taxonID	type	description	language	source
5E47CA08FFB009123CE4FB2F9EE3FB76.taxon	description	The family Chironomidae is usually divided into eleven subfamilies: Aphroteniinae, Buchonomyiinae, Chilenomyinae, Chironominae, Diamesinae, Orthocladiinae, Podonominae, Prodiamesinae, Tanypodinae, Telmatogetoninae, and Usambaromyiinae (Saether 2000; Bánki et al. 2024). Procladius is placed among the Tanypodinae. The phylogenetic relationship of Tanypodinae worldwide outlined by Silva & Ekrem (2016) recognized the tribe Procladiini containing the genera Procladius, Djalmabatista, Laurotanypus, Lepidopelopia and Saetheromyia. The study was based on DNA barcodes and morphology of adults, pupae and larvae. Only Procladius is present in Europe.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFB009123CE4FB2F9EE3FB76.taxon	description	1. Wing crossvein MCu absent. Subfamilies Aphroteniinae, Chironominae, Orthocladiinae, Telmatogetoninae and Usambaromyiinae - Wing crossvein MCu present (Fig. 5) ..................................................................... 2. 2. Wing vein R 2 + 3 absent. Subfamilies Buchonomyiinae, Chilenomyinae, Podonominae, some Tanypodinae - Wing vein R 2 + 3 present (Fig. 5) .......................................................................... 3. 3. Antenna with 5 ‒ 13 flagellomeres, last flagellomere longer than penultimate flagellomere. Subfamilies Diamesinae and Prodiamesinae - Antenna with 14 flagellomeres, last flagellomere more than three times shorter than penultimate flagellomere (Fig. 1). Subfamily Tanypodinae ......................................................................................... 4. 4. Leg fourth tarsomere broadened from base to end and shorter than fifth tarsomere. Hind leg tibial comb double. Coelopynia, Clinotanypus, Coelotanypus and Naelotanypus - Leg fourth tarsomere cylindrical and longer to slightly shorter than fifth tarsomere. Hind leg tibial comb simple (Fig. 10) or absent ............................................................................................... 5. 5. Wing vein Cu fork FCu (division into M 3 + 4 and CuA) proximal to opposite MCu. - Wing vein Cu fork FCu distal MCu (Figs. 5 and 6) ........................................................... 6. 6. Distance between MCu and FCu much less than half as long as M 3 + 4. - Distance between MCu and FCu at least half as long as M 3 + 4 (Figs. 5 and 6). Procladiini ........................... 7. 7. Wing with more the 20 macrotrichia. Gonostylus with or without inner lobe ....................................... 8. - Wing without or less than 5 macrotrichia. Gonostylus with distinct convex inner lobe with strong setae. Procladius subgenus Psilotanypus 8. Gonostylus with protruding convex inner lobe with strong setae ................................................. 9. - Gonostylus without distinct inner lobe, inner margin straight or concave (rarely slightly convex) without strong setae ..... 10. 9. Scutal tubercle absent. Abdomen tergites light and mostly with darker median longitudinal band. Djalmabatista - Scutal tubercle present. Abdomen tergites not with darker median longitudinal band. Procladius subgenus Laurotanypus 10. Scale-like setae on thorax, legs and wings. Wing vein R 2 + 3 not divided. Gonostylus without indication of outer process. Lepidopelopia - No scale-like setae. Wing vein R 2 + 3 divided into R 2 and R 3 (Fig. 5). Gonostylus with or without outer process ......... 11. 11. Gonostylus without indication of outer process. Scutal tubercle present. Saetheromyia Gonostylus with indicated to very long outer process (Figs. 17 and 18). Scutal tubercle absent. Procladius subgenus Holotanypus and Procladius Species delimitation and identification of European Procladius The iterative process comparing morphological data and barcoded data resulted in a list of 27 species of Procladius present in Europe (Table 6) of which four are new species to science, namely P. exilis Brodin, P. gemma Brodin, P. saeticubitus Brodin and P. tenebricosus Brodin & Hellberg. ...... continued on the next page Barcodes are available for 25 of the species. Table 6 shows that BOLD contained barcodes and BINs for 23 species. P. clavus Roback, 1971 and P. tatrensis have a few good barcodes of more than 550 base pars not yet incorporated into BOLD or GenBank. P. fimbriatus Wülker, 1959 and P. gemma lack known barcodes, but they are readily identified species based on several unique morphological characters. Each of the 42 BINs in Table 6 contains only one species of Procladius, but several species have more than one BIN. Several other studies of Chironomidae (Song et al. 2016; Chimeno et al. 2023) and other insects have shown that it is common that species have several BINs. The BINs for Procladius are constructed by barcodes of 6 414 specimens, ranging from 1 for P. clavus to 2 336 for P. dentus. The BINs contain specimens from 35 countries or autonomous regions. About 75 % of the barcoded specimens are from Canada. P. culiciformis and P. lugens Kieffer, 1915 have four BINs each, while the others have tree to one BIN each. Intraspecific distance ranges from 0 to 3.3 % (Table 6). Only P. ferrugineus (Kieffer, 1918) has an intraspecific distance exceeding that of the interspecific distance. Interspecific distance, expressed as distance to nearest neighbour in BOLD, ranges from 2.0 to 8.8 %. Six species have an interspecific distance below 3.3 %. Notably, five of these are among those most easily identified morphologically. The BIN distance between the sixth species P. tenebricosus and its nearest neighbour P. ferrugineus is only 2.0 %. The species are readily distinguished from each other by the form of the phallapodeme. Some species have BINs with the closest neighbour BIN not of morphologically very similar species but of species with very different morphology. The most striking example is P. appropinquatus (Lundström, 1916). with almost no gonostylus process and P. dentus with a very long gonostylus process. In addition to the 42 BINs thoroughly studied, there are 6 more BINs of Procladius in BOLD which contain at least one specimen from Europe. These BINs (BOLD: ABX 4068, BOLD: ACQ 5869, BOLD: ADA 6883, BOLD: ADA 8287, BOLD: AEE 2525, BOLD: AEE 7861) were not included in the analyses of the present study as they contain morphologically deformed males, contain no adult males, or lack adult males available for studies. Whether or not these BINs represent valid species of Procladius is an open question. Measurement data of the analyses of one hundred morphological characters (Supplement 1: https: // doi. org / 10.6084 / m 9. figshare. 28189610) showed that nine of the Procadius species can be identified by at least one non-overlapping character. Seven species can be identified by one or more non-overlapping genitalia characters, viz. the unique form of the gonocoxite (P. dentus), gonostylus (P. clavus and P. crassinervis), phallapodeme (P. ferrugineus) or medioapodeme (P. gemma, P. signatus and P. dentus). P. flavifrons Edwards, 1929 can be distinguished by the numerous setae on the katepisternum. P. fimbriatus is effortlessly identified by its femalized head and an antenna with a much lower AR than that of all other species. Reliable identification of each of the other eighteen species mostly requires measurements of several morphological characters. The most useful character for species identification is GspR, a ratio constructed as the gonostylus outer length divided by the gonostylus process length (Table 2 item G 22, Fig. 17). GspR is always one of the characters when more than one character is needed for species identification. An advantage is that GspR is usually easily measured for specimens prepared on slides or kept in alcohol. Other frequently useful morphological characters for separation of morphologically similar species are antenna AR (Table 2, item H 2, Fig. 1), macrotrichia on the median anepisternum (Table 2, item T 10, Fig. 4), front leg beard ratio (Table 2, item L 10, Fig. 9), wing length (Table 2, item W 1, Fig. 5), and thickness of the gonostylus measured as GsmR (Table 2, item G 23, Fig. 18). Species key to adult males of European Procladius Each step (couplet) in a key for morphological species identification should preferably be dichotomous and give two options for one morphological character. This approach is often not sufficient for distinguishing species of Procladius because of frequent and sometimes considerable overlaps in male characters. In the key below one to three characters are provided for major divisions, while three characters are always available for species separation. The characters of each division are ranked according to importance (I ‒ III) for species identification. Taking all three characters and the drawings of male genitalia (Figs. 20 ‒ 127) into account is often needed to achieve reliable species identification. Most characters have a code within parenthesis, e. g. (A 1), that is explained in Table 2 and illustrated in Figs. 1 ‒ 19. The key contains 28 of the 100 characters chosen for the present study. Before using the key, measurement of GspR, gonostylus process length divided by gonostylus outer length (G 22, Fig. 17) is recommended as this character forms the base of species identification. If the key does not result in satisfactory species identification, two additional instruments can be consulted. The helpdesk (heading below, Table 7) contains an overview of 21 morphological characters important for species separation. The subheadings “ Diagnostical characters ” and “ Geographical distribution and ecology ” for each species (below under heading “ The species; systematics, distribution and ecology ”) contains information particularly useful to separate morphologically similar species. The helpdesk and diagnostical characters are also useful for quality assurance of species determinations. characters of a species are marked in bold. The characters are explained in Table 2 and illustrated in Fig. 1 ‒ 19. ...... continued on the next page characters of a species are marked in bold. Characters are explained in Table 2 and illustrated in Fig. 1 ‒ 19. ...... continue on the next page ...... continued on the next page 1. I) Wing; membrane with more than 20 macrotrichia (W 3, Fig. 6). II) Genitalia; gonostylus inner margin slightly concave to slightly convex without strong setae (G 28, Fig. 17) ........................................................... 2. - I) Wing; membrane without macrotrichia or rarely 1 ‒ 5 at wing apex (W 3). II) Genitalia; gonostylus inner margin distinctly convex with strong setae (G 28, Fig. 113) .................................................................. 23. 2. I) Head; antenna AR 0.3 ‒ 0.6 (H 2). II) Genitalia; gonostylus process long GspR 0.34 ‒ 0.40 (G 22, Fig. 47). III) Genitalia; gonocoxite width 264 ‒ 311 µm (G 11, Fig. 16). Figs. 20, 47 ‒ 49. P. fimbriatus Wülker, 1959 - I) Head; antenna AR 1.3 ‒ 3.2 (H 2, Fig. 1). II. Genitalia; gonostylus process very short to very long GspR 0.02 ‒ 0.48 (G 22, Figs. 56 and 65). III) Genitalia; gonocoxite width 185 ‒ 532 µm (G 11, Fig. 16) ......................................... 3. 3. I) Genitalia; medioapodeme long and inner section strongly curved backwards 30 ‒ 105 ° (G 3, Fig. 14) .................. 4. - I) Genitalia; medioapodeme short and inner section not or slightly curved backwards 0 ‒ 10 ° (G 3, Fig. 15) ............... 7. 4. I) Genitalia; gonostylus process short GspR 0.10 ‒ 0.22 (G 22, Fig. 18) ............................................ 5. - I) Genitalia; gonostylus process long to very long GspR 0.36 ‒ 0.48 (G 22, Fig. 17) .................................. 6. 5. I) Genitalia; medioapodeme anterior margin undulating (Fig. 21). II) Abdomen; tergite IX hind mid margin with a ball-formed projection (A 7, Fig. 21). III) Genitalia; gonostylus process length / width 0.8 ‒ 1.1 (G 21, Fig. 50). Figs. 21, 50 ‒ 52. P. gemma Brodin, new species - I) Genitalia; medioapodeme anterior margin straight (Fig. 22). II) Abdomen; tergite IX hind mid margin with a semicircular projection (A 7, Fig. 22). III) Genitalia; gonostylus process length / width 0.5 ‒ 0.8 (G 21, Fig. 53). Figs. 22, 53 ‒ 55. P. clavus Roback, 1971 6. I) Genitalia; gonocoxite with a marked dorsal ridge (Fig. 23). II) Genitalia; medioapodeme mid with long anteriorly directed process (Fig. 23). III) Head; antenna AR 2.5 ‒ 3.0 (H 2). Figs. 14, 23, 56 ‒ 58. P. dentus Roback, 1971 - I) Genitalia; gonocoxite without a marked dorsal ridge (Fig. 24). II) Genitalia; medioapodeme mid without anteriorly directed process (Fig. 24). III) Head; antenna AR 1.9 ‒ 2.5 (H 2). Figs. 17, 24, 59 ‒ 61. P. signatus (Zetterstedt, 1850) 7. I) Genitalia; gonostylus process long GspR 0.35 ‒ 0.43 (G 22, Fig. 63). II) Wing; length 3.5 ‒ 4.5 mm (W 1). III) Thorax; median anepisternum with 7 ‒ 21 setae on each side (T 10). Figs. 11, 25, 62 ‒ 64. P. tatrensis Gowin, 1944 - I) Genitalia; gonostylus process very short to long GspR 0.02 ‒ 0.39 (G 22), if> 0.34 then II) Wing; length <3.5 mm (W 1) and III) Thorax; median anepisternum with 0 ‒ 6 setae on each side (T 10, Fig. 4) ....................................... 8. 8. I) Genitalia; gonostylus process slightly indicated to short GspR 0.02 ‒ 0.10 (G 22, Figs. 66 and 69). II) Genitalia; gonostylus process length / base width 0.1 ‒ 0.5 (G 21, Fig. 66). III) Genitalia; gonocoxite width 282 ‒ 451 µm (G 11, Fig. 16) ........... 9. - I) Genitalia; gonostylus process short to very long GspR 0.10 ‒ 0.39 (G 22). II) Genitalia; if gonostylus process length / base width <0.6 (G 21, Fig. 17) then III) Genitalia; gonocoxite width <265 µm (G 11, Fig. 16) ............................ 10. 9. I) Genitalia; gonocoxite width 377 ‒ 451 µm (G 11). II) Leg; hind leg each tibial comb with 8 ‒ 10 spines (L 18). III) Wing; length 3.7 ‒ 4.6 mm (W 1). Figs. 9, 26, 65 ‒ 67. P. simplicistilus Freeman, 1948 - I) Genitalia; gonocoxite width 282 ‒ 350 µm (G 11). II) Leg; hind leg each tibial comb with 11 ‒ 16 spines (L 18, Fig. 10). III) Wing; length 2.5 ‒ 3.7 mm (W 1). Figs. 27, 68 ‒ 70. P. appropinquatus (Lundström, 1916) 10. I) Leg; hind leg tibial comb with 6 ‒ 9 spines (L 18). II) Leg; tarsi of front leg with very long setae BRI 5.5 ‒ 8 (L 10, Fig. 9). III) Thorax; median anepisternum with 10 ‒ 18 setae on each side (T 10) ............................................. 11. - I) Leg; hind leg tibial comb with 9 ‒ 14 spines (L 18, Fig. 10), if <10 spines then II) Leg; tarsi of front leg with BRI <5 (L 10, Fig. 9). III) Thorax; median anepisternum with 0 ‒ 26 setae on each side (T 10, Fig. 4) .................................. 12. 11. I) Genitalia; gonostylus process GspR 0.23 ‒ 0.31 (G 22, Fig. 28). II) Wing; Cu stem vein before MCu without setae (W 11, Fig. 5). III) Head; palpomere five length / width 7.2 ‒ 8.5 (H 9). Figs. 28, 71 ‒ 73. P. lugubris (Zetterstedt, 1850) - I) Genitalia; gonostylus process GspR 0.14 ‒ 0.19 (G 22, Fig. 29). II) Wing; Cu stem vein before MCu with 4 ‒ 11 setae (W 11). III) Head; palpomere five length / width 10.7 ‒ 11.3 (H 9, Fig. 2). Figs. 29, 74 ‒ 76. P. exilis Brodin, new species 12. I) Genitalia; phallapodeme inner section about as dark and basally as broad as outer section (G 6, G 7, Fig. 30). II) Genitalia; phallapodeme mid not to moderately angled 0 ‒ 30 ° (G 5, Fig. 30). III) Genitalia; gonostylus process moderately long GspR 0.27 ‒ 0.33 (G 22, Fig. 77). Figs. 30, 77 ‒ 79. P. ferrugineus (Kieffer, 1918) - I) Genitalia; phallapodeme inner section lighter to much lighter and basally distinctly thinner than outer section (G 6, G 7, Fig. 15). II) Genitalia; phallapodeme mid moderately to strongly angled 25 ‒ 90 ° (G 5, Fig. 15). III) Genitalia; gonostylus process short to long GspR 0.10 ‒ 0.39 (G 22) ...................................................................... 13. 13. I) Genitalia, gonostylus process strongly oriented upwards 30 ‒ 60 ° (G 33, Fig. 19). II) Genitalia; gonostylus process strongly diverging 40 ‒ 75 ° (G 31, Fig. 80). III) Genitalia; gonostylus process long GspR 0.32 ‒ 0.39 (G 22, Fig. 80). Figs. 19, 31, 80 ‒ 82. P. crassinervis (Zetterstedt, 1838) - I) Genitalia; gonostylus process not to moderately oriented upwards 0 ‒ 25 ° (G 33, Fig. 19). II) Genitalia; gonostylus process not to strongly diverging 0 ‒ 50 ° (G 31, Fig. 83). III) Genitalia; gonostylus process short to long GspR 0.10 ‒ 0.37 (G 22) ....... 14. 14. I) Thorax; median anepisternum with 5 ‒ 26 setae (T 10). II) Head; palpomere five length / width 8.0 ‒ 9.3 (H 9). III) Genitalia; gonostylus process moderately long GspR 0.25 ‒ 0.33 (G 22, Fig. 83). Figs. 32, 83 ‒ 85. P. frigidus (Holmgren, 1869) - I) Thorax; median anepisternum with 0 ‒ 6 setae (T 10, Fig. 4). II) Head; palpomere five length / width 9.1 ‒ 12.3 (H 9, Fig. 2), if <9,4 then median anepisternum with <4 setae (T 10). III) Genitalia; gonostylus process short to long GspR 0.10 ‒ 0.37 (G 22). .................................................................................................. 15. 15. I) Genitalia; gonostylus process moderately to strongly diverging 30 ‒ 50 ° (G 31, Fig. 86). II) Body; length 3.6 ‒ 4.4 mm (B 1). III). Genitalia; gonostylus process rather long to long GspR 0.28 ‒ 0.37 (G 22, Fig. 86). Figs. 33, 86 ‒ 88. P. floralis Kieffer, 1915 - I) Genitalia; gonostylus process not to moderately diverging 0 ‒ 30 ° (G 31, Fig. 89), if> 25 ° then II) Body; length> 4.4 mm (B 1). III) Genitalia; gonostylus process short to long GspR 0.10 ‒ 0.34 (G 22) .......................................... 16. 16. I) Genitalia; gonostylus process medium long to long GspR 0.23 ‒ 0.34 (G 22, Fig. 89). II) Genitalia; gonostylus outer length / mid width GsmR 5.2 ‒ 6.9 (G 23, Fig. 89). III) Genitalia; gonostylus process length / width 1.2 ‒ 1.9 (G 21, Figs. 17 and 89) ... 17. - I) Genitalia; gonostylus process short to medium long GspR 0.10 ‒ 0.25 (G 22, Fig. 18), if> 0.23 then II) Genitalia; gonostylus outer length / mid width GsmR <5.2 (G 23, Fig. 107) and III) Genitalia; gonostylus process length / width <1.4 (G 21, Figs. 17 and 18) ................................................................................................ 19. 17. I) Wing; with distinct to very distinct dark patch in anal cell (W 7, Fig. 6). II) Leg; hind leg tibia length 0.84 ‒ 1.17 mm (L 16). III) Genitalia; gonostylus process medium to rather long GspR 0.23 ‒ 0.29 (G 22, Fig. 89). Figs. 6, 12, 34, 89 ‒ 91. P. tenebricosus Brodin & Hellberg, new species - I) Wing; with no or faint dark patch in anal cell (W 7, Fig. 5). II) Leg; hind leg tibia length 1.21 ‒ 1.71 mm (L 16). III) Genitalia; gonostylus process medium to long GspR 0.23 ‒ 0.34 (G 22) ................................................... 18. 18. I) Genitalia; outer section length / gonocoxite base width 0.34 ‒ 0.46 (G 10, Fig. 13). II) Genitalia; gonostylus process medium to long GspR 0.25 ‒ 0.34 (G 22, Fig. 92). III) Genitalia; gonostylus outer length / mid width GsmR 5.5 ‒ 6.9 (G 23, Fig. 92). Figs. 19, 35, 92 ‒ 94. P. longistilus (Kieffer, 1916) - I) Genitalia; phallapodeme outer section length / gonocoxite base width 0.27 ‒ 0.36 (G 10, Fig. 13). II) Genitalia; gonostylus process medium long to rather long GspR 0.23 ‒ 0.30 (G 22, Fig. 95). III) Genitalia; gonostylus outer length / mid width GsmR 5.2 ‒ 6.3 (G 23, Fig. 95). Figs. 5, 8, 16, 36, 95 ‒ 97. P. pruinosus (Kieffer, 1924) 19. I) Genitalia; gonostylus outer length / mid width GsmR 5.9 ‒ 6.9 (G 23, Figs. 18 and 98). II) Wing; Cu stem vein before MCu with 5 ‒ 33 setae (W 11). III) Genitalia; gonostylus process short to rather short GspR 0.14 ‒ 0.20 (G 22, Fig. 98). Figs. 37, 98 ‒ 100. P. saeticubitus Brodin, new species - II) Genitalia; gonostylus outer length / mid width GsmR 4.2 ‒ 6.4 (G 23, Figs. 18 and 101), if> 5.8 then II) Wing; Cu stem vein before MCu with 0 ‒ 4 setae (W 11, Fig. 5) and / or III) Genitalia; gonostylus process GspR> 0.20 (G 22, Fig. 101) ......... 20. 20. I) Genitalia; gonostylus outer length / mid width GsmR 5.3 ‒ 6.4 (G 23, Figs. 18 and 101). II) Leg; front leg BR 3 ‒ 6.5 (L 10, Fig. 9). III) Genitalia; gonostylus process rather short GspR 0.18 ‒ 0.24 (G 22, Fig. 101). Figs. 7, 15, 38, 101 ‒ 103. P. islandicus (Goetghebuer, 1931) - I) Genitalia; gonostylus outer length / mid width GsmR 4.2 ‒ 5.7 (G 23, Figs. 18 and 104), if> 5.2 then II) Leg; front leg BR <3 (L 10, Fig. 9) and / or III) Genitalia; gonostylus process GspR <0.18 (G 22, Fig. 104) ................................ 21. 21. I) Genitalia; gonostylus process short GspR 0.10 ‒ 0.16 (G 22, Fig. 104). II) Leg; mid leg tibia length 1.05 ‒ 1.26 mm (L 13). III) Genitalia; gonostylus process length / width 0.5 ‒ 0.8 (G 21, Figs. 18 and 104). Figs. 39, 104 ‒ 106. P. breviatus Remmert, 1953. - I) Genitalia; gonostylus process short to medium long GspR 0.13 ‒ 0.25 (G 22, Fig. 107), if <0.16 then II) Leg; mid leg tibia length <1.05 mm (L 13) and / or III) Genitalia; gonostylus process length / width> 0.8 (G 21, Figs. 17 and 107) ............ 22. 22. I) Genitalia; gonostylus process rather short to medium long GspR 0.18 ‒ 0.24 (G 22, Fig. 107). II) Genitalia; gonostylus outer length / mid width GsmR 4.2 ‒ 5.2 (G 23, Figs. 18 and 107). III) Body; length 4.4 ‒ 5.7 mm (B 1). Figs. 1, 3, 13, 40, 107 ‒ 109. P. culiciformis (Linnaeus, 1767) - I) Genitalia; gonostylus process short to rather short GspR 0.13 ‒ 0.20 (G 22, Fig. 110). II) Genitalia; gonostylus outer length / mid width GsmR 4.9 ‒ 5.6 (G 23, Figs. 18 and 110). III) Body; length 2.9 ‒ 4.4 mm (B 1). Figs. 2, 18, 41, 110 ‒ 112. P. choreus (Meigen, 1804) 23. I) Genitalia; gonostylus process short GspR 0.10 ‒ 0.19 (G 22, Fig. 113). II) Genitalia; gonostylus process length / width 0.4 ‒ 0.9 (G 21, Fig. 113) ...................................................................................... 24. - I) Genitalia; gonostylus process absent to very short GspR 0 ‒ 0.06 (G 22, Fig. 119). II) Genitalia; gonostylus process length / width 0 ‒ 0.4 (G 21, Fig. 119) ............................................................................ 25. 24. I) Genitalia; gonostylus outer length / mid width GsmR 2.6 ‒ 3.3 (G 23, Figs. 18 and 113). II) Genitalia; gonostylus outer length / gonocoxite length 0.42 ‒ 0.50 (G 25, Figs. 16 and 113). IIII) Genitalia; gonocoxite base width 206 ‒ 285 µm (G 11, Fig. 16). Figs. 42, 113 ‒ 115. P. bellus (Loew, 1866) ....................................................................... - I) Genitalia; gonostylus outer length / mid width GsmR 3.2 ‒ 3.9 (G 23, Figs. 18 and 116). II) Genitalia; gonostylus outer length / gonocoxite length 0.50 ‒ 0.59 (G 25, Fig. 16). IIII) Genitalia; gonocoxite base width 178 ‒ 216 µm (G 11, Fig. 16). Figs. 43, 116 ‒ 118. P. nudipennis Brundin, 1947 25. I) Thorax; katepisternum with 9 ‒ 18 setae on each side (T 11, Fig. 4). II) Head; antenna AR 1.3 ‒ 1.7 (H 2, Fig. 1). III) Wing; length 1.7 ‒ 2.3 mm (W 1, Fig. 5). Figs. 4, 44, 119 ‒ 121. P. flavifrons Edwards, 1929 - I) Thorax; katepisternum without setae (T 11). II) Head; antenna AR 1.7 ‒ 2.6 (H 2, Fig. 1). III) Wing; length 2.1 ‒ 3.4 mm (W 1, Fig. 5) ............................................................................................. 26. 26. I) Genitalia; gonostylus outer margin smoothly curved in basal section (Fig. 122). II) Leg; front leg tibia length 0.83 ‒ 1.17 mm (L 1). III) Head; antenna AR 1.7 ‒ 2.3 (H 2, Fig. 1). Figs. 45, 122 ‒ 124. P. lugens Kieffer, 1915 - I) Genitalia; gonostylus outer margin abruptly angled in basal section (Fig. 125). II) Leg; front leg tibia length 1.21 ‒ 1.39 mm (L 1). III) Head; antenna AR 2.2 ‒ 2.6 (H 2, Fig. 1). Figs. 10, 46, 125 ‒ 127. P. imicola Kieffer, 1922 Species identification helpdesk Consulting the key above might not always result in unambiguous species identification of European Procladius. Cases where two, or more rarely even three, species are possible may occur because of overlaps in measurements of morphological characters. The key might also have a limited functionality if diagnostic species characters of a specimen studied are obscured, deformed or completely lost, such as antennae or tarsomere setae for measurement of leg BR (Tabel 2, L 10). Table 7 functions as a helpdesk for species identification if the key is not sufficient. The helpdesk contains 21 important diagnostic characters, of which two are not in the key. The fictive example in Table 8 shows how Table 7 by adopting a stepwise procedure can be used for species identification of a difficult specimen. Usually 7 ‒ 10 steps may be enough, but in a few cases it might be necessary to go through all 21 steps to reach a conclusion. Table 7 may also serve as a general quality assurance tool for reliable species identification. A test including a hundred specimens indicates that the most problematic species separation issues are those of the “ species pairs ” P. choreus / P. culiciformis, P. choreus / P. islandicus, P. culiciformis / P. pruinosus, P. pruinosus / P. tenebricosus and P. imicola / P. lugens. TABLE 8. Stepwise procedure to reveal species identity of a fictive Procladius male. Step 1 a: Measure GspR of the fictive male, e. g. 0.20. Step 1 b: Mark all species in Table 7 A 1 with a GspR-interval covering 0.20. Step 2 ‒ 7: Mark agreement with fictive male e. g. with green and disagreement e. g. with red figures. Step 8 ‒ 21: Continue to Table 7 B ‒ C if necessary for reliable species identification. Here, P. islandicus is the most likely species as it is the only one with all figures marked green. Species Character G 22) W 3) wing G 23) A 2) tergite G 31) gonostylus W 1) wing T 10) gonostylus membrane gonostylus II ‒ IV posterior process length, mm anepisternum GspR setae GsmR colour divergence ° setae Step 1 Step 2 Step 3 Step 4 Step 5 Step 6 Step 7 fictive male 0.20 3 5.5 2 10 3.1 2 P. choreus 0.13 ‒ 0.20 2 ‒ 3 4.9 ‒ 5.6 0 ‒ 1 0 ‒ 15 1.8 ‒ 2.8 0 ‒ 2 P. saeticubitus 0.14 ‒ 0.20 3 5.9 ‒ 6.9 0.5 ‒ 2.5 0 ‒ 15 2.6 ‒ 3.4 0 ‒ 3 P. gemma 0.17 ‒ 0.22 2 5.4 ‒ 5.7 2 ‒ 2.5 25 ‒ 45 3.7 ‒ 4.3 0 P. islandicus 0.18 ‒ 0.24 2.5 ‒ 3 5.3 ‒ 6.4 1 ‒ 2.5 0 ‒ 15 2.6 ‒ 3.5 0 ‒ 3 P. culiciformis 0.18 ‒ 0.25 2 ‒ 3 4.2 ‒ 5.2 0.5 ‒ 2 0 ‒ 25 2.4 ‒ 3.5 0 Species identification of other than European Procladius Twelve species not known from Europe, but might be found there in the future, have also been studied using several characters of the 100 - character analysis (Supplement 1). The most important characteristics to identify these species are presented in Table 9. from Europe but possibly found there in the future. Characters are explained in Table 2 and illustrated in Fig. 1 ‒ 19.