Quedius, Stephens, 1829

Hansen, Aslak Kappel, Brunke, Adam, Simonsen, Thomas & Solodovnikov, Alexey, 2022, Revision of Quedius sensu stricto (Coleoptera: Staphylinidae), Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 62 (1), pp. 225-299 : 287-290

Key to species and species groups of Quedius sensu stricto

1 Presence of 4 or 6 additional setiferous punctures between anterior frontal punctures ( Fig. K1A ). Palearctic, two species introduced to the Nearctic. Q. fuliginosus -group .................................................. 5

– Presence of 2 or no additional setiferous punctures between anterior frontal punctures ( Fig. K1B ). ..... 2

2 Presence of 2 additional setiferous punctures between anterior frontal punctures ( Fig. K2A ). Nearctic. Q. molochinoides -group ..................................... 15

– No additional setiferous punctures between anterior frontal punctures ( Fig. K2B ). ................................. 3

3 Additional 1 to 3 punctures in the posterior half of the sublateral row ( Fig. K3A ). W Palearctic. Q. pallipes - group ...................................................................... 8

– No additional puncture in the posterior half of the sublateral row ( Fig. K3B ). ..................................... 4

4 Palearctic. Q. molochinus -group .......................... 9

– Nearctic. Q. laticollis -group and Q. molochinus introduced from Palearctic. .................................. 19

5 Scutellum punctured and pubescent ( Fig. K5A ). W Palearctic, usually preferring warmer drier habitats. ................................. Quedius levicollis Brullé, 1832

– Scutellum unpunctured and glabrous ( Fig. K5B ). ......................................................................................6

6 Head with 4 additional punctures between posterior frontal punctures, but without additional puncture between anterior and posterior frontal punctures ( Fig. K6A ). Median lobe without apical ridge. Paleartic. Introduced to multiple Nearctic localities. ............. 7

– Presence of 4 additional punctures between posterior frontal punctures and one additional puncture on either side between anterior and posterior frontal punctures ( Fig. K6B ). Median lobe with apical ridge ( Fig. 14A ). Palearctic, known only from North Africa and the islands of Sardinia and Corse. ...................... .................. Quedius afrofuliginosus Gusarov, 1991

7 Basal three antennomeres at least slightly darkened. Eyes clearly protruding making the head seem more rectangular wider across ( Fig. K7A ). Apex of median lobe not laterally flattened ( Fig. 14D ). Palearctic. Introduced to E Canada. Prefers wetter humus rich habitats. Paleartic. Introduced to multiple Nearctic localities. ................................................................... ................. Quedius fuliginosus ( Gravenhorst, 1802)

– Basal three antennomeres completely pale. Eyes not clearly protruding making the head seem round ( Fig. K7B ). Apex of median lobe laterally flattened ( Fig. 14C ). Paleartic. Introduced to multiple Nearctic localities. ..... Quedius curtipennis Bernhauer, 1908

8 Sides of median lobe apex folded with distinct tooth situated at lower part ( Fig. 15C ). Western Europe along Atlantic Coast from Denmark to Iberia and inland in Southern France and Iberian Peninsula, also on the islands of Madeira, Azores, Canaries, and Balearic ( Fig. K8A ). .................................................. ....................... Quedius hispanicus Bernhauer, 1898

– Sides of median lobe apex folded with distinct tooth situated at medial part ( Fig. 15D ). North Africa, Italy and islands of Sardinia and Corse ( Fig. K8B ). .......... ................................... Quedius pallipes Lucas, 1846

9 Elytra shortened, distinctly shorter than pronotum ( Fig. K9A ), obviously brachypterous species without whitish apical seam on abdominal tergite VII ( Fig. 11F ). East Palearctic. ................................................. .......................... Quedius sundukovi Smetana, 2003

– Elytra normal, not shortened, about as long as pronotum ( Fig. K9B ), with whitish apical seam on abdominal tergite VII. ............................................................ 10

10 Basal three antennomeres at least slightly darkened ( Fig. K10A ). ......................................................... 11

