Umbyquyra gen. nov., a new tarantula spider genus from the Neotropical region (Araneae, Mygalomorphae, Theraphosidae), with a description of eight new species

Umbyquyra gen. nov., a new Theraphosinae genus with stridulatory bristles on the palpal trocanther of pedipalp trochanter and fi rst leg, is proposed. The genus differs from the other genera with stridulatory bristles on the same segments, Acanthoscurria Ausserer, 1871, Cyrtopholis Simon, 1892, Longilyra Gabriel, 2014 and Nesipelma Schmidt & Kovarik, 1996, by having a palpal bulb with a very short and acuminate embolus and four short keels; separated tibial apophysis; and female spermathecae resembling those of Cyrtopholis, with two seminal receptacles with elongated ducts emerging from a common area. Cyrtopholis palmarum Schiapelli & Gerschman, 1945 and C. schmidti Rudloff, 1996 from Brazil and Acanthoscurria acuminata Schmidt & Tesmoingt in Schmidt, 2005 from Bolivia are transferred to the new genus. The female of Umbyquyra palmarum (Schiapelli & Gerschman, 1945) gen. et comb. nov. and the male of U. schmidti (Rudloff, 1996) gen. et comb. nov. are described for the fi rst time. Cyrtopholis zorodes Mello-Leitão, 1923 is considered a junior synonym of Acanthoscurria gomesiana Mello-Leitão, 1923 and Cyrtopholis meridionalis (Keyserling, 1891) is considered a nomen dubium. Eight new species from Brazil are described: Umbyquyra paranaiba gen. et sp. nov., U. cuiaba gen. et sp. nov., U. araguaia gen. et sp. nov., U. sapezal gen. et sp. nov., U. belterra gen. et sp. nov., U. caxiuana gen. et sp. nov., U. tucurui gen. et sp. nov. and U. tapajos gen. et sp. nov. Data and maps on the geographic distribution are provided.


Introduction
The family Theraphosidae Thorell, 1869 currently has eleven subfamilies (Guadanucci 2014), four of them with species described from tropical and subtropical regions: Aviculariinae, Theraphosinae, Schismatothelinae and Ischnocolinae.The latter was established by Simon (1903) based on the presence of a divided tarsal scopula on the posterior legs.He included several genera in this subfamily, among them Cyrtopholis Simon, 1892, which he cited as being the only genus with a type IV stridulatory apparatus (stridulatory bristles on palp trochanter and leg I).Raven (1985) transferred Cyrtopholis to Theraphosinae based on the large subtegulum, presence of keels on palpal bulb and retrolateral femur IV with scopula.
While revising the four Brazilian species of Cyrtopholis, we found that none of them could be included in this genus.Studies of the type species, Cyrtopholis cursor (Ausserer, 1875), revealed differences in the spermathecae when compared to the Brazilian species.Also, a very different morphology of the male palp bulb was observed when compared to the Caribbean species of Cyrtopholis.
The study of type material and large samples of specimens from several Brazilian collections allowed us to establish the new genus Umbyquyra gen.nov., which includes eleven species: two transferred from Cyrtopholis, another from Acanthoscurria and eight new species.The female of Umbyquyra palmarum (Schiapelli & Gerschman, 1945) gen.et comb.nov.and the male of U. schmidti (Rudloff, 1996) gen.et comb.nov.are herein described for the fi rst time.Data on the geographical distribution are provided for all species.

Material and methods
The examined specimens are deposited in the following taxonomic collections (curators indicated between parentheses): Male palpal bulb terminology follows Bertani (2000) with some modifi cations in the abbreviations.Spine number and disposition follow Petrunkevitch (1925), with modifi cations proposed by Bertani (2001).All measurements are in millimeters and were taken using a Leica DFC425 stereo microscope with camera lucida.Length of leg segments were measured between joints, in dorsal view.Total body length excludes chelicerae and spinnerets.Extended focal range images were composed with Leica Application Suite version 2.5.0 (available from http://www.heliconsoft.com/heliconsoft-products/helicon-focus/).All photos of scanning electronic microscopy (SEM) were taken under high vacuum with a FEI Quanta 250 SEM at the Laboratório de Biologia Celular of the Instituto Butantan.Spermathecae were dissected and immersed in enzyme (Ultrazyme ® ) for 72 hours for soft tissue digestion to allow observation of internal structures.Geographic coordinates were obtained using a GPS, for specimens collected by us, or through information on the Museum original labels.Localities from museum samples without coordinates were georeferenced using Google Earth ® .(MNHNSD 09.1454) (Fig. 2).

