17636_Paredes_Munguia_Brescovit_&_Teixeira_2024_Zoo_5414_1_83 (1)

Zootaxa 5414 (1): 001–083 ISSN 1175-5326 (print edition)
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Copyright © 2024 Magnolia Press
Monograph ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5414.1.1
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ZOOTAXA
5414
Revision of Neotropical wolf spider genus Arctosa C.L. Koch, 1847 (Araneae:
Lycosidae), with description of seven new species
WILLIAMS PAREDES-MUNGUÍA1, 2, 4, ANTONIO D. BRESCOVIT3, 5, RENATO A.
TEIXEIRA1,6
1
Pontifícia Universidade Católica do Rio Grande do Sul, Escola de Ciências da Saúde e da Vida, Museu de Ciências e Tecnologia,
Laboratório de Aracnologia, Porto Alegre, Brazil.
2
Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Laboratorio de Entomología, Av. Arenales 1256,
Lima 14, Peru.
3
Laboratório de Coleções Zoológicas, Av. Vital Brasil, 1500, Butantã, 05503-900, São Paulo, SP, Brazil.
4�
williams.paredes@acad.pucrs.br; https://orcid.org/0000-0002-3320-4611
5�
antonio.brescovit@butantan.gov.br; https://orcid.org/0000-0002-1511-5324
6�
renatoaug.tx@gmail.com; https://orcid.org/0000-0002-1756-9821
Magnolia Press
Auckland, New Zealand
Accepted by A. Santos: 19 Jan. 2024; published: 19 Feb. 2024

WILLIAMS PAREDES-MUNGUÍA, ANTONIO D. BRESCOVIT, RENATO A. TEIXEIRA
Revision of Neotropical wolf spider genus Arctosa C.L. Koch, 1847 (Araneae: Lycosidae), with
description of seven new species
(Zootaxa 5414)
83 pp.; 30 cm.
19 Feb. 2024
ISBN 978-1-77688-992-1 (paperback)
ISBN 978-1-77688-993-8 (Online edition)
FIRST PUBLISHED IN 2024 BY

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ISSN 1175-5326 (Print edition)

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· Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Table of Contents
Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Material and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Taxonomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Lycosidae Sundevall, 1833. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Arctosa C.L. Koch, 1847 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Identification key for Neotropical Arctosa species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Arctosa ayaymama sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Arctosa conflicta sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Arctosa costenola sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Arctosa denticulata Jiménez & Dondale 1984. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Arctosa jibarosa sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Arctosa littoralis (Hentz, 1844). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Arctosa mineira sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Arctosa pacaya sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Arctosa sapiranga Silva & Lise 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Arctosa serii Roth & Brown, 1976. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Arctosa tenebrosa (Keyserling, 1877) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Arctosa tenella (Keyserling, 1877) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Arctosa villa sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Misplaced species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Hogna andina (Chamberlin, 1916) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Hogna pugil (Bertkau, 1880) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Prolycosides aussereri (Keyserling, 1877) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Trochosa fusca (Keyserling, 1877) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Trochosa humicola Bertkau, 1880 comb. rest. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Abstract
The genus Arctosa C.L. Koch is redescribed and diagnosed. Seven new species are described, four of them based on
both sexes (A. conflicta sp. nov., A. costenola sp. nov., A. jibarosa sp. nov. and A. villa sp. nov.), two only on females
(A. ayaymama sp. nov. and A. mineira sp. nov.), and one only on the male (A. pacaya sp. nov.). We also propose the
transference of Arctosa humicola (Bertkau, 1880) and Arctosa fusca (Keyserling, 1877) to Trochosa C.L. Koch, 1847, and
the new synonymy of Arctosa inconspicua (Bryant, 1948) with Trochosa humicola (Bertkau, 1880) comb. nov. Besides,
Arctosa andina (Chamberlin, 1916) and Arctosa pugil (Bertkau, 1880) are transferred to Hogna Simon, 1885, and Arctosa
aussereri (Keyserling, 1877) to Prolycosides Mello-Leitão, 1942. Additionally, eight lycosid species are synonymized with
Prolycosides aussereri: Schizocosa proletaria (Tullgren, 1905); Arctosa workmani (Strand, 1909); Hogna planithoracis
(Mello-Leitão, 1938); Hogna variolosa (Mello-Leitão, 1941); Megarctosa melanostoma (Mello-Leitão, 1941); Hippasosa
huachoi (Mello-Leitão, 1942); Pirata abalosi (Mello-Leitão, 1942); and Pirata soukupi (Mello-Leitão, 1942). We also
transfer Trochosa tenebrosa Keyserling, 1877 to Arctosa. The males of Arctosa tenebrosa (Keyserling, 1877) comb. nov.
and Trochosa humicola (Bertkau, 1880) comb. nov. are described for the first time.
Key words: Diversity, taxonomy, wetlands, Andes, distribution
Introduction
The genus Arctosa C.L. Koch, 1847 was erected to include six European species of wolf spiders, diagnosable by
the presence of opisthosomal gray dots, irregular mottled prosoma pattern, dark annuli on legs and small eyes on
the first row (Koch 1847). The original genus diagnosis was based on imprecise characters and, as a result, Arctosa
became a wastebasket taxon with worldwide distribution. This fact motivated the review of the genus from different
regions of the world, e.g., North America (Gertsch & Wallace 1937; Dondale & Redner 1983a, 1983b), Central
Europe (Lugetti & Tongiorgi 1965, 1966), Japan (Tanaka 2009), and India (Sankaran et al. 2021). The phylogenetic
Revision of Neotropical species of Arctosa Zootaxa 5414 (1) © 2024 Magnolia Press ·
position of the genus has also been controversial, as it has been included in Lycosinae (Dondale 1986) or Arctosinae
(He & Song 1996). However, after several phylogenetic analyses (Zehethofer & Sturmbauer 1998; Vink et al.
2002; Murphy et al. 2006; Piacentini & Ramírez 2019), Arctosinae is considered a junior synonym of Tricassinae.
Besides, in recent phylogenetic hypotheses, Nearctic Arctosa species appear into Tricassinae and one neotropical (A.
sapiranga) is recovered within the subfamily Allocosinae (Gonnet et al., 2021; Laborda et al., 2022).
Arctosa comprises small to medium-sized swift runners with good eyesight (Dondale & Redner 1990), and with
long and slender legs (Dondale & Redner 1983a). Some species prefer seashores or live under stones on riverbanks
(Tanaka 1985), while others live near rivers or lakes (Doleschall 1852; Kulczyński 1885;). Heathlands, lichen, and
arctic tundra habitats are frequently occupied as well, as reported by Dondale & Redner (1983a) for Arctosa alpigena
(Doleschall, 1852). On the other hand, Arctosa cinerea (Fabricius, 1777) and Arctosa villa sp. nov. inhabit coastal
salt marshes (respectively Framenau et al. 1996, and Paredes-Munguia 2012 [sub Arctosa sp. nov.]). Furthermore,
sandy habitats are suitable to build diagonal retreats as reported by Dondale & Redner (1983a) for Arctosa perita
(Latreille, 1799) and A. littoralis (Hentz, 1844).
Currently, Arctosa comprises 172 species worldwide (around 7% of all lycosids), with 11 Neotropical species
(World Spider Catalog 2023), all having an unsteady taxonomic background. The taxonomic history of neotropical
species has been progressing southward starting with Arctosa littoralis (Hentz, 1844), which was described from the
western USA, near Mexico’s border, and initially confused with European representatives of A. cinerea by Emerton
(1885; 1902), Montgomery (1902) and Chamberlin (1908). The difficult differentiation among these two species
was solved by Chamberlin (1924) by comparing individuals from USA and Canada with A. cinerea from Europe.
Three species were subsequently included in the genus from the Caribbean and South America, Arctosa fusca
(Keyserling, 1877) (Cuba), Arctosa aussereri (Keyserling, 1877) (Puerto Rico) and Arctosa bogotensis (Keyserling,
1877) (Colombia); the original names Tarentula bogotensis and Lycosa aussereri was transferred to Arctosa by
Rower (1955), and Tarentula fusca was transferred to Arkalosula by Rower (1955), posteriorly synonymized to
Arctosa by Dondale and Redner (1983a). Arctosa pugil (Bertkau, 1880) and Arctosa humicola (Bertkau, 1880) were
described from Brazil and Guyana, respectively. Then Arctosa minuta F.O. Pickard-Cambridge, 1902 was described
from Guatemala and Panama; Arctosa andina (Chamberlin, 1916) from Andean mountains in Peru, at almost 3500
m altitude. Posteriorly, the genus current composition was completed by the description of Arctosa inconspicua
Bryant, 1948 from Haiti, two species from western Mexico, Arctosa serii Roth & Brown, 1976 and Arctosa
denticulata Jiménez & Dondale, 1984, and Arctosa sapiranga Silva & Lise, 2009 from Brazil, the southernmost
record of the genus.
After revising specimens available in several collections, including the type-material of Neotropical species,
we perceived the need to update the taxonomy of the genus. During this process, we not only concluded that some
species should be synonymized, but we have also found undescribed species that should be included within the
genus. In this paper, we illustrate and redescribe some species, and propose a dichotomous key for identification of
Arctosa species.
Material and Methods
Abbreviations and terminology. The description format and terminology follow Dondale & Redner (1983a, 1990),
Piacentini & Laborda (2013) and Wang et al. (2012). Synonymic lists will include only references that present
morphological or habitat descriptions, or taxonomic notes. Species lists or catalog are not included. We use the
terms “embolic division” and “palea region”, following Sierwald (2000). All measurements are in millimeters. The
abbreviations used in the text are the following: AER, anterior eye row; ALE, anterior lateral eyes; ALS, anterior
lateral spinnerets; AME, anterior median eyes; C, conductor; CD, copulatory duct; CCH, carapace cephalic height;
CCW, carapace cephalic width; CL, carapace length; CTH, carapace thoracic height; CTW, carapace thoracic
width; CYM, cymbium; DMA, depression of median apophysis; FD, fertilization duct; H, hood; HS, head of
spermatheca; MA, median apophysis; MS, median septum; Pa, posterior arm of median apophysis; PLE, posterior
lateral eyes; PLS, posterior lateral spinnerets; PME, posterior median eyes; POQ, posterior ocular quadrangle;
Pp, palea protuberance; pp, pars pendula; Ra, retrolateral arm of median apophysis; SF, stridulatory files; ST,
subtegulum; STA, subterminal apophysis; SS, stalk of spermatheca; TA, terminal apophysis; TL, total body length;
TLB, tegular lobe [modified from Dondale 1986, TL: tegular lobe]; TS, transversal septum; VC, vulval chamber.

Google Earth was used to tag the occurrences, then a .kml file was generated and read into Quantum GIS to create
the distribution map. When no specific geographic coordinates were available in the specimen labels, approximate
coordinates were attributed using Google Earth and named with "ca." (circa).
Specimens and figures. To examine the female genitalia, they were mechanically dissected from the opisthosoma
(following Levi 1965), cleaned using a thin brush, and then immersed in a pancreatin solution for 5 hours to
dissolve soft tissues (Álvarez-Padilla & Hormiga 2007). The internal female genitalia were immersed in clove oil
for photographing and drawing. Both epigyne and pedipalp drawings were made using a camera lucida mounted
on a Zeiss STEM SR stereoscopic microscope. The sketches were scanned and outlined in Inkscape (Inkscape
project 2020) with a Wacom graphics tablet. Photographs were taken with a Nikon AZ100M Multipurpose Zoom
Microscope, and a DMC 2900 digital camera attached to a Leica M205-A stereomicroscope. All specimens selected
for Scanning Electron Microscopy (SEM) were gradually dehydrated at increasing concentrations of 80%, 85%, and
90% up to absolute ethanol. Once dehydrated, they passed through critical-point drying in a Baltec CPD 030 Point
Dryer and immediately mounted on pin over SEM stubs with copper-backed tape. Finally, stubs were sputter coated
for 120 seconds with gold using a QUORUM 150R ES-Plus Vacuum Sputter. SEM images were obtained through
a Quanta F50 (FEI) Field Emission Scanning Electron Microscope from Centro de Microscopia e Microanálises
(CEMM) of the Pontifícia Universidade Católica do Rio Grande do Sul (PUCRS). All plates were composed using
GIMP 2.10.12 (The GIMP Development Team 2019) image editor.
All the material examined is deposited in the following collections (abbreviations and curators in
parentheses):
● American Museum of Natural History, New York, USA (AMNH, L. Prendini)
● Centro de Coleções Taxonômicas, Universidade Federal de Minas Gerais, Belo Horizonte, Brazil (UFMG, A.J.
Santos)
● Coleção de Aracnídeos, Departamento de Zoologia da Universidade Nacional de Brasília, Brazil (UnB, P.C.
Motta)
● Coleção de Aracnologia da Universidade Federal de Pernambuco, Recife, Brazil (UFPE, L. Iannuzzi)
● Colección de Arácnidos de la Universidad de Cartagena, Bolívar, Colombia (CUDC, G.R. Navas-Suarez)
● Colección de Insectos de la Universidad del Quindío, Quindío, Colombia (CIUQ, A.L. García-Hernández)
● Colección Entomológica Universidad de Nariño, Pasto, Colombia (CEUN-PSO, G. Castillo)
● Colección Nacional de Arácnidos, Departamento de Zoología, Instituto de Biología, Universidad Nacional
Autónoma de México (CNAN, E. González Santillán)
● Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (IRSNB, J. Kekenbosch)
● Instituto Butantan, São Paulo, Brazil (IBSP, A.D. Brescovit)
● Instituto de Ciencias Naturales de la Universidad de Colombia, Bogotá, Colombia (ICN-Ar, E. Florez)
● Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Boyacá, Colombia (IAvH, J.C.
Neita)
● Inventario Biológico Nacional, Museo Nacional de Historia Natural del Paraguay, San Lorenzo, Paraguay
(IBNP, J.A. Kochalka)
● Laboratorio de Entomología, Universidad de la Amazonia, Florencia, Colombia (LEUA, Y. Ramos-Pastrana)
● Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, Argentina (MACN, M.J.
Ramírez)
● Museo de Entomología “Klaus Raven Büller”, Lima, Peru (UNALM, C. Vergara)
● Museo de Historia Natural, Universidade Nacional Mayor de San Marcos, Lima, Peru (MUSM-ENT, D. Silva
Dávila)
● Museo de la Plata, Universidad Nacional de La Plata, La Plata, Argentina (MLP, L. Pereira & C. Damborenea)
● Museo de la Pontificia Universidad Católica de Quito, Quito, Ecuador (QCAZ, F. Salazar)
● Museo de la Pontificia Universidad Javeriana, Bogotá, Colombia (MPUJ, D. Forero)
● Museo de la Universidad del Valle del Cauca, Cauca, Colombia (MUSENUV, J. Cabra)
● Museo La Salle, Bogotá, Colombia (MSL, A. Rodríguez)
● Museu de Ciências Naturais, SEMA, Porto Alegre, Brazil (MCN, R. Ott)
● Museu de Ciência e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Brazil
(MCTP, R.A. Teixeira)
● Museu de História Natural Capão da Imbuia, Curitiba, Brazil (MHNCI, M. Zamoner)
● Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP, R. Pinto da Rocha)
● Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ, A.B. Kury)
● Museum of Comparative Zoology, Harvard University, Cambridge, USA (MCZ, G. Giribet)
● Naturhistoriska Riksmuseet, Stockholm, Sweden, (NRS, J. Stigenberg)
● Peabody Museum of Natural History, Yale University New Heaven, USA (PMNH, L.C. Dunn)
● The Natural History Museum, London, UK (BMNH, J. Beccaloni)
Taxonomy
Lycosidae Sundevall, 1833
Arctosa C.L. Koch, 1847
Arctosa C.L. Koch, 1847: 94. Type-species by subsequent designation (Simon 1937: 1089), Aranea cinerea Fabricius, 1777.
Bonnet 1955: 640; Dondale & Redner 1990: 278; Tanaka 1991: 290.
Leaena Simon, 1885: 11. Type-species by original designation, Lycosa personata L. Koch, 1872. Bonnet 1957: 2385; Roewer
1960: 939. Synonymized with Arctosa by Lugetti & Tongiorgi 1965: 186.
Tricca Simon, 1889: 250. Type-species by original designation, Tricca japonica Simon, 1889 [= Arctosa ipsa (Karsch, 1879)].
Bonnet 1959: 4684; Roewer 1960: 950. Synonymized with Arctosa by Dondale & Redner 1983a: 2.
Arctosella Roewer, 1960: 671. Type-species by original designation, Aranea perita Latreille, 1799. Synonymized with Arctosa
by Lugetti & Tongiorgi 1965: 175.
Arkalosula Roewer, 1960: 231. Type-species by original designation, Arctosa sanctaerosae Gertsch & Wallace, 1935. Roewer
1960: 759. Synonymized with Arctosa by Dondale & Redner 1983a: 2.
Triccosta Roewer, 1960: 298. Type-species by original designation, Tarentula japonica Bösenberg & Strand, 1906 [= Arctosa
ipsa (Karsch, 1879)]. Roewer 1960: 866. Synonymized with Tricca Simon, 1889 by Braun 1963: 81.
Leaenella Roewer 1960: 264. Type-species by original designation, Trochosa intricaria C.L. Koch, 1847. Roewer 1960: 867.
Synonymized with Arctosa by Wunderlich 1984: 23.
Bonacosa Roewer, 1960: 237. Type-species by original designation, Lycosa meinerti Thorell, 1875. Roewer 1960: 933.
Synonymized with Arctosa by Wunderlich 1984: 23.
Tetrarctosa Roewer, 1960: 296. Type-species by monotypy, Tarentula brevispina Kulczyński, 1908. Roewer 1960: 948.
Synonymized with Arctosa by Lugetti & Tongiorgi 1965: 194.
Diagnosis. Arctosa species can be recognized by having the posterior arm of the male median apophysis very
protruding (Figs 1c, 16c), as well as an extremely reduced palea (Figs 1b, 6a, 16a). Neotropical species of the
genus can be further recognized by the straight embolus, which is twisted on its own axis (Figs 1b, d). Females are
recognized by the triangular to trapezoidal epigynal plate; and the copulatory openings located at the convergence
of the median and transversal septum (Fig. 2a, Dondale & Redner 1983a; Paik 1994).
Species composition: There are 172 species of Arctosa worldwide, with almost half of them distributed in the
Afrotropical region (World Spider Catalog 2023). After this taxonomic revision, 13 are known from the neotropics:
Arctosa ayaymama sp. nov.; A. conflicta sp. nov.; A. denticulata Jiménez & Dondale, 1984; A. jibarosa sp. nov.; A.
littoralis (Hentz, 1844); A. costenola sp. nov.; A. pacaya sp. nov.; A. sapiranga Silva & Lise, 2009; A. serii Roth &
Brown, 1976; A. tenella comb. nov.; A. tenebrosa comb. nov. (Keyserling, 1877); A. mineira sp. nov. and A. villa
sp. nov.
To facilitate the identification of these species, we propose two groups with shared characteristics that can
further simplify their recognition. The “cinerea group” gathers eight widely distributed species: A. sapiranga, A.
ayayamama, A. costenola, A. villa, A. pacaya, A. denticulata, A. serii and A. littoralis. These species are possibly
closely related with Arctosa cinerea (type species of the genus) by the following diagnostic characters: males with
triangular posterior arm of median apophysis (Figs 29e, 44a, b) and acuminated retrolateral arm of median apophysis
(Figs 12a, 24b), and females with trapezoidal epigynal plate (Figs 4d, 31b, 47b). The “jibarosa group” consists of
five species with predominantly Andean distribution: Arctosa jibarosa, A. tenebrosa, A. mineira, A. conflicta and A.
tenella. The males of this group have a blunt retrolateral arm of median apophysis (Figs 7e, 34a), which is directed
ventrally, and a patch of setae on the dorsal tip of the cymbium; females have a quadrangular epigynal plate (Figs
23b, 42b) and round spermathecae with thin stalks (Figs 20b, 37e).

Distribution. About 40% of the species in each of the Afrotropical and Asian regions. Likewise, almost 20%
in Europe. In this taxonomical revision, 13 are considered Neotropical distributed, from Tijuana, in Mexico, to the
Parque Nacional El Palmar, Entre Ríos, Argentina (Figs 62–68).
Identification key for Neotropical Arctosa species
cinerea group, males

1 Depression of median apophysis (DMA) wide (Figs 14b, 47b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Depression of median apophysis (DMA) narrow (Fig. 7d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Terminal apophysis (TA) broad (Fig. 25b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa pacaya sp. nov.
- Terminal apophysis (TA) narrow (Fig. 15e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Palea protuberance ending in an acute tip, prolateral arm of the median apophysis bulging (Figs 14a, b), embolus straight (Fig.
16a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa costenola sp. nov.
- Palea protuberance bulging (Fig. 47e), embolus rotated on its own axis (Figs 46a, 47b) . . . . . . . . . . . . . Arctosa villa sp. nov.
4 Posterior arm of median apophysis (Pa) bulging (Roth & Brown 1976: fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Posterior arm of median apophysis (Pa) triangular (Jiménez & Dondale 1984: fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5 Palea protuberance absent (Dondale & Redner 1983a: fig. 73) . . . . . . . . . . . . . . . . . . . . . . . . .Arctosa littoralis (Hentz, 1844)
- Palea protuberance bulging (Dondale & Redner 1983a: fig. 57). . . . . . . . . . . . . . . . . . . . . . Arctosa serii Roth & Brown, 1976
6 Tip pointed palea protuberance acuminated and sclerotized (Jiménez & Dondale, 1984, fig. 5). . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Arctosa denticulata Jiménez & Dondale, 1984
- Tip pointed palea protuberance bulging and membranous (Figs 31e, 33a) . . . . . . . . . . . . Arctosa sapiranga Silva & Lise, 2009
cinerea group, females

1 Median septum (MS) completely flat (Fig. 5a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa ayaymama sp. nov.
- Median septum (MS) with a longitudinal raised ridge (Fig. 28i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Transversal septum projected posteriorly (Dondale & Redner 1983a: fig. 70). . . . . . . . . . . . . . Arctosa littoralis (Hentz, 1844)
- Transversal septum aligned with epigastric furrow (Figs 44a, 45b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Vulval chamber positioned below to the head of spermatheca, i.e., visible in dorsal view (Roth & Brown 1976: fig. 7) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa serii Roth & Brown, 1976
- Vulval chamber positioned behind to the head of spermatheca, i.e., not visible in dorsal vew (Fig. 28h) . . . . . . . . . . . . . . . . . .4
4 Head of spermatheca rectangular (Figs 12b, 13e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Head of spermatheca triangular or pear-shaped (Figs 28e, 44d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5 Epigynal plate projected into the atrium (Figs 12a, 13b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa costenola sp. nov.
- Epyginal plate not projected (Jiménez & Dondale 1984: fig. 1). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa denticulata Jiménez & Dondale, 1984
6 Head of spermatheca pear-shaped (Figs 28e, 32b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa sapiranga Silva & Lise 2009
- Head of spermatheca triangular (Figs 44d, 45b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Arctosa villa sp. nov.
jibarosa group, males

1 Depression of Median apophysis (DMA) shallow (Fig. 20f)…. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Arctosa jibarosa sp. nov.
- Depression of median apophysis (DMA) deep (Fig. 7d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Retrolateral arm of median apophysis (Ra) bulging and pronounced (Fig. 6a). . . . . . . . . . . . . . . . . . .Arctosa conflicta sp. nov.
- Retrolateral arm of median apophysis (Ra) with an acute tip (Fig. 40c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Terminal apophysis (TA) curved (Figs 41e, 43a). . . . . . . . . . . . . Arctosa tenella (F.O. Pickard-Cambridge, 1902) comb. nov.
- Terminal apophysis (TA) straight (Figs 34a, 35d). . . . . . . . . . . . . . . . . . . . . Arctosa tenebrosa (Keyserling, 1877) comb. nov.
jibarosa group, females

1 Median septum (MS) with a longitudinal raised ridge (Fig. 17a)… . . . . . . . . . . . . . . . . . . . . . . . . . . . .Arctosa jibarosa sp. nov.
- Median septum (MS) completely flat (Fig 22a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Transversal septum posteriorly projected (Figs 38a, 39b) . . . . . Arctosa tenella (F.O. Pickard-Cambridge, 1902) comb. nov.
- Transversal septum aligned with epigastric furrow (Figs 17a, 18e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Head of spermatheca rounded (Figs 8d, 9b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Arctosa conflicta sp. nov.
- Head of spermatheca differently shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4 Vulval chamber almost the same size of head of spermathecae (Figs 22b, 23d). . . . . . . . . . . . . . . . . . Arctosa mineira sp. nov.
- Vulval chamber shorter than the head of spermatheca (Fig. 37b). . . . . . . . . . . . . . . . . . . . . . . . . Arctosa tenebrosa comb. nov.
Arctosa ayaymama sp. nov.
Figs 2b, 3‒5, 62
Type Material. Female holotype from Yuracyacu, 5°56'34.11"S 77°08'53.44"W, 805 m, 7.ix.2008, Moyobamba,
San Martín, Peru, W. Paredes & C. Albujar leg. (MUSM-ENT 500411). Female paratype, near the same locality,
5°54'06.42"S 77°11'18.92"W, 816 m, 11.ix.2008. C. Albujar leg. (MUSM-ENT 505143).
Other material examined. COLOMBIA: Caquetá: 1♀, Curillo, Vereda El Conquistador, Laguna la Cocha,
ca. 1°20'16"N 75°54'49.7"W, 226 m, F. Arcos-Valencia leg. (LEUA 88163). PERU: San Martín: 1♀, Moyobamba,
Yuracyacu, 5°54'6.42"S 77°11'18.92"W, 816 m, 11–12.ix.2008, C. Albujar leg. (MUSM-ENT 500358); 4♀, ditto,
11.ix.2008, C. Albujar leg. (MUSM-ENT 506993).
Etymology. The species name comes from a Peruvian Amazon folk tale. In this story, one stepmother in the
middle of the jungle abandoned two stepsons, and their lament is heard on moonlit nights looking for his dead
biological mother. The sound of these children's wailing is folklorically known as "ayaymama".
Diagnosis. Females of A. ayaymama can be recognized among all Neotropical species of Arctosa by the
clearly triangular median septum (Figs 3a, 4d), like in Arctosa tenebrosa comb. nov., but with a deeper hood in A.
ayaymama (Fig. 3b). Likewise, A. ayaymama can be distinguished by the longitudinal median band (LMB) of the
carapace covered by white setae (Fig. 4a) and by the pyriform-shaped head of the spermatheca (HS), which narrows
anteriorly (Figs 3b, 5b).
Description. Male. Unknown.

