AnimaliaColeopteraHydrophilidaeGirónJennifer C.ShortAndrew Edward Z.The Acidocerinae (Coleoptera, Hydrophilidae): taxonomy, classification, and catalog of speciesZookeys18620211045123610.3897/zookeys.1045.63810 Acidocerinae Zaitzev, 1908 Acidocerini Zaitzev, 1908: 353, as subfamily. Acidocerina as subtribe Acidocerina [of tribe Hydrophilini, subfamily Hydrophilinae] in Hansen (1991: 282; 1999b: 155). Acidocerina as tribe [of subfamily Hydrophilinae] in Short and Fikáček (2011: 85). Acidocerina as subfamily in Short and Fikáček (2013: 741). Helopeltini Horn, 1873: 118; synonymized by Hansen (1991: 282); unavailable: generic name is preoccupied (ICZN 1999, Code Art. 39). Type genus: Helopeltis Horn, 1873: 137 [synonym of Helobata Bergroth, 1888: 221]. Helocharae d’Orchymont, 1919c: 147; described as subtribe, synonymized by Hansen (1991: 282). Type genus: Helochares Mulsant, 1844a: 197. Horelophopsinae Hansen, 1997: 108. Type genus: Horelophopsis Hansen, 1997: 109; synonymized by Short and Fikáček (2013: 15, in table, discussed along the text). Globulina García, 2001: 153; emended to Globulosina by Short and Hebauer (2006: 338); synonymized with tribe Acidocerini by Short and Fikáček 2011: 84. Type genus: Globulosis García, 2001: 153. Type genus.

Acidocerus Klug, 1855: 649.

Diagnosis.

Body length 1.2–14.0 mm. Body shape oval in dorsal view, dorsoventrally flattened, or weakly to strongly convex in lateral view (Fig. 1); surface even (without elevations or depressions), granulate (e.g., Figs 17, 33) or smooth on head and pronotum. From yellowish to dark brown in coloration (Fig. 1), usually uniform, sometimes different regions of body colored differently. Shape of head variable (trapezoid, subquadrate, round; Fig. 11E–L). Anterior corners of frons sometimes extended posteriorly forming canthus and emarginating anterior margin of eyes (e.g., Tobochares, Helobata; e.g., Fig. 11B, C). Eyes varying in size, shape, degree of emargination, and degree of projection from outline of head (Fig. 11E–L); absent only in cavernicolous genus Troglochares Spangler, 1981a. Clypeus variable in shape (rectangular to trapezoid; Fig. 11E–L), with anterior margin from straight to mesally emarginate. Labrum usually exposed; concealed by clypeus in Helobata (Fig. 11L) and Helopeltarium (Fig. 1H). Mentum usually wider than long, with strong median anterior depression, may be limited by low transverse carina (Fig. 12A–C); surface of mentum with variable sculpture, ranging from smooth (Fig. 12A) to roughly punctate or obliquely striate (Fig. 12B). Antennae with eight or nine antennomeres (Fig. 12D, E), with cupule varying in symmetry and shape. Maxillary palps curved inward, ranging from very short (nearly half width of the head; e.g., Quadriops reticulatus, Fig. 12C) and stout, to very long and slender (nearly twice the width of the head; e.g., Peltochares, Fig. 11K). Pronotum evenly convex, usually with systematic punctures forming paired anterolateral semicircles and paired short posterolateral transverse bands. Elytra with or without sutural striae, with outer margins simple, slightly flared, or laterally explanate; elytral punctation variable (Fig. 13). Hind wings usually well developed (Fig. 15A–F), seldom reduced along apical region (Fig. 15G). Surface of prosternum flat (e.g., Fig. 14A, B), convex or rarely medially carinate (e.g., Crucisternum; Fig. 29C), with anterior margin straight or anteriorly projected. Posterior elevation of mesoventrite either only weakly bulging or with transverse (e.g., Fig. 14E, G) or longitudinal ridge (e.g., Fig. 14D, F); with strongly produced, anteriorly pointed and longitudinally carinate transverse ridge in Crucisternum (Fig. 14C). Anapleural sutures variable in shape and orientation. Metaventrite rather uniformly covered by hydrofuge pubescence (e.g., Fig. 14E), sometimes with posteromesal glabrous patch (e.g., Fig. 14D, F, G), sometimes also with posterolateral glabrous patches (e.g., Fig. 14D). Protibiae with anterior row of spines varying in shape and development; apical spurs of protibiae varying in development. Metafemora with tibial grooves of varying development; hydrofuge pubescence on anterior surface of metafemora absent, reduced to only basal or dorsal patch, or increasingly covering most of surface. Tarsomeres 5-5-5; tarsomeres variable in size, proportions, and dorsal and ventral coverage. Abdomen with five pubescent ventrites, density of setae ranging from sparse to very dense. Fifth abdominal ventrite with apex either rounded (Fig. 15I), truncate (Fig. 15J), or emarginate (Fig. 15H); apex with or without fringe of flat and stout setae. Aedeagus usually symmetrical (Fig. 16), with basal piece varying in size from longer than parameres (e.g., Primocerus, Fig. 47; Batochares, Fig. 22D), to reduced and virtually absent (e.g., Peltochares, Fig. 45); parameres highly variable in shape, either slender and only connected to each other at base of ventral surface (e.g., Fig. 16A–D, G, H), or fused together forming tube-like structure (e.g., Fig. 16E, F); apex of parameres either simple, or bifurcated and modified with hooks and spines (e.g., Fig. 16C, D); median lobe either simple or with dorsal and ventral lobes, with well-developed lateral basal apodemes; further modifications (longitudinal divisions, presence of internal hooks and spines, development of gonopore) widespread.

