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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: 67207, figs. 1-468, tabs. 1-3. [ISSN 0040-7496]. Published 1 June 2003.
This study includes a revisionary, phylogenetic and biogeographical analysis of Neotropical components 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 including the type species Aeshna grandis Fabricius. In the present synopsis the generic name Rhionaeschna Förster is assigned to the New World group characterized by the presence of a conical tubercle 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 Cockerell, 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-Pacific 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; phylogeny; biogeography.
CONTENTS
Introduction .........................................................68
Material and methods
Phylogenetic analysis.............................................69
Biogeographic analysis...........................................70
Taxonomic analysis ...............................................70
Taxonomic part
Rhionaeschna Förster .............................................72
Keys
Keys to males of Rhionaeschna ...............................73
67
T  E,  146, 2003
Keys to females of Rhionaeschna ............................77
Rhionaeschna species accounts
1. R. draco group ...................................................80
Rhionaeschna draco (Rácenis).................................80
2. Marmaraeschna group .......................................81
Rhionaeschna brevicercia (Muzón & von Ellenrieder).
.......................................................................81
Rhionaeschna brevifrons (Hagen)............................82
Rhionaeschna fissifrons (Muzón & von Ellenrieder) 83
Rhionaeschna intricata (Martin) .............................84
Rhionaeschna obscura (Muzón & von Ellenrieder) .84
Rhionaeschna pallipes (Fraser) ................................85
Rhionaeschna vigintipunctata (Ris) .........................86
3. Schizuraeschna group.........................................86
Rhionaeschna dugesi (Calvert) ................................86
Rhionaeschna jalapensis (Williamson).....................87
Rhionaeschna multicolor (Hagen) ...........................88
Rhionaeschna mutata (Hagen)................................89
4. Neureclipa group ...............................................89
Rhionaeschna absoluta (Calvert) .............................89
Rhionaeschna bonariensis (Rambur)........................90
Rhionaeschna diffinis (Rambur)..............................91
Rhionaeschna elsia (Calvert) ...................................92
Rhionaeschna galapagoensis (Currie) .......................93
5. R. variegata group .............................................94
Rhionaeschna brasiliensis (von Ellenrieder & Martins
Costa) .............................................................94
Rhionaeschna californica (Calvert) ..........................94
Rhionaeschna confusa (Rambur) .............................95
Rhionaeschna marchali (Rambur)...........................96
Rhionaeschna peralta (Ris)......................................97
Rhionaeschna tinti (von Ellenrieder).......................99
Rhionaeschna variegata (Fabricius) .........................99
6. R. cornigera group ...........................................101
Rhionaeschna cornigera (Brauer)...........................101
Rhionaeschna haarupi (Ris) ..................................102
Rhionaeschna manni (Williamson & Williamson)103
Rhionaeschna nubigena (De Marmels)..................103
Rhionaeschna pauloi (Machado)...........................104
Rhionaeschna planaltica (Calvert).........................104
Rhionaeschna psilus (Calvert) ...............................106
Rhionaeschna vazquezae (Gonzalez) .....................106
7. R. punctata group ............................................107
Rhionaeschna biliosa (Kennedy ) ..........................107
Rhionaeschna condor (De Marmels) .....................108
Rhionaeschna decessus (Calvert) ............................108
Rhionaeschna demarmelsi von Ellenrieder sp. n. ...109
Rhionaeschna eduardoi (Machado) .......................110
Rhionaeschna joannisi (Martin) ............................110
Rhionaeschna punctata (Martin)...........................111
Phylogenetic analysis...........................................112
Biogeographic analysis.........................................113
Acknowledgements .............................................115
References ...........................................................115
Tables .................................................................121
Illustrations .........................................................124
68
INTRODUCTION
The genus Aeshna was described by Fabricius in
1775 to include four species: A. forcipata Linnaeus, A.
grandis Linnaeus, A. variegata Fabricius and A. clavata
Fabricius. Selys (1883) fixed A. juncea as the type
species of Aeshna, but as this species was not included
in the original description of the genus, its designation
is invalid according to the ICZN (1999). Latreille
(1810) mentioned A. forcipata as the type species of
Aeshna, but Aeshna forcipata of Latreille is a synonym
of Libellula vulgatissimus and not the same species as
A. forcipata Linnaeus, and thus it is also ineligible as
the type species of Aeshna. Therefore, the first valid
type designation is that of Cowley (1934), who designated Aeshna grandis as type species of Aeshna.
Aeshna was the first described genus of Aeshnidae,
and the current number of genera within this family
now stands at 50. Nineteen genera (40% of the total
number of currently accepted genera) have been created to include species originally described as Aeshna as
follows: Amphiaeschna Selys, 1871, Anaciaeschna
Selys, 1878, Andaeshna De Marmels, 1994, Basiaeschna Selys, 1883, Boyeria McLachlan, 1896, Caliaeschna Selys, 1883, Castoraeschna Calvert, 1952,
Coryphaeschna Williamson, 1903, Epiaeschna Hagen,
1877, Gomphaeschna Selys, 1871, Gynacantha Rambur, 1842, Hemianax Selys, 1883, Nasiaeschna Selys
in Förster, 1900, Neuraeschna Hagen, 1867, Oplonaeschna Selys, 1883, Planaeschna McLachlan, 1896,
Polycanthagyna Fraser, 1933, Remartinia Navás, 1911,
and Staurophlebia Brauer, 1865. No unique characters
define ‘Aeshna’, its species being usually identified as
those not belonging to other genera. Peters (1987) analyzed the European genera of Aeshnidae and concluded that Aeshna was not monophyletic, with ‘Aeshna’ affinis van der Linden, 1820 and A. mixta Latreille,
1805 more closely related to Anaciaeschna and Anax
than to the remaining European Aeshna species.
The most recent treatment of the Neotropical
species of ‘Aeshna’ was by Calvert (1956). His study
comprised species currently assigned to Andaeshna, Remartinia, Castoraeschna and Coryphaeschna. Calvert
(1956) included detailed redescriptions of all species
but no diagnoses were provided outside the keys. The
keys were based almost exclusively on thoracic color
pattern, which can be intraspecifically variable, and
several of his species accounts were composite descriptions of more than one species. For example, I found
that A. peralta included specimens of A. peralta, A. variegata and A. brasiliensis; A. variegata included A. variegata and A. marchali; A. cornigera included A. cornigera
and A. planaltica; A. planaltica included A. planaltica
and A. nubigena; A. vigintipunctata included A. vigintipunctata, A. obscura and A. pallipes; A. intricata included A. intricata, A. brevicercia, A. fissifrons, A. intri-
 E: Rhionaeschna synopsis
cata and A. vigintipunctata; A. brevifrons included A.
brevifrons and A. intricata; and A. diffinis included A.
absoluta and A. diffinis. Since Calvert’s monograph
(1956), 12 new species have been added to the 26
Neotropical ‘Aeshna’ species he included, bringing it to
a total of 38 described species. Proper identification of
the Neotropical species of this group has been difficult
not only because of problems enumerated above, but
also due to the paucity of specimens of several species
in the world’s collections; many species appear to be
rare or are seldom encountered. I treat the mostly
Neotropical components of Calvert’s genus ‘Aeshna’
cladistically as well as allowing for correct identification
of all its members. My results are based on an examination of over 3000 specimens, including primary
types or paratypes of 31 out of 39 distinguished
species. This study includes keys and diagnoses supported by extensive annotated illustrations, which
should facilitate identification of all species presently
included in the resurrected genus Rhionaeschna Förster.
MATERIAL AND METHODS
Phylogenetic analysis
The ingroup taxa included 68 species out of the 85
currently assigned to Aeshna, two species of Andaeshna
and two of Anaciaeschna. Polarity of characters was established with reference to the outgroup Oreaeschna
dictatrix Lieftinck (Oreaeschna constitutes the sister
group of the remaining Aeshninae according to von Ellenrieder 2002: fig. 437). Most parsimonious trees
were sought using branch-swapping search (mh*, bb*
command) in Hennig86 (version 1.5, Farris 1988).
Implicit enumeration commands were not used due to
the large number of taxa. Autapomorphies of terminal
taxa were excluded from the analysis. All question
marks in the matrix refer to missing data. Multistate
characters were treated as unordered. All characters
were initially weighted equally, and successive weighting was performed a posteriori. Cladograms were examined by manually plotting the proposed synapomorphies.
Character analysis
The lack of information on the preimaginal stages
of several species precludes using larval characters for
phylogenetic analyses; comparative study was therefore focused on adult characters. Thirty-nine external
morphological characters and associated character
states are listed below. The data matrix is shown in
table 2. Characters were chosen and scored from direct examination of voucher specimens.
Characters
Head
1. Frons transverse carina (frontal view): (0) linear or convex
(figs. 1, 7, 9-38c); (1) cleft (figs. 2-6, 8c)
2. Occipital triangle (B)/ interocular suture length (A): (0)
0.25-0.65 (fig. 415); (1) 0.7-1 (fig. 414); (2) 0.1-0.22
(fig. 416)
3. Margin between horizontal and vertical portions of frons
(lateral view): (0) angled (figs. 1, 9-38b); (1) rounded
(figs. 2-8b)
4. Frons (lateral view): (0) not flattened (figs. 1-2, 5-6, 838b); (1) flattened (figs. 3-4, 7b)
5. T-spot of frons (dorsal view): (0) well defined (figs. 1-39a;
414-416); (1) not defined
Thorax
6. Thoracic color pattern: (0) with pale stripes (figs. 48-58,
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 the extension of the stripes is variable being reduced or absent in some specimens (i.e. figs. 59, 75, 90);
these species were scored as (0).
7. Supratriangular crossveins: (0) present (figs. 396-398,
400-407); (1) absent (fig. 399). In species scored as lacking supratriangular crossveins, some specimens (about 115% of the examined specimens) possessed 1-2 crossveins
in 1-2 of the supratriangles.
8. Bending of MA: (0) slight (fig. 405, 407); (1) marked (figs.
396-404, 406)
9. Origin of two cell rows between RP1 and RP2: (0) proximal to pterostigma or under it (figs. 396-397, 401-405);
(1) distal to pterostigma or at its distal end (figs. 398-400,
406-407)
10. Course of RP2: (0) markedly bent (figs. 397-405, 407);
(1) kinked at distal end of pterostigma (fig. 406); (2)
evenly curved (fig. 396)
11. IRP2 fork: (0) asymmetrical with inferior branch as a
continuation of IRP2 and upper branch weak (fig. 405);
(1) symmetrical (figs. 396-404); (2) asymmetrical with
upper branch as a continuation of IRP2 and inferior
branch weak (fig. 406)
12. Cells of anal triangle: (0) 3 (exceptionally 4 in some
specimens) (figs. 396-404, 406-407); (1) 2 (fig. 405)
13. Anal angle: (0) angled (figs. 396-406, 408-412); (1)
rounded (fig. 407, 413)
14. Fusion of basal and distal veins of anal triangle: (0) parallel for a long distance (about 30% of anal margin) (figs.
408, 411); (1) parallel for a short distance (less than 25%
of anal margin) (figs. 409-410, 412); (2) at a single point
at angle (fig. 413)
15. Membranule length: (0) 50-66% of anal wing margin
(figs. 408-411); (1) 75-100% of anal wing margin (figs.
412-413)
Abdomen
16. Hamular anterior process in ventral view: (0) short (fig.
422); (1) long and triangular (fig. 423); (2) absent (figs.
424-425b,c)
17. Hamular posterior process in ventral view: (0) absent
(figs. 276-315b, 422-423); (1) tuberculate (fig. 424a,b);
(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)
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) triangular and wide: twice or less as wide at base (A) than at distal 30% (B) (figs. 317-324, 353-354); (2) cylindrical (fig.
423); (3) vestigial (figs. 347-348)
20. Genital lobe: (0) low (figs. 234-235, 237, 239, 241,
243-273, 418); (1) high and projected ventrally (figs.
236, 238, 240, 242, 274-275, 419); (2) very high and
projected postero-ventrally (fig. 421); (3) no higher than
margin of fossa (fig. 417)
21. Denticles on genital lobe: (0) absent (fig. 421); (1) on
margin (figs. 234-269, 417-418); (2) widespread (figs.
270-275, 419)
22. Auricle denticles: (0) 2 or 3 (figs. 195-233, 430-433);
(1) 4 to 5 (figs. 428-429)
23. Auricle shape (ventral view): (0) triangular or quadrangular (figs. 428-431); (1) narrow parallel-sided (figs. 432433)
24. Basal fold on lateral lobe of vesica spermalis IVth segment: (0) present (figs. 434, 436); (1) absent (fig. 435)
25. Medio-longitudinal fold on ventral lobe of vesica
spermalis IVth segment: (0) absent (figs. 434-435); (1)
present (fig. 436)
26. Light blue spot on male sterna IX (posterior to genital
opercula) and X: (0) absent (figs. 119-141b, 142c, 151156b); (1) present (figs. 143c-e, 144-150b)
27. Conical tubercle on abdominal sternum I: (0) absent (fig.
421); (1) with denticles widespread (figs. 234-249, 252275); (2) with denticles restricted to apex (figs. 250-251)
28. Transverse ridge on abdominal sternum I: (0) absent; (1)
present (fig. 417)
29. Constriction of abdominal segment III: (0) marked (figs.
119-120, 123, 125-156); (1) slight (figs. 121-122, 124)
30. Subbasal tooth or carina in dorsal surface of male X segment: (0) present (figs. 119-141b, 142c-d, 143-156b,
427a); (1) absent (fig. 426a)
31. Lateral carina of male cercus: (0) convex (figs. 390-395b,
426a); (1) linear (figs. 359-365); (2) concave (figs. 358,
366-389, 427a)
32. Subbasal tooth of male cerci: (0) absent (figs. 382-395,
426a); (1) present (small to prominent) (figs. 358-381,
427a)
33. Apical spine ventrally directed projected from inner
margin of cercus: (0) absent (fig. 427a); (1) present (figs.
426a-c)
34. Row of denticles on dorso-distal crest of cercus: (0) absent (fig. 426b); (1) present (Fig 427a-b)
35. Dorso-distal crest of cercus: (0) absent; (1) linear in dorsal view and evenly curved in lateral view (figs. 358-365,
370-383, 385-387-392, 394-395, 427a); (2) linear in dorsal view and angled in lateral view (figs. 366-369, 384,
388-389, 393); (3) curved in dorsal view (fig. 426c)
36. Distal concavity on cercus inner margin: (0) absent (fig.
427b); (1) present (fig. 426c)
37. Subapical ventral process of inner margin of male cerci:
(0) absent (figs. 358-365, 370-395, 426-427); (1) present
(figs. 366-369)
38. Spines of epiproct tip: (0) small and directed dorsally (fig.
427c); (1) large and directed anteriorly (fig. 426d)
39. Tip of cercus (lateral view): (0) in same plane with cercus
(figs. 358-365, 370-381); (1) extreme apex bent ventrally
(figs. 366, 382-389); (2) whole tip ventrally bent (figs.
367-369, 390-395).
70
Biogeographic analysis
Endemicity areas for Rhionaeschna were delimited
based on the distribution of the more restricted species
(fig. 466). An analysis was carried out to establish the
degree of similarity among areas of endemicity, using
Jaccard’s association coefficient and average linking. A
parsimony analysis of endemicity (PAE; Rosen 1988,
Cracraft 1991, Morrone 1994) was performed in
which areas (analogous to taxa) are classified by their
shared taxa (analogous to characters) according to the
most parsimonious cladogram. Data for PAE consist of
area 3 taxa matrices and the resulting cladograms represent nested sets of areas of endemicity (Morrone &
Crisci 1995). Cladistic information is then incorporated by adding supraspecific monophyletic groups to
the matrix. PAE recognizes generalized tracks through
the discovery of nested sets of areas. In order to undertake the PAE, species and species groups were coded for
their absence (0) or presence (1) in each area of endemism in the data matrix (table 3). The cladogram
obtained was rooted with a hypothetical area coded by
all zeros. Most parsimonious trees were sought using
implicit enumeration (ie* command) in Hennig86
(version 1.5, Farris 1988).
Taxonomic analysis
Each species account includes synonymy, diagnosis,
brief description, biology data, remarks, and distribution. Synonymies include only references to the taxon
under consideration that incorporate descriptions, redescriptions, illustrations or new distribution records.
Wing venation terms and abbreviations follow Riek &
Kukalová-Peck (1984), as amended by Bechly (1996).
The term ‘hamuli’ used throughout text refers to the
anterior pair of hamuli. Total length includes caudal
appendages, maximum width of the wings was taken
at the level of the nodus, and the length of the
pterostigma was measured from the anterior costal
end to the posterior anal end. All dimensions are in
millimeters. The range of measurements was taken
from 10 males and 10 females whenever available and
all specimens available of each species were examined
to establish the variability of characters. All diagnostic
characters were illustrated, including variation when
observed. Illustrations were made with the aid of a
camera lucida, and are not to scale.
The following acronyms are used under species accounts, he: head, th: pterothorax, ab: abdomen, te:
ventral terga, ge: genital fossa, tg: ventral tubercle and
genital lobe, ha: anterior hamuli, as: anterior lamina
spine, ce: cerci, wi: wings, au: auricle; ve: vesica
spermalis distal segment, and Mp: map, and for countries in the plates, AR: Argentina, BO: Bolivia, BR:
Brazil, CH: Chile, CO: Colombia, CR: Costa Rica, EC:
Ecuador, GU: Guatemala, MX: Mexico, PE: Peru, PR:
Puerto Rico, US: U.S.A.
Little is known of the biology of most species of this
genus; I have extracted notes from specimen labels and
comments from collectors on the behavior of certain
species. Under each species I included a tentative
ranking (rare; common) based on its occurrence and
abundance in collections and in the field. Full locality
data are given only for new or revalidated species; for
remaining species, country, state, and repository have
been indicated in the distribution lists. Full locality
data for those species are available from the author.
Asterisks in distribution lists indicate material that has
been illustrated. Maps represent distribution records
from collections and reliable references. Distribution
data for some common and well-known species in the
U.S.A. (e.g. R. californica, R. dugesi, R. multicolor and
R. mutata) are not exhaustive. Maps were created electronically from the Digital Chart of the World
(1:1,000,000) using ArcView 3.1 (ESRI 1996). Elevation data and longitude/latitude coordinates were
culled from the Global Gazetteer website (http://
www.calle.com/world/) and placed into a Microsoft
FoxPro Data base linked to the ArcView program.
Acronyms used for collections are those given in
the website ‘The insect and spider collections of the
world http://hbs.bishopmuseum.org/codens/codensr-us.html and codens not included there are as follows: ABMMC – Dr. A. B. M. Machado personal collection, Belo Horizonte, Brazil; GSVC – Dr. Graham
S. Vick personal collection, Hans, U.K.; IADIZA – Instituto de Investigaciones de Zonas Áridas, Mendoza,
Argentina; IEUM – Instituto de Entomología, Universidad Metropolitana de Ciencias de la Educación,
Santiago, Chile; KJTC – Dr. Kenneth J. Tennessen
personal collection, Florence, U.S.A.; WBC – Dr. Wilfried Braun personal collection, Berlin, Germany.
Characters
Discrimination among species of this group is difficult since combinations of characters must be used
to identify most species. The following characters are
useful in evaluating specific status:
Head (figs. 1-38). – Contour of frons in lateral view
dorsally projected (fig. 1b) or not (i.e. fig. 2b); frontal
carina linear to convex (i.e. fig. 7c) or with a median
concavity in frontal view (i.e. figs. 2, 4c), anteriorly
angled (fig. 18a) or smoothly curved in dorsal view
(i.e. fig. 19a), and located at anterior margin of frons
(i.e. fig. 8b) or recessed into horizontal portion of
frons (i.e. fig. 9b); clypeal lobes angled (i.e. fig. 14c) or
rounded (i.e. fig. 16c); frons and vertex shorter (i.e.
fig. 24a) or longer (i.e. fig. 22a) than interocular suture and occipital triangle; T-spot of frons complete
(i.e. fig. 2a) or incomplete (i.e. fig. 4a), sides of stem
wide (i.e. fig. 34a) or narrow (i.e. fig. 33a), parallel (i.e.
fig. 14a) or converging anteriorly (i.e. fig. 13a); frontoclypeal groove with (i.e. fig. 17b) or without (i.e. fig.
16b) a dark stripe, and fronto-ocular groove with a
dark stripe which may be of uniform width (i.e. fig.
35b) or widened toward point of contact between
frontoclypeal groove and eyes (i.e. fig. 37b).
Thorax (figs. 39-118). – Extension and contour of
pale mesanepisternal, mesepimeral, and metepimeral
stripes show intraspecific as well as interspecific variation, but can be used as diagnostic characters for some
species (i.e. figs. 92, 95). Color of wing membranule
can also aid in identification (i.e. dark except basal
20% pale in R. bonariensis, pale except distal 20-33%
dark in R. absoluta). Wing venation provides no specific characters (table 1), but presence (i.e. fig. 396) or
absence (fig. 399) of crossveins in supratriangles, and
level of beginning of two rows of cells between RP1
and RP2 with respect to pterostigma (i.e. proximal, fig.
402 vs. distal, fig. 399), are good characters for groups
of species.
Abdomen (figs. 119-156). – Presence or absence
and confluence or separation of certain pale spots are
sometimes diagnostic (i.e. figs. 133, 140), as well as
presence or absence of pale or dark medio-longitudinal stripes on female dorsum of segment II (i.e. figs.
131-132). Useful structural characters of the secondary genitalia include: anterior margin of ventral
tubercle of segment II concave (i.e. fig. 267), linear
(i.e. figs. 264-265), or convex (i.e. figs. 261-262); genital lobe low (i.e. fig. 237) or high (i.e. fig. 240), with
denticles on margin (i.e. fig. 268) or spread over lateral surface as in punctata group (figs. 270-275); anterior lamina spine wide (i.e. fig. 317) or narrow (i.e. fig.
325), short (i.e. fig. 354) or long (i.e. fig. 350), triangular (i.e. fig. 355) or needle-shaped (figs. 356-357),
with dorsal margin linear (i.e. fig. 330) or concave (i.e.
fig. 332), tip directed dorsally (i.e. fig. 326) or ventrally (i.e. fig. 325); anterior hamuli with long anterior
process in Marmaraeschna group (i.e. figs. 277-284),
short in remaining species (i.e. fig. 288), with pointed
(i.e. fig. 311) or rounded tip (i.e. fig. 312), dorsal fold
short (i.e. fig. 312) or long (i.e. fig. 311). Ventral terga (delimited by inner and outer latero-ventral longitudinal and postero-ventral transverse carinae) can be
narrow (i.e. fig. 181) or wide (i.e. fig. 180), and lateral carinae can be linear (i.e. fig. 191) or sinuous (i.e.
fig. 192). Male cerci (figs. 358-395) with dorso-distal
crest low (i.e. fig. 384) or high (i.e. fig. 382), short (i.e.
fig. 364) or long (i.e. fig. 365), smoothly curved (i.e.
fig. 393) or angled (i.e. fig. 392); sub-basal tooth absent (i.e. fig. 382) or present (i.e. fig. 380), low (i.e. fig.
363) or prominent (i.e. fig. 364); lateral margin linear
(i.e. fig. 359), concave (i.e. fig. 358) or convex (i.e. fig.
390) in lateral view; tips pointed (i.e. fig. 391) or blunt
(i.e. fig. 393), ventrally bent (i.e. figs. 367, 385, 390)
or in the same plane as cerci (i.e. fig. 359); inner mar71
T  E,  146, 2003
gin with (figs. 366-369) or without (i.e. fig. 370) a
ventro-distal process. Female cerci tips angled (i.e. fig.
368) or rounded (i.e. fig. 366), with a long apical
spine (i.e. fig. 358), mucronate (i.e. fig. 364), incised
(i.e. fig. 395), or lacking such structures (i.e. fig. 365);
sides parallel (i.e. fig. 365), diverging (i.e. fig. 364), or
converging (i.e. fig. 383) distally.
TAXONOMIC PART
Rhionaeschna Förster
Rhionaeschna Förster, 1909: 220-223 (diagnosis, R. maita
only species included); Martin 1911: 13 (generic diagnosis from Förster); Ris 1913: 83 (indication of close relationship of maita with variegata and multicolor groups);
Hincks 1934: 80 (Peru): Schmidt 1952: 256 (Peru).
Neureclipa Navás, 1911: 476, 478 (brief description of
genus). Syn. n.
Aeshna (Hesperaeschna) Cockerell, 1913: 581 (brief description of subgenus in key; type species Aeshna (Hesperaeschna) californica); Calvert 1952: 253-255 (addition of
subgeneric characters and species); Calvert, 1956: 10 (in
key), 21-22 (subgeneric diagnosis), 217-218 (discussion
of origin), 222-224 (discussion of relationships).
Aeshna (Marmaraeschna) Calvert, 1952: 256 (characters of
subgenus, intricata designated as subgeneric type);
Calvert, 1956: 10 (in key), 109-110 (subgeneric diagnosis), 217-218 (discussion of origin), 222-224 (discussion
of relationships).
Aeshna (Schizuraeschna) Calvert, 1952: 256-257 (characters
of subgenus, multicolor designated as subgeneric type);
Calvert, 1956: 10 (in key), 100-102 (subgeneric diagnosis), 217-218 (discussion of origin), 222-224 (discussion
of relationships).
Aeshna (Neureclipa): Calvert, 1952: 257-258 (relegated to
subgeneric rank, bonariensis designated as type species, addition of subgeneric characters); Calvert 1956: 10 (in key),
125-126 (subgeneric diagnosis), 217-218 (discussion of
origin), 222-224 (discussion of relationships).
Aeshna (Rhionaeschna): Calvert, 1956: 11 (in key), 98-99
(relegated to subgeneric rank, subgeneric diagnosis), 217218 (discussion of origin), 222-224 (discussion of relationships).
Type species. – Rhionaeschna maita Förster, 1909 (=R. brevifrons (Hagen 1861)) by monotypy.
Diagnosis
The autapomorphy characterizing Rhionaeschna is
the presence of a conical tubercle bearing denticles on
abdominal sternum I (figs. 234-275, 418-419); sternum I is planar in Oreaeschna, Aeshna sensu stricto,
several ‘Aeshna’ species of uncertain position ‘A.’ affinis, ‘A.’ mixta, ‘A.’ brevistyla Rambur, 1842, ‘A.’ ellioti
Kirby, 1896, ‘A.’ williamsoniana Calvert, 1905, ‘A.’
moori Pinhey, 1981, ‘A.’ rileyi Calvert, 1892 and ‘A.’
subpupillata McLachlan, 1896, and Andaeshna, and
has a transverse ridge in ‘Aeshna’ isoceles Müller, 1767
and Anaciaeschna (fig. 417). Other characters that differentiate it from related Aeshnidae genera are: T-spot
of frons present (absent in Andaeshna and ‘A.’ isoceles);
72
occipital triangle of moderate length, measured as triangle length/interocular suture length: 0.25-0.65 (fig.
415) (shorter in Anaciaeschna; fig. 416); bend of MA
prominent (figs. 396-404, 406) (slight in Oreaeschna,
Aeshna s.str. and ‘A.’ isoceles; figs. 405, 407); IRP2 fork
symmetrical (figs. 396-404) (asymmetrical with inferior branch as a continuation of IRP2 stem in Oreaeschna, Aeshna s.str., Andaeshna, fig. 405; asymmetrical with upper branch as a continuation of IRP2 stem
in Anaciaeschna, fig. 406); anal triangle with three cells
(figs. 396-404, 406-407) (two in Aeshna s.str. except
for constricta group, fig. 405); anal angle angled (figs.
396-406, 408-412) (rounded in Andaeshna and ‘A.’
isoceles, 407, 413), with basal and distal veins parallel
and fused for a distance before angle (figs. 408-412)
(fused at a single point in Andaeshna and ‘A.’ isoceles,
fig. 413); membranule length 50-66% of wing margin
(figs. 408-411) (75-100% of wing margin in Anaciaeschna, Andaeshna and ‘A.’ isoceles, figs. 412-413);
hamular fold separated from wall of genital fossa (figs.
423-425) (not differentiated from wall of genital fossa
in Andaeshna and constricta group of Aeshna s.str., fig.
422). Genital lobe elevated above rest of ventral margin of tergum I (figs. 418-419, 421) (at same level as
ventral margin of tergum I in Anaciaeschna, fig. 417);
auricle triangular or quadrangular (figs. 428-431)
(narrow parallel-sided in Anaciaeschna and ‘A.’ isoceles,
figs. 432-433), with two or three denticles (usually
four or five denticles in Aeshna s.str.); basal fold of posterior lobes of vesica spermalis IVth segment present
(figs. 434, 436) (absent in Andaeshna and ‘A.’ isoceles,
fig. 435); and dorso-distal crest of cercus present (figs.
358-395) (absent in Oreaeschna, ‘A.’ affinis, ‘A.’ mixta,
‘A.’ brevistyla, and ‘A.’ ellioti, Andaeshna, Anaciaeschna, and ‘A.’ isoceles).
The remaining Aeshninae genera, Castoraeschna,
Coryphaeschna and Remartinia, can be easily distinguished from Rhionaeschna by the elongated discal triangles (width/length 0.35 or less) (shorter in Rhionaeschna, width/length 0.40 or more), male anal
triangle with two cells, and distal segment of vesica
spermalis lacking lateral and ventral folds and bearing
patches of michrotrichiae. Coryphaeschna and Remartinia have a smooth abdominal sternum I, while Castoraeschna has a cylindrical tubercle lacking denticles
accompanied in the male by a pair of medio-posterior
projections of the ventral tergum I (fig. 420).
Remarks
The oldest available name for the monophyletic
group comprising the Neotropical species of ‘Aeshna’
is Rhionaeschna Förster, 1909, in spite of the different
original definition of the genus. Rhionaeschna (from
Greek rhion = a jutting part of a mountain, peak,
promontory, headland) was erected by Förster (1909)
to include R. maita, and its distinction from Aeshna
was based in following structural characters: frons
wide (ratio head/frons width 1.5), dorsal margin of
frons rounded, and abdomen slightly constricted at
segment III. However, the ratio head/frons width is intraspecifically variable, and a wide frons is found
among several species of Rhionaeschna (i.e. R. brevifrons: 1.5-1.9 mm, R. fissifrons: 1.6-2.1 mm, R. intricata: 1.7-2.4 mm, R. pallipes: 1.7-2.5 mm, R. confusa:
1.6-2.5 mm, R. marchali: 1.5-1.9 mm, R. peralta: 1.71.85 mm, R. tinti: 1.7-2.2 mm, R. variegata: 1.6-2.3
mm, R. absoluta: 1.6-2.3 mm, R. bonariensis: 1.7-2.4
mm, R. diffinis: 1.4-2.3 mm, R. elsia: 1.7-2.2 mm). A
rounded frons is characteristic of several species (R.
brevicercia, R. brevifrons, R. fissifrons, R. intricata, R.
obscura, R. pallipes, and R. vigintipunctata) and the
slightly constricted abdomen is found in R. brevifrons,
R. fissifrons and R. pallipes. My examination of the lectotype of R. maita demonstrated that Rhionaeschna
maita Förster, 1909 is a junior subjective synonym of
R. brevifrons (Hagen, 1861).
Four other names, one originally for a genus and
three for subgenera, have been assigned to some of the
species which I presently include in Rhionaeschna.
Resolution of these names is given as follows.
Neureclipa was originally created by Navás (1911)
to include Aeshna bonariensis Rambur, 1842 and A.
litigatrix Navás, 1911 and was characterized by the absence of cross veins in the supratriangle, abdominal
segment X of male with a dorsal tooth, and male cerci
bearing a subbasal tooth. The subgenus Hesperaeschna
was created by Cockerell (1913) to include A. californica Calvert, 1905. The diagnostic characters included
in his key were: RP3-4 and MA separated by one cell
only at wing margin, but a short distance before by
two rows of cells, owing to the deflection of MA from
the straight course; cell formula of triangles 2, 1, 1; upper branch of radial sector in a line with stem; IRP2
separated from supplementary vein below by only
three rows of cells; IRP2 fork a short distance before
level of beginning of pterostigma.
Calvert (1952) relegated Neureclipa to subgeneric
rank, fixed A. bonariensis as its type including besides
Aeshna bonariensis, A. diffinis, A. elsia and A. galapagoensis, and added as subgeneric characters abdominal
segment I with a ventral tubercle, vein IRP1 beginning
at distal end of pterostigma, thorax with two lateral
oblique pale stripes, discoidal triangles with the formula 1, 1, 1, inner triangles usually with one cell, anal
triangle with three cells, and male cerci in lateral view
not bifid nor with an anteapical point. He also listed
more characters to differentiate Hesperaeschna species
from the European species of Aeshna (presence of a
ventral tubercle in the first abdominal segment; supratriangular crossveins; thorax dark with two pale lateral
stripes; male cerci in lateral view not bifid nor with an
anteapical ventral tubercle; dorso-medial carina in ab-
dominal segment X, and anal triangle with three cells).
Finally Calvert (1952) erected another two subgenera for Aeshna: Marmaraeschna and Schizuraeschna.
Marmaraeschna, with M. intricata as generotype, was
diagnosed by the pterothoracic marbled color pattern,
abdominal segment I with a ventral tubercle, supratriangular cross veins present, vein IRP1 beginning proximal to the level of the pterostigma or under the proximal end or proximal half of the pterostigma, anal
triangle 3-celled, male with a middorsal longitudinal
carina on segment X and male cerci with the apex not
bifid nor with an anteapical ventral tubercle. Schizuraeschna, including A. multicolor (generotype), A. mutata, A. dugesi and A. jalapensis, was characterized by
abdominal segment I with a ventral tubercle, supratriangular cross veins present, thorax dark with two lateral oblique pale stripes, vein IRP1 beginning distal to the
level of the pterostigma or under the distal end, discoidal triangles with the formula 2, 1, 1, inner triangles
two celled, anal triangle with three cells, and male cerci in lateral view bifid in the apical fourth or less, the
lower division much shorter than the upper and in one
species reduced to an anteapical point.
In the present synopsis, some of these names are
used in an informal sense to make reference to certain
species-groups, but I do not recognize subgenera.
Keys
Comparison with illustrations and diagnosis is advised to aid in identifications especially when trying to
identify poorly preserved or incomplete material (i.e.
specimens with bad color preservation, with broken
cerci or lacking head).
Key to males of Rhionaeschna
1. Pterothorax and abdominal segments VII-X without pale markings (figs. 39, 119a-b); hamulus
with medial digit-like projection on antero-ventral margin (fig. 276); frons projected dorsally,
higher than vertex in lateral view (fig. 1b);
Venezuelan tepuis (fig. 439)........................draco
– Pterothorax with or without pale stripes, abdominal segments IV-VIII to X with pale spots (figs. 41118, 120-156a-b); hamulus with no medial projection on antero-ventral margin (figs. 277-315);
frons not projected dorsally, lower than vertex
(figs. 2-38b) ......................................................2
2. Pterothorax with marbled pattern of black markings and pale areas (figs. 40-47); hamular anterior process long: distance between ventral tip of
process and axis of hamulus (A) longer than distance between dorsal end of hamular fold and
axis (B) (figs. 277-284) .........................Key M-1
– Pterothorax uniform in color or with pale stripes
73
T  E,  146, 2003
3.
–
4.
–
5.
–
6.
–
or spots (figs. 48-118); hamular anterior process
short: distance between ventral tip of process and
axis of hamulus (A) shorter than distance between
dorsal end of hamular fold and axis (B) (figs. 285315)..................................................................3
Sterna X and IX posterior to genital opercula with
bright light-blue spot (figs. 143-150b); surface of
abdominal ventral terga delimited by inner and
outer carinae narrow (width measured at narrowest point/length of ventral terga IV less than 0.1
except for R. vazquezae (fig. 188) and R. pauloi
(fig. 185) with widths of 0.13 mm and 0.18 mm,
respectively) (figs. 181-188); HW with two rows
of cells between RP1 and RP2 beginning under
pterostigma or proximal to it (fig. 401-402); subbasal tooth of cercus absent and extreme tip bent
ventrally (figs. 382-389b)......................Key M-2
Sterna of X and IX posterior to genital opercula
yellow or pale brown to black (figs. 127-142, 151156b); surface delimited by inner and outer carinae of abdominal ventral terga wide (width measured at narrowest point/ length of ventral terga IV
larger than 0.13) (figs. 165-180, 189-194); HW
with origin of two rows of cells between RP1 and
RP2 variable; cercus with other combination of
characters (figs. 366-381, 390-395b) .................4
Inner margin of cercus with a ventral process at
distal 25%, cercus with dorso-distal crest triangular in lateral view, sub-basal tooth low and blunt
(figs. 366-369) .....................................Key M-3
Cercus without ventral process on inner margin,
dorso-distal crest and sub-basal tooth variable
(figs. 370-381, 390-395)...................................5
Cercus with entire tip ventrally bent in lateral
view; sub-basal tooth absent; cercus in dorsal
view gradually widening to distal end (maximum
width at distal 12-16%) (figs. 390-395), denticles
of genital lobe small, numerous, and spread over
ventral and lateral surfaces of lobe (figs. 270-275)
.............................................................Key M-4
Cercus tip not ventrally bent, sub-basal tooth
present (but vestigial in R. californica and R. confusa); cercus in dorsal view variable (figs. 370381); denticles of genital lobe confined to ventral
surface of lobe (figs. 247-258)...........................6
Supratriangles usually free, sometimes one or two
supratriangles with 1-2 cross veins (in 2-10% of
specimens examined) (fig. 399).............Key M-5
Supratriangles with 1-3 cross veins, exceptionally
one supratriangle without cross veins (in 1% of R.
brasiliensis and R. variegata specimens examined)
(fig. 400)...............................................Key M-6
Key M1
1. Cerci short: 3.5-4.8 mm (figs. 360-362); constriction of abdominal segment III slight (figs.
121-122, 124)...................................................2
74
–
2.
–
3.
–
4.
–
5.
–
6.
–
Cerci long: 4.9-6.5 mm (figs. 359, 363-365);
constriction of abdominal segment III marked
(figs. 120, 123, 125-126) ..................................4
Frontal carina linear in frontal view (fig. 7c); Argentina (Buenos Aires and Mendoza north to
Salta province) (fig. 441)..........................pallipes
Frontal carina concave in frontal view (figs. 3-4c)
.........................................................................3
Frontal carina with a slight medial concavity (fig.
3c); inner and outer carina of ventral terga VI sinuous (fig. 159a-b); black spots of pterothorax reduced to 1 on mesanepisternum, 3 on mesepimeron-metepisternum, and 1 on metepimeron
(fig. 41-42); cercus with sub-basal tooth low (figs.
360b, e-f); Valdivia province in Chile north to
Peru (fig. 440) ......................................brevifrons
Frontal carina with a deep medial cleft (fig. 4c);
inner and outer carina of ventral terga VI linear,
parallel (fig. 160a); black spots of pterothorax: 2-3
on mesanepisternum, 4-5 on mesepimeron-metepisternum, and 3 on metepimeron (fig. 43); cercus with sub-basal tooth prominent (fig. 361b);
northwestern Argentina north to Peru (fig. 440)..
...............................................................fissifrons
Sub-basal tooth of cercus low (figs. 359B, 363b,
d); abdominal segment I with black spot extending
to tip of auricles in ventral view (figs. 196, 199);
smaller species (total length 66-73.5 mm)...........5
Sub-basal tooth of cercus prominent (figs. 364365b); black spot not extending to tip of auricles
in ventral view (figs. 200, 202); larger species (total length 72-80.5 mm).....................................6
Stem of T-spot strongly narrowed at its confluence with transverse arms (fig. 5a); ventral tubercle of abdominal segment I high (about as high as
50% of its length in lateral view) (fig. 238); Peru
to Ecuador (fig. 442)..............................intricata
Stem of T-spot widely confluent with transverse
arms (fig. 2a); ventral tubercle of abdominal segment I low (about as high as 30% of its length in
lateral view) (fig. 236); Ecuador to Venezuela
(fig. 442).............................................brevicercia
Stem of T-spot strongly narrowed at its confluence with transverse stem, vertex yellow with posterior 30% black (fig. 8a); genital lobe approximately triangular in lateral view (fig. 242);
dorso-distal crest of cercus extended along distal
30% of cercus (fig. 365b); northwestern Argentina to Peru (fig. 443).....................vigintipunctata
Stem of T-spot widely confluent with transverse
stem, vertex black with a small rounded yellow
spot at each anterolateral edge (fig. 6a); genital
lobe subquadrate in lateral view (fig. 240); dorsodistal crest of cercus extended along distal 25% of
cercus (fig. 364b); Bolivia north to Peru (fig.
443).........................................................obscura
Key M-2
1. Dorso-distal crest of cercus rising gradually, angulate and lower than base of cercus in lateral
view (figs. 384, 388, 389b) ...............................2
– Dorso-distal crest of cercus rising abruptly,
smoothly convex and higher than base of cercus
in lateral view (figs. 382-383, 385-387b) ..........4
2. Anterior lamina spine well developed (fig. 349);
pale mes- and metepimeral stripes of thorax wide
with anterior (upper) portion as wide as 50-66%
of their respective sclerites (figs. 107-109); dark
stripe on frontoclypeal groove (figs. 32b-c);
Guerrero and Nayarit states in Mexico (fig. 456)
............................................................vazquezae
– Spine of anterior lamina reduced (figs. 347-348);
pale mes- and metepimeral stripes relatively narrow, as wide at anterior portion as 30-50% of
their respective sclerites (figs. 97-99, 106); no
dark stripe on frontoclypeal groove (figs. 27, 32bc) ......................................................................3
3. Inner and outer margins of cercus approximately
parallel to each other along medial 30% (fig.
388a); pale thoracic stripes bright green,
mesanepisternal stripe complete (fig. 106); northern Argentina to southern USA (fig. 455).......psilus
– Inner and outer margins of cercus diverging gradually along medial 30% (fig. 384a); pale thoracic
stripes light blue, mesanepisternal stripe incomplete, as long as 50% of mesanepisternum (figs.
