2urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7Eurn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727AZooKeysZK1313-29891313-2970Pensoft Publishers10.3897/zookeys.737.2119121191Research ArticleAnimaliaCollembolaNeanuridaeSystematicsCenozoicAsiaEuropeThe genera Albanura Deharveng, 1982 and Persanura Mayvan et al., 2015 are no longer monotypic: description of new species from the Caucasus (Collembola, Neanuridae, Neanurinae, Neanurini)SmolisAdrianadrian.smolis@uwr.edu.pl1KuznetsovaNataliyahttps://orcid.org/0000-0002-5042-91932Institute of Environmental Biology, Department of Invertebrate Biology, Evolution and Conservation, University of Wrocław, Przybyszewskiego 65, 51-148 Wrocław, PolandUniversity of WrocławWrocławPolandInstitute of Biology and Chemistry, Moscow State Pedagogical University, Moscow 129164, RussiaMoscow State Pedagogical UniversityMoscowRussia
Corresponding author: Adrian Smolis (adrian.smolis@uwr.edu.pl)
Academic editor: W.M. Weiner
201812022018737112FFB7EA07-FFD2-ED49-FFD0-FFE6FF915B04E5A60FD4-289B-4B2D-86B1-C7BB20F0FE7812222902509201716012018Adrian Smolis, Nataliya KuznetsovaThis 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.http://zoobank.org/E5A60FD4-289B-4B2D-86B1-C7BB20F0FE78
Two new species from the Caucasus belonging to the genera Albanura and Persanura are described and illustrated in detail. Albanurasecundasp. n. is distinctive because of the presence of chaetae E on the head as well as three ordinary chaetae on tubercles De of thorax III and abdomen I–III. Additionally, the species can be recognized by the absence of chaeta O on the head and presence of 3+3 chaetae Di on abdomen V. The most important characters that can be used to distinguish Persanuralencaranasp. n. are the labral formula, an increased number of chaetae De on thorax II and III, and the number of chaetae Di on the thorax and abdomen V. Comments on the status of the genera and the affinities of the Caucasian fauna of Neanurinae are also given.
AzerbaijanRussiataxonomyCitation
Smolis A, Kuznetsova N (2018) The genera Albanura Deharveng, 1982 and PersanuraMayvan et al., 2015 are no longer monotypic: description of new species from the Caucasus (Collembola, Neanuridae, Neanurinae, Neanurini). ZooKeys 737: 1–12. https://doi.org/10.3897/zookeys.737.21191
Introduction
Creating and establishing new genera, especially monotypic ones, may be very problematic and complicated for taxonomists. Very often such decisions are questioned by other specialists, and authors who decide on such choices are often labelled as ‘splitters’. Interestingly, the term ‘splitter’ was used for the first time by one of the most famous biologists, Charles Robert Darwin. Among Neanurini, one of the six tribes within the subfamily Neanurinae (Cassagnau, 1989), there are both large and “megadiverse” genera, e.g., Deutonura Cassagnau, 1979 and Endonura Cassagnau, 1979 comprising 57 and 51 valid species, respectively (Cassagnau 1979, Smolis and Kaprus’ 2009, Porco et al. 2010, Smolis et al. 2011, Deharveng et al. 2015, Smolis et al. 2016, Smolis and Kuznetsova 2016, Smolis et al. 2017), and several monotypic ones, e.g. Albanura Deharveng, 1982, Cansilianura Dallai & Fanciulli, 1983, Edoughnura Deharveng et al., 2007, Xylanura Smolis, 2011, orPersanura Mayvan et al., 2015 (Deharveng 1982, Dallai and Fanciulli 1983, Deharveng et al. 2007, Smolis 2011, Mayvan et al. 2015).
The examination of rich Neanurinae materials from the Caucasus collected during primarily ecological research has revealed two unknown species that should be classified within the so far monotypic genera Albanura and Persanura. The genus Albanura was created by Deharveng (1982) for Neanura (Deutonura) nana Cassagnau & Péja, 1979, from Albania (Cassagnau and Péja 1979), whilst the genus Persanura was established recently by Mayvan et al. (2015) for the species Persanurahyrcanica Mayvan et al., 2015 from Iran. Illustrated descriptions of the new species with remarks on the status of both corresponding genera and the possible origin of the Neanurinae fauna of the region are provided herewith.
