Leiolima iberica, a new harvestman genus and species from the Iberian Peninsula (Arachnida, Opiliones, Sclerosomatidae)

Abstract

Recent studies have shown that the speciose Holarctic genera of Leiobunum C.L. Koch, 1839 and Nelima Roewer, 1910 are polyphyletic taxa, and therefore, the traditional diagnostic characters for these European genera of Leiobuninae Banks, 1893 (respectively, the presence or absence of tubercle rows on leg coxae) are unsuitable. We present the description of Leiolima iberica gen. et sp. nov., a new endemic harvestman from the north-western part of the Iberian Peninsula. The newly established genus shows intermediate characters between Leiobunum and Nelima. In addition, the new genus is characterized by shorter legs compared to Leiobunum and the presence of trichomes on all leg femora and pedipalpal patellae, a structure that is absent in all other western Palearctic genera of the subfamily Leiobuninae.

Full text

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European Journal of Taxonomy 609: 1–20 ISSN 2118-9773
https://doi.org/10.5852/ejt.2020.609 www.europeanjournaloftaxonomy.eu
2020 · Prieto C.E. & Wijnhoven H.
This work is licensed under a Creative Commons Attribution License (CC BY 4.0).
Research article
urn:lsid:zoobank.org:pub:F50120F1-767A-40E3-9B86-80DBA1BA041E
Leiolima iberica, a new harvestman genus and species
from the Iberian Peninsula
(Arachnida, Opiliones, Sclerosomatidae)
Carlos E. PRIETO 1, * & Hay WIJNHOVEN 2
1 Departamento de Zoología y Biología Celular Animal, Universidad del País Vasco
(UPV / EHU), PO Box 644, E-48080 Bilbao, Spain.
2 Groesbeeksedwarsweg 300, NL-6521 DW Nijmegen, Netherlands.
* Corresponding author: carlos.prieto@ehu.eus
2 Email: hayw@xs4all.nl
1 urn:lsid:zoobank.org:author:CAC20D21-78FF-4B8A-8023-F06A06EA031F
2 urn:lsid:zoobank.org:author:484CDCCE-5F82-4D84-B9BB-BE7AE7145AD7
Abstract. Recent studies have shown that the speciose Holarctic genera of Leiobunum C.L. Koch, 1839
and Nelima Roewer, 1910 are polyphyletic taxa, and therefore, the traditional diagnostic characters for
these European genera of Leiobuninae Banks, 1893 (respectively, the presence or absence of tubercle
rows on leg coxae) are unsuitable. We present the description of Leiolima iberica gen. et sp. nov., a
new endemic harvestman from the north-western part of the Iberian Peninsula. The newly established
genus shows intermediate characters between Leiobunum and Nelima. In addition, the new genus is
characterized by shorter legs compared to Leiobunum and the presence of trichomes on all leg femora
and pedipalpal patellae, a structure that is absent in all other western Palearctic genera of the subfamily
Leiobuninae.
Keywords. Leiolima new genus, Leiolima iberica new species, Portugal, Spain, trichomes.
Prieto C.E. & Wijnhoven H. 2020. Leiolima iberica, a new harvestman genus and species from the Iberian Peninsula
(Arachnida, Opiliones, Sclerosomatidae). European Journal of Taxonomy 609: 1–20.
https://doi.org/10.5852/ejt.2020.609
Introduction
The subfamily Leiobuninae Banks, 1893 is represented by three genera in Europe, Leiobunum
C.L. Koch, 1839, Cosmobunus Simon, 1879 and Nelima Roewer, 1910 (Martens 1978; Rambla 1970).
Geographically close to Europe are Microliobunum Roewer, 1912, with its two species from Lebanon
and SE-Turkey, and the monotypic Goasheer Snegovaya et al., 2018 erected for Microliobunum
iranum Roewer, 1952 (Kurt 2018; Snegovaya et al. 2018). Cosmobunus is also a monotypic genus,
with Cosmobunus granarius Lucas, 1846 ranging from northern Algeria and Morocco to Catalonia
NE-Spain, although it has recently been proposed that a fossil Miocene specimen from Spain belongs
to the genus Cosmobunus (Palencia et al. 2019). Cosmobunus granarius is similar to Leiobunum, but

