Casas & Rheims 2023 Zoo 5360 1 43

Zootaxa 5360 (1): 001–043 ISSN 1175-5326 (print edition)
https://www.mapress.com/zt/
Copyright © 2023 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5360.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:C97810F4-A6EB-4040-8DC4-0BBCD687E9AF
Nativus gen. nov., a new huntsman spider genus from South America (Araneae:
Sparassidae: Heteropodinae)
CRISTIAN M. CASAS1 & CRISTINA A. RHEIMS2
1
Grupo de Investigación Biodiversidad del Caribe Colombiano, Semillero de Investigación Sistemática de Artrópodos Neotropicales,
Departamento de Biología, Universidad del Atlántico, Barranquilla, Colombia.
� ccasas@est.uniatlantico.edu.co; https://orcid.org/0000-0002-0507-264X
2
Laboratório de Coleções Zoológicas, Instituto Butantan, Av. Vital Brasil, 1500, 05503-900, São Paulo, SP, Brazil.
� carheims@gmail.com; https://orcid.org/0000‑0003‑4418‑0552
Abstract
In this paper, the new genus, Nativus gen. nov. is proposed to include the type species, Nativus tupana sp. nov. (♂♀)
from Brazil and nine new species: N. carare sp. nov. (♂), N. hazzii sp. nov. (♂♀), N. janai sp. nov. (♂), N. nocaima sp.
nov. (♂♀), N. tawu sp. nov. (♂♀) and N. yurupari sp. nov. (♂♀) from Colombia, N. juruti sp. nov. (♂) and N. mariua sp.
nov. (♂♀) from Brazil, and N. napo sp. nov. (♂♀) from Peru. The genus is placed in the subfamily Heteropodinae due to
the presence of intermarginal denticles on the chelicerae, a long-toothed female palpal claw, two recurved eye rows with
median eyes smaller than laterals, and only one lateral spine on each side of metatarsi I−II, as well as a ventral branch
on the RTA (vRTA) in the male palp. Within Heteropodinae, it seems to be more closely related to Guadana Rheims and
Sparianthina Banks due to the presence of intermarginal denticles along the entire cheliceral groove and a dorsal tegular
apophysis (DTA) in the male palp. All species are described and illustrated. An identification key and distribution maps
are provided for all species of the genus.
Key words: Taxonomy, Amazon, Brazil, Colombia, Peru
Introduction
Heteropodinae Thorell, 1873 is a subfamily of Sparassidae Bertkau, 1872, characterized by the presence of
intermarginal denticles in the chelicerae, a long toothed female palpal claw, and two recurved eye rows with median
eyes smaller than laterals (Jäger 1998), as well as a ventral branch of the RTA (vRTA) in the male palp and only
one lateral spine on each side of metatarsi I−II (Rheims 2010a). Jäger et al. (2009) restricted the subfamily to the
Asian and Australian genera in what was called Heteropodinae s.str. However, the authors admitted to an extended
Heteropodinae (Heteropodinae s.l.), including Heteropodinae s.str., the African genus Berlandia Lessert, 1921
and the South American genera Anaptomecus Simon, 1903, Caayguara Rheims, 2010b, Guadana Rheims, 2010a,
Sparianthina Banks, 1929 and Sadala Simon, 1880.
Moradmand et al. (2014) provided a summary of the classification of Heteropodinae including thirteen
genera, nine in Heteropodinae s.str (Barylestis Simon, 1909, Bhutaniella Jäger, 2000, Heteropoda Latreille, 1804,
Martensopoda Jäger, 2006, Pandercetes L. Koch, 1875, Pseudopoda Jäger, 2000, Sinopoda Jäger, 1999, Spariolenus
Simon, 1880 and Yiinthi Davies, 1994) that together with four additional genera (Anaptomecus, Caayguara, Guadana
and Sparianthina) comprised Heteropodinae s.l.. The analysis carried out in that study, recovered a monophyletic
Heteropodinae s.str. but no representatives of the four Heteropodinae s.l. were included in the analysis. Pinto &
Rheims (2016) and Rheims (2021) argued that Anaptomecus, Guadana and Sparianthina should be considered true
Heteropodinae because they show all characters considered diagnostic for the subfamily, but Caayguara should be
excluded, because it lacks the characteristic long-toothed female palpal claw and recurved eye rows observed in the
other genera.
Accepted by C. Muster: 28 Sept. 2023; published: 25 Oct. 2023

With the recent description of Borniella Grall & Jäger, 2022, Martensikara Jäger, 2021, Menarik Grall & Jäger,
2022, Micropoda Grall & Jäger, 2022, Platnickopoda Jäger, 2020a, Thunberga Jäger 2020b and Tiomaniella Grall
& Jäger, 2022, the current composition of Heteropodinae includes 708 species in twenty genera (World Spider
Catalog 2023). Nevertheless, despite the generic diversity, over 80% of the known species belong to only three
genera: Heteropoda (189 spp.), Pseudopoda (252 spp.) and Sinopoda (141 spp.), distributed mostly in the Australian
and Oriental regions. From the Neotropical region only 29 species are known, six from the genus Anaptomecus,
twelve from Guadana and eleven from Sparianthina (World Spider Catalog 2023).
In this paper we propose the new genus, Nativus gen. nov., including the description of ten new species from
Brazil, Colombia and Peru. The new genus exhibits all the distinctive characters of Heteropodinae, and is therefore
considered a member of the subfamily. In addition, we provide an identification key and distribution maps for all
species of the genus.
Material and methods
Format of descriptions follows Rheims (2010a). Species are listed in alphabetical order. Only characters that differ
from the generic pattern are mentioned in species descriptions. Leg spination pattern is expressed according to
Petrunkevitch (1925). The material was examined under a LEICA MZ12.5 and a LEICA S8APO. All measurements
are in millimeters. Leg measurements are listed as: total length (femur, patella, tibia, metatarsus, tarsus); eye diameters
as AME, ALE, PME, PLE and interdistances as AME-AME, AME-ALE, PME-PME, PME-PLE, AME-PME, ALE-
PLE. Positions of tegular appendages are given according to clock positions, based on the left palp in ventral view.
Female epigynes were dissected and immersed in clove oil for better visualization of internal structures, following
Levi (1965), or submerged in a solution of pancreatin for 24 hours, following Álvarez-Padilla & Hormiga (2007).
Illustrations were made using a LEICA MZ 165 C stereomicroscope, with camera-lucida. Left male palps
were illustrated in prolateral, ventral and retrolateral views, epigynes in ventral view and vulva in dorsal view.
Photographs of genital structures and specimens were taken using a Leica DFC 500 digital camera, mounted on a
Leica MZ 16A stereomicroscope. Extended focal range images were composed using the program Leica Application
Suite version 2.5.0. Scanning electron micrographs (SEM) were taken using a scanning electron microscope of the
Laboratório de Biologia Celular (Butantan Institute). Material used for SEM was dehydrated through a series of
graded ethanol (80% to 100%), mounted on metal stubs and sputter coated with gold. In schematic courses of female
internal duct system, copulatory openings are marked with a circle and the end of the fertilization duct in direction
of uterus externus with an arrow. Geographical coordinates of collection localities were obtained from the labels
(given in round brackets) or from Google Earth (given in square brackets). Distribution maps were created using a
geographic information system (QGIS Team & Quantum, 2023; version 3.16.3, http://www.qgis.org/es/site/).
Abbreviations used throughout the text: ALE—anterior lateral eyes; ALS—anterior lateral spinnerets; AME—
anterior median eyes; bp—projection at base of embolus; C—conductor; CD—copulatory duct; CO—copulatory
opening; COd—copulatory opening depression; d—dorsal; dRTA—dorsal branch of RTA; DTA—dorsal tegular
apophysis; E—embolus; EF—epigynal field; FD—fertilization duct; GP—glandular projection; LL—lateral lobe;
MS—median septum; PLE—posterior lateral eyes; PLS—posterior lateral spinnerets; PME—posterior median
eyes; p—prolateral; PMS—posterior median spinnerets; PTA—prolateral tibial apophysis; r—retrolateral; RdP—
retrodistal tegular protrusion; RpP—retroproximal cymbial projection; RTA—retrolateral tibial apophysis; vRTA—
ventral branch of RTA; v—ventral; VTA—ventral tibial apophysis. Collection acronyms (curator in parenthesis):
AMNH—American Museum of Natural History, New York, USA (L. Prendini); IAvH-I— Instituto Alexander Von
Humboldt, Villa de Leyva, Boyacá, Colombia (J. C. Neita); IBSP—Instituto Butantan, São Paulo, Brazil (A.D.
Brescovit); ICN-Ar—Instituto de Ciencias Natrulares, Universidad Nacional de Colombia, Bogotá, Colombia (E.
Florez); INPA—Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (M. Oliveira); MPEG—Museu
Paraense Emilio Goeldi, Belém, Brazil (A.B. Bonaldo); MPUJ_ENT—Colección Entomológica del Museo
Javeriano de Historia Natural Lorenzo Uribe, Bogotá, Colombia (G. Fagua); MUSM—Museo de Historia Natural,
Universidad Nacional Mayor de San Marcos, Lima, Peru (D. Silva); MZSP—Museu de Zoologia, Universidade de
São Paulo, São Paulo, Brazil (R. Pinto da Rocha).
· Zootaxa 5360 (1) © 2023 Magnolia Press Casas & Rheims

