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38 Accepted by W.M. Weiner: 23 Sept. 2008; published: 29 Oct. 2008
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2008 · Magnolia Press
Zootaxa 1917: 38–54 (2008)
www.mapress.com/zootaxa/
Definition of Lepidocyrtus lusitanicus Gama, 1964 species-complex (Collembola,
Entomobryidae), with description of new species and color forms from the
Iberian Peninsula
EDUARDO MATEOS
Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Avinguda Diagonal, 645, 08028 Barcelona (Spain).
E–mail: emateos@ub.edu
Abstract
Lepidocyrtus lusitanicus is one of the species with greater color pattern variability within the genus Lepidocyrtus. Four
subspecies have been described in relation to its body color variation. The study of various populations of Lepidocyrtus
from the Iberian Peninsula has allowed me to describe the new chromatic form L. lusitanicus form A, as well as the new
species L. bilobatus, which is very closely related to L. lusitanicus, and also shows high color pattern variability. The
high similarity among species L. lusitanicus, L. selvaticus and L. bilobatus sp. nov., the high color pattern variability of
their populations, the overlapping geographical distribution of many studied populations, and the presence of three dorsal
macrochaetae between trichobothria m2 and a5 of the second abdominal tergum, led me to define the group Lepidocyrtus
lusitanicus species-complex. This group included the three abovementioned species with all of their color forms.
Key words: Collembola, Lepidocyrtus lusitanicus species-complex, taxonomy, new species, color pattern variability
Resumen
Lepidocyrtus lusitanicus es una de las especies con mayor variabilidad cromática dentro del género Lepidocyrtus. Cuatro
subespecies han sido descritas en función de su patrón pigmentario. El estudio de varias poblaciones de Lepidocyrtus de
la Península Ibérica me ha permitido describir la nueva forma cromática L. lusitanicus forma A, así como la nueva espe-
cie Lepidocyrtus bilobatus, muy próxima a L. lusitanicus, y también con variabilidad cromática acentuada. La gran simil-
itud entre las especies L. lusitanicus, L. selvaticus y L. bilobatus sp. nov., la gran variabilidad cromática de sus
poblaciones, la distribución geográfica solapada de muchas de las poblaciones estudiadas, y el hecho de tener tres
macrosedas dorsales entre los tricobotrios m2 y a5 del el segundo terguito abdominal, me han llevado a definir el com-
plejo de especies de Lepidocyrtus lusitanicus. Este grupo incluye las tres especies mencionadas con todas sus formas
cromáticas.
Palabras clave: Collembola, Lepidocyrtus lusitanicus species-complex, taxonomía, nueva especie, variabilidad
cromática
Introduction
The European Lepidocyrtus fauna consists of 25 described species (Mateos 2008), among which Lepidocyrtus
lusitanicus Gama, 1964 is one of those with greater color pattern variability. In the original description of this
species, Gama defined two subspecies based on color pattern, both from Portuguese localities: L. lusitanicus
lusitanicus and L. lusitanicus coloratus. Simón (2007) described two new subspecies from several Spanish
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
localities based on different color patterns: L. lusitanicus nigrus and L. lusitanicus piezoensis. Since its
description, L. lusitanicus has been found in many localities of the Iberian Peninsula (Spain and Portugal) and
is currently regarded as a species with a wide Iberian distribution (see: Jordana et al. 1990, Simón 2007).
There are only two records outside the Iberian Peninsula, one in the Balearic Islands (Arbea & Jordana 1990)
and one in the French Pyrenees (Bonnet et al. 1979). Recently, Arbea and Ariza (2007) have described from
various localities of the Northeastern Spain the species L. selvaticus, which is closely related to L. lusitanicus.
For the present paper, I have studied specimens of Lepidocyrtus from different populations distributed across
the Iberian Peninsula including all known L. lusitanicus subspecies and the species L. selvaticus. One of the L.
lusitanicus populations studied shows a different color pattern from that described for the four known subspe-
cies, and it is herein described as a new color form. Another population studied shows significant chaetotaxic
differences with L. lusitanicus and L. selvaticus, allowing the definition of a new species. The high morpho-
logical and chaetotaxic similarity among these species and subspecies, the high color pattern variability of
their populations, and the overlapping geographical distribution of many studied populations, led me to define
the group Lepidocyrtus lusitanicus species-complex.
Material and methods
I examined specimens from different populations of the species and subspecies L. lusitanicus lusitanicus, L.
lusitanicus coloratus, L. lusitanicus nigrus, L. lusitanicus piezoensis and L. selvaticus, as well as several pop-
ulations of two new Lepidocyrtus taxa (see: Tab. 1 and Fig. 1). From each population, at least ten specimens
were studied under phase contrast microscope. For the dorsal macrochaetae and labial chaetotaxy notation I
followed Gisin (1963, 1964a, 1964b), except for head macrochaetae row A codified following Barra (1975),
and head macrochaetae R1s codified following Wang and Christiansen (2003). For the dorsal chaetotaxy of
thoracic and abdominal segments I used the notation established by Szeptycki (1979), except for seta d2 (abd.
