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Integrative taxonomy reveals three new species of European Lepidocyrtus lignorum-group (Collembola, Entomobryidae)

February 2022
Zootaxa 5100(4):451-481

DOI:10.11646/zootaxa.5100.4.1

Authors:

Eduardo Mateos

University of Barcelona

Marta Alvarez-Presas

Institute of Evolutionary Biology

Lepidocyrtus is one of the genera of springtails with the largest number of species in the world. Molecular studies carried out to date on this genus have revealed the existence of a large number of cryptic species. Molecular and morphological studies done with four European populations of the genus have allowed us to describe three new species within the L. lignorum-group: L. fuscocephalus sp. nov., L. milagrosae sp. nov. and L. semicoloratus sp. nov. One of these species is made up of two geographically separate populations that represent two clearly separate but morphologically indistinguishable haplotypes.

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ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)

Accepted by W. M. Weiner: 29 Jan. 2022; published: 22 Feb. 2022 451

Zootaxa 5100 (4): 451–481

https://www.mapress.com/zt/

https://doi.org/10.11646/zootaxa.5100.4.1

http://zoobank.org/urn:lsid:zoobank.org:pub:74EEFDED-EEB9-46DF-83D5-2FB5693F920E

Integrative taxonomy reveals three new species of European

Lepidocyrtus lignorum-group (Collembola, Entomobryidae)

EDUARDO MATEOS1,2,* & MARTA ÁLVAREZ-PRESAS3

1Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals; Facultat de Biologia; Universitat de Barcelona; Avinguda Di-

agonal 643, 08028 – Barcelona; Spain.

�

emateos@ub.edu; https://orcid.org/0000-0001-9741-5744

2Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona, Barcelona, Spain.

3University of Bristol, School of Life Sciences, Bristol, United Kingdom.

�

marta.alvarez-presas@bristol.ac.uk; https://orcid.org/0000-0002-4825-9965

*Corresponding author

Abstract

Lepidocyrtus is one of the genera of springtails with the largest number of species in the world. Molecular studies carried

out to date on this genus have revealed the existence of a large number of cryptic species. Molecular and morphological

studies done with four European populations of the genus have allowed us to describe three new species within the

L. lignorum-group: L. fuscocephalus sp. nov., L. milagrosae sp. nov. and L. semicoloratus sp. nov. One of these

species is made up of two geographically separate populations that represent two clearly separate but morphologically

indistinguishable haplotypes.

Key words: Chaetotaxy, colour pattern, COXII, EF-1α, Lepidocyrtinae, mPTP, ASAP, bPTP

Resumen

Lepidocyrtus es uno de los géneros de colémbolos con un mayor número de especies a escala mundial. Los estudios

moleculares realizados hasta la fecha en este género han revelado la existencia de un gran número de especies crípticas.

Estudios moleculares y morfológicos llevados a cabo con cuatro poblaciones europeas del género nos han permitido describir

tres nuevas especies en el L. lignorum-group: L. fuscocephalus sp. nov., L. milagrosae sp. nov. and L. semicoloratus sp.

nov. Una de estas especies está compuesta por dos poblaciones geográficamente alejadas que representan dos haplotipos

claramente separados pero morfológicamente indistinguibles.

Palabras clave: Quetotaxia, patrón de color, COXII, EF-1 α, Lepidocyrtinae, mPTP, ASAP, bPTP

Introduction

Lepidocyrtus Bourlet, 1839, with ~250 species, is one of the genera of springtails with a greater number of species,

and some investigations using molecular markers have demonstrated the presence of an important cryptic diversity

within the genus (Cicconardi et al. 2010, 2013, Mateos et al. 2018, Zhang B. et al. 2018, 2019). Morphological char-

acters currently used in species descriptions of Lepidocyrtus seem insufficient to describe the diversity discovered

by molecules, and looking for new characters is a necessary task (Mateos et al. 2021).

Colour pattern has always been used as an important character for species diagnoses in Lepidocyrtus and other

genera of Entomobryidae (Bonet 1934; Gisin 1960; Yosii 1961), although it is a trait to be taken with caution and

always with other associated characters (Soto-Adames 2002; Zhang B. et al. 2019). Using chaetotaxic characters,

the European members of Lepidocyrtus have been grouped into five species groups, and molecular analyses have

confirmed their validity (Mateos et al. 2018, Winkler et al. 2020). The most species-rich is the lignorum-group

MATEOS & ÁLVAREZ-PRESAS

which, currently, is composed of 16 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. labyrinthi Baquero & Jordana, 2021 (in Baquero et al. 2021), L. lignorum (Fabricius, 1793), L.

paralignorum Baquero & Jordana, 2021 (in Baquero et al. 2021), L. peisonis Traser & Christian, 1992, L. pulchellus

Denis, 1926, 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 (Geoffroy, 1762). All of them have trunk macrochae-

totaxy formula 00/0101+3 (Mateos 2011). 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) (Mateos

2020). Fundamental diagnostic characters other than pigmentation, dorsal macrochaetotaxy, unguis morphology

and scale distribution, are still unknown for the species L. instratus and L. vexillosus, and their diagnoses are mainly

based on the original descriptions (which are very concise). More studies about these two species are necessary in

order to elaborate an adequate morphological description that allows their comparison with the other species in the

group.

Recently, Mateos et al. (2021) detected three haplotypes within the European lignorum-group that clearly dif-

fered from the haplotypes of the known species in the group. Each of these three lineages had a characteristic colour

pattern, and was named as Lepidocyrtus spJ, spK, and spL, respectively. In the present paper we analyse the genetic

sequences of these three morphotypes using more molecular species delineation analyses than in the previous work

and describe them as three new species of the European Lepidocyrtus lignorum-group. Also, some notes on Lepido-

cyrtus instratus, a poorly described species in the literature, are provided.

Materials and methods

Specimens. Specimens of the new species were hand collected from soil litter (using micro aspirator) or herbaceous

vegetation (beating branches) and stored in microvials with absolute alcohol. Mitochondrial cytochrome oxidase

subunit II (CoxII) and nuclear Elongation Factor 1-alpha (EF) gene sequences were downloaded from the NCBI

GenBank public database (Table 1).

Molecular methods. Alignments were produced with the retrieved sequences in BioEdit (Hall 1999) using

amino acids as a guide since both genes are coding. In the case of EF, the intronic regions were aligned separately

using the online MAFFT program v7 (Rozewicki et al. 2019) with the G-INS-i refinement method and the other

options by default.

Phylogenetic trees were inferred for the three datasets (CoxII, EF and Concatenated) under the Maximum

Likelihood criterium (ML) using the software IQtree2 (Minh et al. 2020). The best model fit was inferred with the

TESTMERGE option, resembling PartitionFinder (Lanfear et al. 2017) by just considering the invariable site and

Gamma rate heterogeneity. For the individual alignments 100,000 ultrafast bootstrap (UFB) replicates (Minh et al.

2013) were run, while for the Concatenated dataset 10,000 UFB were inferred. For the analysis of the Concatenated

dataset, a partition scheme was taken into account by gene. A Bayesian Inference (BI) tree was inferred for the

Concatenated dataset using the software MrBayes v3.2. (Ronquist et al. 2012) for 8 million generations and two

independent runs, diagnosing MCMC convergence with Tracer v1.7. (Rambaut et al. 2018).

Molecular species delimitation was assessed using three different approaches. First, the multi-rate Poisson Tree

Process (mPTP) method (Kapli et al. 2017) under the maximum likelihood criterium, cropping the outgroup, and

using the individual gene ML trees in newick format previously inferred as input on the website (http://mptp.h-its.

org/#/tree). Second, the ASAP method (Puillandre et al. 2021), using the CoxII alignment as input and setting the

Kimura-80 as the model and the rest of parameters by default. Finally, we applied the Bayesian Poisson Tree Process

(bPTP) method (Zhang J. et al. 2013) on the website (http://species.h-its.org/ptp/) using the individual ML trees as

input and default parameters.

Morphology methods

For morphological studies specimens were cleared with Nesbit fluid and slide-mounted in Hoyer medium. Observa-

tions were made with a phase-contrast microscope with a digital camera attached.

TABLE 1. Species included in molecular analysis indicating population code, locality (province, country), and GenBank

accession numbers of sequences used.

