Rusetria (K. Viets, 1956)

Goldschmidt, Tom, 2007, Studies on Latin American water mites of the genus Torrenticola Piersig, 1896 (Torrenticolidae, Hydrachnidia, Acari), Zoological Journal of the Linnean Society 150 (3), pp. 443-678 : 460-475

‘ RUSETRIA ’-LIKE SPECIES

Previously known species: T. gradaticoxalis K. O. Viets, 1977 ( Guatemala) .

New species from Costa Rica: T. costaricense , T. fontinale , T. harpagophora .

Differential diagnosis of the group: Antero-lateral dorsal platelets fused with main dorsal plate; dorsal plate reddish to red or with pale colour pattern ( Figs 6A, 6 B-2, 6B-3); idiosoma mid-sized to large, rounded to rounded-oval; capitulum with mid-sized to long rostrum; palp with +/– cone-shaped ventral projections at P2/3; genital skeleton apically relatively short, cella proximalis relatively large.

Discussion: These species are characterized (and clearly separated from all other neotropical species) by the fusion of the antero-lateral dorsal platelets with the large dorsal plate. Until now, no Torrenticola species with these characters has been documented from Latin America. Even though the fusion of the antero-lateral dorsal platelets of T. gradaticoxalis from Guatemala has not been recognized (or at least not mentioned in the description) by K.O. Viets (1977/ 78 Teil I), this species also clearly belongs to the ‘ Rusetria ’-like species (see discussion there). In the material from Costa Rica three new species were found. Originally species from the old world with this feature were placed in a subgenus Rusetria (K. Viets, 1956) , which has been synonymized with the subgenus Torrenticola by K.O. Viets (1987). Bader (1988) re-established the subgenus, however, in a cladistic analysis Wiles (1997a) stated that variations in the number of free dorsal platelets evolved independently in different clades and should therefore not be used to differentiate subgenera. In the present paper, the name ‘ Rusetria ’ is used only for practical reasons to separate a group of species easy to recognize from all the others. Hence, the use of the term ‘ Rusetria ’ in this study does not aim at the reintroduction of this subgenus. The – apart from the organization of the dorsal plates – quite variable morphology (regarding palps, rostrum and coxal field) of the four species known so far emphasizes the opinion of an independent fusion of the antero-lateral dorsal platelets with the main dorsal plate. Furthermore, in several specimens of other species partly fused antero-dorsal platelets can be found. Therefore, most probably the ‘ Rusetria ’-like species do not comprise a phylogenetic group.

Key to the species

1a Lateral margins of Cx-I/II/III very sharply graded, antero-lateral corner of Cx-II forming an anterior pointed edge, medial margin of Cx-II/III in male very short, in female pointed (Cx-I nearly extended to genital field) ( Fig. 15A, E ) ................................................................................................................................................................... T. gradaticoxalis

b Lateral margins of Cx-I/II/III various in shape, but antero-lateral corner of Cx-II never forming a pointed edge, medial margin of Cx-II/III in male mostly longer, in female short but present ( Figs 11A , 12A , 13A , 14A ) ...............2

2a Antero-medial platelets long and narrow; rostrum relatively short; P4 strongly curved, ventral setae on distinct tubercle ( Fig. 16 ) ....................................................................................................................................... T. harpagophora

b Antero-medial platelets medium to short, broad ( Fig. 11B ); rostrum mid-sized to long ( Figs 11C , 13C ); P4 only distally curved, ventral setae on flat hump .......................................................................................................................... 3

3a Idiosoma large (> 650 µm); coxal field broad, Cx-I tips triangular, slender, antero-lateral corner of Cx-III directed rostro-laterally; capitular bay elongated U-shaped; ventral margin of capitulum with sharp bend, rostrum mid-sized ( Figs 11 , 12 )................................................................................................................................................... T. costaricense

b Idiosoma mid-sized (<650 µm); coxal field slender, however, Cx-I tips broad, truncated, antero-lateral corner of Cx-III directed laterally; capitular bay wide-U-shaped, lateral margins +/– convex; ventral margin of capitulum sigmoid, rostrum elongated; P2 and P3 relatively long ( Figs 13 , 14 ) ........................................................... T. fontinale

TORRENTICOLA COSTARICENSE SP. NOV.

( FIGS 11A–E , 12A–D; TABLE 2)

Type series: Holotype male, CR 142, Guanacaste, ACG, Maritza, Quebrada Mata Redonda, small stream, 700 m asl, 28.ii.1996, mounted; paratypes, same locality and date, 1/0/0 mounted, 2/0/0 unmounted; CR 152, Guanacaste, ACG, Maritza , Quebrada Marilin , small stream, 560 m asl, 02.iii.1996, 0/1/0 mounted; CR 234, Guanacaste, ACG, Maritza , affluent to Laguna Mata Redonda (outlet of springs CR 233, 80 m below), small stream, 520 m asl, 04.iv.1996, 2/1/0 mounted, 5/3/0 unmounted; CR 234-IIa, Guanacaste, ACG, Maritza (outlet of springs CR 233, 10 m below) spring brook, 530 m asl, 21.ii.1997, 1/0/0 mounted, 0/2/0 unmounted; CR 234-IIb, Guanacaste, ACG, Maritza , affluent Laguna Mata Redonda (outlet of springs CR 233, 100 m below), small stream, 520 m asl, 21.ii.1997, 1/0/0 mounted.

