The Italian species of Pritha (Araneae, Filistatidae): a critical revision and description of two new species

Abstract

The species of Pritha Lehtinen, 1967 in Italy are revised for the first time, and a preliminary putative diagnosis for the genus is provided. Two new Italian species are described: Pritha parva sp. nov. (male and female) and P. sagittata sp. nov. (male and female). P. pallida (Kulczyński, 1897) and the type species P. nana (Simon, 1868) are entirely re-described. The copulatory organs, some other somatic characters and the habitus of each sex of the four Italian species are illustrated and photographed at high magnifications. P. debilis (Simon, 1911) and P. vestita (Simon, 1873) are removed from the synonymy with P. nana and revalidated to species status, and a lectotype is designated for the latter. P. pallida is recorded from Spain and Greece for the first time. Distribution, behavioural aspects and ecology of the four Italian species are discussed and photographs of live specimens, webs and habitats are provided.

Full text

Accepted by F. Labarque: 24 Jan. 2017; published: 15 Mar. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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Copyright © 2017 Magnolia Press
Zootaxa 4243 (2): 201
–
248
http://www.mapress.com/j/zt/
Article
201
https://doi.org/10.11646/zootaxa.4243.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:29EF88AB-493A-47B9-BF6B-B27E54438CF2
The Italian species of Pritha (Araneae, Filistatidae): a critical revision and
description of two new species
CARLO MARIA LEGITTIMO
1,2
, ENRICO SIMEON
1,3
, PIERGIORGIO DI POMPEO
1,4
& ALESSANDRO KULCZYCKI
1,5
1
ARACNOFILIA
—
ASSOCIAZIONE ITALIANA DI ARACNOLOGIA
—
WWW.ARACNOFILIA.ORG
2
Città della Pieve (PG), Italy. E-mail: carlomarialegittimo@yahoo.it
3
Cervignano del Friuli (UD), Italy. E-mail: enricosimeon@gmail.com
4
Pescara (PE), Italy. E-mail: piergiorgio.dipompeo@alice.it
5
Roma (RM), Italy. E-mail: alessandro.kulczycki@gmail.com
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Family Filistatidae Ausserer, 1867 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Subfamily Prithinae Gray, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .203
Pritha Lehtinen, 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Pritha nana (Simon, 1868). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Pritha pallida (Kulczyński, 1897) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Pritha parva sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
Pritha sagittata sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
Pritha vestita (Simon, 1873) spec. reval.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Pritha debilis (Simon, 1911) spec. reval.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238
General ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Abstract
The species of Pritha Lehtinen, 1967 in Italy are revised for the first time, and a preliminary putative diagnosis for the
genus is provided. Two new Italian species are described: Pritha parva sp. nov. (male and female) and P. sagittata sp.
nov. (male and female). P. pallida (Kulczyński, 1897) and the type species P. nana (Simon, 1868) are entirely re-described.
The copulatory organs, some other somatic characters and the habitus of each sex of the four Italian species are illustrated
and photographed at high magnifications. P. debilis (Simon, 1911) and P. vestita (Simon, 1873) are removed from the syn-
onymy with P. nana and revalidated to species status, and a lectotype is designated for the latter. P. pallida is recorded
from Spain and Greece for the first time. Distribution, behavioural aspects and ecology of the four Italian species are dis-
cussed and photographs of live specimens, webs and habitats are provided.
Key words: taxonomy, ecology, redescription, revalidation, Palearctic, Europe, calamistrum, Haplogynae

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Introduction
The Filistatidae Ausserer, 1867 are a globally distributed family presently comprised by 19 genera and 148 species
(World Spider Catalog 2016). They are the sole entirely cribellate family of the traditional Haplogynae, and differ
from cribellate entelegynes by having the labium fused to the sternum and chelicerae ventrally connected with a
thin membrane. Some members of the family are also uniquely characterized by the presence of a triseriate
calamistrum, a character state that is not found elsewhere in the order (Gray 1995). The Filistatidae are presently
divided in two well-defined subfamilies, differentiated by the structure and shape of the palpal cymbium, of the
seminal duct, and of the calamistrum. The Prithinae are characterized by the presence of a distinctively short,
reduced cymbium, a seminal duct formed by loosely arranged coils and setae with silk combing teeth on the
calamistrum. The Filistatinae are instead distinguished by having a longer, cylindrical cymbium, the seminal duct
always formed by tightly screwed coils and the calamistrum always located on the metatarsal crest.
The family has historically received little attention and most contributions are from the more recent decades.
Wang (1987a) described the genus Tricalamus Wang, 1987 from the Chinese region, Gray (1994) revised the
Australian genera Wandella Gray, 1994 and Yardiella Gray, 1994, Ramírez & Grismado (1997) revised the family
in Argentina, and Brescovit et al. (2016) described new species of Misionella Ramírez & Grismado, 1997 from
northern Brazil. Palearctic taxa have been comparatively more neglected and most species in this region are in dire
need of a revision. In fact, more than half of the species described for the Eurasian region are currently known only
from specimens of a single sex (Zonstein et al. 2013).
Two cosmopolitan genera have been traditionally recorded for Europe: Filistata Latreille, 1810 and Pritha
Lehtinen, 1967. Only recently were new species of Zaitunia Lehtinen, 1967 described and recorded for eastern
Mediterranean areas (Zonstein & Marusik 2016). The genus Pritha is comprised by 21 species (World Spider
Catalog 2016) widely distributed in the Palearctic and Indo-Malayan regions. In Europe, the genus has yet to be
recorded from the central and northern areas and presently only occurs in the temperate southern regions of the
continent (Nentwig et al. 2016). Most extant species were described more than 100 years ago during the19th
century. The majority of these descriptions are based on single specimens and lack diagnostic figures of the
genitalia. As such, this genus has been largely neglected throughout the 20th century and no systematic revision is
currently available. Other factors that may have discouraged researchers include: 1) the general shortage of male
specimens, short lived and only collectable during brief mating periods; 2) the fact that many species share a
similar habitus; 3) the overall reduced size of the spiders that makes analyzing internal female genitalia
challenging; 4) the lack of an external epigyne; 5) the difficulty of maintaining specimens in good conditions of
preservation in spirit collections, as previously noted by other authors (Brignoli 1982; Zonstein et al. 2013).
According to the World Spider Catalog (2016), in mainland Europe the genus Pritha is represented by two
species: Pritha nana (Simon, 1868) and Pritha pallida (Kulczyński, 1897). While the latter has always been
uniformly defined, P. nana has been historically misinterpreted a number of times and, as of today, lacks clear
taxonomic identity. Much of the confusion can be associated with Ledoux (1977), who redefined the species on the
basis of wrong morphological characters. This error, reiterated through time and unknowingly referenced by
several subsequent authors, has led to the present day confusion that surrounds this taxon.
As a result, extant species of the genus Pritha that occur in Italy and in the surrounding European countries are
presently poorly known and are in dire need of a revision. Describing well-defined diagnostic characters for each
while neglecting those that vary even within the same population is of utmost importance. The revision of the
morphological variants ascribed to the Pritha nana-group by Brignoli (1982) and identified as Pritha nana in both
private and museum collections in Italy, revealed the existence of two additional distinct species.
In this revisionary work we provide a putative diagnosis of the genus Pritha, re-describe P. nana and P. pallida,
describe two new species (P. parva sp. nov. and P. sagittata sp. nov.), revalidate Pritha vestita (Simon, 1873) and
Pritha debilis (Simon, 1911), and designate a lectotype for the former. All descriptions are based on a thorough
analysis of the genital morphology of both sexes. The descriptions are well-supported by figures and photographs
of all diagnostic features discussed, and provide a clear and definitive way on how to distinguish the European
species. We also provide notes on the observed ecology and distribution of the congeners for the Italian mainland.

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A CRITICAL REVISION: THE GENUS PRITHA IN ITALY
Materials and methods
More than 300 specimens were analyzed during this study. These were either hand collected by the authors or
obtained as loans from European and Italian museums. All have been deposited and preserved in 75% ethanol.
Photos of the habitus were obtained utilizing an Olympus E-M1 camera equipped with a Sigma 150mm macro +
Raynox DCR250 and an extension tube. Imagery for specimens examined using a LEICA MZ16 stereomicroscope
was obtained using a Leica IC80HD camera or an Olympus E-M1. Specimens were also examined using a LEICA
DMLB light microscope and related photography was obtained using a LEICA DC200 camera or the Olympus E-
M1. A LEICA STEREOSCAN 430i scanning electron microscope, of the University of Trieste’s Life Sciences
department, was utilized to obtain high-magnification imagery for both the pedipalps and the calamistrum. All
photographs were processed and elaborated using Helicon Focus 5.3 and Adobe Photoshop CS6.
The vulvae of females were extracted using handheld needles of varying thickness (0.2–0.6 mm) and after
having accurately removed the surrounding abdominal setae. The genitals were then cleared using either 10%
potassium hydroxide (KOH), clove oil or a house-made solution comprised by two parts chloral hydrate, one part
phenol, one part lactic acid and one part glycerol.
If not stated otherwise, the colors discussed for each species describe how they appear when the specimens are
immersed in ethanol. All measurements are reported in millimeters and were obtained under stereomicroscopic
magnification using the LEICA APPLICATION SUITE v.4.7 software. All appendages were measured along the
dorsal plane, from the median point of the anterior margin to the median point of the posterior margin. The palpal
bulbs were measured along the dorsal plane from the notch of the horse-shoe shaped cymbium to the tip of the
embolus. In the descriptions, the first measurements reported pertain to the average, standard specimen, while the
following range in parenthesis reports the observed intraspecific variability. Species descriptions are not
exclusively based on the analysis of the sole type, but also take into account the variability of morphological
features observed in the rest of the type series and in the numerous other specimens analyzed.
The following abbreviations are used throughout the text: m = male; f = female; BL = body length; CL =
carapace length; CW = carapace width; AL = abdominal length; AW = abdominal width; CIL = clypeal length;
EGW = eye group width; MOAL = median ocular area length; AME = anterior median eyes; ALE = anterior lateral
eyes; PLE = posterior lateral eyes; PME = posterior median eyes; ALS = anterior lateral spinnerets; PMS =
posterior median spinnerets; PLS = posterior lateral spinnerets; PTR = palpal tibia ratio (ratio between palpal tibia
width and length as seen prolaterally).
Types and remaining specimens analyzed in the study are currently deposited in the following institutions: CI,
private collection of Marco Isaia stored at Dept Life Sciences and Systems Biology University of Torino, Torino,
Italy; CL, private collection of Carlo Maria Legittimo, Trieste, Italy. MCSNBG, Natural History Museum “E.
Caffi", Bergamo, Italy; MNHN, Museum National d’Histoire Naturelle, Paris, France; MNHT, Museum of Natural
History of Trieste, Trieste, Italy; MSNF, Natural History Museum “La Specola”, Florence, Italy; NHMW,
Naturhistorisches Museum, Vienna, Austria; OUMNH, Hope Entomological Collections, Oxford University
Museum of Natural History, Oxford, U.K.
Taxonomy
Family Filistatidae Ausserer, 1867
Subfamily Prithinae Gray, 1995
Pritha Lehtinen, 1967
Pritha Lehtinen, 1967: 260. Type species Filistata nana Simon, 1868.
Putative diagnosis. small to medium sized spiders. Females with a lighter fishbone abdominal pattern, males with
a single dorsal spot of whitish setae. PME > AME. Spines present. Cribellum bipartite. Calamistrum triseriate,
formed by three rows of variable length. Median row always shorter and reduced. Male palpal tibia distinctly
thickened. Cymbium horseshoe-shaped. Bulb distally and gradually tapered. Embolus thin and straight or weakly

