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Rhithrops capensis gen. et sp. nov. is described from the Matjies River, in the Cedeberg range of the Western Cape Province, South Africa. The new genus is apparently related to the South African endemic genera Rapnus Grouvelle, 1899 and Strina Redtenbacher, 1867, but exhibits a number of distinct character states, including gibbosities outside the sublateral pronotal furrows and a transversely upturned labral apex, unique within the family. Rhithrops gen. nov. was collected from around the roots of aquatic plants in fast flowing deep water runs, as well as being sampled in nocturnal drift nets. The new species has relatively long legs and well-developed tarsal claws, and appears to be adapted to an entirely aquatic, benthic existence. A revised key is provided to the dryopid genera of South Africa.

Revised key to South African Dryopidae genera The key below is based on that of Nelson (2007), adapted to incorporate Rhithrops gen. nov. and Pedestrodryops Kodada, 2001. 1 Almost entire dorsum and venter with long, fine, sparse erect sensillae; terrestrial, in plant litter away from water … Pedestrodryops Kodada, 2001 - Cuticular vestiture shorter and denser; aquatic taxa.......................................................... 2 2 Eyes conspicuously pubescent.......................................................................... 3 - Eyes not conspicuously pubescent, at most with scattered peg-like setae between facets............................. 4 3 First antennomere elongate; second antennomere large, transverse, more or less rectangular, distal border rounded......................................................................................... Ahaggaria Bollow, 1938 - First antennomere not elongate; second antennomere enlarged, ear-shaped, distal border angular...... Dryops Olivier, 1791 4 Antennae with 10 antennomeres; second antennomere enlarged, ear-shaped, covering most of rest of antennomeres; pronotum with sublateral furrows but no carinae..................................................... Rhithrops gen. nov. - Antennae with fewer than 10 antennomeres; second antennomere smaller; pronotum with sublateral stepped carinae...... 5 5 Antennae with 9 antennomeres; sublateral pronotal carinae with lateral flange; prosternal process large, gradually narrowed apically; mesotibial cleaning fringe a dense tuft of golden setae; ovipositor with coxites dorsally serrate Strina Redtenbacher, 1867 - Antennae with 7 antennomeres; lateral margin of pronotum without lateral flange; prosternal process large, abruptly narrowed apically; mesotibial cleaning fringe not a dense tuft of golden setae; ovipositor with coxites not serrate dorsally........................................................................................... Rapnus Grouvelle, 1899, Published as part of Bilton, David T. & Shepard, William D., 2022, Rhithrops capensis gen. et sp. nov., a new aquatic dryopid beetle from the Western Cape of South Africa (Coleoptera: Dryopoidea: Dryopidae), pp. 539-553 in Zootaxa 5195 (6) on page 552, DOI: 10.11646/zootaxa.5195.6.3, http://zenodo.org/record/7223884, {"references":["Nelson, H. G. (2007) Chapter 16 Dryopidae. In: Stals, R. & de Moor, I. J. eds. Guides to the Freshwater Invertebrates of Southern Africa. Vol. 10. Coleoptera. Water Research Commision., Pretoria, pp. 153 - 157.","Kodada, J. (2001) Pedestrodryops endroedyi gen. et sp. nov., a new terrestrial, wingless dryopid from South Africa (Coleoptera: Dryopidae). Entomological Problems, 33, 37 - 44.","Bollow, H. (1938) Monographie der palaearktischen Dryopidae, mit Berucksichtigung der eventuell transgredierenden Arten (Coleoptera). Mitteilungen der Munchener Entomologischen Gesellschaft, 28, 147 - 187.","Olivier, G. A. (1791) Encyclopedie methodique. Historie naturelle. Insectes. Vol. 6. hez Panckoucke, Imprimeur-Libraire, hotel de Thou, rue des Poitevins, Paris, 368 pp.","Redtenbacher, L. (1867) Reise des osterreichischen Fregatte Novara und die Erde in den Jahren 1857, 1858, 1859, unter der befehlen des Commodore B. von Wullerstorf- Urbair. Zoologischer Theil. Zweiter Band: Coleopteren. Kaiserlich- Ko ¨ nigliche Hof- und Staatsdruckerei; in Commission bei K. Gerold's Sohn, Wien, 249 pp.","Grouvelle, M. A. (1899) Decriptions de Clavicornes d'Afrique et de la region Malgache. 2 me Memoire. Annales de la Societe Entomologique de France, 68, 136 - 185."]}

Rhithrops gen. nov. (Figs 1–11) Type species. Rhithrops capensis sp. nov., by monotypy. Diagnosis. Rhithrops is unique amongst aquatic dryopid genera in that the labrum is transversely upturned apically. In South Africa, Rhithrops can be separated from Ahaggaria and Dryops by the lack of long erect setae on the eyes and dorsum of the body. Rhithrops can be separated from Strina and Rapnus by the antennae having 10 antennomeres as opposed to nine in Strina and seven in Rapnus. Additionally, Rhithrops has shallow sublateral pronotal furrows rather than sublateral step-like carinae, with discrete gibbosities on the pronotum outside of the sublateral furrows. The dorsally serrate coxites of the ovipositor are only shared with Strina, another South African dryopid genus. Abdominal ventrite 5 lacking a plastron is a character state commonly seen in the genus Helichus Erichson, 1847., Published as part of Bilton, David T. & Shepard, William D., 2022, Rhithrops capensis gen. et sp. nov., a new aquatic dryopid beetle from the Western Cape of South Africa (Coleoptera: Dryopoidea: Dryopidae), pp. 539-553 in Zootaxa 5195 (6) on page 540, DOI: 10.11646/zootaxa.5195.6.3, http://zenodo.org/record/7223884, {"references":["Erichson, W. H. (1847) Naturgeschichte der Insekten Deutschlands. Abteilung 1. Coleoptera, 3, 481 - 640."]}

The species of Elmomorphus Sharp, 1888 occurring in Japan and Korea are redescribed and illustrated: E. brevicornis Sharp, 1888 (Japan, Korea) and E. amamiensis Nomura, 1959 (Japan). The standard barcoding fragment of the mitochondrial gene coding for cytochrome c oxidase subunit I (COI) was sequenced and used together with morphological characters to delimit the taxonomic boundaries of the two species. To assess their morphometric variation, eight morphometric characters were measured and statistically evaluated using principal component analysis. The two species of Elmomorphus formed distinct and well-separated clusters in the COI tree. Their interspecific divergence is very high, ranging from 22.7 to 23.9%. On the other hand, morphometric characters, including those previously presumed to be diagnostic, overlap and per se do not allow unambiguous species identification. Reliable morphological distinguishing characters are described for males and females. Molecular data along with the morphological evidence strongly confirm the species status of E. amamiensis. An identification key to the Japanese and Korean species is provided.

Elmomorphus amamiensis Nomura, 1959 Figs 5B, 6B, D, F, 7E, 8C���D, 9C, E, 10B Elmomorphus brevicornis amamiensis Nomura, 1959: 33 (original description). Elmomorphus brevicornis amamiensis ��� Sat�� 1960: 252; 1965: 90; 1977: 1. ��� Kodada & J��ch 2006: 442 (catalogue). Elmomorphus amamiensis ��� Jung & Bae 2014: 5 (new status, distribution). ��� Kodada & J��ch 2016: 606 (catalogue). ��� Nakajima et al. 2020: 178, 322 (diagnosis, identification key, photographs of living specimens). Type locality Amami Island, Ryukyu Archipelago, Japan. Type material (not examined) See Jung & Bae (2014: 5). Material examined JAPAN ��� 1 ♂; ��� Amami-��shima Shinmura | 13. VI. 1962 M. Sato ���; NHMW ��� 1 ♂; ���(Ryukyus) Yona Okinawa 13.VIII.1969 Y. Hori leg.���; NHMW ��� 1 ♂; ��� Nago-shi, Okinawa-ken 2-5.VIII.1980 Col. K. Baba | Elmomorphus brevicornis amamiensis Nomura Det. M. Sato 1992 ���; NHMW ��� 4 ♂♂, 4 ♀♀; ��� Shinokawa, Setouchi-cho, Amami-��shima, Kagoshima, Japan, 28.22513, 129.29986, 5.III.2020, H. Yoshitomi leg.���; JKCB. Diagnosis Elmomorphus amamiensis is medium-sized (TL 2.80���3.22 mm), elongated oval, dorsally moderately convex, with the highest point slightly before the middle of the elytral length (lateral view); body outline slightly discontinuous between pronotum and elytra; plastron microscales covering the entire surface of the cranium, lateral portions of the pronotum and the whole dorsal surface of the elytra (Fig. 5B). Elytron with nine rows of moderately large, round punctures in shallow striae. Legs long and robust, somewhat shorter than the combined length of pronotum and elytra, claws moderately curved. Differences to the morphologically most similar E. brevicornis are discussed above. Measurements (mm) TL: ♂♂ 2.80���2.99 (2.91 �� 0.06, n = 9), ♀♀ 3.09���3.22 (3.16 �� 0.05, n = 7); PL: ♂♂ 0.68���0.77 (0.74 �� 0.03, n = 9), ♀♀ 0.70���0.78 (0.76 �� 0.03, n = 7); PW: ♂♂ 1.14���1.24 (1.20 �� 0.03, n = 9), ♀♀ 1.25���1.35 (1.31 �� 0.03, n = 7); EL: ♂♂ 2.18���2.34 (2.27 �� 0.05, n = 9), ♀♀ 2.42���2.60 (2.50 �� 0.05, n = 7); EW: ♂♂ 1.38���1.46 (1.42 �� 0.02, n = 9), ♀♀ 1.48���1.59 (1.54 �� 0.04, n = 7); MsTL: ♂♂ 0.84��� 0.96 (0.92 �� 0.03, n = 9), ♀♀ 0.83���0.87 (0.85 �� 0.01, n = 7); MtTL: ♂♂ 1.03���1.10 (1.07 �� 0.03, n = 9), ♀♀ 0.99���1.04 (1.02 �� 0.02, n = 7); PhL: 0.91���1.19 (1.02 �� 0.08, n = 7); PrL: 0.39���0.43 (0.42 �� 0.01, n = 7). Redescription BODY. Elongated oval, widest behind mid-length of elytra, moderately convex dorsally. Colouration black; mouthparts, antennae, tibiae and tarsi reddish-brown; femora dark brown. Main pubescence consisting of very short and thin, decumbent yellowish setae, which arise from small punctures. Dorsal plastron present on the entire surface except for the pronotal disc. Ventral surface except for prosternal process and disc of metaventrite with hair-like plastron setae. HEAD. With plastron and round setiferous punctures with a diameter of ca 0.75�� a facet diameter, punctures separated by distance 0.50���1.00�� a facet diameter. Labrum transverse, anterior margin straight, setose; anterolateral angles rounded; exposed portion microreticulate and micropunctate; male with a transverse row of strong, densely set conspicuous setae near middle (Fig. 6B), longest setae laterally, in length subequal to two-third of interocular distance; female with shorter and rather sparse setae, setal row hardly discernible. Anterior margin of clypeus straight, and not in the same plane as labrum. Eyes large, oval, protruding; interfacetal setae sparse, short; ID: ♂♂ 0.37���0.43 mm (0.40 �� 0.02, n = 9), ♀♀ 0.43���0.48 mm (0.44 �� 0.02, n = 7). Antennae 10-segmented, shorter than eye length. THORAX. Pronotum transverse, moderately convex, widest at base, PW/PL: ♂♂ 1.60���1.71 (1.62 �� 0.04, n = 9), ♀♀ 1.66���1.91 (1.73 �� 0.08, n = 7); plastron on entire lateral portions, mesally reaching level of third elytral row; pronotal disc smooth, with large, round setiferous punctures; punctures slightly smaller than facets, nearly regularly spaced, interstices 0.75���1.20�� a facet diameter. Anterior angles strongly protruding, acute and deflexed; lateral sides of pronotum convergent anteriad, slightly arcuate or nearly straight, narrowly explanate. Prosternal process with lateral margins divergent, moderately arcuate; posterior margin rounded; lateral portion slightly elevated, separated from median keel by longitudinal depressions; median keel nearly flat, surface finely punctate and irregularly wrinkled. Male with two clusters of conspicuous long setae in anterolateral portions of prosternal process (Fig. 6D), longest setae as long as prosternum in front of prosternal process. Scutellum longer than wide, surface smooth, with small, sparse punctures. Metaventrite along middle ca twice as long as prosternal process; disc flat in males, slightly elevated in females, discrimen present nearly along the entire length, surface finely sparsely punctate, and irregularly wrinkled. Metaventral process as long as prosternal one, with lateral sides keel-like, anterior margin not straight, flat; male with two clusters of conspicuous long setae in posterolateral portions of metaventrite (Fig. 6F). Elytra ovate, widest behind middle; sides moderately arcuate, more convergent posteriad than anteriad, narrowly explanate; EL/EW: ♂♂ 1.57��� 1.64 (1.60 �� 0.02, n = 9), ♀♀ 1.57���1.65 (1.62 �� 0.02, n = 7); entire surface with plastron; each elytron with nine striae, strial punctures smaller than facets, separated by distance of 1.00���2.00�� puncture diameter; punctures smaller laterad and toward apex. Tibiae all straight, slender; protibia ca 1.25�� as long as protarsus; PrTL/PL: ♂♂ 1.12���1.25 (1.19 �� 0.04, n = 9), ♀♀ 1.03���1.21 (1.09 �� 0.05, n = 7). Terminal protarsomere in male ca 1.20�� as long as all preceding tarsomeres combined; male foreclaws moderately curved, not widened or thickened, slightly less than half length of the terminal tarsomere length, both similar to each other and similar to female foreclaws. ABDOMEN. Ventrites completely covered with plastron; ventrites 1���5 with length ratio 1.00: 0.74: 0.52: 0.48: 1.66 in male, and 1.00: 0.88: 0.65: 0.57: 1.84 in female. Intercoxal process wide, about twice as wide as long, subtriangular, sides distinctly arcuate; admedian keels indistinct, feebly raised in male, reaching posterior margin of ventrite, but nearly absent in female. Mesal portion of male ventrite 5 flattened on anterior third and strongly deflected posteriad (lateral view); apex with distinct triangular excision and cluster of several long semi-erect setae on each side of excision (Fig. 7E). Female ventrite 5 evenly convex and only indistinctly flattened anteriorly, with minute longitudinal smooth keel at apex and few longer setae alongside keel; apex rounded. Aedeagus: phallobase long, narrow, expanded proximally (Fig. 8C), PhL/PrL: 2.32���2.74 (2.43 �� 0.12, n = 7); parameres strongly curved ventrad, especially in apical half (Fig. 8D), moderately narrowed, apices subacute; penis with apex slightly expanded and rounded in lateral aspect; sclerotized fibula absent. Ovipositor heavily sclerotized; right coxite ca 1.30�� as long as left one; paraproct ca 1.52�� as long as right coxite (Fig. 9D). Bursa copulatrix with cluster of minute spines on each side of distal portions and with several larger sclerites scattered over proximal portion (Fig. 9E). Sexual dimorphism Females are on average larger than males, and the apex of their fifth ventrite is rounded, while in males it is triangularly excised. Moreover, males have long conspicuous setae arranged in a transverse row on the labrum and in clusters situated on lateral sides of the prosternal process, lateral sides of the metaventrite and on the apex of the fifth ventrite; females lack these groups of long setae. Distribution Elmomorphus amamiensis has been so far recorded only from the Ryukyu Archipelago in Japan (Fig. 10B): Amami Island, Tokunoshima, Okinawa-jima, Iheya-jima, Zamami-shoto, Aka-jima, and Kume-jima (Nakajima et al. 2020)., Published as part of Kodada, J��n, Selnekovi��, D��vid, J��ch, Manfred A., Goffov��, Katar��na & V��a��n��, Peter, 2021, Taxonomic revision of the genus Elmomorphus Sharp, 1888 I. Japanese and Korean species (Coleoptera: Dryopidae), pp. 97-121 in European Journal of Taxonomy 758 (1) on pages 114-117, DOI: 10.5852/ejt.2021.758.1427, http://zenodo.org/record/5088151, {"references":["Nomura S. 1959. Notes on the Japanese Dryopoidea (Coleoptera) II. Toho-Gakuho 9: 33 - 38.","Sato M. 1960. Aquatic Coleoptera from Amami-Oshima of the Ryukyu Islands (I). Kontyu 28 (4): 251 - 254.","Sato M. 1965. Dryopoidea of the Ryukyu Archipelago, I (Coleoptera). Journal of the Nagoya Women's College 11: 76 - 94.","Sato M. 1977. Family Dryopidae. Check-List of Coleoptera of Japan 8: 1.","Kodada J. & Jach M. A. 2006. Family Dryopidae Billberg, 1820. In: Lobl I. & Smetana A. (eds) Catalogue of Palaearctic Coleoptera. Volume 3. Scarabaeoidea, Scirtoidea, Dascilloidea, Buprestoidea, Byrrhoidea: 441 - 443. Apollo Books, Stenstrup.","Jung S. W. & Bae Y. J. 2014. First record of the family Dryopidae (Coleoptera) in Korea with taxonomic notes on Elmomorphus brevicornis and Elmomorphus amamiensis. Animal Systematics, Evolution and Diversity 30 (1): 1 - 8. https: // doi. org / 10.5635 / ASED. 2014.30.1.001","Nakajima J., Hayashi M., Ishida K., Kitano T. & Yoshitomi H. 2020. Dryopidae. In: Aquatic Coleoptera and Hemiptera of Japan: 177 - 179. Bun-ichi Sogo Shuppan, Tokyo. [In Japanese.]","Sharp D. 1888. Descriptions of some new Coleoptera from Japan. Annals and Magazine of Natural History Series 6 2: 242 - 245. https: // doi. org / 10.1080 / 00222938809460918"]}

