 2011). Oldest records of winged
insects in the world are one undetermined species of Archaeorthoptera from the Paskov Mine (circa
324-325 Ma), Czech Republic (Prokop & Nel, 1996), and a species of Palaeodictyoptera from Delitzsch
(320-323 Ma), Germany (Brauckmann & Schneider, 1996; Zhang et al., 2013). Concerning oldest
Odonatoptera, there are four genera with four species of Neodonatoptera from the German locality
Hagen-Vorhalle (320 Ma) (Zessin et al., 2011) and Erasipteron Pruvost, 1933 from Horní Suchá (318 Ma),
Czech Republic (Zessin, 2008).
One of the specimens described here (see below) is the most complete and prettiest female
dragonfly from the Carboniferous, with part of head, thorax with 6 wings and abdomen with ovipositor.
The preservation of prothoracic wings is unusual for basal Odonatoptera, being the only other species
worldwide Erasipteroides valentini Brauckmann, 1985 from the Carboniferous of Germany (Brauckmann
et al., 1985; Bechly et al., 2001). The specimen has been part of a research about fly adaptations in
Carboniferous insects (Wotton et al., 1998; Vogel, 1998; Wotton & Kukalová-Peck, 2000) and has been
discussed and figured in several contributions (Wotton et al., 1998; Gutiérrez et al., 2000; Bechly et al.,
2001; Bechly, 2007; Kukalová-Peck, 2008; Staniczek et al., 2011). In this work it is described, named
and included in a new classification.
Materials and methods
In this work, we follow the wing venation nomenclature of Kukalová-Peck (1983), amended by
Kukalová-Peck (1991, 2009), and contributions by Riek & Kukalová-Peck (1984), Nel et al. (1993), and
Bechly (1996). Abreviations of the wing venation used in the text and figure are: CP (Costa Posterior),
ScA (Subcosta Anterior), ScP (Subcosta Posterior), RA (Radial Anterior), RP (Radial Posterior), MA
(Media Anterior), MP (Media Posterior), CuA (Cubital Anterior), CuP (Cubital Posterior), CuAc (Cubital
Anterior crossing), CuPc (Cubital Posterior crossing), AA (Anal Anterior), AP (Anal Posterior). Crossveins
are in lower case letter.
We use the groundplan method of phylogenetic Systematics sensu Hennig (1968, 1981) and
Kukalová-Peck (2009). In this sense, the groundplan plesiomorphies and synapomorphies could be
retained by basal and successive intermediate taxa and could be not visible in the crown group. As
example of typical groundplan character of Odonatoptera, the presence of “prothoracic winglets
obliquely directed anteriorly with venation and articulated” is a synapomorphy of Odonatoptera only
preserved in the most basal taxa: Argentinoptera ord. nov. and Eomeganisoptera, and supposedly
present in Geroptera and Kukaloptera ord. nov.
The protowing evolutionary model of wing articulation was proposed by Kukalová-Peck (1983,
1998, 2008) after an extensive study of extinct and extant insects (Kukalová-Peck et al., 2009). The
higher phylogenetical classification of basal Odonatoptera proposed here adds new taxa and new
characters and discusses with some coincidence with the phylogenetic system of Bechly (1996, 2007).
ARQUIVOS ENTOMOLÓXICOS, 16: 341-358
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Specimens are deposited in two institutions of Argentina. The Museo Argentino de Ciencias
Naturales Bernardino Rivadavia, Ciudad Autónoma de Buenos Aires, Argentina, under the acronym
MACN-In, and the Museo de La Plata, División Paleozoología Invertebrados, La Plata, Argentina, under
the acronym MLP.
Age and strata. Specimens described and studied here come from two localities and
formations 200 km distant, in La Rioja Province, Argentina. The new species described come from the
base of Guandacol Formation in the Paganzo Basin (Gutiérrez et al., 2000). They were collected by one
of us (P.G.) in Guandacol 1 locality, Quebrada de las Libélulas (South East from Quebrada Las
Blanquitas), Cerro Guandacol. Guandacol Formation strata are deposited in the interval between 325 and
318.79 Ma (Gulbranson et al., 2010; Césari et al., 2011). The insects were found in the very base of the
formation and could be restricted to 325-324 Ma, which corresponds to the upper Serpukhovian sensu
Cohen et al. (2013; updated 2016).
The remaining species (Riek & Kukalová-Peck, 1984) were exhumed from Cuestita de la
Herradura locality from the Malanzán Formation. The Malanzán Formation lacks absolute datation until
now, though in previous literature, based on flora and microflora, it is considered to be a lateral
equivalent of Guandacol Formation (Césari & Gutiérrez, 2001). The discovery in the last fieldtrip of a
specimen of Megasecoptera: Xenopteraidae assignable to Xenoptera in Guandacol Formation
(Petrulevičius & Gutiérrez, in prep.) is coincident with this equivalence.
New classification of basal Odonatoptera (see below)
1 Superorder Odonatoptera Lameere, 1900
Included taxa: Eugeroptera ord. nov. and Palaeodonatoptera taxon nov.
1.1 Eugeroptera ord. nov.
Included taxa: Eugeropteridae Riek, 1983 with Eugeropteron Riek, 1983: Eugeropteron lunatum
Riek, 1983 and Tupacsala niunamenos gen. nov. et sp. nov.
1.2 Palaeodonatoptera taxon nov.
Included taxa: Kukaloptera ord. nov. and Plesiodonatoptera taxon nov.
1.2.1 Kukaloptera ord. nov.
Included taxa: Kirchneralidae fam. nov. with Kirchnerala treintamil gen. nov. et sp.
nov.
1.2.2 Plesiodonatoptera taxon nov.
Included taxa: Argentinoptera ord. nov. and Apodonatoptera taxon nov.
1.2.2.1 Argentinoptera ord. nov.
Included taxa: Argentinalidae fam. nov. with Argentinala cristinae gen.
nov. et sp. nov.
1.2.2.2 Apodonatoptera taxon nov.
Included taxa: Geroptera Brodsky, 1994 and Neodonatoptera Bechly,
1996
1.2.2.2.1 Order Geroptera Brodsky, 1994
Included taxa: Geropteridae fam. nov. with Geropteron
arcuatum Riek, 1983
1.2.2.2.2 Neodonatoptera Bechly, 1996
Petrulevičius & Gutiérrez (2016): New basal Odonatoptera from the lower Carboniferous (Serpukhovian) of Argentina.
