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Citation: Makarkin VN, Yang Q, Shi C, Ren D (2013) The presence of the recurrent veinlet in the Middle Jurassic Nymphidae (Neuroptera): a unique character condition in Myrmeleontoidea. ZooKeys 325: 1–20. doi: 10.3897/zookeys.325.5453
A well-developed recurrent veinlet is found in the forewing of two species of Nymphidae from the Middle Jurassic locality of Daohugou (Inner Mongolia, China), Liminympha makarkini Ren & Engel and Daonymphes bisulca gen. et sp. n. This is the first record of this trait in the clade comprised of the superfamilies Myrmeleontoidea and Chrysopoidea. We interpret the recurrent veinlet in these species as a remnant of the condition present more basally in the psychopsoid + ithonoid + chrysopoid + myrmeleontoid clade (i.e., as a plesiomorphy). Other venational character states of Daonymphes bisulca of interest include the configuration of subcosta anterior (ScA), which is very similar to that of extant Nymphidae. We consider the short ScA terminating on ScP to be an autapomorphy of Neuroptera.
Neuroptera, Nymphidae, recurrent veinlet, subcosta anterior, Daohugou, Middle Jurassic
Understanding the evolution of particular morphological characters during the broad historical development of higher insect taxa is a very interesting and important question. In fact, the phylogeny of a taxon may be realistic only if the evolutionary transformations of all available characters are correctly interpreted (
The forewing humeral veinlet (i.e., the proximal-most veinlet of ScP) in all Neuropterida (and generally in all Neoptera) is plesiomorphically a simple, crossvein-like. However, in some families of Neuroptera it is curved, obviously directed towards wing base (i.e., recurrent), and often profusely branched. This condition (the recurrent veinlet for short) is one of the significant informative venational characters found within this order. It is characteristic of several families and subfamilies, the vast majority of which have the costal space basally dilated (see below). However, it was hitherto not found in any of numerous species of the clade comprised the superfamilies Myrmeleontoidea (Nymphidae, Nemopteridae, Ascalaphidae, Myrmeleontidae, Palaeoleontidae and Babinskaiidae) and Chrysopoidea (Chrysopidae, Mesochrysopidae and Ascalochrysidae) (
Nymphidae are generally accepted to be the most basal family in the superfamilies Myrmeleontoidea, and the only myrmeleontoid family known from the Jurassic. In general, 18 fossil (named) species are known in the family (
This paper is based on two specimens collected from the Daohugou locality and housed in the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China (CNUB; Dong Ren, curator). Daohugou Village is situated in Shantou Township, Ningcheng County, Inner Mongolia, China. The insect-bearing beds of the Daohugou locality are considered to belong to the Jiulongshan Formation, and are dated as Bathonian, Middle Jurassic (
The specimens were examined under a Leica MZ12.5 dissecting microscope; line drawings were prepared with CorelDraw 12 graphics software and Adobe Illustrator CS5 with the aid of Adobe Photoshop CS3; photographed by a Nikon SMZ1000 stereomicroscope; the whole specimens were photographed by Nikon D100 Digital Camera.
Terminology and abbreviations. Wing venation terminology in general follows
Venational abbreviations. AA, analis anterior; AP, analis posterior; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; M, media; MA, media anterior; MP, media posterior; R, radius; RA, radius anterior; RP, radius posterior; RP1, proximal-most branch of RP; rv, recurrent veinlet; ScP, subcosta posterior.
Daonymphes bisulca sp. n.
Forewing broad proximally [strongly narrowed proximally in Liminympha]; vast majority of proximal subcostal veinlets forked [simple in all other Mesozoic genera]; crossveins between branches of MP absent [present in Nymphites Haase, 1890, Mesonymphes Carpenter, 1929]; CuP space broad, nearly two times as wide as intracubital space [nearly as wide as intracubital space in Liminympha].
From Daohugou, the locality of the type species, and Nymphes, a genus-group name. Gender: feminine.
