Research Article |
Corresponding author: Grzegorz Paśnik ( pasnik@isez.pan.krakow.pl ) Academic editor: Michael Schmitt
© 2024 Adrian Smolis, Nataliya Kuznetsova, Grzegorz Paśnik.
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.
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With 54 described species, Endonura is one of the most species-rich genera of the Neanurinae. The taxon is characterised by the presence of 0–2 ocelli, separate tubercles Di and De with the non-crossed type of chaetotaxy on the head, and two or three tubercles on the fifth abdominal tergite. Four new species from the Crimea and the Caucasian region have been described based on adult specimens: Endonura armeniaca sp. nov., E. cochlearifera sp. nov., E. crimica sp. nov., and E. duplex sp. nov. A phylogenetic analysis of the genus Endonura using 70 characters of adult external morphology is presented. Two methods were chosen to examine different approaches to the reconstruction of evolutionary relationships: Maximum Parsimony (MP) and Bayesian Inference (BI). Besides supporting Endonura monophyly both analyses failed to resolve any of the deeper relationships within the genus. All known species within the genus are grouped into two main clades, A and D. Both of these clades have members distributed throughout Europe, the Caucasus and the western part of Central Asia. Endonura crimica sp. nov. can be considered as an intermediate between these two clades.
Cladistic analysis, Bayesian inference, Springtails, taxonomy
Springtails belonging to the subfamily Neanurinae are an example of evolutionary success, with over 800 currently described taxa, representing almost one tenth of all the known Collembola (
The evolutionary success of this subfamily is probably due to a number of factors. The first is the presence of body tubercles and well-developed chaetae, which in turn provides an important mechanical barrier for potential predators. The second is protection of toxic volatile chemicals, e.g. phenols (
The genus Endonura Cassagnau, 1979 is one of the most thoroughly studied and the largest genera within Neanuridae, with 54 valid species currently (
Material for taxonomic descriptions was collected during field trips to the Crimea (by Alexander Sharikov and Dmitry Shitikov; February 2015 and 2016) and the Caucasus (by: Boris Efeikin, Alexander Kremenitsa, Ivan Kuchiev, Nataliya Kuznetsova and Mikhail Potapov; September 1980, July 2015 and May 2016). Samples were taken from soil, litter and moss and extracted using a Berlese-Tullgren apparatus. After extraction, the specimens were preserved in alcohol. For detailed morphological analysis, they were cleared in Nesbitt’s fluid, mounted in Swan’s medium and examined using a Nikon Eclipse E600 phase-contrast microscope. Figures were drawn using a camera lucida and prepared for publication using Adobe Photoshop CS3.
Abbreviations used. General morphology: Abd. – abdomen; Ant. – antenna; AOIII – sensory organ of antennal segment III; Cx – coxa; Fe – femur; Scx2 – subcoxa 2; T – tibiotarsus; Th. – thorax; Tr – trochanter; VT – ventral tube. Groups of chaetae: Ag – antegenital; An – chaetae of anal lobes; ap – apical; ca – centroapical; cm – centromedial; cp – centroposterior; d – dorsal; Fu – furcal; vc – ventrocentral; Ve or ve – ventroexternal; Vea – ventroexternoanterior; Vem – ventroexternomedial; Vep – ventroexteroposterior; Vel – ventroexternolateral; Vec – ventroexternocentral; Vei – ventroexternointernal; Vi or vi – ventrointernal; Vl – ventrolateral. Tubercles: Af – antenno–frontal; Cl – clypeal; De – dorsoexternal; Di – dorsointernal; Dl – dorsolateral; L – lateral; Oc – ocular; So – subocular. Types of chaetae: Ml – long macrochaeta; Mc – short macrochaeta; me – mesochaeta; mi – microchaeta; ms – s–microchaeta; S or s – chaeta s; bs – s–chaeta on Ant. IV; miA – microchaetae on Ant. IV; iv – ordinary chaetae on ventral Ant. IV; or – organite of Ant. IV; brs – border s–chaeta on Ant. IV; i – ordinary chaeta on Ant. IV; mou – cylindrical s–chaetae on Ant. IV („soies mousses”); x – labial papilla x; L’ – ordinary lateral chaeta on Abd. V; B4, B5 – ordinary chaetae on tibiotarsi.
Terminology. Terminology and layout of the tables used in the paper follow
Depositories.
Department of Invertebrate Biology, Evolution and Conservation, University of Wrocław, Poland (DIBEC);
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland (
To investigate the phylogenetic relationships between species of the genus Endonura, 43 of the 54 known species were selected for analyses (Table 1). Because of the unavailability of specimens for examination and insufficient descriptions, the following species were not included in the analyses: E. caeca (Gisin, 1963), E. granulata (Cassagnau, Delamare & Deboutteville, 1955), E. ichnusae Dallai, 1983, E. immaculata Deharveng, 1980, E. ludovicae (Denis, 1947), E. occidentalis Deharveng, 1979, E. pejai Deharveng, 1980, E. poinsotae Deharveng, 1980, E. tartaginensis (Deharveng, 1980), E. tetrophthalma (Stach, 1929) and E. tyrrhenica Dallai, 1983.
