Oreiallagma gen. nov. with a redefinition of Cyanallagma Kennedy 1920 and Mesamphiagrion Kennedy 1920, and the description of M. dunklei sp. nov. and M. ecuatoriale sp. nov. from Ecuador (Odonata: Coenagrionidae) Author Ellenrieder, Natalia Von Author Garrison, Rosser W. text Zootaxa 2008 1805 1 51 journal article 10.5281/zenodo.182666 d1698b4a-ec6b-4a5f-96cf-956535545e85 1175-5326 182666 Cyanallagma Kennedy 1920 Figures 1 a–2a; 3a–b; 4a; 6; 9–14; 30–37; 54–59; 95–97; 102; 105–106 Cyanallagma Kennedy 1920 : 87 (diagnosis; designation of Acanthagrion interruptum Selys 1876 as type species). Bulla 1973 : 95–104, figs. 1–21 (discussion of generic diagnosis, redescription of adults of C. bonariense and C. interruptum and description of larvae, illustration of diagnostic characters). Davies 1981 : 5 (generic listing with type species listed). Davies & Tobin 1984 : 66 (synonymic list). De Marmels 1988 : 100 (synonymy of Archaeallagma ). De Marmels 1989 : 246–248 (generic characterization). Bridges 1994 : III.13 (synonymic list). De Marmels 1997 : 135–156, figs. 1–85 (generic diagnosis, keys, maps, and illustrations for northern species, description of the larva of C. gaianii ). Steinmann 1997 : 247 (synonymic list). Tsuda 2000 : 31 (synonymic list). Lencioni 2001 : 345–349, figs. 1–14 (generic key and illustrations for southern species). De Marmels 2003 : 103–106, figs. 8–16, 21 (description of C. ferenigrum , illustrations of male S10, genital ligula, head, thorax, and S1–2, S7–10, pt, male and female posterior prothoracic lobe, map, discussion of generic placement). Lencioni 2006 : 25, 33, fig. B7 (key to Brazilian genera; diagnosis). Type species: Acanthagrion interruptum Selys 1876 by original designation ( Kennedy 1920: 87 ). Other species included: C. angelae Lencioni 2001 , C. bonariense ( Ris 1913 ) , C. ferenigrum De Marmels 2003 , C. nigrinuchale ( Selys 1876 ) , and C. trimaculatum ( Selys 1876 ) . FIGURE 5. Color pattern of head, dorsal view, and body, lateral view, Oreiallagma thelkterion , male paratype, Venezuela, Monte Zerpa. Specimens examined. C. angelae (2 ɗ, 1 Ψ). BRAZIL . São Paulo State: Salesópolis, 2.xi.2001 , F.A.A. Lencioni leg., 2 ɗ, 1 Ψ (RWG). C. bonariense (9 ɗ, 3 Ψ). ARGENTINA . Buenos Aires Prov.: Arroyo Caracol Chico, N of Berisso, 23.xi.1979 , C.M. & O.S. Flint, Jr. leg., 1 ɗ, 2 Ψ (RWG); Berisso, 24.ii.1989 , G. Jurzitza leg., 1 ɗ (RWG); Villa Elisa, 26.xi.1979 , C.M. & O.S. Flint, Jr. leg., 3 ɗ (RWG); Campana, Delta del Paraná, San Fernando, 8.i.1999 , J. Muzón & N. von Ellenrieder leg., 3 ɗ, 1 Ψ (MLP). Córdoba Prov.: Arroyo Las Vacas, Embalse Río Tercero, 24.xi.1981 , A. Rodrigues Capítulo leg., 1 ɗ (MLP). C. ferenigrum (1 ɗ, 1 Ψ). BRAZIL . Mato Grosso State: Utiariti, viii.1961 , K. Lenko leg., 1 ɗ holotype , 1 Ψ paratype (ABMM). C. interruptum (21 ɗ, 10 Ψ). CHILE . Aisén Region: 19 km N de Puerto Ibáñez, 590 m , 22.i.1992 , J. Muzón leg., 2 ɗ, 1 Ψ (RWG); same 1 ɗ (MLP). Osorno Region: Maicolpue, spring-fed pools, 22.xii.1993 , C.M. & O.S. Flint, Jr. leg., 1 ɗ (RWG). de los Lagos Region: Cuesta Moraga, 9 km N Villa Santa Lucía , 24.i.1987 , C.M. & O.S. Flint, Jr. leg., 3 ɗ (RWG); 10 km SW Futaleufú, 28.i.1987 , C.M. & O.S. Flint, Jr. leg., 1 ɗ, 1 Ψ (RWG); Lago Lonconao, just W of Futaleufú, 17.i.1987 , R.W. Garrison leg., 10 ɗ, 2 Ψ (RWG); Ensenada, 12-14.xii.1926 , R.C. & E.S. Shannon leg., 1 ɗ (RWG). Malleco region: Cordillera de Nahuelbuta, marsh N of Coimalín, 16.xii.1993 , C.M. & O.S. Flint, Jr. leg., 1 ɗ, 1 Ψ (RWG). Maule Region: Reloca, S of Constitución, damp area near road, 16.xii.1976 , Gurney & Barria leg., 3 ɗ, 1 Ψ (RWG). Valparaiso region: Valparaiso, M. Mac Lachlan leg., 1 ɗ lectotype (BMNH); same data 1 ɗ, 1 Ψ paralectotypes (IRSNB). ARGENTINA . Santa Cruz Prov.: El Calafate, Ayo. Los Perros (estepa extra-andina), 19.i.1992 , J. Muzón leg., 1 ɗ (RWG). Chubut Prov.: Esquel, Laguna La Zeta, 19.i.1989 , J. Muzón leg., 2 Ψ (RWG); Parque Nacional Nahuel Huapi, charca camino a Los Alerces, 24.i.1988 , J. Muzón leg., 2 ɗ (RWG); Languineo, pond and stream nr. Tecka, by route 62, 680 