The first Triassic Diptera (Insecta) from South America, with review of Hennigmatidae.

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ZOOTAXA

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Zootaxa 3710 (1): 081–092

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http://dx.doi.org/10.11646/zootaxa.3710.1.6

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The first Triassic dipteran (Insecta) from South America,

with review of Hennigmatidae

MARÍA BELÉN LARA1 & ELENA D. LUKASHEVICH 2,3

1Area Entomología, Universidad Nacional del Nordeste and Area Paleontología, Centro de Ecología Aplicada del Litoral

(CONICET), Casilla de Correo 128, 3400 Corrientes, Argentina. E-mail: [email protected] Paleontological Institue RAS, Moscow, Russia. E-mail: [email protected] author

Abstract

The first Triassic dipteran from South America is described based on an isolated wing from the lower Upper Triassic de-

posits of Argentina (Mendoza Province, Potrerillos Formation, Quebrada del Durazno locality). Trihennigma zavattierii

gen. et sp. nov. is a member of the Mesozoic family Hennigmatidae, previously recorded only from Eurasia. A key for

the genera and species of Hennigmatidae is provided and systematic position of the taxa is discussed.

Key words: Diptera, new genus, Upper Triassic, Argentina, basiala, SEM

Introduction

The Diptera, or true flies, are known since the early Middle Triassic (Anisian) Buntsandstein of Spain (Calafat Colom 1988; Zessin 2008), France (Grès à Voltzia Formation; Krzemiński et al. 1994), and Germany (Röt Formation; Bashkuev et al. 2012). The oldest assemblage of Diptera from France is very diverse: both suborders (Nematocera and Brachycera) and four infraorders of nematocerans (Tipulomorpha, Psychodomorpha sensu

Hennig, Culicomorpha, and Bibionomorpha) have been described based on adults and immatures (Krzemiński & Krzemińska 2003; Lukashevich et al. 2010).

Later in the Triassic, dipterans became more diverse and widespread: impressions of other Triassic Diptera have been described from the Ladinian, Ladinian–Carnian, and Norian–Rhaetian of Asia (Kovalev 1983a; Shcherbakov et al. 1995; Hong & Guo 2003), Carnian of Australia (Kovalev 1983b; Blagoderov 1999; Lukashevich & Shcherbakov 1999) and North America (Krzemiński 1992; Blagoderov et al. 2007), Rhaetian of Europe (Krzemiński & Jarzembowski 1999; Krzemiński & Krzemińska 2002); and also recorded but not yet described from the Norian of Europe (Barth et al. 2011) and Carnian of Africa (Anderson & Anderson 1993). Additionally, the first nematoceran inclusion has been recently found in the Late Carnian amber of northeastern Italy, but it cannot be formally described due to partial preservation (Schmidt et al. 2012). Up to now, 31 genera with 44 species of Triassic dipterans have been described (listed in Blagoderov et al. 2007, updated by Lukashevich et al. 2010); most of them found in the Northern Hemisphere. Only four monotypic genera of Triassic Diptera, based on four isolated wings, have been described from the Southern Hemisphere, all from the Mount Crosby Formation, Queensland, Australia (Crosaphis anomala Evans, 1971, Tillyardiptera prima Lukashevich et

Shcherbakov, 1999, Austrocramptonomyia minuta Blagoderov, 1999 and Veriplecia handlirschi Blagoderov, 1999).

Although the rich Triassic entomofauna of Argentina has been studied for a long time, yielding 12 orders recorded and 81 species described (Brauckmann et al. 2010; Gallego et al. 2011; Martins-Neto et al. 2011; Lara et

al. 2012), until recently South America was one of only two continents (the other one being Antarctica) with no records of Triassic Diptera. The first insects recorded from the Triassic of South America (Argentina) have been described as crane flies, Tipuloidea rhaetica Wieland, 1925 and Tipulodites affinis Wieland, 1925; however now

Accepted by V. Blagoderov: 29 Aug. 2013; published: 11 Sept. 2013 81


they are considered to be members of Homoptera and Mecoptera, respectively (Tillyard 1926; Martins-Neto et al.