Differential diagnosis.

Acidocerines can be generally recognized by their oval and moderately convex body shapes with slender maxillary palps and uniformly slender tibiae (usually strongly convex and sometimes rounded in Cylominae and Sphaeridiinae, with short and stout maxillary palps and stout to apically broadened tibiae). The maxillary palps are always curved inwards in Acidocerinae (maxillary palpomere 2 with inner margin straight to concave; Fig. 12F–J), with palpomeres 2–4 similar in length and proportions (curved outwards, zig-zag oriented, or with shorter palpomere 3 in most Enochrinae and Chaetarthriinae). In addition, Acidocerines always bear five tarsomeres on the meso- and metatarsi (four in some enochrines).

Selected references.

Hansen 1991: diagnosis of the group (at the time as a subtribe, and including some genera now placed in the subfamily Enochrinae), list of genera and subgenera with synonyms, key to genera, and description of each genus (8 out of the 23 recognized in this paper). Hansen 1999b: catalog with full list of species at the time (nearly 300), synonyms and references. Short and Fikáček 2013: Acidocerinae as a subfamily excluding enochrine genera, with Horelophopsinae as synonym, list of genera, general diagnosis. Short et al. (2021): molecular phylogeny and biogeography of the subfamily, groups of genera.

Remarks.

The subfamily Acidocerinae is a group with many contrasts. It includes some of the largest as well as smallest hydrophilids; some genera are either strikingly different from, or extremely similar to others; the external morphology of some genera is extremely uniform and species can only be recognized by characters of the male genitalia, or so variable that is difficult to diagnose the group as a unit; at the species level, the distributions can be very narrow and restricted to one or a few fairly close localities, or very broadly widespread across several continents. There is a trend for species living in the same kind of habitats to have certain shared morphological features. For example, species that live in aquatic habitats tend to have slender and relatively long maxillary palps and metafemora mostly covered by hydrofuge pubescence, whereas species living in hygropetric habitats tend to have shorter and stouter maxillary palps and reduced or absent coverage of hydrofuge pubescence on the metafemora.