Female. Holotype (MUSM-ENT 500411). Carapace brown with black radial pattern, covered with tiny black
setae. Longitudinal median band dark orange, covered by white setae, and tapering towards the opisthosoma (Fig.
4a, e), submarginal bands pale. Ocular area light orange, covered by sparse white setae (Fig. 4f). Sternum yellow,
sparsely covered by long brown setae, labium pale brown. Anterior row of eyes procurved, shorter than the middle
row, anterior median eyes larger than anterior lateral eyes (Fig. 4f). Chelicerae brown, darker than the endites and
labium (Fig. 4c), and condyle short. Legs, dorsum of femora yellowish with black annuli, patellae, tibiae, and
metatarsi yellowish with black annuli; tarsi yellow, with no annuli (Fig. 4e). A sparse scopulae on legs I−II, absent
in III− IV. Spinules present only on tarsi III−IV. All tibiae, metatarsi, and tarsi with trichobothria increasing in size
distally. Opisthosoma dark dusky with arrow-shaped cardiac mark flanked by yellow bands, covered by tiny white
setae, with an irregular yellow median band, distal portion with a chevron yellow pattern (Fig. 4a). Venter yellow as
the sternum with a dusky irregular band of spots. Spinnerets yellow.
Genitalia (Figs 2b, 3a‒d, 4b, d) with median and transversal septum of epigyne not distinguishable from each other
in ventral view, TS with rounded lateral borders; atrium narrow, MS not depressed. Fertilization ducts membranous
and fusiform (Fig. 3b). Copulatory ducts narrow. Spermatheca with stalk broad and curved outward, quite as thick
as the base of spermatheca; head of spermatheca pear-shaped, narrowing anteriorly visible in posterolateral view
(Fig. 3d).
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-2 d1-1-1 r0-d1-1ap, II r0-d1-d1 d1-1-1, r0-d1-d1 III
p0-d1-d1 d1-1-1 r1-1-1, IV pd1-0-d1 d1-1-1 r0-0-1; patella I p1 d 1bristle-1 bristle r1, II p1 d1 bristle – 1 bristle
r1, III p1 r1 d1-1bristle ap, IV p1 r1 d1bristle - 1ap; tibia I p1-v1 r0-1-1 v2-2-2ap, II pd1-1 r1-1 v2-2-2ap III,
p1-1 d1-1 r1-1 v1-2-2ap, IV p1-v1 d1-1 r1-1 v1-2-2ap; metatarsus I p0-1-1ap r0-1-2ap v2-2-1ap, II p1-1-2ap r0-
1-2ap v2-2-1ap, III p1-1-2ap rd1-1-2ap v2-2-1ap, IV pd1-d1-2ap rd1-v1-d1-0-2ap v2-2-1ap.
Measurements: TL 5.90, CL 2.80, CCW 1.64, CTW 2.50, CCH 1.24, CTH 1.32. Eyes: AME 0.20, ALE 0.10,
PME 0.38, PLE 0.30, interdistances: AME–AME 0.12, AME–ALE 0.08; PME–PME 0.30, PME–PLE 0.38, PLE–
PLE 0.88, POQ length 0.66, POQ posterior width 1.04, POQ anterior width 0.60. Anterior eye row procurved, as
long as the PMEs. Chelicerae: 1.18 length. Opisthosoma: length 2.76, width 1.98. Legs: length of segments (femur
+ patella+tibia + metatarsus+tarsus = total length): pedipalp 1.10 + 0.56 + 0.80 + - + 0.90 = 3.36, I 2.30 + 1.10 +
1.94 + 1.78 + 1.22 = 8.34, II 2.10 + 0.98 + 1.68 + 1.66 + 1.16 = 7.58, III 2. 00 + 0.82 +1.54 + 2.02+1.10 = 7.48, IV
2. 68 + 1.10 + 2. 38 + 3.20+1.48 = 10.84.
Variation. Females (N = 6, range, mean±s.d.): TL 5.67–7.92, 6.51±0.33; CL 3.00–3.67, 3.31±0.09; CW 2.17–
2.58, 2.46±0.06.
Distribution. Amazonian Colombia (Caquetá) and Peru (San Martín) (Fig. 62).

FIGURE 1. Arctosa spp. Female Arctosa conflicta sp. nov. a, c, e. Habitus with measurement patterns (a, lateral; c, frontal; e,
dorsal). b, f. male palpus Arctosa villa sp. nov. (b, ventral; f, retrolateral). d. Male Arctosa tenebrosa comb. nov. Scale bars: a,
c, e 2 mm; b, d, f, 0.25 mm. ANT anal tubercle, ALE anterior lateral eye, AME anterior median eye, C conductor, CH prosoma
height, CL prosoma length, CW prosoma wide, CYM cymbium, DMA depression of MA, E embolus, LMB longitudinal
median band, MA median apophysis, OA ocular area ,Pa Posterior arm, Pp palea protuberance, PLE postero lateral eye, PME
postero median eye, PSB pale submarginal band, Ra Retrolateral arm, SD seminal duct, SP spinnerets, T tegulum, TA Terminal
Apophysis, ST subtegulum.
FIGURE 2. Epigynum. a, c, f. Arctosa villa sp. nov (a, ventral; c, dorsal; f, posterolateral). b. Arctosa ayaymama sp. nov. d.
Arctosa villa sp. nov. (anterior). e. Arctosa mineira sp. nov. (posterior). g. Arctosa jibarosa sp. nov. (posterolateral). Scale bars:
a–g, 0.25 mm. AT, atrium, BS, base of spermatheca, CO, copulatory opening, FD, fertilization duct, H, hood, HS, head of
spermatheca, MS, median septum, SS, stalk of spermatheca, TS, transversal septum, VC, vulval chamber.
10 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Arctosa conflicta sp. nov.
Figs 1a–c, 6‒11, 63
Type material. Male holotype from Poblado de Carahuasi, ca. 7°3'14.21''S 79°10'6.93''W, 1301 m, 25.iii.2010,
Cajamarca, Peru, C. Espinoza leg. (MUSM-ENT 501400). Female paratype from Hualgayoc, 6°45'S 78°36'W,
±3720 m, Cajamarca, Peru, no further data, no collector (MUSM-ENT 505361).
Other material examined. COLOMBIA: Cundinamarca: 1♂, Pantano de Martos, Vereda Monquetiva,
Guatavita, 4°54'30.09''N 73°43'49.6''W, 3030 m, 27.x.2014, K. Pulido leg. (ICN-Ar 12746); 1♂, Pantano de
Martos, Guatavita, Vereda Monquetiva, 4°54'05.4''N 73°43'33.92''W, 2940 m, 26.x.2014, K. Pulido leg. (ICN-Ar
12817); 1♂, ditto (ICN-Ar 12818); 1♂, ditto (ICN-Ar 12748); 1♂, Vereda Monquetiva, Guatavita, 4°43'36.6''N
73°51'53''W, 3400 m, 12.xi.2014, D. Martinez leg. (ICN-Ar 12747); 1♂1♀, La Calera, Finca Tierraleja, Vereda
Jerusalén, 4°38'52.86''N 73°56'7.85''W, (human feces), 3580 m, 24.xi.2014, K. Pulido leg. (ICN-Ar 12814); 1♂,
ditto (ICN-Ar 12815); 1♂, Guasca, Vereda Camino del Oso, Finca Suasie, 4°43'55.7''N 73°51'49.5''W, 3500 m,
12.xi.2014, D. Martinez leg. (ICN-Ar 12819). Pasto: 1♀, no specific locality, ca. 1°12'10.36''N 77°18'07.04''W,
2714 m, (CEUN-PSO). ECUADOR: Pichincha: 1♀, Amaguaña, Volcán Pasochoa, 0°26'50.73"S 78°28'36.11"W,
3320 m, 9.xi.2013, E. Salazar leg. (QCAZ); 1♀, Cotopaxi, Reserva Acosa, 3500 m, 0º40'S 78º26'W, pine plantation
páramo, 29.i.2000, L.L. Tapia leg. (QCAZ); 1♀, Quito, pine plantation, 2810 m, 0º11'22"S 78º29'38"W, 12.x.2006,
J. Mejia leg. (QCAZ); 1♀, Quito, Parque, Rumipampa, 2810 m, ca. 0º10'48.26"S 78º29'58.09"W, 29.i.1994, J.
Durango leg. (QCAZ 1285); 1♂, Papallacta, 0°22'S 78°09'W, 3200 m, 17.i.1999, S. Castelo leg. (QCAZ). Santo
Domingo: 1♀, Guajalito, 0º13'59"S 78º49'00.1"W, 1800 m, 9.xi.2013, A. Gortaire leg. (QCAZ). Cañar: 1♀, ca.
2º33'55.91"S 78º56'19.77"W, 2500 m, viii.1986, Onore leg. (QCAZ).
Etymology. The species epithet is a noun with means "conflict" and refers to the social conflicts between the
Hualgayoc peasant and the mining companies which caused environmental degradation in the area.
FIGURE 3. Arctosa ayaymama sp. nov. Epigynum, holotype a–b (MUSM-ENT 500411) (a, ventral; b, dorsal); paratype c–d
(MUSM-ENT 505143) (c, ventral; d, posterolateral) Scale bars: a–d, 0.25 mm. CO copulatory opening, FD fertilization duct, H
hood, HS head of spermatheca, SS stalk of spermatheca, TS transversal septum.
Revision of Neotropical species of Arctosa Zootaxa 5414 (1) © 2024 Magnolia Press · 11
FIGURE 4. Arctosa ayaymama sp. nov. Female habitus (MUSM-ENT 500411) a, c, e–f (a, dorsal; c, ventral; e, lateral, f,
frontal) Epigynum b, d (b, dorsal; d, ventral) Scale bars: a, c, e, 1 mm, b, d 100 µm, f, 500 µm.
Diagnosis. Arctosa conflicta males are recognizable among all Neotropical species by the pronounced
retrolateral arm of the median apophysis, and bifid sclerotized palea protuberance (Pp) (Figs 6a, 7d). Females can be
distinguished from all other members of Arctosa by the large and rounded head of spermatheca (Figs 8c, d, 9), and
by the stalk of the spermatheca (SS) strongly curved, U-shaped (Fig. 9b). Both sexes have a dark brown sternum,
but females have an incomplete yellow band on the middle (Figs 7b, 10c).
Description. Male. Holotype (MUSM-ENT 501400). Carapace brown, covered by short white and black bristles,
except on the yellow longitudinal median band, which is broad at the front and tapering towards the back (Fig.7a).
Sternum longer than wide, light brown with a transverse yellow band in the anterior half, covered by short black
bristles, which are more abundant at the marginal area and sparse in the central area. Ocular area black, covered with

FIGURE 5. Arctosa ayaymama sp. nov. Epigynum (MUSM-ENT 506993) a–c (a, ventral; b, dorsal; c, dictynoid pore) d, Male
leg I, prolateral. Scale bars: a, 150 µm, b, 400 µm, c, 15 µm, d, 200 µm. CO copulatory opening, FD fertilization duct, H hood,
HS head of spermatheca, DP dictynoid pore, MS median septum.
FIGURE 6. Arctosa conflicta sp. nov. Male bulb (ICN-Ar 12814) (a, ventral; b, retrolateral; c, apical. Scale bars: a–c, 200 µm.
E embolus, Pa Posterior arm, Pp Palea protuberance, Ra Retrolateral arm, TA terminal apophysis.
FIGURE 7. Arctosa conflicta sp. nov. Male habitus (ICN-Ar 12814) a–c (a, dorsal; b, ventral; c, lateral), pedipalpal bulb d–g
(d, ventral; e, prolateral; f, anterior; g, posterior). Scale bars: a–g, 200 µm. DMA depression of MA, E embolus, Pa posterior
arm, Pp Palea protuberance, Ra retrolateral arm, SD seminal duct, ST subtegulum, TA terminal apophysis.
white bristles, cephalic portion of the carapace covered by sparse, large white bristles. First row of eyes straight,
shorter than the second row; AME slightly larger than the ALE; PME separated a radius apart. Chelicerae, creamy,
covered frontally with long, brown setae and some shorter, white setae, boss present and clearer than the chelicerae;
retromarginal and promarginal fang furrow with three teeth each, chillum membranous, divided, with the mesial
portion sclerotized. (Fig. 10e). Endites creamy and convergent; labium square, darker than the endites; endites and
labium clearer in their distal portion. All leg segments yellow, except tarsi that are darker, all femora and tibia with
black spots. Opisthosoma entirely olive-colored, covered by tiny white bristles and sparse black bristles mainly on
the dorsum, cardiac mark present with a group of strong white bristles on the dorsal proximal part, near the prosoma.
Venter clearer than the dorsal part, sparsely covered by black bristles; spinnerets yellow both cylindrical (Fig. 7b).
Pedipalp (Fig. 7d‒g), tibia cylindrical, slightly wider apically. Cymbium fusiform (Fig. 7b). Subtegulum
trapezoidal and noticeable in ventral view, with a curved sperm duct crossing the tegulum from base to the apex
(Fig. 7e, g). Median apophysis with retrolateral arm bulging and pointed, prolateral arm triangular (Fig. 7f). Basal
portion of the TA partially sclerotized, Pp bifid and sclerotized (Figs 6a, 7d). Embolus straight (Figs 6c, 7d).
Leg formula IV>I>III>II. Spination pattern: femur I p0-0-1 d1-1-1, II p0-0-1 d1-1-1 r0-d1-0, III pd1-d1 d1-
1-1 rd1-d1, IV pd1-d1 d1-1-1 r0-d1; patella I d1(bristle)-0, II p1 d0-1-ap(1 bristle), III p1 d1bristle-1ap r1, IV p1
d1bristle-1ap r1; tibia I p1-1v v2-2-2ap r0-1-0, II p1-1v d1bristle-1bristle v2-2-2ap, III p1-1v d 1 r1-1 v2-2-2ap,
IV p1-1 d1r-1 r1-1 v2-2-2ap; metatarsus I p0-0-1 r0-0-1 d1bristle-1bristle v2-2-1ap, II p0-0-2 d1 bristle-0 r1-1-2
v2-2-1ap, III p1d-1-2 d1bristle-0 r1-1v-2 v2-2-1ap, IV p1d-1-2ap d1bristle-0 r1d-1-2ap v2-2-1ap.

FIGURE 8. Arctosa conflicta sp. nov. Epigynum (MUSM-ENT 505361) a, c, d (a, ventral; c, dorsal; d, basal), prosoma, b,
e (b, lateral; e, frontal) Scale bars: a, c, d, 0.25 mm, b, e, 2 mm. CO copulatory opening, FD fertilization duct, HS head of
spermatheca, VC vulval chamber.
FIGURE 9. Arctosa conflicta sp. nov. Epigynum (MUSM-ENT 505361) (a, ventral; b, dorsal) Scale bars: a–b, 0.1 mm. HS
head of spermatheca, SS stalk of spermatheca.
FIGURE 10. Arctosa conflicta sp. nov. Female prosoma (MUSM-ENT 505361) a, c, e (a, dorsal; c, ventral; e, lateral), female
opisthosoma b, d, f (b, dorsal; d, ventral; f, lateral) Scale bars: a–f, 1 mm.
PME 0.26, PLE 0.21, interdistances: AME-AME 0.09; AME-ALE 0.05; PME-PME 0.14; PME-PLE 0.30; PLE-PLE
0.50. POQ long 0.52, POQ posterior width 0.71, POQ anterior width 0.43. Chelicerae: length 1.31. Opisthosoma:
length 2.67, width 1.83. Legs: length of segments (femur + patella + tibia + metatarsus + tarsus = total length):
pedipalp 1.04 + 0.52 + 0.67 - + 0.79 = 3.02, I 2.05 + 0.76 + 1.78 + 1.62 + 1.05 = 7.26, II 1.86 + 0.83 + 1.45 + 1.45
+ 1.05 = 5.19, III 1.74 + 0.76 + 1.29 + 1.79 + 0.97 = 6.55, IV 2.24 + 1.02 + 2.00 + 2.62 + 1.21 = 9.09.

FIGURE 11. Arctosa conflicta sp. nov. Epigynum SEM (a, ventral; b, copulatory opening detail; c, conic papillae) Scale bars:
a, 150 µm, b, 25 µm, c, 50 µm. CO copulatory opening, H hood, MS median septum, TS transversal septum.
Female. Paratype (MUSM-ENT 505361). Carapace, eye pattern and proportions, and chelicerae as in male (Fig.
10c); visible boss, clearer than the chelicerae. Sternum dark brown with an incomplete yellow median band (Fig.
10c); endites light brown and convergent; labium darker than the endites; both endites and labium yellowish in their
distal portion. All femora yellow, tibia, metatarsi, and tarsi darker than femora, legs with black spots, except on the
metatarsus and tarsus. Opisthosoma, color as in male, cardiac mark paler.
Genitalia (Figs 8a, c, d, 9a‒b, 11a‒c), median and transversal septum of epigyne not distinguishable from each
other, lateral sclerotized margins converging in a bulb-like shape. Atrium narrow and slightly trapezoidal in ventral
view. Copulatory opening narrow (Figs 8a, 11b). Fertilization ducts short and thin (Fig. 8c). Head of spermathecae
large and rounded; stalk long and curved; copulatory opening narrow; vulval chambers ectally rounded and smaller
than the spermatheca (Fig. 8d).
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1 d1-1-1, II p0-1-1 d1-1-1, III p0-d1-d1 d1-1-1 r0-
d1-d1, IV p0-0-d1ap d1-1-1 r0-0-d1ap; patella I d1-ap(1 bristle), II d1-ap(1 bristle), III p1 dbas (1bristle)-1ap r1,
IV p1 d(1 bristle)-1ap r1; tibia I p0-1-d1 d(1 bristle)-0 v2-1-2ap, II pd1-1 v2-2-2ap, III pd1-1 d(1bristle)-1 r1-1
v 2-2-2ap, IV pd1-1 dr1-0-1 r1-1 v2-2-2ap; metatarsus I p0-d1-2 r0-v1-1ap v2-2-1ap, II p0-d1-2 r0-0-1 v2-2-1ap,
III pd1-d1-2 rd1-1-2 v2-2-1ap, IV pd1-d1-2ap rd1-v1-d1-0-2ap v2-2-1ap.
0.56. POQ long 0.44, POQ posterior width 0.76, POQ anterior width 0.44. Chelicerae: length 1.40. Opisthosoma:
length 1.66, width 1.06. Legs: length of segments (femur + patella + tibia + metatarsus + tarsus = total length):
pedipalp 1.20 + 0.54 + 0.66 - + 0.80 = 3.26 (1.04 + 0.52 + 0.67 - + 0.79 = 3.02), I 2.20 + 1.20 + 1.74 + 1.50 + 1.04
= 8.68, II 1.92 + 1.14 + 1.60 + 1.42 + 1.02= 8.12, III 2.00 + 1.00 + 1.40 + 1.72 + 1.04 = 7.69, IV 2.50 + 1.20 + 2.02
+ 2.60 + 1.44 = 10.91.
Variation. Females N = 6 (males N = 4) (range, mean±s.d.): TL 5.90 – 8.50, 7.26±1.01; CL 3.20 – 3.80,
3.38±0.21; CW 2.40– 2.80, 2.49±0.14; (TL 4.40 – 6.90, 5.83±1.03; CL 2.60 – 3.80, 3.20±0.47; CW 1.80 – 2.60,
2.20±0.35).
Distribution. Colombia (Cundinamarca), Ecuador (Pichincha, Santo Domingo, Cañar) and Peru (Cajamarca)
(Fig. 63).
FIGURE 12. Arctosa costenola sp. nov. Male pedipalp holotype (ICN-Ar 1378) a–c (a, anterior; b, ventral; c, retrolateral).
Scale bars: a–c, 0.25 mm. CMS cymbial macrosetae, DMA depression of median apophysis, E embolus, Pa posterior arm, Pp
palea protuberance, Ra retrolateral arm, ST subtegulum, T tegulum, TA terminal apophysis
Arctosa costenola sp. nov.

