Research Article |
Corresponding author: Alexander V. Martynov ( centroptilum@gmail.com ) Academic editor: Lyndall Pereira-da-Conceicoa
© 2021 Alexander V. Martynov, C. Selvakumar, Dmitry M. Palatov, K. A. Subramanian, K. G. Sivaramakrishnan, M. Vasanth, Luke M. Jacobus.
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Citation:
Martynov AV, Selvakumar C, Palatov DM, Subramanian KA, Sivaramakrishnan KG, Vasanth M, Jacobus LM (2021) Overview of Indian and Nepali representatives of the Cincticostella nigra (Uéno, 1928) complex (Ephemeroptera, Ephemerellidae), with discussion about Cincticostella Allen, 1971 species complexes. ZooKeys 1040: 123-166. https://doi.org/10.3897/zookeys.1040.64280
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The concept of the Cincticostella nigra (Uéno, 1928) (Ephemeroptera: Ephemerellidae) complex is clarified and Indian and Nepali representatives of the complex are reviewed. Four new species are described viz. Cincticostella changfai Martynov & Palatov, sp. nov., Cincticostella funki Martynov, Selvakumar, Palatov & Vasanth, sp. nov., Cincticostella shinichii Martynov & Palatov, sp. nov. and Cincticostella wangi Selvakumar, Martynov & Subramanian, sp. nov. The larva of C. corpulenta (Braasch, 1981) is re-described, based on the holotype and paratypes. Cincticostella gosei (Allen, 1975) is recorded from India for the first time. Morphological differences of the species complexes of Cincticostella Allen, 1971 are analysed. A new species complex, Cincticostella gosei complex, is proposed. Summaries of distribution data and habitat preferences, as well as new larval diagnoses, are presented for all species of the Cincticostella nigra complex.
China, Ephemerellinae, Ephemerelloidea, Indomalayan Region, new records, new species, Pannota
This article is a further contribution to a series of papers (
Due to uncertain relationships of species in the genus Cincticostella and possible polyphyly, we use the term “complex” to indicate distinctly different groups of species – the C. insolta complex (see
The C. nigra complex, as well as the family Ephemerellidae in general, has been relatively poorly investigated in India and Nepal. Only one species of the C. nigra complex has been reported from these countries: C. corpulenta described from Nepal by
Materials from India and Nepal were used in this research. New larval material was collected by kick-net sampling and hand-picking. All of this material is stored in 80–95% ethanol. Some specimens were mounted on slides with Canada balsam or Hoyer’s medium.
The holotype of C. corpulenta from the Stuttgart State Museum of Natural History [SSMNH] (Stuttgart, Germany) was also examined, along with two paratypes from the Purdue University Entomological Research Collection [
Photographs of specimens and their body parts were taken using a Leica M205A microscope, Zeiss Stemi 2000 binocular with Canon Power Shot A 640 and Ulab XY-B2T microscope with Canon Power Shot A 630. Some figures were subsequently improved with Adobe Photoshop CS5 and Helicon Focus 6.
Our hypotheses of species are based on morphological species concepts. Two middle instar larvae from a single morphotype were collected in Nepal in 2014. These larvae belong to the C. nigra complex and slightly differ from all known species. However, the poor material and absence of late larval instars do not allow us to describe a new species adequately and with confidence. Therefore, we give a provisional species designation only, along with diagnostic features.
The analysis of original material, historical collections and literature data showed that there are seven species belonging to C. nigra and C. gosei complexes (see details below) within India and Nepal: C. corpulenta (Braasch, 1981), C. gosei (Allen, 1975), four new species (C. changfai Martynov & Palatov, sp. nov., C. shinichii Martynov & Palatov, sp. nov., C. wangi Selvakumar, Martynov & Subramanian, sp. nov. and C. funki Martynov, Selvakumar, Palatov & Vasanth, sp. nov.) and one provisional species that we designate as Cincticostella sp. A. The taxonomic status of C. indica (Kapur & Kripalani, 1961) is discussed below. In addition, we report the first record of C. gosei for India.
Diagnoses of species provide characters that distinguish them within their corresponding complexes.
The following nominal species are treated in alphabetical order, with the provisional species listed at the end.
