Abstract
Nemoura kozari sp. n. is described on the basis of morphology of male adults collected in the Eastern Carpathians. The new species is classified as a member of the Nemoura marginata species group. It was found in medium elevations of the Krasna Mts, Ukraine, and the Rodna Mts, Romania, inhabiting slowly flowing, small brooks. New diagnosis of the marginata group and enumeration of its members are provided.
Introduction
The genus Nemoura Latreille, 1796 is a large and clearly paraphyletic assemblage of euholognathan stoneflies, distributed in the Holarctic and Oriental realms (Gamboa et al., 2019; Mo et al., 2020). Most West Palaearctic Nemoura belongs to the sensu stricto lineage, consisting of closely related morphological species groups and isolated species as well (Murányi, 2007; Mo et al., 2020). The species diversity is relatively evenly distributed within the subregions of the West Palaearctic, though most narrow endemics are restricted to mountainous regions (Graf et al., 2009; DeWalt et al., 2023). In regards of the Carpathians, the occurrence of 19 species is confirmed (Kis, 1974; Žiak, 2016). Four of these are Carpathian endemics and three are subendemics (Kis, 1974).
More than twenty years ago, a single male of an unknown Nemoura was found along a small open brook of the Krasna Mts, a medium high range of the Ukrainian Eastern Carpathians. Unfortunately, the epiproct of the specimen was lost during mounting, and no subsequent trip was taken to its hardly accessible habitat. Six years later, a second male specimen was found along a small forest brook at medium elevations of the Rodna Mts, Romania, about hundred kilometres far from the Krasna Mts. Collecting in various other localities of different elevations of the Rodna Mts yielded no further specimens. Due to the very limited material and the taxonomic problems around the Nemoura marginata species group, to which this new species belongs, we postponed its description. Recently, new species description and redescription of some marginata group members clarified morphological traits and limits within this complicated lineage (Vinçon and Ruffoni, 2021). Thus, herein we describe this East Carpathian Nemoura and give a new diagnosis of the marginata group, together with an enumeration of its members.
Material and methods
Specimens were collected by hand or beating sheet and stored in 75% ethanol. All materials were deposited in the Hungarian Natural History Museum, Budapest, Hungary (HNHM).
Figure 1 was made by a drawing tube mounted on a Nikon SMZ1500 microscope, and on the basis of photos taken by different microscopes. Figure 2 was made with a Keyence VHX 5000 digital microscope. To avoid destruction of the holotype male, terminalia was cleared in 10% KOH solution but the epiproct was not slide mounted.
Male terminalia of Nemoura kozari sp. n. — A: terminalia, ventral view; B: same, lateral view; C: same, dorsal view; D: epiproct, left side dorsal, right side ventral view; E: apical sclerite of epiproct, caudal view — Abbreviations: Sternum 9: hy (hypoproct), ve (vesicle); Paraproct: il (inner lobe), ol (outer lobe); Cercus: ah (apical hook), vs (vestigial second segment); Tergum 9: 9h (strong hairs); Tergum 10: 10s (medial spot); Epiproct: as (apical sclerite), bc (basal cushion), bp (spines of basal plate), la (lateral arm), lk (lateral knob), ri (ring) — scale 0.5 mm for Figs. A–C, 0.1 mm for Figs D–E
Citation: Acta Phytopathologica et Entomologica Hungarica 58, 1; 10.1556/038.2023.00177
Male morphology of Nemoura kozari sp. n. — A: head and thorax, dorsal view; B: terminalia, dorsocaudal view; C: same, dorsolateral view — scale 1 mm for Fig. A, 0.5 mm for Figs B–C
Citation: Acta Phytopathologica et Entomologica Hungarica 58, 1; 10.1556/038.2023.00177
Morphological terminology follows our previous Nemouridae papers (Murányi, 2007; Mo et al., 2020).
