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Ameghiniana

versión On-line ISSN 1851-8044

Ameghiniana v.43 n.3 Buenos Aires jul./sep. 2006

 

The Triassic insect fauna from Argentina. Coleoptera from the Los Rastros Formation (Bermejo Basin), La Rioja Province

Rafael Gioia Martins-Neto1, Oscar F. Gallego2 and Adriana C. Mancuso3

1 Programa de Pós-graduação em Ciências Biológicas, Comportamento e Biologia Animal, Universidade Federal de Juiz de Fora-UFJF. Sociedade Brasileira de Paleoartropodologia- SBPr/Campus Universitário-Martelos. 36036-900, Juiz de Fora, MG, Brasil. martinsneto@terra.com.br
2 Facultad de Ciencias Exactas y Naturales y Agrimensura, Universidad Nacional del Nordeste y Área Paleontología (Centro de Ecología Aplicada del Litoral-CONICET), CC 128, 3400 Corrientes, Argentina. ofgallego@hotmail.com
3 Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) and Departamento de Geología y Paleontología del IANIGLA-CRICYT/CONICET, Av. Adrián Ruiz Leal s/n, Parque Gral. San Martín, 5500 Mendoza, Argentina. amancu@gl.fcen.uba.ar

Abstract. New fossil insect taxa from the Los Rastros Formation (early Late Triassic), La Rioja Province (Argentina) are described. Four new genera and fourteen new species of the order Coleoptera are proposed: Ademosyne arcucciae Martins-Neto and Gallego sp. nov., Ademosyne elongatus Martins-Neto and Gallego sp. nov., Ademosyne hexacostata Martins-Neto and Gallego sp. nov., Ademosyne punctuada Martins-Neto and Gallego sp. nov., Argentinosyne frenguellii Martins-Neto and Gallego gen. et sp. nov., Argentinosyne gonaldiae Martins-Neto and Gallego sp. nov., Argentinosyne gualoensis Martins-Neto and Gallego sp. nov., Argentinosyne rugosa Martins-Neto and Gallego sp. nov., Argentinosyne losrastrosensis Martins-Neto and Gallego sp. nov., (Permosynidae Tillyard), Argentinocupes pulcher Martins-Neto and Gallego gen. et sp. nov., Argentinocupes abdalai Martins-Neto and Gallego sp. nov. (Cupedidae Lacordaire), Gemelina triangularis Martins-Neto and Gallego gen. et sp. nov., Cardiosyne obesa Martins-Neto and Gallego gen. et sp. nov., and Cardiosyne elegans Martins-Neto and Gallego sp. nov. (Elateridae? Leach). Additionally new specimens assigned to Argentinosyne rugosa Martins-Neto and Gallego sp. nov. and Argentinosyne gonaldiae Martins-Neto and Gallego sp. nov., from the Cacheuta Formation (early Late Triassic), Uspallata, Mendoza Province (a new locality for insects) are included in this paper.

Resumen. La fauna de insectos triásicos de la Argentina. Coleoptera de la Formación Los Rastros (Cuenca del Bermejo), Provincia de la Rioja . En esta contribución se describen nuevos taxones de insectos fósiles provenientes de la Formación Los Rastros (Triásico Tardío bajo), provincia de La Rioja, Argentina. Se proponen cuatro géneros y catorce especies nuevas pertenecientes al orden Coleoptera: Ademosyne arcucciae Martins-Neto y Gallego sp. nov., Ademosyne elongatus Martins-Neto y Gallego sp. nov., Ademosyne hexacostata Martins-Neto y Gallego sp. nov., Ademosyne punctuada Martins-Neto y Gallego sp. nov., Argentinosyne frenguellii Martins-Neto y Gallego gen. et sp. nov., Argentinosyne gonaldiae Martins-Neto y Gallego sp. nov., Argentinosyne gualoensis Martins-Neto y Gallego sp. nov., Argentinosyne rugosa Martins- Neto y Gallego sp. nov., Argentinosyne losrastrosensis Martins-Neto y Gallego sp. nov, (Permosynidae Tillyard), Argentinocupes pulcher Martins-Neto y Gallego gen. et sp. nov., Argentinocupes abdalai Martins- Neto y Gallego sp. nov. (Cupedidae Lacordaire), Gemelina triangularis Martins-Neto y Gallego gen. et sp. nov., Cardiosyne obesa Martins-Neto y Gallego gen. et sp. nov., y Cardiosyne elegans Martins-Neto y Gallego sp. nov. (Elateridae? Leach). Adicionalmente, se incluyen en este trabajo nuevos especímenes asignados a Argentinosyne rugosa Martins-Neto y Gallego sp. nov. y Argentinosyne gonaldiae Martins-Neto y Gallego sp. nov., provenientes de la Formación Cacheuta (Triásico Tardío bajo), Uspallata, provincia de Mendoza (una nueva localidad para insectos).

Key words. Triassic insects; Coleoptera; Los Rastros Formation; Argentina.
Palabras clave. Insectos triásicos; Coleoptera; Formación Los Rastros; Argentina.

Introduction

The present contribution on Triassic insect faunas from Argentina describes fossil insects of the order Coleoptera from the Los Rastros Formation (Río Gualo, Picos Gemelos, Agua Escondida, and quebrada de Ischichuca Chica localities, La Rioja Province) (figure 1). Additionally, two species from the Cacheuta Formation (Cuyo Basin) at Paramillos de Uspallata (Mendoza Province, Argentina) are recorded, the first unquestionable insects from this formation (see Gallego, 1997). The history of research and the previously described species were discussed by Gallego (1997), Gallego and Martins- Neto (1999), Martins-Neto and Gallego (1999, 2001) and Martins-Neto et al. (2003, 2005).

