The European Lepidocyrtus lanuginosus group (Collembola: Entomobryidae), definition and description of a new species from Spain

Abstract

The Lepidocyrtus lanuginosus group is formed by the species L. lanuginosus and L. cyaneus, which can be differentiated by the body color pattern. In the present paper several populations of these two species from the Northeastern Iberian Peninsula have been studied. This study has allowed the detection of chaetotaxic and morphological differences distinguishing the two mentioned species. Also the description of the new species L. bicoloris has been made (also characterized by the body color pattern), as well as a redefinition of the European Lepidocyrtus lanuginosus group.

Full text

Accepted by W.M. Weiner: 25 Oct. 2012; published: 3 Dec. 2012
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Copyright © 2012 · Magnolia Press
Zootaxa 3570: 69–81 (2012)
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Article
69
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The European Lepidocyrtus lanuginosus group (Collembola: Entomobryidae),
definition and description of a new species from Spain
EDUARDO MATEOS
Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona. Avinguda Diagonal, 643, 08028 Barcelona (Spain).
E-mail: emateos@ub.edu
Abstract
The Lepidocyrtus lanuginosus group is formed by the species L. lanuginosus and L. cyaneus, which can be differentiated
by the body color pattern. In the present paper several populations of these two species from the Northeastern Iberian Pen-
insula have been studied. This study has allowed the detection of chaetotaxic and morphological differences distinguishing
the two mentioned species. Also the description of the new species L. bicoloris has been made (also characterized by the
body color pattern), as well as a redefinition of the European Lepidocyrtus lanuginosus group.
Key words: taxonomy, chaetotaxy, body color pattern
Resumen
El grupo Lepidocyrtus lanuginosus está compuesto por las species L. lanuginosus y L. cyaneus, las cuales se diferencian
por el patrón de color del cuerpo. En el presente trabajo se han estudiado varias poblaciones de estas especies procedentes
del Nordeste de la Península Ibérica. Este estudio nos ha permitido detectar las diferencias quetotáxicas y morfológicas
de las dos especies mencionadas. Asimismo se ha realizado la descripción de la nueva especie L. bicoloris (también car-
acterizada por el patrón pigmentario del cuerpo), así como una redefinición del grupo Lepidocyrtus lanuginosus de Euro-
pa.
Palabras clave: taxonomía, quetotaxia, patrones de color corporal
Introduction
L. lanuginosus (Gmelin, 1788) is the oldest species of the genus and, together with L. cyaneus Tullberg, 1871, form
the L. lanuginosus group (sensu Gisin1964b, Hüther 1986). Of these two species there is no complete published
descriptions and only details of the chaetotaxy and some morphological characters can be found (Christiansen &
Bellinger 1980, Fjellberg 2007, Gisin 1963b, 1964a, 1964b, Hüther 1986, Loksa & Bogojevi
1967, Mateos 2008a,
Snider 1967). According to these descriptions the only character that differentiates L. lanuginosus and L. cyaneus is
the color pattern. In this paper we have examined several specimens of Iberian Peninsula populations of both
species in order to determine the existence of more characters (besides the pigmentation) that allow their
differentiation. The study of these L. lanuginosus group populations has allowed to describe a new species
characterized by the body color pattern.
Material and methods
Samples were taken at 10 locations in Northeastern Iberian Peninsula (Table 1). The Lepidocyrtus specimens
collection was performed by hand sorting, except on one occasion in which the specimens were extracted from soil
samples using Berlese-Tullgren devices. A total of 353 specimens of Lepidocyrtus lanuginosus group were
obtained of which 50 were cleared and mounted for observation with phase contrast optical microscope (Table 2).

