Research Article |
Corresponding author: Thomas Kaltenbach ( thomas.kaltenbach@bluewin.ch ) Academic editor: Ben Price
© 2017 Thomas Kaltenbach, Jean-Luc Gattolliat.
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:
Kaltenbach T, Gattolliat J-L (2017) New species of Indocloeon Müller-Liebenau from South-East Asia (Ephemeroptera, Baetidae). ZooKeys 723: 43-60. https://doi.org/10.3897/zookeys.723.20578
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One new species of Indocloeon Müller-Liebenau from Brunei, I. spathasetis sp. n., and one new species from Indonesia, I. timorense sp. n., are described and illustrated based on their larvae. The total number of known Indocloeon species increases from two to four and the generic attributes of the larvae are amended based on the examination of the new species. Results on the genetics of some species (COI) as well as comments on the distribution of I. indonesiae Kluge are also provided.
Brunei, COI, distribution, Indonesia, Oriental Region, taxonomy
The genus Indocloeon was established by
The specimens were collected by Kate Baker (King’s College London, UK) in the Ulu Temburong National Park in Brunei Darussalam with surber and kick-sampling methods (
The specimens were preserved in 70%–96% ethanol. The dissection of larvae was done in Cellosolve (2-Ethoxyethanol) with subsequent mounting on slides with Euparal liquid, using an Olympus SZX7 stereomicroscope.
The DNA of part of the specimens was extracted using non-destructive methods allowing subsequent morphological analysis (see
Species | Locality | Specimen catalog # | GenBank # (COI) | GenSeq Nomenclature |
---|---|---|---|---|
I. spathasetis sp. n. | Brunei | GBIFCH 00280816 | MF414701 | genseq-1 COI |
I. spathasetis sp. n. | Brunei | GBIFCH 00280817 | MF414702 | genseq-2 COI |
I. indonesiae | Flores | GBIFCH 00280818 | MF414703 | genseq-4 COI |
I. indonesiae | Flores | GBIFCH 00280819 | MF414704 | genseq-4 COI |
I. indonesiae | Sumbawa | GBIFCH 00280820 | MF414705 | genseq-4 COI |
I. indonesiae | Sumbawa | GBIFCH 00280821 | MF414706 | genseq-4 COI |
I. timorense sp. n. | Timor | GBIFCH 00280822 | MF414707 | genseq-1 COI |
Drawings were made using an Olympus BX43 microscope. Photographs of larvae were taken using a Canon EOS 6D camera and the Visionary Digital Passport imaging system (http://www.duninc.com) and processed with the programs Adobe PhotoShop Lightroom (http://www.adobe.com) and Helicon Focus version 5.3 (http://www.heliconsoft.com). Photographs were subsequently enhanced with Adobe Photoshop Elements 13.
The distribution map was generated with the program Simple Mapper (http://research.amnh.org/pbi/maps) and the program GEOLocate (http://www.museum.tulane.edu/geolocate/web/WebGeoref.aspx) was used to attribute approximate GPS coordinates to sample locations of
The taxonomic descriptions presented herein were generated with a DELTA (
Larva. Following combination of characters: A) middle length of antenna with conspicuous large spines at outer lateral margin (Fig.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown (Fig.
Antenna (Fig.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Terga (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the noticeable spatulate submarginal dorsal setae of the labrum.
Only known from Brunei, but presence in other regions of Borneo such as Sarawak and Sabah (Malaysia) is possible as their fauna remains poorly known.
The specimens were collected in lowland tropical rainforest at an altitude of about 100 m a.s.l., directly at the confluence of small tributaries with large rivers (Belalong, Temburong) as well as in upstream pools of these tributaries (Fig.
Holotype. Larva (on slide, GBIFCH 00280816), Brunei, Temburong District, Ulu Temburong National Park, 4°32.77'N, 115°09.52'E, May 2014, leg. Kate Baker. Paratypes. Brunei, Temburong District, Ulu Temburong National Park, May 2014, leg. Kate Baker: larva (on slide, GBIFCH 00280817), 4°32.77'N, 115°09.52'E; two larvae (one on slide, GBIFCH 00465131; one in alcohol, GBIFCH 00515214), 4°33.67'N, 115°08.87'E; two larvae (one on slide, GBIFCH 00465130; one in alcohol, GBIFCH 00515213), 4°33.64'N, 115°09.07'E; larva (on slide, GBIFCH 00465132), 4°33.39'N, 115°10.03'E; larva (on slide, GBIFCH 00465133), 4°33.21'N, 115°09.31'E; two larvae (one on slide, GBIFCH 00465134; one in alcohol, GBIFCH 00515215), 4°32.87'N, 115°09.47'E. All material deposited in the Museum of Zoology Lausanne (MZL).
