Garrison & Von Ellenrieder - 2024 - Taxonomic Revisions of Neotropical Zygoptera

Zootaxa 5405 (1): 001–042 ISSN 1175-5326 (print edition)
https://www.mapress.com/zt/
Copyright © 2024 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5405.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:6298292C-F06A-4DB7-BBE8-07D47AE4F9DF
Taxonomic revisions of some Neotropical Zygoptera (Odonata)

Rosser Garrison1,2 & Natalia von Ellenrieder1,3
1
Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento, CA 95832-1448,
U. S. A.
2
Research Associate
� argiavivida@gmail.com; https://orcid.org/0000-0001-8586-2723
3
Senior Insect Biosystematist
� natalia.ellenrieder@gmail.com; https://orcid.org/0000-0002-1159-2019
Table of contents
Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Resumen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Material and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Proposed taxonomic changes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Lestes debellardi De Marmels, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Lestes scalaris Gundlach, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Palaemnema brevignoni Machet, 1990. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Hetaerina duplex Selys, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Hetaerina longipes Hagen in Selys, 1853. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Mnesarete guttifera (Selys, 1873). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Acanthagrion obsoletum (Förster, 1914). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Ischnura sobrina Schmidt, 1943. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Leptagrion Selys, 1876. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Leptagrion macrurum (Burmeister, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Oxyagrion ablutum (Calvert, 1909). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Telebasis Selys, 1865. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Tepuibasis De Marmels, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Tepuibasis demarmelsi (Machado & Lencioni, 2011). New combination. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Possible synonyms pending further study. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Philogenia carrillica Calvert, 1907. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Leptagrion andromache Hagen in Selys, 1876. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Phoenicagrion flammeum (Selys, 1876). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Tuberculobasis croceum (Burmeister, 1839). New combination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Incertae sedis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Agrion rufovittatum Blanchard, 1846. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Abstract
We provide updated synonymies for various genera and species of Neotropical Zygoptera based on examination of
specimens and literature accompanied by justifications and supported by illustrations. The following generic synonyms
are proposed: Austrotepuibasis Machado & Lencioni, 2011 and Pseudotepuibasis Stand-Pérez & Pérez-Gutiérrez,
2020 are junior synonyms of Tepuibasis De Marmels, 2007; Fredyagrion Lencioni, 2022, Kiautagrion Lencioni, 2022,
Nathaliagrion Lencioni, 2022, and Machadagrion Lencioni, 2022 are junior synonyms of Leptagrion Selys, 1876;
Juenagrion Lencioni, 2023 is a junior synonym of Telebasis Selys, 1865. The following specific synonyms are proposed:
Lestes fernandoi Costa, Souza & Muzón, 2006 is a junior synonym of Lestes debellardi De Marmels, 1992; Lestes
tikalus Kormondy, 1959 is a junior synonym of Lestes scalaris Gundlach, 1888; Palaemnema brasiliensis Machado,
Accepted by M. Marinov: 14 Dec. 2023; published: 26 Jan. 2024