Figs 12‒16, 64
Type Material. Male holotype from Isla Barú, ca. 10°15'00.44"N 75°34'59.24"W, x.1999, Cartagena de Indias,
Colombia, E. Florez et al leg. (ICN-Ar 1378); female paratype with same locality and same data (ICN-Ar 12745).
Etymology. “Costeñol” is a regional dialect formed by a mixture of Spanish and Colombian Caribbean slang
(Lambraño 2011).
Diagnosis. Males of Arctosa costenola sp. nov. can be recognized among all the Neotropical species by the
straight and untwisted embolus (Fig. 13e), the bulging posterior arm of the median apophysis (Fig. 12a), and the
curved, beak-like, and keeled retrolateral arm of the median apophysis (Fig. 13b). Females of Arctosa costenola are
distinguishable among all Neotropical species by the hood notched (Fig. 14a) and by the head of spermatheca (HS)
rectangular in dorsal view, with two rounded bulges at the tip (Figs 14b, 15e).
Description. Male. Holotype (ICN-Ar 1378). Carapace brown, with a yellow longitudinal, median band,
wider anteriorly to fovea. MLB tapering from fovea towards posterior margin of opisthosoma; submarginal bands
discontinuous. Two parallel brown stripes behind the ocular area. Carapace covered with little black setae, MLB
with sparse white setae; ocular area black, covered with long black setae and some short white ones. Sternum paler
brown, with an incomplete yellow median band, and sparsely covered with brown bristles, which more abundant on

FIGURE 13. Arctosa costenola sp. nov. Male pedipalp (ICN-Ar 1378) a–c (a, prolateral; b, ventral; c, retrolateral). Pedipalp
bulb d–h (d, prolateral; e, ventral; f, retrolateral; g posterior; h, anterior). i dorsal cymbium macrosetae. Scale bars: a–c, 0.20
mm, d–h, 100 µm, i, 200 µm. C conductor, DMA depression of median apophysis, E embolus, Ra retrolateral arm, Pa posterior
arm, T tegulum, TA terminal apophysis, TK tegular keel, ST subtegulum,
the margins (Fig. 15d). Four long black bristles on the clypeal condyle anteriorly of AER. First row of eyes straight
and shorter than the second row (Fig. 15h); AME slightly bigger than ALE; ALE resting on tubercles distally
oriented; PME separated by more than half their radius. Labium light brown covered with brown bristles the same
color as the chelicerae, boss short and poorly visible. Legs yellow, darker distally. All femora yellowish with black
irregular patches; leg I and II light brown from patellae to tarsi (Fig. 15g). All tarsi with no scopula, tarsi III and IV
with spinules; trichobothria in tibiae, tarsi and metatarsi distally increasing in size. Opisthosoma dark olive-gray,
dorsally darker near the pedicel, with long white bristles in the middle and black bristles on both sides, cardiac mark
pale yellow, lanceolate, flanked with irregular but symmetric yellow spots at both sides; venter dark yellow with
some brown patches; spinnerets yellow.
Pedipalp (Fig 12‒13, 16) with cylindrical tibia, about two times longer than wide, with a ventral patch of bristles
on the femur. Cymbium piriform with a strong apical spine; embolus straight, transversally oriented and not twisted
(Figs 12a, 13e, 16a); subtegulum squared, in a notch in the tegulum, TA fine-pointed; Pa of median apophysis bulging
and Ra with a dorsal keel, beak-like and curved in ventral view. Left palp with sperm ducts running diagonally from
the prolateral to retrolateral, drawing a “V” pattern through the tegulum in a clockwise way (Fig. 13b).
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1 d1-1-1, II p0-0-1 d1-1-1, III d1-2-1-3, IV d1-1-3;
patella I d1bristle-1bristle ap, II p1 d1bristle-1bristle ap III p1 d1bristle-1 bristle ap r1, IV p1 d1bristle -1bristle ap
r1; tibia I p1v-1 d1bristle v2-1-2ap, II pd1-1 d1bristle v1-1-2ap III, p1-1 d1r-1 r1-1 v2-2-2ap, IV p1-1 d1r-1 r1-1
v2-2-2ap; metatarsus I p1ap r1ap v2-2-2, II p0-2-2ap r0-0-2ap v2-1r-1axp, III p1-1ap d2-0-2 r1-1ap v2-2-1ap, IV
p1-1-2ap r1-1-0-2ap v3-2-1ap.
FIGURE 14. Arctosa costenola sp. nov. Female epigynum (ICN-Ar 12745) a–d (a, ventral; b, dorsal; c, anterior; d, posterolateral).
Scale bars: a–d, 0.25 mm. FD fertilization duct, H hood, HS head of spermatheca, MS median septum, SS stalk of spermatheca,
VC vulval chamber.
PME 0.26, PLE 0.18, interdistances: AME-AME 0.08, AME-ALE 0.04; PME-PME 0.16, POQ length 0.32, POQ
posterior width 0.54, POQ anterior width 0.36. Chelicerae: length 1.16. Legs: length of segments femur + patella

+ tibia + metatarsus + tarsus = total length: pedipalp 1.00 + 0.48 +0.64 + - + 0.80 = 2.92, I 1.74 + 0.86 + 1.50 +
1.32 + 0.92 = 6.34, II 1.68 + 0.88 + 1.30 + 1.32 + 0.88 = 6.06, III 1.64 + 0.84 + 1.14 + 1.50 + 0.86 = 5.98, IV 2.36
+ 1.02 + 1.78 + 2.36 + 1.16 = 8.68.
Female. Paratype (ICN-Ar 12745). Coloration and pattern as in male. Anterior eye row slightly procurved, ALE
on small tubercles. All legs with the same color patterns as in males. Coloration and setae arrangement as in male
(Figs 15c, f, i).
FIGURE 15. Arctosa costenola sp. nov. Male habitus (ICN-Ar 1378) a, d, g–h (a, dorsal; d, ventral; g, lateral; h, frontal).
Epigynum (ICN-Ar 12745) b, e (b, ventral; e, dorsal). Female habitus (ICN-Ar 12745) c, f, i (c, dorsal; f, ventral; i, lateral) Scale
bars: a, d, g, 1 mm, b, e, 0.1 mm, h, 500 µm, c, f, i, 2 mm. H, hood, HS head of spermatheca, MS median septum, SS stalk of
spermatheca, TS transversal septum, VC vulval chamber.
Genitalia with the median septum and transversal septum (TS) forming a trapezoidal pattern (Fig. 15b); TS with
rounded lateral borders; atrium narrow, parallel to median septum; lateral lobes slightly sclerotized; anterior part of
the MS depressed. FD fusiform and membranous. Copulatory ducts broad located at each rounded border of the TS.
Spermatheca rectangular, with two small, rounded bulges (Figs 14d, 15e), SS thin, curved distally, VC wide (Fig.
15e). fertilization ducts thin, membranous, and postero-dorsal origin
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1d d1-1-0, II p0-0-1 d1-1-1, III p0-1d-d1 d1-1-1
r0-1d-1d, IV p0-1d-0-1d d1-1-1 r0-0-1d; patella I d1bristle-1bristle ap, II d1bristle-1bristle ap III p1 d1bristle-ap1
r1, IV p1 d1bristle-ap1 r1; tibia I p1v-1 d1bristle v2-1-2ap, II p1d-1 d1bristle v1-1-2ap III, p1-1 d1r-1 r1-1 v2-2-
2ap, IV p1-1 d1r-1 r1-1 v1-2-2ap; metatarsus I p0-1v-1 d1bristle-0 v2-1-2, II p0-1v-1ap d1 bristle-0 v2-0-1ap, III
p1-1-2ap d1bristle-0 r1-1-0-2ap v2-2-1ap, IV p1-1-2ap r1-1-2ap v1-3-2-1ap.
posterior width 0.66, POQ anterior width 0.46. Chelicerae: length 1.38. Legs: length of segments femur + patella
+ tibia + metatarsus + tarsus = total length: pedipalp 1.16 + 0.66 + 0.58 + - + 0.84 = 3.24, I 2.06 + 1.14 + 1.60 +
1.42 + 0.96 = 7.18, II 1.98 + 1.00 + 1.36 + 1.42 + 0.92 = 6.68, III 1.90 + 0.96 + 1.20 + 1.64 + 0.88 = 6.58, IV 2.46
+ 1.20 + 2.12 + 2.70 + 1.34 = 9.82.
2.20±0.28; CW 1.27 – 1.80, 1.60±0.17; (TL 2.82 – 5.20, 4.03±0.65; CL 1.62 – 2.73, 2.29±0.31; CW 1.17 – 1.97,
1.60±0.23)
Distribution. Colombia (Bolívar, Fig. 64)
FIGURE 16. Arctosa costenola sp. nov. Male. Pedipalpal bulb SEM (ICN-Ar 1378) a–c (a, ventral; b, retrolateral details, c,
detail of Pa). Scale bars: a, 150 µm, b–c, 50 µm. E embolus, MA median apophysis, Pa posterior arm, Ra retrolateral arm, ST
subtegulum, T tegulum, TA Terminal apophysis.
Arctosa denticulata Jiménez & Dondale 1984

Fig. 65
Arctosa denticulata Jiménez & Dondale, 1984: 115, figs 1–6. Male holotype from near Aguacachil Cave, Taxco, Guerrero, Mexico,
5 June 1981, M. Ramirez leg. (see remarks). Female paratype, same locality of holotype. Examined by photography.
Remarks. This species is the only Arctosa collected inside a cave. It could be an accidental occurrence since the
description made by the authors does not indicate any morphological adaptation for subterranean environments.
Regarding the depository of holotype, after consulting Dra. Jiménez, she reported that may be deposited in the
UNAM. However, the current curator Dr. E. González-Santillan provided information and pictures of paratypes,
from same locality and date, but none of them corresponds to the holotype. Therefore, here we conclude the holotype
is lost.
Emended diagnosis. The structure referred as a stout spur and a small tooth in the median apophysis by Jiménez &
Dondale (1984: fig. 3) is what we call here as posterior arm (Pa), and the small tooth, the retrolateral arm (Ra). Another
small tooth on the terminal apophysis, used diagnostically too, is recognized here as the minute palea protuberance.

This protuberance is also present in Arctosa jibarosa sp. nov., Arctosa costenola sp. nov. and Arctosa tenebrosa.
Females can be distinguished by the largest and sinuous stalk of spermatheca (Jimenez & Dondale 1984: fig. 2).
Description. Detailed description in Jiménez & Dondale (1984: 115‒116).
Distribution. Mexico (Aguacachil Cave, Guerrero State, Fig. 65)
FIGURE 17. Arctosa jibarosa sp. nov. Male pedipalp holotype (MUSM-ENT 500410) a–c (a, ventral; b, retrolateral; c, anterior).
Scale bars: a–c, 0.25 mm. CMS, cymbium macrosetae, E embolus, Pa posterior arm, Pp palea protuberance, Ra Retrolateral
arm, TA terminal apophysis.
Arctosa jibarosa sp. nov.

Figs 2g, 17‒21, 64
Type Material. Male holotype form Yuracyacu, 5°57'0.91"S 77°14'1.34"W, 828 m, 9.ix.2008, Moyobamba, San
Martín, Peru, W. Paredes & C. Albujar leg., (MUSM-ENT 500410). Female paratype from Campamento Cashiriari
2, La Convención, Cusco, Peru, 11°51'51.3"S 72°45'45.6"W, 578 m, 2.ix.1997. S. Cordova leg. (MUSM-ENT
505269).
Other material examined. COLOMBIA: Meta: 5♂, Acacias, Vereda La Esmeralda, Finca Versalles,
4°01'14.85"N 73°42'32.37"W, 514 m, 21–25.iv.2004, E. Florez et al. leg. (ICN-Ar 2754); 1♂, Cubarral, Vereda
Vergel Alto, ca. 3°47'47.92"N 73°51'12.99"S, 1100 m, 1–3.vi.1996 (ICN-Ar 2723); 1♂, ditto, (ICN-Ar 6622); 2♂,
Villavicencio, Vereda la Vanguardia, Pozo Azul, ca. 4°10'42.45"N 73°37'18.09"W, 16–17.iv.2005, (ICN-Ar 5642);
8♂, San Martín, Vereda el Manantial, Finca la Marina, ca. 3°40'44.91"N 73°40'58.24"W, 28.x.2006, N. Rodríguez
leg. (ICN-Ar 5650). Boyacá: 5♂, Santa María, Vereda Caño Negro, Cuchilla Negra (pastizal), ca. 5°57'51.12"N
74°19'52.86"W, A. Gomez leg. (ICN-Ar 5642). ECUADOR: Napo, 1♀, Tena, Puerto Napo, Estación Biológica
Jatun Sacha, 1°03'57.5"S 77°37'00.2"W, 410 m, 1–5.xii.2009, C. Grismado & F. Labarque leg. (MACN-Ar 37822).
PERU: San Martín: Moyobamba, Yuracyacu, 5°56'34.11"S 77°08'53.44"W, 805 m, 7.ix.2008, W. Paredes & C.
FIGURE 18. Arctosa jibarosa sp. nov. Male pedipalp (MUSM-ENT 504111) a–c (a, prolateral; b, ventral; c, retrolateral).
Pedipalp bulb d–i (d, prolateral; e, ventral; f, retrolateral; g, anterior; h, basal expanded; i, posterior). Scale bars: a–c, 0.2 mm,
d–i, 200 µm. C conductor, DMA depression of median apophysis, E embolus, Pp palea protuberance, Pa posterior arm, Ra
retrolateral arm, RS row of setae, ST subtegulum, TA terminal apophysis.

FIGURE 19. Arctosa jibarosa sp. nov. Epigynum, paratype (MUSM-ENT 505269) a–d (a, ventral; b, lateral; c, posterior; d,
anterior). Scale bars: 91–97, 0.25 mm. CD copulatory duct, FD fertilization duct, H hood, HS head of spermatheca, MS median
septum, SS stalk of spermatheca, TS transversal septum, VC vulval chamber.
Albujar leg. (MUSM-ENT 500411); 1♂1♀, Moyobamba, Yuracyacu. 5°57'41"S 77°14'01.34"W, 828 m, 09.ix.2008,
C. Albujar leg. (MUSM-ENT 505000). Huánuco: 7♂2♀, Panguana Biological Station Rio Yuyapichis, 9°37'S
74°56'W, ca. 236 m, 21.i–18.ii.1984, M. Verhaagh leg. (MUSM-ENT 504111); 1♂, ditto, 26.xi–24.xii.1983, M.
Verhaagh leg. (MUSM-ENT 505001). Junin: 1♀, Chanchamayo, Fundo La Génova, Campo Guaba, ca. 11°05'S
75°20'W, ca. 771 m, xi.2009, A. Giraldo leg. (UNALM 326). Madre de Dios: 1♂, Nueva Arequipa, 12º54'24.3"S
69º59'32.2"W, 231 m, C. Penha & F. Meza leg. (MUSM-ENT 500975).
Etymology. The specific name is a noun in apposition honoring the “Jibaros” indigenous people, who established
populations after leaving their Chachapoyas native area in the northern of Peru. “Jibaros” were famous for shrinking
the heads of enemies, which are kept as trophies (Karsten 1923). This name is chosen in reference to the reduced
size of the species, if compared to other Arctosa recorded herein.
Diagnosis. Males of Arctosa jibarosa resembles those of Arctosa sapiranga Silva & Lise, 2009 by the shape
of the median apophysis, but can be distinguished by the posterior arm (pa) of the median apophysis with a wider
base and a pointed tip (Fig. 17a); the terminal apophysis beak-shaped (Fig. 18b), and a minute, triangular palea
protuberance (Fig. 18b, d). Females of Arctosa jibarosa are easily recognizable among all the neotropical species
by the MS elevated, visibly larger than the small transversal septum, which is straight in the outer border (Fig. 19a),
and the atrium triangular and sinuous (Fig. 19a). Additionally, the stalk of spermatheca (SS) is thrice as narrow than
the head of spermatheca (Figs 2g, 19b, 20b).
Description. Male. Holotype (MUSM-ENT 500410). Carapace light brown with an irregular, yellow longitudinal
median band with two wider parts, the first one behind the ocular area and the second, surrounding the thoracic
groove (Fig. 20a), and with discontinuous submarginal bands (Fig. 20g). Carapace covered with short white setae,
except in the longitudinal median band. Ocular area black, covered with long white setae and some black ones.
Sternum yellow, sparsely covered with brown bristles, more abundant on the margins (Fig. 20d). First row of
eyes slightly procurved and shorter than the second row (Fig. 20h); AME equal in size with ALE, ALE on small
tubercles, PME separated by more than half their radii. Labium light brown, covered with brown bristles the same
color as the endites; endites light brown at the base and yellow at the borders. Boss short and poorly visible,
chillum membranous and divided. Legs yellow, darker distally. All femora yellowish with annuli; patellae, tibiae,
and metatarsi darker than the femora, with annuli (Figs 20d, g). All tarsi without scopula, nor spinules on tarsi III
and IV; trichobothria in tibiae, tarsi and metatarsi distally increasing in size. Dorsum of opisthosoma olive-gray,
darker near the pedicel, with long white bristles in the middle and black on both sides, cardiac mark pale-yellow,
lanceolate; venter yellow with some brown dusk patches; spinnerets yellow, AS larger than PS.
FIGURE 20. Arctosa jibarosa sp. nov. Male habitus (MUSM-ENT 504111) a, d, g–h (a, dorsal; d, ventral; g, lateral; h, frontal).
Epigynum (MUSM-ENT 504111) b, e (b, dorsal; e, ventral). Female habitus paratype (MUSM-ENT 505269) c, f, i (c, dorsal; f,
ventral; i, lateral). Scale bars: a, d, g, 1 mm, b, e, 0.1 mm, h, 200 µm, c, f, i, 1 mm. SS stalk of spermatheca.
Pedipalp (Figs 17‒18, 21) with a cylindrical tibia, about two times longer than wide, a ventral patch of thin
bristles on the femur. Cymbium piriform (Figs 17a, 18c); subtegulum rounded, tegulum with a curved basal notch
with a membranous distal border. Terminal apophysis sclerotized, palea protuberance a sharp projection directed
distally (Figs 18b, 21a), rounded and membranous at the base. Base of the embolus lobulated, straight and tapering
towards its thin extreme and directed transversally, not twisted in any part; median apophysis with a Pa pyramidal,
with a wide base, dorsal groove poorly visible in apical view (Figs 17c, 18g,); retrolateral arm projection of the
median apophysis rounded in retrolateral view (Fig. 17b). Sperm ducts slightly sinuous through the tegulum in a
clockwise way. Conductor membranous (Fig. 18e, g).
Leg formula IV>II>III>I. Spination pattern: femur I p0-0-1, d1-1-1, II p0-0-1 d1-1-1, r0-0-1, III p0-1-1, d1-
1-1, r0-1-1, IV p0-1-1, d1-1-1, r 0-0-1; patella I d1bristle-1bristle ap, II d1bristle-1bristle ap III p1 d1bristle-1
bristle ap r1, IV p1 d1bristle-1ap r1; tibia I p1-1 d1bristle v2-2-2ap r1v-1, II p1-1 d1bristle v2-2-2ap III, p1d-1

d1r-1 r1-1 v2-2-2ap, IV p1d-1 d1r-1 r1-1 v2-2-2ap; metatarsus I pv1-1-2ap d1bristle r0-1-2ap v2-2-2, II p1-1-2ap
r0-1-2ap v2-2-1ap, III p1d-1d-2ap r1d-1-2ap v2-2-1ap, IV p0-1d-1ap d2-0-2 r1d-v1-1d-1v-1ap v2-2-1ap.
PME 0.24, PLE 0.18, interdistances: AME-AME 0.06, AME-ALE 0.04; PME-PME 0.14, PME-PLE 0.20; POQ
length 0.34, POQ posterior width 0.40, POQ anterior width 0.30. Chelicerae: length 0.80. Legs: length of segments
femur + patella + tibia + metatarsus + tarsus = total length: pedipalp 0.74 + 0.36 +0.38 + - + 0.66 = 2.14, I 1.26 +
0.60 + 1.08 + 1.02 + 0.64 = 4.60, II 1.30 + 0.68 + 1.00 + 1.04 + 0.68 = 4.70, III 1.20 + 0.58 + 0.92 + 1.10 + 0.68 =
4.48, IV 1.64 + 0.74 + 1.38 + 1.76 + 0.90 = 6.42.
Female. Paratype (MUSM-ENT 505269). Prosoma, opisthosoma and leg coloration and setae arrangement as
in male.
Genitalia (Figs 19, 20b, e) with transversal septum broad and rounded laterally, anterior part of the MS depressed
in apical view (Fig. 19d). Atrium narrow, parallel to median septum, which is longitudinally bulged. Copulatory
opening located at border of the TS (Fig. 19b). Fertilization ducts thin, membranous, originated posterodorsally;
spermatheca rounded, stalks short, curved distally, quite thrice as thin as the HS, base of spermatheca wide.
FIGURE 21. Arctosa jibarosa sp. nov. Male pedipalp SEM (MUSM-ENT 504111) a–e (a, ventral; b, TA detail; c, MA detail;
d, apical cymbium setae; e, papillae on MA). Scale bars: a, 100 µm, b–d 50 µm, e, 15 µm. E embolus, C conductor, P palea, Pa
posterior arm, Ra retrolateral arm, ST subtegulum, T tegulum, TA terminal apophysis.
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-d1 d1-1-1 r0-0-1d, II p0-0-1d d1-1-1 r0-1-1 , III
p0-1d-1d d1-1-1 r0-1d-1d, IV p0-0-1d d1-1-1 r0-0-1d; patella I p1 d1bristle-1bristle ap, II p1, d1bristle-1bristle
ap, r1; III p1 d1bristle-ap1 r1, IV p1 d1bristle-ap1 r1; tibia I p1-0 d1bristle-1bristle v2-2-2 r0-1-0, II p0-1d-1
d0-1bristle v2-2-2ap; III, p1-1 d1r-1 r1-1 v1-2-2ap, IV p1-1 d1r-1 r1d-1 v1-2-2ap; metatarsus I pd1-v1-1d-0-2
r0-1-2 v2-0-1, II p1d-1d-2ap r0-2-2ap v2-0-1ap, III p1d-1-2ap d1bristle-0 r1-1-2ap v2-2-1ap, IV p1-1-2ap rd1-
1v-1d-1v-2ap v2-2-1ap.
PME 0.24, PLE 0.16, interdistances: AME-AME 0.04, AME-ALE 0.04; PME-PME 0.12, PME-PLE 0.20; POQ
length 0.36, POQ posterior width 0.54, POQ anterior width 0.30. Chelicerae: length 0.78. Legs: length of segments
femur + patella + tibia + metatarsus + tarsus = total length: pedipalp 0.70 + 0.34 + 0.42 + - + 0.54 = 2.00, I 1.30 +
0.64 + 1.10 + 0.98 + 0.66 = 4.68, II 1.22 + 0.60 + 0.96 + 0.98 + 0.60 = 4.36, III 1.10 + 0.52 + 0.82 + 0.94 + 0.54 =
3.92, IV 1.60 + 0.68 + 1.34 + 1.64 + 0.76 = 6.02.
Variation. Males N = 9 (range, mean±s.d.): TL 3.41 – 3.92, 3.57±0.06; CL 1.92 – 2.17, 2.00±0.03; CW 1.33
– 1.58, 1.44±0.03.
Distribution. Colombia (Meta), Ecuador (Napo) and Peru (San Martín, Huánuco, Junín, Cusco and Madre de
Dios) (Fig. 64).
Arctosa littoralis (Hentz, 1844)

Fig. 65
Lycosa littoralis Hentz, 1844: 388, pl. 17, fig. 9. (Female holotype from North Carolina, USA, probably deposited at BMNH,
following Saunier (1980), lost according to Dondale & Redner 1983a).
Lycosa maritima Hentz, 1844: 389, pl. 17, fig. 10. Females syntypes from “South Carolina, Massachusetts and probably all
the Atlantic coast”, USA, probably deposited at BSNH, lost according to Dondale & Redner 1983a). Synonymized by
Montgomery 1902.
Lycosa cinerea: Emerton 1885: 488, pl. 47, fig. 3; Emerton 1902: 73, figs 177–178; Montgomery 1902: 555, pl. 29, figs 17–18;
Chamberlin 1908: 281, pl. 20, fig. 6. All misidentified.
Arctosa trifida F.O. Pickard-Cambridge, 1902: 330, pl. 31, figs 24–25. Male holotype from Teapa, Mexico, female paratypes
from Santa Ana and Guatemala City, Guatemala, BMNH, not examined). Gertsch & Wallace 1935: 5. Synonymized by
Dondale & Redner 1983a.
Trochosa cinerea: F.O. Pickard-Cambridge 1902: 331, pl. 31, figs 28–29; Montgomery 1904: 305, pl. 20, fig. 43.
Arctosa littoralis: Chamberlin 1924: 673; Gertsch 1934: 7; Kaston 1948: 320, figs 1050, 1070–1071; Roth & Brown 1976: 61,
figs 3, 6, 8; Dondale & Redner 1983a: 24, figs 65–74; Dondale & Redner 1990: 294, figs 479–487; Paquin & Dupérré 2003:
159, figs 1751–1753; Jiménez, Chamé-Vasquez & Palacios-Cardiel 2022: 198, figs 9–10.
Arctosa panamana Petrunkevitch, 1925: 179, fig. 97. Female holotype from Remedios, Panama, 1924, J.M. Valentine leg.,
PMNH, not examined. Synonymized by Dondale & Redner 1983a.
Arkalosula panamana: Roewer 1955: 232.
Remarks. Specimens of Arctosa littoralis dig burrows about 25 cm deep in suitable conditions (Gertsch 1934). It
was the first Neotropical species described from sandy and humid habitats. The similarities between A. littoralis and
Arctosa serii led to several misidentifications, and the former has been confused with Arctosa cinerea as well. The
correction of these identification errors (Chamberlin 1924) lead to the realization that the A. littoralis records from
eastern United States were in fact A. cinerea, and that A. littoralis is distributed north-westward from the USA to
Central America (Petrunkevitch 1925; Dondale & Redner 1990).
Emended diagnosis. Females of A. littoralis are distinguished from other neotropical species by having anchor-
shaped median and transverse septa of the epigyne (Roth & Brown 1976: fig. 6). Males have the posterior arm of the
median apophysis spatulate (Roth & Brown 1976, fig. 3).
Description. See Dondale & Redner (1983a: 294‒296).
Distribution. Nearctic and Neotropical regions, in the neotropics from western Mexico to Panama. Updated
Neotropical records obtained from species previously synonymized (Dondale & Redner 1983a; Jiménez et al. 2022;
Petrunkevitch 1925) (Fig. 65).

Arctosa mineira sp. nov.
Figs 2e, 22‒23, 66
Type material. Female holotype from Parque Nacional da Serra do Cipó, 19°37'S 43°21'W, 21–27.vii.2002,
Jaboticatubas, Minas Gerais, Brazil, E.S.S. Álvares leg. (UFMG 6319).
Etymology. The species epithet refers to people native from the Brazilian state of Minas Gerais.
Diagnosis. Arctosa mineira sp. nov. can be distinguished from all other members of the genus by the extremely
reduced transversal and median septa (MS, Figs 22a, 23b), and head of spermatheca piriform, with vulval chamber
as large as the head of spermatheca (HS, Figs 22d, 23b).
FIGURE 22. Arctosa mineira sp. nov. Epigynum (UFMG 3619) a–d (a, ventral; b, dorsal; c, posterior; d, anterior) Scale
bars: a–d 0.25 mm. FD fertilization duct, HS head of spermatheca, MS median septum, SS stalk of spermatheca, VC vulval
chamber.
Description. Male. Unknown.

