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Neoelmis guarani Shepard & Barr, a sexually dimorphic new species from
Paraguay (Insecta: Coleoptera: Elmidae: Elminae)

Article  in  Zootaxa · February 2016

DOI: 10.11646/zootaxa.4083.3.6

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 Zootaxa 4083 (3): 418–430 ISSN 1175-5326 (print edition)
http://www.mapress.com/j/zt/
Copyright © 2016 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://doi.org/10.11646/zootaxa.4083.3.6
http://zoobank.org/urn:lsid:zoobank.org:pub:06CBA73B-1D2F-46B5-9A38-E37F91BE1C43

Neoelmis guarani Shepard & Barr, a sexually dimorphic new species from
Paraguay (Insecta: Coleoptera: Elmidae: Elminae)

WILLIAM D. SHEPARD1 & CHERYL B. BARR2

Essig Museum of Entomology, 1101 Valley Life Sciences Bldg. #4780, University of California, Berkeley, CA 94720 USA.
E-mail: 1william.shepard@csus.edu; 2cbarr@berkeley.edu

Abstract

Neoelmis guarani new species is described and illustrated from specimens collected in streams of the Cordillera de los
Altos, southeast of Asunción and near the towns of Piribebuy and Chololó, Paraguay. Males and females of this species
exhibit strong secondary sexual dimorphism not found in other known species of Neoelmis. Males have striking modifi-
cations of the pro- and mesothoracic legs and bear a pair of ventrally projecting processes on both the mesoventrite and
the second abdominal ventrite. Females have the elytra modified with a pair of dorsal projections.

Key words: riffle beetle, sexual dimorphism, South America

Resumen

Neoelmis guarani especies nuevas se describe e ilustra a partir de ejemplares recolectados en arroyos de la Cordillera de
los Altos, al sureste de Asunción y cerca de las ciudades de Piribebuy y Chololó, en Paraguay. A diferencia de otras espe-
cies conocidas de Neoelmis, los machos y hembras de esta especie presentan fuerte dimorfismo sexual secundario. Los
machos poseen notables modificaciones en el primer y segundo par de patas, así como un par de proyecciones que se ex-
tienden ventralmente en el mesoventrito y segundo ventrito abdominal. Por su parte, las hembras tienen los élitros modi-
ficados con un par de proyecciones dorsales.

Introduction

During a preliminary survey of the aquatic Byrrhoidea of Paraguay in 2006 (Shepard & Aguilar Julio 2010), a
unique new species of Neoelmis was discovered by WDS. Additional specimens were collected by the authors
during a repeat visit in 2011. The external morphology of the new species is very unusual compared to other known
species of Neoelmis, but it readily keys to Neoelmis in the available keys to Neotropical elmid genera (Manzo
2005; Brown in litt.).
Neoelmis is one of the larger elmid genera, with 49 described species in the New World (Manzo &
Archangelsky 2012). In addition, there are numerous undescribed species known from Central and South America.
Three species of Neoelmis are currently known to occur in Paraguay: N. maculata Hinton, N. nelo Hinton, and N.
opis Hinton (Shepard & Aguilar Julio 2010). The geographic distribution of Neoelmis extends from North America
(southwestern USA) to southern South America.

Material and methods

The description of N. guarani is based on seven specimens, three males and four females, which were collected by
the authors during trips to Paraguay in 2006 and 2011. We were guided in the field by author and scarab researcher
Carlos Aguilar Julio, and our collecting was enabled under his project “Inventario de Scarabaeidae, Elateridae,

