Zootaxa 3753 (2): 187–195 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press
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ISSN 1175-5326 (print edition)
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http://dx.doi.org/10.11646/zootaxa.3753.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:3161924A-23A6-4FB7-83FD-8614D4C36D85
Two new Larainae species from Guayana region, Venezuela (Coleoptera: Elmidae) KRISTÍNA LAŠŠOVÁ1, FEDOR ČIAMPOR Jr & ZUZANA ČIAMPOROVÁ-ZAŤOVIČOVÁ Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, SK-84506, Bratislava, Slovakia. E-mail:
[email protected],
[email protected],
[email protected] 1 Corresponding author
Abstract Two new species of the subfamily Larainae (Insecta: Coleoptera: Elmidae), Hexanchorus angeli n. sp. and Hypsilara autanai n. sp., are described from Guyana region in Venezuela. We provide habitus photographs, detail drawings of both male and female genitalia, and description of morphological features important for discrimination of the new species. Molecular differences within genera were measured using 816bp fragment of mtDNA gene for cytochrome oxidase c subunit I. Sequence divergences among species are discussed. Key words: Coleoptera, Larainae, Elmidae, Hexanchorus, Hypsilara, Venezuela, DNA
Introduction The Elmidae fauna in Venezuela is very rich regarding number of species and also the density of the beetles in the streams. Although large areas have already been destroyed by deforestation, Venezuela still provides various preserved aquatic biotopes, which together with its tropical climate predetermine this country to be home of vast number of elmid species. They occur not only in lowlands, but also in the specific habitats like table mountains (Aubrecht et al. 2012). The Guyana region, with its bizarre isolated flat-topped table mountains is known for its huge diversity and high level of endemism. This concerns also aquatic insects (e.g. Čiampor Jr et al. 2013, Kodada et al. 2012, Kodada & Jäch 1999, Derka 2002, Derka et al. 2009). However, contrasting with an expected high diversity, only some 61 species of Elmidae are recorded from whole Venezuela (Segura et al. 2013). The Larainae, which are characteristic by semiaquatic life cycle, form smaller subfamily of Elmidae with 28 described genera. Ovipositing and larvae development takes place in the water, while adults occur often above the water line on partly submerged wood or in spray zones of waterfalls and cascades. The genus Hexanchorus was described by Sharp (1882). Spangler & Staines (2003) described three new species, based on the revision of older museum material, and the most recent work including description of new species of Hexanchorus is Maier 2013, raising the number of species to nineteen. Now, the genus is known throughout the whole Central and South America, from Mexico to Argentina. Except the new species described herein, six other Hexanchorus species occur in Venezuela (Maier 2013). The genus Hypsilara was described only recently (Maier & Spangler 2011). Currently, the genus includes two species found only in Venezuela (Čiampor Jr et al. 2013). In this paper we describe two new species, Hexanchorus angeli n. sp. and Hypsilara autanai n. sp., and we also provide comparison with related congeneric species based on morphological and molecular characters.
Material and methods Specimens prepared for the study were cleaned and examined under a Leica M205C stereomicroscope with a Planapo 1.0 lens, by using diffuse lighting at magnifications up to 160×. Male genitalia and pregenital segments
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were studied as temporary glycerine slides at magnifications up to 600× by using a Leica DM1000 light microscope. Drawings were made with a drawing tube. Habitus photographs were made using Leica M205C with Nikon D3s digital camera attached. Morphological terms generally follow Kodada & Jäch (2005). Metric characters were measured with an ocular grid to nearest 0.05 mm. Abbreviations used in the text: CL— body length (measured from anterior margin of pronotum to elytral apices), EL—elytral length, EW—maximum elytral width, PL—pronotal length, PW—maximum pronotal width, ID—interocular distance, NMW—Natural History Museum Vienna, CCB—collection of Fedor Čiampor Bratislava, MIZA—Museo del Instituto de Zoología Agrícola, Facultad de Agronomía, Universidad Central de Venezuela, Maracay, ZSM—Zoologische Staatssammlung München. For the analysis of DNA, several specimens of Hypsilara and Hexanchorus were used. Additionaly, few samples representing other neotropical elmid genera were also included (see Fig. 22). DNA was isolated from the whole specimens by the DNeasy Blood & Tissue Kit (Qiagen, Hilden Germany) according to the manufacturer´s protocol. 618bp fragment of the mitochondrial gene for cytochrome oxidase c subunit I (cox1) was amplified using primers Pat (CAA CAT TTA TTT TGA TTT TTT GG) and Jerry (TCC AAT GCA CTA ATC TGC CAT ATT A) (Simon et al. 1994). Amplification products were purified and sequenced in Macrogen Europe Inc. (Amsterdam, Netherlands) and subsequently assembled and edited in Sequencher v.5.0.1. The measure of the genetic distance (uncorrected p-distance) was calculated in MEGA software v.5.2.1 (Tamura et al. 2011). Vouchers are deposited in CCB.