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For the dorsal macrochaetae and labial chaetotaxy the notation of Gisin (1963a, 1964a, 1964b, 1967), Barra (1975),
and Wang et al. (2003) have been used. For the dorsal chaetotaxy of thoracic and abdominal segments the notation
established by Szeptycki (1979), Snider (1967), Wang et al. (2003), and Mateos (2008a) have been used. The ratio
C1-B4/B4-B6 on abd.IV (i.e. distance between the C1 and B4 setae divided by the distance between the B4 and B6
setae) has been used following Mateos (2008a).
Abbreviations used. The following abbreviations have been used: ant.—antennal segment, abd.—abdominal
segment, cx.—coxae, th.—thorax segment, tr.—trochanter, I–VI—segments, bcm—blunt broad ciliated
macrochaeta, tcm—acuminate thin ciliated macrochaeta.
TABLE 1. Lepidocyrtus lanuginosus group. Localities and collection data. Loc: locality code; Coordinates: lat long
coordinates in decimal degrees format; Mount. Chain: mountain chain of the locality; Habitat: habitat type and specimens
collection methodology: h—hand sorting, b—berlese extraction; Altitude: meters over sea level of the locality.
TABLE 2. Lepidocyrtus lanuginosus group. Specimens and species studied. Loc: locality code; Lot: specimens lot code;
Species: Lla—Lepidocyrtus lanuginosus, Lbi—Lepidocyrtus bicoloris sp. nov., Lcy—Lepidocyrtus cyaneus; Slid / alc:
specimens conservcation method, Slid—slide, alc—absolute alcohol; Date: collection date.
Loc Coordinates Mount. Chain Province Town Habitat Altitude
110 N 41.54550
E 2.35850
Serra Marina Barcelona Orrius Ivy above ground (h) 345
114 N 41.66449
E 2.56279
Serra Montnegre Barcelona Sant Celoni Oak litter (h) 610
115 N 41.65830
E 2.51990
Serra Montnegre Barcelona Vallgorguina Holm and cork oak litter (h) 390
119 N 41.66740
E 2.60080
Serra Montnegre Barcelona Tordera Holm and cork oak litter (h) 390
121 N 41.53797
E 2.38423
Serra Marina Barcelona Cabrera de Mar Pine and oak litter (h) 225
122 N 41.54661
E 2.36532
Serra Marina Barcelona Cabrils Pine and oak litter (h) 425
148 N 42.22890
E 1.93800
Serra Catllaràs Barcelona Pobla de Lillet Holm oak soil (b) 1150
210 N 42.67648
E 0.08542
Ordesa Nat Park Huesca Bielsa Beech litter (h) 1280
202 N 42.68407
W 0.11188
Ordesa Nat Park Huesca Torla Beech and spruce litter (h) 1300
249 N 41.75090
E 2.30553
Serra Montseny Barcelona Tagamanent Herbs above ground (h) 1030
Loc Lot Species Slid / alc Date
110 LP120 Lla 2 / 4 22/02/2007
121 LP146 Lla 1 / 6 05/06/2007
114 LP126 Lla 2 / 48 18/04/2007
115 LP130 Lla 2 / 28 18/04/2007
119 LP137 Lla 2 / 12 18/04/2007
148 LP202 Lla 3 / 27 05/04/2008
122 LP147 Lbi 6 / 32 05/05/2007
122 LP190 Lbi 7 /16 28/11/2007
249 LP295 Lbi 9 /21 15/10/2011
210 LP271 Lcy 10 / 70 30/05/2009
202 LP277 Lcy 6 / 39 31/05/2009

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FIGURES 1–4. Lepidocyrtus lanuginosus group: 1, Lepidocyrtus lanuginosus habitus; 2 Lepidocyrtus lanuginosus dorsal
head; 3, Lepidocyrtus bicoloris sp. nov. habitus; 4, Lepidocyrtus cyaneus habitus.