– Basal three antennomeres pale ( Fig. K10B ). ....... 13

11 Elytra most often reddish, very rarely darkened. Appendages only slightly darkened. Tibia, tarsi and apical antennomeres of antennae pale reddish brown ( Figs K11A, 11E ). Found near water bodies in Central-Eastern Europe and Middle Asia (Fig. 23). .. .................................. Quedius balticus Korge, 1960

– Elytra always dark. Appendages darkened. Tarsi slightly paler brown to dark brown at most ( Figs K11B , 10E–F ). Found at higher elevations or latitudes (Figs 25, 27). .......................................... 12

12 Head disc with fine isodiametric microsculpture ( Fig. K12A ). Without C-sclerite of the internal sac ( Fig. 16A ). Found at alpine sites of Central and Southern Europe ( Fig. K12C ). .................................................. ...................... Quedius unicolor Kiesenwetter, 1847

– Head disc most often with microsculpture of fine transverse waves, rarely isodiametric ( Fig. K12B ). With clear C-sclerite of the internal sac ( Fig. 15C ). Found disjunct in boreal Scandinavia and Russia, as well as alpine in Altai Mountains and Transbaikalia ( Fig. K12D ). ..... Quedius subunicolor Korge, 1961

13 Body brown. Elytra yellowish red. Apex of paramere pointing ventral, away from median lobe in form of a small blunt hook ( Figs K13B , 15A ). Central Anatolia, Middle East, Transcauscasus, Central Asia ( Fig. K13A ). ..... Quedius vicinus ( Ménétriés, 1832)

– Body most often black, or at most brownish. Elytra brick or brownish red. Apex of paramere straight, not pointing ventrad, away from median lobe ( Figs K13C–D , 15D, 15F ). Distribution more northern ( Figs K14A–B )...................................................... 14

14 C-shaped sclerite of internal sac without spine-like basal extension ( Figs K13C , 15F ). SE European distribution ( Fig. K14A ). ........................................... ........... Quedius meridiocarpathicus Smetana, 1958

– C-shaped sclerite of internal sac with spine-like basal extension ( Figs K13D , 15D ). N Palearctic distribution ( Fig. K14B ). ........................................... ............... Quedius molochinus ( Gravenhorst, 1806)

15 Basal three antennomeres pale and antennae more elongate (antennomere 10 clearly longer than wide) ( Fig. K15A ). Elytra light to dark brown. Generally larger. Forebody length 5.49–5.93 (5.78) mm ( Fig. 13F ). Boreal North America (Fig. 28). ...................... ..................... Quedius labradorensis Smetana, 1965

– Basal three antennomeres darkened and antennae more robust (antennomere 10 more or less transverse) ( Fig. K15B ). Elytra black or reddish. Generally smaller. Forebody length 4.97–5.86 (5.39) mm. ........................................................... 16

16 Paramere with basal attenuation and broadening towards apex forming a spoon like shape, peg setae situated in a semi-circle along the edge of the paramere ( Figs K16A , 8 , 17B ). Coastal to mid elevation from Oregon to Southern Alaska including Vancouver and Haida Gwaii. ..................................... ...................................... Quedius horni Hatch, 1957

– Paramere only with at most slight attenuation forming a lanceolate shape ( Figs K16B–D ). ...................... 17

17 Distributed across Boreal North America. South of the Northern Rocky Mountains ( Fig. K17A ). Teeth of median lobe clearly folded inwards ( Figs K16B , 17E ). ......... Quedius molochinoides Smetana, 1965

– Central and Southern Rocky Mountains, Cascades, Blue Mountains and Sierra Nevada ( Fig. K17B ). Teeth of median lobe either protruding or not clearly formed ( Figs K16C–D ). ....................................... 19

18. Found in the Cascades, Blue Mountains and Sierra Nevada ( Fig. K18A ). Paramere and median lobe clearly broader ( Figs K16C , 17D ). ............................ ....................................... Quedius lanei Hatch, 1957

– Found in Central and Southern Rocky Mountains ( Fig. K18B ). Paramere and median lobe clearly more slender ( Figs K16D , 17C ). ........................................ ........................................... Quedius altanai sp. nov.

