taxonID	type	description	language	source
03B3067F0C0B1B08BAE07792471DFAA2.taxon	diagnosis	Diagnosis of larva. Surface of head capsule smooth. Nasale with three to four distinct teeth (Figs 2 C, 4 A, 5 B). Both epistomal lobes projecting anteriorly, slightly asymmetrical, rounded or angulate (Figs 2 C, 4 A, 5 B). Epistomal lobe bearing a few cuticular spines; gFR 2 composed of two or three setae (Fig. 2 C). Frontal lines lyriform, not convergent at base; coronal line absent (Fig. 2 A). Sensilla PA 26 – 28 not closely grouped; PA 27 between PA 26 and PA 28 (Fig. 2 B). Antenna, maxilla and labium not bearing long cuticular hairs (Figs 3 A, 4 B, 6 A). Antennomere 1 longer than second (Figs 3 A, 4 B, 6 A). Antennal SE 1 long, as long as antennomere 3 (Figs 3 A, 4 B, 6 A). Mandibles symmetrical, with two or three inner teeth (Figs 3 B, 4 C, 6 B). Ligula present, as a small tubercle or long (Figs 3 D, 4 E, 6 D). Prosternal sclerite divided into two plates (Fig. 5 E). Mesonotal sclerites closely placed (Fig. 5 F). Leg five segmented, short (Fig. 5 H). Abdominal segments more or less lobed laterally and with transverse rows of tubercles dorsally and laterally (Fig. 1 C, D, F – I). Spiracular atrium developed, without acrocercus and prostylus (Fig. 5 I). Tracheal gills absent (Fig. 1 C, D).	en	Minoshima, Yûsuke N., Watanabe, Kohei (2024): Morphology of the egg-case and larva of Paracymus aeneus (Germar) (Coleoptera: Hydrophilidae). Zootaxa 5541 (2): 200-214, DOI: 10.11646/zootaxa.5541.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5541.2.5
03B3067F0C0B1B08BAE07792471DFAA2.taxon	discussion	Remarks. The tribe Laccobiini comprises two lineages: the Laccobius - group and the Paracymus - group. Paracymus belongs to the latter, and its larvae exhibit greater similarity with the genera of the Paracymus - group than with the Laccobius - group. The Paracymus - group comprises five genera, Afrotormus Hansen, 1999, Paracymus, Scoliopsis Orchymont, 1919, Tormus Sharp, 1884 and Tritonus Mulsant, 1844, with the larval morphology of three of these genera (Paracymus, Tormus and Tritonus) having been described (Archangelsky 1997, 1999; Short & Fikáček 2013; Fikáček et al. 2013, 2017; this study). Larvae of the Laccobius - group genera have a piercing-sucking type (sensu Rodriguez et al. (2020 )) morphology of the mandible and the epistomal lobe, which are considered adaptations to a benthic lifestyle in aquatic habitats (Rodriguez et al. 2020). In contrast, larvae of the Paracymus - group genera have more ‘ typical’ features within Hydrophilidae. Among the larvae of the Paracymus - group, the New Zealand endemic terrestrial genus Tormus is the most morphologically similar to Paracymus, as indicated by previous phylogenetic studies revealing a sister group relationship (Short & Fikáček 2013; Toussaint et al. 2016; Toussaint & Short 2018). Fikáček et al. (2013) discussed the morphological similarity and evolutionary implications of Tormus and Paracymus larvae based on their general morphology. The most notable differences between the larvae are that Tormus bears numerous long cuticular hairs on the mouthparts (excluding the mandibles) and lacks a developed ligula (Fikáček et al. 2013), whereas such long cuticular hairs are absent (Fig. 6) and a more or less developed ligula is present (Fig. 4 E; Archangelsky 1997, 1999) in Paracymus. Larvae of the Afrotropical genus Tritonus also share characters with those of Paracymus, but Tritonus is differentiated by having V-shaped frontal lines, the presence of a coronal line, the position of PA 26 – 28 (forming a group), a small antennal sensorium SE 1, and an undivided prosternum (Fikáček et al. 2017).	en	Minoshima, Yûsuke N., Watanabe, Kohei (2024): Morphology of the egg-case and larva of Paracymus aeneus (Germar) (Coleoptera: Hydrophilidae). Zootaxa 5541 (2): 200-214, DOI: 10.11646/zootaxa.5541.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5541.2.5
