ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2014 Magnolia Press

Zootaxa 3878 (4): 390–400

www.mapress.com/zootaxa/Article

http://dx.doi.org/10.11646/zootaxa.3878.4.5

http://zoobank.org/urn:lsid:zoobank.org:pub:08CC020F-7D39-414A-9158-6C42036660EB

Nomenclature and taxonomy of Telmatoscopus Eaton and Seoda Enderlein;

with a discussion of parameral evolution in Paramormiini and Pericomaini

(Diptera: Psychodidae, Psychodinae)

GUNNAR MIKALSEN KVIFTE1,2

1Department of Limnology, Institute of Biology, University of Kassel. Heinrich-Plett Str. 40, 34132 Kassel–Oberzwehren, Germany2Department of Natural History, University Museum of Bergen, P.O. Box 7800, University of Bergen, N-5020 Bergen, Norway.

E-mail: Gunnar.Kvifte@um.uib.no

Abstract

Historically, Telmatoscopus Eaton, 1904 has been a nomenclaturally and taxonomically problematic taxon as different au-

thors have used different type species to define their concepts of the genus. Here it is shown that Pericoma advena Eaton,

1893 is the valid type species rather than Pericoma morula Eaton, 1893. Furthermore, the genus Seoda Enderlein, 1935

is revived for the genus comprising Pericoma labeculosa Eaton, 1893, P. morula and their relatives. The differences be-

tween Telmatoscopus and Seoda are described in detail based on historical and freshly collected material of the three pu-

tative type species. Four new synonymies are proposed: Panimerus havelkai Wagner, 1975 and Telmatoscopus seguyi

Vaillant, 1990 are synonymized with Telmatoscopus advena, and Telmatoscopus incanus Nielsen, 1964 and Telmatosco-

pus vaillanti Withers, 1986 are synonymized with Seoda morula.

A potential phylogenetic pattern in the male genital sclerites is discussed in detail. In Telmatoscopus, the jointed ap-

pendages of the gonocoxally derived parameral complex are separate small sclerites found near the bases of the distiphal-

lic lobes of the aedeagus. In Seoda, they are fused medially to form a small, moveable triangular or arrow-shaped sclerite.

Medial parameral sclerite fusion in Psychodinae is otherwise known to occur only in Pericomaini and the paramormiine

genus Psychomasina Ježek, 2004; however, many genera of Paramormiini show an apparently intermediate condition

where the parameres are fused in one end to form a V- or U-shaped "furca". It is hypothesized that Paramormiini is para-

phyletic with respect to Pericomaini, as suggested in a previous phylogenetic hypothesis based on molecular data.

Key words: moth flies, new synonymy, male genitalia, homology

Introduction

Tribe Paramormiini Enderlein, 1935 (=Telmatoscopini Vaillant, 1971, partim) is distributed in all biogeographical regions apart from Antarctica and comprises roughly 300 described species in an uncertain number of genera. Moth flies in this group are recognized by their antennal flagellomeres being asymmetrically nodiform without significant reductions in size of the apical flagellomeres, labella bulbous, apical segment of maxillary palpus striated, R

2+3 without connection to R

4, aedeagal complex nearly always symmetrical and dorsal gonocoxal

condyles fused medially. Many of these characters are of dubious phylogenetic value as they are widespread in Psychodinae, and no clearly synapomorphic characters have been identified. Thus, the monophyly of Paramormiini may be uncertain.

Some taxa within Paramormiini are readily recognized by means of specialized sensory appendages of the head and antennae, e.g. Panimerus and Jungiella. The majority of genera, however, are still unsatisfactorily characterized; particularly the group referred to as the ”telmatoscopoid group” of genera by Duckhouse (1978). Species in this group typically have paired, unbranched, digitate ascoids and symmetrical genitalia with distally jointed aedeagal appendages.