Figure 1. A, Geologic map of the Bermejo Basin (modified from Stipanicic and Bonaparte, 1979 and Kokogian et al., 2001) / mapa geológico de la Cuenca del Bermejo (modificado de Stipanicic y Bonaparte, 1979 y Kokogian et al., 2001). B, Schematic section of the Los Rastros Formation indicating fossiliferous horizons / sección esquemática de la Formación Los Rastros indicando los horizontes fosilíferos.

The record of fossil coleopterans in the Argentinean Triassic includes studies by Frenguelli (1934, 1948) and Fossa Mancini (1941). Frenguelli (1948) mentioned the presence of coleopteran elytra in the Ischigualasto Formation (San Juan Province), but these levels probably correspond to the Los Rastros Formation (sensu Stipanicic and Bonaparte, 1979). In a recent field trip to the Ischigualasto park (San Juan Province), Gallego et al. (2004) found specimens probably assignable to the new genera Argentinosyne and Argentinocupes from the Los Rastros Formation. The only west Gondwanan coleopteran records are from the Jurassic of Antarctica (Zeuner, 1959; Tasch, 1973). Gallego and Covacevich (1998) mentioned the unique occurrences of fossil insects (probably coleopterans) in the Triassic of Punta Puquén and Los Molles areas in Chile (Fuenzalida, 1937; Cecioni and Westermann, 1968). Additional records referable to Ademosyne hexacostata sp. nov. (Permosynidae) came from the Upper Triassic of Bio-Bío river from southern central Chile (Gallego et al., 2005) and from the Ischichuca Formation (late Middle Triassic - early Late Triassic) from La Rioja Province, Argentina.
The stratigraphic and geographic sources (figure 1) of these paleoentomofaunas are well documented in the Triassic literature and were briefly treated in other papers by the authors (Martins Neto et al., 2003, 2005). The coleopterans specimens came from the lake facies assemblages of the first to fifth sedimentary cycles from the Los Rastros Formation, exposed at the localities mentioned above (figure 1).
The samples from the Los Rastros Formation were collected by OFG, A. Arcucci, C. Forster, C. May and R. Rogers during 1995 and by ACM, A. Arcucci and C. Marsicano in 1999, and from the Cacheuta Formation by OFG and R. Herbst during 1995.
The terminology adopted here conforms mainly that of Ponomarenko (1969).
The repository and institutional abbreviations used here are as follows. PULR-I: Paleontology Collection, Museo de Ciencias Naturales, Universidad Nacional de La Rioja, La Rioja, Argentina. CTES-PZ: Paleozoological Collection, Facultad de Ciencias Exactas y Naturales y Agrimensura, Universidad Nacional del Nordeste, Corrientes, Argentina. MCNAM- PI: Museo de Ciencias Naturales y Antropológicas "Cornelio Moyano", Mendoza, Argentina.

Systematic paleontology (by Martins-Neto and Gallego)

Order COLEOPTERA Linnaeus, 1758
Family PERMOSYNIDAE Tillyard, 1924 (including Ademosynidae Ponomarenko, 1969)

Genus Ademosyne Handlirsch, 1906

Type species. Ademosyne major Handlirsch, 1906, Pl. 39, Fig. 14, from the Australian Triassic.

Ademosyne arcucciae Martins-Neto and Gallego sp. nov. Figures 2.A1-A10, 4.A


Figure 2. A1-A10, Ademosyne arcucciae Martins-Neto and Gallego sp. nov. A1, holotype / holotipo, PULR-I 298. A2-A10, paratypes / paratipos, PULR-I 224a, b (A2, A6), PULR-I 302 (A8), PULR-I 300a, b, c, d (A4, A5, A9, A10), PULR-I 301a, b (A3, A7). B1-B6, Ademosyne hexacostata Martins-Neto and Gallego sp. nov. B1, holotype/ holotipo, PULR-I 305. B2-B6, paratypes/ paratipos, PULR-I 308a-e. C1-C6, Ademosyne punctuada Martins-Neto and Gallego sp. nov., C1, holotype / holotipo, PULR-I 309. C2-C6, paratypes / paratipos, PULR-I 313a-e. D1-D2, Ademosyne elongatus Martins-Neto and Gallego sp. nov. D1, paratype / paratipo, PULR-I 304. D2, holotype / holotipo, PULR-I 303. E1-E5, Argentinosyne gualoensis Martins-Neto and Gallego sp. nov. E1-E4, paratypes / paratipos, PULR-I 311a-d. E5, holotype / holotipo, PULR-I 320. F, Argentinosyne rugosa Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 319. G, Schematic drawing of the coleopteran elytron, showing the main diagnostic characters used in the text / dibujo esquemático del élitro de un coleóptero, mostrando los principales caracteres diagnósticos usados en el texto. Scale bar / Escala = 1 mm. H, Paleogeography of known Ademosyne species (map adapted from Czerkas and Czerkas, 1990) / paleogeografía de las especies de Ademosyne (mapa adaptado de Czerkas y Czerkas, 1990).

1997. Ademosyne sp. Gallego, p. 513, lám. I, fig. G.
1999. Ademosyne arcucciae Gallego and Martins-Neto (nomen nudum), p. 88, Fig. 7.

Etymology. Dedicated to Andrea Arcucci for her great contribution to the knowledge of the Los Rastros fossil biota.

Material. Holotype: PULR-I 298 (figures 2.A1, 4.A). Paratypes: from Picos Gemelos (5th cycle), fifty-five specimens, PULR-I 300a, b, c, d (figures 2.A4, A5, A9, A10), CTES-PZ 7299; from Agua Escondida (4th cycle) eleven specimens, PULR-I 301a, b (figures 2.A3, A7); from Río Gualo (2nd, 3rd and 5th cycle) twenty-nine specimens, PULR-I 224a, b (figures 2.A2, A6), PULR-I 302a (figure 2.A8); from quebrada Ischichuca Chica (1st and 4th cycle) two specimens, PULR-I 306, (see table I).

