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Research Article
Revisionary notes on Feuerborniella Vaillant, 1971, with the first record of the genus from the Afrotropical region (Diptera, Psychodidae)
expand article infoGunnar Mikalsen Kvifte‡, Santiago Jaume-Schinkel§
‡ Nord University, Steinkjer, Norway
§ Leibniz-Institut für Biodiversität der Tiere, Bonn, Germany

Corresponding author: Gunnar Mikalsen Kvifte ( gunnar.mikalsen-kvifte@nord.no )

Academic editor: Dominique Zimmermann
© 2023 Gunnar Mikalsen Kvifte, Santiago Jaume-Schinkel.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Kvifte GM, Jaume-Schinkel S (2023) Revisionary notes on Feuerborniella Vaillant, 1971, with the first record of the genus from the Afrotropical region (Diptera, Psychodidae). Deutsche Entomologische Zeitschrift 70(1): 121-127. https://doi.org/10.3897/dez.70.97465
ZooBank: urn:lsid:zoobank.org:pub:5C970D95-0F4E-43D0-A6AB-CEB51E84124E
Open Access

Abstract

We establish a diagnosis for Feuerborniella Vaillant, 1971, based on a re-description of its type species, Feuerborniella obscura (Tonnoir, 1919) and comment on earlier diagnoses. Feuerborniella sinefurcata Kvifte & Jaume-Schinkel, sp. nov. is described, based on material from Tanzania, representing the second Afrotropical species of the genus following Psychoda morogorica Wagner & Andersen, 2007 which we treat as Feuerborniella morogorica comb. nov. We furthermore review earlier combinations, transferring Philosepedon ensiger Quate, 1996 and Philosepedon longistylus Quate, 1996 to Feuerborniella comb. nov., and briefly discuss generic limits with Quatiella Botosaneanu & Vaillant, 1970 and Nielseniella Vaillant, 1971.

Key Words

moth flies, moth fly, Psychodinae, Psychodini

Introduction

The genus Feuerborniella was named by Vaillant (1971), with the European species F. obscura (Tonnoir, 1919) as the type species. The original circumscription of the genus also comprised two species from Brazil and one from the Malay Archipelago, but the definition remained unclear for many years due to confusion with the closely-related Quatiella Botosaneanu & Vaillant, 1970 and different opinions amongst specialists regarding generic limits in Psychodini/Trichopsychodina (see discussions in, for example, Kvifte (2015); Ježek and Le Pont (2016); Kvifte et al. (2018); Kvifte (2019)).

Feuerborniella was redefined by Ibáñez-Bernal (2004), Cordeiro et al. (2014) and Cordeiro et al. (2015); the latter definition was not recognised by Ježek & Le Pont (2016) or in the classifications used in Brown et al. (2018) and Borkent et al. (2018). This is due to many characters within the diagnosis being polymorphic within the genus concept and one (the fusion of flagellomeres 11 and 12) is even polymorphic within species of Psychodini (see, for example, Quate (1955), p. 231). Furthermore, Feuerborniella obscura, the type species of the genus, differs from their diagnosis in at least two characters: flagellomere 11 and 12 are separated (e.g. Vaillant (1974), fig. 267) and the female cerci are equal or slightly shorter in length than the width of the female genital plate (Gunnar Mikalsen Kvifte, pers.obs.).

In the present paper, we thus follow the diagnoses given by Ježek (1985) and Ibáñez-Bernal (2004); however, in order to expand upon these, we also provide a supplementary re-description of the type species of the genus, Feuerborniella obscura from Europe. We also describe Feuerborniella sinefurcata sp. nov. from Tanzania and transfer Psychoda morogorica Wagner & Andersen, 2007, Philosepedon ensiger Quate, 1996 and Philosepedon longistylus Quate, 1996 to Feuerborniella, comb. nov.

Materials and methods

The studied specimens are deposited at the Royal Belgian Institute of Natural Sciences (RBINS), the Department of Natural History, University Museum of Bergen, Bergen, Norway (ZMBN) and the personal collection of Rüdiger Wagner (RW). In the material examined section, the holding institution is indicated at the end of each record and between square brackets ([]).

Measurements were made with an ocular micrometer in a microscope Leitz model Dialux 20, measurements are given in millimetres (mm). Head width was taken at the widest part, approximately above the insertion of the antennal scape, whereas the length was taken from the vertex to the lower margin of the clypeus; wing length was measured from the base of the wing at the start of the costal node to the apex of the wing-tip, while the width was taken approximately at an imaginary vertical line crossing the radial and medial forks; palpal proportions are given considering the length of the first palpal segment as a unit (1.0).

Morphological terminology is according to Kvifte and Wagner (2017).