Elmomorphus brevicornis Sharp, 1888 Figs 4A���C, 5A, 6A, C, E, 7A���D, 8A���B, E���F, 9A���B, D, 10A Elmomorphus brevicornis Sharp, 1888: 243 (original description, type locality). Helichus ��� 862 spec.?��� Lewis 1879: 12 (first indication). Elmomorphus brevicornis ��� Zaitzev 1910: 19 (catalogue). ��� K��no 1934: 125 (catalogue). ��� Bollow 1940: 59���62, figs 271���276 (description, Japanese records). ��� Ch��j�� & Sat�� 1964: 193. ��� Del��ve 1968: 151 (differential diagnosis). ��� Sat�� 1981: 53. ��� J��ch 1984: 314 (checklist). ��� Kodada & J��ch 2006: 442 (catalogue). ��� Hayashi & Shimada 2006: 129 (Japanese records, in Japanese). ��� Yoshioka 2007: 236, 242, fig. 17a (Japanese records, in Japanese); 2008: 224 (Japanese records, in Japanese). ��� Hayashi 2011: 93. ��� Kamezawa & Nomura 2013: 29 (Japanese records, in Japanese). ��� Jung & Bae 2014: 2���7, figs 2a���c, 3���5 (description, identification key, Korean records). ��� Kodada & J��ch 2016: 606 (catalogue). ��� Yoshitomi & Haga 2018: 341���342 (identification key). ��� Nakajima et al. 2020: 177, 322 (diagnosis, identification key, photographs of living specimens). non Elmomorphus brevicornis ��� Devi et al. 2016: 371���373 (misidentification, invalid lectotype designation)]. Type locality Kobe, Hyōgo Prefecture, southern Honshu, Japan. Material examined Syntypes JAPAN ��� 1♂ (Fig.4C);���Kobe 18.V.71 [handwritten]|Japan.G Lewis1910���320.[printed]| Elmomorphus brevicornis [handwritten] | Elmomorphus brevicornis. [handwritten] | Elmomorphus brevicornis Sharp det. H. Bollow 1934 [handwritten and printed]���; NHMUK ��� 1 ♀ (Fig. 4A���B); ��� Elmomorphus brevicornis Type D.S. [David Sharp] Jokio [sic]. Japan Lewis. [handwritten] | Japan. G.Lewis [printed] | Sharp Coll. 1905-313. [printed] | Type [printed, round label with red frame]���; NHMUK. Additional material JAPAN ��� 1 ♀; ��� Shimakatsu Mie-Pref. 16.XI.1958 Coll. Shozo Ishida | Elmomorphus brevicornis Sharp Det. M. SATO 1971 ���; NHMW ��� 1 ♂; ��� Nagara-River Gifu-City 26.VII.1951 Coll. Zen. Naruse ���; NHMW ��� 1 ♂; ��� Kurokawa, N-Echigo 2.IX.1960 Col. K. Baba ���; NHMW ��� 1 ♂; ��� Kabuchi-gawa Gifu Pref. 18.VIII.1967 M. Sato leg.���; NHMW ��� 10 ♂♂, 6 ♀♀; ��� Japan, Shimane Pref.: Matsue: Yakumo 26.���30. iii.2019 M. Hayashi ���; JKCB ��� 5 ♂♂, 3 ♀♀; ��� Japan, Shimane Pref.: Izumo, Nishitani 30.iii.2019 M. Hayashi ���; JKCB ��� 1 ♀; ��� NE-JAPAN Miyagi Pr., Mts. Abukuma, 6.6. Uchikawa riv., leg. Ohmomo 1987 | Elmomorphus brevicornis Sharp ���; NHMW. SOUTH KOREA ��� 1 ♂, 1 spec. sex not examined; ��� South Korea, Jeollabuk-do, Buan-gun, Byeonsanmyeon, Junggye-ri, Jiko Falls 15.VI.2011, leg. S.W.Jung ���; NHMW. Note The original description was based on two specimens collected by G. Lewis. Both specimens are still preserved in the collection of David Sharp in the NHMUK. The female specimen is glued on a larger rectangular card with the following handwritten text: ��� Elmomorphus brevicornis Type D.S. Jokio. Japan Lewis.���. This style of labelling and label text is typical of David Sharp���s type material. The male specimen and the dissected aedeagus are glued on one card, while its detached appendages are glued on a second card, pinned below the specimen. The two syntypes are conspecific. However, while the type locality mentioned in the original description (���Kob�����) is written on the original label of the male specimen (Fig. 4C), the name ���Jokio��� [Sharp���s spelling of Tokyo] is found on the label of the female specimen (Fig. 4A). Since no other species is known from Honshu and since taxonomic confusion is improbable, a lectotype designation is considered unnecessary (see ICZN 1999: Recommendation 74G, https://www.iczn.org/the-code/declaration-44-amendment-of-article-74-7-3/). In the abstract of their most remarkable article, Devi et al. (2016: 371) wrote that ���A lectotype is designated for this species [Elmomorphus brevicornis]���. However, the term ��� lectotype ��� is not used anywhere else in their article, and it does not contain sufficient information to ensure recognition of the specimen designated. Therefore, the requirements of the International Code of Zoological Nomenclature (ICZN 1999: Art. 74.7) are not fulfilled. Diagnosis Elmomorphus brevicornis is a medium-sized (TL 2.59���3.51 mm), elongated oval, dorsally moderately convex species, with the highest point at the anterior third of the elytral length (lateral view). Body outline discontinuous between pronotum and elytra; yellowish plastron microscales cover the entire surface of the cranium, lateral portions of the pronotum and the entire dorsal surface of the elytra, so most of the dorsal surface appears to be matt (Fig. 5A). Each elytron has nine rows of large round punctures arranged in shallow striae. Legs conspicuously long and robust, only moderately shorter than the combined length of pronotum and elytra. Elmomorphus brevicornis closely resembles E. amamiensis in external morphology, and it can be distinguished by the following characteristics: (1) elytral sides in E. brevicornis subparallel in the anterior half, while in E. amamiensis they are arcuate and more convergent (Fig. 5B), thus the outline appears to be generally broader and less oval in E. brevicornis; (2) sides of pronotum usually more rounded in E. brevicornis, while they are almost straight in E. amamiensis; (3) males of E. brevicornis with several longer setae in a transverse row on the labrum, and with similar setae in two pairs of clusters on each prosternal process and on the admedian portions of metaventrite, row and clusters of setae rather inconspicuous (Fig. 6A, C, E); similar, longer setae also near the apex of the fifth ventrite (Fig. 7D); in the male of E. amamiensis the setal row and all setal clusters are well developed, conspicuous (Figs 6B, D, F, 7E); (4) parameres weakly curved ventrad in E. brevicornis (Fig. 8B), but strongly curved in E. amamiensis (Fig. 8D); (5) bursa copulatrix of the two species distally with one cluster of microspines on each side (Fig. 9D), while in E. amamiensis, besides these microspines there are several additional larger sclerites (Fig. 9E). Except for differences in the morphological characters, the two species differ by 22.7���23.9% in the partial mtDNA encoding for cytochrome c oxidase subunit I. Measurements (mm) TL: ♂♂ 2.89���3.38 (3.10 �� 0.13, n = 14), ♀♀ 3.15���3.51 (3.34 �� 0.12, n = 7); PL: ♂♂ 0.65���0.85 (0.78 �� 0.05, n = 14), ♀♀ 0.75���0.90 (0.83 �� 0.05, n = 7); PW: ♂♂ 1.20���1.43 (1.33 �� 0.06, n = 14), ♀♀ 1.30���1.47 (1.40 �� 0.06, n = 7); EL: ♂♂ 2.22���2.57 (2.42 �� 0.09, n = 14), ♀♀ 2.50���2.86 (2.66 �� 0.11, n = 7); EW: ♂♂ 1.44���1.66 (1.54 �� 0.05, n = 14), ♀♀ 1.51���1.77 (1.65 �� 0.09, n = 7); MsTL: ♂♂ 0.82��� 0.94 (0.88 �� 0.03, n = 14), ♀♀ 0.87���0.97 (0.91 �� 0.03, n = 7); MtTL: ♂♂ 0.91���1.09 (1.02 �� 0.05, n = 14), ♀♀ 0.97���1.12 (1.06 �� 0.05, n = 7); PhL: 0.73���0.89 (0.83 �� 0.04, n = 11); PrL: 0.32���0.39 (0.37 �� 0.02, n = 11). Redescription BODY. Elongated oval, widest behind mid-length of elytra, moderately convex dorsally, with the highest point at anterior third of elytral length (lateral view).Colouration black, except reddish-brown mouthparts, antennal clubs, tarsi, claws, trochanters, and ventral portions of femora. Tibiae and remainder of femoral surface dark brown to black. Dorsal pubescence consists mainly of short, thin decumbent yellowish setae arising from small punctures. Plastron microscales (Fig. 7B) on the entire cranium, on pronotum in two lateral bands, covering ca 0.2 of pronotal width on each side, and on the entire surface of elytra. Ventral surface with dense, thin plastron hair-like setae (Fig. 7C) except for prosternal process and median part of metaventrite. HEAD. Dorsally entirely covered with plastron scales and with round, setiferous punctures; puncture diameter ca 0.75�� diameter of an eye facet, and their distance varies between 0.50���1.00�� of a facet diameter. Labrum transverse, with anterior margin straight; anterolateral angles rounded; exposed portion microreticulate, with small setiferous punctures; two types of hair-like setae present, short decumbent setae on most of the surface, and several longer semi-erect setae in a hardly discernible transverse row near mid-length (Fig. 6A), setae nearly half as long as ID and more obvious in males (in older material long setae often abraded). Clypeus with anterior margin straight and with a row of the fine short hairs. Eyes oval, protruding and large, interfacetal setae short; ID: ♂♂ 0.38���0.43 mm (0.41 �� 0.02, n = 14), ♀♀ 0.42���0.48 mm (0.44 �� 0.02, n = 7); APD/ID: ♂♂ 1.93���2.09 (2.01 �� 0.04, n = 14), ♀♀ 1.89���2.04 (1.96 �� 0.05, n = 7). Antennae short, reaching slightly behind middle of eyes, 10-segmented; densely covered by plastron; each club segment with a conspicuous peg-like sensillum on anterior face and numerous hair-like, peg-like and branched sensilla; scape and pedicel small and subequal in length, not enlarged. THORAX. Pronotum transverse, widest at base, strongly convex, PW/PL: ♂♂ 1.60���1.85 (1.72 �� 0.07, n = 14), ♀♀ 1.60���1.76 (1.68 �� 0.05, n = 7); plastron in lateral band on each side along entire pronotal length, mesally reaching level of third elytral row; disc smooth, shiny, with large round setiferous punctures, punctures nearly as large as facets, separated by 0.50���1.00�� of a facet diameter; punctures on plastron area smaller, similar to those on head. Anterior angles strongly deflexed, protruding and acute, third as long as interocular distance; pronotal sides convergent anteriad, distinctly arcuate near middle, moderately explanate along entire length. Hypomeron widest behind mid-length. Prosternal process wide and short, lateral margins divergent and moderately arcuate; posterior margin widely rounded; lateral portion raised, wide and in male with group of longer setae (Fig. 6C) forming a hardly discernible setal cluster (these setae only moderately longer than other hair-like ventral setae and often abraded); median keel moderately convex, ca ⅓ as wide as width of prosternal process. Scutellum longer than wide, smooth, with small, sparse punctures. Metaventral process with lateral sides strongly raised, anterior margin not raised. Metaventral disc flat in female, finely depressed in male; longitudinal step-like elevation along sides delimits lateral area with plastron from smooth central one, surface irregularly and moderately sparsely punctate and in some specimens finely irregularly wrinkled; discrimen well impressed, distinct; a small, hardly discernible setal cluster present in front of metakatepisternal suture in males on each side of metaventral disc (Fig. 6E). Elytra oblong, widest behind middle; moderately convex dorsally, in dorsal view with the highest point at anterior third; sides subparallel in anterior half, more strongly arcuate in posterior half; EL/EW: ♂♂ 1.53���1.69 (1.57 �� 0.04, n = 14), ♀♀ 1.53���1.66 (1.61 �� 0.04, n = 7); surface entirely covered by plastron scales and with numerous minute punctures distinctly smaller than a facet diameter, punctures separated by about 0.50���1.00�� a facet diameter. Strial punctures large and deep on elytral disc and subequal in size with facet size, laterad and posteriad progressively smaller. All tibiae straight, to slightly bent near mid-length; protibia ca 1.25�� as long as protarsus; PrTL/PL: ♂♂ 1.01���1.20 (1.10 �� 0.05, n = 14), ♀♀ 1.00���1.08 (1.05 �� 0.03, n = 7). Terminal tarsomere in male foreleg ca 1.20�� as long as all preceding tarsomeres combined; male foreclaws strongly curved, not widened or thickened, about half of terminal tarsomere length, both similar to each other (Fig. 7A) and similar to female foreclaws. ABDOMEN. All ventrites completely covered with plastron; ventrites 1���5 with a length ratio about 1.00: 0.76: 0.60: 0.40: 1.26 in male, and 1.00: 0.90: 0.73: 0.60: 1.60 in female; intercoxal process wide, about twice as wide as long, subtriangular, sides moderately arcuate; admedian keels feebly raised, reaching posterior margin of ventrite. Male ventrite 5 near anterior margin declivous and then shortly flattened, laterad and posteriad evenly deflexed; apex with distinct triangular excision (Fig. 7D); several longer semi-erect setae on each side of excision in hardly discernible clusters. Female ventrite 5 evenly convex and not flattened anteriorly, with minute longitudinal smooth keel at apex and few longer setae alongside keel; apex rounded. Sternite VIII moderately sclerotized laterally, membranous medially, in males with short strut on anterior margin (Fig. 8E), in females with strut almost as long as abdomen (Fig. 9A). Male sternite IX narrow, with sclerotized anterior strut (Fig. 8F). Aedeagus: phallobase long, narrow, expanded proximally (Fig. 8A), PhL/PrL: 2.14���2.36 (2.24 �� 0.07, n = 11); parameres slightly curved ventrad along apical half, nearly straight, with apices narrowly rounded (Fig. 8B); apex of penis slightly expanded and rounded in lateral aspect, not reaching apex of parameres; ventral sclerotized fibula absent. Ovipositor heavily sclerotized; right coxite ca 1.40�� as long as left one; paraproct ca 1.80�� as long as right coxite (Fig. 9B). Bursa copulatrix with a cluster of numerous minute spines arranged on each side of distal portions (Fig. 9D). Spermatheca tubular; accessory gland large, semi-tubular. Sexual dimorphism Females are, on average longer and broader than males. For an unambiguous distinction between males and females, the apex of the fifth ventrite should be examined: it is distinctly excised in males (Fig. 7D), while it is rounded, with a short, smooth keel in females. The presence of several longer semi-erect setae on the labrum, prosternal process, metaventral disc and on the apex of the fifth ventrite in males is hard to observe even in well-preserved specimens, and even under a high-quality microscope at a magnification of about 100��. Distribution Japan: Honshu, Izu-��shima, Oki, Shikoku, Kyushu, Tsushima, and Tanegashima (Nakajima et al. 2020); South Korea (Fig. 10A). Devi et al. (2016) erroneously recorded E. brevicornis from India. Their record was published in the unreviewed ��� Journal of Entomology and Zoology Studies ���, a so-called predatory journal (see https://predatoryjournals.com/journals/). The ���redescription��� provided in this article does not allow any conclusions to be drawn on the identity of the specimens. The ���Aedagues��� [sic] depicted in their fig. 1c is, in fact, an ovipositor! Judging from their habitus photograph (fig. 1a), these specimens, beyond any doubt, do not belong to E. brevicornis or any other species similar to E. brevicornis. Immature stages The larva was described by Hayashi & Kadowaki (2008) and Hayashi (2015). It was found in a river on submerged roots of Salix L. and Carex L. (Hayashi pers. com.)., Published as part of Kodada, J��n, Selnekovi��, D��vid, J��ch, Manfred A., Goffov��, Katar��na & V��a��n��, Peter, 2021, Taxonomic revision of the genus Elmomorphus Sharp, 1888 I. Japanese and Korean species (Coleoptera: Dryopidae), pp. 97-121 in European Journal of Taxonomy 758 (1) on pages 106-114, DOI: 10.5852/ejt.2021.758.1427, http://zenodo.org/record/5088151, {"references":["Sharp D. 1888. Descriptions of some new Coleoptera from Japan. Annals and Magazine of Natural History Series 6 2: 242 - 245. https: // doi. org / 10.1080 / 00222938809460918","Lewis G. 1879. A Catalogue of Coleoptera from the Japanese Archipelago. Taylor and Francis, London.","Zaitzev P. 1910. Dryopidae, Cyathoceridae, Georyssidae, Heteroceridae. In: Schenkling S. (ed.) Coleopterorum Catalogus 17: 1 - 68. W. Junk, Berlin. Available from https: // www. biodiversitylibrary. org / item / 189656 page / 3 / mode / 1 up [accessed 4 Jun. 2021].","Kono H. 1934. Die Dryopiden aus Japan. Insecta Matsumurana 8 (3): 124 - 128.","Bollow H. 1940. Monographie der palaearktischen Dryopidae mit Berucksichtung der eventuell transgredierenden Arten (Col.). Mitteilungen der Munchner Entomologischen Gesellschaft 30 (1): 24 - 71.","Chujo M. & Sato M. 1964. Family Dryopidae. In: Kira T. & Umesao T. (eds) Nature and Life in Southeast Asia. Volume III: 193 - 194. Fauna and Flora Research Society, Kyoto.","Deleve J. 1968. Dryopidae et Elminthidae (Coleoptera) du Vietnam. Annales Historico-Naturales Musei Nationalis Hungarici 60: 149 - 181.","Sato M. 1981. Dryopidea (Coleoptera) of Nepal I. Family Dryopidae. Bulletin of Natural Science Museum Tokyo, Series A (Zoology) 7 (1): 51 - 56.","Jach M. A. 1984. Die Koleopterenfauna der Bergbache von Sudwest-Ceylon (Col.). Archiv fur Hydrobiologie Supplement 69 (2): 228 - 332.","Kodada J. & Jach M. A. 2006. Family Dryopidae Billberg, 1820. In: Lobl I. & Smetana A. (eds) Catalogue of Palaearctic Coleoptera. Volume 3. Scarabaeoidea, Scirtoidea, Dascilloidea, Buprestoidea, Byrrhoidea: 441 - 443. Apollo Books, Stenstrup.","Hayashi M. & Shimada T. 2006. Records on Elmidae and Dryopidae from Eastern Area of Shimane Prefecture and Oki Islands, Japan. Bulletin of the Hoshizaki Green Foundation 9: 127 - 143. [In Japanese with English abstract.]","Yoshioka M. 2007. Records on Dryopidae and Elmidae from water systems of Iinashi-gawa and Iu-gawa River, Eastern Area of Shimane Prefecture. Bulletin of the Hoshizaki Green Foundation 10: 235 - 256. [In Japanese with English abstract.]","Hayashi M. 2011. Aquatic Coleoptera of Shimane Prefecture. Special Bulletin of the Hoshizaki Green Foundation 1: 1 - 117.","Kamezawa H. & Nomura S. 2013. Records of Elmomorphus brevicornis Sharp from Izu-Oshima, Izu Islands, Japan. Sabyabane, New Series 9: 29. [In Japanese.]","Jung S. W. & Bae Y. J. 2014. First record of the family Dryopidae (Coleoptera) in Korea with taxonomic notes on Elmomorphus brevicornis and Elmomorphus amamiensis. Animal Systematics, Evolution and Diversity 30 (1): 1 - 8. https: // doi. org / 10.5635 / ASED. 2014.30.1.001","Yoshitomi H. & Haga K. 2018. A new record of Dryops nitidulus (Coleoptera, Dryopidae) from Japan. Elytra, Tokyo, New Series 8 (2): 341 - 342.","Nakajima J., Hayashi M., Ishida K., Kitano T. & Yoshitomi H. 2020. Dryopidae. In: Aquatic Coleoptera and Hemiptera of Japan: 177 - 179. Bun-ichi Sogo Shuppan, Tokyo. [In Japanese.]","Devi M. B., Devi O. S. & Wanghengbam L. 2016. Redescription of the first record of the species Elmomorphus brevicornis from the family Dryopidae (Coleoptera) in Manipur. Journal of Entomology and Zoology Studies 4 (3): 371 - 373.","ICZN 1999. International Code of Zoological Nomenclature. 4 th Ed. Tipografia La Garangola, Padova.","Nomura S. 1959. Notes on the Japanese Dryopoidea (Coleoptera) II. Toho-Gakuho 9: 33 - 38.","Hayashi M. & Kadowaki H. 2008. Records on aquatic Coleoptera from rivers in and around Mt. Daisen, Tottori Prefecture, Japan, part II. Bulletin of the Hoshizaki Green Foundation 11: 269 - 286. [In Japanese with English abstract.]","Hayashi M. 2015. Description of larva of Elmomorphus brevicornis Sharp (Coleoptera, Dryopidae). Bulletin of the Hoshizaki Green Foundation 18: 219 - 223. [In Japanese with English abstract.]"]}

