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European Journal of Taxonomy 865: 1–94 ISSN 2118-9773
https://doi.org/10.5852/ejt.2023.865.2097 www.europeanjournaloftaxonomy.eu
2023 · Rivera-Quiroz F.A. & Álvarez-Padilla F.
This work is licensed under a Creative Commons Attribution License (CC BY 4.0).
Monograph
urn:lsid:zoobank.org:pub:845BAE1A-0E6E-484B-BE6D-17F866777B51
Integration or minimalism: twenty-one new species of ghost spiders
(Anyphaenidae: Anyphaena) from Mexico
F. Andrés RIVERA-QUIROZ 1,* & Fernando ÁLVAREZ-PADILLA 2
1 Understanding Evolution Research Group, Naturalis Biodiversity Center,
Darwinweg 2, 2333 CR Leiden, the Netherlands.
2 Laboratorio de Aracnología, Departamento de Biología Comparada, Facultad de Ciencias,
Universidad Nacional Autónoma de México, Distrito Federal, México Ciudad Universitaria,
México D.F. Del. Coyoacán, C.P. 04510, Mexico.
* Corresponding author: andres.riveraquiroz@naturalis.nl
2 Email: fap@ciencias.unam.mx
1 urn:lsid:zoobank.org:author:970DAA18-987A-4819-BE46-A3D399F77409
2 urn:lsid:zoobank.org:author:0B5D4EFE-71E9-4C44-AF70-98C2E7BB37E6
Table of contents
Abstract ................................................................................................................................................... 2
Introduction ............................................................................................................................................ 2
Material and methods ............................................................................................................................ 4
Specimen localities ................................................................................................................................ 4
Descriptive methods .............................................................................................................................. 4
Molecular methods ................................................................................................................................ 5
Abbreviations used in text and gures ................................................................................................... 5
Results ..................................................................................................................................................... 6
Taxonomy .............................................................................................................................................. 6
Celer group ......................................................................................................................................... 6
Anyphaena natachae sp. nov. ........................................................................................................... 9
Anyphaena adnani sp. nov. .............................................................................................................. 13
Anyphaena fernandae sp. nov. ......................................................................................................... 17
Pacica group ................................................................................................................................... 19
Anyphaena triangularis sp. nov. ...................................................................................................... 19
Anyphaena urieli sp. nov. .. ............................................................................................................. 24
Pectorosa group ................................................................................................................................ 28
Anyphaena jimenezi sp. nov. ........................................................................................................... 29
Anyphaena franciscoi sp. nov. ......................................................................................................... 31
Anyphaena bifurcata sp. nov. .......................................................................................................... 34
Anyphaena epicardia sp. nov. .......................................................................................................... 38
Anyphaena dulceae sp. nov. ............................................................................................................ 42
Anyphaena rebecae sp. nov. ............................................................................................................ 47
European Journal of Taxonomy 865: 1–94 (2023)
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Anyphaena miniducta sp. nov. ......................................................................................................... 51
Anyphaena quadrata sp. nov. ......................................................................................................... 56
Anyphaena soae sp. nov. ............................................................................................................... 58
Anyphaena noctua sp. nov. .............................................................................................................. 61
Porta group ....................................................................................................................................... 63
Anyphaena porta sp. nov. ............................................................................................................... 63
Anyphaena salgueiroi sp. nov. ......................................................................................................... 68
Anyphaena ibarrai sp. nov. .............................................................................................................. 70
Anyphaena tonoi sp. nov. ................................................................................................................ 73
Anyphaena megamedia sp. nov. ...................................................................................................... 77
Anyphaena stigma sp. nov. ............................................................................................................. 80
Molecular analyses ............................................................................................................................ 83
Discussion ............................................................................................................................................. 87
Anyphaena genital morphology ........................................................................................................... 87
Integrative vs minimalistic taxonomic approaches ............................................................................. 87
Acknowledgements .............................................................................................................................. 89
References ............................................................................................................................................. 89
Supplementary material ..................................................................................................................... 94
Abstract. The rhythm of biodiversity loss vastly surpasses the number of new species described per
year, with several taxa going extinct without us even knowing about their existence. After more than
250 years of traditional taxonomy, it is clear that the rate of biodiversity description and discovery needs
to be improved. Molecular data has greatly increased the speed of species discovery and accuracy of
taxonomic delimitation. Phenotypic documentation, although relatively slower, is still crucial to identify
species and communicate taxonomic discoveries to a broader audience. Here, we integrate these data
sources to describe a relatively large number of new species of the spider genus Anyphaena and look
into its internal phylogenetic relationships. Our ndings support the existence of several species groups
within Anyphaena (as currently dened), but failed to recover the monophyly of this genus, suggesting
a more comprehensive revision of its species groups and closely related anyphaenid genera is necessary.
The 21 new species described here are: Anyphaena adnani sp. nov., A. bifurcata sp. nov., A. dulceae
sp. nov., A. epicardia sp. nov., A. fernandae sp. nov., A. franciscoi sp. nov., A. ibarrai sp. nov., A. jimenezi
sp. nov., A. megamedia sp. nov., A. miniducta sp. nov., A. natachae sp. nov., A. noctua sp. nov., A. porta
sp. nov., A. quadrata sp. nov., A. rebecae sp. nov., A. salgueiroi sp. nov., A. soae sp. nov., A. stigma
sp. nov., A. tonoi sp. nov., A. triangularis sp. nov. and A. urieli sp. nov.
Keywords. Diversity, Neotropic, Araneae, cybertaxonomy, species online documentation.
Rivera-Quiroz F.A. & Álvarez-Padilla F. 2023. Integration or minimalism: twenty-one new species of ghost spiders
(Anyphaenidae: Anyphaena) from Mexico. European Journal of Taxonomy 865: 1–94.
https://doi.org/10.5852/ejt.2023.865.2097
Introduction
The debate about the taxonomic impediment and how it should be addressed has generated fruitful
discussions about taxonomy’s philosophy, methods and relevance in an evermore data-hungry scientic
context (Godfray 2002, 2007; Knapp et al. 2002; Wilson 2003, 2004; Godfray & Knapp 2004; Wheeler
et al. 2004; Ebach & Holdrege 2005; Hebert & Gregory 2005; Carvalho et al. 2007, 2008; Miller
2007; Godfray et al. 2008; Knapp 2008; Wheeler 2008a, 2008b, 2008c; Clark et al. 2009; Zauner 2009;
Wheeler & Valdecasas 2010; Orr et al. 2020; Sharkey et al. 2021a; Meier et al. 2021). Gathering and
reviewing fundamental data about new taxa can take decades in many cases (Fontaine et al. 2012).
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
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Taxonomic documentation requires a combination of artistic and scientic skills that allows the
generation of hypotheses about species relations and delimitations.
The shadow of the ongoing biodiversity crisis makes it essential to expedite discovery and description of
new species, and the revision of old and disorganized taxa. In the last few decades, taxonomic descriptions
have been supplemented with digital photography, molecular data, and other types of information. This
has transformed taxonomy from a classication system to a data-driven hypothesis-testing science.
Recently, an effort to use DNA barcodes to supersede traditional taxonomic descriptions and diagnoses
resulted in an outstanding work describing more than 400 species of rare parasitoid wasps (Sharkey
et al. 2021a). This work minimized the use of morphological characters and abandoned altogether the
morphological description of the treated species, substituting it for the processed barcodes used in their
analysis. Nevertheless, its approach has sparked a still ongoing discussion (Meier et al. 2021; Zamani
et al. 2021 et al.; Sharkey et al. 2021a, 2021b), and in a reanalysis of this data, Meier et al. (2021)
found inconsistencies when parameters of Sharkey et al. (2021a) were found to have been modied,
suggesting that some of their freshly described species are already in need of revision.
Other attempts to expedite species identications and descriptions include the publication of online
catalogs for morphological documentation applied to spider diversity inventories (Ramirez 2003;
Alvarez-Padilla 2012; Miller & Sac 2014; Álvarez-Padilla et al. 2020; Garcilazo-Cruz & Álvarez-
Padilla 2022). These online catalogs allow the recognition of the existence of taxa, sharing their relevant
data even before the taxon in question has been formally described. Furthermore, this model allows the
world-wide collaboration between experts for new species discovery without the need of sharing physical
specimens (Miller et al. 2014; Álvarez-Padilla et al. 2020). Lastly, the implementation of prospective
data sharing (Agosti & Egloff 2009; Miller et al. 2012; Agosti et al. 2019; Chester et al. 2019) as
implemented by this and a number of other taxonomic journals aids in the access and dissemination of
biodiversity data within global databases such as Plazi (Plazi 2020) and GBIF (GBIF 2022). This can
ultimately aid in the detection of biogeographical patterns, inferring taxa phenologies, among other
applications.
Here, we illustrate the integration of spider inventory online catalogs, barcoding and biodiversity data
sharing to describe 21 new species of the ghost spider genus Anyphaena Sundevall, 1833. Currently,
the family Anyphaenidae Bertkau, 1878 is divided into two subfamilies, Amaurobioidinae Hickman,
1949 and Anyphaeninae Bertkau, 1878, and includes 614 species divided in 57 genera. Of these genera,
Anyphaena is among the three with the largest diversity within this family with 87 described species.
This genus has a wide distribution, being found in Asia, Europe, and the Americas, but more than 72% of
the currently valid species has been found in North and Central America, and the Caribbean (WSC 2022).
A revision of the Nearctic and some Neotropical species recognized four species groups (accentuata
[palearctic], celer, pectorosa and pacica [nearctic and neotropical]) based on genital morphology and
somatic sexual dimorphic characters (Platnick 1974). The majority of species of Anyphaena have good-
quality illustrations and descriptions that facilitate the comparison with the new species described here
(Keyserling 1879; Pickard-Cambridge O. 1896; Pickard-Cambridge F. 1900; Kraus 1955; Platnick 1974,
1977; Dondale & Redner 1982; Sierwald 1988; Brescovit & Lise 1989; Brescovit 1997; Durán-Barrón
et al. 2016). However, the utilization of somatic and genital descriptions, standard views in photography,
illustrations, and molecular data greatly facilitate taxonomic comparisons. This combination of methods
gives multiple lines of evidence for the description of new species, making it easy to test them and
subsequently minimizing taxonomic errors.
European Journal of Taxonomy 865: 1–94 (2023)
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Material and methods
Specimen localities
All species were collected as part of four diversity inventories using standardized protocols for spiders
that specify collecting methods applied, sampling area sizes, and collecting effort units (Coddington et al.
1991, 1996; Scharff et al. 2003; Malumbres-Olarte et al. 2018; and references therein). All specimens
were collected in 96% ethanol. The rst inventory was conducted in San Luis Potosi Sate, municipality
of Xilitla, inside the ecotouristic park Las Pozas. The area is dominated by tropical vegetation fragments
as part of the Sierra Madre Oriental. Here, a 1 ha plot was established with the following central
coordinates and elevation 21.39722° N, 98.99388° W, alt. 662 m. Four expeditions were carried out
from August 2011 to June 2012 (Rivera-Quiroz et al. 2016). The second inventory was made in an oak
forest located 15 km from the Pico de Orizaba Volcano, Veracruz Sate, Municipality Calcahualco, at
2 km in a SW direction from the town of Atotonilco. Two one-hectare plots were established with the
following central coordinates and elevations: Plot I: 19.12569° N, 97.06756° W, alt. 2300 m and Plot
II: 19.29483° N, 97.2045° W, alt. 2388; they are separated by 400 meters in a NW direction. Plot I was
dominated by oak sprouts from old tree stumps, Plot II was a mixture between the same oak sprouts and
pine trees. Three expeditions were made during one year from May 2012 to February 2013. The third
inventory was done in the same Municipality of Calcahualco, 1.6 km in a SE direction from the town of
Xamaticpac. Two contiguous one-hectare plots were established with the following central coordinates
and elevations: Plot I: 19.14172° N, 97.20597° W, alt. 1710 m and Plot II: 19.12614° N, 97.06708° W,
alt. 1700 m. Both plots were dominated by a mixture of oak sprouts from old stumps, tropical vegetation
and agricultural land with 25 years of vegetation overgrowth. Three expeditions were made from April
2013 to February 2014. The plots at Atotonilco and Xamaticpac are separated by ca 15 km in an E
direction. The fourth inventory was done in a primary tropical wet forest at the Biological Station Los
Tuxtlas IB-UNAM, in the municipality of San Andres Tuxtlas, Veracruz State. A single one-hectare plot
was established with central coordinates 18.58225° N, 95.07558° W, alt. 217–172 m. Four expeditions
were made from May 2016 to February 2018 (2017 was sampled only once for logistical reasons)
(Álvarez-Padilla et al. 2020).
Descriptive methods
Online phenotypic documentation for all species was done following a recent protocol that organizes
digital images as standard views across spider diversity inventories. It uses a few high-resolution images
of standard views that cover the main genital and somatic diagnostic characters (Álvarez-Padilla et al.
2020). Images were taken with the following Nikon equipment: digital cameras, a DS-Fi1 connected
to the dissecting microscope SMZ1270 and a DS-Fi2 connected to the glass slide microscope E200,
both cameras controlled with a desktop Dell Inspiron 660s; SMZ1000 and E200 microscopes, both
sharing a DS-Fi3 connected to a HP Slimline 270-a0xx. Custom-made lamps of two 1.5 watts LEDs of
2 cm in diameter were used for illumination. Individual images were stacked using Helicon Focus ver.
6.8.0 (radius 15, smoothing 4 and rendering method B: depth map). More details of the photography
methods are available at www.unamfcaracnolab.com. Drawings were done with the drawing tubes for
the E200 (Y-IDT) and dissecting microscopes (P-IDT). Female genitalia were dissected and digested
with pig pancreatic digestive enzymes overnight (Alvarez-Padilla & Hormiga 2007). Internal structures
were made translucent with methyl salicilate and observed using semi-permanent slide preparations
(Coddington 1983) under the E200 microscope. All measurements and scale bars are in millimeters.
Species descriptions are based on type material unless specied otherwise. Numbers of specimens
measured to establish the species variation are given in each description. Type and examined material
are deposited in the Colección Nacional de Arácnidos, Instituto de Biología, Universidad Nacional
Autónoma de México. PFX000 codes are for voucher specimens that are illustrated or photographed,
ANYM000 codes are for specimens measured, CNAN specimen codes pending. The following species,
A. bromelicola Platnick, 1977, A. catalina Platnick, 1974, A. alachua Platnick, 1974 and A. fraterna
(Banks, 1896), were collected in the diversity inventories mentioned above, were used for comparisons
in this study, and are documented with standard views at www.unamfcaracnolab.com.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
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Molecular methods
Tissues were sampled for most of the new species described here to accurately match the sexes,
infer their relationships, and test the monophyly of the species group within Anyphaena. One or two
legs of every sampled specimen were used. Genomic DNA was extracted using the Qiagen DNeasy
Blood & Tissue Kit (Qiagen, Valencia, CA). A fragment of the commonly used barcode COI was
amplied using the following combination of primers: 1) LCO1490 (5′-ggtcaacaaatcataaagatattgg-3′)
and HCO2198 (5′- taaacttcagggtgaccaaaaaatca-3′) (Folmer et al. 1994); and 2) J1718 spider
(5′-ggnggatttggaaattgrttrgttcc-3′) (Vink et al. 2005) or J1751 (5′-ggatcacctgatatagcattccc-3′) and N2568
(5′-gctacaacataataagtatcatg-3′) (Hedin & Maddison 2001). The reaction mix included 3 µL of template
DNA, 0.25 µL of each primer, 0.5 µL of MgCl2, 0.12 µL Taq DNA polymerase and 3.5 µL of buffer 5 ×
(Mg with ddNTPs). PCR amplication was done as follows: 5 min initial denaturation (95°C); 35 cycles
of 45 s denaturation (95°C), 1 min (48°C) annealing, and 1 min (72°C) extension; 7 min nal extension
(72°C). Sequencing was done at the Laboratorio de Secuenciación Genómica de la Biodiversidad y
de la Salud of the IBUNAM. Sequences were edited in Geneious Prime ver. 2020.0.5 and deposited
in GenBank; accession numbers are reported in Supp. le 1. The COI data was supplemented with
sequences from several anyphaenid and related species: Clubiona canadensis Emerton, 1890 (as an
outgroup), three species of Hibana Brescovit, 1991, two of Aysha Brescovit, 1997 and Oxysoma Nicolet,
1849, and one of 13 other genera (Supp. le 1). A total of 107 sequences were analyzed, including the
type species Anyphaena accentuata (Walckenaer, 1802) and representatives of Platnick’s species groups
(Platnick 1974, 1977; Platnick & Lau 1975): Anyphaena pacica (Banks, 1896), Anyphaena celer
(Hentz, 1847) and Anyphaena pectorosa L. Koch, 1866. All COI sequences were aligned with MAFFT
ver. 7.450 online (https://mafft.cbrc.jp/alignment/server/) with default parameters (matrix available in
Supp. le 1). The best model t was obtained via jModelTest2 (Darriba et al. 2012); GTR+I+G was
selected. SequenceMatrix ver. 1.8 was used to transform the dataset formats. Our barcode dataset was
analyzed under Maximum Likelihood (ML) and Bayesian Inference (BI) approaches using respectively:
RaXML (Stamatakis 2014) in CIPRES (GTR, bootstrap = 1000), and MrBayes ver. 3.2.6 (Ronquist &
Huelsenbeck 2003) (two independent runs with one cold and three heated chains, mcmc = 5 000 000 gen,
samplefreq = 1000, burnin = 20 000). The program Tracer ver. 1.7.1 (Rambaut et al. 2018) was used to
analyze the performance of our BI analyses (Supp. le 2).
Abbreviations used in text and gures
Text
ALE = anterior lateral eyes
AME = anterior median eyes
BEAT = collecting method beating as dened in Scharff et al. (2003)
BERL = leaf litter concentrated and specimens extracted with Berlese funnels (Álvarez-Padilla et al.
2020)
CNAN = Colección Nacional de Arácnidos, Instituto de Biología, Universidad Nacional Autónoma
de México
CRP = cryptic
d = dorsal
LUD = looking down
LUP = looking up (Scharff et al. 2003)
p = prolateral
PF = Pitfall trap
PLE = posterior lateral eyes
PME = posterior median eyes
r = retrolateral
RTA = male pedipalp retrolateral tibial apophysis
v = ventral
European Journal of Taxonomy 865: 1–94 (2023)
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Figures
a = atrium
aRTA = retrolateral tibial apophysis anterior branch
cd = copulatory ducts
co = copulatory openings
e = embolus
fd = fertilization ducts
h = hood
lb = epigynal atrium lateral borders
ma = median apophysis
mp = epigynum midpiece
pRTA = retrolateral tibial apophysis posterior branch
ptp = prolaetral tegular projection
rtp = retrolateral tegular projection (tegular projection homologies follow Brescovit 1997)
s = spermatheca
sr = seminal receptacle
st = subtegulum
vtp = ventral tegular projection
t = tegulum
vMTA = median tibial apophysis ventral branch
Results
Taxonomy
Class Arachnida Cuvier, 1812
Order Araneae Clerck, 1757
Family Anyphaenidae Bertkau, 1878
Genus Anyphaena Sundevall, 1833
Type species
Aranea accentuata Walckenaer, 1802; gender feminine.
Celer group
Diagnosis
The celer group can be separated from other Anyphaena species groups by the following characters.
Males
Coxae lacking ventral spurs (spine-like modications) but might present clusters of setae (Fig. 3F)
or tubercles (cuticle blunt protuberances of various shapes) (Fig. 41E–F). The palp of the celer group
(Fig. 1A–C) (Platnick 1974; Platnick & Lau 1975) presents a tegulum divided in three lamella-shaped
projections, described as prolateral tegular projection, retrolateral tegular projection and ventral tegular
projection following the homologies in Brescovit (1997) and Oliveira & Brescovit (2021). The embolus is
a visible sclerite (in the un-expanded palp), tubular in shape, and curved dorsally. The median apophysis
(or conductor in Platnick & Lau 1975) is a thin, transparent and curved sclerite, articulated at the base of
the ventral tegular projection by exible cuticle. In expanded pedipalps, the retrolateral tegular projection
and prolateral tegular projection are lamella-shaped and partially articulated. In the unexpanded bulb,
these projections lie dorsally like two folded tortillas around the other sclerites (Fig. 1A–C). Their RTA
is dominated by the anterior and posterior branches, the former usually larger and shaped as a folded
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
7
cuticular lamella, the latter is usually smaller and its shape varies considerably between species (Platnick
1974, 1977; Platnick & Lau 1975) (Fig. 5A, C).
Females
Externally the epigynum has a hood close to the anterior margin, a sclerotized median plate below the
hood, and a transparent atrium delineated by two sclerotized lateral borders. The copulatory openings
are difcult to observe; however, they are located near the lateral border's posterior edge (Figs 2A,
4E, 5E, 7E, 8E, 9E, 18A). Internally, the copulatory and fertilization ducts are usually sclerotized and
Fig. 1. Anyphaena spp., expanded pedipalps. A–C. A. natachae sp. nov., ♂ (PXF302). A. Ventral
view. B. Retrolateral view. C. Prolateral view. – D–F. A. urieli sp. nov., allotype, ♂ (CNAN-T01521).
