Research Article |
Corresponding author: Alexander V. Martynov ( centroptilum@gmail.com ) Academic editor: Ben Price
© 2022 Alexander V. Martynov, T. Sivaruban, Dmitry M. Palatov, Pandiarajan Srinivasan, S. Barathy, Rajasekaran Isack, Michel Sartori.
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, Sivaruban T, Palatov DM, Srinivasan P, Barathy S, Isack R, Sartori M (2022) Contribution to the knowledge of Teloganodidae (Ephemeroptera, Ephemerelloidea) of India. ZooKeys 1113: 167-197. https://doi.org/10.3897/zookeys.1113.85448
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Two new species of Dudgeodes Sartori, 2008 and a new species of Teloganodes Eaton, 1882 are described from India; they are Dudgeodes selvakumari Martynov & Palatov, sp. nov. from Himalayan region (Uttarakhand), Dudgeodes molinerii Sivaruban, Martynov, Srinivasan, Barathy & Isack, sp. nov., and Teloganodes barathyae Sivaruban, Martynov, Srinivasan & Isack, sp. nov. from the Tamil Nadu part of the Western Ghats. Thus, for now, the Teloganodidae fauna of India includes 11 species. Dudgeodes selvakumari sp. nov. appears to be significantly extend northward the known distribution of Dudgeodes. Partial COI sequences were used as an initial clustering method to show the relationships of D. selvakumari sp. nov. with other sequenced operational taxonomic units (OTU) of the genus.
COI, distribution, imago, larva, morphology, Pannota, Tamil Nadu, Uttarakhand
The superfamily Ephemerelloidea is a relatively diverse group within Indian subcontinent. This article is the next contribution in a series of papers on the superfamily of the region. Ephemerelloidea has been actively studied during the last years, and, as a result, series of new species mainly of the family Ephemerellidae have been described from this territory (
Teloganodes and Dudgeodes occur within Indomalayan realm only. Teloganodes consists of eight species distributed in the Indian subcontinent (
Representatives of Teloganodes are well distinguished at larval stages from other genera of the family (
In the present contribution we describe three new species from India: Dudgeodes selvakumari Martynov & Palatov, sp. nov. based on larval, imaginal, and egg stages; Dudgeodes molinerii Sivaruban, Martynov, Srinivasan, Barathy & Isack, sp. nov. based on larval and egg stages, and Teloganodes barathyae Sivaruban, Martynov, Srinivasan & Isack, sp. nov. based on the larval stage only.
Larvae were collected by hand picking and kick-net sampling in Uttarakhand Pradesh and Tamil Nadu, India. Winged stages were reared from larvae in Martynov-designed grow nets (Fig.
Specimens of Dudgeodes selvakumari sp. nov. and their body parts unmounted on slides were photographed using a Leica M205A microscope with a Leica Z16 APO apochromatic zoom system and Leica DFC450 camera. The photographs were processed with LAS Core v. 3.8. Body parts mounted on slides were photographed with a Ulab XY-B2T microscope with a Canon Power Shot A 630 camera. These photographs were processed with Adobe Photoshop CS5 and Helicon Focus v. 6. Larvae studied with a scanning electron microscope were dehydrated in ethanol and then critical-point dried. Scanning electron microscopy was done on a Vega3 Tescan SEM.
Larval morphological characters of D. molinerii sp. nov. and Teloganodes barathyae sp. nov. were studied using a LABOMED Luzeo 6Z stereo zoom microscope with an AR 6 Pro camera. Specimens studied under SEM were dehydrated in ethanol and critical-point dried, then examined using a Zeiss EVO 18 SEM. The photographs were processed using Adobe Photoshop 7.0 when necessary.
Names of protuberances of thorax (excluding lateral anterior tubercles, LAs) are given according to
Type material on D. selvakumari sp. nov. is deposited in collection of first author in the National Museum of Natural History of the National Academy of Sciences of Ukraine, Kyiv, Ukraine (
Total genomic DNA was extracted from three specimens of D. selvakumari sp. nov. using the BioSprint 96 extraction robot (Qiagen Inc., Hilden, Germany) following the supplier’s instructions. We used the non-destructive protocol described by
Alignment of analyzed sequences was made in BioEdit v. 7.0.5.3. Recently, both tree- and distance-based methods of species delimitation based on single-locus data have been used.