Larva. Late instars: body length 10.7–14.0 mm; caudal filaments length 6.0–9.1 mm. Body robust, yellowish-brown to brown (Fig.
Head: Without tubercles and ridges. Genae moderately developed, rounded (Fig.
Thorax: Dorsal surface of thorax covered with few scattered short, hair-like setae. Pronotum expanded laterally, with broad, rounded, anterolateral projections (Fig.
In late instars, femora of all legs slightly flattened (length/width ratio = fore femur 2.17–2.38; middle femur 2.56–2.86; hind femur 2.63–2.86), each one with longitudinal ridge, especially visible on middle and hind femora (Fig.
Whole dorsal surface of fore femur with scattered middle-sized, hair-like setae; basal part of surface with group of spine-like setae, bifurcated stout setae and stout hair-like setae. Distal part of dorsal surface with narrow, transverse band consisting of irregular, sparse row of mainly long and middle-sized, pointed, bifurcated and only sometimes rounded apically, stout setae (some of them situated on chalazae) (Fig.
Dorsal surfaces of middle and hind femora covered with few hair-like setae and numerous short and several middle-sized rounded or bluntly pointed (sometimes bifurcated) apically stout setae (Fig.
Inner margins of middle and hind tibiae and tarsi densely covered with spine-like setae and stout, hair-like setae; inner margins of middle and hind tibiae also with long, pointed and bifurcated, stout setae. Outer margins of middle and hind tibiae with rows of long, pointed and bifurcated, stout setae. Outer margins of middle and hind tarsi with hair-like setae (solitary and in tufts) only.
Ventral surfaces of all tibiae and tarsi with hair-like setae (solitary and in tufts) and long, thin, pointed, stout setae; stout setae situated in apical parts of tibiae and tarsi and along their inner margins.
Tarsal claws of all legs hooked, usually with two subequal denticles (seldom with one denticle) (Fig.
Abdomen: Central parts of dorsal surfaces of terga II–IX with two medial fields of middle-sized and small stout setae (Fig.
Posterior margins of terga I–II each with row of long hair-like setae; of tergum III with few long thin bluntly pointed or bifurcated apically stout setae and hair-like setae; of terga IV–VII with several rounded and pointed (sometimes bifurcated) stout setae near paired projections; of terga VIII–X with rows of short and middle-sized, mainly rounded apically, stout setae (these rows extending from paired projections to lateral margins). Lateral margins of terga IV–VIII covered with numerous short stout setae.
Pairs of pointed, not bifurcated projections present on abdominal terga II–X, with those on terga II–IV and X smaller than others; those on terga V–IX strongest (Fig.
Dorsal surfaces of abdominal gills covered with hair-like setae and scale sockets; shapes of gills as in Fig.
Caudal filaments subequal in length, with elongated, bluntly pointed or rounded (sometimes bifurcated) apically stout setae and hair-like setae at articulations (Fig.
Unknown.
The new species is named in honour of Dr. Chang-Fa Zhou (Nanjing Normal University, China), who contributed significantly to the study of the genus Cincticostella.
The new species can be easily distinguished from other representatives of the C. nigra complex by the following combination of characters: (i) genae moderately developed, rounded (Fig.
Known only from Nepal and northern India.
Larvae of this species were collected in middle-sized rivers (wide 4–15 m) in deep valleys, at an altitude of about 1000 m a.s.l. on the South slope of the Great Himalaya Range (India, Uttarakhand State and Bagmati Zone, Central Nepal) (Fig.
Holotype: Nepal: larva, Bagmati zone, Shivapuri Nagarjun National Park, Gohare Khola River (near Mahankal village), 27.885842°N, 85.531386°E, h ~ 1050 m a.s.l., 4.iii.2007, Chertoprud M.V. leg. – IN Nepa5Cinsp1/1 [NMNH
Ephemerella (Drunella) corpulenta Braasch, 1981
Cincticostella corpulenta
(Braasch, 1981) in
A supplemental narrative description is provided, based primarily on the study of a slide of the holotype (larva, male) and two paratypes in ethanol. Some characters are given simply as in the original description (
Larva (male). Body brown according to
Dorsal surface of head covered with numerous very short scales, empty scale sockets and short, hair-like setae. Mouthparts (Fig.
