Phrynus calypso, Joya, Daniel Chirivi, 2017

Joya, Daniel Chirivi, 2017, A new species of Phrynus Lamarck, 1801 (Arachnida: Amblypygi), from Trinidad and Tobago, and Venezuela, with a redescription of Phrynus pulchripes (Pocock, 1894), Zootaxa 4254 (3), pp. 357-374 : 359-365

Phrynus calypso sp. nov.

Figures: 1A; 2E–F; 3–4; 7C, D, F; 8 A, C.; Table 1

Type material: Holotype (female): Trinidad and Tobago: Trinidad: Laja Trace (10.741194, -61.271472): tropical rain forest of low mountain, 180–550 masl, Victor Townsend, Paul Reslar, Dan Proud, 22 July 2006, female (CASENT 9060553).

Paratypes (16 females, 17 males, 8 juveniles): Trinidad and Tobago, Trinidad, Laja Trace (10.741194, - 61.271472): tropical rain forest of low mountain, 180–550 masl, Victor Townsend, Paul Reslar, Dan Proud, 22 July 2006, 1 female and 1 male (CASENT 9060553) . Trinidad and Tobago, Trinidad, Petite Tacarib (10.741194, - 61.225750): Crapo-cocorite sessional forest-jungle, 0–10 masl, Victor Townsend, Dan Proud, Rebeca Hunter, 12 July 2005, 2 females (CASENT 9060537) . Trinidad and Tobago, Trinidad, North Road, 5km from Maracas Bay Port of Spain (10.750593, -61.478858): Fernando Marques, manual collection, 1 October 2014, 1 juvenile, 2 males ( MZUSP 70892 ) . Venezuela, Estado de Monagas, Cave over Cueva del Guacharo (10.200000, -63.639722): Abel Pérez, Alessandro Giupponi, 8 December 2002, female ( MNRJ 09310 ) . Venezuela, Estado de Monagas, Cave over Cueva del Guacharo (10.200000, -63.639722): Abel Pérez, Alessandro Giupponi, 8 December 2002, 5 females and 6 males ( MNRJ 09315 ) . Venezuela, Estado de Monagas, Cueva del Guacharo, (10.200000, -63.639722): Abel Pérez, Alessandro Giupponi, 8 December 2002, 1 female and 3 males ( MNRJ 09312 ) . Venezuela, Estado de Monagas, Quebrada Cambural (10.042500, -63.558861): Abel Pérez, Alessandro Giupponi, 8 December 2002, 2 females and 4 males ( MNRJ 09317 ) . Venezuela, Estado de Monagas, Way between Cueva del Guacharo and Caripe (10.169900, -63.505800): 1000 masl, Abel Pérez, Alessandro Giupponi, 3 December 2002, 1 female, 1 male, 5 juvenile ( MNRJ 09316 ) . Venezuela, Estado de Sucre, Santa Maria (10.631944, -63.077777) : La Vega, Quebrada Macha, Abel Pérez, Alessandro Giupponi, 6 December 2002, 3 females ( MNRJ 09313 ) . Venezuela, Estado de Sucre, Santa Maria (10.631944, -63.077777) : La Vega, Quebrada Macha, Abel Pérez, Alessandro Giupponi, 6 December 2002, 2 juvenile ( MNRJ 09314 ) .

Distribution. Trinidad Island, in Trinidad and Tobago, and Coast of Venezuela ( Fig. 8A , C).

Etymology. Species name is dedicated to music rhythm Calypso , typical from Trinidad and Tobago. In commemoration to cultural expression of this country. Name in apposition.

