a new unexpected species of chrysopa leach from cyprus

A new unexpected species of Chrysopa Leach from Cyprus with biogeographic remarks (Neuroptera: Chrysopidae)A new unexpected species of Chrysopa Leach from Cyprus with biogeographic remarks (Neuroptera: Chrysopidae)
Davide Badano a,b* & Christodoulos Makrisc
aDISTAV, University of Genoa, Corso Europa 26 I-16132, Genoa, Italy; bDepartment of Biology and Biotechnology “Charles Darwin”, Sapienza University of Rome, Piazzale A. Moro 5 I-00185, Rome, Italy; cEthnikis Antistaseos 21 C-3022, Lemesós, Cyprus
(Accepté le 14 avril 2020; publié en ligne le 18 mai 2020)
Summary. Cyprus is rich in endemics; however, all but two lacewing species are shared with the mainland. Thus, the discovery of a new morphologically distinct species of chrysopid belonging to the extensively studied genus Chrysopa, Chrysopa niki n. sp., appears remarkable. While most Chrysopa species differ in relatively subtle morphological characters, the new taxon is set apart from all Western Palaearctic congeners due to a unique combination of pattern, venational, leg and genital characters. The characteristics of C. niki n. sp. suggest that it might be an endemic to Cyprus, not strictly related to other congeners, underlining the importance of the discovery. The biogeography of Cypriot Neuroptera is discussed in light of this new finding.
Résumé. Une nouvelle espèce inattendue de Chrysopa Leach de Chypre avec des remarques biogéographiques (Neuroptera : Chrysopidae). Chypre est riche en espèces endémiques, mais tous les névroptères à l’exception de deux sont partagés avec le continent. À cet égard, la découverte d’une nouvelle espèce de chrysopemorphologiquement distincte appartenant au genre Chrysopa, Chrysopa niki n. sp., est remarquable. Tandis que la plupart des espèces de Chrysopa se distinguent par des caractères morphologiques relativement subtils, le nouveau taxon se differencie de tous ses congénères du Paléarctique occidental grâce à une combinaison unique de caractères de coloration, des nervures et des genitalia. Les caractéristiques de C. niki n. sp. suggèrent qu’il s’agit d’une espèce endémique de Chypre, non strictement affine d’autres congénères, soulignant l’importance du nouveau taxon. La biogéographie des névroptères chypriotes est discutée en fonction de cette découverte.
http://www.zoobank.org/urn:lsid:zoobank.org:act:C6E3397B-19BA-431A-8F3A-4B872A72CED1
Keywords: Neuropterida; green lacewings; Mediterranean; biogeography; endemics; new species
Mediterranean islands are biodiversity hotspots (Myers et al. 2000) and among them Cyprus, the third largest, had a long geologic history with important biogeographic implications. Cyprus originated from the uplift of the sea bed during the Late Cretaceous but reached its actual shape and extension only in the Late Pliocene– Pleistocene (Robertson 1977; Hadjisterkotis et al. 2000; Kinnaird et al. 2011). A direct connection between Cyprus and the mainland is debated (see Hadjisterkotis et al. 2000), although geological data suggest a union with the Asian coast during the Late Messinian (Jolivet et al. 2006; Bache et al. 2012). Probably the first wave of colonizers reached the island at that time, while during the Pliocene and Pleistocene Cyprus was not connected to the mainland; this history may help explain the relatively high number of endemic species (Poulakakis et al. 2013).
The Neuropterida of Cyprus comprise 54 species, belonging to the orders Raphidioptera (2 species) and Neuroptera (52 species), only two of which are endemic to the island: the chrysopid Pseudomallada cyprinus
(Navás, 1932) and the myrmeleontid Delfimeus limassolicus (Navás, 1931) (Aspöck & Hölzel 1996; Aspöck et al. 2001; Kral & Devetak 2016). Chrysopidae, or green lacewings, are the most diverse family in Cyprus, with 17 reported species, four of which are in the genus Chrysopa Leach in Brewster, 1815, i.e. C. dubitans McLachlan, 1887, C. pallens (Rambur, 1838), C. viridana Schneider, 1845 and C. walkeri McLachlan, 1893 (Tjeder 1958; Aspöck & Hölzel 1996; Hölzel & Ohm 1999; Canard 2007; Kral & Devetak 2016).
