“Flying” or digging? The trapdoor spider genus Neocteniza Pocock, 1895: Redescription of three species, new records from Brazil, notes on natural history and first record of ballooning for Idiopidae (Araneae, Mygalomorphae)

Abstract

The Neotropical trapdoor spider genus Neocteniza Pocock, 1895 is distributed throughout Central and South America. The genus currently comprises 18 species and little is known about the biology of these spiders. We provide an amended diagnosis for the genus Neocteniza and the species: N. minima Goloboff, 1987, N. australis Goloboff, 1987 and N. toba Goloboff, 1987, and also redescribe them. We include new records of these species from Brazil and notes on natural history of N. toba, providing data on burrow structure, diet, development and the first record of ballooning behavior for Idiopidae. We propose a terminology for the genitalia and consider homology among genital structures and among possibly stridulatory structures. We also discuss relationships of Neocteniza with other Idiopidae, stressing the uniqueness of the genus and its differences from all other Idiopidae including other Genysinae.

Full text

ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by R. Raven: 3 Aug. 2021; published: 20 Aug. 2021 451
Zootaxa 5023 (4): 451–485
https://www.mapress.com/j/zt/
Copyright © 2021 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5023.4.1
http://zoobank.org/urn:lsid:zoobank.org:pub:600DF2C1-7249-45E7-A0C6-A6FAEE76A3C6
“Flying” or digging? The trapdoor spider genus Neocteniza Pocock, 1895:
Redescription of three species, new records from Brazil, notes on natural history
and first record of ballooning for Idiopidae (Araneae, Mygalomorphae)
GIULLIA DE F. ROSSI1, VICTOR M. GHIROTTO2*, ARTHUR GALLETI-LIMA1,
RAFAEL P. INDICATTI2,3 & JOSÉ P. L. GUADANUCCI2
1Programa de Pós-graduação em Ciências Biológicas (Zoologia), Departamento de Biodiversidade, Instituto de Biociências,
Universidade Estadual Paulista, Rio Claro, São Paulo, Brazil;
2Departamento de Biodiversidade, Instituto de Biociências, Universidade Estadual Paulista, Rio Claro, São Paulo, Brazil;
3Laboratório de Coleções Zoológicas, Instituto Butantan, São Paulo, São Paulo, Brazil.
https://orcid.org/0000-0002-3909-0168
https://orcid.org/0000-0002-6596-3612
https://orcid.org/0000-0001-9050-1623
https://orcid.org/0000-0002-1984-8621
https://orcid.org/0000-0001-9724-9010
*Corresponding author.
�
Victorghirotto@hotmail.com
Abstract
The Neotropical trapdoor spider genus Neocteniza Pocock, 1895 is distributed throughout Central and South America.
The genus currently comprises 18 species and little is known about the biology of these spiders. We provide an amended
diagnosis for the genus Neocteniza and the species: N. minima Goloboff, 1987, N. australis Goloboff, 1987 and N. toba
Goloboff, 1987, and also redescribe them. We include new records of these species from Brazil and notes on natural
history of N. toba, providing data on burrow structure, diet, development and the first record of ballooning behavior
for Idiopidae. We propose a terminology for the genitalia and consider homology among genital structures and among
possibly stridulatory structures. We also discuss relationships of Neocteniza with other Idiopidae, stressing the uniqueness
of the genus and its differences from all other Idiopidae including other Genysinae.
Key words: Genysinae, stridulatory apparatus, sexual dimorphism, haematodocha, Cerrado, Atlantic Forest, behavior
Resumo
“Voando” ou cavando? O gênero de aranhas-de-alçapão Neocteniza Pocock, 1895: redescrição de três espécies,
registros novos do Brasil, notas sobre a história natural e primeiro registro de balonismo para Idiopidae (Araneae,
Mygalomorphae). O gênero neotropical de aranhas de alçapão Neocteniza Pocock, 1895 é distribuído pela América
Central e do Sul. O gênero atualmente é composto por 18 espécies e pouco é conhecido sobre a biologia destas aranhas.
Nós apresentamos uma diagnose modificada para o gênero Neocteniza e as espécies: N. minima Goloboff, 1987, N.
australis Goloboff, 1987 e N. toba Goloboff, 1987, também as redescrevendo. Incluímos novos registros destas
espécies para o Brasil e notas sobre a história natural de N. toba, providenciando dados sobre a estrutura da toca, dieta,
desenvolvimento e o primeiro registro do comportamento de balonismo para Idiopidae. Propomos uma terminologia para
as genitálias, considerando a homologia entre as estruturas genitais e entre estruturas possivelmente estridulatórias. Além
disso, discutimos as relações entre Neocteniza e outros Idiopidae, enfatizando a singularidade do gênero e suas diferenças
perante todos os outros Idiopidae incluindo outros Genysinae.
Palavras-chave: Genysinae, aparelho estridulatório, dimorfismo sexual, hematodoca, Cerrado, Mata Atlântica,
comportamento

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Introduction
Several spiders from different families of the Infraorder Mygalomorphae construct trapdoor burrows for living
and sheltering. These doors and burrows offer physical protection, including from UV rays in open areas, and
keep temperature and humidity stable compared to the external environment (Mason et al. 2013). Some species
camouflage the trapdoor lid with leaf fragments, sticks or moss, hiding from predators and enhancing their sit-and-
wait strategy to catch prey (Mason et al. 2013; Gupta et al. 2015). Among the spider families that build specialized
trapdoor burrows are the Idiopidae (Coyle et al. 1992; Raven 1985; Rix et al. 2017). Spiders of this family are
distributed worldwide, mainly on parts of the former supercontinent Gondwana (Raven 1985), and comprise 414
species in 23 genera (World Spider Catalog 2021).
Two genera occur in the Americas: Idiops Perty, 1833 (Idiopinae Simon, 1889a) and Neocteniza Pocock, 1895
(Genysinae Simon, 1903) (Raven 1985). The latter are widely distributed throughout the Neotropics, and currently
comprise 18 species (World Spider Catalog 2021). They are yellowish-brown to entirely dark, small sized (4.7 –
27.5 mm in body length), and exhibit strong sexual dimorphism, with females usually three to four times larger than
males (Platnick & Shadab 1976; Goloboff 1987, fig. 1). Spiders belonging to Neocteniza are distinguished from
other idiopid genera by the following combination of features: two eye rows, strong rastellum, recurved tripartite
thoracic fovea (T-shaped) and male palpal bulb with separated (not fused) apical sclerites and well developed
median haematodocha (Goloboff 1987). Apart from a few observations in taxonomical studies, little is known about
their biology and behavior (Goloboff 1987).
In this study, we present an amended diagnosis and proposition of five stridulatory apparatus’ for Neocteniza,
a discussion on relationships with other Idiopidae as well as other Genysinae, and new records from Brazil for: N.
australis Goloboff, 1987 from the Atlantic Forest, N. minima Goloboff, 1987 from the Cerrado (Brazilian savannah)
and N. toba Goloboff, 1987 from both biomes. We also present an amended diagnosis for these species, and notes
on natural history for N. toba, with the first record of ballooning behavior for Idiopidae.
Material and methods
Specimens studied herein are deposited in the following institutions: Coleção Aracnológica de Diamantina,
Universidade Estadual Paulista ‘’Júlio de Mesquita Filho’’, Rio Claro, Brazil (CAD, J.P.L. Guadanucci); Laboratório
de Estudos Subterrâneos, Universidade Federal de São Carlos, São Carlos, Brazil (LES, M.E. Bichuette); Instituto
Butantan, São Paulo, Brazil (IBSP, A.D. Brescovit); Museu de Ciências e Tecnologia, Pontifícia Universidade
Católica, Porto Alegre, Brazil (MCTP, R.A. Teixeira); Museu de Zoologia da Universidade de São Paulo, São Paulo,
Brazil (MZSP, R. Pinto da Rocha); Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires,
Argentina (MACN, C. Scioscia; M. Ramírez). Measurements are given in millimeters. Images and measurements
were obtained with a Leica M205C Stereomicroscope with the program Leica Application Suite V4.12. The length
of leg segments was measured between joints in dorsal view. Total body length includes chelicerae but not the
pedicel and spinnerets. Micrographs were taken with a Hitachi TM-3000 scanning electron microscope (SEM) at
the Laboratório de Microscopia, Departamento de Biologia, UNESP and Hitachi TM-1000 SEM at the Queensland
Museum, South Brisbane.
Terminology for general structures, bulb positions and spine notation follows Goloboff (1987), except for
genital structures. We use new terminology for the relatively complex genitalia of this genus. See discussion for
further comments on the genitalia.
Abbreviations and acronyms. General structures: Bt, bothrium; CG, cheliceral grooves; F, fovea; FT, filiform
trichobothrium; It, integument with scaly cuticle; ITC, inferior tarsal claw; Pa, patella; PLS, posterior lateral
spinnerets; PMS, posterior median spinnerets; Ra, rastellum; STC, superior tarsal claws; Ti, palpal tibia; TO, tarsal
organ. Spination and setae positioning: ant, anterior; ap, apical; b, basal; D, dorsal; inf, inferior; P, prolateral;
post, posterior; R, retrolateral; sup, superior; V, ventral. Spermathecae: At, spermathecal atrium; Du, spermathecal
duct; IDu, inferior spermathecal duct; R, receptaculum; SB, spermathecal base; SDR, spermathecal duct ring; SDu,
superior spermathecal duct. Male palp and bulb: AH, apical haematodocha; AK, apical keel; ALoK, anterior
locking keel; BH, basal haematodocha; Cy, cymbium; EA, embolic aperture; EC, embolic crack; Em, embolus;
MH, median haematodocha; PA, paraembolic apophysis; PLoK, posterior locking keel; RTG, retrolateral palpal

