Research Article |
Corresponding author: Liza M. González-Rodríguez ( lizmgr287@gmail.com ) Academic editor: Mariano Michat
© 2022 Liza M. González-Rodríguez, Andrew Edward Z. Short.
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:
González-Rodríguez LM, Short AEZ (2022) Revision of the water scavenger beetle genus Notionotus Spangler, 1972 in the Neotropical Region (Coleoptera, Hydrophilidae, Enochrinae). ZooKeys 1109: 141-191. https://doi.org/10.3897/zookeys.1109.80775
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The Neotropical species of the water scavenger beetle genus Notionotus Spangler, 1972 are revised using an integrative taxonomic approach combining morphology with DNA sequence data from two genes. Support exists for four putative species groups into which 18 species are placed, including twelve that are described here as new: N. bicolor sp. nov. (Suriname), N. bifidus sp. nov. (Venezuela), N. brunbadius sp. nov. (Brazil), N. garciae sp. nov. (Brazil), N. giraldoi sp. nov. (Brazil), N. insignitus sp. nov. (Venezuela), N. juma sp. nov. (Brazil), N. parvus sp. nov. (Suriname), N. patamona sp. nov. (Guyana), N. peruensis sp. nov. (Peru), N. retusus sp. nov. (Guyana), and N. vatius sp. nov. (Brazil). Four new synonymies are created: N. shorti Queney syn. nov. is found to be conspecific with N. dilucidus Queney; N. edibethae García syn. nov., N. nucleus Perkins syn. nov., and N. perijanus García syn. nov. are found to be conspecific with N. tricarinatus Perkins. New records are provided for all previously described species except N. mexicanus Perkins. Within the Neotropical region, the range of the genus is greatly expanded and now known from as far south as Bolivia and the Brazilian state of Mato Grosso do Sul. While a few species are found in hygropetric habitats, most are associated with the margins of forested streams. Genitalia and habitus images are provided for nearly all species, as well as a key to the four species groups.
Las especies de escarabajos acuáticos detritívoros neotropicales del género Notionotus Spangler, 1972 son revisadas usando taxonomía integrativa, combinando morfología con datos de secuencias de ADN para dos genes. Se encontró soporte para cuatro grupos de especies conformados por 18 especies, incluyendo 12 que son aquí descritas como nuevas: N. bicolor sp. nov. (Suriname), N. bifidus sp. nov. (Venezuela), N. brunbadius sp. nov. (Brazil), N. garciae sp. nov. (Brazil), N. giraldoi sp. nov. (Brazil), N. insignitus sp. nov. (Venezuela), N. juma sp. nov. (Brazil), N. parvus sp. nov. (Suriname), N. patamona sp. nov. (Guyana), N. peruensis sp. nov. (Peru), N. retusus sp. nov. (Guyana), N. vatius sp. nov. (Brazil). Se sinonimizan cuatro especies: N. shorti Queney syn. nov. se considera conespecífico con N. dilucidus Queney; N. edibethae García syn. nov., N. nucleus Perkins syn. nov., y N. perijanus García syn. nov. son conespecíficos con N. tricarinatus Perkins. Nuevos registros son provistos para todas las especies previamente descritas excepto para N. mexicanus Perkins. Hasta el momento, en la región neotropical, el rango de distribución del género es ampliamente expandido, se conoce desde el Sur de Bolivia hasta el estado brasileño de Mato Grosso do Sul. Si bien algunas especies son encontradas en hábitats higropétricos, la mayoría de ellas están asociadas a las orillas de arroyos boscosos. Se proveen imágenes del hábito dorsal y genitalia para la mayoría de las especies, al igual que la clave para los cuatro grupos de especies.
aquatic insects, integrative taxonomy, new species, South America
The genus Notionotus Spangler, 1972 is a group of small to minute water scavenger beetles that occur in streams and seepages in the tropics of the New World and Southeast Asia. The original concept of Notionotus was based on two new species from the Venezuelan Andes (
During the last two decades, extensive fieldwork for aquatic beetles in the Neotropics has resulted in the collection of many new specimens of Notionotus, including some that represent new species. We here use an integrative approach, combining adult morphological data with DNA sequences from two gene loci, to revise the Neotropical species of the genus. This revision provides a taxonomic foundation not only for future descriptive work on the genus, but also for evolutionary studies on the diversity and biogeography of the Neotropical region.
We amplified and sequenced the mitochondrial gene COI and the nuclear ribosomal gene 28S for 56 specimens of Notionotus, including 55 specimens from the Neotropical region and one unidentified specimen from India which we used as an outgroup to root the tree. We sampled specimens from most localities for which we had appropriate material, including a broad geographic sampling of N. tricarinatus and N. dilucidus as these two species had large ranges with some observed variation in the aedeagus. All specimens were preserved as frozen tissue samples since collection with the exception of the specimen from Costa Rica (SLE2381) which was pinned. Molecular extraction and sequencing methods follow those of
List of DNA voucher specimens and GenBank accession numbers used in this study. “N/A” indicates the gene fragment was not successfully amplified or sequenced.