D. Ventral view. E. Retrolateral view. F. Prolateral view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
8
Fig. 2. Anyphaena spp., epigynum anatomies. A–B. A. adnani sp. nov., paratype, ♀ (CNAN-T01546).
A. Epigynum, ventral view. B. Epigynum, dorsal view. – C–D. A. rebecae sp. nov., holotype, ♀
(CNAN-T01536). C. Epigynum, ventral view. D. Epigynum, dorsal view. – E–F. A. salgueiroi sp. nov.,
holotype, ♀ (CNAN-T01537). E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars:
0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
9
variable in length and shape (Platnick 1974, 1977; Platnick & Lau 1975). The seminal receptacles are
cylindrical in shape and also located near the posterior edge. The spermathecae are usually spherical to
oval and close to the posterior margin (Figs 2B, 4F, 5F, 7F, 8F, 9F, 18B).
Anyphaena natachae sp. nov.
urn:lsid:zoobank.org:act:AA547563-9B11-4AF3-BFEA-0738DBB895D4
Figs 1A–C, 3–5, 52
Differential diagnosis
Males of A. natachae sp. nov. have the RTA posterior branch single and cone-shaped, the anterior RTA
branch rectangular with a curved apical margin (Figs 4C–D, 5C–D). Both RTA branches are similar to
those of A. hespar Platnick, 1974 (Platnick & Lau 1975: gs 39–40), A.wanlessi Platnick & Lau, 1975
(Platnick & Lau 1975: gs 103–104) and A. inferens Chamberlin, 1925 (Brescovit 1997: gs 7–8, 11–
13), but differ from these species by the ventral tegular projection tip being at and squared, the slightly
curved median apophysis, and the retrolateral tegular projection margin not overlapping the median
apophysis (Figs 4A–D, 5A, D). Females of this species are differentiated from those of A. inferens
(Platnick & Lau 1975: gs 115–116; Brescovit 1992: gs 1–2, 1997: gs 9–10) by the acute posterior
edge of the epigynal midpiece and the atrium lateral borders being longer and less curved (Figs 4E–F,
5E–F).
Etymology
The species epithet is dedicated to Natacha Merritt, friend of the authors and a taxonomy supporter.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m;
4–17 Feb. 2014; Arcanolab team leg.; oak and tropical wet forest fragment; LUP; CNAN-T01532.
Allotype
MEXICO • ♂; same locality as for holotype; 2–11 Oct. 2013; BEAT; CNAN-T01516.
Paratypes
MEXICO • 2 ♀♀, 1 ♂; same collection data as for allotype; CNAN-T01562.
Additional material
MEXICO • 1 ♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
19–27 Apr. 2013; Arcanolab team leg.; oak and tropical wet forest fragment; LUP • 19 ♀♀, 2 ♂♂; same
collection data as for preceding; 2–11 Oct. 2013; BEAT • 2 ♂♂; same collection data as for preceding •
2 ♂♂; same collection data as for preceding; ANYM067 • 2 ♂♂; same collection data as for preceding;
ANYM068 • 4 ♀♀; same collection data as for preceding; ANYM072 • 3 ♀♀; same collection data as
for preceding; CRP • 5 ♀♀, 2 ♂♂; same collection data as for preceding; LUP • 3 ♀♀; same collection
data as for preceding; 4–17 Feb. 2014; BEAT • 1 ♀; same collection data as for preceding; CRP • 1 ♀;
Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m; 19–27 Apr. 2013; Aracnolab team leg.;
oak and tropical wet forest fragment; LUP • 21 ♀♀, 2 ♂♂; same collection data as for preceding; 2–11
Oct. 2013; BEAT • 3 ♀♀; same collection data as for preceding; ANYM071 • 1 ♂, expanded palp; same
collection data as for preceding; PXF302 • 3 ♀♀; same collection data as for preceding; LUP • 4 ♀♀;
same collection data as for preceding; 4–17 Feb. 2014; LUP
European Journal of Taxonomy 865: 1–94 (2023)
10
Description
Female
Total length 4.88. Carapace light yellow, pattern with light brown longitudinal marks over the ocular
quadrangle, same pattern but darker delineating cephalic area and center, two dark longitudinal bands
from ocular quadrangle lateral sides, around thoracic area and carapace margins, clypeus light yellow
(Fig. 3A, D). Sternum yellow, darker at margins, intercoxal triangles present on all legs. Labium
brown, white at tip, longer than wide. Endites dark yellow, rectangular, slightly broader at tip (Fig. 3C).
Chelicerae darker than cephalothorax, paturon dorsal surface with brown reticulated pattern (Fig. 3D),
promargin with four teeth, retromargin with seven to eight denticles. Leg coloration: light yellow with
scattered brown patches irregularly distributed from femora to tarsi (Fig. 3A–C). Abdomen light yellow,
homogenously covered with brown patches concentrated in transverse chevrons over posterior central
line, ventral surface with clear rectangle and dark longitudinal midline cut by tracheal spiracle at center
Fig. 3. Anyphaena natachae sp. nov., paratypes (CNAN-T01562). A. ♀, dorsal habitus. B. ♀, lateral
habitus. C. ♀, ventral habitus. D. ♀, prosoma, anterior view. E. ♂, prosoma, anterior view. F. ♂,
prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
11
of abdomen (Fig. 3A–C). Epigynum atrium longer than wide, delineated by slightly curved sclerotized
borders. Hood elliptical, wider than long. Midpiece same width as hood, rectangular, posterior half
tapering. Copulatory openings inside atrium posterior margins. Copulatory ducts hook-shaped, entering
posterior margins of spermathecae. Fertilization ducts entering spermathecae dorsal to copulatory ducts.
Seminal receptacles curved and near posterior atrium border (Figs 4E–F, 5E–F). Cephalothorax length
2.23, thoracic width 1.61, cephalic width 1.12. Clypeus height 0.09. Eye diameters: AME 0.09, ALE
0.12, PME 0.12, PLE 0.12. Eye interdistances: AME–AME 0.09, AME–ALE 0.04, ALE–PLE 0.09,
PME–PME 0.17, PME–PLE 0.11. Femur lengths: I 1.71, II 1.63, III 1.41, IV 1.83. Leg spination: femur I
d1-1-1, p0-0-2, r0-0-1. Tibia I v3-2-2, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II
d1-1-1, p0-0-1, r0-0-1. Tibia II v2-2-1, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III
d1-1-1, p0-0-1, r0-0-1. Tibia III v1-1-2, p1-1-0, r1-1-0. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV
d1-1-1, p0-0-1, r0-0-1. Tibia IV v1-1-2, p1-1-0, r1-1-0. Metatarsus IV v1-1-2, p1-1-2, r1-1-2.
Fig. 4. Anyphaena natachae sp. nov., paratypes (CNAN-T01562). A. ♂, pedipalp, ventral view. B. ♂,
pedipalp, prolateral view. C. ♂, pedipalp, dorsal view. D. ♂, pedipalp, retrolateral view. E. ♀, epigynum,
ventral view. F. ♀, epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
12
Male
Total length 4.68. Cephalothorax and abdomen reticulated patterns as in female, coloration dark
brown (Fig. 3E). All coxae covered with scattered setae (Fig. 3F). Chelicerae promargin with six
teeth, retromargin with eight to nine denticles. Pedipalp ventral tegular projection straight, tip roughly
rectangular (Figs 1A, 4A–B, 5A–B). Prolateral tegular projection curved and surrounding ventral
tegular projection (Figs 1A–B, 4A–B, 5A–B). Retrolateral tegular projection enclosing other two tegular
apophyses, ventral margin straight (Figs 1A–B, 4A, D, 5A, D). Median apophysis slightly curved and
projected apically (Figs 4A, 5A). Embolus black, thicker than median apophysis, and coiling towards
Fig. 5. Anyphaena natachae sp. nov., paratypes (CNAN-T01562). A. ♂, pedipalp, ventral view. B. ♂,
pedipalp, prolateral view. C. ♂, pedipalp, dorsal view. D. ♂, pedipalp, retrolateral view. E. ♀, epigynum,
ventral view. F. ♀, epigynum, dorsal view. Scale bars: A–D = 0.2 mm; E–F = 0.1 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
13
apex (Figs 1A–B, 4A, 5A). RTA anterior branch at, curved and longer than wide, RTA posterior branch
single, spine-shaped and located at base (Figs 4C–D, 5A, C–D). Prolateral apophysis absent (Figs 4B–
C, 5BC). Pedipalp tibia slightly longer than wide, ventral surface with large white tubercle at base
(Figs 4A–D, 5A–D). Cephalothorax length 2.26, thoracic width 1.71, cephalic width 0.93. Clypeus
height 0.07. Eye diameters: AME 0.09, ALE 0.12, PME 0.11, PLE 0.12. Eye interdistances: AME–
AME 0.05, AME–ALE 0.01, ALE–PLE 0.06, PME–PME 0.13, PME–PLE 0.8. Femur lengths: I 2.37,
II 2.15, III 1.78, IV 2.29. Leg spination as in female.
Variation
Females (N = 10): total length 4.69 (± 0.5), cephalothorax length 2.15 (± 0.07), thoracic width 1.57
(± 0.05), cephalic width 1.07 (± 0.05), femur I 1.69 (± 0.04). Males (N = 10): total length 4.44 (± 0.19),
cephalothorax length 2.15 (± 0.09), thoracic width 1.67 (± 0.05), cephalic width 0.91 (± 0.03), femur I
2.23 (± 0.09).
Distribution
This species is found in oak mixed with tropical wet forest fragments around Pico de Orizaba Volcano
National Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray. This species
is present year-round.
Anyphaena adnani sp. nov.
urn:lsid:zoobank.org:act:44F20A0E-E32D-43DC-B7B2-1AE4AA62AC21
Figs 2A–B, 6–8, 52
Differential diagnosis
Males of A. adnani sp. nov. have the RTA anterior and posterior branches similar to those of A. bromelicola
Platnick, 1977 (Platnick 1977: gs 5–6), but differ from this species by the sharper point of the RTA
anterior branch and the ventral tegular projection tip acute and being less curved (Figs 7A, D, 8A, D).
Females of this species are differentiated from those of A. bromelicola (Platnick 1977: gs 7–8) by the
attened hood, shorter midpiece, and wider atrium (Figs 7E, 8E).
Etymology
The species epithet is dedicated to Dr Adnan Selimovic, friend of the authors and taxonomy supporter.
Material examined
Holotype
MEXICO • ♂; Veracruz, Calcahualco, Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m;
4–17 Feb. 2014; Arcanolab team leg.; oak and tropical wet forest fragment; BEAT; CNAN-T01522.
Allotype
MEXICO • ♀; same locality as for holotype, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m; 19–27 Apr.
2013; BEAT; AR_084; GenBank: ON619641; CNAN-T01499.
Paratypes
MEXICO • 2 ♀♀; same collection data as for allotype; CNAN-T01543 • 2 ♂♂; same collection data as
for allotype; CNAN-T01544 • 1 ♂; same collection data as for allotype; LUP; CNAN-T01545 • 2 ♀♀;
same collection data as for holotype; 19–27 Apr. 2013; CNAN-T01546.
European Journal of Taxonomy 865: 1–94 (2023)
14
Additional material
MEXICO • 3 ♀♀; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
19–27 Apr. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT • 2 ♀♀; same
collection data as for preceding; ANYM057 • 2 ♀♀, 1 ♂; same collection data as for preceding; CRP
• 1 ♂; same collection data as for preceding; LUP; ANYM064 • 2 ♀♀; same collection data as for
preceding; 4–17 Feb. 2014; BEAT • 3 ♀♀; same collection data as for preceding; Xamaticpac, Plot II;
19.12614° N, 97.06708° W; alt. 1700 m; 19–27 Apr. 2013; Aracnolab team leg.; oak and tropical wet
forest fragment; BEAT • 3 ♀♀, 1 ♂; same collection data as for preceding; ANYM060 • 1 ♂; same
collection data as for preceding; ANYM063 • 1 ♀; same collection data as for preceding; CRP • 3 ♀♀;
same collection data as for preceding; LUP • 1 ♂; same collection data as for preceding; AR_0811;
GenBank: ON619640 • 1 ♀; same collection data as for preceding; ANYM059 • 1 ♀; same collection
Fig. 6. Anyphaena adnani sp. nov. A–D. Paratype, ♀ (CNAN-T01546). E–F. Paratype, ♂
(CNAN-T01545). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view.
E. Prosoma, anterior view. F. Prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
15
data as for preceding; 4–17 Feb. 2014; BEAT • 1 ♂; same collection data as for preceding; AR_081;
GenBank: ON619640 • 2 ♀♀; same collection data as for preceding; ANYM058 • 1 ♂; same collection
data as for preceding; ANYM065 • 1 ♀; same collection data as for preceding; CRP.
Description
Female
Total length 4.15. Carapace yellow, pattern with brown longitudinal bands around cephalic area extending
over most of thoracic area and clypeus (Fig. 6A, D). Sternum surface yellow, covered with brown
patches, intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide. Endites
yellow, rectangular, slightly broader at tip (Fig. 6C). Chelicerae dark yellow, paturon covered with brown
patches (Fig. 6B, D), promargin with four teeth, retromargin with six to seven denticles. Leg coloration:
light yellow with scattered brown patches irregularly distributed from femora to tarsi (Fig. 6A–C).
Abdomen white, hirsute, dorsal surface with two central brown chevrons, lateral and ventral surfaces
white, covered with isolated dark patches, ventral surface with clear rectangle and dark longitudinal
Fig. 7. Anyphaena adnani sp. nov. A–D. Paratype, ♂ (CNAN-T01545).E–F. Paratype, ♀ (CNAN-
T01546). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp,
retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
16
midline cut by tracheal spiracle at center of abdomen (Fig. 6A–C). Epigynum atrium rectangular,
slightly longer than wide, delineated by almost straight sclerotized borders. Hood attened, midpiece
semicircular. Copulatory openings inside atrium posterior border (Figs 7F, 8F). Copulatory ducts coiled
360° around spermathecae, entering dorsal surface of spermathecae. Spermathecae dorsally projected.
Fertilization ducts entering central margins of spermathecae and partially covered by copulatory ducts.
Fig. 8. Anyphaena adnani sp. nov. A–D. Paratype, ♂ (CNAN-T01545). E–F. Paratype, ♀
(CNAN-T01546). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars:
A–D = 0.2 mm; E–F = 0.1 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
17
Seminal receptacles curved and near posterior atrium border (Fig. 8E–F). Cephalothorax length 1.81,
thoracic width 1.32, cephalic width 0.85. Clypeus height 0.07. Eye diameters: AME 0.06, ALE 0.1, PME
0.09, PLE 0.1. Eye interdistances: AME–AME 0.04, AME–ALE 0.04, ALE–PLE 0.06, PME–PME 0.2,
PME–PLE 0.07. Femur lengths: I 1.29, II 1.22, III 1.15, IV 1.46. Leg spination: femur I d1-1-1, p0-0-2,
r0-0-1. Tibia I v2-2-2, p1-1-1, r1-1-1. Metatarsus I v2-2-2, p1-1-1, r1-1-1. Femur II d1-1-1, p0-0-1, r0-0-
1. Tibia II v2-2-2, p1-1-1, r1-1-1. Metatarsus II v2-2-2, p1-1-1, r1-1-1. Femur III d1-1-1, p0-0-1, r0-0-1.
Tibia III v1-1-2, p1-1-0, r1-1-0. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1.
Tibia IV v1-1-2, p1-1-1, r1-1-1. Metatarsus IV v2-1-2, p1-1-2, r1-1-2.
Male
Total length 4.2. Cephalothorax (Fig. 6E) and abdomen coloration as in female. Ventral surface of coxa
smooth (Fig. 6F). Pedipalp ventral tegular projection straight, tip narrow and slightly curved (Figs 7A,
8A). Prolateral tegular projection curved and dorsally surrounding ventral tegular projection (Figs 7A–B,
8A–B). Retrolateral tegular projection enclosing other two tegular apophyses, ventral margin projected
over median apophysis. Medial apophysis slightly curved, projected towards prolateral tegulum edge
(Figs 7A, 8A). Embolus very sclerotized, same width as median apophysis, and coiling towards apex
(Figs 7A–B, 8A, D). RTA anterior branch at, curved and longer than wide, RTA posterior branch length
almost half that of anterior branch, basal spine with small sharp point (Figs 7D, 8D). Prolateral apophysis
absent (Figs 7B–C, 8B–C). Pedipalp tibia slightly longer than wide, with small white tubercle on basal
ventral surface (Figs 7A, D, 8A, D). Cephalothorax length 2.1, thoracic width 1.58, cephalic width 0.9.
Clypeus height 0.09. Eye diameters: AME 0.09, ALE 0.12, PME 0.09, PLE 0.07. Eye interdistances:
AME–AME 0.03, AME–ALE 0.01, ALE–PLE 0.05, PME–PME 0.18, PME–PLE 0.09. Femur lengths:
I 1.95, II 1.66, III 1.46, IV 1.95. Leg spination as in female except: femur I p0-1-2. Tibia I v2-2-1.
Femur II p0-1-2. Tibia II v2-2-1. Tibia III p1-1-1, r1-1-1. Metatarsus IV v2-2-2.
Variation
Females (N = 10): total length 4.17 (± 0.39), cephalothorax length 1.79 (± 0.08), thoracic width 1.34
(± 0.07), cephalic width 0.86 (± 0.03), femur I 1.32 (± 0.06). Males (N = 10): total length 4.13 (± 0.11),
cephalothorax length 1.94 (± 0.1), thoracic width 1.53 (± 0.06), cephalic width 0.86 (± 0.03), femur I
1.57 (± 0.13).
Distribution
This species is found in oak mixed with tropical wet forest fragments around Pico de Orizaba Volcano
National Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray. This species
is present all year-round.
Anyphaena fernandae sp. nov.
urn:lsid:zoobank.org:act:350499A4-08EA-4C45-8265-F55302FE61A3
Figs 9, 18A–B, 52
Differential diagnosis
Females of A. fernandae sp. nov. are differentiated from those of A. autumna Platnick, 1974 (Platnick
1974: gs 39, 45) and the other species of the celer group by the following features: ellipsoid hood,
wider than long, midpiece rhomboidal and tapering posteriorly, spermathecae wider than long, and short
copulatory ducts (Figs 9E–F, 18A–B).
Etymology
The species epithet is dedicated to Fernanda Lia Rivera, sister of the rst author.
European Journal of Taxonomy 865: 1–94 (2023)
18
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m; 4–14
Oct. 2012; Arcanolab team leg.; oak forest fragment; BEAT; CNAN-T01526.
Description
Female
Total length 5.5. Carapace yellow, more sclerotized over ocular area, pattern with brown longitudinal
bands around cephalic area extending to thoracic area and clypeus (Fig. 9A, D). Sternum surface yellow,
margins darker, intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide.
Fig. 9. Anyphaena fernandae sp. nov., holotype, ♀ (CNAN-T01526). A. Dorsal habitus. B. Lateral
habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view. F. Epigynum, dorsal
view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
19
Endites yellow, rectangular, slightly broader at tip (Fig. 9C). Chelicerae slightly dark yellow, paturon
dorsal surface covered with two darker lines (Fig. 9B, D), promargin with four teeth, retromargin with
eight to nine denticles. Leg coloration: yellow with scattered brown patches irregularly distributed from
femora to tarsi (Fig. 9A–C). Abdomen dark brown, dorsal surface with four central darker chevrons,
lateral and ventral surfaces yellow, covered with reticulated dark lines, ventral surface with clear
rectangle and dark longitudinal midline cut by tracheal spiracle at center of abdomen (Fig. 9A–C).
Epigynum atrium wider than long, delineated by curved sclerotized lateral borders. Copulatory openings
inside posterior margins of atrium (Fig. 9E–F). Copulatory ducts short and comma-shaped, entering
spermathecae ventrally. Fertilization ducts entering spermathecae dorsally. Seminal receptacles not
visible (Fig. 18A–B). Cephalothorax length 2.26, thoracic width 1.61, cephalic width 1.05. Clypeus
height 0.09. Eye diameters: AME 0.07, ALE 0.13, PME 0.11, PLE 0.13. Eye interdistances: AME–
AME 0.05, AME–ALE 0.02, ALE–PLE 0.06, PME–PME 0.15, PME–PLE 0.11. Femur lengths: I 1.95,
II 1.76, III 1.46, IV 2.05. Leg spination: femur I d1-1-1, p0-0-2, r0-0-0. Tibia I v2-1-1, p1-1-1, r0-0-1.
Metatarsus I v2-2-0, p1-1-0, r1-1-0. Femur II d1-1-1, p0-0-2, r0-0-0. Tibia II v2-1-1, p1-1-1, r0-0-1.