Species | Number of examined specimens | Status of examined types | Collections | Other sources of data e.g. literature |
Endonura agnieskae | 24 | Holotype, Paratypes |
|
— |
E. aibgai | 12 | Holotype, Paratypes |
DIBEC, |
— |
E. annae | 22 | Holotype, Paratypes |
|
— |
E. alavensis | — | — | — |
|
E. alticola | 1 | Holotype |
|
— |
E. arbasensis | — | — | — |
|
E. asiatica | 11 | Holotype, Paratypes |
|
— |
E. baculifer | — | — | — |
|
E. cantabrica | — | — | — |
|
E. carpatica | 8 | Holotype, Paratypes | DIBEC | — |
E. centaurea | 3 | Syntypes |
|
— |
E. ceratolabralis | 7 | Holotype, Paratypes | DIBEC | — |
E. colorata | — | — | — |
|
E. cretensis | 7 | Holotype |
|
— |
E. cryptopyga | 4 | Holotype |
|
— |
E. dalensi | — | — | — |
|
E. dentifera | 7 | Holotype, Paratypes |
DIBEC, |
— |
E. deharvengi | 2 | Syntypes |
|
— |
E. dichaeta | 14 | Holotype, Paratypes | DIBEC | — |
E. diminutichaeta | 11 | Holotype, Paratypes |
|
— |
E. dobrolyubovae | 8 | Holotype, Paratypes |
|
— |
E. dudichi | 40 | — | DIBEC | — |
E. gracilirostris | 5 | Holotype, Paratypes |
DIBEC, |
— |
E. gladiirostra | 1 | Holotype | DIBEC | — |
E. gladiolifer | — | — | — |
|
E. incolorata | 26 | Lectotype, Paralectotype |
|
— |
E. kremenitsai | 11 | Holotype, Paratypes |
|
— |
E. levantica | 2 | Holotype, Paratype |
DIBEC, |
— |
E. longirostris | 4 | Holotype, Paratypes | DIBEC | — |
E. lusatica | 38 | Holotype, Paratype |
|
— |
E. ossetica | 2 | Holotype, Paratype |
|
— |
E. paracantabrica | 5 | Holotype, Paratypes |
|
— |
E. paracentaurea | 4 | Holotype, Paratypes | DIBEC | — |
E. persica | 14 | Holotype, Paratypes | DIBEC | — |
E. reticulata | 52 | — |
DIBEC, |
— |
E. saleri | — | — | — |
|
E. schwendingeri | 9 | Holotype, Paratypes |
|
— |
E. tatricola | 162 | Lectotype |
|
— |
E. taurica | 11 | Holotype, Paratypes |
|
— |
E. transcaucasica | 1 | Lectotype |
|
— |
E. turkmenica | 2 | Holotype, Paratype | DIBEC | — |
E. quadriseta | 17 | Holotype, Paratype |
|
— |
E. urotuberculata | 2 | Holotype, Paratype |
|
— |
Neanura muscorum | 177 | — | DIBEC | — |
Kalanura babenkoi | 34 | Holotype, Paratypes |
DIBEC, |
— |
Xylanura oregonensis | 47 | Holotype, Paratypes |
|
— |
The four new species described in this paper were included in the phylogenetic analysis. Forty-seven species were analysed out of 58. The trees were rooted using Neanura muscorum (Templeton, 1836), Kalanura babenkoi Smolis, 2007 and Xylanura oregonensis
Species were selected based on specimen availability, with a preference for the type-specimens. Most characters were based on personal observations of specimens. These observations were supplemented by previous taxonomic/phylogenetic studies (
A total of 70 characters were scored for the study taxa, including 61 binary characters and 9 multistate characters. Missing data were coded as ‘?’ in the matrix (Table 2). All characters were treated as unordered (
Taxon | Characters | Characters |
11111111112222222222333333 12345678901234567890123456789012345 |
33334444444444555555555566666666667 67890123456789012345678901234567890 |
|
N. muscorum | 00010100000110001000000010100010100 | 01010000010100101100000001001101100 |
K. babenkoi | 00010000000110001000000010000010100 | 01100000000000001100000001000011110 |
X. oregonensis | 00011011110010001030100010000010121 | 1110010101010110000101?001001011000 |
E. agnieskae | 10011000000110001000000000010110100 | 01111110000100101110000001101111110 |
E. aibgai | 10011000000111121001?00010010110100 | 01001111000100101110000001111010110 |
E. alavensis | 100110011001100110201000100101102?? | ?1001?1???0????0??100010?00??011111 |
E. alticola | 10011000000110001000000010010110210 | 011011110001001111100000??111?11111 |
E. annae | 10011001000110021100110000011200000 | 0100101001010100001101?001101010110 |
E. arbasensis | 111?100000011000100000001001111???? | ?1001111000000?0??100010?000?011111 |
E. armeniaca | 10011001000112121101?001?1?10100200 | 000011111?111?10111101?0?1111111111 |
E. asiatica | 10001000000110001000000010010110100 | 00001111000100101110101000111011111 |
E. baculifer | 10011001100111111121?1001001110???? | ?0001010010101?0111010100110?0111?? |
E. cantabrica | 10011001000110011001?0001001011???? | ?1001111000100?0??100000?100?0111?? |
E. carpatica | 111?1000100110021121?10000011111100 | 00001010010101001110000001101011111 |
E. centaurea | 11011001001110011101?00000011110110 | 000010100101010011100010?0101011110 |
E. ceratolabralis | 10011001000110011100100000010110221 | 111010100101010011100010?0101011111 |
E. cochlearifera | 10011000000110011000000000010110200 | 00001111000100101110100001001111110 |
E. colorata | 10011001110110001021?0000001011???? | ??001?1???0????0??101000?100?011110 |
E. cretensis | 10011000100012121020000010010111210 | 011111111?111?10221101?101000111111 |
E. crimica | 10011001000100011001?00000010110100 | 010010100001001011100000?1000011110 |
E. cryptopyga | 10011000000111121101?00010010100200 | 010111110001001011100000?1001110010 |
E. dalensi | 1001100000001001100000001001111???? | ?1001111000000?0??100010?100?011111 |
E. deharvengi | 10011000000111121000000010011?10110 | 00001010010101001110100001100011111 |
E. dentifera | 10011000000110001100000000010110100 | 00001111000100101110100001001110001 |
E. dichaeta | 11011001100111121121?00000011?10210 | 00001010010101001111000000101011111 |
E. diminutichaeta | 10011001000111101000100010010?00200 | 01101111000100102210001001001111010 |
E. dobrolyubovae | 10011001000111121101?00000010?00100 | 00001111000100101110100011001?10110 |
E. dudichi | 11011001000110000101?00000011?10100 | 01001010010101011110000000100111110 |
E. duplex | 10011000000110001000000000010110100 | 01001111000100101110100001000011110 |
E. gladiirostra | 10011011110110011130100010010200221 | 111110100101010000100010?1101?11110 |
E. gladiolifer | 10011000100110001021?000?001010???? | ?11110100111???0111011?0?000?010111 |
E. gracilirostris | 10011000100100011020000000010?10220 | 00001111000100101110001001101?11010 |
E. incolorata | 11011000000110000101?00000011110100 | 00001010??0101001120001000100011111 |
E. kremenitsai | 10011000000111101000000010010110100 | 01101111000100101110000011001111110 |
E. levantica | 10011000000110021000000000011110120 | 000110100101010011100000?0101011111 |
E. longirostris | 10011000000112121101?00010011110220 | 000011110000001012100010?1100011111 |
E. lusatica | 10011000000111011000000000010110200 | 00001111000100101110100001001010010 |
E. ossetica | 10011001010110021110100010011100200 | 011010100001011010101010?1001011111 |
E. paracantabrica | 10011000000110001001?00010010110100 | 01101111000100101110000001001011010 |
E. paracentaurea | 10011001000110021101?00000011110120 | 010010100101010011100000?0101011111 |
E. persica | 10011000000111121100000010010200100 | 01001111000100100010000001001010010 |
E. quadriseta | 10011000100110011020000010010110100 | 00001111000100101110001011000011111 |
E. reticulata | 10011000000110001000000000010110100 | 00001111000000001110100001000011110 |
E. saleri | 10011000000111111111?00000011111120 | 000010100001000011100000?1001111111 |
E. schwendingeri | 10011001000111121101?001?1?10100100 | 00001111000100101110001011111110110 |
E. tatricola | 11011001000110000101?00000011110100 | 00001010010101001110000000100011111 |
E. taurica | 10011001100100011120100010010110000 | 00001111000100101110000001001110011 |
E. transcaucasica | 10011001000110021100111000011110210 | 010010100101010011100010?1101011111 |
E. turkmenica | 10011000000110021100000000010110220 | 00001010010101001110000001101011110 |
E. urotuberculata | 111?1000000010011100000000011110100 | 00001111000000101110001100101011111 |
The morphological dataset was analysed using both Maximum Parsimony (MP) and Bayesian Inference (BI).