10.3897/zookeys.1045.63810.figure1

Variation across Acidocerinae, dorsal and lateral views AColossochares ellipticusBPeltochares sp. CPeltochares conspicuusDAulonochares tubulusEHelochares sp. FHelochares tristisGNovochares sp. HHelopeltarium ferrugineumIBatochares sp. JHelobata larvalisKRadicitus sp. LNanosaphes tricolorMAgraphydrus cf. attenuatusNTobochares luteomargoOTobochares sulcatusPQuadriops similarisQCrucisternum ouboteriRPrimocerus neutrumSAgraphydrus coomaniTAgraphydrus sp. UGlobulosis flavusVCrephelochares nitescens.

https://binary.pensoft.net/fig/557400
10.3897/zookeys.1045.63810.figure17

Habitus of Acidocerus aphodioidesA dorsal habitus B lateral habitus C head. Scale bar: 1 mm.

https://binary.pensoft.net/fig/557416
10.3897/zookeys.1045.63810.figure33

Habitus of Helobata spp. A–CH. larvalis: A dorsal habitus B lateral habitus C ventral habitus DH. quatipuru (from Clarkson and Almeida 2018) EH. amazonensis (from Clarkson and Almeida 2018) FH. pantaneira (from Clarkson et al. 2016). Scale bars: 1 mm.

https://binary.pensoft.net/fig/557432
10.3897/zookeys.1045.63810.figure11

Head of miscellaneous AcidocerinaeA–D anterolateral view: ATobochares luteomargo with white arrow pointing to straight anterior margin of eye BTobochares emarginatus with white arrow pointing to canthus emarginating anterior margin of eye CQuadriops politus with white arrow pointing to canthus fully dividing the eye in dorsal and ventral faces DBatochares sp. black arrow pointing to transverse carina on labrum E–L dorsal view of head: EBatochares sp. FHelochares tristisGCrephelochares nitescens, HChasmogenus australis with black arrow pointing to preclypeal membrane IColossochares ellipticusJAulonochares tubulusKPeltochares conspicuusLHelobata larvalis.

https://binary.pensoft.net/fig/557410
10.3897/zookeys.1045.63810.figure12

Head structures A–C scanning electron micrographs of ventral view of head: ATobochares pallidus with smooth mentum and white arrow pointing to transverse carina limiting posterior margin of antero-medial depression BNanosaphes tricolor with top white arrow pointing to oblique crenulations of mentum, mid white arrow pointing to flat and smooth anterior surface of submentum, and bottom white arrow pointing to concave posterior surface of submentum CQuadriops reticulatus with white arrow pointing to antero-medial depression of mentum D, E light micrographs of antenna: DAulonochares tubulus (9 antennomeres) EChasmogenus cremnobates (8 antennomeres) F–J light micrographs of maxillary palps: FQuadriops reticulatusGAgraphydrus insidiatorHHelochares sp. IHelochares lividusJAulonochares tubulus. Scale bars: 100 μm (A–C)

https://binary.pensoft.net/fig/557411
10.3897/zookeys.1045.63810.figure13

Elytral punctation ATobochares communis with red arrow pointing to systematic puncture BTobochares sipaliwini with red arrow pointing to systematic puncture, white arrow pointing to serial puncture, and black arrow pointing to ground/interserial puncture CTobochares striatus with red arrow pointing to systematic puncture, white arrow pointing to serial puncture, and black arrow pointing to ground/interserial puncture DTobochares communis elytron with all kinds of punctures similar in size and degree of impression, seemingly evenly distributed (to longitudinally aligned) EQuadriops similaris with serial punctures longitudinally aligned FPrimocerus maipure with sutural stria GTobochares striatus with impressed serial striae. Scale bars: 100 μm (A); 200 μm (B, C); 500 μm (D–G).

https://binary.pensoft.net/fig/557412
10.3897/zookeys.1045.63810.figure15

Hind wing and abdominal ventrite 5 A–G hind wings: AColossochares ellipticusBPrimocerus gigasCHelobata larvalisDCrucisternum ouboteriETobochares sipaliwiniFQuadriops similarisGTobochares micropsH–J abdominal ventrite 5: HAulonochares tubulusIPrimocerus neutrumJEphydrolithus hamadae. Scale bars: 1 cm (A); 3 mm (B, C); 1 mm (D–F); 0.5 mm (G).