344
Systematic Paleontology
Hydropalaeoptera Rohdendorf, 1968 sensu Kukalová-Peck, 2009 (= Eupalaeoptera Bechly, 2003)
Palaeoptera Martynov, 1923
1. Superorder Odonatoptera Lameere, 1900
1.1. Eugeroptera ord. nov.
Included taxa: Eugeropteridae Riek, 1983 with Eugeropteron Riek, 1983 including only Eugeropteron
lunatum Riek, 1983, and Tupacsala niunamenos gen. nov. et sp. nov.
Phylogenetic definition: Eugeroptera ord. nov. shall include all the Odonatoptera more closely related
to Eugeropteron lunatum Riek, 1983 than to any of the type species of the other type genera of the
Odonatoptera group taxa (stem-based definition).
Diagnosis: Characters from mesothoracic and metathoracic wing venation (see below). (1) wings with
relatively undeveloped anal field (more marked in forethoracic wing); (2) MP unbranched; (3) CuP with a
kink at the point of contact with AA; (4) archaedictyon reduced; (5) presence of a subcostal brace; (6)
Presence of an anal brace in fore- and hindwings; (7) presence of a cubital cell; (8) ScP short (up to RP1
and RP2 bifurcation) in hindwings; (9) free ScA(+) relatively short, forked into a ScA1+2 branch fused
with the anterior margin, and a strong crossvein like branch ScA3+4; (10) subcostal brace formed by
ScA+ScA fork+scp-ra+ra-rp; (11) ScA3+4+scp-ra+ra-rp aligned; (12) MP linked by a crossvein to CuA in
hindwings; (13) MP bending anteriorly before and in the connection (via crossvein) with CuA; (14) RA
bending to wing margin and running adjacent at the level of RP1-RP2 bifurcation (no place to
pterostigma); (15) cubital cell in (forewings ? and) hindwings formed by five elements: CuA, CuA
crossing, CuP crossing, CuP, cua-cup; (16) AA1+2 not fused with CuP in fore- and hindwings; (17) anal
brace composed of five venal elements: AA, AA1+2, cup-aa1+2 crossvein, kink in CuP, cua-cup crossvein;
(18) pectination of AA1+2 in mesothoracic wing; (19) RP+MA elevated basally to convex level.
Phylogenetic systematic: Eugeroptera ord. nov. retains nearly all the synapomorphies of the stem
group of the Odonatoptera (Characters: 1-7). Character 8 is a synapomorphy of the new order
(convergent to Eomeganisoptera, Bechly, 2007). Characters 9-18 are groundplan symplesiomorphies of
Odonatoptera. Character 19 seems to be a synapomorphy for Hydropalaeoptera (see Remarks of
Neodonatoptera). Kukalová-Peck (2016) considered the RP elevated basally to convex level a
synapomorphy for Palaeoptera.
Etymology: Named after the family Eugeropteridae Riek, 1983.
Remarks: Eugeropteron and Tupacsala gen. nov. remain the only genera of Eugeropteridae. Geropteron
placed by Riek & Kukalová-Peck (1983) in Eugeropteridae Riek, 1983 is transferred to Geropteridae
fam. nov. and maintained in the Order Geroptera Brodsky, 1994 (see below).
One of the notable characters of Odonatoptera basal to Neodonatoptera is the R stem
separated into RA and RP from the very base. The separated closely parallel RA and RP is only present
(partially preserved) somewhere else in Bojophlebia prokopi (Kukalová-Peck, 1985; Sroka et al., 2014). In
the Odonatoptera basal to Neodonatoptera these veins are not only independent but not parallel;
basally RA bends anteriorly and RP bends posteriorly and between them they could have even one
crossvein, character that is unique (autapomorphy) from these basal forms (Figs. 1-2). Other unique
character from Odonatoptera basal to Neodonatoptera is that free RP is convex (+) in the base, getting
concave (-) just after the emergence of the MA (Figs. 1-2, 5-6) (see Remarks in Neodonatoptera). RP
ARQUIVOS ENTOMOLÓXICOS, 16: 341-358
345
was considered concave by Riek in Riek & Kukalová-Peck (1984: text and fig. 1) and Kukalová-Peck (1991:
fig. 6.15A) but rectified to convex in Kukalová-Peck (2009: see fig. 7) and considered an autapomorphy
of Palaeoptera by Kukalová-Peck (2016).
1.1.1. Eugeropteridae Riek, 1983
Type genus: Eugeropteron Riek, 1983.
1.1.1.1. Eugeropteron Riek, 1983
Type species: Eugeropteron lunatum Riek, 1983.
1.1.1.1.1. Eugeropteron lunatum Riek, 1983
Type material: Holotype specimen MLP 12886. Cuestita de La Herradura locality, province of La Rioja,
Northwest Argentina, at paleolatitude ≈ 60º. Malanzán Formation, La Divisoria Member, Serpukhovian
(circa 325-324 (?) Ma) (Césari et al., 2011; Césari & Gutiérrez, 2001).
Remarks: The specimen was named by Riek in Riek & Kukalová-Peck (1983). In the description and
analysis of the species in the cited paper, and in Kukalová-Peck (1991), the RP was considered concave in
all its length. In a more recent paper, Kukalová-Peck (2008) noted the convexity of RP(+) before the
arising of MA. So, both RA and RP are convex. After the arising of MA, RP becomes concave. See below
our consideration that while RP is convex, MA is fused to it. In present contribution we interpret the
cup-aa1+2 as a crossvein as in Kukalová-Peck (2009) and contrary to Riek & Kukalová-Peck (1983), where
they interpret it as AA1.
1.1.1.1.2 Tupacsala gen. nov.
Type species: Tupacsala niunamenos sp. nov.
Diagnosis: Characters from metathoracic wing venation (see below). (1) CuP kink; (2) Cubital cell; (3) MP
single; (4) anal brace; (5) CuA is not fused with CuP in the hindwings; (6) Crossvein between CuP and AA
long; (7) Cubital cell oblique; (8) CuA with four branches; (9) and MA bifurcated well basal to RP
bifurcation.
Etymology: Dedicated to the memory of José Gabriel Condorcanqui Noguera, “Túpac” Amaru II (1738-
1781) and to Milagro Amalia Ángela “Sala” (1963-). Túpac Amaru in 1780-1781 initiated a revolt against
Spanish State and its rules. He was tortured (forced to witness the execution of the sentences
imposed on his family), executed and quartered to be exposed (Cline, 2015). Milagro Sala (1963-) is a
prominent Argentine social leader, Secretary of the “Organización Barrial Túpac Amaru” and
Parliamentary of the Parlasur imprisoned with other members of the organization since January 16,
2016.