Parsimony analyses utilized New Technology heuristic searches implemented in the program TNT v. 1.5 (
Some argue (e.g.
Clade supports were assessed based on Bremer support (BS) (
The Symmetric Resampling (SR) support calculated the differences in the frequencies of a given group and its most frequent contradictory group (GC). The analyses were run in TNT with the traditional search, using 10,000 replications, change probability of 0.33, two initial Wagner trees, and holding three trees per replicate.
The following values were applied to support the clades: weak (SR<50%, BS 1–2), moderate (SR 51–75%, BS 3–6), good (SR 76–90%, BS 7–8), and strong (SR>90%, BS 9–10).
The synapomorphies were mapped in WinClada onto the most parsimonious tree using an option showing unambiguous changes only.
Bayesian inferences were performed in MrBayes v3.2.7 (
In tree resulting from Bayesian inference, Posterior Probability (PP) was interpreted as statistical support values.
Neanura (Endonura) Cassagnau, 1979: 192.
Achorutes tetrophtalmus Stach, 1929: 282.
0–2 + 0–2 eyes, pigmented or not. Colour of body blue, purple brown, white, never yellow or red. Dorsal tubercles present, well developed. Mouthparts feebly developed, maxilla styliform, mandible slender or rarely strong. Labral chaetotaxy: 4/2, 4; sometimes prelabral or labral chaetae reduced. Sensilla S on Ant. IV of similar size. Head with 10 or 12 tubercles. Tubercles Af and Cl separate or fused. Tubercles Di and De on head separate, sometimes Di not differentiated. Arrangement of chaetae Di and De on head of the „non-croisé” type (
Holotype: adult female on slide, Armenia, road to Agveran village (40°29´32.6´´ N; 44° 35´35.1´´ E), mountain oak forest, litter and soil, 24.V.2016, leg. B. Efeikin (DIBEC). Paratype: adult female on slide, same data as holotype.
2+2 pigmented eyes. Buccal cone long, labrum ogival. Head with chaetae B, C and D. Chaetae A, O, Ocp and E absent. Tubercles Cl and Af separate. Tubercles Dl and (L+So) on head with 4 and 7 chaetae respectively. Tubercles Di on Th. I absent. Tubercles De on Th. II and III with 3 chaetae. Tubercles L on Abd. III and IV with 3 and 5 chaetae respectively. Abd. IV and V with 8 and 3 tubercles respectively. Claw without inner tooth. Tibiotarsi with chaetae B4 and B5 short.
Body length
: 1.1 to 1.2 mm (without antennae) (holotype: 1.2 mm). — Colour: pale bluish grey (alive and in alcohol). 2+2 black eyes, in a typical arrangement for the genus (one anterior, one posterior, Fig.
Chaetotaxy of Endonura armeniaca sp. nov.: cephalic chaetotaxy–dorsal side.
Tubercle | Number of chaetae | Types of chaetae | Names of chaetae |
Cl | 4 | Ml | F |
Mc | G | ||
Af | 6 | Ml | B |
Mc | C, D | ||
Oc | 2 | Ml | Ocm |
mi | Oca | ||
Di | 2 | Ml | Di1 |
Mc | Di2 | ||
De | 2 | Ml | De1 |
Mc | De2 | ||
Dl | 4 | Ml | Dl5, Dl1 |
Mc | Dl4 | ||
mi | Dl2 | ||
(L+So) | 7 | Ml | L1, L4, So1 |
me | So3–6 |
Segment, Group | Number of chaetae | Segment, Group | Number of chaetae adult |
I | 7 | IV | or, 8 S, i, 12 mou, 6 brs, 2 iv |
II | 11 | ||
III | 5 sensilla AO III | ||
ve | 5 | ap | 8 bs, 5 miA |
vc | 4 | ca | 2 bs, 3 miA |
vi | 4 | cm | 3 bs, 1 miA |
d | 5 | cp | 8 miA, 1 brs |
Terga | Legs | ||||||||
Di | De | Dl | L | Scx2 | Cx | Tr | Fe | T | |
Th. I | 1 | 2 | 1 | — | 0 | 3 | 6 | 13 | 19 |
Th. II | 3 | 2+s | 3+s+ms | 3 | 2 | 7 | 6 | 12 | 19 |
Th. III | 3 | 2+s | 3+s | 3 | 2 | 8 | 6 | 11 | 18 |
Sterna | |||||||||
Abd. I | 2 | 2+s | 2 | 2 | VT: 4 | ||||
Abd. II | 2 | 2+s | 2 | 2 | Ve: 3–6; chaeta Ve1 present | ||||
Abd. III | 2 | 2+s | 2 | 3 | Vel: 3–5; Fu: 5 me, 0 mi | ||||
Abd. IV | 2 | 1+s | 3 | 5 | Vel: 4; Vec: 1–2; Vei: 2; Vl: 4 | ||||
Abd. V | (2+2) | 4+s | Ag: 3; Vl: 1 | ||||||
Abd. VI | 7 | Ve: 10–11; An: 2mi |
Endonura armeniaca sp. nov.: 1 dorsal chaetotaxy of head and Th. I–II; 2 mandible; 3 maxilla; 4 apical bulb, ventral view; 5 apical bulb, dorsal view; 6 ventral chaetotaxy of head; 7 S– chaeta of Ant. IV; 8 sensillum sgd; 9 ventral chaetotaxy of Ant. III; 10 dorsal chaetotaxy of Abd. III–VI; 11 ventral sclerifications of labrum; 12 chaetotaxy of labrum; 13 sensillum of Abd. V; 14 chaeta Di1 of Abd. V; 15 tubercle L of Abd. IV.
The name armeniaca is derived from Armenia, the country where the species was found.
Among the known members of the genus, E. armeniaca sp. nov. is distinguished by an extraordinary reduction of dorsal cephalic chaetotaxy. This includes the absence of chaetae A, O, E, Ocp, Dl3, L2, L3 and So2, as well as the presence of an ogival labrum and the absence of an internal tooth on the claws. Morphologically, the new species appears to be most similar to E. schwendingeri Smolis and Skarżyński, 2020, a species recently described from northwestern Iran (
E. armeniaca sp. nov. is also similar to E. cretensis (Ellis, 1976) and E. quadriseta Cassagnau and Peja, 1979, which were described from Greece and recently redescribed (
Morphological differences between E. armeniaca sp. nov., E. schwendingeri, E. cretensis and E. quadriseta.
Characters | E. armeniaca sp. nov. | E. schwendingeri | E. cretensis | E. quadriseta |
Shape of labrum | ogival | not ogival | ogival | not ogival |
No. of labial chaetae | 11 | 11 | 8 | 11 |
No. of prelabral chaetae | 4 | 4 | 2 | 4 |
Cephalic chaetae A, E and O | absent | absent | present | present |
No. of cephalic chaetae Dl | 4 | 5 | 3 | 6 |
No. of cephalic chaetae (L+So) | 7 | 7 | 7 | 9 |
No. of cephalic chaetae Oc | 2 | 2 | 3 | 3 |
Tubercle Di on Th. I | absent | absent | present | absent |
No. of chaetae De on Th. III | 2+s | 3+s | 2+s | 3+s |
No. of chaetae De on Abd. I–III | 2+s | 3+s | 2+s | 3+s |
No. of chaetae L on Abd. III and IV | 3 and 5 | 2 and 4 | 4 and 7 | 4 and 6–8 |
No. of chaetae Di on Abd. V | 2+2 | 3+3 | 2+2 | 3+3 |
Cauliflower-like tubercles on Abd. IV–VI | absent | absent | present | absent |
Male ventral organ | unknown | present | absent | present |
Internal tooth on claws | absent | present | absent | absent |
Holotype: adult female on slide, Russia, Caucasus, Northern Ossetia, North Ossetia Nature Reserve, surroundings of the village Tsey, green moss pine forest, in mosses, 23.IX.1980, leg. I. Kuchiev (DIBEC). Paratypes: 4 adult females, subadult male and 2 juveniles on slides, same data as holotype (DIBEC and
2+2 pigmented eyes. Buccal cone long, labrum ogival. Head with chaetae A, B, C, D, E and O. Tubercles Cl and Af separate. Tubercles Dl and (L+So) on head with 6 and 9 chaetae respectively. Tuberles Di on Th. I absent. Tubercles De on Th. II and III with 3 and 4 chaetae respectively. Tubercles L on Abd. III and IV with 4 and 6–7 chaetae respectively. Abd. IV and V with 8 and 3 tubercles respectively. Claw without inner tooth. Tibiotarsi with chaetae B4 and B5 short.
Body length
: 0.55 (juvenile) to 1.45 mm (without antennae) (holotype: 1.45 mm). — Colour: body pale bluish grey (alive and in alcohol). 2+2 black eyes, in a typical arrangement for the genus (Fig.
Chaetotaxy of Endonura cochlearifera sp. nov.: cephalic chaetotaxy-dorsal side.
Tubercle | Number of chaetae | Types of chaetae | Names of chaetae |
Cl | 4 | Ml | F |
Mc | G | ||
Af | 11 | Ml | B |
Mc | A, C, D, E, O | ||
Oc | 3 | Ml | Ocm |
Mc | Ocp | ||
mi | Oca | ||
Di | 2 | Ml | Di1 |
Mc | Di2 | ||
De | 2 | Ml | De1 |
Mc | De2 | ||
Dl | 6 | Ml | Dl5, Dl1 |
Mc | Dl2, Dl3, Dl4, Dl6 | ||
(L+So) | 9 | Ml | L1, L4, So1 |
Mc | L2 | ||
mi | So2 | ||
me | So3–6 |
Segment, Group | Number of chaetae | Segment, Group | Number of chaetae adult |
I | 7 | IV | or, 8 S, i, 12 mou, 6 brs, 2 iv |
II | 12 | ||
III | 5 sensilla AO III | ||
ve | 5 | ap | 8 bs, 5 miA |
vc | 4 | ca | 2 bs, 3 miA |
vi | 4 | cm | 3 bs, 1 miA |
d | 5 | cp | 8 miA, 1 brs |
Chaetotaxy of Endonura cochlearifera sp. nov.: postcephalic chaetotaxy.
Terga | Legs | ||||||||
Di | De | Dl | L | Scx2 | Cx | Tr | Fe | T | |
Th. I | 1 | 2 | 1 | — | 0 | 3 | 6 | 13 | 19 |
Th. II | 3 | 2+s | 3+s+ms | 3 | 2 | 7 | 6 | 12 | 19 |
Th. III | 3 | 3+s | 3+s | 3 | 2 | 8 | 6 | 11 | 18 |
Sterna | |||||||||
Abd. I | 2 | 3+s | 2 | 3 | VT: 4 | ||||
Abd. II | 2 | 3+s | 2 | 3 | Ve: 5; chaeta Ve1 present | ||||
Abd. III | 2 | 3+s | 2 | 4 | Vel:5–6; Fu: 5–7 me, 0 mi | ||||
Abd. IV | 2 | 2+s | 3 | 6–7 | Vel: 4; Vec: 2; Vei: 2; Vl: 4 | ||||
Abd. V | (3+3) | 8–9 +s | Ag: 3; Vl: 1 | ||||||
Abd. VI | 7 | Ve: 14; An: 2mi |
Endonura cochlearifera sp. nov.: 16 dorsal chaetotaxy of head and Th.; 17 dorsal chaetotaxy of Ant. III–IV; 18 ventral chaetotaxy of Ant. III; 19 mandible; 20 maxilla; 21 dorsal chaetotaxy of Abd. III–VI; 22 ventral sclerifications of labrum; 23 chaetotaxy of labrum; 24 sensillum of Abd. V; 25 chaeta Di1 of Abd. V.
The name cochlearifera refers to the spoon-like shape of the labral sclerifications of this species.
E. cochlearifera sp. nov. is unique and differs from other species of the genus by the spoon-like shape of the apical labral sclerifications. In general appearance (e.g. dorsal chaetotaxy of head and thorax) the new species strongly resembles E. annae Smolis & Skarżyński, 2020, E. dentifera
Morphological differences between E. cochlearifera sp. nov., E. lusatica, E. dentifera and E. annae.