Mandibles (Fig.
Thorax: Dorsal surface of thorax covered with scattered short, waved and hooked, stout hair-like setae, thin and stout, hair-like setae and scale sockets (Fig.
Legs slightly flattened (length/width ratio = fore femur 2.08; hind femur 2.56), each one with longitudinal ridge (Fig.
Tarsal claws of all legs hooked, with one large denticle and several subapical setae (Fig.
Abdomen: Central part of dorsal surface of terga II–IX with two medial fields of mainly short stout setae with rounded or bluntly pointed, sometimes bifurcated, apices (Fig.
Pairs of not bifurcated projections present on abdominal terga II–IX, relatively weakly developed (Fig.
Abdominal gills dorsal surfaces covered with hair-like setae and scale sockets; gill III oval, with somewhat extended posteromedial angles without medial, transverse band of weakened membrane. Due to slide-mounting of holotype, gills deformed and not separated from abdomen.
Caudal filaments subequal in length; middle parts with elongated stout setae with bluntly pointed or bifurcated apices (Fig.
This species can be distinguished from all other species of the C. nigra complex by the following combination of characters: (i) genae poorly developed, head oval dorsally (Fig.
Nepal (
Adult stages unknown. The holotype (on slide) has the middle legs and one hind leg missing.
No data, but assumed to be cold water rivers and streams, based on what we know about the Trisuli River near Dhunche.
Holotype: Nepal: larva on slide, Himalaya, Trisuli Khola vor Dhunche, 1950 m a.s.l. NN, 30.04.1978, Leg. I. Sivec [SSMNH]. Paratypes: two larvae, same data as holotype [
Larva. Late instars: body length 10.5–11 mm; caudal filaments length 6–7 mm. Body robust, yellowish-brown, abdomen darker (Fig.
Head: Without tubercles and ridges. Genae rounded, moderately developed (Fig.
Thorax: Dorsal surface of thorax covered with scattered mainly short, waved and hooked, stout hair-like setae (Fig.
All femora slightly flattened (length/width ratio = fore femur 2.0–2.1; middle femur 2.0–2.2; hind femur 2.0–2.2) (Fig.
Whole dorsal surface of fore femur covered with scattered middle-sized, hair-like setae and scale sockets with small scales in some of them; basal part of surface with group of spine-like setae, bifurcated, stout setae and stout, hair-like setae. Additionally, numerous long, bifurcated apically, stout setae covering dorsal surface of fore femora along basal half of its inner margin. Distal part of surface with relatively wide, transverse, band consisting of irregular, sparse rows of different-sized (mainly long), rounded apically (only sometimes bluntly pointed with shallow bifurcation of apex), stout setae; some of setae situated on chalazae (Fig.
Dorsal surface of middle and hind femora covered with few scattered hair-like setae and scale sockets with small scales in some of them; also, surface covered with numerous short, rounded apically, stout setae (Fig.
Inner margins of middle and hind tibiae and tarsi densely covered with spine-like setae and stout, hair-like setae; inner margins of middle and hind tibiae also with long, pointed and stout setae. Outer margin of middle and hind tibiae with irregular row of long, pointed and bifurcated, stout setae. Outer margin of middle and hind tarsi with hair-like setae (solitary and in tufts) only.
Ventral surfaces of all tibiae and tarsi with hair-like setae (solitary and in tufts) and long, thin, pointed, stout setae; stout setae situated in apical parts of tibiae and tarsi and along their inner margins.
Tarsal claw of all legs hooked, with two (rarely three) denticles distanced from each other; basal denticle (rarely two denticles) distinctly larger; distal denticle directed angled forward (Fig.
Abdomen: Posterior margins of tergum I with thin and stout, hair-like setae only; several similar setae present on surface of tergum I. Submedian areas of terga II–IX surfaces, posterior margins of terga VIII–X (excluding central area between submedian projections) and all paired submedian projections covered with small and middle-sized, oval or, sometimes, with slightly divergent margins, stout setae with rounded apices. Additionally, all terga surfaces covered with scattered, small, stout hair-like setae, short, thin, hair-like setae, scale sockets and short scales in some of them. Sterna covered with scattered thin and stout, hair-like setae and scale sockets.