1.1.1.1.2.1 Tupacsala niunamenos sp. nov. (Figs. 1-2)
Previous references to Tupacsala niunamenos gen. et sp. nov.:
2000: Eugeropteridae [Gutiérrez et al. (2000)]
Type material: Holotype specimen MACN-In 2678C. Guandacol 1 locality, Quebrada de las Libélulas,
Cerro Guandacol, province of La Rioja, Northwest Argentina, at paleolatitude ≈ 60º. Lowermost part of
Guandacol Formation (Gutiérrez et al., 2000), Serpukhovian (circa 325-324 Ma) (Césari et al., 2011).
Diagnosis: That of the genus by monotypy.
Petrulevičius & Gutiérrez (2016): New basal Odonatoptera from the lower Carboniferous (Serpukhovian) of Argentina.
346
Description: A basal fragment of a metathoracic wing. Only the counterpart (reverse) is known.
Venation three-dimensionally preserved. Wide anal area and development of anal veins indicate a
hindwing.
Metathoracic wing fragment; length of the fragment 19.9 mm; width of the fragment 12.9 mm.
RA shortly preserved well distal to discoidal cell; RP shortly preserved diverging from RA. MA
bifurcated well basal to RP bifurcation. MP single. Cubital cell partially preserved, less vertical and
narrower than Argentinala. Cubital cell with four partially preserved elements: CuA crossing curved, CuP
crossing slightly curved, CuP (0.6 mm long) (nearly straight and longer than cua-cup), and cua-cup quite
long (0.3 mm) (similar than Eugeropteron). CuP kink strong. CuA with three veins. CuP bifurcated basal
to CuA. AA1+2 connected to CuP via a long crossvein. AA not fused with CuP connected by cup-aa (1.1 mm
long). AA1+2 with three or four branches. AA3 sinuous. AP long and single.
Etymology: Dedicated to “Ni Una Menos” (No one less), a collective against gender violence. It is a
collective campaign that arose from the need to say “enough femicides”, because in Argentina every 30
hours a woman is killed just by being a woman.
Discussion: We decided to describe and name this fragmentary specimen due to the rarity of preserved
winged insects from lower Carboniferous worldwide and to highlight the diversity of these strange stem
dragonflies. Although it is not evident at first sight, fortunately it preserves some interesting
characters to place it within basal Odonatoptera. The specimen is an Odonatoptera because has 1) a CuP
kink, 2) a cubital cell, 3) a MP single, and 4) an anal brace. It is an Odonatoptera basal to
Apodonatoptera taxon nov. because the CuA is not fused with the CuP in the hindwings. The crossvein
between CuP and AA is long and the cubital cell is oblique contrary to Argentinoptera ord. nov. which
have a short crossvein and a vertical cubital cell. The new species seems to fit better with Eugeroptera
ord. nov. because of the oblique cubital cell. Eugeroptera ord. nov. has both forewing and hindwing
cubital cells oblique and quite broad like the new species and unlike Kukaloptera ord. nov. having it
horizontal and narrower. In spite of the fragmentary condition of the wing that makes difficult the
comparison with Kukaloptera ord. nov. we place the new species provisionally in Eugeroptera ord. nov.:
Eugeropteridae. Differences with Eugeropteron are the MA bifurcated well basal to RP bifurcation and
a CuA with four branches. These differences allows us to erect a new genus Tupacsala gen. nov.
1.2. Palaeodonatoptera taxon nov.
Included taxa: Kukaloptera ord. nov. and Plesiodonatoptera taxon nov.
Synapomorphies: Fusion of MP with CuA in forewings. This character is tentative as only Kirchnerala
gen. nov. and Argentinala gen. nov. preserve this part of the forewing among the most basal
Odonatoptera. Eugeropteron lacks the antero-basal part of the forewing.
1.2.1. Kukaloptera ord. nov.
Included family: Kirchneralidae fam. nov.
Phylogenetic definition: Kukaloptera ord. nov. shall include all the Odonatoptera more closely related
to Kirchnerala treintamil gen. et sp. nov. than to any of the type species of the other type genera of
the Palaeodonatoptera taxon nov. group taxa (stem-based definition).
Etymology: In honour of the most influential paleoentomologist Jarmila Kukalová-Peck. Kukalová-Peck's
theories about monophyly of Arthropoda based on fossil and recent species are foundational of modern
view of these animals. Her theories include the homologation of the multirrameous Arthropoda limb and
detailed morphologies of specialization in head, thorax and abdomen. One of her outstanding results is
the comprehension of the limb origin of the insect wing and the homologation of wing articulation and
venation.
ARQUIVOS ENTOMOLÓXICOS, 16: 341-358
347
Diagnosis: Characters from mesothoracic and metathoracic wing venation (see below). (1) anterior
articular plate (AAP), with an “h” shaped basialare (B-C), a large bi-lobed fulcalare (F-C) narrowly
separated from the axalare (AX-C) by a deeply incised groove-like suture; (2) wings with relatively
undeveloped anal field (more marked in mesothoracic wing); (3) MP unbranched; (4) CuP with a kink at
the point of contact with AA; (5) archaedictyon reduced; (6) presence of a subcostal brace; (7)
presence of an anal brace in forewings; (8) presence of a cubital cell; (9) reduction of pectination of
AA1+2 and AA3+4 pectinated in forewing; (10) cubital cell long, straight and horizontal in forewings; (11)
MP shortly fussed to CuA in forewings; (12) ScA3+4+scp-ra+ra-rp aligned; (13) MP bending anteriorly
before and in the connection with CuA; (14) RA bending to wing margin and running adjacent at the level
of RP1-RP2 bifurcation (no place to pterostigma).
Phylogenetic systematic: Eugeroptera ord. nov. retains nearly all the synapomorphies of the stem
group of the Odonatoptera (Characters 1-8). Character 9-10 are synapomorphies of the new order.
Character 11 is a synapomorphy of Palaeodonatoptera taxon nov. Characters 12-14 are groundplan
symplesiomorphies of Odonatoptera.
1.2.1.1. Kirchneralidae fam. nov.
Type genus: Kirchnerala gen. nov.