European Journal of Taxonomy 609: 1–20 (2020)
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easily diagnosed by its having a smooth pedipalpal claw, a rough sclerotized dorsal surface and a broad
yellowish longitudinal band (Rambla 1970).
Currently, the species of Nelima are separated from Leiobunum merely by a negative criterion: the leg
coxae lack any row of tubercles, whereas the species of Leiobunum are dened as having at least one
row (Martens 1978). Martens (1969, 1978) has already mentioned that this single discriminating feature
is not sufcient to reveal true phylogenetic relationships, since penial morphology is not even taken into
account. Martens & Schonhöfer (2016) provide a recent example demonstrating this problem by creating
a group of species around Leiobunum rupestre (Herbst, 1799) for four European species, including
L. apenninicum (Martens, 1969), originally described under Nelima. In fact, an extensive molecular
analysis of the New World complex of ‘Leiobunum/Nelima’ (Hedin et al. 2012) has clearly demonstrated
that the classical taxonomical approach is highly supercial and that the entire group (Sclerosomatidae)
needs comprehensive revisions. In that paper, Gyas titanus Simon, 1879 (Sclerosomatidae) belonged
to the clade of Phalangiidae Simon, 1879, thus suggesting a new placement for the subfamily Gyinae
Šilhavý, 1946, while the remaining Sclerosomatidae were recovered as a monophyletic group. Within
this clade, the seven included European taxa (three Leiobunum spp., three Nelima spp. and Astrobunus
grallator Simon, 1879, but no species of Cosmobunus) were recovered in basal positions, with the
exception of Nelima doriae (Canestrini, 1871), which was grouped with some New World Gagrellinae
Thorell, 1889. However, neither Leiobunum nor Nelima, two genera with Holarctic ranges, have been
recovered as monophyletic groups. Interestingly, the two European taxa included in the combined
analysis (N. doriae and Astrobunus) were proven to form a clade, although being placed in different
subfamilies, which is a perfect demonstration of the title by Hedin et al. (2012), that is, in this group
"geography is better than taxonomy" in the prediction of phylogeny.
The Iberian Peninsula is known for its high level of endemism (Prieto 2003). So far, eight species of
Leiobunum and ve species of Nelima have been reported from the region (Martens 1969; Prieto &
Fernández 2007; Prieto 2008; Prieto & Wijnhoven 2017), of which six and two species, respectively, are
endemic. Several representatives of Leiobuninae from the Iberian Peninsula await descriptions (Prieto
2008).
In this contribution, we present a new species of Leiobuninae from Northern Spain/Portugal, whose
general appearance is reminiscent of that of Nelima. However, since it also has distinct coxal tubercle
rows, by denition, we ought to use the generic name Leiobunum for this taxon. To face this conictive
situation, we introduce the name Leiolima gen. nov., being aware of the fact that the European
Sclerosomatidae should be thoroughly revised. The species was rst mentioned and gured by Prieto
(2008: g. 10) as Leiobunum sp. nov.
Material and methods
Institutional abbreviations
AMNH = American Museum of Natural History, New York, USA
CHW = Collection Hay Wijnhoven, Nijmegen, the Netherlands
CJM = Collection Jochen Martens, Mainz, Germany
MNCN = Museo Nacional de Ciencias Naturales (CSIC), Madrid, Spain
MNHN = Muséum national d’histoire naturelle, Paris, France
Naturalis = Naturalis Biodiversity Center, Leiden, the Netherlands
SMF = Senckenberg Natural History Museum, Frankfurt am Main, Germany
ZUPV = Departamento de Zoología y Biología Celular Animal, University of the Basque Country
(UPV / EHU), Leioa, Bizkaia, Spain

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
3
Abbreviations for morphological terms
A = anterior row of coxal tubercles (lowercase for weakly developed ones) in the formula for leg 1–4
coxae; measurements in mm.
BLI = ratio between length of femur I and prosomal width (measured between lateral protuberances
placed between the 2nd and 3rd coxae) (Staręga 1972: 463)
Ch = sensillum chaeticum
F = female
fe = femur
L = leg
le = body length
M = male
mt = metatarsus
P = posterior row of coxal tubercles (lowercase for weakly developed ones) in the formula for leg 1–4
coxae; measurements in mm
pa = patella
ta = tarsus
ti = tibia
Tr = trichome
tr = trochanter
wi = prosomal width
Results
Taxonomy
Class Arachnida Lamarck, 1801
Order Opiliones Sundevall, 1833
Family Sclerosomatidae Simon, 1879
Subfamily Leiobuninae Banks, 1893
Leiolima gen. nov.
urn:lsid:zoobank.org:act:D808528C-9131-4A73-B0B5-48DA9132A183
Type species
Leiolima iberica gen. et sp. nov.
Diagnosis
A genus of the subfamily Leiobuninae having the following combination of characters: prosomal shield
fully granulated; supra-cheliceral lamellae with two cylindrical, apically granulated protuberances;
ozopores being rounded and minute; ocularium is deep black, granulated, canaliculate and does not tilt
backwards; cephalothorax with a median raised area in front of ocularium; opisthosoma densely covered
with coarse granules; saddle as paramedian pairs of diffuse patches; leg coxae margins with short tubercle
rows, except on rear margin of third leg coxae; trochanter concolorous with coxae; legs of medium
length, brownish, with silvery spots on dorsal sides; leg femora and patellae covered with trichomes;
tibia II with pseudoarticulations; pedipalpal patella, tibia and tarsus covered with trichomes; pedipalpal
male tarsus without ventral row of microdenticles, claw pectinate; slender penis, with broadened alate
portion; ventral membranes reaching base of glans, further distal from the dorsal membranes.
This genus can easily be differentiated from the three currently accepted European genera of Leiobun-
inae. The monotypic Cosmobunus presents a smooth tarsal claw, large body size and long legs (males

