#ZooKeys ZooKeys 1218: 81-98 (2024) DOI: 10.3897/zookeys.1218.135558 Research Article Two new rove beetle genera in Staphylininae that reduce “Heterothops’ and “Quedius’ taxonomic wastebaskets (Coleoptera, Staphylinidae) José L. Reyes-Hernandez'®, Alexey Solodovnikov'® 1 Natural History Museum of Denmark, University of Copenhagen, Zoological Museum, Universitetsparken 15, 2100, Copenhagen, Denmark Corresponding author: José L. Reyes-Herndndez (j/.reyeshdez@gmail.com) OPEN Qaccess Academic editor: Adam Brunke Received: 27 August 2024 Accepted: 19 October 2024 Published: 15 November 2024 ZooBank: https://zoobank. org/8C9034A6-1 E8D-4FF6-8721- 004826BB7797 Citation: Reyes-Hernandez JL, Solodovnikov A (2024) Two new rove beetle genera in Staphylininae that reduce “Heterothops” and “Quedius” taxonomic wastebaskets (Coleoptera, Staphylinidae). ZooKeys 1218: 81-98. https://doi.org/10.3897/ zookeys.1218.135558 Copyright: © José L. Reyes-Hernandez & Alexey Solodovnikov. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract The here-provided description of the new genera Chiquiticus gen. nov. and Nitidocolpus gen. nov. was necessitated by a phylogenetic study of Staphylininae (to be published separately), which will be used for the proper characterization of their respective new suprageneric lineages in an upcoming update of the higher classification of this sub- family. Both new genera are erected for species that had been previously described but misplaced in the highly polyphyletic “taxonomic wastebasket” genera Heterothops (Am- blyopinina) and Quedius (Quediina), resulting in the following new combinations: Chigq- uiticus arizonicus (Smetana, 1971), comb. nov. ex. Heterothops; Chiquiticus campbelli (Smetana, 1971), comb. nov. ex. Heterothops; tChiquiticus cornelli (Chatzimanolis & Engel, 2013), comb. nov. ex. Heterothops; Chiquiticus gemellus (Smetana, 1971), comb. nov. ex. Heterothops; tChiquiticus infernalis (Chatzimanolis & Engel, 2013), comb. nov. ex. Heterothops; Chiquiticus occidentis (Casey, 1886), comb. nov. ex. Heterothops; Chig- uiticus pusio (LeConte, J. L., 1863), comb. nov. ex. Heterothops; Chiquiticus rambouseki (Blackwelder, 1943), comb. nov. ex. Heterothops; Nitidocolpus aurofasciatus (Bernhauer, 1917), comb. nov. ex. Quedius; Nitidocolpus championi (Sharp, 1884), comb. nov. ex. Quedius; Nitidocolpus columbinus (Bernhauer, 1917), comb. nov. ex. Quedius; Nitido- colpus germaini (Bernhauer, 1917), comb. nov. ex. Quedius; Nitidocolpus illatus (Sharp, 1884), comb. nov. ex. Quedius; Nitidocolpus laeticulus (Sharp, 1884), comb. nov. ex. Que- dius; Nitidocolpus triangulum (Fauvel, 1891), comb. nov. ex. Quedius. Additionally, sev- eral more Neotropical Quedius species, which resemble Nitidocolpus, have been revised and transferred to the amblyopinine genus Cheilocolpus or the cyrtoquediine genus Cyrtoquedius, with the following new combinations: Cheilocolpus forsteri (Scheerpeltz, 1960), comb. nov. ex. Quedius; Cheilocolpus speciosus (Bernhauer, 1917), comb. nov. ex. Quedius; Cheilocolpus viridulus (Erichson, 1840), comb. nov. ex. Quedius; Cyrtoquedi- us viridipennis (Fauvel, 1891), comb. nov. ex. Quedius. The undescribed species diversity of the newly described genera is also highlighted. Key words: Cheilocolpus, Chiquiticus gen. nov., Cyrtoquedius, Neotropical fauna, new combinations, new lineages, Nitidocolpus gen. nov., taxonomy 81 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Introduction During a large-scale systematic study of Staphylininae rove beetles (in the broad sense of Newton 2022), which integrates phylogenomic, morphologi- cal, and biogeographic evidence for the first time (Reyes-Hernandez et al. in prep.), we discovered two new genera, Chiquiticus gen. nov. (Figs 1-3, 4A-C) and Nitidocolpus gen. nov. (Figs 5, 6A—C), both from the New World. These genera represent two previously unrecognized phylogenetic lineages with- in Staphylininae. The formal recognition of these lineages addresses a tax- Onomic gap, with further details to be presented in a forthcoming paper by Reyes-Hernandez et al. (in prep.). Currently, both new genera are classified as Staphylininae incertae sedis. Chiquiticus gen. nov. belongs to a newly recovered lineage that also includes the genus Ctenandropus Cameron, 1926 (now in the subtribe Amblyopinina of Staphylinini), the genus Amazonothops Jenkins Shaw, Orlov & Solodovnikov, 2020 (now Staphylinini incertae sedis), and a few extant and extinct (from Do- minican amber) Nearctic and Neotropical species of the genus Heterothops Stephens, 1829 (now in the subtribe Amblyopinina). The small Australo-Asian genus Ctenandropus is morphologically and biogeographically peculiar and poorly known. Amazonothops is also a small Neotropical genus that was dis- covered only recently (Jenkins Shaw et al. 2020). Chiquiticus gen. nov. is erect- ed for the few species of Heterothops that in fact do not belong to that genus and even to the subtribe Amblyopinina. Nitidocolpus gen. nov. forms a monogeneric lineage comprising several Neotropical species of “Quedius” Stephens, 1829, primarily those related to Q. columbinus Bernhauer, 1917. The phylogenetic analysis of Reyes-Hernandez et al. (in prep.) reveals that this lineage is distinct from true Quediina, as defined by Brunke et al. (2021). However, some