GenBank Accession Number

Species Code Locality CoxII EF

Lepidocyrtus barbulus LP239_1 Crete, Greece MF095498 MF095594

LP239_2 MF095499 MF095595

Lepidocyrtus chorus LP383_1 Miljacka, Croatia MF095522 MF095609

LP383_2 MF095523 -

Lepidocyrtus fuscocephalus sp. nov. LP125_1 Barcelona, Spain MT345927 MT345994

LP125_2 MT345928 MT345995

LP125_3 MT345929 MT345996

LP125_4 MT345930 MT345997

LP125_5 MT345931 MT345998

Lepidocyrtus intermedius LP114_1 Barcelona, Spain MF095458 -

LP114_2 MF095459 MF095569

Lepidocyrtus juliae LP229_1 Crete, Greece MF095490 MF095590

LP229_2 MF095491 -

Lepidocyrtus lanuginosus LP130_1 Barcelona, Spain MF095474 MF095579

LP130_2 MF095475 MF095574

Lepidocyrtus lignorum LP118_1 Barcelona, Spain MF095465 MF095570

LP118_2 MF095466 MT345988

Lepidocyrtus lusitanicus LP100_1 Tarragona, Spain MF095537 MF095614

LP100_2 MF095540 -

Lepidocyrtus milagrosae sp. nov. LP250_4 Rhodes, Greece MT345941 -

LP250_5 MT345942 MT346008

LP250_6 MT345943 -

LP250_7 MT345944 -

Lepidocyrtus montseniensis LP129_1 Barcelona, Spain MF095470 MF095571

LP129_2 MF095471 MF095572

Lepidocyrtus pallidus LP362_3 Vestfold, Norway MF095519 MF095608

LP362_4 MF095520 -

Lepidocyrtus pulchellus LP389_7 La Spezia, Italy MT345946 -

LP389_8 MT345947 MT346010

Lepidocyrtus semicoloratus sp. nov. LP108_2 Barcelona, Spain MT345918 MT345984

LP108_3 MT345919 MT345985

LP108_4 MT345920 MT345986

LP108_5 MT345921 MT345987

LP122_1 Valencia, Spain MT345922 MT345989

LP122_2 MT345923 MT345990

LP122_3 MT345924 MT345991

LP122_4 MT345925 MT345992

LP122_5 MT345926 MT345993

Lepidocyrtus tellecheae LP106_2 Barcelona, Sain MF095529 MF095564

LP106_3 MF095530 MF095565

Lepidocyrtus traseri LP546_1 Pest, Hungary MT345963 MT346024

LP546_3 MT345965 MT346026

Lepidocyrtus violaceus LP096_1 A Coruña, Spain MT345910 MT345977

LP096_3 MT345912 MT345979

MATEOS & ÁLVAREZ-PRESAS

In January 2019 one slide was revised (deposited at the National Museum of Natural History-NMNH, Paris,

France) containing one specimen of L. instratus prepared and identified by Gisin in 1948. With the study of this

specimen along with the information provided in several publications (Handschin 1924, Gisin 1964a, 1964b), a

diagnosis of the species is proposed in the present paper.

For the taxonomic descriptions, the following terms and codes were used: For dorsal cephalic chaetotaxy the

“AMS” nomenclature system (see Soto-Adames 2010). For Ant.I-organ Hüther (1986). For labial papillae and

maxillary palp Fjellberg (1999). For clypeal chaetotaxy Yoshii & Suhardjono (1992) and Zhang F. et al. (2016). For

labial chaetotaxy Gisin (1964b). 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, Mateos (2008), and Zhang F. et al. (2019). For tergal specialized chaetae (S-chaetae)

Zhang F. & Deharveng (2015). For pseudopores distribution Mateos et al. (2021).

Abbreviations and symbols in text and figures: BP4—basal plate of the fourth abdominal segment, Ant.—an-

tennal segment, Th.—thoracic segment, Abd.—abdominal segment, I–VI—segment numbers, pse—pseudopore.

Results

Molecular section

Molecular results. CoxII dataset is composed of 44 sequences all belonging to the Lepidocyrtus genus with a length

of 688 bp. EF dataset contains 36 sequences and a length of 632 bp. The Concatenated dataset bears 44 sequences

and a total length of 1320 bp. The topology of the two individual gene trees is different, perhaps due to different

taxon sampling, so it is difficult to establish a good comparison. In both phylogenies, however, all known species

are monophyletic. Regarding the Concatenated dataset results, both ML (Fig. 1) and BI give similar topologies.

Lepidocyrtus lusitanicus, L. lanuginosus and L. pallidus appear at the base of the tree, sister to L. montseniensis, that

is sister to all the other species (representing the L. lignorum-group on the tree). The new species L. milagrosae sp.

nov. belongs to the same clade as L. juliae, and both group with the other species described here, L. fuscocephalus

sp. nov.. All the individuals from the third new species, L. semicoloratus sp. nov. form a single cluster, showing

long branches separating both populations (Montseny – named LP108, and Valencia – named LP122). Sister to this

cluster appears a big clade formed by the species L. pulchellus, L. tellecheae, L. traseri, L. intermedius, L. lignorum,

L. violaceus, L. barbulus and L. chorus. All the species are shown as monophyletic with high support; however, the

relationships of big clades lack statistical support in most of the cases.

Molecular species delimitation. The mPTP results disagree between both molecular markers used. In the

EF analysis, all the specimens seem to belong to the same species, while the CoxII results show all the previously

morphologically defined species as such. The same occurs with the other species delimitation methods applied, so

it is impossible to comment on the results of the EF marker for species delimitation. Regarding the CoxII results,

all three methods agree in the species delimitation, showing 17 entities as putative species (Fig. 1). The number of

putative species retrieved by the 10 best partition schemes produced through ASAP ranges from 12 to 27. The best

composite score (named asap-score) is 1.5, and it corresponds to a partition with 17 putative species.

Taxonomic section

Family Entomobryidae Tömösváry, 1882

Genus Lepidocyrtus Bourlet, 1839

Lepidocyrtus instratus Handschin, 1924

Figs 2–7, Table 2

Material examined. One slide (deposited at the National Museum of Natural History-NMNH, Paris, France) con-

taining one specimen of L. instratus prepared and identified by Gisin in 1948. The slide was labeled with the code

“1b:Rr4/2”, and “Autriche, Tirol” as locality.

FIGURE 1. Phylogenetic tree inferred with Maximum Likelihood (ML) using the Concatenated dataset. Stars at nodes cor-

respond to ultrafast bootstrap values, and their size is proportional to the value, showing only values > 70%. Vertical bars to the

right of the phylogeny correspond to the molecular species delimitation methods results (ASAP, bPTP and mPTP from left to

right). Scale bar represents number of substitutions per site.

Diagnosis based on the original description of Handschin (1924), Gisin (1964a, 1964b) and specimen from

the slide “1b:Rr4/2”. Length 2.0 mm (maximum). With dark blue pigment on Ant.II–IV, buccal area, dorsal side of

Th.III to Abd.III (and dispersed over Abd.IV–V) and coxae (Fig. 2). Th.II sligthly projecting over head (Fig. 3). Ant.

I–II, legs, ventral tube and posterior region of manubrium with scales. Labial chaetae (M1M2REL1L2) in “p row”

well developed and ciliated, M1 a little shorter than M2, R half in length than M2 (Fig. 4). Dorsal cephalic and body

macrochaetae formula as A0A2A3Pa5/00/0101+3 (Fig. 5). Anterior trichobotrium of Abd.IV with accessory chaetae

a, D1 and m fan-shaped, without chaeta s (Fig. 6). Ungues with a small basal tooth at about 40% of the inner edge;

unguiculi lanceolate, with smooth and curved outer edge (Fig. 7).

Discussion. In the original description of Handschin (1924) the chaetotaxy was not described and the species

was characterised by the colour pattern, the ungues without inner teeth and the unguiculi truncate. The localities of

MATEOS & ÁLVAREZ-PRESAS

the specimens assigned for the type series are located in the Engadine region of Swiss Alps, at altitudes between

2600 and 2800 m a.s.l.; these specimens were collected under stones on the edge of a snowy field and in groundhog

droppings.

FIGURES 2–7. Lepidocyrtus instratus: 2, habitus lateral (from Handschin 1924); 3 Head and mesothorax in lateral view (pic-

ture from specimen in the slide “1b:Rr4/2”, NMNH); 4, labial chaetotaxy (sensu Gisin 1964b); 5, dorsal cephalic and body mac-

rochaetae (black filled dots), trichobothria (lines) and pseudopores (empty circles) (after Gisin 1964a); 6, anterior trichobotrium

of Abd.IV complex (after Gisin 1964b); 7 unguis and unguiculus (after Gisin 1964a).

