Gelanor Thorell, 1869

Ligia R. Benavides & Gustavo Hormiga, 2016, Taxonomic revision of the Neotropical pirate spiders of the genus Gelanor Thorell, 1869 (Araneae, Mimetidae) with the description of five new species, Zootaxa 4064 (1), pp. 1-72 : 9-14

Gelanor Thorell, 1869

Type species by original designation: Galena zonata C.L. Koch, 1845

Galena C.L. Koch, 1845 . Type species by original designation: Galena zonata C.L. Koch, 1845

N.B. Thorell (1869:37) erroneously argued that the name Galena was preoccupied and proposed the replacement name Gelanor . Pickard-Cambridge (1905) clarified that Galene De Haan, 1833 (a crustacean of family Goneplacidae ), and not Galena , is the name that Thorell erroneously took as preoccupied and therefore the valid genus name should be Galena because Gelanor was a junior synonym. We believe that the prevailing use of the genus name Gelanor should be maintained because the use of the older synonym, barely used since 1905, would cause confusion. Because of the use of the genus-name Galena by Pickard-Cambridge (1905) we cannot apply the automatic Reversal of Precedence (Article 23.9 of the ICZN), as the condition described in Article 23.9.1.1 is not met. We will refer the matter to the Commission for a ruling under its plenary power [Art. 81]. While the case is under consideration we will maintain the use of the junior name Gelanor as recommended by the Code.

Diagnosis. Gelanor species can be easily differentiated from other mimetids by the presence of two white to light yellow circular or oval spots with dark red-brown margins (absent in some species) located on the anterior part of the abdomen. Spot size varies from small (e.g., G. consequus ) to large (e.g., G. latus ), margins from thin (e.g., G. s i q u i r re s) to broad (e.g., G. l a t u s). With six to ten horizontal transversal lines, posterior to abdominal spots, colored as spot margins. In live or recently collected specimens, a whitish band can be seen between each pair of dark red- black lines. Colors in live specimens more vivid than those of ethanol specimens (in the latter the spots and the black and white lines tend to vanish). Male palpal length (involving elongation of femur, patella and tibia) usually twice the length of the body ( Fig. 4 a) (unique to Gelanor among mimetids). In addition, Gelanor is the only mimetid genus with laterigrade legs ( Fig. 1 ). Thoracic fovea very conspicuous and semicircular ( Figs. 8 I, 27K, 46L, 49K). Males lack the characteristic prolateral row of long and short macrosetae of tibia I and II present in females; Gelanor lacks the small spines on the retrolateral proximal side of femur I and prolateral proximal side of femur II characteristic of Australomimetus and Mimetus .

Description. Small to medium size spiders; total length 3.54–5.38 in males and 3.86–7.54 in females. Carapace light yellow to dark brown-orange of a uniform color or with some areas darker than others; 1.99–3.15 long in males, 1.71–4.09 long in females; 1.45–2.53 wide in males and 1.52–2.66 wide in females. Fovea well marked, rhomboidal ( Figs. 8 I, 27K, 46L, 49K). Eyes in two rows with the anterior row nearly straight and the posterior procurved ( Fig. 3 A, E), AME largest and placed on a small tubercle. ALE, PLE and PME similar in diameter. Lateral eyes juxtaposed ( Figs. 3 A, E). Clypeus low, 0.05–0.1. Chelicerae nearly straight, fused at the base ( Fig. 3 E), larger in females than in males; 0.6–1.07 long and 0.34–0.54 wide in males; 0.83–1.36 long and 0.41– 0.62 wide in females; pale yellow to dark orange. Cheliceral promargin with ca.11 peg teeth ( Fig. 3 E, G). Cheliceral retromargin with a single protuberance (mound) and close to it, right under the fang, a conglomerate of ca. nine cuticular pores placed on a low mound ( Fig. 3 H). Two macrosetae crossing each other near to the distal end on their internal cheliceral margin ( Fig. 3 E); an additional setae located towards the middle on the anterior part of each paturon. Sternum longer than wide and prolonged between coxae IV ( Fig. 3 B), pale or with dark patterns. Endites darker than the sternum on its proximal end, lighter towards its distal end; reniform, with the distal end wider than the proximal. Labium wider than long, with a rounded distal edge and light yellow to dark brown. Abdomen cylindrical or rhomboidal, typically larger in females, whitish or darker, usually with two large dorsal white spots, and in some species with a dark red/brown margin; a series of transversal white and dark red lines posterior to the two large spots can be seen in most of the species. Legs laterigrade, yellowish with I and II darker and markedly longer than III and IV; leg formula: I-II-IV-III. Prolateral and retrolateral area of femora I and II with dark brown or distal black spots; tibiae I and II with distal dark areas ( Fig. 3 D). ALS and PLS with serrated macrosetae on its border, surrounding the spigots ( Figs. 10 A, C); these macrosetae are present also on the PMS but are less abundant ( Fig. 10 B). Colulus rhomboidal ( Figs. 21 F, 40A). Tracheal system composed by four cylindrical tracheal trunks with the medium tracheal trunks shorter and their apical end leaf-shaped ( Figs. 40 G–H).

