Selenops banksi Muma, 1953

Crews, Sarah C., Galvis, William & Esposito, Lauren A., 2021, Gliding canopy flatties and relatives from the Selenops banksi group (Araneae: Selenopidae), Journal of Natural History 54 (35 - 36), pp. 2343-2365 : 2348-2357

Selenops banksi Muma, 1953

( Figures 1 (a–d, 2(a–i, w), 3(a–d), 4(a–o), 5(a–c) and 6(a–b, e, h–i, l), Map 1)

Selenops banksi Muma, 1953: 38 , Figs 61–63 (♂, examined).

Selenops banksi Crews, 2011: 25 , Figs 15–16 (♂).

Selenops kikay Corronca, 1996: 69 , Figs 6 –9 (♂, examined; destroyed), syn. n.

Selenops kikay Corronca, 1998: 136 , Figs 27–30 (♀, examined; lost), syn. n.

Type material. Holotype. male, Barro Colorado, Canal Zone, Panama, 26.VII (no year), N. Banks ( MCZ 20436, examined).

Other material examined. BRAZIL: Amazonas: Manaus, Reserva Florestal Adolpho Ducke , 4 .I .1993, col . H Höfer and T Gasnier, 1♂ ( MCN) ; Bahía: Itabuna, 1♂ (type of

S. kikay, MNRJ 1875 – destroyed, examined from photos). ECUADOR: Orellana: Reserva Etnica Waorani, Transect Ent . 1 km S Onkone Gare Camp , 0.65713°S 76.453° W, 216.3 m, 24 . VI.1994, TL Erwin et al., Lot # 737, 1p ♀ ( USNM); same as previous, 2 .II .1995, Lot # 1117, 1♂ ( USNM); 4 .II .1996, Lot # 1419, 1♂ ( USNM); same as previous, 8 .II .1995, Lot # 964, 1♂ ( USNM); same as previous, 23 . VI .1996, Lot # 1607, 1♂ ( USNM); same as previous, Lot # 1619, 1♀ ( USNM); Tiputini Biodiversity Stations near Yasuni National Park , Erwin Transect T/1 Sta .1, 0.63194°S, 76.14416°W, 220–250 m, 21 .X .1998, TL Erwin et al., Lot # 1996, 1♂ ( USNM); same as previous, Lot # 1982, 1♂ ( USNM) . GUYANA: Iwokrama Forest Research Station: 1 km north of Kurupukari , canopy fog of Mora tree, 14–19 .I .1996, W Tschinkel , 1♂ ( CASENT9058003 ) . PANAMÁ: Canal Zone: Barro Colorado Island , II .2008, R Duncan, 2 imm . ( EMEC1140967 , 1140970 ); Juan Mina Station, II .1945, CD Michener, 1 imm . ( AMNH); Panama City: monsoon forest canopy fogging, 15–30 .VII .1979, E Brodhead et al., 1♂ ( USNM) . PERU: Madre de Dios: Rio Tambopata Reserve, 30 km (air) southwest of Puerto Maldonato , 12.83333°S 69.33333°W, 290 m, Smithsonian Institute Canopy Fogging Project, 12 .VII .1984, TL Erwin et al., 1♂ ( MCZ) .

