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First detailed records of Calathocratus from the Iberian peninsula, with description of two new species (Opiliones: Dyspnoi: Trogulidae)

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The genus Calathocratus is reliably recorded from the Iberian Peninsula, encompassing two new species described herein. Both species are small, have a similar overall morphology, like a dorsal median area devoid of papillae, but they are very different in penial morphology. Calathocratus zaragozai sp. nov. from the Valencian-Alicante mountains in Spain has a unique subapical stylus and a glans with large setae, while Calathocratus algarvensis sp. nov. from the Algarve region in Portugal, has a stout apical stylus and a glans with short, robust setae. Descriptions are complemented with the first record of teratological anophthalmia in Tro-gulidae and several observations of citizen science. A discussion on related species is included.
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Revista Ibérica de Aracnología,44 (30/06/2024): 127–139. ARTÍCULO
Grupo Ibérico de Aracnología (S.E.A.). ISSN: 1576 - 9518. http://www.sea-entomologia.org
FIRST DETAILED RECORDS OF CALATHOCRATUS FROM THE
IBERIAN PENINSULA, WITH DESCRIPTION OF TWO NEW SPECIES
(OPILIONES: DYSPNOI: TROGULIDAE)
Carlos E. Prieto¹*, Hay Wijnhoven² & Jochen Martens³
¹ Departamento de Zoología, Universidad del País Vasco (UPV/EHU), Apdo.644 48080-Bilbao, Spain / C. Astarloa 5, 5ºA, 48001-
Bilbao, Spain / carlos.prieto@ehu.eus
² Nijmegen, The Netherlands
³ Johannes Gutenberg-Universität, Institut für Organismische und Molekulare Evolutionsbiologie (iomE), D-55099 Mainz, Germany /
Senckenberg Research Institute, Arachnology, D-60325 Frankfurt am Main, Germany
* Corresponding author
Abstract. The genus Calathocratus is reliably recorded from the Iberian Peninsula, encompassing two new species described here-
in. Both species are small, have a similar overall morphology, like a dorsal median area devoid of papillae, but they are very differ-
ent in penial morphology. Calathocratus zaragozai sp. nov. from the Valencian-Alicante mountains in Spain has a unique subapical
stylus and a glans with large setae, while Calathocratus algarvensis sp. nov. from the Algarve region in Portugal, has a stout apical
stylus and a glans with short, robust setae. Descriptions are complemented with the first record of teratological anophthalmia in Tro-
gulidae and several observations of citizen science. A discussion on related species is included.
Key words: Spain, Portugal, penial morphology, tarsal formula, teratological anophthalmia, citizen science.
Primeras citas detalladas de Calathocratus para la Península Ibérica, con descripción de dos nuevas especies (Opiliones:
Dyspnoi: Trogulidae).
Resumen. Se cita de forma fiable el género Calathocratus para la Península Ibérica con la descripción de dos nuevas especies.
Ambas especies son pequeñas, tienen una morfología general similar, peculiarmente un área dorsal mediana desprovista de
papilas, pero son muy diferentes por la morfología penial. Calathocratus zaragozai sp. nov. de las montañas de Valencia-Alicante
en España tiene un singular estilo subapical y un glande con grandes setas, mientras que Calathocratus algarvensis sp. nov. de la
región del Algarve en Portugal, tiene un estilo apical robusto y un glande con setas cortas y robustas. Las descripciones se com-
plementan con el primer registro de anoftalmía teratológica en la familia Trogulidae y varias observaciones de ciencia ciudadana.
Se incluye una discusión sobre especies relacionadas.
Palabras claves. España, Portugal, morfología penial, fórmula tarsal, anoftalmia teratológica, ciencia ciudadana.
Introduction
The family Trogulidae Sundevall, 1833, erected for the genus
Trogulus Latreille, 1798 (and Caeculus Dufour, 1832, now a
trombidiform mite) and named for the presence of a frontal
head cap that covers and protects the pedipalps and chelicerae
(Sundevall 1833), was further characterized by Simon (1879),
who added the genera Anelasmocephalus Simon, 1879 and
Calathocratus Simon, 1879 and separated the current Di-
cranolasmatidae Simon, 1879 into their own subfamily.
Roewer (1940), in his style, based the Trogulidae systematics
on the segment numbers of leg tarsi, allowing him to erect the
genera Platybessobius Roewer, 1940 (tarsal articles legs I to
IV: 1-1-2-2), Trogulocratus Roewer, 1940 (2-1-3-3) and
Kofiniotis Roewer, 1940 (3-6-3-3). Šilhavý (1967) added the
genus Anarthrotarsus Šilhavý, 1967 (with 1-1-1-1; see also
Kontos & Martens 2022), although the taxonomic signifi-
cance of the tarsal formula was questioned both by Staręga
(1966), who synonymized Platybessobius with Calatho-
cratus, and by Martens & Chemini (1988), who accepted up
to four tarsal formulas in the genus Anelasmocephalus. The
generic systematics of the family was molecularly analyzed
by Schönhofer & Martens (2010), demonstrating that
Platybessobius and Trogulocratus are polyphyletic, passing
them to the synonymy of Calathocratus.
The genus Calathocratus, erected for Trogulus afri-
canus Lucas, 1846 from Algiers (Algeria) and later cited from
Sicily (Marcellino 1965) and Tunisia (Roewer 1950 as Tro-
gulocratus tunetanus Roewer, 1950), presently includes fur-
ther eight species (Table I, Figure 1): C. sinuosus (Sørensen,
1873) from the surroundings of Rome (Italy) and the whole
Italian Peninsula (Schönhofer & Martens 2010); C. singularis
(Roewer, 1940) from Crete Island (Greece) and reported, as
Platybessobius cf. singularis, from Greece and Anatolia by
Schönhofer & Martens (2010); C. intermedius (Roewer,
1940), also from Crete Island although Schönhofer (2013)
states that its type locality is “likely erroneous” because it was
never collected again on Crete; C. rhodiensis (Gruber, 1963)
from the island of Rhodes, C. caucasicus (Šilhavý, 1966)
from the Russian Southwest Caucasus, and C. beieri Gruber,
1968 from Southeast Anatolia (Gruber 1963, 1968; Šilhavý
1966); C. hirsutus Snegovaya, 2011 and C. minutus Sne-
govaya, 2011, both from Krasnodar Krai in the Russian Cau-
casus (Snegovaya & Chumachenko 2011). Chemeris (2013)
described Trogulocratus kyrghyzicus Chemeris, 2013 from
Kyrgyzstan, which Kury et al. (2020) placed under Calatho-
cratus. This presently is the easternmost published record of
the genus.
On the Iberian Peninsula, the family Trogulidae in-
cludes six species of Trogulus and six of Anelasmocephalus
(Schönhofer & Martens 2008; Prieto & Las Heras 2020).
Additionally, Prieto (2008) mentioned the presence in Valen-
cia [lapsus calami; in reality, province of Alicante] of an
unnamed species of Trogulocratus. Based on a molecular
analysis by Schönhofer & Martens (2010) the mentioned
sample was revealed by Schönhofer et al. (2013) as a well
separated lineage within the newly defined Calathocratus.
