Research Article |
Corresponding author: Alexander V. Martynov ( centroptilum@gmail.com ) Academic editor: Lyndall Pereira-da-Conceicoa
© 2020 Alexander V. Martynov, Dmitry M. Palatov.
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:
Martynov AV, Palatov DM (2020) A new species of Indoganodes Selvakumar, Sivaramakrishnan & Jacobus, 2014 (Ephemeroptera, Teloganodidae) from Sri Lanka. ZooKeys 969: 123-135. https://doi.org/10.3897/zookeys.969.56025
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A new species, Indoganodes tschertoprudi sp. nov. is described from Sri Lanka. The genus Indoganodes Selvakumar, Sivaramakrishnan & Jacobus, 2014 was previously known only by one species from the Western Ghats (India). The new species differs from Indoganodes jobini Selvakumar, Sivaramakrishnan & Jacobus, 2014 by the number of denticles on the claws, shape of the femora, shape of the chalazae on the femora, absence of any median tubercles on the terga, and presence of posterolateral processes only on segments VI–IX. The diagnosis of Indoganodes is also emended. Morphological larval affinities of Indoganodes and Ephemerellina Lestage, 1924 and the probable origin and diversification of I. tschertoprudi sp. nov. are discussed.
Indomalayan realm, larva, mayflies, Pannota, type material
Teloganodidae Allen, 1965 is a relatively small family distributed within the Indomalayan realm and southern part of the Afrotropical realm. Endemism is typical for this family, and the many of species in the family have a restricted distribution. Moreover, the Afrotropical and Indomalayan realms are represented by different genera. Four genera of the family occur in Indomalayan realm: Teloganodes Eaton, 1882, Dudgeodes Sartori, 2008, Derlethina Sartori, 2008, and Indoganodes Selvakumar, Sivaramakrishnan & Jacobus, 2014. The family Teloganodidae in the Indomalayan region is currently undergoing a detailed investigation; 22 species and three of four genera mentioned above were described during last 12 years (
Until now, the genus Indoganodes was known only from the Western Ghats (India) and only by the larval stage of the sole species, Indoganodes jobini Selvakumar, Sivaramakrishnan & Jacobus, 2014.
In this paper, a new species of Indoganodes from Sri Lanka is described based on the larval stage. Detailed observations of the larval features of this new species and its comparison with I. jobini, the type species from southern India, allows for the emendation of the generic diagnosis.
All material were preserved in 80–95% EtOH; some paratypes were mounted with Canada balsam on slides.
Administrative districts and geographical coordinates of localities are given according to Google Earth (http://earth.google.com). Photographs were made using a Canon Power Shot A 630 with Ulab XY-B2T microscope in the National Museum of Natural History, National Academy of Sciences of Ukraine (NMNH
The type material now is housed in the NMNH
Holotype : larva (slide 672, mounted with Canada balsam), Sri Lanka, border of Central and Sabaragamuwa provinces, vicinity of Marathenna village, mountain slope, helocrene in valley of large stream, 6.751333, 80.686167, 1390 m a.s.l., Chertoprud M.V. leg., 5.ii.2017 – IN Sri1Ingsp. Paratypes: 1 larva (slide 673, mounted with Canada balsam), ibid., Chertoprud M.V. leg., 5.ii.2017 – IN Sri1Ingsp. 1 larva (in slide 674 with Euparal), Sri Lanka, Central Province, vicinity of Holmwood Estate, stream (section with almost no current), 6.826389, 80.724444, 1660 m a.s.l., Chertoprud M.V. leg., 4.ii.2017 – IN Sri2Ingsp.
This species is named after Dr Mikhail V. Chertoprud (Moscow, Russia), who provided the material for this study.
Indoganodes tschertoprudi sp. nov. can be distinguished from the only other known representative of Indoganodes, I. jobini, by the following combination of characters: (i) tarsal claw with row of 5–8 large, blunt denticles and several (1–3) small, pointed denticles among the large ones (Fig.
Larva: body length 8.7–12.5 mm; caudal filaments partially detached, their length ratio to body unknown, paracercus not rudimental. Body light brown (Fig.
Head. Genae small; head without any protuberances; surface of head covered with small hair-like setae and small scale sockets (Fig.
Thorax. Dorsal surface covered with short, thin, hair-like setae and scattered empty scale sockets; tubercles and ridges absent (Fig.
Femora of all legs robust, with longitudinal ridge; outer margin without apical projections (Fig.
Indoganodes tschertoprudi sp. nov., larva, type specimens A fore femur, dorsal view B middle femur, dorsal view C hind femur, dorsal view D transversal band of stout setae on fore femur, dorsal view E stout setae of dorsal surface of fore femur F chalazae with stout setae on inner margin of hind femur G, H tarsal claws.
