Research Article |
Corresponding author: Jindřich Roháček ( rohacek@szm.cz ) Academic editor: Owen Lonsdale
© 2021 Jindřich Roháček.
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:
Roháček J (2021) The Herniosina story continues in the Mediterranean: H. calabra sp. nov. from Calabria and H. erymantha Roháček, new female from the Peloponnese (Diptera, Sphaeroceridae). ZooKeys 1061: 165-190. https://doi.org/10.3897/zookeys.1061.72235
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A study of recently acquired material of Herniosina Roháček, 1983 (Diptera: Sphaeroceridae: Limosininae) in the Mediterranean subregion revealed a new species, H. calabra sp. nov. (Italy: Calabria: Serre Calabresi Mts) and the first females of H. erymantha Roháček, 2016 (Greece: southern Peloponnese: Taygetos Mts). Herniosina calabra sp. nov. (both sexes) and the female of H. erymantha are described and illustrated in detail including structures of terminalia, their relationships are discussed and new information on their biology (habitat association) is given. An update of a key to all know species of Herniosina species is presented.
Biology, distribution, Europe, Herniosina Roháček, key, Limosininae, relationships, taxonomy, terminalia
The genus Herniosina Roháček, 1983 (Sphaeroceridae: Limosininae) has recently been reviewed by
Herniosina was originally described during the re-classification of the previous genus Limosina Macquart, 1835 by
The genus Herniosina has been newly diagnosed by
All the material examined (including specimens of Herniosina bequaerti used for illustrations but not listed below) is deposited in SMOC, Slezské zemské muzeum, Opava, Czech Republic.
Abdomens of some specimens were detached, cleared by boiling several minutes in 10% solution of potassium hydroxide (KOH) in water, then neutralised in 10% solution of acetic acid (CH3COOH) in water, washed in water and subsequently transferred to glycerine. Postabdominal structures were dissected and examined in a drop of glycerine under binocular microscopes (Reichert, Olympus). Detailed examinations of genital structures were performed with a compound microscope (Zeiss Jenaval). After examination, all dissected parts were put into small plastic tubes containing glycerine, sealed with hot forceps and pinned below the respective specimens. Specimens with abdomen removed and terminalia dissected are indicated in the list of material by the abbreviation genit. prep.
Legs were drawn on squared paper using a Reichert binocular microscope with an ocular screen. Details of the male and female genitalia were drawn by means of Abbe’s drawing apparatus on a compound microscope (Zeiss Jenaval) at larger magnification (130–500×). Wings were photographed on the compound microscope Olympus BX51 with an attached digital camera (Canon EOS 1200D). Whole adult (dry-mounted) specimens and wings were photographed by means of a digital camera Canon EOS 5D Mark III with a Nikon CFI Plan 10×/0.25NA 10.5 mm WD objective attached to a Canon EF 70–200 mm f/4L USM zoom lens. The specimen photographed by means of the latter equipment was repositioned upwards between each exposure using a Cognisys StackShot Macro Rail and the final photograph was compiled from multiple layers (20–40) using Helicon Focus Pro 7.0.2. The final images were edited in Adobe Photoshop CS6.
Six main characteristics of the new species were measured: body length (measured from anterior margin of head to end of cercus, thus excluding the antenna), index t2 : mt2 (= ratio of length of mid tibia : length of mid basitarsus), wing length (from wing base to wing tip), wing width (maximum width), C-index (Cs2 : Cs3) (= ratio of length of 2nd costal sector : length of 3rd costal sector) and index rm\dm-cu : dm-cu (= ratio of length of section between rm and dm-cu on discal cell : length of dm-cu). All type specimens and also all newly collected specimens of H. erymantha were measured.
Label data of primary-type specimens are presented strictly verbatim including information on form and colour of all associated labels. Data from paratypes of the new species and also from formerly unpublished non-type specimens are standardised and presented in full. Phenological and other biological information obtained from the material examined and literature are given in the Biology paragraph.
