Research Article |
Corresponding author: Nikita J. Kluge ( n.kluge@spbu.ru ) Academic editor: Lyndall Pereira-da-Conceicoa
© 2020 Nikita J. Kluge, Roman J. Godunko, Marek Svitok.
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:
Kluge NJ, Godunko RJ, Svitok M (2020) Nomenclatural changes in Centroptella Braasch & Soldán, 1980 (Ephemeroptera, Baetidae). ZooKeys 914: 81-125. https://doi.org/10.3897/zookeys.914.46652
|
The genus Centroptella Braasch & Soldán, 1980 is accepted here in a wide sense, i.e., including Chopralla Waltz & McCafferty, 1987. This genus concept is similar to the concept of the genus Bungona Harker, 1957 proposed by
India, mayflies, new species, South Africa, systematics, Thailand, Vietnam
Initially, the genus Centroptella Braasch & Soldán, 1980 was established for a single species, C. longisetosa Braasch & Soldán, 1980 described from China. Subsequently, other species of Centroptella were described from the Oriental Region, i.e., C. ceylonensis Müller-Liebenau, 1983, C. similis Müller-Liebenau, 1983 and C. soldani Müller-Liebenau, 1983 from Sri Lanka, C. pusilla Müller-Liebenau, 1984 from Borneo, C. liebenauae Soldán, Braasch & Muu, 1987 and C. colorata Soldán, Braasch & Muu, 1987 from Vietnam.
Imagines were reared from larvae in cages placed in natural flowing water and in containers with stagnant water. Part of material, including the holotype of Centroptella ingridae sp. nov., will be permanently deposited in the Russian Academy of Sciences, Zoological Institute, Zoological Museum (Saint Petersburg, Russia) (
The term “microlepide” is used according to
For scanning electron microscopy (Figs
Other samples (Figs
Originally, the genus Bungona was established for a single species, Bungona narilla Harker, 1957, which was described from Coal and Candle Creek, Ku-ring-gai Chase National Park, Sydney (Australia). This species description was based on one male imago (holotype), one female subimago and one larva. The reason these three specimens were placed in one species was not reported. The description contains evident errors (tarsi of middle and hind legs were regarded to be 5-segmented, gonostyli were regarded to be 4-segmented, paraprocts were confused with the penis); the combination of other characters is different from any known species. The holotype and paratypes of B. narilla were stated to be housed in the British Museum (Natural History) (
1) “A neotype is validly designated when there is an exceptional need and only when that need is stated expressly and when the designation is published with the following particulars: 75.3.1. a statement that it is designated with the express purpose of clarifying the taxonomic status or the type locality of a nominal taxon ...”. All species taken into account in the publication, where the neotype of B. narilla was designated, i.e., narilla [Bungona], fustipalpus [Cloeodes] and illiesi [Cloeodes], were regarded as belonging to one species, and all their characters hitherto regarded as species-specific, were regarded as individual variability. In this situation, neotype designation is unnecessary, because it does not serve to clarify the taxonomic status of any nominal taxon.
2) “A neotype is validly designated when ... the designation is published with the following particulars: 75.3.5. evidence that the neotype is consistent with what is known of the former name-bearing type from the original description and from other sources”. There are no sources of knowledge about the holotype of B. narilla other than its original description, so the neotype can be compared only with the description given by
In the holotype description, coloration is characterized as follows: “First two abdominal segments light brown, segments 3–7 yellow, the posterior segments light brown”; in the neotype description—coloration is characterized as follows: “abdominal segments 1–2 with central cream marking, 3 dark brown, 4 cream, 5–6 dark brown, 7–10 light brown”.
