NATALIA VON ELLENRIEDER

Natural History Museum of Los Angeles County

A SYNOPSIS OF THE NEOTROPICAL SPECIES OF ‘AESHNA’ FABRICIUS: THE GENUS RHIONAESCHNA FÖRSTER (ODONATA: AESHNIDAE)

von Ellenrieder, N., 2003. A synopsis of the Neotropical species of ‘Aeshna’ Fabricius: The Genus Rhionaeschna Förster (Odonata: Aeshnidae). – Tijdschrift voor Entomologie 146: 67- 207, figs. 1-468, tabs. 1-3. [ISSN 0040-7496]. Published 1 June 2003. This study includes a revisionary, phylogenetic and biogeographical analysis of Neotropical com- ponents of Aeshna Fabricius characterized by a midventral tubercle on abdominal sternum I. Phylogenetic relationships of the Neotropical species of Aeshna were inferred based on 39 adult characters. Ingroup taxa included 68 out of the 85 species currently assigned to Aeshna, and two species each of Andaeshna De Marmels and Anaciaeschna Selys. Oreaeschna dictatrix Lieftinck was chosen as outgroup. The strict consensus tree obtained after successive weighting revealed that Aeshna is not monophyletic; some of its species are more closely related to Anaciaeschna or Andaeschna. The name Aeshna should consequently be restricted to the Holarctic group includ- ing the type species Aeshna grandis Fabricius. In the present synopsis the generic name Rhion- aeschna Förster is assigned to the New World group characterized by the presence of a conical tu- bercle on abdominal sternum I, comprising 39 species formerly assigned to Aeshna. The synopsis includes keys to adults of both sexes, diagnoses, biological notes, distribution maps and more than 400 diagnostic illustrations. Rhionaeschna demarmelsi sp. n. is described, R. maita Förster is considered a junior synonym of R. brevifrons (Hagen), R. peralta (Ris) is considered a valid species, not a synonym of R. variegata (Fabricius), R. planaltica (Calvert) is raised to specific rank, ‘Aeshna’ williamsoniana Calvert, formerly included in the subgenus Hesperaeschna Cock- erell, is excluded from Rhionaeschna, and lectotypes are designated for R. maita, R. intricata (Martin), R. multicolor (Hagen), R. bonariensis (Rambur), R. diffinis (Rambur), and R. peralta. Females of three species and larvae of 16 species are still unknown. Rhionaeschna occurs from southern Argentina to southern Canada, but is primarily Neotropical with its highest diversity along the Andean mountain range between Venezuela and Bolivia. It is absent from the Amazon basin, only three species occur north to the Neotropical region. The sister group of Rhionaeschna includes some African species of ‘Aeshna’ (A. rileyi Calvert, A. subpupillata McLachlan and A. moori Pinhey). Rhionaeschna plus the African clade constitute the sister group of Andaeshna, Anaciaeschna, Anax Leach, Hemianax Selys and several species of ‘Aeshna’ of uncertain affinities (i.e. A. affinis Vander Linden, A. brevistyla Rambur, A. ellioti Kirby, A. mixta Latreille, A. isoceles Müller and A. williamsoniana); the phylogenetic relationships within this complex are not yet known and their resolution is beyond the scope of this study. Rhionaeschna is absent from the Brazilian shield. Its related species and genera (‘A.’ rileyi, ‘A.’ subpupillata, ‘A.’ moori in Africa; ‘A.’ brevistyla in Australia and New Zealand, Andaeshna in the Andes and ‘A.’ williamsoniana in Central America, ‘A.’ isoceles and highest species numbers of Anaciaeschna, Hemianax and Anax species in the Indo-Australian region) display a low diversity in Africa, which suggests a trans-Pa- cific rather than trans-Atlantic (Gondwanian) track, as has been hypothesized for other groups of similarly distributed odonates. Dr. N. von Ellenrieder, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, CA 90007, U.S.A. E-mail: [email protected] Key words. – Odonata; Aeshnidae; Aeshna; Rhionaeschna; systematics; keys; cladistics; phyloge- ny; biogeography.

CONTENTS Taxonomic analysis...... 70 Introduction ...... 68 Taxonomic part Material and methods Rhionaeschna Förster ...... 72 Phylogenetic analysis...... 69 Keys Biogeographic analysis...... 70 Keys to males of Rhionaeschna...... 73

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Keys to females of Rhionaeschna ...... 77 INTRODUCTION Rhionaeschna species accounts 1. R. draco group...... 80 The genus Aeshna was described by Fabricius in Rhionaeschna draco (Rácenis)...... 80 1775 to include four species: A. forcipata Linnaeus, A. 2. Marmaraeschna group ...... 81 grandis Linnaeus, A. variegata Fabricius and A. clavata Rhionaeschna brevicercia (Muzón & von Ellenrieder). Fabricius. Selys (1883) fixed A. juncea as the type ...... 81 species of Aeshna, but as this species was not included Rhionaeschna brevifrons (Hagen)...... 82 in the original description of the genus, its designation Rhionaeschna fissifrons (Muzón & von Ellenrieder) 83 is invalid according to the ICZN (1999). Latreille Rhionaeschna intricata (Martin)...... 84 (1810) mentioned A. forcipata as the type species of Rhionaeschna obscura (Muzón & von Ellenrieder) .84 Aeshna, but Aeshna forcipata of Latreille is a synonym Rhionaeschna pallipes (Fraser) ...... 85 of Libellula vulgatissimus and not the same species as Rhionaeschna vigintipunctata (Ris) ...... 86 A. forcipata Linnaeus, and thus it is also ineligible as 3. Schizuraeschna group...... 86 the type species of Aeshna. Therefore, the first valid Rhionaeschna dugesi (Calvert) ...... 86 type designation is that of Cowley (1934), who desig- Rhionaeschna jalapensis (Williamson)...... 87 nated Aeshna grandis as type species of Aeshna. Rhionaeschna multicolor (Hagen) ...... 88 Aeshna was the first described genus of Aeshnidae, Rhionaeschna mutata (Hagen)...... 89 and the current number of genera within this family 4. Neureclipa group ...... 89 now stands at 50. Nineteen genera (40% of the total Rhionaeschna absoluta (Calvert) ...... 89 number of currently accepted genera) have been creat- Rhionaeschna bonariensis (Rambur)...... 90 ed to include species originally described as Aeshna as Rhionaeschna diffinis (Rambur)...... 91 follows: Amphiaeschna Selys, 1871, Anaciaeschna Rhionaeschna elsia (Calvert) ...... 92 Selys, 1878, Andaeshna De Marmels, 1994, Basi- Rhionaeschna galapagoensis (Currie) ...... 93 aeschna Selys, 1883, Boyeria McLachlan, 1896, Cali- 5. R. variegata group ...... 94 aeschna Selys, 1883, Castoraeschna Calvert, 1952, Rhionaeschna brasiliensis (von Ellenrieder & Martins Coryphaeschna Williamson, 1903, Epiaeschna Hagen, Costa)...... 94 1877, Gomphaeschna Selys, 1871, Gynacantha Ram- Rhionaeschna californica (Calvert)...... 94 bur, 1842, Hemianax Selys, 1883, Nasiaeschna Selys Rhionaeschna confusa (Rambur) ...... 95 in Förster, 1900, Neuraeschna Hagen, 1867, Oplon- Rhionaeschna marchali (Rambur)...... 96 aeschna Selys, 1883, Planaeschna McLachlan, 1896, Rhionaeschna peralta (Ris)...... 97 Polycanthagyna Fraser, 1933, Remartinia Navás, 1911, Rhionaeschna tinti (von Ellenrieder)...... 99 and Staurophlebia Brauer, 1865. No unique characters Rhionaeschna variegata (Fabricius) ...... 99 define ‘Aeshna’, its species being usually identified as 6. R. cornigera group ...... 101 those not belonging to other genera. Peters (1987) an- Rhionaeschna cornigera (Brauer)...... 101 alyzed the European genera of Aeshnidae and con- Rhionaeschna haarupi (Ris) ...... 102 cluded that Aeshna was not monophyletic, with ‘Aesh- Rhionaeschna manni (Williamson & Williamson)103 na’ affinis van der Linden, 1820 and A. mixta Latreille, Rhionaeschna nubigena (De Marmels)...... 103 1805 more closely related to Anaciaeschna and Anax Rhionaeschna pauloi (Machado)...... 104 than to the remaining European Aeshna species. Rhionaeschna planaltica (Calvert)...... 104 The most recent treatment of the Neotropical Rhionaeschna psilus (Calvert) ...... 106 species of ‘Aeshna’ was by Calvert (1956). His study Rhionaeschna vazquezae (Gonzalez) ...... 106 comprised species currently assigned to Andaeshna, Re- 7. R. punctata group...... 107 martinia, Castoraeschna and Coryphaeschna. Calvert Rhionaeschna biliosa (Kennedy ) ...... 107 (1956) included detailed redescriptions of all species Rhionaeschna condor (De Marmels) ...... 108 but no diagnoses were provided outside the keys. The Rhionaeschna decessus (Calvert) ...... 108 keys were based almost exclusively on thoracic color Rhionaeschna demarmelsi von Ellenrieder sp. n. ...109 pattern, which can be intraspecifically variable, and Rhionaeschna eduardoi (Machado) ...... 110 several of his species accounts were composite descrip- Rhionaeschna joannisi (Martin)...... 110 tions of more than one species. For example, I found Rhionaeschna punctata (Martin)...... 111 that A. peralta included specimens of A. peralta, A. var- Phylogenetic analysis...... 112 iegata and A. brasiliensis; A. variegata included A. varie- Biogeographic analysis...... 113 gata and A. marchali; A. cornigera included A. cornigera Acknowledgements ...... 115 and A. planaltica; A. planaltica included A. planaltica References...... 115 and A. nubigena; A. vigintipunctata included A. vigin- Tables ...... 121 tipunctata, A. obscura and A. pallipes; A. intricata in- Illustrations...... 124 cluded A. intricata, A. brevicercia, A. fissifrons, A. intri-

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cata and A. vigintipunctata; A. brevifrons included A. Characters brevifrons and A. intricata; and A. diffinis included A. Head absoluta and A. diffinis. Since Calvert’s monograph 1. Frons transverse carina (frontal view): (0) linear or convex (1956), 12 new species have been added to the 26 (figs. 1, 7, 9-38c); (1) cleft (figs. 2-6, 8c) 2. Occipital triangle (B)/ interocular suture length (A): (0) Neotropical ‘Aeshna’ species he included, bringing it to 0.25-0.65 (fig. 415); (1) 0.7-1 (fig. 414); (2) 0.1-0.22 a total of 38 described species. Proper identification of (fig. 416) the Neotropical species of this group has been difficult 3. Margin between horizontal and vertical portions of frons not only because of problems enumerated above, but (lateral view): (0) angled (figs. 1, 9-38b); (1) rounded also due to the paucity of specimens of several species (figs. 2-8b) in the world’s collections; many species appear to be 4. Frons (lateral view): (0) not flattened (figs. 1-2, 5-6, 8- 38b); (1) flattened (figs. 3-4, 7b) rare or are seldom encountered. I treat the mostly 5. T-spot of frons (dorsal view): (0) well defined (figs. 1-39a; Neotropical components of Calvert’s genus ‘Aeshna’ 414-416); (1) not defined cladistically as well as allowing for correct identification of all its members. My results are based on an exami- Thorax nation of over 3000 specimens, including primary 6. Thoracic color pattern: (0) with pale stripes (figs. 48-58, types or paratypes of 31 out of 39 distinguished 60-74, 76-89, 91-118); (1) uniform in color (Fig 39); (2) with dark spots and pale areas (figs. 40-47). In some species. This study includes keys and diagnoses sup- species the extension of the stripes is variable being re- ported by extensive annotated illustrations, which duced or absent in some specimens (i.e. figs. 59, 75, 90); should facilitate identification of all species presently these species were scored as (0). included in the resurrected genus Rhionaeschna Förster. 7. Supratriangular crossveins: (0) present (figs. 396-398, 400-407); (1) absent (fig. 399). In species scored as lack- ing supratriangular crossveins, some specimens (about 1- MATERIAL AND METHODS 15% of the examined specimens) possessed 1-2 crossveins in 1-2 of the supratriangles. Phylogenetic analysis 8. Bending of MA: (0) slight (fig. 405, 407); (1) marked (figs. 396-404, 406) The ingroup taxa included 68 species out of the 85 9. Origin of two cell rows between RP1 and RP2: (0) proxi- currently assigned to Aeshna, two species of Andaeshna mal to pterostigma or under it (figs. 396-397, 401-405); and two of Anaciaeschna. Polarity of characters was es- (1) distal to pterostigma or at its distal end (figs. 398-400, tablished with reference to the outgroup Oreaeschna 406-407) 10. Course of RP2: (0) markedly bent (figs. 397-405, 407); dictatrix Lieftinck (Oreaeschna constitutes the sister (1) kinked at distal end of pterostigma (fig. 406); (2) group of the remaining Aeshninae according to von El- evenly curved (fig. 396) lenrieder 2002: fig. 437). Most parsimonious trees 11. IRP2 fork: (0) asymmetrical with inferior branch as a were sought using branch-swapping search (mh*, bb* continuation of IRP2 and upper branch weak (fig. 405); command) in Hennig86 (version 1.5, Farris 1988). (1) symmetrical (figs. 396-404); (2) asymmetrical with upper branch as a continuation of IRP2 and inferior Implicit enumeration commands were not used due to branch weak (fig. 406) the large number of taxa. Autapomorphies of terminal 12. Cells of anal triangle: (0) 3 (exceptionally 4 in some taxa were excluded from the analysis. All question specimens) (figs. 396-404, 406-407); (1) 2 (fig. 405) marks in the matrix refer to missing data. Multistate 13. Anal angle: (0) angled (figs. 396-406, 408-412); (1) characters were treated as unordered. All characters rounded (fig. 407, 413) were initially weighted equally, and successive weight- 14. Fusion of basal and distal veins of anal triangle: (0) par- allel for a long distance (about 30% of anal margin) (figs. ing was performed a posteriori. Cladograms were exam- 408, 411); (1) parallel for a short distance (less than 25% ined by manually plotting the proposed synapomor- of anal margin) (figs. 409-410, 412); (2) at a single point phies. at angle (fig. 413) 15. Membranule length: (0) 50-66% of anal wing margin Character analysis (figs. 408-411); (1) 75-100% of anal wing margin (figs. The lack of information on the preimaginal stages 412-413) of several species precludes using larval characters for phylogenetic analyses; comparative study was there- Abdomen fore focused on adult characters. Thirty-nine external 16. Hamular anterior process in ventral view: (0) short (fig. morphological characters and associated character 422); (1) long and triangular (fig. 423); (2) absent (figs. 424-425b,c) states are listed below. The data matrix is shown in 17. Hamular posterior process in ventral view: (0) absent table 2. Characters were chosen and scored from di- (figs. 276-315b, 422-423); (1) tuberculate (fig. 424a,b); rect examination of voucher specimens. (2) longer than width of ventral portion of hamulus (fig. 425a-c) 18. Hamular fold: (0) separated from wall of genital fossa (figs. 423-425); (1) not differentiated from wall of genital

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fossa (fig. 422) Biogeographic analysis 19. Spines of anterior lamina: (0) triangular and narrow: at base (A) wider than twice at distal 30% (B) (figs. 316, 325-346, 349-352, 355-357, 422, 424-425); (1) triangu- Endemicity areas for Rhionaeschna were delimited lar and wide: twice or less as wide at base (A) than at dis- based on the distribution of the more restricted species tal 30% (B) (figs. 317-324, 353-354); (2) cylindrical (fig. (fig. 466). An analysis was carried out to establish the 423); (3) vestigial (figs. 347-348) degree of similarity among areas of endemicity, using 20. Genital lobe: (0) low (figs. 234-235, 237, 239, 241, Jaccard’s association coefficient and average linking. A 243-273, 418); (1) high and projected ventrally (figs. 236, 238, 240, 242, 274-275, 419); (2) very high and parsimony analysis of endemicity (PAE; Rosen 1988, projected postero-ventrally (fig. 421); (3) no higher than Cracraft 1991, Morrone 1994) was performed in margin of fossa (fig. 417) which areas (analogous to taxa) are classified by their 21. Denticles on genital lobe: (0) absent (fig. 421); (1) on shared taxa (analogous to characters) according to the margin (figs. 234-269, 417-418); (2) widespread (figs. most parsimonious cladogram. Data for PAE consist of 270-275, 419) area ϫ taxa matrices and the resulting cladograms rep- 22. Auricle denticles: (0) 2 or 3 (figs. 195-233, 430-433); (1) 4 to 5 (figs. 428-429) resent nested sets of areas of endemicity (Morrone & 23. Auricle shape (ventral view): (0) triangular or quadran- Crisci 1995). Cladistic information is then incorpo- gular (figs. 428-431); (1) narrow parallel-sided (figs. 432- rated by adding supraspecific monophyletic groups to 433) the matrix. PAE recognizes generalized tracks through 24. Basal fold on lateral lobe of vesica spermalis IVth seg- the discovery of nested sets of areas. In order to under- ment: (0) present (figs. 434, 436); (1) absent (fig. 435) take the PAE, species and species groups were coded for 25. Medio-longitudinal fold on ventral lobe of vesica spermalis IVth segment: (0) absent (figs. 434-435); (1) their absence (0) or presence (1) in each area of en- present (fig. 436) demism in the data matrix (table 3). The cladogram 26. Light blue spot on male sterna IX (posterior to genital obtained was rooted with a hypothetical area coded by opercula) and X: (0) absent (figs. 119-141b, 142c, 151- all zeros. Most parsimonious trees were sought using 156b); (1) present (figs. 143c-e, 144-150b) implicit enumeration (ie* command) in Hennig86 27. Conical tubercle on abdominal sternum I: (0) absent (fig. (version 1.5, Farris 1988). 421); (1) with denticles widespread (figs. 234-249, 252- 275); (2) with denticles restricted to apex (figs. 250-251) 28. Transverse ridge on abdominal sternum I: (0) absent; (1) Taxonomic analysis present (fig. 417) 29. Constriction of abdominal segment III: (0) marked (figs. Each species account includes synonymy, diagnosis, 119-120, 123, 125-156); (1) slight (figs. 121-122, 124) brief description, biology data, remarks, and distribu- 30. Subbasal tooth or carina in dorsal surface of male X seg- tion. Synonymies include only references to the taxon ment: (0) present (figs. 119-141b, 142c-d, 143-156b, 427a); (1) absent (fig. 426a) under consideration that incorporate descriptions, re- 31. Lateral carina of male cercus: (0) convex (figs. 390-395b, descriptions, illustrations or new distribution records. 426a); (1) linear (figs. 359-365); (2) concave (figs. 358, Wing venation terms and abbreviations follow Riek & 366-389, 427a) Kukalová-Peck (1984), as amended by Bechly (1996). 32. Subbasal tooth of male cerci: (0) absent (figs. 382-395, The term ‘hamuli’ used throughout text refers to the 426a); (1) present (small to prominent) (figs. 358-381, anterior pair of hamuli. Total length includes caudal 427a) 33. Apical spine ventrally directed projected from inner appendages, maximum width of the wings was taken margin of cercus: (0) absent (fig. 427a); (1) present (figs. at the level of the nodus, and the length of the 426a-c) pterostigma was measured from the anterior costal 34. Row of denticles on dorso-distal crest of cercus: (0) ab- end to the posterior anal end. All dimensions are in sent (fig. 426b); (1) present (Fig 427a-b) millimeters. The range of measurements was taken 35. Dorso-distal crest of cercus: (0) absent; (1) linear in dor- from 10 males and 10 females whenever available and sal view and evenly curved in lateral view (figs. 358-365, 370-383, 385-387-392, 394-395, 427a); (2) linear in dor- all specimens available of each species were examined sal view and angled in lateral view (figs. 366-369, 384, to establish the variability of characters. All diagnostic 388-389, 393); (3) curved in dorsal view (fig. 426c) characters were illustrated, including variation when 36. Distal concavity on cercus inner margin: (0) absent (fig. observed. Illustrations were made with the aid of a 427b); (1) present (fig. 426c) camera lucida, and are not to scale. 37. Subapical ventral process of inner margin of male cerci: The following acronyms are used under species ac- (0) absent (figs. 358-365, 370-395, 426-427); (1) present (figs. 366-369) counts, he: head, th: pterothorax, ab: abdomen, te: 38. Spines of epiproct tip: (0) small and directed dorsally (fig. ventral terga, ge: genital fossa, tg: ventral tubercle and 427c); (1) large and directed anteriorly (fig. 426d) genital lobe, ha: anterior hamuli, as: anterior lamina 39. Tip of cercus (lateral view): (0) in same plane with cercus spine, ce: cerci, wi: wings, au: auricle; ve: vesica (figs. 358-365, 370-381); (1) extreme apex bent ventrally spermalis distal segment, and Mp: map, and for coun- (figs. 366, 382-389); (2) whole tip ventrally bent (figs. tries in the plates, AR: Argentina, BO: Bolivia, BR: 367-369, 390-395). Brazil, CH: Chile, CO: Colombia, CR: Costa Rica, EC:

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Ecuador, GU: Guatemala, MX: Mexico, PE: Peru, PR: fig. 14a) or converging anteriorly (i.e. fig. 13a); fron- Puerto Rico, US: U.S.A. toclypeal groove with (i.e. fig. 17b) or without (i.e. fig. Little is known of the biology of most species of this 16b) a dark stripe, and fronto-ocular groove with a genus; I have extracted notes from specimen labels and dark stripe which may be of uniform width (i.e. fig. comments from collectors on the behavior of certain 35b) or widened toward point of contact between species. Under each species I included a tentative frontoclypeal groove and eyes (i.e. fig. 37b). ranking (rare; common) based on its occurrence and Thorax (figs. 39-118). – Extension and contour of abundance in collections and in the field. Full locality pale mesanepisternal, mesepimeral, and metepimeral data are given only for new or revalidated species; for stripes show intraspecific as well as interspecific varia- remaining species, country, state, and repository have tion, but can be used as diagnostic characters for some been indicated in the distribution lists. Full locality species (i.e. figs. 92, 95). Color of wing membranule data for those species are available from the author. can also aid in identification (i.e. dark except basal Asterisks in distribution lists indicate material that has 20% pale in R. bonariensis, pale except distal 20-33% been illustrated. Maps represent distribution records dark in R. absoluta). Wing venation provides no spe- from collections and reliable references. Distribution cific characters (table 1), but presence (i.e. fig. 396) or data for some common and well-known species in the absence (fig. 399) of crossveins in supratriangles, and U.S.A. (e.g. R. californica, R. dugesi, R. multicolor and level of beginning of two rows of cells between RP1 R. mutata) are not exhaustive. Maps were created elec- and RP2 with respect to pterostigma (i.e. proximal, fig. tronically from the Digital Chart of the World 402 vs. distal, fig. 399), are good characters for groups (1:1,000,000) using ArcView 3.1 (ESRI 1996). Eleva- of species. tion data and longitude/latitude coordinates were Abdomen (figs. 119-156). – Presence or absence culled from the Global Gazetteer website (http:// and confluence or separation of certain pale spots are www.calle.com/world/) and placed into a Microsoft sometimes diagnostic (i.e. figs. 133, 140), as well as FoxPro Data base linked to the ArcView program. presence or absence of pale or dark medio-longitudi- Acronyms used for collections are those given in nal stripes on female dorsum of segment II (i.e. figs. the website ‘The insect and spider collections of the 131-132). Useful structural characters of the sec- world http://hbs.bishopmuseum.org/codens/codens- ondary genitalia include: anterior margin of ventral r-us.html and codens not included there are as fol- tubercle of segment II concave (i.e. fig. 267), linear lows: ABMMC – Dr. A. B. M. Machado personal col- (i.e. figs. 264-265), or convex (i.e. figs. 261-262); gen- lection, Belo Horizonte, Brazil; GSVC – Dr. Graham ital lobe low (i.e. fig. 237) or high (i.e. fig. 240), with S. Vick personal collection, Hans, U.K.; IADIZA – In- denticles on margin (i.e. fig. 268) or spread over later- stituto de Investigaciones de Zonas Áridas, Mendoza, al surface as in punctata group (figs. 270-275); anteri- Argentina; IEUM – Instituto de Entomología, Univer- or lamina spine wide (i.e. fig. 317) or narrow (i.e. fig. sidad Metropolitana de Ciencias de la Educación, 325), short (i.e. fig. 354) or long (i.e. fig. 350), trian- Santiago, Chile; KJTC – Dr. Kenneth J. Tennessen gular (i.e. fig. 355) or needle-shaped (figs. 356-357), personal collection, Florence, U.S.A.; WBC – Dr. Wil- with dorsal margin linear (i.e. fig. 330) or concave (i.e. fried Braun personal collection, Berlin, Germany. fig. 332), tip directed dorsally (i.e. fig. 326) or ventral- ly (i.e. fig. 325); anterior hamuli with long anterior Characters process in Marmaraeschna group (i.e. figs. 277-284), Discrimination among species of this group is dif- short in remaining species (i.e. fig. 288), with pointed ficult since combinations of characters must be used (i.e. fig. 311) or rounded tip (i.e. fig. 312), dorsal fold to identify most species. The following characters are short (i.e. fig. 312) or long (i.e. fig. 311). Ventral ter- useful in evaluating specific status: ga (delimited by inner and outer latero-ventral longi- Head (figs. 1-38). – Contour of frons in lateral view tudinal and postero-ventral transverse carinae) can be dorsally projected (fig. 1b) or not (i.e. fig. 2b); frontal narrow (i.e. fig. 181) or wide (i.e. fig. 180), and later- carina linear to convex (i.e. fig. 7c) or with a median al carinae can be linear (i.e. fig. 191) or sinuous (i.e. concavity in frontal view (i.e. figs. 2, 4c), anteriorly fig. 192). Male cerci (figs. 358-395) with dorso-distal angled (fig. 18a) or smoothly curved in dorsal view crest low (i.e. fig. 384) or high (i.e. fig. 382), short (i.e. (i.e. fig. 19a), and located at anterior margin of frons fig. 364) or long (i.e. fig. 365), smoothly curved (i.e. (i.e. fig. 8b) or recessed into horizontal portion of fig. 393) or angled (i.e. fig. 392); sub-basal tooth ab- frons (i.e. fig. 9b); clypeal lobes angled (i.e. fig. 14c) or sent (i.e. fig. 382) or present (i.e. fig. 380), low (i.e. fig. rounded (i.e. fig. 16c); frons and vertex shorter (i.e. 363) or prominent (i.e. fig. 364); lateral margin linear fig. 24a) or longer (i.e. fig. 22a) than interocular su- (i.e. fig. 359), concave (i.e. fig. 358) or convex (i.e. fig. ture and occipital triangle; T-spot of frons complete 390) in lateral view; tips pointed (i.e. fig. 391) or blunt (i.e. fig. 2a) or incomplete (i.e. fig. 4a), sides of stem (i.e. fig. 393), ventrally bent (i.e. figs. 367, 385, 390) wide (i.e. fig. 34a) or narrow (i.e. fig. 33a), parallel (i.e. or in the same plane as cerci (i.e. fig. 359); inner mar-

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gin with (figs. 366-369) or without (i.e. fig. 370) a occipital triangle of moderate length, measured as tri- ventro-distal process. Female cerci tips angled (i.e. fig. angle length/interocular suture length: 0.25-0.65 (fig. 368) or rounded (i.e. fig. 366), with a long apical 415) (shorter in Anaciaeschna; fig. 416); bend of MA spine (i.e. fig. 358), mucronate (i.e. fig. 364), incised prominent (figs. 396-404, 406) (slight in Oreaeschna, (i.e. fig. 395), or lacking such structures (i.e. fig. 365); Aeshna s.str. and ‘A.’ isoceles; figs. 405, 407); IRP2 fork sides parallel (i.e. fig. 365), diverging (i.e. fig. 364), or symmetrical (figs. 396-404) (asymmetrical with inferi- converging (i.e. fig. 383) distally. or branch as a continuation of IRP2 stem in Ore- aeschna, Aeshna s.str., Andaeshna, fig. 405; asymmetri- cal with upper branch as a continuation of IRP2 stem TAXONOMIC PART in Anaciaeschna, fig. 406); anal triangle with three cells (figs. 396-404, 406-407) (two in Aeshna s.str. except Rhionaeschna Förster for constricta group, fig. 405); anal angle angled (figs. Rhionaeschna Förster, 1909: 220-223 (diagnosis, R. maita 396-406, 408-412) (rounded in Andaeshna and ‘A.’ only species included); Martin 1911: 13 (generic diagno- isoceles, 407, 413), with basal and distal veins parallel sis from Förster); Ris 1913: 83 (indication of close rela- and fused for a distance before angle (figs. 408-412) tionship of maita with variegata and multicolor groups); Hincks 1934: 80 (Peru): Schmidt 1952: 256 (Peru). (fused at a single point in Andaeshna and ‘A.’ isoceles, Neureclipa Navás, 1911: 476, 478 (brief description of fig. 413); membranule length 50-66% of wing margin genus). Syn. n. (figs. 408-411) (75-100% of wing margin in Anaci- Aeshna (Hesperaeschna) Cockerell, 1913: 581 (brief descrip- aeschna, Andaeshna and ‘A.’ isoceles, figs. 412-413); tion of subgenus in key; type species Aeshna (Hesper- hamular fold separated from wall of genital fossa (figs. aeschna) californica); Calvert 1952: 253-255 (addition of subgeneric characters and species); Calvert, 1956: 10 (in 423-425) (not differentiated from wall of genital fossa key), 21-22 (subgeneric diagnosis), 217-218 (discussion in Andaeshna and constricta group of Aeshna s.str., fig. of origin), 222-224 (discussion of relationships). 422). Genital lobe elevated above rest of ventral mar- Aeshna (Marmaraeschna) Calvert, 1952: 256 (characters of gin of tergum I (figs. 418-419, 421) (at same level as subgenus, intricata designated as subgeneric type); ventral margin of tergum I in Anaciaeschna, fig. 417); Calvert, 1956: 10 (in key), 109-110 (subgeneric diagno- auricle triangular or quadrangular (figs. 428-431) sis), 217-218 (discussion of origin), 222-224 (discussion of relationships). (narrow parallel-sided in Anaciaeschna and ‘A.’ isoceles, Aeshna (Schizuraeschna) Calvert, 1952: 256-257 (characters figs. 432-433), with two or three denticles (usually of subgenus, multicolor designated as subgeneric type); four or five denticles in Aeshna s.str.); basal fold of pos- Calvert, 1956: 10 (in key), 100-102 (subgeneric diagno- terior lobes of vesica spermalis IVth segment present sis), 217-218 (discussion of origin), 222-224 (discussion (figs. 434, 436) (absent in Andaeshna and ‘A.’ isoceles, of relationships). fig. 435); and dorso-distal crest of cercus present (figs. Aeshna (Neureclipa): Calvert, 1952: 257-258 (relegated to 358-395) (absent in , ‘ .’ ‘ ’ subgeneric rank, bonariensis designated as type species, ad- Oreaeschna A affinis, A. mixta, dition of subgeneric characters); Calvert 1956: 10 (in key), ‘A.’ brevistyla, and ‘A.’ ellioti, Andaeshna, Anaci- 125-126 (subgeneric diagnosis), 217-218 (discussion of aeschna, and ‘A.’ isoceles). origin), 222-224 (discussion of relationships). The remaining Aeshninae genera, Castoraeschna, Aeshna (Rhionaeschna): Calvert, 1956: 11 (in key), 98-99 Coryphaeschna and Remartinia, can be easily distin- (relegated to subgeneric rank, subgeneric diagnosis), 217- guished from Rhionaeschna by the elongated discal tri- 218 (discussion of origin), 222-224 (discussion of rela- tionships). angles (width/length 0.35 or less) (shorter in Rhion- Type species. – Rhionaeschna maita Förster, 1909 (=R. brev- aeschna, width/length 0.40 or more), male anal ifrons (Hagen 1861)) by monotypy. triangle with two cells, and distal segment of vesica spermalis lacking lateral and ventral folds and bearing Diagnosis patches of michrotrichiae. Coryphaeschna and Remar- The autapomorphy characterizing Rhionaeschna is tinia have a smooth abdominal sternum I, while Cas- the presence of a conical tubercle bearing denticles on toraeschna has a cylindrical tubercle lacking denticles abdominal sternum I (figs. 234-275, 418-419); ster- accompanied in the male by a pair of medio-posterior num I is planar in Oreaeschna, Aeshna sensu stricto, projections of the ventral tergum I (fig. 420). several ‘Aeshna’ species of uncertain position ‘A.’ affi- nis, ‘A.’ mixta, ‘A.’ brevistyla Rambur, 1842, ‘A.’ ellioti Remarks Kirby, 1896, ‘A.’ williamsoniana Calvert, 1905, ‘A.’ The oldest available name for the monophyletic moori Pinhey, 1981, ‘A.’ rileyi Calvert, 1892 and ‘A.’ group comprising the Neotropical species of ‘Aeshna’ subpupillata McLachlan, 1896, and Andaeshna, and is Rhionaeschna Förster, 1909, in spite of the different has a transverse ridge in ‘Aeshna’ isoceles Müller, 1767 original definition of the genus. Rhionaeschna (from and Anaciaeschna (fig. 417). Other characters that dif- Greek rhion = a jutting part of a mountain, peak, ferentiate it from related Aeshnidae genera are: T-spot promontory, headland) was erected by Förster (1909) of frons present (absent in Andaeshna and ‘A.’ isoceles); to include R. maita, and its distinction from Aeshna

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was based in following structural characters: frons dominal segment X, and anal triangle with three cells). wide (ratio head/frons width 1.5), dorsal margin of Finally Calvert (1952) erected another two subgen- frons rounded, and abdomen slightly constricted at era for Aeshna: Marmaraeschna and Schizuraeschna. segment III. However, the ratio head/frons width is in- Marmaraeschna, with M. intricata as generotype, was traspecifically variable, and a wide frons is found diagnosed by the pterothoracic marbled color pattern, among several species of Rhionaeschna (i.e. R. brev- abdominal segment I with a ventral tubercle, supratri- ifrons: 1.5-1.9 mm, R. fissifrons: 1.6-2.1 mm, R. intri- angular cross veins present, vein IRP1 beginning proxi- cata: 1.7-2.4 mm, R. pallipes: 1.7-2.5 mm, R. confusa: mal to the level of the pterostigma or under the proxi- 1.6-2.5 mm, R. marchali: 1.5-1.9 mm, R. peralta: 1.7- mal end or proximal half of the pterostigma, anal 1.85 mm, R. tinti: 1.7-2.2 mm, R. variegata: 1.6-2.3 triangle 3-celled, male with a middorsal longitudinal mm, R. absoluta: 1.6-2.3 mm, R. bonariensis: 1.7-2.4 carina on segment X and male cerci with the apex not mm, R. diffinis: 1.4-2.3 mm, R. elsia: 1.7-2.2 mm). A bifid nor with an anteapical ventral tubercle. Schizu- rounded frons is characteristic of several species (R. raeschna, including A. multicolor (generotype), A. mu- brevicercia, R. brevifrons, R. fissifrons, R. intricata, R. tata, A. dugesi and A. jalapensis, was characterized by obscura, R. pallipes, and R. vigintipunctata) and the abdominal segment I with a ventral tubercle, supratri- slightly constricted abdomen is found in R. brevifrons, angular cross veins present, thorax dark with two later- R. fissifrons and R. pallipes. My examination of the lec- al oblique pale stripes, vein IRP1 beginning distal to the totype of R. maita demonstrated that Rhionaeschna level of the pterostigma or under the distal end, dis- maita Förster, 1909 is a junior subjective synonym of coidal triangles with the formula 2, 1, 1, inner triangles R. brevifrons (Hagen, 1861). two celled, anal triangle with three cells, and male cer- Four other names, one originally for a genus and ci in lateral view bifid in the apical fourth or less, the three for subgenera, have been assigned to some of the lower division much shorter than the upper and in one species which I presently include in Rhionaeschna. species reduced to an anteapical point. Resolution of these names is given as follows. In the present synopsis, some of these names are Neureclipa was originally created by Navás (1911) used in an informal sense to make reference to certain to include Aeshna bonariensis Rambur, 1842 and A. species-groups, but I do not recognize subgenera. litigatrix Navás, 1911 and was characterized by the ab- sence of cross veins in the supratriangle, abdominal Keys segment X of male with a dorsal tooth, and male cerci bearing a subbasal tooth. The subgenus Hesperaeschna Comparison with illustrations and diagnosis is ad- was created by Cockerell (1913) to include A. califor- vised to aid in identifications especially when trying to nica Calvert, 1905. The diagnostic characters included identify poorly preserved or incomplete material (i.e. in his key were: RP3-4 and MA separated by one cell specimens with bad color preservation, with broken only at wing margin, but a short distance before by cerci or lacking head). two rows of cells, owing to the deflection of MA from the straight course; cell formula of triangles 2, 1, 1; up- Key to males of Rhionaeschna per branch of radial sector in a line with stem; IRP2 separated from supplementary vein below by only 1. Pterothorax and abdominal segments VII-X with- three rows of cells; IRP2 fork a short distance before out pale markings (figs. 39, 119a-b); hamulus level of beginning of pterostigma. with medial digit-like projection on antero-ven- Calvert (1952) relegated Neureclipa to subgeneric tral margin (fig. 276); frons projected dorsally, rank, fixed A. bonariensis as its type including besides higher than vertex in lateral view (fig. 1b); Aeshna bonariensis, A. diffinis, A. elsia and A. galapa- Venezuelan tepuis (fig. 439)...... draco goensis, and added as subgeneric characters abdominal – Pterothorax with or without pale stripes, abdom- segment I with a ventral tubercle, vein IRP1 beginning inal segments IV-VIII to X with pale spots (figs. 41- at distal end of pterostigma, thorax with two lateral 118, 120-156a-b); hamulus with no medial pro- oblique pale stripes, discoidal triangles with the for- jection on antero-ventral margin (figs. 277-315); mula 1, 1, 1, inner triangles usually with one cell, anal frons not projected dorsally, lower than vertex triangle with three cells, and male cerci in lateral view (figs. 2-38b)...... 2 not bifid nor with an anteapical point. He also listed 2. Pterothorax with marbled pattern of black mark- more characters to differentiate Hesperaeschna species ings and pale areas (figs. 40-47); hamular anteri- from the European species of Aeshna (presence of a or process long: distance between ventral tip of ventral tubercle in the first abdominal segment; supra- process and axis of hamulus (A) longer than dis- triangular crossveins; thorax dark with two pale lateral tance between dorsal end of hamular fold and stripes; male cerci in lateral view not bifid nor with an axis (B) (figs. 277-284) ...... Key M-1 anteapical ventral tubercle; dorso-medial carina in ab- – Pterothorax uniform in color or with pale stripes

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or spots (figs. 48-118); hamular anterior process – Cerci long: 4.9-6.5 mm (figs. 359, 363-365); short: distance between ventral tip of process and constriction of abdominal segment III marked axis of hamulus (A) shorter than distance between (figs. 120, 123, 125-126)...... 4 dorsal end of hamular fold and axis (B) (figs. 285- 2. Frontal carina linear in frontal view (fig. 7c); Ar- 315)...... 3 gentina (Buenos Aires and Mendoza north to 3. Sterna X and IX posterior to genital opercula with Salta province) (fig. 441)...... pallipes bright light-blue spot (figs. 143-150b); surface of – Frontal carina concave in frontal view (figs. 3-4c) abdominal ventral terga delimited by inner and ...... 3 outer carinae narrow (width measured at narrow- 3. Frontal carina with a slight medial concavity (fig. est point/length of ventral terga IV less than 0.1 3c); inner and outer carina of ventral terga VI sin- except for R. vazquezae (fig. 188) and R. pauloi uous (fig. 159a-b); black spots of pterothorax re- (fig. 185) with widths of 0.13 mm and 0.18 mm, duced to 1 on mesanepisternum, 3 on mesepi- respectively) (figs. 181-188); HW with two rows meron-metepisternum, and 1 on metepimeron of cells between RP1 and RP2 beginning under (fig. 41-42); cercus with sub-basal tooth low (figs. pterostigma or proximal to it (fig. 401-402); sub- 360b, e-f); Valdivia province in Chile north to basal tooth of cercus absent and extreme tip bent Peru (fig. 440) ...... brevifrons ventrally (figs. 382-389b)...... Key M-2 – Frontal carina with a deep medial cleft (fig. 4c); – Sterna of X and IX posterior to genital opercula inner and outer carina of ventral terga VI linear, yellow or pale brown to black (figs. 127-142, 151- parallel (fig. 160a); black spots of pterothorax: 2-3 156b); surface delimited by inner and outer cari- on mesanepisternum, 4-5 on mesepimeron-met- nae of abdominal ventral terga wide (width mea- episternum, and 3 on metepimeron (fig. 43); cer- sured at narrowest point/ length of ventral terga IV cus with sub-basal tooth prominent (fig. 361b); larger than 0.13) (figs. 165-180, 189-194); HW northwestern Argentina north to Peru (fig. 440).. with origin of two rows of cells between RP1 and ...... fissifrons RP2 variable; cercus with other combination of 4. Sub-basal tooth of cercus low (figs. 359B, 363b, characters (figs. 366-381, 390-395b) ...... 4 d); abdominal segment I with black spot extending 4. Inner margin of cercus with a ventral process at to tip of auricles in ventral view (figs. 196, 199); distal 25%, cercus with dorso-distal crest triangu- smaller species (total length 66-73.5 mm)...... 5 lar in lateral view, sub-basal tooth low and blunt – Sub-basal tooth of cercus prominent (figs. 364- (figs. 366-369) ...... Key M-3 365b); black spot not extending to tip of auricles – Cercus without ventral process on inner margin, in ventral view (figs. 200, 202); larger species (to- dorso-distal crest and sub-basal tooth variable tal length 72-80.5 mm)...... 6 (figs. 370-381, 390-395)...... 5 5. Stem of T-spot strongly narrowed at its conflu- 5. Cercus with entire tip ventrally bent in lateral ence with transverse arms (fig. 5a); ventral tuber- view; sub-basal tooth absent; cercus in dorsal cle of abdominal segment I high (about as high as view gradually widening to distal end (maximum 50% of its length in lateral view) (fig. 238); Peru width at distal 12-16%) (figs. 390-395), denticles to Ecuador (fig. 442)...... intricata of genital lobe small, numerous, and spread over – Stem of T-spot widely confluent with transverse ventral and lateral surfaces of lobe (figs. 270-275) arms (fig. 2a); ventral tubercle of abdominal seg- ...... Key M-4 ment I low (about as high as 30% of its length in – Cercus tip not ventrally bent, sub-basal tooth lateral view) (fig. 236); Ecuador to Venezuela present (but vestigial in R. californica and R. con- (fig. 442)...... brevicercia fusa); cercus in dorsal view variable (figs. 370- 6. Stem of T-spot strongly narrowed at its conflu- 381); denticles of genital lobe confined to ventral ence with transverse stem, vertex yellow with pos- surface of lobe (figs. 247-258)...... 6 terior 30% black (fig. 8a); genital lobe approxi- 6. Supratriangles usually free, sometimes one or two mately triangular in lateral view (fig. 242); supratriangles with 1-2 cross veins (in 2-10% of dorso-distal crest of cercus extended along distal specimens examined) (fig. 399)...... Key M-5 30% of cercus (fig. 365b); northwestern Argenti- – Supratriangles with 1-3 cross veins, exceptionally na to Peru (fig. 443)...... vigintipunctata one supratriangle without cross veins (in 1% of R. – Stem of T-spot widely confluent with transverse brasiliensis and R. variegata specimens examined) stem, vertex black with a small rounded yellow (fig. 400)...... Key M-6 spot at each anterolateral edge (fig. 6a); genital lobe subquadrate in lateral view (fig. 240); dorso- Key M1 distal crest of cercus extended along distal 25% of 1. Cerci short: 3.5-4.8 mm (figs. 360-362); con- cercus (fig. 364b); Bolivia north to Peru (fig. striction of abdominal segment III slight (figs. 443)...... obscura 121-122, 124)...... 2