418 Accepted by F. Ciampor: 19 Jan. 2016; published: 22 Feb. 2016

Buprestidae y Otros Coleoptera en Varios Lugares del Paraguay” (SEAM, Direccion de Vida Silvestre - DCPCB,
Dictamen AJ No. 888/2009).
Aquatic insect collecting techniques of either kick-sampling or manual substrate disturbance were used to
dislodge the beetles from the stream substrate, from which they were washed by the water current into rectangular
or D-frame aquatic nets. Specimens were then individually handpicked from the nets or were included in bulk
samples along with debris. The samples were preserved in 95% ethanol to be examined later in the laboratory.
Due to their small size, the specimens were not recognized to be an undescribed species until the samples were
sorted and examined in the laboratory. The following descriptions are based on the examination of seven intact
adult specimens using a Leica MZ 12.5 stereo microscope fitted with an ocular micrometer. Measurements of body
length represent the length of the pronotum plus the length of the elytra, excluding the head, which is retractile, and
the variable space between the pronotum and elytra. Measurements of body width are composed of both elytra at
their widest point. Specimen photographs were taken by CBB with a Visionary Digital BK Plus Lab System fitted
with a Canon EOS 7D camera, and by Robert Sites (University of Missouri) using a Leica MZ16 stereo microscope
coupled with the Leica Application Suite v4.4 Extended Depth of Focus module. Line drawings of the genitalia
were done by illustrator Kevin Wiseman (Alameda, California) using a Leica MZ 12.5 microscope fitted with a
camera lucida. The specimens are mounted on pinned card points. Dissected genitalia are placed in glycerol in a
genitalia vial pinned beneath the specimen.
Specimens will be deposited in the collections of the Essig Museum of Entomology at the University of
California, Berkeley, California, USA (EMEC) and in the Museo Nacional de Historia Natural del Paraguay, San
Lorenzo, Paraguay (INBP).

Description

Neoelmis guarani sp. n.

Figs. 1–12

Holotype, adult male. Body slender, 2.1 mm long (pronotum + elytra), 0.7 mm wide at widest point; body color
testaceous, appendages (antennae, palpi, legs) paler (Figs. 1–5).
Head with frons moderately setose, setae long and pale; genae covered with fine, adpressed, plastron setae;
eyes large, protuberant; antennal ridges produced, especially anteriorly. Antennae filiform, each with 11 segments;
antennomeres 1–2 twice as long as wide, antennomeres 3–10 more than twice as long as wide; antennomere 11
clavate, slightly curved, longer than antennomeres 9+10 (Figs. 1, 2). Clypeus setose, transverse, apex feebly
emarginate. Labrum transverse, apex arcuate; surface shiny with few setae; apicolateral margins with fringe of
pale, dense setae. Maxillary and labial palpi each with three palpomeres.
Pronotum (Fig. 1) elongate, 0.6 mm long, 0.5 mm wide at widest point about 1/3 distance from base; surface
shiny, smooth, and sparsely clothed with pale, recumbent setae except for central disc where finely granulate and
more densely setose; lateral margins bisinuate and irregularly crenulate; apicolateral angles acute, produced;
posterior border bisinuate, notched medially to receive scutellum; prominent, wide transverse sulcus at 1/3 distance
from apex, deeper fovea at midline with longitudinal groove which tapers posteriorly; two prominent, sinuate,
sublateral carinae; broad, shallow depression between each sublateral carina and lateral margin, with a large fovea
ventrally adjacent to transverse sulcus. Hypomeron with shallow depression on each side at basal 1/3 and a large
fovea ventrally adjacent to fovea between lateral margin and sublateral carina; without plastron setae.
Scutellum subovoid, longer than wide, feebly convex.
Elytra (Fig. 1) elongate, 1.5 mm long, 0.7 mm wide at widest point 1/3–1/2 distance from elytral apices;
shallow linear punctures and setae present. Each elytron with a short, prominent, basal carina on third interval
separating shallow lateral and medial depressions; two long sublateral carinae extending posteriorly from humeral
angle, inner one extending to apical 1/3, outer one extending nearly to apex. Epipleuron with plastron setae,
notched just before apex to receive tooth from abdominal ventrite 5.
Prosternum (Fig. 2) longer than wide, with flat, evenly and lightly setose, shiny disc bordered by two
longitudinal carinae 3/4 length of prosternum, fading out anteriorly; episternum clothed with plastron setae;
prosternal process wide, apex extending posterior to procoxae, margins raised and rimmed with narrow sulcus

A NEW SPECIES OF NEOELMIS FROM PARAGUAY Zootaxa 4083 (3) © 2016 Magnolia Press · 419
FIGURE 1. Neoelmis guarani, male dorsal habitus; length 2.1 mm.