In the holotype description, hind leg proportions are characterized as follows: “tibia and tarsus equal in length, being about three-quarters length of femur. Tarsal segments of hind leg in decreasing order of length: 2, 3, 5, 4, 1 (fused with tibia)”; in the neotype description, hind leg proportions are characterized as follows: 1.00 : 0.74 : 0.09 : 0.18 : 0.10 : 0.08 : 0.15. That means, that the neotype has a femur/tibia/tarsus ratio of 1 : 0.75 : 0.6 (i.e., tibia and tarsus are not equal in length), and its tarsal segments in decreasing order of length are 2, 5, 3, 4, 1. The meaning of these numbers is unclear, because hind legs of all Baetidae have only 4 tarsal segments (including the first one, which is immovably fused with the tibia); but in any case, in the holotype the pen-penultimate segment is longer than the claw-bearing segment, while the neotype has the pen-penultimate segment shorter than the claw-bearing segment.
The drawing of gonostyli included with the holotype description (
3) “A neotype is validly designated when ... the designation is published with the following particulars: 75.3.6. evidence that the neotype came as nearly as practicable from the original type locality”. In the publication where this neotype was designated (
Later,
The possibility to designate a new neotype after respective request to the International Commission of Zoological Nomenclature (according to Article 75.5 of the ICZN) can be a reasonable step for the rectification of this situation and taxonomic stability within the genus Bungona. Nevertheless, such a step should be taken only when new material of reared imaginal and larval specimens (preferably close to the type locality) is available. Despite considerable effort, such material is not available yet. Consequently, usage of the generic name Bungona is questionable and as such does not meet the requirements of the Article 23.9.1 of the ICZN.
The Australian Baetidae remain poorly known, with only 20 species described to date.
Given the inadequate nature of the original description, the loss of the type material, the improper assignment of a neotype, and the poorly documented diversity of related species in Australia, Bungona narilla (the type species of the genus Bungona) should be regarded as a nomen dubium. It then follows that the senior generic name for the species described below should be Centroptella.
= Chopralla Waltz & McCafferty, 1987: 182, syn. nov.
= Crassolus Salles, Gattolliat & Sartori, 2016: 104, syn. nov.
Centroptella longisetosa Braasch & Soldán, 1980.
Centroptella is characterized by an unusual combination of characters: on one hand, it undoubtedly belongs to the holophyletic taxon Baetovectata Kluge & Novikova, 2011, based on (1) presence of two marginal intercalaries in each space of wing (Fig.
Besides this paradoxical combination of baetovectatan and protopatellatan characters, Centroptella has an evident autapomorphy: secondary swimming setae on the outer sides of the larval cerci in the distal part of the cercus have oval transverse bases and form a regular row (Figs
Another peculiar character of Centroptella is the presence of a pair of spaced transverse rows of long bifurcate setae on certain abdominal sterna of the larva (Figs
The genus Crassolus Salles, Gattolliat & Sartori, 2016 was established for a single South African species Crassolus inzingae (Crass, 1947), which was originally described in the genus Pseudocloeon (
Examination of reared material of saxophilum [Pseudocloeon] collected in the Western Cape Province in 2019 (Figs
Character “3” (distance between prostheca and incisors of right mandible) was said to have increased from 0.00 (ancestral condition reported for Crassolus inzingae) to 0.04 (Node 72). Actually, according to the matrix of characters (Appendix S3), among the species attributed to “Bungona”, this characters varies from 0.00 to 0.26. The condition “0.00” was reported for the larvae determined as “Bungona (Chopralla) liebenauae” and actually belongs to the new species Centroptella ingridae sp. nov. described below (Fig.
Character “9” (length of fore femur / distance between base of fore femur and base of most distal setae of fore femur) was said to have increased from 0.92 (ancestral condition reported for Crassolus inzingae) to 0.95 (Node 72). Actually, according to the matrix of characters, among the species attributed to “Bungona”, this characters varies from 0.92 (in three species included in the matrix) to 1.00.
Character “40” (slender process on prostheca of right mandible) was said to have changed from “0=absent” (ancestral condition reported for Crassolus inzingae) to “1=present” (Node 72). Actually, according to the matrix of characters, this process is absent in the larvae determined as “Bungona (Chopralla) liebenauae” and actually belongs to the new species Centroptella ingridae sp. nov. described below (Fig.