Taxon | Extraction | Locality | COI Accession | 28S Accession |
---|---|---|---|---|
N. bicolor | SLE1810 | Suriname: Sipaliwini: Kabalebo Nature Resort | ON239446 | ON243733 |
N. bicolor | SLE2120 | Suriname: Sipaliwini: Kabalebo Nature Resort | ON239447 | N/A |
N. bifidus | SLE1113 | Venezuela: Amazonas: Tobogán de la Selva | ON239437 | ON243725 |
N. bifidus | SLE2369 | Venezuela: Amazonas: Tobogán de la Selva | ON239438 | ON243726 |
N. brunbadius | SLE1553 | Brazil: Amazonas: Ducke Reserve | ON239441 | ON243731 |
N. brunbadius | SLE2102 | Brazil: Amazonas: Ducke Reserve | ON239442 | ON243732 |
N. dilucidus | SLE0505 | Suriname: Brokopondo: Brownsberg Nature Park | ON239422 | N/A |
N. dilucidus | SLE0506 | Suriname: Brokopondo: Brownsberg Nature Park | ON239420 | ON243688 |
N. dilucidus | SLE1799 | Suriname: Sipaliwini: Kabalebo Nature Resort | ON239418 | ON243695 |
N. dilucidus | SLE1811 | Suriname: Sipaliwini: Kabalebo Nature Resort | ON239419 | ON243696 |
N. dilucidus | SLE2107 | Guyana: Region IX: Kusad Mountain | ON239411 | ON243702 |
N. dilucidus | SLE2108 | Guyana: Region VI: Upper Berbice | ON239417 | ON243692 |
N. dilucidus | SLE2113 | French Guiana: Crique Eau Chire | ON239406 | ON243693 |
N. dilucidus | SLE2114 | Suriname: Sipaliwini: Sipaliwini Savanna Reserve | ON239413 | ON243703 |
N. dilucidus | SLE2121 | Suriname: Brokopondo: Brownsberg Nature Park | ON239421 | ON243694 |
N. dilucidus | SLE2330 | Brazil: Roraima: ca. 13 km NE of Caroebe | N/A | ON243704 |
N. dilucidus | SLE2335 | Guyana: Region 9: Kusad Mountain | ON239412 | ON243705 |
N. dilucidus | SLE2339 | French Guiana: St. Laurent du Maroni (ca. 15 km SW) | ON239415 | ON243690 |
N. dilucidus | SLE2366 | Venezuela: Amazonas: Tobogán de la Selva | ON239404 | ON243687 |
N. dilucidus | SLE2368 | Guyana: Region 8: 7 km NW Chenapau | ON239409 | ON243699 |
N. dilucidus | SLE2374 | Brazil: Roraima: Rio Cocal, near Tepequem | ON239408 | ON243698 |
N. dilucidus | SLE2375 | Suriname: Sipaliwini: Wehepai | ON239416 | ON243706 |
N. dilucidus | SLE2377 | Brazil: Roraima: Serra do Tepequém | ON239410 | ON243700 |
N. dilucidus | SLE2378 | Venezuela: Bolívar: Piedra de la Virgen | ON239405 | ON243701 |
N. dilucidus | SLE2380 | Guyana: Region IX: Parabara | ON239414 | ON243707 |
N. dilucidus | SLE2383 | French Guiana: Carbet ONF Grillon | ON239407 | ON243697 |
N. dilucidus | SLE2389 | Suriname: Sipaliwini: Raleighvallen | ON239423 | ON243689 |
N. dilucidus | SLE2394 | Suriname: Sipaliwini: Raleighvallen | N/A | ON243691 |
N. garciae | SLE1900 | Brazil: Amazonas: Pres. Fig. (ca. 57 km E) | ON239440 | ON243730 |
N. giraldoi | SLE2088 | Brazil: Rondonia: Vale do Cachoeiras | ON239428 | ON243718 |
N. giraldoi | SLE2332 | Brazil: Rondonia: Ji-Parana (27 km SW) | ON239427 | ON243719 |
N. giraldoi | SLE2334 | Brazil: Rondonia: Ji-Parana (27 km SW) | ON239429 | ON243720 |
N. insignitus | SLE1115 | Venezuela: Bolívar: La Escalera | ON239443 | ON243734 |
N. juma | SLE1269 | Brazil: Amazonas: Novo Airão | ON239435 | ON243727 |
N. juma | SLE2100 | Brazil: Amazonas: Ducke Reserve | ON239436 | ON243728 |
N. liparus | SLE2111 | Venezuela: Aragua: Henri Pittier National Park | ON239403 | ON243714 |
N. liparus | SLE2123 | Venezuela: Barinas: Barinitas (ca. 13 km NW) | ON239401 | ON243715 |
N. liparus | SLE2124 | Venezuela: Mérida: Santo Domingo (ca. 2 km SE) | ON239400 | ON243716 |
N. liparus | MSC1820 | Venezuela: Aragua: Henri Pittier National Park | ON239402 | KC992598 |
N. lohezi | SLE2337 | French Guiana: Savane Roche Virginie | ON239444 | ON243737 |
N. lohezi | SLE2387 | French Guiana: Savane Roche Virginie | ON239445 | ON243738 |
N. parvus | SLE2388 | Suriname: Sipaliwini: Grensgeberte Moutains | ON239434 | ON243729 |
N. retusus | SLE2110 | Guyana: Region 8: ca. 7 km NW Chenapau | N/A | ON243735 |
N. retusus | SLE2372 | Guyana: Region 8: Ayanganna Airstrip | ON239439 | ON243736 |
N. tricarinatus | SLE1112 | Venezuela: Zulia: Tukuko | ON239425 | ON243710 |
N. tricarinatus | SLE2371 | Venezuela: Zulia: Tukuko | ON239426 | ON243711 |
N. tricarinatus | SLE2381 | Venezuela: Aragua: Henri Pittier National Park | ON239424 | ON243712 |
N. tricarinatus | SLE2391 | Venezuela: Portuguesa: Biscucuy | N/A | ON243713 |
