Tengella perfuga Dahl, 1901

Leister, Matthew, Mallis, Rachael & Miller, Kelly, 2013, The male of Tengella perfuga Dahl, 1901 with re-description of the female and comparisons with T. radiata (Kulczynski, 1909) (Araneae: Tengellidae), Zootaxa 3709 (2), pp. 185-199 : 187-193

Tengella perfuga Dahl, 1901

Figs. (2, 3, 4A, C–F 5, 6A–C, G, H, 7A, B)

Diagnosis. Tengella perfuga can best be distinguished from other Tengella species by genitalic characters in both males and females. T. perfuga can be distinguished from T. thaleri by smaller overall body size ( T. thaleri male = 6.0mm, female 8.1mm, T. perfuga see Description), as well as the lack of an inner lobe in the male palpal RTA, the presence of a denticulate median apophysis, the presence of thick ventral tarsal scopulae and different septum shape and size in the female epigynum ( Platnick 2009). Whereas only the male is known from T. albolineata , it differs from other Tengella species in carapace colour pattern and palpal structure, particularly the sclerotised portion of the embolus which appears to have a terminal apophysis ( Wolff 1977; Platnick 2009). Both of these species have been collected in or associated with caves in Mexico, whereas T. perfuga and T. radiata are not known from caves.

Whereas Tengella perfuga and T. radiata have similar morphology, spin highly similar webs (pers. obs.), and occupy similar habitats, they can still readily be distinguished from each other. Colour and patterning is more defined in T. radiata ( Figs. 3A , 4 ). The male palp of T. perfuga has three prolateral tibial spines, whereas T. radiata has two ( Figs. 5A, B , 6A, B, D, E ). T. perfuga has a large cylindrical RTA bearing terminal denticles, whereas the RTA of T. radiata is slender and bears a three-lobed anterior projection ( Figs. 6 G–J). The sclerotised median apophysis has a large prominent basal tooth along with numerous small serrations in T. perfuga , whereas T. radiata lacks the basal tooth ( Figs. 5B, C , 6B, C, E, F ) ( Wolff 1977; Platnick 2009). The female epigyna are even more distinguishable, with the septum of T. perfuga being more rounded and narrowed abruptly anteriorly, and the septum broader and more angular in T. radiata ( Figs. 5D–E , 7 ).

Lectotype designation. The type series of Tengella perfuga Dahl, 1901 includes two female syntypes from “Süd Amerika?”, deposited in the Museum für Naturkunde at Humboldt-Universität, Berlin, each of which were examined for this study. One female specimen (ZMB 34656) was used by Dahl (1901) for internal dissection and due to its poor condition, serves little diagnostic or morphologic use. The second specimen (ZMB 34657) is mostly intact with the exception of a dissected epigynum that has since been lost. This specimen, Tengella perfuga (“Süd Amerika?”) (ZMB 34657), is designated as the lectotype.

Other material examined. Nicaragua: Matagalpa Dpto: Selva Negra, near Matagalpa, (12.9836°N, 85.9002°W), 1251m 0 9 November 2010, K.B. Miller, J. Smith coll., 1 male ( MSBA 023929), 3 females ( MSBA 24989, 24990, 24991); same locality, 25–30 May 2012, K.B. Miller, R. Mallis, M. Leister coll., 3 males ( MSBA 24980, 24981, 24982), 2 females ( MSBA 24985, 24986); Fuente Pura, 17 August 1994, J. M. Maes coll., 1 male ( MSBA 24983, 29083).