Additional specimens examined: CR 5, Alajuela, stream, 900 m asl, 17.vi.1995, 0/1/0 mounted; CR 8, Alajuela, helocrene, 1600 m asl, 18.vi.1995, 1/0/0, mounted; CR 10, Heredia, El Tirol, stream, 1780 m asl, 19.vi.1995, 2/1/0 mounted; CR 25, Puntarenas, Ecolodge San Luis, spring next to Quebrada Alondra, rheocrene, 1020 m asl, 26.vi.1995, 4/0/0 mounted, 3/6/0 unmounted; CR 27, Puntarenas, Ecolodge San Luis, affluent to Río San Luis, spring brook, 1000 m asl, 27.vi.1995, 0/1/0 mounted; CR 28, Heredia, Varablanca, spring brook, 1960 m asl, 29.vi.1995, 1/0/0 mounted, 0/1/0 unmounted; CR 34, Alajuela, Río Sardinal, stream, 420 m asl, 30.vi.1995, 1/0/0 mounted; CR 40, Alajuela, Río Barranca, small stream, 1540 m asl, 01.vii.1995, 1/0/0 mounted; CR 45, Limón, Río Corinto , stream, 500 m asl, 04.vii.1995, 1/0/0 unmounted; CR 49, Cartago, affluent Río Coliblanco , small stream, 1540 m asl, 05.vii.1995, 1/0/0 unmounted; CR 51, Cartago, 5 km north Capellades, small stream, 1660 m asl, 05.vii.1995, 1/1/0 unmounted; CR 57, Alajuela, San Ramon Field Station , hygropetric area below waterfall at Río San Lorencito , 1040 m asl, 09.vii.1995, 2/0/0 mounted, 2/2/0 unmounted; CR 58, Alajuela, San Ramon Field Station , small stream, 940 m asl, 10.vii.1995, 1/0/0 mounted; CR 59, Alajuela, San Ramon Field Station , left affluent Río San Lorencito , small stream, 1000 m asl, 10.vii.1995, 3/0/0 mounted, 1/3/0 unmounted; CR 60, Alajuela, San Ramon Field Station , right affluent to Río San Lorencito , spring brook, 1080 m asl, 10.vii.1995, 5/2/0 mounted, 13/2/0 unmounted; CR 66, Puntarenas, Ecolodge San Luis, bank of Río San Luis, rheopsammocrene, 1230 m asl, 16.vii.1995, 1/1/0 mounted; CR 107, Puntarenas, Las Alturas Field Station , Río Colon , stream, 1340 m asl, 01.viii.1995, 1/ 0/0 mounted, 0/1/0 unmounted; CR 123, Guanacaste, ACG, Gongora, Río La Yengua , small stream, 560 m asl, 22.ii.1996, 2/0/0 mounted, 0/3/0 unmounted; CR 136, Guanacaste, ACG, Cacao, Quebrada Florcita, spring brook, 1180 m asl, 27.ii.1996, 2/0/0 mounted, 2/1/0 unmounted; CR 137, Guanacaste, ACG, Cacao, rheocrene, 1260 m asl, 27.ii.1996, 2/0/0 mounted, 1/3/0 unmounted; CR 138, Guanacaste, ACG, Cacao, spring brook, 1170 m asl, 27.ii.1996, 3/0/0 mounted, 3/0/0 unmounted; CR 139, Guanacaste, ACG, Cacao, rheocrene, 1150 m asl, 27.ii.1996, 1/0/0 mounted, 1/0/0 unmounted; CR 140, Guanacaste, ACG, Cacao, Quebrada Florcita, small stream, 740 m asl, 28.ii.1996, 2/0/0 mounted, 1/2/0 unmounted; CR 145, Guanacaste, ACG, Las Pailas, Rincon de la Vieja, small sulphurous stream, 740 m asl, 29.ii.1996, 0/1/0 mounted; CR 158, Cartago, NP Tapanti, small stream, 1420 m asl, 06.iii.1996, 1/2/0 mounted; CR 160, Cartago, NP Tapanti, Quebrada Palmitas, flowing small stream, 1500 m asl, 06.iii.1996, 1/0/0 mounted; CR 164, San José, La Fonda, Carretera Braulio, spring brook, 1480 m asl, 09.iii.1996, 2/0/0 mounted, 2/10/0 Male Female