03B3067F0C0B1B00BAE072EA4728FB1E.taxon	description	(Figs 1 – 6) Description of egg-case. Egg-case (Fig. 1 A, B, E). Whitish, sac-shaped, composed by globular part and long, slender mast. Mast narrowing apically. Description of larvae. First instar. General morphology similar to second instar larva. Thorax slightly wider than head capsule (Fig. 1 C, F, G). Spiracles on mesothorax and abdominal segments 1 – 7 proportionally more strongly protruded than in second instar (Fig. 1 P). Head. Frontal lines similar to second instar (Fig. 2 A). Epistomal lobe weakly angulate at apex (Fig. 2 C). Maxilla (Fig. 3 C). Borderline between sclerite and membrane of head appendages hardly visible in several cases, but generally identical with later instar. Cuticular scale-like structures on dorsal membrane of stipes present but very indistinct (Fig. 3 C). Labium (Fig. 3 D). Anterior membranous part of mentum bearing cuticular spines laterally. Primary chaetotaxy of head. Frontale (Fig. 2 A, C). Median part with three pairs of sensilla (FR 1 – 3); FR 1 long seta on midlength, close to frontal line; FR 2 pore-like, anteromesal to FR 1; FR 3 very short seta, anterior to FR 2. Three sensilla (FR 4 – 6) closely aggregated behind inner margin of antennal socket; FR 4 pore-like, mesal to FR 6; FR 5 short seta behind FR 6; FR 6 long seta. FR 7 short seta on inner margin of antennal socket. Short seta FR 8 and pore-like sensillum FR 15 behind nasale, between antennal sockets. Sensilla FR 9 – 14 on epistome; FR 11 pore-like, mesal, very close to lateral seta of gFR 1; FR 12 very short seta, lateral to FR 11; FR 13 pore-like behind FR 12; FR 9 long seta, very close and posterolateral to FR 13; FR 10 short seta behind FR 9 and FR 13; FR 14 pore-like, posterolateral to FR 9. Nasale with eight setae (gFR 1); one short seta on each lateral area; two short, stout mesal setae on anterior margin; two very short, stout setae behind mesal setae; two very short ventral setae on anterior margin. Apex of epistomal lobe with two very short, rather stout setae (gFR 2). Parietale (Fig. 2 A – B). Dorsal surface with five sensilla (PA 1 – 5) in posterior area, irregularly longitudinally distributed; PA 1, PA 2, PA 4 and PA 5 minute setae, PA 3 pore-like; PA 1 behind PA 3; PA 3 lateral to PA 2; PA 2 posterior to PA 4; PA 4 posteromesal to PA 5; PA 5 between PA 4 and PA 7. PA 6 pore-like sensillum, basal, close to frontal line, mesal to PA 2. PA 7, PA 12 – 17 forming transverse row at midlength on lateral area of head capsule; from dorsal to ventral, PA 7 (very long seta), PA 12 (short seta), PA 13 (very long seta), PA 14 (very long seta), PA 15 (pore-like), PA 16 (very long seta), PA 17 (pore-like). PA 10 pore-like sensillum between PA 7 and PA 8. PA 8 very long seta behind antennal socket, close to frontal line. PA 9 very long seta behind lateral margin of antennal socket. PA 11 short seta between PA 9 and PA 13. Anterior corner of head capsule with seven sensilla (PA 19 – 25). PA 19 pore-like, dorsal; PA 20 somewhat short seta on lateral face, lateral to PA 19; PA 21 long seta, lateral, ventral to PA 20; PA 22 long seta, lateroventral, mesal to PA 21. Pore-like sensilla PA 23 – 25 close to ventral mandibular articulation; PA 23 lateral to PA 24; PA 24 between PA 23 and PA 25; PA 25 on mesal margin. PA 26 long seta on anterior third, mesal to PA 17. Long seta PA 18 ventral, on midlength of lateral area, posterior to PA 17. PA 30 pore-like, posterior to PA 18. PA 27 and PA 28 on midlength and midwidth of ventral parietale, close to each other; PA 27 pore-like, anterior to very long seta PA 28. Pore-like sensillum PA 29 on posterior fifth, somewhat close to gular sulcus. Antenna (Fig. 3 A). Antennomere 1 with five pore-like sensilla (AN 1 – 5). AN 1 dorsal on posterior third; AN 2 – 5 subapical; AN 2 dorsal; AN 3 on outer face; AN 4 on inner face; AN 5 ventral. Antennomere 2 with six sensilla (AN 6 – 8, AN 10, AN 11 and SE 1). AN 6 pore-like, on subapical area of sclerite. AN 7, AN 8, AN 10, AN 11 and SE 1 on intersegmental membrane between antennomeres 2 and 3; AN 7 very short seta, on outer face, behind SE 1; AN 8 minute seta, dorsolateral, behind SE 1; AN 9 absent; AN 10 and AN 11 on inner face; AN 10 very long seta, anterior to very short seta AN 11. Sensorium SE 1 on outer face, long, rather stout, weakly curved, as long as or slightly longer than antennomere 3. Antennomere 3 with six apical sensilla of variable length and shape (gAN) on apical membranous area. Mandible (Fig. 3 B). Mandible with