Male

Leg spination. All femora with dorsal macrosetae arranged in two parallel alternating lines and three macrosetae very close to the distal articulation point ( Fig. 3 D). Number of dorsal macrosetae on femora decreases from femur I to femur IV. (Femur I has ca. 21, femur II ca. 16, femur II ca. 10 and femur IV ca. 6). Patellae I and II with a distal retrolateral macroseta. Tibiae I and II with three prolateral, three dorsal and one retrolateral macrosetae ( Fig. 6 H). Metatarsi I and II with dorsal macrosetae linearly arranged, sub-equal in size ( Fig. 6 I), absent in females. Metatarsus I with four long prolateral macrosetae (Prol-mt-I); metatarsus II with three prolateral macrosetae (Prolmt-II); these long macrosetae are intercalated by a series of shorter macrosetae (7–10 short macrosetae in between two long macrosetae). Each series of shorter macrosetae gradually increase in length from the proximal to the distal end. Although the number of long prolateral macrosetae in metatarsi I and II is the same across all Gelanor species, the number of short macrosetae in between these long macrosetae varies among specimens of the same species and even in the same individual this number may differ between left and right leg.

Spinnerets and epiandrous fusules. ALS ( Fig. 18 F) with a MAP, a nubbin, more than 40 piriform gland spigots (Pi) and two very distinctive spigots that are of a different shape than the other Pi (modified Pi spigots (MoPi, Figs. 18 F–G) sensu Townley et al. (2013)). These modified spigots are conical, wider and with larger openings than the Pi spigots. PMS with four aciniform gland spigots (Ac), one minor ampullate gland spigot (mAP), a tartipore and a nubbin ( Fig. 18 H). PLS with ca. 20 Ac ( Fig. 18 I). Epiandrous fusules distributed in a transverse line and leveled with the surrounding cuticle. The fusules are inside pits, each one of them containing between one and four fusules. The posterior edge of the epiandrous area appears swollen when compared with the surrounding cuticle.