Characters “42” and “48” are “setae between prostheca and mola of right mandible” and “long setae between prostheca and mola of left mandible”. The both characters were said to have changed from “1=present” (ancestral condition) to “0=absent” (in the Node 72). In the matrix of characters (
The monotypic genus Crassolus was said to be characterized by six apomorphies under the numbers 0, 2, 5, 7, 20 and 22 (
The type species of Crassolus is closely related to Centroptella ingridae sp. nov. described below; these species have many common characters, including peculiar halberd-like tips of gonovectes not found in other taxa (Figs
Based on the above, the following generic synonymy is suggested: Centroptella = Crassolus, syn. nov. Within the genus Centroptella, several Asian and African species, including C. inzingae and C. ingridae sp. nov., constitute a natural species group, characterized by the halberd-like tips of the gonovectes and other common characters in imaginal and larval structure.
Among them, the subgenus Chopralla was an artificial group, because one of its species belongs to the inzinagae-ingridae group, while another species of the inzinagae-ingridae group was placed in a separate genus Crassolus (see above).
Two other subgenera, Centroptella and “Bungona” had not been separated one from another by any currently recognized characters.
According to the diagnosis of the subgenus Bungona, “Dorsal surface of labrum with two setae on anterolateral corner” (character “1”). In contrast, according to the original descriptions by
According to the diagnoses of the subgenera Bungona and Centroptella, they differ by the distance between the prostheca and mola of the right mandible (character “3”). However, the structure and position of the right prostheca is the same in the larva determined as “Bungona narilla” and in Centroptella soldani (
According to the diagnosis of the subgenus Centroptella, it has “few setae on outer margin of fore femur (around six)”, in contrast to 10 in Bungona (character “14”). However, according to original descriptions by
According to the diagnosis of the subgenus Bungona, it has “angle of row of long setae on posterior surface of fore tibia around 60°”, in contrast to 30° in Centroptella (character “13”). Actually this angle is around 30° both in the species attributed to Bungona (Fig.
Thus, the subgeneric classification proposed by
Considering the factors discussed above, the genus Centroptella should be accepted as comprising the following nominal species (alphabetically): Centroptella bifida (Shi & Tong, 2019) comb. nov.; Centroptella bintang (Marle, Salles & Gattolliat, 2016) comb. nov.; Centroptella ceylonensis Müller-Liebenau, 1983; Centroptella colorata Soldán, Braasch & Muu, 1987; Centroptella fusina (Tong & Dudgeon, 2003) comb. nov.; Centroptella fustipalpus (Lugo-Ortiz & McCafferty, 1998) comb. nov.; Centroptella illiesi (Lugo-Ortiz & McCafferty, 1998) comb. nov.; Centroptella inzingae Crass, 1947 comb. nov.; Centroptella liebenauae Soldán, Braasch & Muu, 1987; Centroptella longisetosa Braasch & Soldán, 1980; Centroptella ovata (Shi & Tong, 2019) comb. nov.; Centroptella papilionodes (Marle, Salles & Gattolliat, 2016) comb. nov.; Centroptella pontica (Sroka, Godunko & Gattolliat) (in
Under the name “Centroptella liebenauae”,
(1) Mature male larva with labels: “VIETNAM, Vinn Phu Prov., Soui Bac – Tam Dao, 10–16.X.1985 T. Soldán”, “Centroptella liebenauae T. Soldán det. 1985” and “HOLOTYPE”;
(2) 46 larvae with labels: “VIETNAM, Vinn Phu Prov., Suoi Bac Stream, Tam-Dao, 10–16.10.1984 T. Soldán”, “Centroptella liebenauae T. Soldán det. 1985” and “PARATYPES”; many of these larvae are late instars, and some are ready to moult to subimago;
(3) tube with 3 male imagines (one without genitalia), 1 male subimago, 1 female imago, 1 male larval exuviae and 1 abdomen of female subimago extracted from mature larva, with labels: “VIETNAM, stream, Tam-Dao 60 km NW of Hanoi, 23–25.5.1982 T. Soldán”, “Centroptella, T. Soldán det. 1982” and “PARATYPE”; now larval exuviae, parts of one male imago and parts of male subimago are mounted on slides in Canadian balsam. The larval exuviae are designated as the neotype of Centroptella liebenauae (see below).