Results and discussion
Nemoura kozari sp. n. (Figs 1–3)
Habitat and distribution of Nemoura kozari sp. n. — A: habitat of the holotype, downstream view; B: same, upstream view; C: distribution in the Carpathian Basin (red ring: locality of the holotype; red dot: locality of the paratype)
Citation: Acta Phytopathologica et Entomologica Hungarica 58, 1; 10.1556/038.2023.00177
Type material: Holotype male: ROMANIA, Maramureş county, Rodna Mts, Săcel, forest brook above the Iza Spring, 1155 m a.s.l., N 47°34.830′ E 24°32.108′, leg. J. Béres, Cs. Csuzdi, J. Kontschán, D. Murányi, V. V. Pop, 20.v.2008 (HNHM: PLP2391). Paratype: UKRAINE, Zakarpatska region, Tyachivskyi district, Krasna Mts, small, open spring and its outlet in the upper valley of Luzanka River, 1350 m a.s.l., N 48°22.971′ E 23°45.039′, leg. K. Balogh, B. Cser, D. Murányi, 19.v.2002: 1 male (HNHM: PLP926; epiproct lost).
Diagnosis. Small sized species of the Nemoura marginata group. Male: paraproct outer lobe with elongated, slightly bent tip; cercus distinctly bent and hairy, with large and erect apical hook; epiproct with short and strong ring, apically widened lateral arms, large and plate-like apical sclerite that is rectangular in caudal view.
Description. Small sized species, macropterous. Forewing length: holotype male 5.2 mm, paratype male 5.8 mm. General colour brown, pilosity short and indistinct (Fig. 2A). Head brown with indistinct pattern, mouthparts and palpi yellowish, antennae light brown (Fig. 2A). Pronotum rectangular with rounded corners, wider than long, pale brown with indistinct rugosities (Fig. 2A). Meso- and metanotum brown, ventral aspect of thorax brown with whitish membranous portions (Fig. 2A). Legs pale brown, but apex of femora, basal half of tibiae and all tarsi darker. Wings hyaline, venation brown. Abdominal segments pale but terminalia brown to dark brown.
Male abdomen: Sternum 8 membranous with small, paired anterolateral sclerites. Hypoproct rounded, longer than wide, apical part very small and acute (Fig. 1A–B); vesicle spoon shaped, twice longer than wide (Fig. 1A–B). Paraproct inner lobe thin, half as long as outer lobe (Fig. 1A); the outer lobe is with convex inner margin and strongly curved outer margin, tip is elongated with apex blunt and slightly bent medially (Fig. 1A–B). Cercus long and with large apical hook (Fig. 1A–C, 2B–C); laterally well sclerotized, membranous in its inner portion and to the vestigial second segment; bent in lateral view (Fig. 1B), distinctly bent medially in dorsal and ventral views (Figs 1A–1C); apex reminds a chicken-head, with strong, nearly erect hook, and rounded, lightly sclerotized inner portion (Figs 1C, 2C); the whole cercus bears distinct, long hairs (Figs 1A, 1C, 2B). Terga 8–9 mostly membranous but with strong antecosta and light brown anterior and lateral sclerotized portions; tergum 9 with posteromedial row of 5–6 strong hairs (Figs 1C, 2B–C). Tergum 10 wide and strongly sclerotized, with a medial light spot under the tip of epiproct, lacking projections and distinct hairs (Figs 1C, 2B–C). Epiproct elongated elliptical, with large basal cushion, with short and strong ring and large, plate-like apical sclerite (Fig. 1B–E, 2B–C). Dorsal sclerite with large lateral arms, apically widened in half-moon shape (Fig. 1C–D–D, 2B). Ventral sclerite basally with pair of small lateral knobs, after the moderately broad base continued in a thin, elongated basal plate that bears rows of 7–8 small spines (Fig. 1D); ring of the ventral sclerite short but strong, regularly curved (Fig. 1C–D, 2B–C); apical sclerite erect, large and plate-like (Figs 1B, 1D–E), bears three or four very small lateral spines, rectangular in caudal view (Fig. 1E).
Female and larva: unknown.