420 · Zootaxa 4083 (3) © 2016 Magnolia Press SHEPARD & BARR
FIGURE 2. Neoelmis guarani, male ventral habitus; length 2.2 mm. Arrows indicate paired processes on the mesothorax and
abdominal ventrite 2.

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FIGURES 3–5. Neoelmis guarani, male. 3. Left prothoracic leg, femur, dorsal view. 4. Left prothoracic leg, posteroventral
view. 5. Left mesothoracic leg, anteroventral view.

422 · Zootaxa 4083 (3) © 2016 Magnolia Press SHEPARD & BARR
FIGURE 6. Neoelmis guarani, aedeagus, A—dorsal view, B—lateral view. Scale bar = 0.1 mm.

between. Mesoventrite (Fig. 2) shorter than wide; with a pair of large, posterolateral, ventrally projecting processes
(Fig. 2, arrow) adjacent to and extending below mesocoxae; processes with ventromedially oriented, flattened
surfaces; area between processes excavated; mesanepisternum with plastron setae. Metaventrite (Fig. 2) shorter
than prosternum, longer than mesosternum; disc flat, shiny, sparsely setose, with distinct discrimen; with two
sublateral carinae extending from mesocoxae to metacoxae and small, carinate tubercle near each posterolateral
margin; plastron setae present laterad to carinae and on metanepisternum. Pro- and mesocoxae globular; metacoxae
transverse. Pro- and mesotrochanters of similar size, metatrochanters larger.
Prothoracic leg (Figs. 1–4) with profemur abruptly and deeply excavated on both anterior and posterior
surfaces 1/3 distance from apex (Fig. 3), excavations lined with short, stiff setae (Fig. 4); anterior surface bearing a
large, blunt, distally directed tooth at inner edge of excavation; plastron setae present except in excavations.
Protibia (Fig. 4) slender, wider apically; posterior face with long setae present basally and apically and a deep
excavation near apex; cleaning fringe of long, dense, pale setae on anterior surface. Protarsus (Fig. 4) shiny,
tarsomeres 1–4 with long setae ventrally, tarsomere 5 with a pair of long, projecting setae at apex; tarsomere 5
longer than tarsomeres 1–4 combined; claws simple. Mesothoracic leg (Figs. 1, 2, 5) with mesofemur (Fig. 5)
widened medially; a deep, longitudinal groove present on ventral surface to receive tibia when folded, posterior
margin of groove with numerous, long, curved setae; plastron setae present except inside of groove. Mesotibia
(Fig. 5) in lateral view with ventrally directed expansion at middle and shallow excavation on anterior face about 1/
3 distance from apex; with anterior and posterior cleaning fringes (Fig. 1) having long, tuft-like setae that extend

A NEW SPECIES OF NEOELMIS FROM PARAGUAY Zootaxa 4083 (3) © 2016 Magnolia Press · 423
FIGURE 7. Neoelmis guarani, female dorsal habitus; length 2.3 mm. Arrow indicates paired elytral protuberances.