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFB009123CE4FB2F9EE3FB76.taxon	description	Geographical distribution and ecology The literature search identified about 3 560 papers with adequate information on Procladius. The search included comprehensive information in databases, particularly BOLD (Ratnasingham et al. 2024). Loans and contacts with museums, scientists and hobby collectors provided additional unpublished geographical and ecological information, as also information on slide labels, alcohol sample notes, and in computer files. Procladius is reported from 135 of the about 200 countries or autonomous regions of the world (Table 10). Canada, the United States, Finland, England, Scotland, Norway and Spain had most localities with findings. Records of Procladius could not be found from six (Afghanistan, Libya, Mali, Mauretania, Mozambique and Venezuela) of the 42 countries with land area exceeding 640 000 km 2. There are very few studies of Chironomidae species in these countries. Recent findings of Procladius are from New Zealand (Krosch et al. 2022), New Caledonia (Moubayed-Breil et al. 2021) and Vanuatu (Strandberg et al. 2023). The possibly most isolated findings are from the Azores (Murray et al. 2004; Raposeiro et al. 2009) in the Atlantic Ocean and Réunion (Freeman 1958) in the Indian Ocean. The northernmost findings of a Procladius species are from 82 ° N latitude on Ellesmere Island in Canada (Oliver 1963) and the southernmost findings seems to be from 46 ° S in southern Argentina (Massaferro et al. 2014). The geographical range of Procladius choreus stands out among the European Procladius species with reports from 58 countries or autonomous regions (Table 11), mostly from Europe including the remote Azores, but also a wide geographical belt stretching from the Canary Islands, to Egypt, Iran, India, Mongolia, Japan and the United States. P. choreus is the only of the European Procladius with quality assured findings in Africa. Other European species with wide geographical range covering Europe, Asia and North America are P. culiciformis and P. ferrugineus reported from 37 and 30 countries or autonomous regions, respectively. As much as 20 of the 27 European species of Procladius are also found in Asia and / or North America. Table 11 furthermore shows that six of the Procladius species of Europe reach above 2 000 meters altitude. P. tatrensis has the highest record of 2 670 m and might reach 2 800 m in the Alps (Heiri & Lotter 2008). Worldwide it is surpassed by P. brevipetiolatus which reaches altitude 3 500 m in Uganda (Eggermont & Verschuren 2007) and an unknown species of Procladius at 4 730 m in Himalayan China (Echeverría-Galindo et al. 2023). None of the European Procladius species is only confined to high altitude. All, but for P. fimbriatus with lowest record at 410 m, occur at altitudes of less than 80 m. The coldest conditions with findings of Procladius are on Ellesmere Island of northernmost Canada (Oliver 1963) and the New Siberian Islands of Russia (Lundström 1915; Krasheninnikov 2013; Nazarova et al. 2015) with mean annual temperature of about ‒ 14 to ‒ 18 ° C. Coldest conditions known for Procladius in Europe is ‒ 8 ° C mean annual temperature prevailing during the 1920 s on the Svalbard islands of northernmost Norway (Edwards 1924; Stur & Ekrem 2020). As many as 8 of the 27 identified species of Procladius in Europe have been found in areas with annual mean annual temperature as low as ‒ 10 to ‒ 18 ° C (Table 11). P. clavus (Roback 1971), P. exilis and P. frigidus (Lundström 1915; Oliver 1963; Saether 2004) reach the coldest records. Four other species of Procladius (P. desis, P. jeris, P. paragretis and P. prolongatus), all from North America, have also been found at sites with mean annual temperature from ‒ 10 ° C or colder. This means that Procladius, with 12 species, is one of the most species-rich genera of aquatic insects in extremely cold conditions, such as in Canada (Oliver 1963; Danks 1992; Pentinsaari et al. 2020), Greenland (Edwards 1931; Böcher et al. 2015), Russia (Lundström 1915; Saether 2004; Coulson et al. 2014) and Alaska of the United States (Butler et al. 1980; Butler 1982; Lougheed et al. 2011). All species in Europe can thrive in areas with a mean annual temperature below + 6 ° C and are thus able to withstand ice cover on lakes or smaller water bodies. Warmest conditions with records of Procladius worldwide are in Niger (Alhou et al. 2012) and the United Arab Emirates (Reeves & Epler 2016) with mean annual temperature about + 29 ° C. Table 11 shows that only P. choreus of the European species reach mean annual temperature above + 20 ° C, with the highest record at + 25 ° C in Saudi Arabia (Cranston & Judd 1989). Six species of Procladius species found in Europe have temperature ranges of more than 20 ° C, with the most extended range noted for P. lugens (Table 11). All species but four have ranges of 10 ‒ 20 ° C. Among these, three species, P. fimbriatus, P. gemma and P. saeticubitus, might be rare as they have been recorded from less than eleven localities each. Most reports of Procladius species are from lentic freshwater bodies of 0.01 km 2 or more, thus lakes and reservoirs. Records are rather frequent from other standing freshwater such as springs, ponds, puddles, marshes, swamps, bogs and human constructions such as rice fields, sewage water treatment plants, sewage ponds and rainwater barrels. Larvae of Procladius also frequently inhabit salty conditions such as estuaries, brackish sea water and even salt lakes with salinity higher than that of the sea. They are, however, not known from true marine environments. All 27 European Procladius species have been found in lakes and reservoirs, most of them also in other kinds of water (Table 12). Brackish and slightly marine water have representatives of nine species of European Procladius, while moderately fast running water only three. ...... continued on the next page All 25 species with water depth notes have been found in shallow water between 0 to 2 m, while findings beyond 100 m are only reported for P. lugubris and P. culiciformis, of which the latter has a probable record from 206 m in Lake Vierwaldstätter in Switzerland (Zschokke 1905). Table 12 furthermore shows that most Procladius species, as most other species of Chironomidae, have a wide adaptability to different trophic levels in lakes and reservoirs encompassing oligotrophic, mesotrophic and eutrophic conditions (Saether 1979; Ruse 2010). No Procladius species is, however, noted for all five categories of trophic conditions. Five species which mainly or solely inhabit lakes in northern Europe are only found in ultraoligotrophic or oligotrophic conditions, thus with low levels of available nutrient for primary production. Species of Procladius have proven useful as indicators of environmental problems caused by human activity, including eutrophication (Zinchenko 1992; Ruse 2015; Takamura et al. 2021), contamination by heavy metals or toxic organic compounds (Pettigrove 1989; Warwick 1991; Aliyev et al. 2013), climate-change effects (Engels et al. 2019; Rigterink et al. 2022, Brodin & Hellberg 2023) or biological diversity changes (Alhou et al. 2012; Bista et al. 2017; Gadawski et al. 2022). Indicator usefulness of Procladius is however often hampered by the fact that identification could only be achieved to the genus level. Species of Procladius are generally not among those reported to cause trouble for humans, e. g. because of mass-development or allergy problems. An unidentified species of Procladius is suspected to be a vector of cholera (Maheshwari et al. 2010). The species; systematics, distribution and ecology The 27 European species are presented in alphabetical order. After the heading with the species name and author follows in chronological order a list of Latin names, including synonyms, with pertinent information on literature, countries or autonomous regions with findings, life stages and available keys, illustrations and photos. At the end of the list, a question mark in front of the name indicates questionable synonym. Material examined is presented in the following way and order; 1) in alphabetic order, country or autonomous region in capital letters, 2) numbers and life stage, 3) within parentheses (type-status if it is a type, if another name is assigned to the specimen, code of four letters of the museum or institution where the specimen is deposited) (Table 13), 4) site with WGS 84 geographic coordinates, 5) altitude in meters above sea level abbreviated m a. s. l., 6) mean annual temperature abbreviated m. a. t. of the locality of the year of sampling, 7) date of sampling, 8) leg. collector name, 9) within brackets [barcode code]. Several specimens, sometimes also types, are in private collections and don’t have a museum or institution code. ...... continued on the next page Diagnostic characters: Reference to figures of the species and couplet of appearance in the key above are given. The most important morphological information to distinguish the species in question from those most similar ones is highlighted. Notes on taxonomy and the existence of barcodes are also provided. The helpdesk above contains twenty-one characters useful to distinguish the species from other species. Supplement 1 serves as a comprehensive description of the species based on one hundred characters. Geographical distribution and ecology: A synthesis based on important information from published literature and other sources including unpublished works and information only on slides, pins or of alcohol samples is given. Most of the information is also available in Table 11 and 12. Only information from sources where the species in question can be reliably identified is considered. References: This part contains a list of published scientific literature or other works with important information about the species in question, provided that species identification can be regarded as reliable, e. g. done be an expert on Chironomidae taxonomy. The species in question is identified with another name or only as Procladius sp. in several works, but the species identity is revealed by drawings, photos, specimen loans or correspondence by email.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF9909133CE4FAEF9CCBF88D.taxon	description	Procladius sp. B — Halvorsen et al. (1982), Norway, adult male, illustration.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF9909133CE4FAEF9CCBF88D.taxon	materials_examined	Material examined (n = 29). CANADA, 1 adult male (Paratype of P. ruris, CNCC), Banff Alta, 51.24 ° N 115.56 ° W, 1 383 m a. s. l., ‒ 1 ° C m. a. t., 25. vii. 1922, leg. C. B. D. Garrett; 1 adult male (Paratype of P. ruris, CNCC), Lake Cooking, 53.43 ° N 113.01 ° W, 732 m a. s. l., + 3 ° C m. a. t., 24. v. 1937, leg. F. O. Morrison; 1 adult male (Paratype of P. ruris, CNCC), Ottawa, 45.36 ° N 75.84 ° W, 70 m a. s. l., + 7 ° C m. a. t., 2. v. 1952, leg. J. E. Mitchell; 1 adult male (Holotype of P. ruris, CNCC), Lake Elkwater, 49.39 ° N 110.17 ° W, 1 234 m a. s. l., + 2 ° C m. a. t., 2. vi. 1955, leg. V. R. Vockeroth; 1 adult male (Paratype of P. ruris, CNCC), Lake Scout, 49.36 ° N 105.99 ° W, 830 m a. s. l., + 4 ° C m. a. t., 17. vi. 1955, leg. V. R. Vockeroth; 1 adult male (as P. cf. ruris, NTNU), Churchill town, 58.77 ° N 94.16 ° W, 8 m a. s. l., ‒ 6 ° C m. a. t., 21. vii. 2007, leg. J. McGovan [Barcode CHURC 487 - 08]; 1 adult male (as P. cf. ruris, NTNU), River Churchill, 58.68 ° N 94.17 ° W, 1 m a. s. l., ‒ 6 ° C m. a. t., 25. vii. 2007, leg. T. Ekrem [Barcode CHURC 507 - 08]. — FINLAND, 1 adult male, Lake Saanajärveniompolo, 69.04 ° N 20.88 ° E, 659 m a. s. l., ‒ 4 ° C m. a. t., viii. 1956, leg. P. Virtanen; 4 adult males (MZHF), Tvärminne, rockpools, 59.83 ° N 23.30 ° E, 0 ‒ 1 m a. s. l., + 5 ° C m. a. t., vii. 1957, leg. B. Lindeberg; 1 adult male (ZMUO), Enontekiö, Lake Toskaljärvi, 69.19 ° N 21.44 ° E, 704 m a. s. l., ‒ 5 ° C m. a. t., 1. viii. 2014, leg. L. Paasivirta; 2 adult males (ZMUO), Liminka, Liminganlahti, Liminka, 64.83 ° N 25.33 ° E, 1 m a. s. l., + 3 ° C m. a. t., 4. viii. 2015, leg. L. Paasivirta [Barcode CHIFI 412 - 16 and CHIFI 413 - 16]; 1 adult male (ZMUO), Termislehto, Enontekiö, 68.69 ° N 23.73 ° E, 436 m a. s. l., ‒ 2 ° C m. a. t., 15. vi. 2016, leg. L. Paasivirta. — NORWAY, 2 adult males (as P. sp. A and P. sp. B, ZMBM), River Ekse, 60.52 ° N 6.15 ° E, 580 m a. s. l., + 2 ° C m. a. t., 10 ‒ 23. viii. 1976, leg. G. A. Halvorsen; 1 adult male (as P. cf. ruris, NTNU), Sølandet, Kildebeck springbrook, 62.69 ° N 11.83 ° E, 785 m a. s. l., ‒ 1 ° C m. a. t., 11 ‒ 22. vi. 2006, leg. O. Hanssen [Barcode MIDGE 701 - 08]. — RUSSIA, 3 adult males (IANR), tundra ponds, east of Naryan-Mar, Yarey-shor, 67.62 ° N 53.70 ° E, 18 m a. s. l., ‒ 3 ° C m. a. t., 29. vii. 1990, leg. A. Rybakova. — SWEDEN, 1 adult male (Holotype of P. appropinquatus, NHRS), Sarek valley, Pelajauratjah, 67 ° N 18 ° E, 810 m a. s. l., ‒ 5 ° C m. a. t., 2. viii. 1907, leg. B. Poppius; 1 adult male (as P. simplicistilis, ZSMG), Norrby archipelago, Ängerån stream, 63.3 ° N 19.5 ° E, 1 ‒ 3 m a. s. l., + 3 ° C m. a. t., 1 ‒ 10. viii. 1981, leg. K. Müller; 1 adult male (NHRS), Baltic Sea, Sillviksskatan Bay, 63.76 ° N 20.46 ° E, 0 m a. s. l., + 3 ° C m. a. t., 27. vii. 2011, leg. N. Ericson [Barcode BSCHI 390 - 17]; 1 adult male (NHRS), Lake Kåbdalisjaure, Kåbdalis, 66.14 ° N 19.99 ° E, 330 m a. s. l., 0 ° C m. a. t., vii. 1993, leg. B. Viklund; 1 adult male (NHRS), Torne lappmark, 68 ° N 19 ° E, 650 m a. s. l., ‒ 1 ° C m. a. t., 1. vii. 2016, leg. H. Vårdal [Barcoded]; 2 adult males (NHRS), Lake Stor-Björsjön, 63.61 ° N 12.23 ° E, 566 m a. s. l., 0 ° C m. a. t., 16 ‒ 23. vi. 2019, leg. S. Persson [Barcoded]. Diagnostic characters. Figs. 27, 68 ‒ 70, key couplet 9. Male P. appropinquatus and P. simplicistilus have an only slightly indicated gonostylus process which separates them from all other European Procladius species with wing macrotrichia. The gonostylus process of P. appropinquatus is on average somewhat longer than that of P. simplicistilus (GspR 0.04 ‒ 0.10 versus 0.02 ‒ 0.06). P. appropinquatus is smaller than P. simplicistilus exemplified by wing length (2.5 ‒ 3.7 mm versus 3.7 ‒ 4.6 mm), body length (4.3 ‒ 6.0 mm versus 6.5 ‒ 8.1 mm) and gonocoxite width (282 ‒ 350 µm versus 377 ‒ 451 µm). The antenna AR-ratio of P. appropinquatus is mostly lower than that of P. simplicistilus (AR 1.8 ‒ 2.6 versus 2.5 ‒ 3.2). Hairiness is often also useful to distinguish P. appropinquatus from P. simplicistilus (scutellum 40 ‒ 73 setae versus 74 ‒ 117 setae, wing vein Cu stem 4 ‒ 36 setae versus 0 ‒ 4 setae). P. appropinquatus has often been identified as P. ruris in Canada. Studied males including types, both with and without barcodes, shows that P. ruris is a synonym. The adult female, pupa and larva have been described as P. ruris. Barcodes of adult males, adult females and larvae are available. Geographical distribution and ecology. P. appropinquatus has been found in northern Europe from Slovakia at latitude 49 ° N to northern Finland and Russia at latitude 69 ° N. It has been found from 45 ° N to 59 ° N in North America. The species has records from sites in boreal coniferous forests, subalpine mountain birch forests, taiga and completely treeless tundra. The 34 sites with findings have a mean annual temperature ranging from + 7 to ‒ 6 ° C, with the highest temperature at sea level in the Baltic Sea and the coldest in the arctic tundra. Highest altitude is at about 1 900 m above sea level in a high plain above the tree limit in northern Slovakia. Larvae of P. appropinquatus have been collected at water depth from 0 to 5 m in oligohaline to mesohaline brackish water of bays and rockpools in the Baltic Sea. In freshwater, findings have been reported from ponds, bog systems, shallow lakes, tarns and small streams with low velocity. All findings are from ultraoligotrophic to mesotrophic conditions. Adults have been caught from late May to mid-August. Countries with records of P. appropinquatus in Europe are Finland, Norway, Poland, Russia, Slovakia and Sweden. It is also known from Canada, Japan and the United States.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF97091E3CE4FF709CC7FC2D.taxon	description	Procladius profundorum Kieffer, 1923 — Kieffer (1923 b), Germany, adult male, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF97091E3CE4FF709CC7FC2D.taxon	description	? Procladius anomalus Kieffer, 1906 — Kieffer (1918 a), Lithuania, adult female, key, description.? Procladius anomalus Kieffer, 1906 — Goetghebuer (1927), France, adult female, key, description.? Psilotanypus anomalus Kieffer, 1906 — Goetghebuer & Lenz (1936 a), France, adult female, key, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF97091E3CE4FF709CC7FC2D.taxon	materials_examined	Material examined (n = 29). AUSTRIA, 1 adult male (as P. rufovittatus, ZSMG), Stift Schlägl, Böhmer Wald, Glashüttenteich pond, 48.64 ° N 13.97 ° E, 552 m a. s. l., + 8 ° C m. a. t., 2. vi. 1982, leg. H. Malicky. — CANADA, 1 adult male (Holotype of P. malifero, CNCC), Cranbrook, 49.50 ° N 115.79 ° W, 919 m a. s. l., + 5 ° C m. a. t., 20. v. 1925, leg. C. B. D. Garrett; 2 adult males (Paratypes of P. malifero, CNCC), Cranbrook, 49.50 ° N 115.79 ° W, 919 m a. s. l., + 5 ° C m. a. t., 20. v. 1925, leg. C. B. D. Garrett. — CZECHIA, 4 adult males (as P. rufovittatus, NHRS), Divcice, Lake Blatec, 49.11 ° N 14.31 ° E, 397 m a. s. l., + 7 ° C m. a. t., 19. vi. 1991, leg. Y. Brodin. — ENGLAND, 1 adult male (as P. rufovittatus, BMNH), Ruislip Lido Reservoir, 51.59 ° N 0.43 ° W, 50 m a. s. l., + 11 ° C m. a. t., 1948. — FINLAND, 2 adult males (as P. nudipennis, MZHF), Lake Puruvesi, Punkasalmi, Akonniemi, 61.74 ° N 29.35 ° E, 75 m a. s. l., + 3 ° C m. a. t., 10. viii. 1974, leg. B. Lindeberg; 2 adult males (as P. rufovittatus, ZMUO), Pihlava, Lake Enäjärvi, 61.55 ° N 21.61 ° E, 5 m a. s. l., + 4 ° C m. a. t., 4. vi. 2015, leg. L. Paasivirta [Barcode LEFIJ 3499 - 16 and LEFIJ 3500 - 16]. — GERMANY, 1 adult male (as P. crassinervis, ZSMG), Plön, Eutinerstrasse, 54.15 ° N 10.62 ° E, 18 m a. s. l., + 9 ° C m. a. t., vi. 1964, leg. E. J. Fittkau. — NETHERLANDS, 1 adult male (as P. rufovittatus), Almere-Verzetswijk, 52.37 ° N 5.24 ° E, ‒ 3 m a. s. l., + 10 ° C m. a. t., 14. v. 2022, leg. P. Hoekstra. — RUSSIA, 1 adult male (as P. rufovittatus, IBIB), Rybinsk Reservoir, 58.1 ° N 38.6 ° E, 97 m a. s. l., + 4 ° C m. a. t., viii. 1972, leg. A. I. Shilova. — SLOVAKIA, 5 adult males (as P. rufovittatus, NHRS), Lake Liptovska Mara, Liptovsky Mikulas, 49.10 ° N 19.57 ° E, 555 m a. s. l., + 6 ° C m. a. t., 19. vi. 1991, leg. Y. Brodin. — SWEDEN, 1 adult male (as P. rufovittatus, NHRS), Lake Boren, 58.58 ° N 15.13 ° E, 74 m a. s. l., + 6 ° C m. a. t., 10 ‒ 11. vi. 1980, leg. Y. Brodin; 1 adult male (as P. rufovittatus, NHRS), Lake Mälaren, Drottningholm, 59.33 ° N 17.87 ° E, 3 m a. s. l., + 6 ° C m. a. t., 26 ‒ 29. v. 1989, leg. T. Sandberg; 3 adult males (as P. rufovittatus, NHRS), Lake Grytsjö, 58.59 ° N 15.52 ° E, 96 m a. s. l., + 6 ° C m. a. t., 28. v. 1990, leg. Y. Brodin. — UNITED STATES, 2 adult males (Paratypes of P. riparius, INHS), Havana, Chautauqua Lake, 40.37 ° N 49.99 ° W, 147 m a. s. l., + 9 ° C m. a. t., 24. iv. 1914, leg. J. R. Malloch; 1 adult male (USNM), Edgerton, Lake Koshkanong, 42.86 ° N 89.00 ° W, 312 m a. s. l., + 8 ° C m. a. t., 9. viii. 1947, leg. H. S. Dybas. Diagnostic characters. Figs. 42, 113 ‒ 115, key couplet 24. The gonostylus with its distinctly convex inner margin in combination with a distinct outer process readily separates P. bellus and P. nudipennis from all other European Procladius. The gonostylus of P. bellus is comparatively broader (gonostylus GsmR 2.6 ‒ 3.3 versus 3.2 ‒ 3.9), more bulging and usually with more strong setae (6 ‒ 12 versus 4 ‒ 8) than that of P. nudipennis, while the gonostylus process is on average slightly shorter (gonostylus GspR 0.10 ‒ 0.17 versus 0.13 ‒ 0.19). P. bellus can usually also be separated from P. nudipennis by the relatively short gonostylus in relation to the length of the gonocoxite (0.42 ‒ 0.50 versus 0.50 ‒ 0.59). P. bellus is on average somewhat larger than P. nudipennis exemplified by an often broader gonocoxite base (206 ‒ 285 µm versus 178 ‒ 216 µm), wing length (2.0 ‒ 2.7 mm versus 1.8 ‒ 2.4 mm) and body length (3.5 ‒ 4.5 mm versus 3.1 ‒ 3.8 mm). Some light-coloured specimens of P. bellus are, as its name applies, regarded as nice-looking owing to a whitish to light yellowish scutellum, mid-section of tibia and abdomen tergites I ‒ V, contrasting with darker parts of the body. Oher specimens of P. bellus are as P. nudipennis more uniformly brownish. P. bellus has since about one hundred and fifty years been referred to as P. rufovittatus in Europe. Studied specimens from North America and Europe, in some cases also barcoded, reveals that P. rufovittatus is a synonym. The adult female, pupa and larva of P. bellus have been described in several papers. Barcodes of adult males, adult females and larvae are available. Geographical distribution and ecology. The southernmost finding of P. bellus in Europe is at latitude 47 ° N in France and the most northern at 63 ° N in Finland. Climate conditions in Europe range from + 11 ° C mean annual temperature in the south and to + 2 ° C in the north. It has a wider geographical range in the United States and Canada, namely from 39 ° N to 68 ° N, and with a temperature span from + 12 ° to ‒ 8 ° C on the Arctic coast of northern Canada. Findings in Europe are from altitude 3 m below to 560 m above sea level. The highest altitude records are from a lake in Canada at 1570 m and one in Mongolia at about 1 650 m. P. bellus larvae are mostly found in lakes and lake-like reservoirs. They are also reported from smaller stagnant water bodies such as ponds or puddles in peatlands, fens, gravel pits or sand dunes along the sea. A few records are from slow running water. In Europe and North America most findings are from mesotrophic to eutrophic conditions where the larvae can be among the dominant benthic species in terms of abundance and biomass, e. g. 1 600 individuals and 2 g dry weight per m 2. Larvae have less frequently been recorded from oligotrophic lakes and hypereutrophic lakes or reservoirs. P. bellus larvae have been found in the littoral zone at 0 ‒ 4 m in lakes with vegetation dominated by plants such as Elodea and Myriophyllum. They have been frequently found in the profundal and emerging from 3 to 15 m. The larvae have been reported to have a diapaus during summer stagnation with strong oxygen shortage in deeper zones of eutrophic or hypereutrophic lakes and reservoirs. A kind of diapause with very little growth is also reported to occur during cold winter months when the larvae mostly are at their 3 rd stage, but some also in their 2 nd or last 4 th stage. Gut content studies of P. bellus indicate that the larvae are detritivore or omnivore and consume algae, bacteria and animals. The amount of animals remains increases in the final fourth instar. Adult P. bellus are known from late April to August and sometimes also until late September. One to three generations per year have been reported in Europe. Emergence may sometimes take place during daytime but mainly during night with a peak at dawn. Ascending pupae can emerge to adults immediately after they reached the water surface. Adult P. bellus have been found to feed on honeydew from aphids. Countries or autonomous regions with records of P. bellus in Europe are Austria, Czechia, Denmark, England, Estonia, Finland, France, Germany, Ireland, Latvia, Netherlands, Northern Ireland, Poland, Russia, Scotland, Slovakia, Sweden and Wales. The species has also been found in Canada, Mongolia and the United States.