Diagnosis
Umbyquyra gen.nov.presents stridulatory bristles on the palpal trocanther and fi rst leg as the genera Acanthoscurria, Cyrtopholis, Nesipelma and Longilyra.It differs from these genera by a projected and well marked cephalic region (Fig. 5) and mainly by the aspect of the genitalia: male palpal bulb compact and globose, tegulum short, embolus, palpal bulb with a short embolus presenting four keels, tibial apophysis with two branches not originating from a common base, and metatarsus I, when fl exed, touches the retrolateral branch laterally (Figs 7C, 8D); female spermathecae with SR connected to a sclerotized basal plate with conspicuous ducts (Fig. 7D).

Etymology
Umbyquyra gen.nov.originates from the Tupi Indian language and means 'pointed bird beak', relative to the aspect of the short embolus of the palpal bulb.The gender is neuter.

Description
Total length: males 18-40, females 20-44.General color is dark brown with legs lighter brown.The dorsal side of abdomen covered with long reddish or golden hairs with a dark patch (Fig 16F), ventral side of abdomen dark brown, legs with long yellowish or reddish hairs (Fig. 16F).Carapace longer than wide.Cephalic region with a projection slightly raised defi ned by the thoracic grooves.Fovea procurve (Fig. 5A-B, D), straight (Fig. 5C, E) or recurve (Fig. 5F), some species with projection over the fovea (Fig. 15E).Eye tubercle slightly raised, wider than long.Eight eyes, ringed with black.Anterior row of eyes procurved, posterior row slightly recurved or straight (Fig. 16E).Labium as wide as long, quadrate, with 130 cuspules (Fig. 6E).Endites with distinct anterior lobe, with 90-230 cuspules on internal basal angle.Sternum oval with six sigilla.Tarsal scopulae dense and integral, only tarsus IV with some bristles in longitudinal line.STC with 4-5 small teeth (Fig. 6A).Tarsi I-IV with two rows of clavate trichobotria (Fig. 6C-D).Scopulae on all tarsi and metatarsi I-II integral and only apical on metatarsi III-IV.Stridulatory bristles on palpal trochanter with 13-25 bristles and 30-38 on trocanther of leg I, sometimes with very long bristles similar as the described for Longilyra, occupying all the length of trochanter in other diffi cult to observe as in Umbyquyra acuminata (Schmidt & Tesmoingt in Schmidt, 2005) gen.et comb.nov.(Fig. 14E-F).Umbyquyra palmarum gen.et comb.nov.(Fig. 12E-F), U. tapajos gen.et sp.nov.(Fig. 24E-F), U. tucurui gen.et sp.nov.(Fig. 26E-F), U. caxiuana gen.et sp.nov.(Fig. 28E-F) have bristles long and very large, varying in size and number.PLS threesegmented with digitiform apical segment.Male palpal bulb with oval tegulum, short embolus, four or fi ve keels resembles Acanthoscurria, but differs by distribution of keels.Tibia I with two branches, not originating from common base, retrolateral and prolateral branch with strong spines; palpal bulb with oval tegulum (Figs 7A-C, 8A-D), in general with four keels (PS, PI, SA, A), but two species have a prolateral accessory keel (PAc).Umbyquyra palmarum gen.et comb.nov.have four keels.Female genitalia: spermatheca with heavy sclerotized basal area (Fig. 7D), in U. cuiaba gen.et sp.nov.basal area reduced, SR with short (Fig. 11D) or elongated (Figs 7D, 15D) ducts emerging or not from the basal area.Urticating hairs of type I and III present in both sexes.

Diagnosis
Males of U. paranaiba gen.et sp.nov.differ from those of the other species by the short and straight fovea (Fig. 5E), a palpal bulb with a short embolus, the prolateral inferior (PI) and apical keel (A) parallel near the rear distancing from each other in the middle region, becoming more evident and narrowing again at the rear (Figs 7A, 8B) and tibial apophysis with a very short prolateral branch with two strong distinct spines (Figs 7C, 8D).Females resemble those of U. palmarum gen.et comb.nov.by an SR with short ducts, but differ by a rectangular and broader sclerotized basal area (Fig. 7D).

Note
Bertani (2000: 30-31) identify a male as C. palmarum (IBSP 4730).The specimen was examined by us and identifi ed as U. paranaiba gen.et sp.nov.and we detected that the number of keels is four (PS, PI, A and SA, Fig. 8) and not three.

Etymology
The species epithet is a noun in apposition taken from the type locality.

Natural History
Very aggressive when disturbed; specimens raise the abdomen resembling the defensive display of the species of Avicularia and of some spiders of Nemesiidae, or raise pedipalps and legs I resembling species of Acanthoscurria (Fig. 18D, H; Gonzalez-Filho et al. 2012) and Aguapanela Perafán, Cifuentes & Estrada-Gomez, 2015 (Perafán et al. 2015).