424 · Zootaxa 4083 (3) © 2016 Magnolia Press SHEPARD & BARR
FIGURE 8. Neoelmis guarani, female lateral habitus; length 2.3 mm.

beyond tibial apex. Mesotarsus similar to protarsus; right mesotarsus missing tarsomeres 3–5 due to breakage.
Metathoracic leg (Figs. 1, 2) with metafemur unmodified. Metatibia with cleaning fringe on posterior surface at
central 1/3; inner surface flat at apical 1/4, glabrous and shiny with a small peg near ventral margin. Metatarsus
similar to pro- and mesotarsus.
Abdomen (Fig. 2) with five ventrites; ventrites 1–4 decreasing in length posteriorly, ventrite 5 longer than all
but ventrite 1. Ventrite 1 longest, with basomedial depression bordered by two short carinae which extend from
posterior margin of metacoxae nearly to posterior margin of ventrite; surface shiny and sparsely setose between
carinae, plastron setae present laterad to carinae. Ventrite 2 with a pair of processes (Fig. 2, arrow) projecting
ventrally from anteromedial border, size of each process less than half that of each mesoventral process; surface
between processes shiny, plastron setae present elsewhere. Ventrites 3–5 covered with plastron setae; ventrite 5
with a dorsally projecting tooth on each lateral margin which serves to link with a notch in the epipleuron. Ventrite
5 removed from abdomen and mounted on a card point.
Aedeagus (Fig. 6) in dorsal view with phallobase twice as long as parameres, widely open in basal half;
parameres widely open at apical 2/3 exposing penis; penis stout, sides faintly arcuate at basal 1/2, then moderately
constricted and nearly parallel-sided to apical 1/8, thereafter evenly converging to narrowly rounded apex which
extends slightly past paramere apices. Penis in ventral view with fibula at middle. Aedeagus in lateral view with
apices of parameres and penis acutely pointed. Aedeagus removed from abdomen and placed in genitalia vial.
Allotype, adult female. Body slender, 2.3 mm long (pronotum + elytra), 0.8 mm wide at widest point about 1/
3 distance from elytral apices (Fig. 7). Slightly larger but generally similar to male except as follows: Each elytron
(Figs. 7, 8) with a dorsally projecting protuberance in the third interval at about 1/3 distance from apex (Fig. 7,
arrow), pointed and subtriangular in lateral view (Fig. 8). Mesoventrite and abdominal ventrite 2 lacking
modifications. All legs (Figs. 7, 8) without special modifications, similar to each other; tibiae with cleaning fringes.
Ovipositor very elongate; valvifers more than 10 times longer than wide, thin basally then widest apically by
coxites; coxites twice as long as wide, rectangular; styli 2-segmented, second segment very short.

A NEW SPECIES OF NEOELMIS FROM PARAGUAY Zootaxa 4083 (3) © 2016 Magnolia Press · 425
FIGURE 9. Arroyo Naranjo, type locality of Neoelmis guarani.

426 · Zootaxa 4083 (3) © 2016 Magnolia Press SHEPARD & BARR
FIGURE 10. Arroyo Naranjo, type locality of Neoelmis guarani.