Taxonomy

Feuerborniella Vaillant, 1971

Feuerborniella Vaillant, 1971: 119. Type species: Psychoda obscura Tonnoir, 1919.

Diagnosis

(modified from Vaillant (1974) and Ibáñez-Bernal (2004)). Eyes separated; flagellomeres 11–14 reduced in size, fused or separated; labellum of both sexes fleshy, carrying spines and hairs, but not blunt teeth; wing membrane without pilosity; R5 ending in wing apex; male genitalia with gonocoxites widely separated; aedeagus and parameres symmetrical; surstylus longer than epandrium and carrying single terminal tenaculum on the surstylus.

Remarks

Apart from Feuerborniella, two other genera of Psychodini taxa have a fleshy labellum and a single terminal tenaculum of the surstylus. Nielseniella Vaillant, 1971 has asymmetric aedeagus and parameres, setae on the wing membrane and often one of the posterior branches of the ascoids reduced (as in Threticus Eaton, 1904). The Neotropical/Nearctic Quatiella Botosaneanu & Vaillant, 1970 is separated from Feuerborniella on the gonocoxites touching medially and the surstyli being as short as or shorter than the tenacula they carry.

Species included

F. amblytes (Quate, 1999), F. ancepitis (Quate, 1996), F. bicuspis (Quate, 1996), F. ensiger (Quate, 1996) comb. nov., F. hamata (Quate, 1996), F. longistylus (Quate, 1996) comb. nov., F. morogorica Wagner & Andersen, 2007 comb. nov., F. obscura (Tonnoir, 1919), F. pandiculata (Quate, 1996), F. plaumanni (Duckhouse, 1968), F. retusus (Quate, 1996), F. sinefurcata sp. nov., F. spathipenis (Duckhouse, 1968), F. veracruzana Ibañez-Bernal, 2004.

Feuerborniella obscura (Tonnoir, 1919)

Fig. 1

Psychoda obscura Tonnoir, 1919: 140.

Psychoda eximia Feuerborn, 1923: 200.

Philosepedon uniretinacleum Krek, 1971: 92.

Material examined

Lectotype female. “Uccle Av. Defré, 21. Mai 1917, A.Tonnoir”. Designated by J. Ježek (1985). 3 female paralectotypes, one with the same data as holotype, two from “Forêt Soignes, 3 Juin 1918, A. Tonnoir”. All in coll. [RBINS]. Belgium: “Linkebeek, 26.V.[19]20, A.Tonnoir”, 2 females (RBINS). “Falaën, Juin [19]21, A.Tonnoir”, 1 male [RBINS]. “Ohain, 23.V.[19]20, R. Mayné”, 2 females [RBINS]. Germany: Hessen, Vogelsbergkreis, Schlitz, 13.VI.1971, R. Wagner leg. 1 male. [RW]

Diagnosis

Feuerborniella obscura can be separated from other described species of Feuerborniella by the following combination of characters: Wing forks complete, ejaculatory apodeme narrow, parameres dorsally connected, with elongate projections reaching apical ⅕ of aedeagus, aedeagus without subapical constriction.

Re-description

Measurements in mm (n = 1). Wing length 1.63, width 0.65; Head length 0.30, width 0.32; Antennal segments, scape: 0.06, pedicel: 0.04, flagellomere 1-9 0.10; Palpomeres 1: 0.06, 2: 0.08, 3: 0.10, 4: 0.12.

Male. Head about the same length as width, vertex about ⅓ of head length; eyes separated by 2 facet diameters, eye bridge with four facet rows, interocular suture as an inverted Y, length of suture about 2 facet diameters. Antennal scape about the same length as width, subquadrate; pedicel spherical, about the same length of scape, flagellomeres vaguely asymmetrical and nodiform, nodes progressively decreasing in size and internode increasing in length up to apical flagellomeres which are reduced in size and globular, not fused; ascoids with one anterior branch and to posterior branches, Y-shaped; frontal alveoli patch undivided, anterior margin extending almost to interocular suture reaching the second facet; labella bulbous, setose, without teeth. Palpal segments sclerotised, palpal proportions 1.0:1.4:1.7:2.1.

Figure 1. 

Feuerborniella obscura A. Wing; B. Genitalia, aedeagus, gonocoxites, gonostyli; C. Genitalia, epandrium and surstyli. Scales in millimetres (mm). Abbreviations: aed: aedeagus; eja: ejaculatory apodeme; ep: epandrium; gns: gonostyli; gnx: gonocoxite; hyp: hypandrium; pm: paramere; surs: surstyli; ten: tenaculum.

Wing hyaline, except costal cell which is infuscated; wing length 2.8 times its width; Sc short ending at the base of R1; Radial fork apical to Medial fork, M2 not connected to M1; R5 looks more sclerotised than the rest of wing veins, ending at wing apex.