97-99); Baja California state in Mexico (fig. 456)
.................................................................manni
4. Pale mesepimeral and metepimeral stripes with
deep rounded indentations (figs. 95-96, 103105); abdominal PL spots large (figs. 144b, 147148b)................................................................5
– Pale mesepimeral and metepimeral stripes lacking
indentations (figs. 92-94) or with broad and shallow indentations (figs. 101-102); abdominal PL
spots small to absent (figs. 143a-c, 146b) ..........7
5. Ventral tubercle of abdominal segment I lower
than 30% of its length (fig. 266); cercus in dorsal
view with tip directed posteriorly, distal 60% of
cercus parallel-sided (fig. 387a); Paraná and Minas Gerais states in Brazil (fig. 453) ...........pauloi
– Ventral tubercle of abdominal segment I about as
high as 50% of its length (figs. 259, 264-265);
cercus variable (figs. 383, 386a) ........................6
6. Stem of T-spot of frons gradually narrowed distally (fig. 30); pale mesanepisternal stripes complete
(figs. 103-105), as wide as 50% of mesanepisternum; Argentina north to Venezuela (fig.
453) .....................................................planaltica
– Stem of T-spot of frons abruptly narrowed distally (fig. 26); pale mesanepisternal stripes complete
(fig. 95) or incomplete (fig. 96), as wide as 2530% of mesanepisternum; northwestern Argenti-
na (fig. 453) ............................................haarupi
7. Pale mesepimeral stripe narrowed at midlength to
50% of its basal width (figs. 101-102); anterior
side of ventral tubercle concave (fig. 267); dorsodistal crest of cercus gradually rising at base (fig.
385b); Venezuelan tepuis (fig. 454) .......nubigena
– Pale mesepimeral stripe approximately uniform
in width (figs. 92-94); anterior and posterior
sides of ventral tubercle convex (figs. 261-262);
dorso-distal crest of cercus abruptly rising at base
(fig. 382b, g); Bolivia north to Mexico (fig. 454)
..............................................................cornigera
Key M-3
1. Ventral process of inner margin of cercus small,
not projected posteriorly (fig. 366b); tip of anterior lamina spine directed ventrally (fig. 325);
Mexico to southern USA (fig. 444)..............dugesi
– Ventral process of inner margin of cercus projected posteriorly (figs. 367-369b); tip of anterior lamina spine curved dorsally (figs. 326-328).............2
2. No pale abdominal PL spots in segments IV-V, PD
spots small (fig. 128a-b); only extreme tip of cercus curved ventrally (fig. 367c); Costa Rica to
Mexico (fig. 444) .................................jalapensis
– Pale abdominal PL spots present in segments IV-V
and confluent with large PD spots (figs. 129130a-b); entire tip of cercus curved ventrally
(figs. 368-369c) ................................................3
3. Distance between posterior base of ventral process
and base of dorsal crest of cercus (A) larger than
distance between it and tip of cercus (B) (fig.
369c); ventral tubercle of abdominal segment I
low (about as wide as 3 times its height) (fig. 245);
base of T-spot stem wider than vertex; vertex black
with anterior 30% pale (fig. 12a); eastern USA to
southern Canada (fig. 445).........................mutata
– Distance between posterior base of ventral
process and base of dorsal crest (A) shorter than
distance between it and tip of cercus (B) (fig.
high (about as wide as 2 times its height) (fig.
244); base of T-spot stem narrower than vertex,
vertex pale with posterior 30% black (fig. 11a);
Mexico to southern Canada through western USA
(fig. 445) .............................................multicolor
Key M-4
1. Black stripe on frontoclypeal groove strongly
widened toward eyes (figs. 37-38); pale pterothoracic stripes complete, linear, and narrower than
50% of their sclerites (114, 118); genital lobe
prominent, higher than 30% of its width in lateral view (figs. 274-275); tip of hamular anterior
process blunt in ventral view (unknown in decessus) (figs. 314-315)............................................2
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–
2.
–
3.
–
4.
–
5.
–
6.
Black stripe on frontoclypeal groove not strongly
widened toward eyes or absent (figs. 33-36); pale
pterothoracic stripes with deep indentations, divided into spots, or wider than 50% of their sclerites (figs. 110-113, 115-117); genital lobe lower
than 30% of its width in lateral view (figs. 270273); tip of hamular anterior process pointed in
ventral view (figs. 310-313) ..............................4
Anterior lamina spine absent; labrum, clypeus
and frons orange; pale abdominal PL spots present
on segments IV-IX, fussed with ML on IV-VIII;
known only from Itatiaia, Rio de Janeiro state,
Brazil (fig. 459) (characters abstracted from
Calvert 1953) ..........................................decessus
Anterior lamina spine present (figs. 355-357);
labrum, clypeus and frons light blue or yellow;
pale abdominal PL spots absent on segments VIVII, separated from ML on IV-V, fused or not with
ML on VIII (figs. 155-156b) ...............................3
Ventral tubercle of abdominal segment I low and
smoothly curved in lateral view (fig. 275); black
of T-spot not extended on antefrons; vertex yellow with lateral edges black (fig. 38); São Paulo,
Rio de Janeiro, Espirito Santo and Minas Gerais
states in Brazil (fig. 459) ........................punctata
Ventral tubercle of abdominal segment I high
and trapezoidal in lateral view (fig. 274); black of
T-spot extended on dorsal surface of antefrons;
vertex yellow with lateral and posterior 30%
black (fig. 37); Minas Gerais state in Brazil (fig.
459).......................................................eduardoi
Dorso-distal crest of cercus smoothly convex in
lateral view (fig. 393b); anterior lamina spine as
long as 30% of its basal width (fig. 353-354);
ventral portion of hamuli (A) shorter than hamular fold (B) (fig. 311a); Táchira State in
Venezuela (fig. 458) ...........................demarmelsi
Dorso-distal crest of cercus angulate in lateral
view (figs. 390-392b); anterior lamina spine as
long as 2 times or more its basal width (figs. 350352); ventral portion of hamuli (A) longer than
hamular fold (B) (figs. 310, 312-313) ...............5
Transverse arms of T-spot represented by a fine
line (fig. 33a); labrum orange, clypeus and frons
bright yellow, no dark stripe on frontoclypeal
groove (figs. 33b-c); auricle bearing two teeth
(fig. 228); Peru to Ecuador (fig. 457) ........biliosa
Transverse arms of T-spot wide (figs. 34, 36a);
labrum pale brown, clypeus and frons light blue
to light green, dark stripe on frontoclypeal groove
(figs. 34, 36b-c); auricle bearing three teeth (figs.
229, 231)..........................................................6
Stem of T-spot wider than vertex (fig. 34a); pale
pterothoracic stripes as wide as 50-66% of their
sclerites (fig. 112); anterior tip of hamular process
in posterior view recurved (fig. 313); Táchira
76
–
state in Venezuela (fig. 457) ......................condor
Stem of T-spot narrower than vertex (fig. 36a);
pale mesepimeral and metepimeral stripes as wide
as 25-33% of their sclerites with an anterior and
a posterior indentation (figs. 115), or divided
into spots (figs. 116-117); anterior tip of hamular
process in posterior view not recurved (fig. 312);
Bolivia to Colombia (fig. 458) ................joannisi
Key M-5
1. Clypeal lobes rounded (figs. 16-17c); dorsal margin of anterior lamina spine concave (figs. 332333); ventral tubercle of abdominal segment I
with a few denticles restricted to its apex (fig.
250-251)...........................................................2
– Clypeal lobes angled (figs. 13-15c); dorsal margin
of anterior lamina spine linear (figs. 329-331);
ventral tubercle of segment I with numerous denticles spread over its posterior surface (fig. 247249)..................................................................3
2. No dark stripe on frontoclypeal groove (fig. 16bc); dorso-distal crest at distal 25% of cercus
abruptly rising at its base (fig. 373b); tip of
hamular anterior process pointed in ventral view
(fig. 292b); smaller species (total length 49-54
mm, male cerci 4.3-4.6 mm); central Chile to
Ecuador (fig. 448) ........................................elsia
– Dark stripe on frontoclypeal groove (fig. 17b-c);
dorso-distal crest at distal 30% of cercus gradually rising at its base (fig. 374b); tip of hamular anterior process rounded in ventral view (fig. 293b);
larger species (total length 58-60 mm, male cerci
5 mm); Galápagos Islands in Ecuador (fig. 448) .
........................................................galapagoensis
3. Black stripe on fronto-ocular groove narrow and
uniform in width, frontoclypeal groove not bordered by black stripe (fig. 14b-c); outer base of
cercus with a pale spot (fig. 371a-b); membranule
dark except basal 20% pale; stem of T-spot of
frons approximately parallel-sided (fig. 14a); Argentina to Brazil (fig. 447)..................bonariensis
– Black stripe on fronto-ocular groove widening towards frontoclypeal groove, frontoclypeal groove
bordered by black stripe (figs. 13, 15a); membranule pale except distal 20-33% dark; cercus
without pale spots (figs. 370, 372a-b); stem of Tspot of frons gradually widened basally (figs. 13,
15a) ..................................................................4
4. Ventral tergum IV of abdomen relatively narrow,
ratio length/maximum width of abdominal terga
IV: 4.2-4.6; basal 30% of outer lateral carinae of
segment IV linear to concave (fig. 169a); southern
Argentina to Ecuador (fig. 446) ..............absoluta
– Ventral tergum IV of abdomen relatively wide, ratio length/maximum width of abdominal terga
IV: 2.4-2.6, basal 30% of outer lateral carina of
segment IV convex (fig. 171a); southern and central Chile and southwestern Argentina (fig. 447).
.................................................................diffinis
Key M-6
1. Subbasal tooth of cercus vestigial (figs. 376377b)................................................................2
– Subbasal tooth of cercus well developed (figs.
375, 378-381b).................................................3
2. Pale thoracic mesanepisternal stripes complete
(fig. 81); outer base of cercus with a pale spot (fig.
376a-b); membranule dark except basal 20%
pale; abdominal segments V-IX with PL spot (fig.
138b); stem of T-spot of frons approximately
parallel-sided (fig. 20a); frontoclypeal groove
lacking a black stripe (figs. 20b-c); Argentina and
Chile north to Brazil (fig. 450).................confusa
– Pale thoracic mesanepisternal stripes incomplete
(figs. 79-80); cercus without pale spots (fig. 377ab); membranule pale except distal 30-50% dark;
abdominal segments V-IX without PL spot (fig.
137b); stem of T-spot of frons gradually widening
basally (fig. 19a); frontoclypeal groove with a black
stripe (figs. 19b-c); Baja California state in Mexico
north to southwestern USA (fig. 449) .....californica
3. Pterothorax without pale markings or with small
rounded pale spots at ventral and/or dorsal edges
of pleura (figs. 86-90); membranule dark except
basal 20% pale; subbasal tooth of cercus prominent and with acute angle (fig. 380b); northern
Chile (fig. 451).............................................tinti
– Pterothorax with pale stripes (figs. 76-78, 82-85,
93); membranule pale except distal 30-50% dark;
subbasal tooth of cercus with orthogonal angle
(figs. 375, 378-379, 381b) ................................4
4. Frontal carina in dorsal view angled anteriorly
(fig. 18a); pale metepimeral stripe constricted or
divided into a ventral rounded spot and a dorsal
T-shaped spot (figs. 76-78); southeastern Brazil
(fig. 450).............................................brasiliensis
– Frontal carina in dorsal view evenly curved (figs.
21, 23-24a); pale metepimeral stripe complete
(figs. 82-84, 93) ................................................5
5. Frontoclypeal groove lacking a black stripe (fig.
21b-c); anterior process of hamulus prominent,
projecting as high as 50% of its width (fig. 297b);
cercus with a marked dorsal constriction at distal
25% (fig. 378a); Bolivia to Venezuela (fig. 451) ..
..............................................................marchali
– Frontoclypeal groove with a black stripe (figs. 22,
24b-c); anterior process of hamulus not prominent, projecting only as high as 30% of its width
(fig. 298-299b); cercus with a slight dorsal constriction at distal 25% (figs. 379, 381a) ............6
6. Confluence of eyes long (frons + vertex/eyes + occipital triangle less than 1; eyes/occipital triangle
–
equal to 2) (fig. 24a-b); distal portion of ventral
margin of anterior lamina spine convex (fig. 339);
larger species (total length 57.8-67.9 mm, cercus
length 4.45-5.85 mm); Chile and Argentina (fig.
452)........................................................variegata
Confluence of eyes short (frons + vertex/eyes +
occipital triangle larger than 1; eyes/occipital triangle less than 2) (fig. 22a); distal portion of ventral margin of anterior lamina spine linear (fig.
338); smaller species (total length 51.75-56.5
mm, cercus length 3.75-4.5 mm); Bolivia and
Peru (fig. 452) ..........................................peralta
Key to females of Rhionaeschna
Females of R. condor, R. decessus, and R. vazquezae
are unknown, but I tentatively include them here
based on characters shared by males and females in
other species.
1. Pterothorax and abdominal segments VII-X without pale markings (figs. 39, 119c); frons projected
dorsally, higher than vertex (fig. 1b); cercus tip
bearing spine (fig. 358c, not to be confused with
mucron, i.e. figs. 359c, 360h, 361-362c, 363c,
364c, 385c,d); Venezuelan tepuis (fig. 439)..draco
– Pterothorax with or without pale stripes, abdominal segments IV-VIII to X with pale spots (figs. 40118, 120-156c-d); frons not projected dorsally,
lower than vertex (figs. 2-38b); cercus tip lacking
spine (figs. 359c, 360c, g-h, 361-362c, 363c,
364-365c, 366-369d, 370-395c).......................2
2. Pterothorax with marbled pattern of black markings and pale areas (figs. 40-47) ..............Key F-1
– Pterothorax uniform in color (figs. 59, 67, 90) or
with pale stripes or spots (figs. 48-58, 60-66, 6889, 91-118).......................................................3
3. Surface of abdominal ventral terga delimited by inner and outer carinae narrow (width measured at
narrowest point/length of ventral terga IV less than
0.09) (figs. 181-188B); HW with two rows of cells
between RP1 and RP2 beginning under pterostigma
or proximal to it (figs. 401-402)............................
.................................................................Key F-2
– Surface delimited by inner and outer carinae of
abdominal ventral terga wide (width measured at
narrowest point/length of ventral terga IV larger
than 0.10) (figs. 165-180, 189-194B); HW with
origin of two rows of cells between RP1 and RP2
variable .............................................................4
4. Two rows of cells between RP1 and RP2 in HW beginning under pterostigma or proximal to it (fig.
403) .......................................................Key F-3
– Two rows of cells between RP1 and RP2 in HW beginning at level of pterostigma distal end or distal
to it (figs. 398-400)...........................................5
5. Frontoclypeal groove lacking a dark stripe (figs.
77
T  E,  146, 2003
–
9-12, 14, 16, 20-21b-c) ..........................Key F-4
Frontoclypeal groove with a dark stripe (figs. 13,
15, 17-19, 22-24b-c) ..............................Key F-5
Key F-1
1. Constriction of abdominal segment III slight
(figs. 121-122, 124c-d); cercus length: 1.7-2.3
mm (figs. 360c, g-h, 361-362c) ........................2
– Constriction of abdominal segment III marked
(figs. 120, 123, 125-126c-d); cercus length: 2.76.5 mm (figs. 359, 363-365c) ...........................4
2. Frontal carina linear in frontal view (fig. 7c); Argentina (Buenos Aires and Mendoza north to
Salta province) (fig. 441)..........................pallipes
– Frontal carina with a medial cleft in frontal view
(figs. 3-4c) ........................................................3
3. Cleft of frontal carina slight (fig. 3c); inner and
outer carina of ventral terga VI sinuous (fig.
159c); black spots of pterothorax reduced to 1 on
mesanepisternum, 2 on mesepimeron, 1-2 on
metepisternum and 1 on metepimeron (figs. 4142); Valdivia province in Chile north to Peru (fig.
440) .....................................................brevifrons
– Cleft of frontal carina deep (fig. 4c); inner and
outer carina of ventral terga VI linear, parallel (figs.
160b); black spots of pterothorax represented by
2-3 on mesanepisternum, 4-5 in mesepimeron
and 3 on metepimeron (Fig 43); northwestern Argentina north to Peru (fig. 440) ..............fissifrons
4. Shorter cercus (2.7-3.7 mm) (figs. 355, 359c);
smaller species (total length 66-73.5 mm).........5
– Longer cercus (4.9-6 mm) (figs. 360-361c); larger species (total length 72-80.5 mm) .................6
5. Stem of T-spot strongly narrowed before its confluence with transverse arms (fig. 5a); ventral tubercle of abdominal segment I high (about as
high as 50% of its length in lateral view) (fig.
238); cercus: 3.7 mm (fig. 363c); Peru to
Ecuador (fig. 442)..................................intricata
– Stem of T-spot widely confluent with transverse
arms (fig. 2a); ventral tubercle of abdominal segment I low (about as high as 30% of its length in
lateral view) (as in fig. 236); cercus: 2.7-3 mm (fig.
359c); Ecuador to Venezuela (fig. 442) .................
..............................................................brevicercia
6. Stem of T-spot widely confluent with transverse
arms; vertex black with a small rounded yellow
spot on each anterior lateral edge (fig. 6a); cercus
widening distally with pointed tip (Fig 364c);
Bolivia north to Peru (fig. 443)................obscura
– Stem of T-spot strongly narrowed before its confluence with transverse arms; vertex yellow with
posterior 30% black (fig. 8a); cercus with inner
and outer margins approximately parallel, tip not
pointed (fig. 365c); northwestern Argentina to
Peru (fig. 443)..............................vigintipunctata
78
Key F-2
1. Pale mesanepisternal and mesepimeral stripes
complete and as wide as 60% of their sclerites
(figs. 108-110); Guerrero and Nayarit states in
Mexico (fig. 456) .................................vazquezae
– Pale mesanepisternal stripes complete or incomplete, mesepimeral complete, as wide as 30-50%
of their sclerites (fig. 95-107) ............................2
2. Cercus longer than segments VIII-X and wider
than 1.15 mm (figs. 384, 388c) ........................3
– Cercus shorter than segments VIII-X and as wide
as or narrower than 1.1 mm (figs. 382-383, 385387c) ................................................................4
3. Mesanepisternal stripes yellowish-light blue and
incomplete (fig. 99-100); Baja California state in
Mexico (fig. 456) ......................................manni
– Mesanepisternal stripes bright green and complete (fig. 107); northern Argentina to southern
USA (fig. 455) ..............................................psilus
4. Pale mesepimeral and metepimeral stripes with
deep rounded indentations (figs. 97-98, 101,
104-106)...........................................................5
– Pale mesepimeral and metepimeral stripes lacking
indentations (figs. 92-94) or with broad and shallow indentations (figs. 101-102) .......................7
5. Stem of T-spot of frons abruptly narrowed anteriorly (fig. 26a); cercus as long as or slightly shorter than abdominal segments IX-X (fig. 383c);
northwestern Argentina (fig. 453) ...........haarupi
– Stem of T-spot of frons gradually narrowed anteriorly (figs. 29-30); cercus longer than abdominal
segments IX-X (figs. 386c-e, 387c) .....................6
6. Pale mesanepisternal stripes incomplete to absent
(fig. 100), as wide as 25-33% of mesanepisternum; ventral tubercle of abdominal segment I
lower than 30% of its length (fig. 265); cercus
with rounded tip bearing a minute medial spine
(fig. 387c); Paraná and Minas Gerais states in
Brazil (fig. 453)..........................................pauloi
– Pale mesanepisternal stripes complete (figs. 104106), as wide as 50% of mesanepisternum; ventral
tubercle of abdominal segment I about as high as
50% of its length (as in figs. 261-262); cercus with
pointed medial tip (fig. 386c-e); Argentina north
to Venezuela (fig. 453) .........................planaltica
7. Mesepimeral stripe narrowed to 50% of its basal
width at midlength (figs. 101-102); cercus with
mucron on external margin (figs. 385c-d); anterior side of ventral tubercle concave (fig. 267);
Venezuelan tepuis (fig. 454)..................nubigena
– Mesepimeral stripe approximately uniform in
width or slightly narrowed at midlength (figs. 9294); cercus tip rounded or medially pointed (figs.
382c-e); anterior side of ventral tubercle convex
(figs. 261-262); Bolivia north to Mexico (fig.
454) ......................................................cornigera
Key F-3
1. Black stripe on frontoclypeal groove strongly
widened toward the eyes (figs. 37-38b-c);
pterothoracic stripes complete, linear, and narrower than 50% of their sclerites (figs. 114, 118)......2
– Frontoclypeal groove not bordered by black
stripe (figs. 33-36b-c); pterothoracic stripes with
deep indentations (figs. 113, 115), divided into
spots (figs. 110, 111, 116, 117), or wider than
50% of their sclerites (fig. 112) .........................4
2. Labrum, clypeus and frons orange; pale abdominal
PL spots present on segments IV-IX, fused with ML
on IV-VIII; only known from Itatiaia (type locality),
Rio de Janeiro state, Brazil (fig. 459) (characters
abstracted from Calvert, 1953...................decessus
– Labrum, clypeus and frons light blue or yellow;
pale abdominal PL spots absent on segments VIVII, fused or not with ML on VIII (as in figs. 155156c-d).............................................................3
3. Ventral tubercle of abdominal segment I low and
rounded in lateral view (fig. 275); black of T-spot
not extended on antefrons; vertex yellow with lateral edges black (figs. 38b-c); cercus with rounded tip, longer than abdominal segments IX-X and
shorter than VIII-IX (fig. 395c); São Paulo, Rio de
Janeiro and Minas Gerais states in Brazil (fig.
459).......................................................punctata
– Ventral tubercle of abdominal segment I high and
trapezoidal shaped in lateral view (fig. 274); black
of T-spot extended on dorsal surface of antefrons;
vertex yellow with lateral and posterior 30% black
(figs. 37b-c); cercus with acute mucronate tip, as
long as abdominal segments VIII-X; Minas Gerais
state in Brazil (fig. 459) ..........................eduardoi
4. Transverse arms of T-spot represented by a fine
line (fig. 33a); labrum orange, clypeus and frons
bright yellow; no dark stripe on frontoclypeal
groove (figs. 33b-c); Peru to Ecuador (fig. 457) ....
..................................................................biliosa
– Transverse arms of T-spot wide (fig. 34-36a);
labrum, clypeus and frons light blue to green or
yellow; dark stripe on frontoclypeal groove (figs.
34-36b-c)..........................................................5
5. Stem of T-spot wider than vertex (fig. 34a); pale
pterothoracic stripes wide (as wide as 50-80% of
their sclerites) and complete (fig. 112); Táchira
state in Venezuela (fig. 457) ......................condor
– Stem of T-spot narrower than vertex (figs. 3536); pale mesepimeral and metepimeral stripes
narrow (as wide as 30-50% of their sclerites) and
complete with an anterior and a posterior indentation (figs. 113, 115), or divided into spots (figs.
116-117) ..........................................................6
6. PD spots absent in segments VIII-X (figs. 154c-e);
outer margin of segment VII concave (fig. 192b);
maximum width of cercus at medial 30% (fig.
–
391c); larger species (cercus 6.5 mm, total length
68-72.5 mm); Bolivia to Colombia (fig. 458) .....
...............................................................joannisi
PD spots present in segments VIII-X; outer margin
of segment VII linear (fig. 191b); maximum width
of cercus at distal 30% (fig. 393c); smaller species
(cercus 5 mm, total length 58.3 mm); Táchira
State in Venezuela (fig. 458) ..............demarmelsi
Key F-4
1. Medio-longitudinal pale stripe on abdominal segment II complete (figs. 132, 134, 138-139c-d);
South America .................................................2
– Medio-longitudinal pale stripe on abdominal segment II incomplete or interrupted at midlength
(figs. 127-130c-d); Central and North America 5
2. Supratriangles always crossed (1-4 crossveins)
(fig. 400)...........................................................3
– Supratriangles generally free, sometimes one or
two with 1-2 crossveins (in 2-10% of specimens
examined) (fig. 399) .........................................4
3. Pale mesanepisternal stripe complete (fig. 81);
Chile and Argentina north to Brazil (fig. 450) ....
................................................................confusa
– Pale mesanepisternal stripe incomplete or absent
(fig. 79-80); Bolivia to Venezuela (fig. 451) ........
..............................................................marchali
4. Clypeal lobes rounded (fig. 16c); stem of T-spot
widening basally (fig. 16a); membranule dark except basal 30% pale; cercus tip not pointed (fig.
with few denticles restricted to apex (fig. 250);
central Chile to Ecuador (fig. 448) ...............elsia
– Clypeal lobes angled (fig. 14c); stem of T-spot
parallel sided (fig. 14a); membranule dark except
basal 15% pale; cercus tip pointed (fig. 371c);
ventral tubercle of abdominal segment I with numerous denticles spread over posterior surface (as
fig. 248); Argentina to Bolivia and Brazil (fig.
447) ...................................................bonariensis
5. Posterior margin of inner carina of ventral terga
V-VII convergent to outer carina (fig. 165b); cercus tip rounded (fig. 366d); Mexico to southern
USA (fig. 444) .............................................dugesi
– Posterior margin of inner carina of ventral terga
V-VII parallel to outer carina (figs. 166-168b); cercus tip pointed (figs. 367-369d)........................6
6. No pale abdominal PL spots on segments IV-V
(fig. 128d); Costa Rica to Mexico (fig. 444)........
.............................................................jalapensis
– Pale abdominal PL spots of segments IV-V present
(figs. 129-130d) ................................................7
7. Ventral tubercle of abdominal segment I low (as
wide as about 3 times its height) (fig. 245); base
of stem of T-spot wider than vertex, anterior of
vertex half pale, posterior 50% black (fig. 12a);
79
T  E,  146, 2003
–
outer margin of cercus linear (fig. 369d); eastern
USA to southern Canada (fig. 445)............mutata
Ventral tubercle of abdominal segment I high (as
wide as 2 times its height) (fig. 244); base of stem
of T-spot narrower than vertex, vertex pale with
posterior 30% black (fig. 11A); outer margin of
cercus slightly convex (figs. 368D-E); Mexico to
southern Canada through western USA (fig. 445)
............................................................multicolor
Key F-5
1. Pale PL spots absent on segments V-IX (fig. 137d);
Baja California state in Mexico north to Canada
through southwestern USA (fig. 449).....californica
– Pale PL spots present on segments V-IX (figs. 131,
133, 135-136, 140-142d); South America ........2
2. Clypeal lobes angled (figs. 13, 15, 18, 22-24c);
ventral tubercle of abdominal segment I with numerous denticles spread over its posterior surface
(figs. 247, 249, 252, 256-258) ..........................3
– Clypeal lobes rounded (fig. 14c); ventral tubercle
of abdominal segment I with few denticles restricted to its apex (fig. 251); Galápagos Islands
(fig. 448) .........................................galapagoensis
3. Tip of cercus rounded (figs. 372, 379-381); ventral tergum VI widest point at basal 30% (figs.
171, 178-180b).................................................4
– Tip of cercus pointed (figs. 370, 375); ventral tergum VI widest point at distal 60% (figs. 169, 174)
.........................................................................7
4. Larger species (total length 55.8-67.2 mm, cercus
4.3-5.5 mm); southern Argentina and Chile to
southern Peru (fig. 452) ........................variegata
– Smaller species (total length 47-54.2 mm, cercus
3-4.35 mm) ......................................................5
5. MD spots widely confluent to ML in segments IIIIX (fig. 141c-d); stem of T-spot abruptly narrowing at distal 30% (fig. 23a); northern Chile (fig.
451) .............................................................tinti
– MD spots separated from ML in segments III-IX
(figs. 133, 140c-d); stem of T-spot gradually narrowing to distal 30% (figs. 22, 15a) ..................6
6. ML and PL spots separated on segments IV-V (fig.
140d); supratriangles always crossed (1-2 cross
veins) (fig. 397); Bolivia and Peru (fig. 452) .......
.................................................................peralta
– ML and PL spots confluent on segments IV-V (fig.
133d); supratriangles usually free, sometimes one
or two with 1-2 cross veins (in 10% of specimens
examined as in fig. 396); Chile and southern Argentina (fig. 447)......................................diffinis
7. Abdominal segment II lacking a medio-longitudinal black stripe (fig. 136c); pale metepimeral
stripe divided into a ventral rounded spot and a
dorsal T-shaped spot (fig. 76-78); frontal carina in
dorsal view angled anteriorly (fig. 18a); supratri80
–
angles always crossed (1-2 cross veins) (fig. 400);
southeastern Brazil (fig. 450) ..............brasiliensis
Abdominal segment II with a complete mediolongitudinal black stripe between transverse carina and posterior margin (fig. 131c); pale
metepimeral stripe complete to absent, but never
divided into ventral and dorsal spots (figs. 5459); frontal carina in dorsal view smoothly curved
(figs. 13a); supratriangles generally free, sometimes one with 1 cross vein (in 10% of specimens
examined) (fig. 399); southern Argentina to
Ecuador (fig. 446) ..................................absoluta
Rhionaeschna species accounts
In order to facilitate species treatment I have allocated the 39 species into seven groups. The groups are
based on male characters used to build the subordinate keys, and do not necessarily imply clades.
1. R. draco group
Frons projected dorsally, higher than vertex (fig.
1b); pterothorax and abdominal segments VII-X without pale markings (figs. 39, 119a-b); hamulus with a
medial digitiform projection on its antero-ventral
margin (fig. 276); female cercus with long terminal
spine (fig. 439). One species: R. draco.
Rhionaeschna draco (Rácenis) comb. n.
(figs. 1-he, 39-th, 119-ab, 157-te, 195-ge, 234-tg, 276-ha, 316-as,
358-ce, 397-wi, 439, 460-Mp)
Aeshna draco Rácenis, 1958: 323, 326-330, figs. 1, 3a-c (description ?: holotype ? VENEZUELA: El Dragón-Auyantepui, 1700 m (Bolivar dept.), 15-IX-1958, Trebbau leg.
(MIZA) [examined: allotype / (GSVC)]); Rácenis 1970:
31; De Marmels 1989b: 35; De Marmels 1990a: 3-5 (description larva); De Marmels 1992b: 41; Vick 1993: 9399 (description /).
Diagnosis. – Rhionaeschna draco is distinguished
from all other species of Rhionaeschna by the projected
edge of frons (fig. 1b), uniformly colored reddish
brown thorax (fig. 39), abdomen with pale spots only
in segments II-VI (fig. 119), antero-ventral margin of
hamuli with a medial digitiform projection (fig. 276),
and tip of female cerci bearing a long spine (fig. 358c).
Description. – Anteclypeus pale brown, postclypeus
and frons bright light blue, grayish white area surrounding T-spot stem; T-stem widening basally with
convex sides; frontoclypeal groove lacking dark stripe;
vertex light blue, with anterior and posterior margins
black; edge of frons angled in lateral view and projected antero-dorsally (higher than vertex) (figs. 1a-c).
Thorax reddish brown, without pale stripes or spots
(fig. 39). Abdomen dark reddish brown, pale abdominal spots bright light blue, restricted to PD in II, AD, AL
and ML in III, and AL in IV-VI (figs. 119a-c). Ventral tubercle of abdominal segment I with anterior margin
concave in lateral view (fig. 234); spines of anterior
lamina long (as long as 37% of its basal width) and triangular (fig. 316); anterior margin of hamuli with a
medial digitiform projection, tip of hamular anterior
process rounded in posterior and ventral view (fig.
276); auricles with two teeth (fig. 234). Lateral carina
of male cercus concave; dorso-distal crest lower than
base of cercus in lateral view and smoothly curved
along distal 25% of cercus; sub-basal tooth low and
blunt (fig. 358b); maximum width at medial 30%; tip
directed externally (fig. 358a). Female cercus as long as
abdominal segment IX; maximum width at distal
25%; tip bearing a spine (fig. 358c). Dimensions:
head width: 10.65-11.2; HW length: 49-55.5; HW
width: 14.1-15.2; HW pterostigma length: 2.54-3; cerci length: ? 6.5, / 2.52; female cerci maximum
width: 0.75; total length: 77.5- 82.
Biology. – De Marmels (1990a) described last larval instar by supposition based on larvae and exuviae
from Bolivar and Amazonas, Venezuela, and reported
adults and larvae as common in Cerro Neblina. The
species breeds in mountain creeks and rock pools in
the tepuis. Rare in collections.
Distribution [0-5°N, 60-66°W, 1700-2800 m]
(figs. 439, 460). – Venezuela: Bolivar (GSVC*; MLPA*;
Rácenis 1958, 1970; De Marmels 1990a, 1992b;
Vick 1993) and Amazonas (USNM; De Marmels
1989b, 1990a).
2. Marmaraeschna group
(= subgenus Marmaraeschna Calvert, 1952).
Frontal carina recessed, located in the horizontal
portion of the frons, giving a rounded contour to antero-dorsal margin of head (figs. 2-8b); pterothorax
with marbled pattern of black markings and pale areas
(figs. 40-47); two rows of cells between RP1 and RP2 in
HW beginning proximal to pterostigma or under it
(fig. 396); hamular anterior process very high (distance between ventral tip of process and axis of hamulus higher than distance between dorsal end of hamular fold and axis) (figs. 277-284); anterior lamina spine
wide throughout its length in lateral view (twice or less
as wide at base as at distal 30% (figs. 317-324); external margin of male cercus approximately linear in lateral view (with the exception of intricata, where it is
slightly concave) (figs. 359-365). Seven species: R. brevicercia, R. brevifrons, R. fissifrons, R. intricata, R. obscura, R. pallipes, R. vigintipunctata.
Rhionaeschna brevicercia (Muzón & von
Ellenrieder) comb. n.
359-ce, 442-Mp)
Aeshna (Marmaraeschna) brevicercia Muzón & von Ellenrieder, 2001: 98-102, 122 (description ? /: holotype ?
ECUADOR: Pichincha, between Calacali and Nanegalito,
ca. 30 km W Quito, 2000 m, 04-XI-1990, O.S. Flint Jr.
leg. (USNM) [examined: holotype ? allotype / paratypes
? / (RWGC; USNM; MIZA; MLPA)]).
[Aeshna (Marmaraeschna) intricata. – Martin 1908: 59-60
(in part); Calvert 1956: 112-119 (in part). Misidentification]
[Aeshna (Marmaraeschna) vigintipunctata. – De Marmels
1988: 101; De Marmels 1990b: 337; De Marmels 1994:
437; De Marmels 2001a: 130-132 (description larva).
Misidentification]
Diagnosis. – Rhionaeschna brevicercia shares with R.
obscura the wide confluence of stem and transverse
arms of T-spot but can be easily distinguished from it
by the anterior half of vertex yellow (vertex black with
small yellow antero-lateral spots in R. obscura, figs. 2,
6a), male cercus shape (cercus with sub-basal tooth low
and dorso-distal crest lower, sub-basal tooth prominent and dorso-distal carina higher in R. obscura, figs.
359, 364b), and female cercus size (not longer than 3
mm in R. brevicercia, 5.65 mm or longer in R. obscura,
figs. 359, 364c). It has been confused with R. vigintipunctata but is separated by the wide confluence of Tspot stem and transverse arms (narrow in R. vigintipunctata, figs. 2, 8a), sub-basal tooth of male cercus
low (prominent in R. vigintipunctata, figs. 359, 365b),
lower dorso-distal crest (higher in R. vigintipunctata,
figs. 359, 365b), and female cercus mucronate with inner and outer margins diverging distally (tip not mucronate and margins parallel along medial 30% in R.
vigintipunctata, figs. 359, 365c). Rhionaeschna brevicercia is most similar to R. intricata in size and shape of
cercus but it differs by the wide confluence of stem and
transverse arms of T-spot (narrow in R. intricata, figs.
2, 5a), the lower ventral tubercle of abdominal segment
I (higher in R. intricata, figs. 236, 238), and the shorter female cercus (2.7-3 mm in R. brevicercia, 3.7 mm in
R. intricata, figs. 359, 363c).
Description. – Clypeus and frons light blue and yellow to grayish blue, grayish blue spots lateral to light
yellow area surrounding T-spot stem; T-stem maximum width at mid-length, sides convex, widely confluent with transverse arms (fig. 2a); wide dark brown
stripe on frontoclypeal groove (figs. 2b-c); anterior
50% of vertex yellow, posterior 50% black. Frontal
carina with a slight concavity in frontal view (fig. c);
frons not flattened (fig. 2b). Pterothorax pale yellowbrown 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 ventral abdominal terga II extended over auricle; auricles
81
T  E,  146, 2003
with two teeth (fig. 196); constriction of segment III
marked (figs. 120a-d). Inner and outer carinae of ventral terga VI sinuous (figs. 158a-b). Ventral tubercle of
abdominal segment I lower than 30% of its length in
lateral view, with anterior margin convex in lateral
view; genital lobe as high as twice the tubercle height
(fig. 236). Sub-basal tooth of male cercus low; dorsodistal crest lower than 50% of cercus width at base, developed at distal 20% of cercus (fig. 359b); tip on external margin (fig. 359a). Female cercus maximum
width at distal 30%; tip mucronate (fig. 359c). Dimensions: head width: 8.9-10.2; HW length: 48.2-52;
HW width: 13.6-16.3; HW pterostigma length: 2.6-3.2;
cerci length: ? 5.4-6, / 2.7-3; female cerci maximum
width: 0.75; total length: 66.6-73.5.
Biology. – De Marmels (2001) described the larva
based on reared specimens found along small to
medium-sized streams with dense riparian vegetation
and submerged roots bordered by bushes and scattered trees outside closed forest. He reported adults
patrolling over open pasture lands along or away from
small streams and occasionally perching on vertical
rock surfaces where they were almost invisible due to
their lichen-like color. From Ecuador it has been reported from clear, sandy and rocky bottom streams
with good current, surrounded by marsh plants and
shrubs, and one larger pool (Dr. D. R. Paulson pers.
comm.). A common species.
Remarks. – Martin (1908) described R. intricata
based on several specimens from Ecuador, Colombia
and Venezuela. Examination of these specimens revealed that the type series of R. intricata included several specimens of R. brevicercia, from Venezuela
(without localities), Colombia and Ecuador.
Distribution [3°S-9°N, 70-79°W, 604-3020 m]
(fig. 442). – Ecuador: Tungurahua (RWGC*), Esmeraldas (ISNB; MLPA), Imbabura (ANSP; ISNB; QCAZ), Napo
(KJTC*), Manabi (UMMZ), and Pichincha (DRPC; FSCA;
MLPA*; ZMHB; QCAZ; USNM*) – Colombia: Santander
(ISNB); Cundinamarca (UMMZ) – Venezuela: Trujillo
(MIZA), Mérida (MIZA; MLPA), Barinas (MIZA), and
Táchira (MIZA; De Marmels 2001a).
Rhionaeschna brevifrons (Hagen) comb. n.
(figs. 3-he, 41-42-th, 121-ab, 159-te, 197-ge, 235,237-tg, 279-280ha, 318-319-as, 360-ce, 396-wi, 440-Mp)
Aeshna brevifrons Hagen, 1861: 129, 314 (description ? /:
lectotype ? MEXICO: Acapulco (MCZ) [examined: lectotype ? paralectotype / (MCZ)]; Hagen 1875: 36; Ris
1904: 4, 29; Calvert 1905:181, 186; Calvert 1908: 488;
Martin 1908:58, 59; Martin 1911: 12; Calvert 1952: 256
(Marmaraeschna n. subgn.); Schmidt 1952: 238 (in part);
Calvert 1956: 13, 110-112 (in part); Paulson 1977: 175
(in part); Muzón & von Ellenrieder 2001: 144-148 (redescription ? /).
Rhionaeschna maita Förster, 1909: 220-223 (description ?:
82
lectotype ? (here designated) PERU, Arequipa, 2300 m,
14-IV-1907, K. Seyd. leg. (MWNH) [examined]); Martin
1911: 13; Ris 1913: 83; Hincks 1934: 80: Schmidt 1952:
256. Syn. n.
Aeshna (Rhionaeschna) maita. – Calvert 1956: 11, 99-100.
[Aeshna (Marmaraeschna) intricata. – Calvert 1956: 112119 (in part); Fraser 1957: 155, 159, 163, fig. 4g; Jurzitza
1989: 9 (in part). Misidentification]
Diagnosis. – Rhionaeschna brevifrons shares with R.
pallipes and R. fissifrons a flattened frons (figs. 3-4, 7b),
stem of T-spot widest at posterior end and narrowly
confluent or separated from transverse arms (figs. 3-4,
7a), stocky abdomen (constriction of segment III slight,
figs. 121-122, 124), and short cercus (less than 4.8 mm
in males, 2.3 mm in females, figs. 360-362). The three
species can be distinguished from each other by the
shape of the frontal carina (linear in R. pallipes, fig. 7c,
concave in R. brevifrons, fig. 3c, deeply cleft in R. fissifrons, fig. 4c); and shape of hamular anterior process
tip (figs. 280-282b-c). In addition, the sub-basal tooth
of male cercus in R. brevifrons is low (figs. 360b, e-f),
not prominent as in R. fissifrons and R. pallipes (figs.
361-362b); the pterothorax has fewer black spots (figs.
41-44); and in R. fissifrons the inner and outer carinae
of abdominal ventral terga VI are linear and parallel to
each other (fig. 160) (sinuous in R. brevifrons and R.
pallipes, figs. 159, 163), and in R. pallipes the black areas of ventral abdominal terga II do not extend over
auricles (figs. 197-198, 201).
Description. – Clypeus and frons light blue and yellow to grayish blue, grayish dark blue spots lateral to
light blue area surrounding T-spot stem; T-stem maximum width at posterior end, sides converging anteriorly, narrowly or not confluent with transverse arms
(figs. 3a, c); narrow black stripe on frontoclypeal
groove (figs. 3b-c, d-e); vertex yellow with posterior
margin black. Frontal carina with a slight concavity in
frontal view (figs. 3c, e); frons flattened (figs. 3b, d).
