European Lepidocyrtus lignorum-group, new findings and redescritpion of Lepidocyrtus pulchellus Denis, 1926 (Collembola, Entomobryidae)

Abstract

Some European species of the genus Lepidocyrtus were described almost a century ago and are only known from the type locality. This is the case of the Italian species L. pulchellus Denis, 1926, whose original description is very brief and does not allow its proper identification according to the current standards of the genus taxonomy. The study of the specimens of the type series, as well as other fresh specimens collected in several Italian locations, has allowed us to redescribe the species and expand its distributional range.

Full text

https://doi.org/10.11646/zootaxa.4790.3.10
http://zoobank.org/urn:lsid:zoobank.org:pub:164CFD6B-8CB9-41AC-9FBC-20C5861A0AAB
564 Accepted by W.M. Weiner: 21 May 2020; published: 12 Jun. 2020
Article ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Zootaxa 4790 (3): 564–576
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Copyright © 2020 Magnolia Press
European Lepidocyrtus lignorum-group, new findings and redescritpion of
Lepidocyrtus pulchellus Denis, 1926 (Collembola, Entomobryidae)
EDUARDO MATEOS
Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals; Facultat de Biologia; Universitat de Barcelona; Avinguda Diago-
nal 643, 08028—Barcelona; Spain.
�
emateos@ub.edu; https://orcid.org/0000-0001-9741-5744
Abstract
Some European species of the genus Lepidocyrtus were described almost a century ago and are only known from the type
locality. This is the case of the Italian species L. pulchellus Denis, 1926, whose original description is very brief and does
not allow its proper identification according to the current standards of the genus taxonomy. The study of the specimens
of the type series, as well as other fresh specimens collected in several Italian locations, has allowed us to redescribe the
species and expand its distributional range.
Key words: Lepidocyrtinae, chaetotaxy
Introduction
Lepidocyrtus Bourlet, 1839 is one of the most species-rich and widespread genera of Collembola, and up to the pres-
ent a total of 35 species, distributed in five species-groups, have been described in Europe (see Mateos et al. 2018).
The lignorum-group (sensu Mateos 2011) is the group that contains the largest number of described species, some
of them with a wide distribution on the continent, while others are only known of their type locality. Mateos (2008),
in a study of European Lepidocyrtus Bourlet, 1839 stated that several species were poorly known and difficult to
include in the identification keys due to a lack of diagnostic characters beyond body pigmentation. They are histori-
cal descriptions that lack many characters currently used for species diagnosis. One of this species was Lepidocyrtus
pulchellus Denis, 1926 of which only the citation of the original description is known.
Denis (1926) described the species based on two specimens from Campigna (Italy), one of them incomplete.
This original description was only a short diagnosis (without any image), and the author announced that the full
description would appear in a future publication. Denis (1927) “formally” described the species (based on the same
two specimens from Campigna) offering descriptions and drawings of body pigmentation, unguis morphology and
apical end of dentes and mucro. Denis (1941) indicates that L. pulchellus could be a synonym for L. ruber Schött,
1902. This synonymy, although with doubts, was accepted by Gisin (1960) and Salmon (1964) and that is why in the
catalog of Italian springtails Dallai et al. (1995) cited L. ruber and not L. pulchellus. Until now no more references
of this species have been published.
The study of the L. pulchellus specimens deposited at the National Museum of Natural History (NMNH) of
Paris (France), as well as new specimens found in two Italian locations, will allow us to expand our knowledge of
this species.
Materials and methods
In a visit to the NMNH on January 2019, the author revised three slides labelled as Lepidocyrtus pulchellus iden-
tified by J.R. Denis, two of them from the type locality (Campigna, Italy), and the third one from another Italian
locality (Bertinoro) (Fig. 1, localities 1 and 2). The four specimens contained in these three slides were studied under

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phase contrast microscope. All four specimens were in very bad conditions and only body pigmentation, mucro,
unguis and unguiculi morphology were visible.
FIGURE 1. Lepidocyrtus pulchellus, map of Italy showing the four localities were the species has been found. 1, Campigna
(Type locality); 2, Bertinoro; 3, Viterbo; 4, Rapallo.