European Journal of Taxonomy 609: 1–20 (2020)
4
with BLI>3), opisthosomal sternites with denticle rows, and a conspicuous, lighter and broad medial
stripe (contrasting with a dark lateral pattern). The European-Maghrebian species of the Holarctic spe-
ciose genus Leiobunum, which include the type species L. rotundum (Latreille, 1798), have very long
legs (males with BLI>3) and an ocularium of variable color; the absence of trichomes on leg femora and
patellae is the major character that distinguishes it from Leiolima. Like Leiobunum, Nelima is another
Holarctic speciose genus with several European-Maghrebian species, including the type species Nelima
silvatica (Simon, 1879); the absence of tubercle rows along the margins of the leg coxae is the main
diagnostic feature.
Etymology
The generic epithet is a combination of the generic names Leiobunum and Nelima, and the gender is
feminine.
Key to the European genera of Leiobuninae
1. Pedipalpal claws smooth. Opisthosomal sternites with transversal rows of granules ........................
.................................................................................................................. Cosmobunus Simon, 1879
– Pedipalpal claws pectinate (Fig. 5B). Opisthosomal sternites smooth ............................................. 2
2. All leg coxae margins without tubercle rows .................................................. Nelima Roewer, 1910
– Some, at least one, leg coxae margins with tubercle rows ............................................................... 3
3. Leg femora covered with trichomes (Fig. 3A, D). Legs of medium length (males with BLI<2) ......
............................................................................................................................... Leiolima gen. nov.
– Leg femora with ‘sensilla chaetica’ only (Fig. 3C, F). Long legs (males with BLI>3) ......................
...............................................................................................................Leiobunum C.L. Koch, 1839
Leiolima iberica gen. et sp. nov.
urn:lsid:zoobank.org:act:AFA32744-CAE4-4DF0-88FC-5122775787E1
Figs 1–10
Leiobunum sp. – Prieto 2008: g. 10.
Diagnosis
Same as for genus.
Etymology
The specic epithet refers to its supposed endemicity for the Iberian Peninsula.
Type material
Holotype
SPAIN • ♂; León Prov., San Feliz de las Lavanderas; 42.6869° N, 5.9765° W; 1182 m a.s.l.; 12 Oct.
2006; C. Prieto and K. Altonaga leg.; pitfall traps; MNCN 20.02/17372.
Paratypes
SPAIN – León Prov. • 1 ♂, 1 ♀; same data as for holotype; MNCN 20.02/17373 to 17374 • 2 ♂♂; same
data as for holotype; ZUPV 3809. • 1 ♂; Ponferrada, Campus Universidad; 42.5479° N, 6.5829° W; 560 m
a.s.l.; 13 Oct. 2006; C. Prieto leg.; grasses; ZUPV 3821 • 1 ♂; Robledo de la Valduerna; 42.3270° N,
6.0696° W; 820 m a.s.l.; 8 Nov. 1988; K. Altonaga and R. Gorrotxategi leg.; riverside; ZUPV 1373 •
1 ♂; Susañe del Sil; 42.8308° N, 6.5047° W; 894 m a.s.l.; 30 Oct. 2009; C. Prieto and K. Altonaga leg.;

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
5
ruderal, grass; ZUPV 4261 at CHW. ‒ Ávila Prov. • 1 ♀; Piedrahita, Eta.Virgen de la Vega; 40.4876° N,
5.3303° W; 1000 m a.s.l.; 22 Sep. 1987; K. Altonaga and A.I. Puente leg.; ZUPV 1446. ‒ Salamanca
Prov. • 1 ♂; Peña de Francia; 40.5127° N, 6.1584° W; 1350 m a.s.l.; 20 Sep. 2002; Sergio Montagud
leg.; pasture, pinewood; ZUPV 3279. ‒ Zamora Prov. • 2 ♂♂; Cubelo; 42.0982° N, 6.6509° W; 1000 m
a.s.l.; 9 Nov. 1988; K. Altonaga and R. Gorrotxategi leg.; orchards, oaks; ZUPV 1471 • 1 ♂; Lubián;
42.0332° N, 6.9239° W; 1030 m a.s.l.; 9 Nov. 1988; K. Altonaga and R. Gorrotxategi leg.; pasture,
chestnuts; ZUPV 1076 • 1 ♂; Tabara; 41.8509° N, 5.9880° W; 790 m a.s.l.; 9 Nov. 1988; K. Altonaga and
R. Gorrotxategi leg.; pasture, poplars; ZUPV 1378.
PORTUGAL – Bragança Distr. • 2 ♂♂; Freixo de Espada, Mazouco; 41.1671° N, 6.7989° W; 730 m
a.s.l.; 7 Feb. 2001; Pedro Cardoso leg.; Cistus, Genista; ZUPV 3324 • 1 ♀; Freixo de Espada, Palao;
41.1261° N, 6.8217° W; 630 m a.s.l.; 10 Oct. 2001; Pedro Cardoso leg.; ZUPV 5001 • 2 ♀♀; Freixo de
Espada, Picotino; 41.2051° N, 6.7619° W; 740 m a.s.l.; 7 Feb. 2001; Pedro Cardoso leg.; ZUPV 5000 •
2 ♂♂, 1 ♀; same collection data as for preceding; 21 Feb. 2001; Pedro Cardoso leg.; pinewood; ZUPV
3330 • 3 ♂♂; same collection data as for preceding; CHW • 1 ♂; Malhadas; 41.5429° N, 6.3205° W;
740 m a.s.l.; 10 Nov. 1988; K. Altonaga and R. Gorrotxategi leg.; pasture, ash; ZUPV 1648 • 1 ♀;
Mogadouro, Algozinho; 41.3046° N, 6.5590° W; 620 m a.s.l.; 21 Feb. 2001; Pedro Cardoso leg.;
ZUPV 5002 • 1 ♂, 2 ♀♀; Mogadouro, Bruçó; 41.2522° N, 6.7268° W; 760 m a.s.l.; 7 Feb. 2001; Pedro
Cardoso leg.; Douglas r; ZUPV 3333 • 1 ♂, 2 ♀♀; same collection data as for preceding; CJM • 2 ♂♂;
Porrais; 41.400° N, 6.805° W; 360 m a.s.l.; 10 Nov. 1988; K. Altonaga and R. Gorrotxategi leg.; ruderal;
Fig. 1. Leiolima iberica gen. et sp. nov. Dorsal view of the male holotype (MNCN 20.02/17372) (left,
body length 3.15 mm) and a female paratype (MNCN 20.02/17374) (right, body length 5.58 mm). Scale
bars: 1 mm.