Pairs of pointed, not bifurcated, projections present on abdominal terga II–IX; those on terga V–VIII strongest; those on terga II–IV distinctly smaller than others (Fig.
Dorsal surface of abdominal gills covered with scattered hair-like setae and scale sockets; shape of gills as in Fig.
Caudal filaments subequal in length, with mainly elongated, rounded apically, stout setae and hair-like setae at articulations.
Unknown.
The new species is named in honour of Dr. David Funk (Stroud Water Research Center, USA), who contributed significantly to the study of Ephemerellidae.
The species is morphologically close to C. shinichii sp. nov. (see below), but can be distinguished from this species and other representatives of the complex by the following combination of the characters: (i) genae rounded and moderately developed (Fig.
Northern India and the India-China border region (Uttarakhand State and Arunachal Pradesh).
In Arunachal Pradesh, larvae of C. funki sp. nov. were collected from the Rike River (type locality) (2–5 m wide) at intermediate high mountain areas, on the Eastern Himalayan Range (Fig.
In Uttarakhand State, larvae of C. funki sp. nov. were collected from a small river (2–4 m wide) in medium high mountains, on the southern slope of the Great Himalayan Range (Fig.
Holotype: India: larva, Arunachal Pradesh, Papumpare District, vicinity of Parang Village, Rike River, 27.32797°N, 93.50308°E, h ~1285 m a.s.l., 14.xii.2018, Coll. Bikramjit Sinha – IN
Ephemerella (Cincticostella) gosei Allen, 1975
Serratella thailandensis Allen, 1980 (junior objective synonym, Edmunds and Murvosh 1995)
This species can be distinguished from other Cincticostella species by the following combination of characters: (i) head brown, with three white to yellow spots near ocelli (spots near lateral ocelli the largest) (Fig.
Thailand (
The larva of this species was properly described from Thailand by
Cold fast-flowing river with cobbles and gravel. The Ranga River habitat is shown in
India: 17 larvae, Arunachal Pradesh, Lower Subansiri District, Ranga River, 27.396404°N, 93.757378°E, h ~ 625 m a.s.l., 06.xi.2015, Coll. Bikramjit Sinha – IN 5346/H13 [
Ephemerella indica Kapur & Kripalani, 1961
Cincticostella indica
(Kapur & Kripalani, 1961) in
Cincticostella indica was described only from the female adult stage (
Larva. Middle and late instars: body length of mature larva 12.0 mm, caudal filaments length 7.8 mm. Body yellowish-brown to brown, robust, covered with scale sockets and small scales in some of them.
Head: Without tubercles and ridges. Genae moderately developed, rounded (Fig.
Thorax: Dorsal surface of thorax covered with numerous small stout setae with divergent margins, rounded or bifurcated apices and less numerous short, strait, thin and stout, hair-like setae and scale sockets (Fig.
In late instars, femora of all legs slightly flattened (length/width ratio = fore femur 2.00–2.17; middle femur 2.17–2.33; hind femur 2.33–2.44) and bearing longitudinal ridge, especially visible on middle and hind femora (Fig.
Larva of Cincticostella shinichii Martynov & Palatov, sp. nov., paratype (A–C, E, F, H) and holotype (D, G) A fore femur B middle femur C hind femur D, E stout setae of dorsal surface of fore femur F stout setae of outer margin of hind femur G stout setae of outer margin of middle tibia H tarsal claw.
Dorsal surface of fore femur with several elongated, pointed and bifurcated, stout setae with feathered margins near basal margin; also apical half of dorsal surface of fore femur with transverse, relatively wide and dense band of mainly middle-sized and short, bifurcated, bluntly pointed or rounded apically, stout setae (some situated on chalazae) (Fig.
Dorsal surface of middle and hind femora covered with numerous mainly middle-sized and short, rounded or bifurcated apically, stout setae (Fig.
Outer margins of middle and hind tibiae with regular rows of long, pointed and bifurcated, stout setae and few hair-like setae (solitary and in tufts) amongst them (Fig.
Tarsal claws of all legs hooked, with one large denticle and several subapical setae (Fig.
Abdomen: Central part of dorsal surface of terga II–X with two medial fields of middle-sized and small, stout setae with bifurcated, bluntly pointed or rounded apices (Fig.
