Zootaxa 3786 (5): 541–556 www.mapress.com / zootaxa / Copyright © 2014 Magnolia Press

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3786.5.3 http://zoobank.org/urn:lsid:zoobank.org:pub:A4F2E880-D01E-4FBD-B439-2D2DABDCFCF4

Aquatic Empididae (Diptera: Hemerodromiinae and Clinocerinae) of the Sierra Nevada, Spain, with the description of five new species MARIJA IVKOVIĆ1,4, CARMEN ZAMORA-MUÑOZ2, MARTA SAINZ-BARIAÍN2 & BRADLEY J. SINCLAIR3 1

Department of Zoology, Division of Biology, Faculty of Science, University of Zagreb, Rooseveltov trg 6, 10000 Zagreb, Croatia. E-mail: [email protected] 2 Department of Zoology, University of Granada, Granada, Spain. E-mail: [email protected]; [email protected] 3 Canadian National Collection of Insects & Canadian Food Inspection Agency, Ottawa Plant Laboratory – Entomology, CFIA, Ottawa, Canada. E-mail: [email protected] 4 Corresponding author

Abstract In total 24 species of aquatic Empididae (Clinocerinae and Hemerodromiinae) are known from the Sierra Nevada Mountains of Spain, including five new species (Hemerodromia planti Ivković & Sinclair sp. nov., Kowarzia nevadensis Sinclair & Ivković sp. nov., Wiedemannia darioi Sinclair & Ivković sp. nov., W. horvati Ivković & Sinclair sp. nov. and W. vedranae Ivković & Sinclair sp. nov.). The new species are described, illustrated and distribution of all species listed. Wiedemannia (Philolutra) angelieri Vaillant and W. (Roederella) ouedorum Vaillant are newly recorded in Sierra Nevada and nine species are endemic to this region. A key to all 24 species of aquatic empidids is presented. Key words: Clinocerinae, Hemerodromiinae, Empididae, Diptera, new species, Sierra Nevada, Spain

Introduction The high mountain systems of Europe (e.g., Alps, Pyrenees) have greatly influenced the evolution of species during the cyclic climatic changes (Schmitt 2009). The mountains have acted both as barriers and refugia contributing to allopatric speciation of species endemic to these ranges. The Alps have been studied most extensively and shelter the highest number of endemics (Schmitt 2009). The highest number of endemic plants in Europe and also vast numbers of endemic invertebrates are confined to the Sierra Nevada range (Gonzalez-Megias et al. 2008). Although Europe has been relatively well studied in comparison to other parts of the world, some mountain ranges, including the Sierra Nevada remain poorly known with regard to aquatic Empididae (Diptera). The Sierra Nevada Mountains are situated in the southern region of Andalusia, Spain and have Mediterranean high mountain ecosystems that are mostly protected by a national park. This mountain range is less than 2000 km2, about 100 km in length and includes more than 20 peaks higher than 3000 m a.s.l. It is the highest mountain chain in the Iberian Peninsula and the second highest in Western Europe after the Alps. Its axis is orientated in a southwest to north-east direction, and the centre of the chain forms an elongated and depressed arch, constituting a series of rounded hillsides on southern slopes, and a clear glacial geomorphology with cirques and headwalls on the northern slopes. The peripheral zones of the Sierra Nevada, although of lower altitudes, present a radically different aspect with abrupt drops, steep slopes, and deep canyons (Martín et al. 2008). Due to its combination of alpine elevation and rather low latitude (at least in comparison to other European high mountains), the Sierra Nevada range is characterized by hard snowy winters and extremely warm summers with drought periods that result in some streams and rivers drying up and disappearing. Since 1986 the Sierra Nevada has been designated a Mountain Biosphere Reserve by UNESCO (MBRs) for studying the consequences of climate change (UNESCO 2014). Thirty-four streams and rivers were surveyed for aquatic Empididae (Clinocerinae and Hemerodromiinae) in

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the Sierra Nevada range from April to August, 2013. Here we describe one new species of Hemerodromia Meigen, one new species of Kowarzia Mik and three new species of Wiedemannia Zetterstedt discovered during this survey. In addition, we provide a detailed analysis of all publications on aquatic dance flies from this region and contribute with additional records resulting from research conducted during 2013 and material in the Canadian National Collection of Insects (CNC). This study builds on the recent Empididae summary (Ventura-Pérez 2013) that appeared in Ruano et al. (2013).

Material and methods This paper is based on a review of literature data and on unpublished data from our own research on the aquatic dance fly fauna in Sierra Nevada. The names of taxa presented in this checklist reflect current nomenclature and classifications (Sinclair 1995; Yang et al. 2007). A list of locality names including latitude, longitude, altitude, slope and number codes (site ID) that were studied in this research are presented in Table 1 and a map with sites is also provided (Fig. 1). Specimens were collected using sweep nets and by aspirator, preserved in ethanol (EtOH). For the purpose of determination male terminalia were dissected, macerated in hot 85% lactic acid and stored in ethanol along with the remaining body parts in the same tube. All specimens under material examined sections were collected by M. Ivković, unless otherwise stated.

FIGURE 1. Map of sampling sites of aquatic Empididae recorded from Sierra Nevada, Spain (See Table 1 for explanation of codes).

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Label data for primary types are cited from the top downward, with the data from each label in quotation marks. Labels are cited in full, with original spelling, punctuation, and date, and label lines are delimited by a slash (/). Additional information is included in square [ ] brackets. The repository of each type is given in parentheses. Secondary type data are abridged and listed alphabetically. This study is based on material from the following institutions: Canadian National Collection of Insects, Ottawa, Canada (CNC); col. M. Ivković, University of Zagreb, Croatia (UZC); University of Granada, Spain (UGS). Terms used for adult structures primarily follow those of McAlpine (1981) and Cumming & Wood (2009), except for the antenna and wing venation where the terms of Stuckenberg (1999) and Saigusa (2006) are used respectively. In the system outlined by Saigusa (2006), the dipteran wing vein A1 (as used in McAlpine 1981) is homologized with the mecopteran CuP, and consequently CuA1 (of McAlpine) is termed M4, whereas CuA2 is CuA, the anal cell is cell cua and the anal vein (A1+CuA2) is CuA+CuP. Homologies of the male terminalia follow those of Sinclair & Cumming (2006). Species of Wiedemannia described herein will not be assigned to subgenus because of the reasons mentioned in Ivković et al. (2012) as we consider current subgeneric concepts confused and mostly not monophyletic.