Female. Holotype (UFMG 6319). Carapace brown covered by white, short, and black bristles except into
the irregular yellow longitudinal median band. Sternum yellow longer than wide, covered by short black bristles
abundant in the marginal area and sparsely covered in the central area (Fig. 23c). Ocular area black and the cephalic
portion of the prosoma covered by sparse large black bristles (Fig. 23a); four long black bristles on the clypeal
condyle anterior to AER. First row of eyes straight, shorter than the second row; AME slightly larger than the ALE;
PME separated by a length greater than their radii. Chelicerae brown covered frontally with long brown hairs and
the rest of it by long white hairs; chillum conspicuous and membranous, divided with the mesial portion sclerotized;
condyle notorious and clearer than the chelicerae; endites light brown and convergent; labium darker than the
endites; both endites and labium yellowish in their distal portion. All segments of legs yellow except on tarsi which
are darker; annuli on all portions of legs except on tarsi (Fig. 23a). Opisthosoma all covered by tiny white bristles,
the dorsum is olive and covered by tiny black large bristles; cardiac mark absent; venter yellow sparsely covered by
black bristles; spinnerets yellow; anterior spinnerets and posterior spinnerets cylindrical and equal in size.
Epigyne (Figs 22, 23b, d), median septum square covered by white bristles, transversal septum thin and less
noticeable; the atrium is absent. Copulatory opening is located at the intersection of both septa. In the dorsal view,
the fertilization duct is membranous, curved, and thin. (Figs 22b, c). The head of spermatheca are notoriously
piriform and large; the stalk is narrow than the head of spermatheca and curved directing outwards; vulval chamber
ventrally directed visible in apical view (Fig. 22d), rounded and as bigger than the head of spermatheca (Fig 22b).
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1 d1-1-1, II p0-0-1 d1-1-1 r0-0-d1, III p0-d1-d1 d1-
1-1 r0-d1-d1, IV p0-0-d1ap d1-1-1 r0-0-d1ap; patella I p1 d1-ap(1 bristle), II p1 d1-ap(1 bristle), III p1 d1bristle-
1ap r1, IV p1 d1 bristle-1ap r1; tibia I p2-1 d1 bristle-1 bristle v2-1-2ap r0-0-1, II pd1-1 d1bristle v1-1-2ap, III
p1-v1 dr1-1 r1-1 v2-1-2ap, IV p1-d1 d1-1 r1-1 v2-2-2ap; metatarsus I p0-2-2 r0-2-2 d1bristle-1bristle v2-0-1ap,
II p0-2-2 d1bristle-1bristle r1-2-2 v2-0-1ap, III pd1-d1-2 d1bristle-1bristle rd1-1-2 v2-2-1ap, IV p0-0-1-1-2ap
d0-r1-1ap r1-1-1-4ap v1-p1-r1-0-0-r1-p1-2ap.
FIGURE 23. Arctosa mineira sp. nov. Female habitus (UFMG 3619) a, c, e (a, dorsal; c, ventral; e, lateral) Epigynum b, d (b,
ventral; d, dorsal) f, eggsac. Scale bars: a, c, e 1mm, b, d 0.1 mm, f 1 mm. HS head of spermatheca.

0.56. POQ long 0.56, POQ posterior width 0.80, POQ anterior width 0.50. Chelicerae: 1.50 length. Opisthosoma:
length 1.72, width 0.52. Legs: (femur + patella+tibia + metatarsus + tarsus = total length): pedipalp 1.10 + 0.60 +
0.70 - + 0.80 = 3.26, I 2.06 + 1.00 + 1.60 + 1.70 + 1.06 = 8.68, II 1.90 + 0.98 + 1.46 + 1.62 + 1.00= 8.12, III 1.84
+ 0.94 + 1.36 + 1.88 + 1.02 = 7.69, IV 2.52 + 1.10 + 2.00 + 2.74 + 1.34 = 10.91.
Distribution. Known only for the type locality (Fig. 66).
Natural history. The female holotype was collected with a two-valved egg sac, which is covered by a thin layer
of silk (Fig. 23f).
Arctosa pacaya sp. nov.

Figs 24‒27, 62
Type Material. Male holotype from Pithecia, 5°06'S 74°50'W, 100 m, 11.viii.1989, Parque Nacional Pacaya Samiria,
Loreto, Peru, D. Silva D. leg (MUSM-ENT 505356).
Etymology. The species epithet is dedicated to the Pacaya Samiria Natural Reserve (PSNR), a protected natural
wetland area. PSNR is the biggest protected flooded forest in the Peruvian Amazon, the biggest RAMSAR site in
Peru, and serves as habitat to 25% of the bird fauna reported from Peru (Isola 2007).
FIGURE 24. Arctosa pacaya sp. nov. Holotype male pedipalp (MUSM-ENT 505356) a–c (a, ventral; b, retrolateral; c, anterior).
Scale bars: a–c, 0.25 mm. CMS cymbium macrosetae, Pa posterior arm, Ra retrolateral arm, TA terminal apophysis.
FIGURE 25. Arctosa pacaya sp. nov. Male pedipalp (MUSM-ENT 505356) a–c (a, retrolateral; b, ventral; c, prolateral).
Pedipalp bulb d–f, h (d, retrolateral; e, ventral; f, prolateral; h, anterior). g, apical cymbium setae, i, protuberance on male
chelicerae. Scale bars: a–c, 0.2 mm, d–h, 100 µm, i, 125 µm. C conductor, DMA depresion of median apophysis, E embolus, P
palea, Pa posterior arm, SD seminal duct, ST subtegulum, T tegulum, TA terminal apophysis.

FIGURE 26. Arctosa pacaya sp. nov. Male habitus (MUSM-ENT 505356) a–c (a, dorsal; b, ventral; c, lateral). Pedipalp,
expanded d, e (d, prolateral; e, anterior). Scale bars: a–c, 1 mm, d, e, 200 µm. BH basal hematodocha, CMS cymbial macrosetae,
DH distal hematodocha, E embolus, Pa posterior arm, SD seminal duct, ST subtegulum, T tegulum, TA terminal apophysis.
Diagnosis. Males of Arctosa pacaya resemble Arctosa sapiranga in dorsal pattern of habitus but can be
recognized by the wider carapace and the tibia/metatarsus I and II whitish (Fig. 26a, c); the male palpus of A.
pacaya have the embolus twisted once in the middle part (Fig. 25b), and a T-shaped posterior arm (Pa) of the median
apophysis (Figs 24a, c).
FIGURE 27. Arctosa pacaya sp. nov. Male pedipalp SEM (MUSM-ENT 505356) a–c (a, ventral; b, TA detail; c, Pa detail).
Scale bars: a, 150 µm, b, 100 µm, c, 50 µm. C conductor, E embolus, MA median apophysis, P palea, Pa posterior arm, Ra
retrolateral arm, ST subtegulum, T tegulum, TA terminal apophysis.
Description. Female. Unknown.

Male. Holotype (MUSM-ENT 505356). Dorsum of Prosoma light brown sparsely covered with tiny black setae
and a pale yellow longitudinal median band tapering to fovea; non-recognizable submarginal bands (Fig. 26a), four
long black bristles on the clypeal condyle. Ocular area is the same color as the longitudinal median band, covered
by a sparse white seta. First row of eyes procurved and shorter than the second row; AME slightly bigger than
ALE; ALE resting on tubercles distally oriented; PME separated by more than half the length of its radii. anterior to
AER. Sternum yellow sparsely covered with brown bristles (Fig. 26b). Labium yellow covered with brown bristles,
border of the labium and endite clearer than the base; chelicerae dark brown, darker than the endites and labium.
Condyle short and poorly notorious, pointed bulge in the outer margin of the cheliceral claw (Fig. 25i). Legs yellow,
becoming lighter distally. All femora are yellow with dusky irregular patches; only the leg I white from tibiae to tarsi
(Fig. 26c). All tarsi with no scopula, spinules on tarsi III and IV; trichobothria in tibiae, tarsi and metatarsi distally
increasing in size. Opisthosoma dorsal and lateral shield olive gray; a broad yellow longitudinal band runs along the
spinnerets finishing in not defined chevron pattern stripes, long black bristles in the yellow band; venter yellow with
some irregular dusky patches; spinnerets yellow.
Pedipalp (Figs 25a‒h, 26d, e, 27) femora slightly curved in the dorsal view. Cymbium with a short but strong
subterminal spine in the apical area (Fig. 26d). Sperm duct crossing the tegulum in a C-shaped pathway, subtegulum
visible and bowl shaped (Figs 25b, d). In ventral view, Median apophysis with an evident posterior arm T-shaped
ventrally projected (Figs 25b, e, 27c). The basal portion of the terminal apophysis is more sclerotized than the distal
and ribbed with its distal part sharpened (Figs 24a, 25b). Embolus is fusiform, sclerotized, and twisted once in its
median part (Figs 25b, e, 27a).

Leg formula IV>I>II>III. Spination pattern: femur I p0-0-2 d1-1-1 r0-1-1, II p0-0-2 d1-1-1 r0-1-1, III p0-1-1
d1-1-1 r0-1-1, IV p1-0-1 d1-1-1 r0-0-1; patella I p1 d1bas-1ap r1, II p1 d1bas-1ap r1, III p1 d1bas-1ap r1, IV
p1 d1bas-1ap r1; tibia I p1-1v v2-2-2ap r1-1v, II p1-1 d1-(1 bristle) r1-1 v2-2-2ap III, pd1-1 d1-1 r1-1 v2-2-2ap,
IV p1-1 d1-1 r d1-1 v2-2-2ap; metatarsus I p0-1-2ap r0-12ap v2-2-1ap, II p0-1-2ap r0-1-2ap v2-2-1ap, III p1d-1d
r1d-1v-1d-0-1ap v2-2-1ap, IV p1d-1d-2ap r1d-1v-1d-0-2ap v2-2-1ap.
posterior width 0.90, POQ anterior width 0.56. Chelicerae: 0.92 length. Opisthosoma: length 2.17, width 1.33. Legs:
length of segments femur + patella+tibia + metatarsus + tarsus = total length: pedipalp 1.16 + 0.60 + - + 0.82 + 1.00
= 3.58, I 2.00 + 0.80 + 1.70 + 1.64+1.08 = 7.22, II 1.80 + 0.82 + 1.52 + 1.56+1.02 = 6.72, III 1.76 + 0.66 +1.30 +
1.66+0.92 = 6.30, IV 2. 40 + 0.92 + 2. 10 + 2.74+1.30 = 9.46.
Distribution. Known only for the type locality, Loreto, Peru (Fig. 62).
Arctosa sapiranga Silva & Lise 2009

Figs 28‒33, 66
Arctosa sapiranga Silva & Lise, 2009: 27, figs 124–133. Male holotype from Arroio Feitoria, 29°35'43"S 51°2'56"W, 234 m,
Sapiranga, Rio Grande do Sul, Brazil, 19.ii.2008, E.L.C. Silva leg., deposited in MCTP 19998, examined. Female paratype,
same data, MCTP 19999, examined.
Other material examined. BRAZIL. Tocantins: 1♂, Sandovalandia/Araguaçu, Fazenda Praia Alta II, ca.
10°47'27.99"S 49°37'13.25"W, 186 m, 5–13.vii.2001, L.S.Rocha leg. (IBSP 12008); Distrito Federal: 1♂, Fercal,
Mata (MT-2.6), ca. 15°36'07.14"S 47°52'26.02"W, 909 m, Â.S. Zerbini leg. (UnB 5456); Goiás: 7♂, Cavalcante,
Fazenda Miraflores, ca. 13°48'49.78"S 47°25'51.47"W, 804 m, 19.xii.2012, P.C. Motta et al. leg. (UnB 6843). Minas
Gerais: 1♀, Lavras, Cachoeira de Farias, ca. 17°26'20.81"S 41°50'9.50"W, (soil, riverbank, gallery forest), 931 m,
18.ii.1993, R.L.C. Baptista leg. (MNRJ 6512). São Paulo: 1♂1♀, Salesópolis, Estação Biológica de Boracéia,
23°37'51"S 45°52'11"W, (in grasses on the riverside), ~983 m, 28.ii–3.iii.1992, R.L.C. Baptista leg. (MNRJ 6501).
Paraná: 1♀, Pinhais, ca. 25°23'55.33"S 48°55'36.50"W, Serra da Farinha Seca, 1019 m, 15–20.ix.1995 (MCTP
7666). Rio Grande do Sul: 1♀, Santa Maria, Santo Antão, ca. 29°39'23"S 53°50'49.89"W, ~222 m, 2.ii.1989, J.S.
Fernandes leg. (MCTP 39791). ARGENTINA. Misiones: 1♀, San Javier, Arroio Guerrero, ca. 27°47'16.28"S
55°9'8.83"W, 106 m, 11–21.iv.1989, Equipe Garabi leg. (MCTP 3055). Entre Ríos: 1♀, Parque Nacional El Palmar,
ca. 31°51'51.42"S 58°15'32.41"W, ii.1981, P. Gollobof leg. (MACN-Ar 24197).
Emended diagnosis. Males of Arctosa sapiranga resemble Arctosa serii Roth & Brown, 1976 by the similar
palea region, but differ by the large, bulging, membranous palea protuberance (Pp) of A. sapiranga (Figs 29e, 32a;
Silva & Lise 2009: fig. 6, missindicated as term). Additionally, the combination of a triangular posterior arm and
bulging retrolateral arm of the median apophysis distinguishes A. sapiranga from A. serii. Females of A. sapiranga
are recognized from other neotropical species by a keel on median septum (Fig. 30a, f, g, i; Silva & Lise 2009:
fig. 9) like Arctosa perita (Dondale & Redner 1983a: fig. 46) but differs from the latter by the pear-shaped head of
spermatheca and very small vulval chamber (Figs 30e, 33b).
Description. Male and female were described by Silva & Lise, 2009. However, some remarks are presented
here since somatic and genitalic details were passed over or misinterpreted. The median apophysis is not corkscrew
shape (Silva & Lise, 2009; fig. 6) but an excavated triangle (Figs 28a, f, 31b, e). Likewise, the structure misnamed
as “term” (Silva & Lise 2009: figs 6, 12) is certainly a pronounced palea protuberance, diagnostic for males of A.
sapiranga (Figs 29e, h, 32a) and not a terminal apophysis. The conical papilla on the female epigynal plate is visible
and richly illustrated here (Figs 33a, d).
Variation. As we observed, São Paulo female specimens shown variation in the atrium which was wider than the
southern Brazilian specimens (Figs 30g‒j). Male pedipalps present slight variation on posterior arm and retrolateral
arm proportions (Figs 28a–c) when compared with São Paulo specimens (Figs 28f–h). Slight variations on the
posterior arm and retrolateral arm of specimens from São Paulo show a smaller projection and a slightly wider base
(Figs 28f‒h).
Distribution. Brazil (Rio Grande do Sul, and new records from the states of Tocantins, Distrito Federal, Goiás,
Minas Gerais, São Paulo, and Paraná) and Argentina (Misiones and Entre Ríos) (Fig. 66).
FIGURE 28. Arctosa sapiranga Silva & Lise, 2009. Male pedipalp. a–c. Pedipalp, holotype, Rio Grande do Sul (MCTP 19999)
(a, ventral; b, retrolateral; c, prolateral). d–e. Palpal bulb (d, anterior; e posterior). f–h. Paratype, São Paulo (MNRJ 6501), (f,
retrolateral, g, ventral, h, anterior). Scale bars: a–h, 0.25mm. DMA depression of MA, E embolus, MA median apophysis, Pa
Posterior arm, Ra Retrolateral arm, SD seminal duct, ST subtegulum, T tegulum, TA Terminal Apophysis.

FIGURE 29. Arctosa sapiranga Silva & Lise, 2009. Male. Pedipalp a–c. Pedipalp, holotype, Rio Grande do Sul (MCTP 19999)
(a, prolateral; b, ventral; c, retrolateral). d–h. Palpal bulb (MCTP 19997) (d, prolateral; e, ventral; f, retrolateral; g, posterior; h,
anterior). i. Chelicerae. Scale bars: a–c, 200 µm; d–h, 100 µm; i, 500 µm. E, embolus, DMA, depression of MA, Pa posterior
arm, Ra retrolateral arm, SF striae file, TA terminal apophysis, TF tegular furrow.
FIGURE 30. Arctosa sapiranga Silva & Lise 2009. Epigynum a–c. Paratype, Sapiranga, Rio Grande do Sul (MCTP 19999)
(a, ventral; b, dorsal; c, anterolateral). d–f. Variation, Sapiranga, Rio Grande do Sul (MCTP 39791) (d, ventral; e, anterior; f,
lateral). g–j. Variation Salesópolis, São Paulo (MNRJ 6501) (g, ventral; h, dorsal; i, lateral; j, anterodorsal). Scale bars: a–j,
0.25 mm. CO copulatory opening, FD fertilization duct, H hood, HS head of spermatheca, MS median septum, TS, transversal
septum, VC vulval chamber.

FIGURE 31. Arctosa sapiranga Silva & Lise, 2009. Male, habitus (MCTP 19998) a, b, g–h. Holotype, Rio Grande do Sul (a,
dorsal; d, ventral; g, lateral; h, frontal). Female, epigynum (MCTP 19999) b, e (b, ventral; e, dorsal), c, f, i. Female paratype
(MCTP 19999) (c, dorsal; f, ventral; i, lateral). Scale bars: a, d, g 1 mm; h, 500 µm; b, e, 200 µm; c, f, i, 2 mm.
Arctosa serii Roth & Brown, 1976

Fig. 65
Arctosa serii Roth & Brown, 1976: 61, figs 1‒2, 4‒5, 7. Male holotype from El Desemboque de los Seris, Sonora, Mexico,
29°30'N 112°34'W, 4.iii.1974, H. Flanders leg., AMNH, not examined). Dondale & Redner 1983a: 26, figs 57‒59, 61.
Arctosa littoralis: Chamberlin 1924: 673 (some specimens from Gulf of California misidentified, according to Roth & Brown
1976).
Remarks. Arctosa serii is sympatric with A. littoralis, but the latter prefers mostly sandy beach environments
(Esquivel-Bobadilla et al. 2013). Chamberlin (1924) confused some specimens of A. serii with A. littoralis because
both came from similar localities. However, the larger size of A. serii (about one third larger than A. littoralis) and
the paler body pattern also served to differentiate it.
Emended diagnosis. Because of its sympatry with A. littorais, males of A. serii are recognized among all
neotropical species by the shorter and sharp posterior arm of median apophysis, which is also not spatulated as in
A. littoralis (Roth & Brown 1976: fig. 2). Females of A. serii are distinguished from all neotropical species by the
shortest median septum in the epigyne and the narrow transversal septum (Roth & Brown 1976: figs 4, 5).
Description. See Roth & Brown (1976: 61‒63).
Distribution. Mexico (Baja California, Sonora), updated records based on Roth & Brown (1976: 63) (Fig.
65).
FIGURE 32. Arctosa sapiranga Silva & Lise, 2009. Male. Pedipalp (MCTP 28548) (a, ventral; b, detail TA; c, close up of
MA; d, detail Pa; e, protuberance on tegulum). Scale bars: a, 250 µm; b, e 50 µm; c, 100 µm; d, 25 µm. CMS cymbium seate,
E embolus, MA median apophysis, Pa posterior arm, Pp palea protuberance, Ra retrolateral arm, ST subtegulum, T tegulum,
TA terminal apophysis.
Arctosa tenebrosa (Keyserling, 1877) comb. nov.

Figs 1d, 34‒37, 64
Trochosa tenebrosa Keyserling, 1877: 665, pl. 8, fig. 41 (female holotype from St. Fé de Bogotá, Colombia, should be deposited
in the BMNH, not found).
Lycosa tenebrosa: Petrunkevitch 1911: 568.
Remarks. Keyserling described T. tenebrosa from one female from Bogotá, Colombia and presented drawings
containing the diagnostic pointed internal projection on epigynal plate (Keyserling 1877: fig. 41). We examined male
and female specimens collected near the type locality and the females’ epigyne matched the original illustration of
the species, but also correspond to the typical trapezoidal shape seen in Arctosa, instead of the T-shaped septum seen
in Trochosa (Brady 1979; Dondale & Redner 1990). Additionally, the males collected with A. tenebrosa females

does not have the curved posterior arm spur of the median apophysis, which is characteristic of Nearctic and
Neotropical Trochosa C.L. Koch, 1847.
Other material examined. COLOMBIA: Cundinamarca: 7♂3♀, Mosquera, Laguna La Herrera, ca.
4°41'28.69"N 74°16'36.50"W, 2600 m, 9.ii.2002, J. Martinez et al. leg. (ICN-Ar 3927); 1♀, ditto, (ICN-Ar 1937);
3♂, ditto, 9.iii.2002, J. Martinez et al. leg. (ICN-Ar 6221); Bolívar: 2♀, Galerazamba, near El Totumo, ca.
10°44'36.64"N 75°14'29.51"W, 5 m, x.1999, E. Florez et al. leg. (ICN-Ar 1449).
FIGURE 33. Arctosa sapiranga Silva & Lise, 2009. Female, epigynum (MCTP 28548) (a, ventral; b, dorsal; c, plumose setae;
d, conic papillae). Scale bars: a, 150 µm; b, 200 µm; c, 25 µm; d, 20 µm. CO copulatory opening, HS head of spermatheca, MS
median septum.
Diagnosis. Males of Arctosa tenebrosa can be recognized among all neotropical species by a lump at the base
of the median apophysis (Fig. 35d), gutter-shaped terminal apophysis (Fig. 35d, h) and tip of posterior arm of the
median apophysis (Pa) slightly curved apically (Figs 34b, 35d). Females can be recognized by the pointed internal
projection of the epigynal plate (Fig. 37b), and the head of spermatheca rounded in dorsal view, with the stalk of
spermatheca broader than in Arctosa conflicta (Figs 36c, 37e).
Description. Male. Holotype (ICN-Ar 1378). Carapace brown with dark brown radial pattern, yellow longitudinal
median band expanded in the fovea, pale submarginal bands, dorsal shield of prosoma covered with tiny black setae,
four long black bristles on the clypeal condyle anterior to AER (Fig. 37h). Sternum sparsely covered by long brown
setae. Ocular area black, covered by sparse black setae, two parallel yellow lines beside the longitudinal median
band just behind the ocular area. AME slightly larger than ALE; PME separated a radius apart. Anterior eye row
slightly procurved, quite equal to the length of the PME. Three promarginal and retromarginal teeth each. Brown
chelicerae darker than the endites and labium. Intense yellow sternum and coxae yellow. Labium and endites are
light brown. All femora with pale annuli, tibiae, metatarsus, and tarsus I-II dark brown different from the III–IV
which are lighter brown (Fig. 37g). Sparse scopulae on legs I and II, absent in III and IV. Spinules present only on
tarsi III and IV. All tarsi with trichobothria present. Opisthosoma dusky, with three paired yellow dots in the cardiac
mark area (Fig. 37a). Four triad of parallel yellow dots in the posterior half of the dorsal area; venter yellow with
brown dusky scattered spots.
Pedipalp (Figs 34a‒c, 35), tibia slightly curved in dorsal view. Cymbium with two large spines in the dorsal
distal area (Fig. 34c) and a row of seven pairs of macrosetae (Fig. 35i). Squared and visible subtegulum, with
a broad sperm duct diagonally crossing the tegulum from base to the apex (Fig. 35d). Basal lamellar mound on
median apophysis with an evident sclerotized prolateral arm. The basal portion of the terminal apophysis is more
sclerotized than the distal and ribbed in shape with its distal part peak shaped (Figs 35d, h). Embolus fusiform,
sclerotized and straight (Figs 34a, 35d).
FIGURE 34. Arctosa tenebrosa (Keyserling, 1877) comb. nov. Male. Pedipalp (ICN-Ar 1378) a–c (a, ventral; b, retrolateral; c,
anterior). Scale bars: a–c, 0.25 mm. CMS cymbium macrosetae, Pa posterior arm, Pp palea protuberance, Ra retrolateral arm,
T tegulum, TA terminal apophysis.
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1 d1-1-1 rd1, II d1-3-3, III d1-3-3, IV p1d-0 d1-1-3;
patella III p1 r1 d1bas-1ap, IV p1 r1 d1bas-1ap; tibia I p1-1v r0-1-0 v2-2-2ap, II p0-1-1 r0-1-1 v2-2-2ap III, p1-1
d1-1 r1-1 v2-2-2ap, IV p1-1 d1-1 r1-1 v2-2-2ap; metatarsus I p0-1-1ap r0-1-1ap v2-2-1ap, II p1-1-2ap r1-1-2ap
v2-2-1ap, III p1-2-2ap r1-2-2ap v2-0-1ap, IV p1-1-2ap r1-1v-1-0-0-2ap v2-0-1ap.

