Research Article |
Corresponding author: Marc Pollet ( mpollet.doli@gmail.com ) Academic editor: Marija Ivković
© 2018 Marc Pollet, Andreas Stark.
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:
Pollet M, Stark A (2018) The quest for the identity of Orthoceratium lacustre (Scopoli, 1763) reveals centuries of misidentifications (Diptera, Dolichopodidae). ZooKeys 782: 49-79. https://doi.org/10.3897/zookeys.782.26329
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Recently, a species of Orthoceratium was collected in Greece that differs morphologically from the European species commonly presumed to be Orthoceratium lacustre (Scopoli, 1763). Verification of the identity of the Greek species through comparison with 460 specimens of Orthoceratium from 17 West Palaearctic and one Afrotropical country, and examination of existing type material, revealed that the species recognized as O. lacustre in northwestern Europe for over 250 years is actually O. sabulosum (Becker, 1907), the other known species in the genus, which was originally described from Tunisia. Although the types of O. lacustre have been lost, a comparison of the distribution ranges of both species in Europe provided evidence that the species collected in Greece is conspecific with O. lacustre. Both species have distinct distributions in the West Palaearctic, with O. lacustre largely restricted to the northern border of the Mediterranean basin, and O. sabulosum more widespread, occurring in northwestern Europe, the western, southern, and eastern Mediterranean, the Middle East, and the Afrotropical Region (Tanzania). Both species are redescribed and fully illustrated, a neotype is designated for O. lacustre and a lectotype for O. sabulosum, and a key to males and females is provided. The misidentifications that lasted for over two centuries are explained by the omission by previous authors to study the type specimens, and inaccuracies in species descriptions and keys.
distribution, Dolichopodidae , ecology, Europe, North Africa, Mediterranean basin, Orthoceratium lacustre , Orthoceratium sabulosum , taxonomy, types
Orthoceratium Schrank, 1803 is a nearly exclusively West Palaearctic dolichopodid genus in the subfamily Hydrophorinae (
1 | Seven dorsocentral bristles. Two large inner and two smaller outer scutellar bristles. Two basal postpronotal bristles. One proepisternal bristle. Proepimeron simple. Male: fore femur with posteroventral pollinose spot. Fore tarsus simple. Abdomen with 5th tergite with lateroventral process. Fore tarsus with one claw. Hypopygium with robust, large cercus | Orthoceratium Schrank, 1803 |
- | Six dorsocentral bristles. Six equally strong scutellar bristles. One basal postpronotal bristle. Proepisternal bristles absent. Proepimeron with distinct ventral acute process. Male: fore femur simple. Fore tarsus with 2nd tarsomere flattened. Abdomen with 5th tergite simple. Fore tarsus with two claws. Hypopygium with small cercus with apical filiformous process | Liancalus Loew, 1857 |
In the process of verifying the identity of ‘species B’, the depository and availability of the type specimens of both described species was checked. This revealed that the Scopoli types of O. lacustre had been lost (Lorenzo Munari, pers. comm.), and that the status of O. sabulosum type specimens in Becker’s collection could be questioned. Moreover, series of Orthoceratium specimens from different European museums contained both O. sabulosum and ‘species B’. It thus appeared crucial to establish whether ‘species B’ actually corresponded with O. lacustre or represented a new, third species.
In the present paper, we present the results of this study, and give (re)descriptions of the species and information on their distribution and ecology. We finally discuss the plausible reasons for the continuous series of misidentifications, and the significance of type specimen examination.
Two specimens of Orthoceratium sabulosum from the Becker collection (Museum für Naturkunde,
(Re)descriptions are based on a large number of representative specimens of each species, both in alcohol and pin-mounted. A total of 173 character states was scored, with 35, 61, and 77 related to the head, thorax/abdomen/wing, and legs respectively. This allowed us to determine the most reliable and consistent decisive diagnostic features that were subsequently applied in the key.
Relevant non-genitalic diagnostic characters in collected specimens were photographed by the junior author. The hypopygium and tergite V of each species were drawn using a camera lucida. The left lateral view of the hypopygium is illustrated here. In describing the hypopygium, ‘dorsal’ and ‘ventral’ refers to the morphological position prior to genitalic rotation and flexion. Thus, in the drawings showing a lateral view of the hypopygium, the top is morphologically ventral, while the bottom is dorsal.