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tibia grooves; SD, seminal duct; SLoK, subapical locking keel; St, subtegulum; T, tegulum; TE, tibial excavation;
TG, tegular grooves; TS, ventro-retrolateral tibial spines; TT, tegular teeth; VTA, ventral tegular apophysis; VTG,
ventral palpal tibia grooves. Burrow structure: Cha, chamber; CDC, cemented disposal chamber.
Taxonomy
Family Idiopidae Simon, 1889
Subfamily Genysinae Simon, 1903
Genus Neocteniza Pocock, 1895
Neocteniza Pocock, 1895: 193 (type species by monotypy, N. sclateri Pocock, 1895). Platnick & Shadab, 1976: 5; Raven, 1985:
17, 138; Goloboff, 1987: 33; World Spider Catalog, 2021.
Amended diagnosis: Neocteniza can be distinguished from other idiopid genera (and from all remaining
Mygalomorphae (Goloboff 1987)) by an embolic crack (EC) on the palpal bulbs of males from where the embolus
naturally breaks off once it is inserted and attached in the duct of the female’s spermathecae. Additionally, the
following combination of features also distinguish Neocteniza from other Idiopidae: two eye rows, strong rastellum,
recurved tripartite thoracic fovea (T-shaped) and lacking clavate trichobothria on all tarsi and palps in both sexes,
male prolateral region of cymbium without angular lobe and palpal bulb with separated (not fused) apical sclerites
and well developed median haematodocha (Goloboff 1987). See more details in the discussion.
Distribution: Central and South America: Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador,
Guatemala, Guyana, Hondura, Panama, Paraguay, Peru, Venezuela.
Neocteniza minima Goloboff, 1987
(Figs 1–3)
Neocteniza minima Goloboff, 1987: 37, figs 6, 7, 13–16, 20–24, 32–36. Female holotype from Argentina: Jujuy, Yuto, 13.v.1983,
M. Vinãs & P. Goloboff coll., MACN 7947 and male paratype from 3 km W. Campamento Vespucio, same province, date,
MACN 7946, not examined. Goloboff & Platnick, 1992: 6; World Spider Catalog, 2021.
Additional material examined: Brazil, Mato Grosso do Sul: Corumbá, V. Ferreira coll., vi.2003, 1 male (MCTP
17590). Goiás: Mambaí, M.E. Bichuette; J.E. Gallão; D.M. Schimonsky coll., 29.iv.2013, 1 female (LES 015324);
same data, (Gruta da Tarimba), 1 female (LES 015323); Rio Verde, 1 male, without data (CAD 792); 1 juvenile
(CAD 793).
Diagnosis: Females of N. minima are distinguished from those of the australis group species (sensu Goloboff
1987) by the very sclerotized inferior spermathecal duct (IDu) (Fig. 1A), and from other species of the sclateri
group (sensu Goloboff 1987) (except N. agustinea Miranda & Arizala, 2013; N. coylei Goloboff & Platnick, 1992;
N. platnicki Goloboff, 1987 and N. spinosa Goloboff 1987) by the receptacle being uniformly rounded and of
smaller size in relation to the spermathecal base, and symmetrical lobes (Goloboff 1987, figs 13, 14) (Fig. 1A).
They are further distinguished from remaining species by the conspicuous constriction of the superior spermathecal
duct (SDu) (Fig. 1A). Males of N. minima are distinguished from the australis group species by the thinner embolus
of palpal bulb. Males are distinguished from those of N. fantastica Platnick & Shadab, 1976 by the presence of a
paraembolic apophysis (PA) on the bulb, and from those of N. pococki Platnick & Shadab, 1976 by the rounded PA
(Fig. 1E–G), and additionally, from both by bearing fewer tibial ventro-retrolateral spines (TS), ca. 20 in N. minima
(Fig. 1H, I) vs ca. 32 in N. pococki and 58 in N. fantastica.
Female (LES 00015323)
Color in ethanol: Carapace light brown (Fig. 2A), abdomen light grayish (Fig. 2C). Legs and palps light brown
(Fig. 2A, B).
Total length 14.38. Carapace 4.55 long, 3.39 wide; cephalic region strongly convex, fovea deep, recurved (T-

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shaped) (Fig. 2A). Anterior ocular row slightly procurved, posterior ocular row recurved (Fig. 2A). Eye sizes: AME
0.15, ALE 0.27, PME 0.17, PLE 0.23. Eye interdistances: AME–ALE 0.38, PME–PLE 0.15, AME–AME 0.17,
ALE–PLE 0.23, PME–PME 1.13. Eye tubercle 1.51 long, 0.58 wide. Chelicerae with rastellum formed by 12 thick,
blunt spines (Fig. 2D). Chelicerae row with 9 teeth on the promargin, 10 central teeth, 8 larger teeth on retromarginal
row. Labium 0.80 long, 0.82 wide, without cuspules (Fig. 2B). Palpal coxae with a single blunt cuspule on anterior
prolateral edge. Sternum 2.59 long, 2.35 wide. Sigilla barely discernible (Fig. 2B). Palp measurements: trochanter
0.34/ femur 0.52/ patella 1.40/ tibia 0.75/ tarsus 1.11/ total 4.12. Leg measurements: I trochanter 0.26/ femur 2.36/
patella 1.55/ tibia 1.43/ metatarsus 1.14/ tarsus 0.61/ total 7.35; II 0.28/ 2.46/ 1.87/ 1.17/ 1.30/ 0.58/ 7.66; III 0.25/
2.20/ 1.45/ 1.03/ 1.53/ 0.80/ 7.26; IV 0.46/ 2.83/ 1.42/ 1.44/ 2.23/ 0.66/ 9.04. ITC on all legs. Preening combs on
all metatarsi, I, II, 2 VP; III, 3 VP, 3 VR; IV, 2 VP, 3 VR. Integument with flattened scaly cuticle with highest distal
ends on all legs and palp. Tarsi I–IV integral. All tarsi without scopula. Spinnerets short; PMS with ca. 50 spigots,
basal article of PLS with ca. 90, medial article with ca. 55, apical article with 43 larger on apex. Spinnerets length:
PMS 0.71, basal article of PLS 0.97, medial article 0.35, apical article 0.30, short, domed (Fig. 2E). Spermathecae
curved, presenting sclerotized SB, bearing IDu, SDu, and with thin receptacle (R) (Fig. 1A–D).
Trichobothria: Tibia: legs: I, prolateral row 6, retrolateral 5; II, prolateral row 4, retrolateral 6; III, prolateral
row 4, retrolateral 6; IV, prolateral row 6, retrolateral 4; palp, prolateral row 5. Metatarsi: I, retrolateral row 5; II,
prolateral row 2, retrolateral 3; III, retrolateral row 7; IV, retrolateral row 7. Tarsi: I, both rows 2; II, prolateral row
1, retrolateral 2; III, both rows 3; IV, both rows 2; palp, prolateral row 2, retrolateral 3.
Spination: Femora: IV, 15 D ant ap, huddled. Patellae: III, 23/21 D ant ap, forming a triangle, 0/1 R ap; IV, 11/4
D ant b, huddled. Tibia: I, 1/2 D; III, 16/19 D; palp, 5/15 D ant Metatarsi: I, row of 9/10 D ant, 9/12 D post; II, 7 D
ant, 9/7 D post; III, 19/16 D, huddled, 1 V ap medial; IV 3/5 D ant, 2/4 D post, 5/3 V ap, 2/3 V. Tarsi: I, 7/6 D ant,
6/5 D post; II, 5/4 D ant, 4 D post; palp, 17/9 D, 2 V.
Male (CAD 792)
Color in ethanol: Carapace light brown (Fig. 3A), abdomen light gray (Fig. 3C). Legs and palps light brown
(Fig. 3A, B).
Total length 5.60. Carapace 2.00 long, 1.60 wide; cephalic region strongly convex, fovea deep, recurved (T-
shaped) (Fig. 3A). Anterior and posterior ocular row slightly recurved (Fig. 3A). Eye sizes: AME 0.16, ALE 0.14,
PME 0.10, PLE 0.10. Eye interdistances: AME–ALE 0.04, PME–PLE 0.06, AME–AME 0.08, ALE–PLE 0.07,
PME–PME 0.30. Eye tubercle 0.69 long, 0.37 wide. Chelicerae with rastellum formed by 3 thick, blunt spines on
edge of cheliceral prolongation and two laterally adjacent smaller spines outside cheliceral prolongation (Fig. 3D).
Intercheliceral tumescence absent. Cheliceral promarginal row with 6 teeth, central teeth absent, 8 smaller teeth on
retromarginal row. Labium 0.20 long, 0.44 wide, without cuspules. Palpal coxae without cuspules. Sternum 1.31
long, 1.03 wide. Sigilla barely discernible (Fig. 3B). Palp measurements: trochanter 0.27/ femur 0.78/ patella 0.38/
tibia 0.60/ cymbium 0.44/ total 2.47. Leg measurements: I trochanter 0.45/ femur 1.66/ patella 0.75/ tibia 1.27/
metatarsus 1.16/ tarsus 0.66/ total 5.95; II 0.41/ 1.50/ 0.72/ 1.00/ 1.05/ 0.64/ 5.32; III 0.42/ 1.14/ 0.70/ 0.76/ 1.21/
0.77/ 5.00; IV 0.47/ 1.79/ 0.79/ 1.60/ 1.87/ 0.88/ 7.40. ITC on all legs. Preening combs on all metatarsi, I, II, 2 VP;
III, IV, 3 VP, 3 VR. Integument with flattened scaly cuticle with highest distal ends on all legs and palp (similar to
Pseudonemesia spp., Microstigmatidae, see Raven & Platnick 1981, figs 5, 6; Indicatti & Villarreal-M 2016, fig.
4A, C, D). Tarsi I–IV integral. All tarsi without scopula (Fig. 3F). Spinnerets short; PMS with ca. 18 spigots, basal
article of PLS with ca. 68, medial article with ca. 36, apical article with 26 larger on apex. Spinnerets length: PMS
0.27, basal article of PLS 0.47, medial article 0.25, apical article 0.34, short, domed (Fig. 3E). Palpal tibia incrassate
and with spines (Fig. 1H–J). Epigastric area with ca. 16 epiandrous spigots. Bulb with large and strongly twisted
tegulum and thin embolus, filiform, presenting: Haematodochae BH, MH, AH; Tegular grooves (TG); Tegular teeth
(TT); Paraembolic apophysis (PA); keel AK; EC in the embolus (Fig. 1E–J).
Trichobothria: Tibia: I, both rows 4; II, both rows 3; III, both rows 3; IV, prolateral row 5, retrolateral 4; palp,
retrolateral row 2, prolateral 4. Metatarsi: I, prolateral row 1, retrolateral 3; II, retrolateral row 3; III, median row 3;
IV, retrolateral row 5. Tarsi: I, III and IV, both rows 2; II, prolateral row 2, retrolateral 3.
Spination: Femora: IV, 6/7 D–P sup ap, huddled. Patellae: III, 5/6 D–P ap and 8/5 D–P IV, 4/3 P b. Tibia: III, 2
P (ant 2:2), and 2 D, 1 R ap sup, 3 very thin V; IV, 1 very thin, V. Metatarsi: III, 3 D ap, ca. 5 very thin, V; IV, ca. 5
very thin.