Type locality. Picos Gemelos, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytra length varying from 1.4 to 9.3 mm (media 3.9, standard deviation +/- 0.4) and width varying from 0.3 to 3.2 mm (media 1.52, standard deviation +/- 0.12). Length/width ratio varying from 1.4 to 3.6 (media 2.47, standard deviation +/- 0.28). Elytron with nine costae. Costae ornamentation smooth. Space between costae bears granulated ornamentation.
Description of the holotype (figures 2.A1, 4.A). Elytron 9.25 mm long and 3.02 mm wide (L/W relation = 3), with narrow lateral border. Nine smooth, non-convergent costae. Space between costae bearing small granules homogeneously distributed over the whole elytron surface.
Discussion. In the original diagnosis of Ademosyne by Dunstan (1923), the elytra are costate without any abnormally wide borders, while the number of costae or intervals, together with size, shape, and other features provide specific differences. The number of costae or intervals varies from eight to eleven, with nine generally present, and a variation in length of elytra between 1.8 mm and 6.2 mm. Ademosyne arcucciae sp. nov. has all the generic characters mentioned above. Its length range attains around 9.5 mm, and the smallest size is around 1.5 mm, differing from all the Ademosyne species proposed by Handlirsch (1906), Tillyard (1916) and Dunstan (1923), but not enough for the erection of a new genus. In this respect, Ademosyne arcucciae sp. nov. is more similar to Ademosyne speciosa Riek (1974), from the Molteno Formation (South Africa), but differing in having more conspicuous costae (faint in Ademosyne speciosa). Ademosyne curvata Dunstan (1923) is an Australian species with a more similar elytra shape.

Ademosyne elongatus Martins-Neto and Gallego sp. nov. Figures 2.D1-D2, 4.B

Etymology. From Latin, elongatus, elongated.

Material. Holotype: PULR-I 303 (figures 2.D1, 4.B). Paratypes: PULR-I 304a-b (figure 2.D1), two specimens from Picos Gemelos (5th cycle).

Type locality. Gualo, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation. Age. Early Late Triassic.
Diagnosis. Elytron subelliptical, elongated in shape with length between 6 mm and 9 mm, width between 2 mm and 3 mm. Length/width ratio around 3. Eleven smooth, non-convergent costae. Space between costae bearing small square granules homogeneously distributed over the whole elytron surface.
Description of the holotype (figures 2.D2, 4.B). Elytron subelliptical elongated in shape with the proximal margin similar to the distal one, both semicircular, 9 mm long and 2.8 mm wide (L/W relation = 3.2, see table 1). Lateral border narrow. Eleven smooth costae. Ornamentation between costae consisting of small square granules homogeneously distributed over the whole elytron surface.
Discussion. Ademosyne elongatus sp. nov. differs from the previously described species by having eleven costae (nine in A. arcucciae) and a more elongated elytra (around three times longer than wide in A. elongatus and around two and a half times in A. arcucciae). A. elongatus slightly resembles Simmondsia cylindrica Dunstan (1923) from the Triassic of Australia, in general outline and square ornamentation between costae, but differs in the number of costae (eight in S. cylindrica), well marked scutellary margin, more conspicuous and strong ornamentation and less length/width ratio.

Ademosyne hexacostata Martins-Neto and Gallego sp. nov. Figures 2.B1-B6, 4.C

Etymology. Refers to the number (six) of costae.

Material. Holotype: PULR-I 305 (figures 2.B1, 4.C). Paratypes: from Pico Gemelos (5th cycle), three specimens, PULR-I 307, from Río Gualo (5th cycle), twelve specimens (PULR-I 308a-e, figures 2.B2 - B6).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron elliptical, elongated in shape with length varying between 1.1 and 9 mm (media 2.7, standard deviation +/- 1.31), width varying between 0.33 and 3.3 mm (media 0.99, standard deviation +/- 0.5). Length/width ratio varying between 2.2 and 3.5 (media 2.72, standard deviation +/- 0.33). Six smooth, subparallel costae. Space between costae bearing small granules homogeneously distributed over the whole elytron surface.
Description of the holotype (figures 2.B1, 4.C). Elytron elliptical elongated in shape with the proximal margin similar to the distal one, both semicircular, 5.3 mm long and 1.58 mm wide (L/W ratio = 3.3, see table 1). Lateral border narrow. Six subparallel costae. Ornamentation consisting of small granules homogeneously distributed over the whole surface of elytron.
Discussion. Ademosyne hexacostata sp. nov. differs from the previously described species by the costae number (six in A. hexacostata, nine in A. arcucciae, eleven in A. elongatus and eight to eleven in Australian species, sensu Dunstan, 1923, p. 73). The L/W ratio in A. hexacostata (2.72) is similar to that of A. arcucciae (2.65), but differs in having the costae subparallel (divergent in the previously described species).

Ademosyne punctuada Martins-Neto and Gallego sp. nov. Figures 2.C1-C6, 5.A

Etymology. Refers to the ornamentation of punctated costae.

Material. Holotype: PULR-I 309 (figures 2.C1, 5.A). Paratypes: from Picos Gemelos (5th cycle), three specimens (PULR-I 309a; CTES-PZ 7298), from Río Gualo (2nd cycle), five specimens (PULRI 313a, c-e figures 2.C2, C4-C6) and from Agua Escondida (4th cycle), one specimen (PULR-I 313b, figure 2.C3).