We describe this new Spanish species, dedicating it to the
128
Figure 1. Geographic distribution of the Calathocratus species, indicated by the initial of the specific epithet (see it in Table 1, except g=
C. singularis); moreover z= C. zaragozai sp. nov. and α= C. algarvensis sp. nov. The locality of C. kyrghyzicus is located 2000 km east of
the specified symbol and the record of C. africanus for Corsica (Simon 1879) is cited by Delfosse (2018) as Calathocratus sp. We add
the first record of Calathocratus sp . (+) f or Mor occo. / Figura 1. Distribución geográfica de las especies de Calathocratus, indicada por la
inicial del epíteto específico (ver en la Tabla 1, excepto g= C. singularis); además z= C. zaragozai sp. nov. y α= C. algarvensis sp. nov.
La localidad de C. kyrghyzicus se ubica 2000 km al este del símbolo especificado y el registro de C. africanus para Córcega (Simon
1879) es citado por Delfosse (2018) como Calathocratus sp. Añadimos la primera cita de Calathocratus sp. (+) para Marruecos.
Table I. Specific diversity and geographical distribution of the genus Calathocratus according to the tarsal formula FT (formerly, 1122 in
Platybessobius, 2133 in Trogulocratus). Body size (mm) is given for males/females. SMA refers to the presence of a smooth dorsal me-
dian area, devoid of papillae. / Tabla I. Diversidad específica y distribución geográfica del género Calathocratus según la fórmula tarsal
(1122 y 2133 es la de los antiguos géneros Platybessobius y Trogulocratus). El tamaño del cuerpo (mm) se da para machos/hembras.
SMA se refiere a la presencia de un área media dorsal lisa, desprovista de papilas.
FT Species Geographical distribution Body size: M/F SMA
1122 C. singularis (Roewer, 1940) Greece, Crete I., Anatolia 3 -
1122 C. caucasicus (Silhavy, 1966) S-Russia: Kuban 4.5 -
2122 C. africanus (Lucas, 1846) Algeria, Tunisia, Sicily 6.2 / 6.8 +
2122 C. beieri Gruber, 1968 Anatolia: Vilayet 4.7 -
2122 C. hirsutus Snegovaya, 2011 S-Russia: Krasnodar 6.4-7.2 / 7.3-8.2 -
2122 C. minutus Snegovaya, 2011 S-Russia: Krasnodar 4.1-5 / 4.7-5.2 -
2133 C. sinuosus (Sørensen, 1873) Italian Peninsula 4.8 / 5.3 +
2133 C. intermedius (Roewer, 1940) Crete Island? 5 -
2133 C. rhodiensis (Gruber, 1963) Rhodes Island 6.1 / 7.2 -
2133 C. kyrghyzicus (Chemeris, 2013) Kyrghyzstan 5.2 -
memory of a friend, Juan Antonio Zaragoza Miralles, recently
deceased. Amazingly, a sample of Calathocratus from the
southern Portuguese Algarve region, collected in 2019,
proved to belong to a morphologically different and new
species, which is also described herein. Both species clearly
differ in genital morphology and are geographically isolated
from their assumed closest relatives Calathocratus africanus
and C. sinuosus, with which they share a defined median area
largely devoid of papillation.
Material & methods
The specimens of C. zaragozai were studied, measured and
photographed by a SMZ1500 Nikon stereomicroscope and
attached hardware Digital Sight DS-L1 Nikon; serial photo-
graphs were stacked, and backgrounds edited with the Heli-
con Focus (v.7.1.1) software. The photographs of T. al-
garvensis were taken with a video camera attached to a Zeiss
SV 6 stereomicroscope and stacked with the Automon-
tage©System. Original drawings were based on sketches
directly drawn from an Olympus stereo light microscope and
a Euromex dissecting microscope, with the aid of a calibrated
drawing mirror.
The studied material has been or will be deposited in:
AXLS = Collection Axel Schönhofer, Mainz, Germany.
CHW = Collection Hay Wijnhoven, Nijmegen, The Nether-
lands. CJM = Collection Jochen Martens, deposited in the
Senckenberg Research Institute, Arachnology, Frankfurt am
Main, Germany. MNCN = Museo Nacional de Ciencias
Naturales (CSIC), Madrid, Spain. SMF = Senckenberg Re-
search Institute and Natural History Museum, Frankfurt am
Main, Germany. ZUPV = Departamento de Zoología y Bi-
ología Celular Animal, University of the Basque Country
(UPV / EHU), Leioa, Bizkaia, Spain.
Measurements in mm.
Abbreviations: M = male(s), F = female(s), J = juve-
nile(s), EMS = underground sampling station, gr. MSS =
Mesovoid Shallow Substratum group.
Additional specimens studied:
Calathocratus africanus, two males (5.95, 6.03 mm) and one
female (7.06 mm) from Fienzza, Sicily, Italy [SMFꞏ1389].
Calathocratus africanus, a juvenile (2.95 mm) from Albano,
Rom, Italy [SMFꞏ1390], referred to C. sinuosus by A.
Schönhofer (pers.comm.).
Calathocratus africanus, a female (6.9 mm) from Algiers,
Algeria [SMFꞏ1391].
Trogulocratus tunetanus Roewer, 1950, holotype (female,
7.07 mm) from Sfax, Tunisia [SMFꞏ9101].
Calathocratus sp., 1 adult, 2 juveniles from Béja, Teboursouk
129
Pass, Tunesia, 820 m, 36.4861 9.1514 (TF 1-1-2-2), H.
Meybohm leg. 1.3.2011 [CJMꞏ7132].
Calathocratus sp., 2 adults, 1 juvenile, from Tafoughalt,
Morocco, 560 m, 34.82397 2.42985, H. Wijnhoven leg.
30.9.2010 [CJMꞏ7019].
Systematics
Family TROGULIDAE Sundevall, 1833
Genus Calathocratus Simon, 1879
Calathocratus zaragozai sp. nov.
Trogulocratus sp. nov.? – Prieto, 2008: 62; fig. 17 (Valencia =
lapsus calami: Alicante).
Trogulocratus sp. Schönhofer & Martens, 2010: 62 (Alicante,
Alcoy, Santuario Font Rotja).
Calathocratus sp. – Schönhofer et al., 2013: 309 (Spain).
TYPE LOCALITY. Serra del Menejador, Parque Natural de la
Font Roja, Alcoi (Alicante, Spain; 30SYH131818, 1265 m
asl)
Holotype: male [MNCN 20.02/38432, ex-coll. ZUPVꞏ5654]
Paratypes: 1M, 3F, 4J [ZUPVꞏ3753]: Alcoi: P.N. Font Rotja:
Santuario (Alicante; 30SYH144826, 960 m), J. Zaragoza &
V. Ripoll leg. 13ꞏ05ꞏ2006 [+ sequences in GenBank for 28S
rRNA: GQ466314 and cytb: GQ466239] 1M, 1F, 1J
[ZUPVꞏ5654]: Alcoi: P.N. Font Roja: Serra del Menejador
(Alicante; 30SYH131818, 1265 m), V. Ortuño & gr. MSS
leg. 10ꞏ05ꞏ2013 (EMS-1) 3M, 1F [ZUPVꞏ5752] + 2M
[CHWꞏ496, ex ZUPVꞏ5752]: =, VIIꞏ2012 (EMS-1/3) • 2M,
3F [MNCN 20.02/38433-38437, ex-coll. ZUPVꞏ5753]: =,
Xꞏ2012 (EMS-1, 3, 4) • 3M [ZUPVꞏ5755]: Agres: Sierra de
Mariola (Alicante; 30SYH173945, 1170 m), V. Ortuño & gr.