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FIGURE 1. A–J. A–C, E–J Neocteniza minima Goloboff, 1987. D Neocteniza sp., sclateri group: A–D. Female, spermathecae:
A. Dorsal view; B. Frontal view; C, D. Posterior view; D. Arrow showing a broken embolus. E–J. Right male palpal bulb
(mirrored): E. retrolateral view; F. prolateral view; G. Palp and PA and EC in detail, ventral view. H–J. Male, left palpal tibia:
H. Ventro-prolateral view; I. Frontal view; J. Prolateral view; Abbreviations and acronyms. Spermathecae: At, spermathecal
atrium; IDu, inferior spermathecal duct; R, receptaculum; SB, spermathecal base; SDu, superior spermathecal duct. Male palp
and bulb: AH, apical haematodocha; AK, apical keel; BH, basal haematodocha; Ti, palpal tibia; Cy, cymbium; EA, embolic
aperture; EC, embolic crack; Em, embolus; MH, median haematodocha; PA, paraembolic apophysis; RTG, retrolateral palpal
tibia grooves; SD, seminal duct; St, subtegulum; T, tegulum; TE, tibial excavation; TG, tegular grooves; TS, ventro-retrolateral
palpal tibia spines; TT, tegular teeth; VTG, ventral palpal tibia grooves. Scale bars: A–D, H, I = 0.2 mm; E–G, J = 0.5 mm.

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FIGURE 2. A–E. Neocteniza minima Goloboff, 1987, female. A, B. Prosoma: A. Dorsal view; B. Ventral view; C. Opisthosoma,
dorsal view; D. Chelicerae, ventral view; E. Spinnerets, ventral view. Abbreviations and acronyms. F, fovea; PLS, posterior
lateral spinnerets; PMS, posterior median spinnerets; Ra, rastellum. Scale bars: A–C = 1 mm; D, E = 0.5 mm.

NEOCTENIZA: REDESCRIPTION, RECORDS AND NATURAL HISTORY Zootaxa 5023 (4) © 2021 Magnolia Press · 457
FIGURE 3. A–F. Neocteniza minima Goloboff, 1987, male. A, B. Prosoma: A. Dorsal view; B. Ventral view; C. Opisthosoma,
dorsal view; D. Chelicerae, ventral view; E. Spinnerets, ventral view; F. Leg I, retrolateral view. Abbreviations and acronyms.
CG, cheliceral grooves; F, fovea; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets; Ra, rastellum. Scale bars:
A–C, F = 1 mm; D, E = 0.5 mm.

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Variation: Females (n=3): total length 11.00–14.38; carapace 4.05–6.88 long; abdomen 5.33–7.48 long. Palpal
coxae with 1–3 blunt cuspules. Males (n=4): total length 5.60–6.48; carapace 2.00–2.93 long; abdomen 2.60–2.88
long.
Distribution: Argentina, Bolivia (Goloboff 1987); herein recorded for the first time for Brazil, in the Cerrado
(Brazilian savanna) (Fig. 13).
Neocteniza australis Goloboff, 1987
(Figs 4, 5)
Neocteniza australis Goloboff, 1987: 44, figs 1–4, 9, 10, 27–31, 37–41. Female holotype from Argentina: Buenos Aires, 10
km Puerto Obligado, 20.i.1983, MACN 5983, not examined and male paratype, same locality as holotype, 20.iii.1983, A.
Zanetic & P. Goloboff coll., MACN 7959, examined by photos. Goloboff & Platnick, 1992: 9, figs 5, 6 (male); Grismado
& Goloboff, 2014: 109, figs 6–8, 12, 13 (female/male); World Spider Catalog, 2021.
Additional material examined: Brazil, Santa Catarina: Urussanga (Rio Molha), collected with pitfall traps, R.A.
Teixeira coll., 28.xii.2006, 1 male (IBSP 113434, Grismado & Goloboff 2014, fig. 7); Rio Grande do Sul: Augusto
Pestana, 12.xi.2008, 1 male (MCTP 26144); 27.xi.2008, 1 male (MCTP 26146); 12.xii.2008, 5 males (MCTP
26147–26151); 21.iii.2009, 1 male (MCTP 27376); 04.iv.2009, 1 male (MCTP 27377), all without collector.
Diagnosis: Males of N. australis are distinguished from those of species of the sclateri group by the thick
embolus (Goloboff 1987, figs 37–40) (Fig. 4A–G), from N. myriamae Bertani et al., 2006 by the thinner embolus,
from N. chancani Goloboff & Platnick, 1992 by the absence of a basal constriction of the embolus, and from N.
toba by the straighter embolus (Goloboff & Platnick 1992, figs 5, 6; Bertani et al. 2006) (Fig. 4A–J). Females not
examined, see Goloboff (1987: 44) for female diagnosis.
Male (MCTP 27377)
Color in ethanol: Carapace light brown (Fig. 5A), abdomen light gray (Fig. 5C). Legs and palps light brown (Fig.
5A, B).
Total length 8.91. Carapace 3.67 long, 3.16 wide; cephalic region strongly convex, fovea deep, recurved (T-
shaped) (Fig. 5A). Anterior ocular row slightly procurved, posterior ocular row slightly recurved (Fig. 5A). Eye
sizes: AME 0.22, ALE 0.22, PME 0.17, PLE 0.19. Eye interdistances: AME–ALE 0.03, PME–PLE 0.01, AME–
AME 0.06, ALE–PLE 0.04, PME–PME 0.44. Eye tubercle 0.71 long, 1.12 wide. Chelicerae with rastellum formed
by 8 thick, blunt spines (Fig. 5D). Intercheliceral tumescence absent. Cheliceral promarginal row with 6 teeth, 21
central teeth, 2 larger teeth on retromarginal row. Labium 0.40 long, 0.78 wide, without cuspules. Palpal coxae
without cuspules. Sternum 2.39 long, 1.74 wide. Sigilla barely discernible (Fig. 5B). Palp measurements: trochanter
0.75/ femur 1.98/ patella 0.72/ tibia 1.88/ cymbium 1.15/ total 6.48. Leg measurements: I trochanter 0.57/ femur
2.81/ patella 0.64/ tibia 1.63/ metatarsus 2.07/ tarsus 1.25/ total 8.97; II 0.38/ 2.06/ 1.46/ 2.05/ 1.62/ 0.98 / 8.55; III
0.54/ 1.90/ 0.56/ 1.54/ 2.23/ 1.50/ 8.27; IV 0.45/ 2.88/ 1.71/ 3.20/ 3.12/ 1.67/ 13.03. ITC on all legs. Preening combs
on metatarsi II, 2 VP; III, 4 VP, 4 VR; IV 5 VP, 7 VR. Integument with flattened scaly cuticle with highest distal
ends on all legs and palp. Tarsi I–IV integral. All tarsi without scopula (Fig. 5F). Spinnerets short; PMS with ca. 15
spigots, basal article of PLS with ca. 26, medial article with ca. 24, apical article with 13 larger on apex. Spinnerets
length: PMS 0.41, basal article of PLS 0.63, medial article 0.26, apical article 0.19, short, domed (Fig. 5E). Palpal
tibia incrassate and with spines (Fig. 4D, F, I). Epigastric area with ca. 41 epiandrous spigots. Bulb with large,
slightly twisted tegulum and lanceolate embolus, presenting: Haematodochae BH, MH, AH; Tegular grooves (TG);
Tegular teeth (TT); ventral tegular apophysis (VTA); keels ALoK, PLoK, SLoK and AK; EC near or reaching the
tegulum (Fig. 4A–J). Note the empty, unexpanded haematodochae in Fig. 4I, J.
Trichobothria: Tibia: I, both rows 3; II, prolateral row 4, retrolateral 6; III, prolateral row 4, retrolateral 6; IV,
prolateral row 6, retrolateral 4; palp, prolateral row 5. Metatarsi: I, retrolateral row 5; II, prolateral row 2, retrolateral
3; III, retrolateral row 7; IV, retrolateral row 7. Tarsi: I, both rows 2; II, prolateral row 1, retrolateral 2; III, both rows
3; IV, both rows 2; palp, prolateral row 2, retrolateral 3.
Spination: Femora: IV, 22 D–P sup ap, huddled. Patellae: III, 11 sup ap, huddled; IV, 16 D–P. Tibia: I, 3 V ap;
II, 2 V ap; III, 2 P, 3 D, 4 V; IV, 9 V. Metatarsi: I, 4 V; II, 6 V; III, 12 D, 5 V; IV, 10 V.
Variation: Males (n=5): total length 8.91–9.61; carapace 3.30–3.67 long; abdomen 3.96–4.67 long. Palpal coxae
without cuspules. Some individuals also with 2 VP preening combs on leg I.