We calculated genetic distances within and between species and other taxa were calculated in MEGA v. 11 (
GenBank accession numbers for newly derived sequences are given in Table
Species | Specimen catalogue number | Locality | GPS Coordinates | Date | GenBank ID | GenSeq nomenclature |
---|---|---|---|---|---|---|
Dudgeodes selvakumari sp. nov. | GBIFCH00970940 | India, Uttarakhand, Mailani Range, vicinity of Garjiya village, unnamed river – left tributary of Kosi River | 29.4732°N, 79.1640°E | 1.v.2018 | ON255658 | genseq-2 COI |
Dudgeodes selvakumari sp. nov. | GBIFCH00970941 | ON255659 | genseq-2 COI | |||
Dudgeodes selvakumari sp. nov. | GBIFCH00970942 | ON255660 | genseq-2 COI |
Holotype
: imago ♂, with corresponding larval and subimaginal exuviae, India, Uttarakhand, Mailani Range, vicinity of Garjiya village, unnamed river – left tributary of Kosi River, 29.4732°N, 79.1640°E, 430 m a.s.l., 22.v.2018, A.V. Martynov & D.M. Palatov leg., Indi9Telsp/1 (
The new species is named in honour of Dr C. Selvakumar of India, who contributed significantly to the study of mayflies in India.
Mature larva. Body length 3.0–5.5 mm; cerci length 3.5–6.2mm. Dorsal surface of body yellowish with brown-black spots and strokes (Fig.
Head
dirty yellow with indistinct brown smudges. Antennae also dirty yellow, distal segments of flagellum and distal part of scapus blackish. Dorsal part of male eyes brown (Fig.
Mouthparts. Labrum wide and compact, width/length ratio 2.64–2.65, with smooth medial concavity on anterior margin (Fig.
Mandibles slender with few small setae along outer margin and one stout, hair-like seta in middle of margin. Right mandible (Fig.
Maxilla (Fig.
Hypopharynx with long, feathered setae on the rounded apexes of superlinguae, and very short setae on lingua (Fig.
Labial palp three-segmented, slightly constricted towards apex; articulation between segments clearly visible; segment III elongate and rounded apically, length/width at base ratio 1.9–2.2 (Fig.
Thorax. Pronotum with three pairs of tubercles: SMs, SLs, and Ls; with a few short globular stout setae; M tubercle absent (Fig.
Larva of Dudgeodes selvakumari Martynov & Palatov, sp. nov., paratypes A head and pronotum, dorsal view B row of setae at outer margin of head C thorax, dorsal view D sub-median tubercle (SM) of pronotum E thorax, lateral view. Abbreviations: white arrows show tubercles of pronotum and mesonotum. Scale bars: 0.3 mm (A, C, E); 0.1 mm (B, D).
Forefemur moderately slender, ca 2.1 times longer than wide; outer margin covered with regular row of long, stout, hair-like setae, and few thin, hair-like setae (some setae in bunches); submarginal row of setae distinct, composed of stout setae (elongate and short) with slightly divergent margins (Fig.
Middle and hind femora (Fig.
Tarsal claws moderately hooked, lacking subapical denticles, with 4–6 medial denticles and several (3–5) subapical setae (Fig.
Abdomen. All terga with moderately developed, narrowed (especially on terga V–X) median tubercles (Fig.
Larva of Dudgeodes selvakumari Martynov & Palatov, sp. nov., paratypes A–C abdomen, dorsal A ventral B and lateral C views D sublateral area of posterior margin of tergum VI E median tubercle of tergum VII F terga VIII–X, dorsal view G, H dorsal surface of tergum VIII, SEM microscopy G and light microscopy (H) I caudal filaments. Scale bars: 0.3 mm (A–C); 0.03 mm (D, G); 0.1 mm (E, F, I).
Posterior margin of terga I–V with row of long, stout, hair-like setae; posterior margin of terga VI–IX with row of elongate (on tergum VI) and short (all other terga) stout setae with rounded apices (Fig.
Gills on segments II–V (Fig.
Cerci length subequal to the body length, posterior margins of central segments with hair-like and forked stout setae; length of the stout setae less than length of corresponding segment (Fig.
Subimagos. Wings wholly grey, semitransparent in subimagos of both sexes.
