Phrynus operculatus Pocock, 1902

Joya, Daniel Chirivi, 2021, Four new species of Phrynus, Lamarck (Arachnida: Amblypygi) from Mexico, Zootaxa 4948 (2), pp. 151-183 : 153-157

Phrynus operculatus Pocock, 1902

Figures. 1–3 , 17–18 . Table. 1, 6.

Phalangium palmatum Herbst , in Lichtenstein & Herbst 1797: 82–84, plate IV fig. 2.

Phrynus palmatus (Herbst) : Latreille 1804: 136–137; Gervais 1844: 6; Wood 1863: unnumbered fig. on p. 680; Butler 1873: 118, plate 6 fig. 5; Butler 1879: 314, 315; Mello-Leit„o 1931: 42–4.

Admetus palmatus (Herbst) : C.L. Koch 1850: 81; Hansen 1893: 161.

Neophrynus palmatus (Herbst) : Kraepelin 1895: 30–34, figs 3, 12, 15, 19, 25, 27–28, 30.

Tarantula palmata (Herbst) : Pereyaslawzewa 1897a: 319; Pereyaslawzewa 1897b: 377; Kraepelin 1901: 264; Börner 1902a: fig. 6; Börner 1902b: 214; Börner 1904: 5, figs 6, 18, 23, 26, 32, 57, 102–114, figs 10–11, 38, 46, 64, 85; Comstock 1913: 20, fig. 19; Petrunkevitch 1925: 60; Kästner 1932: figs 8, 23, 28, 31, 73–74, 81; Werner 1935: 475, figs 25, 34–35, 40, 47, 56, 59, 72–73, 100, 110, 113, 141a–b, 182b; Speijer 1936: 260–261, figs 12–15; Millot 1949a: figs 68, 71; Millot 1949b: figs 326, 328, 333, 344; Petrunkevitch 1949, fig. 17; Caporiacco 1951: 36; Zakhvatkin 1952: fig. 28b; Weygoldt 1977: 273–275, figs 1a–c, 3a.

Phrynus operculatus Pocock, 1902: 52 , plate 10 figs 8, 8a–c; Franganillo 1926: 66–67 (probable misid); Franganillo 1931: 120; Mello-Leit„o 1931: 42; Franganillo 1936: 149; Quintero 1981: 155–157, figs 102–108, 154–155, 157, 162, map; Coddington et al. 1990: 11; Vázquez-Rojas 1995: 30 (as Phrynus opercularis [sic]); Vázquez-Rojas 1996: 72; Ávila-Calvo & Armas 1997: 32; Armas 1999: 29 (synonymized with Phalangium palmatum by Quintero (1981), but treated as senior synonym); Armas & Víquez 2001: 15; Armas & Pérez 2001: 48; Jiménez & Gutierrez 2004: 863; Harvey, 2003: Order Amblypygi , 28–29; Armas & Gadar 2004: 134–136, fig. 8; Víquez & Armas 2006: 307–3016, figs. 9, 13; Armas 2006a: 232; Armas 2006b: 345–358, fig. 18, Table 1; Chirivi & Armas 2012: 396; Armas et al. 2013: 25–30; Armas 2014: 30–35; Armas et al. 2014: 3–6, fig. 3; Guzmán et al. 2015: 478.

Not Phrynus palmatus (Herbst) : Koch 1840: 13–15, fig. 601 (see Phrynus gervaisii (Pocock)) .

Type locality: Guatemala .

Revised material: (4 females, 1 male): MEXICO. Chiapas: Arriaga (16.235833, -93.899722): 10 km at north, 305 masl, 23 August 1972, C. Mullinex, K. Lucas, female ( CASENT 9060540 ) ; 10 km at north, 305 masl, 23 August 1972, C. Mullinex, K. Lucas, female ( CASENT 9060527 ) ; 10 km at north, 305 masl, 23 August 1972, C. Mullinex, K. Lucas, male ( CASENT 9060512 ) . Tonalá (16.089722, -93.754722): 13 miles South, slope of Cerro Vernal , 610 masl, 5 October 1974, D.E. Breedlove, J.A. Breedlove, female ( CASENT 9060511 ) . Federal highway 190, near to Santa Isabel (16.494444, -94.081944): 805 masl, 1 September 2005, O. Francke, M. Jaimes, A. Valdez, H. Montaño, female (CNAN-Am000407) .

