Procladius ferrugineus (Kieffer, 1918)

Procladius ferrugineus (Kieffer, 1918) View in CoL

Trichotanypus ferrugineus Kieffer, 1918 — Kieffer (1918a), Hungary, adult male, key.

Trichotanypus parvulus Kieffer, 1918 — Kieffer (1918a), Lithuania, adult male, adult female, description, illustration.

Trichotanypus choreus ( Meigen, 1804) — Goetghebuer (1921), Belgium, adult male, description, illustration.

Trichotanypus ferrugineus Kieffer, 1918 View in CoL — Kieffer (1924), adult male, adult female, key, description.

Trichotanypus fulvus Kieffer, 1924 View in CoL — Kieffer (1924), Germany, adult female, key, description.

Trichotanypus parvulus Kieffer, 1918 View in CoL — Kieffer (1924), adult male, adult female, key.

Trichotanypus profundorum Kieffer, 1924 — Kieffer (1924), Germany, adult male, key.

Trichotanypus ferrugineus Kieffer, 1918 View in CoL — Goetghebuer (1927), France, adult male, illustration, key, description.

Trichotanypus parvulus Kieffer, 1918 — Goetghebuer (1927), France, adult male, adult female, key, description.

Procladius ferrugineus (Kieffer, 1918) View in CoL — Goetghebuer & Lenz (1936a), Goetghebuer & Lenz (1936b), Hungary, adult male, adult female, key, description, illustration.

Procladius fulvus ( Kieffer, 1924) View in CoL — Goetghebuer & Lenz (1936a), Goetghebuer & Lenz (1936b), Belgium, adult female, key, description, illustration.

Procladius parvulus (Kieffer, 1918) View in CoL — Goetghebuer & Lenz (1936a), Goetghebuer & Lenz (1936b), Lithuania, adult female, key, description, illustration.

Procladius profundorum ( Kieffer, 1924) — Goetghebuer & Lenz (1936a), Germany, adult male, key, description.

Procladius cf. nigriventris ( Kieffer, 1924) View in CoL — Brundin (1949), Sweden, adult male, pupa in part, key, description, illustration.

Procladius ferrugineus (Kieffer, 1918) View in CoL — Muragina-Koreneva (1957), Russia, adult male, adult female, pupa, larva, keys, descriptions, illustrations.

Procladius ferrugineus (Kieffer, 1918) — Shilova (1969), Russia, adult male, key, illustration.

Procladius ferrugineus (Kieffer, 1918) — Izvekova (1973), Russia, adult male, illustration.

Procladius ferrugineus (Kieffer, 1918) — Pankratova (1977), Russia, larva, illustration.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Pinder (1978), England, adult male, key, illustrations.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Fittkau & Murray (1986), pupa, illustrations.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Langton (1991), pupa, key, illustrations.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL —Sasa & Kikushi (1995), Japan, adult male, key, illustration.

Procladius choreus ( Meigen, 1804) View in CoL — Kobayashi (1998), Japan, adult male in part, description, photo.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Langton & Visser (2003), pupa, key, illustrations.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Langton & Pinder (2007), England, adult male, key, illustrations.

Procladius choreus ( Meigen, 1804) View in CoL — Ratnasingham et al. (2024), China, adult male.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Ratnasingham et al. (2024), Germany, Montenegro, Poland, Sweden and China, adult males, adult females, photos.

Procladius ferrugineus (Kieffer, 1918) View in CoL — Ratnasingham et al. (2024), Germany and Canada, adult males, adult females, photos.

Procladius sp. — Ratnasingham et al. (2024), Austria, Finland, Germany, Montenegro, Norway, Russia, Slovakia, China, Kazakhstan, Canada and United States, adult males, adult females, larvae, photos.

Procladius rugulosus Saether, 2010 View in CoL — Saether (2010), Canada, adult male, description, illustration.

Procladius crassinervis ( Zetterstedt, 1838) View in CoL — Langton et al. (2013), pupa, key, illustrations.

? Tanypus distans Kieffer, 1909 — Kieffer (1909), Germany, adult male, key, description.

? Trichotanypus distinguendus Kieffer, 1915 — Kieffer (1915c), Germany, adult male, adult female, description.

? Trichotanypus distans Kieffer, 1909 — Kieffer (1918a), Lithuania, adult male, key, description.

? Trichotanypus distinguendus Kieffer, 1909 — Kieffer (1918a), Lithuania, adult male, key, description.

? Trichotanypus distans Kieffer, 1909 — Kieffer (1924), adult male, key.

