Liocanthydrus clayae ( J. Balfour-Browne, 1969 ) J. Balfour-Browne, 1969

Baca, Stephen M., Gustafson, Grey T., Toledo, Mario & Miller, Kelly B., 2014, Revision of the Neotropical burrowing water beetle genus Liocanthydrus Guignot (Coleoptera: Noteridae: Noterinae: Noterini) with the description of two new species, Zootaxa 3793 (2), pp. 231-246 : 241-243

Liocanthydrus clayae ( J. Balfour-Browne, 1969) n. comb.

Siolius clayae J. Balfour-Browne (1969: 5) (orig. descr.); Nilsson (2005: 124) (cat.); Nilsson (2011: 31) (cat.).

Type locality. Guyana, Kanuku Mountains, Rupununi.

Type material examined. Holotype (male): “ BRIT. GUIANA:/Kanuku Mts./Rupununi.// 21.ii.1961. T. Clay” [printed; “ BRIT. GUIANA:” with green underline], “Debris edge of/forest creek.” [printed], “Brit. Mus./1961– 188.” [printed], “ Siolius / clayae Type!/J. Balfour-Browne det./ VII.1967 ” [handwritten except for “J. Balfour- Browne det./196” (printed)], “ Type [red bordered, circular NHM label]” ( NHM). Paratypes: Same data as holotype except for “ Siolius / clayae P’type!/J. Balfour-Browne det./1967 [handwritten except for “J. Balfour-Browne det./ 196” (printed)], and “Para-/ type ” [circular, yellow bordered NHM label; printed] (2 females exs., NHM, FSCA); “ BRIT. GUIANA:/Moco-Moco/Kanuku Mts./ 21.ii.1961. T. Clay.” [printed; “ BRIT. GUIANA:” with green underline], “At light edge/of forest.” [printed], “ Siolius / clayae P’type/J. Balfour-Browne det./1967” [handwritten except for “J. Balfour-Browne det./196” (printed)] (1 female ex., FSCA). Note: All label descriptions are included in brackets.