Results Taxonomy Hemerodromia planti Ivković & Sinclair, sp. nov. (Figs 2, 3) Type material. HOLOTYPE ♂ (in alcohol) labelled: “HOLOTYPE/ Hemerodromia/ planti IVKOVIĆ & SINCLAIR/ SPAIN, Sierra Nevada,/ Río Aguas Blancas,/ Área Recreativa,/ 37°13'42''N,/ 03°24'32''W, 29.v.2013, leg. M. Ivković” (UZC). PARATYPE: 1 ♂, Río Aguas Blancas, Fuente de los Trucheros, 37°13'25''N, 03°24'17''W, 1155 m, 29.v.2013 (1 ♂ UZC). Diagnosis. A yellow-brown species, characterized in the male by dark brown to black terminalia with oval cerci, with a small posterior, inwardly directed subapical pointed process. Description. Male. Body length 2 mm. Head dorsoventrally flattened, dark brown, all setae whitish, ocellar triangle black, antenna whitish. Eyes iridescent black, ommatidia conspicuously enlarged anteriorly; narrowly separated on face. Frons with 1–2 frontal setulae. Pair of fine vertical setae midway between anterior ocellus and eye margin, contiguous with 2–3 postocular setae. Occiput bearing scattered fine hairs; clypeus with rather dense short downwardly directed pile. Antenna with scape, pedicel and postpedicel bearing distinct short dorsal setulae; postpedicel about 1.5X as long as wide, stylus much shorter than postpedicel. Thorax elongated; yellow, all setae yellowish. Mesonotum with brown longitudinal stripe. Acrostichal setae

small and fine, biserial; 1−2 distinct intra-alar setae; 3−4 supra-alar setae.

Wing (length 2.4 mm) membrane faintly yellowish tinged, veins yellow; fork R4+5 less than 90°, slightly distal of fork M1+2; R5 and M1 convergent apically; second submarginal cell (r4) rather long, vein R5 about 3X as long as R4. Halter whitish yellow. Legs whitish yellow. Fore coxa about as long as distance between fore and mid coxae, 5X as long as wide with several pale dorsoapical setae. Fore femur subequal in length to fore coxa, 4X as long as wide, evenly inflated, widest at middle. Fore femur with the following chaetotaxy: 6 anteroventral denticles, 10 anteroventral spines, 6 posteroventral denticles, 10 posteroventral spines, 4 basal posteroventral spines; denticles black, becoming closer spaced, rows converging distally; anteroventral spines dark brown to black, becoming weaker and shorter distally; 2 basal posteroventral spines short and dark and 2 long and yellow. Fore tibia 0.8X as long as fore femur, evenly curved. Mid and hind legs slender, lacking strong setae. Abdomen yellow ventrally and brown dorsally with pale setae most conspicuous on hind margin of posterior sternites. Terminalia: Cercus (Fig. 3) black, somewhat narrow, concave in middle (in lateral view); apically incurved, pointed subapical process directed internally 0.5 from base also visible in lateral view, distinctly setose; anterior part of cercus pointed apically, with strong setae; apex of cercus expanded, nearly as wide as epandrium (Fig. 2). Epandrium dark brown to black, medially slightly convex, distally blunt-ended with posterior margin

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pronounced and bearing distinct setae; entire epandrium covered in numerous setae (Fig. 2). Hypandrium brown, rather oval, not reaching end of epandrium, with strong setae laterally; postgonite apical and sharply pointed, emergent above cercus in lateral view. Phallus whitish and broad. Female. Unknown. Remarks. Hemerodromia planti is known only from the Sierra Nevada Mountains, Spain. This species appears to be close to H. wagneri Cobo & Carreira, 2003 also described from Spain, but H. planti is distinguished from H. wagneri by the apically rounded epandrium and shorter anterior projection of the cercus. Etymology. The new species is named after Dr Adrian Plant, colleague and leading expert on Hemerodromiinae.

FIGURES 2–3. Hemerodromia planti Ivković & Sinclair, sp. nov., male terminalia. 2. Lateral view; 3. Cercus, dorsal view. Abbreviations: cerc—cercus, epand—epandrium, hypd—hypandrium, pgt—postgonite, ph—phallus. Scale bar = 0.2 mm.

Kowarzia nevadensis Sinclair & Ivković, sp. nov. (Figs 4, 6, 7) Type material. HOLOTYPE ♂ (dried material) labelled: “SPAIN: Sierra Nevada/ Barranco Frío, Hoya Carlos/ 1560 m, 17.iv.2013/ 36°56'47''N, 03°23'44''W/ leg. M. Ivković”; “HOLOTYPE/ Kowarzia/ nevadensis/ Sinclair & Ivković [red label]” [dissected] (CNC). Diagnosis. This species is characterized by the pale yellowish brown coxae and femora; straight, broad clasping cercus; apically forked surstylus; and distinctive apex of the phallus. Description. Male. Body length 3 mm (Fig. 4). Head dark brown, face with bright blue pruinescence. Face with 3−4 setulae along inner margin of each eye. Arista-like stylus long and slender, slightly shorter than height of eye. Thorax dark brown, lacking distinct dorsal vittae; pleurae with very pale blue pruinescence. Chaetotaxy long and slender: several short acrostichal setulae anterior to first dorsocentral setae; 5 dorsocentral setae; 1 postpronotal seta; 1 presutural supra-alar seta; 2 notopleural setae, lower seta more slender; 1 postalar seta; 1 pair of apical scutellar setae.