The genus Chrysopa has a largely Holarctic distribution, including more than 60 described species, although the affinities of some non-Holarctic species need further investigations as this taxon was once a “waste basket” including most species of Chrysopinae (Brooks & Barnard 1990). Phylogenomic analyses corroborate morphological clues, strongly supporting a close relationship among Chrysopa and the smaller genera Plesiochrysa Adams, 1982 and Ceratochrysa Tjeder, 1966, suggesting that they might be merged within a broader Chrysopa (Tauber et al. 2001; Winterton et al. 2019). On
*Corresponding author. Email: [email protected]
© 2020 Société entomologique de France
Published online 18 May 2020
morphological grounds, the species of Chrysopa are often recognizable at a first glance due to their head and thor- acic pattern, although the genus monophyly relies on genital and larval characters (Principi 1940–1941; Brooks & Barnard 1990; Tsukaguchi 1995; Penny & Tauber 2000). The adults of Chrysopa are characterized by unusual traits for green lacewings, being predatory (in contrast to most green lacewings, which feed on nectar and pollen) and secreting skatole from the prothoracic glands for defense (Principi 1947, 1954; Canard et al. 1984; Canard 2001; Aldrich & Zhang 2016). The larvae of Chrysopa are mostly “naked” (i.e. not camouflaging with debris) with a few exceptions, such as the North American C. slossonae Banks, 1924 (Principi 1947, 1954; Eisner et al. 1978; Tsukaguchi 1995; Penny & Tauber 2000; Monserrat & Díaz-Aranda 2012; Tauber et al. 2014).
The Western Palaearctic contains 25 species (taking into account the recently revised status of some taxa), including some of the most common European chrysopids, and these species are mostly widespread in the region, with rare exceptions such as the Iberian endemic C. nierembergi Navás, 1908 (Aspöck et al. 1980, 2001). The European species of this genus are relatively well known, while a few Middle Eastern species belonging to the C. formosa Brauer, 1851 and C. phyllochroma Wesmael, 1841 species- groups are in need of revision (Aspöck et al. 2001). Most European Chrysopa species were described in the nine- teenth and in the first decades of the twentieth century, while several Iranian species were named in the 1960s (Hölzel 1967); the most recent described species are C. nigrescens Hölzel & Ohm, 1983 and the recently re-eval- uated C. gibeauxi Leraut, 1992. Given this information, the finding of a new Chrysopa species from Cyprus is note- worthy and unexpected. The aim of the present paper is to describe the morphology of this new species and compare it with other Western Palaearctic congeners.
Materials and methods
Specimens were examined and measured with a Leica® (Wetzlar, Germany) MZ 9.5 stereomicroscope equipped with an optical micrometer. Photographs were taken with a Canon® (Tokyo, Japan) EOS 600D digital camera equipped with Canon® (Tokyo, Japan) lens MP-E 65 mm; the resulting images were then processed and focus-stacked with the software Zerene® (Richland, WA, USA) Stacker. The color and pattern of chrysopids quickly fade after death; therefore, photos of live specimens are provided to show the habitus of the new species in life. Measurements were taken following the protocol of Tauber (2010): vertex width (V), pronotum length (PrL), pronotum width (PrW), forewing length (FL), forewing width (FW), hind wing length (HL), hind wing width (HW); body length (BL) was taken from vertex to abdomen tip. Genitalia were cleared in a solution of 10% KOH (potassium hydroxide) at room temperature and later rinsed in a solution of glacial acetic acid and water. Finally, they were stained in a solution of
chlorazol black and alcohol to enhance the contrast of the sclerites. The specimens and their genitalia were preserved in alcohol. Terminology for genitalia follows the interpretation of the homology of genital sclerites of Aspöck & Aspöck (2008); however, traditional terminology, i.e. following Adams & Penny (1985) and Tauber (2010), is given in square brackets to ease comparisons with previous literature. Wing venation largely follows Breitkreuz et al. (2017), also taking account of recent updates (Tauber 2019).