MSS leg., 16ꞏ01ꞏ2012 (EMS-1, 2) • 2M, 1F [ZUPVꞏ5756]: =,
Xꞏ2012 (EMS-1, 2) • 1F [ZUPVꞏ5655]: =, 09ꞏ05ꞏ2013 (pit-
fall) 2F [ZUPVꞏ5652]: Barxeta: Rio Barxeta (Valencia;
30SYJ268210, 187 m), V. Ortuño & gr. MSS leg.,
08ꞏ03ꞏ2012 (EMS-1) • 2F [ZUPVꞏ5651]: Cocentaina: Mas de
Llopis (Alicante; 30SYH187927, 1220 m), V. Ortuño & gr.
MSS leg., 09ꞏ05ꞏ2013 (EMS-2) • 2M, 3F [ZUPVꞏ5750]: =,
Xꞏ2012 (EMS-1, 2) • 2F [ZUPVꞏ5653]: Confrides: Sierra de
Aitana (Alicante; 30SYH350822, 1330 m), V. Ortuño & gr.
MSS leg., 16ꞏ05ꞏ2013 (pitfall) 1M [ZUPVꞏ5751]: =,
VIIꞏ2012 (EMS-7) 1M, 4F [ZUPVꞏ5656]: Denia: P.N.
Montgó (Alicante; 31SBD495000, 322 m), V. Ortuño & gr.
MSS leg., 14ꞏ05ꞏ2013 (EMS-2, pitfall) • 5F [ZUPVꞏ5757]: =,
27ꞏ01ꞏ2012 (EMS-3, 4) 2M, 2F [ZUPVꞏ5758]: =, X-
XIꞏ2012 (EMS-3, 4) • 1F [ZUPVꞏ5657]: Quatretondeta: La
Serrella (Alicante; 30SYH351887, 1000 m), V. Ortuño & gr.
MSS leg., 14ꞏ05ꞏ2013 (pitfall) 2M, 2F [CJMꞏ9642, ex
ZUPVꞏ5759]: =, X-XI ꞏ2012 (EMS-1, 2) • 1F [ZUPVꞏ5728]:
Tibi: Sierra del Maigmó (Alicante; 30SYH060646 1080 m),
V. Ortuño & gr. MSS leg., Xꞏ2012 (EMS-1) 1F
[ZUPVꞏ5754]: =, Xꞏ2012 (pitfall) • 1M [ZUPVꞏ5650]: Xaló:
Serra de Bèrnia (Alicante; 30SYH564836, 900 m), V. Ortuño
& gr. MSS leg., 14ꞏ05ꞏ2013 (EMS-3) 1M, 2F, 1J
[ZUPVꞏ5749]: =, X-XIꞏ2012 (EMS-3, 4). 1F,1J [AXLSꞏ878]:
Bicorp, road to Casas Benalí, Cueva de los Moros (Valencia;
39.11716, -0.77613, 266 m), S. Huber & A. Schönhofer leg.,
01ꞏ06ꞏ2010 • 1M [AXLSꞏ879]: Pego/Val de Ebo (Alicante;
38.81772, -0.12256, 405 m), S. Huber & A. Schönhofer leg.,
02ꞏ06ꞏ2010 • 3F,2J [AXLSꞏ892]: Pego/Val de Ebo, cavities
(Alicante; 38.82610, -0.10824, 256 m), NE facing Pinus for-
est on slope, in dead wood, S. Huber & A. Schönhofer leg.,
03ꞏ06ꞏ2010 • 1J [AXLSꞏ895]: W Alcoy, road to Santuario de
la Font Roja (Alicante; 38.66504, -0.53734, 1142 m), leaf
litter in dense evergreen Quercus forest, S. Huber & A.
Schönhofer leg., 05ꞏ06ꞏ2010.
Total: 26 males, 41 females, 10 juveniles. Adult sex
(male/female) ratio, 0.63.
DERIVATIO NOMINIS. The species name is dedicated to the
memory of Juan Antonio Zaragoza Miralles (1953 - October
18, 2022), a friend and a Spanish arachnologist and biospele-
ologist from Alicante who, with his wife Vicen Ripoll, col-
lected the first specimens of this species.
DIAGNOSIS. A medium sized species (males 5.1-5.9 mm,
females 6.0-7.1 mm body length) with an elongated dorsal
median area free of papillae, tarsal formula 2-1-3-3, glans
with large, robust spines and subapical straight stylus, unique
in the genus.
DESCRIPTION (Figures 2-6, 7 B). Male (holotype). Body
(Figure 2), length 5.85, width 2.12, colour yellowish brown,
partly due to soil incrustations. Body elongated (length/width
ratio 2.75), abdomen slightly wider behind coxae IV. Body
integument papillose, partly obscured by soil incrustations,
dispersed papillae, each with an erect seta on top. Dorsal
surface (Figure 2A) densely covered with papillae app. 40 µm
in diameter; behind the eyes a median area almost devoid of
papillae (only scattered single papillae, more abundant in the
prosomal region), reaching almost to the posterior edge of the
scutum, the band narrowing, then widening in the interseg-
mental areas, thus forming a sequence of six bare patches,
bordered on both sides by closely placed, elevated papillae
forming an irregular crest. Tergite VI with posterior median
incision, and densely covered with slender papillae, especially
on the ribbed medial area.
Eyes and head cap (Figure 2 A, B). Ocularium plus
head cap length 0.68, head cap length (to the eyes) 0.44, head
cap width 0.60, eye interdistance 0.28, interocular distance
0.10, eye diameter 0.13. Ocularium situated near front of
prosoma, shallow and ill defined, covered by papillae smaller
than those on dorsal surface. Eyes standing very close togeth-
er , se parate d by les s than o ne e ye d iamet er. Hea d ca p branc h-
es relatively short, diverging then converging, covered with
cylindrical papillae, longer in the median area, some with a
seta on top.
Ventral surface (Figure 2C) densely covered with papil-
lae. Coxae covered with slender setose papillae.
Chelicera small (Figure 3A, 4A). Basal segment 0.57,
second segment 0.39, chela 0.37. Basal segment long and
cylindrical, with sparse setae. Chela almost as long as the
second segment.
Pedipalp (Table II, Figure 3B, 4B) small and short, pro-
truding from head cap. Trochanter cylindrical. Femur twice as
long as the trochanter, slightly curved apically, with few se-
tae, longer setae on the ventral side. Patella half as long as the
femur, convex dorsally and with long dorsal setae. Tibia 3/4
as long as femur, slightly thickened distally and with longer
dorsal setae. Tarsus shorter than trochanter, cone-shaped with
long setae on distal half, no distal claw.
Legs (Table II) with papillation on all segments except
tarsi, papillae with erect fine setae, setae length half of leg
diameter, up to leg diameter on metatarsus III and IV. Coxa,
femur, patella, tibia and metatarsus distally with grouped
130
Figure 2. Calathocratus zaragozai sp. nov. Holotype,
male 5.85 mm [ZUPVꞏ5654, Alcoi: P.N. Font Rotja:
Serra del Menejador]. Body. A, side view. B, dorsal view.