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Taxonomic section
Family Entomobryidae Schött, 1891
Genus Lepidocyrtus Bourlet, 1839
Lepidocyrtus lanuginosus (Gmelin, 1788)
Figs 1–2, 5–24; Tabs 1–3
Material examined. 12 specimens in slides from six localities located at the “Serra Marina, “Serra del Montnegre
and “Serra del Catllaràs mountains (Barcelona province, Spain) (Tabs 1 and 2). One specimen of the lot LP130
(slide code CRBA10695), and one specimen of the lot LP137 (slide code CRBA10696) saved in the collection of
the Centre de Recursos de Biodiversitat Animal, Faculty of Biology, University of Barcelona (http://
www.crba.ub.edu); other specimens kept in the E. Mateos’ slides collection.
Other material. 125 specimens preserved in absolute alcohol and kept in the E. Mateos’ collection (Tables 1
and 2).
Description. Adult body length (without head and furca) 0.9–1.2 mm. Mesothorax not projecting over the
head. Body dorsoventrally compresed with grey-orange background color. Blue pigment only present on ant.II–IV
(with increasing colour intensity towards the distal part of each segment), and cx.I–III; densely black pigmented
ocular areas (Fig. 1). Some individulas have blue pigment in the mid-dorsal head. The foremost part of the eye-
patches connected by a pigmented band (Fig. 2).
FIGURES 5–10. Lepidocyrtus lanuginosus: 5, ant.I dorsal microchaetae; 6, ant.II distal rod-like sensory seta; 7, ant.III dorsal
view showing ant.III organ; 8, labrum; 9, outer labial papilla; 10, maxillary palp.

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FIGURES 11–12. Lepidocyrtus lanuginosus: 11, labium (left side) and ventral cephalic groove (p/a, seta present or absent
depending on the specimen); 12, scheme of dorsal macrochaetae (circles), trichobothria (lines) and pseudopori (#).

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FIGURES 13–15. Lepidocyrtus lanuginosus: 13, dorsal head chaetotaxy, broad circles—long ciliated macrochaetae, small
circles—short ciliated macrochaetae, lines—smooth mesoch
aetae; 12, interocular chaetotaxy (right ocular area); 15, th.II
complete chaetotaxy (left side), broad circles—broad ciliated macrocha
etae, lines—mesochaetae, pse—pseudoporus.
Antenna without scales. Ratio antenna:cephalic diagonal 1.1–1.7. Ratio ant.I:II:III:IV as 1:2:2:3. Basis of ant.I
dorsally and ventrally with three microchaetae arranged in triangle (Fig. 5). Ant.II dorsally with one distal rod-like
sensory seta (Fig. 6). Ant.III organ composed of two subcylindrical sensory rods partially covered by an
integumentary fold, and with three lateral sensory setae (Fig. 7). Without apical ant.IV bulb.
Ciliated prelabral setae and smooth labral setae in typical number 4/5,5,4; setae of apical row thicker than
those in other rows. Inverted V-shaped labral apical intrusion (Fig. 8). Four rounded smooth labral papillae. Lateral

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process (sensu Fjellberg 1999) of outer labial papilla curved, tip not reaching the apex of the papilla (Fig. 9).
Maxillary palps with two lobal smooth setae and three sublobal smooth setae (Fig 10).
Labial anterior row formed by five smooth setae (a1–a5); posterior row formed by ciliated setae with formula
[M
1
*] M
2
R*EL
1
L
2
; seta M
1
present only in four out of twelve examined individuals, setae R and M
1
half in length
of other setae (marked with *). Ventral cephalic groove with 4+4 ciliated setae (Fig. 11).
FIGURES 16–17. Lepidocyrtus lanuginosus: 16, abd.II complete chaetotaxy (left side), broad circles—broad ciliated
macrocha
etae, lines—mesochaetae, pse—pseudoporus; 17, abd.III complete chaetotaxy (left side), broad circles—broad
ciliated macrocha
etae, small circles—thin ciliated macrochaetae, lines—mesochaetae, pse—pseudoporus.
Dorsal macrochaetae formula such as R
0
R
1s
R
1
R
2
STSo/10/0101+2 (Fig. 12); a pair of supplementary
macrochaetae R
1s
between R
0
and R
1
present. Maximum number of macrochaetae A between ocular areas 9+9 (Fig.
13). Interocular chaetotaxy with ciliated setae (s, t, p), and 2–3 scales (Fig 14). Eyes G and H somewhat smaller
than the other. Th.I chaetotaxy as in Fig. 15; seta p3 ciliated macrochaeta; anterior margin of the tergite bordered
with ciliated macrochaetae. Abd.II–III chaetotaxy as in Figs 16–17; abd.II seta ml absent; abd.III setae ll and d3
absent; abd.III seta p8p smooth mesochaeta. Abd.IV chaetotaxy as in Fig. 18-a; trichobothrium T2 without
accessory seta s; seta D1 ciliated and double in length than setae a and m (Fig. 19); seta E4p ciliated macrochaeta in