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FIGURES 4–5. Male habitus images, holotype. 4. Kowarzia nevadensis Sinclair & Ivković, sp. nov., terminalia removed. 5. Wiedemannia darioi Sinclair & Ivković, sp. nov. Scale bar = 1.0 mm.

Wing (length 3.6 mm) narrow, lightly infuscate. Pterostigma narrow, elongate, faint. Basal costal seta long, extending to humeral crossvein. Legs with coxae and femora yellowish brown; tibiae and tarsi dark brown; coxae with pale blue pruinescence on dorsal half. Fore femur with preapical anterior comb. Fore tibia with row of stout anterodorsal setae. Hind tibiae with erect dorsal and ventral stout setae on apical half; setae usually shorter than width of tibia. Hind tarsomere 1 with pair of erect basal setae and stout setae along ventral margin. Abdominal sclerites dark brown. Terminalia (Figs 6, 7): Hypandrium cone-shaped, with short setae. Phallus somewhat curved, with broadly rounded apex; distiphallus sinuous, whip-like; ejaculatory apodeme small, slender. Epandrium oval, subequal in length to hypandrium. Cercal plate with 4−5 strong setae and numerous slender setae confined to dorsolateral region. Clasping cercus straight, digitiform; apex somewhat produced into short beak-like tip; apical half of inner face with stout, peg-like setae (Fig. 7). Surstylus narrow, straight, similar in shape to clasping cercus, apex narrowed to long subterminal beak-like tip, apex produced into sharp crown; apex clothed in setae. Female. Unknown. Remarks. This new species of Kowarzia is known only from the Sierra Nevada mountains, distinguished from related species by features outlined in the key. Etymology. The new species is named after the Sierra Nevada mountains.

Wiedemannia darioi Sinclair & Ivković, sp. nov. (Figs 5, 8, 9) Type material. HOLOTYPE ♂ (dried material) labelled: “SPAIN: Sierra Nevada; Río/ Aguas Blancas, Cenes de la/ Vega, 760 m, 13.v.2013/ 37°09'55''N, 03°30'53''W/ leg. M. Ivković”; “HOLOTYPE/ Wiedemannia/ darioi/ Sinclair & Ivković [red label]” [dissected] (CNC). Diagnosis. This species is distinguished by yellowish brown legs and clasping cercus and a distinct erect preapical anterior seta on the fore femur. Description. Male. Body length 2.5 mm (Fig. 5). Head brown, in lateral view longer than wide; gena broad, less than half as deep as height of eye, with pale blue pruinescence. Frons short, broader than face. Face wide, with distinct carina on lower margin, silvery-grey pruinescent. Pair of ocellar setae slightly longer than postpronotal seta; 2 vertical setae and 3–5 distinct upper postoculars similarly sized and black, with numerous shorter, thinner NEW AQUATIC EMPIDIDAE

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setae. Lower postocular setae much finer and paler merging with pale, longer setae on middle and lower occiput. Few dark setulae present on vertex and between ocellar area and eye margin. Antenna dark brown; postpedicel and stylus minutely pubescent; pedicel >0.5X long as scape, with complete circlet of apical setae; postpedicel apically pointed, stylus 2.5X length of postpedicel.

FIGURES 6–9. Male terminalia, lateral view. 6. Kowarzia nevadensis Sinclair & Ivković, sp. nov. 7. K. nevadensis Sinclair & Ivković, sp. nov., clasping cercus, inner view 8. Wiedemannia darioi Sinclair & Ivković, sp. nov. 9. W. darioi Sinclair & Ivković, sp. nov., clasping cercus, inner view. Abbreviations: cerc pl—cercal plate, cl cerc—clasping cercus, epand—epandrium, hypd—hypandrium, ph—phallus, sur—surstylus. Scale bar = 0.1 mm.

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Scutum brown, iridescent (viewed anteriorly); median stripe extending onto scutellum (viewed posteriorly). Pleura with pale blue pruinescence. Mesonotum with 5 pairs of dorsocentral setae, without interspersed setae. Acrostichal setulae indistinct, fine, biserial, extending around prescutellar depression; 1 postpronotal seta, 2 notopleural setae; 1 presutural supra-alar seta and numerous scattered setulae. Antepronotum with pair of pale setae. Proepisternum with patch of fine pale setae. Laterotergite with fine pale setae. Pair of strong marginal scutellar setae and several setulae; disc with numerous short fine setulae. Wing (length 3.0 mm) membrane lightly infuscate, veins darker. Basal costal seta extending to humeral crossvein. Cell dm produced anteroapically. M1+2 petiolate, stem longer the m-m crossvein. Vein CuA+CuP absent. Pterostigma elongate, pale brown. Squama with pale fringes. Halter light brown. Legs mostly yellowish brown, tarsi somewhat darker, tarsomeres 4−5 dark brown. Coxae with long pale setae anteriorly. Legs uniformly clothed with rows of small dark setae. Fore femur with erect preapical anterior seta. Abdomen concolorous with thorax. Terminalia (Figs 8, 9): Hypandrium very small, cylindrical with pair of setae. Epandrium subquadrate, small, subequal in length to hypandrium; small spine-like surstylus emerging from inner face apically. Clasping cercus yellowish brown, broad, leaf-like, with thumb-like posterior lobe; oval anterior lobe with short tapered lobe near posterior lobe; slender setae mostly confined to outer margin; inner face with stout, wavy, round tipped setae on anteroventral region (Fig. 9). Cercal plate 3–4 long setae on knob-like projection. Phallus mostly straight with slight preapical bend near base of distiphallus; distiphallus slightly curved, slender in dorsal view. Female. Unknown. Etymology. The new species is named after the husband of the senior author, Dario Bjelkanović, for his immense support.

