Zootaxa 4459 (2): 201–234 ISSN 1175-5326 (print edition)

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Copyright © 2018 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4459.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:B545851E-6B10-4D3F-B17B-2643BED06EFA

Taxonomic review of Epicadinus Simon, 1895 (Araneae: Thomisidae)

ANDRÉ WANDERLEY DO PRADO1,3, RENNER LUIZ CERQUEIRA BAPTISTA1 & MIGUEL MACHADO2
1
Laboratório de Diversidade de Aracnídeos, Universidade do Brasil/Universidade Federal do Rio de Janeiro. Av. Carlos Chagas
Filho 373, 21941-902, Ilha do Fundão, Rio de Janeiro, BRAZIL. E-mail: [email protected]; [email protected]
2
Laboratório de Aracnologia, Faculdade de Biociências, Pontifícia Universidade Católica do Rio Grande do Sul (PUCRS), Porto
Alegre, RS, BRAZIL. E-mail: [email protected]
3
Corresponding author

Abstract

The genus Epicadinus Simon, 1895 can easily be distinguished from the other Neotropical Stephanopines by their abun-
dant and robust setiferous tubercles, topped by elongated macrosetae, which cover most of the tegument. Additionally, the
genus can be recognized by a pair of conical ocular mounds above the ALE, the anterior eye row very recurved, posterior
one slightly procurved; carapace flattened and without tubercles, and opisthosoma with three conical projections (“tuber-
cles”) of variable size and shape. This work is a taxonomic review of the 12 valid species of Epicadinus as listed in version
19 of the World Spider Catalogue (2018), whose hitherto known distribution included few records from Brazil, French
Guiana, Bolivia and Peru, and only one from Mexico. Four valid species are recognized: Epicadinus biocellatus Mello-
Leitão, 1929; E. trispinosus (Taczanowski, 1872) [with two junior synonyms E. trifidus (Pickard-Cambridge, 1893) syn.
nov. and E. cornutus (Taczanowski, 1872) syn. nov.]; E. spinipes (Blackwall, 1862) [with two junior synonyms E. albi-
maculatus Mello-Leitão, 1929 syn. nov. and E. gavensis Soares, 1946 syn. nov]; and E. villosus [with two junior syn-
onyms E. helenae Piza, 1936 syn. nov. and E. marmoratus Mello-Leitão, 1947 syn. nov.]. Epicadinus polyophthalmus
Mello-Leitão, 1929 and Epicadinus tuberculatus Petrunkevitch, 1910 are transferred to Epicadus. Epicadus polyophthal-
mus (Mello-Leitão, 1929) comb. nov. is considered a nomen dubium. Epicadus tuberculatus (Petrunkevitch, 1910) comb.
nov. is a senior synonym of Epicadus pustulosus (Mello-Leitão, 1929) syn. nov.

Key words: Neotropical Region, Amazon, Atlantic Forest, crab-spiders

Introduction

Thomisidae Sundevall, 1833 is one of the most diverse spider families, with more than 2.168 species described in
170 genera (World Spider Catalog 2018). Thomisids are commonly known as crab spiders. They are ambush
predators, which do not use silk for hunting, and are usually found in shrubs, tree bark and flowers, where they
often camouflage. Some genera even have the ability to change color (Homann 1934, Comstock 1948, Silva-
Moreira & Machado 2016, Machado et al. 2017, Machado et al. 2018).
Although its monophyly is well supported by both morphological and molecular analysis (Benjamin et al.
2008, Benjamin 2011, Wheeler et al. 2017), the systematics of Thomisidae still needs more work with respect to
intrafamiliar relationships. As there is no comprehensive phylogenetic analysis aiming at the recognition of
subfamilies within Thomisidae, the tradidional division in seven subfamilies by Ono (1988) is still in use. Among
them, Stephanopinae is diagnosed by the presence of cheliceral teeth, parallel endites, a short and truncated labium,
robust anterior legs with ventral macrosetae on the tibiae and metatarsi, and well-developed posterior median eyes,
which are usually larger than the posterior lateral eyes (Mello-Leitão 1929, Ono 1988). Several papers have been
published by Brazilian arachnologists since the mid 20th century, helping to better understand the generic
taxonomic boundaries in the subfamily: Lise (1973) on Sidymella Strand, 1942, Lise (1981) on Onocolus Simon,
1895, Bonaldo & Lise (2001) on Stephanopoides Keyserling, 1880, Machado et al. (2015) on Tobias Simon, 1895,
and Silva-Moreira & Machado (2016) and Machado et al. (2018) on Epicadus Simon, 1895. However, Epicadinus
Simon, 1895 is still poorly known and in need of revision.

Accepted by C. Muster: 6 Jul. 2018; published: 15 Aug. 2018 201

 Epicadinus includes small crab spiders (females up to 8.5 mm; males up to 4 mm of body length) with
Neotropical distribution. The genus currently has 12 described species, 11 of them recorded from Brazil and
bordering countries, while only one occurrs in Mexico, E. trifidus (O. Pickard-Cambridge, 1893) (Melo-Leitão
1929, World Spider Catalog 2018). The type-species of the genus is Epicadinus trispinosus (Taczanowski, 1872),
which is based on a male holotype from French Guiana. In Brazil, most species have been recorded from the
Southeast region (E. albimaculatus Mello-Leitão, 1929, E. gavensis Soares, 1946, E. helenae Piza, 1936, E.
polyophthalmus Mello-Leitão, 1929, E. spinipes Blackwall, 1862, E. tuberculatus Petrunkevitch, 1910), with other
species known from the North (E. biocellatus Mello-Leitão, 1929, E. cornutus Taczanowski, 1872, E. trispinosus),
Northeast (E. albimaculatus Mello-Leitão, 1929, E. villosus Taczanowski, 1872) and South (E. helenae Piza, 1936,
E. marmoratus Mello-Leitão, 1947) regions (Mello-Leitão 1929, Piza 1936, Soares 1946, Mello-Leitão 1947).
According to the literature, the genus is diagnosed by the ocular mound high and with two acute tips, anterior eye
row very recurved, posterior one slightly procurved, carapace flat, without tubercles, opisthosoma with three
conical projections (tubercles) of variable size and shape, and paired claws of the anterior tarsus pectinate, with a
series of long and numerous teeth (Simon 1895; Mello-Leitão 1929).
Among the 12 described species of Epicadinus, five were described by Mello-Leitão (1929, 1947), who also
redescribed species from other authors and added new distribution records. Besides original descriptions, species
of the genus were cited only in a few inventories of araneofauna (e.g. Santos 1999, Castanheira et al. 2016). An
unidentified species of Epicadinus was also included in a molecular phylogenetic analysis of Thomisidae by
Benjamin et al. (2008). The classical subfamily Stephanopinae was not fully recovered in that study, with members
distributed in several clades belonging to different traditional subfamilies and without considerable morphological
similarities between them. The position of Epicadinus in particular was not clearly resolved. The genus emerged as
part of the Stephanopis-clade in some analyses, but it was placed near the Thomisus-clade in others. More recently,
Silva-Moreira & Machado (2016) suggested close relationships among Epicadus, Epicadinus, Onocolus,
Rejanellus Lise, 2005 and Tobias. Subsequently, Machado et al. (2017) tested the monophyly of Epicadus (and its
synonym Tobias) in their morphological phylogenetic analysis, including representatives of all the aforementioned
genera as outgroups. Although it was not the main objective of their work, these authors also discussed the
relationships between these Neotropical Stephanopinae, proposing an “Epicadus clade”, including Epicadinus,
Epicadus and Onocolus, but excluding Rejanellus.
The absence of a review of Epicadinus entails serious difficulties to identify its component species, since most
of them have been described succinctly, with insufficient illustrations and lack of accurate diagnoses. In this paper,
we revise and redescribe the species of Epicadinus, provide a key for species identification, and a map the
distribution area of each species, correlating biomes and areas of endemism.

Material and methods

This study was based in 243 samples comprising 508 specimens (257 females, 149 males and 102 juveniles) from
the following institutions (abbreviations and curators within parentheses): Facultad de Ciencias, Instituto de
Biología, Universidad de la Republica, Montevideo, Uruguay (FCE, M. Simó), Museu de Ciências Naturais da
Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre (FZBRS, R. Ott), Instituto Butantan, São Paulo (IBSP,
A. Brescovit), Instituto Nacional de Pesquisas da Amazônia, Manaus (INPA, C. Magalhães), Laboratório de
Biodiversidade de Aracnídeos/Universidade Federal do Rio de Janeiro, Rio de Janeiro (UFRJ, R. Baptista), Museo
Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires (MACN, M. Ramírez), Museum of
Comparative Zoology, Harvard University, Cambridge (MCZ, G. Giribet), Museu de História Natural do Capão do
Imbuia, Curitiba (MHNCI, A. Silva), Museum and Institute of Zoology, Polska Akademia Nauk, Warsaw (Polish
Academy of Sciences) (MIZ, W. Wawer), Musée National d´Histoire Naturelle, Paris (MNHN, C. Rollard), Museu
Nacional do Rio de Janeiro, Rio de Janeiro (MNRJ, A. Kury), Museu Paraense Emilio Goeldi, Belém (MPEG, A.
Bonaldo), Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP, R. Pinto-da-Rocha), Peabody
Museum of Natural History, Yale University, New Haven (PMNH, R. Pupedis), Museu de Ciências e Tecnologia,
Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre (PUCRS, R. Teixeira), Oxford University
Museum of Natural History, Oxford (OUMNH, Z. Simmons), and Universidade Federal de Minas Gerais, Belo
Horizonte (UFMG, A. Santos).

202 · Zootaxa 4459 (2) © 2018 Magnolia Press PRADO ET AL.

 Most of the holotypes were analyzed in loco except for E. cornutus (Taczanowski, 1872), E. trispinosus
(Taczanowski, 1872) and E. tuberculatus Petrunkevitch, 1910 which were examined by photographic registers. The
holotypes of E. trifidus (O. Pickard-Cambridge, 1893) and E. polyophthalmus Mello-Leitão, 1929 were not found
in their respective house collections (OUMNH and MNRJ) and are herein considered lost. Type material of E.
polyophthalmus had already been considered as lost by Silva-Moreira (2010).
Images and descriptions of external morphology and genitalia were obtained from specimens preserved in 75%
ethanol under a LEICA M205 C binocular stereoscopic microscope with a Leica DFC 450 digital camera attached.
The female genitalia were dissected and clarified in 10% KOH at 27 ºC for approximately 24 hs, in order to remove
soft tissues. For scanning electron microscopy (SEM), specimens were critical-point dryed. The images were
obtained through a Jeol JSM 6510 microscope in the Laboratório de Microscopia do Instituto de Biologia - UFRJ.
Descriptions and terminology follow Ono (1988), Benjamin (2011) and Machado et al. (2018). To indicate the
position of the transversal furrow in the tegulum of the male genitalia, we used clock positions in ventral view of
the left male palp, as in Machado et al. (2015, 2018) and Silva-Moreira & Machado (2016). Only non-expanded
bulbs were analyzed to determine furrow position, since the tegular rotation in an expanded bulb alters the position
of the furrow (compare, for example, Fig. 12D and 12E, see also Lise 1981). All measurements are in millimeters
and were obtained through the software LEICA Application Suite vers. 4.12. As an exception, the measurements of
the type-material unavailable for analysis were taken from the original descriptions.
The maps were constructed using QGIS 2.14 (QGIS Development Team 2018), based on the coordinates
obtained on Google Earth (https://www.google.com/earth). Inaccurate locations were presented on the maps
approximately, using the narrowest geographic location possible. Illustrations were made in Adobe Illustrator CC
2015, and the editing of images and making of plates were made in Adobe Photoshop CS6.
Anatomical abbreviations: A, atrium; AC, anterior curve of the copulatory duct; E, embolus; EA, embolus
apex; ALE, anterior lateral eyes; AME, anterior median eyes; BP, basal pouch; CD, copulatory duct; CO,
copulatory opening; CV, superior curve of the copulatory duct; FD, fertilization duct; DTA, distal tibial apophysis;
MF, marginal furrow of the cymbium; MS, median septum; MOQ, median ocular quadrangle; PLE, posterior
lateral eyes; PME, posterior median eyes; PST, paired setiferous tubercles; RTA, retrolateral tibial apophysis; S,
spermatheca; TF, tegular furrow; TK, translucent keel; TM, tibial macrosetae.

Taxonomy

Thomisidae Sundevall, 1833

Stephanopinae O. Pickard-Cambridge, 1871

Epicadinus Simon, 1895

Epicadinus Simon, 1895: 1052–1053. Mello-Leitão 1929: 98, pl. 7, figs 12–20.
Type species. Thomisus trispinosus Taczanowski, 1872, by original designation.