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF9509183CE4FBD99C75FD45.taxon	materials_examined	Material examined (n = 27). ENGLAND, 1 adult male (as P. sagittalis), Sandscale Haws, 54.16 ° N 3.24 ° W, 0 m a. s. l., + 9 ° C m. a. t., 18. iv. 1964. — FINLAND, 1 adult male, Baltic Sea, Storskär Island, Lake Käringsund, 63.43 ° N 21.07 ° E, 0 m a. s. l., + 4 ° C m. a. t., 30. v ‒ 13. vi. 2017, leg. L. Paasivirta. — GERMANY, 1 adult male (as P. sagittalis, ZSMG), Odesloe, Brenner Moor mire and saline ponds, 53.8 ° N 10.4 ° E, 7 m a. s. l., + 10 ° C m. a. t., 14. vi. 1923, leg. A. Thienemann; 2 adult males (Paratypes of P. breviatus, ZSMG), Baltic Sea, Howacht Bay, 54.32 ° N 10.67 ° E, 0 m a. s. l., + 10 ° C m. a. t., 23. iv. 1952, leg. H. Remmert. — IRELAND, 5 adult males (as P. sagittalis, NMID), Lake Reenydonegan, 51.70 ° N 9.45 ° W, 3 m a. s. l., + 9 ° C m. a. t., 19. v. 1969, leg. J. Bracken. — MONTENEGRO, 1 adult male (as P. sagittalis, LUIZ), Karuc springs at Lake Skadar, 42.36 ° N 19.11 ° E, 8 m a. s. l., + 15 ° C m. a. t., 1. vi. 2018, leg. P. Gadawski [Barcode BIOUG 56531 - FO 4]; 1 adult male (as P. sagittalis, LUIZ), Vrijesko Vrelo, at spring, 42.48 ° N 19.15 ° E, 31 m a. s. l., + 14 ° C m. a. t., 30. v. 2018, leg. A. Zawal et al. [Barcode BIOUG 56221 - AO 8]; 1 adult male (as P. sagittalis, LUIZ), Malo Blato, 42.35 ° N 19.18 ° E, 6 m a. s. l., + 15 ° C m. a. t., 6. vi. 2018, leg. P. Gadawski [Barcode BIOUG 57312 - D 12]. — NETHERLANDS, 3 adult males (NHRS), Atlantic Sea, Katwijk aan Zee, 52.20 ° N 4.39 ° E, 0 m a. s. l., + 11 ° C m. a. t., 19. vi. 1993, leg. Y. Brodin. — SPAIN, 1 adult male (as P. sagittalis, UBCL), Galindo, Sestao, Atlantic coast, 43.30 ° N 2.99 ° W, 2 m a. s. l., + 15 m. a. t., vii. 2014, leg. R. Quintana. — SWEDEN, 2 adult males (NHRS), Baltic Sea, Hamnefjärden Bay, Simpevarp, 57.42 ° N 16.67 ° E, 0 m a. s. l., + 9 ° C m. a. t., 5. v. 1978, leg. Y. Brodin; 2 adult males (as P. sagittalis, NHRS), Baltic Sea, Furusundsfjärden islands, 59.70 ° N 18.90 ° E, 0 ‒ 1 m a. s. l., + 5 ° C m. a. t., 15. viii. 2008, leg. C. Essenberg [Barcode BSCHI 081 - 11 and BSCHI 082 - 11]; 1 adult male (as P. sagittalis, NHRS), Baltic Sea, Rödkobbehamnskär Island, 59.47 ° N 19.12 ° E, 1 m a. s. l., + 6 ° C m. a. t., 4. ix. 2010, leg. G. Hjertstrand [Barcode BSCHI 098 - 11]; 3 adult males (as P. sagittalis, NHRS), Baltic Sea, Södra Skräplen island, 59.14 ° N 18.75 ° E, 0 m a. s. l., + 6 ° C m. a. t., 28. vii. 2011, leg. G. Hjertstrand [Barcode BSCHI 363 - 17, BSCHI 364 - 17 and BSCHI 421 - 17]; 1 adult male (as P. sagittalis, NHRS), Baltic Sea coast, Pataholm, 56.92 ° N 16.43 ° E, 0 m a. s. l., + 6 ° C m. a. t., 2. vi. 2012, leg. Y. Brodin [Barcode BSCHI 683 - 17]; 1 adult male (as P. sagittalis, NHRS), Baltic Sea coast, Orrvarp, Gaviksfjärden Bay, 62.84 ° N 18.21 ° E, 0 m a. s. l., + 3 ° C m. a. t., 13. viii. 2013, leg. Y. Brodin [Barcode BSCHI 819 - 17]. Diagnostic characters. Figs. 39, 104 ‒ 106, key couplet 21. P. breviatus has a short gonostylus process with a GspR that overlaps that of seven other species of Procladius in Europe. Of these, P. appropinquatus, P. bellus, P. nudipennis and P. clavus are easily morphologically separated from P. breviatus as evident from the illustrations of genitalia and several characters in the key and the helpdesk. The GspR of P. breviatus to some degree overlaps that of P. saeticubitus (0.10 ‒ 0.16 versus 0.14 ‒ 0.20). P. breviatus can be distinguished from P. saeticubitus by the less slender gonostylus (GsmR 4.6 ‒ 5.7 versus 5.9 ‒ 6.9), wing vein Cu stem setae numbers (0 versus 5 ‒ 33) and the gonostylus process length compared to width (0.5 ‒ 0.8 versus 0.8 ‒ 1.1). The GspR of P. breviatus to some degree overlaps that of P. exilis (0.10 ‒ 0.16 versus 0.14 ‒ 0.19). P. breviatus can be distinguished from P. exilis by the less slender gonostylus (GsmR 4.6 ‒ 5.7 versus 5.9 ‒ 6.9), median anepisternum setae numbers (0 versus 10 ‒ 18) and antenna AR (1.7 ‒ 2.1 versus 2.6 ‒ 2.7). P. breviatus is sometimes very difficult to distinguish from P. choreus in cases where the GspR is overlapping (0.10 ‒ 0.16 versus 0.13 ‒ 0.20). When overlapping, the species can almost always be separated by a combination of characters related to size. P. breviatus is often bigger than P. choreus expressed as wing length wing length (2.3 ‒ 3.2 versus 1.8 ‒ 2.8), mid leg tibia length (1.05 ‒ 1.26 mm versus 0.72 ‒ 1.14) and body length (3.9 ‒ 5.0 mm versus 2.9 ‒ 4.4). In addition, P. breviatus has an on average broader gonostylus process relative to length compared with P. choreu s (0.5 ‒ 0.8 versus 0.7 ‒ 1.2). P. breviatus has almost only been found in brackish or saline water of the sea or water bodies within 30 km of the seacoast. P. choreus is also found in brackish water of the sea, but more often in inland waters far from the seacoast. P. breviatus Remmert, 1953, has for more than fifty years been regarded as a synonym of P. sagittalis Kieffer, 1909. The drawing of Kieffer’s P. sagittalis and other information in Remmert (1953) reveals that it is a synonym of P. culiciformis. The adult female of P. breviatus has been briefly described, the pupa in detail, whereas the larva is undescribed. Barcodes of adult males, adult females and larvae are available. Geographical distribution and ecology. In Europe, P. breviatus has been found from latitude 41 ° N in North Macedonia to 63 ° N in Finland. The southernmost findings of the species are from the Mediterranean coast of Morocco at 35 ° N. A great majority of the 42 sites with findings are in the sea near the coast, stagnant water bodies connected to the sea or inland water within 30 km from the sea. Confirmed findings are from altitude ‒ 1 to 40 m above sea level, which means that P. breviatus has the most limited altitude range of all European Procladius, except for P. clavus. Mean annual temperature of localities with findings of P. breviatus range from + 18 ° C in northern Morocco to + 3 ° C in Finland. Larvae of P. breviatus inhabit shallow parts of sea bays, estuaries or nearshore zones of the Atlantic Ocean, the Baltic Sea and the Mediterranean Sea. Near the sea, findings are from lakes, ponds, permanent or temporary pools, springs, man-made ditches and in a few cases slow-flowing sections in rivers. Most of these waters have a salinity ranging from slightly brackish conditions to somewhat higher than on average in the Mediterranean Sea (38 ‰) and the Atlantic Ocean (35 ‰). P. breviatus can be considered the best adapted to saline and marine conditions of all Procladius species in Europe. Most findings of P. breviatus are from the littoral zone with eutrophic to hypereutrophic conditions at water depth 0 ‒ 4 m, sometimes with very low oxygen concentration. No confirmed records from the profundal are known. Some findings are from mesotrophic conditions and a few from oligotrophic ones. Larvae of P. breviatus are usually found in waters with pH above 7, but there are a few findings from acid conditions including a very acid peat pool with pH 4.2. Larvae inhabit mud bottoms with dense to sparse vegetation of e. g. Phragmites, Juncus and Scirpus. It is not known what kind of food items the larvae consume. Adults have been caught from late April to mid-September. Countries or autonomous regions with records of P. breviatus in Europe are Albania, Denmark, England, Estonia, Finland, France, Germany, Ireland, Montenegro, Netherlands, Northern Ireland, North Macedonia, Norway, Poland, Portugal, Russia, Scotland, Spain, Sweden and Wales. The species is also known to occur in Morocco, Japan and Asian Russia.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF93091B3CE4FD319C25F8B1.taxon	materials_examined	Material examined (n = 47). BELGIUM, 1 adult male (as Procladius sp., RBNS), Mol, Postel, 51.29 ° N 5.19 ° E, 40 m a. s. l., + 10 ° C m. a. t., vii. 1925, leg. G. Severin. — CYPRUS, 2 adult males (as Procladius sp., LUIZ), Polis town, 35.03 ° N 32.42 ° E, 20 m a. s. l., + 19 ° C m. a. t., 23. vii. 2007, leg. J. Kazimierczak. — CZECHIA, 1 adult male (NHRS), Lake Blatec, Divcice, 49.11 ° N 14.31 ° E, 397 m a. s. l., + 7 ° C m. a. t., 19. vi. 1991, leg. Y. Brodin and M. Gransberg. — EGYPT, 2 adult males (as Procladius sp., CBGG), Alexandria, Smouha, Antoniades Gardens, Lake Farm, 31.20 ° N 29.95 ° E, ‒ 4 m a. s. l., + 21 ° C m. a. t., 22 ‒ 29. v. 2013, leg. O. El-Ansary [Barcode GMESA 103 - 14 and GMEEO 10040 - 21]. — ENGLAND, 1 adult male (as P. sagittalis), Lewes, 50.87 ° N 0.01 ° E, 2 m a. s. l., + 10 ° C m. a. t., 1948; 4 adult males (UUZM), South Chingford, Banbury Reservoir, 51.60 ° N 0.03 ° W, 10 m a. s. l., + 11 ° C m. a. t., 18. vii. 1982, leg. T. Samman; 2 adult males (UUZM), South Hanningfield, Hanningfield Reservoir, 51.65 ° N 0.51 ° E, 53 m a. s. l., + 11 ° C m. a. t., 23. vii. 1982, leg. T. Samman and Y. Brodin; 2 adult males (WIFE), London, Ashford Common, 51.42 ° N 0.44 ° W, 12 m a. s. l., + 11 ° C m. a. t, 3. ix. 1989, leg. R. S. Wotton. — FRANCE, 1 adult male (NHRS), La Guiel stream, 49 ° N 0 ° E, circa 1 m a. s. l., + 11 ° C m. a. t., 6. vi. 2007, leg. J. Moubayed-Breil [Barcoded]. — GERMANY, 1 adult male (as P. sagittalis, ZSMG), Berlin, pond at Tegel, 52.57 ° N 13.28 ° E, 38 m a. s. l., + 8 ° C m. a. t., 1932; 2 adult males (ZMLU), Lake Grosser Plönen See, Plön, 54.14 ° N 10.44 ° E, 18 m a. s. l., + 9 ° C m. a. t., 1947, leg. A. Thienemann. — IRAN, 1 adult male (as P. sagittalis, UKSI), Qeshlagh reservoir, River Qeshlagh, 35.43 ° N 47.00 ° E, 1 394 m a. s. l., + 12 ° C m. a. t., 16. vii. 2018, leg. H. Mohammadi and E. Ghaderi. — ITALY, 1 adult male (NHRS), Lake Lago di Landro, Carbonin Senluderb, 46.63 ° N 12.23 ° E, 1 403 m a. s. l., + 5 ° C m. a. t., 29. vi. 1991, leg. Y. Brodin; 1 adult male (MTSN), Lago di Garda, Riva del Garda, 45.87 ° N 10.83 ° E, 65 m a. s. l., + 13 ° C m. a. t., 29. ix. 2004, leg. L. Marziali. — JAPAN, 2 adult males (KUHY), Lake Ikuta, 35.3 ° N 139.3 ° E, 60 m a. s. l., + 16 ° C m. a. t., vii. 1992, leg. T. Kobayashi. — LEBANON, 3 adult males (NHRS), Anjar-Chamsine, affluent Ghozayel du Litani, cold springs, 33.74 ° N 35.96 ° E, 940 m a. s. l., + 14 ° C m. a. t., 15. iii. 1982, leg. J. Moubayed-Breil; 2 adult males (NHRS), River Damour, Damour, 33.70 ° N 35.46 ° E, 50 m a. s. l., + 18 ° C m. a. t., 17. vi ‒ 17. vii. 1979, leg. J. Moubayed-Breil; 2 adult males (NHRS), Jib-Jennine, stream, 33.54 ° N 35.69 ° E, 800 m a. s. l., + 15 ° C m. a. t., 10. v. 1982, leg. J. Moubayed-Breil; 2 adult males (NHRS), Nahr Beyroth Chyah Reservoir, 33.9 ° N 35.5 ° E, 39 m a. s. l., + 18 ° C m. a. t., 22. v. 1982, leg. J. Moubayed-Breil. — PORTUGAL, 2 adult males (NHRS), Albuferira, Santa Clara Reservoir, 37.52 ° N 8.44 ° W, 127 m a. s. l., + 16 ° C m. a. t., 5. v. 1996, leg. Y. Brodin and K. Murray-Brodin. — ROMANIA, 2 adult males (NHRS), Malaie, Lake Bradisar, 45.35 ° N 24.08 ° E, 450 m a. s. l., + 8 ° C m. a. t., 23. vi. 1991, leg. Y. Brodin and M. Gransberg. — RUSSIA, 2 adult males (as Procladius sp., TIEV), Kuybyshev Reservoir, 53.6 ° N 49.0 ° E, 47 m a. s. l., + 6 ° C m. a. t., vii. 1977; 1 adult male (SMUS), River Volga, Saratov, 51.3 ° N 45.9 ° E, 9 m a. s. l., + 8 ° C m. a. t., vi. 1995, leg. I. V. Sergeeva. — SPAIN, 1 adult male (as Procladius sp., DEBE), Cordoba, Retortillo Reservoir, 37.50 ° N 5.21 ° W, 189 m a. s. l., + 17 ° C m. a. t., 24. xi. 1974, leg. N. Prat; 1 adult male (as P. rivulorum, DEBE), Orense, Velle Reservoir, 42.22 ° N 7.09 ° W, 108 m a. s. l., + 14 ° C m. a. t., 12. v. 1974, leg. N. Prat. — SWEDEN, 1 adult male (NHRS), Baltic Sea, Hamnefjärden Bay, Simpevarp, 57.42 ° N 16.67 ° E, 0 m a. s. l., + 9 ° C m. a. t., 5. v. 1978, leg. Y. Brodin; 2 adult males (NHRS), Askö island, west of Askötorp, 58.81 ° N 17.67 ° E, 5 m a. s. l., + 6 ° C m. a. t., 24. vii ‒ 19. viii. 2011, leg. B. - E. Bengtsson [Barcode BSCHI 556 - 17]; 1 adult male (NHRS), Baltic Sea, Stora Karlsö island, Älmar, 57.29 ° N 17.97 ° E, 3 m a. s. l., + 7 ° C m. a. t., 13. viii. 2011, leg. M. Forshage [Barcode BSCHI 187 - 17]. — TUNISIA, 1 adult male (as Procladius sp., LHUC), Magsbaya, 37.09 ° N 9.26 ° E, 150 m a. s. l., + 18 ° C m. a. t., 2005, leg. S. Boulaaba. — UNITED STATES, 2 adult males (as P. freemani, USNM), Edgerton, Lake Koshkanong, 42.86 ° N 89.00 ° W, 312 m a. s. l., + 8 ° C m. a. t., 9. vii. 1947, leg. H. S. Dybas. Diagnostic characters. Figs. 2, 18, 41, 110 ‒ 112, key couplet 22. P. choreus has a short gonostylus process with a GspR that overlaps that of nine other species of Procladius in Europe. Of these, P. bellus, P. nudipennis, P. clavus and P. gemma are easily morphologically separated from P. choreus as evident from the illustrations of genitalia and several characters in the key and the helpdesk. P. choreus is sometimes very difficult to distinguish from P. breviatus in cases where the GspR is overlapping (0.13 ‒ 0.20 versus 0.10 ‒ 0.16). When overlapping, the species might be separated by a combination of characters related to size. P. choreus is often smaller than P. breviatus expressed as wing length (1.8 ‒ 2.8 versus 2.3 ‒ 3.2), mid leg tibia length (0.72 ‒ 1.14 mm versus 1.05 ‒ 1.26) and body length (2.9 ‒ 4.4 mm versus 3.9 ‒ 5.0 mm). In addition, P. choreus has an on average narrower gonostylus process relative to length compared with P. breviatus (0.7 ‒ 1.2 versus 0.5 ‒ 0.8). The GspR of P. choreus entirely overlaps that of P. exilis (0.13 ‒ 0.20 versus 0.14 ‒ 0.19). P. choreus can be distinguished from P. exilis by the less slender gonostylus (GsmR 4.9 ‒ 5.6 versus 5.9 ‒ 6.9), median anepisternum setae numbers (0 ‒ 2 versus 10 ‒ 18) and size expressed as wing length (1.8 ‒ 2.8 versus 3.1 ‒ 3.5). The GspR of P. choreus entirely overlaps that of P. saeticubitus (0.13 ‒ 0.20 versus 0.14 ‒ 0.20). P. choreus can be distinguished from P. saeticubitus by the less slender gonostylus (GsmR 4.9 ‒ 5.6 versus 5.9 ‒ 6.9), wing vein Cu stem setae number (0 versus 5 ‒ 33) and mostly also size e. g. mid leg tibia length (0.72 ‒ 1.14 mm versus 1.13.1 ‒ 1.43). The GspR of P. choreus partly overlaps that of P. islandicus (0.13 ‒ 0.20 versus 0.18 ‒ 0.24). If overlapping, the species can mostly be distinguished by a combination of other characters. P. choreus has shorter hairs on the front leg compared with P. islandicus (BR 1.5 ‒ 3 versus 3 ‒ 6.5), usually shorter wing length (1.8 ‒ 2.8 mm versus 2.6 ‒ 3.5) and usually lighter colour exemplified by the posterior colour of tergite II ‒ IV (whitish to light brown contrasting with the clearly darker anterior part versus contrasting light brown to dark brown as the rest of the tergites). P. choreus is sometimes difficult to distinguish from P. culiciformis in cases where the GspR overlaps (0.13 ‒ 0.20 versus 0.18 ‒ 0.25). When overlapping, the species can mostly be distinguished by a combination of other characters. P. choreus often has a slenderer gonostylus (GsmR 4.9 ‒ 5.6 versus 4.2 ‒ 5.2), and mostly smaller size as expressed by body length (2.9 ‒ 4.4 mm versus 4.4 ‒ 5.7 mm) and wing length (1.8 ‒ 2.8 mm versus 2.4 ‒ 3.5 mm). P. choreus is confused with P. culiciformis in several publications, although correctly identified in the key of Pinder (1978), Sergeeva (1995) and Langton & Pinder (2007). The adult female, pupal exuvia and larva have been described in several papers. Barcodes of adult males and adult females are available. Geographical distribution and ecology. P. choreus is among the most recorded and best studied species of Procladius in Europe and the world. Quality assured records are from about 149 sites. It has a wide geographical distribution with the southernmost findings in the Canary Islands (Spain), Saudi Arabia and India at latitude 27 ‒ 28 ° N. The southernmost findings in Europe are at 35 ‒ 36 ° N in Malta and Cyprus, and the northernmost in Finland at 60 ° N. The findings worldwide comprise about 57 countries or autonomous regions from the subtropical to the boreal zone, and even a semidesert area of southern Russia. Mean annual temperature ranges from + 25 to + 5 ° C. There is a distinct trend that specimens of P. choreus are gradually darker, larger and somewhat hairier with colder climate conditions. Records of the species are from at least 38 countries or autonomous regions in Europe, with a notable occurrence on the isolated Azore Islands at 28 ° W. Outside Europe, findings of P. choreus are reported from about 20 countries or autonomies stretching from the Canary Islands, Jordan, Iran, India to the most western findings in Japan. Most findings are from altitudes from below sea level at – 5 m to about 300 m above. In Mediterranean countries some records are from above 1 000 m above sea level with 1 400 m in Italy as the highest point. Almost 1 400 m above sea level is also reached in Iran. P. choreus is among the most common of all chironomid species in European lakes and lake-like reservoirs. Literature studies imply that the species can be expected from all European lakes with mesotrophic, eutrophic and hypereutrophic conditions. P. choreus is rather common in oligotrophic conditions but possibly never reported from ultraoligotrophic environments. P. choreus has frequently been reported to be the most abundant chironomid species or even of all macrobenthos in eutrophic to hypereutrophic lakes and reservoirs. In line with this, the larvae have been pointed out as very important food items for various fish such as carps (Cyprinidae) and salmonids (Salmonidae). Adult P. choreus have been reported to be important food items for birds and bats. Larvae of P. choreus are mainly littoral at water depths from 0 to 8 m in vegetation referred to as reed (Phragmites, Arundo, Glyceria and others) or water lilies (Nymphaeaceae), but also in zones with only submerged plants and macroalgae. The larvae are mostly on mud bottoms, but sometimes on sand bottoms. Larvae of P. choreus are less frequently found in the profundal zone with low oxygen conditions in eutrophic or hypereutrophic lakes and reservoirs where they seem to be outnumbered by P. ferrugineus. P. choreus might have the most extensive adaptability to different kinds or waters among the species of Procladiu s. Except for P. breviatus, it seems to be the best adapted Procladius to saline conditions, such as brackish coastal waters at the Black Sea and the Baltic Sea and estuaries of the Baltic Sea, the Atlantic Ocean or the Mediterranean Sea. Larvae of P. choreus inhabit lakes or ponds with much higher salt content than in the sea, such as in small coastal waters in Scotland, France and Tunisia. It has not been found in true marine conditions. Rivers and streams are frequent habitats of P. choreus larvae, but not the swift flowing upper reaches. Springs, ponds and puddles are also colonized by P. choreus larvae, even those which dry out completely during summer. One of these ponds is defined as thermal as it is fed by hot water from a natural spring. The larvae are well suited to withstand strong human impact such as severe eutrophication and heavy metal contamination. P. choreus are among the first chironomids to colonize new ponds and reservoirs only a few days after these have been constructed. Even completely human constructions in urban areas are colonized by P. choreus larvae, such as garden or park ponds in Austria, Germany and Spain, fountain ponds in Denmark and sand filter beds in constructed pools for water purification in England and Japan. Some of these ponds are only 1 m in diameter, less than 1 m deep and contain no water during winter months. Larvae of P. choreus are furthermore known to inhabit flooded rice fields in Italy, Hungary and Romania. The omnivorous diet of P. choreus includes detritus, algae and animals such as other chironomids, oligochaetes and crustaceans. Mainly animal food promotes faster growth and earlier emergence, while larvae feeding only on algae and detritus seem not to be able to emerge to adults. The species seems to be univoltine in northern Europe with adults flying from mid-April to mid-September. In southern Europe P. choreus fly all months and may develop three generations per year, or even four in southern Spain. Countries with records of P. choreus in Europe are Albania, Austria, Belgium, Bulgaria, Croatia, Czechia, Cyprus, Denmark, England, Estonia, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Latvia, Lithuania, Malta, Moldova, Montenegro, Netherlands, Northern Ireland, North Macedonia, Poland, Portugal (including the Azores), Romania, Russia, Scotland, Serbia, Slovakia, Spain, Sweden, Switzerland, Ukraine and Wales. Outside Europe P. choreus is recorded from Algeria, Azerbaijan, Canada, China, Egypt, India, Iran, Israel, Japan, Jordan, Lebanon, Mongolia, Morocco, North Korea, Russia (Asian part), Saudi Arabia, South Korea, Tunisia, Turkey and the United States. It might also be present in Bangladesh, Pakistan, Philippines and Thailand.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8F09043CE4FF709AF6FB8F.taxon	materials_examined	Material examined (n = 5). CANADA, 1 adult male (Holotype of P. clavus, CNCC), Spence Bay, Taloyoak, 69.5 ° N 93.3 ° W, 3 m a. s. l., ‒ 14 ° C m. a. t., 22. vii. 1951, leg. J. G. Chillcott; 1 adult male (Paratype of P. clavus, CNCC), Lake Ceillini, Taloyoak, 69.5 ° N 93.4 ° W, 20 m a. s. l., ‒ 14 ° C m. a. t., 19. vii. 1952, leg. J. G. Chillcott. — RUSSIA, 2 adult males (IANR), tundra ponds, east of Naryan-Mar, Yarey-shor, 67.62 ° N 53.70 ° E, 20 m a. s. l., ‒ 3 ° C m. a. t., 29. vii. 1990, leg. A. Rybakova; RUSSIA (Asia), 1 adult male (NHRS), Chaunskaya Gulf, Ajon Island, 69.55 ° N 169.18 ° E, 10 m a. s. l., ‒ 10 ° C m. a. t., 15 ‒ 18. vii. 2015, leg. P. Mortensen [Barcoded]. Diagnostic characters. Figs. 22, 53 ‒ 55, key couplet 5. P. clavus is easily distinguished from all other Procladius of Europe and North America. The semicircular projection posteriorly of tergite IX, the short, broad gonostylus and the long medioapodeme with a strongly curved inner section form a unique combination of characters. Description of P. clavus male genitalia and pupal exuvia in Egan & Langton (2018) might be another species considering the small size. The adult female has been described, but the larva is unknown. No barcoded material has been identified. Geographical distribution and ecology. Confirmed records of P. clavus are from four sites only, so knowledge of its ecology and geographical distribution is very limited. The sites are in the tundra at 68 ‒ 70 ° N in Russia and Canada with mean annual temperature between ‒ 3 to ‒ 14 ° C. This means that P. clavus is the only species of Procladius that has not been found at mean annual temperature of 0 or warmer, and only P. exilis and P. frigidus are known from colder conditions. The sites with P. clavus are in environments with small oligotrophic to possibly mesotrophic ponds at 3 ‒ 20 m above sea level within 60 km from the Arctic Ocean. Food requirements of the larvae are not known. Countries with records of P. clavus in Europe is only Russia. P. clavus is also present in Canada and the Asian part of Russia.