C, ventral view. The specimen has been cleaned of
adhering earth particles. Note that injured papillae leave
scarred black dots and a large scarred area in third free
sternite. Photo C. Prieto. / Figura 2. Calathocratus
zaragozai sp. nov. Holotipo, macho 5.85 mm
[ZUPVꞏ5654, Alcoi: P.N. Font Rotja: Serra del Meneja-
dor]. Cuerpo. A, vista lateral. B, vista dorsal. C, vista
ventral. El ejemplar se ha limpiado de las partículas de
tierra adheridas. Nótese que las papilas lesionadas
dejan puntos cicatrizados negros y una gran área cica-
trizada en el tercer esternito libre. Fotos, C. Prieto.
Figure 3. Calathocratus zaragozai sp. nov. Holotype
[ZUPVꞏ5654, Alcoi: P.N. Font Rotja: Serra del Meneja-
dor]. Appendices. A, right chelicera, lateral view. B, right
pedipalp, medial view. C-F: tarsi of legs I-IV, prolateral
view. Photo C. Prieto. / Figura 3. Calathocratus zarago-
zai sp. nov. Holotipo [ZUPVꞏ5654, Alcoi: P.N. Font Rotja:
Serra del Menejador]. Apéndices. A, quelícero derecho,
vista lateral. B, pedipalpo derecho, vista medial. C-F,
tarsos de las patas I-IV, vista prolateral. Fotos, C. Prieto.
131
Figure 4. Calathocratus zaragozai sp. nov. Paratype male
[ZUPVꞏ5752, Alcoi: P.N. Font Rotja: Serra del Menejador].
Appendices. A, right chelicera, lateral view. B, right pedipalp,
lateral view. C-F, left tarsi of legs I-IV, prolateral view. Scale
bar 0.5 mm. / Figura 4. Calathocratus zaragozai sp. nov.
Macho paratipo [ZUPVꞏ5752, Alcoi: P.N. Font Rotja: Serra del
Menejador]. Apéndices. A, pedipalpo derecho, vista medial.
B, quelícero, vista lateral. C-F, tarsos de las patas I-IV, vista
prolateral. Escala, 0.5 mm.
Table II. Measurements of palp and legs of the holotype of Calathocratus zaragozai sp. nov. /
Tabla II. Medidas del palpo y las patas del holotipo de Calathocratus zaragozai sp. nov.
Troch. Femur Patella Tibia Metat. Tarsus Length
Palp 0.15 0.35 0.20 0.27 0.13 1.10
Leg I 0.41 1.02 0.57 0.70 0.68 0.34 3.72
Leg II 0.61 2.01 0.75 1.23 1.36 0.64 6.60
Leg III 0.45 1.17 0.65 0.89 0.95 0.30 4.41
Leg IV 0.66 1.71 0.83 1.24 1.32 0.33 6.09
Table III. Measurements of the most relevant morphological parameters of Calathocratus zaragozai sp. nov. The coefficient of variation
(CV, defined as the ratio of the standard deviation to the mean) is expressed as percentage. / Tabla III. Medidas de los parámetros mor-
fológicos más relevantes de Calathocratus zaragozai sp. nov. El coeficiente de variación (CV, relación entre la desviación típica y la me-
dia) se expresa en porcentaje.
MALES (n=15) FEMALES (n=20)
Mean Range CV (%) Mean Range CV (%)
Body (length) 5.59 5.17 – 5.88 3.48 6.46 6.00 – 7.08 4.12
Body (width) 2.12 1.96 – 2.29 4.38 2.50 2.29 – 2.74 4.56
Leg II (length) 6.49 6.15 – 6.86 2.82 6.85 6.36 – 7.34 3.46
Tarsus II (length) 0.63 0.59 – 0.70 4.88 0.60 0.56 – 0.65 3.90
Tarsus II (ratio l/w) 2.66 2.44 – 2.83 4.64 2.65 2.46 – 2.91 4.95
Ocularium + hood 0.71 0.62 – 0.79 6.11 0.78 0.66 – 0.85 23.04
Hood branches 0.44 0.37 – 0.50 7.35 0.51 0.42 – 0.60 8.97
Ocularium (width) 0.30 0.28 – 0.33 4.80 0.33 0.29 – 0.37 22.69
B/w eyes distance 0.11 0.07 – 0.13 16.12 0.13 0.10 – 0.17 15.72
Eye (diameter) 0.13 0.11 – 0.15 11.00 0.12 0.10– 0.15 10.04
long papillae shielding the corresponding articulation joints.
Dorso-distal metatarsus margin is protruding, especially in
metatarsi III and IV (Fig. 3 E-F) and covered with long papil-
lae (Fig. 3 C-D for legs I-II). Leg tarsi I-IV with tarsal formu-
la 2-1-3-3. Tarsus II large and cylindrical, 2.65 times (range
for 15 males 2.44-2.91) its width.
Penis (Figure 5 A-B). Length 1.35. Penis truncus slen-
der, 1.25 mm long, its base drawn out in two short indistinct
roots, dorsoventrally flattened, more distally cylindrical.
Glans penis (Figure 5 C-E) covered with large and robust
setae (25 µm) evenly distributed (dorsal, lateral and ventral
side each with about 14 setae), ventrobasal glans area without
setae. Stylus attached subapically (50 µm long) at an angle of
approximately 50°, almost straight and parallel-sided, in lat-
eral view with a rectangular tip.
132
Figure 5. Calathocratus zaragozai sp.
nov. Paratype [ZUPVꞏ5752, Alcoi: P.N.
Font Rotja: Serra del Menejador]. Penis.
A-B, ventral and lateral view. C-E: glans
penis in ventral, dorsal and lateral view.
Scale bar: A-B 0.5 mm, C-E 0.05 mm. /
Figura 5. Calathocratus zaragozai sp.
nov. Paratipo [ZUPVꞏ5752, Alcoi: P.N.
Font Rotja: Serra del Menejador]. Pene.
A-B, vista ventral y lateral. C-E: glande
en vistas ventral, dorsal y lateral. Esca-
las: A-B, 0.5 mm; C-E, 0.05 mm.
Female. As in the male, but larger (6.0-7.1 mm) and wider
(mean 2.5 mm), longer leg II (mean 6.85 mm), but propor-
tionally longer in males (1.16 times the body length vs 1.06 in
females); the head cap and eye dimensions seem quite varia-
ble compared to males (Table III). Ovipositor (Figure 6A) as
a frustoconical and pleated trunk, distally with two lateral
lobes flanking two internal lips. Surface covered with sensil-
lar papillae, more sclerotized distally.
GEOGRAPHICAL DISTRIBUTION AND ECOLOGY. This Iberian
endemic is known from the northern mountainous part of the
province of Alicante (12 localities) and the southern part of
the province of Valencia (Barxeta and Bicorp), an area limited
to the north by the Serpis River and to the south by the
Vinalopó River (Figure 8). The species has been found both
in the summit areas of the Alicante calcareous mountains,
between 900 and 1330 m asl, as well as at lower altitudes
(P.N. of Montgó at 322 m, Barxeta at 187 m), suggesting a
more extensive geographical distribution than currently
known.
Most samples of C. zaragozai sp. nov. were obtained by
the entomologist Vicente Ortuño and his team in the calcare-
ous mountains north of Alicante, by means of MSS samplings
(Milieu Souterrain Superficial, currently Mesovoid Shallow
Substratum, traps buried up to 1 m deep in pebbled slopes),
and pitfall traps, sampling invertebrates active on soil surface.