European Journal of Taxonomy 609: 1–20 (2020)
6
ZUPV 1419 • 1 ♂; Quintanilha; 41.720° N, 6.589° W; 600 m a.s.l.; 11 Nov. 1988; K. Altonaga and R.
Gorrotxategi leg.; ruderal; ZUPV 1532 • 1 ♂; Vimioso; 41.593° N, 6.558° W; 500 m a.s.l.; 11 Nov.
1988; K. Altonaga and R. Gorrotxategi leg.; ruderal; ZUPV 1497. ‒ Guarda Distr. • 1 ♂; Seia, Cabeça;
40.324° N, 7.746° W; 550 m a.s.l.; Dec. 1998; Grosso-Silva leg.; ZUPV 4104 • 2 ♂♂, 2 ♀♀; same
collection data as for preceding; Jan. 1999; Grosso-Silva leg.; ZUPV 4108.
Remarks
Paratype specimens (male and female) from Freixo de Espada (Bragança, Portugal; samples ZUPV
3324, 3330, 5000 and 5001) have been deposited at the SMF: ♂ (ZUPV 3324), ♀ (ZUPV 5000);
Fig. 2. Leiolima iberica gen. et sp. nov. Lateral view of the male holotype (MNCN 20.02/17372) (left is
front side), showing coxae I to IV with coxal denticle rows.
Table 1. Leiolima iberica gen. et sp. nov. Leg lengths of male holotype (in bold type) and average leg
lengths of males (n = 4) and females (in parentheses, n = 4).
tr fe pa ti mt ta Total
I0.372.98 0.80 2.19 2.40 5.67 14.41
0.4 (0.4) 3.3 (2.9) 1.0 (0.9) 2.6 (2.3) 2.8 (2.3) 6.8 (5.1) 16.5 (13.5)
II 0.36 5.00 1.03 4.37 4.42 11.92 27.10
0.4 (0.4) 5.7 (4.9) 1.2 (1.2) 5.0 (4.5) 5.0 (4.3) 13.5(11.2) 30.4 (26.1)
III 0.37 3.17 0.88 2.28 2.78 5.91 15.39
0.4 (0.4) 3.6 (3.1) 1.0 (0.9) 2.6 (2.3) 3.2 (2.7) 6.6 (5.2) 17.0 (14.2)
IV 0.41 5.27 1.05 3.44 4.68 7.77 22.62
0.4 (0.4) 5.7 (4.9) 1.2 (1.1) 3.9 (3.4) 5.2 (4.5) 8.7 (6.8) 24.7 (20.7)

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
7
Naturalis: ♂ (ZUPV 3324), ♀ (ZUPV 5000); MNHN: ♂, ♀ (ZUPV 3330); AMNH: ♂ (ZUPV 3330), ♀
(ZUPV 5001).
Description
Male (holotype, Fig. 1)
Body length 3.15, prosoma width 1.85, and BLI index 1.61. Leg lengths (Table 1).
Prosoma. Anterior margin slightly concave on both sides and midsection protruding (Fig. 1). Cheliceral
apodemes appear as two narrow stripes. Supracheliceral lamellae with two cylindrical protuberances,
each having three or four apical granules (Fig. 4C). Ozopores close to anterolateral margin, rounded and
minute (0.04–0.05 diameter).
Fig. 3. Male femur I of three European species of Leiobuninae. A, D. Leiolima iberica gen. et sp. nov.
(ZUPV 3330 at CHW). B, E. Nelima doriae (Canestrini, 1871) (from the Netherlands). C, F. Leiobunum
rotundum (Latreille, 1798). A–C. Dorsal view. D–F. Lateral view (dorsal side on the right). Ch =
sensillum chaeticum; Tr = trichome (trichomes in D drawn smaller). Scale bars: A–C = 0.05 mm; D–F
= 0.5 mm.

European Journal of Taxonomy 609: 1–20 (2020)
8
Dorsum. Dorsal integument with a microtuberculate-microgranulate morphology, consist of distinct
obtuse to acute granules, approximately separated by their width. Granules towards sides more prominent.
Coloration of Dorsum. Dorsum dark reddish-brown with black granules and numerous scattered
yellowish to silvery spots, especially prominent near sides of prosoma (Fig. 1). Saddle as faint pattern of
two paramedian longitudinal bands of diffuse and blackish segmental patches. Muscle insertion plaques
dark brown. Area in front of ocularium dark brown. Prosoma bordered with black.
oCularium. Shallow, basally constricted canaliculate, covered with large granules and dorsally armed
with two rows of pointed black denticles, with a few sensilla chaetica (Fig. 4D). Eyes (diameter 0.15) on
a black ocularium (0.32 long, 0.35 wide), base reddish-brown. Ocularium located about its length from
anterior margin of prosoma.
Venter anD Coxae. Coxae smooth, pale yellowish, armed with numerous black sensilla chaetica, distal
pro- and posteriolateral sides with short rows of tubercles (Fig. 2). Posteriolateral side of coxa III
without or rarely with sparse tubercles (Table 2). The resulting coxal tubercle formula can be expressed
as Ap Ap A- aP [A, P, anterior, posterior row (lowercase for weakly developed ones) from 1st to 4th leg
Table 2. Leiolima iberica gen. et sp. nov., number of tubercles on lateral margins of leg coxae (n = 11
males, 7 females). Averages on the right are from right coxae, and averages on the left come from the
combined results on both sides. Catalogue numbers refer to the ZUPV collection.
Leg I II III IV I (r) II (r) III (r) IV (r)
MALES A P A P A P A P A P A P A P A P
3279 12 8 8 2 8 1 3 8 12 5 12 6 7 3 6 7
3324 8 4 9 4 9 0 0 7 10 4 7 4 7 0 0 8
3330 13 0 10 3 9 0 3 7 12 0 12 1 9 0 3 7
3330 12 0 10 7 8 0 3 6 11 2 9 6 9 0 8 6
3333 12 2 6 6 8 0 6 5 11 2 8 2 8 0 2 7
3809 9 2 9 4 9 0 4 7 12 2 10 4 8 0 2 6
3821 11 4 12 7 8 0 3 7 9 2 10 6 11 0 4 6
4108 14 7 10 9 12 0 2 9 16 4 11 –11 1 3 10
4108 13 8 10 9 8 0 3 10 14 9 11 9 10 0 3 9
4108 12 10 12 9 10 0 4 8 12 5 11 7 12 0 4 7
4261 7 0 7 0 7 0 2 4 9 0 9 0 7 0 1 7
average 11.2 4.1 9.4 5.5 8.7 0.1 3 7.1 11.6 3.2 10 4.5 9 0.4 3.3 7.3
FEMALES
1446 14 8 11 6 9 0 7 11 12 8 10 8 8 0 6 11
3330 10 1 8 4 8 0 3 7 8 0 8 2 7 0 3 8
3333 12 0 10 4 6 0 3 7 11 011 2 10 0 2 7
3333 11 2 10 2 8 0 1 7 11 011 4 8 0 1 6
3809 9 2 9 4 9 0 4 7 12 2 10 4 8 0 2 6
4108 13 2 10 7 7 0 0 8 13 2 11 7 9 0 2 11
4108 12 7 10 8 9 0 2 11 13 8 11 10 8 0 0 11
average 11.6 3.1 9.7 5 8 0 2.9 8.3 11.4 2.9 10.3 5.3 8.3 0 2.3 8.6