Terminalia. Hypandrium sclerotised and plate-like; gonocoxites about the same length of gonostylus, cylindrical; gonostylus simple, tapering towards apex, incurved. Aedeagus symmetrical, extending towards the apex of gonostylus, parameres with a broad triangular base, tapering towards the apex, connected by a bridge morphologically ventral to the aedeagus, out-curved, ejaculatory apodeme narrow. Epandrium about the same length as its width, with both anterior and posterior margins concave; surstylus conical, tapering towards the apex with a single spatulate tenacula, about half the length of surstylus; Hypoproct with posterior margin rounded, subquadrate, covered in small setulae.

Remarks

The description is based on the male from Germany; the other specimens listed under “material examined” have been consulted only to confirm the generic diagnosis. Previous re-descriptions of F. obscura by Vaillant (1971), Ježek (1985) and Krek (1999) show some discrepancies in whether the ascoids have a posterior branch or not. Vaillant (1974) describes the ascoids as Y-shaped (i.e. three branches) on the first ten flagellomeres and as either Y-shaped or V-shaped on the following flagellomeres. Ježek (1985) describes and figures the ascoids as possessing two branches, similar to Vaillant’s (1974) illustration of the V-shaped condition. Krek (1999) figures three branches and lists the ascoids as Y-shaped, not mentioning any V-shaped ascoids on the distal flagellomeres at all. We deem it likely that this is a variable character as described by Vaillant (1974).

Feuerborniella morogorica (Wagner & Andersen, 2007), comb. nov.

Psychoda morogorica Wagner & Andersen, 2007: 293.

Diagnosis

Feuerborniella morogorica can be separated from all other Feuerborniella species by the following combination of characters: Forks complete, ejaculatory apodeme narrow, parameres not reaching beyond gonocoxites, aedeagus without subapical constriction.

Remarks

Feuerborniella morogorica is transferred to Feuerborniella, based on the labellum bulbous, aedeagus symmetric with symmetric parameres, gonocoxites widely separate and surstylus with a single tenaculum.

Feuerborniella sinefurcata sp.nov.

https://zoobank.org/1D5795FC-48BB-4ECF-9022-C81AB1F8B304
Fig. 2

Type material

Holotype male. Tanzania: Tanga Region, West Usambara Mountains, Mazumbai Forest Reserve, “Loc G & F”. 2–6.XI.1990 (Malaise trap), “ZMBs Tanzania Expedition” leg. [ZMBN].

Diagnosis

Feuerborniella sinefurcata can be separated from all other Feuerborniella species by the following combination of characters: Wing forks incomplete, ejaculatory apodeme narrow, parameres separated, reaching apical ⅕ of aedeagus, aedeagus without subapical constriction.

Description

Measurements in mm (n = 1). Wing length 1.50, width 0.63; head length 0.28, width 0.29; Antennal segments, scape: 0.06, pedicel: 0.04, flagellomere 1-3: 0.09; Palpomeres 1: 0.04, 2: 0.08, 3: 0.08, 4: 0.11.

Male. Holotype. Head about the same length as width, vertex about ⅓ of head length; eyes separated by approximately 1.5 facet diameters, eye bridge with 4 facet rows, interocular suture angular arch-shaped. Antennal scape about the same length as width, subquadrate; pedicel spherical, about the same length of scape, flagellomeres asymmetrical and nodiform, apical flagellomeres missing in revised material. Ascoids are absent in the revised material. Frontal alveoli patch undivided, anterior margin extending almost to interocular suture reaching the second facet; labella bulbous, setose, without teeth. Palpal segments sclerotised, palpal proportions 1.0:1.6:1.7:2.5.

Figure 2. 

Feuerborniella sinefurcata sp. nov. A. Head; B. Wing; C. Genitalia, epandrium and surstyli; D. Gentalia, aedeagus, gonocoxites, gonostyli. Scales in millimetres (mm). Abbreviations: aed: aedeagus; eja: ejaculatory apodeme; ep: epandrium; gns: gonostyli; gnx: gonocoxite; hyp: hypandrium; pm: paramere; surs: surstyli; ten: tenaculum.

Wing hyaline, except costal cell which is infuscated; wing length 2.6 times its width; Sc short ending at the base of R1; Radial fork apical to Medial fork, R2 not connected to R1, M2 not connected to M1; R5 looks more sclerotised than the rest of wing veins, ending at wing apex.