Etymology
The species epithet is a noun in apposition taken from the type locality.

Female
Unknown.

Diagnosis
Males and females of U. palmarum gen.et comb.nov.differ from those of the other species by the strongly procurved fovea, like a half moon, delimitating the very long projection of the cephalic region (Fig. 5D) and male palpal bulb with well-developed A and SA keels (Figs 11A, 12A-B).Female spermathecae with antero-medially excavated base and SR with very short ducts (Fig. 11D).

Natural history
Jean Vellard, in 1945, collected two males in a rotten palm tree.According to the author, the spiders were very aggressive and no webs in the rotten palm tree, were was found.

Diagnosis
Males and females of U. schmidti gen.et comb.nov.differ from those of the other species by a carapace with large procurve fovea and a cephalic region projected over the same (Fig. 5A), less than 15 stridulatory bristles on palp trochanter, less than 30 stridulatory bristles on leg I (Fig. 14E-F), conspicuous SA and A (Fig. 13A−B) and tibial apophysis with prolateral reduced branch reduced (Figs 13C, 14D).Females differ from those of the other species by spermathecae with a cordiform basal area and SR close to each other emerging from the middle basal area (Fig. 13D).

Remark
The other female from the same locality cited in Rudloff (1996), not designated as a type, was not located.

Diagnosis
Males and females of U. acuminata gen.et comb.nov.differ from those of the other species by a procurve fovea overlapped by a large projection of the cephalic region (Fig. 15E) and by a retrolateral branch of the tibial apophysis with three strong spines (Fig. 15C).Female spermathecae have a cordiform basal area resembling those of U. schmidti gen.et comb.nov., but differs from this species by the RS being more apart when compared to the other species (Fig. 15D).

Diagnosis
Males and females of Umbyquyra cuiaba gen.et sp.nov.resemble those of U. palmarum gen.et comb.nov.by a large procurved fovea delimitating the projection of the cephalic region (Fig. 5D) and by a male palpal bulb with the shape and size of keels, but differ by the tegulum being more compact (Figs 16A-B, 17A-C) and by females having a spermathecae with a very narrow basal area (Fig. 16D).

Etymology
The species epithet is a noun in apposition taken from the type locality.

Natural history
This species is very aggressive, raising the abdomen or legs I when disturbed (Fig. 18D, H).The female from Parque Nacional da Chapada dos Guimarães (IBSP 167419) was collected in a dirt hill, inside a hole with a 3 cm opening diameter which did not contain any silk.20D) and a retrolateral branch with two strong spines, and by a straight fovea without any projections on the carapace (Fig. 5E).

Etymology
The species epithet is a noun in apposition taken from the type locality.

Female
Unknown.

Diagnosis
Males and females of Umbyquyra belterra gen.et sp.nov.differ from those of all other species by a recurved fovea (Fig. 5F) and a very long prolateral inferior keel of the male palpal bulb which extends for more than half of the prolateral face of the tegulum (Fig. 22C).This species presents fi ve keels, including the PAc keel near the PS keel.This is a character shared only between two species of Umbyquyra gen.nov.Females differ from those of the other species by a spermathecae with SR curved sideways, with long ducts and apart from each other by more than fi ve times their diameter (Fig. 21D).

Remarks
The female is in bad condition, so total length and carapace could not be measured.

Diagnosis
Males of Umbyquyra tapajos gen.et sp.nov.resemble those of U. belterra gen.et sp.nov.by the presence of a PAc (Fig. 23C), but differ by a more compact tegulum and keels being parallel to each other (Figs 23A-C, 24A-C).Umbyquyra tapajos gen.et sp.nov.presents a short and straight fovea while U. belterra have a recurve fovea.

Etymology
The species epithet is a noun in apposition taken from the type locality.

Female
Unknown.

Etymology
The species epithet is a noun in apposition taken from the type locality.

Diagnosis
Species of Umbuquyra caxiuana gen.et sp.nov.are distinguished from those of the other species by the large and procurve fovea (Fig. 5C), male palpal bulb with a shortened and compact tegulum, a superior keel very evident being parallel to prolateral accessory keels (Fig. 28B).The female can be distinguished by a spermathecae with globular SR, close to each other emerging from a common base forming a V shape (Fig. 27D).

Remark
The original description indicates MZSP as the type repository; however, no specimen of Cyrtopholis zorodes was located.Bücherl et al. (1971: 124) and Silva-Moreira et al. (2010: 68) agree that the specimen MNRJ 29, holotype ♀ from São Paulo, in the state of São Paulo, Brazil is the true holotype.The holotype is in bad condition without SR and legs.Bücherl mentioned stridulatory bristles on the palp trochanter and fi rst leg.The aspect of the SR and the type locality (São Paulo, state of São Paulo, Brazil) allow us to establish Cyrtopholis zorodes as a junior synonymy of Acanthoscurria gomesiana Mello-Leitão, 1923.