Diagnosis. Selenops banksi can be separated from S. micropalpus and S. ducke by the greenish to green, lichen-like appearance in life and the yellow and white abdomen with a foliate pattern when preserved ( Figures 1 (a–d) and 2(a–i)). S. micropalpus is greyish in life and not lichen-like, and with a darker grey-brown abdomen, rather than yellow and white, when preserved ( Figures 1 (f) and 2(s–u)). S. ducke Corronca, 1996 is mahogany when preserved ( Figure 2 (k–p)) (it is unknown what the animal looks like in life). S. banksi has numerous white, flat setae on the legs both dorsally and ventrally (also present in S. curruganja sp. nov.), whereas S. micropalpus only has a few, and they are neither long nor dense ( Figures 1 (a–d, f) and 2(d–e)). While there are differences in the epigynes and endogynes of S. banksi and S micropalpus , we have many female specimens of S. micropalpus and only a single female of S. banksi , making any sort of generalisation difficult as there is some amount of variation apparent in S. micropalpus . The copulatory ducts of S. micropalpus have more turns than those of S. banksi ( Figure 6 (a–c)). And in the single female specimen of S. banksi , the lateral lobes are further apart towards the posterior, whereas they are usually very close, nearly touching, in S. micropalpus . S. banksi can be separated from S. curruganja sp. nov. by the epigynal plate, which is longer than wide in S. banksi , and about as long as wide in S. curruganja sp. nov. ( Figure 6 (a–b, d)).

Males of S. banksi can be differentiated from other males of the group by the palps: In S. banksi , the tip of the palp is densely covered in chemosensory setae, like a brush, and the tip of the cymbium is pointed ( Figures 3 (a–d), 4(a) and 5(a–c)). In S. micropalpus , the

S. ducke , holotype male, cephalothorax (MCN 25527) (n) S. ducke , holotype male, abdomen (MCN 25527) (o) S. ducke , paratype female, cephalothorax (MCN 21487) (p) S. ducke , paratype female, abdomen (MCN21487) (q) S. curruganja sp. nov., holotype female, ventral, sternum (ICN-Ar-8289) (r) S. curruganja sp. nov., holotype female, abdomen, ventral (ICN-Ar-8289) (s) S. micropalpus , holotype male, Laudat, Dominica (AMNH) (t) S. micropalpus , paratype female, Laudat, Dominica (AMNH) (u) S. micropalpus , female, Dennery, St. Lucia (CAS) (v) S. micropalpus , male, King’s Hill Forest Reserve , St. Vincent and the Grenadines (CAS) (w) S. kikay syn. nov., label from holotype ( MNRJ – destroyed) .

setae at the tip of the pedipalp are not dense, and the cymbium tip is rounded ( Figures 3 (e) and 5(f)). In S. ducke , the chemosensory setae are dense but not brushlike (at least on the single specimen available) ( Figures 3 (f–h) and 5(d)). Males can also be easily and reliably distinguished by the embolic complex, consisting of the embolus and an embolic sclerite. In S. banksi , the oval embolic base is oriented diagonally, whereas it is more vertically oriented in S. micropalpus ( Figure 5 (c,f)). Additionally, the embolus of S. micropalpus is wider than that of S. banksi ( Figure 5 (c,f)). Finally, the embolus and embolic apophysis originate from the base very near to each other and remain close distally in S. banksi ( Figure 5 (a–c)). In S. micropalpus , the embolus and embolic apophysis are separated at the base for most of their length ( Figure 5 (f)). In S. ducke , the embolus and embolic process are not separated but connected by a thin, lightly sclerotised area ( Figure 5 (d–e)).

Description. Female from ECUADOR: Orellana: Reserva Etnica Waorani, Transect Ent. 1 km S Onkone Gare Camp, 0.65713°S 76.453° W, 216.3 m, 24.VI.1994, TL Erwin et al., lot #1619 (USNM). Colour ( Figures 1 (a–d) and 2(d–e)). Carapace orange-tan with white setae, especially anterolaterally in frontal view next to the PLE ( Figure 1 (b)), some dark, thicker setae, many setae rubbed off preserved specimen; sternum yellow with orange-brown border; chelicerae orange-tan; maxillae yellow-brown, lightening distally; labium slightly darker than maxillae, lightening distally; abdomen, in life, green or with green areas, with dark and light setae; preserved, dorsally yellow-white for anterior 2/3, darker posteriorly with dark flecks, darker cardiac mark that terminates in a patch of white setae, darker laterally with a few speckles, yellowish ventrally; no markings on spinnerets; legs green with light and dark setae, preserved legs are orangish brown, slightly lighter than carapace, lighter ventrally with faint annulations, lots of long and short, flat, white setae on the ventral and dorsal or effectively prolateral and retrolateral parts of legs, particularly the femora, trochanters, and coxae, occurring in tufts on the distal parts of the legs.