Description of male. Body length (excluding chelicerae and spinnerets) = 11.15–15.50mm (N = 5, mean = 13.85mm); carapace light brown with thin black marginal line, thick white sub-marginal lines, thin white medial line, pair of angled white lines anteriorly, and faint black lines radiate from thoracic groove; abdomen dark brown with white mottling that increases posteriorly, discontinuous white longitudinal lines present anteriorly, three pairs of white spots extending posteriorly, and faint chevron pattern on the posterior portion of the abdomen ( Figs. 3A , 4C ); legs pale yellow, darker distally, femora with four dark annulations, tibia with three to four dark annulations; metatarsi IV with reduced calamistrum, scopula extending half length of all metatarsi; tarsi with thick scopula, two primary toothed tarsal claws and a small third claw lacking teeth. Carapace pyriform in shape, length = 5.80– 7.80mm (mean = 6.92mm), width at anterior coxae II = 4.21–5.71mm (mean = 5.14mm) ( Fig. 3A , 4C , Table 1), abruptly narrowed at anterior half of coxae I, thoracic groove in middle of carapace, deep, longitudinal; surface covered in short black setae, sparse longer black setae surrounding thoracic groove, carapace margins, and ocular area; eight eyes in two straight rows ( Fig. 3A , 4C ), anteriorly, PER slightly procurved ( Fig. 3B ); all eyes round and subequal in size, AME slightly larger, spacing and measurements as in Table 1; corners of clypeus enclose prominent cheliceral boss ( Fig. 3B ); chilum divided; chelicerae vertical and setose; promargin with three equally spaced teeth, median tooth largest, retromargin with four larger, equally sized and spaced teeth, retromarginal teeth more distally placed on the chelicerae than promarginal teeth; endites rectangular, longer than wide, lateral margin narrowed at approximately one third of length, broader distally, diverging, laterodistal portion rounded with a distinct angle at level of palp coxae, slightly darkened at proximal medial edge near labium, surface covered with dense setae on distomedial margin, long erect setae covering lateral portion of endites; labium rectangular, approximately two thirds length of endites, proximal margins angled, with shallow notch at base, dense setae covering anterior margin; sternum shield-shaped, narrowing at level of coxae III, forming a point, divided medially by pale longitudinal stripe with radiating darker lines extending towards coxae; pedicel long, thin, and sclerotised ( Fig. 4D ).

Cribellum bipartite; ALS conical and two-segmented, PLS cylindrical and two-segmented, PLS more slender and slightly shorter than ALS, PMS small, cylindrical.

Leg formula 1 4 2 3 ( Table 1). Leg spination as follows, with only spine-bearing surfaces lised: Femora: I d1- 1-1, p1-2-1, r1-2-1; II d1-1-1, p1-2-1, r1-2-1; III d1-1-1, p1-2-1, r1-2-1; IV d1-1-1, p1-1-1, r0-2-1; Palp d0-1-2, p0- 0-1; Tibiae: I p0-2-0, v2-4 -2; II p1-0-1, v2-4 -2, r1-0-1; III d1-0-0, p1-1-1, v2-2 -2, r1-1-1; IV d1-0-1, p1-0-1, v2-2 - 2, r1-1-1; Palp d0-1-0, p2-0-1; Metatarsi: I p0-0-1, v2-2 -1, r0-0-1; II d0-0-1, p1-1-1, v2-2 -1, r0-1-1; III d0-1-2, p1- 1-1, v2-2 -2, r1-1-1; IV d2-0-2, p0-1-1, v2-2 -1, r0-2-1.

Mean(range) Mean(range) Body Leg II

Clypeus height 0.56(0.49–0.71) Coxa II 2.18(1.86–2.50) Carapace length 6.92(5.80–7.80) Trochanter II 1.02(0.75–1.15) Carapace width 5.14(4.21–5.71) Femur II 8.42(7.18–9.32) Abdomen length 6.93(5.35–7.89) Patella II 2.73(1.90–3.30) Body length 13.85(11.15–15.50) Tibia II 7.81(6.67–9.48) Eyes Metatarsus II 11.26(7.18–9.15) PME width 0.26(0.21–0.32) Tarsus II 3.57(3.25–3.80) AME width 0.35(0.30–0.41) Total leg II 33.80 (28.79–38.65) PLE width 0.30(0.30–0.31) Leg III