Other material examined. Guyana: Lethem, 25 km SE, Kumu Falls, 3°15.9'N, 59°43.6'W, 4.iv.1994, leg. P.J. Spangler, collection #16, log jam in stream (8 exs., USNM); same data except collection #14, side pool at base of falls (209 exs., USNM); same data except 5.iv.1994, collection #18, from root mats trailing between boulders in cascade (111 exs., USNM); same data except collection #25, 28.iv.1995, leg. P.J. Spangler & S.A. Perry (81 exs., USNM); same data except 30.iv.1995, collection #28, leaf packs in stream near falls (130 exs., USNM); Lethem, 30 km SE, 3°18.2'N, 59°39'W, Moco Moco River, collection #26, 29.iv.1995, P.J. Spangler & S.A. Perry (35 exs., SEMC); Suriname: Para District, 05°27'N, 55°14'W, Zanerijsavanne, 1st trib. of Colakreek, 15.ix.1969, leg. N. Neiser, (SN 146) (1 ex., NHMW); Saramacca District, 05°23'N, 55°21'W, 4th trib. of Troelikreek, 12.i.1970, leg N. Nieser, (SN 289) (1 ex., NHMW). Venezuela: Amazonas State: Sumariapo, 18 km N, Rio Cuima, 21.i.1989, netted from leaves and roots, leg. P.J. Spangler, R.A. Faitoute & C.B. Barr (104 exs., USNM); 5°36.250'N, 67°34.955'W, 96 m, just S. of Puerto Ayacucho, HG-vapor light, 4.i.2006, AS-06-010, leg. A.E.Z. Short (1 ex., SEMC); 5°20.497'N, 67°45.358'W, 72 m, ca. 12 km N. Samariapo, small stream at road crossing, 6.i.2006, AS-06- 0 13, leg. A.E.Z. Short (1 ex., SEMC); 5°48.414'N, 67°26.313'W, 80 m, near Iboruwa, “Tobogancito”, 13.i.2009, leg. K.B. Miller, VZ09-0113-02E, leaf choked detrital pools in forest (11 exs., SEMC); same data except leg. A. Short, shallow backwater of streamlet over bare rock, VZ09-0113-02A (8 exs., SEMC); same data except VZ09- 0113-02C, river margin (7 exs., SEMC); 5°03.707'N, 67°46.768'W, 92 m, River near Orinco/Spipapo confl., 15.i.2009, leg. Short, Miller, Garcia, Camacho, & Joly, VZ09-0115- 01X (1 ex., SEMC). Apure State: 6°30.900'N, 67°32.604'W, 68 m, Btw Orinco & Cinaruco Rivers, 17.i.2009, leg. Short, Miller, & Camacho, VZ09-0117- 01X, morichal (40 exs., SEMC). Bolivar State: 6°35.617'N, 66°49238'W, 60 m, Los Pijiguaos, 15.ix.2007, HG-vapor light at morichal outcrop, AS-07-014, leg. A.E.Z. Short (1 ex., SEMC); 5°2.623'N, 61°6.083, 885 m, S. San Francisco, 3.viii.2008, AS-08-068, leg. A. Short & M. Garcia, morichal stream/marsh (7 exs., SEMC); same data except 1 km S. San Francisco (21 exs., SEMC); 6°13'4.6'N, 67°14'26.4"W, 60 m, ca. 25 km E. El Burro, 7.viii.2008, leg. A. Short, M. Garcia, & L. Joly, AS-08-077, rocky morichal (24 exs., SEMC); same data except 12.i.2009, leg. Short et al., VZ09-0113- 01X, (4 exs., SEMC); 6°35.617'N, 66°49.238'W, 80 m, Los pijiguaos, outcrop/morichal, 12.i.2009, leg Short, Garcia, Camacho, Miller, & Joly, VZ09-0112-01A, morichal margin (39 exs SEMC); same data except VZ09-0112001B, marginal algae (1 ex., SEMC); same data except VZ09-0112-01D, isolated forest pool in streambed (6 exs., SEMC); same data except 8.vii.2010, leg Short, Tellez, & Arias, small stream on outcrop VZ10-0708-01B (2 exs., SEMC); same data except at rock out crop, seep scrubbing, VZ10-0708-01C (1 ex., SEMC); 6.58694°N, 67.02912°W, Rio Caripito, nr. Rio Orinoso, 12.i.2009, leg. Short & Miller, VZ09-0112-02A, river margin (11 exs., SEMC); same data except VZ09-0112-02B, detrital pool (3 exs., SEMC); 04°37.362'N, 61°05.679'W, 876 m, Gran Sabana, N. Santa Elena, Rio Guara at Rt. 10, 17.vii.2010, leg. Short, Tellez, & Arias, marshy area, VZ10-0717-02A (4 exs., SEMC); 04°41.878'N, 61°04.246, 815 m, Gran Sabana, N. Santa Elena, marshy area along Rt. 10, leg. Short, Tellez, & Camacho, 17.vii.2010, VZ10-0717-03A (5 exs., SEMC). Guárico State: Calabozo, Biológica, 23.vi.1963, leg. C. Bordon (1 ex., FSCA); Calabozo, 16 km S, Rio Orituco, 8.iii.1986, leg. P.J. Spangler, collection #33 (13 exs., USNM); 8°8.296'N, 66°24.459'W, San Nicolasito Field Station, 10.i.2009, leg. Short & Miller, VZ09-0100- 02X, morichal (27 exs., SEMC); same data except VZ09-0110-02 (1 ex. KBMC); same data except 62 m, leg. Short, Tellez, & Arias, 5.vii.2010, lights at morichal, VZ10-0705-02B (1 ex., SEMC); 8°6.226'N, 66°26.228'W, 52 m, UCV San Nicolasito Field Station, Rio Aguaro, 10.i.2009, leg. Short, Miller, Joly, Garcia, & Camacho, VZ09-0110-01A (66 exs., MALUZ, MIZA, SEMC).

Diagnosis. Elytra maculate. Distinguishable from other members of Liocanthydrus by the following combination of characters: a) slender appearance ( Fig. 7 ), L/GW = 2.08–2.29, PNWpost/PNWant = 1.53–1.62; b) elytral patterning as in Fig. 7 , posterior bands oblique;anterior bands slightly oblique, nearly perpendicular to elytral suture; c) eye size large relative to head capsule ( Figs 7 , 14 ), HW/EW = 1.60–1.67; d) aedeagus as in Figs 20 a–e; median lobe slender and not sinuate in dorsal aspect ( Fig. 20 b), distally elongate and attenuate in lateral aspect ( Figs 20 a, c); right lateral lobe triangular with distal angle acute, not projected ( Fig. 20 e)

Externally L. clayae appears similar to L. octoguttatus and L. nanops n. sp. However, L. clayae is distinguished from both species by its much larger relative eye size and by its less obliquely oriented posterior elytral bands that do not cross onto the basal halves of the elytra. Liocanthydrus clayae also differs by its distinct aedeagus, with a median lobe that is slender and not sinuate in dorsal aspect and is distally attenuate in lateral aspect. Also L. clayae has a triangular right lateral lobe with an acute distal angle, similar to L. octoguttatus , but not subtriangular with the distal angle projected and rounded as in L. nanops .

