Research Article |
Corresponding author: Simon Vitecek ( simon.vitecek@wcl.ac.at ) Academic editor: Ralph Holzenthal
© 2015 Simon Vitecek, Ana Previšić, Mladen Kučinić, Miklós Bálint, Lujza Keresztes, Johann Waringer, Steffen Pauls, Hans Malicky, Wolfram Graf.
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:
Vitecek S, Previšić A, Kučinić M, Bálint M, Keresztes L, Waringer J, Pauls S, Malicky H, Graf W (2015) Description of a new species of Wormaldia from Sardinia and a new Drusus species from the Western Balkans (Trichoptera, Philopotamidae, Limnephilidae). ZooKeys 496: 85-103. https://doi.org/10.3897/zookeys.496.9169
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New species are described in the genera Wormaldia (Trichoptera, Philopotamidae) and Drusus (Trichoptera, Limnephilidae, Drusinae). Additionally, the larva of the new species Drusus crenophylax sp. n. is described, and a key provided to larval Drusus species of the bosnicus-group, in which the new species belongs. Observations on the threats to regional freshwater biodiversity and caddisfly endemism are discussed.
The new species Wormaldia sarda sp. n. is an endemic of the Tyrrhenian island of Sardinia and differs most conspicuously from its congeners in the shape of segment X, which is trilobate in lateral view. The new species Drusus crenophylax sp. n. is a micro-endemic of the Western Balkans, and increases the endemism rate of Balkan Drusinae to 79% of 39 species. Compared to other Western Balkan Drusus, males of the new species are morphologically most similar to D. discophorus Radovanovic and D. vernonensis Malicky, but differ in the shape of superior and intermediate appendages. The females of D. crenophylax sp. n. are most similar to those of D. vernonensis, but differ distinctly in the outline of segment X. Larvae of D. crenophylax sp. n. exhibit toothless mandibles, indicating a scraping grazing-feeding ecology.
Caddisfly, Europe, larval key, taxonomy, conservation, Mediterranean, hydropower
The Mediterranean area is a flora and fauna biodiversity hot-spot. The Tyrrhenian islands and the Balkans, in particular, are noteworthy for their high number of plant endemics (
The genus Wormaldia currently comprises 204 species (
The genus Drusus is in the subfamily Drusinae Banks, and comprises 84 species (
In this paper we describe a new species of Wormaldia and a new grazer Drusus species, including a key to the hitherto known larval stages of the bosnicus-group, in which Drusus crenophylax sp. n. belongs.
Adults were collected using sweep nets and immature stages by handpicking. Collected specimens were stored in 70% and 96% EthOH, for morphological and molecular analyses, respectively.
Male and female genitalia were examined after being cleared in either KOH or lactic acid. Nomenclature of male genitalia of Wormaldia McLachlan follows
Molecular genetic sequence data were used to support larval association and assess relationships to previously described Drusus species. DNA extraction and amplification of a 541-bp-long fragment of the mtCOI gene using standard primers (forward primer: Jerry,
Detailed list of Drusus specimens used for mtCOI analysis. Abbreviations: M adult male, F female; L larva; U unknown.
Species | Locality | Specimen ID/Stage | Accession # | Collectors |
---|---|---|---|---|
D. crenophylax | 44°32.932'N, 17°23.562'E | fDsp4501M/M | KP793082 | Dmitrović, Šukalo |
D. crenophylax | 44°33.003'N, 17°23.580'E | fDsp4502L/L | KP793083 | Dmitrović, Šukalo |
D. crenophylax | 44°33.003'N, 17°23.580'E | fDsp4503L/L | KP793081 | Dmitrović, Šukalo |
D. crenophylax | 44°33.003'N, 17°23.580'E | fDsp3401F/F | KP793084 | Dmitrović, Šukalo |
D. crenophylax | 44°33.003'N, 17°23.580'E | fDsp3402F/F | KP793085 | Dmitrović, Šukalo |
D. vernonensis | 41°0.887'N, 21°10.448'E | DdphPEIM1/M | KC881524 | Kučinić, Graf |
D. vernonensis | 41°0.887'N, 21°10.448'E | DdphPEIM2/M | KP793087 | Kučinić, Graf |
D. vernonensis | 41°0.887'N, 21°10.448'E | DdphPEIM3/M | KP793086 | Kučinić, Graf |
D. discophorus | Macedonia, Jablanica Mts. | fDds0110M/M | KP793089 | Kučinić |
D. discophorus | Macedonia, Jablanica Mts. | fDds0112F/F | KP793088 | Kučinić |
Holotype. 1 male pupa, holotype: Sardinia, Gola di Gorruppo; 40°11.122'N, 9°30.104'E; 350 m a.s.l.; 28.03.2001; leg. Monika Hess, Ulrich Heckes; currently in coll. W. Graf, will deposited in the Biologiezentrum des Oberösterreichischen Landesmuseums, Linz, Austria.