Larva of Cincticostella shinichii Martynov & Palatov, sp. nov., holotype (A and I) and paratype (B–H) A abdomen, dorsal view B tergum X and posterior margin of tergum IX (black arrow shows posterior margin of tergum X lack of paired submedian projections) C–G gills III–VII H caudal filament, middle part I caudal filament, proximal part.
Posterior margins of tergum I with row of long, hair-like setae; of terga II–III (especially tergum II) with row of long, thin, bluntly pointed or rounded apically, stout setae and hair-like setae; of terga IV–VII with several elongated rounded or bifurcated apically stout setae near paired projections; of terga VIII–X with rows of elongated or short, stout setae with rounded or bifurcated apices (these rows extending from paired projections to lateral margins). Lateral margins of terga IV–VIII covered with numerous rounded or bifurcated apically stout setae. Posterolateral projections of abdominal segments IV–IX present (Fig.
Pairs of pointed, not bifurcated projections present on abdominal terga II–IX (Fig.
Dorsal surfaces of abdominal gills covered with hair-like setae (mainly in apical part) and scale sockets; shape of gills as in Fig.
Caudal filaments subequal in length, with elongated stout setae with bluntly pointed, bifurcated or pointed apices and hair-like setae at articulations (Fig.
Unknown.
The new species is named in honour of Dr. Shin-ichi Ishiwata (Kanagawa Environmental Research Center, Japan), who contributed significantly to the study of Cincticostella species.
This new species is close to C. corpulenta, but can be differentiated from this and other species of the complex by the following combination of characters: (i) genae moderately developed, rounded (Fig.
Known only from Nepal.
Larvae of C. shinichii sp. nov. were collected from stones with algal fouling in sections with current velocity of about 0.3–0.8 m/s of a large river (10–17 m wide) within low mountains on the southern slope of the Great Himalayan Range (Bagmati Zone, Central Nepal). Investigated rivers are under the significant anthropogenic pressure and can be classified as alpha- or beta-mezosaprobic waterbodies. Apparently, larvae inhabit rhithral zones of waterbodies with relatively high average current velocity and the predominance of large stones at the bottom. Larvae of Stenopsyche, Glossosomatidae, Hydropsychidae, Baetis (s.str.) sp. and representatives of other Ephemerellidae genera were collected along with the new species.
Holotype: Nepal: larva (on slide 647), Bagmati Zone, Shivapuri Nagarjun National Park, Melamchi River (1 km below Talamarang village), 27.844497°N, 85.557433°E, h ~ 900 m a.s.l., 4.iii.2007, Chertoprud M.V. leg. – IN Nepa7Cinsp [NMNH
Larva. Body length 6.0–7.7 mm. Caudal filaments length 5.0–5.5 mm. Body colour yellowish-brown (Fig.
Head: Ocelli, clypeus dorsal surface and vertex without tubercles and ridges (Fig.
Mandibles (Fig.
Thorax: Pronotum expended laterally, with anterolateral angles small and projecting forward (Fig.
Femora of all legs flattened (length/width ratio = fore femur 2.0–2.1; middle femur 2.0–2.1; hind femur 2.0–2.2), with longitudinal ridges (Fig.
Fore femur with distinct transverse band of numerous, mainly extremely long to middle-sized, stout setae with deeply bifurcated apices (some situated on chalazae) (Fig.
Dorsal surfaces of middle and hind femora almost completely covered with different-sized, bifurcated, stout setae, some very long (Fig.
Dorsal surface of all tibiae with longitudinal rows of long, bifurcated stout setae; in middle and hind legs, these rows situated closer to inner margins of tibiae. Outer margin of fore tibia with thin hair-like setae only; in middle and hind tibiae also with regular rows of long, bifurcated, stout setae.
Claws of all legs with one denticle each and several subapical setae (Fig.
Abdomen: Paired, pointed projections present on abdominal terga II–VIII; posterior margin of tergum VIII almost straight, its protuberances very small and smooth; posterior margin of terga IX and X without projections (Fig.
Gill III without medial transverse band of weakened membrane, dorsal lobe of gills III–V similar in shape (Fig.
