AnimaliaColeopteraHydrophilidaeGirónJennifer C.ShortAndrew Edward Z.The Acidocerinae (Coleoptera, Hydrophilidae): taxonomy, classification, and catalog of speciesZookeys18620211045123610.3897/zookeys.1045.63810 Helobata Bergroth, 1888Figs 1J, 2, 5, 11L, 33, 34 Helopeltis Horn, 1873: 137. Type species: Helopeltis larvalis Horn, 1873: 137; by monotypy. Helobata Bergroth, 1888: 221 – Replacement name for Helopeltis Horn, 1873. Helopeltina Cockerell, 1906: 240 – Replacement name for Helopeltis Horn, 1873. Type species: Helopeltis larvalis Horn, 1873: 137. Gender.

Feminine.

Type species.

Helopeltis larvalis Horn, 1873: 137; by monotypy.

Diagnosis.

Medium sized beetles, body length 4–7 mm. Body shape oval in dorsal view, dorsoventrally flattened, with dorsal outline nearly straight along medial third in lateral view (Fig. 33); surface even and granulate. From yellowish, orange brown to dark brown in coloration, usually with patterns along elytra, with different areas of head and pronotum darkened. Shape of head somewhat trapezoid (Fig. 11L). Anterior corners of frons extended laterally and posteriorly, emarginating anterior margin of eyes. Eyes of moderate size, somewhat oval, anteriorly deeply emarginated, not projected from outline of head. Clypeus somewhat pentagonal, laterally explanate, with anterior margin usually straight (Fig. 11L). Labrum concealed by clypeus (Fig. 11L). Mentum with surface variably sculptured, usually with oblique and transverse striae (Fig. 33C). Antennae with eight antennomeres, with cupule strongly asymmetric and oval in outline. Maxillary palps slender, slightly longer than greatest width of head; inner margin of maxillary palpomere 2 weakly and evenly curved, and outer margin weakly curved along apical third (Fig. 33C). Pronotum with surface of lateral areas flat. Elytra without sutural striae, with outer margins laterally explanate; serial punctures clearly aligned in longitudinal rows (Fig. 33A). Scutellar shield U-shaped. Surface of prosternum flat, to medially bulging, smooth to irregularly sculptured. Posterior elevation of mesoventrite only weakly bulging, with pair of lateral, longitudinal, low ridges; anapleural sutures nearly parallel along anterior section, separated anteriorly by distance slightly shorter than anterior margin of mesepisternum. Metaventrite uniformly covered by hydrofuge pubescence, with medial, narrow, and slightly carinate glabrous patch; posterolateral glabrous patches reduced. Protibiae with spines of anterior row short and semi erect; apical spurs of protibiae reduced, much shorter than protarsomere 1. Metafemora with tibial grooves moderately developed; hydrofuge pubescence covering 5/6 of anterior surface (Fig. 33C). Tarsomeres 1–4 ventrally densely covered by setae; metatarsomere 2 longer than 3 and 4 combined, 1 nearly as long as 3, and 5 nearly as long as 2–4 combined. Fifth abdominal ventrite apically emarginate, with fringe of flat and stout setae. Aedeagus divided (Fig. 34), parameres separated from each other for most of their lengths; median lobe divided in dorsal and ventral plates; dorsal plate usually strongly sclerotized; ventral plate bilaterally bifurcated, forming thick lateral lobes along apical region; basal piece nearly 0.2 × the length of parameres, always noticeable; gonopore not clearly visible.

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.

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10.3897/zookeys.1045.63810.figure34

Aedeagi of Helobata spp. A, BH. pantaneira (from Clarkson et al. 2016): A dorsal view B lateral view C, DH. quatipuru (Clarkson and Almeida 2018) A dorsal view D lateral view EH. sp. (Ecuador), dorsal view. Scale bars: 0.5 mm.

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Differential diagnosis.

Helobata is one of the most conspicuous genera of acidocerines, especially in the New World. The flattened and broadly explanate body shape and concealed labrum, accompanied by granulose surface, long and slender maxillary palps and well-defined elytral serial punctures, are quite unique in the subfamily. The only genus that shares some of these features is Helopeltarium, except that the latter has short maxillary palps, smooth surface and lacks serial punctures along the elytra. The configuration of the aedeagus (Fig. 34), in particular the thickness of the lateral lobes of the ventral plate of the median lobe, is also unique among acidocerines.

Distribution.

Nearctic: United States (California, Florida, Louisiana, Mississippi, North Carolina, South Carolina, Texas, Virginia). Neotropical: Argentina, Bolivia, Brazil (Amazonas, Ceará, Mato Grosso, Mato Grosso do Sul, Pará, Rio de Janeiro, Roraima), Cuba, Guatemala, Mexico, Paraguay, Suriname, Venezuela; Fig. 5.

Natural history.

Species of Helobata occur primarily in open habitats with abundant vegetation. According to Clarkson et al. (2016), specimens of Helobata are uncommonly encountered and occur in marshes, swamps, and ponds, most often in small numbers, although they are rarely found in modest amounts (dozens of individuals; Short, pers. obs.). According to Archangelsky (1997), they can be found in slow moving creeks or rivers, living among the littoral vegetation or on floating plants. They are attracted to lights. Females have been observed carrying their egg cases attached to the ventral side of their abdominal ventrites (Archangelsky 1997).