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
9
coxae]. Largest individual coxal tubercles 25–30 μm in height, separated from each other by 20–30 μm,
with upper ange wider than basal width, bevelled towards their coxa and trilobed, being higher than
central lobe. Ventral side pale yellowish. Epistome conical and 0.3 long. Genital operculum with sensilla
chaetica, laterally with few obtuse denticles (Fig. 4A); internal side of anterior end of genital operculum
without sclerotized lobes or any other cuticular structure.
CheliCerae. Smooth, pale yellowish, dorsal area of rst segment darkened and provided with ‘sensilla
chaetica’ (Fig. 4B).
PeDiPalPs. Pale yellowish, distal areas of femur darkened, as well as patella and tibia. Femur covered
with large black sensilla chaetica, medially with black-tipped blunt denticles (Fig. 5); a single spine
occurs dorsodistally. Pedipalpal tarsus almost straight, only very slightly curved inward, claw pectinate
(Fig. 5B), without ventral rows of denticles. Pedipalpal patella, tibia and tarsus covered with trichomes
(non-sensorial, not socketed “setae”). Dorsal regions of patella and tibia with elds of robust, transparent
trichomes (Fig. 5D).
legs. Medium long and slender (Table 1; BLI = 1.62 in holotype), orange-brown, mottled with small
silvery spots, especially on dorsal sides of patella and tibia. Trochanters with bright patches on anterior
Fig. 4. Leiolima iberica gen. et sp. nov. (male) (holotype, MNCN 20.02/17372). A. Genital operculum.
B. Right chelicera (median view). C. Supracheliceral lamellae (dorsal view). D. Ocularium (lateral
view; front side on the right). Scale bars: A–B = 0.5 mm; C–D = 0.1 mm.

European Journal of Taxonomy 609: 1–20 (2020)
10
and posterior sides. Femur I conspicuously shorter (59.5% in holotype) and more robust than femur II.
Femora with irregular rows of small, black-tipped spines, each accompanied by a ‘sensillum chaeticum’.
Tibia II has 5–6 silvery bordered pseudoarticulations. Patellae with three pointed granules on dorsodistal
margin. All femora, except ventrally, densely covered with trichomes (Fig. 3A, D), which also occur on
patella, tibiae, metatarsi and tarsi of all legs.
Penis. Truncus slightly and regularly narrowing towards top (Fig. 6). Alate portion well developed and
elongated, consisting of two dorsal and two ventral membranes which are fused only in proximal one-
quarter section, thus forming two pockets that open to exterior distally as well as laterally. Ventral
membranes attached at truncus/glans junction and have a characteristically curved distal margin.
Dorsal membranes attached more proximally, extending on to ventral side, forming two wings that
partly enclose ventral membranes. Distal margins of ventral membranes have an irregular ‘worn-down’
appearance (Fig. 6D). Glans and stylus with characteristic shape (Fig. 6F–G) and four sensilla chaetica
located near tip. Measurements: penis length 1.83, glans + stylus length 0.29, truncus length from base
to pockets 0.91, penial muscle length 0.77, basal truncus width 0.23, width of alae 0.32.
Female (Fig. 1)
Body length 5.58, prosoma width 2.17, opisthosoma width 3.29, BLI index 1.24.
Fig. 5. Right male pedipalp of Leiolima iberica gen. et sp. nov. (holotype, MNCN 20.02/17372). A.
Median view. B. Lateral view with detail of claw. C. Detail of ventral side of femur showing sensilla
chaetica and imbricate microsculpture. D. Detail of dorsal side of patella showing robust trichomes (tr)
and sensilla chaetica (ch). Scale bars: A–B = 0.5 mm; C–D = 0.05 mm.

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
11
legs. Long, with legs I, III and IV more robust than in male, leg II slender. Femur of leg I more than
twice as wide as that of leg II (Table 1).
seminal reCePtaCles (Fig. 7). In fourth to fth ovipositor segments.
Fig. 6. Penis of Leiolima iberica gen. et sp. nov. A. Ventral view. B. Detail of distal part (ventral view).
C. Lateral view. D. Dorsal view. E. Attachment sites of dorsal and ventral membranes (dorsal view).
F. Stylus (lateral view, ventral side to left). G. Stylus of two males in ventral (left) and dorsal (right)
view. A, G: paratypes, ZUPV 3330 at CHW; B–F: paratype, ZUPV 4261 at CHW. Scale bars: A, C–D =
0.5 mm; B = 0.1 mm; F–G = 0.05 mm.