Prosoma. 0.91 times longer than broad; clypeus 0.06 high.

Eyes. AER slightly recurved; PER recurved; PME and AME about the same size, PLE largest, ALE smallest; eye diameters, AME 0.26, ALE 0.18, PME 0.25, PLE 0.34, interdistances AME-PME 0.10, PME-ALE 0.21, ALE-PLE 0.31, PME-PME 1.42, ALE-ALE 1.87; ocular quadrangle AME-AME 0.79, PLE-PLE 2.32.

Sternum. 1.09 times as long as wide, posteriorly indented.

Mouthparts. Chelicerae extend anteriorly somewhat, with lots of setae, darker setae proximally and proximolaterally, lighter setae distally; maxillae longer than broad with setal tuft distally; labium truncate with rounded edges anteriorly.

Palp. Fm, spination 0-1-4; claw with 8 teeth increasing in size distally.

S. kikay syn. nov., holotype male, palpal tibia and RTA, ventral, image has been flipped horizontally (MNRJ – destroyed) (j) S. banksi , ex- S. kikay syn. nov. male, palpal tibia and RTA, ventral, image has been flipped horizontally (MCN 25526) (k) S. banksi , ex- S. kikay syn. nov. male, palpal tibia and RTA, ventral (MCN 25526) (l) S. banksi , palpal tibia and RTA, retrolateral, Orellana, Ecuador (NMNH) (m) S. banksi , palpal tibia and RTA, retrolateral, Orellana, Ecuador (NMNH) (n) S. banksi , palpal tibia and RTA, retrolateral, Peru, image has been flipped horizontally (MCZ) (o) S. banksi , palpal tibia and RTA, retrolateral, Guyana (CAS).

Epigyne ( Figure 6 (a–b, l)). Plate longer than broad, LLs closer medially than posteriorly, with EPs located medially, then LLs slightly separate again posteriorly; ML narrows abruptly anteriorly, COs at juncture between LLs and ML.

Endogyne ( Figure 6 (a–b, l)). COs mostly obscured by strongly sclerotised CDs with several turns, CDs wide, with PS located posteriorly, FDs arise from the bottom of PS, PF absent.

Opisthosoma. With terminal setal tufts, more easily seen on live specimens ( Figure 1 (a–d))

Legs. Leg formula 2314; scopulae present on ta and mt of legs I and II; ta I–IV with strong claw tufts; prolateral claws I–IV with ~11 teeth increasing in length distally; spination: Leg I, Fm d 1-1-1, pr 1-1-0, Ti v 2-2-2, Mt v 2-2; Leg II, Fm d 1-1-1, pr 0, Ti v 2-2-2 Mt v 2-2; Leg III, Fm d 1-1-1, pr 0, Ti v 1-1, Mt v 2-1; Leg IV, Fm d 1-1-1, pr 0, Ti v 1-1, Mt v 1-0.

Measurements. Total length 9.93. Carapace length 4.47. Carapace width 4.93. Sternum length 2.32. Sternum width 2.14. Abdomen length 5.46. Abdomen width 4.04. Pedipalp: Fm 1.51, Pt 0.77, Ti 0.99, Ta 1.42, total 4.70. Leg I Fm 5.41, Pt 2.09, Ti 4.59, Mt 3.80, Ta 1.67, total 17.53; Leg II Fm 6.42, Pt 1.98, Ti 4.93, Mt 3.94, Ta 1.64, total 18.81; Leg III Fm 6.03, Pt 1.77, Ti 4.53, Mt 4.01, Ta 1.72, total 18.07; Leg IV Fm 5.77, Pt 1.61, Ti 3.97, Mt 3.88, Ta 1.71, total 16.94.