II) codified following Snider (1967), seta m7a (abd. III) according to Wang et al. (2003) and seta p8p (abd.
III) as in Mateos (2008). For a revision of the characters used in Lepidocyrtus species descriptions, see Mari
Mutt (1986), Soto-Adames (2000) and Mateos (2008).
Abbreviations used: The following abbreviations are used: ant.––antennal segment; th.––thoracic seg-
ment; abd.––abdominal segment; I–VI––segments.
Taxonomic section
Family Entomobryidae Schäffer, 1896
Genus Lepidocyrtus Bourlet, 1839
Lepidocyrtus bilobatus sp. nov.
Figs 2–20, Tab. 1
Type material. Spain, Salamanca (LOC139, see Tab. 1 and Fig. 1), on herbaceous vegetation along the
Alayón river, 2.viii.2007, E.Mateos coll. Holotype: female (CRBA1272) on 1 slide, paratypes: lot LP175
(unpigmented specimens) composed by 11 specimens prepared on slides and 80 specimens in alcohol, lot
LP176 (pigmented and unpigmented specimes) composed by 4 specimens mounted on slides and 16 speci-
mens in alcohol. Holotype and 7 paratypes mounted on slides deposited at the Centre de Recursos de Biodi-
versitat Animal, Faculty of Biology, University of Barcelona (http://www.crba.ub.edu): Slide CRBA1341
with one pigmented specimen, slides CRBA1342 and CRBA1343 with one unpigmented specimen each, slide
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40 · Zootaxa 1917 © 2008 Magnolia Press
CRBA1344 with two pigmented and two unpigmented specimens. Other paratypes deposited in the E. Mateos
collection.
Etymology. The species name refers to the bilobed morphology of the ant. IV apical bulb, which is a diag-
nostic character for the new species within the European Lepidocyrtus fauna.
FIGURE 1. Localities (in letters, see: column “ref” in Table 1) and taxa (in numbers). 1––L. lusitanicus lusitanicus, 2––
L. lusitanicus coloratus, 3––L. lusitanicus nigrus, 4––L. lusitanicus piezoensis, 5––L. lusitanicus form A, 6––L. selvati-
cus, 7––L. bilobatus sp. nov.
Description. Holotype body length (without head nor furca) 1 mm, paratypes 0.7–1.0 mm. Holotype body
unpigmented; paratypes body color highly variable with yellow-white (form A) and purple (form B) speci-
mens present in the same population (Figs 2–3); all specimens with purple pigment on anterior region of the
head and ant. II–IV. Ocular areas densely black pigmented. Antenna, legs, and dorsal side of manubrium with-
out scales. Body dorsoventrally compressed; mesothorax not projecting over the head.
Ratio antenna:cephalic diagonal = 1.5 for holotype, for paratypes 1.4–1.7; ant. I:II:III:IV = 1:1.9:2.1:3.6.
Basis of ant. I dorsally with three microchaetae arranged in triangle. Ant. III organ as in Fig. 4. With bilobed
nonretractile ant. IV apical bulb (Fig. 5). 8+8 eyes of equal size.
Smooth prelabral and labral setae in typical number 4/554; prelabral setae ciliated; the four labral setae of
the third row are shorter, more curved, and with more developed insertion than setae of the other two rows;
inverted U-shaped labral apical intrusion; with four rounded labral papillae (Fig. 6). Subapical seta of outer
maxillary palp smooth and slightly longer than or subequal to apical seta, with three smooth setae on sublobu-
lar plate (Fig. 7). Outer differentiated seta of labial appendage curved, tip almost reaching the apex of the
papilla (Fig. 8).
Labium anterior row (a1–a5) formed by smooth setae; posterior row formed by ciliated setae
(M2R*EL1L2), with R half in length on seta M2 (marked as R*) (Fig. 9). Ventral cephalic groove with 3+3 cili-
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
ated macrochaetae and 2+2 scales.
The dorsal macrochaetae formula is R0R1R2STSo/10/0301+3, with a pair of supplementary macrochaetae
R1s between R0 and R1 (Fig. 10). Maximum number of macrochaetae A on the head 9+9 (Fig. 11). Interocular
chaetotaxy with s, t, r, q, p ciliated setae, without scales (Fig. 12). The th. II dorsal macrochaeta correspond to
p3 (Fig. 13). Abd. II–III chaetotaxy as in Figs 14–15; abd. II setae a2, m3 and m3e ciliated macrochaetae, seta
a2p absent; abd. III seta m7a acuminate thin ciliated macrochaeta. Abd. IV chaetotaxy as in Fig. 16-A; tricho-
bothrium T2 without accessory seta s; seta D1p ciliated and double in length than other setae of the trichobo-
thrial complex (Fig. 17); seta Fe4 can be thin ciliated macrochaeta or smooth mesochaeta depending on the
specimens (Fig. 16-B). Bilateral asymmetries can be observed in some specimens in which Fe4 is thin ciliated
macrochaeta on one side and smooth mesochaeta on the other; without correlation between the morphology of
seta Fe4 and other characters like size, color pattern nor locality. Three posterior smooth mesochaetae on abd.