Taxonomy
Nativus gen. nov.
Etymology. The generic name honors all indigenous communities from the great Amazon rainforest as most of the
described species herein are distributed in the region. This also commemorates the native Amazonians’ key role in
protecting Earth’s biodiversity; gender is masculine.
Type species. Nativus tupana sp. nov.
Diagnosis. Species of Nativus gen. nov. resemble those of Guadana Rheims and Sparianthina Banks in having
intermarginal denticles along the entire cheliceral groove, two pairs of ventral spines on tibiae I−II and one lateral
spine on each side on metatarsi I−II (Fig. 1), male palps with strong DTA (e.g. Figs 16−17, 106−107) and female
palp with a long-toothed claw (Fig. 3). They further resemble Guadana in having a vRTA, DTA acutely tapering
(e.g. Figs 29, 48, 92) (vRTA absent and DTA roundly blunt or distally widened in Sparianthina) and female vulva
with GP absent and FD short (e.g. Figs 32, 63, 78) (GP present and FD long in Sparianthina). They are distinguished
from both genera by the palps with E without basal projection, filiform along its entire length (e.g. Figs 40, 75,
122), or distally filiform with wide base, in N. carare sp. nov. and N. mariua sp. nov. (Figs 17, 60) (E not filiform
with basal projection in the latter genera) and by the epigyne with LL fused along posterior half (e.g. Figs 77,
94, 109) (completely separated in Guadana and separated or touching each other, leaving a median fissure in
Sparianthina).
Description. Total length of males 5.51−8.71, of females 5.5−9.07. Dorsal shield of prosoma generally longer
than wide, can be as wide as long. Cephalic region slightly higher than thoracic region, flattening posteriorly. Fovea
conspicuous on posterior third of prosoma. Eyes arranged in two rows, the anterior recurved and the posterior
straight; AME smaller than ALE, more distant from each other than from laterals; PME smaller than PLE, all mostly
equidistant (e.g. Figs 11, 19, 24). Clypeus low, less than AME diameter. Chelicerae longer than wide with three
promarginal teeth, median one largest, and 6−8 retromarginal teeth, three subequal, others smaller. Between 10−20
intermarginal denticles clustered in front of promarginal teeth and in a row along rest of groove. Promargin with
one escort seta (Fig. 1). Labium rebordered as wide as long. Endites slightly convergent, longer than wide, with
dense scopula on internal margin. Serrula with single row of denticles (Fig. 2). Sternum longer than wide, slightly
projected between coxae IV. Female palp with single pectinate claw with 5−6 long teeth (Fig. 3); sensory setae
long, distally curved, with barbules along the entire setae and with distal region bearing a large rounded pore and
a single filiform extension, scattered dorsally along palpal tarsus (Fig. 4). Legs laterigrade, generally 2143, rarely
1423. Spination pattern in males: femora I−III: p1-1-1, d0-1-1, r1-1-1; femur IV: p1-1-1, d0-1-1, r0-0-1; patellae
I−IV: 0; tibiae I−II: p1-0-1, d1-1-1, r1-0-1, v2-2-0; tibiae III−IV: p1-0-1, d0-0-1, r1-0-1, v2-2-0; metatarsi I−II: p1-
0-0, r1-0-0, v2-2-0; metatarsus III: p1-1-0, r1-1-0, v2-2-1; metatarsus IV: p1-1-2, r1-1-2, v2-2-0; palp: femur: p0-
0-1, d0-1-2, r0-0-1; patella: p1, r1; tibia: p2-1-0, d1-0-0; in females: femora I−III: p1-1-1, d0-1-1, r1-1-1; patellae
I−IV: 0; tibiae I−III: p1-0-1, r1-0-1, v2-2-0; tibia IV: p1-0-1, d0-0-1, r1-0-1, v2-2-0; metatarsi I−II: p1-0-0, r1-0-0,
v2-2-0; metatarsus III: p1-1-0, r1-1-0, v2-2-0; metatarsus IV: p1-1-2, r1-1-2, v2-2-0; palp: femur: p0-0-1, d0-1-2,
r0-0-1; patella: p1, r1; tibia: p2-1-0, d1-0-0, r1-1-0; tarsus p2-1-0, r2-1-0. Trochanter deeply notched. Metatarsi I–IV
distally with dorsal trilobate membrane with median hook slightly shorter than lateral projections (Fig. 5). Tarsi
and distal half of metatarsi scopulate. Trichobothria present on dorsal tibiae, metatarsi and tarsi, arranged in several
rows on tarsi, converging to a single row on metatarsi. Dorsal plate of trichobothria with one incomplete transversal
groove, projecting over a smooth proximal plate (Fig. 6). Tarsal organ capsulate, with drop-shaped opening, located
dorsally, at distal end of tarsi (Fig. 7). Tarsi with pair of pectinate claws with 10−12 short teeth and claw tufts (Figs
8−9). Opisthosoma oval, longer than wide, with two pairs of muscle sigilla located in the anterior region. Male
epiandrium with scattered small groups of epiandrous spigots (Fig. 10). Six spinnerets: ALS contiguous, conical
and bi-segmented; basal segment elongated and cylindrical; distal segment short and truncated. PMS conical and
short. PLS conical and bi-segmented; basal segment elongate and cylindrical; distal segment short and truncated.
Male palp: tibia slightly longer than half cymbium length with three prolateral and one dorsal spines (e.g. Figs 16,
47, 91); VTA present, slightly displaced retrolaterally (e.g. Figs 40, 92, 137) or reduced (e.g. Figs 48, 75, 107); RTA
distal with dRTA single (e.g. Figs 18, 75) or bifid (e.g. Figs 29, 48, 60) and vRTA rounded cleaver-shaped, best seen
in lateral view (e.g. Figs 93, 108, 138); cymbium slightly elongate with large rounded alveolus, dorsal elongate
scopula and rectangular RpP; subtegulum ring-shaped bearing a subtegular locking lobe that fits into an indentation
NATIVUS gen. nov. Zootaxa 5360 (1) © 2023 Magnolia Press ·

in the DTA (e.g. Figs 28, 74, 91); tegulum generally smooth (Fig. 17), some species can have a small triangular
protrusion close to bp (Figs 60, 107) and/or RdP (e.g. Figs 29, 75, 137); E arising from tegulum at 9 o’clock position,
filiform, sinuous (e.g. Figs 48, 75, 137) or straight (e.g. Figs 17, 60, 92); C hyaline with slender base, fanning out
to accommodate the embolus distally; DTA arising from tegulum between 8−9 o’clock position acutely tapering
and curved retrolaterally, generally smooth (e.g. Figs 39−40, 59‑60, 91−92),with median projection in N. tupana sp.
nov. (Figs 121−122); bp hyaline, laminar, of varied shapes (e.g. Figs 48, 60, 122). Female epigyne: EF wider than
long or as long as wide without conspicuous muscle attachment bands, divided into LL and MS; LL fused medially
(e.g. Figs 31, 62, 109) or slightly posteriorly (e.g. Figs 77, 94, 139), smooth, except N. tawu sp. nov. with lateral
pockets (Fig. 109); MS smooth; CO situated anteriorly, in a depressed area (COd). Vulva: Internal ducts with first
winding antero-laterad and then strongly convoluted, encapsulated within a sclerotized, elongated structure that
swells posteriorly (ducts can be seen by transparency in less sclerotized specimens, e.g. Figs 90, 120); FD short and
curved (e.g. Figs 32, 63, 95).
Composition. Ten species: Nativus carare sp. nov., N. hazzii sp. nov., N. janai sp. nov., N. juriti sp. nov., N.
mariua sp. nov., N. napo sp. nov., N. nocaima sp. nov., N. tawu sp. nov., N. tupana sp. nov. and N. yurupari sp.
nov.
Distribution. Northern South America: Brazil, Colombia and Peru.
Identification key for species of Nativus gen. nov.
1 Males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2(1) Tegulum with triangular RdP, arising between 1−2 o’clock; E sinuous (e.g. Figs 29, 48, 92) . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Tegulum without triangular RdP; E roughly straight (Figs 17, 60, 107) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3(2) dRTA distally bifid, with two pointed branches (e.g. Figs 30, 49, 93) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- dRTA distally truncated without pointed branches (Figs 75−76) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. napo sp. nov.
4(3) dRTA with ventral branch shorter than dorsal branch (e.g. Figs 41, 93, 123) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- dRTA with ventral branch as long as or longer than dorsal branch (e.g. Figs 30, 138) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
5(4) DTA without median projections (e.g. Figs 39, 47, 91) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- DTA with large median projection (Figs 121−122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tupana sp. nov.
6(5) RdP small, wider than long or as wide as long, rounded at tip (Figs 48, 92) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- RdP pronounced, longer than wide, pointed (Fig. 40) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. janai sp. nov.
7(6) RdP small, much wider than long; bp as wide as long (Fig. 92) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nocaima sp. nov.
- RdP as wide as long; bp slightly elongate, longer than wide (Fig. 48) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. juruti sp. nov.
8(4) dRTA with two pointed tips, in ventral view (Fig. 30); RdP small, wider than long; bp large, roughly triangular, slightly wider
than long (Fig. 29) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. hazzii sp. nov.
- dRTA with one blunt and one pointed tip, in ventral view (Fig. 138); RdP large, triangular, as long as wide; bp small, wider than
long (Fig. 137) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. yurupari sp. nov.
9(2) Tegulum bearing small tp close to TBE (between 8‑9 o’clock); dRTA bifid, with ventral branch long and slender, curved in
ventral view (Figs 60, 107) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Tegulum without triangular projection; dRTA single, truncated (Figs 17−18) . . . . . . . . . . . . . . . . . . . . . . . . . N. carare sp. nov.
10(9) bp keel-like, wider than long (Fig. 60); dRTA with ventral branch straight in retrolateral view (Fig. 61) . . . . . N. mariua sp. nov.
- bp irregular-shaped, slightly longer than wide (Fig. 107); dRTA with ventral branch hooke-shaped in retrolateral view (Fig.
108). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tawu sp. nov.
11(1) MS situated anterior to fused part of LL (Figs 78, 139) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- MS situated posterior to fused part of LL (e.g. Figs 31, 62, 124) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12(11) Fused part of LL extending until 1/3 of EF length; MS roughly as wide as long (Fig. 77) . . . . . . . . . . . . . . . . N. napo sp. nov.
- Fused part of LL extending until almost half EF length; MS two times longer than wide (Fig. 139) . . . . . N. yurupari sp. nov.
13(11) Fused part of LL forming an inverted T shape (e.g. Figs 62, 94, 109) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Fused part of LL forming an inverted Y shape (Fig. 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. hazzii sp. nov.
14(13) LL fused medially; MS large (Figs 62, 109) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- LL fused medio-posterially; MS small (Figs 94, 124) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15(14) MS hexagonal, as wide as long; fused part of LL without epigynal pockets (Fig. 62) . . . . . . . . . . . . . . . . . . N. mariua sp. nov.
- MS trapezoid, wider than long; fused part of LL with pair of semi circular-shaped pockets (Fig. 109) . . . . . . N. tawu sp. nov.
16(14) COd C-shaped, seprated from each other by half MS width (Fig. 94) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nocaima sp. nov.
- COd J-shaped, separated from each other by almost MS width (Fig. 124) . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tupana sp. nov.

FIGURES 1−6. Nativus tupana sp. nov. (IBSP 8828). 1 Male, left chelicerae, ventral view (promarginal teeth in yellow;
intermarginal denticles in blue; retromarginal teeth in red; arrow points to retrolateral escort seta); 2 Male, left endite, serrula,
ventral view; 3 Female, left palp, claw, prolateral view; 4 Female, palp, tarsus, sensory setae; 5 Male, left leg I, metatarsus,
trilobate membrane, dorsal view; 6 Male, left leg I, tarsus, trichobothrium, dorsal view. DP = distal plate; PP = proximal plate.

FIGURES 7−10. Nativus tupana sp. nov. (IBSP 8828). 1 Male, left leg I, tarsus, tarsal organ, dorsal view; 2 Male, left leg I,
tarsus, claws, prolateral view; 3 Female, left leg I, tarsus, claws, prolateral view; 4 Male, epiandrium, epiandrous spigots (arrows
point to clusters of spigots).
Nativus carare sp. nov.