2003; Bashkuev pers. comm.).Only recently the first Triassic member of Diptera was found in South America. This specimen was collected

from the upper section of the Potrerillos Formation (lower Upper Triassic) at the Quebrada del Durazno locality (33º04´90''S; 68º07´13''W), south of Cerro Cacheuta, Mendoza Province, Argentina (for further details on the locality, see Martins-Neto & Gallego 1999; Martins-Neto et al. 2008). More than two hundred insect specimens have been collected in the Potrerillos Formation, and 21 species have already been described (see Lara et al. 2012), eight of them from the Quebrada del Durazno locality: Argentinocicada magna Martins-Neto et Gallego, 1999, Argentinocicada minima Martins-Neto et Gallego, 1999, Cacheutacicada kurtzae Martins-Neto et Gallego, 2008 and Gallegomorphoptila acostai (Martins-Neto et Gallego, 1999), and Potrerillia nervosa Martins-Neto et

Gallego, 1999 (Hemiptera), Miomina mendozina Martins-Neto et Gallego, 1999 (Miomoptera), Argentinosyne

duraznoensis Martins-Neto et Gallego, 2008 (Coleoptera: Schizocoleidae), and Notopamphagopsis? sp. 1 (Orthoptera: Haglidae). A photo of the specimen described herein has been published by Lara et al. (2012, Fig. 6k) with the caption “indet. material.”

Material and methods

The specimen MCNAM-PI 24607 described below is housed in the collection of the Museo de Ciencias Naturales y Antropológicas “Juan Cornelio Moyano” (Mendoza city). Photographs were taken using an Olympus SZ51 stereomicroscope with an Olympus SP-350 digital camera (8.0 megapixels), and a Leica M1 65C stereomicroscope equipped with a Leica DFC425 digital camera, with subsequent adjustments made using the Adobe Photoshop® CS 10.0 software. Scanning electron micrographs of the uncoated holotype of Metatrichopteridium cladistorum

were taken on a Tescan Vega XMU instrument using a secondary electron (SE) detector.The classification of Nematocera accepted here follows that of Shcherbakov et al. 1995 and Blagoderov et al.

2002. The vein nomenclature mostly follows Wootton & Ennos 1989 (the vein traditionally named 1A is, in their opinion, CuP), as modified by Shcherbakov et al. 1995.

Systematics

Infraorder Psychodomorpha sensu Hennig, 1968

Superfamily Hennigmatoidea Shcherbakov in Shcherbakov, Lukashevich et Blagoderov, 1995

Family Hennigmatidae Shcherbakov in Shcherbakov, Lukashevich et Blagoderov, 1995

The family, comprising two subfamilies, Kuperwoodiinae and Hennigmatinae, has been established based on three isolated wings from the Triassic and Cretaceous of Asia (Shcherbakov et al. 1995). The subfamily Kuperwoodinae, established based on a single wing from the Ladinian or Carnian of Kyrgyzstan, so far remains unknown elsewhere, whereas the nominative subfamily appears to be rare but widespread in the Mesozoic. The type genus Hennigma Shcherbakov, 1995 was described from the Early Cretaceous of Mongolia. Later, Ansorge (2001) synonymized it under Metatrichopteridium Handlirsch, 1939 from the Early Jurassic (Early Toarcian) of Germany, previously erroneously referred to Trichoptera (Handlirsch 1939). Another genus of this subfamily, Daohennigma

Lukashevich, Huang et Lin, 2006, close in its wing venation to Metatrichopteridium, was described from the Middle or Late Jurassic of China based on a single body fossil (Lukashevich et al. 2006). The third genus of this subfamily, Anemeca Shcherbakov, 1995—of debatable systematic position—is known from the Triassic (Ladinian or Carnian) of Kyrgyzstan (see Discussion).

The species described below can be assigned to Hennigmatidae based on the following characters diagnostic of the family: wing with main forks and crossveins more or less shifted to base; Sc apex beyond wing midlength; R base up to MA weak; Rs originating very early, close to MA; R

2 free; R

2+3+4 aligned with Rs stem, forking

symmetrically into R2+3

and R4; R

5 separated before wing midlength, earlier than R

4;

alular incision distinct and anal

LARA & LUKASHEVICH82 · Zootaxa 3710 (1) © 2013 Magnolia Press


lobe prominent. Another character, included in the original diagnosis of the family as “strong convex M base aligned with CuA” (Shcherbakov et al. 1995), should be amended as “CuA continued proximal to MA as strong convex CuA base,” because the actual M base, more or less aligned with M

3+4, is clearly visible in the new genus

and Anemeca and is also traceable in M. cladistorum (Figs 1, 2, 5, 6). The strength of the CuA base, contrasting to the relative weakness of the R base remains one of the most important diagnostic characters of Hennigmatidae.