Materials and methods
The specimens were cleared in Nesbitt’s fluid, subsequently mounted on slides in Phoera liquid (200 g of chloral hydrate, 30 g of arabic gum and 20 g of glycerol dissolved and mixed into 50 g of distilled water) and studied using a Nikon Eclipse E600 phase contrast microscope. Figures were drawn with camera lucida and prepared for publication using Adobe Photoshop CS3.
Institutions of depository of materials:
DIBEC Department of Invertebrate Biology, Evolution and Conservation, Institute of Environmental Biology, University of Wrocław, Poland
MSPUMoscow State Pedagogical University, Institute of Biology and Chemistry, Moscow, Russia
Terminology. The terminology and abbreviations used in the paper are those of Deharveng (1983), Deharveng and Weiner (1984), Smolis and Deharveng (2006) and Smolis (2008).
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, Veorve – ventroexternal, Vea – ventroexternoanterior, Vem – ventroexternomedial, Vep – ventroexteroposterior, Vel – ventroexternolateral, Vec – ventroexternocentral, Vei – ventroexternointernal, Viorvi – 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, Mcc – very short macrochaeta, me – mesochaeta, mi – microchaeta, ms – s-microchaeta, Sors – 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.
Holotype: male on slide, Russia, Caucasus,Karachaevo-Cherkessiya, Teberdinsky Reserve, spruce forest, litter, Oct.1978, leg. E. Dobrolyubova (DIBEC). Paratype: female on slide, same data as for holotype.
Etymology.
The name secunda refers to the fact that the new species is the second member of the genus.
Diagnosis.
Habitus similar to that of Albanuranana. Dorsal tubercles present and well developed. Buccal cone long, labrum ogival. Head with chaetae A, B, C, D and E. Chaeta O absent. Tubercles Dl and (L+So) on head with six and nine chaetae respectively. Tubercles De on Th. II and III with three and four chaetae respectively. Tubercles De on Abd. I–III with four chaetae. Tubercles L on Abd. III and IV with four and seven chaetae respectively. Abd. IV and V with seven and three tubercles respectively. Abd. V with 3+3 chaetae Di.
Description.
General. Body length (without antennae): 0.8 mm (holotype), 0.9 mm (paratype). Colour of the body white. 2+2 rather large black eyes, in a typical arrangement for the genus (one anterior and one posterior, Fig. 1).
Albanurasecunda sp. n.: 1 chaetotaxy of head, Th. and Abd. I (holotype), dorsal view 2 chaetotaxy of tubercles Dl and (L+So) on head, ventrolateral view 3 mandible 4 maxilla 5 chaetotaxy and ventral sclerifications of labrum 6 apical bulb, ventral view 7 dorsal chaetotaxy of Ant. III–IV 8 ventral chaetotaxy of Ant. III 9 dorsal chaetotaxy of Abd. IV–VI 10 tubercle L of Abd. IV 11 sensillum of Abd. V 12 chaeta Di1 of Abd. V 13 furca rudimentary.
https://binary.pensoft.net/fig/185675
Chaetal morphology. Dorsal ordinary chaetae of five types: long macrochaetae, short macrochaetae, very short macrochaetae, mesochaetae and microchaetae. Long macrochaetae relatively thick and short, straight, narrowly sheathed, feebly serrated, apically arc-like or rounded (Figs 1, 9, 12). Macrochaetae Mc and Mcc morphologically similar to long macrochaetae, but much shorter. Mesochaetae similar to ventral chaetae, thin, smooth and pointed. Microchaetae similar to mesochaetae, but shorter. S-chaetae of tergites thin, smooth and short, distinctly shorter than nearby macrochaetae (Figs 1, 9, 11).
Antennae. Typical of the genus. Dorsal chaetotaxy of Ant. III–IV as in Fig. 7 and Table 2. S-chaetae of Ant. IV of medium length and moderately thickened (Fig. 7). Apical vesicle distinct and trilobed (Fig. 6). Ventral chaetotaxy of Ant. III–IV as in Fig. 8 and Table 2.