two setae (MN 1 and MN 5) and three pore-like sensilla (MN 2 – 4); MN 6 undetectable. MN 1 – 4 at midlength; somewhat short seta MN 1 on posterior third of lateral margin, posterolateral to MN 2; MN 2 – 4 forming a triangle; MN 2 between MN 1 and MN 3; MN 3 mesal to MN 2; MN 4 on lateral margin, anterior to MN 1. MN 5 very short seta, on anterior third of lateral margin. Maxilla (Fig. 3 C). Cardo with one short ventral seta (MX 1). Stipes with a row of five setae (MX 7 – 11) along dorsal inner face; MX 7 short, basal; MX 8 – 11 short, stout, almost equidistant. MX 2 and MX 3 ventral pore-like sensilla at midlength; MX 2 on median area; MX 3 on inner area. MX 4 – 6 lateral, subapical; MX 4 pore-like, behind MX 6; MX 5 very long seta, ventral to MX 6; MX 6 long seta. Dorsal surface of palpomere 1 with one somewhat long, stout seta (MX 16) on inner face. Lateroventral area with three sensilla (MX 12 – 14) closely aggregated; MX 12 pore-like, lateral to MX 14; MX 13 very long seta, posterior to MX 12 and MX 14; MX 14 long seta. MX 15 and MX 17 pore-like, on membrane behind inner appendage; MX 17 dorsal; MX 15 ventral. Inner appendage with one somewhat long seta and a few (possibly four) setae (gAPP). Palpomere 2 with two pore-like sensilla (MX 18 and MX 19) and one very short seta (MX 27). MX 18 ventral on lateral face; MX 19 dorsal on inner face of intersegmental membrane between palpomeres 2 and 3; MX 27 basal on outer face. Palpomere 3 with four sensilla (MX 20 – 23). Pore-like sensillum MX 20 and long seta MX 23 on outer face; MX 20 ventral to MX 23. Long seta MX 21 and pore-like sensillum MX 22 on inner face; MX 22 posterior to MX 21. Palpomere 4 with one somewhat long seta (MX 24) at base of inner face and with digitiform (MX 25) and pore-like (MX 26) sensilla on subapical part of outer face; MX 25 dorsal; MX 26 ventral. Apical membranous area of palpomere 4 with several very short setae of variable shape (gMX). Labium (Figs 2 B, 3 D). Submentum (Fig. 2 B) with two pairs of setae (LA 1 – 2); LA 1 very long on lateral corner of submentum; LA 2 short on anterolateral area. Mentum (Fig. 3 D) with two pairs of ventral sensilla (LA 3 and LA 4) on median part of anterior margin; LA 3 short seta, lateral to pore LA 4. Prementum with four pairs of sensilla (LA 8 – 11) on dorsal surface, three pairs (LA 5 – 7) on ventral surface. LA 8 pore-like on median part, on borderline of sclerite. LA 9 – 11 on median area of anterior membranous area, behind ligula; LA 9 pore-like, small, behind LA 10; LA 10 long seta, between LA 11 and LA 9; LA 11 pore-like, very close to or on base of ligula. LA 5 very short seta on basal margin of lateral face. LA 6 and LA 7 on anterior margin; LA 6 somewhat long, mesal to LA 7; LA 7 pore-like. LA 12 elongate pore-like, on dorsal apex of ligula. Labial palpomere 1 with two sensilla (LA 13 and LA 14); LA 13 very short seta, ventral, on basal margin; LA 14 pore-like, dorsal, on intersegmental membrane of palpomeres 1 and 2. Palpomere 2 with pore-like sensillum LA 15 on midlength of outer face and group of apical sensilla (gLA) on anterior membranous area. Second instar. General morphology similar to third instar. Frontal lines of head capsule clearly visible, lyriform, not convergent at base; coronal line absent (see first instar, Fig. 2 A). Labium (Fig. 4 E). Cuticular spines on anterior membrane of mentum reduced in size and number, often undetectable. Chaetotaxy of head, compared with first instar (see third instar, Fig. 5 C). Head capsule. Frontale: one rather long secondary seta behind FR 1. Parietale: one or two rather long secondary setae anterior to PA 6; two short secondary setae close to PA 8, one behind PA 8, one between PA 9 and PA 8; one short secondary seta lateral to PA 9; one rather short secondary seta anterior to PA 16. Mandible (Fig. 4 C) with four secondary setae on outer face; two short setae on basal part; one minute seta anterior to short setae; one minute seta behind MN 1. Maxilla (Fig. 4 D). Stipes with two long lateral secondary setae, one close to MX 5, one on subbasal part. Labium (Fig. 4 E). Mentum with three to four pairs of stout, short secondary setae on lateral part of membrane between mentum and prementum. Third