Metatarsus II v2-2-0, p1-1-0, r1-1-0. Femur III d1-1-1, p0-0-1, r0-0-1. Tibia III v2-2-2, p1-1-0, r1-1-0.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-0, r1-1-0.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Unknown.
Variation
Only type specimen known.
Distribution
Only known from the type locality (Fig. 52).
Natural history
Collected by beating vegetation.
Pacica group
Diagnosis
The pacica group can be separated from other species groups of Anyphaena by the following characters.
Males
Both of the species described here show modications in the venter of coxae II, III and IV (Figs 12E–
F, 15E–F). Coxae II show a small and relatively inconspicuous bump. Coxae III and IV have fully
developed ventral spurs. The palp of the pacica group shows a morphology similar to that of the
pectorosa and porta groups. It can be separated from both by the large dorsal cymbium bulge, and the
larger RTA posterior branch that projects dorso-laterally (Figs 11A, C–D, 14A, C–D).
Females
Both new species differ from other Anyphaena by having an enlarged epigynal plate bulging anteriorly
(Figs 10F, 13F). This enlarged plate shows two sclerotized spots that might serve as an anchor point
for male palp structures; the spermathecae are usually oval to spherical (Figs 11E–F, 12G–H, 14E–F,
15G–H).
European Journal of Taxonomy 865: 1–94 (2023)
20
Anyphaena triangularis sp. nov.
urn:lsid:zoobank.org:act:99F21C7F-3CC1-413F-9269-57931511CE81
Figs 10–12, 52
Differential diagnosis
The epigynal plate in A. triangularis sp. nov. is broad, more than two thirds of the epigastric furrow
length, anteriorly projected and ventrally distended, epigynum anterior projection with two deep comma-
shaped pits (Figs 10C, F, 11E–F, 12G (arrow), H). Similar epigyna are found in A. urieli sp. nov., but they
differ by the atter anterior projection and copulatory duct paths. Males of A. triangularis sp. nov. are
differentiated from those of A. urieli sp. nov. by their larger cymbium basal projection (Figs 11C, 12C
(arrow)), ventral tegular projection retrolateral margin without a translucent section, median apophysis
distal hook smaller, the RTA posterior branch wider, and the coxa II tubercle larger (Figs 11A–D, 12A–
F).
Etymology
The species epithet refers to the triangular shape of the epigynal plate in ventral view.
Material examined
Holotype
MEXICO • ♀; San Luis Potosi, Xilitla City, Las Pozas; 21.39722° N, 98.99388° W; alt. 662 m; 10–15
Jun. 2012; Aracnolab team leg.; tropical wet forest fragment; BEAT; CNAN-T01541.
Allotype
MEXICO • ♂; same collection data as for holotype; CNAN-T01520.
Paratypes
MEXICO • 1 ♀; same collection data as for holotype; LUP; CNAN-T01579 • 1 ♀; same collection data
as for holotype; CNAN-T01580 • 1 ♂; same collection data as for holotype; CNAN-T01577 • 1 ♂; same
collection data as for holotype; CNAN-T01578.
Additional material
MEXICO • 3 ♂♂; San Luis Potosi, Xilitla City, Las Pozas; 21.39722° N, 98.99388° W; alt. 662 m;
10–15 Jun. 2012; Aracnolab team leg.; tropical wet forest fragment; BEAT • 1 ♀; same collection data
as for preceding; ANYM085 • 1 ♂; same collection data as for preceding; CRP • 1 ♀; same collection
data as for preceding; ANYM086 • 1 ♀; same collection data as for preceding; ANYM087 • 3 ♂♂;
same collection data as for preceding; LUD • 2 ♀♀; same collection data as for preceding; ANYM090 •
2 ♂♂; same collection data as for preceding; ANYM091 • 4 ♂♂; same collection data as for preceding;
ANYM092 • 5 ♂♂; same collection data as for preceding; LUP • 1 ♀; same collection data as for
preceding; ANYM088 • 1 ♀; same collection data as for preceding; ANYM089 • 1 ♀; same collection
data as for preceding; 27–31 Jul. 2011; CRP; AR_002; GenBank: ON619642.
Description
Female
Total length 6.9. Carapace yellow, ocular area slightly darker, pattern with two darker longitudinal
brown parallel bands around cephalic area and fovea, clypeus dark yellow (Fig. 10A, D). Sternum
surface white, intercoxal triangles present on all legs. Labium yellow, white at tip, longer than wide.
Endites yellow, rectangular, slightly broader at tip (Fig. 10C). Chelicerae dark yellow, paturon dorsum
without pattern (Fig. 10B, D), promargin with ve teeth, retromargin with eleven to twelve denticles.
Leg coloration: yellow covered with darker patches from femora to tarsi. Abdomen yellow, hirsute,
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
21
dorsal surface delineated with light brown pattern and several chevrons over posterior half, anterior
central half with darker median band, lateral surfaces light brown pattern only at top, lateral sides and
ventral surface white, tracheal spiracle at middle (Fig. 10A–C). Copulatory openings under semicircular
and sclerotized hood. Hood with straight edge slightly projected at center. Atrium short, lateral borders
sinuous, extended in posterior region (Figs 11E–F, 12G–H). Copulatory ducts sclerotized, short,
entering spermathecae on anterior surfaces. Seminal receptacles rounded and located in anterior third of
copulatory ducts. Fertilization ducts short, straight, and emerging from lateral surfaces of spermathecae
(Figs 11E–F, 12G–F). Cephalothorax length 2.93, thoracic width 2.1, cephalic width 0.97. Clypeus
height 0.09. Eye diameters: AME 0.07, ALE 0.13, PME 0.12, PLE 0.12. Eye interdistances: AME–
AME 0.07, AME–ALE 0.04, ALE–PLE 0.06, PME–PME 0.12, PME–PLE 0.09. Femur lengths: I 3.38,
II 3.05, III 2.29, IV 3.43. Leg spination: femur I d1-1-1, p0-2-1, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1.
Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1.
Fig. 10. Anyphaena triangularis sp. nov. A–D, F. Paratype, ♀ (CNAN-T01579). E. Paratype, ♂
(CNAN-T01577). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior
view. E. Prosoma, anterior view. F. Epigynum, lateral view. Scale bars: A–C = 1.0 mm; D–E = 0.5 mm;
F = 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
22
Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p0-1-1, r0-1-1.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p0-1-1, r0-1-1.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Total length 6.9. Cephalothorax and abdomen coloration as in female (Fig. 10E). Retromargin of
chelicerae with seven to eight denticles. Coxa III tubercle roughly rectangular with posterior apical
spine. Coxa IV with a long, curved spur, located at retrolateral basal corner (Fig. 12E–F). Ventral tegular
apophysis bifurcated (Fig. 12A (arrow)). Median apophysis hook-shaped in ventral view. Embolus long,
translucent and liform (Figs 11A, D, 12A, D). RTA anterior branch curved with sclerotized margin,
posterior branch of RTA large, distal edge with translucent keel (Figs 11A, C–D, 12A, C–D). Prolateral
apophysis of palpal tibia small, conical (Figs 11B–C, 12B–C). Pedipalp tibia proportions longer than
wide, stridulatory ridges extending from ventral branch of median tegular apophysis to tibial distal
margin (Figs 11A–B, D, 12A–B, D). Cephalothorax length 2.78, thoracic width 2.15, cephalic width
Fig. 11. Anyphaena triangularis sp. nov. A–D. Paratype, ♂ (CNAN-T01577). E–F. Paratype, ♀
(CNAN-T01579). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
23
Fig. 12. Anyphaena triangularis sp. nov. A–F. Paratype, ♂ (CNAN-T01577). G–H. Paratype, ♀
(CNAN-T01579). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
24
0.97. Clypeus height 0.11. Eye diameters: AME 0.09, ALE 0.16, PME 0.16, PLE 0.15. Eye interdistances:
AME–AME 0.05, AME–ALE 0.04, ALE–PLE 0.05, PME–PME 0.1, PME–PLE 0.09. Femur lengths: I
3.52, II 3.19, III 2.38, IV 3.57. Leg spination as in female except: femur IV r0-1-1.
Variation
Females (N = 10): total length 6.94 (± 0.36), cephalothorax length 2.73 (± 0.11), thoracic width 2.06
(± 0.07), cephalic width 1.12 (± 0.06), femur I 3.32 (± 0.12). Males (N = 10): total length 6.66 (± 0.26),
cephalothorax length 2.74 (± 0.1), thoracic width 2.19 (± 0.1), cephalic width 1.06 (± 0.06), femur I 3.6
(± 0.06).
Distribution
This species is found in primary tropical wet forest fragments (Fig. 52).
Natural history
Most specimens were collected over vegetation by beating, direct searching and cryptic search on the
ground.
Anyphaena urieli sp. nov.
urn:lsid:zoobank.org:act:82E29AF5-6DE2-4561-B7F1-4B88170DBDF6
Figs 1D–F, 13–15, 52
Differential diagnosis
The epigynal plate in A. urieli sp. nov. is broad, more than two thirds of epigastric furrow length,
globular, anteriorly projected and ventrally compressed (Figs 13C, F, 14E–F), epigynum anterior half
with two deep pits (Figs 14E (arrow), F, 15G (arrow), H). Similar epigyna are found in A. triangularis
sp. nov., but differ from those in A.urieli sp. nov. by their anterior projection being ventrally distended,
the shape of the anterior pits, and the path of the copulatory ducts extremely short (Figs 14A–B, 15G–
H). Males of A. urieli sp. nov. are differentiated from those of related species by their ventral tegular
projection base being bifurcated with a process shaped like a bird head (Figs 14A, 15A (arrows)), and
from A. triangularis sp. nov. by the smaller basal projection of the cymbium (Figs 14C–D, 15D (arrows)),
the ventral tegular projection of the retrolateral margin with a transparent cuticular edge, the narrower
RTA posterior branch (Figs 14A, C–F, 15A, C–F), and the smaller coxa II tubercle (Fig. 15E–F).
Etymology
The species epithet is dedicated to Uriel Garcilazo-Cruz, Mexican arachnologist, and former member of
the second author’s lab.
Material examined
Holotype
MEXICO • ♀; Veracruz, San Andres Tuxtla, Estación de Biología Tropical Los Tuxtlas IB-UNAM;
18.58225° N, 95.07558° W; alt. 217–172 m; 16–22 May 2016; Aracnolab team leg.; primary tropical
wet forest; LUP; CNAN-T01542.
Allotype
MEXICO • ♂; same collection data as for holotype; 20–27 Nov. 2017; CNAN-T01521.
Paratypes
MEXICO • 1 ♂, 1 ♀; same collection data as for allotype; BEAT; CNAN-T01582 • 1 ♀; same collection
data as for holotype; BEAT; CNAN-T01581.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
25
Additional material
MEXICO • 3 ♀♀, 2 ♂♂; Veracruz, San Andres Tuxtla, Estación de Biología Tropical Los Tuxtlas IB-
UNAM; 18.58225° N, 95.07558° W; alt. 217–172 m; 16–22 May 2016; Aracnolab team leg.; primary
tropical wet forest; BEAT • 1 ♀; same collection data as for preceding; ANYM073 • 1 ♀; same collection
data as for preceding; ANYM074 • 2 ♂♂; same collection data as for preceding; ANYM083 • 1 ♀,
1 ♂; same collection data as for preceding; LUD • 1 ♀; same collection data as for preceding; LUP;
ANYM077 • 1 ♂; same collection data as for preceding; ANYM080 • 1 ♂; same collection data as for
preceding; ANYM081 • 3 ♂♂; same collection data as for preceding; ANYM084 • 3 ♀♀, 1 ♂; same
collection data as for preceding • 1 ♀; same collection data as for preceding; ANYM075 • 1 ♀, 1 ♂;
same collection data as for preceding; LUP • 1 ♀; same collection data as for preceding; ANYM078
• 1 ♀, 2 ♂♂; same collection data as for preceding; 20–27 Nov. 2017; BEAT • 2 ♀♀; same collection
data as for preceding; ANYM076 • 1 ♂; same collection data as for preceding; ANYM082 • 2 ♀♀, 1 ♂;
Fig. 13. Anyphaena urieli sp. nov. A–D. Holotype, ♀ (CNAN-T01542). E. Paratype, ♂ (CNAN-T01582).
F. Paratype, ♀ (CNAN-T01581). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma,
anterior view. E. Prosoma, anterior view. F. Epigynum, lateral view. Scale bars: A–C = 1.0 mm;
D–E = 0.5 mm; F = 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
26
same collection data as for preceding; LUP • 1 ♀; same collection data as for preceding; ANYM079 •
1 ♀, 2 ♂♂; same collection data as for preceding; 9–16 Feb. 2018; BEAT • 4 ♂♂; same collection data
as for preceding; LUP.
Description
Female
Total length 5.0. Carapace yellow, ocular area slightly darker, pattern with two darker longitudinal
brown parallel bands around cephalic area and fovea, clypeus yellow (Fig. 13A, D). Sternum surface
white, intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide. Endites
yellow, rectangular, slightly broader at tip (Fig. 13C). Chelicerae yellow, paturon dorsum without
pattern (Fig. 13B, D), promargin with ve teeth, retromargin with eight to nine denticles. Leg coloration:
yellow, darker from distal section of femora to tarsi. Abdomen yellow, hirsute, dorsal pattern brown,
Fig. 14. Anyphaena urieli sp. nov. A–D. Paratype, ♂ (CNAN-T01582). E–F. Paratype, ♀ (CNAN-
T01581). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp,
retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
27
Fig. 15. Anyphaena urieli sp. nov. A–F. Paratype, ♂ (CNAN-T01582). G–H. Paratype, ♀
(CNAN-T01581). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
28
surface delineated with wide band and several chevrons over posterior central half, anterior central
half without this pattern, brown pattern of lateral surfaces only at top, lateral sides and ventral surface
white covered with faint darker spots, tracheal spiracle at middle (Fig. 13A–C). Copulatory openings
under straight hood slightly projected at center. Copulatory ducts sclerotized, short, slightly curved,
entering spermathecae on lateral surfaces. Seminal receptacles in anterior third in copulatory ducts
inconspicuous. Fertilization ducts short, straight, and emerging from lateral surfaces of spermathecae
(Figs 14E–F, 15G–H). Cephalothorax length 2.24, thoracic width 1.8, cephalic width 0.97. Clypeus
height 0.07. Eye diameters: AME 0.09, ALE 0.12, PME 0.12, PLE 0.12. Eye interdistances: AME–
AME 0.05, AME–ALE 0.02, ALE–PLE 0.04, PME–PME 0.12, PME–PLE 0.09. Femur lengths: I 2.58,
II 2.42, III 1.81, IV 2.65. Leg spination: femur I d1-1-1, p0-2-1, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1.
Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1.
Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v1-1-2, p0-1-1, r0-1-1.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p0-1-1, r0-1-1.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Total length 5.8. Cephalothorax and abdomen coloration as in female (Fig. 13E). Chelicerae retromargin
with eleven to twelve denticles. Coxa III tubercle roughly rectangular with posterior apical spur. Coxa
IV with a long, curled spur curved at tip (Fig. 15E–F). Median apophysis hook-shaped in ventral view.
Embolus long, translucent and liform (Figs 14A, 15A). RTA anterior branch curved with sclerotized
margin, apical edge of posterior branch of RTA with two triangular tips (Figs 14A, C–D, 15A, C–D).
Palpal tibia prolateral apophysis small, conical (Figs 14B–C, 15B–C). Pedipalp tibia proportion slightly
longer than wide, stridulatory ridges extending from ventral branch of median tegular apophysis to
tibial distal margin (Figs 14A–B, D, 15A–B, D). Cephalothorax length 2.54, thoracic width 2.05,
cephalic width 0.9. Clypeus height 0.06. Eye diameters: AME 0.07, ALE 0.17, PME 0.15, PLE 0.15.
Eye interdistances: AME–AME 0.05, AME–ALE 0.02, ALE–PLE 0.05, PME–PME 0.11, PME–PLE
0.09. Femur lengths: I 3.24, II 2.9, III 2.19, IV 3.1. Leg spination as in female except: femur I p0-1-2.
Femur II p0-1-2. Metatarsus II r1-1-0. Femur III r0-0-1. Tibia III v1-2-2, p1-1-0, r1-1-0. Femur IV r0-
1-1. Tibia IV p1-1-1, r1-1-1.
Variation
Females (N = 10): total length 5.83 (± 0.44), cephalothorax length 2.43 (± 0.14), thoracic width 1.86
(± 0.11), cephalic width 0.99 (± 0.05), femur I 2.89 (± 0.33). Males (N = 10): total length 5.48 (± 0.3),
cephalothorax length 2.44 (± 0.1), thoracic width 1.99 (± 0.07), cephalic width 0.88 (± 0.03), femur I
3.15 (± 0.16).
Distribution
This species is found in primary tropical wet forest (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching and beating vegetation.
Pectorosa group
Diagnosis
The pectorosa group can be separated from other Anyphaena species groups by the following characters.
Males
Coxae II, III and IV ventral surface armed with spurs and/or tubercles. Coxae II tubercle sometimes
reduced. Coxae III and IV commonly with spurs, the spurs on coxa III may be bifurcated (Fig. 35E–F).
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
29
The general shape of the male palp is similar to that in the porta and pacica groups. It can be separated
from the former by the relative length of the palpal tibia and the general shape of the RTA. It differs from
the latter by the relative length and direction of the RTA posterior branch.
Female
Epigynum with an atrium anterior to the spermathecae. Copulatory openings anterior to the atrium usually
under the hood. Copulatory ducts very sclerotized and usually longer than 1.5 times the spermathecae
diameter and always projecting posteriorly, ‘encircling’ the atrium in dorsal view (A. epicardia sp. nov.,
A. dulceae sp. nov., A. soae sp. nov., A. rebecae sp. nov. and A. franciscoi sp. nov.). Exceptions to the
standard length and shape of the copulatoy ducts, and sometimes the overall shape of the atrium, can be
found in A. noctua sp. nov., A. jimenezi sp. nov, and A. bifurcata sp. nov. Spermatheca well sclerotized,
oval to spherical except in A. soae sp. nov. (Figs 18C, 21C, 38C).
Anyphaena jimenezi sp. nov.
urn:lsid:zoobank.org:act:44A36DC8-80BF-4197-95E8-B4D86F42D9F4
Figs 16, 18C–D, 52
Differential diagnosis
Females of A. jimenezi sp. nov. are differentiated from those of all species of the pectorosa and pacica
groups (Keyserling 1879; Pickard-Cambridge O. 1896; Pickard-Cambridge F. 1900; Kraus 1955;
Platnick 1974; Dondale & Redner 1982; Sierwald 1988; Brescovit & Lise 1989; Brescovit 1997; Durán-
Barrón et al. 2016) by the curved mustache-shaped lateral borders, and the genital openings being
adjacent at the center of the epigynal plate (Figs 16E, 18C). Central and adjacent genital openings are
also present in females of A. tonoi sp. nov. (Fig. 46E–F), but both species differ by the path and length
of the copulatory ducts (Fig. 18B–C).
Etymology
The species epithet is dedicated to the Mexican arachnologist Dr María Luisa Jiménez.
Material examined
Holotype
MEXICO • ♀; Veracruz, San Andres Tuxtla, Estación de Biología Los Tuxtlas IB-UNAM; 18.58225° N,
95.07558° W; alt. 217–172 m; 9–16 Feb. 2018; Arcanolab team leg.; primary tropical wet forest; LUP;
CNAN-T01529.
Description
Female
Total length 6.07. Carapace yellow, with few dark patches of sclerotized cuticle and setae, delineating
cephalic area and around fovea (Fig. 16A, D). Sternum surface white, intercoxal triangles present on all
legs. Labium light brown, white at tip, longer than wide. Endites light yellow, rectangular, broader at
tip (Fig. 16C). Chelicerae slightly darker than cephalothorax (Fig. 16B, D), promargin with ve teeth,
retromargin with eleven to twelve denticles. Leg coloration darker than cephalothorax and scattered
brown patches irregularly distributed from femora to tarsi (Fig. 16A–C). Abdomen yellow-gray, dorsal
and lateral surfaces with scattered darker patches, ventral surface without patterns, tracheal spiracle
closer to epigastric furrow (Fig. 16A–C). Epigyum hood short, slightly curved, and over both genital
openings. Anterior plate area delineated by two sclerotized curved notches at corners (Figs 16E–F, 18C–
D). Seminal receptacles in anterior third of copulatory ducts. Copulatory duct path U-shaped, bending
dorsally, and curving again to enter external lateral surface of spermathecae. Fertilization ducts short,
cylindrical, entering spermathecae dorsally. Spermathecae longer than wide and between copulatory
European Journal of Taxonomy 865: 1–94 (2023)
30
ducts coils (Fig. 18C–D). Cephalothorax length 2.7, thoracic width 2.2, cephalic width 1.2. Clypeus
height 0.1. Eye diameters: AME 0.1, ALE 0.13, PME 0.13, PLE 0.15. Eye interdistances: AME–AME
0.05, AME–ALE 0.04, ALE–PLE 0.06, PME–PME 0.16, PME–PLE 0.13. Femur lengths: I 3.32, II
3.27, III 2.39, IV 3.37. Leg spination: femur I d1-1-1, p0-1-1, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1.
Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1.
Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-0-1, r0-0-1. Tibia III v2-2-2, p1-1-0, p1-1-0.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-1, r1-1-1.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Unknown.
Fig. 16. Anyphaena jimenezi sp. nov., holotype, ♀ (CNAN-T01529). A. Dorsal habitus. B. Lateral
habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view. F. Epigynum, dorsal
view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
31
Variation
Only type specimen known.
Distribution
Only known from the type locality (Fig. 52).
Natural history
Collected at night by direct searching over vegetation.
Anyphaena franciscoi sp. nov.
urn:lsid:zoobank.org:act:D00F2524-98DA-4755-82FE-FC064FCD1CCD
Figs 17, 18E–F, 52
Differential diagnosis
The female of A. franciscoi sp. nov. is differentiated from all those of species of the pectorosa and
pacica groups by the following features: atrium trapezoidal, hood broad and posteriorly curved at the
center (Figs 17E–F, 18E–F). The female of A. rebecae sp. nov. also has a similar trapezoidal atrium,
and copulatory duct paths (Fig. 28E–F), but it differs from that of A. franciscoi sp. nov. by the notched
hood and smaller size (7.23 ± 1.1). Anyphaena soae sp. nov. shares similar copulatory duct paths
and spermathecal proportions with both species, but differs by the oval atrium and copulatory ducts
extending beyond the hood (Fig. 36E–F).
Etymology
The species epithet is dedicated to Francisco Rivera, father of the rst author.
Material examined
Holotype
MEXICO • ♀; San Luis Potosi, Xilitla City, Las Pozas; 21.39722° N, 98.99388° W; alt. 662 m; 26–30
Mar. 2012; Arcanolab team leg.; tropical wet forest fragment; LUP; CNAN-T01527.
Description
Female
Total length 9.6. Carapace light yellow, with two faint darker bands delineating cephalic area and around
fovea (Fig. 17A, D). Sternum surface white, intercoxal triangles present on all legs. Labium brown,
white at tip, longer than wide. Endites yellow, rectangular, broader at tip (Fig. 17C). Chelicerae brown
without dorsal pattern (Fig. 17B, D), promargin with ve teeth, retromargin with nine to ten denticles.
Legs femora base yellow, orange-brown distally and from patella to tarsi (Fig. 17A–C). Abdomen
dorsal surface white and delineated by two light brown parallel longitudinal lines, lateral surfaces
white and dorsally delineated by light brown longitudinal lines, ventral surface center with faint light
brown longitudinal band from epigastric furrow to spinneret bases, tracheal spiracle closer to epigastric
furrow (Fig. 17A–C). Anterior area of epigynal plate delineated laterally by two faint triangular pits.
Genital openings at both sides of anterior edge of atrium under hood. Copulatory duct slightly curved
and entering surface of anterior spermathecae. Seminal receptacles oval in anterior third of copulatory
ducts. Fertilization ducts short, cylindrical, and entering lateral surface of spermathecae (Figs 17E–F,
18E–F). Cephalothorax length 4.37, thoracic width 2.73, cephalic width 1.55. Clypeus height 0.13. Eye
diameters: AME 0.11, ALE 0.15, PME 0.16, PLE 0.16. Eye interdistances: AME–AME 0.07, AME–
ALE 0.04, ALE–PLE 0.07, PME–PME 0.15, PME–PLE 0.13. Femur lengths: I 4.24, II 3.71, III 2.98, IV
4.29. Leg spination: femur I d1-1-1, p0-1-1, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0,
p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0,
European Journal of Taxonomy 865: 1–94 (2023)
32
p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p1-1-0, p1-1-0. Metatarsus III v2-2-2,
p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-1, r1-1-1. Metatarsus IV v2-2-2,
p1-1-2, r1-1-2.
Male
Unknown.
Variation
Only type specimen known.
Distribution
Only known from the type locality (Fig. 52).
Fig. 17. Anyphaena franciscoi sp. nov., holotype, ♀ (CNAN-T01527). A. Dorsal habitus. B. Lateral
habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view. F. Epigynum, dorsal
view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
33
Natural history
Collected at night by direct searching over vegetation.
Fig. 18. Anyphaena spp. A–B. A. fernandae sp. nov., holotype, ♀ (CNAN-T01526). A. Epigynum,
ventral view. B. Epigynum, dorsal view. – C–D. A. jimenezi sp. nov., holotype, ♀ (CNAN-T01529).
C. Epigynum, ventral view. D. Epigynum, dorsal view. – E–F. A. franciscoi sp. nov., holotype, ♀
(CNAN-T01527). E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
34
Anyphaena bifurcata sp. nov.
urn:lsid:zoobank.org:act:35864EEB-BF52-411A-82FC-C5BC0AD40F7E
Figs 19–21, 52
Differential diagnosis
The epigynum in A. bifurcata sp. nov. is broad, the anterior edge is more than two thirds of the abdomen
width, the shape triangular and projected below the epigastric furrow (Fig. 19C), posterior third of the
epigynal plate narrower, the tip excavated with a small central atrium where the copulatory openings
are located (Figs 20E–F, 21G–H). Similar epigyna plates and atria are found in females of A. pectorosa,
but differ from those of A.bifurcata sp. nov. by the trapezoidal shape and convex posterior margin
(Dondale & Render 1982: gs 328–329). Broad epigynal plates are also present in A. triangularis
sp. nov. and A. urieli sp. nov., but they are anteriorly projected in both species (Figs 10C, F, 13C, F).
Males of A. bifurcata sp. nov. are differentiated from those of related species by their ventral tegular
Fig. 19. Anyphaena bifurcata sp. nov. A–D. Holotype, ♀ (CNAN-T01523). E–F. Paratype, ♂
(CNAN-T01549). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view.
E. Prosoma, anterior view. F. Prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
35
projection base being excavated and bifurcated with a thin rectangular process (Figs 20A–B, 21A–
B, arrows), median apophysis with a small hook in ventral and retrolateral view, and cymbium base
dorsally projected (Figs 20A–D, 21A–D). A similar cymbium and bifurcated ventral tegular projections
are found in males of A. triangularis sp. nov. (Fig. 12A (arrow)) and A. urieli sp. nov. (Fig. 15A (arrow)),
but they differ from those of A. bifurcata sp. nov. by not being excavated.
Etymology
The species epithet is taken from the Spanish word bifurcado and refers to the bifurcated ventral tegular
projection diagnostic of this species.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m; 15–
24 Feb. 2013; Aracnolab team leg.; oak forest fragment; LUP; CNAN-T01523.
Fig. 20. Anyphaena bifurcata sp. nov. A–D. Paratype, ♂ (CNAN-T01549). E–F. Paratype, ♀
(CNAN-T01550). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
36
Allotype
MEXICO • ♂; same collection data as for holotype; AR_035; GenBank: ON619650; CNAN-T01512.
Paratypes
MEXICO • 3 ♂♂; same collection data as for holotype; CNAN-T01547 • 2 ♀♀; same collection data as
for holotype; CNAN-T01548 • 1 ♂; same collection data as for holotype; CNAN-T01549 • 1 ♀; same
collection data as for holotype; 21–30 Mar. 2012; CNAN-T01550.
Additional material
MEXICO • 2 ♂♂; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m;
15–24 Feb. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BERL • 4 ♂♂, 1 ♀; same
collection data as for preceding; CRP • 1 ♀; same collection data as for preceding; AR_063; GenBank:
ON619629 • 1 ♀; same collection data as for preceding; ANYM023 • 3 ♀♀, 4 ♂♂; same collection data
as for preceding; LUP • 1 ♂; same collection data as for preceding; AR_036; GenBank: ON619651 •
2 ♂♂; same collection data as for preceding; ANYM019 • 1 ♂; same collection data as for preceding;
PF; AR_037; GenBank: ON619652 • 1 ♀; same collection data as for preceding; 21–30 May 2012;
LUP; AR_062; GenBank: ON619628 • 1 ♂; same collection data as for preceding; 4–14 Oct. 2012;
BEAT; AR_034; GenBank: ON619649 • 1 ♀; same collection data as for preceding; ANYM021 • 1 ♀;
same collection data as for preceding; BERL; ANYM022.
Description
Female
Total length 7.1. Carapace yellow, pattern with darker longitudinal bands over cephalic and thoracic
areas, carapace margins and clypeus yellow (Fig. 19A, D). Sternum surface white, intercoxal triangles
present on all legs. Labium brown, white at tip, longer than wide. Endites dark yellow, rectangular,
slightly broader at tip (Fig. 19C). Chelicerae orange, paturon dorsum with darker and diffuse line pattern
(Fig. 19B, D), promargin with four teeth, retromargin with eight to nine denticles. Leg coloration: light
yellow with scattered dark patches, pattern slightly darker at metatarsi and tarsi. Abdomen yellow,
hirsute, dorsal surface covered with scattered small brown patches, pattern lighter at center, forming
longitudinal band, lateral surfaces with same pattern faded ventrally, ventral surface white, tracheal
spiracle at middle (Fig. 19A–C). Copulatory openings inside small internal edges of atrium. Copulatory
ducts sclerotized, short, slightly curved, entering spermathecae via lateral surfaces below seminal
receptacles. Fertilization ducts short, straight, and emerging from anterior surface of spermathecae
(Figs 20E–F, 21G–H). Cephalothorax length 2.68, thoracic width 2.2, cephalic width 1.19. Clypeus
height 0.11. Eye diameters: AME 0.07, ALE 0.15, PME 0.12, PLE 0.13. Eye interdistances: AME–
AME 0.07, AME–ALE 0.06, ALE–PLE 0.1, PME–PME 0.17, PME–PLE 0.1. Femur lengths: I 2.94,
II 2.68, III 2.1, IV 2.77. Leg spination: femur I d1-1-1, p0-1-2, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1.
Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1.
Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p0-1-1, r0-1-1.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-1, r1-1-1.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Total length 5.8. Cephalothorax and abdomen coloration as in female except yellow chelicerae (Fig. 19E).
Coxa II ventral surface with broad and shallow tubercle. Coxa III tubercle roughly rectangular with
spine at middle. Coxa IV spur located at retrolateral basal corner (Figs 19F, 21E–F). Embolus long,
translucent and liform. RTA branches short, anterior branch curved with sclerotized margin, posterior
branch small, triangular in dorsal and ventral views (Figs 20A, C–D, F 21A, C–D). Prolateral apophysis
of palpal tibia small, conical (Figs 20B–C, 21B–C). Pedipalp tibia longer than wide, ventral surface with
median stridulatory ridges. Median tibial apophysis absent (Figs 20C, 21A, D). Cephalothorax length
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
37
2.68, thoracic width 2.2, cephalic width 0.9. Clypeus height 0.11. Eye diameters: AME 0.07, ALE 0.11,
PME 0.13, PLE 0.13. Eye interdistances: AME–AME 0.06, AME–ALE 0.05, ALE–PLE 0.09, PME–
PME 0.15, PME–PLE 0.12. Femur lengths: I 3.1, II 2.94, III 2.26, IV 2.9. Leg spination as in female
except: femur II r0-2-1. Femur IV p0-1-1, r0-1-1.
Fig. 21. Anyphaena bifurcata sp. nov. A–F. Paratype, ♂ (CNAN-T01549). G–H. Paratype, ♀
(CNAN-T01550). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
38
Variation
Females (N = 10): total length 7.2 (± 0.67), cephalothorax length 2.84 (± 0.12), thoracic width 2.19
(± 0.07), cephalic width 1.2 (± 0.04), femur I 2.88 (± 0.08). Males (N = 10): total length 5.79 (± 0.17),
cephalothorax length 2.79 (± 0.09), thoracic width 2.21 (± 0.06), cephalic width 1.0 (± 0.05), femur I
3.13 (± 0.19).
Distribution
This species is found in oak forest fragments around Pico de Orizaba Volcano National Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching, Berlese funnels and cryptic searching
on the ground vegetation.
Anyphaena epicardia sp. nov.
urn:lsid:zoobank.org:act:D75F5ABC-6E2F-4EA7-8594-45670D3F3632
Figs 22–24, 52
Differential diagnosis
Females from Anyphaena epicardia sp. nov. can be differentiated from those of all other American
species of the pectorosa and pacica groups by the heart-shaped atrium of the epigynum (Fig. 23E–F),
and from those of A. dulceae sp. nov. by the straight copulatory ducts (Fig. 24G–H). Males of this species
resemble those of A. fraterna (Platnick 1974: gs 52, 56, 60) and A. simoni Becker, 1878 (Brescovit &
Lise 1989: gs 1–9) by their elongated ventral tegular projection, but differ from both species by the
presence of an apical translucent curved lamella (Figs 23A–D, 24A–D (arrows)), a broader median
apophysis in ventral view (Figs 23A, 24A), an acute spine on the ventral corner of the trapezoidal RTA
posterior branch (Figs 23D, 24D) and the coxa III tubercle bifurcated (Figs 22F, 24E–F), this last feature
more similar to A. simoni than A. fraterna.
Etymology
The species epithet is a combination from the Spanish words epigino and cardia, referring to the heart-
shaped epigynum atrium, a diagnostic feature of this species.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m; 21–
30 May 2012; Aracnolab team leg.; oak and tropical wet forest fragment; LUP; CNAN-T01525.
Allotype
MEXICO • ♂; same collection data as for holotype; 15–24 Feb. 2013; CNAN-T01514.
Paratypes
MEXICO • 1 ♀; same collection data as for holotype; AR_073; GenBank ON619623; CNAN-T01557 •
1 ♂; same collection data as for preceding; Atotonilco, Plot II; 19.29483° N, 97.2045° W; alt. 2388 m;
21–30 May 2012; Arcanolab team leg.; oak and pine forest fragment; BEAT; AR_069; GenBank:
ON619619; CNAN-T01556.
Additional material
MEXICO • 2 ♂♂; same collection data as for holotype; 15–24 Feb. 2013; BEAT • 7 ♂♂; same collection
data as for preceding; LUP • 1 ♂; same collection data as for preceding; PF • 1 ♂; same collection data
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
39
as for preceding; 21–30 May 2012; BEAT; AR_72; GenBank: ON619622 • 1 ♂; same collection data
as for preceding; CRP; AR_071; GenBank: ON619621 • 1 ♀; same collection data as for preceding;
ANYM002 • 1 ♀, 8 ♂♂; same collection data as for preceding; LUP • 3 ♂♂; same collection data as
for preceding; ANYM006 • 1 ♂; same collection data as for preceding; PF • 1 ♂; same collection data
as for preceding; 4–14 Oct. 2012; BEAT • 4 ♂♂; same collection data as for preceding; Atotonilco, Plot
II; 19.29483° N, 97.2045° W; alt. 2388 m; 15–24 Feb. 2013; Aracnolab team leg.; oak forest fragment;
BEAT • 1 ♂; same collection data as for preceding; BERL; ANYM005 • 1 ♂; same collection data as
for preceding; CRP • 1 ♂; same collection data as for preceding; LUP • 7 ♂♂; same collection data as
for preceding; 21–30 May 2012; BEAT • 1 ♀; same collection data as for preceding; ANYM003 • 1 ♀,
2 ♂♂; same collection data as for preceding; LUP • 1 ♀; same collection data as for preceding; AR_076;
GenBank: ON619624 • 1 ♂; same collection data as for preceding; AR_70; GenBank: ON619620 • 1 ♀;
Fig. 22. Anyphaena epicardia sp. nov. (♀: CNAN-T01525; ♂: CNAN-T01514). A. ♀, dorsal habitus.
B. ♀, lateral habitus. C. ♀, ventral habitus. D. ♀, prosoma, anterior view. E. ♂, prosoma, anterior view.
F. ♂, prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
European Journal of Taxonomy 865: 1–94 (2023)
40
same collection data as for preceding; ANYM001 • 1 ♀; same collection data as for preceding; 4–14
Oct. 2012; BEAT; ANYM004.
Description
Female
Total length 9.6. Carapace dark yellow, hirsute, with darker bands over cephalic, thoracic area and
clypeus (Fig. 22A, D). Sternum yellow, slightly darker at margins, intercoxal triangles present on all
legs. Labium brown, white at tip, longer than wide. Endites brown, rectangular, slightly broader at
tip. Chelicerae dark brown (Fig. 22C–D), promargin with four teeth, retromargin with eight to nine
denticles. Abdomen dorsum white, hirsute, with few scattered light brown patches around edges, lateral
sides and center, ventral surface with diffuse longitudinal line between tracheal spiracle and spinnerets,
tracheal spiracle in center of abdomen (Fig. 22A–C). Leg coloration: dark yellow with brown patches
at distal end of femora, patella and tibia. Metatarsus and tarsus brown. Epigynum hood absent, lateral
border sinuous. Copulatory openings on anterior internal margin of heart-shaped atrium. Copulatory
Fig. 23. Anyphaena epicardia sp. nov. A–D. Paratype, ♂ (CNAN-T01556). E–F. Holotype, ♀
(CNAN-T01557). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
41
Fig. 24. Anyphaena epicardia sp. nov. A–F. Paratype, ♂ (CNAN-T01556). G–H. Holotype, ♀
(CNAN-T01557). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
42
ducts parallel. Seminal receptacles cylindrical, projected dorsally and located at junction between
spermathecae and copulatory duct entrance (Figs 23E–F, 24G–H). Copulatory and fertilization ducts
attached to posterior margin of spermathecae. Fertilization ducts short and slightly curved (Fig. 24G–
H). Cephalothorax length 3.27, thoracic width 2.44, cephalic width 1.35. Clypeus height 0.11. Eye
diameters: AME 0.11, ALE 0.15, PME 0.15, PLE 0.15. Eye interdistances: AME–AME 0.06, AME–
ALE 0.05, ALE–PLE 0.07, PME–PME 0.16, PME–PLE 0.11. Femur lengths: I 3.56, II 3.27, III 2.49, IV
3.41. Leg spination: femur I d1-1-1, p0-1-2, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0,
p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0. p1-1-1, r1-1-1. Metatarsus II v2-2-0,
p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p0-1-1, r1-1-1. Metatarsus III v2-2-2,
p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-1, r1-1-1. Metatarsus IV v2-2-2,
p1-1-2, r1-1-2.
Male
Total length 7.8. Cephalothorax coloration as in female except dark yellow chelicerae and endites
(Fig. 22E). Coxa II tubercle roughly triangular, coxa IV spur base broad (Figs 22F, 24E–F). Abdomen
yellow with darker patches covering dorsal surface and lateral sides, ventral surface with longitudinal
line between tracheal spiracle and spinnerets. Pedipalp ventral tegular projection in prolateral view
with distal translucent keel (Figs 23B, 24B), Embolus liform and translucent (Fig. 24A). RTA anterior
branch semicircular with sclerotized edge and translucent center, RTA posterior branch cuticle as in
tibia, more sclerotized and roughly rectangular (Figs 23A, C–D, 24A, C–D). Prolateral apophysis small
and cone-shaped (Figs 23B, 24B). Pedipalp tibia slightly longer than wide. Median tibial apophysis
ventral branch displaced towards proximal border of tibia (Figs 23A, D, 24A, D). Cephalothorax length
3.37, thoracic width 2.73, cephalic width 1.10. Clypeus height 0.15. Eye diameters: AME 0.11, ALE
0.13, PME 0.09, PLE 0.15. Eye interdistances: AME–AME 0.05, AME–ALE 0.04, ALE–PLE 0.06,
PME–PME 0.15, PME–PLE 0.11. Femur lengths: I 4.20, II 3.76, III 2.98, IV 3.95. Leg spination as in
female except: femur II p0-1-2, r0-2-2. Femur IV p0-1-1.
Variation
Females (N = 9): total length 9.1 (± 0.82), cephalothorax length 3.26 (± 0.15), thoracic width 2.6
(± 0.15), cephalic width 1.37 (± 0.07), femur I 3.63 (± 0.14). Males (N = 10): total length 7.66 (± 0.43),
cephalothorax length 3.38 (± 0.23), width 2.79 (± 0.19), cephalic width 1.21 (± 0.09), femur I 4.20
(± 0.28).
Distribution
This species is found in oak and oak-pine forests around Pico de Orizaba Volcano National Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray; some male
specimens were caught in PF traps and Berlsese funnels. This species is present year-round.
Anyphaena dulceae sp. nov.
urn:lsid:zoobank.org:act:3452AA1C-E9A1-43E2-88BD-7EACA6AA1898
Figs 25–27, 52
Differential diagnosis
Females of Anyphaena dulceae sp. nov. can be differentiated from those of all other species of the
pectorosa and pacica groups by the heart-shaped epigynal atria (Fig. 26E–F), and from A. epicardia
sp. nov. by the middle coil of the copulatory ducts (Fig. 27G–H). Males of this species resemble those
of A. fraterna (Platnick 1974: gs 52, 56, 60) and A. simoni (Brescovit & Lise 1989: gs 1–9) by their
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
43
elongated ventral tegular projection with a rounded tip and the hook-shaped median apophysis, but
differ from these species by the presence of a rectangular process at the ventral tegular projection base
(Figs 26A, 27A (arrows)), the more elongated and curved median apophysis hook (Figs 26A, 27A), the
RTA posterior branch being hook-shaped (Figs 26D, 27D) and the large spurs of coxa II–IV (Figs 25F,
27E–F).