2009 is a junior synonym of Palaemnema brevignoni Machet, 1990; Hetaerina aurora Ris, 1918 is a junior synonym of
Hetaerina duplex Selys, 1853; Hetaerina hebe Selys, 1853 is a junior synonym of Hetaerina longipes Hagen in Selys,
1853; Mnesarete mariana Machado, 1996 is a junior synonym of Mnesarete guttifera (Selys, 1873); Acanthagrion hartei
Muzón & Lozano, 2005 is a junior synonym of Acanthagrion obsoletum (Förster, 1914); Oxyagrion bruchi Navás, 1924
is a junior synonym of Oxyagrion ablutum (Calvert, 1909); Austrotepuibasis alvarengai Machado & Lencioni, 2011
and Austrotepuibasis manolisi Machado & Lencioni, 2011 are junior synonyms of Tepuibasis demarmelsi (Machado &
Lencioni, 2011). Helveciagrion Machado, 1980 is again considered a junior synonym of Telebasis Selys, 1865; Leptagrion
auriceps St. Quentin, 1960 is again considered a junior synonym of Leptagrion macrurum (Burmeister, 1839), and
Homeoura sobrina (Schmidt, 1943) is transferred back to Ischnura Charpentier. The following synonymies are tentatively
proposed pending further examination of specimens: Philogenia lankesteri Calvert, 1924 is a possible junior synonym
of Philogenia carrillica Calvert, 1907; Leptagrion jeromei Lencioni, Vilela & Furieri in Vilela, Garcia Júnior, Furieri &
Lencioni, 2021 is a possible junior synonym of Leptagrion andromache Hagen in Selys, 1876; Phoenicagrion flavescens
Machado, 2010 is a possible junior synonym of Phoenicagrion flammeum (Selys, 1876); Tuberculobasis mammilaris
(Calvert, 1909) is a possible junior synonym of Tuberculobasis croceum (Burmeister, 1839) new combination. The status
of Agrion rufovittatum Blanchard, 1846 is briefly discussed.
Key words: Odonata, Zygoptera, damselfly, genera, species, synonymy, taxonomy, Neotropical
Resumen
Proveemos sinonimias actualizadas para varios géneros y especies de Zygoptera neotropicales sobre la base de la
examinación de ejemplares y literatura, acompañadas de justificaciones y soportadas por ilustraciones. Los siguientes
sinónimos genéricos son propuestos: Austrotepuibasis Machado & Lencioni, 2011 y Pseudotepuibasis Stand-Pérez &
Pérez-Gutiérrez, 2020 son sinónimos junior de Tepuibasis De Marmels, 2007; Fredyagrion Lencioni, 2022, Kiautagrion
Lencioni, 2022, Nathaliagrion Lencioni, 2022 y Machadagrion Lencioni, 2022 son sinónimos junior de Leptagrion
Selys, 1876; Juenagrion Lencioni, 2023 es un sinónimo junior de Telebasis Selys, 1865. Los siguientes sinónimos
específicos son propuestos: Lestes fernandoi Costa, Souza & Muzón, 2006 es un sinónimo junior de Lestes debellardi De
Marmels, 1992; Lestes tikalus Kormondy, 1959 es un sinónimo junior de Lestes scalaris Gundlach, 1888; Palaemnema
brasiliensis Machado, 2009 es un sinónimo junior de Palaemnema brevignoni Machet, 1990; Hetaerina aurora Ris, 1918
es un sinónimo junior de Hetaerina duplex Selys, 1853; Hetaerina hebe Selys, 1853 es un sinónimo junior de Hetaerina
longipes Hagen in Selys, 1853; Mnesarete mariana Machado, 1996 es un sinónimo junior de Mnesarete guttifera (Selys,
1873); Acanthagrion hartei Muzón & Lozano, 2005 es un sinónimo junior de Acanthagrion obsoletum (Förster, 1914);
Oxyagrion bruchi Navás, 1924 es un sinónimo junior de Oxyagrion ablutum (Calvert, 1909); Austrotepuibasis alvarengai
Machado & Lencioni, 2011 y Austrotepuibasis manolisi Machado & Lencioni, 2011 son sinónimos junior de Tepuibasis
demarmelsi (Machado & Lencioni, 2011). Helveciagrion Machado, 1980 es considerado nuevamente un sinónimo junior
de Telebasis Selys, 1865; Leptagrion auriceps St. Quentin, 1960 es considerado nuevamente un sinónimo junior de
Leptagrion macrurum (Burmeister, 1839), y Homeoura sobrina (Schmidt, 1943) es transferida de regreso a Ischnura
Charpentier. Las siguientes sinonimias son propuestas tentativamente pendiendo la examinación de más ejemplares:
Philogenia lankesteri Calvert, 1924 es un possible sinónimo junior de Philogenia carrillica Calvert, 1907; Leptagrion
jeromei Lencioni, Vilela & Furieri in Vilela, Garcia Júnior, Furieri & Lencioni, 2021 es un posible sinónimo junior de
Leptagrion andromache Hagen en Selys, 1876; Phoenicagrion flavescens Machado, 2010 es un posible sinónimo junior
de Phoenicagrion flammeum (Selys, 1876); Tuberculobasis mammilaris (Calvert, 1909) es un posible sinónimo junior
de Tuberculobasis croceum (Burmeister, 1839) nueva combinación. El estado de Agrion rufovittatum Blanchard, 1846 se
discute brevemente.
Palabras claves: Odonata, Zygoptera, libélula, géneros, especies, sinonimia, taxonomía, Neotropical
Introduction
Availability of additional specimens of some species described in the past based on just one or a few specimens
from a limited geographic area and a further understanding of the infraspecific variability to be expected in several
taxa have led us to the recognition of several specific synonyms of New World Zygoptera. We also suggest that
some new generic names were introduced without valid justifications and without any regard to recognition of the
female sex resulting in names to which specimens cannot be reliably adjudicated. We realize that generic concepts
are subjective divisions allowing for the inclusion of groups of species presumably sharing a common ancestor.
However, independently whether one favors a more inclusive or exclusive generic concept, we consider that a genus
· Zootaxa 5405 (1) © 2024 Magnolia Press GARRISON & VON ELLENRIEDER
should be clearly diagnosable and that the placement of species within it should be achieved without having to first
key out the specimens to species. Below, we provide suggested synonymies for these various names of New World
Zygoptera with our justification for doing so.
Material and methods
Taxa are treated by families, these arranged phylogenetically, and then alphabetically within families following the
classification outlined in Dijkstra et al. (2013). Nomenclature follows Garrison et al. (2010). Abbreviations used
throughout the text are as follows: Fw: forewing; Hw: hindwing; Pt: pterostigma; pq: postquadrangular cells; px:
postnodal crossveins; S1–10: abdominal segments 1 to 10. Illustrations were made with the aid of a camera lucida
coupled to a Zeiss SteREO discovery V20 stereomicroscope. Macrophotographs of dried specimens were edited
with Adobe Photoshop to match colors from photographs of live specimens when possible. Photographs, scans, and
illustrations are not to scale.
Specimens examined are listed under each respective taxon, and are deposited in the collection of Rosser W.
Garrison (RWG) unless otherwise mentioned as follows: ABMM: Angelo B. M. Machado, Centro de Coleções
Taxonômicas da Universidade Federal de Minas Gerais (CCT-UFMG), Belo Horizonte, Brazil; BMNH: British
Museum of Natural History, London, England; CSCA: California State Collection of Arthropods, Sacramento,
California; DZRJ: Departamento de Zoologia, Universidade Federal do Paraná, Curitiba; HZ: Haomiao Zhang
collection, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China; KPMNH: Kanagawa
Prefectural Museum of Natural History, Kanagawa, Japan; MNRJ: Museu Nacional, Universidade Federal do Rio
de Janeiro; MZLU: Lund University, Lund, Sweden; MZUSP: Museu de Zoologia da Universidade de São Paulo;
RBINS: Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium; RMNH: Naturalis Biodiversity
Center (formerly Rijksmuseum van Natuurlijke Historie), Leiden, The Netherlands; UG: University Guiana;
UMMZ: Museum of Zoology, University of Michigan, Ann Arbor, Michigan; USNM: United States National
Museum, Washington, D. C; UT: University of Texas, Austin, Texas. Approximate GPS data are shown in curly
brackets ({ }).
Proposed taxonomic changes
Lestes debellardi De Marmels, 1992
Lestes fernandoi Costa, Souza & Muzón, 2006. New synonymy
Several years ago, De Marmels (in litt.) pointed out to us that Lestes fernandoi was likely a junior synonym of L.
debellardi, and figures of both species taken from De Marmels (1992) (Figs. 1‒3) and Costa et al. (2006) (Figs. 4‒6)
bear this out, showing that appendage morphology and thoracic venter pattern are basically the same. The minor
differences between the two sets of figures in the size of the dark spot on thoracic venter and slightly different contour
of appendages fit within the usual intraspecific variability observed in species of this genus and slightly different
orientation of appendages. Costa et al. (2006) did not include L. debellardi in their key to Brazilian species since
it was described from Venezuela, nor did they compare L. fernandoi with that species. The types of L. fernandoi,
originally in the MNRJ, were lost in the 2008 fire that consumed that institution (Lencioni et al. 2021). De Marmels
(in litt.) recently opined that both names might be synonyms of the poorly known Lestes quadristriatus Calvert,
1909 described from four incomplete specimens from Chapada, Brazil. The specific name no doubt refers to the
four dark spots on the venter of its thorax (Fig. 7). We have not seen the male described by Calvert but reproduce
his three figures here (Figs. 7‒9). Lestes quadristriatus has appendages similar to those of L. debellardi (Figs. 1‒3)
and L. fernandoi (Figs. 4‒6) but differs in having four dark spots on the venter of the thorax (instead of two as in the
other two taxa) and in having the tip of the cercus "...bent downward so strongly as to form almost a right angle with
the superior margin, the extreme inferior tip being moderately acute [Fig. 8])" (tip of cercus not bent ventrad and tip
rounded in the other two taxa, Figs. 2, 5). Costa et al. (2006) in their key to the Brazilian species of Lestes separated
L. fernandoi from L. quadristriatus by the two (L. fernandoi) versus four (L. quadristriatus) thoracic markings on
the venter of the thorax. Based on this difference, we continue to rank L. debellardi as a distinct species but consider
L. fernandoi to represent a junior synonym of the latter.
TAXONOMIC REVISIONS OF NEOTROPICAL ZYGOPTERA Zootaxa 5405 (1) © 2024 Magnolia Press ·
FIGURES 1–11. Lestes names treated here. 1–3: ♂ L. debellardi; 4–6: ♂ L. fernandoi; 7–9: ♂ L. quadristriatus; 10: ♂ L.
scalaris; 11: ♂ L. tikalus. 1, 4, 7: pterothorax, ventral view; 2, 3, 5, 6, 8–11: ♂ caudal appendages; 2, 5, 8: lateral view; 3,
6: dorsal view; 9–11: mediodorsal view.
FIGURES 12–15. Lestes and Palaemnema names treated here. 12: ♂ L. tikalus; 13: ♂ L. scalaris; 14: ♂ P. brasiliensis;
15: ♂ P. brevignoni. 12, 13: pterothorax, lateral view; 14a, 15a: ♂ head, frontal view; 14b, 15b: ♂ thorax, lateral view.
Lestes scalaris Gundlach, 1888
Lestes tikalus Kormondy, 1959. New synonymy
Kormondy (1959) described Lestes tikalus from Guatemala and noted its similarity to L. scalaris described from
Cuba. We have been unable to find specific differences between a topotype male and description of the series of
males described from the type locality of L. tikalus and males from other Central American localities and matching
the description of L. scalaris from the Bahamas (San Salvador, pond 6.6 mi south of Cockburn Town, ca. 23.97°
N, 74.548° W; information from Dennis Paulson in litt.) and Jamaica. Kormondy compared his specimens with
Calvert's (1909) redescription and figure of L. scalaris and stated that they differed in the color of thorax ("thoracic
color and color pattern of tikalus is different from scalaris, but grossly similar to that in forficula"), shape of the
inner edge of cercus ("...similar inner edge except that the lamina in tikalus terminates in an acute spine much as
in L. mediorufus"), and relative length of paraproct to cercus ("The inferior appendages in tikalus are as long as
the superiors, whereas in scalaris they are noticeably shorter."). The thoracic pattern in our series of L. scalaris is
variable (Figs. 12, 13); that of our Jamaican male matches that of our topotype (Guatemala) and illustration of the
holotype given by Kormondy (Fig. 12). Other males (Fig. 13) have a narrower metallic mesepisternal stripe and
only a hint of a dark mesepimeral stripe (Belize), or a metallic mesepisternal stripe occupying the medial 2/3 of the
sclerite with the entire mesepimeron metallic (Costa Rica, Puerto Viejo). The alleged thoracic pattern differences
noted by Kormondy cannot be used as a specific differential. The paraprocts in our series of males including a
topotype (e. g., Fig. 11) show overlap in their length with our Jamaican male (Fig. 10), and the acute basal spine
also shows no significant differences beyond individual variability. We agree with Dunkle (1991) in treating Central
American populations as L. scalaris and consider L. tikalus a junior synonym of the latter, a conclusion that was
also arrived at by Dennis Paulson (in litt.) based on his comparison of a male of L. scalaris from the Bahamas with
his series of males from Central America.
Material Examined: JAMAICA, Trelawny Parish: 1 ♂, Windsor Caves upper pond {18.3555°N, 77.647°W},
102 m, 8 April 1962, Thomas W. Donnelly leg.;—BELIZE, Cayo District: 2 ♂♂, pond & sedge marsh 4.9 miles south
of Western highway on Chiquibul Road, 17.1323°N, 88.9605°W, 170 m, 30 May 1993, Sidney W. Dunkle leg.;—
HONDURAS, Olancho Department: 1 ♂, pond 7 km east of San Francisco de Becerra, 14.6542°N, 86.0528°W,
354 m, 18 December 1987, Sidney W. Dunkle leg.;—GUATEMALA, El Peten Department: 1 ♂, Tikal; 69x036,
17.225°N, 89.6133°W, 194 m, 27 August 1969, Thomas W. Donnelly leg.;—COSTA RICA, Heredia Province:
2 ♂♂, 1 ♀, Puerto Viejo, La Selva Biological Station; L01-439, 10.4667°N, 84.0167°W, 60 m, 26 August 1988,
Carlos Esquivel-Herrera leg.; 1 ♀, same data but 19 September 1966, Dennis R. & Mary Lynn Paulson leg.; 1 ♂,
same data but 23 September 1966.
Palaemnema brevignoni Machet, 1990
Palaemnema brasiliensis Machado, 2009. New synonymy
Palaemnema brasiliensis was described from the Serra do Navio, Amapá State, Brazil (Machado 2009) based on
the holotype male and two paratype males. From the original description, the species appeared to be close to, if not
synonymous, with P. brevignoni known from Venezuela (De Marmels 1990) and French Guiana (Machet 1990) and
later recorded also from Guyana (von Ellenrieder et al. 2017). We received a paratype of P. brasiliensis from Dr.
Machado several years ago and compared this male with males of P. brevignoni in our collection from Venezuela,
Guyana, and French Guiana. Machado (2009) provided a table listing eight diagnostic characters from P. brevignoni,
based on the original description and figures, to separate both taxa (characters for P. brevignoni in parentheses)
as follows: anteclypeus dark brown (bluish yellow), pronotum hind lobe reddish brown (light brown), tooth of
dorsal margin of cercus in lateral view absent (hardly visible), cercus in relation to paraprocts only slightly longer
(much longer), ventral margin of distal half of cercus straight (rounded), tip of penis (recte paraproct) pin-shaped
