Zootaxa 4845 (2): 275–282 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4845.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:748422C9-347E-4830-99A8-4D4309107E23 Moth flies (Diptera, Psychodidae) living in the dark of caves in the Dinaric Karst RÜDIGER WAGNER1* & TONĆI RADA2 1University of Kassel, FB 10, Biology-Zoology, Heinrich-Plett-Straße 40, D-34132 Kassel, Germany Parkstraße 65, D-36110 Schlitz, Germany. [email protected]; https://orcid.org/0000-0002-2024-1827 2Speleološko Društvo “Špiljar”, Zajdenica Tehničke Kulture Grada Splita, 21000 Split, Varardinska 53, Croatia [email protected]; https://orcid.org/0000-0001-5287-6336 *Corresponding author Abstract Seoda cavernicola sp. nov. and Psychoda glamocensis sp. nov., are new species and cave dwellers from Bosnia and Hercegovina, and Croatia. Adults of S. cavernicola are pale and small; the eye bridge is reduced, ommatidia irregularly arranged, epandrium with a pair of setose excrescences. The eye bridge of P. glamocensis is likewise reduced with 2 or 3 irregularly ordered facet rows, palpus segments of some individuals are malformed; its closest relative is Psychoda alticola Vaillant based on the morphology of male and female genitalia as well as on COI barcodes. Key words Psychodidae, troglobites, new species, Balkan peninsula Introduction The systematic research on cave dwelling animals has the best development worldwide on the Balkan Peninsula; the greatest amount of troglophilic animals is known from the Dinaric Karst. Among the cave living species found there are representatives of fish, amphibia, insects, millipeds, centipeds, spiders, crustaceans and other groups (e.g. Sket 2016). Organisms found in caves are classified troglobites (cave dwellers), troglophiles (species that sometimes oc- cur outside of caves), and trogloxenes (species that cannot live permanently in caves). Life in caves affects species ecology, morphology, and physiology, and has further implications on evolutionary processes. Insect collections from caves usually contain few Psychodidae, at most species typically live near the entrance of the caverns with permanent contact to the surface environment (troglophiles, trogloxenes). They use caves as refugium against sunlight, high air temperature, low moisture, and to avoid predators. Thus, representatives of all extant psychodid subfamilies (Phlebotominae, Bruchomyiinae, Trichomyiinae, Sycoracinae, Psychodinae) have been reported from caves on almost all continents (e.g. Bravo et al. 2008, Bravo & Barata 2012a, 2012b, Khadri Shahar et al. 2011, Sarà 1950, 1962, Polseela et al. 2011, Quate 1962, 2000, Vaillant 1966). Usually most Psychodidae from collections in caves belong, besides Phlebotominae, to the subfamily Psycho- dinae, tribus Psychodini, genus Psychoda (sensu lato); ecologically these species are assumed to be euryecious. Larvae feed on feces of bats, birds, and any other decomposing organic material (biofilms, plant and animal cadav- ers). Adults of these species are without particular adaptation to permanent life in caves. Below we describe new species of the genera Seoda Enderlein and Psychoda (s.l.) Latreille, that show remark- able morphological variation that are probably adaptations to life in caves. Material and methods Specimens were collected with hand net in caves at high altitudes in Croatia and Bosnia and Hercegovina. Adults were put in situ in 80% ethanol and used for morphological and genetic studies. For morphological study, wings of specimens were dissected and kept in clove oil; the body was macerated in Accepted by D. Cordeiro: 12 Aug. 2020; published: 2 Sept. 2020 275 10% KOH for 12-18 hrs, washed in acetic acid and put with drops of acetic acid to the clove oil with wings. After 1 day the acid evaporated and head, thorax abdomen and wings were arranged in drops of Canada balsam under 4 separate coverslips. Inspection and line drawing were made with a Leitz binocular microscope MZ 125 and Leitz MZ 20 EB with drawing mirror attached. Morphological terminology has changed and follows Kvifte & Wagner (2017). Taxonomy Seoda cavernicola sp. nov. (Figs 1–9) Etymology. The name refers to the species’ preference for caves (caverna Latin). Diagnosis. Seoda cavernicola sp. nov. is distinguished from congeners by the pale body, the eye bridge of only 2–3 irregularly ordered facet rows, scape short, pedicel globular, and 1st flagellomere elongate, ventral surface of the subepandrial plate with 2 large setose excrescences; genitalia with robust outgrowths of gonocoxites, distiphallus lobes bowed out. Material. Holotype, 1♂, Croatia, Seget Gornji, Jama I. u. Gospinu Gaju, 17 February 2019; paratypes, 2♂, col- lected at the type locality with the holotype; (one paratype with wings destroyed). Further material (all paratypes): 2♂, 3♀, Croatia, Zelovo, Jama na Vranjinoj Glavici, 19 May 2019; 2♂, Croatia, Radošić, Baračeva jama, 31 March 2019; 1♂, Croatia, Zelovo, Elezova jama 19 May 2019; all leg. T. Rada. Holotype and 1 paratype (slide mounted) and remaining material in ethanol deposited in the Natural History Museum & Zoo, Split, Croatia. 