Distribution. BELIZE. GUATEMALA. MEXICO: States of Chiapas, Colima, Guanajuato, Guerrero, Jalisco, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Sinaloa. UNITET STATES OF AMERICA: Texas. Many of these records are not supported with deposited material in collections and have no specific localities ( Quintero, 1981; Armas 2006b; Armas et al., 2014). ( Fig. 18 ).

Diagnosis. Phrynus operculatus can be distinguished from other species of the genus by the presence of the following set of characters: only one conspicuous tooth in the ectal row of the base of chelicerae, with an acuminated apex ( Fig. 1F ), does not present an inconspicuous spine in the dorsomedial area of the pedipalp tarsus, Pd2 is smaller than Pd4, Fv3 smaller than Fv5, Td1 is very small but longer than Td4 ( Fig. 2 ). Female genitalia have tubular and elongated gonopod bases, with claw-like sclerites of the gonopod, curved at the apical region ( Fig. 17A ). In the male genitalia, the ornamentations over LoL 1 and 2 are tubular and long, and LoD is very thick and pronounced ( Fig. 3 ). Almost the wide of the base of LoL 1 ( Table 6).

Species comparisons: Phrynus operculatus is morphologically similar to P. jalisco , however, the female genitalia are different, in P. jalisco the gonopod bases are more elongated, the sclerites are thicker and have a pronounced inward curvature ( Fig. 17F ). Phrynus operculatus is also similar to P. aliciae sp. nov., however, in this species the base of the sclerites of the female genitalia are straighter and surrounds the lateral margin of the base ( Fig. 17C ) whereas in P. operculatus the base is smaller. Is also similar to P. alfonsoi sp. nov., but in this species, the sclerite of female gonopod is bigger with a more pronounced curvature in the apical region (17B). In the male genitalia, the projections on LoL 2 are completely tubular ( Fig. 3B ) whereas in P. alfonsoi sp. nov. the projections bases are wider ( Fig. 6B ). The LoD in P. operculatus is thicker than in the other species compared ( Fig. 3C , Table 6).

The spine Pd1 is conspicuous (a quarter of the size of Pd2) ( Fig. 2D ) similar to P. alfonsoi sp. nov., P. jalisco , and P. aliciae sp. nov.; Tv1 is shorter than Tv3 ( Fig. 2E ) similar to P. tresmarias sp. nov. and P. aliciae sp. nov. ( Fig. 8E , 14E ) whereas, in P. alfonsoi sp. nov. Tv1 is longer ( Fig. 5E ) and, in P. giseae sp. nov. and P. jalisco , are subequal ( Fig. 11E ). Td1 is a quarter of size Td2, so it differs from other species here compared ( Fig. 2F ).

Complementary description. Female (CNAN-Am000407): Total length 16 mm. Carapace and opisthosoma color brown, pedipalps and legs are brown with red tonalities. Dimensions of prosoma, opisthosoma, pedipalps segments, and all leg femora are provided in Table 1.

Carapace. The frontal margin is almost straight. Carapace presents a group of granules widely spaced; anterior, lateral, and posterior margins with a white border, posterior margin concave. Lateral and anteromedial eyes clearly visible, ocular tubercle black ( Fig. 1A ).

Sternum. Tri-segmented, all segments poorly sclerotized, the area around segments is slightly less sclerotized. Tritosternum projected anteriorly, elongated, conical, with eight setae on the basal region, four on the medial region, and two more in the apex. Second segment (tetrasternum) oval, with two setae on the medial region. Third segment (pentasternum) oval, slightly smaller than the second segment, with two setae on the medial region. Metasternum longitudinally divided, with two setae on the posterior margin of each half ( Fig. 1B ).

Abdomen. Oblong, color dark brown. Presents patches of light coloration on both sides of each tergite. The carapace is wider than the abdomen ( Fig. 1C ).

Chelicera. The mesal row of the basal segment of chelicerae with three teeth, the first is bilobed, placed in the proximal portion, lobe 1b bigger than 1a, followed by one tooth shorter in medial portion, and the third, bigger than the others, placed in the distal region ( Fig. 1D ). The ectal row with one conspicuous tooth (tooth 2) with an acuminated apex, placed in the medial region. There is a small keel on the most proximal region (tooth 1) ( Fig. 1F ). The mobile segment of chelicerae with four teeth, the first one is the biggest, followed by the third, and the fourth is the smallest.

