Research Article |
Corresponding author: Andrew Edward Z. Short ( aezshort@gmail.com ) Academic editor: Mariano Michat
© 2017 Alex T. Kohlenberg, Andrew Edward Z. Short.
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The genus Tobochares Short & García, 2007 is revised. A combination of morphological and molecular data provide support for ten distinct species. Five new species are described: T. canaliculatus sp. n. (Venezuela), T. canthus sp. n. (Venezuela), T. emarginatus sp. n. (Suriname), T. kusad sp. n. (Guyana), and T. pallidus sp. n. (Venezuela). All four preexisting species are redescribed. A tenth species, known from a single female from Venezuela, is left undescribed pending the collection of additional specimens. New collecting records are provided for T. sulcatus Short & García, 2007 and T. kasikasima Short, 2013. Tobochares sipaliwini Short & Kadosoe, 2011 is newly recorded from Guyana. All species are associated with seepage or wet rock habitats, although some species have also been found along the margins of streams that flow over rocky substrates. High-resolution images including scanning electron micrographs are provided, as well as a key to species and habitat photographs.
South America, Guiana Shield, taxonomy, new species
The genus Tobochares Short & García, 2007 was erected for a single unusual species found living under leaves on wet rock along a creek in southern Venezuela (
Depositories of examined material.
Morphological methods. Specimens were examined using an Olympus SZX16 microscope (to 110× magnification). Specimens for dissection were relaxed in warm water, and their genitalia were removed and placed in glycerin on a glass slide, which was then viewed and imaged using an Olympus BX51 to 200× magnification. Genitalia were mounted beneath the specimens in microvials with glycerin. Scanning electron micrographs were taken by mounting specimens on carbon tape and coated in gold. Habitus photographs were taken with a Visionary Digital imaging system. All final images were created by stacking multiple individual photographs from different focal planes using the software Zerene Stacker. Morphological terminology largely follows
Molecular methods. Total genomic DNA was extracted from entire beetles using a DNeasy kit (Qiagen, Alameda, CA). All vouchers (Table
List of specimens and GenBank accession numbers used in this study. All vouchers are deposited in
Taxon | Extraction | Locality | Coordinates | GenBank Accession |
---|---|---|---|---|
T. emarginatus | SLE424 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679835 |
T. emarginatus | SLE482 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679836 |
T. emarginatus | SLE483 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679837 |
T. kasikasima | SLE1045 | Suriname: Kappel Airstrip | 3.791317, -56.149467 | KY679850 |
T. kasikasima | SLE1046 | Suriname: Kappel Airstrip | 3.791317, -56.149467 | KY679851 |
T. kasikasima | SLE1048 | Suriname: Tafelberg Summit | 3.926667, -56.188332 | KY679849 |
T. kasikasima | SLE1049 | Suriname: Tafelberg Summit | 3.926667, -56.188332 | KY679852 |
T. kasikasima | SLE1050 | Suriname: Tafelberg Summit | 3.926667, -56.188332 | KY679848 |
T. kusad | SLE1021 | Guyana: Kusad Mts. | 2.80885, -59.865 | KY679846 |
T. pallidus | SLE525 | Venezuela: Tobogan de la Selva | 5.386783, -67.615364 | KY679853 |
T. sipaliwini | SLE422 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679838 |
T. sipaliwini | SLE478 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679839 |
T. sipaliwini | SLE497 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679840 |
T. sipaliwini | SLE1020 | Guyana: Kusad Mts. | 2.80885, -59.865 | KY679841 |
T. sipaliwini | SLE1023 | Suriname: Kwamala | 2.182883, -56.787251 | KY679842 |
T. striatus | SLE423 | Suriname: Kasikasima | 2.976883, -55.411385 | KY679847 |
T. sulcatus | SLE0035 | Venezuela: Tobogan de la Selva | 5.386783, -67.615364 | KC935327 |
T. sulcatus | SLE1027 | Venezuela: Tobogan de la Selva | 5.386783, -67.615364 | KY679845 |
T. sulcatus | SLE1035 | Venezuela: Tobogan de la Selva | 5.386783, -67.615364 | KY679843 |
T. sulcatus | SLE1037 | Venezuela: Pijiguaos | 6.593617, -66.820633 | KY679844 |
T. sp. A | SLE526 | Venezuela: Tobogan de la Selva | 5.386783, -67.615364 | KY679854 |
The results of the maximum likelihood analysis (Fig.