Diagnosis. Epicadinus is related to Epicadus and Onocolus in the general shape of the male palpus, presenting a
discoid tegulum and filiform embolous encircling it, and in the canoe-shaped RTA fused to the DTA. Some
Epicadus (e.g. E. heterogaster (Guérin-Menéville, 1829) and E. rubripes Mello-Leitão, 1924) share the presence of
a pair of conical projections above the ALE with Epicadinus (Fig. 1A), but the flat carapace of Epicadinus (Fig.
1C) contrasts with the presence of a median dorsal projection in the foveal area, at least in females, of most
Epicadus species (heterogaster-clade sensu Machado et al. 2017). Epicadinus species can be recognized by their
spiny appearance, since most of their tegument is covered by robust setiferous tubercles, surmounted by elongated,
needle-shaped macrosetae (Figs 1D, E, I; 8A–C, G). Additionally, the paired macrosetae present on the tibiae and
metatarsi I–II are not quite mobile and inserted directly on the leg surface as in most Thomisidae, but rather located
on the largest setiferous tubercles (Fig. 8G). Some Australian species described in Sidymella and juvenile
specimens of species in the pustulosus-clade of the genus Epicadus also have spiny tegument, but the setiferous
tubercles are not elongated.
Description. Spiders with clear sexual dimorphism. Males about 3/5 of females size. Carapace (Fig. 1B) rather

TAXONOMIC REVIEW OF EPICADINUS Zootaxa 4459 (2) © 2018 Magnolia Press · 203
longer than wide, almost flat, without dorsal projections, only with ocular region elevated, presenting two clear
projections above ALE, forming a high ocular mound with two acute tips. Dorsum with three median rows of
setiferous tubercles and six rows between radial furrows, and others irregularly distributed. Posterior slope of
carapace without tubercles or macrosetae. Clypeus high, at least 1.5x AME diameter. Ocular area with anterior row
strongly recurved and posterior slightly procurved. MOQ trapezoid. ALE size equal to or slightly larger than AME
(Figs 1A, B). Labium up to twice as wide as long. Sternum cordiform, almost as wide as long, bearing abundant
spiny or feathery bristles. Endites setaceous.
Opisthosoma generally pentagonal (but with trapezoidal anterior half and semicircular posterior half in E.
biocellatus), covered by abundant tubercles of variable size. Dorsum with three prominent, conical projections,
with apex directed laterally or perpendicular to midline of opisthosoma. First pair of conical projections of similar
size and located at median portion of opisthosoma. Posterior conical projection strongly variable in size and shape
in females and slightly variable in males. Anterior border generally with projected angles (except E. biocellatus),
usually covering posterior slope of carapace. Venter covered by thick feathery bristles, more abundant anteriorly,
without setiferous tubercles, and with two longitudinal and parallel rows of sigils, sometimes almost
inconspicuous. Small, triangular spinnerets covered by abundant bristles (Fig. 1H).
Small chelicerae covered by abundant bristles. Fang furrows toothless and with abundant, long and erect
bristles. Fangs short and curved (Fig 1A).
Leg formula 1243, with legs I and II up to twice as long as legs III and IV. Legs with abundant setiferous
tubercles, larger on tibiae and metatarsi, each one topped by an elongated macroseta. Legs I–II with setiferous
tubercles larger and more abundant than in posterior legs. Tibia I-II ventrally with five pairs of macrosetal tubercles
and metatarsus I-II with four pairs. Paired macrosetae not mobile and located on top of largest tubercles present in
body. Macrosetal tubercles slightly offset to sides and directed externally, nearly lateral to longitudinal axis of
article. Areas without setiferous tubercles or bristles with abundant small granules (Fig. 1D). Larger granules,
similar to warts, grouped in distinct areas, such as lateral clusters in femurs, two dorsal bands between rows of
setiferous tubercles on dorsum of patella and tibia (wider in first article), and two marginal bands on tibia, near
patella joint (Figs 1E, F). Tarsus with one pair of unequal pectinated claws; prolateral claw (mesal) with 5–6 acute,
sharp and straight teeth in basal portion, 5 thick, blunt and straight teeth in apical portion; retrolateral claw (ectal)
with only 5 thick apical teeth, and basal teeth replaced by a gap and a longer basal keel (Fig. 1G). Claw tuft with
about 25–30 spatulated setae.
Palpus of male with roundish cymbium, with pointed apex. Alveolus deep, forming a pronounced convexity in
dorsal face of cymbium. Tegulum roundish and flattened, covering apex of embolus in some species (Figs 9E–F).
When uncovered (Figs 3H, 6E), apex of embolus protrudes into a deep marginal furrow on the triangular, apical
glabrous area of tegulum (Fig. 3H). Embolus encircling the margin of tegulum, varying from half a turn up to six
turns around tegulum. RTA with central concavity, limited by marginal folds, generally elongated (except E.
helenae). Base of RTA partially fused to DTA, both variable in size. DTA hook- or horn-shaped, usually quite
distinct and smaller than RTA (Figs 3G, 6G, 9F, 12F).
Epigynum with atrium (or median field) usually very conspicuous (except E. biocellatus) and median septum
either covering or not copulatory openings (Figs 2E, 5D, 8E). Septum located posteriorly to atrium (except E.
biocellatus). Vulva structure variable, with helicoidal copulatory ducts presenting six to ten turns (E. spinipes and
E. villosus, Fig. 8F), or forming an expanded anterior curve (or secondary spermathecae), followed by a few
posterior curves (Figs 2F, G; 5F). Spermathecae roundish and well developed, located near epigastric furrow.
Color of specimens variable, from light yellow to orange and dark brown in 70% ethanol (Figs 6A–C, 11A–C,
G). Live specimens can show light green hues (Fig. 1I) or more vivid colors (Fig. 1J).
Composition. Four species. Epicadinus biocellatus Mello-Leitão, 1929, Epicadinus trispinosus (Taczanowski,
1872), Epicadinus spinipes (Blackwall, 1862), and Epicadinus villosus Mello-Leitão, 1929.
Distribution. Neotropical region from southern Mexico to northern Uruguay. In South America, there are
records only east of the Andes, from Amazonian Ecuador at west, to Bahia, Brazil at east.
Natural history. Despite the scarce knowledge about the ecology and behavior of Epicadinus species, the
specimens are usually found on leaves and twigs of different plants, where they hunt by the sit-and-wait behavior,
as many other thomisids, and use their strong anterior legs to catch and hold prey (Fig. 2H). Many specimens were
captured near ground, over herbs and small bushes in open areas, inside or at the border of forests.

204 · Zootaxa 4459 (2) © 2018 Magnolia Press PRADO ET AL.

FIGURE 1. A Epicadinus villosus Mello-Leitão, 1929, female (FZBRS 39201), carapace frontal; ,B E. spinipes (Blackwall,
1862), female (UFRJ 1326), carapace dorsal; C E. trispinosus (Taczanowski, 1872), male (FZBRS 8939), carapace lateral; D–F
E. spinipes, female (UFRJ 1336) (D femur I dorsal E patella I dorsal view with two rows of scale-like structures, F detail of E),
G–H E. villosus, female (UFRJ 1330) (G pair of tarsal claws with unequal teeth, H spinnerets, ventral); I–L E. villosus, live
specimens (I female (UFRJ 1330), J male (FCE-Ar 8444)). Scale bars: A–C, 1 mm; D, E, H, 0.1 mm; F,G, 0.25 mm. Photo
Credits: J, Alvaro Laborda.

TAXONOMIC REVIEW OF EPICADINUS Zootaxa 4459 (2) © 2018 Magnolia Press · 205
 Species groups. The four species of Epicadinus fall in two species groups, clearly set apart by morphological
and genitalic similarities. The trispinosus-group includes E. biocellatus and E. trispinosus. Both species have
females presenting the posterior conical projection perpendicular to the opisthosoma (Figs 2B, 5C), epigynum with
median septum totally fused, visible copulatory openings and sinuous and non-spiral, short copulatory ducts (Figs
2D–G, 5D–F). The males of E. trispinosus and E. biocellatus have a V- or U-shaped notch between the RTA and the
DTA, RTA with ventral fold forming a deep basal pouch, and embolus with a clearly visible apex and forming an
incomplete turn around the tegulum (Figs 3E–H, 6D–G). This group occurs mainly in the Amazon. E. biocellatus is
restricted to eastern Amazonia, while E. trispinosus is a widespread species, occurring in western Amazonia and
also in several other biomes, from Mexico to the Bolivian chaco and the Brazilian state of São Paulo. The last two
records are from open vegetation biomes (chaco and an ecotone between the Brazilian Cerrado and Atlantic Forest,
respectively, Fig. 15). The second group is the spinipes-group, including E. spinipes and E. villosus. Females of
both species have an posterior conical projection arranged transversely to the opisthosoma (Figs 8C, 11C),
epigynum with medial portion of the median septum covering copulatory openings, and inner genitalia with spiral
copulatory duct, composed of six or more coils (Figs 8D–F, 11D–F). Males of these species present the dorsal fold
of the RTA directly fused to the base of the DTA, without basal pouch, and a long embolus, with at least three turns
around the tegulum, and with apex hidden by the tegulum (Figs 9D–F, 12D–F). This group is restricted to the
Brazilian Atlantic Forest and its ecotones. E. spinipes is found near the coast of northeastern and southeastern
states of Brazil (Bahia to São Paulo), while E. villosus occurs away from the coast in the northern portion of its
range, from Bahia to São Paulo, but is found also near the coast from southern São Paulo to Uruguay (Fig. 15). The
records from Brazilian Amazon are considered doubtful (see species description below).

Key to the species

Females
1 Opisthosoma with posterior conical projection transversal to venter (Figs 8C, 11C), epigynum with central portion of median
septum covering copulatory opening, genitalia with a spiral copulatory duct composed of six or more coils (Figs 8E, F, 11E, F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Opisthosoma with posterior conical projection perpendicular to venter (Figs 2B; 5C), epigynum with median septum totally
fused, copulatory openings visible and copulatory ducts sinuous and non-spiral (Figs 2D–G, 5D–F). . . . . . . . . . . . . . . . . . . . . 3
2 Carapace with a thin, yellowish median stripe and two dark brown stripes, two yellowish lateral stripes on dorsum of opistho-
soma, sternum yellowish or with brown marginal spots (Figs 8A, B). Paired macrosetal tubercles of tibia and metatarsus longer
than or equal to half the article width (Fig. 8G). Epigynum with triangular median septum with anterior base at least 2x wider
than posterior one (Fig. 8E). Copulatory duct spiral, with about 10 coils, the last one partially covering dorsum of spermatheca
(Fig. 8F). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. spinipes
- Carapace with a long, median longitudinal brown stripe, bifurcate posteriorly, dorsum of opisthosoma without white lateral
bands, dark brown sternum with yellowish central area (Figs 11A, B). Paired macrosetal tubercles of tibia and metatarsus
shorter than half the width of the article. Epigynum with rectangular median septum, bearing anterior base almost as wide as
posterior one (Fig. 11F). Copulatory duct spiral, with about seven coils, the last one not covering dorsum of spermatheca (Fig.
11E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .E. villosus
3 Carapace with a longitudinal, median brown stripe, bifurcate posteriorly, opisthosoma with semicircular posterior portion,
anterior margin straight and not projected on carapace, posterior conical projection half as high as opisthosoma (Figs 2A, B).
Epigynum very small, wider than long, with atrium almost inconspicuous, median septum and copulatory openings located
inside atrium, posterior epigynal area wide (Fig. 2D, E). Copulatory ducts contouring inner margin of spermatheca (Figs 2F, G)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. biocellatus
- Carapace with a longitudinal, median yellowish stripe between two light brown stripes, opisthosoma with triangular posterior
portion, anterior margin slightly concave and projected on carapace, posterior conical projection much higher than opistho-
soma (Figs 5A, C). Epigynum longer than wide, with deep and well-delimited atrium, median septum and copulatory openings
located outside atrium (Fig. 5D). Copulatory duct not contouring margin of spermatheca (Fig. 5F) . . . . . . . . . . . E. trispinosus

Males
1 Palpus with external margin of RTA fused to the basis of DTA, ventral fold of RTA slightly pronounced (Figs 9F, 12F),
embolus long, forming at least three turns around tegulum, and with apex covered by tegulum (Figs 9E, F; 12E, F) . . . . . . . . 2
- Palpus with a V- or U-shaped notch between the RTA and DTA, ventral fold of RTA forming a deep basal pouch (Figs 3G, 6F,
G), short embolus, not forming a complete turn around tegulum, with clearly visible apex (Figs 3G, H, 6E). . . . . . . . . . . . . . 3
2 Sternum yellowish (Fig. 9C), RTA about 2x longer than wide, RTA apex forming an acute and elongated curve (Figs 9E, F),
embolus with about 4 turns around tegulum (in semi-distended palpus, Fig. 9F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. spinipes
- Sternum reddish brown (Fig. 12C), radially bordered by distinct dark brown spots, RTA slightly longer than wide, and with its

206 · Zootaxa 4459 (2) © 2018 Magnolia Press PRADO ET AL.

 apex forming a wide and rounded curve (Figs 12E, F), embolus with about 6 turns around tegulum (in semi-distended palpus,
Figs 12E, F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .E. villosus
3 Carapace yellowish with a dark brown longitudinal stripe, bifurcate posteriorly, legs yellowish (Fig. 3A). Opisthosoma with
semicircular posterior half, posterior conical projection similar in size to median pair (Figs 3A, B). RTA sinuous, at least 3x
longer than wide, margin of ventral fold S-shaped, basal pouch barely visible in ventral view, DTA short and horn-shaped (Fig.
3F, G). Palpal tibia dorsally with large setiferous tubercle directed retrolaterally, harboring very thick macrosetae (Figs 3E, F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. biocellatus
- Carapace reddish brown, with a light-yellow stripe separating two dark brown stripes, femur and patella I and II brownish,
strongly contrasting to other yellowish articles (Figs 6A–C). Opisthosoma pentagonal, with posterior conical projection clearly
smaller than median pair (Figs 6A, C). RTA about 2x longer than wide, margin of ventral fold forming a long, convex curve,
basal pouch very conspicuous in ventral view, DTA very long, almost as long as hook-shaped RTA (Figs 6E–G) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. trispinosus

Epicadinus biocellatus Mello-Leitão, 1929

Figs 2–4, 15

Epicadinus biocellatus Mello-Leitão, 1929: 104.