Idiosoma L 701 753 667 849 44.3 800 746 868 40.2 Idiosoma W 594 618 544 755 52.6 672 613 721 34.8 Idiosoma L / W 1.18 1.20 1.11 1.30 0.05 1.18 1.13 1.29 0.05 Cx-I tL 314 314 284 358 14.5 321 304 329 8.4 Cx-III W 412 412 368 500 28.9 432 417 461 12.9 Cx-I tL/Cx-III W 0.76 0.75 0.67 0.82 0.04 0.73 0.69 0.77 0.03 Ds L 608 652 559 760 46.6 706 628 731 35.7 Dp L 574 618 525 706 44.8 666 594 697 35.7 Ds W 481 527 461 628 38.8 570 520 613 28.5 Ds L / W 1.27 1.24 1.16 1.29 0.03 1.23 1.17 1.27 0.03 Dp L / W 1.19 1.17 1.08 1.23 0.03 1.16 1.11 1.20 0.03 A-m platelet L 184 189 159 218 13.1 187 169 206 10.7 A-m platelet W 61 69 58 83 5.9 71 64 76 4.0 A-m platelet L/ W 3.00 2.69 2.44 3.00 0.1 2.71 2.46 2.93 0.1 Capitular bay L 164 164 135 181 10.2 178 159 184 9.2 Capitular bay W 69 76 64 102 8.1 87 74 98 7.6 Cb L / W 2.39 2.13 1.51 2.44 0.2 2.07 1.63 2.38 0.2 Dist cb – gf 216 224 203 279 15.7 181 174 191 5.6 Cx-I mL 152 157 135 194 10.6 145 132 153 5.6 Cx-II + III mL 54 59 39 78 8.0 27 25 42 5.0 Cx-I tL/Cx-II/III mL 5.82 5.35 4.00 8.63 0.9 11.83 7.42 13.41 1.7 Cx-I/Cx-II + III mL 2.82 2.67 1.87 4.25 0.4 5.50 3.41 5.90 0.8

unmounted; CR 208, Alajuela, San Ramon Field Station, left affluent to Río San Lorencito, small stream, 1000 m asl, 26.iii.1996, 9/2/0 mounted, 55/29/0 unmounted; CR 211, Alajuela, San Ramon, small stream, 700 m asl, 27.iii.1996, 5/1/0 mounted, 8/17/0 unmounted; CR 212, Alajuela, 12 km north San Ramon, Río Balsa , stream, 960 m asl, 27.iii.1996, 1/0/0 mounted; CR 216, Alajuela, Arenal, Río Agua Caliente , stream, 620 m asl, 29.iii.1996, 0/1/0 mounted; CR 223, Puntarenas, Ecolodge San Luis, Quebrada Alondra, small stream, 1100 m asl, 31.iii.1996, 1/0/0 mounted; CR 233, Guanacaste, ACG, Maritza, rheopsammocrene, 530 m asl, 04.iv.1996, 1/2/0 mounted, 0/1/0 unmounted; CR 233-II, Guanacaste, ACG, Maritza, rheopsammocrene, 530 m asl, 21.ii.1997, 4/4/0 mounted, 5/6/2 unmounted; CR 288, Guanacaste, Dos Ríos, Quebrada La Gato , small stream, 520 m asl, 03.ii.1997, 2/0/0 mounted; CR 289, Guanacaste, ACG, Nueva Zelandia, Río Cucaracho , small stream, 640 m asl, 03.ii.1997, 1/0/0 mounted, 0/2/0 unmounted; CR 291, Guanacaste, Río Sabalo , small stream, 540 m asl, 06.ii.1997, 1/0/0 unmounted; CR 299, Heredia, Zona Norte, affluent to Río Toro , small stream, 35 m asl, 09.ii.1997, 1/0/0 mounted, 1/4/0 unmounted; CR 309, Limón, Hitoy Cerere, spring brook, 200 m asl, 12.ii.1997, 0/1/0 unmounted; CR 310, Limón, Hitoy Cerere, small stream, 200 m asl, 12.ii.1997, 4/4/1 unmounted; CR 311, Limón, Hitoy Cerere, spring brook, 200 m asl, 12.ii.1997, 2/0/0 mounted, 13/8/1 unmounted; CR 312, Limón, Hitoy Cerere, left bank of Río Hitoy Cerere , rheopsammocrene, 190 m asl, 13.ii.1997, 2/0/0 mounted, 1/3/1 unmounted; CR 314, Limón, Hitoy Cerere, left affluent to Río Hitoy Cerere , small stream, 190 m asl, 13.ii.1997, 1/1/0 unmounted; CR 321, Limón, Bribri, waterfall, 110 m asl, 15.ii.1997, 1/0/0 mounted, 1/2/0 unmounted; CR 323, Guanacaste, ACG, Cacao, Quebrada San Josecito, small stream, 980 m asl, 20.ii.1997, 3/1/1 unmounted; CR 324, Guanacaste, ACG, affluent Quebrada San Josecito, small stream, 1000 m asl, 20.ii.1997, 0/2/0 unmounted; CR 325, Guanacaste, ACG, Cacao, bank of affluent to Quebrada San Josecito, rheopsammocrene, 1000 m asl, 20.ii.1997, 1/0/0 mounted, 1/0/0 unmounted; 01AA- MM-96, Guanacaste, ACG, Maritza , small stream, 05.v.1996, 1/5/0 unmounted, 02AA-MM-96, Guanacaste, ACG, Maritza , helocrene, 05.v.1996, 1/1/0 mounted, 3/3/0 unmounted; 03AA-MM-96 Guanacaste, ACG, Maritza , small stream, 05.v.1996, 77/79/0 unmounted; 05AA-MM-96, Guanacaste, ACG, Maritza , rheohelocrene, 05.v.1996, 11/12/0 unmounted .