Dorsal surface of fore femur with indistinct wide, transversal band of short and medium-sized, oval, stout setae bearing feathered margins and short and medium-sized, feathered, stout setae with divergent margins and cleft at apex (Figs
Ventral surfaces of fore tibia and tarsus with numerous differently shaped, stout setae on inner margin and along it; main types of stout setae are: long, stout setae with feathered margins and pointed apex (Fig.
Dorsal surfaces of middle and hind femora covered with oval and rounded, medium-sized, feathered, stout setae with cleft at apex in some (Fig.
Patella-tibial suture on tibiae of middle and hind legs distinctly shorter than that on fore leg. Setation of middle and hind tibiae and tarsi near that of fore leg, but in contrast to fore leg, outer margins of these tibiae bear short, feathered, oval, stout setae; in immature larvae, row of stout setae more dense and distinct.
Tarsal claw of all legs robust, hooked, its surface covered with several medium-sized, thin, hair-like setae. Claw with row of 5–8 large, blunt denticles and several (1–3) small, pointed denticles among the larger ones (Fig.
Abdomen. All terga without any median tubercles (Fig.
Segment I without gills; gills present on abdominal segments II–VI (Fig.
Basal part of caudal filaments with feathered, stout setae and stout, hair-like setae at articulations; stout setae shorter, mainly oval on dorsal side of the filaments and elongated on ventral side.
Winged stages : unknown.
Larvae of new species were found in wooded gullies in the mountains of Sri Lanka in the subtropical altitudinal zone (altitude 1390–1660 m a.s.l.) (Fig.
Distribution of Indoganodes representatives (A), and habitats of Indoganodes tschertoprudi sp. nov. (B, C) A map of genus Indoganodes distribution B type locality, helocrene spring in valley of large stream, vicinity of Marathenna village, border of Central and Sabaragamuwa Provinces, Sri Lanka (February 2017, photo by M.V. Chertoprud) C small stream, section with almost no current, vicinity of Holmwood Estate, Central Province, Sri Lanka (February 2017, photo by M.V. Chertoprud).
The new species of Indoganodes reveals features that enable us to emend the diagnosis of the genus as follows: characters (i) and (iv–vi) stay unchanged. Emended characters are (ii) abdominal posterolateral processes well developed on segments VI–IX; the processes on segments I–V absent or poorly developed; (iii) claw with one row of denticles, claw with up to eight large denticles and three small denticles, they might alternate in row. Additional characters are: (vii) glossae and paraglossae deeply divided and bluntly pointed; (viii) forefemur not flattened, without distinct and regular, transversal row of stout setae; (ix) outer margin of fore femora without long stout setae; (x) paracercus not reduced.
The Gondwanan origin of the Teloganodidae (McCafferty and Wang 1977) and the close relationship of Indoganodes with Ephemerellina Lestage 1924 (
The genus Indoganodes is most similar to Ephemerellina in the combination of some larval characters (winged stages of both Indoganodes species are not described): (i) shape of labrum, (ii) fore femur not significantly flattened, (iii) absence of distinct, narrow, transversal row of stout setae on fore femur, (iv) inner margin without a row of setae continuing on dorsal surface near articulation with trochanter, (v) absence of filamentous gill I, (vi) gills present on segments II–VI, (vii) semi-operculate dorsal lobe of gill II, (viii) deep division of glossae and paraglossae, (ix) unreduced paracercus, and some other characters.
Indoganodes and Ephemerellina are isolated biogeographically, with Ephemerellina from the Afrotropical realm and Indoganodes from the Indomalayan realm. It is probable that they share a common ancestor from the African continent. After eastern Gondwana, including also India and Sri Lanka, had broken free of Africa about 100 million years ago, these genera evolved separately. Sri Lanka later split from India, and since the Pliocene (5.33–2.58 million years ago), the geographic position of Sri Lanka has been similar to that at present. However, during the periodic low sea levels in the Pleistocene (2.58–0.0117 million years ago), there was a land bridge between India and Sri Lanka, and two-way dispersal of mainly terrestrial fauna was facilitated. The last land bridge was cut off by rising sea levels 5,000–8,000 years ago as the Pleistocene gave way to warmer climates and northern glaciers retreated during the Holocene (
Presently, only the narrow Palk Strait separates Sri Lanka and India. Although mayflies have winged stages capable of dispersal, the teloganodid fauna of the island shares no species with India or other countries of Indian subregion, which is in contrast to the vast number of other mayfly families (
The authors are grateful to Dr M.V. Chertoprud (Moscow, Russian Federation) for sampling material on described here species. The authors are also grateful to Dr C. Selvakumar (The Madura College, Madurai, Tamil Nadu, India) for providing additional photographs of the type specimen of Indoganodes jobini.