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two out of 12 examined individuals, smooth mesochaeta on the other 10 (Fig. 18-b); seta Fe4 smooth mesochaeta;
macrochaeta E3 inserted below F2; ratio C1-B5/B5-B6 as 1.2–1.6; dorsal macrochaetae of two distinct
morphologies: B5, B6, C1, D3, E2, E3, E4, F1, F2, F3 broader and with broad socket (bcm in Fig. 16-b); T6, T7,
D2, De3, E1, E4p (depending on the specimens), Fe5, E4p2 shorter or longer but always thinner and with socket of
minor diameter (tcm in Fig. 18-b).
Ventral tube without scales; 9+9 ciliated setae on anterior side (Fig. 20); 7+7 ciliated setae on posterior side,
and each lateral flap with a maximum of 7 ciliated setae and 2 smooth setae (Fig. 21).
FIGURES 18–19. Lepidocyrtus lanuginosus: 18-a, abd.IV complete chaetotaxy, broad black circles—broad ciliated
macrocha
etae, small black circles—thin ciliated macrochaetae, lines—smooth mesochaetae and microchaetae, triangles—fan-
shaped setae, black square—variable E4p seta, x—trichobothria, #—pseudoporus; 18-b, bcm—blunt broad ciliated
macrochaeta, tcm—acuminate thin ciliated macrochaeta, E4p—two different morphologies of seta E4p; 19, abd.IV
trichobothrial complex.
Legs without scales. V shaped trochanteral organ (leg III) with a maximum of 16 smooth straight setae
arranged in triangular shape (Fig. 22). Unguis with basal pair of teeth at 50% of the inner edge, and with one inner

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EUROPEAN LEPIDOCYRTUS LANUGINOSUS GROUP
teeth at 67% from the base of the inner edge respectively. Unguiculus lanceolate with smooth outer margin.
Spatulate tibiotarsal tenent hair (Fig. 23).
Furca without scales on dorsal surface. Mucro bidentate; mucronal basal spine without spinelet. Ratio
manubrium:dens:mucro as 17:16:1. Manubrial plate with two pseudopora, 2 inner setae, and 3–4
outer setae (Fig.
24).
FIGURES 20–24. Lepidocyrtus lanuginosus: 20, ventral tube, anterior side (right side); 21, ventral tube, postero-lateral with
left lateral flap; 22, trochanteral organ; 23, third leg apex with unguis and unguiculus; 24, manubrial plate (left side),
circles—ciliated macrochaetae, filled circles—pseudopora.
Lepidocyrtus cyaneus Tullberg, 1871
Figs 4, 5–25, 27; Tabs 1–3
Material examined. 16 specimens in slides from two localities located at the Ordesa National Park (Huesca
province, Spain) (Tabs 1 and 2). One specimen of the lot LP271 (slide code CRBA10699), and one specimen of the
lot LP277 (slide code CRBA10700) saved in the collection of the Centre de Recursos de Biodiversitat Animal,
Faculty of Biology, University of Barcelona (http://www.crba.ub.edu); other specimens kept in the E. Mateos’
slides collection.