ALE width 0.31(0.30–0.31) Coxa III 1.93(1.78–2.10) PME-PME 0.18(0.08–0.21) Trochanter III 0.91(0.69–1.05) PME-PLE 0.33(0.30–0.35) Femur III 7.29(5.58–8.30) PLE-ALE 0.10(0.08–0.11) Patella III 2.38(1.85–2.61) AME-ALE 0.09(0.08–0.11) Tibia III 5.83(4.90–6.90) AME-AME 0.09(0.07–0.11) Metatarsus III 6.95(6.08–7.51) OQL 0.87(0.70–0.95) Tarsus III 2.90(2.45–3.35) OQW 1.87(1.60–2.15) Total leg III 28.18(23.61–30.85) Leg I Leg IV

Coxa I 2.47(2.13–2.70) Coxa IV 2.09(1.83–2.30) Trochanter I 1.14(0.95–1.20) Trochanter IV 1.09(0.88–1.20) Femur I 10.29(9.20–10.90) Femur IV 9.44(8.00–10.23) Patella I 3.07(2.61–3.40) Patella IV 2.49(2.00–2.89) Tibia I 11.19(9.72–13.18) Tibia IV 8.09(7.55–8.89) Metatarsus I 11.26(9.93–12.70) Metatarsus IV 10.48(8.20–11.55) Tarsus I 4.76(4.48–5.10) Tarsus IV 3.63(3.25–4.19) Total leg I 44.17(39.02–48.89) Total leg IV 37.30(31.71–41.25)

Pedipalp

Palp trochanter 0.60(0.38–0.70)

Palp femur 3.28(2.45–3.70)

Palp patella 1.38(1.05–1.60)

Palp tibia 1.81(1.47–2.00)

Palp tarsus 2.36(1.85–2.70)

Total palp 9.43(7.20–10.41) Palp tibia widened distally; RTA large, heavily sclerotised, distally darkened, bearing large denticles on distal portion ( Figs. 5A–C , 6A–C , G, H); cymbium densely setose with hooked setae; tegulum with large median apophysis, sclerotised, curved dorsomedially, with prominent basal tooth followed by row of small serrated denticles which begin at middle and extend along medial edge; embolus broadened at base, strongly curving distally above median apophysis; hyaline conductor extending medially between embolus and median apophysis, dorsally cupping the embolus distally ( Figs. 5C , 6C ).

Female. Total body length (excluding chelicerae and spinnerets) = 13.76–18.05 (N= 5, mean = 15.72mm, Table 2 ); colour and pattern of carapace, abdomen and legs as in male, but patterning more distinct ( Fig. 4A, E ); calamistrum more prominent than in male, beginning at one-fourth and terminating at one-half length of metatarsus IV ( Fig. 2C ), scopula extending half length of metatarsi; tarsi scopulate with two primary toothed tarsal claws and a single small claw lacking denticles as in male. Carapace pyriform in shape as in male, length = 5.81–7.55mm (mean = 6.85mm), width = 4.55–5.55mm (mean = 5.07mm, Table 2 ); thoracic groove as in male; surface setae and eight eyes in two rows as in male (eye measurements in Table 2 ); clypeus enclosing cheliceral boss as in male; chilum, cheliceral setae and shape, marginal teeth, endites, labium and sternum as in male ( Fig. 4A, E, F ).

Cribellum bipartite ( Fig. 2B ), more prominent than in male; ALS and PLS as in male.

Leg formula 1 4 2 3 ( Table 2 ). Leg spination as follows, with only spine-bearing surfaces lised: Femora: I d1- 1-1, p0-2-1, r0-2-2; II d1-1-1, p1-1-1, r1-1-2; III d1-1-1, p1-2-1, r1-2-1; IV d1-1-1, p0-1-1, r0-1-1; Palp d0-0-4, p0- 0-1; Tibiae: I v2-4 -2; II p0-1-1, v2-4 -2; III d1-1-0, p0-1-1, v2-2 -2, r0-1-1; IV d1-0-0, p0-1-1, v2-2 -2, r0-1-1; Palp d0-1-1, p2-0-1; Metatarsi: I v2-2 -3; II p0-1-2, v2-2 -1, r0-0-1; III d1-1-0, p1-1-2, v2-2 -1, r1-1-2; IV d0-1-2, p1-1-1, v2-3 -1, r1-1-1; Palp tarsus: p2-1-0, r0-1-0.