Results Hexanchorus angeli n. sp. Figs 1–14 Type locality: environment of the Río Churun, large stream ca. 15 m wide and mostly about 0.5 m deep with orange-brown coloured humic water (specimens were collected at light). Type material: holotype male (NMW): ”Venezuela, Auyán Tepui env., Salto Angel camp, Río Churun env., at light, 506 m a.s.l., 05°58'35.5'' N, 062°30'58.1'' W, 5.12.2012”, paratypes (CCB, MIZA, NMW, ZSM): 11 specimens with the same locality data as holotype. Diagnosis. Hexanchorus angeli n. sp. is externally similar to all other known Hexanchorus species. It differs in combination of the following characters: 1) presence of a greenish iridescence on dorsal surface, 2) without median longitudinal sulcus at base of pronotum, 3) elytral apices feebly (males) or distinctly (females) upturned, 4) third ventrite of females with distinct median process, 5) ventrite 5 slightly emarginated apically in males, 6) aedeagus with median lobe short, narrow apically, parameres long abruptly constricted basally. Description. Body elongate, subparallel (Figs 1, 2); length 3.16 mm; width 1.15 mm; dorsum moderately convex, dorsal surface with very dense short recumbent setae with greenish iridescence and more sparse, longer, dark, semierected setae. Body black, scape and pedicel dark brown, maxillae, trochanters and basal portion of femora pale brown, tarsal claws reddish-brown. Venter dark with light yellowish-brown setae. Head. Head partly retractable into prothorax. Labrum densely setose, with largely rounded anterior angles, feebly emarginate anteromedially; clypeus densely setose, shorter and narrower than labrum, about 3.0 times as wide as long, frontoclypeal suture visible, almost straight. Eyes well developed, suboval in lateral view, protruding from head outline, bordered with long black curved setae arising near dorsal and ventral side and extend toward middle of eye, cranial surface not raised around eyes. Frons between eyes flat. Antennae eleven segmented, densely setose, scape curved, about 2.0 times as long as pedicel, remaining segments about three times longer than scape and pedicel combined, segments 3–10 short, subtriangular, terminal segment subglobular with slightly pointed apex. Thorax. Pronotum widest behind middle, PW: 0.88 mm, PL: 0.78 mm; partly shiny with dense smaller and larger setigerous punctures; disc convex, divided by broad transversal U-shaped depression in the first third; triangular median depression at base shallow, reaching basal third of pronotum. Sides of pronotum convex in posterior 2/3, slightly constricted before base, feebly convex in anterior third before transverse constriction; lateral margins narrowly rimmed, posterior angles orthogonal, anterior angles not produced. Lateral sides decliving to the
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ventral side. Hypomeron narrow, sinuate. Prosternal process narrow, triangular, with apex rounded; prosternum in front of coxae extremely narrow, reduced. Mesoventrite short and wide, disc narrowed posteriad, posteromedially with deep triangular impression for reception of prosternal process. Metaventrite wide, metaventral discrimen thin, present along basal 2/3; disc raised on sides, surface reticulate and densely setose (Fig. 3). Elytra (EL: 2.42 mm, EW: 1.15 mm), in anterior 2/3 parallel, then tapering toward apices, disc moderately convex, sides strongly declivous; surface glabrous with very dense short setae with greenish iridescence and sparser longer semierect hairs; elytral margin only very narrowly rimmed; humeri rounded, slightly produced; epipleuron widest in anterior third, then reduced, very thin. Each elytron with ten rows of small punctures, striae impressed on disc, impression diminishes toward lateral margin and elytral apex. Scutellum wide, subtriangular, anterior portion flat, apical triangular part raised. Legs moderately long, mesotibia markedly flattened, longest.
FIGURES 1, 2. Habitus of Hexanchorus angeli n. sp.: 1) holotype, male, 2) paratype female.