Much of the confusion regarding the taxonomy of Paramormiini is due to differing opinions about which species constitutes the type species of Telmatoscopus. The International Code of Zoological Nomenclature (ICZN

390 Accepted by G. Curler: 9 Oct. 2014; published: 28 Oct. 2014

1999, § 67.2) states that a species needs to be one of the originally included species in a genus to be eligible for type species designation. No species are explicitly mentioned as members of the genus in the original description of Telmatoscopus by Eaton (1904); however it is stated that the taxon is "nearly co-extensive with Sections 3 C, 3 D, and 4 A of Pericoma in the Supplement of my Synopsis", referring to Eaton (1893–1894). Due to disagreements on what constitutes an "originally included species", four different nomina have been proposed as type species by subsequent authors. Tonnoir (1933) considered that the species thus referred to were to be treated as originally included, and designated Pericoma morula Eaton, 1893 as the type species as it was the first mentioned by Eaton (1893). This designation was followed by Quate (1955), Duckhouse (1966), Ježek (1984, 1989), Krek (1999) and Bravo et al. (2011).

An implicit listing of species such as that given by Eaton (1904), however, does not constitute a valid species listing according to the ICZN (§67.2.3, see also §69). For a species to be available for type species designation, it needs to be explicitly mentioned as belonging to the genus. The earliest explicit species placements in Telmatoscopus were made by Eaton (1912), who described Telmatoscopus rothschildi Eaton, 1912 and compared it to Telmatoscopus advena (Eaton, 1893) and Telmatoscopus meridionalis (Eaton, 1894) (=Clogmia albipunctata

(Williston, 1893)). In recognition of ICZN (§67.2.3), Quate (1965) designated Pericoma advena Eaton, 1893 as the type species of Telmatoscopus. This concept was later followed by Duckhouse (1978), Vaillant (1983), Wagner (1990) and Kvifte (2012).

Two additional designations of type species for Telmatoscopus exist. Enderlein (1935, 1937) designated Pericoma soleata Walker, 1856, a species which was later designated as the type of Jungiella Vaillant, 1972. Enderlein's (1935) designation is invalid because P. soleata was not included in Telmatoscopus by Eaton (1912). Finally Vaillant (1971) designated Psychoda albipunctata Williston, 1893 as the type of Telmatoscopus; however, this is invalid due to the existence of a previous valid designation.

As shown in the above synopsis, Pericoma advena is the validly designated type species of Telmatoscopus (see ICZN §67.2.3). As suggested by Ježek & Mogi (1995) and Ježek (2001), Pericoma advena and Pericoma morula

are too different in structure to be considered congeneric. With this in mind, P. morula must be transferred from Telmatoscopus to a different genus. Seoda Enderlein, 1935 is therefore revived to accomodate some species previously placed in Telmatoscopus auct.

Material and methods

Material from the following collections was examined:

BMNH The Natural History Museum, London, United KingdomRW Private collection of Rüdiger Wagner, University of Kassel, GermanyZMUB Natural History Collections, University Museum of Bergen, Norway

Illustrations and measurements were based on specimens in the RW collection. Wing measurements are given in mm with an accuracy of 0.03 mm, other measurements in µm with an accuracy of 5 µm. Measurements are given as ranges followed by the mean. Morphological terminology is derived from Quate & Brown (2004), modified as follows: The aedeagus in Psychodinae consists of a basiphallus and a distiphallus, and the distiphallus is comprised of two phallomeres. The gonocoxites of the gonopods have mostly membranous inner appendages which are referred to here as parameres; these often give rise to jointed, paired or unpaired parameral sclerites and frequently also a membranous parameral sheath (=phallic sheath of Kvifte et al. 2013). The often sclerotized morphodorsal margin of the parameres (=chitinous inner bridge of Vaillant 1971-1983) are called the gonocoxal condyles.