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8F09063CE4FB479DB6F959.taxon	materials_examined	Material examined (n = 36). BELGIUM, 1 adult male (Syntype of P. bifasciatus, RBNS), Namur district, Falaën, 50.28 ° N 4.80 ° E, 204 m a. s. l., + 9 ° C m. a. t., 12. vi. 1921, leg. M. Goetghebuer; 1 adult male (Syntype of P. cinereus, RBNS), Postel, 51.29 ° N 5.19 ° E, 40 m a. s. l., + 10 ° C m. a. t., 11. vi. 1923, leg. G. Severin; 1 adult male, (as Procladius sp., RBNS), Lake Zillebekevijver, 50.84 ° N 2.92 ° E, 23 m a. s. l., + 10 ° C m. a. t., 12. vii. 1939, leg. M. Goetghebuer. — CANADA, 1 adult male (Syntype of P. abetus, CNCC), Alta, Wabamun, 53 ° N 114 ° W, 721 m a. s. l., + 2 ° C m. a. t., 6. vii. 1931, leg. F. H. Strickland; 1 adult male (as P. culiciformis, FWIM), Lake Winnipeg, Selkirk Island, 53.30 ° N 99.01 ° W, 217 m a. s. l., 0 ° C m. a. t., 30. vi. 1969, leg. P. S. S. Chang; 1 adult male (as P. paragretis, ZMBN), Lake Winnipeg, Matheson Island, 51.76 ° N 96.89 ° W, 216 m a. s. l., + 1 ° C m. a. t., 6. vii. 1931, leg. P. S. S. Chang. — CHINA, 1 adult male (as Procladius sp., NUCL), Yuyao, Siminghu Reservoir, 29.93 ° N 121.07 ° E, 27 m a. s. l., + 17 ° C m. a. t., 21. vii. 2011, leg. X. Lin [Barcode TANCH 097 - 16]; 1 adult male (as Procladius sp., NUCL), Yunhe, Xiaoshuncun, 28.20 ° N 119.64 ° E, 173 m a. s. l., + 16 ° C m. a. t., 29. vii. 2012, leg. W. Liu [Barcode TANCH 123 - 16]; 1 adult male (as Procladius sp., NUCL), Lishui, Yunhe, 28.12 ° N 119.57 ° E, 131 m a. s. l., + 17 ° C m. a. t., 29. vii. 2012, leg. X. Lin [Barcode TANCH 182 - 16]; 1 adult male (as Procladius sp., NUCL), Kaihua, Qianjiangyuan, 29.10 ° N 118.40 ° E, 120 m a. s. l., + 17 ° C m. a. t., 31. vii. 2012, leg. X. Lin [Barcode TANCH 131 - 16]; 1 adult male (as Procladius sp., NUCL), Huguan, Taihang Mountain, 35.93 ° N 113.61 ° E, 645 m a. s. l., + 12 ° C m. a. t., 11. x. 2014, leg. C. Song [Barcode TANCH 155 - 16]; 1 adult male (as Procladius sp., NUCL), Xiangxiang, Maohushuixiang, 27.75 ° N 112.52 ° E, 52 m a. s. l., + 16 ° C m. a. t., 6. v. 2015, leg. C. Song [Barcode TANCH 100 - 16]; 1 adult male (as Procladius sp., NUCL), Poyang, Niejia, 29.09 ° N 116.58 ° E, 27 m a. s. l., + 17 ° C m. a. t., 12. v. 2015, leg. C. Song [Barcode TANCH 026 - 16]. — CZECHIA, 4 adult males (NHRS), Cheb, Vodni Nadrz Skalka Reservoir, 50.08 ° N 12.32 ° E, 439 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin and M. Gransberg. — FINLAND, 2 adult males (as Procladius sp., HECH), Lake Vanajavesi, Aidassaari, 61.17 ° N 24.08 ° E, 79 m a. s. l., + 5 ° C m. a. t., 10 ‒ 12. vii. 1977, 17 ‒ 19. vii. 1977, leg. P. H. Kansanen; 3 adult males (as Procladius. sp., HECH), Lake Vanajavesi, K-niemi, 61.20 ° N 24.10 ° E, 79 m a. s. l., + 5 ° C m. a. t., 12 ‒ 14. viii. 1977, 18 ‒ 21. viii. 1977, leg. P. H. Kansanen; 1 adult male (as Procladius. sp., HECH), Lake Karvastenlahti, 61.16 ° N 24.03 ° E, 80 m a. s. l., + 5 ° C m. a. t., 25. v. 1978, leg. P. H. Kansanen; 1 adult male, Kiikala, Lake Tervakas, 60.49 ° N 23.70 ° E, 117 m a. s. l., + 5 ° C m. a. t., 9. v. 2023, leg. L. Paasivirta. — FRANCE, 1 adult male (as Procladius sp., NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 850 m a. s. l., + 7 ° C m. a. t., iv. 2019, leg. J. Moubayed-Breil. — POLAND, 3 adult males (LUIZ), Harcerska park, Spala, 51.54 ° N 20.14 ° E, 157 m a. s. l., + 9 ° C m. a. t., 5 ‒ 9. vi. 2007, leg. M. Płóciennik. — SCOTLAND; 1 adult male (as P. choreus, USNM), Arran Island, Lake Coire Fhionn Lochan, 55.66 ° N 5.34 ° W, 330 m a. s. l., + 7 ° C m. a. t., 29. v. 1919, leg. F. W. Edwards; 1 adult male (as Procladius sp., USNM) Beinn Heasgarnich, 56.5 ° N 4.6 ° W, 950 m a. s. l., + 5 ° C m. a. t., 11. vi. 1939, leg. F. W. Edwards; 1 adult male (BMNH), Lake Garten, 57.24 ° N 3.71 ° W, 238 m a. s. l, + 7 ° C m. a. t., 1947, leg. R. Coe. — SWEDEN, 1 adult male (Lectotype of Tanypus crassinervis, ZMLU), Lycksele, 64.6 ° N 18.7 ° E, 211 m a. s. l., + 1 ° C m. a. t., 2. viii. 1832, leg. J. W. Zetterstedt; 1 adult male (as Procladius sp., SLUU), Lake Mälaren, Västeråsfjärden, 59.55 ° N 16.55 ° E, 3 m a. s. l., + 6 ° C m. a. t., vii. 1971, leg. T. Wiederholm; 1 adult male (NHRS), Lake Färnebofjärden, Östa, 60.17 ° N 16.80 ° E, 60 m a. s. l., + 5 ° C m. a. t., 15. viii. 2007, leg. Y. Brodin; 2 adult males (NHRS), Valmbäcken rivulet and wetland, 60.30 ° N 16.84 ° E, 62 m a. s. l., + 5 ° C m. a. t., 7. ix. 2007, leg. A. Hagelin [1 Barcoded]. Diagnostic characters. Figs. 19, 31, 80 ‒ 82, key couplet 13. P. crassinervis has a long gonostylus process with a GspR that overlaps that of eight other species of Procladius in Europe. Of these, P. ferrugineus, P. fimbriatus, P. signatus and P. dentus are easily separated from P. crassinervis as evident from the illustrations of genitalia and several characters in the key and the helpdesk. Identification of P. crassinervis is sometimes difficult as the strongly upward oriented gonostylus process is frequently deformed and appears much shorter on slides than in reality, e. g. in macerated specimens or if slide preparation is not meticulously done. Moving the microscope ocular upwards and downwards to find out where the base and the top of the gonostylus process appear sharp can reveal the strongly upward direction of the gonostylus process. The GspR of P. crassinervis slighly overlaps that of P. frigidus (0.32 ‒ 0.39 versus 0.25 ‒ 0.33). P. crassinervis can easily be separated from P. frigidus by other characters, particularly the strongly upward oriented gonostylus process (30 – 60 ° versus 0 ‒ 20 °) but also the number of median anepisternum setae (0 versus 5 ‒ 26) and antenna AR (1.6 ‒ 2.1 versus 2.2 ‒ 2.8). P. crassinervis is found in temperate climate with mean annual temperature ranging from + 18 to + 1 C °, while P. frigidus is mainly found in the Artic with mean annual temperature below 0 C °. The GspR of P. crassinervis somewhat overlaps that of P. longistilus (0.32 ‒ 0.39 versus 0.25 ‒ 0.34). If overlapping, the species can usually be separated by the more strongly upward oriented gonostylus process of P. crassinervis (30 – 60 ° versus 0 ‒ 20 °), the stronger gonostylus process divergence (40 ‒ 75 ° versus 5 ‒ 30 °) and on average shorter body length (3.5 ‒ 5.1 mm versus 4.2 ‒ 5.7 mm). P. crassinervis is sometimes difficult to distinguish from P. floralis. The GspR of P. crassinervis considerably overlaps that of P. floralis (0.32 ‒ 0.39 versus 0.28 ‒ 0.37). The species can usually be separated by a combination of characters. P. crassinervis has a more strongly upward oriented gonostylus process (30 – 60 ° versus 10 ‒ 30 °), an on average broader gonostylus (GsmR 5.2 ‒ 5.9 versus 5.6 ‒ 6.8), a stronger gonostylus process divergence (40 ‒ 75 ° versus 30 ‒ 50 °) and a lower length ratio between the gonostylus and the gonocoxite (0.47 ‒ 0.55 versus 0.53 ‒ 0.57). The GspR of P. crassinervis substantially overlaps that of P. tatrensis (0.32 ‒ 0.39 versus 0.35 ‒ 0.43). P. crassinervis can be separated from P. tatrensis by several other characters, such as the number of median anepisternum setae (0 versus 7 ‒ 21), the more strongly upward oriented gonostylus process (30 – 60 ° versus 0 ‒ 20 °) and wing length (2.4 ‒ 3.3 mm versus 3.5 ‒ 4.5 mm). P. crassinervis has been identified under several other names, often because the gonostylus process has been deformed on slides. The name P. crassinervis has frequently been used for P. ferrugineus e. g. in the keys of adult males in Pinder (1978) and Langton & Pinder (2007). The female, pupal exuvia and larva have been briefly described. Barcodes of adult males and adult females are available. Geographical distribution and ecology. In Europe P. crassinervis is found from France and Austria at latitude 47 ° N to northern Sweden at latitude 65 ° N. In Asia the species known distribution is more southern ranging from 28 ° N in mid China to 45 ° N in southern Russia and northern Japan. In Canada records are from 53 ° N. The species has been found at altitudes ranging from 3 m to 950 m above sea level in Europe, with the highest point in Scotland. Records from northern Pakistan are at 1 370 m above sea level. Mean annual temperature at sites with records of P. crassinervis in Europe are from + 10 to + 1 ° C. In Asia and North America, the temperature range is from + 18 ° C in China to + 1 ° C in Canada. Larvae of P. crassinervis inhabit shallow water in temporary inundated swamps, small ponds and lakes. Some findings are from deeper zones of lakes, down to at least 14 m depth, and from slowly floating water including a forest ditch. Most sites with records of larvae are mesotrophic, eutrophic to hypereutrophic, while a few records are from oligotrophic conditions. Adults are found from early June to mid-October. Countries or autonomous regions with records of P. crassinervis in Europe are Austria, Belgium, Czechia, England, Finland, France, Germany, Lithuania, Russia, Scotland, Sweden and Switzerland. P. crassinervis is also present in Canada, China, Japan, Pakistan, Russia (Asia), South Korea and the United States.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8D09023CE4F90D9CFFF8E9.taxon	materials_examined	Material examined (n = 95). BELGIUM, 1 adult male (as P. choreus, RBNS), Gand, 51.05 ° N 3.72 ° E, 12 m a. s. l., + 10 ° C m. a. t., 19. iv. 1914, leg. M. Goetghebuer; 1 adult male (as Procladius sp., RBNS), Heusden, 51.03 ° N 3.80 ° E, 5 m a. s. l., + 11 ° C m. a. t., 23. v. 1943, leg. M. Goetghebuer. — CANADA, 1 adult male (as. P. freemani, CNCC), Val Marie, 49.24 ° N 107.73 ° W, 795 m a. s. l., + 4 m. a. t., 14. vi. 1956, leg. V. R. Vockeroth. — CZECHIA, 1 adult male (NHRS), Lake Rozmberk, Stara Hlina, 49.05 ° N 14.79 ° E, 423 m a. s. l., + 7 ° C m. a. t., 19. vi. 1991, leg. Y. Brodin; 3 adult males (NHRS), Cheb district, Vodni Nadrz Skalka Reservoir, 50.08 ° N 12.32 ° E, 439 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin and M. Gransberg; 1 adult male (NHRS), Cheb district, Lake Pomezsky rybnik, 49.72 ° N 16.29 ° E, 568 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin and M. Gransberg; 1 adult male (NHRS), Lake Blatec, Divcice, 49.11 ° N 14.31 E °, 397 m a. s. l., + 7 ° C m. a. t., 19. vi. 1991, leg. Y. Brodin and M. Gransberg; 1 adult male (NHRS), Lnare district, Vesky Ryb Reservoir, 49.44 ° N 13.82 ° E, 451 m a. s. l., + 7 ° C m. a. t., 18. vi. 1991, leg. Y. Brodin. — FAROE ISLANDS, 1 adult male (NHRS), Lake Mjauvotn, 62.12 ° N 7.00 ° W, 80 m a. s. l., + 5 ° C m. a. t., 27. vii. 1980, leg. G. Brodin-Lindsten; 4 adult males (NHRS), Lake Eithis, 62.17 ° N 7.06 ° W, 135 m a. s. l., + 5 ° C m. a. t., 15. viii. 2002, leg. Y. Brodin. — FINLAND, 2 adult males (as Procladius sp., MZHF), Lake Puruvesi, 61.73 ° N 29.35 ° E, 75 m a. s. l., + 3 ° C m. a. t., 16. vii. 1974, leg. B. Lindeberg; 2 adult males (as P. pectinatus, ZMUO), Hameenlinna, Lammin puhdistamo, 61.02 ° N 24.42 ° E, + 5 ° C m. a. t., 71 m a. s. l., 9. v. 2015, leg. L. Paasivirta [Barcode LEFIJ 3930 - 16 and LEFIJ 3931 - 16]; 1 adult male (as Procladius sp., ZMUO), Baltic Sea, Kolpanlahti Bay 61.61 ° N 21.63 ° E, 0 m a. s. l., + 5 ° C m. a. t., 4. vii. 2015, leg. L. Paasivirta [Barcode LEFIJ 3596 - 16]; 2 adult males (as P. cf. vesus), Malax, Rönnskären Island, 63.06 ° N 20.83 ° E, + 4 ° C m. a. t., 5 m a. s. l., 30. vii ‒ 5. viii. 2018, leg. L. Paasivirta. — FRANCE, 1 adult male (as Procladius sp., LHST), Lake d’Aumar, 42.84 ° N 00.15 ° E, 2 191 m a. s. l., + 2 ° C m. a. t., 24. viii. 1964, leg. H. Laville; 2 adult males (as Procladius sp., LHST), Lake Mare Lac Long, 42.82 ° N 0.12 ° E, 2 094 m a. s. l., + 3 ° C m. a. t., 25. vii. 1965, leg. H. Laville; 1 adult male (as Procladius sp. 1, LHST), Lake Gourg Nére Inferior, 42.87 ° N 0.18 ° E, 2 201 m a. s. l., + 2 ° C m. a. t., 26. vii. 1966, leg. H. Laville; 1 adult male (as Procladius sp., LHST), Lake Vert, 42.86 ° N 0.19 ° E, 2 114 m a. s. l., + 3 ° C m. a. t., 27. vii. 1966, leg. H. Laville; 1 adult male (as Procladius sp. 1, LHST), Lake Gourg Nére Moyen, 42.87 ° N 0.19 ° E, 2 211 m a. s. l., + 2 ° C m. a. t., 9. viii. 1966, leg. H. Laville; 1 adult male (as P. cf. scapularis, LHST), Lake Port-Bielh, 42.87 ° N 0.19 ° E, 2 285 m a. s. l., + 2 ° C m. a. t., 13. viii. 1968, leg. H. Laville; 1 adult male (as Procladius sp., LHST), River Lot, Golinhac, 44.61 ° N 2.60 ° E, 253 m a. s. l., + 12 ° C m. a. t., 22. vii. 1977, leg. H. Laville; 3 adult males (NHRS), Corsica, Lake Melu, 42.21 ° N 9.02 ° E, 1 708 m a. s. l., + 6 ° C m. a. t., 25. viii. 2015, leg. J. Moubayed-Breil; 3 adult males (NHRS), Corsica, Lake Calacuccia, 42.32 ° N 9.01 ° E, 774 m a. s. l., + 10 ° C m. a. t., 25. viii. 2015, leg. J. Moubayed-Breil; 1 adult male (as Procladius sp., NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 849 m a. s. l., + 7 ° C m. a. t., iv. 2019, leg. B. Tissot [Barcoded]. — GERMANY, 1 adult male (as Procladius sp., MNHB), Berlin, Pichelberg, 52.51 ° N 13.22 ° E, 33 m a. s. l., + 8 ° C m. a. t., viii. 1907, leg. B. Lichtwardt; 1 adult male (ZSMG), Münster, Dortmund-Ems canal, 51.95 ° N 7.66 ° E, 54 m a. s. l., + 10 ° C m. a. t., 1956; 1 adult male (as Procladius sp., ZFMK), Winningen, in forest, 50.31 ° N 7.51 ° E, 66 m a. s. l., + 10 ° C m. a. t., 4. x. 2012, leg. B. Rulik [Barcode GBMWN 781 - 15]; 1 adult male (as P. choreus, ZFMK), River Elz, Emmendingen, 48.11 ° N 7.85 ° E, 213 m a. s. l., + 10 ° C m. a. t., 10. ix. 2018 [Barcode GBOL- 2625211]. — HUNGARY, 4 adult males (NHRS), Derzsitavak Reservoir, 47.61 ° N 21.01 ° E, 86 m a. s. l., + 12 ° C m. a. t., 21. vi. 1991, leg. Y. Brodin; 1 adult male (NHRS), Hortobagy Reservoir III, Hortobagyi-halasto, 47.65 ° N 21.09 ° E, 84 m a. s. l., + 12 ° C m. a. t., 21. vi. 1991, leg. Y. Brodin. — ICELAND, 1 adult male, (as Procladius sp., RBNS), Myvatn, 65.62 ° N 16.94 ° W, 276 m a. s. l., + 3 ° C m. a. t., 20. viii. 1929; 2 adult males (NHRS), Lake Masvatn, 65.62 ° N 17.24 ° W, 266 m a. s. l., + 3 ° C m. a. t., 12. vii. 1978, leg. Y. Brodin; 2 adult males (NHRS), Lake Laugarvatn, 64.21 ° N 20.73 ° W, 76 m a. s. l., + 8 ° C m. a. t., 12. viii. 2000, leg. Y. Brodin. — IRELAND, 1 adult male (as Procladius sp., UCDZ), Lake Gougane Barra, 51.84 ° N 9.32 ° W, 165 m a. s. l., + 8 ° C m. a. t., 26. vii. 1966, leg. D. A. Murray. — LEBANON, 2 adult males (as P. cf. choreus, NHRS), Jib-Jennine stream, 33.54 ° N 35.69 ° E, 800 m a. s. l., + 15 ° C m. a. t., 10. v. 1982, leg. J. Moubayed-Breil. — POLAND, 1 adult male (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. J. Hellberg and Y. Brodin; 2 adult males (NHRS), west of Rabka, Lake Lebsko, 54.75 ° N 17.44 ° E and 54.76 ° N 17.47 ° E, ‒ 1 m a. s. l., + 9 ° C m. a. t., 8. viii. 2012, leg. Y. Brodin and L. Brodin [Barcode BSCHI 696 - 17 and BSCHI 597 - 17]. — ROMANIA, 1 adult male (NMNB), Zigoneni Reservoir, 45.09 ° N 24.66 ° E, 379 m a. s. l., + 9 ° C m. a. t., vii. 1974, leg. P. Albu; 1 adult male (NMNB), Lake Vidraru, 45.40 ° N 24.62 ° E, 803 m a. s. l., + 7 ° C m. a. t., vii. 1974, leg. P. Albu. — RUSSIA, 4 adult males (as Procladius sp.), Tsimlyansk Reservoir, 47.73 ° N 42.26 ° E, 31 m a. s. l., + 9 ° C m. a. t., vii. 1979, leg. M. P. Miroshnichenko; 3 adult males (IANR), Lake Kolodenskoe, 59.01 ° N 37.06 ° E, 113 m a. s. l., + 4 ° C m. a. t., 27. v. 1989, leg. A. Rybakova; 1 adult male (TUSF), Lake Ladoga, Sortavala, 61.74 ° N 30.77 ° E, 4 m a. s. l., + 5 ° C m. a. t., 4. viii. 1991, leg. G. Söderman. — SLOVAKIA, 1 adult male (as Procladius sp., ZSMG) Sládkovičovo kons., Lake Vincov les, 48.2 ° N 17.7 ° E, 120 m a. s. l., + 11 ° C m. a. t., 20. ix. 1962, leg. Lastovka; 2 adult males (NHRS), Lake Liptovska Mara, Liptovsky Mikulas, 49.10 ° N 19.57 ° E, 555 m a. s. l., + 6 ° C m. a. t., 19. vi. 1991, leg. Y. Brodin. — SPAIN, 1 adult male (as P. rivulorum, DEBE), El Vellon Reservoir, 40.46 ° N 3.37 ° W, 830 m a. s. l., + 13 ° C m. a. t., 11. xi. 1974, leg. N. Prat; 1 adult male (as Procladius sp., DEBE), Loriguilla Reservoir, 39.40 ° N 0.53 ° W, 322 m a. s. l., + 15 ° C m. a. t., 6. xii. 1974, leg. N. Prat; 1 adult male (as Procladius sp., DEBE), Guadalajara, Buendia Reservoir, 40.24 ° N 2.46 ° E, 714 m a. s. l., + 14 ° C m. a. t., 9. xi. 1974, leg. N. Prat. — SWEDEN, 1 adult male (as Procladius sp., NHRS), Lake Vitalampi, 59.55 ° N 15.13 ° E, 322 m a. s. l., + 5 ° C m. a. t., 12. vii. 1974, leg. P. Mossberg; 4 adult males (NHRS), Lake Vänern, Mariestadsfjärden Bay, Mariestad, 58.70 ° N 13.77 ° E, 44 m a. s. l., + 6 ° C m. a. t., 7 ‒ 9. v. 1987, leg. Y. Brodin and K. Nielsen; 1 adult male (NHRS), Baltic Sea, Furusundsfjärden islands, 59.70 ° N 18.90 ° E, 1 m a. s. l., + 5 ° C m. a. t., 15. viii. 2008, leg. C. Essenberg [Barcode BSCHI 085 - 11]; 1 adult male (NHRS), Baltic Sea coast, south of Obbola, Fläsebadet, 63.66 ° N 20.29 ° E, 0 m a. s. l., + 3 ° C m. a. t., 14. vii. 2011, leg. N. Ericson [Barcode BSCHI 287 - 17]; 1 adult male (NHRS), Baltic Sea coast, Sillviken Bay, Sillviken, 63.76 ° N 20.45 ° E, 0 m a. s. l., + 3 ° C m. a. t., 27. vii. 2011, leg. N. Ericson [Barcode BSCHI 360 - 17]; 1 adult male (NHRS), Baltic Sea, Ratakär Island, 63.99 ° N 20.89 ° E, 1 m a. s. l., + 3 ° C m. a. t., 3. viii. 2011, leg. N. Ericson [Barcode BSCHI 433 - 17]; 1 adult male (NHRS), Baltic Sea coast, Pataholm, 56.92 ° N 16.43 ° E, 1 m a. s. l., + 6 ° C m. a. t., 2. vi. 2012, leg. Y. Brodin [Barcode BSCHI 684 - 17]; 1 adult male (NHRS), Baltic Sea coast, Hamnefjärden Bay, Simpevarp, 57.42 ° N 16.67 ° E, 0 m a. s. l., + 9 ° C m. a. t., 2. vi. 2012, leg. Y. Brodin [Barcode BSCHI 638 - 17]; 1 adult male (NHRS), Baltic Sea coast, Vallvik beach, 61.18 ° N 17.19 ° E, 0 m a. s. l., + 5 ° C m. a. t., 25. vii. 2012, leg. Y. Brodin and G. Lindberg [Barcode BSCHI 611 - 17]; 1 adult male (NHRS), Baltic Sea coast, Sillvikskatan, 63.56 ° N 19.83 ° E, 0 m a. s. l., + 3 ° C m. a. t., 13. viii. 2013, leg. Y. Brodin [Barcode BSCHI 825 - 17]; 1 adult male (NHRS), Eriksberg nature reserve, Bruket pond, 56.16 ° N 15.00 ° E, 5 m a. s. l., + 7 ° C m. a. t., 4. vi. 2015, leg. J. Wolgast [Barcoded]. — SWITZERLAND, 4 adult males (as P. sagittalis, MCSN), Lake Geneva, 46.2 ° N 6.1 ° E, 372 m a. s. l., + 10 ° C m. a. t., 20 ‒ 24. vi. 1994, 20 ‒ 24. v. 1995, 23 ‒ 28. vi. 1995, leg. B. Lods-Crozet; 1 adult male (as P. cf. sagittalis, MCSN), small stream, Le Bainoz, Montet, 46.82 ° N 6.87 ° E, 505 m a. s. l., + 9 ° C m. a. t., 28. vi. 2006, leg. P. Stucki; 1 adult male (as P. choreus, MCSN), small stream, Moulin Veigy, Srce, 46.14 ° N 6.03 ° E, 410 m a. s. l., + 10 ° C m. a. t., 22. vi. 2006, leg. P. Stucki. — UNITED STATES, 1 adult male (as P. freemani, USNM), Dayton, Lake Flathead, 47.86 ° N 114.25 ° W, 882 m a. s. l., + 5 m. a. t., 13. vii. 1935, leg. Melander; 1 adult male (Holotype of P. freemani, USNM), San Bruno, 37.61 ° N 112.42 ° W, 8 m a. s. l., + 12 m. a. t., 23. viii. 1957, leg. R. P. Maynard. Diagnostic characters. Figs. 1, 3, 13, 40, 107 ‒ 109, key couplet 22. P. culiciformis has a medium long gonostylus process with a GspR that overlaps that of eleven other species of Procladius in Europe. Of these, P. exilis, P, saeticubitus, P. tenebricosus, P. lugubris and P. longistilus are distinctly separated from P. culiciformis by a slenderer gonostylus, reflected by the GsmR, and one or more other morphological characters in the key and the helpdesk. Also P. nudipennis and P. gemma are easily distinguished from P. culiciformis by several characters, although GspR overlaps. P. culiciformis is sometimes difficult to distinguish from P. choreus in cases where the GspR overlaps (0.18 ‒ 0.25 versus 0.13 ‒ 0.20). When overlapping, the species might be separated by the mostly thicker gonostyus of P. culiciformis (GsmR 4.2 ‒ 5.2 versus 4.9 ‒ 5.6), and mostly bigger size as expressed by body length (4.4 ‒ 5.7 mm versus 2.9 ‒ 4.4 mm) and wing length (2.4 ‒ 3.5 mm versus 1.8 ‒ 2.8 mm). The GspR of P. culiciformis entirely overlaps that of P. islandicus (0.18 ‒ 0.25 versus 0.18 ‒ 0.24). The species can be separated by the thicker gonostyus of P. culiciformis (GsmR 4.2 ‒ 5.2 versus 5.3 ‒ 6.4), the on average shorter setae of the front leg (BR 2 ‒ 4.5 versus 3 ‒ 6.5) and the on average lighter mid-section of the front leg tibia (whitish to light brown versus light brown to dark brown). The GspR of P. culiciformis to some degree overlaps that of P. pruinosus (0.18 ‒ 0.25 versus 0.23 ‒ 0.30). When overlapping, the species can usually be separated by the thicker gonostylus of P. culiciformis (GsmR 4.2 ‒ 5.2 versus 5.2 ‒ 6.3) and an on average lighter mid-section of the front leg tibia (whitish to light brown versus light brown to dark brown). The GspR of P. culiciformis slightly overlaps that of P. frigidus (0.18 ‒ 0.25 versus 0.25 ‒ 0.33). P. culiciformis can be distinguished from P. frigidus by the number of median anepisternum setae (0 versus 5 ‒ 26), the length of palpomere five divided with its width (9.3 ‒ 12.2 versus 8.0 ‒ 9.3) and an on average thicker gonostyus (GsmR 4.2 ‒ 5.2 versus 4.9 ‒ 5.8). The status of P. culiciformis has been unclear ever since Linneus’ description in 1767. It has frequently been confused with P. choreus in Europe and North America. The holotype of P. freemani from the United States is in fact P. culiciformis, but at least some of the paratypes might be P. choreus. The adult female, pupal exuvia and larva of P. culiciformis have been described. Barcodes of adult males, adult females, pupae and larvae are available. Geographical distribution and ecology. The circumpolar P. culiciformis has a more northern range than its possibly closest relative and morphologically similar P. choreus. In Europe P. culiciformis has been recorded from latitude 37 ° N in southern Spain to 66 ° N on Iceland and 68 ° N in northern Norway. It has been reported from 32 countries or autonomous regions in Europe. The most western outposts with findings in Europe are Portugal in the south and Iceland in the north. The southernmost records outside Europe are from 34 ° N in Lebanon and 35 ° N in Japan, and the northernmost from 62 ° N in northern Canada. Sites with P. culiciformis have a range from + 18 to ‒ 5 ° C mean annual temperature, which is on average colder than that of P. choreus but about the same as that of the also morphologically similar P. pruinosus. It has a higher altitude record than that of the other two, reaching 2 210 m in the Pyrenees of southern France. Males of P. culiciformis, as the genus Procladius in general, become smaller, lighter and less hairy with increasing mean annual temperature. P. culiciformis can be considered to be one of the most common species of Chironomidae in lakes and reservoirs in Europe, but for those with ultraoligotrophic conditions where no findings of the species are known. It is common in brackish water along the coast of the Baltic Sea and the Atlantic Ocean. Quality assured findings of P. culiciformis are from 129 sites, mainly lakes and lake-like reservoirs. Larvae of the species have been found in vegetation near the shore and down to 20 m depth, and in the profundal from 2 m down to a depth record of 206 m in a clear lake in Switzerland. Larvae of P. culiciformis can be common in slow-running streams and rivers, ponds and oligohaline to mesohaline water as in estuaries along the Baltic Sea, the Atlantic coast and the Black Sea. The larvae have sometimes been recorded in huge numbers from limnocrene springs. Man-made canals and ditches are other habitats inhabited by the larvae. P. culiciformis, possibly together with P. pruinosus, seems to be the Procladius species which is best adapted to water with high humus content. Larvae inhabit polyhumic lakes, tarns and pools of bog systems where water transparency might be less than 0.2 m. Conditions in these habitats are often acidic with pH sometimes as low as around 4. In line with this, P. culiciformi s holds the acidity record for a Procladius species with pH 3.3 in a ditch close to the Baltic coast of northern Sweden. P. culiciformis larvae are found in a wide range of trophic conditions, being common in oligotrophic to eutrophic conditions but less common in hypereutrophic conditions. Substantial decrease in numbers of P. culiciformis larvae have been noted for the profundal of reservoirs in Russia when the water changed from eutrophic to hypereutrophic accompanied by strongly deteriorated oxygen conditions. P. culiciformis females produce jelly covered balls with several hundred eggs which are laid in shallow water. Larvae of P. culiciformis are known to be omnivorous feeding on algae such as diatoms, detritus and animals smaller than themselves such as crustaceans, oligochaete worms and insects including other chironomids. In the laboratory, larvae of P. culiciformis have been observed to attack much bigger larvae of the chironomid genus Chironomus and suck out their bodies. Larvae which feed mainly on animals are reported to develop more rapidly and become bigger as adults. Larvae of P. culiciformis can be important food items for common fish species such as rainbow trout (Oncorhynchys mykiss) and three-spined stickleback (Gasterosteus aculeatus). Adults can be food of songbirds such as the Eurasian reed warbler (Acrocephalus scirpaceus). Adults of P. culiciformis are mostly found from mid-April to October, however until mid-December in southern Spain with up to four generations per year. A special case is emergence of adults in late February from a bay of the Baltic Sea in southern Sweden with winter water temperature + 5 to + 10 ° C higher than the surroundings because of cooling water discharges from a nuclear power plant. Sex ratios in P. culiciformis are sometimes very unbalanced. A report from Japan noted that males made up less than 1 % of about 16 700 collected specimens. Countries or autonomous regions with records of P. culiciformis in Europe are Albania, Austria, Belarus, Belgium, Bulgaria, Croatia, Czechia, England, Estonia, Faroe Islands, Finland, France, Germany, Greece, Hungary, Iceland, Ireland, Italy, Latvia, Lithuania, Montenegro, Netherlands, Norway, Poland, Portugal, Romania, Russia, Scotland, Slovakia, Spain, Sweden and Switzerland. P. culiciformis has also been recorded from Canada, China, Japan, Lebanon, Mongolia, Russia (Asia), the United States and Uzbekistan, possibly also India, Morocco and Tunisia.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8809033CE4FF709BB4F82B.taxon	description	? Procladius sp. — Langton et al. (2013), Norway, pupa, key, illustration.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8809033CE4FF709BB4F82B.taxon	materials_examined	Material examined (n = 25). CANADA, 1 adult male (Paratype of P. dentus, CNCC), Baffin Island, Camp Kungovik, 65 ° N 35 ° W, 0 m a. s. l., ‒ 10 ° C m. a. t., 16. vii. 1929, leg. J. D. Soper; 1 adult male (ISUA), Great Caribou Island, 52.27 ° N 55.59 ° W, 5 m a. s. l., + 4 ° C m. a. t., 8. vii. 1943, leg. J. M. Harrison; 1 adult male (Paratype of P. dentus, CNCC), Great Deer, 52.58 ° N 107.05 ° W, 529 m a. s. l., + 2 ° C m. a. t., 20 ‒ 21. v. 1949, leg. V. R. Vockeroth; 1 adult male (Paratype of P. dentus, CNCC), River Great Whale, 54.75 ° N 75.10 ° W, 200 m a. s. l., ‒ 1 ° C m. a. t., 30. vi. 1949, leg. V. R. Vockeroth; 1 adult male (Holotype of P. dentus, CNCC), Manitoba, Fort Churchill, 58.77 ° N 94.22 ° W, 0 m a. s. l., ‒ 6 ° C m. a. t., 12. vii. 1952, leg. J. G. Chillcott; 1 adult male (Paratype of P. dentus, CNCC), Manitoba, Fort Churchill, 58.77 ° N 94.22 ° W, 0 m a. s. l., ‒ 6 ° C m. a. t., 12. vii. 1952, leg. J. G. Chillcott. — FINLAND, 4 adult males, Baltic Sea, Storskär Island, Lake Käringsund, 63.43 ° N 21.06 ° E, 0 m a. s. l., + 4 ° C m. a. t., 9. vi ‒ 23. vii. 2009, leg. L. Paasivirta; 1 adult male (ZMUO), Pelkosenniemi, Kaetkaeaapa, 67.09 ° N 27.81 ° E, 179 m a. s. l., 0 ° C m. a. t., 15. vi. 2015, leg. L. Paasivirta [Barcode CHIFI 114 - 16]; 4 adult males, Baltic Sea, Storskär Island, Lake Käringsund, 63.43 ° N 21.07 ° E, 1 m a. s. l., + 4 ° C m. a. t., 30. v ‒ 13. vi. 2017, leg. L. Paasivirta; 4 adult males, Suvakoski, small lake, 67.31 ° N 28.16 ° E, 175 m a. s. l., + 1 ° C m. a. t., 9. vi ‒ 23. vii. 2009, leg. L. Paasivirta; 1 adult male, Tulppio, Ainijärvi, 67.76 ° N 29.45 ° E, 269 m a. s. l., 0 ° C m. a. t., 10 ‒ 14. vi. 2018, leg. J. Salmela. — NORWAY, 1 adult male (NTNU), Stabbursdalen, Rohkosjavri, 70.17 ° N 24.90 ° E, 12 m a. s. l., 0 ° C m. a. t., 15. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 499 - 11]; 1 adult male (NTNU), Sør-Varanger, Venstre Jakobselv, rockpools, 69.79 ° N 30.80 ° E, 7 m a. s. l., 0 ° C m. a. t., 21. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 445 - 11]. — SWEDEN, 2 adult males (NHRS), Torne lappmark, 68 ° N 19 ° E, 650 m a. s. l., ‒ 1 ° C m. a. t., 1. vii. 2016, leg. H. Vårdal [1 Barcoded]. — United States, 1 adult male (Paratype of P. dentus, CNCC), Alaska, Nome, 64.61 ° N 165.42 ° W, 10 m a. s. l., ‒ 4 ° C m. a. t., 3. vi. 1951, leg. D. P. Williams. Diagnostic characters. Figs. 14, 23, 56 ‒ 58, key couplet 6. The form of the medioapodeme and the distinct dorsal ridge on the gonocoxite readily separates P. dentus from other species of Procladius including its presumably closest relative P. signatus. The antennae AR-value of P. dentus is higher (AR 2.6 ‒ 3.0) than that of P. signatus (AR 1.9 ‒ 2.4). Adult males of P. dentus seem not to have been confused with other species. The adult female and larva have not been described, but the pupal exuvia has. Barcodes of adult males, adult females, pupae and larvae are available. Geographical distribution and ecology. P. dentus has only been found in six of the most northern countries of the world. Findings in Europe are from 63 to 70 ° N and outside Europe from 52 to 82 ° N in Canada and Alaska in the United States. Mean annual temperature at the twenty-two sites with quality assured findings ranges from + 3 ° C in mid Finland to ‒ 14 ° C in northernmost Canada. Several sites are at 0 ‒ 10 m above sea level close to the Arctic Ocean or the Baltic Sea where the larvae live in brackish water conditions in rock pools or shallow lakes influenced by brackish sea water. The highest altitude recorded for P. dentus is in Canada at 1 130 m above sea level. High tolerance towards, or possibly even preference for saline conditions can be illustrated by the findings of P. dentus larvae in a lake in Canada with salinity higher than that of the Atlantic Ocean and the Arctic Ocean. Larvae of P. dentus have been reported to inhabit shallow water of 0 ‒ 2 m depth in lakes, pools, mires and swamps and in one case a tarn without fish. The habitats can be classified as oligotrophic, mesotrophic or eutrophic. No findings are known from profundal water. A few findings are from slowly running water. Adults are known to fly from the middle of May to the middle of August. Countries with records of P. dentus in Europe are Finland, Norway, Russia and Sweden. It has also been recorded from Canada and the United States (Alaska).	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF87090F3CE4FABE9DF6F921.taxon	description	Trichotanypus parvulus Kieffer, 1918 — Kieffer (1918 a), Lithuania, adult male, adult female, description, illustration.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF87090F3CE4FABE9DF6F921.taxon	description	Trichotanypus parvulus Kieffer, 1918 — Goetghebuer (1927), France, adult male, adult female, key, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF87090F3CE4FABE9DF6F921.taxon	description	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.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF87090F3CE4FABE9DF6F921.taxon	description	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.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF87090F3CE4FABE9DF6F921.taxon	description	? Trichotanypus distinguendus Kieffer, 1915 — Goetghebuer (1927), France, adult male, adult female, key, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF87090F3CE4FABE9DF6F921.taxon	materials_examined	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; 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. — 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; 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. — 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. — ENGLAND, 2 adult males (as P. crassinervis, BMNH), Hawkshead, Three Dubs Tarn, 54.37 N 2.96 W, 218 m a. s. l., + 7 ° C m. a. t., 7 ‒ 15. v. 1947, leg. T. T. Macan. — 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. — 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. — 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. — 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; 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; 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 GBMTM 2155 - 15]; 1 adult male (ZFMK), east of Karlsruhe, Wilhelma, 48.98 ° N 8.43 ° E, 121 m a. s. l., + 10 m. a. t., 8. x. 2014 [Barcode GBMTB 294 - 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. — 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. — 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. — 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. — 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 [1 Barcode BSCHI 697 - 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; 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; 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. — 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; 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. — 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; 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 [Barcode BSCHI 083 - 11 and BSCHI 084 - 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 [Barcode BSCHI 563 - 17, BSCHI 570 - 17 and BSCHI 570 - 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 [Barcode BSCHI 222 - 17 and BSCHI 223 - 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 [Barcode BSCHI 256 - 17]; 1 adult male (NHRS), Baltic Sea coast, Östra Sikvik, 60.67 ° N 17.29 E °, 0 m a. s. l., + 5 ° C m. a. t., 15. vii. 2011, leg. Y. Brodin [Barcode BSCHI 368 - 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 [Barcode BSCHI 520 - 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 [BSCHI 629 - 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 [Barcode BSCHI 584 - 17 and BSCHI 585 - 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. Diagnostic characters. Figs. 30, 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.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8409083CE4F8D59FCFFBBD.taxon	materials_examined	Material examined (n = 8). FINLAND, 1 adult male (Holotype of P. fimbriatus, ZSMG), Lake Saanajärvi, 69.05 ° N 20.88 ° E, 679 m a. s. l., ‒ 4 ° C m. a. t., viii. 1956, leg. W. Wülker; 1 adult male (Paratype of P. fimbriatus, ZSMG), Lake Saanajärvi, 69.05 ° N 20.88 ° E, 679 m a. s. l., ‒ 4 ° C m. a. t., viii. 1956, leg. W. Wülker; 1 adult male (MZHF), Lake Ala-Kilpisjärvi, 68.95 ° N 20.92 ° E, 473 m a. s. l., ‒ 2 ° C m. a. t., 25. viii. 1983, leg. J. Tuiskunen; 2 adult males, Lake Kilpsiärvi, Saana, 69.02 ° N 20.84 ° E, 491 m a. s. l., ‒ 2 ° C m. a. t., 16. vii ‒ 24. viii. 2007. — SWEDEN, 1 adult male (ZSMG), Abisko district, lake at stream draining, 68.3 ° N 18.8 ° E, 410 m a. s. l., ‒ 1 ° C m. a. t., 26. vi. 1958, leg. D. R. Oliver; 2 adult males (NHRS), Lake Sitasjaure, 67.87 ° N 17.61 ° E, 613 m a. s. l., ‒ 3 ° C m. a. t., 7. viii. 1992, leg. Y. Brodin. Diagnostic characters. Figs. 20, 47 ‒ 49, key couplet 2. The female-like male antenna with its low antennal ratio (AR) and reduced plume separates P. fimbriatus from all other known species of Procladius. The species can readily be separated also from other species of Procladius in Europe and North America by several additional characters although the gonostylus process GspR overlaps that of six other species of Procladius present in Europe. Absence of any signs of parasitism, e. g. by nematodes, in the body of all eight studied specimens suggest that P. fimbriatus is a valid species and not an intersex which also may have female-like antenna but normal male genitalia (Wülker 1959; Aagaard 1974). Wülker, who first described P. fimbriatus, was a specialist in parsitism in Chironomidae. The adult female and pupal exuvia have been described. The larva is not known. No barcoded specimens are known. Geographical distribution and ecology. All findings of P. fimbriatus are from northern regions of Norway, Sweden and Finland, at 68 ‒ 70 ° N and altitude 150 ‒ 680 m above sea level. All six sites with findings of adults are above the tree limit in the taiga with a subarctic climate of + 1 to ‒ 4 ° C mean annual temperature and ice cover on lakes for 7 ‒ 10 months annually. Pupal exuviae and adults of P. fimbriatus have been found in or close to ultraoligotrophic or oligotrophic clearwater lakes and tarns without or with very sparse vegetation. One finding is close to a stream. Adults have been found from mid-June to late August. Larvae and larval habitats are unknown. Countries with records of P. fimbriatus in Europe are Finland, Norway and Sweden.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF82090A3CE4FF709F48FDFD.taxon	description	? Procladius albinervis Kieffer, 1918 — Goetghebuer (1927 a), France and Germany, adult female, key, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF82090A3CE4FF709F48FDFD.taxon	materials_examined	Material examined (n = 27). DENMARK, 1 adult male (as Trichotanypus sp., ZMLU), Copenhagen district, 55.67 ° N 12.57 ° E, 5 m a. s. l., + 9 ° C m. a. t., circa 1920. — ENGLAND, 3 adult males (Syntypes of P. flavifrons, BMNH), Lake Windermere, 54.35 ° N 2.94 ° W, 39 m a. s. l., + 9 ° C m. a. t., 9. vii. 1923, leg. F. W. Edwards. — FINLAND, 2 adult males (WIFE), Lake Kuusijärvi, 60.31 ° N 25.11 ° E, 44 m a. s. l., + 5 ° C m. a. t., 18. vii. 1965, leg. P. Armitage; 1 adult male (MZHF), Lake Kuusijärvi, 60.32 ° N 25.09 ° E, 44 m a. s. l., + 5 ° C m. a. t., 28. vii. 1966, leg. B. Lindeberg; 4 adult males (as P. cf. lugens, MZHF), Lake Puruvesi, Punkasalmi, Akonniemi, 61.74 ° N 29.35 ° E, 75 m a. s. l., + 3 ° C m. a. t., 10. vii. 1974, leg. B. Lindeberg; 2 adult males (ZMUO), Asikkalanselkä, 61.26 ° N 25.60 ° E, 74 m a. s. l., + 4 ° C m. a. t., 20. vii. 2015, leg. L. Paasivirta [Barcode CHIFI 548 - 16 and CHFI 549 - 16]. — FRANCE, 1 adult male (NHRS), Remoray National Reserve, 46.78 ° N 6.27 ° E, 853 m a. s. l., + 7 ° C m. a. t., 23. iii. 2017, leg. B. Tissot; 1 adult male (NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 850 m a. s. l., + 7 ° C m. a. t., 7. iv. 2017, leg. B. Tissot; 2 adult males (NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 850 m a. s. l., + 7 ° C m. a. t., iv. 2019, leg. B. Tissot [1 Barcoded]; 1 adult male (NHRS), springs at Lake Remoray, 46.77 ° N 6.26 ° E, 852 m a. s. l., + 7 ° C m. a. t., iv. 2019, leg. B. Tissot [Barcoded]. — NORWAY, 2 adult males (NTNU), Lake Jonsvatn, near Flaten, 63.40 ° N 10.55 ° E, 149 m a. s. l., + 4 ° C m. a. t., 3 ‒ 17. vii. 2014, leg. E. Stur [Barcode CHMNO 216 - 15 and CHMNO 217 - 15]; 1 adult male (NTNU), Lake Jonsvatn, near Flaten, 63.40 ° N 10.55 ° E, 149 m a. s. l., + 4 ° C m. a. t., 31. vii ‒ 14. viii. 2014, leg. E. Stur [Barcode CHMNO 343 - 15]. — SPAIN, 2 adult males (as Procladius sp., DEBE), La Minilla Reservoir, 37.44 ° N 6.10 ° W, 165 m a. s. l., + 17 ° C m. a. t., 27. xi. 1974, leg. N. Prat. — SWEDEN, 1 adult male (NHRS), Lake Färnebofjärden, Dragsheden, 60.24 ° N 16.79 ° E, 60 m a. s. l., + 5 ° C m. a. t., 22. vi. 2008, leg. Y. Brodin [Barcoded]; 1 adult male (NHRS), Lake Vättern, Motala, Råssnäs, 58.53 ° N 14.98 ° E, 88 m a. s. l., + 6 ° C m. a. t., 30. vii. 2012, leg. Y. Brodin [Barcode BSCHI 776 - 17]; 2 adult males (NHRS), Lake Siljan, 60.78 ° N 14.93 ° E, 162 m a. s. l., + 4 ° C m. a. t., 6. vii. 2020, leg. Y. Brodin [1 Barcoded]. Diagnostic characters. Figs. 4, 44, 119 ‒ 121, key couplet 25. P. flavifrons is efficiently identified by the numerous distinct setae of the katepisternum and the lack of macrotrichia on the wing membrane. Other species of Procladius in Europe without wing membrane microtrichia lack katepisternal setae. Two of these species, P. lugens and P. imicola, have similar gonostyli, but P. flavifrons is mostly smaller (wing length 1.7 ‒ 2.3 mm versus 2.1 ‒ 3.4 mm) and has a lower antennal ratio (AR 1.3 ‒ 1.7 versus 1.7 ‒ 2.6). The adult female has been briefly described and the pupal exuvia in detail. The larva is not known. Barcodes of adult males and adult females are available. Geographical distribution and ecology. The geographical distribution of P. flavifrons extends from southern Spain at latitude 37 ° N to Norway at 63 ° N and Finland at 64 ° N. Findings are reported from 17 countries or autonomous regions in Europe. It is mainly an inland freshwater species found up to 920 m above sea level, but some findings are from brackish water at or near the Atlantic coast of Ireland and the Baltic coast of Poland. The climate of sites with findings ranges from + 17 ° C and almost subtropical to temperate and + 4 ° C. Larvae of P. flavifrons inhabit standing freshwater habitats in lakes, reservoirs, ponds and sometimes brackish bays of the sea. There are reports of occurrence in manmade constructions such as fountain ponds in Denmark. P. flavifrons has also been found close to a mountain spring in France and in a rice field in Hungary. Emergence has been reported from 1 ‒ 4 m water depth from bottoms with mud or vegetation dominated by Cladophora. Nutrient conditions in lakes and reservoirs with findings of P. flavifrons are mostly mesotrophic to eutrophic, but same are from oligotrophic conditions. Food sources of the larvae are unknown. Adults have been reported from late March in Ukraine to early December in southern Spain where it can develop at least two generations per year. Countries or autonomous regions with records of P. flavifrons in Europe are Denmark, England, Finland, France, Germany, Hungary, Ireland, Lithuania, Netherlands, Northern Ireland, Norway, Poland, Russia, Spain, Sweden, Ukraine and Wales. It is also known from Asian Russia.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF81090B3CE4FD699B7FFC41.taxon	description	Procladius floralis Kieffer, 1915 — Kieffer (1918 a), Faroe Islands, adult male, adult female, key, description. Psilotanypus floralis (Kieffer, 1915) — Goetghebuer & Lenz (1936 a), Faroe Islands, adult male, adult female, key, description. Procladius sp. — Ratnasingham et al. (2024), Bulgaria, France, Montenegro, Poland, Serbia and Slovakia, adult males, adult females, larvae, photos.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF81090B3CE4FD699B7FFC41.taxon	materials_examined	Material examined (n = 15). CZECHIA, 1 adult male (as Procladius sp., NHRS), Franzensbad, Frantiskovy Lazne, 50.12 ° N 12.34 ° E, 442 m a. s. l., + 7 ° C m. a. t., v. 1879, leg. F. Kowarz; 1 adult male (as Procladius sp., NHRS), Cheb district, Udolni Nadrz Jesenice Reservoir, 50.08 ° N 12.46 ° E, 437 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin; 2 adult males (as Procladius sp., NHRS), Cheb district, Udolni Nadrz Jesenice Reservoir, 50.08 ° N 12.46 ° E, 437 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin; 1 adult male (as Procladiu s sp., NHRS), Cheb district, Udolni Nadrz Jesenice Reservoir, 50.08 ° N 12.46 ° E, 437 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin; 1 adult male (as Procladius sp., NHRS), Cheb district, Lake Vodni Nadrz Skalka, 50.09 ° N 12.32 ° E, 439 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin and M. Gransberg. — ESTONIA, 1 adult male (as Procladius sp., 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. — FINLAND, 1 adult male (as Procladius sp., HECH), Aidassaari, Lake Vanajavesi, 61.17 ° N 24.08 ° E, 79 m a. s. l., + 5 ° C m. a. t., 12 ‒ 15. viii. 1978, leg. P. H. Kansanen. — FRANCE, 2 adult males (as Procladius sp., NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 850 m a. s. l., + 7 ° C m. a. t., iv. 2017, leg. B. Tissot [1 Barcoded]. — GERMANY, 1 adult male (as P. culiciformis, ZSMG), Lake Chiemsee, 47.87 ° N 12.48 ° E, 518 m a. s. l., + 9 ° C m. a. t., 9. vi. 1988, leg. C. Orendt. — SWEDEN, 1 adult male (as Procladius sp., NHRS), Lake Boren, Borenshult, 58.57 ° N 15.10 ° E, 74 m a. s. l., + 6 ° C m. a. t., 11. vi. 1980, leg. Y. Brodin; 1 adult male (as Procladius sp., NHRS), Lake Hålsjön, Marsveden, 59.81 ° N 17.24 ° E, 39 m a. s. l., + 6 ° C m. a. t., 23. v. 1988, leg. J. de Jong; 1 adult male (as Procladius sp., NHRS), Lake Färnebofjärden, Edsviken Bay, 60.29 ° N 16.81 ° E, 59 m a. s. l., + 5 ° C m. a. t., 15. viii. 2007, leg. Y. Brodin; 1 adult male (as Procladius sp., NHRS), Eriksberg nature reserve, Lake Färsksjön, 56.17 ° N 14.99 ° E, 7 m a. s. l., + 7 ° C m. a. t., 13. viii. 2015, leg. J. Wolgast. Diagnostic characters. Figs. 33, 86 ‒ 88, key couplet 15. P. floralis has a rather long gonostylus process with a GspR that overlaps that of ten other species of Procladius in Europe. Of these, P. tenebricosus, P. lugubris, P. frigidus, P. crassinervis, P. fimbriatus and P. signatus can be distinguished from P. floralis by one or more other morphological characters in the key and the helpdesk. The GspR of P. floralis partly overlaps that of P. pruinosus (0.28 ‒ 0.37 versus 0.23 ‒ 0.30). If overlapping, P. floralis can be separated from P. pruinosus by the stronger gonostylus process divergence (30 ‒ 50 ° versus 10 – 25 °) and mostly shorter body length (3.6 ‒ 4.4 mm versus 4.3 ‒ 5.9 mm). The GspR of P. floralis considerably overlaps that of P. longistilus (0.28 ‒ 0.37 versus 0.25 ‒ 0.34). If overlapping, P. floralis can usually be separated from P. longistilus by the stronger gonostylus process divergence (30 ‒ 50 ° versus 5 ‒ 30 °) and usually shorter body length (3.6 ‒ 4.4 mm versus 4.2 ‒ 5.7 mm). P. floralis can be difficult to separate from P. ferrugineus if the phallapodeme of P. ferrugineus is deformed or bleached. The gonostylus process divergence of P. floralis is usually stronger than that of P. ferrugineus (30 ‒ 50 ° versus 10 – 35 °) and the mid leg tibia mostly somewhat longer (0.99 ‒ 1.30 mm versus 0.78 ‒ 1.07 mm). The GspR of P. floralis considerably overlaps that of P. crassinervis (0.28 ‒ 0.37 versus 0.32 ‒ 0.39). The species can nonetheless be separated by a combination of characters. The gonostylus process of P. floralis is less strongly oriented upwards than that of P. crassinervis (10 ‒ 30 ° versus 30 – 60 °), the gonostylus on average narrower (GsmR 5.6 ‒ 6.8 versus 5.2 ‒ 5.9), the gonostylus process divergence weaker (30 ‒ 50 ° versus 40 ‒ 75 °) and the length ratio between the gonostylus and the gonocoxite higher (0.53 ‒ 0.57 versus 0.47 ‒ 0.55). The adult female has been briefly described, but the pupa and larva not. Barcodes of adult males, adult females and larvae are available. Geographical distribution and ecology. P. floralis has been found in Europe from Montenegro at 42 ° N to southern Finland at 61 ° N and the Faroe Islands at 62 ° N. Findings in Bulgaria, Montenegro and Slovakia are based on barcodes. The findings suggest that the species is widespread in Europe. Mean annual temperature of the 19 sites with quality assured findings ranges from + 15 to + 5 ° C and the altitude from 5 to 850 m above sea level. Knowledge of larval habitats of P. floralis is lacking, but they can be expected to inhabit mesotrophic to hypereutrophic lakes and reservoirs judging from the findings of adults. It is possible that the larvae of the species inhabit slowly flowing rivers as well. Adults have been recorded from April to late August. Countries or autonomous regions with records of P. floralis in Europe are Bulgaria, Czechia, Estonia, Faroe Islands, Finland, France, Germany, Montenegro, Poland, Serbia, Slovakia and Sweden.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FF8009753CE4FC359DDBFEDD.taxon	materials_examined	Material examined (n = 17). CANADA, 1 adult male (Holotype of P. gretis, CNCC), Baffin Island, Lake Nettilling, 66.25 ° N 70.05 ° W, 30 m a. s. l., ‒ 9 ° C m. a. t., 8. viii. 1956, leg. J. G. Chillcott; 1 adult male (as Procladius sp. A, CNCC), Ellesmere Island, Lake Hazen, Hazen Camp, 81.5 ° N 71.2 ° W, 158 m a. s. l., ‒ 14 ° C m. a. t., 17. vii. 1961, leg. D. R. Oliver. — FINLAND, 1 adult male (as P. pectinatus, ZMUO), Lake Kankareenjärvi, 60.44 ° N 22.96 ° E, 80 m a. s. l., + 5 ° C m. a. t., 20. iv. 2015, leg. L. Paasivirta [Barcode LEFIJ 3520 - 16]; 2 adult males (as Procladius sp., ZMUO), Lake Nummijärvi, 60.59 ° N 23.25 ° E, 97 m a. s. l., + 4 ° C m. a. t., 4. v. 2015, leg. L. Paasivirta [Barcode LEFIJ 3905 - 16 and LEFIJ 3906 - 16]; 2 adult males (as Procladius sp., ZMUO), Lake Iso-Ruostejärvi, 60.69 ° N 23.80 ° E, 117 m a. s. l., + 4 ° C m. a. t., 5. v. 2015, leg. L. Paasivirta [Barcode LEFIJ 3670 - 16 and LEFIJ 3671 - 16]. — GREENLAND, 3 adult males (as P. choreus, ZMUC), Cape Oswald Peninsula, Franz Josef Fjord district, 72.5 ° N 25.0 ° W, 10 m a. s. l., ‒ 7 ° C m. a. t., vii ‒ viii. 1934, leg. F. S. Andersen. — NORWAY, 1 adult male (Holotype of Tanypus frigidus, NHRS), Bear Island, Mount Misery, 74.42 ° N 19.20 ° E, 300 m a. s. l., ‒ 3 ° C m. a. t., 25. viii. 1868, leg. A. E. Holmgren; 1 adult male (as P. choreus, MTSN), Svalbard, Progetto, Londonelva 3, 78.58 ° N 12.04 ° E, 355 m a. s. l., ‒ 6 ° C m. a. t., 13. viii. 1997, leg. L. Marziali; 1 adult male (NTNU), Svalbard, Adventdalen, Todalen below mine cave 5, 78.16 ° N 15.83 ° E, 100 m a. s. l., ‒ 5 ° C m. a. t., 26. vii. 2005, leg. O. Frengen [Barcode MIDGE 069 - 06]; 1 adult male (NTNU), Svalbard, Kapp Linné, Lake Søndre Borgdam, 78.07 ° N 13.79 ° E, 35 m a. s. l., ‒ 5 ° C m. a. t., 8. viii. 2008, leg. T. Ekrem and K. Hårsaker [Barcode CHRSV 120 - 08]; 1 adult male (NTNU), Kautokeino, Lahpoluoppal, lake, 69.21 ° N 23.76 ° E, 321 m a. s. l., ‒ 2 ° C m. a. t., 12. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 033 - 10]; 1 adult male (NTNU), Porsanger, Lake Øvrevatn, Skoganvarre, 69.84 ° N 26.08 ° E, 76 m a. s. l., 0 ° C m. a. t., 16. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 426 - 11]. — RUSSIA, 1 adult male (as Tanypus signatus, ZISP), Kotelny Island, River Wosnessenje, 75.16 ° N 145.15 ° E, 300 m a. s. l., ‒ 18 ° C m. a. t., 23. vii. 1903, leg. M. I. Brussnew. Diagnostic characters. Figs. 32, 83 ‒ 85, key couplet 14. P. frigidus has a medium long gonostylus process with a GspR that overlaps that of eight other species of Procladius in Europe. Of these, P. culiciformis, P. tenebricosus, P. ferrugineus, P. floralis and P. crassinervis are easily separated from P. frigidus by the absence of setae on the median anepisternum and one or more other characters in the key and helpdesk. It is neither a problem to distinguish P. frigidus from P. lugubris by characters such as gonostylus process divergence (10 ‒ 25 ° versus 25 ‒ 45 °), front leg tarsi BR (3 ‒ 4.5 versus 5.5 ‒ 8) and the number of spines of the hind leg tibial comb (10 ‒ 14 versus 6 ‒ 9). The GspR of P. frigidus strongly overlaps that of P. pruinosus (0.25 ‒ 0.33 versus 0.23 ‒ 0.30) and completely that of P. longistilus (0.25 ‒ 0.33 versus 0.25 ‒ 0.34). P. frigidus can be separated from the other species by a combination of other characters, particularly the number of median anepisternal setae (5 ‒ 26 versus 0 ‒ 6), antenna AR (2.2 ‒ 2.8 versus 1.7 ‒ 2.3) and the length of palpomere five divided with its width (8.0 ‒ 9.3 versus 9.1 ‒ 12.2). P. frigidus is mainly an Arctic, cold-adapted species (mean annual temperature + 5 to ‒ 15 ° C), while P. pruinosus mostly lives in warmer temperature conditions (mean annual temperature + 18 to ‒ 1 ° C). P. frigidus has been regarded as a synonym of P. crassinervis for a hundred years (Edwards 1924) until recently when Stur & Ekrem (2020) reestablished P. frigidus as a valid species name. The adult female and larva have been described. The pupa has not been described. Barcodes of adult males, adult females, a pupa and larvae are available. Geographical distribution and ecology. As its Latin name implies, P. frigidus is a cold-adapted species. It is in fact the most northerly distributed of all Procladius species in the northern hemisphere, with findings up to latitude 79 ° N in Svalbard Island of Norway and 82 ° N in Ellesmere Island of Canada. The southernmost records are from southern Finland at 60 ° N. Altitude of the 33 sites with quality assured records ranges from about 2 m to about 360 m above sea level. Mean annual temperature extends from + 5 in Finland to ‒ 18 ° C on an Arctic island near the Arctic Ocean coast of Siberian Russia. This is the coldest condition in terms of mean annual temperature known for a species of Procladius and other species of Chironomidae, in fact among the coldest known for any insect species. Noteworthy is that there are at least fifty species of Chironomidae known from sites with mean annual temperature ranging from ‒ 14 to ‒ 18 ° C. The northernmost ponds and lakes where the larvae of P. frigidus live can have a complete ice cover lasting for eight to more than ten months a year. Larvae have been found from 0 to 9 m water depth and are able to hibernate in a frozen state in bottom sediments for at least five months. Bottoms of these lakes can be devoid of oxygen and contain poisonous hydrogen sulfide. The larvae of P. frigidus have been reported to be fast swimmers, even swimming backwards, and to feed on smaller chironomids and crustaceans. The ponds and lakes with larvae are ultraoligotrophic to oligotrophic and mostly free of fish, but some contain Arctic char (Salvelinus alpinus). The development of P. frigidus from the first instar larvae to adult may take three years or more in the coldest lakes with water temperature that rarely exceed + 4 ° C. They may overwinter as inactive prepupae. As for other Chironomidae of Arctic lakes and ponds, emergence of P. frigidus can take place as soon as cracks in the ice appear in mid-June or as late as early July. Emergence can continue for about three weeks, which is substantially longer than that for most other insect species in these lakes. Summer weather even at the coldest sites can reach more than + 20 ° C, so conditions for activity of the adults such as mating and egg-laying can be favourable. Adults have been caught from mid-April to early June in southern Finland and from mid-June to mid-August in colder climate conditions. Adults of P. frigidus have been found to be common in the diet of snow buntings (Plectrophenax nivalis) in Svalbard, Norway, and in southern Greenland. Adults are also known to be food items of other birds such as dunlins and sanderlings (Calidris alpina and C. alba). Countries with records of P. frigidus in Europe are Finland and Norway. It is also present in Canada, Greenland and Russia (Asia).	