FIGURE 35. Arctosa tenebrosa (Keyserling, 1877) comb. nov. Male. Pedipalp (ICN-Ar 1378) a–c, i (a, ventral; b, retrolateral;
c, prolateral; i, dorsal setae). Palpal bulb d–h (d, ventral; e, retrolateral; f, prolateral; g, posterior; h, anterior). Scale bars: a–c,
0.2 mm, d–i, 100 µm. C Conductor, DMA depression of median apophysis, E embolus, Pa posterior arm, ST subtegulum, T
tegulum, TA terminal apophysis, TK tegular keel. * basal hump on median apophysis.
FIGURE 36. Arctosa tenebrosa (Keyserling, 1877) comb. nov. Female. Epigynum (ICN-Ar 12745) a–d (a, ventral; b, lateral;
c, dorsal d, anterior). Scale bars: a–d, 0.25 mm. FD fertilization duct, H hood, HS head of spermatheca, MS median septum, SS
stalk of spermatheca, VC vulval chamber (* pointed lateral side of atrium).
Measurements: TL 6.25, CL 3.09, CW 2.38, CH 0.91, TH 0.94. Eyes: AME 0.09, ALE 0.09, PME 0.22, PLE
0.19, POQ long 0.38, POQ posterior width 0.63, POQ anterior width 0.34. Chelicerae: length 1.16. Opisthosoma:
length 2.75, width 1.59. Legs: femur + patella+tibia + metatarsus + tarsus = total length: pedipalp 1.19 + 0.47 +
- + 0.56 = 2.22, I 2.25 + 1.25 + 2.09 + 1.97+1.09 = 8.65, II 2.09 + 1.09 + 1.75 + 1.84+1.03 = 7.80, III 1. 94 + 0.97
+1.47 + 1.88+1.00 = 7.26, IV 2. 41 + 1.16 + 2. 09 + 2.72+1.28 = 9.66.
Female. Paratype (ICN-Ar 12745). General pattern as in male, except by the coloration of the opisthosoma
where the pattern is fuzzy (Figs 37c, f, i). Anterior eye row straight, chillum divided. Dentition as in male.
Genitalia (Fig. 36, 37b, e) longitudinal and transversal septum not distinguishable from each other, with plate
covered by long white setae. Median septum is divided by a thin and perceptible sclerotized elevation (Figs 36a,
37b). Copulatory opening, narrow, tiny membranous and thinner fertilization duct than in Arctosa tenella comb.
nov. (Fig. 36d). Head of speratheca rounded and wider than the stalk of spermatheca. Spermatheca with curved
stalks outwards thicker than base of spermatheca; Vulval chamber visible in lateral view and directed ventrally (Fig.
36b).
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1 d1-1-1, II p0-0-1 d1-1-1, III p0-1d-1d d1-1-1
r0-1d-1d, IV p0-1d-1d d1-1-1 r0-0-1d; patella I d(1ap bristle), II d(1ap bristle), III p1 d1bas-1ap r1, IV p1 d(1
bristle)-1ap r1; tibia I p0-1-0 v2-2-2ap, II p1d-1 v2-2-2ap, III p1d-1 d1-1 r1-1 v2-2-2ap, IV p1-1 d1-0-1 r1-1 v2-
2-2ap; metatarsus I pv1-2-1 r0-1-1ap v2-0-1ap, II p0-0-1 r1-2-2 v2-2-1ap, III p1-2-2 r1d-1v-1d-2 v2-0-1ap, IV
p1d-1v-1d-0-2ap r1-1-2 v2-2-1ap.
Measurements: TL 7.33, CL 3.58, CW 2.67, CH 1.17, TH 1.08. Eyes: AME 0.12, ALE 0.12, PME 0.24, PLE
0.18, POQ long 0.34, POQ posterior width 0.64, POQ anterior width 0.40. Chelicerae: length 1.52. Opisthosoma:
length 3.33, width 2.17. Legs: femur + patella+tibia + metatarsus + tarsus = total length: pedipalp 1.19 + 0.63 +
0.75 - + 0.69 = 3.26, I 2.44 + 1.25 + 2.00 + 1.86 + 1.13 = 8.68, II 2.22 + 1.25 + 1.81 + 1.81 + 1.03= 8.12, III 2.03
+ 1.06 + 1.63 + 2.00 + 0.97 = 7.69, IV 2.91 + 1.25 + 2.41 + 3.06 + 1.28 = 10.91.
Distribution. Known only from Cundinamarca and Bolívar, Colombia (Fig. 64).

FIGURE 37. Arctosa tenebrosa (Keyserling, 1877) comb. nov. Male, habitus (ICN-Ar 1378) a, d, g, h (a, dorsal; d, ventral; g,
lateral; h, frontal). Female, epyginum b, e (b, ventral; e, dorsal) Female habitus c, f, i (c, dorsal; f, ventral; i, lateral). Scale bars:
a, d, g, 1 mm, h, 500 µm, b, e, 0.1 mm, c, f, i 2 mm. SS stalk of spermatheca. (* pointed lateral side of atrium).
Arctosa tenella (Keyserling, 1877) comb. nov.

Figs 38‒43, 68
Trochosa tenella Keyserling, 1877: 667, pl. 8, figs 42‒43. Male holotype from St Fé de Bogotá, Colombia, BMNH 1890.1036,
examined.
Lycosa tenella: Petrunkevitch 1911: 568.
Arctosa minuta F.O. Pickard-Cambridge, 1902: 331, pl. 31, figs 26‒27. Male holotype and female paratype from Guatemala,
BMNH, examined by photographs. Dondale & Redner 1983a: 21, figs 50‒56; Jiménez et al. 2018: 5, figs 9‒10. Syn.
Nov.
Arctosa cheluncata Petrunkevitch, 1925: 177, figs 94‒96. Female syntype from Remedios, Panama; two male, three females,
and one juvenile syntypes from Santiago, Panama; one male, one female, and one juvenile syntypes from Wilcox camp,
San Lorenzo River, Panama; one juvenile syntype from La Mesa, Panama, all in PMNH, not examined. Synonymized by
Dondale & Redner 1983a.
Arkalosula cheluncata: Roewer 1955: 232.
Other material examined: COLOMBIA: Bolívar: Turbaco: 4♂5♀, Jardín Botánico Guillermo Piñeres, 118 m,
14.ii.2020, D. V. Moreno, W. P. Munguia & A. Segovia leg. (ICN-Ar). Cauca: 1♂1♀, Guapi, Parque Nacional Isla
Gorgona, ca. 2°58'6.0"N 78°11'4.00"W, 5 m, vii.2003, A. Rico et al. leg. (ICN-Ar 2577). Caquetá: 1♀, Puerto Rico,
El Lobo, ca. 1°30'55.36"N 74°50'6.04"W, 232 m, 10.i.2020, M. Ribero leg. (ICN-Ar 12758). Guaviare: 1♀, San
José del Guaviare, Isla de la Laguna Negra, Vereda Playa Guio, ca. 2°34'19.44"N 72°42'53.85"W, 212 m, 27.x.2012,
(ICN-Ar 10071). Meta: 1♂, Villavicencio, ca. 4°08'36.26"N 73°55'56.69"W, 408 m, 15.x.2014, E. Quiñones leg.
(ICN-Ar 10061). Tolima: 1♀, San Luis, Cultivo de Guanabana, 4°08'25.26"N 75°06'10.32"W, 420 m, 27.iv.1992,
A. Castillo leg. (ICN-Ar 10045); 1♂, San Luis, 4°08'01.93"N 75°06'11.80"W, 400 m, 10.vi.1992, A. Castillo leg.
(ICN-Ar 10049). ECUADOR: Loja: Malacatos, ca. 4°12'58.0"S 79°11'04.00"W, 1481 m, 27.xii.2006, D. Mora leg.
(QCAZ). Pichincha: 1♂, Mindo, 0°02'S 78°46'4"W, 1500 m, 19.xi.1997, T. Romero leg. (QCAZ); 1♀, Palmeras,
ca. 0°13'44.53"N 78°42'08.35"W, 1930 m, 7.xi.1992, J. Mancheno leg. (QCAZ). Manabi: 1♀, El Carmen, Km 40
Via Santo Domingo-Chone, 250 m, ca. 0°16'31.42"S 79°35'06.70"W, 250 m, 28.xii.1995, Jaramillo leg. (QCAZ);
1♀, Rio Ayampe, 20 m, ca. 1°40'24.46"S 80°42'10.77"W, 27.xii.1992, L. de la Torre leg. (QCAZ). PERU: Tumbes:
2♂1♀, Corrales, Facultad de Ciencias Agrarias UNT, xi.2018, P. Castillo leg. (MUSM-ENT 513519). Cajamarca.
1♂, Huarapomayo, Choropampa, 6°21'29.61"S 78°19'30.46"W, 619 m, Bosque Seco, 16.ix.2015, J. Pacheco leg.
(MUSM-ENT 508928).
FIGURE 38. Arctosa tenella (Keyserling, 1877) comb. nov. Male palp (ICN-Ar 2577) (a, ventral; b, retrolateral; c, apical).
Scale bars: a–c, 0.25 mm. Pa posterior arm, Ra Retrolateral arm, TA terminal apophysis.
Diagnosis. Males of Arctosa tenella are recognizable among all other neotropical species by the embolus
concealed by the median apophysis, retrolateral arm of median apophysis grooved, curved, and pointed (Figs 38c,
39d); posterior arm of median apophysis lamellar and ventrally projected (Figs 39d, 43c); terminal apophysis
(TA) curved, thin and flat, with its basal part more sclerotized than the distal end (Figs 38a, 39b). Females of are
recognized by the transversal septum (TS) projected posteriorly and curved (Figs 42b), the head of the spermatheca
rounded, the stalk of spermatheca thin, and the fertilization ducts robust (Figs 41d, 42d).

FIGURE 39. Arctosa tenella (Keyserling, 1877) comb. nov. Male palp (ICN-Ar 2577) a–c, i (a, retrolateral; b, ventral; c,
prolateral; i, cymbium dorsal view). Pedipalp bulb (MUSM-ENT 508928) d–h (d, retrolateral; e, ventral; f, prolateral, g, basal,
h, apical,) Scale bars: a–c, 0.2 mm, d–h, 100 µm, i, 200 µm. C conductor, CMS cymbium setae, DMA depression of median
apophysis, E embolus, Pa posterior arm, Ra retrolateral arm, ST subtegulum, T tegulum, TA terminal apophysis.
FIGURE 40. Arctosa tenella (Keyserling, 1877) comb. nov. Male habitus (ICN-Ar 2577) a–c (a, dorsal; b, lateral; c, ventral).
Pedipalp, expanded d–e (d, prolateral; e, anterior). Scale bars: a–c 1 mm, d–e 150 µm. BH basal hematodocha, DH distal
hematodocha, MA median apophysis, Pa posterior arm, Ra retrolateral arm, ST subtegulum, TA terminal apophysis.
Description. Male (MUSM-ENT 508928). Carapace dorsally covered with little black hairs, dark yellow, with
irregular yellow longitudinal median band from the ocular area until fovea, discontinuous submarginal bands.
Sternum yellow sparsely covered with brown bristles (Fig. 40c). Ocular area black covered by sparse black setae
and some short white ones; four long black bristles on the clypeal condyle anteriorly of AER. First row of eyes
procurved and shorter than the second row; AME slightly bigger than ALE; ALE resting on tubercles distally
oriented; PME separated by more than half the length of its radii. Labium brown covered with brown bristles
the same color as the chelicerae; chelicerae with two promarginal teeth and three retromarginal all equal in sizes,
condyle short and poorly notorious; protuberance on cheliceral fang. Legs yellow; femora yellowish annuli (Fig.
40b, c). All tarsi with no scopula, spinules on tarsi III‒IV stronger than in tarsi I‒II; trichobothria in tibiae, tarsi and

metatarsi distally increasing in size. Opisthosoma olive gray, dorsally the area near the pedicel is with long white
bristles, pale gray lanceolate mark on the cardiac area flanked with irregular but symmetric yellow spots at both
sides, venter yellow, spinnerets dark yellow.
Pedipalp (Fig 38a‒c, 39a‒i, 40d‒e) with a cylindrical tibia, a ventral patch of bristles arranged in a dispersal
way on the femur. Cymbium piriform with six apical spines (Fig. 39i); tegulum with a rounded basal notch, lamellar
curved terminal apophysis (Figs 40d, 43a); posterior arm lamellar and a retrolateral arm of median apophysis with
dorsally keeled, beaklike, and curved in ventral view, conductor lamellar (Fig. 43b) The sperm duct runs diagonally
through the tegulum in a clockwise way (Fig. 39e). Embolus straight directed transversally from prolateral origin
and not twisted (Figs 39b, 43a)
FIGURE 41. Arctosa tenella (Keyserling, 1877) comb. nov. Epigynum (ICN-Ar 2577) (a, ventral; b, dorsal; c, posterolateral;
d, posterior). Scale bars: a–d, 0.25 mm. CO copulatory opening, HS head of spermatheca, FD fertilization duct, MS median
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-3 d1-1-1 r0-0-d1, II p1d-1d d1-1-1 r 1d-1d, III p1d-
1d d1-1-1 rd1-1d,IV p1d-0-1d d1-1-1 r0-0-1d; patella I p1 d1bristle-1bristle ap r1, II p1 d1bristle-1bristle ap r1,
III p1 d1bristle-1 bristle ap r1, IV p1 d1bristle -1bristle ap r1; tibia I p1-1 v2-2-2ap r1-1, II p1d-1 v2-2-2ap r1-1,
III p1d-1 d1-1 r1d-1 v2-1-2ap, IV p1-1 d1-1 r1-1 v2-2-2ap; metatarsus I p1d-1d-2ap r1d-1d-2ap v2-2-1ap, II p1d-
1d-2ap r1d-1d-2ap v2-2-1ap, III p1d-1-2ap r1d-1-2ap v2-2-1ap, IV p1d-1d-2ap r1d-1d-2ap v2-2-1ap.
PME 0.31, PLE 0.26, interdistances: AME-AME 0.11, AME-ALE 0.05, PME-PME 0.24, POQ length 0.54, POQ
posterior width 0.80, POQ anterior width 0.54. Chelicerae: length 1.55. Legs: length of segments (femur + patella
+ tibia + metatarsus + tarsus = total length): pedipalp 1.45 + 0.36 + 0.69 + - + 0.71 = 3.21, I 2.57 + 1.14 + 2.02 +
1.98 + 1.26 = 8.97, II 2.24 + 1.12 + 1.86 + 2.00 + 1.21 = 8.43, III 2.21 + 0.97 + 1.57 + 2.17 + 1.09 = 8.01, IV 3.12
+ 1.24 + 2.62 + 3.50 + 1.52 = 12.00.
Female (ICN-Ar 2577). Coloration as in male. All legs with the same color patterns. Coloration and setae
arrangement as in male.
FIGURE 42. Arctosa tenella (Keyserling, 1877) comb. nov. Female habitus (ICN-Ar 2577) a, c (a, dorsal; c, lateral). Epigynum,
b, d (b, ventral; d, dorsal) Scale bars: a, c, 2 mm, b, d, 0.1 mm. HS head of spermatheca, MS median septum, SS stalk of
spermatheca, TS transversal septum, VC vulval chamber.
FIGURE 43. Arctosa tenella (Keyserling, 1877) comb. nov. Male palpal bulb SEM (MUSM-ENT 508928) a–c (a, ventral; b,
conductor detail; c, MA details). Scale bars: a, 150 µm, b–c, 50 µm. C conductor, E embolus, Pa posterior arm, Ra retrolateral
arm, ST subtegulum, TA terminal apophysis.

pigyne hoodless, transversal septum curved (Fig. 42b); fertilization duct visible big and membranous (Figs 41b,
d). Copulatory duct narrows at extremes of transversal septum. Internally spermatheca is rounded, (Figs 41b, 42d),
vulval chamber easily to recognize and separate the stalk of spermatheca (Figs 41c, 42b).
Variation. Females N=5 (males N=5) (range, mean±s.d.): TL 6.07 – 9.40, 7.41±0.79; CL 3.43 – 4.04, 3.86±0.17;
CW 2.74 – 3.33, 3.03±0.15; (TL 5.10 – 7.74, 6.94±0.83; CL 2.80 – 4.04, 3.51±0.49; CW 2.30 – 3.03, 2.82±0.21).
Distribution. USA (Dondale & Redner 1983a), Mexico (Jiménez et al., 2018), Guatemala (F.O. Pickard-
Cambridge 1902), Colombia, Ecuador and Peru (Fig. 68).
Arctosa villa sp. nov.

Figs 44‒49, 64
Arctosa sp. nov.: Paredes-Munguia 2012: 42
Type Material. Male holotype from Pantanos de Villa, Chorrillos, Lima, Peru, 13.xi.1994. S. Córdova & J. Duarez
leg. (MUSM-ENT 500104), female paratype, from Totorales, Pantanos de Villa, Chorrillos, Lima, Peru, 12°12'22.0"S
76°59'19.5"W, ±24 m, 12.xi.2006. W. Paredes & C. Ruiz leg. (MUSM-ENT 500107).
Other material examined. PERU: Lima: 1♂, Pantanos de Villa—Totorales, 12°12'22"S 76°59'19.5"W,
24 m, 12.xi.2006, W. Paredes & C. Ruiz leg. (MUSM-ENT 514161); 1♂2♀, ditto, 23.xii.2003, W. Paredes leg.
(MUSM-ENT 505002); 5♀, ditto, 12.x.2003, W. Paredes leg. (MUSM-ENT 500108); 1♂2♀, Pantanos de Villa
21.vii.1994, D. Silva & J. Duarez leg. (MUSM-ENT 500106); 1♂1♀, Cañete, Planta de Licuefacción Pampa
Melchorita, 13°14'55,8"S 76°17'16.5"W, 146 m, 9–12.i.2013, V. Borda leg. (MUSM-ENT 507296). Lambayeque:
1♀, Lambayeque, Campo de cultivo Instituto de Desarrollo agrícola de Lambayeque (IDAL), Road to Ferreñafe,
6°40'12.32"S 79°47'6.58"W, 37 m, C. Albujar leg. (MUSM-ENT 505355).
Etymology. The specific name is a noun in apposition from the type-locality, the Pantanos de Villa Reserved
Zone, a salt marsh at the southern Lima, one of the most important salt marshes in the Peruvian central coast.
Diagnosis. Males of Arctosa villa can be recognized among all the neotropical species by the wide base of
posterior arm of the median apophysis (Pa) (Fig. 44a, f), and the membranous, thick palea protuberance (Pp) (Fig.
45a, e). The Arctosa sapiranga median apophysis resemble that of A. villa, but the base of the posterior arm of the
median apophysis is thinner in the former. Females of A. villa can be recognized by the transversal septum straight
(Fig. 46a), and the triangular head of spermatheca (Fig. 46b). Female epigynal plate of Arctosa perita (Latreille,
1799) (Dondale & Redner 1983a: figs 46, 49) and A. sapiranga (Silva & Lise 2009: figs 9, 10) look like that of A.
villa sp. nov., but it is wider anteriorly in A. villa.
Description. Male. Holotype (MUSM-ENT 500104). Carapace brown to light brown with pale black radiating
lines covered with short black setae; median band yellow with a few bristles, two blurry parallel yellow lines in
the longitudinal median band just behind the ocular area and pale yellowish irregular sub marginal band. Sternum
longer than wide, yellow, and covered with short brown bristles in the marginal area; the central are sparsely covered
by short brown bristles too (Fig. 47d). Ocular area black covered with short white bristles; four long black bristles
on the clypeal condyle anterior to AER; one large black bristle between the AME. First row of eyes is straight and
shorter than the first row; AME slightly larger than ALE; PME separated by the length of its radii. Chelicerae strong
and pale brown covered frontally with long brown hairs; SF present (Fig. 45i). Chillum membranous divided but not
conspicuous; boss easy visible. endites and labium pale yellow, whitish on its distal portions. All legs pale brown
except in coxae, trochanter and femora which are yellow. A sparse scopulae on legs I and II, absent in III and IV.
Spinules present only on tarsi III and IV. All tibiae, tarsi, and metatarsi with trichobothria present. Opisthosoma
dorsum light yellow covered with dusk brown mottled patches and brown bristles; poorly visible yellow lanceolate
mark on the cardiac area; five pairs of black spots that cover the rear half of the opisthosoma; venter all yellow,
cylindrical yellow spinnerets (Fig. 47d).
Pedipalp (Fig. 44a‒c, 44f, 45a‒f, 45h), cylindrical tibia slightly wider apically. Cymbium fusiform. Trapezoidal
subtegulum visible in ventral view, with a broad sinuous sperm duct diagonally crossing the tegulum from base to
the apex (Figs 45e‒f). Median apophysis with posterior arm evidently pyramidal (Figs 44f, 45h) and a sclerotized
mound basal in position. The basal portion of the terminal apophysis, sclerotized, sharpened, and curved, its distal
part membranous and lamellar shaped. Embolus fusiform, sclerotized and twisted once in its mesial part (Figs 44b,
45b).
FIGURE 44. Arctosa villa sp. nov. Male pedipalp (MUSM-ENT 500104) a–c, f (a, ventral; b, retrolateral; c, sub anterior;
f, anterior). Male prosoma d–e (d, lateral; e, frontal). Scale bars: a–c, f 0.25 mm, d–e 1 mm. CHI chillum, CND condylum,
DMA depression of median apophysis, E embolus, Pa posterior arm, Pp palea protuberance, Ra retrolateral arm, TA terminal
apophysis.

FIGURE 45. Arctosa villa sp. nov. Male pedipalp (MUSM-ENT 500104) a–c (a, retrolateral; b, ventral; c, prolateral). Pedipalp
bulb (MUSM-ENT 505002) d–f, h (d, retrolateral; e, ventral; f, prolateral; h, anterior). g, spinnerets; i, chelicerae. Scale bars:
a–c, 0.2 mm, d–f, h, 250 µm, g, 200 µm, i, 500 µm. DMA depression of median apophysis, E embolus, Pa posterior arm, Pp
palea protuberance, Ra retrolateral arm, SD seminal duct, SF striae file, TA terminal apophysis.
FIGURE 46. Arctosa villa sp. nov. Epigynum, paratype (MUSM-ENT 500107) (a, ventral; b, dorsal; c, anterior; d, posterolateral)
Scale bars: a–d, 0.25mm. BS base of spermatheca, FD fertilization duct, H hood, HS head of spermatheca, MS median septum,
VC vulval chamber.
Leg formula IV>I>III>II. Spination pattern: femur I p0-0-1 d1-1-1 r0-1d-1d, II p0-1-1 d1-1-1-1 r0-1-1-0-0-1
, III p0-1-1 d1-1-1 r0-1-1, IV p1d-1d d 1-1-1 r0-0-1d; patella I d1bristle-1 bristle II p1 d1bristle-1 bristle III p1
r1 d1bristle bas-1bristle ap, IV p1 r1 d1bristle bas-1bristle ap; tibia I p1-v1 d1 bristle- 1 bristle r1-1 v2-2-2ap, II
p1-1v r1-1 v2-2-2ap III, p1d-1 d1-1 r1-1 v2-1-2ap, IV p1-v1 d1-1 r1-1 v2-2-2ap; metatarsus I p1-1-1ap r0-0-1ap
v2-2-0, II p1-2-2ap r0-1-1ap v2-1-1ap, III p1-1-2ap d1-1-2ap v2-2-1ap, IV pd1-2-2ap v2-1r-1 r1d-1v-1d-1v-2ap.
0.54. POQ length 0.50, POQ posterior width 0.68, POQ anterior width 0.44. Chelicerae: length 1.22. Opisthosoma:
length 3.08, width 2.25. Legs: femur + patella+tibia + metatarsus + tarsus = total length: pedipalp 1.28 + 0.68 + - +

0.68 + 1.08 = 3.72, I 2.34 + 1.10 + 2.04 + 1.92+1.20 = 8.60, II 2.10 + 1.04 + 1.66 + 2.18+1.12 = 8.10, III 2.16 +
1.00 +1.76 + 2.12+1.18 = 8.22, IV 2. 84 + 1.22 + 2.56 + 3.38+1.50 = 11.50.
Female. Paratype (MUSM-ENT 500107). General pattern (Figs 47c, f, i, 49b) as in male, except by the
coloration of the opisthosoma which is less visible. Anterior eye row straight, chillum divided (Fig. 47h), dentition
as in male.
Genitalia (Fig. 46a‒d, 47b, e, 48a‒c), longitudinal and transversal septum not distinguishable from each other,
the plate is covered by a long white seta. In retrolateral view the anterior part of the transversal septum is depressed
ventrally (Fig. 46d). In ventral view, the atrium is divided by a thin and perceptible median septum keel (Fig. 46a).
Copulatory opening laterally located (Figs 48a, b) roughed surface on transversal septum (Fig. 48c). Fertilization
ducts thick and curved (Fig. 46b). Head of spermathecae triangular, base of spermatheca expanded, stalk of
spermathecae sinuous (Figs 46b, d). A short vulval chamber is visible in lateral view (Fig. 46d).
Leg formula IV>I>II>III. Spination pattern: femur I p0-0-1 d1-1-1, II p0-0-1 d1-1-1 r0-1-0, III p0-1d-1d d1-
1-1 r0-1d-1d, IV p0-0-1d d1-1-1 r0-1d-1d; patella I d1 bristle - 1bristle ap, II d1 bristle-1 bristle ap, III p1 d1bristle
bas-1ap r1, IV p1 d1 bristle bas-1ap r1; tibia I v2-2-2ap, II pd1-1 d1 bristle-1 bristle v1-2-2ap, III pd1-1 d1-1 r1d-
1d v1-2-2ap, IV p1d-1 d1-1 r1-1v v2-2-2ap; metatarsus I p0-1-2 r0-0-2ap v2-2-1ap, II p0-1d-2 r0-0-2 v2-2-1ap,
III p1-1-2 r1d-1d-2 v2-2-1ap, IV pd1-0-1d-2ap r1-1v-2 v2-2-1ap.
0.68. POQ length 0.56, POQ posterior width 0.84, POQ anterior width 0.58. Chelicerae: length 1.60. Opisthosoma:
length 1.48, width 1.08. Legs: (femur + patella + tibia + metatarsus + tarsus = total length): pedipalp 1.19 + 0.63 +
0.75 - + 0.69 = 3.26, I 2.44 + 1.25 + 2.00 + 1.86 + 1.13 = 8.68, II 2.22 + 1.25 + 1.81 + 1.81 + 1.03= 8.12, III 2.03
+ 1.06 + 1.63 + 2.00 + 0.97 = 7.69, IV 2.91 + 1.25 + 2.41 + 3.06 + 1.28 = 10.91.
FIGURE 47. Arctosa villa sp. nov. Male habitus (MUSM-ENT 500104) a, d, g (a, dorsal, d, ventral; g, lateral). Epigynum
(MUSM-ENT 500107) b, e (b, dorsal; e, ventral) Female habitus c, f, h–i (c, dorsal; f, ventral; h, frontal; i, lateral). Scale bars:
a, d, g, 1 mm, b, e, 0.2 mm, h, 500 µm, c, f, i, 2 mm.
FIGURE 48. Arctosa villa sp. nov. Epigynum SEM (MUSM-ENT 500108) a–c (a, ventral, b, Copulatory opening detail; c,
roughened surface on TS). Female tarsus I d (d, ventral leg spinula). Scale bars: a, 150 µm, b, 15 µm, c, 50 µm, d, 250 µm. CO
copulatory opening, H hood, MS median septum.
FIGURE 49. a. Pantanos de Villa semiaquatic vegetation, type locality of Arctosa villa sp. nov. b. Female live specimen
carrying eggsac of Arctosa villa sp. nov. a. modified from Paredes-Munguia (2012).