European Journal of Taxonomy 609: 1–20 (2020)
12
Sexual dimorphism
Leiolima iberica gen. et sp. nov. shows minor sexual dimorphism. Females present smaller silvery
patches on the prosoma and the length of the rst thoracic tergite is about twice that in males. Legs are
1/6 shorter than in males, but female bodies are almost twice as long as male ones, with the opisthosoma
greatly enlarged in egg-developing females.
Distribution
Leiolima iberica gen. et sp. nov. is an Iberian endemic restricted to the north-western corner of the
Iberian Peninsula (Fig. 8). The known distribution extends along the mountainous areas ranging from
the north-western part of Castilla y León (Spanish provinces of León and Zamora) and the eastern part
of the old Tras-os-Montes e Alto Douro region (Portuguese district of Bragança) to the mountain range
that separates the Duero/Douro basin from the Tajo/Tejo basin (Serra da Estrela in Portugal, and Sierra
de Francia and Sierra de Gredos in Spain). Galicia, Asturias or Extremadura have no records, although
it may exist in the border areas.
Fig. 7. Female Leiolima iberica gen. et sp. nov.; seminal receptacles, distance between left and right
receptacles to scale. Paratypes: top: ZUPV 3333 at CHW, bottom ZUPV 4108 at CHW. Scale bar: 50 μm.

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
13
It is possible to recognize a northern concentration and a southern block, separated by less than 100 km.
Although both are separated by the deep canyons of the Duero/Douro River bordering Spain and
Portugal, here we assume a continuous distribution of the species. In the southern area, the only known
locality of the Portuguese Serra da Estrela is 150 km west of the Sierra de Gata region of Spain. The
similar orogeny and geography, as well as the absence of obvious physical barriers, make their presence
in the comprised area probable. The eastern border of its known distribution area in Castilla y León is
marked by the Órbigo River (a tributary of the Esla River), which separates the Montes de León from
the vast plains of the Duero basin, which are mainly cultivated with cereals.
Habitat
Leiolima iberica gen. et sp. nov. is a species that lives in dry open biotopes, mainly pastures, grasslands
and ruderal biotopes. It seems to ll the characteristic Nelima niches (ground and herb layer, sheltering
under stones, wood and vegetation). The altitudinal range is 360–1350 m a.s.l. (average 782 m), with
two somewhat divergent localities: the Porrais site (360 m a.s.l.) is near the bottom of the Sabor River
(a tributary of the Douro River), whereas the Peña de Francia site (1350 m a.s.l.) is near the mountain
top of the Sierra de Francia. The average altitude of the 129 sampled localities from the concerned
provinces or districts represented in the ZUPV collection is 975 m, which on average is distinctly higher,
suggesting that L. iberica gen. et sp. nov. is a ‘lowland’ species.
Fig. 8. Geographical distribution of Leiolima iberica gen. et sp. nov. on the Iberian Peninsula. Some
names of autonomic or regional entities (in capitals, without borders), mountain ranges and larger rivers
(in italics) are given.

European Journal of Taxonomy 609: 1–20 (2020)
14
Phenology
Adults of Leiolima iberica gen. et sp. nov. have been obtained from September to February, mainly
during malacological sampling trips. Because sampling outside of the appropriate season for land snails
was infrequent, phenological information is biased. The collected specimens resulted in a sex ratio of
2.07, which is about two males per female. The bimonthly sex ratio gures (7 males / 3 females in Sep.–
Oct.; 11 / 0 in Nov.–Dec.; 11 / 11 in Jan.–Feb.) does not allow further conclusions about its life cycle,
which is also obscured by the total absence of juvenile specimens in the samples.
Fig. 9. Body size and leg length measurements (in mm apart from the MBLI ratio) of all European/
Mahgrebian species of Leiobunum C.L. Koch, 1839 (grey bars, diamonds on averages) and Leiolima
iberica gen. et sp. nov. (red rectangles). Only the measured parameters for L. iberica gen. et sp. nov.
are shown. Source of data: Caporiacco (1929), Karaman (1996), Komposch (1998), Martens (1978),
Prieto & Fernández (2007), Prieto & Wijnhoven (2017) and Šilhavý (1965).

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
15
Discussion
Leiolima iberica gen. et sp. nov. represents a new Iberian species with some peculiar features. Evidently,
the generic name itself is an expression of our reservations regarding the placement of this species
within Leiobuninae. However, as mentioned earlier, it reects the need for a general revision of the
European Sclerosomatidae.
Therefore, due to the presence of coxal tubercle rows, the new species should be included in Leiobunum,
as suggested by Prieto (2008), where this taxon is mentioned for the rst time as an undescribed species.
As mentioned earlier, Roewer (1923) based the separation between Leiobunum and Nelima on the single
criterion of the presence (Leiobunum) or absence (Nelima) of rows of tubercles on both, anterior and
posterior, margins of the leg coxae. The genus Cosmobunus was not considered a related genus at that
time because Roewer (1923, 1957) included it in the North American subfamily Leptobuninae, which
was dismantled by Cokendolpher (1985), transferring Cosmobunus to the subfamily Leiobuninae.
Martens (1978) retained this system due to the absence of alternatives.
The fuzzy taxonomy of the family Sclerosomatidae has been revealed by a molecular analysis of
Hedin et al. (2012). The placement of the sclerosomatid Gyas titanus (Gyinae) in the Phalangiidae
ingroup, highly supported by combined Bayesian analysis and recently conrmed by Fernández et al.
(2017) through transcriptomic analysis, is an outstanding result that agrees with the absence of coxal
tubercle rows, a key feature of the family Phalangiidae (Tsurusaki 2007). Besides Gyas titanus, Hedin’s
phylograms show a limited representation of seven European sclerosomatids; that is, three Leiobunum
and three Nelima species plus Astrobunus grallator (although Cosmobunus is missing). Interestingly,
in the 28S Bayesian phylogram, these seven taxa were recovered in basal positions within a clade that
has several New World gagrellines but not a single New World Leiobuninae. In the combined Bayesian
analysis, all the extra-European leiobunines and Japanese gagrellines constitute a highly supported
monophyletic clade, whereas the two European taxa (Nelima doriae and Astrobunus) constitute the sister
clade of the New World gagrellines (Trachyrhinus Weed, 1892 and Krusa Goodnight & Goodnight,
Fig. 10. Orices of repugnatorial glands of the type species of European genera of Leiobuninae in dorsal
(upper) and lateral (lower) views. A. Leiolima iberica gen. et sp. nov. (male, ZUPV 4104). B. Nelima
silvatica (Simon, 1879) (male, ZUPV 1316: Castarnés, Huesca, Spain). C. Leiobunum rotundum
(Latreille, 1798) (male, ZUPV 5065: Oña, Burgos, Spain). D. Cosmobunus granarius Lucas, 1846
(male, ZUPV 1932: Hornos, Jaén, Spain). Coxae I were removed to improve vision and images have
been resized in proportion to their respective prosomal width (A = 111 %; B = 125 %; C = 82 %; D =
73 %). Arrowheads in upper views point to the orices. Scale bars: 0.1 mm.