Distribution. Brazil, Ecuador, Guyana, Panama, Peru (Map 1).

Life history and habitat preferences. This species has almost always been collected via canopy fogging; however, photographs and a few collections of juveniles indicate that indeed they do venture down the tree trunks and even make their eggsacs there, out in the open, 100–180 cm high ( Figure 1 (a)). As in many other selenopids, the female guards the eggsac, but rather than a smooth, white eggsac, the eggsac of S. banksi has dark flecks that look like crosshatching ( Figure 1 (a), inset). Only a single female has been collected, and this was from canopy fogging, but the species does not seem uncommon given the amount of observations made. This phenomenon of being common, yet poorly represented in collections, is typical for the family. The species tends to be found on mossy trees or trees with lichen, and the animal very much resembles lichen in both the colouration and the setal covering that provides texture. They are one of a few known green spiders, and one of the only green selenopids. Given the extent of the distribution, their ability to glide ( Yanoviak et al. 2015), and the fact that all collections are from canopy fogs, it is possible that they disperse via the canopy; thus, forests that remain intact would be most conducive to this lifestyle, and a lack of connectivity due to forest fragmentation would have conservation implications.

Variation. S. banksi , male, N = 10; Size: 6.6–8.0; S. micropalpus , female, N = 19, 6.6–13.5; S. micropalpus , male, N = 7; 6.5–10.5. The dRTA is the only variable part of the male palp based on the specimens of S. banksi available to us. The specimens from Ecuador are similar to one another, with the dRTA slightly longer and more curved ventrally than in specimens from other localities, although there is also variation among the individuals collected in Ecuador ( Figure 4 (a–o)). The dRTA of the holotype appears to differ from ex- S. kikay syn. nov. in being longer and appearing more ridged than toothed; however, we were unable to control for the position of the palp during the imaging of S. kikay syn. nov. ( Figure 4 (a–o)). The dRTA of the Guyana specimen differs from the holotype by being shorter and curving less ( Figure 4 (e, g, o)). In the specimen from Peru, the dRTA is not as long as in the holotype and is only slightly curved ( Figure 4 (e–f, n)).

Notes. We assign this female to S. banksi because it was collected with several males of the species. S. lavillai Corronca, 1996 is superficially similar to S. banksi , and the female is also undescribed. This species is lichen-like, yellow and white with a foliate pattern when preserved, and found in the canopy and on tree trunks, collected from canopy fogs simultaneously with S. banksi ; however, the male genitalia are very different from those of the banksi group. Additionally, we have specimens of the female of S. lavillai that were collected with the male of that species which will be described in a subsequent publication. Corronca (1996) described S. kikay syn. nov. based on a male and female from Itabuna, Bahía, Brazil. Although S. banksi and S. micropalpus were already described at the time ( Muma 1953), their similarity to S. kikay syn. nov. went unmentioned. The holotype of the male of S. kikay syn. nov. was examined via digital image; the female paratype could not be located. Both were indicated to be in MNRJ; however, the majority of specimens from that institution were unfortunately lost in a fire ( Kury et al. 2018). Corronca (1996) differentiates the male of S. kikay syn. nov. from others by having ‘grooves in the retrolateral tibial apophysis and a large keel ending in three teeth, and by a spatulate prolateral tibial apophysis with rounded lateral projection.’ S. banksi has a spatulate prolateral keel, as do other species of selenopids, including S. micropalpus ( Figures 3 (a,e), 4(h–k) and 5(f)). Additionally, after examination of multiple males of S. banksi , some specimens have a toothed dRTA and some do not, and some of these teeth are more pronounced in some specimens (3a, c–d, 4a–o). For the diagnosis of the female of S. kikay , syn. nov., Corronca (1996) provided no characters exclusive to this species.

The holotype of S. kikay syn. nov. was purportedly collected in Itabuna, Brazil; however, there was no other information regarding collector, year, etc. ( Figure 2 (w)). Given the distance from any other collection of S. banksi group species, it is possible that this is an erroneous locality, but the rarity of collections of these spiders also hints to its validity. Additionally, there is a hypothesised corridor between the Mata Atlântica and the Amazon that has existed since the uplift of the Andes ( Ledo and Colli 2017). According to molecular clock data ( Crews and Esposito 2020), the banksi group has been around for at least 20 my, or around the time of the beginning of the uplift of the Andes, and the group members appear to have no trouble dispersing. Thus, they have had ample time and have ample means to disperse throughout the rainforests of Central America, the Amazon, and Mata Atlântica. It is possible the Itabuna locality is on the edge of the group’s range in the Mata Atlântica. Additional efforts should be made to confirm the presence of species there.

In general, S. banksi is smaller than S. micropalpus and S. curruganja sp. nov.; this mostly applies to males of S. banksi and S. micropalpus , because males of S. curruganja sp. nov. are unknown, and there is only a single female specimen of S. banksi . Males of S. banksi range from 6.6 to 8.0, with the holotype at 7.55, the holotype of S. kikay syn. nov. at 7.2 (N = 11), and S. ducke at 6.5 ( Corronca 1996). S. micropalpus females range from 6.9 to approximately 13.45 (the specimen is damaged) (N = 18). The holotype and the second-largest female are both from Dominica. Females from St. Lucia, Martinique, and St. Vincent and the Grenadines tend to be smaller. The males of S. micropalpus are more similar in size to males of S. banksi , ranging from 6.65 to 10.61 (N = 7). The female of S. curruganja sp. nov. is large (12.2) compared to the single female specimens of S. banksi , S. ducke (9.0), and ex- S. kikay syn. nov. (7.55 – ( Corronca 1996)), and many of the S. micropalpus females.

Figures

Figure 1. (a) Selenops banksi, female, guarding eggsac, Costa Rica; inset shows crosshatching pattern on eggsac (photos by Gernot Kunz) (b) S. banksi, Costa Rica (photo by Gernot Kunz). (c) S. banksi, eating a queen ant of Camponotus sp., Reserva Nacional Allpahuayo-Mishana, Loreto, Peru (photo by Projeto Mantis – https://www.projetomantis.com/) (d) S. banksi, on a tree, Iwokrama Rainforest Reserve, Guyana (photo by Paul Bertner) (e) S. curruganja sp. nov., on mossy guava tree, Lloró, Chocó, Colombia (photo by Sarah Crews) (f) S. micropalpus, on Bursera simaruba, St. Lucia (photo by Sarah Crews).

Figure 2. (a) Selenops banksi, paratype male, from Panama (MCZ) (b) S. banksi, holotype male, from Panama (MCZ 20436) (c) S. banksi, ex-S. kikay syn. nov., holotype male, from Itabuna, Brazil (MNRJ – destroyed) (d) S. banksi, female described here, from Orellana, Ecuador (NMNH) (e) S. banksi, female, ventral, Orellana, Ecuador (NMNH) (f) S. banksi, male, Orellana, Ecuador (NMNH) (g) S. banksi, male, ventral, Orellana, Ecuador (different specimen than f but in same vial) (NMNH) (h) S. banksi, male, Rio Tambopata Reserve, Madre de Dios, Peru (MCZ) (i) S. banksi, male, Iwokrama,Guyana (CAS 9058003) (j) S. curruganja sp. nov., holotype female, Corpoica, Tumaco, Dept. Nariño, Colombia, (ICN-Ar-8289) (k) S. ducke, holotype male, sternum (MCN 25527) (l) S. ducke, paratype female, sternum (MCN 21487) (m)