Figs 11−18
Type material: Holotype: COLOMBIA: Santander: ♂, Municipio Puerto Araujo (6.5164, ‑74.0969), Veredas Las
Marías, Hacienda Los Manantiales, 1 April 2003, S. Bustamante leg. (MPUJ_ENT0070589).
Additional material examined. COLOMBIA: Boyacá: 1♂, Otanche, Serranía Las Quinchas (5.8463,
‑74.2709), April 1997, G. Alarcón leg. (ICN−Ar-12998).
Etymology. The specific name refers to the Carare River, located close to the collection locality of the holotype;
noun in apposition.
Diagnosis. Males of N. carare sp. nov. resemble those of N. mariua sp. nov. (Figs 59−61) and N. tawu sp. nov.
(Figs 106−108) by the palps with E roughly straight and tegulum lacking RdP. They are distinguished from both
species by the dRTA single and by the tegulum lacking small triangular protrusion close to bp (Figs 17−18) (dRTA
bifid, with ventral branch hook-shaped in retrolateral view and triangular protrusion present in the latter species).
Females unknown.

FIGURES 11−15. Nativus carare sp. nov., male (MPUJ_ENT0070589). 11−12 Habitus (11 dorsal, 12 ventral); 13−15 Left palp
(13 prolateral, 14 ventral, 15 retrolateral). Scale lines: 11−12 = 2 mm, 13−15 = 1 mm.
Description. Male (holotype): Prosoma orange brown with brown margins; cephalic region with three pale
brown longitudinal lines extending posteriorly from behind ALE and from between PME; fovea and thoracic striae
dark brown; eye borders black. Chelicerae brown, lighter than prosoma. Legs and palps brown, lighter than prosoma.
Endites brownish orange, distally lighter. Labium brown, distally pale brown. Sternum pale yellow, with orange
margins. Opisthosoma yellowish cream colored; dorsally with few white marks scattered laterally and around cardiac
mark; ventrally with pair of U-shaped muscle impressions. Spinnerets yellowish cream colored (Figs 11−12). Total

length 8.79. Prosoma: 4.15 long, 4.07 wide. Opisthosoma: 4.47 long, 2.52 wide. Eyes diameters: 0.35, 0.41, 0.40,
0.60; interdistances: 0.53, 0.60, 0.76, 0.73, 0.76, 0.81. Legs: I: 25.35 (5.62, 1.95, 7.90, 7.32, 2.56); II: absent; III:
19.06 (5.54, 1.69, 5.61, 4.80, 1.45); IV: 24.37 (6.47, 1.57, 6.50, 7.31, 2.52). Spination follows the generic pattern,
except femur III: r1-0-1; tibia III: d1-0-1; metatarsus III: v2-2-0. Palp: PTA roughly triangular, with rounded tip,
slightly wider than long; VTA triangular, displaced retrolaterally; vRTA two times longer than wide in retrolateral
view; dRTA spade-shaped with small subdistal triangular projection; tegulum smooth without protrusions; bp keel-
like, at least 7 times wider than long; DTA without projections (Figs 13−18).
FIGURES 16−18. Nativus carare sp. nov., male (MPUJ_ENT0070589), left palp (16 prolateral, 17 ventral, 18 retrolateral).
Scale line: 1 mm. bp = projection at base of embolus; C = conductor; dRTA = dorsal branch of RTA; DTA = dorsal tegular
apophysis; E = embolus; PTA = prolateral tibial apophysis; RpP = retroproximal cymbial projection; STlb = subtegular locking
lobe; vRTA = ventral branch of RTA; VTA = ventral tibial apophysis.
Female: Unknown.
Variation. Males (n = 2): Total length 8.74−8.79; prosoma length 4.13−4.15; femur I length 5.59−5.62.
Distribution. Only known from Boyacá and Santander departments, central-northern Colombia (Fig. 142).

Nativus hazzii sp. nov.
Figs 19−33
Type material: Holotype: COLOMBIA: Amazonas: ♂, corregimiento La Pedrera, quebrada El Ayo (‑1.5833,
‑69.4666), April 2001, J. Pinzón & A. Sabogal leg. (ICN-Ar-12999). Paratypes: COLOMBIA: Amazonas: 1♂,
2 ♀, same locality as for holotype, April 2001, J. Pinzón & A. Sabogal leg. (ICN-Ar-5789); 1♂, 2 ♀, same locality
as holotype, May 2002, J. Pinzón leg. (ICN-Ar-13000). Vaupés: 1♀, Taraira, Serranía Taraira (‑0.8988, ‑69.7021),
April 2012 (ICN-Ar-13001).
FIGURES 19−23. Nativus hazzii sp. nov., male (ICN-Ar-12999). 19−20 Habitus (19 dorsal, 20 ventral); 21−23 Left palp (21
prolateral, 22 ventral, 23 retrolateral). Scale lines 19−20 = 2 mm; 21−23 = 1 mm.

FIGURES 24−27. Nativus hazzii sp. nov., female (ICN-Ar-12999). 24−25 Habitus (24 dorsal, 25 ventral); 26−27 Epigyne/
vulva (26 ventral, 27 dorsal). Scale lines: 24−25 = 2 mm; 26−27 = 1 mm.
Etymology. The specific name honors arachnologist Dr. Nicolas Hazzi (George Washington University), for his
contributions to the knowledge of biogeography, systematics and taxonomy of Neotropical ctenids; name in genitive
case.
Diagnosis. Males of N. hazzii sp. nov. resemble those of N. juruti sp. nov. (Figs 47−49), N. nocaima sp. nov.
(Figs 91−93) and N. tupana sp. nov. (Figs 119−121) by the palps with tegulum bearing small, wider than long RdP
and sinuous E. They are distinguished from N. tupana sp. nov. by the DTA smooth with no medial projections (Figs
28−29) (present in N. tupana sp. nov.) and from N. juruti sp. nov. and N. nocaima sp. nov. by the dRTA with two
similar sized branches (ventral branch shorter than dorsal branch in N. juruti sp. nov. and N. nocaima sp. nov.).
Females are distinguished from all congeners by LL in roughly an inverted Y-shape (Fig. 31).
Description. Male (holotype): Prosoma orange brown, with slightly darker lateral margins; cephalic region
with longitudinal stripes extending posteriorly from behind posterior eyes; fovea dark brown, thoracic striae slightly
lighter; eye borders black. Chelicerae pale orange brown. Legs pale brown, lighter than prosoma. Endites pale
brown, distally lighter. Labium orange brown, distally pale brown. Sternum yellowish cream colored with orange
10 · Zootaxa 5360 (1) © 2023 Magnolia Press Casas & Rheims

FIGURES 28−33. Nativus hazzii sp. nov. 28−30 Male (ICN-Ar-12999), left palp (28 prolateral, 29 ventral, 30 retrolateral);
31−33 Female (ICN-Ar-5789), epigyne/vulva (31 ventral, 32 dorsal, 33 schematic course of internal duct system). Scale lines:
1 mm. bp = projection at base of embolus; C = conductor; CO = copulatory opening; COd = copulatory opening depression;
dRTA = dorsal branch of RTA; DTA = dorsal tegular apophysis; E = embolus; ed = encapsulated part of internal ducts; FD
= fertilization duct; fw = first winding of copulatory duct; LL = lateral lobe; MS = median septum; PTA = prolateral tibial
apophysis; RdP = retrodistal tegular protrusion; RpP = retroproximal cymbial projection; STlb = subtegular locking lobe; vRTA
= ventral branch of RTA; VTA = ventral tibial apophysis.
NATIVUS gen. nov. Zootaxa 5360 (1) © 2023 Magnolia Press · 11

margins. Opisthosoma yellowish cream colored; dorsally with few scattered white marks laterally and two pairs of
round, brown muscle impressions; ventrally with V-shaped lines of muscle impressions. Spinnerets pale yellowish
cream colored (Figs 19−20). Total length 5.51. Prosoma: 2.6 long, 2.84 wide. Opisthosoma: 2.76 long, 1.84 wide.
Eyes: diameters: 0.15, 0.17, 0.28, 0.36; interdistances: 0.51, 0.52, 0.70, 0.63, 0.69, 0.68. Legs (2143): I: 17.44 (4.38,
1.22, 5.08, 4.9, 1. 86); II: 20.96 (5.42, 1.02, 5.87, 6.05, 2.6); III: 13.01 (3.61, 0.81, 3.94, 3.34, 3.84, 1.41); IV: 17.27
(4.9, 0.94, 3.78, 5.55, 2.1). Spination follows the generic pattern, except femur IV: p1-1-1, d0-1-1, r0-1-1; tibiae
I−II: p1-0-1, d1-1-1, r1-0-1, v2-2-0; tibia III: p1-0-1, d1-0-1, r1-0-1, v2-2-0; tibia IV: p1-0-1, d1-1-1, r1-0-1, v2-2-0.
Palp: PTA finger-like, roughly 1.5 times longer than wide; VTA triangular, displaced retrolaterally; vRTA two times
longer than wide in retrolateral view; dRTA bifid, resembling a small bird (in retrolateral view), with two pointed
branches; tegulum with small RdP at least three times wider than long; bp roughly triangular, as long as wide,
membranous at center with hyaline margins; DTA with no projections (Figs 21−23, 28−30).
Female (paratype): Prosoma orange-brown; cephalic area with longitudinal brown stripes extending posteriorly
from behind PLE, PME and from between AME; fovea and thoracic striae brown; eye borders black. Chelicerae
orange-brown. Legs orange brown, lighter than prosoma. Endites pale orange, distally lighter. Labium brown, distally
orange brown. Sternum yellowish cream colored with orange margins. Opisthosoma brownish gray; dorsally with
few scattered white marks laterally on anterior half and two pairs of brown, round muscular impressions; ventrally
with two roughly parallel lines of muscle impressions. Spinnerets orange brown, distally lighter (Figs 24−25).
Total length: 7.11. Prosoma: 2.60 long, 3.00 wide. Opisthosoma: 4.11 long, 2.51 wide. Eyes: diameters: 0.23, 0.27,
0.27, 0.30; interdistances: 0.33, 0.35, 0.46, 0.44, 0.46, 0.47. Legs (2143): I: 14.58 (3.93, 1.38, 4.23, 3.41, 1.63); II:
15.71 (4.05, 1.60, 4.61, 3.96, 1.49); III: 11.90 (3.38, 1.15, 3.08, 3.11, 1.18); IV: 14.45 (4.12, 1.14, 3.53, 3.98, 1.68).
Spination follows the generic pattern, except tibiae I−II: p0-0-1, r0-0-1; tibiae III−IV: d1-0-1; metatarsus IV: p1-1-2,
r1-1-2, v2-2-0. Epigyne: EF roughly as wide as long; LL fused medially, converging anteriorly; MS triangular, as
wide as long, posterior to fused part of LL; pair of long elliptical COd, bearing pair of CO situated posteriorly (Figs
26, 31). Vulva: CD with first winding posteriad then bent antero-laterad; encapsulated part of internal ducts more
than two times longer than wide, more dilated posteriorly; FD anteriad (Figs 27, 32−33).
Variation. Males (n = 3): Total length 5.51−6.78; prosoma length 2.6−3.17; femur I length 4.04−4.38. Females
(n = 4): Total length 6.78−7.11; prosoma length 2.60−3.17; femur I length 3.93−4.04.
Distribution. Only known from the type locality, in Amazonas, Colombia (Fig. 142).
Nativus janai sp. nov.