Etymology
The species epithet is dedicated to Dulce Flor Piedra-Jiménez, Mexican arachnologist, and former
member of the second author’s lab.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m; 21–
30 May 2012; Aracnolab team leg.; oak and tropical wet forest fragment; LUP; CNAN-T01524.
Allotype
MEXICO • ♂; same collection data as for holotype; BEAT; CNAN-T01513.
Paratypes
MEXICO • 1 ♀; same collection data as for holotype; CRP; AR_033; GenBank: ON619655;
CNAN-T01555 • 1 ♀; same collection data as for preceding; 15–24 Feb. 2013; CNAN-T01553 • 1 ♀;
same collection data as for preceding; Atotonilco, Plot II; 19.29483° N, 97.2045° W; alt. 2388 m; 21–
30 May 2012; Arcanolab team leg.; oak and pine forest fragment; LUP; CNAN-T01552 • 1 ♂; same
collection data as for preceding; CNAN-T01551 • 1 ♀; same collection data as for preceding; 15–24
Feb. 2013; BEAT; AR_031; GenBank: ON619653; CNAN-T01554.
Additional material
MEXICO • 1 ♀, 2 ♂♂; same collection data as for holotype; 15–24 Feb. 2013; BEAT • 1 ♀; same
collection data as for preceding; CRP, ANYM010 • 5 ♀♀, 2 ♂♂; same collection data as for preceding;
LUP • 1 ♀; same collection data as for preceding; ANYM011 • 2 ♂♂; same collection data as for
preceding; 21–30 May 2012; BEAT • 2 ♀♀, 2 ♂♂; same collection data as for preceding; LUP • 1 ♀;
same collection data as for preceding; AR_032; GenBank: ON619654 • 1 ♀, 1 ♂; same collection data
as for preceding; 4–14 Oct. 2012; LUP • 4 ♂♂; same collection data as for preceding; Atotonilco, Plot
II; 19.29483° N, 97.2045° W; alt. 2388 m; 15–24 Feb. 2013; Aracnolab team leg.; oak forest fragment;
BEAT • 4 ♀♀, 12 ♂♂; same collection data as for preceding; LUP • 1 ♀; same collection data as for
preceding; ANYM012 • 1 ♀, 4 ♂♂; same collection data as for preceding; ANYM013 • 2 ♂♂; same
collection data as for preceding; ANYM014 • 2 ♂♂; same collection data as for preceding; ANYM015
• 4 ♂♂; same collection data as for preceding; ANYM014; 21–30 May 2012; BEAT • 7 ♀♀, 1 ♂; same
collection data as for preceding; LUP • 1 ♂; same collection data as for preceding; 4–14 Oct. 2012; LUP.
Description
Female
Total length 8.8. Carapace dark yellow, hirsute, with darker areas delineating cephalic area, around
thoracic groove, ocular quadrangle, and clypeus (Fig. 25A, D). Sternum yellow, darker at margins,
intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide. Endites yellow,
rectangular, slightly broader at tip (Fig. 25C). Abdomen dorsum white, hirsute, covered with brown
patches, lateral sides with same brown patches diffusing ventrally, ventral surface with dark longitudinal
line at center, tracheal spiracle in center of abdomen (Fig. 25A–C). Chelicerae dark brown (Fig. 25D),
promargin with four teeth, retromargin with eight to nine denticles. Leg coloration: dark yellow with
brown patches irregularly distributed from distal end of femora to tarsi, tibia IV with two dark bands
at opposite ends (Fig. 25A). Epigynum hood absent, lateral borders sinuous. Copulatory openings
European Journal of Taxonomy 865: 1–94 (2023)
44
on anterior internal margin of heart-shaped atrium (Fig. 26E–F). Copulatory ducts entering lateral
margins of spermathecae. Seminal receptacles cylindrical, projected dorsally and located at junction
between spermathecae and copulatory duct entrance. Fertilization ducts attached to posterior margin
of spermathecae (Fig. 27G–H). Cephalothorax length 3.8, thoracic width 3.07, cephalic width 2.39.
Clypeus height 0.13. Eye diameters: AME 0.13, ALE 0.18, PME 0.17, PLE 0.2. Eye interdistances:
AME–AME 0.04, AME–ALE 0.11, ALE–PLE 0.09, PME–PME 0.2, PME–PLE 0.12. Femur lengths:
I 4.1, II 4.0, III 3.17, IV 4.0. Leg spination: femur I d1-1-1, p0-0-2, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1.
Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-2, r0-1-1. Tibia II v2-2-0. p1-1-1, r1-1-1.
Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p2-1-0, r1-1-1.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-1-1, r0-1-1. Tibia IV v2-2-2, p2-1-0, r1-1-1.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Fig. 25. Anyphaena dulceae sp. nov. A–D. Paratype, ♀ (CNAN-T01553). E–F. Paratype, ♂
(CNAN-T01551). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view.
E. Prosoma, anterior view. F. Prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
45
Male
Total length 8.2. Cephalothorax coloration as in female except yellow chelicerae (Fig. 25E). Spurs of
coxa II to IV large, covered with scattered setae (Figs 25F, 27E–F). Abdomen pattern as in female but
paler, dark longitudinal line absent. Pedipalp ventral tegular projection in prolateral view without dorsal
keel (Figs 26B, 27B). Embolus liform and translucent (Fig. 27A). RTA anterior branch semicircular
with sclerotized edge and translucent center, RTA posterior branch sclerotized and hook-shaped with
translucent tip (Figs 26 A, C–D, F, 27A, C–D). Prolateral apophysis small and cone-shaped (Fig. 27B).
Pedipalp tibia slightly longer than wide. Ventral branch of median tibial apophysis displaced towards
proximal border of tibia (Figs 26 A, D, 27A, D). Cephalothorax length 3.32, thoracic width 2.68,
cephalic width 1.80. Clypeus height 0.1. Eye diameters: AME 0.11, ALE 0.18, PME 0.15, PLE 0.15.
Eye interdistances: AME–AME 0.06, AME–ALE 0.04, ALE–PLE 0.07, PME–PME 0.15, PME–PLE
0.11. Femur lengths: I 3.9, II 3.61, III 2.88, IV 3.8. Leg spination: as in female except: femur I p0-1-2.
Fig. 26. Anyphaena dulceae sp. nov. A–D. Paratype, ♂ (CNAN-T01551). E–F. Paratype, ♀
(CNAN-T01552). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
46
Fig. 27. Anyphaena dulceae sp. nov. A–F. Paratype, ♂ (CNAN-T01551). G–H. Paratype, ♀
(CNAN-T01552). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
47
Variation
Females (N = 10): total length 8.16 (± 0.97), cephalothorax length 3.49 (± 0.27), thoracic width 2.69
(± 0.32), cephalic width 2.17 (± 0.27), femur I 3.8 (± 0.28). Males (N = 10): total length 7.47 (± 0.39),
cephalothorax length 3.3 (± 0.13), thoracic width 2.64 (± 0.12), cephalic width 1.79 (± 0.08), femur I
3.97 (± 0.18).
Distribution
This species is found in oak and oak-pine forests around Pico de Orizaba Volcano National Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray. This species
is present year-round.
Anyphaena rebecae sp. nov.
urn:lsid:zoobank.org:act:DA4ABB40-A590-44F2-8591-60B8F70C5CF8
Figs 2C–D, 28–30, 52
Differential diagnosis
Females of A. rebecae sp. nov. are differentiated from those of all species of the pectorosa and pacica
groups by the following features: atrium trapezoidal, hood notched at the center (Figs 29E–F, 30G–H).
The female of A. franciscoi sp. nov. also has a similar trapezoidal atrium and copulatory duct paths
(Fig. 17E–F), but differs from that of A. rebecae sp. nov. by the central part of the atrium being convex
and its larger size (9.6 ± 1.1). The females of A. soae sp. nov. share with both species similar copulatory
duct paths and spermathecae proportions, but differ by the oval atrium and copulatory ducts extending
beyond the hood (Fig. 36E–F). Males can be differentiated by the presence of the following features:
median apophysis hook-shaped and with a ventral transparent lamella, ventral tegular projection distal
section widest, middle section constricted, retrolateral border with a transparent cuticular edge, prolateral
border with a basal protuberance (Figs 29A–B, D, 30 A–B, D), RTA posterior branch roughly squared
with a spine-shaped process on its anterior border (Figs 29D, F, 30D). Ventral tegular projections with
broad middle sections and translucent retrolateral edges are also present in A. zorynae and A. zuyelenae
Durán-Barrón, Pérez & Brescovit, 2016 (Durán-Barrón et al. 2016: gs 5, 9, 12, 16), A. pectorosa and
A. pacica (Platnick 1974: gs 51, 55, 59, 63), A. porta sp. nov. (Fig. 41A), and A. quadrata sp. nov.
(Fig. 35A). Median apophysis with transparent lamella is present in A. pectorosa, A. fraterna, A. lacka
and A. alachua (Platnick 1974: gs 51–58). Similar RTA are present in A. epicardia sp. nov. (Fig. 24D).
Etymology
The species epithet is dedicated to Maria Rebeca Quiroz, mother of the rst author.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m;
19–27 Apr. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; LUP; CNAN-T01536.
Allotype
MEXICO • ♂; same collection data as for holotype; CNAN-T01518.
European Journal of Taxonomy 865: 1–94 (2023)
48
Paratypes
MEXICO • 3 ♂♂; same collection data as for holotype; CNAN-T01567 • 1 ♀; same collection data as
for holotype; CNAN-T01569 • 1 ♂ same collection data as for holotype; Atotonilco, Plot I; 19.12569° N,
97.06756° W; alt. 2300 m; BEAT; CNAN-T01568.
Additional material
MEXICO • 2 ♂♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
19–27 Apr. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT • 1 ♀; same collection
data as for preceding; ANYM035 • 1 ♀; same collection data as for preceding; ANYM036 • 3 ♂♂;
same collection data as for preceding; LUP • 1 ♀; same collection data as for preceding; ANYM032
• 1 ♀; same collection data as for preceding; ANYM033 • 1 ♀; same collection data as for preceding;
ANYM034 • 1 ♂; same collection data as for preceding; 4–17 Feb. 2014; BEAT • 1 ♂; same collection
data as for preceding; LUP • 1 ♀; same collection data as for holotype; BEAT • 1 ♀; same collection
Fig. 28. Anyphaena rebecae sp. nov. A–D. Holotype, ♀ (CNAN-T01536). E–F. Paratype, ♂
(CNAN-T01568). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view.
E. Prosoma, anterior view. F. Prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
49
data as for preceding; ANYM031 • 2 ♂♂; same collection data as for preceding; ANYM040 • 1 ♂;
same collection data as for preceding; ANYM043 • 4 ♂♂; same collection data as for preceding; LUP
• 1 ♀; same collection data as for preceding; ANYM037 • 1 ♀; same collection data as for preceding;
ANYM038 • 2 ♂♂; same collection data as for preceding; ANYM041 • 1 ♂; same collection data as for
preceding; 4–17 Feb. 2014; BEAT.
Description
Female
Total length 7.8. Carapace light yellow, with two darker bands over cephalic area, around fovea
and clypeus, lateral sides without pattern (Fig. 28A, D). Sternum surface white, intercoxal triangles
present on all legs. Labium brown, white at tip, longer than wide. Endites white, rectangular, broader
at tip (Fig. 28C). Chelicerae brown without dorsal pattern (Fig. 28B, D), promargin with four teeth,
Fig. 29. Anyphaena rebecae sp. nov. A–D. Paratype, ♂ (CNAN-T01568). E. Holotype, ♀
(CNAN-T01536). F. Paratype, ♀ (CNAN-T01569). A. Pedipalp, ventral view. B. Pedipalp, prolateral
view. C. Pedipalp, dorsal view. D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum,
dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
50
Fig. 30. Anyphaena rebecae sp. nov. A–F. Paratype, ♂ (CNAN-T01568). G –H. Paratype, ♀
(CNAN-T01569). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
51
retromargin with eight to nine. Legs femora base yellow and covered with darker patches from patella
to tarsi. Abdomen dorsum white, hirsute, covered with brown patches, lateral sides with same brown
patches diffusing ventrally, ventral surface white, tracheal spiracle in center of abdomen (Fig. 28A, C).
Epigynal plate anterior area delineated laterally by two faint pits. Atrium trapezoidal. Genital openings
at both lateral sides under hood. Copulatory duct slightly curved, with seminal receptacles at half their
length, and entering surface of anterior spermathecae. Fertilization ducts short, comma-shaped, and
entering lateral surface of spermathecae (Figs 29E–F, 30G–H). Cephalothorax length 3.7, thoracic width
3.24, cephalic width 2.34. Clypeus height 0.13. Eye diameters: AME 0.1, ALE 0.15, PME 0.13, PLE
0.13. Eye interdistances: AME–AME 0.06, AME–ALE 0.05, ALE–PLE 0.09, PME–PME 0.16, PME–
PLE 0.13. Femur lengths: I 3.51, II 3.27, III 2.54, IV 3.32. Leg spination: femur I d1-1-1, p0-1-1, r0-1-1.
Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia
II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia
III v2-2-2, p0-1-1, r0-1-1. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-1-1, r0-1-1. Tibia
IV v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Total length 6.3. Cephalothorax and abdomen coloration as in female, except yellow chelicerae (Fig. 28E).
Coxa II tubercle bifurcated. Coxa III tubercle roughly rectangular with median spine. Coxa IV spur bent
and located at retrolateral basal corner (Figs 28F, 30E–F). Embolus liform and translucent. Palpal tibia
prolateral apophysis present (Figs 29E–F, 30B–C). Pedipalp tibia slightly longer than wide. Ventral
branch of median tibial apophysis displaced towards proximal border of tibia (Figs 29A, D, 30A, D).
Cephalothorax length 2.83, thoracic width 2.2, cephalic width 1.0. Clypeus height 0.07. Eye diameters:
AME 0.11, ALE 0.15, PME 0.15, PLE 0.15. Eye interdistances: AME–AME 0.04, AME–ALE 0.04,
ALE–PLE 0.04, PME–PME 0.17, PME–PLE 0.09. Femur lengths: I 3.41, II 3.12, III 2.39, IV 3.17. Leg
spination as in female except: femur I r0-2-1.
Variation
Females (N = 10): total length 7.23 (± 1.01), cephalothorax length 2.97 (± 0.2), thoracic width 2.35
(± 0.3), cephalic width 1.19 (± 0.08), femur I 3.28 (± 0.15). Males (N = 10): total length 6.49 (± 0.23),
cephalothorax length 2.85 (± 0.12), thoracic width 2.27 (± 0.11), cephalic width 0.99 (± 0.03), femur I 3.44
(± 0.18).
Distribution
This species is found in oak and tropical wet forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray. This species
is present year-round.
Anyphaena miniducta sp. nov.
urn:lsid:zoobank.org:act:A95B18CA-F021-4829-9F50-4EE5E14AFA9F
Figs 31–33, 52
Differential diagnosis
Females of A. miniducta sp. nov. are differentiated from most of those of the pectorosa and pacica
groups by the following features: atrium T-shaped, lateral borders short and parallel, copulatory ducts
short, slightly curved (Figs 32E–F, 33G–H). Similar atria are present in females of A. simplex (Pickard-
Cambridge O. 1894: pl. 18 g. 3a) and A. porta sp. nov. (Fig. 39E), but they differ by the atrium
European Journal of Taxonomy 865: 1–94 (2023)
52
proportions and the length of the lateral sides, respectively. Males can be differentiated by the presence
of the following features: median apophysis hook-shaped, transparent lamellae absent, middle section
of ventral tegular projection broad, retrolateral border with a narrow cuticular transparent edge, distal
section pointed, RTA anterior branch ear-shaped, larger than the lamella-shaped posterior branch in
retrolateral view (Figs 32A–D, 33A–D).
Etymology
The species epithet refers to the short copulatory ducts of this species in relation to other members of
the pectorosa group.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m;
19–27 Apr. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; LUP; CNAN-T01531.
Allotype
MEXICO • ♂; same collection data as for holotype; BEAT; CNAN-T01515.
Paratypes
MEXICO • 1 ♂, 1 ♀; same collection data as for holotype; CNAN-T01561 • 1 ♀; same collection data
as for holotype; CNAN-T01560.
Additional material
MEXICO • 1 ♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
19–27 Apr. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT; ANYM047 • 1 ♂;
same collection data as for holotype; BEAT; ANYM048 • 1 ♀; same collection data as for preceding;
ANYM049 • 1 ♀; same collection data as for preceding; CRP; ANYM050 • 1 ♀; same collection data
as for preceding; LUP; ANYM051.
Description
Female
Total length 9.0. Carapace light yellow, with two darker and faint bands over cephalic area, around
fovea and clypeus (Fig. 31A, D). Sternum surface white, intercoxal triangles present on all legs. Labium
brown, white at tip, longer than wide. Endites white, rectangular, broader at tip (Fig. 31C). Chelicerae
brown without dorsal pattern (Fig. 31B, D), promargin with ve, retromargin with eight to nine denticles.
Abdomen dorsum light yellow, hirsute, delineated by dark yellow longitudinal bands, central anterior
half white, lateral sides light yellow, darker band only at top, ventral surface white, tracheal spiracle
closer to epigastric furrow (Fig. 31A, C). Copulatory duct slightly curved and entering lateral surface
of spermathecae. Seminal receptacles above genital openings. Fertilization ducts short, comma-shaped,
and entering lateral surface of spermathecae (Figs 32E–F, 33G–H). Cephalothorax length 3.76, thoracic
width 2.83, cephalic width 1.52. Clypeus height 0.15. Eye diameters: AME 0.11, ALE 0.15, PME 0.15,
PLE 0.15. Eye interdistances: AME–AME 0.06, AME–ALE 0.05, ALE–PLE 0.11, PME–PME 0.2,
PME–PLE 0.1. Femur lengths: I 4.39, II 4.05, III 3.17, IV 4.59. Leg spination: femur I d1-1-1, p0-2-1,
r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-
2. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1.
Tibia III v2-2-2, p0-1-1, r0-1-1. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV 1-1-1, p0-0-1, r0-0-1.
Tibia IV v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
53
Male
Total length 8.0. Cephalothorax and abdomen pattern as in female, except darker and yellow chelicerae
(Fig. 31E). Coxa II ventral surface smooth. Coxa III tubercle bifurcated, with posterior bent spine and
anterior tubercle. Coxa IV spur bent and with broad base (Figs 31F, 33E–F). Embolus liform and
translucent. Pedipalp tibia longer than wide. RTA posterior branch short and shaped as bent lamella.
Prolateral apophysis of palpal tibia present. Ventral branch of median tibial apophysis present
(Figs 32A–D, 33A–D). Cephalothorax length 3.41, thoracic width 2.59, cephalic width 1.19. Clypeus
height 0.12. Eye diameters: AME 0.09, ALE 0.12, PME 0.12, PLE 0.15. Eye interdistances: AME–AME
0.05, AME–ALE 0.04, ALE–PLE 0.11, PME–PME 0.15, PME–PLE 0.12. Femur lengths: I 4.63, II 4.1,
III 3.17, IV 4.63. Leg spination as in female except: femur I r0-2-1. Femur II p0-1-2.
Variation
Females (N = 6): total length 8.9 (± 0.39), cephalothorax length 3.8 (± 0.11), thoracic width 2.89 (± 0.06),
cephalic width 1.52 (± 0.06), femur I 4.41 (± 0.04). Males (N = 4): total length 8.03 (± 0.37), cephalothorax
length 3.44 (± 0.21), thoracic width 2.63 (± 0.12), cephalic width 1.21 (± 0.07), femur I 4.61 (± 0.13).
Fig. 31. Anyphaena miniducta sp. nov. A–D. Paratype, ♀ (CNAN-T01560). E–F. Allotype, ♂
(CNAN-T01515). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view.
E. Prosoma, anterior view. F. Prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
European Journal of Taxonomy 865: 1–94 (2023)
54
Distribution
This species is found in oak and tropical wet forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray. This species
is present year-round.