species currently classified as Quedius, although superficially similar to Q. columbinus and its allies, were found to be more closely related to the genus Cheilocolpus Solier, 1849, both phylogenetical- ly and morphologically. As a result, these species are transferred here to Chei- locolpus. Below, we present, justify, and discuss all these taxonomic novelties. Material and methods The studied specimens are deposited in the following collections: AMNH (American Museum of Natural History, New York, USA; D.A. Grimaldi and A. Pierwola), CNC (Canadian National Collection, Ontario, Canada; A. Brunke), FMNH (Field Museum of Natural History, Chicago, Illinois, USA; M. Turcatel, A.F. Newton, M.K. Thayer), IRSNB (Institut Royal des Sciences Naturelles de Bel- gique, Brussels, Belgium; W. Dekoninck), NHM (The Natural History Museum, London, United Kingdom; M. Barclay and D. Telnov), MCZ (Museum of Compar- ative Zoology, Harvard University, Cambridge, Massachusetts, USA; C. Maier), and MFNB (Museum fir Naturkunde an der Humboldt Universitat, Berlin, Ger- many; B. Jager). Specimens were examined with a Leica M125 dissecting microscope (Lei- ca Microsystems, Switzerland). Photographs were captured using a Canon 5D ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 89 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Mark III camera with a Canon MP-E 65mm f/2.8 1—5x macro lens (Canon, Ja- pan) and a StackShot 3x (Cognisys, USA). Images were then stacked using Zerene Stacker (Zerene Systems, USA) with the PMax function. Further image processing, including cropping, lightening, drawing lines, and adding scales, was performed in Adobe Photoshop 2023. Morphological terminology mainly follows Li and Zhou (2011), Brunke et al. (2019), and Reyes-Hernandez et al. (2024), with the following modifica- tions in prothorax and pterothorax terminology to align with that of Herman (2023). In the ventral view of the prothorax, what was formerly referred to as the basisternum and furcasternum, separated by the sternacostal ridge, is now termed the upper probasisternum (UBS, Fig. 1E) and lower probasister- num (LBS, Fig. 1E), respectively, separated by the intercoxal carina (ICC, Fig. 1E). In Staphylininae, the profurcasternum is a significantly reduced scler- ite defined by an internal ridge connecting the apophyseal invaginations (Herman 2023). In the mesothorax (ventral view), the anterior margin is re- ferred to as the external part of prepectus (EXPP, Fig. 1F), the sternopleural (anapleural) suture is termed as mesanapleural sutures (MNPS, Fig. 1F), and in the metathorax, the marginal carina of the mesocoxal acetabuli is called the pericoxal ridge. Regarding the frontoclypeal (epistomal) “suture”, here we clarify that it represents the connection of the anterior tentorial arms (CATA, Fig. 4A) inside the head capsule, which is superficially visible externally as a dark line (Figs 2B, 3B; see also fig. 7D, E in Reyes-Hernandez et al. 2024). This “suture” or ridge is more clearly visible in teneral or chem- ically clarified specimens, while in highly sclerotized species it appears as a shallow line on the surface as well. The supra-antennal ridge (SAR) is the term used here for a structure referred to as the supra-antennal carina in the figure legend of Brunke (2022: fig. 3A, B) but described as the suprantennal ridge in the main text. It represents the fold of the anterior or anterolateral margin of the supra-antennal tubercle which may or may not be carinated (Figs 4, 6). Additionally, new characters are introduced and illustrated here, such as the anterolateral clypeal punctures (ACP, Figs 4, 6), which are setif- erous punctures laterally adjacent to the frontoclypeal punctures on the dor- sal portion of the head. On the abdominal segments, the medial macroseta (MMA, Fig. 1D) and posteromarginal large macroseta (PMA, Fig. 1D) are also indicated. In certain Staphylininae species, the base of the paramere (on its outer side around the area of its attachment to the median lobe) exhibits an upward or forward projection that is most distinctly visible in lateral view (Fig. 61, L, O). Abbreviations for measurements are as follows: EYL (eye length in lateral view), HL (head length from the apex of the clypeus to the nuchal ridge, or when the latter is dorsally absent, then to a hypothetical line joining the sides of the nuchal ridge or the groove marking the nuchal constriction), HW (head width at the widest point, including eyes), GL (gena length), NW (neck width at the widest point), PL (pronotum length along the median line), PW (pronotum width at the widest point), and TL (total length from the anterior margin of the clypeus to the posterior margin of segment VIII). All measurements were taken in millimeters using an ocular micrometer on a dissecting microscope. ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 83 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Results Family Staphylinidae Latreille, 1802 Subfamily Staphylininae Latreille, 1802 Tribe incertae sedis Genus Chiquiticus gen. nov. https://zoobank.org/4BCF1 2DF-8231-4269-B13B-917E194D43B9 Figs 1-3, 4A-C Type species. Heterothops pusio J.L. LeConte, 1863, here designated. Included species. Chiquiticus arizonicus (Smetana, 1971), comb. nov. ex. Het- erothops [holotype and 2 paratypes from CNC examined]; Chiquiticus campbelli (Smetana, 1971), comb. nov. ex. Heterothops [holotype, and 7 paratypes from CNC examined]; tChiquiticus cornelli (Chatzimanolis & Engel, 2013), comb. nov. ex. Heterothops [photos of the holotype and one paratype from AMNH exam- ined]; Chiquiticus gemellus (Smetana, 1971), comb. nov. ex. Heterothops [holo- type from MCZ and 2 paratypes from CNC 2 examined]; tChiquiticus infernalis (Chatzimanolis & Engel, 2013), comb. nov. ex. Heterothops [moved to Chiquiti- cus based on the data in the publication]; Chiquiticus occidentis (Casey, 1886), comb. nov. ex. Heterothops [holotype of Heterothops mediocris Fall, 1907, from MCZ examined, H. mediocris is a junior synonym of H. occidentis according to Smetana (1971) and non-type material identified by A. Smetana in CNC]; Chig- uiticus pusio (LeConte, J. L., 1863), comb. nov. ex. Heterothops [1 presumed syntype or at least historical specimen examined, (MCZ) conspecific with the type material according to Smetana (1971) and non-type material identified by A. Smetana in CNC]; Chiquiticus rambouseki (Blackwelder, 1943), comb. nov. ex. Heterothops [1 syntype from NHM examined]. Diagnosis. Small Staphylininae mainly around 2.5-3.5 mm long (Fig. 1A, B) with subconical head with ventral basal ridge but without postgenal ridge (Fig. 1C); without supra-antennal punctures (Fig. 4A); pronotum transverse (PW/PL = 1.1), with its maximum width in posterior half, with paired punctures on dorsal series widely separated from each other (Fig. 4B); mesanapleural sutures trans- verse not reaching or fusing with external part of prepectus (Fig. 1F); males with black combs on first mesotarsomeres, but without black combs on mesotro- chanters; tergites VII and VIII without broader, foliose setae in addition to usual acuminate, simple setae; tergite X not fused to lateral tergal sclerites in males. Description. Body small (TL = 2.3-3.9 mm). Head (Fig. 1C): frontoclypeal (epistomal) “suture” complete; head about as wide as long (HW/HL > 0.9 but < 1.1); neck from moderately (NW/HW > 0.75 but < 0.90) to distinctly wide (NW/ HW = 0.90); dorsal macrosetae: anterolateral clypeal punctures present; fron- toclypeal punctures present; supra-antennal punctures absent; parocular punc- tures present as one on each side or absent; basal punctures present, single; posterior frontal punctures located posterior to temporal punctures; gular su- tures separated, gula with distinct transverse basal impression; ventral basal ridge (VBR) present, postgenal ridge absent; postmandibular ridge (PMR) ex- tended diagonally towards gula; infraorbital ridge extends to PMR; nuchal ridge absent or rudimentary, at most present as a linear impression but not a ridge; eye from medium large (EYL/HL ratio > 1/2 but < 3/4) to very small (EYL/HL ra- tio less than 3/10); antennomere 1 distinctly shorter than antennomeres 3 and ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 84 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Figure 1. Some Chiquiticus species and characters A C. sp. nr. pusio B C. rambouseki (Blackwelder, 1943) C C. pusio (Le- Conte, J. L., 1863), head ventral view D C. pusio, abdomen E C. pusio, prosternum F C. pusio, mesothorax. Abbreviations: ASP, antesternal plate; EXPR, external part of prepectus; FS, profurcasternum; ICC, intercoxal carina of the probasister- num; INPR, internal part of prepectus; IOR, infraorbital ridge; LBS, lower probasisternum; MMA, medial macroseta; MNPS, mesanapleural sutures; PMA, posteromarginal large macrosetae; PMR, postmandibular ridge; POR, postoccipital ridge; UBS, upper probasisternum; VBR, ventral basal ridge. Scale bars: 1 mm (A, B, D); 0.1 mm (C, E, F). ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 85 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets 2 combined; male and female antennomere 11 at least two times longer than antennomere 10; left mandible with proximal, not bifid tooth; maxillary palps: palpomere 4 (apical) subulate, distinctly shorter than palpomere 2, palpomere 3 markedly dilated compared to palpomere 4; labial palps: palpomere 3 (apical) more or less cylindrical, distinctly narrower and longer than palpomere 2, paral- lel-sided and needle-shaped, palpomere 2 markedly dilated at apex. Thorax (Fig. 1E, F): prothorax with slightly transverse pronotum (PW/PL = 1.1) with one pair of setiferous punctures in dorsal series widely separated from each other, the distance between them being about equal to the distance be- tween them and the lateral margin; sublateral setiferous punctures present or absent; sternacostal (transverse carina) ridge medially not protruding; antester- nal membrane with distinct semisclerotized plate (Fig. 1E); pronotum and pros- ternum not fused in procoxal cavity, pronotosternal suture complete in cavity; pronotal hypomeron not setose, without postcoxal hypomeral process; upper probasisternum with or without pair of macrosetae. Mesothorax: mesoscutel- lum without posterior scutellar carina, without sub-basal ridge; elytron without humeral spines or spine-like setae, with evenly setiferous punctation on disc and epipleuron; mesanapleural sutures (Fig. 1F) transverse not reaching or fusing with external part of prepectus, medially fused or nearly touching each other; apex of mesobasisternal intercoxal process