FIGURE 1. Wing venation of Hennigmatidae (holotypes): A, Kuperwoodia benefica Lukashevich, 1995; B, Anemeca liya

Shcherbakov, 1995; С, Metatrichopteridium cladistorum (Shcherbakov, 1995); D, Daohennigma panops Lukashevich et al.,

2006; modified after Shcherbakov et al., 1995 (A–C) and Lukashevich et al., 2006 (D).

Key to genera of Hennigmatidae

1 (2) Nodal flexion line (with R2+3+4

bend) and pterostigma distinct, M stem as long as d cell, CuA bent at m-cu and then strongly

diverging from M4, m-cu before wing midlength (Figs 1A, 2A) . . . . . . . . . . . . . . . . . . . . . Kuperwoodiinae Lukashevich, 1995

1 sp., Kuperwoodia benefica Lukashevich in Shcherbakov et al., 1995; Kyrgyzstan, Dzhailoucho, Madygen Fm., T2-3

2 (1) Nodal flexion line untraceable, pterostigma at most vestigial, M stem much shorter than d cell, CuA straight at m-cu and

curved backwards before apex, m-cu beyond wing midlength . . . . . . . . . . . . . . . . . . . . . . . . Hennigmatinae Shcherbakov, 1995

3 (4) Wing elongate with anal lobe reduced, bifurcation M1+2

proximal to bifurcation M3+4

(Figs 1B, 2B) . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Anemeca Shcherbakov, 1995

1 sp., A. liya Shcherbakov, 1995; Kyrgyzstan, Dzhailoucho, Madygen Fm., T2-3

4 (3) Wing broad, with anal lobe produced, bifurcation M1+2

distal to bifurcation M3+4

5 (10) R2 shorter than R

2+3, anal loop narrow (unknown for Daohennigma)

6 (7) Pterostigma vestigial, supernumerary crossvein r2+3-r4 and spots on crossveins present (Figs 1D, 3B). . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Daohennigma Lukashevich, Huang et Lin, 2006

1 sp., D. panops Lukashevich, Huang et Lin, 2006; China, Daohugou, J2-3

7 (6) Pterostigma, supernumerary crossvein r2+3-r4, and spots on crossveins absent. . . . . . . Metatrichopteridium Handlirsch, 1939

8 (9) R2 slightly shorter than R

2+3. . . . . . . . M. confusum Handlirsch, 1939; Germany, Dobbertin, Braunschweig, Lower Toarcian, J

1

9 (8) R2 twice shorter than R

2+3 (Figs 1C, 3A) . . M. cladistorum (Shcherbakov, 1995); Mongolia, Khutel-Khara, Tsagantsab Fm., K

1

10 (5) R2 longer than R

2+3, anal loop broad (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Trihennigma gen. nov.

1 sp. T. zavattierii sp. nov.; Argentina, Quebrada del Durazno, Potrerillos Fm., T3

Subfamily Hennigmatinae Shcherbakov in Shcherbakov, Lukashevich et Blagoderov, 1995

The new species can be assigned to Hennigmatinae based on the following characters diagnostic of the subfamily: first radial space of usual width; M stem extremely short (so that the base of the d cell is close to MA); CuA straight at m-cu and curved backwards before apex; m-cu beyond wing midlength. Although the median wing area,

Zootaxa 3710 (1) © 2013 Magnolia Press · 83TRIASSIC DIPTERAN FROM SOUTH AMERICA


including im, is not preserved, the presence of a very long d cell, typical of the subfamily, can be confidently hypothesized.

FIGURE 2. A, Kuperwoodia benefica Lukashevich, 1995: holotype PIN 2344/490; B, Anemeca liya Shcherbakov, 1995:

holotype PIN 2069/3354.

Trihennigma gen. nov.

Type species. Trihennigma zavattierii sp. nov.Differential diagnosis. Wing rather broad, with vestigial pterostigma and without spots at crossveins; long

strong Sc ending just before R2+3

bifurcation and possibly distal to M1+2

bifurcation; postsectoral sc-r probably

nearly level with R5 origin; supernumerary crossveins r-rs, r3-r4, r4-r5 and r5-m1; R

2 longer than R

2+3, R

4 almost as

long as half of wing, entering margin at wing tip; im not preserved, m-cu just before M3+4

fork; M4 curved

backwards before apex; CuA section before m-cu twice as long as section beyond it; anal loop broad, subequal to 1A beyond it, the latter terminating at deep alular incision, anal lobe possibly produced.



FIGURE 3. A, Metatrichopteridium cladistorum (Shcherbakov, 1995): holotype PIN 3965/2789; B, Daohennigma panops

Lukashevich et al., 2006: holotype NIGPAS 133706.



