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Alyculus Kazantsev, 1999 Alyculus Kazantsev, 1999: 252. Type species: Alyculus kurbatovi Kazantsev, 1999 (by monotypy). Diagnosis. Alyculus is the only net-winged beetle with shortened elytra (Figs 1���2). Additionally, the very small body (1.1���2.7 mm) and shape of male genitalia (Figs 9���12) distinguish the genus from other lycids. Redescription. Male. Body tiny, dark testaceous or brownish black, never brightly colored (Figs 1���2). Head small, hypognathous, cranium with prominent frontal part, antennae inserted on frontal tubercles, cranium sometimes with apparent cell-like structure, covered with sparse long pubescence (Fig. 5). Eyes large, hemispherically prominent, eye diameter slightly larger than interocular distance. Antennae slender to moderately robust, filiform or antennomere 3 and subsequent ones slightly compressed; antennomere 1 long, antennomere 2 very small, approximately as long as wide, antennomere 3 slender, parallel-sided (Fig. 7) or sometimes compressed, wide (Fig. 6), subsequent antennomeres becoming gradually more slender and parallel-sided (Figs 6���8). Mouthparts tiny, reduced, with very small, vestigial mandibles, labrum minute, maxillary palpi 4 -segmented, both palpi with apical palpomeres pointed at apex. Pronotum strongly transverse, widest at basal margin, with posterior angles acutely prominent, lateral and basal margins variable in shape; pronotum either without any keels or costae, or longitudinal keel present along midline (Figs 3���5); disc with punctures and sparse, long, erected pubescence. Scutellum bilobed apically, distal processes slender, projected obliquely backwards (Fig. 5). Elytra without costae or reticulate cells, shortened, tapering to apex, rounded apically, densely covered with minute papillae bearing setae, which form dense pubescence (Fig. 1). Wings long, without veins, lightly infuscate. Legs slender, tarsomeres similar in width, slender. Male genitalia trilobate, phallus slender, usually slightly shorter than parameres, parameres widest at basal two fifths to one fourth, connected dorsally, approaching each other ventrally, phallobase simple (Figs 9���12). Body length: 1.1���2.7 mm. Remark. Only Lyropaeus and Platerodrilus have been collected in females and their neoteny documented (Bocak & Matsuda 2003). Females of the remaining Lyropaeinae genera, including Alyculus, are unknown. Although a limited number of Alyculus specimens are known, and therefore the female neoteny cannot be inferred from the absence of females in the available material, the neotenic development is hypothesized on the basis of male morphology and molecular data, which place the genus in relationships with Lyropaeus Waterhouse, 1878 and other neotenic lineages (Bocak et al. 2008, Bocak & Bocakova 2008). Distribution. Alyculus is at present known from the Sunda Shelf Islands, but without any record from the Malay Peninsula, whose fauna is zoogeographically very close to Sumatra and Borneo (Bocak & Yagi 2010). Three species occur in the mountain chain spreading from Northern Sumatra (A. wittmeri reported from the Aceh Province) through the central part of the island (A. tujuhensis sp. nov., Barisan Mts) to Western Java (A. kurbatovi from Mt. Gede and environs). The later mountain region was repeatedly connected with Sumatra by the subaerial Sunda Strait (Voris 2000) and shares fauna with Southern Sumatra. Although Sumatra in the present form has fairly recent origin, the presence of a limited extent of dry land during the time when most of Sumatra was submerged (~ 25 ��� 15 mya, Hall 2002) was supported by the dated evolution of another neotenic lineage, Scarelus Waterhouse, 1879 (Malohlava & Bocak 2010). The highly morphologically diversified species of Alyculus in Sumatra and Java also support the continuous presence of some dry land, as the neotenic lineages do not disperse and occupy for a long time very restricted ranges (Bocak & Malohlava 2010). A single species is known from Borneo, which is a much older island and rain forests were preserved in the