Pedipalp. Trochanter: Prolateral face with four spines; spines Tr1 and Tr3 placed in medial region, Tr2 placed near ventral margin. Tr3 and Tr4 subequal in length. Spine lengths: Tr2>Tr1>Tr4ŻTr3. Dorsal oblique series of five setiferous tubercles. Dorsomedial area without spines but with one big tubercle ( Fig. 1E ). Femur: Ventral face with five major spines, Fv4, Fv3a, Fv5a, and Fv7 are small. Spine lengths: Fv1>Fv2>Fv5>Fv3>Fv6>Fv4>Fv7>F v5a>Fv3a ( Fig. 2A ). Dorsal face with five major spines; Fd4 small, with the spines Fd5a, Fd7, and Fd8 present as tubercles; Fd1 and Fd2 share the same base, Fd3 is separated from Fd2. Spine lengths: Fd2>Fd3>Fd5>Fd1>Fd6>Fd 4=Fd5a>Fd7>Fd8 ( Fig. 2B ). Patella: Ventral face with five major spines; with conspicuous spines Pv1-a and Pv1a, and one small spine between Pv2–Pv3, Pv4–Pv5, and Pv5–Pv6. Spine lengths: Pv2>Pv5>Pv1>Pv4>Pv6>Pv7>Pv1- a=Pv1a>Pv3 ( Fig. 2C ). Dorsal face with seven major spines, proximal to Pd1 there are two tubercles, Pd8 is small. Spine lengths: Pd5>Pd3>Pd4>Pd2>Pd6>Pd7>Pd1>Pd8 ( Fig. 2D ). Tibia: Ventral face with three major spines; has one tubercle and one small spine between Tv2–Tv3. Spine lengths: Tv2>Tv3>Tv1 ( Fig. 2E ). Dorsal face with three major spines, one tubercle between Td2–Td3, two conspicuous spines over Td3, distal to Td3 there are two small spines, the most distal spine is a little bigger and is here named Td4; Td1 is almost a half of Td3 length. Spine lengths: Td2>Td3>Td1>Td4 ( Fig. 2F ). Tarsus-metatarsus: Internal face with two lines of dorsomedial bristles; suture between the tarsus and metatarsus is not visible. Tarsus without an inconspicuous spine on the proximal end of the dorsomedial surface.

Legs. Femora brown. Femora lengths: I>III>II>IV ( Table 1). Leg I: Tibia with 29 segments and tarsus with 62 segments in both legs. Leg IV: Basitibia with three segments. Basitibia-distitibia lengths: BT1>DT>BT3>BT2. Basitarsus and Telotarsus subequal in size. Tarsus tetramerous.

Female genitalia. Genital operculum pentagonal, it extends to the second segment of the opisthosoma; gonopod bases tubular, dorsal surface poorly sclerotized; on the most proximal area, there is a strongly sclerotized line joining both gonopod bases, coloration brown on the most proximal region, and white over entire the dorsal surface. Sclerites of the gonopods claw-like, the base is more than two times wider than medial and apical regions, apex slightly curved. Sclerite coloration brown ( Fig. 17A , Table 6).

Variation. Measurements and the number of segments in the basitibia IV, Tibia I, and tarsus I, are summarized in Table 1. Sternum: Setae over all segments of the sternum are variable in number and position. Pedipalp: Trochanter: line of dorsal tubercles varies between four and six tubercles. Femur: Ventral face: tubercles are constant in most individuals, however, the size is variable, as well as the size of spines Fv4, Fv3a, Fv4a, and Fv5a, which are always smaller than major spines, but the relative sizes between them are variable. Dorsal face: the size and presence of tubercles mentioned in the complementary description are variable. Patella: Ventral face: Tubercles and small spines are more conspicuous in larger individuals. Dorsal face: tubercles can be more conspicuous, in some individuals the tubercles can be present between each spine, in larger specimens a small spine Pd8 can be present. Tibia: Ventral face: between Tv2–Tv3, two or three small spines can be present. Dorsal face: tubercles between spines can be more conspicuous. Legs. The number of segments of the tibia and tarsus of leg I, was variable among the individuals, the number of tibial segments varies between 29 and 42, and the tarsal segments between 62 and 69. In specimens studied, the number of segments did not depend on sex, frequently there are more tarsal segments in larger specimens ( Table 1).

