Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818 (Araneae, Theraphosidae, Aviculariinae) with description of three new aviculariine genera

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 history 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 characters. 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, Ybyrapora 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 medially, 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 avicularia (Linnaeus, 1818), Avicularia glauca Simon, 1891, Avicularia variegata (F. O. Pickard-Cambridge, 1896) stat. n., Avicularia minatrix Pocock, 1903, Avicularia taunayi (Mello-Leitão, 1920), Avicularia juruensis Mello-Leitão, 1923, Avicularia rufa Schiapelli & Gerschman, 1945, Avicularia purpurea Kirk, 1990, Avicularia hirschii Bullmer et al. 2006, Avicularia merianae sp. n., Avicularia lynnae sp. n., and Avicularia caei sp. n.. Avicularia species are distributed throughout Mexico, Costa Rica, Panama, Trinidad and Tobago, 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 laeta (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 Avicularia avicularia; Avicularia bicegoi Mello-Leitão, 1923 is a junior synonym of Avicularia 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 Simon, 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 dubium; 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 nomen 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 nomen 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 Avicularia avicularia, Avicularia variegata stat. n., Avicularia juruensis, Caribena laeta comb. n., Euathlus affinis comb. n. and a neotype is established for Caribena versicolor comb. n.


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 confusion 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 Avicularia fasciata (Latreille, 1804). Under the name Avicularia canceridea he listed Aranea avicularia, a species described by Linnaeus (1758) in Systema Naturae.
At that time, most mygalomorph spiders were described in the genus Mygale Latreille, 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) followed 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-  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). Apparently, 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 little bird-plus aria, meaning "which refers to" (Beechhold 1997apud Smith 2000a. Thorell (1870) explained the genus name as a derivation of avicularium, meaning birdkeeper, 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 Zoological Nomeclature (ICZN 1928) was the genus Avicularia included in the Official List of Generic Names. The Direction 67 of this same Commission (ICZN 1957) established that the specific name of the type species of Avicularia is avicularia Linnaeus, 1758 as published in the combination Aranea avicularia. 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 despite 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 transferred 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 plantation, resulting in the largest ecological dataset on A. avicularia and one of the most detailed studies of aviculariine ecology. Stradling (1994) presented important conclusions 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 maturation 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 (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, andHaplopelma 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 recorded from Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, Bolivia, and Chile as well across the Caribbean archipelago including Cuba, Dominican Republic, Puerto Rico, Martinique and Guadeloupe, 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 understudied 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 descriptions 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 related 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 collections 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                               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*.
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 separately from its "character-variable" characters to preserve transformational independence 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 cladistics. In order to find new characters, we have tried many approaches. We took measurements of leg and palp articles such as tibia, metatarsi and femora, and established 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 . 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.
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.
Most outgroup species have only slight variations in the general color and abdominal 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 maturity, 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 carapace, 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". 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.
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. 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.
Males of Y. sooretama comb. n., Y. gamba comb. n., A. lynnae sp. n., and I. hirsutum have a single dorsal central stripe on abdomen dorsum. This pattern is very distinct from those found in females of these same species and also very distinct from patterns commonly found in other aviculariine males. Patterns other than dorsal central 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.
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.
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 normally homogeneously dark or present a gradual lightening toward apex. But, in some species, these setae are dark and present a contrasting whitish apex.
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.
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 . It is not a well-defined lobe as in other taxa, such as Y. sooretama comb. n. (Fig. 257) or in A. taunayi . Due to this, we considered C. laeta comb. n. as having state 0. State 1 is widespread among theraphosid genera.
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 midwidth 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 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 sclerotization degree, even in adult females , and they were coded as state 2. Melloina 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 "?".
The presence of a process on the cymbium was considered by Raven (1985) a synapomorphy of Aviculariinae. However, more recent analyses showed that the structure 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  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 . 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). (1) thick (Fig. 307).