(subrectangular) and S10 black with ventrolateral pale spot (dorsally bluish, laterally brown). We have not observed
any of these differences between the paratype of P. brasiliensis and the specimens of P. brevignoni. The anteclypeus
in both is pale (almost white, Figs. 14a, 15a), and thoracic pattern (Figs. 14b, 15b) and caudal appendages (Figs.
16a, 17a), including the tips of the paraprocts in
FIGURES 16–17. Palaemnema names treated here. 16: ♂ P. brasiliensis; 17: ♂ P. brevignoni. 16a, 17a: ♂ caudal
appendages, mediodorsal view; 16b, 17b: ♂ paraproct tip, posterior view; 17c: ♂ habitus, lateral view.
posterior view (Figs. 16b, 17b), are essentially the same. Our paratype of P. brasiliensis is missing the genital ligula,
but Machado’s drawing of this structure matches that of P. brevignoni. Accordingly, we consider P. brasiliensis a
junior synonym of P. brevignoni. We also include a scan of P. brevignoni from Venezuela (Fig. 17c) showing its
characteristic color pattern.
Material Examined: VENEZUELA, Bolivar State: 1 ♂, small streams vic. Piedra de la Virgen, 14‒17 km S of
Kilometer 88 (highway), 87x050 {6.0867°N, 61.3994°W}, 1000 m, 30 July‒1 August 1987, Thomas W. Donnelly
leg.;—FRENCH GUIANA, Saint-Georges Arrondissement: 1 ♂, Approuague-Kaw, Kaw Mountain, malaise trap,
FRG MF3, 4.5506°N, 52.1944°W, 104 m, 23 January‒7 February 2007, N. Jönsson leg.;—2 ♂♂, same data but
4‒12 February 2007, N. Jönsson leg.;—GUYANA, Potaro-Siparuni Region: 2 ♂♂, Camp 1 to Chenepau, creek
[very teneral], 5.01°N, 59.6364°W, 470 m, 18 March 2014, N. von Ellenrieder & N. Johnson; 7 ♂♂ [teneral], same
data [CSCA];—BRAZIL, Amapá State: 1 ♂ (paratype of P. brasiliensis), Serra do Navio {0.9833°N, 52.05°W},
86 m, January 1957, Angelo Barbosa Monteiro Machado.
Hetaerina duplex Selys, 1853
Hetaerina aurora Ris, 1918. New synonymy
Garrison (1990) separated H. aurora from H. duplex primarily by the presence (H. aurora) or absence (H. duplex)
of red wingtips. He was unable to distinguish between females of these two and he (Garrison 1990: 210) discussed
variation in H. duplex noting that "It is possible that H. aurora is a subspecies of H. duplex." A more recent series of
specimens of H. aurora collected from western Ecuador (Pinchincha Province, Mindo, streams near town, 0°3'20"
S, 78°46'17" W, 28 January 2009) shows wingtip coloration to be variable, even within a single specimen. Three
males have both fore and hind wingtips red (as in Fig. 18c) while two others have hyaline wing tips (as in Fig. 18a).
One other male has three red wingtips while the right fore wingtip is hyaline (Fig. 18b). The caudal appendages
are the same in all and match those both for H. aurora and H. duplex, suggesting the synonymy of both names. We
consider H. duplex to be variable as to wing markings. We illustrate one of the clear-winged morphs of H. duplex
from Colombia (Fig. 19) that was referenced in Garrison (1990).
Material examined: COLOMBIA, Antioquia Department: 1 ♂, 1 ♀, 12 km NW of Medellin, road to San
Pedro, 6.41°N, 75.611°W, 2554 m, 13 February 1984, Carol M. & Oliver S. Flint, Jr. leg.; 1 ♂, tributary to Quebrada
Potreros, W of La Fe (small development at Fizebad, southeast of Medellin on road to La Ceja)—in forest, 6.09°N,
75.5067°W, 2000 m, 26 February 1984, Carol M. & Oliver S. Flint, Jr. leg.; Cauca Department: 1 ♂, Monte Redondo
{3.2168°N, 76.2002°W}, 1600 m, 23 February 1957; Meta Department: 3 ♂♂, 2 ♀♀, stream W of Villavicencio
{4.1419°N, 73.6604°W}, 600 m, 5 September 1980, Sidney W. Dunkle leg.;—ECUADOR, El Oro Province: 2♂♂,
2 ♀♀, Moromoro {3.6815°S, 79.7424°W}, 1500 m, 14–21 July 1941, David B. Laddey leg.; 1 ♀, Piñas {3.6802°S,
79.6857°W}, 1000 m, 13 July 1941, David B. Laddey leg.; 5 ♀♀, Portobello {3.7151°S, 79.6138°W}, 600 m,
1 August 1941, David B. Laddey leg.; Loja Province: 2 ♀♀, Malacatos, 15 mi S, 3 mi W of Loja {4.2333°S,
79.2833°W}, 1500 m, 23–25August 1941, David B. Laddey leg.; Napo Province: 1 ♀, Cotundo, {0.85°S, 77.8°W},
730 m, 5 September 1980; Pichincha Province: 8 ♂♂, Mindo, swamps and streams near town, 0.0556°S, 78.7714°W,
1244 m, 28 January 2009, R.W. Garrison & N. von Ellenrieder leg.; Santo Domingo de los Tsáchilas Province: 1 ♂,
3.2 km on road to Chiriboga from Alluriquin turnoff {0.3°S, 78.9667°W}, 780 m, 10 December 1989, Herman G.
Real leg.; 2 ♂♂, 2 ♀♀, Tandapi, {0.4141°S, 78.7982°W}, 1450 m, 15‒20 June 1965; 2 ♂♂, Tinalandia, 12 km east
of Santo Domingo de los Colorados, streams south of river, 0.2839°S, 79.05°W, 760 m, 5 October 1988, Sidney W.
Dunkle leg.; Zamora-Chinchipe Province: 2 ♂♂, Zamora, 4.8667°S, 79.15°W, 1100 m, 15–24 October 1941, David
B. Laddey leg.
Hetaerina longipes Hagen in Selys, 1853
Hetaerina hebe Selys, 1853. New synonymy
A similar condition as stated above for H. duplex exists for H. hebe (fore and hind wingtips red) versus H. longipes
(only hind wingtips red); Garrison (1990: 215) suggested a possible synonymy of these two names ("...the overall
morphology of the superior appendage allies it with the variable H. longipes, and it is possible that they represent
the same species.").
FIGURES 18–19. Wing color variability in Hetaerina duplex. 18: ♂♂ from Ecuador; 19 ♂ from Colombia.
Material examined: BRAZIL, Bahia State: 2 ♂♂, Reserva Serra Bonita; Fazenda Paris, Riberao Braço do
Sul, 15.4228°S, 39.5406°W, 190 m, 2 January 2013, Carol M. & Oliver S. Flint, Jr. leg.; 2 ♂♂, same data but 26
December 2012; Espirito Santo State: 1 ♂, Pôrto Catherina do Santa Leopoldina {20.1017°S, 40.5294°W}, 80 m;
Minas Gerais State: 1 ♂, Serra do Cipo, Santana do Riacho {19.5°S, 43.75°W}, 1300 m, 10 March 1956, Angelo
B. M. Machado leg.; 1 ♂, same data but November 1982; Pará State: 4 ♂♂, 4 ♀♀, Jacareacanga {6.2667°S,
57.65°W}, 88 m, October–December 1969, F. R. Barbosa leg.; Rio de Janeiro State: 4 ♂♂, 3 ♀♀, Cachoeira de
Macacu, Santana de Japuiba, Rio Sao Juao, 22.5681°S, 42.6567°W, 8–10 December 2000, Rosser W. Garrison
leg.; 1 ♂, 3 ♀♀, Guanabara, Represa do Rio Grande, 22.9167°S, 43.4167°W, 81 m, January 1972, F. M. Oliveira
leg.; 1 ♂, Parque Nacional do Itatiaia, Cachoeira Véu da Noiva, 22.4228°S, 44.6178°W, 1150 m, 27 November
2000, Rosser W. Garrison leg.; 1 ♂, 1 ♀, Rio de Janeiro vicinity of Alta Boa Vista on Mt. Tijuca {22.9628°S,
43.2619°W}, 500 m, 12 September 1922, Jesse H. Williamson leg.; 1 ♂, Rio Pirai {22.4°S, 47.55°W}, 608 m, 8
April 1977, Carol M. & Oliver S. Flint, Jr. leg.; 1 ♂, Vista Chinesa, Reserva Florestal Estadual Vista, Alto Boa
Vista, 22.8668°S, 43.25°W, 370 m, 12 December 2000, Rosser W. Garrison leg.; Rio Grande do Sul State: 1 ♂,
Porto Alegre {30.0346°S, 51.2176°W}, 15 m, 3 February 1938; 1 ♂, 1 ♀, San Francisco de Paula, Inst. Nacional
do Piedras {50.5578°S, 50.5578°W}, 650 m, 7 March 1959, O. A. Pereira leg.; Santa Catarina State: 8 ♂♂, 12 ♀♀,
Nova Teutonia {27.1623°S, 52.4172°W}, 400 m, May 1967–March 1980, Fritz Plaumann leg.; São Paulo State: 1
♂, Campos do Jordao, 22.6911°S, 45.4865°W, 1452 m, 7 March 1998, F. A. A. Lencioni leg.; 1 ♂, Horto Florestal
Augusto Ruschi, Sao Jose dos Campos {23.1833°S, 45.8833°W}, 645 m, 16 June 1996, F.A.A. Lencioni leg.; 1 ♂,
Nova Friburgo, 22.2667°S, 42.5332°W, 913 m, P. Sandig leg.;—PARAGUAY, Amambay Department: 1 ♀, Arroyo
Guavira, route 5, 10 km W of Cerro Cora, 22.6168°S, 55.9833°W, 266 m, 28–29 November 1973, Oliver S. Flint,
Jr.;—ARGENTINA, Misiones Province: 1 ♂, Arroyo Mbocay, Puerto Iguazú; 97x039, 25.6246°S, 54.5865°W, 250
m, 22 January 1997, Thomas W. Donnelly leg.; 1 ♂, Dos de Mayo, 27.0333°S, 64.65°W, 521 m, 1 January 1980,
R. Foerster leg.; 3 ♂♂, Iguazú {25.5667°S, 54.5667°W}, 161 m, January 1979–December 1980, R. Foerster leg.; 1
♂, Parque Nacional Iguazú, Arroyo Yacuy {25.63°S, 54.5886°W}, 160 m, 5 January 1979, Gerhard Jurzitza leg.; 1
♂, stream 6 km E of Eldorado, on route 17, 26.4°S, 54.6333°W, 228 m, 22 November 1973, Oliver S. Flint, Jr. leg.;
2 ♂♂, Eldorado {26.4°S, 54.6333°W}, 228 m, April 1991; 4 ♂♂, Puerto Rico {26.81°S, 55.0253°W}, 181 m, 4–8
April 1971, Carol M. & Oliver S. Flint, Jr. leg.
Mnesarete guttifera (Selys, 1873)
Mnesarete mariana Machado, 1996. New synonymy
Garrison (2006) suggested a possible synonymy between M. mariana and M. guttifera based on examination of a
topotype of M. mariana with several variable examples of M. guttifera, and he illustrated the variability of paraproct
shape that showed a gradation from a blunt to a pointed paraproct. We have been unable to unambiguously assign
all males of M. guttifera to either this species or M. mariana based on the only suggested diagnostic character (tip
of paraproct), nor have we been able to distinguish between females of the two taxa. Accordingly, we consider M.
mariana a junior synonym of M. guttifera.
Material examined: Listed in Garrison (2006).
Acanthagrion obsoletum (Förster, 1914)
Acanthagrion hartei Muzón & Lozano, 2005. New synonymy
We have been unable to find any significant differences between two paratypes of A. hartei and the original description
of this taxon and specimens of A. obsoletum in the RWG collection. We presented our evidence to Federico Lozano
who responded (in litt. August 2011) that he was also unable to find any differences in the male genital ligula but
that there were differences in the male appendages: the cerci of A. hartei being longer with the inner margin straight
whereas in A. obsoletum they are shorter with the inner margin forming an obtuse angle and S10 being slightly wider
than S9 in A. hartei. These differences were based on his examination of two males of A. obsoletum from Peru and
Ecuador. Upon sending photos of the terminalia of the specimens in the RWG collection (Figs. 22–25), Lozano agreed
10 · Zootaxa 5405 (1) © 2024 Magnolia Press GARRISON & VON ELLENRIEDER
FIGURES 20–25. Acanthagrion names treated here, ♂ caudal appendages. 20, 22, 23: A. hartei; 21, 24, 25: A. obsoletum;
20–25 upper row: posterior view; 20–25 lower row: lateral view.
TAXONOMIC REVISIONS OF NEOTROPICAL ZYGOPTERA Zootaxa 5405 (1) © 2024 Magnolia Press · 11
FIGURES 26–30. Ischnura and Leptagrion names treated here. 26–28: I. sobrina; 29: L. aculeatum; 30: L. acutum. 26,
29, 30: ♂ habitus; 27: ♀ habitus; 28: genital ligula, lateral view (right) and ectal view (left).
that they looked much the same and were intermediate between his examples of both taxa. Lozano suggested the
possibility that our two paratype males of A. hartei might really be A. obsoletum. We have found no significant
differences between our specimen of A. obsoletum from Madre de Dios (from where A. obsoletum was described)
in Peru and the illustrations of the holotype of A. hartei (Fig. 20); cercus length and inner margin of cercus are
indistinguishable (Fig. 25), and there is no character or combination of characters that would allow recognition of
two species among our extensive series of specimens of A. obsoletum. The minor differences observed between the
illustrations of the holotype of A. hartei (from Muzón & Lozano 2005; Fig. 20) and of A. obsoletum (Fig. 21) are
bridged by paratypes of A. hartei and other specimens of A. obsoletum, eliminating any apparent distinction. We
interpret this as intraspecific variability, which is to be expected in a species distributed over a large geographic area
(Southern Peru through Ecuador to Colombia; Leonard 1977).
Material examined: COLOMBIA, Meta Department: 2 ♂♂, road to Puerto Ariari {3.7837°N, 73.8516°W},
580 m, 6 February 2010, Melissa Sanchez leg.;—ECUADOR, Napo Province: 2 ♂♂, Río Arajuno, Dayuma Camp,
1.1°S, 77.5832°W, 434 m, 22–25 September 1990, Jean Belle leg.; 2 ♂♂, Río Sindy on Jatun Sacha Road, 5.3
km east of Puerto Napo, 1.05°S, 77.7833°W, 522 m, 15 June 1995, Sidney W. Dunkle leg. [ABMM]; 2 ♂♂, Río
Sindy 2, small muddy stream, 1.0467°S, 77.7422°W, 427 m, 21 January 2020, Rosser W. Garrison & Natalia von
Ellenrieder leg.; 15 ♂♂, pond and small vegetated tributary stream to Río Lupi at El Establo de Tomás Lodge,
0.9763°S, 77.8588°W, 559 m, 17 January 2020, Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA,
RWG];—ECUADOR, Napo Province: 4 ♂♂, 2 ♀♀, small rocky rivulet and associated pond 3.5 km SW of Puerto
Napo, 1.0592°S, 77.8158°W, 454 m, 22 January 2020, Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA,
RWG]; Orellana Province: 2 ♂♂, Scientific Research Station, Yasuni National Park, palm swamp just S of Bogi
Este River, S of km 42 on Yasuni Road, 0.6333°S, 76.5°W, 12 June 1995, Sidney W. Dunkle leg.; Pastaza Province:
1 ♂, Puyo, 1.4666°S, 77.9833°W, 1000 m, 21 April 2008, T. & K. Miyashita leg. [KPMNH]; Sucumbios Province:
2 ♂♂ (paratypes of A. hartei), stream east edge of Lago Agrio, 0.1°N, 76.8667°W, 396 m, 18 August 1980, Sidney
W. Dunkle leg.; Zamora-Chinchipe Province: 2 ♂♂, stream crossing road just before village of Guadaloupe #2302,
3.9114°S, 78.8575°W, 972 m, 20–21 August 2006, John C. Abbott leg. [UT];—PERU, Madre de Dios Region:
1 ♂, Manu, Erika Hostel (nr. Salvacion) along water supply by chicken coop, 12.8264°S, 71.3888°W, 550 m, 4
September 1989, Nancy E. Adams leg.; 13 ♂♂, Hotel Amazonia, across from Atalaya, exposed pond, 12.8703°S,
71.376°W, 414 m, 27 June 1993, Rosser W. Garrison leg. [RWG, USNM]; 16 ♂♂, Pakitza Reserved Zone, approx
4.5 km E of Pakitza, Aguajal, 11.93°S, 71.255°W, 250 m, 5 July 1993, Rosser W. Garrison leg. [BMNH, CSCA,
RWG]; 1 ♂, Pakitza, Trocha 1 to Aguajal, forest 5 km east JAL88-015, 11.93°S, 71.255°W, 250 m, 20 September
1988, Jerry A. Louton leg.; 2 ♂♂, Pakitza, Aguajal, approx. 4.5 trail km east, JAL88-018, 11.93°S, 71.255°W, 250
m, 22 September 1988, Jerry A. Louton leg.
Ischnura sobrina Schmidt, 1943
Homeoura sobrina: von Ellenrieder 2008b (transfer to Homeoura Kennedy).
von Ellenrieder (2008b) transferred I. sobrina to the genus Homeoura based on cercal morphology and Schmidt’s
note of its similarity to 'Ischnura' nepos (currently Homeura nepos). The types are apparently lost and Schmidt did
not describe the genital ligula beyond stating that it was "….durchaus Ischnura-artig ("….of the Ischnura-type."),
nor structural characters of the prothorax. The recent rediscovery of this species by José Martin Meléndez Quinto
in Peru and subsequent examination of specimens (Figs. 26–28) collected by Drs. Holger Hunger and Franz-Josef
Schiel showed that the genital ligula lacks the diagnostic triangular ental processes of Homeoura and that the
original placement of this species in the genus Ischnura was correct.
Material Examined: PERU, Pasco Región: 3 ♀♀, pond about 6.3 km NE of Grapanazu, 10.4327°S,
75.4569°W, 2390 m, 10 December 2015, Holger Hunger & Franz-Josef Schiel leg. [RWG]; 4 ♂♂, 7 ♀♀, 2.1 km NE
of Restaurante Turístico Pozucino along Camino Vecinal San Alberto, 10.5481°S, 75.3736°W, 2156 m, 9 December
2015, Holger Hunger & Franz-Josef Schiel leg. [CSCA, RWG].
Leptagrion Selys, 1876
Fredyagrion Lencioni, 2022. New synonymy