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Key M-2 na (fig. 453) ...... haarupi 1. Dorso-distal crest of cercus rising gradually, an- 7. Pale mesepimeral stripe narrowed at midlength to gulate and lower than base of cercus in lateral 50% of its basal width (figs. 101-102); anterior view (figs. 384, 388, 389b) ...... 2 side of ventral tubercle concave (fig. 267); dorso- – Dorso-distal crest of cercus rising abruptly, distal crest of cercus gradually rising at base (fig. smoothly convex and higher than base of cercus 385b); Venezuelan tepuis (fig. 454) ...... nubigena in lateral view (figs. 382-383, 385-387b) ...... 4 – Pale mesepimeral stripe approximately uniform 2. Anterior lamina spine well developed (fig. 349); in width (figs. 92-94); anterior and posterior pale mes- and metepimeral stripes of thorax wide sides of ventral tubercle convex (figs. 261-262); with anterior (upper) portion as wide as 50-66% dorso-distal crest of cercus abruptly rising at base of their respective sclerites (figs. 107-109); dark (fig. 382b, g); Bolivia north to Mexico (fig. 454) stripe on frontoclypeal groove (figs. 32b-c); ...... cornigera Guerrero and Nayarit states in Mexico (fig. 456) ...... vazquezae Key M-3 – Spine of anterior lamina reduced (figs. 347-348); 1. Ventral process of inner margin of cercus small, pale mes- and metepimeral stripes relatively nar- not projected posteriorly (fig. 366b); tip of ante- row, as wide at anterior portion as 30-50% of rior lamina spine directed ventrally (fig. 325); their respective sclerites (figs. 97-99, 106); no Mexico to southern USA (fig. 444)...... dugesi dark stripe on frontoclypeal groove (figs. 27, 32b- – Ventral process of inner margin of cercus project- c) ...... 3 ed posteriorly (figs. 367-369b); tip of anterior lam- 3. Inner and outer margins of cercus approximately ina spine curved dorsally (figs. 326-328)...... 2 parallel to each other along medial 30% (fig. 2. No pale abdominal PL spots in segments IV-V, PD 388a); pale thoracic stripes bright green, spots small (fig. 128a-b); only extreme tip of cer- mesanepisternal stripe complete (fig. 106); north- cus curved ventrally (fig. 367c); Costa Rica to ern Argentina to southern USA (fig. 455)...... psilus Mexico (fig. 444) ...... jalapensis – Inner and outer margins of cercus diverging grad- – Pale abdominal PL spots present in segments IV-V ually along medial 30% (fig. 384a); pale thoracic and confluent with large PD spots (figs. 129- stripes light blue, mesanepisternal stripe incom- 130a-b); entire tip of cercus curved ventrally plete, as long as 50% of mesanepisternum (figs. (figs. 368-369c) ...... 3 97-99); Baja California state in Mexico (fig. 456) 3. Distance between posterior base of ventral process ...... manni and base of dorsal crest of cercus (A) larger than 4. Pale mesepimeral and metepimeral stripes with distance between it and tip of cercus (B) (fig. deep rounded indentations (figs. 95-96, 103- 369c); ventral tubercle of abdominal segment I 105); abdominal PL spots large (figs. 144b, 147- low (about as wide as 3 times its height) (fig. 245); 148b)...... 5 base of T-spot stem wider than vertex; vertex black – Pale mesepimeral and metepimeral stripes lacking with anterior 30% pale (fig. 12a); eastern USA to indentations (figs. 92-94) or with broad and shal- southern Canada (fig. 445)...... mutata low indentations (figs. 101-102); abdominal PL – Distance between posterior base of ventral spots small to absent (figs. 143a-c, 146b)...... 7 process and base of dorsal crest (A) shorter than 5. Ventral tubercle of abdominal segment I lower distance between it and tip of cercus (B) (fig. than 30% of its length (fig. 266); cercus in dorsal 368c); ventral tubercle of abdominal segment I view with tip directed posteriorly, distal 60% of high (about as wide as 2 times its height) (fig. cercus parallel-sided (fig. 387a); Paraná and Mi- 244); base of T-spot stem narrower than vertex, nas Gerais states in Brazil (fig. 453) ...... pauloi vertex pale with posterior 30% black (fig. 11a); – Ventral tubercle of abdominal segment I about as Mexico to southern Canada through western USA high as 50% of its length (figs. 259, 264-265); (fig. 445) ...... multicolor cercus variable (figs. 383, 386a) ...... 6 6. Stem of T-spot of frons gradually narrowed distal- Key M-4 ly (fig. 30); pale mesanepisternal stripes complete 1. Black stripe on frontoclypeal groove strongly (figs. 103-105), as wide as 50% of mesan- widened toward eyes (figs. 37-38); pale pterotho- episternum; Argentina north to Venezuela (fig. racic stripes complete, linear, and narrower than 453) ...... planaltica 50% of their sclerites (114, 118); genital lobe – Stem of T-spot of frons abruptly narrowed distal- prominent, higher than 30% of its width in later- ly (fig. 26); pale mesanepisternal stripes complete al view (figs. 274-275); tip of hamular anterior (fig. 95) or incomplete (fig. 96), as wide as 25- process blunt in ventral view (unknown in deces- 30% of mesanepisternum; northwestern Argenti- sus) (figs. 314-315)...... 2

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– Black stripe on frontoclypeal groove not strongly state in Venezuela (fig. 457)...... condor widened toward eyes or absent (figs. 33-36); pale – Stem of T-spot narrower than vertex (fig. 36a); pterothoracic stripes with deep indentations, di- pale mesepimeral and metepimeral stripes as wide vided into spots, or wider than 50% of their scle- as 25-33% of their sclerites with an anterior and rites (figs. 110-113, 115-117); genital lobe lower a posterior indentation (figs. 115), or divided than 30% of its width in lateral view (figs. 270- into spots (figs. 116-117); anterior tip of hamular 273); tip of hamular anterior process pointed in process in posterior view not recurved (fig. 312); ventral view (figs. 310-313) ...... 4 Bolivia to Colombia (fig. 458) ...... joannisi 2. Anterior lamina spine absent; labrum, clypeus and frons orange; pale abdominal PL spots present Key M-5 on segments IV-IX, fussed with ML on IV-VIII; 1. Clypeal lobes rounded (figs. 16-17c); dorsal mar- known only from Itatiaia, Rio de Janeiro state, gin of anterior lamina spine concave (figs. 332- Brazil (fig. 459) (characters abstracted from 333); ventral tubercle of abdominal segment I Calvert 1953) ...... decessus with a few denticles restricted to its apex (fig. – Anterior lamina spine present (figs. 355-357); 250-251)...... 2 labrum, clypeus and frons light blue or yellow; – Clypeal lobes angled (figs. 13-15c); dorsal margin pale abdominal PL spots absent on segments VI- of anterior lamina spine linear (figs. 329-331); VII, separated from ML on IV-V, fused or not with ventral tubercle of segment I with numerous den- ML on VIII (figs. 155-156b) ...... 3 ticles spread over its posterior surface (fig. 247- 3. Ventral tubercle of abdominal segment I low and 249)...... 3 smoothly curved in lateral view (fig. 275); black 2. No dark stripe on frontoclypeal groove (fig. 16b- of T-spot not extended on antefrons; vertex yel- c); dorso-distal crest at distal 25% of cercus low with lateral edges black (fig. 38); São Paulo, abruptly rising at its base (fig. 373b); tip of Rio de Janeiro, Espirito Santo and Minas Gerais hamular anterior process pointed in ventral view states in Brazil (fig. 459) ...... punctata (fig. 292b); smaller species (total length 49-54 – Ventral tubercle of abdominal segment I high mm, male cerci 4.3-4.6 mm); central Chile to and trapezoidal in lateral view (fig. 274); black of Ecuador (fig. 448) ...... elsia T-spot extended on dorsal surface of antefrons; – Dark stripe on frontoclypeal groove (fig. 17b-c); vertex yellow with lateral and posterior 30% dorso-distal crest at distal 30% of cercus gradual- black (fig. 37); Minas Gerais state in Brazil (fig. ly rising at its base (fig. 374b); tip of hamular an- 459)...... eduardoi terior process rounded in ventral view (fig. 293b); 4. Dorso-distal crest of cercus smoothly convex in larger species (total length 58-60 mm, male cerci lateral view (fig. 393b); anterior lamina spine as 5 mm); Galápagos Islands in Ecuador (fig. 448) . long as 30% of its basal width (fig. 353-354); ...... galapagoensis ventral portion of hamuli (A) shorter than hamu- 3. Black stripe on fronto-ocular groove narrow and lar fold (B) (fig. 311a); Táchira State in uniform in width, frontoclypeal groove not bor- Venezuela (fig. 458) ...... demarmelsi dered by black stripe (fig. 14b-c); outer base of – Dorso-distal crest of cercus angulate in lateral cercus with a pale spot (fig. 371a-b); membranule view (figs. 390-392b); anterior lamina spine as dark except basal 20% pale; stem of T-spot of long as 2 times or more its basal width (figs. 350- frons approximately parallel-sided (fig. 14a); Ar- 352); ventral portion of hamuli (A) longer than gentina to Brazil (fig. 447)...... bonariensis hamular fold (B) (figs. 310, 312-313) ...... 5 – Black stripe on fronto-ocular groove widening to- 5. Transverse arms of T-spot represented by a fine wards frontoclypeal groove, frontoclypeal groove line (fig. 33a); labrum orange, clypeus and frons bordered by black stripe (figs. 13, 15a); mem- bright yellow, no dark stripe on frontoclypeal branule pale except distal 20-33% dark; cercus groove (figs. 33b-c); auricle bearing two teeth without pale spots (figs. 370, 372a-b); stem of T- (fig. 228); Peru to Ecuador (fig. 457) ...... biliosa spot of frons gradually widened basally (figs. 13, – Transverse arms of T-spot wide (figs. 34, 36a); 15a) ...... 4 labrum pale brown, clypeus and frons light blue 4. Ventral tergum IV of abdomen relatively narrow, to light green, dark stripe on frontoclypeal groove ratio length/maximum width of abdominal terga (figs. 34, 36b-c); auricle bearing three teeth (figs. IV: 4.2-4.6; basal 30% of outer lateral carinae of 229, 231)...... 6 segment IV linear to concave (fig. 169a); southern 6. Stem of T-spot wider than vertex (fig. 34a); pale Argentina to Ecuador (fig. 446) ...... absoluta pterothoracic stripes as wide as 50-66% of their – Ventral tergum IV of abdomen relatively wide, ra- sclerites (fig. 112); anterior tip of hamular process tio length/maximum width of abdominal terga in posterior view recurved (fig. 313); Táchira IV: 2.4-2.6, basal 30% of outer lateral carina of

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segment IV convex (fig. 171a); southern and cen- equal to 2) (fig. 24a-b); distal portion of ventral tral Chile and southwestern Argentina (fig. 447). margin of anterior lamina spine convex (fig. 339); ...... diffinis larger species (total length 57.8-67.9 mm, cercus length 4.45-5.85 mm); Chile and Argentina (fig. Key M-6 452)...... variegata 1. Subbasal tooth of cercus vestigial (figs. 376- – Confluence of eyes short (frons + vertex/eyes + 377b)...... 2 occipital triangle larger than 1; eyes/occipital tri- – Subbasal tooth of cercus well developed (figs. angle less than 2) (fig. 22a); distal portion of ven- 375, 378-381b)...... 3 tral margin of anterior lamina spine linear (fig. 2. Pale thoracic mesanepisternal stripes complete 338); smaller species (total length 51.75-56.5 (fig. 81); outer base of cercus with a pale spot (fig. mm, cercus length 3.75-4.5 mm); Bolivia and 376a-b); membranule dark except basal 20% Peru (fig. 452) ...... peralta pale; abdominal segments V-IX with PL spot (fig. 138b); stem of T-spot of frons approximately Key to females of Rhionaeschna parallel-sided (fig. 20a); frontoclypeal groove lacking a black stripe (figs. 20b-c); Argentina and Females of R. condor, R. decessus, and R. vazquezae Chile north to Brazil (fig. 450)...... confusa are unknown, but I tentatively include them here – Pale thoracic mesanepisternal stripes incomplete based on characters shared by males and females in (figs. 79-80); cercus without pale spots (fig. 377a- other species. b); membranule pale except distal 30-50% dark; 1. Pterothorax and abdominal segments VII-X with- abdominal segments V-IX without PL spot (fig. out pale markings (figs. 39, 119c); frons projected 137b); stem of T-spot of frons gradually widening dorsally, higher than vertex (fig. 1b); cercus tip basally (fig. 19a); frontoclypeal groove with a black bearing spine (fig. 358c, not to be confused with stripe (figs. 19b-c); Baja California state in Mexico mucron, i.e. figs. 359c, 360h, 361-362c, 363c, north to southwestern USA (fig. 449).....californica 364c, 385c,d); Venezuelan tepuis (fig. 439)..draco 3. Pterothorax without pale markings or with small – Pterothorax with or without pale stripes, abdom- rounded pale spots at ventral and/or dorsal edges inal segments IV-VIII to X with pale spots (figs. 40- of pleura (figs. 86-90); membranule dark except 118, 120-156c-d); frons not projected dorsally, basal 20% pale; subbasal tooth of cercus promi- lower than vertex (figs. 2-38b); cercus tip lacking nent and with acute angle (fig. 380b); northern spine (figs. 359c, 360c, g-h, 361-362c, 363c, Chile (fig. 451)...... tinti 364-365c, 366-369d, 370-395c)...... 2 – Pterothorax with pale stripes (figs. 76-78, 82-85, 2. Pterothorax with marbled pattern of black mark- 93); membranule pale except distal 30-50% dark; ings and pale areas (figs. 40-47)...... Key F-1 subbasal tooth of cercus with orthogonal angle – Pterothorax uniform in color (figs. 59, 67, 90) or (figs. 375, 378-379, 381b)...... 4 with pale stripes or spots (figs. 48-58, 60-66, 68- 4. Frontal carina in dorsal view angled anteriorly 89, 91-118)...... 3 (fig. 18a); pale metepimeral stripe constricted or 3. Surface of abdominal ventral terga delimited by in- divided into a ventral rounded spot and a dorsal ner and outer carinae narrow (width measured at T-shaped spot (figs. 76-78); southeastern Brazil narrowest point/length of ventral terga IV less than (fig. 450)...... brasiliensis 0.09) (figs. 181-188B); HW with two rows of cells – Frontal carina in dorsal view evenly curved (figs. between RP1 and RP2 beginning under pterostigma 21, 23-24a); pale metepimeral stripe complete or proximal to it (figs. 401-402)...... (figs. 82-84, 93)...... 5 ...... Key F-2 5. Frontoclypeal groove lacking a black stripe (fig. – Surface delimited by inner and outer carinae of 21b-c); anterior process of hamulus prominent, abdominal ventral terga wide (width measured at projecting as high as 50% of its width (fig. 297b); narrowest point/length of ventral terga IV larger cercus with a marked dorsal constriction at distal than 0.10) (figs. 165-180, 189-194B); HW with 25% (fig. 378a); Bolivia to Venezuela (fig. 451) .. origin of two rows of cells between RP1 and RP2 ...... marchali variable ...... 4 – Frontoclypeal groove with a black stripe (figs. 22, 4. Two rows of cells between RP1 and RP2 in HW be- 24b-c); anterior process of hamulus not promi- ginning under pterostigma or proximal to it (fig. nent, projecting only as high as 30% of its width 403) ...... Key F-3 (fig. 298-299b); cercus with a slight dorsal con- – Two rows of cells between RP1 and RP2 in HW be- striction at distal 25% (figs. 379, 381a) ...... 6 ginning at level of pterostigma distal end or distal 6. Confluence of eyes long (frons + vertex/eyes + oc- to it (figs. 398-400)...... 5 cipital triangle less than 1; eyes/occipital triangle 5. Frontoclypeal groove lacking a dark stripe (figs.

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9-12, 14, 16, 20-21b-c)...... Key F-4 Key F-2 – Frontoclypeal groove with a dark stripe (figs. 13, 1. Pale mesanepisternal and mesepimeral stripes 15, 17-19, 22-24b-c)...... Key F-5 complete and as wide as 60% of their sclerites (figs. 108-110); Guerrero and Nayarit states in Key F-1 Mexico (fig. 456)...... vazquezae 1. Constriction of abdominal segment III slight – Pale mesanepisternal stripes complete or incom- (figs. 121-122, 124c-d); cercus length: 1.7-2.3 plete, mesepimeral complete, as wide as 30-50% mm (figs. 360c, g-h, 361-362c) ...... 2 of their sclerites (fig. 95-107) ...... 2 – Constriction of abdominal segment III marked 2. Cercus longer than segments VIII-X and wider (figs. 120, 123, 125-126c-d); cercus length: 2.7- than 1.15 mm (figs. 384, 388c) ...... 3 6.5 mm (figs. 359, 363-365c) ...... 4 – Cercus shorter than segments VIII-X and as wide 2. Frontal carina linear in frontal view (fig. 7c); Ar- as or narrower than 1.1 mm (figs. 382-383, 385- gentina (Buenos Aires and Mendoza north to 387c) ...... 4 Salta province) (fig. 441)...... pallipes 3. Mesanepisternal stripes yellowish-light blue and – Frontal carina with a medial cleft in frontal view incomplete (fig. 99-100); Baja California state in (figs. 3-4c) ...... 3 Mexico (fig. 456) ...... manni 3. Cleft of frontal carina slight (fig. 3c); inner and – Mesanepisternal stripes bright green and com- outer carina of ventral terga VI sinuous (fig. plete (fig. 107); northern Argentina to southern 159c); black spots of pterothorax reduced to 1 on USA (fig. 455) ...... psilus mesanepisternum, 2 on mesepimeron, 1-2 on 4. Pale mesepimeral and metepimeral stripes with metepisternum and 1 on metepimeron (figs. 41- deep rounded indentations (figs. 97-98, 101, 42); Valdivia province in Chile north to Peru (fig. 104-106)...... 5 440) ...... brevifrons – Pale mesepimeral and metepimeral stripes lacking – Cleft of frontal carina deep (fig. 4c); inner and indentations (figs. 92-94) or with broad and shal- outer carina of ventral terga VI linear, parallel (figs. low indentations (figs. 101-102) ...... 7 160b); black spots of pterothorax represented by 5. Stem of T-spot of frons abruptly narrowed ante- 2-3 on mesanepisternum, 4-5 in mesepimeron riorly (fig. 26a); cercus as long as or slightly short- and 3 on metepimeron (Fig 43); northwestern Ar- er than abdominal segments IX-X (fig. 383c); gentina north to Peru (fig. 440) ...... fissifrons northwestern Argentina (fig. 453) ...... haarupi 4. Shorter cercus (2.7-3.7 mm) (figs. 355, 359c); – Stem of T-spot of frons gradually narrowed ante- smaller species (total length 66-73.5 mm)...... 5 riorly (figs. 29-30); cercus longer than abdominal – Longer cercus (4.9-6 mm) (figs. 360-361c); larg- segments IX-X (figs. 386c-e, 387c) ...... 6 er species (total length 72-80.5 mm)...... 6 6. Pale mesanepisternal stripes incomplete to absent 5. Stem of T-spot strongly narrowed before its con- (fig. 100), as wide as 25-33% of mesanepister- fluence with transverse arms (fig. 5a); ventral tu- num; ventral tubercle of abdominal segment I bercle of abdominal segment I high (about as lower than 30% of its length (fig. 265); cercus high as 50% of its length in lateral view) (fig. with rounded tip bearing a minute medial spine 238); cercus: 3.7 mm (fig. 363c); Peru to (fig. 387c); Paraná and Minas Gerais states in Ecuador (fig. 442)...... intricata Brazil (fig. 453)...... pauloi – Stem of T-spot widely confluent with transverse – Pale mesanepisternal stripes complete (figs. 104- arms (fig. 2a); ventral tubercle of abdominal seg- 106), as wide as 50% of mesanepisternum; ventral ment I low (about as high as 30% of its length in tubercle of abdominal segment I about as high as lateral view) (as in fig. 236); cercus: 2.7-3 mm (fig. 50% of its length (as in figs. 261-262); cercus with 359c); Ecuador to Venezuela (fig. 442) ...... pointed medial tip (fig. 386c-e); Argentina north ...... brevicercia to Venezuela (fig. 453) ...... planaltica 6. Stem of T-spot widely confluent with transverse 7. Mesepimeral stripe narrowed to 50% of its basal arms; vertex black with a small rounded yellow width at midlength (figs. 101-102); cercus with spot on each anterior lateral edge (fig. 6a); cercus mucron on external margin (figs. 385c-d); anteri- widening distally with pointed tip (Fig 364c); or side of ventral tubercle concave (fig. 267); Bolivia north to Peru (fig. 443)...... obscura Venezuelan tepuis (fig. 454)...... nubigena – Stem of T-spot strongly narrowed before its con- – Mesepimeral stripe approximately uniform in fluence with transverse arms; vertex yellow with width or slightly narrowed at midlength (figs. 92- posterior 30% black (fig. 8a); cercus with inner 94); cercus tip rounded or medially pointed (figs. and outer margins approximately parallel, tip not 382c-e); anterior side of ventral tubercle convex pointed (fig. 365c); northwestern Argentina to (figs. 261-262); Bolivia north to Mexico (fig. Peru (fig. 443)...... vigintipunctata 454) ...... cornigera

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Key F-3 391c); larger species (cercus 6.5 mm, total length 1. Black stripe on frontoclypeal groove strongly 68-72.5 mm); Bolivia to Colombia (fig. 458) ..... widened toward the eyes (figs. 37-38b-c); ...... joannisi pterothoracic stripes complete, linear, and narrow- – PD spots present in segments VIII-X; outer margin er than 50% of their sclerites (figs. 114, 118)...... 2 of segment VII linear (fig. 191b); maximum width – Frontoclypeal groove not bordered by black of cercus at distal 30% (fig. 393c); smaller species stripe (figs. 33-36b-c); pterothoracic stripes with (cercus 5 mm, total length 58.3 mm); Táchira deep indentations (figs. 113, 115), divided into State in Venezuela (fig. 458) ...... demarmelsi spots (figs. 110, 111, 116, 117), or wider than 50% of their sclerites (fig. 112)...... 4 Key F-4 2. Labrum, clypeus and frons orange; pale abdominal 1. Medio-longitudinal pale stripe on abdominal seg- PL spots present on segments IV-IX, fused with ML ment II complete (figs. 132, 134, 138-139c-d); on IV-VIII; only known from Itatiaia (type locality), South America ...... 2 Rio de Janeiro state, Brazil (fig. 459) (characters – Medio-longitudinal pale stripe on abdominal seg- abstracted from Calvert, 1953...... decessus ment II incomplete or interrupted at midlength – Labrum, clypeus and frons light blue or yellow; (figs. 127-130c-d); Central and North America 5 pale abdominal PL spots absent on segments VI- 2. Supratriangles always crossed (1-4 crossveins) VII, fused or not with ML on VIII (as in figs. 155- (fig. 400)...... 3 156c-d)...... 3 – Supratriangles generally free, sometimes one or 3. Ventral tubercle of abdominal segment I low and two with 1-2 crossveins (in 2-10% of specimens rounded in lateral view (fig. 275); black of T-spot examined) (fig. 399) ...... 4 not extended on antefrons; vertex yellow with lat- 3. Pale mesanepisternal stripe complete (fig. 81); eral edges black (figs. 38b-c); cercus with round- Chile and Argentina north to Brazil (fig. 450) .... ed tip, longer than abdominal segments IX-X and ...... confusa shorter than VIII-IX (fig. 395c); São Paulo, Rio de – Pale mesanepisternal stripe incomplete or absent Janeiro and Minas Gerais states in Brazil (fig. (fig. 79-80); Bolivia to Venezuela (fig. 451) ...... 459)...... punctata ...... marchali – Ventral tubercle of abdominal segment I high and 4. Clypeal lobes rounded (fig. 16c); stem of T-spot trapezoidal shaped in lateral view (fig. 274); black widening basally (fig. 16a); membranule dark ex- of T-spot extended on dorsal surface of antefrons; cept basal 30% pale; cercus tip not pointed (fig. vertex yellow with lateral and posterior 30% black 373c); ventral tubercle of abdominal segment I (figs. 37b-c); cercus with acute mucronate tip, as with few denticles restricted to apex (fig. 250); long as abdominal segments VIII-X; Minas Gerais central Chile to Ecuador (fig. 448) ...... elsia state in Brazil (fig. 459) ...... eduardoi – Clypeal lobes angled (fig. 14c); stem of T-spot 4. Transverse arms of T-spot represented by a fine parallel sided (fig. 14a); membranule dark except line (fig. 33a); labrum orange, clypeus and frons basal 15% pale; cercus tip pointed (fig. 371c); bright yellow; no dark stripe on frontoclypeal ventral tubercle of abdominal segment I with nu- groove (figs. 33b-c); Peru to Ecuador (fig. 457) .... merous denticles spread over posterior surface (as ...... biliosa fig. 248); Argentina to Bolivia and Brazil (fig. – Transverse arms of T-spot wide (fig. 34-36a); 447) ...... bonariensis labrum, clypeus and frons light blue to green or 5. Posterior margin of inner carina of ventral terga yellow; dark stripe on frontoclypeal groove (figs. V-VII convergent to outer carina (fig. 165b); cer- 34-36b-c)...... 5 cus tip rounded (fig. 366d); Mexico to southern 5. Stem of T-spot wider than vertex (fig. 34a); pale USA (fig. 444) ...... dugesi pterothoracic stripes wide (as wide as 50-80% of – Posterior margin of inner carina of ventral terga their sclerites) and complete (fig. 112); Táchira V-VII parallel to outer carina (figs. 166-168b); cer- state in Venezuela (fig. 457)...... condor cus tip pointed (figs. 367-369d)...... 6 – Stem of T-spot narrower than vertex (figs. 35- 6. No pale abdominal PL spots on segments IV-V 36); pale mesepimeral and metepimeral stripes (fig. 128d); Costa Rica to Mexico (fig. 444)...... narrow (as wide as 30-50% of their sclerites) and ...... jalapensis complete with an anterior and a posterior inden- – Pale abdominal PL spots of segments IV-V present tation (figs. 113, 115), or divided into spots (figs. (figs. 129-130d)...... 7 116-117) ...... 6 7. Ventral tubercle of abdominal segment I low (as 6. PD spots absent in segments VIII-X (figs. 154c-e); wide as about 3 times its height) (fig. 245); base outer margin of segment VII concave (fig. 192b); of stem of T-spot wider than vertex, anterior of maximum width of cercus at medial 30% (fig. vertex half pale, posterior 50% black (fig. 12a);

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outer margin of cercus linear (fig. 369d); eastern angles always crossed (1-2 cross veins) (fig. 400); USA to southern Canada (fig. 445)...... mutata southeastern Brazil (fig. 450) ...... brasiliensis – Ventral tubercle of abdominal segment I high (as – Abdominal segment II with a complete medio- wide as 2 times its height) (fig. 244); base of stem longitudinal black stripe between transverse cari- of T-spot narrower than vertex, vertex pale with na and posterior margin (fig. 131c); pale posterior 30% black (fig. 11A); outer margin of metepimeral stripe complete to absent, but never cercus slightly convex (figs. 368D-E); Mexico to divided into ventral and dorsal spots (figs. 54- southern Canada through western USA (fig. 445) 59); frontal carina in dorsal view smoothly curved ...... multicolor (figs. 13a); supratriangles generally free, some- times one with 1 cross vein (in 10% of specimens Key F-5 examined) (fig. 399); southern Argentina to 1. Pale PL spots absent on segments V-IX (fig. 137d); Ecuador (fig. 446) ...... absoluta Baja California state in Mexico north to Canada through southwestern USA (fig. 449).....californica Rhionaeschna species accounts – Pale PL spots present on segments V-IX (figs. 131, 133, 135-136, 140-142d); South America ...... 2 In order to facilitate species treatment I have allo- 2. Clypeal lobes angled (figs. 13, 15, 18, 22-24c); cated the 39 species into seven groups. The groups are ventral tubercle of abdominal segment I with nu- based on male characters used to build the subordi- merous denticles spread over its posterior surface nate keys, and do not necessarily imply clades. (figs. 247, 249, 252, 256-258)...... 3 – Clypeal lobes rounded (fig. 14c); ventral tubercle 1. R. draco group of abdominal segment I with few denticles re- stricted to its apex (fig. 251); Galápagos Islands Frons projected dorsally, higher than vertex (fig. (fig. 448) ...... galapagoensis 1b); pterothorax and abdominal segments VII-X with- 3. Tip of cercus rounded (figs. 372, 379-381); ven- out pale markings (figs. 39, 119a-b); hamulus with a tral tergum VI widest point at basal 30% (figs. medial digitiform projection on its antero-ventral 171, 178-180b)...... 4 margin (fig. 276); female cercus with long terminal – Tip of cercus pointed (figs. 370, 375); ventral ter- spine (fig. 439). One species: R. draco. gum VI widest point at distal 60% (figs. 169, 174) ...... 7 Rhionaeschna draco (Rácenis) comb. n. 4. Larger species (total length 55.8-67.2 mm, cercus (figs. 1-he, 39-th, 119-ab, 157-te, 195-ge, 234-tg, 276-ha, 316-as, 4.3-5.5 mm); southern Argentina and Chile to 358-ce, 397-wi, 439, 460-Mp) southern Peru (fig. 452) ...... variegata Aeshna draco Rácenis, 1958: 323, 326-330, figs. 1, 3a-c (de- – Smaller species (total length 47-54.2 mm, cercus scription (: holotype ( VENEZUELA: El Dragón-Auyan- 3-4.35 mm) ...... 5 tepui, 1700 m (Bolivar dept.), 15-IX-1958, Trebbau leg. 5. MD spots widely confluent to ML in segments III- (MIZA) [examined: allotype & (GSVC)]); Rácenis 1970: IX (fig. 141c-d); stem of T-spot abruptly narrow- 31; De Marmels 1989b: 35; De Marmels 1990a: 3-5 (de- scription larva); De Marmels 1992b: 41; Vick 1993: 93- ing at distal 30% (fig. 23a); northern Chile (fig. 99 (description &). 451) ...... tinti – MD spots separated from ML in segments III-IX Diagnosis. – Rhionaeschna draco is distinguished (figs. 133, 140c-d); stem of T-spot gradually nar- from all other species of Rhionaeschna by the projected rowing to distal 30% (figs. 22, 15a) ...... 6 edge of frons (fig. 1b), uniformly colored reddish 6. ML and PL spots separated on segments IV-V (fig. brown thorax (fig. 39), abdomen with pale spots only 140d); supratriangles always crossed (1-2 cross in segments II-VI (fig. 119), antero-ventral margin of veins) (fig. 397); Bolivia and Peru (fig. 452) ...... hamuli with a medial digitiform projection (fig. 276), ...... peralta and tip of female cerci bearing a long spine (fig. 358c). – ML and PL spots confluent on segments IV-V (fig. Description. – Anteclypeus pale brown, postclypeus 133d); supratriangles usually free, sometimes one and frons bright light blue, grayish white area sur- or two with 1-2 cross veins (in 10% of specimens rounding T-spot stem; T-stem widening basally with examined as in fig. 396); Chile and southern Ar- convex sides; frontoclypeal groove lacking dark stripe; gentina (fig. 447)...... diffinis vertex light blue, with anterior and posterior margins 7. Abdominal segment II lacking a medio-longitudi- black; edge of frons angled in lateral view and project- nal black stripe (fig. 136c); pale metepimeral ed antero-dorsally (higher than vertex) (figs. 1a-c). stripe divided into a ventral rounded spot and a Thorax reddish brown, without pale stripes or spots dorsal T-shaped spot (fig. 76-78); frontal carina in (fig. 39). Abdomen dark reddish brown, pale abdomi- dorsal view angled anteriorly (fig. 18a); supratri- nal spots bright light blue, restricted to PD in II, AD, AL

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and ML in III, and AL in IV-VI (figs. 119a-c). Ventral tu- Rhionaeschna brevicercia (Muzón & von Ellenrieder) comb. n. bercle of abdominal segment I with anterior margin (figs. 2-he, 40-th, 120-ab, 158-te, 196-ge, 236-tg, 277-ha, 317-as, concave in lateral view (fig. 234); spines of anterior 359-ce, 442-Mp) lamina long (as long as 37% of its basal width) and tri- Aeshna (Marmaraeschna) brevicercia Muzón & von Ellen- angular (fig. 316); anterior margin of hamuli with a rieder, 2001: 98-102, 122 (description (&: holotype ( medial digitiform projection, tip of hamular anterior ECUADOR: Pichincha, between Calacali and Nanegalito, process rounded in posterior and ventral view (fig. ca. 30 km W Quito, 2000 m, 04-XI-1990, O.S. Flint Jr. 276); auricles with two teeth (fig. 234). Lateral carina leg. (USNM) [examined: holotype ( allotype & paratypes of male cercus concave; dorso-distal crest lower than (&(RWGC; USNM; MIZA; MLPA)]). base of cercus in lateral view and smoothly curved [Aeshna (Marmaraeschna) intricata. – Martin 1908: 59-60 (in part); Calvert 1956: 112-119 (in part). Misidentifica- along distal 25% of cercus; sub-basal tooth low and tion] blunt (fig. 358b); maximum width at medial 30%; tip [Aeshna (Marmaraeschna) vigintipunctata. – De Marmels directed externally (fig. 358a). Female cercus as long as 1988: 101; De Marmels 1990b: 337; De Marmels 1994: abdominal segment IX; maximum width at distal 437; De Marmels 2001a: 130-132 (description larva). 25%; tip bearing a spine (fig. 358c). Dimensions: Misidentification] head width: 10.65-11.2; HW length: 49-55.5; HW Diagnosis. – Rhionaeschna brevicercia shares with R. width: 14.1-15.2; HW pterostigma length: 2.54-3; cer- obscura the wide confluence of stem and transverse ci length: ( 6.5, & 2.52; female cerci maximum arms of T-spot but can be easily distinguished from it width: 0.75; total length: 77.5- 82. by the anterior half of vertex yellow (vertex black with Biology. – De Marmels (1990a) described last lar- small yellow antero-lateral spots in R. obscura, figs. 2, val instar by supposition based on larvae and exuviae 6a), male cercus shape (cercus with sub-basal tooth low from Bolivar and Amazonas, Venezuela, and reported and dorso-distal crest lower, sub-basal tooth promi- adults and larvae as common in Cerro Neblina. The nent and dorso-distal carina higher in R. obscura, figs. species breeds in mountain creeks and rock pools in 359, 364b), and female cercus size (not longer than 3 the tepuis. Rare in collections. mm in R. brevicercia, 5.65 mm or longer in R. obscura, Distribution [0-5°N, 60-66°W, 1700-2800 m] figs. 359, 364c). It has been confused with R. vigin- (figs. 439, 460). – Venezuela: Bolivar (GSVC*; MLPA*; tipunctata but is separated by the wide confluence of T- Rácenis 1958, 1970; De Marmels 1990a, 1992b; spot stem and transverse arms (narrow in R. vigin- Vick 1993) and Amazonas (USNM; De Marmels tipunctata, figs. 2, 8a), sub-basal tooth of male cercus 1989b, 1990a). low (prominent in R. vigintipunctata, figs. 359, 365b), lower dorso-distal crest (higher in R. vigintipunctata, figs. 359, 365b), and female cercus mucronate with in- 2. Marmaraeschna group ner and outer margins diverging distally (tip not mu- (= subgenus Marmaraeschna Calvert, 1952). cronate and margins parallel along medial 30% in R. Frontal carina recessed, located in the horizontal vigintipunctata, figs. 359, 365c). Rhionaeschna brevicer- portion of the frons, giving a rounded contour to an- cia is most similar to R. intricata in size and shape of tero-dorsal margin of head (figs. 2-8b); pterothorax cercus but it differs by the wide confluence of stem and with marbled pattern of black markings and pale areas transverse arms of T-spot (narrow in R. intricata, figs. (figs. 40-47); two rows of cells between RP1 and RP2 in 2, 5a), the lower ventral tubercle of abdominal segment HW beginning proximal to pterostigma or under it I (higher in R. intricata, figs. 236, 238), and the short- (fig. 396); hamular anterior process very high (dis- er female cercus (2.7-3 mm in R. brevicercia, 3.7 mm in tance between ventral tip of process and axis of hamu- R. intricata, figs. 359, 363c). lus higher than distance between dorsal end of hamu- Description. – Clypeus and frons light blue and yel- lar fold and axis) (figs. 277-284); anterior lamina spine low to grayish blue, grayish blue spots lateral to light wide throughout its length in lateral view (twice or less yellow area surrounding T-spot stem; T-stem maxi- as wide at base as at distal 30% (figs. 317-324); exter- mum width at mid-length, sides convex, widely con- nal margin of male cercus approximately linear in lat- fluent with transverse arms (fig. 2a); wide dark brown eral view (with the exception of intricata, where it is stripe on frontoclypeal groove (figs. 2b-c); anterior slightly concave) (figs. 359-365). Seven species: R. bre- 50% of vertex yellow, posterior 50% black. Frontal vicercia, R. brevifrons, R. fissifrons, R. intricata, R. ob- carina with a slight concavity in frontal view (fig. c); scura, R. pallipes, R. vigintipunctata. frons not flattened (fig. 2b). Pterothorax pale yellow- brown with 2-3 black spots in mesanepisternum, 3-4 in mesepimeron, 2-3 in metepisternum and 3-4 in metepimeron (fig. 40). Abdomen pale brown and grayish light blue with black spots; black areas of ven- tral abdominal terga II extended over auricle; auricles

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lectotype ( (here designated) PERU, Arequipa, 2300 m, with two teeth (fig. 196); constriction of segment III 14-IV-1907, K. Seyd. leg. (MWNH) [examined]); Martin marked (figs. 120a-d). Inner and outer carinae of ven- 1911: 13; Ris 1913: 83; Hincks 1934: 80: Schmidt 1952: tral terga VI sinuous (figs. 158a-b). Ventral tubercle of 256. Syn. n. abdominal segment I lower than 30% of its length in Aeshna (Rhionaeschna) maita. – Calvert 1956: 11, 99-100. lateral view, with anterior margin convex in lateral [Aeshna (Marmaraeschna) intricata. – Calvert 1956: 112- view; genital lobe as high as twice the tubercle height 119 (in part); Fraser 1957: 155, 159, 163, fig. 4g; Jurzitza (fig. 236). Sub-basal tooth of male cercus low; dorso- 1989: 9 (in part). Misidentification] distal crest lower than 50% of cercus width at base, de- Diagnosis. – Rhionaeschna brevifrons shares with R. veloped at distal 20% of cercus (fig. 359b); tip on ex- pallipes and R. fissifrons a flattened frons (figs. 3-4, 7b), ternal margin (fig. 359a). Female cercus maximum stem of T-spot widest at posterior end and narrowly width at distal 30%; tip mucronate (fig. 359c). Di- confluent or separated from transverse arms (figs. 3-4, mensions: head width: 8.9-10.2; HW length: 48.2-52; 7a), stocky abdomen (constriction of segment III slight, HW width: 13.6-16.3; HW pterostigma length: 2.6-3.2; figs. 121-122, 124), and short cercus (less than 4.8 mm cerci length: ( 5.4-6, & 2.7-3; female cerci maximum in males, 2.3 mm in females, figs. 360-362). The three width: 0.75; total length: 66.6-73.5. species can be distinguished from each other by the Biology. – De Marmels (2001) described the larva shape of the frontal carina (linear in R. pallipes, fig. 7c, based on reared specimens found along small to concave in R. brevifrons, fig. 3c, deeply cleft in R. fis- medium-sized streams with dense riparian vegetation sifrons, fig. 4c); and shape of hamular anterior process and submerged roots bordered by bushes and scat- tip (figs. 280-282b-c). In addition, the sub-basal tooth tered trees outside closed forest. He reported adults of male cercus in R. brevifrons is low (figs. 360b, e-f), patrolling over open pasture lands along or away from not prominent as in R. fissifrons and R. pallipes (figs. small streams and occasionally perching on vertical 361-362b); the pterothorax has fewer black spots (figs. rock surfaces where they were almost invisible due to 41-44); and in R. fissifrons the inner and outer carinae their lichen-like color. From Ecuador it has been re- of abdominal ventral terga VI are linear and parallel to ported from clear, sandy and rocky bottom streams each other (fig. 160) (sinuous in R. brevifrons and R. with good current, surrounded by marsh plants and pallipes, figs. 159, 163), and in R. pallipes the black ar- shrubs, and one larger pool (Dr. D. R. Paulson pers. eas of ventral abdominal terga II do not extend over comm.). A common species. auricles (figs. 197-198, 201). Remarks. – Martin (1908) described R. intricata Description. – Clypeus and frons light blue and yel- based on several specimens from Ecuador, Colombia low to grayish blue, grayish dark blue spots lateral to and Venezuela. Examination of these specimens re- light blue area surrounding T-spot stem; T-stem max- vealed that the type series of R. intricata included sev- imum width at posterior end, sides converging anteri- eral specimens of R. brevicercia, from Venezuela orly, narrowly or not confluent with transverse arms (without localities), Colombia and Ecuador. (figs. 3a, c); narrow black stripe on frontoclypeal Distribution [3°S-9°N, 70-79°W, 604-3020 m] groove (figs. 3b-c, d-e); vertex yellow with posterior (fig. 442). – Ecuador: Tungurahua (RWGC*), Esmeral- margin black. Frontal carina with a slight concavity in das (ISNB; MLPA), Imbabura (ANSP; ISNB; QCAZ), Napo frontal view (figs. 3c, e); frons flattened (figs. 3b, d). (KJTC*), Manabi (UMMZ), and Pichincha (DRPC; FSCA; Pterothorax pale yellow-brown with 1 black spot in MLPA*; ZMHB; QCAZ; USNM*) – Colombia: Santander mesanepisternum, 2 in mesepimeron, 1-2 in metepis- (ISNB); Cundinamarca (UMMZ) – Venezuela: Trujillo ternum and 1 in metepimeron (figs. 41-42). Ab- (MIZA), Mérida (MIZA; MLPA), Barinas (MIZA), and domen pale brown to pale yellow and grayish light Táchira (MIZA; De Marmels 2001a). blue with dark reddish brown to black spots; black ar- eas of ventral abdominal terga II extended over auricle (fig. 197); constriction of segment III very slight (figs. Rhionaeschna brevifrons (Hagen) comb. n. 121a-d). Inner and outer carinae of ventral terga VI (figs. 3-he, 41-42-th, 121-ab, 159-te, 197-ge, 235,237-tg, 279-280- sinuous (figs. 159a-c). Ventral tubercle of abdominal ha, 318-319-as, 360-ce, 396-wi, 440-Mp) segment I lower than 30% of its length, with anterior Aeshna brevifrons Hagen, 1861: 129, 314 (description (&: margin convex in lateral view; genital lobe approxi- lectotype ( MEXICO: Acapulco (MCZ) [examined: lecto- mately as high as tubercle (figs. 235, 237); auricles type ( paralectotype & (MCZ)]; Hagen 1875: 36; Ris 1904: 4, 29; Calvert 1905:181, 186; Calvert 1908: 488; with two teeth (fig. 197). Sub-basal tooth of male cer- Martin 1908:58, 59; Martin 1911: 12; Calvert 1952: 256 cus low; dorso-distal crest lower than 50% of cercus (Marmaraeschna n. subgn.); Schmidt 1952: 238 (in part); width at base, developed at distal 25% of cercus (fig. Calvert 1956: 13, 110-112 (in part); Paulson 1977: 175 360e); tip on external margin (fig. 360a, d). Female (in part); Muzón & von Ellenrieder 2001: 144-148 (re- cercus maximum width at distal 80%; tip mucronate description (&). or not (figs. 360c, g-h). Dimensions: head width: 8.6- Rhionaeschna maita Förster, 1909: 220-223 (description (: 9.5; HW length: 41.7-46; HW width: 13.5-15; HW

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pterostigma length: 2.6-3.1; cerci length: ( 4.2-4.8, holotype, & allotype, (¶types (FSCA; DRPC; MLPA; UMMZ; RWGC)]). & 1.9-2.3; female cerci maximum width: 0.5; total [Aeshna (Marmaraeschna) brevifrons. – Schmidt 1952: 238 length: 57-69.5. (in part). Misidentification] Biology. – Collection labels indicate that in North- [Aeshna (Marmaraeschna) intricata. – Calvert 1956: 13, ern Chile specimens were collected flying over ther- 112-119 (in part); Rácenis 1959: 494 (in part); Paulson mal creeks in areas of naturally warm waters used for 1977: 175 (in part); Rodrigues Capítulo et al. 1991: 62 valley irrigation in the middle of the desert, and in a (in part); Rodrigues Capítulo 1992: 38, 57 (in part). Misidentification] valley surrounded by high mountains with small irri- gation streams for alfalfa crops. A common species. Diagnosis. – The deeply cleft frontal carina (fig. Larva unknown. 4c) and linear contour of ventral terga (fig. 160) are Remarks. – The type locality of Rhionaeschna brev- unique character states of Rhionaeschna fissifrons. Fur- ifrons (Mexico, Acapulco) is probably a labeling mis- ther distinction from the other species is given under take, since no Marmaraeschna species has ever been R. brevifrons. found north of Venezuela, and no specimen of R. bre- Description. – Clypeus and frons pale brown, gray- vifrons north of Peru. Förster (1909) described Rhion- ish dark blue spots lateral to yellow area surrounding aeschna maita based on two males from Arequipa, T-spot stem; T-stem maximum width at posterior Peru, one of them deposited in the Museum Wies- end, sides converging anteriorly, separated from trans- baden and the second one in his collection. The sec- verse arms (fig. 4a); wide black stripe on frontoclypeal ond male is apparently lost (Calvert 1956: 99), and groove (fig. 4b-c); vertex yellow with lateral and poste- the first one is here designated as lectotype. Examina- rior margins black. Frontal carina with a deep medial tion of the lectotype showed that Rhionaeschna maita cleft in frontal view (fig. 4c); frons flattened (fig. 4b). Förster 1909 is a junior synonym of Rhionaeschna bre- Pterothorax pale yellow-brown with 2-3 black spots in vifrons (Hagen 1861). The specimen is poorly pre- mesanepisternum, 3-4 in mesepimeron, 3-4 in served; the head is flattened, thorax partly broken, and metepisternum and 3-4 in metepimeron (fig. 43). Ab- cerci seem to be atrophied (left) and malformed domen grayish light blue and pale yellow with dark (right), but all diagnostic characters of R. brevifrons reddish brown to black areas; black areas of ventral ab- can be distinguished and wing dimension and vena- dominal terga II extended over auricle; auricles with tion characters agree completely with those for R. bre- two teeth (fig. 198); constriction of segment III very vifrons. Notes on the types of R. maita made by E. slight (figs. 122a-d). Inner and outer carinae of ventral Schmidt in Calvert (1956) include some errors (i.e. terga VI linear and parallel to each other (figs. 160a-b). stated absence of frontal carina and anterior lamina Ventral tubercle of abdominal segment I lower than spines), which are probably responsible for the mis- 30% of its length in lateral view, with anterior margin placement of this species until now. convex in lateral view; genital lobe approximately as Distribution [9-44°S, 68-77°W, 510-3750 m] (fig. high as tubercle (fig. 239). Sub-basal tooth of male 440). – Chile: De Los Lagos: Valdivia (IEUM), Gener- cercus prominent; dorso-distal crest lower than 50% al O´Higgins: Cachapoal (USNM), Metropolitan: of cercus width at base, developed at distal 20% of cer- Chacabuco (MNHN), Cordillera (IEUM), and Santiago cus (fig. 361b); tip on external margin (fig. 361a). Fe- (IMLA; IEUM; RWGC*; USNM), Valparaiso: Valparaiso male cercus maximum width at distal 66%; tip mu- (IEUM; MCZ*; MNHN), Atacama: Huasco (FSCA*), Co- cronate (figs. 361c). Dimensions: head width: 8.2-9.6; quimbo: Elqui (IEUM) and Choapa (DRPC; FSCA; HW length: 41.3-48.8; HW width: 13.8-16.1; HW IEUM; MLPA; RWGC; UMMZ), Antofagasta: El Loa pterostigma length: 2.3-2.9; cerci length: ( 3.5-4.5, (MLPA; MNHN; UMMZ), and Tarapacá: Parinacota & 1.7-2.2; female cerci maximum width: 0.45; total (FSCA; MLPA; UMMZ), Arica (FSCA; MLPA; UMMZ) and length: 56-59. Iquique (FSCA; MLPA; MNHN; UMMZ) – Peru: Arequipa Biology: In Peru adults of Rhionaeschna fissifrons (ANSP; MCZ; MWNH*; USNM; Förster 1909), Ayacucho were seen flying over the steep part of a rocky stream (FSCA), Apurimac (MLPA; RWGC*; WB), Lima (DRPC; and grassy areas nearby, and landing on vertical or IMLA*; RWGC*), and Ancash (DRPC). horizontal rocks at the stream (DRPC), and in North- western Argentina patrolling mountain streams and Rhionaeschna fissifrons (Muzón & von Ellenrieder) creeks in Preandean and Andean localities. Rare in comb. n. collections. Larva unknown. (figs. 4-he, 43-th, 122-ab, 160-te, 198-ge, 239-tg, 281-ha, 320-as, Distribution [11-27°S, 65-75°W, 1200-4057 m] 361-ce, 440-Mp) (fig. 440). – Argentina: Catamarca (FSCA) and Salta Aeshna (Marmaraeschna) fissifrons Muzón & von Ellen- (MACN; MLPA*; ZMHB) – Chile: Antofagasta: El Loa rieder, 2001: 148-150 (description (&: holotype ( AR- (UMMZ; MLPA*) – Bolivia: Cochabamba: Chapare GENTINA: 7 km S Minas Capillitas (Catamarca prov.), ca. (FSCA) – Peru: Puno (UMMZ), Junín (IMLA; RWGC; 3000, 27-XII-1072, J. Schultz leg. (FSCA) [examined: ( USNM), Ayacucho (FSCA), and Cuzco (DRPC).

