The Neotropical genus Cyriocosmus Simon, 1903 and new species from Peru, Brazil and Venezuela (Araneae: Theraphosidae: Theraphosinae)

Abstract Cyriocosmus peruvianus sp. nov., C. itayensis sp. nov., C. aueri sp. nov., C. giganteus sp. nov. from Peru, Loreto region, C. hoeferi sp. nov. from Brazil, C. williamlamari sp. nov. and C. nicholausgordoni sp. nov. from Venezuela are described, illustrated, diagnosed and both sexes, if known, keyed. New Peruvian species can be distinguished from all congeners, except C. bertae Pérez-Miles, 1998 and C. pribiki Pérez-Miles and Weinmann, 2009, by the uniformly coloured carapace and abdomen without stripes and by the short paraembolic apophysis in male palpal bulb. The female of Cyriocosmus ritae Pérez-Miles, 1998 is described for the first time from a known Peruvian population near Iquitos.


Introduction
Tobago, Trinidad and Venezuela, including Isla Margarita ( Figure 36). Spiders of the genus inhabit tropical and non-tropical areas, lowland rainforests as well as high-altitude forests at the elevation of 3000 m asl Pérez-Miles and Weinmann 2009).

Description
The genus Cyriocosmus comprises small to medium-sized spiders, with total length 10-32 mm, excluding chelicerae and spinnerets. Carapace oval, uniformly coloured or with bicoloured pattern. Caput moderately domed. Eye tubercle oval, flattened, distinctly wider than longer, with eight eyes, anterior eye row slightly procurved, posterior row slightly recurved in dorsal view, a group of strong setae present on the median anterior margin of the tubercle. Clypeus absent to very narrow. Fovea transverse, straight to slightly procurved. Chelicerae without rastellum and stridulatory bristles, with teeth on promargin (7)(8)(9)(10), first basal teeth are complemented with granulation. Labium domed, wider than longer, with 30-100 cuspules in anterior third, maxillae with 100-360 cuspules in basal half on ventral side, maxillary lobe pronounced into conical process. Variability in number of labial and maxillary cuspules in Cyriocosmus perezmilesi was described by Kaderka (2010). Labiosternal groove distinct, shallow and flat, with two slightly separate or joined elongated sigilla. Sternum oval, with three pairs of small, oval sigilla located near coxae III, coxae II and coxae I, posteriorly separated from the margin approximately by their own diameter. Legs uniformly hirsute, with (Figures 12b,17b,34) or without (Figures 1b, 7) whitish or yellowish longitudinal striation on dorsal side. Leg pattern (from longest to shortest): I>IV>II>III in Cyriocosmus ritae or IV>I>II>III in all congeners. Leg segments: generally uniform to slightly incrassate on femur III. Incrassate tibia I is present in males of C. ritae only (Figure 24b).
Dense scopulae on ventral side of all tarsi, metatarsi partly scopulate, scopulae more extended on anterior than on posterior legs. Tarsal scopulae I, II usually undivided, on tarsi III, IV usually divided by longitudinal band of setae. Retrolateral side of femur IV and prolateral side of femur I without pad of plumose setae. Maxillary and trochanteral stridulatory setae or bristles absent. Spination as in species descriptions. Dorsal side of all tarsi with two irregular longitudinal rows of very short claviform trichobothria. Paired tarsal claws without teeth, third claw absent in all tarsi. Claw tufts dense, bilobate, present on all tarsi.
Females with spermathecae composed of two separated spiral seminal receptacles, distally terminating with caliciform ( Figure 29a, f, g) or globular extension (Figure 29b, c, e, h), with or without sclerotized basal plates. If present they can be flat (Figure 29a, f) or convex (Figures 29c,g,35a,b).

Etymology
The specific name is derived from a distribution area covering a part of the Peruvian Amazon rainforest.