Habitat: Mainly slow to fast flowing spring brooks, small streams, streams (also one polluted stream), rheocrenes, rheopsammocrenes, rheohelocrenes, some very fast flowing small streams, streams and waterfalls, some hygropetric areas and standing helocrenes (the only species also found in standing water!) at 35–1960 m asl (mainly 500–1500 m asl); mesolithal, akal, psammal (to lower proportions: macropelal, micropelal, macrolithal, lithophytal, leaf packages, terrestrial vegetation, phytal); temperature 15.0– 30.8 °C; conductivity 17–146 µS cm −1 (one site 1047 µS cm −1).

Distribution: Costa Rica (in all regions, mainly at mid elevations of the Cordillera de Guanacaste and Cordillera de Tilarán, also some sites in the Cordillera Central, northern, southern and Caribbean slope of the Cordillera de Talamanca).

Derivatio nominis: costaricense (Spanish = Costa Rican); referring to the wide distribution of the species all over the country.

Diagnosis: Characters of the ‘ Rusetria ’-like species; relatively variable species: idiosoma +/– rounded, midsized to large; antero-medial dorsal platelets large; coxal field broad, relatively short, medial margin of Cx-II/III relatively short; genital field compact; capitulum high, ventrally with sharp bend, posterior part ventrally straight; P2/P3 with strong cone-shaped projections, P4 distally curved; genital skeleton with elongated cella proximalis.

Description – Male (N = 78): Idiosoma mid-sized to relatively large [L 701 µm (667–849 µm)], rounded to oval [L/ W 1.18 (1.11–1.30)]; dorsal plate reddish or with pale red pattern ( Fig. 6 B-3), antero-medial platelets large, medially straight to convex, laterally slightly oblique; Dgl-4 medial to Dgl-5 ( Fig. 11B ); Cx-I tips relatively short, rounded, Cx-I/II/III laterally sharply graded; Cxgl-4 at the anterior tip of Cx-I; capitular bay U-shaped, lateral sides +/– straight, basely rounded; medial margin of Cx-II/III short ( Fig. 11A , Table 2); posterior margin of Cx-IV postero-lateral of caudal end of genital field, directed slightly from postero-medially to antero-laterally; excretory pore between Vgl-2, slightly posterior caudal margin of primary sclerotization; genital field anterior truncated, lateral margins slightly convex, posterior rounded, slightly tapering ( Fig. 11A ); genital skeleton relatively large, apically short, cella proximalis large with long, slender processus proximalia (aL/tL 0.28– 0.44), brachia distalia strong and short, brachia proximalia long ( Fig. 11E ); capitulum basely high, ventrally with sharp bend, rostrum mid-sized, basely high; palp strong, P2 ventrally slightly convex, dorsally convex in the proximal, straight in the distal part, ventro-distal projection strong, cone-shaped, pointed towards distal, P3 relatively long, ventrodistal projection smaller than projection of P2, coneshaped, pointed towards ventral, P4 distally slightly tapering, slightly curved, ventral setae on flat hump ( Fig. 11C, D ).

Female (N = 22): Idiosoma similar to male ( Table 2, Fig. 12A, B ), on an average slightly larger (L 746– 868 µm); medial margin Cx-II/III short; genital field rounded-rhombic ( Fig. 12A ); gnathosoma similar to male ( Fig. 12C, D ).

Discussion: This widespread species is relatively variable ( Table 2) and very euryoecious. According to the measurements no clear differentiation between populations are visible. A wide variety exists in the width of the capitular bay (L/ W 1.51 –2.44), the proportions of the genital field (L/ W 1.09 –1.33 in males, 0.99–1.07 in females), the palp segments and the genital skeleton. However, these differences are continuous and cannot be related to certain habitats (springs, streams), elevations or different geographical regions.

TORRENTICOLA FONTINALE SP. NOV.

( FIGS 13A–E , 14A–D, TABLE 3)

Type series: Holotype male, CR 201, Puntarenas, Peninsula de Osa, above Los Angeles de Drake , Los Migueles, rheopsammocrene, 150 m asl, 23.iii.1996, mounted; paratypes, same locality and date, 2/3/0 mounted, 4/7/0 unmounted.

Additional specimens examined: CR 184, Puntarenas, Peninsula de Osa, Finca de Juan Francisco Sanchez , rheocrene, 130 m asl, 17.iii.1996, 1/0/0 mounted .

Habitat: Slow flowing rheopsammocrenes and rheocrenes at low elevations (130–150 m asl) in rainforest; akal, psammal, mesolithal; temperature 24.9 °C; conductivity 160–194 µS cm −1.

Distribution: Costa Rica (Peninsula de Osa).