Gisin (1964a), based on several topotypes, corrected the original description of Handschin (1924) and noted

the presence in the ungues of a small basal teeth (at about 40% of the inner edge) and a tiny tooth (at about 55% of

the inner edge), and the lanceolate morphology of the unguiculi rather than truncate. However, Gisin noted that the

unguiculi has a curved outer edge, giving the appearance of being truncated. Gisin (1964a) also described the dorsal

macrochaetotaxy of head and body, and confirmed the colour pattern of the species as the one described by Hand-

schin in the original description. Both Handschin and Gisin noted the dark pigmented buccal area as characteristic of

the species. In the same year, Gisin (1964b) described the chaetotaxy of the basal labium and the accessory chaetae

of the anterior trichobothrium of Abd.IV.

The slide from the NMNH revised by the author is, supposedly, part of the material used by Gisin (1964a) for

the revision of the species. The specimen was in very bad condition, and the chaetotaxy was completely invisible.

However, the morphology of several ungues and unguiculi was visible and was the same as that described by Gisin

(1964a).

As noted by the above paragraphs (and also previously noted by Mateos 2008) L. instratus is poorly described

in the literature. However, enough characters are known to ascribe the species to the European lignorum-group

(sensu Mateos (2011). Several fundamental chaetotaxic characters are still unknown, which makes difficult com-

parisons between this species and the other species of the lignorum-group with the standards currently required in

the genus Lepidocyrtus.

Lepidocyrtus fuscocephalus Mateos sp. nov.

Figs 8–23, Tabs 1–2

Zoobank: urn:lsid:zoobank.org:act:63B8C59B-4F13-45A9-A26B-3A71E784ADA4

Type material. Holotype: Male on two slides (CRBA-90744a (head) and CRBA-90744b(body)), Montnegre Natu-

ral Park, Sant Celoni, Barcelona (Spain), 610 m above sea level, lat/long coordinates N41.66449 E2.56279, on

soil litter under oak trees, hand collecting, 18.iv.2007, leg. E. Mateos. Paratypes: 8 specimens on slides (3 males, 1

female, and 4 without visible sexual plate) and 17 specimens preserved in absolute alcohol, same data as holotype.

Holotype and paratype slides CRBA-90745a and CRBA-90745b saved in the collection of the Centre de Recursos

de Biodiversitat Animal, Faculty of Biology, University of Barcelona (http://www.crba.ub.edu); other paratypes

kept in the E. Mateos’ collection (lot LP125).

Other material. 40 specimens preserved in absolute alcohol, Montnegre Natural Park, Sant Celoni, Barcelona

(Spain), 427 m above sea level, lat/long coordinates N41.6663 E2.5793, on soil litter under oak trees, 18.iv.2007,

leg. E. Mateos. All material kept in the E. Mateos’ collection (lot LP134).

FIGURE 8. Lepidocyrtus fuscocephalus sp. nov.: Habitus lateral (specimen in alcohol, without scales).

MATEOS & ÁLVAREZ-PRESAS

Diagnosis. With dark blue pigment on all head, Ant.II–IV, coxae-trocanter-femur of legs I–III, and ventral tube;

disperse blue pigment also on Abd.IV and Abd.V. Th.II slightly projecting over the head. Ant.I–III, legs, ventral tube

and posterior region of manubrium with scales. Labial chaetotaxy M1M2REL1L2, R shortened. Dorsal cephalic and

body macrochaetae formula A0[A2a]A2A3Pa5/00/0101+3. Abd.IV without chaeta s. Unguiculus lanceolate and with

serrated outer margin.

Molecular diagnosis. This species includes all populations that cluster with CoxII and EF sequences of the

individuals LP125-1 to LP125-5 (Table 1), with significant support in an adequate molecular delimitation model.

Etymology. The species name refers to the dark blue colour of the head. In latin “fuscus” means dark, and

“cephalus” means head.

Description. Holotype body length (without head nor furca) 2.2 mm, paratypes 1.8–2.2 mm. Body colour pat-

tern (Fig. 8) with dark blue pigment on all head, Ant.II–IV (with increasing colour intensity towards the distal part

of each segment), coxae, and ventral tube; disperse blue pigment also on trochanter-femur of legs I–III, Abd.IV and

Abd.V; densely black pigmented ocular areas. Mesothorax slightly projected over the head.

Antenna with scales on dorsal and ventral region of Ant.I–III. Ratio antenna:cephalic diagonal = 1.8–1.9 (head

diagonal measured from cervical edge to apex of mouth part); ratio Ant.I:II:III:IV as 1:1.7:1.6:2.5. Basis of Ant.I

dorsally with three microchaetae arranged in triangle (Ant.I-organ); apex of Ant.I with a short curved S-chaeta in the

membranous area of the ventral region. Ant.III organ composed of two subcilindrical and curved sensory rods. Ant.

IV without apical bulb. Eyes 8+8; eyes A to F of equal size, G and H slightly smaller, ratio A/G and A/H = 1.6.

FIGURES 9–12. Lepidocyrtus fuscocephalus sp. nov.: 9, clypeal and labral chaetotaxy; 10, maxillary palp (right side); 11, outer

labial papilla (right side); 12, labial and postlabial chaetotaxy (the arrow points to a supernumerary M chaeta present in one

paratype). Black dots––ciliated chaetae, cross––scales, p.c.—proximal chaetae of labial palp.

FIGURE 13. Lepidocyrtus fuscocephalus sp. nov.: Dorsal head chaetotaxy (left side). Broad circles––long ciliated macrocha-

etae, small circles––short ciliated macrochaetae.

MATEOS & ÁLVAREZ-PRESAS

FIGURES 14–16. Lepidocyrtus fuscocephalus sp. nov. dorsal chaetotaxy (left side): 14, Th.II; 15, Th.III; 16 Abd.I. Circles––

ciliated chaetae.

FIGURES 17–18. Lepidocyrtus fuscocephalus sp. nov. dorsal chaetotaxy (left side): 17, Abd.II; 18, Abd.III. Broad circles––

broad ciliated macrochaetae, small circles––thin ciliated macrochaetae.

Clypeus (Fig. 9) with three prefrontal chaetae (1 pf0 and 2 pf1), four facial chaetae (f), and four lateral chaetae

(2 L1 and 2 L2), all these chaetae ciliated. Prelabral and labral chaetae in typical number 4/554 (Fig. 9), prelabral

chaetae ciliated, all labral chaetae smooth and pointed, apical row curved; inverted U-shaped labral apical intrusion;

four rounded labral papillae with a central small pointed expansion. Maxillary palp outer lobe with smooth apical

appendage and basal chaeta, and three smooth sublobal appendages (Fig. 10). Lateral process of outer labial papilla

finger-shape, slightly curved, tip not reaching apex of papilla (Fig. 11).

Labial and postlabial chaetotaxy as in Fig. 12; with five smooth proximal chaetae at the base of labial palp;

labial anterior row formed by five smooth chaetae (a1–a5); posterior row formed by ciliated chaetae with formula

M1M2REL1L2; chaeta R shorter, ratio M2/R ≈ 2.4; one paratype with a supernumerary reduced M chaeta (as long

as chaeta R) between M1 and M2 present on one side (Fig. 12); postlabial chaetaxy with all chaetae ciliated, row I

(along ventral cephalic groove) with 4 chaetae.

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 15+15. Interocular chaetotaxy with s, t, p

ciliated chaetae and 5–7 scales (Fig. 13).

MATEOS & ÁLVAREZ-PRESAS

FIGURE 19. Lepidocyrtus fuscocephalus sp. nov.: Abd.IV chaetotaxy (left side). Broad black circles––broad ciliated macro-

chaetae, small black circles––thin ciliated macrochaetae.

FIGURES 20–22. Lepidocyrtus fuscocephalus sp. nov.: 20, Abd.V chaetotaxy (left side), broad circles––long ciliated chaetae,

small circles––short ciliated or smooth chaetae; 21, trochanteral organ; 22, third leg unguis and unguiculus.

Dorsal body macrochaetae formula 00/0101+3 (macrochaetae m3 on Abd.II, and Sm+B4, B5, B6 on Abd.IV).