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Rhionaeschna intricata (Martin) comb. n. Martin collections. In the Martin collection at the (figs. 5-he, 44-th, 123-ab, 161-te, 199-ge, 238-tg, 278-ha, 321-as, 363-ce, 442-Mp) MNHN there are no type specimens of R. intricata (Dr. J. Legrand pers. com.), and nine specimens with type Aeshna (Marmaraeschna) intricata Martin, 1908: 59-60, fig. labels are deposited in the Selys collection at the ISNB. 55 (in part; description (&: lectotype & (here designat- ed) ECUADOR: Nanegal, 0°7’0N 78°40’0W 2093 m Examination of this series revealed that it includes two (Pichincha prov.) (ISNB) [examined]); Martin 1911: 12; species, which according to the last synopsis of the Calvert 1952: 256; Calvert 1956: 112-119 (in part); group would correspond to R. intricata and R. bre- Rácenis 1959: 494 (in part); Muzón & von Ellenrieder vicercia (Muzón & von Ellenrieder 2001). In that pa- 1998: 23; Muzón & von Ellenrieder 2001: 150-154 (re- per, identification of R. intricata was based on the il- description (&). lustration of the male cerci from the original description (Martin 1908). However, no male speci- Diagnosis. – Rhionaeschna intricata is characterized men belonging to this species was found among the by the high ventral tubercle of abdominal segment I (as types at the ISNB and all are R. brevicercia except for high or higher than 50% of its length, fig. 238), which one female, which is herein designated as lectotype of is low in the remaining Marmaraeschna species (figs. R. intricata. In addition to the mixed series of nine 235-237, 239-242). Further differences from other specimens at the ISNB collection there is also a pinned species are found under R. brevicercia and R. brevifrons. type label [no sex stated] without an associated speci- Description. – Clypeus and frons gray with yellow men. This label may correspond to the lost male illus- margins, light yellow area surrounding T-spot stem; T- trated by Martin, or it may belong with one of the stem maximum width at mid-length, sides convex, other missing specimens mentioned by Martin (1908) strongly narrowed at confluence with transverse arms of his collection from Mexico, Peru, Bolivia, Chile (fig. 5); wide black stripe on frontoclypeal groove (fig. and Brazil. 5b-c); anterior 50% of vertex yellow, posterior 50% Distribution [0-6°S, 78°W, 2248-2833 m] (fig. black. Frontal carina with a slight concavity in frontal 442). – Peru: Cajamarca (FSCA*; UMMZ) – Ecuador: view (fig. 5c); frons not flattened (fig. 5b). Pterothorax Chimborazo (ANSP; MLPA*) and Pichincha (ANSP*; pale yellow-brown with 2 black spots in mesanepister- ISNB). num, 3-4 in mesepimeron, 2-3 in metepisternum and 2-3 in metepimeron (fig. 44). Abdomen pale reddish Rhionaeschna obscura (Muzón & von Ellenrieder) brown to yellow and light blue with black spots; black comb. n. areas of ventral abdominal terga II extended over auri- (figs. 6-he, 45-th, 125-ab, 162-te, 200-ge, 240-tg, 283-ha, 322-as, cle; auricles with three teeth (fig. 199); constriction of 364-ce, 443-Mp) segment III marked (figs. 123a-d). Inner and outer cari- Aeshna (Marmaraeschna) obscura Muzón & von Ellenrieder, nae of ventral terga VI sinuous (figs. 161a-b). Ventral 2001: 154-156 (description (&: holotype ( BOLIVIA: 4 tubercle of abdominal segment I as high or higher than km W Chulimani (La Paz dept., sud Yungas prov.), 25-V- 50% of its length in lateral view, with anterior margin 1989, T.C. Emmel leg. (FSCA) [examined: holotype (, al- lotype &, paratypes (&(FSCA; ANSP; MLPA; UMMZ)]). convex in lateral view; genital lobe approximately as [Aeshna (Marmaraeschna) vigintipunctata. – Calvert 1956: high as tubercle (fig. 238). Sub-basal tooth of male cer- 119-123 (in part). Misidentification] cus very low and blunt; dorso-distal crest lower than 50% of cercus width at base, developed at distal 25% Diagnosis. – The dark vertex of R. obscura (fig. 6a) of cercus (figs. 363b, d); tip medial (fig. 363a). Female is unique to this species within the marmaraeschna- cercus maximum width at distal 66%; tip mucronate group (figs. 2-5, 7-8a). Further differences from other (fig. 363c). Dimensions: head width: 9-9.5; HW length: species are found under R. intricata and R. brevifrons. 46.4-50.25; HW width: 14.1-16.4; HW pterostigma This species is the largest marmaraeschna together length: 2.8-3.35; cerci length: ( 4.9-5.5, & 3.7; fe- with R. vigintipunctata, both of which have a similarly male cerci maximum width: 0.8; total length: 66-68. shaped male cercus (figs. 364-365b-c). However, the Biology. – The following notes by Felix Woyt- widely confluent T-spot stem (fig. 6a) (narrow in R. kowski on envelopes of specimens he collected in vigintipunctata, fig. 8a), shape of genital lobe (sub- Peru state: ‘This species has been taken sitting on the quadrate in R. obscura, fig. 240; subtriangular in R. wall on a rock some 300 m higher in the mountains vigintipunctata, fig. 242), and shape of female cercus east of Celendin far from any water. This is a new (mucronate tip and margins diverging distally in R. species for the higher Andes, rare and difficult to get’. obscura, fig. 364; no mucronate tip and margins paral- Rare in collections. Larva unknown. lel to each other in R. vigintipuncata, fig. 365), allow Remarks. – In his description of R. intricata Martin for the distinction between these two species. (1908) indicated that the specimens he examined Description. – Clypeus and frons light blue, light from Mexico, Peru, Bolivia, Chile, Ecuador, yellow area surrounding T-spot stem; T-stem maxi- Venezuela and Brazil were deposited in the Selys and mum width at mid-length, sides convex, widely con-

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fluent with transverse arms (fig. 6a); wide black stripe 123-125; Paulson 1977: 175; Rodrigues Capítulo et al. 1991: 62; Rodrigues Capítulo 1992: 38, 57; Muzón & von on frontoclypeal groove (fig. 6b-c); vertex black with a Ellenrieder 1998: 23; Muzón & von Ellenrieder 2001: small yellow antero-lateral spot on each side. Frontal 157-160 (redescription &, description (); von Ellenrieder carina with a slight concavity in frontal view (6c); & Muzón 2003: 95-98 (description larva). frons not flattened (fig. 6b). Pterothorax pale yellow- [Aeshna (Marmaraeschna) vigintipunctata. – Calvert 1956: brown with 1 black spot in mesanepisternum, 3 in 119-123 (in part); Rodrigues Capítulo & Muzón 1989b: mesepimeron, 2-3 in metepisternum and 2-3 in 147-148; Rodrigues Capítulo et al. 1991: 62; Rodrigues Capítulo 1992: 38, 57 (in part). Misidentification] metepimeron (fig. 45). Abdomen pale brown to red- [Aeshna (Marmaraeschna) intricata. – Paulson 1977: 175; dish brown and light blue to yellowish green with Rodrigues Capítulo et al. 1991: 62; Rodrigues Capítulo black spots; black areas of ventral abdominal terga II 1992: 38, 57. Misidentification] not extended over auricle; auricles with three teeth (fig. 200); constriction of segment III marked (figs. Diagnosis. – The linear frontal carina of R. pallipes 125a-b). Inner and outer carinae of ventral terga VI (fig. 7c) is unique within the marmaraeschna-group sinuous (figs. 162a-b). Ventral tubercle of abdominal (figs. 2-6, 8c). Further distinction from remaining segment I lower than 30% of its length in lateral view, species is provided under R. brevifrons. with anterior margin convex in lateral view; genital Description. – Clypeus and frons light blue, light lobe as high as twice the tubercle height and sub- blue spots lateral to light yellow area surrounding T- quadrate (fig. 240). Sub-basal tooth of male cercus spot stem; T-stem maximum width at posterior end, prominent and angled; dorso-distal crest lower than sides converging anteriorly, separated from transverse cercus width at base, developed at distal 25% of cercus arms (fig. 7a); narrow black stripe on frontoclypeal (fig. 364b); tip medial (fig. 364a). Female cercus max- groove (fig. 7b-c); vertex light blue with posterior imum width at distal 30%; tip mucronate (fig. 364c). margin black. Frontal carina linear in frontal view (fig. Dimensions: head width: 10-10.7; HW length: 47.8- 7c); frons flattened (fig. 7b). Pterothorax pale yellow- 53.9; HW width: 14.8-17.3; HW pterostigma length: 3- brown with 1-2 black spots in mesanepisternum, 2-3 3.8; cerci length: ( 5.8-6.5, & 5.6-6; female cerci in mesepimeron, 2 in metepisternum and 2-3 in maximum width: 1.4-1.5; total length: 72-80.5. metepimeron (fig. 46). Abdomen grayish light blue Biology. – Dr. K. Tennessen, who collected this and light yellow with reddish brown to black spots; species in Bolivia, kindly provided following com- black areas of ventral abdominal terga II not extended ments: ‘Exuviae were found on a large boulder of a over auricle; auricles with two teeth (fig. 201); con- small stream, and larvae in slow-flowing places. This striction of segment III very slight (figs. 124a-d). Inner stream was about 1-2 m wide, shaded by vegetation and outer carinae of ventral terga VI linear and parallel and the substrate was mostly rocks with some gravel to each other (figs. 163a-b). Ventral tubercle of ab- and mud. The stream was receiving siltation because dominal segment I lower than 30% of its length in lat- the water became very silty when disturbed by our eral view, with anterior margin convex in lateral view; feet; the hillsides were being clear cut and burned for genital lobe approximately as high as tubercle (fig. agriculture. We saw cattle and hogs in the area. Un- 241). Sub-basal tooth of male cercus prominent; dor- doubtedly the species can withstand the siltation, but so-distal crest lower than 50% of cercus width at base, may not be able to tolerate removal of the streamside developed at distal 20% of cercus (fig. 362b); tip on vegetation. The adult males fly along trails where it is external margin (fig. 362a). Female cercus maximum still wooded, and we saw a couple fly rapidly over the width at distal 66%; tip mucronate (fig. 362c). Di- small stream’. Rare in collections. Larva undescribed. mensions: head width: 8.7-10.4; HW length: 43.3- Distribution [10-18°S, 63-76°W, 400-2700 m] 50.05; HW width: 13.3-16.1; HW pterostigma length: (fig. 443). – Chile: without locality (MNHN) – Peru: 2.5-3.4; cerci length: ( 4.1-4.65, & 1.8-2.2; female Huanuco (UMMZ) – Bolivia: La Paz: Sud Yungas cerci maximum width: 0.5; total length: 59-59.5. (FSCA*) and Nor Yungas (ANSP; MLPA). Cochabamba: Biology. – The species has been found in sympatry Chapare (FSCA*; KJTC*), Santa Cruz: Ichilo (FSCA) with R. fissifrons in Preandean mountain range locali- and Florida (KJTC). ties in Northwestern Argentina. Males were observed patrolling streams and creeks and females ovipositing Rhionaeschna pallipes (Fraser) comb. n. in marginal vegetation. They were also occasionally (figs. 7-he, 46-th, 124-ab, 163-te, 201-ge, 241-tg, 282-ha, 323-as, seen perching on rocks and vertical banks of the 362-ce, 441-Mp) streams exposed to the sun where they were camou- Aeschna laticeps Hagen, 1875: 39 (nomen nudum). flaged due to their marbled color. Larvae found under Aeshna pallipes Fraser, 1947: 433, 443-445 (description &: stones and clinging to submerged roots of marginal holotype & ARGENTINA: Amaicha del Valle, 26°36’0S vegetation in these streams were described by von El- 65°55’0W 1800 m (Tafí dept., Tucumán prov.), 01-II- lenrieder & Muzón (2003). Abundant in the field, 1946, A. Willink leg. (IMLA) [examined]); Calvert 1956:14, but rare in collections.

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Distribution [25-38°S, 60-68°W, 12-2500 m] (fig. range forest in Argentina and Bolivia to dry Prean- 441). – Argentina: Buenos Aires (FSCA; MLPA*; USNM), dean environments. Adults were seen flying along the Santa Fe (MLPA; ZMHB), Mendoza (IADIZA), Córdoba streams close to the water. Larva unknown. A com- (IMLA; FSCA; MACN; MCZ), La Rioja (MACN; MLPA), mon species. Catamarca (DRPC), Tucumán (IMLA*), and Salta Distribution [5-29°S, 63-79°W, 170-3139 m] (fig. (MLPA*; ZMHB; RWGC; UMMZ). 443). – Argentina: Catamarca (DRPC; IMLA; MACN; Ris 1918), La Rioja (MACN), Tucumán (IMLA; MLPA*; Rhionaeschna vigintipunctata (Ris) comb. n. ZMHB; USNM), Salta (IMLA; MLPA*; RWGC*), and Jujuy (figs. 8-he, 47-th, 126-ab, 164-te, 202-ge, 242-tg, 284-ha, 324-as, (MLPA; UMMZ) – Bolivia: Cochabamba: Chapare 365-ce, 443-Mp) (FSCA) and Santa Cruz: Ichilo (FSCA*) – Peru: Piura Aeshna vigintipunctata Ris, 1918: 163-166 (description (&: (Ris 1918) and Amazonas (Calvert 1956). lectotype ( ARGENTINA: Cerro de Aconquija, 2400 m, La Paz (Catamarca prov.), 19-II-1915, P. Joergensen leg. (SMFD16055); Fraser 1947: 433, 446; Calvert 1952: 256; 3. Schizuraeschna group Calvert 1956: 119-123 (in part); Paulson 1977: 175; Ro- (= subgenus Schizuraeschna Calvert, 1952). drigues Capítulo et al. 1991: 62 (in part); Rodrigues Capí- tulo 1992: 38, 57 (in part); Donnelly et al. 1998: 115; Cercus inner margin with a ventral process at distal Muzón & von Ellenrieder 1998: 23; Muzón & von Ellen- 25%, dorso-distal crest triangular in lateral view, later- rieder 2001: 160-164 (redescription (&). al carina concave, and sub-basal tooth low and blunt [Aeshna (Marmaraeschna) intricata. – Calvert 1956: 112-119 (figs. 366-369a-c); two rows of cells between RP1 and (in part); Rácenis 1959: 494 (in part). Misidentification] RP2 in HW beginning distal to end of pterostigma (fig. 398). Four species: R. dugesi, R. jalapensis, R. multicol- Diagnosis. – Distinction between R. vigintipuncta- or, R. mutata. ta and its congeners is found under R. brevicercia, R. brevifrons, R. intricata and R. obscura. Rhionaeschna dugesi (Calvert) comb. n. Description. – Clypeus and frons light blue and yel- (figs. 9-he, 48-th, 127-ab, 165-te, 203-ge, 243-tg, 285-ha, 325-as, low, gray blue spots lateral to light yellow area sur- 366-ce, 398-wi, 444-Mp) rounding T-spot stem; T-stem maximum width at Aeshna dugesi Calvert, 1905: 180, 184-185 (description (: mid-length, sides convex, strongly narrowed at conflu- holotype ( MEXICO: Guanajuato, 19º48’0N 104º ence with transverse arms (fig. 8a); wide black stripe 20’60W 966 m, A. Dugès leg. (USNM; 66420) [exam- on frontoclypeal groove (fig. 8b-c); vertex yellow with ined]); Martin 1908: 49-50, 83; Calvert 1956: 107-108; latero-posterior margins black. Frontal carina with a González Soriano & Novelo Gutiérrez 1991: 73, 81, 93- 94; Novelo Gutiérrez & González Soriano 1991: 114, slight concavity in frontal view (fig. 8c); frons not flat- 141-145; González Soriano 1993: 296; González Soriano tened (fig. 8b). Pterothorax pale yellow-brown with 1 & Novelo Gutiérrez 1996: 164; Dunkle 2000: 51; Need- black spot in mesanepisternum, 3-4 in mesepimeron, ham et al. 2000: 122, 125, 133, 138-139. 2-3 in metepisternum and 4-5 in metepimeron (fig. 47). Abdomen pale brown to light yellowish green and Description. – Clypeus and frons light blue to yel- grayish light blue with dark reddish brown to black low, grayish blue spots lateral to white area surround- spots; black areas of ventral abdominal terga II not ex- ing T-spot stem; T-stem narrower than vertex, gradu- tended over auricle; auricles with three teeth (fig. ally narrowing anteriorly; no dark stripe on 202); constriction of segment III marked (figs. 126a- frontoclypeal groove; latero-posterior margin of vertex b). Inner and outer carinae of ventral terga VI sinuous black (figs. 9a-c). Pale stripes light blue to white, com- (figs. 164a-b). Ventral tubercle of abdominal segment plete, and as wide as 30% of their sclerites (fig. 48). I lower than 30% of its length in lateral view, with an- Abdomen reddish brown with light blue and yellow to terior margin convex in lateral view; genital lobe as greenish yellow spots; PL spots in III-V (figs. 127b, d); high as twice the tubercle height and subtriangular posterior portion of inner carina of ventral terga V-VII (fig. 242). Sub-basal tooth of male cercus prominent concave in females (fig. 165b). Ventral tubercle of ab- and angled; dorso-distal crest lower than cercus width dominal segment I as high as 40% of its length (fig. at base, developed at distal 30% of cercus (fig. 365b); 243); distal portion of ventral margin of anterior lam- tip medial (fig. 365a). Female cercus maximum width ina spine linear (fig. 325); tip of hamular anterior at medial 30%; tip not mucronate (fig. 365c). Dimen- process pointed in ventral view (fig. 285b); auricles sions: head width: 9.4-10.3; HW length: 47.2-54.4; with two-three teeth (fig. 203). Male cercus with a HW width: 14.8-18.8; HW pterostigma length: 2.5-3.7; small ventral process at distal 25% not projected pos- cerci length: ( 5.8-6.4, & 4.9-5.65; female cerci max- teriorly (fig. 366b), distance between base of crest and imum width: 1.0-1.1; total length: 72-77. posterior base of ventral process shorter than distance Biology. – Rhionaeschna vigintipuncata was found between base of process and tip of cercus, tip not bent at mountain streams ranging from humid mountain ventrally, dorso-distal crest higher than base of cercus

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in lateral view (fig. 366c). Female cercus external mar- groove; vertex black with central portion light blue gin slightly convex, tip rounded (fig. 366d). Dimen- (figs. 10a-c). Pale stripes light blue to white or yellow; sions: head width: 8.7-9; HW length: 44-53; HW mesanepisternal complete to incomplete at basal 30- width: 13.5-14.3; HW pterostigma length: 3.3-3.5; 50% of mesanepisternum; mesepimeral and cerci length: ( 5.6-6.1, & 5.8-6.2; female cerci maxi- metepimeral complete and as wide as 25% of their mum width: 1.2; total length: 65-74. sclerites (figs. 49-50). Abdomen reddish brown with Diagnosis. – Rhionaeschna dugesi is easily distin- light blue and yellow to greenish yellow spots; PL spots guished from other Schizuraeschna species by the male in III (figs. 128b, d); posterior portion of inner carina cercus with ventral process of inner margin not pro- of ventral terga V-VII linear in female (fig. 166b). Ven- jected posteriorly and tip not projected ventrally (fig. tral tubercle of abdominal segment I as high as 40% of 366b), distal portion of ventral margin of anterior its length (fig. 246); distal portion of ventral margin of lamina spine linear (fig. 325), and posterior portion of anterior lamina spine convex (fig. 326); tip of hamular inner carina of female ventral terga V-VII concave (fig. anterior process pointed in ventral view (fig. 286b); 165). auricles with two-three teeth (fig. 204). Male cercus Biology. – Dunkle (2000) reported males pa- with ventral process at distal 25% projected posterior- trolling low along edges of stream pools edged with ly (fig. 367b), distance between base of crest and pos- grass and trees in pine and oak tree zones. Gonzáles & terior base of ventral process shorter than distance be- Novelo (1991) observed females ovipositing in emer- tween base of process and tip of cercus, only extreme gent or submerged vegetation of streams in some cas- tip ventrally bent, dorso-distal crest higher than base es still in tandem. They described the last larval instar of cercus in lateral view (fig. 367c). Female cercus ex- (Novelo & Gonzáles 1991) based on reared larvae ternal margin slightly convex, tip pointed (fig. 367d). from Durango, MX. A common species. Dimensions: head width: 8.5-9.5; HW length: 42- Distribution [16-36°N, 92-111°W, 240-2400 m] 45.3; HW width: 12.5-13.8; HW pterostigma length: (fig. 444). – Mexico: Oaxaca (DRPC), Puebla (USNM), 3.4-3.5; cerci length: ( 5.7-6.2, & 5.3-5.9; female Guanajuato (USNM; Calvert 1905), Zacatecas (USNM), cerci maximum width: 1.3; total length: 67-72. Baja California Sur (Calvert 1956), Durango (DRPC), Biology. – Larva described by supposition by (Novelo & Gonzáles 1991), and Nuevo León (USNM) Calvert (1956) based on exuviae from Guatemala, – USA: Arizona (LACM; RWGC*; UAIC), Texas (RWGC), Costa Rica, and larvae from Cuba and Venezuela. A and New Mexico (Needham et al. 2000). common species. Adults were collected at small, marshy ponds and streams through open and woods; Rhionaeschna jalapensis (Williamson) comb. n. muddy bottom streams with abundant vegetation; (figs. 10-he, 49-50-th, 128-ab, 166-te, 204-ge, 246-tg, 286-ha, springs; weedy drainage ditch widening into shallow 326-as, 367-ce, 444-Mp) marsh; rain ponds choked with grass in shallow areas, Aeshna jalapensis Williamson, 1908: 307-308 (description open in center; over pasture; some more-or-less boggy (: holotype ( MEXICO: Jalapa, 19°31’60N 96°55’0W areas in hollow tiny forest streams to large, open, 1381 m, IX-1906, P.P. Calvert leg. (ANSP)); Walker 1912: rocky rivers. Females were observed ovipositing in 198; Calvert 1956: 13, 106-107; Machado 1985b: 328; Scirpus and in sedge at and below water level, but González Soriano 1993: 296; González Soriano & Nov- keeping wings dry; males flying very low, bouncing elo Gutiérrez 1996: 164. just over vegetation; and pairs in copula flying along [Aeshna multicolor. – Calvert 1905: 180, 183-184 (in part); Calvert 1907: 400 (in part). Misidentification] shore several times and hanging low in sedges or 3.5-4 [Aeshna punctata. – Martin 1908: 54 (in part: & from Mex- m up in trees (DRPC). ico); Calvert 1956: 82 (in part: records from Mexico); Remarks. – Calvert (1956) included not full-grown Santos 1966a (in part: records from Mexico). Misidenti- larvae from Cuba and Venezuela in his description of fication] the larva. Those records were not included here be- cause no adults of this species are known from these Diagnosis. – Rhionaeschna jalapensis can be distin- countries. Since no reliable diagnostic descriptions guished from the remaining Schizuraeschna species by exist for half-grown larvae it is not possible to identi- the male cercus bent ventrally only at extreme tip (fig. fy the larvae from Venezuela with certainty. 367c), and the presence of PL spots only in abdominal Distribution [8-23°N, 82-105°W, 28-2040 m] (fig. segment III (128b, d). It differs also from the sym- 444). – Panama: Bocas del Toro (Walker 1912) – Cos- patric R. dugesi and R. multicolor by the T-spot stem ta Rica: Puntarenas (DRPC*), San José (DRPC; USNM), wider than vertex (fig. 10a), vs. narrower (figs. 9, 11a). Cartago (DRPC; USNM; Calvert 1956), Heredia (DRPC; Description. – Clypeus and frons light blue to yel- INBC; USNM), Alajuela (USNM; DRPC), and Guanacaste low, grayish blue spots lateral to white area surround- (LACM; RWGC*) – Nicaragua: Matagalpa (Beckemeyer ing T-spot stem; T-stem wider than vertex, gradually 2001) – Honduras: without locality (Förster 2001) – narrowing anteriorly; no dark stripe on frontoclypeal El Salvador: without locality (Förster 2001) –

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Guatemala: Santa Rosa (Calvert 1956), Suchitepéquez plete to incomplete at basal 30-50% of mesanepister- (USNM), Retalhuleu (Calvert 1956), Guatemala num; mesepimeral and metepimeral complete and as (MLPA*; Calvert 1956), Chimaltenango (RWGC), Solola wide as 25% of their sclerites (figs. 51-52). Abdomen (DRPC), Quezaltenango (Calvert 1956), Zacapa reddish brown with light blue and yellow to greenish (Calvert 1956), Alta Verapaz (DRPC), and Huehuete- yellow spots; PL spots in III-V (figs. 129b, d); posterior nango (DRPC) – Mexico: Chiapas (DRPC; RWGC; portion of inner carina of ventral terga V-VII linear in USNM), Guerrero (Calvert 1905), Veracruz (DRPC; female (fig. 167b). Ventral tubercle of abdominal seg- RWGC; USNM; Calvert 1905; Williamson 1908), Puebla ment I as high as 50% of its length (fig. 244); distal (USNM), Morelos (RWGC*; USNM), Michoacan (DRPC), portion of ventral margin of anterior lamina spine Jalisco (USNM), and Durango (DRPC; RWGC). convex (fig. 327); tip of hamular anterior process pointed in ventral view (fig. 287b); auricles with two- Rhionaeschna multicolor (Hagen) comb. n. three teeth (fig. 205). Male cercus with ventral process (figs. 11-he, 51-52-th, 129-ab, 167-te, 205-ge, 244-tg, 287-ha, at distal 25% projected posteriorly (fig. 368b), dis- 327-as, 368-ce, 445-Mp) tance between base of crest and posterior base of ven- Aeshna multicolor Hagen, 1861: 121-122 (description (&: tral process shorter than distance between base of lectotype ( (here designated) U.S.A.: Pecos River, Western process and tip of cercus, whole cercus tip ventrally Texas (Capt. Pope exp.) (MCZ; 8833) [examined: lecto- bent, dorso-distal crest higher than base of cercus in type (, syntypes &]); Calvert 1905: 180, 183-184 (in lateral view (fig. 368c). Female cercus external margin part); Calvert 1907: 400 (in part); Williamson 1908: 301; slightly convex, tip pointed (fig. 368d). Martin 1908: 48-49, 83; Walker 1912: 62, 66, 69, 71, 190-198 (redescription adults, description larva); Calvert Dimensions: head width: 9-10; HW length: 42-45; 1956: 13, 102-106; Walker 1958: 51-52, 55, 117-121 HW width: 13-13.8; HW pterostigma length: 3.5-3.8; (redescription (&larva); González Soriano & Novelo cerci length: ( 6.4-6.6, & 5.5-5.8; female cerci max- Gutiérrez 1991: 73, 80, 91; Novelo Gutiérrez & González imum width: 1.2; total length: 65-72. Soriano 1991: 114, 145-146; González Soriano 1993: Biology. – Adults feed over open fields from before 296; González Soriano & Novelo Gutiérrez 1996: 164; sunrise to after sunset, hanging under trees and bush Dunkle 2000: 50; Needham et al. 2000: 122, 126, 133, 144-145. during hot midday hours; males patrol large areas of Aeschna furcifera Karsch, 1891: 310. water in an irregular pattern (Dunkle 2000). They re- produce mainly in temporary pools, ponds, lakes, and Diagnosis. – Rhionaeschna multicolor and R. mutata slow streams, with emergent or floating plants differ from R. dugesi by the long ventral process of (González & Novelo 1991). One of the most com- male cercus (figs. 368-369c vs. fig. 366c) and the lin- mon species of Aeshnidae in western United States. ear inner lateral carina of ventral terga V-VII (figs. 167- Larva described by supposition by Walker (1912), 168 vs. fig. 165), and from R. jalapensis by the entire based on exuviae from Washington and by Musser tip of male cercus ventrally bent (figs. 368-369c vs. fig. (1962), based on larvae from Utah, USA. 367c) and presence of pale spots in abdominal seg- Remarks. – This species was described (Hagen ments III-V (figs. 129-130b, d vs. fig. 127b, d). They 1861: 122) from an unspecified number of specimens differ from each other by the height of ventral tubercle from ‘Pecos River, Western Texas (Capt. Pope); Up- I (as high as 50% of its length in R. multicolor, fig. 244; per Missouri; Mexico; Cordova (Saussure)’ without as high as 30% of its length in R. mutata, fig. 245), mention of the repository of type specimens. From the distance separating the posterior base of the ventral original type series, only four specimens are still in the process of male cercus from base of crest and tip of cer- Hagen collection in the MCZ (one male and two fe- cus (closer to base of crest in R. multicolor, fig. 368c; to males from Pecos River and one female from Upper tip of cercus in R. mutata, fig. 369c) and height of dor- Missouri). Williamson (1908) established that the fe- so-distal crest (higher than base width in R. multicolor, male from Upper Missouri belonged to a different fig. 368c; as high as base width in R. mutata, fig. species and was included in Walker’s description of 369c), external margin of female cercus (slightly con- Aeshna interrupta lineata (Walker 1912). Although the vex in R. multicolor, fig. 368d; linear in R. mutata, fig. locality mentioned by Hagen for the lectotype is the 369d), and T-spot stem width (narrower than vertex Pecos River in western Texas (32°N 104°W), it was in R. multicolor, fig. 11a; wider in R. mutata, fig. 12a). apparently collected in the vicinity of the present town Description. – Clypeus and frons light blue to yel- of Roswell, New Mexico, where Pope’s expedition oc- low, grayish blue spots lateral to white area surround- curred in 1854-1855 (see discussion in Garrison ing T-spot stem; T-stem narrower than vertex, gradu- 1994: 220). ally narrowing anteriorly; no dark stripe on Distribution [19-51°N, 98-125°W, 0-2650 m] (fig. frontoclypeal groove; vertex light blue or yellow with 445). – Mexico: Puebla (Walker 1912), Michoacan latero-posterior margins black (figs11a-c). Pale stripes (LACM; Calvert 1956), Distrito Federal (Calvert light blue to white or yellow; mesanepisternal com- 1956), Hidalgo (RWGC), Durango (Novelo &

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Gonzáles 1991), Baja California Sur (UAIC; Calvert gled (fig. 369d). Dimensions: head width: 9-10; HW 1956; Walker 1912), Jalisco (DRPC) – U.S.A.: Wash- length: 44-51; HW width: 13-13.9; HW pterostigma ington (Walker 1912), Oregon (RWGC; Walker 1912), length: 3.6-4.1; cerci length: ( 5.8-6.3, & 6.5-6.7; fe- Idaho (RWGC; Walker 1912), Colorado (Walker male cerci maximum width: 1.4; total length: 67-75. 1912), Utah (Walker 1912), California (RWGC*; UAIC; Biology. – Males patrol low over vegetation with a Walker 1912), Arizona (RWGC*; UAIC), New Mexico leisurely erratic flight. Females usually perch on water (RWGC; Hagen 1861; Walker 1912), Texas (RWGC; lily flowers to lay eggs in the underwater part of the Hagen 1861; Walker 1912), Nebraska (RWGC*), stem. They breed in fishless ponds, usually with water Nevada (RWGC), Massachusetts (Needham et al. lilies, and occasionally in bog ponds (Dunkle 2000). A 2000), Iowa (Needham et al. 2000), Kansas (Need- rare species. The larva was described by Walker (1958). ham et al. 2000), Montana (Needham et al. 2000), Remarks. – According to the original description, Oklahoma (Needham et al. 2000), South Dakota the holotype female was deposited at the NMW. How- (Needham et al. 2000), and Wyoming (Needham et ever, Calvert (1905) found a female labeled as type at al. 2000) – Canada: British Columbia (Walker 1912) the MCZ and considered it the holotype, a decision and Alberta (Needham et al. 2000). with which Williamson (1908) agreed. Distribution [38-41°N, 74-84°W, 0-243 m] (fig. Rhionaeschna mutata (Hagen) comb. n. 445). – U.S.A.: Ohio (Walker 1912), Indiana (Walk- (figs. 12-he, 53-th, 130-ab, 168-te, 206-ge, 245-tg, 288-ha, 328-as, er 1912), Pennsylvania (MLPA; RWGC*), Ohio (RWGC), 369-ce, 445-Mp) New Jersey (RWGC*), Connecticut (Needham et al. Aeshna mutata Hagen, 1861: 124-125 (description &: holo- 2000), Illinois (Needham et al. 2000), Kentucky type & North America (MCZ; 8832)); Williamson 1908: (Needham et al. 2000), Maine (Needham et al. 2000), 301-306; Walker 1912: 62, 66, 71 198-202 (redescrip- Maryland (Needham et al. 2000), Michigan (Need- tion (&); Walker 1958: 52, 55, 121-125 (redescrip- ham et al. 2000), Missouri (Needham et al. 2000), tion(&, description larva); Beatty & Beatty: 1969: 149- New Hampshire (Needham et al. 2000), New York 151; Dunkle 2000: 51-52; Needham et al. 2000: 122, 125, 133, 145-146. (Needham et al. 2000), Rhode Island (Needham et al. [Aeshna multicolor. – Calvert 1905: 180, 183-184 (in part); 2000), Virginia (Needham et al. 2000), West Virginia Calvert 1907: 400 (in part); Martin 1908: 48 (in part). (Needham et al. 2000), and Wisconsin (Needham et Misidentification] al. 2000) – Canada: Ontario (Needham et al. 2000).