Description. Measurements. TL = 2.7–3.15 mm; male max. = 3.08 mm, female max. = 3.15 mm, male min. = 2.75 mm, female min. = 2.7 mm, GW = 1.2–1.45. L/GW = 2.08–2.29, HW = 0.75–0.9 mm, EW = 0.45–0.55 mm, HW/EW = 1.60–1.67, PNWpost = 1.2–1.4 mm, PNWant = 0.75–0.9 mm, PNWpost/PNWant = 1.53–1.62.

Coloration. Elytra maculate, with transverse bands; color of elytra ranging from brown to very dark reddish brown to nearly black; color of head pronotum and elytral bands yellow or brownish yellow to brownish red; elytral bands as in Fig. 7 , often constricted or reduced medially, sometimes reduced to spots, bands discontinuous across elytral suture, anterolateral extensions of posterior bands never crossing to basal half of elytra; color of venter yellowish brown to darker reddish brown.

Body. Body outline as in Fig. 7 , oval, narrow, attenuate posteriorly; slightly convex in lateral aspect ( Fig. 14 ).

Thorax. Elytra with lateral margins slightly ascendant medially in lateral aspect ( Fig. 14 ).

Head. Eyes large relative to head capsule, HW/EW = 1.60–1.67 ( Figs 7 , 14 ).

Abdomen. Aedeagus as in Figs 20 a–e; median lobe slender, not sinuate in dorsal aspect ( Fig. 20 b), distally elongate and attenuate in lateral aspect; left lateral lobe narrowed from base, moderately curved; setal fringe restricted to distal third ( Fig. 20 d); right lateral lobe triangular, distal angle acute and not projected ( Fig. 20 e).

Variability. This species exhibited some variation in overall color, size, and prominence of maculae. The vast majority of specimens were similarly colored, with dark brown or reddish brown elytra, brownish yellow maculae, pronota, and heads, and reddish brown venters. However, some individuals are lighter or darker in overall coloration with colors ranging as stated above (‘coloration’). Variation was also observed in the prominence of the elytral bands. Most often, members of this species appear as in Fig. 7 , with more or less complete bands, often constricted, but in some individuals reduced to a series of spots. Despite this, the respective oblique orientation of the bands or spots remained consistent within all members of the species. The last notable source of variation was size, with individuals ranging from 2.7–3.15 mm in length.

Biology. Liocanthydrus clayae has been collected in a wide variety of aquatic habitats, including marshes and forest pools, however, the majority of specimens were found in some type lotic environment. In Guyana, individuals were commonly found in streams, sometimes near waterfalls, that held debris such as leaf packs, log jams, or root mats. A single collecting event from “pools at the base of falls” yielded over 200 individuals. In Venezuela, specimens were taken from many habitat types, but the largest numbers of individuals were taken from morichal habitats.

Distribution. This species occurs in the northern region of South America. Specimens examined were collected from Venezuela, Guyana, and Suriname

Figures

FIGURES 3 – 9. Liocanthydrus species, dorsal habitus: 3) L. angustus; 4) L. uniformis; 5) L. bicolor; 6) L. armulatus sp. n.; 7) L. clayae; 8) L. nanops sp. n.; 9) L. octoguttatus (illustration). Scale bar = 1.0 mm.

FIGURES 10 – 15. Liocanthydrus species, lateral aspect: 10) L. angustus; 11) L. uniformis; 12) L. bicolor; 13) L. armulatus sp. n.; 14) L. clayae; 15) L. nanops sp. n. Scale bar = 1.0 mm.

FIGURES 16 – 22. Aedeagi of Liocanthydrus species. a) median lobe, right lateral aspect; b) median lobe, dorsal aspect; c) median lobe, left lateral aspect; d) left lateral lobe, right lateral aspect; e) right lateral lobe, left lateral aspect: 16) L. angustus; 17) L. uniformis; 18) L. bicolor; 19) L. armulatus sp. n.; 20) L. clayae; 21) L. nanops sp. n.; 22) L. octoguttatus. Scale bars = 0.2 mm. Notes: Terminology follows K. B. Miller & Nilsson (2003). Illustrations of lateral lobes depict medial (inner) surfaces.