Epicadinus trispinosus (Taczanowski). Mello-Leitão 1929: 99 (in part)

Type material: Holotype: female, BRASIL or PERU, “Para ou Pebas ♀ type”, [M. de Mathan] (MNHN AR-
15077, Coll. Simon 2044b), examined.
Additional material examined. BRAZIL: Maranhão: 1 female, Bom Jardim, Reserva Biológica do Gurupi,
3°55'34.31''S, 46°46'19.21''W, 21 October 2011, M. A. Aguiar-Neto (MPEG 31329); 1 female, Centro novo do
Maranhão, Reserva Biológica do Gurupi, 3°40'54.09''S, 46°45'15.24''W, 15 March 2012, M. A. Aguiar-Neto
(MPEG 31322); idem, 1 female, 3°41'7.92''S, 46°45'46.08''W, 9 November 2011 (MPEG 31327); idem, 1 male
(MPEG 31328); 1 male, Centro novo do Maranhão, Cândido Mendes, Fazenda Sete Irmãos 1°51'48.08''S,
45°45'39.02''W, 08 May 2014, M. A. Aguiar-Neto (MPEG 31324), idem, 2 females, 8 May 2014, M. A. Aguiar-
Neto (MPEG 31330). Pará: 1 female, Barcarena, ca. 1°34'47.68"S, 48°35'55.92"W, 19 November 2001, Domingos
(MPEG 31332); 1 male, Belém (MNHN Coll. Simon 2044); 1 male, Belém, Reserva Mocambo, 01°26'28.07''S,
48°24'46.02''W, 11 December 2007, G. Ruiz (MPEG 31333); 1 female, Belém, Bosque Rodrigues Alves,
1°25'49.00''S, 48°27'22.03''W, 22 April 2009, E. G. S. Cafofo (MPEG 31334); 2 females, Belém, Museu Paraense
Emílio Goeldi, Campus de Pesquisa, 1º27'04.44"S, 48°26'39.32"W, 08 February 2018, P. Pantoja (MPEG 34862); 1
female, Cametá, Vila de Curuçambaba, 2°06'27.02''S, 49°18'33.1''W, 26 June 2014, Y. Shimano (MPEG 31335); 1
male, Santarém, Alter do Chão, ca. 2°30'36.96"S, 54°57'10.56"W, 26 January 1994, A. D. Brescovit (FZBRS
25051), idem, 1 female, 26 June 2007, M. L. N. Sirotheau (MPEG 31336), idem, 1 female, 02 July 2007, B. J. F.
Silva (MPEG 31337), idem, 1 female, 02 July 2007, M. L. N. Sirotheau (MPEG 31338), idem, 1 female, 30 June
2007 (MPEG 31339), idem, 1 male, 04 July 2007, B. J. F. Silva (MPEG 31340); 1 male, Viseu, São José do Gurupi,
1°34'02.03''S, 46°16'47.04''W, 11 March 2014, M. A. Aguiar-Neto (MPEG 31323), idem, 1 female , (MPEG
31326); idem, 1 male, -1°33'16.09''S, 46°16'18.06''W (MPEG 31325).
Diagnosis. E. biocellatus resembles E. trispinosus, but females of E. biocellatus are distinguished by the
carapace presenting a wide brown, median longitudinal stripe, bifurcate posteriorly, and by the opisthosoma with
semicircular posterior portion, anterior margin straight and not projected towards the carapace and by the posterior
conical projection about half as high as the opisthosoma (Fig. 2A). The epigynal plate is reduced, wider than long,
with an almost inconspicuous atrium, median septum and copulatory openings are located within the atrium and
presenting a large posterior fold (Fig. 2D, E). The copulatory ducts contour the inner margin of the spermathecae
(Fig. 2G). Males of E. biocellatus are distinguished by the yellowish-brown carapace with a dark brown, median
longitudinal stripe, bifurcate posteriorly, yellowish legs, and roundish opisthosoma (Fig. 3A, B). The palpus
presents a RTA sinuous and very long, at least 3x longer than wide, with ventral fold S-shaped, DTA short and
horn-shaped and dorsum of tibia with large setiferous tubercle, directed retrolaterally, harboring a very thick
macroseta (Fig. 3E, F).
Notes. Mello-Leitão (1929) cited “Para” as the type locality of E. biocellatus. However, the analysis of the
holotype in Simon Collection (accession number 2044b, MNHN, Figs 4A–B) shows that its label, handwritten by
Mello-Leitão himself, clearly states "Para ou Pebas ♀ type" (Fig. 4C), indicating that the specimen was collected in
one of those two localities. The word "Para" or "Le Para" was a reference used by Simon and other nineteenth

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century collectors to refer not to the Brazilian state of Pará, but to the old name of the city of Belém: Grão Belém
do Pará (e.g. Levi 1964: 72, Papavero 1973: 377, Levi 1996: 108, 142). Simon normally used the word “Prov.” or
“Province” when referring to the current state of Pará (e.g. Simon 1895: 159). The other locality is Pebas, a city
near the Amazon River, in the department of Loreto, Peru, near the border with Brazil. This specimen most
probably was collected by Marc de Mathan, a French collector who followed the Amazon River and its tributaries,
from Belém to northern Peru, reaching up to Ecuador, during the end of the 19th and the beginning of the 20th
centuries (Papavero 1971: 161). Mathan was the sole collector cited for Pebas and he also collected in “Le Para”
(e.g. Simon 1893: 310, 326; 1895: 135, 153). Unfortunately, spider samples deposited in MNHN, identified as
belonging to a single species, often refer to more than one locality and may have been mixed in just one flask (R.
Baptista, personal observation). Simon probably had identified three specimens as E. trispinosus, all included
under accession number 2044, although they belonged to different samples from Belém and Pebas (Fig. 4D).
Mello-Leitão (1929) considered these specimens as belonging to three different species and splitted this mixed
sample into three log numbers: one with a male (2044) that he also identified as E. trispinosus, agreeing with
Simon (Fig. 4D), another with a female of E. cornutus (2044a, Fig. 4E), and the last one with the female holotype
of E. biocellatus (2044b, Figs 4A–C). The male identified as E. trispinosus (2044) by Mello-Leitão is, in fact, the
then undescribed male of E. biocellatus, as many samples of the same males and females from several localities
Pará suggest. The other female (2044a) actually is E. cornutus, but Mello-Leitão committed a lapsus while copying
the label, including only "Para," without referring to "Pebas". In MNHN, there is a single female specimen of E.
biocellatus (2044b), indicating that the species description was not based on a type series, but on a single specimen
collected either in Pará or Pebas. The male of E. biocellatus (2044) included in the same flask was not described by
Mello-Leitão, since this author mistakenly attributed it to E. trispinosus.
We assume that the true locality of the species is actually Belém, since many specimens were collected in this
locality and surrounding areas of Pará and Maranhão states, both in Brazil. In addition, no specimens of E.
biocellatus from western Amazon, a region where Pebas is located, were found in the collections examined.
Conversely, the female of E. cornutus (2044a) is most probably the specimen that was originally collected in Pebas,
since no sample of this species has been collected in Pará or other areas of eastern Amazon, and there are several
records from western Amazon, surrounding the Peru locality, i. e., Ecuador and the Brazilian states of Amazonas
and Acre.
Description. Female (MPEG 31330): Total length: 4.39. Carapace: length 1.92, width 1.80, height 1.19.
Clypeus: height 0.25. Chelicera: length 0.57, width 0.27. Sternum: length 0.87, width 0.69. Endites: length 0.28,
width, 0.25. Labium: length 0.15, width 0.30. Eye diameters and interdistances: AME 0.07, ALE 0.08, PME 0.06,
PLE 0.05, AME–AME 0.09, AME–ALE 0.07, ALE–ALE 0.23, PME–PME 0.12, PME–PLE 0.06, PLE–PLE 0.36.
MOQ trapezoidal. Leg formula: 1243. Leg I – femur 1.69, patella 0.94, tibia 1.85, metatarsus 1.46, tarsus 0.69,
total 6.63; II: 1.46-0.74-1.68-1.33-0.63-5.84; III: 0.66-0.51-0.73-0.60-0.40-2.90; IV: 1.08-0.48-0.86-0.70-0.45-
3.57. Paired setal tubercles on tibiae and metatarsi with length less than or equal to half the article width.
Opisthosoma: length 2.47, width 2.81, height 3.28, with trapezoidal anterior half and semicircular posterior half in
dorsal view, covered by abundant macrosetae of variable size. Anterior dorsal margin not projecting over posterior
portion of carapace, straight in dorsal view, with anterior angles not projected. Paired conical projections smaller in
median portion, directed posterolaterally. Posterior conical projection placed a bit more posteriorly, in almost
central position, and dorsally directed, nearly perpendicular to venter, about twice as long as paired ones and
measuring about half the height of the opisthosoma itself. In lateral view, dorsum very convex both before and after
posterior conical projection. Venter with anterior portion profusely setaceous.
Epigynum (Figs 2D, E) slightly wider than long, with septum and copulatory openings located within atrium,
separating it into two parts, and a large flattened posterior area, about 1.5x longer than atrium. Epigynum small,
occupying a much smaller area than the one occupied by internal genitalia. Atrium reniform, very wide,
approximately 3x wider than long, very shallow, almost inconspicuous, delimited by a light marginal depression,
strongly demarcated on anterior margin and inconspicuous on most of posterior margin. Median septum almost as
wide as long, placed almost entirely inside and fused to median area of atrium, which is restricted to only two free
lateral areas. Anterior base of septum entirely within atrium and slightly larger than posterior base, which is located
just below atrium. Copulatory openings clearly visible, located in depressions at each side of median portion of
septum. Vulva (Fig. 2F, G) with short ventral ducts, connected posteriorly to copulatory openings and anteriorly to
internal posterior edge of large reniform anterior curve, which is very expanded. From external posterior angle of

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anterior curve, a long duct originates, directed internally at its initial portion, after which there is a closed curve,
directed ventrally and contouring internal margin of spermatheca, reaching down close to ventral surface. A 90o
curve follows, directed posteriorly, forming a long and almost straight path, clearly visible by transparency (Fig.
2D), and reaching back to near epigastric furrow, then an abrupt and long curve directed dorsally occurs, followed
by another one directed anteriorly, originating a short straight path and a curve directed externally, which finally
connects to internal posterior margin of large and roundish spermatheca.
Coloration. Carapace yellow with a brown median longitudinal stripe, bifurcate posteriorly, not reaching
margin of carapace. Light brown irregular stripes at margins. Ocular mound brown. Sternum, endites and labium
mostly light yellow. Labium and sternum with light brown margins. Chelicerae light yellow with brown median
stripe at each one. Legs mostly yellow with light brown irregular spots. Legs III and IV lighter than I and II and
with less spots. Opisthosoma mostly yellow, lighter in folium and conical projections, where there are visible
guanine crystals. Dorsally, one brown median stripe reaching up to posterior conical projection, and two black
roundish spots near to base of each paired conical projections of opisthosoma. Laterally, black irregular spots, and
ventrally yellow and white hues.
Description. Male (MPEG 31328): Total length: 2.48. Carapace: length 1.27, width 1.21, height 0.71.
Clypeus: height 0.15. Chelicera: length 0.37, width 0.17. Sternum: length 0.54, width 0.52. Endites: length 0.15,
width, 0.18. Labium: length 0.06, width 0.17. Eye diameters and interdistances: AME 0.05, ALE 0.06, PME 0.05,
PLE 0.04, AME–AME 0.01, AME–ALE 0.06, ALE–ALE 0.18, PME–PME 0.11, PME–PLE 0.07, PLE–PLE 0.31.
MOQ trapezoidal. Legs 1243. Leg I: femur 1.11, patella 0.47, tibia 0.94, metatarsus 0.82, tarsus 0.46, total 3.80; II:
1.02-0.44-0.90-0.70-0.44-3.50; III: 0.50-0.28-0.43-0.33-0.27-1.81; IV: 0.62-0.32-0.53-0.39-0.30-2.13. Paired setal
tubercles in tibiae and metatarsi near half to almost as long as the width of the article. Opisthosoma: length 1.21,
width 1.67, height 1.28, with trapezoidal anterior half and semicircular posterior half in dorsal view, covered by
abundant macrosetae of variable size. Anterior margin of dorsum straight, slightly projecting over posterior portion
of carapace, with anterior angles not projected. Paired projections in median portion of opisthosoma, directed
posterolaterally. Posterior conical projection placed more posteriorly than in female, and directed both dorsally and
posteriorly. In lateral view, dorsum very convex both before and behind posterior conical projection.
Palpus (Figs 3 D–H). Tegulum with transversal furrow in 3 o’clock position. Embolus basis gradually
separating from tegulum, initially forming a wide stripe, that tapers up to filiform apex, describing about 3/4 of a
turn around lateral margin of tegulum, and with apex located on a furrow placed on prolateral margin glabrous
apical area of cymbium. RTA fused to DTA, both connected directly to tibia and not constricted at base. Short V-
shaped notch between RTA and DTA. RTA wide and very elongated, about 3x longer than wide, with curved lateral
margins, arranged transversally to longitudinal axis of cymbium, large median concavity, wide marginal folds, and
rounded apex thinner than its median portion. RTA with longitudinal axis very slanted downward and also
projecting retrolaterally, especially at its distal half. Its dorsal fold describes a long and convex curve and projects
over median concavity, forming a deep marginal trough. Ventral fold describes a S-shaped curve, with concave
distal portion and convex basal portion, projected over median depression only at basal area, where it extends
dorsally and fuses to common base RTA + DTA, forming a deep basal pouch around DTA and dorsal fold of RTA.
Basal pouch conspicuous in ventral view and with evident grooves, numerous and close to each other along its
length, which give a striated aspect to surface. DTA very short, horn shaped, a little blunt, with slightly thicker
base. Dorsum of tibia with large setal tubercle, surmounted by a very strong spiniform macroseta, inclined
retrolaterally.
Coloration. Carapace orange yellow with a wide longitudinal dark brown median stripe, bifurcated at its
posterior portion, and with irregular dark brown stripes at margins. Sternum entirely yellow; endites and labium
predominantly yellow with brown margin; chelicerae light yellow with a median longitudinal brown stripe near the
fang. Legs predominantly yellow with irregular light brown rings and spots. Legs III and IV lighter than I and II,
and with less spots. Opisthosoma pale brown, dorsum with a trapezoidal brown folium between a white area
stained by guanine. Conical projections dark yellow, also with basal guanine spots. Sides predominantly pale
brown with black hues and venter pale brown with yellow hues.
Distribution. Only know from eastern Amazon, from the Brazilian states of Pará and Maranhão (Fig. 15).

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FIGURE 2. Epicadinus biocellatus Mello-Leitão, 1929. A–G female (MPEG 31330); H female (MPEG 34862). A–C habitus
(A dorsal, B lateral, C ventral); D–E, epigynum (D photo ventral, E SEM ventral); F–G vulva (F photo ventral, G dorsal); H
live specimen in hunting position. Abbreviations, A, atrium; AC, anterior curve of copulatory duct; CO, copulatory opening;
FD, fertilization duct; MS, median septum. Scale bars: A–C, 1 mm; D–G, 0.1 mm. Photo credits: H, César Favacho.

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FIGURE 3. Epicadinus biocellatus Mello-Leitão, 1929, male (MPEG 31328). A–C habitus (A dorsal, B lateral, C ventral); D–
H left male palpus (D ventral, E–F retrolateral, G slanted retrolateral, H cymbium apex, ventral). Abbreviations, BP, basal
pouch; DTA, distal tibial apophysis; E, embolus; EA, embolus apex; MF, marginal furrow of cymbium; RTA, retrolateral tibial
apophysis; TF, tegular furrow; TM, tibial macrosetae. Scale bars: A–C, 1 mm; D–H, 0.1 mm.

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FIGURE 4. Epicadinus biocellatus female holotype (MHNH AR-15077, Coll. Simon 2044b) [total length: 4 mm]. A dorsal
habitus; B venter of opisthosoma; C–E original labels of the splitted typical lot (C female holotype 2044b, D misidentified male
of E. biocellatus 2044, E female of E. cornutus 2044a). Photo credits: Christine Rollard and MNHN Laboratory of
Arachnology crew.

Epicadinus trispinosus (Taczanowski, 1872)

Figs 5–7, 15

Thomisus cornutus Taczanowski, 1872: 88, pl. 4, fig. 8. New synonymy.