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Studying Lepidocyrtus specimens from samples collected in two Italian localities in 2014 and 2015 (Fig. 1,
localities 3 and 4), 18 specimens appeared with the body pigmentation and unguiculi morphology as described for
L. pulchellus by Denis (1926, 1927). The complete chaetotaxy and morphology of seven specimens were studied in
detail. For this morphological study, the specimens were cleared using Nesbitt fluid and then slide-mounted, with
the head separated from the body, in Hoyer medium. The slides were studied under a phase contrast microscope
with a digital camera attached.
For the taxonomic descriptions the following terms and codes were used: For dorsal cephalic chaetotaxy the
“AMS” nomenclature system (see Soto-Adames 2010). For labial palp Fjellberg (1999). For labial chaetotaxy Gisin
(1964). For postlabial chaetotaxy Soto-Adames (2010). For interocular chaetotaxy Mari-Mutt (1986). For dorsal
chaetotaxy schemes of thoracic and abdominal segments Gisin (1967), Szeptycki (1972, 1979), Wang et al. 2003
and Mateos (2008). For tergal specialized chaetae (S-chaetae) Zhang & Deharveng (2015).
Abbreviations and symbols in text and figures: ant.—antennal segment, th.—thoracic segment, abd.—ab-
dominal segment, ungui. —unguiculi, I–VI—segment numbers, psp—pseudopore.
Taxonomic section
Family Entomobryidae Schäffer, 1896
Genus Lepidocyrtus Bourlet, 1839
Lepidocyrtus pulchellus Denis, 1926
Figs 1–17, Table 1
Material examined. Two specimens (of indeterminate sex) on two slides from the type locality (Campigna, Italy,
1000 m above sea level) deposited at the MNHN Paris (France), J.R. Denis det., M.P. Zangheri leg., 01.vii.1924.
Two specimens (of indeterminate sex) on one slide from Bertinoro (Italy, 250 m a.s.l.) deposited at the MNHN Paris
(France), J.R. Denis det., M.P. Zangheri leg, 31.x.1926. One male on slide from Monte Cimino, Viterbo municipal-
ity, Lazio province (Italy, 1014 m a.s.l.), position N42°24’33.9” E12°12’14.3”, E. Mateos leg., 04.v.2014, sample
code LP521. Six specimens (4 females and 2 with indeterminate sex) on six slides and eleven specimens on absolute
alcohol form Rapallo, Varese Ligure municipality, La Spezia province (Italy, 680 m a.s.l.), position N44°22’48.53
E9°30’40.63”, B. Zhang leg., 04.v.2015, sample code LP389.
Diagnosis. Body colour yellow with a characteristic dark blue design: A wide transverse dark blue band in abd.
II, abd.III, and most of abd.IV. Th.II slightly projecting over head. Ant.I–III, legs, ventral tube and posterior region
of manubrium with scales. Apical bulb on ant.IV absent. Labial chaetae M1M2REL1L2 in “p row” well developed
and ciliated, R shortened. Dorsal cephalic and body macrochaetae formula as A0[A2a]A2A3Pa5/00/0101+3. Without
chaeta s on abd.IV. Unguiculi truncate, with serrate outer edge.
Redescription. Body length 1.5–1.7 mm. Specimens from the slides of MNHN and all new collected speci-
mens showed a very characteristic colour pattern (Fig. 2): a wide dark blue (almost black) transverse band in abd.
II–abd.III and most of abd.IV, tips of ant.II–IV and coxae. Ocular areas dark pigmented. Th.II slightly projecting
over the head. Scales densely covering dorsal and ventral surfaces of head, trunk, legs, ventral tube and manubrium,
dorsal surface of ant.I–III, and anterior surface of dens.
Antennal length to head diagonal length ratio (head diagonal measured from cervical edge to apex of mouth
part) 1.6–1.9. Relation of antennal joints I–IV as 1:1.9:1.8:2.8. Ant.I with three dorsolateral small acute chaetae in
a triangle (ant.I-organ sensu Hüther, 1986). Ant.III sense organ composed of two bent sensory rods partially behind
a cuticular fold (Fig. 3). Ant. IV without apical bulb.
Arrangement of chaetae on labrum 4/554, prelabral chaetae ciliated, first and second row of labral chaetae
smooth, apical row bi-branched (detail in Fig. 4). Labrum intrusion inverted U-shaped, labral edge with four minute
rounded labral papillae with two/three-pointed end (Fig. 4). Labial palp with lateral process on papilla E slightly
curved, with pointed apical end and not reaching apex of papilla (Fig. 5). Outer maxillary palp with two smooth
chaetae and three smooth sublobal chaetae (Fig. 6).