Pit falls contained only 6 females, the MSS traps 24 males, 26
females and 2 juveniles. Samples from Barxeta (in alluvial
shallow substratum, as described by Ortuño et al. 2013) pro-
vided two females. Thus, C. zaragozai sp. nov. likely favours
endogean habitats.
This harvestman species was found year-round: summer
2012 (12 %: 6M, 1F), autumn 2012 (45 %: 11M, 15F, 1J),
winter 2013 (17 %: 3M, 7F) and spring 2013 (27 %: 4M, 11F,
1J). In spring and autumn, they are more active, due to the
Mediterranean climate, with dry and hot summers and pre-
cipitation predominant in spring and autumn. At higher alti-
tudes this effect is less distinct (Chatzaki et al. 2009). Juve-
niles occurring in spring and autumn suggest an eurychronous
reproduction strategy with adults surviving more than one
year and overlapping generations (Belozerov, 2012), as in
other trogulids.
Calathocratus algarvensis sp. nov.
TYPE LOCALITY: São Marcos da Serra (Algarve, Portugal;
29SNB55253263, 97m asl), H. Wijnhoven leg. 3.5.2019.
Holotype: male [CJM 8153].
Paratypes: 1M, 2F, 2J [CJM 9643]; same data as holotype.
DERIVATIO NOMINIS. The species name is derived from the
Portuguese Algarve district.
DIAGNOSIS: A small and slender trogulid species (5.15-5.27
body length, 2.13-2.15 body width; n = 2) with slender legs;
elongated dorsal median area devoid of papillae, tarsal formu-
la 2-1-3-3, penis nearly parallel-sided, robust glans with many
massive setae.
DESCRIPTION (Figures 9-11). Male (holotype). Measure-
ments in Table IV.
Body male body length 5.27, width 2.15; yellowish
brown, partly due to soil incrustations. Body small and elon-
gated (length/width ratio 2.45), with the abdomen slightly
wider behind leg coxae IV. Dorsal surface (Figure 9 B) dense-
ly covered with papillae app. 50 µm in diameter; behind the
eyes a median area devoid of papillae, ranging almost to the
posterior edge of the scutum, the area bordered on both sides
by closely placed, elevated papillae forming an irregular crest.
Tergite VI with posterior median cleft, covered with similar
papillae as the posterior scutum region (Fig. 9).
Eyes and head cap (Figure 9). Eyes plus head cap length
0.53; head cap length (to the eyes) 0.42; head cap width 0.60
mm; eye interdistance 0.30; interocular distance 0.08; eye
diameter 0.11. Eyes situated at front of prosoma, ocularium
shallow and ill defined, covered by papillae smaller than those
on dorsal surface. Eyes standing close together, separated by
approximately one eye diameter. Head cap branches relatively
short, covered with long and slender papillae, each with a seta
on top, surface morphology obscured by soil incrustations.
Ventral surface densely covered with papillae. Few soil
incrustations, more along the ventral side. Coxae covered
with thin setose papillae.
Chelicera small (Figure 10 C). Chelicera: basal segment
0.86, second segment 0.35, chela 0.38. Basal segment long
and cylindrical, with sparse setae. Chela almost as long as the
second segment.
133
Figure 6. Calathocratus zaragozai sp.
nov. A, ovipositor of a paratype female
6.6 mm long [ZUPVꞏ5753, Serra del
Menejador]. B, eyeless ocularium and
head cap branches of a teratological
female 6.47 mm long [ZUPVꞏ5749,
Serra de Bèrnia]. C, leg pairs I-II of a
teratological female 6.33 mm long
[ZUPVꞏ5759]. Photo C. Prieto. / Figura
6. Calathocratus zaragozai sp. nov. A,
ovipositor de una hembra paratípica de
6.6 mm [ZUPVꞏ5753, Serra del
Menejador]. B, oculario sin ojos y
ramas del capuchón de una hembra
teratológica de 6.47 mm [ZUPVꞏ5749,
Serra de Bèrnia]. C, patas I-II de una
hembra teratológica de 6.33 mm
[ZUPVꞏ5759]. Fotos, C. Prieto.
Figure 7. A, Calathocratus sp., female
6.18 mm [ZUPVꞏ5597, Grândola]. B,
Calathocratus zaragozai sp. nov. male
paratype 5.31 mm [ZUPVꞏ5752, Serra
del Menejador]. C-F, Calathocratus
africanus (Lucas, 1846). C, male 6.03
mm [SMFꞏ1389, Fienzza, Sicily]. D,
female 6.7 mm [SMFꞏ1391, Algiers,
Algeria]. E, female 7.07 mm, holotype
of Trogulocratus tunetanus Roewer,
1950 [SMF-9101, Sfax, Tunisia]. F,
female (7.06 mm) [SMFꞏ1389, Fienzza,
Sicily]. Photo C. Prieto. / Figura 7. A,
Calathocratus sp., hembra 6.18 mm
[ZUPVꞏ5597, Grândola]. B, Calatho-
cratus zaragozai sp. nov., paratipo
macho 5.31 mm [ZUPVꞏ5752, Serra
del Menejador]. C-F, Calathocratus
africanus (Lucas, 1846). C, macho 6.03
mm [SMFꞏ1389, Fienzza, Sicilia]. D,
hembra 6.7 mm [SMFꞏ1391, Argel,
Argelia]. E, hembra 7.07 mm, holotipo
de Trogulocratus tunetanus Roewer,
1950 [SMF-9101, Sfax, Túnez]. F,
hembra (7.06 mm) [SMFꞏ1389,
Fienzza, Sicilia]. Fotos, C. Prieto.
Table IV. Measurements of pedipalp and legs of Calathocratus algarvensis sp. nov. Male holotype (in brackets female paratype). /
Tabla IV. Medidas del pedipalpo y las patas de Calathocratus algarvensis sp. nov. Macho holotipo (en paréntesis, paratipo hembra)
Troch. Femur Patella Tibia Metat. Tarsus Length
Palp 0.15 (0.15) 0.3 (0.35) 0.15 (0.2) 0.25 (0.25) 0.15 (0.12) 1.00 (1.07)
Leg I 0.5 (0.4) 1.10 (1.1) 0.5 (0.4) 0.8 (0.8) 0.75 (0.8) 0.35 (0.3) 4.00 (3.80)
Leg II 0.7 (0.65) 2.0 (2.05) 0.6 (0.65) 1.3 (1.3) 1.4 (1.4) 0.6 (0.6) 6.60 (6.65)
Leg III 0.5 (0.5) 1.2 (1.2) 0.5 (0.6) 0.9 (0.9) 1.0 (0.95) 0.3 (0.3) 4.40 (4.45)
Leg IV 0.7 (0.6) 1.9 (1.85) 0.65 (0.7) 1.4 (1.35) 1.4 (1.35) 0.4 (0.4) 6.45 (6.25)
134
Figure 8. Geographical distribution of Calathocratus zaragozai sp.
nov. in the Comunidad Valenciana. Red dots, specimen-based re-
cords; white squares, photographic records (BV, Biodiversidad Virtual;
iN, iNaturalist). A few sites are cited and the type locality is underlined.