Male imago. Body length: 5.6–6.5 mm; forewing length: 5.6–5.9 mm; cerci length: 11.0–13.3. General coloration brown; thorax dark brown. Turbinate eyes brick-colored (Fig.
Imago male of Dudgeodes selvakumari Martynov & Palatov, sp. nov., holotype A total lateral view B abdominal segments VII–X, lateral view C genitalia, ventral view D abdomen, ventral view E head and fore leg, lateral view. Abbreviations: white arrows show postero-lateral projections of segments; black arrows show remnants of gill sockets. Scale bars: 1 mm (A); 0.2 mm (B, C); 0.5 mm (D, E).
Fore leg (Fig.
Main area of fore wing transparent (Figs
Abdominal terga IV–VIII with small pointed median tubercles, in some specimens these tubercles distances on terga IV–V only (Fig.
Female imago. Body length: 6.0 mm; forewing length: 5.9 mm; cerci length: 12.8 mm. General coloration brown. Legs coloration as in male imago. Turbinate eyes brown.
On fore leg outer tarsal claw hooked and pointed and inner claw blunt. On middle and hind legs outer tarsal claw blunt and inner hooked and pointed. Wings venation as in male imagos, but longitudinal veins browner. Only abdominal segments VII–IX with distinct rounded apically postero-lateral projections, largest on segments VIII and IX. Abdominal terga IV–VIII with small pointed median tubercles. Tergum X with longitudinal distinctly divergent median concavity that reach posterior margin. Segments II–V with remnants of gill sockets. Subgenital plate not elongate, with wide and shallow concavity. Subanal plate rounded.
Egg
(dissected from mature larva). Shape (Fig.
Himalaya (Uttarakhand, India). All Indian representatives of Dudgeodes, excluding D. selvakumari sp. nov., are known from the Western Ghats only. Dudgeodes selvakumari sp. nov., which is distributed in the lower down part of Great Himalayan mountain range, is the most northern representative of the genus and family within India. This new species and D. lugens (Navás, 1933), which is known by single female subimago from Zhou Shan Island, in Zhejiang province, China (
Larvae of this species were collected in a mid-sized river (6–10 m wide) in a shallow woodland valley at an altitude of about 400 m a.s.l. in the southern foothills of the Great Himalaya Range (Nainital District, Uttarakhand state, India). The river was relatively warm (24–26 °C), had a current of moderate velocity (ca 0.3–0.7 m/s), and was with a mainly stony or rocky substrate. The river is located in the recreational zone of the Jim Corbett National park with a weak anthropogenic load. Larvae were collected from the riparian zone from stones or vegetation at local current velocity 0.05–0.2 m/s (Fig.
The new species can be distinguished from other representatives of the genus by the following combination of characters. Larva: (i) dorsal part of male eyes brown; (ii) antennae length 1.15 times head width, flagellum with about 15 or more segments; (iii) labrum with transversal band of long, stout, hair-like setae; (iv) prothorax with three pairs of tubercles: SMs, SLs, and Ls; mesothorax with an MP, pair of SMMs and pair of LAs; (v) forefemur without transversal row of stout setae; (vi) outer margin of forefemur covered with a regular row of long, stout, hair-like setae, a few bunches, and single, thin, hair-like setae; (vii) submarginal row of setae of forefemur distinct, consisting of elongate and short, stout setae with slightly divided margins; (viii) tarsal claw of all legs with 4–6 medial denticles and without subapical denticles; (ix) terga I–X with moderately developed, narrowed (especially on terga V–X) median tubercles; the largest tubercles on terga V–VIII; tubercle of tergum X narrow and pointed; in lateral view, median tubercles of terga I–IX distinctly elongate; (x) posterolateral projections moderately developed on segments VI–IX, slightly marked on segments II–V. Imago male: (i) fore wing with numerous cross-veins; (ii) hind wing with 3–4 cross-veins between Sc and RA, and two cross-veins between RA and IRA; (iii) penis lobes maximum width on about 0.7 of their length; (iv) abdominal terga IV–VIII with small pointed median tubercles; in some specimens these tubercles distinct on terga IV–V only; (v) abdominal segments VI–IX with rounded apically postero-lateral projections. Egg: (i) without spines on pole opposite to polar cap; (ii) surface covered with microgranules.