Palp. Palp length between 7.8 and 10.73 mm and 1.86 to 2.41 times the body length. The tarsus is rotated, therefore the cymbium is prolateral and the bulb is retrolateral in a forward-extended pedipalp. All segments of the palp are similar in width, with the distal end of the tibia wider. Macrosetae present on the tibia as follows: dorsal side with three long (ca. 1 mm) macrosetae, placed in a row, equidistant one from the other and towards its distal end ( Fig. 4 A); ca. 14 short setae on the tibia (approximately half the length of the bulb) surrounding the palpal bulb and two long setae pointing backwards. Promarginal and retromarginal edges of the cymbium are sclerotized with the retromarginal edge much more sclerotized and darker ( Fig. 4 C). Macrosetae present on the cymbium with higher density on the apex and the lower half ( Fig. 4 E). Cymbium bearing a flap along its retrolateral margin ( Figs. 4 C, E); this flap varies in shape across species and therefore is useful for species identification. Paracymbium integral (continuous with the cymbium and not attached to it by means of a membrane), with a concave shape, darker than the cymbium; the paracymbium can have one of its margins attached to one of the edges of the cymbial flap, forming and “L” shaped with the cymbial retromargin (most Gelanor species, Fig. 4 C) or be more conspicuous, having a triangular concave shape and be detached and shifted towards the apex ( G. altithorax and G. fortuna ; Figs. 7 C and 23G respectively); tegulum sclerotized, ring-like shaped. Conductor connected to tegulum by means of a membrane ( Figs. 5 B, F), its apex sclerotized and darker than its base ( Figs. 5 A, E). Median apophysis absent. Embolus conical and connected to tegulum by means of a membrane, heavily sclerotized and black ( Figs. 5 A, E). Cymbial tarsal organ base diameter smaller than base of nearby setae ( Figs. 17 G, 28F).

Female

Leg spination. Femora with one to three scattered dorsal macrosetae subequal in size. Tibia I with seven long prolateral macrosetae; tibia II with five long prolateral macrosetae; metatarsus I with four prolateral macrosetae with and metatarsus II with three prolateral macrosetae. As in males, the long macrosetae are intercalated by a series of shorter macrosetae increasingly longer from the proximal to the distal end. The series of long and short prolateral macrosetae of tibiae I and II are absent in males. Number of short macrosetae on each series between two long macrosetae varies across specimens of the same species and in some cases between left and right legs of the same specimen.

Spinnerets. ALS with a MAP, nubbin and ca. 90 Pi ( Fig. 10 A) PMS ( Fig. 10 B) with 4–5 Ac, an anterior Cyl, posterior mAP and nubbin. PLS with ca. 19 Ac and two Cyl (one anterior and one posterior). Cyl with long and wide base, almost equal in size to the shaft. Cyl shaft grooved, conically shaped, its base wider than its distal end; the diameter of the Cyl opening is almost as the total diameter of its apex. (e.g. Figs. 10 B, 30O). PMS Cyl similar in morphology to PLS Cyl. Aggregate and flagelliform silk gland spigots absent.

Epigynum . Epigynum with a longitudinal septum that varies in shape and width across species and separates two pit-like copulatory openings that are ventrally oriented ( Fig. 4 F). The septum is not as evident in G. consequus as in the other Gelanor species. Anterior lateral lobes on each side of the septum, darker and covering the copulatory openings (absent in G. consequus ). Copulatory openings ventrally oriented and circular shaped. Spermathecae ( Fig. 4 G) spherical ( G. altithorax , G. f o r t u n a, G. innominatus , G. latus , G. siquirres , G. waorani , G zonatus ), oval ( G. juruti ) or in the shape of an interrogation sign ( G. consequus ). Spermathecae can be externally fused (G. i n n o m i n a t u s, G. j u r u t i, G. l a t u s) or separated less than one spermathecae diameter ( G. altithorax , G. consequus , G. fortuna , G. siquirres , G. waorani , G. zonatus ). Spermathecal walls sclerotized; accessory glands clustered with its base diameter between one and two times less than the duct diameter. Copulatory ducts originate anteriorly, are less than a half of the spermathecae in length, rigid and sclerotized. Fertilization ducts originate posteriorly, are less than half the spermathecae in length and are slightly curved. Copulatory plugs were observed in the copulatory openings of G. latus and G. zonatus .