This genus is most closely related to four other Mesozoic genera (Nymphites, Mesonymphes, Liminympha and Sialium Westwood, 1854) but clearly distinguished from these as indicated in the diagnosis. The latter genus is only known from single hind wing from the early Berriasian of Purbeck, England. In general, Nymphidae occur very rare in the Jurassic, only six specimens are known. The monotypic genera Liminympha (Middle Jurassic [Bathonian/Callovian] of Daohugou, China) and Mesonymphes (Late Jurassic [Tithonian] of Solnhofen, Germany) are represented by almost complete specimens possessing both fore- and hind wings. The type species of Nymphites from Solnhofen is represented by only two incomplete hind wings. Two species from Daohugou are assigned to this genus: one species (two specimens) have almost complete fore- and hind wings; the other only hind wings (
http://zoobank.org/61D8C637-96AD-4D41-BEF6-270FE5F3A6BA
http://species-id.net/wiki/Daonymphes_bisulca
Figs 1, 2, 6AAs for the genus.
Forewing 29 mm long as preserved (estimated complete length about 40 mm), 11.5 mm wide. Costal space relatively broad for entire length, narrowed basally. Humeral veinlet recurrent, with one short branch. Subcostal veinlets somewhat curved, mostly forked once or twice, shallowly, few deeply; several proximal veinlets simple. ScA well developed, terminating on ScP within humeral area. Subcostal space narrow, with several scarce crossveins. RA space relatively narrow, slightly narrowed towards wing apex, with relatively numerous crossveins. RP with nine preserved branches, becoming more closely spaced towards wing apex. Crossveins rather numerous over entire radial space, irregularly spaced. M forked distal of origin of RP. MA incomplete, simple for entire preserved length. MP distally pectinate, with six preserved branches, most of these forked once or twice; no crossveins detected between branches. Cu divided into CuA, CuP rather far from wing base. CuA distally pectinate, with six preserved branches, all forked once or twice; one series of crossveins between branches of CuA, continued CuP (‘pseudo-CuP’). CuP long, strongly pectinately branched with nine long branches; of four proximal-most branches, two forked; all distal branches dichotomously forked; no crossveins detected. Eleven crossveins between CuA, CuP rather irregularly spaced. AA3+4 and its posterior branch deeply forked. Basal crossvein between CuP, AA3+4 very short; distal crossvein strongly oblique. AP1+2 deeply forked. AP3+4 not preserved. Membrane around one crossvein between MA, MP near MP and three crossveins between CuA, CuP near CuA broadly heavily shaded; most crossveins between RA, RP apparently broadly shaded, pale fuscous.
Daonymphes bisulca gen. et sp. n. Holotype CNU-NEU-NN2011119 as preserved. Scale bar = 10 mm.
Daonymphes bisulca gen. et sp. n. Forewing venation of the holotype CNU-NEU-NN2011119. Scale bar = 10 mm.
Holotype CNU-NEU-NN2011119, deposited in CNUB; an incomplete forewing.
Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China. Jiulongshan Formation, Middle Jurassic.
From the Latin bisulcus, -a, -um, forked, divided into two parts, in reference to its forked subcostal veinlets.
http://species-id.net/wiki/Liminympha
Liminympha makarkini Ren & Engel, 2007, by original designation.
Forewing strongly narrowed proximally [broad proximally in Nymphites, Daonymphes gen. n. and Mesonymphes]; all veinlets of ScP simple [at least distal veinlets forked once or twice in Nymphites, Daonymphes gen. n.]; CuP space narrow, nearly as wide as intracubital space [nearly two times as wide as intracubital space in Nymphites, Daonymphes gen. n. and Mesonymphes]. Hind wing MP with four pectinate branches [eight in Sialium]; branches of CuA not connected by crossveins [at least proximal branches of CuA connected by crossveins in Nymphites]; anterior branch of CuP simple [deeply forked in Nymphites and Mesonymphes].
http://species-id.net/wiki/Liminympha_makarkini
Figs 3 – 5, 6BBody (metathorax, abdomen) poorly, fragmentarily preserved. First abdominal tergite rather long; distally with distinct transverse suture, probably heavily sclerotized; medially with mediolongitudinal short suture; portion of 1st tergite distal to transverse suture (‘Transversalnaht’ of
Liminympha makarkini Ren & Engel, 2007. Holotype CNU-NEU-NN1999024 (counterpart) as preserved.Scale bar = 5 mm.