All specimens in tubes (1) and (2) belong to one and the same species, which is described below as C. ingridae sp. nov., and all specimens in tube (3) belong to a single, different species, which is C. longisetosa.
According to the original description, the holotype is a larva collected 23–25.V.1982 together with an additional 18 larvae and 5 reared winged insects (three male imagines, one female imago and one male subimago), while larvae collected 10–16.X.1984 and 17.X.1984 are paratypes. This means that the larva labelled as “holotype” was actually collected 10–16.X.1984 (not 16.X.1985), and is not the holotype, but a paratype. We speculate that the true holotype (i.e., the specimen designated as the holotype in the original publication) is mixed among the 18 other larvae collected on the same dates (23–25.V.1982) and now cannot be recognized among them.
Judging by the list of specimens examined in the original description, among the specimens contained in the tube (3), the male imago without genitalia is “paratype No. 1”, the single female imago is “paratype No. 2” and the single male subimago is “paratype No. 3”; the single set of male larval exuviae belongs to one of four males in this tube. The location of the 19 larvae from this series (including the true holotype) is unknown. The lost holotype was an intact larva; no structures were mounted on any slide, so its details were not examined, and the authors of the original description could not have known to which of the two species it belonged.
Sometimes larvae of different species of Centroptella can be collected at the same place (NJK; unpublished data). In the case of the larvae of the two species described under the name “Centroptella liebenauae”, they differ in size, shape and coloration; these differences are easily visible if they lie together, but such differences can be overlooked if they are examined separately. The fact that the authors of the original description did not notice these differences indicates that they never saw larvae of these two species side-by-side. Judging by the fact that all 47 larvae collected 10–16.X.1985 belong to C. ingridae sp. nov., and all five reared imagines and subimagines collected 23–25.V.1982 belong to C. longisetosa, we assume that all larvae collected 23–25.V.1982, including the lost holotype of C. liebenauae, also belong to C. longisetosa. It is well known, that even in unimpaired rivers composition of mayfly communities varies considerably over time (e.g.,
Complete set of last instar male larval exuviae (Figs
Based on this neotype designation, we propose a new synonym: Centroptella longisetosa = Centroptella liebenauae syn. nov.; another species, described under the name “Centroptella liebenauae”, is a new species, and it is described here under the name C. ingridae sp. nov. (see below).
According to the Article 75.3 of the International Code of Zoological Nomenclature, a neotype is validly designated when there is an exceptional need and only when that need is stated expressly and when the designation is published with the particulars listed in the paragraphs 75.3.1–75.3.7. In the present case, such exceptional need is present, because usage of the same name Centroptella liebenauae for two distant species causes confusion; all particulars required in the paragraphs 75.3.1–75.3.7 are published here as the following:
“75.3.1. a statement that it is designated with the express purpose of clarifying the taxonomic status ...”. This purpose is to choose, which of two different species originally described under the name Centroptella liebenauae, should bear this name.
“75.3.2. a statement of the characters that the author regards as differentiating from other taxa the nominal species-group taxon for which the neotype is designated ...” These characters are give below, in the discription of C. longisetosa.
“75.3.3. data and description sufficient to ensure recognition of the specimen designated”. These data are given above.
“75.3.4. the authors’ reasons for believing the name-bearing type specimen(s) (i.e., holotype, or lectotype, or all syntypes, or prior neotype) to be lost or destroyed, and the steps that had been taken to trace it or them”. The steps that had been taken to trace the holotype, are reported above; the lost holotype is an intact larva, whose individual features have never been reported or figured; even its sex is unknown. If this specimen is found in future, it will be impossible to prove that it is the holotype designated in the original publication, as it could be any other specimen among 19 lost larval specimens, which have one and the same geographical label.