Type specimens were deposited in the following collections:
MSNG Museo Civico di Storia Naturale “G. Doria”, Genoa, Italy
MZUR Museum of Zoology, Sapienza Università di Roma, Rome, Italy
DBRC Davide Badano Research Collection, Taggia (IM), Italy
Taxonomy
Tribe Chrysopini Schneider, 1851 Genus Chrysopa Leach in Brewster, 1815
Diagnosis Male genitalia without gonocoxites 11 [tignum]; gonapophyses 9 [entoprocessus] with gonocornua; dorsal apodeme of the ectoproct curved upward; sternite 8 and 9 not fused; ventral apodeme of sternite 9 protruding apically; apex of sternite 9 with gonosetae.
Chrysopa niki n. sp.
Paratypes: 1, 1, CYPRUS/Koilani, Agia Mavri, 722 m/34°5043.56N 32°528.56E/2.VIII.2018/C. Makris leg.//PARATYPUS/Chrysopa niki Badano & Makris, sp. nov./D. Badano det. 2020. Alcohol preserved (MZUR). 1, 1, CYPRUS/Koilani, Agia Mavri, 722 m/ 34°5043.56N 32°528.56E/2.VIII.2018/C. Makris leg.// PARATYPUS/Chrysopa niki Badano & Makris, sp. nov./ D. Badano det. 2020. Alcohol preserved (MSNG). 2, 2, CYPRUS/Koilani, Agia Mavri, 722 m/34°5043.56N 32°528.56E/2.VIII.2018/C. Makris leg.//PARATYPUS/ Chrysopa niki Badano & Makris, sp. nov./D. Badano det. 2020. Alcohol preserved (DBRC). 1, 2 (of which one dissected), CYPRUS/near Pera Pedi village and Pano Platres, 920 m/34°5043.56N 32°528.56E/4. VIII.2018, C. Makris leg.//PARATYPUS/Chrysopa niki Badano & Makris, sp. nov./D. Badano det. 2020. Alcohol preserved (DBRC). 1, Pafos Panagia, 800 m/
2 D. Badano & C. Makris
6.VI.2010, light trap/C. Makris leg.//Pafos Panagia, 800 m; 6.VI.2010; light trap; C. Makris leg.; PARATYPUS/Chrysopa niki Badano & Makris, sp. nov./D. Badano det. 2020. Dry pinned (DBRC).
All specimens were collected in Mediterranean mid- mountain biotopes in the western part of the island.
Diagnosis Relatively small-sized Chrysopa with boldly marked body (Figure 1). Head with reddish brown x-shaped interantennal marking; vertex yellow with a pair of reddish brown stripes; gena and labrum with black stripe (Figure 2). Thorax with a pair of transverse dark brown bands (Figure 2). Pretarsal claws with
basal trapezoidal dilatation. Wing veins largely dark brown.
Description
Measurements (in mm). Males (6 specimens measured): body length mean 7.73 (min–max 5.87–9.37); vertex width 0.68 (0.50–0.85); pronotum length 0.39 (0.35–0.40); pronotum width 0.92 (0.75–1.00); forewing length 9.82 (7.75– 10.62); forewing width 3.64 (3.12–4.00); hind wing length 8.69 (7.25–9.75); hind wing width 3.21 (2.75–3.75). Females (5 specimens measured): body length mean 9.25 (min–max 8.37–10); vertex 0.75 (0.75–0.75); pronotum length 0.46 (0.40–0.50); pronotum width 1.01 (0.85– 1.10); forewing length 11.42 (10.75–12.25); forewing
Figure 1. Chrysopa niki n. sp., live specimens, habitus. A, (Cyprus, Kalo Chorio, 17.VI.2016, not collected). B, paratype, pale morph (Koilani, Agia Mavri). C, paratype (Koilani, Agia Mavri). All photos by C. Makris.