Figs 34−41
Type material: Holotype: COLOMBIA: Leticia: ♂, Reserva Forestal del Rio Calderon, Estación Biologica
el Zafire (‑4.0058, ‑69.9125), 146 m, 5−7 December 2007, L. Franco & S. Florez leg. (MPUJ_ENT0070440).
Paratypes: COLOMBIA: Leticia: 2 ♂, same data as for holotype, (MPUJ_ENT0086798).
Additional material examined: COLOMBIA: Leticia: 1♂, Leticia, Comunidad Monilla Amena Km 9
(-4.1121, -69.928), 70 m, 30 October 2004, A. Daza & E. Torres leg. (MPUJ_ENT0086786); 1♂, same locality as
for previous specimen, 35m, October 2005, Marin & M. Salgado leg. (MPUJ_ENT0087072)
Etymology. The specific name refers to Janai (meaning incomprehensible to the senses), a general designation
of the indigenous Uitoto tribe, from southeastern Colombia and northern Peru, for ghostly or supernatural creatures;
noun in apposition.
Diagnosis. Males of N. janai sp. nov. resemble those of N. napo sp. nov. (Figs 74−76) and N. yurupari sp. nov.
(Figs 136−138) by the tegulum with large triangular RdP, at least 1.5 times longer than wide and E sinuous in the
male palps. It is distinguished from N. napo sp. nov. by the dRTA bifid with two branches (Figs 40−41) (single and
distally truncated in N. napo sp. nov.) and from N. yurupari sp. nov. by the bp large, irregularly-shaped, slightly
wider than long, with anterior margin medially depressed (Fig. 40) (keel-like, over two times wider than long in N.
yurupari sp. nov.). Females unknown.
Description. Male (holotype): Prosoma orange brown with darker margins, slightly darker at eye area; fovea
dark brown; eye borders black. Chelicerae pale brown, lighter than prosoma. Legs and palps pale brownish orange.
Endites pale brown. Labium brown, anteriorly lighter. Sternum yellowish cream colored with pale brown margins.
Opisthosoma grey; dorsally with two pairs of brownish gray, rounded muscle impressions. Spinnerets cream

colored (Figs 34−35). Total length 5.77. Prosoma: 2.95 long, 3.15 wide. Opisthosoma: 2.89 long, 1.86 wide. Eyes:
diameters: 0.31, 0.33, 0.31, 0.42; interdistances: 0.49, 0.52, 0.70, 0.63, 0.67, 0.68. Legs (2143): I: 19.97 (5.13, 1.43,
5.87, .31, 2.23); II: 23.26 (5.97, 1.56, 6.85, 6.36, 2.52); III: 13.61 (3.74, 1.10, 3.76, 3.53, 1.48); IV: 18.87 (4.91, 1.20,
4.96, 5.60, 2.20). Spination follows the generic pattern, except femur III: r0-0-1; tibia III: p0-0-1; tibia IV: d1-0-1;
metatarsus III: p0-1-0. Palp: PTA finger-like, bent distally, two times longer than wide; VTA triangular, displaced
retrolaterally; vRTA roughly 2.5 times longer than wide in retrolateral view; dRTA with dorsal branch large, spoon-
shaped, slightly longer than wide, widest medially, and ventral branch small, distally pointed (Figs 36−41).
FIGURES 34−38. Nativus janai sp. nov., male (MPUJ_ENT0070440). 34−35 Habitus (34 dorsal, 35 ventral); 36−38 Left palp
(36 prolateral, 37 ventral, 38 retrolateral). Scale lines: 34−35 = 2 mm; 36−38 = 1 mm.

FIGURES 39−41. Nativus janai sp. nov., male (MPUJ_ENT0070440), left palp (39 prolateral, 40 ventral, 41 retrolateral).
Scale lines: 1 mm. bp = projection at base of embolus; C = conductor; dRTA = dorsal branch of RTA; DTA = dorsal tegular
apophysis; E = embolus; PTA = prolateral tibial apophysis; RdP = retrodistal tegular protrusion; RpP = retroproximal cymbial
projection; STlb = subtegular locking lobe; vRTA = ventral branch of RTA; VTA = ventral tibial apophysis.
Female: Unknown.
Distribution. Only known from the type locality in Leticia, southern Colombia (Fig. 142).
Nativus juriti sp. nov.

Figs 42−49
Type material. Holotype: BRAZIL: Pará: ♂, Juruti, Sítio Barroso (‑2.4616, ‑56.0032), 14 August 2006, N.F. Lo
Man Hung leg. (MPEG 8561). Paratype: BRAZIL: Pará: 1♂, same locality as for holotype (‑2.4643, ‑56.0024),
8 February 2007, J.A.P. Barreiros leg. (MPEG 30726).
Etymology. The specific name refers to the Juriti bird (genus Leptotila Swaison), which used to be present in
large numbers at the time of the founding of the Juruti municipality; noun in apposition.

FIGURES 42−46. Nativus juriti sp. nov., male (MPEG 8561). 42−43 Habitus (42 dorsal, 43 ventral); 44−46 Left palp (44
prolateral, 45 ventral, 46 retrolateral). Scale lines: 42−43 = 2 mm; 44−46 = 1 mm.
Diagnosis. Males of N. juruti sp. nov. resemble those of N. hazzii sp. nov. (Figs 28−30), N. nocaima sp. nov.
and E sinuous. They are distinguished from N. tupana sp. nov. by the DTA smooth with no medial projections (Figs
47−48) (present in N. tupana sp. nov.); from N. hazzii sp. nov. by the dRTA with ventral branch smaller than dorsal
branch (Fig. 49) (similar-sized in N. hazzii sp. nov.) and from N. nocaima sp. nov. by the bp rectangular, longer than
wide and RdP roughly two times wider than long (bp irregularly-shaped, as wide as long and RdP smaller, more than
three times wider than long in N. nocaima sp. nov.). Females are unknown.

FIGURES 47−49. Nativus juriti sp. nov., male (MPEG 8561), left palp (47 prolateral, 48 ventral, 49 retrolateral). Scale lines:
1 mm. bp = projection at base of embolus; C = conductor; dRTA = dorsal branch of RTA; DTA = dorsal tegular apophysis; E =
embolus; PTA = prolateral tibial apophysis; RdP = retrodistal tegular protrusion; RpP = retroproximal cymbial projection; STlb
= subtegular locking lobe; vRTA = ventral branch of RTA; VTA = ventral tibial apophysis.
Description. Male (holotype): Prosoma orange brown with dark brown margins and brown thoracic striae;
cephalic region with brown lines extending posteriorly from PLE, PME and between AME; fovea dark brown; eye
borders black. Legs and palps pale brownish orange. Endites pale brownish orange, distally lighter. Labium brown,
distally orange brown. Sternum pale range brown with brown margins. Opisthosoma yellowish cream colored;
dorsally with small grayish brown marks and two pairs of rounded, brown muscular impressions; ventrally with
slightly conspicuous dark gray median longitudinal band and U-shaped lines of muscular impressions. Spinnerets
yellowish cream colored, distally lighter (Figs 42−43). Total length 6.3. Prosoma: 3.3 long, 3.1 wide. Opisthosoma:
3.0 long, 2.0 wide. Eyes: diameters: 0.22, 0.25, 0.20, 0.30; interdistances: 0.11, 0.04, 0.26, 0.26, 0.26, 0.20. Legs
(2143): I: 19.2 (5.0, 1.4, 5.4, 5.2, 2.2); II: 22.6 (5.8, 1.5, 6.5, 6.2, 2.6); III: 15.0 (4.3, 1.2, 4.0, 4.0, 1.5); IV: 18.4
(5.1, 1.1, 4.7, 5.5, 2.0). Palp: PTA, finger-like, as long as wide; VTA reduced, inconspicuous; vRTA roughly 1.5
times longer than wide in retrolateral view; dRTA with two pointed branches, dorsal one roughly three times longer
than ventral branch; RdP triangular, distally rounded, two times wider than long; DTA with no projections (Figs
44−49).
Female: Unknown.
Distribution. Only known from the type locality, in western Pará, Brazil (Fig. 142).

Nativus mariua sp. nov.
Figs 50−64
FIGURES 50−54. Nativus mariua sp. nov., male (INPA). 50−51 Habitus (50 dorsal, 51 ventral); 52−54 Left palp (52 prolateral,
53 ventral, 54 retrolateral). Scale lines: 50−51 = 2 mm; 52−54 = 1 mm.
Type material. Holotype: BRAZIL: Amazonas: ♂, Barcelos [1.1833, ‑64.9667], 29 September 2006, R. Braga
Neto leg. (INPA). Paratypes: BRAZIL: Amazonas: all same locality as for holotype, 2 ♂, 14 October 2006, A.A.
Nogueira leg. (INPA); 1♂, 16 October 2006, A.A. Nogueira leg. (INPA); 1♂, 17 October 2006 A.A. Nogueira leg.

(IBSP 273022); 1♂, 30 September 2006, A.A. Nogueira leg. (IBSP 273023); 1♀, 17 October 2006, A.A. Nogueira
leg. (IBSP 273024); 1♀, 17 October 2006, R. Braga Neto leg. (IBSP 273025); 1♀, 3 October 2006, R. Braga Neto
leg. (INPA-Df); 1♀ 13 October 2006, R. Braga Neto leg. (INPA); 2 ♀, 6 October 2006, A.A. Nogueira leg. (INPA);
1♂, 1♀, 10 October 2006, A.A. Nogueira leg. (INPA); 1♂, 29 September 2006, A.A. Nogueira leg. (MZSP); 1♀, 4
October 2006, A.A. Nogueira leg. (MZSP).
FIGURES 55−58. Nativus mariua sp. nov., female (INPA-Df). 55−56 Habitus (55 dorsal, 56 ventral); 57−58 Epigyne/vulva (57
ventral, 58 dorsal). Scale lines: 57−58 = 2 mm, 59−60 = 1 mm.
Additional material examined. BRAZIL: Amazonas: 1♂, São Gabriel da Cachoeira, [‑0.1188, ‑67.0850],
Pico da Neblina, Cachoeira do Tucano, 23 September 2007, N. Lo Man Hung leg. (IBSP 278829); 1♂, Barcelos
[1.1833, ‑64.9667], Serra do Tapirapecó, 14 October 2006, A.A. Nogueira leg. (INPA); 1♂, 17 October 2006, R.
Braga Neto leg. (INPA); 1♀, 4 October 2006, A.A. Nogueira leg. (INPA); 1♂, 17 October 2006, R. Braga Neto leg.
(INPA); 1♂, 3 October 2006, A.A. Nogueira leg. (INPA); 1♂, 1♀, October 2006, R. Braga Neto leg. (INPA); 1♂, 6
October 2006, R. Braga Neto leg. (INPA); all same locality.

FIGURES 59−64. Nativus mariua sp. nov. 59−61 Male (INPA), left palp (59 prolateral, 60 ventral, 61 retrolateral); 62−64
Female (INPA-Df) (62 epigyne, ventral, 63 vulva, dorsal, 64 schematic course of internal duct system). Scale lines: 1 mm.
bp = projection at base of embolus; C = conductor; CO = copulatory opening; Cod = copulatory opening depression; dRTA =
dorsal branch of RTA; DTA = dorsal tegular apophysis; E = embolus; ed = encapsulated part of internal ducts; FD = fertilization
duct; fw = first winding of copulatory duct; LL = lateral lobe; MS = median septum; PTA = prolateral tibial apophysis; RpP =
retroproximal cymbial projection; STlb = subtegular locking lobe; tp = tegular protrusion; vRTA = ventral branch of RTA; VTA
= ventral tibial apophysis.