Diagnosis
Cyriocosmus peruvianus sp. nov. can be distinguished from all other congeners, except C. itayensis sp. nov., C. aueri sp. nov., C. giganteus sp. nov., C. bertae and C. pribiki, by its uniformly coloured carapace and abdomen and by the short paraembolic apophysis in male palpal bulb. It differs from C. itayensis sp. nov., C. aueri sp. nov., C. bertae and C. pribiki by having the caliciform extension in female seminal receptacles, from C. itayensis sp. nov. also by the different morphology of male palpal bulb, from C. aueri sp. nov. also by the different coloration and the flat basal plates of female seminal receptacles, from C. giganteus sp. nov. by having the prolateral superior keel in male palpal bulb and the embolus approximately twice as long, from C. bertae also by having a smooth prolateral superior keel and by the presence of a retrolateral process on male palpal tibia, from C. pribiki also by the different coloration, flat basal plates of female seminal receptacles, cymbium without spiniform setae and by the metatarsal flexion between the two tibial apophyses.
Distribution (Figures 36,37) Known only from Maynas province in Loreto region, Rio Nanay near Iquitos in Peru. The region is a part of the Amazonian lowland originally covered with rainforest.
Palpal organ as in Figure 25a and b, embolus with short PA and with smooth PS keel. Tegulum with distinct granulated TP, projecting prolaterally. Retrolateral face of cymbium without basal field of spiniform setae. Retrolateral face of palpal tibia with distinct subapical protuberance covered with cluster of numerous spiniform setae (Figure 2e). Two unequal subapical apophyses are present on tibia I ( Figure 27a): a longer retrolateral tibial apophysis with very short apical spine, a shorter prolateral tibial apophysis with single, well-developed retrolateral spine at base and approximately of the same length as prolateral tibial apophysis. Metatarsus I not sigmoidly curved and without basal or median protuberance on retrolateral face. Metatarsus I flexion is between both tibial apophyses.

Variability
The variability in morphology of male palpal bulbs is shown in Figure 30, in the shape of spermathecae in Figure 33a-d. The variability in the length of the carapace, the number and the arrangement of the cheliceral teeth, the number of labial and maxillary cuspules, the leg pattern and the spination of tibial apophyses is shown in Table 3.

Etymology
The specific name is derived from the type locality near the Itaya River in the Peruvian Amazon.

Diagnosis
Cyriocosmus itayensis sp. nov. can be distinguished from all other congeners, except C. peruvianus sp. nov., C. aueri sp. nov., C. giganteus sp. nov., C. bertae and C. pribiki, by its uniformly coloured carapace and abdomen and by the short paraembolic apophysis in the male palpal bulb. Differs from C. peruvianus sp. nov. by having the globular extension in female seminal receptacles, from C. aueri sp. nov. by the different coloration and by having the flat basal plates in seminal receptacles, from C. giganteus sp. nov. by having the prolateral superior keel and approximately two times longer embolus in male palpal bulb, from C. bertae by having a smooth prolateral superior keel and by the presence of the retrolateral process on male palpal tibia, from C. pribiki by the different coloration, cymbium without spiniform setae and by the metatarsal flexion between the two tibial apophyses.

Distribution (Figures 36, 37)
Known only from Peru, Maynas province in Loreto region, Rio Itaya near Iquitos. The region is originally covered with lowland rainforest.

Variability
The variability in morphology of male palpal bulbs is shown on Figure 31e-h, in the shape of spermathecae on Figure 33e, f. In adult females the seminal receptacles have slightly convex basal sclerotized plates which are less extended and less sclerotized in juvenile specimens. The variability in the length of the carapace, the number and the arrangement of the cheliceral teeth, the number of labial and maxillary cuspules, the leg pattern and the spination of tibial apophyses is shown in Table 3.

Etymology
The specific name is a patronym in honour of Hans-Werner Auer, who found all new species here described from Amazon rainforest in the Loreto region, Peru.