Variation. Females N=13 (males N=3) (range, mean±sd): TL 6.92-8.75 7.88±0.50; CL 3.5-4.75, 3.90±0.36;
CW 2.67-3.67, 2.96±0.29; (TL 6.25-7.58 6.83±0.68; CL 3.33-3.92 3.61±0.29; CW 2.58-2.92, 2.75±0.17).
Natural history. Arctosa villa has been collected in the type-locality on marshes, under debris of cattail
(Typha domingensis). Cattails are constantly falling and decomposing becoming debris which facilitates building
of spider retreats. All the records for this species are associated with a certain preference whose flooded vegetation
environments facilitates its permanence (Paredes-Munguia 2012: 42, fig. 3A–B). Adults were collected from August
to December. Females were more numerous than males and observed with egg sacs during August to September
(Fig. 49).
Distribution: Peruvian coastal region, between Lima and Lambayeque (Fig. 64).
Misplaced species
Hogna andina (Chamberlin, 1916) comb. nov.

Figs 50‒51, 62
Lycosa andina Chamberlin, 1916: 286, pl. 24, figs 2–3. Male holotype and female paratype from Tincochaca, Cuzco, Peru,
deposited in MCZ 285 and 286, respectively, examined by photos.
Arctosa andina: Roewer 1955: 230.
FIGURE 50. Hogna andina Chamberlin, 1916 comb. nov. Male pedipalp, holotype a–c (a, retrolateral; b, ventral; c, prolateral)
Collecting and ID labels d–e (d, type locality label; e, ID label). Scale bars: a–c, 1 mm. E, embolus, DMA depression of MA,
MA median apophysis, P palea, SD seminal duct, T tegulum, TA terminal apophysis, TL tegular lobe, VP ventral process.
FIGURE 51. Hogna andina Chamberlin, 1916 comb.nov. Female habitus, paratype a–b, d (a, ventral; b, prosoma dorsal; d,
opisthosoma dorsal) Epigynum c (c dorsal) Collecting and ID labels e–f (e, type locality; f, ID label). Scale bars: a, f, 5 mm, b,
1 mm, c, 0.5 mm. CM cardiac mark, H hood, HS head of spermatheca, MS median septum, SS stalk of spermatheca.
Remarks. The male and female description by Chamberlin (1916) presented detailed information on morphology
for both sexes. Holotype photos confronted with drawings of both epigynum and pedipalp (Chamberlin 1916:
286, pl. 24, figs 2–3) suggest transferring to Hogna. The male pedipalp has a rounded palea and a sickle shaped
terminal apophysis (Fig. 50b), which is diagnostic for the majority of Hogna species (Dondale & Redner 1990).
The epigynum presents an inverted T-shaped pattern and a smooth atrium (Fig. 51c). Different from Trochosa,
Hogna median septum is longer than wide, the transversal septum is narrow and about half the length of the median
septum (Dondale & Redner 1990). Additionally, H. andina hoods are parallel (Fig. 51c), different from Trochosa or
Varacosa Chamberlin & Ivie, 1942 species, which have curved hoods. The two parallel bands inside the longitudinal
median band are absent in both sexes (Fig. 51b), which also excludes placing this species in Trochosa. The hoods

in Birabenia Mello-Leitão, 1941, also called anterior pockets (Piacentini & Laborda 2013) are deep, as opposed
to the shallow ones in Hogna females. Likewise, the long legs and thick body (Fig. 51a, d) are diagnostic of
Hogna Dondale & Redner 1990), along with the three, instead of four retromarginal teeth of Birabenia. Finally, this
species should not be kept in Arctosa because the males do not have the reduced palea or TA in different degrees
of sclerotization (Dondale & Redner 1983a), and females possess the inverted T-shape of the epigynal plate, which
is triangular to trapezoidal in Arctosa (Dondale & Redner 1983a). The synapomorphies of Hogna used to remove
species from this genus and be transferred to Tigrosa are summarized in Brady (2012:182).
Diagnosis: Males of Hogna andina can be recognized by the curved cymbium (Fig. 50b) and short ventral
projection of median apophysis (Fig. 50c). Females can be recognized by the dilated and rounded head of spermatheca
(Fig. 51c).
Distribution. Known from four localities Cuzco, Peru including the type locality (Fig. 62, Chamberlin 1916)
Hogna pugil (Bertkau, 1880) comb. nov.

Fig. 67
Tarentula pugil Bertkau, 1880: 71, pl. 2, fig. 23 (syntypes from Teresópolis, Rio de Janeiro and São João del Rei, Minas Gerais,
Brazil, should be deposited in IRSNB, probably lost).
Allocosa pugil: Mello-Leitão 1943: 161.
Arctosa pugil: Roewer 1955: 230.
Arctosella pugil: Roewer 1960: 671.
Remarks. According to Bertkau’s (1880) description, this species has body size, coloration, scopulated tarsi on legs
and proportions of the prosoma, as well as the absence of internal markings in the dorsal median band, consistent
with diagnostic characters listed for Hogna by Brady (2012:182) and Logunov (2020:349). The rounded palea, the
sickle-shaped terminal apophysis, and the triangular and transversely positioned median apophysis (Bertkau 1880:
fig. 23a) also support the transfer of this species to Hogna (see characters in Logunov 2020).
Distribution. Brazil (Fig. 67, Bertkau 1880)
Prolycosides aussereri (Keyserling, 1877) comb. nov.

Figs 52‒56, 67
Tarentula aussereri Keyserling, 1877: 657, pl. 7, fig. 33. Female holotype from St. Fé de Bogota, Colombia, deposited in the
BMNH-1890/999, examined by photographs (Fig. 56).
Lycosa aussereri: Banks 1901: 223.
Arctosa aussereri: Roewer 1955: 230.
Lycosa proletaria Tullgren, 1905: 65, pl. 8, fig. 31. Female holotype from Tatarenda, ca. 19°6'23.24"S 63°30'51.22"W,
Chuquisaca, Bolivia, 1901-1902, E. Nordeskiöld leg., deposited in NRS, Sweden, 18, examined. Mello-Leitão 1941a: 134,
fig. 28. Syn. nov.
Avicosa proletaria: Roewer 1955: 236.
Schizocosa proletaria: Chamberlin & Ivie 1942: 38.
Tarentula workmani Strand, 1909: 277. Female holotype from Paraguay, ix.1880, should be in National Museum of Ireland,
Dublin, Ireland, lost). Syn nov.
Hygrolycosa workmani: Roewer 1955: 261.
Arctosella workmani: Roewer 1960: 671.
Lycosa planithoracis Mello-Leitão, 1938: 98, figs 12–13. Female holotype from Chacras de Coria, ca. 32°58'51.27"S
68°52'36.84"W, Mendoza, Argentina, Don P. Jorgensen leg., deposited in MLP 14059, examined. Syn. nov.
Hogna planithorax: Roewer 1955: 256. (unacceptable emendation).
Lycosa variolosa Mello-Leitão, 1941a: 134, fig. 32, pl. VI, fig. 26. Female holotype from Villa Nueva, ca. 32°26'41.65"S
63°15'16.17"W, Córdoba, Argentina, A. Parko leg., deposited in MLP 14702, examined. Syn. nov.
Lycorma variculosa: Roewer, 1955: 267 (Transferred from Lycosa, lapsus).
Lycosa melanostoma Mello-Leitão, 1941b: 208, fig. 7. Female holotype from Reconquista, ca. 29°8'5.20"S 59°38'32.11"W,
Santa Fé, Argentina, M. Birabén leg., deposited in MLP 15076, examined. Syn. nov.
Megarctosa melanostoma: Roewer 1955: 278.
Lycosa huachoi Mello-Leitão, 1942a: 431, fig. 5. Female holotype from Huacho, 11°6′S 77°36′W, Peru, deposited in MNRJ
02001, examined, and subsequently destroyed in the 2018 fire. Syn. nov.
Ocyale huachoi: Roewer 1955: 281.
Lycosa abalosi Mello-Leitão, 1942b: 391, fig. 4. Female holotype from Colonia Dora, ca. 28°36'25.32"S 62°57'9.57"W, Santiago
del Estero, Argentina, deposited in MLP 15310, examined. Syn. nov.
Sosilaus abalosi: Roewer 1955: 296.
Pirata abalosi: World Spider Catalog 2023
Lycosa soukupi Mello-Leitão, 1942a: 431, fig. 4 (female holotype from Huacho, 11°6′S 77°36′W, Peru, deposited in MNRJ
01364, examined, and subsequently destroyed in the 2018 fire. Syn. nov.
Sosilaus soukupi: Roewer 1955: 296.
Pirata soukupi: World Spider Catalog 2023
Other material examined. CUBA: Piñar del Rio: 1♂, La Bajada, Peninsula de Guanahacabibes, ca. 21°54'17.19"N
84°23'46.51"W, 12.v.2002, A. Sanchez leg. (IBSP 169614). Sabaera: 1♀, Maisí, Guantánamo, ca. 20°13'32.99"N
74°9'12.08"W, 12.iv.1998, A. Sanchez leg. (IBSP 169619), 1♀, ditto, 4.iv.1998, A. Sanchez leg. (IBSP 169610),
Santiago de Cuba: 2♂, La Puya, III Frente, ca. 20°8'45.38"N 76°17'38.79"W, 19.iii.2000, A. Sanchez-Ruiz leg. (IBSP
233858). VENEZUELA: Maracay: 1♀, Radio Sertaneja, ca. 10°15'32.50"N 67°38'27.46"W, 454 m, 16.iv.1984, W.
Manya leg. (MUSM-ENT 500166). COLOMBIA: Cundinamarca: 1♀, Cáqueza, confluence of Cáqueza and Rio
Negro rivers, ca. 4°23'53.50"N 73°54'20.33"W, 1520 m, 4.vi.2002, A. Rodriguez leg. (ICN-Ar 1867); 1♀, Sasaima,
19.x.1997, R. Casallas leg. (MLS 302); 1♀, Fusagasugá, ca. 4°20'42.55"N 74°21'42.56"W, 14.vi.1905, R. Guevara leg.
(MLS 291). Cauca: 1♀, El Tambo, ca. 2°27'07.59"N 76°48'41.21"W, 600 m, 29.xii.1998, R. Casallas leg. (MLS 436).
PERU: Tumbes: 1♂, Tumbes, Corrales, ca. 3°34'S 80°28'W, 5 m, ii.1983, P. Vinces & P. Castillo leg. (MUSM-ENT
500151); 1♀, Palmas, Rio Queiroz, ca. 3°34'00.22"S 80°12'50.71"W, 600 m, 8.x.1955, F. Blancas leg. (MUSM-ENT).
Piura: 1♀, near Huancabamba, Valle N. Pucara, 5°56.15'S 79°16.19'W, 1058 m, 20.viii.2006, G. Binford et al. leg.
(MUSM-ENT); 1♀, Ayabaca, ca. 4°38'S 79°43'W, 2715 m, 24.ix.2019, F. Blancas leg. (MUSM-ENT); 1♂, Chocan,
Ayabaca, ca. 4°41'31.23"S 80°34'04.42"W, 114 m, 2.x.1955, F. Blancas leg. (MUSM-ENT 500131). Amazonas:
1♀, Rodriguez de Mendoza, Limabamba, ca. 6°29'52.43"S 77°29'55.71"W, 1674 m, 6.xi.1986, R. Fernandez leg.
(MUSM-ENT 500133). Cajamarca: 2♂, Celendín, road to Llangoat, 6°42'51"S 77°51'55.9"W, 2048 m, 7.xii.2004,
W. Paredes leg. (MUSM-ENT); 1♂, La Victoria, ca. 6°57'37.85"S 78°06'53.24"W, 2638 m, 23.vi.1986, H. Ortega
leg. (MUSM-ENT 500159); 1♂, Baños del Inca, ca. 7°10'S 78°28'W, 2659 m, 6.viii.1984, I. Bohorquez leg. (MUSM-
ENT 500144); 1♂, San Andres de Cutervo, ca. 6°22'S 78°49'W, 2866 m, 16.iii.1989, D. Silva leg. (MUSM-ENT);
2♂, Rio Mashcón, ca. 6°12'16.65"S 78°38'16.03"W, 240 m, 4.x.1965, CC & BB leg. (MUSM-ENT); 1♀, Hacienda
Sunchubamba, 7°29'S 78°23'W, 2657 m, 2.xi.2009, D. Silva leg. (MUSM-ENT); 3♀, Cumbemayo, ca. 7°11'26.07"S
78°34'43.23"W, 3592 m, 27.xi.2004, W. Paredes leg. (MUSM-ENT 500154). Lambayeque: 1♂, Motupe, ca.
6°45'07.24"S 79°51'07.68"W, 24 m, 2.ix.2009 (MUSM-ENT); 1♀, Chiclayo, ca. 6°47'24.35"S 79°51'30.35"W, 21
m, 22.vii.2006, A. Albujar leg. (MUSM-ENT); 1♀, Motupe, Distrito de Olmos, ca. 5°59'15.81"S 79°44'42.91"W,
142 m, 21.ii.1955, Vellard leg. (MUSM-ENT 500129); 1♀, Lambayeque, Santa Maria dos Vales de Chiclayo, ca.
6°45'47.00"S 79°50'12"W, 27 m, 2.viii.2006, A. Albujar leg. (MUSM-ENT). La Libertad: 3♀, Chicama, Valle de
Chicama, ca. 7°52'59.94"S 79°13'00.04"W, 61 m, 1.ix.1978, P. Hocking & G. Martinez leg. (MUSM-ENT 500138);
3♂1♀, Pataz, Yalen, ca. 7°45'32.08"S 77°33'37.13"W, 2850 m, 26.iii.1988, D. Silva leg. (MUSM-ENT 500135); 1♀,
Trujillo, Victor Larco Herrera, Santiago de Huamán, ca. 8°08'18.98"S 79°02'23.28"W, 10 m, iii.1983, Sinuas leg.
(MUSM-ENT). Ucayali: 1♀, Tipishca, Tamaya, ca. 8°27'56.84"S 74°14'36.38"W, 149 m, 13–19.vi.1958, Koepcke
& Koepcke leg. (MUSM-ENT 500165); 1♂1♀, Ivita, near to Pucallpa, ca. 8°38'S 74°58'W, 196 m, 23–24.vii.1986,
D. Silva leg. (MUSM-ENT 500145); 1♀, ditto, 4.x.2002, R. Ramirez leg. (MUSM-ENT 500158). Huánuco: 1♀,
Rio Yuyapichis, Panguana Biological Station, 9º37'S 74º56'W, 236 m, 26.ii–24.xii.1983, M. Verhaghi leg. (IBSP
23680); 1♂, Huacaybamba, Road Huambu-Huanuco, near Hui Cuibambu, ca. 9°31'36.18"S 76°46'02.24"W,
3511 m, x.1984, R. Tejada leg. (MUSM-ENT); 1♂1♀, Tingo Maria, ca. 9°17'39.51"S 75°59'14.64"W, 705 m,
14–16.vii.1948 (MUSM-ENT). Pasco: 1♂, Oxapampa, ca. 10°34'30.09"S 75°24'00.04"W, 1817 m, 22.vi.1986,
D. Silva leg. (MUSM-ENT 500150). Lima: 1♀, San Vicente de Cañete, Valle de Mala, 13º5'S 76º24'W, 19 m,
18.ix.1997, M.A.B. Pino leg. (IBSP 14893); 4♂1♀, ditto (IBSP 14899); 1♀, ditto, 9.xi.1997 (IBSP 14895); 1♂,
ditto, 7.xi.1997 (IBSP 14906); 3♂, ditto (IBSP 14913); 1♂, ditto, 19.x.1997 (IBSP 14915); 2♂, ditto (IBSP 14919);
1♂2♀, ditto, 7.ix.1997 (IBSP 14921); 1♀, Chorillos, Los Pantanos de Villa, ca. 12°12'22.00"S 76°59'20.00"W, 2
m, 19.x.2003, W. Paredes leg. (MUSM-ENT 500142); 1♂, ditto, 20.viii.2006 (MUSM-ENT); 1♀, Road to Canta,
Km40, Trapiche, Recreo La Retama, 11°43'20.80"S 76°57'46.7"W, 546 m, 23.vi.2006, W. Paredes leg. (MUSM-
ENT); 1♀, Huacho, Rio Huacho, 11°04'39.16"S 77°36'03.33"W, 47 m, 21.x.2006, W. Paredes leg. (MUSM-ENT);
1♂, Hurin, La Tablada, Rio Lurin, ca. 12°12'27.83"S 76°52'12.27"W, 84 m, 12.vii.1988, G. Soza leg. (MUSM-ENT);

FIGURE 52. Prolycosides aussereri (Keyserling, 1877) comb. nov. Male pedipalp. (MUSM-ENT 500135) a–c (a, retrolateral;
b, ventral; c, prolateral); i, stridulatory organ; palp bulb d–h (d, retrolateral; e, ventral; f, prolateral; g, anterior; h, posterior)
Scale bars: a–i 200 µm. DMA depression of MA, E embolus, MA median apophysis, P palea, pp pars pendula, S scrapper,
ST subtegulum, STA subterminal apophysis, T tegulum, TA terminal apophysis, TDF tibial distal file, TF tegular furrow, TK
tegular keel, TL tegular lobe, VP ventral process.
FIGURE 53. Prolycosides aussereri (Keyserling, 1877) comb. nov. Male habitus (MUSM-ENT 500135) a, d, g–h (a, dorsal;
d, ventral; g, lateral; h, prosoma, eyes). Female epigynum b, e (b, ventral; e, dorsal). Female habitus c, f, i (c, dorsal; f, ventral;
i, lateral) Scale bars: a, c, d, f–i, 1 mm, b, e, 200 µm.
1♀, Cañete, Rio Cañete, ca. 13°07'29.65"S 76°21'44.05"W, 49 m, 15.ii.1996, S. Cordova & J. Duarez leg. (MUSM-
ENT 500152); 1♀, San Antonio de Chaclla, Jicamarca, ca. 11°54'23.68"S 76°42'12.53"W, 450 m, 1.ix.1968, A.
Hoempper leg. (MUSM-ENT 500137); 1♀, La Molina, 12°5'S 76°57'W, 236 m, 1.i.2006, A. Aparicio leg. (MUSM-
ENT 500126); 1♀, ditto, 15.i.2006, A. Aparicio leg. (MUSM-ENT 500127). Ica: 2♀, Huacachina, ca. 14°05'16.27"S
75°45'40.72"W, 401 m, 4.x.1985, F. Blancas leg. (MUSM-ENT). Cusco: 1♀, Santa Maria, Quillabamba, ca. 12°52'S
72°42'W, 1070 m, ix.2002, G. Binford et al. leg. (MUSM-ENT). Puno: 1♀, Puno, Pukará, ca. 15°02'30.51"S
70°21'48.11"W, 3867 m, 6.viii.2004, G. Binford et al. leg. (MUSM-ENT). Arequipa: 1♀, 7 km E. Cháparra, ca.
15°41'S 73°49'W, 1450 m, 15.ix.1988, R. Tejada leg. (MUSM-ENT 500134); 1♀, ditto, 1496 m, 19.ix.1988, R.
Tejada leg. (MUSM-ENT). BOLIVIA: Cochabamba: 1♀, Near Monte Puncu, ca. 17°35'13.82"S 65°18'23.36"W,
2700 m, 22.x.1983, A. Roig leg. (MACN); 1♀, ditto (MACN). BRASIL: Pernambuco: 1♂, Brejão, Fazenda
Santa Rita, Vila Santa Rita, ca. 7°33'19.85"S 35°0'5.91"W, v–x.2002, S.D. Bella leg., (IBSP 41085). Acre: 2♂1♀,
Rio Branco, Reserva Extrativista Catuaba, ca. 10°48'29.08"S 68°43'22.02"W, 262 m, 9.iv.1996, Equipe IBSP &
SMNK leg. (IBSP 15953). Amapá: 1♂2♀, Laranjal do Jari, Vila Cachoeira de Santo Antônio, ca. 0°49'54.59"S
52°30'30.83"W, 15 m, 18–24.ii.2003, J.A.P. Barreiros leg. (IBSP 53787); 1♂, Serra do Navio, 0°58'N 52°02'W,
110 m, 16–17.vi.1966, Galiano leg. (MACN); 1♂ ditto, (MACN). Amazonas: 1♀, Coari, Rio Urucu, Porto Urucu,
ca. 4°50'55.47"S 65°20'28.17"W, 61 m, 13.vii.2003, A.B. Bonaldo leg. (IBSP 53789); 1♀, Benjamin Constant, ca.
4°23'33.29"S 70°01'19.25"W, 79 m, ix.1962, K. Lenko leg. (MZUSP 20517). Pará: 1♀, Joanes, Ilha de Marajó,
ca. 0°52'42.65"S 48°30'33.22"W, 12 m, 7.x.2002, A.B. Bonaldo leg. (IBSP 53799); 2♀, Santarém, Fátima de
Urucurituba, ca. 2°26'28.78"S 54°48'46.64"W, 73 m, 24.i.1994, A.D. Brescovit leg. (MCN 25021). Bahia: 1♀,
Xique-Xique, ca. 10°49'16.44"S 42°43'01.57"W, 405 m, iii.1966 (IBSP 1984); 1♀, Iraquara, Chapada Diamantina,