Dorsal chaetotaxy of Th.II–III and Abd.I as in Figs 14–16. Th.II with 2 lateral S-chaetae (al and ms) and without

macrochaetae 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 17–18. Abd.II chaeta ml present or

absent depending on the specimens; macrochaeta m3 with socket diameter 1.6 times higher than macrochaeta m5.

Abd. III chaeta mi present or absent depending on the specimens, with chaeta d3 between macrochaetae pm6 and

p6, and with S-chaetae as and ms. All chaetae associated with the trichobothria on Abd.II–III strongly ciliate and

partially fan-shaped. Chaetotaxy of Abd.IV as in Fig. 19; macrochaetae Sm, B4, B5, B6, D3, E2, E3, E4, F1, F2,

F3 broader and with broad socket; macrochaetae T6, T7, D2, De3, E1, E4p, Fe4, Fe5, F3p shorter or longer but

always thinner and with socket of minor diameter; macrochaeta F2 inserted above macrochaeta E3; the ratio of

distances between macrochaetae Sm–B4/B4–B6 is 0.8–0.9; the ratio of distances between macrochaetae B4–B5/

B5–B6 is 1.1–1.3; accessory chaeta s associated with trichobotrium T2 absent; chaetae D1, m, pe and pi associated

with trichobotria T2 and T4 fan-shaped and strongly ciliate, chaeta a pointed and strongly ciliate; sens chaetotaxy

composed of 2 anterior dorsomedial elongate S-chaetae, and short chaetae as and ps; posterior margin with 9+9

smooth mesochaetae; lateral region BP4 without pseudopori. Dorsal chaetotaxy of Abd.V with S-chaetae as, acc.p4

and acc.p5 (Fig. 20).

Legs with scales except in claws. Rectangular-shaped trochanteral organ formed by a maximum of 28 smooth

straight chaetae (Fig. 21). Unguis (Fig. 22) with basal pair of teeth at 49% from base of the inner edge, and with

two inner unpaired teeth at 71% and 88% from the base of the inner edge respectively; on unguis I–II the apical in-

ner tooth tiny and sometimes difficult to see; one external tooth and a pair of lateral teeth also present. Unguiculus

lanceolate with serrated outer margin (serration less evident on unguiculus III). Tibiotarsal tenent hair spatulate,

smooth and as long as claw; ratio of supra-empodial chaeta (smooth chaeta on tibiotarsus III opposite to tenent hair)

/ unguiculus ≈ 1.2.

Ventral tube with 6+6 ciliated chaetae on anterior side (4+4 proximal and 2+2 distal) and 11+11 weakly ciliated

chaetae on posterior side; scales present on anterior and posterior sides; lateral flap with a maximum of 26 latero-

distal chaetae (20–23 ciliated and 3 smooth).

Manubrium with scales on anterior and posterior surfaces, with 2+2 ciliated apical chaetae on anterior side.

The ratio manubrium:dens:mucro is 19:20:1. Manubrial plate with 3–4 inner chaetae and a maximum of 16 outer

chaetae. Dental tubercle absent. Mucro with the two teeth of the same size, without spinelet on basal spine.

Pseudopores distribution on dorsal and ventral positions as in Figs 23a–b.

Ecology and distribution. All specimens were obtained by sifting soil surface litter under oak trees.

Discussion. Morphological characters clearly assign Lepidocyrtus fuscocephalus sp. nov. to the Lepidocyrtus

lignorum-group (sensu Mateos 2011). By the characteristic body colour pattern L. fuscocephalus sp. nov. clearly

differs from all the other species of the L. lignorum-group. Other differences between all species included in the

group are summarized in Table 2. By having body partially blue pigmented and lanceolate unguiculus L. fuscoceph-

alus sp. nov. is close to species L. instratus, L. labyrinthi, L. traseri, L. milagrosae sp. nov. and L. semicoloratus

MATEOS & ÁLVAREZ-PRESAS

sp. nov. Of these, the new species differ by the colour pattern, the presence of scales on Ant.III and pointed labral

apical chaetae.

FIGURE 23. Lepidocyrtus fuscocephalus sp. nov, L. milagrosae sp. nov. and L. semicoloratus sp. nov.: Pseudopores distribu-

tion: a, dorsal pseudopores; b–c, ventral pesudopores.

TABLE 2. Characters differentiating between the European Lepidocyrtus species of lignorum-group.

(+)––present; (-)––absent; ?––no data available; B––broad ciliated macrochaeta; band–– body white with some datk blue bands; bl––body dark blue pigmented; dots––body white

with some dark blue spots; part––body partially dark blue pigmented; wh––body white; T––thin ciliated macrochaeta; Ceph: cephalic chaetae; Labial M: number of labial M

chaetae; Labial R: number of labial R chaetae; Labrum apical: morphology of apical labral chaetae, bra––chaeta with branched apical end, poi––chaeta with pointed apical end;

Labrum papilla: number of denticles in the labral papillae; BP4 pse: presence/absence of pseudopores on the basal plate of fourth abdominal segment; Unguis bp%: basal pair of

teeth position on the inner edge of unguis-III measured in percentage from base of this inner edge; Unguis unp: number of unpaired teeth on the inner edge of unguis-III; Unguicu-

lus: morphology of unguiculi apex, acu––acuminate, tru––truncate. 1Data from Mateos et al. (2021). *new species.

Body colour

Ant.III scales

Ceph A3

Ceph Pa5

Interocular q

Labial M

Labial R

Labrum apical

Labrum papilla

Abd.IV T6

Abd.IV B6

BP4 pse

Unguis bp%

Unguis unp

Unguiculus

barbulus wh/bl + + + - 3–4 2–5 poi 4 T B - 50% 2 acu

chorus dots - + + - 2 1 bra 1–3 B B - 54% 2 acu

fuscocephalus*part + + + - 2 1 poi 1 T B - 49% 2 acu

instratus part - + + ? 2 1 ? ? T B - 40% 1 acu

intermedius wh - - + - 2 1 bra 3 T B - 56% 2 acu

juliae dots - + + + 2 1 bra 3 T B + 52% 2 acu

labyrinthi part - + + - 2 1 bra 1 T B - 50% 2 acu

lignorum wh - + + - 2 1 bra 3 T B +/-150% 1–2 acu

milagrosae*part - + + - 2 1 bra 3 T B + 48% 2 acu

paralignorum wh - + + - 2 1 bra 1–3 T B - 50% 2 acu

peisonis wh +/- + + - 2 1 poi 0 T B - 54% 2 tru

pulchellusband + + + - 2–3 1 bra 2–3 T B - 50% 2 tru

ruber wh/bl - + - ? 2 1 poi 0 T B - 60% 1 tru

semicoloratus*part - + + - 2 1 bra 2 T B - 50% 2 acu

tellecheae wh + + + - 2 1 poi 1 T B - 50% 1 acu

traseri part - + + - 2 1 poi/bra 1 T T - 52% 2 acu

uzeli bl - + + ? 2 1 poi 1 T B - 50% 0 tru

vexillosus dots ? - + ? 2 1 ? ? T B - 60% 1–2 acu

violaceus bl - + + - 2 1 bra 1–3 T B/T1+/-150% 2 acu

MATEOS & ÁLVAREZ-PRESAS

Lepidocyrtus milagrosae Mateos sp. nov.

Figs 24–40, Tabs 1–2

Zoobank: urn:lsid:zoobank.org:act:1AE4D1FD-A06D-491B-A9EA-47AB98D233B1

Type material. Holotype: male on slide (CRBA-90746), Laerma, Rhodes (Greece), 253 m above sea level, lat/long

coordinates N36.146639 E27.920374, on herbaceous vegetation and soil litter, hand collecting, 3.iv.2009, leg. E.

Mateos. Paratypes: 9 specimens without visible sexual plate on slides and 9 specimens preserved in absolute al-

cohol, same data as holotype. Holotype and paratype slide CRBA-90747 saved in the collection of the Centre de

Recursos de Biodiversitat Animal, Faculty of Biology, University of Barcelona (http://www.crba.ub.edu); other

paratypes kept in the E. Mateos’ collection (lot LP250).

Other material. 8 specimens preserved in absolute alcohol, Butterfly Valley, Petaloudes, Rhodes (Greece), 225

m above sea level, lat/long coordinates N36.337087 E28.062473, on herbaceous vegetation (lot LP248) and soil

litter (lot LP246), 3.iv.2009, leg. E. Mateos. All material kept in the E. Mateos’ collection.