region during the drier Pliocene and Pleistocene periods to a large extent (Canon et al. 2009). Therefore, Alyculus may be widespread also in the Crocker Range and the Kinabalu Massif. Biology. These lycids were sifted (A. kurbatovi), intercepted by Malaise traps (A. barclayi sp. nov., A. wittmeri) or swept from lower vegetation under the rain forest canopy (A. tujuhensis sp. nov., A. kurbatovi). All species of Alyculus are generally rare and individual species are known in a single or a few specimens from very limited ranges, typically represented by mountain tops. These small-bodied and feebly sclerotized neotenic lycids show a tendency to remain in humid conditions at the lowest herb stratum under canopy and they seldom fly. Although a low number of specimens have been collected, we can say that these neotenic lycids fly close to the surface of soil (typical height of flight up to 20 cm) and have never been collected in higher strata of vegetation. Their flight is very slow and the flying activity is restricted to weather conditions without wind, at high temperatures and humidity. Probably, the flight of males serves only for finding a conspecific female as low dispersal ability of males must be expected as a consequence of feeble, small body, flightlessness of larviform females, rareness and the dependence on stable rainforest habitats. The extensive distribution and vicariant ranges of Alyculus resemble the distribution of neotenic net-winged beetle Scarelus (Malohlava & Bocak 2010) and the species ranges correspond with the uninterrupted presence of the South East Asian rain forests in the mountains of Sumatra, western Java, and Northwestern Borneo during drier periods of the Pleistocene (Voris 2000). All recently collected species of Lyropaeinae inhabit undisturbed humid forests (Bocakova 2006) and are seldom collected in ecosystems affected by human activities. Considering their dependence on the original habitats and presumed low dispersal propensity, these animals are deeply threatened by the undergoing deforestation in the region (Sodhi et al. 2004)., Published as part of Ata, Va C L Av Pa L & K, L A D I S L Av B O C A, 2012, Revision of the genus Alyculus Kazantsev, 1999 (Coleoptera: Lycidae), pp. 61-67 in Zootaxa 3564 on pages 62-63, DOI: 10.5281/zenodo.282964, {"references":["Kazantsev, S. V. (1999) Alyculus, new genus, first brachypterous male lycid (Insecta: Coleoptera). Raffles Bulletin of Zoology, 47, 251 - 255.","Bocak, L. & Matsuda, K. (2003) Review of the immature stages of the family Lycidae (Insecta: Coleoptera). Journal of Natural History, 37, 1463 - 1507.","Bocak, L. & Yagi, T. (2010) Evolution of mimicry patterns in Metriorrhynchus (Coleoptera: Lycidae): the history of dispersal and speciation in South East Asia. Evolution, 64, 39 - 52.","Voris, H. K. (2000) Maps of Pleistocene sea levels in south-east Asia: shorelines, river systems and time duration. Journal of Biogeography, 27, 1153 - 1167.","Malohlava, V. & Bocak, L. (2010) Evidence of extreme habitat stability in a Southeast Asian biodiversity hotspot based on the evolutionary analysis of neotenic net-winged beetles. Molecular Ecology, 19, 4800 - 4811.","Bocakova, M. (2006) Review of the tribe Lyropaeini (Coleoptera: Lycidae). European Journal of Entomology, 103, 127 - 136.","Sodhi, N. S., Koh, L. P., Brook, B. W. & Ng, P. K. L. (2004) Southeast Asian biodiversity: an impending disaster. Trends in Ecology and Evolution, 19, 654 - 660."]}

Alyculus tujuhensis sp. nov. (Figs 4, 7, 10) Type material. Holotype, Sumatra, Jambi, Kersik Tua, Gunung Kerinci N. P., 19.