The inset shows the mapped area for C. zaragozai and the sites of the
other Iberian species of Calathocratus. / Figura 8. Distribución geo-
gráfica de Calathocratus zaragozai sp. nov. en la Comunidad Valen-
ciana. Círculos rojos, registros basados en especímenes; cuadrados
blancos, registros fotográficos (BV, Biodiversidad Virtual; iN, iNatu ra-
list). Se citan algunas localidades y se subraya la localidad tipo. El
recuadro muestra el área cartografiada para C. zaragozai y las locali-
dades de las otras especies ibéricas de Calathocratus.
Pedipalp (Figure 10 A, Table IV) small and robust. Trochan-
ter cylindrical, unarmed. Femur slightly enlarged distally,
twice as long as the trochanter, with few setae, longer setae on
the ventral side. Patella half as long as the femur, with long
dorsal setae. Tibia slightly shorter than femur, slightly en-
larged distally, with setae that are longer on dorsal side. Tar-
sus as long as trochanter, cone-shaped with long setae on
distal half, no distal claw.
Legs (Table IV). Papillation on all segments except tarsi,
papillae with erect fine and long setae, setae length half of leg
diameter, and up to leg diameter. Coxa, femur, patella, tibia
and metatarsus distally with grouped long papillae shielding
the corresponding articulation joints. Dorso-distal metatarsus
margin is protruding, especially in metatarsi III and IV (Fig-
ure 11 C-D) and covered with long papillae. Leg tarsi I-IV
with tarsal formula 2-1-3-3 (Figure 11 A-D). Tarsus II large
and cylindrical, 2.70 times its width. Claws hook-shaped,
claw on leg I, III and IV most robust, in leg I almost as large
as last article, larger in legs III and IV; smooth setae on all
tarsi, including single long setae dorsally and distally.
Penis (Figure 10 H-I). Penis length 1.40. Truncus slender,
almost parallel-sided, its base drawn out in two short indis-
tinct roots, dorsoventrally flattened, more distally cylindrical,
only slightly broadening below the glans. Glans penis (Figure
10 E-G) elongated and robust, covered with short, robust and
curved setae on dorsal, ventral and lateral side. Glans contin-
uously tapering into the stylus. Stylus slightly curved dorsally,
with blunt top.
Female (Figure 9, 10 B, D, 11 E-H). As in the male, but
slightly larger (5.49) and wider (2.32), with longer legs (Table
IV).
GEOGRAPHICAL DISTRIBUTION AND ECOLOGY. All speci-
mens were collected from under a decomposing tree trunk
(Figure 12) on a hill side close to a watercourse, linked with
th e Od elouca riv er, a tr ibu tary of the Arade r ive r, with typ ical
Mediterranean vegetation characterized by holly oak (Quer-
cus ilex), common gorse (Ulex europaeus) and rockrose (Cis-
tus sp.). The location is part of the Monchique Range, so
probably this species is more wide-spread in the calcareous
mountainous regions of the Algarve.
Calathocratus sp.
MATERIAL: 1M, 1F [ZUPVꞏ5597]: Grândola: Herdade Ribei-
ra Abaxo (Setúbal, Portugal; 29SNC363171, 241 m), R.
Plăiașu leg., 05ꞏ11ꞏ2016. The removed penis was missing.
These two specimens differ in certain aspects from C. al-
garvensis sp. nov. They are larger (male is 5.57 vs 5.15-5.27;
female (Figure 7 A) is 6.18 vs 5.49). In addition, these speci-
mens from Grândola have longer, thinner hood branches, eyes
larger and set closer together, a tuft of longer papillae behind
eyes, and slenderer tarsus II (3+ times its width) than C. al-
garvensis. Possibly, a third Iberian species is involved; addi-
tional material is needed for a sound decision.
Discussion
Variability in C. zaragozai sp. n ov. Samples of C. zaragozai
(Table III) show no significant morphological variability
(variation coefficient is less than 10%). The two specimens
from the lowest altitude (Barxeta) are the largest and speci-
mens from Montgó have slightly smaller eyes with larger
interocular distance (ratio 0.6-0.85) than the other measured
specimens (ratio 0.9-1.3).
Male/female ratio is 0.63, increasing with altitude, being
0.24 (4M, 17F) at lower altitudes, 0.50 (5M, 10F) around
1000 m asl, and 1.21 (17M, 14F) at altitudes above 1170 m.
An increasing sex ratio at higher altitudes has been found for
Odonata (Crowley & Johansson 2002). Generally, males
move around more than females, resulting in a biased sex
ratio for pitfall traps. Body size tends to decrease with altitude
(not significant): mean for males / females is 5.79 / 6.6 mm at
lower altitudes, 5.66 / 6.35 mm around 1000 m asl and 5.53 /
6.33 mm at higher altitudes.
The juvenile ratio (no. of immatures / total number =
0.13), agrees with figures for other taxa from Alicante moun-
tains such as Dicranolasma soerensenii (0.092, 119 juveniles,
1176 adults), but differs significantly from Trogulus cf. lusi-
tanicus (0.43, 183 juveniles, 244 adults) (unpublished results
based on the collection ZUPV). This different pattern could
be related to the large size of T. lusitanicus, with males larger
than 10 mm and females reaching 15 mm long, whose propor-
tionately larger juveniles are probably more active than small-
er immature stages of Dicranolasma or Calathocratus.
Teratologies in C. zaragozai sp. nov. A female specimen
from Serra de Bèrnia [ZUPV5749] presents teratological
anophthalmia (Figure 5B); the lack of the eyes makes the
hood branches appear thinner at the base. Blindness is rarely
reported in Opiliones; it was not reported by Mitov (1995) in
135
Figure 9. Calathocratus algarvensis s p.
nov. Female paratype, 5.49 mm [CJM
8153]. Body. A, side view. B, dorsal
view. C, ventral view. D, head cap and
prosomal region. Photo P. Schwen-
dinger. / Figura 9. Calathocratus
algarvensis sp. nov. Paratipo hembra,
5.49 mm [CJM 8153]. Cuerpo. A, vista
lateral. B, vista dorsal. C: vista ventral.
D, capuchón y región prosómica.
Fotos, P. Schwendinger.
Table V. Photographic records of Calathocratus ssp. from citizen science in two repositories (BV, www.biodiversidadvirtual.org; iN,
www.inaturalist.org) are identified by their observation number. . Only records from Alicante and Valencia can be assigned with certain-
ty to C. zaragozai sp.nov. They are ordered chronologically by the observation date. Decimal geographic coordinates of iNaturalist rec-
ords have been converted to MGRS coordinates with hectare precision. / Tabla V. Registros fotográficos de Calathocratus ssp. proced-
entes de ciencia ciudadana en dos repositorios (BV, www.biodiversidadvirtual.org; iN, www.inaturalist.org) identificados por su nº de ob-
servación. Sólo los registros de Alicante y Valencia pueden asignarse con certeza a C. zaragozai sp.nov.Están ordenados cronológi-
camente por su fecha de observación. Las coordenadas geográficas decimales de los registros de iNaturalist se han transformado en
coordenadas MGRS con precisión hectométrica.