? Trichotanypus distinguendus Kieffer, 1915 — Kieffer (1924), adult male, adult female, key.

? Trichotanypus distinguendus Kieffer, 1915 — Goetghebuer (1927), France, adult male, adult female, key, description.

? Procladius distans ( Kieffer, 1909) — Goetghebuer & Lenz (1936a), Germany, adult male, key, description.

? Procladius distinguendus (Kieffer, 1915) — Goetghebuer & Lenz (1936a), Germany, adult male, key, description.

? Procladius distans ( Kieffer, 1909) — Fritz (1983), Germany, adult male, illustration.

? Procladius rugulosus Saether, 2010 View in CoL — Saether (2010), Canada, adult female, key, description, illustration.

Material examined (n = 56). AUSTRIA, 1 adult male (as P. choreus, ZSMG ), Lake Lunzer Untersee , 47.85°N 15.05°E, 608 m a.s.l., +6 °C m.a.t., 1940‒1942, leg. F. Krüger and A. Thienemann GoogleMaps ; 3 adult males (as P. choreus, IZUW ), Lake Lunzer Untersee , 47.84°N 15.04°E, 608 m a.s.l., +6 °C m.a.t., 1940‒1942, leg. F. Krüger. GoogleMaps — CANADA, 1 adult male (Paratype of P. rugulosus, FWIM ), Lake Winnipeg, Gimli , 50.63°N 96.96°W, 217 m a.s.l., +1 °C m.a.t., 14.vii.1971, leg. P.S.S. Chang GoogleMaps ; 2 adult males (as P. rugulosus, FWIM ), Lake Winnipeg, Gimli , 50.62°N 96.96°W, 217 m a.s.l., +1 °C m.a.t., 5.viii.1971, leg. N. Hooper and P. Johnson. GoogleMaps — CZECHIA, 3 adult males ( NHRS), Lake Pomezsky rybnik, Pomezy , 49.72°N 16.29°E, 565 m a.s.l., +6 °C m.a.t., 17.vi.1991, leg. Y. Brodin. GoogleMaps — ENGLAND, 2 adult males (as P. crassinervis, BMNH ), Hawkshead, Three Dubs Tarn , 54.37N 2.96W, 218 m a.s.l., +7 °C m.a.t., 7‒15.v.1947, leg. T. T. Macan GoogleMaps .— ESTONIA, 1 adult male ( EAST), Lake Peipsi-Pihkva , 58.83°N, 26.96°E, 28 m a.s.l., +5 °C m.a.t., 16.vii.1980, leg. V. Timm and T. Timm GoogleMaps .— FINLAND, 2 adult males (as P. culiciformis -t), Turku, Island Ruissalo , 60.4°N 22.2°E, 8 m a.s.l., +5 °C m.a.t., 29.vii‒5.viii.2009, leg. L. Paasivirta. GoogleMaps — FRANCE, 1 adult male (as Procladius sp. , NHRS), Corsica, Lake Melu , 42.21°N 9.02°E, 1 715 m a.s.l., +6 °C m.a.t., 25.viii.2015, leg. J. Moubayed-Breil. GoogleMaps — GERMANY, 1 adult male (as P. parvulus, ZSMG ), Eider , small stream, 54.31°N 9.96°E, 2 m a.s.l., +10 °C m.a.t., 1950 GoogleMaps ; 1 adult male (as Procladius sp. , ZSMG), Eider Achterwehr, canal, Silo , 54.3°N 9.9°E, 1 m a.s.l., +10 °C m.a.t., 29.vii.1969 GoogleMaps ; 1 adult male ( ZFMK), Rügen Island, Kniepow , small lake, 54.35°N 13.35°E, 9 m a.s.l., +9 °C m.a.t., 16.viii.2014 [Barcode GBMTM2155-15 ] GoogleMaps ; 1 adult male ( ZFMK), east of Karlsruhe, Wilhelma , 48.98°N 8.43°E, 121 m a.s.l., + 10 m GoogleMaps .a.t., 8.x.2014 [Barcode GBMTB294-15 ] .— ITALY, 1 adult male (as Procladius sp. , ZSMG), Lake Lago di Garda , 45.8°N 10.8°E, 65 m a.s.l., +13 °C m.a.t., 23.viii.1966, leg. F. Reiss. GoogleMaps — JAPAN, 2 adult males (as P. choreus, KUHY ), Lake Ikuta , 35.3°N E 139.3°E, 60 m a.s.l., +16 °C m.a.t., vi.1991, leg. T. Kobayashi GoogleMaps .