��� 22. Jan. 2005, 1600���2000 m, Bolm lgt. (Voucher number of the holotype UPOLVP0029, LMBC). Paratypes, 5 males, the same locality data (Voucher number of the paratype UPOLVP0028, LMBC). Diagnosis. The species resembles in dark coloration A. kurbatovi, but it is easily recognizable by the presence of the sharp longitudinal keel in the pronotum and smaller body. The pronotal keel reaches five sixth of the way to the midline and ends abruptly before the posterior margin. Male genitalia resemble those of A. barclayi in the convex basal part of parameres, but A. barclayi differs in the shape of the pronotum, antennae, and the absence of the pronotal keel. Description. Body tiny, dark brown to black. Head small, hypognathous, cranium with apparent cell-form structure; eyes large hemispherically prominent. Eye diameter 1.06 times interocular distance. Antennae slender, filiform, antennomere 1 long, 2 very small, approximately as long as wide (Fig. 7), antennomere 3 parallel-sided, 2.2 times longer than width, subsequent antennomeres becoming gradually more slender. Mouthparts with mandibles reduced, labrum minute, maxillary palpi 4 -segmented, apically pointed. Pronotum transverse, widest at basal margin, anterior and lateral margins almost straight, posterior margin bisinuate, projected in middle, posterior angles acutely prominent; strong longitudinal keel along midline reaches five sixths of pronotal length, pronotal disc with dense irregular punctures (Fig. 4). Scutellum bilobed apically, distal tips projected obliquely backwards. Elytra without costae, shortened, tapering to apex, distal portion rounded. Elytra densely covered with minute papillae bearing fine setae, elytra without costae or reticulate cells. Legs slender. Male genitalia with slender phallus, parameres widest at basal third (Fig. 10). Measurements. Body length 1.65 mm, pronotal width 0.41 mm, pronotal length 0.24 mm, length of elytra 0.9 mm, maximum eye diameter in lateral view 0.137 mm, minimum frontal eye distance 0.121mm. Distribution. Sumatra: Jambi Province, Gunung Tujuh. Etymology. The specific name is derived from the local name Gunung Tujuh, where the species occurs., Published as part of Ata, Va C L Av Pa L & K, L A D I S L Av B O C A, 2012, Revision of the genus Alyculus Kazantsev, 1999 (Coleoptera: Lycidae), pp. 61-67 in Zootaxa 3564 on pages 65-66, DOI: 10.5281/zenodo.282964

Key to males of the genus Alyculus 1. Pronotum with strong longitudinal keel reaching five sixths of the pronotal midline length (Fig. 4), cranium with apparent celllike structure, male genitalia with slender phallus (Fig. 10)..................................... A. tujuhensis sp. nov. - Pronotum without apparent longitudinal keel (Figs 3, 5), cranium shiny or with fine irregular punctures, without apparent celllike structure......................................................................................... 2 2. Antennae slender, antennomere 3 at least 1.5 times longer than its width at apical part (Figs 1, 8)...................... 3 - Antennae robust, antennomere 3 apparently compressed, widest apically, 1.1 times longer than width at apex, subsequent antennomeres gradually slenderer, parallel-sided, less compressed to filiform (Fig. 6), male genitalia with slender phallus, paramerae widest at basal third (Fig. 11).................................................... A. barclayi sp. nov. 3. Body very small, less than 1.1 mm, testaceous, antennomere 3 is 1.5 times longer than width at apex (Fig. 1), male genitalia with slender phallus widened basally (Fig. 12).............................................. A. wittmeri Bocakova - Body large, about 2.5 mm long, dark brown to black, antennae slender, antennomere 3 is 1.8 times longer than width at apex (Fig. 8), male genitalia with short phallus (Fig. 9).......................................... A. kurbatovi Kazantsev, Published as part of Ata, Va C L Av Pa L & K, L A D I S L Av B O C A, 2012, Revision of the genus Alyculus Kazantsev, 1999 (Coleoptera: Lycidae), pp. 61-67 in Zootaxa 3564 on page 66, DOI: 10.5281/zenodo.282964