19 Paramere rather slender and clear C-sclerite of the internal sac ( Figs K19A , 15D ). Introduced in parts of Eastern North America, seems to be spreading. ........ ............... Quedius molochinus ( Gravenhorst, 1806)

– Paramere rather broad and No C-sclerite of internal sac ( Figs K19B–C ). Q. laticollis -group .............. 20

20 Peg setae of paramere forming a solid triangular field, usually with two wide and short extensions posteriorly ( Figs K20A , 16E ). E North America ( Fig. K20B ). ........ Quedius laticollis ( Gravenhorst, 1802)

– Peg setae of paramere arranged into two short longitudinal rows, which more or less diverge posteriorly ( Figs K20C , 16F ). Great American Plains ( Fig. K20D ). ........... Quedius strenuus Casey, 1915

Figures

Fig. 1. Phylogenetic tree of Quedius s. str. based on preliminary target capture molecular dataset of Bඋඎඇĸൾ et al. (2021) and Hൺඇඌൾඇ et al. (in prep). Species groups shown in color. Q. altanai Hansen & Brunke, sp. nov. of the Q. molochinoides-group was not included in the dataset. Habitus sketch is of Q. fuliginosus (Gravenhorst, 1802). Lettering denotes species distribution within their respective groups.

Fig. 2. Habitats of Quedius s. str. A – Q. fuliginosus (Gravenhorst, 1802) in mossy habitat in a forest on Zealand, Denmark; B – Oak forest habitat of Q. levicollis Brullé, 1832 found in litter in Extremadura, Spain; C – Saline lake habitat of Q. meridiocarpathicus Smetana, 1958 found in bank litter on Peloponese, Greece; D – River bank habitat of Q. hispanicus Bernhauer, 1898 found in flood debris in Extremadura, Spain; E – Wet mixed forest habitat of Q. fuliginosus found in litter in Southern Ural, Russia; F – Alpine near tree line habitat of Q. subunicolor Korge, 1961 found under stones at 1700 m in Altai Mountains, Russia; G – Alpine mossy creek habitat of Q. unicolor Kiesenwetter, 1847 found in moss in Italian Alps (Photo:A. Zanetti).

Fig. 3. Phylogeny of Quedius s. str. based on Bayesian Inference (BI) and Maximum Likelihood (ML) analyses of COI barcodes (part 1). BI posterior probabilities (PP), ML ultrafast bootstrap support (UFB) and ML SH-aLRT test are shown. Nodes coloured as: green, PP ≥ 0.90, and either UFB ≥ 95 or SH-aLRT ≥ 80; yellow, one of PP ≥ 0.90, UFB ≥ 95 or SH-aLRT ≥ 80; red, PP <0.90, UFB <95 and SH-aLRT <80.

Fig. 5. Network analysis of Quedius molochinoides-group: Quedius horni Hatch, 1957 (purple), Q. lanei Hatch, 1957 (green), Q. molochinoides-group 1 (dark blue), Q. molochinoides-group 2 (light blue), and Q. molochinoides-group 3 (teal).

Fig.6. Head and pronotal chaetotaxy of Quedius s. str.A – general scheme; B – Q. curtipennis Bernhauer, 1908, C – Q. fuliginosus (Gravenhorst, 1802); D – Q. levicollis Brullé, 1832; E – Q. afrofuliginosus Gusarov, 1991; F – Q. molochinus- and Q. laticollis-groups; G – Q. molochinoides-group. Abbreviations as follows: anterior frontal puncture (AFP); basal punctures (BP); interocular punctures (IOP); lateral puncture (LP); lateral row (LR); posterior frontal puncture (PFP); parocular punctures (POP); sublateral row (SLR).

Fig. 14. Aedeagus of Quedius s. str. A – Q. unicolor Kiesenwetter, 1847; B – Q. levasseuri Coiffait, 1964 (syn. of Q. pallipes Lucas, 1846); C – Q. hispanicus Bernhauer, 1898; D – Q. pallipes Lucas, 1846; E – Q. laticollis (Gravenhorst, 1802); F – Q. strenuus Casey, 1915. I – Parameral view of whole aedeagus, II – apices of paramere in antiparameral view with peg setae, III – lateral view of whole aedeagus, IV – apices of median lobe at an angle between parameral and lateral view.