Caudal filaments subequal in length, with pointed, stout setae and hair-like setae on posterior edge of each segment, setae shorter than corresponding segment.
Setae on dorsal surface of thorax of some representatives of Cincticostella nigra (Uéno, 1928) complex A dominant kind of stout setae – numerous small stout setae with divergent margins, bifurcated apices B dominant kind of stout setae – short, waved and hooked, stout hair-like setae C dominant kind of stout setae – scattered short, relatively strait, stout hair-like setae.
Unknown.
The new species is named in honour of Dr. T.-Q. Wang (formerly Purdue University, USA), who contributed significantly to the study of Ephemerelloidea.
Habitats of Cincticostella changfai Martynov & Palatov, sp. nov. (A) and Cincticostella funki Martynov, Selvakumar, Palatov & Vasanth sp. nov. (B and C). A Ramganga River (300 m above Patangoari Village), Uttarakhand State, Almora District, India B Rike River (vicinity of Parang Village), Arunachal Pradesh, Papumpare District, India C second order left tributary of the Ramganga River (10.1 km north-eastwards of Chaukhutia Town), Uttarakhand State, Almora District, India.
This species can be distinguished from other representatives of C. nigra complex by the following combination of characters: (i) genae moderately developed, rounded; (ii) labrum with deep and wide anteromedian emargination (Fig.
India-China border region.
Cold fast-flowing stream with cobbles and gravel. The type locality of C. wangi sp. nov. is shown in
Holotype: India: larva, Arunachal Pradesh, Lower Subansiri District, Tale Valley, unnamed stream, 27.537201°N, 93.959883°E, h ~ 2370 m a.s.l., 14.iv.2015, Coll. K.A. Subramanian – IN 5575/H13 [
Immature larva: (i) genae moderately developed, rounded; (ii) anteromedian emargination of labrum shallow (labrum height in emargination/maximum labrum height ratio – 0.85); (iii) maxillary palp well-developed; (iv) segment III of maxillary palp, thin, elongated, rounded apically; (v) group of 13–16 setae different-sized, stout, hair-like situated on surface of galea-lacinia near base; (vi) prothoracic anterolateral projections small; (vii) mesothoracic anterolateral projections well-developed, subparallel to lateral aspect of body, not notched; (viii) surface of thorax covered with scattered short, thin, hair-like setae and few scale sockets and very short rounded scales with feathered margins in some of them; few waved and hooked setae cover mainly wing pads; (ix) dorsal surface of fore femur with sparse, transverse row of mainly long, stout setae with slightly convergent margins and bifurcated apices; (x) tarsal claw with one large denticle and several subapical setae; (xi) pairs of pointed projections present on abdominal terga II–IX; those on terga V–VIII strongest; those on tergum IX moderately developed, rounded apically.
Nepal.
Two middle larval instars (body length 5.5–6.5 mm; caudal filaments length 5.2–5.3 mm) of this operational taxonomic unit (OTU) were collected in Nepal in 2014. However, the poor material and absence of late larval instars do not allow us to describe a new species adequately. We hope that this account will facilitate discovery of additional material of this OTU.
Larvae of Cincticostella sp. A inhabit cold, moderate flowing springs with stones, gravel and clumps of moss. Larvae were collected from such a spring (up 1.3 m wide) in medium high mountains, on the southern slope of the Great Himalayan Range. This was a small, lotic waterbody with relatively low water temperature (10 °C in sampling period), average current velocity (0.2–0.6 m/s), mosaic bottom and a low degree of anthropogenic pressure.