Wiedemannia horvati Ivković & Sinclair, sp. nov. (Fig. 10) Type material. HOLOTYPE ♂ (in alcohol) labelled: “HOLOTYPE/ Wiedemannia/ horvati IVKOVIĆ & SINCLAIR/ SPAIN, Sierra Nevada,/ Río Guarnón, Corral del/ Veleta, 37°03'39''N,/ 03°22'04''W,/ 19.viii.2013,/ leg. C. Zamora-Muñoz” (UZC). PARATYPES: 3 ♂, same locations and data as holotype (1 ♂ UZC, 1 ♂ (dried) CNC, 1 ♂ UGS) Diagnosis. This species of Wiedemannia is characterized by the elongate clasping cercus with its apical fingerlike prolongation. Description. Male. Body length ~ 4.5 mm. Head dark brown, in lateral view higher than wide; gena broad, less than half as wide as height of eye, with blue pruinescence. Face wide, with blue pruinescence; distinct carina on lower margin. Pair of ocellar and one pair of vertical setae, about 3–5 distinct upper postoculars; all similarly sized and black, with interspersed shorter setae. Lower postocular setae much finer and paler merging with pale, longer setae on middle and lower occiput. Few dark setulae present on vertex and between ocellar area and eye margin. Antenna dark brown to black; postpedicel and stylus minutely pubescent; pedicel >0.5X long as scape, with complete circlet of apical setae; postpedicel apically pointed, stylus longer; scape with setulae dorsally. Scutum dark brown; prescutellar depression with blue pruinescence, extending onto concolorous scutellum; postpronotal lobe and lower margin of notopleuron with blue pruinescence. Pleura with blue pruinescence. Mesonotum with 5 pairs of dorsocentral bristles with shorter setae interspersed between setae. Acrostichal setae, small and fine, irregularly biserial, diverging around prescutellar depression; 1 postpronotal seta, 2 notopleural setae, lower shorter and narrower; 1 presutural supra-alar seta and numerous scattered setulae; 1 postalar seta. Antepronotum with pair of dark setae and some fine setulae. Proepisternum with some fine setulae. Laterotergite with fine, pale setulae. Pair of strong marginal scutellar setae. Wing (length 5.4−5.6 mm) membrane infuscate, veins darker. Basal costa seta extending beyond humeral crossvein. Cell dm produced anteroapically. Origin of veins M1 and M2 widely separated at end of cell dm. Vein CuA+CuP short. Pterostigma elongate, brown. Squama with pale fringes. Halter dark brown. Legs entirely dark brown, rather slender, fore femur narrowed apically, posterior femur more or less linear. Rather uniformly covered with rows of small dark setulae. All coxae with longer setae anteriorly. Fore femur with 4−5 stout anterior setae on apical 0.25.

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FIGURES 10–12. Male terminalia, lateral view. 10. Wiedemannia horvati Ivković & Sinclair, sp. nov. 11. W. vedranae Ivković & Sinclair, sp. nov. 12. W. vedranae Ivković & Sinclair, sp. nov., clasping cercus, inner view. Abbreviations: cl cerc—clasping cercus, distph—distiphallus. Scale bar = 0.1 mm.

Abdomen concolorous with thorax. Terminalia (Fig. 10): Hypandrium subequal in length to epandrium,

bearing 2−3 setae. Epandrium rounded, covered with numerous dark long setae especially ventrally and laterally;

small spine-like surstylus emerging from inner face apically. Clasping cercus unilobed, basal half expanded

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medially clothed with fine setulae; apical half prolonged and finger-like, arched medially; posterior margin of lower half with row of long setae. Phallus more or less linear, brownish; distiphallus without swelling in middle. Female. Identical to male except terminalia; cercus short ovate and minutely pilose. Remarks. Wiedemannia horvati is known only from Sierra Nevada, Spain. Etymology. The new species is named after entomologist Dr Bogdan Horvat, for his teaching and guidance of the senior author on the taxonomy of aquatic Empididae.