Biometrics were generally based on five specimens (wet = preserved in alcohol solution) of each gender in each of the two species unless otherwise mentioned, and include: (i) face width, (ii) body length, (iii) wing length (= distance between basis of basicosta and wing apex), (iv) relative wing width, (v) proximal versus apical section of vein M1, (vi) proximal versus apical section of vein CuA1, (vii) CuAx ratio (= crossvein dm-cu versus apical section of vein M1) and (viii) relative lengths ratio of femur, tibia and tarsomeres of each leg. The latter relative lengths were recalculated so that the shortest leg part represents a value of “1”. Wing length was measured in both dry and wet specimens. All values given in this paper are average values, unless otherwise mentioned. Palp and proboscis size is compared to the eye size, measured as the vertical diameter (from about ocellar tubercle to the lower eye margin). Wing length was measured in 76 and 142 specimens of O. lacustre and O. sabulosum resp., to find out if differences occurred between both species and separate populations (see Table
Wing lengths (in mm) in males and females of O. lacustre and O. sabulosum. Measurements per country are given for those countries where at least five specimens of each sex were examined.
Biometrics | Mean (min–max) | No. specimens | Mean (min–max) | No. specimens |
---|---|---|---|---|
Sex | Male | Female | ||
Orthoceratium lacustre | ||||
BULGARIA | 5.8 (5.5–6.2) | 9 | 6.3 (5.8–6.7) | 11 |
FRANCE | 5.7 (5.4–6.0) | 13 | 6.1 (5.2–6.7) | 13 |
ITALY | 5.8 (5.4–6.2) | 5 | 6.2 (5.7–6.5) | 5 |
GREECE | 5.8 (5.2–6.1) | 7 | 6.4 (5.9–6.6) | 8 |
All specimens | 5.7 (5.2–6.2) | 35 | 6.2 (5.2–6.7) | 41 |
Orthoceratium sabulosum | ||||
BELGIUM | 5.6 (5.3–5.9) | 20 | 6.1 (5.8–6.4) | 20 |
Dudzele | 5.6 (5.3–5.9) | 5 | 6.1 (6–6.3) | 5 |
Lissewege | 5.8 (5.6–5.9) | 5 | 6.2 (5.9–6.4) | 5 |
Knokke (Het Zwin) | 5.7 (5.5–5.9) | 5 | 6.1 (5.8–6.3) | 5 |
GREAT BRITAIN | 5.5 (5.2–5.8) | 19 | 5.9 (5.2–6.4) | 17 |
NETHERLANDS | 5.8 (5.5–6.0) | 6 | 6.1 (5.7–6.5) | 5 |
SPAIN | 5.4 (4.5–5.8) | 14 | 5.8 (5.5–6.1) | 5 |
All specimens | 5.6 (4.5–6.1) | 77 | 6 (5.2–6.5) | 65 |
Capture locations of Orthoceratium specimens are given in Figure
Distribution ranges of Orthoceratium lacustre (red symbols, incl. type locality of neotype) and O. sabulosum (black symbols, incl. type locality of lectotype) in the West Palaearctic (Iranian records not included). Information related to the site codes is given in Suppl. material
The general morphological terminology follows
ac acrostichal bristles;
ad anterodorsal;
ant pprn anterior postpronotal (= humeral sensu
ap apical;
apv apicoventral;
av anteroventral;
bas pprn basal postpronotal (= posthumeral sensu
bv basoventral;
dc dorsocentral bristle pairs;
ds dorsal;
MSSC(s) male secondary sexual character(s);
npl notopleural;
pal postalar;
pd posterodorsal;
psut ial presutural intra-alar (= presutural sensu
pv posteroventral;
S abdominal sternite;
spal supra-alar;
sut ial sutural intra-alar (= sutural sensu
ta tarsomere, 1-5 in the descriptions of tarsi refers to basal (1) to apical (5) tarsomeres;
T abdominal tergite;
vt ventral;
I, II, III refers to fore, mid and hind leg;
I–VI in the descriptions of abdominal segments (tergites/sternites) refers to basal (I) to caudal (VI) segments.