Diagnosis. – Discrimination of this species from 4. Neureclipa group the closely related R. multicolor is provided under that (= subgenus Neureclipa Calvert, 1952) species. Description. – Clypeus and frons light blue to yel- Supratriangles usually free, sometimes one or two low, grayish blue spots lateral to white area surround- with 1 or 2 crossveins (in 2-10% of specimens exam- ing T-spot stem; T-stem wider than vertex, gradually ined); two rows of cells between RP1 and RP2 in HW narrowing anteriorly; no dark stripe on frontoclypeal beginning at distal end of pterostigma or distal to it groove; vertex black with central portion light blue (fig. 399); male cerci with high dorso-distal crest (as (figs. 12a-c). Pale stripes light blue to white or yellow; high or higher than width of cercus at base), sub-basal mesanepisternal complete to incomplete at basal 30- tooth prominent, and lateral carina concave (figs. 50% of mesanepisternum; mesepimeral and 370-374). Five species: R. absoluta, R. bonariensis, R. metepimeral complete and as wide as 25% of their diffinis, R. elsia, R. galapagoensis. sclerites (fig. 53). Abdomen reddish brown with light blue and yellow to greenish yellow spots; PL spots in Rhionaeschna absoluta (Calvert) comb. n. III-V (figs. 130b, d); posterior portion of inner carina (figs. 13-he, 54-59-th, 131-ab, 169-te, 207-ge, 247-tg, 289-ha, of ventral terga V-VII linear (fig. 168b). Ventral tuber- 329-as, 370-ce, 446-Mp) cle of abdominal segment I as high as 30% of its length Aeshna diffinis absoluta Calvert 1952: 258-260 (description (fig. 245); distal portion of ventral margin of anterior (&: holotype ( PERU: vicinity of Concepción, 07/08- lamina spine convex (fig. 328); tip of hamular anteri- IV-1935, F. Woytkowski leg. (UMMZ) [examined: holo- or process pointed in ventral view (fig. 288b); auricles type (, allotype &, paratypes (&]); Calvert 1956: 15, with two-three teeth (fig. 206). Male cercus with ven- 133-134; Rácenis 1959: 494. Aeshna absoluta Jurzitza 1990a: 353-372; Rodrigues Capítu- tral process at distal 25% projected posteriorly (369b), lo 1992: 37, 56; Muzón 1995: 5 (in part); Muzón 1997: distance between base of crest and apex of ventral 127-128, 132-133; Muzón & von Ellenrieder 1998: 23; process longer than distance between apex of process von Ellenrieder 2001a: 301-305; b: 423-426, 431-433. and tip of cercus, whole cercus tip ventrally bent, dor- [Aeshna diffinis. – Ris 1904: 24, 26-29 (in part); Ris 1908: so-distal crest as high as base of cercus in lateral view 523, 526-527 (in part); Ris 1913: 84-85 (in part); Navás (fig. 369c). Female cercus outer margin linear, tip an- 1920d: 53; Navás 1930a: 125; Fraser 1947: 433, 446; Ro-

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drigues Capítulo 1980: 9, 16-17, 20; Abenante & Philip- head width: 6.9-8.4; HW length: 32-41.1; HW width: pi 1982: 151; Rodrigues Capítulo & Muzón 1989b: 147; Rodrigues Capítulo et al. 1991: 62, 66 (in part); Ro- 10.5-13.1; HW pterostigma length: 2.8-4.3; cerci drigues Capítulo 1992: 37, 57 (in part); Pérez D’A. & length: ( 4.3-5.1, & 3.8-4.9; female cerci maximum Mutschke 1993-94: 63-68; Muzón 1995: 5-6 (in part); width: 0.9; total length: 43.1-59.8. Muzón & von Ellenrieder 1998: 23 (in part). Misidentifi- Biology. – Rhionaeschna absoluta occupies one of cation] the largest distribution areas within Rhionaeschna, [Aeshna diffinis diffinis. – Calvert 1956: 127, 233 (in part); breeding in a wide variety of environments from tem- Rácenis 1959: 494. Misidentification] [Aeshna elsia. – Rodrigues Capítulo 1992: 37, 57; Daigle et porary ponds and pools to creeks, streams and small al. 1997: 12 (in part). Misidentification] rivers. I collected foraging adults flying along an At- lantic beach in Chubut province. They patrolled Diagnosis. – Rhionaeschna absoluta and R. diffinis small temporary ponds on sides of a road, creeks and can be distinguished from the other Neureclipa species small streams in the Patagonian steppe, dams for irri- by the stem of T-spot with concave sides (figs. 13, gation and banks of small rivers in Northwestern Ar- 15a), and the dorso-distal crest of male cerci higher gentina. Larvae were found under stones or crawling than cerci width at base (figs. 370, 372b). They differ among aquatic vegetation (von Ellenrieder 2001b). A from each other by the comparatively narrower ventral common species. terga (ratio length/width of IV 4.2-4.5) and maximum Distribution [0-52°S, 56-78°W, 0-4500 m] (fig. width of V-VI occupies the distal portion in R. absolu- 446). – Argentina: Santa Cruz (Muzón 1995), ta (fig. 169). In R. diffinis the ventral terga are com- Chubut (DRPC; IMLA; MACN; MLPA*; RWGC), Neuquén paratively wider (ratio length/width of IV 2.4-2.6) and (DRPC; MLPA*; RWGC; USNM; Muzón 1995), Río Ne- maximum width of V-VI occupies the basal portion gro (DRPC; MLPA*; RWGC; USNM), Buenos Aires (DRPC; (fig. 171). Females differ by the dorsal color pattern of MACN; MLPA*; RWGC; USNM), La Pampa (MLPA), Santa second abdominal segment; longitudinal black stripe Fe (MACN), Entre Ríos (MLPA), Mendoza (DRPC; IMLA; between transverse carina and posterior margin in R. IADIZA; MLPA; USNM; Ris 1908; Fraser 1947; Jurzitza absoluta (fig. 131c) but only at posterior margin in R. 1990a), San Juan (IMLA); La Rioja (MACN; MLPA), diffinis (fig. 133c), and shape of cercus; tips pointed in Catamarca (DRPC), Santiago del Estero (MLPA), Cór- R. absoluta, fig. 370c, and rounded in R. diffinis, 372c. doba (Navás 1930a), Tucumán (DRPC; IMLA; MLPA; Description. – Clypeal lobes angled (fig. 13c); RWGC), Salta (IMLA; MLPA*; USNM), and Jujuy (IMLA; clypeus light blue or yellow, frons yellow, grayish MLPA) – Chile: Magallanes: Magallanes (Perez D’A. & green spots lateral to yellow area surrounding T-spot Mutschke 1993-94), and Santiago: Santiago (MNNS), stem; T-stem widening posteriorly, sides slightly con- Biobío: Arauco (UMMZ) – Uruguay: San José (Calvert cave; vertex yellow with latero-posterior margins black 1956), and Montevideo (ANSP; Calvert 1956) – Bo- (fig. 13a); wide black stripe on frontoclypeal groove; livia: La Paz: Sud Yungas (RWGC) - Peru: Lima (DRPC), wide black stripe on fronto-ocular groove (figs. 13b- Junín (RWGC; UMMZ; Calvert 1952; 1956), Cuzco c). Pale mesanepisternal stripes at basal 25-33% of (UMMZ; USNM; Calvert 1956), Ayacucho (IMLA; mesanepisternum to absent; mesepimeral and UMMZ), and Arequipa (Calvert 1952; 1956) - metepimeral stripes light blue to yellow, linear and Ecuador: Imbabura (KJTC). narrow (except in teneral specimens where they are wide: fig. 54), complete to absent (figs. 55-59). Mem- Rhionaeschna bonariensis (Rambur) comb. n. branule dark except basal 30% pale. Abdomen reddish (figs. 14-he, 60-th, 132-ab, 170-te, 208-ge, 248-tg, 290-ha, 330-as, brown with light blue and yellow spots; female seg- 371-ce, 399-wi, 447-Mp) ment II with a medio longitudinal dorsal black stripe Aeshna bonariensis Rambur, 1842: 204 (description (&: between transverse carina and posterior margin (fig. lectotype ( (here designated) ARGENTINA: Buenos Aires 131c). Abdominal ventral terga narrow (ratio length/ (ISNB) [examined: lectotype (, paralectotypes &]); Hagen width of V at basal 25% higher than 4), maximum 1861: 314; Hagen 1875: 39; Ris 1904: 24-25; Martin width of V-VI at distal 66%, basal 30% of inner and 1908: 51-52; Ris 1908: 523, 525-526; Calvert 1909: 221; Martin 1911: 12; Navás 1911: 476, 478; Cockerell 1913: outer lateral carinae of IV concave (figs. 169a-b). Ven- 580; Ris 1913: 85; Navás 1917a: 187; Ris 1918: 158; tral tubercle of abdominal segment I bearing numer- Navás 1920a: 152; Navás 1920c: 267; Martin 1921: 23; ous denticles spread over its posterior surface (fig. Campion 1922: 292-293; Martin 1923: 109; Navás 1927: 247); dorsal margin of anterior lamina spine linear 23; Gazulla & Ruiz 1928: 290; Navás 1929c: 220; Pirión (fig. 329); tip of hamular anterior process pointed in 1933: 82; Fraser 1947: 433, 448; Calvert 1952: 257-258; ventral view (fig. 289); auricles with two teeth (fig. Herrera et al. 1955-66: 81; Calvert 1956: 14, 140-144, 227; Fraser 1957: 159; Martins Costa 1971: 194; Paulson 207). Male cercus lacking pale basal spot in outer sur- 1977: 175; Abenante 1978: 29-48; Abenante 1980: 105- face (fig. 370a); dorso-distal crest rising suddenly, 149; Rodrigues Capítulo 1980: 1-21; Abenante & Philip- higher than base of cercus in lateral view (fig. 370b). pi 1982: 151; Jurzitza 1989: 7; Jurzitza 1990a: 353-372; Tip of female cercus pointed (fig. 370c). Dimensions: Rodrigues Capítulo et al. 1991: 62, 66; Watson 1992:

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455, 657-462; Pitzke-Widdig 1992: 116, 122; Rodrigues (1978), Rodrigues Capítulo (1980) and von Ellen- Capítulo 1992: 37, 56; Mola & Papeschi 1994: 185-188; Mola 1995: 47-54; Muzón 1995: 5; Muzón & von Ellen- rieder (2001b). It breeds in ponds and temporary rieder 1997: 146; Carvalho & Nessimian 1998: 7; Muzón pools. Adults are common in grasslands of central Ar- & von Ellenrieder 1998: 23; Martins Costa & Santos gentina, where they are usually found flying together 1999: 4; Martins Costa, et al. 2000: 13; von Ellenrieder with Rhionaeschna confusa. I observed a mass migration 2001a: 301, 305-308; b: 424, 426-427, 431-433. of hundreds of teneral R. bonariensis and R. confusa Aeschna dicrostigma Selys in Martin, 1908: 53 (nomen perching on bushes and grasses near the shore of the nudum). Neureclipa bonariensis. – Navás 1911: 476, 478. Rio de La Plata (February 1998). An abundant species. Aeshna bonaerensis var. lutea Navás, 1920b: 11; Navás Distribution [15-41°S, 41-72°W, 0-2000 m] (fig. 1920c: 267; Navás 1928: 340. 447). – Argentina: Río Negro (MLPA; Hagen 1875), Buenos Aires (DRPC; MACN; MLPA*; RWGC; USNM; Campion 1922; Navás 1927), Entre Ríos (MACN; Diagnosis. – Rhionaeschna bonariensis shares with R. MLPA; USNM; Fraser 1947), Mendoza (Ris 1908), San absoluta and R. diffinis a linear dorsal margin of ante- Juan (Calvert 1956), Córdoba (MACN; MLPA; UMMZ; rior lamina spine (figs. 329-331) and ventral tubercle Calvert 1956; Navás 1929c), Santa Fé (MACN; MLPA; of I with numerous denticles (figs. 247-249), and with USNM), Santiago del Estero (MLPA*; Navás 1920b; Ris R. elsia and R. galapagoensis the dorso-distal crest of 1913); Chaco (Calvert 1956; Ris 1913), Formosa cercus as high as width of cercus at base (figs. 371, (IMLA; FCN), Catamarca (RWGC; Ris 1918), La Rioja 373-374b) and a complete medio-dorsal longitudinal (MLPA; MACN), Tucumán (IMLA; Calvert 1956), Salta yellow stripe of female abdominal segment II (figs. (IMLA; MLPA*; USNM; Pitzke-Widdig 1992), Jujuy 132-134c). The narrow stripe on fronto-ocular groove (Calvert 1956), Corrientes (MACN; MLPA*; Calvert (fig. 14b), parallel sided T-spot (fig. 14a) and pale spot 1956), and Misiones (MLPA; USNM) – Uruguay: San on outer base of male cercus (fig. 371a-b) distinguish José (Calvert 1956), Montevideo (MNRJ; Abenante R. bonariensis from all other Neureclipa species. 1978; Calvert 1956; Hagen 1875; Ris 1904; 1908), Description. – Clypeal lobes angled (fig. 14c); Colonia (Mola 1995; Mola & Papeshi 1994), and clypeus and frons light blue and yellow, grayish blue Florida (UMMZ) – Brazil: Rio Grande do Sul (DRPC; spots lateral to yellow area surrounding T-spot stem; MNRJ; RMNH; USNM; ZMH; Hagen 1861; Martins Cos- T-stem usually parallel sided, in some cases slightly ta 1971), Santa Catarina (Calvert 1956), Paraná narrowed anteriorly; vertex yellow with latero-posteri- (MNRJ; Calvert 1956), Sâo Paulo (DRPC; MNRJ; USNM; or margins black (fig. 14a); no dark stripe on fronto- Calvert 1956; Martins Costa et al. 2000), Minas clypeal groove; black stripe on fronto-ocular groove Gerais (MNRJ; Calvert 1909), Río de Janeiro (Carvalho narrow (figs. 14b-c). Pale mesanepisternal stripes at & Nessimian 1998; Martins Costa & Santos 1999), basal 25-33% of mesanepisternum to absent; and Goias (MNRJ) – Paraguay: Paraguarí (Calvert mesepimeral and metepimeral stripes yellow to light 1956; RWGC), Central (DRPC) – Bolivia: Santa Cruz blue, linear, complete and narrow (fig. 60). Mem- dept.: Florida (RWGC) – Chile: Araucanía: Cautín branule dark except basal 20% pale. Abdomen pale (Herrera et al. 1955-66; Jurzitza 1989), and Val- reddish brown with light blue and yellow spots; female paraiso: Quillota (Pirión 1933) and Valparaiso (MNNS; segment II with a complete medio longitudinal dorsal Gazulla & Ruiz 1928; Jurzitza 1989; Martin 1923). yellow stripe (figs. 132c). Abdominal ventral terga narrow (ratio length/width of V at basal 25% higher Rhionaeschna diffinis (Rambur) comb. n. than 4), maximum width of V-VI at distal 66%, basal (figs. 15-he, 61-67-th, 133-ab, 171-te, 209-ge, 249-tg, 291-ha, 30% of inner and outer lateral carinae of IV concave 331-as, 372-ce, 447-Mp) (figs. 170a-b). Ventral tubercle of abdominal segment Aeshna diffinis Rambur, 1842: 203-204 (description (&: I bearing numerous denticles spread over its posterior lectotype & (here designated) Chile (ISNB) [examined]); surface (fig. 248); dorsal margin of anterior lamina Hagen 1861: 314; Hagen 1875: 38-39 (in part); Selys spine linear (fig. 330); tip of hamular anterior process 1895: 61; Porter 1897: 13; Porter 1899: 181; Ris 1904: rounded in ventral view (fig. 290b); auricles with two 24, 26-29 (in part: a form); Martin 1908: 43-44, fig. 40 (in part); Ris 1908: 523, 526-527 (in part); Navás 1917b: teeth (fig. 208). Male cercus with a pale basal spot in 38; Campion 1922: 292-293; Martin 1923: 109; Pirión outer surface (fig. 371a); dorso-distal crest rising grad- 1928: 96-97; Gazulla & Ruiz 1928: 290; Navás 1929a: ually, as high as base of cercus in lateral view (fig. 145; Navás 1929b: 326; Navás 1930b: 350; Pirión 1933: 371b). Tip of female cercus pointed (fig. 371c). Di- 81; Ureta 1935: 93; Needham & Bullock 1943: 358-359; mensions: head width: 6.9-8; HW length: 33.9-39.7; Herrera et al. 1955-66: 81; Peña 1962: 4; Fraser 1957: HW width: 10.5-13; HW pterostigma length: 3.2-4.2; 159; Paulson 1977: 175 (in part); Davies & Tobin 1985: ( & 3; Jurzitza 1990a: 353-372, figs. 2-6; Rodrigues Capítulo cerci length: 4.5-5.2, 4.5-5.7; female cerci maxi- et al. 1991: 62, 66 (in part); Rodrigues Capítulo 1992: 37, mum width: 0.95; total length: 50-57.8. 57; fig. 175a, b (in part); Muzón 1995: 5-6 (in part); Biology. – The larva was described by Abenante

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Muzón 1997: 127-128, 132-133; Muzón & von Ellen- and Río Negro, Argentina. An abundant species. rieder 1997: 146; Muzón & von Ellenrieder 1998: 23 (in part); von Ellenrieder 2001a: 301, 309-312; b: 424, 427- Remarks. – The description of R. diffinis (Rambur 428, 431-433. 1842) mentioned only female characters and two fe- Aeschna configurata Hagen, 1861: 314 (nomen nudum). males with type labels were found in the collection of [Aeschna bonariensis. – Bolivar 1884: 5-6. Misidentification] the ISNB. The two females belong to two different Aeshna diffinis diffinis. – Calvert 1952: 258; Calvert 1956: species; the one here designated as lectotype is in poor 14, 126-133, 201-203, 232 (in part). Böttger & Jurzitza condition and has only head and part of thorax, and 1967: 35-37; Jurzitza 1975: 10-11; Jurzitza 1989: 8-9 (in part); Watson 1992: 455, 657-462. has the locality label of ‘Chili’ mentioned by Rambur [Aeshna absoluta. – Muzón 1995: 5 (in part). Misidentifica- in his description. The paralectotype female, which tion] belongs to R. absoluta, is labeled ‘Ae. diffinis?’ in Ram- bur’s hand. Distribution [27-46°S, 67-74°W, 80-1850 m] (fig. Diagnosis. – Rhionaeschna diffinis is distinguished 447). – Chile: Aisén: Aisén (Jurzitza 1989; Ureta from all the other Neureclipa species by the wide ven- 1935), Coihaique (MLPA; RWGC; Calvert 1956), and tral terga (fig. 171). Further differences are discussed General Carrera (MNNS), De los Lagos region: Valdivia under R. absoluta. (IEUM; MNNS; MLPA*; RWGC; USNM; Ris 1904; Böttger Description. – Clypeal lobes angled (15c); clypeus & Jurzitza 1967; Jurzitza 1975; 1989; 1990a; Navás and frons light blue to yellow, grayish green spots lat- 1930b), Osorno (IEUM; MNNS; MLPA; USNM), Llanqui- eral to yellow area surrounding T-spot stem; T-stem hue (IEUM; MLPA; MNNS; USNM), Palena (UMMZ; widening posteriorly, sides slightly concave; vertex yel- RWGC), and Chiloe (IEUM; UMMZ; USNM), Araucanía: low with latero-posterior margins black (fig. 15a); Cautín (IEUM; RWGC; USNM), and Malleco (IEUM; wide black stripe on frontoclypeal groove; wide black MLPA*; RWGC; UMMZ; USNM; Calvert 1956; Navás stripe on fronto-ocular groove (Figs15b-c). Pale 1929b), Biobío: Arauco (IEUM; MLPA*; UMMZ; USNM; mesanepisternal stripes at basal 25-33% of mesanepis- Watson 1992), Biobío (IEUM; UMMZ), Concepción ternum to absent; mesepimeral and metepimeral (RWGC; USNM; ZMH; Calvert 1956; Jurzitza 1975), and stripes yellow to light blue, linear and narrow (except Ñuble (IEUM; UMMZ; USNM; Navás 1929a), Maule: in teneral specimens where they are wide, figs. 61-62), Linares (IEUM; MNNS; RMNH; UMMZ; USNM), complete to absent (figs. 63-67). Membranule dark Cauquenes (UMMZ; USNM), Talca (IEUM; UMMZ; except basal 30% pale. Abdomen reddish brown to USNM), and Curicó (IEUM; MNNS; RWGC; UMMZ), Gen- black with light blue and yellow or greenish yellow eral O’Higgins: Colchagua (UMMZ; USNM; Jurzitza spots; female segment II with a medio longitudinal 1975; 1989), Cardenal Caro (IEUM), and Cachapoal dorsal black stripe at posterior 25% of segment (figs. (IEUM; MNNS; UMMZ), Santiago: Maipo (IEUM; MNNS; 133c). Abdominal ventral terga wide (ratio length/ UMMZ), Melipilla (UMMZ), Cordillera (IEUM; UMMZ; width of V at basal 25% less than 3.5 mm), maximum USNM), Talagante (IEUM), Santiago (IEUM; MNNS; width of V-VI at basal 30%, basal 30% of inner lateral UMMZ; USNM), and Chacabuco (IEUM; MNNS; MNNS), carinae of IV concave, of outer convex (figs. 171a-b). Valparaiso: San Antonio (IEUM; UMMZ), Valparaiso Ventral tubercle of abdominal segment I bearing nu- (IEUM; MNNS), Quillota (RWGC; UMMZ), and Liberta- merous denticles spread over its posterior surface (fig. dor Aconcagua (UMMZ), Coquimbo: Coquimbo 249); dorsal margin of anterior lamina spine linear (RWGC), Choapa (IEUM; UMMZ), Limarí (IEUM; MNNS), (fig. 331); tip of hamular anterior process pointed in and Elqui (MNNS), and Atacama: Huasco (ZMH). – Ar- ventral view (fig. 291b); auricles with two teeth (fig. gentina: Chubut (IMLA; MLPA*; RWGC*; USNM; Campi- 209). Male cercus lacking pale basal spot in outer sur- on 1922; Muzón 1995), Río Negro (LACM; MLPA; face (fig. 372a); dorso-distal crest rising suddenly, USNM; Muzón 1995), and Neuquén (MLPA*; USNM; higher than base of cercus in lateral view (fig. 372b). Muzón 1995). Tip of female cercus rounded (fig. 372c). Dimensions: head width: 7.6-8.4; HW length: 34.2-39.4; HW width: Rhionaeschna elsia (Calvert) comb. n. 11-13.7; HW pterostigma length: 2.5-3.7; cerci length: (figs. 16-he, 68-73-th, 134-ab, 172-te, 210-ge, 250-tg, 292-ha, ( 4.3-5.3, & 3.4-4.3; female cerci maximum width: 332-as, 373-ce, 448-Mp) 0.95; total length: 49.6-59.9. Aeshna (Neureclipa) elsia Calvert, 1952: 260-262 (descrip- Biology. – I have seen adults flying along streams, tion (&: holotype ( PERU: vicinity of Pacasmayo, small rivers, and shores of lakes in Southern Argentina 7°24’0S 79°34’0W 7 m, 20-V-1936, F. Woytkowski leg. and Chile, where the species is common and usually (UMMZ) [examined: holotype (, allotype &, paratypes ( found with R. variegata. Calvert (1956) provided a de- (UMMZ; RWGC)]); Calvert 1956: 14, 134-137; Rácenis 1959: 494; Paulson 1977: 175; Davies & Tobin 1985: 3; scription of the larva based on material from Chile, Jurzitza 1989: 9; Daigle et al. 1997: 12 (in part); von El- and von Ellenrieder (2001b) provided an additional lenrieder 2000: 353-357 (redescription (&). description based on reared specimens from Neuquén

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Diagnosis. – Rhionaeschna elsia and R. galapagoensis Calvert 1952, 1956), and Iquique (Calvert 1952, are distinguished from the remaining Neureclipa 1956). Not confirmed records: Maule: Curicó species by the T-stem with convex sides (figs. 16-17a), (Calvert 1952, 1956) and Talca (Calvert 1952, 1956) clypeal lobes rounded (figs. 16-17c) and dorsal margin – Peru: Arequipa (RWGC), Lima (IMLA; UMMZ*; USNM; of anterior lamina spine concave (figs. 332-333). They Calvert 1952, 1956), Huanuco (Calvert 1956), and differ from each other in (character states of R. elsia La Libertad (IMLA; RWGC*; USNM; Calvert 1952, mentioned first): shape of dorso-distal crest of male 1956) – Ecuador: Imbabura (KJTC). cercus (rising suddenly, fig. 373b vs. gradually, fig. 374b), tip of female cercus (pointed, fig. 373c vs. Rhionaeschna galapagoensis (Currie) comb. n. rounded, fig. 374c), tip of hamular anterior process in (figs. 17-he, 74-75-th, 135-ab, 173-te, 211-ge, 251-tg, 293-ha, posterior and ventral view (pointed, fig. 292 vs. 333-as, 374-ce, 448-Mp) rounded, fig. 293), and color of frontoclypeal groove Aeshna galapagoensis Currie, 1901: 382-385 (description ( (lacking black stripe, fig. 16b vs. bearing black stripe, &: lectotype ( ECUADOR: Chatham Island, Islas Galápa- fig. 17), and membranule (dark with basal 30% pale gos, 0°49’60S 89°25’60W 143 m, 26-V-1899 (USNM) vs. dark with external margin pale). [examined: lectotype (, paralectotype &]); Needham Description. – Clypeal lobes rounded (fig. 16c); 1904: 695-696 (description larva); Martin 1908: 50, 84; Campos 1922: 6, 33; Calvert 1956: 14, 137-140; Belle clypeus and frons light blue to light yellow, grayish 1991: 2. green spots lateral to yellow area surrounding T-spot stem; T-stem widening posteriorly, with convex sides; Diagnosis. – See under R. elsia. vertex yellow with latero-posterior margins black (fig. Description. – Clypeal lobes rounded (fig. 17c); 16a); no dark stripe on frontoclypeal groove; black clypeus and frons light blue to pale brown, brown stripe on fronto-ocular groove wide (figs. 16b-c). Pale spots lateral to yellow area surrounding T-spot stem; mesanepisternal stripes at basal 25% of mesanepister- T-stem widening posteriorly, with convex sides; vertex num to absent; mesepimeral and metepimeral stripes yellow or light blue with latero-posterior margins light blue to yellow and linear, from complete to re- black (fig. 17a); black stripe on frontoclypeal groove; duced to basal spots (figs. 68-73). Membranule dark wide black stripe on fronto-ocular groove (figs. 17b- except basal 30% pale. Abdomen pale reddish brown c). Pale mesanepisternal stripes absent; mesepimeral with light blue and yellow spots; female segment II with and metepimeral stripes light blue, complete to absent a complete medio longitudinal dorsal yellow stripe (fig. (figs. 74-75). Membranule dark except basal 20-25% 134c). Abdominal ventral terga narrow (ratio length/ pale. Abdomen dark brown with light blue spots; fe- width of V at basal 25% higher than 4), maximum male segment II with a complete medio longitudinal width of V-VI at distal 0.66, basal 30% of inner and dorsal yellow stripe. Abdominal ventral terga narrow outer lateral carinae of IV concave (figs. 172a-b). Ven- (ratio length/width of V at basal 25% higher than 4), tral tubercle of abdominal segment I bearing few denti- maximum width of V-VI at distal 66%, basal 30% of cles (10 or less) restricted to its apex (fig. 250); dorsal inner and outer lateral carinae of IV concave (figs. margin of anterior lamina spine concave (fig. 332); tip 173a-b). Ventral tubercle of abdominal segment I of hamular anterior process rounded in posterior and bearing few denticles (10 or less) restricted to its apex ventral view (fig. 292b); auricles with two teeth (fig. (fig. 251); dorsal margin of anterior lamina spine con- 210). Male cercus lacking pale basal spot in outer sur- cave (fig. 333); tip of hamular anterior process point- face (fig. 373a); dorso-distal crest rising suddenly, as ed in posterior and ventral view (fig. 293b); auricles high as base of cercus in lateral view (fig. 373b). Tip of with two teeth (fig. 211). Male cercus lacking pale female cercus pointed (fig. 373c). Dimensions: head basal spot in outer surface (fig. 374a); dorso-distal width: 7.8-8.5; HW length: 35.8-39; HW width: 12- crest rising suddenly, as high as base of cercus in later- 13.6; HW pterostigma length: 2.8-3.4; cerci length: ( al view (fig. 374b). Tip of female cercus rounded (fig. 4.3-4.6, & 4.2-4.5; female cerci maximum width: 374c). Dimensions: head width: 8-8.3; HW length: 0.65; total length: 49.85-53.75. 39.5-40.5; HW width: 12-13.2; HW pterostigma Biology. – The type series was collected at a pond length: 3.2-3.4; cerci length: ( 5-5.3, & 4.5; female between sand dunes in Peru (Calvert 1952). Rare in cerci maximum width: 0.65; total length: 58-60. collections. Larva is still unknown. Biology. – Needham (1904) provided a brief de- Remarks. – Calvert’s (1952; 1956) records from scription based on half grown larvae from Chatham Southern Chile are considered doubtful and have not Island. He mentioned the absence of an apical tooth been included here, since no specimens have been in labial palps and small lateral spines on abdominal found among the abundant material examined from segments VI-VII as diagnostic, but since these charac- this area. ters can change through different instars it is impossi- Distribution [0-19°S, 69-79°W, 7-1544 m] (fig. ble to know if they are characteristic of the last instar. 448). – Chile: Tarapacá: Arica (IEUM; MNHN; UMMZ; Rare in collections.

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Distribution [0°S, 89-91°W, 143-600 m] (fig. two characters mentioned above are unique for this 448). – Ecuador: Colón (USNM*; Currie 1901; Belle species within the variegata group. 1991; Calvert 1956). Description. – Clypeal lobes angled; clypeus and frons yellow, grayish blue spots lateral to yellow area surrounding T-spot stem; T-stem gradually narrowing 5. R. variegata group anteriorly, with sides approximately linear; frontal ca- Two rows of cells between RP1 and RP2 in HW be- rina angled in dorsal view; confluence of eyes long ginning at distal end of pterostigma or distal to it (fig. (frons + vertex/eyes + occipital triangle less than 1; 400); male cerci with high dorso-distal crest (as high eyes /occipital triangle equal to 2); vertex yellow with or higher than width of cercus at base), and lateral ca- latero-posterior margins black (fig. 18a); black stripe rina concave (figs. 375-381). Seven species: R. brasi- on frontoclypeal groove; narrow black stripe on fron- liensis, R. californica, R. confusa, R. marchali, R. peral- to-ocular groove (figs. 18b-c). Pale mesanepisternal ta, R. tinti, R. variegata. stripes at basal 30-50% of mesanepisternum; mesepimeral and metepimeral stripes yellow, con- Rhionaeschna brasiliensis (von Ellenrieder & stricted or divided at mid-length (figs. 76-78). Mem- Martins Costa) comb. n. branule dark except basal 50-60% pale. Abdomen pale (figs. 18-he, 76-78-th, 136-ab, 174-te, 212-ge, 252-tg, 294-ha, reddish brown with light blue and yellow spots; medio 334-as, 375-ce, 450-Mp) longitudinal dorsal yellow stripe of female segment II Aeshna brasiliensis von Ellenrieder & Martins Costa, 2002: 1- incomplete; segments V-IX with PL spots; ML and MD 8 (description (&larva: holotype ( Brazil: Rio Tainha- spots separated from each other (figs. 136a-d). Ab- 900 m, 29º26’60S 54º34’60W (Rio Grande do Sul state), dominal ventral terga narrow (ratio length/ width of V 20-I-1958, N.D. Santos leg. (MNRJ) [examined: holotype at basal 25% higher than 4) (figs. 174a-b). Ventral tu- (, allotype &, paratypes (&(MNRJ, MLPA, RWGC)]). [Aeshna peralta. – Calvert 1956: 67-72 (in part); Santos bercle of abdominal segment I bearing numerous den- 1966a: 97; Santos 1966b: 123-124 (description of larva); ticles spread over its posterior surface and higher than Santos 1988: 266, 276; Muzón & von Ellenrieder 1997: genital lobe (fig. 252); distal portion of ventral margin 143-146; Carvalho & Nessimian 1998: 3-28; Martins of anterior lamina spine concave to linear (fig. 334); Costa et al. 2000: 13. Misidentification] tip of hamular anterior process projected in ventral Aeshna sp. – von Ellenrieder 2001a: 318; 2001b: 430. view, projection lower than its width at base (fig. 294b); auricles with two teeth (fig. 212). Male cercus Diagnosis. – Rhionaeschna brasiliensis shares with R. inner margin diverging gradually from outer margin californica and R. confusa narrow ventral terga (ratio to mid-length of cercus, distal portion of inner margin length/width of V at basal 25% higher than four, figs. linear; tip pointed (fig. 375a); no pale basal spot in 174-176), male cercus with inner margin diverging outer surface; dorso-distal crest rising suddenly at dis- gradually from outer margin and tip of cercus pointed tal 20% of cercus, as high as base of cercus in lateral in both sexes (figs. 375-377a, c). It also shares with R. view; sub-basal tooth pointed (fig. 375b). Tip of fe- californica the stem of T-spot with sides approximately male cercus pointed (fig. 375c). Dimensions: head linear (figs. 18-19), vs. convex in R. confusa (fig. 20), width: 8.3-9; HW length: 35.7-41; HW width: 11.5-14; dark stripe on frontoclypeal groove (figs. 18-19b-c) vs. HW pterostigma length: 2.8-4; cerci length: ( 4.58- dark stripe absent in R. confusa (figs. 20b-c), mem- 5.1, & 4.6-5.6; female cerci maximum width: 1.15- branule dark with basal 50-60% pale vs. basal 20% 1.3; total length: 59.9-65.5. pale in R. confusa, mesanepisternal stripes incomplete Biology. – Larva described by Santos (1966b) and at basal 30-50% of mesanepisternum (figs. 76-80) vs. von Ellenrieder & Costa (2002) based on reared lar- complete in R. confusa (fig. 81), and distal portion of vae collected at small pools with abundant aquatic anterior lamina spine linear to concave (figs. 334-335) vegetation formed by cascading waters characteristic vs. convex in R. confusa (fig. 336). Rhionaeschna of the highlands within the region of Itatiaia. Rare in brasiliensis and R. confusa possess PL spots on segments collections. VII-IX (figs. 136, 138b, d) vs. absent in R. californica Distribution [22-29ºS, 44-51ºW, 752-2200 m] (137b, d). Rhionaeschna brasiliensis can be distin- (fig. 450). – Brazil: Rio Grande do Sul (MNRJ*; guished from R. californica and R. confusa by the angled RWGC), Paraná (MNRJ), São Paulo (MNRJ*), Rio de frontal carina in dorsal view (fig. 18a) vs. rounded (figs. Janeiro (MLPA; MNRJ*; NMW; RWGC). 19-20a), epimeral stripes constricted or divided at mid- length (figs. 76-78) vs. linear (figs. 79-80), dorso-distal Rhionaeschna californica (Calvert) comb. n. carina of cercus rising abruptly at distal 20% of cercus (figs. 19-he, 79-80-th, 137-ab, 175-te, 213-ge, 253-tg, 295-ha, (fig. 375b) vs. rising gradually at distal 30% (figs. 376- 335-as, 377-ce, 449-Mp) 377b), and presence of a sub-basal tooth in male cercus Aeshna californica Hagen, 1875: 33 (nomen nudum); Calvert, (fig. 375b), vs. tooth vestigial (figs. 376-377). The first

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1895: 504-507 (description (&: holotype ( U.S.A.: Mt. emergent plants (Dunkle 2000). Collection records Tamalpais (California state), 25-V, C. Jack (ANSP; 9251)); Needham & Hart 1901: 45 (description larva); Martin show this to be an early spring species in the western 1908: 47, 51, 84; Walker 1912: 61, 66, 69, 71, 184-190 United States. A common species. (redescription adults and larva); Calvert 1956:10, 22-25; Distribution [31-54°N, 110-131°W, 0-1720 m] Walker 1958: 51, 54, 113-117; Musser 1962: 25-26, fig. (fig. 449). – Mexico: Baja California Norte (Dunkle 16; Dunkle 2000: 52, Pl. 5; Needham et al. 2000: 122, 2000; Needham et al. 2000) – U.S.A.: Arizona (Need- 125, 133, 138-139. ham & Hart 1901); California (DRPC; LACM; RWGC*; Diagnosis. – Discrimination of R. californica from Calvert 1895; Kennedy 1917), Oregon (DRPC; other species is given under R. brasiliensis. RWGC*; Walker 1912), Nevada (RWGC), Idaho (DRPC; Description. – Clypeal lobes angled; clypeus and RWGC), Utah (RWGC; Calvert 1895), Washington frons yellow to greenish yellow or light blue, grayish (DRPC; RWGC; Calvert 1956; Needham et al. 2000), blue spots lateral to yellow area surrounding T-spot Colorado (Needham et al. 2000), Montana (Need- stem; T-stem gradually narrowing anteriorly, with ham et al. 2000), South Dakota (Needham et al. sides approximately linear; frontal carina rounded in 2000), Wyoming (Needham et al. 2000) – Canada: dorsal view; confluence of eyes long (frons + vertex/ British Columbia (DRPC; LACM; Walker 1912). eyes + occipital triangle less than 1; eyes/occipital tri- angle equal to 2); vertex yellow with latero-posterior Rhionaeschna confusa (Rambur) comb. n. margins black (fig. 19a); black stripe on frontoclypeal (figs. 20-he, 81-th, 138-ab, 176-te, 214-ge, 254-tg, 296-ha, 336-as, groove; wide black stripe on fronto-ocular groove 376-ce, 450-Mp) (figs. 19b-c). Pale mesanepisternal stripes at basal 30- Aeshna confusa Rambur, 1842: 205 (description (&: holo- 50% of mesanepisternum; mesepimeral and type & ARGENTINA: Buenos Aires (ISNB) [examined]); Ha- metepimeral stripes yellow to light blue and linear gen 1861: 314; Hagen 1875: 39; Ris 1904: 24-25; Martin mesepimeral complete or incomplete, metepimeral 1908: 52-53; Ris 1908: 523, 525; Martin 1911: 12; Ris complete (figs. 79-80). Membranule dark except basal 1913: 85; Navás 1916: 17; Navás 1917a: 187; Martin 1921: 22; Martin 1923: 109; Navás 1927: 23; Gazulla & 50-60% pale. Abdomen pale reddish brown with light Ruiz 1928: 290; Navás 1929c: 220; Pirión 1933: 81; blue to greenish yellow spots; medio longitudinal dor- Needham & Bullock 1943: 358-359; Fraser 1947: 433, sal yellow stripe of female segment II incomplete (fig. 448; Mielewczyk 1978: 29; Calvert 1956: 11, 30-34, 230; 137c); PL spots absent in segments V-IX; ML and MD Fraser 1957: 159; Paulson 1977: 175; Abenante & Philip- spots separated from each other (figs. 137b-d). Ab- pi 1982: 151; Davies & Tobin 1985: 2; Jurzitza 1989: 7- dominal ventral terga narrow (ratio length/width of V 8; Mola 1991: 10; Rodrigues Capítulo et al. 1991: 62, 66; Watson 1992: 454, 657-462; Rodrigues Capítulo 1992: at basal 25% higher than 4) (figs. 175a-b). Ventral tu- 39, 56; Mola & Papeschi 1994: 185-188; Mola 1995: 47- bercle of abdominal segment I bearing numerous den- 48, 54; Muzón 1997: 127-128, 132; Muzón & von Ellen- ticles spread over its posterior surface and higher than rieder 1998: 23; von Ellenrieder 2001a: 301, 312-314; b: genital lobe (fig. 253); distal portion of ventral margin 424, 428-429, 431-433. of anterior lamina spine concave to linear (fig. 335); tip of hamular anterior process projected in ventral Diagnosis. – Discrimination from other species view, projection lower than its width at base (fig. provided under R. brasiliensis. 295b); auricles with two teeth (fig. 213). Male cercus Description. – Clypeal lobes angled; clypeus and inner margin diverging gradually from outer margin frons yellow to light blue, grayish blue spots lateral to to mid-length of cercus, distal portion of inner margin yellow area surrounding T-spot stem; T-stem approx- linear; tip pointed (fig. 377a); no pale basal spot in imately parallel sided, slightly narrowed anteriorly, outer surface; dorso-distal crest rising gradually at dis- with convex sides; frontal carina evenly curved in dor- tal 0.30 of cercus, as high as base of cercus in lateral sal view; confluence of eyes long (frons + vertex/eyes + view; sub-basal tooth vestigial (fig. 377b). Tip of fe- occipital triangle less than 1; eyes/occipital triangle male cercus pointed (fig. 377c). equal to 2); vertex black with central portion yellow Dimensions: head width: 7.3-8.5; HW length: (fig. 20a); no dark stripe on frontoclypeal groove; 32.5-41.5; HW width: 11.2-14.4; HW pterostigma black stripe on fronto-ocular groove wide (figs. 20b- length: 2.8-4.6; cerci length: ( 3.8-4.5, & 3-4.8; fe- c). Pale mesanepisternal stripes complete; mesepimer- male cerci maximum width: 1; total length: 48.2-64. al and metepimeral stripes yellow to light blue, linear, Biology. – Larvae breed in small lakes, ponds, complete and narrow (fig. 81). Membranule dark ex- stream pools, and marshes. Last larval instar was de- cept basal 20% pale. Abdomen pale reddish brown scribed by Needham & Hart (1901) based on one with light blue and yellow spots; medio longitudinal reared larva from Tombstone, Arizona, USA, and by dorsal yellow stripe of female segment II complete (fig. Musser (1962), based on larvae from Utah. Adults for- 138c); segments V-IX with PL spots; ML and MD spots age over roads and clearings. Males patrol close to wa- separated from each other (figs. 138b, d). Abdominal ter without a definite beat along shore and among ventral terga narrow (ratio length/width of V at basal

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25% higher than 4) (figs. 176a-b). Ventral tubercle of Janeiro, 22º52’0S 43º15’0W 370 m (Navás 1916). abdominal segment I bearing numerous denticles spread over its posterior surface and higher than geni- Rhionaeschna marchali (Rambur) comb. n. tal lobe (fig. 254); distal portion of ventral margin of (figs. 21-he, 82-th, 139-ab, 177-te, 215-ge, 255-tg, 297-ha, 337-as, anterior lamina spine convex (fig. 336); tip of hamular 378-ce, 451-Mp) anterior process projected in ventral view, projection Aeshna marchali Rambur, 1842: 203 (description (: holo- lower than its width at base (fig. 296b); auricles with type ( COLOMBIA (ISNB) [examined]); Hagen 1861: 314; two teeth (fig. 214). Male cercus inner margin diverg- 1875: 34; Martin 1908: 45, 84; Ris 1913: 83; Ris 1918: ing gradually from outer margin to mid-length of cer- 159-160; Campos 1922: 6, 33; Navás 1934: 3; 1935: 33; cus, distal portion of inner margin with a slight con- Schmidt 1952: 257; Rácenis 1953: 35, 37; Soukup 1954: 16; Calvert 1956: 12, 58-67, Map 3; Rácenis 1958; cavity; tip pointed (fig. 376a); pale basal spot in outer Limongi 1983: 95, 101, 104, 106-107 (description lar- surface; dorso-distal crest rising gradually at distal va); De Marmels 1988: 101; Daigle et al. 1997: 11. 30% of cercus, as high as base of cercus in lateral view; [Aeshna variegata. – Calvert 1956: 90 (in part: teneral ( sub-basal tooth vestigial (fig. 376b). Tip of female cer- from Chapare). Misidentification] cus pointed (fig. 376c). Dimensions: head width: 7.3- [Aeshna cornigera. – Martin 1908: 46 (in part). Misidentifi- 8.1; HW length: 32.5-41.5; HW width: 11.2-14.4; HW cation] pterostigma length: 2.8-4.6; cerci length: ( 3.8-4.5, & 3-4.8; female cerci maximum width: 1; total length: Diagnosis. – Rhionaeschna marchali, R. peralta, R. 48.2-59.8. tinti, and R. variegata differ from R. brasiliensis, R. cal- Biology. – Larva occurs in ponds and temporary ifornica, and R. confusa by several characters detailed pools and was described by von Ellenrieder (2001b) under R. brasiliensis. The following combination of based on reared specimens from Buenos Aires, Ar- characters allow for the separation of R. marchali from gentina. Adults are commonly seen flying in company R. peralta, R. tinti, and R. variegata (second contrast- with R. bonariensis over grasslands in central Argenti- ing character states for other species): frontoclypeal na. Mielewczyk (1978) reported a mass migration of groove lacking a dark stripe (fig. 21b-c) vs. stripe pre- R. confusa 60 miles from the shore of the Rio de La sent (figs. 22-24b-c), projection of tip of hamular an- Plata and I observed another in which hundreds of terior process 50% as high (C) as width (D) at anteri- teneral specimens of R. bonariensis and R. confusa ar- or margin (fig. 297b) vs. 30% as high as width at base rived and perched in bushes and grasses near shore in (figs. 298-300b), and male cercus with marked distal the Río de La Plata (February 1998). concavity at distal portion of inner margin (fig. 378a) Distribution [22-41°S, 43-73°W, 5-1402 m] (fig. vs. slight concavity (figs. 379-381a). 450). – Chile: Aisén: Cohiaique (Calvert 1956), De Description. – Clypeal lobes angled; clypeus and Los Lagos: Llanquihue (MNNS), Valdivia (MNNS; Jurz- frons yellow to greenish yellow, greenish blue spots itza 1989), and Osorno (USNM), Araucanía: Cautín lateral to yellow area surrounding T-spot stem; T- (IEUM; Calvert 1956; Fraser 1957; Jurzitza 1989) and stem approximately parallel sided, slightly narrowed Malleco (USNM; Calvert 1956; Jurzitza 1989), Biobío: anteriorly, with linear sides; frontal carina evenly Biobío (UMMZ), Concepción (UMMZ; USNM; Calvert curved in dorsal view; confluence of eyes long (frons + 1956; Jurzitza 1989), Arauco (USNM), and Ñuble vertex/eyes + occipital triangle less than 1; eyes/occip- (IEUM; UMMZ; USNM), Maule: Cauquenes (UMMZ; ital triangle equal to 2); vertex yellow with latero-pos- USNM), Talca (IEUM; Calvert 1956; Jurzitza 1989), terior margins black (fig. 21a); no dark stripe on fron- and Curicó (Hagen 1875; Ris 1904), Santiago: Santi- toclypeal groove; black stripe on fronto-ocular groove ago (IEUM), and Valparaiso: Valparaiso (MNNS; Gazul- wide (figs. 21b-c). Pale mesanepisternal stripes incom- la & Ruiz 1928; Jurzitza 1989) and Quillota (MNNS; plete at basal 30-50% of mesanepisternum; Jurzitza 1989; Pirión 1933) – Argentina: Buenos Aires mesepimeral and metepimeral stripes yellow, (MACN; MLPA*; UMMZ; USNM; Calvert 1956; Hagen mesepimeral complete or incomplete and sinuous, 1875; Martin 1908; Mola 1995; Mola & Papeshi metepimeral complete and with posterior side concave 1994; Ris 1904; 1908; 1913; Scott 1934), Entre Ríos (fig. 82). Membranule dark except basal 30% pale. (MACN; MLPA; Fraser 1947), Santa Fé (Navás 1917a), Abdomen pale reddish brown with light blue and yel- Córdoba (Hagen 1875; Navás 1927; 1929c), Tu- low spots; medio longitudinal dorsal yellow stripe of cumán (IMLA), and Misiones (MACN) – Uruguay: female segment II complete (fig. 139c); segments V-VI, Montevideo (Hagen 1875; Mielewczyk 1978; Ris VIII-IX with PL spots; ML and MD spots separated from 1904; 1908), Colonia (Mola 1995; Mola & Papeshi each other (figs. 139b, d). Abdominal ventral terga 1994) – Brazil: Rio Grande do Sul state: Pelotas, wide (ratio length/width of V at basal 25% less than 31º46’0S 52º19’60W (RMNH; Watson 1992). Santa 3.8) (figs. 177a-b). Ventral tubercle of abdominal seg- Catarina state: Nova Teutonia, 27º11’0S 52º13’0 W ment I bearing numerous denticles spread over its pos- 400 m (Calvert 1956). Rio de Janeiro state: Rio de terior surface and higher than genital lobe (fig. 255);