Diagnosis. Phrynus calypso sp. nov. is distinguished from most Phrynus species by the presence of two teeth in the external line of the base of chelicerae, both with an acuminated apex, and a small spine on dorso-mesal surface of pedipalp tarsus. These two conditions are present together in just four more species: Phrynus araya Colmenares & Villarreal, 2008 , Phrynus panche Armas & Angarita, 2008 , Phrynus pulchripes ( Pocock, 1894) and Phrynus exsul Harvey, 2002 . P. calypso differs from P. araya on the number o spines between Fv3 and Fv5. P. araya presents three spines between Fv3 and Fv5; Fv4 and two more, according to nomenclature proposed by Quintero (1981). P. calypso has usually just one Fv4 spine and at most can present two spines between Fv3 and Fv5. P. araya has Fv3 equal or longer than Fv5, in P. calypso Fv 3 is shorter than Fv5. The female genitalia of P. panche , P. exsul and P. araya are very similar, all have the sclerites of gonopods with a rectangular elongated shape, this sticks out from seminal receptacles ( Fig. 1 D), P. calypso has short, fusiform sclerites with an acuminated apex ( Fig. 1 A). In P. exsul spine Fd4 is absent, and in this position there is a small setal tubercle. P. exsul , P. panche and P. pulchripes present three small spines placed over the Td3, P. calypso presents at most one spine on the base of Td3. The most similar species to P. calypso is P. pulchripes . They differ mainly in the female genitalia: in P. pulchripes sclerites do not have an acuminated apex, and are shorter and thicker, as well as more sclerotized. Seminal receptacles in P. pulchripes are completely sclerotized ( Fig. 1 B), in P. calypso the sclerotized region covers just the basal area. Female genitalia in P. calypso can be confused with the genital of Phrynus barbadensis ( Pocock, 1894) , which inhabit the same localities, however this species has the gonopod apex sclerite at least three times thinner than the base ( Fig. 1 C), a condition that is not present in P. calypso .

Female genitalia present many characters frequently used for to identify species in Phrynidae ( Quintero 1981) . However, little is known about ontogenetic variation of female genitalia ( Weygoldt 2000). One reasonable question is if differences between two putative species could be attributed to variation between smaller and larger individuals of the same species. In this research all available female genitalia of P. calypso and P. pulchripes were examined, even in young exemplars. In all cases pattern of shape, coloration, proportions and position were constant, with some variations in size. Some characteristics can be accentuated in larger specimens, like curvature in sclerites of P. pulchripes ( Fig.1 B, 2A–D), and size of sclerites in both species. Also, in both species, coloration of sclerites and seminal receptacles could be a little lighter in smaller specimens ( Fig. 1 A–B, 2). Differences between these two species were constant, a posterior view of female sclerites allowed a clearer identification of the differences in shape, and small details like curvatures in P. pulchripes and the less sclerotized ending in P. calypso (cf. P. pulchripes Fig. 2 B, D, vs P. calypso Fig. 2 F). These differences were also visible when we compared with illustrations of type material of P. pulchripes , (See Fig. 126 in Quintero 1981). Recently, P. gervaisii was reported as a synonym of P. barbadensi s ( Armas 2013). Specimens of P. gervaisii were considerably smaller than specimens of P. barbadensis , in this case female genital of both are almost equal, with just some differences in size and accentuation of coloration ( Armas 2013). Unlike this case, differences between P. calypso and P. puclhripes are in relation to shape, patterns of sclerotization and proportions. There are no intermediate character states between larger females of P. calypso and smaller females of P. pulchripes . In spite of the somatic similarities, the female genitalia of P. calypso and P. pulchripes present clear differences between them, as interspecific differences with genitals of other species such as P. barbadensis and P. panche .

TABLE ·. Measures of bođy, peđipalp segments anđ femur of legs in Phrynus calypso sp. nov., measures in mm. When character is missing cell is empty. Paratypes (1-10): 1 anđ 2 corresponđ to CASENT 9060553, 3 anđ 4 to CASENT 9060537, 5 to MZUSP 70892-1, 6 to MZUSP 70892-2, 7 anđ 8 MNRJ 69315, 9 anđ 10 to MNRJ 69312.