— LITHUANIA, 1 adult male ( IBIB), Lake Vistytis , 54.43°N 22.75°E, 168 m a.s.l., +6 °C m.a.t., vii.1983, leg. A.I. Shilova. GoogleMaps — NETHERLANDS, 1 adult male (as P. crassinervis ), Haarlem, Tuinbouwgebied-zuid , 52.38°N 4.60°E, 3 m a.s.l., +11 °C m.a.t., 26.iii.2023, leg. M. van Wieringen. GoogleMaps — POLAND, 3 adult males ( NHRS), west of Dzwirzyno, Lake Resko Przymoirskie , 54.15°N 15.34°E, ‒ 2 m a.s.l., +9 °C m.a.t., 8.viii.2012, leg. Y. Brodin and J. Hellberg GoogleMaps [1 Barcode BSCHI697-17 ] .— RUSSIA, 5 adult males ( NHRS), Lake Shchuch , 55.75°N 32.18°E, 174 m a.s.l., +5 °C m.a.t., vii.1975, leg. P.I. Persson GoogleMaps ; 3 adult males (as P. grp. ferrugineus, TIEV ), Kuybyshev Reservoir , 53.6°N 49.0°E, 47 m a.s.l., +6 °C m.a.t., vii.1977 GoogleMaps ; 3 adult males, Lake Onega, Petrozavodsk , 61.83°N 34.32°E, 33 m a.s.l., +4 °C m.a.t., 26.vii.1989 GoogleMaps .— SPAIN, 2 adult males (as Procladius sp. , DEBE), Cordoba district, La Brena Reservoir , 37.51°N 5.03°W, 121 m a.s.l., +18 °C m.a.t., 23.xi.1974, leg. N. Prat GoogleMaps ; 1 adult male (as Procladius sp. , DEBE), La Minilla Reservoir, Sevilla , 37.44°N 6.10°W, 165 m a.s.l., +17 °C m.a.t., 27.xi.1974, leg. N. Prat. GoogleMaps — SWEDEN, 1 adult male ( NHRS), Marsveden, Lake Hålsjön , 59.81°N 17.24°E, 39 m a.s.l., +6 °C m.a.t., 17.v.1988, leg. J. de Jong GoogleMaps ; 2 adult males ( NHRS), Baltic Sea, Furusundsfjärden islands , 59.70°N 18.90°E, 0 m a.s.l., +5 °C m.a.t., 15.viii.2008, leg. C. Essenberg GoogleMaps [ Barcode BSCHI083-11 and BSCHI084-11 ] ; 3 adult males ( NHRS), Baltic Sea, Askö Island, west of Askötorp , 58.81°N 17.67°E, 5 m a.s.l., +6 °C m.a.t., 19.vii‒2.viii.2011, leg. B.-E. Bengtsson GoogleMaps [ Barcode BSCHI563-17 , BSCHI570-17 and BSCHI570-17 ] ; 2 adult males ( NHRS), Baltic Sea coast, Bönan , 60.74°N 17.32°E, 0 m a.s.l., +5 °C m.a.t., 15.vii.2011, leg. Y. Brodin GoogleMaps [ Barcode BSCHI222-17 and BSCHI223- 17 ] ; 1 adult male ( NHRS), Baltic Sea coast, Gävle harbor, 60.68°N 17.19°E, 2 m a.s.l., +5 °C m.a.t., 15.vii.2011, leg. Y. Brodin GoogleMaps [ Barcode BSCHI256-17 ] ; 1 adult male ( NHRS), Baltic Sea coast, Östra Sikvik , 60.67°N 17.29E °, 0 m a.s.l., +5 °C m.a.t., 15.vii.2011, leg. Y. Brodin GoogleMaps [ Barcode BSCHI368-17 ] ; 1 adult male ( NHRS), Lake Ottnaren, Tummen , 60.50°N 16.58°E, 67 m a.s.l., +5 °C m.a.t., 22.vii.2011, leg. Y. Brodin GoogleMaps [ Barcode BSCHI520-17 ] ; 1 adult male ( NHRS), Baltic Sea, Norrfjärden Bay, Axmar bridge, 61.05°N 17.16°E, 0 m a.s.l., +5 °C m.a.t., 25.vii.2012, leg. Y. Brodin GoogleMaps [ BSCHI629-17 ] ; 2 adult males ( NHRS), Ljusne harbor, Storstenshavet , 61.20°N 17.15°E, 1 m a.s.l., +5 °C m.a.t., 25.vii.2012, leg. Y. Brodin GoogleMaps [ Barcode BSCHI584-17 and BSCHI585-17 ] ; 1 adult male ( NHRS), Lake Mälaren, Vårby beach, 59.26°N 17.88°E, 4 m a.s.l., +6 °C m.a.t., 1.viii.2012, leg. Y. Brodin. GoogleMaps