Morphological terminology follows that used for Sphaeroceridae by
A1 anal vein;
ac acrostichal (seta);
ads additional (setulae) on frons;
C costa;
ce cercus;
Cs2, Cs3 -2nd, 3rd costal sector;
CuA1 cubitus;
dc dorsocentral (seta);
dm discal medial cell;
dm-cu discal medial-cubital (= posterior, tp) cross-vein;
dp distiphallus;
ea ejacapodeme;
ep epandrium;
f1, f2, f3 fore, mid, hind femur;
g genal (seta);
gs gonostylus;
hu humeral (= postpronotal) (seta);
hy hypandrium;
ifr interfrontal (seta);
M media;
mt2 mid basitarsus;
oc ocellar (seta);
occe outer occipital (seta);
occi inner occipital (seta);
ors fronto-orbital (seta);
pg postgonite;
pha phallapodeme;
pp phallophore;
pvt postvertical (seta);
R1 1st branch of radius;
R2+3 2nd branch of radius;
R4+5 3rd branch of radius;
r-m radial-medial (= anterior, ta) cross-vein;
S1–S10 abdominal sterna;
sc scutellar (seta);
stpl sternopleural (= katepisternal) (seta);
T1–T10 abdominal terga;
t1, t2, t3 fore, mid, hind tibia;
va ventroapical seta on t2;
vi vibrissa;
vte outer vertical (seta);
vti inner vertical (seta).
Holotype
♂ labelled: “ITALY: W Calabria: Serre Calabresi Mts, Mongiana 2.4 km N, 38°32'05"N, 16°19'06"E “, ”1000 m, 25.5.2018, in tufts of Juncus in alder forest, J. Roháček leg.“, ”Holotypus ♂ Herniosina calabra sp. n., J. Roháček det. 2021“ (red label). The specimen is dry-mounted on pinned triangular card, intact (SMOC 06/001/2018-1, Fig.
The name of the new species is an adjective derived from Calabria (a region in southern Italy) where the type locality of the new species is situated.
Male (Fig.
Thorax
dark brown to black, mesonotum relatively shining because of sparse microtomentum, pleuron more densely microtomentose and duller (Fig.
Legs
dark brown, coxae, trochanters, knees and tarsi brown to pale brown. f1 with sparse and relatively short setae in posterodorsal and posteroventral rows. f2 with a row of 4–6 curved but relatively short ventral setae in basal third (Fig.
Wing
(Fig.
Abdomen
blackish brown to black, with only some postabdominal sclerites brown. Preabdominal terga (Figs
Herniosina calabra sp. nov. (male paratype). 3 Abdomen, laterally 4 mid tibia, dorsally 5 mid femur and tibia, anteriorly 6 preabdominal sterna, ventrally 7 S5, ventrally. Abbreviations: ce – cercus, ep – epandrium, gs – gonostylus, S – sternum, T – tergum. Scale bars: 0.2 mm (3, 6); 0.3 mm (4, 5); 0.1 mm (7).
Genitalia. Epandrium (Figs
Herniosina calabra sp. nov. (male paratype). 8 Genitalia, laterally 9 external genitalia, caudally 10 gonostylus, laterally 11 aedeagus, dorsally 12 ditto, laterally 13 aedeagal complex, laterally. All scale bars 0.1 mm. Abbreviations: ce – cercus, dp – distiphallus, ea – ejacapodeme, ep – epandrium, gs – gonostylus, hy – hypandrium, ma – medandrium, pg – postgonite, pha – phallapodeme, pp – phallophore.
Female (Fig.
Postabdomen
(Figs
Herniosina calabra sp. nov. seems to be morphologically intermediate between H. bequaerti and H. erymantha. Although seemingly more similar to H. erymantha (smaller body size, shorter male T5 and S8, male S5 with deeply forked medial process, anterior process of male cercus long and robust, gonostylus ventrally rounded, not emarginate, spermathecae with conical basal part distally ringed) it is probably most closely related to H. bequaerti. Sister-species relationship of H. calabra and H. bequaerti seems to be particularly demonstrated by the following putative synapomorphies: very similar construction of the male aedeagal complex, including the short distiphallus (with both lateral lobes and unpaired ventral process short) and surprisingly similarly formed, short and robust postgonite. In the female postabdomen there is also a shared synapomorphy: the modified (posterodorsally more or less incised) lateral part of S10 (cf. Fig.