N. tricarinatus | SLE2392 | Venezuela: Portuguesa: Biscucuy | N/A | ON243708 |
N. tricarinatus | SLE2397 | Costa Rica: Cartago: Tapanti National Park | N/A | ON243709 |
N. vatius | SLE2104 | Brazil: Bahia: Cachoeira Domingos Lopes | ON239430 | ON243722 |
N. vatius | SLE2324 | Brazil: Mato Grosso do Sul: Aquidauana (ca. 27 km S) | ON239432 | ON243724 |
N. vatius | SLE2327 | Brazil: Mato Grosso do Sul: Aquidauana (ca. 15 km E) | ON239433 | ON243723 |
N. vatius | SLE2385 | Brazil: Bahia: Livramento de Nossa Senhora | ON239431 | ON243721 |
N. sp. | SLE0092 | India: Tamil Nadu: Bodi Hills, Western Ghats | ON239399 | KC992599 |
N. sp. | SLE2140 | Peru: Cusco: 1 km N. Quince Mil | ON239398 | ON243717 |
We examined more than 900 specimens for this work, including the holotypes of most previously described Neotropical species. The specimens were examined using Olympus SZ61 and SZX7 microscopes and preparation of the specimens and dissection of the genitalia were carried out following the methodology of
SCC Private collection of Simon Clavier, Kourou, French Guiana;
From our integrated approach of combining morphological data with DNA sequence data from two genes, we found support for 18 species in the Neotropical region, including twelve new species. At the same time, we found that two of the most widespread species have been described multiple times, resulting in four new synonymies. For the 14 species that we had molecular data, the maximum likelihood analysis (Fig.
Among the 14 species for which we had molecular data, the minimum uncorrected pairwise distances in COI between any two species was 5.0% (between N. giraldoi and N. vatius) with the exception of N. dilucidus, in which the two specimens from Venezuela (SLE2366 and SLE2378) were 6% divergent from the remaining populations further to the east (Guyana, Brazil, Suriname, French Guiana). We sequenced one specimen from Peru that is known only from a single female, and we were unable to associate it with confidence to any of the described species for which we did not have DNA; however, due to the importance of the male genitalia in identifications, we refrain from describing this species here.
Notionotus shorti Queney, 2010, syn. nov.
Notionotus edibethae García, 2000, syn. nov.
Notionotus nucleus Perkins, 1979, syn. nov.
Notionotus perijanus García, 2000, syn. nov.
Notionotus Spangler, 1972: 139.
Notionotus rosalesi Spangler, 1972: 141; by original designation.
Small to very small beetles, total body length 1.5–2.0 mm. Color yellow, reddish brown, dark and pale brown to black. Body shape oval in dorsal view; moderately convex to convex in lateral view. Antennae with eight antennomeres. Maxillary palps short, nearly half the width of the head, second segment bending outwards, apical segment ~ 2 × as long as the penultimate segment (Fig.
As this revision only treats the New World species, we did not comprehensively examine the Old World species to generate a global genus description. Therefore, the diagnosis above should be considered for Neotropical species only. The Old World species of Notionotus are generally similar to the New World species, but some species do differ in significant characters: for example, N. suturalis Hebaeur, 2003 has a sutural stria and antennae with 9 antennomeres.
Body shape and coloration. The degree of convexity between species is variable; some are moderately convex, others weakly convex. The general dorsal coloration of the body among species ranges from yellow (e.g., N. tricarinatus, Fig.
Habitus of Notionotus spp.: A N. bicolor (paratype) B N. patamona (holotype) C N. brunbadius (holotype) D N. insignitus (paratype) E N. juma (paratype) F N. parvus (holotype) G N. garciae (paratype) H N. vatius (holotype) I N. giraldoi (paratype) J N. bifidus (paratype) K N. peruensis (holotype).
Mesoventrite. In Notionotus the elevation of the mesoventrite is composed of two or three laminae: one transverse ridge and one longitudinal (Fig.
Elytral punctation. The density of the ground punctation is typically sparse, and the degree of impression is variable between species within the genus. In some species, the ground punctation is very weakly impressed and may almost appear absent and low magnification (Fig.