Natural history. Unfortunately, very little is known on the natural history of Gelanor . Gelanor species are found between sea level and 1,600 m, primarily in tropical rain forests, or cloud forests. Most of the Gelanor specimens collected and photographed by us were found in the understory, sitting on a few silk lines on the reverse side of leaves of plants up to one meter with large leaves (e.g., Arecaceae ), or on shrubs up to two meters, although based on the material examined, Gelanor species can be found also in the canopy. Gelanor species have a crab-like laterigrade posture, in which the prolateral side of the legs I and II face up.

Distribution. Northeast Mexico to southern Uruguay ( Fig. 2 ). Gelanor species are found exclusively in the Neotropics. No Gelanor species are found in southern South America or in the Nearctic region, most probably due to ecological barriers ( Sanmartín and Ronquist 2004). Although most of the species of Gelanor seem to have a restricted distribution rage (which might be an artifact due to insufficient sampling), three species, are widely distributed ( G. consequus , G. latus and G. zonatus ; Figs 22 , 42 and 57 respectively). Based on the specimens that we have studied, G. zonatus is the only Gelanor found on islands, with a record from Port of Spain ( Trinidad and Tobago).

Phylogenetics. The monophyly of Gelanor is supported by the following putative synapomorphies: laterigrade legs; fovea well marked, semicircular and transversal; abdomen with two white to light yellow round or oval spots with dark red–brown margins, located on the anterior part of the abdomen, which is wider anteriorly; male palp approximately twice the total body length; embolus conical, dark with a thick base; cymbium bearing a flap that is less sclerotized than the rest of the cymbium; epigynum with a medial septum and rounded genital openings.

Composition. The genus Gelanor groups ten species: Gelanor fortuna new species, Gelanor juruti new species, Gelanor moyobamba new species, Gelanor siquirres new species, Gelanor waorani new species, Gelanor altithorax Keyserling, 1893 (= Gelanor lanei Soares, 1941 new synonymy), Gelanor consequus O. P.- Cambridge, 1902 (= Gelanor depressus Chickering, 1956 new synonymy, Gelanor gertschi Chickering, 1947 new synonymy and Gelanor heraldicus Petrunkevitch, 1925 new synonymy), Gelanor innominatus Chamberlin, 1916 , Gelanor latus ( Keyserling, 1881) (= Gelanor mixtus O. P.- Cambridge, 1899 new synonymy, Gelanor mabelae Chickering, 1947 new synonymy, Gelanor ornatus Schenkel, 1953 new synonymy and Gelanor proximus Mello- Leitão, 1929 new synonymy) and Gelanor zonatus ( C.L. Koch, 1845) (= Gelanor distinctus O-P. Cambridge, 1899 new synonymy, Gelanor insularis Mello-Leitão, 1929 new synonymy and Gelanor obscurus Mello-Leitão, 1929 new synonymy).

Figures

FIGURE 1. Gelanor latus. A, Male. B, Female. C, Male and female. All from Caicubí, Rio Jufarí, Médio Rio Negro, (Brazil) (Photos, L. Benavides).

FIGURE 2. Distribution records of Gelanor species.

FIGURE 3. Standard measurements used in this study. Asterisks correspond to peg teeth.

FIGURE 4. Palp and epigynum showing the morphological terms used.

FIGURE 5. Schematic drawings of Gelanor male palps (drawn after right palps; reversed to show as left palps), G. latus (A – D); G. consequus, partially expanded with KOH (E – G). A, Ectoventral view; arrow points to alveolar cavity. B, Tegular division, proximal (basal) view; arrow points to partially collapsed membranous sac (due to methyl salicylate treatment). C, Proximal (basal) view; arrow points to embolus base-tegulum sclerotized hinge. D, Detail of embolus base-tegulum connection (right pointing arrow, membranous connection; left pointing arrow, sclerotized connection). E, Ectoventral view; bottom pointing arrow, embolus; up pointing arrow, conductor (part). F, Mesal view arrow, embolus base. G, Ectal view; arrow, embolus.

FIGURE 6. Gelanor altithorax, Male (A – I, LB 308); A – C, Habitus; D – E, