110 - Avicularia, Zmiany 2017

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A peer-reviewed open-access journal

ZooKeys 659: 1–185 (2017)


Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 1
doi: 10.3897/zookeys.659.10717 MONOGRAPH
http://zookeys.pensoft.net Launched to accelerate biodiversity research

Taxonomic revision and cladistic analysis of Avicularia


Lamarck, 1818 (Araneae, Theraphosidae, Aviculariinae)
with description of three new aviculariine genera

Caroline Sayuri Fukushima1, Rogério Bertani1

1 Laboratório Especial de Ecologia e Evolução, Instituto Butantan, Av. Vital Brazil, 1500 CEP 05503-900,
São Paulo, Brazil

Corresponding author: Caroline Sayuri Fukushima ([email protected])

Academic editor: I. Agnarsson | Received 4 October 2016 | Accepted 2 February 2017 | Published 2 March 2017

http://zoobank.org/79A6393D-8021-41B8-BF1A-2A3723AFECFB

Citation: Fukushima CS, Bertani R (2017) Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818
(Araneae, Theraphosidae, Aviculariinae) with description of three new aviculariine genera. ZooKeys 659: 1–185.
https://doi.org/10.3897/zookeys.659.10717

Abstract
The genus Avicularia Lamarck, 1818 is revised and all species are rediagnosed. The type species, described
as Aranea avicularia Linnaeus, 1758, is the oldest mygalomorph species described and its taxonomic his-
tory is extensive and confusing. Cladistic analyses using both equal and implied weights were carried out
with a matrix of 46 taxa from seven theraphosid subfamilies, and 71 morphological and ecological char-
acters. The optimal cladogram found with Piwe and concavity = 6 suggests Avicularia and Aviculariinae
are monophyletic. Subfamily Aviculariinae includes Avicularia Lamarck, 1818, Typhochlaena C. L. Koch,
1850, Tapinauchenius Ausserer, 1871, Stromatopelma Karsch, 1881, Ephebopus Simon, 1892, Psalmopoeus
Pocock, 1895, Heteroscodra Pocock, 1899, Iridopelma Pocock, 1901, Pachistopelma Pocock, 1901, Ybyra-
pora gen. n., Caribena gen. n., and Antillena gen. n. The clade is supported by well-developed scopulae
on tarsi and metatarsi, greatly extended laterally. Avicularia synapomorphies are juveniles bearing black
tarsi contrasting with other lighter articles; spermathecae with an accentuated outwards curvature medi-
ally, and male palpal bulb with embolus medial portion and tegulum’s margin form an acute angle in
retrolateral view. Avicularia is composed of twelve species, including three new species: Avicularia avicu-
laria (Linnaeus, 1818), Avicularia glauca Simon, 1891, Avicularia variegata (F. O. Pickard-Cambridge,
1896) stat. n., A. minatrix Pocock, 1903, Avicularia taunayi (Mello-Leitão, 1920), Avicularia juruensis
Mello-Leitão, 1923, Avicularia rufa Schiapelli & Gerschman, 1945, Avicularia purpurea Kirk, 1990, A.
hirschii Bullmer, Thierer-Lutz & Schmidt, 2006, Avicularia merianae sp. n., A. lynnae sp. n., and A. caei
sp. n. Avicularia species are distributed throughout Mexico, Costa Rica, Panama, Trinidad and Tobago,

Copyright Caroline S. Fukushima, Rogério Bertani. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
2 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Venezuela, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Bolivia, and Brazil. Three new
genera are erected to accommodate former Avicularia species: Caribena gen. n., composed of Caribena la-
eta (C. L. Koch, 1842), comb. n. and Caribena versicolor (Walckenaer, 1837), comb. n.; Antillena gen. n.,
with a single species, Antillena rickwesti (Bertani & Huff, 2013), comb. n., both from the Caribbean; and
Ybyrapora gen. n., composed of Ybyrapora sooretama (Bertani & Fukushima, 2009), comb. n., Ybyrapora
gamba (Bertani & Fukushima, 2009), comb. n. and Ybyrapora diversipes (C. L. Koch, 1842), comb. n.
from Brazilian rainforest. The subspecies Avicularia avicularia variegata F. O. Pickard-Cambridge, 1896
is elevated to species status, resulting in the combination Avicularia variegata (F. O. Pickard-Cambridge,
1896) stat. n. The following new synonymies are established: Avicularia velutina Simon 1889, Avicularia
exilis Strand, 1907, Avicularia ancylochyra Mello-Leitão, 1923, Avicularia cuminami Mello-Leitão, 1930,
and Avicularia nigrotaeniata Mello-Leitão, 1940 are junior synonyms of A. avicularia; Avicularia bicegoi
Mello-Leitão, 1923 is a junior synonym of A. variegata stat. n., and Avicularia urticans Schmidt, 1994 is a
junior synonym of Avicularia juruensis Mello-Leitão, 1923. Species transferred to other genera: Avicularia
affinis (Nicolet, 1849) is transferred to Euathlus Ausserer, 1875, making the new combination Euathlus
affinis (Nicolet, 1849), comb. n.; Avicularia subvulpina Strand, 1906 is transferred to Grammostola Si-
mon, 1892, making the new combination Grammostola subvulpina (Strand, 1906), comb. n.; Avicularia
aymara (Chamberlin, 1916) is transferred to Thrixopelma Schmidt, 1994, making the new combination
Thrixopelma aymara (Chamberlin, 1916), comb. n.; Avicularia leporina (C. L. Koch, 1841) and Avicularia
plantaris (C. L. Koch, 1842) are transferred to Iridopelma Pocock, 1901, making the new combinations
Iridopelma leporina (C. L. Koch, 1841), comb. n. and Iridopelma plantaris (C. L. Koch, 1842), comb. n.;
the two last species are considered nomina dubia. The following species are considered nomina dubia:
Avicularia hirsutissima (C. L. Koch, 1842) nomen dubium; Ischnocolus hirsutum Ausserer, 1875 nomen
dubium; Ischnocolus gracilis Keyserling, 1891 nomen dubium; Avicularia arabica (Strand, 1908) nomen
dubium; Araneus hirtipes (Fabricius, 1787) nomen dubium; Avicularia ochracea (Perty, 1833) nomen du-
bium; Avicularia walckenaerii (Perty, 1833) nomen dubium; Avicularia testacea (C. L. Koch, 1841) nomen
dubium; Avicularia detrita (C. L. Koch, 1842) nomen dubium; Ischnocolus doleschalli Ausserer, 1871 no-
men dubium; Avicularia metallica Ausserer, 1875 nomen dubium; Avicularia rapax (Ausserer, 1875) nomen
dubium; Avicularia holmbergi Thorell, 1890 nomen dubium; Avicularia aurantiaca Bauer, 1996 nomen
dubium; Avicularia azuraklaasi Tesmoingt, 1996 nomen dubium; Avicularia huriana Tesmoingt, 1996 no-
men dubium; Avicularia ulrichea Tesmoingt, 1996 nomen dubium; Avicularia braunshauseni Tesmoingt,
1999 nomen dubium; Avicularia geroldi Tesmoingt, 1999 nomen dubium; Avicularia soratae Strand, 1907
nomen dubium; Avicularia fasciculata Strand, 1907 nomen dubium; Avicularia fasciculata clara Strand,
1907 nomen dubium; and Avicularia surinamensis Strand, 1907 nomen dubium. Avicularia vestiaria (De
Geer, 1778) is considered nomen nudum. Keys are provided for identification of all aviculariine genera, as
well as to Avicularia, Caribena gen. n., Ybyrapora gen. n. and Antillena gen. n. species. Maps with records
and information on species habitat are also presented. Lectotypes are newly designed for A. avicularia,
A. variegata stat. n., A. juruensis, C. laeta comb. n., E. affinis comb. n. and a neotype is established for
C. versicolor comb. n.

Keywords
Birdspider, Linnaeus, morphology, Mygalomorphae, systematics, tarantula
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 3

Table of contents
Introduction.............................................................................................................. 4
Material and methods............................................................................................. 10
Cladistic analysis..................................................................................................... 11
Taxonomy............................................................................................................... 28
Family Theraphosidae Thorell, 1869.................................................................... 28
Subfamily Aviculariinae Simon, 1889................................................................... 28
Genus Avicularia Lamarck, 1818....................................................................... 31
Avicularia avicularia (Linnaeus, 1758)........................................................... 41
Avicularia glauca Simon, 1891....................................................................... 60
Avicularia variegata (F. O. Pickard-Cambridge, 1896), stat. n........................ 61
Avicularia minatrix Pocock, 1903................................................................... 70
Avicularia taunayi (Mello-Leitão, 1920)......................................................... 76
Avicularia juruensis Mello-Leitão, 1923.......................................................... 80
Avicularia rufa Schiapelli & Gerschman, 1945............................................... 89
Avicularia purpurea Kirk, 1990....................................................................... 98
Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006.......................... 104
Avicularia merianae sp. n.............................................................................. 109
Avicularia lynnae sp. n.................................................................................. 114
Avicularia caei sp. n...................................................................................... 117
Taxonomic remarks......................................................................................... 120
Avicularia vestiaria (De Geer, 1778), nomen nudum..................................... 120
Iridopelma leporina (C. L. Koch, 1841), comb. n., nomen dubium................ 120
Iridopelma plantaris (C. L. Koch, 1842), comb. n., nomen dubium............... 122
Euathlus affinis (Nicolet, 1849), comb. n...................................................... 122
Ischnocolus hirsutum Ausserer, 1875, nomen dubium..................................... 123
Avicularia metallica Ausserer, 1875, nomen dubium...................................... 123
Ischnocolus gracilis Keyserling, 1891, nomen dubium..................................... 123
Grammostola subvulpina (Strand, 1906), comb. n......................................... 124
Avicularia arabica (Strand, 1908), nomen dubium......................................... 124
Thrixopelma aymara (Chamberlin, 1916), comb. n....................................... 125
Avicularia aurantiaca Bauer, 1996, nomen dubium........................................ 125
Araneus hirtipes (Fabricius, 1787), nomen dubium........................................ 126
Avicularia testacea (C. L. Koch, 1841), nomen dubium................................. 126
Avicularia detrita (C. L. Koch, 1842), nomen dubium................................... 126
Avicularia hirsutissima (C. L. Koch, 1842), nomen dubium........................... 126
Avicularia holmbergi Thorell, 1890, nomen dubium...................................... 127
Ischnocolus doleschalli Ausserer, 1871, nomen dubium.................................... 127
Avicularia rapax (Ausserer, 1875), nomen dubium......................................... 127
Avicularia ochracea (Perty, 1833), nomen dubium.......................................... 128
Avicularia walckenaerii (Perty, 1833), nomen dubium................................... 128
Avicularia azuraklaasi Tesmoingt, 1996, nomen dubium............................... 129
4 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Avicularia braunshauseni Tesmoingt 1999, nomen dubium............................ 129


Avicularia geroldi Tesmoingt, 1999, nomen dubium...................................... 129
Avicularia huriana Tesmoingt, 1996, nomen dubium.................................... 129
Avicularia ulrichea Tesmoingt, 1996, nomen dubium.................................... 129
Avicularia soratae Strand, 1907, nomen dubium............................................ 129
Avicularia fasciculata Strand, 1907, nomen dubium....................................... 130
Avicularia fasciculata clara Strand, 1907, nomen dubium............................... 130
Avicularia surinamensis Strand, 1907, nomen dubium................................... 130
Caribena gen. n............................................................................................... 130
Caribena laeta (C. L. Koch, 1842), comb. n................................................. 132
Caribena versicolor (Walckenaer, 1837), comb. n.......................................... 138
Ybyrapora gen. n.............................................................................................. 144
Ybyrapora sooretama (Bertani & Fukushima, 2009), comb. n........................ 147
Ybyrapora gamba (Bertani & Fukushima, 2009), comb. n............................ 148
Ybyrapora diversipes (C. L. Koch, 1842), comb. n......................................... 151
Antillena gen. n. Bertani, Huff & Fukushima.................................................. 153
Cladistics............................................................................................................... 160
Acknowledgements................................................................................................ 170
References............................................................................................................. 171
Supplementary material 1...................................................................................... 184
Supplementary material 2...................................................................................... 184
Supplementary material 3...................................................................................... 184
Supplementary material 4...................................................................................... 185
Supplementary material 5...................................................................................... 185

Introduction
The genus Avicularia Lamarck, 1818 was erected for some species formerly included
in Mygale Latreille, 1802. The type species, described as Aranea avicularia Linnaeus,
1758, was the first mygalomorph species described. Thus, its taxonomic history is
extensive. It is also confusing; reflecting the knowledge and history of arachnology
throughout the centuries.
The original description of Avicularia is vague (Lamarck 1818), resulting in confu-
sion about which species should be included in the genus. Lamarck (1818) described
Avicularia, among other characters, as large spiders, with eight eyes in St. André’s cross
format (i. e. in an “X” shape), lacking rastellum on the chelicerae and tarsi with velvet
scopulae. Lamarck (1818) also stated that these spiders can be found in cavities on the
ground or in trees, and are mostly wandering animals, which do not build permanent
retreats unlike the “Mygale” species. He included three species in the genus, in order:
Avicularia canceridea (Latreille, 1806); Avicularia blondii (Latreille, 1804); and Avicu-
laria fasciata (Latreille, 1804). Under the name Avicularia canceridea he listed Aranea
avicularia, a species described by Linnaeus (1758) in Systema Naturae.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 5

At that time, most mygalomorph spiders were described in the genus Mygale La-
treille, 1802. The name Mygale (“les Mygales”, in French) is a non-scientific name used
by Walckenaer (Bonnet 1957) to distinguish the “mineuses” and “aviculaires” spiders
from the others (Walckenaer 1802). This name was subsequently used by Latreille
(1802) to designate a spider genus, and the first species mentioned as an example
of a Mygale species was Aranea avicularia Linnaeus, 1758. Olivier (1811) followed
Walckenaer’s system, but he only considered in Mygale what he called the “mineuses”
spiders. For the “aviculaire” spiders he used the name Aranea. Lamarck (1818) fol-
lowed Olivier (1811), naming as Mygale only “les Araignées mineuses”; for “le Araignées
aviculaires” he erected a new genus, Avicularia. Even after Lamarck (1818) erected
Avicularia, many arachnologists continued to use the name Mygale when describing
new Avicularia species. From 1833 to 1849, ten Avicularia species were originally de-
scribed as Mygale: Mygale walckenaerii Perty, 1833; Mygale ochracea Perty, 1833; Myg-
ale versicolor Walckenaer, 1837; Mygale leporina C. L. Koch, 1841; Mygale caesia C.
L. Koch, 1842; Mygale detrita C. L. Koch, 1842; Mygale diversipes C. L. Koch, 1842;
Mygale laeta C. L. Koch, 1842; Mygale plantaris C. L. Koch, 1842; and Mygale affinis
Nicolet, 1849. However, the name Mygale was unavailable; it is a junior homonym of
a mammal genus described by Cuvier (1800).
About 50 years later the name Avicularia was used again (Ausserer 1871). Ap-
parently, there was a gradual acceptance of this name together with the awareness
that the name Mygale should not be used for spiders. The reluctance in accepting the
name Avicularia proposed by Lamarck had cultural foundations. According to Smith
(2000a), the name Avicularia is derived from the earlier usage of the word by Linnaeus
(1758) when he described the species Aranea avicularia—from the latin avicula, or lit-
tle bird—plus aria, meaning "which refers to" (Beechhold 1997 apud Smith 2000a).
Thorell (1870) explained the genus name as a derivation of avicularium, meaning bird-
keeper, but in the signification adopted, bird-catcher. With the name “avicularia”
Linnaeus (1758) directly referenced the famous illustration made by Maria Sibylla
Merian (1705) in Metamorphosis Insectorum Surinamensium (Smith 2000a), in which
she drew a tarantula feeding on a bird (Fig. 1). The German word Vogelspinne is used
to refer to spiders of the infraorder Mygalomorphae and translated literally as “bird
spider”, which is likely also derived from Merian’s engraving. When Lamarck (1818)
erected the new genus, he also mentioned that these spiders could feed on little birds
on their nests, perhaps based on Merian’s illustration and explanations. The idea of a
bird-eating spider was ridiculed by Langsdorff and others at the time, who considered
it a regression to a science full of mythology and legends (Smith 2000a,b). However,
now we know that Merian drew a real predation event.
Even though Avicularia was a nomenclatorally available and valid name, only in
1928 with the publication of Opinion 104 of the International Commission on Zoo-
logical Nomeclature (ICZN 1928) was the genus Avicularia included in the Official
List of Generic Names. The Direction 67 of this same Commission (ICZN 1957) es-
tablished that the specific name of the type species of Avicularia is avicularia Linnaeus,
1758 as published in the combination Aranea avicularia.
6 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 1–2. Historical aviculariine drawings. 1 Maria Sybilla Merian’s plate from Metamorphosis insec-
torum Surinamensium (1705) depicting an Avicularia avicularia eating a bird 2 Clusius’ (1611) drawing
of a possible Pachistopelma sp., the oldest known illustration of a tarantula (Theraphosidae, Aviculariinae)
in the New World.

Some of the most renowned arachnologists have studied the genus throughout
the 19th Century and beginning of the 20th Century. Ausserer (1875) described two
new species, Avicularia rutilans Ausserer, 1875 from Colombia and Avicularia me-
tallica Ausserer, 1875 from Suriname; Thorell (1890) described Avicularia holmbergi
Thorell, 1890 from French Guiana; Simon (1889) described Avicularia velutina Si-
mon, 1889 from Venezuela and few years afterwards, Avicularia glauca Simon, 1891
from Panama (Simon 1891). Some years afterwards, F. O. Pickard-Cambridge (1896)
described Avicularia avicularia variegata F. O. Pickard-Cambridge, 1896 from Bra-
zil; Pocock (1903) described Avicularia minatrix Pocock, 1903 from Venezuela; and
Strand (1906, 1907abcd) described five more species and one subspecies of the ge-
nus—Avicularia subvulpina Strand, 1906, Avicularia exilis Strand, 1907, Avicularia
fasciculata Strand, 1907, Avicularia fasciculata clara Strand, 1907 (all from “South
America”); Avicularia soratae Strand, 1907 from Bolivia, and Avicularia surinamensis
Strand, 1907 from Suriname.
Mello-Leitão studied the genus between 1920–1945. In the revision of Brazilian
“theraphosoidea”, Mello-Leitão (1923) described three species: Avicularia ancylochyra
Mello-Leitão, 1923; Avicularia bicegoi Mello-Leitão, 1923; and Avicularia juruensis
Mello-Leitão, 1923. Some years later he described Avicularia cuminami Mello-Leitão,
1930 and Avicularia pulchra Mello-Leitão, 1933, both from Brazil; Avicularia nigrotae-
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 7

niata Mello-Leitão 1940, from Guyana; and Avicularia palmicola Mello-Leitão, 1945
also from Brazil (Mello-Leitão 1930, 1933, 1940, 1945). Subsequently, Schiapelli and
Gerschman (1945) described a new species from Brazil, Avicularia rufa Schiapelli &
Gerschman, 1945. Until 1942, Roewer’s Catalog counted 29 species and two subspe-
cies, which were distributed throughout Trinidad, Martinique and Guadeloupe Islands,
USA, Puerto Rico, Panama, Bolivia, Suriname, Colombia and Brazil (Roewer 1942).
Despite the many authors who have studied the Avicularia species, only a few
of them proposed a diagnosis for the genus. Ausserer (1871) was the first to offer a
proposal, diagnosing Avicularia as having well-developed tarsal and metatarsal scopula
and males with spines on the tibia (likely referring to male tibial apophysis). F. O.
Pickard-Cambridge (1896) diagnosed Avicularia as having an “anterior row of eyes
strongly procurved”, “legs of first pair shorter than those forth”, and “habits arboreal”.
Simon (1903) characterized the genus as having posterior lateral eye smaller than an-
terior, anterior row of eyes strongly procurved, and males with tibial apophysis oblicu-
ous, obtuse, with small spines. Mello-Leitão (1923) also published a diagnosis for the
genus, but it is mainly a Portuguese translation of Simon’s (1903) diagnosis. Valerio
(1979) diagnosed the genus as lacking stridulatory apparatus and males presenting
medial spur on tibia I.
Over the course of many years, contributions to taxonomy of Avicularia were
sparse. A significant alteration occurred only when Raven (1985) considered Eurypel-
ma C. L. Koch, 1850 as a junior synonym of Avicularia. In fact, Thorell (1870) al-
ready had considered Eurypelma C. L. Koch as a junior synonym of Avicularia, but
apparently this was overlooked by subsequent authors. Eurypelma was an old specious
unrevised genus having about 50 species that were transferred to Avicularia at once,
significantly raising the number of species in the genus.
However, Raven (1985) did not analyze all the Eurypelma species transferred, only
the type species of the genus. Thus, the identity of the species transferred to Avicularia
was not checked one by one and the taxonomic confusion was merely transferred and
added from Eurypelma to Avicularia. As the general concept of Eurypelma was signifi-
cantly distinct from Avicularia (since Eurypelma was considered formerly a grammosto-
line, now theraphosine, genus), it was clear that most of the species should not be includ-
ed in Avicularia, and a series of transferences took place thereafter. Several species were
transferred from Avicularia to Aphonopelma Pocock, 1901 (Schmidt 1993, Smith 1995),
to Brachypelma Simon, 1891 (Smith 1993, 1986; Hancock and Hancock 1989) and to
Chromatopelma Schmidt, 1995 (Schmidt 1995b). Despite this, several species formerly
included in Eurypelma still remained in Avicularia: Avicularia affinis (Nicolet, 1849);
Avicularia alticeps (Keyserling, 1878); Avicularia anthracina (C. L. Koch, 1842); Avicu-
laria aymara (Chamberlin, 1916); Avicularia borelli (Simon, 1897); Avicularia diversipes
(C. L. Koch, 1842); Avicularia doleschalli (Ausserer, 1871); Avicularia gracilis (Keyser-
ling, 1891); Avicularia guyana (Simon, 1892); Avicularia hirsuta (Ausserer, 1875); Avicu-
laria obscura (Ausserer, 1875); Avicularia ochracea (Perty, 1833); Avicularia panamensis
(Simon, 1891); Avicularia parva (Keyserling, 1878); Avicularia plantaris (C. L. Koch,
1842); Avicularia rapax (Ausserer, 1875); and Avicularia tigrina (Pocock, 1903).
8 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

In this same work, Raven (1985) synonymized the genus Ancylochirus Mello-Leitão,
1920 with Avicularia and the single species Ancylochirus taunayi Mello-Leitão, 1920
was transferred, making the new combination Avicularia taunayi (Mello-Leitão, 1920).
Only in 1990 was a new species of Avicularia described: Avicularia purpurea Kirk,
1990, from Ecuador (Kirk 1990). Since then, many other species were described such
as Avicularia urticans Schmidt, 1994, Avicularia aurantiaca Bauer, 1996, and Avicu-
laria azuraklaasi Tesmoingt, 1996, all from Peru (Schmidt 1994; Bauer 1996; Tesmo-
ingt 1996a); Avicularia recifiensis Struchen & Brändle, 1996 and Avicularia ulrichea
Tesmoingt, 1996, both from Brazil (Struchen and Brändle 1996; Tesmoingt 1996b);
Avicularia huriana Tesmoingt, 1996, from Ecuador (Tesmoingt 1996bc); Avicularia
braunshauseni Tesmoingt, 1999 and Avicularia geroldi Tesmoingt, 1999, both also
from Brazil (Tesmoingt 1999ab). At the same time, Lucas, Silva & Bertani (1992)
transferred Ephebopus violaceus Mello-Leitão, 1930 to Avicularia, making the new
combination Avicularia violacea (Mello-Leitão, 1930).
More recently, Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006 was
described from Ecuador (Bullmer et al. 2006); and two years after, Avicularia viola-
cea (Mello-Leitão, 1930) was transferred to Tapinauchenius Ausserer, 1871 by West
et al. (2008); A. obscura (Ausserer, 1875) was transferred to Ami Pérez-Miles, 2008
(Pérez-Miles et al. 2008) and Gallon (2008) considered the monotypic genus Avicusco-
dra Strand, 1908 (Strand 1908) as a junior synonym of Avicularia, making the new
combination Avicularia arabica (Strand, 1908). In 2009, Avicularia borelli (Simon,
1897) was transferred to Grammostola Simon, 1892 (Theraphosinae) (Gabriel 2009a);
Avicularia guyana (Simon, 1892) was transferred to Eupalaestrus Pocock, 1901 (Ga-
briel 2009b) and Avicularia panamensis was transferred to Sericopelma Ausserer, 1875
(Gabriel 2009b). In this same year, two new species were described from the Atlantic
Rainforest of Brazil (Bertani and Fukushima 2009): Avicularia sooretama Bertani &
Fukushima, 2009 and Avicularia gamba Bertani & Fukushima, 2009. In 2011, after a
study of all Avicularia species from Uruguay, Avicularia anthracina (C. L. Koch, 1842)
was transferred to Grammostola Simon, 1892 and Avicularia parva was transferred to
Catumiri Guadanucci, 2004 by Fukushima et. al (2011). The names Pterinopelma ti-
grinum Pocock, 1903 and Ischnocolus alticeps Keyserling, 1878, formerly belonging to
Avicularia, were considered as nomina dubia (Fukushima et al. 2011).
The last taxonomic changes in the genus were proposed by Bertani (2012), after a
revision of three Aviculariinae genera. The author considered Avicularia pulchra Mello-
Leitão, 1933 and Avicularia recifiensis Struchen & Brändle, 1996 as junior synonyms
of Pachistopelma rufonigrum Pocock, 1901, and Avicularia palmicola Mello-Leitão,
1945 as a junior synonym of Iridopelma hirsutum Pocock, 1901 (Bertani 2012).
The last Avicularia species described was Avicularia rickwesti Bertani & Huff, 2013,
a very distinct species endemic of the Dominican Republic (Bertani and Huff 2013).
The most recent diagnosis for the genus was proposed by Gallon (2008). He con-
sidered Avicularia close to the genera Pachistopelma Pocock, 1901 and Iridopelma Po-
cock, 1901 based on the presence of urticating setae type II on the abdomen. Accord-
ing to Gallon (2008), the males of Avicularia can be distinguished from males of Iri-
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 9

dopelma by lacking tibial apophysis on leg II and the females by the absence of lobes on
spermathecae. Besides, both sexes of Avicularia would have had leg IV longer than leg
I, the opposite of Iridopelma; and both sexes of Avicularia differ from Pachistopelma by
having anterior row of eyes strongly procurved, not straight. Also, according to Gallon
(2008), only Avicularia, Pachistopelma and Iridopelma would belong to the subfamily
Aviculariinae. This idea was contradicted by West et al. (2008) and Bertani (2012),
who presented cladistics analyses containing diverse related genera and proposed that
subfamily Aviculariinae is more inclusive than proposed by Gallon (2008).
Avicularia species have an arboreal habit, making their silk retreats on vegetation
and human structures (Stradling 1994). Studies with A. avicularia pointed out that de-
spite being sedentary, this species has a well-defined pattern of daily activities. During
the day, the specimens remain inside their retreats while during the night they emerge
to catch their prey (Stradling 1994).
An interesting defense mechanism was observed by Bertani and Marques (1996).
They observed that the abdominal urticating setae type II (Cooke et al. 1972) are trans-
ferred by direct contact when the spiders are disturbed, and are not expulsed in the
air as happens in theraphosines. Type II setae of Avicularia species is stouter than the
slender setae of theraphosines, preventing them from being carried by the air (Bertani
and Marques 1996). However, some years later, an interesting exception was seen,
constituting a case of convergence between aviculariines and theraphosines. Bertani et
al. (2003) observed that A. versicolor (Walckenaer, 1837) expulse the urticating setae
in a similar way of theraphosines. The type II urticating setae of this particular species
are very slender, therefore, they can be carried by air.
Detailed studies on habit, life cycle, and reproduction as well as biogeography and
conservation are practically nonexistent. In one of the few ecological studies, Stradling
(1994) analyzed the distribution and behavioral ecology of A. avicularia in Trinidad.
He carried out this study in the field for 18 months, in an abandoned tonka bean plan-
tation, resulting in the largest ecological dataset on A. avicularia and one of the most
detailed studies of aviculariine ecology. Stradling (1994) presented important conclu-
sions about distribution, dispersion, courtship and mating behavior of this species.
A more recent report on the ecology of an Avicularia species was conducted by
Maréchal et al. (2009). The authors studied the behavioral ecology and population
structure of A. versicolor over several years and suggested that this endemic species from
Martinique should be included in the CITES’s list (The Convention on International
Trade in Endangered Species of Wild Fauna and Flora).
Desco-Derouet and Gros (1972) studied weight increase and linear growth of A.
avicularia and concluded that changes in these variables cannot be used to determine
sex in juveniles. Dresco-Derouet (1970) published a short note about the development
cycle of A. avicularia in captivity. Stradling (1978) also studied the growth and matu-
ration of A. avicularia, but he did it in both laboratory and field conditions, resulting
in a more extensive and accurate work.
Regarding applied studies using the Avicularia species as a model, an antifungal
peptide from the venom of the A. rufa (misidentified as A. juruensis) was discovered
10 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

(Ayroza et al. 2012). This peptide, named juruin, lacks haemolytic activity on human
erythrocytes at the antimicrobial concentrations and it is very similar to peptides found
in Selenocosmia Ausserer, 1871, Chilobrachys Karsch, 1892, and Haplopelma Simon,
1892 species (Ayroza et al. 2012). Juruin has very potent activity against the majority
of the fungal and yeast strains, which can be used for development of new drugs and
antibiotics (Ayroza et al. 2012).
To date, the genus Avicularia includes 47 species and two subspecies and is re-
corded from Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French
Guiana, Brazil, Ecuador, Peru, Bolivia, and Chile as well across the Caribbean archi-
pelago including Cuba, Dominican Republic, Puerto Rico, Martinique and Guade-
loupe, and Trinidad and Tobago (World Spider Catalog 2016).
Even though the genus has a large number of species and has historical importance,
it remains unrevised. To date, only short, taxonomic works were published, with the
exception of Bertani and Fukushima (2009), who discussed some aspects of behavior,
distribution, and conservation of the Avicularia species. With 31 nomina dubia and
three nomina nuda (World Spider Catalog 2016), in addition to the many understud-
ied valid species, it is an urgent task to revise the genus. Thus, the aim of this work is to
revise Avicularia and propose a cladistic hypothesis for the genus and its close relatives.
Taking into account the historical taxonomic difficulties, poor informative descrip-
tions allied to old or lost type material and the extreme morphological homogeneity of
the species, the present work does not intend to solve all the taxonomic problems re-
lated to the genus but rather to give a foundation for future and more detailed studies.
An additional difficulty of this work is the existence of relatively few specimens
housed in scientific collections and available for taxonomic research. Avicularia is one
of the most popular genera in the pet trade, with thousands of specimens spread out all
over the world. However, only few specimens from the pet trade are deposited in col-
lections and these are usually lacking locality or other data, therefore restricting their
use in taxonomic studies.

Material and methods


Specimens from the following institutions were examined:
American Museum of Natural History, New York (AMNH); The Natural History
Museum, London (BMNH); California Academy of Sciences, San Francisco (CAS);
Museu do Departamento de Zoologia da Universidade de Brasília, Brasília (DZUB);
Instituto Butantan, São Paulo (IBSP); Universidad Nacional de Colombia, Bogota
(ICN–AR); Museu do Instituto Nacional de Pesquisas da Amazônia, Manaus (INPA);
Museu Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires
(MACN–AR); Museu de Ciências e Tecnologia da Pontifícia Universidade Católica
do Rio Grande do Sul, Porto Alegre (MCP); Museum of Comparative Zoology, Har-
vard University, Cambridge (MCZ); Instituto de Zoologia, Universidad Central de
Venezuela, Maracay (MIZA); Muséum National d’histoire Naturelle, Paris (MNHN–
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 11

AR); Museu Nacional, Rio de Janeiro (MNRJ); Museu Paraense Emílio Goeldi, Be-
lém (MPEG); Universidad Nacional Mayor de San Marcos, Lima (MUSM–ENT);
Museum Wiesbaden (Naturwissenschaftliche Sammlung), Wiesbaden (MWNH);
Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP); Forschung-
sinstitut und Naturmuseum Senckenberg, Frankfurt (SMF); Museo de Entomologia
Klaus Raven Büller, Universidad Nacional Agraria La Molina, Lima (UA); Museum
of Evolution, University of Uppsala, Uppsala (UUZM); Museum für Naturkunde
der Humboldt-Universität, Berlin (ZMB); and Museu de Zoologia da Universidade
Estadual de Campinas, Campinas (ZUEC).
The AMNH and CAS collections do not assign numbers to specimens or sets.
Thus, in order to individualize and facilitate localization of the specimens by subse-
quent researchers, we gave informal codes and numbers after Museum acronyms.
The general description format follows Raven (2005), with modifications, mainly
of setae and thrichobothria patterns, which were not, herein, studied in detail. All
the measurements are in millimeters (mm). A stereomicroscope Leica M205C and a
microscope Leica DM2500, both with a DFC 450 camera attached, combined with
Leica LAS Montage and LAS 3D modules, were used to obtain images and measure-
ments of small body parts and urticating setae, respectively. Large body parts as leg
articles and carapace were measured with a digital Mitutoyo callipter with an error of
0.005, rounded up to two significant decimals. The measurements of legs and palps
were taken on the dorsal aspect of the left side, unless appendages were lost, damaged,
or obviously regenerated. Structures of the left side of the specimens were chosen for
descriptions. When using structures of the right side, the figures were mirrored to
show them as of the left side and allow for easy comparison. A Scanning Electron
Microscope JEOL JSM840A from Laboratório de Microscopia Eletrônica of Instituto
de Física of Universidade de São Paulo (USP) was used to obtain microphotographs.
Male palpal bulb terminology follows Bertani (2000). Urticating setae terminology
follows Cooke et al. (1972). Geographical coordinates: primary sources are between
round brackets, and secondary sources obtained with Google Earth™, between square
brackets. The coordinates from the secondary source were obtained from the center of
the municipality cited in the specimen label and are in DMS (Degrees, Minutes, and
Seconds), with format rounded off to minutes. Maps were made with SimpleMappr,
an online tool to produce maps (Shorthouse 2010). Abbreviations: ALE = anterior
lateral eye; AME = anterior median eye; CI = consistence index; ITC = inferior tarsal
claw; L = length; PLE = posterior lateral eye; PLS = posterior lateral spinneret; PME
= posterior median eye; PMS = posterior median spinneret; RI = retention index; and
STC = superior tarsal claws. Missing leg articles are indicated by “–” in descriptions.

Cladistic analysis
A data matrix with 46 taxa and 71 characters (Table 1) was analyzed with two pro-
grams using equal (Nona 2.0 for Windows (Goloboff 1998)) and implied character
12
Table 1. Data matrix showing the distribution of character states in cladistic analysis. (?= unknown, - = non-applicable; both codification treated as missing data).

Taxon \ Character 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35
M. santuario 0 0 0 0 0 0 0 0 0 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 - 0 0 0 0 0 0 0 0 0 0
H. rondoni 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 - 0 0 0 0 0 0 0 1 0 0
Pterinochilus sp. 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 1 0 0 0 0 0 1 0 0 1 - 0 0 0 0 0 0 0 0 0 0
P. muticus 0 1 0 0 0 1 0 1 1 1 1 0 1 1 0 0 0 0 1 0 0 1 0 0 0 - 0 0 0 0 0 0 0 1 0 0
Haplopelma sp. 0 1 0 0 0 1 0 0 1 1 1 0 0 0 0 1 1 0 0 0 0 1 0 0 1 - 0 0 0 0 0 0 2 - - -
Phlogiellus sp. 1 0 0 2 0 1 0 0 1 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 - 0 0 0 0 0 0 0 0 0 0
Lasiodora sp. 0 1 0 0 0 0 0 0 1 1 1 0 0 0 1 0 0 0 1 0 0 1 0 0 0 - 0 0 0 0 0 0 1 0 - -
P. vulpinus 0 0 0 1 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 3 - 0 0 0 0 0 0 0 1 - -
Poecilotheria sp. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 1 1 0 0 0 1 0 0 2 - 0 0 0 0 0 0 2 - - -
E. olivacea 0 0 1 1 0 0 0 0 1 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 0 0 1 - - - -
Stromatopelma sp. 1 0 1 0 0 0 1 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 1 0 1 - 0 0 0 0 0 0 0 0 0 0
H. maculata 1 0 1 0 0 0 1 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 1 0 1 - 0 0 0 0 0 0 0 0 0 0
Psalmopoeus sp. 1 0 0 0 0 0 1 0 1 2 1 0 1 0 0 0 0 1 0 0 0 1 0 1 4 0 0 1 0 0 0 0 0 1 0 0
Tapinauchenius sp. 1 0 0 0 0 0 1 0 1 2 1 0 1 0 0 0 0 0 0 0 0 1 0 1 4 0 0 1 0 0 0 0 0 0 0 0
E. murinus 1 0 0 0 0 0 0 0 1 2 1 0 1 0 0 0 0 0 0 0 1 1 0 1 0 - 0 1 0 0 0 0 0 0 0 0
E. uatuman 1 0 0 0 0 0 0 0 1 2 1 0 1 0 0 0 0 0 0 0 0 2 0 1 0 - 0 1 0 0 0 0 0 0 0 0
T. seladonia 0 0 0 0 1 0 0 0 1 2 1 0 1 0 0 0 0 0 0 1 0 0 0 0 4 0 1 0 0 0 0 0 0 0 0 1
T. amma 0 0 0 0 1 0 1 0 1 2 1 1 - 0 0 0 0 0 0 1 0 1 0 ? ? ? ? ? 0 0 0 0 0 0 1 0
T. costae 0 0 0 0 1 0 0 0 1 2 1 1 - 0 0 0 0 0 0 1 0 1 0 0 4 0 0 0 0 0 0 0 0 0 0 0
T. curumim 0 0 0 0 1 0 1 0 1 2 1 1 - 0 0 0 0 0 0 1 0 1 0 0 4 2 0 1 0 0 0 0 0 1 0 1
T. paschoali 0 ? 0 0 1 0 ? 0 1 2 ? 0 1 0 0 0 0 0 0 1 0 1 0 0 5 - 0 1 ? 0 0 0 0 1 0 0
P. rufonigrum 1 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 0 0 1 4 2 1 0 0 0 0 0 0 0 0 0
P. bromelicola 1 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 0 0 1 4 0 1 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 5 - 1 0 1 0 0 0 0 1 0 1
Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

I. hirsutum
I. zorodes 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 5 - 1 0 0 0 0 0 0 0 0 0
I. vanini 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 ? ? - ? ? 0 0 0 0 0 1 0 1
I. katiae 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 0 0 1 5 - 0 1 0 0 0 0 0 1 0 0
I. oliveirai 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 5 - ? ? 0 0 0 0 0 1 0 1
Taxon \ Character 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35
I. marcoi 0 ? 0 0 0 0 ? 0 1 2 ? 1 - 0 0 0 0 0 0 0 0 1 0 ? ? - ? ? ? 0 0 0 0 1 0 1
C. laeta comb. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 4 2 1 0 0 0 0 0 0 0 0 0
C. versicolor comb. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 0 0 1 4 0 1 0 0 0 0 0 0 0 0 0
Y. sooretama comb. n. 0 0 0 0 0 ? 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 6 - 1 0 1 0 0 0 0 1 0 0
Y. gamba comb. n. 0 0 0 0 0 ? 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 6 - 1 0 1 0 0 0 0 0 0 0
Y. diversipes comb. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 2 0 1 6 - 1 0 0 0 0 0 0 0 0 0
A. rickwesti comb. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 0 5 - 0 0 0 0 0 0 0 0 - -
A. avicularia 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 ? 0 1 4 2 0 1 0 1 ? 0 0 0 0 0
A. variegata stat. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 ? 0 1 4 2 0 1 0 1 1 0 0 0 1 0
A. taunayi 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 4 1 0 1 0 0 0 0 0 1 1 0
A. juruensis 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 ? 0 1 4 2 0 1 0 ? ? 0 0 0 1 0
A. rufa 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 2 0 1 4 2 0 1 0 1 1 0 0 0 0 0
A. purpurea 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 4 0 1 0 0 0 0 0 0 0 0 0
A. merianae sp. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 4 1 0 1 0 0 0 0 0 0 0 0
A. hirschii 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 1 7 - 0 1 ? 0 0 0 0 0 0 1
A. minatrix 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 0 4 0 0 1 0 0 0 0 0 0 0 0
A. lynnae sp. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 ? ? ? ? ? 1 ? 0 ? ? ? ? ?
A. caei sp. n. 0 0 0 0 0 0 0 0 1 2 1 1 - 0 0 0 0 0 0 0 0 1 0 ? ? ? ? ? 0 ? 0 ? ? ? ? ?
Taxon \ Character 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70
M. santuario ? 0 0 - - 0 0 0 0 - 2 0 0 0 0 - 0 0 0 0 0 0 - 0 0 0 - - 0 0 0 0 0 0 -
H. rondoni 0 0 0 - - 0 1 0 1 0 0 0 0 0 0 - 0 0 0 0 0 0 - 0 0 0 - - 0 0 0 0 0 3 -
Pterinochilus sp. 0 0 0 - - 1 0 0 0 - 0 0 0 0 0 - 0 0 0 0 0 1 - 0 0 0 - - 0 0 0 2 1 3 -
P. muticus 0 0 0 - - 1 0 0 1 0 0 0 0 0 1 2 0 0 0 0 1 - - 0 0 0 - - 0 0 0 0 0 3 -
Haplopelma sp. 0 - 0 - - 1 1 1 0 - 0 1 2 0 1 0 1 1 1 1 0 2 1 0 0 0 - - 0 0 0 1 1 3 -
Phlogiellus sp. 0 0 0 - - 1 0 0 1 1 0 0 0 0 0 - 0 0 0 0 1 - - 0 0 0 - - 0 0 0 1 1 3 -
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818...

Lasiodora sp. 0 - 0 - - 0 1 1 0 - 1 1 1 0 0 - 1 1 1 1 0 0 - 0 1 0 - - 1 0 0 1 1 3 -
P. vulpinus 0 - 0 - - 0 1 1 0 - 0 0 0 0 0 - 1 1 0 0 0 0 - 0 0 0 - - 1 1 0 2 2 3 -
Poecilotheria sp. 0 - 0 - - 1 0 0 0 - 1 1 0 0 0 - 1 1 1 0 1 - - 0 0 0 - - 0 0 0 1 1 1 0
E. olivacea - - 0 - - 1 0 0 ? ? 0 1 2 0 0 - 1 1 0 0 1 - - 0 0 0 - - 0 0 0 1 1 1 ?
13

Stromatopelma sp. 0 0 0 - - 1 0 0 0 - 0 0 0 0 0 - 0 0 0 0 1 - - 0 0 0 - - 0 0 0 1 1 1 0
14

Taxon \ Character 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70
H. maculata 0 0 0 - - 1 0 0 0 - 0 0 0 0 0 - 0 0 0 0 1 - - 0 0 0 - - 0 0 0 1 0 1 0
Psalmopoeus sp. 0 0 0 - - 1 0 0 1 1 2 0 0 0 0 - 0 0 0 0 0 0 - 0 0 0 - - 0 0 0 1 1 2 -
Tapinauchenius sp. 0 0 0 - - 1 0 0 0 - 2 0 0 0 0 - 0 0 0 0 0 0 - 0 0 0 - - 0 0 0 1 1 2 -
E. murinus 0 0 0 - - 1 0 0 1 1 0 0 0 0 0 - 0 0 0 0 0 0 - 0 0 0 - - 0 0 1 2 2 3 -
E. uatuman 0 0 0 - - 1 0 0 1 1 2 0 0 0 0 - 0 0 0 0 0 0 - 0 0 0 - - 0 0 1 1 1 3 -
T. seladonia ? 0 0 - - 1 0 0 1 0 3 0 0 0 2 1 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 1 1 2
T. amma 0 1 0 - - 1 0 0 0 - 0 0 0 0 2 1 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 1 1 ?
T. costae 0 0 0 - - 1 1 0 0 - 0 0 0 0 1 2 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 1 1 ?
T. curumim 0 0 0 - - 1 0 0 0 - 0 0 0 0 1 2 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 0 1 2
T. paschoali 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 0 0 ? 1 ? ?
P. rufonigrum 0 0 1 0 0 1 0 0 0 - 2 0 0 0 2 0 0 0 0 0 0 2 0 0 0 1 1 0 0 0 0 0 0 1 3
P. bromelicola 0 0 1 0 0 1 0 0 0 - 2 0 0 0 2 0 0 0 0 0 0 2 0 0 0 1 1 0 0 0 0 0 0 1 3
I. hirsutum 0 0 1 1 0 1 0 0 0 - 0 0 0 0 2 2 0 0 0 0 0 2 0 1 0 1 0 0 0 0 0 2 1 1 1
I. zorodes 0 0 1 1 0 1 0 0 0 - 0 0 0 0 2 2 0 0 0 0 0 2 0 1 0 1 0 0 0 0 0 1 1 1 1
I. vanini 0 0 1 1 0 1 0 0 0 - 2 0 0 0 2 2 0 0 0 0 0 2 0 1 0 1 0 0 0 0 0 1 1 1 ?
I. katiae 0 0 1 1 0 1 0 0 0 - 2 0 0 0 2 2 0 0 0 0 0 2 0 1 0 1 0 0 0 0 0 1 1 1 3
I. oliveirai 0 0 1 1 0 1 0 0 0 - 0 0 0 0 2 2 0 0 0 0 0 2 0 1 0 1 0 0 0 0 0 2 1 1 ?
I. marcoi 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? 1 - 0 0 0 ? 1 1 ?
C. laeta comb. n. 0 0 1 0 1 1 0 0 0 - 2 0 0 0 1 2 0 0 0 0 0 2 0 0 0 1 0 1 0 0 0 1 1 1 0
C. versicolor comb. n. 0 1 1 0 1 1 0 0 1 1 2 0 0 0 2 1 0 0 0 0 0 2 0 0 0 1 0 1 0 0 0 1 1 1 0
Y. sooretama comb. n. 2 1 0 - - 1 0 0 1 1 2 0 0 1 2 2 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 0 1 1
Y. gamba comb. n. 2 1 0 - - 1 0 0 0 - 2 0 0 0 1 2 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 1 1 1
Y. diversipes comb. n. 2 1 1 1 0 1 0 0 0 - 3 0 0 0 2 0 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 1 1 1
A. rickwesti comb. n. 0 - 1 1 0 1 0 0 0 - 1 0 2 0 1 2 1 1 1 0 0 2 1 0 0 1 0 0 0 0 0 1 1 1 0
A. avicularia 0 1 1 1 0 1 0 0 1 1 2 0 0 1 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 0 0 1 0
Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

A. variegata stat. n. 0 1 1 1 0 1 0 0 1 2 2 0 0 1 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 1 1 1 0
A. taunayi 0 1 1 1 0 1 0 0 1 1 2 0 0 1 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 1 1 1 0
A. juruensis 0 1 1 1 0 1 0 0 1 2 2 0 0 1 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 1 1 1 0
A. rufa 0 1 1 1 0 1 0 0 1 1 2 0 0 1 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 0 0 1 0
Taxon \ Character 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70
A. purpurea 1 1 0 - - 1 0 0 1 1 2 0 0 0 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 1 1 1 0
A. merianae sp. n. 1 1 0 - - 1 0 0 1 1 2 0 0 0 2 1 0 0 0 0 0 2 1 0 0 1 0 0 0 0 0 1 1 1 ?
A. hirschii 0 1 1 0 0 1 0 0 1 1 2 0 0 0 2 1 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 0 1 0
A. minatrix 0 1 0 - - 1 0 0 0 - 2 0 0 1 2 1 0 0 0 0 1 - - 0 0 1 0 0 0 0 0 1 1 1 0
A. lynnae sp. n. ? ? 1 0 0 1 0 0 1 1 3 0 0 0 2 1 0 0 0 0 1 - - 0 0 1 ? 0 0 0 0 1 ? 1 ?
A. caei sp. n. ? ? 1 0 0 1 0 0 1 0 3 0 0 0 2 1 0 0 0 0 1 - - 0 0 1 ? 0 0 0 0 1 ? ? ?
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818...
15
16 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

weighting (Piwe 3.0 for Windows (Goloboff 1997)). For Piwe, concavities from 1 to
6 were used. All characters were treated as unordered. Commands used to search for
trees were h10000, h/1000, amb-, mult*500. Absolute and relative Bremer supports
(Bremer 1994) were estimated in Piwe using the following command: h1000, find*,
bsupport50, bsupport*.
In the cladistic analyses, representative species were included of seven of the eight
theraphosid subfamilies recognized by Raven (1985), as well as species of all aviculariine
genera and other available arboreal taxa with uncertain position, e.g., Poecilotheria Si-
mon, 1885, Heteroscodra Pocock, 1899, and Stromatopelma Karsch, 1881, in order to
test Aviculariinae monophyly. Melloina santuario Bertani, 2013 (Paratropididae) was
used to root the cladogram. The choice of this genus was done based on two considera-
tions: paratropidids were at the beginning of this study considered as the sister-group of
Theraphosidae in morphological analysis (Raven 1985), and Melloina is the paratro-
pidid genus which retains more plesiomorphic characters shared with the Theraphosi-
dae (Raven 1985). However, recent molecular analyses indicate Barychelidae as the
sister-group of Theraphosidae (Bond et al. 2012, Hamilton et al. 2016, Garrison et al.
2016, Wheeler et al. 2016), so future studies should aim to include more taxa based on
molecular analyses. The ingroup specimens used in cladistics analysis are those used in
the descriptions. Specimens used as outgroups are: Paratropididae, Glabropelmatinae:
Melloina santuario Bertani, 2013, holotype male (MIZA 520) and paratype female
(MNRJ 12965), both from Venezuela, Lara, Cueva El Santuario [9°49'N, 70°03'W],
O. Villarreal col., 19 April 2000. Theraphosidae, Aviculariinae: Ephebopus murinus
(Walckenaer, 1837), male (IBSP 9650) and female (IBSP 9658), Brazil, state of Pará,
Tucuruí, U.H.E. Tucuruí, [4°20'S, 49°31'W], Equipe de Resgate de Fauna col., 1984;
Ephebopus uatuman Lucas, Silva & Bertani, 1992, male holotype (IBSP 4939) and
paratype female (IBSP 4940), Brazil, state of Amazonas, Presidente Figueiredo [2°02'S,
60°01'W], Uatuman River, Balbina Hydroelectric Power Station, M. Costa col., 19
February 1988; Heteroscodra maculata Pocock, 1899, male, Africa, pet trade (IBSP
9642); female, Guinea-Bissau, pet trade (IBSP 9644); Iridopelma hirsutum Pocock,
1901, male, Brazil, state of Paraíba, João Pessoa [7°07'S, 34°52'W], P. F. L. Duarte col.,
30 November 1979 (IBSP 8078); female, Brazil, state of Alagoas, Murici, E. E. Murici,
UFAL (09°14'1.73"S, 35°50'1.61"W), R. Bertani, D. R. M. Ortega and R. H. Naga-
hama col., 16 August 2006 (MNRJ 06252); Iridopelma katiae Bertani, 2012, holotype
female, Brazil, state of Bahia, Mucugê, Parque Nacional da Chapada Diamantina
(12°45'7.00"S, 41°30'2.18"W), 1399 m a.s.l., R. Bertani, C. S. Fukushima and R. H.
Nagahama col., 18 February 2008 (MZUSP 36887) and paratype male, same data,
(12°45'3.77"S, 41°30'4.01"W), 1377 m a.s.l., R. Bertani, C. S. Fukushima and R. H.
Nagahama col., 17 February 2008 (MZUSP 36888); Iridopelma marcoi Bertani, 2012,
holotype female, Brazil, state of Bahia, São Desidério (12°28'52"S, 45°09'10"W), 724
m a.s.l, M. A. Freitas col., October 2009, under tree bark, 1 m above the ground, area
of carrasco vegetation (MZUSP 36891); Iridopelma oliveirai Bertani, 2012, holotype
male, Brazil, state of Bahia, Central, Toca dos Pilões [11°08'S, 42°06'W], A. D. Bres-
covit col., July 2000 (IBSP 10100) and paratype female, at night in “Macambira” bro-
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 17

meliad (Bromelia laciniosa), same data (IBSP 8714); Iridopelma vanini Bertani, 2012,
holotype female, Brazil, state of Piauí, Parnaíba [2°53'S, 41°41'W], 5 m a.s.l., R. Ber-
tani col., November 1994 (IBSP Ref. 74.595) and paratype male, Brazil, state of Mara-
nhão, Barreirinhas, Parque Nacional dos Lençóis Maranhenses [2°41'S, 42°55'W), 32
m a.s.l., Equipe Biota col., 12–18 October 2001 (IBSP 11328); Iridopelma zorodes
(Mello-Leitão, 1926), male, Brazil, state of Bahia, Mata de São João, RPPN Camuru-
jipe, in a leaf at night, R. Bertani, C. S. Fukushima and R. H. Nagahama col., 04 Oc-
tober 2007 (MNRJ 6254); female, Brazil, state of Sergipe, Areia Branca, Mosqueiro, I.
Matos col., 01 November 1992, ref. 79901 (IBSP 11760); Pachistopelma bromelicola
Bertani, 2012, holotype male, Brazil, state of Bahia, Elísio Medrado, RPPN Jequitibá
(12°52'3.20"S, 39°28'9.09"W), R. Bertani, C. S. Fukushima and R. H. Nagahama col.,
07 October 2007, collected at night, found immature inside bromeliads, matured in
captivity in May 2010 (MNRJ 06241) and paratype female, same data (MNRJ 06242);
Pachistopelma rufonigrum Pocock, 1901, male and female, Brazil, state of Alagoas, Mu-
rici, Estação Ecológica de Murici (9°14'9.52"S, 35°48'0.25"W), 245 m a.s.l., R. Ber-
tani, R. H. Nagahama and D. R. M. Ortega col., 11 August 2006 (MNRJ 6246
AL1100); Psalmopoeus cambridgei Pocock, 1895, male, Trinidad-Tobago, pet trade
(IBSP 9653); Psalmopoeus sp., female, Venezuela, pet trade (IBSP 9655); Stromatopelma
sp., male, Sierra Leone, pet trade (IBSP 9665); female, Africa, pet trade (IBSP 11136);
Tapinauchenius violaceus (Mello-Leitão, 1930), holotype female of T. purpureus
Schmidt, 1995 (SMF 38042) and paratype male of T. purpureus Schmidt, 1995, French
Guiana (SMF 38046); Typhochlaena amma Bertani, 2012, holotype female, Brazil, state
of Espírito Santo, Santa Teresa, Estação Ecológica de Santa Lúcia [19°58'S, 40°32'W],
672 m a.s.l, A. P. L. Giupponi col., February 2008 (MNRJ 06239) and paratype male,
same locality, no further data (MNRJ 12926); Typhochlaena costae Bertani, 2012, holo-
type female, Brazil, state of Tocantins, Palmas, U.H.E. Luís Eduardo Magalhães
[10°12'S, 48°24'W], 211 m a.s.l, during faunal rescuing, M. Costa and D. Cândido
col., 12 January 2002 (IBSP unnumbered) and paratype male, Brazil, Tocantins, La-
jeado (9°46'4.85"S, 48°21'6.69"W), 226 m a.s.l, G. Puorto, R. Martins and I. Knysak
col. (pitfall trap), April 2002 (IBSP unnumbered); Typhochlaena curumim Bertani,
2012, holotype female, Brazil, state of Paraíba, Areia, Reserva Ecológica Estadual Mata
do Pau-Ferro [6°58'S, 35°42'W], 500 m a.s.l., under tree bark, A. D. Brescovit, R.
Bertani, A. B. Bonaldo and S. C. Dias col., September 1999 (IBSP 8701) and paratype
female (IBSP 8354), same data; male, Brazil, state of Rio Grande do Norte, Baía For-
mosa, lying over a leaf in a bush at night, S. N. Migliore col., 19 April 2014, S90
(MNRJ 06915); Typhochlaena paschoali Bertani, 2012, holotype female and immature
paratype, Brazil, state of Bahia, Camacan [15°24'S, 39°30'W], no further data (MNRJ
13723); paratypes 1 female, 1 subadult male, 9 immatures, Brazil, state of Bahia, Jus-
sari [15°10'S, 39°29'W], no further data (MNRJ 12928 – R2981); Typhochlaena sela-
donia (C. L. Koch, 1841), 1 male, 1 female exuvium, Brazil, state of Bahia, Camaçari,
Jacuipe [12°42'S, 38°07'W], T. Brazil ded., August 1980 (IBSP 4551); 1 female, Brazil,
state of Bahia, Salvador, Alphaville [12°56'S, 38°21'W], G. G. Montingelli col., 11
December 2001 (IBSP 109718); Eumenophorinae: Pelinobius muticus Karsch, 1885,
18 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

male (IBSP 8530), female (IBSP 9643), both from Kenya, born in captivity; Harpac-
tirinae: Pterinochilus sp., male, Angola, Biula-Dala [11°10'S, 20°12'E] (IBSP 9647);
female, Africa, pet trade (IBSP 8765); Ischnocolinae: Holothele rondoni (Lucas &
Bücherl, 1972), holotype male and paratype female, Brazil, state of Amazonas, Iauaretê
[0°36'N, 69°11'W] (IBSP 4090); Ornithoctoninae: Haplopelma longipes von Wirth &
Striffler, 2005, male, Cambodia, Skuon [1°33'N, 104°55'E], A. Anderson col., 10 July
2003 (MZUSP 28761); Haplopelma minax (Thorell, 1897), female, Thailand, 1 mi east
of Bangkok [13°43'N, 100°31'E] (IBSP 9645); Selenocosmiinae: Phlogiellus sp, male,
Vietnam, R. Blauman col., April 1989 (MZUSP 28762); female (AMNH), pet trade;
Poecilotheria sp., male, India, pet trade (IBSP 9660); Poecilotheria ornata Pocock, 1899,
female, Sri Lanka, pet trade (IBSP 8767); Theraphosinae: Phrixotrichus vulpinus (Karsch,
1880), 3 males, Chile (IBSP 3817–A); 1 female, Chile, Osorno [40°34'S, 73°09'W]
(IBSP 3817–B); Lasiodora sp., male, Brazil, state of Paraíba, João Pessoa [7°07'S,
34°52'W] (IBSP 11143); female, Brazil, state of Pernambuco, Jaboatão dos Guararapes
[8°06'S, 35°00'W], Conjunto Murebeca (IBSP 10293); Subfamily Incertae sedis: Encyo-
cratella olivacea Strand, 1907, holotype female, Tanzania, Amani (05°09'S, 38°36'E),
Vosseler leg., November 1903 (ZMB 10484); male, Tanzania, mountains near Arusha
(03°23'S, 36°40'E) wild caught, reared to maturity, 2004 (BMNH); female holotype
and 2 females paratypes of Xenodendrophila gabrieli Gallon, 2003, northern Tanzania,
mountains near Arusha (03°23'S, 36°40'E), Joe Beraducci leg., circa April 1999
(BMNH 2005.123).
In the cladistic analyses we used the characters proposed by West et al. (2008),
Bertani (2012), and 15 new characters. Forty characters (0–3, 5–18, 20–22, 31–33,
41, 43, 47–48, 52–57, 60–61, 64–66, and 69–70) are the same used by West et al.
(2008) or were slightly modified here. Ten characters (4, 19, 23, 26, 35, 37–38, 59,
67 and 68) are strictly the same used by Bertani (2012). Characters 24, 44–46 and 50
were adapted from that work. The newly introduced characters are 25, 27–30, 34, 36,
39–40, 42, 49, 51, 58, 62–63. In characters marked with an asterisk (*, viz 7–8, 14,
20, 31, 60 and 65) fit and steps are not given as the character is not significant to the
analyses but were kept to indicate autapomorphies for terminal taxa.
Character coding is always challenging, especially regarding problems concerning
“part” coding (e.g., presence of tibial apophysis) versus “character-variable” coding
(e.g., development of tibial apophysis). We decided to code “part” characters sepa-
rately from its “character-variable” characters to preserve transformational independ-
ence between them, assuring that primary homology statements of both characters
were included in the analyses (Lee and Bryant 1999). Fusing coding results in a loss
of phylogenetic information and also in the determination of taxa synapomorphies
by others characters (Lee and Bryant 1999). The characters that were splited are: 8/9,
11/12, 24/25, 31/32, 44/45, 56/57/58, 61/62, and 61/63.
Theraphosids are notorious for their morphological homogeneity (Bertani 2001).
Due to this, it is extremely difficult to find characters suitable for taxonomy and clad-
istics. In order to find new characters, we have tried many approaches. We took meas-
urements of leg and palp articles such as tibia, metatarsi and femora, and established
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 19

ratios between them and ratios between one of these articles and the carapace. We also
calculated the ratio between the length of tibial apophysis and tibia. Unfortunately, all
results were not significant enough to be used in the analyses. Measurements of male
palpal bulb, such as the ratio between the length of embolus and carapace, showed the
same problems. Besides this, the morphology of the Aviculariinae male palpal bulb is
a problem per se. It is very difficult to position the specimen in order to make reliable
and replicable embolus measurements.
Another feature we have tried to use is the length of type II urticating setae. Some
species, such as A. hirschii, A. minatrix, A. taunayi, Y. sooretama comb. n. and Y. gamba
comb. n., apparently have unusual urticating setae length. However, we could not
confirm this suspicion since we have just one or two specimens of those species. Thus,
it was impossible to determine the intraspecific length variation and establish a reliable
length range for each species. Beside setae length, another urticating setae feature that
could be used is the distribution of barbs along setae in females (Figs 16–17). Usually,
females present barbs only in a short area near hair stalk (Stradling 1978). In contrast,
morphotype 5 of A. avicularia presents barbs along 1/3 of the seta length (Fig. 17).
However, since we have only three immature females, it is not possible to state if that
is a typical feature of the morphotype population.
Thus, after trying different attempts and approaches, we consider the following
characters suitable to be used:

0. Anterior row of eyes: (0) procurved (Fig. 13), (1) straight (Fig. 12).
1. Clypeus: (0) absent or narrow; (1) present, wide.

West et al. (2008) and Bertani (2012) used three states for this character: “ab-
sent”, “narrow” and “wide”. Herein, we consider that the states “absent” and “nar-
row” should be merged due to difficulty in establishing limits between them. The state
“wide”, on the other hand, is easily distinguishable.

2. Fovea, closure: (0) slit like; (1) pit like.


3. Labial cuspules, number: (0) 30–300; (1) 0–20; (2) 350–450.
4. Sternum, shape: (0) longer than wide, not truncated behind (Fig. 9); (1) as long
as wide, truncated behind (Fig. 7).
5. Sigilla, posterior pair, position: (0) marginal, less than 1.5 diameters from mar-
gin; (1) close to the center, more than 2 diameters from margin.
6. Setae on metatarsi and tibia I–IV, length, males: (0) same length as other arti-
cles (Fig. 299); (1) longer setae laterally projected, forming a brush (Fig. 301).
7. Scopulae on metatarsi IV, division: (0) divided by setae or spiniform setae; (1)
not divided.*
8. Tarsal scopulae, occurrence: (0) no true scopula; (1) true scopula.*
9. Tarsal scopulae, setae development: (0) scopula composed of sparse setae; (1) dense
scopula that does not extend much laterally; (2) scopula very extensive laterally,
giving the tarsi and metatarsi I and II a spatulate appearance (Figs 299–301).
20 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

10. Tarsi IV, division, males: (0) cracked; (1) integral.


11. Leg spines, occurrence: (0) present; (1) absent.
12. Leg spines, distribution: (0) in whole tibiae and/or metatarsi; (1) only in ven-
tral apical tibiae and/or metatarsi.
13. Palpal femora, scopula on retrolateral face, occurrence: (0) absent, (1) present.
14. Femora IV, scopulae on retrolateral face, occurrence: (0) absent; (1) present.*
15. Chelicerae, scopulae on retrolateral face, occurrence: (0) absent, (1) present.
16. Maxillae, spiniform setae on lower prolateral face: (0) absent, (1) present.
17. Stridulatory bristles form maxillae lyra: (0) absent, (1) present (Fig. 11).
18. Stridulatory bristles on coxae I, occurrence: (0) absent, (1) present.
19. Posterior lateral spinnerets, distal article, shape: (0) digitiform (Fig. 10);
(1) domed (Fig. 8).
20. Patellae and tibia, stripes, color: (0) same color of the rest of the segment, (1) white.*
21. Leg rings on distal femora, tibiae and metatarsi, coloration: (0) same color of
the rest of the segment, (1) white (Fig. 299), (2), yellow or orange (Fig. 300).

Avicularia avicularia and A. variegata stat. n. have morphotypes with different


colors of leg rings. Due to this intraespecific variability, these two taxa were coded as
“?” in the matrix.

22. Tibiae, metatarsi and tarsi, dorsal coloration: (0) homogeneous color, (1) with
black marking (Fig. 5).
23. Color pattern, ontogenetic change: (0) pattern remains practically the same
during ontogeny; (1) pattern presents drastic changes during ontogeny.

Most outgroup species have only slight variations in the general color and abdomi-
nal pattern during their ontogeny. We consider a drastic change when legs, carapace,
and abdomen change their color during ontogeny. Most aviculariines have abdomen
dorsum pattern with stripes in their early life stages, losing it when reaching matu-
rity, which is, herein, considered as state 1. Other species, such as Ephebopus murinus,
despite not having heterogeneous abdominal pattern during all ontogeny, have some
conspicuous changes: immatures have light brown legs with black tarsi and black cara-
pace, while adults have black legs with very conspicuous white stripes and light brown
carapace. This condition is also considered as a drastic change and codified as “1”.

24. Dorsal abdominal pattern in immatures (see figures in Bertani 2012): (0) ho-
mogeneous; (1) herringbone; (2) with a zigzag central longitudinal dark stripe
over a clear spot, which is marginated in black and connects to five narrow
transversal black stripes; (3) two median dorso-lateral spots; (4) central longi-
tudinal black stripe with 5–6 lateral stripes, connecting or not with the central
stripe (Fig. 37); (5) leaf pattern: black with a large central clear area having a
longitudinal dark stripe, area closer to longitudinal dark stripe usually red-
dish; (6) central longitudinal reddish stripe inside a dark area with zigzag bor-
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 21

ders connected to transverse dark stripes (Fig. 295); (7) longitudinal central
stripe of a different color of remaining abdomen (Fig. 184).

Character modified from Bertani (2012). A new state was added (state 7), which
was applied here only to A. hirschii. State 4 is present in most Avicularia species. This
abdominal stripped pattern is also spread over the subfamily. It can also be found in
Caribena gen. n. species, in both Pachistopelma species, in T. curumim, T. costae and
T. seladonia as well as Tapinauchenius sp. and Psalmopoeus sp.

25. Abdominal pattern, immatures, central longitudinal stripe, connection with


lateral stripes: (0) connected with all lateral stripes (Fig. 167); (1) connected
only with first and second lateral pair of stripes (Fig. 101); (2) disconnected
from lateral stripes (Fig. 44).

In species with abdominal pattern corresponding to state 4 of character 24, the


central longitudinal black stripe can connect or not with the lateral stripes. State 0 is
found in many Aviculariinae species such as A. purpurea and A. minatrix as well as in
C. versicolor comb. n., T. costae, T. seladonia, P. bromelicola, Tapinauchenius sp. and
Psalmopoeus sp. State 1 is found only in two Avicularia species from the early stages: A.
merianae sp. n. and A. taunayi. Central stripe disconnected from lateral stripes (state
2) can be found in A. rufa, A. juruensis, A. variegata stat. n. and A. avicularia, as well
as in C. laeta comb. n., T. curumim and in P. rufonigrum.

26. Body coloration, juveniles: (0) brownish or grayish (Fig. 44); (1) metallic
green or blue (Fig. 239).
27. Tarsi, coloration, juveniles: (0) same color of other articles (Fig. 167); (1)
black (Fig. 44).

Black tarsi contrasting with other lighter articles in legs and palpi are present in
juveniles of some Tapinauchenius and Ephebopus species. It also occurs in all Avicularia
species with known immature stages (except A. purpurea) as well as in I. katiae and T.
paschoali. Since Psalmopoeus sp. has black tarsi contrasting with clear metatarsi, it was
codified as state 1 despite most articles having a dark color. Typhochlaena curumim
has black tarsi and metatarsi; but as the other articles are lighter and there is a strong
contrast between them, it is considered as having state 1.

28. Dorsal abdominal pattern, single dorsal stripe, male, occurrence: (0) absent
(Fig. 43); (1) present (Fig. 297).

Males of Y. sooretama comb. n., Y. gamba comb. n., A. lynnae sp. n., and I. hirsu-
tum have a single dorsal central stripe on abdomen dorsum. This pattern is very dis-
tinct from those found in females of these same species and also very distinct from pat-
terns commonly found in other aviculariine males. Patterns other than dorsal central
22 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

stripe were all codified as state 0. Avicularia hirschii is coded as “?” since the holotype,
the single known male, has the abdomen in poor condition. The description also does
not allow us to recognize its abdominal pattern.

29. Abdominal setae, distribution, females: (0) homogeneous (Fig. 304); (1) het-
erogeneous, with long guard-setae grouped on lateral and dorso anterior areas
(Figs 302–303).

Presence of long guard-setae grouped on lateral and dorsal anterior areas of the
abdomen is a characteristic found only in females of A. avicularia, A. rufa, A. juruensis
and A. variegata stat. n. These longer guard-setae have different coloration from the
shorter body setae found on remaining abdominal areas. In males, the distribution
pattern is homogeneous, with long guard-setae spread all over the dorsal abdomen.
This character is coded “?” in A. juruensis since each morphotype has a different
type of abdominal setae pattern.

30. Legs and palp, long guard-setae, coloration: (0) setae with homogeneous col-
oration or gradually lightening along its length (Fig. 299); (1) setae with dark-
er base and contrasting whitish apex (grizzled setae) (Fig. 300).

Theraphosid spiders have two main covering setae types in legs and pedipalps; the
short setae that densely covers most of the article area, providing a velvety aspect, is
known as short body setae (Foelix 2010). The other type is the very long tactile setae,
which presents insertion into a socket, allowing them to move, and is more spaced out
among each other; this is known as guard-setae (Foelix 2010). These long setae are nor-
mally homogeneously dark or present a gradual lightening toward apex. But, in some
species, these setae are dark and present a contrasting whitish apex.
All examined outgroup taxa have setae with homogeneous coloration along its
length. In the ingroup, some species such as A. avicularia, A. rufa, A. juruensis and A.
variegata stat. n. have the long setae with a contrasting whitish apex, giving a grizzled
body aspect (Fig. 300). However, this character is coded “?” in A. juruensis and A.
avicularia since some morphotypes (morphotypes 1 and 3 for A. juruensis and mor-
photypes 1, 2, 4, and 5 for A. avicularia) have homogeneous coloration along most of
the guard-setae.

31. Spermathecae, occurrence: (0) present; (1) absent.*


32. Spermathecae, number: (0) two, completely separated (Fig. 14), (1), two,
fused at base; (2) one, totally fused.
33. Spermathecae, walls, shape: (0) without projections or lobes (Fig. 21); (1) with
projections or lobes (Fig. 91).

Of the several specimens of A. rufa, A. juruensis, A. avicularia, and A. variegata stat.


n. examined, a few specimens have one or more discrete lobes in the spermathecae.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 23

We consider them an extreme morphological variation, as the typical spermathecae of


those species lack any type of lobes.
Caribena laeta comb. n. spermathecae have a slight intumescence in their apex
(Figs 227–229). It is not a well-defined lobe as in other taxa, such as Y. sooretama
comb. n. (Fig. 257) or in A. taunayi (Figs 91–92). Due to this, we considered C. laeta
comb. n. as having state 0. State 1 is widespread among theraphosid genera.

34. Spermathecae, midwidth: (0) as wide as its base, or midwidth more slender
than the base width but wider than apex width (Fig. 21); (1) midwidth ex-
panded, about 1.5 times its basal and apical portion widths, or wider (Fig. 54).

We consider the spermathecae with midwidth expanded when this region is 1.5
times or more wider than both apex and base widths. Spermathecae of A. variegata
stat. n. (Fig. 54) and A. juruensis (Fig. 106) clearly have state 1. In other aviculariine
species, it is also possible to recognize this pattern despite their unusual spermathecae
morphology, as in spermathecae with lobes in A. taunyai (Fig. 91). Spermathecae mid-
width of Y. sooretama comb. n., Y. gamba comb. n. and A. merianae sp. n. are slightly
wider than their bases’ width, but not enough to consider them as state 1.
In other taxa such as Haplopelma sp., Poecilotheria sp., Lasiodora sp., P. vulpinus
and A. rickwesti comb. n., spermathecae (Fig. 284) are very modified or completely
fused. In these cases we consider the character inapplicable.
35. Spermathecae, shape: (0) non-spiraled or not-twisted (Fig. 21); (1) twisted or
spiraled (Fig. 183).
36. Spermathecae, weakly-sclerotized area, size: (0) weakly-developed or shorter
than half the length of the well-sclerotized area (Fig. 21); (1) twice the length
or the same length of the well-sclerotized area (Fig. 154); (2) spermatheca
virtually non-sclerotized (Figs 257–259).

In some species, the spermathecae have a weakly-sclerotized basal portion, lighter


and softer than the more sclerotized distal area, and without large visible pores. As this
weakly-sclerotized area is barely present in most outgroup species, we consider them as
having state 0. State 1 is found only in A. purpurea (Fig. 154) and A. merianae sp. n. (Fig.
196). All three species of Ybyrapora gen. n. have spermathecae with a very small scleroti-
zation degree, even in adult females (Figs 257–259), and they were coded as state 2. Mel-
loina santuario and T. seladonia have spermathecae virtually non-sclerotized, very similar
to Ybyrapora gen. n. spermathecae. However, we know a single female of each species,
both very small in size, in which the non-sclerotized spermathecae condition may be due
to a not fully mature life stage. Thus, we decided to code both species in matrix as “?”.

37. Spermathecae, curvature: (0) straight or almost so, curved inward (Fig. 14);
(1) with an accentuated outwards curvature medially (Fig. 21).
38. Cymbium, process in retrolateral lobe, occurrence: (0) absent or very weakly-
developed (Figs 162, 305); (1) present, well-developed (Figs 306–307).
24 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

The presence of a process on the cymbium was considered by Raven (1985) a


synapomorphy of Aviculariinae. However, more recent analyses showed that the struc-
ture is lacking in many Aviculariinae genera (West et al. 2008, Bertani 2012). Despite
this, the character is informative and was studied in detail, herein. The process is a
projection on the apical area of cymbium retrolateral lobes (Figs 306–307) and shows
a considerable level of interspecific variation in size, shape, and associated setae. If on
one hand this variation provides more information for cladistic analysis, on the other
hand sometimes is difficult to establish limits for the states. We consider the process
well-developed only when there is a clear and well-defined projection in the cymbium
lobes (Figs 306–307). If there is a small bump without clear delimitation on cymbium
retrolateral area, it was considered a very weakly-developed process, or state 0 (Figs
162, 305). We consider that Pachistopelma spp., Iridopelma spp., Caribena gen. n.
spp., Y. diversipes comb. n., A. rickwesti comb. n., A. avicularia, A. rufa, A. juruensis, A.
variegata stat. n. and A. taunayi have well-developed process (state 1).

39. Cymbium, setae covering the retrolateral process, thickness: (0) thin (Fig. 306);
(1) thick (Fig. 307).

Thin setae covering the retrolateral process are found both in some Avicularia spe-
cies as A. hirschii, A. lynnae sp. n. and A. caei sp. n., as well as in species of Caribena
gen. n. and in Pachistopelma species.

40. Cymbium, process in retrolateral lobe, shape: (0) rounded (Fig. 307); (1) sharp
(Fig. 306).

The apex of the process on retrolateral lobe is usually rounded, except in both spe-
cies of Caribena gen. n., in which the process apex is sharp.

41. Cymbium, prolateral lobe, shape: (0) rounded; (1) subtriangular.


42. Tegulum, format: (0) globous (Fig. 29); (1) piriform (Fig. 320).

We detected two distinct tegula shapes. The globous tegulum is short, narrowing
abruptly and giving origin to a slender embolus since its beginning. The pyriform
tegulum is longer and tapers to form the embolus.
All Aviculariinae possess globous tegulum, except T. costae.

43. Subtegulum, length: (0) small; (1) large, extending down the bulb for half of
the tegulum.
44. Palpal bulb, prominence on tegulum (frontal view), occurrence: (0) absent
(Fig. 312); (1) present (Figs 313–315).

The tegula of some species have a groove in its prolateral side, which forms a
prominence near it (Bertani 2012). This prominence can be better seen in frontal view
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 25

(Figs 313–315). Presence of a prominence is shared by all Avicularia species except


A. minatrix and also in species of Holothele, Pelinobius, Phlogiellus, Ephebopus, and
Psalmopoeus. In other aviculariine species this groove is lacking and the tegulum pro-
lateral area is rounded. This state (0) is found in species of Iridopelma, Pachistopelma,
Typhochlaena (except T. seladonia), Ybyrapora gen. n. (except in Y. sooretama comb.
n.), C. laeta comb. n. and A. rickwesti comb. n. Encyocratela olivacea is coded as “?”
since this character was included after the specimen examination and we could not re-
examine it in the proper position.

45. Palpal bulb, prominence on tegulum, development: (0) weakly-developed


(Fig. 313); (1) developed (Fig. 314); (2) well-developed (Fig. 315).

We found that prominence on tegulum can have different development degrees.


There are species in which prominence is well-developed (state 2), with a deep groove,
as seen in A. variegata stat. n. and in A. juruensis (Fig. 315). When there is a distinct
groove and it is clearly possible to determine a tegulum prominence, though it is not
well-developed, it was coded as state 1. This state is found in most species analyzed
(Fig. 314). However, if the groove is discrete, hard to see, and consequently the promi-
nence is weakly-developed (Fig. 313), it was coded as state 0, as in H. rondoni, P. mu-
ticus, T. seladonia and A. caei sp. n.

46. Embolus, length, retrolateral view: (0) 1.5 to 2.5 times the tegulum’s length;
(1) smaller than tegulum’s length (Fig. 320); (2) 3.0 to 3.5 times the tegulum’s
length (Fig. 323); (3) more than 4 times the tegulum’s length (Fig. 277).

Most Aviculariinae have emboli from 3 to 3.5 times the tegulum’s length (state 2),
including all Avicularia species, except A. lynnae sp. n. and A. caei sp. n., which have
emboli described as very long (state 3).

47. Embolus, distal width: (0) thin, shorter than 1/5 tegulum’s length (Fig. 323);
(1) thick, more than 1/3 tegulum’s length (Fig. 320).
48 Embolus, shape: (0) not flattened (Fig. 323); (1) slightly flattened (Fig. 320);
(2) very flattened (Fig. 286).
49. Embolus, tip: (0) tapers (Figs 318–319); (1) narrows abruptly (Figs 316–317).

In some Aviculariinae taxa such as A. minatrix, A. taunayi, A. rufa, A. juruensis,


A. avicularia, A. variegata stat. n. and Y. sooretama comb. n., the embolus tip narrows
abruptly near 1/5 distal portion (better visualized in dorsal view). The remaining taxa
show a gradual embolus tapering.

50. Embolus, proximal part in frontal view, shape: (0) straight; (1) slightly curved
(Fig. 233); (2) very curved (Fig. 30).
26 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Most outgroups (except Haplopelma sp. and P. muticus) and some Aviculariinae,
such as Psalmopoeus sp., Tapinauchenius sp., Ephebopus spp., Stromatopelma sp. and
Heteroscodra sp., have straight proximal portion of embolus in frontal view. All the
other Aviculariinae taxa have some curvature degree in proximal portion of embolus.
Caribena laeta comb. n. (Fig. 233), Y. gamba comb. n., T. costae, T. curumim and A.
rickwesti comb. n. have slightly curved proximal embolus. The other Aviculariinae taxa
have very curved proximal embolus (state 2) (Fig. 30).

51. Embolus, curved, retrolateral view, angle between tegulum’s margin and em-
bolus medial portion: (0) very acute angle (Fig. 321) (1) acute angle (Fig. 323)
(2) right or obtuse angle (Fig. 322).

We found that the angle between tegulum’s margin and embolus medial portion
in retrolateral view can be informative for cladistic analysis. In Haplopelma sp., Pachis-
topelma spp. and Y. diversipes comb. n., the angle formed is very acute (state 0) (Fig.
321). In other taxa, such as T. costae, T. curumim, Iridopelma spp., Ybyrapora gen. n.
(except Y. diversipes comb. n.), A. rickwesti comb. n., C. laeta comb. n. and P. muticus
sp., an obtuse angle is formed between embolus medial portion and tegulum’s margin
(state 2) (Fig. 322). In the remaining aviculariine species, including all Avicularia, an
acute angle is formed between these two structures (state 1) (Fig. 323).

52. Bulb, prolateral inferior keel on embolus: (0) absent; (1) present.
53. Bulb, prolateral superior keel on embolus: (0) absent;(1) present.
54. Bulb, apical keel on embolus: (0) absent;(1) present.
55. Bulb, retrolateral keel on embolus: (0) absent; (1) present.
56. Tibial apophysis on leg I, occurrence: (0) present (Figs 310–311); (1) absent
or weakly-developed (Figs 308–309).

Specimens of Y. sooretama comb. n., Y. gamba comb. n., Typhochlaena spp.,


P. muticus, Phlogiellus sp., Poecilotheria sp., E. olivacea, Heteroscodra sp., and Stro-
matopelma sp. do not present any type of apophysis or tibial modification. Some
aviculariine species have a discrete elevation on prolateral tibia that can be covered
or not by a cluster of setae but clearly does not form a branch. This discrete elevation
on apical prolateral tibia is found in Y. diversipes comb. n., A. hirschii, A. minatrix,
A. lynnae sp. n., and in A. caei sp. n., and was coded as state 0 since it cannot be
considered as a true apophysis.

57. Tibial apophysis on leg I, shape: (0) 2 branches; (1) 1 branch with a megas-
pine; (2) 1 branch with setae (Figs 310–311).

Males of many barychelid and theraphosid species have tibial apophysis composed
of two branches on leg I (Raven 1985) (state 0). This type of apophysis should have
been present on the ancestral Theraphosoidina, suffering modifications and losses
in all derivate groups (Raven 1985). This condition is present in Theraphosinae
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 27

(represented here in the analysis by Lasiodora sp. and P. vulpinus), in Ischnocolinae


(here represented by H. rondoni) and in Paratropididae (represented by Melloina
santuario). The presence of tibial apophysis composed of single branch with a
megaspine (state 1) is found exclusively in Harpactirinae (Pterinochilus sp. in our
analysis) (Gallon 2003).
The presence of a tibial apophysis with a single branch bearing spiniform setae
(state 2, figs 310–311) is a condition widely distributed in Aviculariinae. All Pachis-
topelma spp., Iridopelma spp., Caribena gen. n. spp. as well as A. rickwesti comb. n. and
most Avicularia species have this condition. This condition also appears in outgroup
taxon Haplopelma minax (Ornithoctoninae).

58. Tibial apophysis on leg I, one branch with setae, spiniform setae on a branch,
branch development; (0) weakly-developed branch (Fig. 310); (1) well-devel-
oped branch (Fig. 311).

We detected different degrees of branch development in species that have a single


and well-defined branch covered with spiniform setae (state 1 for character 57). The
branch can be weakly-developed, not projecting far from tibia longitudinal axis. This
is the condition of Pachistopelma spp., Iridopelma spp. and Caribena gen. n. A well-de-
veloped branch projected far from tibia longitudinal axis is present in most Avicularia
species (A. avicularia, A. rufa, A. juruensis, A. variegata stat. n., A. taunayi, A. purpurea
and A. merianae sp. n.), in A. rickwesti comb. n., and also in Haplopelma sp.

59. Tibial apophysis in leg II, occurrence: (0) absent; (1) present.
60. Type I urticating setae, occurrence: (0) absent; (1) present.*
61. Type II urticating setae, any life stage, occurrence: (0) absent; (1) present.

Despite females of Pachistopelma spp. lacking type II urticating setae, immature stages
and males of both Pachistopelma species do have the type II urticating setae. As we do not
know immature stages nor males of Iridopelma marcoi, this species was coded as “?”.

62. Type II urticating setae, adult female, occurrence: (0) present; (1) absent.

Mature females of Pachistopelma spp. and of I. marcoi lack type II urticating setae.

63. Type II urticating setae, length, females: (0) up to 0.90mm (Figs 16–17); (1)
more than 1 mm (Fig. 243).

According to Cooke et al. (1972), urticating setae type II are approximately 0.5–
1.5 mm in length. Bertani et al. (2003) observed that C. versicolor comb. n. has type
II setae that is more slender and longer than the Avicularia species—up to 1.37 mm
in females. In fact, this species as well as C. laeta comb. n. has longer and slender setae
when compared with Avicularia spp., what was confirmed by comparison of scanning
microscopy images of setae of both genera (Figs 17–18).
28 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

When urticating setae are analyzed, the gender of specimens examined should be
considered because males have setae significantly longer than females (Stradling 1978,
Bertani and Guadanucci 2013).

64. Type III urticating setae, occurrence: (0) absent; (1) present.
65. Type IV urticating setae, occurrence: (0) absent; (1) present.*
66. Type V urticating setae, occurrence: (0) absent; (1) present.
67. Legs, ratio between length of leg IV and I, males: (0) leg IV more than 10%
longer than leg I; (1) leg IV roughly the same length as leg I; (2) leg IV more
than 10% shorter than leg I.
68. Legs, ratio between length of leg IV and I, females: (0) leg IV more than 10%
longer than leg I; (1) leg IV roughly the same length as leg I; (2) leg IV more
than 10% shorter than leg I.
69. Habits, females: (0) retreat within surface layers of soil; (1) arboreal; (2) op-
portunistic; (3) fossorial.
70. Arboreal retreat made by adults: (0) built on tree trunk or on palm tree leaf
base; (1) built in leaves, normally with two or more leaves connected by silk;
(2) built under loosened tree bark; (3) bromelicolous.

Taxonomy
Family Theraphosidae Thorell, 1869

Subfamily Aviculariinae Simon, 1889

Aviculariae Simon, 1889: 213; 1891: 312.


Avicularieae Simon, 1892: 170; 1903: 918; Pocock 1895: 229; 1901: 547.
Aviculariieae Simon, 1903: 958.
Aviculariinae Petrunkevitch, 1928: 34, 81; 1929: 48; 1939: 152, 274; Schiapelli &
Gerschman de Pikelin 1945: 209; Bonnet 1955: 833; Bücherl 1957: 380; Raven
1985: 119; West et al. 2008: 37, 53, 55, 56; Bertani 2012: 5, 73.
Avicularias Mello-Leitão, 1923: 314.

Diagnosis. Aviculariines can be distinguished by the conjunction of the following


characters: legs aspinose or with few apical spines on ventral tibiae and metatarsi;
metatarsi and tarsi with scopulae very extended laterally, mainly on anterior legs, giv-
ing a spatulate appearance (Figs 299–301); two completely separated spermathecae
(Fig. 21); absence of spiniform setae on lower prolateral maxillae.
Included genera: Antillena gen. n., Avicularia Lamarck, 1818, Caribena gen. n., Ephe-
bopus Simon, 1892, Heteroscodra Pocock, 1899, Iridopelma Pocock, 1901, Pachistopelma
Pocock, 1901, Psalmopoeus Pocock, 1895, Stromatopelma Karsch, 1881, Tapinauchenius
Ausserer, 1871, Typhochlaena C. L. Koch, 1850, Ybyrapora gen. n.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 29

Figure 3. Approximate distribution area of Aviculariinae. ? = dubious record.

Distribution (Fig. 3): from state of Vera Cruz in Mexico, throughout Central
America and some Caribbean islands, northern and central South America to state of
São Paulo, Brazil. They can be found from the Atlantic to Pacific coasts. Two genera
(Heteroscodra and Stromatopelma) are found in West and Central Africa.

Key to Aviculariinae genera

(Females of A. lynnae sp. n. and A. caei sp. n. are unknown)

1 Presence of a paddle of urticating setae (type V) on prolateral distal palpal


femora (Fig. 4)............................................................................. Ephebopus
– Absence of a paddle of urticating setae (type V) on prolateral distal palpal
femora.........................................................................................................2
2 Presence of black marks on dorsal tibiae, metatarsi and tarsi (Africa) (Fig. 5)..... 3
– Absence of black marks on dorsal tibiae, metatarsi and tarsi (New World)..... 4
3 Tibia IV incrassate (Fig. 6)...................................................... Heteroscodra
– Tibia IV not incrassate.......................................................... Stromatopelma
4 Sternum as long as wide, truncated behind (Fig. 7); posterior lateral spinner-
ets with domed, short distal article (Fig. 8)............................. Typhochlaena
– Sternum longer than wide (Fig. 9), posterior lateral spinnerets with digiti-
form distal article (Fig. 10)..........................................................................5
5 Spines present on ventral apical tibiae and/or metatarsi...............................6
– Spines absent in all legs................................................................................7
30 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

6 Stridulatory setae forming a lyra on prolateral maxilla (Fig. 11).......Psalmopoeus


– Stridulatory setae absent on maxilla.....................................Tapinauchenius
7 Anterior row of eyes straight or very slightly procurve (Fig. 12); dorso-ven-
trally flattened abdomen in females........................................ Pachistopelma
– Anterior row of eyes strongly procurve (Fig. 13); abdomen not flattened.....8
8 Spermathecae very short and broad, with distal half strongly sclerotized (Fig.
284); male palpal bulb with well-developed keels (Figs 285, 288) (Hispaniola
Island)................................................................................Antillena gen. n.
– Spermatheca long, not strongly sclerotized; male palpal bulb lacking keels..... 9
9 Urticating setae type II very slender (Fig. 18) on a conspicuous patch on ab-
domen dorsum (Fig. 254) (Caribbean Islands: Martinique, Puerto Rico and
U. S. Virgin Islands) (Fig. 226)...........................................Caribena gen. n.
– Urticating setae type II stout (Figs 15–17) spread over most abdominal area
(Figs 302–304)..........................................................................................10
10 Spermathecae lacking an accentuated outwards curvature medially, with dis-
tal portion far from base (Fig. 14); male with tibial apophyses on legs I and
II.................................................................................................Iridopelma
– Spermathecae with an accentuated outwards curvature medially (Fig. 21);
male lacking tibial apophysis on leg II........................................................11
11 Female.......................................................................................................12
– Male..........................................................................................................13
12 Spermathecae virtually non-sclerotized (Figs 257–259) (southeastern and
part of northeastern Brazil)...............................................Ybyrapora gen. n.
– Spermathecae with a well-sclerotized area (Fig. 21).......................Avicularia
13 Tibial apophysis on leg I absent (Fig. 308) or tibia with discrete elevation
covered by a cluster of setae on apical portion (Fig. 309)...........................14
– Tibial apophysis on leg I present, well-developed (Fig. 311)......Avicularia (part)
14 Cymbium with a weakly-developed process on retrolateral lobe or process
lacking (Fig. 305)......................................................................................15
– Cymbium with a well-developed process on retrolateral lobe (Figs 177, 280).... 16
15 Abdomen dorsum with a single stripe (Fig. 297) (southeastern and part of
northeastern Brazil)............................................... .Ybyrapora gen. n. (part)
– Abdomen dorsum with orange spots on its sides (Figs 89) (Venezuela)..........
........................................................................................... Avicularia (part)
16 Male palpal bulb embolus strongly curved (Figs 276–278); tarsi and metatar-
si with dorso-medial conspicuous orange stripe and leg rings on femora, tibia
and metatarsi orange (Bertani and Fukushima 2009, figs A1–A2) (southeast-
ern and part of northeastern Brazil)........................ Ybyrapora gen. n. (part)
– Male palpal bulb embolus curved (Figs 203, 212); tarsi and metatarsi without
orange stripes, and leg rings on femora, tibia and metatarsi whitish or yellow
(Figs 209, 218)................................................................... Avicularia (part)
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 31

Genus Avicularia Lamarck, 1818


Figs 1, 9–10, 13, 15–17, 19–218, 299–300, 302–304, 307, 309, 311–317

Aranea Linnaeus, 1758: 622 (in part: A. avicularia); Fabricius 1775: 438 (in part: A.
avicularia); 1793: 424 (in part: A. avicularia); De Geer 1778: 313, pl. 38, fig. 8 (in
part: A. avicularia); 1783: 122, pl. 38, figs 8–10 (in part: A. avicularia); Latreille
1802: 49 (in part: A. avicularia). Name abandoned as generic name to be used for
the Order name (Thorell 1870: 151).
Mygale Latreille, 1802: 49, 1804: 152, pl. LXII, fig.1 (in part: M. avicularia); 1806: 85
(in part: M. avicularia); Walckenaer 1805: 4 (in part: M. avicularia); Hahn 1820:
5 (in part: M. avicularia); Simon 1864: 70, figs 26–28 (in part: M. avicularia).
Name preoccupied in Mammalia by Mygale Cuvier, 1800 (Thorell 1870: 162).
Avicularia Lamarck, 1818: 107 (type species Aranea avicularia Linnaeus, 1758 by sub-
sequent designation in direction 67 of ICZN (1957: 116)); Thorell 1870: 168;
Ausserer 1871: 128, 201; 1875: 138; Simon 1889: 213; 1892: 171; 1903: 958,
960; F. O. Pickard-Cambridge 1896: 740, 746; 1899: 42; Pocock 1901: 548;
Mello-Leitão 1923: 320, 376; ICZN 1928: 395; Petrunkevitch 1928: 81; Roewer
1942: 253; Bonnet 1955: 826; Valerio 1979: 307; Raven 1985: 119; Gallon 2008:
243; World Spider Catalog 2016.
Eurypelma C. L. Koch, 1850: 73, 74 (in part); Simon 1864: 67 (in part); Roewer 1942:
238 (in part); Bonnet 1955: 1828; Raven 1985: 146, 153. Considered objective
synonym of Avicularia by Raven (1985: 146).
Avicuscodra Strand, 1908: 771 (type species by original designation Avicuscodra ara-
bica Strand, 1908, female, Egypt, El-Tor [28°14'N, 33°37'E], (Tor, Arabien [sic])
Rüppell, SMF 2660, examined); Petrunkevitch 1928: 81; Roewer 1942: 256; Gal-
lon 2008: 243. First synonymized by Gallon (2008: 243).
Ancylochiros Mello-Leitão, 1920: 41 (type species by original designation Ancylochi-
ros taunayi Mello-Leitão, 1920, immature male, Brazil, Minas Gerais, Mariana
[20°22'S, 43°25'W], J. P. Fonseca leg., MZUSP 327, examined,); 1923: 318; 376;
Roewer 1942: 225; Raven 1985: 149. First synonymized by Raven (1985).
Anchylochyrus Petrunkevitch, 1928: 83 (unjustified emendation per Bonnet 1955);
Strand 1929: 12.

Type species. Aranea avicularia Linnaeus, 1758, by subsequent designation (ICZN 1957).
Species included. Avicularia avicularia, A. caei sp. n., A. glauca, A. hirschii, A. ju-
ruensis, A. lynnae sp. n., A. merianae sp. n., A. minatrix, A. purpurea, A. rufa, A. taunayi
and A. variegata stat. n.
Diagnosis. Avicularia resembles Caribena gen. n., Ybyrapora gen. n., Iridopelma
and Typhochlaena by the procurve anterior row of eyes (Fig. 13). It can be distin-
guished from Typhochlaena by the digitiform apical article of PLS (Fig. 10). It dif-
fers from Iridopelma by male lacking tibial apopysis on tibiae II and female by sper-
mathecae long, with accentuated outwards curvature medially (Fig. 21). It differs from
Caribena gen. n. by stout urticating setae on abdomen dorsum of male and female
32 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 4–14. Aviculariinae characters. 4 Ephebopus murinus, palp with paddle of urticating setae type V
(arrow) on prolateral side 5–6 Heteroscodra sp. 5 black marks (arrow) on dorsal tibiae, metatarsi and tarsi
6 tibia IV incrassate 7 Typhochlaena curumim, sternum as long as wide 8 Typhoclaena seladonia, posterior
lateral spinnerets with domed article (arrow) 9–10 Avicularia merianae sp. n. 9 sternum longer than wide
10 posterior lateral spinnerets with digitiform distal article (arrow) 11 Psalmopoeus sp., stridulatory setae
forming a lyra (arrow) on prolateral maxilla 12 Tapinauchenius sp., straight anterior row of eyes 13 Avicu-
laria avicularia, anterior row of eyes strongly procurve 14 Iridopelma hirsutum, spermathecae.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 33

(Figs 15–17) and by rounded process on cymbium (Fig. 307). From Ybyrapora gen. n.
it can be distinguished by sclerotized spermathecae in females (Fig. 21) and presence
of developed tibial apophysis in males (Fig. 311), except A. minatrix, A. lynnae sp. n.,
A. caei sp. n., and A hirschii that lack tibial apophysis. Males of these Avicularia species
can be distinguished from Y. diversipes comb. n. by the embolus less curved in frontal
view (Fig. 30). From Y. sooretama comb. n. and Y. gamba comb. n. they differ in terms
of the presence of a well-developed process on cymbium (Fig. 307), except A. minatrix.
Avicularia minatrix can be distinguished from Y. sooretama comb. n. and Y. gamba
comb. n. by the abdomen dorsum black having orange spots on its side (Fig. 89).
Description. Carapace slightly longer than wide, cephalic region slightly raised.
Cephalic and thoracic striae inconspicuous due to setae density. Fovea deep or shallow,
slightly recurve (most species) or straight. Chelicera without rastelum. Eye tubercle
distinct, raised or slightly raised, wider than long. Anterior row of eyes procurve (Fig.
13), posterior recurve or slightly recurve (most species). Clypeus narrow (most species)
or absent. Labium subquadrate, longer than wide, with 50–133 cuspules spaced by one
or two diameters from each other on the anterior third center. Maxillary lyra absent.
Maxilla subretangular, anterior lobe distinctly produced into conical process, inner
angle bearing 85–215 cuspules. Sternum longer than wide, posterior angle acute, not
separating coxae IV. Three pairs of sigilla, some pairs sometimes not evident. Anterior
ellipsoidal or rounded, middle rounded, posterior ellipsoidal (most species) or round-
ed. All positioned one diameter or less from margin. Leg formula: I=IV II III (most
species) or IV I II III (A. avicularia, A. rufa and A. hirschii female). Clavate trichoboth-
ria on distal 2/3 or 1/2 of tarsi I–IV. Tarsi I–IV fully scopulate, IV divided by a band
of setae or divided by a bald stripe. Metatarsi I–II fully scopulate in most species, III
1/3 to 1/2 distal scopulate and IV 1/5 to 1/2 distal scopulate. Metatarsi IV divided by
a row of setae. Scopulae of tarsi and metatarsi I–II very extended laterally giving them a
spatulate appearance (Figs 299–300). Femora IV without retrolateral scopulae. Stridu-
latory setae absent. Legs aspinose. ITC absent; STC without denticles. Posterior lateral
spinnerets digitiform (Fig.10). Typical stout type II urticating setae on dorsal abdo-
men in females and males, 0.36 to 0.72 mm long, 0.11–0.019 mm wide in females
and 0.78 to 1.11 mm long, 0.013–0.025 mm wide in males (Figs 15–17). Male tibiae I
with apophysis (most species) (Fig. 311) or with discrete elevation covered by a cluster
of setae on apical portion (Fig. 309), on prolateral side. Tibial apophysis with a single
branch having well-developed base and grouped spiniform setae distally (Figs 33–35).
Tibiae II lacking apophysis. Male metatarsus I touches retrolaterally tibial apophysis’
setae when folded. Globous bulb with small subtegulum; prominence on prolateral
tegulum developed (most species) (Fig. 314), well-developed (A. variegata stat. n. and
A. juruensis) (Fig. 315), weakly-developed (A. caei sp. n.) (Fig. 313) or absent (A. mi-
natrix) (Fig. 312). Embolus not flattened, without keels, about 3.0 to 3.5 times tegu-
lum’s length (most species) or more than 4 times tegulum’s length (A. lynnae sp. n. and
A. caei sp. n.) in retrolateral view. Embolus medial portion and tegulum’s margin form
an acute angle in retrolateral view (Fig. 323). Embolus with basal part very curved in
frontal view (Figs 312–315), thin distal width and tip narrowing abruptly (Figs 316–
34 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 15–18. Urticating setae type II in Aviculariinae. 15 Avicularia juruensis male (CAS 4), showing
well-developed barbs along all its length 16 Avicularia variegata stat. n. female (IBSP 7900), showing
weakly-developed barbs only near the stalk 17 Avicularia avicularia female morphotype 5 (MNRJ 06916)
from Pando, showing developed barbs along almost entire length 18 Caribena versicolor comb. n. male
(MNHN−AR 4904), very slender setae with barbs along all length. Scale bars = 0.1 mm.

317) or tapering (Figs 318–319). Cymbium subtriangular with subequal lobes, having
a well-developed rounded process on retrolateral lobe (most species) (Fig. 307) or lack-
ing any process (Fig. 162). Cymbium process bearing thick setae (most species) (Fig.
307) or thin setae (Fig. 205). Spermathecae with an accentuated outwards curvature
medially, completely separated, and long (Fig. 21). Spermathecae not-twisted (except
A. hirschii) with walls lacking projections or lobes (except A. taunayi) (Figs 21–27).
Spermathecae midwidth as wide as its base width (most species) (Fig. 21) or midwidth
expanded, about 1.5 times the apex width (A. variegata stat. n., A. juruensis and A. tau-
nayi) (Fig. 54). Spermathecae with weakly-sclerotized area shorter than half the length
of well-sclerotized area (most species) (Fig. 21) or at least same size of well-sclerotized
area (A. purpurea and A. merianae sp. n.) (Fig. 154). Abdomen dorsum of females
with homogeneously distributed setae (most species) (Fig. 304) or with long guard-
setae grouped on lateral and dorsal anterior areas (Figs 302–303). Legs and palps with
long guard-setae having homogeneous coloration along its length (A. avicularia, most
morphotypes; A. juruensis morphotype 3, A. purpurea, A. taunayi and A. merianae sp.
n.) (Fig. 299) or long guard-setae with darker base and contrasting whitish apex (A.
avicularia morphotypes 6 and 7; A. variegata stat. n.; A. rufa; and A. juruensis mor-
photype 2) (Fig. 300). Leg rings on distal femora, tibiae and metatarsi whitish (most
species) (Fig. 299) or yellowish (A. avicularia, morphotype 5; A. variegata stat. n.,
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 35

Figure 19. Map showing records of Avicularia Lamarck, 1818 species in Central and South America.

morphotype 2; A. juruensis morphotype 2; and A. rufa) (Fig. 300). All species except A.
minatrix showing drastic ontogenetic changes on abdominal color pattern. Brownish
juveniles lacking metallic green or blue sheen, with black tarsi contrasting with other
lighter articles (except A. purpurea) (Fig. 44). Juveniles having dorsal reddish abdomen
with a single central longitudinal stripe and some transversal black stripes on each side
(except A. hirschii). Central longitudinal stripe disconnected from transversal stripes
(A. avicularia, A. variegata stat. n., A. juruensis and A. rufa) (Fig. 44), connected with
all transversal stripes in each side of abdomen (A. purpurea and A. minatrix) (Fig. 167),
connected only with the anterior pair of transversal stripes (A. merianae sp. n.), or con-
nected with first and second anterior pair of transversal stripes (A. taunyai) (Fig. 101).
Distribution and habitat. Costa Rica, Panama, Trinidad and Tobago, Venezuela,
Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Bolivia and Brazil. In
36 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Brazil, it occurs in the states of Roraima, Amapá, Amazonas, Pará, Maranhão, To-
cantins, Acre, western Bahia, Rondônia, Mato Grosso, Goiás, Minas Gerais, São Paulo,
and the Distrito Federal (Fig. 19). Valerio (1979) reports presence of Avicularia species
in Costa Rica (see note on A. avicularia redescription and on A. glauca taxonomy).
Beside this, there are reports in Lago Catemaco and other localities in the state of Verac-
ruz, as well as in the state of Chiapas, Mexico (R. West and J. Mendoza, pers. comm.),
but no Mexican specimens could be examined. Thus, despite the reliable records, it is
not possible to know which species are in Mexico.
Remarks. When erecting the genus Avicularia, Lamarck (1818) included in it three
species described originally in Aranea, in this order: Avicularia canceridea (Lamarck,
1818), Avicularia blondii (Latreille, 1804), and Avicularia fasciata (Latreille, 1804).
The first species, A. canceridea, could be found in the Antilles in “Meridional Ameri-
ca” (South America)(Lamarck 1818). As synonyms of A. canceridea, Lamarck (1818)
cited Aranea avicularia Linnaeus, 1758 (sensu Linnaeus 1758 and Fabricius 1793)
and Mygale avicularia Latreille, 1804 (sensu Latreille 1804 and Walckenaer 1805). It
is unclear why Lamarck (1818) did not use the pre-existent specific name avicularia
instead of canceridea.
Thorell (1870) considered Aranea avicularia Linnaeus “as the type for Avicularia”,
and Avicularia canceridea Lamarck and Aranea vestiaria De Geer as its synonyms. Ap-
parently, this was the reason for the Lamarckian combination Avicularia canceridea
being buried.
Ausserer (1871) considered Avicularia canceridea Lamarck, Aranea avicularia Lin-
naeus, Aranea vestiaria De Geer and Mygale avicularia Latreille as synonyms; however,
he considered valid the binomy Avicularia vestiaria and stated it as the type species.
F. O. Pickard-Cambridge (1899) also considered Avicularia canceridea Lamarck as a
synonym of Avicularia avicularia, but stated this last one as the type species. Despite
the specific epithet avicularia being the most used for the type species of Avicularia
over the years, it was only officially considered as such in 1928 through Opinion 104
(ICZN 1928) and introduced in the official list of specific names in Zoology through
Direction 67 of ICZN (ICZN 1957).
Herein, we tried to establish the real identity of Avicularia avicularia as the original
description is uninformative and useless for species identification. The type locality,
America, is also vague and the existence of types was controversial. Taking into con-
sideration that Aranea avicularia is a very old species described by Linnaeus in the 10th
Edition of Systema Naturae (Linnaeus 1758), it would be a difficult task to track a type.
Apparently, Linnaeus (1758) did not designate holotypes. According to Papavero
and Llorente-Bousquets (1995), the oldest citation regarding the type concept as we
use, nowadays, is dated 1817 by the German dipterologist C. R. W. Wiedmann. In
his publication, Wiedmann (1817 apud Papavero and Llorente-Bousquets 1995) ap-
proved the idea about designation of types in description of new species and creation
of a Normalmuseum where all types should be deposited. Therefore, Linnaeus could
not determine types since this concept was elaborated many years after publication of
Systema Naturae (Papavero and Llorente-Bousquets 1995).
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 37

Linnean holotypes are also a problem for other specialists. Many botanists, for
example, argue it is very difficult to know when Linnaeus used a well-defined speci-
men to describe a plant species. Thus, The Linnean Plant Name Typification Project
was created to choose types from the specimens and illustrations that Linnaeus used in
arriving at his concept of a species in question (Linnean Society of London 2016). The
creation of this project strengthens the ideia that Linnaeus did not designate holotypes.
Specifically in Avicularia’s case, this idea is supported by the expressed citation by F.
O. Pickard-Cambridge (1896) that the type is a figure of two specimens drawn by
Merian (1726), and not a specimen.
Even with a doubtful existence of the A. avicularia holotype, Gabriel et al. (2007)
tried to locate it in The Linnean Society of London Collection. The authors found two
specimens of Aranea avicularia considered by the curator as Linnean material (Gabriel
et al. 2007) and observed they are two different species: a female of Stromatopelma
calceatum Fabricius, 1793; and a male of Harpactira atra Latreille, 1832, both African
species (Gabriel et al. 2007). They concluded that none of the African specimens in
the Linnean collection could be considered types of A. avicularia (Gabriel et al. 2007).
The ICZN states on the article 72.1.1 of its Code that “in the absence of holotype
designation, or the designation of syntypes, or the subsequent designation of a lecto-
type, all [specimens] are syntypes and collectively they constitute the name-bearing
type” (ICZN 1999). Additionally, the article 72.4.1. (ICZN 1999) states that “the
type series of a nominal species-group taxon consists of all the specimens included by
the author in the new nominal taxon (whether directly or by bibliographic reference)”.
Thus, as there is no designation of a specific holotype in the original description, all
the specimens used in the publications mentioned by Linnaeus in the description of
Aranea avicularia are part of a syntypic series.
In that description, Linnaeus (1758) cited his own previous work (Linnaeus 1749),
as well as Clusius (1611), Laet (1633), Piso and Marcgrave (1648), Worm (1655),
Olearius (1666) and Merian (1705).
The spider drawn by Clusius (1611) seems, in fact, to be an aviculariine, since it
has a hairy body and very developed subungueal tufts (Fig. 2). The spider was illus-
trated with low cephalic region and could be found in Baya de todos los Sanctos [sic]
(Clusius 1611). The animal depicted resembles specimens of Pachistopelma spp., which
can be found in Baía de Todos-os-Santos, a bay in the city of Salvador, state of Bahia,
northeastern Brazil. Laet (1633) and Piso and Marcgrave (1648) also reproduced this
illustration and indicated that this species is very common in Saint Aleixo Island (state
of Pernambuco, Brazil), area where Pachistopelma rufoniger specimens are frequent and
where also there is no record of Avicularia spp. Other authors (Olearius 1666, Worm
1655) used the same Clusius’ spider illustration to refer to this Brazilian species which
we found to be a Pachistopelma species.
Merian (1705), on the other hand, illustrated two spiders with very developed and
conspicuously colored subungueal tufts, unmistakably Avicularia sp. specimens (Fig. 1).
One of them is leaving a typical retreat and feeding on ants and the other specimen is
feeding on a small bird. Smith (2000b) stated that, probably because of this illustration,
38 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Linnaeus used the specific name “avicularia”, meaning “small bird” in Latin. Merian
(1705) had drawn these spiders based on specimens observed during her expedition to
Suriname. In the illustration’s comments, there is no indication of locality, nor any infor-
mation that could lead to an inequivocal identification of the illustrated species.
Linnaeus (1758), in his Aranea avicularia description, also cited two illustrations of
Seba (1734) of a spider from Ceylon, now Sri Lanka, which is probably a Poecilotheria
specimen, and an unidentified specimen from America, which clearly is not an aviculariine.
Thus, among the syntypic A. avicularia series are specimens of Pachistopelma spp.
(Clusius 1611, Laet 1633, Piso and Margrave 1648, Worm 1655, Olearius 1666),
Avicularia (Merian 1705) and unidentified specimens (Seba 1704). Apparently, the
specimens used and/or illustrated by these authors are lost or have never been deposit-
ed in a zoological collection, but only observed in their natural habitat. Smith (2000a)
believe that Linnaeus (1758) used the engraving done by Merian (1705) for describing
Aranea avicularia, which was considered perfectly acceptable at that time. However,
Smith (2000a) did not discard the possibility that Linnaeus had examined specimens
collected by Merian, which could be deposited in particular collections, although the
author states that much of Merian’s material is lost.
The ICZN Code article 72.4.1.1 states that “for a nominal species or subspecies
established before 2000, any evidence, published or unpublished, may be taken into
account to determine what specimens constitute the type series” (ICZN 1999). The
Code also considered part of the syntypic series specimens of which there is evidence
that they were known to the author and recognized by him when the nominal species
was established (see example in article 72.4.1.1 of ICZN 1999). After searching some
collections, we found three specimens of Aranea avicularia belonging to the Linnean
collection at the Museum of Evolution of Uppsala University. Even though there is no
irrefutable proof that these specimens once belonged to Linnaeus and were examined
by him when describing the species (Dr. Mats Eriksson, pers. comm.), we think that
two of three specimens found in the collection were used by Linnaeus in his work. The
spiders’ donor was Adolf Fredrik, King of Sweden in the time Linnaeus was writing
his Systema Naturae (Kullander 2001). The King had an extensive biological collection
composed of plants and animals from all over the world, a common habit of wealthy
and significant people of that time (Kullander 2001). During nine weeks between
1751 and 1754, Linnaeus, during that time a professor at Uppsala University, was
housed in the Royal Castle to catalogue the royal collection; the result of this work
was published in a book (Kullander 2001). It is probable that the King donated some
material to Linnaeus, which were brought with him to Uppsala instead of going with
most of the King’s material to the Swedish Museum of Natural History in Stockholm.
Thus, those specimens of Aranea avicularia could have been really used by Linnaeus
(1758) to describe the species and, therefore, they are part of the syntypic series.
King Adolf’s specimens are in a dry, pinned collection, thus they cannot be proper-
ly examined and handle (Fig. 20). However, even by photography, it is clear that both
are Avicularia avicularia specimens, probable from the morphotype found in the state
of Pará, Brazil, since morphological features such as size and coloration match with
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 39

those seen in specimens found in that area. Therefore, herein, we establish the speci-
men UUZM 61 from the Uppsala collection as lectotype and the specimen UUZM
62 as paralectotype, solving the oldest and one of the most vexing problems in myga-
lomorph taxonomy.

Key to Avicularia species

Male of A. glauca and female of A. caei sp.n. and A. lynnae sp. n. are unknown.

Female

1 Spermathecae long and twisted (Fig. 183).................................... A. hirschii


– Spermathecae not-twisted............................................................................2
2 Spermathecae with lobes from median to distal portions (Fig. 91).....A. taunayi
– Spemathecae lacking lobes...........................................................................3
3 Spermathecae midwidth expanded, 1.5 times its base and apical widths (Fig.
54)...............................................................................................................4
– Spermathecae midwidth as wide as its base width (Fig. 21)..........................5
4 Legs with or without grizzled setae, brownish guard-setae on dark abdomen,
and whitish or pale yellow rings on distal femora, tibiae and metatarsi (Figs
119, 122) (see morphotype characteristics and occurence areas).... A. juruensis
– All legs with grizzled setae, vivid reddish guard-setae on black abdomen, and
whitish rings on distal femora, tibiae and metatarsi (Fig. 70); or all legs with
grizzled setae, brownish guard-setae on black abdomen and pale yellow rings
on distal femora, tibiae and metatarsi (Fig. 74).............. A. variegata stat. n.
5 Leg IV longer (more than 10%) than leg I...................................................6
– Leg IV as long as leg I..................................................................................7
6 All legs always with grizzled setae, vivid yellow rings on distal femora, tibiae
and metatarsi, and abdomen with grey guard-setae grouped on lateral dorsal
anterior areas and black short body setae (Fig. 145)............................A. rufa
– Legs with or without grizzled setae, whitish or pale yellow rings on distal fem-
ora, tibiae and metatarsi and guard-setae grouped on lateral dorsal anterior
area (Figs 36, 38–42, 45, 48) (see morphotype characteristics and occurence
areas)........................................................................................A. avicularia
7 Abdomen dorsum black with lateral orange spots on its sides (Fig. 88), Ven-
ezuela.......................................................................................... A. minatrix
– Abdomen dorsum with homogeneous color pattern, lacking orange spots.......8
8 Carapace and legs with green sheen (Fig. 52), Panama.................... A. glauca
– Carapace and legs without green sheen, western South America..................9
9 Carapace and legs with intense purple sheen (Figs 168, 170), abdomen black
velvety (Fig. 304), Ecuador, Colombia and Peru........................ A. purpurea
– Carapace and legs with discrete pink sheen, abdomen with red brownish setae
(Fig. 199), Peru............................................................... A. merianae sp. n.
40 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Males

1 Embolus long, more than 4 times tegulum’s length (Figs 202, 211)............2
– Embolus 2.5 to 3.5 times tegulum’s length (Fig. 29)...................................3
2 Abdomen dorsum with lateral stripes (Fig. 218) and tegulum with weakly-
developed prominence (Fig. 313)............................................. A. caei sp. n.
– Abdomen dorsum with a single longitudinal stripe and tegulum with devel-
oped prominence (Fig. 209)................................................. A. lynnae sp. n.
3 Tibia I with a discrete elevation covered by cluster of setae in apical portion,
on prolateral side (Fig. 309).........................................................................4
– Tibia I apophysis with well-developed branch bearing thick setae (Fig. 311).... 5
4 Tegulum without prominence (Fig. 312), cymbium lacking process on retro-
lateral lobe (Fig. 84) and abdomen dorsum black with lateral orange spots on
its sides (Fig. 89)......................................................................... A. minatrix
– Tegulum with developed prominence (Fig. 175), cymbium with developed
process on retrolateral lobe (Fig. 177)........................................... A. hirschii
5 Cymbium with retrolateral lobe lacking any process or with weakly-developed
process (Fig. 162)............................................................................................ 6
– Cymbium with retrolateral lobe with well-developed process bearing thick
setae (Fig. 307)............................................................................................7
6 Carapace and legs with intense purple sheen (Figs 169, 171), abdomen black
velvety, Ecuador, Colombia and Peru........................................ A. purpurea
– Carapace and legs with discrete pink sheen, abdomen with red brownish
guard-setae, Peru............................................................. A. merianae sp. n.
7 Leg IV longer (more than 10%) than leg I...................................................8
– Leg IV as long as leg I..................................................................................9
8 All legs always with grizzled setae, distal femora, tibiae and metatarsi with vivid
yellow rings and grey guard-setae on black abdomen (Fig. 300).............A. rufa
– Legs with or without grizzled setae, distal femora, tibiae and metatarsi with
whitish or pale yellow rings (Figs 43, 46, 49) (see morphotype characteristics
and occurrence areas)................................................................A. avicularia
9 Tegulum with developed prominence (Fig. 95), abdomen with three pairs of
light brown spots extending from the dorsum to lateral region (Fig. 103)......
..................................................................................................... A. taunayi
– Tegulum with well-developed prominence (Fig. 315), abdomen dorsum
lacking spots..............................................................................................10
10 Legs with or without grizzled setae, abdomen dark with brownish guard-setae,
and distal femora, tibia and metatarsi with whitish or pale yellow rings (Fig.
121) (see morphotype characteristics and occurrence areas)........... A. juruensis
– Legs with grizzled setae, abdomen black with brownish guard-setae and distal
femora, tibia and metatarsi with whitish or pale yellow rings (Figs 71, 75)....
...................................................................................... A. variegata stat. n.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 41

Avicularia avicularia (Linnaeus, 1758)


Figs 1, 13, 17, 19–51, 299, 303, 311, 314, 316, 323

Aranea avicularia Linnaeus, 1758: 622 (lectotype UUZM 61 and paralectotypes


UUZM 62, UUZM 235 here designated, A. Fredrik leg., examined by photos);
Fabricius 1775: 438; 1787: 346; 1793: 424; De Geer 1778: 313, pl. 38, fig. 8;
1783: 122, pl. 38, fig. 8; Latreille 1802: 49; ICZN 1957: 116.
Mygale avicularia: Latreille 1804: 152, pl. LXII, fig. 1; 1806: 32; Walckenaer 1805:
4; Hahn 1820: 5, t. 3; 1833: 101, tab. 25, fig. 75; C. L. Koch 1841: 73, tab. CC-
CXIII, fig. 737.
Mygale scoparia C. L. Koch, 1841: 54, fig. 725. First synonymized by F. O. Pickard-
Cambridge 1896: 741.
Eurypelma avicularia: C. L. Koch 1850: 73; Simon 1864: 67. Considered objective
synonym per Raven 1985: 146.
Avicularia velutina Simon, 1889: 213 (types not found; topotypes female and male,
Venezuela, state of Carabobo, Puerto Cabello [10°27'N, 68°00'W], Collection
E. Simon, MNHN–AR 4883, examined); Petrunkevitch 1911: 51; Mello-Leitão
1923: 377; Roewer 1942: 255; Schiapelli and Gerschman 1945: 191, pl. XVII;
Bonnet 1955: 833. Syn. n.
Avicularia avicularia: Simon 1892: 171, fig. 120; F. O. Pickard-Cambridge 1896:
741, pl. 33, figs 10–11, pl. 34, fig. 19, pl. 35, fig. 13; Strand 1907a: 224; Strand
1907b: 89; Petrunkevitch 1911: 49; ICZN 1928: 395; Mello-Leitão 1923: 320,
324, 376; Bücherl 1957: 404, figs 91–91a; Valerio 1979: 307, figs 7–10; Roewer
1942: 253; Bonnet 1955: 827; 1957: 119; World Spider Catalog 2016.
Avicularia exilis Strand, 1907a: 220 (holotype, male, South America, perhaps Suri-
name, no further information, Coll. Kirschbaum, MWNH 418, examined by
photo); Petrunkevitch 1911: 50; 1939: 286; Mello-Leitão 1923: 377; Roewer
1942: 254; Bonnet 1955: 830; World Spider Catalog 2016. Syn. n.
Avicularia ancylochyra Mello-Leitão, 1923: 330 (holotype, immature male, Brazil, state
of Pará, Rio Tapajós [4°44"S, 56°36'W], E. Garbe col., E2977 C3141, examined);
Petrunkevitch 1939: 284; Schiapelli & Gerschman de Pikelin 1945: 210, lam.
XXIV; Bonnet 1955: 827. Syn. n.
Avicularia cuminami Mello-Leitão, 1930: 56, fig. 3 (holotype, immature, Brazil, Rio
Cuminá [1°21'N, 56°04'W], G. Cruls col., MNRJ 21, examined); Petrunkevitch
1939: 286; Roewer 1942: 254; World Spider Catalog 2016. Syn. n.
Avicularia nigrotaeniata Mello-Leitão, 1940: 178, fig. 1 (holotype, immature male,
Guyana, Kamara, Rio Cuyuni [6°24'N, 58°46'W], C. W. Richards col., Oxford
University expedition, 22 November 1929, n°. 6290, BMNH 1930.12.14.168,
examined); Brignoli 1983: 134; World Spider Catalog 2016. Syn. n.
Avicularia ancylochira: Roewer 1942: 253; World Spider Catalog 2016 (considered as
the spelling apparently intended by Mello-Leitão, 1923).
Avicularia cuminamensis: Bonnet 1955: 830 (invalid emendation).
42 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Remarks. Avicularia velutina syntypes are two females from the forest of San Esteban,
Venezuela, and a specimen from a mountain in North Venezuela (Simon 1889). After
a search on Simon’s collection at MNHN, it was found 4 specimens from Venezuela
labeled as A. velutina. One is an immature from San Esteban (MNHN–AR4888), but
its carapace measurement (4.6 mm) is not compatible with the female used on descrip-
tion (13.8 mm). Other two, one male and one female (MNHN–AR4883), are from
Puerto Cabello, a town very close to San Esteban, locality mentioned on the descrip-
tion and where recently was created a National Park. Despite the female having the
carapace measurement (13.25 mm) compatible with the female used in the description
(13.8 mm), its collection data is not exactly the same of the one of the syntype series.
Adding to this, its pair on the vial is a male, not other female as described by Simon
(1889). The other female found in collection is from Caracas (MNHN–AR4894), on
North Venezuela, and it is smaller than the Puerto Cabello’s female. Thus, herein, we
consider the syntypes of A. velutina Simon, 1889 lost. However, considering we exam-
ined topotypes and that their characteristics match with those found in A. avicularia,
only with slight differences on coloration, we decide to establish Avicularia velutina
Simon, 1889 as a junior synonym of A. avicularia (Linnaeus, 1758).
Avicularia exilis holotype is clearly an Avicularia since it has well-developed scop-
ula, anterior row of eyes procurved and tibial apophysis with a single branch having
well-developed base and grouped spiniform setae distally. The specimen has palpal
bulb with developed prominence, leg IV longer than leg I, not grizzled guard-setae on
palps and legs, and overall reddish brown coloration. These characteristics match A.
avicularia, which is found in Suriname, the probable type locality A. exilis. Thus, we
consider A. exilis Strand, 1907 as junior synonym of A. avicularia (Linnaeus, 1758).
Avicularia ancylochyra holotype has tarsi with well-developed scopula, anterior row
of eyes procurved, no spines on legs, urticating setae of type II on abdomen dorsum,
and abdominal pattern similar to A. avicularia, A. rufa, A. juruensis, and A. variegata
stat. n. The leg IV is longer than leg I, it bears whitish legs rings (not vivid yellow)
and type locality is in the state of Pará, Brazil. All these characters match A. avicularia;
therefore, we consider A. ancylochyra Mello-Leitão, 1923 a junior synonym of A. avicu-
laria (Linnaeus, 1758).
Avicularia cuminami holotype has tarsal and metatarsal scopulae laterally expand-
ed, anterior row of eyes procurve and type locality is state of Pará, Brazil. These char-
acteristics match A. avicularia; thus, we consider A. cuminami Mello-Leitão, 1930 a
junior synonym of A. avicularia (Linnaeus, 1758).
Avicularia nigrotaeniata holotype is an immature male lacking spines on legs and
with spatulated scopulae on tarsi and metatarsi. It has characteristics of Avicularia
genus such as anterior row of eyes procurve and abdomen dorsum with lateral stripes
and a black central longitudinal stripe. In Guyana, the type locality, A. avicularia is
the only species found. Thus, we consider A. nigrotaeniata Mello-Leitão, 1940 a junior
synonym of A. avicularia (Linnaeus, 1758).
Diagnosis. Male and female of A. avicularia resemble A. rufa and female of A.
hirschii by having leg IV longer than leg I. Females of A. avicularia can be distin-
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 43

Figure 20. Avicularia avicularia (Linnaeus, 1758), types from Uppsala Collection (UUZM). Above,
lectotype female (UUZM 61), below paralectotype (UUZM 62). Photo: Mats Eriksson, published with
UUZM permission.

guished from those of A. hirschii by the non-twisted spermathecae. Males and females
of A. avicularia can be distinguished from those of A. rufa by having whitish rings on
distal femora, tibiae and metatarsi (most morphotypes, Figs 36, 38–40) or pale yellow
rings (morphotype 5, Fig. 41) combined with legs and palps with long guard-setae
with homogeneous coloration (all morphotypes except 6 and 7, but these have intense
green metallic sheen on carapace and legs, absent in A. rufa).
Material examined. 1 female, Brazil, Pará, Altamira, Paratipuã, Ponto 5, A. P. L.
Giupponi & D. Pedroso col., 12 April 2009 (MNRJ 13995); 2 males and 1 female,
Amapá, Macapá, Parque Zoobotânico (IEPA), C. Costa, P. Magno & C. A. Júlio col.,
20 August 1996 (MNRJ 13659A).
44 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Additional material. No further information: 1 juvenile female (MPEG 5140); 1


male, 1 female (IBSP 2419); 1 male, died in 27 October 1985 (IBSP IBA 414); 1 fe-
male (IBSP 8842); 1 female, IBAMA, died in January 2000, ref. 84526 (IBSP 8031);
1 immature male (IBSP 9916); 1 female (MNHN–AR Box 301); VENEZUELA: 1
juvenile female, no further information, Sullivan det. 2007 (AMNH Ve23); 1 female,
no further information, Steve Brody coll. (AMNH Ve28); Bolívar: Hato La Vergareña
(6°45'N, 63°30'W) 400–500 ft., 25 October 1954, J. J. Wurdack & N. G. L. Guppy
col. (AMNH Ve22); Carabobo: Puerto Cabello [10°27'N, 68°00'W], 1 female, 1
male, no further information, Collection E. Simon (MNHN–AR 4883 Box 302);
Distrito Capital: Caracas [10°29'N, 66°54'W], 1 female, no further information, Col-
lection E. Simon (MNHN–AR 4894); Monagas: Caripito [10°06'N, 63°06'W], 1 fe-
male, 10 May 1942, Tropical Research Expedition, N. Y. Zoological Society, W. H.
Beebe col. (AMNH Ve30); TRINIDAD AND TOBAGO: no further information, 1
female (AMNH TR133); 1 female, M. Nieves col. (AMNH TR131); 1 male (AMNH
TR129); probably from Port Spain sent in by T. H. G. Aigken [sic] (Aitken), 1 male,
1 immature male, Gertsch det. (AMNH TR130); 2 immatures, 3–4 instar group [sic],
A. Bordes col., 1972 (AMNH TR tube); several spiderlings, A. Bordes col. (AMNH
TR165); Couva-Tabaquite-Talparo: Freeport [10°26'N, 61°24'W], 1 female, E. N.
Kjellesvig-Waering col., 1 May 1966 (AMNH TR181); Diego Martin: Fort George
[10°41'N, 61°32'W], POS, 1 male, M. Nieves col., June 1957 (AMNH TR147);
Goodwood Park [10°41'N, 61°34'W] (Goodward Park [sic]), 5 miles NW of Port of
Spain, 300 m elevation, 1 female, 1 male, E. N. Kjellesvig-Waering col., 1 May 1965
(AMNH TR143); 1 immature, E. N. Kjellesvig-Waering col., 30 September 1965,
(AMNH TR176); Port of Spain [10°39'N, 61°31'W]: 1 male, E. N. Kjellesvig-Waer-
ing col., 1 May 1954 (AMNH TR136); 1 female, E. N. Kjellesvig-Waering col., 18
September 1963 (AMNH TR126); 1 female, E. N. Kjellesvig-Waering col., 16 July
1965 (AMNH TR139); 1 male, 1 female, E. N. Kjellesvig-Waering col., 18 June
1968 (AMNH TR178); 1 male, R. S. Mathews col., April 1890 (AMNH 1.12); Rio
Claro-Mayaro: La Horquette, Valley Road, 1 immature female, T. H. G. Aitken col.,
6 April 1960, #78 (AMNH TR138); Maraval [10°41'N, 61°31'W], 3 males, 2 fe-
males, R. C. West col., 12 May 1981 (AMNH RW09, RW11, RW12, RW13, RW10,
respectively); Mayaro Beach area [10°14'N, 61°00'W], 2 miles N of Plaisance, inside
helicopter hangar, 2 juvenile females, E. N. Kjellesvig-Waering col., 20 December
1964 (AMNH TR182); under coconut husks, 2 males, E. N. Kjellesvig-Waering col.,
22 March 1964 (AMNH TR157); 2 m elevation, 1 male, E. N. Kjellesvig-Waering
col., 22 April 1964 (AMNH TR134); ex House, 2 m, 1 male, E. N. Kjellesvig-Waer-
ing col., 1 November 1964 (AMNH TR171); 1 juvenile female, E. N. Kjellesvig-
Waering col., 5 September 1965 (AMNH TR167); 1 male, E. N. Kjellesvig-Waering
col., 10 June 1966 (AMNH TR174); 3 m, 1 male, E. N. Kjellesvig-Waering col., 5
January 1966 (AMNH TR152); Saint George: Carenage [10°41'N, 61°35'W], 1 juve-
nile female, 29 April 1966, E. N. Kjellesvig-Waering col. (AMNH TR177); Nariva
Swamp, bush bush forest [10°22'N, 61°02'W], Trinidad Regional Virus Lab., 1 juve-
nile female, T. H. G. Aitken col., 20 August 1962 (AMNH TR51); 1 male, T. H. G.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 45

Aitken coll., ex Boar, 12 February 1964 (AMNH TR156); 2 males, T. H. G. Aitken


col., 29 September 1963 (AMNH TR132); Saddle Road (W61°, N10°), 1 juvenile T.
H. G. Aitken col., May 1964 (AMNH TR166); Sangre Grande: Brigand Hill [10°29'N,
61°04'W], 1 juvenile female, L. N. Sorkin col., 21 July 1979, shed 31 July 79, 7
March 80, 10 February 81 and died 28 May 82 (AMNH TR141); Mount Harris
[10°29'N, 61°06'W], 1 male, M. Nieves col., July 1959 (AMNH TR168); 1 male, M.
Nieves col., June 1959 (AMNH TR151); 1 male, M. Nieves col., July 1959 (AMNH
TR135); 1 female, 1 immature, M. Nieves col., July 1959 (AMNH TR145); Siparia:
Fyzabad [10°10'N, 61°32'W] (Fyzbad [sic]), Bamboo stump, no. 226, 1 female, T. S.
Jones col., 21 February 1945 (AMNH TR172); GUYANA: no further information, 1
female, 1914 (CAS13); 1 female, 2 immature males, 1914 (CAS12); Berbice Oriental-
Corentyne: Canje (5°70'N, 57°50'W), Ikuruna River, forest savanna, 2 males, August−
December 1961, G. Bentley col. (AMNH BG78, BG79); Lonsdale Village [6°11'N,
57°31'W] (Larsdale Village [sic]), 26 spiderlings, 19 juveniles, 2 females, E. B. Berbice
col., 9 September 1946 (AMNH BG83); Cuyuni-Mazaruni: Bartica [6°24'N,
58°37'W], 15 milles above, Rio Moraballi and Rio Esequibo, 1 immature, R. W. G.
Hingston col., labeled as A. nigrotaeniata n. 43 (BMNH 1930.4.15.53); Kartabo
[6°24'N, 58°37'W], 2 males, 1 female, captured in a small bush (AMNH BG42); 1
female, 1924 (AMNH BG73); 1 male, 1924 (AMNH BG93); 2 males, 1 female,
Beebe col., “A. surinamensis” det. Di Caporiacco (AMNH BG67); Demerara-Mahai-
ca: Georgetown [6°48'N, 58°09'W], 1 small female, 7 immatures, J. Moral col., 1954
(AMNH BG82); 2 males, L. van Sertima col., 7 May 1991 (AMNH RW65, RW66);
2 immatures, A. McKee col., 19 April 1986 (AMNH RW64); Potaro-Siparuni: Tu-
matumari [5°15'N, 59°08'W], 1 male, 1 female, 5 August 1913, Acc 4435 (AMNH
1.9); 1 immature, August 1913, Acc: 4435 (AMNH 1.11); Upper Takutu-Upper Es-
sequibo Superior: Dadanawa [2°50'N, 59°31'W], Ishertor, 5 females, 2 immatures, in
webbed holes in ground, Terry-Holden Expedition, 7–9 November 1937 (AMNH
BG61); 2 females, 2 immature males, 1 immature, 1 spiderling, 7–9 November 1937,
Terry–Holden Expedition, Snedigar & Hassler col. [hand note: original label and part
of material in another jar] (AMNH BG76); SURINAME: no further information, 1
female, Box 306 3739 (MNHN–AR 4887); in house, 1 female, Geiskes col. (AMNH
Su62); 1 male, Box 306 3739 (MNHN–AR 4887); 1 female, C. Heller col., Decem-
ber 1908, jar no.1632/08 (ZMB); Paramaribo: Paramaribo [5°49'N, 55°10'W], 1
male, Reed col., 29 June 1910, Acc 3792 (AMNH Su60); 2 females, H. Heyde col.,
30 July 1980 (AMNH RW61, RW62); 1 female, 1 immature, 7 March 1939, Geiskes
col. (AMNH Su59); 1 female, C. Heller col., 16 June 1908, jar no.1136/08 (ZMB);
Agronomic Station, 1 male, V. Doesburg col., August 1962 (AMNH Su101); Marowi-
jne: Langamankondre [5°42'N, 54°01'W], 1 female, B. Malkin col., 15–30 August
1965 (MZUSP 10861); 7 females, 1 male, 5 immature males, 8 juveniles, 1 spider-
ling, B. Malkin col., 15–30 August 1965 (AMNH Su97); Para: no further informa-
tion, 1 female, C. Heller col., 13 March 1909, jar no. 541/09 (ZMB); 1 female and 1
immature male, C. Heller col., 10 March 1909, jar no. 541/09 (ZMB); 1 female, C.
Heller col., February 1908, jar no. 684/08 (ZMB); Brokopondo: Kabelstation [4°54'N,
46 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

55°08'W] Riheroever [?, unreadable], in bromeliad, 1 juvenile female, Gerskes col., 25


September 1938 (AMNH Su63); in a house, 1 female, Gerskes col., 26 September
1938 (AMNH Su62); FRENCH GUIANA: Cayenne: Camopi [3°09'N, 52°20'W],
Yanioré, high Oyapock, 100 to 150 m, en amont de Saut, 1 male, Mission E. Aubert
de La Rüe col., February 1949 (MNHN–AR 4886); Cayenne [4°55'N, 52°19'W], 1
female, “Avicularia anthracina” det. E. Simon, Milinon (MNHN–AR 155h-77); 1
male, “Avicularia anthracina” det. E. Simon (MNHN–AR 1719-79); 1 male, J.
Moonen col., February 1992 (AMNH RW60); Table Du Mahury [4°52'N, 52°16'W],
1 female, J. Geay col.; 1902 (MHNP 3089); Montabo [4°56'N, 52°18'W], (Bordat
Montabo [sic]) 1 immature male, Expedição Instituto Butantan col., 4 July 1955
(IBSP 3458); 1 female, Expedição Instituto Butantan col. (IBSP 3379); 1 female, Dr.
Hoge col., 5 July 1951, in nest (IBSP 3377); Montsinery [4°53'N, 52°30'W], Emer-
ald Jungle Village, 1 female, 3 males, J. Moonen col., 12 August 1998 (AMNH
RW57, RW56, RW58, RW59, respectively), RW59 and RW58 on lodge roof beam;
1 female, R. C. West col., April 1999 (AMNH RW55); Saint-Georges-de-l’Oyapock
[3°53'N, 51°48'W] (Oyapock [sic]), 1 male, F. Geay, 1900 (MNHN–AR 1293); Sin-
namary [5°22'N, 52°57'W] (Linamany [sic]), 1 female, 1 male, 1 juvenile female,
Expedição Instituto Butantan col. (IBSP 3396, IBSP 3398, IBSP 3397); Saint Laurent
du Maroni: Saint Laurent du Maroni [5°29'N, 54°01'W] (du Maroni [sic], ?), 1 fe-
male, Gambey col. (MNHN–AR 4901); Saint Jean [5°24'N, 54°4'W] (Saint-Jean-du-
Maroni [sic]), 1 male, 1 spiderling, R. Benoit col., March–April 1914 (MNHN–AR
4896); BRAZIL: Amapá: no further information, 1 male, F. Baia col., ref. 65469
(IBSP 8846); 2 males, 2 females, Dr. Hoge col., 4 August 1966 (IBSP 3760); Macapá
[0°02'N, 51°04'W], Fazendinha, 1 male, 1 immature male, Dr. Hoge col., 4 August
1965 (IBSP 3736); 1 juvenile female, P. Magno col., October 1997 (MNRJ 12923);
1 male (MNRJ 12912); 6 males, C. Costa, P. Magno & C. A. Júlio col., 20 August
1996 (MNRJ 13659); Parque Zoobotânico (IEPA), 2 immatures, P. Magno, C. E.
Alvarenga & C. Costa col., October 1995 (MNRJ 13653); Pacoval [0°33'N, 51°03'W],
1 male, Dr. Hoge col., 21 January 1966 (IBSP 3755); 1 female, 2 juvenile females, Dr.
Hoge col., 26 September 1966 (IBSP 3795, IBSP 3797); Serra do Navio [1°38'S,
52°16'W], 1 male (IBSP 3685); 3 males, K. Lenko col., 8 November 1957 (IBSP
3490); Paredão Pacoval, 1 male, Dr. Hoge col., 4 March 1965 (IBSP 3704); 1 female,
March 1985 (MNRJ 13526); 1 juvenile female, 7 immatures, 1 spiderling, March
1985 (MNRJ 13597); Laranjal do Jari [1°05'N, 53°13'W], Cachoeira do Santo
Antônio, Rio Jari, 2 juvenile females, J. A. P. Barreiros col., 18–24 February 2003
(MPEG 0174); Amazonas: Fazenda São Francisco, 300 km subindo o Rio Negro, 1
male, G. Ilutseh col., 27 May 1982 (IBSP 4814); Pará: no further information, 1 fe-
male, J. C. Branner col., 1881 (CAS); Altamira [3°11'S, 52°12'W], trilha caverna cli-
nas, Ponto 2, 1 female, A. P. L. Giupponi & D. Pedroso col., 10 April 2009 (MNRJ
13993); Ponto 2 , Cararaô, 2 females, A. P. L. Giupponi & D. Pedroso col., 10 April
2009 (MNRJ 13994, MNRJ 1400); Ponto 5, Paratipuã, 1 female, A. P. L. Giupponi
& D. Pedroso col., 12 April 2009 (MNRJ 13995); Acampamento do Juruá, 2 females,
2 immatures, A. J. Cardoso, C. F. B. Haddad & M. Gordo col., 03–18 December
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 47

1986 (ZUEC); Ananindeua [1°21'S, 48°22'W], 1 female, R. F. da Silva col., 30 Janu-


ary 1975 (MPEG 5399); BR 316, Km 06, R. F. da Silva col., 1 male, 10 November
1996 (MPEG 5127); Socêgo [sic], Bacia do Sol, 1 female, 1 immature, March 1956
(IBSP 3781); Barcarena [1°34'S, 48°35'W], Ilha das Onças, Rio Piramanha, 2 males,
E. Santos col., 19–20 April 2003 (MPEG 0172); Belém [1°27'S, 48°30'W], 3 females,
1 juvenile female, P. Cerveira col., refs 63979, 63999 (IBSP 8849, IBSP 8854, IBSP
8853, IBSP 8855, respectively); 1 juvenile female, Eq. Resgate de Fauna col., ref.
55334-3 (IBSP 8571); 1 female, 2 males (IBSP 3578); 1 immature male, 4 females
(IBSP 3772); 1 male, 2 females and 1 immature (IBSP 3780); 1 male, 1 female, 15
June 1966 (IBSP 3770); 3 females, 1 immature, 15 June 1966 (IBSP 3771); 2 females,
15 June 1966 (IBSP 3779); 1 male, R. Cerqueira col., 1972 (ZUEC); 1 male, F. Lima
col., 18 April 1989 (MCP 2702); 1 immature female, W. França col., 30 March 1987
(MCP 2697); 1 male, 2 January 2003 (MPEG 1108); C. Galeno col., 1 immature,
June 1998 (AMNH RW14); Bairro de Souza, 1 male, L. M. Cunha col., 07 March
2005 (MPEG 5147); Base Aérea, 1 male, 1 female (IBSP 3687); Bragança, 1 imma-
ture, 15 June 1966 (IBSP 3777); campus de Pesquisa, 1 juvenile, J. Dias col., 30
March 1992 (MCP 2695); CPATU (em dendê), 2 males, Izaías col., 15 February
1984 (MPEG 135); Empresa Brasileira de Pesquisa Agropecuária (EMBRAPA), 1 fe-
male, R. F. da Silva col., 03 November 1977 (MPEG 5089); EMBRAPA in dendê
area, 1 female, 3 juveniles, 1 spiderling, October–November 1981, H. N. da Cunha
col. (MNRJ 13012); Instituto Agronômico do Norte (IAN), 1 immature male, 1 fe-
male, 4 immatures, A. R. Hoge col., 29 October to 5 December 1958 (IBSP 3672);
IPEAN, 1 male, J. Bushell col., 23 May 1967, Exline–Peck collection (CAS); Mocam-
bo, 1 female, R. F. da Silva col., 14 November 1977 (MEPG 3074); Mocambo,
Parque Regional de Manutenção, Parque Ambiental de Belém, CPPTU, 1 male, M.
A. S. de Azevedo & J. N. Ferreira col., 31 March 2005 (MPEG 5155); Serraria, IPAN,
1 male, P. Waldir col., 06 December 1966 (MPEG 5107); Universidade Federal Ru-
ral da Amazônia (UFRA), 1 female, W. C. Carrasco col., 08 July 2004, F18 A43 B3
(MPEG 5295); 1 juvenile female, Mara col., 15 June 2005, F4 A18 B3 (MPEG
14221); 1 female, 14 June 2004 (MPEG 5188); Vila Ajutei, 1 male, Augusto Côrrea
col., 14 July 2002 (MPEG 0170); Museu Paraense Emílio Goeldi, campus de pesquisa
[01°27'S, 48°26'W], 1 male, W. França col., 04 June 1984 (MPEG 5120); 1 male,
Jarilson col., June 2009 (MPEG 5299); 1 male, J. Dias col., 16 June 1986 (MPEG
5126); 1 female, D. F. Candiani col., April 2006 (MPEG 5300); 1 female, M. C.
Santos-Costa col., 02 March 2005 (MPEG 5221); 1 male, L. S. Carvalho col., 18
Abril 2006 (MPEG 5098); 1 male, A. B. Bonaldo col., 24 July 2002 (MPEG 227); 1
male, D. F. Candiani col., 22 Abril 2004 (MPEG 5094); 1 male, 08 April 2002
(MEPG 5117); 1 male, 31 March 2009 (MPEG 5632); 1 male, 11 April 2005 (MPEG
5141); 2 males, D. F. Candiani col., 12 July 2004 (MPEG 5281); 1 male, A. B. Bon-
aldo col., 12 July 2002 (MPEG 160); 2 immatures, 25 May 1912, Bluntschli-Peyer
coll. (AMNH 1.27, AMNH 1.3); 1 immature, Bluntschli-Peyer col., 25 June 1912,
(AMNH 1.18); Icoaraci [1°18'S, 48°28'W], 1 male, R. F. da Silva col., 10 March
1977 (MEPG 3082); 1 juvenile female, R. F. da Silva col., 05 July 1984, (MPEG
48 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

5131); 1 male, R. F. da Silva col., 8 July 1977 (AMNH RW15); 1 male, R. F. da


Silva col., 18 May 1977 (AMNH RW18); 1 female, R. F. da Silva, 8 April 1977
(AMNH RW16); 1 male, R. F. da Silva col., 8 July 1977 (AMNH RW17); Outeiro
[1°16'S, 48°28'W], 1 juvenile female, R. F. da Silva col., 13 July 1978 (MEPG 5096);
Universidade Federal do Pará (01°28'S, 48°27'W), 1 female, C. Castro col., 24 June
2005 (MPEG 4820); 1 male, L. T. Miglio col., 15 September 2002 (MPEG 1109);
Utinga [1°25'S, 48°24'W], 1 immature, Oliveira & Wygodzinsky col., 10–21 No-
vember 1963 (AMNH 1.2); Belém, Ilha de Cotijuba (01°14'S, 48°35'W), 1 female,
27 August 2003 (MPEG 4701); 1 female, Cotijuba 0011 (MPEG 5291); 1 female, R.
F. da Silva col., 17 March 1977 (MPEG 3073); 1 female, R. F. da Silva col., 26 De-
cember 1977 (MEPG 3030); 1 male, R. F. da Silva col., 11 September 1977 (MPEG
3056); 1 male, R. F. da Silva col., 11 September 1977 (MPEG 3057); Icoaraci,1 fe-
male, 01 July 2002 (MPEG 1111); Belém, Ilha de Jutuba [1°14'S, 48°31'W], 2 males,
3 females, R. F. da Silva col., 30 June 1977 (MPEG 3047, MPEG 3046, MPEG
3045, MPEG 3034, MPEG 3033, respectively); Belém, Ilha do Mosqueiro [1°06'S,
48°23'W], 1 female, 1 immature male, P. Cerveira col., December 1990, ref. 63979
(IBSP 7884, IBSP 7883); 1 male, M. L. Mocambira col., 15 December 1989 (MCP
2699); 2 juvenile females, B. Mascarenhas & team col., 19–25 April 1998 (MPEG
5392, MPEG 5395); Belém, Ilha Nova [2°12'S, 49°27'W], 2 males, 2 females, R. F.
da Silva col., 20 June 1977 (MPEG 3040, MPEG 3043, MPEG 3071, MPEG 5090);
1 male, R. F. da Silva col., 24 June 1977 (MPEG 3041); Belém, Ilha de Paquetá
[1°16'S, 48°32'W], 2 females, R. F. da Silva col., 15 November 1977 (MPEG 3025,
MPEG 3023); 1 juvenile female, R. F. da Silva col., 28 May 1977 (MPEG 5088); 1
male, R. F. da Silva col., 15 September 1977 (MPEG 3052); 1 male, 1 female, R. F.
da Silva col., 19 June 1977 (MPEG 3078, MPEG 3062); 1 male, R. F. da Silva col.,
15 May 1977 (AMNH RW21); 1 male, R. F. da Silva col., 20 May 1977 (AMNH
RW22); 1 immature male, R. F. da Silva col., 15 November 1977 (AMNH RW20);
1 juvenile female, R. F. da Silva col., 26 November 1977 (AMNH RW19); Belém,
Ilha de Tatuoca [1°12'S, 48°30'W], 1 female, 23 March 1978 (MPEG 5105); 2 males,
EPA col., August 1969 (MZUSP 10849); Belém, Ilha de Urubuoca [1°19'S, 48°27'W],
1 female, 1 male, R. F. da Silva col., 28 July 1977 (MEPG 3036, MPEG 3313, res-
pectively); Belterra [3°08'S, 55°03'W], 1 male, 18 December 2003 (MPEG 4703);
Rio Assuá [?, unreadable], 1 female, H. Sioli col., May 1952 (IBSP 3130); Benevides
[1°21'S, 48°14'W], Pratinha, Estrada do Açucareiro, 1 female, O. Cunha col., 19
February 1975 (MPEG 141); Breves [1°40'S, 50°28'W], Corcovado, 1 male, Expedi-
ção CDZ col., 18–19 October 1965 (IBSP 5688); Castanhal [1°17'S, 47°55'W], 1
female, O. Feassi col., Ref. 52599 (IBSP 8863); Ilha de Marajó [0°58'S, 49°35'W], 1
male, 1 female, 16 immatures, Instituto Agronômico col., 15 June 1966 (IBSP 3787,
IBSP 3786, respectively); Santo André, 1 male, June–September 1912, Bluntschli-
Peyer (AMNH 1.23); Cachoeira do Arari [1°0'S, 48°57'W], Jabuti, 1 female, L.
Macambira col., 12 December 1990 (MPEG 0138); Itaituba [04°16'S, 55°59'W], 1
female (MZUSP 27659); Itupiranga [5°08'S, 49°19'W], campus avançado da UFPA,
3 females, 2 immatures, A. R. Hoge & P. Villela col., December 1971 (IBSP 4097);
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 49

Jacareacanga [07°15'S, 57°26'W], 1 immature male (IBSP 3538); (Jacare-acanga


[sic]), 1 spiderling, M. Alvarenga col., December 1968 (AMNH 1.20); Aproeste, km
350 da Transamazônica, 1 female, B. V. Rodrigues & N. Abrahim col., 27 October
2009 (MPEG 15640); Jatobal [5°08'S, 56°51'W], 3 males, January 1975 (IBSP
7896); Juruti [2°09'S, 56°05'W], Ferrovia Km 07, 1 immature female, A. C. Lima &
F. E. Pimenta col., 17 Maio 2007, JURU 006 0286 (MPEG 15635); Ferrovia km 36,
1 female, A. Lima & F. Pimenta col., 24 May 2007, JURU 0060268 (MPEG 15636);
Platô Capiranga, Linha 168E (02°28'22.1"S, 56°12'29.4"W), 1 male (MPEG 15639);
Juruti, Acampamento Barroso (02°28'10.5"S, 56°00'3.5"W), 1 male, D. F. Candiani
col., 07 August 2004, ref. JURU002 0044 (MPEG 2000); 2 males, 1 female, D. R.
Santos-Souza col., 12 July 2002, ref. JURU 002 0084 (MPEG 2003, MPEG 2006,
respectively); 1 female, D. F. Candiani col., 12 July 2002, ref. JURU 002 0083
(MPEG 2009); Juruti, Acampamento Mutum (01°36'44.77"S, 56°11'39.2"W), 1 ju-
venile female, 1 immature male, D. F. Candiani and D. R. Santos-Souza col., 03
August 2004, ref. Juruti 2004 (MPEG 15634); Marabá [5°22'S, 49°07'W],
Transamazônica, 3 males, J. Navas col., 15 March 1975 (IBSP 4190); 1 female, M. F.
Torres col., 14 September 1985 (MPEG 4231); 1 male, P. Tumma col., 8 October
1973 (IBSP 2495); Melgaço, Estação Científica Ferreira Penna, FLONA Caxiuanã
[2°01'S, 51°39'W], 1 female, A. B. Bonaldo col., 21–30 November 2000 (MPEG
1923); Mocajuba [2°34'S, 49°30'W], Praia do Imbaubal, 1 male, 17 June 1984
(MPEG 5104); Monte Alegre [1°59'S, 54°04'W], Lagoa Grande, 1 immature, P. de
Biasi col., 10 March 1979, ref. 24.434 (IBSP 4408); Óbidos [1°54'S, 55°31'W], 1
female, E. Garbe col., 1920 (MZUSP 10856, old collection number 556); Oriximiná
[1°45'S, 55°51'W], 1 immature male, 1 March 1968, EPA col. (MZUSP 10850); 1
immature, in pineapple, EPA col., 13 January 1968 (MZUSP 10854); Porto Trombe-
tas [1°28'S, 56°22'W], 1 male (IBSP 8847); Ourém [1°29'S, 47°10'W], Patauateua, 1
juvenile female, D. D. Guimarães col., 06 December 2002 (MPEG 213); Paragomi-
nas [3°00'S, 47°21'W], Fazenda Brejeiro, 1 male, C. Junqueira Netto col., 5 October
1970, ref. 5741 (IBSP 84); Canindé [2°33'S, 46°30'W] (Canindé, Maranhão [sic]), 1
immature female, May to August 1965, J. Carvalho col. (AMNH 1.16); Rio Cuminá
Mirim [1°14'N, 55°52'W], As Pedras, 1 female, 1 immature, EPA col., 29 September
to 4 October 1969 (MZUSP 1085); Rio Tocantins [02°04'S, 49°18'W], Ilhas das
Cobras, Castanheira, 1 female, A. J. da Silva col. (MPEG 5100); Rio Trombetas
[1°28'S, 56°22'W], 3 males, 2 females, 1 immature male, F. Palinger col., (IBSP 8762,
IBSP 8761, IBSP 8759, respectively); Jacaré, 3 males, female, Expedição CDZ col., 20
September to 13 October 1965 (IBSP 5687); Santarém [2°26'S, 54°41'W], 1 female,
E. Garbe col. (MZUSP 10864, old collection number 557); 1 male, G. Puorto col.,
December 1996 (IBSP 8578); Fazenda Taperinha, 1 female, EPA col., 1–11 February
1968 (MZUSP 10859); 1 female, 1–11 February 1968, E.P.A. col. [handwritten la-
bel], or Manaus, Lenko col., 12 September 1962, forest [museum label] (MZUSP
10858); Santo Antônio do Tauá [1°09'S, 48°07'W], Uxitena, 1 male, R. F. da Silva
col., 07 April 1977 (MPEG 3076); Tomé-Açu [2°25'S, 48°09'W], Roda D’Água (ca-
poeira), 1 female, J. Dias col., 19 June 1994 (MEPG 143); UHE Tucuruí [3°46'S,
50 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

49°40'W], Equipe de Resgate de Fauna col.: 2 males, 1 female, ERF col., 1984, IBA
243 (IBSP 7898); 1 female, ERF col., ref. 48286 (IBSP 8845); 1 female, ERF col.,
1984, ref. IBA 383 (IBSP 7880); 1 male, ERF col.,1984, ref. IBA 1002 (IBSP 7879);
1 female, ERF col., 1984, ref. 47607 (IBSP 7882); 2 females, ERF col., 1984, ref. IBA
283 (IBSP 7887); 1 female, ERF col., ref. IBA 600 (IBSP 8851); 1 female, ERF col.,
ref. CB-1 (IBSP 8568); 1 female, ERF col., (IBSP IBA 380); 1 female, born in capitiv-
ity, died in 06 July 1993 (IBSP IBA 083-43); 1 female, ERF col., IBA 599 (IBSP
8850); 1 female, ERF col., 12 December 1984 (IBSP 4832); 1 female, ERF col., ref.
34C XXXV (IBSP 8857); 1 male, ERF col., ref. 48286 (IBSP 8848); 1 male, ERF col.,
ref. 47607 (IBSP 8573); 1 male, ERF col., 18 November 1986, IBA 860 (IBSP 4925-
B); 2 males, ERF col., ref. CB-1 (IBSP 8579); 1 male, ERF col.(IBSP 8575); 2 fe-
males, ERF col. (IBSP 8843); 1 male, ERF col., ref. B5677 (IBSP 8580); 1 male, ERF
col., 4 June 1987, IBA 277 (IBSP 8566); 1 female, ERF col., 1984, ref. IBA 283
(IBSP 7888); 1 female, ERF col., August 1984, ref. IBA 734 (IBSP 7877); 1 female,
ERF col., ref. IBA 283 (IBSP 8862); 1 male, in bromeliad in a tree (21 m from the
soil), C. Pantoja col., 28 June 1984 (IBSP 7916); 1 male, Equipe de Resgate de Fauna
col., 1984, IBA 292 (IBSP 7885); 1 male, Equipe de Resgate de Fauna col., 1984, ref.
47607 (IBSP 7881); picada do Inajá, A3, lote 8, 1 female, B. Mascarenhas col., 20
June 1980 (MZUSP 9480); Base 4, Bravo, 1 female, Operação Resgate Faunístico
col., 12 December 1984 (IBSP 4832); Vila Bravo, 1 female, Equipe de Resgate de
Fauna col., ref. XXXVII-87 base 4 (IBSP 8844); Vila Cisipé, 1 male, Equipe Resgate
de Fauna col., ref. 47607 (IBSP 8572); Ilha de Tocantins, Equipe de Resgate de Fau-
na col.: 1 female, ref. IBA 389 (IBSP 8861); 1 female, ERF col., ref. IBA 599 (IBSP
8569); 1 female, 2 June 1986, ERF col., ref. IBA 1013 (IBSP 8852); 1 female (IBSP
ref. IBA 283); Breu Branco [3°45'S, 49°33'W], 1 male, died 20 February 1986 (IBSP
CB1 - XV); 1 male, Equipe de Resgate de Fauna col., ref CBI V (IBSP 8565); Vigia
[0°51'S, 48°08'W] (Vigia de Nazaré [sic]), Hospital UBSO IV, 1 male, 03 July 2005
(MPEG 5189); UBS V, 1 male, J. F. Maia col., 30 August 2005 (MPEG 5144); Ma-
ranhão: São Luís [2°31'S, 44°18'W], 1 female, G. R. Ristan Filho col., 29 February
1996, ref. 78226 (IBSP 8244); 1 female, ref. 78226 (IBSP unnumbered); Bacabal
[4°13'S, 44°47'W], Lago Verde, 1 female, R. Neto col., 03 February 1983 (MPEG
5106); Palmeirândia [2°40'S, 44°54'W], 1 female, Dr. Hoge, Pedro and Joaquim col.,
7–30 May 1962 (IBSP 3622); Fazenda São Luís, 1 female, 1 immature male, Dr.
Hoge, Pedro & Joaquim col., 7–30 May 1962 (IBSP 3621); São Bento [2°41'S,
44°49'W], 1 immature male, 1 female, Dr. Hoge, Pedro & Joaquim col., 7–30 May
1962 (IBSP 3623, IBPS 3618); Mato Grosso: Alta Floresta [9°52'S, 56°05'W], 1 male,
F. Palinger col. (IBSP 8768); PERU: Madre de Dios: Cuenca Rio Los Amigos [12°34'S,
70°06'W], 1 male (UA 003/2005); 1 female (UA 042/2006); 1 female (UA 007/2005);
Aguajal [12°15'S, 69°16'S], CICRA, 1 female, 21 March 2006 (UA 041/2006); 1
male (UA 005/2005); 1 male (UA 002/2005); Puerto Maldonado [12°36'S, 69°41'W],
15 km East, Rio Madre de Dios, Albergue Lodge Cuzco Amazônico, 200 m, 1 female,
G. C. Hunter col, 8 June 1983 (CAS10); La Cachuela (12°33'S, 69°11'W), 1 male
and 2 females, H.–W. Auer col., September–October 2013 (MUSM–ENT 0506819);
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 51

Tambopata [12°34'S, 69°11'W], 1 female, R. Bennett col., August 1994, in silk re-
treat on palm trees (AMNH RW47); Reserva Tambopata–Candamo [12°59'S,
69°36'W], Albergue Explorer’s Inn, 1 male and 1 female, K. J. N. Villa leg., 2005
(MZUSP 70948); BOLIVIA: Beni: Yucumo [15°08'S, 67°02'W], Aserradero Chapa-
rina, 7 km from Yucumo, path to San Borja, prov. J. Ballivian, 1 male, J. Peñaranda
col., 9 July 1991 (MNRJ 06918); 1 female, same data (MNRJ 06919); Pando: Ma-
nuripi, Camalho 15 km northwest Puesto Castañero, Chivé [12°23'S, 68°34'W],
bosque alto, 1 female, F. Guerra col., 30 September 1991 (MNRJ 06916); La Paz:
Iturralde [12°54'S, 67°45'W], Estância El Dorado, UMSA, Instituto de Ecologia, 1
immature male, S. Beck col., 28 February 1984 (MNRJ 06917); Santa Cruz: San
Antonio del Parapeti [20°01'S, 63°13'W], 1 male, B. Malkin col., 24–25 May 1985
(AMNH Bo01).
Female. Redescription. MNRJ 13995. Carapace: 19.39 long, 17.68 wide, 5.5 high.
Chelicera: 8.40 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 14.66, 8.55,
11.65, 10.56, 8.25, 53.67. II: 14.04, 8.15, 10.56, 10.22, 3.55, 46.55. III: 13.55, 8.20,
10.55, 10.66, 3.11, 46.01. IV: 17.56, 8.65, 14.65, 13.66, 8.12, 62.64. Palp: 10.42,
8.65, 8.76, -, 8.62, 36.45. Midwidth: femora I–IV= 3.45, 3.21, 4.05, 3.50, palp= 2.56;
patellae I–IV= 3.66, 3.55, 3.15, 3.50, palp= 3.05; tibiae I–IV= 3.37, 2.56, 3.26, 3.17,
palp= 2.62; metatarsi I–IV= 2.45, 2.35, 2.40, 2.35; tarsi I–IV= 2.61, 2.65, 2.53, 2.30,
palp= 2.55. Abdomen: 26.37 long, 17.56 wide. Spinnerets: PMS, 2.99 long, 1.62 wide,
0.1 apart; PLS, 2.04 basal, 1.71 middle, 4.07 distal; width 2.73, 2.3, 1.72, respectively.
Carapace: 1.10 times longer than wide; cephalic region slightly raised, thoracic
striae inconspicuous.
Fovea: deep, straight, 2.27 wide.
Eyes: eye tubercle 1.30 high, 2.62 long, 3.52 wide. Clipeus: 0.53. Anterior eye
row procurve, posterior slightly recurve. Eye size and interdistances: AME 0.73, ALE
0.85, PME 0.26, PLE 0.74, AME–AME 0.57, AME–ALE 0.48, AME–PME 0.27,
ALE–ALE 2.35, ALE–PME 0.58, PME–PME 2.15, PME–PLE 0.14, PLE–PLE 2.86,
ALE–PLE 0.38; AME–PLE 0.60.
Maxilla: length to width 1.86. Cuspules: 164 spread over ventral inner heel. Labi-
um: 2.07 long, 3.46 wide, with 82 cuspules spaced by one diameter from each other on
anterior third. Labio-sternal groove swallow, flat, with two slightly separate, large sigilla.
Chelicera: basal segment with 11 teeth in row and some small teeth on promargin.
Sternum: 9.07 long, 8.08 wide. Sigilla: three pairs, posterior and middle rounded,
anterior small, all less than one diameter from margin.
Legs: Formula: IV I II III. Length leg IV to leg I: 1.16. Clavate trichobothria:
2/3 distal tarsi I–IV. Scopulae: Tarsi I–IV fully scopulate; IV with a few sparse setae.
Metatarsi I–II fully scopulate; III 2/3 distal; IV 1/3 distal. IV divided by rows of setae.
Type II urticating hairs: 0.62–0.69 long, 0.017–0.019 wide.
Spermathecae (Fig. 21): two completely separated, not-twisted and long sper-
mathecae, with walls lacking projections or lobes and accentuated outwards curvature
medially. Midwidth as wide as its base width and weakly-sclerotized area shorter than
half the length of well-sclerotized area.
52 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 21–27. Avicularia avicularia (Linnaeus, 1758), spermathecae variation. 21 morphotype 1, Al-
tamira, state of Pará, Brazil (MNRJ 13995) 22 morphotype 2, UHE Tucuruí, state of Pará, Brazil (IBSP
4885) 23 morphotype 3, Puerto Cabello, state of Carabobo, Venezuela (MNHN–AR 4883) 24 morpho-
type 4, Cuenca Rio Los Amigos, department of Madre de Dios, Peru (UA 0042/2006) 25 morphotype
5, Yucumo, department of Beni, Bolivia (MNRJ 06919) 26 morphotype 6, Paramaribo, district of Para-
maribo, Suriname (AMNH Su59) 27 morphotype 7, Juruti, state of Pará, Brazil (MPEG 15640). Scale
bars = 1 mm.

C 6): carapace brown with brown short body setae with green sheen. Carapace
border with long reddish brown setae with pink sheen. Coxae, labium, sternum and
maxillae brown, slightly darker than ventral femora. Legs and palps with brown short
body setae with green sheen and reddish brown guard-setae with homogeneous dark
coloration on anterior legs and guard-setae with darker base and contrasting whitish
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 53

apex on posterior legs. Leg rings on distal femora, tibiae and metatarsi whitish. Abdo-
men dorsum with long reddish brown guard-setae with pink sheen grouped on lateral
dorsal anterior areas and dark short body setae. Ventral abdomen brown.
Male. Redescription. MNRJ 13659A. Carapace: 17.23 long, 16.81 wide, 4.9
high. Chelicera: 7.2 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 15.4,
8.6, 11.8, 11.7, 6.9, 54.4. II: 14.8, 7.7, 11.7, 11.5, 6.7, 52.4. III: 13.2, 6.7, 10.2,
11.5, 6.6, 48.2. IV: 16.2, 7.9, 14.7, 15.6, 6.9, 61.3. Palp: 9.4, 5.3, 7.0, –, 2.9, 24.6.
Midwidths: femora I–IV= 3.0, 2.8, 3.4, 3.1, palp= 2.2; patellae I–IV= 3.1, 3.1, 2.8,
3.2, palp= 2.3; tibiae I–IV= 2.3, 2.2, 2.2, 2.6, palp= 2.2; metatarsi I–IV=1.7, 1.6, 1.5,
1.8; tarsi I–IV= 2.0, 1.9, 1.8, 1.6, palp= 2.2. Abdomen: 19.83 long, 13.39 wide. Spin-
nerets: PMS, 1.75 long, 0.80 wide, 0.1 apart; PLS, 2.82 basal, 2.04 middle, 3.56 distal;
width: 1.88, 1.55, 1.15, respectively.
As in female, except:
Carapace: 1.03 times longer than wide; cephalic region slightly raised, thoracic
striae inconspicuous.
Fovea: deep, straight, 1.70 wide.
Eyes: eye tubercle 1.60 high, 2.23 long, 3.33 wide. Clypeus 0.22 wide. Eyes size
and interdistances: AME 0.71, ALE 0.77, PME 0.27, PLE 0.700, AME–AME 0.62,
AME–ALE 0.48, AME–PME 0.22, ALE–ALE 2.24, ALE–PME 0.78, PME–PME
1.99, PME–PLE 0.12, PLE–PLE 2.58, ALE–PLE 0.46, AME–PLE 0.51.
Maxilla: length to width: 2.13. Cuspules: 157 spread over ventral inner heel. La-
bium: 1.73 long, 2.40 wide, with 94 cuspules.
Chelicerae: basal segment with 10 teeth in row and some small teeth on promar-
gin. Sternum: 8.58 long, 7.35 wide. Sigilla: three pairs, all rounded and large, less than
one diameter from margin.
Legs: Length leg IV to leg I: 1.13. Metatarsi I–II fully scopulate, III scopulate in
distal 2/3; IV, in distal 1/3. IV divided by wide row of setae.
Type II urticating hairs: 0.94–1.09 long, 0.020–0.025 wide.
Palp (Figs 28–31): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 4.99 long in retrolateral
view, about 3.0 times tegulum’s length. Medial portion and tegulum’s margin form an
acute angle in retrolateral view (Fig. 323). Proximal part very curved in frontal view;
thin distal width, abruptly narrowing distally; basal, middle, and distal width 0.49,
0.21, 0.07, respectively. Tegulum: 2.66 long, 1.72 high in retrolateral view. Cymbium
subtriangular with subequal lobes, with well-developed rounded process on retrolateral
lobe, bearing thick setae (Fig. 32).
Tibial apophysis (Figs 33–35): single branch on prolateral leg I, with well-devel-
oped base and grouped spiniform setae distally. Male metatarsus I touches retrolater-
ally tibial apophysis’ setae when folded.
Color pattern: carapace brown with brown short body setae with green and golden
sheen. Carapace border with long setae the same color as dorsal carapace short body
setae. Coxae, labium, sternum and maxillae brown, slightly darker than ventral fem-
ora. Legs and palps with brown short body setae with green sheen and reddish brown
54 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 28–35. Avicularia avicularia (Linnaeus, 1758), male (MNRJ 13659A). 28–31 right palpal bulb
(mirrored) 28 prolateral 29 retrolateral 30 frontal 31 dorsal 32 right cymbium, dorsal view (mirrored)
33–35 right tibial apophysis of leg I (mirrored) 33 prolateral 34 ventral 35 retrolateral. Scale bars = 1 mm.

guard-setae with homogeneous dark coloration on anterior legs and guard-setae with
darker base and contrasting whitish apex on posterior legs. Leg rings on distal femora,
tibiae and metatarsi whitish. Abdomen dorsum with long reddish brown guard-setae
with pink sheen and dark short body setae. Ventral abdomen brown.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 55

Figures 36–43. Avicularia avicularia (Linnaeus, 1758) morphotypes. 36–37 morphotype 1 36 female
from Castanhal, state of Pará, Brazil 37 immature from Santarém, state of Pará, Brazil 38 morphotype 2,
female from Tucuruí, state of Pará, Brazil 39 morphotype 3, female from Caracas, Distrito Capital, Ven-
ezuela 40 morphotype 4, female from Tambopata, department of Madre de Dios, Peru 41 morphotype
5, preserved female from Manuripi, department of Pando, Bolivia 42–43 morphotype 6 42 female from
Montsinery, department of Cayenne, French Guiana 43 male from Georgetown, department of Deme-
rara-Mahaica, Guyana. Photos: 36 R. Bertani; 37 M. Gamache, 38 W. Bokermann; 39–40, 42–43 R. C.
West; 41 C. S. Fukushima.
56 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 44–49. Avicularia avicularia (Linnaeus, 1758) morphotypes. 44–46 morphotype 6, from Trini-
dad and Tobago 44 immature 45 female 46 male 47–49 morphotype 7, from Juruti, state of Pará, Brazil
47 immature 48 female 49 male. Photos: 44–46 R. C. West; 47–49 F. E. Pimenta.

Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi
contrasting with other lighter articles and abdomen dorsum reddish, with dorsal cen-
tral longitudinal black stripe disconnected from transversal black stripes (Figs 37, 44,
47). When mature, both males and females lose this pattern.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 57

Figure 50. Map showing records of Avicularia avicularia (Linnaeus, 1758) and Avicularia rufa Schiapelli
& Gerschman, 1945.

Distribution. Venezuela, Trinidad and Tobago, Guyana, Suriname, French Guia-


na, Brazil (states of Amapá, Pará, Maranhão, Amazonas, Mato Grosso), and populations
in Peru and Bolivia (Fig. 50). Possibly, on an opposite pattern to A. rufa, A. avicularia
is rarer in western Brazil, the area connected to the region where occur A. avicularia in
Peru and Bolivia. This disjunct distribution pattern of A. avicularia can be better under-
stand only after sampling or having access to material from poorly sampled areas such as
western Amazon (near Brazil-Peru-Colombia border), and Bolivia and Peru as a whole.
Natural history. Stradling (1994) studied distribution and behavioral ecology of a
population of A. avicularia in Trinidad. The author reported that retreats were found
in different situations on vegetation and in human constructions. He found first eight
instars specimens predominantly between leaves of low-growing plants (especially in
Heliconia Linnaeus), holding edges of leaves together with silk to create retreat. After
this life stage, Stradling (1994) noted spiders tended to build their retreats in more
elevated structures, not incorporating leaves in retreat. Stradling (1994) recorded cap-
58 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 51. Map showing records of Avicularia avicularia (Linnaeus, 1758) morphotypes.

ture of katydids, cockroaches, scarabs and lizards by A. avicularia and also observed
courtship behavior with males maturing in beginning of rainy season, in May and
June. Females lay the eggsac in May, and after an incubation period ranging from 29
to 62 days, about 125 spiderlings hatch (Stradling 1994).
Bates (1863) observed a scene similar to the one depicted by Merian (1705) in
Cametá, state of Pará, Brazil. He observed A. avicularia specimen actively feeding on
a small bird inside its webbed retreat on a tree, and also the existence of another small
bird inside the spider’s retreat.
F. O. Pickard-Cambridge (1896) observed that adults constructed their retreat in
“almost any locality which offered a more or less vacant cylindrical space”, including
hollow stumps of Açai palm, folded leaves of bananas, in the hollow center of a pineap-
ple plant and among the palm-leaf thatch of native houses.
Villa (2004) observed in areas at Tambopata district in Peru retreats of A. avicu-
laria built on trunks and in axils of palm tree Attalea butyracea (Mutis ex L.f.) Wess.
Boer and also in Heliconia psittacorum L. plant. The reproduction period in that area is
in September and October (Villa 2004).
Variation. Specimens of A. avicularia show different patterns of coloration which
seems to be correlated with their geographical distribution. However, genitalic and so-
Table 2. Avicularia avicularia morphotypes: characteristics and geographical distribution.

Morpho Morpho Morpho Morpho Morpho Morpho Morpho


1 (Fig. 36) 2 (Fig. 38) 3 (Fig. 39) 4 (Fig. 40) 5 (Fig. 41) 6 (Figs 42–46) 7 (Figs 47–49)
Brazil, Guyana, French
Brazil, Guyana, French Brazil: states of Pará, Bolivia: departments
Occurrence area Peru: department of Guiana, Suriname, Brazil: states of Pará
Guiana, Venezuela and Maranhão and Mato Venezuela of Santa Cruz, La Paz
(Fig. 51) Madre de Dios. Venezuela and and Amapá.
Trinidad and Tobago Grosso and Beni.
Trinidad and Tobago
Overall aspect brownish brownish brownish brown reddish greyish greyish light brownish
Legs and palps discrete grizzled or not discrete grizzled or discrete grizzled or discrete grizzled or not
not grizzled very grizzled very grizzled
guard-setae grizzled not grizzled not grizzled grizzled
Legs and palps short brown with very intense brown with discrete brown with brown with green brown with very brown with very brown with discrete
body setae green sheen golden sheen golden sheen and pink sheen intense green sheen intense green sheen golden sheen
Leg rings whitish whitish whitish whitish pale yellowish whitish whitish
brown, gradually
Females: abdomen, orange brownish,
reddish with pink sheen reddish brown reddish brown reddish brown reddish lightening from base
guard-setae, color with whitish tip
to tip
Females: posterior
legs, guard-setae vivid red vivid red reddish vivid red greyish greyish vivid orange
color
dark brownish,
Males: abdomen, dark brownish, some
dark brownish dark brownish dark brownish dark brownish dark brownish some with whitish
guard-setae, color with whitish tip
tip
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818...
59
60 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

matic characters are very homogeneous along the distribution of the species. Variations
among different populations concerning color patterns are, herein, discussed. We do not
consider the detected differences sufficient to elevate each morphotype to species status.
Nonetheless, we do not discard the possibility of they corresponding to criptic species. In
order to establish more accurate limits in A. avicularia, it is necessary to employ multiple
approaches, considering molecular, ecological, behavioral and geographic data.
Besides body coloration, there is urticating setae variation. Females of morphotype
5 have urticating setae with developed barbs along almost all lengths (Fig. 17), differ-
ent from the other morphotypes (Fig. 16). However, as we had only access to three
specimens it is not possible to make more considerations about it.
Remark. Valerio (1979) reported the presence of A. avicularia in Costa Rica
(Fig. 19). In fact, the male palpal bulb and spermathecae illustrated by the author are
compatible with Avicularia species. However, the specimens from Costa Rica reported
by Valerio (1979) were examined and are more similar to A. purpurea than to the larger
species of Avicularia. See discussion in Avicularia glauca taxonomy.

Avicularia glauca Simon, 1891


Figs 19, 52–53

Avicularia glauca Simon, 1891: 312 (holotype female, Panama, MNHN–AR 4897,
examined); F. O. Pickard-Cambridge 1896: 744, 1899: 42; Petrunkevitch 1911:
50; Mello-Leitão 1923: 377; Roewer 1942: 254; Bonnet 1955: 831; World Spider
Catalog 2016.

Remark. The examined specimen is not labeled as holotype. However, it is the only
specimen in Simon’s collection in which locality and size are compatible with the de-
scription (Simon 1891). Therefore, we consider it as the holoype.
The small specimen (carapace length 9.7 mm) has tarsal and metatarsal scopulae
expanded, giving a spatulated aspect, characteristic of Aviculariinae (Fig. 52). A single
spermatheca is present: the other is supposed lost. It has an accentuated outwards
curvature medially and lacks lobes, as most species of Avicularia (Fig. 53). Thus, it
undoubtedly belongs to Avicularia. Its spermatheca has a short, well-sclerotized area
that resembles the spermathecae of A. purpurea, with which it seems to be very closely
related. Furthermore, it has the overall metallic green color exhibited by young and
small specimens of A. purpurea. The specimens found in Costa Rica and reported by
Valerio (1979) as A. avicularia were examined by the second author (RB). They are,
in fact, definitely not A. avicularia. Despite many searches, we did not find any other
specimen from Panama or Costa Rica in other arachnological collections, indicating
Avicularia is a rare genus in southern Central America. Therefore, we conclude that
A. glauca can be a valid species from Panama and Costa Rica. The identity of Costa
Rica’s Avicularia will be better discussed in another paper, in preparation with a Costa
Rican colleague.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 61

Figures 52–53. Avicularia glauca Simon, 1891, holotype female (MNHN–AR 4897). 52 habitus
53 spermathecae.

Avicularia variegata (F. O. Pickard-Cambridge, 1896), stat. n.


Figs 16, 19, 54–77, 315, 317

Avicularia avicularia variegata F. O. Pickard-Cambridge, 1896: 743, pl. 33, fig. 12 (lec-
totype female, here designated, Brazil, Amazonas, Itacoatiara [3°07'S, 58°26'W],
Lower Amazon, January 1896, BMNH 1896.12.13.16; and paralectotype female,
here designated, Brazil, Amazonas, Itacoatiara [3°07'S, 58°26'W], in banana tree,
7 February 1896, BMNH 1896.12.13.17; examined); Strand 1907b: 90; Petrunk-
evitch 1911: 49; Mello-Leitão 1923: 326, 376; Roewer 1942: 254; Bücherl 1957:
404; World Spider Catalog 2016.
Avicularia bicegoi Mello-Leitão, 1923: 329, figs 187, 189 (holotype subadult female,
Brazil, Amazonas, Manaus [03°06'S, 60°01'W], Bicego col., MZUSP 133, exam-
ined); Roewer 1942: 254; Bonnet 1955: 830; Bücherl 1957: 404, figs 92–92a;
World Spider Catalog 2016. Syn. n.

Remarks. Avicularia bicegoi holotype has spermatheca midwidth expanded, about 1.5
times its basal and apical portion widths (Fig. 57); leg IV as long as leg I; whitish leg
rings on distal femora, tibiae and metatarsi, and the type locality is Manaus, Amazonas.
These characteristics match A. variegata stat. n.; thus, we consider A. bicegoi Mello-
Leitão, 1923 junior synonym of A. variegata (F. O. Pickard-Cambridge, 1896) stat. n.
Diagnosis. Females of A. variegata stat. n. resemble A. juruensis and A. taunayi
by the spermatheca midwidth expanded, about 1.5 times its basal and apical portion
widths (Figs 54–57). They differ from females of A. taunayi by lacking lobes on sper-
mathecae. Females can be distinguished from those of A. juruensis by the abdomen
with vivid reddish brown guard-setae with pink sheen grouped on lateral and dorsal
anterior areas, contrasting with the black short body setae and black venter (morpho-
type 1) (Fig. 70) or by legs and carapace with golden short body setae with green sheen,
and pale yellow rings on distal femora, tibiae and metatarsi (morphotype 2) (Fig. 74).
Males of A. variegata stat. n. resemble those of A. avicularia, A. rufa, A. juruensis, A.
taunayi, A. purpurea and A. merianae sp. n. by having tibial apophysis on leg I with
well-developed base and grouped spiniform setae distally (Fig. 63). They differ from
males of all these species except A. juruensis by the well-developed prominence on
tegulum (Fig. 60). Males and females of A. variegata stat. n. can be distinguished from
62 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 54–57. Avicularia variegata (F. O. Pickard-Cambridge, 1896) stat. n., spermathecae varia-
tion. 54 morphotype 1, Manaus, state of Amazonas, Brazil (INPA 4894) 55 holotype, Itacoatiara, state
of Amazonas, Brazil (BMNH 1896.12.13.16) 56 morphotype 2, Alto Alegre, state of Roraima, Brazil
(MZUSP 70946) 57 holotype of Avicularia bicegoi Mello-Leitão, 1923, Manaus, state of Amazonas,
Brazil (MZUSP 133). Scale bars = 1 mm.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 63

A. juruensis by lacking intense purple sheen on carapace and legs (Figs 70, 71, 74, 75)
or by occurrence area: Brazil, state of Amazonas (especially Manaus region) to state of
Roraima, and Venezuela (Fig. 76).
Material examined. 1 female, Brazil, Amazonas, Manaus [03°06'S, 60°01'W], Par-
quejo, R. Oliveira-Filho col., 27 May 2008 (INPA 4894); 1 male, Brazil, Amazonas,
Manaus [03°06'S, 60°01'W], Praça 14, M. P. Sena col., 27 March 1980 (INPA 4897).
Additional material. VENEZUELA: 1 juvenile female, D. Grimaldi col., March
1989 (AMNH Ve21); Distrito Capital: Caracas [10°29'N, 66°54'W], Avila Mountains,
1 juvenile female and 1 immature, on trees, C. Siederman col., August 1991, (AMNH
Ve18); Monagas: Caripito [10°06'N, 63°06'W], 1 male, 15–31 March 1942, Venezue-
la Expedition, Dept. Tropical Research, N. Y. Zool. Society, W. Beebe col., high and
low jungle trails (AMNH Ve31); Amazonas: Puerto Ayacucho [5°39'N, 67°38'W],
reg. 22, 2 males, in trees, C. Siederman col., May 1993 (AMNH Ve29); BRAZIL:
Roraima: Alto Alegre [2°53'N, 61°29'W], 1 female, C. M. Moraes ded., April 2013,
(MZUSP 70946); Amajari, Vila Tepequém (03°47'54"N, 61°44'57"W), 1 male,
17 November 2008, Yamaguti & Pinto da Rocha col. (MZUSP 70945); Boa Vista
[2°49'N, 60°40'W], Balneário Água Boa, 1 male, 6h00, S. M. B. Lima col., 02 January
2002 (MNRJ 12968); Caracaraí, Estação Ecológica Niquiá [1°49'N, 61°07'W], Hotel
Ecotur, 1 immature male, 07 October 2001 (IBSP 11270); Ilha de Maracá [3°25'N,
61°39'W], 2 males, A. B. Bonaldo col., 31 January to 14 February 1992 (MCP 1969);
2 males, A. Lise leg., 13 January to 14 February 1992 (MCP 1968); Amapá: Macapá,
4 km from Pacoval [0°02'N, 51°04'W], 1 female, Dr. Hoge col., 4 August 1965 (IBSP
3837); [0°03'S, 49°33'W], 1 female, 4 August 1965, Dr. Hoge col., died 26 June
1968 (IBSP 3873); Amazonas: Coari, Porto Urucu, Base de Operações Geólogo Pedro
de Moura, (4°45'47"S, 65°02'41"W), 1 male, Dias et al. col., 2006 (MPEG 15633);
(4°48'45"S, 65°02'01"W ), 1 male, L. T. Miglio col., 09 July 2006 (MPEG 15641);
Itacoatiara [3°07'S, 58°26'W], 1 female, Dirings col., March 1961 (IBSP 4265); Ita-
piranga [2°44'S, 58°01W], 1 immature male, EPA col., 11 September 1968 (MZUSP
10.860); Manaus [03°06'S, 60°01'W], 2 males, T. Gasnier col. (INPA 4952, INPA
4948); 1 male, H. Höfer leg., 28 March 1988 (INPA 4891); 1 female, 2 February
1948 (AMNH 1.13); 1 juvenile female, February 1943 (AMNH 1.26); Balneário do
SESC, 1 female, R. Freitas col., 07 December 1975 (INPA 4896); conjunto Suframa,
1 juvenile female, Larissa col., 13 November 1997 (INPA 4893); conjunto Acariquara,
1 male, D. M. M. Mendes col., 18 March 2005 (INPA 4895); campus INPA, 1 male,
Albuquerque col., May 1995 (INPA 4887); Estação Ecológica Experimental de Silvi-
cultura Tropical, INPA, 1 female, A. L. R. Barreto col., 29 July 1980 (ZUEC 015);
Estrada Manaus–Caracaraí, km 45, 2 immature males, J. Vasconcellos Neto col., July
1978 (ZUEC 019); INPA, Mata do Laguinho, 1 male and 1 female, 25 April 1959
(IBSP 3503); INPA, campus Peralta, 1 female, F. J. A. col., 13 January 1988 (INPA
4881); Manaus Airport, 1 male, A. Barros col., 27 April 1980 (INPA 4892); Peralta,
1 male, F. J. A. col., 05 June 1993 (INPA 4889); 1 female, F. J. A. col., 03 December
1990 (INPA 4883); Prédio do INPA, V8, 3°andar, Silvicultura, 1 male, P. A. Celes-
64 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 58–65. Avicularia variegata (F. O. Pickard-Cambridge, 1896) stat. n., male (INPA 4897).
58–61 left palpal bulb 58 prolateral 59 retrolateral 60 frontal 61 dorsal 62 left cymbium, dorsal
63–65 left tibial apophysis of leg I 63 prolateral 64 ventral 65 retrolateral. Scale bars = 1 mm.

te col., 19 May 1986 (INPA 4888); Rio Tarumá, off Rio Negro, Manaus [03°06'S,
60°01'W], 1 juvenile female, N. Gordon col., June 1995, in house (AMNH RW32);
Terra Nova [Manaus neighborhood or Amazonas’s town?], 1 male, Mancelu col.,
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 65

07 September 1975 (INPA 4890); Vivenda Verde [Manaus neighborhood?], 1 male,


H. Brandão col., 29 March 2002 (INPA 4885); Maraã [2°17'S, 65°00'W], Rio Japurá,
Maguari, 1 male, R. Constantino col., 02 December 1998 (MPEG 5182); 1 juvenile
female (MPEG 5504); Presidente Figueiredo [02°01'S, 60°01'W], UHE Balbina, 1
female, 2 juvenile females, faunal rescue team col., February 1988, ref. 56112 (IBSP
7872, IBSP 7871, IBSP 7875, respectively); 2 immatures, November 1987, collection
Bokermann no. 886 (MZUSP 32173); 1 female, 1987, ref. 55569-12 (IBSP 7900); 2
juvenile females, 1 immature male, 1987, ref. 56681 (IBSP 7901); 1 juvenile female,
November 1987, ref. IB34 (IBSP 7876); 1 male, 20 November 1987, ref. 55569-20
(IBSP 7873); 1 male, ref. 55854-2 (IBSP 7878); 3 females, 1 male, 2 immatures,
1987, ref. 56681 (IBSP 7902, IBSP 7903); 1 male, 1988, ref. 56572-7 (IBSP 7874);
3 juvenile females, 2 immature males, Eletronorte col., April 1988 (MNRJ 13818);
Cachoeira do Tucumã, Rio Uatumã [2°36'S, 58°05'W], 1 immature (IBSP 7934);
Margens do Rio Uatumã [2°36'S, 58°05'W], 1 female, 1 juvenile female, C. F. Alva-
renga col., October 1987 (MNRJ 13652); Pará: Belém [1°26'S, 48°28'W] 1 female,
Dr. Hoge & João col., September 1952 (IBSP 3119).
Female. Redescription. INPA 4894. Carapace: 19.27 long, 19.06 wide, 4.20
high. Chelicera: 7.25 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I:
16.22, 10.06, 12.58, 10.94, 7.27, 57.07. II: 15.21, 9.21, 11.43, 10.37, 6.93, 53.15.
III: 13.91, 8.39, 11.34, 10.72, 6.95, 51.31. IV: 16.89, 9.04, 14.17, 14.00, 6.94, 61.04.
Palp: 11.19, 7.10, 7.84, –, 8.88, 35.01. Midwidths: femora I–IV= 3.40, 3.67, 3.63,
3.78, palp= 3.08; patellae I–IV= 3.68, 3.86, 4.03, 3.89, palp= 3.18; tibiae I–IV= 3.41,
3.21, 3.24, 3.38, palp= 2.98; metatarsi I–IV= 2.28, 2.52, 2.29, 2.58; tarsi I–IV= 3.20,
3.00, 2.97, 2.97, palp= 3.03. Abdomen: 24.77 long, 16.94 wide. Spinnerets: PMS,
2.34 long, 0.93 wide, 0.31 apart; PLS, 2.70 basal, 1.59 middle, 3.50 distal; midwidths
2.22, 1.83, 1.43, respectively.
Carapace: 1.01 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Fovea: deep, slightly recurve, 2.35 wide.
Eyes: eye tubercle 0.95 high, 2.70 long, 3.71 wide. Clypeus 0.67. Anterior row of
eyes procurve. Posterior row of eyes recurve. Eye size and interdistances: AME 0.86,
ALE 0.89, PME 0.39, PLE 0.58, AME–AME 0.71, AME–ALE 0.64, AME–PME
0.22, ALE–ALE 2.35, ALE–PME 0.91, PME–PME 2.45, PME–PLE 0.08, PLE–PLE
3.10, ALE–PLE 0.55, AME–PLE 0.68.
Maxilla: length to width: 1.84. Cuspules: 100–200 spread over ventral inner heel.
Labium: 2.68 long, 3.26 wide, with 103 cuspules spaced by one diameter in third dis-
tal area. Labium-sternal groove shallow, flattened, with two sigilla.
Chelicera: basal segment with 15 teeth and some small teeth on promargin. Ster-
num: 9.99 long, 8.39 wide. Sigilla: only posterior pair evident, rounded, less than one
diameter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.07. Clavate trichobothria: dis-
tal 2/3 tarsi I–IV. Scopulae: Tarsi I–IV fully scopulate. Metatarsi I–II fully scopulate;
III 2/3; IV 1/3 distal scopulate. IV divided by a wide row of setae.
66 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 66–73. Avicularia variegata stat. n., morphotype 1, habitus 66 immature on leaf retreat 67 im-
mature 68 juvenile 69 juvenile female 70 female 71 male 72 adult inside retreat on tree bark 73 female
in coconut tree. Photos: 66–69, 71–73 Marlus Almeida; 70 R. C. West.

Type II urticating setae (Fig. 16): 0.64–0.72 long, 0.016–0.019 wide.


Spermathecae (Fig. 54): two completely separated, not-twisted long spermathecae,
with walls lacking projections or lobes and accentuated outwards curvature medially.
Midwidth expanded, about 1.5 times its basal and apical portion widths, and weakly-
sclerotized area shorter than half the length of well-sclerotized area.
Color pattern (Fig. 70): carapace brown with greyish short body setae with green
sheen. Carapace border with long setae the same color as dorsal carapace short body
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 67

setae. Coxae, labium, sternum and maxillae dark brown, darker than ventral femora.
Legs and palps with greyish short body setae with green and pink sheen and brown
guard-setae with darker base and contrasting whitish apex. Leg rings on distal femora,
tibiae and metatarsi whitish. Abdomen dorsum with vivid reddish brown guard-setae
with vivid pink sheen grouped on lateral and dorsal anterior areas and black short body
setae. Abdomen venter velvety black.
Male. Description. INPA 4897. Carapace: 19.38 long, 19.08 wide, 5.27 high.
Chelicera: 6.69 long. Legs (femur, patella, tibia, metatarsus, tarsus and total): I: 19.21,
10.36, 14.03, 14.46, 7.62, 65.68. II: 18.52, 9.55, 14.57, 14.15, 7.94, 64.73. III:
16.41, 8.40, 13.17, 13.82, 7.60, 59.40. IV: 19.64, 8.88, 16.32, 18.13, 7.72, 70.69.
Palp: 11.53, 6.64, 9.10, –, 3.78, 31.05. Midwidths: femora I–IV= 3.92, 3.90, 4.09,
3.60, palp= 2.85; patellae I–IV= 3.87, 4.03, 3.86, 3.83, palp= 3.03; tibiae I–IV= 3.06,
3.31, 2.85, 3.18, palp= 2.71; metatarsi I–IV= 2.28, 2.12, 2.00, 1.95; tarsi I–IV= 2.49,
2.45, 2.25 2.31, palp= 2.61. Abdomen: 22.45 long, 14.42 wide. Spinnerets: PMS,
2.31 long, 0.79 wide, 0.16 apart; PLS, 2.35 basal, 1.20 middle, 3.56 distal; midwidths
1.95, 1.63, 1.25, respectively.
As in female, except:
Carapace: 1.02 times longer than wide.
Fovea: 2.59 wide.
Eyes: eye tubercle 1.59 high, 2.67 long, 3.48 wide. Clypeus 0.44. Eye size and
interdistances: AME 0.84, ALE 0.79, PME 0.29, PLE 0.68, AME–AME 0.45, AME–
ALE 0.52, AME–PME 0.27, ALE–ALE 2.53, ALE–PME 0.70, PME–PME 2.23,
PME–PLE 0.20, PLE–PLE 2.68, ALE–PLE 0.52, AME–PLE 0.72.
Maxilla: length to width: 2.05. Labium: 2.19 long, 3.19 wide, with 100 cuspules
spaced by one diameter in third area. Labio-sternal groove with no evident sigilla.
Chelicera: basal segment with 9 teeth and some small teeth on promargin. Ster-
num: 9.93 long, 7.45 wide. Sigilla: only posterior evident, rounded, less than one
diameter from margin.
Legs: Length leg IV to leg I: 1.08. Scopula: Metatarsi III 1/2 distal scopulate.
Type II urticating setae: 0.91–1.00 long, 0.019–0.020 wide.
Palp (Figs 58–61): globous bulb with small subtegulum and well-developed prom-
inence on tegulum. Embolus: not flattened, lacking keels, 5.65 long in retrolateral
view, about 3.5 times tegulum’s length. Medial portion and tegulum’s margin form an
acute angle in retrolateral view. Proximal part very curved in frontal view; thin distal
width, abruptly narrowing distally; basal, middle, and distal width 0.91, 0.31, 0.05,
respectively. Tegulum: 3.04 long, 1.65 high in retrolateral view. Cymbium subtrian-
gular with subequal lobes, having a well-developed rounded process on retrolateral
lobe, bearing thick setae (Fig. 62).
Tibial apophysis (Figs 63–65): single branch on prolateral leg I, with well-devel-
oped base and grouped spiniform setae distally. Male metatarsus I touches retrolater-
ally tibial apophysis’ setae when folded.
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tar-
si contrasting with other lighter articles and abdomen dorsum reddish, with dorsal
68 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 74–75. Avicularia variegata stat. n., morphotype 2, habitus 74 female 75 male. Photos: 74 C.
S. Fukushima; 75 R. Bertani.

central longitudinal black stripe disconnected from transversal black stripes (Fig. 66).
When mature, both males and females lose this pattern.
Distribution. Venezuela and Brazil (states of Roraima, Amapá, Amazonas and
Pará) (Fig. 76).
Natural history. Silva and Meirelles (2016) reported a predation on Troglodytes
musculus (Naumann, 1823) bird by an specimen of A. variegata stat. n. in a urban park
at Manaus, state of Amazonas, Brazil.
Variation. All examined specimens of A. variegata stat. n. have grizzled setae on
palps and legs (except those found in Venezuela), but with slight differences in body
coloration. We detected three morphotypes. Morphotype 1 is found especially near
Manaus (Fig. 77) and females have carapace, legs, and palps with greyish short body
setae with very intense green sheen and whitish leg rings (Fig. 70). Abdomen has vivid
reddish guard-setae grouped on lateral and dorsal anterior areas, black short body setae
and venter velvety black. Males have black abdomen with some whitish tipped setae
homogeneously distributed (Fig. 71). Morphotype 2 is found in same area (Fig. 77),
but has overall greyish coloration, with short body setae with discrete green sheen and
yellowish leg rings. Females have abdomen with light brown guard-setae grouped on
lateral and dorsal anterior areas, black short body setae, and venter dark brown (Fig.
74). Males are very similar to morphotype 1 but with yellowish rings (Fig. 75). Males
from Puerto Ayacucho (AMNH Ve29) and from Caripito (AMNH Ve31), Venezuela,
have genitalia and legs with proportions compatible with those from Brazil, as well as
whitish leg rings. However, they do not have white tipped hairs on legs and abdomen
as specimens of A. variegata stat. n. from Manaus, Brazil (Fig. 77), and the prominence
on palpal bulb is not well-developed as in specimens from Manaus (morphotype 1). The
juvenile female AMNH Ve21, also from Venezuela, has short body setae with golden
sheen and very discrete white tipped setae in legs and palps as well as whitish rings. We
prefer not to consider these forms as separate species because these slight differences
can be due to populational variation since the specimens were collected in the northern
limit of the species distribution. It is here called morphotype 3. Despite being collected
in Caracas, also in the northern limit of the species, the specimen AMNH Ve18 seems
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 69

Figure 76. Map showing records of Avicularia variegata (F. O. Pickard-Cambridge, 1896) stat. n. and
Avicularia juruensis Mello-Leitão, 1923.

to be a typical morphotype 1. It is necessary to examine more material to understand


morphological variation and species boundaries of Venezuelan Avicularia.
In his expedition on Lower Amazonas, F. O. Pickard-Cambridge (1896) made
interesting observations about color variations along the river. According to him,
there was a tendency in Avicularia spp. to have grizzled setae as he went further west
(F. O. Pickard-Cambridge 1896). When F. O. Pickard-Cambridge (1896) described
A. variegata stat. n., he considered it as a subspecies of A. avicularia and pointed
out that some could consider A. variegata stat. n. a species in the differentiation
process—as further westwards it extended, the more evident the differential char-
acters might be. He also observed the presence of intermediate forms at Santarém,
state of Pará, which had the setae of the first two pairs of legs decidedly grizzled
(F. O. Pickard-Cambridge 1896). We failed to find specimens with this pattern
in our studied sample. Some specimens we examined had posterior legs with more
grizzled setae than anterior, but none without any grizzled setae. Despite these two
morphotypes being very distinct, F. O. Pickard-Cambridge (1896) stated a careful
70 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 77. Map showing records of Avicularia variegata F. O. Pickard-Cambridge, 1896 stat. n.
morphotypes.

comparasion was needed of both forms to prove that there are more differences than
just coloration. In fact, we found other differences. Leg IV of Avicularia avicularia is
more than 10% longer than leg I and spermathecae midwidth is as wide as its base
width; whereas leg IV of A. variegata stat. n. has roughly the same length of leg I and
spermathecae midwidth is expanded. Accordingly, we can consider them as separate
taxa. Thus, we elevate the former subspecies Avicularia avicularia variegata F. O.
Pickard-Cambridge, 1896 to species status, resulting in Avicularia variegata (F. O.
Pickard-Cambridge, 1896) stat. n.

Avicularia minatrix Pocock, 1903


Figs 19, 78–90, 312

Avicularia minatrix Pocock, 1903: 81 (holotype female, Venezuela, Lara, Duaca


[10°17'N, 69°09'W], BMNH 1903.7.1.120, examined); Petrunkevitch 1911: 50;
Mello-Leitão 1923: 377; Roewer 1942: 255; Bonnet 1955: 832; Smith 1992: 22,
figs 1–11; Tinter 1993: 10, figs 1–3, World Spider Catalog 2016.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 71

Diagnosis. Females and males of A. minatrix can be distinguished from all other
Avicularia species by the abdominal pattern, vivid orange lateral spots over black
background (Figs 88–89). Additionally, females of A. minatrix resemble females of A.
avicularia, A. rufa, A. purpurea, A. hirschii and A. merianae sp. n. by the spermathecae
midwidth as wide as its base (Fig. 78). They can be distinguished from A. avicularia
and A. rufa by leg IV as long as leg I; from A. purpurea and A. merianae sp. n. by the
spermathecae weakly-sclerotized area shorter than half the length of well-sclerotized
area, and from A. hirschii by the non-twisted spermathecae (Fig. 78). Males of A. mi-
natrix resemble those of A. hirschii, A. lynnae sp. n. and A. caei sp. n. by the tibia I with
a discrete elevation covered by a cluster of setae in apical portion, on prolateral side
(Fig. 86), and can be distinguished from all of them by lacking process on retrolateral
lobe of the cymbium.
Material examined. VENEZUELA: 1 female, M. Baumgarten leg. 1994, ref.
73485 (IBSP 12886); 1 male, pet trade (MZUSP 70949); 1 male, L. Koschorreck col.,
1991 (IBSP 11596).
Female. Redescription. IBSP 12886. Carapace 12.85 long, 12.30 wide, 4.50 high.
Chelicera: 4.09 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 9.38, 6.10,
6.91, 6.39, 4.24, 33.02. II: 8.84, 5.85, 6.56, 6.27, 4.26, 31.78. III: 7.82, 5.15, 5.23,
6.05, 3.49, 27.74. IV: 9.99, 5.07, 7.79, 7.42, 3.93, 34.20. Palp: 6.87, 4.31, 4.08, –,
5.00, 20.26. Midwidths: femora I–IV= 1.99, 2.00, 1.62, 2.13, palp= 2.11; patellae I–
IV= 2.39, 2.24, 2.48, 2.59, palp= 2.26; tibiae I–IV= 2.01, 2.25, 2.01, 2.32, palp= 2.05;
metatarsi I–IV= 1.95, 1.58, 1.59, 1.92; tarsi I–IV= 2.05, 2.02, 1.76, 2.02, palp= 2.12.
Abdomen 14.10 long, 8.91 wide. Spinnerets: PMS, 1.72 long, 0.95 wide, 0.15 apart;
PLS 2.57 basal, 1.68 middle, 2.34 distal; midwidths 1.57, 1.18, 0.93, respectively.
Carapace: 1.05 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous. Carapace covered by long setae.
Fovea: deep, slightly recurve, 2.46 wide.
Eyes: eye tubercle: 0.90 high, 1.97 long, 2.77 wide. Clypeus 0.46. Anterior row
of eyes procurve, posterior row of eyes slightly recurve. Eye size and interdistances:
AME 0.64, ALE 0.59, PME 0.24, PLE 0.58, AME–AME 0.27, AME–ALE 0.36,
AME–PME 0.14, ALE–ALE 1.70, ALE–PME 0.55, PME–PME 1.58, PME–PLE
0.15, PLE–PLE 2.02, ALE–PLE 0.40, AME–PLE 0.47.
Maxilla: length to width: 1.48. Cuspules: 190 spread over inner heel. Labium:
1.87 long, 2.31 wide, with about 50 cuspules spaced by one diamenter of each other
on anterior half (malformed). Labium sternal groove shallow, flat, with two separate,
large sigilla.
Chelicera: basal segment with 8 teeth and some small teeth on promargin.
Sternum: 7.27 long, 4.32 wide. Sigilla: only posterior evident, rounded, very close
to margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.04. Clavate trichobothria:
distal 2/3 of tarsi I–II; distal 1/2 of tarsi II–IV. Scopula: Tarsi I–IV fully scopulate.
Metatarsi I–II fully scopulate, III 2/3, IV 1/3 distal scopulate. IV divided by row of
setae.
72 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 78–79. Avicularia minatrix Pocock, 1903, spermathecae variation. 78 Venezuela (IBSP 12886)
79 holotype, Duaca, state of Lara, Venezuela (BMNH 1903.7.1.120). Scale bars = 1 mm.

Type II urticating setae: 0.38–0.51 long and 0.011–0.013 wide (measured in holo-
type BMNH 1903.7.1.120).
Spermathecae (Fig. 78): two completely separated, not-twisted long spermathecae,
with walls lacking projections or lobes and accentuated outwards curvature medially.
Midwidth as wide as its base width and weakly-sclerotized area shorter than half the
length of well-sclerotized area.
Color pattern (Fig. 88): carapace orange brown with golden short body setae
with pink sheen. Carapace border with long setae the same color as dorsal carapace
short body setae. Coxae, labium, sternum and maxillae orange brown, same color of
ventral femora. Legs and palps with brown short body setae with pink sheen and long
orange guard-setae. Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 73

Figures 80–87. Avicularia minatrix Pocock, 1903, male (MZUSP 70949). 80–83 right palpal bulb
(mirrored) 80 prolateral 81 retrolateral 82 frontal 83 dorsal 84 right cymbium, dorsal (mirrored)
85–87 left tibia I 85 prolateral 86 ventral 87 retrolateral. Scale bars = 1 mm.

dorsum with orange guard-setae homogeneously distributed, vivid orange short body
setae forming lateral spots and black short body setae forming dark background.
Ventral abdomen brown.
74 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 88–89. Avicularia minatrix Pocock, 1903, habitus. 88 female, state of Falcón, Venezuela
89 male. Photos: 88 J. Huff; 89 J. Newland.

Male. Redescription. IBSP 11596. Carapace 10.08 long, 9.49 wide, 2.32 high.
Chelicera 2.84 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 9.40,
5.29, 7.46, 6.76, 3.97, 32.88. II: 8.97, 4.76, 5.73, 5.92, 3.55, 28.93. III: 7.30, 3.89,
5.08, 5.92, 3.58, 25.77. IV: 9.76, 4.66, 7.98, 8.61, 4.00, 35.01. Palp: 5.83, 3.43,
4.17, –, 2.47, 15.9. Midwidths: femora I–IV= 2.02, 1.77, 2.04, 1.60, palp= 1.61;
patellae I–IV= 1.85, 1.76, 1.82, 1.87, palp= 1.48; tibiae I–IV= 1.73, 1.65, 1.72, 1.70,
palp= 1.48; metatarsi I–IV= 1.25, 1.19, 1.19, 1.18; tarsi I–IV= 1.54, 1.33, 1.42,
1.21, palp= 1.75. Abdomen 9.78 long, 5.73 wide. Spinnerets: PMS, 1.06 long, 0.59
wide, 0.18 apart; PLS, 1.66 basal, 0.69 middle, 1.58 distal; midwidths 0.73, 0.59,
0.49, respectively.
As in female, except:
Carapace: 1.06 times longer than wide.
Fovea: 1.56 wide.
Eye: eye tubercle: 1.51 long, 2.24 wide, 0.71 high. Clypeus: 0.20 wide. Eye size
and interdistances: AME 0.58, ALE 0.60, PME 0.26, PLE 0.54, AME–AME 0.25,
AME–ALE 0.18, AME–PME 0.16, ALE–ALE 1.25, ALE–PME 0.45, PME–PME
1.33, PME–PLE 0.13, PLE–PLE 1.69, ALE–PLE 0.10, AME–PLE 0.42.
Maxilla: length to width: 1.98. Cuspules: 100–115 spread over inner heel. La-
bium: 1.04 long, 1.58 wide, with about 80 cuspules spaced by two diameters from
each other on anterior half.
Chelicera: basal segment with 10 teeth decreasing in size from distal area. Ster-
num: 4.95 long, 3.72 wide. Sigilla: three pairs, posterior oval.
Legs: Length leg IV to leg I: 1.07. Scopula: tarsi IV with some sparse setae.
Type II urticating setae: not seen due to abdomen in poor conditions.
Palp (Figs 80–83, 312): globous bulb with a small subtegulum, lacking promi-
nence on tegulum. Embolus: not flattened, lacking keels, 2.66 long in retrolateral
view, about 3.5 times tegulum’s length. Medial portion and tegulum’s margin form an
acute angle in retrolateral view. Proximal part very curved in frontal view; thin distal
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 75

Figure 90. Map showing records of A. minatrix Pocock, 1903, A. hirschii Bullmer, Thierer-Lutz &
Schmidt, 2006, A. lynnae sp. n. and A. caei sp. n.

width, abruptly narrowing distally; basal, middle, and distal width of 0.48, 0.15, 0.04,
respectively. Tegulum: 1.25 long, 0.88 high in retrolateral view. Cymbium subtrian-
gular with subequal lobes, lacking process on retrolateral lobe (Fig. 84).
Tibia I with discrete elevation covered by a cluster of setae in apical portion, on
prolateral side (Figs 85–87).
Color pattern (Fig. 89): same as female, but the dorsal central black area is wider
than in female.
Color pattern ontogeny. Adults maintain the same coloration pattern of imma-
tures. There is no drastic ontogenetic changes in this species.
Distribution. Known only from Venezuela (Fig. 90).
Natural history. Silken tubes are built by specimens inside tree bark and hollow
branches and in the center of large bromeliads, which are in xenophyte bush grassland
clearings of tropical forest (Smith 1992).
76 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 91–92. Avicularia taunayi (Mello-Leitão, 1920), spermathecae variation. 91 Sobradinho, Brasília,
Distrito Federal, Brazil (DZUB 352) 92 Brasíla, Distrito Federal, Brazil (DZUB 1979). Scale bars = 1 mm.

Avicularia taunayi (Mello-Leitão, 1920)


Figs 19, 91–105

Ancylochiros taunayi Mello-Leitão, 1920: 142 (holotype immature male, Brazil, Minas
Gerais, Mariana [20°22'S, 43°25'W], J. P. Fonseca leg., MZUSP 327, examined);
1923: 319, 376, figs 41–44, 160; Roewer 1942: 225; Bonnet 1955: 316.
Anchylochiros taunayi: Petrunkevitch 1939: 291.
Avicularia taunayi: Raven 1985: 149; Bertani 2012: 78, fig. 176; Bertani and Motta
2013: 108, figs 1–13; World Spider Catalog 2016.

Diagnosis. Females of A. taunayi resemble those of A. juruensis and A. variegata stat.


n. by the spermathecae with midwidth expanded; about 1.5 times its basal and apical
portions widths (Fig. 91). They differ from all these species by the spermathecae with
lobes from median to distal portions (Fig. 91). Males of A. taunayi resemble A. avicu-
laria, A. variegata stat. n., A. juruensis, A. rufa, A. purpurea, and A. merianae sp. n. by
tibial apophysis on leg I with well-developed base and grouped spiniform setae distally
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 77

Figures 93–100. Avicularia taunayi (Mello-Leitão, 1920), male (DZUB 1675; except cymbium, DZUB
4542). 93–96 left palpal bulb 93 prolateral 94 retrolateral 95 frontal 96 dorsal 97 left cymbium, dorsal
98–100 right tibial apophysis of leg I (mirrored) 98 prolateral 99 ventral 100 retrolateral. Scale bars = 1 mm.