The following abbreviations and colour codings are used in the illustrations:

aed j—aedeagal joint (sclerotized basal part of distal phallomere, linking distiphallus to basiphallus and ventral epandrial plate); colour-coded by dark hatchingbp—basiphallusCn—costal node

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gcx c—gonocoxal condyle; colour-coded by light hatchingpar—parameral sclerites; colour-coded dark greyphm—phallomerepms—parameral sheath; colour-coded light grey.

Taxonomy

Telmatoscopus Eaton, 1904

Telmatoscopus Eaton, 1904: 58. Type species: Pericoma advena Eaton, 1893, by subsequent designation of Quate (1965)= Sciria Enderlein, 1935: 247. Type species: Pericoma advena Eaton, 1893, by original designation.

Diagnosis. Frons and clypeus clearly separate, not protruding over eye margin; flagellomeres asymmetrically nodiform, with paired, curved, leaf-shaped to digitiform ascoids; flagellomere 13 with elongate neck, flagellomere 14 with elongate apiculus; R

2+3 not connected to R

4; apex of R

5 ending in apex of wing; basiphallus narrow in dorsal

view and distally ending in two short branches with membranous connection to distiphallus; distiphallus composed of paired, symmetrical phallomeres; aedeagal complex encapsuled in parameral sheath, parameral sclerites separate, usually slightly curved in dorsal and ventral views, usually narrow, arranged transversely near bases of phallomeres; surstylus with indistinctly fringed tenacula; epandrium with single aperture.

Species included:

Telmatoscopus advena (Eaton, 1893) Telmatoscopus dendrophilus Vaillant, 1983 Telmatoscopus laurencei Freeman, 1953

Telmatoscopus pappi (Wagner, 1979) Telmatoscopus patibulus Quate, 1955

Telmatoscopus tanegashimensis (Ježek & Mogi, 1995) comb. nov.

Telmatoscopus thuringicus Beran, Doczkal, Pfister & Wagner, 2010

Discussion. The definition of the advena group as distinct from other Telmatoscopus auct. was first pointed out by Vaillant (1989), and followed upon by Ježek & Mogi (1995). Due to the latter authors' incorrect assertion that Pericoma morula is available for type species designation in Telmatoscopus, however, they used the name Sciria

Enderlein, 1935 for the advena group. Some authors, e.g. Salmela (2005) and Svensson (2009) followed this arrangement; however, since Pericoma advena is the type species of both Telmatoscopus and Sciria, the latter name falls as an objective junior synonym. For the taxa similar to Pericoma morula, the name Seoda Enderlein, 1935 is available (type species Pericoma labeculosa, see below).

Telmatoscopus advena (Eaton, 1893)

Pericoma advena Eaton, 1893: 127= Panimerus havelkai Wagner, 1975: 1, syn.nov.

? = Telmatoscopus seguyi Vaillant, 1990: 378. syn. nov.

Material examined. Holotype male for Panimerus havelkai Wagner, 1975. GERMANY: Hessen, Schlitz, light trap, 23.-28.V.1974 (50.669609°N, 9.568590°E)

GERMANY: Rheinland-Pfalz, Winnweiler, Imsbach, Weisse Grube (N 49°35'32.3'' E 7°54' 0.7''), 355 m.a.s.l., 30.IV.2007, D. Weber leg. (RW). Hessen, Schlitz area, Rimmelswiesen (N°50.698860, E°9.484820), 2.VI.1975, R. Wagner leg. (RW). Bayern, Schweinfurt area, Oberes Garstadter Holz (N°49.978984, E° 10.179700 )

GREAT BRITAIN: Devon, Branscombe, 15.V.1901, A.E. Eaton leg. (BMNH)NORWAY: Rogaland, Finnøy, Helgøy, 5.VI-5.VII.2011. 1 m, T. Jonassen leg.; Hordaland, Bergen, Milde,

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Hatlehaugen. 25.V.2011. 1 m, B. Røttingen & A. Schrøder-Nielsen leg.; Sogn og Fjordane, Aurland, Fretheim, 12.VI.1939, 1 m, Knaben leg. (ZMUB)