Discussion
Spiders of Theraphosinae in general present few morphological characters and several taxonomic problems (Raven 1990).Simon (1903) proposed the use of the stridulatory bristles, present in some Theraphosinae (Aviculariinae) to distinguish genera, according the localization on legs segments and pedipalps.He established four types of stridulatory bristles and the type IV, on the retrolateral face of the pedipalp and prolateral face of trochanter of leg I are nowadays recognized as distinctive characters in four genera: Acanthoscurria, Cyrtopholis, Nesipelma and Longilyra.According to this character, three species from South America were described: Cyrtopholis palmarum, C. schmidti and Acanthoscurria acuminata.Fig. 30.Geographic distribution records of species of Umbyquyra gen.nov.: U. acuminata (Schmidt & Tesmoingt in Schmidt, 2005) gen.et comb.nov., U. palmarum (Schiapelli & Gerschman, 1945) gen. et comb. nov., U. araguaia gen. et sp. nov., U. paranaiba gen. et sp. nov. and U. sapezal gen. et sp. nov.Historically, the relationship between Cyrtopholis and Acanthoscurria has been referenced in the literature (Pérez-Miles et al. 1996;Bertani 2001), but Nesipelma and Longilyra have never been phylogenetically tested.Umbyquyra gen.nov.presents the same disposition of the stridulatory bristles, showing their relationship with these genera and this character could be a synapomorphy for this group within Theraphosinae.
It is noteworthy that the three species mentioned above, Umbyquyra palmarum gen. et comb. nov., U. schmidti gen. et comb. nov. and U. acuminata gen. et comb. nov.show a prominent projection of the cephalic region (Fig. 15E), another characteristic common in species of the Antillean genus Cyrtopholis (see Schmidt 2003: 268), were mistakenly included in Acanthoscurria.This character should be treated as plesiomorphic in Theraphosidae, as it also occurs in Ceratogyrus Pocock, 1897, an African Harpacterinae (De Wet & Dippenaar-Schoeman 1991;Peters 2000) while Umbyquyra gen.nov. is included in Theraphosinae.Unfortunately, this character, highly variable among species, could hardly be used as diagnostic for these genera (Gallon, 2001).These structures range from almost absent to protruding projections on the back of the carapace (Fig. 5; De Wet & Dippenaar-Schoeman 1991: fi g. 4A-H) and their homology has not yet been studied among the various occurrences in Theraphosidae.
On the other hand, this study shows that in Umbyquyra gen.nov., as in general for Theraphosidae, the shape of the palp of the males and their structures are fundamental in the determination of the genera and their relations (Bücherl 1957;Pérez-Miles et al. 1996;Bertani 2000;Pérafan & Pérez-Miles 2014;Ortiz & Francke 2015).The pattern presented by Umbyquyra gen.nov. is very different from that of Cyrtopholis and Nesipelma, where the male palp bulb lacks an apophysis and has an elongated and tapered embolus (Figs 2A-B, 4C).In Longylira the males are still unknown, only females have been described.The palps of the males of Umbyquyra gen.nov.appear closer to Acanthoscurria since they Fig. 31.Geographic distribution records of species of Umbyquyra gen.nov.: U. cuiaba gen.et sp.nov.and U. schmidti (Rudloff, 1996) gen. et comb. nov.present an apophysis, but are distinguished by the bulb of the palp with an acuminate and protruding bulb, and two tibial apophyses, whereas in the species of Acanthoscurria the apex of the embolus forms a spoon or furrow and presents only one tibial apophysis (Fig. 1C).The female genitalia in this group appears to be more conservative, but in Umbyquyra gen.nov. the seminal receptacles of the spermathecae have short ducts emerging from a large, sclerotic base (Fig. 7D).
Although we found some putative relationships, a phylogeny of this group is still necessary, but it would be equally important to review of Cyrtopholis, a genus with a wide diversity in the Antilles, and to increase the knowledge of the monotypic Nesipelma and Longilyra, the latter known only by females.
Umbyquyra tucurui gen.et sp.nov.resemblethose of U. paranaiba gen.et sp.nov.by a straight fovea (Fig.5E), but this species can be distinguished from U. paranaiba gen.et sp.nov.and the other species by a retrolateral branch of the tibial apophysis with 5-6 strong distal spines (Figs 25C, 26D), and prolateral inferior and accessory keels being parallel to each other (Figs25A, 26A-B).The female spermathecae resemble those of Umbyquyra paranaiba gen.et sp.nov., but in U. tucurui gen.et sp.nov. the base is narrower (Fig.