Wiedemannia vedranae Ivković & Sinclair, sp. nov. (Figs 11, 12) Type material. HOLOTYPE ♂ (in alcohol): “HOLOTYPE/ Wiedemannia/ vedranae IVKOVIĆ & SINCLAIR/ SPAIN, Sierra Nevada,/ Río Laroles, Área/ Recreativa Laroles,/ 37°03'56''N,/ 03°02'45''W, 27.iv.2013,/ leg. M. Ivković” (UZC). PARATYPES: 23 ♂, same location and data as holotype (9 ♂, UZC, 7 ♂ (dried) CNC, 6 ♂ UGS). Additional material. SPAIN: Río Chico, Puente Palo, 36°58'12''N, 03°24'27''W, 1720 m, 17.iv.2013 (UZC); Fuente Agria, Pórtugos, 36°56'35''N, 03°18'23''W, 1274 m, 17.iv.2013 (UZC); Barranco Moja Caco, Antes de llegar al Puerto de la Ragua, 37°07'09''N, 03°01'50''W, 2028 m, 27.iv.2013 (UZC); Arroyo de Palancón, Posada de los Arrieros, 37°04'43''N, 03°01'14''W, 1720 m, 27.iv.2013 (UZC); Río Válor, Cortijo de la Sacristía, 37°03'15''N, 03°06'00''W, 1820 m, 27.iv.2013 (UZC); Río Mecina, Área Recreativa Las Chorreras, 37°03'20''N, 03°07'52''W, 1866 m, 27.iv.2013 (UZC); Río Grande de los Bérchules, Cortijo de Las Jeromillas, 37°02'38''N, 03°11'07''W, 1797 m, 27.iv.2013 (UZC); Río Mecina, Mecina-Bombarón, 37°03'18''N, 03°08'05''W, 1140 m, 27.iv.2013 (UZC); Río Nechite, Mecina-Alfahar, 37°00'36''N, 03°04'02''W, 840 m, 27.iv.2013 (UZC); Barranco de los Pasillos, Los Pasillos, 37°06'55''N, 03°04'05''W, 1646 m, 15.v.2013 (UZC); Barranco de los Tejos, Aldeire, 37°07'38''N, 03°05'24''W, 1660 m, 15.v.2013 (UZC); Barranco del Gallego, Aldeire, 37°07'56''N, 03°06'18''W, 1680 m, 15.v.2013 (UZC); Río del Pueblo, La Cabañuela, 37°08'12''N, 03°09'23''W, 1547 m, 15.v.2013 (UZC); Arroyo del Alhorí, Central eléctrica, 37°08'59''N, 03°11'56''W, 1514 m, 15.v.2013 (UZC); Río Aguas Blancas, Área Recreativa, 37°13'42''N, 03°24'32''W, 1159 m, 29.v.2013 (UZC); Río Monachil, Fuente Alta, 37°06'20''N, 03°24'33''W, 2160 m, 10.vi.2013 (UZC); Sierra Nevada Highway, 2100 m, 9.viii.1986, J.R. Vockeroth (CNC). Diagnosis. This species of Wiedemannia is distinguished by the acrostichal setae extending only to the prescutellar depression and distiphallus with spine-like projections on the median swelling. Description. Male. Body length 2.9–3.5 mm, wing length 3.0–3.7 mm . Head dark brown to black, in lateral view longer than wide; gena broad, one-third as deep as height of eye. Frons short, broader than face. Face wide with pale iridescence, with distinct carina on lower margin, bare, lacking distinctive setae. Pair of ocellar and one pair of vertical setae, about 3–4 distinct upper postoculars; all similarly sized and black. Lower postocular setae finer and merging with some black, longer setae on middle and lower occiput. Few dark setulae present on vertex and between ocellar area and eye margin. Antenna dark brown; postpedicel and stylus minutely pubescent; pedicel almost as long as scape, with complete circlet of apical setae; postpedicel apically pointed, stylus longer; scape with setulae dorsally. Scutum dark brown, postpronotal lobe and postalar ridge pale brown. Mesonotum with 5 pairs of dorsocentral setae. Acrostichal setae small and fine, irregularly biserial, extended only to prescutellar depression; 1 postpronotal seta, 2 notopleural setae, lower shorter and narrower; 1 presutural supra-alar seta and several anterior setulae; 1 postalar seta. Antepronotum with 2 pairs of dark setae. Proepisternum with some fine setulae. Katepisternum with some small fine setulae on posterior margin. Laterotergite with fine, pale setulae. Pair of strong marginal scutellar setae with some finer setulae on disc. Wing membrane infuscate, veins darker. One long basal costal seta extending beyond humeral crossvein. Cell dm produced anteroapically. Veins M1 and M2 originating together or sometimes very short stem vein proximal to M1+2 fork present. Vein CuA+CuP extremely faint. Pterostigma elongate, extremely faint, almost invisible. Squama with dark setulae. Halter brownish. Legs mostly dark brown, except coxae and ventral margins of femora light brown; fore femur without preapical anterior setae; uniformly covered with rows of small dark setulae. All coxae with longer setae anteriorly. Fore and mid femora ventrally with some longer setulae on proximal 0.5. Abdomen concolorous with thorax, covered in small setae. Pubescence darker on tergites than sternites.

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Terminalia (Figs 11, 12): Hypandrium subequal in length with epandrium; not very large, dark brown. Epandrium elongate, irregularly rectangular, covered with dark long setae especially ventrally and laterally; small curved spine-like surstylus on inner face apically. Clasping cercus bilobed, somewhat oval, with short thumb-like anterior lobe and expanded and pale brown posterior lobe (Fig. 11); small setae apically; stronger dark and shorter setae on inner face apically and laterally with longer finer setae basally; outer margin with fringe of setae (Fig. 12). Phallus more or less linear, yellowish; distiphallus with spine-like median swelling. Female. Identical to male except terminalia; cercus short ovate and minutely pilose. Remarks. Wiedemannia vedranae is known only from Sierra Nevada mountains, Spain. Etymology. The species is named after the senior author’s oldest friend, Vedrana Pretković, who was with the senior author when the species was collected for the first time.

Key to aquatic Empididae of the Sierra Nevada mountains (key written primarily for male specimens) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 -

Eyes bare, lacking ommatrichia; labellum not sucker-like; costal margin with simple setae . . . . HEMERODROMIINAE … 2 Eyes with dense ommatrichia; labellum usually sucker-like; anterior costal margin of wing with stout erect setae intermixed among longitudinal rows of simple setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .CLINOCERINAE ... 5 Cell cua (anal cell) and crossvein bm-cu absent (discal cell absent); R1 meeting costa before middle of wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemerodromia Meigen ... 3 Cells cua (anal cell) and dm present (crossvein bm-cu present); R1 meeting costa beyond middle of wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelifera Macquart … 4 Male cercus with inner digitiform process bearing 4 apical peg-like setae; apex of cercus slender, not expanded (Collin 1961, fig. 301) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemerodromia baetica Collin Male cercus with slender inner process with 2 lateral setae, lacking peg-like apical setae; apex of cercus expanded, nearly as broad as epandrium (Figs 2, 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemerodromia planti Ivković & Sinclair, sp. nov. Male cercus held erect, very long, broad and apex rounded with digitiform process at middle on posterior edge (Collin 1961, fig. 288) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelifera stigmatica (Schiner) Male cercus held horizontal, flat, narrow, inner dorsal margin with row of spine-like setae; apex tapered and lacking additional lobes (Vaillant & Chvála 1973, fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelifera vockerothi (Vaillant & Chvála) Neck arising high on occiput, from near top of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dolichocephala guttata (Haliday) Neck near centre of occiput or level with centre of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Lower margin of face lacking notch or deep cleft above mouthparts; phallus lacking apical filament . . Clinocera Meigen … 7 Lower margin of face with notch or deep cleft; phallus with articulated apical filament. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Comb of preapical anterior setae on fore femur absent; postpronotal seta reduced, shorter and thinner than notopleural setae; postsutural supra-alar setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nigra Meigen Comb of preapical anterior setae on fore femur present; postpronotal seta well developed, similar to scutal setae; postsutural supra-alar setae present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Base of R4 with short spur or appendix (Sinclair 1999, fig. 1); wing without clouding; femora entirely dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. appendiculata (Zetterstedt) Base of R4 without short spur or appendix; wings with faint clouding about crossveins; apex of femora (“knees”) light brown, compared to bluish pruinescent femur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stagnalis (Haliday) Face with setulae along inner margin of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kowarzia Mik … 10 Face bare, without setulae along inner margin of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Apex of male surstylus forked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Apex of male surstylus simple, unforked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Scutum with a narrow, dull, dark central stripe between two more shining bluish stripes; clasping cercus strongly bent; surstylus deeply forked (Collin 1961, fig. 314c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kowarzia bipunctata (Haliday) Scutum very indistinctly striped, dull; clasping cercus straight, apex slightly hooked; surstylus with shallow apical fork (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kowarzia nevadensis Sinclair & Ivković, sp. nov. Surstylus as broad as clasping cercus; clasping cercus strongly curved at middle, generally similar in width until apex (Vaillant 1965, figs 3b, d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kowarzia madicola (Vaillant) Surstylus long and slender, much thinner than clasping cercus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Surstylus with apex clothed in short dense setae (Collin 1961, fig. 314b) . . . . . . . . . . . . . . . . . . . Kowarzia tenella (Wahlberg) Surstylus with setae confined to dorsal margin (Collin 1961, fig. 314a) . . . . . . . . . . . . . . . . . . . . . . . Kowarzia barbatula Mik Wings with distinct spots; pterostigma clearly outlined, elliptical . . . . . . . . . . . . . . . . Phaeobalia lyneborgi Vaillant & Chvála Wings lacking spots; pterostigma either both faint and elongate or dark and circular . . . . . . . . Wiedemannia Zetterstedt … 15 Pterostigma rounded, usually very distinct; male clasping cercus elongate and slender, extending obliquely from epandrium (Wagner 1990, fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia (Wiedemannia) nevadensis Wagner Pterostigma elongate and usually faint; clasping cercus variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