Pterothorax pale yellow-brown with 1 black spot in
mesanepisternum, 2 in mesepimeron, 1-2 in metepisternum and 1 in metepimeron (figs. 41-42). Abdomen pale brown to pale yellow and grayish light
blue with dark reddish brown to black spots; black areas of ventral abdominal terga II extended over auricle
(fig. 197); constriction of segment III very slight (figs.
121a-d). Inner and outer carinae of ventral terga VI
sinuous (figs. 159a-c). Ventral tubercle of abdominal
segment I lower than 30% of its length, with anterior
margin convex in lateral view; genital lobe approximately as high as tubercle (figs. 235, 237); auricles
with two teeth (fig. 197). Sub-basal tooth of male cercus low; dorso-distal crest lower than 50% of cercus
width at base, developed at distal 25% of cercus (fig.
360e); tip on external margin (fig. 360a, d). Female
cercus maximum width at distal 80%; tip mucronate
or not (figs. 360c, g-h). Dimensions: head width: 8.69.5; HW length: 41.7-46; HW width: 13.5-15; HW
pterostigma length: 2.6-3.1; cerci length: ? 4.2-4.8,
/ 1.9-2.3; female cerci maximum width: 0.5; total
length: 57-69.5.
Biology. – Collection labels indicate that in Northern Chile specimens were collected flying over thermal creeks in areas of naturally warm waters used for
valley irrigation in the middle of the desert, and in a
valley surrounded by high mountains with small irrigation streams for alfalfa crops. A common species.
Larva unknown.
Remarks. – The type locality of Rhionaeschna brevifrons (Mexico, Acapulco) is probably a labeling mistake, since no Marmaraeschna species has ever been
found north of Venezuela, and no specimen of R. brevifrons north of Peru. Förster (1909) described Rhionaeschna maita based on two males from Arequipa,
Peru, one of them deposited in the Museum Wiesbaden and the second one in his collection. The second male is apparently lost (Calvert 1956: 99), and
the first one is here designated as lectotype. Examination of the lectotype showed that Rhionaeschna maita
Förster 1909 is a junior synonym of Rhionaeschna brevifrons (Hagen 1861). The specimen is poorly preserved; the head is flattened, thorax partly broken, and
cerci seem to be atrophied (left) and malformed
(right), but all diagnostic characters of R. brevifrons
can be distinguished and wing dimension and venation characters agree completely with those for R. brevifrons. Notes on the types of R. maita made by E.
Schmidt in Calvert (1956) include some errors (i.e.
stated absence of frontal carina and anterior lamina
spines), which are probably responsible for the misplacement of this species until now.
Distribution [9-44°S, 68-77°W, 510-3750 m] (fig.
440). – Chile: De Los Lagos: Valdivia (IEUM), General O´Higgins: Cachapoal (USNM), Metropolitan:
Chacabuco (MNHN), Cordillera (IEUM), and Santiago
(IMLA; IEUM; RWGC*; USNM), Valparaiso: Valparaiso
(IEUM; MCZ*; MNHN), Atacama: Huasco (FSCA*), Coquimbo: Elqui (IEUM) and Choapa (DRPC; FSCA;
IEUM; MLPA; RWGC; UMMZ), Antofagasta: El Loa
(MLPA; MNHN; UMMZ), and Tarapacá: Parinacota
(FSCA; MLPA; UMMZ), Arica (FSCA; MLPA; UMMZ) and
Iquique (FSCA; MLPA; MNHN; UMMZ) – Peru: Arequipa
(ANSP; MCZ; MWNH*; USNM; Förster 1909), Ayacucho
(FSCA), Apurimac (MLPA; RWGC*; WB), Lima (DRPC;
IMLA*; RWGC*), and Ancash (DRPC).
Rhionaeschna fissifrons (Muzón & von Ellenrieder)
comb. n.
361-ce, 440-Mp)
Aeshna (Marmaraeschna) fissifrons Muzón & von Ellenrieder, 2001: 148-150 (description ? /: holotype ? ARGENTINA: 7 km S Minas Capillitas (Catamarca prov.), ca.
3000, 27-XII-1072, J. Schultz leg. (FSCA) [examined: ?
holotype, / allotype, ? / paratypes (FSCA; DRPC; MLPA;
UMMZ; RWGC)]).
[Aeshna (Marmaraeschna) brevifrons. – Schmidt 1952: 238
(in part). Misidentification]
[Aeshna (Marmaraeschna) intricata. – Calvert 1956: 13,
112-119 (in part); Rácenis 1959: 494 (in part); Paulson
1977: 175 (in part); Rodrigues Capítulo et al. 1991: 62
(in part); Rodrigues Capítulo 1992: 38, 57 (in part).
Misidentification]
Diagnosis. – The deeply cleft frontal carina (fig.
4c) and linear contour of ventral terga (fig. 160) are
unique character states of Rhionaeschna fissifrons. Further distinction from the other species is given under
R. brevifrons.
Description. – Clypeus and frons pale brown, grayish dark blue spots lateral to yellow area surrounding
T-spot stem; T-stem maximum width at posterior
end, sides converging anteriorly, separated from transverse arms (fig. 4a); wide black stripe on frontoclypeal
groove (fig. 4b-c); vertex yellow with lateral and posterior margins black. Frontal carina with a deep medial
cleft in frontal view (fig. 4c); frons flattened (fig. 4b).
Pterothorax pale yellow-brown with 2-3 black spots in
mesanepisternum, 3-4 in mesepimeron, 3-4 in
metepisternum and 3-4 in metepimeron (fig. 43). Abdomen grayish light blue and pale yellow with dark
reddish brown to black areas; black areas of ventral abdominal terga II extended over auricle; auricles with
two teeth (fig. 198); constriction of segment III very
slight (figs. 122a-d). Inner and outer carinae of ventral
terga VI linear and parallel to each other (figs. 160a-b).
Ventral tubercle of abdominal segment I lower than
30% of its length in lateral view, with anterior margin
convex in lateral view; genital lobe approximately as
high as tubercle (fig. 239). Sub-basal tooth of male
cercus prominent; dorso-distal crest lower than 50%
of cercus width at base, developed at distal 20% of cercus (fig. 361b); tip on external margin (fig. 361a). Female cercus maximum width at distal 66%; tip mucronate (figs. 361c). Dimensions: head width: 8.2-9.6;
HW length: 41.3-48.8; HW width: 13.8-16.1; HW
/ 1.7-2.2; female cerci maximum width: 0.45; total
length: 56-59.
Biology: In Peru adults of Rhionaeschna fissifrons
were seen flying over the steep part of a rocky stream
and grassy areas nearby, and landing on vertical or
horizontal rocks at the stream (DRPC), and in Northwestern Argentina patrolling mountain streams and
creeks in Preandean and Andean localities. Rare in
collections. Larva unknown.
Distribution [11-27°S, 65-75°W, 1200-4057 m]
(fig. 440). – Argentina: Catamarca (FSCA) and Salta
(MACN; MLPA*; ZMHB) – Chile: Antofagasta: El Loa
(UMMZ; MLPA*) – Bolivia: Cochabamba: Chapare
(FSCA) – Peru: Puno (UMMZ), Junín (IMLA; RWGC;
USNM), Ayacucho (FSCA), and Cuzco (DRPC).
83
T  E,  146, 2003
Rhionaeschna intricata (Martin) comb. n.
363-ce, 442-Mp)
Aeshna (Marmaraeschna) intricata Martin, 1908: 59-60, fig.
55 (in part; description ? /: lectotype / (here designated) ECUADOR: Nanegal, 0°7’0N 78°40’0W 2093 m
(Pichincha prov.) (ISNB) [examined]); Martin 1911: 12;
Calvert 1952: 256; Calvert 1956: 112-119 (in part);
Rácenis 1959: 494 (in part); Muzón & von Ellenrieder
1998: 23; Muzón & von Ellenrieder 2001: 150-154 (redescription ? /).
Diagnosis. – Rhionaeschna intricata is characterized
by the high ventral tubercle of abdominal segment I (as
high or higher than 50% of its length, fig. 238), which
is low in the remaining Marmaraeschna species (figs.
235-237, 239-242). Further differences from other
species are found under R. brevicercia and R. brevifrons.
Description. – Clypeus and frons gray with yellow
margins, light yellow area surrounding T-spot stem; Tstem maximum width at mid-length, sides convex,
strongly narrowed at confluence with transverse arms
(fig. 5); wide black stripe on frontoclypeal groove (fig.
5b-c); anterior 50% of vertex yellow, posterior 50%
black. Frontal carina with a slight concavity in frontal
view (fig. 5c); frons not flattened (fig. 5b). Pterothorax
pale yellow-brown with 2 black spots in mesanepisternum, 3-4 in mesepimeron, 2-3 in metepisternum and
2-3 in metepimeron (fig. 44). Abdomen pale reddish
brown to yellow and light blue with black spots; black
areas of ventral abdominal terga II extended over auricle; auricles with three teeth (fig. 199); constriction of
segment III marked (figs. 123a-d). Inner and outer carinae of ventral terga VI sinuous (figs. 161a-b). Ventral
tubercle of abdominal segment I as high or higher than
50% of its length in lateral view, with anterior margin
convex in lateral view; genital lobe approximately as
high as tubercle (fig. 238). Sub-basal tooth of male cercus very low and blunt; dorso-distal crest lower than
50% of cercus width at base, developed at distal 25%
of cercus (figs. 363b, d); tip medial (fig. 363a). Female
cercus maximum width at distal 66%; tip mucronate
(fig. 363c). Dimensions: head width: 9-9.5; HW length:
46.4-50.25; HW width: 14.1-16.4; HW pterostigma
length: 2.8-3.35; cerci length: ? 4.9-5.5, / 3.7; female cerci maximum width: 0.8; total length: 66-68.
Biology. – The following notes by Felix Woytkowski on envelopes of specimens he collected in
Peru state: ‘This species has been taken sitting on the
wall on a rock some 300 m higher in the mountains
east of Celendin far from any water. This is a new
species for the higher Andes, rare and difficult to get’.
Rare in collections. Larva unknown.
Remarks. – In his description of R. intricata Martin
(1908) indicated that the specimens he examined
from Mexico, Peru, Bolivia, Chile, Ecuador,
Venezuela and Brazil were deposited in the Selys and
84
Martin collections. In the Martin collection at the
there are no type specimens of R. intricata (Dr.
J. Legrand pers. com.), and nine specimens with type
labels are deposited in the Selys collection at the ISNB.
Examination of this series revealed that it includes two
species, which according to the last synopsis of the
group would correspond to R. intricata and R. brevicercia (Muzón & von Ellenrieder 2001). In that paper, identification of R. intricata was based on the illustration of the male cerci from the original
description (Martin 1908). However, no male specimen belonging to this species was found among the
types at the ISNB and all are R. brevicercia except for
one female, which is herein designated as lectotype of
R. intricata. In addition to the mixed series of nine
specimens at the ISNB collection there is also a pinned
type label [no sex stated] without an associated specimen. This label may correspond to the lost male illustrated by Martin, or it may belong with one of the
other missing specimens mentioned by Martin (1908)
of his collection from Mexico, Peru, Bolivia, Chile
and Brazil.
Distribution [0-6°S, 78°W, 2248-2833 m] (fig.
442). – Peru: Cajamarca (FSCA*; UMMZ) – Ecuador:
Chimborazo (ANSP; MLPA*) and Pichincha (ANSP*;
ISNB).
MNHN
Rhionaeschna obscura (Muzón & von Ellenrieder)
comb. n.
364-ce, 443-Mp)
Aeshna (Marmaraeschna) obscura Muzón & von Ellenrieder,
2001: 154-156 (description ? /: holotype ? BOLIVIA: 4
km W Chulimani (La Paz dept., sud Yungas prov.), 25-V1989, T.C. Emmel leg. (FSCA) [examined: holotype ?, allotype /, paratypes ? / (FSCA; ANSP; MLPA; UMMZ)]).
[Aeshna (Marmaraeschna) vigintipunctata. – Calvert 1956:
119-123 (in part). Misidentification]
Diagnosis. – The dark vertex of R. obscura (fig. 6a)
is unique to this species within the marmaraeschnagroup (figs. 2-5, 7-8a). Further differences from other
species are found under R. intricata and R. brevifrons.
This species is the largest marmaraeschna together
with R. vigintipunctata, both of which have a similarly
shaped male cercus (figs. 364-365b-c). However, the
widely confluent T-spot stem (fig. 6a) (narrow in R.
vigintipunctata, fig. 8a), shape of genital lobe (subquadrate in R. obscura, fig. 240; subtriangular in R.
vigintipunctata, fig. 242), and shape of female cercus
(mucronate tip and margins diverging distally in R.
obscura, fig. 364; no mucronate tip and margins parallel to each other in R. vigintipuncata, fig. 365), allow
for the distinction between these two species.
Description. – Clypeus and frons light blue, light
yellow area surrounding T-spot stem; T-stem maximum width at mid-length, sides convex, widely con-
fluent with transverse arms (fig. 6a); wide black stripe
on frontoclypeal groove (fig. 6b-c); vertex black with a
small yellow antero-lateral spot on each side. Frontal
carina with a slight concavity in frontal view (6c);
frons not flattened (fig. 6b). Pterothorax pale yellowbrown with 1 black spot in mesanepisternum, 3 in
mesepimeron, 2-3 in metepisternum and 2-3 in
metepimeron (fig. 45). Abdomen pale brown to reddish brown and light blue to yellowish green with
black spots; black areas of ventral abdominal terga II
not extended over auricle; auricles with three teeth
(fig. 200); constriction of segment III marked (figs.
125a-b). Inner and outer carinae of ventral terga VI
sinuous (figs. 162a-b). Ventral tubercle of abdominal
segment I lower than 30% of its length in lateral view,
with anterior margin convex in lateral view; genital
lobe as high as twice the tubercle height and subquadrate (fig. 240). Sub-basal tooth of male cercus
prominent and angled; dorso-distal crest lower than
cercus width at base, developed at distal 25% of cercus
(fig. 364b); tip medial (fig. 364a). Female cercus maximum width at distal 30%; tip mucronate (fig. 364c).
Dimensions: head width: 10-10.7; HW length: 47.853.9; HW width: 14.8-17.3; HW pterostigma length: 33.8; cerci length: ? 5.8-6.5, / 5.6-6; female cerci
maximum width: 1.4-1.5; total length: 72-80.5.
Biology. – Dr. K. Tennessen, who collected this
species in Bolivia, kindly provided following comments: ‘Exuviae were found on a large boulder of a
small stream, and larvae in slow-flowing places. This
stream was about 1-2 m wide, shaded by vegetation
and the substrate was mostly rocks with some gravel
and mud. The stream was receiving siltation because
the water became very silty when disturbed by our
feet; the hillsides were being clear cut and burned for
agriculture. We saw cattle and hogs in the area. Undoubtedly the species can withstand the siltation, but
may not be able to tolerate removal of the streamside
vegetation. The adult males fly along trails where it is
still wooded, and we saw a couple fly rapidly over the
small stream’. Rare in collections. Larva undescribed.
(fig. 443). – Chile: without locality (MNHN) – Peru:
Huanuco (UMMZ) – Bolivia: La Paz: Sud Yungas
(FSCA*) and Nor Yungas (ANSP; MLPA). Cochabamba:
Chapare (FSCA*; KJTC*), Santa Cruz: Ichilo (FSCA)
and Florida (KJTC).
Rhionaeschna pallipes (Fraser) comb. n.
362-ce, 441-Mp)
Aeschna laticeps Hagen, 1875: 39 (nomen nudum).
Aeshna pallipes Fraser, 1947: 433, 443-445 (description /:
holotype / ARGENTINA: Amaicha del Valle, 26°36’0S
65°55’0W 1800 m (Tafí dept., Tucumán prov.), 01-II1946, A. Willink leg. (IMLA) [examined]); Calvert 1956:14,
123-125; Paulson 1977: 175; Rodrigues Capítulo et al.
1991: 62; Rodrigues Capítulo 1992: 38, 57; Muzón & von
Ellenrieder 1998: 23; Muzón & von Ellenrieder 2001:
157-160 (redescription /, description ?); von Ellenrieder
& Muzón 2003: 95-98 (description larva).
[Aeshna (Marmaraeschna) vigintipunctata. – Calvert 1956:
119-123 (in part); Rodrigues Capítulo & Muzón 1989b:
147-148; Rodrigues Capítulo et al. 1991: 62; Rodrigues
Capítulo 1992: 38, 57 (in part). Misidentification]
[Aeshna (Marmaraeschna) intricata. – Paulson 1977: 175;
Rodrigues Capítulo et al. 1991: 62; Rodrigues Capítulo
1992: 38, 57. Misidentification]
Diagnosis. – The linear frontal carina of R. pallipes
(fig. 7c) is unique within the marmaraeschna-group
(figs. 2-6, 8c). Further distinction from remaining
species is provided under R. brevifrons.
Description. – Clypeus and frons light blue, light
blue spots lateral to light yellow area surrounding Tspot stem; T-stem maximum width at posterior end,
sides converging anteriorly, separated from transverse
arms (fig. 7a); narrow black stripe on frontoclypeal
groove (fig. 7b-c); vertex light blue with posterior
margin black. Frontal carina linear in frontal view (fig.
7c); frons flattened (fig. 7b). Pterothorax pale yellowbrown with 1-2 black spots in mesanepisternum, 2-3
in mesepimeron, 2 in metepisternum and 2-3 in
metepimeron (fig. 46). Abdomen grayish light blue
and light yellow with reddish brown to black spots;
black areas of ventral abdominal terga II not extended
over auricle; auricles with two teeth (fig. 201); constriction of segment III very slight (figs. 124a-d). Inner
and outer carinae of ventral terga VI linear and parallel
to each other (figs. 163a-b). Ventral tubercle of abdominal segment I lower than 30% of its length in lateral view, with anterior margin convex in lateral view;
genital lobe approximately as high as tubercle (fig.
241). Sub-basal tooth of male cercus prominent; dorso-distal crest lower than 50% of cercus width at base,
developed at distal 20% of cercus (fig. 362b); tip on
external margin (fig. 362a). Female cercus maximum
width at distal 66%; tip mucronate (fig. 362c). Dimensions: head width: 8.7-10.4; HW length: 43.350.05; HW width: 13.3-16.1; HW pterostigma length:
2.5-3.4; cerci length: ? 4.1-4.65, / 1.8-2.2; female
cerci maximum width: 0.5; total length: 59-59.5.
Biology. – The species has been found in sympatry
with R. fissifrons in Preandean mountain range localities in Northwestern Argentina. Males were observed
patrolling streams and creeks and females ovipositing
in marginal vegetation. They were also occasionally
seen perching on rocks and vertical banks of the
streams exposed to the sun where they were camouflaged due to their marbled color. Larvae found under
stones and clinging to submerged roots of marginal
vegetation in these streams were described by von Ellenrieder & Muzón (2003). Abundant in the field,
but rare in collections.
85
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441). – Argentina: Buenos Aires (FSCA; MLPA*; USNM),
Santa Fe (MLPA; ZMHB), Mendoza (IADIZA), Córdoba
(IMLA; FSCA; MACN; MCZ), La Rioja (MACN; MLPA),
Catamarca (DRPC), Tucumán (IMLA*), and Salta
(MLPA*; ZMHB; RWGC; UMMZ).
Rhionaeschna vigintipunctata (Ris) comb. n.
365-ce, 443-Mp)
Aeshna vigintipunctata Ris, 1918: 163-166 (description ? /:
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;
Calvert 1956: 119-123 (in part); Paulson 1977: 175; Rodrigues Capítulo et al. 1991: 62 (in part); Rodrigues Capítulo 1992: 38, 57 (in part); Donnelly et al. 1998: 115;
Muzón & von Ellenrieder 1998: 23; Muzón & von Ellenrieder 2001: 160-164 (redescription ? /).
[Aeshna (Marmaraeschna) intricata. – Calvert 1956: 112-119
(in part); Rácenis 1959: 494 (in part). Misidentification]
Diagnosis. – Distinction between R. vigintipunctata and its congeners is found under R. brevicercia, R.
brevifrons, R. intricata and R. obscura.
Description. – Clypeus and frons light blue and yellow, gray blue spots lateral to light yellow area surrounding T-spot stem; T-stem maximum width at
mid-length, sides convex, strongly narrowed at confluence with transverse arms (fig. 8a); wide black stripe
on frontoclypeal groove (fig. 8b-c); vertex yellow with
latero-posterior margins black. Frontal carina with a
slight concavity in frontal view (fig. 8c); frons not flattened (fig. 8b). Pterothorax pale yellow-brown with 1
black spot in mesanepisternum, 3-4 in mesepimeron,
2-3 in metepisternum and 4-5 in metepimeron (fig.
47). Abdomen pale brown to light yellowish green and
grayish light blue with dark reddish brown to black
spots; black areas of ventral abdominal terga II not extended over auricle; auricles with three teeth (fig.
202); constriction of segment III marked (figs. 126ab). Inner and outer carinae of ventral terga VI sinuous
(figs. 164a-b). Ventral tubercle of abdominal segment
I lower than 30% of its length in lateral view, with anterior margin convex in lateral view; genital lobe as
high as twice the tubercle height and subtriangular
(fig. 242). Sub-basal tooth of male cercus prominent
and angled; dorso-distal crest lower than cercus width
at base, developed at distal 30% of cercus (fig. 365b);
tip medial (fig. 365a). Female cercus maximum width
at medial 30%; tip not mucronate (fig. 365c). Dimensions: head width: 9.4-10.3; HW length: 47.2-54.4;
HW width: 14.8-18.8; HW pterostigma length: 2.5-3.7;
cerci length: ? 5.8-6.4, / 4.9-5.65; female cerci maximum width: 1.0-1.1; total length: 72-77.
Biology. – Rhionaeschna vigintipuncata was found
at mountain streams ranging from humid mountain
86
range forest in Argentina and Bolivia to dry Preandean environments. Adults were seen flying along the
streams close to the water. Larva unknown. A common species.
443). – Argentina: Catamarca (DRPC; IMLA; MACN;
Ris 1918), La Rioja (MACN), Tucumán (IMLA; MLPA*;
ZMHB; USNM), Salta (IMLA; MLPA*; RWGC*), and Jujuy
(MLPA; UMMZ) – Bolivia: Cochabamba: Chapare
(FSCA) and Santa Cruz: Ichilo (FSCA*) – Peru: Piura
(Ris 1918) and Amazonas (Calvert 1956).
3. Schizuraeschna group
(= subgenus Schizuraeschna Calvert, 1952).
Cercus inner margin with a ventral process at distal
25%, dorso-distal crest triangular in lateral view, lateral carina concave, and sub-basal tooth low and blunt
(figs. 366-369a-c); two rows of cells between RP1 and
RP2 in HW beginning distal to end of pterostigma (fig.
398). Four species: R. dugesi, R. jalapensis, R. multicolor, R. mutata.
Rhionaeschna dugesi (Calvert) comb. n.
366-ce, 398-wi, 444-Mp)
Aeshna dugesi Calvert, 1905: 180, 184-185 (description ?:
holotype ? MEXICO: Guanajuato, 19º48’0N 104º
20’60W 966 m, A. Dugès leg. (USNM; 66420) [examined]); Martin 1908: 49-50, 83; Calvert 1956: 107-108;
González Soriano & Novelo Gutiérrez 1991: 73, 81, 9394; Novelo Gutiérrez & González Soriano 1991: 114,
141-145; González Soriano 1993: 296; González Soriano
& Novelo Gutiérrez 1996: 164; Dunkle 2000: 51; Needham et al. 2000: 122, 125, 133, 138-139.
Description. – Clypeus and frons light blue to yellow, grayish blue spots lateral to white area surrounding T-spot stem; T-stem narrower than vertex, gradually narrowing anteriorly; no dark stripe on
frontoclypeal groove; latero-posterior margin of vertex
black (figs. 9a-c). Pale stripes light blue to white, complete, and as wide as 30% of their sclerites (fig. 48).
Abdomen reddish brown with light blue and yellow to
greenish yellow spots; PL spots in III-V (figs. 127b, d);
posterior portion of inner carina of ventral terga V-VII
concave in females (fig. 165b). Ventral tubercle of abdominal segment I as high as 40% of its length (fig.
243); distal portion of ventral margin of anterior lamina spine linear (fig. 325); tip of hamular anterior
process pointed in ventral view (fig. 285b); auricles
with two-three teeth (fig. 203). Male cercus with a
small ventral process at distal 25% not projected posteriorly (fig. 366b), distance between base of crest and
posterior base of ventral process shorter than distance
between base of process and tip of cercus, tip not bent
ventrally, dorso-distal crest higher than base of cercus
in lateral view (fig. 366c). Female cercus external margin slightly convex, tip rounded (fig. 366d). Dimensions: head width: 8.7-9; HW length: 44-53; HW
width: 13.5-14.3; HW pterostigma length: 3.3-3.5;
cerci length: ? 5.6-6.1, / 5.8-6.2; female cerci maximum width: 1.2; total length: 65-74.
Diagnosis. – Rhionaeschna dugesi is easily distinguished from other Schizuraeschna species by the male
cercus with ventral process of inner margin not projected posteriorly and tip not projected ventrally (fig.
366b), distal portion of ventral margin of anterior
lamina spine linear (fig. 325), and posterior portion of
inner carina of female ventral terga V-VII concave (fig.
165).
Biology. – Dunkle (2000) reported males patrolling low along edges of stream pools edged with
grass and trees in pine and oak tree zones. Gonzáles &
Novelo (1991) observed females ovipositing in emergent or submerged vegetation of streams in some cases still in tandem. They described the last larval instar
(Novelo & Gonzáles 1991) based on reared larvae
from Durango, MX. A common species.
(fig. 444). – Mexico: Oaxaca (DRPC), Puebla (USNM),
Guanajuato (USNM; Calvert 1905), Zacatecas (USNM),
Baja California Sur (Calvert 1956), Durango (DRPC),
(Novelo & Gonzáles 1991), and Nuevo León (USNM)
– USA: Arizona (LACM; RWGC*; UAIC), Texas (RWGC),
and New Mexico (Needham et al. 2000).
Rhionaeschna jalapensis (Williamson) comb. n.
(figs. 10-he, 49-50-th, 128-ab, 166-te, 204-ge, 246-tg, 286-ha,
326-as, 367-ce, 444-Mp)
Aeshna jalapensis Williamson, 1908: 307-308 (description
?: holotype ? MEXICO: Jalapa, 19°31’60N 96°55’0W
1381 m, IX-1906, P.P. Calvert leg. (ANSP)); Walker 1912:
198; Calvert 1956: 13, 106-107; Machado 1985b: 328;
González Soriano 1993: 296; González Soriano & Novelo Gutiérrez 1996: 164.
[Aeshna multicolor. – Calvert 1905: 180, 183-184 (in part);
Calvert 1907: 400 (in part). Misidentification]
[Aeshna punctata. – Martin 1908: 54 (in part: / from Mexico); Calvert 1956: 82 (in part: records from Mexico);
Santos 1966a (in part: records from Mexico). Misidentification]
Diagnosis. – Rhionaeschna jalapensis can be distinguished from the remaining Schizuraeschna species by
the male cercus bent ventrally only at extreme tip (fig.
367c), and the presence of PL spots only in abdominal
segment III (128b, d). It differs also from the sympatric R. dugesi and R. multicolor by the T-spot stem
wider than vertex (fig. 10a), vs. narrower (figs. 9, 11a).
Description. – Clypeus and frons light blue to yellow, grayish blue spots lateral to white area surrounding T-spot stem; T-stem wider than vertex, gradually
narrowing anteriorly; no dark stripe on frontoclypeal
groove; vertex black with central portion light blue
(figs. 10a-c). Pale stripes light blue to white or yellow;
mesanepisternal complete to incomplete at basal 3050% of mesanepisternum; mesepimeral and
metepimeral complete and as wide as 25% of their
sclerites (figs. 49-50). Abdomen reddish brown with
light blue and yellow to greenish yellow spots; PL spots
in III (figs. 128b, d); posterior portion of inner carina
of ventral terga V-VII linear in female (fig. 166b). Ventral tubercle of abdominal segment I as high as 40% of
its length (fig. 246); distal portion of ventral margin of
anterior lamina spine convex (fig. 326); tip of hamular
anterior process pointed in ventral view (fig. 286b);
auricles with two-three teeth (fig. 204). Male cercus
with ventral process at distal 25% projected posteriorly (fig. 367b), distance between base of crest and posterior base of ventral process shorter than distance between base of process and tip of cercus, only extreme
tip ventrally bent, dorso-distal crest higher than base
of cercus in lateral view (fig. 367c). Female cercus external margin slightly convex, tip pointed (fig. 367d).
Dimensions: head width: 8.5-9.5; HW length: 4245.3; HW width: 12.5-13.8; HW pterostigma length:
3.4-3.5; cerci length: ? 5.7-6.2, / 5.3-5.9; female
cerci maximum width: 1.3; total length: 67-72.
Biology. – Larva described by supposition by
Calvert (1956) based on exuviae from Guatemala,
Costa Rica, and larvae from Cuba and Venezuela. A
common species. Adults were collected at small,
marshy ponds and streams through open and woods;
muddy bottom streams with abundant vegetation;
springs; weedy drainage ditch widening into shallow
marsh; rain ponds choked with grass in shallow areas,
open in center; over pasture; some more-or-less boggy
areas in hollow tiny forest streams to large, open,
rocky rivers. Females were observed ovipositing in
Scirpus and in sedge at and below water level, but
keeping wings dry; males flying very low, bouncing
just over vegetation; and pairs in copula flying along
shore several times and hanging low in sedges or 3.5-4
m up in trees (DRPC).
Remarks. – Calvert (1956) included not full-grown
larvae from Cuba and Venezuela in his description of
the larva. Those records were not included here because no adults of this species are known from these
countries. Since no reliable diagnostic descriptions
exist for half-grown larvae it is not possible to identify the larvae from Venezuela with certainty.
Distribution [8-23°N, 82-105°W, 28-2040 m] (fig.
444). – Panama: Bocas del Toro (Walker 1912) – Costa Rica: Puntarenas (DRPC*), San José (DRPC; USNM),
Cartago (DRPC; USNM; Calvert 1956), Heredia (DRPC;
INBC; USNM), Alajuela (USNM; DRPC), and Guanacaste
(LACM; RWGC*) – Nicaragua: Matagalpa (Beckemeyer
2001) – Honduras: without locality (Förster 2001) –
El Salvador: without locality (Förster 2001) –
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Guatemala: Santa Rosa (Calvert 1956), Suchitepéquez
(USNM), Retalhuleu (Calvert 1956), Guatemala
(MLPA*; Calvert 1956), Chimaltenango (RWGC), Solola
(DRPC), Quezaltenango (Calvert 1956), Zacapa
(Calvert 1956), Alta Verapaz (DRPC), and Huehuetenango (DRPC) – Mexico: Chiapas (DRPC; RWGC;
USNM), Guerrero (Calvert 1905), Veracruz (DRPC;
RWGC; USNM; Calvert 1905; Williamson 1908), Puebla
(USNM), Morelos (RWGC*; USNM), Michoacan (DRPC),
Jalisco (USNM), and Durango (DRPC; RWGC).
Rhionaeschna multicolor (Hagen) comb. n.
327-as, 368-ce, 445-Mp)
Aeshna multicolor Hagen, 1861: 121-122 (description ? /:
lectotype ? (here designated) U.S.A.: Pecos River, Western
Texas (Capt. Pope exp.) (MCZ; 8833) [examined: lectotype ?, syntypes /]); Calvert 1905: 180, 183-184 (in
part); Calvert 1907: 400 (in part); Williamson 1908: 301;
Martin 1908: 48-49, 83; Walker 1912: 62, 66, 69, 71,
190-198 (redescription adults, description larva); Calvert
1956: 13, 102-106; Walker 1958: 51-52, 55, 117-121
(redescription ? / larva); González Soriano & Novelo
Gutiérrez 1991: 73, 80, 91; Novelo Gutiérrez & González
Soriano 1991: 114, 145-146; González Soriano 1993:
296; González Soriano & Novelo Gutiérrez 1996: 164;
Dunkle 2000: 50; Needham et al. 2000: 122, 126, 133,
144-145.
Aeschna furcifera Karsch, 1891: 310.
Diagnosis. – Rhionaeschna multicolor and R. mutata
differ from R. dugesi by the long ventral process of
male cercus (figs. 368-369c vs. fig. 366c) and the linear inner lateral carina of ventral terga V-VII (figs. 167168 vs. fig. 165), and from R. jalapensis by the entire
tip of male cercus ventrally bent (figs. 368-369c vs. fig.
367c) and presence of pale spots in abdominal segments III-V (figs. 129-130b, d vs. fig. 127b, d). They
differ from each other by the height of ventral tubercle
I (as high as 50% of its length in R. multicolor, fig. 244;
as high as 30% of its length in R. mutata, fig. 245),
distance separating the posterior base of the ventral
process of male cercus from base of crest and tip of cercus (closer to base of crest in R. multicolor, fig. 368c; to
tip of cercus in R. mutata, fig. 369c) and height of dorso-distal crest (higher than base width in R. multicolor,
fig. 368c; as high as base width in R. mutata, fig.
369c), external margin of female cercus (slightly convex in R. multicolor, fig. 368d; linear in R. mutata, fig.
369d), and T-spot stem width (narrower than vertex
in R. multicolor, fig. 11a; wider in R. mutata, fig. 12a).
Description. – Clypeus and frons light blue to yellow, grayish blue spots lateral to white area surrounding T-spot stem; T-stem narrower than vertex, gradually narrowing anteriorly; no dark stripe on
frontoclypeal groove; vertex light blue or yellow with
latero-posterior margins black (figs11a-c). Pale stripes
light blue to white or yellow; mesanepisternal com88
plete to incomplete at basal 30-50% of mesanepisternum; mesepimeral and metepimeral complete and as
wide as 25% of their sclerites (figs. 51-52). Abdomen
reddish brown with light blue and yellow to greenish
yellow spots; PL spots in III-V (figs. 129b, d); posterior
portion of inner carina of ventral terga V-VII linear in
female (fig. 167b). Ventral tubercle of abdominal segment I as high as 50% of its length (fig. 244); distal
portion of ventral margin of anterior lamina spine
convex (fig. 327); tip of hamular anterior process
pointed in ventral view (fig. 287b); auricles with twothree teeth (fig. 205). Male cercus with ventral process
at distal 25% projected posteriorly (fig. 368b), distance between base of crest and posterior base of ventral process shorter than distance between base of
process and tip of cercus, whole cercus tip ventrally
bent, dorso-distal crest higher than base of cercus in
lateral view (fig. 368c). Female cercus external margin
slightly convex, tip pointed (fig. 368d).
Dimensions: head width: 9-10; HW length: 42-45;
HW width: 13-13.8; HW pterostigma length: 3.5-3.8;
cerci length: ? 6.4-6.6, / 5.5-5.8; female cerci maximum width: 1.2; total length: 65-72.
Biology. – Adults feed over open fields from before
sunrise to after sunset, hanging under trees and bush
during hot midday hours; males patrol large areas of
water in an irregular pattern (Dunkle 2000). They reproduce mainly in temporary pools, ponds, lakes, and
slow streams, with emergent or floating plants
(González & Novelo 1991). One of the most common species of Aeshnidae in western United States.
Larva described by supposition by Walker (1912),
based on exuviae from Washington and by Musser
(1962), based on larvae from Utah, USA.
Remarks. – This species was described (Hagen
1861: 122) from an unspecified number of specimens
from ‘Pecos River, Western Texas (Capt. Pope); Upper Missouri; Mexico; Cordova (Saussure)’ without
mention of the repository of type specimens. From the
original type series, only four specimens are still in the
Hagen collection in the MCZ (one male and two females from Pecos River and one female from Upper
Missouri). Williamson (1908) established that the female from Upper Missouri belonged to a different
species and was included in Walker’s description of
Aeshna interrupta lineata (Walker 1912). Although the
locality mentioned by Hagen for the lectotype is the
Pecos River in western Texas (32°N 104°W), it was
apparently collected in the vicinity of the present town
of Roswell, New Mexico, where Pope’s expedition occurred in 1854-1855 (see discussion in Garrison
1994: 220).
445). – Mexico: Puebla (Walker 1912), Michoacan
(LACM; Calvert 1956), Distrito Federal (Calvert
1956), Hidalgo (RWGC), Durango (Novelo &
Gonzáles 1991), Baja California Sur (UAIC; Calvert
1956; Walker 1912), Jalisco (DRPC) – U.S.A.: Washington (Walker 1912), Oregon (RWGC; Walker 1912),
Idaho (RWGC; Walker 1912), Colorado (Walker
1912), Utah (Walker 1912), California (RWGC*; UAIC;
Walker 1912), Arizona (RWGC*; UAIC), New Mexico
(RWGC; Hagen 1861; Walker 1912), Texas (RWGC;
Hagen 1861; Walker 1912), Nebraska (RWGC*),
Nevada (RWGC), Massachusetts (Needham et al.
2000), Iowa (Needham et al. 2000), Kansas (Needham et al. 2000), Montana (Needham et al. 2000),
Oklahoma (Needham et al. 2000), South Dakota
(Needham et al. 2000), and Wyoming (Needham et
al. 2000) – Canada: British Columbia (Walker 1912)
and Alberta (Needham et al. 2000).
Rhionaeschna mutata (Hagen) comb. n.
369-ce, 445-Mp)
Aeshna mutata Hagen, 1861: 124-125 (description /: holotype / North America (MCZ; 8832)); Williamson 1908:
301-306; Walker 1912: 62, 66, 71 198-202 (redescription ? /); Walker 1958: 52, 55, 121-125 (redescription? /, description larva); Beatty & Beatty: 1969: 149151; Dunkle 2000: 51-52; Needham et al. 2000: 122,
125, 133, 145-146.
[Aeshna multicolor. – Calvert 1905: 180, 183-184 (in part);
Calvert 1907: 400 (in part); Martin 1908: 48 (in part).
Misidentification]
Diagnosis. – Discrimination of this species from
the closely related R. multicolor is provided under that
species.
Description. – Clypeus and frons light blue to yellow, grayish blue spots lateral to white area surrounding T-spot stem; T-stem wider than vertex, gradually
narrowing anteriorly; no dark stripe on frontoclypeal
groove; vertex black with central portion light blue
(figs. 12a-c). Pale stripes light blue to white or yellow;
mesanepisternal complete to incomplete at basal 3050% of mesanepisternum; mesepimeral and
metepimeral complete and as wide as 25% of their
sclerites (fig. 53). Abdomen reddish brown with light
blue and yellow to greenish yellow spots; PL spots in
III-V (figs. 130b, d); posterior portion of inner carina
of ventral terga V-VII linear (fig. 168b). Ventral tubercle of abdominal segment I as high as 30% of its length
(fig. 245); distal portion of ventral margin of anterior
lamina spine convex (fig. 328); tip of hamular anterior process pointed in ventral view (fig. 288b); auricles
with two-three teeth (fig. 206). Male cercus with ventral process at distal 25% projected posteriorly (369b),
distance between base of crest and apex of ventral
process longer than distance between apex of process
and tip of cercus, whole cercus tip ventrally bent, dorso-distal crest as high as base of cercus in lateral view
(fig. 369c). Female cercus outer margin linear, tip an-
gled (fig. 369d). Dimensions: head width: 9-10; HW
length: 44-51; HW width: 13-13.9; HW pterostigma
length: 3.6-4.1; cerci length: ? 5.8-6.3, / 6.5-6.7; female cerci maximum width: 1.4; total length: 67-75.
Biology. – Males patrol low over vegetation with a
leisurely erratic flight. Females usually perch on water
lily flowers to lay eggs in the underwater part of the
stem. They breed in fishless ponds, usually with water
lilies, and occasionally in bog ponds (Dunkle 2000). A
rare species. The larva was described by Walker (1958).
Remarks. – According to the original description,
the holotype female was deposited at the NMW. However, Calvert (1905) found a female labeled as type at
the MCZ and considered it the holotype, a decision
with which Williamson (1908) agreed.
445). – U.S.A.: Ohio (Walker 1912), Indiana (Walker 1912), Pennsylvania (MLPA; RWGC*), Ohio (RWGC),
New Jersey (RWGC*), Connecticut (Needham et al.
2000), Illinois (Needham et al. 2000), Kentucky
(Needham et al. 2000), Maine (Needham et al. 2000),
Maryland (Needham et al. 2000), Michigan (Needham et al. 2000), Missouri (Needham et al. 2000),
New Hampshire (Needham et al. 2000), New York
(Needham et al. 2000), Rhode Island (Needham et al.
2000), Virginia (Needham et al. 2000), West Virginia
(Needham et al. 2000), and Wisconsin (Needham et
al. 2000) – Canada: Ontario (Needham et al. 2000).
4. Neureclipa group
(= subgenus Neureclipa Calvert, 1952)
Supratriangles usually free, sometimes one or two
with 1 or 2 crossveins (in 2-10% of specimens examined); two rows of cells between RP1 and RP2 in HW
beginning at distal end of pterostigma or distal to it
(fig. 399); male cerci with high dorso-distal crest (as
high or higher than width of cercus at base), sub-basal
tooth prominent, and lateral carina concave (figs.
370-374). Five species: R. absoluta, R. bonariensis, R.
diffinis, R. elsia, R. galapagoensis.
Rhionaeschna absoluta (Calvert) comb. n.
329-as, 370-ce, 446-Mp)
Aeshna diffinis absoluta Calvert 1952: 258-260 (description
? /: holotype ? PERU: vicinity of Concepción, 07/08IV-1935, F. Woytkowski leg. (UMMZ) [examined: holotype ?, allotype /, paratypes ? /]); Calvert 1956: 15,
133-134; Rácenis 1959: 494.
Aeshna absoluta Jurzitza 1990a: 353-372; Rodrigues Capítulo 1992: 37, 56; Muzón 1995: 5 (in part); Muzón 1997:
127-128, 132-133; Muzón & von Ellenrieder 1998: 23;
von Ellenrieder 2001a: 301-305; b: 423-426, 431-433.