Italy, Sardinia.
Morphology of the male terminalia suggests placement of the new species in Wormaldia. The species is unique in the European Trichoptera fauna, and easily differentiated from all other Wormaldia species by the combination of the following characters: (1) presence of median subtriangular protrusion in the distal half of the harpago, (2) membraneous dorsoproximal portion and trilobate lateral portions of segment X, and (3) distinct sclerotized structures visible on the invaginated phallus.
Adults (in pupa). Habitus dark, sclerites and tergites brown; cephalic and thoracic setal areas pale; cephalic, thoracic and abdominal setation dark brown; legs light brown, proximally darker; haustellum and intersegmental integument pale cream. Wings brown mottled with golden patches. Male maxillary palp 5-segmented. Spurformula 2–4–4 in males.
Male genitalia (Fig.
Mature pupa (Fig.
Pupal characteristics of Wormaldia sarda sp. n., Philopotamus montanus, and Wormaldia spp. A generalized pupal abdomen in dorsal view, depicting the position of the dorsal sclerites B dorsal sclerites of Philopotamus montanus C dorsal sclerites of Wormaldia occipitalis D dorsal sclerites of W. sarda sp. n. E head of W. sarda pupa in left lateral view F head of W. sarda pupa in ventral view G pupal mandibles of Philopotamus montanus in ventral view H pupal mandibles of Wormaldia copiosa in ventral view. Scale bars: 100 µm (B); 50 µm (C, D).
Female and larva unknown.
The species epithet refers to the island of Sardinia, the type locality.
Holotype. 1 male: Bosnia and Herzegovina, Cvrcka river; 44°32.932'N 17°23.562'E; 393 m a.s.l.; 01.10.2014; leg. Dejan Dmitrović, Goran Šukalo; specimen identifier: fDsp4501M. Paratypes: 2 females: Bosnia and Herzegovina, Spring of Cvrcka river, Vilenjska vrela; 44°33.003'N, 17°23.580'E; 456 m a.s.l.; 12.09.2012; leg. Dejan Dmitrović; specimen identifiers: fDsp3401F, fDsp3402F. 4 males, 3 females, 19 larvae: Bosnia and Herzegovina, Spring of Cvrcka river, Vilenjska vrela; 44°33.003'N, 17°23.580'E 456 m a.s.l.; 12.09.2012; leg. Dejan Dmitrović, Goran Šukalo; specimen identifiers for 3 larvae: fDsp4502L, fDsp4503L, fDsp4504L. Holotype and paratypes currently in coll. W. Graf, will deposited in the Biologiezentrum des Oberösterreichischen Landesmuseums, Linz, Austria.
Bosnia and Herzegovina, Republika Srpska, Cvrcka River.
Males of the new species are most similar to Drusus discophorus Radovanovic and D. vernonensis Malicky, but exhibit (1) subtriangular superior appendages in lateral view, (2) subtriangular, low tip of the intermediate appendage in lateral view, and (3) simple, rounded tips of intermediate appendages in caudal view. Drusus discophorus males have suboval superior appendages and a high round tip of the intermediate appendage in lateral view; D. vernonensis males have round superior appendages in lateral view and trilobate tips of intermediate appendages in caudal view.
Females of the new species show the reduced median lobe of the vaginal sclerite and high base of the lateral lobe of segment IX as typical for Balkan Drusinae, and are most similar to Drusus vernonensis, but exhibit (1) a sharp dorsal notch of segment X in lateral view, and (2) segment X with 2 round median lobes in dorsal view. Drusus vernonensis females have a rounded dorsal outline of segment X and lack the median lobes of segment X.
Larvae of the new species are most similar to Drusus klapaleki Marinković-Gospodnetić and D. serbicus Marinković-Gospodnetić, but exhibit (1) a semicircular area dorsomedially on the pronotum anterior the pronotal ridge void of white recumbent setae, (2) lateral gills, and (3) a subtriangular pronotal ridge in lateral view. Larvae of D. klapaleki have white recumbent setae covering the whole pronotum, and larvae of D. serbicus lack lateral gills and have an annular pronotal ridge.