Record Nº Locality, Province coord. MGRS Photographer Observed Uploaded
BVꞏ310435 Villajoyosa, Alicante 30SYH 380684 Daniel García 18ꞏ02ꞏ2012 18ꞏ02ꞏ2012
BVꞏ454462 Xátiva, Valencia 30SYJ 151179 Francisco Rodriguez 29ꞏ04ꞏ2013 02ꞏ05ꞏ2013
iNꞏ138469635 [Xàtiva,] Valencia 30SYJ 152177 faluke 29ꞏ04ꞏ2013 12ꞏ10ꞏ2022
iNꞏ107212534 Arenas de San Pedro, Ávila 30TUK 201522 jmsandicor18 20ꞏ02ꞏ2022 21ꞏ02ꞏ2022
iNꞏ111201258 Sta. Margarida da Serra, Grândola, Setúbal 29SNC 372168 joaodpontes 29ꞏ03ꞏ2022 12ꞏ04ꞏ2022
iNꞏ117050416 Teulada, Alicante 31SBC 481905 eva_b_s 24ꞏ04ꞏ2022 15ꞏ05ꞏ2022
his extensive search for teratologies in Bulgarian harvestmen.
Only two records are known, both belonging to Phalangioi-
dea, Togwoteeus biceps (Thorell, 1877) by Holmberg & Kok-
ko (1983), and two specimens of Lophopilio palpinalis
(Herbst, 1977) by Noordijk et al. (2016). Thus, this report is
the first observation for the suborder Dyspnoi and the family
Trogulidae.
The other observed teratology is the shortened right leg
II of a female (body length 6.33 mm, Figure 5C) from La
Serrella [ZUPVꞏ5759]; right leg is 4.93 mm long (left leg 6.69
mm), with a normal femur, a little shorter patella and tibia,
and much shorter metatarsus and tarsus, which, being fused,
barely reach a quarter of the corresponding parts on the left
leg.
Citizen Science records of Calathocratus spp. First citizen
science record for the Iberian Peninsula was uploaded in 2012
at the Biodiversidad Virtual repository, while most recent
records were uploaded in 2022 at iNaturalist repository (Ta-
ble V; Figure 13). All photographic records show the large
and undivided tarsus II, a diagnostic character for Calatho-
cratus, and the smooth median dorsal area delimited laterally
by a sinuous crest formed by longer papillae, which is an
exclusive character of Western Calathocratus taxa. Three of
the five records (note that ones by Francisco Rodríguez and
“faluke”, his nickname, are the same) come from the provinc-
es of Valencia and Alicante and match the distribution area of
C. zaragozai sp. nov. Although identification of trogulids
136
Figure 10. Calathocratus algarvensis sp. nov. A, C, E-I:
holotype male [CJMꞏ8153], B, D: paratype female
[CJMꞏ9643]. A-B, right pedipalp, retrolateral view. C-D,
right chelicera, retrolateral view. E-G, glans penis in
ventral, dorsal and lateral view. H-I, penis in ventral and
lateral view. Scale bar: A-D, H-I 0.5 mm; E-G 0.05 mm. /
Figure 10. Calathocratus algarvensis sp. nov. A, C, E-I:
macho holotipo [CJMꞏ8153], B, D: hembra paratipo
[CJMꞏ9643]. A-B, pedipalpo derecho, vista retrolateral.
C-D, quelícero derecho, vista retrolateral. E-G, glande
del pene en vistas ventral, dorsal y lateral. H-I, pene en
vistas ventral y lateral. Escalas: A-D, H-I, 0.5 mm; E-G,
0.05 mm).
Figure 11. Calathocratus algarvensis sp. nov. A-D: left
tarsi of legs I-IV of the holotype male [CJMꞏ8153], pro-
lateral view. E-H, left tarsi of legs I-IV of the paratype
female [CJMꞏ9643]. Scale bar 0.5 mm./ Figura 11.
Calathocratus algarvensis sp. nov. Tarsos izquierdos de
las patas I-IV, vista prolateral. A-D: macho holotipo
[CJMꞏ8153]. E-H, hembra paratipo [CJMꞏ9643]. Escala,
0.5 mm.
137
Figure 12. Habitat of Calathocratus algarvensis sp. nov.
at the type locality, São Marcos da Serra (Algarve,
Portugal), with holly oak (Quercus ilex), common gorse
(Ulex europaeus) and rockrose (Cistus sp.). The speci-
mens were found under the tree trunk in the bottom
right. Photo H. Wijnhoven, 3.5.2019. / Figura 12. Hábitat
de Calathocratus algarvensis sp. nov. en la localidad
tipo, São Marcos da Serra (Algarve, Portugal), con
encina (Quercus ilex), aulaga (Ulex europaeus) y jara
(Cistus sp.). Los especímenes fueron encontrados
debajo del tronco de la parte inferior derecha. Foto H.
Wijnhoven, 3.5.2019.
Figure 13. Iberian photographic records in two citizen
science repositories (Table 5), Biodiversidad Virtual
(top image, the first photographic record) and iNaturalist
(bottom images connected by an Iberian map with their
provenances). Note in each photo the smooth median
area delimited by two undulating crests and the diagnos-
tic undivided tarsus II, and that the specimen
iNꞏ138469635 seems to be eating a small land snail. /
Figura 13. Registros fotográficos ibéricos en dos reposi-
torios de ciencia ciudadana (Tabla 5), Biodiversidad
Virtual (imagen superior, el primer registro fotográfico) e
iNaturalist (imágenes inferiores conectadas por un
mapa de la península con sus procedencias). Nótese en
cada fotografía el área mediana lisa delimitada por dos
crestas ondulantes y el diagnóstico tarso II indiviso, y
que el espécimen iNꞏ138469635 parece estar comiendo
un pequeño caracol terrestre.
from field photographs is not reliable, we consider that those
three records likely represent C. zaragozai sp. nov.
There is a record from Santa Margarida da Serra, less
than one km apart the site of Calathocratus sp. from Herdade
Ribeira Abaxo [ZUPVꞏ5597], both from Grândola (Setúbal,
Portugal). Although the distance to the type locality of C.
algarvensis sp. nov. is barely 90 km, the above-mentioned
morphological differences suggest it is a different species.
The latest record is from Arenas de San Pedro (Ávila,
Spain), a site separated by more than 400 km from Alican-
te/Valencia and Algarve regions. Taking in account that the
habitat of the Ávila population is not Mediterranean (it was
found among lichens and mosses in a humid gorge of a granit-
ic mountain range), it could belong to a fourth species.
138
Systematics. Schönhofer & Martens (2010) synonymized
Trogulocratus and Platybessobius with Calathocratus, based
on polyphyly by molecular analysis, and, as a consequence,
the genus now contains ten species (Table I).
Beyond the generic systematics, species delimitation of these
small trogulids was a challenge ever since. Though only few
species have been described to date, a revision of the classical
material of Lucas, Simon and Sørensen was never achieved.
Thus, it is unproven how many species actually occur in
North Africa and, even more strange, if an African species, C.
africanus, actually occurs in southern Italy as suggested by
determinations by Roewer (see above) and indication by
Marcellino (1965). Being aware that the much larger species
of the genus Trogulus put difficult problems to taxonomists,
which up to now could only partly be solved by means of
molecular genetics, the taxonomic situation in Calathocratus
seems to be even harder. Fairly rich and geographically wide-
ly scattered material from Turkey was attributed to highly
distinct clades resulting from molecular genetic studies, ap-
parently indicating species status (Schönhofer & Martens
2010). But a species characterization by (genital) morphologi-
cal data was hardly possibly and thus was not further fol-
lowed. Enormous cryptic diversity of Calathocratus in the
Near East, and, in addition, in the Caucasus range in Russia,
Georgia, Azerbaijan and Armenia is to be expected (Sne-
govaya & Chumachenko 2011, J.M. unpublished data).