Aedeagus. The shape of the aedeagus is the most important and often crucial feature to identify species of Notionotus. Most species in the Neotropics exhibit two different generalized aedeagal forms: in some species, the median lobe and basal piece have the same length, or the median lobe is slightly longer than the parameres (e.g., N. tricarinatus, Fig.
The species of this group can be diagnosed by the following combination of characters: (1) the shape of the elevation mesoventrite, having one transverse ridge and one longitudinal ridge (Fig.
Notionotus dilucidus Queney, 2010: 130.
Paratype (male): “♂ ”, “Notionotus/dilucidus n. sp./ PARATYPE/ P. QUENEY descr. 2010”, “Guyane [= French Guiana]: Roura,/ Cacao, Chemin/ Molokoi, crique,/ 16-IX-2009/ leg. P. Queney” (
Holotype (male): “♂ ”, “Notionotus shorti/n. sp. HOLOTYPE/P. QUENEY descr.2010”, “Guyana: Mazaruni-Potaro/District, Takutu Mountains,/ stream debris berlesed, 18-/XII-1983, leg. P.J. Spangler,/ W.E. Steiner & M. Levine” (
(362 exs.). Brazil: Roraima State: Serra do Tepequém, 3°47.334'N, 61°42.570'W, 14.i.2018, leg. Short, Benetti, & Santana, small forested stream, BR18-0114-02A (1 ex.,
The general coloration of Notionotus dilucidus is similar to N. giraldoi, N. mexicanus, and N. tricarinatus and these species are very difficult to differentiate with external characters alone. However, the shape of the aedeagus in N. dilucidus is quite distinct: the outer and inner margins of the parameres are sinuate, the apex of the parameres presents an indentation in which the depth can vary from very deep, slightly deep, or almost not distinguishable and pointing outwards (Fig.
Aedeagi of Notionotus dilucidus A N. dilucidus (paratype) B N. shorti (paratype) C–J N. dilucidus C specimen from Roraima, Brazil D specimen from Guyana E specimen from French Guiana F specimen from Suriname G specimen from Roraima, Brazil H specimen from Bolívar State, Venezuela I specimen from Amazonas State, Venezuela J specimen from Suriname.
Size and form
: Body length 1.6–1.8 mm. Body form elongate oval, moderately convex in lateral view (e.g., Fig.
There is significant variation in the apex of the parameres of the aedeagus. The paramere tip is weakly sclerotized, as evidenced by its very pale to white appearance in cleared specimens (e.g., Fig.
This species is widely distributed throughout the Guiana Shield region, from Tobogán de la Selva in Venezuela east to the coast of French Guiana (Fig.
This species is the most common and abundant species of the genus in the Guiana Shield region and is frequently collected in leaf packs or along the margins of forested streams. They are typically found in streams that are lined with detritus and have rocky or sandy substrates. Although some specimens have been collected in seepage-like habitats or adjacent to seepages, this does not seem to be a primary or favored habitat for this species.
To help interpret these morphological observations, we sequenced 22 specimens that span the entire width of the Guiana Shield, including specimens from Venezuela, Brazil, Guyana, Suriname, and French Guiana. These specimens also represented a range of paramere form diversity. In general, we found little meaningful molecular divergence among these populations (Fig.
Among all sequenced specimens, the maximum pairwise divergence in the gene COI was 6.0%. Although this is on the higher end of intraspecific divergence observed in hydrophilids, it is seen in some taxa (see
As both N. dilucidus and N. shorti were proposed in the same work (
Holotype (male): “BRAZIL: Rondonia: Novo [sic: Nova] Uniao/ -10.91764°, -62.377°, 359 m/ Vale do Cachoeiras; 10.vii.2018/ leg. Short; Margin of rocky/ stream; BR18-0710-02B” (
The dorsal coloration, shape of the elevation of the mesoventrite, area of pubescence on the metafemur and degree of impression of the ground punctation of Notionotus giraldoi are very similar to N. dilucidus, N. mexicanus, N. tricarinatus. It can be distinguished only by its aedeagus, including the unique shape of the parameres with a depression of the inner margin, as well as by the abrupt narrowing in the midlength of the median lobe (Fig.
Size and form
: Body length 1.7–1.9 mm. Body form elongate oval, convex in lateral view (Fig.
L. M. González-Rodríguez names this species in honor of Juan José Giraldo Gutiérrez in gratitude for the encouragement and support in her career.
Known from several localities in the Brazilian States of Rondonia and Mato Grosso do Sul (Fig.
This species was collected along the margins of two adjacent streams, one with a sandy bottom and one with a rocky bottom (Fig.
Notionotus liparus Spangler, 1972: 144.
Holotype (male): “VENEZUELA/ Bar., 24 Km. NW/ Barinitas/II-23-1969/P.&P. Spangler”, “Collected with/ 207 Oocyclus”, “HOLOTYPE/ Notionotus/liparus/P.J. Spangler” (
(139 exs.). Venezuela: Aragua State: Henri Pittier National Park, Río Curucuruma, 10°21.070'N, 67°34.920'W, 11.i.2006, leg. A.E.Z. Short, waterfall/seep, AS-06-023 (75 exs.,
Notionotus liparus can be recognized by the distinct black coloration among the other dark (reddish brown) species such as N. brunbadius, N. parvus and N. retusus, also for being the only dark species in the liparus group (Fig.