varies from sharply point- ed to obtuse angle, without V-shaped projection medially; mesocoxal cavities contiguous; pericoxal ridge absent. Metathorax with wings present, with veins CuA and MP4 fused in one vein; metakatepisternal processes divided. Legs: protarsomeres expanded in both sexes; apical tarsomere of all legs with one empodial seta distinctly shorter than tarsal claws; protibial row of laterodorsal spines always present in females but maybe absent in males in some species; ventral tibial spur resting at base of apical excavation connected to tibial mar- gin by thin membranous region; first segment of mesotarsi in males with black comb lateroventrally, without pale adhesive setae; mesotrochanters in both sex- es without black comb. Abdomen: tergites III-V with only anterior transverse basal carina; tergites III-VI with a large posteromarginal macroseta on each side (Fig. 1D); tergites VII and VIII without broader, foliose setae in addition to the usual acuminate, simple setae; anterior transverse basal carina of tergite VII not continuing to paraterg- ites; lateral tergal sclerites IX cylindrical; male sternite VIII with medial apical emargination; male sternum IX symmetrical. Aedeagus with rounded apex. Distribution. Nearctic region, Central America, the Caribbean. Introduced into Central Europe: C. pusio in Germany (Schilke and Renner 2020) and anoth- er species related to C. pusio collected in Austria with a car net. Ecology. The North American species occur in various types of debris like leaf litter and similar substrates, with some species also occurring facultatively in mammal burrows or nests (Neotoma Say & Ord, 1825), especially in drier regions (Smetana 1971; Adam Brunke pers. comm.). In Europe, where they are introduced, they have been collected using car nets. Comparison. Chiguiticus gen. nov. can be distinguished from all other genera of Staphylininae in broad sense (Newton 2022) by the combination of charac- ters mentioned in the diagnosis. From species close to the type of Heterothops, it is distinguished by the paired setiferous macropunctures of the dorsal series being distinctly separated from each other, by a black comb on first mesotar- ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 86 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets if | ee ; a | a i : EPs sad | i gee s - = -, ‘ ww \ Ne as 5 a Ae: Na Figure 2. Chiquiticus arizonicus (Smetana, 1971) (Holotype #CNC935711) A dorsal habitus B head C pronotum. Abbrevi- ations: CATA, connection of the anterior tentorial arms (frontoclypeal “suture”); PPDS, paired punctures on dorsal series. Photos of # CNC935711 (J. Buffam, Canadian National Collection of Insects, Arachnids and Nematodes). somere of males, and by presence of only one posteromarginal macrosetae on each side of tergites III-VI. In species close to the type of Heterothops, the paired setiferous macropunctures of the dorsal series are close together, the distance between them being distinctly less than the distance between dorsal row and respective lateral margin of pronotum. Also, in the true Heterothops, males do not have a black comb on the first mesotarsomere and their pattern of posteromarginal large macrosetae on tergites III-VI is 1, 1, 2 and 2 per side. For more details on the differentiation between Chiquiticus gen. nov. and Heterotho- ps species from Central Europe, see Schiilke and Renner (2020). Presence of a frontoclypeal “suture” is a rare trait in Staphylininae, which in Chiguiticus gen. nov. is always visible. In Staphylininae this character is found in few basal lin- eages, like Arrowinus Bernhauer, 1935, several genera of Amblyopinina including Loncovilius Germain, 1903 (Reyes-Hernandez et al. 2024), and in the relatives to Chiquiticus, Amazonothops (Jenkins Shaw et al. 2020), and Ctenandropus. ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 87 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Figure 3. tChiquiticus cornelli (Chatzimanolis & Engel, 2013) (paratype AMNH DR-10-1214) A dorsal habitus B head C pro- notum. Abbreviations: CATA, connection of the anterior tentorial arms (frontoclypeal “suture”); PPDS, paired punctures on dorsal series. Photo of AMNH DR-10-1214 (A. Pierwola, American Museum of Natural History). Scale bars: 0.1 mm. Additionally, Chiquiticus gen. nov. displays an antesternal plate, currently known as a unique character of groups such as Arrowinus and several tribes united in Xatholininae sensu Zyta and Solodovnikov (2020). However, this plate some- times is also found in Amblyopinina among Staphylinini, for example in species that are close to the types of the currently non-monophyletic genera Heterotho- ps and Cheilocolpus (Reyes-Hernandez et al. 2024). Furthermore, Chiquiticus exhibits unique characters shared only with the related genera Ctenandropus and Amazonothops, such as the presence of transverse mesanapleural sutures distinctly separated from the external portion of the prepectus (Fig. 1F and fig. 3E in Jenkins Shaw et al. (2020) misinterpreted as “transvere ridge (TR)”) and distinctly elongated protergal glands (fig. 3F in Jenkins Shaw etal. (2020)). Chig- uiticus is distinguished from Ctenandropus by the subconical head with ventral basal ridge; the transverse pronotum (PW/PL = 1.1) with its maximum width in the