AnimaliaColeopteraHydrophilidaeGirónJennifer C.ShortAndrew Edward Z.The Acidocerinae (Coleoptera, Hydrophilidae): taxonomy, classification, and catalog of speciesZookeys18620211045123610.3897/zookeys.1045.63810 Quadriops Hansen, 1999Figs 1P, 2, 6, 11C, 48, 49A–D Quadriops Hansen, 1999a: 131.Gender.

Masculine.

Type species.

Quadriops depressus Hansen, 1999a: 136; by original designation.

Diagnosis.

Small to very small beetles, body length 1.6–2.6 mm. Body shape oval in dorsal view; moderate to strongly convex in lateral view, dorsal outline evenly convex or nearly straight along median region (Fig. 48). Color orange brown to dark brown, uniform along body regions; ground punctation shallow to moderately marked (Fig. 48). Shape of head somewhat rectangular. Frons lateral and posteriorly expanded, forming canthus completely dividing eyes in dorsal and ventral portions (Fig. 11C). Eyes very small in dorsal view. Clypeus laterally expanded in front and around outer margin of eyes; anterior margin of clypeus straight (Fig. 11C). Labrum partly exposed. Mentum rather smooth and medially depressed; median anterior depression marked by a transverse carina (Fig. 48C, F). Antennae with nine antennomeres, cupule slightly asymmetric with rounded outline. Maxillary palps rather short and stout, nearly half as long as width of head; maxillary palpomere 4 slightly longer than palpomere 3; inner margin of maxillary palpomere 2 straight to convex, outer margin strongly curved along apical 2/3. Elytra without sutural striae, with punctures either irregularly distributed or forming well defined longitudinal rows; elytra narrowly explanate anteriorly, explanation gradually broader towards apex (Fig. 48). Surface of prosternum flat. Posterior elevation of mesoventrite, usually with well-defined transverse ridge, seldom with acute tooth; anapleural sutures concave, separated at anterior margin by distance nearly 0.7 × width of anterior margin of mesepisternum. Metaventrite usually uniformly densely pubescent, sometimes with reduced posteromedian glabrous patch. Protibiae with spines of anterior row hair-like, semi erect, relatively long, and thick; apical spurs of protibia moderately stout, reaching apex of protarsomere 3. All tarsomeres with thick hair-like spines on ventral face of tarsomeres 2–4; metatarsomeres 1–4 similar in length, 5 nearly as long as 3 and 4 combined. Metafemora with tibial grooves moderately developed; anterior surface of metafemora mostly glabrous, with few very scattered small setae (Fig. 48C, F). Fifth abdominal ventrite apically rounded and without fringe of stout setae. Aedeagus trilobed (Fig. 49A–D), with basal piece about half length of parameres; median lobe wider than base of each paramere, with narrow, triangular, longitudinal sclerite, usually extending along apical third; parameres as long as, to longer than median lobe, and nearly half as wide; gonopore situated preapically; basal piece with lateral margins straight to sinuate, apically slightly diverging.

10.3897/zookeys.1045.63810.figure48

Habitus of Quadriops spp. A–CQ. acroreius: A dorsal habitus B lateral habitus C ventral habitus D–FQ. clusia: D dorsal habitus E lateral habitus F ventral habitus. Scale bars: 1 mm.

https://binary.pensoft.net/fig/557447
10.3897/zookeys.1045.63810.figure49

Aedeagi of Quadriops, Radicitus and Sindolus spp. AQ. clusiaBQ. depressusCQ. reticulatusDQ. similarisES. sp. (Venezuela) F. S. sp. (Venezuela) G, HR. ayacucho: G dorsal view H lateral view I, JR. cf. granitum (Suriname): I dorsal view J lateral view K, LR. surinamensis: K dorsal view L lateral view. Scale bars: 0.1 mm (A–D); 0.5 mm (E–L).

https://binary.pensoft.net/fig/557448
Differential diagnosis.