Variation. Although seven specimens (three males and four females) is a small sample with which to examine
variation, some differences were noted. Most striking of these are the different external modifications of males and
females, detailed in the descriptions and Diagnosis. In addition, the females (2.2–2.3 mm long, 0.7–0.8 mm wide)
are slightly larger than the males (2.1–2.2 mm long, 0.7 mm wide). In some specimens mineral deposits on the
dorsum obscure details of the surface, causing semi-glabrous, shiny areas to appear granulate. Specimens also
exhibit varying degrees of surface abrasion which can lessen the number of setae present in the abraded area
compared to that of an unabraded specimen.
Diagnosis. The distinctive secondary sexual characters of N. guarani, present in both males and females, serve
to distinguish this species from all other known species of Neoelmis. Males (Figs. 1–5) have strong modifications
of the profemora, protibiae, mesofemora, mesotibiae, and metatibiae, and bear a pair of ventral processes on both
the mesoventrite and second abdominal ventrite. Females (Figs. 7, 8) have a dorsal pair of elytral protuberances
and unmodified legs.
The aedeagus of N. guarani most closely resembles that of N. simoni (Grouvelle) which is known only from
Venezuela (Hinton 1939). However, in N. guarani the phallobase is open only in the basal half and the penis barely
projects beyond the tips of the parameres, whereas in N. simoni the phallobase is open entirely and the penis
projects well beyond the tips of the parameres. The aedeagus of N. guarani bears no resemblance to those of N.
maculata, N. nelo, nor N. opis, other species occurring in Paraguay (Hinton 1940a, 1972).
Type material. Holotype (male): PARAGUAY: Paraguarí [Dpto.], Arroyo Naranjo at Balneario Salto Cristal
7.5 km S Piribebuy, 17 Feb. 2011, C. B. Barr // 25º32.026’ S 57º01.717’ W; elevation 214 m // HOLOTYPE
Neoelmis guarani Shepard & Barr [red label, handwritten]. Deposited in EMEC. Allotype (female): PARAGUAY:
Cordillera [Dpto.], Piribebuy-Barrio Santa Ana, unnamed stream, 820 ft [250 m], 25º27.95’S 57º01.99’W, 18 VI
2006, [WDS-A-1687, on underside of label] // W. D. Shepard, leg. // ALLOTYPE Neoelmis guarani Shepard &
Barr

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FIGURE 11. Arroyo Mborebí, Neoelmis guarani collection site.

428 · Zootaxa 4083 (3) © 2016 Magnolia Press SHEPARD & BARR
FIGURE 12. Capilla Cue, Neoelmis guarani collection site.

[red label, handwritten]. Deposited in EMEC. Paratypes (2MM, 3FF): PARAGUAY: Cordillera [Dpto.],
Piribebuy-Barrio Santa Ana, unnamed stream, 820 ft [250 m], 25º27.95’S 57º01.99’W, 18 VI 2006 [WDS-A-1687,
on underside of label] // W. D. Shepard, leg. // PARATYPE Neoelmis guarani Shepard & Barr [yellow label] (1M,
EMEC); Cordillera [Dpto.], Piribebuy–B. Sta. Ana, 17 II 2011, 230 m, Arroyo Mborebi, S25º28.075’ W57º02.330’
// W. D. Shepard, leg. [WDS-A- 1830, on underside of label] // PARATYPE Neoelmis guarani Shepard & Barr
[yellow label] (1M, INBP); Paraguarí [Dpto.], Arroyo Naranjo at Balneario Salto Cristal 7.5 km S Piribebuy, 17
Feb. 2011, C.B. Barr // 25º32.026’ S 57º01.717’ W, elevation 214 m // PARATYPE Neoelmis guarani Shepard &
Barr [yellow label] (2 FF; EMEC, INBP); Paraguari [Dpto.], 15.3 km NE Paraguari, 18 II 2011, 282 m, Capilla
Cue, S25º33.210’ W57º02.829’ // W. D. Shepard, leg. [WDS-A-1834, on underside of label] // PARATYPE
Neoelmis guarani Shepard & Barr [yellow label] (1F, EMEC).
Etymology. Guarani, a noun in apposition, was chosen to honor the indigenous Guaraní people who are native
to the region and whose language is widely spoken in Paraguay.
Distribution. Neoelmis guarani is known from four localities in the highlands area of the Cordillera de los
Altos southeast of Asunción in the departments of Cordillera and Paraguarí. Although 42 streams have been
sampled during our survey of the aquatic Byrrhoidea of Paraguay, the species has been found only in this small
area. The elevations at the collection sites range from 214–282 masl and they are all less than 10 km apart (straight
line distance) near the towns of Piribebuy and Chololó.
Habitat. The type locality, Arroyo Naranjo (Figs. 9, 10), is a small to medium-sized stream with slightly turbid
water and a substrate of orange sand with sparse gravel/cobbles, numerous bedrock outcrops and ledges, and small
waterfalls. The three other streams are similar in size and substrate, except that Arroyo Mborebí (Fig. 11) lacks
extensive bedrock outcrops, at least at the collection site (Figs. 11, 12). Other elmids collected in association with
N. guarani include Heterelmis sp., Hexacylloepus sp., Hexanchorus sp., Macrelmis sp., Microcylloepus longipes