The larva of D. selvakumari Martynov & Palatov, sp. nov. is easily distinguished from other Indian Dudgeodes species by: (i) absence of tubercles on head; (ii) number of tubercles on pro- and mesonotum; (iii) forefemur setation; (iv) shape of fore femur; (v) absence of subapical denticles on tarsal claws; (vi) shape of gill II; (vii) shape of median tubercles of abdominal terga.
Holotype : mature ♀ larva, India, Tamil Nadu, Theni district, Kurangani hills, Kottakudi River, 10.0809°N, 77.2552°E, 632 m a.s.l., 28.x.2020, Pandiarajan Srinivasan & Isack Rajasekaran leg., ZSI–SRC/I/E/654. Paratypes: 5 larvae, ibid., 28.x.2020, Pandiarajan Srinivasan & Isack Rajasekaran leg., ZSI–SRC/I/E/655 (1 larva), AMC ZN 237 (4 larvae).
The new species is named in honour of Dr Carlos Molineri of Argentina, who contributed significantly to the study of mayflies.
Mature larva. Body length 4.7–4.9 mm; cerci length subequal to body length. General coloration of the dorsal side of head dirty yellow, with dark brown to blackish maculations; thorax and abdomen dark brown to blackish, with dirty yellow maculations (Fig.
Larva of Dudgeodes molinerii Sivaruban, Martynov, Srinivasan, Barathy, Isack, sp. nov., paratypes A total dorsal view B total ventral view C head, thorax and abdomen, dorsal view D head, dorsal view E row of setae at outer margin of head F head and thorax, dorso-lateral view G tarsal claw H gill II I gill III J gill IV K gill V. Abbreviations: white arrows show tubercles.
Head
with pair of occipital tubercles (Fig.
Mouthparts. Labrum compact, width/length ratio 2.51–2.53; with smooth anterior emargination; dorsal surface with transversal row of scattered, stout, hair-like setae (Fig.
Thorax. Pronotum with three pairs of tubercles: SMs, SLs, and Ls; tubercles with a few short, rounded setae. Mesonotum with three pairs of tubercles: two pairs of SMMs, a pair of LAs, and unpaired MP (Fig.
Forefemur broad, ca 1.3 times longer than wide (Fig.
Larva of Dudgeodes molinerii Sivaruban, Martynov, Srinivasan, Barathy & Isack, sp. nov., paratypes A, D fore femur B middle femur C hind femur E, F stout setae of transversal row on forefemur G outer margin of fore femur H abdomen I eggs, light microscopy J egg, SEM microscopy K cluster of spines on pole of egg. Abbreviations: sII – abdominal segment II, sX – abdominal segment X. Scale bars: 0.2 mm (H); 0.02 mm (J); 0.01 mm (K).
Middle and hind femora, in contrast to fore femur, more slender, ca 1.8–2 times longer than wide, with denser submarginal row of short, stout setae (Fig.
Tarsal claw moderately hooked, bearing 3–6 medial denticles, 1–2 subapical denticles (if two subapical denticles present, they are situated each on opposite sides of claw) and a row of 3–4 subapical setae on dorsal and ventral sides (Fig.
Abdomen. All terga with median tubercles that bear short, stout setae with slightly divergent margins. Median tubercles moderately developed on terga IV–VIII, and slightly marked on terga I–III, IX, and X (Fig.
Gills on segments II–V (Fig.
Cerci length subequal to the body length; posterior margin of proximal half segments with elongate, stout setae with rounded apices; posterior margin of distal half segments with long, spine-like setae on the lateral margins; length of the stout setae less than length of corresponding segment. Paracercus absent.
Egg.
Ovoid, ca 100–110 μm long, with numerous micropyles. Egg with one polar cap, on the opposite pole with a cluster of 18–20 spines (Fig.
Winged stages. Unknown.
Western Ghats (Tamil Nadu, India).
The larvae of D. molinerii sp. nov. inhabit cobble and pebble substrates of rivers with a strong current (Fig.