Derivatio nominis: fontinale (Latin = dedicated to the god of springs); as the species has exclusively been found in several springs.

Diagnosis: Characters of the ‘ Rusetria ’-like species; idiosoma oval; antero-medial dorsal platelets relatively long; dorsal plate reddish; coxal field slightly elongated, anterior tips of Cx-I noticeably truncated, very broad; capitular bay very wide, lateral margins slightly convex; rostrum elongated (especially in males, female bear thicker rostrum), ventral margin of capitulum flatly sigmoid curved; P2 with strong coneshaped ventro-distal projection.

Description – Male (N = 4): Idiosoma mid-sized [L 613 µm (628–647 µm)], oval [L/ W 1.36 (1.32–1.35)]; dorsal plate rounded-oval [L/ W 1.18 (1.15–1.18)], antero-medial platelets relatively large, medially rounded, laterally straight, slightly oblique, Dgl-4 lateral to Dgl-5 (Dgl-3, -4, -5 lay on a straight line) ( Fig. 13B ); coxal field elongated [Cx-I tL/Cx-III W 0.86 (0.84–0.87)], especially Cx-II/III elongated, Cx-I more compact, anterior tips truncated, very broad; Cxgl-4 antero-laterally on the truncated Cx-I tips; capitular bay large, very wide, lateral margins slightly convex; medial margin of Cx-II/III relatively short; posterior margins of Cx-IV postero-lateral of genital field, oblique towards antero-lateral; excretory pore on caudal margin of primary sclerotization, slightly anterior to the very close Vgl-2; genital field in posterior half of ventral shield, small [L/Id L 0.22 (0.21–0.22)], elongated, anterior truncated, lateral margins straight, posterior tapering ( Fig. 13A ); genital skeleton apically small, triangular, cella proximalis large, without processus proximalia [aL/tL 0.36 (0.25–0.34)], brachia proximalia well developed, elongated, brachia distalia very weak, merely visible ( Fig. 13E ); capitulum ventrally flatly curved, basely nearly straight, at base of rostrum heavily sculptured, rostrum straight, long; P2 and P3 very long, ventro-distal projection of P2 large cone-shaped, that of P3 smaller, truncated; P4 relatively small [P2/P4 1.44 (1.44)], without ventral projections, distally tapering, relatively straight ( Fig. 13C, D ).

Female (N = 3): Idiosoma similar to male, larger (L 647–726); dorsal plate broader (L/ W 1.12 –1.15), anterior truncated ( Fig. 14B ); coxal field wider than in male (Cx-I tL/Cx-III W 0.75 –0.85), Cx-II/III laterally more graded; capitular bay even wider, lateral margin clearly convex; medial margin of Cx-II/III very short ( Table 3, Fig. 14A ); genital field large, rhombic, anterior truncate, laterally +/– straight, posterior tapering; excretory pore between to slightly posterior Vgl-2, pore and glands posterior to caudal margin of primary sclerotization ( Fig. 14A ); capitulum and rostrum higher, shorter than in males ( Fig. 14C ), palp similar to male ( Fig. 14C, D ).

Discussion: The combination of a ‘ Rusetria -like’ dorsal shield and a long and slender rostrum characterizes this species, furthermore, it is typified by the very wide capitular bay and the very broad truncated anterior tips of the Cx-I. The absence of processus proximalia at the cella proximalis is also very characteristic. Together with T. amalgamada these are the only species of the genus in their distribution restricted to rainforest springs on the Peninsula de Osa in south-western Costa Rica. These two species can be regarded as relicts of the ancient fauna of the Nicoya Complex, the oldest geological formation in southern Central America ( Savage, Flowers & Porras, 2005).

TORRENTICOLA GRADATICOXALIS K.O. VIETS, 1977

( FIG. 15A–E; TABLE 4)

Type series: Holotype male, Guatemala, km 148 road Guatemala Ciudad to Cobán , at Niño Perdido, Quebrada del Niño, 11.vii.1974, leg. Böttger, prep. no. Male Female