ca. 12°14'44.11"S 41°37'25.99"W, 691 m, 15.xii.1998, L.S. Rocha leg. (IBSP 53876). Mato Grosso do Sul: 3♀,
Serra da Bodoquena, ca. 21°05'07.82"S 56°39'40.16"W, 616 m, 20–22.xi.2001, J. Raizer leg. (IBSP 33023). Rio de
Janeiro: 1♂1♀, Maricá, Praia de Bambuí, ca. 22°56'44.27"S 42°45'25.71"W, 2 m, 9.v.1977, C.J. Becker leg. (MCN
6179); 1♀, São João da Barra, Atafona, ca. 21°38'45.77"S 41°01'09.92"W, 4 m, 07.viii.1963, Exp. Dpto. Zool. leg.
(MZUSP 20481). São Paulo: 1♀, Barão de Itapetininga, ca. 23°35'28.72"S 48°02'44.27"W, 663 m, 2.vii.1964 (IBSP
1888); 1♀, São Paulo, 779 m, (IBSP 1948); 1♀, São Paulo, 23º32'51"S 46º38'10"W, 789 m, 17.vii.1969, D. Martins
leg. (IBSP 2283); 1♀, Araçoiaba da Serra, ca. 23°30'43.03"S 47°36'30.44"W, 625 m, ii.1987, R. Klink leg. (IBSP
4552); 1♀, Osasco, 23º31'19"S 46º48'07.77"W, 720 m, iii.1990, Equipe IBSP leg. (IBSP 5041); 1♂, Natividade da
Serra, ca. 23°22'45.81"S 45°26'37.54"W, 746 m, 23–26.vi.2001, M. Calleffo leg. (IBSP 29171); 1♀, Guarulhos, ca.
23°27'17.58"S 46°31'48.61"W, 767 m, 01.iv.2003, B. Imporato leg. (IBSP 37144); 1♂, São Paulo, ca. 23°32'38.07"S
46°38'30.61"W, 790 m, xi.1981, M.A. Klaussner leg. (IBSP 53807); 1♀, Itapevi, 23º32'51.80"S 46º56'24.55"W,
742m, 08.iv.1999, R. Sales leg. (IBSP 23650). 1♂1♀, Presidente Prudente, Rosana, Usina Hidrelétrica Porto
Primavera, ca. 22°06'47.06"S 51°27'53.42"W, 394 m, 11.xii.1998–i.1999, Eq. IBSP leg. (IBSP 53808); 1♀, ditto,
Eq. IBSP leg. (IBSP 53836); 2♀, Rosana, ca. 22°09'07.56"S 51°24'33.29"W, 424 m, 21.ii.2001, Usina Hidrelétrica
Sérgio Motta, R.P. Indicatti & D.F. Candiani leg. (IBSP 53817); 7♀, ditto, (IBSP 53818); 1♀, ditto (IBSP 53838);
4♂6♀, ditto, 11.xii.1998–i.1999 (IBSP 53846); 1♀, ditto, (IBSP 53850); 2♂2♀, Rosana, 22º34'47"S 53º03'33"W,
xii.1986, Equipe IBSP leg. (IBSP 4499). Rio Grande do Sul: 1♂, Rio Pardo, ca. 29°58'54.75"S 52°23'39.11"W,
17 m, 20.ii.1967, A.A. Lise leg. (IBSP 53791); 1♀, Herval, Santa Eliza, ca. 32°01'38.49"S 53°40'47.37"W, 101
m, 28.vii.1973, J. Sbruzzi leg. (MCN 3565); 1♀, Salto do Jacuí, Itaúba, ca. 29°04'58.58"S 53°13'12.95"W, 289 m,
11.iv.1978, A.A. Lise leg. (MCN 7933); 1♀, Santa Maria, Camobi, ca. 29°42'58.58"S 53°43'36.55"W, 112 m, i–
vi.1982, M. Ceolin leg. (MCN 14319); 1♀, Eldorado do Sul, ca. 30°00'15.96"S 51°18'10.79"W, 7 m, 30.ii.1996, G.S.
Carvalho & R. Silva Jr. Leg. (MCTP 12585); 1♂, ditto, 11–17.i.1997 (MCTP 12604); 9♀, Reserva Biológica de São
Donato, São Borja, ca. 29°02'49.09"S 56°03'05.09"W, 93 m, 23.vii.2012, M. Machado leg. (MCTP 36211– 36214);
1♀, ditto, 11.vii.2012 (MCTP 36226). PARAGUAY: Central: 1♀, Villeta, ca. 25°30'25.26"S 57°33'32.18"W, 92
m, 30.viii.1984, J. Kochalka leg. (IBNP); 1♀, ditto, (IBNP); 1♂, ditto, 24.vii.1982 (IBNP); 1♂, San Lorenzo, ca.
25°20'27.34"S 57°30'44.39"W, 118 m, 1–10.x.1985, J. Kochalka leg. (IBNP); 1♂ 2♀, Asunción, 14.ix.1984, J.
Kochalka leg. (IBNP); 1♀ ditto, 14.ix.1984 (IBNP); 1♀ ditto, 14.ix.1984 (IBNP). Boquerón: 4♂8♀, Estancia
La Gama, ca. 23°04'32.66"S 61°28'11.82"W, 191 m, 19–20.x.1990, J. Kochalka leg. (IBNP); 1♂, ditto, (IBNP).
Itapúa: 1♂, Isla Yacyretá, ca. 27°26'00.35"S 56°45'55.70"W, 73 m, 12–15.ix.1992, J. Kochalka leg. (IBNP); 1♀,
Near Arroyo Guapó, General Artigas, San Miguel Potrero, Coronel Bogado, ca. 27°10'14.89"S 56°13'03.97"W, 105
m, 29–30.ii.1996, J. Kochalka leg. (IBNP). 1♂, Isla Yacyretá, near Estrada Melgarejo, 27°25′48"S 56°46′12"W, 70
m, 12–15.ix.1992, J. Kochalka leg. (IBNP); 1♀, ditto, 25.iv.1993, D. Torres leg. (IBNP). Paraguarí: 3♂, Presidente
Hayes, 25 Leguas, ca. 22°47'50.74"S 59°36'34.56"W, 121 m, 11–12.vii.1983, J. Kochalka leg. (IBNP); 3♂, 25
Leguas (currently Teniente Primero Manuel Irala Fernández), 22°48′50.4″S 59°35′9.6″W, 120 m, 11–12.vii.1963,
J. Kochalka leg. (IBNP); 1♂, Ruta Trans-Chaco, km 20, ca. 25°10'41.50"S 57°33'07.91"W, 60 m, 25.viii.1981, J.
Kochalka leg. (IBNP). ARGENTINA: Salta: 1♀, Tartagal, ca. 22°31'00.34"S 63°48'20.19"W, ii.1963, Apostoli
leg. (MACN); 1♀, Rio Santa Maria, ca. 26°01'28.45"S 65°52'05.24"W, vii.1947, A.G. Giai leg. (MACN 2275);
1♀, Lorenzo, ca. 24°43'57.72"S 65°29'12.85"W, 6.vi.1927, Zotta leg. (MACN). Buenos Aires: 4♀, Berazategui,
ca. 34°45'45.50"S 58°12'37.49"W, 16.vi.1940, F. Monrós leg. (MACN). Corrientes: 1♀, Itati, ca. 27°16'22.07"S
58°14'33.23"W, 26.viii.1963, M.E. Galiano leg. (MACN). Santiago Del Estero: 1♀, Beltran, ca. 27°49'46.28"S
64°03'39.75"W, vi.1940, Birabén leg. (MLP). Tucuman: 1♀, Güemes, ca. 27°18'50.65"S 65°19'51.43"W, 26.x.1998,
(MACN). 4♀, Burruyacú, ca. 26°30'02.96"S 64°44'30.95"W, xi.1960, Birabén leg. (MACN); 1♀, ditto, xi.1960, M.
Birabén leg. (MACN). Entre Rios: 1♀, Rincón de Nogoyá, ca. 32°46'45.70"S 59°54'36.08"W, 27.ii.1982, A. Roig
leg. (MACN); 1♀, Parque Nacional El Palmar, 31°51'11"S 58°19'21"W, 18.iv.1981, P. Goloboff leg. (MACN).
Formosa: 1♀, Parque Nacional Pilcomayo, ca. 25°04'01.10"S 58°07'48.16"W, ix.1990, M.J. Ramírez leg. (MACN).
Chaco: 1♀, Resistencia, ca. 27°25'16.11"S 58°59'17.68"W, xii.1976, Aguillera leg. (MACN); 1♀, ditto, Denier leg.
(MACN 82).
Remarks: Prolycosides was erected by Mello-Leitão (1942b) from a single female based on the following
diagnostic characters: sparse spinulation on tibiae I and II, atypical compared to the Lycosa pattern, and five small
retromargin equal teeth lower border as well as the broad, smooth, and rectangular atrium on epigynum. Although
spinulation patterns are not always useful for distinguishing between genera, as demonstrated in other families (e.g.,
Hippa & Lehtinen 1983 [Lycosidae]; Ramírez 2003 [Anyphaenidae]; Miller et al. 2012 [Eresidae]) Mello-Leitão
succeeded in separating Prolycosides from other genera based on this evidence. After examination of the type
specimen and paired with males, we added diagnostic characters for the genus based on the males: embolus so long
that it bends at the tegular lobe and absence of basal apophysis at the median apophysis.
Different from Hogna, Prolycosides species have a straight prolateral arm of the median apophysis, which
is curved in Hogna, As mentioned, the tip of the embolus rests on the tegular lobe but is not wrapped by it as in
Pavocosa. In Prolycosides females the median septum is twice as long as the transversal septum (Fig. 53b), and
not thrice as it usually is in Pavocosa. By contrast the median septum in Hogna is quite the same length as the
transversal septum, with different degrees of dilatation on median septum (Dondale & Redner 1990). The atrium in
Prolycosides is smooth and wider (Fig. 55b) than in Hogna, and not narrow or striated as in Pavocosa (Fig. 55a).
Diagnosis. The males of Prolycosides have a very large embolus with a bent tip at the tegular lobe (Fig. 54e)
and lack a basal protuberance on the median apophysis (Fig. 54b). Females differ from other genera in having a
broad, rectangular atrium (Fig. 53b) and an ovoid spermatheca head (Fig. 53e).
FIGURE 54. Prolycosides aussereri (Keyserling, 1877) comb. nov. Male palpal bulb (MUSM-ENT 500135) a–c (a, ventral;
b, palear area; c, MA detail). Scale bars: a, 250 µm, b, 200µm, c, 100 µm. E embolus, MA median apophysis, P palea, pp
pars pendula, ST subtegulum, STA subterminal apophysis, T tegulum, TA terminal apophysis, TL tegular lobe, VP ventral
process.
Description. Male (MUSM-ENT 500135). Carapace brown, covered by short black bristles except at longitudinal
median band and submarginal bands, both yellow (Fig. 53a), broad at the front, tapering towards the back. Sparse
black bristles between the ocular area and the thoracic groove. Ocular area black, covered with white bristles,
cephalic portion of the carapace covered by sparse, large white bristles (Fig. 53h). Sternum longer than wide,
uniformly light brown, covered by short black bristles, which are sparse in the marginal area (Fig. 53d). First row
of eyes slightly procurved, shorter than the second row; AME slightly larger than the ALE; PME more than a radius
apart. Endites yellow, teardrop-shaped and convergent; labium square, darker than the endites; both endites and
labium yellowish in their distal portion. Chelicerae brown, frontally covered with sparse black setae and dense
white setae in all its extension; striae file on its frontal part; chillum membranous, divided with the mesal portion
sclerotized; condyle large and clearer than the chelicerae. Legs: All segments uniformly yellow except on tarsi and

metatarsi I and II, which are darker. Opisthosoma entirely covered by alternated tiny, black, and white bristles, the
background dark olive; cardiac mark dark, flanked by yellow longitudinal bands (Fig. 53a). Venter uniformly pale
yellow from epigastric furrow to spinnerets; anterior spinnerets wider than posterior spinnerets, both cylindrical
(Fig. 53d).
Pedipalp (Figs 52, 54), tibia cylindrical. Cymbium piriform (Fig, 52b), subtegulum lenticular (Fig. 52h). Tegulum
with a curved, C-shaped sperm duct. Terminal apophysis sclerotized, laminar, sickle-shaped, and distally sharp (Figs
52e, 54a), subterminal apophysis membranous, shorter than terminal apophysis (Figs 52g, 54b). Embolus thin, with
tip curved and embraced by the lamellar part of tegular lobe (Figs 52e, g, 54b). Median apophysis triangular and
with a sharp tip (Figs 52d, 54a); ventral projection (VP) short and slightly curved ventrally (Figs 52a, 54a).
Leg formula IV>I>II>III. Spination pattern: femur I p 0-0-2 d 1-1-1 r 1-1d-1d, II p 1d-1d d 1-1-1 r 1-1-1,
III-IV p 1d-1d d 1-1-1 r 1d-1d; patella I-IV p 1 d 1bristle-1bristle ap r 1; tibia I-II p d1-d1 d 1bristle-1 v 2-2-2ap
r 1-1d, III-IV p 1d-1d d r1-1 r 1-1 v 2-2-2ap; metatarsus I-II p 1d-1d-2 r 1d-1d-2 d 2bristle-2bristle v 2-2-1ap, III
p 1d-1d-2 d 1bistle-bristle-0 r 1d-1-2 v 2-2-1ap, IV p 1d-1d-2 d 1bistle-1bristle-0 r 1d-1d-2 v 2-1-2-1ap.
PME 0.60, PLE 0.45, interdistances: AME-AME 0.17; AME-ALE 0.06; PME-PME 0.40; PLE-PLE 1.21. POQ long
0.95, POQ posterior width 1.54, POQ anterior width 0.98. Chelicerae: length 2.55. Opisthosoma: length 6.78, width
4.66. Legs: femur + patella + tibia + metatarsus + tarsus = total length: pedipalp 2.55 + 1.22 + 1.39 + - + 1.83 =
6.99, I 5.94 + 2.78 + 5.22 + 5.44 + 3.00 = 22.38, II 6.17 + 2.61 + 4.78 + 5.00 + 2.89 = 21.45, III 5.00 + 2.22 + 4.22
+ 5.67 + 2.78 = 19.89, IV 6.33 + 2.56 + 5.44 + 7.78 + 3.33 = 25.44.
Female (MUSM-ENT 500135). General pattern as in male. First row of eyes straight, shorter than the second
row; AME slightly larger than the ALE; PME separated by a length same as their radii. Chelicerae with three
promarginal teeth, the median the largest; retromarginal with three teeth, the basal the shortest. Chelicerae is darker
brown than in males; chillum as in males; boss larger compared with males. Sternum, endites and labium yellow
(Fig. 53f). Legs and opisthosoma as in males (Fig. 53i).
Genitalia (Figs 53b, e, 55), LS and TS forming a T-shaped plate (Fig. 53b). LS wider than TS, atrium smooth
and wide (Fig. 55a). CO at the border of TS (Fig. 55c). Hoods parallel and triangular (Figs 53b, 55a). HS kidney-
shaped, SS narrow; BS bulbous, VC absent (Figs 53e).
FIGURE 55. Prolycosides aussereri (Keyserling, 1877) comb. nov. Female, epigynum (MUSM-ENT 500135) a–c (a, ventral;
b, atrium detail; c, copulatory opening). Scale bars: a, 250 µm, b, 200 µm, c, 100 µm. AT atrium, H hood, CO copulatory
opening, MS median septum, TS transversal septum.
Leg formula IV>I>II>III. Spination pattern: femur I p 0-0-2 d 1-1-1, II p 0-d1-d1 d 1-1-1 r 0-1d-0, III p 0-
1d-1d d 1-1-1 r 1d-1d-1d, IV p 0-0-1d d 1-1-1 r 1d-1d; patella I-II d 0-1bristle, III-IV p 1 d 1bristle-1bristle r
1; tibia I p 0-1-0 d 1bristle v 2-2-2ap, II p 1d-1 d 1bristle-1bristle v 2-2-2ap, III-IV p 1-1v d 1-1 r 1-1 v 2-2-2ap;
metatarsus I p 0-0-1 d 1bristle-1bristle v 2-2-1 r 0-0-d1, II p 0-0-2 d 2bristle-1bristle r 0-0-1 v 2-2-1ap, III-IV p
1d-1d-2 r 1d-1d-2 v 2-2-1ap.
4.78. Legs: femur + patella + tibia + metatarsus + tarsus = total length: pedipalp 2.56 + 1.22 + 1.56 + - + 1.78 =
7.12, I 5.00 + 2.78 + 4.00 + 3.78 + 2.11 = 17.67, II 4.89 + 2.33 + 3.78 + 3.78 + 2.22 = 17.09, III 4.33 + 2.11 + 3.11
+ 3.72 + 2.11 = 15.38, IV 5.67 + 2.44 + 4.61 + 6.22 + 2.67 = 21.61.
6.68±0.69; CW 4.22 – 6.11, 5.02±0.55; (TL 11.67 – 13.33, 12.59±0.69; CL 6.11 – 8.89, 7.26±1.18; CW 4.78 – 6.45,
5.41±0.74).
Distribution. Cuba, Venezuela, Colombia, Peru, Bolivia, Brazil, Paraguay, and Argentina (Fig. 67)
FIGURE 56. Tarentula aussereri Keyserling, 1877. Female holotype, a. ventral opisthosoma, b. epigynum ventral view.
Collecting and ID labels c–d (c, type locality label; d, ID label).
Trochosa fusca (Keyserling, 1877) comb. nov.

Figs 57–58, 65
Tarentula fusca Keyserling, 1877: 640, plate 7, fig. 22 (female holotype from Cuba, no additional information, BMNH 1636413,
not examined).
Lycosa fusca: Banks 1901: 223; Petrunkevitch 1929: 91, figs 76–79; Bryant 1940: 280, pl. 2, figs 23–24.
Arkalosula fusca: Roewer 1955: 232.
Arctosa fusca: World Spider Catalog 2023

Remarks. Trochosa fusca female was first described by Keyserling (1877) and the male was subsequently described
by Bryant (1940), both based on specimens from Cuba. The “pair of small dark spots in stripe” mentioned by Bryant
(1940) for both the male and female is a typical character in Trochosa. On the other hand, a dense scopula on tarsi,
metatarsi, and distal half of tibiae of legs I–II is not typical in Arctosa. Bryant (1940) described the epigynal plate
of this species as a “cross piece”, wider than long with a “circular depression each side of the septum” (Keyserling
1877: fig 22; Bryant 1940: fig 23). The epigynal atrium of Arctosa females lack this depression, which is not
compatible with the diagnostic Arctosa trapezoidal epigynum plate (Dondale & Redner 1990: figs 439, 446, 457).
In addition, the anterior pockets (hoods) of T. fusca are not as deep as in Birabenia but shallow like in Trochosa. In
Bryant's drawings, the ventral process of the median apophysis is minute, corresponding to one of the diagnostic
characters of Trochosa (Bryant 1940: fig 24). Furthermore, the terminal apophysis is larger and flattened in Trochosa
and not slender as in Birabenia. Finally, Birabenia species have diagnostic four promarginal teeth, T. fusca have
only three.
Material examined: CUBA: Santiago de Cuba: 1♀, Jardin Botánico, ca. 22°59'29.51"N 82°20'13.38"W,
17.vi.1998, A. Sánchez leg. (IBSP 169626); 1♂, ditto, 6.ii.1998, A. Sánchez-Ruiz leg. (IBSP 233748); 1♂, Guamá,
Pico Cuba, ca. 19°58'43.28"N 76°31'32.52"W, 27.ix.1999, A. Sánchez leg. (IBSP 169618).
Diagnosis: The ventral projection of median apophysis of T. fusca is as short as in T. humicola, but the median
apophysis is clearly shorter (Figs 57b, h) and the former has only two macrosetae on tip of cymbium (Fig. 57b)
instead of three as in T. humicola. Females are recognized by the vulval chamber larger than the head of spermathecae
(Fig. 58e).
Description. Male (IBSP 169618). Carapace brown, covered by short black bristles except on the white
longitudinal median and submarginal bands, (Fig. 58a), longitudinal median band yellow, broad at the front and
tapering towards the back. Ocular covered by sparse black bristles. Sternum longer than wide, uniform brown in
color, covered by sparse, short, black bristles (Fig. 58d); endites yellow, teardrop-shaped, and convergent; labium
square, darker than the endites; both endites and labium yellowish distally. First row of eyes slightly procurved,
shorter than the second row; AME slightly larger than the ALE; PME separated by less than their radii (Fig. 57g).
Chelicerae with striated, brown with SF, covered frontally with sparse black and white setae in the frontal part;
chillum membranous, divided with the mesial portion sclerotized; visible boss clearer than the chelicerae. Legs:
all segments uniformly dully brown, except tarsi and metatarsi I and II, which are darker. Dorsum of opisthosoma
dully olive, covered by alternated black bristles; cardiac mark dark brown flanked by yellow longitudinal bands
(Fig. 58a). Ventrally uniformly pale yellow from epigastric furrow to spinnerets, speckled with scattered black dots;
anterior spinnerets wider than posterior spinnerets, both cylindrical (Fig. 58d).
Pedipalp (Figs 57a–f, h, i). Tibia cylindrical. Cymbium piriform, subtegulum lenticular in ventral view (Fig.
57e). Tegulum traversed by a C-shaped, curved sperm duct. Ventral projection of median apophysis short, triangular,
with sharp tip in retrolateral view (Figs 57e, f, h); median apophysis short, slightly curved ventrally (Fig. 57b).
Terminal apophysis laminar, sclerotized, sickle-shaped (Figs 57c, e, h). Embolus thin, with straight tip lying in the
dorsal channel of median apophysis (Figs 57c, e, f).
Leg formula IV>III>II>I. Spination pattern: femur I p 0-1-2 d 1-1-1 r 2-d1-d1, II p 0-1-2 d 1-1-1 r d1-d1-d1,
III-IV p d1-d1-d1 d 1-1-1 r d1-d1-d1; patella I-IV p 1 d 0-1bristle ap r 1; tibia I-II p 1d-1 d 0-1 v 2-2-2ap r 0-1-1,
III-IV p 1-1 d 1-1 r 1-1 v 2-2-2ap; metatarsus I-II p d1-1-2ap r d1-d1-2 d 1bristle-21bristle v 2-2-1ap, III p 1d-1d-2
d 1bistle-1bristle-1bristle r d1-d1-2 v 1-2-2-1ap, IV p d1-d1-2 d 1bistle-1bristle-1bristle r 1v-1d-2 v 2-2-1ap.
Measurements: TL 11.67, CL 6.78, CCW 3.44, CTW 5.00, CCH 2.55, CTH 2.77. Eyes: AME 0.36, ALE 0.24
posterior width 1.79, POQ anterior width 1.07. Chelicerae: length 2.74. Legs: length of segments femur + patella +
tibia + metatarsus + tarsus = total length: pedipalp 2.61 + 1.17 +1.33 + - + 2.00 = 7.11, I 6.00 + 2.56 + 5.76 + 6.44
+ 2.89 = 23.65, II 6.00 + 2.78 + 5.67 + 6.33 + 2.88 = 23.66, III 5.33 + 2.11 + 4.44 + 5.78 + 2.67 = 21.33, IV 6.56
+ 2.44 + 6.11 + 7.89 + 3.11 = 26.11.
Female (IBSP 169626). General pattern as in male. First row of eyes slightly procurved, shorter than the second
row; AME larger than the ALE; PME separated by less than their radii. Chelicerae with three promarginal teeth, the
median the largest; retromarginal with three teeth, the basal the shorter. Chelicerae is darker brown and more robust
than in males; chillum and boss larger compared with those of the male. Sternum, endites, and labium darker than
in males (Fig. 58f). Legs as in males except by a uniform color in each segment; venter of opisthosoma dark brown
uniform (Fig. 58i).
FIGURE 57. Trochosa fusca (Keyserling, 1877) comb. nov. (IBSP 169618) Male pedipalp a–c (a, prolateral; b, ventral; c,
retrolateral) Male palpal bulb d–f, h (d, prolateral; e, ventral; f, retrolateral; h, anterior), g, male chelicerae, i, stridulatory
organ. Scale bars: a–c, 500 µm, d–f, h, i, 200 µm, g, 2 mm. CHI chilum, CMS cymbium macrosetae, E embolus, MA median
apophysis, P palea, pp pars pendula, T tegulum, TA terminal apophysis, TDF Tibial distal file, TF tegular furrow, TK tegular
keel, TL tegular lobe, S scraper, SF striae file, ST subtegulum, VP ventral process.