Etymology. The specific name refers to the Vila de Mariuá, built by the Manaus Indians at the right margin of
the Negro River, that later became the city of Barcelos; noun in apposition.
Diagnosis. Males of N. mariua sp. nov. resemble those of N. carare sp. nov. (Figs 16−18) and N. tawu sp. nov.
(Figs 106−108) by the palps with E roughly straight and tegulum lacking RdP (Fig. 60). They are distinguished from
N. carare sp. nov. by the dRTA bifid, with two pointed branches and triangular protrusion close to bp (Figs 60−61)
(dRTA distally truncated and tegulum lacking small triangular protrusion close to bp in N. carare sp. nov.) and from
N. tawu sp. nov. by the bp keel-like, wider than long and dRTA with ventral branch straight in retrolateral view (Figs
60‑61) (bp irregular-shaped, slightly longer than wide and dRTA with ventral branch hook-shaped in retrolateral
view in N. tawu sp. nov.). Females resemble those of N. tawu sp. nov. (Figs 109−111) by the LL fused medially with
pair of anterior COd bearing posterior CO and large, posterior MS. They are distinguished by the MS trapezoid, as
long as wide and fused part of LL as long as anterior COd (Fig. 62) (MS rectangular, wider than long and fused part
of LL shorter than anterior COd in the latter species).
Description. Male (holotype): Prosoma brown with dark brown margins and darker thoracic striae; cephalic
region with brown longitudinal stripe extending posteriorly from behind AME; fovea dark brown; eye borders
black. Chelicerae, legs and palps pale yellowish brown. Endites pale brownish yellow. Labium brown, distally pale
brownish yellow. Sternum pale yellowish cream colored with brown margins. Opisthosoma gray; dorsally with
pair of rounded darker gray muscle impressions; ventrally with V-shaped lines of muscle impressions. Spinnerets
grayish cream colored, distally lighter (Figs 50−51). Total length 6.7. Prosoma: 3.3 long, 3.2 wide. Opisthosoma:
3.3 long, 1.9 wide. Eyes: diameters: 0.22, 0.26, 0.18, 0.30; interdistances: 0.15, 0.05, 0.32, 0.31, 0.28, 0.26. Legs
(2143): I: 19.6 (5.0, 1.4, 6.6, 5.3, 2.3); II: 22.7 (5.8, 1.4, 6.5, 6.4, 2.6); III: 15.0 (4.1, 1.2, 4.0, 4.1, 1.6); IV: 18.2
(5.0, 1.1, 4.8, 5.2, 2.1). Palp: PTA resembling the top part of a thumb, longer than wide; VTA small, triangular,
slightly displaced retrolaterally; vRTA slightly over 1.5 times longer than wide in retrolateral view; dRTA with
ventral branch acuminate, three times longer than wide and dorsal branch spoon-shaped with subdistal triangular
projection; tegulum with small triangular protrusion close to bp; DTA without projections (Figs 52−54, 59−61).
Female (paratype, INPA-Df): Prosoma brown with darker brown reticulated margins and dark brown thoracic
striae; cephalic region with dark brown lines extending posteriorly from behind PLE, PME and between AME;
fovea dark brown; eye borders black. Chelicerae, legs and palps pale yellowish brown. Endites pale yellow. Labium
pale brown, distally pale yellow. Sternum pale yellow with slightly darker margins. Opisthosoma brownish gray;
dorsally with pair of rounded brown muscle impressions; ventrally with two parallel lines of muscle impressions.
Spinnerets dorsally brownish gray, ventrally cream colored (Figs 55−56). Total length 7.5. Prosoma: 3.0 long, 3.0
wide, Opisthosoma: 4.1 long, 2.5 wide. Eyes: 0.20, 0.29, 0.20, 0.31; interdistances: 0.12, 0.04, 0.22, 0.30, 0.29,
0.27. Legs (2413): I: 13.2 (3.6, 1.5, 3.6, 3.1, 1.4); II: 14.8 (4.1, 1.5, 4.1, 3.5, 1.6); III: 10.9 (3.3, 1.1, 2.7, 2.7, 1.1); IV:
13.3 (3.9, 1.1, 3.2, 3.6, 1.5). Epigyne: EF slightly wider than long; anterior COd long, oval (Figs 57, 62). Vulva: first
winding of copulatory ducts posteriad and gently curving antero-laterally; encapsulated part of internal duct system
roughly 2.5 times longer than wide and mostly the same width throughout; FD postero-laterad (Figs 58, 63−64).
Variation. Males (n = 10): Total length 5.9−6.9; prosoma length 2.9−3.4; femur I length 4.8−5.1. Females (n =
9); Total length 6.1−8.6; prosoma length 2.8−3.1; femur I length 3.5−3.9.
Distribution. Only known from the type locality, in northern Amazonas, Brazil (Fig. 143).
Nativus napo sp. nov.

Figs 65−79
Type material: Holotype: PERU: Loreto: ♂, Maynas, Rio Napo (-1.8703, ‑74.7904, 200 m), 11 January 2009, W.
Yawaromi leg. (MUSM ENT-505866). Paratypes: PERU: Loreto: 1♂, from the same vial as holotype (MUSM
ENT-505866); 1♂, 2 juv., Estirón [‑3.3166, -71.8500], Río Ampiacu, 13 November−9 December 1961, B. Malkin
leg. (AMNH); 1♀, Pebas, Río Amazonas (‑3.1333, ‑71.8167), 12−15 April 1977, C.W. Myers leg. (AMNH).
Etymology. The specific name refers to the type locality; noun in apposition.
Diagnosis. Males of N. napo sp. nov. resemble those of N. janai sp. nov. (Figs 39−41) and N. yurupari sp. nov.
(Figs 136−138) by the tegulum with large triangular RdP, at least 1.5 times longer than wide and E sinuous in the
male palps. It is distinguished from both species by the palps with dRTA single and distally truncated (Figs 75−76)
(bifid in both latter species). Females resemble those of N. yurupari sp. nov. (Figs 139−141) by the MS diamond

shaped, anterior to fused part of LL. They are distinguished from the latter species by the fused part of LL extending
until 1/3 of EF length and MS roughly as wide as long (Fig. 77) (fused part of LL extending until almost half EF
length and MS two times longer than wide in the latter species).
FIGURES 65−69. Nativus napo sp. nov., male (MUSM 5058866). 65−66 Habitus (65 dorsal, 66 ventral); 67−69 Left palp (67
prolateral, 68 ventral, 69 retrolateral). Scale lines: 65−66 = 2 mm; 67−69 = 1 mm.

FIGURES 70−73. Nativus napo sp. nov., female (AMNH). 70−71 Habitus (70 dorsal, 71 ventral); 72−73 Epigyne/vulva (72
ventral, 73 dorsal). Scale lines: 70−71 = 2 mm, 72−73 = 1 mm.
Description. Male (holotype): Specimen slightly dried out. Prosoma brown, darker along thoracic striae; fovea
dark brown; eye borders black. Chelicerae legs and palps pale brown. Endites pale brown, distally slightly lighter.
Labium brown, distally pale brown. Sternum pale brown with darker brown margins. Opisthosoma brownish gray;
dorsally with one pair of rounded muscle impressions. Ventrally with two V-shaped lines of muscle impressions.
Spinnerets brownish gray (Figs 65−66). Total length 7.0. Prosoma: 3.7 long, 3.3 wide. Opisthosoma: 3.1 long, 1.6
wide. Eyes: diameters: 0.26, 0.24, 0.20, 0.28; interdistances: 0.12, 0.04, 0.26, 0.30, 0.20, 0.24. Legs (2413): I: 21.9
(5.9, 1.5, 6.4, 6.4, 2.7); II: 27.9 (7.2, 1.6, 7.6, 8.1, 3.4); III: 17.8 (4.9, 1.3, 4.7, 5.0, 1.9); IV: 22.3 (6.0, 1.2, 5.5, 7.0,
2.6). Palp: PTA triangular, distally rounded, as wide as long; VTA reduced; vRTA two times longer than wide in
retrolateral view; bp of irregular shape, roughly 1.5 times longer than wide (Figs 67−69, 74−76).
Female (paratype): Prosoma brown; fovea and thoracic striae dark brown; eye borders black. Chelicerae, palps
and legs brown, slightly lighter than prosoma; legs with faint brown marks at the base of spines. Labium brown,
distally pale brown. Endites pale brown, distally lighter. Sternum pale brown with brown margins. Opisthosoma
brown; dorsally with slightly lighter heart mark and two pairs of small rounded muscular impressions; ventrally
lighter brown with two more or less parallel lines of muscle impressions. Spinnerets pale brown, distally cream
colored (Figs 70−71). Total length 7.6. Prosoma 3.0 long, 2.7 wide. Opisthosoma: 4.6 long, 2.6 wide. Eyes: diameters:
0.23, 0.28, 0.22, 0.29; interdistances: 0.10, 0.04, 0.26, 0.27, 0.20, 0.22. Legs (2143): I: 12.8 (3.5, 1.3, 3.6, 3.1, 1.3);
II: 14.4 (4.1, 1.3, 4.1, 3.5, 1.4); III: 10.7 (3.2, 1.0, 2.7, 2.7, 1.1); IV: 12.7 (3.8, 0.9, 3.1, 3.5, 1.4). Epigyne: EF as

wide as long; MS with CO situated anteriorly (Figs 72, 77). Vulva: CD with first winding gently curving laterally;
encapsulated part of internal ducts 2.5 times longer than wide, dilated posteriorly; FD anteriad (Figs 73, 78−79).
Distribution. Known from the state of Loreto, in north and northeastern Peru (Fig. 143).
FIGURES 74−79. Nativus napo sp. nov. 74−76 Male (AMNH), left palp (74 prolateral, 75 ventral, 76 retrolateral); 77−79
Female (INPA), epigyne/vulva (77 ventral, 78 dorsal, 79 schematic course of internal duct system). Scale lines: 1 mm. bp =
projection at base of embolus; C = conductor; CO = copulatory opening; dRTA = dorsal branch of RTA; DTA = dorsal tegular
apophysis; E = embolus; ed = encapsulated part of internal ducts; FD = fertilization duct; fw = first winding of copulatory
duct; LL = lateral lobe; MS = median septum; PTA = prolateral tibial apophysis; RdP = retrodistal tegular protrusion; RpP =
retroproximal cymbial projection; STlb = subtegular locking lobe; vRTA = ventral branch of RTA.

FIGURE 80. Nativus nocaima sp. nov., male (MPUJ_ENT0086707).
Nativus nocaima sp. nov.