Idiosoma L 613 633 613 647 14.6 716 647 726 42.8 Idiosoma W 451 468 451 491 16.2 540 520 549 15.0 Idiosoma L / W 1.36 1.35 1.32 1.36 0.02 1.30 1.25 1.35 0.1 Cx-I tL 280 284 280 289 5.7 309 280 324 22.5 Cx-III W 324 334 324 343 8.0 383 373 383 5.7 Cx-I tL/Cx-III W 0.86 0.85 0.84 0.87 0.01 0.81 0.75 0.85 0.05 Ds L 491 510 491 530 17.9 559 520 579 30.0 Dp L 461 483 461 500 17.2 540 495 549 28.7 Ds W 392 414 392 432 16.2 481 432 481 28.3 Ds L / W 1.25 1.24 1.21 1.25 0.02 1.20 1.16 1.20 0.02 Dp L / W 1.18 1.17 1.15 1.18 0.01 1.14 1.12 1.15 0.01 A-m platelet L 135 142 135 149 6.3 154 152 156 1.9 A-m platelet W 49 53 49 54 2.3 51 51 54 1.4 A-m platelet L/ W 2.75 2.75 2.59 2.77 0.1 2.95 2.86 3.02 0.1 Capitular bay L 123 129 123 132 4.2 147 145 152 3.7 Capitular bay W 81 85 81 91 4.2 108 105 110 3.5 Cb L / W 1.52 1.51 1.46 1.54 0.04 1.35 1.33 1.37 0.03 Dist cb – gf 225 232 221 243 10.3 191 172 192 11.7 Cx-I mL 159 158 145 164 8.3 162 142 176 17.2 Cx-II + III mL 59 70 59 71 5.8 18 17 20 1.7 Cx-I tL/Cx-II/III mL 4.75 4.07 4.07 4.75 0.3 16.03 15.77 16.30 0.4 Cx-I/Cx-II + III mL 2.71 2.26 2.11 2.71 0.3 8.27 8.25 8.29 0.0 Genital field L 132 134 130 140 4.2 159 147 174 13.5 Gf L /Cx-II + III mL 2.25 1.93 1.89 2.25 0.2 8.35 8.13 8.57 0.3 Genital field W 96 102 96 105 5.1 147 140 152 6.2 Genital field L/ W 1.38 1.33 1.28 1.38 0.04 1.08 1.05 1.15 0.05 Gf L / Id L 0.22 0.21 0.21 0.22 0.01 0.23 0.22 0.24 0.01 Gf L /dist cb – gf 0.59 0.58 0.57 0.59 0.01 0.86 0.83 0.91 0.04 Dist gf – expo 74 92 74 105 15.9 152 135 159 12.6 Dist gf – cauda 137 142 130 147 8.3 208 186 233 23.3 Gs L 213 216 211 221 4.5

Gs aL 76 69 54 76 9.5

Gs W 118 116 108 137 12.6

Gs aL/tL 0.36 0.32 0.25 0.36 0.05

Gs tL/ W 1.81 1.85 1.54 2.05 0.2

Capitulum vL 299 313 299 321 9.2 363 341 363 12.7 Capitulum dL 225 232 225 238 5.1 274 257 274 9.9 Rostrum L 126 128 120 132 5.3 140 132 142 5.1 Capitulum H 115 121 115 125 4.2 157 145 159 7.9 R L/c dL 0.56 0.56 0.52 0.56 0.02 0.51 0.51 0.52 0.00 R L/c vL 0.42 0.41 0.38 0.42 0.02 0.39 0.39 0.39 0.00 Gn bend depth 15 15 15

Chelicera L 360 376 360 380 9.2 443 436 451 10.4 Chelicera H 29 31 29 32 1.4 34 32 39 3.7 Chelicera L /H 12.25 12.09 11.69 12.92 0.5 12.60 11.50 13.69 1.6 Chelicera bs L 301 314 301 316 6.6 366 360 372 8.7 Chelicera claw L 59 61 59 66 3.7 78 76 81 2.4 Chel bs/claw L 5.13 5.04 4.74 5.33 0.3 4.75 4.74 4.75 0.01 P1 dorsal L 37 37 37 38 0.6 44 44 44 0.00 P2 dL 116 120 116 120 1.8 142 132 145 6.5 P3 dL 64 67 64 69 2.3 78 74 78 2.8 P4 dL 81 83 81 83 1.2 93 88 93 2.8 5678 SMF; allotype female, Guatemala, km 150–151 Road Guatemala Ciudad to Cobán, Río Lima, 1520 m asl, 16.viii.1974, leg. Böttger, prep. no. 5727 SMF .

Further material: Guatemala, leg. Böttger: near Finca Sacté , north-west Cobán, Río Cuxja, 800 m asl, 24.vii.1974, 1/0/0 mounted, prep. no. 5963 SMF; km 150–151 road Guatemala Ciudad to Cobán, Río Lima, 1520 m asl, 17.viii.1974, 1/0/0 mounted, prep. no. 6274 SMF. (According to slide material at SMF, not all mentioned in K.O. Viets, 1977/78 Teil I) .

Habitat: Small mountain streams.

Geographical distribution: Guatemala.

Published records: K.O. Viets 1977/78 Teil I.

Diagnosis: Characters of the ‘ Rusetria ’-like species; idiosoma rounded-oval; dorsal plate reddish, anteromedial dorsal platelets large; lateral margins of coxal field very sharply graded, antero-lateral corner of Cx- II forms an anterior pointed edge, medial margin of Cx-II/III in male very short, in female Cx-III medially meet at genital field; capitulum basely high, ventral margin proximally oblique, with sharp bend towards mid-sized rostrum.

Description: See K.O. Viets 1977/78 Teil I. Discussion: The species is clearly characterized by the shape of the lateral margin of the coxal field. In the description of T. gradaticoxalis, K.O. Viets (1977/78 Teil I) mentioned that the caudal end of the anterolateral dorsal platelets would be unclear, actually – according to new investigations of the slide material at the SMF (see above) – these platelets are completely fused with the main dorsal plate.

TORRENTICOLA HARPAGOPHORA SP. NOV.