(Fig. 98). They differ from A. purpurea and A. merianae sp. n. by well-developed pro-
cess on cymbium retrolateral lobe (Fig. 97); from A. avicularia and A. rufa by leg IV
as long as leg I and from A.variegata stat. n. and A. juruensis by developed prominence
78 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 101–104. Avicularia taunayi (Mello-Leitão, 1920), habitus and retreat. 101 immature 102 female
103 male 104 retreat inside trunk of Myrcia tormentosa (Aubl.). Arrow indicating retreat opening. Photos:
101–103 R. Bertani; 104 C. S. Fukushima.

on tegulum (Fig. 95). Additionally, they can be distinguished from all these species by
light brown spots extending from the dorsum to lateral region of abdomen (Fig. 103).
Material examined. Female, Brazil, Minas Gerais, Barão de Cocais [19°56'S,
43°28'W], J. P. Couto col., 5 February 1971, ref. 4336 (IBSP 199); male, Brazil, Bra-
sília D.F. [15°46'S, 47°55'W], Gláucia & Reuber col., 14 May 2002 (DZUB 1675).
Additional material. BRAZIL: Pará: Tucuruí [3°46'S, 49°40'W], 1 male,
Equipe de Resgate de Fauna col., IBA 285 (IBSP 8570); 1 male, 3 immatures, 27
January 1986 (IBSP ref. 48014); Mato Grosso: Barra do Bugres [15°03'S, 57°10'W],
Serra das Araras, 1 juvenile female, C. Strusman col., September 1992 (MCP
2293); Chapada dos Guimarães [15°27'S, 55°45'W], 1 female, M. Acuarensa col.,
November 1963 (AMNH 1.7); Barra do Tapirapé [10°38'S, 50°36'W] (Barro
do Tapirapé [sic]), 2 females, B. Malkin col., 1962 (AMNH 1.5); Goiás: Ipameri
[17°43'S, 48°09'W], 1 male, 1 female, F. R. Alves col., February 1996, ref. 78206
(IBSP 14397); Distrito Federal: Brasília [15°47'S, 47°53'W], 1 female, 15 Novem-
ber 2002, F. Brasil leg. (DZUB 1979); Paranoá [15°43'S, 47°44'W], 1 female, 1
immature, R. Bertani, C. S. Fukushima, R. H. Nagahama, P. C. Motta, P. Jotta,
12 July 2007 (DZUB 4707); Sobradinho [Colorado, Córrego do Urubu, 15°42'S,
47°51'W], 1 female, 1 January 1999, J. Marinho-Filho leg. (DZUB 352); Minas
Gerais: Santa Vitória [18°51'S, 50°07'W], 1 female, M. Rosa col., 28 July 1981
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 79

Figure 105. Map showing records of Avicularia taunayi (Melo-Leitão, 1920), Avicularia purpurea Kirk,
1990 and Avicularia merianae sp. n.

(ZUEC 018); Monte Alegre de Minas [18°52'S, 48°52'W] (Monte Alegre [sic]), 1
male, A. Lourenço col., June 2005, ref. 95480 (IBSP 12780).
Description, color pattern ontogeny and distribution. See Bertani and Motta (2013).
Complementary description. Spermathecae (Figs 91–92): two completely sepa-
rated not-twisted long spermathecae, with walls having lobes from median to distal
portions and accentuaded outwards curvature medially. Midwidth expanded, about
1.5 times its basal and apical portion widths and weakly-sclerotized area shorter than
half the length of well-sclerotized area.
Palp (Fig. 93–96): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 2.62 long in retrolateral
view, about 3.0 times tegulum’s length. Medial portion and tegulum’s margin form an
acute angle in retrolateral view. Proximal part very curved in frontal view; thin distal
width, abruptly narrowing distally; basal, middle, and distal widths 0.42, 0.24, 0.03,
respectively. Tegulum: 1.76 long, 0.96 high in retrolateral view. Cymbium subtrian-
gular with subequal lobes, having well-developed rounded process on retrolateral lobe,
bearing thick setae (Fig. 97).
80 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Tibial apophysis (Figs 98–100): single branch on prolateral leg I, with well-devel-
oped base and grouped spiniform setae distally. Male metatarsus I touches retrolater-
ally tibial apophysis’ setae when folded.
Type II urticating setae: 0.362–0.407 long, 0.009–0.013 wide in females; 0.840–
0.968 long, 0.014–0.020 wide in males.
Color pattern (Figs 102–103): both male and female have long guard-setae on legs
and palps not grizzled.
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi
contrasting with other lighter articles, and abdomen dorsum reddish, with dorsal cen-
tral longitudinal black stripe connected with first two pairs of transversal black stripes
(Fig. 101). When mature, part of the pattern remains (Figs 102–103). Adult females
have abdomen with reddish brown guard-setae homogeneously distributed and black
short body setae with spots of reddish brown short body setae (Fig. 102). Males have
same abdominal pattern as females but reddish brown spots of short body setae are
ill-defined (Fig. 103).
Distribution. Brazil, states of Tocantins, Goiás, Pará, São Paulo, Mato Grosso, west
of Bahia, Minas Gerais and in Brasília (Distrito Federal), in savannah areas (Fig. 105).
Natural history. A small population of A. taunayi was found at Distrito Federal, in
a mountain area that had savannah areas mixed with anthropized areas with houses and
farms (Bertani and Motta 2013). Specimens were found in tree holes within retreats
similar to the ones made by other species of Avicularia (Fig. 104). This is a unique spe-
cies of this genus in the bioma Cerrado (savannah) (Bertani and Motta 2013).

Avicularia juruensis Mello-Leitão, 1923


Figs 15, 19, 76, 106–127, 307

Avicularia juruensis Mello-Leitão, 1923: 321, 377, figs 156, 188 (syntypes 4 females
and 1 male, Brazil, Amazonas, Juruá [4°47'S, 68°38'W], Garbe col., 1902, MZUSP
125A–D, examined; lectotype male (MZUSP 125C) and paralectotype female
(MZUSP 125B), here designated); Roewer 1942: 255; Bücherl 1953: 128, figs
9–11 (misidentification, probably a Tapinauchenius); Bonnet 1955: 831; World
Spider Catalog 2016.
Avicularia urticans Schmidt, 1994: 5, figs 1–2 (holotype female, from Peru, Krasa leg.,
1989, SMF 38035 and spermathecae in microslides, SMF 58243-84 21/11, SMF
58243-84, examined), 1995c: 2, figs 1–2; World Spider Catalog 2016. Syn. n.

Remarks. Avicularia urticans holotype is in poor conditions since the specimen died
during moulting process. Its spermatheca is preserved in slides, but unfortunately it lost
most of its natural shape. Despite this, it was possible to observe that spermatheca have
midwidth expanded, about 1.5 times its basal and apical portion widths. Spermatheca
morphology and overall body coloration match with large specimens found in Peru and
Ecuador. Well-preserved material were examined and despite some differences in color
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 81

Figures 106–108. Avicularia juruensis Mello-Leitão, 1923, spermathecae variation. 106 paralectotype,
Rio Juruá, state of Amazonas, Brazil (MZUSP 125C) 107 paralectotype, Rio Juruá, state of Amazonas,
Brazil (MZUSP 125A) 108 morphotype 3, Pebas, department of Loreto, Peru (MNHN–AR 4902). Scale
bars = 1 mm.

features, A. urticans is indistinguishable from A. juruensis. Thus, we consider A. urticans


Schmidt, 1994 as junior synonym of A. juruensis Mello-Leitão, 1923.
Diagnosis. Females of A. juruensis resemble those of A. variegata stat. n. and A.
taunayi by the spermathecae having midwidth expanded, about 1.5 times its basal
and apical portion widths (Fig. 106). They can be distinguished from A. taunayi by
the spermathecae lacking lobes (Fig. 106). Males of A. juruensis resemble those of A.
avicularia, A. rufa, A. variegata stat. n., A. taunayi, A. purpurea, and A. merianae sp. n.
by the tibial apophysis on leg I with well-developed base and grouped spiniform setae
distally (Fig. 114). They differ from males of all these species except A. variegata stat.
n. by its well-developed prominence on tegulum (Fig. 111). Males and females of A.
juruensis can be distinguished from A. variegata stat. n. by intense purple sheen on
carapace and legs (morphotype 2, Figs 120–121) or intense golden sheen on carapace
and legs (morphotype 1, Fig. 122). They can also be distinguished from A. variegata
stat. n. by the occurrence area: western part of South America, in Brazil, Colombia,
Ecuador, and Peru (Fig. 76).
Additional material. COLOMBIA: Vaupés: Vaupés [0°5'N, 70°48'W], low Río
Apaporis, Lago Toraima, Estación Biológica Caparu, 200 m asl, 1 male, Col. Jaime
Pinzól (AP3-5) (ICN–Ar-2006); Putumayo: Puerto Leguízamo, Parque Nacional Na-
82 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 109–116. Avicularia juruensis Mello-Leitão, 1923, male lectotype (MZUSP 125B). 109–112 left
palpal bulb 109 prolateral 110 retrolateral 111 frontal 112 dorsal 113 left cymbium, dorsal 114–116 right
tibial apophysis of leg I (mirrored) 114 prolateral 115 ventral 116 retrolateral. Scale bars = 1 mm.

tural La Paya [0°28'N, 75°49'W], Mamansoyá (Mamangaya [sic]), 1 male, 21 Septem-


ber 2001, D. Campos col. (ICN–Ar-1972); Amazonas: Letícia, km 2 Via Taparaca
(trapaca [sic]), (4°12'19.25'S, 69°55'58.07"W), 100 m asl, 1 male, Col. Est. Sist. Anim.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 83

I-2002, 25 April 2002 (13001) (ICN–Ar-1970); [4°12'S, 69°56'W], km 11, Carretera


a Tarapaca, bosque en interior de hija enrollada con casulo de seda a 50 m del suelo, co-
leta manual, 100 m asl, 1 immature, E. Flórez col., 27 October 1997 (ICN–Ar-1978);
km 10, via Terapacos, Finca La Arerosa, 95 m asl, 1 immature, Col. Est. Sist. Animal
II.03, 06 November 2003 (ICN–Ar-2369); La Pedrera, Resguardo Indígena Curaril–
Los Ingleses, colectada em el interior de una vivenda, em horas nocturnas, 1 male, Z.
Cordero col., 24 April 2004 (ICN–Ar-6819); cerca de Letícia [4°12'S, 69°56'W], 100
m asl, D. Campos col., August 1997 (ICN–Ar-5002); BRAZIL: Amazonas: between
Benjamin Constant [4°22'S, 70°01'W] and São Paulo de Olivença [3°22'S, 68°52'W],
1 male, P. L. Conti col., 3 August 1972, ref. 9971 (IBSP 2829); Alto Solimões, 2
males, 31 August 1972 (IBSP 3389); Igarapé Belém, near confluence with Rio Soli-
mões [3°05'S, 60°08'W], 1 male, B. Malkin col., 5–30 April 1966 (AMNH 1.29); Rio
Negro [3°09'S, 59°57'W], 1 female, J. Coffey col., September 1994 (AMNH RW31);
Carauari, left margin of Rio Juruá, Comunidade Esperança, RESEX Médio Juruá
(05°05'31"S, 67°10'03"W), 1 male, F. F. Xavier Filho & A. L. Henriques col., 27
June to 16 July 2005 (INPA 4886); Pará: Breves [1°40'S, 50°28'W], margem W, Área
2, 1 male, J. Dias col., 2 February 1988 (MPEG 5398); Acre: Cruzeiro do Sul [7°37'S,
72°40'W], 1 female, S. Albuquerque (by photo); ECUADOR: 1 female, M. Baumgar-
ten leg., 1994 (IBSP 12887); Napo: Parque Nacional Yasuní, Catholica Field Station
(0°40'54"S, 76°23'9.33"W), 2 males, A. I. Ognato col., 15 July 1996 (CAS 6, CAS 4);
Puerto Napo, 20 km east, Aliñahuí (1°0'S, 77°25'O), 450 m, 1 immature male, V. D.
& B. Roth col., January 1994 (CAS 11A); 1 immature male, A. urticans det. R. West
in August 94, V. D. & B. Roth col. (CAS 8); 1 male, A. urticans det. R. West in August
94, V. D. & B. Roth col., June 1994 (CAS 5); Puerto Napo, 25 km East, Selva Aliña-
huí [1°0'S, 77°25'W], 450 m, 2 immature males, E. Ross col., January–February 1991;
Avicularia sp. near juruensis det. J. Ledford 1997 (CAS); Pastaza: Tiguino [1°12'S,
7°51'W], 1 female, W. Lamar col., September 1990 (AMNH RW53); Morona-Santia-
go: Los Tayos, (3°05'S, 78°02'W), 1 female, in grass area near mil. camp, in afternoon,
5 July 1976 (IBSP 12884); same locality, in hole in tree trunk live but part rotten c.1
m from base, 1 male (IBSP 12888); PERU: spermathecae in microslides, no further
information (SMF 58257-84); Rio Bombo, alto Tapiche, 1 female, 2 immatures, H.
Bassler col, January 1928 (AMNH Pe55); R. Marañon [6°24'S, 76°05'W], 1 female,
Bristol [col.?], October 1927 (AMNH Pe5); Marañón (Marauon [sic]),1 female,1 im-
mature male, Bristol [col.?], October 1927 (AMNH Pe96); no data, probably Loreto,
INRENA confiscation, 1 female (UA 088/2004); Peruvian jungle, confiscation, 1 male
(UA 098/2004); Loreto: no further information, 1 female, Collection Bluntschili-Pe-
yes, 1912 (AMNH Pe115); Cashiboya [7°39'S, 74°55'W], 1 female, February 1927
(AMNH Pe111); Estirón [4°07'S, 70°43'W], Rio Ampiacu, 3 females, 1 male, 2 juve-
niles, B. Malkin col., 15–22 May 1966 (AMNH Pe58); Iquitos [3°44'S, 73°15'W], 1
female, J. Huff col., November 1995 (AMNH RW51); Iquitos, Rio Momon, Amazon
Camp (3°41'13.00"S, 73°16'48'00W), 2 females, 1 male, R. C. West col., 9 Novem-
ber 1993 (AMNH RW35, AMNH RW36, AMNH RW37, respectively); 1 male, T.
Mason col., March 1993 (AMNH RW39); 2 females, R. C. West col., 6 November
84 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 117–122. Avicularia juruensis Mello-Leitão, 1923, habitus. 117–118 morphotype 2 117 im-
mature 118 juvenile 119 female with yellowish leg ring, department of Napo, Ecuador 120 female with
whitish leg ring, department of Loreto, Peru 121 male, department of Loreto, Peru 122 morphotype 1,
female, Peru. Photos: 117 Tanya Stewart; 118 W. Lamar; 119–122 R. C. West.

1993 (AMNH RW40, AMNH RW41); 1 male, E. Cooper col., May 1993 (AMNH
RW43); Pebas [3°19'S, 71°51'W], 1 female, De Mathan col. (MNHN–AR 4902); Rio
Tahuayo [4°53'S, 73°08'W], 1 male, W. Lamar col., September 1990, found para-
site on forest floor (AMNH RW46); San Martín: Tarapoto [06°07'S, 75°57'W], 1
female, (UA 668/2005); Valley of Cainarachi, 40 miles east of Tarapoto, 1 female,
700–1500 m a.s.l., December 1925 (AMNH Pe54); Ucayali: Pampa Hermosa, Rio
Ucayali [7°34'S, 74°19'W], 1 male, 1 female, January or June 1927 (AMNH Pe34);
(Pompa Hermosa [sic]; Ucayoli [sic]), 8 females, February 1927 (AMNH Pe116); Rio
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 85

Utiquinea [8°131'S, 74°32'W] (Upper Utoguinia [sic]), La frontera, 1 juvenile female,


H. Bassler col., 1928 (AMNH Pe113); (Rio Utuguinea [sic]), Peru–Brazil frontier,
1 immature, August 1927 (AMNH Pe109); Crenze Zwischen Peru und Brazil, sud
Crenze, Beim Uberer Utoquinia, 1 immature male, “Scolopenda”, 10 February 1928
(AMNH 1.10); Río Ucayali [7°34'S, 74°19'W], Suhuaya + Rean Rean [?], 2 juvenile
females, 1 immature male, 12–16 December 1926 (AMNH Pe122).
Male. Description. MZUSP 125C. Carapace: 15.23 long, 15.31 wide, 4.67 high.
Chelicera: 5.60 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 15.92, 8.47,
12.61, 11.81, 7.31, 56.12. II: lost. III: lost. IV: 16.63, 6.87, 14.78, 15.72, 6.16, 60.16.
Palp: 9.11, 6.03, 8.09, –, 3.51, 26.74. Midwidths: femora I–IV= 3.21, –, –, 3.08, palp=
2.23; patellae I–IV= 2.74, –, –, 2.60, palp= 1.91; tibiae I–IV= 2.34, –, –, 2.43, palp=
2.18; metatarsi I–IV= 1.54, –, –, 1.70; tarsi I–IV= 1.67, –, –, 1.93, palp= 2.06. Abdo-
men: 16.66 long, 12.25 wide. Spinnerets: PMS, 1.20 long, 0.54 wide, 0.23 apart; PLS,
1.83 basal, 1.79 middle, 2.96 distal; midwidths 1.41, 1.41, 0.93, respectively.
Carapace: as long as wide; cephalic region slightly raised, thoracic striae conspicuous.
Fovea: deep, slightly recurve, 2.27 wide.
Eyes: ocular tubercle 1.05 high, 2.19 long, 3.22 wide. Clypeus 0.56. Anterior eye
row procurve. Posterior slightly recurve. Eye size and interdistances: AME 0.69, ALE
0.77, PME 0.25, PLE 0.65, AME–AME 0.59, AME–ALE 0.51, AME–PME 0.22,
ALE–ALE 2.14, ALE–PME 0.95, PME–PME 1.97, PME–PLE 0.21, PLE–PLE 2.47,
ALE–PLE 0.54, AME–PLE 0.56.
Maxilla: length to width: 2.31. Cuspules: 100–200 spread over ventral inner heel.
Labium: 2.51 long, 2.75 wide, with 87 cuspules spaced by one diameter from each
other on anterior third. Labio-sternal groove shallow, flat, with no visible sigilla.
Sternum: 7.18 long, 6.71 wide. Sigilla: not evident.
Legs: Formula: IV=I – –. Length leg IV to leg I: 1.07. Clavate trichobothria: 2/3
distal on tarsi I, IV. Scopulae: Tarsi I and IV fully scopulate. IV lacking sparse setae.
Metatarsi I fully scopulate, II–III ?; IV on distal 1/3. IV divided by a row of setae.
Type II urticating setae: 0.94–1.01 long, 0.017–0.021 wide.
Palp (Figs 109–112): globous bulb with small subtegulum and well-developed
prominence on tegulum. Embolus: not flattened, without keels, 5.02 long in retro-
lateral view, about 3.0 times tegulum’s length. Medial portion and tegulum’s margin
form an acute angle in retrolateral view. Proximal part very curved in frontal view;
thin distal width, abruptly narrowing distally; basal, middle, and distal width of 0.89,
0.17, 0.03, respectively. Tegulum: 1.62 long, 2.72 high in retrolateral view. Cymbium
subtriangular with subequal lobes, and well-developed rounded process on retrolateral
lobe, bearing thick setae (Figs 113, 307).
Tibial apophysis (Figs 114–116): single branch on prolateral leg I, with well-devel-
oped base and grouped spiniform setae distally. Male metatarsus I touches retrolater-
ally tibial apophysis’ setae when folded.
Color pattern: carapace brown with golden short body setae. Carapace border with
long setae the same color as dorsal carapace short body setae. Coxae, labium, sternum
and maxillae brown, slightly darker than ventral femora. Legs and palps with golden
86 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

brown short body setae and brown long dark guard-setae. Leg rings on distal femora,
tibiae and metatarsi whitish. Abdomen dorsum with reddish brown guard-setae and
black short body setae. Ventral abdomen brown.
Female. Redescription. MZUSP 125B. Carapace: 19.26 long, 16.94 wide, 5.22
high. Chelicera: 8.77 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I:
15.30, 9.13, 11.53, 9.92, 6.66, 52.54. II: 14.21, 8.26, 10.39, 10.37, 6.25, 49.48. III:
12.99, 7.49, 9.86, 9.68, 6.43, 46.45. IV: 15.75, 8.48, 13.59, 13.19, 6.62, 57.63. Palp:
10.61, 6.74, 7.07, –, 8.34, 32.76. Midwidths: femora I–IV= 3.64, 3.71, 3.84, 3.45,
palp= 2.84; patellae I–IV= 3.55, 3.68, 3.69, 3.62, palp= 2.90; tibiae I–IV= 2.97, 3.13,
3.00, 2.72, palp= 2.88; metatarsi I–IV= 2.34, 2.67, 2.21, 2.20; tarsi I–IV= 2.89, 2.44,
2.80, 2.62, palp= 2.78. Abdomen: 24.29 long, 16.59 wide. Spinnerets: PMS, 2.32
long, 1.54 wide, 0.1 apart; PLS, 3.43 basal, 2.14 middle, 2.86 distal; widths 1.82,
1.45, 1.28, respectively.
As in male, except:
Carapace: 1.14 times longer than wide.
Fovea: 2.19 wide.
Eyes: eye tubercle 1.15 high, 2.76 long, 3.56 wide. Clypeus 0.48. Eye size and
interdistances: AME 0.78, ALE 0.80, PME 0.34, PLE 0.75, AME–AME 0.64, AME–
ALE 0.56, AME–PME 0.29, ALE–ALE 2.53, ALE–PME 0.91, PME–PME 2.13,
PME–PLE 0.14, PLE–PLE 2.95 ALE–PLE 0.66, AME–PLE 0.66.
Maxilla: length to width: 1.70. Cuspules: 100–200 spread over ventral inner heel.
Labium: 2.00 long, 2.71 wide, with 88 cuspules spaced by one diameter on anterior third.
Chelicera: basal segment with 10 teeth. Sternum: 8.69 long, 7.79 wide.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.10. Clavate trichobothria: 2/3
distal on tarsi I–IV. Scopulae: Tarsi I and IV fully scopulate. IV lacking sparse setae.
Metatarsi I–II fully scopulate, III on distal 2/3; IV on distal 1/3. III divided by a bald
area, IV divided by a row of setae.
Type II urticating setae: 0.54–0.66 long, 0.014–0.018 wide (measured MZUSP 125A).
Spermathecae (Fig. 106): two completely separated, not-twisted very long sper-
mathecae, with walls lacking projections or lobes and accentuated outwards curvature
medially. Midwidth expanded, about 1.5 times its basal and apical portion widths and
weakly-sclerotized area shorter than half the length of well-sclerotized area.
Color pattern: dorsal abdomen with long brown guard-setae grouped on lateral
and dorsal anterior areas, and dark short body setae.
Variation. We found two different morphotypes. Morphotype 1 is found near Rio
Juruá, state of Amazonas and Acre, Brazil, and in some areas of Peru (Fig. 127). Fe-
males have discrete grizzled setae in palp and legs, whitish leg rings and carapace, legs
and palps with short body setae with golden and pink sheen, and abdomen with long
light brown guard-setae homogeneously distributed over dark brown body short setae
(Fig. 124). Males examined are not in good conditions but they do not seem to have
white tipped setae on dorsal abdomen. Despite having yellowish leg rings, a female
from Iquitos, Peru (Fig. 122) is considered as morphotype 1 since it has discrete griz-
zled guard-setae and short body setae with intense golden sheen on legs and carapace.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 87

Figures 123–126. Avicularia juruensis Mello-Leitão, 1923, retreat, habitus and behavior. 123 morphotype
2, retreat 124 morphotype 1, female from Cruzeiro do Sul, state of Acre, Brazil, habitus 125 female swim-
ming on Rio Momon, Peru 126 female swimming on Rio Marañon, Peru. Photos: 123, 125–126 R. C.
West; 124 S. Albuquerque.

Morphotype 2 is the most common morphotype found in the material examined and it
was formerly known as Avicularia urticans. Both sexes have very grizzled setae on palps
and legs, and carapace, legs and palps with dark short body setae with intense purple
sheen, and yellowish leg rings (Figs 119–121). Females have dorsal abdomen with long
reddish brown guard-setae grouped on lateral and dorsal anterior areas, and dark short
body setae (Fig. 120). Males have white tipped setae homogenously distributed on dor-
sal abdomen (Fig. 121). They can be found mostly in Ecuador and Peru, but a specimen
with the same characteristic pattern was found in Breves, state of Pará, Brazil.
Remarks. For many years the name A. juruensis has been applied to specimens that
have vivid yellow leg rings and grizzled setae on legs and palps, that are commonly found
in the states of Mato Grosso and Rondônia, Brazil. They have spermathecae with mid-
width not expanded, developed prominence on palpal bulb and leg IV longer than leg I.
However, A. juruenesis syntypes have whitish leg rings and lack setae with conspicuous
88 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 127. Map showing records of Avicularia juruensis Mello-Leitão, 1923 morphotypes.

whitish apex on legs, spermathecae with midwidth expanded, palpal bulb with well-de-
veloped prominence, and leg IV as long as leg I. The characters found in these specimens
formerly known as A. juruensis match, in fact, with those of A. rufa. Thus, we conclude
the name A. juruensis is being mistakenly applied to specimens of A. rufa.
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi
contrasting with other lighter articles and abdomen dorsum reddish, with dorsal cen-
tral longitudinal black stripe disconnected from transversal black stripes (Fig. 117).
When mature, both males and females lose this pattern.
Distribution. Brazil (states of Amazonas, Acre and Pará), Colombia, Ecuador and
Peru (Fig. 76).
Natural history. Silken retreats of A. juruensis are similar to other Avicularia spe-
cies (Fig. 123). An adult female A. juruensis was reported feeding on a greater sac-
winged bat (Saccopteryx bilineata (Temminck, 1838)) on the side of a palm tree near
Rio Yarapa, Peru (Nyffeler and Knörnschild 2013). Another interesting behavior was
the ability to swim through large rivers such as Marañon and Momon rivers in Peru
(Figs 125–126) (R. C. West, pers. comm.), which can explain why rivers do not seem
to act as natural barriers to Avicularia.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 89

Avicularia rufa Schiapelli & Gerschman, 1945


Figs 19, 50, 128–153, 300, 302

Avicularia rufa Schiapelli & Gerschman, 1945: 190, pls. XV–XVI, XXV (holotype
female, Brazil, Mato Grosso, Alto Ji Paraná [10°52'N, 61°55'W] (Gy Paraná [sic]),
Dr. Vellard col., December 1938, MACN–Ar 845, examined,); Brignoli 1983:
134; World Spider Catalog 2016.
Avicularia juruensis (misidentification): West et al. 2008: 37, 52–54; Ayroza et al.
2012: 2; Bertani 2012: 5, 79–80.

Diagnosis. males and females of A. rufa resemble A. avicularia and female of A. hirschii
by the leg IV longer than leg I. Females of A. rufa can be distinguished from those of
A. hirschii by the non-twisted spermathecae. Females and males of A. rufa differ from
those of A. avicularia by having vivid yellow rings on distal femora, tibiae and meta-
tarsi combined with legs and palps with very grizzled guard-setae (Fig. 145) (from all
morphotypes of A. avicularia except 6 and 7, but these have intense green metallic
sheen on carapace and legs, absent in A. rufa).
Material examined. 1 female, Brazil, Mato Grosso, between Vale de São Domin-
gos [15°17'S, 59°03'W] (Vale de São Lourenço [sic]) and Pontes & Lacerda [15°12'S,
59°19'W], Operação Coatá, I. Knysak col., 14 December 2002, RGI 1963 (IBSP
10264) and 2 males and 1 immature female, Brazil, Mato Grosso, Operação Coatá
col., 1–20 July 2002 (MCP 13592).
Additional material. BRAZIL: no further data: 1 female, 1 immature (IBSP
2157); 1 female (IBSP 3170); 7 immatures, G. Raidar col., 14 May 1951 (IBSP 2620);
1 female (IBSP 4368); 2 males (MZUSP 22501); 2 females (IBSP 1393); Amapá:
Serra do Navio [1°38'N, 52°12'W], 2 immature males (IBSP 3703); Amazonas: Ro-
dovia Transamazônica, km 530, 1 male, 23 October 1975 (IBSP 7895); Transamazôni-
ca Humaitá–Porto Velho, 1 immature, Dr. A. H. Hoge col., ref. 4102, July 1972
(IBSP 4102); Barcelos [0°57'S, 62°55'W], Parque Nacional do Jaú, Lago do Miratuca,
1 female, R. Andriazze, W. L. S. Costa & L. Aquino col., 14 July 1993 (INPA 4884);
Boca do Jacaré [5°45'S, 63°41'W], Rio Solimões, 1 female (IBSP 3110); Boca do Tefé
[3°19'S, 64°43'W], Rio Solimões, 1 female (IBSP 3104); 2 females, 1 immature male
(IBSP 3099); 1 female, 3 immatures (IBSP ref.189); 1 male (IBSP 3101); 1 female
(IBSP 3103); 1 immature male (IBSP 3102); 2 immatures (IBSP 3100); Boca do Tefé
[3°19'S, 64°43'W], Rio Solimões, 5 females, 1 male, 2 immatures males, 5 immatures
(IBSP 3104, IBSP 3099, IBSP ref. 189, IBSP 3103, IBSP 3101, IBSP 3102, IBSP
3100); Cucuí [1°11'N, 66°49'W], 1 juvenile female, P. Serveira col., November 1975,
ref.17.595 (IBSP 2569); Humaitá [7°30'S, 63°01'W], 1 female, in silk retreat in a
palm tree (IBSP 7932); 1 male, 4 females, L. R. Latorre col., November 1981, ref.
40348 (IBSP 4686, IBSP 4687); pottery and pineapple crop, 1 male, 1 juvenile fe-
male, 1 immature male, 2 immatures, Dr. A. R. Hoge col., July 1972, IB 24.8.72
(IBSP 4099); Transamazônica, km 86, 1 male, 23 July 1972 (IBSP 4100); Puruzinho,
Rio Madeira [6°52'S, 62°05'W], 1 female, EPA col., 4 December 1975 (MZUSP
90 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 128–130. Avicularia rufa Schiapelli & Gerschman, 1945, spermathecae variation. 128 between vale
de São Domingos and Pontes & Lacerda, state of Mato Grosso, Brazil (IBSP 10264) 129 Rio Negro, state
of Amazonas, Brazil (AMNH RW29) 130 Tomé-Assu, state of Pará, Brazil (IBSP 3105). Scale bars = 1 mm.

27610); Rio Negro [3°09'S, 59°57'W], 2 males, 1 female, A. McKee col., 19 April
1989 (AMNH RW67, AMNH RW25, AMNH RW27); 2 females, A. McKee col., 29
June 1988 (AMNH RW29, AMNH RW28); 1 immature male, R. Mascarino col., 18
September 1996 (AMNH RW30); Tapera [0°55'N, 67°26'W], Rio Negro, 1 imma-
ture male, EPA col., 4 November 1972, P. E. Vanzolini col., 722574 (MZUSP un-
numbered); Tefé [3°20'S, 64°43'W], 1 male, 1 female, De Mathan col. (MNHN–AR
4903); 3 females, 1 immature (IBSP 193); 2 immature females, 2 immatures (IBSP
184); Pará: Belém [1°27'S, 48°30'W], IAN, 1 immature male (IBSP 3123); Belterra
[3°07'S, 55°03'W], 1 female, 15 March 2003 (MPEG 4800); Itaituba [4°16'S,
55°59'W], Rio Maropa, 1 male, W. G. Ravem col., ref. 41844 (IBSP 8576); 1 female,
dead 25 April 1990 (IBSP ref. 41.844); Óbidos [1°54'S, 55°31'W], Igarapé Jaramaca-
ru, Campos do Ariramba, 1 immature male, EPA col., 24 January 1960 (MZUSP
14873); São Félix do Xingu [6°37'S, 51°58'W], 1 male, 1 female, G. Whitaker col.,
July 1980, ref. 28300 (IBSP 4562); Tomé-Assu [2°25'S, 48°09'W] (Thomé Assu [sic]),
Rio Acará Mirim, 1 female (IBSP 3105); Tucuruí [3°46'S, 49°40'W], 1 female, Equi-
pe de Resgate de Fauna col., Ref. IBA 218 (IBSP 8574); Vila Bravo, 1 female, Equipe
de Resgate de Fauna col., ref. B410-12C (IBSP 8577); Acre: Rio Branco [9°58'S,
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 91

67°48'W], Catuaba, 1 immature male, A03, April 1996 (IBSP unnumbered); Mare-
chal Thaumaturgo, Terra Indígena Jaminawa-Arara, Rio Bagé, Aldeia Buritizal,
[8°56'S, 72°47'W], 1 immature, 02 September 2000, 9AM, Coleção de Simone La-
deia Andrade (MZUSP 70947); Rondônia: Candeias do Jamari, UHE Samuel [8°45'S,
63°27'W], 1 female, W. Bokerman col., June 1989 (AMNH RW24); Jaru [10°26'S,
62°28'W], Santa Cruz da Serra, 2 females, Expedição Polo Noroeste col., 23–27 De-
cember 1964 (MZUSP 11065); Monte Negro [10°14'S, 63°17'W], 1 immature male,
1 female, 1 juvenile female, R. Bertani & P. I. da Silva Junior col., 22 July to 03 August
2002 (IBSP 10042, IBSP 10218, IBSP 10945, respectively); Núcleo Avançado de
Pesquisa de Monte Negro, 1 male, L. M. A. Camargo col., June 2002, ref. 89431
(IBSP 1028); 1 juvenile female, L. M. A. Camargo col., 06 April 2002, ref. 89430
(IBSP 10205); Porto Velho [8°43'S, 63°53'W], 1 male, G. Insley col., 31 August
1973, ref. 12732 (IBSP 2454); 1 male, G. Insley col., 9 October 1973, ref. 12922
(IBSP 2483); Santa Luzia d’Oeste [11°54'S, 61°46'W], 1 immature female, R. Mote-
rani col., July 1999 (IBSP 9538); Mato Grosso: Alto Xingu [7°16'S, 52°36'W], 1 male,
H. Schulze col., 21 October 1964 (IBSP 3693); Barra do Bugres [15°03'S, 57°10'W],
Cia Paulista de Ferro Ligas, 1 immature, 26 July 1971, ref. 7624 (IBSP 4103); 1
immature, A. Cerrutti col., November 1984 (MNRJ 12942); Chapada dos Guimarães
[15°26'S, 55°44'W], UHE Rio Manso, 1 female, Faunal Rescue Team of Furnas col.,
2000 (IBSP 9087); 1 female, H. N. da Cunha col., December 1976 (IBSP 13805); Rio
Kuluene (Koluene [sic]) [13°22'S, 52°59'W], 1 immature male, J. M. C. Carvalho col.,
1947 (MNRJ 13.773); Lagoa Ipavu [12°7'S, 53°26'W] (Lagoa Ipavi [sic]), 1 immatu-
re male, 1 juvenile female, P. Vanzolini col., 1965 (MZUSP E3474 C3918); Nova
Mutum [13°49'S, 56°05'W], 1 female, R. K. Ribeiro col., 06–15 July 2001 (IBSP
10929); Parque Nacional Xingu [11°59'S, 54°00'W], 1 male, C. Andreatta col., Mar-
ch 1968 (IBSP 39A); 2 immatures, Alvarenga & Werner col., November 1961
(AMNH 1.19); São José do Rio Claro [13°25'S, 56°42'W], 2 males, M. Caleffo col.,
June 1997 (IBSP 10309); Sinop [11°52'S, 55°29'W], 1 immature, 4 March 1977, ref.
20109 (IBSP 4455); Vila Bela da Santissima Trindade [15°00'S, 59°57'W] (Villebela
[sic]), Rio Serra Azul, 1 male, Dr. Hoge col., 27 June 1962 (IBSP 3614); Maloca Feia,
1 female, Dr. Hoge, Pedro & Joaquim col., 27 June 1962 (IBSP 3615); Between Vale
de São Domingos [15°17'S, 59°03'W] ([sic] Vale de São Lourenço) and Pontes &
Lacerda [15°14'S, 59°19'W], UHE Guaporé, Operação Coatá, I. Knysak col.: 1 juve-
nile female, 17 May 2002, RGI 074 (IBSP 10274); 1 juvenile female, Operação Coatá,
I. Knysak col., 18 May 2002, RGI 086 (IBSP 10253); 1 juvenile female, Operação
Coatá, I. Knysak col., 14 September 2002, RGI 1871 (IBSP 10262); 2 females, Oper-
ação Coatá, I. Knysak col.,1 October 2002, RGI 2527, RGI 2528 (IBSP 10241,
IBSP10242, respectively); 2 females, 5 juvenile females, 3 immature males, Operação
Coatá, I. Knysak col., 03 October 2002, RGI 2299, RGI 2516, RGI 2313, RGI 2518,
RGI 2300, RGI 2314, RGI 2517, RGI 2315, RGI 2316, RGI 2302 (IBSP 10232,
IBSP 10224, IBSP 10246, IBSP 10226, IBSP 10233, IBSP 10247, IBSP 10225, IBSP
10248, IBSP 10249, IBSP 10235, respectively); 1 female, Operação Coatá, I. Knysak
col., 05 October 2002, RGI 1871 (IBSP 10220); 1 immature female, Operação Coatá,
92 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 131–138. Avicularia rufa Schiapelli & Gerschman, 1945, male (MCP 13592). 131–134 right
palpal bulb (mirrored) 131 prolateral 132 retrolateral 133 frontal 134 dorsal 135 right cymbium, dorsal
(mirrored) 136–138 right tibial apophysis of leg I (mirrored) 136 prolateral 137 ventral 138 retrolateral.
Scale bars = 1 mm.

I. Knysak col., 7 October 2002, RGI 1871 (IBSP 10230); 1 female, 2 juvenile females,
Operação Coatá, I. Knysak col., 08 October 2002, RGI 1773, RGI 1774, RGI 1775
(IBSP 10275, IBSP 10279, IBSP 10278, respectively); 1 immature, S2, Operação
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 93

Coatá, I. Knysak col., 8 October 2002 (IBSP unnumbered); 2 females, 1 juvenile fe-
male, 1 immature male, Operação Coatá, I. Knysak col., 14 October 2002, RGI-, RGI
1962, RGI 1964, RGI 1966 (IBSP 10256, IBSP 10263, IBSP 10255, IBSP 10257,
respectively); 1 female, Operação Coatá, I. Knysak col., 01 December 2002, RGI 2526
(IBSP 10240); 1 immature male, 03 December 2002, RGI 2301 (IBSP 10234);
U.H.E. Guaporé [15°16'S, 59°04'W], Operação Coatá, 2 females, 1 juvenile female,
4–14 October 2002 (MCP 13595); 1 immature male, 2 immatures, Operação Coatá,
14 October 2002 (MCP 13598); 4 immatures, Operação Coatá, 10 June 2002 (MCP
13593); 3 females,1 immature female, Operação Coatá, 14 February 2002 (MCP
13596, MCP 13599); 1 immature, Operação Coatá, 01–07 September 2002 (MCP
13555); 2 immature males, 1 female, Operação Coatá, 4–14 December 2002 (MCP
13594); 1 female, 1 immature, Operação Coatá, 14 December 2002 (MCP 13597);
Xingu, Suia-Missu [11°39'S, 51°25'W] (Tuiã Missu [sic]), 5 immatures, Whytaker
col., 05 February 1980, ref. 27055 (IBSP 4523); ECUADOR: Napo: Puerto Napo
[1°01'S, 77°43'W], 25 km east, Selva Aliñahuí, 450 m, 1 juvenile female, E. S. Ross
col., March 1992 (CAS 9); PERU: Madre de Dios: Iñapari [10°57'S, 69°34'W], Esper-
anza, 2 females, 4 immature males, 245 m a.s.l., average temperature 32.4°C, J. Mo-
rant Araque col., 22 September 1992 (MUSM); Junin: Rio Tambo [11°10'S, 74°14'W],
Shevaja, in short palm trees, 3 females, P. Hocking col., 12 October 2011 (MUSM);
300 m a.s.l., in palm trees, 1 female, P. Hocking col., 23 October 2010 (MUSM); 1
female, P. Hocking col., October 2010 (MUSM–ENTO 504260); 1 female, H.
Bassler col., February 1928 (AMNH Pe121); Cusco: Cashiriari (11°52'S, 72°39'W), 1
female, S. Córdova col., 25 November 1997 (MUSM–ENTO); Cusco [department or
city?], Timpia Rio Urubamba [12°28'S, 72°29'W], 1 female, P. Hocking & L. Cam-
pos col., March 2009 (MUSM–ENTO 500676); BOLIVIA: Santa Cruz: San Ignácio
de Velasco [16°22'S, 60°55'W], 1 female (IBSP 3552).
Female. Redescription. IBSP 10264. Carapace: 17.53 long, 17.69 wide, 4.26 high.
Chelicera: 7.70 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 15.59, 8.80,
11.42, 9.78, 6.41, 51.63. II: 14.52, 8.76, 10.23, 9.58, 6.86, 49.95. III: 13.61, 7.99,
10.21, 10.37, 6.14, 48.32. IV: 17.59, 8.68, 15.93, 14.81, 6.67, 63.68. Palp: 10.20, 6.51,
7.18, –, 7.48. Midwidths: femora I–IV= 3.69, 3.18, 3.42, 3.68, palp= 2.85; patellae I–
IV= 3.44, 3.41, 3.36, 3.78, palp= 2.96; tibiae I–IV= 2.89, 3.18, 3.02, 3.24, palp= 2.80;
metatarsi I–IV= 2.61, 2.51, 2.82, 2.73; tarsi I–IV= 2.89, 2.93, 2.95, 3.05, palp= 2.95.
Abdomen: 24.70 long, 17.11 wide. Spinnerets: PMS, 2.73 long, 1.44 wide, 0.4 apart;
PLS, 3.31 basal, 1.56 middle, 3.78 distal; midwidths 2.44, 2.06, 1.61, respectively.
Carapace: 0.99 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Fovea: deep, slightly recurved, 3.52 wide.
Eyes: eye tubercle 1.61 high, 2.71 long, 3.39 wide. Clypeus 0.18. Anteriore eye
row procurve. Posterior eye row slightly recurve. Eye sizes and interdistances: AME
0.78, ALE 0.87, PME 0.32, PLE 0.76, AME–AME 0.56, AME–ALE 0.42, AME–
PME 0.22, ALE–ALE 2.13, ALE–PME 0.56, PME–PME 2.15, PME–PLE 0.18,
PLE–PLE 2.90, ALE–PLE 0.28, AME–PLE 0.64.
94 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 139–141. SEM microphotographs of cymbium process in male of Avicularia rufa. 139 dorsal
view 140 process, detail, dorsal 141 well-developed setae of process.

Maxilla: length to width: 1.69 longer than wide. Cuspules: 100–200 spread over
ventral inner heel. Labium: 2.02 long, 3.41 wide, with 133 cuspules spaced by one
diameter from each other on anterior third. Labio-sternal groove shallow, flattened,
without evident sigilla.
Chelicera: basal segment with 10 teeth in a row and some small teeth on promar-
gin. Sternum: 7.75 long, 7.45 wide. Sigilla: rounded posterior, less than one diameter
from margin; other sigilla not evident.
Leg: Formula: IV I II III. Length leg IV to leg I: 1.23. Clavate trichobothria: distal
2/3 tarsi I–IV. Scopula: Tarsi I–IV fully scopulate. Metatarsi I fully scopulate; II 4/5;
III 1/3 and IV 1/2 distal scopulate. IV divided by a row of setae.
Type II urticating setae: 0.50–0.58 long, 0.013–0.019 wide (measured from
IBSP 10233).
Spermathecae (Fig. 128): two completely separated, not-twisted long spermathe-
cae, with walls lacking projections or lobes and accentuated outwards curvature medi-
ally. Midwidth as wide as its base width and weakly-sclerotized area shorter than half
the length of well-sclerotized area.
Color pattern (Fig. 145): carapace brown with golden short body setae with sligthtly
purple sheen. Carapace border with long setae the same color as dorsal carapace short body
setae. Coxae, labium, sternum and maxillae darker than ventral femora. Legs and palps
with brown short body setae having pink sheen and brown long guard-setae with darker
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 95

base and contrasting whitish apex. Leg rings on distal femora, tibiae and and metatarsi
vivid yellow (Fig. 300). Abdomen with long greyish brown guard-setae grouped on lateral
and dorsal anterior areas and dark short body setae (Fig. 302). Abdomen venter brown.
Male. Description. MCP 13592 (larger specimen). Carapace: 14.15 long, 14.45
wide, 3.3 high. Chelicera: 6.6 long. Legs (femur, patella, tibia, metatarsus, tarsus, total):
I: 13.56, 7.50, 11.67, 11.46, 7.55, 51.74. II: 13.02, 7.16, 11.14, 11.15, 6.45, 48.92. III:
13.44, 6.45, 10.55, 11.46, 6.66, 48.59. IV: 18.14, 7.35, 14.16, 13.55, 8.15, 61.65. Palp:
3.62, 3.45, 8.51, –, 3.54, 19.12. Midwidths: femora I–IV= 2.62, 3.07, 2.65, 5.56; palp=
2.16; patellae I–IV= 3.50, 3.15, 2.56, 2.66, palp= 2.42; tibiae I–IV= 2.66, 2.25, 2.26, 2.46,
palp= 2.15; metatarsi I–IV= 1.67, 1.75, 1.70, 1.13; tarsi I–IV= 2.12, 2.15, 2.26, 2.33.
palp= 2.16. Abdomen: 19.07 long, 13.05 wide. Spinnerets: PMS, 1.90 long, 0.98 wide, 0.2
apart; PLS, 2.70 basal, 1.40 middle, 2.95 distal; midwidths 1.50, 1.40, 1.16 respectively.
As in female, except:
Carapace: 0.98 times longer than wide.
Fovea: 2.55 wide.
Eyes: eye tubercle 1.3 high, 2.56 long, 3.00 wide. Clypeus 0.34. Eye size and inter-
distances: AME 0.75, ALE 0.80, PME 0.28, PLE 0.50, AME–AME 0.58, AME–ALE
0.37, AME–PME 0.19, ALE–ALE 1.92, ALE–PME 0.63, PME–PME 1.92, PME–
PLE 0.11, PLE–PLE 2.39, ALE–PLE 0.31, AME–PLE 0.52. Ratio of the eye group
width to length 1.84.
Maxilla: length to width: 1.94. Labium: 1.79 long, 2.45 wide, with 109 cuspules
spaced by one diameter from each other on anterior third.
Chelicera: basal segment with 12 teeth in a row and some small teeth on promar-
gin. Sternum: 7.72 long, 6.44 wide. Sigilla: anterior rounded, middle fusiform, both
less than one diameter from margin.
Legs: Length leg IV to leg I: 1.19. Scopula: Tarsi I–IV fully scopulate. Metatarsi
I–II 3/4; III 1/2, IV 1/3 distal scopulate. IV divided by a bald area.
Type II urticating setae: 1.01–1.11 long, 0.021–0.023 wide.
Palp (Figs 131–134): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 4.38 long in retrolateral
view, about 3 times tegulum’s length. Medial portion and tegulum’s margin form an
acute angle in retrolateral view. Proximal part very curved in frontal view; thin distal
width, abruptly narrowing distally; basal, middle, and distal width, 0.45, 0.24, 0.12,
respectively. Tegulum: 2.68 long, 1.57 high in retrolateral view. Cymbium subtrian-
gular with subequal lobes, having a well-developed rounded process on retrolateral
lobe, bearing thick setae (Fig. 135, 139–141).
Tibial apophysis (Figs 136–138): single branch on prolateral leg I, with well-developed
base and grouped spiniform setae distally. Male metatarsus I touches retrolaterally tibial
apophysis’ setae when folded.
Coloration (Fig. 146): abdomen with long greyish brown guard-setae homogeneously
distributed and dark short body setae. Abdomen venter brown.
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi
contrasting with other lighter articles (Fig. 142) and abdomen dorsum reddish, with
96 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 142–147. Avicularia rufa Schiapelli & Gerschman, 1945, habitus and retreat. 142 immature
143 juvenile 144 old juvenile 145 female 146 male 147 retreat on bromelid. Photos: 142–145, 147 R.
Bertani; 146 W. Bokermann.

dorsal central longitudinal black stripe disconnected from transversal black stripes
(Fig. 143). When mature, both males and females lose this pattern (Figs 145–146).
Distribution. Brazil (states of Amazonas, Amapá, Pará, Acre, Rondônia and Mato
Grosso), Ecuador, Peru and Bolivia (Fig. 50).
Natural history. In northern region of Brazil, such as in state of Rondônia, speci-
mens are commonly found in babaçu palms (Fig. 148). They build their retreats in
palm axis or using underside of the palms leaves (Figs 149–150, 152). They can also
build retreats over larger tree bark (Fig. 151) or inside bromeliads (Fig. 147). Speci-
mens are usually found on human-made structures, such as on houses and other build-
ings (Fig. 153).
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 97

Figures 148–153. Avicularia rufa Schiapelli & Gerschman, 1945, habitat and retreats. 148 Babaçu
palms, a common habitat of A. rufa in state of Rondônia, Brazil 149 retreat of immature on a palm tree
150 detail of retreat 151 retreat over a tree bark 152 use of intern side of babaçu palm to build retreat
153 in human buildings. Photos: 148–152 R. Bertani; 153 C. S. Fukushima.
98 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Avicularia purpurea Kirk, 1990


Figs 19, 105, 154–172, 304

Avicularia purpurea Kirk, 1990: 15, figs 1–5 (holotype female, Ecuador, Tena [0°58'S,
77°48'W], approximately 500 m a.s.l., H. Hirschi col., 1989, BMNH 1990.5.22.1
and paratype female, Peru, H. Hirschi col., 1990, BMNH 1990.5.22.2; exam-
ined); Bullmer et al. 2006: 7, figs 16–19, 21; World Spider Catalog 2016.

Diagnosis. Females of A. purpurea resemble those of A. merianae sp. n. by the sper-


mathecae with weakly-sclerotized area at least same length of well-sclerotized area (Figs
154–157). They differ from A. merianae sp. n. by the velvety black abdomen as well
as legs and palps with dark short body setae with very intense purple sheen (Figs 168,
170). Males of A. purpurea resemble those of A. avicularia, A. rufa. A. juruensis, A.
variegata stat. n., A. taunayi, and A. merianae sp. n. by tibial apophysis on leg I with
well-developed base and grouped spiniform setae distally (Fig. 163). They can be dis-
tinguished from all these species except A. merianae sp. n. by cymbium lacking well-
developed process on retrolateral lobe (Fig. 162). It differs from A. merianae sp. n. by
velvety black abdomen, and legs and palps with dark short body setae with very intense
purple sheen (Figs 169, 171).
Material examinad. 1 female, Ecuador, Napo, 20 km East of Puerto Napo, Aliña-
huí (1°0'S, 77°25'W), 450 m, V. D. & B. Roth col., January 1994 (CAS 11); 1 male,
Ecuador, Napo, 25 km East of Puerto Napo [1°01'S, 77°43'W], jungle Aliñahuí, 450
m, E. S. Ross col, January–February 1991 (CAS 3).
Additional material. COLOMBIA: Putumayo: Mocoa [1°09'N, 76°39'W], Vda.
Pepino, 500 m asl, 1 female, T. Sanjuan col., 30 April 1997 (ICN–Ar-1990); ECUA-
DOR: Napo: Puerto Napo, 20 km East, Aliñahuí (1°0'S, 77°25'W), 450 m, 1 male, V.
D. & B. Roth col., June 1994 (CAS 7); [1°01'S, 77°43'W], 1 male, V. Roth col., June
1994 (AMNH RW52); Río Napo [2°00'S, 74°20'W], 1 female, Gerhard col., 1994
(IBSP 11597); Tena, Cabañas Aliñahui (1°02'54.00"S, 77°36'05.00"W), 1 female,
R. Baxter col., August 1994, in silk retreat on side of tree (AMNH RW54); PERU:
Loreto: Iquitos, Rio Momon, Amazon Camp (3°41'16.00"S, 73°16'48"W), T. Mason
col., May 1994 (AMNH RW33); 1 male, W. Lamar col., 20 August 1997 (MNRJ
06913); Río Yarapa [4°31'S, 73°22'W], 1 female, R. C. West col., 16 November 1993,
in a palm tree at night (AMNH RW44); Río Nanay [3°48'S, 73°23'W], 1 female,
R. C. West col., 5 November 1993 (AMNH RW45); Ucayali Río Ucayali [7°34'S,
74°20'W], 1 female, R. C. West col., 14 November 1993, in silk retreat on a citrus
tree (AMNH RW48).
Female. Redescription. CAS. Carapace: 15.4 long, 14.02 wide, 3.98 high. Cheli-
cera: 5.38 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 11.69, 7.20,
8.54, 7.25, 5.15, 39.83. II: 10.74, 6.84, 7.99, 7.13, 4.89, 37.59. III: 9.75, 5.93, 7.33,
7.15, 4.42, 34.58. IV: 12.08, 6.47, 10.14, 9.51, 4.31, 42.51. Palp: 8.10, 5.10, 4.72, –,
5.41, 23.33. Midwidths: femora I–IV= 2.81, 3.08, 3.07, 2.90, palp= 2.38; patellae I–
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 99

Figures 154–157. Avicularia purpurea Kirk, 1990, spermathecae variation. 154 Puerto Napo, department
of Napo, Ecuador (CAS 11) 155 Tena, department of Napo, Ecuador (AMNH RW54) 156 holotype, Tena,
department of Napo, Ecuador (BMNH 1990.5.22.1) 157 Iquitos, department of Loreto, Peru (AMNH
RW33). Scale bars = 1 mm.

IV= 2.95, 2.70, 2.87, 2.98, palp= 2.57; tibiae I–IV= 2.39, 2.30, 2.56, 2.82, palp= 2.49;
metatarsi I–IV= 1.92, 1.72, 1.86, 1.71; tarsi I–IV= 2.18, 2.19, 2.01, 2.13, palp= 2.36.
Abdomen: 19.26 long, 13.62 wide. Spinnerets: PMS, 1.81 long, 1.19 wide, 1.07 apart;
PLS, 2.28 basal, 1.09 middle, 2.74 distal; midwidths 2.38, 1.78, 1.41, respectively.
Carapace: 1.10 times longer than wide; cephalic region moderately raised, thoracic
striae conspicuous.
Fovea: deep, recurve, 2.03 wide.
Eyes: eye tubercle 1.17 high, 2.21 long, 3.43 wide. Clypeus 0.12. Anterior row of
eye procurve. Posterior row of eyes slightly recurve. Eye sizes and interdistances: AME
100 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 158–165. Avicularia purpurea Kirk, 1990, male (CAS 3; except palpal bulb in prolateral view,
CAS 7). 158–161 left palpal bulb 158 prolateral 159 retrolateral 160 frontal 161 dorsal 162 left cymbium,
dorsal 163–165 left tibial apophysis of leg I 163 prolateral 164 ventral 165 retrolateral. Scale bars = 1 mm.

0.75, ALE 0.72, PME 0.27, PLE 0.62, AME–AME 0.6, AME–ALE 0.47, AME–PME
0.25, ALE–ALE 2.19, ALE–PME 0.63, PME–PME 2.06, PME–PLE 0.23, PLE–PLE
2.67, ALE–PLE 0.48, AME–PLE 0.67.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 101

Maxilla length to width: 1.89. Cuspules: 100–200 spread over ventral inner heel.
Labium: 2.03 long, 2.46 wide, with 108 cuspules spaced by one diameter from each
other on anterior half. Labio-sternal groove shallow and flattened, sigilla not evident.
Chelicera: basal segment with 14 teeth and some small teeth on promargin. Ster-
num: 7.56 long, 6.27 wide. Sigilla: three pairs, elipsoidal posterior, in 45°angle, less
than one diameter from margin; fusiform median, less than one diameter from margin;
anterior not evident.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.07. Clavate trichobothria:
distal 1/2 of tarsi I–IV. Scopula: Tarsi I–IV fully scopulate, IV divided by a wide band
of setae. Metatarsi I–II fully scopulate; III on distal 2/3; IV, on distal 1/3. IV divided
by wide band of setae.
Type II urticating setae: 0.53–0.58 long, 0.011–0.014 wide.
Spermathecae (Fig. 154): two completely separated, not-twisted long spermathe-
cae, with walls lacking projections or lobes and accentuated outwards curvature me-
dially. Midwidth as wide as its base width and weakly-sclerotized area at least same
length of well-sclerotized area.
Color pattern (Figs 168): carapace brown with golden short body setae with very
intense purple sheen. Carapace border with long setae the same color as dorsal cara-
pace short body setae. Coxae, labium, sternum and maxillae brown, with the same
color of ventral femora. All ventral parts, specially sternum, covered by longer setae.
Legs and palps with brown short body setae with very intense purple sheen and brown
long guard-setae. Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen
dorsum with long brown guard-setae homogeneously distributed and black short body
setae, with velvety aspect (Fig. 304). Abdomen venter brown.
Male. Description. CAS. Carapace: 11.98 long, 10.74 wide, 2.85 high. Chelicera:
3.79 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 12.59, 6.43, 9.70,
9.13, 5.68, 43.53. II: 11.82, 5.86, 9.35, 8.78, 5.18, 40.99. III: 10.47, 5.26, 8.36, 8.54,
4.77, 37.40. IV: 13.16, 5.27, 11.04, 11.25, 4.68, 45.40. Palp: 6.97, 4.35, 5.55, –, 2.32,
19.15. Midwidths: femora I–IV= 2.08, 2.49, 2.18, 2.20, palp= 1.71; patellae I–IV=
2.22, 2.21, 2.15, 2.28, palp= 1.68; tibiae I–IV= 1.64, 1.73, 1.86, 1.89, palp= 1.65;
metatarsi I–IV= 1.35, 1.31, 1.25, 1.23; tarsi I–IV= 1.63, 1.58, 1.65, 1.53, palp= 1.69.
Abdomen: 13.09 long, 7.88 wide. Spinnerets: PMS, 1.57 long, 0.79 wide, 0.29 apart;
PLS, 2.00 basal, 1.12 middle, 2.78 distal; midwidths 1.42, 1.29, 0.92, respectively.
As in female, except:
Carapace: 1.12 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Fovea: straight, 1.14 wide.
Eyes: eye tubercle 0.60 high, 1.73 long, 2.54 wide. Clypeus 0.30. Eye size and
interdistances: AME 0.58, ALE 0.67, PME 0.22, PLE 0.42, AME–AME 0.48, AME–
ALE 0.36, AME–PME 0.20, ALE–ALE 1.66, ALE–PME 0.74, PME–PME 1.52,
PME–PLE 0.07, PLE–PLE 1.95, ALE–PLE 0.40, AME–PLE 0.43.
Maxilla: length to width: 2.07. Labium: 1.33 long, 1.92 wide, with 120 cuspules
spaced by one diameter from each other on the anterior half.
102 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 166–171. Avicularia purpurea Kirk, 1990, habitus and retreat. 166 silken retreat over tree bark
167 immature 168–169 northern form, from Tena, Ecuador 168 female 169 male 170–171 southern
form, from Rio Momon, Peru 170 female 171 male. Photos: 166, 168, 170 R. C. West; 167 R. Bertani;
169 R. Baxter; 171 W. Lamar.

Chelicera: basal segment with 11 teeth and some small teeth on promargin. Ster-
num: 5.78 long, 4.81 wide. Sigilla: three pairs, posterior and median rounded, large,
less than one diameter from margin; anterior not evident.
Legs: Length leg IV to leg I: 1.04. Scopula: Metatarsi IV scopulate in distal 1/4.
IV divided by band of setae.
Type II urticating setae: 0.78–0.83 long, 0.015–0.017 wide.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 103

Figure 172. Map showing records of Avicularia purpurea Kirk, 1990 morphotypes.

Palp (Figs 158–161): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 3.5 long in retrolateral view,
about 3.0 times tegulum’s length. Medial portion and tegulum’s margin form an acute
angle in retrolateral view. Proximal part very curved in frontal view; thin distal width,
104 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

tapering distally; basal, middle, and distal width 0.68, 0.18, 0.05, respectively. Tegu-
lum: 1.96 long, 1.66 high in retrolateral view. Cymbium subtriangular with subequal
lobes, lacking well-developed process on retrolateral lobe (Fig. 162).
Tibial apophysis (Figs 163–165): single branch on prolateral leg I, with well-devel-
oped base and grouped spiniform setae distally. Male metatarsus I touches retrolater-
ally tibial apophysis’ setae when folded.
Color pattern ontogeny. Juveniles present green metallic sheen, all articles with
blackish color and abdomen dorsum with central longitudinal black stripe connected
with all transversal black stripes of each side (Fig. 167). When mature, both males and
females lose this pattern (Figs 168 –171).
Variation. We found two different morphotypes among the examined material
(Fig. 172). The northern form is the same holotype’s morphotype (Figs 168–169).
The southern form has very discrete grizzled setae in palps and legs, and less intense
purple sheen in short body setae of carapace, legs, and palps (Figs 170–171). Females
of the southern form have abdomen with light brown guard-setae homogeneously dis-
tributed, with a couple of reddish brown lateral stripes until penultimate or ultimate
molt, distinct from the velvety black dorsal abdomen of the northern morphotype.
Males from southern and northern form showed no morphological difference among
them (Figs 169, 171).
Distribution. Colombia, Ecuador, and Peru (Fig. 105).
Natural history. Specimens were found in anthrophized areas, such as cultivated
areas and cattle pastures with dispersed trees, near Tena, Ecuador (Kirk 1990). They are
arboreals, making their retreats in tree holes, epiphytes, and inside houses and other hu-
man constructions (Kirk 1990). Specimens of the southern form were found in natural
cavities of living palms or citrus-like trees in open forested fringes or in gallery forest areas
(Rick West, pers. comm.). They use bark and organic debris to camouflage the partially
exposed webbed retreat, which lies about 25 cm from soil (Rick West, pers. comm.).

Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006


Figs 19, 90, 173–187, 309

Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006: 3, figs 1–10, 20, 22 (holo-
type male, Ecuador, Oriente, nahe Misahualli [1°02'S, 77°41'W], Bullmer leg.,
August 1997, SMF 57125 and paratype female, Ecuador, Oriente, nahe Misahualli
[1°02'S, 77°41'W], Bullmer leg., August 1997, SMF 57126, examined); World
Spider Catalog 2016.

Diagnosis. Females of A. hirschii resemble those of A. avicularia and A. rufa by the


leg IV longer than leg I. They can be distinguished from them by twisted spermatheca
(Fig. 181). Males of A. hirschii resemble those of A. minatrix, A. lynnae sp. n. and A.
caei sp. n. by tibia I with discrete elevation covered by a cluster of setae in apical por-
tion, on prolateral side (Fig. 179). They can be distinguished from males of A. lynnae
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 105

Figures 173–180. Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006, male holotype (SMF
57125). 173–176 right palpal bulb (mirrored) 173 prolateral 174 retrolateral 175 frontal 176 dorsal
177 right cymbium, dorsal (mirrored) 178–180 left tibia I 178 prolateral 179 ventral 180 retrolateral.
Scale bars = 1 mm.

sp. n. and A. caei sp. n. by shorter embolus, about 3.0 to 3.5 times tegulum’s width
in retrolateral view (Fig. 174) and from A. minatrix by having prominece on tegulum
(Fig. 175) and cymbium with process on retrolateral lobe (Fig. 177).
106 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Additional material. BRAZIL: Acre: Senador Guiomard [10°08'S, 67°44'W], 1


female, C. Alexandre col., 12–17 July 2013 (MNRJ 06912); Estação Ecológica Rio
Acre [10°59'S, 70°13'W], M. A. de Freitas col., 4–18 February 2016 (MNRJ 06911);
ECUADOR: Pastaza: 25 km east of Puerto Napo [1°01'S, 77°43'W], selva Aliñahuí,
450 m, I–II, 1 female, 1 immature, E. S. Ross col., 1991 (CAS 1, CAS 2); PERU: Lore-
to: confluence of Rio Zumun and Rio Yaguasyacu [3°21'S, 71°58'W] (Rio Yahnasyacu
[sic]), 2 juvenile females, J. Becker col. (MNRJ 13759); Iquitos, Rio Nanay [3°48'S,
73°23'W], 1 juvenile female, R. C. West col., 5 November 1993 (AMNH RW50); 1
juvenile female, Rio Tigre, Cristo Rey village [3°58'S, 74°16'W], W. Lamar col., 19
July 1998, in light silken retreat between broad leaves 1.5 m above ground in forest
behind village (MNRJ 06910).
Male. Redescription. SMF 57125. Carapace: 10.83 long, 9.32 wide, 2.50 high.
Chelicera: 2.65 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 10.02, 5.63,
7.80, 7.72, 4.51, 35.68. II: 9.66, 4.99, 7.44, 6.88, 4.48, 33.45. III: 8.44, 4.27, 6.17,
6.94, 3.66, 29.48. IV: 10.20, 4.64, 8.79, 9.30, 3.68, 36.61. Palp: 6.08, 3.63, 4.57, –,
1.78, 16.06. Midwidths: femora I–IV= 1.94, 1.77, 2.14, 2.15, palp= 1.45; patellae I–
IV= 2.9, 1.87, 1.97, 1.76, palp= 1.56; tibiae I–IV= 1.53, 1.66, 1.53, 1.60, palp= 1.67;
metatarsi I–IV= 1.02, 1.22, 1.04, 1.17; tarsi I–IV= 1.24, 1.47, 1.38, 1.50, palp= 1.57.
Abdomen: 10.95 long, 6.04 wide. Spinnerets: PMS, 1.02 long, 0.46 wide, 0.1 apart;
PLS, 1.82 basal, 0.93 middle, 2.21 distal; midwidths 1.04, 0.82, 0.60, respectively.
Carapace: 1.16 times longer than wide; cephalic region not raised.
Fovea: deep, straight, 1.29 wide.
Eyes: eye tubercle 0.66 high, 1.59 long, 2.30 wide. Clypeus absent. Anterior row
of eyes procurve, posterior slightly recurve. Eye size and interdistances: AME 0.69,
ALE 0.72, PME 0.30, PLE 0.44, AME–AME 0.34, AME–ALE 0.16, AME–PME
0.10, ALE–ALE 1.45, ALE–PME 0.55, PME–PME 1.50, PME–PLE 0.03, PLE–PLE
1.75, ALE–PLE 0.23, AME–PLE 0.43.
Maxilla: length to width: 2.53. Cuspules: 112 spread over ventral inner heel. La-
bium: 1.00 long, 1.64 wide, with 54 cuspules spaced by one diameter from each other
on anterior third. Labium sternal groove shallow, flat, sigilla not evident.
Chelicera: basal segment with 9 teeth and some small teeth on promargin. Ster-
num: 5.01 long, 3.93 wide. Sigilla: only posterior pair evident, rounded, less than one
diameter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.03. Clavate trichobothria:
2/3 distal tarsi I–IV. Scopula: Tarsi I–IV fully scopulate. IV with some sparse setae.
Metatarsi I–II fully scopulate; III 2/3; IV 1/2 distal scopulate. IV divided by a wide
row of setae.
Type II urticating setae: 0.10–0.11 long (according to original description, since
holotype abdomen is bald).
Palp (Figs 173–176): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 2.7 long in retrolateral view,
about 3.5 times tegulum’s length. Medial portion and tegulum’s margin form an acute
angle in retrolateral view. Proximal part very curved in frontal view; thin distal width,
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 107

Figures 181–183. Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006, spermathecae variation.
181 paratype, Oriente, department of Napo, Ecuador (SMF 57126) 182 Estação Ecológica Rio Acre,
Assis Brasil, state of Acre, Brazil (MNRJ 06911) 183 Senador Guiomard, state of Acre, Brazil (MNRJ
06912). Scale bars = 1 mm.

tapering distally; basal, middle, and distal width of 0.35, 0.11, 0.03, respectively. Teg-
ulum: 1.51 long, 0.93 wide in retrolateral view. Cymbium subtriangular with subequal
lobes. Cymbium with well-developed rounded process on retrolateral lobe, bearing
thin setae (Fig. 177).
Tibia I with discrete elevation covered by cluster of setae in apical portion, on
prolateral side (Figs 178–180, 309).
Color pattern: carapace brown with golden short body setae with pink sheen. Cara-
pace border with long setae the same color as dorsal carapace short body setae. Coxae,
labium, sternum and maxillae light brown, same color of ventral femora. Legs and palps
with brown short body setae with golden sheen and reddish brown long guard-setae.
Posterior legs darker, blackish. Tarsi III and IV with reddish central well-developed
tufts (not detected, but informed on original description). Tarsi with “U” shaped or-
ange stripe. Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen dorsum
with long reddish guard-setae and dark short body setae. Ventral abdomen light brown.
Female. Redescription. SMF 57126. Carapace: 11.92 long, 11.37 wide, 3.46
high. Chelicera: 3.84 long. Legs (femur, patella, tibia, metatarsus, tarsus and total): I:
108 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 184–187. Avicularia hirschii Bullmer, Thierer-Lutz & Schmidt, 2006, habitus. 184 immature,
Tiguino, department of Pastaza, Ecuador 185 juvenile, Senador Guiomard, state of Acre, Brazil 186 fe-
male, Estação Ecológica Rio Acre, Assis Brasil, state of Acre, Brazil 187 female feeding on an insect, Rio
Momón, department of Loreto, Peru. Photos: 184 W. Lamar; 185 S. Albuquerque, 186 M. A. de Freitas;
187 R. C. West.

9.69, 5.45, 6.78, 6.43, 4.81, 33.16. II: 9.11, 5.39, 6.69, 6.05, 4.38, 31.62. III: 8.23,
4.90, 5.49, 6.53, 3.94, 29.09. IV: 10.19, 5.64, 8.73, 9.22, 4.57, 38.35. Palp: 6.99,
4.14, 4.40, –, 5.08, 20.61. Midwidths: femora I–IV= 2.00, 1.97, 2.49, 2.21, palp=
1.93; patellae I–IV= 2.10, 1.92, 2.27, 2.47, palp= 1.74; tibiae I–IV= 1.90, 1.83, 2.09,
2.27, palp= 1.73; metatarsi I–IV= 1.60, 1.39, 1.58, 1.82; tarsi I–IV= 1.91, 1.71, 1.81,
1.96, palp= 1.79. Abdomen: 12.61 long, 10.24 wide. Spinnerets: PMS, 1.38 long,
1.07 wide, 0.07 apart; PLS damaged.
As in male, except:
Carapace: 1.05 times longer than wide; thoracic striae conspicuous.
Fovae: recurve, 1.46 wide.
Eyes: eye tubercle 0.94 high, 2.02 long, 2.88 wide. Clypeus 0.08. Eye size and
interdistances: AME 0.65, ALE 0.67, PME 0.22, PLE 0 .68, AME–AME 0.41, AME–
ALE 0.40, AME–PME 0.14, ALE–ALE 1.74, ALE–PME 0.75, PME–PME 1.57,
PME–PLE 0.03, PLE–PLE 1.99, ALE–PLE 0.24, AME–PLE 0.50.
Maxilla: length to width: 2.03. Cuspules: 146. Labium: 1.50 long, 2.40 wide, with
64 cuspules spaced by one diameter. Labio-sternal groove with two separate, large sigilla.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 109

Chelicera: basal segmente with 12 teeth and some small teeth on promargin. Ster-
num: 6.66 long, 4.30 wide. Sigilla: only posterior pair evident, elipsoidal, less than one
diameter from margin.
Legs: Formula: IV I II III. Length leg IV to leg I: 1.16. Scopula: Tarsi IV divided
by some setae on base; metatarsi IV scopulate on distal 1/4.
Type II urticating setae: 0.38–0.50 long, 0.011–0.014 wide
Spermathecae (Fig. 181): two completely separated, twisted long spermatheca,
with walls lacking projections or lobes and accentuated outwards curvature medially.
Midwidth as wide as its base width and weakly-sclerotized area shorter than half the
length of well-sclerotized area.
Color pattern: dorsal abdomen with vivid reddish guard-setae grouped on lateral
area and black short body setae (Fig. 186).
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi
contrasting with other lighter articles and black central longitudinal stripe on abdomen
dorsum (Fig. 184).
Distribution. Brazil (state of Acre), Ecuador and Peru (Fig. 90).
Natural history. Types were collected in trees in an old pasture, in different trees
about 5 m from each other, surrounding by grass (Bullmer et al. 2006). Immature male
was in a tree in a retreat about 1.6 m above the ground and female was also in a tree; both
retreats were made by web, soil and moss, 60 cm from ground (Bullmer et al. 2006). A
found eggsac hatched in captivity and contained 38 specimens (Bullmer et al. 2006). In
the state of Acre, Brazil, specimens were found in silken retreats constructed inside tree
trunks (C. Alexandre, pers. comm.; M. A. Freitas and A. Zanotti, pers. comm.).

Avicularia merianae sp. n.


http://zoobank.org/C2A7B023-1FA4-4C74-B205-0549E650C4DA
Figs 9–10, 19, 105, 188–200

Diagnosis. Females of A. merianae sp. n. resemble those of A. purpurea by the sper-


mathecae with weakly-sclerotized area at least same length of well-sclerotized area
(Fig. 196). They differ from A. purpurea by dark brown dorsal abdomen, and by lack-
ing dark short body setae with very intense purple sheen on legs and palps. Males of A.
merianae sp. n. resemble those of A. avicularia, A. rufa. A. juruensis, A. variegata stat.
n., A. taunayi and A. purpurea by tibial apophysis on leg I with well-developed base and
grouped spiniform setae distally (Fig. 194). They can be distinguished from all these
species except A. purpurea by cymbium lacking well-developed process on retrolateral
lobe (Fig. 192). It differs from A. purpurea by lacking velvety black abdomen and dark
short body setae with very intense purple sheen on legs and palps.
Etymology. It was named after Maria Sybilla Merian, the German-born naturalist
who drew the famous engraving of a specimen of Avicularia eating a bird, in recognition
to her importance for Natural Sciences. This extraordinary woman was one of the pioneer-
ing female scientists and a remarkable artist. This name is considered feminine in gender.
110 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 188–195. Avicularia merianae sp. n., male holotype (AMNH Pe102). 188–191 left palpal bulb
188 prolateral 189 retrolateral 190 frontal 191 dorsal 192 left cymbium, dorsal 193–195 left tibial
apophysis of leg I 193 prolateral 194 ventral 195 retrolateral. Scale bars = 1 mm.

Material examined. Holotype male, 1 female and 3 immatures paratypes, Peru,


department of San Martín, Hara, 20 miles SE from Moyobamba [6°01'S, 76°58'W],
F. Woytkowski col., 1–30 June 1947 (AMNH Pe102).
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 111

Additional material. PERU: San Martín: Tarapoto [6°07'S, 75°57'W], 830 m


a.s.l., 7 females, 1 juvenile female, 2 immature males, 15 immatures, 1 spiderling, F.
Woytkowski col., 10–18 February 1947 (AMNH Pe106, Pe105, Pe108); Ekin, west
of Tarapoto, 1 young female, under fallen tree, 890 m a.s.l., F. Woytkowski col., 9–21
March 1947 (AMNH Pe104); 1 immature male, under fallen bark, same collector
and date (AMNH Pe107); Moyobamba [6°01'S, 76°58'W], Hara, 20 miles southeast
of Moyobamba, 4 males, 12 females, 3 juvenile females, 22 immatures, 1 spiderling,
F. Woytkowski, 1–30 June 1947 (AMNH Pe98, Pe103, Pe100, Pe53, Pe99, Pe101,
AMNH tube, AMNH tube); Rio Huallaga [7°47'S, 76°11'W], 1 immature male, J. C.
Pallister col., 15 December 1946 (AMNH Pe112).
Male. Description. AMNH Pe102. Carapace: 13.49 long, 12.80 wide, 3.48 high.
Chelicera: 3.46 long. Legs (femur, patella, tibia, metatarsus, tarsus and total): I: 13.41,
7.59, 10.39, 9.96, 6.21, 47.56. II: 12.75, 7.14, 9.89, 9.44, 5.48, 44.70. III: 11.01,
5.86, 8.57, 9.13, 5.46, 40.03. IV: 14.15, 7.03, 12.05, 11.85, 5.54, 50.62. Palp: 7.98,
4.78, 5.87, –, 2.45, 21.08. Midwidths: femora I–IV=2.45, 2.51, 2.39, 2.55, palp=
1.85; patellae I–IV= 2.58, 2.90, 2.51, 2.89, palp= 1.99; tibiae I–IV= 2.09, 2.05, 2.06,
2.16, palp= 1.69; metatarsi I–IV= 1.68, 1.57, 1.53, 1.33; tarsi I–IV= 1.56, 1.70, 1.67,
1.51, palp= 1.88. Abdomen: 16.35 long, 10.45 wide. Spinnerets: PMS, 2.15 long,
0.94 wide, 0.44 apart; PLS, 2.38 basal, 1.88 middle, 3.00 distal; midwidths 1.30, 1.26,
1.17, respectively.
Carapace: 1.06 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Fovea: deep, recurve, 1.78 wide.
Eyes: eye tubercle 1.16 high, 1.97 long, 2.61 wide. Clypeus absent. Anterior eye
row procurve. Posterior eye row slightly recurve. Eye size and interdistances: AME
0.72, ALE 0.72, PME 0.21, PLE 0.54, AME–AME 0.32, AME–ALE 0.37, AME–
PME 0.12, ALE–ALE 1.78, ALE–PME 0.90, PME–PME 1.71, PME–PLE 0.07,
PLE–PLE 2.07, ALE–PLE 0.39, AME–PLE 0.40.
Maxilla: length to width: 1.96. Cuspules: 236 spread over ventral inner heel. Labi-
um: 1.62 long, 1.89 wide, with 101 cuspules spaced by one diameter from each other,
on anterior half. Labio-sternal groove shallow, flat, with no evident sigilla.
Chelicera: basal segment with 12 teeth and some small teeth on promargin. Ster-
num: 7.04 long, 6.04 wide. Sigilla: anterior pair not evident, middle fusiform, poste-
rior ellipsoidal, in a 45°angle, both less than one diameter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.06. Clavate trichobothria:
distal 2/3 of tarsi I–IV. Scopula: Tarsi I–IV fully scopulate. Metatarsi I fully scopulate,
II–III 2/3; IV 1/4 distal scopulate. IV divided by a row of setae.
Type II urticating setae: 0.82–0.92 long, 0.016–0.019 wide.
Palp (Figs 188–191): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 3.83 long in retrolateral
view, about 3.0 times tegulum’s length. Medial portion and tegulum’s margin form an
acute angle in retrolateral view. Proximal part very curved in frontal view; thin distal
112 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 196–198. Avicularia merianae sp. n., spermathecae variation. 196 paratype, Moyobamba, de-
partment of San Martín, Peru (AMNH Pe102) 197 Tarapoto, department of San Martín, Peru (AMNH
Pe106) 198 Moyobamba, department of San Martín, Peru (AMNH Pe100). Scale bars = 1 mm.

width, tapering distally; basal, middle, and distal width 0.81, 0.21, 0.03, respectively.
Tegulum: 2.07 long, 1.20 high in retrolateral view. Cymbium subtriangular with sub-
equal lobes, lacking process on retrolateral lobe (Fig. 192).
Tibial apophysis (Figs 193–195): a single branch on prolateral leg I, with well-
developed base and grouped spiniform setae distally. Male metatarsus I touches retro-
laterally tibial apophysis’ setae when folded.
Color pattern: carapace brown with golden short body setae with pink sheen.
Carapace border with long setae the same color as dorsal carapace short body setae.
Coxae, labium, sternum and maxillae light brown, same color of ventral femora. Legs
and palps with golden brown short body setae with pink sheen and brown long guard-
setae. Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen with orange
brown guard-setae homogeneously distributed and dark brown body setae. Abdomen
venter brown.
Female. Description. AMNH Pe102. Carapace: 14.57 long, 13.15 wide, 3.83
high. Chelicera: 6.14 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I:
10.75, 6.90, 7.99, 6.72, 5.15, 37.51. II: 9.99, 6.52, 7.31, 6.41, 4.67, 34.90. III: 9.04,
5.59, 6.88, 6.54, 4.88, 32.93. IV: 11.37, 6.50, 9.66, 8.82, 4.38, 40.73. Palp: 7.95,
4.77, 4.70, –, 5.35, 22.77. Midwidths: femora I–IV= 2.68, 3.02, 3.05, 2.88, palp=
2.09; patellae I–IV= 2.82, 2.79, 2.77, 2.90, palp= 2.34; tibiae I–IV= 2.29, 2.35, 2.59,
3.11, palp= 2.29; metatarsi I–IV= 2.07, 2.05, 1.90, 2.05; tarsi I–IV= 2.24, 2.27, 2.09,
2.12, palp= 2.37. Abdomen: 17.49 long, 11.96 wide. Spinnerets: PMS, 2.08 long,
1.01 wide, 0.16 apart; PLS, 2.37 basal, 1.11 middle, 2.65 distal; midwidths 1.78, 1.49,
1.21, respectively.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 113

Figures 199–200. Avicularia merianae sp. n., habitus and retreat. 199 female 200 silken retreat over
tree bark. Photos: H.-W. Auer.

As in male, except:
Carapace: 1.11 times longer than wide.
Fovea: slightly recurve, 1.77 wide.
Eyes: eye tubercle 0.70 high, 2.07 long, 2.91 wide. Clypeus 0.30. Eye size and
interdistances: AME 0.72, ALE 0.70, PME 0.28, PLE 0.55, AME–AME 0.50, AME–
ALE 0.47, AME–PME 0.19, ALE–ALE 2.03, ALE–PME 0.71, PME–PME 1.64,
PME–PLE 0.16, PLE–PLE 2.11, ALE–PLE 0.49, AME–PLE 0.52.
Maxilla: length to width: 2.02. Labium: 1.77 long, 2.41 wide, with 96 cuspules
spaced by one diameter from each other, on anterior half.
Chelicera: basal segment with 9 teeth and some small teeth on promargin. Ster-
num: 7.17 long, 5.92 wide.
Legs: Length leg IV to leg I: 1.09 Clavate trichobothria: on distal 1/2 tarsi I–IV.
Scopula: Tarsus IV with sparse setae. Metatarsi II fully scopulate; III 1/2, IV 1/3 distal
scopulate.
Type II urticating setae: 0.49–0.60 long, 0.014–0.017 wide.
Spermathecae (Fig. 196): two completely separated, not-twisted long spermathe-
cae, with walls lacking projections or lobes and accentuaded outwards curvature me-
dially. Midwidth expanded, about 1.5 times its basal and apical portion widths and
weakly-sclerotized area at least same length of well-sclerotized area.
Color pattern (Fig. 199): as in male.
Color pattern ontogeny. brownish juveniles lacking metallic sheen, black tarsi
contrasting with other lighter articles and abdomen dorsum reddish, with dorsal cen-
tral longitudinal black stripe connected only with anterior pair of transversal black
stripes. When mature, both males and females lose this pattern.
Distribution. Peru, department of San Martín (Fig. 105).
Natural history. Unknown.
114 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Avicularia lynnae sp. n.


http://zoobank.org/4087E14F-2413-4DF6-84CE-F7D0C7B710E5
Figs 19, 90, 201–209

Diagnosis. Males of A. lynnae sp. n. resemble those of A. minatrix, A. hirschii and A.


caei sp. n. by tibia I with discrete elevation covered by a cluster of setae in apical por-
tion, on prolateral side (Fig. 207). They can be distinguished from all species except
A. caei sp. n. by very long embolus, more than 4 times tegulum’s width in retrolateral
view (Fig. 202). Males of A. lynnae sp. n. differ from male A. caei sp. n. by having
developed prominence on tegulum (Fig. 203) and by abdomen dorsum with single
central longitudinal dark stripe (Fig. 209). Female unknown.
Etymology. It was named after Lynn West, wife of mygalomorph expert Rick
West. This name is considered feminine in gender.
Material examined. Holotype male, Peru, Loreto, Rio Tigre, Cristo Rey village
[3°58'S, 74°16'W] near Iquitos, R. C. West col., 21 November 1993, crossing trail by
day (AMNH RW49); paratype male, Peru, Loreto, Brillo Nuevo [3°09'S, 71°46'W]
(Brillo Neuvo [sic]), Rio Yaguasyacu, B. Lamar col. (AMNH–RCW).
Additional material. ECUADOR: Pastaza: Tigüino [1°10'S, 76°57'W], 1 male,
B. Lamar col., September 1990, found in a bird capture net (AMNH–RCW); PERU:
Marañón (Marauon [sic] [river or province?]), 1 male, Bristol, October 1927 (AMNH
Pe96); Madre de Dios: Zona Reservada Pakitza [11°56'S, 71°17'W], 356 m asl, 1 male,
Igidio & D. Silva col., 13 August 1992 (MUSM-ENTO 500685).
Male. Description. AMNH RW49. Carapace: 10.87 long, 10.25 wide, 2.40 high.
Chelicera: 3.07 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 11.9, 5.4,
9.2, 9.3, 5.0, 40.8. II: 11.1, 5.2, 8.9, 8.6, 4.7, 38.5. III: 10.0, 4.6, 8.0, 8.6, 4.6, 35.8.
IV: 12.2, 4.9, 10.6, 12.6, 4.0, 44.3; Palp: 6.8, 3.7, 5.5, –, 2.1, 18.1; Midwidths: fem-
ora I–IV= 1.18, 1.7, 2.14, 1.9, palp= 1.6; patellae I–IV= 1.9, 2.2, 2.0, 2.1, palp= 1.8;
tibiae I–IV= 1.7, 1.6, 1.4, 1.5, palp= 1.4; metatarsi I–IV= 1.1, 1.2, 1.1, 1.2; tarsi I–IV=
1.2, 1.3, 1.4, 1.2, palp= 1.3. Abdomen: 10.77 long, 8.49 wide. Spinnerets: PMS, 1.05
long, 0.46 wide, 0.13 apart; PLS, 1.50 basal, 0.85 middle, 2.17 distal; midwidths 0.96,
0.76, 0.68, respectively.
Carapace: 1.06 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Fovea: shallow, straight, 0.64 wide.
Eyes: eye tubercle 0.91 high, 1.80 long, 2.41 wide. Clypeus absent. Anterior row
of eyes slightly procurve, posterior slightly recurve. Eye size and interdistances: AME
0.62, ALE 0.61, PME 0.19, PLE 0.55, AME–AME 0.41, AME–ALE 0.27, AME–
PME 0.14, ALE–ALE 1.54, ALE–PME 0.39, PME–PME 1.50, PME–PLE 0.04,
PLE–PLE 1.96, ALE–PLE 0.19, AME–PLE 0.34.
Maxilla: length to width: 2.57. Cuspules: about 85 spread over ventral inner heel.
Labium: 1.08 long, 1.72 wide, with 54 cuspules spaced by more than one diameter
from each other, on anterior half. Labio-sternal groove shallow, flat, two slightly sepa-
rate, large sigilla.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 115

Figures 201–208. Avicularia lynnae sp. n., male holotype (AMNH RW49). 201–204 left palpal bulb
201 prolateral 202 retrolateral 203 frontal 204 dorsal 205 left cymbium, dorsal 206–208 left tibia I
206 prolateral 207 ventral 208 retrolateral. Scale bars = 1 mm.

Chelicera: basal segment with 12 teeth and some small teeth on promargin. Ster-
num: 6.0 long, 3.91 wide. Sigilla: three pairs, anterior rounded, middle fusiform, pos-
terior rounded, set at 45°angle, all close to margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.09. Clavate trichobothria:
distal 2/3 tarsi I–IV. Scopula: Tarsi I–IV fully scopulate. Metatarsi I–II fully scopulate;
III 1/2, IV 1/5 distal scopulate. IV divided by a bald area.
116 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 209. Avicularia lynnae sp. n., habitus, male from department of Loreto, Peru. Photo: R. C. West.

Type II urticating setae: 0.73–0.87 long, 0.013–0.019 wide.


Palp (Figs 201–204): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 3.91 long in retrolateral view,
about 4.5 times tegulum’s length. Medial portion and tegulum’s margin form an acute
angle in retrolateral view. Proximal part very curved in frontal view; thin distal width,
tapering distally; basal, middle and distal width of 0.26, 0.18, 0.03, respectively. Tegu-
lum: 1.51 long, 0.89 high in retrolateral view. Cymbium subtriangular with subequal
lobes, with well-developed process bearing thin setae on retrolateral lobe (Fig. 205).
Tibial I with a discrete elevation covered by a cluster of setae in apical portion, on
prolateral side (Figs 206–208).
Color pattern (Fig. 209): carapace brown with golden short body setae and thick
dark longer setae spread over the carapace. Carapace border long setae the same color
as dorsal carapace short body setae. Coxae, labium, sternum and maxillae light brown,
same color as ventral femora. Legs and palps with brown short body setae and orange
brown long guard-setae. Leg rings on distal femora, tibiae and metatarsi whitish. Ab-
domen dorsum with long guard-setae homogeneously distributed, lateral orange short
body setae and black short body setae forming a central longitudinal stripe. Ventral
abdomen light brown.
Color pattern ontogeny. Immatures are unknown.
Distribution. Ecuador and Peru (Fig. 90).
Natural history. Specimens were found in a silken retreat in a curled living leaf
(W. Lamar, pers. comm. to R. C. West).
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 117

Female unknown.
Remarks. Aviculariinae diversity in Ecuador, Peru and Colombia is poorly known,
and certainly underestimated; specimens are rare in arachnological collections. Thus,
the identity of some specimens collected in these countries should be analyzed carefully.
Avicularia lynnae sp. n. specimens were collected in Peru and Ecuador, and its female is
unknown. The species is sympatric with A. hirschii and resembles it by having tibia I with
discrete elevation covered by a cluster of setae in apical portion on prolateral side, and
by having cymbium with thin setae covering the process on retrolateral lobe. The differ-
ence lies in embolus length, much greater in A. lynnae sp. n. (Fig. 204) than in A. hirschii
(Fig. 176). Pairing female and immature to males of each one of these sympatric species
is a problem. Paratype female of A. hirschii has very long twisted spermatheca, which is
morphologically more compatible with the very long embolus of males of A. lynnae sp. n.
Immatures of A. hirschii were described as having single dorsal black stripe on abdomen
(Fig. 184). However, this immature pattern could fit to adults of either A. hirschii and
A. lynnae sp. n. Thus, it is necessary to further collect specimens to solve this query.

Avicularia caei sp. n.


http://zoobank.org/AE612016-7D21-48F6-8366-B289560ED3BC
Figs 19, 90, 210–218, 313

Diagnosis. males of A. caei sp. n. resemble those of A. minatrix, A. hirschii and A. lyn-
nae sp. n. by tibia I with discrete elevation covered by a cluster of setae in apical por-
tion, on prolateral side (Fig. 216). They can be distinguished from all species except A.
lynnae sp. n. by very long embolus, more than 4 times tegulum’s width in retrolateral
view (Fig. 211). Males of A. caei sp. n. can be distinguished from males of A. lynnae sp.
n. by weakly-developed prominence on tegulum (Fig. 212) and abdomen with lateral
dark stripes (Fig. 218). Female is unknown.
Etymology. This species is named after Carlos Eduardo Gurgel Paiola, aka Caê, in
honor to his continuous support to one of the authors (CSF). This name is masculine
in gender.
Material examined. Holotype male, Brazil, Pará, Juruti, Acampamento Mutum
(01°36'44.7"S, 56°11'39.2"W), L. T. Miglio col., 08 August 2008, ref. JURU 009
0043 (MPEG 15637).
Male. Description. MPEG 15637. Carapace: 10.44 long, 9.96 wide, 3.08 high.
Chelicera: 3.45 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 11.01, 5.79,
8.44, 8.21, 4.47, 37.92. II: 10.37, 5.18, 8.05, 8.30, 4.35, 36.25. III: 9.19, 4.34, 7.17,
7.92, 4.02, 32.64. IV: 10.85, 5.26, 10.26, 10.66, 4.59, 41.62. Palp: 6.39, 3.96, 5.29, –,
2.62, 18.26. Midwidths: femora I–IV= 2.08, 2.08, 2.44, 1.89, palp= 1.44; patellae I–
IV= 1.98, 2.10, 2.17, 1.90, palp= 1.48; tibiae I–IV= 1.25, 1.50, 1.47, 1.45, palp= 1.57;
metatarsi I–IV= 1.10, 1.46, 1.02, 1.01; tarsi I–IV= 1.15, 1.37, 1.33, 1.15, palp= 1.64.
Abdomen: 12.51 long, 7.59 wide. Spinnerets: PMS, 1.32 long, 0.63 wide, 0.19 apart;
PLS, 2.31 basal, 1.16 middle, 2.04 distal; midwidths 1.17, 0.93, 0.76, respectively.
118 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 210–217. Avicularia caei sp. n., male holotype (MPEG 015637). 210–213 right palpal bulb
(mirrored) 210 prolateral 211 retrolateral 212 frontal 213 dorsal 214 right cymbium, dorsal (mirrored)
215–217 right tibia I (mirrored) 215 prolateral 216 ventral 217 retrolateral. Scale bars = 1 mm.

Carapace: 1.05 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Fovea: shallow, recurve, 1.14 wide.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 119

Figure 218. Avicularia caei sp. n, holotype, habitus, preserved male (MPEG 015637). Detail: lateral
abdomen. Arrow indicating stripes. Photo: C. S. Fukushima.

Eyes: eye tubercle 0.76 high, 1.72 long, 2.16 wide. Clypeus absent. Anterior row
of eyes procurve, posterior slightly recurve. Eye size and interdistances: AME 0.50,
ALE 0.57, PME 0.20, PLE 0.49, AME–AME 0.37, AME–ALE 0.20, AME–PME
0.07, ALE–ALE 1.41, ALE–PME 0.34, PME–PME 1.35, PME–PLE 0.02, PLE–PLE
1.70, ALE–PLE 0.29, AME–PLE 0.28.
Maxilla: length to width: 2.14. Cuspules: about 101 spread over ventral inner heel.
Labium: 1.23 long, 1.65 wide, with 64 cuspules spaced by one diameter from each
other on anterior half. Labium sternal groove shallow, flat, with two slightly separate,
slim sigilla.
Chelicera: basal segment with 9 teeth and some small teeth on promargin. Ster-
num: 5.57 long, 4.49 wide. Sigilla: only posterior pair evident, oval, set at 45°angle,
one diameter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.10. Clavate trichobothria:
distal 1/2 tarsi I–II; 2/3 tarsus III, 1/2 tarsus IV. Scopula: Tarsi I–IV fully scopulate,
IV with few sparse setae. Metatarsi I–II scopulate in distal 3/4; III 1/2; IV 1/4 distal
scopulate. IV divided by row of setae.
Type II urticating setae: 0.99–1.05 long, 0.018–0.024 wide.
Palp (Figs 210–213, 313): globous bulb with small subtegulum and weakly-de-
veloped prominence on tegulum. Embolus: not flattened, lacking keels, 3.78 long in
retrolateral view, about 4 times tegulum’s length. Medial portion and tegulum’s mar-
gin form an acute angle in retrolateral view. Proximal part very curved in frontal view;
thin distal width, tapering distally; basal, middle, and distal width of 0.28, 0.15, 0.03,
respectively. Tegulum: 1.62 long, 0.96 high in retrolateral view. Cymbium subtrian-
gular with subequal lobes having well-developed rounded process bearing thin setae on
retrolateral lobe (Fig. 214).
Tibia I with discrete elevation covered by cluster of setae in apical portion, on
prolateral side (Figs 215–217).
120 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Color pattern (Fig. 218): carapace orange with golden brown short body setae,
thick longer setae spread over the carapace. Carapace border with long setae the same
color as dorsal carapace short body setae. Coxae, labium, sternum and maxillae light
brown, same color as ventral femora. Legs and palps with dark brown long guard-setae.
Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen dorsum with light
brown guard-setae, light brown short body setae forming lateral stripes, and black
short body setae forming a dark background. Ventral abdomen dark brown.
Female is unknown.
Color pattern ontogeny. Immatures are unknown.
Distribution. Brazil, known only from type locality (Fig. 90).
Natural history. Unknown. A single specimen was found walking on forest
ground (L. Miglio, pers. comm.).

Taxonomic remarks
Avicularia vestiaria (De Geer, 1778), nomen nudum

Aranea vestiaria De Geer, 1778: 313; F. O. Pickard-Cambridge 1896: 741 (under Avi-
cularia avicularia); Petrunkevitch 1911: 49 (under Avicularia avicularia); Roewer
1942: 253 (under Avicularia avicularia); Bonnet 1955: 828 (under Avicularia avi-
cularia); World Spider Catalog 2016 (under Avicularia avicularia).
Avicularia vestiaria: Ausserer 1871: 201 (as senior synonym of Aranea avicularia), 1875:
184; Hasselt 1888: 166 (as senior synonym of Mygale avicularia sensu Latreille).

Remarks. As stated by F. O. Pickard-Cambridge (1896), the name “vestiaria” is solely


a word used for De Geer (1778) in his redescription of Aranea avicularia Linnaeus
(1758) (Simon 1892: 172; F. O. Pickard-Cambridge 1896: 741), in litt. “Aranea ves-
tiaria hirsutissima nigro-fusca S. rufescens, plantis amplis tomentosis” (De Geer 1778:
313). It is not a specific name as considered by Ausserer (1871). Even though other
authors such as Ausserer (1875) considered it a valid name, we understand a formal de-
scription was not presented or intended and, therefore, Avicularia vestiaria (De Geer,
1778) is, herein, considered a nomen nudum.

Iridopelma leporina (C. L. Koch, 1841), comb. n., nomen dubium


Fig. 219

Mygale leporina C. L. Koch, 1841: 55 tab. CCCVI, fig. 726 (syntypes 4 dry, pinned
immatures, Bahia, Brazil, Freir leg., ZMB 2028, examined,); Ausserer 1871: 217;
F. O. Pickard-Cambridge 1896: 744. Syn. n.
Eurypelma leporina: C. L. Koch 1850: 74.
Avicularia leporina: Simon 1892: 172; Roewer 1942: 255; Bonnet 1955: 832; World
Spider Catalog 2016.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 121

Figures 219–224. Species formerly included in Avicularia. 219 syntype of Mygale leporina C. L. Koch,
1841, immature, from state of Bahia, Brazil, (ZMB 2028), habitus 220 syntype of Mygale plantaris
C. L. Koch, 1842, immature, from Brazil (ZMB 2026), habitus 221 Euathlus affinis (Nicolet, 1849)
comb. n., lectotype female, from Santiago, Chile (MNHN–AR 4871B), habitus 222 Avicularia arabica
(Strand, 1908) nomen dubium, holotype juvenile female, from El-Tor, Egypt (SMF 2660), spermathecae
223–224 Thrixopelma aymara (Chamberlin, 1916) comb. n., holotype female, from Aymara, Peru (MCZ
145) 223 spermatheca 224 habitus.

Remarks. The syntypes are specimens of subfamily Aviculariinae because they have
tarsi with well-developed scopulae, with a spatulated aspect (Fig. 219). They have an-
terior row of eyes procurved, spinnerets with the last article digitiform and no spines
on legs. The color pattern of the immatures and the type locality, Bahia (Brazil), match
with Iridopelma species. Thus, we decided to transfer Mygale leporina to Iridopelma,
making the new combination Iridopelma leporina (C. L. Koch, 1841) comb. n. How-
ever, as the specimens are immatures and in poor conditions we consider Iridopelma
leporina (C. L. Koch, 1841) comb. n. nomen dubium.
122 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Iridopelma plantaris (C. L. Koch, 1842), comb. n., nomen dubium


Fig. 220

Mygale plantaris C. L. Koch, 1842; 71 tab. CCCXII, fig. 736 (syntypes 2 immatures,
Brazil, ZMB 2026, examined). Syn. n.
Eurypelma plantaris: C. L. Koch 1850: 74.
Avicularia plantaris: Simon 1892: 172; World Spider Catalog 2016.

Remarks. C. L. Koch (1842) published the description of Mygale plantaris in the


same year of Mygale leporina. As M. leporina, the procurve anterior row of eyes, the
digitiform posterior article of spinnerets, the color pattern and type locality (Brazil) of
the syntypes match with Iridopelma (Fig. 220). Therefore, we transfer Mygale plantaris
to Iridopelma, making the new combination Iridopelma plantaris (C. L. Koch, 1842)
comb. n. Like M. leporina, the syntypes of M. plantaris are immature specimens.
Iridopelma species present very intense ontogenetic changes in color pattern, thus
different life stages present very distinct abdominal color pattern. This could be the
reason why C. L. Koch (1842) described both forms as two different species when they
could be the same. As we can’t assuredly affirm they are the same species, since the types
are not adults or even in good condition, we decided to keep them as separate species
and to consider Iridopelma plantaris (C. L. Koch, 1842) comb. n. nomen dubium.

Euathlus affinis (Nicolet, 1849), comb. n.


Figs 221, 225

Mygale affinis Nicolet, 1849: 333, pl. 1, fig. 6 (female lectotype and immature paralec-
totype designated, herein, from Chile, Santiago [33°28'S, 70°38'W]), MNHN–
AR 4871B, examined).
Brachypelma affinis: Mello-Leitão 1936d: 118; Bonnet 1955: 2989.
Eurypelma affine: Roewer 1942: 238.
Avicularia affinis: World Spider Catalog 2016.

Remarks. The lectotype female designated, herein, is a small specimen, with carapace
5.3 mm long (Fig. 221). It does not belong to any Aviculariinae species since it has
spines on the legs and lacks the characteristic developed scopula on tarsi. Its sper-
mathecae (Fig. 225) are compatible with Euathlus Ausserer, 1875 species (Perafán and
Pérez-Miles 2014), which have species recorded for the area where the specimen was
collected (Santiago, Chile). Furthermore, the original illustration published by Nicolet
(1849, plate 1) shows a spider with reddish coloration and lacking two urticating setae
patches, which is also compatible with Euathlus species, such as E. parvulus (Pocock,
1903) and Euathlus condorito Perafán & Pérez-Miles, 2014. Therefore, we transfer
Avicularia affinis (Nicolet, 1849) to Euathlus, resulting in Euathlus affinis (Nicolet,
1849) comb. n.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 123

Figure 225. Euathlus affinis (Nicolet, 1849) comb. n., lectotype (MNHN–AR 4871B), spermathecae.

Ischnocolus hirsutum Ausserer, 1875, nomen dubium

Ischnocolus hirsutus Ausserer, 1875: 170 (syntypes 11 small specimens from Cuba,
BMNH 1890.7.1.347, and 2 small specimens from Bogota, Colombia [4°35'N,
74°04'W], in another vial, BMNH 1890.7.1.339+347, examined).
Eurypelma hirsutum: Roewer 1942: 240.
Avicularia hirsuta: World Spider Catalog 2016.

Remarks. Ausserer (1875) mentioned several females from Cuba and two specimens
from “Santa Fé de Bogotá” (known, nowadays, as Bogota, capital of Colombia) in
Keyserling collection (housed at the BMNH). All examined specimens from Cuba have
urticating setae type III on abdomen dorsum, and all specimens from Bogota have setae
type I. Therefore, they belong to subfamily Theraphosinae and it is, herein, removed
from the genus Avicularia. All syntypes are immatures lacking diagnostic characters
and, therefore, we consider Ischnocolus hirsutum Ausserer, 1875 a nomen dubium.

Avicularia metallica Ausserer, 1875, nomen dubium

Avicularia metallica Ausserer, 1875: 185; Simon 1892: 171; Strand 1907b: 92; Mello-
Leitão 1923: 377; Roewer 1942: 255; Bonnet 1955: 832; Petrunkevitch 1911: 50;
1939: 287; World Spider Catalog 2016.

Remarks. The type was not found in Vienna collection where it should be deposited accord-
ing to description. Thus, we consider Avicularia metallica Ausserer, 1875 as nomen dubium.

Ischnocolus gracilis Keyserling, 1891, nomen dubium

Ischnocolus gracilis Keyserling, 1891: 11 (holotype immature male, Brazil, Rio de Janeiro,
Monte Verde [21°27'S, 41°55'W], BMNH 90.7.1.346, examined).
Eurypelma gracile: Roewer 1942: 239.
Avicularia gracilis: World Spider Catalog 2016.
124 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Remarks. The specimen is very small, the epigastric area is dissected, but there are
no spermathecae. Abdomen dorsum is bald. Since the description mentions spines
on posterior articles, we, herein, remove the species from the genus Avicularia. As the
specimen is immature, damaged, and lacking diagnostic characters, therefore we con-
sider, herein, Ischnocolus gracilis Keyserling, 1891 a nomen dubium.

Grammostola subvulpina (Strand, 1906), comb. n.

Avicularia subvulpina Strand, 1906a: 22 (holotype male, no further information,


MWNH 362, examined by photo); 1907j: 233; Mello-Leitão 1923: 377; Petrunk-
evitch 1939: 288; Roewer 1942: 254; Bonnet 1955: 833; World Spider Catalog
2016.

Remarks. A specimen that fits Strand’s (1906a) description was found in Wiesbaden
collection and is, herein, considered as the holotype. The male resembles a charac-
teristic Grammostola Simon, 1892 species from Brazil. Thus, we transfer Avicularia
subvulpina to Grammostola, making the new combination Grammostola subvulpina
(Strand, 1906) comb. n.

Avicularia arabica (Strand, 1908), nomen dubium


Fig. 222

Avicuscodra arabica Strand, 1908: 771 (holotype female, Egypt, El-Tor [28°14'N,
33°37'E] (Tor, Arabien [sic]), Rüppell, SMF 2660, examined); 1916: 20; Roewer
1942: 256.
Chaetopelma arabica: Raven 1985: 149.
Avicularia arabica: Gallon 2008: 243, figs 39–41; World Spider Catalog 2016.

Remarks. The holotype of Avicuscodra arabica was recently rediscovered by Gallon


(2008) in SMF. The specimen, supposedly from Tor, Egypt, is clearly an Avicularia as
proposed by Gallon (2008). It has two spermathecae lacking lobes and with accentu-
ated outwards curvature medially. Urticating setae of type II are present on abdomen
dorsum. These characters are present in Avicularia species, which are restricted to Cen-
tral and South America. As Gallon (2008) stated, the presence of a tropical American
specimen together with Arabian material is most likely a mistake made by a nonspe-
cialist before Strand’s examination.
Even though it is possible to identify the specimen as belonging to the genus Avicu-
laria, it is a small female in poor condition, without evident color pattern and no other
diagnostic characteristic. It is not possible to assure the specimen identity since spermathe-
cae morphology is similar in most Avicularia species (Fig. 222). Furthermore, there is no
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 125

indication of where the specimen came from. Thus, Avicularia arabica (Strand, 1908) is,
herein, considered nomen dubium.

Thrixopelma aymara (Chamberlin, 1916), comb. n.


Figs 223–224

Eurypelma aymara Chamberlin, 1916: 201 (holotype female, Peru, Aymara, Dr. W. H.
Jones col., MCZ 145, examined); Bonnet 1955: 1831.
Eurypelma aymarum: Roewer 1942: 238.
Avicularia aymara: World Spider Catalog 2016.

Remarks. The specimen is not an aviculariine since it has no spatulate scopulae, bears
two rounded spermathecae, and has no type II urticating setae (Fig. 224). The type
locality, spermathecae morphology (Fig. 223) and presence of urticating setae type
III match with the diagnostic characteristics of Thrixopelma species. Thus, we transfer
Eurypelma aymara to Thrixopelma, making the new combination Thrixopelma aymara
(Chamberlin, 1916) comb. n.

Avicularia aurantiaca Bauer, 1996, nomen dubium

Avicularia aurantiaca Bauer, 1996: 2, figs 1–4; World Spider Catalog 2016.

Material examined. female spermathecae exuvia in microscope slide, Peru, G. Schmidt


ded. (SMF 58271-84).
Remarks. The original description (Bauer 1996) stated he intended to deposit a female
specimen at SMF as holotype, a slide prepared with spemathecae from an exuvia and a para-
type male. The holotype and paratype specimen were never deposited in SMF collection.
The description states female has leg IV longer than leg I, spermathecae with midwidth not
expanded and orange leg rings (explaining why it was called “aurantiaca”). Thus, A. auran-
tiaca is likely to be a junior synonym of A. rufa. However, as there is no holotype and the
description does not allow for a precise identification and characterization of the species, the
name Avicularia aurantiaca Bauer, 1996 is considered, herein, as nomen dubium.
Types not found. The following types have been unsuccessfully searched for within
many arachnological collections and their descriptions do not allow for reliable identifica-
tion. Thus, we consider the following species all nomina dubia: Araneus hirtipes Fabricius,
1793 nomen dubium; Avicularia testacea (C. L. Koch, 1841) nomen dubium; Avicularia de-
trita (C. L. Koch, 1842) nomen dubium; Avicularia hirsutissima (C. L. Koch, 1842) nomen
dubium; Avicularia holmbergi Thorell, 1890 nomen dubium; Ischnocolus doleschalli Ausserer,
1871 nomen dubium; Avicularia rapax (Ausserer 1875) nomen dubium; Avicularia ochracea
(Perty, 1833) nomen dubium; and Avicularia walckenaerii (Perty, 1833) nomen dubium.
126 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Araneus hirtipes (Fabricius, 1787), nomen dubium

Aranea hirtipes Fabricius, 1787: 346, 1793: 428; Latreille 1806: 83 (under Mygale avicu-
laria); Hahn 1833: 10 (under Mygale avicularia); Walckenaer 1837: 212 (under
Mygale versicolor); World Spider Catalog 2016 (under Avicularia versicolor).
Mygale hirtipes: C. L. Koch 1836: 38.
Eurypelma hirtipes: C. L. Koch 1850: 74.
Avicularia hirtipes: Bonnet 1955: 831.

Remarks. Considered synonym of Mygale avicularia by Latreille (1806) and Hahn


(1833) as well as of Mygale versicolor by Walckenaer (1837). Although very uninforma-
tive, the description shows an eye disposition that suggests this species is not a mygalo-
morph spider. Thus, we considered Araneus hirtipes (Fabricius, 1787) nomen dubium.

Avicularia testacea (C. L. Koch, 1841), nomen dubium

Mygale testacea C. L. Koch, 1841: 45, Tab. CCCIII, fig. 719; F. O. Pickard-Cambridge
1896: 741 (under A. avicularia); World Spider Catalog 2016 (under A. avicularia).

Remarks. F. O. Pickard-Cambridge (1896) considered it as a probable synonym of


A. avicularia. The specimen drawned by C. L. Koch (1841) resembles a specimen of
Avicularia since it has well-developed scopula, anterior row of eyes procurve and long
and curved palpal embolus. However, it is not possible to establish its identity without
analyze the type. Therefore, herein, we removed it from the synonymy with A. avicu-
laria and consider Avicularia testacea (C. L. Koch, 1841) nomen dubium.

Avicularia detrita (C. L. Koch, 1842), nomen dubium

Mygale detrita C. L. Koch, 1842: 86, Tab. CCCXVIII, fig. 746; F. O. Pickard-Cam-
bridge 1896: 744; Petrunkevitch 1911: 49; 1939: 286; Mello-Leitão 1923: 377;
Roewer 1942: 254.
Avicularia detrita: Simon 1903: 958; Bonnet 1955: 830; World Spider Catalog 2016.

Remarks. Analyzing Koch’s drawing, it is possible to note the absence of spatulated scopula,
typical of Aviculariinae. Male palpal bulb also seems different from Aviculariinae species.
Thus, it is probable that this species belongs to another subfamily, such as Theraphosinae.

Avicularia hirsutissima (C. L. Koch, 1842), nomen dubium

Mygale hirsutissima C. L. Koch, 1842: 76, fig. 738; Petrunkevitch 1911: 49 (under
Avicularia avicularia); Roewer 1942: 254 (under Avicularia avicularia); Bonnet
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 127

1955: 829 (under Avicularia avicularia); World Spider Catalog 2016 (under Avicu-
laria avicularia).

Remarks. The drawn probably represents a specimen of Avicularia since it has well-
developed scopula and the anterior row of eyes procurve. However, it is not possible to
establish its identity without examining the type. Herein, we removed it from the syn-
onymy with A. avicularia established by Petrunkevitch (1911) and we consider Avicu-
laria hirsutissima (C. L. Koch, 1842) nomen dubium.

Avicularia holmbergi Thorell, 1890, nomen dubium

Avicularia holmbergi Thorell, 1890: 399; F. O. Pickard-Cambridge 1896: 744; Simon


1892: 170; Petrunkevitch 1911: 50; 1939: 287; Mello-Leitão 1923: 377; Roewer
1942: 254; Bonnet 1955: 831; World Spider Catalog 2016.

Remarks. This species was described in a work on the Indonesian fauna, but type lo-
cality was considered uncertain (Thorell 1890).

We consulted the curator of Arachnoidea Collection in the Natural History Museum


of Vienna, where Ausserer’s specimens below were likely deposited. Unfortunately, the
types are not there and descriptions do not allow reliable identification of the species.

Ischnocolus doleschalli Ausserer, 1871, nomen dubium

Ischnocolus doleschalli Ausserer, 1871: 189.


Eurypelma doleschalli: Roewer 1942: 239.
Avicularia doleschalli: World Spider Catalog 2016.

Remarks. Ausserer (1871) renamed as Ischnocolus doleschalli the species Theraphosa


bicolor Doleschal (probably from Brazil), and added to Doleschal’s description that
the specimen has spines on tibiae and metatarsi. Thus, it is not an Aviculariinae and,
herein, we consider Ischnocolus doleschalli Ausserer, 1871 a nomen dubium.

Avicularia rapax (Ausserer, 1875), nomen dubium

Eurypelma rapax Ausserer, 1875: 200, pl. 7, fig. 45; Petrunkevitch 1911: 63; 1939:
264; Roewer 1942: 241; Bonnet 1955: 1833.
Avicularia rapax: World Spider Catalog 2016.

Remarks. Ausserer (1875) described a male with two-branched tibial apophysis, and
having setae on palpal tibia. It resembles a theraphosine specimen, but Ausserer did
128 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

not give information on diagnostic characters. Thus, we consider Avicularia rapax


(Ausserer, 1875) a nomen dubium.

We failed to find holotypes of both Perty’s species below. However, Perty (1833) drew
in detail the legs of these species, showing well-developed scopulae on tarsi and meta-
tarsi, with the spatulated aspect characteristic of aviculariines.

Avicularia ochracea (Perty, 1833), nomen dubium

Mygale ochracea Perty, 1833: 191, pl. XXXVIII, fig. 2; Walckenaer 1837: 215; C. L.
Koch 1841: 48, pl. CCCIV, fig. 721.
Eurypelma ochracea: Ausserer 1871: 215; C. L. Koch: 1842: 48; 1850: 74; Petrunk-
evitch 1911: 63; 1939: 264.
Eurypelma ochraceum: Roewer 1942: 240; Bonnet 1955: 1832.
Avicularia ochracea: World Spider Catalog 2016.

Remarks. Mygale ochracea was described as having an overall ocher coloration and a
reddish abdomen. The specimen was found around Rio Negro River, in the province of
the same name. At this time, the Rio Negro Province comprised the cities Barcelos and
Vila da Barra do Rio Negro, now the city of Manaus, capital of the state of Amazonas
(Instituto Histório e Geográfico Brasileiro in Spix and Martius 1823). The aviculariine
species that fit Perty’s illustration and that occur in that area are A. avicularia and A. ju-
ruensis. However, based in Perty’s description it is not possible to determine A. ochracea
identity. Thus, we considered Avicularia ochracea (Perty, 1833) nomen dubium.

Avicularia walckenaerii (Perty, 1833), nomen dubium

Mygale walckenaerii Perty, 1833: 191, pl. XXXVIII, fig. 3; Walckenaer 1837: 217; C.
L. Koch 1841: 46, pl. CCCIV, fig. 720.
Avicularia walckenaeri: Simon 1892: 171; Petrunkevitch 1911: 766; 1939: 288; Mello-
Leitão 1923: 377; World Spider Catalog 2016.

Remarks. Mygale walckenaerii was found in equatorial Brazil by Spix and Martius. The
equatorial part of Brazil they explored were the states of Amazonas, Maranhão and Pará.
Mygale walckenaerii was described as having an overall dark coloration, with greyish se-
tae covering its body and “scarlet feet” (Perty 1833), in reference to vivid reddish rings
on tarsi, metatarsi and tibiae. Based on the description (Perty 1833) and considering
the probable occurrence area, the specimen depicted by Perty could be an A. rufa or A.
variegata stat. n. As it is not possible to positively identify the species described by Perty
(1833), we considered Avicularia walckenaerii (Perty, 1833) nomen dubium.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 129

The holotypes of the species described by Tesmoingt (1996abc; 1999ab) are not at
the Musée d’Histoire Naturelle de Lille, where they should be deposited according to
descriptions. We contacted the author and he said the types have never been deposited in
any collection. As it is not possible to be sure about the identity of this species only by the
information contained in the descriptions, we consider Avicularia azuraklaasi Tesmo-
ingt, 1996 nomen dubium, Avicularia braunshauseni Tesmoingt, 1999 nomen dubium,
Avicularia geroldi Tesmoingt, 1999 nomen dubium, Avicularia huriana Tesmoingt, 1996
nomen dubium, and Avicularia ulrichea Tesmoingt, 1996 nomen dubium.

Avicularia azuraklaasi Tesmoingt, 1996, nomen dubium

Avicularia azuraklaasi Tesmoingt, 1996a: 8, figs 1–8; World Spider Catalog 2016.

Avicularia braunshauseni Tesmoingt, 1999, nomen dubium

Avicularia braunshauseni Tesmoingt, 1999a: 14, figs 1–5; World Spider Catalog 2016.

Avicularia geroldi Tesmoingt, 1999, nomen dubium

Avicularia geroldi Tesmoingt, 1999b: 17, figs 1–8; World Spider Catalog 2016.

Avicularia huriana Tesmoingt, 1996, nomen dubium

Avicularia huriana Tesmoingt, 1996b: 6, figs 1–18; 1996c: 2, figs 1A, 2–7, Bullmer et
al. 2006: 7, figs 11–14; World Spider Catalog 2016.

Avicularia ulrichea Tesmoingt, 1996, nomen dubium

Avicularia sp. Tesmoingt, 1996b: 6, fig. unnumbered.


Avicularia ulrichea Tesmoingt, 1996c: 2, fig. 1B; World Spider Catalog 2016.

Types considered lost, herein.

Avicularia soratae Strand, 1907, nomen dubium

Avicularia soratae Strand, 1907d: 556; 1907e: 260; Petrunkevitch 1911: 50; 1939:
288; Mello-Leitão 1923: 377; Roewer 1942: 255; World Spider Catalog 2016.
Avicularia soratensis: Bonnet 1955: 832.
130 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Remarks. Strand (1907e) indicated the type is deposited in Lübeck Museum. How-
ever, the former collections of the old museum were destroyed in 1942 during World
War II (S. Füting, pers. comm.). Thus, the type is lost and Avicularia soratae Strand,
1907 is considered, herein, as nomen dubium.
The three species below were deposited in Staatliches Museum für Naturkunde
Stuttgart (SMF). Its arachnological collection was destroyed in World War II and
contained at least 169 types of Strand, 9 types of C. Koch and 3 types of Keyserling
(Renner 1988). The names Avicularia fasciculata Strand, 1907, Avicularia fasciculata
clara Strand, 1907 and Avicularia surinamensis Strand, 1907 are, herein, considered as
nomina dubia.

Avicularia fasciculata Strand, 1907, nomen dubium

Avicularia fasciculata Strand, 1907b: 92; 1907c: 465; Mello-Leitão 1923: 377; Petrunkevitch
1939: 286; Roewer 1942: 254; Bonnet 1955: 831; World Spider Catalog 2016.

Avicularia fasciculata clara Strand, 1907, nomen dubium

Avicularia fasciculata clara Strand, 1907c: 467; Roewer 1942: 254; World Spider Catalog
2016.

Avicularia surinamensis Strand, 1907, nomen dubium

Avicularia surinamensis Strand, 1907b: 90; 1907b: 465; Mello-Leitão 1923: 377;
Petrunkevitch 1939: 288; Roewer 1942: 255; Bonnet 1955: 833, World Spider
Catalog 2016.

Note. The Avicularia species considered nomina dubia and nomina nuda by World
Spider Catalog (2016) do not have their status changed.

Caribena gen. n.
http://zoobank.org/45A06052-FAED-48FB-8307-72E3A74048ED

Diagnosis. Females and males differ from those of all Aviculariinae genera by having type
II urticating setae longer (more than 1 mm long) and slender (less than 0.009 mm wide)
(Figs 18, 243) on a conspicuous patch on abdomen dorsum (Figs 241, 254). Additionally,
males have a sharp spiniform process in the retrolateral lobe of cymbium (Fig. 306).
Etymology. From the Spanish word “caribeña”, meaning “from Caribbean”, the
occurrence area of this genus, and is considered feminine in gender.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 131

Description. Carapace longer than wide, cephalic region slightly raised. Cephalic
and thoracic striae inconspicuous or evident. Fovea deep, straight or recurve. Cheli-
cera without rastelum. Eye tubercle distinct, slightly raised to raised, wider than long.
Anterior row of eyes procurve, posterior slightly recurve. Clypeus narrow or absent.
Labium subquadrate, longer than wide, with 95–115 cuspules spaced by one diameter
from each other on anterior third center. Maxillary lyra absent. Maxilla subretangular,
anterior lobe distinctly produced into conical process, inner angle bearing 100–200
cuspules. Sternum longer than wide, with posterior angle acute, not separating coxae
IV. Anterior pair of sigilla not evident, middle fusiform, posterior rounded or ellipsoi-
dal; all less than one diameter from margin. Leg formula: I=IV II III. Clavate trichobo-
thria on distal 2/3 or 1/2 of tarsi I–IV. Tarsi I–IV fully scopulate, IV divided or not by
a band of setae. Metatarsi I–II fully scopulate, III 1/2 to 2/3 distal scopulate, IV 1/2 to
1/3 distal scopulate. Metatarsi IV divided by row of setae. Scopulae of tarsi and meta-
tarsi I–II extended very laterally giving them spatulate appearance. Femora IV with-
out retrolateral scopulae. Stridulatory setae absent. Leg aspinose. ITC absent; STC
without denticles. Posterior lateral spinneret distally elongating, digitiform. Type II
urticating setae very slender on dorsal abdomen, with barbs along entire length in both
sexes (Figs 18, 243). Tibial apophysis single branch on prolateral leg I, with weakly-
developed base and grouped spiniform setae distally (Fig. 310). Tibial apophysis on leg
II absent. Male metatarsus I touches retrolaterally tibial apophysis’ setae when folded.
Globous bulb with small subtegulum, lacking prominence (C. laeta comb. n., Fig.
233) or with developed prominence on tegulum (C. versicolor comb. nov., Fig. 246).
Embolus not flattened, lacking keels, about 3.0 to 3.5 times tegulum’s length. Embo-
lus medial portion and tegulum’s margin forming an obtuse (C. laeta comb. n., Fig.
232) or an acute angle (C. versicolor comb. n., Fig. 245) in retrolateral view. Embolus
proximal part slightly curved (C. laeta comb. n., Fig. 233) or very curved (C. versicolor
comb. n., Fig. 246) in frontal view; thin distal width, tapering distally (Figs 234; 318).
Cymbium subtriangular with subequal lobes, having well-developed sharp process on
retrolateral lobe, bearing thin setae (Figs 235, 306). Two completely separated, not-
twisted, short (C. laeta comb. n., Fig. 227) or long spermathecae (C. versicolor comb.
n., Fig. 230); with apex with intumescence (C. laeta comb. n., Fig. 227) or lacking
it (C. versicolor comb. n., Fig. 230). Spermathecae almost straight (C. laeta comb. n.,
Fig. 227) or with an accentuated outwards curvature medially (C. versicolor comb.
n., Fig. 230). Spermatheca midwidth as wide as its base width and weakly-sclerotized
area shorter than half the length of well-sclerotized area. Drastic ontogenetic changes
on abdominal color pattern. Juveniles have green metallic sheen, no black tarsi (Figs
240, 252) and abdomen with dorsal central longitudinal black stripe connected (C.
versicolor comb. n., Fig. 252) or not (C. laeta comb. n., Fig. 240) with conspicuous
transversal black stripes. When mature, they lose this pattern.
Type species. Mygale laeta C. L. Koch, 1842, herein, designated.
Species included. Caribena laeta (C. L. Koch, 1842) comb. n. and Caribena ver-
sicolor (Walckenaer, 1837) comb. n.
Distribution. Puerto Rico, the U. S. Virgin Island and Martinique (Fig. 226).
132 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 226. Map showing records of Caribena gen. n. and Antillena gen. n. ? = uncertain record.

Key to males

1 Embolus with proximal part slightly curved in frontal view (Fig. 233), medial
portion and tegulum’s margin form an obtuse angle in retrolateral view (Fig.
232); tegulum lacking prominence (Fig. 233).................... C. laeta comb. n.
– Embolus with proximal part very curved in frontal view (Fig. 246), medial por-
tion and tegulum’s margin form an acute angle in retrolateral view (Fig. 245);
tegulum with developed prominence (Fig. 246).............. C. versicolor comb. n.

Key to females

1 Spermathecae almost straight, with intumescence on its apex (Fig. 227)........


.......................................................................................... C. laeta comb. n.
– Spermathecae with accentuated outwards curvature medially, lacking intu-
mescence on its apex (Fig. 230)..................................C. versicolor comb. n.

Caribena laeta (C. L. Koch, 1842), comb. n.


Figs 226–229, 231–243, 310

Mygale laeta C. L. Koch, 1842: 66, pl. CCCX, fig. 732 (lectotype female and paralec-
totype immature, here designated, Puerto Rico, Moritz leg., ZMB 2045, exam-
ined); Ausserer 1871: 217; F. O. Pickard-Cambridge 1896: 744.
Mygale caesia C. L. Koch, 1842: 83, tab. CCCXVII, fig. 744 (syntypes 4 immatures,
Puerto Rico, ZMB 2034, examined); F. O. Pickard-Cambridge 1896: 744. Syn. n.
Avicularia caesia: Simon 1892: 171; Petrunkevitch 1911: 49, 1929: 20, 51, figs 32–35,
1939: 285; Roewer 1942: 254; Bonnet 1955: 830; World Spider Catalog 2016.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 133

Avicularia laeta: F. O. Pickard-Cambridge 1896: 744; Petrunkevitch 1911: 50, 1929:


20, 31, 48, fig. 31, 1939: 287; Mello-Leitão 1923: 377; Roewer 1942: 255; Bonnet
1955: 832; World Spider Catalog 2016.

Diagnosis. Females can be distinguished from females of C. versicolor comb. n. by al-


most straight spermathecae with intumescence on their apex (Figs 227–229). Males dif-
fer from males of C. versicolor comb. n. by lacking prominence on tegulum, by having
embolus medial portion and tegulum’s margin forming an obtuse angle in retrolateral
view and by slightly curved shape of basal part of embolus in frontal view (Fig. 233).
Material examined. Puerto Rico, Corozal [18°18'N, 66°19'W], 1 female, M.
Martinez col., 19 September 1994 (MNRJ 06909); Puerto Rico. Bayamon [18°22'N,
66°09'W], 1 male, 19 November 1932 (AMNH PR).
Additional material. CUBA: 1 female (ZMB 30732); PUERTO RICO: 1 fe-
male (ZMB 30757); Road 10, km 21, 1 female, V. Roth col., 12 July 1977, in foliage
of Cecropia tree (AMNH RW05); Aibonito [18°08'N, 66°15'W], 1 juvenile female,
1–3 June 1915, ref. 3553 (AMNH PR); 1 immature male (AMNH 1.15); Bayamon
[18°22'N, 66°09'W], 1 male, T. Fuentes col., 12 September 1989 (AMNH RW04);
Corozal [18°18'N, 66°19'W], 1 male, M. Martinez col., 26 September 1994 (MNRJ
06908); 1 male, same collector, 17 September 1991 (MNRJ 06906); 1 male, same
collector, 21 September 1991 (MNRJ 06907); Mayaguez [18°12'N, 67°08'W], Tela:
Ninguna, 1 female, A. Ruiz col., 15 February 1962 (AMNH PR); Rio Grande, El
Yunque [18°20'N, 65°45'W], 1 female, J. Coffey col., 11 October 1993, in silk retreat
in bromeliad (AMNH RW03); 1 male, north slope of Turquillo mountains near El
Yunque National Forest [18°17'N, 65°48'W], A. Sanchez col., 29 September 1997
(MNRJ 06905); VIRGIN ISLANDS: Saint John [18°20'N, 64°47'W], 1 female, A.
Sanchez col., April 1999, in silken retreat in bromeliad (AMNH RW02); Cinnamon
Bay [18°21'N, 64°45'W] trail, about 700 m, 1 immature male, R. Zueifel col., in for-
est, in a bark, during night, 9 July 1980 (AMNH VI); 1 male, 1 female, Island Project
staff col., 18 December 1965 (AMNH VI).
Remarks. The syntypic series of Mygale laeta is constituted by specimens poorly
conserved: a specimen without spermatheca (thereafter considered immature) and fe-
male with characteristic spermathecae, as reason why it was chosen to be lectotype.
The syntypic series of Mygale caesia is composed of four very small specimens, with
central dark longitudinal stripe and dark transversal stripes on each side of abdomen.
They retain the green-blue sheen illustrated on the drawings of C. L. Koch (1842),
a pattern characteristic of juveniles from many aviculariines. C. L. Koch (1842) de-
scribed these specimens as being a second species of Mygale in Puerto Rico (the first
one was M. laeta). However, it is very probable that both are the same species as it was
not found any other aviculariine species in Puerto Rico and these immatures are very
similar to C. laeta comb. n. immatures. Thus, we consider M. caesia as the immature
stage of M. laeta. Since both species were described in the same paper, by page priority
Mygale caesia C. L. Koch, 1842 is, herein, considered junior synonym of M. laeta C.
L. Koch, 1842 new synonymy.
134 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 227–230. Caribena gen. n., spermathecae 227–229 Caribena laeta (C. L. Koch, 1842) comb.
n., spermathecae variation. 227 Corozal, Puerto Rico (MNRJ 06909) 228 syntype, Puerto Rico (ZMB
2045) 229 Saint John, Virgin Islands (AMNH VI) 230 Caribena versicolor (Walckenaer, 1837) comb.
n., Martinique (MNHN–AR 4904). Scale bars = 1 mm.

Female. Redescription. AMNH RCW. Carapace: 16.98 long, 14.82 wide, 5.01
high. Chelicera: 7.55 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I:
13.84, 8.48, 11.16, 9.24, 5.44, 48.16. II: 12.68, 7.52, 9.63, 8.80, 4.84, 43.47. III:
11.87, 6.85, 8.75, 8.73, 4.51, 40.71. IV: 13.88, 7.54, 11.18, 10.89, 3.92, 47.41. Palp:
9.55, 5.71, 6.27, –, 5.97, 27.50. Midwidths: femora I–IV= 3.13, 2.62, 3.08, 3.08,
palp= 2.58; patellae I–IV= 3.16, 2.99, 3.18, 3.01, palp= 2.50; tibiae I–IV= 2.45, 2.56,
2.43, 2.73, palp= 2.01; metatarsi I–IV= 1.98, 1.94, 1.89, 1.87; tarsi I–IV= 2.24, 2.07,
2.14, 2.15, palp= 2.28. Abdomen: 23.50 long, 17.64 wide. Spinnerets: PMS, 1.73
long, 0.90 wide, 0.43 apart; PLS, 1.87 basal, 1.67 middle, 2.80 distal; midwidths 1.30,
1.17, 0.76, respectively.
Carapace: 1.15 times longer than wide; cephalic region not raised, thoracic striae
inconspicuous.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 135

Figures 231–238. Caribena laeta (C. L. Koch, 1842) comb. n., male (AMNH PR). 231–234 left palpal
bulb 231 prolateral 232 retrolateral 233 frontal 234 dorsal 235 left cymbium, dorsal 236–238 left tibial
apophysis of leg I 236 prolateral 237 ventral 238 retrolateral. Scale bars = 1 mm.

Fovea: deep, straight, 2.06 wide.


Eyes: eye tubercle 1.07 high, 2.43 long, 3.51 wide. Clypeus 0.20. Anterior row of
eyes procurve, posterior row slightly recurve. Eye size and interdistances: AME 0.80,
136 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

ALE 0.86, PME 0.39, PLE 0.65, AME–AME 0.60, AME–ALE 0.51, AME–PME
0.17, ALE–ALE 2.00, ALE–PME 0.96, PME–PME 1.86, PME–PLE 0.28, PLE–PLE
2.38, ALE–PLE 0.49, AME–PLE 0.57.
Maxilla: length to width: 1.68. Cuspules: 100–200 spread over ventral inner heel.
Labium: 2.28 long, 2.73 wide, with 97 cuspules spaced by one diameter from each
other on anterior third. Labio-sternal groove shallow, flat, sigilla not evident.
Chelicera: basal segment with 12 teeth and some small teeth on promargin. Ster-
num: 8.34 long, 6.71 wide. Sigilla: anterior pair not evident, median fusiform, poste-
rior elipsoidal, both less than one diameter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 0.98. Clavate trichobothria:
distal 2/3 tarsi I–IV. Scopula: Tarsi I–IV fully scopulate. IV divided by row of setae.
Metatarsi I–II fully scopulate; III–IV 1/2 distal scopulate. IV divided by row of setae.
Type II urticating setae (Fig. 243): 1.02–1.33 long, 0.006–0.009 wide.
Spermathecae (Fig. 227): two short, almost straight spermathecae bearing intu-
mescence (not a well-defined lobe) in their apex. Midwidth as wide as its base width
and weakly-sclerotized area shorter than half the length of well-sclerotized area.
Color pattern (Fig. 241): carapace brown with golden short body setae. Carapace
border long setae the same color as dorsal carapace short body setae. Coxae, labium,
sternum and maxillae brown, slightly darker than ventral femora. Legs and palps with
golden short body setae with green sheen and dark brown long guard-setae, gradually
lightening from base to tip. Leg rings on distal femora, tibiae and metatarsi whitish.
Tarsi with an “U” shaped orange stripe. Abdomen with golden brown guard-setae
homogeneously distributed and golden short body setae. Ventral abdomen brown.
Urticating setae form very distinctive small golden patch on dorso posterior area
of abdomen.
Male. Description. AMNH. Carapace: 10.22 long, 8.86 wide, 3.05 high. Cheli-
cera: 2.96 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 11.12, 5.80,
9.30, 8.75, 4.43, 39.40. II: 10.20, 5.03, 8.86, 8.01, 4.77, 36.87. III: 9.20, 4.14, 7.14,
7.91, 4.06, 32.45. IV: 10.57, 4.85, 9.54, 9.48, 4.30, 38.74. Palp: 6.68, 3.90, 5.01, –,
1.76, 17.35. Midwidths: femora I–IV= 1.94, 1.80, 1.77, 1.90, palp= 1.53; patellae I–
IV= 1.69, 1.75, 1.72, 1.70, palp= 1.38; tibiae I–IV= 1.53, 1.35, 1.41, 1.40, palp= 1.42;
metatarsi I–IV= 1.16, 1.01, 1.02, 1.08; tarsi I–IV= 1.16, 1.07, 1.27, 1.27, palp= 1.29.
Abdomen: 9.79 long, 6.92 wide. Spinnerets: PMS, 0.92 long, 0.41 wide, 0.18 apart;
PLS, 1.47 basal, 0.80 middle, 1.56 distal; midwidths 0.78, 0.68, 0.48, respectively.
As in female, except:
Carapace: 1.15 times longer than wide.
Fovea: 1.02 wide.
Eyes: eye tubercle 0.73 high, 1.65 long, 2.16 wide. Clypeus absent. Eye size and
interdistances: AME 0.61, ALE 0.61, PME 0.27, PLE 0.48, AME–AME 0.31, AME–
ALE 0.25, AME–PME 0.11, ALE–ALE 1.27, ALE–PME 0.72, PME–PME 1.17,
PME–PLE 0.07, PLE–PLE 1.67, ALE–PLE 0.24, AME–PLE 0.37.
Maxilla: length to width: 1.90. Cuspules: 174. Labium: 1.22 long, 1.45 wide, with
112 cuspules spaced by one diameter from each other on anterior half.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 137

Figures 239–242. Caribena laeta (C. L. Koch, 1842) comb. n., Puerto Rico, habitus. 239 immature
240 juvenile 241 female 242 male. Arrow: Urticating setae type II on a conspicuous patch on dorsal
abdomen. Photos: R. C. West.

Chelicera: basal segment with 8 teeth and some small teeth on promargin. Ster-
num: 4.94 long, 3.73 wide. Sigilla: three pairs, posterior rounded, less than one diam-
eter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 0.98. Metatarsus III scopulate
in distal 2/3; IV, in distal 1/2.
Type II urticating setae: 1.41–1.71 long and 0.007–0.009 wide.
Palp (Figs 231–234): globous bulb with small subtegulum, lacking prominence on
tegulum. Embolus: not flattened, lacking keels, 2.32 long in retrolateral view, about 3
times tegulum’s length. Medial portion and tegulum’s margin form an obtuse angle in
retrolateral view. Proximal part slightly curved in frontal view; thin distal width, taper-
ing distally; basal, middle, and distal width of 0.44, 0.12, 0.04, respectively. Tegulum:
1.25 long, 0.74 high in retrolateral view. Cymbium subtriangular with subequal lobes,
having well-developed sharp process on retrolateral lobe, bearing thin setae (Fig. 235).
Tibial apophysis (Figs 236–238, 310): single branch on prolateral leg I, with weak-
ly-developed base and grouped spiniform setae distally. Male metatarsus I touches
retrolaterally tibial apophysis’ setae when folded.
Color pattern ontogeny. Juveniles with metallic sheen, all articles with same
blackish color (Fig. 240), and abdomen dorsum with dorsal central longitudinal black
138 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 243. Caribena laeta (C. L. Koch, 1842) comb. n., SEM microphotographs of slender urticating setae
type II, with barbs along all its length. Detail: region near stalk, female (MNRJ 06909). Scale bar = 100 μm.

stripe disconnected from transversal black stripes. When mature, they lose this pattern
(Figs 241–242).
Distribution. Puerto Rico and the U. S. Virgin Island (Lesser Antilles) (Fig. 226).
There is a dubious record of a single specimen from Cuba lacking any information on
collectors and date.
Natural history. Petrunkevitch (1929) mention that three females captured in
Puerto Rico were found on retreats inside bromeliad leaves. The author also mentioned
retreats of this species are similar to retreats of Avicularia species, consisting of a dense
silk tube found in leaves and even in human constructions (Petrunkevitch 1929).

Caribena versicolor (Walckenaer, 1837), comb. n.


Figs 18, 226, 230, 244–255, 306

Mygale versicolor Walckenaer, 1837: 211 (holotype considered lost).


Avicularia rutilans Ausserer, 1875: 184, t. 7, fig. 34 (holotype male, N. Granada, Key-
serling collection, BMNH 1890.7.1.359, examined); F. O. Pickard-Cambridge
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 139

1896: 744; Petrunkevitch 1911: 50, 1939: 28; Mello-Leitão 1923: 377; Roewer
1942: 255; Bonnet 1955: 832; World Spider Catalog 2016. Syn. n.
Avicularia versicolor: Simon 1892: 171, 172; Petrunkevitch 1911: 51, 766, 1939: 288;
Mello-Leitão 1923: 377; Roewer 1942: 255; World Spider Catalog 2016.

Remarks. Avicularia rutilans holotype has same morphology of palpal bulb, tibial
apophysis and urticating setae II and the characteristic intense green sheen found in
specimens from Martinique. Despite labeled as coming from N. Granada (Colombia,
see discussion below), it is undoubtedly a specimen of C. versicolor comb. n. Thus, we
consider A. rutilans Ausserer, 1875 as junior synonym of C. versicolor (Walckenaer,
1837) comb. nov.
Diagnosis. Females can be distinguished from those of C. laeta comb. n. by sper-
mathecae with accentuated outwards curvature medially and by lacking intumescence
on apex (Fig. 230). Males differ from males of C. laeta comb. n. by developed promi-
nence on tegulum (Fig. 246), embolus medial portion and tegulum’s margin form an
acute angle in retrolateral view (Fig. 245) and by very curved shape of basal part of
embolus in frontal view (Fig. 246).
Type material. Holotype considered, herein, lost since it is not in MNHN–AR
where it should be deposited according to description (Walckenaer 1837). Here, we
establish the male AR 4904 as the neotype since no holotype is believed to be extant
and a type is considered necessary to define the taxon objectively because the original
name is involved in a complex taxonomic problem (article 75.1, ICZN 1999).
Material examined. Martinique, 6 females, 2 males (one male established here as
neotype), 1 immature male, Gambey col., (MNHN–AR 4904, Box 314).
Additional material. CUBA: 1 male, 2 females (MNHN–AR); MARTINIQUE:
Le Carbet [14°42'N, 61°10'W], Morne Carbet, 1500 m, forest, 2 males, 18 March
1967 (AMNH); 1 male, Beatty col., 1967 (AMNH), from trees, 1 female (AMNH);
Absalon [14°41'N, 61°6'W], 1 immature male, Vaunes, 16 June 1960 (AMNH Ma);
Anse Céron [14°50'N, 61°13'W], 1 km SE, 1 male, J. Paterson col., 7 July 1998
(AMNH RW07), same data, 1 male (AMNH RW06); 1 male, Le Prêcheur, west of
Mountain Pelée, P. Charbonnet col., 21 August 1998, on side of tree (MNRJ 06914).
Remarks. In order to describe Mygale versicolor, Walckenaer (1837) used a female
from Guadeloupe and a male from Brazil. Even though he considered Aranea hirtipes
Fabricius, 1787 described from French Guiana as a synonym of M. versicolor, Walck-
enaer (1837) did not include Cayenne nor Brazil in the distribution area, only Guade-
loupe and Martinique. Concerning the description, the female corresponds to C. ver-
sicolor comb. n., but the male did not, since it was described as having a two-branched
tibial apophysis and occurs in Brazil. Therefore, the syntype series is composed of two
different species, causing a complex taxonomic problem that lasted for years.
C. L. Koch (1842) apparently followed Walckenaer (1837) but formally extended
the species distribution to Brazil. A few years later, the same author (C. L. Koch 1850)
transferred this species to Lasiodora, creating the new combination Lasiodora versicolor.
140 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 244–251. Caribena versicolor (Walckenaer, 1837) comb. n., male neotype (MNHN–AR 4904).
244–247 left palpal bulb 244 prolateral 245 retrolateral 246 frontal 247 dorsal 248 left cymbium, dorsal
249–251 left tibial apophysis of leg I 249 prolateral 250 ventral 251 retrolateral. Scale bars = 1 mm.

In 1871, Ausserer (1871) erected the new genus Homoeomma Ausserer, 1871 with
a single species, Homoeomma versicolor (Walckenaer, 1837). Ausserer (1871) examined
a male and a female from Rio de Janeiro, Brazil and considered Mygale versicolor sensu
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 141

C. L. Koch, 1842 and Lasiodora versicolor (C. L. Koch, 1850) synonyms of H. versi-
color. Years later, O. Pickard-Cambridge (1881) explicitly considered as being distinct
the concepts of M. versicolor from C. L. Koch (1842) and Walckenaer (1837).
He described a new species, Homoeomma stradlingi O. Pickard-Cambridge, 1881,
based on the concept of Mygale versicolor used by C. L. Koch, and in a female and
two males from Brazil (O. Pickard-Cambridge 1881). The species H. versicolor was
included in the synonymic list with a question mark by O. Pickard-Cambridge (1881).
Simon (1892) considered that M. versicolor Walckenaer was erroneously reported
as Homoeomma and it was, in fact, an Avicularia species very common in Lesser Antil-
les. As a consequence, it is likely Simon (1892) ignored the combination H. versicolor
and considered H. stradlingi as the type species of Homoeomma.
A problem lies with the identity of Mygale versicolor Walckenaer, 1837. This name is
constantly applied to aviculariine species found in Martinique. However, as mentioned
before, the type series consists of two different species: one aviculariine; and the other a
theraphosine. As the description is vague and syntype series is here considered lost, we
decide to establish a neotype in order to preserve the name’s stability and its usage.
Female. Redescription. AR 4904. Carapace: 19.07 long, 16.75 wide, 5.33 high.
Chelicera: 6.90 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 14.29,
8.76, 10.86, 9.49, 6.12, 49.52. II: 13.16, 7.74, 10.32, 9.23, 5.92, 46.37. III: 11.96,
7.64, 9.49, 9.40, 5.18, 43.67. IV: 14.86, 7.98, 12.21, 13.25, 5.99, 54.29. Palp: 10.16,
6.58, 6.79, –, 7.42, 30.95. Midwidths: femora I–IV= 3.61, 3.74, 3.80, 3.59, palp=
2.90; patellae I–IV= 3.67, 3.62, 3.30, 3.48, palp= 3.06; tibiae I–IV= 3.08, 2.69, 2.91,
3.06, palp= 2.83; metatarsi I–IV= 2.37, 2.27, 2.60, 2.03; tarsi I–IV= 2.86, 2.58, 2.75,
2.89, palp= 2.92. Abdomen: 22.09 long, 17.02 wide. Spinnerets: PMS, 1.45 long,
1.01 wide, 0.10 apart; PLS, 2.56 basal, 0.77 middle, 3.04 distal; midwidths 2.10, 1.75,
1.52, respectively.
Carapace: 1.14 times longer than wide; cephalic region not raised, carapace striae
inconspicuous.
Fovea: deep, slightly recurve, 2.33 wide.
Eyes: eye tubercle 1.11 high, 2.20 long, 3.18 wide. Clypeus 0.64. Anterior row of
eyes procurve, posterior slightly recurve. Eye size and interdistances: AME 0.78, ALE
0.83, PME 0.36, PLE 0.82, AME–AME 0.47, AME–ALE 0.51, AME–PME 0.22,
ALE–ALE 1.83, ALE–PME 0.75, PME–PME 1.78, PME–PLE 0.12, PLE–PLE 2.38,
ALE–PLE 0.43, AME–PLE 0.67.
Maxilla: length to width: 1.84. Cuspules: 100–200 spread over ventral inner heel.
Labium: 2.07 long, 3.19 wide, with 115 cuspules spaced by one diameter from each
other on anterior third. Labium sternal groove shallow, flat, sigilla not evident.
Chelicera: basal segment with 11 teeth and some small teeth on promargin. Ster-
num: 10.21 long, 6.83 wide. Sigilla: only posterior pair evident, rounded, less than one
diameter from margin.
Legs: Formula: IV=I II III. Length leg IV to leg I: 1.10. Clavate trichobothria: dis-
tal 1/2 tarsi I–IV. Scopulae: Tarsi I–IV fully scopulate. Metatarsi I–II fully scopulate,
III 1/2, IV 1/3 distal scopulate. IV divided by a row of setae.
142 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 252–255. Caribena versicolor (Walckenaer, 1837) comb. n., habitus. 252 immature 253 juvenile
254 female 255 male. Arrow: urticating setae type II on conspicuous patch on dorsal abdomen. Photos:
R. C. West.

Type II urticating setae: 1.33–1.58 long, 0.006–0.009 wide.


Spermathecae (Fig. 230): two completely separated, not-twisted long spermathe-
cae, with walls lacking projections or lobes and accentuated outwards curvature medi-
ally. Midwidth as wide as its base width and weakly-sclerotized area shorter than half
the length of well-sclerotized area.
Color pattern (Fig. 254): carapace brown with golden short body setae with very
intense green sheen. Carapace border with long setae the same color as dorsal carapace
short body setae, but with iridescent sheen. Coxae, labium, sternum and maxillae light
brown, same color of ventral femora. Legs and palps with gold short body setae with
green sheen and brown long guard-setae with very intense iridescent sheen. Leg rings
on distal femora, tibiae and metatarsi same color as the rest of segment. Dorsal abdo-
men with vivid red long guard-setae with very intense iridescent sheen homogeneously
distributed and black, short body setae. Ventral abdomen brown. Urticating setae
form very distinctive small bronze patch on dorso posterior area of abdomen.
Male. Description. AR 4904. Carapace: 16.01 long, 15.05 wide, 4.50 high. Cheli-
cera: 6.16 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 14.39, 8.80, 11.38,
10.76, 6.18, 51.51. II: 13.64, 8.34, 11.03, 10.31, 5.80, 49.12. III: 12.28, 6.97, 9.43,
10.27, 5.57, 44.52. IV: 15.01, 7.65, 12.92, 13.89, 5.41, 54.88. Palp: 9.26, 5.83, 7.00,
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 143

–, 2.93, 25.02. Midwidths: femora I –IV= 3.08, 3.13, 3.20, 2.92, palp= 2.24; patellae I–
IV= 3.27, 2.88, 2.92, 3.06, palp= 2.30; tibiae I–IV= 2.59, 2.43, 2.32, 2.59, palp= 2.17;
metatarsi I–IV= 1.61, 1.76, 1.85, 1.71; tarsi I–IV= 2.02, 2.01, 2.06, 2.07, palp= 2.23.
Abdomen: 18.41 long, 13.01 wide. Spinnerets: PMS, 1.94 long, 0.92 wide, 0.30 apart;
PLS, 2.40 basal, 1.14 middle, 2.64 distal; midwidths 1.52, 1.36, 1.14, respectively.
As in female, except:
Carapace: 1.06 times longer than wide.
Fovea: 1.27 wide.
Eyes: eye tubercle 1.14 high, 2.30 long, 3.01 wide. Clypeus 0.54. Eye size and
interdistances: AME 0.78, ALE 0.70, PME 0.31, PLE 0.62, AME–AME 0.41, AME–
ALE 0.51, AME–PME 0.13, ALE–ALE 1.94, ALE–PME 0.68, PME–PME 1.73,
PME–PLE 0.16, PLE–PLE 2.14, ALE–PLE 0.18, AME–PLE 0.55.
Maxilla: length to width: 1.77. Labium: 1.89 long, 2.73 wide, with 95 cuspules.
Labium sternal groove with two large separate sigilla.
Chelicera: basal segment with 10 teeth and some small teeth on promargin. Ster-
num: 7.49 long, 6.41 wide. Sigilla: three pairs, large posterior.
Legs: Length leg IV to leg I: 1.07. Scopula: tarsus IV with few, sparse setae. Meta-
tarsus IV divided by a row of setae.
Type II urticating setae (Fig. 18): 1.61–1.75 long, 0.007–0.011 wide.
Palp (Figs 244–247): globous bulb with small subtegulum and developed promi-
nence on tegulum. Embolus: not flattened, lacking keels, 4.64 long in retrolateral view,
about 3 times tegulum’s length. Medial portion and tegulum’s margin form an acute an-
gle in retrolateral view. Proximal part very curved in frontal view; thin distal width, taper-
ing distally; basal, middle, and distal width of 0.70, 0.22, 0.08, respectively. Tegulum:
2.42 long, 1.51 high in retrolateral view. Cymbium subtriangular with subequal lobes,
having developed sharp process on retrolateral lobe, bearing thin setae (Figs 248, 306).
Tibial apophysis (Fig. 250): single branch on prolateral leg I, with weakly-devel-
oped base and grouped spiniform setae distally. Male metatarsus I touches retrolater-
ally tibial apophysis’ setae when folded.
Color pattern ontogeny. Juveniles with metallic sheen, all articles with same
blackish color (Fig. 252) and abdomen dorsum with dorsal central longitudinal black
stripe connected with all transversal black stripes.(Fig. 253). When mature, they lose
this pattern (Figs 254–255).
Distribution. Martinique (Fig. 226).
Remark. Contrasting with Walckenaer’s description (Walckenaer 1837), this spe-
cies is not found on Guadeloupe Island (P. Marechal, pers. comm.). Thus, it is possible
that Walckenaer (1837) used a specimen from Martinique, which is close to Guade-
loupe Island and also French territory, but recorded as coming from Guadeloupe.
Similar situations were very common in past centuries.
Occurrence of C. versicolor comb. n. in Cuba is still doubtful since the specimens
here analyzed are very old and the only ones known from Cuba in the entirety of arach-
nological collections examined. The record of A. rutilans, junior synonym of C. versicolor
comb. n. in Nova Granada, former name of Colombia, is also doubtful. There is no other
144 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

record of this species in Colombia nor in any other continental territory. An explanation
could be that this specimen was dispatched from Bogota, an important South Amercian
commercial center in eighteenth and nineteenth centuries, which is near the Antilles.
Natural history. It was poorly known until a few years ago. The situation changed
since an extensive and detailed ecological and populational study was carried out by
Maréchal et al. (2009) on Martinique. They pointed out that C. versicolor comb.
n. prefers mesophyll forests, building its retreat in Bromeliacea leaves, between tree
branches, inside bamboo trunks, over tree trunks and also within human construc-
tions (Maréchal et al. 2009). Its life cycle is about two years, reaching sexual maturity
around 18 months, after approximately ten molts; the reproductive period is in March
and eggsac eclosion is in May and June (Maréchal et al. 2009).
Remarks. Two color forms are known, one with specimens having leg and palp
hairs in bright red and the other with specimens with darker hairs on legs and palps.

Ybyrapora gen. n.
http://zoobank.org/68B8A50A-E657-4091-8931-58859EF55AAC
Figs 256–283, 295–298, 305, 308, 319

Diagnosis. Ybyrapora gen. n. can be distinguished from all other aviculariine genera, ex-
cept Avicularia, Caribena gen. n., Iridopelma and Typhochlaena by procurve anterior row of
eyes. It can be distinguished from Typhochlaena by digitiform apical article of PLS. From
Iridopelma by males lacking tibial apophysis on tibiae II and females by long spermathe-
cae, with accentuated outwards curvature medially. It differs from Caribena gen. n. by
stout urticating setae on abdomen dorsum of male and female, and by lacking sharp pro-
cess bearing thin setae on retrolateral lobe of cymbium in males. From Avicularia females
it can be distinguished by having virtually non-sclerotized spermathecae (Figs 257–259).
From Avicularia males (except A. minatrix, A. lynnae sp. n., A. caei sp. n., and A hirschii) it
differ by lacking developed tibial apophysis (Fig. 308). Males of Y. diversipes comb n. can
be distinguished from those of four species by having very long embolus strongly curved in
frontal view (Fig. 278). Males of Y. sooretama comb. n. and Y. gamba comb. n. differ from
those of A. lynnae sp. n., A. caei sp. n., and A. hirschii by lacking well-developed process
on cymbium (Fig. 305). They can be distinguished from A. minatrix by color pattern of a
single red (Fig. 297) or dark longitudinal stripe on the abdomen dorsum.
Etymology. From the ancient Tupi, the classic indigenous language from Brazil,
“ybyrá”, meaning “tree”; and “pora”, meaning “those that lives in”. Thus, Ybyrapora
means “those that lives in trees”, regarding the arboreal habit of the three species of
the genus.
Description. Carapace slightly longer than wide, cephalic region slightly raised.
Cephalic and thoracic striae inconspicuous or conspicuous. Fovea shallow and straight.
Chelicera without rastelum. Eye tubercle distinct, slightly raised, wider than long. An-
terior row of eyes procurve, posterior slightly recurve or recurve. Clypeus absent or
narrow. Labium subquadrate, wider than long, with 35–130 cuspules on anterior third
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 145

center, spaced by more than one diameter from each other on anterior third. Maxil-
lary lyra absent. Maxilla subretangular, anterior lobe distinctly produced into conical
process, inner angle bearing 80–200 cuspules. Sternum longer than wide, posterior
angle acute, not separating coxae IV. Sigilla anterior not evident, middle fusiform,
posterior ellipsoidal, one to 1.5 diameter from margin. Leg formula: I=IV II III (most
species) or IV I II III (Y. sooretama comb. n. female). Clavate trichobothria on the
distal 2/3 to 1/2 of tarsi I–IV. Tarsi I–IV fully scopulate, IV divided or not by band
of setae. Metatarsi I–II 4/5 to fully scopulate, III 2/3, IV 2/3 to 1/3 distal scopulate.
Metatarsi IV divided by row of setae. Scopulae of tarsi and metatarsi I–II extended very
laterally giving them spatulate appearance. Femora IV without retrolateral scopulae.
Stridulatory setae absent. Spiniform setae absent. ITC absent; STC without denticles.
Posterior lateral spinneret distally elongating, digitiform. Typical type II urticating
setae on dorsal abdomen. Tibia I with discrete elevation covered by a cluster of setae in
apical portion, on prolateral side (Y. diversipes comb. n.) (Fig. 282) or lacking modi-
fications (other species) (Fig. 266). Tibial apophysis on leg II absent. Globous bulb
with small subtegulum and developed prominence on tegulum (Y. sooretama comb.
n.) (Fig. 262) or without prominence (other species) (Figs 270, 278). Embolus not
flattened, lacking keels, about 3 to 3.5 times tegulum’s length (Y. sooretama comb. n.
and Y. gamba comb. n.) (Figs 261, 269) or more than 4 times tegulum’s length (Y.
diversipes comb. n.) (Fig. 277). Embolus medial portion and tegulum’s margin form-
ing an obtuse angle in retrolateral view (Y. sooretama comb. n. and Y. gamba comb. n.)
(Figs 261, 269) or forming very acute angle in retrolateral view (Y. diversipes comb. n.)
(Fig. 277). Embolus proximal part very curved in frontal view (Y. sooretama comb. n.
and Y. diversipes comb. n.) (Figs 262, 278) or slightly curved (Y. gamba comb. n.) (Fig.
270). Embolus with thin distal width, tapering distally (Y. diversipes comb. n. and Y.
gamba comb. n.) (Figs 279, 271) or narrowing abruptly (Y. sooretama comb. n.) (Fig.
263). Cymbium subtriangular with subequal lobes, having well- developed rounded
process on retrolateral lobe, bearing thick setae (Y. diversipes comb. n.) (Fig. 280) or
lacking any process (Y. sooretama comb. n. and Y. gamba comb. n.) (Figs 264, 272).
Spermathecae with accentuated outwards curvature medially, virtually not sclerotized,
completely separated, not-twisted and with midwidth as wide as its base width (Figs
257–259). Spermathecae long (Y. sooretama comb. n. and Y. gamba comb. n.) (Figs
257–258) or very long (Y. diversipes comb. n.) (Fig. 259), with multilobular apex (Y.
sooretama comb. n.) (Fig. 257) or lacking lobes (other species) (Figs 258−259). Drastic
ontogenetic changes on abdominal color pattern. Immatures with abdomen with a
central longitudinal reddish stripe inside a dark area with zigzag borders connected to
transverse dark stripes (Fig. 295). Males with central longitudinal stripe on dorsal ab-
domen (Y. sooretama comb. n. and Y. gamba comb. n.) or lacking stripes (Y. diversipes
comb. n.).
Type species. Avicularia sooretama Bertani & Fukushima, 2009, here designated.
Species included. Ybyrapora sooretama (Bertani & Fukushima, 2009) comb. n.,
Ybyrapora diversipes (C. L. Koch, 1842) comb. n. and Ybyrapora gamba (Bertani &
Fukushima, 2009) comb. n.
146 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 256. Map showing records of Ybyrapora gen. n.