Diagnosis. With dark blue pigment on the dorsal and ventral sides of Th.II to Abd.III, Ant.II–IV, and cx.I–III.

Th.II slightly projecting over head. Ant.I–II, legs, ventral tube and posterior region of manubrium with scales. Api-

cal bulb on Ant.IV absent. Labial chaetotaxy M1M2REL1L2, R slightly shortened. Dorsal cephalic and body mac-

rochaetae formula A0[A2a]A2A3Pa5/00/0101+3. Abd.IV without chaeta s and with 4–6 lateral pseudopori on BP4.

Unguiculus lanceolate and with serrated (or finely serrated) outer margin.

Molecular diagnosis. This species includes all populations that cluster with CoxII and EF sequences of the

individuals LP250-4 to LP250-7 (Table 1), with significant support in an adequate molecular delimitation model.

Etymology. The species is named after author’s wife Milagros (in apposition).

Description. Holotype body length (without head nor furca) 1.2 mm, paratypes 1.0–1.2 mm. Body colour pat-

tern (Fig. 24) with dark blue pigment on the dorsal and ventral sides of Th.II to Abd.III (including ventral tube),

Ant.II–IV (with increasing colour intensity towards the distal part of each segment), and cx.I–III; densely black

pigmented ocular areas. Mesothorax slightly projected over the head.

FIGURE 24. Lepidocyrtus milagrosae sp. nov.: Habitus lateral (specimen in alcohol, without scales).

Antenna with scales on dorsal region of Ant.I–II. Ratio antenna:cephalic diagonal = 1.4–1.6 (head diagonal

measured from cervical edge to apex of mouth part); ratio Ant.I:II:III:IV as 1:1.8:1.8:2.8–3.2. Basis of Ant.I dorsally

with three microchaetae arranged in triangle (Ant.I-organ); apex of Ant.I with a short curved S-chaeta in the mem-

branous area of the ventral region. Ant.III organ composed of two subcilindrical and curved sensory rods. Ant.IV

without apical bulb. 8+8 eyes; eyes A–F of equal size, eyes G and H a little bit smaller, ratio A/F and A/G = 1.3.

Clypeus (Fig. 25) with three prefrontal chaetae (1 pf0 and 2 pf1), facial area with four chaetae and three scales,

and four lateral chaetae (2 L1 and 2 L2), all these chaetae ciliated. Prelabral and labral chaetae in typical number

4/554 (Fig. 25), prelabral chaetae ciliated, first and second rows of labral chaetae smooth, apical row of labral

chaetae branched (Fig. 26); inverted U-shaped labral apical intrusion; four rounded labral papillae with three small

pointed expansions (Fig. 26). Maxillary palp outer lobe with smooth basal chaeta and apical appendage; sublobal

plate with three smooth appendages and a smaller fourth one (Fig. 27). Lateral process of outer labial papilla finger-

shape, slightly curved, tip not reaching apex of papilla (Fig. 28).

FIGURES 25–29. Lepidocyrtus milagrosae sp. nov.: 25, clypeal and labral chaetotaxy; 26, apical labral chaetae and papillae;

27, maxilary palp (right side); 28, outer labial papilla (right side); 29, labial and postlabial chaetotaxy (right side).

Labial and postlabial chaetotaxy as in Fig. 29; with five smooth proximal chaetae at the base of labial palp;

labial anterior row formed by five smooth chaetae (a1–a5); posterior row formed by ciliated chaetae with formula

M1M2REL1L2; chaeta R shorter, ratio M/R ≈ 2.3; postlabial chaetotaxy with all chaetae ciliated, row I (along ven-

tral cephalic groove) with 4 chaetae.

Dorsal cephalic macrochaetae formula A0A2A3Pa5, but also with pair of smaller supplementary macrochaetae

A2a between A0 and A2, this chaeta A2a can have two different morphologies depending on the specimens (Fig. 30) ;

maximum number of macrochaetae An on head 11+11 (Fig. 30). Interocular chaetotaxy with s, t, p ciliated chaetae

and 2–3 scales.

Dorsal body macrochaetae formula 00/0101+3 (macrochaetae m3 on Abd.II, and Sm+B4, B5, B6 on Abd.IV).

MATEOS & ÁLVAREZ-PRESAS

Dorsal chaetotaxy of Th.II–III and Abd.I as in Figs 31–33. Th.II with 2 lateral S-chaetae (al and ms) and without

macrochaetae 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 34–35. Abd.II chaeta ml present

or absent depending on the specimens, chaeta p5p absent; macrochaetae m3 and m5 with equal socket diameter.

Abd. III chaeta mi present or absent depending on the specimens, chaeta d3 absent, with S-chaetae as and ms, one

specimen with a supernumerary associate ciliated chaeta between ll and a6 on Abd.III trichobothria a5. All chaetae

associated with the trichobothria on Abd.II–III acuminate and strongly ciliate. Chaetotaxy of Abd.IV as in Fig. 36;

macrochaetae Sm, B4, B5, B6, D3, E2, E3, E4, F1, F2, F3 broader and with broad socket; macrochaetae T6, T7,

D2, De3, E1, E4p, Fe4, F3p shorter or longer but always thinner and with socket of minor diameter; macrochaeta

F2 inserted above macrochaeta E3; the ratio of distances between macrochaetae Sm–B4/B4–B6 is 0.6–0.8; the ratio

of distances between macrochaetae B4–B5/B5–B6 is 1.0–1.4; accessory chaeta s associated with trichobotrium T2

absent; chaetae a, D1, m, pe and pi associated with trichobotria T2 and T4 acuminate and strongly ciliate; sens

chaetotaxy composed of one anterior dorsomedial elongate S-chaetae, and short chaetae as and ps; posterior margin

with 5+5 smooth mesochaetae; lateral region with 4–7 small pseudopori on BP4 (Fig. 37). Dorsal chaetotaxy of

Abd.V with S-chaetae as, acc.p4 and acc.p5 (Fig. 38).

FIGURE 30. Lepidocyrtus milagrosae sp. nov.: Dorsal head chaetotaxy (left side). On the right side, the arrangement and mor-

phology of the apical chaetae is drawn (note the different morphology that chaeta A2a may present). Broad circles––long ciliated

macrochaetae, small circles––short ciliated macrochaetae.

FIGURES 31–33. Lepidocyrtus milagrosae sp. nov. dorsal chaetotaxy (left side): 31, Th.II; 32, Th.III; 33, Abd.I. Circles––cili-

ated chaetae.

MATEOS & ÁLVAREZ-PRESAS

FIGURES 34–35. Lepidocyrtus milagrosae sp. nov. dorsal chaetotaxy (left side): 34, Abd.II; 35, Abd.III. Broad circles––broad

ciliated macrochaetae, small circles––thin ciliated macrochaetae.

Legs with scales except in claws. V-shaped trochanteral organ formed by a maximum of 9 smooth straight

chaetae (Fig. 39). Unguis with basal pair of teeth at 48% from base of the inner edge, and with two inner unpaired

teeth at 62% (the bigger) and 81% from base of the inner edge respectively; one external tooth and a pair of lateral

teeth also present. Unguiculus lanceolate with smooth or finely serrated outer margin. Tibiotarsal tenent hair spatu-

late and smooth (Fig. 40); ratio of tibiotarsal tenant hair / unguis inner edge ≈ 1.3; ratio of supra-empodial chaeta /

unguiculus ≈ 1.

Ventral tube with 6+6 ciliated chaetae on anterior side (4+4 proximal, 2+2 distal) and 10+10 weakly ciliated

chaetae on posterior side; scales present only on anterior side; lateral flap with a maximum of 12 laterodistal chaetae

(7 ciliated and 5 smooth).

Manubrium with scales on anterior and posterior surfaces, with 2+2 ciliated apical chaetae on anterior side. The

ratio manubrium:dens:mucro is 17:19:1. Manubrial plate with 2–3 inner chaetae and a maximum of 8 outer chaetae.

Dental tubercle absent. Mucro with the two teeth of the same size, without spinelet on basal spine.

Pseudopores distribution on dorsal and ventral positions as in Figs 23a,c.

Ecology and distribution. All specimens were obtained by beating the herbaceous vegetation and sifting soil

surface litter under holm oak trees.