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curved. Ejaculatory ducts with tridimensional helical development. Bulb without a dorsal lamelliform paraembolic
process (present in Wandella). Post tegulum without teeth (present in Wandella and Yardiella). Embolus without
deep prolateral grooves (present in Yardiella). Female genitalia bilaterally structured with two or three pairs of
glandular tissue-covered diverticula per seminal receptacle.
Notes. Lehtinen (1967) misidentified the specimen he illustrated and the palp portrayed as P. nana belongs to
an unidentified Pritha sp. In addition, he did not explicitly state the exact origin and identity of those specimens of
P. nana he directly utilized for his description.
Pritha nana (Simon, 1868)
Figs 1, 2A, 3A, 4A, 4E, 5, 6A–B, 7A–B, 14A–B, 15A–B, 16A, 16C, 17A–E, 19, 21, 22, 23C–D, 25B, 26A–E.
Filistata nana Simon, 1868: 455. [holotype not designated in the original description; male types from Vaucluse, France, in
MNHN, apparently lost; six females syntypes from France, Italy and Algeria, in MNHN, vial Rif. AR5416-A of Jar 477
labeled as “2747 Filistata nana E.S. Gall., Ital., Alg”, examined.]
Pritha nana—Lehtinen 1967: 260 (transferred from Filistata)
Filistata (Pritha) nana—Heimer 1990: 4, figs 4–5.
Notes on type. Simon originally described a male and female from an unknown number of specimens collected in
Vaucluse (France). Vial Rif. AR5416 of Jar 477 (MNHN) labeled as “2747 Filistata nana E.S. Gall., Ital., Alg”
contained one Agroeca sp. (Liocranidae) and a total of 14 specimens of Pritha belonging to four distinct species.
No male specimen was found inside and the male syntype is probably lost. Of the remaining female specimens,
only six can be identified as P. nana. The rest include five P. p a r v a sp. nov, one P. cf. sagittata, and two
undetermined Pritha sp. (potentially P. debilis). The label includes no date of collection or deposition, three vague
localities and no indication that among its content are elements of the type series. Several decades later, Simon
(1911) reported the presence of P. nana for France, Italy and Algeria, the same three localities that happen to be
listed on the label of the vial. Thus, while still impossible to prove with utmost certainty, it is very probable and
reasonable to assume that among this mix of specimens and taxa are the female syntypes from Vaucluse. Our intent
was to typify this very problematic species. It is one of great taxonomic relevance (the type of its genus) in dire
need of taxonomic stability. A valid, clear name-bearing physical representation would ensure its correct
interpretation for future revisions of both congeneric species and the genus itself. However, since vial Rif. AR5416
(Jar 477 MNHN) likely contains at least part of the original type series, it’s impossible to consider the type material
as being definitively lost.
At the same time, however, it is simply impossible to identify which of the female specimens contained within
are the French syntypes. In 1911 Simon records the presence of P. nana for both Italy and France. Thus, if we
exclude the undetermined, lightly colored specimens most definitely of north-African origin, in no way can any of
the specimens we identified as the real P. nana be correctly and reliably assigned to either one of the two localities.
It is simply impossible to determine which females are the ones collected from Vaucluse, France and, consequently,
impossible to correctly designate a lectotype from the specimens deposited in Simon’s collection.
Finally, it is extremely important to note that, while quite distinctive and consistent with our specimens, the
bulb morphology as originally described and worded by Simon for P. nana is not diagnostic for the species. Pritha
vestita (Simon, 1873) is described by the same author as having a “pyriform bulb” that is “as long as the tibia and
gradually tapered”. The descriptions of the bulbs of these two species are easily interchangeable and neither is
diagnostic. Without taking into consideration unreliable characters such as size or ocular measurements, small male
specimens of P. nana cannot be possibly and unequivocally discerned from similar-sized males of P. vestita when
utilizing the two descriptions of Simon. Ultimately, only the direct analysis of the type series of the latter allowed
us to taxonomically discern the two species and diagnostically differentiate and redescribe their bulbs and overall
taxonomic identities.
P. nana is a type species that lacks taxonomic identity, has been continuously misidentified, has no valid and
reliable name-bearing type specimen and has an original description that is not diagnostic. Despite the species
being in clear, indisputable and extreme need of typification, neither a neotype nor a lectotype could be designated
in the present redescription. As such, the authors will reserve the right to present the matter to the International
Commission on Zoological Nomenclature, proposing both the removal of the syntype status from any specimen

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contained in vial Rif. AR5416 (Jar 477 MNHN) and the designation of a neotype in the form of a well-preserved
male from southern France (Fig. 21), recently collected by P. Oger.
Notes on misidentifications. Pritha nana has been historically misidentified a number of times and has lacked
a clear taxonomic identity since its original description. The male palp illustrated and labeled as P. nana by
Lehtinen (1967: 260, fig. 22) was a misidentification and belongs to an unidentified Pritha sp. All illustrations
presented by Ledoux (1977) in his redescription are misidentifications, some of unidentified Pritha sp. (1977: 69,
figs 3C–3D) but most portray P. parva sp. nov. (1977: 67-69, figs 1–2, 3A–3B).. Some of these drawings are
republished and again mislabeled by Trotta (2005: 164, figs 266–268), Wunderlich (2011: 173, figs 3–7) and Le
Peru (2011: 109, fig. 109). The female vulva illustrated by Brignoli (1982: 71, fig 7) was a misidentification and
corresponds to that of P. parva sp. nov. The female genitalia illustrated by Heimer & Nentwig (1991: 38, fig. 75.4)
belongs to P. p a r v a sp. nov, while the rest of the illustrations correspond to an unidentifiable Pritha sp. (1991: 38,
fig. 75.1, 75.5–75,6). The palp and vulva in Gray (1995: 84–85, figs 14, 21) represent a misidentification and
portray an unidentifiable Pritha sp. The unpublished male palp illustrated by Pesarini and presented as P. nana in
Nentwig et al. (2016) is again a misidentification. While it may be that of P. sagittata sp. nov. due to the lack of
sufficient evidence we consider it a Pritha sp. for now.
Material examined. ITALY: Abruzzo: L’Aquila, 1♂, VI/2012, P. Di Pompeo (MCSNBG); 18♂, 30/VI/2015,
P. Di Pompeo (MCSNBG); 1♀, VI/2012, P. Di Pompeo (MCSNBG); 7♀, VI/2012, P. Di Pompeo (MCSNBG); 2♂,
VI/2012, P. Di Pompeo (CL); 1♂, VI/2013, P. Di Pompeo (MNHT); 2♂, 18/VI/2013, P. Di Pompeo (MNHT); 2♂,
30/VI/2015, P. Di Pompeo (MCSNBG); 1♂, V/2013, P. Di Pompeo (MCSNBG); 2♂, V/2012, P. Di Pompeo
(MCSNBG); 4♂, VI/2012, P. Di Pompeo (MCSNBG); 16♀, V/2012, P. Di Pompeo (CL); Avezzano: 2♂, 18/VI/
2013, P. Di Pompeo (MCSNBG); 1♀, 18/VI/2013, P. Di Pompeo (MCSNBG); Magliano dei Marsi: 1♂, 18/VI/
2013, P. Di Pompeo (MCSNBG); 1♀, 18/VI/2013, P. Di Pompeo (MCSNBG); Massa d'Albe: 1♂, II/2014, P. Di
Pompeo (MCSNBG); Tagliacozzo: 2♀, 16/VI/2013, P. Di Pompeo (MCSNBG); Lazio: Corvaro suburb of
Borgorose: 3♀, VII/2013, P. Di Pompeo (MCSNBG); Acquapendente: 4♀VIII/2014, L. Colonnelli (MCSNBG);
1♀, VIII/2014, L. Colonnelli (CL); Umbria: Assisi, Valle del Tescio, Ponte Grande: 2♀, 5/VI/2013, P. Salerno
(MCSNBG); Castiglione del Lago, Polvese Island, service centre: 4♀, 18/V/2013, P. Salerno (MCSNBG); 2♀, 2/
V/2013, P. Salerno (MCSNBG); 2♀, 7/V/2013, P. Salerno (MCSNBG); 3♂, 18/V/2013, P. Salerno (MCSNBG);
1♀, 23/V/2013, P. Salerno (MCSNBG); Castiglione del Lago, Polvese Island, mill: 2♂, 23/VI/2013, P. Salerno
(MCSNBG); 3♀, 23/VI/2013, P. Salerno (MCSNBG); Castiglione del Lago, Polvese Island, stable: 6♀, 23/V/2013,
P. Salerno (MCSNBG); Città della Pieve: 4♀, V/2016, C. M. Legittimo (CL); Gualdo Tadino, old town: 1♀, 29/VI/
2013, P. Salerno (MCSNBG); Magione, train station: 1♀, 15/V/2013, P. Salerno (MCSNBG); FRANCE:
Provence-Alpes-Côte d'Azur: Marseille, Island of Tiboulen: 1♂ 8/VI/2012, Pierre Oger & Ph. Ponel, T07 (CL);
Island of Riou, Grand Congloué: 1♂ 21/V/2013, Pierre Oger & Ph. Ponel, 1917A (CL); SPAIN: 1♂, Spain, det. O.
P. Cambridge (OUMNH); UNCERTAIN LOCALITY: 6♀, several localities included in the same vial: “Gall.,
Ital., Alg.” (MNHN AR5416-A, jar 477).
Diagnosis. Males (Figs 1B, 1D, 14B, 15B) can be distinguished from the similar P. vestita by the relatively
thinner palpal tibia and by the shape of the embolus that is similarly distinctly elongated, extending to the base of
the tibia and gradually tapering towards the end, but is significantly larger in size, comparably narrower, more
tapered apically and distally straight or slightly arched upwards. Females (Figs 1A, 1C, 14A, 15A) can be
distinguished by the large vulva, formed by four diverticula, two in each seminal receptacle: the two median are
larger, rounded, distally enlarged and divergent, the two lateral are smaller and globular in shape.
Redescription of male. Measurements: BL 2.68 (2.20–3.08), CL 1.21 (1.10–1.44), CW 0.96 (0.91–1.06), AL
1.53 (1.22–1.72), AW 0.86 (0.77–0.94), CIL 0.20 (0.14–0.26), EGW 0.26 (0.25–0.3), MOAL 0.17 (0.16–0.2),
AME 0.045, ALE 0.095, PME 0.075, PLE 0.079.
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 1.52 1.14 0.96 1.34 0.74
Patella 0.44 0.40 0.40 0.44 0.35
Tibia 1.50 1.04 0.80 1.14 0.53
Metatarsus 1.30 0.98 0.91 1.16 Cymbium 0.24
Tarsus 0.87 0.59 0.49 0.60 Embolus 0.60
Total 5.63 4.15 3.56 4.68

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FIGURE 1. Pritha nana. Habitus of live specimens: female (A), male (B); habitus of preserved specimens under microscopic
light: female (C), male (D). Scale bars: 1 mm.

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FIGURE 2. Male left palps in prolateral view. A: Pritha nana. B: Pritha pallida. C: Pritha parva. sp. nov. D: Pritha sagittata.
sp. nov. Scale bars: 200 µm.

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FIGURE 3. Male left palps in retrolateral view. A: Pritha nana. B: Pritha pallida. C: Pritha parva sp. nov. D: Pritha sagittata
sp. nov. Scale bars: 200 µm.

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FIGURE 4. Male left palps in prolateral (A–D) and retrolateral view (E–H). Size comparison between the four species; A, E:
Pritha nana. B, F: Pritha pallida. C, G: Pritha parva sp. nov. D, H: Pritha sagittata sp. nov. Scale bar: 400 µm.
Cephalothorax. Carapace ovoid-subcircular, posterior margin flattened, lateral margins slightly undulated,
covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles that covers the
preocular and postocular areas respectively. Clypeus frontally rounded with slight medial indentation. ALE > PLE
>= PME > AME. PME separated by more than their diameter. Color not uniform, dark brown with more or less
lighter patches. A darker, generally triangular mark originates from the center and forwardly bi- or trifurcates
towards the ocular area. Margins of carapace dark. Faint and irregular darker radiation present. Ocular area dark.
Clypeus covered by a depigmented patch, sometimes divided in two by a longitudinal row of setae. Sternum ovoid,
centrally light brownish-yellow, darker anteriorly and along the margins, with two small posterior sigilla,
associated with lighter pigmentation and positioned near the margin, which may be more or less visible or
completely absent. Labium longer than wide, ogival, dark, with two lighter, central spots. Coloration lightens
anteriorly. Labium and sternum darker than coxae. Maxillae elongated and convergent, brownish-orange, lighter
anteriorly. Chelicerae brownish-orange.

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FIGURE 5. SEM pictures of the left male palp of Pritha nana. A: bulb in retrolateral view; B: bulb in prolateral view; C:
horseshoe–shaped cymbium in dorsal view; D: bulb in frontal view; E: palp in retrolateral view; F–H: details of the distal
portion of the bulb and embolus.

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FIGURE 6. SEM pictures of the calamistrum showing its triseriate structure and the setae with silk combing teeth. A, B: Pritha
nana. C, D: Pritha pallida. E, F: Pritha parva sp. nov. G, H: Pritha sagittata sp. nov.

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FIGURE 7. Vulvae. A, B: Pritha nana. C, D: Pritha pallida. E, F: Pritha parva sp. nov. G, H: Pritha sagittata sp. nov. Detail
of single seminal receptacles (B, D, F, H). Scale bars A, C, E, G: 100 µm; B, D, F, H: 50 µm.

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Legs. Leg femora distinctly dark, brown or black, with three lighter dorso-lateral longitudinal bands. Femoral
setae slightly longer ventrally. Remaining leg segments, including patellae, light brownish-yellow. Tibiae not
uniformly colored.
Palps. Palpal tibia and femur dark brown, patella lighter. Tibia moderately thickened. PTR < 0.42 (Fig. 2A).
Cymbium short, horseshoe-shaped and with a prolateral and dorso-distal projection that embraces the basal portion
of the bulb (Fig. 5C).
When viewed with a light microscope, the bulb appears brownish-orange, conical, proximally thickened, very
elongated and apically tapered. The elongated distal portion extends to the base of the tibia and may sometimes
appear slightly arched upwards. The ejaculatory duct, visible on cleared bulbs, is proximally helical and stretches
distally (Figs 2A, 3A).When viewed with a scanning electron microscope the proximal portion of the bulb appears
smooth and helical, and follows the coils of the internal ejaculatory duct. (Fig. 5D) Distally, the bulb is covered by
a series of longitudinal grooves and ends with a flattened, slightly screwed, narrow tip (Figs 5F, 5G, 5H).
Abdomen. Abdomen ovoid, dark brown to dark grey with a narrow, slightly elongated white dorsal mark (Fig.
1B); ventrally brown. Residues of a cream-colored fishbone setae pattern may be present posteriorly. These
correspond to a similarly-shaped mark of lighter pigmentation generally present on the underlying cuticle. Two
pairs of dorsal sigilla present but hardly visible.
Redescription of female. Measurements: BL 3.58 (2.52–4.07), CL 1.5 (1.12–1.82), CW 1.09 (0.88–1.47), AL
2.17 (1.48–2.53), AW1.58 (0.99–1.63), CIL 0.24 (0.19–0.33), EGW 0.3 (0.25–0.38), MOAL 0.19 (0.16–0.21),
AME 0.059, ALE 0.100, PME 0.090, PLE 0.095.
Cephalothorax. Carapace ovoid-subcircular, posterior margin flattened, lateral margins slightly undulated;
covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles that covers the
preocular and postocular areas respectively. Clypeus frontally rounded. ALE > PLE >= PME > AME. PME
separated by more than their diameter. Color not uniform, dark brown with more or less lighter patches. A darker,
generally triangular mark originates from the center and forwardly bi- or trifurcates towards the ocular area.
Margins of carapace dark. Faint and irregular darker radiation present. Ocular area dark. Clypeus covered by a
depigmented patch, sometimes divided in two by a longitudinal row of setae. Sternum ovoid, centrally light
brownish-yellow, darker anteriorly and along the margins, with two small posterior sigilla, associated with lighter
pigmentation and positioned near the margin, which may be more or less visible or completely absent. Labium
longer than wide, ogival, dark, with two lighter, central spots. Coloration lightens anteriorly. Labium and sternum
slightly darker than coxae. Maxillae elongated and convergent, brownish-orange, lighter anteriorly. Chelicerae
brownish-orange.
Legs and palps. Palps brownish-orange, stocky, with tarsus, tibia and patella lighter than femur. Leg femora
dark brown, with lighter longitudinal bands, one of which is ventral. Dorsal bands may appear as one. Femoral
setae distinctly longer ventrally. Remaining leg segments, including patellae, light brownish-yellow. Tibiae not
uniformly colored, mostly pigmented laterally. Calamistrum short, triseriate and formed by setae with silk combing
teeth (Figs 6A, 6B).
Abdomen. Abdomen ovoid, dark brown to dark grey with a lighter dorsal mark that extends posteriorly as a
fishbone pattern formed by 4–6 more or less distinct pairs of chevron markings, ventrally brown to light brown.
The cream-colored setae pattern corresponds to a similarly-shaped mark of lighter pigmentation generally present
on the underlying cuticle. Two pairs of dorsal sigilla present.
Vulva. Appears as a single, membranous copulatory pocket with one seminal receptacle on each side, each
divided into two distinct diverticula. The two copulatory ducts are large and wide, and terminally form a pair of
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 1.50 1.17 1.03 1.32 0.82
Patella 0.50 0.50 0.50 0.50 0.41
Tibia 1.53 1.03 0.80 1.17 0.49
Metatarsus 1.44 0.99 0.93 1.09
Tarsus 0.98 0.73 0.59 0.67 0.67
Total 5.95 4.42 3.85 4.75 2.39