Cymbium
Thin setae covering the retrolateral process are found both in some Avicularia species 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.
The apex of the process on retrolateral lobe is usually rounded, except in both species of Caribena gen. n., in which the process apex is sharp. 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. 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 . 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 prolateral 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 reexamine it in the proper position. (2) well-developed (Fig. 315).

Subtegulum
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 prominence is weakly-developed (Fig. 313), it was coded as state 0, as in H. rondoni, P. muticus, 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).
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., Pachistopelma 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). Specimens of Y. sooretama comb. n., Y. gamba comb. n., Typhochlaena spp., P. muticus, Phlogiellus sp., Poecilotheria sp., E. olivacea, Heteroscodra sp., and Stromatopelma 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. 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 (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).

Bulb
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 Pachistopelma 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).
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-developed 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. 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 "?".
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).
When urticating setae are analyzed, the gender of specimens examined should be considered because males have setae significantly longer than females (Stradling 1978, Bertani andGuadanucci 2013).
Distribution and habitat. Costa Rica, Panama, Trinidad and Tobago, Venezuela, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Bolivia and Brazil. In Brazil, it occurs in the states of Roraima, Amapá, Amazonas, Pará, Maranhão, Tocantins, 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 Veracruz, 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.
Ausserer (1871) considered Avicularia canceridea Lamarck, Aranea avicularia Linnaeus, 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 consideration that Aranea avicularia is a very old species described by Linnaeus in the 10 th 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) approved 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).
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 specimen 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 lectotype, 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.
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 illustrated 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, 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 information 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 deposited 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 properly 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 those seen in specimens found in that area. Therefore, herein, we establish the specimen 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 mygalomorph taxonomy.

Key to Avicularia species
Male of A. glauca and female of A. caei sp.n. and A. lynnae sp. n. are unknown. 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 Abdomen dorsum with lateral stripes (Fig. 218) and tegulum with weaklydeveloped prominence (Fig. 313)  All legs always with grizzled setae, distal femora, tibiae and metatarsi with vivid yellow rings and grey guard-setae on black abdomen (Fig. 300 Tegulum with developed prominence (Fig. 95), abdomen with three pairs of light brown spots extending from the dorsum to lateral region (Fig. 103)

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 description (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 description 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 examined 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 scopula, 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. avicularia (Linnaeus, 1758).
Avicularia cuminami holotype has tarsal and metatarsal scopulae laterally expanded, anterior row of eyes procurve and type locality is state of Pará, Brazil. These characteristics 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).
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.
Type II urticating hairs: 0.62-0.69 long, 0.017-0.019 wide. Spermathecae (Fig. 21): two completely separated, not-twisted and 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. 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 apex on posterior legs. Leg rings on distal femora, tibiae and metatarsi whitish. Abdomen dorsum with long reddish brown guard-setae with pink sheen grouped on lateral dorsal anterior areas and dark short body setae. Ventral abdomen brown.
Chelicerae: basal segment with 10 teeth in row and some small teeth on promargin. Sternum: 8.58 long, 7.35 wide. Sigilla: three pairs, all rounded and large, less than one diameter from margin.
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 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 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.  Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi contrasting with other lighter articles and abdomen dorsum reddish, with dorsal central longitudinal black stripe disconnected from transversal black stripes (Figs 37,44,47). When mature, both males and females lose this pattern.  (Linnaeus, 1758) and Avicularia rufa Schiapelli & Gerschman, 1945. Distribution. Venezuela, Trinidad and Tobago, Guyana, Suriname, French Guiana, 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 understand 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-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 pineapple plant and among the palm-leaf thatch of native houses. Villa (2004) observed in areas at Tambopata district in Peru retreats of A. avicularia 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-   Occurrence area ( Fig. 51 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), different 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. 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 description (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.  (Figs 54-57). They differ from females of A. taunayi by lacking lobes on spermathecae. 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 (morphotype 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) 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).
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 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.
Chelicera: basal segment with 9 teeth and some small teeth on promargin. Sternum: 9.93 long, 7.45 wide. Sigilla: only posterior evident, rounded, less than one diameter from margin.
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi contrasting with other lighter articles and abdomen dorsum reddish, with dorsal central longitudinal black stripe disconnected from transversal black stripes (Fig. 66). When mature, both males and females lose this pattern.
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 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 characters 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 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. Pocock, 1903 Figs 19, 78-90, 312 Avicularia minatrix Pocock, 1903 (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. minatrix 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) 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 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 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.
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 immatures. 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 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) ( Fig. 98). They differ from A. purpurea and A. merianae sp. n. by well-developed process 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 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) : two completely separated 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.
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 central 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).
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 species of this genus in the bioma Cerrado (savannah) (Bertani and Motta 2013).