Kiautagrion Lencioni, 2022. New synonymy
Nathaliagrion Lencioni, 2022. New synonymy
Machadagrion Lencioni, 2022. New synonymy
Lencioni (2022) erected four new genera, Fredyagrion (type species: L. lencioninetoi Lencioni, 2022; including
L. aculeatum Santos, L. andromache Hagen in Selys, L. alfonsoi Machado, L. capixabae Santos, L. dispar Selys,
L. elongatum Selys, L. itabaiana Vilela, Lencioni & Santos, L. jeromei Lencioni, Vilela & Furieri, in Vilela,
Garcia Júnior, Furieri & Lencioni, L. siqueirai Santos, and L. vriesianum Santos; Kiautiagrion (type species: L.
acutum Santos, 1961); Nathalagrion (type species: L. porrectum Selys, 1876; including L. perlongum Calvert), and
Machadagrion (type species: L. cyanostigma Machado, 2012; including L. dardanoi Santos and L. garbei Santos).
All four genera were diagnosed in a table using only four characters: the posterior surface of the head (pale or black)
and three characters of the genital ligula. No consideration, except for the first character, was given to females even
though Garrison et al. (2010) diagnosed and keyed both sexes of Leptagrion and Furieri et al. (2020) presented a
key to eight of the known females. We have examined males of L. aculeatum (Figs. 29, 42), L. acutum (Figs. 30, 43,
55), L. andromache (Figs. 31, 44, 56), L. bocainense (Figs. 32, 45), L. dardanoi (paratype, Figs. 33, 46), L. dispar
(Figs. 34, 47), L. elongatum (Figs. 35, 48), L. garbei (Figs. 36, 49), L. itabaiana (paratype, Figs. 37, 50), L. jeromei
(Figs. 38, 57), L. macrurum (Figs. 39, 51, 52), L. perlongum (Figs. 40, 53) and L. porrectum (Figs. 41, 54) and have
failed to find any characters of the genital ligula or any other characters that would justify erecting new genera.
We believe that the color of the head and the secondary sexual characters of the male genital ligula proposed by
Lencioni (2022) to define his genera reflect only species-specific differences that are known to occur within many
other coenagrionid genera. We also believe that generic definitions should also include diagnostic characters for
the female sex, since otherwise it will be impossible to assign an unknown female to one specific genus. For these
reasons we consider these newly introduced scientific names to be junior synonyms of Leptagrion.
We offer illustrations of the genital ligula and full body scans of the 13 species referenced above. The illustrations
of the genital ligulae in Lencioni (2022) are diagrammatic and we have attempted to illustrate details of the ligula
in entolateral view for some of the species we have. Our illustrations of the genital ligula for our single specimen of
L. acutum (Fig. 43) determined by A. B. M. Machado differ significantly from that illustrated by Lencioni (2006,
figs. 81A‒D; 2017, figs. 209A‒E, 2022, figs. E 4, 69). The caudal appendages of our male (Fig. 55) are close to,
if not identical, to those illustrated by Lencioni (2006: figs. 81A, B; 2017: figs. 209A, B; 2022: figs. E1, 2) but our
male has two medially directed teeth followed ventrally by a pointed lobe (not a tooth) and we suspect that the same
condition exists for the specimen illustrated by Lencioni (2017, 2022). In our opinion, this difference also further
questions the validity of Kiautagrion since the genital ligula of our male significantly differs from that illustrated by
Lencioni on which his generic definition was based.
Leptagrion macrurum (Burmeister, 1839)
Leptagrion auriceps St. Quentin, 1960.—De Marmels & Garrison 2005 (synonymy);—Lencioni 2022 (removed from
synonymy).
After comparing the holotype of L. auriceps with males of L. macrurum, De Marmels & Garrison (2005) proposed
the synonymy of both names since the latter differed from typical examples of L. macrurum only by the pale
coloration of the head, thorax, and first four abdominal segments (fig. 23 in De Marmels & Garrison 2005), and the
appendages and genital ligula were found to be identical (figs. 21, 22 in De Marmels & Garrison 2005) to those of
L. macrurum. Lencioni (2022) removed L auriceps from synonymy with the older name L. macrurum based on the
following: "The shape of the junction between the posterior lobe of the prothorax and the middle lobe (angle of the
junction and the shape of the lateral border of the posterior lobe) proved to be an important and reliable structure to
be used together with the morphology of the cerci and penis and, in L. auriceps it was fundamental to separate this
species from its congener L. macrurum, as shown in Figures 9 [recte 8] and 11C [recte 10] (holotype of L. auriceps
and lectotype of L. macrurum, respectively) the junction is very different, confirming the position of L. auriceps as a
valid species." The differences figured by Lencioni are small and are suggestive of individual variation rather than a
FIGURES 31–34. Leptagrion names treated here, ♂ habitus. 31: L. andromache; 32: L. bocainense; 33: L. dardanoi; 34:
L. dispar.
FIGURES 35–38. Leptagrion names treated here, ♂ habitus. 35: L. elongatum; 36: L. garbei; 37: L. itabaiana; 38: L.
jeromei.
FIGURES 39–41. Leptagrion names treated here, ♂ habitus. 39: L. macrurum; 40: L. perlongum; 41: L. porrectum.
FIGURES 42–45. Leptagrion names treated here, genital ligula: lateral view (right column), entolateral view (central
column), ectal view (left column). 42: L. aculeatum; 43: L. acutum; 44: L. andromache; 45: L. bocainense.
column), ectal view (left column). 46: L. dardanoi; 47: L. dispar; 48: L. elongatum; 49: L. garbei; 50: L. itabaiana.
column), ectal view (left column). 51: L. macrurum; 52: L. auriceps; 53: L. perlongum; 54: L. porrectum.
FIGURES 55–57. Leptagrion names treated here. 55: ♂ L. acutum, caudal appendages, mediodorsal view (right), lateral
view (center), dorsal view (left); 56: ♂ L. andromache; 57: ♂ L. jeromei; a, b: posterior lobe of prothorax, a, lateral view,
b, strict lateral view; c, d: caudal appendages, c, lateral view, d, dorsal view.
specific difference (see additional comments concerning L. andromache and L. jeromei under possible synonymies
below). The morphology of the male caudal appendages and genital ligula are essentially identical. Though De
Marmels & Garrison (2005) failed to flex the genital ligula of the holotype of L. auriceps (Fig. 52), the overall shape
of the genital ligula in the two (Figs. 51, 52) is alike, with the medio-longitudinal laminar process being probably
also present in the holotype of L. auriceps (as in Fig. 51, entolateral view) suggesting that these two names are
synonymous. Accordingly, we still consider L. auriceps a junior subjective synonym of L. macrurum.
Material Examined: Leptagrion aculeatum: SURINAME, Paramaribo District: 1 ♂, Ma Retraite, RMNH.
INS 514697, 5.8667°N, 55.1333°W, 1 m, 30 April 1948, D. Geijskes leg.;—FRENCH GUIANA, Cayenne: 1 ♂, at
mercury vapor light, about 8 km S of N2 at PK82, 4.8733°N, 52.3364°W, 19 February 1988, Rosser W. Garrison
leg.; Leptagrion acutum: BRAZIL, Bahia State: 1 ♂, Amargosa {13.0167°S, 39.6°W}, 359 m, October 1988,
Olaf Mielke leg.; Leptagrion andromache: BRAZIL, Santa Catarina State: 1 ♂, Nova Teutonia, {27.162305°S,
52.417214°W}, 400 m, 28 February 1968, Fritz Plaumann leg.; Leptagrion bocainense: BRAZIL, Rio de Janeiro
State: 5 ♂♂, Serra da Bocaina, Sao Jose do Barreto {22.3333°S, 41.75°W}, 2 m, November 1968, Moachir
Alvarenga leg.; Leptagrion dardanoi: BRAZIL, Pernambuco State: 1 ♂ paratype, Recife, Parque Zoo-Botanico,
Dois Irmaos {8.8333°S, 35.95°W}, 463 m, 18 November 1965, Newton Dias dos Santos leg.; Leptagrion dispar:
BRAZIL, Espirito Santo State: 2 ♂♂, Conceicao da Barra, 18.5833°S, 39.75°W, 6 m, 18‒23 March 1968, Paulo
Elias leg.; Leptagrion elongatum: BRAZIL, Rio de Janeiro State: 3 ♂♂, Ilha de Marambaia, Praia da Armacao (by
boat), 23.0425°S, 43.9517°W, 4 m, 3 December 2000, J. Costa leg.; Leptagrion garbei: BRAZIL, Sergipe State:
1 ♂, Areia Branca, Serra de Itabaiana National Park, C2248, 10.7517°S, 37.3416°W, 179 m, 3 February 2021, J.
C. Santos leg.; Leptagrion itabaiana: BRAZIL, Sergipe State: 1 ♂ paratype, Areia Branca, Serra de Itabaiana
National Park; C2260, 10.7517°S, 37.3416°W, 179 m, 3 February 2021, J. C. Santos leg.; Leptagrion jeromei:
BRAZIL: Minas Gerais State: 1 ♂, Rio Caraça, near Santa Barbara, 20.0228°S, 43.4792°W, 728 m, 11 January
1987, Angelo Barbosa Monteiro Machado leg.; Leptagrion macrurum: BRAZIL, São Paulo State: 1 ♂, Parque
Estadual da Ilha do Cardoso, 25°06'2.67S, 47°55'52.81W, S PEIC-05 Malaise, trilha na mata ombrofila, ao lado do
Rio Pereque bromelias, 25.1008°S, 47.9313°W, 40 m, 20 October 2011, Angelo P. Pinto leg.;—Santa Catarina State:
1 ♂, Rio Vermelho {26.2833°S, 49.3333°W}, 566 m, December 1944‒January 1945, A. Maller leg.;—1 ♂, Joinville
{26.3°S, 48.8333°W}, 123 m, 28 February 1979, Miers leg.; Leptagrion perlongum: BRAZIL, Espirito Santo
State: 3 ♂♂, Castelo {20.6128°S, 41.2185°W}, 190 m, October 1979, Inacio leg.;—São Paulo State: 3 ♂♂, Parque
Estadual da Serra do Mar, Núcleo Caraguatatuba, Caraguatatuba {23.5894°S, 45.42°W}, 86 m, 15 January 2000,
F. A. A. Lencioni leg.; Leptagrion porrectum: BRAZIL, Espirito Santo State: 1 ♂, Coceiçao da Barra: Parque de
Itaunas (Trilha almescar) [compared to type in RBINS], 18.4289°S, 39.7097°W, 20 February 2005, Karina Schmidt
Furieri, R. Kawada, R. R. Silva leg.
Oxyagrion ablutum (Calvert, 1909)
Oxyagrion bruchi Navás, 1924. New synonymy
Oxyagrion bruchi was described by Navás (1924) based on a single male specimen collected in Alta Gracia, Córdoba
province, Argentina. The species was not recorded again, the type was reported to be lost (Costa 1978), and its
taxonomic status was deemed doubtful (Muzón & von Ellenrieder 1998). In her revision of the genus Oxyagrion
Selys, 1876, Costa (1978) suggested that specimens described by Ris (1918) from Bolivia (Coroico and Río Zongo)
as Oxyagrion terminale 'form b', were in fact O. bruchi. von Ellenrieder & Garrison (2006) redescribed O. bruchi
based on a series of specimens reared from larvae from Salta province in Argentina, plus some additional specimens
found in collections from Tucumán and Córdoba provinces, which conformed in every way to Navás' (1924)
original description and to the description of O. terminale 'form b' by Ris (1918). At that time, O. ablutum was still
placed under the genus Acanthagrion Selys, 1876 and the distinction between Acanthagrion and Oxyagrion Selys,
1876 based on color pattern (overall pale body coloration red and lacking postocular spots in Oxyagrion; overall
pale body coloration blue and with well-defined postocular spots in Acanthagrion [Leonard 1977; Costa 1978; De
Marmels 1984]), and we (von Ellenrieder & Garrison 2006) compared specimens identified as O. bruchi only with
other specimens of Oxyagrion, but not with Acanthagrion (now Oxyagrion) ablutum. After that publication, all
our subsequent attempts to find O. bruchi failed. von Ellenrieder & Lozano (2008) redefined these two genera and
transferred A. ablutum to Oxyagrion. Our better understanding of species in this group led us later to reexamine
these specimens and compare them with O. ablutum, which was present at the localities in which O. bruchi had
been previously collected. Specimens identified as both species share the morphology of male caudal appendages,
displaying the same degree of variability (Figs. 58–61), of female mesepisternal plates, carinae and fossae (Figs.
64, 65), and their genital ligulae are identical (Figs. 62, 63). We believe that the specimens identified as O. bruchi
simply represent juveniles of O. ablutum in which the blue or black areas of thorax and abdomen are red and the
postocular spots are not yet defined due to the entire color of the head being still very pale (Figs. 66–69), although
they are faintly hinted in some specimens (e. g., Fig. 67). Aside from the pale colors being reddish instead of blue,
the color pattern is identical in the females, but in the males the black on S2–6 ranges from being restricted to the
posterior ends of the segments (as illustrated for the neotype of O. bruchi, fig. 23 in von Ellenrieder & Garrison
2006) to halfway along them (Fig. 67) rather than extending along the entire length of the segments (Fig. 66). Such
an ontological color change is known in other 'blue' coenagrionids, in which dark areas can be reddish-brown in
tenerals (e. g., Acanthagrion cuyabae Calvert, A. gracile (Rambur), A. hildegarda Gloger, A. lancea Selys and A.
longispinosum Leonard; von Ellenrieder & Lozano 2008), or go from red to black and blue as specimens mature (e.
g., Mesamphiagrion laterale (Selys), M. nataliae Bota-Sierra & Wolf, 2013, M. occultum (Ris), and M. tamaense
(De Marmels); De Marmels 1988; von Ellenrieder & Garrison 2008; Bota-Sierra & Wolf 2013). Navás (1924) did
not state whether his male of O. bruchi was a juvenile or not, although his mention of S8 'strongly compressed'
would seem to indicate so, but Ris (1918) did state so for his O. terminale 'form b' ('Male [teneral]… Female [almost
adult]…'). The larval description of O. ablutum (as Acanthagrion; Pessacq et al. 2005) also agrees overall (shape
of head and caudal lamellae, setation of prementum and labial palp, shape and number of mandibular teeth) with
the description of the larva of O. bruchi (von Ellenrieder & Garrison 2006), although there appears to be some
variability in the shape of the larval cerci. The distribution range of O. bruchi (Bolivia and Jujuy, Salta, Tucumán,
and Córdoba provinces in Argentina) also falls within that known for O. ablutum (Lozano et al. 2020). We can find
no consistent differences between specimens of O. ablutum and O. bruchi other than the color differences mentioned
above, which we attribute to ontogenetic changes between teneral specimens and older adults, and consequently
consider O. bruchi a junior synonym of O. ablutum.
Material examined (includes teneral specimens previously identified as O. bruchi): BOLIVIA, La Paz
Department: 1 ♂, Caranavi, 15.8333°S, 67.5667°W, 660 m, 14 January 2011, Haruki Karube leg.; 1 ♂, Coroico,
South, 16.1981°S, 67.6944°W, 1400 m, 18 January 2011, T. & K. Miyashita leg.; Santa Cruz Department: 1 ♂,
Samaipata, Mama Pasquale, 3.9 km east on Hwy 4 to turnoff, Río el Fuerte, 18.1692°S, 63.8189°W, 1630 m, 5
November 1998, William F. Mauffray leg.; 1 ♀, Río Achira, 8.6 km E of Samaipata, 18.1519°S, 63.8025°W, 1400
m, 25 November 1999, Kenneth J. Tennessen leg.; 1 ♂, Río El Fuerte, E of Samaipata, 18.15°S, 63.8667°W, 1799
m, 5 November 1998, Kenneth J. Tennessen leg.;—ARGENTINA, Catamarca Province: 1 ♂, 1 ♀, Río Potrero, ca.
10 mi. north of Andalgalá, {27.5089°S, 66.3306°W}, 1375 m, 26 January 1974, Mary Lynn Paulson leg.; Córdoba
Province: 1 ♂, Tanti {31.3333°S, 64.6°W}, 1055 m, Bulla & Grosso leg.; Jujuy Province: 1 ♀, Stream on Hwy
9, yunga, pond (emerged in lab II-A-16), 24.4903°S, 65.3061°W, 1300 m, 3 April 1997, Natalia von Ellenrieder
leg. [CSCA]; 2 ♂♂, 2 ♀♀, Reserva El Pantanoso, Río El Pantanoso (wide, stony), 23.5214°S, 64.5969°W, 609 m,
3–4 November 2005, Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA, RWG]; 1 ♂, 1 ♀, Tilcara, laguna
Cerro Chico, 23.5644°S, 65.3836°W, 2432 m, 26 October 2006, Rosser W. Garrison & Natalia von Ellenrieder
leg.; 2 ♂♂, Arroyo Tres Cruces, Parque Nacional Calilegua, 23.7833°S, 64.7833°W, 1130 m, 21–22 March 2006,
Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA, RWG]; 1 ♂, same data but 8 September 2006, Natalia
von Ellenrieder leg. [CSCA]; 4 ♂♂, 1 ♀, same data but 15 January 1997, Javier Muzón & Natalia von Ellenrieder
leg. [CSCA, RWG]; 6 ♂♂, 1 ♀, La Angostura, vegetated stream [collected under permit ref. 119-10884/05, res.
379/07], 24.4503°S, 65.3789°W, 1511 m, 8 January 2006, 5 April 2007, 12 August 2005, Natalia von Ellenrieder
leg. [BMNH, CSCA]; 3 ♂♂, 1 ♀, Río Yuto, 23.51°S, 64.5406°W, 1200 m, 20 March 2006 & 16 May 2006, Natalia
von Ellenrieder leg. [CSCA]; 8 ♂♂, 2 ♀♀, Calilegua, laguna, 23.7833°S, 64.7833°W, 675 m, 21–22 March 2006,
Rosser W. Garrison & Natalia von Ellenrieder leg.; xx??, same data but 8 September 2006, Natalia von Ellenrieder
leg. [CSCA]; 2 ♂♂, Arroyo Zanjón Seco, 23.6864°S, 64.5711°W, 430 m, 16 May 2006, Natalia von Ellenrieder leg.
[CSCA]; Salta Province: 2 ♂♂, Angastaco, Río Angastaco at Rt. 40, 25.6789°S, 66.1586°W, 1877 m, 31 October
2005, Natalia von Ellenrieder & Rosser W. Garrison leg.; 6 ♂♂, Arroyo Brealito on nat. route 40, ca. 1 km north
of Seclantas, 25.315°S, 66.2494°W, 2116 m, 23 April 2009 & 28 December 2006 & 11 March 2010, Natalia von
Ellenrieder & Rosser W. Garrison leg. [CSCA, RWG]; 3 ♂♂, Arroyo sobre ruta a Yacimiento Macueta (PAE) 5.5
km northwest of Piquirenda Viejo, 22.3478°S, 63.8053°W, 582 m, 7 November 2006, Rosser W. Garrison & Natalia
von Ellenrieder leg. [RWG, CSCA]; 1 ♂, Cachi, Río Calchaquí, 25.1°S, 66.1833°W, 2250 m, 29 December 2006,
FIGURES 58–61. Oxyagrion names treated here, ♂ caudal appendages. 58, 60: O. bruchi; 59, 61: O. ablutum. 58, 59:
mediodorsal view; 60, 61: lateral view.
FIGURES 62–69. Oxyagrion names treated here. 62, 64, 67, 69: O. bruchi; 63, 65, 66, 68: O. ablutum. 62, 63: genital
ligula, ectal view (upper row), lateral view (lower row); 64, 65: ♀ mesostigmal plates, dorsal view (upper row), lateral
view (lower row); 66–69: habitus, 66, 67: ♂; 68, 69: ♀.
Natalia von Ellenrieder leg. [CSCA]; 3 ♂♂, 3 ♀♀, Cachi Adentro, acequia cerca de Hostal Samay Huasi, 25.0989°S,
66.2086°W, 2511 m, 28 December 2005, Natalia von Ellenrieder leg. [CSCA]; 2 ♂♂, Cafayate, Quebrada del
Río Colorado, 26.09°S, 66.0067°W, 1750 m, 10–28 March 2006, Rosser W. Garrison & Natalia von Ellenrieder
leg.; 1 ♂, Quebrada de Cafayate, Hwy 68; 97x012, 25.9333°S, 65.7214°W, 1500 m, 10 January 1997, Thomas
W. Donnelly leg. [FSCA]; 7 ♂♂, 2 ♀♀, Chicoana, Quebrada de Tilian, Laguna, 25.1308°S, 65.54°W, 1350 m, 20
April 2006 & 26 January 2012, Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA, RWG]; 2 ♂♂, Campo
Quijano {24.9167°S, 65.65°W}, 1664 m, 6 January 1989, Gerhard Jurzitza leg.; 2 ♂♂, Dique Itiyuro, ponds with
macrophytes below main dam, 22.1014°S, 63.7344°W, 552 m, 27 October 2005, Rosser W. Garrison & Natalia
von Ellenrieder leg. [CSCA, RWG]; 3 ♂♂, 1 ♀, El Gallinato, Arroyo El Gallinato, 24.6794°S, 65.3408°W, 1256
m, 8 March 2006, Natalia von Ellenrieder leg. [CSCA]; 1 ♂, Lagunita de las Catas, km 23 sobre ruta de Piquirenda
Viejo a Yacimiento Macueta, 22.2792°S, 63.8436°W, 843 m, 6 November 2006, Rosser W. Garrison & Natalia
von Ellenrieder leg.; 36 ♂♂, 9 ♀♀, Lesser, stream, 24.6825°S, 65.4775°W, 1312 m, 24 April 2001–5 December
2006 Natalia von Ellenrieder & Rosser W. Garrison leg. [CSCA, RBINS, RWG], some emerged in lab from 19
October–28 November 2005; 10 ♂♂, 2 ♀♀, areas within National Park El Rey, 24.7311°S, 64.6367°W, 861 m, 26–
27 April & 30 October 2006, Natalia von Ellenrieder & Rosser W. Garrison leg. [CSCA, RWG]; 1 ♂, Pool 10 km
southeast of Isla de Cañas, 22.9091°S, 64.6131°W, 775 m, 17 May 2006, Natalia von Ellenrieder leg. [CSCA]; 16
♂♂, 3 ♀♀, unnamed stream 15 km SE of Isla de Cañas, 22.9256°S, 64.5758°W, 706 m, 17 May 2006–8 December
2008, F. Lozano & Natalia von Ellenrieder leg. [CSCA]; 20 ♂♂, unnamed stream 20 km SE of Isla de Cañas,
22.9583°S, 64.5556°W, 661 m, 9 September 2006–1 May 2009, Rosser W. Garrison & Natalia von Ellenrieder leg.
[RWG, CSCA]; 3 ♂♂, 2 ♀♀, Pulares, lagunas en criadero de truchas, 25.0908°S, 65.6139°W, 1468 m, 6 December
2006 & 1 February 2006, Federico Lozano & Natalia von Ellenrieder leg. [CSCA]; 3 ♂♂,1 ♀, Río Anta Muerta,
23.1167°S, 64.498°W, 496 m, 18 May 2006, 22 May 2008, Natalia von Ellenrieder leg. [CSCA]; 13 ♂♂, 6 ♀♀,
Río Caldera, about 5 km south of Campo Alegre Dam, side stream with macrophytes, 24.5947°S, 65.3694°W,
1419 m, 11 November 2005–20 January 2012, Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA, RWG];
1 ♀, stream at km 5.5 on road from Piquirenda Viejo to Yacimiento Macueta (PAE), 22.3478°S, 63.8053°W, 582
m, 27 March 2007, Natalia von Ellenrieder leg. [CSCA]; 1 ♂, 1 ♀, San Lucas, small dam in stream, 25.8664°S,
66.0354°W, 2038 m, 14 April 2001, Natalia von Ellenrieder leg. [CSCA]; 1 ♂, San Antonio, 25.9953°S, 66.0318°W,
1649 m, 27 March 2006, Rosser W. Garrison & Natalia von Ellenrieder leg. [CSCA]; 1 ♂, San Lucas, small dam in
stream, 25.8664°S, 66.0354°W, 2038 m, 14 April 2001, Natalia von Ellenrieder leg.
Telebasis Selys, 1865
Helveciagrion Machado, 1980.—Garrison 2009 (synonymy);—Lencioni 2023 (removed from synonymy).