Institutional, collection and other abbreviations:
ANSC Andreas Stark private collection, Halle/S., Germany;
IBER Institute of Biodiversity and Ecosystem Research, Sofia, Bulgaria;
MAPC Marc Pollet private collection, Welle, Belgium;
MIBC Miroslav Barták private collection, Prague, Czech Republic;
ZLKU Zoology Laboratory, Department of Biology, Faculty of Science, Muğla Sıtkı Koçman University, Muğla, Turkey;
Other abbreviations: MT: Malaise trap, SW: collected by sweepnet.
Label information of mounted specimens is provided in full and with the original spelling. If not indicated otherwise, the label was white and rectangular, and information is from the top side. Label information is given from the top downward, with data from each label between quotation marks, and data from different lines on the same label separated by a slash (/). Information from different labels is separated by a semi-colon (;). The species record is followed by the repository of each specimen between square brackets []. In addition to the label information, for non-type specimens, the most relevant label information is enriched, uniformly structured and given in the following format: “(site code) – COUNTRY: ♂, ♀, province (or equivalent administrative division), locality, location/area, latitude, longitude, altitude, sampling date (start) – sampling date (end), sampling method, collector [collection]” (see Suppl. material
A total of 428 specimens of Orthoceratium from eight European museums and three other collections has been examined, mainly by the senior author; the identity of two, two, five and 23 additional specimens from
To our surprise, the examination of the type material of Orthoceratium sabulosum revealed that the species widely known (and collected) as ‘O. lacustre’ in northwestern Europe was conspecific with this species. This held true for nearly all specimens of ‘O. lacustre’ examined from North Africa and the Middle East (Turkey, Iran) as well. The question evidently raised if ‘species B’ then represented the true O. lacustre or not. As mentioned before, establishing the species concept of the latter species proved difficult due to the loss of the type material. Hence, the original description and other literature sources were studied carefully in search for information on significant diagnostic features that matched those of ‘species B’.
“Diagn. Thorax aeneus. Abdomen viridi-aeneum. Ambulat super aquas stagnantes tanquam Cimex Lacustris [now in Gerris]. Habitat in lacubus. Frons subargentea. Oculi virides. Pili duo divaricati in occipite. Antennae nigrae, clavatae, obtusae. Rostrum palpis subvillosis, parvis. Thorax aeneus, glaber. Alae hyalinae, immaculatae; costa antice ferruginea. Scutellum edentatum, rotundatum, pilosum. Abdomen lineam longum, viridi-aeneum, albido villo adspersum, subtus subfuscum: segmentis lateraliter punctatis. Pedes longi: lamellis unguium pallidis.”
Unfortunately, all listed characters fit both O. sabulosum and ‘species B’. Scopoli’s species, however, seemed to occur in stagnant water bodies (see original description) like e.g., inland lakes, and did not seem to be confined to saltmarshes or brackish marshes like O. sabulosum in northwestern Europe.
As the Scopoli type specimens of O. lacustre were destroyed as early as 1787, it is unlikely that
“Wings hyaline, usually tinged with ferruginous towards the fore edge, … Abdomen of the male … lamella oblong, compressed, broad at tip and truncated. … On waters, both fresh and brackish. (E[ngland]. I[reland]).”
However, it can be assumed that Haliday based this description on ‘O. lacustre’ from England or Ireland, which now appears to be O. sabulosum.
“Beine schwarz mit gelben Knieen und Gelenken. — Metallisch-grün. Untergesicht silberweiss schimmernd. Fühler schwarz. Rückenschild undeutlich gestriemt. Analanhänge länglich, zusammengedrückt, am Ende breit und abgestutzt. Schenkel oben grün, auch der hinterste Metatarsus. Flügel glashell, gegen den Vorderrand gewöhnlich bräunlich, gelblich tingirt, die vierte Längsader gebrochen. 2 2/3 ‘’’. Nach Scopoli in [Herzogtum] Krain; ich erhielt die Art durch Hrn. Micklitz aus dem Küstenlande.”