1. Tobochares canaliculatus sp. n. Venezuela (Amazonas)
2. Tobochares canthus sp. n. Venezuela (Amazonas)
3. Tobochares emarginatus sp. n. Suriname
4. Tobochares kasikasima Short, 2013 Suriname
5. Tobochares kusad sp. n. Guyana
6. Tobochares pallidus sp. n. Venezuela (Amazonas, Bolivar)
7. Tobochares sipaliwini Short & Kadosoe, 2011 Suriname, Guyana
8. Tobochares striatus Short, 2013 Suriname
9. Tobochares sulcatus Short & García, 2007 Venezuela (Amazonas, Bolivar)
10. Tobochares sp. A Venezuela (Amazonas)
Species of Tobochares, in general, are morphologically well-defined and we found relatively little variation within species.
Dorsal coloration. The dorsal coloration of most species is medium to dark brown (Figs
Head. A lateral canthus of the frons emarginates the eye to some extent in all Tobochares species (e.g. Fig.
Thoracic venter. The condition of the mesoventrite slightly varies between species, but is not extremely useful for making identifications. In most species, it possesses a low transverse ridge which may vary in elevation (Fig.
Elytra. The condition of the elytra is extremely useful for separating species (Fig.
Abdomen. The surface of the abdomen is densely pubescent in all species, but some species have cuticular projections mixed in amongst the setae (Fig.
Tobochares Short & García, 2007: 2.
Tobochares sulcatus Short & García, by original designation.
Size small, 1.5–2.4 mm. Antennae with eight antennomeres, including three-segmented club. Elytra without sutural stria, but with serial punctures impressed into distinct grooves (serial punctures present but not impressed into groves in T. canthus, T. emarginatus, and T. pallidus). Median elevation of mesoventrite low, forming a narrow transverse ridge or elevated bulge (Fig.
Head. Antennae with eight antennomeres, including three-segmented pubescent club. Maxillary palps curved inward and moderately long, as long or longer than the width of head just anterior to eyes; inner face of palpomere 2 straight to slightly curved; apical palpomere slightly longer than penultimate. Labial palps short, distinctly shorter than mentum width. Mentum flat and set with a few scattered setae; strongly emarginated anteromedially with a notch extending posteriorly about one-quarter to one-third of its length. Head with ground punctures. Frons with series of irregular systematic, setae-bearing punctures anterior to each eye. Systematic punctures also present on clypeus and labrum but blend with ground punctation. Eyes not bulging, continuous with outline of the head; slightly to strongly emarginated anteriorly by a small extension of the frons (Figs
Larvae. The immature stages are unknown.
Venezuela (Amazonas, Bolivar), Guyana, Suriname.
Nearly all collections of Tobochares are associated with hygropetric habitats, e.g. thin water films on rock. A handful of specimens have been collected from stream or forest pool margins that are fed by or adjacent to such habitats. Most seepages on which Tobochares tend to be found are littered with leaves and detritus (e.g. Figs
Holotype (male): “VENEZULEA: Amazonas State/ 5°23.207'N, 67°36.922'W, 125m/ Tobogan de la Selva; 8.viii.2008/ leg. A. Short, M. García, L. Joly/ AS-08-080b; old “tobogancito”/ on seepage area w/ detritus”, “[barcode]/ SEMC0877724/ KUNHM-ENT” (
The combination of the pale dorsal coloration and deeply sulcate elytra along their entire length (Fig.
Size and form: Body length 1.6–2.0 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head brown to dark brown, anterolateral margins of clypeus with prominent pale preocular patches (Fig.
Dimunuative of canalis, referring to the elytral grooves.