Elmomorphus brevicornis Sharp, 1888 Figs 4A–C, 5A, 6A, C, E, 7A–D, 8A–B, E–F, 9A–B, D, 10A Elmomorphus brevicornis Sharp, 1888: 243 (original description, type locality). Helichus “ 862 spec.?” Lewis 1879: 12 (first indication). Elmomorphus brevicornis – Zaitzev 1910: 19 (catalogue). — Kôno 1934: 125 (catalogue). — Bollow 1940: 59–62, figs 271–276 (description, Japanese records). — Chûjô & Satô 1964: 193. — Delève 1968: 151 (differential diagnosis). — Satô 1981: 53. — Jäch 1984: 314 (checklist). — Kodada & Jäch 2006: 442 (catalogue). — Hayashi & Shimada 2006: 129 (Japanese records, in Japanese). — Yoshioka 2007: 236, 242, fig. 17a (Japanese records, in Japanese); 2008: 224 (Japanese records, in Japanese). — Hayashi 2011: 93. — Kamezawa & Nomura 2013: 29 (Japanese records, in Japanese). — Jung & Bae 2014: 2–7, figs 2a–c, 3–5 (description, identification key, Korean records). — Kodada & Jäch 2016: 606 (catalogue). — Yoshitomi & Haga 2018: 341–342 (identification key). — Nakajima et al. 2020: 177, 322 (diagnosis, identification key, photographs of living specimens). non Elmomorphus brevicornis – Devi et al. 2016: 371–373 (misidentification, invalid lectotype designation)]. Type locality Kobe, Hyōgo Prefecture, southern Honshu, Japan. Material examined Syntypes JAPAN • 1♂ (Fig.4C);“Kobe 18.V.71 [handwritten]|Japan.G Lewis1910—320.[printed]| Elmomorphus brevicornis [handwritten] | Elmomorphus brevicornis. [handwritten] | Elmomorphus brevicornis Sharp det. H. Bollow 1934 [handwritten and printed]”; NHMUK • 1 ♀ (Fig. 4A–B); “ Elmomorphus brevicornis Type D.S. [David Sharp] Jokio [sic]. Japan Lewis. [handwritten] | Japan. G.Lewis [printed] | Sharp Coll. 1905-313. [printed] | Type [printed, round label with red frame]”; NHMUK. Additional material JAPAN • 1 ♀; “ Shimakatsu Mie-Pref. 16.XI.1958 Coll. Shozo Ishida | Elmomorphus brevicornis Sharp Det. M. SATO 1971 ”; NHMW • 1 ♂; “ Nagara-River Gifu-City 26.VII.1951 Coll. Zen. Naruse ”; NHMW • 1 ♂; “ Kurokawa, N-Echigo 2.IX.1960 Col. K. Baba ”; NHMW • 1 ♂; “ Kabuchi-gawa Gifu Pref. 18.VIII.1967 M. Sato leg.”; NHMW • 10 ♂♂, 6 ♀♀; “ Japan, Shimane Pref.: Matsue: Yakumo 26.–30. iii.2019 M. Hayashi ”; JKCB • 5 ♂♂, 3 ♀♀; “ Japan, Shimane Pref.: Izumo, Nishitani 30.iii.2019 M. Hayashi ”; JKCB • 1 ♀; “ NE-JAPAN Miyagi Pr., Mts. Abukuma, 6.6. Uchikawa riv., leg. Ohmomo 1987 | Elmomorphus brevicornis Sharp ”; NHMW. SOUTH KOREA • 1 ♂, 1 spec. sex not examined; “ South Korea, Jeollabuk-do, Buan-gun, Byeonsanmyeon, Junggye-ri, Jiko Falls 15.VI.2011, leg. S.W.Jung ”; NHMW. Note The original description was based on two specimens collected by G. Lewis. Both specimens are still preserved in the collection of David Sharp in the NHMUK. The female specimen is glued on a larger rectangular card with the following handwritten text: “ Elmomorphus brevicornis Type D.S. Jokio. Japan Lewis.”. This style of labelling and label text is typical of David Sharp’s type material. The male specimen and the dissected aedeagus are glued on one card, while its detached appendages are glued on a second card, pinned below the specimen. The two syntypes are conspecific. However, while the type locality mentioned in the original description (“Kobé”) is written on the original label of the male specimen (Fig. 4C), the name “Jokio” [Sharp’s spelling of Tokyo] is found on the label of the female specimen (Fig. 4A). Since no other species is known from Honshu and since taxonomic confusion is improbable, a lectotype designation is considered unnecessary (see ICZN 1999: Recommendation 74G, https://www.iczn.org/the-code/declaration-44-amendment-of-article-74-7-3/). In the abstract of their most remarkable article, Devi et al. (2016: 371) wrote that “A lectotype is designated for this species [Elmomorphus brevicornis]”. However, the term “ lectotype ” is not used anywhere else in their article, and it does not contain sufficient information to ensure recognition of the specimen designated. Therefore, the requirements of the International Code of Zoological Nomenclature (ICZN 1999: Art. 74.7) are not fulfilled. Diagnosis Elmomorphus brevicornis is a medium-sized (TL 2.59–3.51 mm), elongated oval, dorsally moderately convex species, with the highest point at the anterior third of the elytral length (lateral view). Body outline discontinuous between pronotum and elytra; yellowish plastron microscales cover the entire surface of the cranium, lateral portions of the pronotum and the entire dorsal surface of the elytra, so most of the dorsal surface appears to be matt (Fig. 5A). Each elytron has nine rows of large round punctures arranged in shallow striae. Legs conspicuously long and robust, only moderately shorter than the combined length of pronotum and elytra. Elmomorphus brevicornis closely resembles E. amamiensis in external morphology, and it can be distinguished by the following characteristics: (1) elytral sides in E. brevicornis subparallel in the anterior half, while in E. amamiensis they are arcuate and more convergent (Fig. 5B), thus the outline appears to be generally broader and less oval in E. brevicornis; (2) sides of pronotum usually more rounded in E. brevicornis, while they are almost straight in E. amamiensis; (3) males of E. brevicornis with several longer setae in a transverse row on the labrum, and with similar setae in two pairs of clusters on each prosternal process and on the admedian portions of metaventrite, row and clusters of setae rather inconspicuous (Fig. 6A, C, E); similar, longer setae also near the apex of the fifth ventrite (Fig. 7D); in the male of E. amamiensis the setal row and all setal clusters are well developed, conspicuous (Figs 6B, D, F, 7E); (4) parameres weakly curved ventrad in E. brevicornis (Fig. 8B), but strongly curved in E. amamiensis (Fig. 8D); (5) bursa copulatrix of the two species distally with one cluster of microspines on each side (Fig. 9D), while in E. amamiensis, besides these microspines there are several additional larger sclerites (Fig. 9E). Except for differences in the morphological characters, the two species differ by 22.7–23.9% in the partial mtDNA encoding for cytochrome c oxidase subunit I. Measurements (mm) TL: ♂♂ 2.89–3.38 (3.10 ± 0.13, n = 14), ♀♀ 3.15–3.51 (3.34 ± 0.12, n = 7); PL: ♂♂ 0.65–0.85 (0.78 ± 0.05, n = 14), ♀♀ 0.75–0.90 (0.83 ± 0.05, n = 7); PW: ♂♂ 1.20–1.43 (1.33 ± 0.06, n = 14), ♀♀ 1.30–1.47 (1.40 ± 0.06, n = 7); EL: ♂♂ 2.22–2.57 (2.42 ± 0.09, n = 14), ♀♀ 2.50–2.86 (2.66 ± 0.11, n = 7); EW: ♂♂ 1.44–1.66 (1.54 ± 0.05, n = 14), ♀♀ 1.51–1.77 (1.65 ± 0.09, n = 7); MsTL: ♂♂ 0.82– 0.94 (0.88 ± 0.03, n = 14), ♀♀ 0.87–0.97 (0.91 ± 0.03, n = 7); MtTL: ♂♂ 0.91–1.09 (1.02 ± 0.05, n = 14), ♀♀ 0.97–1.12 (1.06 ± 0.05, n = 7); PhL: 0.73–0.89 (0.83 ± 0.04, n = 11); PrL: 0.32–0.39 (0.37 ± 0.02, n = 11). Redescription BODY. Elongated oval, widest behind mid-length of elytra, moderately convex dorsally, with the highest point at anterior third of elytral length (lateral view).Colouration black, except reddish-brown mouthparts, antennal clubs, tarsi, claws, trochanters, and ventral portions of femora. Tibiae and remainder of femoral surface dark brown to black. Dorsal pubescence consists mainly of short, thin decumbent yellowish setae arising from small punctures. Plastron microscales (Fig. 7B) on the entire cranium, on pronotum in two lateral bands, covering ca 0.2 of pronotal width on each side, and on the entire surface of elytra. Ventral surface with dense, thin plastron hair-like setae (Fig. 7C) except for prosternal process and median part of metaventrite. HEAD. Dorsally entirely covered with plastron scales and with round, setiferous punctures; puncture diameter ca 0.75× diameter of an eye facet, and their distance varies between 0.50–1.00× of a facet diameter. Labrum transverse, with anterior margin straight; anterolateral angles rounded; exposed portion microreticulate, with small setiferous punctures; two types of hair-like setae present, short decumbent setae on most of the surface, and several longer semi-erect setae in a hardly discernible transverse row near mid-length (Fig. 6A), setae nearly half as long as ID and more obvious in males (in older material long setae often abraded). Clypeus with anterior margin straight and with a row of the fine short hairs. Eyes oval, protruding and large, interfacetal setae short; ID: ♂♂ 0.38–0.43 mm (0.41 ± 0.02, n = 14), ♀♀ 0.42–0.48 mm (0.44 ± 0.02, n = 7); APD/ID: ♂♂ 1.93–2.09 (2.01 ± 0.04, n = 14), ♀♀ 1.89–2.04 (1.96 ± 0.05, n = 7). Antennae short, reaching slightly behind middle of eyes, 10-segmented; densely covered by plastron; each club segment with a conspicuous peg-like sensillum on anterior face and numerous hair-like, peg-like and branched sensilla; scape and pedicel small and subequal in length, not enlarged. THORAX. Pronotum transverse, widest at base, strongly convex, PW/PL: ♂♂ 1.60–1.85 (1.72 ± 0.07, n = 14), ♀♀ 1.60–1.76 (1.68 ± 0.05, n = 7); plastron in lateral band on each side along entire pronotal length, mesally reaching level of third elytral row; disc smooth, shiny, with large round setiferous punctures, punctures nearly as large as facets, separated by 0.50–1.00× of a facet diameter; punctures on plastron area smaller, similar to those on head. Anterior angles strongly deflexed, protruding and acute, third as long as interocular distance; pronotal sides convergent anteriad, distinctly arcuate near middle, moderately explanate along entire length. Hypomeron widest behind mid-length. Prosternal process wide and short, lateral margins divergent and moderately arcuate; posterior margin widely rounded; lateral portion raised, wide and in male with group of longer setae (Fig. 6C) forming a hardly discernible setal cluster (these setae only moderately longer than other hair-like ventral setae and often abraded); median keel moderately convex, ca ⅓ as wide as width of prosternal process. Scutellum longer than wide, smooth, with small, sparse punctures. Metaventral process with lateral sides strongly raised, anterior margin not raised. Metaventral disc flat in female, finely depressed in male; longitudinal step-like elevation along sides delimits lateral area with plastron from smooth central one, surface irregularly and moderately sparsely punctate and in some specimens finely irregularly wrinkled; discrimen well impressed, distinct; a small, hardly discernible setal cluster present in front of metakatepisternal suture in males on each side of metaventral disc (Fig. 6E). Elytra oblong, widest behind middle; moderately convex dorsally, in dorsal view with the highest point at anterior third; sides subparallel in anterior half, more strongly arcuate in posterior half; EL/EW: ♂♂ 1.53–1.69 (1.57 ± 0.04, n = 14), ♀♀ 1.53–1.66 (1.61 ± 0.04, n = 7); surface entirely covered by plastron scales and with numerous minute punctures distinctly smaller than a facet diameter, punctures separated by about 0.50–1.00× a facet diameter. Strial punctures large and deep on elytral disc and subequal in size with facet size, laterad and posteriad progressively smaller. All tibiae straight, to slightly bent near mid-length; protibia ca 1.25× as long as protarsus; PrTL/PL: ♂♂ 1.01–1.20 (1.10 ± 0.05, n = 14), ♀♀ 1.00–1.08 (1.05 ± 0.03, n = 7). Terminal tarsomere in male foreleg ca 1.20× as long as all preceding tarsomeres combined; male foreclaws strongly curved, not widened or thickened, about half of terminal tarsomere length, both similar to each other (Fig. 7A) and similar to female foreclaws. ABDOMEN. All ventrites completely covered with plastron; ventrites 1–5 with a length ratio about 1.00: 0.76: 0.60: 0.40: 1.26 in male, and 1.00: 0.90: 0.73: 0.60: 1.60 in female; intercoxal process wide, about twice as wide as long, subtriangular, sides moderately arcuate; admedian keels feebly raised, reaching posterior margin of ventrite. Male ventrite 5 near anterior margin declivous and then shortly flattened, laterad and posteriad evenly deflexed; apex with distinct triangular excision (Fig. 7D); several longer semi-erect setae on each side of excision in hardly discernible clusters. Female ventrite 5 evenly convex and not flattened anteriorly, with minute longitudinal smooth keel at apex and few longer setae alongside keel; apex rounded. Sternite VIII moderately sclerotized laterally, membranous medially, in males with short strut on anterior margin (Fig. 8E), in females with strut almost as long as abdomen (Fig. 9A). Male sternite IX narrow, with sclerotized anterior strut (Fig. 8F). Aedeagus: phallobase long, narrow, expanded proximally (Fig. 8A), PhL/PrL: 2.14–2.36 (2.24 ± 0.07, n = 11); parameres slightly curved ventrad along apical half, nearly straight, with apices narrowly rounded (Fig. 8B); apex of penis slightly expanded and rounded in lateral aspect, not reaching apex of parameres; ventral sclerotized fibula absent. Ovipositor heavily sclerotized; right coxite ca 1.40× as long as left one; paraproct ca 1.80× as long as right coxite (Fig. 9B). Bursa copulatrix with a cluster of numerous minute spines arranged on each side of distal portions (Fig. 9D). Spermatheca tubular; accessory gland large, semi-tubular. Sexual dimorphism Females are, on average longer and broader than males. For an unambiguous distinction between males and females, the apex of the fifth ventrite should be examined: it is distinctly excised in males (Fig. 7D), while it is rounded, with a short, smooth keel in females. The presence of several longer semi-erect setae on the labrum, prosternal process, metaventral disc and on the apex of the fifth ventrite in males is hard to observe even in well-preserved specimens, and even under a high-quality microscope at a magnification of about 100×. Distribution Japan: Honshu, Izu-Ôshima, Oki, Shikoku, Kyushu, Tsushima, and Tanegashima (Nakajima et al. 2020); South Korea (Fig. 10A). Devi et al. (2016) erroneously recorded E. brevicornis from India. Their record was published in the unreviewed “ Journal of Entomology and Zoology Studies ”, a so-called predatory journal (see https://predatoryjournals.com/journals/). The “redescription” provided in this article does not allow any conclusions to be drawn on the identity of the specimens. The “Aedagues” [sic] depicted in their fig. 1c is, in fact, an ovipositor! Judging from their habitus photograph (fig. 1a), these specimens, beyond any doubt, do not belong to E. brevicornis or any other species similar to E. brevicornis. Immature stages The larva was described by Hayashi & Kadowaki (2008) and Hayashi (2015). It was found in a river on submerged roots of Salix L. and Carex L. (Hayashi pers. com.).