IV present. All setae associated with the trichobothria on abd. II–IV are acuminate and strongly ciliate, except
pi and pe on abd. IV which are fan-shaped.
TABLE 1. Material studied data (all material collected by the author); ref––locality reference (see: Fig. 1); code––local-
ity code; lat––latitude (all N latitudes); long––longitude (positive values means East, negative values means West); elev–
–elevation above sea level in m; date––collection date in dd-mm-yy format; method (capture methodology): v––beating
herbaceous vegetation, s––active searching in soil by hand with an aspirator (between litter-fall, under stones and under
logs); species: 1––L. lusitanicus lusitanicus, 2––L. lusitanicus coloratus, 3––L. lusitanicus nigrus, 4––L. lusitanicus
piezoensis, 5––L. lusitanicus form A, 6––L. selvaticus, 7––L. bilobatus sp. nov.
ref code locality province country lat long elev date method species
a LOC137 Bandeira Coimbra Portugal 40.2016 -8.8798 241 29-07-07 v 1, 5
b LOC126 Arrimal Leiria Portugal 39.4995 -8.8712 320 12-07-07 v 4
c LOC129 Pragais Leiria Portugal 39.5785 -8.8221 308 18-07-07 v 1
d LOC125 Mira Coimbra Portugal 40.4457 -8.7988 13 08-07-07 v 5
e LOC131 San Jacinto Aveiro Portugal 40.6706 -8.7252 9 21-07-07 v 1, 2
f LOC134 Arzila Coimbra Portugal 40.1830 -8.5462 120 26-07-07 v 1, 2
g LOC133 Taveiro Coimbra Portugal 40.2013 -8.5376 4 26-07-07 v 3
h LOC124 Curia Coimbra Portugal 40.4260 -8.4655 42 08-07-07 v 2
i LOC132 Coimbra Coimbra Portugal 40.2231 -8.4458 34 25-07-07 v 1, 5, 3
j LOC123 Buçaco Coimbra Portugal 40.3796 -8.3763 243 08-07-07 v 1, 2
k LOC135 Coimbra Coimbra Portugal 40.2227 -8.3349 9 28-07-07 v 1, 3
l LOC136 Casal de Santo Amaro Coimbra Portugal 40.2845 -8.3046 165 28-07-07 v 2
m LOC127 Gerês Braga Portugal 41.7371 -8.1579 465 14-07-07 v 1
n LOC128 Gerês Braga Portugal 41.7507 -8.1528 720 14-07-07 v 1
o LOC052 Pardieiros Coimbra Portugal 40.2168 -7.9096 600 22-07-07 v 1
p LOC138 El Casarito Salamanca Spain 40.5218 -6.1382 1065 02-08-07 v 1, 4
q LOC139 Sotoserrano Salamanca Spain 40.4110 -6.0535 364 02-08-07 v 7
r LOC120 Pina de Ebro Zaragoza Spain 41.4793 -0.2415 400 31-05-07 s 3
s LOC091 Vall d'Aran Lleida Spain 42.7755 0.7851 1450 17-07-06 v 1
t LOC097 Taull Lleida Spain 42.5102 0.8779 1800 20-07-06 s 1
u LOC108 Montsant Tarragona Spain 41.2370 0.8843 385 17-02-07 v 3
v LOC156 Vila-seca Lleida Spain 41.6850 0.9434 230 29-05-08 v 3
w LOC102 Prades Tarragona Spain 41.3594 1.0804 990 03-01-07 v 1
x LOC101 Viladecavalls Barcelona Spain 41.5469 1.9601 200 25-10-06 v 6
y LOC144 Blanes Girona Spain 41.6867 2.8165 2 28-02-08 v 6
z LOC143 Tossa de Mar Girona Spain 41.7190 2.9028 2 21-02-08 v 6
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FIGURES 2–10. Lepidocyrtus bilobatus sp. nov.: 2, L. bilobatus form A; 3, L. bilobatus form B; 4, ant. III sensillar cha-
etotaxy; 5, apical antennal bulb; 6, labrum; 7, maxillary palp; 8, outer labial appendage; 9, labium and ventral cephalic
groove; 10, dorsal macrochaetae (circles), trichobothria (lines) and pseudopora (#).
V-shaped trochanteral organ formed by 6–8 smooth straight setae (Fig. 18). Unguis with basal pair teeth at
46 % of the inner edge, and with two inner teeth at 66 % (the bigger) and 83 % of the inner edge respectively;
unguiculus lanceolate with finely serrated outer margin; spatulate tibiotarsal tenent hair (Fig. 19).
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
FIGURES 11–13. Lepidocyrtus bilobatus sp. nov.: 11, dorsal head chaetotaxy, broad circles––long ciliated macrochae-
tae, small circles––short ciliated macrochaetae; 12, interocular chaetotaxy (right ocular area); 13, th. II dorsal chaetotaxy,
large circles––ciliated macrochaetae, pse––pseudopore.