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16 17 18

19 20 21 22 23 -

Acrostichals only present anterior to second dorsocentral setae . . . . . . . . . . . . . Wiedemannia (Roederella) ouedorum Vaillant Acrostichals extending to at least prescutellar depression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Legs yellowish brown; fore femur with distinct preapical anterior seta; clasping cercus very pale brown (Figs 5, 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia darioi Sinclair & Ivković, sp. nov. Legs brown to black, often with blue pruinescence; anterior setae if present one-third to one-fourth from apex of fore femur; clasping cercus brown to black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Fore femur with a small distinct anterior seta at about one-fourth from apex; clasping cercus held vertically, long, slender and parallel-sided, yellow, nearly twice as long as width of epandrium (Collin 1961, fig. 313c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia (Chamaedipsia) lota Walker Fore femur lacking anterior seta on apical third or with several short setae one-third from apex; clasping cercus not parallelsided, less than twice width of epandrium, colour variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Clasping cercus undivided (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia horvati Ivković & Sinclair, sp. nov. Clasping cercus bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Clasping cercus oval, with shallow notch separating lobes; length of anterior lobe less than twice its basal width . . . . . . . . 21 Clasping cercus elongate, longer than wide, with deep notch separating lobes; length of anterior lobe more than twice its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Distiphallus without spinous swelling at middle; posterior margin of clasping cercus rounded (Vaillant 1967, figs 3, 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia (Philolutra) angelieri Vaillant Distiphallus with spinous swelling at middle; posterior margin of clasping cercus truncate or angular (Figs 11, 12). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia vedranae Ivković & Sinclair, sp. nov. Anterior lobe broader than posterior lobe (Vaillant & Chvála 1973, fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiedemannia (Philolutra) veletica Vaillant & Chvála Anterior lobe slender, narrower than posterior lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Anterior lobe arising at base of clasping cercus (Vaillant 1950, fig. 20) . . . . . . . . Wiedemannia (Philolutra) fallaciosa (Loew) Anterior lobe arising at middle of clasping cercus (Vaillant 1950, fig. 19) . . . . . . . . Wiedemannia (Philolutra) aquilex (Loew)

List of aquatic dance flies or Empididae of Sierra Nevada mountain range, Spain (Clinocerinae & Hemerodromiinae) The following format is used for the distributional data: species; literature references: name of the site and in the brackets citation of the reference; New records: name of the site and in the brackets site ID. All the sites and their numbers are listed in Table 1. Hemerodromiinae Chelifera stigmatica (Schiner, 1862) Literature reference. Chestnut groves, on Pteris aquiline (Strobl 1899). New record. Río Genil, debajo del Barranco San Juan (22). Distribution. Widespread in Europe. Chelifera vockerothi Vaillant & Chvála, 1973 Literature references. N slope Mt. Veleta (Vaillant & Chvála 1973); Granada, Alhambra (Wagner 1995). Distribution. Spain (Sierra Nevada). Hemerodromia baetica Collin, 1927 Literature reference. Granada, Alhambra (Wagner 1995). Distribution. Europe. Hemerodromia planti Ivković & Sinclair, sp. nov. New records. Río Aguas Blancas, Fuente de los Trucheros (31); Río Aguas Blancas, Área Recreativa (32). Distribution. Spain (Sierra Nevada). Clinocerinae Clinocera appendiculata (Zetterstedt, 1838) Literature references: Waterfalls in upper part of Genil valley (Strobl 1909); N slope Mt. Veleta (Vaillant & Chvála 1973, as C. schremmeri (Vaillant)); Sierra Nevada Highway (Sinclair 1999). New records: Río Monachil, Fuente Alta (33); Río Guarnón, Corral del Veleta (34). Distribution. Europe. NEW AQUATIC EMPIDIDAE