Known only from the type locality in Venezuela (Fig.
This species has been collected on several occasions on rock seepages along the margin of the Rio Coromoto (Fig.
Holotype (male): “VENEZULEA: Amazonas State/ 5°23.207'N, 67°36.922'W, 125m/ Tobogan de la Selva; 8.viii.2008/ leg. A. Short, M. García, L. Joly/ AS-08-080b; old “tobogancito”/ on seepage area w/ detritus”, “[barcode]/ SEMC0877726/ KUNHM-ENT” (
The lack of impressed striae on the elytra (Fig.
Size and form: Body length 1.7–2.0 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head dark brown, frons darker (nearly black) laterally and around eyes, anterolateral margins of clypeus with paler preocular patches (Fig.
Named after the condition of the eyes, in which a lateral canthus of the frons partially divides them into lower and upper surfaces.
Known only from the type locality in Venezuela (Fig.
This species was collected on a rock seepage along the margin of the Rio Coromoto (Fig.
Holotype (male): “SURINAME: Sipaliwini District/ N2°58.613', W55°24.683', 400m/ Camp 4 (high) Kasikasima/ leg. A. Short; main seepage area/ 24.iii.2012; SR12-0324-01C/ 2012 CI-RAP Survey”, “[barcode]/ SEMC1088469/KUNHM-ENT” (
The lack of impressed striae on the elytra, strongly emarginated eye (Fig.
Size and form: Body length 1.7–2.1 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head brown, frons darker laterally and around eyes, anterolateral margins of clypeus with paler preocular patches (Fig.
Named after the condition of the eyes, in which a lateral canthus of the frons partially divides them into lower and upper surfaces.
Known from several localities in central and southern Suriname (Fig.
The majority of specimens were collected on a large granite seepage on Kasikasima (Fig.
Tobochares kasikasima Short, 2013: 83.
Holotype (male): “SURINAME: Sipaliwini District/ N2°58.613', W55°24.683', 400m/ Camp 4 (high) Kasikasima/ leg. A. Short; main seepage area/ 24.iii.2012; SR12-0324-01C/ 2012 CI-RAP Survey”, “[barcode]/ SEMC1088588/KUNHM-ENT” (
(59). SURINAME: Sipaliwini: Same data as type (1 ex.,
The weakly impressed striae limited to the posterior third of the elytra (Fig.
Size and form. Body length 1.6–2.0 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head very dark brown to black, anterolateral margins of clypeus with very faint paler preocular patches (Fig.
The species was originally described from a single male from Mt. Kasikasima in south-central Suriname. It has subsequently been collected in longer series at several other localities in central and southern Suriname including from the summit of Tafelberg Tepui (Fig.
The first specimen of this species was found in a seepage at the base of Kasikasima (Fig.
Holotype (male): “GUYANA: Region IX/ 2 48.531'N, 59 51.900'W, 170m/ Kusad Mts., Mokoro Creek/ main seepage area/ leg. Short, Isaacs, Salisbury/ 27.x.2013; GY13-1027-03B”, “[barcode]/ SEMC1271353/ KUNHM-ENT” (
This species can be distinguished from most species by the impressed striae running the full length of the elytra and its overall dark brown coloration (Fig.
Size and form: Body length 2.0–2.4 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head very dark brown to black, anterolateral margins of clypeus with paler preocular patches (Fig.
Named after Kusad Mountain in the South Rupununi region of Guyana, from where the species is known.
Only known from the type locality in Guyana (Fig.
The species was collected on a thin rock seepage flowing over granite that was associated with a small creek (Fig.
Holotype (male): “VENEZULEA: Amazonas State/ 5°23.207'N, 67°36.922'W, 125m/ Tobogan de la Selva; 8.viii.2008/ leg. A. Short, M. García, L. Joly/ AS-08-080b; old “tobogancito”/ on seepage area w/ detritus”, “[barcode]/ SEMC0877702/ KUNHM-ENT” (
This small species may be easily distinguished by the combination of its very pale dorsal coloration and lack of impressed elytral striae (Fig.