Abdomen. Ventrites densely setose (Fig. 3). Intercoxal process of first ventrite with apex rounded, lateral sides not raised, sublateral carinae absent; distal margins of first and second ventrite feebly sinuate; third ventrite simple in males, in females with distinct median process (Fig. 12). Ventrite 5 with distal margin emarginated in males (Fig. 7), simple in females (Fig. 10). Sternite 8 weakly sclerotized and reduced as two darkened spots in males (Fig. 8), deeply emarginate in females (Fig. 11). The ninth segment and spiculum gastrale extremely elongate (Fig. 6). Aedeagus (Figs 4, 5) elongate, penis with fibula feebly sclerotized, better visible from lateral view, corona membranous; parameres slightly shorter than penis, in lateral view widest in middle, moderately narrowed in basal half, distal half moderately narrowed toward rounded apex, in ventral view moderately narrowed in apical ca. 0.4, in basal 0.6 parallel-sided, base constricted; penis in ventral view subparallel in basal 0.6 with distinct basal apophyses, apical 0.4 narrowed toward thin rounded apex, in lateral view slender, sinuate, with widened basal 2/3. Phallobase parallelsided, curved in lateral view. Penis and parameres with sparse fine spines. Ovipositor (Fig. 9) with stylus (terminal segment) short, slightly widened basally; preterminal segment short and robust, about 2.7 times as long as stylus, apically with numerous curved blunt and acuminate spines or sensilla; basal segment with baculus membranous, ca. 0.8 times as long as preterminal and distal segments combined, baculus straight, well sclerotized.
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FIGURE 3. Hexanchorus angeli n. sp. male and female ventral view.
Sexual dimorphism. Both sexes are generally similar in shape and size, however males are usually slightly smaller. From females, males can be easily distinguished by elytral apices not produced, by ventrite 3 without distal process, by shape of distal margin of ventrite 5, by ventrites 1–2 concave, not convex. Pronotum of males with feeble median triangular depression in basal third, medio basal portion of female pronotum without visible depression. Distribution. Hexanchorus angeli n. sp. is known only from the type locality. Remarks. One female of Hexanchorus was found also in Río Tarota (Gran Sabana), more than 120 km distant from the type locality of H. angeli n. sp. This female is morphologically very similar to the females of the species described herein. However, observed 4 % molecular distance, measured as uncorrected p-distance on 816bp long fragment of the mitochondrial gene for cytochrome oxidase c subunit I (tree not shown), suggest that the female from the Gran Sabana should belong to another species. Habitat. The specimens were caught on light, so we can only suggest that H. angeli n. sp. adults live in and around rapids of the Río Churún, as do most of other related Larainae species, and their larvae develop in the same river or their tributaries. At the place of collecting, the river was about 15 m wide, relatively shallow. Substrate was formed by gravel with large stones and boulders weekly covered by periphyton (Figs. 13, 14). Water was naturally humic, brown coloured.
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Etymology. The species is named after Jimmie Angel, an American aviator who was the first man flying over the Angel falls. These waterfalls are the highest in the World, falling from the slopes of the Auyán tepui close to the type locality.
FIGURES 4–12. Hexanchorus angeli n. sp.: 4) aedeagus lateral view; 5) aedeagus ventral view; 6) male ninth segment and spiculum gastrale; 7) ventrite 5 male; 8) sternite 8 male; 9) ovipositor; 10) ventrite 5 female; 11) sternite 8 female; 12) ventrite 3 female. Scale bars: 0.1 mm.
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FIGURES 13–16. Type localities: 13, 14) Río Churún, type locality of Hexanchorus angeli n. sp.; 15, 16) small stream near Cerro Autana, type locality of Hypsilara autanai n. sp.
