Research Article |
Corresponding author: Cheryl B. Barr ( cbarr@berkeley.edu ) Academic editor: Mariano Michat
© 2018 Cheryl B. Barr.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Barr CB (2018) Amazonopsis, an unusual new genus of riffle beetle from South America with two new species (Coleoptera, Elmidae, Elminae). ZooKeys 803: 71-92. https://doi.org/10.3897/zookeys.803.28124
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Amazonopsis gen. n. is described to include A. theranyi sp. n. from Peru, Venezuela and French Guiana, and A. camachoi sp. n. from Venezuela. The descriptions are accompanied by figures illustrating the male and female habitus of A. theranyi, the male habitus of A. camachoi, and male genitalia of both species. Amazonopsis theranyi exhibits pronounced secondary sexual dimorphism which likewise may be a characteristic of the genus. Amazonopsis males have modified protarsal and mesotarsal claws, a pair of small spines on the anterior prosternum, and a pair of ventrally directed processes on the posterior metaventrite. Females of A. theranyi display a pair of unique, oval perforations in the cuticle of the pronotum and have unmodified claws; females of A. camachoi are unknown. Descriptions are furnished of the stream habitats and microhabitats where the study specimens were collected.
Aquatic beetles, French Guiana, Neotropics, peri-Amazonian, Peru, sexual dimorphism, taxonomy, Venezuela
Presently 40 genera and over 250 species of the aquatic beetle family Elmidae are known from South America (
Specimens of an unusual new genus and species of elmid were collected in the Amazon Basin of southeastern Peru in 2012 during a water quality survey of stream macroinvertebrates conducted by the Stroud Water Research Center (Avondale, Pennsylvania, USA) in conjunction with the Amazon Center for Environmental Education and Research (ACEER) (West Chester, Pennsylvania, USA) (
The purpose of this paper is to describe this distinctive new genus of elmid, its two new species, and the stream habitats from which they were collected.
Four specimens collected by the Stroud–ACEER project in Peru were taken from artificial leaf packs, consisting of plastic mesh bags filled with 7 g of fresh Inga edulis Martius (Fabaceae) leaves, which were deployed in the stream for about four weeks (
Water quality measurements of the streams were provided by the Stroud Water Research Center for the specimens collected in Peru during the ACEER project (
Specimens were examined in the lab using a Leica MZ 12.5 stereo microscope fitted with an ocular micrometer. Measurements of total body length represent the length of the pronotum plus the length of the elytra, excluding the head and the variable space between the pronotum and elytra; measurements of body width are composed of both elytra at their widest point. The habitus images were taken with a Visionary Digital BK Plus Lab System fitted with a Canon EOS 7D camera. A Syncroscopy AutoMontage system was used for the genitalia images. All of the specimens are double-mounted on card points and pins. Males have the aedeagi removed and stored with glycerin in genitalia vials beneath the donor specimens.
Label data are reported verbatim as found on the specimen labels: “ / ” indicates separate lines on one label and “ // ” indicates separate labels. Brackets “ [] ” indicate additional clarifying information not included on the label.
The distribution map is an alteration of a map of South America provided free on the internet by
Specimens will be deposited in the following institutions:
Amazonopsis theranyi sp. n.
Amazonopsis camachoi sp. n.
The flattened and bent pro- and mesotarsal claws of the males (Figs
Male. Body stout, elongate-oval, at least 2× as long as wide; convex dorsally. Surface of dorsum and parts of venter covered by thin, pale gray, microreticulate plastron; thick, glossy plastron present laterally on sterna and adjacent surfaces of coxae, legs (except tarsi), lower margin of hypomeron, abdominal ventrites (except along midline), and entire epipleuron; head (vertex, frons, clypeus) and pronotum with short, broad, flat, pale yellow setae. Tibial cleaning fringes well-developed, formula 2-2-1. Head. Antenna filiform, with 11 antennomeres. Vertex with V-shaped carina opening anteriorly; frons slightly elevated between eyes; eyes large, subcircular in outline. Clypeus rectangular, wider than long. Labrum rectangular, not as wide as clypeus. Mandible with three short, rounded, apical teeth. Maxillary palpus with four palpomeres. Labial palpus with three palpomeres. Pronotum. Subquadrate, slightly wider than long, widest at midlength; without carinae or gibbosities. Disc, including punctures, covered with pale microreticulate plastron. Scutellum subcircular to ovate, flat. Elytron. Elongate, about 3× as long as wide; without carinae except for swollen, raised base of third interval; humeral angles protuberant. Disc with 10 longitudinal rows of coarse, deep punctures; row 10 near margin with much smaller punctures than rows 