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FIGURES 14–15. Lepidocyrtus bilobatus sp. nov.: 14, abd. II complete chaetotaxy, large circles––blunt broad ciliated
macrochaetae, pse––pseudopore; 15, abd. III complete chaetotaxy, large circles––blunt broad ciliated macrochaetae,
small circles––acuminate thin ciliated macrochaetae, pse––pseudopore.
Ratio manubrium : dens : mucro = 23:21:1. Manubrial plate with 2 inner setae and 4–5 external setae (Fig.
20).Ecology and distribution. All the individuals were obtained beating the herbaceous vegetation along the
Alayón river.
Discussion. The bilobed antennal apical bulb is a diagnostic character that separates L. bilobatus sp. nov.
from all other European species. The new species is very close to L. lusitanicus and L. selvaticus, from which
it can be differentiated by abd. IV chaetotaxy: In L. bilobatus sp. nov. setae E1, De1 and E4p are ciliated mac-
rochaetae (Fig. 16), whereas in the other two species are smooth mesochaetae (Fig. 27). The new species also
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
differs from L. selvaticus by the presence of labial seta R (absent in L. selvaticus) and by having 3+3 ciliated
setae along ventral cephalic groove (2+2 ciliated and 1+1 smooth in L. selvaticus, see: Fig. 29). As L. lusitan-
icus, the new species has high color pattern variability, with white (L. bilobatus form A) and purple pigmented
specimens (L. bilobatus form B) living in the same population. The fact that specimens with different color
pattern live together in the same population, suggests that body pigment distribution cannot be considered a
good subspecific character in L. bilobatus sp. nov.; for this reason L. bilobatus form A and form B have been
described as color forms and not as subspecies.
FIGURE 16. Lepidocyrtus bilobatus sp. nov.: 16-A, abd. IV complete chaetotaxy, large black circles––blunt broad cili-
ated macrochaetae, small black circles––acuminate thin ciliated macrochaetae, white circles––smooth mesochaetae, oval
(Fe4 seta)––thin ciliated macrochaetae or smooth mesochaetae, triangles––fan-shaped setae, lines––microchaetae, x––
trichobothria, #––pseudopore; 16-B, bcm––blunt broad ciliated macrochaeta, tcm––acuminate thin ciliated macrochaeta,
Fe4––two different morphologies of seta Fe4.
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FIGURES 17–20. Lepidocyrtus bilobatus sp. nov.: 17, abd. IV trichobothrial complex; 18, trochanteral organ (other
setae than trochanteral organ omitted); 19, p-III unguis and unguiculus; 20, manubrial plates, black circles––pseudopora.
Lepidocyrtus lusitanicus Gama, 1964
Figs 21–27 Tab. 1
Material examined. Specimens from 22 localities in the Iberian Peninsula of the four subspecies described so
far, as well as the new color form mentioned in this paper: L. lusitanicus lusitanicus, L. lusitanicus coloratus,
L. lusitanicus nigrus and L. lusitanicus piezoensis (see: Tab. 1 and Fig. 1). I found the new color form (A) in
three localities: Portugal (see Tab. 1 and Fig. 1): Mira pine litoral forest (LOC125), 54 specimens, 8.vii.2007,
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
E. Mateos coll.; Mata de Choupal (LOC132), 35 specimens, 25.vii.2007, E. Mateos coll.; Mata Nacional
Prazo de Sta. Marinha (LOC137), 80 specimens, 29.vii.2007, E. Mateos coll.
FIGURES 21–25. Lepidocyrtus lusitanicus-complex: 21, L. lusitanicus lusitanicus; 22, L.lusitanicus coloratus; 23, L.
lusitanicus nigrus; 24, L. lusitanicus piezoensis; 25, L. lusitanicus form A.
Diagnosis. Small Lepidocyrtus (1 mm of maximum body length) with mesothorax not projecting over the
head and body color very variable, from white to purple and with various color patterns taxonomically consid-
ered subspecies (Figs 21–25). The new form (A) possess characteristic color pattern (Fig. 25): diffuse purple
pigment in ant. II–IV, head, th. II–abd. II and coxae, and with the th. II anterior margin conspicuously darker
pigmented. Antennae, legs and dorsal side of manubrium not scaled. Ant. IV with simple nonretractile apical
bulb (Fig. 26). Labium chaetotaxy [M1] M2R*EL1L2 (M1 may be present or absent, R half in length than other
setae). Labrum 4/554; prelabral setae ciliated, labral setae smooth; inverted U-shaped labral apical intrusion.
Ventral cephalic groove with 3+3 ciliated setae. Head with a maximum of 7+7 macrochaetae in row A.
Intraocular chaetotaxy with ciliated setae s, t, r, q, p, without scales. Dorsal macrochaetae formula
R0R1R2STSo/10/0301+3, with supplementary macrochaetae R1s between R0 and R1. Abd. II setae a2, m3 and
m3e ciliated macrochaetae; seta a2p absent. Abd. IV trichobothrium T2 without accesory seta s; seta D1p cil-
iated and double in length than other setae of the trichobotrial complex; setae E1, De1 and E4p smooth
mesochaetae; setae Fe4 ciliated macrochaeta or smooth mesochaeta depending on the specimens (Fig. 27). All
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setae associated with the trichobothria on abd. II–IV are acuminate and strongly ciliate. V-shaped trochanteral
organ formed by 5–8 smooth setae. Inner edge of the unguis with basal pair teeth and one or two odd teeth;
unguiculus with outer edge serrate or smooth. Manubrial plate with 2 internal setae and 2–4 external setae.