Characters | E. cochlearifera sp. nov. | E. lusatica | E. dentifera | E. annae |
Shape of apical labral sclerifications | spoon-like | ogival | not ogival | not ogival |
Shape of labrum | elongated and narrow | elongated and narrow | rather long and narrow | short and wide |
Cephalic chaeta O | present | present | absent | absent |
Cephalic chaetae C | present | present | present | absent |
No. of cephalic chaetae Dl | 6 | 5 | 6 | 6 |
No. of cephalic chaetae (L+So) | 9 | 9 | 10 | 8 |
No. of chaetae L on Abd. IV | 6–7 | 7 | 8–9 | 6 |
No. of chaetae Di on Abd. V | 3+3 | 3+3 | 3+3 | 2+2 |
No. of ordinary chaetae (De+Dl+L) on Abd. V | 8–9 | 7 | 8 | 5 |
Chaeta L’ on Abd. V | absent | present | absent | present |
Length and shape of tibiotarsal chaetae B4 and B5 | short and pointed | elongated and clavate | elongated and clavate | short and pointed |
Internal tooth on claws | absent | present | present | present |
Holotype: adult female on slide, Crimea, Jaltynskyi Nature Reserve (34 24.344 N; 44 55.137 E), 570 m alt., mixed forest (pine, beech), 3.II.2016, leg. D. Shitikov, A. Szarikov (DIBEC). Paratype: subadult male on slide, Crimea, Ajudag near Gurzuf, 450 m alt., oak forest, 6.II.2015, leg. D. Shitikov, A. Szarikov (
2+2 pigmented eyes. Buccal cone relatively short and narrow, labrum not ogival. Head with chaetae A, B, C, D and O. Tubercles Cl and Af separate. Tubercles Dl and (L+So) on head with 6 and 9 chaetae respectively. Tuberles Di on Th. I present. Tubercles De on Th. II and III with 3 and 4 chaetae respectively. Tubercles L on Abd. III and IV with 4 and 6–7 chaetae respectively. Abd. IV and V with 8 and 3 tubercles respectively. Claw without inner tooth. Tibiotarsi with chaetae B4 and B5 short.
Body length
: 1.05 (subadult male) to 1.15 mm (holotype) (without antennae). — Colour: body pale bluish grey (alive and in alcohol). 2+2 black eyes, in a typical arrangement for the genus (Fig.
Chaetotaxy of Endonura crimica sp. nov.: cephalic chaetotaxy–dorsal side.
Tubercle | Number of chaetae | Types of chaetae | Names of chaetae |
Cl | 4 | Ml | F |
Mc | G | ||
Af | 9 |
Ml | B |
Mc | A, C, D, O | ||
Oc | 3 | Ml | Ocm |
Mc | Ocp | ||
mi | Oca | ||
Di
|
2 | Ml | Di1 |
Mc | Di2 | ||
De | 2 | Ml | De1 |
Mc | De2 | ||
Dl | 6 | Ml | Dl5, Dl1 |
Mc | Dl3, Dl4 | ||
mi | Dl2, Dl6 | ||
(L+So) | 9 | Ml | L1, L4, So1 |
Mc | L2 | ||
mi | So2 | ||
me | So3–6 |
Segment, Group | Number of chaetae | Segment, Group | Number of chaetae adult |
I | 7 | IV | or, 8 S, i, 12 mou, 6 brs, 2 iv |
II | 12 | ||
III | 5 sensilla AO III | ||
ve | 5 | ap | 8 bs, 5 miA |
vc | 4 | ca | 2 bs, 3 miA |
vi | 4 | cm | 3 bs, 1 miA |
d | 5 | cp | 8 miA, 1 brs |
Terga | Legs | ||||||||
Di | De | Dl | L | Scx2 | Cx | Tr | Fe | T | |
Th. I | 1 | 2 | 1 | — | 0 | 3 | 6 | 13 | 19 |
Th. II | 3 | 2+s | 3+s+ms | 3 | 2 | 7 | 6 | 12 | 19 |
Th. III | 3 | 3+s | 3+s | 3 | 2 | 8 | 6 | 11 | 18 |
Sterna | |||||||||
Abd. I | 2 | 3+s | 2 | 3 | VT: 4 | ||||
Abd. II | 2 | 3+s | 2 | 3 | Ve: 5; chaeta Ve1 present | ||||
Abd. III | 2 | 3+s | 2 | 4 | Vel: 4–5; Fu: 5–7 me, 0 mi | ||||
Abd. IV | 2 | 2+s | 3 | 6–7 | Vel: 4; Vec: 2; Vei: 2; Vl: 4 | ||||
Abd. V | (3+3) | 7+s | Ag: 2–3; Vl: 1 L‘: 1 | ||||||
Abd. VI | 7 | Ve: 14; An: 2mi |
Endonura crimica sp. nov.: 26 dorsal chaetotaxy of Ant. III–IV; 27 ventral chaetotaxy of Ant. III; 28 dorsal chaetotaxy of head and Th.; 29 chaeta Di1 of Abd. V; 30 sensillum of Abd. V; 31 maxilla; 32 mandible; 33 chaetotaxy and ventral sclerifications of labrum; 34 dorsal chaetotaxy of Abd. III–VI; 35 tubercle L of Abd. IV; 36 tibiotarsus and claw of leg III, lateral view.
The name crimica refers to the geographic area of its collecting.
E. crimica sp. nov. belongs to a very small group of Endonura characterised by a broad and short (equal to or shorter than tubercle Oc) cephalic tubercle Af. In addition to the new species, this group includes E. taurica (Stach, 1951) and E. gracilirostris
Morphological differences between E. crimica sp. nov., E. taurica, E. gracilirostris and E. colorata.