Holotype 1 2 3 4 5 6 7 8 9 10

Female Female Male Female Female Male Male Female Male Male Female length 12.7 12.3 14.6 13.0 11.6 20.8 13.6 17 21.8 17.8 18.3 with chelicerae)

Prosoma

Maximum 7.4 7.2 9.3 8.4 6.8 11.7 7.9 9.4 11.2 10.1 9.7 wiđth

Maximum 4.1 4.1 6.6 6.1 4.7 8.1 5.5 6.6 8.2 6.9 7.2 length

Opisthosoma

Maximum 5.5 5.1 5.8 5.8 6.1 8.4 5.5 7.4 7 6.1 6.2 wiđth

Maximum 7.7 7.1 8.6 7.4 7.4 11.6 7.6 9.6 11.7 9.7 9.8 length

Right Left Right Left Right Left Right Left Right Left Right Left Right Left Right Left Right Left Right Left Right Left

Peđipalp lengths

Trochanter 1.8 1.8 2 2 2.5 2.5 2.1 2.1 1.5 1.5 3.1 3.3 2.4 2.4 2.4 2.6 3.1 3.1 2.4 2.4 3 3 Femur 3.6 3.4 4.8 4.8 5.3 5.6 4.7 4.7 3.4 3.4 7.7 7.6 4.2 4.4 5.4 5 7.7 7.9 6.1 6.1 5.7 5.7 Patella 3.6 3.6 4.6 4.6 6.2 6.3 5.6 5.6 4.1 4.1 8.0 7.7 4.8 4.9 5.5 5.5 8 8.3 6.3 6.4 6.2 6.2 Tibia 2.1 2.0 2.4 2.4 3 2.9 2.7 2.4 1.9 1.7 4.1 4.0 2.3 2.2 3 3 4.2 4.2 3.2 3.2 3.4 3.4 Tarsus+apotele 2.5 2.4 2.5 2.5 2.7 2.7 3.1 3.1 2.0 2.0 4.2 4.1 2.4 2.4 3.9 3.7 4.2 4.1 3.8 3.6 3.6 3.6 Femur length

Leg I 10.2 10.2 11.8 12.0 16.0 15.5 12.8 12.7 9.7 8.0 21.4 21.4 12.3 12.2 16.3 16.1 23.3 23.2 17.5 17.6 17.8 17.8 Leg II 7.6 7.7 8.1 8.1 11.5 11.4 9.1 9.3 6.9 6.8 17.2 16.0 8.6 8.6 10.9 10.8 16.3 16.3 12.6 12.7 12.9 12.5 Leg III 8.8 8.6 9.0 9.2 13.1 12.8 10.5 10.2 7.9 7.5 17.1 17.1 10.0 9.8 12.4 12.5 8.7 8.6 13.3 13.3 13.4 13.4 Leg IV 7.4 7.0 7.8 7.8 10.4 10.4 8.6 8.6 6.5 6.4 13.7 13.6 6.4 7.8 10.4 10.4 14.5 14.4 11.6 11.6 11.3 11.2

. segments 33 31 32 32 32 31 31 31 31 31 31 29 31 30 31

I

. segments 67 68 68 67 75 68 68>58 66>50>64 61 68 65 67 tarsus I

. segments 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 basitibia IV

Description. Female (Holotype): Total length 12.7 mm. Carapace and opisthosoma color dark brown, pedipalps and legs a lighter brown. Dimensions of prosoma, opisthosoma, pedipalpal segments, and all leg femora are provided in Table 1.

Carapace. Frontal margin with one lobe on each side, lobes are not joined in the middle area. Carapace presents a group of granules widely spaced, inferior margin with white regions. Lateral and anteromedial eyes clearly visible, ocular tubercle black ( Fig. 3 A).

Sternum. Tri-segmented; all segments clearly sclerotized, area around segments is sclerotized too. Tritosternum projected anteriorly, elongated, conical, with four setae in basal region, two in medial region and two more in the apex.

Second segment (tetrasternum) hexagonal with one setae on each side of medial region, rounded shape. Third segment (pentasternum) rhomboidal slightly shorter than the second segment, with one setae on each corner of medial region. Metasternum longitudinally divided with two setae on the posterior margin of each half ( Fig. 3 B).