Type locality. Picos Gemelos locality, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron elliptical, elongated in shape with length varying between 2.83 mm and 4.5 mm (media 3.91, standard deviation +/- 0,4), width varying between 1.16 and 1.66 mm (media 1.52, standard deviation +/- 0.12). Ratio of length/width varying from 2.2 to 2.7 (media 2.47, standard deviation +/- 0.28). Six to nine punctate, non-convergent costae. Space between costae bearing small granules homogeneously distributed over the whole elytron surface.
Description of the holotype (figures 2.C1, 5.A). Elytron elliptical, elongated in shape with the proximal margin similar to the distal one, both semicircular, with 4.5 mm long and 1.66 mm wide (relation L/W = 2.7, see table 1). Nine punctate, non-convergent costae. Ornamentation consisting of small granules homogeneously distributed over the whole elytron surface.
Discussion. Ademosyne punctuada sp. nov. has a combination of characters present in the previously described species such as the L/W ratio (2.47 in A. punctuada; 2.72 in A. hexacostata and 2.65 in A. arcucciae), usually nine costae (as in A. arcucciae), occasionally six (as in A. hexacostata), costae not convergent, subparallel (as in A. hexacostata), and the space between costae bearing small, homogeneously distributed granules (as in all previously described species). The presence of punctate costae is unique for Ademosyne punctuada sp. nov., also only previously reported for A. parva Dunstan and A. brevis Dunstan from the Ipswich series (Australia). Both forms differ from A. punctuada sp. nov. in the costae convergent to the elytron apex, L/W ratio (1.8 in A. brevis), and general ovate outline.

Argentinosyne Martins-Neto and Gallego gen. nov.

1997. Tillyardopsis in Gallego, p. 514, lám. IH, partim.
1997. Mesostigmodera? in Gallego, p. 514, lám. IJ, partim.
1999. Mesostigmodera in Gallego and Martins-Neto, p. 87, Fig. 6, partim.

Type species. Argentinosyne frenguellii Martins-Neto and Gallego sp. nov., designated here.

Etymology. Refers to República Argentina, and syne, common suffix for Permosynidae.

Diagnosis. Elytron length varying from 1.4 mm to 18 mm, width varying from 0.5 mm to 5.5 mm. Length / width ratio varying from 1.4 to 4.3. Elytron without costae and ornamentation varying from homogeneous granules to rugose.
Discussion. Argentinosyne gen. nov. differs from Ademosyne Handlirsch (1906) and Platycrossos Dunstan (1923) in the absence of costae (well-defined in Ademosyne as well as Platycrossos), as well as by the great size and shape of the elytra. Non-costate genera from Australia, as Tillyardiopsis Dunstan, Mesostigmodera Handlirsch and Reeveana Dunstan slightly resemble the new genus. Due to this fact, in Gallego (1997) and Gallego and Martins-Neto (1999) different specimens of the new genus were tentatively assigned to the first two genera. The three Australian taxa differ from Argentinosyne gen. nov. in the elytron shape (equal sutural and lateral borders and boat-like outline in Reeveana and Tylliardiopsis), broad sutural border and acutelly apex (in Mesostigmodera) and ornamentation (smooth only in Reeveana, with sigmoid rows of granules in Mesostigmodera and convergent in Tylliardiopsis).

Argentinosyne frenguellii Martins-Neto and Gallego sp. nov. Figures 3.A1-A5, 4.D


Figure 3. A1-A5, Argentinosyne frenguellii Martins-Neto and Gallego gen. et sp. nov. A1, holotype / holotipo, PULR-I 227. A2-A5, paratype / paratipo, PULR-I 225 (A2), PULR-I 326a-c (A3-A5). B-C, Argentinosyne gonaldiae Martins-Neto and Gallego sp. nov. B1, holotype / holotipo, PULR-I 321. B2, paratype / paratipo, PULR-I 322. B3, paratype / paratipo, CTES-PZ 7305a. C, paratype / paratipo, MACNM 24259. D, Permosynidae? indet., CTES-PZ 7305b (D1), PULR-I 332 (D2), PULR-I 333 (D3). E, Argentinocupes pulcher Martins-Neto and Gallego gen. et sp. nov., holotype / holotipo, PULR-I 299. F, Argentinocupes abdalai Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 310. G, Gemelina triangularis Martins-Neto and Gallego gen. et sp. nov. G1, holotype / holotipo, PULR-I 280a. G2, paratype / paratipo, PULR-I 323. H, Gen. et sp. indet. 1. H1-H2, PULR-I 328- 329. I, Cardiosyne obesa Martins-Neto and Gallego gen. et sp. nov. I1, holotype / holotipo, PULR-I 324. I2-I6, paratype / paratipo, PULR-I 325a-e. J, Cardiosyne elegans Martins-Neto and Gallego sp. nov. J1, holotype / holotipo, PULR-I 312. J2-J4, paratype / paratipo, PULR-I 327a-c. K, Isolated and fragmented Coleoptera bodies / fragmentos aislados de Coleoptera. K1-K5, CTES-PZ 7305c; K6-K8, PULR-I 226; K9-K10, PULR-I 330; K11-K12, PULR-I 331. Scale bar / escala = 1 mm.


Figure 4. A, Ademosyne arcucciae Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 298, partially articulated specimen with the right elytron absent showing a dorsal view of the thorax and abdomen / espécimen parcialmente articulado con el élitro derecho ausente mostrando una vista dorsal del tórax y abdomen. B, Ademosyne elongatus Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 304. C, Ademosyne hexacostata Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 305, partially articulated specimen with both elytra, pronotum and three appendages in the left side of the photograph / espécimen parcialmente articulado con ambos elitros, el pronoto y tres apéndices del lado izquierdo de la fotografía. D, Argentinosyne frenguellii Martins-Neto and Gallego gen. et sp. nov., holotype / holotipo, PULR-I 227, partially articulated specimen with both elytra / espécimen parcialmente articulado con ambos elitros. E, Argentinosyne gonaldiae Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 321. F, Argentinocupes abdalai Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 310, partially articulated specimen with both elytra / espécimen parcialmente articulado con ambos elitros. G, Gemelina triangularis Martins-Neto and Gallego gen. et sp. nov., holotype / holotipo, PULR-I 280a. H, Cardiosyne obesa Martins-Neto and Gallego gen. et sp. nov., holotype / holotipo, PULR-I 324. I, Cardiosyne elegans Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 312. Scale bar / escala = 1 mm.