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFFE09753CE4FE899C32FBBD.taxon	materials_examined	Material examined (n = 3). FINLAND, 1 adult male (Paratype of P. gemma, as Procladius sp.), Galddoaivi, Sullamintie, 69.9 ° N 27.0 ° E, 90 m a. s. l., 0 ° C m. a. t., 8. vii. 2003, leg. J. Salmela. — RUSSIA, 1 adult male (Holotype of P. gemma, IANR), tundra ponds, east of Naryan-Mar, Yarey-shor, 67.62 ° N 53.70 ° E, 20 m a. s. l., ‒ 3 ° C m. a. t., 29. vii. 1990, leg. A. Rybakova; 1 adult male (Paratype of P. gemma, IANR), tundra ponds, east of Naryan-Mar, Yarey-shor, 67.62 ° N 53.70 ° E, 20 m a. s. l., ‒ 3 ° C m. a. t., 29. vii. 1990, leg. A. Rybakova. Diagnostic characters. Figs. 21, 50 ‒ 52, key couplet 5. P. gemma is easily distinguished from other species of Procladius by the ball-formed projection of tergite IX and the special form of the medioapodeme. Together with the very different P. flavifrons, it is the only species of Procladius in Europe with more than 4 setae on the katepisternum. Neither the female, pupa nor larva are known. No barcodes have been identified. Geographical distribution and ecology. The two sites where adults of P. gemma have been collected lay at latitude 68 and 70 ° N in northern Russia and northernmost Finland. The altitude of the sites is 90 m and 20 m respectively. Mean annual temperature ranges from 0 to ‒ 3 ° C. Both sites are rather pristine palsa mires with discontinuous permafrost, containing small lakes and ponds in the Finnish site and only small ponds in the Russian one. The sites and their surroundings are treeless or with low birch forest somewhat influenced by domesticated reindeer (Rangifer tarandus). Larval ecology of P. gemma is unknown, but the findings suggest that it is a cold-stenothermal inhabitant of shallow ponds. Adult males have been found during July. P. gemma is probably a rare species, since it is an easily identified and relatively large species of Chironomidae with unique genitalia characters. Countries with records of P. gemma are Finland and Russia.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFFD09703CE4F9379B35FBD1.taxon	materials_examined	Material examined (n = 18). CANADA, 1 adult male (Holotype of P. nietus, CNCC), Atlin, 59 ° N 133 ° W, 670 m a. s. l., 0 ° C m. a. t., 9. vi. 1955, leg. H. Hackel; 1 adult male (as P. nietus, CNCC), Lake Manitoba, Bone Pile Pond, 50.11 ° N 98.19 ° W, 248 m a. s. l., + 3 ° C m. a. t., 18. v. 1980, leg. D. A. Wrubleski; 1 adult male (as P. nietus, ANSP), Lake Manitoba, Bone Pile Pond, 50.11 ° N 98.19 ° W, 248 m a. s. l., + 3 ° C m. a. t., 11. vii. 1980, leg. D. A. Wrubleski. — FINLAND, 1 adult male (as P. appropinquatus, ZMUO), Lake Littoistenjärvi, 60.46 ° N 22.38 ° E, 36 m a. s. l., + 6 ° C m. a. t., 20. v. 2013, leg. L. Paasivirta [Barcode LEFIJ 5880 - 17]; 1 adult male (ZMUO), Hamarijärvi, Salo, 60.21 ° N 22.94 ° E, 34 m a. s. l., + 6 ° C m. a. t., 18. v. 2014, leg. L. Paasivirta [Barcode LEFIJ 3836 - 16]; 2 adult males (ZMUO), Tammela, Lake Suujärvi, 60.77 ° N 23.85 ° E, 97 m a. s. l., + 4 ° C m. a. t., 5. v. 2015, leg. L. Paasivirta [Barcode LEFIJ 3665 - 16 and LEFIJ 3666 - 16]. — GERMANY, 1 adult male (as P. flavifrons, ZSMG), River Unterer Schierenseebach, 54.3 ° N 10.0 ° E, 6 m a. s. l., + 8 ° C m. a. t., 13. v. 1985, leg. U. Holm. — NETHERLANDS, 1 adult male (as P. lugens, NMNL), Leiden, 52.2 ° N 4.5 ° E, 1 m a. s. l., + 11 ° C m. a. t., 2014. — RUSSIA, 3 adult males (IBIB), Rybinsk Reservoir, 58.1 ° N 38.6 ° E, 97 m a. s. l., + 4 ° C m. a. t., vii. 1972, leg. A. I. Shilova. — SCOTLAND, 1 adult male (as P. lugens, BMNH), Aviemore, 57.19 ° N 3.83 ° E, 214 m a. s. l., + 7 ° C m. a. t., 16. v. 1947. — SWEDEN, 1 adult male (NHRS), Lake Tullan, 59.22 ° N 17.70 ° E, 25 m a. s. l., + 6 ° C m. a. t., 16. v. 1989, leg. M. Gransberg; 2 adult males (SLUU), Mälaren, Drottningholm, 59.33 ° N 17.87 ° E, 3 m a. s. l., + 6 ° C m. a. t., 21. v. 1991, leg. L. Eriksson and T. Wiederholm; 1 adult male (NHRS), Tärnsjö, Nordmyra, temporary inundated wetland, 60.16 ° N 16.89 ° E, 64 m a. s. l., + 5 ° C m. a. t., 1. v. 2007, leg. A. Hagelin; 1 adult male (NHRS), Eriksberg nature reserve, Lake Gårdsjön, 56.17 ° N 15.00 ° E, 14 m a. s. l., + 7 ° C m. a. t., 1. v. 2017, leg. J. Wolgast [Barcoded]. Diagnostic characters. Figs. 10, 46, 125 ‒ 127, key couplet 26. P. imicola has an almost unidentifiable to very short gonostylus process with a GspR that overlaps that of four other species of Procladius in Europe. Of these, P. flavifrons, P. simplicistilus and P. appropinquatus are easily separated from P. imicola by several characters including gonostylus form, setae on katepisternum, size and numbers of macrotrichia on the wing membrane. The only similar species in Europe is P. lugens. The genitalia of P. imicola is very similar that of P. lugens, but the outer margin of the base section of the gonostylus is usually distinctly angled in P. imicola, while rounded in P. lugens. P. imicola is on average larger (wing length 2.6 ‒ 3.4 mm versus 2.1 ‒ 2.8 mm, mid leg tibia length 1.1 ‒ 1.4 mm versus 0.8 ‒ 1.1 mm, body length 4.8 ‒ 5.5 mm versus 3.5 ‒ 4.6 mm) and has an antenna AR only slightly overlapping (2.2 ‒ 2.6 versus 1.7 ‒ 2.3) that of P. lugens. P. imicola has mostly been correctly identified in taxonomic literature. The adult female, pupa and larva have been described. Barcodes of adult males are available. Geographical distribution and ecology. The known geographical distribution of P. imicola in Europe extents from the Netherlands and England at 52 ° N to Finland at 66 ° N, encompassing sites with mean annual temperature which range from + 11 to + 2 ° C. Down to ‒ 4 ° C mean annual temperature is reached in Asian Russia and Canada. P. imicola has been found at altitudes from 2 m to about 700 m above sea level in Europe. Findings in Canada are up to 1 020 m, while about 1 640 m in Mongolia. The smaller but morphologically very similar P. lugens is only known from altitudes up to 540 m above sea level. A great majority of the 28 sites with quality assured findings of P. imicola are oligotrophic to hypereutrophic lakes or lake-like reservoirs. Findings of P. imicola larvae are often from the profundal zone from 2 m down to 30 m, while its close relative P. lugens has mostly been found in the littoral and not known from more than 5 m depth in the profundal. P. imicola can be among the most abundant species of the macroscopic bottom fauna in deep water of eutrophic and hypereutrophic lakes and reservoirs, showing a profound ability to withstand low oxygen conditions. More than 1 000 individuals of 3 rd to 4 th stage larvae per m 2 and about 1 g wet weight per m 2 have been reported. Rapid growth of the larvae may take place in autumn due to improved oxygen conditions after summer stagnation. Larvae of P. imicola are also known from littoral bottoms with different kinds of vegetation including mosses, Cladophora, Potamogeton, Typha and Phragmites. There are scattered findings of P. imicola from habitats other than freshwater lakes or reservoirs, namely temporary ponds of an inundated wetland, rivers and canals with slowly floating water and a mountain lake with salinity equal to or even higher than typical seawater. P. imicola have been observed to feed on animals such as Cladocera, Copepoda, Ostracoda and other Chironomidae, but also algae such as diatoms, fungal spores and detritus. Experiments have suggested that the larvae cannot or have difficulties to complete development to adults without some animal food. Adults of P. imicola have been found from mid-April to mid-August. Synchronized emergence during about one week has been reported. Swarming of males over light-coloured objects has been observed. Larvae have been reported to overwinter in their last fourth instar. Countries or autonomous regions with records of P. imicola in Europe are England, Estonia, Finland, Germany, Latvia, Lithuania, Netherlands, Poland, Russia, Scotland and Sweden. Records are also known from Canada, Mongolia and Russia (Asia).	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFFB09723CE4FB859BB4FD45.taxon	materials_examined	Material examined (n = 29). BELGIUM, 2 adult males (as Procladius sp., RBNS), Gand (Ghent), 51.05 ° N 3.72 ° E, 12 m a. s. l., + 10 ° C m. a. t., 19. iv. 1919, 4. viii. 1919, leg. M. Goetghebuer. — CANADA, 1 adult male (Holotype of P. vesus, CNCC), Lake Muskox, 64.63 ° N 108.25 ° W, 343 m a. s. l., ‒ 9 ° C m. a. t., 25. vii. 1953, leg. J. G. Chillcott. — FAROE ISLANDS, 2 adult males (NHRS), Lake Leynavatn, 62.12 ° N 7.03 ° W, 74 m a. s. l., + 5 ° C m. a. t., 27. vii. 1980, leg. G. Brodin-Lindsten; 3 adult males (NHRS), Lake Eithis, 62.17 ° N 7.06 ° W, 135 m a. s. l., + 5 ° C m. a. t., 15. viii. 2002, leg. Y. Brodin [1 Barcoded]; 2 adult males (NHRS), Lake Toftavatn, 62.10 ° N 6.72 ° W, 15 m a. s. l., + 5 ° C m. a. t., 15. viii. 2002, leg., Y. Brodin. — FINLAND, 1 adult male, Galddoaivi, Sullamintie, 69.9 ° N 27.0 ° E, 90 m a. s. l., 0 ° C m. a. t., 8. vii. 2003, leg. J. Salmela; 1 adult male (as P. cf. vesus), Malax, Rönnskären island, 63.06 ° N 20.83 ° E, 5 m a. s. l., + 4 ° C m. a. t., 30. vii ‒ 5. viii. 2018, leg. L. Paasivirta. — GERMANY, 1 adult male (as P. sagittalis, ZSMG), Kossau stream, Kossautal, 54.27 ° N 10.55 ° E, 13 m a. s. l., + 9 ° C m. a. t., 1991, leg. Schröder; 3 adult males (as P. choreus, ZFMK), Wersabe, Fleet, meadow, 52.51 ° N 9.38 ° E, 37 m a. s. l., + 10 ° C m. a. t., 6. vii. 2017 [Barcode GBOL- 2597108, GBOL- 2597110 and GBOL- 2597112]. — ICELAND, 1 adult male (Syntype of P. islandicus, RBNS) Grimsstadir, 65.65 ° N 16.11 ° W, 395 m a. s. l., 0 ° C m. a. t., 20 ‒ 22. vii. 1927, leg. C. H. Lindroth; 1 adult male (NHRS), River Laxá, 65.59 ° N 17.17 ° W, 253 m a. s. l., + 3 ° C m. a. t., 12. vii. 1978, leg. Y. Brodin. — NORWAY, 2 adult males (NTNU), Einarsneset, pond at Risbakk, 58.06 ° N 6.79 ° E, 1 m a. s. l., + 8 ° C m. a. t., 25. viii. 2019, 26. vi. 2020, leg. E. Stur and P. Dominiak [Barcode MMCH 050 - 19 and MMCH 222 - 20]. — SWEDEN, 1 adult male (NHRS), Lake Innaren, 56.98 ° N 14.97 ° E, 174 m a. s. l., + 6 ° C m. a. t., 6. viii. 1943, leg. L. Brundin; 1 adult male (Syntype of P. fuscus, NHRS), Lake Grimsberga göl, 56.97 ° N 14.92 ° E, 193 m a. s. l., + 6 ° C m. a. t., 12. vi. 1947, leg. L. Brundin. — UNITED STATES, 3 adult males (as Procladius sp., NHRS), Crater Lake, Wizard Island, 42.93 ° N 122.01 ° W, 1 883 m a. s. l., + 7 ° C m. a. t., 10. viii. 1957, leg. L. Brundin; 2 adult males (as Procladius sp., NHRS), Crowley Lake, 37.62 ° N 118.74 ° W, 2 063 m a. s. l., + 8 ° C m. a. t., 21. viii. 1957, leg. L. Brundin; 2 adult males (as P. vesus, NDSU), Alaska, Utqiagvik, tundraponds, 71.3 ° N 156.8 ° W, 7 ‒ 12 m a. s. l., ‒ 12 ° C m. a. t., vi. 1975, vi. 1976, leg. M. G. Butler and S. Mozley. Diagnostic characters. Figs. 7, 15, 38, 101 ‒ 103, key couplet 20. P. islandicus has a rather short gonostylus process with a GspR that overlaps that of nine other species of Procladius in Europe. Of these, P. nudipennis, P. gemma, P. tenebricosus and P. lugubris are distinctly separated from P. islandicus by more than one other morphological character in the key and the helpdesk. P. islandicus has a highly variable length of the setae of the front leg tarsi (BR 3 ‒ 6.5). When the BR is 4.5 or more, as mostly in the northern colder part of the species geographical distribution, P. islandicus can usually be separated from all other European species of Procladius with overlapping GspR, but for P. exilis. P. islandicus can however be distinguished from P. exilis by a higher number of hind tibia comb spines (10 ‒ 13 versus 7 ‒ 9) and fewer median anepisternal setae (0 ‒ 3 versus 10 ‒ 18). The GspR of P. islandicus partly overlaps that of P. choreus (0.18 ‒ 0.24 versus 0.13 ‒ 0.20). If overlapping, the species can almost always be separated by a combination of characters. P. islandicus has a higher front leg tarsi BR than that of P. choreus (3 ‒ 6.5 versus 1.5 ‒ 3), it is usually larger (wing length 2.6 ‒ 3.5 mm versus 1.8 ‒ 2.8 mm) and with an on average darker posterior colour of tergite II ‒ IV (brown to dark brown not distinctly contrasting with the anterior part of the tergites versus whitish to light brown contrasting with the clearly darker anterior part of the tergites). The GspR of P. islandicus substantially overlaps that of P. saeticubitus (0.18 ‒ 0.24 versus 0.14 ‒ 0.20), but the species can be distinguished by combining other morphological characters. P. islandicus has frequently a higher front leg tarsi BR than that of P. saeticubitus (BR 3 ‒ 6.5 versus 2 ‒ 4), the gonostylus is on average broader (GsmR 5.3 ‒ 6.4 versus 5.9 ‒ 6.9), the Cu stem setae usually fewer (0 ‒ 14 versus 5 ‒ 33) and palpomere five measured as length divided by width usually broader (9.9 ‒ 11.9 versus 10.7 ‒ 12.3). P. islandicus has an almost uncoloured to slightly greyish wing membrane, whereas the wing membrane of P. saeticubitus is greyish to conspicuously grey. The GspR of P. islandicus entirely overlaps that of P. culiciformis (0.18 ‒ 0.24 versus 0.18 ‒ 0.25). The species can be separated by the narrower gonostylus of P. islandicus compared with that of P. culiciformis (GsmR 5.3 ‒ 6.4 versus 4.2 ‒ 5.2), the on average longer setae of the front leg (BR 3 ‒ 6.5 versus 2 ‒ 4.5) and an on average lighter mid-section of the front leg tibia (light brown to dark brown versus whitish to light brown). The GspR of P. islandicus slightly overlaps that of P. pruinosus (0.18 ‒ 0.24 versus 0.23 ‒ 0.30). If overlapping, P. islandicus can be separated from P. pruinosus by the on average longer front leg tarsi BR (3 ‒ 6.5 versus 2 ‒ 4.5) and the broader gonostylus process (length / width 0.9 ‒ 1.2 versus 1.2 ‒ 1.7). P. islandicus is usually named correctly in literature regarding findings in Europe but named P. vesus in North America. The adult female, pupa and larva have been briefly described. Barcodes of adult males are available. Geographical distribution and ecology. In Europe, P. islandicus has a latitude range stretching from 48 ° N in southern Germany to 70 ° N in northernmost Finland. In Canada and the United States, the range is from 38 ° N to 71 ° N. Altitude records in Europe are from 1 m to around 640 m in Iceland, while in North America up to 2 060 m in the United States. P. islandicus has among the widest climate adaptability of all Procladius species, expressed as a mean annual temperature span of 22 ° C ranging from about + 10 ° C in temperate Germany and Belgium to about ‒ 12 ° C in Arctic Alaska in the United States. Larvae of P. islandicus inhabit shallow water over bottoms with vegetation such as mosses, Carex and Cladophora in puddles, ponds or lakes in wet meadows or tundra. Bottoms with mud or lava sand without vegetation are also habitats for the larvae. No findings below 4 m water depth are known. A few findings of adults have been made close to slowly running to rather fast running water and one finding is from a natural spring with pH 9.5. Ice cover on the water bodies containing P. islandicus larvae may remain 6 ‒ 8 months per year in northern Europe and 7 ‒ 9 months per year in northern North America. Completion of the larval life cycle may take at least four years in the coldest waters. Oligotrophic to eutrophic conditions prevail in the water bodies with findings. P. islandicus larvae have been observed to prey on other smaller Chironomidae larvae such as first and second instars of Chironomus. Adults have been found from early May to the middle of September. Countries or autonomous regions with records of P. islandicus in Europe are Belgium, Faroe Islands, Finland, Germany, Iceland, Lithuania, Norway, Russia, Scotland and Sweden. There are also records from Canada and the United States.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF9097C3CE4FD319B2FFB05.taxon	description	Trichotanypus longistilus Kieffer, 1916 — Kieffer (1918 a), Lithuania, adult male, key, description. Trichotanypus longistilus Kieffer, 1916 — Kieffer (1924), adult male, key. Trichotanypus longistylus Kieffer, 1916 — Goetghebuer (1927), France, adult male, key, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF9097C3CE4FD319B2FFB05.taxon	description	Procladius longistylus (Kieffer, 1916) — Ratnasingham et al. (2024), Netherlands, adult male, photo.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF9097C3CE4FD319B2FFB05.taxon	materials_examined	Material examined (n = 58). CHINA, 1 adult male (as P. nigriventris), Huaping Nature Reserve, 25.56 ° N 109.94 ° E, 1 271 m a. s. l., + 12 ° C m. a. t., 24. vi. 2020, leg. S. Zhao [Barcode CHGX 127 - 20]. — CZECHIA, 2 adult males (NHRS), Cheb district, Lake Vodni Nadrz Skalka, 50.08 ° N 12.32 ° E, 439 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin. — FINLAND, 4 adult males (as P.? nigriventris, MZHF), Lake Kuusijärvi, 60.31 ° N 25.11 ° E, 44 m a. s. l., + 5 ° C m. a. t., 18. vii. 1965, leg. P. Armitage; 1 adult male (as P.? choreus, MZHF), Lake Kuusijärvi, 60.31 ° N 25.11 ° E, 44 m a. s. l., + 5 ° C m. a. t., 28. vii. 1966, leg. P. Armitage; 1 adult male (as Procladius sp., MZHF), Lake Vanajavesi, Vihdassaari Island, 61.18 ° N 24.03 ° E, 79 m a. s. l., + 5 ° C m. a. t., 8. vi. 1978, leg. P. H. Kansanen; 2 adult males (as P. culiciformis), Suvakoski, small lake, 67.31 ° N 28.16 ° E, 175 m a. s. l., + 1 ° C m. a. t., 9. vi ‒ 23. vii. 2009, leg. L. Paasivirta; 1 adult male (as P. culiciformis coll., ZMUO), Lake Pyhäjärvi, Säkylä, 61.00 ° N 22.28 ° E, 45 m a. s. l., + 5 ° C m. a. t., 4. vi. 2015, leg. L. Paasivirta [Barcode LEFIJ 3517 - 16]; 1 adult male (as Procladius sp.), Baltic Sea, Lake Käringsund, Storskär Island, 63.43 ° N 21.08 ° E, 0 m a. s. l., + 4 ° C m. a. t., 15. v. 2017, leg. L. Paasivirta; 1 adult male, Tulppio, Ainijärvi, 67.76 ° N 29.45 ° E, 269 m a. s. l., 0 ° C m. a. t., 10 ‒ 14. vi. 2018, leg. J. Salmela; 2 adult males, Kiikala, Lake Tervakas, 60.49 ° N 23.70 ° E, 117 m a. s. l., + 5 ° C m. a. t., 9. v. 2023, leg. L. Paasivirta. — FRANCE, 3 adult males (as Procladius sp. 1, LHST), Lake Gourg Nére, 42.87 ° N 0.18 ° E, 2 201 m a. s. l., + 2 ° C m. a. t., 10. viii. 1966, leg. H. Laville; 3 adult males (NHRS), Corsica, Lake Melu, 42.21 ° N 9.02 ° E, 1 708 m a. s. l., + 6 ° C m. a. t., 25. viii. 2015, leg. J. Moubayed-Breil. — GERMANY, 1 adult male (as. P. sagittalis, ZSMG), Lake Schalkenmehrener Maar, 50.2 ° N 6.9 ° E, 422 m a. s. l., + 8 ° C m. a. t., 1914, leg. A. Thienemann; 1 adult male (as P. bathocryptus, ZSMG), Lake Schluensee, 54.19 ° N 10.46 ° E, 22 m a. s. l., + 9 ° C m. a. t., 1953, leg. I. Müller-Liebenau; 1 adult male (as Procladius sp., ZSMG), Hödinger Tobel Stauweiher, small reservoir, 47.79 ° N 9.13 ° E, 551 m a. s. l., + 9 ° C m. a. t., 3. vi. 1966, leg. F. Reiss; 1 adult male (as P. cf. choreus, ZSMG), Lahnberge, near Biologisches Institut Marburg, 50.81 ° N 8.81 ° E, 340 m a. s. l., + 9 ° C m. a. t., 10. vi. 1992, leg. A. Dettinger-Klemm; 1 adult male (as Procladius sp., ZFMK), Kniepow, Rügen, small lake, 54.35 ° N 13.35 ° E, 9 m a. s. l., + 9 ° C m. a. t., 16. viii. 2014 [Barcode GBMTM 0349 incomplete]; 1 adult male (as Procladius sp., ZFMK), Brettenbach stream, Brücke Vordere Zaismatt, 48.13 ° N 7.90 ° E, 242 m a. s. l., + 10 ° C m. a. t., 31. iii. 2019. — IRELAND, 2 adult males (as Procladius sp. D, UCDZ), Lacy’s canal at Lake Ennell, 53.49 ° N 7.37 ° W, 78 m a. s. l., + 10 ° C m. a. t., 18. iv. 1974, leg. D. A. Murray. — NORWAY, 1 adult male (as P. cf. nigriventris, NTNU), Kautokeino, Lapholuoppal, lake, 69.21 ° N 23.76 ° E, 320 m a. s. l., ‒ 2 ° C m. a. t., 12. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 040 - 10]; 1 adult male (as P. cf. nigriventris, NTNU), Alta, Lake Sierravannet, 69.84 ° N 23.37 ° E, 43 m a. s. l., ‒ 1 ° C m. a. t., 14. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 079 - 10]; 1 adult male (as P. cf. nigriventris, NTNU), Vardø, Lake Nedre Domsvatn, 70.32 ° N 31.03 ° E, 120 m a. s. l., ‒ 1 ° C m. a. t., 18. vi. 2010, leg. T. Ekrem and E. Stur [Barcode ATNA 531 - 10]; 1 adult male (as P. cf. nigriventris, NTNU), Melhus, Gammelelva nature reserve, 63.22 ° N 10.31 ° E, 28 m a. s. l., + 4 ° C m. a. t., 22. v. 2014, leg. E. Stur [Barcode CHMNO 204 - 15]; 1 adult male (as P. cf. nigriventris, NTNU), Bymarka, Lake Kobberdammen, 63.42 ° N 10.28 ° E, 288 m a. s. l., + 3 ° C m. a. t., 27. v. 2014, leg. T. Ekrem and X. Lin [Barcode CHMNO 097 - 14]. — RUSSIA, 1 adult male (NHRS), Lake Khepoyarech, 60.17 ° N 30.56 ° E, 61 m a. s. l., + 5 ° C m. a. t., 3. vii. 1980, leg. Y. Brodin; 2 adult males (TUSF), Lake Ladoga, Sorvala, 61.74 ° N 30.77 ° E, 4 m a. s. l., + 5 m. a. t., 4. viii. 1991, leg. G. Söderman [Barcoded]. — SLOVAKIA, 1 adult male (NHRS), Lake Strbske Pleso, 49.12 ° N 20.06 ° E, 1 349 m a. s. l., + 3 ° C m. a. t., 20. vi. 1991, leg. M. Gransberg. — SPAIN, 1 adult male (as P. choreus, DEBE), Ebro Reservoir, Burgos-Santander, 42.58 ° N 4.02 ° W, 215 m a. s. l., + 13 ° C m. a. t., 28. i. 1975, leg. N. Prat. — SWEDEN, 1 adult male (as Tanypus sp., ZMLU), Åreskutan, 63.4 ° N 13.0 ° E, 920 m a. s. l., ‒ 1 ° C m. a. t., 31. vii ‒ 4. viii. 1840, leg. J. W. Zetterstedt; 1 adult male (NHRS), Lake Innaren, 56.98 ° N 14.97 ° E, 175 m a. s. l., + 6 ° C m. a. t., 21. v. 1947, leg. L. Brundin; 1 adult male (NHRS), Lake Flarken, north of Kvarnemon, 58.56 ° N 13.67 ° E, 129 m a. s. l., + 6 ° C m. a. t., 1. vi. 1981, leg. Y. Brodin; 1 adult male (NHRS), Lake Vänern, Mariestadsfjärden Bay, 58.70 ° N 13.77 ° E, 43 m a. s. l., + 6 ° C m. a. t., 7 ‒ 9. v. 1987, leg. Y. Brodin; 1 adult male (NHRS), Lake Vänern, Kinneviken Bay, Truveholm, 58.50 ° N 13.26 ° E, 44 m a. s. l., + 7 ° C m. a. t., 21. v. 1987, leg. Y. Brodin and U. Häll; 2 adult males (NHRS), Lake Yngern, 59.14 ° N 17.44 ° E, 41 m a. s. l., + 6 ° C m. a. t., 12. v. 1991, leg. T. Sandberg; 4 adult males (NHRS), Lake Voutner, 65.65 ° N 19.70 ° E, 351 m a. s. l., + 1 ° C m. a. t., 1. viii. 1992, leg. Y. Brodin and K. Murray; 1 adult male (NHRS), Kåbdalis, Kåbdalisjaure, 66.14 ° N 19.99 ° E, 330 m a. s. l., 0 ° C m. a. t., vii. 1993, leg. B. Viklund; 1 adult male (NHRS), Laggarbo wetland, west of Österfärnebo, 60.27 ° N 16.79 ° E, 62 m a. s. l., + 5 ° C m. a. t., 20. vi ‒ 5. vii. 2002, leg. T. Persson Vinnersten; 2 adult males (NHRS), Baltic Sea, Askö island, Askötorp, 58.80 ° N 17.67 ° E, 6 m a. s. l., + 6 ° C m. a. t., 31. v ‒ 16. vi. 2011, leg. B. E. Bengtsson. — UNITED STATES, 2 adult males (as P. culiciformis, ANSP), Lake Lacawac, 41.38 ° N 75.29 ° W, 439 m a. s. l., + 8 ° C m. a. t., 7. viii. 1963, leg. S. S. Roback. Diagnostic characters. Figs. 19, 35, 92 ‒ 94, key couplet 18. P. longistilus has a long gonostylus process with a GspR that overlaps that of nine other species of Procladius in Europe. Of these, P. culiciformis, P. lugubris, P. ferrugineus, P. crassinervis and P. fimbriatus are distinctly distinguished from P. longistilus by more than one other morphological character in the key and the helpdesk. The GspR of P. longistilus considerably overlaps that of P. tenebricosus (0.25 ‒ 0.34 versus 0.23 ‒ 0.29). If overlapping, P. longistilus can be separated from P. tenebricosus by the less dark marked wing (anal cell patch absent to faint versus distinct to very distinct) and mostly larger size (wing length 2.4 ‒ 3.5 mm versus 1.8 ‒ 2.6 mm, mid leg tibia length 1.1 ‒ 1.5 mm versus 0.7 ‒ 1.1 mm, body length 4.2 ‒ 5.7 mm versus 3.2 ‒ 4.3 mm). The GspR of P. longistilus considerably overlaps that of P. pruinosus (0.25 ‒ 0.34 versus 0.23 ‒ 0.30). P. longistilus can usually be separated from P. pruinosus by the ratio between the length of the outer phallapodeme section divided by the length of the sternapodeme (0.34 ‒ 0.46 versus 0.27 ‒ 0.36). The length of the outer phallapodeme is however frequently difficult to measure accurately and requires good slide preparation. In addition, the on average narrower gonostylus and gonostylus process of P. longistilus than that of P. pruinosus (GsmR 5.5 ‒ 6.9 versus 5.2 ‒ 6.3, gonostylus process length / width 1.3 ‒ 1.9 versus 1.2 ‒ 1.7) can be useful for species separation. The GspR of P. longistilus completely overlaps that of P. frigidus (0.25 ‒ 0.34 versus 0.25 ‒ 0.33). P. longistilus can be separated from P. frigidus by combining measurements of other characters, particularly the number of median anepisternal setae (0 ‒ 6 versus 5 ‒ 26), antenna AR (1.8 ‒ 2.3 versus 2.2 ‒ 2.8) and the length of palpomere five divided with its width (9.1 ‒ 11.5 versus 8.0 ‒ 9.3). The GspR of P. longistilus considerably overlaps that of P. floralis (0.25 ‒ 0.34 versus 0.28 ‒ 0.37). If overlapping, P. longistilus can usually be separated from P. floralis by the less marked gonostylus process divergence (5 ‒ 30 ° versus 30 ‒ 50 °) and mostly longer body (4.3 ‒ 5.7 versus 3.6 ‒ 4.4 mm). The pupa and larva, but not the adult female, have been described. Barcodes of adult males, adult females, pupa and larvae are available. Geographical distribution and ecology. P. longistilus has been recorded from sites with mean annual temperature difference as much as 23 ° C, ranging from about + 14 ° C in China to about ‒ 9 ° C in Baffin Island in Canada. In Europe the temperature span is less extensive, ranging from + 13 ° C in Spain to ‒ 2 ° C in northern Sweden. The geographical extension of P. longistilus is also wide with findings in Europe from latitude 42 ° N in the island Corsica of France to 70 ° N in the northern mainland of Norway. Records in North America are from 41 ° N in the United States to 68 ° N in Canada, and in China from 25 ° N to 49 ° N. P. longistilus has been reported to be present at altitudes from 5 m near the coast of the Baltic Sea in Sweden and up to 2 200 m in the Alps and Pyrenees of France. The species has also been found above 2 000 m in Tibet, China. Almost all the 82 quality assured findings of P. longistilus are from lakes, including some defined as polyhumic with water transparency less than one and a half m. It has also been reported from lake-like reservoirs, ponds, temporarily inundated meadows and slow flowing small rivers. The lakes are mostly oligotrophic to mesotrophic, but some are ultraoligotrophic or eutrophic. Studies have shown that the larvae of P. longistilus may decrease substantially in actual and relative numbers in reservoirs experiencing eutrophication and oxygen deficiency due to human activity. Larvae are known from 1 to 4 m water depth in the littoral, e. g. with plants such as Cladophora, and down to 5 m in the profundal. One study noted that the larvae of P. longistilus feed on other small Chironomidae. Adults can be important food items of bats and songbirds such as the pied flycatcher (Ficedula hypoleuca). Adults have been found from early May to late September. Countries with records of P. longistilus in Europe are Czechia, Finland, France, Germany, Italy, Lithuania, Netherlands, Norway, Poland, Russia, Slovakia, Spain and Sweden. There are also records from Canada, China, Greenland, Japan, Russia (Asia) and the United States including Alaska.