FIGURE 1. Telmatoscopus advena (Eaton, 1893). Male from Germany. A, head, B, wing, C, aedeagal complex and gonocoxite. Length of scale bars (in µm): 100, 250, 100

Description. Adult male (n=8). Head (fig. 1A) broader than long; vertex length approximately one fifth that of head, with short neck; postocular setae difficult to distinguish from other vestiture on head, reaching half the level between side and median of head; eyebridge comprised of four facet rows, separated by half a facet diameter; interocular suture broadly U-shaped; frontal alveolar patch shaped like a narrow triangle, reaching interocular suture; length of palpomeres (n=4) 80–105 (90), 165–180 (170), 165–180 (170), 225–255 (240); labellum bulbous, setose; scapus cylindrical, pedicel elongate globular; flagellomeres asymmetrically nodiform; asymmetry less pronounced on first flagellomere; basal flagellomeres almost as long as scapus, distalmost flagellomeres slightly shorter than basal ones; length of scapus and pedicel (n=3) 100–120 (110), 75–80 (75); length of flagellomeres 1–11 (n=4) 115–135 (120), 125–140 (130), 120–150 (135), 125–150 (135), 120–150 (135), 120–145 (130), 115–145 (130), 115–145 (130), 115–140 (125), 110–140 (125), 110–135 (120); length of flagellomeres 12–14 (n=3) 100–120 (110), 90–110 (100), 80–105 (90); Thorax without special features; dorsum with setae in three broad longitudinal stripes; anepisternum, katepimeron and scutellum setose, anepimeron bare; posterior spiracle densely setose; legs without special features; Wing (fig. 1B) length 2.31–2.73 (2.51), breadth 0,87–1,05 (0,93); C thickened near base; Sc reaching level of base of R

2+3; radial and medial forks about in line with each other; CuA2

not reaching level of wing forks; jugum trapezoid; Terminalia (fig. 1C) with hypandrium narrow, setose, with broad unsclerotized stripe; gonostylus (not illustrated) elongate cylindrical, with median curve, slightly longer than gonocoxite, gonocoxite subcylindrical with pointed acuminate dorsal and ventral projections meeting at middle; parameres mostly membranous, but with pair of parameral sclerites at level of basiphallic/distiphallic joint; aedeagus with basiphallus narrow, distiphallus symmetrical, with a sclerotized joint basally and curved, sabre-

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shaped and tubular projections distally; epandrium (not illustrated) rectangular, slightly longer than broad; with single oval aperture; subepandrial plate square with rounded edges, lateral margins with thin sclerotized margins; surstyli elongate, curved, of even width throughout length; distally with 23–29 elongate tenacula which are almost as long as surstylus; tenacula distally indistinctively fringed; proctiger with hypoproct oval, epiproct triangular; both micropilose.

Remarks. Telmatoscopus advena has been recorded from the Czech Republic, Finland, France, Germany, Great Britain, Ireland, Norway, Slovakia and Sweden (Eaton 1893, Withers & O'Connor 1992, Ježek 1998, Withers 2004, Salmela 2005, Svensson 2009, Oboňa & Ježek 2012, Kvifte & Boumans 2014). Records from Germany have hitherto only been published as Panimerus havelkai Wagner, 1975; however, there appears to be no significant differences between this species and Telmatoscopus advena. The synonymy was first proposed by Wagner in

Salmela (2005); however, it has not previously been formalised. Another possible synonym is Telmatoscopus seguyi Vaillant, 1990. Most of the characters cited in the original

description to distinguish this species from T. advena fall well within the variability of T. advena; the exceptions being the number of "transverse tergapodemes" (based on the illustrations, these are apparently the margins of the subepandrial plate) and the shape of the phallomeres (parameres of Vaillant). Both of these characters may be artifacts of specimen preparation rather than characters present in the animal itself. The observed variation in ”tergapodemes” may stem from different degrees of sclerotization/bleaching of the specimens, and the phallomere shape can appear different in specimens oriented differently on microscope slides. This synonymy should, however, be confirmed through study of the type material.