FIGURES 26–27. Lepidocyrtus lusitanicus: 26, apical antennal bulb; 27, abd. IV chaetotaxy, broad circles––broad cili-
ated macrochaetae, small circles––thin ciliated macrochaetae, white circle––pseudopore.
Discussion. All studied specimens showed the morphological and chaetotaxic characters described for
this species by Gama (1964), Simón (2007) and Mateos (2008). These characters match the description of the
new species L. bilobatus (Figs 2–20), except for the apical antennal bulb (simple in L. lusitanicus) and abd. IV
setae E1, De1 and E4p (smooth mesochaetae in L. lusitanicus).
Different subspecies are distinguishable only by the color pattern. According to Simón (2007), subspecies
L. lusitanicus piezoensis have diffuse pigment in th. II–abd. II, with the th. II anterior margin more densely
pigmented, and two dorsal dark spots on abd. III and abd. IV. Specimens of this subspecies examined by me,
only have the abd. III and abd. IV dark spots (Fig. 24).
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
Lepidocyrtus selvaticus Arbea & Ariza, 2007
Figs 28–29, Tab. 1
Material examined. I have examined specimens from three localities in Northeastern Spain (LOC101,
LOC143, LOC144, Tab. 1 and Fig. 1).
Diagnosis. Small Lepidocyrtus (maximum body length: 0.97 mm) with mesothorax not projecting over
the head. Body yellowish (Fig. 28). Antennae, legs and dorsal side of manubrium not scaled. Ant. IV with
simple nonretractile apical bulb. Labium chaetotaxy M2EL1L2; ventral cephalic groove with 2+2 ciliated setae
and 1+1 (posterior) smooth setae (Fig. 29). Labrum 4/554; prelabral setae ciliated; labral setae smooth;
inverted U-shaped labral apical intrusion. Head with 5+5 macrochaetae A. Interocular chaetotaxy with cili-
ated setae s, t, r, q, p, without scales. Dorsal macrochaetae formula R0R1R2STSo/10/0301+3, with supplemen-
tary macrochaetae R1s between R0 and R1. Abd. II with ciliated macrochaetae a2, m3 and m3e; seta a2p absent.
Abd. IV trichobothrium T2 without accesory seta s; seta D1p ciliated and double in length than other setae of
the trichobotrial complex; setae E1, De1 and E4p smooth mesochaetae. All setae associated with the trichobo-
thria on abd. II–IV are acuminate and strongly ciliate. V-shaped trochanteral organ formed by 5 smooth setae.
Inner edge of the unguis with basal pair teeth and one odd teeth; unguiculus with serrate outer edge. Manu-
brial plate with 2 internal and 2 external setae.
FIGURES 28–29. Lepidocyrtus selvaticus: 28, habitus; 29, labium and ventral cephalic groove.
Ecology and distribution. The examined specimens came from two localities on the beach (few meters
away from the sea), which were already described in the species original description (LOC144 and LOC143),
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and one non-coastal locality (LOC101) located 27 km inland. In the beach localities, specimens were col-
lected over the low herbaceous vegetation growing in the sand. In the inland locality, specimens were col-
lected also on herbaceous vegetation.
Discussion. All examined specimens have the morphological and chaetotaxic characters described by
Arbea and Ariza (2007), but with some differences in the abdominal chaetotaxy (see: Arbea & Ariza 2007,
Fig. 4). In abd. II terga, the original description indicates the presence of a smooth mesochaeta between setae
ll and a6, which is absent in the specimens examined by me; in abd. III terga, the presence of two microchae-
tae between setae d3 and p5 is indicated, whereas in my specimens there is only one (microchaeta d2); seta
m7a is drawn as a smooth mesochaeta, while in my specimens it is an acuminate thin ciliated macrochaeta.
Therefore, the abd. II–III chaetotaxy of my L. selvaticus specimens is the same as described for L. bilobatus
and L. lusitanicus (Figs 14–15). With this diagnosis the description of L. selvaticus is extended for the follow-
ing characters: interocular chaetotaxy, trocanteral organ, manubrial plate, and abd. IV total chaetotaxy. The
abd. IV chaetotaxy of L. selvaticus is the same as described for L. lusitanicus (Fig. 27), with the difference
that seta Fe4 is smooth mesochaeta in all examined specimens.
L. selvaticus is a closely related species with L. lusitanicus and L. bilobatus sp. nov., from which it can be
differentiated by the absence of labial seta R and by having smooth 1+1 posterior setae instead of ciliated of
the cephalic groove (Fig. 29). It can be also distinguished from L. bilobatus sp. nov. by having simple apical
antennal bulb and abd. IV setae E1, De1 and E4p smooth mesochaetae.