Diagnostic characters. Figs. 30 View FIGURES 28‒31 , 77‒79 View FIGURES 77‒79 , key couplet 12. The unique form of the inner section of the phallapodeme (not to 30° angled from the outer section, dark and basally as broad as outer section) is usually enough to separate P. ferrugineus from all other Procladius in Europe and North America.

In some specimens of P. ferrugineus mounted on slides after maceration or longtime storage in alcohol, the phallapodeme is more or less deformed, bleached or not distinctly visible. In such cases it can sometimes be difficult to distinguish P. ferrugineus from other species with a medium long gonostylus process and overlapping GspR.

There are seven species with a GspR that overlaps that of P. ferrugineus . Of these, P. lugubris , P. frigidus , P. pruinosus and P. longistilus can mostly be distinguished from P. ferrugineus by characters related to size (wing length 2.4‒4.0 mm versus 1.9‒2.7 mm and mid tibia length 1.05‒1.52 mm versus 0.78‒1.07 mm) if the phallopodeme is in less good condition.

The gonostylus GspR of P. ferrugineus slightly overlaps that of P. crassinervis (0.27‒0.33 versus 0.32‒0.39). P. ferrugineus can nevertheless be separated from P. crassinervis by the not to only slightly upward oriented (0‒20° versus 30–60°) and less diverging (10‒35° versus 40–60°) gonostylus process.

P.ferrugineus can be difficult to separate from P. tenebricosus and P. floralis if the phallapodeme of P.ferrugineus is deformed or bleached. The gonostylus process of P. ferrugineus is on average somewhat longer than that of P. tenebricosus but overlapping (0.27‒0.33 versus 0.23‒0.29), and the anal cell of the wing has no or a faint dark patch while the patch is usually distinct in P. tenebricosus . The gonostylus process divergence of P. ferrugineus is mostly less strong than that of P. floralis (10‒35° versus 30–50°) and the mid leg tibia mostly somewhat shorter (0.78‒1.07 mm versus 0.99–1.30 mm).

P. ferrugineus has often been misidentified as P. crassinervis for records in western Europe which are based on the identification keys for males by Pinder (1978) or Langton & Pinder (2007). Pupal exuviae are also often misidentified as P. crassinervis .

P. ferrugineus is known under several other species names of which four to six can be regarded as synonyms.

One of the synonyms, P. rugulosus ( Saether, 2010) , was described as a new species particularly because of the rugulose or striped medial dorsal surface of the gonocoxite. This character is, however, not uncommon in recently emerged males of several species of Procladius .

The adult female and larva of P. ferrugineus have been briefly described, and the pupal exuvia more detailed. Barcodes of adult males, adult females and larvae are available.

Geographical distribution and ecology. With respect to ecology and geographical distribution, P. ferrugineus is the most studied species of Procladius in the world together with P. choreus . Comprehensive ecological information is available, particularly from Russia. Findings in Europe are from at least 31 countries or autonomous regions with the southernmost in Portugal and Spain at 37°N and the northernmost in mid Russia and Norway at 63°N. Records more southward are from Japan and the United States at 35°N. The mean annual temperature at the 121 sites with quality assured records of P. ferrugineus ranges from + 18 in southern Europe to + 1 in northern Europe, and within this temperature interval also for countries outside Europe.

P. ferrugineus has been recorded from the subtropical to the northern temperate zone including dense forests, steppes and semi-deserts, the latter in southern Russia and southern Spain. Most records are from ‒ 2 m below sea level to 600 m above sea level. A few findings higher than 1 000 m above sea level are known, with a top altitude at 1 720 m above sea level in a lake in Corsica, France.