Species from North Africa and the two new ones from
Iberian Peninsula represent the westernmost localities of the
genus. These populations including such from Sicily appar-
ently form a group of closely related species. C. sinuosus and
C. africanus have to be involved in a comparison with both
new species described here, since only these species share a
characteristic feature with the new species: the mid dorsal
scutal area of opisthosoma being devoid of papillae.
C. zaragozai sp. nov. was first mentioned as Trogulo-
cratus sp. nov. by Prieto (2008; a sample of 1M, 3F and 4J)
based on the tarsal formula 2-1-3-3, and tarsus II with dimen-
sions intermediate between those of C. intermedius and C.
tunetanus. A genetical analysis of a sample of the Valencia-
Alicante population of Calathocratus (Trogulocratus sp.’)
confirmed its status as at least a species-level lineage
(Schönhofer & Martens 2010, Figure 6); six Calathocratus
species were analyzed by them: C. africanus, C. beieri,
Platybessobius caucasicus, P. singularis, Trogulocratus sinu-
osus and Trogulocratus sp. from Spain, showing (i.e.
Schönhofer & Martens 2010, Figure 6) that the three samples
of Trogulocratus sinuosus, two of them from mainland Italy
and coded as “species awaiting formal description” did form a
clade with C. africanus, while Trogulocratus sp.’ from Spain
did not cluster with them.
Calathocratus africanus (Lucas, 1846) was described
from Algeria and reported later from Tunisia as its synonym
Trogulocratus tunetanus Roewer, 1950 and Sicily (Marcelli-
no 1965) (Figure 7 C-F). Body size (M 6.0 mm, F 6.7-7.1
mm) and leg lengths (7.5 and 7.9 mm for leg II) and similar
dorsal habitus demonstrate that C. africanus is closely related
to C. zaragozai sp. nov. A similar general habitus is found
for the specimens from Tafoughalt [CJMꞏ7019], representing
the first record of Calathocratus for Morocco and the most
western record for Northern Africa. Figure 7 C-F shows spec-
imens from Algeria, Tunisia and Sicily where the sinuous line
that borders the smooth median dorsal area seems not raised
into a crest of elongate papillae as in C. zaragozai sp. nov.
The glans penis in C. africanus is provided with spines, more
robust and numerous and with the apex incised (Marcellino
1965), and the stylus is curved and placed on top of the glans,
while in C. zaragozai sp. nov. the stylus is straight and insert-
ed subapically on the glans. Also, C. algarvensis sp. nov.
seems to be closely related to Calathocratus africanus, but
the blunt stylus top and small body size of C. algarvensis sp.
nov. point to a different relationship.
Calathocratus sinuosus (Sørensen, 1873) was described
from Gennazano (Italy), its status was resolved by Schönhofer
& Martens (2010). Morphometric data and description of
Marcellino (1965 sub Trogulocratus apenninicus) indicate
that this species has a small body size (4.8 and 5.3) and a
smooth median dorsal area; the penis has a glans with few
spines and an apical, tapering stylus; the tarsus II is three
times its width (two times in C. africanus). Our specimens of
C. zaragozai sp. nov. have an intermediate tarsus II ratio of
2.65 in mean (range 2.44-2.91).
Although there have been considerable taxonomical
achievements in recent decades on Trogulidae of the Iberian
Peninsula (i.e. Prieto & Las Heras 2020, Schönhofer 2009,
Schönhofer & Martens 2008, 2010), studies on trogulid spe-
cies on this large and mountainous subcontinent continue to
offer fascinating opportunities. ‘Still far from understanding
species boundaries within the CPT complex [Calathocratus
Simon, 1879, Platybessobius Roewer, 1940 and Trogulo-
cratus Roewer, 1940] we glance at the amount of cryptic
diversity, far beyond expectation for the European and Near
East fauna’ (Schönhofer & Martens 2010). In most parts of
the Caucasus range, Calathocratus species are widely distrib-
uted and locally rather common. Also, this fauna with three
species documented (Table 1) is largely unexplored (J.M.
personal observation). Obviously, Calathocratus represents
an ancient group for which the western distributional range
and its species radiation is still poorly understood.
Acknowledgements
To Vicente Ortuño (Universidad de Alcalá, Madrid, Spain) for
putting at CP’s disposal the rich collection of Opiliones collected by
his team by trapping in the MSS of the calcareous mountains of
Alicante. To Rodica Plăiașu (Institute of Speleology "Emil Raco-
vita", Romania) by the donation of the Calathocratus sample from
Grândola. To Daniel García, Francisco Rodríguez faluke, Juan
Manuel Sánchez Díaz-Corralejo –jmsandicor18, joaodpontes, and
eva_b_s for allowing the use of the photographs that support the
citizen science records cited in the text. To Bernard Bal (bbinsecte)
by its first determination at iNaturalist as a Calathocratus population
outside Valencia and Alicante provinces. To Axel Schönhofer (Jo-
hannes Gutenberg Universität Mainz, Mainz, Germany) for his
molecular analysis of the new Spanish species, the authorization to
include the material of C. zaragozai from his collection and for the
improvements suggested after reviewing the final version of the
manuscript. To Peter Schwendinger at the MHNG (Geneva, Switzer-
land), who kindly provided the automontage images of C. algarven-
sis sp. nov. To the Feldbausch Foundation and the Wagner Founda-
tion at the Fachbereich Biologie of Mainz University (Germany)
which continuously sponsored J.M. for fieldwork and lab work in
Europe and Asia. We cordially thank all friends, colleagues and
institutions.
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Article
Information on seasonal adaptations in the life cycles of harvestmen (order Opiliones) summarized in this review reveals a great diversity of these arachnids with respect to duration, voltinism, and phenology of their life cycles, as well as to the number and ontogenetic position of the dormant stages required for survival during the winter season and also for synchronizing development with seasonal climate rhythms. Most harvestmen have stenochronous development with univoltine life cycles synchronized by arrest of development in hibernating eggs or (rarely) diapausing nymphs and adults. The number and arrangement of dormant stages represent stable species-specific traits, with some rare cases of interpopulation differences (e.g., in Phalangium opilio). Eurychronous harvestmen exhibit the year-round occurrence of main ontogenetic stages which show equal abilities for either active development or quiescence (depending upon the external factors). Two alternative types of development arrest are common in univoltine opilionids: (1) diapause induced at the early stage of embryogenesis and terminated during cooling (in Opilio parietinus and eleven other species) with transformation into postdiapause quiescence; (2) cold quiescence enforced by low temperatures at the last stage of egg development just before hatching (in Phalangium opilio and four other species). In conclusion, the systems of seasonal adaptations in Opiliones are compared with those in other arachnids, insects, and crustaceans. Some promising directions in the study of seasonal adaptations in opilionid life cycles are suggested.
Nuovi dati e problemi aperti su alcuni Opilioni Italiani (Arachnida Opiliones) [New data and open problems on some Italian Opiliones
  • P M E Brignoli
  • Rafaelli
BRIGNOLI, P.M. & E. RAFAELLI 1978. Nuovi dati e problemi aperti su alcuni Opilioni Italiani (Arachnida Opiliones) [New data and open problems on some Italian Opiliones (Arachnida Opiliones)].
Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgeführten zoologischen Exkursionen. VII. Scorpiones und Opiliones
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