Distribution and habitat. Brazil, in Atlantic rainforest from Bahia to southern


Rio de Janeiro (see Bertani and Fukushima 2009, appendix A, fig. 17) (Fig. 256).

Key for Ybyrapora gen. n. species

Males

1 Embolus length more than 4 times tegulum’s length (Fig. 277).....................


.................................................................................. Y. diversipes comb. n.
– Embolus length from 3.0 to 3.5 times tegulum’s length (Figs 261, 269)......2
2 Abdomen dorsum with dark central longitudinal stripe, tegulum with devel-
oped prominence, embolus with basal part very curved in frontal view (Fig.
262)...........................................................................Y. sooretama comb. n.
– Abdomen dorsum with red central longitudinal stripe (Fig. 297), tegulum
lacking prominence (Fig. 270), embolus with basal part slightly curved in
frontal view (Fig. 270).................................................... Y. gamba comb. n.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 147

Females
1 Spermathecae extremely long, with distal apex reaching base (Fig. 259).........
.................................................................................. Y. diversipes comb. n.
– Short spermathecae, distal apex not reaching base........................................2
2 Spermathecae with multilobular apex (Fig. 257)........Y. sooretama comb. n.
– Spermathecae lacking lobes on apex (Fig. 258)............... Y. gamba comb. n.

Ybyrapora sooretama (Bertani & Fukushima, 2009), comb. n.


Figs 256–257, 260–267, 298, 308

Avicularia sooretama Bertani & Fukushima, 2009: 29, figs 5–8, 17, Appendix II, figs
B1–B4 (holotype male, Brazil, state of Espírito Santo, Reserva Biológica de So-
oretama [18°59'S, 40°07'W], at night, AMNRJ, 18 April 2006, MNRJ 18435
and paratype female, Brazil, state of Espírito Santo, Pinheiros, Reserva Biológica
Córrego do Veado (18°37'0.16"S, 40°14'1.60"W), 71 m a.s.l., AMNRJ, 22 Oc-
tober 2005, MNRJ 12930, examined); Bertani 2102: 5, 79; 80, 88; World Spider
Catalog 2016.

Diagnosis (amended from Bertani and Fukushima 2009). Females of Y. sooretama


comb. n. resemble those of Y. gamba comb. n. by elongated spermathecae with accentu-
ated outwards curvature medially and by retaining vestiges of characteristic juvenile color
pattern on abdomen. They differ from those of Y. gamba comb. n. by spermathecae bear-
ing multilobular apex (Fig. 257). Males resemble those of Y. gamba comb. n. by embolus
length 3.0 to 3.5 times tegulum’s length and lacking a very strong curvature in frontal
view (Fig. 262). They differ from those of Y. gamba comb. n. by developed prominence
on tegulum (Fig. 262) and by central longitudinal black stripe on dorsal abdomen.
Description, color pattern ontogeny, distribution and natural history. See
Bertani and Fukushima (2009) (Fig. 256).
Complementary description. Male: Palp (Figs 260–263): globous bulb with small
subtegulum and developed prominence on tegulum. Embolus: not flattened, lacking
keels, 3.15 long in retrolateral view, about 3.5 times tegulum’s length. Medial portion
and tegulum’s margin form an obtuse angle in retrolateral view. Proximal part very
curved in frontal view; thin distal width, abruptly narrowing distally; basal, middle, and
distal width of 0.63, 0.13, 0.03, respectively. Tegulum: 1.61 long, 0.94 high. Cymbium
subtriangular with subequal lobes, lacking process on retrolateral lobe (Fig. 264).
Tibial apophysis in leg I absent (Figs 265–267).
Type II urticating setae: 0.633–0.681 long; 0.012–0.016 wide in male; 0.422–
0.490 long; 0.009–0.012 wide in female.
Female: Spermathecae (Fig. 257): two completely separated, not-twisted, long
spermathecae, with multilobular apex and accentuated outwards curvature medially.
Spermatheca midwidth as wide as its base width, virtually non-sclerotized.
148 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 257–259. Ybyrapora gen. n., spermathecae variation. 257 Ybyrapora sooretama (Bertani &
Fukushima, 2009) comb. n., Reserva Biológica de Sooretama, state of Espírito Santo, Brazil, paratype
(MNRJ 12930) 258 Ybyrapora gamba (Bertani & Fukushima, 2009) comb. n., RPPN Jequitibá, Elísio
Medrado, state of Bahia, Brazil, paratype (MZUSP 31116) 259 Ybyrapora diversipes (C. L. Koch, 1842)
comb. n., Ilhéus, state of Bahia, Brazil (IBSP 11754). Scale bars = 1 mm.

Ybyrapora gamba (Bertani & Fukushima, 2009), comb. n.


Figs 256, 258, 268–275, 295–297

Avicularia gamba Bertani & Fukushima, 2009: 32, figs 9–12, 16–17, Appendix III,
figs C1–C6 (holotype male, Brazil, state of Bahia, Elísio Medrado, RPPN Jequiti-
bá, (12°52'3.20"S, 39°28'9.09"W), R. Bertani, C. S. Fukushima and R. H. Na-
gahama col., 7 October 2007, collected at night, found immature inside a retreat
made with silk and leaves, matured in captivity on June 2009, MZUSP 31115,
and paratype, female, Brazil, state of Bahia, Elísio Medrado, RPPN Jequitibá,
(12°52'3.20"S, 39°28'9.09"W), R. Bertani, C. S. Fukushima and R. H. Naga-
hama col., 7 October 2007, collected at night, found immature inside a retreat
made with silk and leaves, MZUSP 31116, examined); Bertani 2012: 5, 79; 80,
88; World Spider Catalog 2016.

Diagnosis (amended from Bertani and Fukushima 2009). Females of Y. gamba


comb. n. resemble those of Y. sooretama comb. n. by elongated spermathecae with
accentuated outwards curvature medially (Fig. 258) and by retaining vestiges of the
characteristic juvenile color pattern on abdomen (Fig. 298). They differ from those
of Y. sooretama comb. n. by spermathecae lacking multilobular apex (Fig. 258). Males
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 149

Figures 260–267. Ybyrapora sooretama (Bertani & Fukushima, 2009) comb. n., male holotype (MNRJ
18435). 260–263 right palpal bulb (mirrored) 260 prolateral 261 retrolateral 262 frontal 263 dorsal
264 right cymbium, dorsal (mirrored) 265–267 left tibia I 265 prolateral 266 ventral 267 retrolateral.
Scale bars = 1 mm.

resemble those of Y. sooretama comb. n. by embolus length 3.0 to 3.5 times tegulum’s
length (Fig. 269) and by lacking strong curvature in frontal view (Fig. 270). They differ
from those of Y. sooretama comb. n. by lacking prominence on tegulum (Fig. 270) and
by having central longitudinal red stripe on dorsal abdomen (Fig. 297).
150 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 268–275. Ybyrapora gamba (Bertani & Fukushima, 2009) comb. n., male holotype (MZUSP
31115). 268–271 left palpal bulb 268 prolateral 269 retrolateral 270 frontal 271 dorsal 272 left cymbium,
dorsal 273–275 left tibia leg I 273 prolateral 274 ventral 275 retrolateral. Scale bars = 1 mm.

Diagnosis, material examined, description, color pattern ontogeny, distribution


and natural history. See Bertani and Fukushima (2009) (Fig. 256).
Complementary description. Male: Palp (Figs 268–271): globous bulb with
small subtegulum lacking prominence on tegulum. Embolus: not flattened, lacking
keels, 2.34 long in retrolateral view, about 3.5 times tegulum’s length. Medial portion
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 151

and tegulum’s margin form an obtuse angle in retrolateral view. Proximal part slightly
curved in frontal view; thin distal width, tapering distally; basal, middle, and distal
width of 0.44, 0.11, 0.03, respectively. Tegulum: 1.03 long, 0.59 high. Cymbium
subtriangular with subequal lobes, lacking process on retrolateral lobe (Fig. 272).
Tibial apophysis in leg I absent (Figs 273–275).
Type II urticating setae: 0.605–0.750 long; 0.012–0.017 wide in male; 0.301–0.352
long; 0.006–0.009 wide in female (ecdise).
Female: Spermathecae (Fig. 258): two completely separated, not-twisted long
spermathecae lacking lobes and accentuated outwards curvature medially. Spermathe-
ca midwidth as wide as its base width, virtually not sclerotized.

Ybyrapora diversipes (C. L. Koch, 1842), comb. n.


Figs 256, 259, 276–283, 319

Mygale diversipes C. L. Koch, 1841: 65, pl. CCCX, fig. 731 (lectotype female, Brazil,
Bahia, Freir. leg., ZMB 2943, examined).
Eurypelma diversipes: C. L. Koch 1850: 73; Simon 1864: 67, 1892: 172; Ausserer
1871: 202; Roewer 1942: 239; Bonnet 1955: 1831; 1957: 2990.
Avicularia diversipes: F. O. Pickard-Cambridge 1896: 744; Bertani and Fukushima
2009: 26 (figs 1–4, 13–15, 17, Appendix I, figs A1–A6); Bertani 2102: 5, 79; 80,
88; World Spider Catalog 2016.

Diagnosis (amended from Bertani and Fukushima 2009). Females differ from those
of Y. sooretama comb. n. and Y. gamba comb. n. by presenting very long, strongly
curved outwards spermathecae with its distal portion almost reaching the spermathe-
cae base (Fig. 259). Males differ from those of Y. gamba sp. comb. n. and Y. sooretama
comb. n. by having embolus more than four times tegulum’s length, with strong curva-
ture in frontal view (Fig. 278) and cymbium with well-developed process bearing thick
setae on retrolateral lobe (Fig. 280).
Material examined. Female, Brazil, state of Bahia, Ilhéus, CEPLAC [14°46'S,
39°13'W], R. Bertani & G. Puorto col., March 1991 (IBSP 11754); male, same local-
ity, collectors and date (IBSP 119271 ref. 64.583).
Diagnosis, other material examined, description, color pattern ontogeny, dis-
tribution and natural history. See Bertani and Fukushima (2009).
Complementary description. Male: Palp (Figs 276–279): globous bulb with
small subtegulum, lacking prominence on tegulum. Embolus: not flattened, lacking
keels, 7.76 long in retrolateral view, about 7 times tegulum’s length. Medial portion
and tegulum’s margin form very acute angle in retrolateral view. Proximal part very
curved in frontal view; thin distal width, tapering distally; basal, middle, and distal
width of 1.05, 0.35, 0.08 respectively. Tegulum: 1.89 long, 1.08 high. (Fig. 319).
Cymbium subtriangular with subequal lobes, with a well-developed rounded process
on retrolateral lobe, bearing thick setae (Fig. 280).
152 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 276–283. Ybyrapora diversipes (C. L. Koch, 1842) comb. n., male (IBSP 119271). 276–279 left
palpal bulb 276 prolateral 277 retrolateral 278 frontal 279 dorsal 280 left cymbium, dorsal 281–283 left
tibia I 281 prolateral 282 ventral 283 retrolateral. Scale bars = 1 mm.

Tibia I with discrete elevation covered by a cluster of setae in apical portion, on


prolateral side (Figs 281–283).
Type II urticating setae: 0.791–0.860 long; 0.017–0.020 wide in male; 0.427–
0.520 long; 0.012–0.016 wide in female.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 153

Female: Spermathecae (Fig. 259): two completely separated, not-twisted very long
spermathecae, with walls lacking lobes and accentuated outwards curvature medially.
Spermatheca midwidth as wide as its base width, virtually non-sclerotized.

Antillena Bertani, Huff & Fukushima, gen. n.


http://zoobank.org/D95B044F-815A-48F1-85D7-37C1669F86B3
Figs 226, 284–294

Avicularia rickwesti Bertani & Huff, 2013: 333, figs 2–19 (holotype female, Domini-
can Republic, Pedernales Province, Parque Nacional Jaragua, track into park (un-
marked) between Manuel Goya (Manuell Goa [sic]) and Oviedo (17°48'41.5"N,
71°26'35.9"W), 83.3 m a.s.l., 09 July 2004, J. Huff and E. S. Volschenk leg.,
(collecting permit #01496), AMNH, and paratype female, Dominican Repub-
lic, Independencia Province: Parque Nacional Sierra de Baoruco, Rabo de Gato
(18°18'39.1"N, 71°34'54.4"W), 408 m a.s.l., 10 July 2004, J. Huff & E. S. Vols-
chenk leg., collection permit #01496, AMNH, examined); Kaderka 2016: 121,
figs 2–11; World Spider Catalog 2016.

Etymology. Named after the type locality, the Antilles, where Dominican Republic is
located, and is considered feminine in gender.
Type species. Avicularia rickwesti Bertani & Huff, 2013, by monotypy.
Species included. Antillena rickwesti (Bertani & Huff, 2013) comb. n.
Diagnosis. Female differs from those of other Aviculariinae species by having two
very short and broad spermathecae, with distal half strongly sclerotized (Fig. 284). Ad-
ditionally, they can be distinguished by spiniform setae in prolateral and/or retrolateral
side of coxae I–IV. Males can be distinguished by the very flattened embolus with keels
(Figs 285–288) as well as abdominal leaf pattern (Fig. 294).
Description. See species description.
Material examined. 1 male, Dominican Republic, Pedernales Province, Ja-
ragua National Park, Los Tres Charcos, road to Fondo Paradi (17°48'7.45"N,
71°26'5.41"W), R. C. West and J. Huff col., 20 February 2012, matured in captivity
10 April 2014 (AMNH).

Figure 284. Antillena rickwesti (Bertani & Huff, 2013) comb. n., Dominican Republic (AMNH), sper-
mathecae. Scale bars = 1 mm.
154 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 285–292. Antillena rickwesti (Bertani & Huff, 2013) comb. n., male (AMNH). 285–288 left
palpal bulb 285 prolateral 286 retrolateral 287 frontal 288 dorsal 289 left cymbium, dorsal 290–292 left
tibial apophysis of leg I 290 prolateral 291 ventral 292 retrolateral. Scale bars = 1 mm.

Additional material. DOMINICAN REPUBLIC: Pedernales: 5 km south of Ma-


nuel Goya [17°52'N, 71°29'W], 1 female, R. C. West col., 20 February 2012, in silk
retreat of scrub tree (AMNH RW01); Jaragua National Park, Los Tres Charcos, road
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 155

Figures 293–294. Antillena rickwesti (Bertani & Huff, 2013) comb. n., habitus. 293 immature
294 male. Photos: 293 A. Tosto; 294 R. C. West.

Figures 295–298. Dorsal abdominal patterns of Ybyrapora gen n. 295–297 Y. gamba (Bertani & Fukushima,
2009) comb. n. 295 immature 296 adult female 297 male 298 Y. sooretama (Bertani & Fukushima, 2009)
comb. n., adult female. Photos: C. S. Fukushima.

to Fondo Paradi (17°48'7.45"N, 71°26'5.41"W), 1 female, R. C. West and J. Huff


col., 20 February 2012 (AMNH).
Female. see Bertani and Huff (2013).
Male. Description. AMNH. Carapace: 10.4 long, 9.9 wide, 2.98 high. Chelicera:
6.7 long. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 9.9, 5.9, 7.7, 8.2, 4.8,
156 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 299–304. Legs and abdominal characters. 299–300 leg rings coloration on metatarsi and tibiae,
and guard-setae coloration on legs 299 Avicularia avicularia, female, whitish leg rings and homogeneous
coloration along guard-setae on legs 300 Avicularia rufa, female, yellow leg rings and guard-setae with
darker base and contrasting whitish apex on legs 301 Psalmopoeus irminia, male, longer setae laterally pro-
jected on legs 302 Avicularia rufa, female, abdominal setae covering heterogeneously 303 Avicularia avicu-
laria, female, abdominal setae covering heterogeneously 304 Avicularia purpurea, female, abdominal setae
covering homogeneously. Photos: 299–301 R. Bertani; 302–303 C. S. Fukushima, 304 R. C. West.

35.0. II: 9.6, 5.5, 7.6, 7.9, 4.6, 33.8. III: 8.1, 4.6, 6.6, 7.1, 4.1, 30.1. IV: 10.2, 5.2,
8.7, 8.4, 3.8, 36.3. Palp: 6.2, 3.6, 4.6, –, 2.5, 21.1. Midwidths: femora I–IV= 2.1,
1.9, 2.2, 2.1, palp=1.6; patellae I–IV=2.0, 2.0, 1.9, 2.0, palp=1.6; tibiae I–IV=1.8,
1.6, 1.6, 1.9, palp=1.7; metatarsi I–IV=1.1, 1.1, 1.0, 1.1; tarsi I–IV=1.1, 1.1, 1.2, 1.3,
palp=1.3. Abdomen 14.9 long, 10.7 wide. Spinnerets: PMS, 0.9 long, 0.3 wide, 0.1
apart; PLS, 1.7 basal, 1.0 middle, 2.0 distal; midwidths 1.1, 0.8, 0.6, respectively.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 157

Figures 305–311. Aviculariine characters. 305–307 cymbium process, dorsal 305 Ybyrapora gamba
comb. n., holotype (MZUSP 31115), absent 306–307 well-developed process 306 Caribena versicolor
comb. n. (MNHN–AR 4904), sharp and bearing thin setae 307 Avicularia juruensis (CAS 6), rounded
and bearing thick setae 308–311 tibia I, retrolateral 308 Ybyrapora sooretama comb. n., holotype (MNRJ
18435), lacking modifications 309 Avicularia hirschii, paratype (SMF 57125), thicker setae on a dis-
crete elevation 310 Caribena laeta comb. n. (AMNH PR), developed branch 311 Avicularia avicularia
(MNHN–AR 4894), well-developed branch. Scale bars = 1 mm. Arrow indicates discrete elevation.

Carapace: 1.24 times longer than wide; cephalic region raised. Fovea: deep, re-
curved, 1.6 wide.
Eyes: eye tubercle 0.6 high, 1.8 long, 2.5 wide. Clypeus absent. Anterior eye row
procurve, posterior slightly recurve. Eye sizes and interdistances: AME 0.60, ALE
0.43, PME 0.28, PLE 0.30, AME–AME 0.34, AME–ALE 0.32, AME–PME 0.16,
ALE–ALE 1.78, ALE–PME 0.65, PME–PME 1.49, PME–PLE 0.07, PLE–PLE 1.94,
ALE–PLE 0.54, AME–PLE 0.42.
Maxilla: length to width: 1.76. Cuspules: 128 spread over ventral inner heel. La-
bium: 1.3 long, 1.8 wide, with 61 cuspules spaced by one diameter from each other on
anterior half. Labio-sternal groove shallow, flat, sigilla not evident.
Chelicera: basal segment with 12 teeth, second, fourth and fifth the larger; parallel
basal row of six tiny teeth on promargin.
Sternum: 5.5 long, 4.5 wide. Sigilla: anterior pair not evident, the other ellipsoidal, less
than half diameter from margin; posterior one time and half the diameter of the middle.
Legs: Formula: I=IV II III. Length leg IV to leg I: 1.04. Clavate trichobothria: 2/3
distal tarsi I–IV. Leg coxae with spiniform setae; absent on leg I and prolateral leg II,
158 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figures 312–315. Male palpal bulb prominence in Avicularia, frontal. 312 Avicularia minatrix (MZUSP
70949), absent 313 Avicularia caei sp. n. holotype (MPEG 015637), weakly-developed 314 Avicularia
avicularia (MPEG 2534), developed 315 Avicularia variegata stat. n. (INPA 4897), well-developed. Scale
bars = 1 mm. Arrows indicate groove that forms bulb prominence.

poorly developed on retrolateral leg I, prolateral and retrolateral leg III and prolateral
leg IV. Scopula: tarsi I–IV fully scopulate. Metatarsi I–II fully scopulate; III 4/5, IV
1/2 distal scopulate. IV divided by a row of setae.
Type II urticating setae: 0.77–0.88 long, 0.014–0.018 wide.
Palp (Figs 285–288): globous bulb with small subtegulum lacking prominence
on tegulum. Embolus: flattened, bearing three sharp keels: a prolateral superior on its
distal half; a prolateral inferior for full embolus length; and an apical for two-thirds of
embolus length. Embolus 1.5 long in retrolateral view. Medial portion and tegulum’s
margin form an acute angle in retrolateral view. Proximal part slightly curved in fron-
tal view; thin distal width, tapering distally; basal, middle, and distal width of 0.44,
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 159

Figures 316–319. Embolus tip of male palpal bulb. 316–317 tip narrowing abruptly 316 Avicularia avicularia
(MNRJ 13659A), frontal view 317 Avicularia variegata stat. n. (INPA 4897), dorsal view 318–319 tip tapering
318 Caribena versicolor comb. n. (MNHN–AR 4904), frontal view 319 Ybyrapora diversipes comb. n. (IBSP
119271), dorsal view. Scale bar = 1mm. Arrow indicates point of abrupt narrowing.

0.37, 0.18, respectively. Tegulum: 1.31 long, 1.18 wide in retrolateral view. Cymbium
subtriangular with subequal lobes. Cymbium with well-developed rounded process on
retrolateral lobe, bearing thick setae (Fig. 289).
Tibial apophysis (Figs 290–292): single branch on prolateral leg I, with well-developed
base and grouped spiniform setae distally. Male metatarsus I touches retrolaterally tibial
apophysis’ setae when folded.
Color pattern (Fig. 294): carapace dark brown covered with abundant light brown
short body setae especially on its margins. Legs, palps and chelicerae dorsally brown
covered with light brown short body setae. Legs and palps with brown long guard-
setae. Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen dorsally dark
brown with light pattern. Coxae, labium, sternum, maxillae and legs ventrally brown
with light brown setae. Tibiae, metatarsi and tarsi ventrally darker. Abdomen ventral-
ly greyish, distal third darker. Living specimens shows intense metallic pinkish short
body setae over carapace, dorsal legs and palps. Abdomen is dorsally black with lighter
leaf pattern.
Color pattern ontogeny. See Bertani and Huff (2013) (Fig. 293).
Distribution. Dominican Republic (Fig. 226).
Natural history. See Bertani and Huff (2013).
160 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Figure 320–323. Male palpal bulb characters. 320 Lasiodora sp., retrolateral, left palp 321–323 an-
gle between medial portion of embolus and tegulum’s margin in retrolateral view 321 Pachistopelma
bromelicola, right palp, very acute angle 322 Iridopelma hirsutum (IBSP 8077), right palp, obtuse angle
323 Avicularia avicularia, right palp, acute angle (MNRJ 13659A). Scale bars = 1 mm. Red bars show
angle between tegulum’s margin and embolus medial portion.

Cladistics

Searches using NONA (hold 10000, mult*500, hold/1000) resulted in 6 cladograms


and their strict consensus (Nelsen, L= 229, CI = 41, RI= 67) is shown in Figure 324.
The consensus cladogram is almost completely resolved. It shows Aviculariinae as non
monophyletic, since Ephebopus spp., Psalmopoeus sp. and Tapinauchenius sp. are in a
separated clade as the sister group of Phlogiellus sp. (Selenocosmiinae). Antillena rick-
westi comb. n. is the sister group of Poecilotheria sp., and both taxa could be considered
as aviculariines. The genus Avicularia is paraphyletic in this analysis, as A. minatrix is
recovered apart from the other species and as one of the most basal taxon of Aviculari-
inae. The Avicularia clade is supported by two homoplastic characters, the presence of
prominence on tegulum (character 44, state 1) and male palpal bulb with embolus me-
dial portion and tegulum’s margin form an acute angle in retrolateral view (character 51,
state 1). The genus is the sister group of (Iridopelma (Pachistopelma + Caribena gen. n.)).
With Piwe, we found one or two cladograms with each concavity used (Table 3). Use
of implied weights and different concavities resulted in more than one cladogram topol-
ogy. The topologies obtained using concavities 1 (Suppl. material 1: concavity 1) and 2
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 161

Figure 324. Strict consensus of 6 trees obtained with NONA and all characters as non-additive. Black
square = synapomorphy, white square = homoplasy. Length = 229, CI = 41, RI = 67.
162 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Table 3. Length, fit and number of cladograms obtained using implied weights and different concavities
on Piwe.

Concavity Number of cladograms Fit Length


1 1 338.7 238
2 2 400.4 238
3 1 439.7 236
4 1 466.9 234
5 1 487.1 232
6 1 503.4 230

(Suppl. material 2: concavity 2) are very similar, differing only in the relationship inside
Typhochlaena genus; the one obtained with concavities 3 (Suppl. material 3: concavity 3)
and 4 (Suppl. material 4: concavity 4) are very similar, differing only in position of Po-
ecilotheria sp. and A. rickwesti comb. n. The topologies obtained with concavities 5 (Suppl.
material 5: concavity 5) and 6 are very distinct from the others.
In topologies obtained using concavities 1 and 2, Aviculariinae is recovered as
paraphyletic since A. rickwesti comb. n. is in a clade with Poecilotheria sp., Haplopelma
sp. and Lasiodora sp., genera belonging to three different theraphosid subfamilies. The
clade (Ephebopus (Psalmopoeus + Tapinauchenius)) was not recovered as basal group,
but in a more derived group of Aviculariinae.
The genus Avicularia is recovered as polyphyletic, with Avicularia minatrix in a
basal position as the sister group of most Aviculariinae genera. Avicularia purpurea and
A. merianae sp. n. were recovered in a distinct clade in the middle of aviculariine, and
the clade with A. hirschii, A. lynnae sp. n. and A. caei sp. n. is in a trichotomy with
Caribena gen. n. and Pachistopelma + I. marcoi.
The topologies obtained using concavities 3 and 4 are very similar, differing in
the position of A. rickwesti comb. n. and Poeciolotheria sp. In topology obtained with
concavity 4, they form a clade inside Aviculariinae; in concavity 3, A. rickwesti comb.
n. is in a clade with some genera of the outgroup, making Aviculariinae paraphyletic.
Their topologies resemble those obtained with concavities 1 and 2, differing mainly
in position of the clade Ephebopus (Tapinauchenius + Psalmopoeus), which is basal in
Aviculariinae with concavities 3 and 4, and differing in A. minatrix position, which is
basal in Aviculariinae with concavities 1 and 2.
The topology obtained with concavity 5 resembles the topology obtained using
concavity 6. It differs mainly in Typhochlaena position, which is recovered as a inner
group of Avicularia, as well as in A. rickwesti comb. n. position, which is the sister
group of most aviculariine genera.
In agreement with other studies (Ramírez 2003, West et al. 2008, Bertani 2012),
the shortest cladogram with highest fit on Piwe was obtained when using concavity
6 (Table 3, Fig. 325) and it is chosen as the preferred cladogram, of which the main
discussion is done (Tables 4–5). In some cases, discussion about specific clades can
include comments about topologies obtained with different concavities.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 163

Figure 325. Single tree obtained with Piwe, all characters as non-additive and concavity 6. Fit = 5051.44
and length = 230. Bremer support values (absolute and relative) of each node are in red.

In the preferred cladogram, Avicularia is monophyletic (node 81 in Fig. 325) and


is composed of 11 species, plus A. glauca which was not included in the matrix: A.
avicularia, A. minatrix, A. purpurea, A. hirschii, A. taunayi, A. juruensis, A. variegata
stat. n., A. rufa, A. merianae sp. n., A. lynnae sp. n. and A. caei sp. n. The sister-group of
Avicularia is a clade formed by Antillena gen. n., Caribena gen. n., Pachistopelma, Iri-
dopelma and Ybyrapora gen. n. The characters supporting Avicularia clade are juveniles
with black tarsi contrasting with other lighter articles (character 27, state 1) (Fig. 44),
with a reversion in A. purpurea; spermathecae with accentuated outwards curvature
medially (character 37, state 1) (Fig. 21) and male palpal bulb with embolus medial
portion and tegulum’s margin form an acute angle in retrolateral view (character 51,
state 1) (Fig. 323). All these characters are very homoplastic.
The species A. minatrix was retrieved as the most basal species in Avicularia. Avicu-
laria purpurea and A. merianae sp. n. are very similar species regarding body dimen-
sions, spermathecae and cymbium morphology as well as geographic distribution area,
164 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Table 4. Synapomorphies for cladogram of Fig. 325.

Taxa or
Taxa or Node Character Change Taxa or Node Character Change Character Change
Node
33 0→1 27 0 → 1 Node 55 11 1→0
H. rondoni
42 0→1 33 0 → 1 Node 56 51 2→1
T. curumim (cont.)
1 0→1 35 0 → 1 Node 57 50 2→1
Pterinochilus
15 0→1 68 1→0 4 0→1
sp.
57 0→1 24 4 → 5 Node 58 19 0→1
1 0→1 T. paschoali 27 0→1 70 0→2
7 0→1 33 0→1 0 0→1
13 0→1 P. rufonigrum 25 0 → 2 Node 59 21 1→0
P. muticus
18 0→1 I. hirsutum 28 0→1 68 1→0
33 0→1 I. zorodes 46 2 → 0 Node 60 62 0→1
50 0→1 21 1→0 46 2→0
Node 61
5 0→1 26 1→0 67 1→2
I. katiae
15 0→1 27 0 → 1 Node 62 35 0→1
Haplopelma sp. 16 0→1 70 1 → 3 Node 63 33 0→1
50 0→1 33 0 → 1 Node 64 59 0→1
I. marcoi
57 0→2 35 0→1 40 0→1
Node 65
3 0→2 A. minatrix 49 0→1 63 0→1
Phlogiellus sp. 10 1→0 A. lynnae sp. n. 28 0 → 1 Node 66 39 1→0
17 0→1 A. caei sp. n. 45 1→0 28 0→1
Node 67
14 0→1 25 0→2 38 1→0
C. laeta comb. n.
18 0→1 50 2→1 36 0→2
24 1→0 21 1 → 0 Node 68 37 0→1
32 2→1 37 0→1 56 0→1
Lasiodora sp. C. versicolor comb. n.
41 1→0 44 0 → 1 Node 69 70 0→1
46 0→1 51 2→1 23 0→1
60 0→1 33 0 → 1 Node 70 26 0→1
64 0→1 44 0→1 58 1→0
Y. sooretama comb. n.
3 0→1 49 0 → 1 Node 71 38 0→1
24 1→3 68 1→0 67 1→0
Node 72
33 0→1 Y. gamba comb. n. 50 2→1 68 1→0
P. vulpinus 41 1→0 21 1 → 2 Node 73 45 1→2
64 0→1 Y. diversipes comb. n. 46 2→3 29 0→1
Node 74
65 0→1 51 2→0 30 0→1
68 1→2 46 2 → 1 Node 75 49 0→1
9 1→2 48 0 → 2 Node 76 36 0→1
11 0→1 50 2 → 1 Node 77 46 2→3
A. rickwesti comb. n.
16 0→1 52 0 → 1 Node 78 39 1→0
Poecilotheria sp.
17 0→1 53 0 → 1 Node 79 38 0→1
24 1→2 54 0→1 23 0→1
Node 80
46 0→1 A. taunayi 33 0→1 44 0→1
2 0→1 26 0→1 27 0→1
A. purpurea
3 0→1 27 1 → 0 Node 81 37 0→1
E. olivacea
13 0→1 1 0→1 51 2→1
Node 46
21 1→0 55 0 → 1 Node 82 46 0→2
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 165

Taxa or
Taxa or Node Character Change Taxa or Node Character Change Character Change
Node
31 0→1 56 0→1 0 1→0
Node 47
H. maculata 68 1→0 69 3→1 Node 83 50 0→2
17 0→1 32 0→2 61 0→1
Psalmopoeus sp.
33 0→1 47 0→1 11 0→1
Node 48
Tapinauchenius
44 1→0 67 2 → 1 Node 84 44 1→0
sp.
20 0→1 52 0→1 69 3→1
Node 49
E. murinus 67 1→2 53 0→1 Node 85 9 1→2
68 1→2 24 0→1 0 0→1
Node 50
E. uatuman 21 1→2 67 0→2 Node 86 45 0→1
21 1→0 2 0→1 67 0→1
T. seladonia 26 0→1 Node 51 6 0→1 12 0→1
Node 87
35 0→1 22 0→1 56 0→1
06 0→1 6 0→1 10 0→1
Node 52
T. amma 34 0→1 69 3→2 Node 88 21 0→1
37 0→1 Node 53 66 0→1 41 0→1
T. costae 42 0→1 23 0→1 8 0→1
06 0→1 Node 54 27 0→1 Node 89 46 2→0
T. curumim
25 0→2 56 1→0 69 0→3

condition reflected in their close position on the cladogram, apart from most Avicu-
laria species (node 76 in fig. 325).
Node 78 (Fig. 325) is composed of A. hirschii, A. lynnae sp. n. and A. caei sp. n.
They are very singular species, since they have discrete elevation on prolateral tibia cov-
ered by a cluster of setae which clearly cannot be considered a branch (Fig. 309), and
process on retrolateral lobe of cymbium covered by thin setae (the homoplasy that sup-
ports the node) (Fig. 205). Both characters are distinct from those found in most species
of genus. Characters related to genitalia are also singular in these species since males
have very long embolus (A. lynnae sp. n. and A. caei sp. n.) (Figs 202, 211 respectively)
and females have twisted spermathecae (A. hirschii as in fig. 181). Despite all of these
unusual features, this clade was recovered as belonging to Avicularia in the chosen clad-
ogram as well as in the cladogram obtained using NONA (fig. 324). However, in the
cladograms obtained using concavities 1–4, this clade formed a trichotomy with Caribe-
na gen. n. and Pachistopelma + I. marcoi. Both A lynnae sp. n. and A. caei sp. n. females
are currently unknown. Even without female characters on matrix, these three species
are retrieved together in all cladograms obtained, using equal or different weights. Thus,
their close relationship is a robust hypothesis. However, the position of this clade inside
Avicularia genus can change with the future inclusion on matrix of characters related to
females of A lynnae sp. n. and A. caei sp. n. There is also the possibility that this clade
could be retrieved as a new genus.
The clade defined by node 75 (Fig. 325) and having A. taunayi, A. avicularia, A.
rufa, A. juruensis and A. variegata stat. n. is retrieved in all cladograms, with equal or
166 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Table 5. Fits, Steps and Extra Steps for characters of cladogram of Fig 325.

Character Fit Steps Extra Steps Character Fit Steps Extra Steps
0 7.5 3 2 36 10.0 2 0
1 7.5 3 2 37 6.6 4 3
2 8.5 2 1 38 7.5 3 2
3 8.5 3 1 39 8.5 2 1
4 10.0 1 0 40 10.0 1 0
5 7.5 3 2 41 7.5 3 2
6 6.6 4 3 42 6.6 4 3
7 — — — 43 8.5 2 1
8 — — — 44 4.6 8 7
9 8.5 3 1 45 7.5 4 2
10 8.5 2 1 46 4.0 12 9
11 7.5 3 2 47 10.0 1 0
12 10.0 1 0 48 7.5 4 2
13 8.5 2 1 49 7.5 3 2
14 — — — 50 5.4 7 5
15 8.5 2 1 51 6.0 6 4
16 8.5 2 1 52 8.5 2 1
17 7.5 3 2 53 8.5 2 1
18 8.5 2 1 54 7.5 3 2
19 10.0 1 0 55 10.0 1 0
20 — — — 56 5.0 7 6
21 4.6 9 7 57 8.5 3 1
22 10.0 1 0 58 10.0 1 0
23 7.5 3 2 59 10.0 1 0
24 5.4 12 5 60 — — —
25 6.0 6 4 61 10.0 1 0
26 6.6 4 3 62 10.0 1 0
27 5.4 6 5 63 10.0 1 0
28 7.5 3 2 64 8.5 2 1
29 10.0 1 0 65 — — —
30 10.0 1 0 66 10.0 1 0
31 — — — 67 5.4 7 5
32 10.0 2 0 68 4.2 10 8
33 4.0 10 9 69 8.5 4 1
34 7.5 3 2 70 8.5 4 1
35 6.0 5 4

different weights. The clade is supported by a single homoplasy, embolus tip narrow-
ing abruptly (character 49, state 1) (Figs 316–317). Despite its fragile support on the
cladogram, this group of species surely seems to be closely related, since bulb, cym-
bium, tibial apophysis and spermathecae morphology are very similar. Adding to this,
corporal size, habitat and geographic distribution area are alike.
The sister group of Avicularia is formed of Caribbean and Brazilian Atlantic rainforest
aviculariine species. The result obtained indicates three new genera in Aviculariinae
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 167

(Fig. 325), which were, herein, described. Caribena gen. n. (node 65 in fig. 325) is
composed of two Caribbean species, C. laeta comb. n. and C. versicolor comb. n. The
synapomorphies supporting monophyly of the genus (node 65 in Table 4 and in Fig.
325) are the presence of cymbium with retrolateral sharp process (character 40, state
1) (Fig. 306), and presence of slender type II urticating setae, with more than 1 mm
in length in females (character 63, state 1) (Fig. 243). This long and slender type II
urticating setae also occur in males and immatures of both species. Caribena gen. n. is the
sister group of Pachistopelma + I. marcoi in topology obtained when using concavity 6. In
the topologies obtained using concavities 1–4, the clade Pachistopelma + I. marcoi forms
a group with Caribena gen. n. as well as with the clade (A. hirschii (A. lynnae sp. n. + A.
caei sp. n.)), resulting in a trichotomy.
The other new genus, Ybyrapora gen. n. (node 68 in Table 4 and in Fig. 325),
includes small aviculariine species of Brazilian Atlantic rainforest, Y. diversipes comb.
n., Y. sooretama comb. n. and Y. gamba comb. n. Genus synapomorphies are the pres-
ence of virtually non-sclerotized spermathecae (character 36, state 2), with accentuated
outwards curvature medially (character 37, state 1, a homoplasy with Avicularia) (Figs
257–259), absent or weakly-developed tibial apophysis on leg I (character 56, state 1,
a homoplasy with many Aviculariinae taxa) (Figs 308–309, respectively). This genus
is retrieved as a monophyletic taxon and is the sister group of Iridopelma (except I.
marcoi, a special case discussed below) in all trees obtained using implied weight. In
NONA analysis, the genus was also recovered as monophyletic, but as the sister group
of (Avicularia (Iridopelma (Pachistopelma + Caribena gen. n.))).
Since its description, the classification of A. rickwesti comb. n. has been a point
of discussion. At first, it was included in Avicularia genus since its inclusion in other
aviculariine genera has no support due to the combination of characters other than
spermathecae. The completely aspinose legs, procurve first eye row, digitiform distal
article of the posterior lateral spinnerets and presence of urticating setae type II on
abdomen dorsum indicate the species should be included in either the Avicularia or
Iridopelma genera (Bertani and Huff 2013). The authors included it in Avicularia due
to biological features in common with Avicularia species and due to the fact that Iri-
dopelma only occurs in Brazil (Bertani and Huff 2013). As it was a very distinct species,
researchers were waiting to collect a male to confirm or reject the classification of A.
rickwesti comb. n. In this analysis, this species is always retrieved as a distinct and iso-
lated taxon (node 34 in Table 4 and in Fig. 325), since it has very unusual combination
of characteristics such as small embolus length in retrolateral view (character 46, state
1) (Fig. 286), embolus very flattened (character 48, state 2) and slightly curved (char-
acter 50, state 1) (Fig. 287), and presence of prolateral superior, prolateral inferior and
apical keels on palpal bulb (characters 52, 53 and 54, state 1) (Figs 285, 288). This is
the only species in Aviculariinae that has keels in the male palpal bulb. In fact, general
morphology of male and female genitalia of A. rickwesti comb. n. is unique and very
distinct from the other Aviculariinae, but is frequently found in other subfamilies, such
as Theraphosinae and Ornithoctoninae, and also in Poecilotheria spp. Antillena rickwes-
ti comb. n. switches its position in different cladograms. In those obtained with con-
168 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

cavities 1 and 2, A. rickwesti comb. n. is the sister group of Poecilotheria (Haplopelma +


Lasiodora), in a basal position of the cladogram, and out of Aviculariinae clade. Using
concavity 3, it is also retrieved in a basal position, but its sister group is ((Poecilotheria
+ Encyocratela) + (Haplopelma + Lasiodora)). In topologies obtained with concavities 4
and 5, A. rickwesti comb. n. is retrieved as the sister group of most Aviculariinae gen-
era; in the topology obtained using concavity 4 it is combined with Poecilotheria, but
it is alone in the topology obtained using concavity 5. However, in the preferred tree
obtained using concavity 6 (Fig. 325), A. rickwesti comb. n. is retrieved as the sister
group of Caribbean genera Caribena gen. n. and Brazilian Atlantic rainforest genera
Pachistopelma, Ybyrapora gen. n. and Iridopelma. Thus, based on its isolated position
on the cladogram, allied to its endemic occurrence in Dominican Republic, we decide
to erect a new genus for this species, Antillena gen. n.
The species I. marcoi is a problematic taxon. Since many important characters for
cladistic analysis such as tibial apophysis, cymbium process and palpal morphology as
well as presence of tibial apophysis in tibia II (an usual sinapomorphy for Iridopelma)
are exclusive to males, it is expected that the species would not be retrieved as part of
the genus since its male is unknown. Besides this, I. marcoi female does not have type
II urticating setae on dorsal abdomen, a characteristic of most Aviculariinae species.
Even though this type of setae is lacking in females, it could be present in males and
immatures of this species, like in Pachistopelma species. Unfortunately, since males are
unknown, this information is lacking in the analysis. On the other hand, as pointed
out by Bertani (2012), the spermathecae shape, as well as some somatic characters
indicate I. marcoi is, in fact, an Iridopelma species and the discovery of the male might
confirm its position, solving this question.
The composition of subfamily Aviculariinae (except A. rickwesti comb. n., see dis-
cussion above) is the same in all trees obtained with equal or different weights, except
in the topology obtained with concavity 4, in which Poecilotheria sp. is inside Avicu-
lariinae clade. In the chosen cladrogram (Fig. 325), Aviculariinae subfamily is com-
posed of genera Avicularia, Psalmopoeus, Tapinauchenius, Ephebopus, Stromatopelma,
Heteroscodra, Typhochlaena, Pachistopelma, Iridopelma, Antillena gen. n., Ybyrapora
gen. n. and Caribena gen. n.
The present analysis differs from the last available Aviculariinae cladogram (Bertani
2012), in which the subfamily was not retrieved as monophyletic. In that cladogram,
Aviculariinae was paraphyletic, the clade Encyocratella (Haplopelma + Poecilotheria) was
the sister-group of Aviculariinae and the clade with Psalmopoeus, Tapinauchenius and
Ephebopus spp. was the sister group of Pelinobius+ Phlogiellus. The cladogram obtained,
herein, agrees with the one presented by West et al. (2008), in which the subfamily
is retrieved as a monophyletic clade, supported by a single character—the presence of
well-developed scopulae on tarsi and metatarsi very extended laterally, mainly those of
legs I and II (Fig. 299). Here this character also supports the subfamily clade (node 85
in Fig. 325). The well-developed scopula also appears independently in Poecilotheria sp.,
but this can be a convergency related to the arboreal habit of specimens of Poecilotheria
sp. and most aviculariine taxa.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 169

In the analysis carried out here, the sister group of Aviculariinae is Phogiellus sp., a
selenocosmiine species. They share presence of straight anterior row of eyes (character
0, state 1, with a reversion in a large internal clade and a posterior change to state 1 in
Pachistopelma spp.) (Fig. 12), a developed prominence on tegulum of male palpal bulb
(character 45, state 1) (Fig. 314) and leg IV roughly the same length as leg I (character
67, state 1, with some changes in internal clades). In the cladograms obtained with
concavities 1 and 2, the sister group of Aviculariinae is Phlogiellus sp. + P. muticus.
However, Phlogiellus sp. is a controversial taxon. It is recovered in distinct positions
in different cladistic analysis. In Guadanucci (2014), it forms a clade with Lyrognathus
sp., Poecilotheria sp. (other controversial taxon) and Psalmopoeus sp., making Avicu-
lariinae paraphyletic. In West et al. (2008), Phlogiellus sp. is also the sister group of
Aviculariinae, but in Bertani (2012) this taxon is sister group of Pelinobius muticus and
both form the sister group of the clade ((Psalmopoeus + Tapinauchenius) Ephebopus),
making Aviculariinae paraphyletic. The inclusion of other Selenocosmiinae species in
the cladistic analysis should confirm if subfamily is the sister group of Aviculariinae as
suggested here.
Many clades were recovered in all cladograms despite different approaches used. The
clades that can be considered very stable are the genera Ybyrapora gen. n., Caribena gen.
n., Pachistopelma and Iridopelma (excluding I. marcoi, discussed above), all recovered as
monophyletic using both approaches. Other clades as Typhochlaena, (E. murinus + E.
uatuman + (Psalmopoeus sp. + Tapinauchenius sp.)); (Stromatopelma sp. + Heteroscodra
sp.); and (A. taunayi ((A. variegata st. n. + A. juruensis)(A. avicularia + A. rufa))) were
also recovered using equal or implied weights. The clade ((A. hirschii (A. lynnae sp. n. +
A. caei sp. n.)) is recovered in all topologies except in that obtained using concavity 5.
The decision of erecting new genera was done in order to preserve the taxonomic
stability since these clades are recoverd as monophyletic in all topologies, although
their relationship with other clades can change. Besides, the new genera have very
distinct morphologic, geographic and ecologic characteristics from Pachistopelma and
Iridopelma, which were recently revised and are well-stablished genera (Bertani 2012).
The decision of considering Caribena gen. n. species as belonging to Pachistopelma (an
endemic Brazilian genus which lives exclusively inside bromeliads) and Ybyrapora gen.
n. species as being Iridopelma species, or consider all the species of the three new genera
as Pachistopelma would cause more taxonomic problems in future analyses if they are
recovered as non-related distinct lineages.
Mygalomorph species are one of the most problematic taxa among spiders for
reliable species delimitation (Hamilton et al. 2014). Understanding species limits in
any taxonomic group is a complex and a researcher’s biased task (Satler et al. 2013),
especially when based on morphological traits. It requires examination of as many
specimens as possible, familiarity with sisters groups’ morphology and knowledge of
distribution and variation of the characters used in taxonomy of the specific group.
Mygalomorph spiders are morphologically conserved at shallow phylogenetic levels,
so, it is very difficult to delimit species’ boundaries and consequently to describe a
new species based on morphological aspects (Hamilton et al. 2014, Hendrixson et al.
170 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

2015). This is even moreso relevant when there are few specimens available to analyze,
which is the case of most mygalomorph species. It is common to decide not to describe
a new species because there is no sufficient morphological evidences to do so. Thus,
it is not a surprise when we note many new species of a previously morphologically
analyzed taxon being detected when using molecular approaches (Hedin and Carlson
2011; Hendrixson and Bond 2005; Starret and Hedin 2007). A good example of a
cryptic biodiversity discovery was by Hamilton et al. (2011), when they studied Apho-
nopelma genus. It is a widespread and specious taxon that has received little attention
and “has been largely reliant upon sparse and sometimes poorly defined morphological
data” (Hamilton et al. 2011). Avicularia seems to be exactly the same case. Avicularia
spermathecae, male palpal bulb, and tibial apophysis shape as well as somatic charac-
ters (except coloration traits) are very similar among different populations. Thus, as
occurring with Aphonopelma (Hendrixson et al. 2015), it is very probable that we can
only access the real Avicularia diversity when we use multiple approaches for an ac-
curate definition of species boundaries.
Morphologically cryptic species are an increasingly recurrent problem on tradi-
tional zoological taxonomy (Satler et al. 2013). Here we detected that boundaries of
many Avicularia species, especially Avicularia avicularia, could not be delimited using
the current morphological tools and data. It is necessary to find new morphological
characters and, combined with molecular, geographic and ecological data, to do a
more extensive and integrative analysis of Avicularia.
An accurate assessment of species-level diversity is essential not only to special-
ists; it plays an important role in studies of ecology and biodiversity and consequently
in conservation decisions and policy (Hendrixson and Bond 2005; Hamilton et al.
2011) as well as in the suitable species’ exploration for developing new technologies,
materials, and ideas to confront environmental, medical, and engineering challenges
(Wheeler et al. 2012). A more precise taxonomy and the proposal of a strongest phy-
logenetic hypothesis especially for theraphosid spiders are very urgent since constant
habitat destruction and high rate of pet trade are pressing problems for spiders popula-
tions (Bond et al. 2005; Hamilton et al. 2011).

Acknowledgements
We are especially grateful to Rick West for his continued support to our work in so many
and important ways. We would like to thank collegues who kindly loaned specimens for
study and/or allowed examination of collections under their care, as well as colleagues who
helped with field data and specimens collected: Adriano Kury (MNRJ), Alexandre Bo­
naldo (MPEG), Augusto Henriques, Regiane Saturnino and Ana Lúcia Tourinho (INPA),
Arno Lise (PUCRS), Artur Nishibe Furegatti (ZUEC), Charles Griswold (CAS), Chris-
tine Rollard (MNHN–AR), Diana Silva (UA), Eduardo Flórez (ICN–AR), Francisco
Provenzano (MIZA), Gonzalo Giribet and Laura Leibensberger (MCZ), Janet Beccaloni
(BMHN), Jason Dunlop and Anja Friederichs (MNHP), Mariajosé Deza Bouroncle (UA),
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 171

Mats Eriksson (UUZM), Norman Platnick, Lorenzo Prendini and Lou Sorkin (AMNH),
Paulo Motta (DZUB), Peter Jäger and Julia Altmann (SMF), Ricardo Pinto da Rocha
(MZUSP), Carlos Víquez (UCR), Cléria Mendonça de Moraes, Fred Pallinger, César
Alexa­ndre and Marco Antônio de Freitas. We would also like to thank the many people
whose photographs improved the paper: Saymon Albuquerque, Marlus Almeida, Marco
Antônio de Freitas, Flávio Pimenta, Antonio Tosto, Jason Newland, Tanya Stewart, Wil-
lian Lamar, Hans-Werner Auer, Martin Gamache, Jeremy Huff, Rick C. West, Ronald N.
Baxter, and the late Werner Bokermann. We thank Laboratório de Microscopia of Instituto
de Física, Universidade de São Paulo, for SEM images. We thank all curators and collegues
such as Fritz Geller-Grimm, Ingo Wendt, Volker von Wirth, Jorge Mendoza, Jorge M.
Cañas, Laura Miglio, Ray Gabriel, Martin Huber, Dirk Weinmann, Jean Michel Verdéz,
Chris Hamilton, Pascal de Bleeckere, Olivier Boilly, Patrick Maréchal, Dr. S. Füting, and
Alex Zanotti who shared information about specimens and type locations and photographs
or papers. We would like to thank Prof. Dr. Eduardo Navarro and Maria Beatriz Ribeiro for
helping with Tupi language. We thank Abel Pérez González and Fernando Pérez-Miles for
important comments on a previous draft of the manuscript as well as Brent Hendrixson,
Rick C. West, Ingi Agnarsson and an anonymous reviewer for valuable suggestions on the
manuscript. CSF would like to thank David Candiani and Nancy Lo for their hospital-
ity at Belém; Wenddi P. Burger and Iara Fino for translations, FAPESP 06/58326-5 and
CAPES for financial support and all other people named in her thesis acknowledgements,
particularly Dr. Paulo Nogueira-Neto for his support on obtaining her doctorate degree.
Support: Fapesp 2015/19976-3 and CNPq research fellow – Brazil for RB; CAPES no.
23038.00814/2011-83.

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184 Caroline Sayuri Fukushima & Rogério Bertani / ZooKeys 659: 1–185 (2017)

Supplementary material 1
Tree obtained with Piwe, all characters as non-additive and concavity 1
Authors: Caroline Sayuri Fukushima, Rogério Bertani
Data type: molecular data
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Supplementary material 2
Tree obtained with Piwe, all characters as non-additive and concavity 2
Authors: Caroline Sayuri Fukushima, Rogério Bertani
Data type: molecular data
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Supplementary material 3
Tree obtained with Piwe, all characters as non-additive and concavity 3
Authors: Caroline Sayuri Fukushima, Rogério Bertani
Data type: molecular data
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818... 185

Supplementary material 4
Tree obtained with Piwe, all characters as non-additive and concavity 4
Authors: Caroline Sayuri Fukushima, Rogério Bertani
Data type: molecular data
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Supplementary material 5
Tree obtained with Piwe, all characters as non-additive and concavity 5
Authors: Caroline Sayuri Fukushima, Rogério Bertani
Data type: molecular data
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

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