1–9. Epipleuron with excavation adjacent to marginal lobe of abdominal ventrite 4. Surface of disc, including punctures, with thin, pale microreticulate plastron, often abraded; cuticle beneath very shiny, reddish-brown. Leg. Femur and tibia covered with thin, shiny layer of dense plastron; tarsus without plastron. Prothoracic leg shortest, metathoracic leg longest. Pro- and mesocoxa globose, metacoxa transverse. Pro- and mesotibia each with anterior and posterior cleaning fringes of long setae; metatibia with a single, posterior fringe. Claws long, without basal teeth; protarsal and mesotarsal inner and outer claws dissimilar; protarsal inner and mesotarsal outer claws enlarged, laterally flattened, bent at base; protarsal outer and mesotarsal inner claws smaller, narrower; metatarsal claws shorter, flattened, but basically unmodified. Venter. Pale microreticulate plastron present on ventral surfaces except at midline; plastron yellow and most evident near lateral thoracic margins and on abdominal ventrites. Prosternum slightly shorter than metaventrite; anterior margin curved posteriad, bounded by pair of small, ventrally directed spines; prosternal process about 2× as long as wide, margins raised, bluntly rounded at apex. Mesoventrite shortest; deep cavity present to accommodate prosternal process. Metaventrite longest, slightly longer than prosternum; metathoracic discrimen distinct; posteromedial margin with pair of ventrally directed processes. Abdomen with five ventrites; ventrites 1–4 decreasing in length posteriorly, ventrite 5 longer than ventrite 1; ventrites narrower medially and wider laterally; ventrite 4 lateral margin with lobe to link with groove on epipleuron, posterior margin strongly raised and rounded. Genitalia. Trilobate, typical form.
Female. Although the female of A. camachoi is unknown, it is possible, if not likely, that secondary sexual dimorphism is a generic characteristic. Females of A. theranyi exhibit the following differences (Fig.
“Amazon”, a Greek word for a legendary race of warrior women, refers to the robust, unique features of the beetles as well as the provenance of the genus; plus “-opsis” from the Greek meaning “look, appearance, likeness.” Gender, feminine.
In
Holotype male deposited in
french guiana / ca. 4 km ESE Saül / Cr.[ique] Nouvelle France / 03.6063, −53.1762 / 9-XI-2016, C. B. Barr // Parc Amazonien / de Guyane just / below Point Chaud / coll. from leaf pack (1 F,
Amazonopsis theranyi males (Figs
Holotype male. Length, 4.20 mm; width, 1.90 mm. Cuticle mostly covered with pale, thin microreticulate layer dorsally and ventrally, with thicker, glossy, golden-yellow plastron ventrolaterally on thoracic sterna and abdominal ventrites; cuticle shiny, dark reddish-brown where exposed. Antenna. Yellow-brown. Antennomeres 1–10 clavate, antennomere 11 fusiform; antennomeres 1 and 2 each stouter than 3–10 which are of similar size and shape; antennomeres 3–10 each with dense tuft of setae at apicoventral margin, overlapping base of next; antennomere 11 with an elongated patch of short setae near the ventral apex. Head. Vertex, frons and clypeus covered with pale, microreticulate plastron and broad, flat, yellow setae. Clypeus dark brown, barely emarginate at center of apical margin, setae slightly less dense than on vertex and frons. Labrum dark brown, barely emarginate, apicolateral angles broadly rounded; surface with small, evenly spaced punctures and short, fine setae; apical and lateral margins with fringe of pale, dense setae, longest laterally. Mandible with three short, rounded, apical teeth. Maxillary palpus yellow-brown; palpomere 4 slightly flattened and curved, longer than 1–3 combined, with oval patch of sensillae at apex. Labial palpus with palpomeres 1 and 2 short, dark brown; palpomere 3 longer than 1 and 2 combined, yellow-brown, ovoid and moderately flattened. Pronotum. Length, 1.20 mm; width, 1.50 mm. In dorsal view, lateral margins coarsely granulate, unevenly arcuate; anterior margin trisinuate, strongly arcuate at middle; anterolateral angles acute, depressed. In lateral view, moderately convex. Disc covered with pale microreticulate plastron and deep, closely spaced, coarse punctures; punctures larger towards the lateral margins, smaller towards the midline; punctures generally spaced a diameter apart; punctures lined with plastron and associated with very short, erect setae; anteromedial disc and lateral areas with broad, flat, recumbent, yellow setae. Center of midline with narrow, lightly impressed, bare, longitudinal line; length about ½ that of pronotum. Scutellum subcircular. Elytron. Length, 3.00 mm; width, 0.95 mm. Surface covered with pale, thin microreticulate plastron, abraded at center of disc; punctures striate, deep, coarse, lined with plastron; intervals of striae with fine, sparse setae. Humeral angle with low, knob-like protuberance; base of third interval slightly swollen and raised; lateral margins smooth, recurved with narrow, longitudinal band of hypomeron plastron visible; shallow sulcus