1997. Tillyardopsis sp. Gallego, p. 513, lám. IH.
1997. Mesostigmodera ? sp. Gallego, p. 513, lám. IJ.
1999. Mesostigmodera frenguelli Gallego and Martins-Neto (nomen nudum), p. 88, Fig. 6.

Etymology. In honor of the notable Argentinean naturalist and paleobotanist Joaquín Frenguelli.

Material. Holotype: PULR-I 227 (figures 3.A1, 4.D), CTES-PZ 7297 (counterpart). Paratypes: From Picos Gemelos (5th cycle), PULR-I 225 (figure 3.A2), PULR-I 326a-c (figures 3.A3-A5), four specimens.

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron length varying from 1.4 mm to 18 mm (media 14.88, standard deviation +/- 1.8), width varying from 3.5 to 5.5 mm (media 4.4, standard deviation +/- 0.65). Length/width ratio varying from 3.6 to 4.2 (media 3.48, standard deviation +/- 0.46). Ornamentation of homogeneous granules.
Description of the holotype (figures 3.A1, 4.D). Rather large permosynid, without well-delimited costae, around four-times longer than wide. Elytron 14.5 mm long and 3.5 mm wide (L/W ratio = 4.14, see table 2), with conspicuous lateral border. Ornamentation consisting of small granules homogeneously distributed over the whole elytron surface.
Discussion. Tylliardopsis tuberculata Dunstan and T. variotuberculata Dunstan differ from the new species in boat-like elytron shape, ornamented with 9 to 16 convergent rows of granules (more than 15 straight rows in A. frenguellii) and the L/W ratio (2.6 in both species). Mesostigmodera typica Etheridge and Olliff differs from A. frenguellii sp. nov. in the broad sutural border and the acutelly apex, elytron ornamented with sigmoid rows of granules and the L/W ratio (3.1 in M. typica).

Argentinosyne gonaldiae Martins-Neto and Gallego sp. nov. Figures 3.B1-B3, C, 4.E

Etymology. Dedicated to the anthropologist María Elena Gonaldi from the Museo de Ciencias Naturales, Universidad Nacional de La Rioja.

Material. Holotype: PULR-I 321 (figures 3.B1, 4.E). Paratypes: from Río Gualo (5th cycle form the Los Rastros Formation), two specimens (PULR-I 322, figure 3.B2; CTES-PZ 7305a, figure 3.B3); from Cacheuta Formation (southeast of the Cerro Los Colorados, Paramillos de Uspallata, Mendoza Province, Argentina), one specimen (MCNAM 24259, figure 3.C).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 3rd cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron from 5 to 8 mm long and 2 mm to 3.5 mm wide. L/W ratio 2.4. Ornamentation consisting of rugose waves of granules homogeneously distributed over the whole elytron surface.
Description of the holotype (figures 3.B1, 4.E). Rather medium-sized permosynid, without well-delimited costae, two and half-times longer than wide. Elytron 7.91 mm long and 3.25 mm wide (L/W relation = 2.4, see table 2), with conspicuous lateral border. Ornamentation consisting of rugose waves of granules homogeneously distributed over the whole elytron surface.
Discussion. This new species differs from Argentinosyne frenguellii sp. nov. in having a rather smaller size and distinct ornamentation pattern (rugose in A. gonaldiae sp. nov., granulated in A. frenguellii sp. nov.), and small length/width ratio (2.4 in A. gonaldiae sp. nov., 3.5 in A. frenguellii sp. nov.). Tillyardiopsis granulata Dunstan differs from the new species in the elytron shape (equal sutural and lateral borders and boat-like outline, narrow sutural and moderate wide lateral borders, acutelly apex and smooth or slightly rugose-granulate ornamentation. Reeveana mayor Dunstan and R. minor Dunstan differ from A. gonaldie in the elytron shape (equal sutural and lateral borders and boat-like outline), narrow sutural border and smooth ornamentation.

Argentinosyne gualoensis Martins-Neto and Gallego sp. nov. Figures 2.E1-E5, 5.C


Figure 5. A, Ademosyne punctuada Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 309. B, Argentinosyne rugosa Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 319. C, Argentinosyne gualoensis Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 311. D, Argentinosyne losrastrosensis Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 344. E, Argentinocupes pulcher Martins-Neto and Gallego sp. nov., holotype / holotipo, PULR-I 299. Scale bar / escala = 1 mm.

Etymology. Refers to the Río Gualo locality, from which the material came.

Material. Holotype: PULR-I 320 (figures 2.E5, 5.C). Paratypes: From Río Gualo, twelve specimens (PULR-I 311a-d, figures 1.E1- E4).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 5th from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron elliptical, elongated in shape with length varying from 2.83 mm to 8.16 mm (media 4.91, standard deviation +/- 1.72), width varying from 0.79 to 2.5 mm (media 1.56, standard deviation +/- 0.54). Ratio of length/width varies from 2.7 to 4.3 (media 3.36, standard deviation +/- 0.39). Elytron without costae or ornamentation (smooth).
Description of the holotype (figures 2.E5, 5.C). Elytron elliptical elongated in shape with the proximal margin similar to the distal one, both semicircular, with 5.3 mm long and 1.58 mm wide (L/W relation = 3.3, see table 2), elliptical-elongated shape, without narrow lateral border. Costae absent and smooth ornamentation.
Discussion. Argentinosyne gualoensis sp. nov. differs from the previously described species in the elytra shape, with the anterior border similar to the posterior border, and the elytron more than three times longer than wide. A. gualoensis sp. nov. has an elytron without costae but is totally smooth, whereas Argentinosyne frenguellii sp. nov. has a granulated ornamentation. The Australian species Lobites granulatus Dunstan resembles A. gualoensis in the elytron shape, but differs in the sigmoid lower sutural border and granulate ornamentation.