Figure 3. (a) Selenops banksi, palp, ventral and retrolateral, highlighting various structures present in species belonging to the banksi group; dark pink – dRTA, teal – vRTA, yellow – MA, purple – C, pink – E, orange – EB, green – EA. (b) S. banksi, holotype male, palp, ventral and retrolateral (MCZ 20436) (c) S. banksi, ex-S. kikay syn. nov., palp, ventral, from Manaus, Brazil; image has been flipped horizontally (MCN 25526) (d) S. banksi, ex-S. kikay syn. nov., holotype male, from Itabuna, Brazil (MNRJ – destroyed) (e) S. micropalpus, holotype male, palp, ventral and retrolateral (AMNH) (f) S. ducke, holotype male, palp, ventral, image has been flipped horizontally (MCN 25527) (g) S. ducke, holotype male,palp, ventral, image has been flipped horizontally (MCN 25527) (h) S. ducke, holotype male, palp, retrolateral (MCN 25527).

Figure 4. (a) Selenops banksi, ex-S. kikay syn. nov., holotype male, palp, retrolateral view, image has been flipped horizontally (MNRJ – destroyed) (b) Selenops banksi, ex-S. kikay syn. nov. holotype male, palpal tibia and RTA retrolateral view, image has been flipped horizontally (MNRJ – destroyed) (c) S. banksi, ex-S. kikay syn. nov. holotype male, palpal tibia and RTA, retrolateral view, left palp (MNRJ – destroyed) (d) S. banksi, palpal tibia and RTA, ventral, Orellana, Ecuador (NMNH) (e) S. banksi, holotype male, palpal tibia and RTA, ventral, Panama (MCZ 20436) (f) S. banksi, palpal tibia and RTA, ventral, Peru, image has been flipped horizontally (MCZ) (g) S. banksi, palpal tibia and RTA, ventral, Guyana (CAS) (h) S. banksi, ex-S. kikay syn. nov., holotype male, palpal tibia and RTA, ventral (MNRJ – destroyed) (i) S. banksi, ex-

Figure 5. (a) Selenops banksi, holotype male, palp, lateral, showing embolus (E), embolar sclerite (ES), and embolar base (EB) (MCZ 20436) (b) S.banksi, palp,lateral,Orellana,Ecuador (NMNH) (c) S.banksi, ex-S.kikay syn. nov., holotype male,palp,lateral,Itabuna,Brazil (note that this palp is in a slightly different orientation than preceding) (MNRJ – destroyed) (d) S.ducke, holotype male,palp,lateral,E and ES connected by weakly sclerotised area, flipped horizontally, Manaus, Brazil (MCN 25527) (e) S. ducke, holotype male, palp, lateral, E and ES connected by weakly sclerotised area, Manaus, Brazil (MCN 25527) (f) S. micropalpus, palp, lateral, E and ES are further apart than in S. banksi (A–C), King’s Hill Forest Reserve,St. Vincent and the Grenadines (CAS).

Figure 6. (a) Selenops banksi, epigyne and endogyne, highlighting various structures, epigyne: green – LLs, yellow – ML, violet – COs, pink – EPs, endogyne: purple – CDs, pink – PS, green – FD (b) S. banksi, epigyne and endogyne,Orellana,Ecuador (NMNH) (c) S. micropalpus, epigyne and endogyne, Dennery, St. Lucia (CAS) (d) S. curruganja sp. nov., epigyne and endogyne, Corpoica, Tumaco, Dept. Nariño, Colombia, (ICN-Ar-8289) (e) S. banksi, male, spinnerets, ventral, Orellana, Ecuador (NMNH) (f) S. ducke, paratype female, epigyne, Manaus, Brazil (MCN 21487) (g) S. ducke, paratype female, endogyne, Manaus, Brazil (MCN 21487) (h) S. banksi, female, spinnerets, ventral, Orellana, Ecuador (NMNH) (i) S. banksi, female, spinnerets, caudal, Orellana, Ecuador (NMNH) (j) S. micropalpus, female, spinnerets, caudal, Dennery, St. Lucia (CAS) (k) S. micropalpus, genitalia, caudal, Dennery, St. Lucia (CAS) (l) S. banksi, female, genitalia, caudal, Orellana, Ecuador (NMNH).