Characters | E. crimica sp. nov. | E. taurica | E. gracilirostris | E. colorata |
Shape of labrum | not ogival | not ogival | ogival | not ogival |
Prelabral chaetae | present | present | absent | present |
Shape of manible | thin | thick | thin | thin |
Shape of cephalic tubercle AF | wide | wide | wide | narrow |
Length of cephalic tubercle AF | equal or shorter than tubercle Oc | equal or shorter than tubercle Oc | equal or shorter than tubercle Oc | longer than tubercle Oc |
Cephalic chaetae E | present | absent | absent | present |
Cephalic chaeta O | present | absent | present | present |
Position of cephalic chaetae D | free | located within tubercle Cl | located within tubercle Cl | located within tubercle Cl |
Tubercle Di on Th. I | present | absent | absent | present |
Position of chaetae De2 on Th. II–III | located within tubercle De | free | free | located within tubercle De |
No. of ordinary chaetae (De+Dl+L) on Abd. V | 7 | 8 | 8 | 6 |
Ratio Di1/Di2/Di3 on Abd. V | 50:21:3 | 48:18:2 | 46:11:2 | 40:22:10 |
No. of mesochaetae on furca rudimentary | 5‒7 | 5 | 5‒6 | 8‒9 |
Length and shape of tibiotarsal chaeta B4 | short and pointed | elongated and clavate | elongated and clavate | short and pointed |
Holotype: adult female on slide, Russia, NW Caucasus, Adygeya, Caucasus Nature Reserve, ‘Kamennoye More’ ridge, southern slope, 1850 m. alt., mixed forest (pine, birch), coniferous litter, 21.VII.2015, leg. M. Potapov, N. Kuznetsova, A. Kremenitsa, L.Vanyavina (DIBEC). Paratype: subadult male and 2 juveniles on slides, same data as holotype (DIBEC and
2+2 pigmented eyes. Buccal cone short, labrum not ogival. Head with chaetae A, B, C, D and E. Two chaetae O present. Tubercles Cl and Af separate. Tubercles Dl and (L+So) on head with 6 and 10 chaetae respectively. Tuberles Di on Th. I present. Tubercles De on Th. II and III with 3 and 4 chaetae respectively. Tubercles L on Abd. III and IV with 4 and 7 chaetae respectively. Abd. IV and V with 8 and 3 tubercles respectively. Claw without inner tooth. Tibiotarsi with chaetae B4 and B5 short.
Body length
: 0.55 (juvenile) to 1.55 mm (holotype) (without antennae). — Colour: body pale bluish grey (alive and in alcohol). 2+2 black eyes, in a typical arrangement for the genus (Fig.
Chaetotaxy of Endonura duplex sp. nov.: cephalic chaetotaxy–dorsal side.
Tubercle | Number of chaetae | Types of chaetae | Names of chaetae |
Cl | 4 | Ml | F |
Mc | G | ||
Af | 12 | Ml | B |
Mc | A, C, D, E, O | ||
Oc | 3 | Ml | Ocm |
Mc | Ocp | ||
mi | Oca | ||
Di | 2 | Ml | Di1 |
Mc | Di2 | ||
De | 2 | Mc | De1, De2 |
Dl | 6 | Ml | Dl5, Dl1 |
Mc | Dl2, Dl3, Dl4, Dl6 | ||
(L+So) | 10 | Ml | L1, L4, So1 |
Mc | L2, L3, So2 | ||
me | So3–6 |
Segment, Group | Number of chaetae | Segment, Group | Number of chaetae adult |
I | 7 | IV | or, 8 S, i, 12 mou, 6 brs, 2 iv |
II | 12 | ||
III | 5 sensilla AO III | ||
ve | 5 | ap | 8 bs, 5 miA |
vc | 4 | ca | 2 bs, 3 miA |
vi | 4 | cm | 3 bs, 1 miA |
d | 5 | cp | 8 miA, 1 brs |
Terga | Legs | ||||||||
Di | De | Dl | L | Scx2 | Cx | Tr | Fe | T | |
Th. I | 1 | 2 | 1 | — | 0 | 3 | 6 | 13 | 19 |
Th. II | 3 | 2+s | 3+s+ms | 3 | 2 | 7 | 6 | 12 | 19 |
Th. III | 3 | 3+s | 3+s | 3 | 2 | 8 | 6 | 11 | 18 |
Sterna | |||||||||
Abd. I | 2 | 3+s | 2 | 2–3 | VT: 4 | ||||
Abd. II | 2 | 3+s | 2 | 3 | Ve: 5; chaeta Ve1 present | ||||
Abd. III | 2 | 3+s | 2 | 4 | Vel: 5; Fu: 5 me, 0 mi | ||||
Abd. IV | 2 | 2+s | 3 | 7 | Vel: 4; Vec: 2; Vei: 2; Vl: 4 | ||||
Abd. V | (3+3) | 7+s | Ag: 3; Vl: 1 L‘: 1 | ||||||
Abd. VI | 7 | Ve: 14; An: 2mi |
Endonura duplex sp. nov.: 37 tubercle L of Abd. IV; 38 dorsal chaetotaxy of head and Th.; 39 apical bulb, ventral view; 40 apical bulb, dorsal view; 41 dorsal chaetotaxy of Ant. III–IV; 42 ventral chaetotaxy of Ant. III; 43 dorsal chaetotaxy of Abd. III–VI; 44 ventral sclerifications of labrum; 45 chaetotaxy of labrum; 46 sensillum of Abd. V; 47 chaeta Di1 of Abd. V.
The name duplex is derived from the presence of two chaetae O on head.
E. duplex sp. nov. differs from all other Endonura by the presence of two cephalic chaetae O. Besides this character, the new species strongly resembles E. agnieskae Smolis & Skarżyński, 2020, E. alticola (Stach, 1951) and E. reticulata (Axelson, 1905). However, the differences between the new species and the outlined taxa include a number of characters: Shape of labrum (ogival in alticola, not ogival in others), number of prelabral chaetae (two in alticola, four in others), number of lateral labial chaetae (three in agnieskae, four in others), length of cephalic chaetae A and Ocp (chaeta A longer than Ocp in duplex and alticola, chaeta A shorter than Ocp in reticulata, equal in agnieskae), presence of tubercle Di on Th. I (present in duplex, present and fused with tubercle De in agnieskae and alticola, usually absent in reticulata), position of chaeta Di3 on Th. II–III (within tubercle Di in alticola, free in others), position of chaeta De2 on Abd. I–III (free in reticulata, within tubercle De in others), position of chaeta s on Abd. I–III (line of chaeta De1-chaeta s not parallel to dorsomedial line in reticulata, parallel in others), fusion of tubercles Di on Abd. IV (present in alticola, absent in others) and presence of free chaeta L on Abd. IV (present in duplex, absent in others). The differences between the species mentioned in the remarks are summarised in Table 10.
Morphological differences between E. duplex sp. nov., E. agnieskae, E. alticola and E. reticulata.