Dudgeodes molinerii sp. nov. can be distinguished from other Dudgeodes species by the following combination of characters. Larva: (i) dorsal part of male eyes dark brown to black; (ii) head with pair of small occipital tubercles; (iii) antennae length 1.25 times head width, flagellum with 11 segments; (iv) labrum with transversal row of scattered, stout, hair-like setae; (v) forefemur with transverse row of about 30 long, apically pointed, stout setae; (vi) tarsal claw bearing 3–6 medial denticles, and 1–2 subapical denticles (if two, they are on opposite sides of claw), and 3–4 subapical setae on dorsal and ventral sides; (vii) pronotum bears three pairs of tubercles: SMs, SLs, and Ls; mesonotum bears three pairs of tubercles: two pairs of SMMs, a pair of LAs, and unpaired MP; (viii) median tubercles moderately developed on terga IV–VIII, and slightly marked on terga I–III, IX and X; (ix) posterolateral projections moderately developed on segments VI–IX, and slightly marked on segments II–V. Egg: (i) egg with cluster of 18–20 spines present on pole opposite to polar cap; (ii) surface without microgranules.
Larval stage of this new species can be easily distinguished from other Indian Dudgeodes by: (i) presence of tubercles on head; (ii) number of tubercles on pro- and mesonotum; (iii) shape of femora; (iv) setation of forefemur; (v) size and shape median tubercles on abdomen.
Holotype : ♀ larva, India, Tamil Nadu, Theni District, Kurangani Hills, Kottakudi River, 10.0809°N, 77.2552°E, 632 m a.s.l., 28.x.2020, Pandiarajan Srinivasan & Isack Rajasekaran leg., ZSI–SRC/I/E/652. Paratypes: 2 larvae, ibid., 28.x.2020, Pandiarajan Srinivasan & Isack Rajasekaran leg., ZSI–SRC/I/E/653 (1 larva), AMC ZN 230 (1 larva).
The new species is named in honour of Dr S. Barathy, an assistant professor in the Department of Zoology, Fatima College, Tamil Nadu, India, who contributed to the study aquatic insects of India.
Mature larva. Body length up to 5.4–5.7 mm without cerci; cerci length subequal to body length. General colouration of dorsal side of head, thorax, and terga I–IX brownish to blackish; tergum X yellowish to whitish; ventral side of the head and thorax pale, sterna I–VIII with submedian black tinges; sternum IX pale (Fig.
Larva of Teloganodes barathyae Sivaruban, Martynov, Srinivasan & Isack, sp. nov., paratypes A total view, dorsal view B total view, ventral view C head and thorax, dorsal view D pronotum, dorso-lateral view E head, dorsal view F abdomen, ventral view. Abbreviations: white arrows show tubercles.
Head. Lateral margins of head fringed with a row of long, stout setae, forked near base and with pointed apices, which run from posterior margin of eyes to labrum; anterior margin of clypeus with numerous stout setae of the same type (Fig.
Mouthparts. Labrum compact, ca 2.4 times wider than long, with smooth anterior emargination; dorsal surface with a transversal band of numerous feathered setae (Fig.
Thorax. Pronotum with three pairs of rounded tubercles: SMs, SLs, and Ls. Mesonotum with three pairs of tubercles: two pairs of SMMs and LAs (lateral anterior tubercles), and an unpaired MP tubercle (Fig.
Forefemur (Fig.
Middle and hind femora with ornamentation similar to foreleg (Fig.
Tarsal claw hooked, bearing four medial denticles and two subapical denticles on opposite sides of claw; dorsal and ventral surface of claw with a row of 3–5 subapical, hair-like setae (Fig.
Abdomen. Median tubercles on terga I–X present; on tergum I poorly developed; on terga II–IV moderately developed; on terga V–X most developed. In dorsal view tubercles I–IX broad and rounded apically, tubercle X distinctly slender and bluntly pointed (Fig.
Gills present on abdominal segments II–VI. Gill II with dorsal lamella operculate and covering others, oval with margin entire; gills II–V with well-developed flabelliform ventral lobe; gills III–V with dorsal lamella incised medially (Fig.
Central portion of cerci with elongate stout setae with bluntly pointed apices and few long, hair-like setae; stout setae length less than half length of the corresponding segment (Fig.
Winged stages. Unknown.
Western Ghats (Tamil Nadu, India).
The same as for D. molinerii sp. nov.