Thomisus trispinosus Taczanowski, 1872: 99.
Stephanopis cornuta (Taczanowski). Keyserling 1880: 183, pl. 4, fig. 100.
Stephanopis trispinosus (Taczanowski). Keyserling 1880: 184, pl. 4, fig. 101.
Eripus trifidus O. Pickard-Cambridge, 1893: 121, pl. 15, fig. 12. New synonymy.
Epicadinus trifidus (O. Pickard-Cambridge). Simon 1895: 1052; F. O. Pickard-Cambridge 1900: 159, pl. 10, fig. 33.
Epicadinus trispinosus (Taczanowski). Simon 1895: 1052; Mello-Leitão 1929: 99 (in part).; Machado et al. 2017: 454, figs
S8A, S10D.
Tobias cornutus (Taczanowski). Simon 1895: 1053.
Epicadinus cornutus (Taczanowski). Mello-Leitão 1929: 105.

Type-material: Holotype of Epicadinus cornutus: female, FRENCH GUIANA, Cayenne, 1868–1871, C. Jelski
(MIZ 500 001, Figs 7A–C), photos examined. Holotype of Epicadinus trispinosus: male, FRENCH GUIANA,
Cayenne, 1868–1871, C. Jelski (MIZ 500 000, Figs 7D–G), photos examined. Holotype of Epicadinus trifidus:
female, MEXICO, Veracruz, Atoyac, H. H. Smith (OUMNH 1278.2, ex coll. Goldman & Salvin, not found).
Additional material examined. BOLIVIA: 1 male, “chaco” (MNHN coll. Simon 18020). BRAZIL: 1
female, December 1919, H. S. Parrish (MCZ). Acre: 1 male, Cruzeiro do Sul, Parque Nacional da Serra do Divisor,
ca. 8°23'10"S, 73°09'30"W, 15 March 1997, L. Resende & S. Vieira (IBSP 12368). Amazonas: 1 male, alto [Rio]
Solimões, ca. 4°20'50"S, 67°33'22"W, December 1979-1980, A. Lise et. al. (FZBRS 8939); 1 female, Tefé, ca.
3°19'09"S, 64°42'30"W, December 1919, H. S. Parrish (MCZ). São Paulo: 7 females, Primavera, Usina
Hidrelétrica Sérgio Motta, ca. 22°29'21"S, 52°57'06W, January–February 2000, IBSP (IBSP 29882). ECUADOR:
Orellana: 1 female, 2 juveniles, Parque Nacional Yasuní, ca. 1°05'48"S, 75°48'44"W, 2–5 December 2009, L. R.
Benavides (UFMG 3922). MEXICO: Campeche: 1 male, 1 female, "El tormento" forest station, 18°37'00"N,
90°48'00"W, 11–12 July 1983, W. Maddison (MCZ). Chiapas: 1 female, Palenque Ruins, ca. 17°28'36"N,

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92°02'59"W, 2–11 July 1983, R. S. Anderson (MCZ). Tabasco: 1 female, Grutas de Coconá, 17°33'00"N,
92°55'59"W, 7 July 1983, W. Maddison (MCZ). PANAMA: Arraiján: 1 male, 1 female, ca. 8°58'49"N,
79°40'24"W, 6 July 1950, A. M. Chickering (MCZ). Canal Zone: 3 males, Barro Colorado Island, ca. 9°09'09"N,
79°50'46"W, May 1964, A. M. Chickering (MCZ); idem, 1 male, 2 juveniles, 6 February 1958 (MCZ); idem,
1juvenile 29 June 1939, (MCZ); idem, 1 male July 1939 (MCZ); idem, 3 females, June 1950 (MCZ); idem, 3
females, 20 February 1950 (MCZ); idem, 1 female, 04 August 1939 (MCZ); 1 female, Exp. Gardens, ca.
9°04'24"N, 79°38'49"W, 26 July 1964, A. M. Chickering (MCZ); idem, 2 females, 13 July 1954 (MCZ); 1 female,
Forest Reserve, ca. 9°09'26"N, 79°45'21"W, 25 July 1954, A. M. Chickering (MCZ); idem, 1 male, 1 female, 8
January 1958, (MCZ); idem, 1 juvenile, 23 July 1950 (MCZ); idem, 1 male, 14 February 1958 (MCZ). PERU:
Loreto: 1 female, Pebas, [M. de Mathan] (MHNH, Coll. Simon 2044a). TRINIDAD Y TOBAGO: 1 juvenile,
Balandra Bay, ca. 10°40'36"N, 61°30'57"W, April 1922, Reynolds (MCZ); 1 juvenile, Port of Spain, ca.
10°43'13"N, 61°00'05"W, no date, R. Thaxter (MCZ); 1 male, Simla, ca. 10°21'37"N, 61°18'06"W, 16 April 1964
(MCZ).
Diagnosis. E. trispinosus belongs to the trispinosus-group and is similar to E. biocellatus. Females of E.
trispinosus are distinguished from E. biocellatus by the carapace presenting a longitudinal, median yellowish stripe
between two light brown stripes, and by the opisthosoma with posterior triangular portion, anterior margin slightly
concave and projected on the carapace, and posterior conical tubercle much higher than the opisthosoma (Fig. 5C).
The epigynum is longer than wide, with a deep and well delimited atrium, a median septum and copulatory
openings located at the edges of the epigynal plate (Fig. 5D, E). Its copulatory ducts do not contour the
spermathecae margin (Fig. 5F). Males of E. trispinosus are distinguished from E. biocellatus by their reddish-
brown carapace and by a light strip separating two dark brown stripes, femur and patella I and II brownish,
contrasting strongly with the other yellowish articles, and the pentagonal opisthosoma is distinctive by the posterior
conical projection smaller than the lateral pair (Figs 6A, C). RTA is approximately 2x longer than wide, with
ventral fold margin describing a long convex curve, very conspicuous basal pouch in ventral view, and very long,
hook-shaped DTA (almost as long as RTA, Figs 6F, G).
Notes. Taczanowski (1872) based Thomisus cornutus on a female holotype (Figs 7A–C), deposited in the MIZ.
In the same work, eleven pages later, the author described Thomisus trispinosus (type-species of Epicadinus) based
on a male holotype (Figs 7D–G). The two specimens are from Cayenne, French Guiana. The analysis of abundant
material from western Brazil and bordering countries, as well as Central America and Mexico, revealed the
presence of only one male morphotype, corresponding to E. trispinosus, and one female morphotype, matching
with E. cornutus, often occurring in the same locality or in nearby areas. In addition to the congruence of localities,
the longitudinal median yellowish stripe in the carapace and the compatibility of genital features (short embolus
and short copulatory ducts, long ATD in palpus and deep lateral fovea at the epigynum strongly suggests synonymy
between these species. As both species have been described in the same paper, we had to apply the principle of the
first reviewer (ICZN 1999: article 24.2). Notwithstanding the page priority criterium, we choose to keep
Epicadinus trispinosus as the valid name over its synonym Epicadinus cornutus. Therefore, the valid name of the
type species of Epicadinus remains Epicadinus trispinosus for the sake of the stability and universality of
nomenclature, following ICZN Recommendation 24A.
The specimen described by Mello-Leitão (1929) as the female of E. trispinosus does not belong to this species,
as he indicated that the three opisthosomal projections were of similar size. It may belong to E. biocellatus, but we
are not sure as we had not found any female specimen identified by Mello-Leitão as E. trispinosus. Moreover, the
short description this author provided does not allow a precise identification. Another species described for the area
of occurrence of E. trispinosus is Eripus trifidus, based on a holotype female from Veracruz, Mexico. In spite of the
disappearance of the holotype, the extensive original description and the good illustrations of Eripus trifidus given
in the “Biologia Centrali Americana" (O. Pickard-Cambridge 1893: fig. 12) allowed the proposed synonymy with
Epicadinus trispinosus. This conclusion is corroborated by the distinct shape of the long and vertically oriented
median opisthosomal projection. Besides, the only species of the genus we have been able to found in Mexico and
Central America is E. trispinosus, represented by abundant material from both sexes.
Description. Female (UFMG 3922, Figs 5A–C). Total length: 3.91. Carapace: length 2.07, width 2.05, height
1.60. Clypeus: height 0.20. Chelicerae: length 0.38, width 0.28. Sternum: length 0.96, width 0.87. Endites: length
0.24, width, 0.19. Labium: length 0.07, width 0.35. Eye diameters and interdistances: AME 0.06, ALE 0.07, PME
0.05, PLE 0.04, AME–AME 0.13, AME–ALE 0.10, ALE–ALE 0.31, PME–PME 0.17, PME–PLE 0.09, PLE–PLE

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0.48. MOQ trapezoidal. Legs 1243. Leg I: femur 2.41, patella 0.88, tibia 2.06, metatarsus 1.58, tarsus 0.72, total
7.67; II: 2.34-0.97-1.77-1.41-0.79-7.30; III: 0.87-0.66-0.74-0.54-0.44-3.27; IV: 0.94-0.62-0.78-0.60-0.46-3.42.
Paired macrosetal tubercles in tibiae and metatarsi not especially long or robust, shorter than half the width of the
article. Opisthosoma: length 1.84, width 2.72, height 3.41, pentagonal, ending in a short terminal projection
carrying the spinnerets (due to the posterior tubercle oriented perpendicularly). Anterior margin of opisthosoma
slightly concave and with anterior angles slightly projected over the posterior slope of the carapace. Paired conical
projections with acute apex directed posterolaterally. Posterior conical projection more than twice as long as paired
ones, pointing vertically and much higher than the opisthosoma itself, arranged perpendicularly to venter of the
opisthosoma. In lateral view, dorsum with anterior portion strongly concave up to the posterior conical tubercle and
posterior portion slightly concave.
Epigynum (Fig. 5D, E) longer than wide. The ellipsoid atrium is about ¾ wider than long, located anteriorly,
with a deep central depression and two lateral foveas, which are deeper than central area. Each fovea is covered by
a thin, translucent chitinous keel, which tapers toward posterior portion, where it disappears. Median septum
almost as wide as long, totally fused to surface of plate with anterior basis extending into the atrium and wider than
posterior base, which is located a little behind the copulatory pores and distant from the atrium. Copulatory
openings exposed, appearing as deep pits. Epigynum bordered by cilium-shaped bristles. Vulva (Fig. 5F) with
slightly sclerotized ventral copulatory ducts, which expand to form a large sclerotized, very dilated and reniform
anterior curve (secondary spermatheca). From the external lower angle of this structure, a sinuous duct originates,
curved initially in ventral and internal direction and with the final part curved dorsolaterally , connecting to the
inferior internal margin of the spermathecae, that are rounded and well developed.
Coloration (Figs 5A–C). Carapace yellow with two light brown longitudinal stripes, separated by a thin stripe
of the carapace color, and light brown irregular marginal stripes. Sternum, endites and labium predominantly light
yellow. Light yellow chelicerae each with a medium longitudinal light brown stripe. Legs I and II uniformly light
yellowish. Legs III and IV predominantly light yellow with irregular brown spots. Opisthosoma predominantly
yellow, including the three conical projections, whose tips are white, due to guanine spots. Dorsally, a dark yellow
median stripe between two marginal white stripes (guanine) and two rounded black spots near the bases of each of
the paired conical projections in the median portion of the opisthosoma. Ventrally, predominantly yellow and with
white hues, sigils brown.
Male (FZBRS 8939, Figs 6A–C): Total length: 2.64. Carapace: length 1.08, width 1.29, height 0.78, with
three rows of median macrosetae and six rows of macrosetae placed between the radial furrows, besides other
irregularly distributed macrosetae. Posterior portion of the carapace with a small marginal area without macrosetae.
Clypeus: height 0.12. Chelicera: length 0.30, width 0.21. Sternum: length 0.58, width 0.87. Endites: length 0.23,
width, 0.18. Labium: length 0.09, width 0.18. Eye diameters and interdistances: AME 0.05, ALE 0.06, PME 0.05,
PLE 0.03, AME–AME 0.01, AME–ALE 0.06, ALE–ALE 0.23, PME–PME 0.11, PME–PLE 0.07, PLE–PLE 0.34.
MOQ trapezoidal. Legs 1243. Leg I: femur 1.08, patella 0.42, tibia 0.90, metatarsus 0.67, tarsus 0.45, total 3.52; II:
1.00-0.47-0.73-0.56-0.44-3.20; III: 0.54-0.25-0.46-0.34-0.33-1.92; IV: 0.63-0.29-0.53-0.33-0.30-2.08. Paired
macrosetal tubercles in the tibiae and in metatarsi, with length less than or equal to half the width of the article.
Opisthosoma: length 1.31, width 1.84, height 1.42, pentagonal, covered by abundant macrosetae of variable size.
Anterior margin of opisthosoma almost straight and barely projecting on the posterior portion of the carapace.
Paired conical projections bigger than the posterior one, placed in the median portion, arranged transversally, with
their acute apex directed posterolaterally. Posterior conical projection smaller than median ones, measuring about
1/3 of the height of opisthosoma. In lateral view, dorsum slightly convex, almost flat, with all projections directed
back and upwards and the median one less inclined than the others.
Palpus (Figs 6D–G) with tegular furrow at 4 o’clock position. Embolus basis gradually separating from
tegulum, initially forming a wide stripe, that becomes gradually thinner up to the filiform apex, describing about 3/
5 of a turn around the lateral margin of the tegulum, but with the apex located on a furrow placed on the prolateral
margin of the glabrous apical area of the cymbium. RTA fused to DTA, both connected directly to the tibia. An
openwide U-shaped angle placed between the RTA and DTA. RTA wide and not much elongated, about 2x longer
then wide, almost parallel to the longitudinal axis of the cymbium in retrolateral view, especially in its distal half,
with wide median concavity and wide marginal folds, and a curved apex, elongated and thinner than the median
portion of RTA. Dorsal fold projected on median concavity, forming a marginal trough in its distal half. Ventral
fold large describing a long convex curve, projected gradually over the median concavity at its basal area, where it

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FIGURE 5. Epicadinus trispinosus (Taczanowski, 1872), female (UFMG 3922). A–C habitus (A dorsal, B ventral, C lateral);
D–E epigynum ventral; F vulva dorsal). Abbreviations, A, atrium; AC, anterior curve of copulatory duct; CD, copulatory duct;
CO, copulatory opening; MS, median septum; TK, translucent keel. Scale bars: A–C, 1 mm; D–F, 0.1 mm.

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FIGURE 6. Epicadinus trispinosus (Taczanowski, 1872), male (FZBRS 8939). A–C habitus (A dorsal, B ventral, C lateral);
D–G left male palpus: (D–E ventral, F retrolateral, G tibial apophysis retrolateral). Abbreviations, BP, basal pouch; DTA, distal
tibial apophysis; E, embolus; EA, embolus apex; RTA, retrolateral tibial apophysis; TF, tegular furrow. Scale bars: A–C, 1 mm;
D–G, 0.1 mm.