Characters | E. duplex sp. nov. | E. agnieskae | E. alticola | E. reticulata |
Shape of labrum | not ogival | not ogival | ogival | not ogival |
No. of prelabral chaetae | 4 | 4 | 2 | 4 |
No. of lateral labial chaetae | 4 | 3 | 4 | 4 |
No. of cephalic chaetae O | 2 | 1 | 1 | 1 |
Length of cephalic chaetae A and Ocp | chaeta A longer than Ocp | equal in lenght | chaeta A longer than Ocp | chaeta A shorter than Ocp |
Tubercle Di on Th. I | present and not fused with De | present and fused with De | present and fused with De | usually absent |
Position of chaetae Di3 on Th. II–III | free | free | located within tubercle Di | free |
Position of chaeta De2 on Abd. I–III | located within tubercle De | located within tubercle De | located within tubercle De | free |
Position of chaeta De2 on Abd. I–III | line of chaeta De1–chaeta s parallel to dorsomedial line | line of chaeta De1–chaeta s parallel to dorsomedial line | line of chaeta De1–chaeta s parallel to dorsomedial line | line of chaeta De1–chaeta s not parallel to dorsomedial line |
Fusion of tubercles Di on Abd. IV | absent | absent | present | absent |
Number of chaetae Lon Abd. III and IV | 4 and 7 | 3 and 5‒6 | 4 and 6 | 4 and 6‒9 |
Free chaeta L on Abd. IV | present | absent | absent | absent |
Head eyes
1. Number of eyes on each side of head: (0) 3 eyes present; (1) 0–2 eyes present.
2. Pigmentation of eyes: (0) present; (1) absent.
3. Presence of anterior eyes: (0) present (Fig.
4. Position of anterior eyes: (0) outside tubercle Oc (Fig.
Head tuberculation
5. Tubercle L: (0) separate (Fig.
6. Tubercle Dl: (0) separate (Fig.
7. Tubercle Af: (0) separate (Fig.
8. Elementary tubercles BE: (0) absent; (1) present (Fig.
9. Elementary tubercles DF: (0) absent; (1) present (Fig.
10. Elementary tubercles DE: (0) absent; (1) present (Fig.
11. Non-reticulate area within tubercle Af: (0) absent; (1) present (Fig.
12. Tubercles Di: (0) tubercles absent; (1) tubercles present (Fig.
13. Length of tubercle Af: (0) longer than tubercle Oc (Fig.
Head chaetotaxy
14. Number of chaetae Dl (on half of head): (0) 6 chaetae present (Fig.
15. Chaeta Dl3: (0) present (Fig.
16. Number of chaetae L+So (on half of head): (0) 10 chaetae present (Fig.
17. Position of chaeta L4: (0) included within tubercle L+So (Fig.
18. Chaeta O: (0) present (Fig.
19. Chaetae D: (0) free (not included in tubercle Af or Cl) (Fig.
20. Chaetae E: (0) present (Fig.
21. Position of chaetae E: (0) free, not included in tubercle Af (Fig.
22. Chaetae C: (0) present (Fig.
23. Chaetae Oca: (0) present (Fig.
24. Chaetae Ocp: (0) present (Fig.
25. Length of chaetae Ocp: (0) longer than chaeta A (Fig.
26. Chaetae A: (0) present (Fig.
27. Length of chaetae A: (0) shorter than chaetae B (Fig.
28. Arrangement of chaetae Di2 and De2: (0) “croisée” (line between chaetae Di 2 and De2 crosses line between Di1 and De1,
Body
29. Color in alive: (0) blue, bluish grey or purple brown; (1) white.
30. Placement of the longest macrochaetae: (0) Abd. VI (Fig.
31. Cryptopygy: (0) present (Fig.
Mouthparts
32. Labial chaeta f: (0) present (Fig.
33. Shape of labial apex: (0) truncate (Fig.
34. Number of prelabral chaetae: (0) 4 chaetae present (Fig.
35. Number of labral chaetae: (0) 6 chaetae present (Fig.
36. Number of apical labral chaetae: (0) 4 chaetae present (Fig.
Thorax
37. Tubercles Di on the first tergite: (0) absent; (1) present (Fig.
38. Tubercles Di and De on the first tergite: (0) separate (Fig.
39. Position of chaetae Di3 on the second and third tergites: (0) free (Fig.
40. Number of ordinary chaetae De on the second tergite (on half of tergite): (0) 3 chaetae present (Fig.
41. Position of chaeta De2 on the second tergite: (0) free (Fig.
42. Number of ordinary chaetae De on the third tergite (on half of tergite): (0) 4 chaetae present (Fig.
43. Position of chaeta De2 on the third tergite: (0) free (Fig.
44. Presence of chaeta De3 on the third tergite: (0) present (Fig.
45. Position of chaeta De3 on the third tergite: (0) free (Fig.
Abdomen dorsal
46. Number of ordinary chaetae De on tergites I–III: (on half of tergite): (0) 3 chaetae present (Fig.
47. Position of chaeta De2 on tergites I–III: (0) free (Fig.
48. Presence of chaeta De3 on tergites I–III: (0) present (Fig.
49. Position of chaeta De3 on tergites I–III: (0) free (Fig.
50. Line of chaetae De1 and s-chaeta on each of tergites I–III: (0) perpendicular to midline (Figs
51. Number of tubercles on tergite IV (excluding tubercles L): (0) 5 or fewer tubercles present (Fig.
52. Length of chaeta Di1 on tergite III: (0) longer than chaeta Di1 on tergite V (Fig.
53. Length of chaeta Di1 on tergite IV: (0) longer than chaeta Di1 on tergite V (Fig.
54. Number of tubercles on tergite V: (0) 4 tubercles present (Fig.
55. Number of chaetae Di on tergite V (on half of tergite): (0) 3 chaetae present (Fig.
56. Length of chaeta Di2 on tergite V: (0) less than half the length of chaeta Di1 on tergite V (Fig.
57. Presence of chaeta Di3 on tergite V: (0) present (Fig.
58. Length of chaeta Di3 on tergite V: (0) more than twice shorter than chaeta Di2 on tergite V (Fig.
59. Shape of tubercles on two last segments: (0) normal (Fig.
Abdomen ventral
60. Male ventral organ (secretory structure composed of modified chaetae,
61. Microchaetae on furcal remnant: (0) present (Fig.
62. Number of chaetae L on segment III: (0) more than 3 chaetae present (Fig.
63. Number of chaetae L on segment IV: (0) more than 5 chaetae present (Fig.
64. Position of chaetae L on segment IV: (0) at least 1 chaeta free (Fig.
65. Chaeta L’: (0) present (Fig.
66. Number of chaetae Vei: (0) 6 chaetae (Fig.
Legs
67. Tooth on claw: (0) present (Fig.
68. Clavate chaetae B4 and B5 on tibiotarsus: (0) present (Fig.
Antennae
69. Number of chaetae on antennal segment I: (0) 8 chaetae (Fig.
70. Number of chaetae on antennal segment II: (0) 12 chaetae (Fig.
Phylogenetic analysis with TNT under an equal weighting scheme produced nineteen most parsimonious trees, with a length of 310 steps, consistency index of 36, and a retention index of 69.