“75.3.5. evidence that the neotype is consistent with what is known of the former name-bearing type from the original description and from other sources ...”. The original description contains characters and figures of two different species, C. longisetosa and C. ingridae sp. nov. Based on the original description, we know that the former name-bearing type was collected in spring 1982, and analyzing the collection we know that all specimens collected at that time belong to C. longisetosa. Designating the neotype from specimens collected in autumn 1985 and belonging to a species different from the holotype, would clearly violate this paragraph.
“75.3.6. evidence that the neotype came as nearly as practicable from the original type locality ...”. The neotype comes from the type locality and has the same date of collecting as the holotype.
“75.3.7. a statement that the neotype is, or immediately upon publication has become, the property of a recognized scientific or educational institution, cited by name, that maintains a research collection, with proper facilities for preserving name-bearing types, and that makes them accessible for study”. This institution is the Institute of Entomology (BC CAS) in České Budějovice, Czech Republic, where this specimen was deposited recently.
Besides these formal rules, the Code requires maintainance of prevailing usage of the taxa names which can be done only under plenary power of the Commission (paragraph 75.6). In this case, referring to prevailing usage is impossible, because there are only six publications where the species name liebenauae [Centroptella] has been mentioned (
Our choice of the neotype, being the single one is consistent with the Code. Moreover, the single one provides the both considered species with monosemantic valid names, because the non-monosemantic (equivocal) name C. liebenauae becomes invalid. The designation of a neotype and the new synonymy proposed here will stop further confusion caused by inaccurate descriptions of two different species under the one name, C. liebenauae.
The type species of the genus Centroptella, C. longisetosa, was originally described based on larvae from China. According to the original description (
One of us (RJG) examined the mayfly collection deposited in the Institute of Entomology (BC CAS) and could not find the slide with the holotype of C. longisetosa. Consultations with Tomáš Soldán also did not clarify the fate of the holotype. Instead, there is a tube with larva in alcohol with labels: “CHINA, Liu Chiu, Kuj Fon Shan Mt., stream, 11.12.1959, leg. Ivan Hrdý”, “Centroptella longisetosa T. Soldán det. 1980” and “HOLOTYPE”. Judging from the original description, this intact larva is not the holotype, but one of five paratypes deposited in the Soldán collection. Another paratype was reported by
Centroptella longisetosa Braasch & Soldán, 1980: 123 (larva)
Cloeodes longisetosus:
Bungona (Centroptella) longisetosa:
Centroptella liebenauae Soldán, Braasch & Muu, 1987: 342 (partim: ♂ & ♀ imagines, ♂ subimago; non larva), syn. nov.
Chopralla liebenauae:
Bungona (Centroptella) liebenauae:
Paratypes of Centroptella longisetosa (deposited in the Institute of Entomology, BC CAS, České Budějovice and Purdue University Entomological Research Collection, West Lafayette, Indiana, USA): mature female larva with labels: “CHINA, Liu Chiu, Kuj Fon Shan Mt., stream, 11.12.1959, leg. Ivan Hrdý”, “Centroptella longisetosa T. Soldán det. 1980”, “HOLOTYPE” (now parts of this specimen are mounted on 2 slides, eggs mounted for SEM; one middle larval leg of another specimen, in the same tube (now treated by alkali and mounted on separate slide); one larva, Peoples Republic of China, Liu Chui, Kuj Fon Shan River, 11-12-1959, I. Hrdy, PERC-0063355.