Hypsilara autanai n. sp. Figs 15–21 Type locality: stream ca. 2–3 m wide, situated at the foot of a mountain, quickly flowing in a dense forest with cascades, riffles and pools, with stones, roots and moss. Type material: holotype male (NMW): ”Venezuela, Amazonas Province, Cerro Autana env., stream ca. 2–3m wide, with stones, roots and moss, 131 m a.s.l., 04°49'33.6'' N, 067°28'55.4'' W, 23.11.2012, Čiampor Jr & Čiamporová-Zaťovičová lgt.”, paratypes (CCB, NMW, MIZA, ZSM): 2 specimens with the same locality data as holotype; 7 specimens ”Venezuela, Amazonas Province, Cerro Cuao env., small stream in primary forest, ca. 1m wide, with large boulders, fine gravel, moss and fallen wood, 923 m a.s.l., 05°05'39.1'' N 067°27'41.4'' W, 19.11.2012”. Diagnosis. Hypsilara autanai n. sp. resembles externally both known species. It differs in smallest size, less convex dorsum, pronotal sublateral grooves almost straight, pronotal median longitudinal groove short but deeply impressed, narrower elytra, elytral apices not produced, metaventrite of males without admedian tufts of darkened setae and different shape of aedeagus as follows: a) phallobase ca. 0.63 times as long as penis, b) parameres ca. 0.8 times as long as penis, c) parameres narrowly fused at base, d) penis narrowly triangular in ventral or dorsal view. Hypsilara autanai n. sp. is also well characterized by differences in cox1 gene. Description. Body elongate (Fig. 17), CL: 3.67 mm, ca. 2.26 times longer than wide (CL/EW), dorsum feebly convex, dorsal surface with very dense short recumbent setae and more sparse longer, dark, semierect setae. Coloration of head dark brown, antenna paler. Body dark brown dorsally, venter reddish-brown, coxa and trochanter light reddish-brown, tarsus brown, tibia brown dorsally, ventrally black. Head. Head partly retractable into thorax. Labrum short, partly concealed by clypeus, dark brown, with large punctures and longer fine setae, lateral angles rounded; clypeus longer and wider than labrum, about 2.0 times as wide as long, densely setose; frontoclypeal suture visible, straight. Eyes well developed, slightly protruding from head outline, suboval in lateral view, bordered with long setae along upper margin, cranial surface not raised near dorsal margin of eyes. Antennae eleven segmented, scape long and curved, pedicel shorter, both densely setose, remaining segments forming club, nearly as long as scape and pedicel combined. Thorax. Pronotum widest behind middle, PW: 1.20 mm, PL: 0.89–0.96 mm; disc moderately convex, with deep, short median longitudinal groove in posterior half (not reaching posterior margin), widely V-shaped transversal depression before middle and pair of deep admedian prebasal punctures; sublateral grooves almost straight, well developed from base, reaching middle of pronotum; sides of pronotum moderately convex in
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posterior 2/3, subparallel in anterior third; lateral margins explanate (wider anteriorly); anterior angles rounded, slightly protruding anteriad; posterior angles nearly orthogonal (Fig. 17). Hypomeron subparallel, slightly narrowed posteriad, margin dark brown to black, anterior depression feebly depressed. Prosternal process 1.5 times as long as prosternum in front of coxae, narrowly triangular, lateral margins slightly rounded, apex rounded. Surface of prosternum with sparse shallow punctures, setose, in males with tuft of longer dark brown setae medially behind anterior margin. Mesoventrite short and wide, with large and deep depression for reception of prosternal process; portions between coxae with distinct transverse darkened carina. Metaventrite with median suture thin and darkened, developed from anterior 1/3 to posterior margin; disc flat, densely pubescent, lateral sides with sparse larger punctures, prebasally with row of punctures along metacoxa. Elytra ca. 2.70 mm long (EL), 1.61 mm wide (EW); subparallel in anterior 0.57, then tapering toward rounded apices; disc slightly convex, posterior third and lateral margins strongly declivous, densely pubescent; elytral margin dark brown to black; humeri rounded; epipleuron glabrous, widest anteriorly, narrowed posteriad. Each elytron with ten rows of punctures, punctures larger and deeper on disc, striae 3 and 4 fused before apex. Scutellum flat, wider than long, with pointed apex and sides arcuate. Legs moderately long, tibiae not flattened, hind tibiae longest; metacoxae transverse prebasally with distinct short transverse carina.
FIGURES 17–21. Hypsilara autanai n. sp.: 17) habitus of holotype (male); 18) aedeagus ventral view; 19) aedeagus dorsal view; 20) aedeagus lateral view; 21) ovipositor. Scale bars (FIGURES 18–21): 0.1 mm.