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distal portion of ventral margin of anterior lamina 1956; Ris 1918); Caldas (Calvert 1956; Navás 1935), spine convex (fig. 337); tip of hamular anterior Boyaca (Ris 1918), Santander (Calvert 1956; Ris process projected in ventral view, projection as high as 1918), Norte de Santander (Ris 1918) – Venezuela: its width at base (fig. 297b); auricles with two teeth Táchira (Rácenis 1958; De Marmels 1988), Mérida (fig. 215). Male cercus inner margin diverging sud- (DRPC; Rácenis 1953); Distrito Federal (Limongi denly from outer margin to mid-length of cercus, 1983; Rácenis 1953). where a ‘heel’ is formed, distal portion of inner margin with a marked concavity; tip angled (fig. 378a); no Rhionaeschna peralta (Ris) comb. n. pale basal spot in outer surface; dorso-distal crest ris- (figs. 22-he, 83-85-th, 140-ab, 178-te, 216-ge, 256-tg, 298-ha, ing gradually at distal 30% of cercus, as high as base of 338-as, 379-ce, 400-wi, 452-Mp) cercus in lateral view; sub-basal tooth pointed (fig. Aeshna peralta Ris, 1918: 159-162 (description (&: lecto- 378b). Tip of female cercus rounded (fig. 378c). Di- type ( (here designated) BOLIVIA: La Paz, 3600 m, 1912, mensions: head width: 8-8.4; HW length: 38-40.5; HW A.S. Fassl. leg. (SMFD; 16049) [examined: lectotype (, width: 12-14.2; HW pterostigma length: 2.8-3.12; cer- paralectotypes (&]); Hincks 1934: 80; Schmidt 1952: ci length: ( 4.4-4.8, & 4.88-5.12; female cerci maxi- 258; Calvert 1956: 67-72 (in part: records from Peru). mum width: 1-1.1; total length: 56-63.5. [Aeshna variegata. – Jurzitza 1990: 385-392 (synonymy with variegata; in part, records from Peru); von Ellenrieder Biology. – Campos (1922) recorded it as the most 2001: 317-318 (in part, records from Peru). Misidentifi- abundant aeshnid from the interandean plateaus of cation] Ecuador where he recorded it flying over wet soils, Examined material (17 ( 16 &). – Types (SMFD): 1 ( crop pastures, and coursing along creeks and streams. Lectotype (&16049) Bolivia, La Paz, 3600 m, 1912, A.S. The following notes were recorded by the collector Fe- Fassl.; 1 ( Paralectotype (&16050), 2 & Paralectotypes lix Woytkowski who collected it in Peru: ‘This species (&16051-52): Peru, Apurimac, 3500 m, 4-VI-1910, Gar- lepp. PERU: 1 ( 2 & Cuzco, stream at Sacsayhuaman, above is not faithful to any stable habitat, it prefers running Cuzco, 13-VI-1977, D.R. Paulson leg. (DRPC); 2 ( Cuzco, brooks with water-plants and sheltered by bush, but it 10-VII-1911, Yale Peruvian exp. leg. (USNM), same except is as frequently meet over stagnant ponds and stagnant 3510 m, 7-VII-1911 (DRPC); 1 ( 1 & Puno, Lago Titicaca at ditches, it frequents humid paths bordered with Puno, 3840 m, grassy shore, 23-V-1972, D.L. Pearson leg. shrub, and at last it reaches highest hills and there it (DRPC); 1 & Puno, 3900m, 26-X-1952, F. Blancas leg. lives on the outskirts of bush-clumps and in ravines’. (USNM); 2 ( Matucana, 26/27-V-1920, J.H. Williamson (UMMZ); 1 ( 2 & Camacani, 3700 m 19/21-XI-1955, L.E. Limongi (1983) described the larva by supposition Peña leg. (MLPA; UMMZ); 1 ( Chimo, 3700 m, 18-XI-1955, based on exuviae collected at temporary muddy bot- L.E. Peña leg. (UMMZ); 5 ( 2 & La Huerta, 3800 m, 24/28- tom pools with abundant emergent vegetation in for- XI-1955, L.E. Peña leg. (UMMZ); 1 ( 2 & Dept. Junin, Prov. est areas from Distrito Federal (Venezuela) where it Tarma, Vitor, 5/17-IV-1940, F. Woytkowski leg. (UMMZ). was sympatric with R. cornigera. BOLIVIA: 1 & Belen Station, Altiplano, 8/9-V-1952, J.A. Munro leg. (USNM); 2 & La Paz dept., Manco Capac prov., Remarks. – The type material of this species de- Copacabana, pequeño arroyo desembocadura en Lago Titi- posited at the ISNB consists of a single male. Since caca, 7-I-1972, Bulla leg. (MLPA); 1 ( 1 & Lago Titicaca, Rambur (1842) apparently described this species from 16°09’32S 69°05’05W 3800 m, von Ellenrieder leg. (MLPA). only this specimen, it is considered the holotype. Distribution [17°S-10°N, 65-80°W, 142-3778 m] Diagnosis. – Distinction of R. peralta from R. (fig. 451). – Bolivia: La Paz: Larecaja (MLPA) and brasiliensis, R. californica, R. confusa, and R. marchali is Cochabamba: Chapare (IMLA; UMMZ) – Peru: Are- given under R. brasiliensis and R. marchali. Rhion- quipa (RWGC; Calvert 1956), Cuzco (USNM; Schmidt aeschna peralta shares with R. variegata a T-spot stem 1952; Calvert 1956), Lima (UMMZ; Calvert 1956), that gradually narrows anteriorly (figs. 22, 24a) vs. Junín (Calvert 1956), Pasco (Calvert 1956), Ama- abruptly so in R. tinti (fig. 23a), thorax with stripes zonas (Calvert 1956), and Cajamarca (UMMZ, Calvert (figs. 83-85, 91) vs. basal and/or distal spots or no pale 1956) – Ecuador: Loja (UMMZ; MLPA; Calvert 1956; markings in R. tinti (figs. 86-90), and abdominal ML Ris 1918), Zamora (Ris 1918), El Oro (QCAZ), Azuay and MD spots separated (figs. 140, 142b, d) vs. conflu- (MLPA*; KJTC; USNM), Chimborazo (Ris 1918; Cam- ent in R. tinti (fig. 141b, d), and with a linear to con- pos 1922; Calvert 1956), Bolivar (Ris 1918), Tungu- cave ventral margin of the anterior lamina spine (fig. rahua (QCAZ; UMMZ; USNM; Calvert 1956; Campos 338, 340) vs. convex in R. variegata (fig. 339). It can be 1922), Napo (DRPC; KJTC*; QCAZ; USNM; Navás distinguished from both R. tinti and R. variegata by 1933), Cotopaxi (Campos 1922; Calvert 1956), Man- greater length of frons relative to eyes (ratio frons + ver- abi (UMMZ; MLPA), Pichincha (DRPC; KJTC; MLPA; tex/eyes + occipital triangle greater than 1 in R. peralta, QCAZ; Calvert 1956; Campos 1922), Imbabura (KJTC; less than 1 in R. tinti and in R. variegata; eyes/occipital Calvert 1956), and Carchi (Calvert 1956) – Colom- triangle less than 2 in R. peralta, fig. 22a, equal to 2 or bia: Cauca (USNM; Ris 1918), Valle (DRPC), Meta (Ris larger in R. tinti and R. variegata, figs. 23-24a). 1918), Tolima (DRPC), Cundinamarca (USNM; Calvert Redescription. – Head. Labium yellow to pale

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brown; teeth brown. Labrum yellow; basal margin at base (fig. 298b). Auricles with two teeth (fig. 216). with a transverse black stripe; distal margin bordered Cerci: Male cercus inner margin diverging suddenly with brown. Clypeus yellow; clypeal lobes angled. from outer margin to mid-length of cercus, where a Black stripe on frontoclypeal groove; wide black stripe ‘heel’ is formed, distal portion of inner margin with a on fronto-ocular groove (figs. 22b-c). Frons yellow, slight concavity; tip angled (fig. 379a); no pale basal grayish blue spots lateral to yellow area surrounding spot in outer surface; dorso-distal crest rising gradual- T-spot stem; T-stem gradually narrowing anteriorly, ly at distal 30% of cercus, as high as base of cercus in with concave sides. Black of spot extended ventrally lateral view; sub-basal tooth blunt (fig. 379b). Female on antefrons in some specimens. Frontal carina be- cercus lanceolate, with rounded tip (fig. 379c). Di- tween ante- and posfrons straight in anterior view and mensions (those of lectotype in brackets): head width: evenly curved in dorsal view. Vertex yellow with lat- 7.85-8.5 (8); HW length: 31-39 (36); HW width: 9- ero-posterior margins black. Occipital triangle yellow. 12.9 (12); HW pterostigma length: 2-2.8 (2); cerci Rear of the head black. Confluence of eyes short (frons length: ( 3.75-4.5 (4), & 3-4.5; female cerci maxi- + vertex/eyes + occipital triangle larger than 1; eyes/ mum width: 0.9; total length: 51.5-56.5 (53.3). occipital triangle less than 2) (fig. 22a). Biology. – Adults have been collected on quieter, Pterothorax. Reddish brown with yellow to light weedy, flat parts of streams and adjacent small marshy blue pale stripes, with dark brown areas over the ponds in dry areas of Peru and Bolivia. Rare in collec- grooves and surrounding the base of the pale stripes. tions. Larva unknown. Pale mesanepisternal stripes incomplete, occupying Remarks. – Rhionaeschna peralta was described by between 30-50% of the sclerite, slightly diverging Ris (1918) based on a series of five specimens collect- basally. Yellow mesepimeral stripes incomplete, occu- ed in Bolivia (La Paz) and Peru (Apurimac). In his pying the basal 75% of the sclerite, almost linear to monograph of Neotropical Aeshnidae, Calvert (1956) sinuous. Yellow or light blue metepimeral stripes com- extended the distribution area of this species to plete with posterior side slightly concave (figs. 83-85). Ecuador, Brazil, Chile, and Argentina. Following Small yellow spot ventral to the mesostigma. Legs Calvert’s key and diagnosis (1956), it was later record- black except coxae pale brown, and extensor surfaces ed from Chile by Jurzitza (1989), from Argentina by of femora yellow. Wings (fig. 400). Hyaline. C, ScP, Rodrigues Capítulo & Muzón (1989), and Rodrigues antenodals and nodal veins light yellow, rest black or Capítulo (1992), and from Brazil by Santos (1966b) dark reddish brown. Pterostigma orange-yellow, with and Costa et al. (2000). Santos (1966b) provided a de- black margins except costal margin pale yellow. Mem- scription of the last instar larva based on reared speci- branule basal 50% pale, distal 50% dark. mens from Brazil. Abdomen. Pale reddish brown with light blue and Rhionaeschna peralta was later synonymized with R. yellow spots; pattern as shown in figs. 140a-d. Ab- variegata by Jurzitza (1990b), based on examination domen segments V-IX with PL spots; ML and MD spots of a large series of several specimens from Chile (some separated from each other (figs. 140b, d). Medio lon- of which corresponded to R. peralta according to the gitudinal dorsal yellow stripe of female segment II in- key and diagnostic characters given by Calvert 1956), complete (fig. 140c). Males: Pale lateral spots of seg- eight from Argentina, and two from Peru. Calvert ments I and II and AL of III white-yellowish; pale dorsal (1956) considered the number of cell rows between spots of II-III yellow; lateral pale spots III-X white-light IRP2a-IRP2b (more than three in R. variegata, not blue; MD III-X yellow; PD III-X white light blue. Fe- more than three in R. peralta) and thoracic color pat- males: Pale lateral spots of I-II yellow ventrally, green- tern (pale antehumeral stripe absent in R. variegata, yellow dorsally; lateral spots III-X yellow; anterior por- present in R. peralta) as diagnostic. Jurzitza (1990b) tion of pale dorsal spots of II yellow, posterior portion examined venation, color pattern and shape of vesica light blue; remaining dorsal pale spots yellow. Ventral spermalis and ovipositor and failed to find significant terga wide (ratio length/width of V at basal 20% less differences. However, neither Calvert nor Jurzitza than 3.8 mm) (figs. 178a-b), reddish brown with yel- looked at the types of R. peralta. During a revision of lowish pale spots. Ventral tubercle of abdominal seg- the Patagonian species of Rhionaeschna (von Ellen- ment I bearing numerous denticles spread over its pos- rieder 2001), I found no specimen fitting the original terior surface and higher than genital lobe; genital lobe description of R. peralta among Patagonian represen- with 13-15 small denticles restricted to the distal 0.15 tatives and I suggested the possibility of an incorrect of segment II. (fig. 256). Male genital fossa: Anterior synonymy. lamina spines laterally compressed, pointed and sur- Examination of the type series of Rhionaeschna per- passing caudally the base of the hamuli anteriores, alta demonstrated that it represents a valid species and with distal portion of ventral margin linear to concave that its synonymy with R. variegata was in error. How- (fig. 338). Tip of hamular anterior process not pro- ever, the original description fails to provide good di- jected in ventral view, projection lower than its width agnostic characters and includes some mistakes (i.e. PL

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and PD spots widely confluent in all segments, which ear (fig. 340); tip of hamular anterior process project- are in fact separated in VI-VII or VIII in some of the type ed in ventral view, projection lower than its width at specimens, and cercus with parallel inner and outer base (fig. 299b); auricles with two teeth (fig. 217). margins and dorsal crest arising from midlength) that Male cercus inner margin diverging suddenly from probably contributed to the failure of subsequent outer margin to mid-length of cercus, where a ‘heel’ is workers in recognizing this species. The previous formed, distal portion of inner margin with a slight records of this species from Chile and Argentina cor- concavity; tip angled (fig. 380a); no pale basal spot in respond to R. variegata, those from Brazil to R. outer surface; dorso-distal crest rising gradually at dis- brasiliensis, and those from Peru to R. peralta. Rhion- tal 25% of cercus, as high as base of cercus in lateral aeschna peralta and R. variegata seem to be allopatric, view; sub-basal tooth prominent (fig. 380b). Tip of fe- the first restricted to Bolivia and Peru and the second male cercus rounded (fig. 380c). to Chile and Argentina (fig. 452). Dimensions: head width: 7.4-8.4; HW length: Distribution [11-16°S, 68-76°W, 1771-3960 m] 32.2-38.2; HW width: 10.8-13.1; HW pterostigma (fig. 452). – Peru: Apurimac (SMFD*), Cuzco (DRPC; length: 2.2-3.2; cerci length: ( 3.75-4.3, & 3.8-4.35; USNM*), Junin (UMMZ), Lima (UMMZ), and Puno female cerci maximum width: 0.85; total length: (DRPC; UMMZ; USNM) – Bolivia: La Paz: Manco Capac 43.5-54.8. (SMFD*; MLPA*; USNM*). Biology. – In Northern Chile, specimens were col- lected in desert farming areas as they flew over canals Rhionaeschna tinti (von Ellenrieder) comb. n. used for irrigating alfalfa crops. Larva unknown. Rare (figs. 23-he, 86-92-th, 141-ab, 179-te, 217-ge, 257-tg, 299-ha, in collections, but common in the field. 340-as, 380-ce, 451-Mp) Distribution [20-24°S, 67-70°W, 2350-4200 m] Aeshna tinti von Ellenrieder, 2000: 348-352, 357 (descrip- (fig. 451). – Chile: Antofagasta: Tocopilla (MNNS), El tion (&: holotype ( Chile: Tilopozo, 23°49’S 68°15’W Loa (IEUM*; MNNS*; MLPA*; RWGC; UMMZ*), and (El Loa prov., Antofagasta region), I-1996 (MLPA) [exam- Antofagasta (IEUM; MNNS), and Tarapacá: Iquique ined: holotype (, allotype &, paratypes (&(MLPA; (IEUM; MNNS). IEUM; MNNS; MNHN; RWGC; UMMZ)]). Rhionaeschna variegata (Fabricius) comb. n. Diagnosis. – Thorax color pattern with small basal (figs. 24-he, 91-th, 142-ab, 180-te, 218-ge, 258-tg, 300-ha, 339-as, and/or distal pale spots (figs. 86-90), and abdomen 381-ce, 431-au, 452-Mp) with ML and MD spots confluent (figs. 141b, d), are Aeshna variegata Fabricius, 1775: 425 (description (: holo- unique for this species within the group. Further dif- type ( Tierra del Fuego [type lost, Zimsen 1964]); Hagen ferences are elaborated under R. brasiliensis, R. mar- 1861: 314; Hagen 1875: 38; Martin 1908: 44-45; Martin chali and R. peralta. 1911: 11; Ris 1913: 81-84; Ris 1918: 159,162-163; Mar- Description. – Clypeal lobes angled; clypeus and tin 1921: 23; Martin 1923: 109; Navás 1926: 103-104; Calvert 1952: 258; Herrera et al. 1955-66: 81; Calvert frons light yellow or light blue, gray or light blue spots 1956: 12, 90-98, 230; Fraser 1957: 159; Fraser 1958: 195- lateral to yellow area surrounding T-spot stem; T- 198; Böttger & Jurzitza 1967: 37-38; Jurzitza 1975: 11; stem suddenly narrowing anteriorly, with convex Paulson 1977: 175; Jurzitza 1989: 10; Jurzitza 1990b: 385- sides; frontal carina evenly curved in dorsal view; con- 393; Rodrigues Capítulo et al. 1991: 62; Watson 1992: fluence of eyes long (frons + vertex/eyes + occipital tri- 455, 657-462; Rodrigues Capítulo 1992: 38, 57; Pérez angle less than 1; eyes /occipital triangle equal to 2); D’A. & Mutschke 1993-94: 63-68; Muzón 1995: 6; Muzón 1997: 127-128, 132-133; Muzón & von Ellen- vertex yellow with latero-posterior margins black (fig. rieder 1997: 143-146; Muzón & von Ellenrieder 1998: 23; 23a); black stripe on frontoclypeal groove present; von Ellenrieder 2001a: 301, 314-320; b: 424, 429-433. black stripe on fronto-ocular groove wide (figs. 23b- [Aeschna diffinis. – Mabille 1888: 3-6 (description); Ris c). Pale pterothoracic stripes absent or represented by 1904: 24, 26-29 (in part); Martin 1908: 43-44 (in part); basal and/or distal spots (figs. 86-92). Membranule Martin 1911: 11 (in part). Misidentification] dark except basal 20% pale. Abdomen light blue to [Aeschna diffinis var. risi Enderlein, 1912: 119. Misidentifi- light yellow with dark reddish brown to black spots; cation] [Aeshna diffinis risi. – Davies & Tobin 1985: 9-10. Misiden- medio longitudinal dorsal yellow stripe of female seg- tification] ment II incomplete (fig. 141c); segments V-IX with PL [Aeshna peralta. – Calvert 1956: 67-72 (in part: records from spots; ML and ML spots confluent in III-VII (figs. 141b, Chile and Argentina); Jurzitza 1989: 9-10; Rodrigues d). Abdominal ventral terga wide (ratio length/width Capítulo & Muzón 1989a: 76; Rodrigues Capítulo et al. of V at basal 25% less than 3.8 mm) (figs. 179a-b). 1991: 62. Misidentification] Ventral tubercle of abdominal segment I bearing nu- merous denticles spread over its posterior surface and Diagnosis. – Distinction from other species of the higher than genital lobe (fig. 257); distal portion of group is found under R. brasiliensis, R. marchali and ventral margin of anterior lamina spine concave to lin- R. peralta.

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Description. – Clypeal lobes angled; clypeus and xeric region of Northern Chile and the Patagonian frons light blue or yellow to greenish yellow, greenish steppe (0-100 mm and 100-270 mm respectively). gray or light blue spots lateral to yellow area surround- Larvae occur in streams, seepages and small ponds. ing T-spot stem; T-stem wide (dark form) to narrow The larva of the dark form was described by Muzón & (light form), gradually narrowing anteriorly, with lin- von Ellenrieder (1997) and that of the light form by ear sides; frontal carina evenly curved in dorsal view; von Ellenrieder (2001b) based on reared specimens confluence of eyes long (frons + vertex/eyes + occipital from Chile and Argentina. Rhionaeschna variegata is triangle less than 1; eyes /occipital triangle equal to the only species found in Tierra del Fuego, being the two); vertex yellow with latero-posterior margins most austral species of Odonata known. black (fig. 24a); black stripe on frontoclypeal groove; Distribution [22-54°S, 65-74°W, 0-3650 m] (fig. wide black stripe on fronto-ocular groove (figs. 24b- 452). – Light form: Argentina: Santa Cruz (IMLA; c). Pale mesanepisternal stripes incomplete at basal MLPA; Muzón 1995), Chubut (MLPA), Río Negro 50% of mesanepisternum to absent; mesepimeral and (MLPA; IMLA; Rodrigues Capítulo & Muzón 1989a; metepimeral stripes yellow, mesepimeral incomplete Muzón 1995), Mendoza (MACN; UMMZ), Catamarca and sinuous, metepimeral complete and with posteri- (UMMZ), Salta (Jurzitza 1990b), and Jujuy (MLPA) – or side concave (fig. 91). Membranule dark except Chile: Magallanes: Magallanes (IEUM; MNNS; PDA; basal 30-50% pale. Abdomen dark redish brown to Pérez D’Angelo 1993-94) and Antofagasta: Antofa- black with light blue and yellow to greenish yellow gasta (IEUM; MLPA) and El Loa (UMMZ). – Intermedi- spots; medio longitudinal dorsal yellow stripe of fe- ates: Chile: Libertador Gral. Bernando O’Higgins: male segment II incomplete (fig. 142e); segments V-IX Colchagua (IEUM), Maule: Curicó (IEUM; UMMZ), with (light form) or without (dark form) PL spots; ML Linares (MNNS), and Talca (IEUM; Jurzitza 1990b), and MD spots separated in III-VII (figs. 142c-d, f-g). Santiago: Chacabuco (IEUM; Jurzitza 1975), Santiago Abdominal ventral terga wide (ratio length/ width of V (IEUM; MNNS; UMMZ; Calvert 1956; Jurzitza 1989; at basal 25% less than 3.8 mm) (figs. 180a-b). Ventral 1990b), and Cordillera (IEUM; USNM), Valparaiso: tubercle of abdominal segment I bearing numerous Quillota (UMMZ; Jurzitza 1990b), San Felipe de denticles spread over its posterior surface and higher Aconcagua (Jurzitza 1990b), Valparaiso (IEUM; MNNS; than genital lobe (fig. 258); distal portion of ventral Calvert 1956; Jurzitza 1989; 1990b), and San Anto- margin of anterior lamina spine convex (fig. 339); tip nio (IEUM; UMMZ), and Coquimbo: Limarí (MNNS) of hamular anterior process projected in ventral view, and Choapa (MNNS; UMMZ; Jurzitza 1990b). – Dark projection lower than its width at base (fig. 300b); au- form: Argentina: Tierra del Fuego (IMLA; MLPA*; Ma- ricles with two-three teeth (fig. 218). Male cercus in- bille 1888; Muzón 1995; Ris 1904; 1913), Santa Cruz ner margin diverging suddenly from outer margin to (IMLA; MLPA*; MNRJ; Fraser 1958; Muzón 1995), mid-length of cercus, where a ‘heel’ is formed, distal Chubut (IMLA; Muzón 1995; Muzón & von Ellen- portion of inner margin with a slight concavity; tip an- rieder 1997), Neuquén (IMLA; MLPA*; RWGC; Calvert gled (fig. 381a); no pale basal spot in outer surface; 1956; Muzón 1995), and Río Negro (MLPA*; Muzón dorso-distal crest rising suddenly at distal 25% of cer- 1995). – Chile: Magallanes: Última Esperanza (IEUM; cus, higher than base of cercus in lateral view; sub- MNNS; MLPA; MNRJ; UMMZ; Ris 1913), Magallanes basal tooth prominent (fig. 381b). Tip of female cer- (IEUM; MNNS; UMMZ; Ris 1904; 1913; Watson 1992), cus rounded (fig. 381c). Dimensions: head width: and Tierra del Fuego (Navás 1916), Aisén: Aisén 8.4-9.5; HW length: 36.2-42.9; HW width: 11.4-14.1; (MNNS; MLPA*; RWGC; Jurzitza 1990b; Muzón & von HW pterostigma length: 2.4-3.7; cerci length: ( 4.4- Ellenrieder 1997), Coihaique (MNNS; MLPA*; RWGC), 5.7, & 4.4-5.5; female cerci maximum width: 1.1; to- General Carrera (MLPA; Calvert 1956; Jurzitza 1989; tal length: 55.8-67.2. 1990b), and Capitán Prat (Jurzitza 1990b), De Los La- Biology. – Rhionaeschna variegata is a highly vari- gos: Valdivia (MLPA; MNNS; UMMZ; Böttger & Jurzitza able species. Von Ellenrieder (2001a, b) described a 1967; Jurzitza 1975; 1989; 1990b), Osorno (MNNS; ‘light’ and ‘dark’ form. The dark form is confined to MLPA*; UMMZ; Jurzitza 1975; 1989; 1990b), Llanqui- humid areas along a narrow area of the subantarctic hue (IEUM; UMMZ; Jurzitza 1990b), Chiloe (IEUM; forest from Tierra del Fuego north to 36.5° S in Chile UMMZ), and Palena (MLPA*; RWGC; UMMZ; Muzón & and 39.6º S in Argentina. The light form appears to be von Ellenrieder 1997), Araucanía: Malleco (IEUM; associated with xeric conditions in the Patagonian MNNS; MLPA; UMMZ; Calvert 1956; Jurzitza 1989; steppe and in other dry areas of Argentina and north- 1990b) and Cautín (IEUM), and Biobío: Ñuble (MNNS; ern Chile. Intermediate specimens occur within an Jurzitza 1990b), Concepción (USNM; Jurzitza 1989; area that approximates the Central Chilean biogeo- 1990b; Ris 1918), and Arauco (IEUM; MLPA; UMMZ). graphic province, a zone of intermediate aridity (350- 1330 mm average yearly rainfall) between those of the humid Subantarctic forest (2000-3500 mm) and the

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6. R. cornigera group complete and as wide as 30-50% of mesanepisternum; mesepimeral and metepimeral stripes bright green, Two rows of cells between RP1 and RP2 in HW be- with linear or slightly undulated margins, and approx- ginning proximal to pterostigma or under it (figs. 401- imately uniform in width (fig. 92-94). Abdomen dark 402); male sterna IX posterior to genital opercula and reddish brown to black with bright green or yellowish X with a bright light-blue spot (figs. 143-150b); male green and light blue spots; male PL spots very small to cercus with low dorso-distal crest, no sub-basal tooth absent (figs. 143b-c). Ventral tubercle of abdominal and extreme apex pointed and ventrally bent (figs. segment I with anterior margin convex in lateral view, 382-389b); relatively narrow abdominal ventral terga about as high as 50% of its length (figs. 260-262); (width measured at narrowest point/length of ventral spines of anterior lamina well developed (fig. 341- terga IV less than 1%, except for males of vazquezae 342); tip of hamular anterior process rounded in pos- and pauloi, width 0.13 mm and 0.18 mm, respective- terior view (fig. 301-302a); auricles with two-three ly; narrow ventral terga are also found in draco and teeth (fig. 219-220). Male cercus with a concavity at some Marmaraeschna species) (figs. 181-188). Eight distal 30% or parallel sided, tip directed externally or species: R. cornigera, R. haarupi, R. manni, R. nubige- posteriorly (figs. 382a, f); dorso-distal crest smoothly na, R. pauloi, R. planaltica, R. psilus, R. vazquezae. curved, higher than base of cercus in lateral view, and developed along distal 25-33% of cercus (figs. 382b, Rhionaeschna cornigera (Brauer) comb. n. g). Female cercus longer than abdominal segments IX- (figs. 25,415-he, 92-94-th, 143-ab, 181-te, 219-220,418-ge, 260- X and shorter than VIII-X, with rounded or pointed tip, 262-tg, 301-302-ha, 341-342-as, 382-ce, 402-wi, 434-ve, 454-Mp) and maximum width at medial 30% (figs. 382c-e). Aeshna cornigera Brauer, 1865: 906 (in part, description ( Dimensions: head width: 8.4-9.55; HW length: 39.1- &: lectotype ( COLOMBIA (NMW) [examined: lectotype 46; HW width: 11.9-14; HW pterostigma length: 2.1-3; (, paralectotypes &]); Brauer 1866; 70-71; Calvert cerci length: ( 5-5.9, & 3.45-6.5; female cerci maxi- 1905: 179, 182-183 (in part); Martin 1908: 46, 84; Ris mum width: 0.6-1.1; total length: 58.5-67. 1918: 163-166 (in part); Calvert 1947: 7-8 (in part); St. Remarks. – Brauer described Aeshna cornigera Quentin 1970: 265; De Marmels 1982: 102-105 (de- scription larva). briefly in 1865, and redescribed it more extensively in Aeshna cornigera cornigera. – Calvert 1947: 7-8 (in part); 1866 based on a type series from ‘Columbien’. He did Calvert 1952: 255; Calvert 1956: 11, 34-43; De Marmels not indicate the number of specimens of the type se- 1988: 101; González Soriano 1993: 296; González Sori- ries but according to measurements he provided it in- ano & Novelo Gutiérrez 1996: 164. cluded only one male and two or more females. St. Quentin (1970) mentioned only two type specimens Diagnosis. – Rhionaeschna cornigera is most similar from the original series, and designated the male as to R. nubigena and R. planaltica. It can be distin- lectotype. I found one male and two females with type guished from R. nubigena by the approximately uni- labels in the NMW collection. Examination of the type form width of the mesepimeral stripe (figs. 92-94) vs. specimens revealed that one of the paralectotype fe- narrowed at midlength to 50% of its basal width in R. males belongs to R. planaltica. Calvert did not exam- nubigena (figs. 101-102), convex anterior contour of ine the types of R. cornigera (1947, 1952, 1956); he the ventral tubercle of abdominal segment I (figs. 260- based his distinction of this species from R. psilus 262) vs. concave in R. nubigena (fig. 267), dorso-distal (1947) and R. planaltica (1952) on comments of the crest of male cercus higher with a stronger curvature types sent to him by St. Quentin. According to St. (rising more abruptly, figs. 382b, g) vs. lower and Quentin’s letter (Calvert 1947) the thoracic stripes of gradually rising in R. nubigena (figs. 385b). Separation the types are not constricted so he apparently did not from R. planaltica is given under that species. The col- see the paralectotype female belonging to R. planaltica. or pattern of R. cornigera is similar to that of R. psilus The records of R. cornigera given by Calvert (1905, from which it is easily distinguished by the well-devel- 1947) and Ris (1904) previous to the descriptions of oped spines of the anterior lamina (figs. 341-342) vs. R. psilus (Calvert 1947) and R. planaltica (Calvert vestigial to absent in R. psilus (fig. 348), and much 1952) are not included here in the distribution list shorter and narrower female cercus (3.45-6.5 mm by since they encompassed all three species. The records 0.6-1.1 mm, figs. 382c-e, contra 7.2-8.26 mm by of Ris (1918) are included, since it is possible to dis- 1.15-1.33 mm in R. psilus, fig. 388c). criminate between the three species through his com- Description. – Clypeus and frons light blue to ments on the variability of the thoracic color pattern: bluish yellow, green spots lateral to yellow area sur- his records of R. cornigera from Mexico correspond to rounding T-spot stem; T-stem approximately parallel- R. psilus, the ones from Costa Rica and Colombia to sided; posterior 60% of vertex black (fig. 25a); fronto- R. cornigera, and the ones from Brazil, Bolivia and Ar- clypeal groove bordered by a narrow dark brown or gentina to R. planaltica. black stripe (figs. 25b-c). Pale mesanepisternal stripes Rhionaeschna cornigera shows the highest morpho-

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logical variability within the group: male cercus can be Rhionaeschna haarupi (Ris) comb. n. (figs. 26-he, 96-97-th, 144-ab, 182-te, 221-ge, 259-tg, 303-ha, parallel sided or with inner margin concave at medial 343-as, 383-ce, 453-Mp) 30%, and tip of cercus can be directed posteriorly or externally (figs. 382a-b, g-f); female cercus can be Aeshna haarupi Ris, 1908: 523-525 (description (&: lecto- type ( ARGENTINA: Estación Pedregal, 32°58’0S short with rounded tips or long and pointed (figs. 68°40’60W 695 m (Mendoza prov.), 01-XII-1906, P. Jo- 382d-e). Based on material I examined, northern spec- ergensen leg. (SMFD 16044)); Martin 1909: 212-213; imens (Mexico to Colombia and Venezuela) are on Martin 1911: 12; Calvert 1952: 254; Calvert 1956: 11, average larger, females possess a longer and more 25-27; Rodrigues Capítulo et al. 1991: 62; Rodrigues acuminate cercus, and male cercus is sometimes paral- Capítulo 1992: 38, 57; Muzón & von Ellenrieder 1998: lel sided, whereas southern specimens (Venezuela and 23; Muzón & von Ellenrieder 1999: 124. Ecuador to Bolivia) are on average smaller, female cer- cus shorter and rounded, and male cercus is not paral- Diagnosis. – This species can be distinguished from lel sided. However, I observed intergradation in these all the other species of the group by the abruptly nar- characters and have thus been unable to separate them rowed T-spot stem of frons (fig. 26a) and the short fe- into two distinct groups. male cercus (as long as or shorter than segments IX-X, Biology. – The species has been reported as rare in fig. 383c) vs. longer than IX-X in all other species (figs. Peru (Calvert 1956), where it was seen flying over wa- 382, 384-389c). Its thoracic and abdominal color pat- ter in shrub-covered areas with abundant irrigation tern (figs. 95-96, 144) are most similar to those of R. canals. According to De Marmels (1982; 1988), it is pauloi and R. planaltica (figs. 100, 103-105, 147-148) common and widely distributed in all the Andean but it differs from the latter two species by the longer mountain ranges of Venezuela. It was collected at dorso-distal crest of male cercus (figs. 383, 386b) and marshes and ponds with very soft muddy bottom, ventral terga shape of female (maximum width of ven- surrounded by herbs, ferns, grasses, wiht debris and tral terga IV-V at distal portion in R. haarupi, fig. 182b) grass tufts in Costa Rica, where males were seen hov- vs. at basal portion in R. planaltica (fig. 186B). Further ering and patrolling at about 30 cm above the water differentiation from R. pauloi is given under this (DRPC). Larva inhabits running waters and was de- species. scribed based on exuviae from Miranda, Venezuela Description. – Clypeus and frons light blue to (De Marmels 1982). bluish yellow, grayish light blue spots lateral to light Distribution [16°S-18°N, 67-94°W, 350-3600 m] yellow area surrounding T-spot stem; T-stem sudden- (fig. 454). – Bolivia: La Paz: Nor Yungas (Calvert ly narrowing distally; posterior 30% of vertex black 1956) – Peru: Cuzco (UMMZ; USNM; Calvert 1956), (fig. 26a); frontoclypeal groove bordered by a narrow Junín (Calvert 1956), Pasco (Calvert 1956), Ancash dark brown or black stripe (figs. 26b-c). Pale (Calvert 1956), Cajamarca (Calvert 1956) – Ecuador: mesanepisternal stripes complete or interrupted into Loja (Calvert 1956), El Oro (Calvert 1956), Napo two spots and as wide as 25-33% of mesanepisternum; (KJTC; MLPA*; QCAZ; Calvert 1956), Azuay (Calvert mesepimeral and metepimeral stripes bluish or yellow- 1956), Chimborazo (Calvert 1956), Bolivar (Calvert ish-green, with deep rounded indentations on anterior 1956), Tungurahua (Calvert 1956), Pastaza (UMMZ; margin (metepimeral) or anterior and posterior mar- Calvert 1956), Sucumbios (QCAZ), and Pichincha gins (mesepimeral) (fig. 96-97). Abdomen dark red- (NMW; QCAZ; RWGC) – Colombia: without locality dish brown with yellowish green or yellow and light (NMW*); Tolima (Ris 1918), Boyaca (UMMZ; Ris 1918; blue spots; male PL spots large (figs. 144b). Ventral tu- Calvert 1956), and Magdalena (UMMZ; Calvert 1947; bercle of abdominal segment I with linear or slightly 1956) – Venezuela: Táchira (De Marmels 1988; convex anterior margin in lateral view, about as high 2001), Mérida (NMW; Calvert 1956), Guárico (MLPA*), as 50% of its length (fig. 259); spines of anterior lam- Aragua (DRPC; RWGC; USNM), Miranda (De Marmels ina well developed (fig. 343); tip of hamular anterior 1982), and Distrito Federal (DRPC) – Panama: Chi- process rounded in posterior view (fig. 303a); auricles riquí (Calvert 1956) – Costa Rica: Cartago (INBC; with two teeth (fig. 221). Male cercus maximum USNM; Calvert 1956; Ris 1918), Alajuela (DRPC; INBC), width at medial 30%, tip directed externally (fig. Puntarenas (DRPC), San José (DRPC; UMMZ) – Nica- 383a); dorso-distal crest smoothly curved, higher than ragua: Matagalpa (Beckemeyer 2001) – El Salvador: base of cercus in lateral view, and developed along dis- without locality (Förster 2001) – Honduras: without tal 30% of cercus (fig. 383b). Female cercus as long as locality (Förster 2001) – Guatemala: Sacatepéquez or slightly shorter than abdominal segments IX-X, with (Calvert 1956), Quetzaltenango (Calvert 1956), Baja pointed tip, and maximum width at medial 30% (fig. Verapaz (Calvert 1956); Alta Verapaz (UMMZ, RWGC) – 383c). Dimensions: head width: 8-8.4; HW length: Mexico: Chiapas (UNAM) and Oaxaca (UMMZ). 36.5-41.2; HW width: 12.6-13; HW pterostigma length: 2.8-3; cerci length: ( 4.2-5, & 3.1-3.5; female cerci maximum width: 0.54; total length: 57-59.

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Biology. – This species occurs in arid areas of the tip of hamular anterior process pointed in posterior Preandean mountain range. I found it flying over lat- view (fig. 304a); auricles with three teeth (fig. 222). eral temporary pools and swampy areas with emer- Male cercus maximum width at distal 30%, with in- gent vegetation on the Calchaquí and Santa María ner and outer margins diverging along medial 30%, rivers in Salta and Tucumán provinces (Argentina). tip directed posteriorly (fig. 384a); dorso-distal crest Rare in collections and in the field. Larva unknown. angled, lower than base of cercus in lateral view, and Distribution [33-25°S, 65-68°W, 690-2030 m] developed along distal 0.30 of cercus (fig. 384b). Fe- (fig. 453). – Argentina: Mendoza (NMW; Calvert male cercus longer than abdominal segments VIII-X, 1956; Ris 1908), Catamarca (IMLA), Tucumán (MLPA; with maximum width at medial 30% (fig. 384c). Di- Calvert 1956); Salta (MLPA; RWGC*). mensions: head width: 8.75-9.5; HW length: 43-45; HW width: 12.8-13.6; HW pterostigma length: 2.75- Rhionaeschna manni (Williamson & Williamson) 3.3;cerci length: ( 5.4-6, & more than 6.25; female comb. n. cerci width: 1.6; total length: 62-66.7. (figs. 27-he, 97-99-th, 145-ab, 183-te, 222-ge, 263-tg, 304-ha, Remarks. – The distinction between R. psilus and 347-as, 384-ce, 456-Mp) R. manni based on the different degree of reduction Aeshna manni Williamson & Williamson, 1930: 26-34 (de- of anterior lamina spine (vestigial in R. psilus and ab- scription (&: holotype ( MEXICO: Los Parres [sic: Las sent in R. manni) by Calvert (1947; 1956) is incorrect Parras], 25°56’60N 111°27’0W 244 m (Baja California since there is a short spine in both species. state), 06-X-1923, J. H. Williamson leg. (UMMZ) [exam- Biology. – The type series was collected at irriga- ined: holotype ( allotype &]); Calvert 1947: 8-9; Calvert 1956: 12, 56-58; González Soriano 1993: 296; González tion dams, pools and swampy areas associated with Soriano & Novelo Gutiérrez 1996: 164. mountain creeks (Williamson & Williamson 1930), [Aeshna cornigera. – Calvert 1895: 507-508 (at least in part); and later found patrolling small stream pools in creek Calvert 1905: 182-183 (in part). Misidentification] canyon at bottom of oak zone (DRPC). Rare in collec- tions. Larva unknown. Diagnosis. – Rhionaeschna manni and R. psilus both Distribution [23-28°N, 110-113°W, 200-600 m] have reduced spines of the anterior lamina in the male (fig. 456). – Mexico: Baja California Sur (DRPC*; (figs. 347-348), and a long, wide, cercus in the female UMMZ*; Williamson & Williamson 1930; Calvert (longer than segments VIII-X and wider than 1 mm, 1895; 1947; 1956). figs. 384, 388c). They are differentiated from each other by the mesanepisternal and mesepimeral stripes Rhionaeschna nubigena (De Marmels) comb. n. which are light blue or greenish yellow and incomplete (figs. 28-he, 101-102-th, 146-ab, 184-te, 223-ge, 267-tg, 305-ha, in R. manni (fig. 97-99) vs. bright green and complete 344-as, 385-ce, 454-Mp) in R. psilus (fig. 106). Males also differ by the shape of Aeshna nubigena De Marmels, 1989b: 85-86 (description ( the cercus: inner and outer margins diverge gradually &: holotype ( VENEZUELA: Neblina base camp., toward distal 30% in R. manni (fig. 384a) but are ap- 00º49’50N 66º09’40W 140 m (Río Negro dept., Ama- proximately parallel to each other along distal 0.60 in zonas state), 15-II-1985, P.J. & P.M. Spangler, R.A. Fai- R. psilus (fig. 388a). Female cercus is wider in R. man- toure, W.E. Steiner leg. (MIZA)); De Marmels 1993: 156; (1.6 mm, fig. 384c) vs. 1.15-1.3 mm in De Marmels 1997: 151. ni R. psilus [Aeshna planaltica. – Calvert 1956: 43-50 (in part, & from (fig. 388a) and probably longer (broken in the allo- Mt. Roraima). Misidentification] type, only known female). Description. – Clypeus and frons light blue to pale Diagnosis. – Rhionaeschna nubigena is most similar brown, light blue spots lateral to yellow area sur- to R. cornigera and R. planaltica; distinction from R. rounding T-spot stem; T-stem approximately parallel- cornigera is provided under that species. It can be dis- sided; posterior 50% of vertex black (fig. 27a); no dark tinguished from R. planaltica by the mesepimeral stripe on frontoclypeal groove (figs. 27b-c). Pale stripe narrowed at midlength to half its basal width mesanepisternal stripes incomplete or interrupted into (figs. 101-102), with deep rounded indentations in R. two spots and as wide as 25-33% of mesanepisternum; planaltica (figs. 103-105), and dorso-distal crest of mesepimeral and metepimeral stripes bluish or yellow- male cercus extended for a longer distance (distal 30% ish-green, with linear to undulated margins, of cercus vs. about distal 25% in R. planaltica) and mesepimeral stripe incomplete, metepimeral complete gradual prominence (fig. 385b) vs. abruptly rising (figs. 97-99). Abdomen dark reddish brown to black crest in R. planaltica (fig. 386b). with yellowish green and light blue spots; male PL Description. – Clypeus and frons greenish yellow, spots large (figs. 145b). Ventral tubercle of abdominal grayish light blue spots lateral to yellow area sur- segment I with linear or slightly convex anterior mar- rounding T-spot stem; T-stem approximately parallel- gin in lateral view, about as high as 50% of its length sided; posterior 50% of vertex black (fig. 28a); fronto- (fig. 263); spines of anterior lamina vestigial (fig. 343); clypeal groove bordered by a narrow dark brown or

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black stripe (figs. 28b-c). Pale stripes complete and teriorly (fig. 387a) vs. directed externally in R. haarupi bright green; mesanepisternal as wide as 25-50% of and R. planaltica (figs. 383, 386a), male cercus with mesanepisternum; mesepimeral narrowed at mi- inner and outer margins parallel along distal 60% in dlength to half its basal width, metepimeral approxi- dorsal view (fig. 387a) vs. converging in the other two mately uniform in width (figs. 101-102). Abdomen species (figs. 383, 386a), and overall larger size (length dark reddish brown to black with bright green or yel- of cerci, HW, and body). lowish green and light blue spots; male PL spots small Description. – Clypeus and frons light blue to gray- to absent (figs. 146b). Ventral tubercle of abdominal ish yellow, light blue to bluish gray spots lateral to yel- segment I with concave anterior margin in lateral view, low area surrounding T-spot stem; T-stem gradually about as high as 50% of its length (fig. 267); spines of narrowing anteriorly; vertex black except medial 30% anterior lamina well developed (fig. 344); tip of hamu- yellow (fig. 29a); frontoclypeal groove bordered by a lar anterior process rounded in posterior view (fig. narrow dark brown or black stripe (figs. 29b-c). Pale 305a); auricles with two-three teeth (fig. 223). Male mesanepisternal stripes absent or incomplete and in- cercus maximum width along medial 30%, tip direct- terrupted into two spots and as wide as 25-33% of ed externally (fig. 385a); dorso-distal crest smoothly mesanepisternum; mesepimeral and metepimeral curved, higher than base of cercus in lateral view, and stripes yellowish-green, with deep rounded indenta- developed along distal 30% of cercus (fig. 385b). Fe- tions on anterior margin (mesepimeral) or anterior male cercus longer than abdominal segments IX-X and and posterior margins (metepimeral) (fig. 100). Ab- shorter than VIII-X, with angled tip bearing mucron on domen dark reddish brown with yellowish green and external margin, and maximum width at medial 30% light blue spots; male PL spots large (fig. 147b). Ven- (fig. 385c-d). Dimensions: head width: 8.85-9; HW tral tubercle of abdominal segment I with anterior length: 42-45; HW width: 12.8-14.6; HW pterostigma margin convex in lateral view, lower than 30% of its length: 2.5-2.7; cerci length: ( 5-5.5, & 3-3.73; fe- length (fig. 266); tip of hamular anterior process male cerci maximum width: 0.78; total length: 63-72. pointed in posterior view (fig. 306); spines of anterior Remarks. – Calvert (1956) included a female from lamina well developed (fig. 346). Auricles with two- Mt. Roraima as a doubtful record of R. planaltica for three teeth (fig. 224). Male cercus distal 60% parallel Venezuela. The specimen is preserved in alcohol, and sided, tip directed posteriorly (fig. 387a); dorso-distal its coloration has faded. However, shape and size of crest high (higher than base of cercus in lateral view) cerci (fig. 385c) agree well with those of R. nubigena, and smoothly curved along distal 25% of cercus (fig. and therefore is included here as this species. 387b). Female cercus longer than abdominal segments Biology. – Apparently restricted to the forests of IX-X and shorter than VIII-X, rounded tip with a Venezuelan tepuis. Larva unknown. minute medial spine, and maximum width at medial Distribution [0°S-5°N, 60-66°W, 140-2150 m] 30% (fig. 387c). Dimensions: head width: 9.4-9.6; (fig. 454). – Venezuela: Río Negro (USNM;De HW length: 44-53; HW width: 13.6-14.5; pterostigma Marmels 1989b), Amazonas (RWGC*; De Marmels length: 2.8-3.75; cerci length: ( 5.7-6, & 5-6.2; max- 1993; 1997), and Bolivar (ANSP*; Calvert 1956). imum width female cerci: 1.1; total length: 63-77. Biology. – This is apparently a mountain species in Rhionaeschna pauloi (Machado) comb. n. Brazil restricted to rocky streams and pools at eleva- (figs. 29-he, 100-th, 147-ab, 185-te, 224-ge, 266-tg, 306-ha, 346- tions above 1000 meters where the larvae occur. Rare as, 387-ce, 453-Mp) in collections. Some of the paratypes described by Aeshna pauloi Machado, 1994: 159-168 (description (&: Machado (1994) were reared from larvae but the lar- holotype ( BRAZIL: Serra do Cipó, 19°0’0S 43°38’60W va has not yet been described. The species is not en- 1300-1400 m, Santana do Riacho (Minas Gerais state), demic to the iron-rich streams mentioned by Macha- IV-1983, A. & P.A. & E. Machado leg. (ABBM) [exam- do, since I find a specimen from Sapucay, Paraguay, ined: (¶types (RWGC)]) and further collecting will probably show the species to be more widespread. Diagnosis. – Rhionaeschna pauloi can be distin- Distribution [20-25°S, 43-56°W 190-1400 m] guished from all the other species of the group by the (fig. 453). – Paraguay: Paraguarí (UMMZ) – Brazil: low ventral tubercle of segment I; lower than 30% of Paraná (ABBM) and Minas Gerais (ABBM). its length (fig. 266) vs. as high as 50% of its length in remaining species (figs. 259-265, 267-269). Thoracic Rhionaeschna planaltica (Calvert) comb. n. color pattern is similar to that of R. haarupi and R. (figs. 30-he, 103-105-th, 148-ab, 186-te, 225-ge, 264-265-tg, 307- planaltica (figs. 95-96, 100, 103-105); it differs from ha, 345-as, 386-ce, 401-wi, 453-Mp) these species by the rounded cercus tip of females (fig. Aeshna cornigera planaltica Calvert, 1952: 255-256 (descrip- 387c), which is pointed in R. haarupi and R. planalti- tion (&: holotype ( BRAZIL: Santa Catarina state: Nova ca (figs. 383c, 386c-e), tip of male cercus directed pos- Teutonia, 27º11’S 52º13’W 300-500 m), 01-XII-1935,