posterior half; usually by presence of intercoxal carina separating the up- per probasisternum and lower probasisternum; and by tergites III-V lacking a posterior transverse carina. In Ctenandropus, the head is subquadrate, without postgenal ridge or ventral basal ridge; the pronotum is as long as wide (PW/ PL > 0.9 but < 1.1), with its maximum width in the anterior half; without intercox- al carina separating the upper probasisternum and lower probasisternum; and Zookeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 88 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets A B Figure 4. Morphological details of Chiquiticus and Heterothops A-C Chiquiticus sp. aff. pusio from Panama D-F Hetero- thops sp. undescribed from Costa Rica A, D head B, E pronotum C, F aedeagus in lateral view, the scale to the right is equal to 1 mm. Abbreviations: ACP anterolateral clypeal puncture; AFP, anterior frontal; AMRS, anterior marginal row of setiferous punctures; BP, basal puncture; CATA, connection of the anterior tentorial arms (frontoclypeal “suture”); FCP, frontoclypeal puncture; [ASP internalmost setiferous puncture of the anterior margin; IOR, infraorbital ridge; LLSP large lateral setiferous puncture; LPSP, large posterior setiferous puncture; NR, nuchal ridge; PFP. posterior frontal puncture; POP parocular punctures; PPDS, paired punctures on dorsal series; SAR, supra-antennal ridge; SLSP sublateral setiferous puncture; TP, temporal puncture. tergites III-V have a posterior transverse carina. Chiquiticus is distinguished from Amazonothops by the absence of a black comb on the mesotrochanters of males and foliaceous setae on the tergites in both sexes, by distinctly short empodial setae, by the tergite X not fused to internal face of lateral sclerites and by paramere with rounded apex and short lateroapical setae. Etymology. The name is derived from the Latinization of the word “Chiquiti- co”, which is a term used in some Hispanophone countries to refer to very small things. The gender is masculine. New combinations notes. The extinct species tC. infernalis from Dominican Amber is here restudied based on the original descriptions only. Many details important for the taxonomic placement of this species (e.g. pronotum and me- sanapleural sutures) are not visible in the photos of this fossil there and prob- ably they would be available only with the mCT examination. However, such visible features as habitus including small body size, as well as antennal bases located close to each other, very elongated last antennomere that is almost as long as the two preceding ones, oval and setose preapical maxillary palpomere that is clearly wider than the apical one, and wide neck allowed us to conclude that TC. infernalis is an extinct member of the new genus Chiquiticus. For tC. cornelli, in addition to the original description, we were able to examine better ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 89 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets photos of the holotype and paratype kindly made available at our request (Fig. 3). In these images, in addition to the characters previously described for tC. infernalis, the connection of the anterior tentorial arms (Fig. 3B) and the widely separated paired punctures in the dorsal series (Fig. 3C) are visible. It should be noted that both fossil species are known only from females. Also, two spec- imens of the extant C. rambouseki (Blackwelder, 1943) that we examined were females and not in the best condition; however, the combination of their visible characters allows the species to be placed in Chiquiticus. Biogeographic note. As far as currently known, the Nearctic is the region with the greatest species diversity of Chiquiticus. However, the Chiquiticus fos- sils found in mid-Miocene Dominican amber (Chatzimanolis and Engel 2013) could suggest that Nearctic speciation took place after the genus dispersed there from the southern hemisphere where its closely related genera Amazo- nothops and Ctenandropus are also found (Reyes-Hernandez et al. in prep.). Such dispersal may have occurred via the GAARIandia land bridge (Iturralde-Vi- nent 2006; Masonick et al. 2017; Iturralde-Vinent and MacPhee 2023). Genus Nitidocolpus gen. nov. https://zoobank.org/9BC238C3-B742-4E1 E-8B59-48E8C83288DF Figs 5, 6A-C Type species. Quedius co/umbinus Bernhauer, 1917, here designated. Included species. Nitidocolpus aurofasciatus (Bernhauer, 1917), comb. nov. ex. Quedius [1 syntype from FMNH examined]; Nitidocolpus championi (Sharp, 1884), comb. nov. ex. Quedius [1 syntype from FMNH examined]; Nit- idocolpus columbinus (Bernhauer, 1917), comb. nov. ex. Quedius [2 syntypes from FMNH examined]; Nitidocolpus germaini (Bernhauer, 1917), comb. nov. ex. Quedius [1 syntype from FMNH examined]; Nitidocolpus illatus (Sharp, 1884), comb. nov. ex. Quedius [1 syntype from NHM examined]; Nitidocolpus laeticulus (Sharp, 1884), comb. nov. ex. Quedius [1 syntype from NHM ex- amined]; Nitidocolpus triangulum (Fauvel, 1891), comb. nov. ex. Quedius [1 syntype from IRSNB examined]. Diagnosis. Supra-antennal punctures and paraocular punctures absent (Fig. 6A); pronotum without paired dorsal series of setiferous macropunctures (Fig. 6B); mesanapleural sutures transverse or nearly transverse but reaching the exter- nal part