Implied character weighting analyses resulted in a single most parsimonious cladogram with k = 9.687500 (Best score: 14.85760, tree length: 319, total fit: 49.14240) (Fig.
Single cladogram obtained in the analysis of morphology under implied weights k = 9.687500 (Length = 319; Fit = 49.14240). Node values indicate the frequency of GC groups derived from Bremer support (above) and symmetric resampling (below). The main clades are indicated with capital letters (A–H) on branches.
We use the most parsimonious tree obtained under implied weighting as a hypothetical reconstruction to discuss character transformations and relationships within the ingroup (Fig.
Unambiguous morphological character optimization obtained from analysis of the data (Table 1) under implied weights. The numbers above and below circles on the branches indicate character numbers and states, respectively. White and black circles represent homoplasious and nonhomoplasious character state transformations, respectively.
In the analyses, the species of Endonura have been classified into two primary clades (A and D). Both contains several different subclades (B–C and E–H respectively). (Fig.
Clade A includes nineteen species, and it is supported by four character state but none of them represent synapomorphy (Fig.
Subclade B includes 8 species and is supported only by one character: the labial apex ogival in shape (character 33:2, ambiguous). The remaining species form subclade C and this grouping is supported by the presence of microchaetae on furcal remnant (character 61:0, ambiguous).
Clade D comprises twenty eight remaining species, and it is supported by three character state: the head chaeta ‘O’ present (character 18:0, ambiguous), the presence of tubercle ‘Di’ on first thoracic tergite (character 37:1, ambiguous) and by the line of chaetae De1 and s-chaeta on abdominal tergites I–III parallel to midline (character 50:1, ambiguous).
The phylogenetic position and affiliation of the two species, Endonura colorata and E. crimica within this clade are obscure, despite their basal positions. It is uncertain whether they belong to clade D. These species exhibit the position of ‘De3’ chaetae on the third tergite of the thorax (character 45) and of ‘De1’ chaetae on thoracic tergites I–II (character 50), similar to taxa of this clade. However, they also share the same position of ‘De2’ chaetae on the thorax (characters 41 and 43) as species belonging to clade A. The Endonura agnieskae is placed in the basal position to the remaining species of this clade, but it shares certain characters with species of clade A, as do the two species mentioned above and this grouping has very low support.
The remaining species constitute a distinct group (clade E), which is characterized by the one character: thoracic chaeta ‘De2’ included in tubercle ‘De’ (character 43:1, ambiguous). This group forms 3 separate subclades. The species of subclade F are supported by the following characters: the absence of tubercles ‘Di’ on first thoracic tergite (character 37:0) and the chaeta ‘Di2’ on fifth abdominal tergite shorter than chaeta ‘Di1’ (character 56:1). This group comprises two newly described species. Endonura duplex was clustered with E. reticulata, while E. cochlearifera was placed as a similar species to E. dentifera and E. lusatica.
Subclade G includes ten species, and it is supported by one character state: the presence of chaeta L’ on ventral abdominal sternites (character 65: 0).
Subclade H comprises 10 species that share two common characters: the presence of 5 chaetae ‘Dl’ on head (character 14:1) and the absence of chaeta ‘Dl3’ on the head (character 15:1). The group comprises a newly described species, E. armeniaca, which has been clustered with E. schwendingeri.
In terms of species richness, Endonura is the third most abundant genus in the subfamily. Only two other genera, Deutonura Cassagnau, 1979 and Pronura Dellamare Deboutteville, 1953 contain more taxa, 62 and 58 species, respectively (
It is common taxonomic practice in large genera to create informal groups of species within them, as opposed to subdividing them into subgenera. The formation and naming of these groups are usually on the basis of both geographical and morphological features. Deutonura, the genus referred to above, has five such groups, fodinarum, alticola, centralis, conjuncta and phlegrea (
The genus Endonura has not yet been phylogenetically analysed. The obtained results indicate Endonura to be non-homogeneous genus and all considered species can be grouped into two major clades, A and D (Fig.
In most Collembola groups, including the subfamily Neanurinae to which the analysed genus belongs, chaetotaxy is the main source of characters used in taxonomic and phylogenetic analyses at different levels of classification, as demonstrated by several studies (e.g.
However, chaetotaxy is identified by some authors (e.g.
Although all Collembola have chaetae, only a few, such as the Neanurinae, are characterised by the presence of cuticular protrusions known as tubercles. These tubercles rank second in importance as a source of characters in our analysis. However, the majority of the characters analysed are homoplasies. The only character representing a synapomorphy specific to the E. tatricola, E. dudichi and E. incolorata group is the ‘L4’ chaeta within the ‘So+L’ tubercle.
The insufficient support for the various clades can be attributed to the limited number of morphological characters available for phylogenetic analysis. This problem is prevalent in both the taxonomy of Neanurinae and Collembola as a whole. This issue is unsurprising because springtails are relatively small organisms with a restricted number of morphological characters. The increase in recently described species results in an insufficient number of characters that can be used in taxonomy to identify species and in phylogenetic analyses to infer relationships. In the systematics of springtails, it is now prevalent to use a combination of multiple characters to identify taxa. While this approach remains effective in systematics, it poses significant challenges in phylogenetic analyses. Such an approach results in a deficiency of defining synapomorphies for taxa. Consequently, the support for cladistic analyses is weakened. As a result, the systematics of Collembola (including Neanuridae) is based on the Linnaean system rather than the phylogenetic relationships. The analysis that is presented in this paper is also subject to the same issues. The analysis divides the species of Endonura into distinct groups, however, it is challenging to determine the extent to which this is a result of relationships or similarities among species.
Further research is needed to clarify the relationships among Endonura species, especially through the use of a combination of morphological characters and molecular methods.
Funding statement: The authors have no funding to report.
Conflict of Interest: The authors have no conflicts of interests to declare.Authors’ contributions: A.S. was responsible for the identification of the species, the preparation of the taxonomic section of the paper, interpretation of data and the revision of the manuscript. N.K. collected and determined material for the study and contributed to the present research’s conception and design. G.P. conducted the phylogenetic analyses with constructive discussions and made suggestions and revisions to the manuscript. All authors read and approved the final manuscript.
We wish to express our sincere thanks to Collectors: Boris Efeikin, Dmitry Shitikov, Alexander Sharikov, Mikhail Potapov (Moscow), Alexander Kremenitsa (Essentuki). Ivan Kuchiev (Alagir).