Mouthparts. Buccal cone long with labral sclerifications ogival. Labrum chaetotaxy: 4/2, 4 (Fig. 5). Labium with four basal, three distal and four lateral chaetae, papillae x absent. Maxilla styliform, mandible thin and tridentate (Figs 3, 4).
Dorsal chaetotaxy and tubercles. Chaetotaxy of head as in Figs 1, 2 and Table 1. Chaetotaxy of thorax and abdomen as in Figs 3, 4 and Table 3. Th. III with chaetae De3 free (Fig. 1). On Abd. I–III, the line of chaetae De1–chaeta s is not perpendicular to the dorsomedian line. No cryptopygy, Abd. VI well visible from above (Fig. 9).
Chaetotaxy of Albanurasecunda sp. n.: cephalic chaetotaxy, dorsal side.
Tubercle
Number of chaetae
Types of chaetae
Names of chaetae
Cl
4
Mc
F
Mcc
G
Af
10
Ml
B
Mc
A
Mcc
C, D, E
Oc
3
Ml
Ocm
Mcc
Ocp
mi
Oca
(Di+De)
4
Ml
Di1, De1
Mcc
Di2, De2
Dl
6
Ml
Dl5, Dl1
Mc
Dl3, Dl4
Mcc
Dl2, Dl6
(L+So)
8
Ml
So1
Mc
L1, L2, L4
Mcc
So2, So6
me
So3–5
Chaetotaxy of Albanurasecunda sp. n.: chaetotaxy of antennae.
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 Albanurasecunda sp. n.: 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; Fu: 5 me, 8 mi
Abd. IV
(2+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: 13–14; An: 2mi
Ventral chaetotaxy. On head, groups Vea, Vem and Vep with 3, 3, 4 chaetae respectively. Group Vi on head with 6 chaetae. Group L on Abd. IV with 7 chaetae (Fig. 10). On Abd. IV, furca rudimentary with 8 microchaetae (Fig. 13). On Abd. V, chaetae Vl and L’ present.
Legs. Chaetotaxy of legs as in Table 3. Claw without internal tooth. On tibiotarsus, chaeta M present and chaetae B4 and B5 relatively short and pointed.
Remarks.
Albanurasecunda sp. n is easily distinguished from A.nana Cassagnau & Péja, 1979, the only other known species in the genus, by its dorsal chaetotaxy: presence/absence of chaeta O on the head (absent in secunda; present in nana), presence/absence of chaetae E on the head (present in secunda; absent in nana), number of chaetae Dl on the head (six in secunda; five in nana), number of ordinary chaetae De on Th. III (three in secunda; two in nana), number of ordinary chaetae De on Abd. I–III (three in secunda; two in nana), number of chaetae L on Abd. IV (seven in secunda; five in nana), and number of chaetae Di on Abd. V (3+3 in secunda; 2+2 in nana).
Holotype: female on slide, Azerbaijan, Lankaran District, surroundings of Dashdatuk, forest, under stones, 30.I.1985, leg. Expedition of Moscow Pedagogical State University (MSPU).
Etymology.
The name “lencarana” refers to a region where the new species was found.
Diagnosis.
Habitus similar to that of Persanurahyrcanica. Dorsal tubercles present and well developed. 3+3 pigmented eyes. Buccal cone short and relatively wide, labral sclerifications nonogival. Chaetotaxy of central area of head complete, chaetae A, B, C, D, E and O present. Tubercles Dl and (L+So) on head with six and ten chaetae respectively. Tubercles Di on Th. with two chaetae. Tubercles De on Th. II and III with five and six chaetae respectively. Tubercles De on Abd. I–III with four chaetae. Tubercles L on Abd. III and IV with five and eight chaetae respectively. Tubercles Di on Abd. V with three chaetae.
Description.
General. Body length of holotype (without antennae): 2.15 mm. Colour of the body bluish. 3+3 small black eyes, in a typical arrangement for the genus (Fig. 14).