[Aeshna diffinis. – Ris 1904: 24, 26-29 (in part); Ris 1908:
523, 526-527 (in part); Ris 1913: 84-85 (in part); Navás
1920d: 53; Navás 1930a: 125; Fraser 1947: 433, 446; Ro-
89
T  E,  146, 2003
drigues Capítulo 1980: 9, 16-17, 20; Abenante & Philippi 1982: 151; Rodrigues Capítulo & Muzón 1989b: 147;
Rodrigues Capítulo et al. 1991: 62, 66 (in part); Rodrigues Capítulo 1992: 37, 57 (in part); Pérez D’A. &
Mutschke 1993-94: 63-68; Muzón 1995: 5-6 (in part);
Muzón & von Ellenrieder 1998: 23 (in part). Misidentification]
[Aeshna diffinis diffinis. – Calvert 1956: 127, 233 (in part);
Rácenis 1959: 494. Misidentification]
[Aeshna elsia. – Rodrigues Capítulo 1992: 37, 57; Daigle et
al. 1997: 12 (in part). Misidentification]
Diagnosis. – Rhionaeschna absoluta and R. diffinis
can be distinguished from the other Neureclipa species
by the stem of T-spot with concave sides (figs. 13,
15a), and the dorso-distal crest of male cerci higher
than cerci width at base (figs. 370, 372b). They differ
from each other by the comparatively narrower ventral
terga (ratio length/width of IV 4.2-4.5) and maximum
width of V-VI occupies the distal portion in R. absoluta (fig. 169). In R. diffinis the ventral terga are comparatively wider (ratio length/width of IV 2.4-2.6) and
maximum width of V-VI occupies the basal portion
(fig. 171). Females differ by the dorsal color pattern of
second abdominal segment; longitudinal black stripe
between transverse carina and posterior margin in R.
absoluta (fig. 131c) but only at posterior margin in R.
diffinis (fig. 133c), and shape of cercus; tips pointed in
R. absoluta, fig. 370c, and rounded in R. diffinis, 372c.
Description. – Clypeal lobes angled (fig. 13c);
clypeus light blue or yellow, frons yellow, grayish
green spots lateral to yellow area surrounding T-spot
stem; T-stem widening posteriorly, sides slightly concave; vertex yellow with latero-posterior margins black
(fig. 13a); wide black stripe on frontoclypeal groove;
wide black stripe on fronto-ocular groove (figs. 13bc). Pale mesanepisternal stripes at basal 25-33% of
mesanepisternum to absent; mesepimeral and
metepimeral stripes light blue to yellow, linear and
narrow (except in teneral specimens where they are
wide: fig. 54), complete to absent (figs. 55-59). Membranule dark except basal 30% pale. Abdomen reddish
brown with light blue and yellow spots; female segment II with a medio longitudinal dorsal black stripe
between transverse carina and posterior margin (fig.
131c). Abdominal ventral terga narrow (ratio length/
width of V at basal 25% higher than 4), maximum
width of V-VI at distal 66%, basal 30% of inner and
outer lateral carinae of IV concave (figs. 169a-b). Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface (fig.
247); dorsal margin of anterior lamina spine linear
(fig. 329); tip of hamular anterior process pointed in
ventral view (fig. 289); auricles with two teeth (fig.
207). Male cercus lacking pale basal spot in outer surface (fig. 370a); dorso-distal crest rising suddenly,
higher than base of cercus in lateral view (fig. 370b).
Tip of female cercus pointed (fig. 370c). Dimensions:
90
head width: 6.9-8.4; HW length: 32-41.1; HW width:
10.5-13.1; HW pterostigma length: 2.8-4.3; cerci
length: ? 4.3-5.1, / 3.8-4.9; female cerci maximum
width: 0.9; total length: 43.1-59.8.
Biology. – Rhionaeschna absoluta occupies one of
the largest distribution areas within Rhionaeschna,
breeding in a wide variety of environments from temporary ponds and pools to creeks, streams and small
rivers. I collected foraging adults flying along an Atlantic beach in Chubut province. They patrolled
small temporary ponds on sides of a road, creeks and
small streams in the Patagonian steppe, dams for irrigation and banks of small rivers in Northwestern Argentina. Larvae were found under stones or crawling
among aquatic vegetation (von Ellenrieder 2001b). A
common species.
446). – Argentina: Santa Cruz (Muzón 1995),
Chubut (DRPC; IMLA; MACN; MLPA*; RWGC), Neuquén
(DRPC; MLPA*; RWGC; USNM; Muzón 1995), Río Negro (DRPC; MLPA*; RWGC; USNM), Buenos Aires (DRPC;
MACN; MLPA*; RWGC; USNM), La Pampa (MLPA), Santa
Fe (MACN), Entre Ríos (MLPA), Mendoza (DRPC; IMLA;
IADIZA; MLPA; USNM; Ris 1908; Fraser 1947; Jurzitza
1990a), San Juan (IMLA); La Rioja (MACN; MLPA),
Catamarca (DRPC), Santiago del Estero (MLPA), Córdoba (Navás 1930a), Tucumán (DRPC; IMLA; MLPA;
RWGC), Salta (IMLA; MLPA*; USNM), and Jujuy (IMLA;
MLPA) – Chile: Magallanes: Magallanes (Perez D’A. &
Mutschke 1993-94), and Santiago: Santiago (MNNS),
Biobío: Arauco (UMMZ) – Uruguay: San José (Calvert
1956), and Montevideo (ANSP; Calvert 1956) – Bolivia: La Paz: Sud Yungas (RWGC) - Peru: Lima (DRPC),
Junín (RWGC; UMMZ; Calvert 1952; 1956), Cuzco
(UMMZ; USNM; Calvert 1956), Ayacucho (IMLA;
UMMZ), and Arequipa (Calvert 1952; 1956) Ecuador: Imbabura (KJTC).
Rhionaeschna bonariensis (Rambur) comb. n.
371-ce, 399-wi, 447-Mp)
Aeshna bonariensis Rambur, 1842: 204 (description ? /:
lectotype ? (here designated) ARGENTINA: Buenos Aires
(ISNB) [examined: lectotype ?, paralectotypes /]); Hagen
1861: 314; Hagen 1875: 39; Ris 1904: 24-25; Martin
1908: 51-52; Ris 1908: 523, 525-526; Calvert 1909: 221;
Martin 1911: 12; Navás 1911: 476, 478; Cockerell 1913:
580; Ris 1913: 85; Navás 1917a: 187; Ris 1918: 158;
Navás 1920a: 152; Navás 1920c: 267; Martin 1921: 23;
Campion 1922: 292-293; Martin 1923: 109; Navás 1927:
23; Gazulla & Ruiz 1928: 290; Navás 1929c: 220; Pirión
1933: 82; Fraser 1947: 433, 448; Calvert 1952: 257-258;
Herrera et al. 1955-66: 81; Calvert 1956: 14, 140-144,
227; Fraser 1957: 159; Martins Costa 1971: 194; Paulson
1977: 175; Abenante 1978: 29-48; Abenante 1980: 105149; Rodrigues Capítulo 1980: 1-21; Abenante & Philippi 1982: 151; Jurzitza 1989: 7; Jurzitza 1990a: 353-372;
Rodrigues Capítulo et al. 1991: 62, 66; Watson 1992:
455, 657-462; Pitzke-Widdig 1992: 116, 122; Rodrigues
Capítulo 1992: 37, 56; Mola & Papeschi 1994: 185-188;
Mola 1995: 47-54; Muzón 1995: 5; Muzón & von Ellenrieder 1997: 146; Carvalho & Nessimian 1998: 7; Muzón
& von Ellenrieder 1998: 23; Martins Costa & Santos
1999: 4; Martins Costa, et al. 2000: 13; von Ellenrieder
2001a: 301, 305-308; b: 424, 426-427, 431-433.
Aeschna dicrostigma Selys in Martin, 1908: 53 (nomen
nudum).
Neureclipa bonariensis. – Navás 1911: 476, 478.
Aeshna bonaerensis var. lutea Navás, 1920b: 11; Navás
1920c: 267; Navás 1928: 340.
Diagnosis. – Rhionaeschna bonariensis shares with R.
absoluta and R. diffinis a linear dorsal margin of anterior lamina spine (figs. 329-331) and ventral tubercle
of I with numerous denticles (figs. 247-249), and with
R. elsia and R. galapagoensis the dorso-distal crest of
cercus as high as width of cercus at base (figs. 371,
373-374b) and a complete medio-dorsal longitudinal
yellow stripe of female abdominal segment II (figs.
132-134c). The narrow stripe on fronto-ocular groove
(fig. 14b), parallel sided T-spot (fig. 14a) and pale spot
on outer base of male cercus (fig. 371a-b) distinguish
R. bonariensis from all other Neureclipa species.
Description. – Clypeal lobes angled (fig. 14c);
clypeus and frons light blue and yellow, grayish blue
spots lateral to yellow area surrounding T-spot stem;
T-stem usually parallel sided, in some cases slightly
narrowed anteriorly; vertex yellow with latero-posterior margins black (fig. 14a); no dark stripe on frontoclypeal groove; black stripe on fronto-ocular groove
narrow (figs. 14b-c). Pale mesanepisternal stripes at
basal 25-33% of mesanepisternum to absent;
mesepimeral and metepimeral stripes yellow to light
blue, linear, complete and narrow (fig. 60). Membranule dark except basal 20% pale. Abdomen pale
reddish brown with light blue and yellow spots; female
segment II with a complete medio longitudinal dorsal
yellow stripe (figs. 132c). Abdominal ventral terga
narrow (ratio length/width of V at basal 25% higher
than 4), maximum width of V-VI at distal 66%, basal
30% of inner and outer lateral carinae of IV concave
(figs. 170a-b). Ventral tubercle of abdominal segment
I bearing numerous denticles spread over its posterior
surface (fig. 248); dorsal margin of anterior lamina
spine linear (fig. 330); tip of hamular anterior process
rounded in ventral view (fig. 290b); auricles with two
teeth (fig. 208). Male cercus with a pale basal spot in
outer surface (fig. 371a); dorso-distal crest rising gradually, as high as base of cercus in lateral view (fig.
371b). Tip of female cercus pointed (fig. 371c). Dimensions: head width: 6.9-8; HW length: 33.9-39.7;
HW width: 10.5-13; HW pterostigma length: 3.2-4.2;
cerci length: ? 4.5-5.2, / 4.5-5.7; female cerci maximum width: 0.95; total length: 50-57.8.
Biology. – The larva was described by Abenante
(1978), Rodrigues Capítulo (1980) and von Ellenrieder (2001b). It breeds in ponds and temporary
pools. Adults are common in grasslands of central Argentina, where they are usually found flying together
with Rhionaeschna confusa. I observed a mass migration
of hundreds of teneral R. bonariensis and R. confusa
perching on bushes and grasses near the shore of the
Rio de La Plata (February 1998). An abundant species.
447). – Argentina: Río Negro (MLPA; Hagen 1875),
Buenos Aires (DRPC; MACN; MLPA*; RWGC; USNM;
Campion 1922; Navás 1927), Entre Ríos (MACN;
MLPA; USNM; Fraser 1947), Mendoza (Ris 1908), San
Juan (Calvert 1956), Córdoba (MACN; MLPA; UMMZ;
Calvert 1956; Navás 1929c), Santa Fé (MACN; MLPA;
USNM), Santiago del Estero (MLPA*; Navás 1920b; Ris
1913); Chaco (Calvert 1956; Ris 1913), Formosa
(IMLA; FCN), Catamarca (RWGC; Ris 1918), La Rioja
(MLPA; MACN), Tucumán (IMLA; Calvert 1956), Salta
(IMLA; MLPA*; USNM; Pitzke-Widdig 1992), Jujuy
(Calvert 1956), Corrientes (MACN; MLPA*; Calvert
1956), and Misiones (MLPA; USNM) – Uruguay: San
José (Calvert 1956), Montevideo (MNRJ; Abenante
1978; Calvert 1956; Hagen 1875; Ris 1904; 1908),
Colonia (Mola 1995; Mola & Papeshi 1994), and
Florida (UMMZ) – Brazil: Rio Grande do Sul (DRPC;
MNRJ; RMNH; USNM; ZMH; Hagen 1861; Martins Costa 1971), Santa Catarina (Calvert 1956), Paraná
(MNRJ; Calvert 1956), Sâo Paulo (DRPC; MNRJ; USNM;
Calvert 1956; Martins Costa et al. 2000), Minas
Gerais (MNRJ; Calvert 1909), Río de Janeiro (Carvalho
& Nessimian 1998; Martins Costa & Santos 1999),
and Goias (MNRJ) – Paraguay: Paraguarí (Calvert
1956; RWGC), Central (DRPC) – Bolivia: Santa Cruz
dept.: Florida (RWGC) – Chile: Araucanía: Cautín
(Herrera et al. 1955-66; Jurzitza 1989), and Valparaiso: Quillota (Pirión 1933) and Valparaiso (MNNS;
Gazulla & Ruiz 1928; Jurzitza 1989; Martin 1923).
Rhionaeschna diffinis (Rambur) comb. n.
331-as, 372-ce, 447-Mp)
Aeshna diffinis Rambur, 1842: 203-204 (description ? /:
lectotype / (here designated) Chile (ISNB) [examined]);
Hagen 1861: 314; Hagen 1875: 38-39 (in part); Selys
1895: 61; Porter 1897: 13; Porter 1899: 181; Ris 1904:
24, 26-29 (in part: a form); Martin 1908: 43-44, fig. 40
(in part); Ris 1908: 523, 526-527 (in part); Navás 1917b:
38; Campion 1922: 292-293; Martin 1923: 109; Pirión
1928: 96-97; Gazulla & Ruiz 1928: 290; Navás 1929a:
145; Navás 1929b: 326; Navás 1930b: 350; Pirión 1933:
81; Ureta 1935: 93; Needham & Bullock 1943: 358-359;
Herrera et al. 1955-66: 81; Peña 1962: 4; Fraser 1957:
159; Paulson 1977: 175 (in part); Davies & Tobin 1985:
3; Jurzitza 1990a: 353-372, figs. 2-6; Rodrigues Capítulo
et al. 1991: 62, 66 (in part); Rodrigues Capítulo 1992: 37,
57; fig. 175a, b (in part); Muzón 1995: 5-6 (in part);
91
T  E,  146, 2003
Muzón 1997: 127-128, 132-133; Muzón & von Ellenrieder 1997: 146; Muzón & von Ellenrieder 1998: 23 (in
part); von Ellenrieder 2001a: 301, 309-312; b: 424, 427428, 431-433.
Aeschna configurata Hagen, 1861: 314 (nomen nudum).
[Aeschna bonariensis. – Bolivar 1884: 5-6. Misidentification]
Aeshna diffinis diffinis. – Calvert 1952: 258; Calvert 1956:
14, 126-133, 201-203, 232 (in part). Böttger & Jurzitza
1967: 35-37; Jurzitza 1975: 10-11; Jurzitza 1989: 8-9 (in
part); Watson 1992: 455, 657-462.
[Aeshna absoluta. – Muzón 1995: 5 (in part). Misidentification]
Diagnosis. – Rhionaeschna diffinis is distinguished
from all the other Neureclipa species by the wide ventral terga (fig. 171). Further differences are discussed
under R. absoluta.
Description. – Clypeal lobes angled (15c); clypeus
and frons light blue to yellow, grayish green spots lateral to yellow area surrounding T-spot stem; T-stem
widening posteriorly, sides slightly concave; vertex yellow with latero-posterior margins black (fig. 15a);
wide black stripe on frontoclypeal groove; wide black
stripe on fronto-ocular groove (Figs15b-c). Pale
mesanepisternal stripes at basal 25-33% of mesanepisternum to absent; mesepimeral and metepimeral
stripes yellow to light blue, linear and narrow (except
in teneral specimens where they are wide, figs. 61-62),
complete to absent (figs. 63-67). Membranule dark
except basal 30% pale. Abdomen reddish brown to
black with light blue and yellow or greenish yellow
spots; female segment II with a medio longitudinal
dorsal black stripe at posterior 25% of segment (figs.
133c). Abdominal ventral terga wide (ratio length/
width of V at basal 25% less than 3.5 mm), maximum
width of V-VI at basal 30%, basal 30% of inner lateral
carinae of IV concave, of outer convex (figs. 171a-b).
Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface (fig.
249); dorsal margin of anterior lamina spine linear
(fig. 331); tip of hamular anterior process pointed in
ventral view (fig. 291b); auricles with two teeth (fig.
209). Male cercus lacking pale basal spot in outer surface (fig. 372a); dorso-distal crest rising suddenly,
higher than base of cercus in lateral view (fig. 372b).
Tip of female cercus rounded (fig. 372c). Dimensions:
head width: 7.6-8.4; HW length: 34.2-39.4; HW width:
11-13.7; HW pterostigma length: 2.5-3.7; cerci length:
? 4.3-5.3, / 3.4-4.3; female cerci maximum width:
0.95; total length: 49.6-59.9.
Biology. – I have seen adults flying along streams,
small rivers, and shores of lakes in Southern Argentina
and Chile, where the species is common and usually
found with R. variegata. Calvert (1956) provided a description of the larva based on material from Chile,
and von Ellenrieder (2001b) provided an additional
description based on reared specimens from Neuquén
92
and Río Negro, Argentina. An abundant species.
Remarks. – The description of R. diffinis (Rambur
1842) mentioned only female characters and two females with type labels were found in the collection of
the ISNB. The two females belong to two different
species; the one here designated as lectotype is in poor
condition and has only head and part of thorax, and
has the locality label of ‘Chili’ mentioned by Rambur
in his description. The paralectotype female, which
belongs to R. absoluta, is labeled ‘Ae. diffinis?’ in Rambur’s hand.
447). – Chile: Aisén: Aisén (Jurzitza 1989; Ureta
1935), Coihaique (MLPA; RWGC; Calvert 1956), and
General Carrera (MNNS), De los Lagos region: Valdivia
(IEUM; MNNS; MLPA*; RWGC; USNM; Ris 1904; Böttger
& Jurzitza 1967; Jurzitza 1975; 1989; 1990a; Navás
1930b), Osorno (IEUM; MNNS; MLPA; USNM), Llanquihue (IEUM; MLPA; MNNS; USNM), Palena (UMMZ;
RWGC), and Chiloe (IEUM; UMMZ; USNM), Araucanía:
Cautín (IEUM; RWGC; USNM), and Malleco (IEUM;
MLPA*; RWGC; UMMZ; USNM; Calvert 1956; Navás
1929b), Biobío: Arauco (IEUM; MLPA*; UMMZ; USNM;
Watson 1992), Biobío (IEUM; UMMZ), Concepción
(RWGC; USNM; ZMH; Calvert 1956; Jurzitza 1975), and
Ñuble (IEUM; UMMZ; USNM; Navás 1929a), Maule:
Linares (IEUM; MNNS; RMNH; UMMZ; USNM),
Cauquenes (UMMZ; USNM), Talca (IEUM; UMMZ;
USNM), and Curicó (IEUM; MNNS; RWGC; UMMZ), General O’Higgins: Colchagua (UMMZ; USNM; Jurzitza
1975; 1989), Cardenal Caro (IEUM), and Cachapoal
(IEUM; MNNS; UMMZ), Santiago: Maipo (IEUM; MNNS;
UMMZ), Melipilla (UMMZ), Cordillera (IEUM; UMMZ;
USNM), Talagante (IEUM), Santiago (IEUM; MNNS;
UMMZ; USNM), and Chacabuco (IEUM; MNNS; MNNS),
Valparaiso: San Antonio (IEUM; UMMZ), Valparaiso
(IEUM; MNNS), Quillota (RWGC; UMMZ), and Libertador Aconcagua (UMMZ), Coquimbo: Coquimbo
(RWGC), Choapa (IEUM; UMMZ), Limarí (IEUM; MNNS),
and Elqui (MNNS), and Atacama: Huasco (ZMH). – Argentina: Chubut (IMLA; MLPA*; RWGC*; USNM; Campion 1922; Muzón 1995), Río Negro (LACM; MLPA;
USNM; Muzón 1995), and Neuquén (MLPA*; USNM;
Muzón 1995).
Rhionaeschna elsia (Calvert) comb. n.
332-as, 373-ce, 448-Mp)
Aeshna (Neureclipa) elsia Calvert, 1952: 260-262 (description ? /: holotype ? PERU: vicinity of Pacasmayo,
7°24’0S 79°34’0W 7 m, 20-V-1936, F. Woytkowski leg.
(UMMZ) [examined: holotype ?, allotype /, paratypes ?
(UMMZ; RWGC)]); Calvert 1956: 14, 134-137; Rácenis
1959: 494; Paulson 1977: 175; Davies & Tobin 1985: 3;
Jurzitza 1989: 9; Daigle et al. 1997: 12 (in part); von Ellenrieder 2000: 353-357 (redescription ? /).
Diagnosis. – Rhionaeschna elsia and R. galapagoensis
are distinguished from the remaining Neureclipa
species by the T-stem with convex sides (figs. 16-17a),
clypeal lobes rounded (figs. 16-17c) and dorsal margin
of anterior lamina spine concave (figs. 332-333). They
differ from each other in (character states of R. elsia
mentioned first): shape of dorso-distal crest of male
cercus (rising suddenly, fig. 373b vs. gradually, fig.
374b), tip of female cercus (pointed, fig. 373c vs.
rounded, fig. 374c), tip of hamular anterior process in
posterior and ventral view (pointed, fig. 292 vs.
rounded, fig. 293), and color of frontoclypeal groove
(lacking black stripe, fig. 16b vs. bearing black stripe,
fig. 17), and membranule (dark with basal 30% pale
vs. dark with external margin pale).
Description. – Clypeal lobes rounded (fig. 16c);
clypeus and frons light blue to light yellow, grayish
green spots lateral to yellow area surrounding T-spot
stem; T-stem widening posteriorly, with convex sides;
vertex yellow with latero-posterior margins black (fig.
16a); no dark stripe on frontoclypeal groove; black
stripe on fronto-ocular groove wide (figs. 16b-c). Pale
mesanepisternal stripes at basal 25% of mesanepisternum to absent; mesepimeral and metepimeral stripes
light blue to yellow and linear, from complete to reduced to basal spots (figs. 68-73). Membranule dark
except basal 30% pale. Abdomen pale reddish brown
with light blue and yellow spots; female segment II with
a complete medio longitudinal dorsal yellow stripe (fig.
134c). Abdominal ventral terga narrow (ratio length/
width of V at basal 25% higher than 4), maximum
width of V-VI at distal 0.66, basal 30% of inner and
outer lateral carinae of IV concave (figs. 172a-b). Ventral tubercle of abdominal segment I bearing few denticles (10 or less) restricted to its apex (fig. 250); dorsal
margin of anterior lamina spine concave (fig. 332); tip
of hamular anterior process rounded in posterior and
ventral view (fig. 292b); auricles with two teeth (fig.
210). Male cercus lacking pale basal spot in outer surface (fig. 373a); dorso-distal crest rising suddenly, as
high as base of cercus in lateral view (fig. 373b). Tip of
female cercus pointed (fig. 373c). Dimensions: head
width: 7.8-8.5; HW length: 35.8-39; HW width: 1213.6; HW pterostigma length: 2.8-3.4; cerci length: ?
4.3-4.6, / 4.2-4.5; female cerci maximum width:
0.65; total length: 49.85-53.75.
Biology. – The type series was collected at a pond
between sand dunes in Peru (Calvert 1952). Rare in
collections. Larva is still unknown.
Remarks. – Calvert’s (1952; 1956) records from
Southern Chile are considered doubtful and have not
been included here, since no specimens have been
found among the abundant material examined from
this area.
448). – Chile: Tarapacá: Arica (IEUM; MNHN; UMMZ;
Calvert 1952, 1956), and Iquique (Calvert 1952,
1956). Not confirmed records: Maule: Curicó
(Calvert 1952, 1956) and Talca (Calvert 1952, 1956)
– Peru: Arequipa (RWGC), Lima (IMLA; UMMZ*; USNM;
Calvert 1952, 1956), Huanuco (Calvert 1956), and
La Libertad (IMLA; RWGC*; USNM; Calvert 1952,
1956) – Ecuador: Imbabura (KJTC).
Rhionaeschna galapagoensis (Currie) comb. n.
333-as, 374-ce, 448-Mp)
Aeshna galapagoensis Currie, 1901: 382-385 (description ?
/: lectotype ? ECUADOR: Chatham Island, Islas Galápagos, 0°49’60S 89°25’60W 143 m, 26-V-1899 (USNM)
[examined: lectotype ?, paralectotype /]); Needham
1904: 695-696 (description larva); Martin 1908: 50, 84;
Campos 1922: 6, 33; Calvert 1956: 14, 137-140; Belle
1991: 2.
Diagnosis. – See under R. elsia.
Description. – Clypeal lobes rounded (fig. 17c);
clypeus and frons light blue to pale brown, brown
T-stem widening posteriorly, with convex sides; vertex
yellow or light blue with latero-posterior margins
black (fig. 17a); black stripe on frontoclypeal groove;
wide black stripe on fronto-ocular groove (figs. 17bc). Pale mesanepisternal stripes absent; mesepimeral
and metepimeral stripes light blue, complete to absent
(figs. 74-75). Membranule dark except basal 20-25%
pale. Abdomen dark brown with light blue spots; female segment II with a complete medio longitudinal
dorsal yellow stripe. Abdominal ventral terga narrow
(ratio length/width of V at basal 25% higher than 4),
maximum width of V-VI at distal 66%, basal 30% of
inner and outer lateral carinae of IV concave (figs.
173a-b). Ventral tubercle of abdominal segment I
bearing few denticles (10 or less) restricted to its apex
(fig. 251); dorsal margin of anterior lamina spine concave (fig. 333); tip of hamular anterior process pointed in posterior and ventral view (fig. 293b); auricles
with two teeth (fig. 211). Male cercus lacking pale
basal spot in outer surface (fig. 374a); dorso-distal
crest rising suddenly, as high as base of cercus in lateral view (fig. 374b). Tip of female cercus rounded (fig.
374c). Dimensions: head width: 8-8.3; HW length:
39.5-40.5; HW width: 12-13.2; HW pterostigma
length: 3.2-3.4; cerci length: ? 5-5.3, / 4.5; female
Biology. – Needham (1904) provided a brief description based on half grown larvae from Chatham
Island. He mentioned the absence of an apical tooth
in labial palps and small lateral spines on abdominal
segments VI-VII as diagnostic, but since these characters can change through different instars it is impossible to know if they are characteristic of the last instar.
Rare in collections.
93
T  E,  146, 2003
Distribution [0°S, 89-91°W, 143-600 m] (fig.
448). – Ecuador: Colón (USNM*; Currie 1901; Belle
1991; Calvert 1956).
5. R. variegata group
Two rows of cells between RP1 and RP2 in HW beginning at distal end of pterostigma or distal to it (fig.
400); male cerci with high dorso-distal crest (as high
or higher than width of cercus at base), and lateral carina concave (figs. 375-381). Seven species: R. brasiliensis, R. californica, R. confusa, R. marchali, R. peralta, R. tinti, R. variegata.
Rhionaeschna brasiliensis (von Ellenrieder &
Martins Costa) comb. n.
334-as, 375-ce, 450-Mp)
Aeshna brasiliensis von Ellenrieder & Martins Costa, 2002: 18 (description ? / larva: holotype ? Brazil: Rio Tainha900 m, 29º26’60S 54º34’60W (Rio Grande do Sul state),
20-I-1958, N.D. Santos leg. (MNRJ) [examined: holotype
?, allotype /, paratypes ? / (MNRJ, MLPA, RWGC)]).
[Aeshna peralta. – Calvert 1956: 67-72 (in part); Santos
1966a: 97; Santos 1966b: 123-124 (description of larva);
Santos 1988: 266, 276; Muzón & von Ellenrieder 1997:
143-146; Carvalho & Nessimian 1998: 3-28; Martins
Costa et al. 2000: 13. Misidentification]
Aeshna sp. – von Ellenrieder 2001a: 318; 2001b: 430.
Diagnosis. – Rhionaeschna brasiliensis shares with R.
californica and R. confusa narrow ventral terga (ratio
length/width of V at basal 25% higher than four, figs.
174-176), male cercus with inner margin diverging
gradually from outer margin and tip of cercus pointed
in both sexes (figs. 375-377a, c). It also shares with R.
californica the stem of T-spot with sides approximately
linear (figs. 18-19), vs. convex in R. confusa (fig. 20),
dark stripe on frontoclypeal groove (figs. 18-19b-c) vs.
dark stripe absent in R. confusa (figs. 20b-c), membranule dark with basal 50-60% pale vs. basal 20%
pale in R. confusa, mesanepisternal stripes incomplete
at basal 30-50% of mesanepisternum (figs. 76-80) vs.
complete in R. confusa (fig. 81), and distal portion of
anterior lamina spine linear to concave (figs. 334-335)
vs. convex in R. confusa (fig. 336). Rhionaeschna
brasiliensis and R. confusa possess PL spots on segments
VII-IX (figs. 136, 138b, d) vs. absent in R. californica
(137b, d). Rhionaeschna brasiliensis can be distinguished from R. californica and R. confusa by the angled
frontal carina in dorsal view (fig. 18a) vs. rounded (figs.
19-20a), epimeral stripes constricted or divided at midlength (figs. 76-78) vs. linear (figs. 79-80), dorso-distal
carina of cercus rising abruptly at distal 20% of cercus
(fig. 375b) vs. rising gradually at distal 30% (figs. 376377b), and presence of a sub-basal tooth in male cercus
(fig. 375b), vs. tooth vestigial (figs. 376-377). The first
94
two characters mentioned above are unique for this
species within the variegata group.
Description. – Clypeal lobes angled; clypeus and
frons yellow, grayish blue spots lateral to yellow area
surrounding T-spot stem; T-stem gradually narrowing
anteriorly, with sides approximately linear; frontal carina angled in dorsal view; confluence of eyes long
(frons + vertex/eyes + occipital triangle less than 1;
eyes /occipital triangle equal to 2); vertex yellow with
latero-posterior margins black (fig. 18a); black stripe
on frontoclypeal groove; narrow black stripe on fronto-ocular groove (figs. 18b-c). Pale mesanepisternal
stripes at basal 30-50% of mesanepisternum;
mesepimeral and metepimeral stripes yellow, constricted or divided at mid-length (figs. 76-78). Membranule dark except basal 50-60% pale. Abdomen pale
reddish brown with light blue and yellow spots; medio
longitudinal dorsal yellow stripe of female segment II
incomplete; segments V-IX with PL spots; ML and MD
spots separated from each other (figs. 136a-d). Abdominal ventral terga narrow (ratio length/ width of V
at basal 25% higher than 4) (figs. 174a-b). Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface and higher than
genital lobe (fig. 252); distal portion of ventral margin
of anterior lamina spine concave to linear (fig. 334);
tip of hamular anterior process projected in ventral
view, projection lower than its width at base (fig.
294b); auricles with two teeth (fig. 212). Male cercus
inner margin diverging gradually from outer margin
to mid-length of cercus, distal portion of inner margin
linear; tip pointed (fig. 375a); no pale basal spot in
outer surface; dorso-distal crest rising suddenly at distal 20% of cercus, as high as base of cercus in lateral
view; sub-basal tooth pointed (fig. 375b). Tip of female cercus pointed (fig. 375c). Dimensions: head
width: 8.3-9; HW length: 35.7-41; HW width: 11.5-14;
HW pterostigma length: 2.8-4; cerci length: ? 4.585.1, / 4.6-5.6; female cerci maximum width: 1.151.3; total length: 59.9-65.5.
Biology. – Larva described by Santos (1966b) and
von Ellenrieder & Costa (2002) based on reared larvae collected at small pools with abundant aquatic
vegetation formed by cascading waters characteristic
of the highlands within the region of Itatiaia. Rare in
collections.
Distribution [22-29ºS, 44-51ºW, 752-2200 m]
(fig. 450). – Brazil: Rio Grande do Sul (MNRJ*;
RWGC), Paraná (MNRJ), São Paulo (MNRJ*), Rio de
Janeiro (MLPA; MNRJ*; NMW; RWGC).
Rhionaeschna californica (Calvert) comb. n.
335-as, 377-ce, 449-Mp)
Aeshna californica Hagen, 1875: 33 (nomen nudum); Calvert,
1895: 504-507 (description ? /: holotype ? U.S.A.: Mt.
Tamalpais (California state), 25-V, C. Jack (ANSP; 9251));
Needham & Hart 1901: 45 (description larva); Martin
1908: 47, 51, 84; Walker 1912: 61, 66, 69, 71, 184-190
(redescription adults and larva); Calvert 1956:10, 22-25;
Walker 1958: 51, 54, 113-117; Musser 1962: 25-26, fig.
16; Dunkle 2000: 52, Pl. 5; Needham et al. 2000: 122,
125, 133, 138-139.
Diagnosis. – Discrimination of R. californica from
other species is given under R. brasiliensis.
frons yellow to greenish yellow or light blue, grayish
blue spots lateral to yellow area surrounding T-spot
stem; T-stem gradually narrowing anteriorly, with
sides approximately linear; frontal carina rounded in
dorsal view; confluence of eyes long (frons + vertex/
eyes + occipital triangle less than 1; eyes/occipital triangle equal to 2); vertex yellow with latero-posterior
margins black (fig. 19a); black stripe on frontoclypeal
groove; wide black stripe on fronto-ocular groove
(figs. 19b-c). Pale mesanepisternal stripes at basal 3050% of mesanepisternum; mesepimeral and
metepimeral stripes yellow to light blue and linear
mesepimeral complete or incomplete, metepimeral
complete (figs. 79-80). Membranule dark except basal
50-60% pale. Abdomen pale reddish brown with light
blue to greenish yellow spots; medio longitudinal dorsal yellow stripe of female segment II incomplete (fig.
137c); PL spots absent in segments V-IX; ML and MD
spots separated from each other (figs. 137b-d). Abdominal ventral terga narrow (ratio length/width of V
at basal 25% higher than 4) (figs. 175a-b). Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface and higher than
genital lobe (fig. 253); distal portion of ventral margin
of anterior lamina spine concave to linear (fig. 335);
tip of hamular anterior process projected in ventral
view, projection lower than its width at base (fig.
295b); auricles with two teeth (fig. 213). Male cercus
inner margin diverging gradually from outer margin
to mid-length of cercus, distal portion of inner margin
linear; tip pointed (fig. 377a); no pale basal spot in
outer surface; dorso-distal crest rising gradually at distal 0.30 of cercus, as high as base of cercus in lateral
view; sub-basal tooth vestigial (fig. 377b). Tip of female cercus pointed (fig. 377c).
Dimensions: head width: 7.3-8.5; HW length:
length: 2.8-4.6; cerci length: ? 3.8-4.5, / 3-4.8; female cerci maximum width: 1; total length: 48.2-64.
Biology. – Larvae breed in small lakes, ponds,
stream pools, and marshes. Last larval instar was described by Needham & Hart (1901) based on one
reared larva from Tombstone, Arizona, USA, and by
Musser (1962), based on larvae from Utah. Adults forage over roads and clearings. Males patrol close to water without a definite beat along shore and among
emergent plants (Dunkle 2000). Collection records
show this to be an early spring species in the western
United States. A common species.
(fig. 449). – Mexico: Baja California Norte (Dunkle
2000; Needham et al. 2000) – U.S.A.: Arizona (Needham & Hart 1901); California (DRPC; LACM; RWGC*;
Calvert 1895; Kennedy 1917), Oregon (DRPC;
RWGC*; Walker 1912), Nevada (RWGC), Idaho (DRPC;
RWGC), Utah (RWGC; Calvert 1895), Washington
(DRPC; RWGC; Calvert 1956; Needham et al. 2000),
Colorado (Needham et al. 2000), Montana (Needham et al. 2000), South Dakota (Needham et al.
2000), Wyoming (Needham et al. 2000) – Canada:
British Columbia (DRPC; LACM; Walker 1912).
Rhionaeschna confusa (Rambur) comb. n.
376-ce, 450-Mp)
Aeshna confusa Rambur, 1842: 205 (description ? /: holotype / ARGENTINA: Buenos Aires (ISNB) [examined]); Hagen 1861: 314; Hagen 1875: 39; Ris 1904: 24-25; Martin
1908: 52-53; Ris 1908: 523, 525; Martin 1911: 12; Ris
1913: 85; Navás 1916: 17; Navás 1917a: 187; Martin
1921: 22; Martin 1923: 109; Navás 1927: 23; Gazulla &
Ruiz 1928: 290; Navás 1929c: 220; Pirión 1933: 81;
Needham & Bullock 1943: 358-359; Fraser 1947: 433,
448; Mielewczyk 1978: 29; Calvert 1956: 11, 30-34, 230;
Fraser 1957: 159; Paulson 1977: 175; Abenante & Philippi 1982: 151; Davies & Tobin 1985: 2; Jurzitza 1989: 78; Mola 1991: 10; Rodrigues Capítulo et al. 1991: 62, 66;
Watson 1992: 454, 657-462; Rodrigues Capítulo 1992:
39, 56; Mola & Papeschi 1994: 185-188; Mola 1995: 4748, 54; Muzón 1997: 127-128, 132; Muzón & von Ellenrieder 1998: 23; von Ellenrieder 2001a: 301, 312-314; b:
424, 428-429, 431-433.
Diagnosis. – Discrimination from other species
provided under R. brasiliensis.
frons yellow to light blue, grayish blue spots lateral to
yellow area surrounding T-spot stem; T-stem approximately parallel sided, slightly narrowed anteriorly,
with convex sides; frontal carina evenly curved in dorsal view; confluence of eyes long (frons + vertex/eyes +
occipital triangle less than 1; eyes/occipital triangle
equal to 2); vertex black with central portion yellow
(fig. 20a); no dark stripe on frontoclypeal groove;
black stripe on fronto-ocular groove wide (figs. 20bc). Pale mesanepisternal stripes complete; mesepimeral and metepimeral stripes yellow to light blue, linear,
complete and narrow (fig. 81). Membranule dark except basal 20% pale. Abdomen pale reddish brown
with light blue and yellow spots; medio longitudinal
dorsal yellow stripe of female segment II complete (fig.
138c); segments V-IX with PL spots; ML and MD spots
separated from each other (figs. 138b, d). Abdominal
ventral terga narrow (ratio length/width of V at basal
95
T  E,  146, 2003
25% higher than 4) (figs. 176a-b). Ventral tubercle of
abdominal segment I bearing numerous denticles
spread over its posterior surface and higher than genital lobe (fig. 254); distal portion of ventral margin of
anterior lamina spine convex (fig. 336); tip of hamular
anterior process projected in ventral view, projection
lower than its width at base (fig. 296b); auricles with
two teeth (fig. 214). Male cercus inner margin diverging gradually from outer margin to mid-length of cercus, distal portion of inner margin with a slight concavity; tip pointed (fig. 376a); pale basal spot in outer
surface; dorso-distal crest rising gradually at distal
30% of cercus, as high as base of cercus in lateral view;
sub-basal tooth vestigial (fig. 376b). Tip of female cercus pointed (fig. 376c). Dimensions: head width: 7.38.1; HW length: 32.5-41.5; HW width: 11.2-14.4; HW
/ 3-4.8; female cerci maximum width: 1; total length:
48.2-59.8.
Biology. – Larva occurs in ponds and temporary
pools and was described by von Ellenrieder (2001b)
based on reared specimens from Buenos Aires, Argentina. Adults are commonly seen flying in company
with R. bonariensis over grasslands in central Argentina. Mielewczyk (1978) reported a mass migration of
R. confusa 60 miles from the shore of the Rio de La
Plata and I observed another in which hundreds of
teneral specimens of R. bonariensis and R. confusa arrived and perched in bushes and grasses near shore in
the Río de La Plata (February 1998).
450). – Chile: Aisén: Cohiaique (Calvert 1956), De
Los Lagos: Llanquihue (MNNS), Valdivia (MNNS; Jurzitza 1989), and Osorno (USNM), Araucanía: Cautín
(IEUM; Calvert 1956; Fraser 1957; Jurzitza 1989) and
Malleco (USNM; Calvert 1956; Jurzitza 1989), Biobío:
Biobío (UMMZ), Concepción (UMMZ; USNM; Calvert
1956; Jurzitza 1989), Arauco (USNM), and Ñuble
(IEUM; UMMZ; USNM), Maule: Cauquenes (UMMZ;
USNM), Talca (IEUM; Calvert 1956; Jurzitza 1989),
and Curicó (Hagen 1875; Ris 1904), Santiago: Santiago (IEUM), and Valparaiso: Valparaiso (MNNS; Gazulla & Ruiz 1928; Jurzitza 1989) and Quillota (MNNS;
Jurzitza 1989; Pirión 1933) – Argentina: Buenos Aires
(MACN; MLPA*; UMMZ; USNM; Calvert 1956; Hagen
1875; Martin 1908; Mola 1995; Mola & Papeshi
1994; Ris 1904; 1908; 1913; Scott 1934), Entre Ríos
(MACN; MLPA; Fraser 1947), Santa Fé (Navás 1917a),
Córdoba (Hagen 1875; Navás 1927; 1929c), Tucumán (IMLA), and Misiones (MACN) – Uruguay:
Montevideo (Hagen 1875; Mielewczyk 1978; Ris
1904; 1908), Colonia (Mola 1995; Mola & Papeshi
1994) – Brazil: Rio Grande do Sul state: Pelotas,
31º46’0S 52º19’60W (RMNH; Watson 1992). Santa
Catarina state: Nova Teutonia, 27º11’0S 52º13’0 W
400 m (Calvert 1956). Rio de Janeiro state: Rio de
96
Janeiro, 22º52’0S 43º15’0W 370 m (Navás 1916).
Rhionaeschna marchali (Rambur) comb. n.