Terminalia. Hypandrium narrow, sclerotised, and plate-like; gonocoxites about half the length of gonostylus, cylindrical; gonostylus simple, tapering towards apex, incurved. Aedeagus extending towards the apex of gonostylus, parameres triangular-broad base, tapering towards the apex, weakly curved laterad, not connected by a bridge; ejaculatory apodeme narrow, about half the length of aedeagus. Epandrium about the same length as its width, with both anterior and posterior margins concave; surstylus cylindrical, slightly tapering towards the apex with a single spatulate tenaculum, about half the length of surstylus; Hypoproct with posterior margin rounded, tongue-shaped, covered in small setulae.

Etymology

From Latin sine, meaning without and, furca, meaning fork – referring to the reduced radial and medial forks in the new species.

Remarks

Feuerborniella sinefurcata sp. nov. is the 19th species of Psychodidae to be described from the West Usambara Mountains (see Kvifte (2022)) and the second Afrotropical species of Feuerborniella.

Discussion

Vaillant (1974) recognised four species in his first review of Feuerborniella, spanning the Neotropical, Palearctic and Oriental Regions. Ježek (1985) considered Psychoda plaumanni Duckhouse, 1968 and Psychoda spathipennis Duckhouse, 1968 to belong to his genera Psycha Ježek, 1983 and Psychomora Ježek, 1983 and considered Trichopsychoda malayensis Satchell, 1955 to belong to an undescribed genus. Neither these changes nor his transfer of the Oriental Psychoda nigripennis Brunetti, 1908 to Feuerborniella were supported by mention of specific morphological characters and they have generally not been considered in subsequent works.

Ibáñez-Bernal (2004) followed Vaillant’s (1974) initial species list and added an additional Neotropical species, lifting the world total to five and Cordeiro et al. (2014) recognised six species with their description of F. paramuna. Many additional combinations were made by Cordeiro et al. (2015), but since their diagnosis is not consistent with the type species of Feuerborniella, the validity of these combinations must be considered on a case-by-case basis.

Our current diagnosis adds the criteria of wing membranes without pilosity and symmetry in the male genitalia to previously-published diagnoses. This excludes the Oriental Trichopsychoda malayensis Satchell, 1955 and the Neotropical Feuerborniella pilosella Cordeiro & Bravo, 2015 and F. paramuna Cordeiro, 2014, which would all be Feuerborniella species, based on the diagnoses of Ibáñez-Bernal (2004) and Cordeiro et al. (2014). These species all have pilose wing membranes and (based on illustrations) an asymmetric aedeagus; in the case of F. pilosella, the parameres are asymmetric as well. According to the differential diagnosis given above, these may belong to Nielseniella; however, it would be premature to transfer the species to that genus until the generic limits of Nielseniella are re-assessed, based on study of the type species. Psychoda nigripennis Brunetti, 1908, which was placed in Feuerborniella by Ježek (1985), was insufficiently described in its original description and only a damaged collection of females remained of the type series when it was revised by Quate (1962). That species can, therefore, not be confidently placed in any genus. Of the 14 described species considered by us to form part of Feuerborniella, eleven are Neotropical, one is Palearctic and two are Afrotropical. With the high number of undescribed Feuerborniella species encountered in the Zurquí project, we suggest the genus to have its highest diversity in the Neotropical Region (Borkent et al. 2018; Brown et al. 2018).

The present paper represents the first mentions of Feuerborniella from the Afrotropical Region, which means existing keys will have to be emended. In Kvifte’s (2015) key to Afrotropical genera of Psychodini, Feuerborniella keys to couplet 6 where neither of the two options work (surstylus with two tenacula vs. surstylus with three or more tenacula). The same is the case for Kvifte and Wagner (2017) where Feuerborniella keys to couplet 29 which uses the same character. Feuerborniella can be separated from all other Afrotropical Psychodini by the labellum being bulbous rather than flat and the surstylus carrying one tenaculum only.

Acknowledgements

We are grateful to Trond Andersen for organising the expedition to the West Usambara Mountains and to him and Rüdiger Wagner for giving us access to collections in their care. Maurice Leponce at the Royal Belgian Institute of Natural Sciences kindly facilitated GMK’s visit to the RBINS where he examined Tonnoir’s type material of Feuerborniella obscura. Santiago Jaume-Schinkel’s work on European Psychodidae is supported by the Bundesministerium für Bildung und Forschung, Berlin, Germany, the project “German Barcode of Life III: Dark Taxa” (FKZ 16LI1901A). We extend our gratitude to Morgane A. Kerdoncuf for opening her flat to SJS during his stay at Bergen. Danilo Cordeiro, Sergio Ibáñez-Bernal and an anonymous reviewer offered constructive criticisms of an earlier version of the manuscript, which we found very helpful in improving the scientific merit of this work. We are indebted to the Museum für Naturkunde, Berlin, for waiving the author’s fees for this manuscript.

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