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swollen and divergent median diverticula. A second pair of lateral and globular diverticula is externally connected
to each by a short accessory duct. When viewed from above and along the coronal plane, the two lateral diverticula
are clearly positioned behind the median pair (Fig. 7A). Masses of glandular tissue abundantly cover the median
diverticula, while only sparse spermathecal pores are visible on the lateral diverticula. These pores, and their
protruding tuft-shaped glandular ducts, can be clearly distinguished under high magnification (Fig. 7B).
Distribution and ecology. Most specimens were collected from the city of L’Aquila and surrounding
provinces. In these areas the webs densely cover most urban walls and buildings and are particularly common on
the ruins of the old towns. Specimens were also occasionally collected inside buildings, generally in close
proximity to doors and in window frames.
This species was encountered in several other inland localities of south-central Italy such as Gualdo Tadino
(PG), Assisi (PG), Polvese Island next to Lake Trasimeno (PG), Magione (PG), Acquapendente (VT) and
Borgorose (RI)), however no Italian specimen was ever collected from coastal areas (Fig. 28). In Italy, Pritha nana
appears to be the species that extends to the highest altitude, as the city of L’Aquila and of Tagliacozzo (AQ) are at
721 and 750 m.a.s.l. respectively. On the Italian peninsula it seems to be mostly distributed in areas with a stronger
seasonality and characterized by a harsher winter climate.
Pritha pallida (Kulczyński, 1897)
Figs 2B, 3B, 4B, 4F, 6C–D, 7C–D, 8, 9, 14C–D, 15C–D, 16D, 17F–I, 20A, 25A, 25C, 27.
Filistata pallida Kulczyński in Chyzer & Kulczyński 1897: 277, pl. 10, fig. 54 [two male syntypes from Crikvenica, Croatia.
Type material not examined, probably lost. Searched in: Finnish Museum of Natural History, Helsinki (Dr. Pedro
Cardoso), Hungarian Natural History Museum, Budapest (Dr. László Dányi), Muzeum i Instytutu Zoologii Polskiej
Akademii Nauk, Warszawa (Dr. Mierzwa-Szymkowiak), Croatian Natural History Museum, Zagreb (Martina Pavlek &
Luka Katusic). Also contacted Prof. Jan Buchar and Prof. Jerzy Prószyński.]; Kulczyński 1899: 324, pl. 6, figs 1–2;
Machado 1941: 1, fig. 1.
Pritha pallida—Lehtinen 1967: 260 (transferred from Filistata); Wunderlich 1987a: 67, fig. 47; Wunderlich 2011: 173, figs 8–
9; Le Peru 2011: 110, fig. 110.
Notes. Pritha pallida was described by Kulczyński in 1897 from two male specimens collected in Crkvenica,
Croatia. The species has retained a very clear taxonomic identity since then and has never been misidentified by
successive authors.
Material examined. ITALY: Abruzzo: Francavilla al Mare: 2♀, 29/VI/2016, P. Di Pompeo & F. Tantalo
(MNHT); Montesilvano: 2♀, 3/VII/2016, P. Di Pompeo (MNHT); Pescara: 11♀, V/2012, P. Di Pompeo
(MCSNBG); 1♂, 20/VII/2014, P. Di Pompeo (MCSNBG); 2♂, V/2013, P. Di Pompeo (CL); 3♀, V/2012, P. Di
Pompeo (MCSNBG); Apulia: Francavilla Fontana: 1♂, VI/2013, D. Cinefra (MCSNBG); 1♀, VI/2013, D. Cinefra
(MCSNBG); Lecce: 2♀, 19/IV/2014, V. Giusti (MNHT); Emilia Romagna: Bologna: 1♂, 20/V/2014, C.M.
Legittimo (MCSNBG); Lazio: Roma: 1♂, 10/4/2014, A. Kulczycki (MCSNBG); 1♂, VI/2012, A. Kulczycki
(MNHT); 5♀, 10/IV/2014, A. Kulczycki (MCSNBG); Roma, Villa Borghese: 2♀, 15/VII/2013, A. Kulczycki
(MCSNBG); Palestrina: 1♂ 3/VII/2015, P. Di Pompeo (MCSNBG); 1♀, 3/VII/2015, P. Di Pompeo (MCSNBG);
Capranica Prenestina: 1♀, 4/VII/2015, P. Di Pompeo (CL); Antrodoco: 3♀, 27/VI/2015, P. Di Pompeo & V. Giusti
(MCSNBG); 3♀, VII/2012, V. Giusti (MNHT); Civitavecchia: 1♀, 23/IX/2015, P. Di Pompeo (MCSNBG);
Liguria: Genova, Imperial Villa: 7♀, 30/V/2016, E. Simeon (CL); 2♂, 30/V/2016, E. Simeon (CL); Genova, Via
XX settembre: 1♂, 30/V/2016, E. Simeon (MNHT); Tu s c a n y : Chiusi: 1♂, 27/VI/2016, F. Fabietti (CL); 3♀, 27/
VI/2016, F. Fabietti (CL); Firenze: 1♂, V/2012, P. Di Pompeo (MCSNBG); 2♂, V/2012, P. Di Pompeo (CL); 1♂,
V/2012, P. Di Pompeo (MNHT); 11♀, V/2012, P. Di Pompeo (MCSNBG); 1♂, 22/VII/1996, Berdondini (MSNF
num.3073); 1♂, 28/VI/2001, Berdondini (MSNF num.3076); Firenze, Via Girolami: 5♀, 8/IV/2015, C. M.
Legittimo (MCSNBG); Pisa: 2♀, V/2012, P. Di Pompeo (MNHT); Radda in Chianti: 1♀, V/2012, E. Simeon
(MCSNBG); Villa di Mercatale, Vinci: 1♂, 19/06/1921, collector unknown (MSNF num.3070); Umbria: Amelia:
4♀, 29/VI/2013, A. Kulczycki (MCSNBG); 1♂, 12/VI/2015, A. Kulczycki (MNHT); Orvieto: 2♀, 12/VIII/2014 C.
M. Legittimo (MCSNBG); Perugia: Ponte San Giovanni: 1♀, 23/IV/2013, P. Salerno (MCSNBG); SPAIN:
Barcellona: 2♀, 19/V/2013, A. Kulczycki (MCSNBG); GREECE: Corfù: 1♀, date and collector unknown
(NHMW num.12368).

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FIGURE 8. Pritha pallida. Habitus of live specimens: female (A), male (B); habitus of preserved specimens under
microscopic light: female (C), male (D). Scale bars: 1 mm.

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FIGURE 9. SEM pictures of the left palp of Pritha pallida. A: bulb in retrolateral view. B: bulb in prolateral view. C:
horseshoe–shaped cymbium in dorsal view. D: detail of the dorso–distal portion of the bulb in prolateral view. E, F: details of
the distal portion of the bulb in frontal view. G, H: close up of the dorso–terminal lamellar process located on the distal portion
of the bulb(G: retrolateral view; H: posterior view).

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Diagnosis. Males (Figs 8B, 8D, 14D, 15D) can be distinguished from the similar P. sagittata by the uniform
light-yellow coloration of cephalothorax and legs and by the morphology of the dorso-terminal lamellar process of
the palp that is formed by two translucid longitudinal, differently sized and slightly overlapped lamellar structures.
Females (Figs 8A, 8C, 14C, 15C) can be distinguished from the similar P. sagittata by the uniform light-yellow
coloration of cephalothorax and legs and by the structure of the conically-developed copulatory ducts that appears
not truncated.
Redescription of male. Measurements: BL 2.70 (2.01–2.75), CL 1.20 (0.96–1.23), CW 0.81 (0.72–0.88), AL
1.54 (1.08–1.59), AW 0.92 (0.68–0.95), CIL 0.20 (0.16–0.23), EGW 0.25 (0.23–0.28), MOAL 0.17 (0.14–0.18),
AME 0.050, ALE 0.100, PME 0.070, PLE 0.080.
Cephalothorax. Carapace ovoid-subcircular, posterior margin flattened, lateral margins barely undulated;
covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles that covers the
preocular and postocular areas respectively. Clypeus frontally rounded. ALE > PLE > PME > AME. PME
separated by more than their diameter. Color uniformly light-yellow. A thin, darker mark of varying size originates
from the center and extends anteriorly to the ocular area. Ocular area dark. Clypeus centrally lacking pigmentation
and with dark lateral margins. Sternum ovoid, pale yellow and without pigmentation. Labium longer than wide,
ogival, slightly darker than the sternum. Maxillae elongated and convergent, yellow-orange, distally darker.
Chelicerae brownish-orange.
Legs. Legs with no longitudinal bands or annulations. Color uniformly yellowish-orange, slightly darker than
that of the cephalothorax. Distal leg segments sometimes darker in tone.
Palps. Palps, including cymbium, uniformly yellowish-orange. Tibia thickened. 0.42 < PTR < 0.52 (Fig. 2B).
Cymbium short, horseshoe-shaped and with a prolateral and dorso-distal projection that embraces the basal portion
of the bulb (Fig. 9C). When viewed with a light microscope, the bulb appears yellow and conical. The apex of the
embolus is dark, cylindrical and slightly oriented downwards. Distally, the bulb carries a unique dorsal process
formed by two translucid longitudinal lamellar structures, differently sized and slightly overlapped between each
other. The ejaculatory duct, visible on cleared bulbs, is proximally helical and stretches distally (Figs 2B, 3B).
When viewed with a scanning electron microscope, the proximal portion of the bulb appears smooth and
helical, and follows the coils of the internal ejaculatory duct (Fig. 9C). Distally and prolaterally, the bulb carries
several deep, longitudinal furrows that become more obvious when viewed from the front (Fig. 9F). The spermpore
is visible at the end of the cylindrical structure that contains the ejaculatory duct. The two differently sized lamellae
are inserted dorsally on the embolus: one is retrolateral and folds internally, the other, longer, is prolateral and folds
externally. The two partially overlap and form a canal (Figs 9E, 9G, 9H).
Abdomen. Abdomen ovoid, dark grey, dorsally marked only by a large patch of white setae. (Fig. 8B), ventrally
lighter brown. The remaining setae are dark. The underlying cuticle may carry lighter pigmentation in the form of a
fishbone pattern, never superficially visible on the setae of live specimens. Two pairs of dorsal sigilla present but
barely visible.
Redescription of female. Measurements: BL 3.34 (3.01–3.50), CL 1.48 (1.34–1.51), CW 1.09 (0.99–1.15),
AL 1.93 (1.76–2.17), AW 1.23 (1.18–1.42), CIL 0.25 (0.22–0.27), EGW 0.28 (0.26–0.32), MOAL 0.18 (0.17–
0.21), AME 0.063, ALE 0.095, PME 0.074, PLE 0.086.
Cephalothorax. Carapace ovoid-subcircular, posterior margin flattened, lateral margins barely undulated;
covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles that covers the
preocular and postocular areas respectively. Clypeus frontally rounded. ALE > PLE > PME > AME. PME
separated by more than their diameter. Color uniformly light-yellow. A darker, generally triangular mark of varying
size originates from the center and forwardly bi- or trifurcates towards the ocular area. Ocular area dark. Clypeus
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 1.29 0.98 0.86 1.06 0.72
Patella 0.45 0.37 0.35 0.42 0.31
Tibia 1.33 0.90 0.72 0.98 0.43
Metatarsus 1.03 0.80 0.80 1.03 Cymbium 0.16
Tarsus 0.75 0.55 0.44 0.50 Embolus 0.38
Total 4.85 3.60 3.17 3.99 2.00