Diagnosis
Cyriocosmus aueri sp. nov. can be distinguished from all other congeners, except C. peruvianus sp. nov., C. itayensis sp. nov., C. giganteus sp. nov., C. bertae and C. pribiki, by its uniformly coloured carapace and abdomen and by the short paraembolic  apophysis in male palpal bulb. Differs from C. peruvianus sp. nov. and C. itayensis sp. nov. by the different coloration and by the spermathecae with the globular extension of seminal receptacles and convex basal plates, from C. giganteus sp. nov. by the different coloration and the different shape of male palpal bulb having the prolateral superior keel, from C. bertae by the different coloration, by having a smooth prolateral superior keel and of a different shape, from C. pribiki by the different coloration, cymbium without spiniform setae and by the metatarsal flexion between the both tibial apophyses.
Palpal organ as in Figure 25e, f, embolus with short PA, smooth PS keel is fused with PA. Tegulum with distinct granulated TP, projecting prolaterally. Retrolateral face of cymbium without basal field of spiniform setae. Retrolateral face of palpal tibia with indistinct subapical protuberance covered with cluster of numerous spiniform setae ( Figure 10e). Two unequal subapical apophyses are present on tibia I ( Figure 27c): a longer retrolateral tibial apophysis with very short, stout spine at apex, a shorter prolateral tibial apophysis with single, well-developed retrolateral basal spine reaching approximately two-thirds of its length. Metatarsus I not sigmoidly curved and without basal or median protuberance on retrolateral face. Metatarsus I flexion is between both tibial apophyses.

Variability
The variability in morphology of male palpal bulbs is shown on Figure 31a-d, and in the shape of spermathecae on Figure 33g-h. The variability in the length of the carapace, the number and the arrangement of the cheliceral teeth, the number of labial and maxillary cuspules, the leg pattern and the spination of tibial apophyses is shown in Table 3.

Etymology
The specific name is a patronym in honour of Dr Hubert Höfer, who found and photographed this new species in the Amazon rainforest near Manaus, Brazil.

Diagnosis
Cyriocosmus hoeferi sp. nov. can be distinguished from all other congeners by its black carapace with two lateral whitish stripes, abdomen with four lateral stripes, joined neither basally nor apically with urticating setae patch, and by the short and wide paraembolic apophysis in male palpal bulb. The spermathecae are without sclerotized basal plates, both spiral necks are basally almost joined.

Distribution (Figures 36, 38)
Known only from the two localities in the Central Brazilian Amazon. The region is originally covered with lowland rainforest.
Palpal organ as in Figure 26g, h, embolus with short triangular PA reaching approximately half of embolus, smooth triangular PS keel is fused with PA. Tegulum with distinct granulated TP, projecting prolaterally. Retrolateral face of cymbium without basal field of spiniform setae, prolateral cymbial lobe longer than retrolateral one. Retrolateral face of palpal tibia with distinct subapical protuberance not covered with cluster of numerous spiniform setae but with setae only (Figure 13g). Two unequal subapical apophyses are present on tibia I (Figure 28a): a longer retrolateral tibial apophysis with short, stout subapical spine, a shorter prolateral tibial apophysis with single, well-developed retrolateral spine at base and approximately of the same length as prolateral tibial apophysis. Metatarsus I not sigmoidly curved and without basal or median protuberance on retrolateral face. Metatarsus I flexion is between both tibial apophyses.
Coloration and covering setae: dorsal view (Figures 12b, 14a): carapace black with two narrow whitish lateral stripes, joined neither anteriorly nor posteriorly, coxae, trochantera, femora, patellae, tibiae and metatarsi black with whitish longitudinal stripes. Tarsi black. All femora with two longitudinal stripes without covering setae, one of them on retrolateral face, the other on dorsal face. Patellae I, II with two longitudinal stripes, patellae III, IV with unequal diagonal stripes without covering setae. Femur I prolaterally bare. Abdomen black (Figure 14d

Variability
The variability in morphology of male palpal bulbs is shown on Figure 26e-h. The PS keel in the male from Rio Tarumã is not so pronounced as in the male holotype where it is subtriangular. Further comparative study of the specimens from both populations isolated by Rio Tarumã is necessary. The variability in the length of the carapace, the number and the arrangement of the cheliceral teeth, the number of labial and maxillary cuspules, the leg pattern and the spination of tibial apophyses is shown in Table 3.

Etymology
The specific name is a patronym in honour of Dr William Lamar, who found this new species in Venezuela whilst filming with the BBC.