Annales de la Société entomologique de France (N.S.) 3
width 4.07 (3.75–4.25); hind wing length 10.52 (9.75– 11.12); hind wing width 3.61 (3.50–3.75).
Head. Base coloration pale green with prominent black and reddish markings (Figures 2, 3, 4A). Vertex slightly raised, smooth, without setae (Figure 4A). Frons smooth, shiny throughout. Clypeus flat, unsculptured. Labrum flat, surface smooth; distal margin straight. Vertex yellow with two paired, faded reddish stripes. Frons creamy with a reddish brown, x-shaped marking darker between the antennae, surrounding the anterior and posterior margins of the toruli and bordering the anterior margin of the vertex. Gena and clypeus marked with a lateral dark black stripe reaching the compound eye. Labrum brown.
Torulus creamy white. Scape creamy, with a small dark marking along the internal side. Pedicel creamy. Flagellum pale brown. Maxillary palp brown externally. Labial palp pale (Figure 4A).
Thorax. Base coloration pale yellowish green with dorsal paired dark bands (Figure 2). Pro-, meso- and metanotum with a pair of lateral dark brown stripes running for their entire length (Figures 3 and 4B). Pronotum with short, brown setae. Mesoscutellum with sparse brown setae. Pleurae pale green. Legs pale, unmarked, covered with short black setae. Pretarsal claws curved, base with trapezoidal dilatation (Figure 4C).
Figure 2. Chrysopa niki n. sp., live specimens, details of head and thorax. A, paratype, pale morph (Koilani, Agia Mavri). B, paratype (Koilani, Agia Mavri). All photos by C. Makris.
Wings. Membrane hyaline, stigmas slightly opaque (Figure 4D). Forewing rounded apically. RP relatively straight; first crossvein between RP and MA almost touching the apex of the first intermedial cell; first intermedial cell triangular; inner gradate series touching pseudomedia (i.e. amalgamation of radial and medial veins), with 2–3 crossveins; outer gradate series with 5–6 crossveins. Costal margin, Sc, RA, MP pale; all other longitudinal veins, costal veinlets and crossveins dark brown. Hind wing rounded apically. Inner gradate series with 2 crossveins. Outer gradate series with 5–6 crossveins. Costal margin, Sc and RA pale; all the other longitudinal veins and crossveins pale brown (Figure 4D).
Abdomen. Base coloration pale green. Tergites green with faint paired lateral brown markings, ventral margin slightly tinged with brown. Sternites green, with slightly brown dorsal margins (especially on the anterior seg- ments). Abdomen covered with short black setae.
Male genitalia. Ectoproct relatively short, apically rounded (Figure 5A). Sternite 9 with ventral apodeme extending dorsally in a sub-apical projection with rounded apex (Figure 5A); gonocoxites 9 [gonarcus] arch-like in dorsal view, with a pair of lateral distal projections and a pair of median distal subtriangular projections; gonapophyses 9 [entoprocessus] with distal horn-like projections and with ventral wing-like projections converging medially (Figure 5B–F). Gonocoxite 10 [pseudopenis] hook-like, gently tapered
apically (Figure 5B–F). Gonosaccus ventral to the genital sclerites, laterally bearing clusters of gonosetae rising from elevated bases (Figure 5B, C).
Female genitalia. Genitalia as for genus (Figure 6).
Variability. In some specimens the dark pattern of the body is brown instead of blackish (Figures 1B and 2A), thus the new species has distinct dark and pale morphs.
Etymology The new species is named after Niki Makri, the daughter of one of the authors, C. Makris. The specific name is a noun in apposition.