Lepidocyrtus lusitanicus species-complex
Figs 30–35, Tab. 1
Diagnosis. Small Lepidocyrtus (1 mm of maximum body length) with mesothorax not projecting over the
head and body color very variable, from white to purple and with variable color pattern (Figs 30–34; see: Figs
2–3, Figs 21–25 and Fig. 28). Antennae, legs and dorsal side of manubrium not scaled. Ant. IV with simple or
bilobed nonretractile apical bulb. Labium with all setae of posterior row ciliated, with one or two M setae,
with or without seta R. Labrum 4/554; prelabral setae ciliated, labral setae smooth; inverted U-shaped labral
apical intrusion. Ventral cephalic groove with 3+3 setae, all ciliated or 2+2 ciliated and 1+1 (posterior)
smooth. Head with 5–9 macrochaetae A. Interocular chaetotaxy with ciliated setae s, t, r, q, p, without scales.
Dorsal macrochaetae formula: R0R1R2STSo/10/0301+3, with a pair of supplementary macrochaetae R1s
between R0 and R1. Abd. II setae a2, m3 and m3e ciliated macrochaetae; seta a2p absent. Abd. IV without seta
s, seta D1p ciliated and twice as long as other setae in the trichobotrial complex. Trochanteral organ V-shaped,
formed by 5–8 smooth setae. Inner edge of the unguis with basal pair teeth and one or two unpaired teeth.
Manubrial plate with 2 internal setae and 2–5 external setae.
The species-complex included the following taxa : L. lusitanicus lusitanicus, L. lusitanicus coloratus, L.
lusitanicus nigrus, L. lusitanicus piezoensis, L. lusitanicus form A, L. selvaticus and L. bilobatus sp. nov
(form A and form B).
Ecology and distribution. L. lusitanicus species-complex has a wide distribution in the Iberian Peninsula
(see: Jordana et al. 1990, Arbea & Ariza 2007, Simón 2007). L. lusitanicus s. str. has been cited in one occa-
sion from the Balearic Islands (Menorca Island: Monte Toro, Arbea & Jordana 1990) and in one occasion
from the French Pyrenees (Haute Vallée de l’Aude: Usson, 850 m altitude, Bonnet et al. 1979). L. lusitanicus
species-complex can be thought of as endemic to the Iberian region, including Balearic Island and French
Pyrenees (Fig. 35). The specimens live on herbaceous vegetation or in soil; they can inhabit various types of
forest or on beach herbaceous vegetation, and a very wide altitudinal range (from sea level up to 1800 meters
above sea level). The intestinal contents of all the examined specimens consists of fungal hyphae and spores.
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
FIGURES 30–34. Lepidocyrtus lusitanicus species-complex color patterns: 30, L. lusitanicus lusitanicus, L. selvaticus
and L. bilobatus sp. nov. form A; 31, L. lusitanicus coloratus; 32: L. lusitanicus nigrus and L. bilobatus sp. nov. form B;
33: L. lusitanicus piezoensis; 34, L. lusitanicus form A.
Discussion. The definition of L. lusitanicus species-complex is justified because, in all included taxa, abd.
II setae a2, m3 and m3e are ciliated macrochaetae; in all other European Lepidocyrtus species seta a2 is
smooth or ciliate mesochaeta, and seta m3e is smooth mesochaeta, so their presence as ciliated macrochaetae
is a diagnostic character of L. lusitanicus species-complex within European Lepidocyrtus fauna. Globally, the
L. lusitanicus species-complex shows five different color patterns (Figs 30–35); both L. lusitanicus and L.
bilobatus sp. nov. show body color variability, especially the former, while L. selvaticus only includes unpig-
mented individuals.
Lepidocyrtus lusitanicus species-complex identification key
1 Apical antennal bulb bilobed; abd. IV setae E1, De1 and E4p ciliated macrochaetae.L. bilobatus sp. nov.
- Apical antennal bulb simple; abd. IV setae E1, De1 and E4p smooth mesochaetae ................................. 2
2 Labial seta R absent; ventral cephalic groove with 2+2 ciliated setae and 1+1 smooth setae L. selvaticus
- Labial seta R present; ventral cephalic groove with 3+3 ciliated setae (L. lusitanicus) ............................ 3
3 Body unpigmented.................................................................................................L. lusitanicus lusitanicus
- Dark pigment on th. III–abd. II................................................................................L. lusitanicus coloratus
- Dark pigment on head, thorax, and abd. I–V................................................................L. lusitanicus nigrus
- Patches of dark pigment on abd. III–IV..................................................................L. lusitanicus piezoensis
- Difuse pigment on head and th. II–abd. III with a conspicuous pigmented th. II anterior margin................
.................................................................................................................................... L. lusitanicus form A
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FIGURE 35. Distribution of Lepidocyrtus lusitanicus species-complex: 1––L. lusitanicus lusitanicus, 2––L. lusitanicus
coloratus, 3––L. lusitanicus nigrus, 4––L. lusitanicus piezoensis, 5––L. lusitanicus form A, 6––L. selvaticus, 7––L. bilo-
batus sp. nov. Data from Bonnet et al. (1979), Arbea & Ariza (2007), Arbea & Jordana (1990), Jordana et al. (1990),
Simón (2007) and present paper.