Persanuralencarana sp. n.: 14 chaetotaxy of head and Th. (holotype), dorsal view 15 chaetotaxy of ventrolateral part of head 16 tubercle De of Abd. II 17 tubercle Dl of Th. II, ventral view 18 mandible 19 maxilla 20 furca rudimentary 21 claw and T III, ventrolateral view 22 sensillum sgv and microsensillum of Ant. III 23 apical bulb, dorsal view 24 apical bulb, ventral view 25 dorsal chaetotaxy of Ant. III–IV 26 chaetotaxy and ventral sclerifications of labrum 27 chaeta Di1 of Abd. IV 28 sensillum of Abd. V.
https://binary.pensoft.net/fig/185676
Chaetal morphology. Dorsal ordinary chaetae of three types: long macrochaetae, short macrochaetae and mesochaetae. Long macrochaetae relatively thin, slightly arc-like or straight, narrowly sheathed, feebly serrated, apically rounded or pointed (Figs 14–17, 27). Macrochaetae Mc morphologically similar to long macrochaetae, but shorter. Mesochaetae similar to ventral chaetae, thin, smooth and pointed. S-chaetae of tergites thin, smooth and short, notably shorter than nearby macrochaetae (Figs 14, 16, 17, 28).
Antennae. Typical of the genus. Dorsal chaetotaxy of Ant. III–IV as in Fig. 25 and Table 5. S-chaetae of Ant. IV relatively long and moderately thickened (Fig. 25). Apical vesicle well developed, trilobed (Figs 23, 24). Ventral chaetotaxy of Ant. III–IV as Table 5, sgv long and s-shaped (Fig. 22).
Mouthparts. Buccal cone short and wide, with labral sclerifications nonogival. Labral chaetotaxy: 4/2, 4 (Fig. 26). Labium with four basal, three distal and four lateral chaetae, papillae x absent. Maxilla styliform, mandible strong with two basal and four apical teeth (Figs 18, 19).
Dorsal chaetotaxy and tubercles. Chaetotaxy of head as in Figs 14, 15 and Table 4. Tubercle Cl with chaetae D. Chaetotaxy of thorax and abdomen as in Figs 14, 16, 17 and Table 6. Thorax and abdomen without free chaetae (Figs 14, 16). On Abd. I–III, the line of chaetae De1–chaeta s perpendicular to the dorsomedian line. Cryptopygy poorly developed, Abd. VI visible from above.
Chaetotaxy of Persanuralencarana sp. n.: cephalic chaetotaxy, dorsal side.
Tubercle
Number of chaetae
Types of chaetae
Names of chaetae
Cl
6
Ml
F
Mc
G, D
Af
8
Ml
A, B
Mc
C, E, O
Oc
3
Ml
Ocm, Ocp
Mc
Oca
Di
2
Ml
Di1
Mc
Di2
De
2
Ml
De1
Mc
De2
Dl
6
Ml
Dl5, Dl1
Mc
Dl2–4, Dl6
(L+So)
10
Ml
L1, L4, So1
Mc
L2, L3, So2, So6
me
So3–5
Chaetotaxy of Persanuralencarana sp. n.: chaetotaxy of antennae.
Segment, Group
Number of chaetae
Segment, Group
Number of chaetae adult
I
8
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 Persanuralencarana sp. n.: postcephalic chaetotaxy.
Terga
Legs
Di
De
Dl
L
Scx2
Cx
Tr
Fe
T
Th. I
2
2
1
-
0
3
6
13
19
Th. II
3
4+s
3+s+ms
3
2
7
6
12
19
Th. III
3
5+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: 6; chaeta Ve1 present
Abd. III
2
3+s
2
5
Vel:5–6; Fu: 5 me, 8 mi
Abd. IV
2
2+s
3
8
Vel: 4–5; Vec: 2; Vei: 2; Vl: 4
Abd. V
3
7+s
Ag: 3; Vl: 1, L‘: 1
Abd. VI
7
Ve: 14; An: 2mi
Ventral chaetotaxy. On head, groups Vea, Vem and Vep with 4, 3, 4 chaetae respectively (Fig. 15). Group Vi on head with 6 chaetae. On Abd. IV, furca rudimentary with 8 microchaetae devoid of visible chaetopores (Fig. 20). On Abd. V, chaetae Vl and L’ present.
Legs. Chaetotaxy of legs as in Table 6. Claw without internal tooth. On tibiotarsus, chaeta M present and chaetae B4 and B5 relatively short and pointed, chaeta A6 similarly in length to chaeta B4 (Fig. 21).
Remarks.