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F. Plaumann leg. (MCZ)); Calvert 1956: 11, 43-50; Ráce- 14; HW pterostigma length: 2.1-2.9; cerci length: ( nis 1959: 493-494; Santos 1970: 203; Jurzitza 1981: 117; De Marmels 1989a: 54-55; 1989b: 35; 1989c: 85; 1990b: 4.18-5.35, & 3.95-5.1; female cerci maximum width: 337; Mola 1991: 10; De Marmels 1992a: 64, figs. 47-48 0.7-0.95; total length: 57.5-65. (description larva); Rodrigues Capítulo et al. 1991: 62, Remarks. – Rhionaeschna planaltica was described 66; Rodrigues Capítulo 1992: 39, 57; Mola & Papeschi by Calvert (1952) as a subspecies of R. cornigera based 1994: 185-187; Mola 1995: 47, 51-54; Muzón & von El- on the different thoracic color pattern which he lenrieder 1998: 23; von Ellenrieder 1999: 151-155 (de- (1905) and Ris (1918) previously mentioned. Calvert scription larva); von Ellenrieder 2000: 21, 24, 28; Martins Costa et al. 2000. (1952) applied the name to the most southern popu- [Aeshna cornigera. – Hagen 1875: 39 (at least in part: Brazil lations ranging from Ecuador to Argentina, whereas R. and Uruguay); Selys 1883: 21; Ris 1904: 23; Calvert c. cornigera corresponded to more northerly popula- 1901-1908: 182-183 (at least in part: Uruguay, Paraguay, tions. However, Calvert (1952; 1956) included both Bolivia and Brazil); Ris 1913: 81; Ris 1918: 157-158; subspecies collected from the same localities and, in Navás 1916: 17; Ris 1918: 157-158 (in part: Brazil, Bo- some cases, the same date for several localities in Bo- livia and Argentina); Navás 1922: 358; Fraser 1947: 433; Schmidt 1952: 238; Abenante & Philippi 1982: 151; Ro- livia, Peru, and Ecuador (figs. 453-454). Since sub- drigues Capítulo et al. 1991: 62; Carvalho & Nessimian species are geographic races that occupy different 1998: 8; Martins Costa 1999: 4. Misidentification] ranges of distribution, the wide area of sympatry of R. [Aeshna cornigera cornigera. – Rodrigues Capítulo 1992: 57. cornigera and R. planaltica indicates that they cannot Misidentification] be considered as such. They could belong to the same polymorphic species or to two different species. Since Diagnosis. – Rhionaeschna planaltica differs from R. no intermediates have been found between the two cornigera by the mes- and metepimeral green stripes morphs and the amount of differences between them with deep rounded indentations (figs. 103-105) vs. mirrors that between R. cornigera and R. nubigena, or linear or slightly undulated in R. cornigera (figs. 92- R. planaltica and R. haarupi, I consider them to be dif- 94), and the linear anterior contour of the ventral tu- ferent species. bercle of first abdominal segment (figs. 264-265) vs. Biology. – The final larval instar was described by convex in R. cornigera (figs. 260-262). Pale areas on De Marmels (1992a) based on one reared specimen the first abdominal sternum in the male (figs. 219- from Tamacuari, Venezuela, and by von Ellenrieder 220, 225) and pale spots of abdomen (figs. 143, 148) (1999a) based on reared specimens from Salta, Ar- are wider than in R. cornigera. gentina. Some differences between the Venezuelan Description. – Clypeus and frons light blue to and Brazilian and Argentine larvae were observed (De bluish yellow, grayish light blue spots lateral to yellow Marmels 1992a; von Ellenrieder 1999), but the area surrounding T-spot stem; T-stem approximately adults show no differences (De Marmels, pers. com.). parallel-sided; posterior 50% of vertex black (fig. 30a); A common species. The larvae occur in mountain frontoclypeal groove bordered by a narrow dark creeks in Venezuela and in small ponds and tempo- brown or black stripe (figs. 30b-c). Pale mesanepister- rary pools in Argentina within forest areas. nal stripes complete and about as wide as 50% of Distribution [34°S-6°N, 40°-79°W, 0-3700 m] mesanepisternum; mesepimeral and metepimeral (fig. 453). – Argentina: Buenos Aires (MLPA*), Córdo- stripes bright green, with deep rounded indentations ba (MLPA; Navás 1927), Tucumán (IMLA; DRPC; on anterior margin (metepimeral) or anterior and pos- Calvert 1956; Fraser 1947; Navás 1922; Ris 1913; terior margins (mesepimeral) (figs. 103-105). Ab- 1918; Schmidt 1952), Catamarca (Fraser 1947; Ris domen dark reddish brown to black with bright green 1918; Schmidt 1952), Salta (IMLA; MLPA*; QCAZ; or yellowish green and light blue spots; male PL spots Calvert 1956; von Ellenrieder 1999), Jujuy (DRPC*), large (figs. 148b). Ventral tubercle of abdominal seg- and Misiones (MACN; MLPA; USNM; Jurzitza 1981; ment I with anterior margin linear in lateral view, Mola & Papeschi 1994; Mola 1995) – Uruguay: San about as high as 50% of its length (fig. 264-265); José (Calvert 1952; 1956), Rocha (Calvert 1952; spines of anterior lamina well developed (fig. 345); tip 1956) – Paraguay: Paraguarí (USNM; Calvert 1952- of hamular anterior process rounded in posterior view 56), Guaira (UMMZ), and Central (DRPC) – Brazil: Rio (fig. 307); auricles with two-three teeth (fig. 225). Grande do Sul (MNRJ; NMW; Calvert 1956; Ris 1918), Male cercus maximum width along medial 30%, tip Santa Catarina (MNRJ; RWGC; Calvert 1952; 1956), directed externally (fig. 386a); dorso-distal crest Paraná (MNRJ), São Paulo (MNRJ; RWGC; UMMZ; smoothly curved, higher than base of cercus in lateral USNM; Calvert 1956; Martins Costa et al. 2000; Ris view, and developed along distal 25% of cercus (fig. 1918), Rio de Janeiro (MNRJ*; Calvert 1956), Mato 386b). Female cercus longer than abdominal segments Grosso do sul (MNRJ), Minas Gerais (MNRJ), Espirito IX-X and shorter than VIII-X, pointed tip, and maxi- Santo (MNRJ), and Mato Grosso (MNRJ; Calvert 1956) mum width at medial 30% (fig. 386c-e). Dimensions: – Bolivia: Cochabamba: Chapare (UMMZ; Calvert head width: 8.2-9; HW length: 36.5-44; HW width: 11- 1956) and La Paz: Larecaja (MLPA) and Nor Yungas

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(Ris 1918; Calvert 1956) – Peru: Apurimac (MLPA), 36-44; HW width: 10.6-12.5; HW pterostigma length: Junín (Calvert 1952; 1956), Lima (Calvert 1956), 2.2-2.8; cerci length: ( 4.9-5.9-, & 7.28-8.26; female Pasco (Calvert 1952), Huanuco (DRPC; UMMZ), An- cerci maximum width: 1.15-1.33; total length: 59-62. cash (Calvert 1952; 1956), Cajamarca (Calvert 1952; Biology. – The larva was described by Needham & 1956), Andes (Calvert 1956), and Amazonas (Calvert Westfall (1955) based on material from the USA, and 1952-56) – Ecuador: Azuay (Calvert 1952), Morona by Calvert (1956) based on material from Costa Rica, Santiago (UMMZ), Chimborazo (Calvert 1952), Boli- Guyana and Cuba. Since Calvert did not breed the var (Calvert 1956), Pastaza (UMMZ; USNM; Calvert larvae, it is likely that some of his specimens belong to 1952-56), and Pichincha (DRPC) – Venezuela: Ama- species other than R. psilus (i.e. material from Mt. Ro- zonas (USNM; De Marmels 1989b; 1992a), Atabapo raima could be R. nubigena or R. planaltica, and mate- (De Marmels 1989a), and Bolivar (De Marmels rial from Costa Rica could be R. cornigera). The spec- 1989a) – Colombia: without locality (NMW*). imens from Cuba are probably R. psilus, since this is the only species of the group recorded from that coun- Rhionaeschna psilus (Calvert) comb. n. try. The species was collected at small rain ponds with (figs. 31-he, 106-th, 149-ab, 187-te, 226-ge, 268-tg, 308-ha, 348- grassy margins in open areas, ponds with abundant as, 388-ce, 455-Mp) aquatic vegetation, grassy marshes with Salvinia and Aeshna psilus Calvert, 1947: 1-7 (description (&: holotype Lemna and large aroid and trees at edge, reservoirs, a ( COSTA RICA: stagnant pool, banks of Río Reventazón, wooded pond in sun, drainage flowing ditches with Cachí, 10-III-1910, P.P. Calvert leg. (ANSP, 9273)); Need- grassy banks and mud bottom, flying over cornfields, ham & Westfall 1955: 291, 293-294, 296-297, 312-313; small rocky and sandy streams in stream bed (DRPC). Calvert 1956: 11, 50-55 194-196; De Marmels 1981: 11; Distribution [22°S-32°N, 61-110°W, 0-1500 m] 1988: 101; González Soriano 1993: 296; González Sori- ano & Novelo Gutiérrez 1996: 164; Donnelly et al. 1998: (fig. 455). – Argentina: Salta (MLPA; Donnelly et al. 101; Muzón & von Ellenrieder 1998: 23. 1998) – Peru: Lima (Calvert 1947; 1956), and Junín [Aeshna cornigera. – Calvert 1895: 507-508, figs. 24, 31-32 (Calvert 1947; 1956) – Ecuador: El Oro (UMMZ), (in part); Calvert 1905: 179,182 (in part); Calvert 1907: Tungurahua (Calvert 1947; 1956), Napo (Calvert 400 (in part); Ris 1918: 157-158 (in part); Klots 1932: 1947; 1956), and Manabi (UMMZ) – Venezuela: 18; García-Díaz 1938: 55. Misidentification] Táchira (De Marmels 1988) and Miranda (De Marmels 1981) – Panama: Chiriquí (Calvert 1947; Diagnosis. – Rhionaeschna psilus is most closely re- 1956) – Costa Rica: Puntarenas (INBC; USNM), San lated to R. manni, their separation is given under R. José (DRPC; MN; INBC; Calvert 1947), Cartago manni. (USNM; Calvert 1947; 1956), and Alajuela (USNM) – Description. – Clypeus and frons light blue, green Nicaragua: Matagalpa (Beckemeyer 2001) – Hon- spots lateral to yellow area surrounding T-spot stem; duras: without locality (Förster 2001) – Guatemala: T-stem approximately parallel-sided or slightly con- Guatemala (Calvert 1947; 1956) and Alta Verapaz verging anteriorly; posterior 60% of vertex black (fig. (USNM) – Belize: Cayo (Boomsma & Dunkle 1996) – 31a); no dark stripe on frontoclypeal groove (figs. Puerto Rico: Río Grande (RWGC; Calvert 1956) – Do- 31b-c). Pale mesanepisternal stripes complete and as minican Republic: Barahoa (USNM) and La Vega wide as 25-33% of mesanepisternum; mesepimeral (RWGC; USNM) – Dominica: Dominica (Donnelly and metepimeral stripes bright green, with slightly sin- 1970) – Haiti: de l’Ouest (DRPC) – Jamaica: Surrey uous margins (fig. 106). Abdomen dark reddish (Calvert 1956) – Cuba: Pinar del Río (Peters 1988), brown to black with bright green or yellowish green Oriente (Alayo 1968, Peters 1988) and Camagüey and light blue spots; male PL spots large (fig. 149b). (Calvert 1956) – Mexico: Chiapas (DRPC; RWGC; Ventral tubercle of abdominal segment I with anterior USNM), Tabasco (UMMZ), Veracruz (RWGC; USNM; Ris margin linear or slightly convex in lateral view, about 1918; Calvert 1905; 1947; 1956), Quintana Roo as high as 50% of its length (fig. 268); spines of ante- (LACM; USNM; Calvert 1905; 1947; 1956), Morelos rior lamina vestigial (fig. 348); tip of hamular anterior (Calvert 1905; 1947; 1956; Ris 1918), Aguascalientes process rounded in posterior view (fig. 308); auricles (Calvert 1905; 1947; 1956), Nayarit (DRPC; Calvert with three teeth (fig. 226). Male cercus with inner and 1905; 1956); Jalisco (Calvert 1905; 1947; 1956), Yu- outer margins approximately parallel to each other catán (LACM; Calvert 1947; 1956), Hidalgo (DRPC), along medial 30%, where maximum width is ob- San Luis Potosí (DRPC), and Sinaloa (DRPC) – U.S.A.: served, tip directed posteriorly (fig. 388a); dorso-distal Texas (RWGC) and Arizona (Behrstock 1998). crest angled, lower than base of cercus in lateral view, and developed along distal 30% of cercus (fig. 388b). Rhionaeschna vazquezae (González) comb. n. Female cercus longer than abdominal segments VIII-X, (figs. 32-he, 107-109-th, 150-ab, 188-te, 227-ge, 269-tg, 309-ha, pointed tip, and maximum width at medial 30% (fig. 349-as, 389-ce, 456-Mp) 388c). Dimensions: head width: 7.8-8; HW length:

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Aeshna vazquezae González, 1986: 47-51, figs. 1-4 (descrip- 7. R. punctata group tion (: holotype ( MEXICO: Acahuizotla, 17.5 km S Chilpancingo (Guerrero state), 15-XI-1983, R. Mendoza leg. (UNAM) [examined: ( paratype]); 1993: 296; Two rows of cells between RP1 and RP2 in HW be- González & Novelo 1996: 164. ginning under pterostigma or proximal to it (fig. 403); male cercus widening gradually to distal end (maxi- Diagnosis. – Rhionaeschna vazquezae is unique in mum width at distal 0.12-0.16) with high dorso-distal the group by the very wide mes- and metepimeral crest, lateral carina convex, ventrally bent tip, and sub- stripes (wider than half of their sclerites, figs. 107- basal tooth absent (figs. 390-395); denticles of genital 109). It shares with R. manni and R. psilus the pres- lobe very small, densely spread over the ventral and ence of an angulate dorso-distal carina on the male lateral surfaces of the lobe (figs. 270-275); hamular cercus being lower than the base of cercus (figs. 384, anterior process carinated. Seven species: R. biliosa, R. 388-389) but it is easily distinguished from those condor, R. decessus, R. demarmelsi, R. eduardoi, R. joan- species by the strongly developed anterior lamina nisi, R. punctata. spine (fig. 349), this structure being rudimentary in R. manni and R. psilus (figs. 347-348). Rhionaeschna biliosa (Kennedy) comb. n. Description. – Clypeus and frons greenish light (figs. 33-he, 110-111-th, 151-ab, 189-te, 228-ge, 270-tg, 310-ha, blue, green spots lateral to yellow area surrounding T- 350-as, 392-ce, 457-Mp) spot stem; T-stem gradually narrowing anteriorly; ver- Aeshna biliosa Kennedy, 1938: 573-576 (description (: tex black except for two yellow antero-lateral spots holotype ( ECUADOR: La Ventana, near Baños, 2000 m, (fig. 32a); frontoclypeal groove bordered by a narrow Río Pastaza (Tungurahua prov.), V-1936, W. Clarke- black stripe (figs. 32b-c). Pale stripes complete and McIntyre leg. (UMMZ) [examined: holotype (, allotype bright green; mesanepisternal about as wide as 60% of &, paratype (]); Kennedy 1939: 344-348 (description &); Calvert 1952: 254; Soukup 1954: 15; Calvert 1956: mesanepisternum; mesepimeral and metepimeral 12, 72-76; Rácenis 1959: 493. wider than 50% of their sclerites, with linear anterior margins and posterior margins slightly undulated Diagnosis. – Rhionaeschna biliosa is easily distin- (mesepimeral) or deeply excavated or divided (metepi- guished from its congeners by the bright yellow frons meral) (figs. 107-109). Abdomen black with bright without dark stripe on frontoclypeal groove and the green and blue spots; male PL spots large (figs. 150b). narrow transverse arms of T-spot. It shares with some Ventral tubercle of abdominal segment I with anterior specimens of R. joannisi a similar thoracic pattern of margin linear in lateral view, about as high as 50% of pale spots but these are yellow in R. biliosa and not its length (fig. 269); spines of anterior lamina well de- green as in R. joannisi. Males share similar morpholo- veloped (fig. 349); tip of hamular anterior process car- gies of cercus, hamulus, and anterior lamina spine inated in posterior view (fig. 309a); auricles with three with R. condor and R. joannisi, and females the mu- teeth (fig. 227). Male cercus maximum width at distal cronate condition of the cercus with R. eduardoi. 30%, tip between directed externally (fig. 389a); dor- Description. – Clypeus orange, frons bright yellow, so-distal crest angled, lower than base of cercus in lat- no spots lateral to T-spot; T-stem parallel-sided, nar- eral view, and developed along distal 30% of cercus rower than vertex, transverse arms represented by a fine (fig. 389b). Female unknown. Dimensions: head black line; vertex completely black or with two antero- width: 8.6-9.5; HW length: 45.5-47.7; HW width: lateral yellow spots (fig. 33a); frontoclypeal groove not 13.7-15; HW pterostigma length: 2.37-2.8; cerci bordered by dark stripe (figs. 33b-c). Pale pterotho- length: ( 5.62-6.5; total length: 68-72. racic stripes yellow, divided in spots: mesanepisternal Biology. – Rhionaeschna vazquezae was recently col- into a basal and a distal spot; mesepimeral into four lected for the first time after its original description by spots and metepimeral into two to four spots (figs. Dr. D. R. Paulson, who kindly provided the following 110-111). Abdomen dark reddish brown to black with notes: ‘The specimens were collected at a stream with yellow to pale brown spots. Outer margin of ventral boulders and gravel riffles, muddy pools (one of them terga VII sinuous (figs. 189a-b). Ventral tubercle of ab- long), and steep gradient, in low forest with sun pene- dominal segment I rounded in lateral view; genital lobe trating. Males flew up and down along pool, sometimes lower than 30% of its length (fig. 270); spines of ante- over center, mostly right at shore, at water surface and in rior lamina longer than twice its basal width and acute and out of overhanging branches and roots, with a very pointed (fig. 350); ventral portion of hamuli higher slow and methodical searching, mostly after the sun had than hamular fold (fig. 310a), tip of hamular anterior disappeared’. Rare in collections. Larva unknown. process rounded in posterior view and pointed in ven- Distribution [17°-21°N, 99°-104°W, 950 m] (fig. tral view (figs. 310a-b); auricles with two teeth (fig. 456). – Mexico: Guerrero (UNAM*; González 1986; 228). Dorso-distal crest of male cercus pointed, tip 1993; González & Novelo 1996) and Nayarit (RWGC; pointed (fig. 392b). Maximum width of female cercus DRPC*). at medial 30%; tip mucronate (fig. 392c). Dimensions:

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head width: 9.9-10.4; HW length: 51.1-52; HW width: merged bush and floating grass stems (De Marmels 14.7-17; HW pterostigma length: 2.8-3.3; cerci length: 2001a). Rare in collections. ( 7-7.3, & 7.4; female cerci maximum width: 1.6; to- Distribution [7°N, 72°W, 2000 m] (fig. 457). – tal length: 73-79. Venezuela: Táchira (RWGC*). Biology. – All specimens have been collected in Andean localities. Larva unknown. Rare in collec- Rhionaeschna decessus (Calvert) comb. n. tions. (fig. 459-Mp) Distribution [6-1°S, 77-78°W, 1800-3500 m] (fig. 457). – Peru: Amazonas (Kennedy 1938; Calvert Aeshna decessus Calvert, 1953: 205-207 (description (: 1956) – Ecuador: Tungurahua (MNRJ; RWGC*; Calvert holotype ( Brazil: Südhäng Itatiaia-Gebirg., 22º25’22S 44º37’4W 700 m, X-1931, Zikan leg. (Rio de Janeiro) 1956; Kennedy 1938; 1939) and Manabi (MLPA; [type probably lost]); Calvert 1956: 12, 88-90. UMMZ*). Diagnosis. – According to its original description, Rhionaeschna condor (De Marmels) comb. n. this species is closely related to R. punctata and R. ed- (figs. 34-he, 112-th, 152-ab, 190-te, 229-ge, 271-tg, 313-ha, 351- uardoi, with which it shares in the male similarly as, 390-ce, 457-Mp) shaped cercus. The orange face, the absence of spines Aeshna condor De Marmels, 2001a: 129-134 (description ( in the anterior lamina, and the pale abdominal PL and larva: holotype ( VENEZUELA: San Vicente de La Re- spots present on segments IV-IX (PL spots absent in VI- vancha, road to Las Copas, Tamá National Park, VII in R. eduardoi and R. punctata, figs. 155-156b, d) 7°29’59N 72°20’35W 2000 m (Táchira state), 06-XI- would distinguish it from these species. 1999, J. De Marmels leg. (MIZA) [paratype ( (RWGC)]) Description. – Clypeus orange, postfrons orange, antefrons orange with purplish spots lateral to pale Diagnosis. – Rhionaeschna condor is distinguished area surrounding T-spot stem; T-stem parallel-sided, from the other species of the group by the wide T-spot narrower than vertex, transverse arms wider than a of frons (fig. 34a), wide green thoracic stripes (fig. line; dark stripe on frontoclypeal groove; vertex orange 112), and recumbent tip of hamular anterior process with lateral and posterior margins black. Pale (fig. 313a). mesanepisternal at basal 80% of mesanepisternum Description. – Clypeus and frons light blue, dark stripes greenish, slightly diverging from each other. blue-grayish spots lateral to yellow area surrounding Abdomen reddish brown with pale green and light T-spot stem; T-stem wider than vertex, with convex blue spots. Genital lobe higher than tubercle; spines of sides, transverse arms wider than a line; vertex black anterior lamina absent; auricles with two teeth. Dorso- (fig. 34a); narrow dark stripe on frontoclypeal groove, distal crest of male cercus smoothly curved, tip point- not widened to fronto-ocular groove (figs. 34b-c). Pale ed. Female unknown. Dimensions: head width: pterothoracic stripes yellowish green, complete and 11.44; HW length: 51; HW width: 16; cerci length: ( wider than 50% of their sclerites, mesepimeral stripe 5.73; total length: 78. with a pointed branch extending on posterior 50% of Biology. – Rhionaeschna decessus is only known metepisternum (fig. 112). Abdomen black with yel- from the male holotype collected at Itatiaia. lowish green and blue spots. Outer margin of ventral Remarks. – The diagnosis and comments on this terga VII sinuous (figs. 190a-b). Ventral tubercle of ab- species are based solely on the original description. dominal segment I trapezoidal in lateral view, genital Calvert (1953) mentioned that the holotype was first lobe lower than 30% of its length (fig. 271); spine of identified as R. punctata and that the genital fossa was anterior lamina longer than twice its basal width and coated with a hardened substance which was subse- acute pointed (fig. 351); ventral portion of hamuli quently removed. Since the spines of the anterior lam- higher than hamular fold (fig. 313a), tip of hamular ina in R. punctata and R. eduardoi are slender, it is pos- anterior process recumbent in posterior view and sible that such fragile structures, if originally present, pointed in ventral view (figs. 313a-b); auricles with could have been accidentally removed during clean- three teeth (fig. 229). Dorso-distal crest of male cercus ing. If so, this specimen may represent a pale specimen angled, tip pointed (fig. 390b). Female unknown. Di- of R. punctata or R. eduardoi. mensions: head width: 10; HW length: 49.2-51.3; HW The type specimen is apparently lost. It was origi- width: 14.12; HW pterostigma length: 2.2-2.6; cerci nally deposited in the collection of Dr. E. Schmidt at length: ( 5.3-6; total length: 69-71.3. the Zoologisches Forschungsinstitut und Museum Biology. – Males and exuviae of this species were Alexander König in Bonn. The collection of Schmidt collected in a small pond fed by a minute stream in the was later sold to Dr. S. Asahina, who in turn donated it Mount Tamá range (De Marmels 2001a). Males were to the National Museum of Natural History in Japan. observed patrolling the edge of the pond and females None of these institutions hosts it today (pers. coms. of attempting to oviposit into a branch of a partly sub- Dr. B. Misof, Dr. S. Asahina and Dr. M. Tomokuni).

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Distribution [22°S, 44°W, 700 m] (fig. 459). – er than vertex, approximately parallel sided, transverse Brazil: Rio de Janeiro (Calvert 1953). arms wider than a line; vertex black with a green spot at anterior margin on each side (as in fig. 35a); brown Rhionaeschna demarmelsi von Ellenrieder sp. n. stripe on frontoclypeal groove, not widened to fronto- (figs. 35-he, 113-th, 153-ab, 191-te, 230-ge, 272-tg, 311-ha, 353- ocular groove (as in figs. 35b-c). Thorax. Pale 354-as, 393-ce, 403,411-wi, 458-Mp) pterothoracic stripes yellowish green, divided or with [Aeshna joannisi. – De Marmels 2001a (description (, &, indentations; mesanepisternal divided into a basal larva). Misidentification] short stripe (as long as 25% of mesanepisternum) and a distal spot, mesepimeral and metepimeral each with Type Material: 1( holotype VENEZUELA: San Vi- a postero-basal and an antero-distal indentation; pos- cente de La Revancha, Las Copas (Táchira State) lagu- tero-basal indentation of mesepimeral stripe and an- na, 2000 m, 16-v-1999, De Marmels leg. (MIZA) – 1& tero-distal of metepimeral not reaching half of stripe allotype, 3( paratypes same except Tamá National width (as in fig. 113), yellowish green spot at each side Park, 7°29’59N 72°20’35W 2000 m, 16-v-1999 (al- of antealar crest in antealar sinus, and on each subalar, lotype MIZA &; paratypes MIZA &; paratype MLPA); 3( costal, and radioanal plate, meso-and metascutum; same data except 7/11-xi-1999; 4 ( same data except light blue on metascutum. Legs black except for 7/8-vii-2000 (MIZA&; RWGC). trochanters, coxae and basal 75% of femora reddish- brown, and basal 50% of inner surface of femur I yel- Etymology: Rhionaeschna demarmelsi (noun in the lowish-green. Wings hyaline, tinged with yellow at genitive case), is named in honor of Dr. Jürg De base (basally to first antenodal) and at nodus [or whole Marmels, whose work on the systematics of the wings]; membranule dark except basal 30% pale (fig. Odonata of Venezuela serves as an inspiration to us all. 403). Wing venation: Antenodals FW: 18; HW: 13 Diagnosis. – The short spine of anterior lamina right, 14 left; posnodals FW: 11 right, 10 left; HW: 14; (figs. 353-354) and the combination of smoothly triangle cells FW: 4; HW: 4; supratriangle crossveins FW: curved dorso-distal crest and blunt tip of male cercus 3; HW: 2; subtriangle cells FW: 2; HW: 2; cu-A space (figs. 393b) are unique for R. demarmelsi within the R. crossveins FW: 3; HW: 3; bridge crossveins FW: 4; HW: punctata group, and separate it from all its congeners. 4 right, 3 left; cell rows between IRP2 FW: 3; HW: 3; The female cercus with maximum width at distal 30% cells anal triangle: 3; cells anal loop: 11; rows of cells (fig. 393c) is also characteristic and allows identifying anal loop: 3. Abdomen dark reddish brown to black the female from all the others known for the group (R. with yellowish green and light blue spots as follows: biliosa, R. joannisi, R. eduardoi, and R. punctata, figs. spot at base of genital lobe, lateral stripe on anterior 391-392, 394-395). The overall color pattern of margin of transverse carina and dorsal spot posterior Rhionaeschna demarmelsi is very similar to that of R. to transverse carina in segment II, AD, AL spots of seg- joannisi, but it differs from it in several other charac- ment III, and PD spots of segments III-IV light blue, re- ters: postero-basal indentation of mesepimeral stripe maining pale spots yellowish green (as in figs. 153a-b). and antero-distal of metepimeral stripe not reaching Outer margin of ventral terga VII linear (as in fig. half of stripe width (fig. 113, surpassing half of stripe 191a). Ventral tubercle of abdominal segment I width or dividing the stripes into spots in R. joannisi, rounded in lateral view, genital lobe lower than 30% figs. 115-117); auricles bearing two large teeth (fig. of its length (as in fig. 272); spines of anterior lamina 230, three in R. joannisi, fig. 231); ventral portion of shorter than 1.5 times of its basal width and blunt [or hamuli shorter than hamular fold (higher in R. joan- pointed] (as in figs. 353-354); ventral portion of nisi); shorter female cerci (fig. 393c, 5 mm versus 6.5 hamuli shorter than hamular fold (as in fig. 311a), tip mm in R. joannisi, fig. 391c) and with maximum of hamular anterior process pointed in posterior and width at distal 30% (at medial 30% in R. joannisi); ventral views (as in figs. 311a-b); right auricle with outer margin of ventral terga VII linear (fig. 191, sinu- two teeth (as in fig. 230), left auricle with two large ous in R. joannisi, fig. 192); abdominal spots AL, AD, teeth and a third one smaller and more external. Vesi- ML, and MD present in segments VII-VIII of male (figs. ca spermalis with basal folds in lateral and dorsal lobes 153a-b, absent in R. joannisi, figs. 154a-b) and AL, MD, as typical for the genus (illustrated by De Marmels and PD present in segment VIII of female (absent in R. 2001a). Dorso-distal crest of male cercus smoothly joannisi, fig. 154c-e). convex, tip blunt (as in fig. 393b). Description. – Holotype (variation of paratypes in Allotype: As holotype except frons pale brown; and brackets): Head. Labium brown. Labrum yellow with thorax and abdomen lacking light blue, all pale spots ventral margin dark brown and dorsal margin black. being yellowish green to turquoise or olivaceus (PD Clypeus light blue, with lateral margins yellow. Frons spots). Wing venation: Antenodals FW: 18; HW: 12 yellow [to light blue], dark grayish-blue spots lateral to right, 11 left; posnodals FW: 12 right, 11 left; HW: 15 yellow area surrounding T-spot stem; T-stem narrow- right, 14 left; triangle cells FW: 5 right, 4 left; HW: 4;

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supratriangle crossveins FW: 3; HW: 2 right, 3 left; sub- male cercus smoothly curved, anterior side with denti- triangle cells FW: 2; HW: 2; cu-A space crossveins FW: 4; cles, tip pointed (fig. 394b), diverging from other cer- HW: 3; bridge crossveins FW: 4 right, 5 left; HW: 4 right, cus tip in dorsal view (fig. 394a). Maximum width of 3 left; cell rows at IRP2 fork FW: 3 right, 2 left; HW: 3; female cercus at medial 30%; tip pointed terminating cells anal loop: 12; rows of cells anal loop: 3. Cercus in a minute spine (Machado 1985a). longer than segments IX-X; maximum width at distal Dimensions: head width: 9.6; HW length: 44.1- 30%; tip incised (fig. 387c). Dimensions: head width: 47.9; HW width: 14.07; HW pterostigma length: 3.2- 9.15 [9-9.2]; HW length: 43.5 [40.2-43.5]; HW width: 3.6; cerci length: ( 5.7-6.1, & 6.3-6.7; maximum 14.8 [13.5-14.8]; HW pterostigma length: 2.5 [2-3]; width of female cerci: 1.3; total length: 68-72. cerci length: ( 5.2 [5-5.24], & 5.0; maximum width Diagnosis. – Rhionaeschna eduardoi is most closely of female cerci: 1.2; total length: 67.5 [58.3-67.5]. related to R. punctata, with which it shares the pres- Biology. – De Marmels (2001) reported this ence of complete linear pterothoracic stripes (figs. species as common at a pond in Tamá National Park 114, 118), prominent genital lobe (higher than 30% where it was found in sympatry with R. condor, R. its width, figs. 274, 275), male cercus with a smooth- cornigera, and R. marchali. Males patrolled the pond ly curved crest and acuminate tip (figs. 394-395), and occasionally approaching the shore; females oviposit- abdominal color pattern (figs. 155-156). They differ ed into floating grass stems, and exuviae were found in several characters (second set of character states for clinging to branches of half submerged bushes. Rare R. punctata): black of T-spot extended on antefrons in collections. (fig. 37c) vs. not extended (fig. 38c); vertex black Distribution [7°N, 72°W, 2000-2600 m] (fig. along posterior 30% (fig. 37a) vs. only lateral edges 458). – Venezuela: Táchira (MIZA*; RWGC*; MLPA*; black (fig. 38a); metepimeral stripes wider than De Marmels 2001a). mesepimeral ones (fig. 114) vs. of equal width (fig. 115); ventral tubercle of segment I high and trape- Rhionaeschna eduardoi (Machado) comb. n. zoidal (fig. 274) vs. low and rounded (fig. 275); and (figs. 37-he, 114-th, 155-ab, 193-te, 232, 419-ge, 274-tg, 314-ha, tip of female cercus pointed and bearing a minute 355-as, 394-ce, 459-Mp) spine vs. rounded and incised (fig. 395c). Aeshna eduardoi Machado, 1984: 161 (nomen nudum); Biology. – Rhionaeschna eduardoi seems to be re- Machado 1985a: 45-56 (description (&: holotype ( stricted to iron-rich environments of mountainous re- BRAZIL: Minas Gerais-Brumadinho (S. of Belo Hori- gions of Minas Gerais. The adults have been reported zonte), reserve of Catarina, 20°7’60S 44°13’0W 1200 m, patrolling margins of man-made pools and swamps 21-IV-1983, A. & E. & P. Machado leg. (ABMMC) [ex- fed by streams inside or near the forest (Machado amined: paratype ( (RWGC)]). 1985a). Rare in collections. Larva unknown. [Aeshna punctata. – Martin 1908: 54-55, 83 (in part: ( in Selys Coll.); Santos 1966a: 97-98 (in part: records from Remarks: The species was named in a congress con- Poços de Caldas); Santos 1966c: 65. Misidentification] tribution in 1984, which does not constitute a valid publication (nomen nudum). Therefore, the name Description. – Clypeus light blue, frons bluish yel- Rhionaeschna eduardoi was first made available for this low, with light blue spots lateral to yellow area sur- species in the description published in 1985a, and it rounding T-spot stem; T-stem parallel-sided, narrow- takes authorship from that date. er than vertex, transverse arms wider than a line; vertex Distribution [20-21°S, 44-46°W, 1000-1500 m] yellow with lateral and posterior 30% black (fig. 35a); (fig. 459). – Brazil: Minas Gerais (Machado 1985; dark stripe on frontoclypeal groove, strongly widened RWGC*). to fronto-ocular groove (figs. 35b-c). Pale pterotho- racic stripes yellow, mesanepisternal at basal 80% of Rhionaeschna joannisi (Martin) comb. n. mesanepisternum; mesepimeral and metepimeral lin- (figs. 36-he, 115-117-th, 154-ab, 192-te, 231-ge, 273-tg, 312-ha, ear and complete, metepimeral wider than 352-as, 391-ce, 458-Mp) mesepimeral (fig. 114). Abdomen reddish brown to Aeshna joannisi Martin, 1897: 502 (description &: holotype black with yellowish green and light blue spots. Outer & BOLIVIA: San Antonio (MNHN) [examined: & holo- margin of ventral terga VII linear (fig. 193a). Ventral type]); Martin 1908: 56, 84; Calvert 1956: 12, 76-80 tubercle of abdominal segment I trapezoidal in lateral (description (). view, as high as 60% of its length, genital lobe higher than 30% of its width (fig. 274); spine of anterior lam- Diagnosis. – Provided under R. demarmelsi. ina longer than twice its basal width, narrowly trian- Description. – Clypeus light blue, frons greenish gular and pointed (fig. 355); ventral portion of hamuli yellow, greenish blue spots lateral to yellow area sur- shorter than hamular fold (fig. 314a), tip of hamular rounding T-spot stem; T-stem narrower than vertex, anterior process blunt in ventral view (fig. 314b); auri- approximately parallel sided, transverse arms wider cles with two teeth (fig. 232). Dorso-distal crest of than a line; vertex black with anterior margin green

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(fig. 36a); narrow dark stripe on frontoclypeal groove, dor: Napo (KJTC*), Pastaza (Calvert 1956), and Pich- not widened to fronto-ocular groove (figs. 36b-c). Pale incha (DRPC*) – Colombia: Cauca (Calvert 1956). pterothoracic stripes yellowish green, divided in spots or with deep indentations; mesanepisternal divided Rhionaeschna punctata (Martin) comb. n. into a basal and a distal spot, mesepimeral and (figs. 38-he, 118-th, 156-ab, 194-te, 233-ge, 275-tg, 315-ha, 356- metepimeral each with a postero-basal and an antero- 357-as, 395-ce, 459-Mp) distal indentation, which surpasses half of stripe width Aeshna punctata Martin, 1908: 54-55, 83 (in part; descrip- or divide the stripe into spots (figs. 115-117). Ab- tion (: lectotype ( BRAZIL: Espirito Santo (MNHN)); domen dark reddish brown to black with yellowish Martin 1911: 12; Calvert 1956: 12, 80-88 (in part, green and light blue spots. Outer margin of ventral records from Brazil; description &); Santos 1966a: 97- terga VII sinuous (figs. 192a-b). Ventral tubercle of ab- 100 (in part; description larva); Machado 1985b: 327- 332 (redescription (); Martins Costa et al. 2000: 13. dominal segment I rounded in lateral view, genital Aeschna depravata Hagen, 1861: 314 (nomen nudum). lobe lower than 30% of its length (fig. 273); spine of Aeschna lobata Hagen, 1861: 314 (nomen nudum). anterior lamina longer than twice its basal width and acute pointed (fig. 352); ventral portion of hamuli Diagnosis. – Differentiation of R. punctata from its higher than hamular fold (fig. 312a), tip of hamular closest allies is given under R. eduardoi and R. decessus. anterior process rounded in posterior view and point- Description. – Clypeus light yellow, frons yellow, ed in ventral view (figs. 312a-b), auricles with three light blue spots lateral yellow area surrounding T-spot large teeth, sometimes with a fourth smaller tooth (fig. stem; T-stem parallel-sided, narrower than vertex, 231). Dorso-distal crest of male cercus angled, tip transverse arms wider than a line; vertex yellow with pointed (fig. 391b). Maximum width of female cercus lateral edges black (fig. 38a); dark stripe on fronto- at medial 30%; tip incised (fig. 391c). Dimensions clypeal groove, strongly widened to fronto-ocular (values of holotype in square brackets): head width: groove (figs. 38b-c). Pale pterothoracic stripes yellow, 9.5-9.8 [9.6]; HW length: 46-50 [47]; HW width: 14- mesanepisternal at basal 80% of mesanepisternum; 15.7 [14.5]; HW pterostigma length: 2.2-3.2 [3.2]; mesepimeral and metepimeral linear and complete, of cerci length: ( 5.7, & 6.5 [6.5]; maximum width of equal width (fig. 118). Abdomen dark reddish brown female cerci: 1.5 [1.5]; total length: 68-72.5 [68]. to black with yellowish green and light blue spots. Biology. – Adults have been collected in areas of dry Outer margin of ventral terga VII sinuous (figs. 194a- climate in Bolivia (Dr. K. Tennessen pers. comm.), b). Ventral tubercle of abdominal segment I rounded and in Ecuador in a very wet climate, flying beats up in lateral view, genital lobe higher than half of its and down on the open part of a one-meter wide road- length (fig. 275); spine of anterior lamina longer than side ditch with slight current and pools, with water twice its basal width and needle-shaped to narrowly from hillside stream adjacent to forest (Dr. D. R. Paul- triangular (figs. 356-357); ventral portion of hamuli son pers. comm.). Rare in collections. Larva unknown. higher than hamular fold (fig. 315a), tip of hamular Remarks. – In his description of R. joannisi, Martin anterior process blunt in ventral view (fig. 315b); auri- (1897) stated that it was based on a single female from cles with two teeth (fig. 233). Dorso-distal crest of Bolivia. There are two females from Bolivia labeled as male cercus smoothly curved, anterior side with denti- R. joannisi by Martin at the collection of the MNHN, cles, tip pointed (fig. 395b), converging or parallel to none of which was labeled as type by Martin; one other cercus tip in dorsal view (395a). Maximum from ‘Bolivie’ with an indication that cerci are broken width of female cercus at medial 30%; tip rounded (‘app. brisé’) and the other from ‘Bolivie, S. Antonio’. and incised (395c). Dimensions: head width: 9.45- Following Dr. E. Schmidt’s comments on these speci- 10.5; HW length: 45-50; HW width: 13.1-15.3; HW mens Calvert (1956) considered the first one to be the pterostigma length: 2.9-3.25; cerci length: ( 5.7-6.1, type because Martin stated only ‘Bolivie’ for the type & 5.5; total length: 68-82. locality. However, since a description of the cerci was Biology. – The species has been reported as abun- included (Martin 1897) and the venation characters as dant from Brejo da Lapa, Itatiaia, Brazil, where adults given by Martin in the original description agree well and larvae were collected at ponds with abundant with the specimen of San Antonio, I consider this macrophytes (Santos 1966a). The larva was described specimen the holotype. Fifty-eight localities called San by Santos (1966a) based on reared material from this Antonio were found in Bolivia, and unfortunately the locality. type labels do not include any further information Remarks. – Machado (1985b) examined the type that would allow for knowing which one corresponds series of Rhionaeschna punctata deposited in the to the type locality. MNHN and ISNB collections and concluded that it in- Distribution [10ºS? 1°S-2°N, 77-78°W, 1100- cludes three different species: R. punctata, R. eduardoi 2200 m] (fig. 458). – Bolivia: without locality and R. jalapensis. He designated a lectotype and pro- (NHMP*; Martin 1897; 1908; Calvert 1956) – Ecua- vided a redescription of the male.