FIGURES 36–37. Lepidocyrtus milagrosae sp. nov.: 36, Abd.IV chaetotaxy (left side); 37, basal plate of Abd.IV. Broad black

circles––broad ciliated macrochaetae, small black circles––thin ciliated macrochaetae, crossed circles––pseudoporus.

MATEOS & ÁLVAREZ-PRESAS

FIGURES 38–40. Lepidocyrtus milagrosae sp. nov.: 38, Abd.V chaetotaxy (left side), broad black circles––long ciliated cha-

tae, small black circles––short ciliated or smooth chaetae; 39, trochanteral organ; 40, third leg unguis and unguiculus with three

different possibilities of unguiculus outer margin.

Discussion. Morphological characters clearly assign Lepidocyrtus milagrosae sp. nov. to the Lepidocyrtus lig-

norum-group (sensu Mateos 2011). By the characteristic body colour pattern L. milagrosae sp. nov. clearly differs

from all the other species of the L. lignorum-group except L. labyrinthi. These two species differ by the morphology

of labral papillae (with three pointed expansions in the new species and only one in L. labyrinthi) and the presence

of pseudopores on BP4 in the new species (Table 2). Other characters (not included in Table 2) differentiating L.

labyrinthi and the new species are the absence of chaetae a5, m5, and p6 on Abd.I, and the presence of chaetae p5p

on Abd.II and d3 on Abd.III in L. labyrinthi (Baquero et al. 2021). By having body partially blue pigmented and

lanceolate unguiculus L. milagrosae sp. nov. is close to species L. instratus, L. traseri, L. fuscocephalus sp. nov.,

and L. semicoloratus sp. nov. Of these, the new species differ by the colour pattern and the presence of pseudopores

on BP4. Other differences between all species included in the group are summarized in Table 2.

Lepidocyrtus semicoloratus Mateos sp. nov.

Figs 41–56, Tabs 1–2

Zoobank: urn:lsid:zoobank.org:act:4C8EF42F-8A89-4A7E-AAE5-5B23D310C405

Type material. Holotype: Female on one slide (CRBA-90748), Montseny Natural Park, Aiguafreda, Barcelona

(Spain), 917 m above sea level, lat/long coordinates N41.7892 E2.3113, on herbaceous vegetation, 10.ii.2007, leg.

E. Mateos. Paratypes: 6 specimens (1 male, 2 female, and 3 without visible sexual plate) on slides and 18 specimens

preserved in absolute alcohol, same data as holotype. Holotype and paratype slide CRBA-90749 saved in the col-

lection of the Centre de Recursos de Biodiversitat Animal, Faculty of Biology, University of Barcelona (http://www.

crba.ub.edu); other paratypes kept in the E. Mateos’ collection (lot LP108).

Other material. 5 specimens on slides and 1 specimen preserved in absolute alcohol, Las Hoyuelas, Sinarcas,

Valencia (Spain), 840 m above sea level, lat/long coordinates N39.765328 W1.226981, on herbaceous vegetation,

hand collecting, 02.iv.2007, leg. E. Mateos, all material kept in the E. Mateos’ collection (lot LP122).

Diagnosis. With dark blue pigment on Ant.II–IV, dorsal and ventral sides of Th.III to Abd.III (and with pale

pigmentation on Abd.IV–V), coxae I–III, and ventral tube. Th.II sligthly projecting over head. Ant.I–II, legs, ventral

tube and posterior region of manubrium with scales. Labial chaetotaxy M1M2REL1L2, R shortened. Dorsal cephalic

and body macrochaetae formula as A0[A2a]A2A3Pa5/00/0101+3. Abd.IV without chaeta s. Unguiculus lanceolate

and with serrated outer margin.

Molecular diagnosis. This species includes all populations that cluster with CoxII and EF sequences of the

individuals LP108-1 to LP108-5 (Table 1), with significant support in an adequate molecular delimitation model.

Etymology. The species name refers to the dark blue colour present only on part of the body. In Latin “semi”

means half, and “coloratus” means pigmented.

FIGURE 41. Lepidocyrtus semicoloratus sp. nov.: Habitus lateral (specimen in alcohol, without scales).

Description. Holotype body length (without head nor furca) 1.5 mm, paratypes 1.5–1.8 mm. Body colour pat-

tern (Fig. 41) with dark blue pigment on the dorsal and ventral sides of Th.III to Abd.III (and with pale pigmentation

on Abd.IV–V), Ant.II–IV (on Ant.II only in apical portion), coxae I-III, and ventral tube; densely black pigmented

ocular areas. Mesothorax slightly projected over the head.

Antenna with scales on dorsal region of Ant.I–II. Ratio antenna:cephalic diagonal = 1.8–1.9, ratio Ant.I:II:III:

IV as 1:1.8:1.8:2.7. Basis of Ant.I dorsally with three microchaetae arranged in triangle (Ant.I-organ); apex of Ant.I

with a short curved S-chaeta in the membranous area of the ventral region. Ant.III organ composed of two subcilin-

drical and curved sensory rods. Ant.IV without apical bulb. 8+8 eyes; eyes A–F of equal size, eyes G and H a little

bit smaller, ratio A/F and A/G = 1.2.

Clypeus (Fig. 42) with three prefrontal chaetae (1 pf0 and 2 pf1), five facial chaetae (central one shorter), and

four lateral chaetae (2 L1 and 2 L2), all these chaetae ciliated. Prelabral and labral chaetae in typical number 4/554

(Fig. 42), prelabral chaetae ciliated, labral setae smooth, apical row branched (Fig. 43), inverted U-shaped labral

apical intrusion, four rounded labral papillae with a central very small pointed expansion (difficult to see in several

specimens, Fig 43). Maxillary palp outer lobe with smooth apical appendage and basal chaeta, and three smooth

sublobal appendages (Fig. 44). Lateral process of outer labial papilla finger-shape, slightly curved, tip not reaching

apex of papilla (Fig. 45).

Labial and postlabial chaetotaxy as in Fig. 46; with five smooth proximal chaetae at the base of labial palp;

labial anterior row formed by smooth chaetae (a1–a5), posterior row formed by ciliated chaetae with formula

M1M2REL1L2; chaeta R half in length of chaeta M2, ratio M/R ≈ 2; postlabial chaetotaxy with all chaetae ciliated,

row I (along ventral cephalic groove) with 4 chaetae.

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 (Fig. 47). Interocular chaetotaxy with

s, t, p ciliated chaetae and 2–3 scales..

Dorsal body macrochaetae formula 00/0101+3 (macrochaetae m3 on Abd.II, and Sm+B4, B5, B6 on Abd.IV).

Dorsal chaetotaxy of Th.II–III and Abd.I as in Figs 48–50. Th.II with 2 lateral S-chaetae (al and ms) and without

macrochaetae 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 51–52. Abd.II chaeta ml present or

absent depending on the specimens, chaeta p5p present; macrochaeta m3 with socket diameter 1.3 times higher

than macrochaeta m5. Abd. III chaetae mi and d3 present in all specimens, with S-chaetae as and ms. All chaetae

MATEOS & ÁLVAREZ-PRESAS

associated with the trichobothria on Abd.II–III strongly ciliated and partially fan-shaped (widened in the center).

Chaetotaxy of Abd.IV as in Fig. 53; macrochaetae Sm, B4, B5, B6, D3, E2, E3, E4, F1, F2, F3 broader and with

broad socket; macrochaetae T6, T7, D2, De3, E1, E4p, Fe4, F3p shorter or longer but always thinner and with

socket of minor diameter; macrochaeta F2 inserted above macrochaeta E3; the ratio of distances between macro-

chaetae Sm–B4/B4–B6 is 0.7–0.9; the ratio of distances between macrochaetae B4–B5/B5–B6 is 1.3–1.8; accessory

chaeta s associated with trichobotrium T2 absent; chaetae a, D1, m, pe and pi associated with trichobotria T2 and

T4 strongly ciliate and widened in the center; sens chaetotaxy composed of 2 anterior dorsomedial elongate S-chae-

tae, and short chaetae as and ps; posterior margin with 5+5 smooth mesochaetae; lateral region without pseudopori

on BP4. Dorsal chaetotaxy of Abd.V with S-chaetae as, acc.p4 and acc.p5 (Fig. 54).

Legs with scales except in claws. V-shaped trochanteral organ formed by a maximum of 15 smooth straight

chaetae (Fig. 55). Unguis with basal pair of teeth at 50% from base of the inner edge, and with two inner unpaired

teeth at 64% and 82% from base of the inner edge respectively; one external tooth and a pair of lateral teeth also

present. Unguiculus lanceolate with finely serrated outer margin. Tibiotarsal tenent hair spatulate and smooth (Fig.