Diagnosis: That of the order by monotypy.
1.2.1.1.1 Kirchnerala gen. nov.
Type species: Kirchnerala treintamil sp. nov.
Diagnosis: That of the order by monotypy.
Etymology: In honour of Argentinean President (2003-2007) Néstor Kirchner, who passed away on
October 27th, 2010 and the Latin “ala” meaning wing. On July 16th, 2003 he agreed to a public meeting
requested by two designed delegates of CONICET External Postdoctoral Fellows in the Argentinean
Embassy in Paris. In relation to our concerns, he explained us the promissory future plans of the
government for the scientific system in our country. After our return to Argentina in 2004, his sayings
were corroborated year by year and continued in the same way by the next (twice) President of the
country, Cristina Fernández de Kirchner. At the end of the meeting he proposed to Gabriel Moron and
one of us (JFP) to visit him when returned; a pending issue that will never be carried out.
1.2.1.1.1.1 Kirchnerala treintamil sp. nov. (Figs. 3-4)
Previous references of Kirchnerala treintamil gen. et sp. nov.:
2000: Eugeropteridae [Gutiérrez et al. (2000)]
2009: Eugeropteron sp., the right forewing of a complete adult dragonfly (Wootton & Kukalová-Peck 2000, fig.
10); fig. 7A [Kukalová-Peck (2009)]
2009: Eugeropteron lunatum; fig. 3 [Kukalová-Peck et al. (2009)]
Type material: Holotype MACN-In 2678D, housed in the Museo Argentino de Ciencias Naturales
Bernardino Rivadavia, Buenos Aires, Argentina. Guandacol 1 locality, Quebrada de las Libélulas, Cerro
Guandacol, province of La Rioja, Northwest Argentina, at paleolatitude ≈ 60º. Lower part of Guandacol
Formation (Gutiérrez et al., 2000), Serpukhovian (circa 325-324 Ma) (Gulbranson et al., 2010; Césari et al.,
2011).
Description: The mesothoracic wing of an adult drangonfly, with part of tergum preserved and bearing
two axillary plates. Only the part (obverse) is known. Venation and articulation three-dimensionally
preserved. Narrow anal area indicates a forewing.
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Wing: mesothoracic wing fragment, preserved length 29.2 mm; probable total length 34.6 mm;
maximum width 11.9 mm. The wing is about 2.9 times as long as broad. Anterior margin (PC) slightly
falcate. Curved posterior margin bent slightly concavely between AP and AA and strongly convexly after
that. CP- short and single, joining CA+ close to base. ScA+ forming a prominent recurrent subcostal
brace (ScA+ScA fork+scp-ra). ScA+ relatively short, forked into a ScA1+2 branch fused with the
anterior margin well before 1/12 of wing length, and a strong crossvein like branch ScA3+4, curved and
ending on ScP(-). Several crossveins not aligned between wing margin to ScP (≈8), ScP to RA (≈18) and
RA to RP (≈10). ScP ending on wing margin (?) up to RP1 and RP2 bifurcation. RA(+) separated from RP(+)
from the very base. RA and RP arched and divergent (0.4 mm) in the middle of the segment from the
base to MA emergence. Strong crossvein like veins aligned ScA3+4, scp-ra, and ra-rp. RA bending to
wing margin at the level of RP1-RP2 bifurcation. RP convex up to MA emergence (at the level of distal
half of cubital cell), distally concave. RP(-) forked beyond mid-wing into RP1+2 and RP3+4. RP1+2
bifurcated into RP1 and RP2. IR2 between RP1+2 and RP3+4. MA forked distal of mid-wing in three
branches. MP± bending anteriorly (MP concavely curved) before and in the connection with CuA. MP
shortly fussed with CuA. MP single. Cu- bending to anterior and forked into CuA and CuP forming a
cubital cell. Cubital cell (horizontal) long (4.4 mm) and narrow (0.6 mm) with five elements: CuA (curved
and short) (1.2 mm), CuA crossing (curved and long) (3.4 mm), CuP crossing (curved and long) (3.4 mm),
CuP (slightly bended anteriorly and short) (0.85 mm), cua-cup (very short) (0.07 mm). CuP kink strong.
CuA with three branches and connected via a bifurcated (Y like) crossvein to CuP. CuP not fused with
AA connected by a long crossvein (cup-aa1+2). AA1+2 not pectinated with four branches. AA3+4
pectinated with three branches sinuous at the base. Ap- long and single. Jugal area scalloped. Anal
brace composed of five veinal elements: AA, AA1+2, cup+aa1+2, kink in CuP, cua-cup.
Mesothoracic wing articulation: Anterior articular plate (AAP) partially preserved formed by
precostal and costal pteralia, with an "h" shaped basialare (B-C), a large bi-lobed (?) costal fulcalare (F-
C) narrowly separated from the costal axalare (AX-C) by a deeply incised groove-like suture. Subcostal
pteralia row free with well developed axalare (A-Sc) and fulcalare (F-Sc) and not so noticeable
basivenale (B-Sc) and proxalare (PR-Sc). Posterior articular plate (PAP) formed by radial, medial and
cubital pteralia. A deep groove posterior to cubital pteralia coinciding with the arising of Cu. Anal and
jugal veins arising obliquely from the posterior articular plate and from the posterior side.
Diagnosis: That of the order by monotypy.
Etymology: Dedicated to the 30,000, in Spanish “treinta mil” (presentes, ahora y siempre!), detained-
disappeared by the Argentinean Military-Civil-Ecclesiastic Dictatorship 1976-1983. Mil is also a
linguistic morph in Kakan (Diaguita) language (Nardi, 1979) from La Rioja province whose meaning has
unfortunately disappeared (Nardi, 1979) as well as its speaking (Fabre, 2005).