Elmomorphus amamiensis Nomura, 1959 Figs 5B, 6B, D, F, 7E, 8C–D, 9C, E, 10B Elmomorphus brevicornis amamiensis Nomura, 1959: 33 (original description). Elmomorphus brevicornis amamiensis – Satô 1960: 252; 1965: 90; 1977: 1. — Kodada & Jäch 2006: 442 (catalogue). Elmomorphus amamiensis – Jung & Bae 2014: 5 (new status, distribution). — Kodada & Jäch 2016: 606 (catalogue). — Nakajima et al. 2020: 178, 322 (diagnosis, identification key, photographs of living specimens). Type locality Amami Island, Ryukyu Archipelago, Japan. Type material (not examined) See Jung & Bae (2014: 5). Material examined JAPAN • 1 ♂; “ Amami-Ôshima Shinmura | 13. VI. 1962 M. Sato ”; NHMW • 1 ♂; “(Ryukyus) Yona Okinawa 13.VIII.1969 Y. Hori leg.”; NHMW • 1 ♂; “ Nago-shi, Okinawa-ken 2-5.VIII.1980 Col. K. Baba | Elmomorphus brevicornis amamiensis Nomura Det. M. Sato 1992 ”; NHMW • 4 ♂♂, 4 ♀♀; “ Shinokawa, Setouchi-cho, Amami-Ôshima, Kagoshima, Japan, 28.22513, 129.29986, 5.III.2020, H. Yoshitomi leg.”; JKCB. Diagnosis Elmomorphus amamiensis is medium-sized (TL 2.80–3.22 mm), elongated oval, dorsally moderately convex, with the highest point slightly before the middle of the elytral length (lateral view); body outline slightly discontinuous between pronotum and elytra; plastron microscales covering the entire surface of the cranium, lateral portions of the pronotum and the whole dorsal surface of the elytra (Fig. 5B). Elytron with nine rows of moderately large, round punctures in shallow striae. Legs long and robust, somewhat shorter than the combined length of pronotum and elytra, claws moderately curved. Differences to the morphologically most similar E. brevicornis are discussed above. Measurements (mm) TL: ♂♂ 2.80–2.99 (2.91 ± 0.06, n = 9), ♀♀ 3.09–3.22 (3.16 ± 0.05, n = 7); PL: ♂♂ 0.68–0.77 (0.74 ± 0.03, n = 9), ♀♀ 0.70–0.78 (0.76 ± 0.03, n = 7); PW: ♂♂ 1.14–1.24 (1.20 ± 0.03, n = 9), ♀♀ 1.25–1.35 (1.31 ± 0.03, n = 7); EL: ♂♂ 2.18–2.34 (2.27 ± 0.05, n = 9), ♀♀ 2.42–2.60 (2.50 ± 0.05, n = 7); EW: ♂♂ 1.38–1.46 (1.42 ± 0.02, n = 9), ♀♀ 1.48–1.59 (1.54 ± 0.04, n = 7); MsTL: ♂♂ 0.84– 0.96 (0.92 ± 0.03, n = 9), ♀♀ 0.83–0.87 (0.85 ± 0.01, n = 7); MtTL: ♂♂ 1.03–1.10 (1.07 ± 0.03, n = 9), ♀♀ 0.99–1.04 (1.02 ± 0.02, n = 7); PhL: 0.91–1.19 (1.02 ± 0.08, n = 7); PrL: 0.39–0.43 (0.42 ± 0.01, n = 7). Redescription BODY. Elongated oval, widest behind mid-length of elytra, moderately convex dorsally. Colouration black; mouthparts, antennae, tibiae and tarsi reddish-brown; femora dark brown. Main pubescence consisting of very short and thin, decumbent yellowish setae, which arise from small punctures. Dorsal plastron present on the entire surface except for the pronotal disc. Ventral surface except for prosternal process and disc of metaventrite with hair-like plastron setae. HEAD. With plastron and round setiferous punctures with a diameter of ca 0.75× a facet diameter, punctures separated by distance 0.50–1.00× a facet diameter. Labrum transverse, anterior margin straight, setose; anterolateral angles rounded; exposed portion microreticulate and micropunctate; male with a transverse row of strong, densely set conspicuous setae near middle (Fig. 6B), longest setae laterally, in length subequal to two-third of interocular distance; female with shorter and rather sparse setae, setal row hardly discernible. Anterior margin of clypeus straight, and not in the same plane as labrum. Eyes large, oval, protruding; interfacetal setae sparse, short; ID: ♂♂ 0.37–0.43 mm (0.40 ± 0.02, n = 9), ♀♀ 0.43–0.48 mm (0.44 ± 0.02, n = 7). Antennae 10-segmented, shorter than eye length. THORAX. Pronotum transverse, moderately convex, widest at base, PW/PL: ♂♂ 1.60–1.71 (1.62 ± 0.04, n = 9), ♀♀ 1.66–1.91 (1.73 ± 0.08, n = 7); plastron on entire lateral portions, mesally reaching level of third elytral row; pronotal disc smooth, with large, round setiferous punctures; punctures slightly smaller than facets, nearly regularly spaced, interstices 0.75–1.20× a facet diameter. Anterior angles strongly protruding, acute and deflexed; lateral sides of pronotum convergent anteriad, slightly arcuate or nearly straight, narrowly explanate. Prosternal process with lateral margins divergent, moderately arcuate; posterior margin rounded; lateral portion slightly elevated, separated from median keel by longitudinal depressions; median keel nearly flat, surface finely punctate and irregularly wrinkled. Male with two clusters of conspicuous long setae in anterolateral portions of prosternal process (Fig. 6D), longest setae as long as prosternum in front of prosternal process. Scutellum longer than wide, surface smooth, with small, sparse punctures. Metaventrite along middle ca twice as long as prosternal process; disc flat in males, slightly elevated in females, discrimen present nearly along the entire length, surface finely sparsely punctate, and irregularly wrinkled. Metaventral process as long as prosternal one, with lateral sides keel-like, anterior margin not straight, flat; male with two clusters of conspicuous long setae in posterolateral portions of metaventrite (Fig. 6F). Elytra ovate, widest behind middle; sides moderately arcuate, more convergent posteriad than anteriad, narrowly explanate; EL/EW: ♂♂ 1.57– 1.64 (1.60 ± 0.02, n = 9), ♀♀ 1.57–1.65 (1.62 ± 0.02, n = 7); entire surface with plastron; each elytron with nine striae, strial punctures smaller than facets, separated by distance of 1.00–2.00× puncture diameter; punctures smaller laterad and toward apex. Tibiae all straight, slender; protibia ca 1.25× as long as protarsus; PrTL/PL: ♂♂ 1.12–1.25 (1.19 ± 0.04, n = 9), ♀♀ 1.03–1.21 (1.09 ± 0.05, n = 7). Terminal protarsomere in male ca 1.20× as long as all preceding tarsomeres combined; male foreclaws moderately curved, not widened or thickened, slightly less than half length of the terminal tarsomere length, both similar to each other and similar to female foreclaws. ABDOMEN. Ventrites completely covered with plastron; ventrites 1–5 with length ratio 1.00: 0.74: 0.52: 0.48: 1.66 in male, and 1.00: 0.88: 0.65: 0.57: 1.84 in female. Intercoxal process wide, about twice as wide as long, subtriangular, sides distinctly arcuate; admedian keels indistinct, feebly raised in male, reaching posterior margin of ventrite, but nearly absent in female. Mesal portion of male ventrite 5 flattened on anterior third and strongly deflected posteriad (lateral view); apex with distinct triangular excision and cluster of several long semi-erect setae on each side of excision (Fig. 7E). Female ventrite 5 evenly convex and only indistinctly flattened anteriorly, with minute longitudinal smooth keel at apex and few longer setae alongside keel; apex rounded. Aedeagus: phallobase long, narrow, expanded proximally (Fig. 8C), PhL/PrL: 2.32–2.74 (2.43 ± 0.12, n = 7); parameres strongly curved ventrad, especially in apical half (Fig. 8D), moderately narrowed, apices subacute; penis with apex slightly expanded and rounded in lateral aspect; sclerotized fibula absent. Ovipositor heavily sclerotized; right coxite ca 1.30× as long as left one; paraproct ca 1.52× as long as right coxite (Fig. 9D). Bursa copulatrix with cluster of minute spines on each side of distal portions and with several larger sclerites scattered over proximal portion (Fig. 9E). Sexual dimorphism Females are on average larger than males, and the apex of their fifth ventrite is rounded, while in males it is triangularly excised. Moreover, males have long conspicuous setae arranged in a transverse row on the labrum and in clusters situated on lateral sides of the prosternal process, lateral sides of the metaventrite and on the apex of the fifth ventrite; females lack these groups of long setae. Distribution Elmomorphus amamiensis has been so far recorded only from the Ryukyu Archipelago in Japan (Fig. 10B): Amami Island, Tokunoshima, Okinawa-jima, Iheya-jima, Zamami-shoto, Aka-jima, and Kume-jima (Nakajima et al. 2020).

Key to the adults of the species of Elmomorphus from Japan and Korea 1. Male labrum with several long, rather widely spaced setae arranged in a more or less distinct transverse row (Fig. 6A); similar, moderately longer setae arranged in small clusters on prosternal process and metaventrite (Fig. 6C, E), and on apex of ventrite 5 (Fig. 7D); long setae on labrum maximally half as long as interocular distance, most elongate ventral setae moderately shorter than those on labrum. Parameres in apical half slightly curved ventrad, nearly straight (Fig. 8B). Female: bursa copulatrix with a cluster of minute spines on each side in distal portions (Fig. 9D).................................................................................................................................. E. brevicornis Sharp, 1888 – Male labrum with numerous conspicuously long, closely arranged setae in a transverse row (Fig. 6B); similar setae in extensive clusters on prosternal process, metaventrite (Fig. 6D, F), and on apex of ventrite 5 (Fig. 7E); most elongate setae on labrum about ⅔ of interocular distance, longest ventral setae moderately longer than interocular distance. Parameres in apical half strongly curved (Fig. 8D). Female: bursa copulatrix with cluster of minute spines and with several larger sclerites scattered over its proximal portion (Fig. 9E)...................................... E. amamiensis Nomura, 1959

Key to the adults of the species of Elmomorphus from Japan and Korea 1. Male labrum with several long, rather widely spaced setae arranged in a more or less distinct transverse row (Fig. 6A); similar, moderately longer setae arranged in small clusters on prosternal process and metaventrite (Fig. 6C, E), and on apex of ventrite 5 (Fig. 7D); long setae on labrum maximally half as long as interocular distance, most elongate ventral setae moderately shorter than those on labrum. Parameres in apical half slightly curved ventrad, nearly straight (Fig. 8B). Female: bursa copulatrix with a cluster of minute spines on each side in distal portions (Fig. 9D).................................................................................................................................. E. brevicornis Sharp, 1888 ��� Male labrum with numerous conspicuously long, closely arranged setae in a transverse row (Fig. 6B); similar setae in extensive clusters on prosternal process, metaventrite (Fig. 6D, F), and on apex of ventrite 5 (Fig. 7E); most elongate setae on labrum about ⅔ of interocular distance, longest ventral setae moderately longer than interocular distance. Parameres in apical half strongly curved (Fig. 8D). Female: bursa copulatrix with cluster of minute spines and with several larger sclerites scattered over its proximal portion (Fig. 9E)...................................... E. amamiensis Nomura, 1959, Published as part of Kodada, J��n, Selnekovi��, D��vid, J��ch, Manfred A., Goffov��, Katar��na & V��a��n��, Peter, 2021, Taxonomic revision of the genus Elmomorphus Sharp, 1888 I. Japanese and Korean species (Coleoptera: Dryopidae), pp. 97-121 in European Journal of Taxonomy 758 (1) on page 117, DOI: 10.5852/ejt.2021.758.1427, http://zenodo.org/record/5088151, {"references":["Sharp D. 1888. Descriptions of some new Coleoptera from Japan. Annals and Magazine of Natural History Series 6 2: 242 - 245. https: // doi. org / 10.1080 / 00222938809460918","Nomura S. 1959. Notes on the Japanese Dryopoidea (Coleoptera) II. Toho-Gakuho 9: 33 - 38."]}

Löbl, Ivan, Smetana, Aleš (2020): On the Names of Three Genera of Dryopidae and Elateridae (Coleoptera). The Coleopterists Bulletin 74 (3): 542-543, DOI: 10.1649/0010-065X-74.3.542, URL: http://dx.doi.org/10.1649/0010-065x-74.3.542

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Genus Parygrus Erichson, 1847 Type species. Parygrus erichsoni Waterhouse, 1876 ; designated by subsequent monotypy. Diagnosis. Parygrus shares antennal morphology with Helichus, but the two genera can be separated by the following characters: Parygrus is very pubescent, covered with erect and semi-erect setae, including the eyes; plastron setae are not present; the enlarged 2nd antennomere covers antennomeres 3���4 or 3���5. Helichus is covered with recumbent plastron setae and lacks erect setae, the eyes are bare; the enlarged 2nd antennomere covers all of the distal antennomeres. Because of its pubescence, Parygrus appears most similar to Pelonomus, Microparnus, Onopelmus Spangler, Dryops Olivier and Platyparnus. These genera differ from Parygrus by the following: Pelonomus does not have an enlarged 2nd antennomere; Microparnus has 10 antennomeres and Onopelmus has 13; Dryops and Platyparnus have pronotal sublateral carinae/sulci. Generic redescription. Size from 4���8 mm long; elongate, parallel-sided or somewhat wider at posterior 1/3, moderately convex in lateral view; color uniformly brown; very setose, covered with long, stout, erect and/or semierect setae and shorter, finer, recumbent setae. Head covered with erect setae; frons projecting slightly or prominently forward between antennal bases; vertex flattened or shallowly depressed between eyes. Eyes hemispherical, protuberant, covered with erect setae, finely faceted. Antennae densely setose, pectinate, with 11 antennomeres; antennomere 2 enlarged, trapezoid, covering antennomeres 3���5 or 3���6; antennomere 3 shortest; antennomeres 4���11 forming a loose club; subantennal grooves deep below eyes; insertions approximate. Clypeus sub-triangular, wider than long. Labrum transverse, shorter than clypeus. Mandible with four teeth; ventral two teeth larger, acutely pointed; dorsal two teeth smaller, blunt, closely appressed. Maxillary palpi each with four palpomeres; labial palpi each with three palpomeres. Pronotum wider (~1.25x) than long; narrowly marginate and sometimes explanate laterally; anterior angles very acute; posterior angles barely acute, depressed; carinae absent, disc slightly flattened; punctate with large, distinct punctures interspersed with very small, fine punctures. Hypomeron flat, wide, narrower near anterior margin. Scutellum: shape variable, anterior margin arcuate. Elytron without carinae; finely or coarsely punctate-striate, with nine rows of punctures; narrowly margined. Epipleuron sinuate, broad in basal 1/3, narrow in apical 2/3 with groove to receive lateral edge of abdominal ventrite 5. Hind wing: macropterous (in species examined). Legs with major structures similarly proportioned; proleg shortest, metaleg longest. Procoxae oblong and flat, more widely separated than meso- and metacoxae; mesocoxae globular, separated by about one coxal width; metacoxae transverse, separated by about same distance as mesocoxae, excavate to receive femora; metatrochanters swollen; protibia weakly to strongly arcuate, sometimes deflexed (males); mesotibia weakly to moderately arcuate; metatibia straight; tarsomeres increasingly longer from base to apex, tarsomeres 1 and 2 very short, tarsomere 5 as long as previous 4 combined; claws simple. Venter with short, fine, recumbent setae present on lateral surfaces including hypomera, pro-, meso- and metepisterna, mesepimera, epipleura; long, semi-recumbent setae present medially. Prosternum long; lateral surface angled at about 45�� to central disc; prosternal process broad between procoxae, laterally margined, with a median, longitudinal carina terminating in a low protuberance. Mesoventrite very short, deeply excavate to receive prosternal spine. Metaventrite long, projecting anteriorly between mesocoxae, margined; lateral surface angled at about 45�� to posterior disc; metathoracic discrimen distinct; posterior border bidentate between metacoxae; metanepisternum wider anteriorly than posteriorly. Abdomen convex, with 5 ventrites; punctures aligned with faint, transverse strigae; ventrite 1 with margined, triangular intercoxal process, posterior border straight; ventrites 2���4 transverse, subequal in length; ventrite 5 longer, with broadly rounded apex. Genitalia. Male genitalia, trilobate type; female genitalia, common type (Kodada et al. 2016). Original description. Erichson���s 1847 descriptions of Parygrus and Helichus were presented in an annotated key. He cited the following characters for separating genera with ���auriculate��� antennae: Middle legs close to each other (Parygrus and Parnus); middle legs further apart (Helichus and Dryops). In addition, he stated that in Parygrus the elytra are clearly punctate-striate, and the antenna clearly ���11-membered.��� Waterhouse (1876), in his paper describing the type species P. erichsoni, stated that Erichson had established the genus to include undescribed species from South America with narrow form and strongly striated elytra. Sexual variation and dimorphism. Females are generally larger than males, and have a small, angular protuberance at the middle apex of abdominal ventrite 5 which is formed by a ventrally-directed fold. This feature is absent in males and therefore very useful in externally determining the sex of individuals. In the specimens we have examined, the protibiae of females were not deflexed and were usually less arcuate than those of conspecific males. Reproductive period. Fifteen females were collected along with 11 males from Paraguay, Alto Paraguay, Puerto Esperanza, 2 nd December 2006. Dissection found eggs in 13 of the 15 females. Thus the austral summer is the reproductive period. This is also true for Pelonomus spp. from Paraguay (W.D. Shepard, unpublished data). Geographic distribution. Parygrus is recorded from Mexico, south through Central America and the South American countries of Venezuela, Colombia, Ecuador, Peru, Bolivia, Paraguay, Brazil, and Argentina (Figs 2, 3, 4 ). Key to the genera of Neotropical aquatic and semiaquatic Dryopidae 1 Body dorsally glabrous; lateral margin of each elytron interrupted near apex by a densely pubescent depression................................................................................................ Elmoparnus Sharp ��� Body dorsally pubescent or with plastron; lateral margin of each elytron without densely pubescent depression.......... 2 2 Pronotum with complete sublateral carina/sulcus on each side....................................... Dryops Olivier ��� Pronotum without sublateral carinae/sulci or with only short carinae/sulci........................................ 3 3 Pronotum with sublateral carinae/sulci only in basal 1/4 to 1/3.................................................. 8 ��� Pronotum without sublateral carinae/sulci.................................................................. 4 4 Each antenna with 13 antennomeres...................................................... Onopelmus Spangler ��� Each antenna with 9 or 11 antennomeres.................................................................. 5 5 Each antenna with 9 antennomeres........................................................ Guaranius Spangler ��� Each antenna with 11 antennomeres....................................................................... 6 6 Antennomere 2 enlarged and covering rest of antennomeres..................................... Helichus Erichson ��� Antennomere 2 may be enlarged but not covering rest of antennomeres.......................................... 7 7 Antennomere 2 enlarged and at least covering antennomeres 3 and 4................................ Parygrus Erichson ��� Antennomere 2 not enlarged............................................................. Pelonomus Erichson 8 Length 6���10 mm; each antenna with 11 antennomeres; pro- and mesotarsus with tarsomere 2 wide and flattened in males............................................................................... Platyparnus Shepard and Barr ��� Length 2���3 mm; each antenna with 10 antennomeres; no tarsomeres wide and flattened............. Microparnus Shepard, Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 101-104, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261, {"references":["Erichson, W. F. (1847) Naturgeschichte der Insecten Deutschlands, Erste Abtheilung, Coleoptera. Vol. 3. Verlag der Nicolaischen Buchhandlung, Berlin, 968 pp. https: // doi. org / 10.5962 / bhl. title. 8270","Waterhouse, C. O. (1876) III. On various new genera and species of Coleoptera. Transactions of the Entomological Society of London, 1876 (Part I), 11 - 25. https: // doi. org / 10.1111 / j. 1365 - 2311.1876. tb 01122. x","Kodada, J., Jach, M. A. & Ciampor, F. (2016) 19.3 Dryopidae Bilberg, 1820 (1817). In: Beutel, R. G. & Leschen, R. A. B. (Eds.), Coleoptera, Beetles. Vol. 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). In: Beutel, R. G. & Kristensen, N. P. (Eds.), Handbook of Zoology, Arthropoda: Insecta, Second Edition. Walter de Gruyter, Berlin, pp. 590 - 602. https: // doi. org / 10.1515 / 9783110373929"]}