Liminympha makarkini Ren & Engel, 2007. Metathorax and the basal part of abdomen of the holotype CNU-NEU-NN1999024 (counterpart).Wetted with ethanol. Abbreviations: ms, mediolongitudinal suture of 1st abdominal tergite; mtpn, metapostnotum; mtscl, metascutellum; ts, transverse suture of 1st abdominal tergite; T1, T2, 1st, 2nd abdominal tergites.Scale bar = 1 mm.
Forewing elongate, narrowed in proximal portion, most dilated at distal 3/4 length; about 30 mm long, 8 mm wide. Costal space narrow, basally narrowed, distally dilated. All preserved subcostal veinlets simple, strongly oblique distally; veinlets of ScP+RA dichotomously branched. Humeral veinlet rather strongly recurrent, branching not detected (Fig. 6B). Subcostal space very narrow; two crossveins in distal part detected, others possible. RA spaces slightly narrower than costal space, narrowed towards apex; crossveins irregularly spaced for entire preserved portion. RP originates rather near wing base, with about 16 branches; RP1 originates far from origin of RP; at least four proximal-most branches widely spaced, more distal branches closely spaced. In right forewing, RP2 terminated at RP1; RP3, RP4 fused for short distance (probably aberrations). Crossveins between branches of RP very scarce, restricted to area between RP1 to RP5. M appears fused with R basally; forked at nearly equal distance from origin of RP, origin of RP1. MA long, slightly arched, distally few-branched. MP long, its anterior trace (stem of MP) nearly straight before terminal branching; with five long distal branches, quite strongly inclined. Crossveins between R/RP and MA, MA and MP irregularly spaced, arranged differently in right, left wings. Cu dividing into CuA, CuP rather near to wing base. CuA long, smoothly curved anteriorly, pectinate with four long branches, each dichotomously branched. Between branches of CuA at last three crossveins forming gradate series continued CuP (‘pseudo-CuP’). CuP long, pectinately branched, with seven-eight rather short branches, most simple. AA3+4 rather short; stem simple, with one or two simple branches. AP1+2 incompletely preserved, probably simple. AP3+4 not preserved. One dark rather big spot in distal portion of radial space might be present (apical half of other wing not preserved).
Liminympha makarkini Ren & Engel, 2007. Wing venation of the holotype CNU-NEU-NN1999024 A right forewing B left forewing C right hind wing D left hind wing. Scale bar = 5 mm (all to scale).
Basal portion of forewings of the Middle Jurassic Nymphidae showing the recurrent veinlet (rv) and the anterior subcosta (ScA) A the holotype of Daonymphes bisulca gen. et sp. n., CNU-NEU-NN2011119 B the holotype of Liminympha makarkini Ren & Engel, 2007, CNU-NEU-NN1999024. Both wetted with ethanol. Scale bar = 1 mm.
Hind wing similar in shape to forewing, slightly narrower; about 28 mm long, 7 mm wide. Costal space narrow, dilated distally. Subcostal veinlets simple, becoming more oblique, closely spaced, curved towards apex. Humeral veinlet bent to base, nearly straight, not branched. Subcostal space narrow; three crossveins detected, others possible. ScP, RA fused far from wing apex; preserved veinlets of ScP+RA long, dichotomously forked; crossveins between them not detected. RA space basally as wide as costal space, narrowed towards apex; crossveins rather regularly spaced for entire preserved portion (right wing). RP originated near wing base, with about 15 branches; RP1 originated far from origin of RP (but closer to wing base than in forewing); five proximal-most branches widely spaced, other (distal) branches closely spaced. Crossveins between branches of RP very scarce, restricted to area between RP1 to RP4. Origin of M and its fork not preserved. MA long, nearly straight, distally dichotomously branched. MP long, its anterior trace (stem of MP) slightly incurved, with four long distal branches, quite strongly inclined. Origin of Cu and its dividing into CuA, CuP not present. CuA long, pectinate, with 15 branches, which proximally simple, distally once or twice forked. CuP short, deeply forked. Distal crossvein between CuA, CuP rather short connecting CuA, anterior branch of CuP fork. Anal veins not preserved. Crossveins between R/RP and MA, MA and MP, MP and CuA poorly preserved, irregularly spaced, arranged differently in right and left wings; crossveins between branches of MP, CuA absent.