European Journal of Taxonomy 609: 1–20 (2020)
16
1957). Therefore, if the European and American leiobunine taxa belong to two unrelated groups, each
species group of the polyphyletic ‘Leiobunum’ and ‘Nelima’ from each biogeographical region would
deserve a generic status. Given that the type species of Leiobunum and Nelima are European taxa, a
taxonomical revision would require a complete reorganization of the Japanese and North American
taxa. For example, the so-called “US Leiobunines” clade in Hedin et al. (2012: g. 2) includes 13
‘Leiobunum’ spp., and the two Eumesosoma spp. and ‘Hadrobunus sp.’ would have to be grouped under
Hadrobunus Banks, 1900 or, more likely, the resurrection of Forbesium Weed, 1890, the older name
for New World leiobunums, with priority over Eumesosoma Cokendolpher, 1980, if the respective type
species (not analysed) also belong to this clade. Alternatively, the genus Krusa, which was shown to be
polyphyletic, with two species (one being the type) related to Trachyrhinus and two others in a clade
together with a ‘Leiobunum’ and a ‘Nelima’ nov. sp. 1, would require a new generic name. In this
sense, Gaona-Escamilla et al. (2016), restraining the use of Krusa to the former group, consider that
the presence of a smooth ocularium and a lobed penis justies that ‘Krusa’ ava and ‘Krusa’ metallica
should be placed in another unnamed genus.
As Hedin et al. (2012) also pointed out, some characters are prone to homoplasy in Leiobuninae due to
heterochronic parallelism processes, e.g., the presence of coxal tubercle rows is a plesiomorphic feature
among sclerosomatids, but the absence of these tubercles, diagnostic for Nelima, is a characteristic trait
also for nymphal stages of Leiobunum (Hedin et al. 2012) and Cosmobunus (personal observation), a
feature frequently observed but never reported in the literature. The existence of many endemic species
of ‘Nelima’ in each region of its Holarctic range (North America, East Asia, Mediterranean Region)
suggests multiple independent cases of paedomorphosis on different continents, which is evidenced by
the existence (see phylograms in Hedin et al. 2012) of clusters of species of ‘Nelima’ nested in different
Leiobunum clades from the United States and Japan. However, accepting this hypothesis of repeated
pedomorphy does not imply that every species of Nelima arose by heterochronic processes, so plausibly,
different species of ‘Nelima’ of each regional cluster evolved through other speciation processes, i.e.,
vicariance, from the ancestral pedomorphic species.
All of this reveals a taxonomical maze due to the enormous diversity of the family Sclerosomatidae, more
than 1300 species, for which Hedin et al. (2012) propose a strategy of “sequence everything”, beginning
with a large sample (they suggest starting with about 500 species) to discover natural groups before a
post hoc search for morphological synapomorphies. This would require paralyzing taxonomic research
based on morphological reasons until fresh samples were obtained from the largely unknown East Asian
diversity, including the type species of all genera. Not against that approach, but complementary, may
be the creation of new genera based on new species if they do not t into current genera. They, like any
other taxa of any hierarchical level, are hypotheses that will be conrmed or refuted by future research,
and their creation demands that the hypothesis should be contrasted in future comprehensive analyses.
Shultz (2018) expresses a different point of view when describing a new North American species of
Leiobunum, by recognizing the difculty of its systematic placement because of the articial denition
and composition of genera of Roewerian, even considering its inclusion in other genera by the absence
of coxal tubercles or the reduced pectination of the tarsal claw. Although its nal plea for a broad-scale
morphological and molecular revision of sclerosomatid harvestmen is shared, the situation is different
in the western Palearctic region.
The situation of western Palearctic leiobunines seems less complex for three reasons. First, they
include the type species of Leiobunum, Nelima and Cosmobunus. The latter became monotypic when
Cokendolpher (1985) deprived it of the spurious American C. americanus Roewer, 1957; moreover,
the Near Eastern genus Microliobunum Roewer, 1912 and the Persian Goasheer Snegovaya et al.,
2018, rather distinctive by their very short legs (Roewer 1952; Snegovaya et al. 2018; Kurt 2018),
remain valid taxa. Secondly, nearly all European leiobunines are well-known taxa that have recently