The new species can be easily separated from all known congeners only by postabdominal characters. The most species-specific are as follows: the long, slender and deeply forked medial process of male S5 (Fig.
The entire type series of H. calabra sp. nov. (21 specimens) was collected (aspirated by a pooter) in May under Juncus tufts (Fig.
Hitherto only known from S. Italy (Calabria).
Herniosina erymantha Roháček, 2016: 80 [male only, phylogenetic notes, illustr.]. Type locality: Greece, Peloponnese, Alepochori 0.5 km SE.
Holotype ♂ labelled: ”GREECE: NW Peloponnese: Alepochori 0.5 km SE 37°58'57"N,21°48'10"E“, “590 m, 27.5.2015, sifting leaves under Platanus, J. Roháček leg.“, “Holotypus ♂ Herniosina erymantha sp. n., J. Roháček det. 2016“ (red label). The specimen is dry-mounted on pinned triangular card, with left wing and abdomen detached, genitalia dissected and all removed parts preserved in glycerine in coalesced plastic tube pinned below the specimen (SMOC).
Greece: SW Peloponnese: Taygetos Mts, Nedousa 0.5 km W, 37°08'35"N, 22°12'17"E, 390 m, sweeping riverside vegetation, 5.x.2017, 2♂ (1♂ genit. prep.); Taygetos Mts, Artemisia 1 km E, 37°05'47"N, 22°14'27"E, 655 m, sweeping vegetation along brook, 7.x.2017, 1♂, 9.x.2017, 1♀ (genit. prep.); Taygetos Mts, Saidona 1.5 km NE, 36°53'16"N,22°17'59"E, 820 m, sweeping vegetation along brook, 8.x.2017, 1♂ (genit. prep.), all J. Roháček leg. (SMOC).
Male (Fig.
Thorax. Scutellum relatively large and long (1.5 ~ as wide as long), rounded triangular, with dense fine microsculpture on flat dorsal surface. Thoracic chaetotaxy: 1 or 2 stpl, posterior long, anterior reduced to microseta or absent.
Legs. f2 with a long row of 6–8 curved but relatively short ventral setae in basal half to two-thirds. t2 : mt2 = 1.83–1.90.
Wing. Discal cell (dm) variable, relatively short to medium long, distally usually less tapered than in most relatives, with small process of M beyond dm-cu being continued by a venal fold; posterior outer corner of dm obtuse-angled to rounded, sometimes with small remnant of CuA1. Wing measurements: length 1.87–2.38 mm, width 0.77–1.01 mm, C-index = 0.88–1.09, rm\dm-cu : dm-cu = 2.62–3.15.
Abdomen.
Male S5 (Fig.
Genitalia.
Epandrium besides a group of longer and stronger setae laterally and lateroventrally usually also with 1 longer dorsolateral seta which can sometimes be reduced (as is in the holotype, see
Female (Fig.
Postabdomen
(Figs
Herniosina erymantha sp. nov. has only been known from the male holotype (
On the other hand, female H. erymantha can be easily distinguished from females of both its relatives (and also from all other congeners) by the unusually robust cerci (Fig.
Almost all newly obtained specimens of H. erymantha were swept from above decaying leaf-litter and sparse vegetation under Platanus trees in valleys of montane brooks in the Taygetos Mts (Figs
Habitat of Herniosina erymantha Roháček, 2016. 31 valley of montane brook surrounded by Platanus trees near Nedousa village in Taygetos Mts (Greece: Peloponnese) (5.x.2017) 32 Platanus growth near brook near Artemisia village in Taygetos Mts with the Czech dipterists J. Starý (on left) and M. Vála (on right) in foreground (9.x.2017).
Hitherto only known from Greece: Peloponnese.