Parygrus parallelus (Grouvelle, 1890) (Figs 2, 11) Holotype specimen. BRAZIL: ���Theresopolis (Br��sil)��� (Grouvelle 1890, p. 146); ���Theresopolis (Rio de Janeiro). Collection Grouvelle.��� (Grouvelle 1896a, p. 216). Depository, MNHN. The type was not examined; locality data is from the original and subsequent descriptions. The type could not be located at the MNHN and its current whereabouts is unknown. Additional specimens, photographic images (2). BRAZIL: ���Bresil / Theresop[olis] // Parygrus / parallelus / Grouv. [handwritten]���. Depository, MNHN. Only photographic images and label data from the photos were examined. Additional specimens examined (8). BRAZIL: ���Brasil / Theres[opolis] // Parygrus / parallelus / Grouv. [handwritten]// Coll. Van de Poll / 1937-744. // H.E. Hinton / collection. / B.M. 1977-566. [printed]��� (1 specimen; head, pronotum and abdomen missing); ���Fry / Rio Jan ��. [handwritten] // G. Lewis Coll. / 1915-38. [printed]��� (1 specimen); ���Beske // Fry / Rio Jan ��. [handwritten] // Fry Coll. / 1905-100. [printed]��� (1 specimen); ���5797 // Beske // Fry / Rio Jan ��. [handwritten] // Fry Coll. / 1905-100. [printed] // Parygrus / parallelus / Grouv. [handwritten]��� (1 specimen; abdomen missing); ���Weir // San / Paulo [handwritten] // Fry Coll. / 1905-100. [printed]��� (1 specimen); ���27079 // Weir // San / Paulo [handwritten] // Fry Coll. / 1905-100. [printed] // Parygrus / parallelus / [illegible word] Grouv [handwritten]��� (1 specimen; head and pronotum missing); ���San / Paulo [handwritten] // G. Lewis Coll. / 1915-38 [printed]��� (2 specimens; 1 abdomen missing). Depository, NHMUK. Provisional diagnosis. The type specimen of P. parallelus is much longer (7���8 mm) than the length of the other species except for that of P. angustatus. Considering that an image of the type P. parallelus was not available for examination, the following was observed from images of two non-types (Additional specimens, photographic images) (Fig. 11) in comparison with the type image of P. angustatus (Fig. 8): elytra sinuate laterally, margin explanate at basal third (vs. parallel-sided, not explanate); pronotum widest at base, lateral margins raised and widely explanate (vs. not raised nor explanate), without a median longitudinal carina (vs. with a carina); elytral striae and punctures distinct (vs. striae not distinct, effaced at posterior third, punctures small). General description. Since we were unable to examine or view an image of the holotype, only a limited description is possible based on the original descriptions (Grouvelle 1890) and images of other specimens of P. parallelus from the Grouvelle Collection that we assume are correctly identified. From a translation of the original Latin and French descriptions: 7���8 mm long; elongate parallel; weakly convex; transverse carina between antennal insertions; scutellum pentagonal; pronotal anterior margin emarginate, lateral margins widely explanate and raised, fringed with long hairs; elytra not wider than pronotum, punctate-striate. To this we add that the elytral lateral margin is explanate at the basal third, that contrary to the description the elytra appear a little wider than the pronotum, and that it is not possible to see the transverse carina between the antennal insertions (Fig. 11). The latter character is perhaps generic because the frons and clypeus are set at different angles forming a rounded ridge between. Distribution. Brazil, described from Rio de Janeiro (Fig. 2). Etymology. Grouvelle did not cite an etymology, but the specific epithet probably refers to the parallel-sided body. Comments. Grouvelle (1890) originally described P. parallelus in Parnus and subsequently moved it to Parygrus (Grouvelle 1896a). The original description in Latin is brief and was presented at a meeting of the Entomological Society of France. The later description includes the verbatim original in Latin, as well as a more detailed one in French. The holotype could not be located in the MNHN. We were sent images of two specimens of P. parallelus ( Additional specimens, photographic images), one of which is included here (Fig. 11), and one image of a series of specimens in a museum unit tray. Several of the specimens in the image bear locality label data which match that of the type locality. At least two pins (one pin bears multiple specimens) have determination labels with the same handwriting as the holotype of P. angustatus and possibly P. elateroides, but the labels do not bear the notation ���ty.��� in the lower left corner. If the type remains missing, a lectotype should be designated. We were loaned a series of specimens from the NHMUK (Additional specimens examined), four of which were from the Fry Collection. Three of these bear determination labels with the same handwriting as the specimens labeled as P. parallelus from the Grouvelle collection. Unfortunately, two of them are missing the pronotum (and head), where important diagnostic characters are located. The remaining determined specimen and another Fry specimen, both from Rio de Janeiro, closely resemble those in the MNHN images, as do two other specimens from Sao Paulo collected by Lewis., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 110-111, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261, {"references":["Grouvelle, A. (1890) Deux especes nouvelle de parnides. Annales de la Societe Entomologique de France, Series 6, 10, 146.","Grouvelle, A. (1896 a) Nitidulides, Colydiides, Cucujides et Parnides recoltes par M. E. Gounelle au Bresil et autres Clavicornes nouveaux d'Amerique. Annales de la Societe Entomologique de France, 65, 177 - 216."]}

Parygrus guarani Barr and Shepard, new species (Figs 3, 14, 15) http://zoobank.org/ ADB49206-58A4-48C0-8578-8BC67B175FDB Type material. Holotype male. PARAGUAY. ��� PARAGUAY: ITAPUA / 17 km N Hohenau / Feb. 2-4, 1983 / Coll. E.G.Riley // HOLOTYPE / Parygrus guarani / Barr & Shepard [red label, handwritten] // EMEC49556��� (EMEC). Paratypes (1 M). [as above] // PARATYPE / Parygrus guarani / Barr & Shepard [yellow label, printed] (1 M, EMEC). Additional material examined (2 F). [as above] (1 F, LSAM); PARAGUAY: Cordillera Dept. / Caacup��, Campamento Jack / Morment; UV light trap / S25��22.116���, W057��07.917���/ 11-14-XI-2016, J.E.Eger / 812ft., permit #271/2016 (1 F, FSCA). Diagnosis. The male genitalia of Parygrus guarani are distinctive (Fig. 15); none of the other species (for which the genitalia are known) have a phallobase more than twice as long as the parameres in addition to parameres with inner teeth. In both the males and females, the body is slender, parallel-sided, and narrowly tapered posteriorly (Fig. 14); nearly all of the other species are slightly narrowed anterior to the middle and widened at the posterior 1/3, and are broadly rounded at the elytral apices. Parygrus angustatus is similarly parallel-sided (Fig. 8), but it is much longer (7���8 mm) than P. guarani (less than 5.6 mm). Because of the very small number of specimens on hand, it is possible that the above external characters may not be reliable for diagnosing the species. The two females were assigned to this species due to their association with a male and/or their similar morphology; they are not designated as paratypes. Description. Holotype male. Cuticle dark brown, legs, antennae and mouthparts lighter; slender, elongate, subparallel, elytra dorsally flattened along midline; length 5.50 mm (pronotum + elytra), width 1.90 mm, more than 2x as long as wide; body covered with erect and semi-erect, moderately long, golden setae and much shorter, recumbent setae (Fig. 14). Head dark brown; punctures separated by 1x puncture diameter; setae moderately long; vertex flat; frons protruding slightly forward between antennal bases, anterior margin barely arcuate, distance between antennal bases less than length of antennomere 1. Antenna with antennomeres 1 and 2 dark brown, setose; antennomeres 3���11 yellow-brown, more densely setose; antennomere 1 slightly shorter than antennomere 2, antennomere 2 covering antennomeres 3���6. Clypeus densely setose, deeply punctate, margin nearly straight. Maxillary palpus with terminal palpomere slightly clavate, ventral face flat. Labrum emarginate, densely setose, bordered with dense fringe of short setae. Labial palpus with terminal palpomere dorsoventrally flattened, vaguely kidney-shaped with one side straight, one side curved; length shorter than that of terminal maxillary palpomere. Pronotum dark brown; convex, disc weakly flattened at center; length 1.20 mm, width 1.55 mm, widest at basal 1/3 to 1/2; anterior margin straight except at strongly acute, depressed, anterolateral angles; lateral margins weakly arcuate, sinuate at base, narrowly margined, fringed with erect setae; posterior border trisinuate; posterolateral angles acute, moderately explanate. Disc evenly punctate, punctures separated by about 1x puncture diameter; setae moderately long. Scutellum dark brown, subcordate, wider than long; anterior margin strongly arcuate between two anterolateral notches; disc flat; punctation fine. Elytron dark brown; length 4.30 mm, width 0.95 mm; flattened; little variation in width except at apical 1/3. Humeral angle rounded; lateral margin widened and shallowly sulcate at basal 1/3, entire length narrowly margined; apex acute. Disc punctate and shallowly striate; central disc with moderately deep, closely spaced, punctures; base and apex with punctures shallow, striae effaced; stria 9 at basal 1/3 with larger, deeper punctures; disc at anterior 3/4 flattened at midline between suture and interval 2, other intervals weakly convex; setae shorter than on pronotum, semi-erect and recumbent, uniformly distributed. Hind wing: Macropterous. Legs. Profemur dark red-brown, covered with fine, evenly spaced punctures and long, recumbent golden setae; protibia red-brown, mostly bare and shiny, dorsal surface with sparse short setae and a few very long setae, ventral surface with dense row of short, stiff setae, arcuate with narrow spines at ventral apex; protarsus red-brown, shiny, dorsal surface with a few sparse, very long setae, ventral surface with dense, shorter setae, protarsomeres 4 and 5 each with a few much-longer dorsal setae. Mesofemur similar to profemur; mesotibia red-brown, mostly bare and shiny, dorsal surface with a few long setae, ventral surface with a patch of dense, short setae at apical 1/2, weakly arcuate with spinose ventral apex; mesotarsus similar to protarsus. Metafemur similar to pro- and mesofemur; metatibia weakly arcuate, setation like that of pro- and mesotibia; both metatarsi missing, but in the male paratype the metatarsi are similar to the pro- and mesotarsi. [Note: left metatarsus was lost after being imaged for Fig. 14.] Venter dark brown; heavily setose. Prosternum with anterior border narrowly margined; prosternal process wide between procoxae, widest at midlength, lateral margins weakly arcuate, thickly margined; process with median longitudinal carina, bordered laterally by sulci, terminating in a low, elongate protuberance near apex. Metaventrite with intercoxal process margined and depressed; posterior disc weakly depressed at junction of sulcate metakatepisternal suture and metathoracic discrimen. Abdomen with some setae longer than on rest of body except legs; ventrite 1 with triangular intercoxal process weakly depressed between metacoxae, thickly margined laterally; ventrites 2���4 of equal length, finely and evenly punctate at center of disc, coarser laterally, punctures aligned with faint, transverse strigae; ventrite 5 longest, disc coarsely punctate, covered with long setae prominent at margins. Genitalia. Aedeagus elongate (Fig. 15). Phallobase about 3x longer than parameres and wider than paramere bases together. Parameres each blade-like, inner surface concave; in dorsal view each with an inward-facing tooth near midlength; each paramere laterally sinuate, widest at base, narrowest near midlength, apex narrowly rounded; in lateral view aedeagus curved ventrally, each paramere wide, ventral surface nearly straight, dorsal surface weakly Intraspecific variation. No significant non-sexual variation was noted among the small number of specimens, except that in the females the elytra are more convex in the posterior 1/2 than in the males. The two males, 5.30 and 5.50 mm long, are close in size to the two females, 5.45 and 5.55 mm long. Geographic distribution. Known from only two localities in the departments of Cordillera and Itap��a, Paraguay (Fig. 3). Etymology. This new species is named in honor of the Guaran��, an indigenous people who inhabit Paraguay and neighboring countries., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 114-117, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261

Parygrus zamuco Barr and Shepard, new species (Figs 3, 20, 21) http://zoobank.org/ DE1B3D0C-5A2F-4175-8750-CE47331D2EEA Type material. Holotype male. BOLIVIA. ��� BOLIVIA: Santa Cruz / El Cairo, 5 km W Buena / Vista, 16-18-X- 2004 / J. E. Eger, UV light // HOLOTYPE / Parygrus zamuco / Barr & Shepard [red label, handwritten]��� (FSCA). Paratypes (24; 7 M, 17 F). BOLIVIA. BOL. Cochabamba / Carrasco, El Sacta / 220 m., 26/X/02 / Morris / Wappes (1 F, FSCA); BOLIVIA: Santa Cruz / El Cairo, 5 km W Buena / Vista, 16-18-X-2004 / J. E. Eger, UV light (1 M, EMEC; 1M, 1 F, FSCA); BOLIVIA: Sta. Cruz, / San Osteben [Opispo Santistevan] Prov., / Muyurina, 49 km N // Santa Cruz. XI-3-1959 / 1120 ft. R.B.Cumming // Property of Florida / State Collection of / Arthropods (1 F, EMEC; 1M, 4 F, FSCA); BOLIVIA: Santa Cruz / 3.7km.SSE. Buena Vista / Hotel Flora & Fauna 430m / 17��29.949���S - 63�� 33.152���W / 5-15-XI-2001, M C Thomas / trop.transition forest, BLT (1 F, EMEC); BOLIVIA: Santa Cruz / 3.7 km SSE Buena Vista / Hotel flora y fauna // 17��29���S 63��33���W / 23-30-April-2004; / A.R. Cline, MV&blacklight (2 F, CSCA); BOLIVIA: Dept. Santa Cruz, / 3.7km SSE Buena Vista / Hotel flora y fauna ~ 400m / 17��29���S 63��39���W; A. Cline // and J. Wappes collrs. / Ex: 23-30 APR-2004 / MV& BL (1 F, CSCA); BOLIVIA: Santa Cruz, 3.7 km SSW / Buena Vista, Hotel Flora y Fauna, / 1-12.V.2004, 17��29���S, 63��33���W, / Ex: MV/ Blacklight, Coll. A.R. Cline (1 F, CSCA); BOLIVIA: Santa Cruz / Dpt. 4k SSE B. Vista / 22-25 November 2013 / Wappes & Skillman // Hotel Flora & Fauna / El 350-400meters / 17��29��� S 63��49��� W (1 F, FSCA); BOLIVIA: Santa Cruz / Dpt. Huaico nr. Potrerillo / 21 November, 2013 / Wappes & Skillman // Elevation 430meters / MV/UV lights / 63��26��� W 17��40��� S (1F, FSCA); BOLIVIA: Santa Cruz / Potrerillos del Guend��; / 40km.NW. Santa Cruz / 17��40.3���S - 063��27.4���W / 22-XI to 12-XII- 2005 / B.K.Dozier (1 F, FSCA); BOLIVIA: Santa Cruz Dist. / Potrerillos del Guenda, Preserva / Natural, 17��40���S 63��27���W, 37m / 17-22-OCT-2007, A.R. Cline & J.E. / Wappes collr.; ex: bl/mv (1 F, CSCA); BOLIVIA: Santa Cruz / Potrerillo de Guand�� / 16-22-XII-2004 / coll. E. Nearns, UV trap // macro. (1M, FSCA). PARAGUAY. PARA- GUAY: Alto Paraguay / Puerto La Esperanza / 2 XII 2006 / Carlos Aguilar Julio (1M, EMEC); Concepci��n / San Alfredo / Tagatiy�� / 24 X 2019 (1 F, EMEC). VENEZUELA. VENEZUELA: Guarico / Est. Exp. Nicolasito / UCV ��� Fagro / 8��8���20������N- 66��24���32������W // 15-VI-2000, BL Trap / P.Freytag, M.Gaiani / Q.Arias collectors (1 M, EMEC; 1 M, FSCA). // All paratypes also have the following final label: ��� PARATYPE / Parygrus zamuco / Barr & Shepard��� [yellow label, printed]. Diagnosis. The male genitalia (Fig. 21), with lateral expansions of the parameres at the apical 1/4, are unique among species for which the genitalia are known. In addition, both males and females of Parygrus zamuco n. sp. have distinguishing external characteristics: none of the other species have a row of extremely long setae on the dorsal mesotibia, nor maxillary palpi as long and robust (Fig. 20). Description. Holotype male. Cuticle brown, legs, antennae and mouthparts lighter; elongate, subcylindrical; length 5.75 mm (pronotum + elytra), width 2.20 mm; body covered with erect and semi-erect, moderately long, golden setae and much shorter, recumbent setae (Fig. 20). Head dark brown; punctures shallow, separated by 1���2x puncture diameter; setae moderately long; vertex with shallow median depression; frons protruding slightly forward between antennal bases, anterior margin truncate, distance between antennal bases more than length of antennomere 1. Antenna with antennomeres 1 and 2 light brown, setose, antennomeres 3���11 yellow, densely setose; antennomere 1 shorter than antennomere 2, antennomere 2 covering antennomeres 3���6. Clypeus moderately setose and punctate, weakly emarginate. Maxillary palpus with terminal palpomere clavate, curved, very long, as long as clypeus. Labrum emarginate, moderately setose, bordered with fringe of long setae. Labial palpus with terminal palpomere vaguely triangular with 1 side straight, 1 side curved, ventral face slightly flattened; length nearly as long as that of terminal maxillary palpomere. Pronotum dark brown; strongly convex, disc weakly flattened at center; dark brown; length 1.25 mm, width 1.65 mm, widest at basal 1/3; anterior margin straight except at strongly acute, depressed, anterolateral angles; lateral margins arcuate, sinuate at lateral angles, narrowly margined, fringed with erect setae; posterior border trisinuate; posterolateral angles barely acute, depressed. Disc punctuation similar to that of head, punctures separated by about 1x puncture diameter; setae moderately long. Scutellum: red-brown; subcircular; anterior margin strongly arcuate between two anterolateral notches; disc flat; punctation very fine. Elytron red-brown; length 4.50 mm, width 1.10 mm; broadly convex; narrowest at anterior 1/3, widest at posterior 1/3. Humeral angle rounded; lateral margin widened and sulcate at anterior 1/3, entire length narrowly margined; apex acute. Disc strongly punctate and striate with deep, closely spaced quadrate punctures distinct from near base to apex, intervals weakly convex; setae shorter than on pronotum, semi-erect and recumbent, uniformly distributed. Hind wing: macropterous. Legs. Profemur red-brown, covered with fine, evenly spaced punctures and long, recumbent golden setae; protibia yellow-brown, mostly bare and shiny, dorsal surface with numerous short and a few very long setae, ventral surface with dense row of short, erect setae, arcuate with narrow spines at ventral apex; protarsus yellow-brown, shiny, dorsal surface with a few sparse, very long setae, ventral surface with row of long setae. Mesofemur similar to profemur; mesotibia yellow-brown, mostly bare and shiny, dorsal surface with row of extremely long, golden setae, other surfaces with scattered, short setae, arcuate with spines at ventral apex; mesotarsus similar to protarsus except ventral setae much longer. Metafemur similar to pro- and mesofemora; metatibia straight (in lateral view) with spines at ventral apex, heavily and evenly setose, setae moderately long to very long; metatarsus similar to mesotarsus. Venter red-brown, heavily setose. Prosternum with anterior border narrowly margined; prosternal process wide between procoxae, widest at midlength, convex over procoxae, lateral margins weakly arcuate; prosternal process with median longitudinal carina, bordered laterally by sulci, terminating in an elongate protuberance near apex. Metaventrite with intercoxal process margined and barely depressed; posterior disc depressed at junction of sulcate metakatepisternal suture and metathoracic discrimen. Abdomen with some setae longer than on rest of body except legs; ventrite 1 with triangular intercoxal process thinly margined laterally; ventrites 2���4 of equal length, punctures smaller at center of disc, coarser laterally, aligned with faint, transverse strigae; ventrite 5 longest, disc coarsely punctate, covered with long setae prominent at margins. Genitalia. Aedeagus stout (Fig. 21). Phallobase about 1.5x longer than parameres and wider than paramere bases together. Parameres in dorsal view each laterally sinuate, widest at base, narrowest at midlength, widened at apical 1/4, then narrowed to rounded apex; in lateral view aedeagus strongly curved ventrally, each paramere wide, ventral surface nearly straight, dorsal surface weakly arcuate, paramere tip bluntly rounded. Penis more than 3/4 length of parameres, in dorsal view basal width subequal to paramere at midlength; evenly narrowed from base to broadly rounded apex. Intraspecific variation. Males, 5.40���6.00 mm long (n=8), are smaller than females, 6.15���7.15 mm long (n=14). In both males and females there is variation between individuals in respect to the degree of prosternal process sculpturing. Extremely long mesotibial setae are present in both sexes, although the setation is reduced in two female specimens. Among males, the degree to which the protibiae are curved varies. Some of the specimens are much paler in color and are likely teneral, probably having been collected at lights. Geographic distribution. Known from just a few localities in Bolivia, Paraguay and Venezuela (Fig. 3). Etymology. This new species is named in honor of the Zamuco, also called the Ayoreo, one of the indigenous peoples who live in the northern Gran Chaco region of Bolivia and Paraguay where this species occurs. Comments. The specimens from Venezuela are quite geographically disjunct from those collected in Bolivia and Paraguay (Fig. 3). Regardless, they are indistinguishable based on the male genitalia and external morphology. Females are readily identifiable as this species and the specimens examined have been designated as paratypes., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 122-123, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261