Size and form: Body length 1.5–1.9 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head very pale brown; maxillary palps pale (Fig.
Named for the species’ relatively pale body coloration.
Known from several localities along the northwestern shoulder of the Guiana Shield in Venezuela (Fig.
All six collecting events of this species were from rock seepages. The type locality and longest series of specimens were collected on a rock seepage along the margin of the Rio Coromoto (Fig.
Because this species co-occurs with several other Tobochares in Venezuela, the extremely pale coloration makes specimens easily confused for teneral individuals of other species in the field until they can be examined under higher magnification.
Tobochares sipaliwini Short & Kadosoe, 2011: 85.
Holotype (male): “SURINAME: Sipaliwini District/ 2°10.973'N, 56°47.235'W, 210 m/ Camp 2, on Sipaliwini River/ leg. Short & Kadosoe; Inselberg/ 29–30. viii.2010; SR10-0829-01A/ 2010 CI-RAP Survey” (
(120). SURINAME: Sipaliwini: Same data as type except 1.ix.2010, seep on inselberg, SR10-0901-01A (1 ex.,
This species can be distinguished from most other Tobochares by the elytral striae being impressed only on the posterior half (Fig.
Size and form: Body length 1.7–2.1 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head very dark brown to black, anterolateral margins of clypeus with paler preocular patches (Fig.
Originally described from an inselberg on the Suriname-Guyana boarder, this species has now been found at other localities in both countries (Fig.
The most frequently encountered species of the genus thus far in the eastern Guianas, T. sipaliwini has been found in a variety of seepage habitats, though most have been associated with more permanent flowing water (Fig.
Tobochares striatus Short, 2013: 83.
Holotype (male): “SURINAME: Sipaliwini District/ N2.24554°, W55.77000°, 800m/ Camp 2 Grensgebergte Rock/ leg. A. Short; rock seepages/ 12.iii.2012; SR12-0312-01A/ 2012 CI-RAP Survey” (
(1). Camp 4 (Kasikasima), 24.iii.2012, leg. A. Short, main seepage area, SR12-0324-01C (1 ex.,
See differential diagnosis for T. kusad.
Size and form: Body length 1.8–2.1 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head very dark brown to black, anterolateral margins of clypeus with paler preocular patches (Fig.
Known only from two localities in south-central Suriname (Fig.
Tobochares sulcatus Short & García, 2007: 4.
(100). VENEZUELA: Amazonas: Tobogan de la Selva, 14.i.2009, leg. Short, García, Miller & Joly, wet rock covered with detritus, VZ09-0114-01F (50 exs.,
The dark brown coloration and deep elytral grooves, which are impressed along their entire length (Fig.
Size and form: Body length 1.8–2.2 mm. Body elongate oval, moderately dorsoventrally compressed. Color and punctation. Dorsum of head very dark brown to black, anterolateral margins of clypeus with paler preocular patches (Fig.
Known from a series of localities along the northwestern edge of the Guiana Shield in Venezuela (Fig.
Tobochares sulcatus occurs on wet rocks with fallen leaves and other detritus along stream corridors (Fig.
(1). VENEZUELA: Amazonas: Tobogan de la Selva, leg. M. Balke (1 female,
This species is morphologically very similar to T. pallidus, and shares most diagnostic features of that species (in particular its very pale coloration). This species can be separated from T. pallidus by the faint but distinctly impressed striae in the posterior quarter of the elytra, and the slightly less emarginated eyes.
This species is only known from a single specimen, which we also extracted for DNA. Molecular data also supports this taxon as sister to T. pallidus, to which it is also similar morphologically. We refrain from describing the species until additional specimens, including ideally a male, can be found.