Affinities. The new species can be assigned to the Nemoura marginata species group, according to the diagnosis of the group as discussed below. The nominal N. marginata Pictet (1836) can be easily distinguished from the new species on the basis of triangular paraproct outer lobe, straight cercus in lateral view, and much lower and wider apical sclerite of epiproct in caudal view (see Fig. 1k–m in Zwick, 1970; Figs 83A–D in Kis, 1974; Fig. 3 in Ravizza and Ravizza Dematteis, 1995; Figs on page 343 in Roesti, 2021). The more morphologically similar Carpathian species are Nemoura flexuosa Aubert (1949) and Nemoura hamata Kis, 1965. Nemoura flexuosa can be distinguished on the basis of less elongated tip of the paraproct outer lobe, stout cercus that is straight in lateral view, and the plate-like apical sclerite of epiproct that is rounded instead of rectangular (see Figs 1a–c in Zwick, 1970; Figs 85A–D in Kis, 1974; Fig. 2 in Ravizza and Ravizza Dematteis, 1995; Figs on page 341 in Roesti, 2021). Nemoura hamata differs by the triangular paraproct outer lobe, the large apical hook of cercus bent downwards, and the apical sclerite of epiproct is bilobed (see Figs 2–5 in Kis, 1965; Figs 91A–D in Kis, 1974).
Ecology and distribution. The species was found in the medium elevations of the Krasna Mts, Ukraine (paratype male), and the Rodna Mts, Romania (holotype male) (Fig. 3C). Both specimens were caught in the second half of May, along relatively slowly running, small brooks. The habitat in the Rodna Mts is a montane brook that flows in spruce forest, with dense riparian vegetation dominated by Petasites hybridus along sunny patches where spruce allows it (Figs 3A–B). The brook has a sandy-gravely substrate and steep bank, with spruce twigs traversing. Besides the holotype, one male and three females of Leuctra dalmoni Vinçon and Murányi (2007) were caught, but the stream was not checked for larvae. The habitat in the Krasna Mts is similar in size, substrate and bank, but located in grassland on a steep slope, just above the forested gorge of a large montane stream. The spring and most of the investigated section of the brook is over hanged by tussocks of Nardus stricta. The paratype was the only adult stonefly collected, while pharate, ultimate and penultimate larvae and one exuviae of Nemoura fusca Kis, 1963 were collected from and nearby the water.
Etymology. The species is dedicated to our late colleague, Dr. Ferenc Kozár (1943–2013), renowned world specialist of scale insects. Used as the genitive of a noun of male gender.
Delimitation and composition of the Nemoura marginata species group
The marginata group was first defined on the basis of the cercus armed with a single apical tooth by Aubert (1946). The assemblage he listed included many European species, some that later were transferred to subsequently defined morphological groups. Since then, some of the species closely related to N. marginata were treated as the Nemoura flexuosa-marginata complex (e.g. Ravizza and Ravizza Dematteis, 1995), or briefly discussed as the marginata group (e.g. Zhiltzova, 2003; Murányi, 2007), but the group was not precisely defined. In the genus level revision of the Nemouridae, Baumann (1975) distinguished two species complexes within Nemoura but not treated the marginata group as a taxonomical or operational unit. Recently, Mo et al. (2020) pointed out, that the limits of Nemoura sensu stricto is hard to define morphologically, the paraphyletic Nemoura sensu lato requires erection of further genera, and that species groups within Nemoura are generally hard to define because of intermediate morphology of several species. However, Nemoura presently includes nearly 200 valid species (DeWalt et al., 2023), and species grouping is necessary for operational reasons, even if some of the groups will prove to be not true evolutionary lineages by future molecular studies. Thus, we propose to distinguish the marginata group on the basis of the following morphological characters: cercus with a single, distinct apical hook on the outer lateral portion (vestigial tooth may present on the inner lateral surface); ring of the epiproct's ventral sclerite short but strong, lacks apical lobes; apical sclerite of the epiproct erect in lateral view, at least as wide as the sclerite that is forming the ring, and always bear small, lateral spines. A distinct set of closely related species are distinguished as the palliventris subgroup, on the basis of presence of a plate-like expansion on the epiproct's apical sclerite.