Figs 80−96
Type Material: Holotype: COLOMBIA: Cundinamarca: ♂, Nocaima, Casa de Oscar (5.0672,-74.3831), 188 m,
6 June 2022, O. Enciso leg. (MPUJ_ENT0086707). Paratypes: COLOMBIA: Cundinamarca: 3 ♀, same locality
as for holotype, 6 June 2022, O. Enciso leg. (MPUJ_ENT0086708); 2 ♂, same locality as for holotype, 6 June 2022,
O. Enciso leg. (MPUJ_ENT0086706).
Etymology. The specific name refers to the municipality of Nocaima, type locality of the new species and
hometown of Oscar Enciso, a self-taught and passionate naturalist who collected the specimen; noun in apposition.

FIGURE 81. Nativus nocaima sp. nov., female (MPUJ_ENT0086708).
Diagnosis. Males of N. nocaima sp. nov. resemble those of N. hazzii sp. nov. (Figs 28−30), N. juruti sp. nov.
and E sinuous. They are distinguished from N. tupana sp. nov. by the DTA smooth with no medial projections (Figs
91−92) (present in N. tupana sp. nov.); from N. hazzii sp. nov. by the dRTA with ventral branch smaller than dorsal
branch (Fig. 93) (similar-sized in N. hazzii sp. nov.) and from N. juruti sp. nov. by the bp irregularly-shaped, as
wide as long and RdP smaller, more than three times wider than long (Fig. 92) (bp rectangular, longer than wide and
RdP roughly two times wider than long in N. juruti sp. nov.). Females resemble those of N. tupana sp. nov. (Figs
124−126) by the epigyne with LL in roughly an inverted T-shape, with pair of elongate COd situated anteriorly. It is
distinguished from the latter species by the by the COd C-shaped and separated from each other by half MS width
(Fig. 94) (COd J-shaped and separated from each other by almost MS width in the latter species).

FIGURES 82−86. Nativus nocaima sp. nov., male (MPUJ_ENT0086707). 82−83 Habitus (82 dorsal, 83 ventral); 84−86 Left
palp (84 prolateral, 85 ventral, 86 retrolateral). Scale lines: 82−83 = 2 mm; 84−86 = 1 mm.
Description. Male (holotype): Prosoma pale yellowish brown with brown margins and grayish-brown lines
along thoracic striae and extending posteriorly from behind AME, PME and PLE; fovea orange brown; eye borders
black. Chelicerae, legs and palps pale yellowish brown. Sternum cream colored with brownish-orange margins.

Labium orange brown, slightly lighter distally. Endites pale yellowish cream colored. Opisthosoma pale gray;
dorsally with whitish cardiac mark and two pairs of brown muscle impressions; ventrally with two parallel lines
of muscle impressions; spinnerets cream colored (Figs 82−83). Total length 7.03. Prosoma: 3.25 long, 3.17 wide.
Opisthosoma: 3.39 long, 2.04 wide. Eyes: diameters: 0.24, 0.28, 0.25, 0.29; interdistances: 0.47, 0.43, 0.68, 0.62,
0.65, 0.66. Legs (2143): I: 18.53 (4.75, 1.52, 5.57, 4.79, 1.90); II: 21.60 (5.56, 1.61, 6.42, 5.81, 2.20); III: 14.22
(4.08, 1.18, 3.87, 3.68, 1.41); IV: 17.64 (4.84, 1.10, 4.61, 5.19, 1.90). Spination follows the generic pattern, except
tibiae III-IV: d1-0-1; metatarsus III: v2-2-0. Palp: PTA roughly rectangular, two times longer than wide; VTA
triangular, retrolaterally displaced; vRTA 1.5 times longer than wide in retrolateral view; dRTA bifid with dorsal
branch 1.5 times larger than ventral branch; tegulum with RdP small, barely conspicuous, three to four times wider
than long; bp irregularly-shaped, roughly as wide as long; DTA without projections (Figs 84−86, 91−93).
FIGURES 87−90. Nativus nocaima sp. nov., female (MPUJ_ENT0086708). 87−88 Habitus (87 dorsal, 88 ventral); 89−90
Epigyne/vulva (89 ventral, 90 dorsal). Scale lines: 87−88 = 2 mm; 89‑90 = 1 mm.

FIGURES 91−96. Nativus nocaima sp. nov. 91−93 Male (MPUJ_ENT0086707), left palp (91 prolateral, 92 ventral, 93
retrolateral); 94−96 Female (MPUJ_ENT0086708), epigyne/vulva (94 epigyne, dorsal, 95 vulva, ventral, 96 schematic course
of internal duct system). Scale lines: 1 mm. bp = projection at base of embolus; C = conductor; CO = copulatory opening; Cod =
copulatory opening depression; dRTA = dorsal branch of RTA; DTA = dorsal tegular apophysis; E = embolus; ed = encapsulated
part of internal ducts; FD = fertilization duct; fw = first winding of copulatory duct; LL = lateral lobe; MS = median septum; PTA
= prolateral tibial apophysis; RdP = retrodistal tegular protrusion; RpP = retroproximal cymbial projection; STlb = subtegular
locking lobe; vRTA = ventral branch of RTA; VTA = ventral tibial apophysis.

Female (paratype): Coloration pattern as in male, slightly darker. Legs with faint gray marks at the base
of spines (Figs 87−88). Total length 7.88. Prosoma: long, 3.08 wide. Opisthosoma: 4.61 long, 3.53 wide. Eyes:
diameters: 0.20, 0.23, 0.20, 0.32; interdistances: 0.47, 0.48, 0.69, 0.64, 0.65, 0.67. Legs (1243) I: 18.22 (4.61, 1.46,
5.58, 4.68, 1.89); II: 14.30 (4.04, 1.43, 4.18, 3.33, 1.32); III: 10.77 (3.16, 1.15, 2.87, 2.56, 1.03); IV: 12.56 (3.66,
1.12, 3.22, 3.20, 1.36). Spination follows the generic pattern, except femur III: r1-0-1; patella: d0-0-1; tibia: d1-0-
1. Epigyne: EF slightly longer than wide; LL fused; pair of elongate COd with CO situated posteriorly; MS small,
triangular, two times longer than wide (Figs 89, 94). Vulva: CD with first winding gently curving antero-laterad;
encapsulated part of internal ducts two times longer than wide, slightly more dilated posteriorly; FD anteriad (Figs
90, 95−96).
Distribution. Only known from the type locality in Colombia (Fig. 143).
Nativus tawu sp. nov.

Figs 97−111
Type Material: Holotype: COLOMBIA: Vaupés: 1♂, Mitú, Casa de Don Luis (1.3965, ‑70.0377), 188 m, 21
September 2019, N. Hazzi & L. Martínez leg. (AvH-I-3207). Paratypes: COLOMBIA: Vaupés: 1♀, same data as
holotype (AvH-I-3246).
Etymology. The specific name refers to the táwü masks used during the ónye, a ceremony of mourning
performed by the Cubeo indigenous tribe inhabiting the Vaupés River basin, especially along its tributaries Caduyarí
and Querarí; noun in apposition.
Diagnosis. Males of N. tawu sp. nov. resemble those of N. carare sp. nov. (Figs 16−18) and N. mariua sp.
nov. (Figs 59−61) by the palps with E straight and tegulum lacking RdP (Fig. 90). They are distinguished from N.
carare sp. nov. by the dRTA bifid, with two pointed branches and tegulum with triangular protrusion (tp) close to
bp (Figs 107−108) (dRTA single and tegulum lacking small tp close to bp in N. carare sp. nov.) and from N. mariua
sp. nov. by the bp irregularly-shaped, slightly longer than wide and dRTA with ventral branch curved in retrolateral
view (Figs 107−108) (bp keel-like, wider than long and dRTA with ventral branch straight in retrolateral view in N.
mariua sp. nov.). Females resemble those of N. mariua sp. nov. (Figs 62−64) by the LL fused medially with pair
of anterior COd bearing posterior CO and large, posterior MS. The new species can be distinguished by the MS
rectangular, wider than long, with pair of antero-lateral moon-shaped pockets, and fused part of LL shorter than
anterior COd (Fig. 109) (MS trapezoid, as long as wide and fused part of LL as long as anterior COd in the latter
species).
Description. Male (holotype): Prosoma orange with brown margins and thoracic striae; cephalic region
with longitudinal stripes extending posteriorly from behind PME and PLE; fovea dark brown; eye borders black.
Chelicerae orange brown. Legs and palps orange brown, slightly lighter than prosoma. Endites pale brown, distally
lighter. Labium orange brown, distally paler brown. Sternum pale yellow, with dark orange margins. Opisthosoma
pale yellowish cream colored; dorsally with two rounded, brown muscle impressions; ventrally with U-shaped lines
of muscle impressions. Spinnerets pale yellow (Figs 97−98). Total length 6.85. Prosoma: 3.27 long, 3.38 wide.
Opisthosoma: 3.26 long, 2.09 wide. Eyes diameters: 0.22, 0.24, 0.23, 0.33; interdistances: 0.46, 0.47, 0.71, 0.65,
0.71, 0.70. Legs (2143): I: 21.89 (3.82, 1.26, 4.42, 4.23, 1.81); II: 26.44 (4.89, 1.30, 5.54, 4.96, 2.14); III: 16.79
(3.72, 1.39, 3.37, 3.40, 1.32); IV: 21.18 (4.33, 1.06, 4.06, 4.34, 1.90). Spination follows the generic pattern, except
femur IV: r0-1-1; tibiae I−II: p1-0-1, d1-1-1, r1-0-1, v2-2-0; tibia III: d1-0-1. Palp: PTA distally rounded, as wide
as long; VTA reduced; vRTA two times longer than wide in retrolateral view; dRTA bifid with dorsal branch wide,
concave at ventral margin, and ventral branch long and hook-like; tegulum with small triangular tegular protrusion,
close to bp; DTA with no projections (Figs 99−101, 106−108).

FIGURES 97−101. Nativus tawu sp. nov., male (AvH-I-3207). 97−98 Habitus (97 dorsal, 98 ventral); 99−101 Left palp (99
prolateral, 100 ventral, 101 retrolateral). Scale lines 97−98 = 2mm; 99−101 = 1 mm.
Female (paratype): Prosoma orange brown, slightly more reddish between thoracic striae, with brown margins;
cephalic region with brown stripes extending posteriorly from behind PLE. PME and between AME; thoracic striae
and fovea brown; eye borders black. Chelicerae pale yellowish brown, distally lighter. Labium brown, distally
yellowish brown. Sternum yellowish cream colored with dark orange margins. Opisthosoma grayish brown; dorsally
with few scattered white marks laterally on anterior half and two paisr of rounded brown muscle impressions;

ventrally with scattered white marks laterally and V-shaped lines of muscle impressions. Spinnerets yellowish
orange, distally lighter (Figs 102−103). Total length: 9.07. Prosoma: 3.42 long, 3.27 wide. Opisthosoma: 4.99 long,
3.32 wide. Eyes: diameters: 0.22, 0.24, 0.22, 0.32; interdistances: 0.47, 0.52, 0.71, 0.70, 0.63, 0.68. Legs (2143):
Legs (2143): I: 15.00 (4.30, 1.58, 4.13, 3.45, 1.54); II: 10.71 (4.31, 1.65, 5.54, 3.39, 1.28; III: 11.78 (3.50, 1.41, 2.89,
2.88, 1.10); IV: 12.75 (3.68, 1.13, 3.23, 3.40, 1.31). Spination follows the generic pattern, except tibia III: d0-0-1.
Epigyne: EF wider than long; MS with transverse striations (Figs 104, 109). Vulva: CD broad, with first winding
posteriad and strongly curving anteriad; encapsulated part of internal ducts three times longer than wide, anteriorly
bent mediad and posteriorly dilated; FD antero-laterad (Figs 105, 110−111).
Distribution. Only known from the type locality, in Vaupés, eastern Colombia (Fig. 142).
FIGURES 102−105. Nativus tawu sp. nov., female (AvH-I-3207). 102−103 Habitus (102 dorsal, 103 ventral); 104−105
Epigyne/vulva (104 ventral, 105 dorsal). Scale lines: 102−103: 2mm; 104−105 = 1 mm.