Juenagrion Lencioni, 2023. New synonymy
Garrison (2009) provided a comprehensive and illustrated review of Telebasis, listed diagnostic characters used to
define the genus, acknowledging that no one character can be considered unique, included a key for the reception
of the genus within New World Coenagrionidae with an angulate frons, and compared Telebasis with its probable
Old World sister genus Ceriagrion Selys. He considered Helveciagrion a junior synonym of Telebasis and provided
reasons for doing so. Finally, he presented three keys (two for males and one for females) for the separation of all
species known at that time.
Lencioni (2023) apparently lifted two couplets from Garrison's Telebasis synopsis: Key M-1: "Cerci
approximate….or nearly so…" and Key F-1:"Mesepisternal pits lateral to mid-dorsal carina present…., distance
between bases of cerci ≤ 0.20 basal width of each cercus", which he provided as his only reason for resurrecting
Helveciagrion from synonymy. We do not consider these two characters to merit generic status. One species of
Argia Rambur (A. lilacina Selys), one species of Enallagma Charpentier (E. clausum Morse), and at least three
species of Ceriagrion Selys: C. annulatum Fraser, C. obfuscans Dijkstra, Mézière & Kipping, and C. tenellum (de
Villers), also have approximate cerci and corresponding pinched mesothoracic pits in the female (except for the
female of C. tenellum), yet all these species remain in their respective genera and are accepted by all odonatologists.
Telebasis carminita Calvert and T. dominicana Selys in Sagra have nearly approximate male appendages showing
an intermediate character state between the Helveciagrion (sensu Lencioni) group of species and the remaining
Telebasis species. The species that Lencioni placed in Helveciagrion share genital ligula morphology with species
that he left in Telebasis; e. g., three species (T. boomsmae Garrison, T. collopistes Calvert, T. garrisoni Bick &
Bick) that Lencioni transferred to Helveciagrion share a common genital ligula morphology (Key Ma-4: apical
segment of ligula trowel-shaped, its lateral margin gently expanding to mid-length and, in some species, terminating
in a posteriorly directed lobe; with or without accessory lateral lobe) with eight non-Helveciagrion species (T.
dominicana (Selys in Sagra), T. vulnerata (Hagen), T. livida Kennedy, T. byersi Westfall, T. incolumis Williamson
& Williamson, T. salva (Hagen), and T. abuna Bick & Bick), and four other species he transferred to Helveciagrion
(T. obsoleta (Selys), T. simulacrum (Calvert), T. vulcanoae (Machado) and T. williamsoni Garrison), key out to Key
Ma-5 (apical segment of ligula clubbed, its lateral margin often expanded in lateral view but abruptly narrowing
at apical 0.25) together with 15 non-Helveciagrion species. Lencioni lists Garrison (2009) and Haber (2020) as a
means for identifying all members of Helveciagrion, although all eight species are treated in these papers within
Telebasis. Lencioni's advice for using Garrison's synopsis as a means of identification would mean identifying a
target species first to species before being able to place it into one of his genera.
The larvae of two of the species Lencioni placed in Helveciagrion, T. boomsmae and T. obsoleta, have been
described (Garrison 1994; Lozano et al. 2012) and their characters and illustrations show no generic differences
with known larvae of other species of Telebasis (García-Díaz 1938; Geijskes 1941, 1943; Westfall 1957; Novelo
Gutiérrez & Gómez Anaya 2005; Westfall & May 2006).
In their preliminary molecular phylogeny of Coenagrionoidea, Willink et al. (2023) tentatively concluded that
Telebasis is likely paraphyletic with Minagrion Santos nesting within it but acknowledged that further work is
required to fully resolve the affinities of all taxa attributed to Minagrion. Their analysis included several species of
Telebasis and three of Helveciagrion, and showed Helveciagrion as polyphyletic, with T. collopistes, T. obsoleta and
H. simulacrum nesting in two separate clades together with Telebasis species (Willink et al. 2023: figs. S7, S19),
thus further questioning the value of recognizing Helveciagrion. The same results are shown by an unpublished
phylogenetic tree based on COI sequence data that Tim Faasen (in litt., Fig. 70) kindly shared with us.
Lencioni (2023) did not treat the remaining 15 species currently placed in Telebasis (including T. byersi
Westfall and T. incolumnis Williamson & Williamson, both of which are closely related to the type species T. salva)
presumably saving these for a Part 2 of his publication. Regardless of where he might intend to taxonomically
place the remaining 15 species, further partitioning Telebasis s. l. would not resolve the para- and polyphylies
demonstrated here.
Based on all these data, we continue to follow Garrison (2009) in considering Helveciagrion a junior synonym
of Telebasis.
Lencioni (2023) also established a new genus Juenagrion (type species J. nanae Lencioni, 2023 by original
designation) to receive three previously described species of Telebasis (T. igapocola Faasen, T. leptocyclia Garrison,
and T. selaopyge De Marmels), based on two secondary sexual characters: "Prothorax of female with a pair of short
process on dorsum of posterior lobe directed caudad" and "[genital ligula] with tip of S3 sharp–angled arrow–shaped
(T. igapocola and T. nanae) or slightly rounded with a rim (T. leptocyclia and T. selaopyge), ectal region of S3 with
a pair of lateral folds". The female of T. leptocyclia is however still unknown. All four of these species fit within the
broad definition of Telebasis although the male of T. leptocyclia is aberrant by its different caudal appendages and
there is no convincing proof that it would cluster with T. igapocola and T. selaopyge. Again, Lencioni cited Garrison
(2009) as a means for identifying all members of Juenagrion although all three species described previously are
treated within Telebasis, which as mentioned above, would mean identifying a target specimen first to species
before placing it into a genus.
Garrison previously examined and illustrated both sexes of both T. igapocola and T. selaopyge and recognized
them as sister species (unpublished, as T. igapocola was described after Garrison's 2009 synopsis (Faasen 2017);
the male appendages and female prothorax are depicted here, Figs. 71, 72), but he found no justification to place
these two (or three) species in a separate genus based on these species-specific characters. The female of T. corbeti
Garrison (fig. 6d in Garrison 2009) has also a pair of posteriorly directed lobes, but neither the morphology of the
genital ligula (fig. 13c) nor the male appendages (figs. 17c, 21a, 25i) are consistent with the generic diagnosis of
Juenagrion.
FIGURE 70. Preliminary Neighbor Joining Tree of Telebasis species based on COI data (courtesy of Tim Faasen).
A preliminary molecular phylogeny based on COI data including two species of Helveciagrion, one of
Juenagrion, and ten of Telebasis (Tim Faasen in litt., Fig. 70) shows Juenagrion igapocola as sister species to
Helveciagrion obsoletum nesting in a clade with T. rubricauda Bick & Bick and T. dunklei Bick & Bick, rendering
Telebasis sensu Lencioni paraphyletic, thus further questioning the value of recognizing Juenagrion. Based on
these data, we consider Juenagrion a junior synonym of Telebasis and place J. nanae within the genus Telebasis as
Telebasis nanae (Lencioni, 2023), new combination.
FIGURES 71–75. Telebasis and Tepuibasis names treated here. 71: ♂ Telebasis igapocola, caudal appendages, mediodorsal
view (a), lateral view (b), dorsal view (c); 72: ♀ T. igapocola, hind lobe of pronotum and mesostigmal plates, mediodorsal
view; 73: ♂ Tepuibasis demarmelsi, habitus; 74: ♂ Tepuibasis fulva, habitus; 75: ♂ Tepuibasis garrisoni, habitus.
Finally, Lencioni's characterization of Telebasis (sensu stricto) employed largely secondary sexual characters
(male cerci, genital ligula, female prothorax) and stated that females possess a "mesanepisternal groove". However,
the type species, T. salva (Hagen) and others (e. g., T. brevis Bick & Bick, T. byersi, T. carminita Calvert, T. coccinea
(Selys), and T. incolumis) lack such a groove, and therefore Lencioni's attempt to morphologically define Telebasis
(s. s.) does not work. Garrison (2009) defined other clusters of species within Telebasis based on secondary genital
characters including the T. byersi, T. incolumis, T. salva complex and the T. filiola, Perty, T. inalata (Calvert), T.
willinki Fraser complex (which in some ways provides a link to Aeolagrion Williamson as discussed by Garrison
2009). Garrison (2009) also noted that the genital ligula in all but 11 species (T. carmesina, T. carminita Calvert, T.
carvalhoi Garrison, T. corallina (Selys), T. farcimentum Garrison, T. isthmica Calvert, T. leptocyclia, T. paraensei
Machado, T. racenisi Bick & Bick, T. sanguinalis Calvert, and T. simulata Tennessen) possesses a minute to well-
developed chitinized round or denticulate tubercle at each laterobasal angle of flexure (also shared with Aeolagrion
and present as a pair of isolated sclerotized digits in T. gigantea Daigle), yet he did not suggest that any of these
groups should be accorded generic rank.
Material Examined (in addition to that listed in Garrison 2009): Telebasis igapocola: PERU, Loreto Region: 1
♂, Aguas Negras, 0.5228°S, 75.2567°W, 145 m, 13 March 1994, Jerry Louton leg.; 1 ♂, 1 ♀ (holotype & allotype,
in copula), Tamshiyacu-Tahuayo Reserve, stream and flooded forest in triangle Río Tahuayo Río Blanco, 4.3317°S,
73.2339°W, 97 m, 17 February 2010, Tim Faasen leg. [RMNH].
Tepuibasis De Marmels, 2007
Austrotepuibasis Machado & Lencioni, 2011. New synonymy