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Clinocera nigra Meigen, 1804 Literature reference: Waterfalls in upper part of Genil valley (Strobl 1909). New records: Sierra Nevada Highway (CNC). Distribution. Europe and Asia. Clinocera stagnalis (Haliday, 1833) Literature references. Upper valley of Genil (Strobl 1909); Río Lanjaron, 9 km SW Orgiva (Vaillant & Chvála 1973). Distribution. Europe, North Africa, Asia, and northern North America. Dolichocephala guttata (Haliday, 1833) Literature reference. Lanjaron, Olive Garden (Strobl 1909) [record requires verification]. Distribution. Europe. Kowarzia barbatula Mik, 1880 Literature reference. Waterfalls in upper part of Genil valley (Strobl 1909). New records. Fuente, Barranco de los Pasillos (25); Río Aguas Blancas, Área Recreativa (32). Distribution. Europe. Kowarzia bipunctata (Haliday, 1833) Literature references. Waterfalls in upper part of Genil valley (Strobl 1909); Barranco de Algarrobo, 12 SW Orgiva (Vaillant & Chvála 1973). New records. Fuente, Barranco de los Pasillos (25); Barranco de los Tejos, Aldeire (26); Río Aguas Blancas, Área Recreativa (32). Distribution. Europe and North Africa. Kowarzia madicola (Vaillant, 1965) Literature reference. Stream east from Melina-Bombarón (Wagner 1995). Distribution. Central and southern Europe. Kowarzia nevadensis Sinclair & Ivković, sp. nov. New record. Barranco Frío, Hoya Carlos (4). Distribution. Spain (Sierra Nevada). Kowarzia tenella (Wahlberg, 1844) Literature reference. N slope Mt. Veleta (Vaillant & Chvála 1973). Distribution. Western Europe and North Africa. Phaeobalia lyneborgi (Vaillant & Chvála, 1973) Literature reference. Near Albergue Universitario on road from Granada to Mt. Veleta (Vaillant & Chvála 1973). Distribution. Spain (Sierra Nevada). Wiedemannia (Chamaedipsia) lota Walker, 1851 Literature reference. Barranco de Algarrobo, 12 SW Orgiva (Vaillant & Chvála 1973). New records. Río Sucio, Las Barreras (Órgiva) (1); Río Chico, Soportújar (2); Río Poqueira, Central eléctrica (Pampaneira) (5); Río Laroles, Laroles (17); Río Aguas Blancas, Cenes de la Vega (18). Distribution. Widespread in Europe and Asia. Wiedemannia (Philolutra) angelieri Vaillant, 1967 New records. Río Genil, debajo del Barranco San Juan (22); Río Monachil, Fuente Alta (33). Distribution. Spain, France and Montenegro (Ivković et al. 2013).

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Wiedemannia (Philolutra) aquilex (Loew, 1869) Literature reference. Waterfalls in upper part of Genil valley (Strobl 1909) [record requires verification]. Distribution. Europe. Wiedemannia (Philolutra) fallaciosa (Loew, 1873) Literature reference. Río Grande de los Berchules (Wagner 1995). New records. Río Sucio, Las Barreras (Órgiva) (1); Río Chico, Soportújar (2); Río Trevélez, Molino Altero (Trevélez) (7); Arroyo de Palancón, Posada de los Arrieros (9); Río Laroles, Área Recreativa Laroles (10); Río Nechite, Las Piedras del Márquez (11); Río Válor, Cortijo de la Sacristía (12); Río Mecina, Área Recreativa Las Chorreras (13); Río Grande de los Bérchules, Cortijo de Las Jeromillas (14); Río Mecina, Mecina-Bombarón (15); Río Laroles, Laroles (17); Río Genil, La Fabriquilla (19); Río Genil, Debajo del Barranco San Juan (22); Barranco de los Pasillos, Los Pasillos (24); Barranco del Gallego, Aldeire (27); Río del Pueblo, La Cabañuela (28); Arroyo del Alcázar, Área Recreativa La Tizná (29). Distribution. Europe, Asia, and North Africa. Wiedemannia (Philolutra) veletica Vaillant & Chvála, 1973 Literature reference. N slope Mt. Veleta (Vaillant & Chvála 1973). Distribution. Spain (Sierra Nevada). Wiedemannia (Roederella) ouedorum Vaillant, 1952 New records. Río Sucio, Las Barreras (Órgiva) (1); Río Genil, La Fabriquilla (19); Río San Juan, Desembocadura río Genil (21); Río Genil, Debajo del Barranco San Juan (22); Río Monachil, Cortijo de Diéchar (23); Barranco de los Tejos, Aldeire (26); Barranco del Gallego, Aldeire (27); Río del Pueblo, La Cabañuela (28); Arroyo del Alhorí, Central eléctrica (30). Distribution. North Africa, France (Corsica), Spain (Sierra Nevada). Wiedemannia (Wiedemannia) nevadensis Wagner, 1990 Literature reference. Río Trevélez by Trevélez (Wagner 1990). New records. Río Genil, La Fabriquilla (19); Río Maitena, Desembocadura (20); Río Genil, Debajo del Barranco San Juan (22). Distribution. Spain (Sierra Nevada). Wiedemannia darioi Sinclair & Ivković, sp. nov. New record. Río Aguas Blancas, Cenes de la Vega (18). Distribution. Spain (Sierra Nevada). Wiedemannia horvati Ivković & Sinclair, sp. nov. New record. Río Guarnón, Corral del Veleta (34) Distribution. Spain (Sierra Nevada). Wiedemannia vedranae Ivković & Sinclair, sp. nov. New records. Río Chico, Puente Palo (3); Fuente Agria, Pórtugos (6); Barranco Moja Caco, Antes de llegar al Puerto de la Ragua (8); Arroyo de Palancón, Posada de los Arrieros (9); Río Laroles, Área Recreativa Laroles (10); Río Válor, Cortijo de la Sacristía (12); Río Mecina, Área Recreativa Las Chorreras (13); Río Grande de los Bérchules, Cortijo de Las Jeromillas (14); Río Mecina, Mecina-Bombarón (15); Río Nechite, Mecina-Alfahar (16); Barranco de los Pasillos, Los Pasillos (24); Barranco de los Tejos, Aldeire (26); Barranco del Gallego, Aldeire (27); Río del Pueblo, La Cabañuela (28), Arroyo del Alhorí, Central eléctrica (30); Río Aguas Blancas, Área Recreativa (32); Río Monachil, Fuente Alta (33). Distribution. Spain (Sierra Nevada).