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 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).    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 globous bulb with small subtegulum and well-developed prominence on tegulum. Embolus: not flattened, without keels, 5.02 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 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).
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 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.
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). Females 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 grizzled guard-setae and short body setae with intense golden sheen on legs and carapace. 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 dorsal 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 midwidth 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 whitish apex on legs, spermathecae with midwidth expanded, palpal bulb with well-developed 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 central longitudinal black stripe disconnected from transversal black stripes (Fig. 117). When mature, both males and females lose this pattern.

Avicularia rufa
Chelicera: basal segment with 10 teeth in a row and some small teeth on promargin. Sternum: 7.75 long, 7.45 wide. Sigilla: rounded posterior, less than one diameter from margin; other sigilla not evident.
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 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.
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 promargin. Sternum: 7.72 long, 6.44 wide. Sigilla: anterior rounded, middle fusiform, both less than one diameter from margin.
Color pattern ontogeny. Brownish juveniles lacking metallic sheen, black tarsi contrasting with other lighter articles (Fig. 142) and abdomen dorsum reddish, with dorsal central longitudinal black stripe disconnected from transversal black stripes (Fig. 143). When mature, both males and females lose this pattern (Figs 145-146).

Avicularia purpurea
Chelicera: basal segment with 14 teeth and some small teeth on promargin. Sternum: 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.
Type II urticating setae: 0.53-0.58 long, 0.011-0.014 wide. Spermathecae (Fig. 154): 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 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 carapace 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.
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 distributed, 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).
Chelicera: basal segment with 9 teeth and some small teeth on promargin. Sternum: 5.01 long, 3.93 wide. Sigilla: only posterior pair evident, rounded, less than one diameter from margin.
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. 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 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 orange 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.
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.).
Chelicera: basal segment with 12 teeth and some small teeth on promargin. Sternum: 7.04 long, 6.04 wide. Sigilla: anterior pair not evident, middle fusiform, posterior ellipsoidal, in a 45°angle, both less than one diameter from margin.
Tibial apophysis : a single branch on prolateral leg I, with welldeveloped base and grouped spiniform setae distally. Male metatarsus I touches retrolaterally 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 guardsetae. 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.
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.
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 central longitudinal black stripe connected only with anterior pair of transversal black stripes. When mature, both males and females lose this pattern.
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. Abdomen 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).
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 difference 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.
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.
Tibia I with discrete elevation covered by cluster of setae in apical portion, on prolateral side (Figs 215-217). 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.

Remarks.
The syntypes are specimens of subfamily Aviculariinae because they have tarsi with well-developed scopulae, with a spatulated aspect (Fig. 219) (Strand, 1908) nomen dubium, holotype juvenile female, from El-Tor, Egypt (SMF 2660), spermathecae 223-224 Thrixopelma aymara (Chamberlin, 1916)  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)  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 spermathecae (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.

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 consider, herein, Ischnocolus gracilis Keyserling, 1891 a nomen dubium.