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Other material. 109 specimens preserved in absolute alcohol and kept in the E. Mateos’ collection (Tables 1
and 2).
Description. Adult body length (without head and furca) 1.0–1.2 mm. Body dark blue pigmented on dorsal
and ventral regions (including ventral tube); head light blue pigmented also on dorsal and ventral regions; blue
pigment also present on ant.II–IV, cx.I-III, tr.I–III, and manubrium (Fig. 4).
All morphological and chaetotaxic characters match those explained for L. lanuginosus, with the following
differences: Ratio antenna:cephalic diagonal 1.2–1.7. Labium chaetotaxy [M
1
*] M
2
R*EL
1
L
2
, M
1
present in 13 out
of 16 examined specimens. Head with a maximum of 10+10 macrochaetae in row A. Abd.II seta ml present on four
out of 16 examined specimens (Fig. 25) (usually this seta is present on one side of the body and absent on the other
side). Abd. IV seta Fe4 ciliated macrochaeta (Fig. 18-a). Ventral tube with a maximum of 8 ciliated setae and 2
smooth setae on lateral flaps. V-shaped trochanteral organ formed by a maximum of 17 smooth setae. Unguiculus
with outer edge smooth (in 13 examined specimens), or finely serrated (in 3 specimens) (Fig.27).
FIGURES 25–27. Lepidocyrtus bicoloris sp. nov. and L. cyaneus: 25, abd.II medial chaetotaxy (left side); 26, third leg unguis
and unguiculus (L. bicoloris); 27, third leg unguis and unguiculus (L. cyaneus).