Pseudotepuibasis Stand-Pérez & Pérez-Gutiérrez, 2020. New synonymy
Machado & Lencioni (2011) erected a new genus, Austrotepuibasis (type species: Austrotepuibasis demarmelsi
Machado & Lencioni, 2011, by original designation) for three new species: A. alvarengai, A. demarmelsi, and A.
manolisi, based primarily on differences in genital ligula structure (absence of sclerotized auricle process on apical
segment, two pairs of lateral lobes, pair of long terminal filaments), slight differences in abdominal coloration, and
posterior margin of female S10 entire. We have examined a small series of specimens corresponding to these species
(Figs. 73–75, see below), and they all have a sclerotized auricular structure along the lateral margin of the genital
ligula [two disjunct sclerotized auricular areas in A. demarmelsi, Fig. 76b, and a posteriorly directed process in T.
fulva (Fig. 77b) and Pseudotepuibasis garrisoni Stand-Pérez & Pérez-Gutiérrez (Fig. 78b)], and a deeply cleft S10
in the female, both characters shared with Tepuibasis. These similarities plus others listed by Machado & Lencioni
(2011) overlap considerably with known specimens of Tepuibasis we examined (T. fulva (Needham), T. neblinae
(De Marmels), T. nigra De Marmels), rendering the genus Austrotepuibasis indefinable on morphological grounds.
Just the possession of tubular long terminal filaments in the genital ligula (Fig. 76b) (similar shorter flattened
hyaline terminal filaments present in the type species, T. fulva, Fig. 77b) and occurrence in lower Amazon Forest
regions in Brazil do not, in our opinion, warrant erection of a new genus for these three species. Accordingly, we
consider Austrotepuibasis a junior synonym of Tepuibasis.
Stand-Pérez & Pérez-Gutiérrez (2020) erected a new genus, Pseudotepuibasis (type species by monotypy:
Pseudotepuibasis garrisoni Stand-Pérez & Pérez-Gutiérrez), for a new species based on differences they observed
between it and species of Tepuibasis in color pattern, genital ligula morphology (lacking an auricle-like process
present in Tepuibasis), and Fw quadrangle with costal side equal to distal side and frons angulated (not so in
Tepuibasis). However, the genital ligula (Fig. 78b) and appendage morphology (Fig. 78a) of a small series of males
of P. garrisoni from Peru are similar to those of the type species of Tepuibasis, T. fulva (Figs. 77a, b), differing
from it only by the more acuminate apical membranous lobes of the genital ligula (lobes shorter, abbreviated in
P. garrisoni, Fig. 78b) and more acuminate cercal tip, Fig. 77a (rounded in P. garrisoni, Fig. 78a), both of which
we consider specific, not generic, differences. Tepuibasis fulva also has a posteriorly directed microserrulate spine
on the lateral margin of the apical segment of genital ligula (Fig. 77b), and a similar non-serrulate aciculate spine
is present in the holotype of P. garrisoni (fig. 5b in Stand-Pérez & Pérez-Gutiérrez 2020) and in our series of P.
garrisoni (Fig. 78b). The two species differ in minor details (frons not angulate in T. fulva versus angulate in P.
garrisoni; postquadrangular veins in Fw 2 in T. fulva vs. 3 in P. garrisoni) but other species of Tepuibasis have 3
postquadrangular cells in the Fw, e. g., T. neblina and T. nigra, and the frons is also angulate in T. neblina. Finally,
FIGURES 76–78. Tepuibasis names treated here. 76: ♂ T. demarmelsi; 77: ♂ T. fulva; 78: ♂ T. garrisoni. a: caudal
appendages, mediodorsal view; b: genital ligula, ectal (upper), lateral view (lower); c: FW wing base.
the difference in the Fw quadrangle ("Males and females of Pseudotepuibasis have a FW quadrangle with costal side
equal to distal side…[a] character… that differs from Tepuibasis and Austrotepuibasis that have a FW with costal side
of quadrangle shorter than distal side…") shows an intermediate condition in the Fw quadrangle of T. demarmelsi
(Fig. 76c) compared to T. fulva (Fig. 77c) and P. garrisoni (Fig. 78c). The differences between Pseudotepuibasis
and Tepuibasis seem minor in comparison to their similarities, and they do not vary more than other species of
Tepuibasis vary among themselves according to De Marmels (2007), and we, therefore, consider Pseudotepuibasis
a junior synonym of Tepuibasis and consequently refer to the species described under it as Tepuibasis garrisoni
(Stand-Pérez & Pérez-Gutiérrez, 2020) new combination.
Material Examined: Tepuibasis demarmelsi: 17 ♂♂, 4 ♀♀: BRAZIL, Amapá State: 1 ♀, Alto Amazonas,
São Paulo de Olivença {3.4637°S, 68.9627°W}, 91 m, December 1960, Richard von Diringshofen leg. [MZUSP];
Pará State: 2 ♂♂, Altamira, {3.2035°S, 52.1907°W}, 91 m, 18 September 1963, Richard von Diringshofen leg.
[MZUSP, RWG]; 1 ♂, Floresta Nacional de Carajás, Buritizal II [FLONA Carajás 63], 6.0702°S, 49.95°W, 225
m, 21 March 2006, N. Ferreira leg. [RWG]; 2 ♂♂, 1 ♀, same data but 7 September 2006, N. Ferreira, Jr. & L.L.
Dumas leg. [RWG]; 1 ♂, same data but, 23 September 2007, N. Ferreira, Jr. & V. Alecrim leg. [DZRJ]; 2 ♂♂, same
data but 5 March 2008, N. Ferreira, Jr. & A. Santos leg. [DZRJ, RWG]; 6 ♂♂, Maloquinha, near Itaituba [FLONA
Carajás], 4.3151°S, 56.0792°W, 40 m, 21 March 2006, Expedição Permanente a Amazônia leg. [MZUSP, RWG]; 1
♂, Rio Azul, Pousada Rio Azul on forest trail, 200 yards from river; 14:00 hrs, 9.2444°S, 55.9867°W, 17 November
2008, Martin Reid leg. [RWG]; 1 ♂, 1 ♀, Rio Tapajós, Barreirinha [recte São Luiz do Tapajós, Itaituba], 4.4475°S,
56.2433°W, 23 m, 20 November 1970, Expedição Permanente a Amazônia leg. [MZUSP]; 1 ♂, Rio Xingu,Altamira,
{3.2035°S, 52.1907°W}, 91 m, 18 October 1963, Richard von Diringshofen leg. [MZUSP]; 1 ♀, Rio Xingu Camp,
ca. 60 km south of Altamira, 2nd jungle stream, trail 1 {3.65°S, 52.3667°W}, 138 m, 10 October 1986, Paul Spangler
& Oliver S. Flint, Jr leg. [RWG]; Tepuibasis fulva: 1 ♂, 1 ♀: VENEZUELA, Amazonas State: 1♂, 1♀, Atabao,
Duida, 3.3°N, 65.6167°W, 2100 m, 20 November 1991, Rafael Garcia Peña leg. [RWG]; Tepuibasis garrisoni: 3
♂♂, 2 ♀♀: PERU, Loreto Region: 3 ♂ ♂, 1 ♀, Tamshiyacu-Tahuayo Reserve, Terra Firme; Location code AF; No.
251, No. 264, 4.3728°S, 73.1858°W, 117 m, 6 August 2009, Tim Faasen leg. [RWG]; 1 ♂, Tamshiyacu-Tahuayo
Reserve, Terra Firme (forest and subcanopy Choroy stream); Location code J; No. 914, 4.3564°S, 73.1683°W, 118
m, 19 February 2010, Tim Faasen leg. [RWG]; 1 ♀, same data but Location code J; No. 971, 4.3564°S, 73.1683°W,
117 m, 20 February 2010, [RWG]; Tepuibasis neblinae: 1 ♂, 1 ♀: VENEZUELA, Amazonas State: 1 ♂, Cerro de
la Neblina Camp X, 0.9111°N, 66.0418°W, 1690 m, 13 February 1985, W. E. Steiner leg. [RWG]; 1 ♀, Arakamuni-
N {1.5333°N, 65.8167°W}, 1415 m, 24–30 October 1987, Anibal Chacon & C. Andara & Osuna leg. [RWG];
Tepuibasis nigra: 2 ♂♂: VENEZUELA, Amazonas State: 1 ♂, Cerro Yavi {5.7189°N, 65.8978°W}, 2150 m, 24–28
February 1995, J. L. Garcia leg. [RWG]; 1 ♀, Cerro Yutajé, 5.7596°N, 66.1343°W, 1750 m, 12-19 February 1995,
J. Clavijo leg. [RWG].
Tepuibasis demarmelsi (Machado & Lencioni, 2011). New combination
Austrotepuibasis alvarengai Machado & Lencioni, 2011. New synonymy