FIGURES 106−111. Nativus tawu sp. nov. 106−108 Male (AvH-I-3207), left palp (106 prolateral, 107 ventral, 108 retrolateral);
109−111 Female (AvH-I-3207), epigyne/vulva (109 ventral, 110 dorsal, 111 schematic course of internal duct system). Scale
lines: 1 mm. bp = projection at base of embolus; C = conductor; CO = copulatory opening; Cod = copulatory opening depression;
dRTA = dorsal branch of RTA; DTA = dorsal tegular apophysis; E = embolus; ed = encapsulated part of internal ducts; ep =
epigynal pocket; FD = fertilization duct; fw = first winding of copulatory duct; LL = lateral lobe; MS = median septum; PTA =
prolateral tibial apophysis; RpP = retroproximal cymbial projection; STlbn = subtegular locking lobe; tp = tegular protrusion;
vRTA = ventral branch of RTA.

Nativus tupana sp. nov.
Figs 1−10, 112−126
FIGURES 112−116. Nativus tupana sp. nov., male (IBSP 8828). 112−113 Habitus (112 dorsal, 113 ventral); 114−116 Left palp
(114 prolateral, 115 ventral, 116 retrolateral). Scale lines: 112−113 = 2 mm; 114−116 = 1 mm.
Type material. Holotype: BRAZIL: Amazonas: ♂, Manaus, Reserva de Campina [‑2.4000, ‑59.8667], 31 January
1994, T. Gasnier leg. (IBSP 43567). Paratypes: BRAZIL: Amazonas: 1♀, same data as for holotype (IBSP 43567);
1♀, Borba, Rio Mapiá [‑4.3833, ‑59.5833], 22 April 1996, Equipe IBSP/SMNK leg. (IBSP 8824); 3 ♂, 7 ♀, 1
juv., same data as previous specimen (IBSP 8831, IBSP 8828); 3 ♂ 3 ♀, Rio Preto da Eva [‑2.6983, ‑59.6977],

7−26 January 2015, E. C. Galvão leg. (IBSP 279452, IBSP 279459, IBSP 279462, IBSP 279470, IBSP 279461,
IBSP 279480); 2 ♀, same locality as for previous specimens, January 2016, E.C. Galvão leg. (IBSP 257691, IBSP
257697); 1♀, Manaus, Fazenda Dimona, BR 174 (ZF3) [‑2.3330, ‑60.0640], 16 May 2006, F.B. Baccaro et al. leg.
(INPA 5907); 1♀, Manaus, Reserva Colosso, Reservas do PDBFF [‑2.4000, ‑59.8667], 28 February 2002, F.N.N.
Rego leg. (IBSP 97273); 1♀, Manaus, Reserva Florestal Adolpho Ducke [‑2.9000, ‑60.0833], 1995, C. Martius leg.
(IBSP 14517); 1♂, same locality as for previous specimen, 2000, E.P. Fagundes leg. (IBSP 245997); 1♀, Presidente
Figueiredo, Usina Hidrelétrica de Balbina, Ilha Abusado (‑1.7616, ‑59.6784), 25−27 June 2016, D. Storck-Tonon
leg. (IBSP 231172); 1♂, same locality as previous specimen (-1.5586, -59.8497), 3−5 July 2016, D. Storck-Tonon
leg. (IBSP 231278); 1♂, same locality as for previous specimem, Ilha Beco do Catitu, 26−28 June 2016, D. Storck-
Tonon leg. (IBSP 250186).
FIGURES 117−120. Nativus tupana sp. nov., female (IBSP 8828). 117−120 Habitus (117 dorsal, 118 ventral); 119−120
Epigyne/vulva (119 ventral, 120 dorsal). Scale lines: 117−118 = 2 mm, 119−120 = 1 mm.
Etymology. The specific name refers to Tupana, mother of thunder, revered by the early Indian inhabitants of
the region of Borba, before the arrival of the Portuguese Jesuits; noun in apposition.

FIGURES 121−126. Nativus tupana sp. nov. 121−123 Male (IBSP 8828), left palp (121 prolateral, 122 ventral, 123 retrolateral);
124−126 Female (AMNH), epigyne/vulva (124 epigyne, ventral, 125 vulva, dorsal, 126 schematic course of internal duct
system). Scale lines: 1 mm. bp = projection at base of embolus; C = conductor; CO = copulatory opening; Cod = copulatory
opening depression; dRTA = dorsal branch of RTA; DTA = dorsal tegular apophysis; E = embolus; ed = encapsulated part of
internal ducts; FD = fertilization duct; fw = first winding of copulatory duct; LL = lateral lobe; mp = median projection of
the DTA; MS = median septum; PTA = prolateral tibial apophysis; RdP = retrodistal tegular protrusion; RpP = retroproximal
cymbial projection; STlbn = subtegular locking lobe; vRTA = ventral branch of RTA.

Diagnosis. Males of N. tupana sp. nov. resemble those of N. hazzii sp. nov. (Figs 28−30), N. juruti sp. nov. (Figs
47−49) and N. nocaima sp. nov. (Figs 91−93) by the palps with tegulum bearing small, wider than long RdP and
E sinuous. They are distinguished from the three latter species by the presence of a large median projection on the
DTA (Figs 121−122) (DTA without median projection in the other species). Females resemble those of N. nocaima
sp. nov. (Figs 94−96) by the epigyne with LL in roughly an inverted T-shape, with pair of elongate COd situated
anteriorly. It is distinguished from the latter species by the by the COd J-shaped and separated from each other by
almost MS width (Fig. 124) (C-shaped and separated from each other by half MS width in the latter species).
Description. Male (IBSP 8828, paratype): Prosoma brown with darker brown margins and thoracic striae;
cephalic region with darker brown stripes extending posteriorly from PME and between AME; fovea dark brown;
eye borders black. Chelicerae, legs and palps pale brown. Endites pale orange brown, distally lighter. Labium
brown, distally orange brown. Sternum pale yellow with dark orange margins. Opisthosoma pale gray; dorsally with
two pairs of brown, rounded muscle impressions; ventrally with faint dark gray longitudinal band and two roughly
parallel lines of muscle impressions. Spinnerets pale yellow (Figs 112−113). Total length 6.1. Prosoma: 3.0 long, 2.9
wide. Opisthosoma: 3.0 long, 1.8 wide. Eyes: diameters: 0.22, 0.26, 0.20, 0.28; interdistances: 0.06, 0.02, 0.24, 0.22,
0.20, 0.20. Legs (2143): I: 19.7 (4.9, 1.4, 5.5, 5.4, 2.5); II: 23.2 (5.6, 1.5, 6.7, 6.6, 2.8); III: 15.5 (4.1, 1.2, 4.0, 4.6,
1.6); IV: 18.7 (4.9, 1.2, 4.8, 5.5, 2.3). Palp: PTA roughly squared, distally rounded, as wide as long; VTA reduced,
absent; vRTA two times longer than wide; dRTA with two pointed branches, the dorsal one two times longer than
ventral one; tegulum with RdP roughly 1.5 times wider than long; bp subsquared, as wide as long (Figs 114−116,
121−123).
Female (IBSP 8828, paratype): Coloration pattern as in male except prosoma, with brown reticulated pattern
laterally and brown marks between thoracic striae (Figs 117−118). Total length 6.7. Prosoma: 2.6 long, 2.7 wide.
Opisthosoma: 3.8 long, 2.4 wide. Eyes: diameters: 0.21, 0.27, 0.20, 0.30; interdistances: 0.10, 0.03, 0.26, 0.28, 0.27,
0.24. Epigyne: EF slightly wider than long; LL fused; pair of elongate COd situated anteriorly bearing posterior CO;
MS situated posteriorly to fused part of LL, trapezoidal, almost three times wider than long (Figs 119, 124). Vulva:
CD with first winding laterad; encapsulated part of internal ducts two times longer than wide, slightly more dilated
posteriorly; FD antero-laterad (Figs 120, 125−126).
Variation. Males (n = 9): Total length 5.6−6.7; prosoma length 2.5−3.3; femur I length 4.6−4.9. Females (n =
10): Total length 5.5−7.8; prosoma length 2.5−3.2; femur I length 3.1−3.8.
Distribution. Known from the state of Amazonas, Brazil (Fig. 143).
Nativus yurupari sp. nov.

Figs 127−141
Type Material: Holotype: COLOMBIA: Vaupés: ♂, Mitú, Casa de Don Luis (1.3965, ‑70.0377), 188 m, 21
September 2019, N. Hazzi & L. Martínez leg. (IAvH-I-3156). Paratypes: COLOMBIA: Vaupés: 1♀, same data as
for holotype (IAvH-I-3136); 2 ♂, same data as for holotype (IAvH-I-3126); Amazonas: 1♀, Leticia, Monilla Amena
Km 9 (-4.0837, ‑69.9011), 60 m, manual capture, October 2003, A. Gnaphos leg. (MPUJ_ENT0087070); 1♀, same
locality as for previous specimen, 1 October 2003, Alvarez leg. (MPUJ_ENT0087070).
Additional material examined: COLOMBIA: Amazonas: 1♀, Leticia, Comunidad Monilla Amena (-4.1121,
-69.9280), 60 m, 3 May 2002, J. Jimenez leg. (MPUJ_ENT0086787); 1♀ same locality as the previous specimen
(-4.1121, -69.9280), 6 November 2005, C.Trejo leg. (MPUJ_ENT0086788); 1♀, Leticia, Comunidad Monilla Amena
km 9 (‑4.0837, -69.9011), 1 October 2003, Alvarez leg. (MPUJ_ENT0086797); Leticia, Km 10 Vía Tarapacá, Finca
la Novedosa, 95 m, 23 September 2003, Estudiantes de Zoología Animal leg. (ICN-Ar-2377).
Etymology. The specific name refers to the Yuruparí myth, considered fundamental in the traditional culture
of some Amazonian groups. It narrates about the origins of the Tenui people, conflicts between its inhabitants and
the arrival of Yuruparí (meaning son of the fruit). Currently, this myth constitutes the basis of dances or ceremonies
that are celebrated by several indigenous communities settled in the Vaupés department (e.g. Tukanos); noun in
apposition.