NEW AQUATIC EMPIDIDAE

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Slope

Altitude (m)

Site name

Latitude

Site ID

Longitude

TABLE 1. The list of sampling sites in the mountains of the Sierra Nevada, Spain.

1

Río Sucio, Las Barreras (Órgiva)

N 36° 54' 22''

W 03° 26' 03''

500

S

2

Río Chico, Soportújar

N 36° 55' 42''

W 03° 24' 47''

746

S

3

Río Chico, Puente Palo

N 36° 58' 12''

W 03° 24' 27''

1720

S

4

Barranco Frío, Hoya Carlos

N 36° 56' 47''

W 03° 23' 44''

1560

S

5

Río Poqueira, Central eléctrica (Pampaneira)

N 36° 56' 39''

W 03° 21' 44''

940

S

6

Fuente Agria, Pórtugos

N 36° 56' 35''

W 03° 18' 23''

1274

S

7

Río Trevélez, Molino Altero (Trevélez)

N 37° 00' 32''

W 03° 15' 26''

1460

S

8

Barranco Moja Caco, Antes de llegar al Puerto de la Ragua

N 37° 07' 09''

W 03° 01' 50''

2028

N

9

Arroyo Palancón, Posada de los Arrieros

N 37° 04' 43''

W 03° 01' 14''

1720

S

10

Río Laroles, Área Recreativa Laroles

N 37° 03' 56''

W 03° 02' 45''

1753

S

11

Río Nechite, Las Piedras del Márquez

N 37° 04' 21''

W 03° 05' 28''

1835

S

12

Río Válor, Cortijo de la Sacristía

N 37° 03' 15''

W 03° 06' 00''

1820

S

13

Río Mecina, Área Recreativa Las Chorreras

N 37° 03' 20''

W 03° 07' 52''

1866

S

14

Río Grande de Bérchules, Cortijo de Las Jeromillas

N 37° 02' 38''

W 03° 11' 07''

1797

S

15

Río Mecina, Mecina-Bombarón

N 37° 03' 18''

W 03° 08' 05''

1140

S

16

Río Nechite, Mecina-Alfahar

N 37° 00' 36''

W 03° 04' 02''

840

S

17

Río Laroles, Laroles

N 37° 00' 36''

W 03° 01' 20''

1020

S

18

Río Aguas Blancas, Cenes de la Vega

N 37° 09' 55''

W 03° 30' 53''

760

N

19

Río Genil, La Fabriquilla

N 37° 09' 21''

W 03° 26' 00''

960

N

20

Río Maitena, Desembocadura

N 37° 09' 01''

W 03° 24' 54''

1018

N

21

Río San Juan, Desembocadura río Genil

N 37° 08' 00''

W 03° 23' 24''

1170

N

22

Río Genil, Barranco San Juan

N 37° 08' 08''

W 03° 23' 24''

1200

N

23

Río Monachil, Cortijo de Diéchar

N 37° 06' 13''

W 03° 27' 15''

1416

N

24

Barranco de los Pasillos, Los Pasillos

N 37° 06' 55''

W 03° 04' 05''

1646

N

25

Fuente, Barranco de los Pasillos

N 37° 07' 02''

W 03° 04' 15''

1646

N

26

Barranco de los Tejos, Aldeire

N 37° 07' 38''

W 03° 05' 24''

1660

N

27

Barranco del Gallego, Aldeire

N 37° 07' 56''

W 03° 06' 18''

1680

N

28

Barranco del Pueblo, La Cabañuela

N 37° 08' 12''

W 03° 09' 23''

1547

N

29

Arroyo del Alcázar, Área Recreativa La Tizná

N 37° 08' 23''

W 03° 11' 07''

1494

N

30

Arroyo del Alhorí, Central eléctrica

N 37° 08' 59''

W 03° 11' 56''

1514

N

31

Río Aguas Blancas, Fuente de los Trucheros

N 37° 13' 25''

W 03° 24' 17''

1155

N

32

Río Aguas Blancas, Área Recreativa Aguas Blancas

N 37° 13' 42''

W 03° 24' 32''

1159

N

33

Fuente Alta, Museo Etnográfico

N 37° 06' 20''

W 03° 24' 33''

2160

N

34

Río Guarnón, Corral del Veleta

N 37° 03' 39''

W 03° 22' 04''

3057

N

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Discussion In total 24 species of aquatic dance flies are found on Sierra Nevada. In this research we collected 14 species, five of which are described here and Wiedemannia (Philolutra) angelieri and W. (Roederella) ouedorum are newly recorded in Sierra Nevada. Nine species (or 37% of aquatic empidid species) are endemic and currently found only in these mountains. On the data set gathered in this research and from prior sporadic research (see Ventura (2013) and above references), Sierra Nevada is possibly one of the richest regions of Europe for endemicity of aquatic dance flies. Numerous endemic species of vascular flora (Fernández-Calzado et al. 2013), terrestrial invertebrates (González-Megías et al. 2008; López-Villalta 2011), stoneflies (Tierno de Figueroa et al. 2013), and caddisflies (Sáinz-Bariain et al. 2013) are also known from this mountain range. The endemism of the Sierra Nevada is likely due to many factors, including its range in altitude and its position in the Mediterranean area, isolated from other high mountain ranges creating barriers to species dispersal and promoting allopatric speciation (Schmitt 2009). In a recent survey of the biodiversity of aquatic families in mountainous Spanish national parks, surprisingly no Empididae were listed (Guareschii et al. 2012). The apparent absence of aquatic Empididae was likely due to the sampling protocol where localities were surveyed once using kick-nets. In contrast, the present study was based on the collection of adult specimens using sweep nets and hand collecting.

Acknowledgements We thank the Sierra Nevada National Park and Consejería de Medio Ambiente of Andalucía Government for logistic help and sampling permits. Prof. Mladen Kerovec kindly provided financial support for the travel expenses of the senior author. Scott Brooks (CNC) kindly assisted in producing the digital images. We thank the referees, Dan Bickel and Adrian Plant, for their constructive suggests and comments.

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