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FIGURE 4. A–J. Neocteniza australis Goloboff, 1987, male. A–C. Left male palpal bulb: A. Retrolateral view; B. Palp and detail
of embolus, prolateral view; C. Palp and detail of embolus, prolateral-ventral view. D–F. Right palpal tibia: D. Retrolateral; E.
Prolateral; F. Ventral view. G, H. Palpal bulb, ventral-prolateral view: H. Detail of EC; I, J. Empty bulb: I. Palpal tibia (ventral
view) and broken bulb (ventral-prolateral view); J. Same, detail. Abbreviations and acronyms. AH, apical haematodocha;
AK, apical keel; ALoK, anterior locking keel; BH, basal haematodocha; VTA, ventral tegular apophysis; Fe, femur; Pa, patella;
Ti, palpal tibia; Cy, cymbium; EA, embolic aperture; EC, embolic crack; Em, embolus; MH, median haematodocha; PLoK,
posterior locking keel; RTG, retrolateral palpal tibia grooves; SD, seminal duct; SLoK, subapical locking keel; St, subtegulum;
T, tegulum; TG, tegular grooves; TS, ventro-retrolateral palpal tibia spines; TT, tegular teeth. Scale bars: A–C, G, I = 0.5 mm;
D–F = 1 mm; H, J = 0.2 mm.

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FIGURE 5. A–F. Neocteniza australis Goloboff, 1987, male. A, B. Prosoma: A. Dorsal view; B. Ventral view; C. Opisthosoma,
dorsal view; D. Chelicerae, ventral view; E. Spinnerets, ventral view; F. Leg I and detail of tarsus I, retrolateral view.
Abbreviations and acronyms. CG, cheliceral grooves; Ra, rastellum. Scale bars: A–D, F = 1mm; E = 0.5 mm.

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Distribution: Argentina (Goloboff 1987); herein recorded for the first time for Brazil, inhabiting the Atlantic
Forest (Fig. 13).
Neocteniza toba Goloboff, 1987
(Figures 6–12, 15D, 17–19)
Neocteniza toba Goloboff, 1987: 41, figs 8, 25, 26. Female holotype from Argentina: Salta, La Quena, 15–16.v.1983, M. Viñas
& P. Goloboff coll., MACN 7952, not examined. Goloboff & Platnick, 1992: 7, figs 16–19 (male); World Spider Catalog,
2021.
Neocteniza australis: Fukami et al., 2004: 30 (misidentification).
Additional material examined: Brazil, São Paulo: Jaboticabal, 21°16’S 48°19’W, F.J. Cividanes coll., 2006, 4
males (IBSP 113032); Matão (Fazenda Cambuhy, Mata da Virgínia), J. Ricetti coll., 1 male (IBSP); Araraquara
(Fazenda IBIOTEQ, UNIARA), A. Galleti-Lima & R.P. Indicatti coll., 19.xi.2019, 1 female juv. (CAD 808); 1
female (CAD 809); 1 female, 2 juv. (CAD 810); 1 female (IBSP); 1 female (MZSP); Pirassununga, A. Galleti-Lima,
M.J.A. Morales, R. Fonseca-Ferreira coll., 19.ii.2019, 1 female (CAD 811); Rio Claro (UNESP campus, Fig. 14A,
B), V.M. Ghirotto coll., 19.iii.2018; died 16.v.2018, 1 female (CAD 795); 1 female, 2 nymphs and 5 first instars
fixed with female, 19.iii.2018, died 02.x.2018 (CAD 796); 1 female, 18.iii.2018 (CAD 797); 1 female, 20.vii.2018
(MCTP); 2 males, 24.vi.2019, V.M. Ghirotto & M. Negreiros coll. (CAD 798); 1 female, 13.vii.2019 (CAD 801);
1 male, 19.iii.2019, V.M. Ghirotto coll. (CAD 802); 1 male, 02.v.2019, V.M. Ghirotto coll. (CAD 803); 1 male,
ix.2019, V.M. Ghirotto & E.F. Trova coll. (CAD 804); 2 males, pitfall traps, 25.iii.2018 (CAD 787; 788); 1 male,
pitfall traps, 23.iii.2018 (CAD 789); 1 male, pitfall traps, 29.iii.2018 (CAD 790); 1 male, wandering on the ground,
20.xi.2017 (CAD 794); 02.v.2019, V. Ghirotto & R.P. Indicatti coll., 1 female (MZSP); 1 female (IBSP); Itirapina
(Estação Ecológica de Itirapina), 22°13’01”S, 47°51’11”W, 13.iii.2019, E.F. Trova coll., 1 female (CAD 786); 1
female, 2018, E.F. Trova coll. (CAD 791); 1 female, 13.iii.2019, E.F. Trova coll. (CAD 799); 1 female, 04.xii.2018,
E.F. Trova coll. (CAD 800); Teodoro Sampaio (Parque Estadual Morro do Diabo, Fig. 14C, D), collected with pitfall
traps, 24–31.iii.2003, C.A. Rheims, R.P. Indicatti, D.F. Candiani (Biota) coll., 4 males (IBSP), as in Candiani et
al. (2005: 3); 15–21.ii.2020, A. Galleti-Lima, R.P. Indicatti, J.P.L. Guadanucci coll., 2 female (CAD 805; 806); 1
juvenile (CAD 807).
Diagnosis: Females of N. toba are distinguished from those of the sclateri group species by the membranous
spermathecal base (SB) (Goloboff 1987) (Fig. 6A–D). They are distinguished from those of N. australis by the
larger and straighter spermathecal receptacles (Fig. 6A–D). Males of N. toba are distinguished from those of the
sclateri group species by the thick embolus (Goloboff 1987, figs 37–40) (Fig. 6E–G); and from those of N. australis,
N. chancani and N. myriamae by the curved aspect and apically widened embolus (Goloboff & Platnick 1992, figs
5, 6; Bertani et al. 2006, figs 1–3) (Fig. 6E–G). Additionally, from males of N. myriamae by the lack of a basal
sclerotized, pointed process on the tegulum (Bertani et al. 2006, figs 1–3).
Female (CAD 795)
Color in life (Fig. 8A–D) and in ethanol (Fig. 9A–C): Entirely black, with grayish abdomen, usually fading to
dark reddish brown in ethanol.
Total length 14.56. Carapace 6.99 long, 6.35 wide; cephalic region strongly convex, fovea deep, recurved (T-
shaped) (Fig. 9A). Anterior and posterior ocular row slightly procurved (Fig. 9A). Eye sizes: AME 0.25, ALE 0.37,
PME 0.21, PLE 0.31. Eye interdistances: AME–ALE 0.42, PME–PLE 0.14, AME–AME 0.25, ALE–PLE 0.23,
PME–PME 1.00. Eye tubercle 1.25 long, 2.22 wide. Chelicerae with rastellum formed by 15 thick, blunt spines
on cheliceral prolongation and two laterally adjacent spines outside cheliceral prolongation (Fig. 9D). Cheliceral
promarginal row with 4 larger and 4 smaller teeth, 12 central teeth, 12 teeth on retromarginal row. Labium 1.25 long,
1.33 wide, without cuspules. Palpal coxae with 2–3 pointed cuspules on anterior prolateral edge. Sternum 5.64 long,
3.81 wide. Sigilla barely discernible (Fig. 9B). Palp measurements: trochanter 0.78/ femur 3.01/ patella 1.80/ tibia
1.89/ tarsus 2.11/ total 9.59. Leg measurements: I trochanter 1.14/ femur: 3.75/ patella 2.64/ tibia 2.50/ metatarsus
2.05/ tarsus 0.98/ total 13.06; II 1.16/ 3.55/ 2.51/ 2.22/ 2.10/ 1.00/ 12.54; III 1.40/ 3.15/ 2.51/ 1.69/ 2.50/ 1.40/ 12.65;
IV 2.00/ 4.55/ 2.93/ 3.00/ 3.44/ 1.32/ 17.24. ITC on all legs (Fig. 10A–D). Preening combs on metatarsi II, 3 VP; III, 3
VP, 3 VR; IV, 4 VR (Fig. 10A–D). Integument with flattened scaly cuticle with highest distal ends on all legs and palp.

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FIGURE 6. A–M. Neocteniza toba Goloboff, 1987. A–D. Female, spermathecae: A. Dorsal view and detail of embolus in left
spermathecal duct; B. Without embolus, dorsal view; C. With embolus, lateral view; D. Detail of embolus inside the atrium
of the left spermathecal duct, posterior view. E–J. Male, left palpal bulb: E. Retrolateral view; F. Prolateral view; G. Dorso-
prolateral view. H–J. Male, left palpal tibia: H. Retrolateral view; I. Prolateral view; J. ventral view; K. Detail of EC; L. Bulb
with EC broken, prolateral view; M. Embolus broken, dorso-prolateral view. Abbreviations and acronyms. Spermathecae: At,
spermathecal atrium; Du, spermathecal duct; R, receptaculum; SB, spermathecal base; SDR, spermathecal duct ring; Male palp
and bulb: AH, apical haematodocha; AK, apical keel; BH, basal haematodocha; Pa, patella; Ti, palpal tibia; Cy, cymbium; EA,
embolic aperture; EC, embolic crack; Em, embolus; MH, median haematodocha; PLoK, posterior locking keel; RTG, retrolateral
palpal tibia grooves; SD, seminal duct; SLoK, subapical locking keel; St, subtegulum; T, tegulum; TE, tibial excavation; TG,
tegular grooves; TS, ventro-retrolateral palpal tibia spines; TT, tegular teeth; VTG, ventral palpal tibia grooves. Scale bars: A–C,
H, J = 0.5 mm; D–G, K–M = 0.2 mm.