56); ratio of tibiotarsal tenant hair / unguis inner edge ≈ 1; ratio of supra-empodial chaeta / unguiculus ≈ 1.

Ventral tube with 6+6 ciliated chaetae on anterior side (4+4 proximal, 2+2 distal) and 11+11 weakly ciliated

chaetae on posterior side; scales present on anterior and posterior sides; lateral flap with a maximum of 23 latero-

distal chaetae (17–21 ciliated and 2 smooth).

FIGURES 42–46. Lepidocyrtus semicoloratus sp. nov.: 42, clypeal and labral chaetotaxy; 43, apical labral chaetae and papil-

lae; 44, maxillary palp (right side); 45, outer labial papilla (right side); 46, labial and postlabial chaetotaxy (right side). Black

dots––ciliated chaetae, cross––scales.

FIGURE 47. Lepidocyrtus semicoloratus sp. nov.: Dorsal head chaetotaxy (left side). Broad circles––long ciliated macrocha-

etae, small circles––short ciliated macrochaetae.

MATEOS & ÁLVAREZ-PRESAS

FIGURES 48–50. Lepidocyrtus semicoloratus sp. nov. dorsal chaetotaxy (left side): 48, Th.II, 49; Th.III; 50, Abd.I. Circles––

ciliated chaetae.

FIGURES 51–52. Lepidocyrtus semicoloratus sp. nov. dorsal chaetotaxy (left side): 51, Abd.II; 52, Abd.III. Broad circles––

broad ciliated macrochaetae, small circles––thin ciliated macrochaetae.

Manubrium with scales on anterior and posterior surfaces, with 2+2 ciliated apical chaetae on anterior side. The

ratio manubrium:dens:mucro is 15:17:1. Manubrial plate with 3 inner chaetae and a maximum of 9 outer chaetae.

Dental tubercle absent. Mucro with the two teeth of the same size, without spinelet on basal spine.

Pseudopores distribution on dorsal and ventral positions as in Figs 23a–b.

Ecology and distribution. All specimens were obtained by beating the herbaceous vegetation.

Discussion. Morphological characters clearly assign Lepidocyrtus semicoloratus sp. nov. to the Lepidocyrtus

lignorum-group (sensu Mateos 2011). By the characteristic body colour pattern, L. semicoloratus sp. nov. clearly

differs from all the other species of the L. lignorum-group except L. instratus, with which the new species shares the

same colour pattern design. With the current information about L. instratus, the only characters useful for differenti-

ating this species and L. semicoloratus sp. nov. are 1) the dark pigmented buccal area in L. instratus (not pigmented

in the new species), 2) the size and position of the basal pair teeth on the inner edge of the unguis (tiny and at 40%

in L. instratus vs. higher and at 50% in the new species), and 3) the shape of the unguiculi (smooth in L. instratus vs.

MATEOS & ÁLVAREZ-PRESAS

FIGURE 53. Lepidocyrtus semicoloratus sp. nov.: Abd.IV chaetotaxy (left side). Broad black circles––broad ciliated macro-

chaetae, small black circles––thin ciliated macrochaetae.

FIGURES 54–56. Lepidocyrtus semicoloratus sp. nov.: 54, Abd.V chaetotaxy (left side), broad circles––long ciliated chaetae,

small circles––short ciliated or smooth chaetae; 55, trochanteral organ; 56, third leg unguis and unguiculus.

serrated in the new species). On the other hand, the type locality and the habitat of both species are so different that

they could hardly be the same species. L. instratus comes from the Swiss Alps, at altitudes above 1600 m above

sea level, and inhabiting under stones in snowy areas and in groundhog droppings. L. semicoloratus sp. nov. comes

from Eastern Spain (with marked Mediterranean climate), at altitudes below 1000 m above sea level, inhabiting

herbaceous vegetation (although it must also live in the soil and under stones) in areas dominated by holm-oak and

pine forests. Other differences between all species included in the lignorum-group are summarized in Table 2.

The three molecular species delimitation analyses carried out (Fig. 1) suggest that the two populations studied

of this new species (populations LP108 and LP122) may represent two different species. The detailed morpho-

logical study carried out with the specimens of these two populations has not detected any relevant difference that

allows assigning each population to a different species. For this reason, we consider both populations as the same

morphological species. The geographical distance that separates the two populations is about 375 km, so it is very

likely that there has been no contact between them for quite some time, which would explain the genetic differentia-

tion found between both populations.

References

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Three new species of Collembola from Sicily with first record of the genus Superodontella Stach, 1949 from Italy

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ZOOTAXA

Three new species of soil Collembola from both agricultural lands (vineyards and olive orchards) and a natural habitat (beech forest) located in Sicily (Italy) are described: Superodontella eleonorae sp. nov., first record of this genus for Italy; Lepidocyrtus rapitalai sp. nov. and Pseudosinella francae sp. nov..

Distinctive Collembola Communities in the Mesovoid Shallow Substratum: Entomobryomorpha of the Sierra de Guadarrama National Park (Central Spain)

Article

Full-text available

Jan 2021

The material for this study was obtained after intensive sampling in the colluvial mesovoid shallow substratum, or MSS, of the Sierra de Guadarrama National Park using 33 subterranean sampling devices (SSD). The data were obtained from the first extraction of the traps between May and October of 2015. This paper presents the results for the Entomobryomorpha Börner, 1913, which was part of the Collembola captured. Four families and 12 genera have been studied: Isotomidae Schäffer, 1896 (Folsomia Willem, 1902, Tetracanthella Schött, 1891, Uzelia Absolon, 1901, Folsomides Stach, 1922, Isotomurus Börner, 1903, Parisotoma Bagnall, 1940, Pseudisotoma Handschin, 1924 and Pachyotoma Bagnall, 1949), Orchesellidae Börner, 1906 (Orchesella Templeton, 1835 and Heteromurus Wankel, 1860), Entomobryidae Schäffer, 1896 (Entomobrya Rondani, 1861) and Lepidocyrtidae Wahlgren, 1906 (Lepidocyrtus Bourlet, 1839 and Pseudosinella Schäffer, 1897). The species of Orchesella were studied in a previous paper (Baquero et al. 2017). The richness of the habitat sampled is defined by twenty-one species, eight of which are new: Pachyotoma penalarensis Baquero & Jordana n. sp., Entomobrya guadarramensis Jordana & Baquero n. sp., Entomobrya ledesmai Jordana & Baquero n. sp., Lepidocyrtus labyrinthi Baquero & Jordana n. sp., Lepidocyrtus paralignorum Baquero & Jordana n. sp., Lepidocyrtus purgatori Baquero & Jordana n. sp., Pseudosinella valverdei Baquero & Jordana n. sp. and Pseudosinella gonzaloi Baquero & Jordana n. sp. Entomobrya intermedia Brook, 1884 (England) is discussed and a new name Entomobrya katzi Jordana & Baquero n. sp. is proposed for E. intermedia sensu Katz et al. (2015) based on the American specimens.

Corrigendum to: IQ-TREE 2: New Models and Efficient Methods for Phylogenetic Inference in the Genomic Era

Article

Full-text available

Jun 2020

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

Article

Full-text available

Jun 2020
ZOOTAXA

Eduardo Mateos

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.

New Insight into the Systematics of European Lepidocyrtus (Collembola: Entomobryidae) Using Molecular and Morphological Data

Article

Full-text available

May 2020

The Collembolan genus Lepidocyrtus is subdivided into up to eight subgenera, of which only Lepidocyrtus s.str. (Bourlet, 1839) and Lanocyrtus (Yoshii & Suhardjono, 1989) are represented by European species. The discovery of unique characters in the European species Lepidocyrtus tomosvaryi (rounded dental tubercle) and L. peisonis (lateral tuft of long filiform chaetae in abdomen III) has only described so far for species of the subgenera Setogaster (Salmon, 1951) and Cinctocyrtus (Yoshii & Suhardjono, 1989) and has raised the need to perform a molecular analysis by involving other representative species of the genus. For this study, phylogenetic analysis of 15 Lepidocyrtus species occurring in the Carpathian Basin were carried out. The analyses, which was based on both concatenated datasets of COII and EF1-α sequences and individual gene sequences, clearly placed L. tomosvaryi within the subgenus Lanocyrtus and L. peisonis within Lepidocyrtus s.srt. European species groups defined on the basis of morphological characters were only partly confirmed by the concatenated and COII analyses because of the splitting of the pallidus–serbicus-group, whereas EF1- α sequences weakly supported this group.