( FIG. 16A–D; TABLE 4)

Type series: Holotype female, CR 162, Limon, right affluent of Río Corinto , spring brook, 500 m asl, 07.iii.1996, mounted.

Additional specimens examined: CR 215, Alajuela, Finca Cerro Chato, affluent Río Agua Caliente , small stream, 760 m asl, 29.iii.1996, 0/1/0, mounted; CR 227, Guanacaste, Tenorio, spring brook, 1000 m asl, 02.iv.1996, 0/1/0 unmounted; CR 285, Guanacaste, ACG, Sta. Maria, Rincon de la Vieja, spring at right bank of Río Negro, rheopsammocrene, 750 m asl, 31.i.1997, 0/1/0, mounted .

Habitat: Fast flowing spring, slow and fast flowing spring brooks and small stream at mid elevations gradaticoxalis harpagophora

Idiosoma L 780 932 706 765 706 780 38.9 Idiosoma W 647 824 618 652 618 677 29.6 Idiosoma L / W 1.20 1.13 1.14 1.15 1.14 1.17 0.02 Cx-I tL 353 358 294 314 294 324 15.0 Cx-III W 441 471 422 446 422 456 17.7 Cx-I tL/Cx-III W 0.80 0.76 0.70 0.70 0.70 0.71 0.01 Ds L 697 814 638 687 638 716 39.6 Dp L 643 755 618 662 618 677 30.6 Ds W 589 652 505 525 505 544 19.6 Ds L / W 1.18 1.25 1.26 1.31 1.26 1.32 0.03 Dp L / W 1.09 1.16 1.22 1.24 1.22 1.26 0.02 A-m platelet L 201 218 181 186 181 191 6.9 A-m platelet W 78 86 49 54 49 59 6.9 A-m platelet L/ W 2.56 2.54 3.70 3.48 3.25 3.70 0.3 Capitular bay L 164 172 152 162 152 164 6.5 Capitular bay W 74 88 103 100 96 103 3.7 Cb L / W 2.23 1.94 1.48 1.61 1.48 1.72 0.1 Dist cb – gf 255 212 186 189 186 194 3.7 Cx-I mL 196 194 149 154 149 162 6.2 Cx-II + III mL 51 15 27 25 20 27 3.7 Cx-I tL/Cx-II/III mL 6.86 24.36 10.92 12.81 10.92 16.52 2.8 Cx-I/Cx-II + III mL 3.81 13.17 5.55 6.30 5.55 8.25 1.4 Genital field L 181 213 167 174 167 174 4.2 Gf L /Cx-II + III mL 3.52 14.50 6.18 7.10 6.18 8.88 1.4 Genital field W 147 184 154 163 154 165 5.8 Genital field L/ W 1.23 1.16 1.08 1.07 1.05 1.08 0.01 Gf L / Id L 0.23 0.23 0.24 0.23 0.22 0.24 0.01 Gf L /dist cb – gf 0.89 0.90 0.89 0.92 0.01 Dist gf – expo 145 230 149 154 149 174 13.0 Dist gf – cauda 186 338 208 240 208 252 22.8 Gs L 252

GsaL 96

Gs aL/tL 0.38

Capitulum vL 292 319 317 337 317 355 19.0 Capitulum dL 225 223 240 265 240 270 15.8 Rostrum L 120 108 105 113 105 123 8.6 Capitulum H 149 167 176 167 176 5.7 R L/c dL 0.53 0.48 0.44 0.44 0.43 0.45 0.01 R L/c vL 0.41 0.34 0.33 0.33 0.33 0.34 0.01 Gn bend depth 22 12 17 12 18 3.2 Chelicera L 360 412 439 412 451 20.1 Chelicera H 27 42 42 40 42 0.7 Chelicera L/H 13.36 9.88 10.82 9.88 10.85 0.6 Chelicera bs L 296 343 365 343 380 18.5 Chelicera claw L 64 69 71 69 74 2.5 (500–1000 m asl); mesolithal, akal, leaf packages, macropelal, psammal, lithophytal, macrolithal and micropelal; temperature 20.1–21.1 °C; conductivity 21–86 µS cm −1.

Distribution: Costa Rica (scattered on the Caribbean and Pacific slope of the Cordillera Central, Cordillera de Tilarán and Cordillera de Guanacaste).

Derivatio nominis: harpago (Latin = grapnel), phor - (Greek = carry); referring to the curved P4 and hookshaped P5-bristles.

Diagnosis (only females): Characters of the ‘ Rusetria ’- like species; idiosoma rounded; dorsal plate reddish or with reddish pattern; antero-medial dorsal platelets long and narrow; coxal field short and broad, Cx-II/III medial margin relatively short, capitular bay relatively wide; capitulum ventrally curved with deep bend, rostrum relatively short; chelicera compact; P3 relatively long, P4 compact, curved, with ventral bristles on a strong, characteristic tubercle, bristles at P5 strong.

Description – Male: Unknown.