Diagnosis
Cyriocosmus williamlamari sp. nov. can be distinguished from all other congeners by the reddish-brown carapace with the black caput, the abdomen with four lateral stripes, apically not joined with urticating setae patch, three clear stripes are basally joined, the fourth stripe is located near spinnerets, and by the presence of the dark longitudinal ventral band. Spermathecae have very well-developed convex basal plates.
Distribution (Figure 36) Known only from the type locality.
Coloration and covering setae: dorsal view (Figures 15, 16a): carapace reddish-brown, and covered with golden pubescence, with black caput, coxae and trochantera reddishbrown, chelicerae dark, femora black, patellae, tibiae, metatarsi and tarsi dark, dorsally with whitish longitudinal stripes. Patellae I, II and palpal patella with two unequal parallel longitudinal stripes without covering setae, patellae III, IV with single diagonal stripe. Palpal femur and femur I prolaterally bare. Abdomen (Figure 16d) covered with short black setae, intermixed with long, pale setae in posterior part, except central reddish-brown glossy patch in shape of heart and four lateral stripes on each side, not reaching the patch, the first three lateral stripes basally joined, the fourth pair basally not joined. Length of central patch: 2.2, width 1.8. Ventral view ( Figure 16b): labium, sternum, coxae and trochantera reddish-brown, femora, patellae, tibiae and metatarsi dark. Abdomen ventrally with dark longitudinal band (Figure 16e). Spinnerets light brown.

Types
Male holotype (SMFD) from Venezuela, State of Amazonas, Puerto Ayacucho, Pozo Crystal (inside fallen rotten damp log), 15 February 1995, R. C. West col.; one female paratype (SMFD), the same data as in the male holotype.

Etymology
The specific name is a patronym in honour of the late Nick Gordon. Nick Gordon found this new species together with the collector while making a documentary about tarantulas.

Diagnosis
Cyriocosmus nicholausgordoni sp. nov. can be distinguished from all other congeners, except C. leetzi Vol, 1999, by its reddish-brown carapace with the black caput and seven radially arranged black spots in the thoracic area, and by the presence of the dark longitudinal ventral band. Males have short paraembolic apophysis and weakly developed prolateral superior keel in palpal bulb morphology, female spermathecae have a caliciform extension and convex basal plates. From C. leetzi it differs by the black femora without whitish longitudinal stripes on the dorsal face and the presence of the fourth pair of very narrow lateral stripes near spinnerets. Males also differ by the absence of retrolateral process in palpal tibia, instead there is only a cluster of numerous spiniform setae.
Distribution ( Figure 36) Known only from the type locality. The region is originally covered with the Amazon rainforest.
Palpal organ as in Figure 32c, d, embolus with short PA reaching approximately one sixth of embolus, smooth PS keel is not fused with PA. Tegulum with distinct granulated TP, projecting prolaterally. Retrolateral face of cymbium with basal field of spiniform setae. Protuberance on retrolateral face of palpal tibia is absent, instead there is only a cluster of numerous spiniform setae (Figure 18c, d). Two unequal subapical apophyses are present on tibia I ( Figure 28c): a longer retrolateral tibial apophysis with single subapical spine on dorsal side, a shorter prolateral tibial apophysis with single, welldeveloped retrolateral basal spine reaching approximately two-thirds of its length. Metatarsus I not sigmoidly curved and without basal or median protuberance on retrolateral face. Metatarsus I flexion is between both tibial apophyses.
Coloration and covering setae: dorsal view (Figures 17a, 18a): carapace reddishbrown with black caput and seven radially arranged black spots in thoracic area, coxae and trochantera reddish-brown, femora black, patellae, tibiae, metatarsi black with longitudinal whitish stripes, tarsi black, chelicerae dorsally covered with whitish pubescence. Patellae I, II and palpal patella with two longitudinal stripes, patellae III, IV with two unequal diagonal stripes. Abdomen black with central reddish-brown patch of urticating setae and three pairs of clear lateral stripes, basally joined, the second and the third pair is joined with central patch. The fourth pair of very narrow stripes is located near spinnerets. Length of central patch: 3.46, width 3.26. Ventral view ( Figure 18b): labium, sternum, coxae and trochantera reddish-brown, femora, patellae, tibiae, metatarsi and tarsi dark grey. Abdomen ventrally with dark longitudinal band. Spinnerets fawn.
Coloration and covering setae: dorsal view (Figures 17b, 19a): carapace reddishbrown with black caput and seven radially arranged, fused black spots in thoracic area, coxae and trochantera reddish-brown, femora black with only triangular spot at apex, patellae, tibiae and metatarsi black with whitish longitudinal stripes. Tarsi black. Patellae I, II with two longitudinal stripes, patellae III, IV with two unequal diagonal stripes. Abdomen (Figure 19c) black except central U-shaped reddish-brown patch and three pairs of clear reddish-brown lateral stripes, joined both basally and apically with central U-shaped patch, complemented by fourth pair of very narrow stripes near spinnerets. Length of central patch: 6.7,width 4.5. Ventral view (Figure 19b): labium, sternum, coxae and trochantera reddish-brown, femora, patellae, tibiae, metatarsi and tarsi dark grey. Abdomen ventrally with dark longitudinal band.