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EUROPEAN LEPIDOCYRTUS LANUGINOSUS GROUP
Lepidocyrtus bicoloris sp. nov.
Figs 3, 5–26; Tabs 1–3
Type material. Holotype: male in one slide (CRBA10697), Serra Marina mountains, Cabrils municipality,
Barcelona province, Spain (N41.54661 E2.36532), pine and oak litter, 28.xi.2007, leg. E. Mateos (see Tabs 1 and 2,
Loc122). Paratypes: 6 specimens in slides and 16 specimens in alcohol, the same data as the holotype; 6 specimens
in slides and 32 specimens in alcohol, same locality as the holotype, 05.v.2007; 9 specimens in slides and 21
specimens in alcohol, Serra del Montseny mountains, Tagamanent municipality, Barcelona Province, Spain
(N41.75090 E2.30553), herbs above ground (see Tabs 1 and 2, Loc249). Holotype and one paratype slide from
Loc249 (specimen CRBA10698) saved in the collection of the Centre de Recursos de Biodiversitat Animal,
Faculty of Biology, University of Barcelona (http://www.crba.ub.edu); other specimens kept in the E. Mateos’
slides collection.
Etymology. The subspecies name refers to the body color pattern (body with two colors).
Description. Adult body length (without head and furca) 0.7–1.0 mm. With dark blue pigment on the dorsal
and ventral sides of th.II to abd.II (including ventral tube), ant.I–IV (with increasing colour intensity towards the
distal part of each segment), and cx.I–III; densely black pigmented ocular areas (Fig. 3). The foremost part of the
eye-patches connected by a pigmented band.
All morphological and chaetotaxic characters match those explained for L. lanuginosus, with the following
differences: Ratio antenna:cephalic diagonal 1.2–1.6. Labium chaetotaxy [M
1
*] M
2
R*EL
1
L
2
; one out of 22
examined individuals with two M setae on one side of the labium. Interocular chaetotaxy with ciliated setae (s, t,
p), and 1–3 scales. Abd.II seta ml present in three out of 22 examined individuals (Fig. 25) (one specimen with seta
ml present on one side of the body and absent on the other side). Abd.IV seta E4p ciliated macrochaeta in three out
of 22 examined individuals, smooth mesochaeta on the other 19; ratio C1-B5/B5-B6 as 1.3–1.7. Ventral tube with
7+7 ciliated setae on anterior side; 6+6 ciliated setae on posterior side; each lateral flap with a maximum of 6
ciliated setae and 2 smooth setae. Four out of 22 examined individuals have a fourth inner tooth at 83% of the inner
edge of the unguis, and unguiculus with a finely serrated outer margin (Fig. 26). Ratio manubrium:dens:mucro as
16:15:1.
Ecology and distribution. Specimens were obtained from pine and oak litter and from herbaceous vegetation.
They all have a gut content composed mainly of fungal hyphae and spores.
Discussion. Color pattern is usually a useful discriminating character in Lepidocyrtinae (even in
Entomobryidae). The "color pattern species" coined by Yoshii (1989) for tropical Entomobryoidea argues that in
the absence of diagnostic morphological characters, differences in color pattern were sufficient to define valid
species. The molecular analysis of Soto-Adames (2002) on Lepidocyrtus and Pseudosinella also demonstrated this
point. The new species is very close to L. lanuginosus and L. cyaneus. Of this two species L. bicoloris sp. nov. can
be differentiated by the body color pattern: blue pigment on th.II to abd.II in L. bicoloris, without pigment in L.
lanuginosus, and dark blue pigmented in L. cyaneus. In addition, the new species has a size slightly smaller than
the others, abd.IV seta Fe4 is a smooth mesochaeta, and shows variability in the presence or absence of abd.II seta
ml, the number of unguis teeth, and the empodial denticulation (see Table 3).
Definition of the Lepidocyrtus lanuginosus group
With the descriptions made in the present work, the Lepidocyrtus lanuginosus group can be defined as follows:
Small Lepidocyrtus with body dorsoventrally compressed, mesothorax not projecting over the head and body color
background from grey-orange to dark blue. Antennae, legs, ventral tube, and dorsal side of manubrium not scaled.
Labium chaetotaxy [M
1
*]M
2
R*EL
1
L
2
. Dorsal macrochaetae formula R
0
R
1s
R
1
R
2
STSo/10/0101+2. Abd. IV
trichobothrium T2 without accesory seta s; seta D1 ciliated and double in length than other setae of the trichobotrial
complex. The group is composed by the taxa L. lanuginosus, L. cyaneus, and L. bicoloris
sp. nov.