378-ce, 451-Mp)
Aeshna marchali Rambur, 1842: 203 (description ?: holotype ? COLOMBIA (ISNB) [examined]); Hagen 1861: 314;
1875: 34; Martin 1908: 45, 84; Ris 1913: 83; Ris 1918:
159-160; Campos 1922: 6, 33; Navás 1934: 3; 1935: 33;
Schmidt 1952: 257; Rácenis 1953: 35, 37; Soukup 1954:
16; Calvert 1956: 12, 58-67, Map 3; Rácenis 1958;
Limongi 1983: 95, 101, 104, 106-107 (description larva); De Marmels 1988: 101; Daigle et al. 1997: 11.
[Aeshna variegata. – Calvert 1956: 90 (in part: teneral ?
from Chapare). Misidentification]
[Aeshna cornigera. – Martin 1908: 46 (in part). Misidentification]
Diagnosis. – Rhionaeschna marchali, R. peralta, R.
tinti, and R. variegata differ from R. brasiliensis, R. californica, and R. confusa by several characters detailed
under R. brasiliensis. The following combination of
characters allow for the separation of R. marchali from
R. peralta, R. tinti, and R. variegata (second contrasting character states for other species): frontoclypeal
groove lacking a dark stripe (fig. 21b-c) vs. stripe present (figs. 22-24b-c), projection of tip of hamular anterior process 50% as high (C) as width (D) at anterior margin (fig. 297b) vs. 30% as high as width at base
(figs. 298-300b), and male cercus with marked distal
concavity at distal portion of inner margin (fig. 378a)
vs. slight concavity (figs. 379-381a).
frons yellow to greenish yellow, greenish blue spots
lateral to yellow area surrounding T-spot stem; Tstem approximately parallel sided, slightly narrowed
anteriorly, with linear sides; frontal carina evenly
curved in dorsal view; confluence of eyes long (frons +
vertex/eyes + occipital triangle less than 1; eyes/occipital triangle equal to 2); vertex yellow with latero-posterior margins black (fig. 21a); no dark stripe on frontoclypeal groove; black stripe on fronto-ocular groove
wide (figs. 21b-c). Pale mesanepisternal stripes incomplete at basal 30-50% of mesanepisternum;
mesepimeral and metepimeral stripes yellow,
mesepimeral complete or incomplete and sinuous,
metepimeral complete and with posterior side concave
(fig. 82). Membranule dark except basal 30% pale.
Abdomen pale reddish brown with light blue and yellow spots; medio longitudinal dorsal yellow stripe of
female segment II complete (fig. 139c); segments V-VI,
VIII-IX with PL spots; ML and MD spots separated from
each other (figs. 139b, d). Abdominal ventral terga
wide (ratio length/width of V at basal 25% less than
3.8) (figs. 177a-b). Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface and higher than genital lobe (fig. 255);
distal portion of ventral margin of anterior lamina
spine convex (fig. 337); tip of hamular anterior
process projected in ventral view, projection as high as
its width at base (fig. 297b); auricles with two teeth
(fig. 215). Male cercus inner margin diverging suddenly from outer margin to mid-length of cercus,
where a ‘heel’ is formed, distal portion of inner margin
with a marked concavity; tip angled (fig. 378a); no
pale basal spot in outer surface; dorso-distal crest rising gradually at distal 30% of cercus, as high as base of
cercus in lateral view; sub-basal tooth pointed (fig.
378b). Tip of female cercus rounded (fig. 378c). Dimensions: head width: 8-8.4; HW length: 38-40.5; HW
width: 12-14.2; HW pterostigma length: 2.8-3.12; cerci length: ? 4.4-4.8, / 4.88-5.12; female cerci maximum width: 1-1.1; total length: 56-63.5.
Biology. – Campos (1922) recorded it as the most
abundant aeshnid from the interandean plateaus of
Ecuador where he recorded it flying over wet soils,
crop pastures, and coursing along creeks and streams.
The following notes were recorded by the collector Felix Woytkowski who collected it in Peru: ‘This species
is not faithful to any stable habitat, it prefers running
brooks with water-plants and sheltered by bush, but it
is as frequently meet over stagnant ponds and stagnant
ditches, it frequents humid paths bordered with
shrub, and at last it reaches highest hills and there it
lives on the outskirts of bush-clumps and in ravines’.
Limongi (1983) described the larva by supposition
based on exuviae collected at temporary muddy bottom pools with abundant emergent vegetation in forest areas from Distrito Federal (Venezuela) where it
was sympatric with R. cornigera.
Remarks. – The type material of this species deposited at the ISNB consists of a single male. Since
Rambur (1842) apparently described this species from
only this specimen, it is considered the holotype.
(fig. 451). – Bolivia: La Paz: Larecaja (MLPA) and
Cochabamba: Chapare (IMLA; UMMZ) – Peru: Arequipa (RWGC; Calvert 1956), Cuzco (USNM; Schmidt
1952; Calvert 1956), Lima (UMMZ; Calvert 1956),
Junín (Calvert 1956), Pasco (Calvert 1956), Amazonas (Calvert 1956), and Cajamarca (UMMZ, Calvert
1956) – Ecuador: Loja (UMMZ; MLPA; Calvert 1956;
Ris 1918), Zamora (Ris 1918), El Oro (QCAZ), Azuay
(MLPA*; KJTC; USNM), Chimborazo (Ris 1918; Campos 1922; Calvert 1956), Bolivar (Ris 1918), Tungurahua (QCAZ; UMMZ; USNM; Calvert 1956; Campos
1922), Napo (DRPC; KJTC*; QCAZ; USNM; Navás
1933), Cotopaxi (Campos 1922; Calvert 1956), Manabi (UMMZ; MLPA), Pichincha (DRPC; KJTC; MLPA;
QCAZ; Calvert 1956; Campos 1922), Imbabura (KJTC;
Calvert 1956), and Carchi (Calvert 1956) – Colombia: Cauca (USNM; Ris 1918), Valle (DRPC), Meta (Ris
1918), Tolima (DRPC), Cundinamarca (USNM; Calvert
1956; Ris 1918); Caldas (Calvert 1956; Navás 1935),
Boyaca (Ris 1918), Santander (Calvert 1956; Ris
1918), Norte de Santander (Ris 1918) – Venezuela:
Táchira (Rácenis 1958; De Marmels 1988), Mérida
(DRPC; Rácenis 1953); Distrito Federal (Limongi
1983; Rácenis 1953).
Rhionaeschna peralta (Ris) comb. n.
338-as, 379-ce, 400-wi, 452-Mp)
Aeshna peralta Ris, 1918: 159-162 (description ? /: lectotype ? (here designated) BOLIVIA: La Paz, 3600 m, 1912,
A.S. Fassl. leg. (SMFD; 16049) [examined: lectotype ?,
paralectotypes ? /]); Hincks 1934: 80; Schmidt 1952:
258; Calvert 1956: 67-72 (in part: records from Peru).
[Aeshna variegata. – Jurzitza 1990: 385-392 (synonymy with
variegata; in part, records from Peru); von Ellenrieder
2001: 317-318 (in part, records from Peru). Misidentification]
Examined material (17 ? 16 /). – Types (SMFD): 1 ?
Lectotype (/16049) Bolivia, La Paz, 3600 m, 1912, A.S.
Fassl.; 1 ? Paralectotype (/16050), 2 / Paralectotypes
(/16051-52): Peru, Apurimac, 3500 m, 4-VI-1910, Garlepp. PERU: 1 ? 2 / Cuzco, stream at Sacsayhuaman, above
Cuzco, 13-VI-1977, D.R. Paulson leg. (DRPC); 2 ? Cuzco,
10-VII-1911, Yale Peruvian exp. leg. (USNM), same except
3510 m, 7-VII-1911 (DRPC); 1 ? 1 / Puno, Lago Titicaca at
Puno, 3840 m, grassy shore, 23-V-1972, D.L. Pearson leg.
(DRPC); 1 / Puno, 3900m, 26-X-1952, F. Blancas leg.
(USNM); 2 ? Matucana, 26/27-V-1920, J.H. Williamson
(UMMZ); 1 ? 2 / Camacani, 3700 m 19/21-XI-1955, L.E.
Peña leg. (MLPA; UMMZ); 1 ? Chimo, 3700 m, 18-XI-1955,
L.E. Peña leg. (UMMZ); 5 ? 2 / La Huerta, 3800 m, 24/28XI-1955, L.E. Peña leg. (UMMZ); 1 ? 2 / Dept. Junin, Prov.
Tarma, Vitor, 5/17-IV-1940, F. Woytkowski leg. (UMMZ).
BOLIVIA: 1 / Belen Station, Altiplano, 8/9-V-1952, J.A.
Munro leg. (USNM); 2 / La Paz dept., Manco Capac prov.,
Copacabana, pequeño arroyo desembocadura en Lago Titicaca, 7-I-1972, Bulla leg. (MLPA); 1 ? 1 / Lago Titicaca,
16°09’32S 69°05’05W 3800 m, von Ellenrieder leg. (MLPA).
Diagnosis. – Distinction of R. peralta from R.
brasiliensis, R. californica, R. confusa, and R. marchali is
given under R. brasiliensis and R. marchali. Rhionaeschna peralta shares with R. variegata a T-spot stem
that gradually narrows anteriorly (figs. 22, 24a) vs.
abruptly so in R. tinti (fig. 23a), thorax with stripes
(figs. 83-85, 91) vs. basal and/or distal spots or no pale
markings in R. tinti (figs. 86-90), and abdominal ML
and MD spots separated (figs. 140, 142b, d) vs. confluent in R. tinti (fig. 141b, d), and with a linear to concave ventral margin of the anterior lamina spine (fig.
338, 340) vs. convex in R. variegata (fig. 339). It can be
distinguished from both R. tinti and R. variegata by
greater length of frons relative to eyes (ratio frons + vertex/eyes + occipital triangle greater than 1 in R. peralta,
less than 1 in R. tinti and in R. variegata; eyes/occipital
triangle less than 2 in R. peralta, fig. 22a, equal to 2 or
larger in R. tinti and R. variegata, figs. 23-24a).
Redescription. – Head. Labium yellow to pale
97
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brown; teeth brown. Labrum yellow; basal margin
with a transverse black stripe; distal margin bordered
with brown. Clypeus yellow; clypeal lobes angled.
Black stripe on frontoclypeal groove; wide black stripe
on fronto-ocular groove (figs. 22b-c). Frons yellow,
grayish blue spots lateral to yellow area surrounding
T-spot stem; T-stem gradually narrowing anteriorly,
with concave sides. Black of spot extended ventrally
on antefrons in some specimens. Frontal carina between ante- and posfrons straight in anterior view and
evenly curved in dorsal view. Vertex yellow with latero-posterior margins black. Occipital triangle yellow.
Rear of the head black. Confluence of eyes short (frons
+ vertex/eyes + occipital triangle larger than 1; eyes/
occipital triangle less than 2) (fig. 22a).
Pterothorax. Reddish brown with yellow to light
blue pale stripes, with dark brown areas over the
grooves and surrounding the base of the pale stripes.
Pale mesanepisternal stripes incomplete, occupying
between 30-50% of the sclerite, slightly diverging
basally. Yellow mesepimeral stripes incomplete, occupying the basal 75% of the sclerite, almost linear to
sinuous. Yellow or light blue metepimeral stripes complete with posterior side slightly concave (figs. 83-85).
Small yellow spot ventral to the mesostigma. Legs
black except coxae pale brown, and extensor surfaces
of femora yellow. Wings (fig. 400). Hyaline. C, ScP,
antenodals and nodal veins light yellow, rest black or
dark reddish brown. Pterostigma orange-yellow, with
black margins except costal margin pale yellow. Membranule basal 50% pale, distal 50% dark.
Abdomen. Pale reddish brown with light blue and
yellow spots; pattern as shown in figs. 140a-d. Abdomen segments V-IX with PL spots; ML and MD spots
separated from each other (figs. 140b, d). Medio longitudinal dorsal yellow stripe of female segment II incomplete (fig. 140c). Males: Pale lateral spots of segments I and II and AL of III white-yellowish; pale dorsal
spots of II-III yellow; lateral pale spots III-X white-light
blue; MD III-X yellow; PD III-X white light blue. Females: Pale lateral spots of I-II yellow ventrally, greenyellow dorsally; lateral spots III-X yellow; anterior portion of pale dorsal spots of II yellow, posterior portion
light blue; remaining dorsal pale spots yellow. Ventral
terga wide (ratio length/width of V at basal 20% less
than 3.8 mm) (figs. 178a-b), reddish brown with yellowish pale spots. Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface and higher than genital lobe; genital lobe
with 13-15 small denticles restricted to the distal 0.15
of segment II. (fig. 256). Male genital fossa: Anterior
lamina spines laterally compressed, pointed and surpassing caudally the base of the hamuli anteriores,
with distal portion of ventral margin linear to concave
(fig. 338). Tip of hamular anterior process not projected in ventral view, projection lower than its width
98
at base (fig. 298b). Auricles with two teeth (fig. 216).
Cerci: Male cercus inner margin diverging suddenly
from outer margin to mid-length of cercus, where a
‘heel’ is formed, distal portion of inner margin with a
slight concavity; tip angled (fig. 379a); no pale basal
spot in outer surface; dorso-distal crest rising gradually at distal 30% of cercus, as high as base of cercus in
lateral view; sub-basal tooth blunt (fig. 379b). Female
cercus lanceolate, with rounded tip (fig. 379c). Dimensions (those of lectotype in brackets): head width:
7.85-8.5 (8); HW length: 31-39 (36); HW width: 912.9 (12); HW pterostigma length: 2-2.8 (2); cerci
length: ? 3.75-4.5 (4), / 3-4.5; female cerci maximum width: 0.9; total length: 51.5-56.5 (53.3).
Biology. – Adults have been collected on quieter,
weedy, flat parts of streams and adjacent small marshy
ponds in dry areas of Peru and Bolivia. Rare in collections. Larva unknown.
Remarks. – Rhionaeschna peralta was described by
Ris (1918) based on a series of five specimens collected in Bolivia (La Paz) and Peru (Apurimac). In his
monograph of Neotropical Aeshnidae, Calvert (1956)
extended the distribution area of this species to
Ecuador, Brazil, Chile, and Argentina. Following
Calvert’s key and diagnosis (1956), it was later recorded from Chile by Jurzitza (1989), from Argentina by
Rodrigues Capítulo & Muzón (1989), and Rodrigues
Capítulo (1992), and from Brazil by Santos (1966b)
and Costa et al. (2000). Santos (1966b) provided a description of the last instar larva based on reared specimens from Brazil.
Rhionaeschna peralta was later synonymized with R.
variegata by Jurzitza (1990b), based on examination
of a large series of several specimens from Chile (some
of which corresponded to R. peralta according to the
key and diagnostic characters given by Calvert 1956),
eight from Argentina, and two from Peru. Calvert
(1956) considered the number of cell rows between
IRP2a-IRP2b (more than three in R. variegata, not
more than three in R. peralta) and thoracic color pattern (pale antehumeral stripe absent in R. variegata,
present in R. peralta) as diagnostic. Jurzitza (1990b)
examined venation, color pattern and shape of vesica
spermalis and ovipositor and failed to find significant
differences. However, neither Calvert nor Jurzitza
looked at the types of R. peralta. During a revision of
the Patagonian species of Rhionaeschna (von Ellenrieder 2001), I found no specimen fitting the original
description of R. peralta among Patagonian representatives and I suggested the possibility of an incorrect
synonymy.
Examination of the type series of Rhionaeschna peralta demonstrated that it represents a valid species and
that its synonymy with R. variegata was in error. However, the original description fails to provide good diagnostic characters and includes some mistakes (i.e. PL
and PD spots widely confluent in all segments, which
are in fact separated in VI-VII or VIII in some of the type
specimens, and cercus with parallel inner and outer
margins and dorsal crest arising from midlength) that
probably contributed to the failure of subsequent
workers in recognizing this species. The previous
records of this species from Chile and Argentina correspond to R. variegata, those from Brazil to R.
brasiliensis, and those from Peru to R. peralta. Rhionaeschna peralta and R. variegata seem to be allopatric,
the first restricted to Bolivia and Peru and the second
to Chile and Argentina (fig. 452).
(fig. 452). – Peru: Apurimac (SMFD*), Cuzco (DRPC;
USNM*), Junin (UMMZ), Lima (UMMZ), and Puno
(DRPC; UMMZ; USNM) – Bolivia: La Paz: Manco Capac
(SMFD*; MLPA*; USNM*).
Rhionaeschna tinti (von Ellenrieder) comb. n.
340-as, 380-ce, 451-Mp)
Aeshna tinti von Ellenrieder, 2000: 348-352, 357 (description ? /: holotype ? Chile: Tilopozo, 23°49’S 68°15’W
(El Loa prov., Antofagasta region), I-1996 (MLPA) [examined: holotype ?, allotype /, paratypes ? / (MLPA;
IEUM; MNNS; MNHN; RWGC; UMMZ)]).
Diagnosis. – Thorax color pattern with small basal
and/or distal pale spots (figs. 86-90), and abdomen
with ML and MD spots confluent (figs. 141b, d), are
unique for this species within the group. Further differences are elaborated under R. brasiliensis, R. marchali and R. peralta.
frons light yellow or light blue, gray or light blue spots
lateral to yellow area surrounding T-spot stem; Tstem suddenly narrowing anteriorly, with convex
sides; frontal carina evenly curved in dorsal view; confluence of eyes long (frons + vertex/eyes + occipital triangle less than 1; eyes /occipital triangle equal to 2);
vertex yellow with latero-posterior margins black (fig.
23a); black stripe on frontoclypeal groove present;
black stripe on fronto-ocular groove wide (figs. 23bc). Pale pterothoracic stripes absent or represented by
basal and/or distal spots (figs. 86-92). Membranule
dark except basal 20% pale. Abdomen light blue to
light yellow with dark reddish brown to black spots;
medio longitudinal dorsal yellow stripe of female segment II incomplete (fig. 141c); segments V-IX with PL
spots; ML and ML spots confluent in III-VII (figs. 141b,
d). Abdominal ventral terga wide (ratio length/width
of V at basal 25% less than 3.8 mm) (figs. 179a-b).
Ventral tubercle of abdominal segment I bearing numerous denticles spread over its posterior surface and
higher than genital lobe (fig. 257); distal portion of
ventral margin of anterior lamina spine concave to lin-
ear (fig. 340); tip of hamular anterior process projected in ventral view, projection lower than its width at
base (fig. 299b); auricles with two teeth (fig. 217).
Male cercus inner margin diverging suddenly from
outer margin to mid-length of cercus, where a ‘heel’ is
formed, distal portion of inner margin with a slight
concavity; tip angled (fig. 380a); no pale basal spot in
outer surface; dorso-distal crest rising gradually at distal 25% of cercus, as high as base of cercus in lateral
view; sub-basal tooth prominent (fig. 380b). Tip of female cercus rounded (fig. 380c).
Dimensions: head width: 7.4-8.4; HW length:
length: 2.2-3.2; cerci length: ? 3.75-4.3, / 3.8-4.35;
female cerci maximum width: 0.85; total length:
43.5-54.8.
Biology. – In Northern Chile, specimens were collected in desert farming areas as they flew over canals
used for irrigating alfalfa crops. Larva unknown. Rare
in collections, but common in the field.
(fig. 451). – Chile: Antofagasta: Tocopilla (MNNS), El
Loa (IEUM*; MNNS*; MLPA*; RWGC; UMMZ*), and
Antofagasta (IEUM; MNNS), and Tarapacá: Iquique
(IEUM; MNNS).
Rhionaeschna variegata (Fabricius) comb. n.
381-ce, 431-au, 452-Mp)
Aeshna variegata Fabricius, 1775: 425 (description ?: holotype ? Tierra del Fuego [type lost, Zimsen 1964]); Hagen
1861: 314; Hagen 1875: 38; Martin 1908: 44-45; Martin
1911: 11; Ris 1913: 81-84; Ris 1918: 159,162-163; Martin 1921: 23; Martin 1923: 109; Navás 1926: 103-104;
Calvert 1952: 258; Herrera et al. 1955-66: 81; Calvert
1956: 12, 90-98, 230; Fraser 1957: 159; Fraser 1958: 195198; Böttger & Jurzitza 1967: 37-38; Jurzitza 1975: 11;
Paulson 1977: 175; Jurzitza 1989: 10; Jurzitza 1990b: 385393; Rodrigues Capítulo et al. 1991: 62; Watson 1992:
455, 657-462; Rodrigues Capítulo 1992: 38, 57; Pérez
D’A. & Mutschke 1993-94: 63-68; Muzón 1995: 6;
Muzón 1997: 127-128, 132-133; Muzón & von Ellenrieder 1997: 143-146; Muzón & von Ellenrieder 1998: 23;
von Ellenrieder 2001a: 301, 314-320; b: 424, 429-433.
[Aeschna diffinis. – Mabille 1888: 3-6 (description); Ris
1904: 24, 26-29 (in part); Martin 1908: 43-44 (in part);
Martin 1911: 11 (in part). Misidentification]
[Aeschna diffinis var. risi Enderlein, 1912: 119. Misidentification]
[Aeshna diffinis risi. – Davies & Tobin 1985: 9-10. Misidentification]
[Aeshna peralta. – Calvert 1956: 67-72 (in part: records from
Chile and Argentina); Jurzitza 1989: 9-10; Rodrigues
Capítulo & Muzón 1989a: 76; Rodrigues Capítulo et al.
1991: 62. Misidentification]
Diagnosis. – Distinction from other species of the
group is found under R. brasiliensis, R. marchali and
R. peralta.
99
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frons light blue or yellow to greenish yellow, greenish
gray or light blue spots lateral to yellow area surrounding T-spot stem; T-stem wide (dark form) to narrow
(light form), gradually narrowing anteriorly, with linear sides; frontal carina evenly curved in dorsal view;
confluence of eyes long (frons + vertex/eyes + occipital
triangle less than 1; eyes /occipital triangle equal to
two); vertex yellow with latero-posterior margins
black (fig. 24a); black stripe on frontoclypeal groove;
wide black stripe on fronto-ocular groove (figs. 24bc). Pale mesanepisternal stripes incomplete at basal
50% of mesanepisternum to absent; mesepimeral and
metepimeral stripes yellow, mesepimeral incomplete
and sinuous, metepimeral complete and with posterior side concave (fig. 91). Membranule dark except
basal 30-50% pale. Abdomen dark redish brown to
black with light blue and yellow to greenish yellow
spots; medio longitudinal dorsal yellow stripe of female segment II incomplete (fig. 142e); segments V-IX
with (light form) or without (dark form) PL spots; ML
and MD spots separated in III-VII (figs. 142c-d, f-g).
Abdominal ventral terga wide (ratio length/ width of V
at basal 25% less than 3.8 mm) (figs. 180a-b). Ventral
tubercle of abdominal segment I bearing numerous
denticles spread over its posterior surface and higher
than genital lobe (fig. 258); distal portion of ventral
margin of anterior lamina spine convex (fig. 339); tip
of hamular anterior process projected in ventral view,
projection lower than its width at base (fig. 300b); auricles with two-three teeth (fig. 218). Male cercus inner margin diverging suddenly from outer margin to
mid-length of cercus, where a ‘heel’ is formed, distal
portion of inner margin with a slight concavity; tip angled (fig. 381a); no pale basal spot in outer surface;
dorso-distal crest rising suddenly at distal 25% of cercus, higher than base of cercus in lateral view; subbasal tooth prominent (fig. 381b). Tip of female cercus rounded (fig. 381c). Dimensions: head width:
8.4-9.5; HW length: 36.2-42.9; HW width: 11.4-14.1;
HW pterostigma length: 2.4-3.7; cerci length: ? 4.45.7, / 4.4-5.5; female cerci maximum width: 1.1; total length: 55.8-67.2.
Biology. – Rhionaeschna variegata is a highly variable species. Von Ellenrieder (2001a, b) described a
‘light’ and ‘dark’ form. The dark form is confined to
humid areas along a narrow area of the subantarctic
forest from Tierra del Fuego north to 36.5° S in Chile
and 39.6º S in Argentina. The light form appears to be
associated with xeric conditions in the Patagonian
steppe and in other dry areas of Argentina and northern Chile. Intermediate specimens occur within an
area that approximates the Central Chilean biogeographic province, a zone of intermediate aridity (3501330 mm average yearly rainfall) between those of the
humid Subantarctic forest (2000-3500 mm) and the
100
xeric region of Northern Chile and the Patagonian
steppe (0-100 mm and 100-270 mm respectively).
Larvae occur in streams, seepages and small ponds.
The larva of the dark form was described by Muzón &
von Ellenrieder (1997) and that of the light form by
von Ellenrieder (2001b) based on reared specimens
from Chile and Argentina. Rhionaeschna variegata is
the only species found in Tierra del Fuego, being the
most austral species of Odonata known.
452). – Light form: Argentina: Santa Cruz (IMLA;
MLPA; Muzón 1995), Chubut (MLPA), Río Negro
(MLPA; IMLA; Rodrigues Capítulo & Muzón 1989a;
Muzón 1995), Mendoza (MACN; UMMZ), Catamarca
(UMMZ), Salta (Jurzitza 1990b), and Jujuy (MLPA) –
Chile: Magallanes: Magallanes (IEUM; MNNS; PDA;
Pérez D’Angelo 1993-94) and Antofagasta: Antofagasta (IEUM; MLPA) and El Loa (UMMZ). – Intermediates: Chile: Libertador Gral. Bernando O’Higgins:
Colchagua (IEUM), Maule: Curicó (IEUM; UMMZ),
Linares (MNNS), and Talca (IEUM; Jurzitza 1990b),
Santiago: Chacabuco (IEUM; Jurzitza 1975), Santiago
(IEUM; MNNS; UMMZ; Calvert 1956; Jurzitza 1989;
1990b), and Cordillera (IEUM; USNM), Valparaiso:
Quillota (UMMZ; Jurzitza 1990b), San Felipe de
Aconcagua (Jurzitza 1990b), Valparaiso (IEUM; MNNS;
Calvert 1956; Jurzitza 1989; 1990b), and San Antonio (IEUM; UMMZ), and Coquimbo: Limarí (MNNS)
and Choapa (MNNS; UMMZ; Jurzitza 1990b). – Dark
form: Argentina: Tierra del Fuego (IMLA; MLPA*; Mabille 1888; Muzón 1995; Ris 1904; 1913), Santa Cruz
(IMLA; MLPA*; MNRJ; Fraser 1958; Muzón 1995),
Chubut (IMLA; Muzón 1995; Muzón & von Ellenrieder 1997), Neuquén (IMLA; MLPA*; RWGC; Calvert
1956; Muzón 1995), and Río Negro (MLPA*; Muzón
1995). – Chile: Magallanes: Última Esperanza (IEUM;
MNNS; MLPA; MNRJ; UMMZ; Ris 1913), Magallanes
(IEUM; MNNS; UMMZ; Ris 1904; 1913; Watson 1992),
and Tierra del Fuego (Navás 1916), Aisén: Aisén
(MNNS; MLPA*; RWGC; Jurzitza 1990b; Muzón & von
Ellenrieder 1997), Coihaique (MNNS; MLPA*; RWGC),
General Carrera (MLPA; Calvert 1956; Jurzitza 1989;
1990b), and Capitán Prat (Jurzitza 1990b), De Los Lagos: Valdivia (MLPA; MNNS; UMMZ; Böttger & Jurzitza
1967; Jurzitza 1975; 1989; 1990b), Osorno (MNNS;
MLPA*; UMMZ; Jurzitza 1975; 1989; 1990b), Llanquihue (IEUM; UMMZ; Jurzitza 1990b), Chiloe (IEUM;
UMMZ), and Palena (MLPA*; RWGC; UMMZ; Muzón &
von Ellenrieder 1997), Araucanía: Malleco (IEUM;
MNNS; MLPA; UMMZ; Calvert 1956; Jurzitza 1989;
1990b) and Cautín (IEUM), and Biobío: Ñuble (MNNS;
Jurzitza 1990b), Concepción (USNM; Jurzitza 1989;
1990b; Ris 1918), and Arauco (IEUM; MLPA; UMMZ).
6. R. cornigera group
Two rows of cells between RP1 and RP2 in HW beginning proximal to pterostigma or under it (figs. 401402); male sterna IX posterior to genital opercula and
X with a bright light-blue spot (figs. 143-150b); male
cercus with low dorso-distal crest, no sub-basal tooth
and extreme apex pointed and ventrally bent (figs.
382-389b); relatively narrow abdominal ventral terga
(width measured at narrowest point/length of ventral
terga IV less than 1%, except for males of vazquezae
and pauloi, width 0.13 mm and 0.18 mm, respectively; narrow ventral terga are also found in draco and
some Marmaraeschna species) (figs. 181-188). Eight
species: R. cornigera, R. haarupi, R. manni, R. nubigena, R. pauloi, R. planaltica, R. psilus, R. vazquezae.
Rhionaeschna cornigera (Brauer) comb. n.
(figs. 25,415-he, 92-94-th, 143-ab, 181-te, 219-220,418-ge, 260262-tg, 301-302-ha, 341-342-as, 382-ce, 402-wi, 434-ve, 454-Mp)
Aeshna cornigera Brauer, 1865: 906 (in part, description ?
/: lectotype ? COLOMBIA (NMW) [examined: lectotype
?, paralectotypes /]); Brauer 1866; 70-71; Calvert
1905: 179, 182-183 (in part); Martin 1908: 46, 84; Ris
1918: 163-166 (in part); Calvert 1947: 7-8 (in part); St.
Quentin 1970: 265; De Marmels 1982: 102-105 (description larva).
Aeshna cornigera cornigera. – Calvert 1947: 7-8 (in part);
Calvert 1952: 255; Calvert 1956: 11, 34-43; De Marmels
1988: 101; González Soriano 1993: 296; González Soriano & Novelo Gutiérrez 1996: 164.
Diagnosis. – Rhionaeschna cornigera is most similar
to R. nubigena and R. planaltica. It can be distinguished from R. nubigena by the approximately uniform width of the mesepimeral stripe (figs. 92-94) vs.
narrowed at midlength to 50% of its basal width in R.
nubigena (figs. 101-102), convex anterior contour of
the ventral tubercle of abdominal segment I (figs. 260262) vs. concave in R. nubigena (fig. 267), dorso-distal
crest of male cercus higher with a stronger curvature
(rising more abruptly, figs. 382b, g) vs. lower and
gradually rising in R. nubigena (figs. 385b). Separation
from R. planaltica is given under that species. The color pattern of R. cornigera is similar to that of R. psilus
from which it is easily distinguished by the well-developed spines of the anterior lamina (figs. 341-342) vs.
vestigial to absent in R. psilus (fig. 348), and much
shorter and narrower female cercus (3.45-6.5 mm by
0.6-1.1 mm, figs. 382c-e, contra 7.2-8.26 mm by
1.15-1.33 mm in R. psilus, fig. 388c).
Description. – Clypeus and frons light blue to
bluish yellow, green spots lateral to yellow area surrounding T-spot stem; T-stem approximately parallelsided; posterior 60% of vertex black (fig. 25a); frontoclypeal groove bordered by a narrow dark brown or
black stripe (figs. 25b-c). Pale mesanepisternal stripes
complete and as wide as 30-50% of mesanepisternum;
mesepimeral and metepimeral stripes bright green,
with linear or slightly undulated margins, and approximately uniform in width (fig. 92-94). Abdomen dark
reddish brown to black with bright green or yellowish
green and light blue spots; male PL spots very small to
absent (figs. 143b-c). Ventral tubercle of abdominal
segment I with anterior margin convex in lateral view,
about as high as 50% of its length (figs. 260-262);
spines of anterior lamina well developed (fig. 341342); tip of hamular anterior process rounded in posterior view (fig. 301-302a); auricles with two-three
teeth (fig. 219-220). Male cercus with a concavity at
distal 30% or parallel sided, tip directed externally or
posteriorly (figs. 382a, f); dorso-distal crest smoothly
curved, higher than base of cercus in lateral view, and
developed along distal 25-33% of cercus (figs. 382b,
g). Female cercus longer than abdominal segments IXX and shorter than VIII-X, with rounded or pointed tip,
and maximum width at medial 30% (figs. 382c-e).
Dimensions: head width: 8.4-9.55; HW length: 39.146; HW width: 11.9-14; HW pterostigma length: 2.1-3;
cerci length: ? 5-5.9, / 3.45-6.5; female cerci maximum width: 0.6-1.1; total length: 58.5-67.
Remarks. – Brauer described Aeshna cornigera
briefly in 1865, and redescribed it more extensively in
1866 based on a type series from ‘Columbien’. He did
not indicate the number of specimens of the type series but according to measurements he provided it included only one male and two or more females. St.
Quentin (1970) mentioned only two type specimens
from the original series, and designated the male as
lectotype. I found one male and two females with type
labels in the NMW collection. Examination of the type
specimens revealed that one of the paralectotype females belongs to R. planaltica. Calvert did not examine the types of R. cornigera (1947, 1952, 1956); he
based his distinction of this species from R. psilus
(1947) and R. planaltica (1952) on comments of the
types sent to him by St. Quentin. According to St.
Quentin’s letter (Calvert 1947) the thoracic stripes of
the types are not constricted so he apparently did not
see the paralectotype female belonging to R. planaltica.
The records of R. cornigera given by Calvert (1905,
1947) and Ris (1904) previous to the descriptions of
R. psilus (Calvert 1947) and R. planaltica (Calvert
1952) are not included here in the distribution list
since they encompassed all three species. The records
of Ris (1918) are included, since it is possible to discriminate between the three species through his comments on the variability of the thoracic color pattern:
his records of R. cornigera from Mexico correspond to
R. psilus, the ones from Costa Rica and Colombia to
R. cornigera, and the ones from Brazil, Bolivia and Argentina to R. planaltica.
Rhionaeschna cornigera shows the highest morpho101
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logical variability within the group: male cercus can be
parallel sided or with inner margin concave at medial
30%, and tip of cercus can be directed posteriorly or
externally (figs. 382a-b, g-f); female cercus can be
short with rounded tips or long and pointed (figs.
382d-e). Based on material I examined, northern specimens (Mexico to Colombia and Venezuela) are on
average larger, females possess a longer and more
acuminate cercus, and male cercus is sometimes parallel sided, whereas southern specimens (Venezuela and
Ecuador to Bolivia) are on average smaller, female cercus shorter and rounded, and male cercus is not parallel sided. However, I observed intergradation in these
characters and have thus been unable to separate them
into two distinct groups.
Biology. – The species has been reported as rare in
Peru (Calvert 1956), where it was seen flying over water in shrub-covered areas with abundant irrigation
canals. According to De Marmels (1982; 1988), it is
common and widely distributed in all the Andean
mountain ranges of Venezuela. It was collected at
marshes and ponds with very soft muddy bottom,
surrounded by herbs, ferns, grasses, wiht debris and
grass tufts in Costa Rica, where males were seen hovering and patrolling at about 30 cm above the water
(DRPC). Larva inhabits running waters and was described based on exuviae from Miranda, Venezuela
(De Marmels 1982).
(fig. 454). – Bolivia: La Paz: Nor Yungas (Calvert
1956) – Peru: Cuzco (UMMZ; USNM; Calvert 1956),
Junín (Calvert 1956), Pasco (Calvert 1956), Ancash
(Calvert 1956), Cajamarca (Calvert 1956) – Ecuador:
Loja (Calvert 1956), El Oro (Calvert 1956), Napo
(KJTC; MLPA*; QCAZ; Calvert 1956), Azuay (Calvert
1956), Chimborazo (Calvert 1956), Bolivar (Calvert
1956), Tungurahua (Calvert 1956), Pastaza (UMMZ;
Calvert 1956), Sucumbios (QCAZ), and Pichincha
(NMW; QCAZ; RWGC) – Colombia: without locality
(NMW*); Tolima (Ris 1918), Boyaca (UMMZ; Ris 1918;
Calvert 1956), and Magdalena (UMMZ; Calvert 1947;
1956) – Venezuela: Táchira (De Marmels 1988;
2001), Mérida (NMW; Calvert 1956), Guárico (MLPA*),
Aragua (DRPC; RWGC; USNM), Miranda (De Marmels
1982), and Distrito Federal (DRPC) – Panama: Chiriquí (Calvert 1956) – Costa Rica: Cartago (INBC;
USNM; Calvert 1956; Ris 1918), Alajuela (DRPC; INBC),
Puntarenas (DRPC), San José (DRPC; UMMZ) – Nicaragua: Matagalpa (Beckemeyer 2001) – El Salvador:
without locality (Förster 2001) – Honduras: without
locality (Förster 2001) – Guatemala: Sacatepéquez
(Calvert 1956), Quetzaltenango (Calvert 1956), Baja
Verapaz (Calvert 1956); Alta Verapaz (UMMZ, RWGC) –
Mexico: Chiapas (UNAM) and Oaxaca (UMMZ).
102
Rhionaeschna haarupi (Ris) comb. n.
343-as, 383-ce, 453-Mp)
Aeshna haarupi Ris, 1908: 523-525 (description ? /: lectotype ? ARGENTINA: Estación Pedregal, 32°58’0S
68°40’60W 695 m (Mendoza prov.), 01-XII-1906, P. Joergensen leg. (SMFD 16044)); Martin 1909: 212-213;
Martin 1911: 12; Calvert 1952: 254; Calvert 1956: 11,
25-27; Rodrigues Capítulo et al. 1991: 62; Rodrigues
Capítulo 1992: 38, 57; Muzón & von Ellenrieder 1998:
23; Muzón & von Ellenrieder 1999: 124.
Diagnosis. – This species can be distinguished from
all the other species of the group by the abruptly narrowed T-spot stem of frons (fig. 26a) and the short female cercus (as long as or shorter than segments IX-X,
fig. 383c) vs. longer than IX-X in all other species (figs.
382, 384-389c). Its thoracic and abdominal color pattern (figs. 95-96, 144) are most similar to those of R.
pauloi and R. planaltica (figs. 100, 103-105, 147-148)
but it differs from the latter two species by the longer
dorso-distal crest of male cercus (figs. 383, 386b) and
ventral terga shape of female (maximum width of ventral terga IV-V at distal portion in R. haarupi, fig. 182b)
vs. at basal portion in R. planaltica (fig. 186B). Further
differentiation from R. pauloi is given under this
species.
bluish yellow, grayish light blue spots lateral to light
yellow area surrounding T-spot stem; T-stem suddenly narrowing distally; posterior 30% of vertex black
(fig. 26a); frontoclypeal groove bordered by a narrow
dark brown or black stripe (figs. 26b-c). Pale
mesanepisternal stripes complete or interrupted into
two spots and as wide as 25-33% of mesanepisternum;
mesepimeral and metepimeral stripes bluish or yellowish-green, with deep rounded indentations on anterior
margin (metepimeral) or anterior and posterior margins (mesepimeral) (fig. 96-97). Abdomen dark reddish brown with yellowish green or yellow and light
blue spots; male PL spots large (figs. 144b). Ventral tubercle of abdominal segment I with linear or slightly
convex anterior margin in lateral view, about as high
as 50% of its length (fig. 259); spines of anterior lamina well developed (fig. 343); tip of hamular anterior
process rounded in posterior view (fig. 303a); auricles
with two teeth (fig. 221). Male cercus maximum
width at medial 30%, tip directed externally (fig.
383a); dorso-distal crest smoothly curved, higher than
base of cercus in lateral view, and developed along distal 30% of cercus (fig. 383b). Female cercus as long as
or slightly shorter than abdominal segments IX-X, with
pointed tip, and maximum width at medial 30% (fig.
383c). Dimensions: head width: 8-8.4; HW length:
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
Biology. – This species occurs in arid areas of the
Preandean mountain range. I found it flying over lateral temporary pools and swampy areas with emergent vegetation on the Calchaquí and Santa María
rivers in Salta and Tucumán provinces (Argentina).
Rare in collections and in the field. Larva unknown.
(fig. 453). – Argentina: Mendoza (NMW; Calvert
1956; Ris 1908), Catamarca (IMLA), Tucumán (MLPA;
Calvert 1956); Salta (MLPA; RWGC*).
Rhionaeschna manni (Williamson & Williamson)
comb. n.
347-as, 384-ce, 456-Mp)
Aeshna manni Williamson & Williamson, 1930: 26-34 (description ? /: holotype ? MEXICO: Los Parres [sic: Las
Parras], 25°56’60N 111°27’0W 244 m (Baja California
state), 06-X-1923, J. H. Williamson leg. (UMMZ) [examined: holotype ? allotype /]); Calvert 1947: 8-9; Calvert
1956: 12, 56-58; González Soriano 1993: 296; González
Soriano & Novelo Gutiérrez 1996: 164.
[Aeshna cornigera. – Calvert 1895: 507-508 (at least in part);
Calvert 1905: 182-183 (in part). Misidentification]
Diagnosis. – Rhionaeschna manni and R. psilus both
have reduced spines of the anterior lamina in the male
(figs. 347-348), and a long, wide, cercus in the female
(longer than segments VIII-X and wider than 1 mm,
figs. 384, 388c). They are differentiated from each
other by the mesanepisternal and mesepimeral stripes
which are light blue or greenish yellow and incomplete
in R. manni (fig. 97-99) vs. bright green and complete
in R. psilus (fig. 106). Males also differ by the shape of
the cercus: inner and outer margins diverge gradually
toward distal 30% in R. manni (fig. 384a) but are approximately parallel to each other along distal 0.60 in
R. psilus (fig. 388a). Female cercus is wider in R. manni (1.6 mm, fig. 384c) vs. 1.15-1.3 mm in R. psilus
(fig. 388a) and probably longer (broken in the allotype, only known female).
Description. – Clypeus and frons light blue to pale
brown, light blue spots lateral to yellow area surrounding T-spot stem; T-stem approximately parallelsided; posterior 50% of vertex black (fig. 27a); no dark
stripe on frontoclypeal groove (figs. 27b-c). Pale
mesanepisternal stripes incomplete or interrupted into
two spots and as wide as 25-33% of mesanepisternum;
mesepimeral and metepimeral stripes bluish or yellowish-green, with linear to undulated margins,
mesepimeral stripe incomplete, metepimeral complete
(figs. 97-99). Abdomen dark reddish brown to black
with yellowish green and light blue spots; male PL
spots large (figs. 145b). Ventral tubercle of abdominal
segment I with linear or slightly convex anterior margin in lateral view, about as high as 50% of its length
(fig. 263); spines of anterior lamina vestigial (fig. 343);
tip of hamular anterior process pointed in posterior
view (fig. 304a); auricles with three teeth (fig. 222).