Etymology
The specific name, giganteus (adjectivum, Latin) means giant, and refers to the size of this new species and to the fact that this male holotype is so far the largest known male in the genus.

Diagnosis
Cyriocosmus giganteus sp. nov. can be distinguished from all other congeners by the uniformly coloured carapace, abdomen without stripes and by the male palpal bulb without prolateral superior keel.
Distribution (Figures 36, 37) Known only from the type locality.
Palpal organ as in Figure 25g, h, embolus with short narrow PA reaching approximately one-third of embolus, PS keel is absent. Tegulum with distinct TP, projecting prolaterally. Retrolateral face of cymbium without basal field of spiniform setae, both cymbial lobes approximately of the same length. Retrolateral face of palpal tibia with indistinct subapical protuberance covered with cluster of numerous spiniform setae (Figure 20f). Two unequal subapical apophyses are present on tibia I (Figure 28b): a longer retrolateral tibial apophysis with short, stout subapical spine on dorsal face, a shorter prolateral tibial apophysis, apically flattened, with single, well-developed retrolateral spine at base and approximately of the same length as prolateral tibial apophysis. Metatarsus I not sigmoidly curved and without basal or median protuberance on retrolateral face. Metatarsus I flexion is between both tibial apophyses.

Diagnosis
The species can be distinguished from all other congeners by lacking the striped pattern on dorsal abdomen ( Figure 21c) and by the black carapace with two pale and wide lateral stripes (Figure 21a), in combination with the yellowish longitudinal stripes on the dorsal face of the legs in females. The males have a long paraembolic apophysis and indistinct prolateral superior keel in palpal bulb (Figures 21f, 26c, d, 32e, f).
Coloration and covering setae: dorsal view (Figures 22b, 23a): carapace, coxae and trochantera reddish-brown, and covered with golden pubescence, except dark oval spot covering caput and surroundings of fovea, chelicerae dark brown, and covered with golden pubescence, femora, patellae, tibiae, metatarsi and tarsi dark grey and intermixed with long, dark setae, except patellae of posterior legs which are covered with short whitish setae, femora IV with golden basal pubescence. Patellae I, II and palpal patellae with two distinct longitudinal stripes without covering setae, patellae III, IV with single diagonal stripe. Palpal femur and femur I prolaterally bare. Retrolateral face of femur IV bare. Abdomen ( Figure 23c) covered with short black setae, intermixed with long, pale setae in posterior part, except central reddish-brown glossy patch, three lateral stripes on each side, basally joined, two short stripes on each side of spinnerets and two weak anteriorly located spots near area of urticating setae. Length of central patch: 6.4, width 3.5. Ventral view (Figure 23b): labium, sternum, coxae and trochantera reddish-brown, femora, patellae, tibiae and metatarsi and tarsi dark grey. Abdomen ventrally with dark longitudinal band (Figure 23d). Spinnerets: basal segment reddishbrown, central segment grey, apical segment dark grey.

Variability
The variability in the shape of spermathecae is shown in Figure 33i, j. The variability in the length of the carapace, the number and the arrangement of the cheliceral teeth, the number of labial and maxillary cuspules, the leg pattern and the spination of tibial apophyses is shown in Table 3.

Diagnosis
Differs from all other congeners, except C. nicholausgordoni sp. nov., by its reddishbrown carapace with the black caput and seven radially arranged black spots in the thoracic area, and by the presence of the dark longitudinal ventral band. From C. nicholausgordoni sp. nov. it differs by the black femora with whitish longitudinal stripes on the dorsal face. Males also differ by the presence of the retrolateral process in palpal tibia with a cluster of numerous spiniform setae (Figure 18e).