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF7097D3CE4FAF19AEAF81C.taxon	description	Procladius lugens Kieffer, 1915 — Kieffer (1918 a), Lithuania, adult male, key, description. Procladius lugens Kieffer, 1915 — Goetghebuer (1927), France and Germany, adult male, adult female, key, description. Procladius lugens Kieffer, 1915 — Edwards (1929), England, adult female, description. Psilotanypus lugens (Kieffer, 1915) — Goetghebuer & Lenz (1936 a), England and Germany, adult male, adult female, key, description.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF7097D3CE4FAF19AEAF81C.taxon	description	Procladius lugens Kieffer, 1915 — Langton (1991), England, pupa, key, illustration. Procladius lugens Kieffer, 1915 — Langton & Visser (2003), England, pupa, key, illustration. Procladius lugens Kieffer, 1915 — Langton & Pinder (2007), adult male, key, illustration.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF7097D3CE4FAF19AEAF81C.taxon	materials_examined	Material examined (n = 23). ENGLAND, 1 adult male (BMNH), Timworth, 52.29 ° N 0.71 ° E, 27 m a. s. l., + 10 ° C m. a. t., 1948. — ESTONIA, 4 adult males (NHRS), Lake Ülemiste, 59.24 ° N 24.46 ° E, 36 m a. s. l., + 5 ° C m. a. t., 21. v. 1993, leg. Y. Brodin. — FINLAND, 4 adult males (as Psilotanypus sp., MZHF), Vantaa, Lake Kuusijärvi, 60.31 ° N 25.11 ° E, 44 m a. s. l., + 5 ° C m. a. t., 20. v. 1965, 16. v. 1975, leg. B. Lindeberg; 2 adult males (as Psilotanypus sp., HECH), Aidassaari, Lake Vanajavesi, 61.17 ° N 24.08 ° E, 79 m a. s. l., + 5 ° C m. a. t., 25. viii. 1978, leg. P. H. Kansanen; 2 adult males, Baltic Sea, Storskär Island, Lake Käringsund, 63.43 ° N 21.07 ° E, 1 m a. s. l., + 4 ° C m. a. t., 30. v ‒ 13. vi. 2017, leg. L. Paasivirta; 1 adult male (as P. flavifrons), Kiikala, Lake Tervakas, 60.49 ° N 23.70 ° E, 117 m a. s. l., + 5 ° C m. a. t., 9. v. 2023, leg. L. Paasivirta. — GERMANY, 1 adult male (ZSMG), Plön, 54.1 ° N 10.4 ° E, 18 m a. s. l., + 9 ° C m. a. t., circa 1947, leg. A. Thienemann. — NORWAY, 2 adult males (as P. cf. flavifrons, NTNU), Rohkosjavri, Stabbursdalen, 70.17 ° N 24.90 ° E, 12 m a. s. l., 0 ° C m. a. t., 15. vi. 2010, leg. T. Ekrem and E. Stur [Barcode CHRFI 500 - 11 and CHRFI 501 - 11]. — SWEDEN, 1 adult male (SLUU), Lake Mälaren, Görväln, 59.43 ° N 17.73 ° E, 2 m a. s. l., + 6 ° C m. a. t., vii. 1971, leg. T. Wiederholm; 2 adult males (SLUU), Lake Mälaren, Västeråsfjärden, 59.56 ° N 16.54 ° E, 3 m a. s. l., + 6 ° C m. a. t., vii. 1971, leg. T. Wiederholm; 1 adult male (NHRS), Lake Boren, Borenshult, 58.57 ° N 15.10 ° E, 74 m a. s. l., + 6 ° C m. a. t., 11. vi. 1980, leg. Y. Brodin; 1 adult male (NHRS), Eriksberg nature reserve, Stenåsa bog pond, 56.16 ° N 15.01 ° E, 10 m a. s. l., + 7 ° C m. a. t., 29. iv. 2015, leg. J. Wolgast [Barcoded]. — UNITED STATES, 1 adult male (as. P. macrotrichus, USNM), Lake Eagles Nest, 47.84 ° N 92.10 ° W, 447 m a. s. l., + 5 ° C m. a. t., 27. v. 1959, leg. W. V. Balduf. Diagnostic characters. Figs. 45, 122 ‒ 124, key couplet 26. P. lugens lacks or has a very short gonostylus process with a GspR that overlaps that of three other species of Procladius in Europe. Of these, P. flavifrons and P. simplicistilus are easily separated from P. lugens by several characters including gonostylus form, setae on katepisternum and number of macrotrichia on the wing membrane. The only similar species in Europe is P. imicola. The genitalia of P. lugens is very similar that of P. imicola, but the outer margin of the base section of the gonostylus is rounded in P. lugens, while usually distinctly angled in P. imicola. P. lugens is on average smaller than P. imicola (wing length 2.1 ‒ 2.8 versus 2.6 ‒ 3.4 mm, mid leg tibia length 0.8 ‒ 1.1 versus 1.1 ‒ 1.4 mm, body length 3.5 ‒ 4.6 versus 4.8 ‒ 5.5 mm) and the antenna AR mostly lower (1.7 ‒ 2.3 versus 2.2 ‒ 2.6). P. lugens has mostly been correctly identified in taxonomic literature regarding findings in Europe. The adult female has been briefly described, the pupal exuvia in detail, while the larva seems not to have been described. Barcodes of adult males, adult females and larvae are available. Geographical distribution and ecology. Sites with records of P. lugens have an extraordinary broad temperature spectrum, ranging from about + 14 ° C in Japan to about ‒ 14 ° C in Canada. The temperature span of P. lugens in Europe is more moderate, ranging from + 11 ° C in Ireland to 0 ° C in Norway. The latitude range of P. lugens in Europe stretches from France at latitude 46 ° N in the south to northern Norway at 70 ° N in the north. Worldwide the distribution is from 35 ° N in Japan to 73 ° N in Canada. The altitude records of the species are usually within 0 ‒ 200 m above sea level, but sometimes up to 540 m. Larvae of P. lugens inhabit oligotrophic to eutrophic, sometimes even hypereutrophic, lakes, lake-like reservoirs and ponds. The species is also known from shallow oligohaline water of the Baltic Sea and in samples of insects emerging from puddles in a temporarily inundated swamp. The bathymetric distribution of P. lugens larvae is mainly littoral on bottoms with vegetation such as Cladophora and reed (Phragmites). There are a few findings of P. lugens from the profundal, but not more than down to 5 m water depth, while its closest relative P. imicola is frequently found in the profundal and sometimes down to 30 m. Diatom remains have been found in the gut of P. lugens larvae, but otherwise nothing is known about the feeding of the larvae. Adults are known from late April to early August. Countries with records of P. lugens in Europe are Austria, England, Estonia, Finland, France, Germany, Ireland, Lithuania, Netherlands, Norway, Poland, Russia and Sweden. The species has also been recorded in Canada, China, Japan and the United States including Alaska, possibly also Algeria.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF5097F3CE4FF709C11FE81.taxon	materials_examined	Material examined (n = 21). CANADA, 1 adult male (Holotype of P. johnsoni, ANSP), Lake Muskoka, 45 ° N 79 ° W, 225 m a. s. l., + 5 ° C m. a. t., 5. vi. 1970, leg. M. Johnson. — NORWAY, 2 adult males (as P. barbatus, NTNU), Lake Målsjøen, 63.14 ° N 10.26 ° E, 166 m a. s. l., + 3 ° C m. a. t., 26. v ‒ 2. vi. 1972, 30. v. 1972, leg. K. Aagaard; 4 adult males (as Procladius sp., NHRS), Caerro, Lake Isejavri, 69.67 ° N 24.21 ° E, 390 m a. s. l., ‒ 3 ° C m. a. t., 26. vi ‒ 30. vii. 1980, leg. K. J. Loine; 1 adult male (as P. barbatus, NTNU), below Baktejavri, small pond, Lebesby, 70.44 ° N 26.80 ° E, 80 m a. s. l., ‒ 1 ° C m. a. t., 17. vi. 2010, leg. T. Ekrem and E. Stur [Barcode ARCHR 092 - 11]. — RUSSIA, 2 adult males (as P. barbatus, NHRS), Lake Mojarets, 69.48 ° N 30.60 ° E, 193 m a. s. l., ‒ 1 ° C m. a. t., 15. viii. 2006, leg. M. Pylvänäinen. — SWEDEN, 1 adult male (Holotype of Tanypus lugubris, ZMLU), Åreskutan, 63.4 ° N 13.0 ° E, 800 m a. s. l., ‒ 1 ° C m. a. t., 2. viii. 1840, leg. J. W. Zetterstedt; 1 adult male (as P. crassinervis, ZSMG), Lake Vuolip Njahkajavri, 68.34 ° N 18.78 ° E, 409 m a. s. l., ‒ 1 ° C m. a. t., 1936, leg. A. Thienemann; 2 adult males (as P. barbatus, NHRS), Lake Skären 57.10 ° N 14.52 ° E, 212 m a. s. l., + 6 ° C m. a. t., 7. v. 1946, 20. v. 1947, leg. L. Brundin; 1 adult male (Syntype of P. barbatus, NHRS), Lake Kallsjön, 63.59 ° N 13.07 ° E, 381 m a. s. l., + 2 ° C m. a. t., 14. vi. 1946, leg. L. Brundin; 1 adult male (Syntype of P. barbatus, NHRS), Lake Mesvattnet, 64.84 ° N 14.13 ° E, 498 m a. s. l., ‒ 1 ° C m. a. t., 25. vi. 1946, leg. A. Määr; 1 adult male (Syntype of P. barbatus, NHRS), Lake Leipikvattnet, 64.87 ° N 14.20 ° E, 0 ° C m. a. t., 448 m a. s. l., 6. vii. 1946, leg. A. Määr; 4 adult males (NHRS), Lake Stor-Björsjön, 63.61 ° N 12.23 ° E, 566 m a. s. l., 0 ° C m. a. t., 16 ‒ 23. vi. 2019, leg. S. Persson [2 Barcoded]. Diagnostic characters. Figs. 28, 71 ‒ 73, key couplet 11. P. lugubris has a medium long gonostylus process with a GspR that overlaps that of eight other species of Procladius in Europe. P. lugubris can easily be separated from seven of these species by several characters, particularly the low number of hind leg comb spines (6 ‒ 9 versus 10 ‒ 14), relatively short palpomere five (length / width 7.2 ‒ 8.5 versus 8.9 ‒ 12.2) and mostly higher antenna AR (2.5 ‒ 3.1 versus 1.6 ‒ 2.6). It is not problematic to distinguish P. lugubris from the remaining species P. frigidus by characters such as gonostylus process divergence (25 ‒ 45 ° versus 10 ‒ 25 °), front leg tarsi BR (5.5 ‒ 8 versus 3 ‒ 4.5), and the number of spines of the hind leg tibial comb (6 ‒ 9 versus 10 ‒ 14). P. lugubris has been named P. barbatus for about seventy-five years. Examination of the well-preserved holotype of P. lugubris and syntypes of P. barbatus shows that the latter is a synonym. The adult female and larva have been briefly described, while pupal exuvia in detail. Barcodes of adult males and adult females are available. Geographical distribution and ecology. P. lugubris is a cold-adapted species present from the temperate boreal to the polar zone in northern Europe and Canada, including sites with mean annual temperature from + 6 to ‒ 13 ° C. It has been found at an altitude ranging from 7 to 800 m above sea level. The southernmost findings in Europe are from Sweden at latitude 57 ° N, and in North America from Canada at latitude 45 ° N. The northernmost records are from sites at 70 ° N in the arctic northern Norway and northern Canada. Larvae of P. lugubris are known from the profundal zones of freshwater lakes at water depths ranging from 5 to 110 m which is among the deepest record for larvae of Procladius. Almost all records of P. lugubris are from ultraoligotrophic to oligotrophic conditions, yet it has also been found in some mesotrophic polyhumic lakes. The rather unusual round form of the pupal respiratory organ of P. lugubris, as also of P. ferrugineus, might be an adaptation to deep water emergence. Pupae of the phantom midge Chaoborus flavicans (Chaoboridae) have been shown to have rounded respiratory organs in lakes with deep water, whereas rather narrow respiratory organs in shallow ponds. Adults of P. lugubris have been collected from mid-May to late August along lakes or sometimes near small ponds in forests or tundra. Countries with records of P. lugubris in Europe are Finland, Norway, Sweden and Russia. The species is also present in Canada.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF409783CE4FE759FEDFF15.taxon	materials_examined	Material examined (n = 24). FINLAND, 1 adult male (MZHF), Lake Kuusijärvi, 60.31 ° N 25.1 ° E, 44 m a. s. l., + 5 ° C m. a. t., 18. vii. 1965, leg. P. Armitage; 2 adult males (ZMUO), Lake Naarjärvi, 60.31 ° N 23.29 ° E, 51 m a. s. l., + 5 ° C m. a. t., 5. vi. 2014, leg. L. Paasivirta [Barcode LEFIJ 3841 - 16 and LEFIJ 3842 - 16]; 2 adult males (ZMUO), west of Salo, Iso-Ruona, 60.36 ° N 23.64 ° E, 84 m a. s. l., + 5 ° C m. a. t., 2. vi. 2015, leg. L. Paasivirta [Barcode LEFIJ 3465 - 16 and LEFIJ 3466 - 16]. — RUSSIA, 3 adult males (NHRS), Lake Kolodenskoe, 59.01 ° N 37.06 ° E, 113 m a. s. l., + 4 ° C m. a. t., 27. v. 1989, leg. A. Rybakova; 5 adult males (NHRS), Lake Udebnoe, 59.07 ° N 37.43 ° E, 108 m a. s. l., + 4 ° C m. a. t., 29. v. 1989, leg. A. Rybakova; 3 adult males (NHRS), Lake Ulomskoe, 59.01 ° N 37.21 ° E, 111 m a. s. l., + 4 ° C m. a. t., 27. v. 1989, leg. A. Rybakova. — SWEDEN, 1 adult male (Syntype of P. nudipennis, NHRS), Lake Innaren, 56.98 ° N 14.97 ° E, 175 m a. s. l., + 6 ° C m. a. t., 27. vi ‒ 3. vii. 1945, leg. L. Brundin; 1 adult male (NHRS), Lake Viken, Sätra Bay, 58.64 ° N 14.25 ° E, 94 m a. s. l., + 6 ° C m. a. t., 15. v. 1987, leg. Y. Brodin; 3 adult males (NHRS), Lake Övre Forsasjön, 58.75 ° N 15.05 ° E, 137 m a. s. l., + 5 ° C m. a. t., 28. v. 1990, leg. Y. Brodin; 1 adult male (NHRS), Lake Salstern, 58.63 ° N 15.13 ° E, 107 m a. s. l., + 6 ° C m. a. t., 28. v. 1990, leg. Y. Brodin; 2 adult males (as P. rufovittatus, NHRS), west of Villingsberg, Lake Våtsjön, 59.28 ° N 14.64 ° E, 145 m a. s. l., + 5 ° C m. a. t., 20. v. 2008, leg. Y. Brodin [1 Barcode BSCHI 814 - 17]. Diagnostic characters. Figs. 43, 116 ‒ 118, key couplet 24. The gonostylus with its distinctly convex inner margin in combination with a distinct outer process readily separates P. nudipennis and P. bellus from all other European Procladius. The gonostylus of P. nudipennis is comparatively narrower (gonostylus GsmR 3.2 ‒ 3.9 versus 2.6 ‒ 3.3), less bulging and usually with fewer strong setae (4 ‒ 8 versus 6 ‒ 12) than that of P. bellus, while the gonostylus process is on average slightly longer (gonostylus GspR 0.13 ‒ 0.19 versus 0.10 ‒ 0.17). P. nudipennis can usually also be separated from P. bellus by the longer gonostylus in relation to the length of the gonocoxite (0.50 ‒ 0.59 versus 0.42 ‒ 0.50). P. nudipennis is on average somewhat smaller than P. bellus, exemplified by an often narrower gonocoxite base (178 ‒ 216 µm versus 206 ‒ 285 µm), wing length (1.8 ‒ 2.4 mm versus 2.0 ‒ 2.7 mm) and body length (3.1 ‒ 3.8 mm versus 3.5 ‒ 4.5). The adult female has been briefly described, but not the pupa and larva. Barcodes of adult males are available. Geographical distribution and ecology. P. nudipennis has a latitude distribution from 55 ° N in Russia to 68 ° N in northern Finland. It has been found at altitudes ranging from 15 to 860 m above sea level encompassing temperate to subarctic climate with + 8 to ‒ 1 ° C mean annual temperature. All findings of P. nudipennis are from lakes, mostly those with oligotrophic to mesotrophic condition, but also a few ultraoligotrophic and eutrophic ones. Most of the lakes are mesohumic to polyhumic. Adults have been found to emerge from the littoral zone at 0 ‒ 8 m from bottoms with sparse vegetation of Equisetum, Isoetes or Cladophora. Larvae are also found in the profundal from 1 m in polyhumic lakes to 15 m water depth in oligohumic ones. The larvae of P. nudipennis are omnivorous judging from their gut content of algae, detritus and animals such as crustaceans (Copepoda and Cladocera) and small Chironomidae. Adults have been found flying from the middle of May to the middle of September. Countries with records of P. nudipennis in Europe are Estonia, Finland, Norway, Russia and Sweden. It might be present in other European countries under the name of P. rufovittatus.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF3097A3CE4FEC19DDEFBB9.taxon	description	Procladius sp. — Muragina-Koreneva (1957), Russia, adult male, description, key, illustration.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF3097A3CE4FEC19DDEFBB9.taxon	materials_examined	Material examined (n = 57). AUSTRIA, 1 adult male (as P. choreus, ZSMG), Lake Lünzer Obersee, 47.80 ° N 15.08 ° E, 1 113 m a. s. l., + 2 ° C m. a. t., 1940, leg. F. Krüger; 1 adult male (as Procladius III, ZSMG), Lunz, stream from pond, 47.85 ° N 15.05 ° E, 610 m a. s. l., + 5 ° C m. a. t., 7. v. 1972, leg. N. Caspers. — CZECHIA, 2 adult males (NHRS), Cheb district, Lake Vodni Nadrz Skalka, 50.08 ° N 12.32 ° E, 439 m a. s. l., + 7 ° C m. a. t., 17. vi. 1991, leg. Y. Brodin. — ESTONIA, 1 adult male (NHRS), Lake Ülemiste, 59.24 ° N 24.46 ° E, 36 m a. s. l., + 6 ° C m. a. t., 21. v. 1993, leg. Y. Brodin. — FINLAND, 1 adult male (as P. cfr. nigriventris, WIFE), Lake Kuusijärvi, 60.31 ° N 25.12 ° E, 44 m a. s. l., + 5 ° C m. a. t., 15. vi. 1965, leg. P. Armitage; 3 adult males (as Procladius sp., HECH), Lake Karvastenlahti, Hiittiö, 61.16 ° N 24.03 ° E, 79 m a. s. l., + 5 ° C m. a. t., 25. v. 1978, leg. P. H. Kansanen; 2 adult males (as P.? cinereus, HECH), Aidassaari, Lake Vanajavesi, 61.17 ° N 24.08 ° E, 79 m a. s. l., + 5 ° C m. a. t., 22 ‒ 25. viii. 1978, leg. P. H. Kansanen; 1 adult male (as Procladius sp.), Baltic Sea, Lake Käringsund, Storskär Island, 63.43 ° N 21.07 ° E, 0 m a. s. l., + 4 ° C m. a. t., 30. v ‒ 13. vi. 2017, leg. L. Paasivirta. — FRANCE, 2 adult males (as Procladius sp. I, LHST), Lake de L’ite, 42.85 ° N 0.16 ° E, 2 281 m a. s. l., + 2 ° C m. a. t., 16. vii. 1965, leg. H. Lavillle; 1 adult male (as Procladius sp. I, LHST), Lake de Nere Inferior, 42.87 ° N 0.18 ° E, 2 209 m a. s. l., + 2 ° C m. a. t., 7. vii. 1966, leg. H. Laville. — GERMANY, 1 adult male (as P. bathocryptus, ZSMG), Lake Schluensee, 54.19 ° N 10.46 ° E, 22 m a. s. l., + 9 ° C m. a. t., 1953, leg. I. Müller-Liebenau; 1 adult male (as P. choreus, ZSMG), northeast of Sonneberg, Lake Königsee, 50.38 ° N 11.18 ° E, 627 m a. s. l., + 7 ° C m. a. t., 25 ‒ 27. iv. 1957, leg. W. Engelhardt; 1 adult male (as Procladius sp., ZSMG), Braunkohlerestgewerke, OS Boden, 51.0 ° N 6.5 ° E, 21 m a. s. l., + 11 ° C m. a. t., 13. xii. 1966, leg. Herbst; 1 adult male (as P. choreus, ZSMG), Lake Königsee, 50.38 ° N 11.19 ° E, 628 m a. s. l., + 7 ° C m. a. t., 25. v. 1986, leg. R. Gerstmaier; 1 adult male (as P. crassinervis, ZSMG), Lake Chiemsee, Herrenchiemsee, 47.87 ° N 12.40 ° E, 518 m a. s. l., + 9 ° C m. a. t., 13. vi. 1988, leg. C. Orendt; 2 adult males (as P. choreus, ZFMK), Meyenburg, Fleet, wet meadow, 52.51 ° N 9.38 ° E, 37 m a. s. l., + 10 ° C m. a. t., 21. vi. 2017, leg. E. Resendiz and H. - G. Rudzinski; 1 adult male (as P. choreus, ZFMK), Brettenbach stream, Brücke Vordere Zaismatt, 48.13 ° N 7.90 ° E, 242 m a. s. l., + 10 ° C m. a. t., 31. iii. 2019 [Barcode GBOL- 2625260]. — IRELAND, 2 adult males (as Procladius sp. 1, UCDZ), Lake Gougane Barra, 51.84 ° N 9.32 ° W, 165 m a. s. l., + 8 ° C m. a. t., 26. vii. 1966, leg. D. A. Murray; 2 adult males (as P. pectinatus, UCDZ), Lake Leane, O`Sullivans Cascade and pool, 52.04 ° N 9.58 ° W, 30 m a. s. l., + 10 ° C m. a. t., 17. iv. 1973, leg. L. Bucheen; 1 adult male (as Procladius sp. A, UCDZ), Lake Doolough, 52.79 ° N 9.30 ° W, 83 m a. s. l., + 8 ° C m. a. t., 14. ix. 1973, leg. D. A. Murray; 1 adult male (as Procladius sp., UCDZ), Lake Carra, 53.66 ° N 9.26 ° W, 21 m a. s. l., + 10 ° C m. a. t., 2. iv. 1974, leg. D. A. Murray; 4 adult males (as Procladius sp., UCDZ), Lake Carra, 53.68 ° N 9.25 ° W, 19 m a. s. l., + 10 ° C m. a. t., 10. v. 1974, leg. D. A. Murray. — ITALY, 4 adult males (NHRS), Lake Lago di Centro Cadore, Calazo, 46.44 ° N 12.39 ° E, 683 m a. s. l., + 8 ° C m. a. t., 29. vi. 1991, leg. Y. Brodin; 2 adult males (NHRS), Lake Lago di Valdaora, Monguelfo, 46.76 ° N 12.05 ° E, 1 087 m a. s. l., + 5 ° C m. a. t., 29. vi. 1991, leg. Y. Brodin; 1 adult male (as P. choreus, MTSN), Tre laghi Isole, 45 ° N 8 ° E, 238 m a. s. l., + 11 ° C m. a. t., 2. vii. 1996, leg. L. Marziali. — NORWAY, 1 adult male (as P. cf. fuscus, NTNU), Trondheim, Lake Gjeddtjørna, 63.38 ° N 10.61 ° E, 158 m a. s. l., + 4 ° C m. a. t., 6. vi. 2010, leg. J. K. Skei [Barcode CHMNO 070 - 14]; 1 adult male (as P. cf. fuscus, NTNU), Trondheim, Lake Gjeddevatnet, 63.38 ° N 10.61 ° E, 157 m a. s. l., + 4 ° C m. a. t., 20. vi. 2010, leg. J. K. Skei [Barcode MIDGE 985 - 13]; 2 adult males (as P. cf. fuscus, NTNU), Melhus, Gammelelva nature reserve, 63.21 ° N 10.31 ° E, 28 m a. s. l., + 4 ° C m. a. t., 22. v ‒ 5. vi. 2014, leg. E. Stur [Barcode CHMNO 205 - 15 and CHMNO 354 - 15]. — POLAND, 1 adult male (as Procladius sp., LUIZ), Spala, River Pilica, 51.54 ° N 20.13 ° E, 151 m a. s. l., + 9 ° C m. a. t., 5. vi ‒ 9. vi. 2007, leg. M. Plóciennek; 2 adult males (as Procladius sp., LUIZ), Zabierzow, 51.54 ° N 19.79 ° E, 270 m a. s. l., + 9 ° C m. a. t., 17. v. 2010, leg. M. Plóciennek. — RUSSIA, 1 adult male (as P. culiciformis, NHRS), Kamchatka, Petropavlovsk area, 53 ° N 158 ° E, 150 m a. s. l., ‒ 1 ° C m. a. t., vii. 1921, leg. R. Malaise; 2 adult males (as Procladius sp, IBIB), Rybinsk Reservoir, 58.1 ° N 38.6 ° E, 97 m a. s. l., + 4 ° C m. a. t., vii. 1972, leg. A. I. Shilova. — SWEDEN, 1 adult male (as Procladius sp., NHRS), Lake Vitalampi, 59.55 ° N 15.13 ° E, 322 m a. s. l., + 5 ° C m. a. t., 7. v. 1974, leg. P. Mossberg; 1 adult male (NHRS), Lake Vristulven, Århult, 58.56 ° N 13.72 ° E, 113 m a. s. l., + 6 ° C m. a. t., 22 ‒ 24. iv. 1987, leg. Y. Brodin; 1 adult male (NHRS), Lake Vänern, Sjötorp, 58.84 ° N 13.97 ° E, 44 m a. s. l., + 6 ° C m. a. t., 16. vi. 1987, leg. Y. Brodin; 2 adult males (NHRS), Baltic Sea, Askö island, Askötorp, 58.80 ° N 17.67 ° E, 6 m a. s. l., + 6 ° C m. a. t., 31. v ‒ 16. vi. 2011, leg. B. E. Bengtsson [Barcode BSCHI 536 - 17 and BSCHI 551 - 17]. — SWITZERLAND, 1 adult male (as P. cinereus, ZSMG), Lake Bodensee, Egnach, 47.54 ° N 9.38 ° E, 396 m a. s. l., + 9 ° C m. a. t., 1938, leg. J. Geissbühler. — WALES, 1 adult male (as P. choreus), Trawscoed, 52.33 ° N 3.95 ° W, 70 m a. s. l., + 8 ° C m. a. t., iv. 2020, leg. J. Graham. Diagnostic characters. Figs. 5, 8, 16, 36, 95 ‒ 97, key couplet 18. P. pruinosus has a medium long gonostylus process with a GspR that overlaps that of eight other species of Procladius in Europe. Of these, P. lugubris and P. ferrugineus are distinctly separated from P. pruinosus by more than one other morphological character in the key and the helpdesk. The GspR of P. pruinosus slightly overlaps that of P. islandicus (0.23 ‒ 0.30 versus 0.18 ‒ 0.24). If overlapping, P. pruinosus can be separated from P. islandicus by the on average lower front leg tarsi BR (2 ‒ 4.5 versus 3 ‒ 6.5) and the longer and narrower gonostylus process (length / width 1.2 ‒ 1.7 versus 0.9 ‒ 1.2). The GspR of P. pruinosus to some degree overlaps that of P. culiciformis (0.23 ‒ 0.30 versus 0.18 ‒ 0.25). When overlapping, P. pruinosus can be distinguished from P. culiciformis by the narrower gonostylus (GsmR 5.2 ‒ 6.3 versus 4.2 ‒ 5.2) and the on average darker mid-section of the front leg tibia (light brown to dark brown versus whitish to light brown). The GspR of P. pruinosus almost completely overlaps that of P. tenebricosus (0.23 ‒ 0.30 versus 0.23 ‒ 0.29). P. pruinosus can be separated from P. tenebricosus by the less dark marked wing (anal cell patch absent to faint versus distinct to very distinct) and almost always larger size (wing length 2.6 ‒ 3.5 mm versus 1.8 ‒ 2.6 mm, mid leg tibia length 1.0 ‒ 1.4 mm versus 0.7 ‒ 1.1 mm, body length 4.3 ‒ 5.9 mm versus 3.2 ‒ 4.3 mm). The GspR of P. pruinosus strongly overlaps that of P. frigidus (0.23 ‒ 0.30 versus 0.25 ‒ 0.33). Despite that, there are generally no problems to separate