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centrally lacking pigmentation and with dark lateral margins. Sternum ovoid, pale yellow and without
pigmentation, with two small, hardly visible posterior sigilla near the margin. Labium longer than wide, ogival,
slightly darker than the sternum. Maxillae elongated and convergent, yellow-orange, distally darker. Chelicerae
brownish-orange.
Legs and palps. Legs and stocky palps with no longitudinal bands or annulations. Color uniformly yellowish-
orange, slightly darker than that of the cephalothorax. Distal leg segments sometimes darker in tone. Femoral setae
slightly longer ventrally. Calamistrum short, triseriate and formed by setae with silk combing teeth (Figs 6C, 6D).
Abdomen. Abdomen ovoid, ventrally lighter brown, dorsally brown to grey with a lighter, cream-colored dorsal
mark that extends posteriorly into a fishbone pattern formed by 4–6 pairs of chevron bands. The first two pairs are
generally thicker and curved, while the remaining are gradually thinner and oblique (Fig. 8A). This cream-colored
setae pattern corresponds to a similarly-shaped mark of lighter pigmentation generally present on the underlying
cuticle. Two pairs of dorsal sigilla present and more obvious than in males.
Vulva. Appears as a single, membranous copulatory pocket with one seminal receptacle on each side, each
divided into three different diverticula. Each copulatory duct is short and conical-shaped, and terminally forms a
rounded, median diverticulum that apically carries an additional smaller, perhaps atrophic diverticulum. A third,
large, spherical diverticulum is laterally positioned and externally connected to each duct (Fig. 7C). Masses of
glandular tissue abundantly cover the median diverticula, while only sparse spermathecal pores are visible on the
lateral diverticula. These pores, and their protruding tuft-shaped glandular ducts, can be clearly distinguished under
high magnification (Fig. 7D).
Distribution and ecology. This species was collected primarily from coastal localities in close proximity to
the sea or from areas of low altitude but still governed by a strictly Mediterranean climate (Corfù (Greece),
Barcelona (Spain), Pisa (PI), Rome (RM), Pescara (PE), Montesilvano (PE), Francavilla al Mare (CH), Francavilla
Fontana (BR), Lecce (LE)). The species was not limited to coastal territories as it was also encountered inland in
areas with a favourable continental climate with milder winters. These include Bologna (BO), Firenze (FI), Radda
in Chianti (SI), Perugia (PG), Orvieto (TR), Amelia (TR) and Antrodoco (RI) (Fig. 28). It appears to be more
thermophilic compared to the other three congeneric species and is the most common in the warmer regions of the
country. In Rome and Pescara specimens were also found under the barks of trees (Pinus spp., Quercus spp.) (Fig.
27) that grow along the streets or inside public gardens and parks. No other species was ever encountered in a
similar microhabitat.
Pritha parva sp. nov.
Figs 2C, 3C, 4C, 4G, 6E–F, 7E–F, 10, 11, 14E–F, 15E–F, 18I, 20B–C.
Pritha nana Ledoux 1977: 66, figs 1–2, 3A–3B (mf misidentified); Brignoli 1982: 70, fig. 7 (f misidentified); Heimer &
Nentwig 1991: 38, fig. 75.4 (mf misidentified); Trotta 2005: 164, figs 266–267 (mf misidentified); Wunderlich 2011: 173,
figs 5–6 (mf misidentified); Le Peru 2011: 109, fig. 109 (mf misidentified).
Notes. Ledoux (1977) and Brignoli (1982) erroneously ascribed wrong morphological characters to the name
Pritha nana. This taxon has since then been universally identified by the short, conical bulbs and the strongly
thickened palpal tibia. As discussed previously, this interpretation of the species is wrong. The morphology initially
reported by Ledoux (1977) and subsequently by several other authors, has never been correctly taxonomically
defined until now. This new species, described as Pritha parva sp. nov., is currently the smallest European member
of the genus.
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 1.30 1.04 0.93 1.23 0.76
Patella 0.45 0.44 0.45 0.45 0.34
Tibia 1.46 0.92 0.77 1.10 0.47
Metatarsus 1.19 0.81 0.68 0.98
Tarsus 0.84 0.53 0.45 0.54 0.49
Total 5.24 3.74 3.28 4.30 2.06

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Type material. Holotype ♂—ITALY: Friuli Venezia Giulia: Trieste, 45°39'23.25"N, 13°48'2.37"E: 21/IV/
2015, E. Simeon, Holotype, deposited together with a red marker (MCSNBG). In a crevice of an urban wall.
Paratypes—ITALY: Friuli Venezia Giulia: Trieste: 4♂, VI/2013, E. Simeon (MNHT); 2♀, 25/V/2014, E.
Simeon (MCSNBG); 1♂, V/2012, E. Simeon (MCSNBG); 4♀, 8/VII/2014, E. Simeon (MNHT); 4♀, 8/VII/2014,
E. Simeon (CL); 4♀, 8/VII/2014, E. Simeon (MNHT); 5♀, 2013, E. Simeon (MNHT); 4♀, 25/V/2014, E. Simeon
(MNHT); 1♀, V/2013, E. Simeon (MNHT); Lazio: Rieti: 2♀, 30/VI/2015, P. Di Pompeo & Y. Vavatzianis
(MCSNBG); Lombardy: Bergamo: 2♂, 2004–2009, P. Pantini (MCSNBG); 2♀, 2004–2009, P. Pantini
(MCSNBG); Milano: 2♂, 2011, P. Di Pompeo (MCSNBG); 1♂, V/2012, A. Nardozi (CL); Piedmont: Invorio
Superiore suburb of Invorio: 1♂, 2/IX/2015, J. Casini (MCSNBG); 3♀, 2/IX/2015, J. Casini (MCSNBG);
Tuscany: Firenze: 1♀, 8/IV/2015, C. M. Legittimo (CL); 5♀, V/2013, P. Di Pompeo (CL); SWITZERLAND:
Lugano: 1♂, V/2013, G. Meier (MNHT); 2♂, V/2013, G. Meier (CL); FRANCE: Provence-Alpes-Côte d'Azur:
Avignon: 1♂, 12/IV/2016, P. Moulet (CL).
Additional material examined. ITALY: Friuli Venezia Giulia: Trieste: 1♂, 15/III/1965, Alberti (MNHT);
1♂, 8/VI/1964, Alberti (MNHT); Lombardy: Bergamo: 1♂, 29/V/2002, P. Pantini (MCSNBG); 1♂, 25/III/2002,
Valle (MCSNBG); Calvatone, Le Bine Oasis: 1♂, VI/1996, P. Pantini-Valle (MCSNBG); Tu s c a ny: Bagno a Ripoli,
Terzano loc.: 1♀, 18/III/1996, Berdondini (MSNF num.3072); 1♂, 30/V/2000, Berdondini (MSNF num.3074); 1♂,
12/VI/2000, Berdondini (MSNF num.3075); Veneto: Venezia: 1♂, date and collector unknown (NHMW num.
12366); FRANCE: Provence-Alpes-Côte d'Azur: Avignon: 5♂, 16 ♀, 9/VI/1970, Ledoux (MNHN 835, Ledoux
collection); 1♀, 10/VI/I970, Ledoux (MNHN 838, Ledoux collection); 1♀, 15/X/I970, Ledoux (MNHN 888,
Ledoux collection); Palais des Papes: 1♀, 24/II/I947, R. Riboulet (MNHN 1144, Ledoux collection); 1♂, 1/V/I977,
Ledoux (MNHN 1150, Ledoux collection); Occitanie: Les Angles: 1♀, 2/II/I977, Ledoux (MNHN 1117, Ledoux
collection); 1♂, collected 13/III/I977 and killed 18/IV/1977, Ledoux (MNHN 1145, Ledoux collection);
UNCERTAIN LOCALITY: 5 ♀, several localities reported in the same vial: “Gall., Ital., Alg.” (MNHN AR5416-
B, jar 477).
Etymology. The specific name refers to the minute size of the species and derives from the Latin word
“parvus”, meaning minute; adjective.
Diagnosis. Males (Figs 10B, 10D, 14F, 15F) can be distinguished by the very short conical-shaped palpal bulb,
the strongly thickened palpal tibia and the glossy appearance of the cephalothorax. Females (Figs 10A, 10C, 14E,
15E) can be distinguished by the very small vulva (formed by four diverticula: two median, conical-shaped and two
lateral, globose), the glossy appearance of the cephalothorax and the abdominal pattern consisting of a centrally-
placed white dorsal spot similar to the one of males.
Description of male. Measurements: BL 1.98 (1.56–2.13), CL 0.95 (0.73–1.01), CW 0.70 (0.60–0.78), AL
1.10 (0.86–1.22), AW 0.65 (0.44–0.71), CIL 0.15 (0.14–0.19), EGW 0.21 (0.16–0.22), MOAL 0.13 (0.11–0.14),
AME 0.025, ALE 0.072, PME 0.035, PLE 0.054.
Cephalothorax. Carapace ovoid-subcircular, posterior margin slightly flattened, lateral margins slightly or very
undulated, covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles that covers
the preocular and postocular areas respectively. Clypeus frontally rounded. ALE > PLE > PME > AME. PME
separated by more than their diameter. Color not uniform, dark brown with more or less lighter patches. A darker,
generally triangular and sometimes absent mark originates from the center and forwardly bi- or trifurcates towards
the ocular area. Margins of carapace dark. The cuticle has a glossy, shiny appearance. Ocular area dark. Clypeus
covered by a depigmented patch, sometimes divided in two by a longitudinal row of setae. Sternum ovoid, brown
to dark brown, darker along the margins. Posterior sigilla not visible. Labium longer than wide, ogival, dark, with
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 0.93 0.77 0.69 0.89 0.53
Patella 0.26 0.26 0.25 0.26 0.23
Tibia 1.04 0.76 0.59 0.88 0.33
Metatarsus 0.84 0.64 0.62 0.83 Cymbium 0.12
Tarsus 0.62 0.42 0.38 0.45 Embolus 0.27
Total 3.69 2.85 2.53 3.31 1.48

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two lighter, central spots. Coloration lightens anteriorly. Labium and sternum darker than coxae. Maxillae
elongated and convergent, brown, lighter anteriorly. Chelicerae brownish-orange.
Legs. Leg femora dark brown, with lighter, longitudinal, more of less fused, dorsolateral bands. Remaining leg
segments light brownish-yellow with patella lighter than the rest. Coloration of tibiae not uniform, distally lighter.
Palps. Palpal tibia and femur dark brown, patella lighter. Tibia strongly thickened. PTR > 0.52 (Fig. 2C). Cymbium
short, horseshoe-shaped and with a prolateral and dorsodistal projection that embraces the basal portion of the bulb
(Fig. 11C).
When viewed with a light microscope, the bulb appears brownish-yellow, conical and stocky. The apex of the
embolus is dark, cylindrical and slightly oriented downwards. The bulb does not carry any dorsal process. The
ejaculatory duct, visible on cleared bulbs, is proximally helical and stretches distally (Figs 2C, 3C). When viewed
with a scanning electron microscope, the proximal portion of the bulb appears smooth and helical, and follows the
coils of the internal ejaculatory duct (Figs 11B, 11D). Distally, the bulb carries several deep, dorsal and prolateral
furrows (Figs 11F, 11H). The spermpore is visible at the end of the cylindrical structure that contains the
ejaculatory duct.
Abdomen. Abdomen ovoid, ventrally dark brown, dorsally dark brown to dark grey and marked in center by a
white, oval dorsal spot (Fig. 10B). The underlying cuticle may posteriorly carry marks of lighter pigmentation in
the form of a fishbone pattern, never superficially visible on the setae of live specimens. Two pairs of dorsal sigilla
present but hardly visible.
Description of female. Measurements: BL 2.29 (2.22–2.50), CL 1.02 (0.88–1.23), CW 0.76 (0.67–0.90), AL
1.31 (1.30–1.45), AW 0.90 (0.85–1.00), CIL 0.19 (0.14–0.20), EGW 0.23 (0.20–0.26), MOAL 0.13 (0.12–0.14),
AME 0.040, ALE 0.074, PME 0.051, PLE 0.061.
Cephalothorax. Carapace ovoid-subcircular, posterior margin flattened, lateral margins slightly undulated,
covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles that covers the
preocular and postocular areas respectively. Clypeus frontally rounded. Setae on lateral margins of cephalothorax
inconspicuous. ALE > PLE > PME > AME. PME separated by more than their diameter. Color not uniform, dark
brown with more or less lighter patches. A darker, generally triangular and sometimes absent mark originates from
the center and forwardly bi- or trifurcates towards the ocular area. Margins of carapace dark. The cuticle has a
glossy, shiny appearance. Ocular area dark. Clypeus covered by a depigmented patch, sometimes divided in two by
a longitudinal row of setae. Sternum ovoid, centrally light brownish-yellow, darker anteriorly and along the
margins, with two small posterior sigilla which may be more or less visible or completely absent. Labium longer
than wide, ogival, dark with two central lighter patches. Labium and sternum generally darker than coxae. Maxillae
elongated and convergent, brownish-orange, lighter anteriorly. Chelicerae orange.
Legs and palps. Palps brownish-orange, stocky, with femora sometimes darker than the rest. Leg femora
brown, with lighter, longitudinal, more of less fused dorsolateral bands. Femoral setae slightly longer ventrally.
Remaining leg segments light brownish-yellow with patellae lighter than the rest. Coloration of tibiae not uniform,
distally lighter. Calamistrum short, triseriate and formed by setae with silk combing teeth (Figs 6E, 6F).
Abdomen. Abdomen ovoid, ventrally brown to light brown, dorsally dark brown to dark grey and centrally
marked by a white, oval dorsal spot (Fig. 10A). Residues of a cream-colored fishbone pattern are present
posteriorly. This cream-colored setae pattern corresponds to a similarly-shaped mark of lighter pigmentation
generally present on the underlying cuticle. Two pairs of dorsal sigilla present but hardly visible.
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 0.83 0.67 0.63 0.79 0.50
Patella 0.27 0.28 0.27 0.29 0.21
Tibia 0.84 0.56 0.46 0.75 0.29
Metatarsus 0.70 0.49 0.50 0.63
Tarsus 0.51 0.37 0.32 0.36 0.32
Total 3.15 2.37 2.18 2.82 1.32

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FIGURE 10. Pritha parva sp. nov. Habitus of live specimens: female (A), male (B); habitus of preserved specimens under
microscopic light: female (C), male (D). Scale bars: 1 mm.