New insights into the systematics of Entomobryoidea (Collembola: Entomobryomorpha): first instar chaetotaxy, homology and classification

Article

Full-text available

Oct 2019

The largest superfamily Entomobryoidea is one of the key groups in Collembola. However, incorrect recognition of chaetotaxic homology within Entomobryoidea severely impedes the accurate species comparison of adults and phylogenetic reconstruction. Traditional classification of the superfamily at suprageneric level is disputable in the light of recent advances. Transformational homology of tergal chaetotaxy was traced and revised based on 38 species of first instar and partial early instars. Morphological phylogenetic reconstructions were reconstructed mainly relying on first instar characters using both parsimony and likelihood-based algorithms. Outgroup selection and several rogue taxa impacted on resolution and support of otherwise well-supported clades. Integrating published molecular phylogeny, a revised classification of three families and nine subfamilies was presented: Orchesellidae, Entomobryidae and Paronellidae. Orchesellidae includes all basal taxa having a short fourth abdominal segment. Cyphoderidae and Microfalculidae taxa were sunk into Paronellidae. New Paronellidae was divided into two subfamilies: Paronellinae (Paronellini + Cyphoderini + Bromacanthini) and Salininae (Cremastocephalini + Callyntrurini). Microfalculidae (Microfalcula) was closer to Salina and Akabosia and thus transferred into Cremastocephalini. This study erected a new classification framework for Entomobryoidea based upon comprehensive phylogenies. Chaetotaxic homologization across a wide range provided a standardized, comparable, powerful tool for taxonomy.

MAFFT-DASH: Integrated protein sequence and structural alignment

Article

Full-text available

Jul 2019
NUCLEIC ACIDS RES

Here, we describe a web server that integrates structural alignments with the MAFFT multiple sequence alignment (MSA) tool. For this purpose, we have prepared a web-based Database of Aligned Structural Homologs (DASH), which provides structural alignments at the domain and chain levels for all proteins in the Protein Data Bank (PDB), and can be queried interactively or by a simple REST-like API. MAFFT-DASH integration can be invoked with a single flag on either the web (https://mafft.cbrc.jp/alignment/server/) or command-line versions of MAFFT. In our benchmarks using 878 cases from the BAliBase, HomFam, OXFam, Mattbench and SISYPHUS datasets, MAFFT-DASH showed 10-20% improvement over standard MAFFT for MSA problems with weak similarity, in terms of Sum-of-Pairs (SP), a measure of how well a program succeeds at aligning input sequences in comparison to a reference alignment. When MAFFT alignments were supplemented with homologous sequences, further improvement was observed. Potential applications of DASH beyond MSA enrichment include functional annotation through detection of remote homology and assembly of template libraries for homology modeling.

DNA-based approaches uncover cryptic diversity in the European Lepidocyrtus lanuginosus species group (Collembola: Entomobryidae)

Preprint

Full-text available

Mar 2019

DNA sequence data and phylogenies are useful tools for species delimitation, especially in taxa comprising cryptic species. The Lepidocyrtus lanuginosus species group (Collembola: Entomobryidae) comprises three morphospecies and distinct cryptic species. We applied three DNA-based methods to delimit species boundaries in the L. lanuginosus species group across central and southern Europe. Using cytochrome c oxidase subunit I and II, we identified gaps of genetic distances that indicate species boundaries and found 10 and 9 distinct genetic lineages in L. cyaneus and L. lanuginosus, respectively. The nuclear gene elongation factor 1-a delimited 89% of the lineages but 28S rDNA (D1-2 domain) was too conserved for this purpose. The phylogenetic trees showed that L. cyaneus and L. lanuginosus are polyphyletic, suggesting that body colour is insufficient for delimiting species in the L. lanuginosus species group. Our study challenges the current morphology-based species delimitation in the L. lanuginosus species group and suggests that molecular approaches are needed for fast and accurate determination of Collembola species in both taxonomic and ecological studies. Overall, the results suggest that wide geographic sampling combined with molecular phylogenetic approaches is needed to delimit species and to understand the full range of cryptic diversity in Collembola. Additional keywords: barcoding, colour, cytochrome c oxidase subunit I and II, elongation factor 1-a, ribosomal subunit 28S rDNA D1-2 domain, springtail.

New European Lepidocyrtus Bourlet, 1839 (Collembola, Entomobryidae) with the first description of feeding-related dancing behaviour in Collembola

Article

Full-text available

Jan 2019
ZOOTAXA

The study of a springtail population from Krka National Park (Croatia) has allowed us to describe the new species Lepi-docyrtus chorus sp. nov. and its dance-like behaviour. This study represents the first record of peculiar dancing behaviour related to search for food and feeding among Collembola. The new species is molecularly and morphologically assigned to the European Lepidocyrtus lignorum-group. Sequences of COXII and EF-1α genes, chaetotaxy and behaviour clearly help to differentiate the new species from the other species in the group.

New morphological and molecular data reveal an important underestimation of species diversity and indicate evolutionary patterns in European Lepidocyrtus (Collembola : Entomobryidae)

Article

Jun 2021

The correct identification of morphological species is a key task for species richness estimation of any ecosystem. Although body colour is a widely used character identifying European Lepidocyrtus species, recent investigations using molecular data have revealed that species delineation using body colour can result in an underestimation of real species diversity because of the presence of cryptic species. Lepidocyrtus violaceus is a European species characterised by its dark violet body colour. Its wide distribution leads us to suspect that several cryptic species can be present within this morphospecies. Since traditional morphological characters have appeared insufficient for real diversity identification in Lepidocyrtus, new morphological characters were needed in order to describe the cryptic diversity detected by molecular data in this genus. Pseudopores are integumentary structures present in all Lepidocyrtus species, but the distribution of these structures has not been properly described in the genus, as well as in Entomobryioidea overall. In the present work we aimed to analyse whether L. violaceus is a monophyletic entity in Europe. Moreover, we aimed to determine if the position and number of pseudopores on the different parts of the body and appendages is a phylogenetically useful character in the identification of the species or superspecific entities. Fourteen populations of L. violaceus from five European countries, and another 25 Lepidocyrtus species from nine European countries have been studied. In total, 208 specimens have been analysed morphologically and half of them were studied molecularly using sequences of the genes COXII and EF-1α. Molecular data revealed that the widely distributed Lepidocyrtus violaceus morphospecies is a polyphyletic entity in Europe. Between 6 and 12 diferent cryptic species have been detected within this European morphospecies, and only the presence of pseudopores on the basal plate of the fourth abdominal segment has been found to be a promising diagnostic character between them. A common basal pattern of pseudopore distribution has been recognised in the European members of the genus, and also a diferential pattern within each European species group. As a general trend, an increase in the number of pseudopores has been detected from the most basal to the most derived species groups in the phylogeny of the genus in Europe.

ASAP: Assemble Species by Automatic Partitioning

Article

Oct 2020

We describe ASAP (Assemble Species by Automatic Partitioning), a new method to build species partitions from single locus sequence alignments (i.e. barcode datasets). ASAP is efficient enough to split datasets as large 104 sequences into putative species in several minutes. Although grounded in evolutionary theory, ASAP is the implementation of a hierarchical clustering algorithm that only uses pairwise genetic distances, avoiding the computational burden of phylogenetic reconstruction. Importantly, ASAP proposes species partitions ranked by a new scoring system that uses no biological prior insight of intra‐specific diversity. ASAP is a stand‐alone program that can be used either through a graphical web‐interface or that can be downloaded and compiled for local usage. We have assessed its power along with three others programs (ABGD, PTP and GMYC) on 10 real COI barcode datasets representing various degrees of challenge (from small and easy cases to large and complicated datasets). We also used Monte‐Carlo simulations of a multi‐species coalescent framework to assess the strengths and weaknesses of ASAP and the other programs. Through these analyses, we demonstrate that ASAP has the potential to become a major tool for taxonomists as it proposes rapidly in a full graphical exploratory interface relevant species hypothesis as a first step of the integrative taxonomy process.

Integrative taxonomy reveals three new species of European Lepidocyrtus lignorum-group (Collembola, Entomobryidae)

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