The members of the marginata group of the present sense are listed, with their distribution and available descriptions, below:
Nemoura apollo Zwick (1978): Greece; known only from the original description.
Nemoura braaschi Joost (1970): Bulgaria; known only from the original description.
Nemoura caspica Aubert (1964): Iran; known only from the original description.
Nemoura ceciliae Aubert (1956b): Spain and Portugal; complementary described by Tierno de Figueroa et al. (2003).
Nemoura confusa Zwick (1970): France; complementary described by Ravizza and Ravizza Dematteis (1995).
Nemoura erratica Claassen (1936): Western Europe and Pyrenees, but not in the Alps; redescribed by Vinçon and Pardo (2003).
Nemoura flexuosa Aubert (1949): most of Europe but not in Iberia and the British Isles, extends to Anatolia; redescription by Zwick (1970), complementary descriptions e.g. in Kis (1974), Lillehammer (1974), Ravizza and Ravizza Dematteis (1995), Roesti (2021).
Nemoura hamata Kis, 1965: Romania; complementary described in Kis (1974).
Nemoura marginata Pictet (1836): Central Europe and the Balkans; redescription by Zwick (1970), complementary descriptions e.g. in Kis (1974), Ravizza and Ravizza Dematteis (1995), Roesti (2021).
Nemoura minima Aubert (1946): subendemic to the Alps, spreads to the western Balkans; complementary described e.g. in Kis (1974), Roesti (2021).
Nemoura oropensis Ravizza & Ravizza Dematteis (1980): Italy; known only from the original description.
Nemoura pesarinii Ravizza & Ravizza Dematteis (1979): Italy and Switzerland; complementary described by Roesti (2021).
Nemoura pseudoerratica Vinçon & Pardo (2003): France, Andorra and Spain; known only from the original description.
Nemoura pygmea Braasch & Joost (1972): Bulgaria; known only from the original description.
Nemoura rivorum Ravizza & Ravizza Dematteis (1995): Italy and France; known only from the original description.
Nemoura sabina Fochetti & Vinçon (2009): Italy; known only from the original description.
Nemoura undulata Ris (1902): Alps; redescribed by Aubert (1950), complementary described in Roesti (2021).
Nemoura wittmeri Zwick (1975): Turkey; known only from the original description.
palliventris subgroup:
Nemoura aprutiana Vinçon & Ruffoni (2021): Italy; known from its recent description.
Nemoura flaviscapa Aubert (1956a): Greece; known only from the original description.
Nemoura hesperiae Consiglio (1960): Italy; recently redescribed by Vinçon and Ruffoni (2021).
Nemoura lucana Nicolai & Fochetti (1991): Italy; recently redescribed by Vinçon and Ruffoni (2021).
Nemoura obtusa Ris (1902): Alps, presence in the Carpathians needs confirmation; complementary description e.g. in Kis (1974), Roesti (2021).
Nemoura palliventris Aubert (1953): Italy, France and Switzerland; recently redescribed by Vinçon and Ruffoni (2021) and Roesti (2021).
Nemoura vinconi Murányi (2007): Albania; known only from the original description.
Nemoura zwicki Sivec (1980): Kosovo; known only from the original description.
Acknowledgements
We are indebted to our friends and colleagues who took part in the field trips, and to Jenő Kontschán (Plant Protection Institute, Centre for Agricultural Research, Budapest, Hungary) for allowing to use the Keyence microscope in the PPI. We are also grateful to Wolfram Graf (Institute of Hydrobiology and Aquatic Ecosystem Management, University of Natural Resources and Life Sciences, Vienna, Austria) for his comments on the manuscript. RJG acknowledges the institutional support of the Institute of Entomology (Biology Centre of the Czech Academy of Sciences) RVO: 60077344.
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