Size and form
: Body length 1.6–1.8 mm. Body form elongate oval, convex in lateral view (Fig.
This species is widespread in the Mérida Andes and Coastal Mountains of Venezuela (Fig.
This species is found in rock seepage habitats and wet rocks adjacent to waterfalls (Fig.
We sequenced four specimens from three Venezuelan states across the range of this species (Aragua, Barinas, and Mérida). The sequences are nearly identical (Fig.
Notionotus mexicanus Perkins, 1979: 306.
Holotype (male): “MEXICO, Oaxaca, 8 mi. E./Tapanatepec, tropical/stream with lg. boulders/3-VII-1974/ME&PD Perkins”, “Type No/76326/U S N M”, “HOLOTYPE/Notionotus/mexicanus/P.D.Perkins” (
Notionotus mexicanus is very similar morphologically to N. tricarinatus sharing characters such as body length, yellow dorsal coloration, pronotum and elytra with fine ground punctation, and elevation of the mesoventrite with a transversal and a longitudinal ridge. It can be distinguished by the shape of the aedeagus, specifically the shape of the parameres: inner margins straight and sinuate reaching the apex, parameres narrowing along apical third, and narrower than N. tricarinatus.
Size and form
: Body length 1.8 mm. Body form elongate oval, convex in lateral view (Fig.
Only known from the type locality in Mexico (Fig.
The type series was collected “from plant debris which had become trapped between stones in a rapid stream” (
Notionotus tricarinatus Perkins, 1979: 309.
Holotype (male): “PANAMA, C.Z./Albrook Forest Site/ground, 22-III-/1968, R.S. Hutton”, Type No/76324/U S N M”, “HOLOTYPE/Notionotus/tricarinatus/P.D. Perkins” (
Aedeagi of Notionotus tricarinatus A N. tricarinatus (holotype) B N. nucleus (holotype) C N. perijanus (holotype) D N. edibethae (holotype) E specimen from Barinas State, Venezuela F specimen from Portuguesa State, Venezuela G specimen from Aragua State, Venezuela H specimen from Zulia State, Venezuela I specimen from Costa Rica.
Holotype (male): “VENEZUELA. Trujillo/Mcpo. Rafael Rangel/La Guaira, Quebrada la/amarilla, 530 m./24-VIII-1997/Det. M. García, 199”, “Col:/M. García/J. Camacho/E. Gomez”, “Holotipo ♂/ Notionotus/edibethae/ Dcrip. M. García, 1997” (
Holotype (male): “GUATE., Alta Verapaz/5 mi. W. La Tinta/small tropical brook/6-VI-1974, ME&PD Perkins”, “Type No/76325/U S N M”, “HOLOTYPE/Notionotus/nucleus/P.D. Perkins” (
Holotype (male): [only the permanent slide mount of the aedeagus was examined] (
(173 exs.). Costa Rica: Cartago Province: Tapanti National Park, Building by Río Villegas, 29.v.2006, leg. A.E.Z. Short, HG-vapor light, AS-06-066 (5 exs.,
See differential diagnosis for Notionotus mexicanus.
Size and form
: Body length 1.6–1.9 mm. Body form elongate oval, strongly convex in lateral view (Fig.
Known from Guatemala, Costa Rica, Panama, and Venezuela (Fig.
This species is found along the margins and in leaf packs of streams in the mountains and foothills of the Northern Andes and Central America. It prefers gravelly or rocky streams, especially in the foothills where it may sometimes be abundant (Fig.
We examined more than 155 specimens from a dozen localities of this species from Guatemala to several chains of the Venezuelan Andes. Although there are subtle variations in the apex of the aedeagal parameres, this variation is relatively small and not correlated to geography, other morphological characters, or molecular data. These subtle variations in paramere shape likely explain why this species has been described four times, once each from Guatemala and Panama (
As both N. nucleus and N. tricarinatus were proposed in the same work (
Holotype (male): “BRAZIL: Mato Grosso do Sul/ -20.72281°, -55.69127°; 225 m/Aquidauana (c. 27 km S) on/MS-174; leg. Hamada & team;/27.vi.2018; seepage & debris nr./stream margin; BR18-0627-01E” (
Among the species of liparus group, Notionotus vatius can be recognized by the brown dorsal coloration, and quite unique color pattern of the head frons and medial region of the clypeus dark brown, lateral side of the clypeus pale brown. In addition, the shape of aedeagus, especially the apex of the parameres is broad, blunt, and pointing slightly outwards, and the gonopore has rounded shape and it is situated at midlength of median lobe.
Size and form
: Body length 1.7–2.3 mm. Body form elongate oval, strongly convex in lateral view (Fig.
The name is derived from the Latin word vatius meaning bent outwards, after the form of the parameres slightly pointing outwards of the aedeagus.
This species is known from several localities in Bahia and Mato Grosso do Sul States in Brazil (Fig.
This species was collected on seepages and along the margins of rocky streams (Fig.