1 paratype from Seget Gornji, and 1 paratype from Radošić (slide mounted) in the collection of the senior author. Description. Male: Specimens pale, whitish. Eyes reniform, eyebridge reduced with only 2 or 3 irregular facet rows, interocular suture wide U-shaped, distance between the eyes 2 to 2.5 facet diameter (figs 1–3). Antennae (fig. 4) of all specimens incomplete; scape pipe-shaped 2x longer than wide, pedicel globular 0.5x as long as the pedicel. First flagellomere almost symmetric elongate widest at middle, subsequent flagellomeres asymmetric bottle-shaped with basal bulb and long neck. Ascoids probably lost, 2 circular holes at about middle of segments may indicate the presence of ascoids. Antennal flagellum broken, length of remaining antennal articles: 40-26-73-64-58-62-; abso- lute length: 0.105-0.0685-0.1921-0.1684-0.1526-0.1631- mm. Basal palpus segment (fig. 5) slightly longer than the scape. Four palpus segments, terminal segment flexible; relative length: 30-58-55-95; absolute length: 0.08-0.15- 0.14-0.25 mm. Wing (fig. 6) pale translucent, length 2.30 mm, width 0.74 mm; length/width 3.11. Sc short terminates in wing; radial fork distal of medial fork, at level of Cu tip; no crossveins r4-r5 or r-m. Thorax without specific features. Abdomen with 8 segments and inverted genitalia; hypandrium equally thin, in the middle with slighter sclero- tization, with fine setae in that area. Gonocoxite tubular, about 2x longer than wide, gonostylus bent dorsal about as long as the gonocoxite, basally wider. From the basal inner edges of the gonocoxites outgrowths (gonocoxal apodemes/parameres?) arise that are half as long as the gonocoxites and enclose the dorsal lobes of the basiphallus and the basal part of the distiphallus sclerites. The dorsal parameral bridge consists of two median prolongations of the gonocoxites, converging in the middle forming a fissure or keel. Basiphallus dorsoventrally flattened, bilobed from the middle on. On dorsal side of the bifurcation thin lobes arise directed dorsally converging towards the fis- sure where they are jointed to the dorsal parameral bridge. The ventral lobes of the basiphallus sclerite are apically jointed to claw-shaped laterally bent distiphallus sclerites. At the positions of the joints between basiphallus and distiphallus lobes the aedeagus is jointed to (enclosed by) the large outgrowth of the gonocoxites. Epandrium basally wider with two sharp corners, lateral and hind edges almost straight; basally with 2 small foramina. On the ventral surface of the epandrium is a pair of large setose excrescences, a well-defined subepandrial plate is not discernable. Epandrial processes almost straight in the distal part on inner side with few stronger setae in line, distally with 8 apically serrate retinacula. The hypoproct is large, rhomboid, apically setose; epiproct small setose triangular. Remarks: Small size compared with species of the genera mentioned below and pale appearance of the type specimens are interpreted as adaptation to life in caves. The species shares large apodemes of the gonocoxites and 276 · Zootaxa 4845 (2) © 2020 Magnolia Press WAGNER & RADA a bilobed basiphallus sclerite with Jungiella Vaillant, Vaillantodes Wagner, Seoda Enderlein, and Panimerus Eaton; size and shape of the distiphallus sclerites are similar to Seoda. The mode of connection of the dorsal parameral bridge and the aedeagus is unique. Usually this is by a separated Y- or U-shaped sclerite, the furca, that appears as an individual sclerite jointed with the basiphallus sclerite and dorsally coadunate in the middle with the parameral bridge. The presence of large outgrowth of the gonocoxites is similar to Jungiella Vaillant. In contrast to Jungiella, Vaillantodes, Seoda, and Panimerus which possess a separate furca the dorsal lobes in S. cavernicola sp. nov. arise from the basiphallus sclerite, they are not separated from or jointed to it. However, the task in the mode of operation is the same as for the other species: to support the back and forth movement of the aedeagus and its components. Another striking feature is the pair of setose outgrowth on the ventral surface of the epandrium; a subepandrial plate is not recognizable; however, the setose outgrowths are in the position of the subepandrial plate and therefore are interpreted as part of this. They are positioned exactly dorsal of the outgrowth of the gonocoxites. FIGURES 1–9. Seoda cavernicola. 1. Eyebridge of holotype. 2–3. Eyebridges of 2♂ paratypes. 4. Scape, pedicel, 1st and 2nd flagellomeres. 5. Basal palpus segment. 6. Wing. 7. Hypandrium, styles and aedeagus ventral view. 8. Epandrium, epandrial processes and proctiger, ventral view. 9. Basiphallus sclerite and dorsal parameral bridge with ‘furca’, ventral view. Scale: 1.0 mm (6), 0.1 mm (1–4, 7–9), 0.05 mm (5). MOTH FLIES FROM DINARIC KARST CAVES Zootaxa 4845 (2) © 2020 Magnolia Press · 277 Psychoda glamocensis sp.