Abdomen. Oblong, color dark brown, with soft punctuations of lighter coloration. Presents patches of light coloration at both sides of each tergite. Carapace wider than the Abdomen ( Fig. 3 C).

Female genitalia. Genital operculum pentagonal, it extends to second segment of opistosoma, seminal receptacles rounded, dorsal surface lightly sclerotized on basal area, with coloration darker than the rest of the structure which is mainly white. Sclerite of gonopod, with fusiform shape, acuminated apex which is less sclerotized, almost transparent. Sclerite coloration light brown, with apical region translucent ( Fig. 1 A, 2 E–F).

Chelicera. Mesal row of the basal segment of chelicerae with three teeth, the first bilobed, placed in the proximal portion, followed by one tooth shorter in medial portion, and the third, bigger than the others, placed in the distal region ( Fig. 3 D). Ectal row with two teeth of similar sizes, both with an acuminated apex, placed in the more proximal region, and previous to the medial region. There is a small keel on the distal region ( Fig. 3 F). Mobile segment of chelicerae with five teeth.

Pedipalp. Trochanter: Prolateral face with four spines; spines Tr1 and Tr3 placed in medial region, Tr2 placed near to the ventral margin. Tr3 and Tr4 with subequal lengths. Spine lengths: Tr2>Tr1>Tr4ŻTr3. Dorsal oblique series of five setal tubercles. Dorsomedial area with one spine with a length slightly longer than one tubercle ( Fig. 3 E). Femur: Dorsal face with five conspicuous spines, spine Fd4 is reduced, similar in size to a setal tubercle, and there are two little spines more distal than Fd6, in total there are eight spines in the dorsal femur, Fd1 and Fd2 share the same base, Fd3 is adjacent to Fd2, between Fd5 and Fd6 there is a small tubercle. Spine lengths: Fd3>Fd2>Fd5>Fd1>Fd6>Fd7>Fd8>Fd4 ( Fig. 4 D). Ventral face with six conspicuous spines, Fv4 very small, more distal than Fv6 are two tubercles present on both pedipalps, and there is also one tubercle between Fv5 and Fv6. Spine lengths: Fv1>Fv2>Fv5>Fv3>Fv6>Fv4 ( Fig. 4 C). Patella: Dorsal face with eight major spines, between Pd1 and Pd2 there is a small spine here named Pd1b, to conserve the spines numeration proposed by Quintero (1981): Pd3>Pd5>Pd2>Pd4>Pd6>Pd1>Pd7>Pd8>Pd1b ( Fig. 4 B). Ventral face with five major spines. Pv3 is small, but it is present on both pedipalps, five very small spines of similar size are present between each major spine: one more proximal than Pv1, between, Pv1 and Pv2, Pv4 and Pv5, Pv5 and Pv6, and one more distal than Pv6. Lengths: Pv2>Pv5>Pv1>Pv4>Pv6>Pv3 ( Fig. 4 A). Tibia: Dorsal face with three major spines, between Td2 and Td3 there is one tubercle and one small spine, more distal than Td3 there are two tubercles, and a small spine smaller than others, here named Td4. Spine lengths: Td2>Td3>Td1>Td4 ( Fig. 4 F). Td1 has half the length of Td3. Ventral face with three major spines, and two tubercles between Tv2 and Tv3. Spine lengths: Tv2>Tv3>Tv1 ( Fig. 4 E). Tarsus+Apotele: Internal face with two lines of dorsomedial bristles; suture between tarsus and apotele is not visible. Tarsus with an inconspicuous spine on proximal end of dorso-mesal surface.

Legs. Femora with lighter coloration rings. Femora lengths: I>III>II>IV (Table 1). Leg I: Tibia with 33 segments and tarsus with 67 segments in the right leg; left leg with 31 segments in the tibia and 68 segments in the tarsus. Leg IV: Basitibia with three segments. Basitibia-distitibia lengths: BT1>DT>BT3>BT2. Basitarsus and Telotarsus with subequal sizes. Tarsus tetramerous.