meters, 16.i.1995 , R.W. Garrison leg., 11 ɗ, 5 Ψ (RWG). Neuquen Prov.: Ruca Malén, 23 km N Villa La Angostura, 29.xii.1993 , C.M. & O.S. Flint, Jr. leg., 1 Ψ (RWG); P.N. Lanín, Lago Queñi, 7.ii.1999 , J. Muzón & P. Marino leg., 1 ɗ, 1 Ψ (MLP). Río Negro Prov.: Meseta de Somuncurá, Valcheta, Ea. El Rincón, 28.i.1999 , J. Muzón & N. von Ellenrieder leg., 1 ɗ (MLP); Laguna Cari Lafquen Chica, Aguada, 4.ii.1999 , J. Muzón & P. Marino leg., 1 ɗ, 1 Ψ (MLP). C. nigrinuchale (3 ɗ, 6 Ψ). In order to clarify application of the name, we designate the syntype male from Brazil , Minas Gerais, São João del Rei, xi.1872 , Walthère de Selys leg. as lectotype , and the 5 Ψ syntypes with the same data as paralectotypes (IRSNB). BRAZIL : São Paulo State: Fazenda Santana do Rio Abaixo, Jacareí, 1-6.xi.1999 , F.A.A. Lencioni leg., 2 ɗ (RWG); same but 8.xii.1998 , 1 Ψ (RWG). ARGENTINA . Misiones Prov.: Parque Nacional Iguazú, 13.iv.1991 , J. Muzón leg., 1 Ψ (RWG); San Pedro , junction of Rt. Nac. 14 and 17, charca, 12.iv.1991 , J. Muzón leg., 1 ɗ (RWG). C. trimaculatum (4 ɗ, 2 Ψ). BRAZIL . Rio Grande do Sul State: Santa Cruz, 10.x. , Walthère de Selys leg., 1 ɗ lectotype , 2 ɗ paralectotypes (IRSNB). Rio de Janeiro State: Teresópolis, 20.x. , Walthère de Selys leg., 1 ɗ, 2 Ψ paralectotypes (IRSNB). Generic characterization. Head . Dorsum dark brown to black with pale blue postocular spots, pale postocular bar usually absent (present in some females of C. interruptum and C. angelae ), rear of head surrounding occipital foramen black ( Fig. 1 a) to entirely dark brown in C. ferenigrum and black in C. nigrinuchale . Frons rounded, occipital lobes protruding posteriorly so that posterior most point of head is at their level ( Figs. 3 a–b). Thorax . Posterior lobe of pronotum trilobate with medial lobe developed into a caudally projected plate, especially in males ( Figs. 9 a–14a), not projected beyond lateral lobes in females of C. interruptum , C. nigrinuchale , and C. trimaculatum ( Figs. 9 b; 13b–14b). Female mesostigmal plates triangular and wide, each with maximum width of less than 0.5 of maximum length (in C. interruptum , Fig. 9 c) to rectangular and narrow, each with maximum width of more than 0.5 of maximum length (in C. angelae , C. bonariense , C. ferenigrum , C. nigrinuchale , and C. trimaculatum , Figs. 10 c–14c); mesepisternal carinae arising between mesostigmal plates, anteriorly to their postero-medial edges in C. interruptum ( Fig. 13 c), at posterior edges of mesostigmal plates in C. angelae , C. bonariense , C. ferenigrum , C. nigrinuchale , and C. trimaculatum ( Figs. 10 c–14c). Pterothorax with dark brown to black mid-dorsal and humeral stripes and usually a dark brown to black stripe over metapleural (metepimeral-metepisternal) suture (absent in some specimens of C. interruptum and C. bonariense ), with pale blue antehumeral stripe (occasionally lacking in some C. interruptum ) usually interrupted distally ( Figs. 105–106 ) but complete in C. trimaculatum and some females of C. interruptum and C. angelae . Legs short with femur 1 shorter than distance between eyes at level of antennifer (ratio = 1; Fig. 3 a), tibial spurs shorter than or as long as distance between them ( Figs. 3 a–b), pretarsal claw with well developed supplementary tooth. Wings hyaline, CuP reaching CuPAA proximal to hind margin of wing for a distance as long as CuP or shorter, vein descending from quadrangle not forming a straight line to wing margin ( Fig. 6 ). Abdomen . Black and blue and pale yellow ( Fig. 4 a), relatively short with a ratio of 3.27–4.27 to length of head plus thorax. Genital ligula distal segment lacking inner fold, with paired latero-apical and latero-medial lobes ( Figs. 30–37 ), with one or two ental membranous transverse folds distal to flexure, one between mediolateral lobes and the second between medio-lateral lobes and flexure ( Figs. 31 ; 34c), one of them usually projected into a medial membranous process, which might be folded and hidden in lateral view and is entire in C. angelae , C. bonariense , C. interruptum , C. nigrinuchale , and C. trimaculatum ( Figs. 31 ; 33–37), and bifid in C. ferenigrum ( Fig. 32 ). Second segment of genital ligula lacking wide latero-apical folds with sclerotized margins ( Figs. 30–37 ). Postero-dorsal margin of male S10 with a 'u'-shaped cleft margined by a pair of tubercles in C. angelae , C. bonariense , C. interruptum , C. nigrinuchale , and C. trimaculatum ( Figs. 54 ; 56–59), lacking tubercles in C. ferenigrum ( Fig. 55 ). Male cercus always with a short ventro-basal process ( Figs. 54– 59 ); also with a broadly triangular ventro-apical process only in C. ferenigrum and C. interruptum ( Figs. 54– 55 ). Male paraproct with a branch ending on a sclerotized tip, dorsal in C. angelae , C. bonariense , C. nigrinuchale , and C. trimaculatum ( Figs. 54 a–b; 56–58b; 59c), and medial in C. ferenigrum ( Fig. 55 ). Female with vulvar spine on S8; ovipositor slightly shorter to slightly longer than S10, not reaching tips of cerci. Generic diagnosis. No unique characters are known for Cyanallagma . The combination of a rounded frons, presence of pale postocular spots, a trilobate prothoracic posterior lobe, striped pterothorax, and male cerci armed with some kind of process is common to Apanisagrion Kennedy 1920 , Chrysobasis Rácenis 1959 a, Hesperagrion Calvert 1902 , Homeoura Kennedy 1920 , some species of Ischnura Charpentier 1840 , Leptobasis Selys 1877 , Mesamphiagrion , Oreiallagma , and Telagrion . Cyanallagma differs from Mesamphiagrion and Oreiallagma by having the rear of head surrounding occipital foramen dark ( Fig. 1 a) and by the male cercus lacking a dorsal process ( Figs. 54–59 ). The three genera are further diagnosed in Table 2 . Cyanallagma can be separated from all the remaining genera mentioned above except Hesperagrion and some Ischnura species by its most posterior point of the head located at the level of the postocular lobes ( Figs. 3 a–b) rather than at the level of eyes (as in Fig. 5 ). FIGURE 6. Wings. (a) Cyanallagma interruptum , male, Argentina, Queñi; (b) C. ferenigrum , male holotype, Brazil, Utiariti. The following characters will separate Cyanallagma from Hesperagrion : male cercus lacking a dorsal process, postero-dorsal margin of S10 with a 'u'-shaped cleft margined by a pair of tubercles ( Figs. 54–59 ), genital ligula lacking a pair of large lateral chitinized spines ( Figs. 30–37 ), and female mesanepisterna with well developed carinae forming distinct ridges ( Figs. 9 c–14c). Cyanallagma differs from Ischnura by having a ventro-basal process in male cercus ( Figs. 54–59 ), genital ligula distal segment with paired latero-apical and latero-medial lobes, lacking an inner fold, with distal corners not projected into flagellae ( Figs. 30–37 ), and female mesanepisterna with well developed carinae forming distinct ridges ( Figs. 9 c–14c). Remarks. A patch of differentiated scalariform-like cuticle apparently occurs in all species except C. ferenigrum . We confirmed its presence in C. interruptum with SEM ( Fig. 95 ) and the same appears as a pale corrugated area surrounded by dense setation when viewed at high magnification (>100x) under a binocular microscope in C. angelae , C. bonariense , C. nigrinuchale , and C. trimaculatum as well as in Oreiallagma oreas , O. prothoracicum , and O. quadricolor ( Figs. 52 ; 56–59; 87–89). Since the only available male of C. ferenigrum was the holotype , we did not subject it to SEM examination. However, the appearance of its cercus under the binocular microscope with a pale spot lacking dense setation and with no evident corrugation ( Fig. 55 ) is like the cerci of Mesamphiagrion laterale and Acanthagrion lancea Selys , both of which we examined with SEM, and confirmed a lack of any differentiated scalariform-like cuticular areas. According to Lencioni ( pers. comm. ) immature adults of C. angelae are purplish for some hours after emergence before acquiring the blue color characteristic of adults. Distribution. Southern South America from southern Chile and Argentina to central Brazil , from 0 to 1450 m above sea level ( Fig. 102 ).