The history and original concept of Parygrus Erichson, 1847, is reviewed, the genus is redescribed, and the status of the type species, P. erichsoni Waterhouse, 1876, is clarified. The four original species are reviewed, with photographs provided for the type specimens of P. angustatus Grouvelle, 1896, P. elateroides Grouvelle, 1896, and P. erichsoni, and a non-type of P. parallelus (Grouvelle, 1890). The holotype of P. parallelus is apparently missing. The female holotype of P. erichsoni is redescribed. An associated male P. erichsoni specimen is compared with the holotype and the male genitalia are described and illustrated. General descriptions and provisional diagnoses are given for P. angustatus, P. elateroides and P. parallelus. Five new species (P. guarani new species, P. lengua new species, P. maya new species, P. quechua new species, P. zamuco new species) are proposed, illustrated, and their geographic distributions mapped. For the first time, the male genitalia of Parygrus species are described and illustrated. A provisional key to the species and an updated key to the Neotropical genera of Dryopidae are provided.

Parygrus quechua Barr and Shepard, new species (Figs 2, 18, 19) http://zoobank.org/ 223E963B-8100-4218-A586-FEFDB83734A8 Type material. Holotype male. BOLIVIA. ��� BOLIVIA, Sta.Cruz / 5kmSSE. BuenaVista, / HotelFlora&Fauna, / W 63��39.13��� S 17��29.92��� // UV& Hg vapor lights / 440m. Feb.6, 2007 / C.W.& L.B.O���Brien // HOLOTYPE / Pa- rygrus quechua / Barr & Shepard [red label, handwritten] // EMEC49558��� (EMEC). Paratypes (9 M). BOLIVIA. BOLIVIA: Santa Cruz / Buena Vista vic. / Flora & Fauna Hotel / 23-25/X/00, R.Morris (1 M, EMEC); BOLIVIA: Santa Cruz / Dpt. 4k SSE B. Vista / 22-25 November 2013 / Wappes & Skillman // Hotel Flora & Fauna / El 350- 400meters / 17��29��� S 63��49��� W (1 M, FSCA); BOLIVIA: Sta. Cruz, / San Osteben [Opispo Santistevan] Prov., / Muyurina, 49 km N // Santa Cruz. XI-3-1959 / 1120 ft. R.B.Cumming // Property of Florida / State Collection of / Arthropods (3 M; FSCA). ECUADOR. 0��40���36������S 76��24���2������W / ECUADOR: Napo, Yasuni Scientific / Station; 13.IV.1998; Coll: K.Will / 210m; Headlamping (in wet area; Will, in litt.) (1M, EMEC); ECUADOR: Napo Prov. / Yasuni Research / Station, 250m / 76��36���W, 0��38���S // 17-31.X.1998 / B.K.Dozier / Collector (1 M, FSCA). PERU. PERU: Madre de Dios / Tambopata Wldlf Res / 30 km SW Pto Moldonado [Maldonado] / 12��50��� S 69��20��� W 290m / 1-14 March 1983 / Joseph J. Anderson (2 M, EMEC). All paratypes also have the following final label: PARATYPE / Parygrus quechua / Barr & Shepard [yellow label, printed]. Additional specimens examined (6 F). BOLIVIA. BOLIVIA: Sta. Cruz, / San Osteben [Opispo Santistevan] Prov., / Muyurina, 49 km N // Santa Cruz. XI-3-1959 / 1120 ft. R.B.Cumming // Property of Florida / State Col- lection of / Arthropods (4 F, FSCA). ECUADOR. ECUADOR: Prov. Napo / ca 10 air km E / Coca, on Rio Napo / 240 m III-8/9-86 / S. McKamey collr. (1 F, EMEC); 0��40���36������S 76��24���2������W / ECUADOR: Napo, Yasuni Scientific / Station; 13.IV.1998; Coll: K.Will / 210m; Headlamping (in wet area; Will, in litt.) (1F, EMEC). Diagnosis. The male genitalia of Parygrus quechua n. sp. are distinctive (Fig. 19); none of the other species (for which the genitalia are known) have parameres with inner teeth at midlength and a phallobase subequal in length to the parameres. Parygrus guarani also has parameres with inner teeth at midlength, but the parameres are less than half as long as the phallobase (Fig. 15). Each of the six females were assigned to this species due to their association with males or on the basis of body size and general morphology, but are not designated as paratypes. Description. Holotype male. Cuticle dark brown, legs, antennae and mouthparts lighter; elongate, moderately flattened; length 5.80 mm (pronotum + elytra), width 2.20 mm; body covered with erect and semi-erect, long, golden setae and much shorter, recumbent setae (Fig. 18). Head dark brown; punctuation moderately fine, dense, punctures separated by about 1x puncture diameter; setae moderately long; vertex with shallow median depression; frons slightly protruding forward between antennal bases, anterior margin arcuate, distance between antennal bases less than length of antennomere 1. Antenna with antennomeres 1 and 2 red-brown, setose, antennomeres 3���11 yellow, densely setose; antennomere 1 and 2 lengths subequal 2; antennomere 2 covering antennomeres 3���5. Clypeus densely setose, coarsely punctate, broadly emarginate. Maxillary palpus with terminal palpomere elongate, subcylindrical, weakly curved and flattened. Labrum emarginate, densely setose, border fringed with long setae. Labial palpus with terminal palpomere subrectangular, slightly flattened; length about 1/2 that of terminal maxillary palpomere. Pronotum dark brown; weakly convex, disc weakly flattened at center; length 1.25 mm, width 1.70 mm, widest at basal 1/3. Anterior margin straight except at strongly acute, depressed, anterolateral angles; lateral margins weakly arcuate, slightly sinuate at posterolateral angles, narrowly margined, fringed with erect setae; posterolateral angles acute, produced, explanate; posterior border trisinuate. Disc punctuation moderately fine, similar to that on head, punctures separated by about 1x puncture diameter; setae moderately long. Scutellum: dark brown; subcircular, slightly wider than long, anterior margin strongly arcuate between two anterolateral notches disc flat; punctation very fine. Elytron dark brown; length 4.55 mm, width 1.10 mm; moderately flattened; nearly parallel-sided, narrowest at basal 1/3, widest at apical 1/3. Humeral angle rounded; lateral margin widened and shallowly sulcate at basal 1/3, entire length narrowly margined; apex acute. Disc strongly punctate and striate with deep, closely spaced punctures distinct from near base to apex, intervals weakly convex; setation similar to that of pronotum except for angulation, uniformly distributed. Hind wing: macropterous. Legs. Profemur dark brown, covered with fine, evenly spaced punctures and long, recumbent golden setae; protibia red-brown, mostly bare and shiny, dorsal surface with sparse, very long setae, ventral surface of apical 1/3 with dense row of short, stiff setae; apical 1/3 strongly deflexed and swollen, ventral apex with granulate carina and narrow spines; protarsus red-brown, shiny, dorsal surface with sparse, very long setae, ventral surface with row of short, erect setae, protarsomeres 4 and 5 each with a few much-longer dorsal setae. Mesofemur similar to profemur; mesotibia red-brown, weakly arcuate with spines at ventral apex; mesotarsus similar to protarsus. Metafemur, and metatarsus similar to those of mesoleg, metatibia straight with spines at ventral apex. Venter dark brown; heavily setose; finely to coarsely punctate. Prosternum with anterior border narrowly margined; prosternal process wide between procoxae, convex over procoxae, lateral margins weakly arcuate, narrowly margined; process with very low, rounded, median longitudinal carina terminating in a low, elongate protuberance near apex. Metaventrite with intercoxal process margined and depressed; posterior disc depressed at junction of sulcate metakatepisternal suture and metathoracic discrimen. Abdomen with some setae longer than on rest of body except legs; ventrite 1 with triangular intercoxal process weakly depressed, narrowly margined laterally; ventrites 2���4 of equal length, finely and evenly punctate at center of disc, coarser laterally, aligned with faint, transverse strigae; ventrite 5 longest, disc coarsely punctate, covered with long setae prominent at margins. Genitalia. Aedeagus moderately elongate (Fig. 19). Phallobase same length as parameres and same width as paramere bases together. Parameres each blade-like, inner surface concave; in dorsal view each with an inward-facing tooth at midlength; in lateral view each paramere wide, ventral surface broadly arcuate, dorsal surface weakly sinuate, paramere tip broadly rounded. Penis nearly as long as parameres; bottle-shaped, bulbous at basal 1/2, narrowest just apical to teeth, subparallel to rounded apex. Intraspecific variation. Males ranged from 5.20���6.00 mm long (n=10); females, 5.50���5.95 mm long (n=6). Body color is quite variable among the examined specimens, ranging from dark brown to light red-brown. The holotype and others are mostly dark brown, whereas many are uniformly medium brown or red-brown, and some have the pronotum darker than the elytra. The lighter-colored individuals are likely teneral, having been collected at lights. Such a specimen is shown in Fig. 18. The eight Bolivian and Peruvian males have protibiae each with the apical 1/3 deflexed, swollen at the apex (Fig. 18), and with a ventral granulate carina; the two Ecuadorian males have the protibiae strongly arcuate rather than deflexed. The protibiae of females are arcuate and without ventral carinae. There is also variability in the shape of the pronotal posterolateral angle (produced or not) and the degree of prosternal process sculpturing. Distribution. Bolivia, Ecuador and Peru (Fig. 2). Etymology. This new species is named in honor of the speakers of the indigenous Quechua languages, the Quechua people, who inhabit the Andean countries of South America where this species occurs., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 119-122, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261

Parygrus lengua Barr and Shepard, new species (Figs 3, 16, 17) http://zoobank.org/ 8995529C-3873-46EC-B9F1-E030A1A07509 Type material. Holotype male. PARAGUAY. ��� PARAGUAY: Alto Paraguay / Puerto La Esperanza / 2 XII 2006 / Carlos Aguilar Julio // HOLOTYPE / Parygrus lengua / Barr & Shepard [red label, handwritten] // EMEC49557��� (EMEC). Paratypes (10 M). PARAGUAY. PARAGUAY: Alto Paraguay / Puerto La Esperanza / 2 XII 2006 / Carlos Aguilar Julio (6 M, EMEC; 1 M, LSAM; 1 M, NHMUK; 1 M, USNM); PARAGUAY: Pdte. [Presidente] Hayes / Cruce los Pioneros / 20 I 1996 // Purchased from / Alfredo Ugarte (1 M, EMEC). All paratypes also have the following final label: PARATYPE / Parygrus lengua / Barr & Shepard [yellow label, printed]. Additional specimens examined (18; 3 M, 15 F). ARGENTINA. ARGENTINA: Entre Rios / Federacion / XII 2006 / Zubaron Gaston, leg. (3 M; EMEC). PARAGUAY. PARAGUAY: Alto Paraguay / Puerto La Esperanza / 2 XII 2006 / Carlos Aguilar Julio (10 F, EMEC; 2 F, FSCA; 1 F, LSAM; 1 F, NHMUK; 1 F, USNM). Diagnosis. The male genitalia of Parygrus lengua n. sp. are distinctive (Fig. 17); none of the other species (for which the genitalia are known) have parameres both with inner ridges and without teeth. An external character not seen in other species is that the scutellum is weakly depressed at the anterior half and convex near the posterior angle. The females were assigned to this species due to their association with the males, their similar scutellar morphology, and body size; they are not designated as paratypes. Description. Holotype male. Cuticle brown, legs, antennae and mouthparts lighter; elongate, subcylindrical; length 4.35 mm (pronotum + elytra), width 1.60 mm; body covered with erect and semi-erect, long, golden setae and much shorter, recumbent setae (Fig. 16). Head dark brown; punctuation fine, dense, punctures separated by at least 1x puncture diameter; setae moderately long; vertex with moderately deep median depression; frons raised and protruding slightly forward between antennal bases, anterior margin arcuate, distance between antennal bases less than length of antennomere 1. Antenna with antennomeres 1 and 2 light brown, setose, antennomeres 3���11 yellow, densely setose; antennomere 1 nearly as long as antennomere 2; antennomere 2 covering antennomeres 3���5. Clypeus densely setose, coarsely punctate, barely emarginate. Maxillary palpus with terminal palpomere elongate, subcylindrical, weakly curved. Labrum emarginate, densely setose, border fringed with long setae. Labial palpus with terminal palpomere subrectangular, slightly flattened; length about 2/3 that of terminal maxillary palpomere. Pronotum dark brown; weakly convex, disc weakly flattened at center; dark brown; length 0.90 mm, width 1.25 mm, widest at basal 1/3. Anterior margin straight except at strongly acute, depressed, anterolateral angles; lateral margins weakly arcuate, barely sinuate at lateral angles, narrowly margined, fringed with erect setae; posterolateral angles acute, explanate; posterior border trisinuate. Disc punctuation slightly coarser than on head; setae moderately long. Scutellum dark brown; subcordate, wider than long; anterior margin weakly arcuate between two anterolateral notches; disc weakly depressed at anterior 1/2, convex near posterior angle; punctation very fine. Elytron red-brown; length 3.45 mm, width 0.80 mm; convex; narrowest at basal 1/3, widest at apical 1/3. Base depressed near margin; humeral angle rounded; lateral margin widened at basal 1/3, narrowly margined; apex acute. Disc strongly punctate and striate with deep, closely spaced punctures distinct from near base to apex, intervals weakly convex; setae slightly shorter than on pronotum, semi-erect and recumbent, uniformly distributed. Hind wing: macropterous. Legs. Profemur red-brown, covered with fine, evenly spaced punctures and long, recumbent golden setae; protibia yellow-brown, mostly bare and shiny, dorsal surface with sparse, very long setae, ventral surface with dense row of short, stiff setae, weakly arcuate with narrow spines at ventral apex; protarsus yellow-brown, shiny, dorsal surface with sparse, very long setae, ventral surface with row of shorter, semi-erect setae, protarsomeres 4 and 5 each with a few much-longer dorsal setae. Mesofemur similar to profemur, mesotibia yellow-brown, ventral setae much longer than those of protibia, weakly arcuate with spinose ventral apex; mesotarsus similar to protarsus except ventral setae much longer. Metafemur and metatarsus similar to those of mesoleg, metatibia straight with spines at ventral apex. Venter red-brown; heavily setose; finely to coarsely punctate. Prosternum with anterior border narrowly margined; prosternal process wide between procoxae, swollen over coxal bases, lateral margins weakly arcuate; process with median longitudinal carina terminating in a low, elongate protuberance near apex. Metaventrite with anterior process margined and depressed between mesocoxae; posterior disc depressed at junction of sulcate metakatepisternal suture and metathoracic discrimen. Abdomen with some setae longer than on rest of body except legs; ventrite 1 with triangular intercoxal process weakly depressed, margined laterally; ventrites 2���4 of equal length, finely and evenly punctate at center of disc, coarser laterally, aligned with faint, transverse strigae; ventrite 5 longest, disc coarsely punctate, covered with long setae prominent at margins. Genitalia. Aedeagus moderately elongate (Fig. 17). Phallobase same length as parameres and same width as paramere bases together. Parameres each blade-like, inner surface concave; in dorsal view narrow, lateral margins nearly straight with a slight inward curve at midlength; in dorsolateral view sclerotized inner dorsal margin with short parallel ridges nearly perpendicular to margin; in lateral view each paramere wide, dorsal and ventral surfaces broadly arcuate, paramere tip acuminate and angled ventrally. Penis nearly as long as parameres, bottle-shaped, base constricted, basal 1/3���1/2 bulbous, apical 1/2 evenly tapered to narrowly rounded apex. Intraspecific variation. The males from Paraguay vary from 3.85���4.45 mm long (n=5); females from 4.50��� 4.70 mm long (n=4). The three male specimens from Argentina are much larger (4.95���5.20 mm long), as are their genitalia, but otherwise differences between the groups are minor. Nevertheless, they have not been designated as paratypes. The morphology of the prosternal process is somewhat variable among all of the males examined. Some have the process swollen over coxal bases and not margined; others are not swollen, but margined and have the longitudinal carina bordered laterally by shallow sulci. Among males, the degree to which the protibiae are curved varies. Distribution. Known from three localities in Argentina and Paraguay (Fig. 3). Etymology. This new species is named in honor of the Lengua, one of the indigenous peoples who inhabited the Gran Chaco of northern Paraguay., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 117-119, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261