Many authors write Telmatoscopus advena as Telmatoscopus advenus, but this is invalid. The name advena is latin for "stranger" and is to be treated as a noun in apposition. Nouns do not transform according to the gender of the genus name, and the correct species epithet of T. advena is therefore "advena".

Seoda Enderlein, 1935

Seoda Enderlein, 1935: 248. Type species: Pericoma labeculosa Eaton, 1893, by original designation.= Krekiella Vaillant, 1971: 72. Type species: Pericoma labeculosa Eaton, 1893, by original designation.

Diagnosis. Frons fused with clypeus into single facial plate; protruding over lower median eye margin (fig. 2A); flagellomeres asymmetrically nodiform with ring of filiform ascoids, pair of digitiform ascoids often present as well; flagellomere 13 with long neck, flagellomere 14 with elongate apiculus; R

2+3 not connected to R

4; Apex of

wing between apices of R4 and R

5 (fig. 2B); Aedeagus broad in dorsal and ventral views, symmetrical, distiphallus

consisting of two jointed phallomeres with sclerotized joints and more or less sclerotized lateral appendages; aedeagus encapsulated in parameral sheath; short unpaired median parameral sclerite present (figs 2C, 2D); Proctiger elongate diamond-shaped; Surstylus with fringed tenacula; Epandrium with single aperture.

Species included:

Seoda labeculosa (Eaton, 1893) Seoda ambigua (Eaton, 1893) comb.nov.

Seoda bosnica (Krek, 1978) comb.nov.

Seoda britteni (Tonnoir, 1940) comb.nov.

Seoda carpathica (Ježek, 1989) comb.nov.

Seoda carthusiana (Vaillant, 1972) comb.nov.

Seoda collarti (Vaillant, 1972) comb.nov.

Seoda falcariformis (Wagner, 1978) comb.nov.

Seoda gressica (Vaillant, 1972) comb.nov.

Seoda hajeki (Ježek, 1997) comb.nov.

Seoda ligustica (Sarà & Salamanna, 1967) comb.nov.

Seoda morula (Eaton, 1893) comb.nov.

Seoda mucronata (Vaillant, 1972) comb.nov.

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Seoda orbiculata (Krek, 1971) comb.nov.

Seoda remota (Wagner, Koç, Özgül & Tonguç, 2013) comb.nov.

Seoda schlitzensis (Wagner, 1975) comb.nov.

Seoda similis (Tonnoir, 1922) comb.nov.

Seoda svanetica (Ježek, 1989) comb.nov.

Seoda tetraspiculata (Ježek & Goutner, 1993) comb.nov.

Seoda verbassica (Krek, 1978) comb.nov.

Seoda wagneri (Salamanna, 1982) comb.nov.

Seoda labeculosa (Eaton, 1893)

Pericoma labeculosa Eaton, 1893: 127= Telmatoscopus latinervosus Vaillant, 1964: 63. Synonymized by Vaillant (1972)

Material examined. GREAT BRITAIN, Devon: "Near Parhaynes, Seaton Junction", 18.VI.1904, 2 ♂♂. "Near Seaton Junction", 13.VI.1901, 1 ♂. "Near Pottlake, Shute", 3.VI.1902, 1 ♂., all A.E. Eaton leg. (BMNH)

LUXEMBOURG: "Quelle E6, NW Horas Rheoholokrene 310 m NN" 15.VI.1999, I. Schrankel leg., 1 ♂ (RW)

FIGURE 2. Seoda labeculosa (Eaton, 1893). Male from Luxembourg. A, head, B, wing, C, aedeagal complex and gonocoxite, D, Seoda morula (Eaton, 1893). Male from Germany. Aedeagal complex and gonocoxite. Length of scale bars (in µm): 100, 250, 100, 100