Fig. 7. Habitus of Quedius s. str. A – Q. afrofuliginosus Gusarov, 1991; B – Q. curtipennis Bernhauer, 1908; C – Q. fuliginosus (Gravenhorst, 1802); D – Q. levicollis Brullé, 1832; E – Q. unicolor Kiesenwetter, 1847; F – Q. subunicolor Korge, 1961.

Fig. 15. Variation of the shape of the paramere of Quedius sundukovi Smetana, 2003. Locality and BOLD BIN number are indicated under respective specimens.

Fig. 8. Habitus of Quedius s. str. A – Q. vicinus (Ménétriés, 1832); B – Q. meridiocarpathicus Smetana, 1958; C – Q. molochinus (Gravenhorst, 1806) (red elytra variant); D – Q. molochinus (black elytra variant); E – Q. balticus Korge, 1960; F – Q. sundukovi Smetana, 2003.

Fig. 9. Habitus of Quedius s. str. A – Q. hispanicus Bernhauer, 1898; B – Q. hispanicus (pale sutures variant); C – Q. pallipes Lucas, 1846; D – Q. laticollis (Gravenhorst, 1802) (black elytra variant); E – Q. laticollis (red elytra variant); F – Q. strenuus Casey, 1915.

Fig. 11. Aedeagus ofQuedius s. str. A – Q. afrofuliginosus Gusarov, 1991; B – Q. levicollis Brullé, 1832; C – Q. curtipennis Bernhauer, 1908; D – Q. fuliginosus (Gravenhorst, 1802). I – parameral view of whole aedeagus; II – apices of paramere in antiparameral view with peg setae; III – lateral view of whole aedeagus; IV – apices of median lobe at an angle between parameral and lateral view.

Fig. 10. Habitus of Quedius s. str. A – Q. horni Hatch, 1957; B – Q. lanei Hatch, 1957; C – Q. altanai Hansen & Brunke, sp. nov.; D – Q. molochinoides Smetana, 1965 (red elytra variant); E – Q. molochinoides (black elytra variant); F – Q. labradorensis Smetana, 1965.

Fig. 12. Distribution of Eurasian species introduced to North America: Quedius curtipennis Bernhauer, 1908 (teal), Q. fuliginosus (Gravenhorst, 1802) (green), and Q. molochinus (Gravenhorst, 1806) (orange).

Fig. 16. Aedeagus of Quedius s. str. A – Q. labradorensis Smetana, 1965; B – Q. horni Hatch, 1957; C – Q. altanai Hansen & Brunke, sp. nov.; D – Q. lanei Hatch, 1957; E – Q. molochinoides Smetana, 1965. I – parameral view of whole aedeagus, II – apices of paramere in antiparameral view with peg setae, III – lateral view of whole aedeagus, IV – apices of median lobe at an angle between parameral and lateral view.

Fig. 13. Aedeagus of Quedius s. str. A – Q. vicinus (Ménétriés, 1832); B – Q. sundukovi Smetana, 2003; C – Q. subunicolor Korge, 1961; D – Q. molochinus (Gravenhorst, 1806); E – Q. balticus Korge, 1960; F – Q. meridiocarpathicus Smetana, 1958. I – parameral view of whole aedeagus; II – apices of paramere in antiparameral view with peg setae; III – lateral view of whole aedeagus; IV – apices of median lobe in parameral view.

Fig. 17. Variation in genitalic morphology of species from Quedius molochinoides-group.From left to right for each specimen, first paramere then median lobe (parameral view) and lastly median lobe (lateral view) are given. For each specimen locality is noted underneath.

Fig. 18. Distribution of Quedius afrofuliginosus Gusarov, 1991 (top), Q. curtipennis Bernhauer, 1908 (middle) and Q. fuliginosus (Gravenhorst, 1802) (bottom).

Fig. 19. Distribution of Quedius levicollis Brullé, 1832 (top), Q. hispanicus Bernhauer, 1898 and Q. pallipes Lucas, 1846 (middle), and Q.balticus Korge, 1960 (bottom).

Fig. 20. Distribution of Quedius molochinus (Gravenhorst, 1806), Q. meridiocarpathicus Smetana, 1958, and Q. vicinus (Ménétriés, 1832) (top), Q. subunicolor Korge, 1961 (middle), and Q. sundukovi Smetana, 2003 (bottom).