Neotype and paratypes of Centroptella liebenauae (deposited in the collection of the Institute of Entomology, BC CAS, České Budějovice): one tube containing: 3 male imagines (one without genitalia), 1 male subimago, 1 female imago, 1 male larval exuviae (neotype) and 1 abdomen of female subimago extracted from mature larva, with labels: “VIETNAM, stream, Tam-Dao 60 km NW of Hanoi, 23–25.5.1982 T. Soldán”, “Centroptella, T. Soldán det. 1982” and “PARATYPE”; now larval exuviae (neotype) and parts of male imago and male subimago (one of which possibly was reared from the neotype) are mounted on slides.
INDIA, Tamilnadu, Tirunelveli District, Courtallam, Chittar River near Peraruvi (= Main Falls), 3–7.II.2013, coll. N. Kluge & L. Sheyko: 3 L-S-I♂, 1 L-S-I♀, 1 S-I♀, 1 L/S♂, 1 L/S♀, 1 larva of penultimate instar.
Larva. Cuticular coloration. Head mostly brown (Fig.
Shape and setation. Frontal suture short, nearly semicircular (Fig.
All thoracic terga without protuberances. Metanotum with vestiges of hind protoptera (Fig.
Scales on abdominal terga and sterna numerous, short, semicircular, colourless and delicate (Figs
Tergalii apically pointed and sharply differentiated as follows: tergalius I lanceolate, slightly bent, widened at midlength, with apex stretched and narrowly pointed (Figs
Male genitalia (examined in Indian specimen): In last larval instar, developing subimaginal gonostyli folded under larval cuticle in peculiar pose, with 3rd segments bent medially-proximally (Fig.
Centroptella longisetosa. 40–46 paratype of C. longisetosa: (40, 41) left and right mandibles (42) apex of maxilla (ds1, ds2, ds3 dentisetae) (43) labrum (44) glossa and paraglossa (ventral view) (45) labium (at left ventral view, at right dorsal view; muscles shown by interrupted lines) (46) maxillary palp 47 the same, neotype of C. liebenauae (actually C. longisetosa) 48 the same, specimen from India.
Centroptella longisetosa. 49–51 paratype of C. longisetosa: tibiae of fore, middle and hind legs, view from anterior side (bases of long setae shown both on anterior and posterior sides) 52 neotype of C. liebenauae (actually C. longisetosa): tenth abdominal segment without caudalii, ventral view.
Larval exuviae of Centroptella longisetosa (neotype of C. liebenauae). 53–57 at the same magnification: (53) left half of pro- and mesonotum (54) thoracic pleura, left half of metanotum, fore- and hind legs (55) frons, antenna and labrum (56) tenth uromere and caudalii; (57) abdominal sterna and terga I–IX 58 margins of abdominal sterna IV–VII 59 cercus.
Larval exuviae of Centroptella longisetosa. 60–75 female, paratype of C. longisetosa: (60–68) fragments of abdominal terga II–IX (indicated by Roman numbers) (69) sternum VI (70–75) fragments of abdominal sterna IV–IX (indicated by Roman numbers) 76 male, neotype of C. liebenauae: fragment of abdominal sternum IX. Arrows on Figs
Subimago.
Adequately described by
Imago, male.
Adequately described by
Imago, female.
Adequately described by
Eggs
(extracted from mature female larva, paratype of C. longisetosa). Oval, chorion with numerous irregular small protuberances (Figs
According to original descriptions, specimens from Vietnam (type series of C. liebenauae) smaller, 3.7–4.3 mm; specimen from China (type series of C. longisetosa) larger, 3.9–5.2 mm.
All 8 examined specimens from India have maxillary palp relatively long (about 0.8 of lacinia length), while specimens from China and Vietnam have maxillary palp shorter (0.5–0.6 of lacinia length, Figs
The original description of C. longisetosa contains some errors. Instead of foreleg (
The larval metanotum is figured by
On the figure of male imaginal genitalia (
Chopralla
sp.:
Centroptella liebenauae:
Bungona (Chopralla) liebenauae:
Holotype
: L-S-I♂ {specimen [XV](1)2015}, THAILAND, Mae-Hong-Son Province, Pai, Mhor-Phaeng Falls, 11.II.2015, coll. N. Kluge & L. Sheyko (
This species is named in honour of Ingrid Müller-Liebenau.