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FIGURE 11. SEM pictures of the left male palp of Pritha parva sp. nov. A: bulb in retrolateral view; B: bulb in prolateral
view; C: horseshoe–shaped cymbium in dorsal view; D: bulb in frontal view; E: palp in prolateral view; F–H: details of the
distal portion of the bulb and embolus; F: frontal view; G: dorsal view; H: prolateral view.

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Vulva. Appears as a single, membranous copulatory pocket with one seminal receptacle on each side, each
divided into two distinct diverticula. Each copulatory duct is short and conical-shaped, and terminally forms a
median diverticulum. A second, large, spherical diverticulum is laterally positioned and externally connected to
each duct (Fig. 7E). Masses of glandular tissue abundantly cover the median diverticula, while only sparse
spermathecal pores are visible on the lateral diverticula. These pores, and their protruding tuft-shaped glandular
ducts, can be clearly distinguished under high magnification (Fig. 7F).
Distribution and ecology. It was collected mostly inland (Bergamo (BG), Milano (MI), Firenze (FI), Rieti
(RI), Invorio (NO), Lugano (Switzerland)) and occasionally along the coast (Venezia (VE), Trieste (TS)). It
appears to be mostly distributed in the north-central regions of the Peninsula and its southernmost record is from
Rieti (RI) (Fig. 28). Compared to the congeners, this species seems to prefer microhabitats with greater shade and
humidity such as urban walls covered by creeper plants or similar vegetation. It does, however, appear to be highly
adaptable as it was also collected from the hotter and sunnier setting of Rieti. In Florence, this species was
occasionally observed to very closely coexist with P. pallida on the very same walls. Where they co-occur, the two
would seem to have overlapping ecological niches.
Pritha sagittata sp. nov.
Figs 2D, 3D, 4D, 4H, 6G–H, 7G–H, 12, 13, 14G–H, 15G–H, 16B, 18A–H, 20D–F, 26F–G.
Type material. Holotype ♂—ITALY: Piedmont: Torino, 45° 4'4.42"N, 7°41'25.22"E, V/2013, T. D'Elia,
Holotype, deposited together with a red marker, (MCSNBG). In a crevice of an urban wall. Paratypes—ITALY:
Lazio: Sigillo suburb of Posta: 5♀, 8/XI/2015, P. Di Pompeo & V. Giusti (MCSNBG); Lombardy: Milano: 1♀, 6/
VIII/2014, P. P. Balice (MNHT); 1♀, XI/2014, G. Gargari (CL); 1♀, 9/V/2013, V. Cortese (MNHT); Piedmont:
Torino: 1♂, V/2013, T. D'Elia (MCSNBG); 1♂, V/2013, T. D'Elia (MCSNBG); 1♀, 27/VIII/2014, T. D'Elia
(MCSNBG); 1♀, 27/VIII/2014, T. D'Elia (MCSNBG); 2♀, VI/2013, T. D'Elia (MCSNBG); 1♂, V/2013, T. D'Elia
(MNHT); 1♀, 27/VIII/2014, T. D'Elia (CL); 1♀, V/2013, T. D'Elia (MNHT); Ghiffa: 3♀, X/2015, L. Cavigioli
(MCSNBG); 1♂, III/2016, L. Cavigioli (CL); SWITZERLAND: Lugano: 1♂, V/2013, G. Meier (MCSNBG); 1♂,
V/2013, G. Meier (CL); 1♂, 5/2013, G. Meier (MNHT); CROATIA: Crikvenica: 1♂, VII/2014, C. M. Legittimo
(MCSNBG); 3♀, VII/2014, C. M. Legittimo (MCSNBG); 2♂, VII/2014, C. M. Legittimo (MNHT); 2♀, 29/VI/
2014, C. M. Legittimo (CL).
Additional material examined: ITALY: Piedmont: Ferrere: 1♂, 8/X/2007, S. Beikes (CI); Carmagnola: 1♀,
18/I/1993, G. B. Delmastro (CI); CROATIA: Krk Island, Prniba Point: 1♂, 11/VII/1974, Alberti (MNHT);
Dalmazia: 6♀, date and collector unknown (NHMW num. 12367).
Etymology. The specific name refers to the arrow-shaped mark present on the abdomen of females and derives
from the Latin word “sagitta”, meaning arrow; adjective.
Diagnosis. Males (Figs 12B, 12D, 14H, 15H) can be distinguished from the similar P. pallida by the dark
coloration of cephalothorax and leg femora, and by the morphology of the dorso-terminal lamellar process of the
palp that is formed by just a single translucid lamellar structure. Females (Figs 12A, 12C, 14G, 15G) can be
distinguished from the similar P. pallida by the dark coloration of cephalothorax and leg femora and by the
structure of the conically-developed copulatory ducts that appears truncated and trapezoid in shape.
Description of male. Measurements: BL 2.21 (2.17–2.53), CL 0.99 (0.91–1.19), CW 0.81 (0.69–0.92), AL
1.133 (1.29–1.45), AW (0.73–0.87), CIL 0.17 (0.15–0.25), EGW 0.21 (0.18–0.23), MOAL 0.14 (0.10–0.14), AME
0.030, ALE 0.075, PME 0.050, PLE 0.072.
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 0.92 0.76 0.66 0.82 0.52
Patella 0.30 0.27 0.27 0.29 0.24
Tibia 0.98 0.70 0.59 0.79 0.37
Metatarsus 0.73 0.59 0.61 0.74 Cymbium 0.11
Tarsus 0.62 0.45 0.39 0.43 Embolus 0.29
Total 3.55 2.77 2.52 3.07 1.53

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FIGURE 12. Pritha sagittata sp. nov. Habitus of live specimens: female (A), male (B); habitus of preserved specimens under
microscopic light: female (C), male (D). Scale bars: 1 mm.

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FIGURE 13. SEM pictures of the left male palp of Pritha sagittata sp. nov. Bulb in retrolateral (A) and prolateral (B) view; C:
horseshoe–shaped cymbium in frontal view; D–H: details of the distal portion of the bulb, embolus and dorso–terminal lamellar
process; F: prolateral view; G: frontal view of structure showing the presence of obstructing debris; H: frontal view of a broken
palp with no debris and an unobstructed cavity.

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FIGURE 14. Illustrated male palps in retrolateral view (right) and female genitalia (left) of each species. A, B: Pritha nana. C,
D: Pritha pallida. E, F: Pritha parva sp. nov. G, H: Pritha sagittata sp. nov.
Cephalothorax. Carapace ovoid-subcircular, posterior margin slightly flattened, lateral margins sometimes
slightly undulated; covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles
that covers the preocular and postocular areas respectively. Clypeus frontally rounded. Setae on lateral margins of
cephalothorax inconspicuous. ALE >= PLE > PME > AME. PME separated by more than their diameter. Color not
uniform, dark brown with more or less lighter patches. A darker, generally triangular and sometimes absent mark

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originates from the center and forwardly bi- or trifurcates towards the ocular area. Margins of carapace dark. Faint
and irregular darker radiation present. Ocular area dark. Clypeus covered by a depigmented patch, sometimes
divided in two by a longitudinal row of setae. Sternum ovoid, brown, darker along the margins, with two small
posterior sigilla associated with lighter pigmentation and positioned near the margin, which may be more or less
visible or completely absent. Labium longer than wide, ogival, dark, with two lighter central spots. Coloration
generally lightens anteriorly. Labium and sternum visibly darker than coxae. Maxillae elongated and convergent,
brown, lighter anteriorly. Chelicerae brownish-orange.
Legs. Leg femora distinctly dark, brown, with lighter, longitudinal, more or less fused dorsolateral bands.
Femoral setae slightly longer ventrally. Remaining leg segments uniformly light brownish-yellow and clearly
lighter than the femora.
Palps. Palpal femur dark brown, tibia dorsally lighter and patella lighter than the rest. Tibia thickened. 0.42 <
PTR <0.52 (Fig. 2D). Cymbium short, horseshoe-shaped, and with a prolateral and dorsodistal projection that
embraces the basal portion of the bulb (Fig. 13C).
When viewed with a light microscope, the bulb appears brownish-yellow, conical and slightly inclined. The
apex of the embolus is dark, cylindrical and slightly oriented downwards. Distally, the bulb carries a unique dorsal
process similar to a translucid lamellar structure. The ejaculatory duct, visible on cleared bulbs, is proximally
helical and stretches distally (Figs 2D, 3D). When viewed with a scanning electron microscope, the proximal
portion of the bulb appears smooth and helical, and follows the coils of the internal ejaculatory duct. Distally, the
bulb is traversed by numerous, irregular furrows that gradually merge and take part in the formation of the dorsal
process. The latter, when viewed from the front (Fig. 13G), is diagonally oriented and circumscribes a cavity that
overlies the terminal section of the ejaculatory duct. Debris often accumulates within this cavity (Figs 13D, 13G).
The spermpore is visible at the end of the cylindrical, slightly screwed structure that is traversed by a canal and that
contains the ejaculatory duct.
Abdomen. Abdomen ovoid, dark brown to dark grey and centrally marked by an extended, sometimes
anteriorly pointed mark of white setae (Fig. 12B); ventrally dark. Additional fishbone markings superficially not
present. Two pairs of dorsal sigilla present but hardly visible.
Description of female. Measurements: BL 3.35 (2.53–3.65), CL 1.45 (1.05–1.50), CW 1.12 (0.82–1.14), AL
1.99 (1.64–2.30), AW 1.26 (1.08–1.70), CIL 0.26 (0.19–0.28), EGW 0.27 (0.22–0.30), MOAL 0.17 (0.15–0.18),
AME 0.040, ALE 0.085, PME 0.056, PLE 0.080.
Cephalothorax. Carapace ovoid-subcircular, posterior margin slightly flattened, lateral margins sometimes
slightly undulated; covered by sparse grey setae and with a strong tuft of forward and posterior-oriented bristles
that covers the preocular and postocular areas respectively. Clypeus frontally rounded. Setae on lateral margins of
cephalothorax inconspicuous. ALE >= PLE > PME > AME. PME separated by more than their diameter. Color not
uniform, varies from light to dark brown and with more or less lighter patches. Additional patterns or markings
absent. Margins of carapace dark. Ocular area dark. Clypeus covered by a depigmented patch, sometimes divided
in two by a longitudinal row of setae. Sternum ovoid, light to dark brown, sometimes darker along the margins.
Darker, lateral elongated marks that diagonally radiate from the central axis can be visible. Two small sigilla,
associated with lighter pigmentation and posteriorly positioned near the margin, generally clearly visible. Labium
longer than wide, ogival, variable in coloration but concordant with sternum. Sometimes with two lighter central
spots. Labium, maxillae and sternum darker than coxae. Maxillae elongated and convergent, comparable in
coloration with sternum and labium. Chelicerae brownish-orange.
Legs and palps. Palps brownish-orange, stocky, with dark femora dorsally traversed by one or more lighter
bands. Patella, tibia and tarsi lighter. Leg femora brown, with lighter, longitudinal, more or less fused dorsolateral
Leg 1 Leg 2 Leg 3 Leg 4 Palp
Femur 1.20 0.99 0.89 1.16 0.72
Patella 0.45 0.43 0.43 0.44 0.39
Tibia 1.21 0.85 0.71 1.08 0.47
Metatarsus 0.96 0.76 0.73 0.95
Tarsus 0.79 0.56 0.48 0.55 0.49
Total 4.61 3.59 3.24 4.18 2.07

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bands. Femoral setae distinctly longer ventrally. Remaining leg segments uniformly light brownish-orange,
distinctly lighter than the femora. Calamistrum short, triseriate and formed by setae with silk combing teeth (Figs
6G, 6H).
Abdomen. Abdomen ovoid, ventrally brown to light brown; dorsally dark brown to dark grey and marked by a
wide, arrow-shaped dorsal, whitish mark (Fig. 12A) that extends posteriorly into a series of chevron bands of
decreasing size and color intensity. Two pairs of dorsal sigilla present, but sometimes hardly visible.
FIGURE 15. Illustrated male (B, D, F, H) and female (A, C, E, G) habitus of each species. A, B: Pritha nana. C, D: Pritha
pallida. E, F: Pritha parva sp. nov. G, H : Pritha sagittata sp. nov.

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FIGURE 16. SEM pictures showing, in detail, the setae of the calamistrum with their silk combing teeth (A, B). A: Pritha
nana. B: Pritha sagittata sp. nov. Structure of female cribellum (C, D); C: Pritha nana. D: Pritha pallida. Scale bars C, D: 100
µm.
Vulva. Appears as a single, membranous copulatory pocket with one seminal receptacle on each side, each
divided into three different diverticula. Each copulatory duct is short, wide, trapezoid-shaped and terminally forms
a rounded, distally flattened, median diverticulum that apically carries an additional, smaller, perhaps atrophic
diverticulum. A third, oval diverticulum is laterally positioned and externally and apically connected to each duct
(Fig. 7G). Masses of glandular tissue cover the median diverticula and, in lower quantities, the lateral and apical
pairs. On the accessory diverticula, these are present in greater amounts when compared to the other congeneric
species. These spermathecal pores, and their protruding tuft-shaped glandular ducts, can be clearly distinguished
under high magnification (Fig. 7H).
Distribution and ecology. Specimens of P. sagittata sp. nov. were primarily collected from internal areas of
northern Italy (Torino (TO), Ferrere (AT), Milano (MI)) and Lugano (Switzerland). Notable exceptions include the
south-central village of Sigillo near Posta (RI) and some coastal areas of Croatia where the climate is essentially
Mediterranean (Fig. 28). In fact, this species was abundantly collected in the coastal town of Crikvenica (in a
relatively more humid and wooded area close to the sea) and from the island of Krk.