Adults. Habitus dark; sclerites and tergites brown; cephalic and thoracic setal areas pale; cephalic, thoracic and abdominal setation blond; legs light brown to fawn, proximally darker; haustellum and intersegmental integument pale, whitish. Wings smoky, with dark setae. Male maxillary palp 3-segmented. Forewing length 11–13.2 mm, spur formula 1–3–3 in males; forewing length 13–14.5 mm, spur formula 1–3–3 in females.
Male genitalia (Fig.
Female genitalia (Fig.
Fifth instar larva (Fig.
Larval characteristics of Drusus crenophylax sp. n. A head, frontal view B head, left lateral view C pronotum dorsal view D meso- and metathorax with abdominal segment I, dorsal view E abdominal segments I-V, left lateral view F left thoracic leg I, frontal view G left thoracic leg II; frontal view H left thoracic leg III, frontal view; bottom, gill and lateral line diagram, positions of gills are depicted as black circles, position of lateral line bold.
Analysis of the genetic distance of mtCOI between Drusus crenophylax sp. n. and the in the adult stage morphologically most similar species, D. discophorus and D. vernonensis, clearly supports the recognition of the new species. Uncorrected p-distances recorded in a fragment of the mtCOI gene (ranging from 2–8%; Fig.
Drusinae species typically are members of crenal species communities, and mainly inhabit crenal sections of cold streams. Larval D. crenophylax were collected at eucrenal sections of the Cvrcka River (Fig.
The species epithet is a compound name, combining κρηνον (‘well, spring, fountain’ in Ancient Greek) and φυλαξ (‘guard, keeper, protector’ in Ancient Greek), terms that reflect the high degree of niche specificity of Drusus species, the majority of which inhabit crenal sections of streams (
Drusinae have evolved into three distinct subclades reflecting feeding ecology of larvae (
Larvae of the bosnicus-group also develop, with the exception of D. ramae (
1 | Head with flat vertex |
Drusus bosnicus ( |
– | Vertex evenly rounded | 2 |
2 | Pronotum with thin long, yellow setation |
Drusus radovanovici (fig. 17 in |
– | Pronotum without thin long, yellow setation | 3 |
3 | Pronotum with numerous short, white, recumbent setae | 4 |
– | Pronotum without numerous short, white, recumbent setae |
Drusus septentrionis (figs 4, 5 in |
4 | Dorsal pronotal hump smoothly rounded |
Drusus medianus (fig. 43 in |
– | Dorsal pronotal hump with distinct ridge | 5 |
5 | Anterior metanotal sclerites narrowly subtriangular (width / length ratio ≥ 2.0) |
Drusus vespertinus ( |
– | Anterior metanotal sclerites broadly subtriangular (width / length ratio < 2.0) | 6 |
6 | In lateral view, dorsal pronotal ridge annular, posterior section sharply descending |
Drusus serbicus ( |
– | In lateral view, posterior section of dorsal pronotal ridge gently descending | 7 |
7 | White recumbent setae cover the entire pronotum |
Drusus klapaleki ( |
– | White recumbent setae lacking in a semicircular area anterior to the pronotal ridge | Drusus crenophylax sp. n. |
The Tyrrhenian islands and Sardinia in particular have been renowned for their relictual fauna and flora for a long time (
The distinct apomorphic characters, particularly the modified segment X and the very different pupal characters (mandibles, dorsal abdominal sclerites; Fig.
Endemic freshwater species are particularly vulnerable to global change and (anthropogenic) habitat degradation (
The construction of hydropower dams in emerging economies is currently one of the greatest threats to freshwater biodiversity (
We are most grateful to Monika Hess and Ulrich Heckes, Germany, and Dejan Dmitrović and Goran Šukalo, Bosnia and Herzegovina, for kindly donating their material. The scientific quality of this contribution was greatly improved by the thorough reviews of two anonymous reviewers and Ralph Holzenthal.
This contribution is a result of research on the project “The Drusinae (Insecta: Trichoptera) in a world of global change“ (project number P23687-B17, PI: J. Waringer), funded by the Austrian Science Fund (FWF). This work benefited from EU 7th Framework Directive support, specifically the MARS project (contract number 603378).
S. Vitecek and W. Graf contributed equally to this manuscript. F. Hoppeler, Germany, assisted in the generation of genetic sequence data.