Epigynum with median septum round, narrowed abruptly anteriorly, with sclerotised mating plugs present on all examined females; dorsally, spermathecae short, oval structures with a single twist ( Figs. 5D, E , 7A, B ).

Colour in life. Since some of the specimens used for this description have been preserved in 70% EtOH, the pigmentation in the setae and the exoskeleton is faded. This makes the specimens lighter in colour than they appear in life, with degradation of patterning. In life, the overall colouration is dark brown, with all lines and accessory patterning yellow ( Fig. 4A ). In the male, sclerotised portions of the RTA, cymbium, and portions of the chelicerae and clypeus also have subtle blue green iridescence. This iridescence is present on the chelicerae and clypeus in females, but less distinctly.

Sexually dimorphic features. Males have longer legs than females, most notably leg I, and the abdomen in females is markedly larger than in males ( Table 1, Table 2 ), and both leg I and abdomen are used in courtship behaviour (pers. obs. of the authors). Spination also varies between the sexes.

Intrasexual variation. Within the sexes, there is little variation in colour pattern and spination; however body size and leg lengths vary. For adult males, body length = 11.15–15.50mm, whereas in adult females body length = 13.76–18.05mm ( Table 1, Table 2 ).

Natural history. These are medium to large cribellate spiders that build sheet webs with a funnel retreat and significant cribellate tangle structure above the web. Webs in the field were typically along stream embankments, tree trunks and between stones on a stone bridge and builidngs. These were comprised of a broad more round sheet with a funnel retreat below the web either in a rear corner or in the center of the web. The main sheet and retreat were surrounded by a scaffolding of tangle lines above, and some anchor lines below the sheet. Cribellate silk was integrated throughout all parts of the funnel web. The spider spends most of its time in the retreat or just at the retreat opening and runs out on top of the sheet to capture prey and back to the retreat. Tengella perfuga is found in Nicaraguan high elevation cloud forests (1251m), often associated with coffee plantations, in close proximity to conspecifics along banks of streams, tree trunks, sides of bridges and other manmade structures. Specimens were collected by hand from webs.

Distribution. Nicaragua, high elevation regions in Departamento Matagalpa and Jinotega ( Fig. 1 ).

Figures

FIGURE 1. Map of published localities for Tengella radiata and new localities for Tengella perfuga and T. radiata.

FIGURES 3 A – B. Tengella perfuga; male. A. Carapace and abdomen, dorsal view; B. Carapace and chelicerae, anterior view. (Original illustrations by Matthew Leister).

FIGURES 4 A – J. Tengella spp., habitus. A. T. perfuga female, alive, in field, Nicaragua; B. T. radiata female, alive, in field, Nicaragua; C, D. T. perfuga male, preserved; E, F. T. perfuga female, preserved; G, H. T. radiata male, preserved; I, J. T. radiata female, preserved.

FIGURES 5 A – E. Tengella perfuga genitalia. A – C. Left male palp; A. prolateral view; B. ventral view; C. retrolateral view; D – E. Epigynum; D. ventral view; E. dorsal view. Con—conductor, Em—embolus, MA—median apophysis, RTA—retrolateral tibial apophysis, Sp—spine attachments. (Original illustrations by Matthew Leister).

FIGURES 6 A – F. Comparison of male genitalia, left palp. G – J. Comparison of male RTA. A – C, G, H. Tengella perfuga; D – F, I, J. T. radiata; A, D. Prolateral view; B, E, G, I. Ventral view; C, F, H, J. Retrolateral view. Con—conductor, Em—embolus, MA—median apophysis, RTA—retrolateral tibial apophysis, Sp—spines and spine attachments.

FIGURES 7 A – D. Comparison of female genitalia. A, B. Tengella perfuga; C, D. T. radiata; A, C. Epigynum, ventral view; B, D. Epigynum, dorsal view.

FIGURES 2 A – C. Tengella perfuga. A, B. Posterior tip of abdomen showing cribellum, ventral view; A. male; B, female; C. Metatarsus IV showing calamistrum, lateral view; male top; female bottom.