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Zygoptera: Coenagrionidae) Odonalologica 13 (4): 501-527 December I. 1984 The genus Nehalennia Selys, its species and their phylogenetic relationships (Zygoptera: Coenagrionidae) J. De Marmels Departamento e Institute de Zoología Agrícola, Facultad de Agronomía, Universidad Central de Venezuela, Apartado 4579, Maracay 2101-A, Aragua, Venezuela Received January 12, 1984 / Accepted April 5, 1984 Nehalennia is revised redefined. is that The genus Selys and It demonstrated is of Nehalennia Brazilian is Argiallagma Calvert a synonym Sel. The N. selysi Kirby redescribed and assigned to minuta Sel., as a subspecies. A lectotype is designated($: Sao Joao del Key, Minas Gerais, Brazil; Coll. Selys, Inst. R. Sc. Nat. Belg., Brussels). The is related other with Irons. It includes genus closely to genera anangulate actually six species and one subspecies, which are tentatively arranged into a "phylogenetic tree”, based on differences and similarities of morphological characters (larval and male of adult). The penes and the anal appendages all species are figured. A new interpretation of the superior appendages is given and a terminology for both the superiors and the inferiors is proposed. Figures ofthe female prothorax of all species are provided, as well as drawings of other important structures and colour patterns. minuta For the seven taxa a list of synonyms is given. In the case of N. and pallidula Calv. a more exhaustive bibliography is added. Measures and venational features (males) are presented, as well as a description of the habitat preferences and dates on the distribution. INTRODUCTION the Nehalennia Sei. is Doubtless, genus not among those coenagrionids most in need of revised. This urgently being was not my purpose, indeed. My aim was simply to describe two supposedly new species ofthis genus from Venezuela,which have been set apart by the late Dr J. Râcenis many years ago, but apparently were forgotten by him later. — Themore I advanced in the study ofthese two species, the it of the more became clear that I had to examine as many other species genus Nehalennia as possible ifI wished to arrive at any definite conclusions. A special 502 J. De Marmels need was the reexaminationand redescription of the Brazilian N. selysi Kirby, and a careful study ofthe odd Argiallagma minutum(Sel.). — It was necessary to relax, with the help ofammonia, the penes and the anal appendages ofall species, in order This to allow comparisons. proceeding proved especially useful in the case of the anal appendages, which are very complicated and almost throughout misinterpreted in the literature. However, once relaxed they could be moved around with and it thenbecame whether branch the a pin, easy to see a belonged to superior orto the inferior appendage. When 1 finally brought to an end my comparative studies, I had checked virtually all known believed in Nehalennia. As species currently to -or to- belong the genus a the Calvert to with consequence, generic name Argiallagma had be synonymized Nehalennia Sel. Eitherof the two presumably new species from Venezuelaproved to be identicwith an already well-known species, while N. selysi couldbe given only a subspecific status. The of the would have the and realization present paper been impossible without help kind collaboration of the Dr S. ASAHINA Dr D.A.L, DAVIES following persons: (Tokyo, Japan), (Cambridge. United Kingdom), DrP. DESSART(Institut Royal des Sciences Naturellesde Belgique. Brussels, Belgium),DrT.W. DONNELLY (Binghamton,N.Y., U.S.A.), Dr H.J. DUMONT (Ghent. Belgium). Dr R.W. GARRISON (Azusa, Cal., U.S.A.), Mrs Leonora K. GLOYD (Ann Arbor. Mich., U.S.A.), Dr B. KIAUTA (Utrecht, The Netherlands), Dr J. LEG RAND (Museum National d'Histoire Nalurelle, Paris, France), Dr A.B.M, MACHADO (Belo Horizonte, Brazil), Dr M. MORATORIO(UniversidadNacional Experimentaldelos Llanos Occidentales"EzequielZamora”, Guanare. Venezuela) and Dr M.J. WESTFALL (University of Florida, Gainesville,Fla.. U.S.A.). 1 wish sincere thanks these and the institutions for the to express my to persons to they represent, loan of material, including type specimens, the furnishingof literature and many valuable comments. THE GENUS NEHALENNIA SELYS The name Nehalenniawas established by Selys (SELYS & HAGEN, 1850)for his ”ler Groupe” of the genus Agrion, with the eurasian Agrion speciosum SELYS definitionof his CHarpentier as the "type”. Later (1876) enlarged the now ”Sous-Genre 9. — Nehalennia”, adding new characters. In view of the results of the present work one may characterize the genus Nehalennia of small with frons and as a group coenagrionids a sharply angulate widely separated eyes. Head, thorax and normally the abdomenare more or less metallic extensively marked with brassy black or green. Postocular spots are visible. This line to absent, but a pale occipital line is normally may be reduced a in The second of the very short mark around the neck, some species. segment antenna is marked with a broad centralwhiteband. The legs are short with short to fairly long tibial spines. Thewings are hyaline with a lozenge-shaped pterostigma, cross-veins: in front which covers