Austrotepuibasis manolisi Machado & Lencioni, 2011. New synonymy
Machado & Lencioni (2011) described three new species under Austrotepuibasis: A. alvarengai, A. demarmelsi
(type species by original designation), and A. manolisi. They provided a listing of 10 characters (four color and
six morphological characters, the latter set pertaining only to the male as only the female of A. demarmelsi was
known) to separate the three species. However, their figures of the genital ligula (figs. 13‒15) and male appendages
(figs. 1‒12) show hardly any differences and we have not been able to place our series of specimens matching their
description into one of the three specific names. Accordingly, we consider the three names to be synonyms and, as
first reviser, we have chosen T. demarmelsi to represent the senior synonym and provide illustrations of its male
here (Figs. 73, 76). We also provide renderings of the female mesostigmal plates of T. demarmelsi (Fig. 79), T. fulva
(Fig. 80) and T. garrisoni (Fig. 81), in hopes that they may aid in the recognition of subsequent collections of these
three species.
FIGURES 79–85. Tepuibasis, Philogenia, and Phoenicagrion names treated here. 79: ♀ T. demarmelsi; 80: ♀ T. fulva,
habitus; 81: ♀ T. garrisoni; 82: ♀ Philogenia carrillica; 83: ♀ Philogenia lankesteri; 84, 85: ♂ Phoenicagrion flammeum.
79–81: mesostigmal plates, dorsal view; 82–83: prothorax, lateral view: 84: genital ligula, ectal view (a), entolateral view
(b), lateral view (c).
Material Examined: BRAZIL, Amapá State: 1 ♀, Alto Amazonas, São Paulo de Olivença {3.4637°S,
68.9627°W}, 91 m, December 1960, Richard von Diringshofen leg. [MZUSP];—Pará State: 2 ♂♂, Altamira
{3.2035°S, 52.1907°W}, 91 m, 18 September 1963, Richard von Diringshofen leg. [MZUSP, RWG];—1 ♀, Rio
Xingu Camp, ca. 60 km S of Altamira, 2nd jungle stream, trail 1 {3.65°S, 52.3667°W}, 138 m, 10 October 1986,
Paul Spangler & Oliver S. Flint, Jr leg. [USNM];—6 ♂♂, Maloquinha, near Itaituba [FLONA Carajás], 4.3151°S,
56.0792°W, 40 m, 21 March 2006, Expedição Permanente a Amazônia MZUSP leg. [MZUSP, RWG];—4 ♂♂,
1 ♀, Floresta Nacional de Carajás, Buritizal I [FLONA Carajás 26], 6.0496°S, 49.89°W, 682 m, 7 September
2006, N. Ferreira, Jr. & L.L. Dumas leg. [DZRJ, RWG];—2 ♂♂, same data but, 5 March 2008, N. Ferreira, Jr. &
A. Santos leg. [DZRJ, RWG];—1 ♂, Floresta Nacional de Carajás, Buritizal II [FLONA Carajás 63], 6.0702°S,
49.95°W, 225 m, 21 March 2006, N. Ferreira leg. [RWG];—1 ♂, same data but 23 September 2007, N. Ferreira,
Jr. & V. Alecrim leg. [DZRJ];—1 ♂, Rio Azul, Pousada Rio Azul on forest trail, 200 yards from river, 9.2444°S,
55.9867°W, 17 November 2008, Martin Reid leg.;—1 ♂, Rio Xingu, Altamira {3.2035°S, 52.1907°W}, 91 m, 18
October 1963, Richard von Diringshofen leg. [MZUSP];—1 ♂, 1 ♀, Rio Tapajós, Barreirinha [actually São Luiz do
Tapajós, Itaituba], 4.4475°S, 56.2433°W, 23 m, 20 November 1970, Expedição Permanente a Amazônia MZUSP
leg. [MZUSP].
Possible synonyms pending further study
Philogenia carrillica Calvert, 1907
? syn. Philogenia lankesteri Calvert, 1924.
Paulson & Haber (2021) in their Appendix A ("Species Previously Recorded from Costa Rica") did not consider
P. lankesteri as a valid species ("…described from a single female, not considered a valid species"); however, they
did not indicate to which species it belongs. Calvert (1924) described P. lankesteri based on two females that only
differed from females of P. carrillica (Fig. 82) by the wider extension of the pronotal hind lobe along the sides
of the thorax (Fig. 83). He stated that he collected males of P. carrillica at the type localities of both, and that the
possibility existed that all females corresponded to polymorphic versions of P. carrillica as regards the hind pronotal
lobe shape. The fact that males of no further species other than P. carrillica have been collected in the area in the
intervening years (Paulson & Haber in litt.) seems to agree with the latter possibility. Finding of pairs in tandem or
copula of males of P. carrillica with both female morphs would confirm this likely synonymy.
Leptagrion andromache Hagen in Selys, 1876
? syn. Leptagrion jeromei Lencioni, Vilela & Furieri in Vilela et al., 2021
The original description of L. jeromei Lencioni, Vilela & Furieri (Vilela et al. 2021) puzzled us, as the description of
the holotype male looked much like two males we had identified as L. andromache. We submitted macrophotographs
of our two males (Figs. 56, 57) to Diogo S. Vilela who responded (in litt. 9 March 2023) as follows:
"…your specimen [Fig. 56], is really matching with the lectotype of L. andromache, and also with our male
from Amapá state. Cercus slightly arched, acute apex, shape of pterostigma (roughly squared [Fig. 31]), all of those
characteristics match perfectly. The only thing that is a bit different is the lateral projections of the hind lobe, that
seems to be much reduced in your specimen; at any rate, your specimen does seem to have a slight medial incision
on the hind lobe, as does the lectotype and our male from Amapá"..."your specimen [Fig. 57] may represent L.
jeromei by the following set of characters: cercus arched, with a medial constriction in dorsal view, blunt apex,
shape of pterostigma (roughly rectangular [Fig. 38]), and shape of hind lobe of prothorax, that seem[s] to be linear,
lacking the slight medial incision. In your specimen, the dorsal margin of S10 presents a marked cleft, whereas our
holotype does not have that incision. It's always nice to spot some variations among specimens, as frequently we
deal with very small type series. They [L. andromache, L. jeromei] are close species, but I believe that they represent
two different taxa.”
FIGURES 86–89. ♂ Phoenicagrion flammeum. 86–88: caudal appendages, dorsolateral view (86), lateral view (87),
dorsal view (88). 89: cercus tip, apical view (right column), ventroapical view (central right column), lateral view (central
left column), mediodorsal view (left column).
The lateral projection of the posterior lobe of the prothorax in L. andromache (and in other species) could be
variable and it should be used as a specific character with caution. The genital ligula is the same for both of our
specimens. We consider the distinctness of L. jeromei from L. andromache open to question, pending examination
of more specimens to determine if the differences observed correspond to intraspecific variability or to specific
differences.
Material examined: Listed under Leptagrion treatment above.
Phoenicagrion flammeum (Selys, 1876)
? syn. Phoenicagrion flavescens Machado, 2010
Phoenicagrion was erected by von Ellenrieder (2008a) for its type species, Leptagrion flammeum Selys, 1876 and
a new species, P. paulsoni. Machado (2010) in his generic revision described four new species, P. flavescens, P.
ibseni, P. karaja, and P. megalobus and included diagnostic keys for all six species. Faasen (2014) described P.
trilobum bringing the total number of species to seven. We have doubts about the distinctness of P. flavescens from
P. flammeum as, from Machado's description, it differs only by the flavescent condition of the wings and in having
an angled frons (rounded in P. flammeum), while the posterior lobe of the pronotum, genital ligula and caudal
appendages are essentially the same. Diogo Vilela examined the holotypes of Machado's species described under
this genus and observed (in litt.) that unfortunately the caudal appendages were missing from the holotypes making
a direct comparison impossible. Due to the similarity of secondary genital structures between both taxa, we wonder
if Machado's interpretation of the angulate frons in P. flavescens might be incorrect. Accordingly, we suggest a
possible synonymy between P. flavescens and P. flammeum pending examination of type material.
NOTE: We include detailed illustrations of Selys' lectotype of Leptagrion flammeum (Figs. 84‒88) and
illustrations of the tip of male cerci in other specimens we have identified as conspecific with P. flammeum (Fig. 89)
showing the variability in the placement of the terminal tooth.
Material examined (in addition to that listed in von Ellenrieder 2008b): COLOMBIA, Vaupés Department:
1 ♂, Mitu, Caño Cuduyari [MIT-94], 1.2917°S, 70.2557°W, 170 m, 24 September 2014, Melissa Sanchez Herrera
leg.;—PERU, Loreto Región;—BRAZIL, Amazonas State: 1 ♂, Sao Paulo de Olivença, Alto Rio Solimoes, 3.45°S,
68.8°W, 71 m, April 1932, from F. Wucherpfennig; Pará State: 2 ♂♂, 4 ♀♀, Agropalma, 2.4065°S, 48.7193°W, 58
m, 28–31 October 2016, Adolfo Cordero-Rivera leg.
Tuberculobasis croceum (Burmeister, 1839). New combination
Tuberculobasis mammilaris (Calvert, 1909)
A brief history of this taxon was provided by Calvert (1898). Costa & Garrison (2001) provided an English translation
of Selys' (1876) description who transferred it to Leptobasis. Selys' description reads in part: "Appendices anals
roussâtres, ayant à peine la moitié du 10e segment; les supérieurs en forme de tubercules épais, tronques au bout,
les inferieurs épais; un peu obscurs an bout, qui parait recourbe en haut, un peu aigu.["Caudal appendages russet,
having barely half [the length of] segment 10; the superiors in form of thick tubercles, truncated at the tip, the
inferiors thick, slightly dark at tip, which appears recurved upwardly, slightly pointed." and: " NB. Le Dr Hagen
ayant eu la bonté de me communiquer ce type précieux, qui est en très-mauvais état, j'ai pu le décrire avec soin et en
prendre un dessin exact." ["NB. Dr Hagen having the goodness to send me this precious type, which is in very bad
shape, I have been able to describe it with care and to make an exact drawing [reproduced here in Fig. 91]"
Garrison & von Ellenrieder (2019) examined and illustrated (Figs. 90, 95) the damaged holotype of Agrion
croceum Burmeister, noting: "Current examination of the thorax leaves no doubt that Agrion croceum is a member of
the genus Tuberculobasis Machado, 2009 and is likely a senior synonym of T. mammilaris (Calvert) or T. yanomani
(De Marmels)." Both T. mammilaris and T. yanomani are known from Suriname. Garrison had previously examined
the two syntypes of Leptobasis mammilaris and determined that the female syntype was actually Telagrion longum
Selys. Selys' illustration of the appendages of Agrion croceum in dorsal and lateral views (Figs. 91) are crude and
were most likely based on appendages that were not spread apart, obscuring the characteristic posteroventral branch
FIGURES 90–98. Agrion and Tuberculobasis names treated here. 90, 91, 95: ♂ A. croceum; 92, 97: ♂ T. yanomani;
93, 94, 96: ♂ T. mammillaris; 98: ♂ A. rufovittatum. 90: pterothorax, lateral view; 91: distal end of abdomen, dorsal
view (right), lateral view (left); 92, 93: S10, lateral view; 94: caudal appendages, dorsal view; 95–97: mesanepisternal
tubercles, dorsolateral view; 98: habitus.
of the cercus present in species of this genus. However, they tally slightly more with those of T. mammilaris (Figs.
93, 94) rather than with those of T. yanomani (Fig. 92), especially since the appendages were stated to be "un peu
aigu [slightly pointed]. The mesanepisternal tubercles of the holotype of A. croceum (Fig. 95) approach more those
of T. mammilaris (Fig. 96) than those of T. yanomami (Fig. 97) but appear to be slightly larger and more conical. We
suspect T. mammilaris is a junior synonym of T. croceum but consider that this should be confirmed once a larger
series of specimens from Suriname is available for study to establish if the apparent differences are breached by
intermediate characters as a result of intraspecific variability.
Material Examined: Tubeculobasis mammilaris: SURINAME: Para District: 2 ♂♂, 1♀, Vier Kinderen, RMNH.
INS. 514311, 5.5167°N, 55.5333°W, 14 m, 11 December 1963, Jean Belle leg. [RWG]; 3 ♀♀, Vier Kinderen, RMNH.
INS. 514513, 5.5167°N, 55.5333°W, 14 m, 11 December 1963, Jean Belle leg.; 1♀, Republiek bos, RMNH.INS.
514470, 5.5°N, 55.2°W, 15 m, 23 November 1963, D. C. Geijskes leg.; Tuberculobasis yanomami: SURINAME:
Marowijne District: 1 ♀, Wia Wia, km 10, RMNH.INS. 514037, 5.8881°N, 54.4792°W, 25 November 1948, D.C.
Geijskes leg. [RMNH]; Para District: 3♂♂, Vier Kinderen, RMNH.INS. 513885, 5.5167°N, 55.5333°W, 14 m,
11 December 1963, Jean Belle leg.; Sipaliwini District: 3♂♂, Tapanahoni River, RMNH.INS. 514628, 4.13°N,
54.63°W, 6 April 1963, Jean Belle leg.
Incertae sedis
Agrion rufovittatum Blanchard, 1846
Blanchard (1846) described and illustrated Agrion rufo-vittatum based apparently on a single specimen from Santa
Cruz Province, Bolivia. The Latin description is short ("Supra nigra, infra testacea; capite nigro, macula oculari,
margine, labroque rufotestaceis; prothorace nifo-vittato; alis diaphaneis, nerrvis nigris; abdormine elongalo
ante apicem viridi-cinereo. Long. 38 mill.; enverg., 45 mill.") and the illustration shows a species with numerous
postnodal crossveins (Fig. 98). Kirby (1890) transferred the species to Micronympha (= Ischnura) with doubt ("M.
(?) Rofovittata, Blanch."), and Davies & Tobin (1984) listed it as species of Ischnura, a decision that has been
followed by all subsequent catalogs. The illustration of the holotype appears to show a female of what is a largely
pale coenagrionid. Its status is uncertain judging from the illustration, but we doubt that the specimen can be placed
in the genus Ischnura due to the numerous px (Fw=17, 20, Hw=18, 14) and extreme recession of IR1.
Discussion
Several of the specific synonyms suggested here are a result from recognizing intraspecific variability after examining
additional specimens; variability which had not been previously evident because species were described based on a
limited number of specimens from a limited geographic area. Others were caused from placing too much weight on
differences between specimens described as new species and illustrations of types of previously described species
without accounting for possible nuances resulting from a slightly different orientation or preparation of diagnostic
structures, postmortem or ontological changes, or inaccurate illustrations. The introduction of several of the genera
we are synonymizing here was often based solely on secondary sexual characters of males (often genital ligula),
rendering placement of females, whether described or unknown, into their respective genera difficult or impossible.
Vilela Silva et al. (2019) provided the following comments in diagnosing group B species of Heteragrion: "However,
it should be mandatory for current taxonomic research on females to be as complete as male descriptions, whenever
possible. A complete female description may avoid difficulties in establishing reliable characters, facilitate the
construction of identification keys and allow female identification in the absence of males." We believe this is good
advice to be followed today.
Recognizing mistakes and correcting them is part of science; as Tattersall (2003) stated: ‘In science it is no
crime to be wrong unless you are (inappropriately) laying claim to truth. What matters is that science as a whole is
a self-correcting mechanism in which both new and old notions are constantly under scrutiny’.
Acknowledgments
We appreciate the critical input from the colleagues who have reviewed our manuscript: Drs. Jürg De Marmels (Museo
del Instituto de Zoología Agrícola, Maracay), Tim Faasen (Maarheeze, The Netherlands), Dennis Paulson (Seattle,
Washington), Kenneth J. Tennessen (Wautoma, Wisconsin) and Yiselle P. Cano Cobos (Universidad Nacional de
Avellaneda, Buenos Aires, Argentina). We thank Drs. Holger Hunger and Franz-Josef Schiel for sharing specimens
of I. sobrina with us and Dr. Jan van Tol for the high-resolution scan of Agrion rufovittatum. The nomenclatorial
changes we advocate here are entirely our suggestions, and the reviewers are in no way responsible for any errors
or opinions expressed here.
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Zootaxa 5405 (1): 001–042 ISSN 1175-5326 (print edition)