General discussion
The genus Lepidocyrtus is represented in Europe (including L. bilobatus sp. nov.) by 26 species (Mateos
2008). Fifteen species inhabit the Iberian Peninsula (Spain and Portugal), of which five are endemic to the
area: L. montseniensis Mateos, 1985, L. pseudosinelloides Gisin, 1967, L. tellecheae Arbea & Jordana, 1989,
L. selvaticus Arbea & Ariza, 2007 and L. bilobatus sp. nov. The species L. lusitanicus has been found outside
the Iberian Peninsula only twice and in localities related to the Iberic area: French Pyrenees (Bonnet et al.
1979) and the island of Menorca (Arbea & Jordana 1990). The species L. selvaticus and L. bilobatus sp. nov.
are only known from Spanish localities. Based on these data, L. lusitanicus species-complex can be consid-
ered an endemic group from southern Europe (Pyrenean region, Balearic Islands, and Iberian Peninsula, see:
Fig. 35), where it has undergone extensive evolution in color pattern and some speciation.
Differentiation between species that make up L. lusitanicus species-complex can be accomplished only if
adult specimens are available. Immature Lepidocyrtus specimens, and Entomobryidae in general (see: Barra
1975, Szeptycki 1979), have an undifferentiated chaetotaxy so the species identification is often impossible.
The two diagnostic characters of L. selvaticus (smooth 1+1 posterior setae of cephalic groove and the absence
of labial seta R) are presented in immature L. lusitanicus, so the separation of these two species is only possi-
ble if adult specimens are available. The abd. IV total chaetotaxy clearly separates L. bilobatus sp. nov. from
L. lusitanicus and L. selvaticus. As Mateos (2008) suggested, abd. IV total chaetotaxy seems to be very infor-
mative in separating European Lepidocyrtus species.
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LEPIDOCYRTUS LUSITANICUS SPECIES-COMPLEX
Within Lepidocyrtus European fauna, L. lusitanicus species-complex can be clearly distinguished because
abd. II setae a2 and m3e are ciliated macrochaetae, whereas in other species these are smooth or ciliate
mesochaetae, but never macrochaetae. Another feature of L. lusitanicus species-complex is the high color pat-
tern variability presented by species L. lustianicus and L. bilobatus sp. nov. As a group, five different color
patterns can be distinguished, ranging from head and body yellowish-white to dark purple (Figs 30–35). The
most widespread color pattern is the unpigmented one (with pigment only in antenna and frontal cephalic
spot), which corresponds to L. lusitanicus lusitanicus, L. selvaticus and L. bilobatus sp. nov. form A. In
Europe there are two more species with variable color pattern: L. lignorum (Fabricius, 1793) and L. ruber
Schött, 1902. L. lignorum is basically an unpigmented species, but Gama (1973) described the variety L.
lignorum var. (without naming it) with pigment on the ventral region of the head and abd. II–III terga, and
Ellis (1976) described the variety as L. lignorum forma? with pigmented spots on abd. III–IV. The L. ruber
color pattern has been described as very variable, from yellow to dark violet (Szeptycki 1967), which is the
same variation found in L. bilobatus sp. nov.
The fact that the color patterns of L. lusitanicus and L. bilobatus are mixed in several populations studied
in this paper, rises doubt of whether they should be given the taxonomic range of subspecies or simply chro-
matic forms. The populations of L. lusitanicus with the new color pattern reported here have been considered
as a color form (A). The two color patterns of L. bilobatus sp. nov. have been described as forms (form A and
form B). Soto-Adames (2002) using molecular data (COI sequences) suggested that several Puerto Rican pop-
ulations of L. biphasis Mari-Mutt, 1986 and L. dispar Mari-Mutt, 1986, differing only in color pattern, are dis-
tinct species. Perhaps the application of molecular analysis as those used by Soto-Adames may help decide
the taxonomic status of the different color forms of the L. lusitanicus species-complex taxa.
Acknowledgments
I thank Graça Vale, Paulo Sousa and Maria Manuela da Gama who have welcomed me into the Department of
Zoology at the University of Coimbra (Portugal). Wanda Maria Weiner, Felipe N. Soto-Adames and György
Traser provided helpful comments that improved the manuscript . I also thank Jordi García the linguistic revi-
sion of the manuscript. The author obtained a fellowship from AGAUR (Generalitat de Catalunya) for a one-
month visit to the University of Coimbra.
References
Arbea, J.I & Ariza, E. (2007) Una nueva especie de Lepidocyrtus Bourlet, 1839 (Collembola: Entomobryidae) de
Gerona, España. Boletín de la Sociedad Entomológica Aragonesa, 41, 87–89.
Arbea, J.I. & Jordana, R. (1989) New species of Pseudosinella and Lepidocyrtus from Navarra (Northern Iberian Penin-
sula), Spixiana, 13 (1), 25–31.