one cell or less. Number of postnodal 7-12 wing, 6-11 in hind Numberof antenodal cells in the discoidal field2-3. ofR wing. Origin , in fore wing between third and sixth postnodals, in most species proximal to the fifth. Origin of R, in hind wing between the third and fifth postnodals, normally Taxonomy of Nehalennia 503 to the fourth. IR between the sixth and ninth in proximal 2 originates postnodals in in front wing as well as hindwing. Arculus atsecond antenodalorslightly distally, both wings. The cubito-anal cross-vein is separated from the posterior border of distance of its is the wing by a at least own length. The shape ofthe quadrangle variable, even within species. In thefront wing thecostal side ofthequadrangle may be shorter or longer thanthe proximal side. The penis has an internal and a terminal fold, but lacks any kind offlagella or spines. Only the lateral lobesofthe distal segment do show specific modifications. is No vulvar spine present on the sterniteofthe eighth segmentin the female. Atthis obtuse lamina be observed. reaches the place an projecting may The ovipositor to end of the abdomen.The dorsumofone or more ofthe distalabdominalsegments display a pale blue colour ("Schlusslicht”, see SCHMIDT, 1953). The anal appendages ofthe male are shorter thanoras long as thetenthabdominalsegment. The superiors are two or three-branched. The inferiors are variably shaped, but directed always steep, upwards, and with their modifiedparts in the upper half. in The sharply angulate frons Nehalenniarenders it easy to separateits species at from similar of Ischnura which is characterized a glance species Charp., a genus by a rounded frons and a compact head with a smaller distance between the Generic differences ofthe larval of the two less compound eyes. stage genera are obvious, although in Nehalenniamost species have one principal mentalseta and a varying number ofsmall accompanying setae (as have the allied genera Telebasis Sel., Ceriagrion Sel., etc.), while Ischnura larvae have threetofive principal mental setae (as has Enallagma Charp. and other related genera). Nehalennia is American essentially an genus distributed from Brazil across northern South Central America the northwestern America, and Antilles, up to Canada. has far been found. — Only one palaearctic species (the generotype) so The genus clearly occupies a specific ecological niche. Its habitat requirements are fulfilled only by lentic habitats with flat water, invaded by emergent, erect, but low and dense variably vegetation in swamps, shores of smaller lakes, as well as around ponds (’’Verlandungszonen”, cf. DE MARMELS & SCHIESS, 1977). WHAT IS ARGIALLAGMA? The bibliography concerning Nehalenniaminuta(Sel.) shows how one and the same species has been described several times under different names and has continuously been pushed for and back across the genera, but not in a single case attributed under its in which it legal specific name to that genus actually belongs; Nehalennia Selys. This in spite of multiple evidence, not only of adult characters, but oflarval features well. — the which as In following 1 shallexpose the arguments brought me to the conclusion that Argiallagma Calvert (pro Argiallagma Selys, nom. nud., see COWLEY, 1934) is nothing else than a synonym of Nehalennia Selys. 504 J. De Marmels KENNEDY (1920) thought that the "curious Argiallagma" joins characters at once with his ’’Argia series” "Enallagma — Acanthagrionseries”and "Nehalennia — Telebasisseries”. Still DAVIES (1981) files Argiallagma underthe Ischnurinae, but Nehalenniaunder the Coenagrioninae and the closely related Telebasis under the Pseudagrioninae. The morphological characters historically andcurrently used to definethegenus Trichocnemis Argiallagma (type species: minutaSel.) inits adult stageare: the long tibial the of spines, presence a vulvar spine on theeighth abdominal segment ofthe the of female, wing venation, absence dorsal processes on the tenth abdominal of the male, and colour SELYS ”Le segment pattern. Already (1876, p. 498) says: premier group, compose du seul Enallagma aduncum [= Argiallagma minutum] est bien notablepar la longueur des cils divariques des tibias, qui le feraient classer les si vulvaire parmi Argias, 1’epine de la femelle ne s’y opposait”. CALVERT ’This (1907, p. 376) quotes: genus [Argiallagma] apparently belongs to the same divisionofthe do biserial legion Agrion as Hyponeuraland.Argia, since the hairs on the legs are nearly, or quite, twice as long as the intervals separating them. It differs from those genera [...] in the presence of an apical ventral spine on the eighth abdominal segmentof the female”. Laterauthors followedthis diagnosis (BYERS, 1930, JOHNSON & WESTFALL, p. 168; 1970, p. 60), or exposed criticism, but without presenting nomenclatorial alternatives (GE1JSKES, 1943). — I do not think that the length of the tibial spines can serve as a generic character. In Acanthagrion Sel. most species have short tibialspines, but A. rubrifrons Leonard and A. longispinosum Leonard, as well as two yet undescribed species from in Venezuela (DE MARMELS, press), have very long tibial spines.
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