A NEW SPECIES OF NEOELMIS FROM PARAGUAY Zootaxa 4083 (3) © 2016 Magnolia Press · 429
 (Grouvelle), M. inaequalis (Sharp), Neoelmis nelo Hinton, Phanocerus sp., Stenhelmoides sp., and Xenelmis
micros (Grouvelle).

Discussion

Secondary sexual dimorphism of the type seen in N. guarani has been unknown in the genus until now. Hinton
(1940a, 1940b) listed several less-striking characters that he observed in nine species from Mexico and South
America, but none of these involved the male mesoventrite or leg modifications other than spination.
Because we did not observe male and female N. guarani in copula, we can only comment on aspects of the
secondary sexual dimorphism and speculate on possible function. The inside distance between the elytral
protuberances of the female (Fig. 7) is wider than the outside distance between the processes on male abdominal
ventrite 2 (Fig. 2). Her elytral protuberances are positioned directly above abdominal ventrite 2 (Fig. 8), so that if
the male were mounted directly above her, the processes on his abdominal ventrite 2 would fit between them. The
mesoventral processes of the male would be then positioned at the pronotal/elytral junction on the female. The
modifications of the male legs (Figs. 1–5) could serve to link with those of the female to further secure his position.

Acknowledgments

We greatly thank Carlos Aguilar Julio, who first interested us in the aquatic Byrrhoidea of Paraguay and who has
never failed to be a gracious host and accomplished guide in the field. In addition, we are indebted to Dr. John
Kochalka, Head of the Departamento de Invertebrados, Microbiología y Paleontología of the Museo Nacional de
Historia Natural del Paraguay, for kindly providing a letter of collaboration and assistance with the collecting
permit on which WDS is listed. We appreciate the generosity and expertise of Dr. Robert Sites whose specimen
close-up images enhance this paper, and of Daniel Reynoso-Velasco who greatly improved the Spanish translation
of our abstract, both from the Enns Entomology Museum, University of Missouri-Columbia. Kevin Wiseman of
Alameda, California, is thanked for his skillful illustration of the male genitalia.

References

Hinton, H.E. (1939) On some new and little known South American Neoelmis Musgrave (Coleoptera, Elmidae). The
Entomologist’s Monthly Magazine, 75, 228–234.
Hinton, H.E. (1940a) A synopsis of the Brazilian species of Neoelmis Musgrave (Coleoptera, Elmidae). The Annals and
Magazine of Natural History, Series 11, 5 (26), 129–153.
Hinton, H.E. (1940b) A monographic revision of the Mexican water beetles of the family Elmidae. Novitates Zoologicae, 42
(2), 217–396.
Hinton, H.E. (1972) New species of Neoelmis from South America (Coleoptera, Elmidae). Papéis Avulsos de Zoologia, 26 (9),
117–135, pls. I–II.
Manzo, V. (2005) Key to the South American genera of Elmidae (Insecta: Coleoptera) with distributional data. Studies on
Neotropical Fauna and Environment, 40 (3), 201–208.
http://dx.doi.org/10.1080/01650520500140619
Manzo, V. & Archangelsky, M. (2012) Two new species of Elmidae (Coleoptera) from Argentina. Zootaxa, 3478, 267–281.
Shepard, W.D. & Aguilar Julio, C.A. (2010) Estudio preliminary de las familias de escarabajos acuáticos Dryopidae, Elmidae,
Lutrochidae y Psephenidae conocidos de Paraguay (Coleoptera: Byrrhoidea). Boletin de la Museo Nacional de Historia
Natural del Paraguay, 16 (1), 30–42.

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