Male cercus maximum width at distal 30%, with inner and outer margins diverging along medial 30%,
tip directed posteriorly (fig. 384a); dorso-distal crest
angled, lower than base of cercus in lateral view, and
developed along distal 0.30 of cercus (fig. 384b). Female cercus longer than abdominal segments VIII-X,
with maximum width at medial 30% (fig. 384c). Dimensions: head width: 8.75-9.5; HW length: 43-45;
HW width: 12.8-13.6; HW pterostigma length: 2.753.3;cerci length: ? 5.4-6, / more than 6.25; female
cerci width: 1.6; total length: 62-66.7.
Remarks. – The distinction between R. psilus and
R. manni based on the different degree of reduction
of anterior lamina spine (vestigial in R. psilus and absent in R. manni) by Calvert (1947; 1956) is incorrect
since there is a short spine in both species.
Biology. – The type series was collected at irrigation dams, pools and swampy areas associated with
mountain creeks (Williamson & Williamson 1930),
and later found patrolling small stream pools in creek
canyon at bottom of oak zone (DRPC). Rare in collections. Larva unknown.
(fig. 456). – Mexico: Baja California Sur (DRPC*;
UMMZ*; Williamson & Williamson 1930; Calvert
1895; 1947; 1956).
Rhionaeschna nubigena (De Marmels) comb. n.
344-as, 385-ce, 454-Mp)
Aeshna nubigena De Marmels, 1989b: 85-86 (description ?
/: holotype ? VENEZUELA: Neblina base camp.,
00º49’50N 66º09’40W 140 m (Río Negro dept., Amazonas state), 15-II-1985, P.J. & P.M. Spangler, R.A. Faitoure, W.E. Steiner leg. (MIZA)); De Marmels 1993: 156;
De Marmels 1997: 151.
[Aeshna planaltica. – Calvert 1956: 43-50 (in part, / from
Mt. Roraima). Misidentification]
Diagnosis. – Rhionaeschna nubigena is most similar
to R. cornigera and R. planaltica; distinction from R.
cornigera is provided under that species. It can be distinguished from R. planaltica by the mesepimeral
stripe narrowed at midlength to half its basal width
(figs. 101-102), with deep rounded indentations in R.
planaltica (figs. 103-105), and dorso-distal crest of
male cercus extended for a longer distance (distal 30%
of cercus vs. about distal 25% in R. planaltica) and
gradual prominence (fig. 385b) vs. abruptly rising
crest in R. planaltica (fig. 386b).
Description. – Clypeus and frons greenish yellow,
grayish light blue spots lateral to yellow area surrounding T-spot stem; T-stem approximately parallelsided; posterior 50% of vertex black (fig. 28a); frontoclypeal groove bordered by a narrow dark brown or
103
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black stripe (figs. 28b-c). Pale stripes complete and
bright green; mesanepisternal as wide as 25-50% of
mesanepisternum; mesepimeral narrowed at midlength to half its basal width, metepimeral approximately uniform in width (figs. 101-102). Abdomen
dark reddish brown to black with bright green or yellowish green and light blue spots; male PL spots small
to absent (figs. 146b). Ventral tubercle of abdominal
segment I with concave anterior margin in lateral view,
about as high as 50% of its length (fig. 267); spines of
anterior lamina well developed (fig. 344); tip of hamular anterior process rounded in posterior view (fig.
305a); auricles with two-three teeth (fig. 223). Male
cercus maximum width along medial 30%, tip directed externally (fig. 385a); dorso-distal crest smoothly
curved, higher than base of cercus in lateral view, and
developed along distal 30% of cercus (fig. 385b). Female cercus longer than abdominal segments IX-X and
shorter than VIII-X, with angled tip bearing mucron on
external margin, and maximum width at medial 30%
(fig. 385c-d). Dimensions: head width: 8.85-9; HW
length: 42-45; HW width: 12.8-14.6; HW pterostigma
length: 2.5-2.7; cerci length: ? 5-5.5, / 3-3.73; female cerci maximum width: 0.78; total length: 63-72.
Remarks. – Calvert (1956) included a female from
Mt. Roraima as a doubtful record of R. planaltica for
Venezuela. The specimen is preserved in alcohol, and
its coloration has faded. However, shape and size of
cerci (fig. 385c) agree well with those of R. nubigena,
and therefore is included here as this species.
Biology. – Apparently restricted to the forests of
Venezuelan tepuis. Larva unknown.
(fig. 454). – Venezuela: Río Negro (USNM; De
Marmels 1989b), Amazonas (RWGC*; De Marmels
1993; 1997), and Bolivar (ANSP*; Calvert 1956).
Rhionaeschna pauloi (Machado) comb. n.
(figs. 29-he, 100-th, 147-ab, 185-te, 224-ge, 266-tg, 306-ha, 346as, 387-ce, 453-Mp)
Aeshna pauloi Machado, 1994: 159-168 (description ? /:
holotype ? BRAZIL: Serra do Cipó, 19°0’0S 43°38’60W
1300-1400 m, Santana do Riacho (Minas Gerais state),
IV-1983, A. & P.A. & E. Machado leg. (ABBM) [examined: ? / paratypes (RWGC)])
Diagnosis. – Rhionaeschna pauloi can be distinguished from all the other species of the group by the
low ventral tubercle of segment I; lower than 30% of
its length (fig. 266) vs. as high as 50% of its length in
remaining species (figs. 259-265, 267-269). Thoracic
color pattern is similar to that of R. haarupi and R.
planaltica (figs. 95-96, 100, 103-105); it differs from
these species by the rounded cercus tip of females (fig.
387c), which is pointed in R. haarupi and R. planaltica (figs. 383c, 386c-e), tip of male cercus directed pos104
teriorly (fig. 387a) vs. directed externally in R. haarupi
and R. planaltica (figs. 383, 386a), male cercus with
inner and outer margins parallel along distal 60% in
dorsal view (fig. 387a) vs. converging in the other two
species (figs. 383, 386a), and overall larger size (length
of cerci, HW, and body).
Description. – Clypeus and frons light blue to grayish yellow, light blue to bluish gray spots lateral to yellow area surrounding T-spot stem; T-stem gradually
narrowing anteriorly; vertex black except medial 30%
yellow (fig. 29a); frontoclypeal groove bordered by a
narrow dark brown or black stripe (figs. 29b-c). Pale
mesanepisternal stripes absent or incomplete and interrupted into two spots and as wide as 25-33% of
mesanepisternum; mesepimeral and metepimeral
stripes yellowish-green, with deep rounded indentations on anterior margin (mesepimeral) or anterior
and posterior margins (metepimeral) (fig. 100). Abdomen dark reddish brown with yellowish green and
light blue spots; male PL spots large (fig. 147b). Ventral tubercle of abdominal segment I with anterior
margin convex in lateral view, lower than 30% of its
length (fig. 266); tip of hamular anterior process
pointed in posterior view (fig. 306); spines of anterior
lamina well developed (fig. 346). Auricles with twothree teeth (fig. 224). Male cercus distal 60% parallel
sided, tip directed posteriorly (fig. 387a); dorso-distal
crest high (higher than base of cercus in lateral view)
and smoothly curved along distal 25% of cercus (fig.
387b). Female cercus longer than abdominal segments
IX-X and shorter than VIII-X, rounded tip with a
minute medial spine, and maximum width at medial
30% (fig. 387c). Dimensions: head width: 9.4-9.6;
HW length: 44-53; HW width: 13.6-14.5; pterostigma
length: 2.8-3.75; cerci length: ? 5.7-6, / 5-6.2; maximum width female cerci: 1.1; total length: 63-77.
Biology. – This is apparently a mountain species in
Brazil restricted to rocky streams and pools at elevations above 1000 meters where the larvae occur. Rare
in collections. Some of the paratypes described by
Machado (1994) were reared from larvae but the larva has not yet been described. The species is not endemic to the iron-rich streams mentioned by Machado, since I find a specimen from Sapucay, Paraguay,
and further collecting will probably show the species
to be more widespread.
Distribution [20-25°S, 43-56°W 190-1400 m]
(fig. 453). – Paraguay: Paraguarí (UMMZ) – Brazil:
Paraná (ABBM) and Minas Gerais (ABBM).
Rhionaeschna planaltica (Calvert) comb. n.
(figs. 30-he, 103-105-th, 148-ab, 186-te, 225-ge, 264-265-tg, 307ha, 345-as, 386-ce, 401-wi, 453-Mp)
Aeshna cornigera planaltica Calvert, 1952: 255-256 (description ? /: holotype ? BRAZIL: Santa Catarina state: Nova
Teutonia, 27º11’S 52º13’W 300-500 m), 01-XII-1935,
F. Plaumann leg. (MCZ)); Calvert 1956: 11, 43-50; Rácenis 1959: 493-494; Santos 1970: 203; Jurzitza 1981: 117;
De Marmels 1989a: 54-55; 1989b: 35; 1989c: 85; 1990b:
337; Mola 1991: 10; De Marmels 1992a: 64, figs. 47-48
(description larva); Rodrigues Capítulo et al. 1991: 62,
66; Rodrigues Capítulo 1992: 39, 57; Mola & Papeschi
1994: 185-187; Mola 1995: 47, 51-54; Muzón & von Ellenrieder 1998: 23; von Ellenrieder 1999: 151-155 (description larva); von Ellenrieder 2000: 21, 24, 28; Martins
Costa et al. 2000.
[Aeshna cornigera. – Hagen 1875: 39 (at least in part: Brazil
and Uruguay); Selys 1883: 21; Ris 1904: 23; Calvert
1901-1908: 182-183 (at least in part: Uruguay, Paraguay,
Bolivia and Brazil); Ris 1913: 81; Ris 1918: 157-158;
Navás 1916: 17; Ris 1918: 157-158 (in part: Brazil, Bolivia and Argentina); Navás 1922: 358; Fraser 1947: 433;
Schmidt 1952: 238; Abenante & Philippi 1982: 151; Rodrigues Capítulo et al. 1991: 62; Carvalho & Nessimian
1998: 8; Martins Costa 1999: 4. Misidentification]
[Aeshna cornigera cornigera. – Rodrigues Capítulo 1992: 57.
Misidentification]
Diagnosis. – Rhionaeschna planaltica differs from R.
cornigera by the mes- and metepimeral green stripes
with deep rounded indentations (figs. 103-105) vs.
linear or slightly undulated in R. cornigera (figs. 9294), and the linear anterior contour of the ventral tubercle of first abdominal segment (figs. 264-265) vs.
convex in R. cornigera (figs. 260-262). Pale areas on
the first abdominal sternum in the male (figs. 219220, 225) and pale spots of abdomen (figs. 143, 148)
are wider than in R. cornigera.
bluish yellow, grayish light blue spots lateral to yellow
area surrounding T-spot stem; T-stem approximately
parallel-sided; posterior 50% of vertex black (fig. 30a);
frontoclypeal groove bordered by a narrow dark
brown or black stripe (figs. 30b-c). Pale mesanepisternal stripes complete and about as wide as 50% of
stripes bright green, with deep rounded indentations
on anterior margin (metepimeral) or anterior and posterior margins (mesepimeral) (figs. 103-105). Abdomen dark reddish brown to black with bright green
or yellowish green and light blue spots; male PL spots
large (figs. 148b). Ventral tubercle of abdominal segment I with anterior margin linear in lateral view,
about as high as 50% of its length (fig. 264-265);
spines of anterior lamina well developed (fig. 345); tip
of hamular anterior process rounded in posterior view
(fig. 307); auricles with two-three teeth (fig. 225).
Male cercus maximum width along medial 30%, tip
directed externally (fig. 386a); dorso-distal crest
smoothly curved, higher than base of cercus in lateral
view, and developed along distal 25% of cercus (fig.
386b). Female cercus longer than abdominal segments
IX-X and shorter than VIII-X, pointed tip, and maximum width at medial 30% (fig. 386c-e). Dimensions:
head width: 8.2-9; HW length: 36.5-44; HW width: 11-
14; HW pterostigma length: 2.1-2.9; cerci length: ?
4.18-5.35, / 3.95-5.1; female cerci maximum width:
0.7-0.95; total length: 57.5-65.
Remarks. – Rhionaeschna planaltica was described
by Calvert (1952) as a subspecies of R. cornigera based
on the different thoracic color pattern which he
(1905) and Ris (1918) previously mentioned. Calvert
(1952) applied the name to the most southern populations ranging from Ecuador to Argentina, whereas R.
c. cornigera corresponded to more northerly populations. However, Calvert (1952; 1956) included both
subspecies collected from the same localities and, in
some cases, the same date for several localities in Bolivia, Peru, and Ecuador (figs. 453-454). Since subspecies are geographic races that occupy different
ranges of distribution, the wide area of sympatry of R.
cornigera and R. planaltica indicates that they cannot
be considered as such. They could belong to the same
polymorphic species or to two different species. Since
no intermediates have been found between the two
morphs and the amount of differences between them
mirrors that between R. cornigera and R. nubigena, or
R. planaltica and R. haarupi, I consider them to be different species.
Biology. – The final larval instar was described by
De Marmels (1992a) based on one reared specimen
from Tamacuari, Venezuela, and by von Ellenrieder
(1999a) based on reared specimens from Salta, Argentina. Some differences between the Venezuelan
and Brazilian and Argentine larvae were observed (De
Marmels 1992a; von Ellenrieder 1999), but the
adults show no differences (De Marmels, pers. com.).
A common species. The larvae occur in mountain
creeks in Venezuela and in small ponds and temporary pools in Argentina within forest areas.
Distribution [34°S-6°N, 40°-79°W, 0-3700 m]
(fig. 453). – Argentina: Buenos Aires (MLPA*), Córdoba (MLPA; Navás 1927), Tucumán (IMLA; DRPC;
Calvert 1956; Fraser 1947; Navás 1922; Ris 1913;
1918; Schmidt 1952), Catamarca (Fraser 1947; Ris
1918; Schmidt 1952), Salta (IMLA; MLPA*; QCAZ;
Calvert 1956; von Ellenrieder 1999), Jujuy (DRPC*),
and Misiones (MACN; MLPA; USNM; Jurzitza 1981;
Mola & Papeschi 1994; Mola 1995) – Uruguay: San
José (Calvert 1952; 1956), Rocha (Calvert 1952;
1956) – Paraguay: Paraguarí (USNM; Calvert 195256), Guaira (UMMZ), and Central (DRPC) – Brazil: Rio
Grande do Sul (MNRJ; NMW; Calvert 1956; Ris 1918),
Santa Catarina (MNRJ; RWGC; Calvert 1952; 1956),
Paraná (MNRJ), São Paulo (MNRJ; RWGC; UMMZ;
USNM; Calvert 1956; Martins Costa et al. 2000; Ris
1918), Rio de Janeiro (MNRJ*; Calvert 1956), Mato
Grosso do sul (MNRJ), Minas Gerais (MNRJ), Espirito
Santo (MNRJ), and Mato Grosso (MNRJ; Calvert 1956)
– Bolivia: Cochabamba: Chapare (UMMZ; Calvert
1956) and La Paz: Larecaja (MLPA) and Nor Yungas
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(Ris 1918; Calvert 1956) – Peru: Apurimac (MLPA),
Junín (Calvert 1952; 1956), Lima (Calvert 1956),
Pasco (Calvert 1952), Huanuco (DRPC; UMMZ), Ancash (Calvert 1952; 1956), Cajamarca (Calvert 1952;
1956), Andes (Calvert 1956), and Amazonas (Calvert
1952-56) – Ecuador: Azuay (Calvert 1952), Morona
Santiago (UMMZ), Chimborazo (Calvert 1952), Bolivar (Calvert 1956), Pastaza (UMMZ; USNM; Calvert
1952-56), and Pichincha (DRPC) – Venezuela: Amazonas (USNM; De Marmels 1989b; 1992a), Atabapo
(De Marmels 1989a), and Bolivar (De Marmels
1989a) – Colombia: without locality (NMW*).
Rhionaeschna psilus (Calvert) comb. n.
Aeshna psilus Calvert, 1947: 1-7 (description ? /: holotype
? COSTA RICA: stagnant pool, banks of Río Reventazón,
Cachí, 10-III-1910, P.P. Calvert leg. (ANSP, 9273)); Needham & Westfall 1955: 291, 293-294, 296-297, 312-313;
Calvert 1956: 11, 50-55 194-196; De Marmels 1981: 11;
1988: 101; González Soriano 1993: 296; González Soriano & Novelo Gutiérrez 1996: 164; Donnelly et al. 1998:
101; Muzón & von Ellenrieder 1998: 23.
[Aeshna cornigera. – Calvert 1895: 507-508, figs. 24, 31-32
(in part); Calvert 1905: 179,182 (in part); Calvert 1907:
400 (in part); Ris 1918: 157-158 (in part); Klots 1932:
18; García-Díaz 1938: 55. Misidentification]
Diagnosis. – Rhionaeschna psilus is most closely related to R. manni, their separation is given under R.
manni.
Description. – Clypeus and frons light blue, green
T-stem approximately parallel-sided or slightly converging anteriorly; posterior 60% of vertex black (fig.
31a); no dark stripe on frontoclypeal groove (figs.
31b-c). Pale mesanepisternal stripes complete and as
wide as 25-33% of mesanepisternum; mesepimeral
and metepimeral stripes bright green, with slightly sinuous margins (fig. 106). Abdomen dark reddish
brown to black with bright green or yellowish green
and light blue spots; male PL spots large (fig. 149b).
Ventral tubercle of abdominal segment I with anterior
margin linear or slightly convex in lateral view, about
as high as 50% of its length (fig. 268); spines of anterior lamina vestigial (fig. 348); tip of hamular anterior
process rounded in posterior view (fig. 308); auricles
with three teeth (fig. 226). Male cercus with inner and
outer margins approximately parallel to each other
along medial 30%, where maximum width is observed, tip directed posteriorly (fig. 388a); dorso-distal
crest angled, lower than base of cercus in lateral view,
and developed along distal 30% of cercus (fig. 388b).
Female cercus longer than abdominal segments VIII-X,
pointed tip, and maximum width at medial 30% (fig.
388c). Dimensions: head width: 7.8-8; HW length:
106
36-44; HW width: 10.6-12.5; HW pterostigma length:
2.2-2.8; cerci length: ? 4.9-5.9-, / 7.28-8.26; female
cerci maximum width: 1.15-1.33; total length: 59-62.
Biology. – The larva was described by Needham &
Westfall (1955) based on material from the USA, and
by Calvert (1956) based on material from Costa Rica,
Guyana and Cuba. Since Calvert did not breed the
larvae, it is likely that some of his specimens belong to
species other than R. psilus (i.e. material from Mt. Roraima could be R. nubigena or R. planaltica, and material from Costa Rica could be R. cornigera). The specimens from Cuba are probably R. psilus, since this is
the only species of the group recorded from that country. The species was collected at small rain ponds with
grassy margins in open areas, ponds with abundant
aquatic vegetation, grassy marshes with Salvinia and
Lemna and large aroid and trees at edge, reservoirs, a
wooded pond in sun, drainage flowing ditches with
grassy banks and mud bottom, flying over cornfields,
small rocky and sandy streams in stream bed (DRPC).
(fig. 455). – Argentina: Salta (MLPA; Donnelly et al.
1998) – Peru: Lima (Calvert 1947; 1956), and Junín
(Calvert 1947; 1956) – Ecuador: El Oro (UMMZ),
Tungurahua (Calvert 1947; 1956), Napo (Calvert
1947; 1956), and Manabi (UMMZ) – Venezuela:
Táchira (De Marmels 1988) and Miranda (De
Marmels 1981) – Panama: Chiriquí (Calvert 1947;
1956) – Costa Rica: Puntarenas (INBC; USNM), San
José (DRPC; MN; INBC; Calvert 1947), Cartago
(USNM; Calvert 1947; 1956), and Alajuela (USNM) –
Nicaragua: Matagalpa (Beckemeyer 2001) – Honduras: without locality (Förster 2001) – Guatemala:
Guatemala (Calvert 1947; 1956) and Alta Verapaz
(USNM) – Belize: Cayo (Boomsma & Dunkle 1996) –
Puerto Rico: Río Grande (RWGC; Calvert 1956) – Dominican Republic: Barahoa (USNM) and La Vega
(RWGC; USNM) – Dominica: Dominica (Donnelly
1970) – Haiti: de l’Ouest (DRPC) – Jamaica: Surrey
(Calvert 1956) – Cuba: Pinar del Río (Peters 1988),
Oriente (Alayo 1968, Peters 1988) and Camagüey
(Calvert 1956) – Mexico: Chiapas (DRPC; RWGC;
USNM), Tabasco (UMMZ), Veracruz (RWGC; USNM; Ris
1918; Calvert 1905; 1947; 1956), Quintana Roo
(LACM; USNM; Calvert 1905; 1947; 1956), Morelos
(Calvert 1905; 1947; 1956; Ris 1918), Aguascalientes
(Calvert 1905; 1947; 1956), Nayarit (DRPC; Calvert
1905; 1956); Jalisco (Calvert 1905; 1947; 1956), Yucatán (LACM; Calvert 1947; 1956), Hidalgo (DRPC),
San Luis Potosí (DRPC), and Sinaloa (DRPC) – U.S.A.:
Texas (RWGC) and Arizona (Behrstock 1998).
Rhionaeschna vazquezae (González) comb. n.
349-as, 389-ce, 456-Mp)
Aeshna vazquezae González, 1986: 47-51, figs. 1-4 (description ?: holotype ? MEXICO: Acahuizotla, 17.5 km S
Chilpancingo (Guerrero state), 15-XI-1983, R. Mendoza
leg. (UNAM) [examined: ? paratype]); 1993: 296;
González & Novelo 1996: 164.
Diagnosis. – Rhionaeschna vazquezae is unique in
the group by the very wide mes- and metepimeral
stripes (wider than half of their sclerites, figs. 107109). It shares with R. manni and R. psilus the presence of an angulate dorso-distal carina on the male
cercus being lower than the base of cercus (figs. 384,
388-389) but it is easily distinguished from those
species by the strongly developed anterior lamina
spine (fig. 349), this structure being rudimentary in R.
manni and R. psilus (figs. 347-348).
Description. – Clypeus and frons greenish light
blue, green spots lateral to yellow area surrounding Tspot stem; T-stem gradually narrowing anteriorly; vertex black except for two yellow antero-lateral spots
(fig. 32a); frontoclypeal groove bordered by a narrow
black stripe (figs. 32b-c). Pale stripes complete and
bright green; mesanepisternal about as wide as 60% of
wider than 50% of their sclerites, with linear anterior
margins and posterior margins slightly undulated
(mesepimeral) or deeply excavated or divided (metepimeral) (figs. 107-109). Abdomen black with bright
green and blue spots; male PL spots large (figs. 150b).
Ventral tubercle of abdominal segment I with anterior
margin linear in lateral view, about as high as 50% of
its length (fig. 269); spines of anterior lamina well developed (fig. 349); tip of hamular anterior process carinated in posterior view (fig. 309a); auricles with three
teeth (fig. 227). Male cercus maximum width at distal
30%, tip between directed externally (fig. 389a); dorso-distal crest angled, lower than base of cercus in lateral view, and developed along distal 30% of cercus
(fig. 389b). Female unknown. Dimensions: head
width: 8.6-9.5; HW length: 45.5-47.7; HW width:
13.7-15; HW pterostigma length: 2.37-2.8; cerci
length: ? 5.62-6.5; total length: 68-72.
Biology. – Rhionaeschna vazquezae was recently collected for the first time after its original description by
Dr. D. R. Paulson, who kindly provided the following
notes: ‘The specimens were collected at a stream with
boulders and gravel riffles, muddy pools (one of them
long), and steep gradient, in low forest with sun penetrating. Males flew up and down along pool, sometimes
over center, mostly right at shore, at water surface and in
and out of overhanging branches and roots, with a very
slow and methodical searching, mostly after the sun had
disappeared’. Rare in collections. Larva unknown.
Distribution [17°-21°N, 99°-104°W, 950 m] (fig.
456). – Mexico: Guerrero (UNAM*; González 1986;
1993; González & Novelo 1996) and Nayarit (RWGC;
DRPC*).
7. R. punctata group
Two rows of cells between RP1 and RP2 in HW beginning under pterostigma or proximal to it (fig. 403);
male cercus widening gradually to distal end (maximum width at distal 0.12-0.16) with high dorso-distal
crest, lateral carina convex, ventrally bent tip, and subbasal tooth absent (figs. 390-395); denticles of genital
lobe very small, densely spread over the ventral and
lateral surfaces of the lobe (figs. 270-275); hamular
anterior process carinated. Seven species: R. biliosa, R.
condor, R. decessus, R. demarmelsi, R. eduardoi, R. joannisi, R. punctata.
Rhionaeschna biliosa (Kennedy) comb. n.
350-as, 392-ce, 457-Mp)
Aeshna biliosa Kennedy, 1938: 573-576 (description ?:
holotype ? ECUADOR: La Ventana, near Baños, 2000 m,
Río Pastaza (Tungurahua prov.), V-1936, W. ClarkeMcIntyre leg. (UMMZ) [examined: holotype ?, allotype
/, paratype ?]); Kennedy 1939: 344-348 (description
/); Calvert 1952: 254; Soukup 1954: 15; Calvert 1956:
12, 72-76; Rácenis 1959: 493.
Diagnosis. – Rhionaeschna biliosa is easily distinguished from its congeners by the bright yellow frons
without dark stripe on frontoclypeal groove and the
narrow transverse arms of T-spot. It shares with some
specimens of R. joannisi a similar thoracic pattern of
pale spots but these are yellow in R. biliosa and not
green as in R. joannisi. Males share similar morphologies of cercus, hamulus, and anterior lamina spine
with R. condor and R. joannisi, and females the mucronate condition of the cercus with R. eduardoi.
Description. – Clypeus orange, frons bright yellow,
no spots lateral to T-spot; T-stem parallel-sided, narrower than vertex, transverse arms represented by a fine
black line; vertex completely black or with two anterolateral yellow spots (fig. 33a); frontoclypeal groove not
bordered by dark stripe (figs. 33b-c). Pale pterothoracic stripes yellow, divided in spots: mesanepisternal
into a basal and a distal spot; mesepimeral into four
spots and metepimeral into two to four spots (figs.
110-111). Abdomen dark reddish brown to black with
yellow to pale brown spots. Outer margin of ventral
terga VII sinuous (figs. 189a-b). Ventral tubercle of abdominal segment I rounded in lateral view; genital lobe
lower than 30% of its length (fig. 270); spines of anterior lamina longer than twice its basal width and acute
pointed (fig. 350); ventral portion of hamuli higher
than hamular fold (fig. 310a), tip of hamular anterior
process rounded in posterior view and pointed in ventral view (figs. 310a-b); auricles with two teeth (fig.
228). Dorso-distal crest of male cercus pointed, tip
pointed (fig. 392b). Maximum width of female cercus
at medial 30%; tip mucronate (fig. 392c). Dimensions:
107
T  E,  146, 2003
head width: 9.9-10.4; HW length: 51.1-52; HW width:
14.7-17; HW pterostigma length: 2.8-3.3; cerci length:
? 7-7.3, / 7.4; female cerci maximum width: 1.6; total length: 73-79.
Biology. – All specimens have been collected in
Andean localities. Larva unknown. Rare in collections.
457). – Peru: Amazonas (Kennedy 1938; Calvert
1956) – Ecuador: Tungurahua (MNRJ; RWGC*; Calvert
1956; Kennedy 1938; 1939) and Manabi (MLPA;
UMMZ*).
Rhionaeschna condor (De Marmels) comb. n.
Aeshna condor De Marmels, 2001a: 129-134 (description ?
and larva: holotype ? VENEZUELA: San Vicente de La Revancha, road to Las Copas, Tamá National Park,
7°29’59N 72°20’35W 2000 m (Táchira state), 06-XI1999, J. De Marmels leg. (MIZA) [paratype ? (RWGC)])
Diagnosis. – Rhionaeschna condor is distinguished
from the other species of the group by the wide T-spot
of frons (fig. 34a), wide green thoracic stripes (fig.
112), and recumbent tip of hamular anterior process
(fig. 313a).
Description. – Clypeus and frons light blue, dark
blue-grayish spots lateral to yellow area surrounding
T-spot stem; T-stem wider than vertex, with convex
sides, transverse arms wider than a line; vertex black
(fig. 34a); narrow dark stripe on frontoclypeal groove,
not widened to fronto-ocular groove (figs. 34b-c). Pale
pterothoracic stripes yellowish green, complete and
wider than 50% of their sclerites, mesepimeral stripe
with a pointed branch extending on posterior 50% of
metepisternum (fig. 112). Abdomen black with yellowish green and blue spots. Outer margin of ventral
terga VII sinuous (figs. 190a-b). Ventral tubercle of abdominal segment I trapezoidal in lateral view, genital
lobe lower than 30% of its length (fig. 271); spine of
anterior lamina longer than twice its basal width and
acute pointed (fig. 351); ventral portion of hamuli
higher than hamular fold (fig. 313a), tip of hamular
anterior process recumbent in posterior view and
pointed in ventral view (figs. 313a-b); auricles with
three teeth (fig. 229). Dorso-distal crest of male cercus
angled, tip pointed (fig. 390b). Female unknown. Dimensions: head width: 10; HW length: 49.2-51.3; HW
width: 14.12; HW pterostigma length: 2.2-2.6; cerci
length: ? 5.3-6; total length: 69-71.3.
Biology. – Males and exuviae of this species were
collected in a small pond fed by a minute stream in the
Mount Tamá range (De Marmels 2001a). Males were
observed patrolling the edge of the pond and females
attempting to oviposit into a branch of a partly sub108
merged bush and floating grass stems (De Marmels
2001a). Rare in collections.
Distribution [7°N, 72°W, 2000 m] (fig. 457). –
Venezuela: Táchira (RWGC*).
Rhionaeschna decessus (Calvert) comb. n.
(fig. 459-Mp)
Aeshna decessus Calvert, 1953: 205-207 (description ?:
holotype ? Brazil: Südhäng Itatiaia-Gebirg., 22º25’22S
44º37’4W 700 m, X-1931, Zikan leg. (Rio de Janeiro)
[type probably lost]); Calvert 1956: 12, 88-90.
Diagnosis. – According to its original description,
this species is closely related to R. punctata and R. eduardoi, with which it shares in the male similarly
shaped cercus. The orange face, the absence of spines
in the anterior lamina, and the pale abdominal PL
spots present on segments IV-IX (PL spots absent in VIVII in R. eduardoi and R. punctata, figs. 155-156b, d)
would distinguish it from these species.
Description. – Clypeus orange, postfrons orange,
antefrons orange with purplish spots lateral to pale
area surrounding T-spot stem; T-stem parallel-sided,
narrower than vertex, transverse arms wider than a
line; dark stripe on frontoclypeal groove; vertex orange
with lateral and posterior margins black. Pale
mesanepisternal at basal 80% of mesanepisternum
stripes greenish, slightly diverging from each other.
Abdomen reddish brown with pale green and light
blue spots. Genital lobe higher than tubercle; spines of
anterior lamina absent; auricles with two teeth. Dorsodistal crest of male cercus smoothly curved, tip pointed. Female unknown. Dimensions: head width:
11.44; HW length: 51; HW width: 16; cerci length: ?
5.73; total length: 78.
Biology. – Rhionaeschna decessus is only known
from the male holotype collected at Itatiaia.
Remarks. – The diagnosis and comments on this
species are based solely on the original description.
Calvert (1953) mentioned that the holotype was first
identified as R. punctata and that the genital fossa was
coated with a hardened substance which was subsequently removed. Since the spines of the anterior lamina in R. punctata and R. eduardoi are slender, it is possible that such fragile structures, if originally present,
could have been accidentally removed during cleaning. If so, this specimen may represent a pale specimen
of R. punctata or R. eduardoi.
The type specimen is apparently lost. It was originally deposited in the collection of Dr. E. Schmidt at
the Zoologisches Forschungsinstitut und Museum
Alexander König in Bonn. The collection of Schmidt
was later sold to Dr. S. Asahina, who in turn donated it
to the National Museum of Natural History in Japan.
None of these institutions hosts it today (pers. coms. of
Dr. B. Misof, Dr. S. Asahina and Dr. M. Tomokuni).
Distribution [22°S, 44°W, 700 m] (fig. 459). –
Brazil: Rio de Janeiro (Calvert 1953).
Rhionaeschna demarmelsi von Ellenrieder sp. n.
(figs. 35-he, 113-th, 153-ab, 191-te, 230-ge, 272-tg, 311-ha, 353354-as, 393-ce, 403,411-wi, 458-Mp)
[Aeshna joannisi. – De Marmels 2001a (description ?, /,
larva). Misidentification]
Type Material: 1? holotype VENEZUELA: San Vicente de La Revancha, Las Copas (Táchira State) laguna, 2000 m, 16-v-1999, De Marmels leg. (MIZA) – 1/
allotype, 3? paratypes same except Tamá National
Park, 7°29’59N 72°20’35W 2000 m, 16-v-1999 (allotype MIZA /; paratypes MIZA /; paratype MLPA); 3?
same data except 7/11-xi-1999; 4 ? same data except
7/8-vii-2000 (MIZA/; RWGC).
Etymology: Rhionaeschna demarmelsi (noun in the
genitive case), is named in honor of Dr. Jürg De
Marmels, whose work on the systematics of the
Odonata of Venezuela serves as an inspiration to us all.
Diagnosis. – The short spine of anterior lamina
(figs. 353-354) and the combination of smoothly
curved dorso-distal crest and blunt tip of male cercus
(figs. 393b) are unique for R. demarmelsi within the R.
punctata group, and separate it from all its congeners.
The female cercus with maximum width at distal 30%
(fig. 393c) is also characteristic and allows identifying
the female from all the others known for the group (R.
biliosa, R. joannisi, R. eduardoi, and R. punctata, figs.
391-392, 394-395). The overall color pattern of
Rhionaeschna demarmelsi is very similar to that of R.
joannisi, but it differs from it in several other characters: postero-basal indentation of mesepimeral stripe
and antero-distal of metepimeral stripe not reaching
half of stripe width (fig. 113, surpassing half of stripe
width or dividing the stripes into spots in R. joannisi,
figs. 115-117); auricles bearing two large teeth (fig.
230, three in R. joannisi, fig. 231); ventral portion of
hamuli shorter than hamular fold (higher in R. joannisi); shorter female cerci (fig. 393c, 5 mm versus 6.5
mm in R. joannisi, fig. 391c) and with maximum
width at distal 30% (at medial 30% in R. joannisi);
outer margin of ventral terga VII linear (fig. 191, sinuous in R. joannisi, fig. 192); abdominal spots AL, AD,
ML, and MD present in segments VII-VIII of male (figs.
153a-b, absent in R. joannisi, figs. 154a-b) and AL, MD,
and PD present in segment VIII of female (absent in R.
joannisi, fig. 154c-e).
Description. – Holotype (variation of paratypes in
brackets): Head. Labium brown. Labrum yellow with
ventral margin dark brown and dorsal margin black.
Clypeus light blue, with lateral margins yellow. Frons
yellow [to light blue], dark grayish-blue spots lateral to
yellow area surrounding T-spot stem; T-stem narrow-
er than vertex, approximately parallel sided, transverse
arms wider than a line; vertex black with a green spot
at anterior margin on each side (as in fig. 35a); brown
stripe on frontoclypeal groove, not widened to frontoocular groove (as in figs. 35b-c). Thorax. Pale
pterothoracic stripes yellowish green, divided or with
indentations; mesanepisternal divided into a basal
short stripe (as long as 25% of mesanepisternum) and
a distal spot, mesepimeral and metepimeral each with
a postero-basal and an antero-distal indentation; postero-basal indentation of mesepimeral stripe and antero-distal of metepimeral not reaching half of stripe
width (as in fig. 113), yellowish green spot at each side
of antealar crest in antealar sinus, and on each subalar,
costal, and radioanal plate, meso-and metascutum;
light blue on metascutum. Legs black except for
trochanters, coxae and basal 75% of femora reddishbrown, and basal 50% of inner surface of femur I yellowish-green. Wings hyaline, tinged with yellow at
base (basally to first antenodal) and at nodus [or whole
wings]; membranule dark except basal 30% pale (fig.
403). Wing venation: Antenodals FW: 18; HW: 13
right, 14 left; posnodals FW: 11 right, 10 left; HW: 14;
triangle cells FW: 4; HW: 4; supratriangle crossveins FW:
3; HW: 2; subtriangle cells FW: 2; HW: 2; cu-A space
crossveins FW: 3; HW: 3; bridge crossveins FW: 4; HW:
4 right, 3 left; cell rows between IRP2 FW: 3; HW: 3;
cells anal triangle: 3; cells anal loop: 11; rows of cells
anal loop: 3. Abdomen dark reddish brown to black
with yellowish green and light blue spots as follows:
spot at base of genital lobe, lateral stripe on anterior
margin of transverse carina and dorsal spot posterior
to transverse carina in segment II, AD, AL spots of segment III, and PD spots of segments III-IV light blue, remaining pale spots yellowish green (as in figs. 153a-b).
Outer margin of ventral terga VII linear (as in fig.
191a). Ventral tubercle of abdominal segment I
rounded in lateral view, genital lobe lower than 30%
of its length (as in fig. 272); spines of anterior lamina
shorter than 1.5 times of its basal width and blunt [or
pointed] (as in figs. 353-354); ventral portion of
hamuli shorter than hamular fold (as in fig. 311a), tip
of hamular anterior process pointed in posterior and
ventral views (as in figs. 311a-b); right auricle with
two teeth (as in fig. 230), left auricle with two large
teeth and a third one smaller and more external. Vesica spermalis with basal folds in lateral and dorsal lobes
as typical for the genus (illustrated by De Marmels
2001a). Dorso-distal crest of male cercus smoothly
convex, tip blunt (as in fig. 393b).
Allotype: As holotype except frons pale brown; and
thorax and abdomen lacking light blue, all pale spots
being yellowish green to turquoise or olivaceus (PD
spots). Wing venation: Antenodals FW: 18; HW: 12
right, 11 left; posnodals FW: 12 right, 11 left; HW: 15
right, 14 left; triangle cells FW: 5 right, 4 left; HW: 4;
109
T  E,  146, 2003
supratriangle crossveins FW: 3; HW: 2 right, 3 left; subtriangle cells FW: 2; HW: 2; cu-A space crossveins FW: 4;
HW: 3; bridge crossveins FW: 4 right, 5 left; HW: 4 right,
3 left; cell rows at IRP2 fork FW: 3 right, 2 left; HW: 3;
cells anal loop: 12; rows of cells anal loop: 3. Cercus
longer than segments IX-X; maximum width at distal
30%; tip incised (fig. 387c). Dimensions: head width:
9.15 [9-9.2]; HW length: 43.5 [40.2-43.5]; HW width:
14.8 [13.5-14.8]; HW pterostigma length: 2.5 [2-3];
cerci length: ? 5.2 [5-5.24], / 5.0; maximum width
of female cerci: 1.2; total length: 67.5 [58.3-67.5].
Biology. – De Marmels (2001) reported this
species as common at a pond in Tamá National Park
where it was found in sympatry with R. condor, R.
cornigera, and R. marchali. Males patrolled the pond
occasionally approaching the shore; females oviposited into floating grass stems, and exuviae were found
clinging to branches of half submerged bushes. Rare
in collections.
Distribution [7°N, 72°W, 2000-2600 m] (fig.
458). – Venezuela: Táchira (MIZA*; RWGC*; MLPA*;
De Marmels 2001a).
Rhionaeschna eduardoi (Machado) comb. n.
(figs. 37-he, 114-th, 155-ab, 193-te, 232, 419-ge, 274-tg, 314-ha,
355-as, 394-ce, 459-Mp)
Aeshna eduardoi Machado, 1984: 161 (nomen nudum);
Machado 1985a: 45-56 (description ? /: holotype ?
BRAZIL: Minas Gerais-Brumadinho (S. of Belo Horizonte), reserve of Catarina, 20°7’60S 44°13’0W 1200 m,
21-IV-1983, A. & E. & P. Machado leg. (ABMMC) [examined: paratype ? (RWGC)]).
[Aeshna punctata. – Martin 1908: 54-55, 83 (in part: ? in
Selys Coll.); Santos 1966a: 97-98 (in part: records from
Poços de Caldas); Santos 1966c: 65. Misidentification]
Description. – Clypeus light blue, frons bluish yellow, with light blue spots lateral to yellow area surrounding T-spot stem; T-stem parallel-sided, narrower than vertex, transverse arms wider than a line; vertex
yellow with lateral and posterior 30% black (fig. 35a);
dark stripe on frontoclypeal groove, strongly widened
to fronto-ocular groove (figs. 35b-c). Pale pterothoracic stripes yellow, mesanepisternal at basal 80% of
mesanepisternum; mesepimeral and metepimeral linear and complete, metepimeral wider than
mesepimeral (fig. 114). Abdomen reddish brown to
black with yellowish green and light blue spots. Outer
margin of ventral terga VII linear (fig. 193a). Ventral
tubercle of abdominal segment I trapezoidal in lateral
view, as high as 60% of its length, genital lobe higher
than 30% of its width (fig. 274); spine of anterior lamina longer than twice its basal width, narrowly triangular and pointed (fig. 355); ventral portion of hamuli
shorter than hamular fold (fig. 314a), tip of hamular
anterior process blunt in ventral view (fig. 314b); auricles with two teeth (fig. 232). Dorso-distal crest of
110
male cercus smoothly curved, anterior side with denticles, tip pointed (fig. 394b), diverging from other cercus tip in dorsal view (fig. 394a). Maximum width of
female cercus at medial 30%; tip pointed terminating
in a minute spine (Machado 1985a).