Fig. 32. Anyphaena miniducta sp. nov. A–D. Allotype, ♂ (CNAN-T01515). E–F. Paratypes, ♀♀
(CNAN-T01560, CNAN-T01561). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp,
dorsal view. D. Pedipalp, retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale
bars: 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
55
Fig. 33. Anyphaena miniducta sp. nov. A–F. Allotype, ♂ (CNAN-T01515).G–H. Paratype, ♀
(CNAN-T01561). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
56
Anyphaena quadrata sp. nov.
urn:lsid:zoobank.org:act:DE0EFD57-6E7A-4854-9B6D-2C919B2AF7D7
Figs 34–35, 52
Differential diagnosis
Males of A. quadrata sp. nov. are differentiated from all those of the pectorosa and pacica groups
by the ventral tegular projection distal section being broad and rectangular, the median apophysis also
rectangular, longer than wide (Figs 34E–F, 35A–D). Coxa III tubercle bifurcated, anterior ramication
blunt, posterior ramication spine-shaped (Fig. 35E–F). Similar tegular apophyses are found in males of
A. zuyelenae (Durán-Barrón et al. 2016: gs 12–14, 16) and A. accentuata (Platnick 1974: gs 63, 70;
Dondale & Redner 1982: g. 336; Brescovit 1997: gs 1–6), but they differ by the hook-shaped median
apophysis, the RTA shape and coxal ventral surfaces.
Fig. 34. Anyphaena quadrata sp. nov., paratype, ♂ (CNAN-T01566). A. Dorsal habitus. B. Lateral
habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Pedipalp, ventral view. F. Pedipalp, retrolateral
view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
57
Etymology
The species epithet is derived from the Latin ‘quadrata’ (‘square’), referring to the distinctive broad
semi-squared shape of the ventral tegular projection.
Material examined
Holotype
MEXICO • ♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
2–11 Oct. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; LUP; CNAN-T01535.
Paratype
MEXICO • 1 ♂; Veracruz, Calcahualco, Xamaticpac, Plot II 19.12614° N, 97.06708° W; alt. 1700 m;
2–11 Oct. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT; CNAN-T01566.
Fig. 35. Anyphaena quadrata sp. nov., paratype, ♂ (CNAN-T01566). A. Pedipalp, ventral view.
B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp, retrolateral view. E. Coxae, ventral
view. F. Coxae, lateral view. Scale bars: A–D = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
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Additional material
MEXICO • 1 ♂; same collection data as for paratype; BEAT; ANYM054 • 1 ♂; same collection data as
for preceding; ANYM055.
Description
Male
Total length 4.5. Carapace yellow, with two darker bands over ocular quadrangle, around fovea and
clypeus. Sternum surface white, intercoxal triangles present on all legs. Labium yellow, white at tip,
longer than wide. Endites white, rectangular, broader at tip (Fig. 34A–C). Chelicerae yellow, paturon
without pattern (Fig. 34D), promargin with four teeth, retromargin with eight to nine denticles. Abdomen
hirsute, dorsal surface dark yellow, anterior border white, lateral sides with same dark pattern on top
and diffusing ventrally, ventral surface white. Tracheal spiracle at center of abdomen (Fig. 34A, C).
Distal surface of median apophysis with spine. Prolateral apophysis present. Tibia slightly longer than
wide. Ventral branch of median tibial apophysis displaced towards proximal border of tibia (Figs 34E–F,
35A–B, D). Coxa II with short basal tubercle, coxa IV with bent spur (Fig. 35F-F). Cephalothorax length
2.24, thoracic width 1.76, cephalic width 0.77. Clypeus height 0.07. Eye diameters: AME 0.06, ALE
0.1, PME 0.1, PLE 0.11. Eye interdistances: AME–AME 0.04, AME–ALE 0.02, ALE–PLE 0.09, PME–
PME 0.15, PME–PLE 0.07. Femur lengths: I 2.29, II 2.05, III 1.71, IV 2.2. Leg spination: femur I d1-
1-1, p0-2-1, r0-0-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1,
p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1,
p0-1-1, r0-1-1. Tibia III v2-2-2, p0-1-1, r0-1-1. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1,
p0-0-1, r0-0-1. Tibia IV v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Female
Unknown.
Variation
Males (N = 4): total length 4.63 (± 0.15), cephalothorax length 2.24 (± 0.04), thoracic width 1.76 (± 0.04),
cephalic width 0.81 (± 0.04), femur I 2.37 (± 0.05).
Distribution
Only known from the type locality (Fig. 52).
Natural history
Most specimens collected by beating and direct searching over vegetation.
Anyphaena soae sp. nov.
urn:lsid:zoobank.org:act:8A24E970-6D83-475F-BD2C-9CD4C599C653
Figs 36, 38A–B, 52
Differential diagnosis
Females of A. soae sp. nov. are differentiated from those of all species of the pectorosa and pacica
groups by the following features: copulatory ducts extending beyond the hood, atrium oval, hood faint,
thin and straight (Figs 36E–F, 38A–B).
Etymology
The species epithet is dedicated to Soa Rivera, niece of the rst author.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
59
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot II; 19.29483° N, 97.2045° W; alt. 2388 m; 15–24
Feb. 2013; Arcanolab team leg.; oak and pine forest fragment; BEAT; AR_060; GenBank: ON619648;
CNAN-T01538.
Paratypes
MEXICO • 1 ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m;
21–30 May 2012; Aracnolab team leg.; oak forest fragment; LUP; AR_057; GenBank: ON619646;
CNAN-T01572 • 1 ♀; same collection data as for preceding; AR_059; GenBank: ON619647;
CNAN-T01573.
Fig. 36. Anyphaena soae sp. nov., paratypes, ♀♀ (CNAN-T01572, CNAN-T01573). A. Dorsal
habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view.
F. Epigynum, dorsal view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
60
Additional material
MEXICO • 1 ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m;
15–24 Feb. 2013; Aracnolab team leg.; oak forest fragment; BEAT • 1 ♀; same collection data as for
preceding; LUP • 1 ♀; same collection data as for preceding; 21–30 May 2012; LUP • 1 ♀; same
collection data as for preceding; ANYM025 • 1 ♀; same collection data as for preceding; Atotonilco.
Plot II; 19.29483° N, 97.2045° W; alt. 2388 m; 15–24 Feb. 2013; oak and pine forest fragment; BEAT •
1 ♀; same collection data as for preceding; CRP • 3 ♀♀; same collection data as for preceding; LUP •
1 ♀; same collection data as for preceding; ANYM029 • 1 ♀; same collection data as for preceding;
21–30 May 2012; BEAT; ANYM026 • 1 ♀; same collection data as for preceding; ANYM027 • 1 ♀;
same collection data as for preceding; ANYM028 • 1 ♀; same collection data as for preceding; LUP •
1 ♀; same collection data as for preceding; ANYM030 • 1 ♀; same collection data as for preceding;
4–14 Oct. 2012; LUP.
Description
Female
Total length 7.8. Carapace yellow, pattern with darker bands delineating cephalic area, around fovea
and clypeus (Fig. 36A, D). Sternum surface white, intercoxal triangles present on all legs. Labium
brown, white at tip, longer than wide. Endites yellow, rectangular, broader at tip (Fig. 36C). Chelicerae
brown, without dorsal pattern (Fig. 36B, D), promargin with four teeth, retromargin with eight to nine
denticles. Abdomen dorsal surface white, covered with darker patches and some diffuse chevrons at
center, lateral and ventral surfaces white, pattern diffusing ventrally, tracheal spiracle slightly closer to
epigastric furrow (Fig. 36A–C). Anterior area of epigynal plate delineated laterally by two faint pits near
copulatory ducts bent. Genital openings at both sides of hood. Copulatory duct paths bent posteriorly
beyond hood and entering spermathecae antero-laterally. Seminal receptacles closer to spermathecae
at one third of copulatory ducts’ length. Fertilization ducts short, straight, and entering spermathecae
via postero-lateral surface (Figs 36E–F, 38A–B). Cephalothorax length 2.93, thoracic width 2.29,
cephalic width 1.23. Clypeus height 0.1. Eye diameters: AME 0.09, ALE 0.12, PME 0.12, PLE 0.12.
Eye interdistances: AME–AME 0.06, AME–ALE 0.04, ALE–PLE 0.17, PME–PME 0.17, PME–PLE
0.13. Femur lengths: I 3.13, II 2.9, III 2.26, IV 3.0. Leg spination: femur I d1-1-1, p0-1-2, r0-1-1. Tibia
I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II
v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III
v2-2-2, p0-1-1, r0-1-1. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-1-1, r0-0-1. Tibia IV
v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2, p1-1-2, r1-1-2
Male
Unknown.
Variation
Females (N = 10): total length 8.03 (± 0.62), cephalothorax length 3.41 (± 0.3), thoracic width 2.48
(± 0.19), cephalic width 1.27 (± 0.08), femur I 3.46 (± 0.27).
Distribution
Only known from the type locality (Fig. 52).
Natural history
Most specimens collected by beating and direct searching over vegetation.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
61
Anyphaena noctua sp. nov.
urn:lsid:zoobank.org:act:1C0E8CF3-254F-4193-B505-8272507357B1
Figs 37, 38C–D, 52
Differential diagnosis
Females of A. noctua sp. nov. are differentiated from those of all species of the pectorosa and pacica
groups by the following features: atrium small, lateral borders sinuous, hood triangular, posteriorly
curved at the center, genital openings at the posterior margin of the atrium, copulatory ducts short,
comma-shaped (Figs 37E–F, 38C–D).
Etymology
The species epithet is derived from the Latin ‘noctua’ (‘owl’), in reference to the characteristic shape of
the epigynum and atrium that are reminiscent of a horned owl with the spermathecae as its eyes.
Fig. 37. Anyphaena noctua sp. nov., holotype, ♀ (CNAN-T01533). A. Dorsal habitus. B. Lateral habitus.
C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view. F. Epigynum, dorsal view.
Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
62
Material examined
Holotype
MEXICO • ♀; San Luis Potosi, Xilitla City, Las Pozas; 21.39722° N, 98.99388° W; alt. 662 m; 10–15
Jun. 2012; Arcanolab team leg.; tropical wet forest fragment; LUP; CNAN-T01533.
Description
Female
Total length 6.1. Carapace light yellow, with two faint darker bands delineating cephalic area and around
fovea, ocular quadrangle and clypeus dark brown (Fig. 37A, D). Sternum surface white, intercoxal
triangles present on all legs. Labium yellow, white at tip, longer than wide. Endites yellow, rectangular,
broader at tip (Fig. 37C). Chelicerae brown without dorsal pattern (Fig. 37B, D), promargin with four
Fig. 38. Anyphaena spp. A–B. A. soae sp. nov., paratype, ♀ (CNAN-T01572). A. Epigynum, ventral
view. B. Epigynum, dorsal view. – C–D. A. noctua sp. nov., holotype, ♀ (CNAN-T01533). C. Epigynum,
ventral view. D. Epigynum, dorsal view. Scale bars: A– B = 0.2 mm: C–D = 0.1 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
63
teeth, retromargin with seven to eight denticles. Femora base of legs white, orange-yellow distally,
darker from patella to tarsi. Abdomen dorsal surface white and covered by diffuse darker pattern, pattern
of lateral and ventral surfaces even fainter, tracheal spiracle closer to epigastric furrow (Fig. 37A–C).
Lateral side of espigynal plate anked by two faint pits. Seminal receptacles close to genital openings.
Fertilization ducts short, cylindrical, and entering posterior surface of spermathecae (Figs 37E–F,
38C–D). Cephalothorax length 2.71, thoracic width 2.1, cephalic width 1.1. Clypeus height 0.05. Eye
diameters: AME 0.07, ALE 0.11, PME 0.1, PLE 0.11. Eye interdistances: AME–AME 0.04, AME–ALE
0.02, ALE–PLE 0.05, PME–PME 0.17, PME–PLE 0.07. Femur lengths: I 2.86, II 2.62, III 2.19, IV
2.95. Leg spination: femur I d1-1-1, p0-0-2, r0-0-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0,
p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-0-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0,
p1-1-1, r1-1-1. Femur III d1-1-1, p0-0-1, r0-0-1. Tibia III v1-1-0, p1-1-1, r1-1-1. Metatarsus III v2-2-2,
p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-1, r1-1-1. Metatarsus IV v2-2-2,
p1-1-2, r1-1-2.
Male
Unknown.
Variation
Only type specimen known.
Distribution
Only known from the type locality (Fig. 52).
Natural history
Collected by beating direct searching over ground vegetation.
Porta group
Diagnosis
The porta group can be separated from other Anyphaena species groups by the following characters.
Males
Coxae III and IV smooth or sometimes with ventral tubercles (Figs 41E–F, 51E–F). The general shape of
the male palp is similar to that of the pectorosa and pacica groups. Differs from both of them by having
relatively longer palpal tibiae and in the general shape of the RTA. The RTA base is usually restricted to
the distal portion of the tibia instead of the medial portion (compare Figs 48E, 49A with Figs 23A, 24A).
Female
Atrium considerably longer than wide. Copulatory openings sometimes covered by the lateral borders
of the atrium. Copulatory ducts long and parallel to each other, projecting posteriorly beyond the hood
margin (A. porta sp. nov., A.ibarrai sp. nov., A. salgueiroi sp. nov.), although two species (A. tonoi
sp. nov. and A. alachua) have shorter copulatory ducts (Fig. 46F, see A. alachua in Supp. le 3).
Spermatheca well sclerotized, oval to spherical.
Anyphaena porta sp. nov.
urn:lsid:zoobank.org:act:F69F982D-2304-4E29-B8F0-F9E66CF24314
Figs 39–41, 52
Differential diagnosis
Females of A. porta sp. nov. can be differentiated from females of all described species of Anyphaena
of the pacica and pectorosa groups by the lateral borders of the epigynum being long and parallel,
European Journal of Taxonomy 865: 1–94 (2023)
64
simulating a pair of lengthy doors slightly open with the hood as the door upper frame. Copulatory ducts
weakly sclerotized, path bent 180° at the hood, making two loops (Figs 39E–F, 40E–F, 41G–H). Similar
epigynum shapes are shared with A. salgueiroi sp. nov. and A. ibarrai sp. nov., but differ, respectively,
from A. porta sp. nov. by the almost closed atrium borders (Fig. 42E) and the copulatory ducts being
heavily sclerotized (Fig. 43F). Males have a broad ventral tegular projection with the prolateral edge
enlarged, a retrolateral rectangular transparent edge, and two small apical apophyses (Figs 40A, 41A).
Distal edge of median apophysis bifurcated, ventral extension the largest and hook-shaped in ventral
view (Figs 40A, 41A), dorsal extension translucent and hook-shaped in retrolateral view (Figs 40D,
41D). RTA anterior branch acute, RTA posterior branch tip C-shaped (Figs 40B–D, 41B–D). Coxae II
to IV with broad and at ventral tubercles (Fig. 41E–F). Similar broad ventral tegular projections are
found in A. pectorosa (Platnick 1974: gs 51, 55, 59; Dondale & Redner 1982: g. 327), A. bermudensis
Sierwald, 1988 (Sierwald 1988: g. 1) and A. zorynae Durán-Barrón, Pérez & Brescovit, 2016 (Durán-
Fig. 39. Anyphaena porta sp. nov. A–C. Holotype, ♀ (CNAN-T01534). E–F. Paratype, ♀
(CNAN-T01564). A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view.
E. Epigynum ventral. F. Epigynum ventral cleared. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
65
Barrón et al. 2016: gs 5–6, 9), these last two species with coxae unarmed and with small tubercles,
respectively.
Etymology
The species epithet refers to the Spanish words ‘portón’ or ‘puerta’, referring to the door-shaped
epigynum atrium, a feature diagnostic of this species.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot II; 19.29483° N, 97.2045° W; alt. 2388 m; 15–24
Feb. 2013; Arcanolab team leg.; oak and pine forest fragment; CRP; CNAN-T01534.
Fig. 40. Anyphaena porta sp. nov. A–D. Paratype, ♂ (CNAN-T01563). E–F. Paratype, ♀ (CNAN-
T01565). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp,
retrolateral view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
66
Allotype
MEXICO • ♂; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m; 21–
30 May 2012; oak forest fragment; LUP; AR_066; GenBank: ON619626; CNAN-T01517.
Paratypes
MEXICO • 1 ♀; same collection data as for holotype; BEAT; AR_077; GenBank: ON619656;
CNAN-T01565 • 1 ♂; same collection data as for holotype; 21–30 May 2012; AR_065; GenBank:
ON619625; CNAN-T01563 • 1 ♀; same collection data as for preceding; BEAT; AR_027; GenBank:
ON619643; CNAN-T01564.
Additional material
MEXICO • 8 ♀♀; 6 ♂♂; same collection data as for allotype; 15–24 Feb. 2013; BEAT • 1 ♀; same
collection data as for preceding; AR_029; GenBank: ON619644 • 1 ♀; same collection data as for
preceding; CRP • 8 ♀♀, 7 ♂♂; same collection data as for preceding; LUP • 1 ♀; same collection data as
for preceding; AR_030; GenBank: ON619645 • 1 ♀, 1 ♂; same collection data as for preceding; 21–30
May 2012; BEAT • 2 ♂♂; same collection data as for preceding; LUP • 1 ♀; same collection data as for
preceding; 4–14 Oct. 2012; BEAT • 17 ♀♀, 9 ♂♂; same collection data as for holotype; BEAT • 1 ♂;
same collection data as for preceding; AR_067; GenBank: ON619627 • 1 ♀; same collection data as for
preceding; AR_078; GenBank: ON619657 • 1 ♀; same collection data as for preceding; BERL • 5 ♂♂;
same collection data as for preceding; CRP • 42 ♀♀, 37 ♂♂; same collection data as for preceding •
2 ♂♂; same collection data as for preceding; ANYM016 • 2 ♂♂; same collection data as for preceding;
ANYM017 • 3 ♂♂; same collection data as for preceding; ANYM018 • 1 ♂; same collection data as for
preceding; PF • 8 ♀♀, 3 ♂♂; same collection data as for preceding; 21–30 May 2012; BEAT • 1 ♂; same
collection data as for preceding; BERL • 1 ♀; same collection data as for preceding; CRP • 7 ♀♀, 3 ♂♂;
same collection data as for preceding; LUP • 1 ♀; same collection data as for preceding; 4–14 Oct. 2012;
BEAT • 1 ♀; same collection data as for preceding; LUP • 1 ♀; Calcahualco, Atotonilco, Xamaticpac,
Plot I; 19.14172° N, 97.20597° W; alt. 1710 m; 19–27 Apr. 2013; Aracnolab team leg.; oak and tropical
wet forest fragment; LUP; AR_014; GenBank: ON619664.
Description
Female
Total length 5.6. Carapace yellow, pattern with darker longitudinal bands around cephalic area and fovea
(Fig. 39A, D). Sternum surface white, intercoxal triangles present on all legs. Labium yellow, white at
tip, longer than wide. Endites white, rectangular, slightly broader at tip (Fig. 39C). Chelicerae yellow
(Fig. 39B, D), promargin with four teeth, retromargin with seven or eight denticles. Leg coloration:
light yellow, slightly darker on metatarsi and tarsi (Fig. 39A–C). Abdomen white, hirsute, dorsal surface
with scattered brown patches, lateral and ventral surfaces with same pattern but faded ventrally, tracheal
spiracle close to epigastric furrow (Fig. 39A–C). Epigynum atrium rectangular, longer than wide, door-
shaped. Copulatory openings inside middle section of lateral borders of atrium. Copulatory ducts coiled
around seminal receptacles, entering spermathecae on ventral surface at junction between long seminal
receptacles and fertilization duct origin. Fertilization ducts short, cylindrical (Figs 39E–F, 40E–F, 41G–
H). Cephalothorax length 2.15, thoracic width 1.56, cephalic width 0.81. Clypeus height 0.09. Eye
diameters: AME 0.07, ALE 0.1, PME 0.09, PLE 0.09. Eye interdistances: AME–AME 0.05, AME–ALE
0.04, ALE–PLE 0.1, PME–PME 0.12, PME–PLE 0.09. Femur lengths: I 1.84, II 1.68, III 1.29, IV 1.87.
Leg spination: femur I d1-1-1, p0-0-2, r0-0-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-
1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-0-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1,
r1-1-0. Femur III d1-1-1, p0-1-1, r0-0-1. Tibia III v2-2-2, p1-1-0, r1-1-0. Metatarsus III v2-2-2, p1-1-2,
r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-0, p1-1-1, r1-1-1. Metatarsus IV v2-2-2, p1-1-2,
r1-1-2.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
67
Fig. 41. Anyphaena porta sp. nov. A–F. Paratype, ♂ (CNAN-T01563). G–H. Paratype, ♀
(CNAN-T01565). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
European Journal of Taxonomy 865: 1–94 (2023)
68
Male
Total length 5.3. Cephalothorax and abdomen coloration as in female. Embolus liform and translucent.
Prolateral apophysis of palpal tibia present (Figs 40B–C, 41B–C). Pedipalp tibia longer than wide.
Ventral branch of median tibial apophysis displaced towards proximal border of tibia (Figs 40A, C–D,
41A, C–D). Cephalothorax length 2.39, thoracic width 1.61, cephalic width 0.84. Clypeus height 0.07.