Argentinosyne rugosa Martins-Neto and Gallego Figures 2.F, 5.B

Etymology. Refers to the surface of the elytron.

Material. Holotype: PULR-I 319 (figures 2.F, 5.B). Paratypes: From Río Gualo (Los Rastros Formation), two specimens (PULR-I 319); from the Cacheuta Formation (southeast of the "Cerro Los Colorados", Paramillos de Uspallata, Mendoza Province, Argentina), one specimen (MCNAM 24258).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron elliptical, elongated in shape with length varying from 1.9 mm to 7.4 mm (media 4.4, standard deviation +/- 2.8), width varying from 0.83 mm to 2 mm (media 1.38, standard deviation +/- 0.58). Length/width ratio varies from 2.3 to 3.7 (media 2.96, standard deviation +/- 0.7). Elytron without costae, with rugose ornamentation.
Description of the holotype (figures 2.F, 5.B). Elytron elliptical elongated in shape with the proximal margin similar to the distal one, both semicircular, 7.4 mm long and 2 mm wide (L/W relation = 3.7, see table 1). Costae absent. Ornamentation rugose, homogeneously distributed over the whole elytron surface.
Discussion. Argentinosyne rugosa sp. nov. is similar to A. gualoensis sp. nov. in having an elongated elytron, more than three times longer than wide, but differs in having a rugose ornamentation (without ornamentation in A. gualoensis). In this respect A. rugosa is similar to A. gonaldiae, though it differs in its notably greater L/W ratio.

Argentinosyne losrastrosensis Martins-Neto and Gallego sp. nov. Figure 5.D

Etymology. Relative to Los Rastros, the stratigraphic unit from which the material came.

Material. Holotype PULR-I 344 (figure 5.D). Paratype. From the Picos Gemelos, one specimen (PULR-I 343).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation
Age. Early Late Triassic.
Diagnosis. Elytron boat-like shape, smooth with length varying from 1.25 mm to 4.08 mm, and width from 0.58 mm to 2.08 mm. Length/width ratio average 2.12, standard deviation +/- 0.48. Costae absent and ornamentation smooth.
Description of the holotype (figure 5.D). Rather small permosynid, without costae, around two-times longer than wide. Elytron boat-like shape, smooth, 2.9 mm long and 1 mm wide (L/W ratio = 2.9).
Discussion. A. losrastrosensis sp. nov. iffers from Argentinosyne frenguellii Martins-Neto and Gallego, A. gonaldiae Martins-Neto and Gallego, A. rugosa Martins-Neto and Gallego, and A. gualoensis Martins-Neto and Gallego in having a rather smaller size and distinct ornamentation pattern (rugose in A. gonaldiae and A. rugosa, granulated in A. frenguellii), and the smallest length/width ratio (2.4 in A. gonaldiae, 3.5 in A. frenguellii, 2.9 in A. rugosa and 3.3 in A. gualoensis). A. losrastrosensis sp. nov. is similar to A. gualoensis Martins-Neto and Gallego by having no ornamentation, differing however in having a notably small L/W ratio.

Family Permosynidae? gen. et sp. indet. Figures 3.D1-D3

Material. CTES-PZ 7305b, PULR-I 332, 333, collected at Río Gualo, La Rioja Province, Argentina.

Stratigraphical unit and age. 2nd cycle from the Los Rastros Formation, early Late Triassic.
Description. Fragmentary material lacking the distal and/or proximal elytra margin, but they are too poorly preserved to assign them to any known genus.
Remarks. The specimen PULR-I 333 (figure 3.D3) could belong to Argentinosyne gonaldie.

Family Cupedidae Lacordaire, 1857

Argentinocupes Martins-Neto and Gallego gen. nov.

Type species. Argentinocupes pulcher Martins-Neto and Gallego sp. nov.

Etymology. Refering to República Argentina, from which the material came, and cupes, common suffix for cupedid-like elytra.
Diagnosis. Elytron with seven slightly curved granulated costae. Space between costae filled by eight homogeneous rows with square-granulated ornamentation.
Discussion. Similar to Simmondsia Dunstan, 1923 from the Australian Triassic (Ipswich series) and Moltenocupes Zeuner, 1961 from the Molteno Formation (South Africa) in its ornamentation pattern (homogeneous rows of square granulated ornamentation), but differing in the number of costae (seven in Argentinocupes gen. nov., nine in Simmondsia Dunstan and ten in Moltenocupes Zeuner). Additionally Argentinocupes gen. nov. has costae slightly curved (versus quite straight in Simmondsia, straight upward and curved downward in Moltenocupes).

Argentinocupes pulcher Martins-Neto and Gallego sp. nov. Figures 3.E, 5.E

Etymology. From Greek, pulchrus, beautiful.

Material. Holotype: PULR-I 299 (figures 3.E, 5.E).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 2nd cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron small, elliptical with around 3 mm long, as preserved, and 1.2 mm wide. L/W ratio 2.4. Elytron apex rounded and base slightly convex. Costae narrow.
Description of the holotype (figures 2.E, 5.E). Elytron 2.9 mm long as preserved, and 1.2 mm wide with seven slightly curved granulated costae. Space between costae filled by eight homogeneous rows with square-granulated ornamentation.
Discussion. Simmondsia sub-piriformis Dunstan and Moltenocupes townrowi Zeuner, the more close Triassic cupedid-like coleopterans from Gondwana, differ from the new species in the elytron shape, more acutelly elytron apex and the more regular square ornamentation between costae. Also M. townrowi Zeuner remains are constituted by the head, mouth parts, antennae and sternites; its total length (from the head to the etrytron apex) is 23 mm.