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FIGURE 17. Pritha nana (A–E) and Pritha pallida (F–I). Male (C, D, E, I) and female (A, B, F–H) intraspecific habitus
variation. Scale bars: 1 mm.

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FIGURE 18. Pritha sagittata sp. nov. (A–H) and Pritha parva sp. nov. (I). Male (F–H) and female (A–E, I) intraspecific
habitus variation. Scale bars: 1 mm.

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FIGURE 19. Morphological intraspecific variability of the vulvae of Pritha nana. A: unfertilized with atrophied lateral
diverticula; B, C: fully developed filled with sperm cells; D: unfertilized fully developed with collapsed ejaculatory ducts; E:
fertilized with spillage of sperm cells; F: detail of the membranous areolae that surround the receptacles; G: asymmetrical
fertilization; H: close up of a seminal receptacle. Scale bars: 100 µm.

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FIGURE 20. Morphological intraspecific variability of the vulvae. A: vulva of Pritha pallida; B, C: vulvae of Pritha parva sp.
nov.; D–F: vulvae of Pritha sagittata sp. nov. Scale bars A, B, D, E: 100 µm; C, F: 50 µm.
Pritha vestita (Simon, 1873) spec. reval.
Figs 23A–F.
Filistata vestita Simon 1873: 36; [holotype not designated in the original description; male and female syntype in Vial. Rif.
AR5422 of Jar 477 (MNHN), labeled as “2746 Filistata vestita E.S. Corsica!” (Fig. 23A), examined]; Simon 1892: 254,
fig. 211; Simon 1914: 72.
Pritha vestita—Lehtinen 1967: 260 (transferred from Filistata)
Pritha nana Ledoux 1977: 66, figs 1–3 (mf misidentified) [incorrect synonymy]

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FIGURE 21. Left palp in prolateral (A) and retrolateral view (B) of a specimen of Pritha nana from the Island of Riou
(Marseille, France). Scale bar: 200um. Photo by Pierre Oger.
FIGURE 22. Pritha nana male. A: label of the specimen preserved in O.P.-Cambridge’s collection at OUMNH. B: habitus. C:
male palps.
Notes. This species was originally described by Simon (1873) from a male collected from the island of Corsica. It
was then incorrectly synonymized with an erroneous interpretation of Pritha nana by Ledoux (1977) who
considered differences in size, shape of sternum, color and spination forms of intraspecific variability. Analysis of
the type material (one male and one female) contained within vial Rif. AR5422 of Jar 477 (MNHN) and, more
specifically, of their genitalic and morphological characters clearly confirmed the taxonomic validity of P. vestita.
The taxon is here revalidated to species status and the synonymy proposed by Ledoux (1977) is rejected. The well-
preserved male syntype is here designated as the lectotype of the species and deposited in vial Rif. AR5422-A. The
female paralectotype (vial Rif. AR5422-B) could not be reliably diagnosed due to its mediocre state of
preservation.
Type material: Lectotype ♂—FRANCE: Corsica, Omessa: date and collector unknown (MNHN AR5422-A,
jar 477). Paralectotype ♀—FRANCE: Corsica, Omessa: date and collector unknown (MNHN AR5422-B, jar
477).
Diagnosis. Males (Figs 22B–F) can be distinguished by the similar P. nana by the relatively thicker palpal tibia
and by the shape of the embolus that is similarly distinctly elongated, extending to the base of the tibia and
gradually tapering towards the end, but is significantly smaller in size, comparably stockier, less tapered apically
and distally arched downwards.

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FIGURE 23. Pritha vestita. A: label of the type series preserved at MNHN. B: dorsal view of the male. C: size comparison
between males of Pritha vestita and Pritha nana. D: comparison between the left male palp of Pritha vestita (left) and Pritha
nana (right). E, F: close-up of the left male palp of the lectotype of Pritha vestita. Scale bars: 100um.
Pritha debilis (Simon, 1911) spec. reval.
Figs 24A–F.
Filistata debilis Simon 1911: 301 [holotype not designated in the original description; the male appears to have been lost and
could not be examined. Vial Rif. AR5427 of Jar 477 (MNHN) contains 2 female specimens labeled as “12800 Filistata
debilis E.S. TYPE—Saïda, B. Saada Ain O’Grab” (Fig. 24A), examined].
Pritha debilis—Lehtinen 1967: 260 (transferred from Filistata)
Pritha nana Ledoux 1977: 66, figs 1–3 (mf misidentified) [incorrect synonymy ]
Notes. This species was originally described by Simon (1911) from specimens collected from three Algerian

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localities (Saïda, Bou-Saada and Ain O’Grab, just to the south of Bou-Saada). It was then incorrectly synonymized
with an erroneous interpretation of Pritha nana by Ledoux (1977) who considered differences in size, shape of
sternum, color and spination forms of intraspecific variability. Analysis of the type material (the male appears to
have been lost) contained within vial Rif. AR5427 of Jar 477 (MNHN) and, more specifically, of the color, general
pattern and morphology clearly confirmed the taxonomic validity of P. debilis. The taxon is here revalidated to
species status and the synonymy proposed by Ledoux (1977) is rejected. Due to the poor state of preservation of
the syntypes, no lectotype has been designated at the moment.
Type material. ALGERIA: Saïda, B. Saada Ain O’Grab: 2♀, date and collector unknown (MNHN
AR5427, jar 477).
Diagnosis. Females (Figs 23B–F) can be distinguished from the similarly colored P. pallida by the distinctly
lighter margins of the carapace and by the conspicuous dark annulations of the legs.
FIGURE 24. Filistata debilis. A: label of the type series preserved at MNHN. B, C, D, E: dorsal and lateral view of a female
specimen showing the conspicuous pigmented ring-shaped markings of the legs. F: close-up of the carapace showing the
distinctly lighter margins.
FIGURE 25. A: drawing of a palp of Pritha pallida from Machado (1941). B: label of the vial containing Filistata nana from
Simon’s collection, preserved at MNHN, with the three distinct localities. C: Pritha pallida male (left) and female (right)
drawings of the prosoma showing the cephalic pattern, from Kulczyński (1899).

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FIGURE 26. A: Pritha nana mating (in captivity); B: Pritha nana with egg sac (in captivity). C–E: typical cribellate webs of
Pritha nana built within the cracks and crevices of urban walls; F, G: webs of Pritha sagittata sp. nov. covered by copious
amounts of debris, particulate and dust. H: typical web of Filistata insidiatrix.

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FIGURE 27. A: Quercus sp. photographed near urban structures and home to a population of Pritha pallida; B–D: close ups of
the web-covered trunk showing specimens of Pritha pallida feeding and cribellate webs abundantly covered by prey remains
and organic debris.
Discussion
Systematics
The genus Pritha Lehtinen, 1967 and its affinities. The genus Pritha was rather inaccurately defined by Lehtinen
in 1967. The author designates P. nana (Lehtinen 1967, p. 260) as the type species of the group, but does not
specify whether the specimens he observed were from the type series, or not. The genus as presently defined is
widely distributed from the Palearctic to the Indo-Malayan regions. The list of diagnostic characters defined by
Lehtinen for the genus Pritha has been considered valid by most subsequent authors (Brignoli 1982). However,
while most do find confirmation in our review, some characters are not diagnostic and need to be revaluated,
whereas others were probably superficially and inaccurately defined at the time and are here considered invalid.
One of the more glaringly erroneous characters, commonly utilized throughout the years even to define new
genera, is the structure of the calamistrum. Our thorough SEM-based analyses, carried out on female specimens of
each species including the type species Pritha nana, revealed that the calamistrum is always triseriate and not
biseriate. Retrolaterally placed on the dorso-proximal area of metatarsi IV, it is always formed by 3 rows of toothed
setae (Fig. 6), variable in length but with the middle row always shorter and reduced compared to the lateral two.
The specific number of toothed setae in each row could not be accurately measured as the setae could be either
tightly overlapped and indistinguishable or missing as probably detached from the specimens. The typical silk
combing teeth carried by the setae of the calamistrum of all Prithinae, were easily identified under high

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magnification in all species (Figs 16A, 16B). Similar observations on the correct anatomy of the calamistrum of
Pritha have been recently carried out by both Ramirez (2014: fig. 51E) and by Marusik & Zamani (2015) who
observed the triseriate structure in Pritha garfieldi. However, it was Gray (1995) who initially noticed the
occurrence of a triseriate calamistrum in both Pritha and other Prithinae. He consequently discouraged authors
from heavily relying on its structure to distinguish the various genera of Filistatidae. Gray also suggested that the
taxonomic status of the Chinese genus Tricalamus (Wang 1987a), which has overlapping distribution with very
similar Pritha species but was defined by having the supposedly unique triseriate condition of the calamistrum,
may be unclear and should be investigated.
The genus Tricalamus was described by Wang on the basis of two diagnostic characters: a triseriate
calamistrum and a square-shaped cribellum. We believe the author may have both heavily relied on the inaccurate
diagnosis of Lehtinen (1967) and may have misinterpreted the triseriate structure of the calamistrum of Pritha
ampulla Wang, 1987b and Pritha spinula Wang, 1987b (who he described that same year) as being anatomically
biseriate. Additionally, Wang did not investigate the morphological characters present on the type specimens of
European species of Pritha.
Our analyses and observations also revealed that the cribellum of Pritha species is indeed always bipartite
(Figs 16C, 16D) but also variable in shape and hardly defined by a specific geometric figure. The triangular shape
ascribed by Lehtinen to the cribellum of Pritha should not be regarded as a relevant character for the genera, and
this was also noted by Ledoux (1977). Similarly, figures provided by Wang that portray the supposedly square-
shaped cribellum of Tricalamus only vaguely recall an actual square and appear to be extremely variable between
the various congeneric species.
In conclusion, the structure of the cribellum and, more importantly, of the calamistrum cannot be considered
valid characters anymore and cannot be utilized to define and distinguish the genus Tricalamus. The recent
redescription of the type species of the genus (Zonstein et al. 2016) did not identify any significant morphological
difference that supports the need to assign T. tetragonius and Pritha nana to two different genera. Furthermore, a
triseriate calamistrum has been recently described also for the genus Wandella and Andohorano Lehtinen, 1967
(Magalhaes 2016: fig. 5E; Zonstein & Marusik 2015: figs 5, 14). The amount of similarities present between
species ascribed to the genus Tricalamus and those known for the genus Pritha is substantial and due to the lack of
unique morphological characters, the former cannot be taxonomically discerned from the latter. The authors here
fully support the idea proposed by Gray in 1995 and, pending a phylogenetic assessment of the group, suggest that
the genus Tricalamus may be a junior synonym of Pritha.
Lehtinen also erroneously ascribed to the genus Pritha the lack of a pattern on the carapace. Both sexes of
several species we analyzed carried distinct markings on the cephalic area. These may be less conspicuous in live
specimens due the color of the tegument and the presence of setae, but become distinctly visible and hard to
overlook in specimens immersed in ethanol (Figs 1, 8, 10, 12). In addition, the ejaculatory duct of this genus is not
diagnostically U-shaped as defined by Lehtinen, but distinctly helical and tridimensional in development (Figs 2,
3) as it progresses inside the bulb. This was similarly noted also by Ledoux (1977).
Thus, out of the characters defined by Lehtinen for Pritha, only those related to the structure of the male palp
are here considered as being presently valid. All species analyzed in the present study present a horse-shoe shaped
cymbium (Figs 5, 9, 11, 13), a distally and gradually tapering palpal bulb and a thin, straight to weakly curved
embolus. The palpal tibia is distinctly thick when compared to that of most other Filistatidae. None of the Pritha
species analyzed in this work appear to carry either a paraembolic process on the embolus, found in Wandella and
Yardiella, or large to minute comb-like teeth arrays on the post-tegular palpal surface, found in Wandella, Yardiella
and Pikelinia (Gray 1995).
Other characters described by Lehtinen that found confirmation in our study include: an indistinct fovea, an
abdominal pattern consisting generally of a lighter fishbone pattern in females and a single, whitish variably-sized
and shaped dorsal spot in males, leg femora and tarsi without spines (though Marusik & Zamani (2015) did identify
the presence of weak femoral spines in Pritha garfieldi) and bilaterally-structured female genitalia formed by
either 2 or 3 pairs of diverticula per seminal receptacle.
Even though all the aforementioned characters can be safely ascribed to species of Pritha, there is no particular
character state that can be defined as unique for this genus. All characters listed by Lehtinen that found
confirmation in our study are not exclusive for genus and none can be reliably utilized to unmistakably identify any
given Prithinae as a Pritha species. The genus, thus, is presently poorly taxonomically defined and, pending a