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FIGURE 2. Lepidocyrtus pulchellus, habitus lateral. A and B specimens from the type locality (Campigna, see « Material ex-
amined »); C and D specimens from Bertinoro (see « Material examined »); E specimen from Viterbo (sample LP521), scale
bar = 1 mm.
Labium chaetotaxy formed by 5 smooth chaetae (a1–a5) in anterior row; basal row with ciliated chaetae
M1M2REL1L2, with R smaller than other chaetae (ratio of R/M≈0.4), some specimens with three M chaetae on one
side of the labium. Postlabial chaetotaxy with 4+4 ciliated chaetae along ventral cephalic grove (Fig. 7).
Dorsal cephalic macrochaetae formula A0A2A3Pa5, but also with pair of smaller supplementary macrochaetae
A2a between A0 and A2; maximum number of macrochaetae An on head 13+13. Eye patches dark blue. Diameters
of eyes A–F about the same. Eyes G and H slightly smaller (A:G; A:H ≈ 1.6). Interocular chaetotaxy with s, t, p
chaetae and 3–4 intraocular scales (Fig. 8).
Dorsal body macrochaetae formula 00/0101+3 (macrochaetae m3 on abd.II, and C1+B4, B5, B6 on abd.IV).
Dorsal chaetotaxy of th.II–III and abd.I as in Figs 9–11. Th.II with 2 lateral S-chaetae (al and ms) and without mac-
rochaetae in dorsal position. Th.III with a lateral sensillum (al) close to several ciliated chaetae. Abd.I with a lateral
S-microchaeta (ms) external to a6. Chaetotaxy of abd.II–III as in Figs 12–13. Abd.II without chaeta a2p; chaeta ml
(associated with trichobothrium m2) absent in several specimens; macrochaeta m3 with socket diameter 1.5 times
higher than macrochaeta m5. Abd.III with S-chaetae as and ms, and chaeta d3 between macrochaetae pm6 and p6;
two specimens from sample LP389 with lateral tuft of 10+10 long ciliated filiform chaetae. Chaetae associated with
trichobotria on abd.II–III pointed, thickened in the center and ciliated, only chaeta am6 on abd.III has fan-shaped
morphology. Chaetotaxy of abd.IV as in Fig. 14; macrochaetae B4, B5, B6, C1, D3, De3, E2, E3, E4, F1, F2 and
F3 broader with broad socket, while D2, D3p, E1, E4p, E4p2, F3p, Fe4, Fe5, r3, T6 and T7 thinner with smaller

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socket; macrochaeta F2 above macrochaeta E3; the ratio of distances between macrochaetae C1–B4/B4–B6 is
0.68–0.98; the ratio of distances between macrochaetae B4–B5/B5–B6 is ≈ 1; accessory chaeta s associated with
trichobotrium T2 absent; chaetae a, m, pe, pi associated with trichobotria T2 and T4 fan-shaped; chaeta D1 pointed,
thickened in the center and ciliated; sens chaetotaxy composed by 3 anterior dorsomedial elongate S-chaetae, and
short chaetae as and ps. Dorsal chaetotaxy of abd.V with S-chaetae as, acc.p4 and acc.p5 (Fig. 15).
Trochanteral organ with a maximum of 30 smooth spiny chaetae forming a rectangular shape pattern. Ungues
(Fig. 16) with paired basal teeth at 45–50% from inner edge, one median tooth at 64–68%, and a tiny apical tooth at
83–87% from inner edge, respectively; one external tooth and a pair of lateral teeth also present; ungui.I–III truncate
and with serrate outer edge; tibiotarsal tenent hair spatulate, smooth and as long as claw; ratio of supraempodial
chaeta (smooth chaeta on tibiotarsus III opposite to tenent hair) / unguiculus is ≈ 0.8.
Ventral tube with 5+5 ciliated chaetae on anterior side and 12+12 weakly ciliated chaetae on posterior side;
lateral flap with 17 laterodistal chaetae (13 ciliated and 4 smooth).
Manubrium with 2+2 ciliated apical chaetae on anterior side; manubrial plate with 3–4 inner chaetae and 8–11
chaetae outer to the 2 pseudopores. Dental tubercle absent. Mucro without spinelet on basal spine. Ratio manu-
brium/dens/mucro as 15:18:1.
Ecology and distribution. The only information provided by Denis (1926, 1927) was that the specimens were
collected from the soil at Campigna locality and at an altitude of 1000 m above sea level. The specimens deposited
at the NMNH (Paris) do not have information related to the collection site, and the only information available is the
altitude (250 m) at which Bertinoro municipality is located. Specimens from Rapallo locality (680 m altitude) were
collected form oak forest litter layer, and specimens from Viterbo locality (1014 m altitude) were collected from
beech forest litter layer.
FIGURES 3–7. Lepidocyrtus pulchellus. 3, ant.III apical organ; 4, labrum; 5, outer labial papilla; 6, maxillary palp; 7, labial
and postlabial chaetotaxy.