Male. Males are similar in size to females ( Table 1) spination pattern as in females, genital operculum with the posterior margin oval, and covering part of the third segment of the opisthosoma.

Male genitalia. (CASENT 9060512): LaM longer than Pi. LoL 1 a little longer than LoL 2, both densely covered with minute projections, in LoL 1, projections are tubular, thin, and elongated, placed close together, the insertion base is not visible; in LoL 2 projections are similar to LoL 1, the insertion base is also not visible. In dorsal view, the LoD is thick and elongated with a similar size to LoL1 ( Fig. 3 , Table 6).

Figures

FIGURE 1. Phrynus operculatus. Female (CNAN-Am000407): A. Carapace. B. Sternum. C. Dorsal opisthosoma. D. Chelicera, mesal view. E. Trochanter (Tr). F. Chelicera, ectal view.

FIGURE 2. Phrynus operculatus. Female (CNAN-Am000407): A. Ventral femur (Fv). B. Dorsal femur (Fd). C. Ventral patella (Pv). D. Dorsal patella (Pd). E. Ventral tibia (Tv). F. Dorsal tibia (Td).

FIGURE 3. Male genital: Phrynus operculatus (CASENT 9060512): A. ventral view. B. Detail of ornamentation of LoL 1 and LoL 2. C. Dorsal view. Fi = fistula; LaM = lamina medialis; LoD = lobus dorsalis; LoL 1 = lobus lateralis primus; LoL 2 = lobus lateralis secundus; Pi = processus internus of gonopod. Colored lines illustrate the comparison between the LoD medial width (green line) and the width of the base of LoL 1 (red line).

FIGURE 5. Phrynus alfonsoi sp. nov. Female (CNAN-T01383): A. Ventral femur (Fv). B. Dorsal femur (Fd). C. Ventral patella (Pv). D. Dorsal patella (Pd). E. Ventral tibia (Tv). F. Dorsal tibia (Td).

FIGURE 6. Male genitalia: Phrynus alfonsoi sp. nov. (CNAN-T01445): A. ventral view. B. Detail of ornamentation of LoL 1 and LoL 2. C. Dorsal view. Fi = fistula; LaM = lamina medialis; LoD = lobus dorsalis; LoL 1 = lobus lateralis primus; LoL 2 = lobus lateralis secundus; Pi = processus internus of gonopod. Colored lines illustrate the comparison between the LoD medial width (green line) and the width of the base of LoL 1 (red line).

FIGURE 8. Phrynus aliciae sp. nov. Female (CNAN-T01447): A. Ventral femur (Fv). B. Dorsal femur (Fd). C. Ventral patella (Pv). D. Dorsal patella (Pd). E. Ventral tibia (Tv). F. Dorsal tibia (Td).

FIGURE 11. Phrynus giseae sp. nov. Female (CNAN-T01450): A. Ventral femur (Fv). B. Dorsal femur (Fd). C. Ventral patella (Pv). D. Dorsal patella (Pd). E. Ventral tibia (Tv). F. Dorsal tibia (Td).

FIGURE 14. Phrynus tresmarias sp. nov. Female (CASENT 9060565):A. Ventral femur (Fv). B. Dorsal femur (Fd). C. Ventral patella (Pv). D. Dorsal patella (Pd). E. Ventral tibia (Tv). F. Dorsal tibia (Td).

FIGURE 17. Female genitalia: A. Phrynus operculatus (CNAN-Am000407). B. Phrynus alfonsoi sp. nov. (CNAN-T01383). C. Phrynus aliciae sp. nov. (CNAN-T01447). D. Phrynus giseae sp. nov. (CNAN-T01450). E. Phrynus tresmarias sp. nov. (CASENT 9060565). F. Phrynus jalisco (CASENT 9060519). Sclerotization of the gonopod bases (black arrows); colored lines illustrate the position of the sclerite of the gonopod with respect to the gonopod base, the length comparison between the gonopod base (green line) and sclerite of the gonopod (red line), and the comparison between width of the base of the sclerite (purple line) and the width of the medial region of the gonopod base (blue line).

FIGURE 18. Distribution: Phrynus alfonsoi sp. nov. (blue circle), Phrynus aliciae sp. nov. (yellow circle), Phrynus giseae sp. nov. (purple circle), Phrynus jalisco (light blue circle), Phrynus operculatus (red circle), Phrynus tresmarias sp. nov. (orange circle). Circles with black rings: type localities.