Although the known localities of this species are widely dispersed in Brazil, they are from similar habitats at similar elevations on the Brazilian Shield. Moreover, the specimens from Bahia and Mato Grosso do Sul states are less than 3% divergent in uncorrected pairwise distances in COI.
Diagnosis. The lohezi species group can be distinguishable by the presence of three ridges in the elevation of the mesoventrite, two transverse ridge and one longitudinal (Fig.
Holotype (male): “SURINAME: Sipaliwini District/ 4.42313°N, 57.19198°W, 104 m/ Kabalebo Nature Resort/ Moi Moi Creek; 10–14.iii.2019/ leg Short & Baca small seeps/ SR19-0310-01F” (
Notionotus bicolor shares the bicolorous dorsal coloration that can also be observed in N. garciae, N. patamona, and N. lohezi. It can be recognized by the shape of the aedeagus, the median lobe is much shorter than in N. insignitus (Fig.
Size and form
: Body length 1.7–2 mm. Body form elongate oval, moderately convex in lateral view (Fig.
The name derived from the Latin words bi meaning two and color meaning hue, referring to the dorsal coloration of the species. This species has yellow coloration in the pronotum and black in the elytra.
Known from two localities in central Suriname: Kabalebo and the summit of Tafelberg Tepui (Fig.
At Kabalebo, this species was collected in several streams, including both along the margin, rock pools with detritus in the creek bed, and in seepage habitats (Fig.
Holotype (male): “VENEZUELA: Amazonas State/ 5°23.207'N, 67°36.922'W, 125 m/ Tobogan de la Selva; 8.viii.2008/leg. A. Short, M. García, L. Joly/ AS-08-080b; old “tobogancito”/on seepage area w/detritus” (
Notionotus bifidus can be separated from all other species of lohezi group by being the only species in the group that present uniformly dorsal yellow coloration (Fig.
Size and form
: Body length 1.5–1.7 mm. Body form elongate oval, moderately convex in lateral view (Fig.
The specific name comes from the Latin word bifidus meaning split into two parts, after the form of the median lobe “bifurcated apically” of the aedeagus.
Known only from the type locality in Venezuela (Fig.
This species was collected in seepage habitats that were covered with algae and detritus (Fig.
Holotype (male): “BRAZIL: Amazonas, Manaus/-2.93079, -59.97514, 75 m/ Ducke Reserve/ leg. Short & team; stream margin/ & assoc. backwater swampy area/ 9–10.vi.2018; BR18-0609-03A”, “DNA VOUCHER/ Extraction #/ SLE-1553” (
This species has a particular coloration pattern, dark reddish brown, which makes it easily distinguishable among the other species of the lohezi group. The shape of the parameres is slightly similar to N. lohezi (Fig.
Size and form
: Body length 1.7–1.9 mm. Body form elongate oval, moderately convex in lateral view (Fig.
The name is a combination of two Latin words brun meaning dark and badius meaning reddish brown, highlighting the distinguishable dark reddish brown color of this species.
Known only from the type locality near Manaus, Brazil (Fig.
The only known specimens were collected along the margin of a stream (Fig.
Holotype (male): “BRAZIL: Amazonas: Manaus/ -2.93079, -59.97514, 75 m/ Ducke Reserve, stream nr./ transect trail; leg. Short & team/ stream margins & leaf packs/ 9.vi.2018; BR18-0609-01A” (
Habitat of Notionotus spp. in the Andean region of Venezuela A Venezuela, Barinas State, seepage habitat of N. liparus (collecting event VZ12-0124-02A) B Venezuela: Aragua, Henri Pittier National Park, seepage habitat of N. liparus, (collecting event AS-06-023) C Venezuela: Zulia, Rio Tukuko, habitat of N. tricarinatus (collecting event VZ09-0129-01A) D Venezuela, Portuguesa, tributary of the Rio Guanare, habitat of N. tricarinatus (collecting event VZ09-0119-03X).
Notionotus garciae is very similar to N. juma in the bicolor dorsal coloration, the shape of the elevation of the mesoventrite and the pubescent area of the metafemora. It can be distinguished by the punctation of the pronotum, the elytra shallowly marked (in N. juma is moderately impressed, Fig.
Size and form
: Body length 1.7–1.8 mm. Body form elongate oval, convex in lateral view (Fig.
This species is named after Andrea Lorena García Hernández curator at the Colección de Insectos de la Universidad del Quindío (CIUQ) in recognition to her passion and contribution of the knowledge of the insects, specially hydrophilids in Colombia.
This species is known from several collecting events at the Ducke Reserve near Manaus, Brazil (Fig.
This species was collected along the margins of small streams in dense forest (Fig.
Holotype (male): “VENEZUELA: Bolívar State/ 6°2'10.5"N, 61°23'57.8"W, 630 m/ Along La Escalera; 31.vii.2008/ leg. A. Short, M. García, L. Joly/ AS-08-058; rocky stream” (
Habitat of Notionotus spp. A Venezuela: Amazonas State, type locality and habitat of N. bifidus (collecting event AS-08-080b) B Venezuela: Bolívar State: Type locality and habitat of N. insignitus (collecting event AS-08-058) C Guyana: Region 9: type locality and habitat of N. patamona (collecting event GY13-0318-01C) D Suriname: Kabalebo, habitat and type locality for N. bicolor (collecting event SR19-0310-01F).