Argentinocupes abdalai Martins-Neto and Gallego sp. nov. Figures 3.F, 4.F

Etymology. Dedicated to Fernando Abdala for his contributions on Argentinean Triassic tetrapods.

Material. Holotype: PULR-I 310 (figures 3.F, 4.F).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 2nd cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytron large, boat like shape with 9.3 mm long and 2.6 mm wide. L/W ratio 3.6. Costae broad.
Description of the holotype (figures 3.F, 4.F). Elytron 9.3 mm long and 2.6 mm wide with seven quite straight granulated costae. Space between costae filled by eight homogeneous rows with square-granulated ornamentation.
Discussion. Simmondsia cylindrica Dunstan differs from the new species in the elytra shape, less acutely elytra apex, narrow lateral border and the rectangular ornamentation between costae. Moltenocupes townrowi Zeuner differs also in the characters mentioned above (see discussion of A. pulcher) and in the narrow costae. Argentinocupes abdalai sp. nov. differs from Argentinocupes pulcher sp. nov. in having the elytra three and a half times longer than wide (versus two and a half times longer in Argentinocupes pulchrer sp. nov.) and shape (elliptical in A. pulchrer). Also, it differs in the more wide costae and square ornamentation between them (in A. abdalai).

Family Elateridae? Leach, 1815

Gemelina Martins-Neto and Gallego gen. nov.

Type species. Gemelina triangularis Martins-Neto and Gallego sp. nov.

Etymology. Diminutive of Gemelos, related to Picos Gemelos, the geographic locality from which the material came.
Diagnosis. Elytron two times longer than wide, triangular shape, with granulated surface. Proximal margin roof-like.
Discussion. Similar to Elaterites Dunstan, 1923, from the Australian Triassic in the general elytra habitus, but differing in having a proximal margin of elytron roof-like (versus quite straight in Elaterites Dunstan). Gemelina gen. nov. is also similar to an unnamed species figured by Brauckmann and Schlüter (1993, Fig. 18), from the Triassic of Lower Franconia (Germany), although without the proximal margin of the elytron preserved for comparison.

Gemelina triangularis Martins-Neto and Gallego sp. nov. Figures 3.G1-G2, 4.G

Etymology. Related to the triangular elytron shape.

Material. Holotype PULR-I 280a, part (figures 3.G1, 4.G); counterpart, CTES-PZ 7275a.

Paratype. From quebrada Ischichuca Chica, one specimen (PULR-I 323, figure 2.G2).

Type locality. Picos Gemelos, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. As for the genus. Elytron 9.6 mm long and 4.9 mm wide. L/W relation 1.9
Description of the holotype (figures 3.G1, 4.G). Elytra 9.6 mm long and 4.9 mm wide, covered by a granulated ornamentation. Proximal elytra margin roof-like. Anterolateral margin slightly curved; posterolateral margin straight. Margin border not welldefined. Apex slightly acuminate.
Discussion. Elaterites subulatus Dunstan is one of the four elaterid-like species described from the Ipswich series (Australia). It differs from the new species in the symmetrical shape and the smaller and rugose ornamentation of the elytron surface.

Cardiosyne Martins-Neto and Gallego gen. nov.

Type species. Cardiosyne obesa Martins-Neto and Gallego sp. nov.

Etymology. Related to the heart-shaped elytra.
Diagnosis. Elytron about twice as long as wide, heart shaped, with smooth surface. Proximal margin rooflike with a thick and conspicuous border.
Discussion. Cardiosyne gen. nov. differs from all known Triassic genera in having a peculiar elytra shape, heart-like, thick and smooth. This new genus is apparently endemic to the Argentinean Triassic, and is relatively abundant, mainly at the Gualo locality. The shape of the elytra indicates a very wide coleopteran; the body width could attain circa 15 mm. The assignment to Elateridae is merely tentative and it is probable that this new genus belongs to a new family.

Cardiosyne obesa Martins-Neto and Gallego sp. nov. Figures 3.I1-I6, 4.H

Etymology. Related to the fat elytra.

Material. Holotype PULR-I 324 (figures 3.I1, 4.H), CTES-PZ 7300.Paratype. From Río Gualo, five specimens (PULR-I 325a-e, figures 2.I2-I6), all from the 5th cycle, except PULR-I 325a from the 2nd cycle from the Los Rastros Formation.

Type locality. Picos Gemelos, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytra heart-like shape, 4 mm to 6 mm long and 4.5 mm to 7 mm wide. L/W ratio 0.8 to 1.1. Anterolateral margin slightly sigmoid; posterolateral one convex.
Description of the holotype (figures 3.I1, 4.H). Elytra heart-like shape, 5.83 mm long and 7 mm wide. L/W relation 0.8. Proximal elytra margin rooflike with a thick and conspicuous border. Anterolateral margin slightly sigmoid; posterolateral one convex. Elytra apex slightly acuminate.

Cardiosyne elegans Martins-Neto and Gallego sp. nov. Figures 3.J1-J4, 4.I

Etymology. Related to the elytra shape.

Material. Holotype PULR-I 312 (figures 3.J1, 4.I). Paratype. From Río Gualo, three specimens (PULR-I 327a-c, figures 3.J2-J4).

Type locality. Río Gualo, La Rioja Province, Argentina.
Type stratum. 5th cycle from the Los Rastros Formation.
Age. Early Late Triassic.
Diagnosis. Elytra elongated, 4mm to 6 mm long and 3.5 mm to 6 mm wide. L/W relation 1.2 to 1.4. Anterolateral margin slightly curved; posterolateral one sigmoid.
Description of the holotype. (figures 3.J1, 4.I). Elytra elongated, heart-shaped, 6.08 mm long and 4.16 mm wide. L/W ratio 1.4. Proximal elytra margin roof-like with a thick and conspicuous border. Anterolateral margin slightly curved; posterolateral one sigmoid. Elytra apex slightly acuminate.
Discussion. C. elegans sp. nov. differs from C. obesa sp. nov. in having a more elongate elytra and with a more conspicuously convex posterolateral margin.