Discussion
Although very much has been written about species concepts and species delimitation, discerning different species with unknown distribution, known only from few localities within large biogeographical units, as for example the Amazon basin, remains difficult.
In theraphosid spiders we most often know only one or few specimens from a single population and nothing about neighbouring (sub-)populations, i.e. about the whole metapopulation, as defined by Levins (1969). Thus we are not able to assess the differences between various populations based on their different eco-evolutionary dynamics and local adaptations during the permanently running process of speciation.
How could we assess the distribution area and species delimitation of Cyriocosmus leetzi Vol, 1999? Cyriocosmus leetzi is now known from the type population in Colombia and from Venezuelan population in Táchira. The representatives of both populations share the common reddish-brown carapace with the black caput and seven radially arranged black spots in the thoracic area, the black abdomen with three clear lateral stripes joined with the U-shaped urticating setae patch, and the black legs, dorsally with whitish longitudinal stripes ( Figure 34). The male palpal bulb morphology with short PA is similar and comparable in both populations, as is the shape of spermathecae with caliciform extensions and convex basal plates. A comparative analysis of the species was done, combined with that of the closely related Cyriocosmus nicholausgordoni sp. nov., and the differences are shown in Table 16. In the case of Cyriocosmus leetzi, only two traits show differences that can be assessed as significant: the presence of the fourth pair of very narrow stripes near spinnerets, not basally joined with three clear lateral stripes (present only in the non-type Venezuelan population), and a range of metatarsal scopulae. Moreover, it was recorded in the population from Táchira that a pair of short anterior stripes near pedicelo, well visible in juveniles, disappears during ontogeny. There is no reason to doubt that both populations of Cyriocosmus leetzi and Cyriocosmus nicholausgordoni sp. nov. belong to the same lineage because of the high degree of conformity.
It is also evident that the type population and the population from Táchira may represent populations connected along the eastern side of the mountain range of Andes, but the population of Cyriocosmus nicholausgordoni sp. nov. from Amazonas is       obviously isolated by the Orinoco River. Further research of both species is necessary to understand the process of speciation inside the lineage of Cyriocosmus leetzi + Cyriocosmus nicholausgordoni sp. nov. Cyriocosmus peruvianus sp. nov. and C. itayensis sp. nov. are very closely related species that can be distinguished only by the shape of spermathecae, the size of adults (C. peruvianus sp. nov. < C. itayensis sp. nov.; see Table 3) and the morphology of the male palpal bulb, of which conglomerate of PS keel with PA is much more pronounced in C. itayensis sp. nov. The coloration is almost congruent, as well as the terrestrial life style and geographical affinity. I believe that the two species had a common ancestor and the process of speciation started when the population was split by any natural barrier in the past, e.g. by the Nanay River. C. peruvianus sp. nov. evolved on the left bank of the Nanay River, unlike C. itayensis sp. nov., which evolved on the right bank along the Itaya River. Intrinsic reproductive isolation was not tested. Both species were strictly established according to the phylogenetic species concept (sensu De Queiroz 2007). Further research of neighbouring populations is necessary, just as in the case of Cyriocosmus leetzi and Cyriocosmus hoeferi sp. nov.
Both genders of Cyriocosmus ritae from the Peruvian population near Iquitos are described here, the female specimen being described for the first time. Males from the population near Iquitos that fit into the diagnoses of C. ritae established by Pérez-Miles (1998) and confirmed by Fukushima et al. (2005) were compared with the data from both of the cited papers. The only differences recorded in the material from Iquitos involved the absence of the whitish longitudinal stripes on the dorsal face of the legs that were mentioned as present in the data matrix made by Fukushima et al. (2005), but not included in the original description, as well as the different number of lateral stripes on the abdomen, where only four are mentioned in the original description and five (three basally joined and two short separate stripes near spinnerets) included in the paper of Fukushima et al. (2005, figure 48) and in the material from Iquitos. This discrepancy was solved by re-examining the male holotype. The whitish longitudinal stripes on the dorsal face of male legs were not found, and only patellae and distal halves of metatarsi were covered with short whitish setae.