Taxonomic revisions of some Neotropical Zygoptera (Odonata)

Accepted by M. Marinov: 14 Dec. 2023; published: 26 Jan. 2024 

Material and methods

Proposed taxonomic changes

Lestes debellardi De Marmels, 1992

Lestes fernandoi Costa, Souza & Muzón, 2006. New synonymy

Lestes tikalus Kormondy, 1959. New synonymy

Palaemnema brevignoni Machet, 1990

Palaemnema brasiliensis Machado, 2009. New synonymy

Hetaerina duplex Selys, 1853

Hetaerina aurora Ris, 1918. New synonymy

Hetaerina longipes Hagen in Selys, 1853

Hetaerina hebe Selys, 1853. New synonymy

Mnesarete guttifera (Selys, 1873)

Mnesarete mariana Machado, 1996. New synonymy

Acanthagrion obsoletum (Förster, 1914)

Acanthagrion hartei Muzón & Lozano, 2005. New synonymy

Ischnura sobrina Schmidt, 1943

Homeoura sobrina: von Ellenrieder 2008b (transfer to Homeoura Kennedy).

Fredyagrion Lencioni, 2022. New synonymy

Leptagrion macrurum (Burmeister, 1839)

Oxyagrion ablutum (Calvert, 1909)

Oxyagrion bruchi Navás, 1924. New synonymy

Telebasis Selys, 1865

Helveciagrion Machado, 1980.—Garrison 2009 (synonymy);—Lencioni 2023 (removed from synonymy).

Tepuibasis De Marmels, 2007

Austrotepuibasis Machado & Lencioni, 2011. New synonymy

Tepuibasis demarmelsi (Machado & Lencioni, 2011). New combination

Austrotepuibasis alvarengai Machado & Lencioni, 2011. New synonymy

Possible synonyms pending further study

Philogenia carrillica Calvert, 1907

? syn. Philogenia lankesteri Calvert, 1924.

Leptagrion andromache Hagen in Selys, 1876

Phoenicagrion flammeum (Selys, 1876)

? syn. Phoenicagrion flavescens Machado, 2010

Tuberculobasis croceum (Burmeister, 1839). New combination

Tuberculobasis mammilaris (Calvert, 1909)

Agrion rufovittatum Blanchard, 1846

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Accepted by M. Marinov: 14 Dec. 2023; published: 26 Jan. 2024