Female (N = 3): Idiosoma medium sized, very broad, rounded, nearly circular [L 706 µm (765–780 µm), L/ W 1.14 (1.15–1.17)] ( Fig. 16A ); dorsal plate with reddish pattern ( Fig. 6 B-1, 6B-3, very pale in one specimen), antero-medial platelets long and narrow ( Table 4), medially rounded, laterally oblique ( Fig. 16B ) (in one specimen fused medially); Dgl-4 clearly medial to Dgl-5 ( Fig. 16B ); coxal field wide [Cx-I tL/Cx-III W 0.70 (0.70–0.71)]; Cx-II/III laterally sharply graded; Cxgl-4 slightly posterior the anterior tip of Cx-I; capitular bay +/– U-shaped, diverging, basely rounded; medial margin of Cx-II/ III short; posterior margin of Cx-IV postero-lateral of caudal end of genital field, across ( Table 4, Fig. 16A ); excretory pore between Vgl-2; genital field nearly rounded, broad, only slightly tapering posterior ( Fig. 16A ); ventral margin of capitulum sigmoid curved, rostrum relatively short and thick; palp strong, P1 short, P2 with nearly straight ventral and dorsal margin, ventro-distal projection strong, triangular, distally pointed, P3 relatively long, ventro-distal projection of equal size as projection of P2, hook-shaped, curved to proximal, P4 compact, strongly curved, ventral bristles inserted on a strong, truncated cone, P5 inserted ventro-distally at P4, strong with heavy, hooked claws, inserted nearly ventro-distally at P5 ( Fig. 16C, D ); chelicera strong, compact ( Table 4, Fig. 16C ).

Discussion: At each of the four sample sites only one female specimen was found. Nevertheless, the very slender antero-medial dorsal platelets as well as the characteristic shape of the P4 clearly differentiate this species.

Figures

Figure 6. Colour patterns on the dorsal plates of Torrenticola (schematic). A, complete dorsal plate reddish to red (e.g. T. amalgamada, T. gradaticoxalis, T. fontinale, T. ratoncitoi, T. menudopalpis, T. elhachensis, T. flexirostris, T. lamellipalpis, T. obliquipalpis, T. cirratipalpis, T. australis, T. monticola, T. brevicoxalis, T. conipalpis, T. conirostris, T. semicolor (partim), T. cortobrazo, T. altifontana, T. esferica, T. ambigua (partim), T. torpebrazo, T. fastigata, T. chicacoxalis, T. rubella, T. levicoxalis, T. collina); B, hour-glass-shaped with pale ‘shoulder-patches’ (B-1), in paler specimens reduced to larger posterior and smaller anterior reddish to red patch (B-2, B-3) (e.g. T. harpagophora, T. costaricense, T. guanacastensis, T. keesdavidsi, T. ambigua (partim)]; C, posterior 1/2–2/3 red, margin (sharp to unclear) slightly V-shaped indented (C-1) (e.g. T. gennada, T. amala, T. semicolor (partim), T. pervagata (partim)], +/– straight (C-2) (e.g. T. alargada, T. columbiana, T. semicolor (partim), T. maceripalpis, T. acuticoxalis, T. bicolor, T. curtipalpis), with rostral extension (C-3) (e.g. T. alexandra, T. carlbaderi) or reduced to crescent-shaped, pale red patch in posterior half (C-4) (e.g. T. esquinada, T. tilaranensis, T. pervagata (partim)].

Figure 11. Torrenticola costaricense. A–D, holotype male (CR 142); E, male (CR 25). A, idiosoma, ventral view; B, idiosoma, dorsal view; C, capitulum with right palp, lateral view; D, left palp, medial view; E, genital skeleton, anterior view. Scale bars = 100 µm.

Figure 12. Torrenticola costaricense. A–D, paratype female (CR 152). A, idiosoma, ventral view; B, idiosoma, dorsal view; C, capitulum with right palp, lateral view; D, left palp, medial view. Scale bars = 100 µm.

Figure 13. Torrenticola fontinale. A–E, holotype male (CR 201). A, idiosoma, ventral view; B, idiosoma, dorsal view; C, capitulum with right palp, lateral view; D, left palp, medial view; E, genital skeleton, anterior view. Scale bars = 100 µm.

Figure 14. Torrenticola fontinale. A–D, paratype female (CR 201). A, idiosoma, ventral view; B, idiosoma, dorsal view; C, capitulum with right palp, lateral view; D, left palp, medial view. Scale bars = 100 µm.

Figure 15. Torrenticola gradaticoxalis. A–D, holotype male, prep. no. 5678 SMF, Viets collection; E, allotype female, prep. no. 5727 SMF, Viets collection; after K.O. Viets (1977/78, Teil I). A, idiosoma, ventral view; B, idiosoma, dorsal view; C, capitulum, lateral view; D, left palp, medial view; E, genital field and central part of coxal field. Scale bars = 100 µm.

Figure 16. Torrenticola harpagophora. A–D, holotype female (CR 162). A, idiosoma, ventral view; B, idiosoma, dorsal view; C, capitulum with right palp, lateral view; D, left palp, medial view. Scale bars = 100 µm.