https://binary.pensoft.net/fig/557414
10.3897/zookeys.1045.63810.figure14

Scanning electron micrographs of thorax in ventral view A, B prosternum: ATobochares striatus with white arrow pointing to anterior projection BQuadriops reticulatus with white arrow pointing to anterior projection C–G mesoventrite and metaventrite: CCrucisternum ouboteri with white arrows pointing to anteriorly pointed transverse ridge and longitudinal carina of mesoventrite, metaventrite with median glabrous patch DNanosaphes tricolor with black arrow pointing to longitudinal carina along mesoventrite and white arrows pointing to median and postero-lateral glabrous patches of metaventrite EQuadriops reticulatus with black arrow pointing to transverse carina across mesoventrite and metaventrite uniformly pubescent FTobochares communis with black arrow pointing to longitudinal carina along mesoventrite and white arrow pointing to narrow postero-medial glabrous patch on metaventrite GTobochares kasikasima with black arrow pointing to transverse elevation across mesoventrite and white arrow pointing to broad postero-medial glabrous patch on metaventrite. Scale bars: 100 μm.

https://binary.pensoft.net/fig/557413
10.3897/zookeys.1045.63810.figure29

Habitus of Crucisternum ouboteriA dorsal habitus B lateral habitus C ventral habitus. Scale bar: 1 mm.

https://binary.pensoft.net/fig/557428
10.3897/zookeys.1045.63810.figure16

Aedeagi A–E trilobed: A schematic BChasmogenus schmitsC, D spiked: C schematic DPeltochares foveicollisE, F tubular: E schematic FHelochares politusG, H divided: G schematic HNovochares pallipes.

https://binary.pensoft.net/fig/557415
10.3897/zookeys.1045.63810.figure47

Aedeagi of Primocerus spp. AP. neutrumB, CP. maipure: B dorsal view C lateral view D, EP. pijiguaense: D dorsal view E lateral view FP. gigasGP. petilusHP. striatolatusIP. cuspidis. Scale bars: 0.25 mm.

https://binary.pensoft.net/fig/557446
10.3897/zookeys.1045.63810.figure22

Aedeagi of Aulonochares and Batochares spp. AA. tubulusBA. novoairensisCA. lingulatusDB. sp. Scale bars: 0.5 mm.

https://binary.pensoft.net/fig/557421
10.3897/zookeys.1045.63810.figure45

Aedeagi of Peltochares spp. AP. conspicuusBP. foveicollisCP. sp. (Australia) DP. sp. (Tanzania;). Scale bars: 1 mm.

https://binary.pensoft.net/fig/557444
HansenM (1991) The hydrophiloid beetles. Phylogeny, classification and a revision of the genera (Coleoptera: Hydrophilidae).Biologiske Skrifter40: 1367.HansenM (1999b) World catalogue of insects (Volume 2). Hydrophiloidea (s. str.) (Coleoptera).Apollo Books, Stenstrup, 414 pp.ShortAEZFikáčekM (2011) World catalogue of the Hydrophiloidea (Coleoptera): additions and corrections II (2006–2010).Acta Entomologica Musei Nationalis Pragae51(1): 83122. http://www.aemnp.eu/PDF/51_1/51_1_83.pdfShortAEZFikáčekM (2013) Molecular phylogeny, evolution and classification of the Hydrophilidae (Coleoptera).Systematic Entomology38(4): 723752. https://doi.org/10.1111/syen.12024ShortAEZHebauerF (2006) World Catalogue of Hydrophiloidea – additions and corrections, 1 (1999–2005) (Coleoptera).Koleopterologische Rundschau76: 315359. http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.614.3424andrep=rep1andtype=pdfClarksonBAlmeidaLM (2018) On Brazilian Helobata Bergroth, 1888 (Coleoptera: Hydrophilidae) II: new distribution data. Papeis Avulsos de Zoologia 58 (e20185835): 1–6. https://doi.org/10.11606/1807-0205/2018.58.35ClarksonBSantosADFerreira-JrN (2016) On Brazilian Helobata Bergroth, 1888 (Coleoptera: Hydrophilidae): description of two new species, new records, and key to species.Zootaxa4126(4): 548562. https://doi.org/10.11646/zootaxa.4126.4.6ShortAEZGirónJCToussaintEFA (2021) Evolution and biogeography of acidocerine water scavenger beetles (Coleoptera: Hydrophilidae) shaped by Gondwanan vicariance and Cenozoic isolation of South America.Systematic Entomology46(2): 380395. https://doi.org/10.1111/syen.12467