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taxonomical and phylogenetic analysis of the subfamily, can only be putatively diagnosed at the present time. Due
to the lack of any unique character state, this genus should presently not be considered monophyletic. Despite
acknowledging its poor taxonomic status, we assigned both new species described to the genus Pritha since it
presently is the only Prithinae recorded for both the European continent and the Mediterranean basin.
Character variability and species diagnosis. Several authors (Ledoux 1977; Brignoli 1982) observed
significant intraspecific variability within the genus Pritha and, concordantly, some morphological characters were
found to vary substantially between conspecific individuals. The size of males and females varied significantly,
especially in the latter which continue to molt and marginally grow even after reaching sexual maturity. Size
variability will often directly affect several other characters including leg thickness, clypeal length, distance
between eyes and the general appearance of the vulva. Larger and older female specimens were found to carry
longer and thinner leg tarsi compared to younger, conspecific females.
A remarkable variability was observed in the size and developmental stage of the vulva. Female genitalia
appeared inconspicuous and reduced in very young females, but fully developed and functional in adult specimens.
The vulvae varies substantially also between fertilized and unfertilized adult females. The ejaculatory ducts are
well expanded in females that have recently mated (Figs 19B, 19C) but are visibly reduced and collapsed when
sperm cells are absent (Figs 19A, 19D, 19G, 19H). This phenomenon was observed and reported also by Ledoux
(1977, p. 68, fig. 2A, 2B). Adult females with well-developed genitalia were sometimes found to have one or more
reduced diverticulum, apparently atrophic with no glandular tissue and barely visible without the use of transmitted
light illumination (Fig. 19A).
The pattern and coloration of both sexes was found to slightly vary depending on both the age of the specimen
and its population of origin (Figs 17, 18). Female abdominal patterns vary in color intensity and shape, and while
generally recognizable can be inconsistent between conspecific specimens. The same holds true for the dorsal spot
of conspecific males which can vary in size and shape and carry indentations along the margins. Similarly, the
color intensity and visibility of the cephalic markings in both sexes also varies and the darker prosomal radiation
common in most species may sometimes be indistinguishable from the background color of the carapace. While
they certainly are useful elements of comparison when used in combination with other characters, pattern and
coloration alone should not be reliably utilized to identify the different species of Pritha.
Due to the degree of intraspecific variability of numerous morphological characters, the Pritha species here
discussed can be reliably identified only by using the shape of the bulb and the relative thickness of the palpal tibia
in males and the general structure and conformation of the vulva in females.
In Italy Pritha nana is easily identifiable by the distinctly elongated and gradually tapering palpal bulb which
is significantly longer than that of the other species. Distally, the bulb is devoid of any sort of terminal process, and
its overall length is comparable to that of the palpal tibia (Fig. 2A). The thin, terminal portion of the embolus is
fragile and can sometimes break off, giving the impression of a shorter bulb. Pritha parva sp. nov. can be
distinguished by the short, compact and conical-shaped bulb that terminates with a brief, cylindrical, slightly
downward-inclined embolus (Fig. 2C). The palps of both Pritha pallida and P. sagittata sp. nov. distally carry a
specific terminal process that makes their distinction relatively rapid and straightforward (Figs 2B, 2D). These two
species generally share several similar characters and are probably closely related. Besides the shape of the bulb,
the relative thickness of the palpal tibia can also be utilized to distinguish single or groups of species. P. nana
carries the longest and least thickened tibia as opposed to P. parva sp. nov., which has a distinctly shorter and
thicker segment. P. pallida and P. sagittata share an intermediate state of the character, and carry a similarly sized
and shaped palpal tibia.
The general structure of the vulva can also be utilized to single out each species with exception of P. pallida
and P. sagittata sp. nov. These two share a very similar morphology as both have an additional, third diverticula
that is apically connected to the median pair. (Figs 6C, 6D, 6G, 6H) The vulvae of P. nana and P. pa rv a sp. nov.
have different structure and cannot be confused. The two genitalia also differ in size, with the former being much
larger than the latter. Both share the presence of two pairs of diverticula per seminal receptacle, but the median pair
is wide, distinctly flattened and terminally rounded in P. nana (Figs. 6A, 6B) while it appears apically pointed in P.
parva sp. nov. (Figs 6E, 6F). While the vulvae of P. pallida and P. sagittata sp. nov. share a very similar
morphology, they differ by the shape of the copulatory ducts: these appear conically-developed in the former and
conically-developed but truncated and trapezoid in shape in the latter.

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Pritha nana (Simon, 1868)
In 1868 Eugène Simon first describes Filistata nana (Simon, 1868) from an unspecified number of French
specimens collected in Vaucluse, Provence (near the town of Avignon). This rather simple description includes a
fairly superficial analysis of an adult male, an even shorter analysis of a female and no figures. Among the
characters listed in said description, the following should be considered most relevant: “Male: 3,5mm, a
longitudinal mark of white setae present on the abdomen, posteriorly marked by small white transversal and
centrally-raised lines similar to accent marks, pedipalps very long, femur black, bulb (crochet genital) red and
cylindrical that terminates with an elongated point that extends up to the base of the tibia/patella (et terminé par
une longue pointe, qui s'étend jusqu'à la base de la jambe, sous laquelle ce crochet est replié), under which the
bulb is folded. Female: legs long and thin as in males, tawny and light, with femora more or less terminally
darkened, palps thicker, cephalothorax and abdomen as in males.” (Simon 1868).
From the Muséum national d'Histoire naturelle of Paris, we successfully obtained and analyzed the only vial of
Simon’s collection that contained specimens identified as present-day Pritha nana. In this vial, Rif. AR5416 (Jar
477 MNHN), we found a total of 15 different specimens labeled as “2747 Filistata nana E.S. Gall., Ital., Alg”.
Preliminary analysis revealed the presence of a liocranid (Agroeca sp.) and more importantly the complete absence
of males. An in-depth analysis of external characters and genitalia of the remaining 14 female specimens resulted
in the following breakdown and were divided accordingly: six females of Pritha nana Rif. AR5416-A, five females
of Pritha parva sp. nov. Rif. AR5416-B, one poorly preserved female probably classifiable as Pritha sagittata sp.
nov. Rif. AR5416-C, and two unidentified Pritha sp Rif. AR5416-D.
The description of the female is extremely vague and easily associable to each specimen found within the vial.
The original male used by Simon to describe Filistata nana is absent from the aforementioned vial and is probably
lost. The description that Simon provides for the male is poorly detailed and largely unhelpful. In his early years,
Simon utilized a vague terminology to discern the different leg segments. At the time of the description of F. nana
he utilized the following: “hanche” for the coxa, “trochanter” for the trochanter, “cuisse” for the femur, “jambe”
for the patella and “tarse” for collectively referring to tibia, metatarsus and tarsus (Simon 1864, p.11). When Simon
describes the bulb of F. nana as extending “jusqu'à la base de la jambe” we can safely assume he is describing a
distinctly long bulb that extends at least to the base of tibia.
The further analysis of a well-preserved adult male from Spain labeled as Filistata nana (Fig. 22), and part of
O.P.-Cambridge’s collection (now deposited in the Hope Entomological Collection, Oxford) provided additional
support to the correct interpretation of Simon’s original morphology. This male was found to be completely similar
to those collected and thoroughly analyzed in the present study. Since O.P.-Cambridge was in the process of
describing several similar Filistatidae during the same time period (Pritha albimaculata (O. P.-Cambridge, 1872),
Pritha condita (O. P.-Cambridge, 1873)), it is quite reasonable to assume that the two authors may have consulted
and exchanged information particularly in regards to how correctly and reliably identify the species that Simon had
just recently described.
In 1977, Ledoux redescribes Pritha nana by comparing some French specimens collected in Avignon and Les
Angles with those contained in Simon’s vial at the MNHN. In doing so, he severely misjudges the taxonomic
identity of this species, interprets interspecific differences as “significant individual variations” and ultimately
redescribes wrong morphological characters. Ledoux inexplicably considered all the specimens contained in the
vial of Simon as a single, highly variable species and concurrently also neglected the presence of the Liocranidae.
It must be noted that, at the time of his redescription, the only existing figure that was supposed to depict the male
palp of Pritha nana was the one published by Lehtinen in 1967 (p.432, fig.22). A quick glance at the image reveals
that, once again, a wrong morphological character was attributed to the name Pritha nana. Lehtinen does not
portray a distinctly elongated bulb that extends up to the base of the tibia/patella as described by Simon, but a
shorter, conical bulb that distally carries an apical embolic process very reminiscent of that of Pritha pallida. The
resulting redescription of P. nana by Ledoux is ultimately mostly comprised by a collection of observations and of
generic characters that can be either observed in more than one species or that represent forms of interspecific
variation.
In regards to females, the author reports the following notable characters: “size from 1.77 mm to 4.42 mm,
shiny cephalothorax on live specimens, a white dorsal mark with length/width ratio of 1.5 to 4 and of variable
conspicuity, sometimes poorly marked, sometimes very similar to the one present on males”. The glossy appearance
of the cephalothorax is actually unique to the sole Pritha parva sp. nov. and the male-typical white dorsal spot is

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found only in the females of this species. The vulvae he graphically portrays in the redescription (Ledoux 1977, p.
68, figs 2A, 2B) again belong to P. parva sp. nov. and not P. nana. The shape of the male palp is only superficially
described as thin and with a duct formed by two internal coils. The author also observes a great variability in the
thickness of the palpal tibiae, but considers this to be intraspecific in nature. Finally, he provides several figures of
differently shaped palps as a proof for the extreme intraspecific variability he reports for this species. Some of
them, however, clearly portray the palp of P. parva sp. nov., but not of P. nana (see Ledoux 1977, Figs 3A, 3B).
The private collection of Ledoux was obtained and analyzed from MNHN where it is currently deposited. As
expected, most specimens collected from Avignon and Les Angles proved to be P. parva sp. nov. and not P. nana.
We also requested Dr. Moulet of the Musem d'Histoire Naturelle d'Avignon to kindly collect specimens of Pritha
from the same areas cited by Ledoux in his work (Ledoux 1977). While most were found to be immature and not
reliably identifiable, a mature male of P. p a r v a sp. nov. was indeed collected from a wall in a suburban area of the
town.
After the erroneous redescription of Pritha nana, Ledoux investigates the taxonomic validity of two other
similar Mediterranean species: Pritha vestita (Simon, 1873) and Pritha debilis (Simon, 1911). After having
analyzed the original descriptions of Simon (1911, 1914), he concludes that the two differ from P. nana only by
size, shape of the sternum, color and pattern of spination. Since Ledoux has considered all these characters as
widely variable even within the same species, he proposed the synonymization of these two with P. nana.
In 1982, the systematics of the Western Palearctic Filistatidae is investigated by P. M. Brignoli (1982). The
author acknowledges the presence of substantial problems with the taxonomy of the group and suggests that
several different species maybe concurrently associated to the name Pritha nana. He analyzes five different
populations of presumed P. nana and finds them to be all different from each other. Brignoli concludes that, due to
the insufficient amount of material he could examine, the synonymies previously proposed by Ledoux can neither
be confirmed nor rejected. Interestingly, the illustration he provides for the vulva of a female from Adibe (Brignoli
1982, p. 71, Fig. 7) corresponds to the morphology of P. parva sp. nov. that had already been inadvertently
published previously by Ledoux under the name of P. nana. The figure for the calamistrum of a female collected on
the island of Zannone (Brignoli 1982, p. 71, fig. 10) is, however, incorrect as it portrays the structure as being
anatomically biseriate.
The revisions of Ledoux (1977) and Brignoli (1982) massively influenced the way Pritha nana would be
subsequently interpreted by other authors. The redescription of Ledoux inadvertently associated several diagnostic
characters of P. parva sp. nov. (a short conical bulb, a strongly thickened palpal tibia and a vulva with two pairs of
diverticula, one median apically pointed and one lateral and globular) to P. nana. It comes to no surprise that almost
all subsequent publications either republish the misidentified drawings of Ledoux (Trotta 2005, Wunderlich 2011,
Le Peru 2011) or provide new but incorrect figures and information for P. nana (Brignoli 1982, Heimer & Nentwig
1991, Gray 1995, Nentwig et al. 2016). Unsurprisingly, all the museum material we obtained from around Europe
appeared to have been hastily and carelessly misidentified as P. nana and the same can probably be said for the
remaining museum specimens we could not analyze. Despite the specific name selected by Simon, Pritha nana as
correctly interpreted is presently the largest European member of its genus.
Pritha vestita (Simon, 1873) spec. reval.
In 1873, Simon described the male of Filistata vestita (Simon, 1873) from the island of Corsica near
Caporalino, a small inland town located within the Omessa municipality. He compared it to the similar Filistata
nana, but provided no figure for the palp or description for the female.
In 1892 he provides the first image for the male bulb of this species. The figure is neither particularly detailed
nor informative and strangely portrays short bulbs that differ from the longer (“as long as the tibia”), gradually
tapered morphology he had previously defined in his original description. Filistata vestita is then subsequently
mentioned only in 1914 when Simon, in his “Les
arachnids de France”, provides a simple dichotomous key for the genus and a new geographic locality for this
species in Algeria. No additional figures are presented for either sex.
We successfully obtained and analyzed the vial Rif. AR5422 of Jar 477 from the Muséum national d'Histoire
naturelle of Paris. In the vial we found a male (with both palps) and a female labeled as 2746 Filistata vestita E.S.
Corsica! (Fig. 23A), well preserved but faded in appearance. The label does not specify whether these two
specimens represent the types series or not. They are, however, deposited together in the only vial labeled as F.