Eye diameters: AME 0.09, ALE 0.1, PME 0.1, PLE 0.1. Eye interdistances: AME–AME 0.05, AME–
ALE 0.09, ALE–PLE 0.06, PME–PME 0.09, PME–PLE 0.07. Femur lengths: I 2.54, II 2.44, III 2.1, IV
2.49. Legs as in female except: femur II p0-0-2. Metatarsus II r1-1-1. Femur III p0-0-1. Tibia III v1-1-0,
p0-1-1, r0-1-1. Tibia IV v0-1-0.
Variations
The anterior curvature of the female copulatory ducts and the seminal receptacle length vary among
specimens (Figs 39E–F, 40E–F). Females (N = 10): total length 5.66 (± 0.98), cephalothorax length 2.18
(± 0.11), thoracic width 1.66 (± 0.12), cephalic width 0.9 (± 0.07), femur I 1.96 (± 0.13). Males (N = 10):
total length 5.2 (± 0.29), cephalothorax length 2.4 (± 0.13), thoracic width 1.88 (± 0.14), cephalic width
0.82 (± 0.05), femur I 2.55 (± 0.11). The anterior projections of the spermathecae and proportions of the
atrium vary in shape (Figs 39E–F, 40A-B).
Distribution
This species is found in oak and tropical wet forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray. This species
is present year-round.
Anyphaena salgueiroi sp. nov.
urn:lsid:zoobank.org:act:397A0D4B-A5D3-4B71-A19D-2A004876ED2F
Figs 2E–F, 42, 44A–B, 52
Differential diagnosis
Females of A. salgueiroi sp. nov. can be differentiated from those of A. porta sp. nov. and A. ibarrai
sp. nov. by their larger size, reddish color in alcohol (Fig. 42A–D), epigynal lateral borders almost
touching, atrium greatly reduced, and genital openings inside two large semicircular sclerotized plates
(Fig. 42E–F).
Etymology
The species epithet is dedicated to Dr Francisco Javier Salgueiro-Sepúlveda, former member of the
second author’s lab.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Atotonilco, Plot I; 19.12569° N, 97.06756° W; alt. 2300 m;
15–24 Feb. 2013; Aracnolab team leg.; oak forest fragment, CRP; AR_039; GenBank: ON619631;
CNAN-T01537.
Paratypes
MEXICO • 1 ♀; same collection data as for holotype; 21–30 May 2012; LUP; AR_040; GenBank:
ON619632; CNAN-T01571 • 1 ♀; Atotonilco, Plot II; 19.29483° N, 97.2045° W; alt. 2388 m; 21–30
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
69
May 2012; Arcanolab team leg.; oak and pine forest fragment; BEAT; AR_038; GenBank: ON619630;
CNAN-T01570.
Description
Female
Total length 8.2. Carapace orange over cephalic area with two red longitudinal bands over most of
thoracic area, lateral margins yellow, clypeus red. Sternum pale orange, darker at margins, intercoxal
triangles present on all legs. Labium red, white at tip, longer than wide. Endites pale orange, rectangular,
slightly broader at tip. Chelicerae orange, promargin with ve teeth, retromargin with six to seven
denticles. Leg coloration: yellow at base, turning red after femora to tarsi (Fig. 42A–D). Lateral sides
Fig. 42. Anyphaena salgueiroi sp. nov., holotype, ♀ (CNAN-T01537). A. Dorsal habitus. B. Lateral
habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view. F. Epigynum, dorsal
view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
70
of abdomen dorsum covered with red reticulated pattern leaving clearer central longitudinal band,
lateral sides with same pattern, less dense ventrally. Tracheal spiracle at center, anterior half of abdomen
surface almost without red pattern, posterior half dominated by reddish longitudinal lines (Fig. 42A–C).
Epigynum atrium rectangular, longer than wide, door-shaped. Hood narrow and straight. Copulatory
openings inside two semicircular sclerotized plates (Fig. 42E–F). Copulatory ducts weakly sclerotized
following external edge of semicircular plates, turning 180° beyond hood, and entering spermathecae
ventrally at junction between long seminal receptacles and fertilization duct origin. Fertilization ducts
short and curved (Figs 42E–F, 44A-B). Cephalothorax length 2.98, thoracic width 2.24, cephalic width
1.35. Clypeus height 0.11. Eye diameters: AME 0.1, ALE 0.13, PME 0.13, PLE 0.12. Eye interdistances:
AME–AME 0.06, AME–ALE 0.04, ALE–PLE 0.18, PME–PME 0.17, PME–PLE 0.11. Femur lengths: I
2.52, II 2.42, III 1.97, IV 2.58. Leg spination: femur I d1-1-1, p0-1-1, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-
1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1.
Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p1-1-1, r1-1-1.
Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v1-2-2, p1-1-1, r1-1-1.
Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Unknown.
Variation
Females (N = 3): total length 7.8 (± 0.36), cephalothorax length 2.67 (± 0.27), thoracic width 2.05
(± 0.18), cephalic width 1.25 (± 0.1), femur I 2.3 (± 0.21).
Distribution
This species is found in oak and oak with pine forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Unknown.
Anyphaena ibarrai sp. nov.
urn:lsid:zoobank.org:act:10A02CEE-A85A-4EBE-9781-82FDEF33C799
Figs 43, 44C–D, 52
Differential diagnosis
Females of A. ibarrai sp. nov. can be differentiated from those of A. porta sp. nov. and A. salgueiroi
sp. nov. by the genital openings being shaped as two large sclerotized chambers, copulatory ducts also
well sclerotized, and the atrium lateral borders diverging posteriorly (Figs 43E–F, 44C–D).
Etymology
The species epithet is dedicated to the Mexican arachnologist Dr Guillermo Ibarra.
Material examined
Holotype
MEXICO • ♀; San Luis Potosi, Xilitla City, Las Pozas; 21.39722° N, 98.99388° W; alt. 662 m; 26–30
Mar. 2012; Arcanolab team leg.; tropical wet forest fragment; LUP; AR_001; GenBank: ON619633;
CNAN-T01528.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
71
Description
Female
Total length 6.6. Carapace yellow with brown bands delineating cephalic area and around fovea,
ocular area and clypeus cuticle brown. Sternum brown at borders with longitudinal white irregular
band, intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide. Endites
dark yellow, rectangular, slightly broader at tip. Chelicerae brown, paturon dorsum covered by darker
reticulated pattern (Fig. 43A–D), promargin with four teeth, retromargin with seven to eight denticles.
Leg coloration: yellow with irregular dark band and patches throughout length. Abdomen dorsum white,
covered by gray pattern of chevrons and irregular patches, lateral and ventral surfaces covered with same
dark pattern of irregular patches, except anterior ventral half, where restricted to central band anterior
to racheal spiracle (Fig. 43A–C). Epigynum atrium rectangular, longer than wide, door-shaped. Hood
Fig. 43. Anyphaena ibarrai sp. nov., holotype, ♀ (CNAN-T01528). A. Dorsal habitus. B. Lateral habitus.
C. Ventral habitus. D. Prosoma, anterior view. E. Epigynum, ventral view. F. Epigynum, dorsal view.
Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
72
narrow and curved. Copulatory duct openings at anterior edge of two sclerotized chambers. Copulatory
ducts well sclerotized, tube-shaped, parallel and ventrally entering spermathecae at junction between
long seminal receptacles and fertilization duct origin. Fertilization ducts short and curved (Figs 43E–F,
44C–D). Cephalothorax length 2.32, thoracic width 1.87, cephalic width 1.1. Clypeus height 0.11. Eye
diameters: AME 0.1, ALE 0.15, PME 0.11, PLE 0.12. Eye interdistances: AME–AME 0.04, AME–ALE
0.04, ALE–PLE 0.07, PME–PME 0.16, PME–PLE 0.09. Femur lengths: I 2.29, II 2.1, III 1.45, IV 2.32.
Leg spination: femur I d1-1-1, p0-0-2, r0-0-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-
1, r1-1-1. Femur II d1-1-1, p0-0-1, r0-0-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1,
r1-1-1. Femur III d1-1-1, p0-0-1, r0-0-1. Tibia III v1-1-0, p1-1-1, r1-1-1. Metatarsus III v2-2-2, p1-1-2,
Fig. 44. Anyphaena spp. A–B. A. salgueiroi sp. nov., holotype, ♀ (CNAN-T01537). A. Epigynum,
ventral view. B. Epigynum, dorsal view. – C–D. A. ibarrai sp. nov., holotype, ♀ (CNAN-T01528).
C. Epigynum, ventral view. D. Epigynum, dorsal view. Scale bars: 0.2 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
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r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1. Tibia IV v2-2-2, p1-1-1, r1-1-1. Metatarsus IV v2-2-2, p1-1-2,
r1-1-2.
Male
Unknown.
Variation
Only type specimen known.
Distribution
Only known from the type locality (Fig. 52).
Natural history
Collected at night by direct searching over vegetation.
Anyphaena tonoi sp. nov.
urn:lsid:zoobank.org:act:C6FF6A9C-010D-41A0-8283-2538027BCBD5
Figs 45–47, 52
Differential diagnosis
Females of A. tonoi sp. nov. can be differentiated from those of all described species of Anyphaena of
the pacica and pectorosa groups by the opposing L-shaped copulatory ducts, and the adjacent genital
openings being inside a small longitudinal rectangular atrium (Figs 46E–F, 47G–C). Central genital
openings are also present in females of A. jimenezi sp. nov., but the latter species differs by the moustache-
shaped copulatory ducts. Males can be differentiated by the presence of the following features: distal
edge of median apophysis with two transparent lamellae and a sclerotized central hook (Figs 46D, 47D),
middle section of ventral tegular projection broad, retrolateral border with a transparent cuticular edge,
distal section shaped as a folded lamella. RTA anterior branch rectangular, long and straight, posterior
branch lamella-shaped with a middle notch (Fig. 47C–D), and coxal ventral surfaces smooth (Figs 45E–
F, 46A–D, 47A–F). Ventral tegular projection with broad middle sections and translucent retrolateral
edges are also found in males of A. zorynae (Durán-Barrón et al. 2016: gs 5–6), A. quadricornuta
Kraus, 1955 (Kraus 1955: gs 108–109), A. stigma sp. nov. (Fig. 50E), A. miniducta sp. nov. (Fig. 32A)
and A. bermudensis (Sierwald 1988: g. 1), these latter two species without the lateral edge. A median
apophysis with transparent lamella are present in A. pectorosa, A. fraterna, A. lacka Platnick, 1974 and
A. alachua (Platnick 1974: gs 51–58). Similar RTA are present in A. scopulata F. Pickard-Cambridge,
1900 (Pickard-Cambridge F. 1900: pl. 7 g. 17), A. simplex O. Pickard-Cambridge, 1894 (Pickard-
Cambridge O. 1896: pl. 23 gs 1–2) and A. stigma sp. nov. (Fig. 50F).
Etymology
The species epithet is dedicated to Marcial Antonio Galán-Sánchez, Mexican arachnologist, and current
member of the second author’s lab.
Material examined
Holotype
MEXICO • ♀; Veracruz, Calcahualco, Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m;
2–11 Oct. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT; CNAN-T01540.
Allotype
MEXICO • ♂; same collection data as for holotype; CRP; AR_020; GenBank: ON619669; CNAN-T01519.
European Journal of Taxonomy 865: 1–94 (2023)
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Paratypes
MEXICO • 1 ♂; same collection data as for holotype; 4–17 Feb. 2014; LUP; AR_019; GenBank:
ON619668 CNAN-T01575 • 1 ♀; same collection data as for preceding; AR_013; GenBank: ON619660;
CNAN-T01576.
Additional material
MEXICO • 1 ♀; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
2–11 Oct. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT; AR_012; GenBank:
ON619659 • 1 ♀; same collection data as for preceding; ANYM045 • 1 ♀; same collection data as for
preceding; 4–17 Feb. 2014; BEAT; AR_016; GenBank: ON619670 • 1 ♀; same collection data as for
preceding; Xamaticpac, Plot II; 19.12614° N, 97.06708° W; alt. 1700 m; 19–27 Apr. 2013; Aracnolab
team leg.; oak and tropical wet forest fragment; BEAT • 1 ♀; same collection data as for preceding;
ANYM044 • 1 ♀; same collection data as for preceding; 4–17 Feb. 2014; BEAT.
Fig. 45. Anyphaena tonoi sp. nov. A–D. Paratype, ♀ (CNAN-T01576). E–F. Paratype, ♂ (CNAN-T01575).
A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Prosoma, anterior
view. F. Prosoma, oblique view. Scale bars: A–C = 1.0 mm; D–F = 0.5 mm.
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Description
Female
Total length 5.1. Carapace yellow, pattern with darker longitudinal bands over cephalic, thoracic areas
and clypeus, lateral edges yellow (Fig. 45A, D). Sternum surface light yellow, intercoxal triangles
present on all legs. Labium light brown, white at tip, longer than wide. Endites white, rectangular,
slightly broader at tip (Fig. 45C). Chelicerae yellow, slightly darker than carapace, paturon dorsum with
a diffuse dark pattern (Fig. 45B, D), promargin with four teeth, retromargin with eight or nine denticles.
Abdomen white, hirsute, dorsal surface with scattered brown patches, center with white chevrons and
marks, lateral and ventral surfaces with same pattern but faded ventrally, tracheal spiracle closer to
epigastric furrow (Fig. 45A–C). Epigynum atrium small, rectangular, longer than wide. Anterior lateral
sides delineated by two anterior shallow pits (Fig. 46E–F). Copulatory openings contiguous, inside
anterior edges of atrium. Copulatory ducts coiled over and around spermathecae, entering spermathecae
on lateral surface. Seminal receptacles not visible or inconspicuous. Fertilization ducts short, emerging
Fig. 46. Anyphaena tonoi sp. nov. A–D. Paratype, ♂ (CNAN-T01575). E–F. Paratype, ♀ (CNAN-T01576).
A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp, retrolateral
view. E. Epigynum, ventral view. F. Epigynum, dorsal view. Scale bars: 0.2 mm.
European Journal of Taxonomy 865: 1–94 (2023)
76
Fig. 47. Anyphaena tonoi sp. nov. A–F. Paratype, ♂ (CNAN-T01575). G–H. Paratype, ♀
(CNAN-T01576). A. Pedipalp, ventral view. B. Pedipalp, prolateral view. C. Pedipalp, dorsal view.
D. Pedipalp, retrolateral view. E. Coxae, ventral view. F. Coxae, lateral view. G. Epigynum, ventral
view. H. Epigynum, dorsal view. Scale bars: A–D, G–H = 0.2 mm; E–F = 1.0 mm.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
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from lateral surfaces below copulatory duct entrance (Figs 46E–F, 47G–H). Cephalothorax length 2.29,
thoracic width 1.8, cephalic width 0.97. Clypeus height 0.07. Eye diameters: AME 0.09, ALE 0.12,
PME 0.11, PLE 0.12. Eye interdistances: AME–AME 0.05, AME–ALE 0.02, ALE–PLE 0.09, PME–
PME 0.15, PME–PLE 0.1. Femur lengths: I 2.2, II 1.95, III 1.61, IV 2.2. Leg spination: femur I d1-1-1,
p0-0-2, r0-0-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1,
p0-0-1, r0-0-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1,
p0-1-1, r0-0-1. Tibia III v2-2-2, p1-1-1, r0-1-1. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1,
p0-1-1, r0-0-1. Tibia IV v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Male
Total length 5.6. Cephalothorax and abdomen coloration as in female. Chelicerae paturon pattern darker
(Fig. 45E). Embolus liform and translucent. Prolateral apophysis of palpal tibia present (Figs 46B–C,
47C). Pedipalp tibia longer than wide. Ventral branch of median tibial apophysis present (Figs 46A–D,
47A–D). Cephalothorax length 2.78, thoracic width 2.15, cephalic width 1.03. Clypeus height 0.11. Eye
diameters: AME 0.09, ALE 0.11, PME 0.12, PLE 0.12. Eye interdistances: AME–AME 0.04, AME–
ALE 0.04, ALE–PLE 0.07, PME–PME 0.09, PME–PLE 0.1. Femur lengths: I 3.61, II 2.68, III 1.95,
IV 2.93. Leg spination as in female except: femur II p0-0-2. Tibia III p0-1-1.
Variation
Females (N = 8): total length 5.29 (± 0.46), cephalothorax length 2.29 (± 0.25), thoracic width 1.73
(± 0.12), cephalic width 0.94 (± 0.07), femur I 2.09 (± 0.21). Males (N = 2): total length 5.55 (± 0.07),
cephalothorax length 2.71 (± 0.1), thoracic width 2.12 (± 0.03), cephalic width 1.0 (± 0.05), femur I 3.46
(± 0.21).
Distribution
This species is found in oak and tropical wet forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by beating tray and direct searching. This species is
present year-round.
Anyphaena megamedia sp. nov.
urn:lsid:zoobank.org:act:F1F5319C-F100-4D69-9370-E1A5BF56E6C0
Figs 48–49, 52
Differential diagnosis
Males of A. megamedia sp. nov. are differentiated from all those of the pectorosa and pacica groups by
the enormous size of the median apophysis, larger than the ventral tegular projection, and trapezoidal in
shape (Figs 48E–F, 49A, D).
Etymology
The species epithet refers to the large size of the median apophysis characteristic of these species.
Material examined
Holotype
MEXICO • ♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
4–17 Feb. 2014; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT; AR_024; GenBank:
ON619661; CNAN-T01530.
European Journal of Taxonomy 865: 1–94 (2023)
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Paratypes
MEXICO • 1 ♂; same collection data as for holotype; LUP; CNAN-T01558 • 1 ♂; Xamaticpac,
Plot II; 19.12614° N, 97.06708° W; alt. 1700 m; 2–11 Oct. 2013; AR_018; GenBank: ON619672;
CNAN-T01559.
Additional material
MEXICO • 1 ♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
2–11 Oct. 2013; Aracnolab team leg.; oak and tropical wet forest fragment; BEAT; AR_025; GenBank:
ON619662 • 1 ♂; same collection data as for preceding; AR_026; GenBank: ON619663.
Description
Male
Total length 6.6. Carapace yellow, darker over ocular quadrangle, around fovea and clypeus. Sternum
surface yellow, intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide.
Fig. 48. Anyphaena megamedia sp. nov., paratype, ♂ (CNAN-T01558). A. Dorsal habitus. B. Lateral
habitus. C. Ventral habitus. D. Prosoma, anterior view. E. Pedipalp, ventral view. F. Pedipalp, retrolateral
view. Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
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79
Endites yellow, rectangular, broader at tip (Fig. 48A–C). Chelicerae dark yellow without dorsal pattern
(Fig. 48B, D), promargin with four teeth, retromargin with eight to nine denticles. Abdomen surfaces
white, hirsute, covered with darker spots except posterior to tracheal spiracle located at middle (Fig. 48A,
C). Ventral tegular projection long, curved retro-ventrally, retrolateral edge with small transparent edge,
prolateral base enlarged. Embolus liform (Fig. 49A–B). RTA anterior branch largest, spine-shaped,
RTA posterior branch small and squared. Prolateral apophysis present. Tibia longer than wide. Ventral
branch of median tibial apophysis present (Figs 48E–F, 49A–D). Cephalothorax length 2.93, thoracic
width 2.24, cephalic width 1.06. Clypeus height 0.12. Eye diameters: AME 0.09, ALE 0.11, PME 0.12,
PLE 0.12. Eye interdistances: AME–AME 0.05, AME–ALE 0.04, ALE–PLE 0.12, PME–PME 0.17,
PME–PLE 0.11. Femur lengths: I 3.16, II 2.68, III 2.24, IV 3.02. Leg spination: femur I d1-1-1, p0-0-1,
r0-0-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0, p1-1-1, r1-1-1. Femur II d1-1-1, p0-0-1, r0-0-
1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0, p1-1-1, r1-1-1. Femur III d1-1-1, p0-0-1, r0-0-1.
Tibia III v2-2-2, p0-1-1, r0-1-1. Metatarsus III v2-2-2, p1-1-2, r1-1-2. Femur IV d1-1-1, p0-0-1, r0-0-1.
Tibia IV v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2, p1-1-2, r1-1-2.
Female
Unknown.
Variation
Males (N = 5): total length 6.12 (± 0.49), cephalothorax length 2.79 (± 0.14), thoracic width 2.18 (± 0.06),
cephalic width 1.05 (± 0.06), femur I 3.44 (± 0.14).
Fig. 49. Anyphaena megamedia sp. nov., paratype, ♂ (CNAN-T01558). A. Pedipalp, ventral view.
B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp, retrolateral view. Scale bars: 0.2 mm.
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80
Distribution
This species is found in oak and tropical wet forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Most specimens were collected over vegetation by direct searching or with a beating tray.