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FIGURE 8. Lepidocyrtus pulchellus, dorsal head chaetotaxy (left side).
Discussion
Currently, the L. lignorum-group is composed of 13 species, namely L. barbulus Mateos, 2011, L. chorus Mateos &
Lukić, 2019, L. instratus Handschin, 1924, L. intermedius Mateos, Escuer & Álvarez-Presas, 2018 (in Mateos et al.
2018), L. juliae Mateos, 2011, L. lignorum (Fabricius, 1793), L. peisonis Traser & Christian, 1992, L. ruber Schött,
1902, L. tellecheae Arbea & Jordana, 1990, L. traseri Winkler, 2016, L. uzeli Rusek, 1985, L. vexillosus Loksa &
Bogojević, 1967 and L. violaceus Lubbock, 1873. All of them have trunk macrochaetotaxy formula 00/0101+3. The
dorsal head macrochaetotaxy formula A0A2A3Pa5 is also shared by all species of the group except L. intermedius
(A0A2Pa5), L. ruber (A0A2A3) and L. vexillosus (A0A2Pa5).
Morphological characters clearly assign Lepidocyrtus pulchellus to the Lepidocyrtus lignorum-group. By the
characteristic body colour pattern L. pulchellus clearly differs from all the other species of the L. lignorum-group
(Fig. 17). Other differences between all 14 species included in the group are summarized in Table 1.

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TABLE 1. Lepidocyrtus lignorum-group, characters differentiating between the species. Body colour: bl––body blue, bl band––body with blue bands, bl dots––body with blue
dots, bl part––body partially blue, wh––body white (without blue pigment), bl/wh––body blue or white; Ant.III scales: presence (+) or absence (-) of scales on ant.III; Head A3:
presence (+) or absence (-) of macrochaeta A3 on dorsal head; Head Pa5: presence (+) or absence (-) of macrochaeta Pa5 on dorsal head; Ocular q: presence (+) or absence (-)
of interocular chaeta q; Labial M: number of chaetae M on labial chaetotaxy; Labial R: number of chaetae R on labial chaetotaxy; Labrum apical: shape of the chaetae on the
apical row of the labrum, poin––apically pointed, bran––apically branched, ?––no published information; Labrum papilla: shape and number of denticles of the labrum apical
papillae, blunt––papillae with blunt denticles, poi––papillae with pointed denticles; smooth––papillae without denticles, ?––no published information; Abd.III tuft: presence (+)
or absence (-) of long filaments tuft of lateral abd.III (for L. pulchellus a “?” is added indicating that this tuft is only present in two specimens); Abd.IV T6: morphology of abd.
IV macrochaeta T6, B––broad ciliated macrochaeta, T–– thin ciliated macrochaeta; Abd.IV B6: morphology of abd.IV macrochaeta B6, B––broad ciliated macrochaeta, T–– thin
ciliated macrochaeta; BP4 psp: presence (+) or absence (-) of pseudopores on the basal plate of abd.IV; Unguis basal: location of the basal pair of denticles on the inner edge of
the unguis (measured in percentage of the inner edge longitude); Unguis unp: number of unpaired denticles on the inner edge of the unguis; Unguiculus: shape of unguilucus and
presence/absence of denticles on the outer edge, lan–– unguiculus lanceolate, tru––unguiculus truncate, (+)––outer edge serrate, (-)––outer edge smooth.
barbulus
chorus
instratus
intermedius
juliae
lignorum
peisonis
pulchellus
ruber
tellecheae
traseri
uzeli
vexillosus
violaceus
Body colour wh/bl bl dots bl part wh dots wh wh bl band wh/bl wh bl part bl bl dots bl
Ant.III scales + - - - - - +/- + - + - - ? -
Head A3 + + + - + + + + + + + + - +
Head Pa5 + + + + + + + + - + + + + +
Ocular q - - ? - + - - - ? - - ? ? -
Labial M 3–4 2 2 2 2 2 2 2–3 2 2 2 2 2 2
Labial R 2–5 1 1 1 1 1 1 1 1 1 1 1 1 1
Labrum apical poin bran ? bran bran bran poin bran poin poin po/br poin ? bran
Labrum papilla blunt 4 poi 1–3 ? poi 3 poi 3 poi 3 smooth poi 2–3 smooth poi 1 poi 1 poi 1 ? poi 1–3
Abd.III tuft - - - - - - + +? - - - - - -
Abd.IV T6 T B T T T T T T T T T T T T
Abd.IV B6 B B B B B B B B B B T B B B/T
BP4 psp - - - - + - - - - - - - - -
Unguis basal 50% 54% 40% 56% 52% ±50% 54% 45–50% 60% 50% 52% 50% 60% ±50%
Unguis unp 2 2 1 2 2 1–2 2 2 1 1 2 0 1–2 2
Unguiculus lan (+) lan (+) lan (-) lan (-) lan (+) lan (-) tru (-) tru (+) tru - lan (+) lan (-) tru (-) lan (-) lan (+/-)