See differential diagnosis for Notionotus bicolor. In addition, N. insignitus has a particular and unique color pattern in the elytra among the other congeners. The elytra are mostly dark brown, the posterior margins yellow and in the middle region has a characteristic yellow spot.
Size and form
: Body length 1.6–1.8 mm. Body form elongate oval, convex in lateral view (Fig.
The specific name comes from the Latin word insignitus meaning marked and refers to the distinctive yellow spot in the elytra of this species.
Known only from the type locality in southeastern Venezuela (Fig.
The only known series of this species was collected along the margin of a forested rocky stream (Fig.
Holotype (male): “Brazil: Amazonas: Manaus/ -2.93079, -59.97514, 75 m/ Ducke Reserve, Igarape Barro/ Branco; Short & team; muddy/ pools in swampy area by stream/ 9.vi.2018; BR18-0609-02B” (
See differential diagnosis for Notionotus garciae. In addition, the aedeagus of N. juma has an emargination in the apex of the medium lobe, being a particular feature among the species of the lohezi group.
Size and form
: Body length 1.6–1.9 mm. Body form elongate oval, convex in lateral view (Fig.
Habitat of Notionotus spp. A type locality and habitat of N. giraldoi (collecting event BR18-0710-02A) B type locality and habitat of N. vatius (collecting event BR18-0627-01A) C type locality and habitat of N. brunbadius (collecting event BR18-0609-03A) D type locality and habitat of N. garciae (collecting event BR18-0609-01A) E type locality and habitat of N. juma (collecting event BR18-0609-02B) F type locality and habitat of N. retusus (collecting event GY14-0311-02A).
This species is named after the Juma, an indigenous tribe located in the Açuã River, in the southern part of the state of Amazonas-Brazil.
Known only from the Ducke Reserve near Manaus, Brazil (Fig.
Specimens were collected in two habitats at the same forest reserve: in detrital pools in an area of shallowly flooded forest with detrital and mud substrate (Fig.
Notionotus lohezi Queney, 2010: 135.
Paratype (male): “♂ ”, “Notionotus lohezi/ n. sp. PARATYPE/ P. QUENEY descr. 2010”, “Guyane: Régina,/ Patawa, crique en/ forêt, 170 m,/ 13-IX-2009,/ leg. P. Queney” (
(2 exs.). French Guiana: Savane Roche Virginie, near RN 2, 4.1883, -52.13982, 64 m, Crique Chauve-souris, leg. Short, 10.iii.2020, solid granite substrate, detritus along margins on granite, FG20-0310-01A (2 exs.,
See differential diagnosis for Notionotus bicolor.
Size and form
: Body length 1.6–1.8 mm. Body form elongate oval, convex in lateral view (e.g., Fig.
This species is only known from a few localities in French Guiana (Fig.
This species was collected in rocky streams with detritus along margins.
Notionotus sp. 2 in Short, 2013: 88.
Holotype (male): “SURINAME: Sipaliwini District/ N 2.97731°, W 55.38500°, 200 m/ Camp 4 (low), Kasikasima; sandy/ stream on trail to METS camp/ 20.iii.2012; SR12-0320-02A/ leg. A. Short; 2012 CI-RAP Survey” (
Notionotus parvus can be recognized by the pale reddish yellow color in the head and pronotum and reddish brown in the elytra (Fig.
Size and form
: Body length 1.7–1.9 mm. Body form elongate oval, convex in lateral view (Fig.
The species name is derived from the Latin word parvus meaning little or small in reference to the small aedeagus size of this species.
This species was collected along the margins of a small, sandy-bottomed stream.
Holotype (male): “GUYANA: Region XIII [!sic: Region 8]/ 5°18.264'N, 59°50.257'W; 687 m/ Ayanganna Airstrip; trail from air-/ strip to Ayanganna; seepage area; over rocks in forest flowing into/ stream; leg. A. Short; 18.iii.2014/ GY14-0318-01C” (
See differential diagnosis for Notionotus bicolor.
Size and form
: Body length 1.6–1.8 mm. Body form elongate oval, convex in lateral view (Fig.
This species is named after the Patamona, an indigenous tribe located in the mountainous region from which this species is known.
Known from several closely situated localities in western Guyana (Fig.
This species was collected at several a variety of stream-associated habitats, including along the margins of detritus and sandy-based streams, as well as in rock seepage habitats adjacent to streams (Fig.
Holotype (male): “GUYANA: Region XIII [!sic: Region 8]/ 5°0.730'N, 59°38.965'W, 585 m/ Upper Potaro Camp I (c. 7 km/ NW Chenapau), Ridge Trial/ leg. Short, Baca and Salisbury/ 11.iii.2014; GY14-0311-02A”, “DNA VOUCHER/ Extraction #/ SLE-2110” (
The external characters of Notionotus retusus and N. lohezi are quite similar. The only way they can be separated is by the features of the aedeagus. In this species, the apex of the parameres is wide and rounded (acute in N. lohezi, Fig.