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vestita from Simon’s collection and both collected from the same type locality. Thus, it is very reasonable to
assume that these are the specimens he originally observed and described in 1873. The shape of the male bulb is
unique, elongated as in Pritha nana but different in morphology: it appears comparably stockier and less tapered
distally, with an embolus arched downwards and a palpal tibia that is relatively thicker (Fig. 23D–F). The male is
also strikingly small and just barely reaches 2 mm in body length. This is in stark contrast with the considerably
greater sizes always obtained by males of P. nana (2.20–3.08 mm). Such significant size difference can hardly be
explained by mere intraspecific, geographic variability (Fig. 23C). The cephalic pattern is similar between the two
species, but the chiaroscuro appears to be more distinctly reticulated in F. vestita (Fig. 23B). In regards to eye
pattern, the PME of P. nana appear farther from each other by a distance greater than their diameter, while in F.
vestita these appear closer.
The female of P. vestita is also small and just slightly surpasses 2 mm in body length. The cephalothorax is
dark along the borders, with a central, vaguely triangular mark and with a distinct widespread radial pattern. This
well-marked radiation of the carapace was not observed in any specimen of P. nana that we analyzed. Leg tibia and
femur carry distinct light, longitudinal and dorso-lateral bands. The distance between the PME of the female is
greater than the one observed in the male. A chevron pattern is barely visible on the posterior portion of the
abdomen. It appears to have faded out probably due to the age of the specimen and the years of preservation.
Generally, the coloration is uniformly brown.
The vulva of the female was extracted and placed in a microtube. It is formed by two globular and lateral
diverticula and two poorly delineated, medially-developed atrophic diverticula. The vulva does not appear to be
sclerotized and is barely visible. This may be due either to the years of preservation that may have damaged the
tissue or to the fact that the female is young and not yet sexually mature. Either way, the structure of the vulva
clearly and visibly differs from the one that characterizes P. nana.
Considering the numerous differences we observed between the two species, most importantly within the
genitalia, we reject the synonymy proposed by Ledoux (1977) and revalidate the species status of Pritha vestita
(Simon, 1873). Since the male and its diagnostic palps are intact and well-preserved, we here designate this
specimen as the lectotype of this species. Due to both the impossibility of obtaining additional better-preserved
specimens and the average quality of the two types, we here cannot yet provide a complete redescription of the
taxon. The species will presently retain the original description of Simon pending further, more in-depth analyses.
Pritha debilis (Simon, 1911) spec. reval.
Simon (1911) described Filistata debilis from specimens sampled from three different localities in Algeria:
Saïda, Bou-Saada and Ain O’Grab, just to the south of Bou-Saada. The author provides a largely superficial
description of a male and a female. In the same work, Simon provides a brief dichotomous key for the genus, but
gives no figure for this particular species. Pritha debilis has not been mentioned or discussed in any other
subsequent work following this original description.
We obtained and analyzed the vial Rif. AR5427 of Jar 477 from the Muséum national d'Histoire naturelle,
Paris. Within the vial we found two females labeled as “12800 Filistata debilis E.S. TYPE—Saïda, B. Saada Ain
O’Grab” (Fig. 24A). The three sampled localities listed by Simon in the description and on the label probably
corresponded to at least three specimens one of which appears to have gone lost. In fact, the male originally
described by Simon is missing.
The two females are in a poor state of preservation, with broken legs, a fragile consistency and a uniform
white-yellow coloration. Despite its precarious state of conservation, a chevron pattern is visible on the abdomen of
both specimens. The cephalic pattern clearly differs from the one we observed in the other congeneric species. The
radiation of the carapace is conspicuous and the margins are lighter compared to the variegated darker background
(Fig. 24F). The cephalothorax is more elongated compared to the other Pritha species. The most relevant and
unique character, however, is the presence of conspicuous pigmented ring-shaped bands on the legs that are
distributed as follows: Specimen 1: two rings on the femur III–IV (Figs 24D–E); Specimen 2 (more faded in
coloration): two rings on the tibia III; two rings on the tibia IV, one ring on the femur IV.
Simon appears to have ignored or overlooked most of the aforementioned characters. For example, he has no
mention of the unusual and distinctive ring-shaped, dark leg in neither the description nor the dichotomous key
(Simon 1911). The poor state of conservation of the abdomen of both specimens discouraged the authors from
attempting to extract and analyze the vulvae. Nevertheless, the unique leg bands of the females of P. debilis have

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never been observed or reported in P. nana and the synonymy proposed by Ledoux (1977) can definitely be
rejected. The species status of Pritha debilis (Simon, 1911) must be revalidated. Since the male specimen appears
to have been lost and none of the females are in a good state of preservation, we chose to not designate a lectotype
for this species. Due to both the impossibility of obtaining additional better-preserved specimens and the precarious
quality of the century-old, poorly-preserved and partly dismantled type series, we here cannot yet provide a
complete redescription of the taxon. The species will presently retain the original description of Simon pending
further, more in-depth analyses.
Pritha pallida (Kulczyński, 1897)
The taxonomic history of Pritha pallida (Kulczyński, 1897) is, instead, very simple and straight forward. In
1897, in the famous manuscript “Araneae hungariae”, Władysław Kulczyński thoroughly describes Filistata
pallida from two male specimens collected in Crkvenica (Cirquenizza), a small coastal town located in front of the
island of Krk in Croatia. He provides a clear figure of the male palp that perfectly corresponds to the one we
observed in our specimens of P. pallida. He ultimately also provides a small dichotomous key that he uses to
distinguish F. pallida from the other species he collected in Crkvenica: F. nana. He discerns the two by the leg
coloration which is uniformly pale in F. pallida, and mostly pale but with black femora in F. nana.
In 1899, Kulczyński provides a second description of F. pallida, this time from specimens collected on the
Island of Madeira (Portugal) (Kulczyński 1899). He reaffirms the characters he had already defined for males and
describes the female for the first time, providing two important figures that portray their cephalic pattern (Fig.
25C). Both perfectly correspond to the pattern we observed on all conspecific specimens we examined. Several
decades later, Machado (1941) records the species for the Iberian Peninsula (Porto, Portugal) and provides a new
detailed figure of the palp that is consistent with what had been previously illustrated by Kulczyński (1897, 1899).
Types and other specimens originally described and observed by Kulczyński were thoroughly searched in
numerous European institutions to no avail. Ultimately, the whereabouts of his collection were impossible to
determine and his specimens of P. pallida may have gone lost.
The authors traveled to Crikvenica in an effort to collect topotypic specimens of P. pallida. None were found in
either this locality or the surrounding areas. The only species that was collected from the town of Crikvenica was
Pritha sagittata sp. nov. This unusual finding essentially confirms what had already been indirectly documented by
Kulczyński in the past. In his small dichotomous key he in advertently misidentified specimens of P. sagittata sp.
nov. as F. nana. The leg coloration he reports in the key for F. nana, pale with distinctly black femora (Kulczyński
in Chyzer & Kulczyński 1897), is actually typical of P. sagittata sp. nov.
General ecology
The genus Pritha mainly occurs in the Eurasian and Middle-Eastern regions and unusually extends into Indo-
Malayan areas with a few exceptional species. In Europe, it only occurs in the southern territories and almost
exclusively around the Mediterranean basin. The genus has been recorded for Spain, Portugal, France, Switzerland,
Italy, Croatia, Bulgaria, Macedonia and Greece (Nentwig et al. 2016). Currently, no species has ever been found in
central and northern Europe. In Italy, the genus is widely distributed from north to south, on both the mainland
peninsula and Sardinia. According to the Italian material we analyzed, Pritha species currently appear to be
completely absent from mountainous regions located above an altitude of 800 m.
Microhabitats. All the Pritha species discussed in the present work show very strong synanthropic tendencies
and frequently occur in anthropic, often heavily urbanized environments. Their webs can be commonly
encountered on the external walls of buildings and other man-made structures. They are often associated with
cracks, crevices and other similarly protected hideouts which the spiders use as their primary retreats. Occasionally
and in certain favourable situations, specimens have also been observed building webs indoors. While the genus
has been known to occur also in natural settings and has been recorded under stones in garrigue (Ledoux 1977), all
the specimens collected for this work come from anthropic environments, from both heavily urbanized city centers
and smaller suburban areas.
The web of all species appears as a messy, three-dimensional tangle of cribellate silk. This chaotic, sticky
mesh, variable in thickness and used for prey capture, extends outwards from a small, more distinctly structured

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retreat that the spider generally builds within a wall crevice or crack (Fig. 26C–G). The web differs from that of the
often cohabiting, larger Filistata insidiatrix (Fig. 26H) by both the visibly more chaotic structure and the clearly
smaller, less conspicuous and more peripherally located tubular retreat.
FIGURE 28. Distribution of the six species of Pritha discussed in the present work. The records shown derive exclusively
from the examined material and, thus, mostly pertain to Italy.

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In the wild, the webs of Pritha spp. are virtually always covered by copious amounts of debris, particulate and
dust that, given the disturbed nature of the urban environment, very rapidly amass on the external sticky tangled
mesh. Exuviae and prey remains are also left by the spider on the web and the latter can give a good indication on
the dietary preferences of the occupant. Pritha species appear to be generalized predators of a wide range of
arthropods including small dipterans and coleopterans, microlepidopterans, ants, collembola and psocopterans. In
favourable situations, specimens also appear to be very reciprocally tolerant and even populations of different
species were found to coexist and very closely cohabit on the same urban wall. Webs of both adults and immatures
can be found quite close to each other and in great numbers in the same space.
Phenology and reproduction. No significant differences in phenology were observed between the four
species discussed in the present work and females, subadults and juveniles of all Pritha spp. can be found and
collected throughout the whole year. Adult females are quite long-lived and, like all filistatids, will continue to
molt and marginally grow for several years even after reaching sexual maturity (up to four or five according to
specimens raised in captivity). During the winter season, specimens do not enter diapause and merely limit their
activity to the milder, temperate days. Throughout the rest of the year, Pritha species can be found waiting for prey
inside the tubular retreat, deep within the chaotic tangle of cribellate silk. Prey capture occurs indiscriminately
throughout the 24-hour period, but web reparation and construction generally takes place at dusk and throughout
the night. Wandering females or juveniles were only very rarely encountered in the wild.
Males generally mature during the warmer months of the year so the mating season starts in late spring and
continues throughout the summer (May-August). Notable exceptions are represented by those populations of
Pritha that, due to favourable conditions, thrive inside buildings: heating and the higher temperatures can offset
and extend the mating season and, in these particular situations, wandering males can be observed even during
colder seasons. After reaching sexual maturity, adult males abandon their web and stop feeding. Depending on the
levels of temperature and humidity, they will live on for two or three more weeks. Courtship and mating can occur
both during the night and throughout the day. The latter situation was frequently observed in P. nana, where
wandering males and mating couples were commonly encountered even during the hotter and sunnier hours of the
day. Courtship and mating are generally simple and quite straight forward: after having approached the boundaries
of a web, the male gently plucks the silk threads with its anterior legs. This alerts the inhabiting female that slowly
exits its tubular retreat and moves towards the courting male. The insertion of the male emboli is relatively quick
and lasts only a couple of minutes. The male then rapidly scurries away while the female returns to her tubular
retreat.
Depending on the age of the specimen and on the species, egg sacs can contain from 10 to 30 eggs that hatch
after approximately one month (Figs 26A–B). Deposition occurs deep within the web and generally from late
spring to the end of the summer. During each mating season, adult females can produce two to three egg sacs, each
with a gradually decreasing quantity of eggs. Based on observations made in captivity, newly hatched juveniles
appear to initially have poor weaving capabilities. They prefer to utilize either the maternal web or an adjacent one
and prey mostly on tiny arthropods that are too small for the larger occupants. As they develop, they then proceed
to build their own webs which they use to catch minuscule prey items (mostly pscocopterans, collembola and
mites).
Conclusions
The thorough analyses of morphological, characters observed on types and on both museum and freshly collected
specimens, produced the following: redescription of P. nana (Simon, 1868) and P. pallida (Kulczyński, 1897),
description of P. p a r v a sp. nov. and P. sagittata sp. nov., revalidation of P. vestita (Simon, 1873) and P. debilis
(Simon, 1911) with designation of a lectotype for the former and a putative diagnosis of the genus Pritha. This
revisionary work also officially records, for the first time, the presence of P. pallida in Spain and Greece.
There is still plenty of taxonomic confusion that surrounds both the single species of Pritha and the genus as a
whole. Numerous taxa are in dire need of revision as they lack clear diagnostic characters that allow them to be
readily separated from the other congeners. The lack of distinct characters can easily lead to taxonomic
misidentifications, especially when taxa are superficially similar in appearance. Pritha nana, for example, is a
species that got revised but erroneously associated to a morphology that differed from the one originally described.

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Wrong taxonomic interpretations reiterated through time and referenced by different authors eventually lead to
species which lack taxonomic identity. This was the case with P. nana which was, consequentially, quite
challenging to redescribe.
The new species here described were assigned to the genus Pritha because it currently is the only Prithinae
recorded for both Europe and the Mediterranean basin. The genus is poorly defined, lacks unique diagnostic
character states, cannot be considered monophyletic and, at present, can only be putatively redescribed. A complete
morphological and molecular phylogenetic assessment of both the genus and the subfamily is urgently needed for
the real systematic relationships present within both to be ultimately redefined. The present work will be key for
future studies on this genus as it provides several important updates to the taxonomy of the group including a
modern and final redescription of the type species P. nana.
Acknowledgments
First of all, we wish to thank Andrea Colla for his precious input on the early drafts of the manuscript and for
giving us access to all the necessary equipment at the MNHT. We thank Marie-Cécile De Bellis for acting as an
indispensable intermediary between the various museum institutions. Our gratitude also goes to our friend Fabrizio
Fanti and to Roberto Antonio Pantaleoni for their help and useful comments. We also wish to greatly thank Paolo
Tongiorgi for his invaluable suggestions and for the passion, dedication and great contribution he has given to
Italian arachnology. We also extend our gratitude to: Francesca Vita, Massimo Avian and Mauro Tretiach of the
Università degli Studi di Trieste who allowed us to utilize both the SEM and the stereomicroscopes, Tiziano D’Elia
and Alessio Morelli for the illustrations and Vanni Veronesi and Violet Zhao for accurately translating the various
literature. We would like to send our special thanks to curators Zoë Simmons, Christoph Hörweg and Christine
Rollard, for enabling us to examine the type material deposited in the historical collections of their museums, and
to Paolo Pantini, Marco Isaia and Rainer F. Foelix for their help. We finally extend our gratitude to all the collectors
who kindly provided us with all those much-needed specimens: Vladimiro Cortese, Grégoire Meier, Pier Paolo
Balice, Giorgio Gargari, Valerio Giusti, Yorgos Vavatzianis, Francesca Tantalo, Federica Fabietti, Luca Colonnelli,
Luca Cavigioli, Antonio Nardozi, Jacopo Casini, Dario Cinefra and, in particular, Piero Salerno, Pierre Moulet and
Pierre Oger (who curates the beautiful taxonomic gallery located at http://arachno.piwigo.com/).
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