instar. General morphology. Colour light, transparent yellowish white to grey with yellowish brown sclerites in living individuals (Fig. 1 I); thoracic and abdominal tubercles slightly darker in ethanol fixed specimens (Fig. 1 D). Spiracles: mesothoracic spiracle on lateroventral face, moderately protuberant; first seven abdominal spiracles on dorsolateral face, moderately protuberant (Fig. 1 R). Last pair on segment 8 large and enclosed within spiracular atrium (Fig. 1 S). Head. Head capsule subquadrate; cervical sclerites very small, irregularly shaped (Fig. 5 A). Frontal lines not visible. Gular sulcus only recognized by internal ridge (Fig. 5 D). Surface of head capsule smooth. Six stemmata on each anterolateral corner of head capsule (Fig. 5 A, also see Fig. 1 F – I). Clypeolabrum (Fig. 5 B) slightly asymmetrical. Nasale slightly asymmetrical with three distinct teeth aggregated mesally, almost equal in size and shape. Lateral lobes of epistome slightly asymmetrical, rounded anteriorly, both lobes projecting anteriorly as far as nasale teeth. Anterolateral part of lobes membranous. Both lobes bearing a few small cuticular teeth on anteromesal part, mesal to gFR 2. Ventral anterior margin of head capsule symmetrical. Antenna (Fig. 6 A) three-segmented, short, rather slender. Antennomere 1 longest and stoutest, slightly longer than antennomeres 2 and 3 combined; antennomere 2 twice as long as antennomere 3; antennomere 3 narrowest and shortest. Mandibles (Fig. 6 B) symmetrical, stout, strongly curved in apical half. Mandible with three inner teeth; anterior two teeth equal in size and shape, anterior margin simple, posterior margin finely serrate; posterior tooth smaller. Incisor area finely serrated. Maxilla (Fig. 6 C) six-segmented, longer than antenna. Cardo large, subtriangular. Stipes the longest and widest, distinctly longer than palpomeres 1 – 4 combined. Dorsal surface of stipes mostly membranous, borderline between sclerite and membrane indistinct or hardly visible. Lateral and ventral surface sclerotized excluding anterior area. Sclerotized part of dorsal surface reduced to an anterior transverse strip connecting inner and outer faces and a basal narrow, transverse area extending from base of inner face. Dorsal surface of stipes bearing fine cuticular projections; inner and anterior part bearing spine-like projections; mesal part bearing fine scale-like structures, hardly recognizable (see comments below). Maxillary palpus four-segmented. Palpomere 1 incompletely cylindrically sclerotized dorsally, the widest, slightly longer than wide, slightly shorter than palpomeres 2 to 4 combined. Inner appendage partly sclerotized. Palpomere 2 short, as long as and wider than palpomere 3. Palpomere 3 wider than palpomere 4. Palpomere 4 longer than palpomeres 2 and 3. Labium (Figs 5 D, 6 D) developed, but with reduced ligula. Submentum (Fig. 5 D) fused to head capsule, subpentagonal, wider than mentum. Submental sulcus hardly visible. Mentum (Fig. 6 D) longer than wide, subquadrate in dorsal view, subtrapezoidal in lateral view (see second instar, Fig. 4 E), wider than prementum; dorsal surface bearing ca. five teeth medially; short cuticular spine on anterior corner may be absent (see comments below). Prementum subquadrate; anterior membranous area with a few short lateral cuticular spines dorsally. Ligula reduced as small protuberance, partly sclerotized. Labial palpus rather long. Palpomere 1 short, two-thirds the length of palpomere 2; intersegmental membrane between palpomeres 1 and 2 bearing a few short cuticular spines dorsally. Palpomere 2 almost straight. Thorax. Surface of thoracic membrane covered with fine cuticular asperities (e. g., Fig. 1 N). Prothorax wider than head capsule (Fig. 1 D, I). Proscutum formed by one large plate subdivided by fine sagittal line, with sparsely arranged transverse row of short setae anteriorly, densely covered with fine spine-like cuticular projections (Fig. 1 M); anterior part weakly sclerotized. Prosternal sclerite with a pair of subquadrate plates (Fig. 5 E). Mesonotum with three pairs of sclerites on each side (Fig. 5 F); two transverse sclerites mesally on anterior margin; two small lateral sclerites on anterior