Holotype CNU-NEU-NN1999024 (part, counterpart), deposited in CNUB; an incomplete specimen.
Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China. Jiulongshan Formation, Middle Jurassic.
This redescription is only based on the counterpart of the holotype.
The recurrent veinlet is known in the following extant taxa of Neuroptera: all species of Psychopsidae and Ithonidae (including Polystoechotidae and Rapismatidae), the vast majority of Hemerobiidae; rarely in Berothidae (including Rhachiberothinae) and Mantispidae (only Symphrasinae).
The recurrent veinlet has not been detected in any Permian Neuroptera (Permithonidae, Archeosmylidae) and the majority of Triassic taxa represented by these two families, and ‘Mesoberothidae’; however the basal portion of the forewing in any reported specimen of the latter is poorly preserved (e.g.,
Of the Early Jurassic Neuroptera, the recurrent veinlet is detected in all psychopsoids whose forewing base is well preserved (e.g.,
The costal space in all numerous undescribed Ithonidae from the Middle Jurassic locality of Daohugou is basally narrowed, and therefore, the recurrent veinlet (where well developed) has short simple branches. Sometimes, the humeral veinlet is not branched and crossvein-like (VM, pers. obs.). Younger Ithonidae (Early Cretaceous to Recent) have the costal space usually broader basally and have a well developed recurrent veinlet (e.g.,
In the Mesozoic family Brongniartiellidae, the well-developed recurred veinlet with long branches is detected in two species from the Early Cretaceous of the Baissa locality, Transbaikalia (
The presence of the recurrent veinlet was long considered a plesiomorphic condition in Neuroptera (e.g.,
The occurrence of the recurrent veinlet through Neuroptera families. The phylogeny of the order is based on that of
The two species described above from the Middle Jurassic locality of Daohugou are the oldest known record of Nymphidae, indeed, of Myrmeleontoidea. Myrmeleontoidea are believed to belong to the suborder Myrmeleontiformia, which also includes Psychopsidae (
The recurrent veinlet is known mainly in species whose costal space is basally dilated. The strong dilation of the basal portion of the costal space is characteristic of Neuroptera. It never occurs in other Holometabola including other Neuropterida orders, although in some taxa, the costal space is markedly dilated at some distance from wing base (e.g., Raphidioptera:
The ScA in all Neuroptera (when present) has a similar configuration: a short vein structure terminating on ScP before the recurrent veinlet. The structure of ScA of Daonymphes bisulca gen. et sp. n. is very similar to that found in extant Nymphidae, especially Nymphes Leach, 1814 (VM, pers. obs.) and closely related genera (e.g., Austronymphes sp.: see
The ground-plan Neoptera wing venation is hypothesized to have ScA consisting of two separate veins with no common stem, ScA1+2 and ScA3+4; the former runs to the costal margin, the latter is terminated on ScP (
The configuration of ScA characteristic of Neuroptera (i.e., short and terminating on ScP) does not occur in other orders of Neoptera, and therefore may be considered as an autapomorphy. On the other hand, the similarities of ScA in Neuroptera and some Paleozoic Palaeoptera (see above) may indicate their homology, and therefore a symplesiomorphy. The latter, however, appears to be less probable than the former.
We thank S. Bruce Archibald (Simon Fraser University, Burnaby, Canada) for correction of English; Shaun Winterton (California State Collection of Arthropods, Sacramento, California, U.S.A.) and André Nel (Museum National d’Histoire Naturelle, Paris, France) for critical reading of the manuscript. This work was supported by the National Basic Research Program of China (973 Program) (grant 2012CB821906), National Science Foundation of China (grants 31230065, 31272352, 41272006), Project of Great Wall Scholar and KEY project of Beijing Municipal Commission of Education Project (grants KZ201310028033), China Geological Survey (grant 1212011120115).