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FIGURE 7. A–D. Neocteniza toba Goloboff, 1987. A–D. Male palpal bulb, prolateral view: B. Detail of TT and EC; C. Detail of
PLoK; D. Detail of embolus tip. Abbreviations and acronyms. AH, apical haematodocha; AK, apical keel; EC, embolic crack;
MH, median haematodocha; PLoK, posterior locking keel; TG, tegular grooves; TT, tegular teeth.
Tarsi I–IV integral. All tarsi without scopula (Fig. 10A–D). Spinnerets short; PMS with ca. 50 spigots, basal article
of PLS with ca. 75, medial article with ca. 50, of them 10 larger on apex, apical article with ca. 35 larger on apex.
Spinnerets length: PMS 1.00, basal article of PLS 1.47, medial article 0.57, apical article 0.27, short, domed (Fig.
9E). Spermathecae fairly straight, just slightly curved, presenting membranous SB, bearing SDR, and with enlarged
receptacle (R) (Fig. 6A–D).
Trichobothria: Tibia: I, both rows 5; II, prolateral row 8, retrolateral 6; III, prolateral row 4, retrolateral 7; IV,
prolateral row 8, retrolateral 9; palp, prolateral row 7, retrolateral 10. Metatarsi: I, retrolateral row 12; II, retrolateral
17; III, retrolateral row 7; IV, retrolateral row 16. Tarsi: I, both rows 6; II, prolateral row 4, retrolateral 5; III, both
rows 5; IV, both rows 6; palp, prolateral row 4, retrolateral 5.
Spination: Femora: IV, 14 D ant ap, huddled. Patellae: III, 15/14 D ant ap, forming a triangle; IV, 12 D ant b,
huddled. Tibia: I, 6/9 R inf, 1 P; II, 1 R inf, 1 P; III, 2 D, 1/2 R ap, 4 P; IV, 1 P inf; palp, 1 D P. Metatarsi: I, row of
17 P D ant, 14/17 R D post; II, 14/13 P, 6/7 R, 9/7 D post; III, 9/10 P, 13 R; IV 6 P, 2 R ap. Palp, 9/8 P, 3/5 R. Tarsi:
I, 4/2 P, 4/3 R; II, 4 P, 3/4 R D; palp, 13/15 P, 11/10 R.
Male (CAD 789)
Color in life (Fig. 8D–F) and in ethanol (Fig. 11A–C): Carapace dark brown to black, abdomen slightly lighter.
Leg femora dark grayish to black, rest of the legs are brown. Entire palps dark grayish to black. Sternum, coxae and
trochanter light gray.

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FIGURE 8. A–F. Neocteniza toba Goloboff, 1987, live specimens. A–C. Female: A. Lateral-frontal view; B. Dorsal view; C.
Female internally bearing eggs, yet to be laid, lateral view; D. Female and male, dorsal view, note the strong sexual dimorphism,
male much smaller. E, F. Male: E. Dorsal view; F. Frontal view. Scale bars = 2 mm.
Total length 5.76. Carapace 2.34 long, 2.03 wide; cephalic region strongly convex, fovea deep, recurved (T-
shaped) (Fig. 11A). Anterior and posterior ocular row slightly recurved (Fig. 11A). Eye sizes: AME 0.11, ALE
0.16, PME 0.11, PLE 0.11. Eye interdistances: AME–ALE 0.07, PME–PLE 0.05, AME–AME 0.10, ALE–PLE
0.07, PME–PME 0.33. Eye tubercle 0.37 long, 0.74 wide. Chelicerae with rastellum formed by 3 thick, blunt spines

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on edge of cheliceral prolongation and two laterally adjacent spines outside cheliceral prolongation (Fig. 11D).
Intercheliceral tumescence absent. Cheliceral promarginal row with 6 teeth, 2 central teeth, 8 teeth on retromarginal
row. Labium 0.19 long, 0.44 wide, without cuspules. Palpal coxae without cuspules. Sternum 1.57 long, 1.10 wide.
Sigilla barely discernible (Fig. 11B). Palp measurements: trochanter 0.22/ femur 1.16/ patella 0.44/ tibia 1.22/
cymbium 0.73/ total 3.77. Leg measurements: I trochanter 0.55/ femur 2.12/ patella 0.82/ tibia 1.60/ metatarsus
1.55/ tarsus 0.79/ total 7.43; II 0.53/ 1.76/ 0.81/ 1.32/ 1.37/ 0.80/ 6.59; III 0.57/ 1.38/ 0.78/ 0.96/ 1.69/ 0.93/ 6.31; IV
0.62/ 2.37/ 0.96/ 2.21/ 2.64/ 1.18/ 9.98. ITC on all legs (Fig. 12B, C, F). Preening combs on all metatarsi, I, II, 2 VP;
III, 3 VP, 3 VR; IV, 3 VP, 3 VR. Integument with flattened scaly cuticle with highest distal ends on all legs (Fig. 12C,
D) and palp. Tarsi I–IV integral (Fig. 12B, E). All tarsi without scopula (Fig. 11F). Spinnerets short; PMS with ca.
8 spigots, basal article of PLS with ca. 14, medial article with ca. 11, apical article with ca. 12, of them 5 larger on
apex. Spinnerets length: PMS 0.20, basal article of PLS 0.42, medial article 0.23, apical article 0.16, short, domed
(Fig. 11E). Palpal tibia incrassate and with spines (Fig. 6H–J). Epigastric area with ca. 15 epiandrous spigots. Bulb
with large, slightly twisted tegulum and lanceolate, somewhat flattened embolus, presenting: Haematodochae BH,
MH, AH; Tegular grooves (TG); Tegular teeth (TT); keels PLoK, SLoK and AK; EC near or reaching the tegulum
(Fig. 6E–M; see Fig. 7D for AK in detail).
Trichobothria (Fig. 12D): Tibia: I, both rows 3; II, prolateral row 4, retrolateral 5; III, both rows 3; IV, both
rows 4; palp, prolateral row 3, retrolateral 4. Metatarsi: I, prolateral row 3, retrolateral 4; II, retrolateral row 5; III,
retrolateral row 3; IV, retrolateral row 6. Tarsi: I, prolateral row 2, retrolateral 3; II, prolateral row 1, retrolateral 2;
III, prolateral row 1, retrolateral 3; IV, prolateral row 2, retrolateral 3.
Spination: Femur: IV, 9/8 D–P sup ap, clustered. Patella: III, 5/ 6 D–P ap row and 5 D–P IV. Tibia: I and II, ca.
4/5 very thin V; III, 5 D. Metatarsus: I and II, ca. 6/7 very thin, V; III, 9 small, D. IV, 2 very thin D ap.
Variation: Females (n=10): total length 14.56–24.21; carapace 6.52–9.34 long; abdomen 7.40–10.54 long.
Palpal coxae with 0–3 blunt cuspules. Some individuals also with 2 VP preening combs on leg I. Males (n=6): total
length 5.23–6.38; carapace 2.28–2.58 long; abdomen 2.16–2.96 long. Palpal coxae without cuspules.
Distribution: Argentina (Goloboff 1987) and Paraguay (Goloboff & Platnick 1992); recorded for the first time
for Brazil, in semideciduous Atlantic Forest (Fig. 14) and in seasonal forest areas of the Cerrado (Brazilian savanna)
(Fig. 13).
Natural History and habitat. Specimens of N. toba were found in Cerrado areas in the São Paulo State, but
mostly in the São Paulo State University campus, in Rio Claro, in a small, somewhat ecotonous forested area (Fig.
14A, B), containing both Atlantic Forest and Cerrado plant species (G. Sabino pers. comm.). The specific site has
great anthropogenic influence, containing exotic plant species (G. Sabino pers. comm.) and being in close proximity
to buildings and deforested areas. N. toba co-occurs with other trapdoor spiders of the genera Idiops and Actinopus
Perty, 1833, although only Idiops were seen closely concentrated (from 4 cm to 5 m) in the same area of exposed
clay soil. A single Actinopus burrow was found near Neocteniza burrows. They also coexist with other mygalomorph
spiders, such as Rachias brachythelus (Mello-Leitão, 1937), Rachias sp., Psalistopoides fulvimanus Mello-Leitão,
1934 (Lucas & Indicatti 2006), Stenoterommata sp. (Pycnothelidae) (Ghirotto et al. 2021) and Acanthoscurria
gomesiana Mello-Leitão, 1923 (Theraphosidae) (Gonzalez-Filho et al. 2012) , Idiops camelus (Mello-Leitão, 1937)
(Idiopidae), Actinopus fractus Mello-Leitão, 1920 (Actinopodidae) (Miglio et al. 2020), that are more widely
distributed throughout the University campus and others areas in the state of São Paulo, e.g., Parque Estadual Morro
do Diabo (Fig. 14C, D) and Estação Ecológica de Itirapina (Ghirotto et al. 2021, fig. 7B). Most males were collected
by actively searching, or by using pitfall traps during the rainy season (from November to March). Two juveniles,
which were identified as subadult males by their incrassated palpal tibiae (Fig. 18G), matured in captivity. Juveniles
and females were found by actively searching for their trapdoor burrows on the forest floor. Trapdoors of Neocteniza
were observed to be close to each other in some areas, as near as 5–7 cm from one another. In dry months (June to
August), juveniles and females kept alive in the laboratory sealed the lid of their burrows with silk, probably aiding
in humidity conservation. Males were exposed to female burrows, but unfortunately no copulatory behavior was
observed. The adult male specimen collected died after a few days in captivity.
The burrow of a female of N. toba was meticulously analyzed. The burrow was slightly curved, with a depth
of 10 cm and a diameter of 8.2 mm at the entrance, 15.7 mm at 5 cm deep, 12.5 mm at 8 cm deep and 10 mm at the
bottom. The burrow had a bevelled entrance, with a widening of 4 mm deep and 13–8.5 mm (tapering from outside
to inside) in diameter for lid fitting, and the hinge was 9.7 mm, firmly connecting the lid and the burrow. The lid
was nearly as long as wide, and densely covered with plant material in small pieces, added in wide concentric layers