Larva. Cuticular coloration. Frontal side of head colourless (Fig.
Shape and setation. Frontal suture short, nearly semicircular (as in Fig.
Pronotum with pair of protuberances near posterior margin (Figs
Scales on abdominal terga and sterna numerous, elongate, varying in size and shape, bordered by brown (Figs
Male genitalia. In last larval instar, developing subimaginal gonostyli folded under larval cuticle in peculiar pose, with 3rd segments bent medially-proximally (Fig.
Centroptella ingridae sp. nov. 94–102 fragments of abdominal terga I–IX (indicated by Roman numbers) 103–108 fragments of abdominal sterna IV–IX of male (indicated by Roman numbers) 109 fragment of abdominal sternum IX of female (94–101 and 103–108 male paratype of C. liebenauae; 102 and 109 female specimen from Thailand).
Centroptella of group inzingae-ingridae. Centroptella ingridae sp. nov. (holotype), fore wing 150 Centroptella inzingae (lectotype), genitalia (gv gonovectis us proximal margin of unistyliger) 151–152 Centroptella saxophila, gonovectes (specimens reared from larvae by N. Kluge in the Western Cape Province of South Africa).
Subimago.
Cuticle light brown with darker brown sutures of thorax; hypodermal coloration as in imago. On all legs of male and female all tarsal segments entirely covered by pointed microlepides (Fig.
Imago, male.
Head brown. Turbinate eyes relatively low and wide, with yellow stem and orange-red facetted surface (Fig.
Imago, female.
Head and thorax ochre with reddish markings (Figs
Egg
. Oval; chorion smooth, without relief (Fig.
Forewing length of male 4.7 mm; of female 5.0 mm.
Indochina: known from Thailand and Vietnam; recently reported from China (Yunnan, Guangxi, Guangdong) under the species name C. liebenauae (
Centroptella ingridae sp. nov. belongs to the inzingae-ingridae species group; male imagines of this group differ from all other Centroptella by halberd-like gonovectes and absence of the sterno-styligeral muscle (Fig.
Centroptella colorata Soldán, Braasch & Muu, 1987: 346 (larva)
Chopralla colorata:
Bungona (Chopralla) colorata:
Abdominal tergum IV without denticles on posterior margin, so regular row of denticles present on posterior margin of terga V–IX only. Tergum X without denticles on median part of posterior margin, with one pair of large denticles at sides (as in C. ceylonensis).
This research would not be possible without the invaluable and sincere help, consultations and type material of the recently deceased Tomáš Soldán. We thank Luke M. Jacobus (Purdue University, West Lafayette, Indiana, USA) for the important information and photos of the type specimens of fustipalpus [Cloeodes] deposited in Purdue University. We thank Phillip J. Suter (La Trobe University, Australia) for specimens of illiesi [Cloeodes] and important discussion.
We thank Helen Barber-James (Albany Museum) for the important information and photos of the type specimens of inzingae [Pseudocloeon] and saxophilum [Pseudocloeon] deposited in the Albany Museum. We acknowledge Helen Barber James and Nilgün Kazanci for comments and suggestions that improved the manuscript. Nikita J. Kluge acknowledges the Saint-Petersburg State University for research grant 1.52.1060.2015. Roman J. Godunko acknowledges the Czech Academy of Sciences for institutional support RVO: 60077344 and the Alexander von Humboldt Foundation (Bonn, Germany) for Georg Forster Research Fellowship for Experienced Researchers. Marek Svitok acknowledges the Operation Programme Research and Innovation (NFP: 13010T721). Milan Pallmann (State Museum of Natural History, Stuttgart, Germany) are acknowledged for technical assistance. Scanning electron microscopy was performed at the Laboratory of Electron Microscopy, Biology Centre of the Czech Academy of Sciences and the Center for Molecular and Cell Technologies of St. Petersburg State University.