Discussion: The specimen could be included into Odonatoptera because of the: (1) anterior articular
plate (AAP), with an “h” shaped basialare (B-C), a large bi-lobed fulcalare (F-C) narrowly separated from
the axalare (AX-C) by a deeply incised groove-like suture; (2) wings with relatively undeveloped anal
field (more marked in mesothoracic wing); (3) MP unbranched; (4) CuP with a kink at the point of contact
with AA; (5) archaedictyon reduced; (6) presence of a subcostal brace; (7) presence of an anal brace in
forewings; (8) presence of a cubital cell. It could be included in Palaeodonatoptera taxon nov. because
of the MP shortly fussed to CuA in forewings. It could be excluded from the more derived
Plesiodonatoptera taxon nov. because of the absence of (1) CuA-CuP fused in forewings; (2) loss of the
crossvein between RA and RP; and (3) fusion of CuP with AA1+2 in forewings. It also could not be
included in the derived Apodonatoptera taxon nov. because of the absence of (1) RA parallel one cell
distant to anterior wing margin (place to pterostigma); (2) CuA-CuP fused in hindwings and; (3) AA1+2
fused with CuP in hindwings. The new genus and species has two autapomorphies, i.e., (1) reduction of
pectination of AA1+2 and AA3+4 pectinated in forewing and (2) the cubital cell long, straight and
horizontal in forewings.
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Remarks: The drawing of fig. 7A of Kukalová-Peck (2009) corresponds to Kirchnerala treintamil gen. et
sp. nov. She referred erroneously to it as “Eugeropteron sp., the right forewing of a complete adult
dragonfly (Wootton & Kukalová-Peck 2000, fig. 10)”. Her mention is erroneous since the complete adult
dragonfly of Wootton & Kukalová-Peck (2000: fig. 10) corresponds to the species described below as
Argentinala cristinae gen. nov. et sp. nov.
1.2.2. Plesiodonatoptera taxon nov.
Included taxa: Argentinoptera ord. nov. and Apodonatoptera taxon nov.
Synapomorphies: CuA-CuP fused in forewings. Loss of the crossvein between RA and RP. Fusion of CuP
with AA1+2 in forewings.
1.2.2.1 Argentinoptera ord. nov.
Included family: Argentinalidae fam. nov.
Phylogenetic definition: Argentinoptera ord. nov. shall include all the Odonatoptera more closely
related to Argentinala cristinae gen. et sp. nov. than to any of the type species of the other type
genera of the Plesiodonatoptera taxon nov. group taxa (stem-based definition).
Phylogenetic systematic: Eugeroptera ord. nov. retains nearly all the synapomorphies of the stem
group of the Odonatoptera (Characters 1-7). Character 8 is a synapomorphy of the new order
(convergent to Eomeganisoptera, Bechly, 2007). Characters 9-18 are groundplan symplesiomorphies of
Odonatoptera.
Diagnosis: Characters from female with 6 wings and body (see below). (1) anterior articular plate (AAP),
with an “h” shaped basialare (B-C), a large bi-lobed fulcalare (F-C) narrowly separated from the axalare
(AX-C) by a deeply incised groove-like suture; (2) MP unbranched; (3) CuP with a kink at the point of
contact with AA; (4) archaedictyon reduced; (5) presence of a subcostal brace; (6) presence of an anal
brace in fore- and hind wings; (7) presence of a cubital cell; (8) prothoracic winglets obliquely directed
anteriorly with venation and articulated; (9) abdomen elongated and slender; (10) MP shortly fussed to
CuA in forewings; (11) CuA-CuP fused in forewings; (12) CuP-AA fusion in forewings; (13) loss of
crossvein between RA-RP; (14) MP touching punctually CuA in hindwings (shortening of Crossvein (?));
(15) abdominal segment 10 somewhat longer than 9; (16) segments 9 and 10 modified with a rounded
genitalia; (17) short and sickle like ovipositor; (18) sharp and sickle valve 1, valve 2 string along with
valve 1 and valve 3 curved and sheathing laterally V1 and V2; (19) ScP long (distal to RP1 and RP2
bifurcation) not reaching the anterior wing margin and disappearing before the apex; (20) MP going
anteriorly before and in the connection with CuA; (21) RA bending to wing margin and running adjacent
at the level of RP1-RP2 bifurcation (no place to pterostigma).
Phylogenetic systematic: Argentinoptera ord. nov. retains nearly all the synapomorphies of the
Odonatoptera (Characters: 1-9). Character 10 is a synapomorphy of Palaeodonatoptera taxon nov.
Characters 11-13 are synapomorphies of Plesiodonatoptera taxon nov. Characters 14-18 are
synapomorphies of the new order. Characters 19-21 are groundplan plesiomorphies of Odonatoptera.
Etymology: Named after the family Argentinalidae fam. nov.
Remarks: Even if MP changes its direction at the fusion with CuA, it is still bending posteriorly, unlike in
Neodonatoptera whose MP strongly bends anteriorly (synapomorphy).
1.2.2.1.1 Argentinalidae fam. nov.
Type genus: Argentinala gen. nov.
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Diagnosis: That of the order by monotypy.
1.2.2.1.1.1 Argentinala gen. nov.
Type species: Argentinala cristinae sp. nov.
Diagnosis: That of the order by monotypy.
Etymology: Named after the country, República “Argentina”, and the Latin “ala” meaning wing.
1.2.2.1.1.1.1 Argentinala cristinae sp. nov. (Figs. 5-6)
Previous references to Argentinala cristinae gen. et sp. nov.:
1998: Undescribed eugeropterid from Rioja, Argentina (Carbonifeous, Namurian/Westphalian); fig. 2(A)
[Wootton et al. (1998)]
1998: 320-millon-year-old dragonfly from Argentina; fig. Ancient aviator [Vogel (1998)]
2000: Undescribed geropterid from Westphalian of Rioja, Argentina [Wootton & Kukalová-Peck (2000)]
2000: Eugeropteridae gen. et sp. indet., winged specimen; fig. 3 [Gutiérrez et al. (2000)]
2001: Undescribed Eugeropteridae gen. et spec. nov., Upper Carboniferous of Argentina (reconstruction drawing
by J. Kukalová-Peck) [Bechly et al. (2001)]
2005: Reconstruction of Eugeropteron (Eugeropteridae). Although superficially resembling a
palaeodictyopteran, eugeropterids were early odonatopterans and perhaps stem-group relatives of odonatans;
fig. 6.24 [Grimaldi & Engel (2005)]
2007: Recently discovered and still undescribed specimen (Kukalová-Peck pers. comm., and pers. observ. on a
cast of the referring specimen) [Bechly (2007]
2008: Carboniferous dragonfly with three pairs of wings; fig. 16 [Kukalová-Peck (2008)]
2011: Eugeropteridae gen. et sp. n., cast of undescribed fossil specimen from the Upper Carboniferous of La
Rioja (Argentina) in coll. Javier Muzón (ILPLA); fig. 49 [Staniczek et al. (2011)]
2013: “Rekonstrukce zástupce skupiny †Geroptera, jejich prothorakální křidélka byla namířena směrem dopředu
(převzato z Wootton a Kukalová-Peck, 2000 upraveno Grimaldi a Engel, 2005)”; fig. 3 [Pecharová (2013)]
2016: The most primitive known fossil dragonflies Eugeropteridae had a short ovipositor [Prokop et al. (2016)]
2016: Reconstruction of Eugeropteron (Eugeropteridae); fig. 1 [Hallam (2016)]
Type material: Holotype female specimen MACN-In 2678A/B. Guandacol 1 locality, Quebrada de las
Libélulas, Cerro Guandacol, La Rioja province, Northwest Argentina, at paleolatitude ≈ 60º. Lower part of
Guandacol Formation (Gutiérrez et al., 2000), lower Serpukhovian (circa 325-324 Ma) (Césari et al., 2011).