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Distribution [31-19°S, 43-52°W, 0-2200 m] (fig. aeschna (and Hemianax-Anax, see fig. 437), and ‘A.’ 459). – Brazil: Rio Grande do Sul (MNRJ), Santa isoceles constitutes the sister group of Andaeshna. A Catarina (Calvert 1956), São Paulo (MNRJ*; USNM*; small group of African species of ‘Aeshna’ consisting of Santos 1966a; Martins Costa et al. 2000), Rio de ‘A.’ rileyi and ‘A.’ subpupillata from South Africa, and Janeiro (MNRJ*; Calvert 1956; Hagen 1861; Santos ‘A.’ moori from Central Africa (‘A.’ moori was not ex- 1966a), Minas Gerais (MNRJ; Santos 1966a), and Es- amined in the present analysis but belongs to the same pirito Santo (Martin 1908). group according to its description) would be the sister group of Rhionaeschna. They share with Rhionaeschna the presence of a linear dorso-distal crest of male cer- PHYLOGENETIC ANALYSIS cus in dorsal view (as in fig. 427b) but this is a homo- The first analysis generated eight equally most par- plastic character present in several other Aeshnidae, simonious trees (113 steps, CI 0.49, RI 0.88). Succes- and the support for this grouping is consequently sive weighting resulted in 332 trees (CI 0.72, RI 0.95). weak. The African group (‘A.’ rileyi, ‘A.’ subpupillata, In the strict consensus cladogram of the 332 trees (fig. ‘A.’ moori) is characterized by the greatly enlarged and 438, CI 0.70, RI 0.94), each node is numbered, and posteriorly projected genital lobe devoid of denticles synapomorphies supporting each node are listed with (fig. 421), and the morphology of the last segment of homoplasies indicated with an asterisk. the vesica spermalis, which bears a unique large The analysis shows the genus Aeshna Fabricius to be medio-longitudinal fold on the ventral lobe (figs. paraphyletic with some of its species more closely re- 436a, b). lated to the Andaeshna-Anaciaeschna-Hemianax-Anax The autapomorphy defining Rhionaeschna is the clade (node 13-15, fig. 438). Paraphyly of Aeshna had presence of a conical tubercle bearing denticles on ab- already been suggested by Peters (1987), who based dominal sternum I (figs. 234-275, 418-419); sternum his analysis on wing characters of the European Aesh- I is planar in Oreaeschna, Aeshna s.str., ‘Aeshna’ species nidae. Part of ‘Aeshna’, corresponding mainly to the of uncertain position (‘A.’ affinis, ‘A.’ mixta, ‘A.’ bre- Holarctic species form a natural group (node 1-2, fig. vistyla, ‘A.’ ellioti, ‘A.’ williamsoniana, ‘A.’ moori, ‘A.’ 438) characterized by the abdominal auricles bearing rileyi and ‘A.’ subpupillata), and Andaeshna (fig. 425). four or five denticles (figs. 428-430). The type species Abdominal sternum I has a transverse ridge in Anaci- of Aeshna, A. grandis, belongs to this group, which aeschna and ‘A.’ isoceles (fig. 421). Castoraeschna has therefore retains the name. Two monophyletic groups also a tubercle in sternum I, but it is cylindrical, much can be distinguished within Aeshna s.str., one of them higher than in Rhionaeschna and lacks denticles (fig. (node 2-3, fig. 438) characterized by narrow two- 424); I consider their presence in Rhionaeschna and celled anal triangles (fig. 405), and the second one Castoraeschna due to convergence. (node 2-8, fig. 438) by the following five autapomor- According to the cladogram, Rhionaeschna species phies: (1) absence of sub-basal tooth or carina in dor- form four major groups as follows, but their relation- sum of Xth segment, (2) male cercus with an apical ships with one another are still unclear: ventral spine on inner margin, (3) cercus inner margin First, Rhionaeschna draco shows numerous autapo- concave distally in dorsal view (figs. 426a, b), (4) dor- morphies, i.e. frons projected dorsally, higher than so-distal crest curved in dorsal view (figs. 426c), and vertex (fig. 1b); pterothorax and abdominal segments (5) apical spines of epiproct large and directed anteri- VI-X without pale markings (figs. 39, 119); hamulus orly (fig. 426d). with a medial digitiform projection on its antero-ven- The remaining ‘Aeshna’ species and Aeshninae gen- tral margin (fig. 276); female cercus with long termi- era form a monophyletic group (node 1-10, fig. 438) nal spine (fig. 358c), but except for the presence of characterized by the marked bending of MA (figs. 396- ventral tubercle I, it does not share other derived char- 404, 406-407) and short parallel fusion of veins de- acters with other species of Rhionaeschna. limiting the anal triangle (figs. 409-410, 412) (re- Second, the Marmaraeschna clade (node 20-21, fig. versed to a long fusion in the punctata-group of 438), characterized by five autapomorphies as follows: Rhionaeschna, fig. 411). Within this group, some (1) the recessed frontal carina located on postfrons giv- ‘Aeshna’ species (‘A.’ brevistyla, ‘A.’ affinis, ‘A.’ ing a rounded contour to the antero-dorsal margin of williamsoniana, ‘A.’ ellioti, ‘A.’ mixta and ‘A.’ isoceles) head (figs. 2-8b), (2) pterothorax with marbled pat- are problematical, because they are interspersed at the tern of black markings and pale areas (figs. 40-47), (3) base of different genera (fig. 438, nodes 11-12, 13-14, hamular anterior process high (figs. 277-284), (4) an- 15-17). ‘Aeshna’ brevistyla is the sister group of the re- terior lamina spine wide throughout its length in lat- maining Aeshninae (node 10-11, fig. 438). ‘Aeshna’ eral view (figs. 317-324), and (5) external margin of affinis, ‘A.’ williamsoniana, ‘A.’ ellioti and ‘A.’ mixta (as male cercus approximately linear in lateral view (in R. well as ‘A.’ minuscula not included here) have a basal intricata it can be slightly concave) (figs. 359-365). position in the clade including Andaeshna and Anaci- Two clades are defined within this group, one includ-

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ing R. brevifrons, R. fissifrons and R. pallipes (node 21- ed in the ‘variegata-group’ and between them and 22, fig. 438), which share a flattened frons (figs. 3-4, Schizuraeschna and Neureclipa –groups is still uncertain 7) and slightly constricted abdomen (figs. 121-122, (basal polytomy in node 20-30, fig. 438) since no 124) and the other (node 21-23) comprising R. bre- synapomorphy is known that would join the variegata- vicercia, R. intricata, R. obscura and R. vigintipunctata group species in its own clade. characterized by the high genital lobe (figs. 236, 238, Thus, the only subgenus traditionally defined for 240, 242). the neotropical species of ‘Aeshna’ which is not mono- Third, the cornigera- and punctata-groups consti- phyletic is Hesperaeschna, comprising, according to tute another clade (node 20-24, fig. 438), character- Calvert (1956), draco, the species belonging to the ized by the male cercus lacking a sub-basal tooth (figs. cornigera-punctata clade, the species of the ‘variegata- 382-395). Both groups constitute sister groups; the group’ belonging to the clade Schizuraeschna-Neurecli- synapomorphies of the cornigera-group species (node pa-variegata and ‘Aeshna’ williamsoniana. My exami- 24-28, fig. 438) consist of the male sterna IX posterior nation of the male holotype of ‘A.’ williamsoniana to genital opercula and X with a bright light-blue spot (deposited at UMMZ), revealed the absence of the ven- (figs. 143-150b) and the male cercus with concave lat- tral tubercle in the first abdominal segment, which eral margin and extreme apex pointed and ventrally would exclude this species from the group. Besides, it bent (figs. 382-389). Synapomorphies of the punctata- does not share cerci morphology, wing venation or group species (node 25-26, fig. 438) include the nu- color pattern with any of the groups of Rhionaeschna. merous small denticles widely distributed on the later- Its phylogenetic position is unclear, but it does not be- al and ventral surface of the genital lobe (figs. long to Rhionaeschna; its closest relative in the 270-275) and the whole tip of male cercus ventrally Neotropical region most likely belongs to the clade in- bent (figs. 290-295). Within the cornigera-group, R. cluding Andaeshna and Anax. manni, R. psilus and R. vazquezae form a clade (node The phylogenetic relationships suggested in the pre- 29-30, fig. 438) distinguished by the angled dorso-dis- sent study should be considered tentative; it is clear tal crest of male cercus (figs. 384, 388-389). Two that further work is required to further understand re- clades are defined within the punctata-group, one in- lationships within the main groups discussed here. cluding R. biliosa, R. condor and R. joannisi (node 26- However, it can be concluded that current classifica- 27, fig. 438), which share an angled dorsodistal crest tion of the Neotropical components comprising ‘Aesh- in male cercus (figs. 290-292), and another one with na’ is artificial. A reclassification of the group including R. decessus, R. eduardoi and R. punctata (node 26-28, Rhionaeschna, Andaeshna, Anaciaeschna, Hemianax and fig. 438) characterized by the high male genital lobe Anax is necessary but it will only be possible once the (figs. 274-275). positions of the enigmatic species of ‘Aeshna’ (‘A.’ affi- Fourth, Schizuraeschna, Neureclipa and variegata- nis, ‘A.’ brevistyla, ‘A.’ ellioti, ‘A.’ isoceles, ‘A.’ mixta and group species form a monophyletic unit (node 20-30, ‘A.’ williamsoniana) are elucidated. fig. 438), characterized by the origin of two rows of cells between RP1 and RP2 distal to pterostigma or at its BIOGEOGRAPHIC ANALYSIS distal end (figs. 398-400), lateral carina of male cercus concave in lateral view and sub-basal tooth present (re- Rhionaeschna is a New World genus distributed duced in R. confusa and R. californica) (figs. 366-381). from southern Argentina to southern Canada. The Schizuraeschna and Neureclipa each represent a clade; group is mostly Neotropical, with its highest diversity the Schizuraeschna clade (node 30-31, fig. 438) is char- along the Andean mountain range of South America acterized by the male cercus inner margin with a ven- between Venezuela and Bolivia (fig. 466). It is not tral process at distal 25% and dorso-distal crest trian- known from the Amazonian basin. Generalized tracks gular in lateral view (figs. 366-369) and the Neureclipa of the Rhionaeschna groups used in the present work, clade (node 30-34, fig. 438) by the free supratriangle which with the exception of the variegata-group repre- (Fig 399). Within Schizuraeschna clade, R. jalapensis, sent monophyletic units, are depicted in figs. 460- R. multicolor and R. mutata constitute a subgroup char- 465. Marmaraeschna species are limited to South acterized by the larger ventral process and the tip of America from central Chile and the hills of Central male cercus ventrally curved (figs. 367-369) and R. Argentina along the Andes to Venezuela (fig. 460). multicolor and R. mutata are sister species, having the The punctata-group shows a restricted distribution entire tip of male cercus ventrally curved (figs. 368- forming a narrow semicircle from Venezuela along the 369). Within Neureclipa, R. elsia and R. galapagoensis Andes to Bolivia and Southeastern Brazil (fig. 465) are sister species, the synapomorphy joining them be- whereas the cornigera-group is the most widely distrib- ing the presence of only a few apical denticles on the uted encompassing the whole range of the genus with ventral tubercle of abdominal segment I (figs. 250- the exception of North America (fig. 464). Schizu- 251). The relationship among the species here includ- raeschna species are restricted to Central and North

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America (fig. 461), and Neureclipa species to South same domain (Andino-patagónico of Cabrera & Will- America (fig. 462). ink 1980), or even the same region (Andina of Mor- Several species of Rhionaeschna have a restricted dis- rone 1999) in the biogeographic schemes of South tribution (figs. 439-459), and six areas of endemism America, which agrees well with the high association were established based on them (fig. 466). These areas value found in the present study between these two ar- are (1) Austral South America, encompassing S Ar- eas (figs. 467-468). Yungas, comprising forested areas gentina and Chile, with R. brevifrons, R. pallipes, R. of the E Andes, belongs to the same biogeographic di- vigintipunctata, R. absoluta, R. bonariensis, R. confusa, vision as the Paranense area in all schemes (Amazon- R. variegata, R. planaltica, R. haarupi, and the endem- ian domain or subregion), and this association is ex- ic R. diffinis, (2) Paranense, including Uruguay, S emplified by the distribution of the punctata-group, Brazil, most of Paraguay and NE Argentina, with R. with species occurring either in the Yungas (R. biliosa, bonariensis, R. confusa, R. planaltica, and endemics R. R. condor, R. demarmelsi, R. joannisi), or in the Para- brasiliensis, R. pauloi, R. decessus, R. eduardoi, and R. nense province (R. decessus, R. eduardoi, R. punctata). punctata; (3) Andes, ranging from NW Argentina and Tepuis are considered a domain different from the Chile to Venezuela along the Andes, with R. brev- Amazonian domain by Cabrera & Willink (1980), ifrons, R. pallipes, R. vigintipunctata, R. absoluta, R. and as a province of the Amazonian subregion by cornigera, R. haarupi, R. planaltica, R. psilus, R. varie- Morrone (1999). According to Halffter (1965), when gata, and endemics R. brevicercia, R. fissifrons, R. intri- the Central American land bridge connected South cata, R. obscura, R. elsia, R. galapagoensis, R. biliosa, R. and North America during the upper Cretasic and joannisi, R. demarmelsi, R. condor, R. marchali, R. per- lower Eocene, a major immigration of South Ameri- alta, R. tinti; (4) Tepuis, encompassing SE Venezuela, can insects towards North America took place. During with R. planaltica, and endemics R. draco and R. nubi- the Tertiary, flora and insects of Neotropical origin gena; (5) Central America, ranging from Panama to spread northward to about 49 degrees in western SE U.S.A., with R. californica, R. multicolor, R. cornig- North America and lower latitudes eastward, and pro- era, R. psilus, and endemics R. dugesi, R. jalapensis, R. gressively retracted during Miocene and Pliocene, manni and R. vazquezae; and (6) North America, in- with relicts being found today in southern parts of the cluding U.S.A. and S Canada, with R. californica, R. western and eastern United States. But several multicolor and the endemic R. mutata. Neotropical insects still exist as far north as Alberta, Only a few species share different areas of en- New York and Michigan (Halffter 1965), which are demism as reflected in the low association values ob- related to South American elements (i.e. Triatoma tained in the similarity analysis (fig. 468). Austral [Heteroptera: Reduviidae], several species of Scarabei- South America and Andes form the highest similarity dae and Cerambicidae [Coleoptera]). Thus, North nucleus to which Paranense and later Tepuis are and Central American areas of endemism in Rhion- joined. North America and Central America form a aeschna may be the result of this same process, where second nucleus joined to the first one by a low associ- several species evolved in North and Central America ation value. after the expansion and subsequent isolation from Parsimony analysis of endemicity resulted in two South America. area-cladograms (length 52, CI 86, RI 66). One of Species of Rhionaeschna are lacking from the Brazil- them shows the same relationship among areas ob- ian shield (Amazonic province of Cabrera & Willink tained with the similarity analysis (fig. 467a); the sec- 1980; Morrone 1999). Distribution of Rhionaeschna- ond differs in the position of the Tepuis as the sister related taxa show a low diversity in Africa (‘A.’ rileyi, group of the remaining areas (fig. 467b). ‘A.’ subpupillata, ‘A.’ moori in Africa; ‘A.’ brevistyla in The relationships among areas of endemicity of Australia and New Zealand, Andaeshna in the Andes Rhionaeschna agree broadly with the hierarchy of the and ‘A.’ williamsoniana in Central America, ‘A.’ isoce- biogeographic subdivisions of South America pro- les in S Europe and Middle East, and Anaciaeschna, posed by Cabrera & Willink (1980) and Morrone Hemianax and Anax species with the highest diversity (1999). There are no subantarctic endemics, and in in the Indo-Australian region). These facts suggest a accordance with the schemes suggested by Morrone trans-Pacific rather than trans-Atlantic (Gondwanian) (1999) and Ringuelet (1961), the species occuring in track for the group to which Rhionaeschna belongs, as the Subantarctic province of the Antarctic region of has been hypothesized for other groups of similarly Cabrera & Willink (1980) are also present in Patag- distributed Odonata (i.e. Epallagidae (Euphaeidae + onian and Andean provinces. Patagonian, Monte and Polythoridae), Gynacanthini, De Marmels 2000; Chilean provinces, included in the Austral South Megapodagrion-complex, De Marmels 2001b). America area, and the drier regions of the Andean area (corresponding to the Desierto, Prepuneña, Puneña and Altoandina provinces) are encompassed in the

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ACKNOWLEDGEMENTS manidades y Ciencias (Ciencias Biológicas) 1 (3): 29-48. Abenante, Y. P., 1980. Morfología externa del adulto de Aeshna (N.) bonariensis Rambur, 1842. Odonata: I thank Dr. Rosser W. Garrison for his generous as- Anisoptera. II Contribución. – Revista de la Faultad de sistance, support, and constructive criticism during Humanidades y Ciencias (Ciencias Biológicas) 1 (10): the realization of this work. Thanks are due to Dr. 105-152. Dennis Paulson, Dr. Jürg De Marmels, Dr. Günther Abenante, Y. P. & M. E. Philippi, 1982. Lista preliminar de Peters, Dr. Thomas W. Donnelly, and Mr. Jan van los odonatos del Uruguay. – Notulae odonatologica 1 (9): 151. Tol for critically reading the manuscript and provid- Alayo, D. P., 1968. Las libelulas de Cuba (Insecta-Odona- ing helpful corrections and comments. Dr. Dennis R. ta). – Torreia, Nueva Serie 2 (1): 1-102. Paulson, Dr. Kenneth J. Tennessen and Dr. Thomas Beatty, G. H. & A. F. Beatty, 1969. Evolution and specia- W. Donnelly generously provided unpublished infor- tion in the subgenus Schizuraeschna, with observations on mation concerning biology and distribution of several Aeshna (Schizuraeschna) mutata Hagen (Odonata). – Pro- species. The following curators and individuals placed ceedings of the Pennsylvania Academy of Science 43: 147-152. specimens and information at my disposal: Dr. Jürg Bechly, G., 1996. Morphologische Untersuchungen am De Marmels (MIZA) sent pictures, illustrations and Flügelgeäder der rezenten Libellen und deren Stamm- measurements of the female of Rhionaeschna de- gruppenvertreter (Insecta; Pterygota; Odonata), unter marmelsi and loans; Dr. Jérôme Constant (ISNB) and besonderer Berücksichtigung der Phylogenetischen Sys- Dr. Fritz Geller Grimm (MWNH) provided pho- tematik und des Grundplanes der *Odonata. – Petalura tographs of type specimens and assisted during visits (special volume) 2: 1-402. Beckemeyer, R., 2001. ‘How far to Wiwilí’ ‘Quince minu- to their collections; Dr. Jean Legrand (MNHN), Dr. Ja- tos! (Fifteen minutes)’ A Nicaraguan Adventure. – Argia son Weintraub (ANSP), Dr. Heinz G. Schroeder 13 (3): 9-14. (SMFD), Dr. Bernhard Misof (ZFNB), Dr. Philip D. Belle, J., 1991. A visit to the Galapagos Islands. – Selysia 20 Perkins (MCZ), Dr. Adolfo Cordero Rivera (Ponteve- (1): 2. dra), Dr. Victor Monserrat (Madrid), Dr. Gloria Behrstock, R.A., 1998. Notes on the first record of Masó (Barcelona), Dr. Syoziro Asahina and Dr. Turquoise-tipped darner (Aeshna psilus) in Arizona. – Ar- gia 10 (4): 11-12. Masaaki Tomokuni (Tokyo) provided information Bolivar, I., 1884. Artrópodos del viaje al Pacífico verificado about type specimens in their collections. For their de 1862 a 1865 por una comisión de naturalistas enviada kind assistance during examination of their collections por el gobierno Español. Insectos Neurópteros y Or- or loan of material I also thank Mr. Mark O’Brien tópteros. – Madrid: 1-8. (UMMZ), Dr. Oliver Flint and Dr. Nancy Adams Boomsma, T. & S.W. Dunkle, 1996. Odonata of Belize. – Odonatologica 25 (1): 17-29. (USNM), Dr. Brian Brown and Mr. Brian Harris Böttger, K. & G. Jurzitza, 1967. Beitrag zur Faunistik, (LACM), Dr. Dennis Paulson (Seattle), Dr. Kenneth Ökologie und Biologie der Odonaten von Südchile. – Tennessen (Florence), Dr. Janira M. Costa (MNRJ), Beiträge zur neotropischen Fauna 5 (1): 22-44. Dr. Angelo Machado (Rio de Janeiro), Dr. Enrique Brauer, F., 1865. Vierter Bericht über die auf der Weltfahrt González Soriano (UNAM), Dr. Giuliano Onore der kais. Fregatte Novara gesammelten Neuropteren. – (QCAZ), Dr. Ariel Camousseight (MNNS), Dr. Jorge Verhandlungen der zoologisch-botanischen Gesellschaft in Wien 14: 903-908. Solervicens (IEUM), Dr. Carlos Esquivel (INBC), Mr. Brauer, F., 1866. Neuroptera. – In: Reise der Österreichis- Jan van Tol (RMNH), Dr. David Goodger (BMNH), Dr. chen Fregatte ‘Novara’ um die Erde in den Jahren 1857, Graham S. Vick (Crossfields), Dr. Ulrike Aspöck 1858, 1859 unter der Befehlen des Commodore B. von (NMW), Dr. Günther Peters (ZMHB), Mr. Bill Mauf- Wüllerstorf-Urbair. Zoologischer Theil. Wien, 2 (4): 104 fray (FSCA), Dr. Mitsutoshi Sugimura (Nakamura), pp, 2 pls. Dr. Axel Bachmann (MACN), Mr. Hector Ferreyra Cabrera, A. L. & A. Willink, 1980. Biogeografía de Améri- ca Latina. – Secretaría General O.E.A., (Biol.), Washing- (MLPA), Dr. Sergio Roig (IADIZA), Dr. Eduardo ton D.C.,13: vi + 122 pp. Dominguez and Dr. Carlos Molineri (IMLA). Calvert, P. P., 1895. The Odonata of Baja California, Mex- This research was supported by a fellowship from ico. – Proceedings of the California Academy of Science 2 the FUNDACION ANTORCHAS, and it was carried out (4): 463-558. during a postdoctoral tenure at the Natural History Calvert, P. P., 1905. Insecta Neuroptera. Odonata. – In: Museum of Los Angeles County. I am indebted to R.H. Porter & Dulau Co. Biologia Centrali Americana, both institutions for providing support and facilities London, pp. 145-212. Calvert, P.P., 1907. Insecta Neuroptera. Odonata. – In: necessary to conduct this study. R.H. Porter & Dulau Co. Biologia Centrali Americana, London, pp. 309-404. Calvert, P. P., 1908. The composition and ecological rela- REFERENCES tions of the Odonate fauna of Mexico and Central Amer- Abenante, Y. P., 1978. Morfología externa del tercer estadio ica. – Proceedings of the Academy of Natural Sciences of larval de Aeshna (Neureclipa) bonariensis Rambur 1842 Philadelphia 60: 460-491. (Odonata, Anisoptera). – Revista de la Facultad de Hu- Calvert, P. P., 1909. Contribution to a knowledge of the

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A description of the female of Aeshna dra- Deutsche entomologische Zeitschrift 518-531. co Rácenis, 1958 (Anisoptera: Aeshnidae). – Odonatolog- Ris, F., 1913. Neuer Beitrag zur Kenntnis der Odonaten- ica 22 (1): 93-99. fauna von Argentina. – Mémoiresde la Sociéte royal ento- von Ellenrieder, N., 1999. Description of the last larval in- mologique de Belgique 22: 55-102. star of Aeshna (Hesperaeschna) cornigera planaltica Ris, F., 1918. Libellen (Odonaten) aus der Region der Calvert, 1952 (Odonata: Aeshnidae). – Revista de la So- amerikanischen Kordilleren von Costa Rica bis Catamar- ciedad entomológica Argentina 58 (3-4): 151-156.

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von Ellenrieder, N., 2000. Aeshna tinti nov. spec. from Chile Walker, E. M., 1958. The Odonata of Canada and Alaska, and redescription of Aeshna elsia Calvert, 1952 (Anisoptera: vol. 2. Part III: The Anisoptera, four families. – Universi- Aeshnidae). – Odonatologica 29 (4): 347-358. ty of Toronto Press, Toronto.xi+ 318 pp. von Ellenrieder, N., 2001a. A synopsis of the Patagonian Watson, J. A. L., 1992. The affinities of Aeshna brevistyla species of the genus Aeshna Fabricius (Anisoptera: Aesh- (Rambur) (Anisoptera: Aeshnidae). – Odonatologica 21 nidae). – Odonatologica 30 (3): 299-235. (4): 453-471. von Ellenrieder, N., 2001b. The larvae of Patagonian Aesh- Williamson, E. B., 1908. Three related American species of na Fabricius species (Anisoptera: Aeshnidae). – Odonato- Aeshna (Odonata). – Entomological News 19: 264-271, logica 30 (4): 423-434. 301-308. von Ellenrieder, N., 2002. A phylogenetic analysis of the ex- Williamson, E. B. & J. H. Williamson, 1930. Two new tant Aeshnidae (Odonata: Anisoptera). – Systematic En- Neotropical Aeshnines (Odonata). – Occasional Papers tomology (London) 27: 1-31. of the Museum of Zoology, University of Michigan 218: von Ellenrieder, N. & J. Martins Costa, 2002. A new species 1-15. of Aeshna, A. brasiliensis (Odonata, Aeshnidae) from Zimsen, E., 1964. The type material of I.C. Fabricius. – South and Southeastern Brazil, with a redescription of its Munksgaard, Copenhagen 656 pp. larva. – Neotropical Entomology 31 (3): 1-8. von Ellenrieder, N. & J. Muzón, 2003. Description of the last larval instar of Aeshna (Marmaraeschna) pallipes Fras- er, 1947 (Anisoptera: Aeshnidae). – Odonatologica 32 (1): 95-98. Walker, E. M., 1912. The North American Dragonflies of Received: 12 July 2002 the Genus Aeshna. – University of Toronto Studies, Bio- Accepted: 5 December 2002 logical Series, pp. 1-213.

120 Downloaded from Brill.com10/08/2021 11:25:36PM via free access  E: Rhionaeschna synopsis ? 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3-4 2-3 2-3 2-3 2-3 2-3 2-3 2-3 2-3 2-3 2-3 2-4 3-4 2-3 al rows ? 8-9 5-9 5-8 6-8 7-8 8-12 8-11 9-13 9-12 9-12 8-11 8-11 8-11 9-11 5-11 5-10 7-13 6-12 9-11 4-11 6-12 6-18 9-13 9-10 8-13 9-11 8-11 9-13 8-10 9-11 7-10 9-10 10-11 12-14 13-14 11-14 11-14 13-15 al cells ? 3 3 3 2 2 3 2 3 3 2 3 5 4 3 3 2-3 2-3 1-3 2-4 2-3 2-4 1-3 2-3 2-4 2-3 2-3 2-3 2-4 1-3 2-3 2-4 3-4 2-3 2-3 2-4 2-3 4-6 3-4 br Hw ? 3 3 3 3 3 5 3 3 3-4 2-4 2-3 2-4 2-3 2-5 2-4 2-4 3-4 3-4 3-4 2-3 2-4 2-3 2-3 2-3 2-4 2-3 1-3 1-4 2-5 3-4 3-4 2-4 2-3 3-4 3-4 4-5 5-6 3-5 br Fw ? 3 3 3 3 3 3 3 3 3 3 3 3-4 2-3 2-3 3-4 2-4 2-4 3-4 2-4 2-3 2-4 2-3 2-3 2-3 2-3 2-4 2-3 1-3 2-3 2-4 2-4 3-4 2-4 3-4 3-4 3-4 2-3 3-4 frk Hw ? 3 3 3 3 3 3 3 3 3 3 3 3-4 2-3 2-3 3-4 2-4 2-4 2-4 2-4 2-3 2-4 2-3 2-3 2-3 2-3 2-4 2-3 1-3 1-3 2-4 2-4 2-4 3-4 2-3 3-4 2-3 2-3 2-3 frk Fw ? 3 3 3 3 3 3 3 3 3 3 3 4 3 5 3 3 4 3 2 3 3 3-4 3-4 3-4 3-4 2-3 3-4 3-4 2-3 2-4 2-3 2-3 2-3 3-4 2-5 2-4 3-4 2-4 cua Hw ? 4 3 4 3 3 3 4 4 3 5 4 4 4 3 4-5 3-4 4-5 3-4 3-4 4-5 3-5 3-4 3-4 3-4 3-5 2-3 2-4 3-4 3-5 3-4 4-5 3-5 2-5 3-5 4-5 5-6 3-4 3-4 cua Fw 2 2 2 2 2 2 2 2 2 2 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1-2 1-2 1-2 1-2 1-2 1-2 1-2 2-3 sbtr Hw 2 2 2 2 2 2 2 2 2 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1-2 1-2 1-2 1-2 1-2 1-2 1-2 1-2 sbtr Fw 4 4 4 4 4 4 4 5 4 4 4 4 4 4 6-7 3-4 2-3 3-4 3-4 4-5 3-4 3-4 4-5 2-4 2-4 2-4 2-4 4-6 3-5 3-4 3-4 3-6 4-5 4-5 4-5 4-5 4-5 4-5 3-4 tr Hw 5 4 5 5 5 5 5 5 4 4 4 6-7 3-5 2-4 4-5 4-5 4-5 4-5 4-5 4-5 4-5 4-5 2-5 2-4 2-4 2-4 3-5 4-5 2-4 2-4 3-6 4-5 4-6 4-5 4-5 4-5 4-6 3-4 4-5 tr Fw 3 2 2 2 0 2 2 3 2 3 2 1-2 1-2 1-2 1-2 1-2 1-2 1-2 2-3 0-1 0-1 0-1 0-3 1-3 1-2 1-2 1-3 2-3 1-2 2-3 3-4 2-3 3-5 2-3 2-3 0-1 1-2 1-2 2-3 sptr Hw 4 3 0 2 2 3 2 fork at pterostigma level; br: bridge crossveins; al: anal loop. 1-3 1-2 0-2 1-3 2-3 0-2 2-3 2-3 2-3 2-3 3-4 0-2 0-1 0-2 0-1 0-3 1-4 1-2 1-3 0-3 2-4 2-3 2-4 2-4 1-3 2-4 3-4 4-5 4-6 3-4 2-3 3-4 2 IRP sptr Fw 23 12 6-9 7-9 8-10 9-11 6-11 9-11 7-13 9-11 9-11 8-12 10-12 11-13 10-12 16-19 12-14 10-13 11-12 12-13 12-15 12-14 13-14 12-13 11-13 10-14 11-13 12-13 12-15 13-16 12-14 13-14 13-17 15-17 16-20 12-14 11-16 16-18 13-14 Px Hw 10 12 7-8 7-9 6-7 7-9 5-9 7-9 9-11 9-11 8-10 8-10 9-11 9-12 8-10 9-15 18-19 13-16 12-13 11-12 10-12 12-13 11-13 12-13 10-14 10-11 10-11 10-12 12-13 12-15 10-11 11-12 11-14 11-15 13-15 11-12 10-13 12-15 12-13 Px Fw 14 7-9 8-9 8-10 7-10 8-10 8-10 8-11 9-11 9-11 7-12 7-10 8-12 8-10 8-10 8-12 9-11 8-11 9-10 6-10 8-10 7-13 9-13 9-10 9-13 9-10 8-12 9-11 9-10 13-14 16-17 11-12 11-14 12-13 11-14 12-14 13-15 11-13 12-14 Ax Hw 20 9-14 20-23 13-16 12-14 11-14 13-16 13-17 11-15 12-15 15-18 16-19 15-17 16-19 10-17 11-15 11-15 11-15 13-15 12-17 13-15 11-16 13-15 11-16 11-19 14-18 14-16 16-20 17-21 16-17 13-18 18-19 13-19 19-22 19-22 18-19 13-17 17-22 14-15 Ax Fw Table 1. Wing characters. Fw: fore wing; Hw: hind Ax: antenodal crossveins; Px: posnodal sptr: supratriangle tr: triangle cells; sbtr: subtriangle cua: cubito-anal crossveins; frk: number of cells between draco brevicercia brevifrons fissifrons intricata obscura pallipes vigintipunctata dugesi jalapensis multicolor mutata absoluta bonariensis diffinis elsia galapagoensis brasiliensis californica confusa marchali peralta tinti variegata cornigera haarupi manni nubigena pauloi planaltica psilus vazquezae biliosa condor decessus demarmelsi eduardoi joannisi punctata

121 Downloaded from Brill.com10/08/2021 11:25:36PM via free access T  E,  146, 2003

Table 2. Data matrix of 39 characters for 73 ingroup taxa and the outgroup Oreaeschna dictatrix. All question marks refer to unknown character states.

Species 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 Weight of 0 1 1 1 1 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 0 1 1 1 1 1 1 0 0 1 0 0 1 1 0 1 1 1 0 character 4 0 0 0 0 3 0 3 1 5 6 0 0 6 0 6 5 4 4 4 1 0 0 0 0 0 0 2 3 0 1 1 0 0 4 0 0 0 1 dictatrix 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 isoceles 0 0 0 0 1 0 0 0 1 0 1 0 1 2 1 0 0 0 0 0 1 0 1 1 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 jaspidea 0 2 0 0 1 0 0 1 1 1 2 0 0 1 1 0 0 0 0 3 1 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 triangulifera 0 2 0 0 1 0 0 1 1 1 2 0 0 1 1 0 0 0 0 3 1 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 andresi 0 0 0 0 1 1 0 1 0 2 0 0 1 2 1 0 0 1 0 0 0 0 1 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 rufipes 0 0 0 0 1 1 0 1 0 2 0 0 1 2 1 0 0 1 0 0 0 0 1 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 draco 0 0 0 0 0 1 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 brevicercia 1 0 1 0 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 brevifrons 1 0 1 1 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 0 1 0 0 0 0 0 1 0 1 0 1 1 0 0 1 0 0 0 0 fissifrons 1 0 1 1 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 0 1 0 0 0 0 0 1 0 1 0 1 1 0 0 1 0 0 0 0 intricata 1 0 1 0 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 obscura 1 0 1 0 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 pallipes 0 0 1 1 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 0 1 0 0 0 0 0 1 0 1 0 1 1 0 0 1 0 0 0 0 vigintipunctata 1 0 1 0 0 2 0 1 0 2 1 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 cornigera 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 1 0 0 0 1 planaltica 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 1 0 0 0 1 haarupi 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 1 0 0 0 1 nubigena 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 1 0 0 0 1 psilus 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 2 0 0 0 1 manni 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 2 0 0 0 1 vazquezae 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 0 2 0 0 0 1 absoluta 0 0 0 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 bonariensis 0 0 0 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 diffinis 0 0 0 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 elsia 0 0 0 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 0 2 1 0 0 1 0 0 0 0 galapagoensis 0 0 0 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 0 2 1 0 0 1 0 0 0 0 brasiliensis 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 californica 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 confusa 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 marchali 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 peralta 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 tinti 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 variegata 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 1 0 0 0 0 biliosa 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 2 0 0 0 2 demarmelsi 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 2 condor 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 2 0 0 0 2 joannisi 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 2 0 0 0 2 decessus 0 0 0 0 0 0 0 ? ? 0 ? 0 ? ? 0 ? 0 ? 3 1 2 0 0 ? 0 0 1 0 0 0 0 0 0 0 1 0 0 0 2 eduardoi 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 2 punctata 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 2 dugesi 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 2 0 1 0 0 jalapensis 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 2 0 1 0 1 multicolor 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 2 0 1 0 2 mutata 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 2 1 0 0 2 0 1 0 2 williamsoniana 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 brevistyla 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 affinis 0 0 0 0 0 1 0 1 1 0 1 0 0 1 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 0 0 0 clepsydra 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 eremita 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 interrupta 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 canadensis 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 1 crenata 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 cyanea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1 constricta 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1 walkeri 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1 palmata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1 persephone 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1

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petalura 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1 umbrosa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 1 0 3 1 0 1 1 ellioti 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 2 grandis 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 2 0 1 1 0 0 0 0 0 0 0 0 2 0 0 0 1 0 0 0 0 juncea 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 2 0 2 0 1 1 0 0 0 0 0 0 0 0 2 0 0 0 1 0 0 0 0 tuberculifera 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 2 1 0 0 1 0 0 0 0 viridis 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 verticalis 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 2 0 0 0 2 1 0 0 0 0 0 0 0 0 2 0 0 0 1 0 0 0 1 septentrionalis 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 2 0 2 1 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 sitchensis 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 2 0 2 1 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 2 caerulea 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 2 0 2 1 0 0 0 0 0 0 0 0 ? 0 0 1 1 0 0 0 0 mixta 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 2 nigroflava 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 2 1 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 rileyi 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 2 0 0 0 0 1 0 0 0 0 0 1 1 0 0 1 0 0 0 1 osiliensis 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 1 0 0 0 1 1 0 0 0 0 0 0 0 0 ? 1 0 1 1 0 0 0 0 subpupillata 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 ? ? 0 ? 2 0 ? 0 0 1 0 0 0 0 0 1 1 0 ? 1 0 0 0 ? subartica 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 2 2 0 0 0 1 1 0 0 0 0 0 0 0 0 2 0 0 0 1 0 0 0 1

Table 3. Parsimony analysis of endemicity. Data matrix for six areas (North America, Central America, Andes, Tepuis, Paranense and Austral South America) and 54 species and monophyletic groups of Rhionaeschna: 01 draco, 02 brevicercia, 03 brevifrons, 04 fissifrons, 05 intricata, 06 obscura, 07 pallipes, 08 vig- intipunctata, 09 (brevifrons+pallipes+fissifrons), 10 (intricata+obscura+brevicercia+vigintipunctata), 11 Marmaraeschna-clade, 12 dugesi, 13 jalapensis, 14 multicolor, 15 Schizuraeschna-clade, 16 mutata, 17 (multicolor+mutata), 18 (multicolor+mutata+jalapensis), 19 abso- luta,20bonariensis,21diffinis, 22 elsia, 23 galapagoensis, 24 (elsia+galapagoensis), 25 Neureclipa-clade, 26 californica, 27 confusa, 28 brasiliensis, 29 marchali, 30 peralta, 31 variegata, 32 tinti, 33 (variegata group+Schizuraeschna+Neureclipa), 34 cornigera, 35 haarupi, 36 manni, 37 nubigena, 38 pauloi, 39 planaltica, 40 psilus, 41 vazquezae, 42 (psilus+manni+vazquezae), 43 cornigera-clade, 44 biliosa, 45 condor, 46 demarmelsi, 47 decessus, 48 eduardoi, 49 joannisi, 50 punctata, 51 (biliosa+condor+joannisi), 52 (decessus+eduardoi+punc- tata), 53 punctata-clade, 54 (cornigera+punctata).

Areas 000000000111111111122222222223333333333444444444455555 123456789012345678901234567890123456789012345678901234 root 000000000000000000000000000000000000000000000000000000 North Am 000000000000011111000000010000001000000000000000000000 Central Am 000000000001111011000000010000001101000111100000000001 Andes 011111111110000000100111100011111110001101111100101011 Tepuis 100000000000000000000000000000000000101000100000000001 Paranense 000000000000000000010000101100001000011000100011010111 Austral 001000111110000000111000101000101010001000100000000001

123 Downloaded from Brill.com10/08/2021 11:25:36PM via free access T  E,  146, 2003

frons 1 projected dorsally

a VE: Auyantepui bcVE: Auyantepui VE: Auyantepui draco

margin slight medial 2 stem of T-spot rounded concavity not strongly narrowed anteriorly

abcEC: HolotypeEC: Guama Yacu EC: Guama Yacu brevicercia frons 3 flattened

abcCH: Huasco BajoLectotype brevifrons Lectotype brevifrons

abcPE: Lectotype maita PE: Lectotype maita PE: Lectotype maita brevifrons

deep 4 frons medial flattened cleft

abcAR: Allotype AR: Allotype AR: Allotype fissifrons

Figs. 1-4. Head. – a, dorsal view; b, lateral view; c, frontal view.

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stem of T-spot strongly narrowed anteriorly

abcPE: Celendin EC: Pallatanga EC: Pallatanga intricata

abcBO: HolotypeBO: Chapare BO: Chapare obscura

frons frontal carina flattened linear

abcAR: Río ColoradoAR: Río Colorado AR: Río Colorado pallipes

abcAR: Río ColoradoAR: San Lorenzo AR: San Lorenzo vigintipunctata

Figs. 5-8. Head. – a, dorsal view; b, lateral view; c, frontal view.

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margin angled

abcUS: Leslie CanyonUS: Leslie Canyon US: Leslie Canyon dugesi

abcMX: Cuernavaca MX: Cuernavaca MX: Cuernavaca jalapensis

T-spot stem narrower than vertex

abcUS: Havasu CanyonUS: Havasu Canyon US: Havasu Canyon multicolor

T-spot stem wider than vertex

abcUS: ScotiaUS: Scotia US: Scotia mutata

Figs. 9-12. Head. – a, dorsal view; b, lateral view; c, frontal view.

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T-spot sides converging anteriorly

dark stripe on fronto- clypeal groove

abcAR: Lago Aluminé AR: Lago Aluminé AR: Cafayate absoluta T-spot sides parallel

dark stripe on fronto-ocular groove narrow clypeal lobe angled abcAR: Punta Lara AR: Punta Lara AR: Dique Cabra Corral bonariensis

dark stripe on fronto- ocular groove wide

abcAR: TrevelinAR: Trevelin AR: Trevelin diffinis

no dark stripe on fronto- clypeal groove

clypeal lobe rounded abcPE: nr Villa PE: nr Pacasmayo PE: nr Villa elsia

dark stripe

abcEC: Is. Santa Cruz EC: Is. Santa Cruz EC: Is. Santa Cruz galapagoensis

Figs. 13-17. Head. – a, dorsal view; b, lateral view; c, frontal view.

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frontal carina angled

abcBR: HolotypeBR: Holotype BR: Holotype brasiliensis

frontal carina evenly curved

dark stripe on fronto- clypeal groove

abcUS: Palo AltoUS: Palo Alto US: Palo Alto californica

no dark stripe

abcAR: Punta LaraAR: Punta Lara AR: Punta Lara confusa

abcEC: Cuenca EC: Cuenca EC: Cuenca marchali

Figs. 18-21. Head. – a, dorsal view; b, lateral view; c, frontal view.

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A > B + C C < 2B

A

B C

abcPE: Paralectotype ApurimacBO: Copacabana PE: Paralectotype Apurimac peralta

abcCH: HolotypeCH: San Pedro de Atacama CH: Holotype tinti

A A < B + C

C = or > 2B B C

a AR: Lago Villarino

bcAR: Cari Laufquen AR: Cari Laufquen

a AR: El Calafate variegata

Figs. 22-24. Head. – a, dorsal view; b, lateral view; c, frontal view.

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abcCO: LectotypeVE: Altos de Tiara VE: Altos de Tiara

abcGU: Tamahú GU: Tamahú GU: Tamahú cornigera

stem of T- spot abruptly narrowed

abcAR: San CarlosAR: San Carlos AR: San Carlos haarupi

abcMX: Holotype MX: Holotype MX: Holotype manni

Figs. 25-27. Head. – a, dorsal view; b, lateral view; c, frontal view.

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abcVE: Cerro YavíVE: Cerro Yaví VE: Cerro Yaví nubigena

abcBR: Paratype, Santana do Riacho BR: Paratype, Santana do Riacho BR: Paratype, Santana do Riacho pauloi

stem of T-spot gradually narrowed anteriorly or parallel sided

abcAR: Pereyra IraolaAR: Pereyra Iraola AR: Pereyra Iraola planaltica

abcAR: Dique ItiyuroAR: Dique Itiyuro AR: Dique Itiyuro psilus

Figs. 28-31. Head. – a, dorsal view; b, lateral view; c, frontal view.

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abcMX: Paratype, Acahizotla MX: Paratype, Acahuizotla MX: Paratype, Acahuizotla vazquezae

transverse arms of T-spot narrow

no dark stripe

abcEC: Manabi EC: Manabi EC: Manabi biliosa

T-spot stem wider than vertex

abcVE: Paratype, Tamá NP VE: Paratype, Tamá NP VE: Paratype, Tamá NP condor

transverse arms of T-spot wide

dark stripe not widened

abcVE: Paratype, Tamá NP VE: Paratype, Tamá NP VE: Paratype, Tamá NP demarmelsi

Figs. 32-35. Head. – a, dorsal view; b, lateral view; c, frontal view.