FIGURES 127−131. Nativus yurupari sp. nov., male (IAvH-I-3156). 127−128 Habitus (127 dorsal, 128 ventral); 129−131 Left
palp (129 prolateral, 130 ventral, 131 retrolateral). Scale lines 127−128 = 2mm; 129−131 = 1 mm.
Diagnosis. Males of N. yurupari sp. nov. resemble those of N. janai sp. nov. (Figs 39−41), N. napo sp. nov.
(Figs 74−76) by the palps with tegulum with large triangular RdP, at least 1.5 times longer than wide and E sinuous
in the male palps. It is distinguished from N. janai sp. nov. by the bp keel-like, two times wider than long (Fig. 137)
(irregularly-shaped, as long as wide with anterior margins depressed in N. janai sp. nov.) and from N. napo sp. nov.
by the dRTA bifid (Figs 137−138) (single in N. napo sp. nov.). Females resemble those of N. napo sp. nov. (Figs
77−79) by the MS diamond shaped, anterior to fused part of LL. They are distinguished from the latter species by
the fused part of LL extending until almost half EF length and MS two times longer than wide (Fig. 139) (fused part
of LL extending until 1/3 of EF length and MS roughly as wide as long in the latter species).

FIGURES 132−135. Nativus yurupari sp. nov., female (IAvH-I-3126). 132−133 Habitus (132 dorsal, 133 ventral); 134−135
Epigyne/vulva (134 ventral, 135 dorsal). Scale lines: 132−133 = 2mm; 134−135 = 1 mm.
Description. Male (holotype): Prosoma orange-brown, with brown lateral margins and thoracic striae; cephalic
region with brown lines extending posteriorly from PME and between AME; fovea dark brown; eye borders black.
Chelicerae, legs and palps orange brown, lighter than prosoma. Endites pale yellowish brown, distally lighter.
Labium orange brown, distally brown. Sternum yellowish cream colored with orange margins. Opisthosoma grayish
cream colored; dorsally with two pairs of brown rounded muscle impressions; ventrally with two roughly parallel
lines of muscle impressions. Spinnerets cream colored (Figs 127−128). Total length 5.51. Prosoma: 2.6 long, 2.84
wide. Opisthosoma: 2.76 long, 1.84 wide. Sternum: 1.32 long, 1.54 wide. Eyes diameters: 0.15, 0.17, 0.28, 0.36;
interdistances: 0.46, 0.47, 0.67, 0.63, 0.59, 0.65. Legs (2143): I: 17.44 (4.38, 1.22, 5.08, 4.9, 1. 86); II: 20.96 (5.42,
1.02, 5.87, 6.05, 2.6); III: 13.01 (3.61, 0.81, 3.94, 3.34, 3.84, 1.41); IV: 17.27 (4.9, 0.94, 3.78, 5.55, 2.1). Spination
follows the generic pattern, except femora III−IV: p1-1-1, d0-1-1, r0-1-1; tibiae I−II: p1-0-1, d1-1-1, r1-0-1, v2-2-
0; tibia III: p1-0-1, d1-0-1, r1-0-1, v2-2-0; tibia IV: p1-0-1, d0-0-1, r1-0-1, v2-2-0. Palp: PTA slightly longer than
wide; VTA triangular, displaced retrolaterally; vRTA 1.5 times longer than wide in retrolateral view; dRTA bifid
with dorsal branch trapezoidal and ventral branch conical, distally pointed; RdP slightly over 2 times longer than
wide (Figs 129−131, 136−138).

FIGURES 136−141. Nativus yurupari sp. nov. 136−138 Male (IAvH-I-3156), left palp (136 prolateral, 137 ventral, 138
retrolateral); 139−141 Female (IAvH-I-3126), epigyne/vulva (139 ventral, 140 dorsal, 141 schematic course of internal duct
system). Scale lines: 1 mm. bp = projection at base of embolus; C = conductor; CO = copulatory opening; dRTA = dorsal branch
of RTA; DTA = dorsal tegular apophysis; E = embolus; ed = encapsulated part of internal ducts; FD = fertilization duct; fw =
first winding of copulatory duct; LL = lateral lobe; MS = median septum; PTA = prolateral tibial apophysis; RdP = retrodistal
tegular protrusion; RpP = retroproximal cymbial projection; STlb = subtegular locking lobe; vRTA = ventral branch of RTA;
VTA = ventral tibial apophysis.

Female (paratype, IAvH-I-3136): Prosoma brown with brown thoracic striae and dark brown margins;
cephalic region with longitudinal stripes extending posteriorly from PME and between AME; fovea dark brown;
eye borders black. Chelicerae brown, as prosoma. Legs and palps brown, slightly lighter than prosoma. Labium
and endites orange brown, distally slightly lighter. Sternum pale orange brown with brown margins. Opisthosoma
yellowish brown; dorsally with two pairs of rounded, brown muscle impressions; ventrally with V-shaped lines
of muscle impressions. Spinnerets orange brown (Figs 132−133). Total length: 7.11. Prosoma: 2.60 long, 3.00
wide. Opisthosoma: 4.11 long, 2.51 wide. Eyes diameters: 0.23, 0.27, 0.27, 0.30; interdistance: 0.49, 0.49, 0.72,
0.66, 0.65, 0.71. Legs (2143): Legs (2143): I: 14.58 (3.93, 1.38, 4.23, 3.41, 1.63); II: 15.71 (4.05, 1.60, 4.61, 3.96,
1.49); III: 11.90 (3.38, 1.15, 3.08, 3.11, 1.18); IV: 14.45 (4.12, 1.14, 3.53, 3.98, 1.68). Spination follows the generic
pattern, except femora II−III: r1-0-1; femur IV: p1-1-1, d0-1-1, r0-0-1; tibiae I−II: p1-0-1, r1-0-1, v2-2-0; tibiae
III−IV: d1-0-1; metatarsus IV: p1-1-2, r1-1-2, v1-1-2. Epigyne: EF as long as wide; MS with pair of CO situated
anteriorly (Figs 134, 139) Vulva: CD with first winding curving antero-laterad; encapsulated part of internal ducts
two times longer than wide, dilated posteriorly; FD antero-laterad (Figs 135, 140−141).
FIGURE 142. Record map for Nativus carare sp. nov., N. hazzii sp. nov., N. juriti sp. nov., N. janai sp. nov. and N. tawu sp.
nov.
Distribution. Only known from the type locality (Fig. 143).

FIGURE 143. Record map for Nativus mariua sp. nov., N. napo sp. nov., N. nocaima sp. nov., N. tupana sp. nov. and N.
yurupari sp. nov.
Discussion
The monophyly of Heteropodinae is well substantiated and it is consistently recovered in molecular analyses
(Agnarsson & Rayor 2013, Moradmand et al. 2014, Wheeler et al. 2017, Gorneau et al. 2022). The subfamily is
characterized by the combination of several morphological characters. Nevertheless, only the long-toothed female
palpal claw seems to be exclusive for the group. Other characters, such as eye arrangement, number of cheliceral teeth
and presence of intermarginal denticles can appear independently throughout the family. For example, intermarginal
denticles can be found in one species of the genus Eusparassus (see Moradmand & Jäger 2012) and in the genera
Caayguara (see Rheims 2010b) and Sadala (see Rheims & Jäger 2022), among others. However, they all have a
short-toothed palpal claw.
Currently, Heteropodinae is comprised of twenty genera (World Spider Catalog 2023). Of these, Nativus gen.
nov. seems to be more closely related to the Neotropical members of the subfamily, Anaptomecus, Guadana and
Sparianthina. All four genera have only two pairs of ventral spines on tibiae I−II (see Jäger et al. 2009, Rheims
2010a, Guala et al. 2012), while the remaining Heteropodinae have three pairs. Nativus gen. nov., Guadana and
Sparianthina seem to be more closely related to each other, sharing the presence of intermarginal denticles along
the entire cheliceral groove, a trilobate membrane with median hook slightly smaller than lateral projections and a
strong dorsal tegular apophysis (DTA) on the male palps. Anaptomecus, on the other hand, has clustered denticles,
trilobate membrane with median hook as large as or slightly larger than lateral projections and lacks the DTA on the
male palps, as most of the other Heteropodinae.

Considering Guadana and Sparianthina, Nativus gen. nov. is probably more closely related to Guadana. Both
genera share male palps with a vRTA and a sickle-shaped DTA, and a female vulva with short copulatory ducts and
lacking glandular projections. Nativus gen. nov. is here considered a monophyletic unit, separate from Guadana,
due to the presence of a filiform embolus with hyaline basal projection in the male palps and medially to medial-
posteriorly fused lateral lobes in the female epigyne. Nevertheless, a cladistic analysis, including representatives of
all the subfamily, will be a welcome confirmation of the generic status herein proposed.
Acknowledgements
This study was supported by Fundação de Amparo à Pesquisa de São Paulo (FAPESP no. 2011/18694-3; 2022/12588-
1 to CAR). We wish to thank Beatriz Mauricio (Laboratório de Biologia Celular, Instituto Butantan. São Paulo)
for helping with SEMs, and thanks to Leonel Martinez (Universidad del Atlántico) and Nicolas Hazzi (George
Washington University) for collecting most of the material described here. We are grateful to J.C. Neita and Julian
Clavijo (both latter from Instituto Alexander von Humboldt), Daniela Martinez and William Galvis (both latter from
Instituto de Ciencias Naturales, Universidad Nacional de Colombia) who kindly allowed and facilitated the revision
of the material referenced herein. Also to J. Fagua, D. Forero and M. Rodríguez of the Museo Javeriano de Historia
Natural Lorenzo Uribe, Universidad Pontificia Javeriana, Bogotá, Colombia. This research also benefited from
the support of the Instituto Alexander von Humboldt, especially in the provision of curatorial materials, and the
Universidad del Atlántico (Barranquilla), through the Convocatoria para el Fortalecimiento de las Instituciones de
Educación Superior (Convocatoria 890). CC owes further thanks to Eduardo Villarreal, Andrea Jiménez, and Karla
Marimon for their support and revision of the early version of the manuscript. The final version of this manuscript
greatly benefited from the comments of two anonymous reviewers and Dr. Christoph Muster (editor for Zootaxa)
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Zootaxa 5360 (1): 001–043 ISSN 1175-5326 (print edition)

Key words: Taxonomy, Amazon, Brazil, Colombia, Peru

Accepted by C. Muster: 28 Sept. 2023; published: 25 Oct. 2023 

Material and methods

 · Zootaxa 5360 (1) © 2023 Magnolia Press Casas & Rheims

Nativus gen. nov.

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Identification key for species of Nativus gen. nov.

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Nativus carare sp. nov.

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Nativus janai sp. nov.

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NATIVUS gen. nov. Zootaxa 5360 (1) © 2023 Magnolia Press · 13

Nativus juriti sp. nov.

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Nativus napo sp. nov.

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Nativus nocaima sp. nov.

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Nativus tawu sp. nov.

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Nativus yurupari sp. nov.

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