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Description. Adult male (n=5, for measurements n=1). Head (fig. 2A) longer than broad; vertex between two and three times length of head, tapering; postocular setae indistinct, reaching base of eyebridge; eyebridge of four facet rows, separated by one facet diameter; interocular suture present, Y-shaped with very short stem; frontal alveolar patch trilobate with median "lobe" reaching interocular suture; frontoclypeus widened over mesal margins of eyes, produced to about eyebridge width in front of eyes; length of palps 85 : 130 : 150 : 200; scape barrell-shaped, 2.5 times length of pedicel; flagellomeres slightly asymmetrical, carrying numerous filiform ascoids and one pair of narrow digitiform ascoids; ascoids slightly longer than flagellomere; most flagellomeres positioned such as to make their measurements unfeasible; thorax destroyed in described specimen; wing (fig. 2B) length 2.1, breadth 0.84; apex of wing between R

4 and R

5; wing forks and apices of Sc and CuA

2 widened; Sc approaching C;

medial fork slightly distal of radial fork; dense circular patch of setae present dorsally of jugum, jugum widely triangular; terminalia (fig. 2C); hypandrium produced triangularly; gonostylus (not illustrated) elongate curved cylindrical, longer than gonocoxite; gonocoxite reniform; gonocosal condyles arched, platelike; triangular parameral sclerite present medially, linked basally to distiphalli; aedeagal complex surrounded by parameral sheath; basiphallus broad with wide aperture; distiphallus consisting of two symmetrical phallomeres sclerotized basally, connected to ventral epandrial plate medially; distal parts of phallomeres membranous, shovel-shaped; epandrium (not illustrated) approximately square with single aperture; surstyli elongate of even width, six times as long as wide; tenacula with tips fringed; tenacula slightly less than twice width of surstylus; epiproct round, hypoproct elongately diamond-shaped.

Remarks. Seoda labeculosa is here recorded from Luxembourg for the first time. The species is previously known from Belgium, the Czech Republic, France, Great Britain, Ireland and Lithuania (Tonnoir 1919, Nielsen 1964, Withers & O'Connor 1992, Bernotienè 2002, Ježek 2003).

Seoda morula (Eaton, 1893)

Pericoma morula Eaton, 1893: 127= Telmatoscopus incanus Nielsen, 1964: 153, syn.nov.

?= Telmatoscopus vaillanti Withers, 1986: 229, syn.nov.

Material examined. Paralectotype ♂: GREAT BRITAIN: Somerset: Stoney Stoke, 27.V.1892, 1 ♂, A.E. Eaton leg. (BMNH) Paratype ♂ for Telmatoscopus vaillanti Withers, 1986.

GREAT BRITAIN: Norfolk: Upton Broad, 2.VI–9.VI.1983, A.Irwin & W.Urwin leg. 1 ♂ (Malaise trap, RW)GREAT BRITAIN: Devon, Branscombe, 5.VI.1901, 1 ♂. Dorset, Blackdown, Broadwinsor, 24.V.1904, 2 ♂♂,

Dorset, Bridport, Symondsbury, 20.V.1910, 1 ♂, all A.E. Eaton leg. (BMNH)GERMANY: Schleswig-Holstein: Eutin near Bungsberg, 31.V.1982, no collector. 1 ♂ Schierenseebach,

1.VI.1985, Holm leg. 1 ♂. Hessen, Schlitz, Breitenbach, no date, no collector. 1 ♂. Thüringen: Rosoppe bei Martinfeld, 10.VI.1982, W. Joost leg. 1♂ (all RW)