the species by consulting other morphological characters. P. pruinosus has fewer median anepisternal setae compared to that of P. frigidus (0 ‒ 5 versus 5 ‒ 26), the antenna AR is usually lower (1.7 ‒ 2.3 versus 2.2 ‒ 2.8) and the length of palpomere five divided with its width is lower (9.3 ‒ 12.2 versus 8.0 ‒ 9.3). The GspR of P. pruinosus considerably overlaps that of P. longistilus (0.23 ‒ 0.30 versus 0.25 ‒ 0.34). P. pruinosus can mostly be separated from P. longistilus by the ratio between the length of the outer phallapodeme section divided by the length of the sternapodeme (0.27 ‒ 0.36 versus 0.34 ‒ 0.46). The length of the outer phallapodeme is however frequently difficult to measure accurately and requires good slide preparations. In addition, the on average broader gonostylus and gonostylus process of P. pruinosus than that of P. logngistilus (GsmR 5.2 ‒ 6.3 versus 5.5 ‒ 6.9, gonostylus process length / width 1.2 ‒ 1.7 versus 1.3 ‒ 1.9) can be useful for species separation. The GspR of P. pruinosus partly overlaps that of P. floralis (0.23 ‒ 0.30 versus 0.28 ‒ 0.37). If overlapping, P. pruinosus can be separated from P. floralis by the less marked gonostylus process divergence (10 ‒ 25 ° versus 30 ‒ 50 °) and the mostly longer body (4.3 ‒ 5.9 versus 3.6 ‒ 4.4 mm). P. pruinosus has been described or mentioned under numerous species-names in the literature, frequently as P. choreus, but also P. bathocryptus, P. cinereus, P. culiciformis, P. fuscus, P. niger, P. nigriventris, P. pectinatus and P. zernyi. The adult female and pupal exuvia have been briefly described, but not the larva. The distinct pruinosity of the thorax of male P. pruinosus, which gave the species its name, can be observed on dry or alive specimens. Barcodes of adult males, adult female, pupa and larva are available. Geographical distribution and ecology. In Europe, P. pruinosus is known from latitude 43 ° N in southern France to 68 ° N in northern Norway. In North America, the known latitude range is from 43 ° N to 62 ° N in the United States and Canada. Altitudes records range from 5 m and up to 2 280 m above sea level in the Pyrenean Mountains of France. The 76 sites with quality assured findings of P. pruinosus have a mean annual temperature span of + 14 to ‒ 2 ° C. P. pruinosus can be regarded as a common species in European lakes and lake-like reservoirs classified as oligotropic to eutrophic. It is less frequently found in hypereutrophic reservoirs and not found in ultraoligotrophic lakes or reservoirs. Larvae of P. pruinosus have been recorded from 0 ‒ 6 m water depth in the littoral of lakes over bottoms of sand, stone or mud with vegetation of quillworts (Isoetes), common reed (Phragmites), water lilies (Nymphaea and Nuphar) and green macroalgae of the genus Cladophora. Larvae are also known from the profundal zone at 1 ‒ 16 m water depth. Most findings of P. pruinosus are from lakes, but it has also been recorded from reservoirs, ponds on meadows, pools on peat bogs with Carex, constructed pools in brown coal excavations, slowly running rivers and rivulets. The larvae can thrive and develop into the pupal stage in very small water bodies, such as in a rowing boat filled with lake water full of green algae. The gut content of P. pruinosus larvae indicates that they consume diatoms, green algae, and smaller animals such as crustaceans (Copepoda and Cladocera) and Chironomidae larvae. Adult P. pruinosus can possibly be found throughout the entire year in Europe, at least from late March to late January in southern Europe. Countries or autonomous regions with records of P. pruinosus in Europe are Austria, Belgium, Czechia, England, Estonia, Finland, France, Germany, Ireland, Italy, Norway, Poland, Russia, Slovakia, Sweden, Switzerland and Wales. It is also present in the Asian part of Russia, Canada and the United States including Alaska, possibly also in Iran.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFF009663CE4FBFD9EE9FDD9.taxon	materials_examined	Material examined (n = 34). AUSTRIA, 1 adult male (ZSMG), Lake Gebhartsteich, 48.80 ° N 15.14 ° E, 545 m a. s. l., + 6 ° C m. a. t., 12. vi. 1978, leg. H. Malicky. — CANADA, 2 adult males (as P. denticulatus, CNCC), Lake Muskox, 64.63 ° N 108.25 ° W, 343 m a. s. l., ‒ 9 ° C m. a. t., 17 ‒ 24. vii. 1953, leg. J. G. Chillcott. — ENGLAND, 1 adult male (UUZM), South Hanningfield, Hanningfield Reservoir, 51.65 ° N 0.51 ° E, 53 m a. s. l., + 11 ° C m. a. t., 23. vii. 1982, leg. T. Samman and Y. Brodin. — FINLAND, 1 adult male (WIFE), Lake Kuusijärvi, 60.31 ° N 25.12 ° E, 44 m a. s. l., + 5 ° C m. a. t., 15. vi. 1965, leg. P. Armitage; 4 adult males (HECH), Lake Vanajavesi, 61.17 ° N 25.08 ° E, 79 m a. s. l., + 5 ° C m. a. t., 21. viii. 1978, leg. P. H. Kansanen; 2 adult males (ZMUO), Lake Pyhäjärvi, Säkylä, 61.00 ° N 22.28 ° E, 45 m a. s. l., + 5 ° C m. a. t., 4. vii. 2015, leg. L. Paasivirta [Barcode LEFIJ 3482 - 16 and LEFIJ 3483 - 16]. — FRANCE, 2 adult males (NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 849 m a. s. l., + 7 ° C m. a. t., iv. 2017, leg. B. Tissot. — GERMANY, 1 adult male (as P. pectinatus, ZSMG), south of Wismar, Lake Tanysee, 53.67 ° N 11.45 ° E, 35 m a. s. l., + 9 ° C m. a. t., 1936, leg. A. Thienemann; 1 adult male (as P.? parvulus, ZSMG), Lake Bodensee, 47.67 ° N 9.32 ° E, 394 m a. s. l., + 9 ° C m. a. t., 12 ‒ 17. vii. 1961, leg. F. Reiss; 1 adult male (ZSMG), Lake Bodensee, 47.53 ° N 9.40 ° E, 395 m a. s. l., + 9 ° C m. a. t., 20. vii. 1961, leg. F. Reiss; 1 adult male (ZSMG), Lake Chiemsee, Herrenchiemsee, 47.87 ° N 12.40 ° E, 518 m a. s. l., + 9 ° C m. a. t., 2. vii. 1988, leg. C. Orendt. — IRELAND, 4 adult males (UCDZ), Lake Leane, Castlelough Bay, 52.03 ° N 9.50 ° E, 17 m a. s. l., + 9 ° C m. a. t., 3. ix. 1973, leg. D. J. Douglas and D. A. Murray. — NORWAY, 2 adult males (NTNU), Lebesby, Lake Eastorjavri, outlet to stream, 70.44 ° N 27.35 ° E, 258 m a. s. l., ‒ 2 ° C m. a. t., 28. vii. 2010, leg. T. Ekrem [Barcode CHRFI 245 - 10 and CHRFI 246 - 10]. — RUSSIA, 3 adult males (TUSF), Lake Ladoga, Sorvala, 61.74 ° N 30.77 ° E, 4 m a. s. l., + 5 ° C m. a. t., 4. viii. 1991, leg. G. Söderman. — SWEDEN, 1 adult male (Syntype of P. signatus, as Tanypus signatus, ZMLU), Åreskutan, 63.4 ° N 13.0 ° E, 900 m a. s. l., ‒ 1 ° C m. a. t., 31. vii ‒ 4. viii. 1840, leg. J. W. Zetterstedt; 1 adult male (as P. lundstroemi, ZSMG), Lake Torneträsk, 68.4 ° N 18.8 ° E, 343 m a. s. l., ‒ 1 ° C m. a. t., 19. vi. 1937, leg. A. Thienemann; 2 adult males (NHRS), Lake Färnebofjärden, Östa, 60.17 ° N 16.80 ° E, 60 m a. s. l., + 5 ° C m. a. t., 1. vi. 2007, 7. ix. 2007, leg. Y. Brodin; 1 adult male (NHRS), Lake Färnebofjärden, Edsviken Bay, 60.29 ° N 16.81 ° E, 59 m a. s. l., + 5 ° C m. a. t., 6 ‒ 13. vi. 2007, leg. Y. Brodin; 1 adult male (NHRS), Lake Vättern, Motala, 58.53 ° N 14.98 ° E, 88 m a. s. l., + 6 ° C m. a. t., 30. vii. 2012, leg. Y. Brodin [Barcode BSCHI 770 - 17]; 1 adult male (NHRS), Lake Mälaren, Vårby beach, 59.26 ° N 17.88 ° E, 3 m a. s. l., + 6 ° C m. a. t., 1. viii. 2012, leg. Y. Brodin [Barcode BSCHI 727 - 17]. — SWITZERLAND, 1 adult male (as P. cinereus, ZSMG), Lake Bodensee, Egnach, 47.54 ° N 9.38 ° E, 396 m a. s. l., + 9 ° C m. a. t., 1938, leg. J. Geissbühler. Diagnostic characters. Figs. 17, 24, 59 ‒ 61, key couplet 6. P. signatus is easily distinguished from other species of Procladius in Europe by the form of the medioapodeme, especially its inner apical part with many dents. A similar strongly dented medioapodeme is present in P. wilhmi Roback, 1971 and P. sublettei Roback, 1971 in North America. P. wilhmi has a much shorter gonostylus process. P. sublettei has a similar gonostylus but lacks the bulging inferior volsella present in P. signatus. It is also smaller and has an on the whole more southern geographical distribution compared to that of P. signatus. P. signatus has a very long curved gonostylus process usually with a distinct net-pattern. Such a net-pattern is only occasionally seen on the gonostylus process of other species of Procladius in Europe. P. signatus is generally correctly identified in literature regarding occurrences in Europe. It has been known as P. denticulatus in North America. Holotype and paratypes are destroyed, but other studied specimens from Canada and the United States show that P. denticulatus is a synonym of P. signatus. The adult female, pupal exuvia and larva have been described. Barcodes of adult males, adult females, a pupa and larvae are available. Geographical distribution and ecology. The latitude range of P. signatus in Europe stretches from France and Hungary at 47 ° N to the most northern districts of the mainland in Norway and Finland at 70 ° N. In North America, the species is found near the west coast of the Pacific Ocean of the United States at 41 ° N to near the Arctic Ocean in northern Canada at 68 ° N. This means a mean annual temperature range from + 11 ° C in England, Hungary and the United States to ‒ 13 ° C in northern Canada, thus as much as 24 degrees. Noteworthy is that barcodes of P. signatus from the sites in England and those from the sites in northern Canada only differ by 0.2 %. P. signatus has been found at altitudes in Europe ranging from 3 m near the sea in Sweden to about 850 m above sea level in France, and up to 1 130 m in Canada. Most of the 80 quality assured findings of P. signatus are from lakes, but it does seem to be frequent also in ponds such as those in peat bogs. Findings from reservoirs, puddles or slowly floating water sections of streams or rivers are less frequent. Larvae of P. signatus occur mostly in oligotrophic to eutrophic water bodies, sometimes also in ultraoligotrophic ones down to 28 m water depth. Larval habitats can have low to high content of dissolved organic matter such as in polyhumic ponds with pH reaching down to 5.0. This indicates that the larvae have a good ability to withstand low oxygen concentrations. P. signatus larvae are mostly found on bottoms with mud or sand without or with vegetation of various kinds, including Chara, Cladophora, Elodea and Nymphaea. Animal food items of P. signatus larvae, such as small Chironomidae larvae, Cladocera, Copepoda and Gastrotrichia, seem to be mainly consumed during periods of rapid growth in spring or summer, whereas mostly benthic algae and detritus are found in the larval guts during late autumn, winter and early springs months from November to April. Larvae of P. signatus serve as food for water mites and other aquatic organisms. Adults of P. signatus appear from early May to late October. Studies from Norway show that adult males can be parasitized by nematodes which cause deformation of the male’s genitalia and other body parts. Countries or autonomous regions with records of P. signatus in Europe are Austria, Belgium, Czechia, Denmark, England, Estonia, Finland, France, Germany, Hungary, Ireland, Lithuania, Netherlands, Northern Ireland, Norway, Poland, Russia, Scotland, Slovakia, Sweden, Switzerland and Wales. The species has also been found in Canada, Russian Asia and in the United States.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFED09673CE4FD8D9AD9FC41.taxon	materials_examined	Material examined (n = 18). BELGIUM, 1 adult male (as Procladius sp., ZSMG), Épioux Reservoir, 49.8 ° N 5.3 ° E, 383 m a. s. l., + 9 ° C m. a. t., v. 1964. — CANADA, 2 adult males (as P. ruris var. grandis, CNCC), Lake Altin, Altin, 59.35 ° N 133.42 ° W, 670 m a. s. l., 0 ° C m. a. t., 9. vi. 1955, leg. Huckel. — ENGLAND, 1 adult male (Holotype of P. simplicistilus, BMNH), Hawkshead, Three Dubs Tarn, 54.37 ° N 2.96 ° W, 218 m a. s. l., + 7 ° C m. a. t., 7 ‒ 15. v. 1947, leg. T. T. Macan; 1 adult male (Paratype of P. simplicistilus, BMNH), Hawkshead, Three Dubs Tarn, 54.37 ° N 2.96 ° W, 218 m a. s. l., + 7 ° C m. a. t., 7 ‒ 15. v. 1947, leg. T. T. Macan. — FINLAND, 2 adult males (as Procladius sp., LMMR), Enontekiö, Lake Toskaljärvi, 69.19 ° N 21.46 ° E, 704 m a. s. l., ‒ 5 ° C m. a. t., 4 ‒ 15. vii. 2010, leg. L. Kakko; 2 adult males (ZMUO), Lake Pyhäjärvi, Säkylä, 61.00 ° N 22.28 ° E, 45 m a. s. l., + 5 ° C m. a. t., 4. vi. 2015, leg. L. Paasivirta [Barcode CHIFI 292 - 16 and CHIFI 293 - 16]. — FRANCE, 3 adult males (as Procladius sp., 1, LHST), Lake Long, 42.82 ° N 0.12 ° E, 2 094 m a. s. l., + 3 ° C m. a. t., 13. viii. 1967, leg. H. Laville. — IRELAND, 2 adult males (UCDZ), Lake Sillan, 54.01 ° N 6.92 ° W, 89 m a. s. l., + 9 ° C m. a. t., 14. iv. 1978, leg. D. A. Murray; 1 adult male (NMID), Lake Sandy, 54.35 ° N 8.17 ° W, 333 m a. s. l., + 8 ° C m. a. t., 28. iv. 2005, leg. D. A. Murray and J. - R. Baars. — NORWAY, 1 adult male (as Procladius sp., NHRS), Caerro, Lake Isejavri, 69.67 ° N 24.21 ° E, 390 m a. s. l., ‒ 3 ° C m. a. t., 26. vi ‒ 30 vii. 1980, leg. K. J. Loine. — SWEDEN, 1 adult male (as P. sagittalis, NHRS), Njulja pond, 68.37 ° N 18.70 ° E, 995 m a. s. l., ‒ 5 ° C m. a. t., 15. vii. 1936, leg. A. Thienemann. — SWITZERLAND, 1 adult male (MCSN), Lake Leman, 46.37 ° N 6.27 ° E, 372 m a. s. l., + 10 ° C m. a. t., 7. v. 2009, leg. B. Lods-Crozet. Diagnostic characters. Figs. 27, 68 ‒ 70, key couplet 9. Male P. simplicistilus and P. appropinquatus have an only slightly indicated gonostylus process which separates them from all other European Procladius species with wing macrotrichia. The gonostylus process of P. simplicistilus is on average somewhat shorter than that of P. appropinquatus (GspR 0.02 ‒ 0.06 versus 0.04 ‒ 0.10). P. simplicistilus is larger than P. appropinquatus exemplified by wing length (3.7 ‒ 4.6 mm versus 2.5 ‒ 3.7 mm), body length (6.5 ‒ 8.1 mm versus 4.3 ‒ 6.0 mm) and gonocoxite width (377 ‒ 451 µm versus 282 ‒ 350 µm). The antenna AR-ratio of P. simplicistilus is mostly higher than that of P. appropinquatus (AR 2.5 ‒ 3.2 versus 1.8 ‒ 2.6). Hairiness is often also useful to distinguish P. simplicistilus from P. appropinquatus (scutellum 74 ‒ 117 setae versus 40 ‒ 73 setae, wing vein Cu stem 0 ‒ 4 setae versus 4 ‒ 36 setae). P. simplicistilus is the largest Procladius known worldwide. The pupal exuvia has been described, but probably not the female and larva. Barcodes of adult males, adult females, and larvae are available. Geographical distribution and ecology. P. simplicistilus is known from southern France at latitude 43 ° N to northern Norway at latitude 70 ° N. The species seems to be rather common in Ireland and Northern Ireland, but otherwise less common or rare. Sites with records cover a mean annual temperature interval from + 10 to ‒ 5 ° C, with the coldest conditions in the northern mountains of Finland and Sweden, and globally ‒ 7 ° C in Canada. The species has an altitude range from 5 m to 2 090 m above sea level. P. simplicistilus has only been reported from lakes and smaller water bodies with standing water such as ponds and gravel pits. Records of up to 1 400 individuals per m 2 are present from 0 to 4 m water depth in the littoral with plants such as Potamogeton, Myriophyllum or Elodea. There are a few records of P. simplicistilus larvae in the profundal, where they have been reported from depths down to 78 m. Ultraoligotrophic to eutrophic conditions exist in the lakes with findings of P. simplicistilus. The lakes are classified as oligohumic to polyhumic with pH from 5.5 to 8. Food items of the larvae are not known. P. simplicistilus adults fly from mid-April to mid-August. Countries or autonomous regions with records of P. simplicistilus in Europe are Belgium, Denmark, England, Finland, France, Ireland, Lithuania, Netherlands, Northern Ireland, Norway, Poland, Russia, Scotland, Slovakia, Sweden and Wales. The species has also been recorded from Canada.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFEC09603CE4FC359C44FD31.taxon	materials_examined	Material examined (n = 24). AUSTRIA, 1 adult male (as P. pectinatus, ZSMG) Lünz, 47.85 ° N 15.04 ° E, 604 m a. s. l., + 5 ° C m. a. t., 1940 ‒ 1942, leg. F. Krüger and A. Thienemann; 1 adult male (as Procladius sp., ZSMG), Lake Weissensee, Neusach, 46.71 ° N 13.31 ° E, 929 m a. s. l., + 5 ° C m. a. t., 7. v. 1982, leg. B. Janacek. — FINLAND, 5 adult males (as P.? barbatus, MZHF), Lake Puruvesi, Koivusaari, 61.48 ° N 29.46 ° E, 76 m a. s. l., + 3 ° C m. a. t., 19. v. 1959, leg. B. Lindeberg. — FRANCE, 3 adult males (NHRS), Lake Remoray, 46.77 ° N 6.26 ° E, 849 m a. s. l., + 7 ° C m. a. t., iv. 2019, leg. B. Tissot [Barcoded]; 2 adult males, Remoray, svamp, 46.78 ° N 6.27 ° E, 848 m a. s. l., + 7 ° C m. a. t., iv. 2019, leg. B. Tissot [1 Barcoded]. — ITALY, 1 adult male (as Procladius sp., MTSN), Lake Scuro del Mandrone, 46.21 ° N 10.57 ° E, 2 660 m a. s. l., + 1 ° C m. a. t., 20. viii. 1996, leg. L. Marziali. — NORWAY, 4 adult males (as Procladius sp., NHRS), Lake Fantesteinsvatnet, Sognefjellshytta, 61.56 ° N 8.00 ° E, 1 408 m a. s. l., ‒ 1 ° C m. a. t., 12. viii. 1952, leg. L. Brundin; 2 adult males (as P. cf. jeris, ZSMG), Finse, pond at Hardanger Jökula, 60.6 ° N 7.4 ° E, 1 450 m a. s. l., ‒ 3 ° C m. a. t., 2. viii. 1985, leg. E. J. Fittkau and F. Reiss. — SLOVAKIA, 1 adult male (Syntype of P. tatrensis), Lake Zelené pleso, 49.21 ° N 20.22 ° E, 2 012 m a. s. l., ‒ 1 ° C m. a. t., 18. viii. 1938, leg. S. Hrabe; 3 adult males (FNSB), Nizne Terianske, 49.17 ° N 20.01 ° E, 1 940 m a. s. l., 0 ° C m. a. t., 4. viii. 2008, leg. P. Bitušík. — SWITZERLAND, 1 adult male (as P. pectinatus, ZSMG), Lake Bodensee, Egnach, 47.54 ° N 9.38 ° E, 396 m a. s. l., + 9 ° C m. a. t., 1938, leg. J. Geissbühler. Diagnostic characters. Figs. 11, 25, 62 ‒ 64, key couplet 7. P. tatrensis has a very long gonostylus process with a GspR that overlaps that of five other species of Procladius in Europe. All of them are efficiently separated from P. tatrensis by more than one other morphological character in the key and the helpdesk, particularly the form of the medioapodeme, wing length, number of median anepisternal setae and antenna AR. Male P. jeris Roback, 1971, found in Alaska in the United States, and P. tatrensis have morphologically similar genitalia. In P. tatrensis the caudolateral ends of the ninth tergite are much less produced than in P. jeris, and body size is considerably larger (wing length 3.5 ‒ 4.5 mm versus 2.3 ‒ 2.7 mm). The adult female and larva of P. tatrensis have been briefly described, and the pupal exuvia in detail. Geographical distribution and ecology. The geographical range of P. tatrensis extends from northern Italy at 46 ° N to 68 ° N in northern Sweden, encompassing a temperature interval from + 9 to ‒ 3 ° C expressed as mean annual temperature. P. tatrensis is mainly a mountain species present at altitudes above 350 m with or without natural forests. It holds the altitude record for Procladius species in Europe, with findings in the Alps at altitude 2 660 m above sea level in northern Italy and 2 670 m in eastern France. There are records of P. tatrensis higher than 2 000 m also from Slovakia, Austria and Switzerland. Only a record from Finland at 75 m is beyond 300 m above sea level. P. tatrensis seems to be mainly confined to lakes or ponds with ultraoligotrophic to mesotrophic conditions. There are a few findings from lakes with eutrophic conditions. Larvae have been reported from 3 to 80 m water depth. It is not known what the larvae consume. P. tatrensis larvae can be important food for fish such as Salvelinus in oligotrophic high-altitude lakes. Adults of P. tatrensis are known from mid-May to late September. Countries with records of P. tatrensis in Europe are Austria, Finland, France, Germany, Italy, Norway, Poland, Slovakia, Sweden and Switzerland.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFEB09613CE4F98E9A1EF8E9.taxon	description	P. cf. cinereus — Ratnasingham et al. (2024), Norway, adult male, photo. P. sp. — Ratnasingham et al. (2024), Germany, adult males.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
5E47CA08FFEB09613CE4F98E9A1EF8E9.taxon	materials_examined	Material examined (n = 19). FRANCE, 1 adult male (as P. choreus, LHST), River Truyère, Entraygues, 44.65 ° N 2.57 ° E, 238 m a. s. l., + 12 ° C m. a. t., 22. ix. 1977, leg. H. Laville. — GERMANY, 1 adult male (as P. sagittalis, ZSMG), Tegel small lake, Berlin, 52.57 ° N 13.28 ° E, 38 m a. s. l., + 8 ° C m. a. t., 1932, leg. D. Scheer; 1 adult male (as Procladius sp., ZSMG), Überlingen, Andelshofen Neuweiher Reservoir, 47.8 ° N 9.2 ° E, 450 m a. s. l., + 9 ° C m. a. t., 8. viii. 1964, leg. F. Ringe; 1 adult male (Paratype of P. tenebricosus, as Procladius sp., ZFMK), Winningen, 50.31 ° N 7.51 ° E, 66 m a. s. l., + 10 ° C m. a. t., 20. ix. 2012, leg. B. Rulik [Barcode GBMWN 627 - 15]; 1 adult male (Holotype of P. tenebricosus, as P. choreus, ZFMK), Meyenburg, Fleet, wet meadow, 52.51 ° N 9.38 ° E, 37 m a. s. l., + 10 ° C m. a. t., 21. vi. 2017, leg. E. Resendiz and H. - G. Rudzinski [Barcode GBOL- 2596807]. — GREECE, 1 adult male (as Procladius sp., HCMR), River Pamisos, Vrachopanagista, 37.24 ° N 21.91 ° E, 90 m a. s. l., + 17 ° C m. a. t., 30. vi. 2002, leg. I. Karaouzas. — NORWAY, 1 adult male (as P. cf. cinereus, NTNU), Lake Jonsvatn, near Flaten, 63.40 ° N 10.55 ° E, 150 m a. s. l., + 4 ° C m. a. t., 14 ‒ 28. viii. 2014, leg. E. Stur [Barcode CHMNO 293 - 15]. — PORTUGAL, 1 adult male (as Procladius sp., DEBE), Albureira do Arade Reservoir, 37.24 ° N 8.37 ° W, 41 m a. s. l., + 17 ° C m. a. t., 10. v. 1996, leg. M. Rieradevall. — SPAIN, 1 adult male (as Procladius sp., DEBE), Gabriel Y Galan Reservoir, 40.13 ° N 6.06 ° W, 388 m a. s. l., + 16 ° C m. a. t., 25. vii. 1965, leg. N. Prat; 3 adult males (as Procladius sp., DEBE), Guadalajara, Entrepenas Reservoir, 40.29 ° N 2.43 ° W, 723 m a. s. l., + 14 ° C m. a. t., 9. xi. 1974, leg. N. Prat; 2 adult males (as P. culiciformis, DEBE), Cuenca, Contreras Reservoir, 39.27 ° N 1.38 ° W, 669 m a. s. l., + 14 ° C m. a. t., 7. xi. 1974, leg. N. Prat. — SWEDEN, 1 adult male (as Procladius sp., NHRS), Lake Boren, Borenshult, 58.57 ° N 15.10 ° E, 74 m a. s. l., + 6 ° C m. a. t., 11. vi. 1980, leg. Y. Brodin; 1 adult male (as P.? pruinosus, NHRS), Lake Vänern, Sjötorp boat harbour, 58.84 ° N 13.97 ° E, 44 m a. s. l., + 6 ° C m. a. t., 16. vi. 1987, leg. Y. Brodin; 1 adult male (as Procladius sp., NHRS), Mattön torrents, 60.27 ° N 16.91 ° E, 64 m a. s. l., + 5 ° C m. a. t., 24. vi. 2007, leg. Y. Brodin; 1 adult male (as Procladius sp., NHRS), River Dalälven, Tyttbo torrents, 60.19 ° N 16.68 ° E, 64 m a. s. l., + 5 ° C m. a. t., 7 ‒ 15. vii. 2007, leg. Y. Brodin; 1 adult male (as Procladius sp., NHRS), Gysinge torrents, 60.28 ° N 16.91 ° E, 63 m a. s. l., + 5 ° C m. a. t., 2. viii. 2007, leg. Y. Brodin. Diagnostic characters. Figs. 6, 12, 34, 89 ‒ 91, key couplet 17. P. tenebricosus has a medium long gonostylus process with a GspR that overlaps that of eight other species of Procladius in Europe. Of these, P. culiciformis, P. lugubris P. frigidus, P. ferrugineus and P. floralis are distinctly separated from P. tenebricosus by more than one mostly non-overlapping other morphological character in the key and the helpdesk. P. tenebricosus males can be distinguished morphologically from P. islandicus, P. pruinosus and P. longistilus by the dark marked wing (anal cell patch distinct to very distinct versus absent to faint), usually smaller size (wing length 1.8 ‒ 2.6 mm versus 2.4 ‒ 3.5 mm, mid leg tibia length 0.7 ‒ 1.1 mm versus 1.0 ‒ 1.4, body length 3.2 ‒ 4.3 mm versus 4.2 ‒ 6.0 mm) and frequently also lighter colour of the posterior half of tergite II ‒ IV (whitish to light brown versus light brown to dark brown). Neither the adult female, pupa or larva has been described. Pupal exuviae and larval skins from specimens reared to males are kept in NHRS. Barcodes of adult males are available and part of BIN BOLD: AER 3360 which has incorrectly been merged into BIN BOLD: AAG 5487. Geographical distribution and ecology. P. tenebricosus is probably a less frequent species in Europe found from southern Greece and Portugal at latitude 37 ° N in the south to mid Norway at latitude 63 ° N in the north. Mean annual temperature at the 16 sites with findings range from + 18 to + 4 ° C, thus from almost subtropical to temperate boreal climatic conditions. The altitude of the sites ranges from 40 m to 1 220 m above sea level. Most of the 15 sites with records of P. tenbricosus are at or close to running water, even torrents in one case. This implies that the species has an overall more lotic habitat choice than other species of Procladius in Europe, but there are also findings of P. tenebricosus at sites with stagnant water of freshwater lakes, lake-like reservoirs and ponds. Larvae of P. tenebricosus from samples taken at 0.5 m water depth in an exposed zone near a lake shore have been reared to adults. The habitat of the larvae is otherwise unknown. Judging from the findings of adult males, the larvae inhabit water with oligotrophic to eutrophic conditions. Adults have been collected from the middle of April to the middle of November. Countries with records of P. tenebricosus in Europe are France, Germany, Greece, Norway, Poland, Spain, Sweden and Switzerland. The species is possibly also recorded from Israel.	en	Brodin, Yngve (2025): Procladius (Diptera, Chironomidae) of Europe and a global view. Zootaxa 5591 (1): 1-127, DOI: 10.11646/zootaxa.5591.1.1, URL: https://doi.org/10.11646/zootaxa.5591.1.1