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extends dorsally, forming a deep, wide basal pouch around the ventral margin of RTA and only part of base of
DTA. Basal pouch of RTA very conspicuous in ventral view and with well separated and evident grooves along its
extension, which confer a scaly aspect to its surface. DTA very long, hook-shaped and slightly blunt, with the base
much thicker than the apex and inner face covered with numerous grooves very close together.
Coloration. Carapace reddish-brown with a median pair of dark brown longitudinal stripes, separated by a thin
stripe of the same shade of the carapace, and dark brown marginal irregular stripes. Sternum, endites and labium
predominantly yellow. Yellowish chelicerae each with a medium longitudinal light brown stripe. Legs I and II with
patella and distal half of the femur predominantly dark brown, tibia and metatarsus light yellow and brown tarsus.
Legs III and IV predominantly yellow and with irregular dark brown spots. Opisthosoma with a median dark
yellow pyriform stripe between two marginal white stripes and two black rounded spots on its dorsal median
portion. Conical projections mostly reddish brown, but yellow at its lower side. Laterally yellow and white-guanine
hues predominate, and ventrally yellow and brown hues.
Distribution. Epicadinus trispinosus has the largest distribution in the genus, occurring from Mexico, through
Trinidad and Tobago, French Guiana, Ecuador, Peru and Bolivia, to Brazil, where it reaches south to northwestern
São Paulo state (Fig. 15).

FIGURE 7. A–C Epicadinus cornutus (Taczanowski, 1872), female holotype (MIZ 500 001) (A dorsal habitus, B lateral
habitus, C original label). D–G Epicadinus trispinosus (Taczanowski, 1872), male holotype (MIZ 500 000) (D dorsal habitus, E
cephalotorax ventral, F left palpus retrolateral, G original label). Scale bars: A, B, D 1 mm; E, F 0.1 mm. Photo credits: A–G,
Wioletta Wawer and MIZ Laboratory of Arachnology crew.

Epicadinus spinipes (Blackwall, 1862)

Figs 8–10, 15

Eripus spinipes Blackwall, 1862: 422.

Epicadus spinipes (Blackwall). Petrunkevitch 1911: 405.
Epicadinus espinipes (Blackwall) (sic). Mello-Leitão 1929: 100.
Epicadinus albimaculatus Mello-Leitão, 1929: 104. New synonymy.
Epicadinus gavensis Soares, 1946: 267, figs 1–2. New synonymy.

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Type-material. Holotype of Epicadinus spinipes: female, BRASIL, Rio de Janeiro: Serra dos Órgãos, 1857,
Clark, H. & Gray, J. (OUMNH 1277.3, Figs 10A–C), examined. Lectotype of Epicadinus albimaculatus: male,
BRASIL, Rio de Janeiro: Rio de Janeiro city (MNHN AR-15076, Coll. Simon 7.319, Figs 10D–F), examined.
Syntypes of Epicadinus gavensis: male lectotype and male paralectotype (herein designated), BRASIL, Rio de
Janeiro: Rio de Janeiro city, Gávea, 1944, P. Wygodzinsky (MZUSP 3547, Figs 10G–I), examined.
Additional material examined. BRASIL: Espírito Santo: 1 male, Anchieta, CSV [Companhia Siderúrgica
Vitória], Macega, ca. 20°47'22"S, 40°38'40"W, 20 August 2008, R. Baptista (UFRJ 1340); idem, 1 female, 24 April
2009, G. A. Pérez (UFRJ 1341); 2 males, Linhares, Caliman, Fazenda Santa Terezinha, ca. 19°07'91.1"S,
39°58'43.5"W, 30 November 2011, R. Baptista (UFRJ 1335); 1 female, 1 juvenile, Linhares, Reserva Natural Vale,
Bicuíba, ca. 19°09'11"S, 40°02'50"W, 4 February 2011, D. T. Castro (UFRJ 1339); idem, 2 females, R. Baptista
(UFRJ 1337); idem, 1 female, 22 April 2011, D. T. Castro, (UFRJ 1336); idem, 1 juvenile, 22 April 2011, R.
Baptista (UFRJ 1332); idem, 3 juveniles, 4 August 2011, R. Baptista (UFRJ 1338); idem, 1 juvenile, 28 November
2011, D. T. Castro (UFRJ 1333); 1 juvenile, Sooretama, REBIO Sooretama, Quirinão, ca. 18°59'35"S, 40°08'58"W,
23 April 2011, D. T. Castro (UFRJ 1331); 1 juvenile, 23 April 2011, R. Baptista (UFRJ 1334). Rio de Janeiro: 1
male, Angra dos Reis, Ilha Grande, Vila Abraão, ca. 23°08'32"S, 44°10'26"W, February 1994, M. Ramirez (MNRJ
[ex MACN]); 1 male, Casemiro de Abreu, Barra do Sana, ca. 22°22'25"S, 42°12'14"W, 28 December 2002,
Wienskoski (MNRJ 11484); 1 juvenile, Macaé, Terminal Cabiúnas, Mata da Fazenda, ca. 22°17'18"S, 41°44'05"W,
16 March 2011, R. Baptista (UFRJ 1316); 1 male, Macaé, Terminal Cabiúnas, Mata do Gasoduto, ca. 22°17'18"S,
41°44'05"W, 11 December 2012, G. Miranda (UFRJ 1320); idem, 1 juvenile, 23 May 2013 (UFRJ 1315); idem, 1
female, 21 February 2014, M. O. Villareal (UFRJ 1318); idem, 1 juvenile, 21 February 2014, D. R. Pedroso (UFRJ
1317); 1 juvenile, Macaé, Terminal Cabiúnas, Mata da Odebei, ca. 22°17'09"S, 41°44'06"W, 17 March 2011, R.
Baptista (UFRJ 1322); idem, 1 juvenile, 24 February 2013, G. Miranda (UFRJ 1321); 1 juvenile, 21 November
2013 (UFRJ 1319); 2 males, Mangaratiba, Rio das Pedras, ca. 22°59'23"S, 44°05'47"W, 12 December 2006,
Wienskosk (MNRJ 06865); 2 males, 1 juvenile, Mendes, Centro Marista São José das Paineiras, Sede, 22°30'35"S,
43°45'16"W, 21 November 2014, R. Baptista (UFRJ 1327); idem, 1 male, 1 female, 1 juvenile, 21 November 2014,
A. W. Prado (UFRJ 1328); idem, 1 female, 21 November 2014, R. Baptista (UFRJ 1326); 1 male, 18 November
2014, A. W. Prado (UFRJ 1324); 2 juveniles, Mendes, Fazenda Arvoredo, Encruzilhada, 22°30'17"S, 43°45'35"W,
31 August 2014, R. Baptista (UFRJ 1323); idem, 1 male, 17 November 2014, R. Baptista (UFRJ 1329); 1 male,
Mendes, Fazenda Arvoredo, Porteira, 22°30'10"S, 43°45'14"W, 4 March 2015, A. W. Prado (UFRJ 1325); 1 male,
Rio de Janeiro, Parque Estadual da Pedra Branca, Pau da Fome, Padaria, 22°56'13"S, 43°26'29"W, 23 September
2013, P. Castanheira (UFRJ 1343); 1 juvenile, Rio de Janeiro, Parque Estadual da Pedra Branca, Pau da Fome,
Figueira, 22°54'40"S, 43°30'25"W, 12 June 2013, R. Baptista (UFRJ 1342); 1 juvenile, Santa Maria Madalena,
Desengano, ca. 21°57'14"S, 42°00'56"W, 23–25 March 1998, A. B. Kury (MNRJ 10388). São Paulo: 1 male, São
Sebastião, Praia de Guaecá, ca. 23°49'20"S, 45°27'07"W, September 1998, C. A. Rheims (IBSP 20588).
Diagnosis. E. spinipes belongs to the spinipes-group and resembles E. villosus. Females of E. spinipes are
distinguished from E. villosus by the presence of a yellowish, thin, median stripe along the carapace, located
between two dark brown stripes, and two yellowish white lateral stripes on the dorsum of the opisthosoma (Fig.
8A), and yellowish sternum, sometimes with pale brown marginal spots (Fig. 8B). The paired macrosetal tubercles
of the tibia and metatarsus are robust, with length greater or equal to half the width of the article (Fig. 8G). The
epigygum presents a remarkable triangular median septum, with anterior base at least twice as wide as the posterior
one (Fig. 8E), and spiral copulatory ducts with approximately ten coils, the last one partially covering the anterior
surface of the spermathecae (Fig. 8F). Males of E. spinipes are distinguished from E. villosus by the yellowish
sternum as in the female (Fig. 9C), RTA about twice as long as wide, slanted to the axis of the cymbium (Fig. 9F),
and embolus with about four turns around the tegulum.
Notes. The type-locality of Eripus spinipes, according to its original description, is Serra dos Órgãos in the
surroundings of Rio de Janeiro city. Clark & Gray's journey crossed the municipalities of Teresópolis and
Petrópolis in the Organ Mountains (Clark 1867: 114–173). The two British collectors made more extensive
collections at Constance, an old farm in Teresópolis (Clark 1867: 118–132, Johnson 2015: 9) and Presidency, the
residence of a British immigrant, about 5 km from Petrópolis (Clark 1867: 148–155). Therefore it is not possible to
specify the precise type-locality. According to Mello-Leitão (1929), Epicadinus albimaculatus was based on
females from Terra Nova, Bahia state (near the Baía de Todos os Santos), and the municipality of Rio de Janeiro,
Rio de Janeiro state, both in Brazil, under number 7329 of Coll. Simon. The only specimen of Epicadinus

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FIGURE 8. Epicadinus spinipes (Blackwall, 1862), female (UFRJ 1326). A–C habitus (A dorsal, B ventral, C lateral); D–E
epigynum ventral; F vulva dorsal; G left leg I highlighting the large paired setiferous tubercles. Abbreviations, A, atrium; CD,
copulatory duct; MS, median septum; TK, translucent keel; PST, paired setiferous tubercles. Scale bars: A–C, G, 1 mm; D–F,
0.1 mm.

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albimaculatus found at MNHN is a female from Rio de Janeiro, clearly labelled as “Type” in Mello-Leitão´s
handwriting (Figs 10D–F), erroneously numbered 7319 of Coll. Simon by later curators. This specimen is herein
considered as the lectotype of E. albimaculatus (ICZN 1999, art. 74.7.3). In color and shape of the epigynum, the
lectotype of E. albimaculatus agrees perfectly with abundant specimens of E. spinipes from the city of Rio de
Janeiro and nearby localities. The other syntype from Terra Nova is considered lost. Besides, Terra Nova is situated
in the coastal area of the Atlantic Forest, a preferred area for E. spinipes. Although Terra Nova is the only record of
the species for Bahia, it is close to Sooretama, in the area of the Tabuleiros Forest, northern Espírito Santo state,
near the border with Bahia, where specimens of E. spinipes were collected. The two males of E. gavensis (Fig.
10G–I) also come from the city of Rio de Janeiro and agree perfectly with the specimens of E. spinipes. We choose
and herein designate the specimen that is best preserved and was illustrated in the original description as the
lectotype (ICZN 1999, recommendation 74B). In addition, the only species of Epicadinus collected in the city of
Rio de Janeiro and surrounding areas and throughout the coastal region of the state of Rio de Janeiro is E. spinipes.

FIGURE 9. Epicadinus spinipes (Blackwall, 1862), male (UFRJ 1324). A–C habitus (A dorsal, B lateral, C ventral); D–F, left
male palpus (D–E ventral, F retrolateral). Abbreviations, DTA, distal tibial apophysis; E, embolus; RTA, retrolateral tibial
apophysis; TF, tegular furrow. Scale bars: A–C, 1 mm; D–F, 0.1 mm.

Description. Female (UFRJ 1326, Figs 8A–C, G): Total length: 4.41. Carapace: length 2.00, width 1.77,
height 1.11. Clypeus: height 0.23. Chelicera: length 0.50, width 0.32. Sternum: length 0.98, width 0.89. Endites
very setaceous, especially in the retrolateral portion: length 0.28, width, 0.25. Labium: length 0.28, width 0.43. Eye