Parygrus erichsoni Waterhouse, 1876 (Figs 2, 5, 6, 7) Type material. Holotype female. ���Type [circular off-white label with red border, printed] // Colum/bia [Colombia] 46 /20 [label reverse side] [circular off-white label, handwritten] / Parygrus / Erichsoni, / (Type.) C. Waterh. [Waterhouse][off-white label, handwritten] // NHMUK 010580154 [white bar code label, printed]��� (Fig. 5). Depository NHMUK. In his description, Waterhouse (1876:19 ) simply stated ���Columbia. Brit. Mus.��� Additional specimen examined. Male. COLOMBIA: ���Columbia [Colombia] [circular off-white label, handwritten] // NHMUK 010580155 [white bar code label, printed]��� (Fig. 6). Depository NHMUK. Diagnosis. The male genitalia of Parygrus erichsoni (Fig. 7) are distinctive (among species for which the genitalia are known), although like P. maya n. sp. (Fig. 13) they bear a subapical tooth on the inner margins of each paramere. In the latter species, however, the penis does not closely approach the paramere teeth, while in P. erichsoni the penis nearly reaches the teeth. Also, the paramere tips of P. erichsoni are blunt in lateral view, whereas in P. maya n. sp. they are acute. Externally they are also dissimilar as follows: Parygrus erichsoni (Figs 5, 6) is covered with setae that are long, dark, coarse, and erect on the dorsum; and the pronotum is much wider at base than apex. Parygrus maya n. sp. (Fig. 12) has setae shorter, yellow, finer, and semi-erect; and the pronotum is usually almost as wide at apex as base. Note that this diagnosis of P. erichsoni is based on only two specimens. Redescription. Holotype female. Cuticle dark brown, legs, antennae and mouthparts lighter; elongate, subcylindrical; length 5.75 mm (including head), 5.40 mm (excluding head, pronotum + elytra), 2.10 mm wide; body covered with long, erect, and short, fine, recumbent setae; pronotum slightly narrower than elytra (Fig. 5). Head dark brown; punctation shallow, dense, punctures separated by about 1x puncture diameter; setae moderately long, erect; vertex depressed in middle; frons protruding forward between antennal bases, anterior margin arcuate, distance between antennal bases less than length of antennomere 1. Antenna with antennomeres 1 and 2 red-brown, setose, antennomeres 3���11 yellow-brown, densely setose; antennomere 1 shorter than antennomere 2; antennomere 2 covering antennomeres 3���5. Clypeus barely emarginate, setose, coarsely punctate. Maxillary palpus with terminal palpomere elongate, subcylindrical, weakly curved. Labrum emarginate, coarsely punctate, densely covered with long, erect setae. Labial palpus with terminal palpomere flattened, shorter than terminal maxillary palpomere. Pronotum dark brown; convex, disc slightly flattened at center; length 1.25 mm, width 2.05 mm, widest at basal 1/4���1/3. Anterior border straight except at strongly acute, depressed, anterolateral angles; lateral margins narrowly margined, weakly arcuate, sinuate at moderately explanate, acute, posterolateral angles; posterior border trisinuate. Disc densely punctate, punctures separated by 1x puncture diameter; setae numerous, erect, moderately long. Scutellum: dark brown; subcordate, wider than long; anterior margin arcuate between two anterolateral notches, lateral borders nearly straight; disc flat, punctate. Elytron dark brown; length 4.15 mm, width 1.05 mm; widest at posterior 1/3; depressed at base. Disc strongly striate and punctate, with deep, closely spaced, quadrate punctures separated by less than a puncture diameter, distinct to apex; intervals convex. Semi-erect setae moderately long, widely and evenly spaced; recumbent setae short, closely spaced. Hind wing: macropterous. Legs. Profemur red-brown, covered with fine, evenly spaced punctures and long, recumbent golden setae; protibia weakly arcuate, red-brown, mostly bare and shiny, dorsal surface with sparse, very long setae, ventral surface with dense row of short setae at distal 1/2; protarsus yellow-brown, shiny, dorsal surface with sparse, very long setae, ventral surface with row of short, semi-erect setae, protarsomere 5 with two pairs of longer setae at apex. Mesofemur similar to profemur; mesotibia weakly arcuate, red-brown, ventral setae much longer than those of protibia and more numerous; left mesotarsus similar to protarsus except ventral setae much longer and more numerous [right mesotarsus missing]. Metafemur, metatibia and metatarsus all similar to those of mesoleg. Venter dark red-brown; heavily setose, evenly punctate. Prostenum long; anterior border narrowly margined; prosternal process wide and slightly depressed between procoxae, parallel-sided with thick margins; process with low, median longitudinal carina terminating in a low, oval protuberance near apex. Mesoventrite very short. Metaventrite about as long as prosternum; intercoxal process margined and flat; posterior disc depressed at junction of sulcate metakatepisternal suture and metathoracic discrimen, discrimen angled posteriorly at junction. Abdomen with long setae more dense than on rest of body; ventrite 1 with triangular intercoxal process margined, slightly depressed between metacoxae; ventrites 2���4 of equal length, evenly punctate, punctures aligned with faint, transverse strigae; ventrite 5 longest, punctate, sparsely granulate laterally, disc centrally depressed, apex with small, angular ventrally directed protuberance, covered with long setae prominent at margins. Male specimen. Length 5.25 mm (including head), 5.10 mm (excluding head, pronotum + elytra); 1.85 mm wide; somewhat smaller than the female holotype. The protibia of the male ( Fig. 6) is weakly deflexed at the apical 1/2 and has a ventral, granulate carina, unlike that of the female (Fig. 5), but otherwise is morphologically similar to the female except for the sexual differences exhibited in abdominal ventrite 5. FIGURE 6. Parygrus erichsoni, non-type associated male: dorsal habitus; ventral habitus; lateral habitus; frontal view; specimen labels (specimen images provided by the NHMUK). FIGURE 7. Parygrus erichsoni, male genitalia: a) dorsal view; b) lateral view; c) ventral view. Genitalia. Aedeagus stout (Fig. 7). Phallobase about 1.5x longer and a little wider than paramere bases together. Parameres each blade-like, inner surface concave, in dorsal view each with an inward-facing tooth just before apex; teeth overlapping, apices capable of contacting each other; in lateral view each paramere wide, ventral surface nearly straight, dorsal surface broadly arcuate, paramere tip blunt. Penis 3/4 or more length of paramere, terminating just before paramere teeth; in dorsal view wider than paramere at midlength; evenly tapered to a slightly produced, blunt apex; dorsum longitudinally carinate. Note: The specimen no longer appears as in Fig. 6. In order to dissect the genitalia, the specimen was removed from the original pin and later remounted on a pinned card point along with abdominal ventrites 3���5; the aedeagus is stored in glycerine in a genitalia vial pinned beneath the specimen. Geographic distribution. Known only from Colombia. Because the specific type locality is unknown, the star on the distribution map is positioned mid-country (Fig. 2). Etymology. Waterhouse did not cite an etymology, but the specific epithet undoubtedly honored Dr. W. F. Erichson, the author of the genus and a noted, early German entomologist who specialized in Coleoptera. Comments. Waterhouse (1876) wrote his description of P. erichsoni in Latin followed by an English comparison with the species Parnus prolifericornis Fabricius, 1792 (synonym of Dryops auriculatus (Geoffroy, 1785)). At the end of the Latin description is the line ���Long. 2 �� lin.; lat. 1 lin.��� This is likely the length vs. width body proportions. Waterhouse made no mention of the number of specimens before him. However, the type female and associated male have nearly identical locality labels with the same handwriting and were mounted similarly. Parygrus erichsoni is the type species of the genus., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 104-107, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261, {"references":["Waterhouse, C. O. (1876) III. On various new genera and species of Coleoptera. Transactions of the Entomological Society of London, 1876 (Part I), 11 - 25. https: // doi. org / 10.1111 / j. 1365 - 2311.1876. tb 01122. x","Fabricius, J. C. (1792) Entomologica Systematica emendata et aucta. Secundum Classes, Ordines, Genera, Species. Adjectis Synonimis, Locis, Obervationibus, Descriptionibus. Tome 1. Pars 1. C. G. Proft, Hafniae, 330 pp. https: // doi. org / 10.5962 / bhl. title. 122153","Geoffroy, M. M. & Fourcroy, A. F. (1785) Entomologia Parisiensis; sive catalogus insectorum quae in Agro Parisiensi reperiuntur; secundum methodum Geoffraeanam in sectiones, genera and species distributus: cui addita funt nomina trivalia & fere trecentae novae Species. Pars prima. Via and Aedibus Serpentineis, Parisiis, 231 pp. https: // doi. org / 10.5962 / bhl. title. 36528"]}

Parygrus maya Barr and Shepard, new species (Figs 1, 4, 12, 13) http://zoobank.org/ DCFAFCB8-D6F7-4454-95DA-54A02957EF03 Type material. Holotype male. MEXICO. ��� MEXICO: Quintana Roo / Isla de Cozumel, W side / 4.5km S Chankanaab NP / 12-X-1997 coll. C.B. Barr // shallow roadside / ditch/wet grassy area // HOLOTYPE / Parygrus maya / Barr & Shepard��� [red label, handwritten] (UNAM). Paratypes (78; 43 M, 35 F). BELIZE (17 M, 13 F). BELIZE: Cayo / Dist., Mile 66 / Western Hwy., W.D. Hasse / VII-5-1969 [?] // Property of Florida / State Collection of / Arthropods (1 F, FSCA); BELIZE: Cayo Dist / Belmopan airport / 8.VIII.1993 / at blacklights // William D. / Shepard, leg. (1 M, EMEC); BELIZE: Cayo Dist. / Las Cuevas Field / Station, Millionario / Camp, Tapir Pond / 30-VI-2019, D.L.Post // 16.75275, -89.00840 / elevation 605 m (1 M, 2 F, EMEC); BELIZE: Stann Cr. Dist. / Bocawina ��� Pond / near Silk Grass Creek / 07-VIII-2017, D. L. Post (10 M, 7 F, EMEC; 1 M, 1 F, FSCA; 1 M, 1 F, NHMUK; 1 M, 1 F, USNM); BELIZE: Toledo District: Hickatee / Cottages, 2.6 km W of Punta Gorda / on Ex-Serviceman Rd. 15-21.vi.2008 / 16��05���40������N 88��49���57.9������W 10m, ex: / mercury vapor lamp, F.G. Andrews (2 M, CSCA). COSTA RICA (2 M, 7 F). La Pacifica, 4 Km / N.W. Canas, Costa / Rica, Guanacaste / Prov., VIII-26-71 / P.A.Opler, collr. // U. V. light trap (1 F, EMEC); COSTA RICA: Guanacaste / 8 km NW Filadelfia / 16 VI 2003 280��� / Roadside pool // William D. / Shepard, leg. (1F, EMEC); C.R., Heredia, La / Selva Biol.Sta. 2 Km. / S.Pt.Viejo:VI-3-5-84 / Riley, Rider & LeDoux (1 M, LSAM); C.R., Heredia / 1 Km S. Pt. Viejo / VI-4-5-84: E.Riley / D.Rider & D.LeDoux (3 F, LSAM); COSTA RICA: Heredia / Pr: La Selva Biol.Sta. / 3 km S Pto. Viejo / 10��2���6N 84��01���W // 20. v. 1990 / H.A. Hespenheide (1 F, EMEC); COSTA RICA: Heredia / Est. Biol. La Selva / 03 July 2001 / CL & SL Staines / Taken / at light (1 M, EMEC); COSTA RICA: Heredia / La Selva Biol. St. / 14-15-I-2005 / HG-vapor lights / A.E.Z.Short, leg. (1F, EMEC). GUATEMALA (2 M, 1 F). GUATEMALA: Dept. Iza- / bal, Quirigua Ruins, / near Los Amates, 61m / 4-IX-1976 / Edward S. Ross / Cal. Acad. Sci. Coll. (1M, CASC); GUATAMALA [GUATEMALA]: Izabel [Izabal] / Ruinas Quirigua / VII-5-1977 / E.M.Fisher coll. (1M, CSCA); GUATEMALA: Peten: / Ruinas Tikal / 245m. 7/10 July 77 / E.M. & J.L.Fisher (1 F, CASC). MEXICO (15 M, 14 F). MEXICO: Quintana Roo / 17km NW Felipe / Carrillo Puerto / 17-VI-1990 / coll. M.C. Thomas (1 M, FSCA); MEX: Quintana Roo / 20 Km. N. Felipe Car- / rillo Puerto: VI- / 12-14-83: E.Riley (1 M, 1 F, LSAM); MEXICO: Quintana Roo / Isla de Cozumel, SW side / 2 mi. W Cedral, 6-X-1993 / C.B.Barr & W.D.Shepard // shallow roadside / ditch/wet grassy area (2 M, 2 F, EMEC); MEX- ICO: Quintana Roo / Isla de Cozumel, SW side / 2 mi. W Cedral, 8-X-1993 / C.B.Barr & W.D.Shepard // shallow roadside / ditch/wet grassy area (1 M, 1 F, EMEC); MEXICO: Quintana Roo / Isla de Cozumel, E end / Cross Island Rd., 7-X-1993 / C.B.Barr & W.D.Shepard // shallow / roadside pond (1 F, EMEC); MEXICO: Quintana Roo / Isla de Cozumel, W side / 4.5km S Chankanaab NP / 11-X-1997 coll. C.B. Barr // shallow roadside / ditch/wet grassy area (2 F, EMEC); MEXICO: Quintana Roo / Isla de Cozumel, W side / 4.5km S Chankanaab NP / 12-X-1997 coll. C.B. Barr // shallow roadside / ditch/wet grassy area (3 M, 2 F, EMEC; 1 M, 1 F, UNAM); MEXICO: Quintana Roo / Isla de Cozumel, W side / 4.4km S Chankanaab NP / 13-X-1997 coll. C.B. Barr // shallow roadside / ditch/wet grassy area (4 M, 2 F, EMEC; 3 M, 1 F, UNAM); MEXICO: Veracruz / Experiment Sta. / Cotaxtla / 24.VI.1958 / In light trap (1 F, CASC). NICARAGUA (1 M). NICARAGUA: Granada Prov., Domitila Reserva / Silvestre Privada ESE / Nandaime, elev. 400��� / 11��42.5���N, 85��57.2���W // 9-VI-2005, C.B. Barr / C.S. Chaboo & W.D. / Shepard, collected at / blacklight near river (1 M, EMEC). PANAMA (6 M). PANAMA: Canal Zone / Fort Kobbe: 9 June 1976 / coll: E. G. Riley / black light trap (2 M, UMC); PANAMA: Prov. Cocl�� / 5 rd. km S of Anton / nr. Los Pantones, BL/MV / 29-VIII-2006, C.B.Barr // 08��21.342��� N / 80��16.515��� W / elevation 59 ft. (1M, EMEC); PANAMA: Cocl�� / 5 km S Anton / 29 VIII 06 59��� / Black & MV lights // Doug / Post, leg. (3 M, EMEC; 1M, STRI). All paratypes also have the following final label: PARATYPE / Parygrus maya / Barr & Shepard [yellow label, printed]. Diagnosis. The male genitalia of Parygrus maya n. sp. (Fig. 13) are distinctive (among species for which the genitalia are known), although they are somewhat similar to those of P. erichsoni (Fig. 7) in that they bear a subapical tooth on each paramere. In the latter species, however, the penis is longer, nearly reaches the paramere teeth, while in P. maya n. sp. the penis does not closely approach the teeth. Externally they are also dissimilar as follows: Parygrus maya n. sp. is covered in setae that are moderately long, fine, yellow, and semi-erect on the dorsum; and the pronotum is usually almost as wide at apex as base. Parygrus erichsoni has setae that are longer, coarse, dark, and erect on the dorsum; and the pronotum is wider at base than apex. Description. Holotype male. Cuticle dark brown, legs, antennae and mouthparts lighter; elongate, subcylindrical; length 4.35 mm (pronotum + elytra), width 1.60 mm; body covered with erect and semi-erect, long, golden setae and much shorter, recumbent setae (Fig. 12). Head dark brown; punctuation fine, dense, punctures separated by about 1x puncture diameter; setae moderately long; vertex with shallow median depression; frons raised and protruding forward between antennal bases, anterior margin arcuate, distance between antennal bases less than length of antennomere 1. Antenna with antennomeres 1 and 2 light brown, setose, antennomeres 3���11 yellow, densely setose; antennomere 1 nearly as long as antennomere 2; antennomere 2 covering antennomeres 3���5. Clypeus densely setose, coarsely punctate, barely emarginate. Maxillary palpus with terminal palpomere elongate, subcylindrical, weakly curved. Labrum emarginate, densely setose, border fringed with long setae. Labial palpus with terminal palpomere subrectangular, slightly flattened; length about 2/3 that of terminal maxillary palpomere. Pronotum dark brown; weakly convex, disc slightly flattened at center; length 1.00 mm, width 1.25 mm, widest at middle. Anterior margin straight except at strongly acute, depressed, anterolateral angles; lateral margins nearly parallel, weakly sinuate at lateral angles, narrowly margined, fringed with erect setae; posterolateral angles acute, explanate; posterior border trisinuate. Disc punctuation fine, slightly coarser than on head, punctures separated by slightly less than 1x puncture diameter; setae moderately long. Scutellum: dark brown; subcircular; anterior margin weakly arcuate between two anterolateral notches; disc weakly convex; punctation very fine. Elytron dark brown, length 3.40 mm, width 0.80 mm, convex, narrowest at basal 1/3, widest at apical 1/3. Humeral angle rounded; lateral margin widened and shallowly sulcate at basal 1/3, entire length narrowly margined; apex acute. Disc strongly punctate and striate with deep, closely spaced punctures distinct from near base to apex, intervals weakly convex; setae shorter than on pronotum, semi-erect and recumbent, uniformly distributed. Hind wing: macropterous. Legs. Profemur red-brown, covered with fine, evenly spaced punctures and long, recumbent golden setae; protibia red-brown, mostly bare and shiny, dorsal surface with sparse, very long setae, ventral surface with dense row of short, stiff setae, deflexed and slightly swollen at apical 1/2, ventral apex with ventral granulate carina and narrow spines; protarsus yellow-brown, shiny, dorsal surface with sparse, very long setae, ventral surface with row of short, erect setae, protarsomeres 4 and 5 each with a few much-longer dorsal setae. Mesofemur similar to profemur; mesotibia red-brown, ventral setae much longer than those of protibia; mesotarsus similar to protarsus except ventral setae much longer, weakly arcuate at apical 1/2 with spinose ventral apex. Metafemur, and metatarsus similar to those of mesoleg, metatibia straight with spines at ventral apex. Venter red-brown; heavily setose; finely to coarsely punctate. Prosternum with anterior border narrowly margined; prosternal process wide between procoxae, parallel-sided with thick margins; process with median longitudinal carina, bordered laterally by shallow sulci, terminating in an elongate protuberance near apex. Metaventrite with intercoxal process margined and depressed; posterior disc weakly depressed at junction of sulcate metakatepisternal suture and metathoracic discrimen. Abdomen with some setae longer than on rest of body except legs; ventrite 1 with triangular intercoxal process weakly depressed between metacoxae, thickly margined laterally; ventrites 2���4 of equal length, finely and evenly punctate at center of disc, coarser laterally, aligned with faint, transverse strigae; ventrite 5 longest, disc coarsely punctate, covered with long setae prominent at margins. Genitalia. Aedeagus stout (Fig. 13). Phallobase about 1.5x longer than parameres and a little wider than paramere bases together. Parameres each blade-like, inner surface concave; in dorsal view each with an inward-facing tooth just before apex; teeth overlap with apices not contacting, producing an apical notch; in lateral view each paramere wide, curved ventrally, ventral surface broadly arcuate, dorsal surface sinuate, paramere tip acute. Penis about 2/3 length of parameres, in dorsal view wider at midlength than paramere at midlength; elongate-oval, dorsum strongly longitudinally carinate, apex pointed. Intraspecific variation. Other than size and sexually dimorphic characters, there is very little discernable difference among individuals. There is about 1 mm variation in length among specimens of the same sex, and females are slightly larger than males: males 4.10���4.95 mm long (n=22); females 4.30���5.45 mm long (n=16). Males have deflexed protibiae, each with a granulate carina on the ventral apex; females have protibiae arcuate and lack carinae. In both males and females there is slight variation between individuals in respect to the shape of the pronotal lateral margins, the scutellar shape, and the degree of prosternal process sculpturing. Some specimens are much paler in color and are likely teneral. Geographic distribution. Southern Mexico through Central America (Belize, Guatemala, Nicaragua, Costa Rica, Panama) (Fig. 4). Etymology. This new species is named in honor of the Maya, the indigenous people of northwestern Mesoamerica who inhabit part of the area where this species occurs. Habitat and behavior. At the type locality and two other collection sites on the Isla de Cozumel, Mexico, P. maya n. sp. was found in shallow water in roadside ditches and small ponds with a primary substrate of limestone bedrock. The most productive collecting site was mostly shaded, with cool water and abundant emergent vegetation (Fig. 1). The beetles occurred together with Pelonomus on the submerged stems of emergent plants and could be seen beneath the water surface encased in silvery bubbles of air. One of the beetles captured in a net was released to observe its behavior: It initially floated atop the water for a few seconds, then struggled to break through the surface film in order to submerge itself. After doing so, it crawled upside-down beneath the film until it encountered and grasped the submerged portion of a grass stem. In Panama we collected P. maya n. sp. at black and mercury vapor lights adjacent to a drainage ditch and rice fields. All but one of our collections of P. maya n. sp. have been in conjunction with collections of Pelonomus. Both genera are semiaquatic and appear to share the same types of lentic habitats., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 111-114, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261