Comparative notes Chrysopa niki n. sp. is unmistakable among western Palaearctic chrysopids because of its combination of pale green body color, reddish brown x-shaped interantennal marking and vertex with reddish-brown stripes and thorax with transversal dark bands (Figure 1). Several Western Palaearctic Chrysopa species, such as C. perla (Linnaeus, 1758), C. walkeri, C. dorsalis Burmeister, 1839, C. nigrescens, C. hungarica Klapálek, 1899 and C. fuscostigma Esben-Petersen, 1933 have broad x-shaped black interantennal markings (Figure 7), while they are narrow and reddish-brown in the new species. Moreover, all these species usually have a black or darkened pedicel (Figure 7), in contrast to the entirely green antenna of C. niki n. sp. (Figure 2). These Chrysopa species also share largely black abdominal
Figure 3. Chrysopa niki n. sp., holotype with labels (abdomen and genitalia in Figure 5A) (Koilani, Agia Mavri). Scale bar: 1 mm.
tergites and sternites, while the abdomen is green (and without bands) in C. niki n. sp. In contrast to the above species with dark markings, all the remaining Western Palaearctic Chrysopa species (see Aspöck et al. 2001 for a complete list) have just a small dot between the antennae and inconspicuous head and body markings (see Monserrat 2016: figs 98, 102, 103, 105) or are completely unmarked (see Monserrat 2016: figs 98, 106). C. niki n. sp., the totally green C. viridana and the sparsely spotted C. sogdianica McLachlan in Fedchenko, 1875 are the only Western Palaearctic Chrysopa species with fewer
crossveins in the forewing inner gradate than in the outer series. Finally, at a first glance, C. niki n. sp. might resemble the small-sized, brownish and strongly patterned species of the genera Pseudomallada Tsukaguchi, 1995 and Suarius Navás, 1914, but it is set apart by the venational and genital characters of Chrysopa.
Life history Chrysopa niki n. sp. was mainly collected on Mediterranean oak trees, although it was also found
Figure 4. Chrysopa niki n. sp., details of head and thorax. A, paratype (Koilani, Agia Mavri), head in frontal view. B, paratype (Koilani, Agia Mavri), head and thorax, dorsal view. C, paratype (Koilani, Agia Mavri), pretarsal claw. D, paratype (Koilani, Agia Mavri), wings. Scale bars: 1 mm. Abbreviations: C = costa; Sc = subcosta; RA = radius anterior; RP = radius posterior; MA = media anterior; MP = media posterior; CuA = cubitus anterior; CuP = cubitus posterior; A = anal field; ig = inner gradates; og = outer gradates; psm = pseudomedia; psc = pseudocubitus; im = intramedian cell.
on conifers. In the site of Agia Mavri (34°5043.56N 32°528.56E), located at 722 m asl, specimens of the new species were found on Quercus infectoria veneris (A.Kern.) Meikle heavily infested with aphids, near a river valley with a riparial vegetation of Alnus orientalis Decne., Platanus orientalis L., Salix alba
L. and Rubus sanctus Schreb. In another locality, near Pera Pedi (34°5231.25N 32°520.91E, 722 m asl), C. niki n. sp. was collected on the endemic Quercus alnifolia Poech in a pine forest with Pinus brutia Ten., Quercus alnifolia, Arbutus andrachne L. and Cistus creticus L. Only one specimen was found on
Figure 5. Chrysopa niki n. sp., male abdomen and genitalia. A, Holotype (Koilani, Agia Mavri), abdomen with partly everted genitalia in lateral view. B–D, Paratype (Koilani, Agia Mavri), genitalia in B, lateral, C, ventrolateral and D, ventral views. E, F, Genitalia, schematic drawing. Scale bars, A: 0.5 mm, B–F: 0.2 mm. Abbreviations: ect = ectoproct; dap = dorsal apodeme; gs = genital sclerites; vap = ventral apodeme; gp9 = gonapophyses 9; gx9 = gonocoxites 9; gx10 = gonocoxites 10; gnsc = gonosaccus; gnst = gonosetae; T = tergite; S = sternite; sp = spiracle.
Pinus brutia at Agia Paraskevi locality (34°4959.07N 32°5929.84E, 570 m asl) in a pine forest mixed with broadleaves such as Quercus alnifolia, Quercus coccifera subsp. calliprinos (Webb) Holmboe, Arbutus andrachne, Myrtus communis L., Genista fasselata Decne. and Cistus creticus.