Dimensions: head width: 9.6; HW length: 44.147.9; HW width: 14.07; HW pterostigma length: 3.23.6; cerci length: ? 5.7-6.1, / 6.3-6.7; maximum
width of female cerci: 1.3; total length: 68-72.
Diagnosis. – Rhionaeschna eduardoi is most closely
related to R. punctata, with which it shares the presence of complete linear pterothoracic stripes (figs.
114, 118), prominent genital lobe (higher than 30%
its width, figs. 274, 275), male cercus with a smoothly curved crest and acuminate tip (figs. 394-395), and
abdominal color pattern (figs. 155-156). They differ
in several characters (second set of character states for
R. punctata): black of T-spot extended on antefrons
(fig. 37c) vs. not extended (fig. 38c); vertex black
along posterior 30% (fig. 37a) vs. only lateral edges
black (fig. 38a); metepimeral stripes wider than
mesepimeral ones (fig. 114) vs. of equal width (fig.
115); ventral tubercle of segment I high and trapezoidal (fig. 274) vs. low and rounded (fig. 275); and
tip of female cercus pointed and bearing a minute
spine vs. rounded and incised (fig. 395c).
Biology. – Rhionaeschna eduardoi seems to be restricted to iron-rich environments of mountainous regions of Minas Gerais. The adults have been reported
patrolling margins of man-made pools and swamps
fed by streams inside or near the forest (Machado
1985a). Rare in collections. Larva unknown.
Remarks: The species was named in a congress contribution in 1984, which does not constitute a valid
publication (nomen nudum). Therefore, the name
Rhionaeschna eduardoi was first made available for this
species in the description published in 1985a, and it
takes authorship from that date.
(fig. 459). – Brazil: Minas Gerais (Machado 1985;
RWGC*).
Rhionaeschna joannisi (Martin) comb. n.
352-as, 391-ce, 458-Mp)
Aeshna joannisi Martin, 1897: 502 (description /: holotype
/ BOLIVIA: San Antonio (MNHN) [examined: / holotype]); Martin 1908: 56, 84; Calvert 1956: 12, 76-80
(description ?).
Diagnosis. – Provided under R. demarmelsi.
Description. – Clypeus light blue, frons greenish
yellow, greenish blue spots lateral to yellow area surrounding T-spot stem; T-stem narrower than vertex,
approximately parallel sided, transverse arms wider
than a line; vertex black with anterior margin green
(fig. 36a); narrow dark stripe on frontoclypeal groove,
not widened to fronto-ocular groove (figs. 36b-c). Pale
pterothoracic stripes yellowish green, divided in spots
or with deep indentations; mesanepisternal divided
into a basal and a distal spot, mesepimeral and
metepimeral each with a postero-basal and an anterodistal indentation, which surpasses half of stripe width
or divide the stripe into spots (figs. 115-117). Abdomen dark reddish brown to black with yellowish
green and light blue spots. Outer margin of ventral
terga VII sinuous (figs. 192a-b). Ventral tubercle of abdominal segment I rounded in lateral view, genital
lobe lower than 30% of its length (fig. 273); spine of
anterior lamina longer than twice its basal width and
acute pointed (fig. 352); ventral portion of hamuli
anterior process rounded in posterior view and pointed in ventral view (figs. 312a-b), auricles with three
large teeth, sometimes with a fourth smaller tooth (fig.
231). Dorso-distal crest of male cercus angled, tip
pointed (fig. 391b). Maximum width of female cercus
at medial 30%; tip incised (fig. 391c). Dimensions
(values of holotype in square brackets): head width:
9.5-9.8 [9.6]; HW length: 46-50 [47]; HW width: 1415.7 [14.5]; HW pterostigma length: 2.2-3.2 [3.2];
cerci length: ? 5.7, / 6.5 [6.5]; maximum width of
female cerci: 1.5 [1.5]; total length: 68-72.5 [68].
Biology. – Adults have been collected in areas of dry
climate in Bolivia (Dr. K. Tennessen pers. comm.),
and in Ecuador in a very wet climate, flying beats up
and down on the open part of a one-meter wide roadside ditch with slight current and pools, with water
from hillside stream adjacent to forest (Dr. D. R. Paulson pers. comm.). Rare in collections. Larva unknown.
Remarks. – In his description of R. joannisi, Martin
(1897) stated that it was based on a single female from
Bolivia. There are two females from Bolivia labeled as
R. joannisi by Martin at the collection of the MNHN,
none of which was labeled as type by Martin; one
from ‘Bolivie’ with an indication that cerci are broken
(‘app. brisé’) and the other from ‘Bolivie, S. Antonio’.
Following Dr. E. Schmidt’s comments on these specimens Calvert (1956) considered the first one to be the
type because Martin stated only ‘Bolivie’ for the type
locality. However, since a description of the cerci was
included (Martin 1897) and the venation characters as
given by Martin in the original description agree well
with the specimen of San Antonio, I consider this
specimen the holotype. Fifty-eight localities called San
Antonio were found in Bolivia, and unfortunately the
type labels do not include any further information
that would allow for knowing which one corresponds
to the type locality.
Distribution [10ºS? 1°S-2°N, 77-78°W, 11002200 m] (fig. 458). – Bolivia: without locality
(NHMP*; Martin 1897; 1908; Calvert 1956) – Ecua-
dor: Napo (KJTC*), Pastaza (Calvert 1956), and Pichincha (DRPC*) – Colombia: Cauca (Calvert 1956).
Rhionaeschna punctata (Martin) comb. n.
(figs. 38-he, 118-th, 156-ab, 194-te, 233-ge, 275-tg, 315-ha, 356357-as, 395-ce, 459-Mp)
Aeshna punctata Martin, 1908: 54-55, 83 (in part; description ?: lectotype ? BRAZIL: Espirito Santo (MNHN));
Martin 1911: 12; Calvert 1956: 12, 80-88 (in part,
records from Brazil; description /); Santos 1966a: 97100 (in part; description larva); Machado 1985b: 327332 (redescription ?); Martins Costa et al. 2000: 13.
Aeschna depravata Hagen, 1861: 314 (nomen nudum).
Aeschna lobata Hagen, 1861: 314 (nomen nudum).
Diagnosis. – Differentiation of R. punctata from its
closest allies is given under R. eduardoi and R. decessus.
Description. – Clypeus light yellow, frons yellow,
light blue spots lateral yellow area surrounding T-spot
stem; T-stem parallel-sided, narrower than vertex,
transverse arms wider than a line; vertex yellow with
lateral edges black (fig. 38a); dark stripe on frontoclypeal groove, strongly widened to fronto-ocular
groove (figs. 38b-c). Pale pterothoracic stripes yellow,
mesanepisternal at basal 80% of mesanepisternum;
mesepimeral and metepimeral linear and complete, of
equal width (fig. 118). Abdomen dark reddish brown
to black with yellowish green and light blue spots.
Outer margin of ventral terga VII sinuous (figs. 194ab). Ventral tubercle of abdominal segment I rounded
in lateral view, genital lobe higher than half of its
length (fig. 275); spine of anterior lamina longer than
twice its basal width and needle-shaped to narrowly
triangular (figs. 356-357); ventral portion of hamuli
anterior process blunt in ventral view (fig. 315b); auricles with two teeth (fig. 233). Dorso-distal crest of
male cercus smoothly curved, anterior side with denticles, tip pointed (fig. 395b), converging or parallel to
other cercus tip in dorsal view (395a). Maximum
width of female cercus at medial 30%; tip rounded
and incised (395c). Dimensions: head width: 9.4510.5; HW length: 45-50; HW width: 13.1-15.3; HW
/ 5.5; total length: 68-82.
Biology. – The species has been reported as abundant from Brejo da Lapa, Itatiaia, Brazil, where adults
and larvae were collected at ponds with abundant
macrophytes (Santos 1966a). The larva was described
by Santos (1966a) based on reared material from this
locality.
Remarks. – Machado (1985b) examined the type
series of Rhionaeschna punctata deposited in the
MNHN and ISNB collections and concluded that it includes three different species: R. punctata, R. eduardoi
and R. jalapensis. He designated a lectotype and provided a redescription of the male.
111
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459). – Brazil: Rio Grande do Sul (MNRJ), Santa
Catarina (Calvert 1956), São Paulo (MNRJ*; USNM*;
Santos 1966a; Martins Costa et al. 2000), Rio de
Janeiro (MNRJ*; Calvert 1956; Hagen 1861; Santos
1966a), Minas Gerais (MNRJ; Santos 1966a), and Espirito Santo (Martin 1908).
PHYLOGENETIC ANALYSIS
The first analysis generated eight equally most parsimonious trees (113 steps, CI 0.49, RI 0.88). Successive weighting resulted in 332 trees (CI 0.72, RI 0.95).
In the strict consensus cladogram of the 332 trees (fig.
438, CI 0.70, RI 0.94), each node is numbered, and
synapomorphies supporting each node are listed with
homoplasies indicated with an asterisk.
The analysis shows the genus Aeshna Fabricius to be
paraphyletic with some of its species more closely related to the Andaeshna-Anaciaeschna-Hemianax-Anax
clade (node 13-15, fig. 438). Paraphyly of Aeshna had
already been suggested by Peters (1987), who based
his analysis on wing characters of the European Aeshnidae. Part of ‘Aeshna’, corresponding mainly to the
Holarctic species form a natural group (node 1-2, fig.
438) characterized by the abdominal auricles bearing
four or five denticles (figs. 428-430). The type species
of Aeshna, A. grandis, belongs to this group, which
therefore retains the name. Two monophyletic groups
can be distinguished within Aeshna s.str., one of them
(node 2-3, fig. 438) characterized by narrow twocelled anal triangles (fig. 405), and the second one
(node 2-8, fig. 438) by the following five autapomorphies: (1) absence of sub-basal tooth or carina in dorsum of Xth segment, (2) male cercus with an apical
ventral spine on inner margin, (3) cercus inner margin
concave distally in dorsal view (figs. 426a, b), (4) dorso-distal crest curved in dorsal view (figs. 426c), and
(5) apical spines of epiproct large and directed anteriorly (fig. 426d).
The remaining ‘Aeshna’ species and Aeshninae genera form a monophyletic group (node 1-10, fig. 438)
characterized by the marked bending of MA (figs. 396404, 406-407) and short parallel fusion of veins delimiting the anal triangle (figs. 409-410, 412) (reversed to a long fusion in the punctata-group of
Rhionaeschna, fig. 411). Within this group, some
‘Aeshna’ species (‘A.’ brevistyla, ‘A.’ affinis, ‘A.’
williamsoniana, ‘A.’ ellioti, ‘A.’ mixta and ‘A.’ isoceles)
are problematical, because they are interspersed at the
base of different genera (fig. 438, nodes 11-12, 13-14,
15-17). ‘Aeshna’ brevistyla is the sister group of the remaining Aeshninae (node 10-11, fig. 438). ‘Aeshna’
affinis, ‘A.’ williamsoniana, ‘A.’ ellioti and ‘A.’ mixta (as
well as ‘A.’ minuscula not included here) have a basal
position in the clade including Andaeshna and Anaci112
aeschna (and Hemianax-Anax, see fig. 437), and ‘A.’
isoceles constitutes the sister group of Andaeshna. A
small group of African species of ‘Aeshna’ consisting of
‘A.’ rileyi and ‘A.’ subpupillata from South Africa, and
‘A.’ moori from Central Africa (‘A.’ moori was not examined in the present analysis but belongs to the same
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 cercus in dorsal view (as in fig. 427b) but this is a homoplastic character present in several other Aeshnidae,
and the support for this grouping is consequently
weak. The African group (‘A.’ rileyi, ‘A.’ subpupillata,
‘A.’ moori) is characterized by the greatly enlarged and
posteriorly projected genital lobe devoid of denticles
(fig. 421), and the morphology of the last segment of
the vesica spermalis, which bears a unique large
medio-longitudinal fold on the ventral lobe (figs.
436a, b).
The autapomorphy defining Rhionaeschna is the
presence of a conical tubercle bearing denticles on abdominal sternum I (figs. 234-275, 418-419); sternum
I is planar in Oreaeschna, Aeshna s.str., ‘Aeshna’ species
of uncertain position (‘A.’ affinis, ‘A.’ mixta, ‘A.’ brevistyla, ‘A.’ ellioti, ‘A.’ williamsoniana, ‘A.’ moori, ‘A.’
rileyi and ‘A.’ subpupillata), and Andaeshna (fig. 425).
Abdominal sternum I has a transverse ridge in Anaciaeschna and ‘A.’ isoceles (fig. 421). Castoraeschna has
also a tubercle in sternum I, but it is cylindrical, much
higher than in Rhionaeschna and lacks denticles (fig.
424); I consider their presence in Rhionaeschna and
Castoraeschna due to convergence.
According to the cladogram, Rhionaeschna species
form four major groups as follows, but their relationships with one another are still unclear:
First, Rhionaeschna draco shows numerous autapomorphies, i.e. frons projected dorsally, higher than
vertex (fig. 1b); pterothorax and abdominal segments
VI-X without pale markings (figs. 39, 119); hamulus
with a medial digitiform projection on its antero-ventral margin (fig. 276); female cercus with long terminal spine (fig. 358c), but except for the presence of
ventral tubercle I, it does not share other derived characters with other species of Rhionaeschna.
Second, the Marmaraeschna clade (node 20-21, fig.
438), characterized by five autapomorphies as follows:
(1) the recessed frontal carina located on postfrons giving a rounded contour to the antero-dorsal margin of
head (figs. 2-8b), (2) pterothorax with marbled pattern of black markings and pale areas (figs. 40-47), (3)
hamular anterior process high (figs. 277-284), (4) anterior lamina spine wide throughout its length in lateral view (figs. 317-324), and (5) external margin of
male cercus approximately linear in lateral view (in R.
intricata it can be slightly concave) (figs. 359-365).
Two clades are defined within this group, one includ-
ing R. brevifrons, R. fissifrons and R. pallipes (node 2122, fig. 438), which share a flattened frons (figs. 3-4,
7) and slightly constricted abdomen (figs. 121-122,
124) and the other (node 21-23) comprising R. brevicercia, R. intricata, R. obscura and R. vigintipunctata
characterized by the high genital lobe (figs. 236, 238,
240, 242).
Third, the cornigera- and punctata-groups constitute another clade (node 20-24, fig. 438), characterized by the male cercus lacking a sub-basal tooth (figs.
382-395). Both groups constitute sister groups; the
synapomorphies of the cornigera-group species (node
24-28, fig. 438) consist of the male sterna IX posterior
to genital opercula and X with a bright light-blue spot
(figs. 143-150b) and the male cercus with concave lateral margin and extreme apex pointed and ventrally
bent (figs. 382-389). Synapomorphies of the punctatagroup species (node 25-26, fig. 438) include the numerous small denticles widely distributed on the lateral and ventral surface of the genital lobe (figs.
270-275) and the whole tip of male cercus ventrally
bent (figs. 290-295). Within the cornigera-group, R.
manni, R. psilus and R. vazquezae form a clade (node
29-30, fig. 438) distinguished by the angled dorso-distal crest of male cercus (figs. 384, 388-389). Two
clades are defined within the punctata-group, one including R. biliosa, R. condor and R. joannisi (node 2627, fig. 438), which share an angled dorsodistal crest
in male cercus (figs. 290-292), and another one with
R. decessus, R. eduardoi and R. punctata (node 26-28,
fig. 438) characterized by the high male genital lobe
(figs. 274-275).
Fourth, Schizuraeschna, Neureclipa and variegatagroup species form a monophyletic unit (node 20-30,
fig. 438), characterized by the origin of two rows of
cells between RP1 and RP2 distal to pterostigma or at its
distal end (figs. 398-400), lateral carina of male cercus
concave in lateral view and sub-basal tooth present (reduced in R. confusa and R. californica) (figs. 366-381).
Schizuraeschna and Neureclipa each represent a clade;
the Schizuraeschna clade (node 30-31, fig. 438) is characterized by the male cercus inner margin with a ventral process at distal 25% and dorso-distal crest triangular in lateral view (figs. 366-369) and the Neureclipa
clade (node 30-34, fig. 438) by the free supratriangle
(Fig 399). Within Schizuraeschna clade, R. jalapensis,
R. multicolor and R. mutata constitute a subgroup characterized by the larger ventral process and the tip of
male cercus ventrally curved (figs. 367-369) and R.
multicolor and R. mutata are sister species, having the
entire tip of male cercus ventrally curved (figs. 368369). Within Neureclipa, R. elsia and R. galapagoensis
are sister species, the synapomorphy joining them being the presence of only a few apical denticles on the
ventral tubercle of abdominal segment I (figs. 250251). The relationship among the species here includ-
ed in the ‘variegata-group’ and between them and
Schizuraeschna and Neureclipa –groups is still uncertain
(basal polytomy in node 20-30, fig. 438) since no
synapomorphy is known that would join the variegatagroup species in its own clade.
Thus, the only subgenus traditionally defined for
the neotropical species of ‘Aeshna’ which is not monophyletic is Hesperaeschna, comprising, according to
Calvert (1956), draco, the species belonging to the
cornigera-punctata clade, the species of the ‘variegatagroup’ belonging to the clade Schizuraeschna-Neureclipa-variegata and ‘Aeshna’ williamsoniana. My examination of the male holotype of ‘A.’ williamsoniana
(deposited at UMMZ), revealed the absence of the ventral tubercle in the first abdominal segment, which
would exclude this species from the group. Besides, it
does not share cerci morphology, wing venation or
color pattern with any of the groups of Rhionaeschna.
Its phylogenetic position is unclear, but it does not belong to Rhionaeschna; its closest relative in the
Neotropical region most likely belongs to the clade including Andaeshna and Anax.
The phylogenetic relationships suggested in the present study should be considered tentative; it is clear
that further work is required to further understand relationships within the main groups discussed here.
However, it can be concluded that current classification of the Neotropical components comprising ‘Aeshna’ is artificial. A reclassification of the group including
Rhionaeschna, Andaeshna, Anaciaeschna, Hemianax and
Anax is necessary but it will only be possible once the
positions of the enigmatic species of ‘Aeshna’ (‘A.’ affinis, ‘A.’ brevistyla, ‘A.’ ellioti, ‘A.’ isoceles, ‘A.’ mixta and
‘A.’ williamsoniana) are elucidated.
BIOGEOGRAPHIC ANALYSIS
Rhionaeschna is a New World genus distributed
from southern Argentina to southern Canada. The
group is mostly Neotropical, with its highest diversity
along the Andean mountain range of South America
between Venezuela and Bolivia (fig. 466). It is not
known from the Amazonian basin. Generalized tracks
of the Rhionaeschna groups used in the present work,
which with the exception of the variegata-group represent monophyletic units, are depicted in figs. 460465. Marmaraeschna species are limited to South
America from central Chile and the hills of Central
Argentina along the Andes to Venezuela (fig. 460).
The punctata-group shows a restricted distribution
forming a narrow semicircle from Venezuela along the
Andes to Bolivia and Southeastern Brazil (fig. 465)
whereas the cornigera-group is the most widely distributed encompassing the whole range of the genus with
the exception of North America (fig. 464). Schizuraeschna species are restricted to Central and North
113
T  E,  146, 2003
America (fig. 461), and Neureclipa species to South
America (fig. 462).
Several species of Rhionaeschna have a restricted distribution (figs. 439-459), and six areas of endemism
were established based on them (fig. 466). These areas
are (1) Austral South America, encompassing S Argentina and Chile, with R. brevifrons, R. pallipes, R.
vigintipunctata, R. absoluta, R. bonariensis, R. confusa,
R. variegata, R. planaltica, R. haarupi, and the endemic R. diffinis, (2) Paranense, including Uruguay, S
Brazil, most of Paraguay and NE Argentina, with R.
bonariensis, R. confusa, R. planaltica, and endemics R.
brasiliensis, R. pauloi, R. decessus, R. eduardoi, and R.
punctata; (3) Andes, ranging from NW Argentina and
Chile to Venezuela along the Andes, with R. brevifrons, R. pallipes, R. vigintipunctata, R. absoluta, R.
cornigera, R. haarupi, R. planaltica, R. psilus, R. variegata, and endemics R. brevicercia, R. fissifrons, R. intricata, R. obscura, R. elsia, R. galapagoensis, R. biliosa, R.
joannisi, R. demarmelsi, R. condor, R. marchali, R. peralta, R. tinti; (4) Tepuis, encompassing SE Venezuela,
with R. planaltica, and endemics R. draco and R. nubigena; (5) Central America, ranging from Panama to
SE U.S.A., with R. californica, R. multicolor, R. cornigera, R. psilus, and endemics R. dugesi, R. jalapensis, R.
manni and R. vazquezae; and (6) North America, including U.S.A. and S Canada, with R. californica, R.
multicolor and the endemic R. mutata.
Only a few species share different areas of endemism as reflected in the low association values obtained in the similarity analysis (fig. 468). Austral
South America and Andes form the highest similarity
nucleus to which Paranense and later Tepuis are
joined. North America and Central America form a
second nucleus joined to the first one by a low association value.
Parsimony analysis of endemicity resulted in two
area-cladograms (length 52, CI 86, RI 66). One of
them shows the same relationship among areas obtained with the similarity analysis (fig. 467a); the second differs in the position of the Tepuis as the sister
group of the remaining areas (fig. 467b).
The relationships among areas of endemicity of
Rhionaeschna agree broadly with the hierarchy of the
biogeographic subdivisions of South America proposed by Cabrera & Willink (1980) and Morrone
(1999). There are no subantarctic endemics, and in
accordance with the schemes suggested by Morrone
(1999) and Ringuelet (1961), the species occuring in
the Subantarctic province of the Antarctic region of
Cabrera & Willink (1980) are also present in Patagonian and Andean provinces. Patagonian, Monte and
Chilean provinces, included in the Austral South
America area, and the drier regions of the Andean area
(corresponding to the Desierto, Prepuneña, Puneña
and Altoandina provinces) are encompassed in the
114
same domain (Andino-patagónico of Cabrera & Willink 1980), or even the same region (Andina of Morrone 1999) in the biogeographic schemes of South
America, which agrees well with the high association
value found in the present study between these two areas (figs. 467-468). Yungas, comprising forested areas
of the E Andes, belongs to the same biogeographic division as the Paranense area in all schemes (Amazonian domain or subregion), and this association is exemplified by the distribution of the punctata-group,
with species occurring either in the Yungas (R. biliosa,
R. condor, R. demarmelsi, R. joannisi), or in the Paranense province (R. decessus, R. eduardoi, R. punctata).
Tepuis are considered a domain different from the
Amazonian domain by Cabrera & Willink (1980),
and as a province of the Amazonian subregion by
Morrone (1999). According to Halffter (1965), when
the Central American land bridge connected South
and North America during the upper Cretasic and
lower Eocene, a major immigration of South American insects towards North America took place. During
the Tertiary, flora and insects of Neotropical origin
spread northward to about 49 degrees in western
North America and lower latitudes eastward, and progressively retracted during Miocene and Pliocene,
with relicts being found today in southern parts of the
western and eastern United States. But several
Neotropical insects still exist as far north as Alberta,
New York and Michigan (Halffter 1965), which are
related to South American elements (i.e. Triatoma
[Heteroptera: Reduviidae], several species of Scarabeidae and Cerambicidae [Coleoptera]). Thus, North
and Central American areas of endemism in Rhionaeschna may be the result of this same process, where
several species evolved in North and Central America
after the expansion and subsequent isolation from
South America.
Species of Rhionaeschna are lacking from the Brazilian shield (Amazonic province of Cabrera & Willink
1980; Morrone 1999). Distribution of Rhionaeschnarelated taxa show a low diversity in Africa (‘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 in S Europe and Middle East, and Anaciaeschna,
Hemianax and Anax species with the highest diversity
in the Indo-Australian region). These facts suggest a
trans-Pacific rather than trans-Atlantic (Gondwanian)
track for the group to which Rhionaeschna belongs, as
has been hypothesized for other groups of similarly
distributed Odonata (i.e. Epallagidae (Euphaeidae +
Polythoridae), Gynacanthini, De Marmels 2000;
Megapodagrion-complex, De Marmels 2001b).
ACKNOWLEDGEMENTS
I thank Dr. Rosser W. Garrison for his generous assistance, support, and constructive criticism during
the realization of this work. Thanks are due to Dr.
Dennis Paulson, Dr. Jürg De Marmels, Dr. Günther
Peters, Dr. Thomas W. Donnelly, and Mr. Jan van
Tol for critically reading the manuscript and providing helpful corrections and comments. Dr. Dennis R.
Paulson, Dr. Kenneth J. Tennessen and Dr. Thomas
W. Donnelly generously provided unpublished information concerning biology and distribution of several
species. The following curators and individuals placed
specimens and information at my disposal: Dr. Jürg
De Marmels (MIZA) sent pictures, illustrations and
measurements of the female of Rhionaeschna demarmelsi and loans; Dr. Jérôme Constant (ISNB) and
Dr. Fritz Geller Grimm (MWNH) provided photographs of type specimens and assisted during visits
to their collections; Dr. Jean Legrand (MNHN), Dr. Jason Weintraub (ANSP), Dr. Heinz G. Schroeder
(SMFD), Dr. Bernhard Misof (ZFNB), Dr. Philip D.
Perkins (MCZ), Dr. Adolfo Cordero Rivera (Pontevedra), Dr. Victor Monserrat (Madrid), Dr. Gloria
Masó (Barcelona), Dr. Syoziro Asahina and Dr.
Masaaki Tomokuni (Tokyo) provided information
about type specimens in their collections. For their
kind assistance during examination of their collections
or loan of material I also thank Mr. Mark O’Brien
(UMMZ), Dr. Oliver Flint and Dr. Nancy Adams
(USNM), Dr. Brian Brown and Mr. Brian Harris
(LACM), Dr. Dennis Paulson (Seattle), Dr. Kenneth
Tennessen (Florence), Dr. Janira M. Costa (MNRJ),
Dr. Angelo Machado (Rio de Janeiro), Dr. Enrique
González Soriano (UNAM), Dr. Giuliano Onore
(QCAZ), Dr. Ariel Camousseight (MNNS), Dr. Jorge
Solervicens (IEUM), Dr. Carlos Esquivel (INBC), Mr.
Jan van Tol (RMNH), Dr. David Goodger (BMNH), Dr.
Graham S. Vick (Crossfields), Dr. Ulrike Aspöck
(NMW), Dr. Günther Peters (ZMHB), Mr. Bill Mauffray (FSCA), Dr. Mitsutoshi Sugimura (Nakamura),
Dr. Axel Bachmann (MACN), Mr. Hector Ferreyra
(MLPA), Dr. Sergio Roig (IADIZA), Dr. Eduardo
Dominguez and Dr. Carlos Molineri (IMLA).
This research was supported by a fellowship from
the FUNDACION ANTORCHAS, and it was carried out
during a postdoctoral tenure at the Natural History
Museum of Los Angeles County. I am indebted to
both institutions for providing support and facilities
necessary to conduct this study.
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119
T  E,  146, 2003
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Received: 12 July 2002
Accepted: 5 December 2002
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
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
9-14
11-16
11-19
14-18
14-16
16-20
17-21
16-17
13-18
18-19
13-19
19-22
19-22
20
18-19
13-17
17-22
14-15
13-14
8-10
7-9
7-10
8-9
8-10
8-10
8-11
9-11
16-17
9-11
11-12
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
11-14
9-10
8-12
12-13
11-14
12-14
13-15
14
11-13
9-11
12-14
9-10
Ax Fw Ax Hw
13-16
12-13
11-12
10-12
12-13
11-13
12-13
10-14
10-11
9-11
8-10
9-11
7-8
7-9
8-10
6-7
10-11
10
9-11
7-9
10-12
5-9
7-9
9-12
12-13
8-10
9-15
12-15
12
10-11
11-12
11-14
11-15
13-15
18-19
11-12
10-13
12-15
12-13
16-19
12-14
10-13
11-12
12-13
12-15
12-14
13-14
12-13
10-12
7-13
11-13
8-10
9-11
9-11
6-9
11-13
10-12
9-11
7-9
10-14
6-11
9-11
11-13
12-13
8-12
12-15
13-16
12
12-14
13-14
13-17
15-17
16-20
23
12-14
11-16
16-18
13-14
3
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
0-3
2
1-4
2
1-2
1-3
0-3
2-4
2-3
2-4
2-4
1-3
2-4
3
3-4
4-5
4-6
4
3-4
2-3
3-4
2
2-3
1-2
1-2
1-2
1-2
1-2
1-2
1-2
2
2
2
2-3
0-1
0-1
0-1
0
0-1
0-3
1-2
1-3
1-2
1-2
1-2
1-3
2-3
2
2
3
1-2
2-3
2-3
3-4
2-3
3-5
3
2
2-3
3
2
Px Fw Px Hw sptr Fw sptr Hw
4-6
3-5
2-4
4-5
4-5
4-5
4-5
4-5
4-5
4-5
4-5
5
2-5
2-4
2-4
2-4
3-4
3-5
4
4-5
4
2-4
2-4
3-6
4-5
4
4-6
5
5
4-5
4-5
5
5
5
6-7
4-5
4-5
5
4
tr Fw
4-5
3-4
2-3
3-4
3-4
4-5
3-4
3-4
4
4
4
4-5
2-4
2-4
2-4
2-4
3-4
4-6
4
3-5
4
3-4
3-4
3-6
4
4
4-5
4
4
4-5
4
5
4
4-5
6-7
4
4-5
4-5
4
2
2
1-2
2
1-2
1-2
2
2
2
2
2
2
1
1
1-2
1
1-2
1-2
1-2
2
2
1-2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1-2
2
1-2
2
2
2
2
2
2
2
1
1
1-2
1
1-2
1-2
1-2
2
2-3
1-2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
5-6
4-5
3-4
4-5
4
3-4
3-4
4-5
3-5
3-4
3
4
3
3-4
3
3-4
4
3-5
3-4
3
3-4
2-3
2-4
3-4
3-5
3-4
4
4-5
4
3-5
4
2-5
3
5
?
4
3-5
4-5
3
4
3
3-4
3-4
3-4
3
3-4
2-3
3-4
3
3
3-4
2-3
3
3
3
3
2-4
2-4
3
2
2-3
2-3
2-3
3
3
3
4
3-4
2-5
3
3
5
3
?
3
2-4
3-4
3
3-4
3-4
2-3
2-3
3
3-4
2-4
2-4
2-4
2-4
2-3
2-4
2-3
2-3
2-3
2-3
2-3
2-4
2-3
2-3
2-3
1-3
1-3
2-4
2-4
3
3
3
3
2-4
3
3
3-4
3
?
3
2-3
3
3
3-4
3-4
2-3
2-3
3
3-4
2-4
2-4
3-4
2-4
2-3
2-4
2-3
2-3
2-3
2-3
3
2-4
2-3
2-3
3
1-3
2-3
2-4
2-4
3
3
3-4
3
2-4
3
3
3-4
3
?
3
3-4
3
3-4
tr Hw sbtr Fw sbtr Hw cua Fw cua Hw frk Fw frk Hw
3-4
2-4
2-3
2-4
2-3
2-5
2-4
2-4
3-4
3-4
3
3-4
2-3
2-4
2-3
2-3
2-3
2-4
3
2-3
3
1-3
1-4
2-5
3-4
3-4
2-4
3
2-3
3-4
3-4
3
4-5
5-6
?
5
3
3-5
3
2-3
2-3
1-3
2-4
2-3
2-4
1-3
2-3
3
3
2-4
3
2-3
2-3
2-3
2
2
2-4
3
2
3
1-3
2-3
2-4
3-4
3
2-3
2-3
2
2-4
2-3
3
4-6
5
?
4
3
3-4
3
br Fw br Hw
8-12
10-11
8-11
12-14
8-9
9-13
9-12
9-12
8-11
8-11
8-11
9-11
5-11
5-10
5-9
5-8
6-8
7-13
7-8
6-12
9-11
4-11
6-12
6-18
9-13
9-10
8-13
9-11
8-11
9-13
8-10
13-14
11-14
11-14
?
9-11
7-10
13-15
9-10
3-4
3
3
3
3
3
3
3
3
3
3
3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
2-3
3
2-3
2-3
2-4
3-4
3
3
3
2-3
3
3
3
3
3
?
3
3
3
3
al cells al rows
Table 1. Wing characters. Fw: fore wing; Hw: hind wing; Ax: antenodal crossveins; Px: posnodal crossveins; sptr: supratriangle crossveins; tr: triangle cells; sbtr: subtriangle cells; cua:
cubito-anal crossveins; frk: number of cells between IRP2 fork at pterostigma level; br: bridge crossveins; al: anal loop.
121
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
Weight of
character
0
1
0
4
0
2
1
0
0
3
1
0
0
4
1
0
0
5
1
0
0
6
0
3
0
7
1
0
0
8
0
3
0
9
0
1
1
0
0
5
1
1
0
6
1
2
1
0
1
3
1
0
1
4
0
6
1
5
1
0
1
6
0
6
1
7
0
5
1
8
0
4
1
9
0
4
2
0
0
4
2
1
0
1
2
2
1
0
2
3
1
0
2
4
1
0
2
5
1
0
2
6
1
0
2
7
1
0
2
8
0
2
2
9
0
3
3
0
1
0
3
1
0
1
3
2
0
1
3
3
1
0
3
4
1
0
3
5
0
4
3
6
1
0
3
7
1
0
3
8
1
0
3
9
0
1
dictatrix
isoceles
jaspidea
triangulifera
andresi
rufipes
draco
brevicercia
brevifrons
fissifrons
intricata
obscura
pallipes
vigintipunctata
cornigera
planaltica
haarupi
nubigena
psilus
manni
vazquezae
absoluta
bonariensis
diffinis
elsia
galapagoensis
brasiliensis
californica
confusa
marchali
peralta
tinti
variegata
biliosa
demarmelsi
condor
joannisi
decessus
eduardoi
punctata
dugesi
jalapensis
multicolor
mutata
williamsoniana
brevistyla
affinis
clepsydra
eremita
interrupta
canadensis
crenata
cyanea
constricta
walkeri
palmata
persephone
0
0
0
0
0
0
0
1
1
1
1
1
0
1
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
0
0
0
0
0
0
2
2
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
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
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
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
0
0
0
0
0
0
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
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1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
?
1
1
1
1
1
1
1
1
1
0
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1
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1
1
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1
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1
0
0
0
0
?
0
0
1
1
1
1
1
0
1
0
0
1
0
0
0
1
0
0
0
0
0
1
1
2
2
0
2
2
2
2
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2
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0
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0
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0
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0
0
0
0
0
0
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0
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0
0
0
0
0
0
0
0
0
0
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0
0
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2
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1
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1
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1
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0
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0
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2
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1
1
1
1
1
1
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1
1
1
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1
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0
0
0
0
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0
0
1
1
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0
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0
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1
1
1
1
1
0
0
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0
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0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
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0
0
0
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0
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0
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2
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0
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0
0
0
0
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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
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0
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0
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0
0
0
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0
0
1
1
1
1
1
0
0
0
0
0
0
0
0
0
1
1
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
0
0
0
0
0
0
1
1
1
1
1
0
0
0
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
3
0
0
0
1
0
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
1
1
1
1
1
1
0
2
2
2
2
2
2
2
2
2
2
0
0
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0
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0
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0
0
0
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0
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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
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
0
0
0
0
0
0
0
0
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0
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1
1
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1
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0
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0
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0
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0
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0
0
0
0
0
0
0
0
0
0
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1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
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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
1
1
0
0
1
1
0
0
1
0
0
0
0
0
0
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0
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0
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0
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0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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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
0
0
0
0
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0
1
1
1
1
1
0
1
1
1
1
1
2
1
1
1
2
1
1
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
0
0
0
0
0
0
0
2
2
2
2
1
1
2
2
2
2
2
2
0
0
0
0
0
0
1
0
0
0
0
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
1
1
1
1
0
1
1
0
0
0
0
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0
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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
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0
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0
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0
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0
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1
1
1
1
1
0
0
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0
0
0
0
0
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0
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0
0
0
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0
0
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0
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0
0
0
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0
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0
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0
0
0
0
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1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
2
2
2
1
1
1
1
1
1
1
1
1
1
1
1
2
1
2
2
1
1
1
2
2
2
2
0
0
0
1
1
1
1
1
3
3
3
3
3
0
0
0
0
0
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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
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0
0
0
0
0
0
1
1
1
1
1
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
0
1
1
1
1
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
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
1
1
1
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
2
2
2
2
2
2
2
0
1
2
2
0
0
0
0
0
0
1
0
1
1
1
1
1
122
petalura
umbrosa
ellioti
grandis
juncea
tuberculifera
viridis
verticalis
septentrionalis
sitchensis
caerulea
mixta
nigroflava
rileyi
osiliensis
subpupillata
subartica
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
1
1
1
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
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
?
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
0
0
1
1
1
1
1
1
1
1
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
2
2
2
2
1
1
1
0
2
0
2
?
2
0
0
0
1
2
1
1
2
0
0
0
0
1
0
1
?
2
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
2
0
0
0
2
2
2
0
0
0
0
?
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
2
0
2
2
1
1
1
2
2
2
2
2
2
1
2
0
1
0
1
1
1
0
1
1
1
1
1
1
1
1
0
1
0
1
?
1
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
0
0
0
0
0
0
0
0
1
0
1
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
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
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
2
2
2
1
2
2
1
?
1
1
1
?
1
2
0
0
0
0
0
1
0
0
0
0
0
0
0
1
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
1
0
1
?
0
3
3
0
1
1
1
0
1
1
1
1
0
1
1
1
1
1
1
1
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
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
2
0
0
0
0
1
0
2
0
2
0
1
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 vigintipunctata, 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 absoluta, 20 bonariensis, 21 diffinis, 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+punctata), 53 punctata-clade, 54 (cornigera+punctata).
Areas
000000000111111111122222222223333333333444444444455555
123456789012345678901234567890123456789012345678901234
root
North Am
Central Am
Andes
Tepuis
Paranense
Austral
000000000000000000000000000000000000000000000000000000
0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0
000000000001111011000000010000001101000111100000000001
011111111110000000100111100011111110001101111100101011
100000000000000000000000000000000000101000100000000001
000000000000000000010000101100001000011000100011010111
001000111110000000111000101000101010001000100000000001
123
T  E,  146, 2003
1
frons
projected
dorsally
a
b
VE: Auyantepui
2
stem of T-spot
not strongly
narrowed
anteriorly
a
EC: Holotype
draco
margin
rounded
b
c
VE: Auyantepui
VE: Auyantepui
slight medial
concavity
c
EC: Guama Yacu
brevicercia
EC: Guama Yacu
frons
flattened
3
a
CH: Huasco Bajo
b
a
PE: Lectotype maita
b
Lectotype brevifrons
c
Lectotype brevifrons
PE: Lectotype maita
c
PE: Lectotype maita
c
AR: Allotype
brevifrons
4
deep
medial
cleft
frons
flattened
a
AR: Allotype
b
Figs. 1-4. Head. – a, dorsal view; b, lateral view; c, frontal view.
124
AR: Allotype
fissifrons
stem of T-spot
strongly narrowed
anteriorly
a
PE: Celendin
a
BO: Holotype
b
b
EC: Pallatanga
intricata
obscura
BO: Chapare
AR: Río Colorado
b
a
AR: Río Colorado
b
EC: Pallatanga
c
BO: Chapare
frontal carina
linear
frons
flattened
a
c
AR: Río Colorado
c
AR: Río Colorado
AR: San Lorenzo
c
AR: San Lorenzo
pallipes
vigintipunctata
125
T  E,  146, 2003
margin
angled
a
US: Leslie Canyon
a
MX: Cuernavaca
b
b
US: Leslie Canyon
c
US: Leslie Canyon
MX: Cuernavaca
c
MX: Cuernavaca
US: Havasu Canyon
c
US: Havasu Canyon
US: Scotia
c
US: Scotia
dugesi
jalapensis
T-spot stem
narrower than
vertex
a
US: Havasu Canyon
b
multicolor
T-spot stem
wider than
vertex
a
US: Scotia
b
mutata
126
T-spot sides
converging
anteriorly
dark stripe
on frontoclypeal
groove
a
b
AR: Lago Aluminé
AR: Lago Aluminé
absoluta
c
AR: Cafayate
T-spot sides
parallel
dark stripe on
fronto-ocular
groove narrow
clypeal lobe
angled
a
b
AR: Punta Lara
c
AR: Punta Lara
bonariensis
AR: Dique Cabra Corral
dark stripe
on frontoocular
groove wide
a
b
AR: Trevelin
AR: Trevelin
diffinis
c
AR: Trevelin
c
PE: nr Villa
c
EC: Is. Santa Cruz
no dark
stripe on
frontoclypeal
groove
a
b
PE: nr Villa
clypeal lobe
rounded
elsia
PE: nr Pacasmayo
dark
stripe
a
EC: Is. Santa Cruz
b
galapagoensis
EC: Is. Santa Cruz
127
T  E,  146, 2003
frontal carina
angled
a
b
BR: Holotype
c
BR: Holotype
US: Palo Alto
c
US: Palo Alto
AR: Punta Lara
c
AR: Punta Lara
BR: Holotype
brasiliensis
frontal carina
evenly curved
dark stripe
on frontoclypeal
groove
a
b
US: Palo Alto
californica
no dark
stripe
a
a
AR: Punta Lara
EC: Cuenca
b
b
confusa
marchali
128
EC: Cuenca
c
EC: Cuenca
A>B+C
C < 2B
A
B
C
a
PE: Paralectotype Apurimac
a
CH: Holotype
b
BO: Copacabana
c
CH: San Pedro de Atacama
c
peralta
b
PE: Paralectotype Apurimac
CH: Holotype
tinti
A
B
A<B+C
C = or > 2B
C
a
AR: Lago Villarino
b
a
AR: El Calafate
c
AR: Cari Laufquen
variegata
129
AR: Cari Laufquen
T  E,  146, 2003
a
CO: Lectotype
b
VE: Altos de Tiara
c
VE: Altos de Tiara
a
GU: Tamahú
b
GU: Tamahú
c
GU: Tamahú
cornigera
stem of Tspot abruptly
narrowed
a
AR: San Carlos
b
AR: San Carlos
c
AR: San Carlos
haarupi
a
MX: Holotype
b
MX: Holotype
manni
130
c
MX: Holotype
a
a
VE: Cerro Yaví
BR: Paratype, Santana do Riacho
b
b
VE: Cerro Yaví
nubigena
c
c
VE: Cerro Yaví
pauloi
stem of T-spot gradually
narrowed anteriorly or
parallel sided
a
AR: Pereyra Iraola
b
AR: Pereyra Iraola
c
AR: Pereyra Iraola
c
AR: Dique Itiyuro
planaltica
a
AR: Dique Itiyuro
b
AR: Dique Itiyuro
psilus
131
T  E,  146, 2003
a
MX: Paratype, Acahizotla
b
MX: Paratype, Acahuizotla
c
vazquezae
transverse
arms of
T-spot
narrow
no dark
stripe
a
b
EC: Manabi
EC: Manabi
c
EC: Manabi
c
VE: Paratype, Tamá NP
c
biliosa
T-spot stem
wider than
vertex
a
b
condor
transverse
arms of
T-spot wide
dark
stripe not
widened
a
b
demarmelsi
132
T-spot stem
narrower
than vertex
a
b
BO: Holotype
EC: E Cornejo Astorga
joannisi
dark stripe
strongly
widened
a
BR: Paratype, Catarina
c
c
black of
T-spot
extended on
antefrons
b
eduardoi
black of
T-spot not
extended on
antefrons
a
BR: Umuarama
b
BR: Umuarama
punctata
c
133
BR: Umuarama
T  E,  146, 2003
pterothorax
uniform in color
VE: Auyantepui
black spots
reduced
EC: Guama Yacu
draco
AR: Paratype, Río Colorado
fissifrons
brevifrons
EC: Pallatanga
BO: Río Carmen Mayu
intricata
AR: Lesser
US: Leslie Canyon
US: St. David
US: Brown's Lake
vigintipunctata
dugesi
multicolor
multicolor
AR: San Antonio
AR: Lago Aluminé
absoluta
absoluta
Figs. 39-58. Pterothorax, lateral view.