Parygrus elateroides Grouvelle, 1896 (Figs 2, 10) Holotype specimen. BRAZIL: ��� Pery-Pery / (Pernambuco) / Gounelle 11.12.1892 [off-white label with black bor- der, printed] // Type [off-white label, printed] // TYPE [red label, printed] // MUSEUM PARIS / 1917 / Coll. Grouvelle [light blue label, printed] // Pargrus [sic] elateroides / ty. Grouv [off-white label, handwritten]���. Depository, MNHN. The type specimen itself was not examined, only the photographic image and label data from the photo. Additional specimens examined (2). BRAZIL: ���80.43 // Parnus / elateroides [illegible letters] / Braz. [handwritten] // [red label with no writing]��� (abdomen missing); ���Brazil. / Castlenau [Castelnau?][on mounting card] // Sharp Coll. / 1905-313 [printed] // Pelonomus / elateroides [illegible letters] / Bras.��� [blue label, handwritten]���. Depository NHMUK. These may not be conspecific with the type or with each other; see Discussion below. Provisional diagnosis. The explanate lateral margins of the pronotum and small size (about 5 mm long), serve to separate P. elateroides (Fig. 10) from the other species of Parygrus. The only other species with an explanate pronotum is P. parallelus (Fig. 11), which is 7���8 mm long. General description. Since we were unable to examine the holotype, only a limited description is possible based on the original descriptions (Grouvelle 1896b) and images of the type (Fig. 10). From a translation of the original Latin and French descriptions: 5 mm long; convex; pronotal lateral border raised in a thin rim; scutellum oval, acuminate apically; elytra wider than pronotum, fairly strongly punctate-striate. To this we add that the prono- tum is quite convex and deeply punctate, widest at the base, lateral margins are arcuate and moderately explanate, and posterolateral angles broadly explanate. Distribution. Brazil, described from the Province de Pernambuco (Fig. 2). Etymology. Grouvelle did not cite an etymology, but the name probably refers to the resemblance of the species to an elaterid or click beetle. The ending ���oides, from the Greek ���eides and - eidos, means ���like, resembling, having the form of.��� Comments. Grouvelle (1896b) described the species in both Latin and French, the French description being the more detailed. The specimen was noted as being 5 mm long, however, measurement of the type image using the included scale indicates it is closer to 5.5 mm long. On the determination label of the holotype the genus is misspelled as ���Pargrus��� (Fig. 10). In his description of Parygrus elateroides, Grouvelle (1896b, p. 34) stated the type locality as ��� Br��sil: Province de Pernambuco (Pery-Pery). ��� Collection Gounelle.��� He reported that he had seen specimens of P. elateroides from S. Paul (Br��sil) belonging to the Fry Collection, that were larger and more stocky. We examined two specimens labeled as P. elateroides from the NHMUK, cited above in Additional specimens examined, however, neither of these has Fry Collection labels. Without actually examining the type specimen we are unable to confirm these determinations and, in fact, suspect they are incorrect. At 7.5 and 8.5 mm long the specimens are at least 2.0���3.0 mm longer than the type, and one of them does not resemble the type image. In addition, they are likely not conspecific due to morphological dissimilarities (i.e., puncto-striation, setation, scutellar shape)., Published as part of Barr, Cheryl B. & Shepard, William D., 2020, Hiding in plain sight: rediscovery and review of Parygrus Erichson, 1847, with description of five new species from the Neotropics (Coleoptera: Byrrhoidea Dryopidae), pp. 99-128 in Zootaxa 4755 (1) on pages 108-110, DOI: 10.11646/zootaxa.4755.1.4, http://zenodo.org/record/3724261, {"references":["Grouvelle, A. (1896 b) Note VI. Descriptions de Dryopides (Parnides) et Helmides nouveaux. Notes from the Leyden Museum, 18, 33 - 52."]}

2007-2009 yılları arasında Afyon, Denizli, Kütahya, Uşak ve 2019 yılında Burdur, Isparta illerinde gerçekleştirilen arazi çalışmaları sonucu Coleoptera takımına ait Dryopidae, Elmidae, Heteroceridae ve Hydraenidae familyalarından 1330 örnek toplanarak değerlendirilmiştir. Teşhis işlemleri sonucunda, 4 familyadan 10 cinse ait 34 tür ve 6 alttür tespit edilmiştir. Bu taksonlardan bilim dünyası için yeni olduğu değerlendirilen Hydraena cinsine ait 3 yeni tür ilk kez bu çalışma ile tanımlanmıştır. Çalışmada ele alınan taksonlardan 19 takson (Dryops caspius (Menetries, 1832), D. lutulentus (Erichson, 1847), D. rufipes (Krynicki, 1832), D. similaris Bollow, 1936, D. sulcipennis Costa, 1883, Heterocerus obsoletus Curtis, 1828, Elmis maugetii maugetii Latreille, 1802, E. rioloides (Kuwert, 1890), Grouvellinus caucasicus (Motschulsky, 1839), Limnius opacus opacus P. W. J. Müller, 1806, L. perrisi perrisi (Dufour, 1843), L. satanus Jäch, 1982, Riolus cupreus (P. W. J. Müller, 1806), R. nitens (P.W.J. Müller, 1817), Hydraena attaleiae Ferro, 1984, H. cata Orchymont, 1943, H. schoenmanni Jäch, 1988, Ochthebius maxfischeri Jäch, 1999, O. viridis viridis Peyron,1858) çalışma bölgesi için yeni kayıttır. Dryops similaris Bollow, 1936 türüne ait Denizli'den verilen kayıt bu tür için ülkemizden verilmiş ilk kesin lokalite kaydıdır. Çeşitlilik indeksleri kullanılarak yapılan değerlendirme sonucunda araştırma alanında Isparta Yazılı Kanyon Tabiat Parkı ve Denizli Süleymanlı Gölü'nün tür çeşitliliği açısından en zengin istasyonlar olduğu tespit edilmiştir., In this study, 1330 samples of Dryopidae, Elmidae, Heteroceridae and Hydraenidae families collected from Afyon, Denizli, Kütahya and Uşak provinces between 2007-2009, Burdur and Isparta provinces in 2019 were evaluated. As a result of the diagnosis procedures, 34 species and 6 subspecies of 10 genera belonging to 4 families were identified. Among these taxa, 3 new species belonging to the Hydraena genus were identified for the first time with this study and 19 taxa (Dryops caspius (Menetries, 1832), D. lutulentus (Erichson, 1847), D. rufipes (Krynicki, 1832), D. similaris Bollow, 1936, D. sulcipennis Costa, 1883, Heterocerus obsoletus Curtis, 1828, Elmis maugetii maugetii Latreille, 1802, E. rioloides (Kuwert, 1890), Grouvellinus caucasicus (Motschulsky, 1839), Limnius opacus opacus P. W. J. Müller, 1806, L. perrisi perrisi (Dufour, 1843), L. satanus Jäch, 1982, Riolus cupreus (P. W. J. Müller, 1806), R. nitens (P.W.J. Müller, 1817), Hydraena attaleiae Ferro, 1984, H. cata Orchymont, 1943, H. schoenmanni Jäch, 1988, Ochthebius maxfischeri Jäch, 1999, O. viridis viridis Peyron,1858) are new records for the study area. The record from Denizli is the first definite locality record of the species Dryops similaris Bollow, 1936 from Turkey. As a result of the evaluation made by using diversity indexes, it was determined that Isparta Yazılı Canyon Nature Park and Denizli Süleymanlı Lake are the richest stations in terms of species diversity.

Platyparnus gen. n. is erected to include three species of Helichus Erichson, 1847, from the Guianas of northern South America. Proposed taxonomic reassignments are Platyparnus bollowi (Hinton, 193...

An annotated list of the aquatic Polyphaga (Coleoptera: Helophoridae, Georissidae, Hydrochidae, Spercheidae, Scirtidae, Elmidae, Dryopidae, Limnichidae and Heteroceridae) recorded from Egypt is presented, based on data gathered from earlier literature. The checklist includes data on the type localities, references in which the species are described in detail, world and Egyptian distributions, and previous literature, for 21 valid species belonging to 13 genera: Helophorus (Helophoridae); Georissus (Georissidae); Hydrochus (Hydrochidae); Spercheus (Spercheidae); Contacyphon (Scirtidae); Oulimnius and Potamodytes (Elmidae); Dryops and Parahelichus (Dryopidae); Bothriophorus and Pelochares (Limnichidae); Augyles and Heterocerus (Heteroceridae). The present study provides a summary that can provide a basis for future progress in the knowledge of the Egyptian aquatic Coleoptera.

The byrrhid subfamily Lidryopinae subfam. nov. is proposed for Lidryops occultus gen. et sp. nov. described from Lower Cretaceous “opaque” amber originated from Archingeay-Les Nouillers, Charente-Maritime, southwest France which was studied using propagation phase contrast X-ray synchrotron microtomography at the European Synchrotron Radiation Facility (Grenoble). This byrrhid subfamily differs from members of the closely related groups (Chelonariidae, Limnichidae, Dryopidae, Elmidae, Lutrochidae) in the set of characters including the moderately convex dorsum, the subpectinate antennae, the five-segmented tarsi with strongly lobed tarsomeres 1–4, subsecuriform ultimate maxillary and labial palpomeres. An assumption on probable lifestyle of Lidryops occultus gen. et sp. nov. is proposed on the basis of structural peculiarities. A short review of the byrrhid fossil record and its analysis are made.

The Comal Springs dryopid beetle, Stygoparnus comalensis Barr and Spangler, and the Comal Springs riffle beetle, Heterelmis comalensis Bosse, Tuff, and Brown, are protected by the United States Fish and Wildlife Service (USFWS), and the development of a captive self-propagating refuge is of importance to stakeholders within the Edwards Aquifer. Being able to reliably distinguish the sex of living subjects is desirable for establishing a successful refuge program. Ventrite, elytron, and pronotum measurements of S. comalensis were taken to determine if there were sexually dimorphic allometries. Various lighting techniques were also implemented to see if there were other characters that could potentially be used to distinguish females and males. Measurements were not found to satisfactorily separate sexes; however, lateral lighting was found to consistently illuminate internal abdominal structures of S. comalensis where sternite 8 was viewable in males and the fused gonocoxites were viewable in females. Lateral lighting was used to examine living specimens of H. comalensis, and it was found that sternite 8 could be viewed in both sexes where the anterior strut of females was much longer and discernible from the anterior strut of males. Commentary regarding the use of cameras and photography for observing living subjects is given.

Dryops lutulentus (Erichson, 1947) Material examined. Loc. 1: 1 ♀; Loc. 3: 1 ♀; Loc. 11: 1 ♁. Notes. New record for Oman. Dryops lutulentus has been recorded from the province of Hormozgan in Iran (DELÈVE 1970; MASCAGNI et al. 2016), Yemen, Saudi Arabia and UAE (MASCAGNI 2009a) (Table 2)., Published as part of Ribera, Ignacio, Hernando, Carles & Cieslak, Alexandra, 2019, Aquatic Coleoptera of North Oman, with description of new species of Hydraenidae and Hydrophilidae, pp. 253-272 in Acta Entomologica Musei Nationalis Pragae 59 (1) on page 265, DOI: 10.2478/aemnp-2019-0021, http://zenodo.org/record/4488950, {"references":["DELEVE J. 1970: Contribution a la faune de l'Iran, 19, Coleopteres Dryopoidea. Annales de la Societe Entomologique de France (Nouvelle Serie) 6: 701 - 703.","MASCAGNI A., JACH M. A., OSTOVAN H. & GHAHARI H. 2016: Annotated checklist of Dryopidae and Heteroceridae (Coleoptera: Byrrhoidea) of Iran. Zootaxa 4144: 354 - 364.","MASCAGNI A. 2009 a: Order Coleoptera, family Dryopidae. Pp. 149 - 150. In: HARTEN A. VAN (ed.): Arthropod Fauna of the United Arab Emirates. Volume 2. Multiply Marketing Consultancy Services, Abu Dhabi, 786 pp."]}

Data from the literature and from specimens preserved at the Colección Nacional de Insectos (CNIN) Instituto de Biología, UNAM were used to compile a checklist of the fauna of Coleoptera of the state of Morelos, México. A sum of 70 families, 167 subfamilies, 361 tribes, 1,022 genera, and 2,606 species are recorded; from this 24 species are new records for Morelos. The State of Morelos ranks fourth in Coleoptera species richness for Mexico, following Veracruz (3,176 spp.), Oaxaca (2,148 spp.) and Chiapas (1,734 spp.). The checklist presented here provides a summary that can serve as a basis for future progress in the knowledge of Mexican Coleoptera.

The flight wing shapes of 4,046 specimens representing 62 genera of Elmidae and 21 genera of Dryopidae were examined. Within Elmidae, non-macropterous wings were found only in the subfamily Elminae, while the subfamily Larainae had only macropterous wings. Within Dryopidae, only macropterous wings were present in almost all aquatic or semiaquatic genera, while only non-macropterous wings occurred in almost all terrestrial genera. Thus, flight wing shapes are generally a good indicator of both habitat and phylogenetic relations.

We report the aquatic Coleoptera (families Dryopidae, Dytiscidae, Georissidae, Gyrinidae, Heteroceridae, Hydraenidae, Hydrophilidae and Limnichidae) from North Oman, mostly based on the captures of fourteen localities sampled by the authors in 2010. Four species are described as new, all from the Al Hajar mountains, three in family Hydraenidae, Hydraena (Hydraena) naja sp. nov., Ochthebius (Ochthebius) alhajarensis sp. nov. (O. punc-tatus species group) and O. (O.) bernard sp. nov. (O. metallescens species group); and one in family Hydrophilidae, Agraphydrus elongatus sp. nov. Three of the recorded species are new to the Arabian Peninsula, Hydroglyphus farquharensis (Scott, 1912) (Dytiscidae), Hydraena (Hydraenopsis) quadricollis Wollaston, 1864 (Hydraenidae) and Enochrus (Lumetus) cf. quadrinotatus (Guillebeau, 1896) (Hydrophilidae). Ten species already known from the Arabian Peninsula are newly recorded from Oman: Cybister tripunctatus lateralis (Fabricius, 1798) (Dytiscidae), Hydraena (Hydraena) gattolliati Jäch & Delgado, 2010, Ochthebius (Ochthebius) monseti Jäch & Delgado 2010, Ochthebius (Ochthebius) wurayah Jäch & Delgado, 2010 (all Hydraenidae), Georissus (Neogeorissus) chameleo Fikáček & Trávníček, 2009 (Georissidae), Enochrus (Methydrus) cf. tetraspilus Régimbart, 1903, Laccobius (Hydroxenus) leucaspis Kiesenwetter, 1870, Paracymus relaxus Rey, 1884 (all Hydrophilidae), Dryops lutulentus (Erichson, 1947) (Dryopidae), and Augyles flavidus (Rossi, 1794) (Heteroceridae). A total of 73 species of aquatic Coleoptera are presently known from Oman, although this number is expected to increase substantially with further sampling. A checklist of the species of aquatic Coleoptera from the Arabian Peninsula is provided., We also thank Rocío Alonso and Anabela Cardoso (IBE) for laboratory work, and funding from projects CGL2007-61665, CGL2013-48950-C2-1-P and CGL2016-76705-P (AEI/FEDER, UE).

The diversity of the aquatic and strictly riparian beetles of the Philippines is reviewed. A checklist (with 317 species and subspecies) of the aquatic and riparian Coleoptera of the Philippines is provided for the first time herein. This checklist covers the following families (number of species and subspecies in parentheses): Hydroscaphidae (1), Gyrinidae (15), Dytiscidae (65), Noteridae (4), Haliplidae (2), Hydraenidae (15), Hydrochidae (2), Spercheidae (1), Hydrophilidae (72), Scirtidae (49), Elmidae (23), Dryopidae (2), Psephenidae (16), Eulichadidae (1), Limnichidae (20), Heteroceridae (5), Malachiidae (5), Lampyridae (15), Nitidulidae (4). A total of 201 species and subspecies (63%) are considered to be endemic to the Philippines. Aquatic/riparian beetle genera and families that are still unrecorded but likely to occur in this country are briefly discussed. Based on estimates at the genus level, it is concluded that about 850 aquatic and riparian species can be expected to occur in the Phil...