Variation. Paratype series measurements and number of segments in basitibia IV, Tibia I and tarsus I, are summarized in Table 1, just adult specimens were measured. Sternum: Setae over all segments of sternum are very variable. Pedipalp: Femur: Dorsal face: the size and presence of mentioned tubercles in the holotype is variable in paratypes. Ventral face: the tubercle’s presence is variable, in larger specimens two tubercles can be observed between Fv5 and Fv6, and can be interpreted as small spines. In specimens from Venezuela one or two small tubercles may be observed between Fv3 and Fv5. In one male (MNRJ 09315) there were two spines observed between Fv3 and Fv5, and these were present just in the right pedipalp. Patella: Dorsal face: in some individuals there is a tubercle anterior to spine Pd1, spine Pd1b is present in many individuals but can be less conspicuous, or even be absent in others. In larger individuals there is one spine smaller than the others that was more distal than Pd8. Ventral face: Tubercles were more conspicuous in larger sized individuals; in some cases, a second tubercle was present more proximal than Pv1. Tibia: Dorsal face: presence of tubercles between spines was variable. Ventral face: In some individuals there was just one tubercle between Tv2 and Tv3. Legs. Number of segments of tibia and tarsus of leg I was variable among the individuals, number of tibial segments keeps between 31 and 33, and tarsal segments between 61 and 75. In observed specimens, number of segments did not depend on size or sex (Table. 1).

Male. Observed males were larger in size than females (Table. 1), spination pattern as in females, genital operculum has an inferior margin oval and cover part of third segment of opisthosoma.

Male genitalia. (MZUSP 70892): LaM longer than Pi. LoL 1 and LoL 2 subequal in length, both densely covered with minute projections, in LoL 1 projections are tubular, thin and elongated, placed close together; in LoL 2 are tubular and short, some of these with ramifications at the apex, spaced over a base clearly visible. In dorsal view the LoD rounded and shorter than LoL1 ( Fig. 7 C, D, F).

Figures

FIGURE 1. Compared female genitalia: A. Phrynus calypso sp. nov, Holotype. B. Phrynus pulchripes, ICN-Am 85. C. Phrynus barbadensis. MUD-AM 403. D. Phrynus panche. ICN-AM 64.

FIGURE 2. Variation in female genitalia: Phrynus pulchripes: MZUSP 70924 - 1: A. Dorsal view. B. posterior view. MZUSP 70924 - 2: C. Dorsal view. D. posterior view. Phrynus calypso: CASENT 9060553: E. Dorsal view. F. posterior view.

FIGURE 3. Phrynus calypso sp. nov. Female Holotype: A. Carapace. B. Sternum. C. Dorsal opisthosoma. D. Chelicera, mesal view. E. Trochanter (Tr). F. Chelicera, ectal view.

FIGURE 4. Phrynus calypso sp. nov. Female Holotype: A. Ventral patella (Pv). B. Dorsal patella (Pd). C. Ventral femur (Fv). D. Dorsal femur (Fd). E. Ventral tibia (Tv). F. Dorsal tibia (Td).

FIGURE 7. Male genitalia: Phrynus pulchripes ICN-AM 86: A. Dorsal view. B. Ventral view. E. Detail of ornamentation of LoL 1 and LoL 2. Phrynus calypso sp. nov. MZUSP 70892: C. Dorsal view. D. Ventral view. F. Detail of ornamentation of LoL 1 and LoL 2. Fi = fistula; LaM = lamina medialis; LoD = lobus dorsalis; LoL 1 = lobus lateralis primus; LoL 2 = lobus lateralis secundus; Pi = processus internus of gonopod.

FIGURE 8. A. Distribution of Phrynus calypso sp. nov and Phrynus pulchripes. B. Distribution of P. pulchripes, detail of records in Colombia. C. Distribution of P. calypso, detail of records. P. calypso: yellow circle = records, yellow circle with black border = Type locality. P. pulchripes: red square = records, red square with black border = Type locality.