about one interval wide adjacent to lateral margin, extending from humeral angle to apical 1/5; elytra constricted at apical 1/5 at point of linkage with abdominal ventrite 4 lateral lobe; apex evenly rounded, moderately produced. Legs. Femora and tibiae covered by thin layer of shiny, pale yellow plastron, sparsely setose and shallowly punctate; tarsus red-brown, without plastron. Procoxa posterior surface coarsely punctate; dense patch of long, golden-yellow setae present on lateral face. Prothoracic leg with tibia longer than femur, tarsus shorter; profemur with oval patch of long, recumbent, golden-yellow setae on anterior surface near base; protibia with pair of cleaning fringes nearly ½ as long as tibia, posterior fringe slightly shorter; protarsus with tarsomeres 1–4 bearing dense tufts of moderately long setae in two rows at apicoventral margins; tarsomere 5 longer than the others combined, with moderately long setae on ventral surface and a few longer, golden-yellow setae at apex which barely extend over base of claws. Protarsal claws dissimilarly shaped, long, laterally flattened, sharply acute; inner claw enlarged, base bent outward, tip bent ventrally; outer claw shorter, base and tip not bent. Mesocoxa coarsely punctate and granulate; dense patch of long, golden-yellow setae present on lateral face and adjacent sternum. Mesothoracic leg similar to prothoracic leg except mesofemur with elongate patch of long, recumbent golden-yellow setae on posterior surface extending from near base to half femoral length; mesotibia with pair of cleaning fringes nearly ⅔ as long as tibia. Mesotarsal claws dissimilarly shaped, much longer than protarsal claws, laterally flattened, sharply acute; outer claw enlarged, slightly curved, bent about 90° at base then flattened and widened, more than 2× wider than inner claw; inner claw slightly shorter and much narrower. Metacoxa medial surface with longitudinal, sulcate row of coarse, deep punctures; posterolateral surface with dense patch of long, golden-yellow setae. Metathoracic leg similar to other legs except tibia much longer than femur; single cleaning fringe on posterior face about ⅔ as long as tibia; both claws slightly flattened but basically unmodified, stout, shorter than pro- and mesotarsal claws. Venter. Hypomeron with large, coarse, closely spaced punctures, more than 2× diameter of lateral pronotal punctures; ventral margin with broadly rounded lobe directed toward coxa; longitudinal band of golden-yellow plastron present on central ventral margin. Prosternum anterior margin raised, bearing two small, ventrally directed spines; distance between spines narrower than labrum; anterolateral margin behind each eye having a small, nearly semicircular notch; prosternal process about 2× as long as wide, with elevated margin; prosternal disc covered with pale, microreticulate plastron, scattered broad, flat, yellow setae, and large circular punctures spaced slightly less than a puncture diameter apart; golden-yellow plastron present laterally. Mesoventrite depressed between mesocoxae; punctation similar to that of prosternum; disc with pale, microreticulate plastron, mesepimeron with band of dense, golden-yellow plastron. Metaventrite depressed between mesocoxae; discrimen sulcate; posteromedial margin with two low, obtuse, ventrally directed processes; punctures more oval than circular in shape, closer together near midline; disc with pale yellow plastron except along midline, most dense laterad and on metepisternum. Abdomen with pale yellow plastron on all surfaces except for areas of bare, shiny cuticle at midline; ventrites 1–4 non-setose, ventrite 5 with fine, scattered setae; punctures not as large as those on thoracic sternites, becoming progressively smaller with each succeeding ventrite; punctures evenly spaced but less dense than on thoracic sternites; ventrite 1 anterior margin between metacoxae smoothly arcuate; ventrites 1–4 moderately convex at lateral ¼; ventrite 5 with two basolateral swellings each bordered by a shallow depression, apical ⅓ depressed and margin broadly rounded. Genitalia (Fig.
Allotype female. Length (excluding head), 4.25 mm; width, 2.00 mm. Pronotum 1.25 mm long, 1.55 mm wide; elytron 3.15 mm long, 1.00 mm wide. Secondarily sexually dimorphic as follows: pronotum with two, moderately large, oval perforations of the cuticle on either side of midline; all claws normal, not modified, shorter than those of males; anterior margin of prosternum without paired spines; posterior margin of metaventrite without paired, ventrally directed processes. Otherwise, similar to the male.
The most striking variation is the strong secondary sexual dimorphism exhibited by males and females. Males (Figs
The single male specimen from Venezuela (Figs
Named for Therany Gonzales Ojeda of ACEER, Puerto Maldonado, Madre de Dios, Peru, the collector of the type series.
This species is currently known only from widely separated, single localities in southeastern Peru, southwestern Venezuela, and French Guiana (Fig.