Neither
“Face white, hardly wider than postpedicel (length) (male). Mesonotum dusted whitish grey on dorsum; sternite IV with ventral process (male). Wing entirely hyaline. Size 5 mm | O. lacustre (Scopoli, 1763) |
Face grey, about 2 × as wide as postpedicel (length) (male). Mesonotum dusted yellowish grey on dorsum; sternite IV normal (without process) (male). Wing brownish. Size 6 mm | O. sabulosum” [trans.] |
The only feature of O. lacustre in this key that matches ‘species B’ is the narrower face, compared to O. sabulosum. But specimens of both species show an equally large variation in the colour of the wings and the dusting of their pronotum. This is quite remarkable as we believe that Becker actually had specimens of both species at hand when he described O. sabulosum, as he repeatedly refers to O. lacustre. Why he used doubtful differences in his key or even mixed up features (see further) is unclear, but possibly he only had one single or a small number of O. lacustre specimens to compare with.
Ten years later,
The key to both species by
In a second stage of the verification process the type locality was considered to comprise a possible clue about the identity of O. lacustre. Scopoli collected the species in Carniola (‘Krain’ in German), a historical region that corresponds mainly with inland parts of present-day Slovenia, including mountains. Since Scopoli stated that the species skated on backwaters [“ambulat super aquas stagnantes”] and occurred in or along lakes [“in lacubus”], it could further be assumed that he collected the species in inland wetland habitats (and not along the coast).
To find out exactly where this type locality was situated within the distribution ranges of O. sabulosum and ‘species B’, capture locations of both species were plotted on a map of the West Palaearctic (see Figure
Order Diptera Linnaeus, 1758
Suborder Brachycera Macquart, 1834
Clade Eremoneura Lameere, 1906
Superfamily Empidoidea Latreille, 1804
Family Dolichopodidae Latreille, 1804
Subfamily Hydrophorinae Becker, 1917
Genus Orthoceratium Schrank, 1803 (monotypic)
Musca
lacustris
Scopoli, 1763: 343. Type locality: Carniola (= present-day Slovenia) – presumably transferred to Orthoceratium by
Musca formosa Haliday, 1832 and Medeterus viridipes Macquart, 1834, previously listed as synonyms of O. lacustre, clearly refer to O. sabulosum (see further).
Large, short-bodied, slender, entirely green species with abdomen 1.6 × as long as thorax (Figs
Male. Body length: 5.0–5.7 mm (n = 25); wing length: 5.2–6.2 mm (n = 44), 0.3 × as wide as long. Head (Fig.
ITALY: NEOTYPE (here designated to fix the identity of the species) ♂, [brownish rectangular] “Liancalus/ lacustris Scp”/ “Görz.” [= Gorizia, in Friuli-Venezia Giulia region]; “Zool. Mus.”/ “Berlin”; “Orthoceratium lacustre”/ “(Scopoli, 1763)”/ “det. Marc Pollet, 2017”; [red rectangular] “NEOTYPE” / “des. Marc Pollet, 2018” (2017 on initial label in Figure
See Suppl. material
In order to fix the identity of the species, a neotype of O. lacustre was selected on the basis of the locality (closest to the original type locality or region), and the preservation status of the specimen (see Figure
Of all examined specimens (n = 131) one Trieste specimen featured a strong curved black bristle on the right fore coxa.
As a result of the taxonomic mix-up between both species in the past, previous distribution records of O. lacustre in the literature – many of which refer to O. sabulosum in reality – must be considered unreliable. Our present study revealed that O. lacustre has been collected nearly exclusively along the northern border of the Mediterranean basin (incl. adjacent islands), both in coastal habitats and inland (montane) habitats (see Figure
Previous records from Austria, Ireland, Madeira or Crimea (Ukraine) could not be verified due to a lack of specimens, but it is very likely that the Irish and Madeiran records refer to O. sabulosum (see Figure
With only two clear exceptions, O. lacustre has been recorded from mostly lowland locations in a 25 km zone along the Mediterranean coast, where it seems to occur along inland lakes which also corresponds with the description of the habitat of the type specimens (
Alloeoneurus
sabulosus
Becker, 1907: 112. Type locality: Tunis (Tunisia) [
Musca formosa Haliday, 1832: 356. Type locality: Cheshire (Great Britain) [unknown] syn. n.
Medeterus viridipes Macquart, 1834: 452. Type locality: Bordeaux (France) [unknown] syn. n.