134
PE: Abancay
brevicercia
obscura
CR: Monteverde
jalapensis
US: Scotia
mutata
AR: Puerto Pirámide
absoluta
PE: Lectotype maita
brevifrons
AR: Río Colorado
pallipes
GU: Los Aposentos
jalapensis
AR: Villa Ventana
absoluta
AR: Dique Campo Alegre
absoluta
AR: Lago Aluminé
absoluta
CH: N Valdivia
diffinis
AR: Los Alerces NP
diffinis
CH: Camarones
elsia
AR: Río Santa Lucía
bonariensis
AR: Lago Aluminé
CH: Coñaripe-Carringue
diffinis
diffinis
AR: Lago Tromen
AR: Paso Futaleufú
diffinis
AR: Ayo. Huarenchenque
diffinis
CH: Camarones
diffinis
PE: Samne
elsia
CH: Camarones
elsia
PE: Lima
elsia
PE: Paratype, nr Villa
elsia
EC: Is. Santa Cruz
elsia
galapagoensis
epimeral
stripes
constricted
or divided
EC: Paratype, Is. Santa Cruz
galapagoensis
BR: Allotype
brasiliensis
epimeral
stripes
constricted
or divided
BR: Rio Lajeado Grande
brasiliensis
135
BR: Paratype, Itatiaia
brasiliensis
T  E,  146, 2003
mesanepisternal
stripe
incomplete
US: Snow Peak
mesanepisternal
stripe
complete
US: Bishop
californica
AR: Punta Lara
californica
PE:Paralectotype, Apurimac
peralta
CH: Paratype, San Pedro de Atacama
tinti
confusa
BO: Belen
PE: Cuzco
peralta
peralta
CH: Holotype
tinti
CH: Paratype, Río Loa
tinti
EC: Cuenca
marchali
CH: Paratype, Calama
tinti
CH: Paratype, Chiu-chiu
tinti
stripes linear
or slightly
narrowed
CH: Pucatrihue-Bahía Mansa
variegata
CO: Lectotype
cornigera
VE: Altos de Tiara
GU: Tamahú
cornigera
cornigera
MX: Holotype
MX: Allotype
stripes with
deep
rounded
indentations
AR: San Carlos
AR: San Carlos
haarupi
haarupi
136
manni
manni
MX: Río Zaya
manni
mesepimeral stripe
narrowed at
midlength (broad
and shallow
indentation)
mesanepisternal
stripe complete
mesanepisternal
stripe incomplete
pauloi
VE: Cerro Yaví
VE: Cerro Yaví
nubigena
nubigena
stripes with
deep rounded
indentations
CO: Paralectotype cornigera
AR: Salta, NR 9
planaltica
AR: Calilegua NP
planaltica
MX: NE Huatusco
planaltica
psilus
stripes
wide
MX: La Galinda
vazquezae
vazquezae
stripes
divided into
spots
EC: Baños
vazquezae
condor
PE: Paratype, W Almirante
biliosa
stripes
complete
and sinuous
stripes
complete
and wide
biliosa
stripes
complete
and linear
demarmelsi
eduardoi
stripes with deep
indentations or
divided into spots
joannisi
joannisi
BO: Holotype
BR: Umuarama
joannisi
punctata
137
T  E,  146, 2003
VE: Auyantepui
a
b
c
VE: Auyantepui
VE: Allotype, Mt. Roraima
119
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, lateral view.
138
a
CH: Maipu
b
CH: Maipu
c
d
PE: Abancay
brevifrons
a
b
c
d
121
PE: Abancay
fissifrons
122
Figs. 121-122. Abdomen – a-b, male; c-d, female; a, c, dorsal view; b, d, lateral view.
139
T  E,  146, 2003
a
EC: Pallatanga
constriction of abdominal
segment III marked
b
c
EC: Pallatanga
d
a
PE: Cajamarca
segment III marked
intricata
123
AR: Río Colorado
segment III slight
b
c
PE: Cajamarca
AR: Río Colorado
segment III slight
d
AR: Río Colorado
pallipes
AR: Río Colorado
140
124
a
b
125
obscura
a
AR: San Lorenzo
b
AR: San Lorenzo
c
AR: Lesser
d
AR: Lesser
126
vigintipunctata
141
T  E,  146, 2003
a
US: Leslie Canyon
b
incomplete mediolongitudinal stripe
US: Leslie Canyon
c
US: Leslie Canyon
d
dugesi
a
US: Leslie Canyon
127
CR: Monteverde
b
CR: Monteverde
PD spots small
c
MX: Cuernavaca
PL spots absent
d
jalapensis
MX: Cuernavaca
142
128
a
US: Valentine
PD large and confluent with PL
b
PL spots present
US: Valentine
US: Del Puerto Canyon
c
d
multicolor
a
US: Scotia
b
US: Scotia
c
US: Lake Denmark
d
mutata
US: Lake Denmark
143
129
130
T  E,  146, 2003
AR: N Buta Ranquil
a
AR: Comi Có
b
c
AR: Comi Có
medio-longitudinal black stripe
d
absoluta
131
AR: Cafayate
a
b
AR: Comi Có
AR: Cafayate
medio-longitudinal pale stripe complete
c
AR: Colonia Dora
AR: Colonia Dora
d
bonariensis
144
132
a
CH: E Caramávida
AR: Trevelin
b
c
CH: Angol-Vegas Blancas
ML and PL confluent
d
MD and ML separated
diffinis
CH: Angol-Vegas Blancas
a
b
c
d
PE: Samne
elsia
a
b
133
134
PE: Samne
EC: Is. Santa Cruz
galapagoensis
EC: Is. Santa Cruz
145
135
T  E,  146, 2003
a
BR: Holotype
b
BR: Holotype
BR: Allotype
c
BR: Allotype
d
brasiliensis
a
US: Sonoma, Kruse Ranch Rd
b
136
US: Silverado
c
d
californica
US: Silverado
146
137
a
AR: Punta Indio
b
AR: Punta Indio
medio-longitudinal pale stripe
c
d
AR: Punta Lara
AR: Punta Lara
confusa
a
EC: Cuenca
b
EC: Cuenca
c
d
138
EC: Baeza
139
EC: Baeza
marchali
147
T  E,  146, 2003
a
PE: Paralectotype, Apurimac
b
c
ML and PL separated
d
peralta
a
MD and ML separated
140
CH: Holotype
b
CH: Holotype
c
CH: Allotype
ML and MD confluent
CH: Allotype
d
tinti
148
141
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
variegata
142
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.
149
T  E,  146, 2003
a
VE: Altos de Tiara
b
GU: Tamahú
PL very small to absent
c
VE: Altos de Tiara
d
bright
light
blue spot
CO: Lectotype
e
GU: Tamahú
f
EC: Napo
g
CR: San José
h
EC: Napo
i
cornigera
CR: San José
Figs. 143. Abdomen – a-e, male; f-i, female; a-b, f-g, dorsal view; c-e, h-i, lateral view.
150
143
a
AR: San Carlos
PL large
b
AR: San Carlos
c
d
AR: San Carlos
144
AR: San Carlos
haarupi
a
MX: Holotype
b
MX: Holotype
MX: Allotype
c
d
MX: Allotype
manni
151
145
T  E,  146, 2003
a
VE: Cerro Yaví
b
VE: Cerro Yaví
c
VE: Cerro Yaví
d
VE: Cerro Yaví
nubigena
a
b
c
d
pauloi
152
146
147
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.
psilus
153
149
T  E,  146, 2003
c
MX: Laguna Azul
d
MX: Laguna Azul
psilus
a
b
149
150
vazquezae
c
EC: Baños
d
EC: Baños
biliosa
154
151
a
b
EC: Santa Rosa
biliosa
c
d
EC: Santa Rosa
condor
151
152
a
AD & MD present in VII-VIII
pale spots present in X
153
b
AL & ML present in VII-VIII
demarmelsi
a
AD & MD absent in VII-VIII
pale spots absent in X
154
b
joannisi
AL & ML absent in VII-VIII
Figs. 151-154. Abdomen – a-b, male; a, dorsal view; b, lateral view.
155
T  E,  146, 2003
c
EC: Narupa
d
EC: Narupa
e
joannisi
BO: Holotype
154
a
b
eduardoi
a
BR: Umuarama
b
BR: Umuarama
c
BR: Macieiras
d
punctata
BR: Macieiras
Figs. 154-156. Abdomen – a-b, male; c-e, female; a, c, dorsal view; b, d-e, lateral view.
156
155
156
inner margin
ventral view of abdominal right half
a
seg. III
b
outer margin
seg. IV
VE: Auyantepui
seg. VIII
157
VE: Mt. Roraima
draco
seg. IX
a
EC: Baeza
b
brevicercia
a
b
c
a
b
PE: Lectotype maita
brevifrons
PE: Abancay
inner and outer carinae parallel
fissifrons
159
AR: Río Colorado
AR: Río Colorado
160
EC: Pallatanga
intricata
a
b
CH: Maipu
inner and outer carinae sinuous
a
b
158
EC: Baeza
PE: Cajamarca
161
obscura
BO: Allotype
Figs. 157-162. Ventral terga, ventral view – a, male; b-c, female.
157
162
T  E,  146, 2003
a
AR: Río Colorado
b
pallipes
a
AR: Río Colorado
163
AR: San Lorenzo
b
vigintipunctata
a
posterior portion of inner margin
convergent to outer margin in V-VII
b
dugesi
a
AR: Lesser
164
US: Leslie Canyon
US: Leslie Canyon
165
CR: Monteverde
b
jalapensis
MX: Cuernavaca
166
US: Valentine
a
posterior portion of inner margin
parallel to outer margin in V-VII
b
multicolor
a
167
US: Scotia
b
mutata
Figs. 163-168. Ventral terga, ventral view – a, male; b, female.
158
US: Lake Denmark
168
a
b
AR: Comi Có
anterior portion of outer
margin linear to concave in IV
maximum width at distal 0.60
absoluta
a
b
a
AR: Comi Có
169
AR: Cafayate
AR: Colonia Dora
bonariensis
170
AR: Trevelin
anterior portion of outer
margin convex in IV
AR: Shaman
b
diffinis
maximum width at basal 0.30
171
a
b
elsia
a
b
172
EC: Is. Santa Cruz
galapagoensis
a
b
CH: Arica
EC: Paralectotype, Is. Santa Cruz
173
BR: Holotype
brasiliensis
BR: Allotype
159
174
T  E,  146, 2003
a
b
californica
a
US: Silverado
175
AR: Punta Lara
b
confusa
a
AR: Punta Lara
176
EC: Cuenca
b
marchali
EC: Baeza
177
a
b
peralta
178
a
CH: Holotype
b
tinti
a
variegata
160
179
CH: Queulat NP
wide ventral terga
b
CH: Allotype
CH: Villa La Tapera
180
GU: Tamahú
a
c
VE: Altos de Tiara
narrow ventral terga
b
d
EC: Napo
cornigera
a
b
haarupi
manni
nubigena
MX: Allotype
183
VE: Cerro Yaví
184
pauloi
a
b
182
VE: Cerro Yaví
a
b
AR: San Carlos
MX: Holotype
a
b
181
AR: San Carlos
a
b
CR: San José
185
AR: Salta, NR 9
planaltica
BR: Itatiaia
Figs. 181-186. Ventral terga, ventral view – a, c, male; b, d, female.
161
186
T  E,  146, 2003
a
PR: El Verde st.
b
a
psilus
MX: Laguna Azul
187
vazquezae
188
a
EC: Santa Rosa
b
a
EC: Baños
condor
a
outer margin linear in VII
b
demarmelsi
a
joannisi
191
BO: Holotype
a
eduardoi
190
VE: Allotype
outer margin sinuous in VII
b
189
biliosa
192
193
a
BR: Umuarama
b
punctata
162
BR: Macieiras
194
black extended
over auricle
brevicercia
draco
fissifrons
CH: Paratype, Inacaliri
brevifrons
EC: Baeza
VE: Auyantepui
CH: Maipu
intricata
obscura
EC: Pallatanga
black not
extended over
auricle
pallipes
AR: Río Colorado
jalapensis
CR: Monteverde
vigintipunctata
dugesi
AR: San Lorenzo
US: Leslie Canyon
multicolor
mutata
US: Valentine
US: Scotia
Figs. 195-206. Male abdominal segments I-II, ventral view.
163
T  E,  146, 2003
absoluta
bonariensis
elsia
galapagoensis
AR: Comi-Có
AR: Cafayate
californica
EC: Is. Santa Cruz
confusa
US: Snow Peak
AR: Punta Lara
peralta
164
tinti
diffinis
AR: Trevelin
brasiliensis
BR: Paratype, Campos do Jordão
marchali
EC: Cuenca
variegata
CH: Villa La Tapera
cornigera
cornigera
VE: Altos de Tiara
manni
MX: Holotype
haarupi
GU: Tamahú
AR: San Carlos
nubigena
pauloi
VE: Cerro Yavi
planaltica
psilus
AR: Salta, NR 9
vazquezae
PR: El Verde St.
biliosa
EC: Santa Rosa
condor
165
T  E,  146, 2003
231
230
demarmelsi
joannisi
EC: Narupa
232
233
eduardoi
166
punctata
EC: Pallatanga
ventral tubercle
I
tubercle
low
draco
II
brevifrons
genital lobe
VE: Auyantepui
CH: Maipu
A
genital
lobe low
B
A < 0.33B
brevicercia
brevifrons
EC: Baeza
tubercle
high
PE: Lectotype maita
A
B
A < 0.50B
intricata
fissifrons
EC: Pallatanga
obscura
vigintipunctata
tubercle
high
B
A < 0.50B
pallipes
genital lobe
high and
subquadrate
AR: San Lorenzo
AR: Río Colorado
genital lobe
high and
triangular
dugesi
tubercle
A low
A
US: Leslie Canyon
B
multicolor
A < 0.75B
mutata
US: Merrit
jalapensis
MX: Cuernavaca
US: Scotia
absoluta
AR: Cafayate
numerous
denticles
bonariensis
diffinis
AR: Trevelin
few apical
denticles
elsia
CH: Arica
brasiliensis
galapagoensis
EC: Is. Santa Cruz
californica
US: Snow Peak
Figs. 234-253. Abdominal segments I-II, lateral view of tubercle I and genital lobe.
167
T  E,  146, 2003
254
AR: Punta Lara
256
PE: Cuzco
258
confusa
255
peralta
257
variegata
259
CH: Puerto Ramirez
tubercle
high
EC: Quito
tinti
haarupi
AR: San Carlos
A
260 A =B0.50B
263
cornigera
GU: Tamahú
anterior side
convex
cornigera
CO: Lectotype
261
VE: Altos de Tiara
262
manni
MX: Holotype
anterior side
linear
AR: Salta, NR 9
planaltica
265
anterior side
concave
A tubercle low
A < 0.33B
marchali
B
266
267
pauloi
psilus
268
269
PR: El Verde St.
270
nubigena
VE: Cerro Yaví
vazquezae
MX: Chilpancingo
tubercle
trapezoidal
small denticles over lateral surface of lobe
271
biliosa
EC: Santa Rosa
condor
A
A
A < 0.33B
B
C
tubercle high
and trapezoidal
273 EC: Ejoannisi
Cornejo Astorga
tubercle low
and rounded
C
A
A
D
D
C > 0.66D
B
A < 0.33B
272 VE:demarmelsi
Paratype, Tamá NP
A > 0.33B
274
eduardoi
B
C = 0.33D
275
Figs. 254-275. Abdominal segments I-II, lateral view of tubercle I and genital lobe.
168
264
CO: paralectotype cornigera
A > 0.33B
punctata
BR: Umuarama
B
276
277
medial
digitiform
projection
anterior
anterior
process
medial digitiform
projection
A
c
anterior
process
external
internal
posterior
B
draco
a
hamular
fold
b
b
brevicercia
EC: Baeza
280
278
279
c
c
a
b
a
a
intricata
EC: Pallatanga
brevifrons
CH: Maipu
brevifrons
281
PE: Lectotype maita
c
A
a
c
B
b
283
b
a
A>B
fissifrons
b
282
A
B
A>B
a
A>B
VE: Auyantepui
pallipes
AR: Río Colorado
284
285
c
c
a
obscura
a
vigintipunctata
AR: San Lorenzo
a
b
Figs. 276-285. Hamuli – a, posterior view; b, ventral view; c, latero-external view.
169
dugesi
US: Leslie Canyon
T  E,  146, 2003
286
287
b
b
a
a
jalapensis
CR: Monteverde
289
multicolor
a
absoluta
a
bonariensis
tip rounded
293
294
a
EC: Is. Santa Cruz
297
b
a
californica
projection
low
D
PE: Cuzco
marchali
EC: Cuenca
C
C = 0.33 D
a
variegata
AR: Cari Laufquen
Figs. 286-300. Hamuli – a, posterior view; b, ventral view.
170
a
C = 0.50D
300
b
b
b
peralta
A<B
b
AR: Punta Lara
299
C
D
B
confusa
US: Snow Peak
a
projection
prominent
A
b
brasiliensis
296
298
b
a galapagoensis
CH: Arica
a
diffinis
AR: Trevelin
b
elsia
295
mutata
US: Scotia
b
b
a
a
b
AR: Cafayate
tip pointed
b
291
b
292
B
A<B
US: Merrit
290
a
288
A
a
tinti
301
303
302
tip rounded
b
a
a
cornigera
VE: Altos de Tiara
304
a
cornigera
305
306
b
b
b
a
manni
MX: Holotype
a
nubigena
VE: Cerro Yaví
307
pauloi
308
tip carinated
309
b
b
a
a
planaltica
b
a
psilus
vazquezae
PR: El Verde St.
AR: Salta, NR 9
310
311
312
tip pointed
tip rounded
A
A
B
b
b
A<B
a
a
biliosa
EC: Santa Rosa
313
tip pointed
tip
recurved
haarupi
AR: San Carlos
GU: Tamahú
tip pointed
a
b
b
B
a
demarmelsi
314
b
A>B
joannisi
315
tip blunt
A
B
a
b
b
A>B
condor
a
eduardoi
b
a
punctata
BR: Umuarama
Figs. 301-315. Hamuli – a, posterior view; b, ventral view.
171
T  E,  146, 2003
dorsal
316
317
anterior
319
B
A
ventral
318
posterior
draco
brevicercia
VE: Auyantepui
320
fissifrons
intricata
dugesi
US: Leslie Canyon
328
mutata
tip curved
dorsally
332
dorsal
margin
concave
elsia
329
absoluta
AR: Cafayate
333
galapagoensis
CH: Arica
EC: Is. Santa Cruz
Figs. 316-335. Spine of anterior lamina, latero-inner view.
172
obscura
pallipes
AR: Río Colorado
326
327
B
A > 2B
AR: San Lorenzo
323
325
A
vigintipunctata
brevifrons
PE: Lectotype maita
322
EC: Pallatanga
324
brevifrons
CH: Maipu
321
US: Scotia
A < 2B
EC: Baeza
tip directed
ventrally
tip curved
dorsally
jalapensis
330
331
dorsal
margin
linear
bonariensis
diffinis
AR: Trevelin
334
brasiliensis
multicolor
US: Merrit
CR: Monteverde
tip curved
dorsally
335
ventral margin
linear to
concave
californica
US: Snow Peak
confusa
336
AR: Punta Lara
340
tinti
marchali
337
EC: Baeza
341
cornigera
CO: Lectotype
peralta
338
PE: Cuzco
342
ventral
margin
linear to
concave
variegata
339
CH: Puerto Ramirez
343
cornigera
GU: Tamahú
ventral
margin
convex
haarupi
AR: San Carlos
347
manni
MX: Holotype
spine
vestigial
344
nubigena
planaltica
345
VE: Cerro Yaví
AR: Salta, NR 9
346
pauloi
psilus
348
PR: El Verde St.
B > 2A
A
acute
pointed
tip
B
349
vazquezae
350
biliosa
EC: Santa Rosa
351
condor
352
joannisi
354
353
A
B
B < 1.5A
demarmelsi
demarmelsi
VE: Paratype, Las Copas
eduardoi
355 BR: Paratype,
Catarina
356
punctata
BR: São Paulo
Figs. 336-357. Spine of anterior lamina, latero-inner view.
173
357
punctata
BR: Umuarama
T  E,  146, 2003
a
VE: Auyantepui
a
EC: Holotype
lateral carina
concave
sub-basal
tooth low
lateral carina
linear
b
b
tip bearing spine
VE: Auyantepui
EC: Holotype
EC: Allotype
VE: Allotype, Mt. Roraima
358
c
draco
359
c
brevicercia
atrophied
cercus
a
PE: Lectotype maita
d
CH: Maipu
PE: Matucana
malfomed
cercus
b
sub-basal
tooth low
CH: Maipu
PE: Lectotype maita
tip not
mucronate
c
360
CH: Paralectotype
f
tip
mucronate
CH: Huasco Bajo
g
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.
174
e
PE: Yauyos
h
a
AR: Holotype
a
PE: Celendin
sub-basal tooth low
c
AR: Allotype
PE: Celendin
AR: Holotype
b
b
361
fissifrons
AR: Cafayate
d
a
EC: Pallatanga
c
sub-basal tooth
prominent
c
AR: Cafayate
AR: Cafayate
b
pallipes
362
PE: Celendin
363
intricata
a
a
BO: Holotype
sub-basal tooth
prominent
crest at distal 0.25
sub-basal tooth
prominent
BO: Holotype
b
c
364
cercus
widening
distally
obscura
tip
mucronate
BO: Allotype
b
crest at distal 0.33
inner and outer
margins parallel
AR: Tucumán
c
BO: Buena Vista
tip rounded
vigintipunctata
Figs. 359-365. Cerci – a-c, male; d, female; a, c, dorsal view; b -d, lateral view.
175
365
T  E,  146, 2003
366
367
a
a
US: Leslie Canyon
MX: Los Aposentos
b
b
crest triangular
process projected
posteriorly
c
c
small process
not projected
posteriorly
sub-basal
tooth low and blunt
only extreme tip
ventrally bent
tip rounded
d
d
US: Chiricahua Mountains
dugesi
jalapensis
MX: Cuernavaca
a
a
US: Merrit
b
entire tip
ventrally bent
crest high
b
B
A<B
US: Scotia
crest low
c
A
B
A>B
US: Merrit
US: Lake Denmark
d
A
c
margin linear
US: Alum Rock Park
tip pointed
e
margin
slightly
convex
multicolor
tip pointed
368
US: San Francisco
d
US: Lake Denmark
mutata
Figs. 366-369. Cerci – a-b, d, male; c, e, female; a, d-e, dorsal view; b, mediodorsal view; c, lateral view.
176
369
370
AR: N Buta Ranquil
a
AR: Punta Lara
a
AR: N Buta Ranquil
outer base with pale spot
crest as high
as base
b
b
AR: Punta Lara
tip pointed
sub-basal
tooth
prominent
tip pointed
c
AR: Río Aluminé
c
AR: Punta Lara
371
absoluta
bonariensis
372
CH: Lago Ranco
a
PE: nr Villa
a
CH: Lago Ranco
crest higher
than base
crest suddenly
rising
b
b
PE: nr Villa
tip pointed
c
c
AR: Lago Aluminé
tip rounded
diffinis
373
PE: Lima
elsia
375
a
BR: Holotype
a
EC: Lectotype
crest gradually
rising
inner margin diverging gradually
b
b
BR: Holotype
EC: Is. Santa Cruz
c
tip rounded
EC: Paralectotype
galapagoensis
374
c
BR: Allotype
brasiliensis
Figs. 370-375. Cerci – a-b, male; c, female; a, c, dorsal view; b, lateral view.
177
tip pointed
T  E,  146, 2003
377
AR: Punta Lara
376
US: Snow Peak
tip pointed
a
a
US: Snow Peak
AR: Punta Lara
b
b
tip pointed
sub-basal
tooth
vestigial
AR: Punta Lara
c
US: Silverado
confusa
c
californica
379
a
tip angled
marked concavity
slight concavity
a
PE: Cuzco
EC: Cuenca
BO: Lectotype
b
b
EC: Cuenca
EC: Cotundo
c
d
peralta
c
AR: Laguna Escondida
378
marchali
BO: Copacabana
a
tip angled
CH: Holotype
a
tip angled
heel at inner margin
CH: Holotype
b
AR: Laguna Escondida
b
tip rounded
sub-basal tooth
prominent
380
CH: Allotype
tinti
c
c
381
Figs. 376-381. Cerci – a-b, male; c-d, female; a, c-d dorsal view; b, lateral view.
178
variegata
AR: Cari Laufquen
tip directed
externally
distal 0.60 parallel sided
f
a
tip directed
posteriorly
CO: Lectotype
GU: Tamahú
inner margin concave
b
crest high and smoothly curved
A
no sub-basal tooth
c
g
B
CO: Lectotype
A<B
EC: Napo
GU: Tamahú
pointed tip
e
rounded tip
CR: San José
VE: Colonia Tovar
d
382
cornigera
inner and outer margins diverging
a
a
AR: San Carlos
MX: Holotype
b
A
lateral carina
concave
crest low, long and angled
B
AR: San Carlos
A>B
b
MX: Holotype
pointed tip
c
383
haarupi
AR: San Carlos
384
c
MX: Allotype
manni
Figs. 382-384. Cerci – a-b, f-g, male; c-e, female; a, c-f dorsal view; b, g, lateral view.
179
T  E,  146, 2003
385
386
tip directed
externally
a
a
VE: Cerro Yaví
AR: Salta, NR 9
crest high, long and
smoothly curved
b
A
B
A<B
ventrally bent
pointed apex
b
AR: Salta, NR 9
VE: Cerro Yaví
c
tip pointed
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
a
AR: Salta, NR 9
planaltica
tip rounded
with minute
medial spine
b
pauloi
Figs. 385-387. Cerci – a-b, male; c-e, female; a, c-e dorsal view; b, lateral view.
180
c
a
a
MX: Laguna Azul
b
MX: Laguna Azul
b
vazquezae
tip pointed
c
MX: Laguna Azul
psilus
a
crest angled
b
tip pointed
a
maximum width at medial 0.33
crest angled
b
c
BO: Holotype
condor
joannisi
181
T  E,  146, 2003
a
a
EC: Santa Rosa
crest smoothly convex
crest angled
b
tip blunt
EC: Santa Rosa
maximum width at distal 0.33
b
tip incised
c
VE: Allotype, Tamá NP (after De Marmels, 2001)
tip
mucronate
EC: Baños
demarmelsi
c
tips converging
or parallel
biliosa
a
BR: Umuarama
tips diverging
b
a
BR: Umuarama
crest with
denticles
tip
incised
c
b
eduardoi
182
BR: Macieiras
punctata
sptrg
RP1
RP2 evenly
curved
subtrg
MA
Cu-A space
RP3-4
PE: Abancay
brevifrons
trg
anal
trg
IRP2a
IRP2b
anal
loop
VE: Auyantepui
draco
IRP2 fork
symmetrical
Figs. 396-397. Right pair of male wings.
183
T  E,  146, 2003
bent of RP2
marked
US: Leslie Canyon
dugesi
two rows of
cells beginning
distal to
pterostigma
AR: Punta Lara
bonariensis
peralta
184
AR: Campo Alegre
planaltica
two rows of cells
beginning at
proximal end of
pterostigma
GU: Tamahú
cornigera
demarmelsi
185
T  E,  146, 2003
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
asymmetrical
anal angle
angled
186
bent of RP2
marked
GERMANY: Limsdorf
"Aeshna" isoceles
anal angle
rounded
long
parallel
fusion
US: Angostura
Aeshna palmata
short
parallel
fusion
ITALY: Chiusi
"Aeshna" mixta
US: Leslie Canyon
Rhionaeschna dugesi
fusion
at a single
point
VENEZUELA: Las Copas
Rhionaeschna demarmelsi
JAPAN: Kizyoka
GREECE: Kefallini
"Aeshna" isoceles
187
T  E,  146, 2003
A
A
CANADA: Edson
B
B
A= 0.50B
A= 3-4B
GUATEMALA: Tamahú
Aeshna sitchensis
genital lobe no higher than margin of fossa with
denticles restricted to a marginal row
I
JAPAN: Kizyoka
B
A= 8-9B
JAPAN: Kizyoka
Rhionaeschna cornigera
transverse ridge
a
A
transverse ridge
II
b
genital lobe low with
denticles on ventral surface
conical
tubercle
a
conical
tubercle
GUATEMALA: Tamahú
b
genital lobe high with denticles on
ventral and lateral surface
ventral terga
posterior
processes
cylindric
tubercle
a
BRAZIL: Catarina
Rhionaeschna eduardoi
a
BRAZIL: São Carlos
Castoraeschna januaria
genital lobe
overdeveloped
and projected
posteriorly
a
"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.
188
hamular
anterior
process long
and triangular
spine
triangular and
narrow
posteriorly
directed
b
hamular anterior
process short
hamular fold
not
differentiated
from wall of
fossa
b
a
US: Merriman
Aeshna constricta
spine
cylindrical
posteroventrally
directed
CANADA: Edson
a
US: Lincoln
a
Aeshna sitchensis
b
hamular
anterior
process absent
hamular posterior
process
tuberculate
b
hamular
posterior process
longer than
width of ventral
portion
a
subarctica
US: Rockaway Township
Aeshna clepsydra
Aeshna subarctica
long anteriorly
directed spines
crest
curved
verticalis
short dorsally
directed spines
crest
linear
denticles
on dorsal
crest
apical
spine of
cercus
inner
margin
distal
concavity
b
a
c
c
US: Smith Falls
Aeshna constricta
d
c
sub-basal carena
of seg. X
b
d
US: Anchorage
Aeshna eremita
Figs. 422-425. Hamuli – a, postero-ventral view; b, c, ventral view – 426-427. Male terminalia – a, latero-ventral view; b, lateral view; c, dorsal view; d, lateral view of epiproct.
189
T  E,  146, 2003
auricle broadly
triangular
USA: Merriman
GERMANY: Sassenpfühle
Aeshna constricta
Aeshna grandis
"Aeshna" rileyi
auricle narrow
parallel sided
ARGENTINA: Nahuel Huapi
GERMANY: Limsdorf
Rhionaeschna variegata
a
Ventral lobe lacking
medio-longitudinal fold
Basal fold of
lateral lobe
GUATEMALA: Tamahú
a
Ventral lobe lacking mediolongitudinal fold
Lateral lobe lacking
basal fold
b
FIJI
"Aeshna" isoceles
Basal fold of
lateral lobe
a
Medio-longitudinal fold of ventral lobe
b
GERMANY: Limsdorf
"Aeshna" isoceles
b
"Aeshna" rileyi
Figs. 428-433. Auricle, ventrolateral view – 434-436. Distal segment of vesica spermalis – a, lateral view; b, dorsal view.
190
Castoraeschna
Coryphaeschna
Remartinia
Oreaeschna
"Aeshna"
Anaciaeschna
"Aeshna" isoceles
Andaeshna
Anax
Hemianax
2 3
1
4
5
6
7 8
9 10 11 12
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).
191
T  E,  146, 2003
Strict consensus tree
of 472 MPT obtained
after successive
weighting
[CI 72, RI 95]
1(1)
3
1(0)
1(1)
4
1(1)
2
5
6 7
1(0)
3(1)
1
7(5)
1(0)
2(0)
11
1(0)
9
1(0)
12
4(3)
14
3(3)
1(0)
10
* = homoplasic changes
Node (1,2): 21.2*, 22.1
Node (2,3): 12.1, 16.3, 17.1*
Node (3,4): 35.1*
Node (4,5): 34.1
Node (5,6): 17.0*
Node (6,7): 2.1, 16.2*, 19.2*
Node (2,8): 18.1*, 30.1, 33.1, 35.3, 36.1,
38.1, 39.1*
Node (1,9): 8.1*, 14.1*
Node (9,10): 11.1*
Node (10,11): 9.1*
Node (11,12): 21.1*
Node (12,13): 39.2*
Node (12,14): 5.1, 15.1, 23.1, 28.1*
Node (14,15): 2.2, 10.1, 11.2, 20.3*, 39.1*
Node (14,16): 13.1, 14.2, 24.1
Node (16,17): 6.1*, 10.2*, 18.1*, 29.1*
Node (14,18): 35.1*
Node (18,19): 20.2, 25.1, 32.1*
Node (18,20): 27.1
Node (20,21): 1.1*, 3.1, 6.2, 10.2*, 16.1*,
19.1, 31.1*, 32.1*
Node (21,22): 4.1, 29.1*
Node (21,23): 20.1*
Node (20,24): 32.0*
Node (24,25): 21.2*, 39.2*
Node (25,26): 35.2*
Node (25,27): 20.1*
Node (24,28): 26.1, 31.2*, 39.1*
Node (28,29): 35.2*
Node (20,30): 9.1*, 31.2*, 32.1*
Node (30,31): 35.2*, 37.1
Node (31,32): 39.1*
Node (32,34): 39.2*
Node (30,34): 7.1
Node (34,35): 27.2
Fig. 438. Cladogram of Rhionaeschna species.
192
1(0)
16
4(0)
13
15
17
3(2) 19
8(3)
Character state changes
5(3)
8
2(1)
21
1(0)
18
2(0)
22
23
1(0) 26
25
1(0) 27
1(1)
20
1(0)
24
28
3(1)
1(0)
29
dictatrix
viridis
grandis
tuberculifera
juncea
verticalis
subarctica
clepsydra
eremita
interrupta
canadensis
osiliensis
nigroflava
crenata
septentrionalis
stichensis
caerulea
cyanea
constricta
walkeri
palmata
persephone
petalura
umbrosa
brevistyla
affinis
williamsoniana
ellioti
mixta
jaspidea
triangulifera
isoceles
andresi
rufipes
rileyi
subpupillata
30
2(1)
31
1(0)
32
1(0)
33
34
1(1)
1(1)
35
Aeshna
"Aeshna"
Anaciaeschna
"Aeshna"
Andaeshna
"Aeshna"
draco
brevifrons
fissifrons
pallipes
brevicercia
inticata
obscura
vigintipunctata
demarmelsi
biliosa
condor
joannisi
decessus
eduardoi
punctata
cornigera
planaltica
haarupi
nubigena
psilus
manni
vazquezae
brasiliensis
californica
confusa
marchali
peralta
tinti
variegata
3(0)
Oreaschna
dugesi
jalapensis
multicolor
mutata
absoluta
bonariensis
diffinis
elsia
galapagoensis
Rhionaeschna
7N
69 W
67 W
65 W
63 W
61 W
59 W
VENEZUELA
5N
COLOMBIA
3N
BRAZIL
1N
draco
100 km
80 W
70 W
60 W
50 W
40 W
PERU
10 S
BRAZIL
BOLIVIA
CHILE
PARAGUAY
20 S
ARGENTINA
URUGUAY
30 S
brevifrons
fissifrons
1000 km
Figs. 439-440. Rhionaeschna dot maps.
193
T  E,  146, 2003
71 W
70 W
60 W
50 W
40 W
PARAGUAY
CHILE
ARGENTINA
30 S
URUGUAY
35 S
pallipes
400 km
10 N
83 W
78 W
73 W
68 W
63 W
VENEZUELA
5N
COLOMBIA
0
BRAZIL
ECUADOR
brevicercia
intricata
5S
PERU
500 km
194
80 W
5S
70 W
60 W
500 km
BRAZIL
PERU
15 S
BOLIVIA
vigintipunctata
obscura
PARAGUAY
CHILE
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
195
PANAMA
T  E,  146, 2003
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
95 W
0
65 W
ECUADOR
1000 km
50 W
35 W
PERU
BRAZIL
15 S
BOLIVIA
PARAGUAY
CHILE
30 S
ARGENTINA
URUGUAY
45 S
absoluta
196
85 W
75 W
65 W
PERU
55 W
45 W
BOLIVIA
BRAZIL
PARAGUAY
CHILE
25 S
35 S
URUGUAY
ARGENTINA
bonariensis
diffinis
45 S
500 km
85 W
80 W
75 W
70 W
65 W
60 W
COLOMBIA
0
ECUADOR
5S
BRAZIL
10 S
PERU
15 S
elsia
galapagoensis
BOLIVIA
500 km
CHILE
197
T  E,  146, 2003
115W
125W
105 W
CANADA
45 N
USA
californica
35 N
500 km
MEXICO
70 W
20 S
BOLIVIA
60 W
50 W
BRAZIL
PARAGUAY
CHILE
30 S
URUGUAY
ARGENTINA
confusa
brasiliensis
40 S
500 km
198
90 W
80 W
70 W
60 W
VENEZUELA
10 N
COLOMBIA
ECUADOR
PERU
0
BRAZIL
10 S
BOLIVIA
marchali
tinti
CHILE
900 km
ARGENTINA
80 W
70 W
60 W
50 W
PARAGUAY
40 W
PERU
BOLIVIA
BRAZIL
20 S
PARAGUAY
CHILE
30 S
URUGUAY
ARGENTINA
40 S
peralta
variegata
50 S
1000 km
199
30 W
T  E,  146, 2003
70 W
85 W
VENEZUELA
55 W
35 W
COLOMBIA
ECUADOR
5S
PERU
BRAZIL
BOLIVIA
20 S
PARAGUAY
CHILE
planaltica
haarupi
pauloi
ARGENTINA
35 S
MEXICO
85 W
GUATEMALA
10 N
1000 km
URUGUAY
75 W
65 W
55 W
NICARAGUA
COSTA RICA
VENEZUELA
PANAMA
COLOMBIA
0
ECUADOR
PERU
cornigera
nubigena
BRAZIL
10 S
BOLIVIA
200
1000 km
105 W
USA
30 N
90 W
75 W
60 W
45 W
MEXICO
DOMINICAN REPUBLIC
CUBA
PUERTO RICO
JAMAICA
15 N
GUATEMALA
DOMINICA
NICARAGUA
COSTA RICA
VENEZUELA
PANAMA
COLOMBIA
0
ECUADOR
PERU
psilus
BRAZIL
15 S
BOLIVIA
PARAGUAY
CHILE
1000 km
ARGENTINA
113 W
108 W
103 W
98 W
USA
29 N
300 km
MEXICO
24 N
manni
vazquezae
19 N
201
T  E,  146, 2003
81 W
76 W
71 W
56 W
VENEZUELA
5N
COLOMBIA
0
ECUADOR
biliosa
condor
5S
BRAZIL
PERU
200 km
79 W
74 W
69 W
VENEZUELA
6N
COLOMBIA
1N
ECUADOR
200 km
202
demarmelsi
joannisi
43 W
48 W
53 W
24 S
PARAGUAY
BRAZIL
29 S
decessus
eduardoi
punctata
300 km
URUGUAY
140 W
50 N
110 W
80 W
50 W
20 W
20 N
10 S
40 S
Marmaraeschna group
Draco group
2000 km
Fig. 459. Rhionaeschna dot maps – 460. Rhionaeschna generalized tracks.
203
T  E,  146, 2003
90 W
50 N
60 W
30 W
20 N
10 S
40 S
Schizuraeschna group
2000 km
140 W
50 N
110 W
20 N
10 S
40 S
Neureclipa group
2000 km
Figs. 461-462. Rhionaeschna generalized tracks.
204
80 W
50 W
20 W
140 W
50 N
110 W
80 W
50 W
20 W
80 W
50 W
20 W
20 N
10 S
40 S
Variegata group
2000 km
140 W
50 N
110 W
20 N
10 S
40 S
Cornigera group
2000 km
Figs. 463-464. Rhionaeschna generalized tracks.
205
T  E,  146, 2003
140 W
50 N
110 W
80 W
50 W
20 W
80 W
50 W
20 W
20 N
10 S
40 S
Punctata
group
2000 km
140 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.
206
0.0
North America
Tepuis
Central America
North America
Tepuis
Central America
Paranense
Paranense
Andes
Andes
Austral South
America
Austral South
America
5.0
a
b
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 areas of Rhionaeschna species.
207

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