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diameters and interdistances: AME 0.09, ALE 0.11, PME 0.06, PLE 0.05, AME–AME 0.013, AME–ALE 0.09,
ALE–ALE 0.33, PME–PME 0.19, PME–PLE 0.13, PLE–PLE 0.56. MOQ trapezoidal. Legs 1234. Leg I: femur
2.68, patella 1.14, tibia 2.15, metatarsus 1.67, tarsus 0.99, total 8.63; II: 2.00-1.16-1.89-1.23-0.71-6.99; III: 1.05-
0.70-0.94-0.66-0.50-3.85; IV: 1.06-0.58-0.90-0.68-0.49-3.71. Paired macrosetal tubercles very robust in tibiae and
metatarsi, longer than or equal to half width of the article. Opisthosoma: length 2.55, width 2.98, height 2.56,
pentagonal. Anterior margin of dorsum concave in dorsal view, projecting over posterior portion of carapace, with
projected anterior angles. Pair of lateral conical projections with acute apex directed posterolaterally, placed
transversally in the median portion, and the posterior one placed in a very slanted position, a little longer than the
paired projections and reaching 2/3 of the height of the opisthosoma. In lateral view, anterior portion of the dorsum
nearly flat, aligned to the dorsal face of the posterior conical projection, and posterior portion slightly convex.
Epigynum (Figs 8C, D) longer than wide, with reniform atrium, approximately twice as wide as long, located
anteriorly, formed by a deep depression, deepest at level of lateral pair of foveas, whose margins are covered by a
thin translucent chitinous keel. Anterior portion of atrium rises gradually reaching surface level of opisthosoma.
Triangular median septum, almost as long as wide, with an anterior base at least twice as wide as posterior base and
projecting itself laterally, merging into interior part of atrium. The posterior base fuses to area near to epigastric
furrow, placed far from atrium. Median portion of septum very high, covering copulatory openings and posterior
margins of atrium. Margins of epigynum present long cilium-shaped bristles, which partially cover the atrium.
Vulva (Fig. 8F) with very long spiral copulatory ducts and a pair of posterior rounded spermathecae. Copulatory
ducts initially formed by a simple, curved duct, followed by a spiral portion directed anteriorly, with approximately
nine coils, lighter in hue, ending in a more dilated anterior portion, whose inner margin originates an inner darker
spiral portion. This darker spiral, formed by approximately ten coils, runs posteriorly in opposite direction from
first spiral with a thicker posterior portion, whose last coil partially covers the dorsal surface of the spermathecae,
and forms a sharply curved distal duct connected to the posterior inner face of the rounded and well-developed
spermathecae.
Coloration. Carapace yellow, with two greyish brown stripes, separated by a thin stripe of the carapace hue,
and greyish brown radial stripes. Sternum, endites and labium light yellow. Chelicerae greyish brown. Legs I and II
of the same color as the carapace, covered with irregular greyish brown spots, legs III and IV lighter and with few
spots. Opisthosoma greyish yellow with two median yellowish white (guanine) longitudinal stripes bordered by
black irregular lines; each opisthosomal projection bears one blackish spot at the basis, followed by a blackish grey
stripe up to the light yellow tip. Venter light yellow, sides with black irregular spots near the basis of the median
projections. Posterior conical projection of the same hue as the dorsum, with light yellow tip.
Male (UFRJ 1324, Fig. 9A–C: Total length: 2.89. Carapace: length 1.32, width 1.23, height 0.75. Clypeus:
height 0.16. Chelicera: length 0.33, width 0.19. Sternum: length 0.56, width 0.59. Endites very setaceous,
especially in the retrolateral portion: length 0.22, width, 0.26. Labium: length 0.09, width 0.20. Eye diameters and
interdistances: AME 0.07, ALE 0.07, PME 0.05, PLE 0.04, AME–AME 0.08, AME–ALE 0.05, ALE–ALE 0.16,
PME–PME 0.07, PME–PLE 0.04, PLE–PLE 0.29. MOQ trapezoidal. Legs 1243. Leg I: femur 1.30, patella 0.49,
tibia 0.96, metatarsus 0.82, tarsus 0.58, total 4.15; II: 1.12-0.52-0.95-0.79-0.52-3.90; III: 0.54-0.25-0.49-0.37-0.31-
1.94; IV: 0.59-0.32-0.51-0.39-0.30-2.11. Paired macrosetal tubercles in tibiae and metatarsi very robust and long,
longer than or equal to half the width of the article. Opisthosoma: length 1.57, width 1.90, height 1.30, pentagonal.
Anterior margin of opisthosoma a little concave in dorsal view, slightly projecting over the posterior portion of the
carapace, with anterior angles a little projected. Paired conical projections thin, about as long as the posterior one,
and arranged transversally. Posterior conical projection very slanted and not reaching half the height of the
opisthosoma itself. In lateral view, anterior portion of the dorsum nearly flat and almost aligned to the posterior
conical projection, and posterior portion slightly convex.
Palpus (Figs 9D–E). Tegulum with transversal furrow at 1 o’clock position. Embolus basis gradually
separating from tegulum, initially forming a wide stripe, slightly tapering along its median portion, which describes
about 4 turns around the lateral margin of the tegulum, and ends in a filiform apex, hidden by the margin of
tegulum. RTA fused to DTA, both connected directly to the tibia by an elongated and narrow base, forming a kind
of pedestal. RTA about 2x longer than wide, arranged transversally, downward slanted to the longitudinal axis of
the cymbium, with large median concavity, marginal folds, and an apex much thinner than the median portion,
forming an acute and elongated curve. Dorsal fold slightly projected over the median concavity only in the apical
region, fusing directly to the base of the DTA. Ventral fold strongly projected over the median concavity, but

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gradually decreasing towards the apex. DTA slightly elongated, hook-shaped, with a slightly thicker base,
excavated at the inner surface of hook, its apex forming a rounded and slightly acute tip.
Coloration. Carapace reddish brown with two median longitudinal dark brown stripes in the posterior portion.
Sternum light yellow bordered by reddish brown spots. Endites and labium uniformly reddish brown. Reddish
brown chelicerae with irregular black spots. Legs I and II with the same color as the carapace, III and IV lighter.
Opisthosoma dorsally yellow with a faint transversal stripe of irregular blackish spots between the conical
projections in the median portion. Conical projections reddish brown. Opisthosoma ventrally and laterally yellow
with several black spots near macrosetae. Basis of macrosetae appearing as reddish brown dots.
Distribution. Most records of E. spinipes are from Atlantic Forest areas near the coast of the states of Rio de
Janeiro and Espírito Santo, reaching south to the northeastern coast of São Paulo state, all in southeastern Brazil
(Fig. 15). Its distribution may reach the coastal area of southern Bahia state, northeastern Brazil, as there are
specimens from the Tabuleiros Forest at the border of Espírito Santo and Bahia, and one of the type-localities of the
species is Terra Nova, near Salvador (albeit we had not examined this lost syntype).

FIGURE 10. A–C Epicadinus spinipes (Blackwall, 1862), female holotype (OUMNH 1277.3) [total length: 8.4 mm] (A dorsal
habitus, B ventral habitus, C original label). D–F Epicadinus albimaculatus female lectotype (MNHN AR-15076, Coll. Simon
7.319) [total length: 7 mm] (D specimen retrolateral, E venter with epigynum, F original labels). G–I Epicadinus gavensis male
syntype (MZUSP 3547) (G dorsal habitus, H left male palpus ventral, I original labels). Scale bars: G, 1 mm; H, 0.1 mm. Photo
credits: D–F, Christine Rollard and MNHN Laboratory of Arachnology crew.

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Epicadinus villosus Mello-Leitão, 1929
(Figs 11–13, 15)

Epicadinus villosus Mello-Leitão, 1929: 103.

Epicadinus helenae Piza, 1936: 50, fig. 1. Soares 1943: 12, fig. 10. New synonymy.
Epicadinus marmoratus Mello-Leitão, 1947: 274. New synonymy.

Type-material: Holotype of Epicadinus helenae: subadult female, BRASIL, São Paulo: Piracicaba, 1936, M.
Diniz (IBSP 138.217 [ex. MZLQ 117], Figs 13D–F), examined. Holotype of Epicadinus marmoratus: juvenile,
BRASIL, Paraná: Curitiba, Barigui, x.1944, R.B. Lange (MNHCI 02496 [Coll. Rudolf B. Lange], Figs 13G–I),
examined. Holotype of Epicadinus villosus: female, BRASIL, Bahia: “S. Antônio da Barra” (= Santo Antônio de
Condeúba, MNHN AR-15078 [Coll. Simon 11.511], Figs 13A–C), examined. Paratypes of Epicadinus villosus:
female, PARAGUAY (MNHN [Coll. Simon 9752]); female, PARAGUAY (MNHN [Coll. Simon 19194]).
Additional material examined. ARGENTINA: Misiones: 1 male ca. 27°25'33"S, 55°56'44"W, February
1950 (MACN 19053); idem, 1 female, October 1953, Schiapelli, De Carlo, Viana, Galiano (MACN 3802); idem, 1
male, October 1953 (MACN 3803); 2 females, General Manoel Belgrano, ca. 26°04'26"S, 53°59'35"W, 1954,
Schiapelli & De Carlo (MACN 19054); 1 female, Montecarlo, ca. 26°33'41"S, 54°46'08"W, January 1966, M. E.
Galiano (MACN 19064); idem, 1 female (MACN 19066); 2 females, Parque Nacional Iguazú, ca. 25°41'08"S,
54°27'39"W, August 1985, M. Ramirez (MACN 19058); idem, 1 female, 8–15 February 1995, M. Ramirez
(MACN 19060); 3 females, Parque Nacional Iguazú, Ruta 101, ca. 25°39'12"S, 54°19'36"W, January 1966,
Galiano (MACN 19065); 1 male, Parque Nacional Iguazú, Area Cataratas, ca. 25°41' 05"S, 54°26'46"W, 13–15
September 1992, Goloboff (MACN 19052); idem, 1 male, November 1989, M. Ramirez & L. Lopardo (MACN
19056); idem, 1 male, 22–30 August 1986, M. Ramirez & L. Lopardo (MACN 19057); idem, 1 female, 11–16
December 1999, M. Ramirez & L. Lopardo (MACN 19062); 1 male, Parque Nacional Iguazú, RN 101, E.
seccional Yacuy, ca. 25°36'27"S, 54°18'52"W, 14–16 December 1999, M. Ramirez & L. Lopardo (MACN 19063);
1 female, San Pedro, Tobuna, ca. 26°28'03"S, 53°53'53"W, February 1952, W. Partridge (MACN 04064).
BRASIL: Minas Gerais: 1 female, Belo Horizonte, Estação ecológica da UFMG 19°52'28"S, 43°58'22"W,
January 2001, E.S.S. Álvares (UFMG 129); idem, 1 male, 1 female, 1 juvenile, 14 November 2009, G.H.F.
Azevedo et al. (UFMG 3389); idem, 1 male, 1 female, x.2000, no data (UFMG 5959); 1 female, Catas Altas, Serra
do Caraça, ca. 20°08'33"S, 43°29'07"W, 23–26 November 1960, Araújo & Martim (MZUSP 13626); 1 male, Sete
Lagoas, Serra Santa Helena, 19° 26' 43"S, 44° 16' 20"W, 26 November 2001, E.S.S. Álvares (IBSP 40263). Pará:
1 male, 1 female, Santarém, ca. 2°27'11"S, 54°41'47"W (MNHN Coll. Simon 16064). Paraná: 1 female, ca.
25°25'35"S, 49°15'59"W, 15 April 2005, J. Ricetti (PUCRS 37233); 1 male, 1 female, Candói, Usina Hidrelétrica
de Segredo ca. 25°47'19"S, 52°06'25"W, April 1996, R. Bertani (IBSP 11665); 1 female, Curitiba, ca. 25°25'35"S,
49°15'59"W, 1938, Pe. F. S. Pereira (MZUSP 00004); 1 male, 1 female, Curitiba, Barigui ca. 25°31'22"S,
49°20'52"W, 20 April 1962, Lange (MHNCI); 1 male, Curitiba, Bom Retiro, ca. 25°24'39"S, 49°16'32"W, 10 April
1987, A. D. Brescovit (FZBRS 16739); 1 female, Curitiba, Parque da Cidade, ca. 25°30'12"S, 49°21'07"W, 1
December 1990, A. D. Brescovit (FZBRS 20627); 4 females, Fênix, Parque Estadual Vila Rica do Espírito Santo,
ca. 23°55'00"S, 51°57'18"W, 22 November 1987, A. D. Brescovit (IBSP 14184); 1 female, 2 juvenile, Foz do
Iguaçu, Parque Nacional do Iguaçu, ca. 25°41'08"S, 54°27'39"W, 29 March 1993, A B. Bonaldo (PUCRS 4310); 7
females, Foz do Iguaçu, Refúgio Biológico de Bela Vista, ca. 25°26'55"S, 54°33'22"W, 17 March 1994, A. B.
Bonaldo (FZBRS 20939); 1 female, Morretes, Serra da Graciosa ca. 25°25'19"S, 48°52'05"W, 9–20 January 1995,
Eq. Lab. Aracnologia (PUCRS 7026); idem, 1 male, 2 females, 2 juveniles (PUCRS 7209); 1 male, 5 females, A.
A. Lise (PUCRS 7347); 1 male, 1 female, São José dos Pinhais, 25°36'18"S, 49°11'37"W, 9 December 2015, A. C.
Domahovski (PUCRS 39104); idem, 1 male, October 2015, A. C. Domahovski (PUCRS 39069); 2 females, 1
juvenile, Três Barras do Paraná, ca. 25°25'14"S, 53°11'04"W, 26 February 1993, A. B. Bonaldo (PUCRS 3035); 1
female, Três Barras do Paraná, Rio Guarani, Foz do Córrego Três Barras, ca. 25°26'28"S, 53°07'23"W, 20-26
February 1993, A. B. Bonaldo (FZBRS 23042). Rio de Janeiro: 1 male, 1 female, Itatiaia, Parque Nacional de
Itatiaia, ca. 22°22'12"S, 44°37'46"W, 28 October 1967 (MACN 19055); 1 female, Mendes, Centro Marista São
José das Paineiras, 22°30'35"S, 43°45'16"W, 21 November 2014, A. W. Prado (UFRJ 1330). Rio Grande do Sul: 1
male, Augusto Pestana, ca. 28°30'51"S, 53°59'40"W, 12 December 2008, L. V. Silva (PUCRS 27091), idem,1
female, 6 September 2009, L. V. Silva & L. B. Medeiros (PUCRS 30592); idem, 1 female, 25 October 2009, L. V.
Silva & L. B. Medeiros (PUCRS 30639); 1 female, Bom Jesus, Fazenda Aver, ca. 28°40'23"S, 50°26'24"W, 24