Quadriops is the only known acidocerine with fully divided eyes. Species with uniformly distributed punctures along the elytra may resemble Globulosis, but the moderate punctation of Quadriops is very evident (punctation only shallowly marked in Globulosis; Fig. 32). Some species of Tobochares have nearly divided eyes, and lack impressed striae along the elytra (emarginatus species group, Girón and Short 2021a), resembling species of Quadriops with uniformly distributed punctures along the elytra, but they differ in the shape of the posterior elevation of the mesoventrite (sharply elevated as a tooth or a blunt transverse carina in Quadriops, medially bulging in T. canthus Kohlenberg & Short).

Distribution.

Neotropical: Brazil (Amazonas), Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Suriname, Venezuela; Fig. 6.

Natural history.

Specimens have been caught using flight intercept traps, many long series have been collected on decaying Clusia fruits, which can be somewhat used as bait (Fig. 10). Additional specimens have been collected in rotten logs, sap flows on freshly cut trees, and in the refuse piles of leafcutter ants (Girón and Short 2017).

Larvae.

The immature stages of Quadriops remain unknown.

Taxonomic history.

Hansen (1999a) described the genus with five species, differentiated mostly by the presence and degree of impression of reticulation on the head and clypeus. When he originally described it, Hansen (1999a) was unsure of the taxonomic affinity of the genus, as the morphology of the lineage was somewhat unusual. He placed it in the Acidocerina (now Acidocerinae) almost by default as it shared no characters in common with other lineages, but ultimately, he was correct as this placement as verified by Short et al. (2021). García (2000b) described an additional species from Venezuela. The genus was revised by Girón and Short (2017): two species were synonymized with Quadriops depressus Hansen; two new species were described.

Remarks.

Quadriops is the only fully terrestrial genus of Acidocerinae. There are six described species within the genus.

Species examined.

Quadriops acroreius Girón & Short (holotype and paratype), Q. clusia Girón & Short (holotype, paratypes and additional specimens), Q. dentatus Hansen (holotype and additional specimens), Q. depressus Hansen (holotype and additional specimens), Q. reticulatus Hansen (holotype and additional specimens), Q. similaris Hansen (holotype and additional specimens).

Selected references.

Hansen 1999a: original description; García 2000b: description of one additional species from Venezuela; Girón and Short 2017: generic revision including two synonymies and two new species; 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.

https://binary.pensoft.net/fig/557400
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.

https://binary.pensoft.net/fig/557401
10.3897/zookeys.1045.63810.figure6

Known distribution of genera of Acidocerinae: Novochares, Peltochares, Primocerus, Quadriops, Radicitus, Sindolus, Tobochares, and Troglochares.

https://binary.pensoft.net/fig/557405
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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10.3897/zookeys.1045.63810.figure32

Habitus of Globulosis flavusA dorsal habitus B lateral habitus C ventral habitus. Scale bar: 1 mm.

https://binary.pensoft.net/fig/557431
10.3897/zookeys.1045.63810.figure10

Examples of terrestrial habitat for AcidocerinaeA, B Rotting Clusia fruit, showing Quadriops clusia crawling on the surface (Suriname: SR17-0322-03A) C collecting specimens by submerging rotting fruits in pans of water and waiting for the beetles to float to the surface.

https://binary.pensoft.net/fig/557409
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.12467GirónJCShortAEZ (2021a) New subgenera, new species and new records of the Neotropical water scavenger beetle genus Tobochares Short & García, 2007 (Coleoptera: Hydrophilidae: Acidocerinae).ZooKeys1019: 93140. https://doi.org/10.3897/zookeys.1019.59881GirónJCShortAEZ (2017) Revision of the Neotropical water scavenger beetle genus Quadriops Hansen, 1999 (Coleoptera: Hydrophilidae: Acidocerinae).ZooKeys705: 115141. https://doi.org/10.3897/zookeys.705.19815HansenM (1999a) Fifteen new genera of Hydrophilidae (Coleoptera), with remarks on the generic classification of the family.Insect Systematics and Evolution30(2): 121172. https://doi.org/10.1163/187631200X00228GarcíaM (2000b) Una nueva especie de Quadriops Hansen, 1999 (Coleoptera: Hydrophilidae: Hydrophilinae) de Venezuela.Boletín del Centro de Investigaciones Biológicas34(1): 5965. http://produccioncientificaluz.org/index.php/boletin/article/view/219