Size and form
: Body length 1.8 mm. Body form elongate oval, moderately convex in lateral view. Color and punctation: Dorsally bicolor, head mostly brown, frons brown, clypeus pale brown; pronotum yellow with two small black round spots along posterior margin; elytra dark brown. Ventrally brown; maxillary palps, mouthparts, antennae (antennal club slightly darker), legs brown. Clypeus and labrum with dense, fine, and weakly impressed ground punctation (punctures separated by 2 × their width); pronotum and elytra ground punctation fine, weakly impressed, and sparser than on head (punctures separated by 3 × their width). Head: Clypeus and labrum shallowly emarginate anteromedially, lateral margins of the labrum bearing setae. Thorax: Prosternum carinate medially, strongly raised, acute and pointing anteriorly. Elevation of mesoventrite with two transversal ridges, elevated medially, lateral sides concave; longitudinal ridge broad anteriorly and sharp posteriorly, the point where the three ridges merged wide and blunt (e.g., Fig.
The specific name comes from the Latin word retusus meaning rounded and notched, after the form of the apex of the median lobe of the aedeagus.
This species is only known from the type locality in western Guyana (Fig.
This species was collected in detrital-filled pools in a shallow ravine with dense forest cover (Fig.
Diagnosis. The species of this group can be recognized by the unique dorsal coloration, being almost yellow with a wide brown band in the third anterior of the elytra (Fig.
Notionotus rosalesi Spangler, 1972: 141
Holotype (male): “VENEZUELA/Arag., 10 Km S./Rancho Grande/II-14-1969/P.&P. Spangler”, “TypeNo/71950/U S N M”, “HOLOTYPE/ Notionotus/rosalesi/P.J.Spangler” (
Trinidad: Guanapo State: 4.1 km up Guanapo Valley, trib of Guanapo River, 460 ft, 11-VII-2005 (1 ex.,
Notionotus rosalesi can be distinguished by the wide brown band in the anterior third of the elytra, as well as, the unique shape of the genitalia, having many accessories at the base of the parameres, the apex of the parameres membranous and lanceolate.
Size and form
: Body length 1.8–1.9 mm. Body form elongate oval, moderately convex in lateral view (Fig.
Originally described from the Venezuelan states of Aragua and Barinas (
Although specific habitat information is limited, all specimens were collected in association with streams.
The genitalia of the holotype appears to have some modest fungal growth on the median lobe (Fig.
Diagnosis. The species of peruensis group can be diagnosed by the dorsal coloration completely yellow, the elevation of the mesoventrite with one transverse and one longitudinal (e.g., Fig.
Holotype (male): “PERU: Dept. Madre de/ Dios: Pantiacolla Lodge,/ Alto Madre de Dios R./ 12°39.3'S, 71°13.9'W 420 m/ 14–19-XI-2007 D. Brzoska/ ex. flight intercept trap/ PER1B07 004” (
Notionotus peruensis can be distinguished by the particular shape of the aedeagus, being nearly rectangular in the basal half and abruptly narrow in the apical half (Fig.
Size and form
: Body length 1.6 mm. Body form elongate oval, convex in lateral view (Fig.
The species is named after Peru, the country where it was collected, as well as for being the first species described for the genus in this country.
Known only from the type locality in Peru (Fig.
The single specimen was collected at a flight intercept trap; nothing is known about its habitat.
(2 exs.). Bolivia: Santa Cruz: Amboro National Park, Guarda Parque Mataracu, 21–27.xi.2004, malaise trap, leg. Robertson, García, & Vidaurre (1 female,
We examined two single female specimens from unique localities that we refrain from identifying or describing, due to the lack of male genitalia for comparison. The specimen from Peru likely represents an undescribed species, as suggested by its distant position in our DNA tree (Fig.
Although it is fairly straightforward to key specimens (especially males) to species group, identification to species within each group almost always requires examination of the aedeagus. For this reason, as well as the fact that there are no doubt many yet-to-be-described species, particularly in the southern Amazon region, we do not include a species key here.
1 | Elevation of the mesoventrite composed by two ridges (one transverse and one longitudinal) (e.g., Fig. |
2 |
– | Elevation of the mesoventrite composed by three ridges (two transverse and one longitudinal) (e.g., Fig. |
lohezi species group |
2 | Length of the parameres nearly the same as the length of the basal piece. Length of the median lobe almost the same as the parameres (e.g., N. liparus, Fig. |
liparus species group |
– | Length of the parameres longer than the basal piece. Length of the median lobe almost the same as the parameres | 3 |
3 | Outer margin of the parameres convex and tapering reaching the apex, rounded apex. Median lobe rectangular along basal half then tapering abruptly (e.g., N. peruensis sp. nov., Fig. |
peruensis species group |
– | Outer margin of the parameres sinuate, apex of the parameres lanceolate. Median lobe wide and approximately triangular (e.g., N. rosalesi, Fig. |
rosalesi species group |
Since Notionotus was first described as a hygropetric genus from the Andean Region of Venezuela fifty years ago (
We are grateful for the assistance and support of many colleagues during fieldwork, including Mauricio García (