LUXEMBOURG: "Quelle E1 Gutland, N Haereberg (Die Kirch)" 18.V.1999, I. Schrankel leg. 1 ♂ (RW)POLAND: Łódź highland, Widawka river, Kodrac. 9.VII.1984, S. Niesołowski leg. 1 ♂ (RW)AUSTRIA: Lunz, Teichbach. 29.V.1972 4 ♂♂, 30.V.1972, 2 ♂♂ (RW)Description. Adult male (n=12). Head about as broad as long, vertex a third of total head length; postocular

bristles reaching bases of eyebridge; eyebridge of four facet rows, separated by two to three facet diameters; interocular suture present, U-shaped; frontal alveolar patch trilobate, median lobe reaching interocular suture; frontoclypeus widened over mesal margins of eyes, produced to about eyebridge width in front of eyes; length of palp segments (n=6, 5,5,3) 100–120 (115), 160–170 (165), 155–165 (160), 225–235 (230); labellum bulbous, setose; scape cylindrical, pedicel globular; flagellomeres nodiform assymetrical; basalmost and distalmost flagellomeres shorter than middle flagellomeres; third and fourth flagellomeres with short spines on lateral side; ascoids not observed in examined specimens, but ring of attachment points present above ring of setae; length of antennal segments 1–4 (n=6) 220–250 (235), 65–75 (70), 90–105 (95), 80–100 (90); length of antennal segment 5 (n=5) 110–125 (115); length of antennal segments 6–15 (n=4) 115–130 (120), 115–130 (120), 115–125 (120), 115–120 (120), 115–120 (120), 110–120 (115), 105–115 (110), 100–110 (105), 95–100 (100); length of antennal segment 16 (n=3) 85–95 (90); Thorax without special features; dorsum with setae in three longitudinal stripes; anepisternum, katepimeron and scutellum setose, anepimeron bare; posterior spiracle setose; legs without special

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features; wing length 2.16–2.49 (2.3), breadth 0.9–1.02 (0.95); apex of wing between R4 and R

5; Sc approaching C,

but not reaching it; medial and radial forks aligned; base of medial fork weakened; dense oval patch of setae present dorsal to jugum; jugum U-shaped with squarish edges; terminalia (fig. 2D); hypandrium of even width; gonostylus (not illustrated) acuminate, with knee-like bend at median, about as long as gonocoxite; gonocoxite reniform, gonocoxal condyles circularly arched; triangular parameral sclerite present medially, linked basally to phallomeres; parameral sheath enclosing aedeagal complex except stylets of phallomeres; basiphallus broad with wide aperture, distiphallus consisting of two symmetrical phallomeres sclerotized basally, no obvious connection to epandrium apparent; distal and lateral parts of phallomeres membranous, unsclerotized lateral stylets present; epandrium (not illustrated) square with single aperture; surstyli cylindrical, of even width throughout length, five times as long as broad; with 8–12 tenacula; tenacula faintly fringed apically; hypoproct elongately diamond-shaped, epiproct round.

Remarks. Seoda morula is here recorded from Luxembourg and Poland for the first time. The species has previously been recorded from Belgium, Denmark, Germany, Great Britain, Ireland, Italy, Lithuania, The Netherlands and Sweden (Tonnoir 1919, Barendrecht 1934, Nielsen 1964, Vaillant 1972, Caspers & Wagner 1980, Withers & O'Connor 1992, Bernotienè 2002). A record from Bosnia and Herzegovina by Krek (1971) was later described as a separate species, Seoda verbassica (Krek, 1978). A record from Italy by Sará (1960) was later suggested by Vaillant (1972) to be a misidentification. This is likely to be true because the characters given in Sará (1960) are inconsistent with the characters as observed in the present study.

The synonymy of Telmatoscopus vaillanti Withers, 1986 with Seoda morula is based on the examination of a paratype male. The separation of the two species appears to be based on the presence or absence of lateral distiphallic stylets—a character which is very prone to misinterpretation if tissues are destroyed during bleaching or if the orientation of the specimen on the slide is not perfectly dorsoventral. In all other characters, the paratype is identical to morula specimens from Britain and continental Europe. In the absence of reliable characters, the species is therefore tentatively placed in synonymy.