Alyculus barclayi sp. nov. (Figs 2, 6, 11) Type material. Holotype, male. Brunei, Kuala Balalong FSC, E 115 7 N 4 34, Dipterocarp forest, BM(NH) 1991 - 173, Ground FIT 1 B, 260 m alt., 22. vi. 91, N. Mawdsley, NM 204 (BMNH). Diagnosis. The body size, coloration, and the shape of male genitalia resemble those of A. tujuhensis sp. nov. A. barclayi is recognizable by the compressed antennae and the shorter, wide antennomere 3 (Fig. 6). Additionally, A. barclayi sp. nov. does not have any trace of the longitudinal keel in the pronotum. Description. Body tiny, dark brown to black. Head small, hypognathous, cranium without apparent microstructure, shiny, with sparse long pubescence. Eyes large, maximum eye diameter in lateral view 1.37 times minimal frontal eye distance, hemispherically prominent. Antennae moderately robust, compressed, antennomere 1 long, 2 very small, approximately as long as wide, antennomere 3 widest at apex, 1.1 longer than width (Fig. 6), subsequent antennomeres becoming gradually more slender, becoming parallel-sided. Mouthparts with reduced mandibles, labrum minute, maxillary palpi 4 -segmented, apically pointed. Pronotum strongly transverse, widest at basal margin, width at posterior margin 2.09 times length at midline, anterior and basal margins almost straight, posterior margin only slightly bisinuate, lateral margins concave before angles, posterior angles acutely prominent; pronotal disc without any keels or costae, with dense irregular punctures. Scutellum bilobed apically, distal tips slender, projected obliquely backwards. Elytra without costae, shortened, tapering to apex, distal portion rounded (Fig. 2). Elytra densely covered with minute papillae bearing setae, which form dense, long pubescence, elytra without costae or reticulate cells. Legs slender. Male genitalia with slender phallus, paramerae widest at basal third (Fig. 11). Measurements. Body length 1.35 mm, pronotal length 0.18 mm, pronotal width 0.37 mm, length of elytra 0.59 mm, minimum frontal eyes distance 0.100 mm, maximum eye diameter in lateral view 0.137 mm Distribution. Borneo: Brunei. Etymology. Named in honor of M. L. V. Barclay, the curator of the Coleoptera collection of the Museum of Natural History, London.

Alyculus wittmeri Bocakova, 2006 (Figs 1, 3, 12) Material studied. Holotype, male, Sumatra, Aceh prov., G. Leuser Nat. P., Ketambe, rainforest, 350 m, 3 �� 41��N, 97 �� 39��E, D. C. Darling, Malaise TP. (LMBC). Diagnosis. A. wittmeri differs from other known species in having the whole body light brown, and in male genitalia with slender parameres widest at basal fourth (Fig. 12). Redescription. Body tiny, light brown (Fig. 1). Head small, with large eyes, eye diameter 1.19 times interocular distance. Antennae slender, filiform, antennomere 1 long, 2 small, 3 weakly longer than 1 (Fig. 1). Mouthparts with mandibles reduced, labrum minute, maxillary palpi 4 -segmented, apically pointed. Pronotum transverse, widest at basal margin, length at midline 2.04 times width at posterior margin, anterior margin straight, lateral margins strongly emarginate, posterior margin bisinuate (Fig. 3); pronotal disc without carinae, shiny, with coarse punctures. Scutellum bilobed apically, distal tips projected obliquely backwards. Elytra shortened, tapering to apex, distal portion rounded. Elytra densely punctured, with no trace of costae or reticulate cells. Legs slender. Male genitalia with slender phallus widened basally (Fig. 12). Measurements. Body length 1.1 mm, humeral width 0.3 mm, pronotal length 0.12 mm, pronotal width 0.25 mm, length of elytra 0.4 mm, minimum frontal eye distance 0.084 mm, maximum eye diameter in lateral view 0.100 mm. Distribution. Indonesia: Sumatra, Aceh Province., Published as part of Ata, Va C L Av Pa L & K, L A D I S L Av B O C A, 2012, Revision of the genus Alyculus Kazantsev, 1999 (Coleoptera: Lycidae), pp. 61-67 in Zootaxa 3564 on page 66, DOI: 10.5281/zenodo.282964, {"references":["Bocakova, M. (2006) Review of the tribe Lyropaeini (Coleoptera: Lycidae). European Journal of Entomology, 103, 127 - 136."]}

Ata, Va C L Av Pa L, K, L A D I S L Av B O C A (2012): Revision of the genus Alyculus Kazantsev, 1999 (Coleoptera: Lycidae). Zootaxa 3564: 61-67, DOI: 10.5281/zenodo.282964