Quebrada Santo Rosario (Fig.
Crique Nouvelle France (Fig.
The information for the Cerro de la Neblina site, a tributary Río Baria, was provided by collector Warren Steiner (in litt.) from his field notes: “small whitewater stream [where I] spent an hour collecting … in leafy side pools near flowing part of stream … ”. The label data with the specimens states “small pool full of dead leaves” as well as an elevation of 140 m. In addition,
Aquatic byrrhoid beetles collected at the same localities as Amazonopsis theranyi include: PERU: Gyrelmis brunnea Hinton, 1940, G. longipes Hinton, 1940, G. maculata Hinton, 1940, Hintonelmis Spangler, 1966 , Neoelmis Musgrave, 1935, Pilielmis Hinton, 1971, Portelmis (Elmidae); Psephenops Grouvelle, 1898 (Psephenidae). FRENCH GUIANA: Cylloepus Erichson, 1847, Gyrelmis brunnea, G. nubila Hinton, 1940, G. spinata Hinton, 1940, G. thoracica Hinton, 1940, Heterelmis Sharp, 1882, Hexacylloepus Hinton, 1940, Hintonelmis perfecta (Grouvelle, 1908), Macrelmis tereus (Hinton, 1946), Neoelmis, Pilielmis apama Hinton, 1971, Phanocerus Sharp, 1882 (Elmidae); undescribed genus/species (Protelmidae); Dryops Oliver, 1791, Elmoparnus collinsae Spangler & Steiner, 1983, Platyparnus bollowi (Hinton, 1939), P. frater (Hinton, 1939) (Dryopidae); Lutrochus Erichson, 1847 (Lutrochidae). VENEZUELA: Gyrelmis Hinton, 1940, Hexacylloepus, Neoelmis, Pilielmis, Stegoelmis fera Spangler, 1990, S. geayi (Grouvelle, 1908), S. tuberosa Spangler, 1990 (Elmidae); new genus (Protelmidae); Dryops, Pelonomus Erichson, 1847 (Dryopidae). Note: Venezuelan records are cited from
Holotype male deposited in
Amazonopsis camachoi males (Figs
Holotype male (Figs
Named for Jesús Camacho of La Universidad del Zulia, Maracaibo, Venezuela, who collected the unique type specimen.
This species is currently known only from one locality in southeastern Venezuela (Fig.
The collector of the specimen recalled that the stream was small, shallow, sandy, and shaded, and that its waters were dark, tannin-stained, and contained decaying leaves (Camacho, in litt.).
The only other specimen known from this locality, an unidentified species of Heterelmis (Elmidae), is in the
As pointed out in the generic diagnosis, Amazonopsis is unique in that no other known genera have males with such bizarrely modified claws or females with pronotal perforations. Amazonopsis bears similarities to both Stenhelmoides and Pagelmis, particularly regarding its extensive dorsal plastron and lack of pronotal and elytral carinae, however, the genus is distinguished by the presence of a lateral projection of the fourth abdominal ventrite and lack of distinctive plastron pattern. Amazonopsis also shares some morphological characteristics with Portelmis. At this time, lacking phylogenetic analysis, it is not possible to ascertain to which other genus or genera Amazonopsis is most closely related.
Given the large geographic range indicated by the four known occurrences (Fig.
Secondary sexual dimorphism, which occurs in several elmid genera and species, involves modifications of various surfaces and structures, particularly on the legs and venter (
The A. theranyi specimens examined for this paper are from closed-canopy streams in humid, tropical lowland forests and share habitat similarities (Figs
At first glance it may seem that the genus Amazonopsis displays an unusual geographic distribution pattern (Fig.
It was fortuitous that through Wills Flowers (Florida A&M University, Tallahassee) I heard about the Stroud–ACEER project in Peru and their need for an elmid specialist to do identifications. I owe special thanks to Bernard Sweeney and David Funk at Stroud Water Research Center (Avondale, Pennsylvania), who provided the specimens forming the basis for this study, supplementary information, and the type locality photograph (Fig.
Collecting and export permits in French Guiana were provided by the Collectivité Territoriale de Guyane according to the DIAG project (Convention N°APA-973-13). Hydreco Guyane, the Office de l’Eau de Guyane, the Direction de l’Environnement de l’Aménagement et du Logement de Guyane, and the Parc Amazonien de Guyane are thanked for providing financial and technical support. I am especially grateful to Simon Clavier, formerly of Hydreco (Laboratoire Environnement de Petit Saut, Kourou), who organized and supplied our expeditions, guided us expertly in the field, and attempted to collect additional material for this article.
Charyn Michelli (
Lastly, I appreciate Bill Shepard, my partner in the study of riffle beetles and in life, for his encouragement and support.