Musca formosa Haliday, 1832 and Medeterus viridipes Macquart, 1834 were previously listed as synonyms of O. lacustre by
Diagnosis. Rather large, short-bodied, rather slender (but stouter than O. lacustre), brilliant green species with abdomen 1.3 × as long as thorax (Fig.
Male. Body length: 5.5–6.2 mm (n = 54); wing length: 5.2–6.1 mm (n = 77), 0.3 × as wide as long (n = 15) mm. Head (Fig.
LECTOTYPE (here designated to fix the identity of the species) ♂, TUNISIA: [Tunis governate] “Tunis904” / “Ujhelyi”, [bottom side] “X 26”; “Alloeoneurus” / “sabulosus Beck.” / “det. Becker”; [red rectangular] “Lectotypus”; “Zool. Mus.” / “Berlin” [
The original description of this species by
See Suppl. material
Compared to O. lacustre, O. sabulosum is much more widespread in the West Palaearctic and currently known with certainty from 14 countries, although it has not (yet) been collected in the northern part of the Mediterranean basin (see Figure
Based on the current records, the distribution range of O. sabulosum differs significantly from that of O. lacustre. Despite that, however, both species seem to display a surprisingly similar ecological amplitude. In northwestern Europe (from Great Britain over Belgium and the Netherlands to Germany and Denmark), it is confined to humid coastal habitats, with a strong preference for salt marshes and brackish marshes. In Belgium, the species has only been collected in sea-aster (Aster tripolium) vegetations, bordering shallow brackish to saltwater ponds (
1 | Coxa I with one strong curved black bristle at basal 1/3 (Figure |
O. sabulosum (Becker, 1907) |
– | Coxa I without a black bristle at basal 1/3 (Figures |
O. lacustre (Scopoli, 1763) |
Table
Considering the fair size of Orthoceratium – compared to other dolichopodid lineages, it remains surprising that key features of this genus have been overlooked by previous authors. The ventral process of the 5th tergite was only mentioned by
Even more worrisome is the fact that
Unlike
But most of all, it is remarkable that O. sabulosum has been ignored as a species since its description. Indeed, apart from its Tunis type locality and its inclusion in the
One wonders how this series of mistakes could happen and last for over 250 years? A first reason seems to be that no previous researcher made the effort to compare with the type specimens, not even in the post-World War II era. Next to the fact that the type of O. lacustre was lost, most dolichopodid workers seemed to believe that O. sabulosum was a strictly southern Mediterranean species. With the present study we clearly proved that this species has a considerably larger distribution range. A second equally important reason are the insignificant, even misleading, characters used in species descriptions and identification keys, and the mere copying of those keys by subsequent authors.
Some authors tend to build elaborate and detailed species descriptions (see e.g.,
Here lies the importance of type material that cannot be underestimated. Only after the examination of the types of O. sabulosum did we realize that researchers had been misidentifying O. lacustre for more than 2.5 centuries (and every correct identification might merely be considered as a fortunate coincidence). Indeed, only when the type locality of O. lacustre proved to be situated within the distribution range of ‘species B’, it became apparent that both species were conspecific. If this would not have been the case, and assuming that the face width would have been unreliable like the other four features used in the extant identification keys, O. lacustre might as well have rendered nomen dubium, and ‘species B’ might have been described as new. In this respect, the use of alternatives to physical type specimens deposited in a museum (and photos thereof) like e.g., field images of uncollected specimens as promoted by
For more than 2.5 centuries the Orthoceratium species that occurs in northwestern Europe has indifferently been considered as O. lacustre, while it was, in reality, O. sabulosum. This is all the more surprising since Orthoceratium species are among the larger and more conspicuous dolichopodid species in Europe. The main reasons for this continuous series of misinterpretations seemed to be the mere copying of keys by successive authors that contained misleading information, the omission of examining type specimens of O. sabulosum, and the loss of the type specimens of O. lacustre. The importance of type specimens and the examination thereof is stressed, as well as that of unequivocal, detailed, and well-illustrated descriptions to avoid this kind of taxonomic confusion.
First of all, we are most grateful to Gordon Ramel (UK) who provided the senior author with the Greek samples containing O. lacustre specimens. Curators Duncan Sivell (
List of (non-type) records of Orthoceratium