FIGURE 58. Trochosa fusca (Keyserling, 1877) comb. nov. Male habitus (IBSP 169618) a, d, g (a, dorsal; d, ventral; g, lateral)
Epigynum b, e (b, ventral; e, dorsal) Female habitus (IBSP 169626) c, f, h–i (c, dorsal; f, ventral; h, frontal; i, lateral) Scale bars:
a, g, 2 mm d, 5 mm, b, e, 200 µm, c, f, i, 5 mm, h, 2 mm. CO copulatory opening, H hood, MS median septum, TS transversal
Genitalia (Figs 58b, e), LS and TS forming a T shaped plate (Fig. 58b). The TS height is greater than the width
of LS; a short and smooth atrium (Fig. 58b). CO at the border of TS (Fig. 58b). Internal genitalia with HS triangular,
not wider than SS, BS short, with VC larger than HS (Figs 58e), hoods convergent, triangular (Figs 58b, 58e).
Leg formula IV>II>I>III. Spination pattern: femur I p 0-0-2 d 1-1-1 r d1-d1-d1, II p 0-d1-d1 d 1-1-1 r d1-d1-
d1, III p 0-d1-d1 d 1-1-1 r 0-d1-d1-d1, IV p d1-0-d1 d 1-1-1 r d1-0-d1; patella I-IV d 0-1bristle p 1 r 1; tibia I p
d1-1 d 1bristle-0 v 2-2-2ap r 0-1, II p 1-1 d 1bristle-0 v 2-2-2ap, III-IV p 1-1 d 1-1 r 1-1 v 2-2-2ap; metatarsus I
p 0-0-1 d 1bristle-1bristle v 2-2-1 r 0-0-d1, II p 0-0-2 d 2bristle-1bristle r 0-0-1 v 2-2-1ap, III p d1-d1-2 r d1-v1-
d1-2ap v 2-1-1ap, IV p d1-d1-2 r d1-d1-v1-2ap v 2-1-1ap.
posterior width 1.48, POQ anterior width 0.95. Chelicerae: length 2.55. Legs: length of segments femur + patella +
tibia + metatarsus + tarsus = total length: pedipalp 1.83 + 1.00 + 1.11 + - + 1.33 = 5.27, I 4.11 + 2.22 + 3.44 + 3.33
+ 1.89 = 14.99, II 3.88 + 1.89 + 2.89 + 3.22 + 1.78 = 13.66, III 3.33 + 1.44 + 2.67 + 3.33 + 1.67 = 12.44, IV 4.56
+ 1.89 + 4.00 + 4.89 + 2.22 = 17.56.
Distribution. Caribbean (Cuba and Puerto Rico, Fig. 65), Puerto Rico records were taken from Petrunkevitch
(1929).
Trochosa humicola Bertkau, 1880 comb. rest.
Figs 59‒61, 66
Trochosa humicola Bertkau, 1880: 65, pl. 1, figs 20, 20a (four female syntypes from Rio de Janeiro, Rio de Janeiro, Brazil, lost,
see remarks). Bonnet 1959: 4703.
Lycosa humicola: Mello-Leitão 1947: 262.
Lycosa inconspicua Bryant, 1948: 355, pl. 2, fig. 16 (Female holotype from Ennery, Haiti, 7.ix.1934, Darlington leg., MCZ
21626, examined by photos, Fig. 61) syn. nov.
Arkalosula inconspicua: Roewer 1955: 232.
Triccosta humicola: Roewer 1955: 298.
Tricca humicola: Braun 1963: 81.
Arctosa humicola: World Spider Catalog 2023
Other material examined. BRAZIL: Rio Grande do Norte: 1♂, Atol das Rocas, Parque Nacional do Atol das
Rocas, ca. 3°52'20.88"S 33°48'40.22"W, -1 m, 4.ix.2005, C.E. Almeida leg. (IBSP 12055). Pernambuco: 3♂2♀,
Fernando de Noronha Archipelago, Fernando de Noronha Island, 3º50'S 32º15'W, 16 m, 9–19.x.2005 (IBSP
70343); 3♂2♀, ditto, (IBSP 70345); 1♂4♀, ditto, (IBSP 70344); 1♂4♀, ditto, (IBSP 70346); 1♂4♀, ditto, (IBSP
70347); 1♂4♀, ditto, (IBSP70348); 1♂1♀, ditto, 16 m, 17.iv.2006 (UFPE), all collected manually in houses by
G.C.C. Freitas. Sergipe: 1♀, Aracaju, Atalaia Velha, ca. 10°58'43.32"S 37°2'12.61"W, 1 m, 23.viii.1978, I. Pinho
leg. (IBSP 7648); 1♂, Santa Luzia do Itanhy, Crasto, ca. 11°21'14.76"S 37°27'30.67"W, 55 m, 12–14.xi.1996,
A.D. Brescovit leg. (IBSP 10160); 1♂, São Cristóvão, Campus UFSE, ca. 10°55'35.21"S 37°6'9.19"W, 9 m,
25.ix.1979 (IBSP 10396), 1♂, ditto, 25.iv.1989 (IBSP 10408). Bahia: 1♂ Central, ca. 11°7'54.13"S 42°7'2.39"W,
648 m, 12.vii.1997, E.F. Ramos leg. (IBSP 13393); 1♂1♀, Riachão do Jacuípe, Rio Jacuípe, ca. 11°48'25.92"S
39°23'5.01"W, 221 m, 1980, S. Lucas leg. (IBSP 4607); 1♀, Feira de Santana, ca. 12°11'53.51"S 38°57'52.59"W,
240 m, 1.x.1991, V. dos Santos leg. (IBSP 7120). Minas Gerais: 1♀, Cambuí, ca. 22°36'45.60"S 46°3'8.66"W, 895
m, viii.1995, L.C. Gregorio leg. (IBSP 14170); 4♂3♀, Belo Horizonte, Estação Ecológica da UFMG, 19°52'38"S
43°58'16"W, 845 m, v.1999, E.S.S. Álvares leg. (UFMG 75); 1♀, Belo Horizonte, Campus UFMG, ca. 19°51'41"S
43°57'48"W, 845 m, 6.iv.2000, E.S.S. Álvares et al. leg. (UFMG 195). Rio de Janeiro: 1♀, Rio de Janeiro, (MNRJ).
4♀, ditto (MNRJ 1918). São Paulo: 1♂, Campinas, ca. 22º54'11.15"S 47º1'25.63"W, 699 m, x.1978 (IBSP 4217);
2♀, Tietê, ca. 23°5'52.81"S 47°42'47.41"W, 482 m, 15.v.2002 (IBSP 34996); 1♀, Sorocaba, 23º30'6"S 47º27'29"W,
599 m, viii.1983, M. Ianelle leg. (IBSP 3744); 1♀, São Paulo, 23º33'S 46º38'W, 756 m, 9.xii.1967, A.A. Miranda leg.
(IBSP 2066); 1♀, ditto, 7.vii.1969, N.S. Junior leg. (IBSP 2284); 1♀, ditto, 14.vii.1969, G.R. Sobrinho leg. (IBSP
8427); 1♀, ditto, 3.ix.1969 (IBSP 2357); 10♀, 1.x.1973, ditto, G.C. Netto leg. (IBSP 2711); 3♀, vi.1988, ditto, R.G.
Silva leg. (IBSP 5021); 1♀, iii.1991, ditto, V.J. Topein leg. (IBSP 41790); 1♂, ditto, 3.iv.1995, L.S. Belfi leg. (IBSP
6182); 1♂, xi.1997, ditto, L.S. Rocha leg. (IBSP 13937); 1♀, ditto, 13.viii.2003, L. de França leg. (IBSP 40611);
1♀, Bairro Freguesia do Ó, ca. 23º30'4.99"S 46º41'51.27"W, 770 m, 4.iv.1966, A.M. Baptista leg. (IBSP 1987); 1♀,
Bairro Pinheiros, ca. 23º33'46.49"S 46º41'29.28"W, 747 m, 9.iv.1968, J.R. Cerrato leg. (IBSP 2115); 1♂3♀, Bairro
Butantã, ca. 23º33'56.59"S 46º43'16.48"W, 788 m, iii.1983, W.M. Sportoni leg. (IBSP 3651); 1♀, Campus IBSP,
ca. 23º33'57.83"S 46º43'49.08"W, 786 m, iii.1983, W.M. Sportoni leg. (IBSP 3717); 1♀, Osasco, ca. 23º32'16.93"S
46º47'30.21"W, 765 m, 30.iv.1996, M. Padilha leg. (IBSP 14350); 1♀, Itapevi, 23º32'56"S 46º56'3"W, 768 m,
8.xi.2002, F. Peit leg. (IBSP 37318), 1♀, ditto, 5.xii.2002, W. Valim leg. (IBSP 37323); 1♀, Dois Córregos, ca.
22º22'29.97"S 48º23'19.90"W, 702 m, 1–6.iv.2002, G.Q. Romero leg. (IBSP 34964); 1♀, Louveira, ca. 23º5'31.03"S
46º56'58.74"W, 700 m, 11.x.1995, D. Tremori leg. (IBSP 6413). Rio Grande do Sul: 1♂, Barros Cassal, em riacho,
29°37'22.7"S 52°30'15.6"W, 85 m, 5.viii.2015, V.S.R. Diniz & L.E.C. Schmidt leg. (UFMG 20948).
Remarks. As originally described by Bertkau, the average size of Trochosa humicola is larger than other Arctosa
species (13 mm in T. humicola versus average 7 mm in neotropical Arctosa). We examined non-type specimens
from Rio de Janeiro as well as localities and states bordering the state of Rio de Janeiro, as well as from central
and northeastern regions of Brazil. Although the holotype is lost as mentioned in Levi (1964): "...The whereabouts
of the types and other specimens of this collection are not known. They may have been lost..." referring to the
specimens collected by the Belgian professor Van Beneden in 1873 and given to Bertkau for further examination,
we conclude that the specimens examined correspond to T. humicola. The prosoma pattern described by Bertkau
(1880) correspond to the Trochosa pattern (Brady 1979), as well as the typical inverted T-shaped epigynum. The
specimens examined by us have epigynal hoods curved internally, transversal septum (TS) height equal when

FIGURE 59. Trochosa humicola Bertkau, 1880 comb. nov. (UFMG 75) Male pedipalp a–c (a, prolateral; b, ventral; c,
retrolateral) Male palpal bulb d–f, h (d, prolateral; e, ventral; f, retrolateral; h, anterior), g, male chelicerae, i, stridulatory organ.
Scale bars: a–h, 200 µm, i, 100 µm. CHI chilum, CMS cymbial macrosetae, E embolus, MA median apophysis, P palea, pp
pars pendula, T tegulum, TA terminal apophysis, TDF Tibial distal file, TF tegular furrow, TK tegular keel, TL tegular lobe, S
scraper, SF striae file, ST subtegulum, VP ventral process.
FIGURE 60. Trochosa humicola Bertkau, 1880 comb. nov. Male habitus (UGMF 75) a, d, g (a, dorsal; d, ventral; g, lateral)
Epigynum b, e (b, ventral; e, dorsal) Female habitus c, f, h–i (c, dorsal; f, ventral; h, frontal; i, lateral) Scale bars: a, d, g, 1 mm,
b, 100 µm, c, f, h–i, 1 mm, e, 200 µm. H hood, MS median septum, TS transversal septum.
FIGURE 61. Lycosa inconspicua Bryant, 1948. Female habitus, holotype a–b, d–f (a, prosoma, dorsal; b, opisthosoma, dorsal;
d, prosoma, ventral; e, opisthosoma, ventral; f, prosoma frontal) Epigynum, c, ventral Scale bars: a, b, d–f, 1 mm, c, 0.25 mm.
CM cardiac mark, CO copulatory opening, PS Parallel stripes.

compared with median septum (MS) width. Arctosa epigynum is commonly a trapezoidal piece with MS and TS
undistinguishable among them. The presence of a rounded, visible palea in males also excludes the specimens we
examined from Arctosa, whose species possess a reduced palea. The size and shape of the ventral projection of the
median apophysis in T. humicola is triangular and short, different from a pronounced and acute one seen in Arctosa
species.
Bryant (1948) records this species from the island of Hispaniola, which nowadays comprises Haiti and
Dominican Republic. The specimens described in Bryant (1948) were collected in the eastern side of the island, in
Haiti. We did not found any published record from Guyana, as reported in the World Spider Catalog (2023).
Diagnosis. Males of Trochosa humicola can be recognized by the short, triangular ventral projection and the
folded tip of the median apophysis (Fig. 59c). Females are recognized by curved and deep hoods, which are internally
convergent, and the head of spermatheca narrower than its stalk (Fig. 60e).
FIGURE 62. Geographic distribution records of Arctosa ayaymama sp. nov. (blue dot), Arctosa pacaya sp. nov. (red dot),
Hogna andina comb. nov. (orange dot).
Description. Male (UFMG 75). Carapace brown, covered by short white and black bristles, except on the
longitudinal median band, which is broad, yellow, and tapering towards the back. Sparse black bristles between
the ocular area and the thoracic groove; ocular area black covered with white bristles, cephalic portion of the
prosoma covered by sparse, large white bristles. Sternum longer than wide, uniformly light brown, covered by
short black bristles, sparsely covered in its entire surface (Fig. 60d). First row of eyes straight, shorter than the
second row; AME slightly larger than the ALE; PME separated by a length same as their radii. Chelicerae with three
promarginal teeth, the median the largest; retromargin with three teeth, the basal the shorter. Chelicerae creamy,
covered frontally with sparse black setae in the frontal part and white setae in distal borders; chillum membranous,
divided with the mesal portion sclerotized; boss clearer than the chelicerae. Frontal protuberance on both cheliceral
teeth (Fig. 59g). Endites dully brown and convergent; labium square-shaped, darker than the endites; both endites
and labium yellowish in their distal portion. Legs: All segments dully brown except tarsi and metatarsi, which are
darker. Opisthosoma all covered by tiny white bristles, the background olive, with dispersed black bristles mainly
on dorsal part; cardiac mark indistinguishable; with a group of strong, curved black bristles on dorsum, near the
prosoma. Venter with a longitudinal, yellow tapered band, which is sparsely covered by black bristles and has two
median, pale brown lines from epigastric furrow to spinnerets; anterior spinnerets the same size as the posteriors,
both cylindrical (Fig. 60d).
Pedipalp (Figs 59a–f, h, i), tibia cylindrical. Cymbium fusiform. Subtegulum lenticular and visible in ventral
view. Tegulum with a curved, L-shaped sperm duct (Figs 59b, e). Median apophysis laminar, triangular, and with
unsharpened tip; ventral process short and slightly curved ventrally (Fig. 59d, h). Terminal apophysis laminar,
distally sharp (Figs 59e, h). Embolus thin and curved, not twisted at the tip; pars pendula present (Fig. 59e).
Leg formula IV>I>III>II. Spination pattern: femur I p 0-0-1 d 1-1-1 r 1-1, II p 1-1 d 1-1-1 r 1-1-0-1, III p 1-1
d 1-1-1 r 0-1-1, IV p 1-0-1 d 1-1-1 r 0-0-1; patella I-IV p 1 r 1; tibia I-II p 1-1 v 2-2-2ap r 1-1, III-IV p 1-1 d 1-1
r 1-1 v 2-2-2ap; metatarsus I-II p 0-1-1 r 0-1-1 v 2-2-2ap, III-IV p 1d-1d-1d r 1d-1d-1d v 2-2-2ap.
2.61. Legs: (femur + patella + tibia + metatarsus + tarsus = total length): pedipalp 1.90 + 0.80 + 1.15 + - + 1.38 =
5.23, I 3.56 + 1.43 + 3.21 + 3.02 + 2.07 = 13.29, II 3.00 + 1.33 + 2.86 + 2.86 + 1.98 = 12.02, III 2.89 + 1.33 + 2.71
+ 3.31 + 1.98 = 12.22, IV 4.11 + 1.48 + 3.86 + 5.02 + 1.98 = 16.44.
FIGURE 63. Geographic distribution records of Arctosa conflicta sp. nov. (red triangles)

FIGURE 64. Geographic distribution records of Arctosa costenola sp. nov. (orange dot), Arctosa jibarosa sp. nov. (white dot),
Arctosa tenebrosa comb. nov. (yellow diamond), Arctosa villa sp. nov. (blue diamond).
FIGURE 65. Geographic distribution records of Arctosa denticulata (blue asterisc), Arctosa littoralis (yellow square), Arctosa
serii (light blue star), Trochosa fusca comb. nov. (green dot).
FIGURE 66. Geographic distribution records of Arctosa mineira sp. nov. (rose dot), Arctosa sapiranga (black dot), Trochosa
humicola comb. nov. (cyan diamond).
Female (UFMG 75). General pattern as in male. First row of eyes straight, shorter than the second row; AME
slightly larger than the ALE; PME separated by their radii (Fig. 60h). Chelicerae with three promarginal teeth, the
median the largest; retromargin with three teeth, the basal the shorter. Chelicerae creamy, covered frontally with
sparse black setae in the frontal part and white setae in distal borders; chillum membranous, divided with the mesial
portion sclerotized; boss clearer than the chelicerae. Sternum endites and labium as in males but slightly darker (Fig.
60f). Legs and opisthosoma as in males (Fig. 60i).
Genitalia (Figs 60b, e), median septum and transversal septum forming a T shape plate. in ventral view, hoods
triangular, atrium smooth and wide (Fig. 60b). Copulatory openings at the anterior border of transversal septum
(Fig. 60e). Internally, head of spermatheca as narrow as the stalk; base of spermatheca bulbous, vulval chamber
absent, hoods curved and convergent (Fig. 60e).
Leg formula IV>I>II>III. Spination pattern: femur I p 0-0-2 d 1-1-1 r 0-1-1, II p 0-1d-1d d 1-1-1 r 0-d1-1d,
III p 0-1d-1d d 1-1-1 r 0-1d-1d, IV p 1d-1d-1d d 1-1-1 r 0-0-d1; patella I d 0-1bristle; II p 1 d 0-1bristle, III-IV
p 1 d 1-1 r 1; tibia I-II p 1-1 d 1bristle-1bristle v 2-2-2ap, III-IV p 1d-1d d r1-1 r 1d-1d v 2-2-2ap; metatarsus I p
0-1 d 1bristle-1bristle v 2-2-1 r 0-1d, II p 1-1-2 d 1bristle-1bristle r 0-0-1 v 2-2-1ap, III-IV p 1d-1d-2 r 1d-1d-2 v
2-2-1ap.

5.00. Legs: (femur + patella + tibia + metatarsus + tarsus = total length): pedipalp 2.22 + 1.23 + 1.44 + - + 1.77 =
4.44, I 4.44 + 2.22 + 4.32 + 3.33 + 2.22 = 16.53, II 4.33 + 2.44 + 3.44 + 3.22 + 2.11 = 15.54, III 4.05 + 2.34 + 3.22
+ 3.78 + 1.89 = 15.28, IV 5.33 + 2.33 + 4.72 + 6.16 + 2.33 = 20.87.
Distribution. Brazil (northeast, central and southern regions) and Haiti (Fig. 66)
FIGURE 67. Geographic distribution records of Hogna pugil comb. nov. (red cross) updated records taken from Bertkau
(1880), Prolycosides aussereri comb. nov. (wine dot).
Discussion
The five new Arctosa species described (A. ayaymama, A. conflicta, A. costenola, A. pacaya and A. villa) here came
from humid areas associated to damp and/or flooded habitats, as already reported for other species of the genus
(Buchholz & Schröder 2013; Esquivel-Bobadilla et al. 2013; Relys et al. 2009; Schmidt et al. 2008; Silva & Lise
2009; Yoo et al. 2007). Nocturnal and burrowing behavior has also been reported for some species of Arctosa (e.g.
Arctosa lutetiana (Simon, 1876); Bizuet-Flores et al. 2015; Dolejš et al. 2008; Logunov 2011). Although it was
not recorded in herein, we have no conclusive evidence of structures responsible for building shelters as in some
Neotropical lycosids in genera as Lycosa, Trochosa, Allocosa and Pavocosa (Mendoza Belmontes et al. 2018).
Therefore, the first pair of legs, the robust chelicerae, particularly in females (Zyuzin, 1990), and the cymbium in
males could be used for digging or copulatory behavior.
Epigynal hoods females of Arctosa are variably developed. This may have led Dondale & Redner (1983a,
1990) to consider its absence as a diagnostic feature of the genus. For instance, the hood was not well documented
or illustrated in Arctosa cinerea (Fabricius, 1777), but it is evident, though minute (Buchar et al., 2006: fig. 25;
Wang et al., 2012: fig. 12E). However, as observed in some Neotropical species of Arctosa, both the absence (A.
tenebrosa, A. conflicta, A. mineira, A. tenella and A. littoralis) or poorly visible condition (A. sapiranga and A. serii)
are present. Similarly, some Holarctic species, as Arctosa raptor (Kulczyński, 1885) and A. emertoni Gertsch, 1934;
and Asian species as Arctosa stigmosa (Thorell, 1875) and A. subamylacea (Bösenberg & Strand, 1906) have no
epigynal hoods. Likewise, there are also species from both regions that do possess it (though reduced), as Arctosa
cinerea (Fabricius, 1777), A. perita (Latreille, 1799), and A. ravida Ponomarev, 2007. Therefore, the absence of
hood should not be considered as a diagnostic character of the genus.
FIGURE 68. Geographic distribution records of Arctosa tenella comb. nov. (rose square).
Molecular phylogenetic analyses indicated that Palearctic species of Arctosa are within Tricassinae, along
with Tricassa (T. deserticola Simon, 1910 and T. madagascariensis Jocqué & Alderweireldt, 2001), while Arctosa
sapiranga emerge within Allocosinae (Gonnet et al. 2021; Laborda et al. 2022; Piacentini & Ramírez 2019).
Furthermore, Tricassinae is characterized by grouping males with large median apophysis positioned longitudinally
on the palpus, and females with a simple epigynum with papillose surface on the scape, sinuous copulatory ducts,
and both sexes with large anterior spinnerets. This combination of characters is assumed to be diagnostic characters
for Tricassinae (Alderweireldt & Jocqué 1993; Jocqué & Alderweireldt 2001) and seems doubtful because none of
the above mentioned characters would serve to include Arctosa within this subfamily. Allocosinae on the other hand,
groups species whose males have transversely arranged, bifid median apophysis and females with simple median
septum without papillae on its surface. That is why it is difficult to place Arctosa in one of the two subfamilies
because it possesses characters of both, although it has more diagnostic elements of Tricassinae.

Acknowledgements
We would like to thank Adalberto Santos by the revision of previous versions of the manuscript and all Colombian
curators, especially Eduardo Florez, who allowed the first author to revise material of Instituto de Ciencias Naturales
(ICN-Ar) in Bogotá, as well as gave to WPM the accommodation in his cozy house. To Jimmy Cabra at MUSENUV,
Navas-Suarez at CUDC, A.L. Garcia-Hernandez at CIUQ, G. Castillo at CEUN-PSO, D. Forero and M.A. Rodriguez-
Morales at MPUJ. To the family of David Vergara Moreno, thank you immensely for hosting WPM in Cartagena.
We are also grateful to Taryn Ghia at QCAZ and to Delis Fiallos Gordon for providing a comfortable stay in Quito
to WPM. To each of the Brazilian curators for allowing the loan and examination of material, without their help
this study would not have been possible. Our gratitude to L. Pereira and C. Damborenea at MLP, and M.J. Ramírez
at MACN. To J. Kekenbosch from Belgium, USA curators G. Giribet from MCZ and L.C. Dunn from PMNH to
provide holotype photographs and loans. Our gratitude also to Diana Silva at MUSM-ENT (Peru) for providing the
material she collected by decades. Part of this publication is a product of the first author’s Internship at the Instituto
Butantan, São Paulo, Brazil, and of his doctorate research at MCT/ PUCRS. This study was supported by CNPq
(ADB grant PQ 303903/2019-8) and CAPES PROSUC (Programa Suporte à Pós-Graduação IES Comunitárias—
88887.179657/2018-00).
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Zootaxa 5414 (1): 001–083 ISSN 1175-5326 (print edition)

Accepted by A. Santos: 19 Jan. 2024; published: 19 Feb. 2024

FIRST PUBLISHED IN 2024 BY

© 2024 Magnolia Press

ISSN 1175-5326 (Print edition)

 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Key words: Diversity, taxonomy, wetlands, Andes, distribution

Material and Methods

 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Lycosidae Sundevall, 1833

Arctosa C.L. Koch, 1847

 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Identification key for Neotropical Arctosa species

cinerea group, males

cinerea group, females

jibarosa group, males

jibarosa group, females

Description. Male. Unknown.

 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

10 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

12 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

14 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

16 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Arctosa costenola sp. nov.

18 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

20 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Arctosa denticulata Jiménez & Dondale 1984

22 · Zootaxa 5414 (1) © 2024 Magnolia Press Paredes-Munguía et al.

Arctosa jibarosa sp. nov.

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Arctosa littoralis (Hentz, 1844)

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Description. Male. Unknown.

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Arctosa pacaya sp. nov.

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Description. Female. Unknown.

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Arctosa sapiranga Silva & Lise 2009

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Arctosa serii Roth & Brown, 1976

Arctosa tenebrosa (Keyserling, 1877) comb. nov.

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Arctosa tenella (Keyserling, 1877) comb. nov.

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Arctosa villa sp. nov.

Arctosa sp. nov.: Paredes-Munguia 2012: 42

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Hogna andina (Chamberlin, 1916) comb. nov.

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Hogna pugil (Bertkau, 1880) comb. nov.

Prolycosides aussereri (Keyserling, 1877) comb. nov.

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Trochosa fusca (Keyserling, 1877) comb. nov.

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