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originating from the hinge in its outer side (Fig. 15A–C). On the inner side, the lid was formed of wide concentric
silk layers similar to the outer side (Fig. 15B). The lid had an irregular edge with soil and plant material incorporated
(Fig. 15C). The inner surface of the burrow was covered with dense layers of sheet silk. The entrance of the examined
burrow had only clean silk layering (Fig. 15D), but another examined burrow of a juvenile female had waste (old
exuviae and prey exoskeleton remains, see below) pressed with silk against the wall near the entrance of the burrow,
but also older waste encrusted with soil. Interestingly, upon excavation of the soil contacting the burrow, we found
a wide lateral chamber, the Cemented Disposal Chamber (CDC, Fig. 16A–D) filled with the spider’s exuviae, old
silk coverings and shattered prey exoskeleton. This chamber was encrusted with soil, then covered with silk which
also forms the wall of the burrow (Fig. 16C). Prey exoskeleton remains in this burrow consisted of 30 ants and one
beetle. The ants were identified, consisting of 10 Pachycondyla sp., 2 Odontomachus sp., 15 Pheidole sp. and 3
Neoponera sp. (species that were also observed in the same forested area where the spider was found), and are all
stinging and predatory species (Schmidt & Shattuck 2014; Sarnat et al. 2015). Other burrows made by juveniles and
females were very similar in structure, differing mostly in size, according to the spider size; some lids were slightly
longer than wide, others slightly wider than long; very small individuals were found in simpler burrows, without
much detritus added, and without a conspicuously webbed layer at the burrow border. Another N. toba had similar
values of burrow measures: the lid was 12 mm in diameter, the burrow was 6 mm in diameter at the entrance, and 10
cm deep. We also found prey remains stored in the same manner: 2 Pachycondyla sp., 2 Pheidole sp., 3 Neoponera
sp. and one Diplopoda. The burrow of a very large N. toba female had the lid slightly longer than wide: 23 mm
long, 18 wide; the burrow border measured 23.32 mm long, 22.11 wide, and the burrow entrance measured 15.12
mm long, 14.50 wide. A N. toba juvenile with a body length of 4 mm, which was kept in captivity, built a burrow
with a 1.8 mm lid diameter in March 2018, which grew to 2.7 mm by October 2018, and finally to 4.62 mm by
September 2019, when the spider became an adult male. Only juvenile males were observed to construct trapdoors
in captivity, while adult males only dug holes with the entrance sparsely sealed with silk and irregularly spread soil.
In that way, only two burrows that belong to adult males could be examined—as only two males were obtained by
rearing since earlier development stages, when the male had constructed its trapdoor before maturing. Both burrows
were wider than long, and didn’t have a layer of silk or detritus bordering the burrow entrance, but the soil around
tapered slightly from the lid fitting aperture to a few millimeters above, apparently presenting a specific structural
pattern of funneling of the first few millimeters of the burrow. The male that matured in September 2019 had the lid
of its burrow measuring 3.59 mm long, 4.62 wide; the burrow border measuring 2.95 mm long, 3.58 wide; and the
burrow entrance measuring 3.52 mm long, 4.61 wide. The other captive reared male had its lid measuring 5.05 mm
long, 6.56 wide. For details on burrows of N. australis, N. minima and other populations of N. toba, see Goloboff
(1987).
A female was observed in the field inside its burrow and the entire burrow was removed and brought into the
laboratory without apparent structural damage. The piece of clod that held the burrow was kept in the laboratory,
the soil was sprayed with water each week, and the spider was fed each week with cockroaches. Seven months later,
the burrow was opened because spider activity could no longer be detected; on 02.x.2018 the female was found
dead in the bottom of the burrow but, surprisingly, with a healthy egg sac that was immediately opened to reveal
132 nymphs, 1 withered egg and 3 unhatched eggs. The egg sac was oval (13 mm x 7 mm) at the central part, and
anchored to the walls of the burrow by two thick threads of silk originated from the upper part and two thick threads
between the bottom part of the egg sac and the burrow wall. It was located ca. 5 cm from the burrow opening (Fig.
17A–D). The nymphs were kept together and became first instars (first molt on 27.x.2018), and a few days later
some nymphs (around ten) dug their own burrows in the small container (31 mm diameter x 73 mm deep, with 40
mm of soil). The first instar spiderlings kept together covered their container with a dense silk layer (Fig. 18A–F).
When handled or placed on a stick, almost all spiderlings were observed to drop from the object while anchored
to it by a silk thread, with their abdomen bent anteriorly, exposing and extending the spinnerets (Fig. 19A–D).
When reaching 20–30 cm from the dropping point, they were carried away by soft breezes. This dropping behavior
seemed to be triggered by soft wind or substrate vibration. At some distance, the silk thread detached from the
dropping point and the spiderlings became airborne, with the silk thread attached to their spinnerets (Fig. 19C–E).
When no breeze or wind was present, the spiderlings were seen gliding, and the silk threads spread in many parallel
fibers (Fig. 19C–F). Some individuals reached distances of 3 meters from the dropping point, inside the laboratory,
without wind, representing the first idiopid spiders witnessed performing ballooning. Older spiderlings, especially
those that had already eaten, had a lower tendency to perform ballooning.

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FIGURE 9. A–E. Neocteniza toba Goloboff, 1987, female. A, B. Prosoma: A. Dorsal view; B. Ventral view; C. Opisthosoma,
dorsal view; D. Chelicerae, ventral view; E. Spinnerets, ventral view. Scale bars: A–C = 2 mm; D, E = 1 mm.

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FIGURE 10. A–D. Neocteniza toba Goloboff, 1987, female metatarsi and tarsi, prolateral view: A. Right leg I; B. Right leg II;
C. Left leg III; D. Left leg IV; Arrows showing preening combs. Scale bars = 0.5 mm. Acronyms. STC, superior tarsal claws;
ITC, inferior tarsal claw.
We were able to observe one successful ant predation. A live Pheidole sp. (ant) worker was allowed to wander
in the container of a very small juvenile spider that had already built a trapdoor. The third time that the ant wandered
near the burrow lid, the spider suddenly and rapidly grabbed the ant and returned into the burrow, quickly closing
the lid.
We were able to indirectly expose a single live male of N. toba to a small to medium sized female (14 mm in body
length) from the University campus, by placing the male in the container with the female, where she had constructed
a burrow. We then recorded courtship behavior. The male performed three variations of body vibration (sensu
Ferretti et al. 2013a), one of them was especially distinct from the general patterns exhibited by most mygalomorph
species whose sexual behaviors are known. This distinct body vibration consisted of a discrete irregular movement
of the body and some legs, recoiling or moving slightly backwards and forwards to prepare for an intense, very high
frequency vibration of the body forward and backward, with minimal amplitude, during 1–4 seconds, similar to a
guitar string vibration, then slowing down in frequency or abruptly ending in an apparently release of tension by a
sudden discrete downward movement of the abdomen and palps and slight relaxation of the legs. This vibration was
performed at a much higher frequency than standard body vibrations. The other two variations of body vibration
were similar to the behavior exhibited by most Mygalomorphae: walking rhythmically with body vibrations,
and performing a standard body vibration, with slower frequency and similar to shaking the body forward and
backward. Once placed in the female’s container, the male started to walk, soon reaching the burrow lid with his
anterior tarsi. Immediately after contacting the lid, the male positioned himself in front of the burrow with his
right anterior tarsi touching the lid, then stopped briefly to start performing courtship behavior. All behaviors were
preceded and followed by pauses. The male performed six high-frequency body vibrations, then recoiled the legs,
and turned the other way, walking rhythmically along with standard body vibration, before performing another

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FIGURE 11. A–F. Neocteniza toba Goloboff, 1987, male. A, B. Prosoma: A. Dorsal view; B. Ventral view; C. Opisthosoma,
dorsal view; D. Chelicerae, ventral view; E. Spinnerets, ventral view; F. Left leg I, retrolateral view. Acronym. CG, cheliceral
grooves. Scale bars: A–C = 1mm; D–F = 0.5 mm.

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FIGURE 12. A–F. Neocteniza toba Goloboff, 1987, male. A–D. Leg I, retrolateral view: A–C Detail of tarsi and claws; D.
Detail of integument. E–F. Tarsi IV; F. Detail of tarsi and claws. Abbreviations and acronyms. Bt, bothrium; FT, filiform
trichobothrium; It, integument with scaly cuticle; ITC, inferior tarsal claw; STC, superior tarsal claw; TO, tarsal organ.
high-frequency body vibration, followed by a standard slower frequency body vibration, also raising and lowering
the palps. Then, the male walked rhythmically with standard body vibration, before unfortunately being attacked
and killed by the female, who suddenly dragged the male inside her burrow. The entire encounter lasted 8 minutes.
Discussion
Diet traits. The remains found in the burrows of N. toba specimens were predominantly those of predatory stinging

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ants, suggesting that these spiders may specialize in hunting them. How the spiders manage to deal with such
dangerous prey is an interesting question for ecological, behavior and toxicological studies. These spiders have
very large cheliceral teeth (Figs 2D, 9D) that could be adapted to ant predation, as the action of the strong cheliceral
muscles could quickly neutralize the ant, crushing it between the fangs and the large cheliceral teeth. This indicate
that the strong condition of the retromarginal cheliceral teeth could be an apomorphic trait in Neocteniza as discussed
by Raven (1985), and possibly related to diet specialization. The trapdoor may also play an important role in
predation, as it protects most of the soft parts of the spider’s body, like the abdomen and dorsal and ventral surfaces
of the cephalothorax, leaving only the front of the spider exposed. This method of myrmecophagy, involving a
trapdoor, appears to be unique within spiders (Pekár 2004).
FIGURE 13. Map showing the distribution of Neocteniza species.