Habitat of Tobochares spp. in Suriname. A–B Base of Voltzberg, collecting event SR16-0316-01C, habitat of T. sipaliwini C–D Waterfall in the upper Palumeu watershed, collecting event SR12-0311-03A, habitat of T. emarginatus sp. n. E Base of Voltzberg, collecting event SR16-0316-01A, habitat of T. sipaliwini F Kasikasima, collecting event SR16-0324-01C, habitat of T. sipaliwini, T. kasikasima, T. striatus, and T. emarginatus sp. n. Red arrows point to example microhabitats where specimens were collected
Habitat of Tobochares spp. in Guyana and Suriname. A–B Guyana, Kusad Mt., collecting event GY13-1027-03B, type locality for T. kusad sp. n. C–D Suriname, side of Kappel Airstrip, collecting event SR13-0824-02B, habitat of T. kasikasima E–F Suriname, summit of Tafelberg, Caiman Creek seepage, collecting event SR13-0819-01A, habitat of T. kasikasima. Red arrows point to example microhabitats where specimens were collected.
Habitat of Tobochares spp. in Venezuela. A–B Tobogan de la Selva, upstream, collecting event AS-08-080b, type locality for T. canaliculatus sp. n., T. sulcatus, T. canthus sp. n., and T. pallidus sp. n. C Tobogan de la Selva, collecting event VZ09-0114-01D D Outcrop near Pijiguaos, collecting event VZ10-0708-01B, habitat of T. pallidus sp. n. E–F Rock outcrop near Pijiguaos, collecting event VZ10-0709-01B, habitat of T. pallidus sp. n. Red arrows point to example microhabitats where specimens were collected.
1 | Elytra with impressed grooves along their entire length (e.g. Figs |
2 |
– | Elytra with impressed grooves in posterior half or less, or with grooves absent (e.g. Fig. |
5 |
2 | Apical maxillary palpomere uniformly pale (Fig. |
canaliculatus sp. n. |
– | Apical maxillary palpomere darkened at least at apex, and sometimes on distal half or more (Fig. |
3 |
3 | Punctures within elytral grooves small, grooves appearing fairly smooth (Fig. |
.sulcatus Short & García |
– | Punctures within elytral grooves strongly impressed and distinct (Fig. |
4 |
4 | Apical maxillary palpomere with apex ranging from slightly to distinctly darkened (Fig. |
kusad sp. n. |
– | Apical maxillary palpomere darkened in at least distal half (Fig. |
striatus Short |
5 | Elytra with grooves on posterior third or posterior half, grooves most prominent near elytral suture (e.g. Fig. |
6 |
– | Elytra without grooves or with weak grooves on posterior quarter (e.g. Fig. |
7 |
6 | Elytra with grooves on posterior half (Fig. |
sipaliwini Short & Kadosoe |
– | Elytra with grooves on posterior third (Fig. |
kasikasima Short |
7 | Dorsum of head, pronotum, and elytra uniformly pale (Fig. |
8 |
– | Dorsum of head, pronotum, and elytra brown to dark brown (e.g. Fig. |
9 |
8 | Elytra without grooves (Fig. |
pallidus sp. n. |
– | Elytra with weak grooves on posterior quarter. Eyes emarginate at lateral margin, narrowing to slightly less than half of the width | sp. A |
9 | Aedeagus (Fig. |
canthus sp. n. |
– | Aedeagus (Fig. |
emarginatus sp. n. |
The water beetle seepage fauna of the Guiana Shield was essentially completely undescribed little more than 15 years ago. Since that time, numerous new lineages have been discovered in a variety of families, including Dytiscidae (e.g. Fontidessus), Noteridae (Tonerus Miller, 2009), and Hydroscaphidae (Confossa
Most sampled localities had only a single, or occasionally two species of Tobochares co-occurring in the same habitat. However, two particular sites had exceptional diversity with five and four species respectively: Tobogan de la Selva in Venezuela, and Mt. Kasikasima in Suriname. As there was no species composition overlap between them, a remarkable nine of the ten species covered in this revision could be collected by visiting just these two places.
Given how relatively little of the Guiana Shield has been surveyed, especially for hygropetric taxa, it is almost a certainty many more Tobochares species are left to be described.
The fieldwork that generated these specimens would not have been possible without the invaluable assistance of many colleagues, including Jesus Camacho and Mauricio Garcia (