FIGURE 4. Trihennigma zavattierii gen. et sp. nov.: holotype MCNAM-PI 24607: A, photo, B, drawing, C, reconstruction.

Scale bar 1.7 mm. Abbreviation: ai—alular incision.



FIGURE 5. Basiala of Hennigmatinae: A, B, Trihennigma zavattierii gen. et sp. nov.: holotype MCNAM-PI 24607, part and

inverted counterpart; C, Metatrichopteridium cladistorum (Shcherbakov, 1995): holotype PIN 3965/2789. Abbreviation: ai—

alular incision.



FIGURE 6. Proximal part of the wing of Metatrichopteridium cladistorum (Shcherbakov, 1995): holotype PIN 3965/2789: A,

drawing; B, SEM image (SE). Abbreviation: ai—alular incision.

FIGURE 7. Distribution map of Hennigmatidae.

Species included. Monobasic.Etymology. From the Triassic period and the genus name Hennigma.Remarks. Combination of long R

2+3 fork, curved M

4 and broad anal loop has been known within the family

only in the monobasic superfamily Kuperwoodiinae. The new genus resembles Metatrichopteridium and Daohennigma in the long Sc and extremely short M stem (the correspondingly long d cell is hypothesized), and the positions of R

4 (ending at wing tip) and m-cu (just before M

3+4 bifurcation). Such strong Sc, as well as 1A

terminating at a deep alular incision, has been recorded in both species of Metatrichopteridium (basiala is not seen



on the single specimen of Daohennigma); a similar set of supernumerary crossveins is known in Daohennigma. The wing under description is distinguishable from Daohennigma by the more distal position of unaligned r3-r4

and r4-r5, and the absence of spots on crossveins; in fact, these may be species-level differences. We agree with Ansorge (2001) that the subcostal veinlets mentioned in the original description of Hennigma

are artifacts of preservation and are absent in the holotype of M. cladistorum, as well as in other hennigmatids (Figs 1, 5, 6). However, his redrawing of the wing base is incorrect (Ansorge 2001: Fig. 3c): cu-a is certainly preserved as it was shown in Shcherbakov et al. (1995: Fig. 33) and can be seen in Trihennigma gen. nov.; moreover, in both genera, 2A is not marginal and a small alula is visible in M. cladistorum and is likely to be present in the new genus (Figs 5, 6).

Trihennigma zavattierii sp. nov.

(Figs 4, 5A, B)

Type material. Holotype: MCNAM-PI 24607 (part and counterpart of partly preserved isolated wing). Quebrada del Durazno, southern flank of Cerro Cacheuta, southern end of the Precordillera, Cuyo Basin, Mendoza Province, Argentina; upper section of the Potrerillos Formation; early Late Triassic (equivalent to the Carnian of marine sequences).

Etymology. In honour of the geologist Dr. Ana M. Zavattieri, for her great contributions to the Triassic geology and palaeontology of Argentina.

Description. Wing infuscate. Sc and Rs long, about 70% and 80 % of wing length, respectively; R2 shifted to

R1; R

2+3 divided by r-rs as 1:2; R

4 divided by r3-r4 as 6:5.5 and by r4-r5 as 7:4.

Measurements (mm). Wing length 8.6 long, 2.9 wide (3.3 mm estimated).

Discussion

As the first member of Hennigmatidae found on the Gondwana continent (Fig. 7) and the first record of Triassic Diptera from South America, Trihennigma zavattierii gen et sp. nov. provides important evidence of the wide distribution and diversity of Diptera already in the Late Triassic. This discovery is a great fortune (in most Triassic localities, Diptera occur as single records per thousands of other insects), but not a great surprise. Not only members of the same family have been found in the Triassic deposits of Laurasia and Gondwana (Procramptonomyiidae and Paraxymyiidae from Australia and North America), but even members of the same genus have been described from both ancient supercontinents: Veriplecia handlirschi (Paraxymiidae) from the Carnian of Australia (Mt. Crosby, Mount Crosby Formation; Blagoderov 1999) and V. rugosa Blagoderov et

Grimaldi, 2007 from the Late Carnian of North America (Solite, Cow Branch Formation; Blagoderov et al. 2007).On basiala structure. According to Shcherbakov et al. (1995), the anal veins forming a loop is characteristic

of all non-tipulomorphan Diptera, and in some generalized groups the 1A is traceable distal to the loop up to the alular incision, usually as a colourless channel vein. Recently, the structure of basiala (or wing stalk of authors) was re-interpreted by Stary (2008), who suggested that the true alular incision arises close to the level of the humeral cross-vein, and the longitudinal cord reaching the alular incision is not a rudiment of an anal vein, as Hennig (1968) supposed, but the alular impression—an evolutionary novelty correlated with the development of alula.