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General discussion
In the examined specimens, besides pigmentation, only one more character has been detected for differentiating L.
lanuginosus, L. bicoloris sp. nov., and L. cyaneus (Table 3): In L. lanuginosus and L. bicoloris sp. nov. seta Fe4 of
abd.IV is always smooth mesochaeta, whereas in L. cyaneus it is always a thin ciliated macrochaeta. This character
is sometimes difficult to observe because, in general, due to the procedure used for performing microscopic slides,
this seta is lost and only the socket can be seen. But this socket is different depending on the type of setae, being of
bigger diameter in ciliated macrochaeta than in smooth mesochaeta.
The examined specimens of L. lanuginosus, L. bicoloris sp. nov. and L. cyaneus show variability in the
presence/absence of the labial seta and M
1
and the morphology of seta E4p of abd.IV (Table 3). Typically labial
chaetotaxy of L. lanuginosus and L. cyaneus is described without seta M
1
(Gisin 1964b, Fjellberg 2007, Mateos
2008a, Winkler & Traser 2012), but according to the results of the present work this seta may be present in a high
percentage of the adult specimens. This is an interesting result because labial chaetotaxy is a widely used character
in Lepidocyrtus species separation (see Mateos 2008a).
L. bicoloris sp. nov. and L. cyaneus have high variability in the presence / absence of seta ml of abd.II (Table
3), with frequent bilateral asymmetries in the same specimen. According to Mateos (2008a) in L. cyaneus this seta
is present, while according to Christiansen & Bellinger (1980) it is absent (in North American specimens). The
results of the present study show that there is intrapopulational variability of this character. L. bicoloris sp. nov. and
L. cyaneus have also variability in the unguiculus morphology (Table 3) (smooth / serrated), and L. bicoloris sp.
nov. in the number of odd teeth in the inner side of the unguis (1–2, Table 3).
Some of the published descriptions of several characters of L. lanuginosus contain errors and can lead to
misinterpretation. Gisin (1963b) gives a wrong scheme of dorsal cephalic macrochaetotaxy, and one year later
(Gisin 1964a) he corrects this error. Steve Hopkin, on the website "http://www.stevehopkin.co.uk/collembolamaps/
Entomobryomorpha/199LElan/" provides an image of the abd.IV anterior trichobothrial complex with seta s
present (photo of a UK specimen deposited at the MNH of London); in all published descriptions consulted (and in
the present work) seta s is always absent in this species. Snider (1967), in a description of North American
specimens, gives an abd.III chaetotaxic scheme without setae am6 and a6. Finally Mateos (2008a) gives a wrong
interocular chaetotaxy (s, t, r, q, p) due to a misinterpretation of microscopic preparations.
TABLE 3. Lepidocyrtus lanuginosus group. Some characters of adult specimens. Long: body length in mm without head and
furca; Body pigment: extension of body pigmentation; Cephalic A: number of setae in dorsal cephalic row A; Labium:
chaetotaxy of posterior labial row; Abd.II seta ml: presence/absence of abd.II seta ml; Abd.IV seta Fe4: morphology of abd.IV
seta Fe4, sm—smooth mesochaeta, cM—ciliated macrochaeta; Abd.IV seta E4p: morphology of abd.IV seta E4p, sm—smooth
mesochaeta, cM—ciliated macrochaeta; TV lat flap: number of setae on lateral flap of the ventral tube, c—ciliated seta,
s—smooth seta; Troch org: number of setae on trochanteral organ (third leg); Unguis teeth: number of inner odd teeth of the
unguis; Unguiculus: morphology of the unguiculus outer margin. In brackets number of specimens.
L. lanuginosus (12) L. bicoloris (22) L. cyaneus (16)
Long 0.9–1.2 0.7–1.0 1.0–1.2
Body pigment Absent from th.II to abd.II from th.II to abd.VI
Cephalic A 7–9 7–9 9–10
Labium MR*(8)
M*MR* (4)
MR* (21)
M*MR* (1)
MR*(3)
M*MR* (13)
Abd.II seta ml absent (12) absent (19)
present (3)
absent (12)
present (4)
Abd.IV seta Fe4 sm sm cM
Abd.IV seta E4p sm (10)
cM (2)
sm (18)
cM (4)
sm (3)
cM (13)
TV lat flap 5–7c + 2s 4–6c + 2s 6–8c + 2s
Troch org 8–16 8–16 10–17
Unguis teeth 1 (12) 1 (18)
2 (4)
1 (16)
Unguiculus smooth (12) smooth (18)
serrate (4)
smooth (13)
serrate (3)

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EUROPEAN LEPIDOCYRTUS LANUGINOSUS GROUP
As is the case in the other European Lepidocyrtus groups (L. lusitanicus species complex sensu Mateos 2008b,
L. lignorum group sensu Mateos 2011, L. curvocillis group sensu Mateos & Pettersen 2012, L. pallidus-serbicus
group sensu Winkler & Traser 2012), L. lanuginosus group shows high variability in body color pattern: body
without pigment (L. lanuginosus) body partially dark blue pigmented (L. bicoloris sp. nov.) and body entirely dark
blue pigmented (L. cyaneus).
Acknowledgements
Dr. Wanda M. Weiner, Dr. Gy. Traser and other anonymous reviewer provided helpful comments that improved the
manuscript.
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