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March 1989, A. B. Bonaldo (FBZRS 18426), idem, 1 female, 1 April 1988 (FZBRS 17395); 1 male, 1 female,
Cambará do Sul, Itaimbézinho, ca. 29°09'40"S, 50°04'39"W, 28 May 1983, A.A. Lise (FZBRS 11760); 2 males,
Canela, ca. 29°21'22"S, 50°49'10"W, 3 June 1966, A. A. Lise (FZBRS 0238); idem, 1 female, 20 February 1972
(FZBRS 1026); idem, 1 female, 31 December 1973 (FZBRS 2056); idem, 1 female, 2 December 1973 (FZBRS
2207); idem, 2 males, 1 female, 5 juveniles, 26 December 1974 (FZBRS 2497); idem, 2 males, 20 February 1973
(FZBRS 8062); idem, 1 male, 3 females, 1 juvenile, 19 March 1976 (FZBRS 8063); idem, 1 male, 1 female, 4
juveniles, 26 December 1975 (FZBRS 8064); idem, 1 male, 8 November 1975 (FZBRS 8067); idem, 1 female, 20
March 1976 (FZBRS 8069); idem, 1 male, 2 March 1976 (FZBRS 8071); idem, 1 male, 2 March 1976 (FZBRS
8074); idem, 1 male, 1 female, 20 March 1976 (FZBRS 8078); idem, 1 female, 2 March 1977 (FZBRS 8129); 1
female, Usina Hidrelétrica da Canastra, ca. 29°23'39"S, 50°44'45"W, 20 November 1997, M. A. L. Marques
(FZBRS 28844); 1 female, Caxias do Sul, Faz. Souza, ca. 29°09'07"S, 50°57'21"W, 11–12 November 1995, Eq.
Lab. Aracnologia (PUCRS 7320); 7 males, 5 females, Derrubadas, Parque Estadual do Turvo, ca. 27°12'26"S,
53°56'07"W, 28–31 October 2003, R. Ott (FZBRS 37824); idem, 1 male, 1 female, 27–31 October 2003 (FZBRS
38863); idem, 4 females, 4–08 May 2004 (FZBRS 38867), idem, 1 male, 1 female, 19–22 October 2004 (FZBRS
38875); idem, 7 males, 4 females, 19–22 October 2004 (FZBRS 38881); idem, 6 females, 25–30 April 2005
(FZBRS 39201); 2 males, Estrela Velha, Barragem Itaúba, ca. 29°15'40"S, 53°14'26"W, 7 March 2001, R. Ott
(FZBRS 33698); 1 female, Farroupilha, ca. 2913'33"S, 51°20'18"W, 29 September 1978, A. A. Lise (FZBRS
8271); idem, 2 juveniles, H. Bischoff (PUCRS 8272); 1 female, Guaíba, ca. 30°06'24"S, 51°19'27"W, 3 June 1994,
A. A. Lise (PUCRS 4817); 1 female, 1 juvenile, Guaíba, Fazenda São Maximiano, ca. 30°10'53"S, 51°22'56"W, 15
March 1996, A. A. Lise (PUCRS 8286); 2 males, 1 female, Iraí, ca. 27°11'32"S, 53°14'54"W, 21 November 1975,
A. A. Lise (FBZRS 8065); idem, 11 males, 7 females, 10 juveniles (FZBRS 8061); idem, 1 female, 18 November
1975 (FZBRS 8908); 1 female, Itaara, ca. 29°35'40"S, 53°45'34"W, 16 January 2007, A. A. Lise (PUCRS 21344),
idem, 1 female, 13 February 2007 (PUCRS 21344), idem, 1 female, 12 May 2007 (PUCRS 21346); 1 male, 1
female, Itaúba, Arroio do Tigre, ca. 29°16'25"S, 53°9'01"W, 22 April 1978, A. A. Lise (FZBRS 8037), 2 males, 22
April 1978 (FZBRS 8038); 1 female, Machadinho, Linha do Tigre, ca. 27°35'17"S, 51°36'25"W, 9–10 May 2001,
R. Ott & L. Moura (FZBRS 33874); 1 female, Marcelino Ramos, ca. 27°28'31"S, 51°55'30"W, February 1989, A.
Braul (PUCRS 19807); 1 juvenile, Montenegro, ca. 29°41'03"S, 51°28'20"W, 3 November 1977, M. E. L. de Souza
(FZBRS 7093); idem, 1 male, M. H. Galileo (FZBRS 8070); 1 male, 3 females, 5 juveniles, Nonoai, Parque
Estadual de Nonoai, ca. 27°25'47"S, 53°02'25"W, 14 January 1985, A. A. Lise (FZBRS 13076); 1 female, Novo
Cabrais, Parque Witeck, ca. 29°46'59"S, 52°58'18"W, 3 January 2008, R. G. Buss (PUCRS 20302); idem, 1 female,
19.x.2007 (PUCRS 28327); 2 males, 2 females, 1 juvenile, Porto Alegre, Ponta Grossa, ca. 30°10'48"S,
51°11'11"W, 7 May 1976, A. A. Lise (FZBRS 8073); 1 male, Porto Alegre, Rio Apuaê, BR126, ca. 27°50'47"S,
51°49'59"W, September 1988, Equipe Proj. Ita-Machadinho (PUCRS 0537); 1 female, Salto do Jacuí, Horto
CEEE, ca. 29°04'49"S, 53°12'56"W, 19 October 1998, A. B. Bonaldo (FZBRS 30762); 1 female, Santa Maria,
Perau Velho, ca. 29°39'14"S, 53°46'45"W, 19 January 1995, C. B. Kotzian & L. Indrusiak (PUCRS 10467); idem, 1
female, 19 November 1998 (PUCRS 38376); 1 male, São Borja, Reserva Biológica São Donato, ca. 29°01'06"S,
56°09'43"W, 18 January 2012, M. Machado (PUCRS 36898); 1 male, São Francisco de Paula, ca. 29°26'47"S,
50°34'48"W, 23–25 November 1998, A. Silva (FZBRS 30819); 1 male, São Francisco de Paula, Barragem dos
Bugres, ca. 29°20'41"S, 50°42'07"W, 1–4 February 1999, A. Silva (FZBRS 30854); 1 female, São Francisco de
Paula, Fazenda 3 Cachoeiras, ca. 29°27'29"S, 50°33'38"W, 5 November 1998, A. Silva (FZBRS 30863); 1 female,
3 February 1999, A. B. Bonaldo (FZBRS 30961); 1 female, 2 juveniles, São Francisco de Paula, Usina Hidrelétrica
Passo do Inferno, ca. 29°16'09"S, 50°45'W, 26 September 2000, M. A. L. Marques (FZBRS 33241); idem, 4
females, 1 juvenile, 19 November 1997, E. H. Buckup (FZBRS 28815); 2 males, 2 juveniles, Tenente Portela, ca.
27°22'07"S, 53°45'35"W, 29 November 1978, H. Bischoff (PUCRS 8394); 1 female, Tenente Portela, Parque
Estadual do Turvo, ca. 27°12'26"S, 53°56'07"W, 15 January 1985, A. A. Lise (FZBRS 13074); idem, 1 female, 1
juvenile, 4–6 February 1980 (FZBRS 8983); 7 males, 20 females, 7 juveniles, Tenente Portela, Parque Estadual do
Turvo, Salto do Yucumã, ca. 27°08'37"S, 53°52'57"W, 16 January 1985, A. A. Lise (FZBRS 13056); idem, 7
females, 7 juveniles, 17 January 1985, A. A. Lise (FZBRS 13065); 1 female, Triunfo, ca. 29°56'12"S, 51°42'44"W,
28 November 1989, E. H. Buckup (FZBRS 19130); idem, 1 female, 25 January 1990, A. D. Brescovit (FZBRS
19343); idem, 1 female, 13 January 1994, L. A. Moura (FZBRS 24772); idem, 2 females, 16–17 March 1998, L.
Moura (FZBRS 29239); idem, 1 male, 1 female, 19 May 1977, A. A. Lise (FZBRS 5391); 1 male, 1 female,
Triunfo, Parque Copesul de Proteção Ambiental, ca. 29°55'46"S, 51°42'27"W, 28 November 2002, R. Araujo

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(FZBRS 34998); idem, 1 female, 5 February 2003, R. Ott (FZBRS 35152); idem, 1 female, 29 July 2003, R. Ott &
A. Barcellos (FZBRS 36093); idem, 1 female, 15 March 2007, M. A. L. Marques (FZBRS 43256); 2 females,
Viamão, ca. 30°04'37"S, 51°01'40"W, 1 May 1976, A. A. Lise (FZBRS 8066); idem, 1 male, 7 July 1995 (PUCRS
6669); 2 females, 1 juvenile, Viamão, Águas Belas, ca. 30°02'38"S, 50°59'59"W, 17 January 1977, A. A. Lise
(FZBRS 4978); 1 female, 29 December 1976, A. A. Lise (FZBRS 5861); 1 female, Viamão, Estação Fitotécnica,
ca. 30°02'15"S, 51°01'20"W, 6 May 1994, A. A. Lise (PUCRS 4688); idem, 1 male, 12 August 1994 (PUCRS
5238); 1 male, Viamão, Parque Estadual Itapuã, ca. 30°20'42"S, 51°01'31"W, April 2002, L. Ernesto (PUCRS
15331). Roraima: 1 female, Ilha de Maracá, ca. 3°25'00"N, 61°40'00"W, 21–30 November 1987, J. A. Rafael
(INPA). Santa Catarina: 2 males, 3 females, Rancho Queimado, ca. 27°40'17"S, 49°00'53"W, 13–15 January
1995, A. B. Bonaldo (FZBRS 26470); idem, 2 males, 3 females, 13–18 November 1995 (FZBRS 27046), idem, 2
females, 8–12 October 1994 (PUCRS 5949), idem, 2 males 3 females, 4 juveniles, 13–15 January 1995 (PUCRS
6137). São Paulo: 1 female, Paranapiacaba, ca. 23°46'32"S, 46°17'49"W, 9 October 1956, W. Bockerman (PUCRS
19738); 1 female, Rosana, ca. 22°34'47"S, 53°03'33"W, December 1986, Equipe IBSP (IBSP 4484); 1 female,
Salesópolis, Estação Biológica Boracéia, Trilha Laboratório, ca. 23°39'21"S, 45°53'29"W, 1 April 2001, S.T.P.
Amarante et al. (MZUSP 70593); 1 female, Santo André, Reserva Biológica do Alto da Serra de Paranapiacaba, ca.
23°47'10"S, 46°20'40"W, 14–16 December 2003, C.A. Rheims & R.P. Indicatti (IBSP 52069), idem, 1 male, ca.
23°48'24"S, 47°07'30"W, 28 October 2007, Equipe Butantan (IBSP 88546); 2 males, 1 female, São Paulo, Parque
Estadual do Jaraguá, ca. 23°27'34"S, 46°45'19"W, 27–28 October 2005 (IBSP 71290); 2 females, São Paulo, Horto
Florestal, ca. 23°27'22"S, 46°37'48"W, 27 March 1948, O.P. Forattini (MZUSP 13624). URUGUAY: Cerro Largo:
1 female, Paso Centurión, 32°08'01"S, 53°43'57" W, 13 April 2017, Bruno da Silva (FCE Ar 6184); 1 female, 15
April 2017, S. Teijón et al. (FCE Ar 6352); idem, 1 female (FCE Ar 6382), idem, 1 female, 1 juvenile (FCE Ar
6399); idem, 1 male, 6 November 2017 (FCE Ar 8444); 1 male, Sierra de Ríos, 32°12'55"S, 53°45'40"W, 14 April
2017, S. Teijón et. al. (FCE Ar 6483).
Diagnosis. E. villosus closely resembles E. spinipes, both belonging to the spinipes-group. Females of E.
villosus are distinguished from E. spinipes by the presence of a wide, median, brown longitudinal stripe along the
carapace, bifurcating in the posterior portion, the dorsum of the opisthosoma dark-brown to black, without white
lateral stripes, and dark-brown sternum with central yellowish area (Figs 11A, B). The paired macrosetal tubercles
of the tibia and metatarsus of E. villosus are shorter than in E. spinipes, with a length less than half the width of the
article. The epigynum has a rectangular median septum, with an anterior base almost as wide as the posterior one
(Fig. 11F), and spiral copulatory ducts with approximately seven coils, the last one not covering the spermatheca
(Fig. 11E). Males of E. villosus are distinguished from E. spinipes by the reddish-brown sternum bordered by dark
brown spots (Fig. 12C), by the RTA just slightly longer than wide (Fig. 12F), and by the embolus forming about six
turns around the tegulum (Figs 12E, F).
Notes. According to Mello-Leitão (1929), E. villosus was based on females from "Santo Antônio da Barra"
(now Santo Antônio de Condeúba), Bahia, and Paraguay, and the type was deposited under #11511 from Coll.
Simon. Since vial #11511 contains only one female labelled as “type” and collected in Bahia (Figs 13A–C), it is
considered as the holotype of the species. The two females from Paraguay were only identified as E. villosus by
Mello-Leitão but were not labeled as type material and were deposited under different numbers (Coll. Simon
#19194 and #9752). Taking into account the article 72.4.1 of the ICZN (1999), these females are part of the type-
series and are considered as paratypes of E. villosus. The holotype of Epicadinus helenae (Fig. 13D–F) is not an
adult female, as stated in the original description. It is a subadult female, with only a primordium of epigynum and
not displaying fully-developed genitalic structures. However, this specimen presents the same color pattern and
disposition of macrosetae and relative position of the conical projections of opisthosoma as adult females of E.
villosus. In addition, there are several records of E. villosus for the interior of the state of São Paulo. The only other
records for Epicadinus species for this state are a single specimen of E. spinipes, from northern coast of São Paulo,
and seven specimens of E. trispinosus, from the border with the state of Mato Grosso do Sul. The female identified
as a "homeotype" of E. helenae by Soares (1943), from the city of Curitiba, is also cospecific with E. villosus. The
last synonym is Epicadinus marmoratus, described by Mello-Leitão (1947) based on a "male" holotype from
Barigui, Curitiba. However, the holotype is a juvenile female (Fig. 13G–I), which also shows the typical color
pattern of E. villosus and a wide and long posterior conical projection. Besides, the only species of the genus
recorded from the Brazilian state of Paraná is E. villosus.

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FIGURE 11. Epicadinus villosus Mello-Leitão, 1929, female (FZBRS 39201). A–C habitus (A dorsal, B ventral, C lateral); D,
F epigynum ventral; E vulva dorsal; G female with colour variation (FZBRS 39201). Abbreviations, A, atrium; CD, copulatory
duct; MS, median septum; TK, translucent keel. Scale bars: A–C, G, 1 mm; D–F, 0.1 mm.

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FIGURE 12. Epicadinus villosus Mello-Leitão, 1929, male (MACN 19052). A–C habitus (A dorsal, B lateral, C ventral); D–F
left male palpus (D–E ventral, F retrolateral). Abbreviations, DTA, distal tibial apophysis; E, embolus; RTA, retrolateral tibial
apophysis; TF, tegular furrow. Scale bars: A–C, 1 mm; D–F, 0.1 mm.