Description: Part and counter-part of a complete female specimen with 6 wings. Head partially
preserved with one rounded and conic eye (?). Thorax with six wings, prothoracic ones smaller with
venation and apparent articulation, mesothoracic quite slender without an anal region developed and
metathoracic ones wider with a developed anal field. Mesothorax and wing in dorsal view with preserved
articular plates. Wing span (mesothoracic wings) ≈ 93.7 mm.
Wings: Prothoracic wing with quadrangular/subtriangular shape directed upwards (4.6 mm long, 3
mm wide). Six major longitudinal convex veins. Four veins originating from anterobasal part of the wing.
First and second vein slightly combed posteriorly and ending in the margin. Third and fourth veins
slightly combed anteriorly. Fifth vein well curved and well separated from the fourth. Sixth vein short
and strongly curved.
Mesothoracic wings: right wing almost complete, 44.5 mm long; left wing nearly complete;
maximum width of right wing 12.3 mm, left one 12.5 mm. The wing is about 3.6 times as long as broad.
Anterior margin slightly falcate. Curved posterior margin bent slightly concavely between AP and AA
and strongly convexly after that. CP- long running well separate and parallel to the wing margin, joining
ScA basal to CA+ arrival. ScA+ forming a rather weak subcostal brace in right wing (ScA+ScA fork+scp-
ra). ScA+ relatively short, forked into a ScA1+2 (reaching CP well before 1/12 of wing length) and an
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oblique crossvein like branch ScA3+4 that ends on ScP-. In left wing there are not ScP3+4 and scp-ra
preserved. Several crossveins between wing margin to ScP (≈ 7 on left, ≈ 8 on right), between ScP to
RA (≈ 11) and RA to RP (≈ 15). Crossveins from wing margin to ScP and ScP to RA not aligned. ScP long,
noticeably curved and sinuous at the base, ending on the wing margin at the height of IR2. RA+
separated from RP+ from the very base. RA and RP arched and divergent (0.3 mm) in the middle of the
segment from the base to MA emergence (6.2 mm distant wing base). Absence of crossvein between
RA-RP. Crossvein scp-ra aligned to ScA3+4 (preserved in right wing). RA bending to wing margin at the
level of RP1-RP2 bifurcation running adjacent to wing margin (no space for pterostigma). RA bifurcates
at the wing apex (right wing). RP convex up to MA emergence (at the level of distal half of cubital cell),
distally concave. RP- forked beyond mid-wing into RP1+2 and RP3+4. RP1+2 bifurcated into RP1 and RP2.
RP1 bifurcated. IR1 between RP1 and RP2, 1/3 longer than RP1 biburcation. IR2+ between RP2 and
RP3+4. MA forked distal of mid-wing in three branches. MP± bending anteriorly before and in the
connection with CuA. Free MP 3.8 mm long. MP single fused shortly with CuA for about 1 mm. Cu-
bending to anterior and forked into CuA and CuP forming a cubital cell. Cubital cell (horizontal) (4.2 mm
long and 0.8 mm wide) developed and broad with six elements: CuA (curved and short) (0.3 mm), CuA+MP
(0.96 mm), CuA crossing (curved and long) (2.9 mm), CuP crossing (curved and long) (2.7 mm), CuP +AA1
(0.8 mm in rightwing, 1.1 in leftwing), CuP (nearly straight and short) (1.0 mm in rightwing, 0.5 in
leftwing). CuP kink strong. CuA fused with CuP shortly after cubital cell (0.3 mm long in right wing, 0.5
mm long in left wing). CuA with three terminal branches. CuP slightly falcate and single. CuP+AA1 fused
just to the middle of posterior side of cubital cell. AA1 with five branches, AA2 and AA3 single, AA4
with two branches. AP- single. JA and JP short. Anal brace composed of five veinal elements: AA, AA1,
CuP+AA1, CuP, CuA+CuP.
Metathoracic wings basally broad, left wing complete, 42 mm long; right wing basal third
preserved; maximum width of left wing 14.8 mm. The wing is about 2.8 times as long as broad. Anterior
margin slightly falcate. Posterior margin strongly convex from base to CuA, distal part nearly straight.
ScA+ forming a prominent recurrent subcostal brace (ScA+ScA fork+scp-ra) in leftwing. ScA+ relatively
short, forked into a ScA1+2 (only visible in leftwing) ending in costal margin well before 1/12 of wing
length, and crossvein like branch ScA3+4 (only visible in leftwing), ending on ScP-. About five crossveins
between wing margin to ScP in lefwing (not aligned with that of ScP to RA), ≈15 crossveins from ScP to
RA and ≈13 crossveins from RA to RP. ScP noticeably curved at the base ending on wing margin at the
height of IR2. RA+ separated from RP+ from the very base. RA and RP arched and divergent (0.4 mm) in
the middle of the segment from the base to MA emergence. Absence of crossvein between RA-RP.