Family uncertain gen. et sp. indet. 1 Figures 3.H1-H2

Material. PULR-I 328, collected at Picos Gemelos and PULR-I 329, collected at Agua Escondida locality, La Rioja Province, Argentina.

Stratigraphical unit and age. 3rd cycle from the Los Rastros Formation, early Late Triassic.
Description. PULR-I 328 (figure 3.H1) triangular elytra fragment 3.6 mm long as preserved, and 2 mm wide, with a pronounced posterolateral border. The specimen PULR-I 329 (figure 3.H2) is a triangular elytra fragment 4.5 mm long as preserved, and 1.8 mm wide, also with a pronounced posterolateral border.
Remarks. Two similar fragments collected from two distinct localities have the same preservation stage, elytra shape, ornamentation pattern and a conspicuous posterolateral border. These characters are not found in any previously described species, indicating a possible new genus for this material, but it is too fragmentary for a formal description and taxonomic treatment.

Discussion

The genus Ademosyne Handlirsch has a world wide distribution (figure 2.H) with representatives in Argentina (this paper), Chile (Gallego et al., 2005), South Africa (Riek, 1974, 1976), Australia (Handlirsh, 1906; Tillyard, 1916; Dunstan, 1923), Germany (Brauckmann and Schlütter, 1993), Japan (Fujiyama, 1973), and Russia (Ponomarenko, 1969), consisting of a typical group of Coleoptera for Triassic sediments (although some scant records for Jurassic sediments are also known). The elytra shape and ornamentation pattern of this group are highly variable, which creates a major taxonomic problem. Authors such as Dunstan (1923) considered each variation as a diagnostic character at the specific or sometimes, generic level (number of costae, costae ornamentation, ornamentation between costae, elytra shape and size). Because of this, several new species have been proposed in the past. The lack of consistent characters based only on the morphology of isolated elytra does not allow a phylogenetic approach. Evidently a broad revision of the Ademosyne-like group is necessary. Ademosyne is probably a paraphyletic group.
The situation of the Ademosyne-like specimens collected from the Argentinean Triassic is not verydifferent. Several morphotypes are present, with a great variation in shape, costae number and ornamentation. The size of specimens and their respective variation can demonstrate paleoecological and/or taphonomic problems (sometimes a smooth elytra does not necessarily indicate a distinctive character, but merely a preservational problem of a moult; sometimes the part preserves the ornamentation and the counterpart does not). The presence or absence of punctate costae can indicate merely sexual dimorphism or just a taphonomic problem. The same can be said for the elytra shape, adding other geological problems (not easily detected), such as sediment compaction after burial (resulting in a deformed elytra of unexpected shape), chemical factors destroying the elytron surface, sediments washing and polishing the elytra surface after burial (eliminating the weak ornamentation pattern), and so on. The ademosynid specimens from the Argentinean Triassic sediments indicate a moderate transport before burial (more than 90% of the specimens consist of isolated elytra).
The most useful of the morphometric parameters used here is the length/width ratio, which defines distinct species reasonably well (this parameter is not dependent of the size variability), but evidently it does not solve the phylogeny of the group. A systematic approach based also on body morphology as proposed by Ponomarenko (1969) is certainly the best, but around 99% of the known ademosynid-like fossils consist only of isolated elytra, making if impossible to take this path. Only a single elytron, possibly associated with the body, was found in the Argentinean Triassic, as well as around 2% of isolated and fragmented Coleoptera bodies, CTES-PZ 7305c, PULR I 226, 330, 331- see figures 3.K1-K12).

Apéndice

Table 1. Morphometric variation (mm), ornamentation type and preservation degree on Ademosyne spp. / Variación morfométrica (mm), tipo de ornamentación y grado de preservación en Ademosyne spp. Abbreviations / Abreviaturas: *, as preserved / como se preserva; Bet. costae, between costae / entre costillas; Frag., fragmented / fragmentado; Compl., complete / completo; Art., articulated / articulado; Disart., disarticulated / desarticulado.


Table 2. Morphometric variation (mm), ornamentation type and preservation degree on other Coleoptera species / variación morfométrica (mm), tipo de ornamentación y grado de preservación en otras especies de Coleoptera. Abbreviations / abreviaturas: *, as preserved / como se preserva; Bet. costae, between costae / entre costillas; Frag., fragmented / fragmentado; Compl., complete / completo; Art., articulated / articulado; Disart., disarticulated / desarticulado.


Acknowledgements

We thank A. Arcucci, C. Forster, C. May and R. Rogers from the project of the National Geographic Society, Grant 5317-94. G. Barrios for making the digital figures and R.Z. Rodríguez for his work in the photographic laboratory, M.E. Gonaldi from the Museo de Ciencias Naturales (Universidad Nacional de La Rioja, Argentina) for her assistance with cataloguing the specimens and also to E. Cerdeño from the Museo de Ciencias Naturales y Antropológicas "Cornelio Moyano" (Mendoza, Argentina). Also thanks to the reviewers A. Nel, A. Alexeev, A. Ponomarenko, A. Ashworth, G. Edgecombe and E. Clarkson for their helpful comments and suggestions that enhanced the manuscript. This research was partly supported by the Consejo Nacional de Investigaciones Científicas y Técnicas (PIP-5581) and by the Secretaría General de Ciencia y Técnica, Universidad Nacional del Nordeste (PI-64/04), both to O.F.G.

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Recibido: 29 de diciembre de 2003.
Aceptado: 21 de marzo de 2005.

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