Description. Female (FZBRS 39201, Fig. 11A–C): Total length: 4.87. Carapace: length 2.02, width 2.17,
height 1.20. Clypeus: height 0.21. Chelicera: length 0.56, width 0.35. Sternum: length 0.84, width 0.95. Endites
very setaceous: length 0.45, width, 0.26. Labium: length 0.25, width 0.37. Eye diameters and interdistances: AME
0.09, ALE 0.10, PME 0.07, PLE 0.06, AME–AME 0.013, AME–ALE 0.09, ALE–ALE 0.31, PME–PME 0.16,
PME–PLE 0.10, PLE–PLE 0.56. MOQ trapezoidal. Legs 1234. Leg I: femur 1.91, patella 0.99, tibia 1.63,
metatarsus 1.35, tarsus 0.71, total 6.59; II: 1.50-0.97-1.52-1.21-0.72-5.92; III: 1.00-0.55-0.66-0.48-0.31-3.00; IV:
1.30-0.55-0.70-0.62-0.40-3.57. Paired macrosetal tubercles in tibiae and metatarsi shorter than half width of the
article. Opisthosoma: length 2.85, width 3.37, height 2.10, pentagonal; anterior margin of the dorsum concave in
dorsal view and projecting over the posterior slope of the carapace, with projected anterior angles. Paired
projection with blunt apex and placed transversally in the median portion. Posterior conical projection with blunt
apex, slanted, almost parallel to the venter of opisthosoma, a little longer than the paired ones, and reaching 2/3 of

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the height of opisthosoma. In lateral view, anterior portion of the dorsum nearly flat and not clearly aligned to the
posterior conical projection, and posterior portion slightly convex.
Epigynum (Fig. 11D–F) mushroom-shaped, slightly wider than long. Atrium reniform, approximately twice as
wide as long, located anteriorly, formed by a deep depression, with the deepest areas forming two lateral foveas,
whose margins are covered by a thin translucent chitinous keel. Anterior portion of the atrium rises gradually
reaching the level of the surface of the opisthosoma. Median septum almost as long as wide, with an anterior base
slightly wider than posterior base and projecting itself laterally, covering the copulatory openings and merging into
the interior of the atrium. Its posterior base fuses to an area near the epigastric furrow, placed far from atrium.
Median portion of the septum very high, covering the copulatory openings and the middle portion of the posterior
margins of the atrium, both located in large and deep semicircular depressions. Margins of the epigynal plate
present long cilium-shaped bristles, which partially cover the atrium. Vulva (Fig. 11E) with very long spiral
copulatory ducts and a pair of posterior rounded spermathecae. Copulatory ducts initially formed by a simple,
curved duct, followed by a spiral portion directed anteriorly, with approximately six coils lighter in hue, with a
more dilated apical portion (secondary spermatheca). From the inner margin of this structure originates an inner
darker spiral portion, formed by approximately seven coils, which runs posteriorly in opposite direction to first
spiral, with a thicker ending portion which does not cover the spermathecae, and forming an almost straight distal
duct connected to the posterior inner face of the rounded and well-developed spermathecae.
Coloration. Carapace predominantly dark brown with wide dark brown longitudinal median stripe (Fig. 11A),
bifurcated only in its posterior third, and dark brown wide marginal stripes. Remaining areas yellowish around and
at the splitting of posterior part of central stripe. Sternum mostly dark brown with a central light yellowish area.
Labium, endites and chelicerae dark brown. Legs I and II predominantly dark brown, except for light yellow tarsi.
Legs III and IV light yellow, with irregular dark brown spots and rings. Opisthosoma predominantly dark brown
with venter brownish yellow and with light brown sigilas.
Male (MACN 19052, Fig. 12A–C): Total length: 3.39. Carapace: length 1.59, width 1.46, height 0.88.
Clypeus: height 0.17. Chelicera: length 0.43, width 0.23. Sternum very setaceous: length 0.65, width 0.63. Endites
very setaceous: length 0.28, width, 0.26. Labium: length 0.13, width 0.26. Eye diameters and interdistances: AME
0.07, ALE 0.06, PME 0.05, PLE 0.04, AME–AME 0.13, AME–ALE 0.08, ALE–ALE 0.29, PME–PME 0.15,
PME–PLE 0.11, PLE–PLE 0.46. MOQ trapezoidal. Legs 1243. Leg I: femur 1.28, patella 0.58, tibia 0.96,
metatarsus 0.81, tarsus 0.48, total 4.11; II: 1.23-0.66-0.88-0.73-0.46-2.73; III: 0.63-0.38-0.57-0.39-0.33-2.30; IV:
0.75-0.34-0.62-0.43-0.33-2.47. Paired macrosetal tubercles in tibiae and metatarsi shorter than half the width of the
article. Opisthosoma: length 1.80, width 2.20, height 1.30, pentagonal, covered by abundant macrosetae of variable
size. Anterior margin slightly concave in dorsal view and anterior angles a little projected over the posterior slope
of the carapace. Paired projections thin, with acute apex directed posterolaterally, and arranged transversally in
median portion of opisthosoma. Posterior conical projection very slanted posteriorly, not reaching half the height of
the opisthosoma itself, and similar in size to paired projections. In lateral view, anterior portion of dorsum nearly
flat and almost aligned to posterior conical projection, and posterior portion slightly convex.
Palpus (Fig. 12D–F). Tegulum with transversal furrow at 1 o’clock position. Embolus basis gradually
separating from tegulum, initially forming a wide stripe, slightly tapering during its median portion. Embolus
describes about six turns around lateral margin of tegulum, from which at least four are visible in non-distended
bulb in retrolateral view, and ends in a filiform apex, hidden by margin of tegulum. RTA fused to DTA, both
connected to the tibia by an elongated and narrow base, forming a kind of pedestal. RTA short, slightly longer than
wide; RTA also presents a large median concavity, marginal folds and apex describing a large rounded curve.
Dorsal fold slightly projected over the median concavity, joining directly to base of DTA. Ventral fold slightly more
projected on the median concavity than the dorsal one. DTA short, hook-shaped, with very thick base, excavated at
the inner surface of the hook, and apex forming a blunt tip.
Coloration. Carapace dark brown, with two longitudinal median light brown stripes. Sternum reddish brown,
radially bordered by dark brown spots. Endites, labium and chelicerae dark brown. Legs I and II with femur, patella
and tibia predominantly dark brown, with irregular reddish-brown spots, metatarsus and tarsus reddish brown. Legs
III and IV lighter, reddish brown with irregular dark brown patches and light brown patella. Opisthosoma dorsally
dirty yellow with irregular black spots and a black transversal stripe between the paired conical projections. Two
pairs of rounded reddish brown sigilla in the center of the dorsum of the opisthosoma. Projections dark brown with
reddish brown hues at the sides. Opisthosoma ventrally and laterally almost black, with reddish-brown sigils.

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FIGURE 13. A–C Epicadinus villosus Mello-Leitão, 1929, female holotype (MNHN AR-15078 [Coll. Simon 11.511]) [total
length: 7 mm] (A dorsal habitus, B ventral habitus, C original labels). D–F Epicadinus helenae subadult female holotype (IBSP
138.217 [ex. MZLQ 117]) (D dorsal habitus, E ventral habitus, F type labels). G–I Epicadinus marmoratus young holotype
(MNHCI 02496 [Coll. Rudolf B. Lange]) (G dorsal habitus, H ventral habitus, I type label). Scale bars: D,E,G,H, 1 mm. Photo
credits: A–C, Christine Rollard and MNHN Laboratory of Arachnology crew.

Variation. Females may present opisthosoma predominantly light yellow with a single trapezoidal dark brown
area on dorsum and some lighter irregular spots (Fig. 11G).
Distribution. Most records of E. villosus come from a wide range of the Brazilian Atlantic Forest, from
southern-central Bahia to the north, to Rio Grande do Sul (and also north Uruguay) to the south, reaching northern
Argentina and Paraguay to the west (Fig. 15). Most localities of E. villosus are located in inland areas, especially in

TAXONOMIC REVIEW OF EPICADINUS Zootaxa 4459 (2) © 2018 Magnolia Press · 229
the northern part of their distribution, but there are coastal records from the states of São Paulo to Rio Grande do
Sul, both in Brazil. The two isolated records of E. villosus from the Amazon are considered doubtful, pending
additional material. The record from Santarém, Pará, is represented by a male and a female specimens collected at
the 19th century, belonging to the Simon Collection (MNHN), and may represent a misplaced locality, as extensive
collections by MPEG in that area produced only specimens of E. biocellatus. The other record is for Ilha de
Maracá, Roraima, consisting of a single female, deposited in INPA. As the material of Ilha de Maracá was sorted
and identified at Rio Grande do Sul, an area with abundant records of E. villosus, there is also the possibility of
label exchange and mixing of localities.

Genus transfers

The following species are here transferred from Epicadinus to Epicadus. The transfers and synonymy are
supported by characters extracted from reviews of Epicadus and its synonyms (by Machado et al. 2015, Silva-
Moreira & Machado 2016, Machado et al. 2018) and analysis of specimens belonging to the latter genus.

Epicadinus polyophthalmus (Mello-Leitão, 1929) new combination

Epicadinus polyophthalmus Mello-Leitão, 1929: 102, fig. 46.

Type material. Holotype: juvenile, BRASIL, Rio de Janeiro: Niterói (MNRJ, Col. Mello-Leitão 901), lost.
E. polyophthalmus is based on a female holotype from Niterói (state of Rio de Janeiro, Brazil). The type
material should be deposited in MNRJ, but is considered lost (Silva-Moreira 2010). It was not found in that
collection, not even with new searches done by the authors. In the original description, Mello-Leitão highlights the
following characters: small size (4 mm), very spiny tegument, tubercles between the lateral eyes, elevated crest in
the carapace, and the color pattern, with black spots near the abdominal conical projections. The specimen
illustration (Mello-Leitão 1929: fig. 46) highlights other important features, such as the broad opisthosoma, with a
short and thin posterior conical projection, not exceeding the posterior margin of the opisthosoma, and the presence
of long macrosetae only in the ventral portion of the legs. These features allowed us to recognize that the holotype
of E. polyophthalmus was actually a young specimen, not an adult female. As in Epicadinus, Epicadus specimens
have ocular tubercles separating the lateral eyes and a spiny tegument. However, they may be recognized by the
presence of a spiny or high spiniform crest on the back of the carapace, and by the absence of robust setiferous
tubercles, surmounted by long macrosetae, covering the tegument. It is important to consider that juveniles of
Epicadus may not present the median crest of the carapace as high and prominent as the very conspicuous
spiniform projections of the adults, and exhibit great variation of size and shape of the abdominal tubercles and
color pattern. In juveniles, the paired tubercles may be very small and almost inconspicuous (R. Baptista pers.
obs.). Mello-Leitão (1929) mentioned the presence of two pairs of very weak, ventral paired "spines" (macrosetae)
in the tarsi, but this characteristic is probably an erroneous interpretation of spininiform bristles a little more robust,
since the genera of Stephanopinae do not have paired macrosetae on the tarsi. Among the valid species of
Epicadus, the illustration and description of Epicadinus polyophthalmus indicates that it may be a synonym of
some species found in or near Rio de Janeiro state: Epicadus rubripes Mello-Leitão, 1924, Epicadus taczanowskii
(Roewer, 1951) or Epicadus trituberculatus (Taczanowski, 1872). Epicadus rubripes and Epicadus trituberculatus
are common in Rio de Janeiro and have been recorded from Niterói (Silva-Moreira & Machado 2016), and
Epicadus taczanowskii has been recorded from the nearby states of Espírito Santo and São Paulo (Machado et al.
2018). So, Epicadinus polyophthalmus clearly belongs to Epicadus, and should be refered as Epicadus
polyophthalmus (Mello-Leitão, 1929) comb. nov. Some characteristics mentioned in the description indicate that
Epicadus polyophthalmus may be a synonym of Epicadus taczanowskii, such as the very spiny tegument (mainly
in the carapace), all three abdominal conical projections with thin tips, and the posterior conical projection not too
high. However, as the immature holotype of Epicadus polyophthalmus is lost, we prefer to consider it as a nomen
dubium.

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Epicadus tuberculatus (Petrunkevitch, 1910) new combination
Fig. 14

Epicadinus tuberculatus Petrunkevitch, 1910: 216, pl. 22, figs 20–22.

Tobias pustulosus Mello-Leitão, 1929: 86, figs 183–184. Machado et al. 2015: 570, figs 12–24. New synonymy.
Epicadus pustulosus (Mello-Leitão). Machado et al. 2017: 448.

Type material. Holotype of Epicadinus tuberculatus: Female immature, BRASIL, São Paulo: Juquiá, Fazenda
Poço Grande, Moenkhaus (PMNH, Coll. Mello-Leitão 901), lost.
Epicadinus tuberculatus was based on a holotype female from Poço Grande (Fazenda Poço Grande, Juquiá,
São Paulo, Brazil). The description and illustration of the holotype shows a specimen of large size (longer than 10
mm) that lacks the diagnostic needle-shaped setae of Epicadinus, indicating that this species does not belong to the
genus. The analysis of the photos of the holotype (Figs 14A–D), sent by the curator of the Peabody Museum of
Natural History, allows us to recognize that the holotype is in fact an immature female with only the primordium of
the epigynum. This species clearly belongs to a group of species that Machado et al. (2017) named “pustulosus
clade”, which is composed of spiders with cryptic bark-dweller habitus due to its dark and rough tegument, and by
the presence of large pustular setiferous tubercles in femora I and II (composed of Epicadus caudatus (Mello-
Leitão, 1929), Epicadus pustulosus (Mello-Leitão, 1929) and Epicadus granulatus Banks, 1909). Epicadus
caudatus, based on a female holotype from the city of Rio de Janeiro, has been recorded from the state of São Paulo
(Machado et al. 2015, Machado et al. 2018). Epicadus pustulosus, based on an immature female holotype from
Fonteboa, Amazonas state, northern Brazil, reaches south to the state of São Paulo (Machado et al. 2015, Machado
et al. 2018). As stated in Machado et al. (2015, 2018), the difference in color of the pustular setiferous tubercles on
the femora I and II allows an easy differentiation between Epicadus caudatus (orange/reddish) and Epicadus
pustulosus (yellowish). The large, yellowish setiferous tubercles in the femora I and II, and also the tarsi I and II
lighter than most leg articles of the holotype indicate that Epicadinus tuberculatus Petrunkevitch, 1910 is a senior
synonym of Epicadus pustulosus (Mello-Leitão, 1929) syn. nov. As Petrunkevich´s species has priority, the valid
name is Epicadus tuberculatus (Petrunkevicth, 1910) comb. nov.

FIGURE 14. Epicadus pustulosus (Mello-Leitão, 1929), subadult female holotype [total length: 10.5 mm]. A–D habitus (A
dorsal, B ventral, C frontal, D lateral). Photo credits: R. Pupedis and PMNH Laboratory of Arachnology crew.

TAXONOMIC REVIEW OF EPICADINUS Zootaxa 4459 (2) © 2018 Magnolia Press · 231
FIGURE 15. Geographic distribution records of Epicadinus, highlighting the type-locality of each of the four species (star
symbols).

Acknowledgements

We thank all the curators who contributed to this work by sending specimens. We are also grateful to C. Rollard
and Z. Simmons that were very receptive during visits to MNHN and OUNHM, respectively, and also sent photos
of the type specimens deposited there. W. Wawer (MIZ), R. Pupedis (PMNH), A. Laborda (FCE) and C. Favacho
(MPEG) were also very kind and helpful in sending fotos of types and other specimens. Furthermore, we are
thankful to José Ricardo Miras Mermudes (UFRJ) for the essencial support and instructions during this work, and
Jorge Nessimian for allowing the use of the automontage microscope at Laboratório de Entomologia/UFRJ. We
appreciate the helpful comments and suggestions of the editor (C. Muster), the two anonymous referees, and of our
dear colleagues Pedro Castanheira (UFRJ) and Renato Teixeira (PUCRS). This study was supported through MSc
grants to the first author by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).

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