Five larvae (in alcohol, GBIFCH 00515216). Brunei, Temburong District, Ulu Temburong National Park, near Kuala Belalong Field Studies Centre, tributary to Temburong river, 4°33.21'N, 115°09.31'E, May 2014, leg. Kate Baker. Deposited in the Museum of Zoology Lausanne (MZL).
Larva. Following combination of characters: A) labrum with submarginal arc of setae composed of one central medium, simple seta plus six medium, simple setae (Fig.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown. Head and thorax with bright dorsal line, forewing pads with bright striation (Fig.
Antenna with scape and pedicel subcylindrical; flagellum with broad spines on apex of each segment and with scales. In the middle part of the flagellum without large spines on apex of segments at outer lateral margin.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Middle leg (Fig.
Terga (Fig.
Gills (Fig.
Paraproct (Fig.
After the type locality, the island of Timor (Indonesia).
Indonesia: Timor.
The specimen was collected at an altitude of 1580 m a.s.l.
Holotype. Larva (on slide, GBIFCH 00280822), Indonesia, Timor, Mt. Mutis, 1580 m, 01.10.2011, 9°38.12'S, 124°12.80'E, leg. M. Balke. Temporary deposited in the Museum of Zoology Lausanne (MZL) before definitely housed in the Museum Zoologicum Bogoriense (MZB) in Indonesia.
In addition to the two new species found in Brunei and Timor, the occurrence of Indocloeon indonesiae on two further Indonesian islands (Flores and Sumbawa) is documented, additionally to Lombok, from where it was described by
Distribution of Indocloeon: a World map (http://www.pixabay.com) b Sri Lanka c Indonesia (partim) and Brunei (yellow stars, I. primum; red square, I. spathasetis sp. n.; green triangles, I. indonesiae (from left to right: islands of Lombok, Sumbawa, Flores); blue spot, I. timorense sp. n. (island Timor)).
Species | Locality | GPS coordinates |
I. indonesiae | Sumbawa | 8°38.54'S, 118°30.31'E |
Flores | 8°42.92'S, 122°04.41'E | |
I. timorense sp. n. | Timor | 9°38.12'S, 124°12.80'E |
I. spathasetis sp. n. | Brunei | 4°32.77'N, 115°09.52'E |
4°33.64'N, 115°09.07'E | ||
4°33.67'N, 115°08.87'E | ||
4°33.21'N, 115°09.31'E | ||
4°33.39'N, 115°10.03'E | ||
4°32.87'N, 115°09.47'E |
COI sequences were obtained from two specimens of I. indonesiae from Flores (Indonesia) and two from Sumbawa (Indonesia), one specimen of I. timorense sp. n. from Timor (Indonesia) and two specimens of I. spathasetis sp. n. from Brunei. Only very limited genetic distances were found between specimens of I. indonesiae from Flores and Sumbawa (Table
The generic description was given by
Based on the additional two new species, the following adaptations to the generic diagnosis are proposed: the maxillary palp differs between the species, always with three segments and longer than galea-lacinia, but may be very long and slender (I. primum, fig. 1e in
Taking into account the taxa described herein, the genus Indocloeon now encompasses four species, all of them occurring in the Oriental realm. In general, the different known species of Indocloeon can be easily identified by a combination of a few characters and often even by a single unique character. Overall important characters are the dorsal submarginal arc of setae of the labrum, the shape of the distomedial protuberance of segment II of the labial palp, the shape of the maxillary palp, the number of larger denticles of the claw and also the number of marginal spines of the paraproct.
Indocloeon primum can be recognised by the pointed distomedial protuberance of segment II of the labial palp, the long and slender maxillary palp with a very short segment III and the basally fused spines at the posterior margin of the terga (
Contrary to mouthparts, legs and abdomen, which are showing important specific differences within Indocloeon, both mandibles seem to be quite uniform inside the genus.
Indocloeon indonesiae and I. timorense sp. n. are morphologically the most similar species, especially when referring to the labrum submarginal arc of setae and the labial palps. However, the maxillary palps are clearly different. This similarity is corroborated by the genetic distance (K2P) based on COI (Table
From I. indonesiaeCOI sequences were obtained from specimens of the Indonesian islands Sumbawa and Flores, but not from Lombok from where the species was originally described (
The presently known distribution of the genus Indocloeon encompasses distant areas like the Indian peninsula on one hand and some South Asian islands on the other hand. Similar distributions can be found in other lineages of Baetidae such as Liebebiella Waltz and McCafferty or Chopralla Waltz and McCafferty (
We sincerely thank Kate Baker (King’s College London, UK) for the allocation of precious mayfly material to the Museum of Zoology Lausanne (MZL), for the photos of larval habitats, and for corrections of the English language. This material was collected during ecological studies in Brunei Darussalam, funded by Natural Environment Research council (NERC), and was in collaboration with Universiti Brunei Darussalam. We are also deeply grateful to Michael Balke (Zoologische Staatssammlung München,