The species from Austria recorded as Panimerus incanus by Wagner (1980) shows small, but consistent genitalic differences from specimens collected elsewhere in Europe. They have, therefore, been omitted from the measurements and descriptions, as they may represent a different species.

Discussion

Seoda and Telmatoscopus, in the sense of the present study, were first formally separated by Ježek & Mogi (1995). This decision was further justified in Ježek (2001), whose identification table agrees very well with the characters given herein for separation of the two genera. Some further characters, newly recognized, include (for Seoda) the flagellomeres with a ring of small setiform ascoids in addition to the main digitiform ascoid pair, the frons fused with the clypeus and protruding over the mesal margins of the eyes, and notably the medial fusion of the parameral sclerites. In Telmatoscopus, there appears to be a single digitiform to foliform ascoid pair, the frons is separate from the clypeus and the parameral sclerites are widely separated.

The parameral sclerite differences between Seoda and Telmatoscopus may have phylogenetic and ultimately taxonomic implications. In the ground-plan condition for Psychodinae genitalia, the parameral sclerites are most likely paired, separate and symmetrical. This condition is present in members of several tribes (cf. Vaillant 1982). In most Paramormiini, the sclerites have moved toward the middle and fused. Jungiella, Parajungiella and Vaillantodes have parameral sclerites in the form of a V-shaped sclerite which Vaillant (1972) calls a "furca" (cf. Parajungiella, fig. 3B). The unpaired median sclerite in Seoda is of parameral origin and probably represents a next step in this development, that is, the two sclerites have fused completely at the midline. This also seems to be the case in the paramormiine genus Psychomasina Ježek, 2004 from Madagascar (Ježek 2004, Kvifte unpubl.).

Apart from Seoda and Psychomasina, the only psychodine taxa to posess an unpaired median parameral appendage belong to tribe Pericomaini. In this tribe, the aedeagus mostly has the lateral branches of the distiphallus fused distally to form a spatulate structure; the parameral sclerite is placed medially and can be moved in the dorso-ventral plane (see Kvifte et al. 2013, fig. 1D). Another example of how the aedeagal complex works in this tribe is illustrated in figure 3D.

Zootaxa 3878 (4) © 2014 Magnolia Press · 397TAXONOMY OF TELMATOSCOPUS AND SEODA

FIGURE 3. Proposed evolutionary sequence of parameral sclerites in Paramormiini. Cladogram modified from Espindola et al. (2012). A, Telmatoscopus advena, B, Parajungiella longicornis (Tonnoir, 1922), C, Seoda labeculosa and D, Pneumia stammeri (Jung, 1954)

The presence of an evolutionary trend within Paramormiini from paired lateral parameral sclerites to a single median appendage may indicate that Pericomaini is derived from within Paramormiini. The hypothesised evolutionary sequence is illustrated in figure 3. Interestingly, this idea also matches very well with the general pattern in the molecular phylogeny of Espíndola et al. (2012): This study resolved Paramormiini as paraphyletic with respect to Pericomaini, and found those taxa who have a single median sclerite, such as Parajungiella,

Jungiella and Seoda britteni, to be closer to Pericomaini than to the separate-sclerite taxa such as Telmatoscopus

advena and Paramormia Enderlein, 1935 (fig. 3). This possible paraphyly of Paramormiini may explain the problems earlier authors have had with defining convincing synapomorphies for the taxon (Vaillant 1990, Curler & Courtney 2009).

Acknowledgements

I am grateful to Rüdiger Wagner for granting me access to his research collection and to him and Gregory Curler for valuable discussions and comments to the manuscript. I am furthermore indebted to Peter Chandler, who suggested the synonymy of Telmatoscopus vaillanti with Seoda morula. Finally I wish to thank Zoe Adams at the Natural History Museum, London for the loan of valuable historical specimens.

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