margin; two large, posterior, transverse subtriangular sclerites. Two small, pigmented tubercles on lateral face; lateral one behind mesonotal spiracle, larger than mesal one. Metanotum with a pair of narrow, transverse sclerites on median area of anterior margin (Fig. 5 F). Five pairs of tubercles on posterior part of dorsal and lateral face of mesothorax, forming a transverse row. Mesal-most tubercle very small, indistinct; lateral four small, pigmented, from mesal to lateral: transverse tubercle, two circular tubercles, lateral one larger than remaining ones. Legs short, five-segmented (Fig. 5 H). Abdomen 10 - segmented, weakly lobed laterally (Fig. 1 D, I); surface of membrane covered with fine cuticular asperities (Fig. 1 N). Segments 1 to 7 similar in shape and size, bearing a pair of very small dorsal sclerites (Fig. 5 G) on median part of anterior margin and small, pigmented tubercles behind dorsal sclerites; tubercles forming three transverse rows on dorsal to lateral faces. Lateral face with large tubercles. Spiracular atrium developed (Fig. 5 I). Segment 8 as wide as posterior part of segment 7, bearing one small, pigmented tubercle on each side, with one large subquadrate dorsal plate partly covered with fine spine-like cuticular projections and bearing several setae of variable length (Fig. 1 O); posterior margin of dorsal plate partly membranous medially. Procercus incompletely sclerotized. Segment 9 trilobed, partly sclerotized, without acrocercus; urogomphi short, one-segmented; prostyli absent. Chaetotaxy of head capsule and head appendages (Figs 5 B – D, 6) very similar to second instar. Anterior corner of mentum with four (left) and two (right) setae in the examined specimen (see below Note). Notes on description. The larvae of P. aeneus are small, and some of the head appendages are partly membranous. This posed some limitations on their descriptions. The membranous part of the stipes is prone to distortion during slide preparation, obscuring character observations. Owing to their size and delicate structure, dorsal cuticular structures on the stipes are often challenging to observe, examination of multiple specimens is essential for an understanding of these characters. The number of secondary setae on the anterior corner of the mentum seem to vary intraspecifically. In this study, the examined third instar larva shows four setae on the left and two on the right, although it is possible that some setae on the right side could be broken.	en	Minoshima, Yûsuke N., Watanabe, Kohei (2024): Morphology of the egg-case and larva of Paracymus aeneus (Germar) (Coleoptera: Hydrophilidae). Zootaxa 5541 (2): 200-214, DOI: 10.11646/zootaxa.5541.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5541.2.5
03B3067F0C0B1B00BAE072EA4728FB1E.taxon	biology_ecology	Biology. Adults constructed eggs-cases on algae and stones as substrates (Fig. 1 E). Three larvae hatched from an egg-case. Larvae were observed walking on algae and water surface, breathing by exposing the tip of the abdomen to the water surface (Fig. 1 F). Larvae spent most of their time on wet land. Because they could not feed well in the water, they moved to land while holding the prey and lifted their head and ate the prey as reported in other hydrophilids (Fig. 1 J, K) (e. g., Archangelsky 1997). Larval cannibalism was frequently observed (Fig. 1 J). In containers containing water fleas, larval midgut content was coloured, suggesting that small invertebrates in the water also served as potential prey. Larvae that ceased feeding burrowed through the soil; however, all individuals died without reaching the adult stage. Since adults commenced egg laying three days after rearing began, June is presumed to be part of the reproductive season of this species. Adults can walk under the water surface using a ventral air bubble, a behaviour reported in the other hydrophilids (Takahashi 1922; Kurosawa 1987; Matsuno 1989; Gould & Valdez 2021) (Fig. 1 L).	en	Minoshima, Yûsuke N., Watanabe, Kohei (2024): Morphology of the egg-case and larva of Paracymus aeneus (Germar) (Coleoptera: Hydrophilidae). Zootaxa 5541 (2): 200-214, DOI: 10.11646/zootaxa.5541.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5541.2.5