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FIGURE 14. A–D. Collecting site of Neocteniza toba Goloboff, 1987, São Paulo state, Brazil, all from Atlantic Forest: A.
Universidade Estadual Paulista, Rio Claro; B. Instituto de Biotecnologia, Universidade de Araraquara, Araraquara; C, D. Parque
Estadual Morro do Diabo, Teodoro Sampaio; C. General view. D. Detail.
Distribution and behavior. Although Neocteniza is widely distributed, according to the updated distribution
map (Fig. 13), we observe distributional gaps in several regions in South and Central America, such as Nicaragua,
Chile, almost all of Colombia and Peru, and vast areas in Brazil. This might reflect the relative difficulty in collecting
Neocteniza specimens, as they are small trapdoor spiders and may have low density populations and/or strong
microhabitat preferences. For example, they were not found or were uncommon in other better sampled areas where
Idiops, Actinopus and several Pycnothelidae representatives were present (e.g., Bonaldo et al. 2009; Ferretti et al.
2010; Lemos et al. 2012; Indicatti 2013; Indicatti et al. 2017: 454), as was evident both in our sampling effort and
in museum collections examined by the authors. Neocteniza specimens are rare in scientific collections (Platnick &
Shadab 1981). Moreover, males could be biased, over represented in collections, as they are easily caught in pitfall
traps while wandering to find a female, as occurs with other mygalomorphs (Costa & Pérez-Miles 2002; Gonzalez-
Filho et al. 2012). In order to successfully find juveniles and females, one has to look carefully for their trapdoors,
sweeping the soil or trail ravines.
Our new records reveal that the geographical distribution of the three species herein studied is much wider
than previously known, extending the known distribution towards the east for each respective species. Ballooning
could be in part responsible for these wider distributions, as it greatly increases the dispersal capacities of an
individual (Fisher et al. 2014). This behavior is mainly associated with newly hatched spiders or small species (Bell
et al. 2005), and has already been described for mygalomorph spiders of the families Actinopodidae (Ferretti et al.
2013b), Atypidae (Bristowe 1939; Coyle 1983; Coyle et al. 1985) and Halonoproctidae (Coyle 1985; Coyle et al.
1985; Eberhard 2006). It could be hypothesized that, when ballooning takes place, it may represent the primary
dispersal mode for newly hatched spiders (Bond & Coyle 1995). In the case herein described (N. toba), only a few
adult burrows were found close together. As seen in captivity, as soon as juveniles disperse, they start digging their

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own burrows. However, we did not find small burrows around the burrows of females, suggesting that ballooning
may be the primary mode of dispersal for spiderlings of N. toba, as well.
FIGURE 15. A–D. Neocteniza toba Goloboff, 1987, burrows: A. Arrows showing closed burrows (with leaf litter removed);
B. Opened burrows; C. Burrow lid with attached leaf litter; D. Excavated burrow showing a spider, note cover with concentric
silk layers. Scale bars = 10 mm.
Stridulatory apparatus. Apart from the well-known stridulatory setae that produce audible sound found in large
theraphosids of both sexes (Uetz & Stratton 1982; Galleti-Lima & Guadanucci 2019), males of several mygalomorph
lineages bear structures that are considered stridulatory, in which the friction between them may produce a sound
or vibration that cannot be heard by humans. The function of this supposed stridulation could be defensive and/or
for sexual communication (Marshall et al. 1995). Five areas, each where two grooved regions can make contact,
were identified: 1) cheliceral grooves (CG) with male palpal femora (known in several Araneomorphae families,

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e.g., Linyphiidae (Hormiga 1994; Draney & Buckle 2005), Zodariidae (Jocqué 2005, fig. 4)) (Figs 3A, 5D); 2)
tegular grooves (TG) with ventral palpal tibia grooves (VTG); 3) tegular teeth (TT) with VTG (possibly touching
the female) (Figs 1E–H, 4A–C, G, 6E–G); 4) retrolateral palpal tibia grooves (RTG) with leg I femora (Figs 1H,
4D, 6H); 5) ventro-retrolateral tibial spines (TS) with leg I femora (possibly touching the female) (Figs 1H, 4F, 6J)
The structures TG, VTG and RTG are evident in most Mygalomorphae families (R.P. Indicatti pers. observation).
Currently, there are only three studies that mention TG and VTG regions as possibly stridulatory apparatus (Wishart
2006, fig. 3H, I (only for the tibial excavation of Arbanitis L. Koch, 1874); Indicatti et al. 2015; Indicatti et al. 2017,
figs 8, 13, 17, 20, 23), but it can also be seen in images from different studies (e.g., Schiapelli & Gerschman 1967,
figs 10, 15; Wishart 1992, figs 28–37; Indicatti et al. 2008, fig. 2B; Gonzalez-Filho et al. 2012, figs 2a,b; Ferretti
et al. 2017, fig. 5a–c; Fonseca-Ferreira et al. 2017, figs 5, 34), in which some are very pronounced (e.g., Schiapelli
& Gerschman 1967, figs 10, 15; Indicatti et al. 2008, fig. 2B, F, K). In Neocteniza, the TG region is particularly
enlarged due to the expansion of the medium haematodocha (MH).
FIGURE 16. A–D. Neocteniza toba Goloboff, 1987, female burrow. A, B. Chamber covered with silk layer forming the burrow
wall and disposal cemented chamber; C. Chamber edge detail with prey remains; D. End of chamber of a different burrow with
prey remains and spider exoskeleton. Abbreviations and acronyms. Cha, chamber; CDC, cemented disposal chamber. Scale
bar = 10 mm.
The genitalia of Neocteniza. As discussed by Goloboff (1987), the Neocteniza male genitalia are quite unique.
Besides exhibiting two distinct patterns that delimit the existing species groups (australis and sclateri), they are the
only Mygalomorphae that are known to naturally break the embolus once it is coupled with the duct of the female’s
spermathecae. The embolus breaks at the EC (Figs 1E–H, 4A, C, H, 6F, G, K–M, 7B), which differs greatly in
relative position between the groups. In the australis group, the EC could be observed in already broken bulbs (Fig.
6L, M), as well as in complete, identifiable emboli inside the spermathecae of females (Fig. 6A, C, D). The exact
delimitation of the EC in the sclateri group also could be observed inside the spermathecae (Fig. 1D), but we did not
find any broken bulbs for specimens of this group. A contraction of the haematodocha, similar to that experienced by
some preserved individuals, probably causes the break inside the spermathecae (Goloboff 1987). The haematodochae

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can be divided into basal (BH), median (MH) and apical (AH); the latter two being well developed in the genus. The
AH, in contact with the EC, is most probable the haematodocha responsible for the embolus detachment (Fig. 7A,
B). The embolus break could be achieved by the contact of the paraembolic apophysis (PA) (Fig. 1E–G) with the
strongly sclerotized spermathecal base (SB) (Fig. 1A–D) facilitating the embolus anchoring by the AK. On the other
hand, the ALoK, PLoK, SLoK and AK probably aid in anchorage of the embolus in the female’s SDR (australis
group), as well as AK anchorage in IDu (sclateri group), respectively, which are specifically sclerotized areas of the
spermathecae.
FIGURE 17. A–D. Neocteniza toba Goloboff, 1987. A–C. Egg sac anchored on the lateral part of the burrow: A. Lateral view;
B. Inferior view; C. Superior view; D. Internal area that touches the chamber wall. Scale bars = 10 mm.

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FIGURE 18. A–G. Neocteniza toba Goloboff, 1987. Immature stages: A, C. Spiderlings in nymphal stage in the egg sac; C.
Spiderlings in nymphal stage; D. First instar spiderling in ethanol, ventral view; E. Detail of spiderling spinnerets; F. First instar
spiderling; G. Sub-adult male with thickened palpal tibia. Scale bars: A, B = 2 mm; C, D, F, G = 1 mm; E = 0.2 mm. Acronyms.
PMS, posterior median spinnerets; PLS, posterior lateral spinnerets.

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FIGURE 19. A–F. Laboratory ballooning experiment with Neocteniza toba Goloboff, 1987 first instar spiderlings: A. Specimen
walking on silk threads spun by it and other individuals; B. A spiderling dropping itself anchored by silk threads, with bent
abdomen and exposed spinnerets; C–F. Individual gliding. The spiderlings became airborne, with the silk thread attached to
their spinnerets: E. The arrow indicates the anterior portion of the set of silk threads used for ballooning (only the focused area
is visible); F. Posterior portion of silk threads used for ballooning (only the focused area is visible). Scale bars: A–D = 2 mm; E
= 5 mm; F = 100 mm.

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FIGURE 20. A–N. Morphology of male palp among all subfamilies of Idiopidae, left palp: A, D, G. Arbanitinae, Arbanitis sp.,
ventral, retrolateral and prolateral view, respectively; B, E, H. Idiopinae, Idiops camelus (Mello-Leitão, 1937), ventral, retrolateral
and prolateral view, respectively; C, F, I, J–N. Genysinae: C, F, I. Neocteniza toba Goloboff, 1987, ventral, retrolateral and
prolateral view, respectively; J. Genysa bicalcarata Simon, 1889b, retrolateral view, adapted from Simon (1903); K. Hiboka
geayi Fage, 1922, retrolateral view, adapted from Fage (1922); L–N: Scalidognathus radialis (O. Pickard-Cambridge, 1869):
L. Retrolateral view; M, N: Palpal bulb, ventral and prolateral view, adapted from Raven (1985). Acronyms. RLC, retrolateral
lobe of cymbium; PLC, prolateral lobe of cymbium. Scale bars: A–I = 0.5 mm; J–N = not to scale.
The relationships of Neocteniza. The family Idiopidae presents three synapomorphies as proposed by Raven
(1985): I) male palpal bulb with a distal haematodocha extending ventrally almost to the embolus (thus transforming
the distal sclerite into an open scoop rather than a cone); II) bilobed cymbium with a broad and an acutely pointed
lobe (Figs 20A, B, D, E, G, H, 21A, B); III) male palpal tibia with a ventro-prolateral excavation bordered by spines
(Fig. 20). The family has three subfamilies: Arbanitinae Simon, 1903, Idiopinae Simon, 1889a and Genysinae
Simon, 1903 (Raven 1985). Neocteniza comprises a unique lineage standing out from all other mygalomorphs,
including those from the same family and subfamily. Raven (1985) found uncertainty when trying to establish
its relationships with other groups. Together with the poorly known genera Genysa Simon, 1889b, Hiboka Fage,
1922 (both from Madagascar) and Scalidognathus Karsch, 1892 (India and Sri Lanka), Raven (1985) included
Neocteniza in Genysinae. Interestingly, Raven (1985) was at the time unable to ascertain whether Neocteniza had
angular cymbium lobes, a synapomorphy for Idiopidae. We could not confirm the presence of this synapomorphy
for the genus, as the prolateral lobe, seen in Figs 20A, B, D, E, G, H and 21A, B for Arbanitinae and Idiopinae