On wings of both Metatrichopteridium and Trihennigma, the alula and alular incision are present, and the latter is situated just at the level of the humeral vein (Figs 5, 6; compare to Hennig 1968: Fig. 7). So, even according to Stary, it is a true alular incision. However, a true vein, pigmented, sclerotized, and without any break beyond anal loop—and thus undoubtedly not an “alular impression”—is clearly visible up to this incision. It is worth to mention that the shape of the considerably protruding anal lobe and the incision with callus in Mesozoic Metatrichopteridium are very similar to those in at least some Blephariceridae, which have no true alular incision according to Stary, because this angle at the base of the anal lobe is situated markedly beyond the humeral vein.

The discovery of perfectly preserved basiala of Trihennigma contradicts the interpretation by Stary and, therefore, there is no reason to alter our opinion on the reduction of the distal 1A.

On the systematic position of Kuperwoodia and Anemeca. Some authors disagreed with the placement of



these genera in Hennigmatidae and suggested alternative classifications (Krzemiński & Krzemińska 2003; Grimaldi & Engel 2005). Our counterarguments were published earlier (Lukashevich et al. 2006). The phylogenetic analysis in Blagoderov et al. (2007) recovered Anemeca, Kuperwoodia, and Metatrichopteridium

close to each other on the Psychodomorpha branch of their strict consensus phylogeny of Diptera, and the authors considered these taxa as members of one family.

The assignment of Kuperwoodinae and Hennigmatinae to the same family was based, first of all, on the base of R being weaker than the very strong CuA base, and the following combination of characters, unknown in other Diptera: early separation of both Rs and R

5, free R

2, long Sc, not terminal sc-r, anal loop and anal lobe present (for

further details, see Shcherbakov et al. 1995). The time gap between Kuperwoodia (T2-3

) and other hennigmatids,

initially represented by only Hennigma (K1), decreased considerably after Early Jurassic M. confusum was placed

in that family (Ansorge 2001). The new Triassic record from Argentina closes that gap altogether. The proposal to raise Kuperwoodinae, based on a single wing, to the family level due to “numerous striking differences in shape and venation of wing” without any discussion (Krzemiński & Krzemińska 2003) seems unjustified. The genera of Hennigmatinae described later (Daohennigma and Trihennigma gen. nov.) display main characters of the subfamily (such as the extremely short M stem and long d cell), as well as some characters previously recorded only in Kuperwoodia (pterostigma, long fork R

2+3, curved M

4, broad anal loop).

In spite of the typical strong CuA base, the systematic position of Anemeca is much more disputable due to several characters unknown in other Hennigmatidae, such as the absence of the anal lobe and the fork M

1+2 being

longer than M3+4

. However, unlike Krzemiński and Krzemińska (2003), we consider Anemeca as a certain member

of Diptera: it cannot be a member of Trichoptera (e.g., due to a clearly visible narrow base of its wing and the unforked CuA; V. D. Ivanov, pers. comm. 2012) or Mecoptera (e.g., due to a very proximal forking of its M stem; A.S. Bashkuev, pers. comm. 2013).

Because both Anemeca and Kuperwoodia are known only from single poorly preserved wings, further discussion can be fruitful only after additional material is discovered.

Acknowledgements

We are deeply indebted to D.E. Shcherbakov (Borissiak Paleontological Institute RAS, Moscow) for valuable discussion and assistance in taking digital images, R.A. Rakitov (same institution) for assistance in taking scanning electron images and improving the English of the manuscript, and A.S. Bashkuev (same institution) and V.D. Ivanov (St. Petersburg State University) for discussion of Anemeca. We thank J. Ansorge and W. Krzemiński for comments on the early version of the manuscript.

The study by M.B. Lara was supported by Secretaría General de Ciencia y Técnica, Universidad Nacional del Nordeste (PI-2010/F022) and Agencia Nacional de Promoción Científica y Tecnológica (ANPCyT-Argentina) and Universidad Nacional del Nordeste (Grant PICTO-UNNE 0226). The research by E.D. Lukashevich was partly supported by a grant from the Russian Foundation for Basic Research (no. 13-04-01839).

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The first Triassic Diptera (Insecta) from South America, with review of Hennigmatidae.

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