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FIGURES 9–11. Lepidocyrtus pulchellus. 9, th.II dorsal chaetotaxy (left side), psp—pseudoporus, circles—ciliated chaetae; 10,
th.III dorsal chaetotaxy (left side), psp—pseudoporus; 11, abd.I dorsal chaetotaxy (left side), psp—pseudoporus.

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The lateral tuft of filiform chaetae on abd.III described in this paper for L. pulchellus (composed of a maximum
of 10+10 filaments) was only present on two studied specimens and, for this reason, cannot be considered as a
characteristic of the species. The lateral tuft of filaments on abd.III has been described for several species of non-
European subgenera Setogaster Salmon, 1951 and Cinctocyrtus Yoshii & Suhardjono, 1989, and for the European
species L. peisonis Traser & Christian, 1992 (see Winkler & Mateos 2018). With this latter species L. pulchellus also
shares the truncate morphology of the unguiculi, but it is serrate in L. pulchellus instead of smooth as in L. peisonis.
It is quite possible that some other European species of Lepidocyrtus have a long, thin and ciliated lateral tuft of
chaetae (filaments?) on abd.III, but this is something that has yet to be studied.
Denis (1927) noted that Lepidocyrtus cinctus Schäffer, 1898 (from New Britain, Solomon Islands) is similar to
L. pulchellus in unguis morphology and body pigmentation. But L. cinctus, now considered a member of subgenus
Setogaster Salmon, 1951, has several characters that clearly differ from L. pulchellus, as for example unscaled an-
tennae, legs and ventral tube, and the presence of spinelet on the basal mucronal spine (see Mateos & Greenslade
2015).
FIGURES 12–13. Lepidocyrtus pulchellus. 12, abd.II chaetotaxy (left side), black circles—broad ciliated macrochaetae, psp—
pseudoporus; 13, abd.III chaetotaxy (left side), broad black circles—broad ciliated macrochaetae, small black circles—thin
ciliated macrochaetae, psp—pseudoporus.

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FIGURE 14. Lepidocyrtus pulchellus, abd.IV chaetotaxy, broad black circles—broad ciliated macrochaetae, small black cir-
cles—thin ciliated macrochaetae, psp—pseudoporus.

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FIGURES 15–16. Lepidocyrtus pulchellus. 15, abd.V chaetotaxy, broad black circles—long ciliated macrochaetae, small black
circles—short ciliated macrochaetae or smooth mesochaetae; 16, third leg apex with unguis, unguiculus and tenent hair.
FIGURE 17. Lepidocyrtus lignorum-group, lateral habitus of all European species currently included in the group. On the left
side of the figure, species without body pigmentation are represented, in the center species with pigment forming dots or bands, on
the right side species with pigment spread on the thorax and abdomen (above—less pigmented species, below—more pigmented
species). Image of L. vexillosus extracted from http://v3.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxid=398041. Im-
age of L. instratus extracted from Handschin (1924). The image of L. pulchellus correspond to one specimen of sample LP389
(from Rapallo, Italy).

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Acknowledgements
I would like to thank Dr. Cyrille D’Haese, form the NMNH (Paris, Fance), for making available to the author the
specimens of L. pulchellus deposited at the NMNH collections, and allowing the use of his laboratory during my
stay at the Museum. Also I would like to thank Dr. Bing Zhang for providing the specimens from the locality of
Rapallo (Italy), and Dr. Daniel Winkler, Dr. Javier Arbea, and Dr. Wanda Maria Weiner for their valuable comments
and suggestions to the text.
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