Collembola: Entomobryidae

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Dec 3, 2012 - 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.
Zootaxa 3570: 69–81 (2012) www.mapress.com / zootaxa/ Copyright © 2012 · Magnolia Press

ISSN 1175-5326 (print edition)

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ZOOTAXA ISSN 1175-5334 (online edition)

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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: [email protected]

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. 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 caracterizada por el patrón pigmentario del cuerpo), así como una redefinición del grupo Lepidocyrtus lanuginosus de Europa. 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 & Bogojević 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). Accepted by W.M. Weiner: 25 Oct. 2012; published: 3 Dec. 2012

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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. 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

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

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 eyepatches 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 mesochaetae; 12, interocular chaetotaxy (right ocular area); 15, th.II complete chaetotaxy (left side), broad circles—broad ciliated macrochaetae, 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 [M1*] M2R*EL1L2; seta M1 present only in four out of twelve examined individuals, setae R and M1 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 macrochaetae, lines—mesochaetae, pse—pseudoporus; 17, abd.III complete chaetotaxy (left side), broad circles—broad ciliated macrochaetae, small circles—thin ciliated macrochaetae, lines—mesochaetae, pse—pseudoporus.

Dorsal macrochaetae formula such as R0R1sR1R2STSo/10/0101+2 (Fig. 12); a pair of supplementary macrochaetae R1s between R0 and R1 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 macrochaetae, small black circles—thin ciliated macrochaetae, lines—smooth mesochaetae and microchaetae, triangles—fanshaped 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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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. EUROPEAN LEPIDOCYRTUS LANUGINOSUS GROUP

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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 [M1*] M2R*EL1L2, M1 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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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 [M1*] M2R*EL1L2; 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 [M1*]M2R*EL1L2. Dorsal macrochaetae formula R0R1sR1R2STSo/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 M1 and the morphology of seta E4p of abd.IV (Table 3). Typically labial chaetotaxy of L. lanuginosus and L. cyaneus is described without seta M1 (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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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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EUROPEAN LEPIDOCYRTUS LANUGINOSUS GROUP

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