Nepal: 2 larvae (one on slide 634), Gandaki Zone, Kaski District, stream at the Tolka-Lambruk Road, 28.365000°N, 83.831667°E, h ~ 2000 m a.s.l., 27.i.2014, Chertoprud M.V., Marinskiy V.V. leg. – IN Nepa1Cinsp [NMNH
To date, nine species of the genus Cincticostella are known from India: C. bifurcata Xie, Jia, Chen, Jacobus & Zhou, 2009; C. braaschi Jacobus & McCafferty, 2008; C. changfai Martynov & Palatov, sp. nov.; C. funki Martynov, Selvakumar, Palatov & Vasanth sp. nov.; C. gosei (Allen, 1975); C. insolta (Allen, 1971); C. ranga Selvakumar & Subramanian, 2019; C. richardi Martynov & Palatov, 2019 and C. wangi Selvakumar, Martynov & Subramanian, sp. nov. (
Overall, only three of all known species of Cincticostella occur at latitudes that are more southern than the Himalayan Region, namely C. gosei, C. insolta and C. femorata. The southernmost records of C. insolta are from Thailand, where the species inhabits coldwater flows in northern upland region of the country (
Cincticostella comprises at least 21 valid species (Table
In light of the several new species described since
Based on our research, we conclude that the establishment of the discussed complexes was for expediency. In fact, the C. insolta and C. nigra complexes are more distanced morphologically from each other than previously thought (see
Characters | C. insolta complex | C. nigra complex | C. gosei complex | |
---|---|---|---|---|
1 | Two pairs of suboccipital tubercles | present | absent | absent |
2 | Serration of margins of middle and hind femora | present | absent | absent* |
3 | Presence of numerous large, rounded, scale sockets on body surface | present | absent | present |
4 | Rate of anterolateral emargination of labrum | shallow | from shallow to deep | moderate |
5 | Maxillary palp | reduced, articulations of segments not distinct, especially between segments I and II | mainly well-developed, articulations of all segments distinct** | absent |
6 | Segments I and II of labial palp | wide | wide | relatively narrow, elongated |
7 | Stout setae on outer margin of fore femur | several stout setae only | numerous stout setae | several stout setae only |
8 | Stout setae on dorsal surface of middle and hind femora | absent or up to several stout setae in basal area | surface with numerous stout setae | absent |
9 | Shape of hind femur | strongly or moderately widened | moderately widened | moderately widened |
10 | Stout setae on dorsal surface of abdominal terga and paired projections | absent | present | absent |
The following morphological features place C. gosei closer to the C. insolta complex than to the C. nigra complex: presence of numerous large, rounded, scale sockets on body surface, absence of maxillary palp (this character state may prove to be related to significant reduction of the maxillary palp, including reduction of segments’ articulations), number of stout setae on the outer margin of the fore femur and absence of numerous stout setae on dorsal surfaces of the middle and hind femora, complete absence of stout setae on dorsal surfaces of abdominal terga and paired projections.
C. insolta complex | C. gosei complex | C. nigra complex |
---|---|---|
C. bifurcata Xie, Jia, Chen, Jacobus & Zhou, 2009 | C. gosei (Allen, 1975) | C. changfai Martynov & Palatov, sp. nov. |
C. braaschi Jacobus & McCafferty, 2008 | C. colossa Kang & Yang, 1995 | |
C. femorata (Tshernova, 1972) | C. corpulenta (Braasch, 1981) | |
C. insolta (Allen, 1971) | C. elongatula (McLachlan, 1875) | |
C. ranga Selvakumar & Subramanian, 2019 | C. funki Martynov, Selvakumar, Palatov & Vasanth sp. nov. | |
C. richardi Martynov & Palatov, 2019 | C. fusca Kang & Yang, 1995 | |
C. sivaramakrishnani Martynov & Palatov, 2019 | C. levanidovae (Tshernova, 1952) | |
C. tornata Auychinda & Gattolliat, 2020 | C. nigra (Uéno, 1928) | |
C. orientalis (Tshernova, 1952) | ||
C. shinichii Martynov & Palatov, sp. nov. | ||
C. szechuanensis Xie, Jia, Chen, Jacobus & Zhou, 2009 | ||
C. wangi Selvakumar, Martynov & Subramanian, sp. nov. | ||
Cincticostella sp. A |
The authors are grateful to Dr. Mikhail V. Chertoprud (Lomonosov Moscow State University, Russia) for donation of larval materials from his collection; to Dr. Arnold H. Staniczek (Stuttgart State Museum of Natural History, Stuttgart, Germany) for the opportunity to investigate the holotype of C. corpulenta; to Dr. Jean-Luc Gattolliat (University of Lausanne and Museum of Zoology, Lausanne, Switzerland) and anonymous reviewer whose comments and remarks helped to improve the article. Some equipment used by AVM in this investigation was provided by the IDEA WILD organisation in 2016 and 2018. CS, KAS and MV thank Director,