Anyphaena stigma sp. nov.
urn:lsid:zoobank.org:act:E3896101-5F6D-4CD5-BF28-C484B68151E7
Figs 50–52
Differential diagnosis
Males of A. stigma sp. nov. are differentiated from all those of the pectorosa and pacica groups by
the presence of the following features: middle section of ventral tegular projection broad, retrolateral
edge with a translucent edge, prolateral surface also distended; distal edge of median apophysis at and
curved, ventral edge with a transparent lamella (Figs 50E–F, 51A–D). RTA anterior branch spine-shaped
with a blunt tip, longer than posterior branch, which is shaped as a curved lamella. Coxae II and III with
a mid-ventral tubercle, coxa IV with two tubercles (Figs 50E–F, 51A–F). Ventral tegular projection with
broad middle sections and translucent retrolateral edges are also present in A. zorynae (Durán-Barrón
et al. 2016: gs 5, 9), A. pacica (Dondale & Redner 1982: g. 336), A. tonoi sp. nov. (Fig. 46A) and
A. miniducta sp. nov. (Fig. 32A). Similar RTA are present in A. scopulata (Pickard-Cambridge F. 1900:
pl. 7 g. 17), A. simplex (Pickard-Cambridge O. 1896: pl. 7 g. 11) and A. tonoi sp. nov. (Fig. 47A, D).
Etymology
The species epithet is derived from the Latin ‘stigma’ (‘mark’ or ‘brand’), in reference to the several
dark spots on the scutum and epiandrous region.
Holotype
MEXICO • ♂; Veracruz, Calcahualco, Xamaticpac, Plot I; 19.14172° N, 97.20597° W; alt. 1710 m;
4–17 Feb. 2014; Aracnolab team leg.; oak and tropical wet forest fragment, CRP; AR_022; GenBank:
ON619667; CNAN-T01539.
Paratype
MEXICO • 1 ♂; same collection data as for holotype; AR_021; GenBank: ON619666; CNAN-T01574.
Description
Male
Total length 6.5. Carapace yellow, darker over ocular quadrangle, around fovea and clypeus. Sternum
surface white, intercoxal triangles present on all legs. Labium brown, white at tip, longer than wide.
Endites yellow, rectangular, broader at tip (Fig. 50A–C). Chelicerae brown, paturon dorsal surface
covered with black reticulated pattern (Fig. 50D), promargin with four teeth, retromargin with eight to
nine denticles. Abdomen dorsal and lateral surfaces white, hirsute, covered with black marks, ventral
surface pattern diffused. Tracheal spiracle at center of abdomen (Fig. 50A, C). Prolateral apophysis
present. Tibia longer than wide. Ventral branch of median tibial apophysis present (Figs 50E, 51A–
D). Cephalothorax length 2.98, thoracic width 2.29, cephalic width 1.06. Clypeus height 0.11. Eye
diameters: AME 0.11, ALE 0.13, PME 0.15, PLE 0.15. Eye interdistances: AME–AME 0.05, AME–
ALE 0.02, ALE–PLE 0.05, PME–PME 0.17, PME–PLE 0.09. Femur lengths: I 2.98, II 2.68, III 2.24, IV
2.93. Leg spination: femur I d1-1-1, p0-1-1, r0-1-1. Tibia I v2-2-0, p1-1-1, r1-1-1. Metatarsus I v2-2-0,
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
81
p1-1-1, r1-1-1. Femur II d1-1-1, p0-1-1, r0-1-1. Tibia II v2-2-0, p1-1-1, r1-1-1. Metatarsus II v2-2-0,
p1-1-1, r1-1-1. Femur III d1-1-1, p0-1-1, r0-1-1. Tibia III v2-2-2, p0-1-1, r0-1-1. Metatarsus III v2-2-2,
p1-1-2, r1-1-2. Femur IV d1-1-1, p0-1-1, r0-1-1. Tibia IV v2-2-2, p0-1-1, r0-1-1. Metatarsus IV v2-2-2,
p1-1-2, r1-1-2.
Female
Unknown.
Variation
Males (N = 2): total length 6.4 (± 0.14), cephalothorax length 2.93 (± 0.07), thoracic width 2.29 (± 0.0),
cephalic width 1.05 (± 0.02), femur I 2.98 (± 0.0).
Fig. 50. Anyphaena stigma sp. nov., paratype, ♂ (CNAN-T01574). A. Dorsal habitus. B. Lateral habitus.
C. Ventral habitus. D. Prosoma, anterior view. E. Pedipalp, ventral view. F. Pedipalp, retrolateral view.
Scale bars: A–C = 1.0 mm; D = 0.5 mm; E–F = 0.2 mm.
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Distribution
This species is found in oak and tropical wet forest fragments around Pico de Orizaba Volcano National
Park (Fig. 52).
Natural history
Both specimens were collected by cryptic searching over leaf litter and tree bark.
Fig. 51. Anyphaena stigma sp. nov., paratype, ♂ (CNAN-T01574). A. Pedipalp, ventral view.
B. Pedipalp, prolateral view. C. Pedipalp, dorsal view. D. Pedipalp, retrolateral view. E. Coxae, ventral
view. F. Coxae, lateral view. Scale bars: A–D = 0.2 mm; E–F = 1.0 mm.
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Molecular analyses
Maximum Likelihood (Fig. 53) and Bayesian Inference (Fig. 54)
The COI data recovered a highly supported but unresolved Anyphaena and Wulla clade. The other 26
terminals represent 19 species and 15 genera of Anyphaenidae (Supp. le 1). Both analyses recovered a
paraphyletic Hibana at the base with one of its clades sister to the remaining genera, which are clustered
in three large clades: 1) Anyphaena and Wulla, 2) a cluster of several Anyphaeninae genera (Xiruana
Brescovit, 1997, Hatitia Brescovit, 1997, and Aysha), and 3) one group of species of Amaurobioidinae,
and Josa lutea (Keyserling, 1878) at the base of this triad.
The relations within the Anyphaena+Wulla clade have generally low support (ML) or remained
unresolved (BI). For the ML analysis, Anyphaena is divided in two clades, one formed by the pacica,
accentuata and celer groups; and the other formed by the pectorosa group as a sister to porta+Wulla
(Fig. 53). Our BI was not able to fully resolve the Anyphaena+Wulla branch (Fig. 54). However, the
accentuata+celer branch and the porta group are recovered as monophyletic with moderate to high
support values.
Fig. 52. Localities for the species of Anyphaena. Black square = Atotonilco Plots I and II, separated
by 400 m, nine species registered: A. bifurcata sp. nov., A. bromelicola Platnick, 1977 (ON619634-
38 Supp. le 1), A. catalina Platnick, 1974 (ON619673-75), A. dulceae sp. nov., A. epicardia
sp. nov., A. fernandae sp. nov., A. porta sp. nov., A. salgueiroi sp. nov. and A. soae sp. nov. White
square = Xamaticpac Plots I and II, plots contiguous, eleven species registered: A. adnani sp. nov.,
A. alachua Platnick, 1974 (ON619665), A. fraterna (Banks, 1896) (ON619671), A. megamedia sp. nov.,
A. miniducta sp. nov., A. natachae sp. nov., A. porta sp. nov., A. quadrata sp. nov., A. rebecae sp. nov.,
A. stigma sp. nov. and A. tonoi sp. nov. Black circle = Xilitla, four species registered: A. franciscoi
sp. nov., A. ibarrai sp. nov., A. noctua sp. nov. and A. triangularis sp. nov. White circle = Los Tuxtlas,
two species registered: A. jimenezi sp. nov. and A. urieli sp. nov.
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Fig. 53. Tree topology inferred with Maximum Likelihood in RaxML. Colored boxes indicate groups
of Anyphaena: pacica (gold), accentuata (red), celer (blue), pectorosa (green) and porta (ochre).
Numbers in the nodes indicate bootstrap support. Support values under 50 are marked in red. Note the
position of Wulla O. Pickard-Cambridge, 1895 as a sister of the porta group.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
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Fig. 54. Tree topology obtained with Bayesian Inference in MrBayes. Colored boxes indicate groups of
Anyphaena: pacica (gold), accentuata (red), celer (blue), pectorosa (green) and porta (ochre). Numbers
in the nodes indicate posterior probabilities. Support values under 50 are collapsed. Note that most of
the deeper relations within the Anyphaena+Wulla clade are not resolved, especially for the pacica
and pectorosa groups. The accentuata+celer and the porta groups are recovered as monophyletic clades
with moderate to high support.
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Accentuata+celer groups
The Palearctic and type species Anyphaena accentuata is consistently recovered as sister to the Nearctic
Anyphaena aperta (Banks, 1921), being in turn a sister to the celer group, both in the ML and BI
analyses. Three new species are described here for the celer group: A. natachae sp. nov., A. fernandae
sp. nov. and A. adnanai sp. nov. New sequences are provided for A. catalina and A. bromelicola. Within
this group, both analyses show moderate to high support values (60–100 in ML, and 0.71–1 in BI)
for all of its branches. Morphologically the celer group is also quite different from all other clades of
Anyphaena and could easily be diagnosed by its unique male and female morphological features; yet
species within the celer group are anatomically difcult to separate. For example, A. bromelicola and
A. adnanai sp. nov. are condently separated in both analyses (with ca 90% of internal node support)
and are geographically segregated; but their morphology could only be separated by the combination of
several features.
Pacica group
This group and its relation to other anyphaenas in our phylogenies is vague at best. ML suggests it has a
relation to the Accentuata+celer clade, being sister to the new species A. triangularis sp. nov. However,
the bootstrap support is very low in both cases (14% and 33%, respectively). Similarly, the BI fails to
give a clear picture of the relations of the pacica group, placing both A. pacica and A. triangularis
sp. nov. in collapsed branches.
Pectorosa group
The ML tree recovers the pectorosa as a lowly supported but monophyletic clade, including A. fraterna
and A. pectorosa, and four of our newly described species: A. bifurcate sp. nov., A. epicardia sp. nov.,
A. dulceae sp. nov., and A. soae sp. nov. Most of the deeper nodes within this clade are lowly supported
(> 30%); however, the most recent branches have generally higher supports, over 90% in most cases.
Similarly, our BI analysis condently recovered all of the internal subgroups of the pectorosa group.
However, their relationships to other members of Anyphaena are not clear.
Porta group
Both of our phylogenetic analyses recovered this group as monophyletic with moderate to high support
values (56 in ML, and 0.95 in BI). ML places this clade as sister to Wulla, while BI again fails to
resolve their relation to the rest of Anyphaena. This new group is formed by A. alachua and six new
species: A. porta sp. nov., A. salgueiroi sp. nov., A. tonoi sp. nov., A. megamedia sp. nov., A. ibarrai
sp. nov. and A. stigma sp. nov. Internally, most relations are well supported, rarely falling under 50% and
reaching 90–100% in most cases. A notable exception is A. porta AR14, whose long branch in ML and
paraphyletic position in the BI suggest this might be a separate species. However, careful morphological
observations failed to show any differences between this or other members of this species. Although COI
did not condently resolve the relationships within Anyphaena, there is a clear phylogenetic signal in
the data to support our morphological observations. We consider that Anyphaena as currently delimited
is not a monophyletic clade and new genera have to be erected to accommodate the pectorosa and porta
groups. Still, information from the ‘usual suspects’ (H3, 12s, 16s, 18s, and 28s), or a subset of them, with
a diverse taxonomic sample of other genera of Anyphaenidae would be desirable before making deeper
taxonomical changes within the genus.
Genital morphology is informative but can be quite problematic. In the case of males, diagnosing
Anypharena groups relying solely on male genital anatomy can lead to spurious relationships that are
not supported by other somatic character or COI data. We found the male coxal tubercles or spurs to be
a reliable diagnostic character that can be used in combination with genital features to efciently tell
species groups apart. For the females, the anatomy of the epigynal plate of Anyphaena is more diverse
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
87
than that of male palps. We found four main types of morphologies. However, substantial interspecic
variation is found within the groups, especially in the pectorosa and porta groups. Stretching the
homology to the limit was necessary to t some species within their groups (as suggested by COI and
male genital data).
Discussion
Anyphaena genital morphology
The great heterogeneity within some anyphaenid clades is derived from the very distinctive genital
characters shown by species groups that have a relatively limited geographic distribution. As Platnick
(1974) mentioned, many such groups might deserve a generic status as unambiguous key characters
are available. Nevertheless, a more pragmatic and conservative approach was preferred here selecting
nomenclatural stability over the creation of smaller genera that better reect the natural relationships.
This broad concept of Anyphaena has grouped together the Palearctic, Nearctic and Neotropical faunas
of Anyphaena.
Here, we found molecular and morphological evidence that supports the existence of Platnick’s (1974)
and other species groups. Nevertheless, we argue for a more careful consideration of their classication.
The considerable morphological diversity within Anyphaena (visually summarized in Supp. le 3),
together with the clear phylogenetic signal recovered from our molecular analyses, suggests that only the
celer group is directly related to the Nearctic A. accentuata, and the rest of the species of the Anyphaena
groups actually deserves a genus of their own. Nevertheless, additional molecular information outside the
scope of this work will be necessary to clarify the phylogenetic relationships of the putative Anyphaena
groups.
Integrative vs minimalistic taxonomic approaches
The present work describes 21 new species of Anyphaena from the Neotropics, representing the largest
contribution to the knowledge of this genus in the last four decades (WSC 2022). Previous landmarks on
the taxonomy of Anyphaena can be found in the faunistic work done by several European scientists in
the late 19th and early 20th centuries, and the revisionary work done by N. Platnick and his collaborators
during the 1970’s that resulted in an explosion of new species descriptions. Here, we benet from several
methodological, technological and conceptual breakthroughs that have occurred in the last decades
to increase the knowledge of the genus Anyphaena by describing new species and testing the former
hypotheses about the relationships of several species groups within this genus. The conclusions reached
here could only be achieved by the careful assessment of morphological and molecular information in
tandem. Any attempt at minimizing either one of the data sources would have resulted in a different
picture of the taxonomic relationships between the new species and their position within Anyphaena.
Furthermore, we would have failed to nd and describe all the new species, and greatly complicated –
in some cases – the sex matching of males and females. In other words, the reciprocal enlightenment
between molecules and morphology should not be understated.
Minimization of morphological data greatly speeds up the process of new species description, at the
expense of taxonomic literature usability and functionality. The minimalistic approach suggested by
Sharkey et al.’s (2021a, 2021b) claims as its main point the time and resources needed for generating
images, diagnoses and taxonomic keys. Because of these, they suggest their COI barcode diagnoses that
are considered a suite of character states universally shared by all members of a species and unique to
the species (Sharkey et al. 2021a). The authors highlight that traditional taxonomical approaches are
prone to error and that they are only useful for an ever-dwindling number of experts.
European Journal of Taxonomy 865: 1–94 (2023)
88
We are conscious of the necessary amount of time that needs to be invested in order to produce
morphological and molecular data that support new taxonomic descriptions. Although our 21 new
species are – to use Sharkey et al.’s (2021a) own words – a drop in the bucket, we argue that it is the
integration of multiple lines of evidence that gives robustness and ultimately usefulness to taxonomy.
Minimization of data might reduce the time needed for describing new species, but at the same time, it
reduces their economical support, adequacy, and usability (Janzen et al. 2009; Meier et al. 2021). As put
by Wheeler et al. (2004), “the challenge is not merely to speed data access but to expedite taxonomic
research”. The taxonomic impediment puzzle will not be solved only by accelerating the production of
data but also by facilitating their access and increasing their explanatory power. Besides COI being a
very cost-effective way (both in terms of time and money), other kinds of information are relevant too.
Sharkey et al (2021a) remarked on the need for taxonomy to show governments and the overall public
that there is a reason to bother. We consider that these arguments will not come from the number of new
taxa alone. Although molecular information is an extremely valuable tool, taxonomy – whether we like it
or not – is still an extremely visual science. Unless the knowledge and facilities for analyzing molecular
data are available to all the stakeholders, images and other kinds of biodiversity data (e.g., occurrence
databases) will still be the best way to transmit taxonomical ndings to the public and other scientists.
Well-curated taxonomic catalogs, databases and standardized image libraries expedite the scientic
discussion, allowing taxonomy to become a team effort instead of being dependent on individual expert
authorities. Despite the constant decrease in outlets, funding and positions in taxonomy and biodiversity
(Agnarsson & Kuntner 2007; Carvalho et al. 2007; Ebach et al. 2011; Zeppelini et al. 2020), it is the
democratization of information which has allowed taxonomy to keep increasing the number of described
species at an ever-growing pace. In the case of spiders, between 500 and 1000 new species have been
described every year in the last two decades (WSC 2022).
Morphological data, if done right, is still the best and most accessible way of quickly identifying taxa
with minimal equipment and facilities, despite the obvious exemption of many megadiverse and difcult
groups. One of the examples of Sharkey et al. (2021a) notably illustrates this point by comparing the
species Alabagrus scottshawi Sharkey et al., 2018 and A. genemonroei Sharkey et al., 2018, where
subtle morphological differences were found between both species only after COI indicated the need to
revisit these two morphospecies. A similar case was found in our study, where morphological differences
between A. porta sp. nov. and A. salgueiroi sp. nov. only became evident after the molecular data showed
them to be different. We found other cases where COI data coincided with segregated geographic
distributions resulting in the separation of A. bromelicola and A. adnani sp. nov., both species with
minimal easy-to-miss morphological differences. Still, similarity does not equal invariability and even
when morphology might seem to be redundant, it can become an important ally for parsing even some
of the toughest cases.
For most of the new spider species treated here, somatic and genital morphology offered an easy and
reliable way to identify and classify to species and species groups. This line of evidence (in stark contrast
to molecular data) does not require the application of expensive lab equipment or reagents, being easy
to do by a trained eye with a good microscope (and ideally a digital camera). Still, this step does require
some time and level of familiarity with the taxonomic group and their literature that is not always
readily available. Generating open access image libraries (Álvarez-Padilla 2012; Rivera-Quiroz et al.
2016; Álvarez-Padilla et al. 2020) and taxonomic literature repositories (WSC 2022) allows sharing
information worldwide, facilitating communication and reaching experts who can expedite and improve
species identications, even opening the door to ‘e-specimens’ based collaborations.
RIVERA-QUIROZ F.A. & ÁLVAREZ-PADILLA F., New ghost spiders (Anyphaenidae) from Mexico
89
With this discussion, we do not mean to appeal to one or the other source of information. On the contrary,
we invite the integration of different sources of data and infrastructures that can give taxonomy more
robustness and added scientic and social value. If we want to overcome the taxonomic impediment,
taxonomy must not be based on individual efforts but on a well-integrated team effort that makes use of
the diversity of methods for species documentation, web-based infrastructure, and scientic networks.
Acknowledgements
Thanks to Alejandro Oceguera and Ofelia Delgado for their great help with the DNA sequence
obtainment and analysis. Thanks to Antonio Brescovit for his comments on the early stages of the
species identication through our website. Thanks to the following students and arachnologists who
participated in the eld expeditions: Francisco J. Salgueiro-Sepúlveda, Uriel Garcilazo-Cruz, Dulce
F. Piedra-Jiménez, Mariana Servín-Pastor, Miguel A. Hernández-Patricio, Danielle Polotow, Facundo
M. Labarque, Thiago Silva-Moreira, Omar Caballero-Hernández, Diana E. Alvarez-Martínez, Rigel
S. González-Contreras, Leonel Pérez-Miguel and Maira S. Montejo-Cruz. To Edmundo Gonzalez-
Santillan for his advice and perspective during the early stages of this work. Thanks to the Xilitla
Foundation (Xilitla), Isidoro Contreras, Nicolás and their families (Atotonilco and Xamaticpac) for
their eld assistance. We also would like to thank Armando Fuertes from CONANP for presenting us
to Mr Isidoro Contreras and the Atotonilco community. Thanks to the editor Rudy Jocqué, the reviewer
Luiz Fernando M. Oliveira and an anonymous reviewer for their comments and suggestions. Funding
was provided by the projects UNAM-DGAPA-PAPIIT IN213612 and IN214916. All species described
here were collected under permit SGPA/DGVS/02403/12.
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Manuscript received: 16 June 2022
Manuscript accepted: 15 November 2022
Published on: 25 April 2023
Topic editor: Tony Robillard
Section editor: Rudy Jocqué
Desk editor: Kristiaan Hoedemakers
Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Muséum national d’histoire naturelle, Paris, France; Meise Botanic Garden, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences,
Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity
Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Leibniz
Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National Museum of the
Czech Republic, Prague, Czech Republic.
Supplementary material
Supp. le 1. Genbank Data. Table with the Genbank accession numbers for the sequences used in our
analyses. Other data like taxonomic information and the combination of primers for the new sequences
generated for the present study are also shown. https://doi.org/10.5852/ejt.2023.865.2097.8805
Supp. le 2. Aligned matrix (in Nexus format) used for our BI analysis. The same alignment was
transformed to the FASTA format and used for the ML, Haplotype network and genetic distances
calculations. We also included here the Trace and Marginal Density charts for our BI analysis.
https://doi.org/10.5852/ejt.2023.865.2097.8807
Supp. le 3. Visual summary of the diagnostic characters for the three Anyphaena groups represented
in our study. Ordering is based on our interpretation of the phylogenetic relations suggested by the
molecular analyses plus the morphological characters: green = pectorosa group, gold = pacica group,
blue = celer group, ochre = porta group. https://doi.org/10.5852/ejt.2023.865.2097.8809