1 | Male | 2 |
– | Female | 7 |
2(1) | S1+2 with a strong protruding bulge (Figs |
3 |
– | S1+2 only slightly protruding ( |
6 |
3(2) | S5 with a single long medial process that is apically more or less forked (Figs |
4 |
– | S5 with 2 small digitiform medial processes ( |
H. horrida (Roháček, 1978) |
4(3) | S5 with medial process long and robust, in lateral view sinuous (Fig. |
H. bequaerti (Villeneuve, 1917) |
– | S5 with medial process either short and in lateral view pestle-shaped (Fig. |
5 |
5(4) | S5 with medial forked process short and robust, pestle-shaped in lateral view (Figs |
H. erymantha Roháček, 2016 |
– | S5 with medial forked process long and slender (Fig. |
H. calabra sp. nov. |
6(2) | Preabdominal sterna sparsely setose ( |
H. pollex Roháček, 1993 |
– | Preabdominal sterna more densely setose ( |
H. hamata Roháček, 2016. |
7(1) | T6, T7, S6 and S7 shorter and more transverse ( |
H. hamata Roháček, 2016 |
– | T6, T7, S6 and S7 longer, narrower, less transverse (Figs |
8 |
8(7) | S10 divided into 2 lateral sclerites ( |
9 |
– | S10 undivided, horseshoe-shaped ( |
11 |
9(8) | Cerci short and robust (Figs |
H. erymantha Roháček, 2016 |
– | Cerci long and slender (Figs |
10 |
10(9) | T6 narrow, hardly wider than T7 ( |
H. bequaerti (Villeneuve, 1917) |
– | T6 broad, wider than T7 (Fig. |
H. calabra sp. nov. |
11(8) | T10 longer, elongately triangular ( |
H. horrida (Roháček, 1978) |
– | T10 shorter, transversely subtriangular ( |
H. pollex Roháček, 1993 |
The monophyly of Herniosina was demonstrated by
Herniosina species, comparison of male characters. 33 H. bequaerti, S5, ventrally, 34 same species, process of S5, laterally 35 H. calabra sp. nov., S5, ventrally 36 same species, process of S5, laterally 37 H. erymantha, S5, ventrally 38 same species, process of S5, laterally 39 H. bequaerti, gonostylus, laterally 40 H. calabra sp. nov., gonostylus, laterally 41 H. erymantha, gonostylus, laterally. All scale bars 0.1 m.
The addition of Herniosina calabra sp. nov. did not affect the concept of the genus which remains a compact group of similar species, differing mainly by the structures of the male and female terminalia. This new species seems to be most closely allied to H. bequaerti and H. erymantha, and, consequently the hypothesis of the relationships of species within Herniosina has to be changed as follows. Herniosina hamata is considered a sister-taxon to the five other congeners which belong to a monophyletic group supported by the following putative synapomorphies: male preabdominal sclerites with setosity reduced; male cercus modified to 2 (lateral and medial) processes; gonostylus with dorsal internal projection; spermathecae pyriform, with distinct conical basal part. Herniosina pollex, with the bulge of the male S1+2 small (a plesiomorphy shared with H. hamata) is considered a sister-group to a clade with H. horrida, H. erymantha, H. calabra and H. bequaerti being supported by 2 synapomorphies (male S1+2 strongly bulging; gonostylus with slender dorsal internal projection). Herniosina horrida (having male S5 with a pair of small posteromedial projections = a plesiomorphy shared with H. hamata and H. pollex) seems to branch off the remaining triplet formed by H. erymantha, H. calabra, and H. bequaerti. The close alliance of these three species is supported not only by 3 synapomorphic features in the male terminalia (S5 with posteromedial distally forked projection; medial process of cercus small and apically pointed; funnel-shaped apex of distiphallus short and robust) but also by female S10 medially desclerotised or divided. Finally, H. erymantha is considered a sister-group to H. calabra – H. bequaerti pair, the latter being supported by similar formation of the aedeagal complex (distiphallus shortened, postgonite short and robust) and modified female S10 having lateral parts posterodorsally emarginated to incised.
Discovery of the new species, H. calabra, added new information to the general distribution of the genus Herniosina. The currently known distribution of the genus ranges from Spain in the west to Russia (Kabardino-Balkariya) in the east and from Iceland and Fennoscandia in the north to Spain, S. Italy, Cyprus and Israel in the south (
It is an agreeable duty to thank my companions on collecting trips to the Calabria (2018) and Peloponnese (2017), M. Vála and J. Starý (Olomouc, Czech Republic) (see Fig.