Prieto C.E. & Wijnhoven H., Leiolima (Opiliones, Sclerosomatidae), a new genus from Iberia
17
been described or revised (Martens 1978; Karaman 1996; Komposch 1998; Prieto & Fernández
2007; Martens & Schönhofer 2016; Wijnhoven & Prieto 2017) and noteworthy faunistic novelties are
conspicuous, whereas the Central American fauna is poorly known and many of the already named
species are insufciently described (Gaona-Escamilla et al. 2016). Thirdly, the European/Mahgrebian
leiobunines show a greater penial uniformity; up to today it has not been possible to dene clear groups
of species in Leiobunum, whereas the American or Japanese faunas include such species groups (Suzuki
1976; Burns et al. 2012) which could rightly be recognized as separate genera.
Restraining the discussion to European taxa, we have based the placement of the new species in its
own genus on two character sets, leg length and trichome cover on proximal podomeres. The relative
length of legs with respect to the body, expressed as BLI index, is often used to diagnose genera.
Sclerosomatines are ground-dwelling harvestmen with short legs and a heavier dorsal sclerotization
with paired segmental protuberances on the scutum, a robust prosomal armature of spines or horns and
trochanter spurs, whereas leiobunines are scansorial or rupicolous, without paired scutal protuberances
or prosomal armature or trochanter spurs (Martens 1978; Hedin et al. 2012). These features place the
new species herein described among the leiobunines, whereas the presence of pectination on the palpal
claw precludes its belonging to Cosmobunus, and the presence of pro- and retrolateral rows of coxal
tubercles would force its inclusion in Leiobunum, as has been said before. However, the new species
show strikingly short legs, which is unlike any European Leiobunum (Fig. 9). While the body size of
L. iberica gen. et sp. nov. lies in the size span of Leiobunum, with relatively large females (larger than the
average size of females of Leiobunum), both the BLI index and leg measurements show that L. iberica
gen. et sp. nov. is clearly situated outside the morphological space of the European (species of the) genus
Leiobunum. Within this minor overall length, it is particularly prominent that the rst leg of L. iberica
gen. et sp. nov. is shorter than half the length of the rst leg in the most short-legged Leiobunum.
The presence of a trichome cover on all the proximal leg podomeres, in addition to the normal
ornamentation of denticles and ‘sensilla chaetica’, recorded as just setae or socketed setae in earlier
literature (Willemart et al. 2009), according to our knowledge, is a unique feature among the Western-
Palaearctic Leiobuninae (Wijnhoven 2013). In the three European genera of Leiobuninae known to
date, trichomes occur only on the tibiae, metatarsi and tarsi of legs, lacking on femora and patellae. We
checked this character for the following species: Nelima doriae (Fig. 3B, E), N. gothica Lohmander,
1945, N. sempronii Szalay, 1951, Leiobunum argentipalpe Prieto & Fernández, 2007, L. blackwallii
Meade, 1861, L. levantinum Prieto & Fernández, 2007, L. rotundum (Fig. 3C, F), L. sp. (sensu Wijnhoven
et al. 2007) and Cosmobunus granarius Lucas, 1846. However, in both juveniles and adults of Gyas
titanus, the presence of trichomes on the femur and patella was conrmed (unpublished data). Also, the
trichomes on the pedipalpal patella in addition to its normal occurrence on the tibia and tarsus, by our
knowledge, is a peculiar unique feature within Leiobuninae. Although the taxonomic signicance is
unknown, another discrepant character with respect to Leiobunum, or at least its type species, is the size
of the ozopores. They are circular and minute in L. iberica gen. et sp. nov. and N. silvatica (0.05 mm);
larger and ovate in Leiobunum rotundum (0.12 × 0.6 mm); and in Cosmobunus granarius, ozopores (0.1
× 0.11 mm) appear as indentations of a carapace ange above coxa I (personal observations).
Leiolima iberica gen. et sp. nov. really seems to deserve an assignment between Nelima and Leiobunum.
The dorsal microsculpture consisting of isolated granules is a character for many species of Nelima,
such as N. doriae and N. sempronii. In Leiolima iberica gen. et sp. nov. these granules are rather large,
whereas in species of Nelima these are generally less conspicuous. The European species of Leiobunum
usually have a rugose dorsal microstructure with pointed microdenticles. Also, the ocularium is Nelima-
like, with two rows of teeth, whereas the ocularium is smooth in most species of Leiobunum. The penial
morphology, on the other hand, is Leiobunum-like and the stylus tip also resembles that of Leiobunum.

European Journal of Taxonomy 609: 1–20 (2020)
18
However, in most species of Leiobunum, the ventral membranes of the pockets are proximally attached
to the dorsal membranes rather than distally, such as in Leiolima gen. nov.
Although many parts of the Iberian Peninsula have been poorly sampled, it is surprising that a relatively
large species, with such an extended distribution area (19 localities within an area of approximately
30000 km²) and living in a common biotope, has been overlooked for such a long time.
Acknowledgements
This work was partially funded by the Basque Government through the Research Group on Systematics,
Biogeography and Population Dynamics (GIC10/76; IT575/13) and by the Spanish Ministerio de
Ciencia e Innovación (Ref. CGL2008-01131 ⁄ BOS). HW expresses his sincere gratitude towards the
Uyttenboogaart-Eliasen Stichting for kindly awarding him with a grant (SUB.2012.05.04) to visit C. Prieto
at the Basque University in Bilbao as part of a collecting expedition to Morocco and Spain. The text of the
manuscript checked by Modern Manuscript Editing Services (http://www.mmanuscripteditserv.com/).
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Manuscript received: 25 April 2019
Manuscript accepted: 18 November 2019
Published on: 27 February 2020
Topic editor: Rudy Jocqé
Desk editor: Eva-Maria Levermann
Printed versions of all papers are also deposited in the libraries of the institutes that are members of
the EJT consortium: Muséum national d’histoire naturelle, Paris, France; Meise Botanic Garden,
Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural
Sciences, Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis
Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid,
Spain; Real Jardín Botánico de Madrid CSIC, Spain; Zoological Research Museum Alexander Koenig,
Bonn, Germany; National Museum, Prague, Czech Republic.