Persanuralencarana sp. n. most visibly differs from P.hyrcanica in the presence of a complete chaetotaxy in the central area of the head (reduced chaetae C, E and O absent in hyrcanica), the presence of two chaetae Di on Th. I (one chaeta in hyrcanica), the presence of three chaetae Di on Th. II–III (two chaetae in hyrcanica), the presence of four and five ordinary chaetae De on Th. II and III, respectively (two chaetae in hyrcanica), the presence of three ordinary chaetae De on Abd. I–III (two chaetae in hyrcanica), and the presence of three chaetae Di on the penultimate abdominal segment (two chaetae in hyrcanica). In addition, they differ in the number of labral chaetae (4/2, 4 in lencarana; 0/0, 4 in hyrcanica), the presence/absence of chaetae Dl3 on the head (present in lencarana; absent in hyrcanica), the number of chaetae L of Abd. IV (8 in lencarana; 3–5 in hyrcanica), and the presence/absence of microchaetae on furca rudimentary (present in lencarana; absent in hyrcanica).
Discussion
The discovery of new species, e.g. Albanurasecunda sp. n. and Persanuralencarana sp. n. described herein, in so far monotypic genera undoubtedly enriches and extends their characteristics providing new facts about their morphological differentiation. In some cases, nevertheless, they may reduce the number of distinguishing characters available for the genera. For example, the number and arrangement of labral chaetae in P.lencarana sp. n. is typical within the tribe and significantly different from the number described in the type species, P.hyrcanica Mayvan et al., 2015, where it was used as a one of the important generic feature of Persanura Mayvan et al., 2015 (Mayvan et al. 2015). On the other hand, strong mandibles, which were not used by the authors of this genus for comparison with other morphologically similar genera, like Kalanura Smolis, 2007, Neanura MacGillivray, 1893, and Xylanura Smolis, 2011 (MacGillivray 1893, Smolis 2007, 2011), appear to be a good and distinctive element of its characteristics. Likewise, A.secunda sp. n. is characterized by 3 + 3 chaetae Di on Abd. V as opposed to 2 + 2 chaetae in the type species of Albanura Deharveng, 1982. Thus, this feature cannot be further used as unique and diagnostic. Besides, new data on the morphological characteristics of originally monobasic genera, provide additional information that may support their recognition, such as geographic location. Interestingly, in the case of Albanura, a taxon morphologically closely related to Deutonura Cassagnau, 1979, but geographically separated, the author of the genus (Deharveng 1982) recognized the geographical criterion as important for its establishment. In the light of the discovery of A.secunda sp. n., the author’s decision seems to be justified as the distributional pattern of the genus is completely different from that which is known in Deutonura. Interestingly none species of Deutonura is known from the Caucasus where other genera of Neanurini, more or less widely distributed in the Western Palearctic, were observed e.g. Endonura, Persanura and Caucasanura Kuznetsova & Potapov, 1988. This last genus up to date includes two species, Caucasanurastebayevae Kuznetsova & Potapov, 1988, known from the Caucasus (Kuznetsova and Potapov 1988) and Caucasanurabesucheti Deharveng, 1989, from north-eastern Turkey (Deharveng 1989).
In addition to supporting taxonomic decisions such as the establishing of monobasic genera, the species described in this work shed light on the origin and composition of the Neanurinae fauna of the Caucasus. The fauna of this subfamily in the region, despite our still poor knowledge, seems to contain elements belonging to the eastern part of Mediterranean Sea and Asia Minor as well as to the mountains in Iran. This assumption is consistent with recent observations on the genus Endonura from the Caucasus, where some of its representatives closely resemble both Mediterranean and Iranian species (Smolis et al. 2016a, b). Undoubtedly, further studies are necessary to understand the diversity and history of the subfamily in this extremely rich natural region, one of the two biodiversity hotspots located in the Western Palaearctic (Myers et al. 2000).
Acknowledgments
We thank Wanda Maria Weiner, Louis Deharveng, and László Dányi for useful comments that improved this manuscript. The work was financially supported by the Russian Foundation for Basic Research (grant N16-04-01228) and the Institute of Environmental Biology, Faculty of Biological Science, University of Wrocław, Poland (project no. 1076/Ś/IBŚ/2017).
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