Crossvein scp-ra aligned to ScA3+4 (preserved in right wing). RA bending to wing margin just basal to
the level of RP1-RP2 bifurcation running adjacent to wing margin. RA single. RP convex up to MA
emergence (at the level of distal half of cubital cell) (6.2 mm from the base), distally concave. RP-
forked beyond mid-wing into RP1+2 and RP3+4. RP1+2 bifurcated into RP1 and RP2. RP1 trifurcated. IR1
between RP1 and RP2, equal to RP1 biburcation. IR2+ between RP2 and RP3+4. MA forked distal of mid-
wing in three branches. MP± bending anteriorly (MP concavely curved) before and in the connection with
CuA (4.1 mm from wing base). MP connected to CuA by an extremely short crossvein. Cu- bending
anteriorly and forked into CuA and CuP forming a cubital cell. Cubital cell (vertical) developed, shorter
and broader than forewing with five elements: CuA (straight and short) (0.9 mm in both wings), CuA
crossing (curved and long) (3.7 mm in both wings), CuP crossing (curved and long) (2.7 mm in both wings),
CuP (1.5 mm in rightwing, 1.2 mm in leftwing), and cua-cup (0.2 mm in rightwing, 0.6 mm in leftwing). CuP
kink. CuA connected with CuP by a crossvein. Cubito-anal area very broad. CuA with three terminal
branches. CuP with two short terminal branches. CuP connected to AA1+2 by a short in right wing (long
in left wing) cua-aa1+2 crossvein. AA1+2 with four branches, AA3+4 with three branches. AP- single. JA
and JP single. Anal brace composed of five veinal elements: AA, AA1+2, cup+aa1+2, CuP, cua+cup.
Prothoracic wing articulation: Not well preserved but seems to have two parts, an anterior and a
posterior one.
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Mesothoracic wing articulation: Anterior articular plate (AAP), with an "h" shaped basialare (B-
C), a uni(?)-lobed fulcalare (F-C) narrowly separated from the axalare (AX-C) by a deeply incised
groove-like suture. Subcostal pteralia row free well developed. Posterior articular plate (PAP) formed by
radial, medial, and cubital pteralia. A deep groove posterior to cubital pteralia coinciding with the arising
of Cu vein. Anal and jugal veins arising obliquely at the middle height of the posterior articular plate.
Metathoracic wing articulation: Anterior articular plate (AAP), with an wide triangle shaped basialare
(B-C), a uni(?)-lobed fulcalare (F-C) with a long nose like structure, axalare (AX-C) oblong touching the
B-C and separated from the F-C by a deeply incised groove-like suture.
In forewings and hindwings, axialare of AAP connected by an arched and fine rib. Axialare of
PAP connected by an arched fine double rib.
No apparent division between pro- meso- and metathorax.
Abdomen: elongated and slender with 10 segments preserved. Female terminalia with segments
VIII-X modified and principally segments IX-X by a rounded genitalia. Segment X plate like slightly
longer than IX. Short and sickle like ovipositor between segments IX and X. Sharp and sickle valve 1,
valve 2 string along with valve 1 and valve 3 curved (at some distance from V1 and V2) and sheathing
laterally V1 and V2.
Etymology: In honour of Argentinean (twice) President (2007-2015) Cristina Fernández de Kirchner.
She elevated the Secretary of Science to Ministry rank, creating in 2007 the Ministerio de Ciencia,
Tecnología e Innovación Productiva (Ministry of Science, Technology and Productive Innovation). Within
the Ministry, the CONICET experienced a quali-quantitative leap in all fields: infrastructure, number of
workers, projects, organization and production.
Remarks: The metathoracic AAP in Argentinala gen. nov. is quite similar in shape to that interpreted
by Kukalová-Peck (2009) for the forewing of Alanympha richardsoni Kukalová-Peck, 2009 (Kukalová-
Peck, 2009: fig. 6).
Discussion: The specimen could be included into Odonatoptera because of the: (1) anterior articular
plate (AAP), with an “h” shaped basialare (B-C), a large bi-lobed fulcalare (F-C) narrowly separated from
the axalare (AX-C) by a deeply incised groove-like suture; (2) MP unbranched; (3) CuP with a kink at the
point of contact with AA; (4) archaedictyon reduced; (5) Presence of a subcostal brace; (6) presence of
an anal brace in fore- and hindwings; (7) presence of a cubital cell; (8) prothoracic winglets obliquely
directed anteriorly with venation and articulated; and (9) abdomen elongated and slender. The specimen
could be included into Palaeodonatoptera taxon nov. because of the MP shortly fussed to CuA in
forewings. It could be included into Plesiodonatoptera taxon nov. by the presence of (1) CuA-CuP fused
in forewings; (2) CuP-AA fusion in forewings; and (3) loss of crossvein between RA-RP. It could not be
included in Apodonatoptera taxon nov. because it lacks the (1) RA parallel one cell distant to anterior
wing margin (place to pterostigma); and the (2) CuA-CuP fused in hindwings; AA1+2 fused with CuP in
hindwings. The new genus and species has five autapomorphies, i.e., (1) MP touching punctually CuA in
hindwings (shortening of Crossvein (?)); (2) abdominal segment 10 somewhat longer than 9; (3) segments
9 and 10 modified with a rounded genitalia; (4) short and sickle like ovipositor; (5) sharp and sickle valve
1, valve 2 string along with valve 1 and valve 3 curved and sheathing laterally V1 and V2.
1.2.2.2. Apodonatoptera taxon nov.
Included taxa: Geroptera and Neodonatoptera taxon nov.
Synapomorphies: RA parallel one cell distant to anterior wing margin (place to pterostigma); CuA-CuP
fused in hindwings; AA1+2 fused with CuP in hindwings.
1.2.2.2.1. Order Geroptera Brodsky, 1994
Included family: Geropteridae fam. nov.
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Phylogenetic definition: Geroptera Brodsky, 1994 shall include all the Odonatoptera more closely
related to Geropteron arcuatum Riek, 1983 than to any of the type species of the other type genera of
the Apodonatoptera taxon nov. group taxa (stem-based definition).
Rediagnosis: Characters from metathoracic wing. (1) wings with relatively undeveloped anal field (more
marked in mesothoracic wing); (2) MP unbranched; (3) CuP with a kink at the point of contact with AA;
(4) archaedictyon reduced; (5) presence of a subcostal brace; (6) presence of an anal brace; (7)
presence of a cubital cell; (8) CuA-CuP fused; (9) AA1+2 fused with CuP; (10) RA parallel one cell distant
to anterior wing margin (place to pterostigma); (11) ScP long reaching the RA in the distal fourth of the
wing and related to two crossveins from RA to the wing margin (pseudopterostigma); (12) cubital cell in
hindwing formed by five elements: CuA, CuA crossing, CuP crossing, CuP+AA1+2, CuP; (13) MP linked by a
crossvein to CuA; (14) MP bending anteriorly before and in the connection (via crossvein) with CuA.