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T-spot stem narrower than vertex

abcBO: HolotypeEC: E Cornejo Astorga EC: E Cornejo Astorga joannisi

dark stripe black of strongly T-spot widened extended on antefrons

abcBR: Paratype, CatarinaBR: Paratype, Catarina BR: Paratype, Catarina eduardoi

black of T-spot not extended on antefrons

abcBR: Umuarama BR: Umuarama BR: Umuarama punctata

Figs. 36-38. Head. – a, dorsal view; b, lateral view; c, frontal view.

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pterothorax black spots uniform in color reduced

VE: Auyantepui EC: Guama Yacu PE: Abancay PE: Lectotype maita draco brevicercia brevifrons brevifrons

AR: Paratype, Río Colorado EC: Pallatanga BO: Río Carmen Mayu AR: Río Colorado fissifrons intricata obscura pallipes

AR: Lesser US: Leslie Canyon CR: Monteverde GU: Los Aposentos vigintipunctata dugesi jalapensis jalapensis

US: St. David US: Brown's Lake US: Scotia AR: Villa Ventana multicolor multicolor mutata absoluta

AR: San Antonio AR: Lago Aluminé AR: Puerto Pirámide AR: Dique Campo Alegre absoluta absoluta absoluta absoluta

Figs. 39-58. Pterothorax, lateral view.

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AR: Lago Aluminé AR: Río Santa Lucía CH: Coñaripe-Carringue AR: Lago Aluminé absoluta bonariensis diffinis diffinis

CH: N Valdivia AR: Paso Futaleufú AR: Lago Tromen AR: Ayo. Huarenchenque diffinis diffinis diffinis diffinis

AR: Los Alerces NP CH: Camarones PE: Samne CH: Camarones diffinis elsia elsia elsia

CH: Camarones PE: Lima PE: Paratype, nr Villa EC: Is. Santa Cruz elsia elsia elsia galapagoensis

epimeral stripes constricted or divided

epimeral stripes EC: Paratype, Is. Santa Cruz BR: Allotype BR: Rio Lajeado Grande BR: Paratype, Itatiaia constricted galapagoensis brasiliensis or divided brasiliensis brasiliensis

Figs. 59-78. Pterothorax, lateral view.

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mesanepisternal mesanepisternal stripe stripe incomplete complete

US: Snow Peak US: Bishop AR: Punta Lara EC: Cuenca californica californica confusa marchali

PE:Paralectotype, Apurimac PE: Cuzco BO: Belen CH: Paratype, Calama peralta peralta peralta tinti

CH: Paratype, San Pedro de Atacama CH: Holotype CH: Paratype, Río Loa CH: Paratype, Chiu-chiu tinti tinti tinti tinti

stripes linear or slightly narrowed

CH: Pucatrihue-Bahía Mansa CO: Lectotype VE: Altos de Tiara GU: Tamahú variegata cornigera cornigera cornigera

stripes with deep rounded indentations

AR: San Carlos AR: San Carlos MX: Holotype MX: Allotype haarupi haarupi manni manni

Figs. 79-98. Pterothorax, lateral view.

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mesepimeral stripe mesanepisternal mesanepisternal narrowed at stripe complete stripe incomplete midlength (broad and shallow indentation)

MX: Río Zaya BR: Paratype, Santana do Riacho VE: Cerro Yaví VE: Cerro Yaví manni pauloi nubigena nubigena

stripes with deep rounded indentations

CO: Paralectotype cornigera AR: Salta, NR 9 AR: Calilegua NP MX: NE Huatusco planaltica planaltica planaltica psilus

stripes wide

MX: La Galinda MX: Paratype, Acahuizotla MX: Paratype, Acahuizotla PE: Paratype, W Almirante vazquezae vazquezae vazquezae biliosa

stripes stripes stripes stripes divided into complete complete complete spots and wide and sinuous and linear

EC: Baños VE: Paratype, Tamá NP VE: Paratype, Tamá NP BR: Paratype, Catarina biliosa condor demarmelsi eduardoi

stripes with deep indentations or divided into spots

EC: E Cornejo Astorga EC: E Cornejo Astorga BO: Holotype BR: Umuarama joannisi joannisi joannisi punctata

Figs. 99-118. Pterothorax, lateral view.

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VE: Auyantepui a

b VE: Auyantepui

119 c VE: Allotype, Mt. Roraima draco

a EC: Baeza

b EC: Baeza

c EC: Baeza

EC: Baeza d 120 brevicercia

Figs. 119-120. Abdomen – a-b, male; a, dorsal view; b, lateral view; c-d, female: 119c, lateral view; 120c, dorsal view; d, lat- eral view.

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a CH: Maipu

b CH: Maipu

c PE: Abancay

d PE: Abancay 121 brevifrons

a AR: Paratype, Río Colorado

b AR: Paratype, Río Colorado

c AR: Paratype, Río Colorado

122 d AR: Paratype, Río Colorado fissifrons

Figs. 121-122. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a constriction of abdominal EC: Pallatanga segment III marked

b EC: Pallatanga

c constriction of abdominal PE: Cajamarca segment III marked

d PE: Cajamarca 123 intricata

AR: Río Colorado a constriction of abdominal segment III slight

b AR: Río Colorado

c constriction of abdominal AR: Río Colorado segment III slight

124 AR: Río Colorado d pallipes

Figs. 123-124. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a BO: Río Carmen Mayu

b BO: Río Carmen Mayu 125 obscura

a AR: San Lorenzo

b AR: San Lorenzo

c AR: Lesser

d AR: Lesser 126 vigintipunctata

Figs. 125-126. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a US: Leslie Canyon

b US: Leslie Canyon

incomplete medio- c longitudinal stripe US: Leslie Canyon

127 d US: Leslie Canyon dugesi

a CR: Monteverde

b CR: Monteverde PD spots small

c MX: Cuernavaca

PL spots absent 128 d MX: Cuernavaca jalapensis

Figs. 127-128. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a US: Valentine PD large and confluent with PL

PL spots present b US: Valentine

c US: Del Puerto Canyon

US: Del Puerto Canyon 129 d multicolor

a US: Scotia

b US: Scotia

US: Lake Denmark c

d US: Lake Denmark 130 mutata

Figs. 129-130. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a AR: N Buta Ranquil

AR: Comi Có b

c medio-longitudinal black stripe AR: Comi Có

AR: Comi Có 131 d absoluta

a AR: Cafayate

AR: Cafayate b medio-longitudinal pale stripe complete

c AR: Colonia Dora

AR: Colonia Dora 132 d bonariensis

Figs. 131-132. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a CH: E Caramávida

AR: Trevelin b

c CH: Angol-Vegas Blancas ML and PL confluent MD and ML separated

133 d diffinis CH: Angol-Vegas Blancas

a PE: Paratype, nr Villa

b PE: Paratype, nr Villa

c PE: Samne

PE: Samne 134 d elsia

a EC: Is. Santa Cruz

EC: Is. Santa Cruz 135 b galapagoensis

Figs. 133-135. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a BR: Holotype

b BR: Holotype

BR: Allotype c

d BR: Allotype 136 brasiliensis

US: Sonoma, Kruse Ranch Rd a

b US: Sonoma, Kruse Ranch Rd

US: Silverado c

US: Silverado 137 d californica

Figs. 136-137. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a AR: Punta Indio

b AR: Punta Indio

medio-longitudinal pale stripe

c AR: Punta Lara

d AR: Punta Lara 138 confusa

a EC: Cuenca

b EC: Cuenca

c EC: Baeza

EC: Baeza d 139 marchali

Figs. 138-139. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a PE: Paralectotype, Apurimac

b PE: Paralectotype, Apurimac

c PE: Paralectotype, Apurimac

ML and PL separated MD and ML separated

d PE: Paralectotype, Apurimac 140 peralta

a CH: Holotype

b CH: Holotype

c CH: Allotype

ML and MD confluent

CH: Allotype d 141 tinti

Figs. 140-141. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a CH: Lago Lonconao

b AR: Cari Laufquen

c CH: Lago Lonconao

d AR: Cari Laufquen

e AR: Lago Queñi

f CH: Queulat NP

g AR: El Calafate 142 variegata

Figs. 142. Abdomen – a-d, male; e-g, female; a-b, e, dorsal view; c-d, f-g, lateral view; a, c, e-f, dark form; b, d, g, light form.

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a VE: Altos de Tiara

b GU: Tamahú

PL very small to absent c bright VE: Altos de Tiara light blue spot d

CO: Lectotype

e GU: Tamahú

f EC: Napo

g CR: San José

h EC: Napo

i CR: San José cornigera 143

Figs. 143. Abdomen – a-e, male; f-i, female; a-b, f-g, dorsal view; c-e, h-i, lateral view.

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a AR: San Carlos

PL large b AR: San Carlos

c AR: San Carlos

AR: San Carlos 144 d haarupi

a MX: Holotype

b MX: Holotype

MX: Allotype c

d MX: Allotype 145 manni

Figs. 144-145. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a VE: Cerro Yaví

b VE: Cerro Yaví

c VE: Cerro Yaví

d VE: Cerro Yaví 146 nubigena

BR: Paratype, Santana do Riacho a

BR: Paratype, Santana do Riacho b

c BR: Paratype, Santana do Riacho

147 d BR: Paratype, Santana do Riacho pauloi

Figs. 146-147. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a AR: Salta, NR 9

b AR: Salta, NR 9

c AR: Salta, NR 9

d AR: Salta, NR 9 148 planaltica

a PR: El Verde st.

b PR: El Verde st. 149 psilus

Figs. 148-149. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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c MX: Laguna Azul

d MX: Laguna Azul 149 psilus

a MX: Paratype, Acahuizotla

b MX: Paratype, Acahuizotla 150 vazquezae

c EC: Baños

d EC: Baños 151 biliosa

Figs. 149-151. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.

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a EC: Santa Rosa 151

b EC: Santa Rosa biliosa

c VE: Paratype, Tamá NP 152

d VE: Paratype, Tamá NP condor VE: Paratype, Tamá NP

a AD & MD present in VII-VIII pale spots present in X 153

AL & ML present in VII-VIII b VE: Paratype, Tamá NP demarmelsi EC: E Cornejo Astorga

a AD & MD absent in VII-VIII pale spots absent in X 154

AL & ML absent in VII-VIII EC: E Cornejo Astorga b joannisi

Figs. 151-154. Abdomen – a-b, male; a, dorsal view; b, lateral view.

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c EC: Narupa

d EC: Narupa

BO: Holotype e joannisi 154

a BR: Paratype, Catarina

b BR: Paratype, Catarina eduardoi 155

BR: Umuarama a

b BR: Umuarama

c BR: Macieiras

BR: Macieiras d punctata 156

Figs. 154-156. Abdomen – a-b, male; c-e, female; a, c, dorsal view; b, d-e, lateral view.

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ventral view of abdominal right half inner margin

a outer margin VE: Auyantepui seg. III seg. IV seg. VIII seg. IX

VE: Mt. Roraima 157 b draco a

EC: Baeza

b 158 brevicercia EC: Baeza

a PE: Lectotype maita

CH: Maipu b inner and outer carinae sinuous

159 c brevifrons PE: Abancay

inner and outer carinae parallel a AR: Río Colorado

160 b fissifrons AR: Río Colorado

a EC: Pallatanga

b intricata PE: Cajamarca 161

a BO: Río Carmen Mayu

b obscura BO: Allotype 162

Figs. 157-162. Ventral terga, ventral view – a, male; b-c, female.

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a AR: Río Colorado b 163 pallipes AR: Río Colorado

a AR: San Lorenzo

AR: Lesser 164 b vigintipunctata

US: Leslie Canyon a posterior portion of inner margin convergent to outer margin in V-VII b 165 dugesi US: Leslie Canyon

a CR: Monteverde b MX: Cuernavaca 166 jalapensis

US: Valentine a posterior portion of inner margin parallel to outer margin in V-VII

b 167 multicolor US: Del Puerto Canyon

a US: Scotia

168 b mutata US: Lake Denmark

Figs. 163-168. Ventral terga, ventral view – a, male; b, female.

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a AR: Comi Có anterior portion of outer margin linear to concave in IV

169 b absoluta maximum width at distal 0.60 AR: Comi Có

a AR: Cafayate

AR: Colonia Dora 170 b bonariensis

a AR: Trevelin anterior portion of outer margin convex in IV AR: Shaman

b maximum width at basal 0.30 171 diffinis a

PE: Paratype, nr Villa

b 172 elsia CH: Arica

a EC: Is. Santa Cruz

b 173 galapagoensis EC: Paralectotype, Is. Santa Cruz

a BR: Holotype

b BR: Allotype brasiliensis 174

Figs. 169-174. Ventral terga, ventral view – a, male; b, female.

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a US: Sonoma, Kruse Ranch Rd

b US: Silverado 175 californica

a AR: Punta Lara

b AR: Punta Lara 176 confusa

a EC: Cuenca

b EC: Baeza 177 marchali

a

PE: Paralectotype, Apurimac

b PE: Paralectotype, Apurimac 178 peralta

a

CH: Holotype

b 179 tinti CH: Allotype

a CH: Queulat NP wide ventral terga

180 b variegata CH: Villa La Tapera

Figs. 175-180. Ventral terga, ventral view – a, male; b, female.

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a GU: Tamahú

c narrow ventral terga VE: Altos de Tiara

b EC: Napo

d CR: San José cornigera 181

a AR: San Carlos

AR: San Carlos b haarupi 182

MX: Holotype a

b manni MX: Allotype 183

a VE: Cerro Yaví

b nubigena VE: Cerro Yaví 184

a BR: Paratype, Santana do Riacho

185 b pauloi BR: Paratype, Santana do Riacho

a AR: Salta, NR 9

186 b planaltica BR: Itatiaia

Figs. 181-186. Ventral terga, ventral view – a, c, male; b, d, female.

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a PR: El Verde st.

MX: Laguna Azul 187 b psilus

a vazquezae MX: Paratype, Acahuizotla 188

a EC: Santa Rosa

EC: Baños 189 b biliosa

a VE: Paratype, Tamá NP condor 190

a outer margin linear in VII VE: Paratype, Tamá NP

b 191 demarmelsi VE: Allotype

a outer margin sinuous in VII EC: E Cornejo Astorga

BO: Holotype 192 b joannisi

a BR: Paratype, Catarina 193 eduardoi

a BR: Umuarama

b 194 punctata BR: Macieiras

Figs. 187-194. Ventral terga, ventral view – a, male; b, female.

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black extended over auricle

draco brevicercia brevifrons VE: Auyantepui EC: Baeza CH: Maipu

fissifrons intricata obscura CH: Paratype, Inacaliri EC: Pallatanga BO: Río Carmen Mayu

black not extended over auricle

pallipes vigintipunctata dugesi AR: Río Colorado AR: San Lorenzo US: Leslie Canyon

jalapensis multicolor mutata CR: Monteverde US: Valentine US: Scotia

Figs. 195-206. Male abdominal segments I-II, ventral view.

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absoluta bonariensis diffinis AR: Comi-Có AR: Cafayate AR: Trevelin

elsia galapagoensis brasiliensis PE: Paratype, nr Villa EC: Is. Santa Cruz BR: Paratype, Campos do Jordão

californica confusa marchali US: Snow Peak AR: Punta Lara EC: Cuenca

peralta tinti variegata PE: Paralectotype, Apurimac CH: San Pedro de Atacama CH: Villa La Tapera

Figs. 207-218. Male abdominal segments I-II, ventral view.

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cornigera cornigera haarupi VE: Altos de Tiara GU: Tamahú AR: San Carlos

pauloi manni nubigena BR: Paratype, Santana do Riacho MX: Holotype VE: Cerro Yavi

planaltica psilus vazquezae AR: Salta, NR 9 PR: El Verde St. MX: Paratype, Acahuizotla

biliosa condor EC: Santa Rosa VE: Paratype, Tamá NP

Figs. 219-229. Male abdominal segments I-II, ventral view.

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230 231

demarmelsi joannisi VE: Paratype, Tamá NP EC: Narupa

232 233

eduardoi punctata BR: Paratype, Catarina EC: Pallatanga

Figs. 230-233. Male abdominal segments I-II, ventral view.

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ventral tubercle

I II draco genital lobe brevifrons tubercle VE: Auyantepui CH: Maipu low A genital B lobe low A < 0.33B brevicercia brevifrons tubercle EC: Baeza PE: Lectotype maita high A B A < 0.50B intricata fissifrons EC: Pallatanga CH: Paratype, Inacaliri

obscura genital lobe pallipes BO: Río Carmen Mayu high and AR: Río Colorado subquadrate

genital lobe vigintipunctata dugesi tubercle AR: San Lorenzo high and US: Leslie Canyon high triangular tubercle A A low B B A < 0.50B A < 0.75B multicolor mutata US: Merrit US: Scotia

jalapensis absoluta MX: Cuernavaca AR: Cafayate numerous denticles

bonariensis diffinis AR: Dique Cabra Corral AR: Trevelin few apical denticles

elsia galapagoensis CH: Arica EC: Is. Santa Cruz

brasiliensis californica BR: Paratype, Campos do Jordão US: Snow Peak

Figs. 234-253. Abdominal segments I-II, lateral view of tubercle I and genital lobe.

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confusa marchali 254 AR: Punta Lara 255 EC: Quito

peralta tinti 256 PE: Cuzco 257 CH: San Pedro de Atacama

258 variegata 259 haarupi CH: Puerto Ramirez AR: San Carlos tubercle high A

B manni 260 A = 0.50B cornigera 263 MX: Holotype GU: Tamahú

anterior side anterior side cornigera CO: Lectotype convex linear AR: Salta, NR 9 planaltica VE: Altos de Tiara 261 264 CO: paralectotype cornigera 262 anterior side 265 concave A tubercle low B A < 0.33B pauloi nubigena 266 BR: Paratype, Santana do Riacho 267 VE: Cerro Yaví

psilus vazquezae 268 PR: El Verde St. 269 MX: Chilpancingo small denticles over lateral surface of lobe tubercle trapezoidal

biliosa condor 270 EC: Santa Rosa271 VE: Paratype, Tamá NP

A A

B B A < 0.33B A < 0.33B demarmelsi joannisi 272 VE: Paratype, Tamá NP 273EC: E Cornejo Astorga tubercle high tubercle low C C and trapezoidal A and rounded A D D

A > 0.33B B A > 0.33B B C > 0.66D C = 0.33D eduardoi punctata 274 BR: Paratype, Catarina 275 BR: Umuarama

Figs. 254-275. Abdominal segments I-II, lateral view of tubercle I and genital lobe.

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medial 276digitiform 277 medial digitiform anterior anterior projection projection process A anterior c external process internal

posterior b B

draco A > B a brevicercia b a VE: Auyantepui EC: Baeza hamular fold 278 280 c 279 c

b a a intricata a brevifrons b EC: Pallatanga brevifrons CH: Maipu PE: Lectotype maita 281 282

A c c A

B B b b A > B a a fissifrons A > B pallipes CH: Paratype, Inacaliri AR: Río Colorado 283 284 285 c c

a obscura a vigintipunctata b a dugesi BO: Río Carmen Mayu AR: San Lorenzo US: Leslie Canyon

Figs. 276-285. Hamuli – a, posterior view; b, ventral view; c, latero-external view.

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286287 288 A

B b b b

a jalapensisa multicolor A < B a mutata CR: Monteverde US: Merrit US: Scotia 289290 291

b b b

a a absoluta a bonariensis diffinis AR: Cafayate AR: Dique Cabra Corral AR: Trevelin

tip pointed tip rounded 292 293 294

b b b a a elsia a galapagoensis brasiliensis CH: Arica EC: Is. Santa Cruz BR: Paratype, Campos do Jordão 297 projection 295 296 prominent C A D

b b B b C = 0.50D a A < B a a californica confusa marchali US: Snow Peak AR: Punta Lara EC: Cuenca

projection 298 299 low C 300 D

b b b C = 0.33 D a a a peralta variegata tinti PE: Cuzco AR: Cari Laufquen CH: San Pedro de Atacama

Figs. 286-300. Hamuli – a, posterior view; b, ventral view.

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301 tip rounded 302 303

b b b a a a cornigera cornigera haarupi VE: Altos de Tiara GU: Tamahú AR: San Carlos 304tip pointed 306305

b b b a a a manni nubigena pauloi MX: Holotype VE: Cerro Yaví BR: Paratype, Santana do Riacho

tip carinated 307 309308

b b b

a a a planaltica psilus vazquezae AR: Salta, NR 9 PR: El Verde St. MX: Paratype, Acahuizotla

310tip pointed 312311 tip rounded

A A

b B b b B AB a biliosa a demarmelsi joannisi EC: Santa Rosa VE: Paratype, Tamá NP EC: E Cornejo Astorga tip pointed tip tip blunt 313recurved 315314

A

B b b b

A>B a a a condor eduardoi punctata VE: Paratype, Tamá NP BR: Paratype, Catarina BR: Umuarama

Figs. 301-315. Hamuli – a, posterior view; b, ventral view.

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dorsal 316 317 318 319

B anterior A

posterior ventral draco brevicerciaA < 2B brevifrons brevifrons VE: Auyantepui EC: Baeza CH: Maipu PE: Lectotype maita

320 321 322 323

fissifrons intricata obscura pallipes CH: Paratype, Inacaliri EC: PallatangaBO: Río Carmen Mayu AR: Río Colorado

324 325 326 327

B A

A > 2B tip directed tip curved tip curved ventrally dorsally dorsally vigintipunctata dugesi jalapensis multicolor AR: San Lorenzo US: Leslie Canyon CR: Monteverde US: Merrit

328 329 330 331 dorsal margin linear

mutatatip curved absoluta bonariensis diffinis US: Scotia AR: Cafayate AR: Dique Cabra Corral AR: Trevelin dorsally

332 333 334 335 dorsal margin concave

ventral margin elsia galapagoensis brasiliensislinear to californica CH: Arica EC: Is. Santa Cruz BR: Paratype, Campos do Jordão concave US: Snow Peak

Figs. 316-335. Spine of anterior lamina, latero-inner view.

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confusa marchali peralta ventral variegata 336AR: Punta Lara 337EC: Baeza 338 PE: Cuzco margin 339 CH: Puerto Ramirez linear to ventral concave margin convex

340tinti 341cornigera 342cornigera 343 haarupi CH: San Pedro de Atacama CO: Lectotype GU: Tamahú AR: San Carlos

347 manni MX: Holotype

spine vestigial 346 nubigena planaltica pauloi psilus 344VE: Cerro Yaví 345 AR: Salta, NR 9 BR: Paratype, Santana do Riacho 348 PR: El Verde St.

A B > 2A acute pointed B tip

349vazquezae 350biliosa 351condor 352 joannisi MX: Paratype, Acahuizotla EC: Santa Rosa VE: Paratype, Tamá NP EC: E Cornejo Astorga

354 A 353 B

B < 1.5A demarmelsi demarmelsi VE: Paratype, Las Copas eduardoi punctata punctata VE: Paratype, Tamá NP 355 BR: Paratype, Catarina 356 BR: São Paulo 357 BR: Umuarama

Figs. 336-357. Spine of anterior lamina, latero-inner view.

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VE: Auyantepui a EC: Holotype a

lateral carina lateral carina concave sub-basal tooth low linear

b b

tip bearing spine VE: Auyantepui EC: Holotype

VE: Allotype, Mt. Roraima EC: Allotype c c 358 draco 359 brevicercia

atrophied cercus

a d PE: Lectotype maita CH: Maipu

PE: Matucana e

malfomed cercus sub-basal b tooth low CH: Maipu

PE: Lectotype maita f

tip not mucronate tip mucronate c g h CH: Huasco Bajo PE: Yauyos CH: Paralectotype 360 brevifrons

Figs. 358-360. Cerci – a, b, d-f, male; c, g-h, female; a, c, d, g-h, dorsal view; b, e-f, lateral view.

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a AR: Holotype

a

PE: Celendin

sub-basal tooth low c AR: Allotype b PE: Celendin AR: Holotype b fissifrons 361

AR: Cafayate d a

EC: Pallatanga

sub-basal tooth c prominent c AR: Cafayate PE: Celendin AR: Cafayate b pallipes 362 363 intricata

a a

BO: Holotype

sub-basal tooth crest at distal 0.33 crest at distal 0.25 prominent

sub-basal tooth prominent inner and outer b BO: Holotype margins parallel b AR: Tucumán

c cercus tip c widening mucronate tip rounded distally BO: Allotype BO: Buena Vista 364 obscura vigintipunctata 365

Figs. 359-365. Cerci – a-c, male; d, female; a, c, dorsal view; b -d, lateral view.

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366 367

a a

US: Leslie Canyon MX: Los Aposentos b b

crest triangular process projected posteriorly c c small process not projected posteriorly only extreme tip sub-basal ventrally bent tooth low and blunt

tip rounded

d dugesi US: Chiricahua Mountains d jalapensis MX: Cuernavaca

a a

US: Merrit b

entire tip US: Scotia ventrally bent crest high b crest low B c AB A US: Merrit c

US: Lake Denmark d margin linear US: Alum Rock Park

tip pointed

e tip pointed

margin slightly d convex 368 US: Lake Denmark multicolor US: San Francisco mutata 369

Figs. 366-369. Cerci – a-b, d, male; c, e, female; a, d-e, dorsal view; b, mediodorsal view; c, lateral view.

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370 AR: N Buta Ranquil AR: Punta Lara a a

crest as high AR: N Buta Ranquil outer base with pale spot as base b b AR: Punta Lara

tip pointed

sub-basal tooth tip pointed prominent c c

AR: Río Aluminé AR: Punta Lara absoluta 371 bonariensis CH: Lago Ranco 372 PE: nr Villa a a

CH: Lago Ranco crest suddenly rising crest higher b than base b PE: nr Villa

tip pointed c c AR: Lago Aluminé tip rounded PE: Lima diffinis 373 elsia

a 375 BR: Holotype a

EC: Lectotype

crest gradually inner margin diverging gradually rising b b BR: Holotype EC: Is. Santa Cruz

c

tip rounded EC: Paralectotype c BR: Allotype tip pointed galapagoensis 374 brasiliensis

Figs. 370-375. Cerci – a-b, male; c, female; a, c, dorsal view; b, lateral view.

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AR: Punta Lara 376 377 tip pointed US: Snow Peak a a

AR: Punta Lara US: Snow Peak

b b tip pointed

sub-basal tooth vestigial AR: Punta Lara c US: Silverado c confusa californica 379 tip angled a

marked concavity slight concavity

a PE: Cuzco EC: Cuenca BO: Lectotype b b EC: Cuenca

d BO: Copacabana c EC: Cotundo PE: Paralectotype, Apurimac c peralta AR: Laguna Escondida

marchali 378 a CH: Holotype tip angled a

tip angled heel at inner margin CH: Holotype b AR: Laguna Escondida b

tip rounded

sub-basal tooth prominent c c CH: Allotype AR: Cari Laufquen 380tinti 381 variegata

Figs. 376-381. Cerci – a-b, male; c-d, female; a, c-d dorsal view; b, lateral view.

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tip directed distal 0.60 parallel sided externally a f tip directed posteriorly CO: Lectotype GU: Tamahú inner margin concave

b crest high and smoothly curved A g

CO: Lectotype B no sub-basal tooth A < B GU: Tamahú

EC: Napo c pointed tip e rounded tip

VE: Colonia Tovar CR: San José d 382 cornigera inner and outer margins diverging maximum width at basal 0.30

a a

AR: San Carlos MX: Holotype

lateral carina crest low, long and angled b A concave

B AR: San Carlos b A > B MX: Holotype

pointed tip

c AR: San Carlos c 383 384 MX: Allotype haarupi manni

Figs. 382-384. Cerci – a-b, f-g, male; c-e, female; a, c-f dorsal view; b, g, lateral view.

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tip directed maximum width at basal 0.33 385 386 externally

a a

VE: Cerro Yaví AR: Salta, NR 9 crest high, long and smoothly curved b A

B ventrally bent pointed apex b A < B AR: Salta, NR 9 VE: Cerro Yaví

tip pointed c VE: Mnt. Roraima c

CO: Paralectotype cornigera mucron at external margin d d VE: Cerro Yaví BR: Itatiaia nubigena 387 distal 0.66 parallel sided e

tip directed posteriorly AR: Salta, NR 9 a planaltica BR: Paratype, Santana do Riacho

tip rounded with minute medial spine b c

BR: Paratype, Santana do Riacho BR: Paratype, Santana do Riacho

pauloi

Figs. 385-387. Cerci – a-b, male; c-e, female; a, c-e dorsal view; b, lateral view.

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a a MX: Laguna Azul MX: Paratype, Acahuizotla

crest low, long and angled

crest low, long and angled b MX: Laguna Azul

b MX: Paratype, Acahuizotla

tip pointed vazquezae

c MX: Laguna Azul

psilus a EC: E Cornejo Astorga

crest angled

b

a tip pointed EC: E Cornejo Astorga VE: Paratype, Tamá NP

maximum width at medial 0.33

crest angled

b

VE: Paratype, Tamá NP c BO: Holotype condor joannisi

Figs. 388-391. Cerci – a-b, male; c, female; a, c, dorsal view; b, lateral view.

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a VE: Paratype, Tamá NP a

EC: Santa Rosa crest smoothly convex

crest angled b tip blunt

VE: Paratype, Tamá NP

maximum width at distal 0.33 EC: Santa Rosa b tip incised c VE: Allotype, Tamá NP (after De Marmels, 2001) tip mucronate demarmelsi

EC: Baños c tips converging biliosa or parallel

a BR: Umuarama

tips diverging

a b BR: Umuarama BR: Paratype, Catarina

crest with denticles tip incised

b c BR: Paratype, Catarina BR: Macieiras

eduardoi punctata

Figs. 392-395. Cerci – a-b, male; c, female; a, c, dorsal view; b, lateral view.

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sptrg

RP1

RP2 evenly curved

subtrg

MA RP3-4

Cu-A space PE: Abancay trg brevifrons

anal trg

IRP2a IRP2b

anal loop

VE: Auyantepui draco

IRP2 fork symmetrical

Figs. 396-397. Right pair of male wings.

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bent of RP2 marked

US: Leslie Canyon dugesi

two rows of cells beginning distal to pterostigma

AR: Punta Lara bonariensis

PE: Paralectotype, Apurimac peralta

Figs. 398-400. Right pair of male wings.

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AR: Campo Alegre planaltica

two rows of cells beginning at proximal end of pterostigma

GU: Tamahú cornigera

VE: Paratype, Tamá NP demarmelsi

Figs. 401-403. Right pair of male wings.

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MALAWI: Mughese Forest "Aeshna" rileyi

bending of MA marked

GERMANY: Sassenpfühle Aeshna grandis

bending of MA slight

RP2 kinked

JAPAN: Kizyoka Anaciaeschna jaspidea

IRP2 fork anal angle asymmetrical angled

Figs. 404-406. Right pair of male wings.

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bent of RP2 marked

GERMANY: Limsdorf "Aeshna" isoceles

anal angle rounded

long parallel fusion short parallel fusion

US: Angostura ITALY: Chiusi US: Leslie Canyon Aeshna palmata "Aeshna" mixta Rhionaeschna dugesi

fusion at a single point

VENEZUELA: Las Copas JAPAN: Kizyoka GREECE: Kefallini Rhionaeschna demarmelsi Anaciaeschna jaspidea "Aeshna" isoceles

Figs. 407-413. Right pair of male wings.

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A A A

B B B A= 0.50B A= 3-4B A= 8-9B CANADA: Edson GUATEMALA: Tamahú JAPAN: Kizyoka Aeshna sitchensis Rhionaeschna cornigera Anaciaeschna jaspidea

transverse ridge genital lobe no higher than margin of fossa with transverse ridge denticles restricted to a marginal row

III a JAPAN: Kizyoka Anaciaeschna jaspidea b genital lobe low with denticles on ventral surface

conical tubercle conical GUATEMALA: Tamahú tubercle a Rhionaeschna cornigera b

genital lobe high with denticles on ventral and lateral surface ventral terga cylindric posterior tubercle processes

a BRAZIL: Catarina Rhionaeschna eduardoi a BRAZIL: São Carlos Castoraeschna januaria

genital lobe overdeveloped and projected posteriorly

a MALAWI: Mughese Forest "Aeshna" rileyi

Figs. 414-416. Head, dorsal view – 417-421. Male abdominal segments I-II – a, lateral view of tubercle I and genital lobe; b, ventral view of tubercle I.

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hamular anterior process long and triangular

spine triangular and b narrow posteriorly directed hamular anterior process short spine hamular fold cylindrical not postero- differentiated ventrally from wall of directed fossa b US: Merriman a CANADA: Edson Aeshna constricta Aeshna sitchensis a b

hamular anterior process absent

hamular posterior process tuberculate b hamular subarctica posterior process longer than width of ventral a portion US: Rockaway Township US: Lincoln a Aeshna clepsydra Aeshna subarctica c verticalis long anteriorly short dorsally directed spines crest directed spines crest curved linear

apical denticles spine of on dorsal cercus crest inner distal margin concavity

bc d c d

a US: Smith Falls sub-basal carena b US: Anchorage Aeshna constricta of seg. X Aeshna eremita

Figs. 422-425. Hamuli – a, postero-ventral view; b, c, ventral view – 426-427. Male terminalia – a, latero-ventral view; b, lat- eral view; c, dorsal view; d, lateral view of epiproct.

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auricle broadly triangular

USA: Merriman GERMANY: Sassenpfühle MALAWI: Mughese Forest Aeshna constricta Aeshna grandis "Aeshna" rileyi

auricle narrow parallel sided

ARGENTINA: Nahuel Huapi GERMANY: Limsdorf FIJI Rhionaeschna variegata "Aeshna" isoceles Anaciaeschna jaspidea

a a

Basal fold of lateral lobe Basal fold of Lateral lobe lacking a Ventral lobe lacking Ventral lobe lacking medio- lateral lobe basal fold medio-longitudinal fold longitudinal fold Medio-longitudinal fold of ventral lobe

b b b

GUATEMALA: Tamahú GERMANY: Limsdorf MALAWI: Mughese Forest Rhionaeschna cornigera "Aeshna" isoceles "Aeshna" rileyi

Figs. 428-433. Auricle, ventrolateral view – 434-436. Distal segment of vesica spermalis – a, lateral view; b, dorsal view.

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Castoraeschna 2 3 Coryphaeschna 1 Remartinia Oreaeschna "Aeshna" 4 Anaciaeschna 5 "Aeshna" isoceles 6 Andaeshna 7 8 Anax 9 10 11 12 Hemianax

1. Hind wing membranule longer than one-fourth of anal wing margin

2. Microtrichia patches on ventral surface of distal segment of vesica spermalis

3. Anterior surface of distal segment of vesica spermalis closed 4. Lateral supplementary carinae at least in some abdominal segments (from IV to IX) of females, males or both sexes 5. Medial surface of labial palp fixed hook with a denticle

6. Hind wing membranule longer than three-fourths of anal wing margin

7. Male hindwing anal angle rounded 8. Fusion of AA & AP with AA2b (anterior and posterior veins of male anal triangle) at a single point at anal angle 9. Lateral lobes of distal segment of vesica spermalis unfolded 10. Microtrichia of the head-arrester system with rounded tips, forming two fields on each side of the head 11. Anal triangle not defined (AA2b not thickened)

12. Auricles absent

Fig. 437. Cladogram (strict consensus tree) of Aeshninae (after von Ellenrieder 2002).

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Strict consensus tree dictatrix Oreaschna viridis of 472 MPT obtained grandis after successive tuberculifera 3 juncea weighting 1(1) verticalis [CI 72, RI 95] subarctica 4 1(0) clepsydra eremita interrupta Aeshna canadensis 5 1(1) osiliensis 2 1(1) nigroflava crenata 6 septentrionalis 1(0) 7 stichensis 1 3(1) caerulea cyanea constricta walkeri 8 palmata 7(5) persephone petalura umbrosa brevistyla affinis williamsoniana "Aeshna" 11 ellioti 1(0) 13 12 1(0) mixta 9 1(0) jaspidea 15 Anaciaeschna 2(0) 5(3) triangulifera 14 4(3) isoceles "Aeshna" 16 andresi 3(3) 17 Andaeshna 4(0) rufipes rileyi 10 19 "Aeshna" 1(0) 3(2) subpupillata draco brevifrons 22 fissifrons 2(1) pallipes 21 8(3) brevicercia inticata 23 1(0) obscura 18 Character state changes 1(0) vigintipunctata * = homoplasic changes Node (1,2): 21.2*, 22.1 demarmelsi Node (2,3): 12.1, 16.3, 17.1* biliosa 1(0) 26 Node (3,4): 35.1* condor 25 Node (4,5): 34.1 2(0) joannisi Node (5,6): 17.0* decessus Node (6,7): 2.1, 16.2*, 19.2* 1(0) 27 Node (2,8): 18.1*, 30.1, 33.1, 35.3, 36.1, eduardoi 24 punctata 38.1, 39.1* 20 1(0) Node (1,9): 8.1*, 14.1* 1(1) cornigera Node (9,10): 11.1* planaltica Node (10,11): 9.1* 28 haarupi Rhionaeschna Node (11,12): 21.1* nubigena Node (12,13): 39.2* 3(1) Node (12,14): 5.1, 15.1, 23.1, 28.1* psilus Node (14,15): 2.2, 10.1, 11.2, 20.3*, 39.1* 29 manni Node (14,16): 13.1, 14.2, 24.1 1(0) vazquezae Node (16,17): 6.1*, 10.2*, 18.1*, 29.1* brasiliensis Node (14,18): 35.1* Node (18,19): 20.2, 25.1, 32.1* californica Node (18,20): 27.1 confusa Node (20,21): 1.1*, 3.1, 6.2, 10.2*, 16.1*, marchali 19.1, 31.1*, 32.1* peralta Node (21,22): 4.1, 29.1* tinti Node (21,23): 20.1* variegata Node (20,24): 32.0* 30 Node (24,25): 21.2*, 39.2* 3(0) dugesi Node (25,26): 35.2* 31 jalapensis 2(1) Node (25,27): 20.1* 32 multicolor Node (24,28): 26.1, 31.2*, 39.1* 1(0) 33 mutata Node (28,29): 35.2* 1(0) Node (20,30): 9.1*, 31.2*, 32.1* absoluta Node (30,31): 35.2*, 37.1 bonariensis Node (31,32): 39.1* 34 diffinis Node (32,34): 39.2* 1(1) elsia Node (30,34): 7.1 35 Node (34,35): 27.2 1(1) galapagoensis

Fig. 438. Cladogram of Rhionaeschna species.

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69 W 67 W 65 W 63 W 61 W 59 W 7 N

VENEZUELA 5 N

COLOMBIA

3 N

BRAZIL

1 N

draco

100 km

80 W 70 W 60 W 50 W 40 W

PERU

10 S BRAZIL

BOLIVIA

PARAGUAY CHILE 20 S

ARGENTINA

URUGUAY 30 S brevifrons fissifrons

1000 km

Figs. 439-440. Rhionaeschna dot maps.

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71 W 70 W 60 W 50 W 40 W

PARAGUAY CHILE

ARGENTINA

30 S

URUGUAY

35 S

pallipes

400 km

83 W 78 W 73 W 68 W 63 W 10 N

VENEZUELA

5 N

COLOMBIA

0 BRAZIL

ECUADOR brevicercia intricata

5 S PERU

500 km

Figs. 441-442. Rhionaeschna dot maps.

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80 W 70 W 60 W 500 km 5 S

BRAZIL

PERU 15 S BOLIVIA

vigintipunctata obscura CHILE PARAGUAY

25 S

ARGENTINA

115 W 105 W 95 W 85 W 75 W 500 km USA

30 N

20 N MEXICO

dugesi jalapensis

GUATEMALA NICARAGUA 10 N

COSTA RICA PANAMA

Figs. 443-444. Rhionaeschna dot maps.

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135 W 125 W 115 W 105 W 95 W 85 W 75 W 65 W CANADA

40 N

30 N USA

multicolor mutata MEXICO

1000 km

0 95 W 65 W 50 W 35 W ECUADOR 1000 km

PERU

BRAZIL 15 S

BOLIVIA

CHILE PARAGUAY

30 S

ARGENTINA URUGUAY

45 S

absoluta

Figs. 445-446. Rhionaeschna dot maps.

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85 W 75 W PERU 65 W 55 W 45 W BOLIVIA BRAZIL

PARAGUAY CHILE 25 S

URUGUAY 35 S ARGENTINA

bonariensis diffinis

45 S

500 km

85 W 80 W 75 W 70 W 65 W 60 W

COLOMBIA

0 ECUADOR

5 S

BRAZIL

10 S

PERU

15 S elsia BOLIVIA galapagoensis

500 km CHILE

Figs. 447-448. Rhionaeschna dot maps.

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125W 115W 105 W

CANADA

45 N

USA californica

35 N

500 km MEXICO

70 W 60 W 50 W BOLIVIA 20 S BRAZIL

CHILE PARAGUAY

30 S

URUGUAY

confusa ARGENTINA brasiliensis

40 S

500 km

Figs. 449-450. Rhionaeschna dot maps.

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90 W 80 W 70 W 60 W

VENEZUELA

10 N

COLOMBIA

ECUADOR

PERU 0

BRAZIL

10 S

BOLIVIA marchali tinti

CHILE PARAGUAY 900 km ARGENTINA

80 W 70 W 60 W 50 W 40 W 30 W

PERU

BOLIVIA

20 S BRAZIL

PARAGUAY

CHILE

30 S

URUGUAY

ARGENTINA

40 S

peralta variegata

50 S

1000 km

Figs. 451-452. Rhionaeschna dot maps.

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85 W 70 W VENEZUELA 55 W 35 W

COLOMBIA

ECUADOR

5 S PERU

BRAZIL

BOLIVIA

20 S PARAGUAY CHILE planaltica haarupi pauloi

1000 km 35 S ARGENTINA URUGUAY

MEXICO 85 W 75 W 65 W 55 W

GUATEMALA NICARAGUA

10 N COSTA RICA PANAMA VENEZUELA

COLOMBIA

0 ECUADOR

PERU cornigera nubigena BRAZIL

10 S

1000 km BOLIVIA

Figs. 453-454. Rhionaeschna dot maps.

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90 W 105 W USA 75 W 60 W 45 W 30 N

MEXICO DOMINICAN REPUBLIC CUBA PUERTO RICO JAMAICA 15 N DOMINICA GUATEMALA NICARAGUA

COSTA RICA VENEZUELA PANAMA COLOMBIA

0

ECUADOR

psilus PERU BRAZIL

15 S BOLIVIA

PARAGUAY CHILE 1000 km ARGENTINA

113 W 108 W 103 W 98 W 29 N USA 300 km

MEXICO 24 N

manni vazquezae

19 N

Figs. 455-456. Rhionaeschna dot maps.

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81 W 76 W 71 W 56 W

VENEZUELA

5 N

COLOMBIA

0 ECUADOR

biliosa condor

5 S

PERU BRAZIL 200 km

79 W 74 W 69 W

VENEZUELA

6 N

COLOMBIA

1 N

demarmelsi ECUADOR joannisi

200 km

Figs. 457-458. Rhionaeschna dot maps.

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53 W 48 W 43 W

24 S

PARAGUAY BRAZIL

29 S

decessus eduardoi punctata

URUGUAY 300 km

140 W 110 W 80 W 50 W 20 W 50 N

20 N

10 S

40 S Marmaraeschna group Draco group

2000 km

Fig. 459. Rhionaeschna dot maps – 460. Rhionaeschna generalized tracks.

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90 W 60 W 30 W 50 N

20 N

10 S

40 S Schizuraeschna group

2000 km

140 W 110 W 80 W 50 W 20 W 50 N

20 N

10 S

40 S Neureclipa group

2000 km

Figs. 461-462. Rhionaeschna generalized tracks.

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140 W 110 W 80 W 50 W 20 W 50 N

20 N

10 S

40 S Variegata group

2000 km

140 W 110 W 80 W 50 W 20 W 50 N

20 N

10 S

40 S Cornigera group

2000 km

Figs. 463-464. Rhionaeschna generalized tracks.

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140 W 110 W 80 W 50 W 20 W 50 N

20 N

10 S

40 S Punctata group

2000 km

140 W 80 W 50 W 20 W 50 N

NAm

CAm

20 N

Te

10 S

An Pa

Au 40 S Areas of endemicity

2000 km

Fig. 465. Rhionaeschna generalized tracks – 466. Rhionaeschna areas of endemicity.

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North America Tepuis

Central America North America

Tepuis Central America

Paranense Paranense

Andes Andes

Austral South Austral South America America ab

0.05.0 10.0

North America

Central America

Tepuis

Paranense

Andes

Austral South America

Fig. 467. Area cladograms for species and monophyletic groups of Rhionaeschna – 468. Similarity analysis of endemicity ar- eas of Rhionaeschna species.

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