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appears to be absent or at least strikingly different in Neocteniza. In N. toba, there is a direct connection to the basal
haematodocha from the prolateral side of the cymbium that does not form a distinct lobe (Figs 20C, F, I, 21C–E).
Significantly, N. minima presents a discrete prominence in the prolateral side of the cymbium, but does not form a
distinct, protruding lobe as for other Idiopidae (Fig. 21F, G). The prolateral side of the cymbium in the examined
Neocteniza specimens has a significant less sclerotized portion, resembling a membrane, presenting some setae as
other regions of the cymbium (Fig. 21C, see white arrow), before directly connecting to the basal haematodocha. It
is hard to access if this is a derived state of the prolateral lobe as present in other Idiopidae or if Neocteniza doesn’t
possess that synapomorphy at all. As a practical example of Neocteniza uniqueness even among Genysinae, in
Raven (1985), Neocteniza is one of the most cited genera in the text of that work and is even more cited than its own
family (with 84 citations, contra 18 of Genysa, 10 of Hiboka, 23 of Scalidognathus, 28 of Idiops, 23 of Arbanitis,
31 of Genysinae, 75 of Idiopidae). The Genysinae are distinguished from Idiopinae and Arbanitinae mainly by two
synapomorphies, the eyes widely arranged and the broad recurved fovea (Raven 1985). Genysinae representatives
additionally have a spine comb in patella III, although this is also present in some Idiops species. Neocteniza bears
several singular characteristics, combining character states from different mygalomorph lineages (Raven 1985).
The legs of both sexes are unusual for Genysinae, with females bearing several spines on metatarsi and tarsi of
anterior legs, combined with preening combs on all legs, and males lacking a tibial spur on leg I. Neocteniza
further lack clavate trichobothria on all tarsi and palps, a feature present in Scalidognathus (Sanap & Mirza 2011).
The lack of knowledge and modern descriptions on all the other Genysinae probably was a problem for Raven
(1985) and currently still halts a more substantial analysis regarding Neocteniza relationship with other genera.
Even so, it’s already evident that Neocteniza differs significantly from supposedly related groups. The integument
of Neocteniza presents shorter, flattened and apicallly elevated cuticular scales on all legs (Figs 12A–F, 23F) and
palps (similar to Pseudonemesia spp., Microstigmatidae, as abovementioned in descriptions), differing from at
least five other Idiopidae genera (Fig. 23A–E). In comparison to other Genysinae, and in several cases to all other
Idiopidae (see Fage 1922; Raven 1985; Sanap & Mirza 2011; Ferretti et al. 2017; Fonseca-Ferreira et al. 2017),
Neocteniza species presents weaker, straighter fovea, eyes more widely arranged, reduced dentition of paired tarsal
claws, lack of tarsal scopulae, a diamond-shaped cephalothorax with enlarged anterior region specially in females,
strongly spinose anterior tarsi in females, and very different genitalia in both sexes. The genitalia are different to
that of remaining Genysinae (Figs 20C, F, I–N, 22A–G; see also Raven 1985; Sanap & Mirza 2011 for comparison),
of all other Idiopidae (Figs 20A–I, 22A–F; see also Main 1985; Raven 1985; Ferretti et al. 2017; Fonseca-Ferreira
et al. 2017; Rix et al. 2017; Wilson et al. 2020 for comparison), and of other mygalomorph families (Raven 1985;
Schiapelli & Gerschman 1967; Raven & Platnick 1981; Lucas & Indicatti 2006; Ríos-Tamayo & Goloboff 2012;
Gonzalez-Filho et al. 2012; Fonseca-Ferreira et al. 2017; Indicatti et al. 2017; Zonstein 2018; Miglio et al. 2020).
Neocteniza males have a shorter and thicker palpal tibia (Figs 1J, 4D, 20C, F, I), palpal bulb with no fused sclerites
allowing the expansion of the median and apical haematodochae (Fig. 4B, C vs 4I), a detachable embolus (Fig.
4I, J). Females have a spermathecae with a distinct sclerotized circular band conjoined with membrane, the ring
in australis group (SDR) (Figs 6A–C, 22D, E) or the inferior duct in sclateri group (IDu) (Figs 1A–C, 22F). In
Idiopinae and Arbanitinae the spermathecae does not form sclerotized bands (Fig. 22A–C; see also figs 187–208 in
Main 1985), and other Genysinae appear to have a different sclerotization pattern and to not present a conspicuous
and large membranous apex (Fig. 22G). Additionally, the strong size sexual dimorphism (Fig. 8D; Goloboff 1987,
fig. 1) is apparently exclusive of Neocteniza among more related lineages, differing from all other Idiopidae and
several other mygalomorph families. We recommend Raven (1985) for a more detailed consideration on Neocteniza
relationships. Modern descriptions and revisions of other Genysinae, together with the inclusion of Neocteniza and
other Genysinae in molecular phylogenetic analysis, would help to solve the enigmatic position of these taxa. We
believe that either Neocteniza or Neocteniza + other Genysinae could represent an independent lineage, forming a
separated clade from other idiopids, i.e., a possible new subfamily.

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FIGURE 21. A–G. Morphology of male cymbium among all subfamilies of Idiopidae, left palp: A. Arbanitinae, Arbanitis sp.,
ventral view; B. Idiopinae, Idiops camelus (Mello-Leitão, 1937), ventral view; C–G. Genysinae: C–E. Neocteniza toba Goloboff,
1987: C. Ventral view; D. Ventro-prolateral; E. Dorsal view; F–H. Neocteniza minima Goloboff, 1987: F. Dorsal view; G.
Ventral view. Arrows: Thinner yellow arrows indicate basal region of retrolateral lobe, thicker black arrows indicate apical
region of retrolateral lobe, medium white arrows indicate prolateral lobe. Scale bars: A, B = 0.5 mm; C–F, G = 0.2 mm.
FIGURE 22. A–G. Morphology of spermathecae among all subfamilies of Idiopidae, dorsal view: A, B. Arbanitinae:
A. Cataxia pulleinei (Rainbow, 1914); B. Arbanitis rapax (Karsch, 1878), adapted from Wishart (1992); C. Idiopinae, Idiops
camelus (Mello-Leitão, 1937); D–G. Genysinae: D. Neocteniza toba Goloboff, 1987; E. Neocteniza australis Goloboff, 1987,
adapted from Goloboff (1987); F. Neocteniza minima Goloboff, 1987; G. Scalidognathus tigerinus Sanap & Mirza, 2011, adapted
from Sanap & Mirza (2011). Abbreviations and acronyms. Du, spermathecal duct; Em, embolus; IDu, inferior spermathecal
duct; R, receptaculum; SDR, spermathecal duct ring; SDu, superior spermathecal duct. Scale bars: A–F = 0.5 mm.

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FIGURE 23. A–F. Morphology of integument among all subfamilies of Idiopidae: A–D. Arbanitinae: A. Arbanitis sp., female
tarsus I; B. Cataxia sp., female tarsus I (QMS 19413, stub 35 CAD); C. Euoplos variabilis (Rainbow & Pulleine, 1918), male
tarsus IV; D. Idiosoma sp., female tarsus I; E. Idiopinae, Idiops sp., male metatarsus I; F. Genysinae, Neocteniza toba Goloboff,
1987, male tarsus I.
Acknowledgments
We thank all the curators, Antonio A. Lise (MCTP) and Cristian Grismado (MACN) for the loan of specimens or for
providing a repository for material; Robert J. Raven (Queensland Museum (QM), South Brisbane) for the donation

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of Australian Arbanitinae specimens used in this work and for the use of SEM at the QM. We thank Emílio Calvo
for aiding in fieldwork, Everton F. Trova for helpful comments within the manuscript, Gabriel Sabino for aiding
in environment characterization, and Rony P.S. Almeida for helping with ant identification. Mônika Iamonte and
Antônio Yabuki for the assistance with SEM images in the Laboratório de Microscopia Eletrônica, UNESP. We also
thank Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) and Comissão Técnico-Científica do
Instituto Florestal (COTEC) for issuing the permits for the collecting localities. Funding was provided by Fundação
de Amparo à Pesquisa do Estado de São Paulo (99/05446–8, RPI, JPL; 03/05487-3, RPI; 10/08459-4, Maria E.
Bichuette (MEB) (LES, São Carlos); 12/18287-1, RPI; 15/06406-4, 17/11985-9, JPLG, RPI), Conselho Nacional
de Desenvolvimento Científico e Tecnológico (141062/2007-0, RPI/PPGBA-Universidade Federal Rural do Rio de
Janeiro; 303715/2011-1, MEB; 479377/2012-0, JPLG, RPI; 308557/2014-0, 310378/2017-6, MEB; 168493/2017-
9, AGL). This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior -
Brasil (CAPES) - Finance Code 001. Fundação Grupo Boticário de Proteção à Natureza for the project “Diagnóstico
Ambiental da Área de Influência e Ambientes Subterrâneos do Sistema Cárstico da Gruna da Tarimba (Mambaí-
GO): Proposta Para a Delimitação de Unidade de Conservação de Proteção Integral” (#0941-20121). We are also
thankful for Robert J. Raven (QM) and Jeremy D. Wilson (MACN) for reviewing and helping to improve the
manuscript.
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