Larvae.

The larva (first instar) and egg case are only known for Helobata larvalis; these immature stages were described by Spangler and Cross (1972). A differential diagnosis of the first instar larva was provided by Fikáček (2003).

Taxonomic history.

This genus was described by Horn (1873) under the name Helopeltis, which was preoccupied by Helopeltis Signoret, 1858 (Hemiptera). Bergroth (1888) proposed the name Helobata as a replacement name for Helopeltis Horn, whereas Cockerell (1906a) proposed the name Helopeltina. Helobata has priority, so it is the currently valid name for the genus, which was revised by Fernández and Bachmann (1987).

Remarks.

There are 13 species of Helobata described to date. The type species, Helobata larvalis (Horn), has generally been known under the name Helobata striata (originally published as Hydrophilus striatus Brullé, 1841: 58, which is a primary homonym of Hydrophilus striatus Say, 1825 [now Berosus striatus (Say)]; therefore unavailable. The name Helobata larvalis (Horn) was then reinstated by Hansen (1991: 293). Photos of a syntype of Helopeltis larvalis (Horn) are available at https://mczbase.mcz.harvard.edu/guid/MCZ:Ent:101 (accessed 9 January 2021). The external morphology of members of Helobata is very homogeneous. Some variation can be observed in the shape of the clypeus (e.g., Fernández 1987; Clarkson et al. 2016). Helobata is the only Neotropical genus truly widespread in the New World, as it ranges from southeastern North America, all the way to Argentina and Southern Brazil.

Species examined.

Helobata cuivaum García (paratype), H. larvalis (Horn), and H. lilianae García (paratype).

Selected references.

Horn 1873: original description of the genus and the type species; Spangler and Cross 1972: description of egg case and first instar larva; Fernández and Bachmann 1987: review of the genus, description of four new species from Argentina, Brazil and Paraguay; García 2000: three new species from Venezuela; Makhan 2007: two new species from Suriname; Clarkson et al. 2016: two new species from Brazil, review and new country records of Brazilian species; Clarkson and Almeida 2018: new records from Brazil; Short et al. 2021: phylogenetic placement.

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.

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10.3897/zookeys.1045.63810.figure2

Phylogeny of the Acidocerinae simplified from Short et al. (2021), indicating the distribution, preferred habitat, and currently described number of species for each genus. For habitat, filled black circles indicate that at least some species of the genus are commonly found in this habitat; light grey circles indicate the genus has been found in this habitat, but is rare or not typical for the group; white circles indicate no species have been recorded for the genus in this habitat.

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10.3897/zookeys.1045.63810.figure5

Known distribution of genera of Acidocerinae: Crucisternum, Ephydrolithus, Globulosis, Helobata, Helochares, Helopeltarium, Katasophistes, and Nanosaphes.

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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.

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ShortAEZGirónJCToussaintEFA (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.12467ClarksonBAlmeidaLM (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.4126.4.6ArchangelskyM (1997) Studies on the biology, ecology, and systematics of the immature stages of New World Hydrophiloidea (Coleoptera: Staphyliniformia).Bulletin of the Ohio Biological Survey12(1): 1207.SpanglerPJCrossJL (1972) A description of the egg case and larva of the water scavenger beetle Helobata striata (Coleoptera: Hydrophilidae).Proceedings of the Biological Society of Washington85: 413418. https://biodiversitylibrary.org/page/34559933FikáčekM (2003) Commented review of immature stages of World Hydrophiloidea (Coleoptera: Staphyliniformia). Bachelor Seminar Work, Charles University, Prague.HornGH (1873) Revision of the genera and species of the tribe Hydrobiini.Proceedings of the American Philosophical Society13(90): 118137. https://www.jstor.org/stable/981607BergrothE (1888) Fåhraea nov. gen. Hydrophilidarum. Deutsche entomologische Zeitschrift 32: e221. https://biodiversitylibrary.org/page/33177095CockerellTDA (1906a) Preoccupied Generic names of Coleoptera.Entomological News17: 240244. https://biodiversitylibrary.org/page/2562995FernándezLABachmannAO (1987) Revisión del género Helobata Bergroth (Coleoptera: Hydrophilidae).Revista de la Sociedad Entomológica Argentina44(2): 149159. https://www.biotaxa.org/RSEA/article/view/36822HansenM (1991) The hydrophiloid beetles. Phylogeny, classification and a revision of the genera (Coleoptera: Hydrophilidae).Biologiske Skrifter40: 1367.MakhanD (2007) Helobata soesilae sp. nov. and Helobata aschnakiranae sp. nov. from Suriname (Coleoptera: Hydrophilidae).Calodema Supplementary Paper14: 13. https://www.researchgate.net/publication/321107032_Helobata_soesilae_sp_nov_and_Helobata_aschnakiranae_sp_nov_from_Suriname_Coleoptera_Hydrophilidae