Phylogenetic systematic: The Order Geroptera retains nearly all the synapomorphies present in the
hindwings of the stem group of the Odonatoptera (Characters 1-7). Characters 8-10 are
synapomorphies of the Apodonatoptera taxon nov. Characters 11-12 are synapomorphies of the order.
Characters 13-14 are groundplan symplesiomorphies of Odonatoptera.
1.2.2.2.1.1 Geropteridae fam. nov.
Type genus: Geropteron Riek, 1983
Diagnosis: That of the order by monotypy.
Remarks: The holotype specimen MLP 12885 was collected by Sergio Archangelsky from the Malanzán
Formation, La Divisoria Member, Malanzán, Cuestita de la Herradura locality, La Rioja, Argentina.
Geropteron Riek, 1983
Type species: Geropteron arcuatum Riek, 1983
1.1.1.1. Geropteron arcuatum Riek, 1983
Type material: Holotype specimen MLP 12885. Cuestita de La Herradura locality, La Rioja province,
Northwest Argentina, at paleolatitude ≈ 60º. Malanzán Formation, Serpukhovian (circa 325-324 (?) Ma)
(Césari et al., 2011; Césari & Gutiérrez, 2001).
Remarks: As Eugeropteron, Geropteron arcuatum was named by Riek in Riek & Kukalová-Peck (1983). For
Riek & Kukalová-Peck (1983) the RP was concave (-) in all its length. See below for the consideration
that while RP is convex, MA is fused to it (same situation as for Eugeropteron in Riek & Kukalová-Peck
(1983) work).
1.2.2.2.2.1 Neodonatoptera Bechly, 1996
Included taxa: Eomeganisoptera Rohdendorf, 1962 and Euodonatoptera Bechly, Brauckmann, Zessin &
Gröning, 2001.
Synapomorphies: Wing venation characters: wings very slender and elongate [from Bechly (2007)]; RA
and RP basally strictly parallel and very close together (but only fused to a long double-barrel radial
stem in Nodialata) [from Bechly (2007)]; MP and Cu are at least shortly fused [from Riek & Kukalová-
Peck (1984), Brauckmann & Zessin (1989) and Bechly (1994)]; the longitudinal wing veins MP and CuA are
not straight but undulating or even kinked [from Bechly (2007)]; AA2 of hindwings secondarily supplied
with several pektinate posterior branches (reversed in some †"Erasipteridae" and Nodialata) [from
Bechly (2007)]; MP bending posteriorly (captured) at the fusion with CuA [proposed here].
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Remarks: The character “base of MA lost its connection with the medial stem and secondarily fused
with RP” proposed by Bechly (2007) for Neodonatoptera (and also considered by Riek & Kukalová- Peck
(1984), Brauckmann & Zessin (1989) and Bechly (1994) is actually a synapomorphy of Hydropalaeoptera.
We have to note that Sroka et al. (2014) discussed twice about this matter. In the Phylogenetic
affinities of Bojophlebia prokopi [page 13] they stated that the “MA attached to RP without anterior
connection of MA to MP” is an “autapomorphy [of Bojophlebia], convergent with Ephemerida and higher
Odonatoptera”. In page 18 (Sroka et al., 2014), they stated a synapomorphy of Hydropalaeoptera: “MA
approximated or fused to RP”. These two sentences are ambiguous and our new interpretation tries to
clarify the matter. In this sense we could note that the free RP is convex before the arising of MA and
after that becomes concave in all Odonatoptera basal to Neodonatoptera. In Bojophlebia it could be the
case, but Sroka et al. (2014) do not point the convexity / concavity of veins in their new interpretation
of the taxon. In Kukalová-Peck’s interpretation of Bojophlebia (Riek & Kukalová-Peck, 1984; Kukalová-
Peck, 1985) it is stated that the RP is concave from the very base before MA arising. We have to note
that this interpretation coincides with their erroneous interpretation of the concavity of the RP for
Eugeropteron and Geropteron (Riek & Kukalová-Peck, 1984). The only possibility is the re-study of
Bojophlebia to clear up our thoughts. Our interpretation is that RP is convex while it is fused with MA in
basal Odonatoptera, and when MA arises it becomes concave. The fusion of RP with MA is visible in the
crown group of Odonata in the anterior part of the arculus (Kukalová-Peck, 1991: figs. 6.15 I-J).
Acknowledgements
This research was supported by grants from the Consejo Nacional de Investigaciones Científicas
y Técnicas of Argentina (CONICET): PIP 0377, PIP 0834, and PIP 0585. We are indebted to the
Agencia Nacional de Promoción Científica y Tecnológica of Argentina (ANPCyT) by grant PICT-2012-
1555 for technical support. We thank the Culture Secretary of La Rioja province for permits.
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Fig. 1.- Photograph of Tupacsala niunamenos gen. nov. et sp. nov. from Guandacol 1 locality, Quebrada de las Libélulas, Cerro
Guandacol, province of La Rioja, Northwest Argentina. Lowermost part of Guandacol Formation (circa 325-324 Ma). Holotype
specimen MACN-In 2678C. Scale bar = 2 mm.
Fig. 2.- Camera lucida drawing of Holotype (MACN-In 2678C) of Tupacsala niunamenos gen. nov. et sp. nov. Scale bar = 2 mm.
Fig. 3.- Photograph of Kirchnerala treintamil gen. nov. et sp. nov. from Guandacol 1 locality, Quebrada de las Libélulas, Cerro
Guandacol, province of La Rioja, Northwest Argentina. Lowermost part of Guandacol Formation (circa 325-324 Ma). Holotype
specimen MACN-In 2678D. Scale bar = 3 mm.
Fig. 4.- Camera lucida drawing of Holotype (MACN-In 2678D) of Kirchnerala treintamil gen. nov. et sp. nov. Scale bar = 3 mm.
1
2
3
4
Petrulevičius & Gutiérrez (2016): New basal Odonatoptera from the lower Carboniferous (Serpukhovian) of Argentina.
358
Fig. 5.- Photograph of Argentinala cristinae gen. nov. et sp. nov. from Guandacol 1 locality, Quebrada de las Libélulas, Cerro
Guandacol, province of La Rioja, Northwest Argentina. Lowermost part of Guandacol Formation (circa 325-324 Ma). Holotype
specimen MACN-In 2678B. Scale bar = 5 mm.
Fig. 6.- Composite camera lucida drawing of Holotype (MACN-In 2678A-B) of Argentinala cristinae gen. nov. et sp. nov. Scale
bar = 5 mm.
5
6