Arbea, I. & Jordana, R. (1990) Colémbolos de las Islas Baleares (Insecta, Collembola). Redia, 73, 187–200.
Barra, J.A. (1975) Le dévoloppement postembryonnaire de Pseudosinella decipiens et P. Impediens. Annales Spéléolo-
gie, 30(1), 173–186.
Bonnet, L., Cassagnau, P. & Deharveng, L. (1979) Recherche d'une méthodologie dans l'analyse de la rupture des équili-
bres biocénotiques: application aux Collemboles édaphiques des Pyrénées. Revue d'Ecologie et de Biologie du Sol,
16(3), 373–401.
Bourlet, A. (1839) Mémoire sur les Podures. Mémoires de la Société des Sciences de l' Agriculture de Lille, 1, 377–417.
Ellis, W.N. (1976) Autumn fauna of Collembola from Central Crete. Tijdschrift voor Entomologie, 119, 221–326.
Fabricius, J.C. (1793) Entomologia Systematica Emendata et Aucta, Secundum Classes, ordines, genera, species adjectis
synonimis, locis, observationibus, descriptionibus. Volume 2, C. G. Proft, Hafniae, 519 pp.
Gama, M.M. da (1964) Colêmbolos de Portugal Continental. Phd Universidade de Coimbra (Portugal), 252 pp.
Gama, M.M. da (1973) Systématique évolutive des Pseudosinella. IX (Insecta: Collembola). Revue Suisse de Zoologie,
80(1), 45–63.
TERMS OF USE
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MATEOS
54 · Zootaxa 1917 © 2008 Magnolia Press
Gisin, H. (1963) Collemboles d’Europe V. Revue Suisse de Zoologie, 7, 77–101.
Gisin, H. (1964a) Collemboles d’Europe VI. Revue Suisse de Zoologie, 71(2), 383–400.
Gisin, H. (1964b) Collemboles d’Europe VII. Revue Suisse de Zoologie, 71(4), 649–678.
Gisin, H. (1967) Deux Lepidocyrtus nouveaux pour l’Espagne. Eos, 42(3–4), 393–396.
Jordana, R., Arbea, J.I. & Ariño, A.H. (1990) Catálogo de colémbolos ibéricos. Base de datos. Publicaciones de Biología
de la Universidad de Navarra, Serie Zoológica, 21, 1–231.
Mari Mutt, J.A. (1986) Puerto Rican species of Lepidocyrtus and Pseudosinella (Collembola: Entomobryidae). Carib-
bean Journal of Sciences, 22(1–2), 1–48.
Mateos, E. (1985) Descripción de Lepidocyrtus montseniensis n.sp. (Collembola, Entomobryidae). Actas II Congreso
Ibérico Entomología, 27–34.
Mateos, E. (2008) The European Lepidocyrtus Bourlet, 1839 (Collembola: Entomobryidae). Zootaxa 1769: 35–59.
Schäffer, C. (1896) Die Collembola der Umgebung von Hamburg und benachbarter Gebiete. Mitteilungen aus dem
Naturhistorischen Museum in Hamburg, 13, 149–216.
Schött, H. (1902) Études sur les Collemboles du Nord. Bihang Svenska Vetenskaps-Akademiens Handlingar, 28(4), 1–
48.
Simón, J.C. (2007) Descripción de dos nuevas subespecies del género Lepidocyrtus Bourlet, 1839 (Collembola, Entomo-
bryidae) de la Península Ibérica. Graellsia, 63(2), 315–324.
Snider, R.J. (1967) The Chaetotaxy of North American Lepidocyrtus s. str., (Collembola: Entomobryidae). Contributions
of the American Entomological Institute, 2(3), 1–28.
Soto-Adames, F.N. (2000) Phylogeny of Neotropical Lepidocyrtus (Collembola: Entomobryidae): first assessment of
patterns of speciation in Puerto Rico and phylogenetic relevance of some subgeneric diagnostic characters. System-
atic Entomology, 25, 485–502.
Soto-Adames, F.N. (2002) Molecular phylogeny of the Puerto Rican Lepidocyrtus and Pseudosinella (Hexapoda: Col-
lembola), a validation of Yoshii's 'colour pattern species'. Molecular Phylogenetics and Evolution, 25, 27–42.
Szeptycki, A. (1967) Morpho-systematic studies on Collembola. Part 1. Materials to a revision of the genus Lepidocyrtus
Bourlet, 1839 (Entomobryidae s.l.). Acta Zoologica Cracoviensa, 12(13), 369–377.
Szeptycki, A. (1979) Chaetotaxy of the Entomobryidae and its phylogenetical significance. Morpho-systematic studies of
Collembola, IV. Polska Akademia Nauk, Zakład Zoologii Systematycznej i Do wiadczalnej, Kraków, Poland, 219
pp.
Wang, F., Chen, J.X. & Christiansen, K. (2003) Taxonomy of the genus Lepidocyrtus s.l. (Collembola: Entomobryidae)
in East and Southeast Asia and Malaysia, with description of a new species from the People's Republic of China.
Canadian Entomologist, 135, 823–837.