The gut of all dissected specimens contained uniden- tifiable insect remains, implying that the adults of C. niki are predaceous like other congeners.
Conclusions
During the Pliocene and Pleistocene, Cyprus was not connected to the mainland and colonization was only possible by oversea dispersal, explaining the relatively high number of endemics (Poulakakis et al. 2013). In the case of Neuropterida, the Cypriot fauna appears domi- nated by W-Palearctic elements, usually widely distribu- ted in the Euro-Mediterranean area, as in case of all
Coniopterygidae (3 species), Hemerobiidae (4 species) and most species of Chrysopidae and Myrmeleontidae. (Aspöck & Hölzel 1996; Aspöck et al. 2001). Several species are Eastern Mediterranean faunal elements, including the only species of Berothidae and Mantispidae present on the island, i.e. Isoscelipteron ful- vum Costa, 1863 and Mantispa scabricollis McLachlan in Fedchenko, 1875, one chrysopid, Suarius nanus (McLachlan, 1893), and some Myrmeleontidae and Ascalaphidae, such as Nemoleon poecilopterus (Stein, 1863), Neuroleon assimilis (Navás, 1914), Libelloides rhomboideus (Schneider, 1845), and Deleproctophylla variegata (Klug in Ehrenberg, 1834) (Aspöck et al. 2001; Kral & Devetak 2016; Badano et al. 2018). Myrmeleontiformia are the only group of Cypriot lacewings comprising Middle Eastern elements such as Dielocroce baudii (Griffini, 1895) (Nemopteridae), Distoleon laticollis Navás, 1913, Solter gaudryi Navás, 1914, Cueta kasyi Hölzel, 1969 and Libelloides jungei Aistleitner, 1982 (Myrmeleontidae and Ascalaphidae) (Aspöck et al. 2001; Kral & Devetak 2016; Badano et al. 2018). In contrast to Neuroptera, Raphidioptera are characterized by a low dispersal ability, and they show a high tendency to form endemic species (Aspöck et al. 1991; Aspöck 1998). Nevertheless, the only two Cypriot snakeflies, i.e. Phaeostigma cypricum (Hagen, 1867) and Ulrike syriaca (Steinmann, 1964), are both present in the Middle East (Aspöck et al. 1991, 2001). Therefore, the Neuropterida species of Cyprus suggest a series of faunal exchanges with the mainland. Only two lacewing species are considered as endemic to Cyprus: Pseudomallada cyprinus and Delfimeus limassolicus, although both belong to taxonomically entangled genera and their status needs confirmation by further studies in the Middle East. C. niki n. sp. joins this small group of apparently endemic Cypriot lacewings, in that further research along the Mediterranean coasts of Western Asia are necessary to verify if C. niki n. sp. is actually limited to the island. If confirmed, it would be a remarkable species from a
Figure 6. Chrysopa niki n. sp., paratype (Koilani, Agia Mavri), abdomen, lateral view. Scale bars: 0.5 mm. Abbreviations: ect = ectoproct; gx8 = gonocoxites 8; gx9 = gonocoxites 9; T = tergite; S = sternite; sp = spiracle.
Figure 7. Western Palearctic Chrysopa species with x-shaped interantennal marking. A, C. perla (Linnaeus). B, C. walkeri McLachlan. C, C. dorsalis Burmeister. D, C. fuscostigma Esben-Petersen. Photos A–C by P. Duelli, D by D. Devetak.
biogeographic point of view taking into account the wide distribution of most Western Palaearctic Chrysopa species.
Acknowledgements
We wish to thank Spyros Sfenthourakis (University of Cyprus, Cyprus) for the critical review of the manuscript. Great thanks also to Dusan Devetak (Univerity of Maribor, Slovenia) and Peter Duelli (Swiss Federal Research Institute WSL, Switzerland) for sharing their photos of Chrysopa species. We would like to thank also the editors and the anonymous referees for their valuable comments on the manuscript.
ORCID Davide Badano http://orcid.org/0000-0001-9715-3107
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a new unexpected species of chrysopa leach from cyprus

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