About 90% of the findings of P. ferrugineus are from brackish water of the sea or freshwater lakes and lake-like reservoirs. Most of these habitats are eutrophic or hypereutrophic. The species has been acknowledged as an important indicator of oxygen deficit in deep areas of strongly eutrophic reservoirs and lakes. In fact, P. ferrugineus together with the chironomid Chironomus plumosus are frequently the only remaining macroscopic animals in conditions with very low oxygen concentrations (1‒3% oxygen saturation) such as in the profundal of hypereutrophic reservoirs and lakes. Calculations have shown that the larvae of P. ferrugineus are frequently among the most abundant macroscopic animal species in nutrient-rich environments, reaching up to 2 500 individuals per m 2 in some reservoirs in Russia. Larvae have been caught from 2‒60 m water depth, mostly above mud bottoms but sometimes also above sand-dominated ones. Some findings of P. ferrugineus are from saltwater lakes, such as the shrinking Lake Aral in Kazakhstan and Uzbekistan with salinity of about 20‰.

Larvae of P. ferrugineus are also common in oligotrophic and mesotrophic lakes and reservoirs, but usually in relatively low numbers. No confirmed findings seem to exist from mesohumic to polyhumic conditions. A few findings are from slowly running water including streams, ditches and manmade canals.

Food items of P.ferrugineus include detritus, algae and smaller animals such as crustaceans, worms ( Tubificidae ), Entomostraca, Testacea and other species of Chironomidae .

Adults are found from early May to the middle of October in northern Europe and until late November in Spain in southern Europe. One generation per year is mostly the case for northern Europe, whereas in mid and southern Europe two or three generations per year have been estimated. Warmer water produces adults which on average are smaller than those growing up in colder water.

Countries or autonomous regions with records of P. ferrugineus in Europe are Albania, Austria, Belgium, Bulgaria, Czechia, Denmark, England, Estonia, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Lithuania, Montenegro, Netherlands, Northern Ireland, Poland, Portugal, Romania, Russia, Scotland, Serbia, Slovakia, Spain, Sweden, Switzerland, Ukraine and Wales. It has also been recorded from Armenia, Azerbaijan, Canada, China, Iran, Japan, Kazakhstan, Mongolia, Russia (Asia), the United States and Uzbekistan.

References. Alecsevnina 1988; Alecsevnina 1989; Aliyev et al. 2013; Anikina 2009a; Anikina 2009b; Aydin & Samin 2020; Bazzanti & Seminara 1987; Berczik 1966; Bérg et al. 1962; Botnariuc & Cure 1999; Brodin et al. 2013; Brundin 1949; Carter 1976; Carter 1980a; Carter 1980b; Charles et al. 1974; Charles et al. 1976; Coe 1950; Erbaeva 1989; Erbaeva & Safronov 2016; Essenberg 2009; Fedoruk 1964; Fittkau & Murray 1986; Fritz 1983; Gadawski 2020; Gadawski et al. 2022; Gerstmeier 1989; Goetghebuer 1921; Goetghebuer 1927; Goetghebuer & Lenz 1936a; Goetghebuer & Lenz 1936b; Hempel 2011; Izvekova 1973; Izvekova 1975; Janković 1972; Jónasson 1964; Jónasson 1969; Jónasson & Thorhauge 1976; Kajak 1980; Kieffer 1909; Kieffer 1915c; Kieffer 1918a; Kieffer 1919; Kieffer 1924; Kobayashi 1998; Kobayashi 2000; Kontula & Haldin 2012; Kownacki 1963; Krupa & Grishaeva 2019; Langton 1991; Langton 1993; Langton & Pinder 2007; Langton & Visser 2003; Larsen 1993; Mitropoliskii 1978; Mothes 1966; Moubayed-Breil et al. 2019; Muragina-Koreneva 1957; Murray & Ashe 1983; Murray & O´Connor 1992; Murray et al. 2018; Nazarova 2006; Nietzke 1937; Ntislidou et al. 2019; Orendt 1993; Ostrovsky 1984; Pankratova 1977; Petrova & Zhirov 2014; Pinder 1978; Prat 1979; Prat 1980; Ratnasingham & Hebert 2007; Ratnasingham et al. 2024; Reiss 1968; Roback 1971; Rozenberg & Vykhristiuk 2008; Saether 2010; Sasa & Kikushi 1995; Sergeeva & Anikina 2009; Shcherbina 1989; Shilova 1969; Shilova 1976; Sokolowa 1968b; Sokolowa 1971; Stasiukynas et al. 2024; Tarwid 1969; Tatole 2023; Thienemann 1954; Timm et al. 1994; Timm et al. 1996b; Timm et al. 2007; Vadadi-Fülöp et al. 2007; Vesterinen et al. 2013; Vesterinen et al. 2018; Wesenberg-Lund 1943; Wiederholm 1984; Wonglersak et al. 2021; Zinchenko 1992.