Abdomen. Intercoxal process of ventrite 1 triangular, longer than wide, lateral sides raised, continuing in carinae reaching posterior margin of ventrite, apex subacute; ventrite 5 feebly pointed, margins of ventrites narrowly darkened. Aedeagus elongate, fibula absent, corona membranous; parameres ca 0.8 times as long as penis, widest at the base, narrowed toward narrowly rounded apices, basally fused in ventral view, in lateral view continuously narrowed toward subacute apex, inner side almost straight; penis narrowed in basal 3/4, apical 1/4 abruptly narrowed with subacuminate apex; in lateral view penis feebly curved, slender except widened base. Penis and parameres with sparse fine spines (Figs 18–20). Ovipositor with stylus (terminal segment) short, cylindrical; preterminal segment elongate ca. 7.5 times as long as styli, bearing several small spines; basal segment with baculus ca. 1.3 times as long as preterminal and distal segments combined (Fig. 21). Sexual dimorphism. Both sexes are very similar regarding external morphology. Males can be distinguished by patches of darkened setae on prosternum. Distribution. H. autanai n. sp. is known only from the two closely situated localities. Habitat. The specimens were collected by kick-sampling and individual collecting in preserved, permanently
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flowing submountain forest stream with riffles and pools. The stream at the sampling site was about 2–3 m wide, with stony substrate covered by moss mats (Figs 15, 16) and naturally humic water with pH 4.67. The measured conductivity was 4 μS, total dissolved solids (TDS) 2 ppm and water temperature 24.5 °C. Etymology. The species is named after Cerro Autana, tepui located in the Amazonas state which stands 1,220 m above sea level. The type locality was situated close to the Autana tepui.
FIGURE 22. Molecular distances measured as uncorrected p-distance using 816bp fragment of cox1 gene. In boxes are interspecific distances. Highlighted with colour are species concerned in this paper.
Discussion At present, the subfamily Larainae comprises 28 genera with more than 150 described species (Jäch & Kodada 2010). Almost half of the genera occur in the Neotropics, which indicates extraordinary richness of the Larainae fauna in this region. Interestingly, 11 neotropic genera include 53 species, whereas 51 species known from the Africa is grouped in only 6 genera. This however can be biased by the extent of knowledge on larains in both regions rather than representing real picture of the fauna of mentioned regions. High number of genera in the Neotropics likely reflects more diversified environmental conditions (table mountains, Andes, lowland forests etc.) and we could expect many new species to be described from this region in the future. Hexanchorus has very wide distribution (from Mexico to Argentina). Now it comprises 20 species, from which 7 occur in Venezuela. However, from the large, megadiverse countries like Brazil, only 2 species are known. Based on these facts, it is highly supposed that the genus is much more diverse and many new species are to be discovered and described. The cox1 sequence divergence was around 3% between H. angeli n. sp. and other Hexanchorus species from the Gran Sabana, which suggests close relationships of species from Gran Sabana, however it would be very useful to compare them with more remote species, i.e. from Andes or other river basins. The genus Hypsilara, even with smaller area of distribution, also very likely includes more species than we know today. It is known only from Venezuela, however the genus could be also found in neighboring countries like Colombia, Guyana or Brazil in the future. All known species are morphologically extremely similar, but they (at least some) had to be isolated for a long time. This is supported by molecular differences ranging from around 5% between H. autanai n. sp. and Hypsilara sp. (specimen from Auyán tepui), up to almost 12% between former two species and H. breweri (Fig. 22). Detected genetic distances among Hypsilara species fit well in the range of interspecific distances summarized recently (Hebert et al. 2003), however such a distinct differences in the sequence divergence between species within a small genus are not very common. These differences are likely due to hydrological isolation. The stream, where H. breweri was collected (see Čiampor Jr et al. 2013), belongs to the basin of the Río Cuyuni, which disembogues into Río Essequibo, flowing directly to the Atlantic ocean. The remaining two species, even geographically more distanced, occur in the Orinoco basin, and thus are hydrologically, and also phylogenetically more closely related. The differences in the genetic distance detected among Hypsilara species, which are morphologically almost identical is very interesting, and it would be, for better understanding of the evolution of local fauna, useful to analyse also other closely related species from this point of view.
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Acknowledgements We wish to thank Ján Kodada and Tomáš Derka for valuable comments on the manuscript. The anonymous reviewers are thanked for their comments. This study was supported by the Slovak Research and Development Agency, Project No. APVV-0213-10 and the project implementation: Development and application of the innovative diagnostic approach for the molecular identification of animals (ITMS: 26240220049) supported by the Research & Development Operational Programme funded by the ERDF.
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