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 characteristic Grammostola Simon, 1892 species from Brazil. Thus, we transfer Avicularia subvulpina to Grammostola, making the new combination Grammostola subvulpina (Strand, 1906) comb. n.
Even though it is possible to identify the specimen as belonging to the genus Avicularia, 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 spermathecae morphology is similar in most Avicularia species (Fig. 222). Furthermore, there is no indication of where the specimen came from. Thus, Avicularia arabica (Strand, 1908) is, herein, considered nomen dubium.
Thrixopelma aymara (Chamberlin, 1916) 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.
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 paratype 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. aurantiaca 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 identification. Thus, we consider the following species all nomina dubia: Araneus hirtipes Fabricius, 1793 nomen dubium; Avicularia testacea (C. L. Koch, 1841)  The holotypes of the species described by Tesmoingt (1996abc;1999ab) are not at Remarks. Strand (1907e) indicated the type is deposited in Lübeck Museum. However, 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 andAvicularia surinamensis Strand, 1907 are, herein, considered as nomina dubia.
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) described 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.
Color pattern ontogeny. Juveniles with metallic sheen, all articles with same blackish color (Fig. 240), and abdomen dorsum with dorsal central longitudinal black stripe disconnected from transversal black stripes. When mature, they lose this pattern .
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.

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 spermathecae with accentuated outwards curvature medially and by lacking intumescence on apex (Fig. 230). Males differ from males of C. laeta comb. n. by developed prominence 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). 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, Walckenaer (1837) did not include Cayenne nor Brazil in the distribution area, only Guadeloupe and Martinique. Concerning the description, the female corresponds to C. versicolor 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. 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 Type II urticating setae: 1.33-1.58 long, 0.006-0.009 wide. Spermathecae (Fig. 230): 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. 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 abdomen 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.
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. Diagnosis. Ybyrapora gen. n. can be distinguished from all other aviculariine genera, except 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 spermathecae, 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 process bearing thin setae on retrolateral lobe of cymbium in males. From Avicularia females it can be distinguished by having virtually non-sclerotized spermathecae . 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.
Key for Ybyrapora gen. n. species

Diagnosis (amended from Bertani and Fukushima 2009).
Females of Y. sooretama comb. n. resemble those of Y. gamba comb. n. by elongated spermathecae with accentuated 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 bearing 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).
Tibial apophysis in leg I absent (Figs 265-267 . 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 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). Diagnosis, material examined, description, color pattern ontogeny, distribution and natural history. See Bertani and Fukushima (2009) (Fig. 256).
Complementary description. : 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 Tibia I with discrete elevation covered by a cluster of setae in apical portion, on prolateral side (Figs 281-283).
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.  (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). Additionally, 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).
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 guardsetae. 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 ventrally 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.

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 rickwesti 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 Aviculariinae. 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 medial 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 topology. The topologies obtained using concavities 1 (Suppl. material 1: concavity 1) and 2   (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 Poecilotheria 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.
The species A. minatrix was retrieved as the most basal species in Avicularia. Avicularia purpurea and A. merianae sp. n. are very similar species regarding body dimensions, spermathecae and cymbium morphology as well as geographic distribution area,  condition reflected in their close position on the cladogram, apart from most Avicularia 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 covered 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 supports 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 cladogram 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 Caribena 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  (Figs 316-317). Despite its fragile support on the cladogram, this group of species surely seems to be closely related, since bulb, cymbium, 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 ( 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 presence 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 Iridopelma 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 isolated 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 (character 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 rickwesti comb. n. switches its position in different cladograms. In those obtained with con-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 genera; 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 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.
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 Aviculariinae 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.
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. 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 Aphonopelma 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 characters (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 accurate definition of species boundaries.
Morphologically cryptic species are an increasingly recurrent problem on traditional 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 specialists; 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 phylogenetic hypothesis especially for theraphosid spiders are very urgent since constant habitat destruction and high rate of pet trade are pressing problems for spiders populations (Bond et al. 2005;Hamilton et al. 2011).