Larva of T. barathyae sp. nov. can be distinguished from other species of Teloganodes by the following combination of characters: (i) dorsal surface of labrum with a transversal band of numerous feathered setae; (ii) inner incisor of the left mandible with two teeth inserted transversely, one smaller and pointed, the other large and rectangular; (iii) superlinguae angular laterally, with a row of long, feathered setae at apex; (iv) forefemur moderately broad, ca 2.4 times longer than wide; outer margin with regular row of long, stout, hair-like setae; without any combination of thin and stout setae in a row; (v) forefemur bears submarginal row of numerous short stout setae with divergent margins, (some of them divided near apex into two rounded lobes); same stout setae scattered over whole dorsal surface; (vi) fore femur without transverse row of stout setae; (vii) median tubercles on terga I–X, on tergum I poorly developed; on terga II–IV moderately developed; on terga V–X best developed; in dorsal view tubercles I–IX broad and rounded apically, tubercle X distinctly slender and bluntly pointed; (viii) posterolateral projections on segments II–IX, segments VII–IX well developed but not extremely.
Type habitats A, B, D, E of new species and cages (Martynov’s construction) C used for Dudgeodes selvakumari Martynov & Palatov, sp. nov. winged stages rearing A, B, D type habitat of Dudgeodes selvakumari Martynov & Palatov, sp. nov. C Martynov-designed grow nets for mayfly winged stages rearing E type habitat of Dudgeodes molinerii Sivaruban, Martynov, Srinivasan, Barathy & Isack, sp. nov. and Teloganodes barathyae Sivaruban, Martynov, Srinivasan & Isack, sp. nov. Abbreviations: arrows show microhabitats with the highest density of a new species’ larvae.
Larvae of this new species can be distinguished for other Indian representatives of Teloganodes by: (i) shape of superlinguae; (ii) length of antennae; (iii) absence of transversal row of stout setae on forefemur; (iv) shape of forefemur; (v) shape of median tubercles of abdominal terga.
This operational taxonomic unit is known by one specimen which distinctly differs from other representatives of the genus. We consider this material unacceptable for describing a new species now but provide a diagnosis.
1 larva, India, Tamil Nadu, Theni District, Kurangani Hills, Kottakudi River, 10.0809°N, 77.2552°E, 632 m a.s.l., 28.x.2020, Pandiarajan Srinivasan & Isack Rajasekaran leg., AMC ZN 243.
This OTU can be distinguished from other representatives of Teloganodes by the following combination of characters: (i) superlinguae laterally angular; (ii) forefemur without transversal row of stout setae; (iii) outer margin of femora with a regular row of long, stout hair-like setae only; (iv) dorsal surface of femora with submarginal irregular row of short stout setae; (v) tarsal claw with four median denticles and 1–2 subapical denticles (if two, on opposite sides of claw); (vi) median tubercles on terga III–X, which are indistinct on tergum III; wide on terga IV–VI; largest and elongate on terga VII–IX; elongate and thin on terga X; (vii) sublateral areas of posterior margins of terga IV–VI with bunches of 3–4 extremely long, pointed, stout setae; (viii) surface of cerci covered with long, stout, hair-like setae; setae in posterior margin of cerci segments less than half length of corresponding segment.
In this study, we used all available COI sequences of Dudgeodes species. Notably, 10 Dudgeodes OTUs, which have not yet been described morphologically, have already been sequenced and their independent position is certain (
All sequenced species of Dudgeodes, including undescribed OTUs, from continental Southeast Asia, excluding D. palnius, form a separate clade (Fig.
Genetic distances (COI) between sequenced Dudgeodes species of continental part of Southeast Asia, calculated using the Tamura-Nei (TN93) and Kimura 2-parameter (K2) models with a gamma distribution (G) (TN93+G/K2+G).
Dudgeodes romani | Dudgeodes sp. B (HQ581578) | Dudgeodes sp. A (HM417049) | Dudgeodes selvakumari sp. nov. | |
---|---|---|---|---|
Dudgeodes romani | 0.01/0.01 | |||
Dudgeodes sp. B (HQ581578) | 0.3226/0.3695 | — | ||
Dudgeodes sp. A (HM417049) | 0.0558/0.0545 | 0.3037/0.3585 | — | |
Dudgeodes selvakumari sp. nov. | 0.4616/0.5456 | 0.4642/0.5376 | 0.4310/0.5140 | 0.00/0.00 |
We are grateful to Andrii Khomenko (V.N. Karazin Kharkiv National University, Kharkiv, Ukraine) for his advice on molecular study, and to Céline Stoffel (Museum of Zoology, Lausanne, Switzerland) for her dedicated work with the genetic procedures. We are also grateful to the anonymous reviewers and editors, Robert Forsyth and Ben Price, who helped to improve the paper significantly.