of prepectus (Fig. 5E); mesobasisternal intercoxal process with V-shaped projection medially (Fig. 5E); metatrochanters with apical spine (Fig. 5F/). Description. Small to medium-sized Staphylininae (TL = 4.5-10 mm) (Fig. 5A-D). Head: Frontoclypeal (epistomal) suture absent or distinct only laterally and only in teneral (or chemically cleared) specimens; head slightly transverse (HW/HL = 1.1) without distinct posterior angles; disc smooth; frons with mi- crosculpture as transverse waves, without distinct concavity between antennal insertions; neck moderately wide (NW/HW > 0.75 but < 0.90); dorsal macrose- tae (Fig. 6A): anterolateral clypeal punctures present; frontoclypeal punctures present; supra-antennal punctures absent; parocular punctures absent; basal punctures present, single; ventral macrosetae: postocular punctures present; infraorbital punctures absent; postmandibular punctures present; submentum with two pairs of macrosetae; mentum with seta alpha only, seta beta absent; ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 90 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Figure 5. Some Nitidocolpus species and characters AN. illatus (Sharp, 1884) B N. /aeticulus (Sharp, 1884) C N. germaini (Bernhauer, 1917) D N. aurofasciatus (Bernhauer, 1917) EN. illatus, pterothoraces F N. columbinus (Bernhauer, 1917), metacoxa and metatrochanter, dissected right leg in dorsal view (side faced to the abdomen). Abbreviations: EXPR, external part of prepectus; INPR, internal part of prepectus; MIV, mesobasisternal intercoxal process with V-shaped pro- jection medially; MNPS, mesanapleural sutures; MTMS, metacoxa with dorsomedial spine; MTTAS, metatrochanter with dorsoapical spine. Photo B by J. Jenkins Shaw. Scale bars: 1 mm. ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 91 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets gular sutures separated from each other, gula with distinct transverse basal impression; ventral basal ridge present; postgenal ridge present; postmandib- ular ridge short, extended parallel to margin of eye, without fusion with any other ridge and distinctly separated from eye margin; infraorbital ridge merged with postgenal ridge; nuchal ridge missing dorsally, present laterally, merged with infraorbital ridge; eyes of medium size (EYL/HL ratio more than 1/2 but less than 3/4); gena short (GL/ETL < 0.5); tomentose pubescence begins from fourth antennomere, its density on fourth antennomere as on following anten- nomeres; setation of antennomere 3 almost evenly distributed; antennomere 11 with subapical rounded field, without subapical lateral pits; mandibles with dorso-lateral seta, without dorso-lateral groove, external edge with curved base and apex, straight in middle length; right mandible with proximal tooth and bifid distal tooth, apically deflexed ventrad at an angle of >20°; left mandible rather straight; labrum with transparent wide apical membrane, anterior margin emar- ginate at middle; maxillary palps: palpomere 4 (apical) subconical, with evenly narrowed apex, longer than palpomere 2 and weakly dilated compared to pal- pomere 3; labial palps: palpomere 3 (apical) more or less fusiform or subconi- cal, distinctly longer than, and about as wide as, palpomere 2. Thorax: Prothorax with slightly transverse pronotum (PW/PL = 1.1), with- out dorsal or sublateral series of setiferous punctures; prosternum without longitudinal keel; antesternal membrane without distinct semisclerotized patch or patches; probasisternum triangular, with narrowed lateral arms and disc protruding medially, with pair of macrosetae on the upper proba- siternum; with rounded postcoxal hypomeral process, interrupted by inferior line. Mesothorax: mesoscutellum without posterior scutellar carina, without sub-basal ridge; elytra with humeral spines or spine-like setae, with even setiferous punctation on disc and epipleuron (sometimes with various se- tose color patterns); with setiferous punctures at apical margin of elytral suture (underside); mesanapleural sutures transverse or nearly transverse but reaching and fusing with external part of prepectus; mesobasisternum with intercoxal process narrowly pointed into sharp angle, without V-shaped projection medially; mesocoxal cavities contiguous; pericoxal ridge present and complete. Metathorax: wings present, with veins CuA and MP4 fused; metakatepisternal processes divided; metascutellar mid-longitudinal inter- nal suture well developed (only visible in chemically cleared specimens). Legs: apical tarsomere of all legs without dorsal setae, with one empodi- al seta distinctly shorter than tarsal claws; protarsomeres 1-3 distinctly wider than long in both sexes, with pale adhesive setae ventrally; procoxa with internal ridge running parallel along external ridge; mesotarsi in both sexes without black comb, sometimes in males ventral side of first meso- tarsomere with pale adhesive setae; mesotibiae straight; metatarsomere 1 shorter than metatarsomere 5; metatarsi shorter than metatibiae; metatro- chanter apically rounded, with strong straight dorsoapical spine; metacoxae with four or fewer spines on ventral posterolateral lobe, spines on dorsome- dial disc also present; basal part of metacoxae distinctly wider and more convex than apical part. Abdomen: protergal glands