Alyculus kurbatovi Kazantsev, 1999 (Figs 5, 8, 9) Alyculus kurbatovi Kazantsev, 1999: 253. Material examined. Male. Indonesia, W Java, Puncak Pass, 1250���1600 m, 10 m of Cipanas, 8.��� 20. Oct. 2002, Bolm lgt. (Voucher number UPOL 000543, LMBC). Diagnosis. A. kurbatovi resembles in coloration A. tujuhensis sp. nov. and differs in the absence of the longitudinal pronotal keel and the shape of antennae (Figs 4 ���5, 7��� 8). Redescription. Body tiny, dark brown to black. Head small, hypognathous, cranium with very fine microstructure, shiny, with sparse long pubescence. Eyes large, hemispherically prominent, eye diameter 1.10 times interocular distance. Antennae moderately robust, antennomere 1 long, 2 very small, approximately as long as wide, antennomeres 3 to 4 slightly compressed, antennomere 3 narrowed at base, slightly wider and parallelsided in distal half, 1.8 times longer than width at apex (Fig. 8), subsequent antennomeres becoming gradually more slender, becoming less compressed, parallel-sided. Mouthparts with reduced mandibles, labrum minute, terminal palpomere of maxillary palpi apically pointed. Pronotum strongly transverse, widest at basal margin, 1.81 times longer than width at posterior margin, anterior margin rounded, lateral margins slightly concave, and basal margin bisinuate, posterior angles acutely prominent; pronotum without any keels or costae, punctures larger along midline, fine in remaining part of disc (Fig. 5). Scutellum bilobed apically, distal tips slender, projected obliquely backwards (Fig. 5). Elytra without costae, shortened, tapering to apex, distal portion rounded. Elytra densely covered with minute papillae bearing setae, which form dense, long pubescence, elytra without costae or reticulate cells. Legs slender. Male genitalia with short phallus, paramerae widest at basal third (Fig. 9). Measurements. Body length: 2.7 mm, pronotal length 0.26 mm, pronotal width: 0.47 mm, length of elytra: 0.98 mm, frontal interocular distance 0.142 mm, maximum eye diameter in lateral view 0.158 mm. Distribution. Indonesia: Java, Mt. Gede, Puncak Pass. Remark. The figure of male genitalia was redrawn from Kazantsev (1999) and shows very unusual shape of phallobase and a strange structure at the base of phallus (Fig. 9). Unfortunately, we have not a chance to check these structures as the abdomen of the second known specimen was used for DNA isolation. The shape of phallobase in other species is similar and concerning the variability of phallobase in other net-winged beetles we did not use phallobases in differential diagnoses., Published as part of Ata, Va C L Av Pa L & K, L A D I S L Av B O C A, 2012, Revision of the genus Alyculus Kazantsev, 1999 (Coleoptera: Lycidae), pp. 61-67 in Zootaxa 3564 on page 65, DOI: 10.5281/zenodo.282964, {"references":["Kazantsev, S. V. (1999) Alyculus, new genus, first brachypterous male lycid (Insecta: Coleoptera). Raffles Bulletin of Zoology, 47, 251 - 255."]}

Key to males of the genus Alyculus 1. Pronotum with strong longitudinal keel reaching five sixths of the pronotal midline length (Fig. 4), cranium with apparent celllike structure, male genitalia with slender phallus (Fig. 10)..................................... A. tujuhensis sp. nov. - Pronotum without apparent longitudinal keel (Figs 3, 5), cranium shiny or with fine irregular punctures, without apparent celllike structure......................................................................................... 2 2. Antennae slender, antennomere 3 at least 1.5 times longer than its width at apical part (Figs 1, 8)...................... 3 - Antennae robust, antennomere 3 apparently compressed, widest apically, 1.1 times longer than width at apex, subsequent antennomeres gradually slenderer, parallel-sided, less compressed to filiform (Fig. 6), male genitalia with slender phallus, paramerae widest at basal third (Fig. 11).................................................... A. barclayi sp. nov. 3. Body very small, less than 1.1 mm, testaceous, antennomere 3 is 1.5 times longer than width at apex (Fig. 1), male genitalia with slender phallus widened basally (Fig. 12).............................................. A. wittmeri Bocakova - Body large, about 2.5 mm long, dark brown to black, antennae slender, antennomere 3 is 1.8 times longer than width at apex (Fig. 8), male genitalia with short phallus (Fig. 9).......................................... A. kurbatovi Kazantsev