well manifested as rounded cuticular acetabu- la; tergites III-V in some species with posterior basal transverse carinae, in some species they are only with anterior carinae; on tergite VII anterior trans- ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 92 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets B a: ar ( «—APP NS AMRSP IASP [sd [oe LLSP Figure 6. Morphological details of Nitidocolpus and other relevant genera of Staphylininae A-C Nitidocolpus columbinus D-F Cheilocolpus viridulus G-| Cyrtoquedius sp. J-L Quedionuchus nr. impunctus M-O Quedius nr. advena. A, D, G, J, M head B, E, H, K, N pronotum C, F, I, L, O aedeagus in lateral view, the black arrow pointing down indicates the basal projection of the paramere. Abbreviations: ACP anterolateral clypeal puncture; AFP anterior frontal; AMRSP, anterior marginal row of setiferous punctures; APP, additional paired punctures adjacent to AMRSP; BP. basal puncture; FCP frontoclypeal puncture; IASP, internalmost setiferous puncture of the anterior margin; IOP interocular punctures; IOR, infraorbital ridge; LLSP large lateral setiferous puncture; LPSP, large posterior setiferous puncture; NR, nuchal ridge; PFP, posterior frontal puncture; POP, parocular punctures; PPDS, paired punctures on dorsal series; SAP, supra-antennal punc- ture; SAR, supra-antennal ridge; SLSP sublateral setiferous puncture; TP, temporal puncture. Scale bars: 1 mm. ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 93 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets verse basal carina not continuing to paratergites; tergites III-VI with different patterns of posteromarginal large macroseta (PMM) on each side, IV-VI with more than one PMM per side; punctation of tergites in form of fine to moderate impressions, some species with wide glabrous areas; lateral tergal sclerites IX short and slightly laterally flattened; male sternite VIII with medial apical emar- gination. Aedeagus with paramere fused to median lobe only at base and very closely appressed to median lobe along entire length; base of the paramere flat medially, not projecting upwards in the middle; paramere strongly produced over apex of median lobe. Ovipositor with each second gonocoxite with one medial macroseta, without spine-like setae on outer lateral margin. Distribution. Neotropical Realm: Mexican Transition Zone, Central America, and northern South America. Ecology. This genus has been collected from a variety of microhabitats, in- cluding mushrooms (e.g., Pleurotus spp.), under decomposing logs, beneath bark, and in leaf litter found in cloud, oak, and pine forests. They are also fre- quently captured using flight-intercept and Malaise traps. Additionally, some species are associated with the inflorescences of plants from the families Are- caceae, Betulaceae, and Heliconiaceae. Comparison. Nitidocolpus gen. nov. can be distinguished from all other gen- era of Staphylininae by the combination of characters mentioned in the diagno- sis. Nitidocolpus is further distinguished from the superficially similar Quediina and Cytoquediina (Fig. 6) by the lack of supra-antennal punctures, the presence of a single carina on the mesoscutellar shield (posterior carina absent), by hind wings with fused CuA and MP4 veins, by a single empodial setae, and by the base of the paramere, which has no upward or forward projection in the mid- dle. Compared to other Neotropical genera that resemble Nitidocolpus, such as Cheilocolpus, Mimosticus Sharp, 1884, and Heterothops, Nitidocolpus is dis- tinguished by the absence of paraocular punctures on the head, the lack of setiferous punctures in or near the center of the pronotal disc, and the presence of postcoxal hypomeral processes. In some species of Nitidocolpus, such as N. laeticulus (Fig. 5B) and N. aurofasciatus (Fig. 5D), the basal punctures have shifted anteriorly and thus may be erroneously considered paraocular punc- tures. However, they can be easily distinguished as basal punctures, because they are the only punctures located near the posterior frontal punctures, aside from the temporal punctures. In other species of Nitidocolpus, such as N. illatus (Fig. 5A), the innermost macrosetae on the anterior margin maybe erroneously considered as a puncture of the dorsal row because it is located far from the anterior margin of pronotum. Etymology. The name is derived from the Latin words “nitidus” and “colpus”, which mean “shiny” and “hit”, respectively. The name refers to the great clarity with which the head and pronotum punctures are seen. The gender is masculine. New combinations in the genera Cheilocolpus Solier, 1849 and Cyrtoquedius Bernhauer, 1917 As shown above, describing the genus Nitidocolpus required careful examina- tion of a number of the poorly known species of Quedius, some of which, as we should stress, do not belong to Nitidocolpus. They belong to neither Quedius nor Quediina. Our examination revealed that most of them belong to the ambly- ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 94 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets C Figure 7. Some Cheilocolpus species A Ch. pyrostoma (Solier, 1849) B Ch. speciosus (Bernhauer, 1917) C Ch. viridulus (Erichson, 1840). opinine genus Cheilocolpus, as it is defined in Coiffait and Saiz (1966) and Saiz (1971), with adjustments by Reyes-Hernandez et al. (2024); i.e., they are close to the type species Cheilocolpus pyrostoma (Solier, 1849) (Fig. 7A). As a result, here we move these species as follows: Cheilocolpus forsteri (Scheerpeltz, 1960), comb. nov. ex. Quedius [although the type material was not examined, the de- tailed original description and distribution data provided by Scheerpeltz (1960) allow this species to be placed within Cheilocolpus; it has a combination of fea- tures typical for Cheilocolpus from the paramos: brachypterous habitus, entirely brownish-black body, rectangular head with transverse microsculpture and small convex eyes, paraocular punctures, and tergites III-V with posterior transverse basal carina]; Cheilocolpus speciosus (Bernhauer, 1917), comb. nov. ex. Quedi- us [1 syntype from FMNH examined (Fig. 7B)]; Cheilocolpus viridulus (Erichson, 1840), comb. nov. ex. Quedius [2 syntypes from MFNB examined (Fig. 7C)]. Furthermore, we propose the following new combination for the species Cyrtoquedius viridipennis (Fauvel, 1891), comb. nov. ex. Quedius. Although the type material was not examined, the original description and distribution data provided by Fauvel (1891) support placing this species within Cyrtoquedius, in accordance with the diagnosis by Brunke et al. (2016). Fauvel noted that it is closely related to Cy. labiatus (Erichson, 1840) but differs in coloration. He also described the elytra as having three rows of larger punctures, with the remain- ing punctures faint and vaguely marked, which are typical characteristics of this genus. This contrasts with Nitidocolpus and Cheilocolpus, where the elytra are evenly setose rather than mostly glabrous except for rows of macrosetae. ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 95 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Discussion The description of Chiquiticus gen. nov. and Nitidocolpus gen. nov. was neces- sitated due to the upcoming formal taxonomic embedding of the newly discov- ered lineages in Staphylininae in Reyes-Hernandez et al. (in prep.). Also, it is a step towards badly needed taxonomic clean-up of the polyphyletic large waste- basket genera Heterothops (Amblyopinina) and Quedius (Quediina), especially for the New World fauna, because here we based our genus descriptions solely on the species earlier described in those genera. These are eight species of Het- erothops (of which two are extinct) for Chiquiticus and eight species of Quedius for Nitidocolpus. Globally, many more non-related convergently similar species remain misplaced in both Heterothops and Quedius. In the American fauna, fol- lowing the establishment of Chiquiticus and Nitidocolpus, the number of mis- classified Heterothops species has significantly decreased. Moreover, in the course of this study we reclassified a number of the Neotropical Quedius spe- cies which superficially resemble Nitidocolpus but in fact, belong to the ambly- opinine genus Cheilocolpus or the cyrtoquediine genus Cyrtoquedius. It should be noted that in both Chiguiticus and Nitidocolpus there are also several new species to be described; as well as in the genus Cheilocolpus and other genera of the former “southern quediines”. Due to the above-mentioned pragmatic tax- onomic purposes of the current generic descriptions, here we considered solely the described species based on the type material or their original descriptions if they displayed enough data to adequately identify them as either new genus. Descriptions of all new species as well as the update of the taxonomy including lectotype designations, redescriptions, summary of the distribution, and bion- omics information on the earlier described species in both new and other in- volved genera will be provided in future revisionary work. Acknowledgements We thank Adam Brunke, Aslak Kappel Hansen, Josh Jenkins Shaw, Alfred New- ton and Qinghao Zhao, who contributed in one way or another to the completion of this work. We would like to thank all the curators listed in the Material and methods section, particularly Agnieszka Pierwola of the American Museum of Natural History, New York, who photographed type material of tChiquiticus cor- nelli. Images of C. arizonicus are provided by the Canadian National Collection of Insects, Arachnids, and Nematodes (CNC), ©His Majesty the King in Right of Canada, as represented by the Minister of Agriculture and Agri-Food, and licensed under the Open Government Licence — Canada. We extend our thanks to Stylianos Chatzimanolis, and an anonymous reviewer for their valuable com- ments, which improved the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. ZooKeys 1218: 81-98 (2024), DOI: 10.3897/zookeys.1218.135558 96 José L. Reyes-Hernandez & Alexey Solodovnikov: Reducing "Heterothops" and "Quedius" taxonomic wastebaskets Funding We thank Copenhagen University DATA+ program funded the PHYLORAMA project and the doctoral scholarship for the first author. Author contributions Both authors have contributed equally. Author ORCIDs José L. Reyes-Hernandez © https://orcid.org/0000-0002-4726-4439 Alexey Solodovnikov © https://orcid.org/0000-0003-2031-849X Data availability All of the data that support the findings of this study are available in the main text. 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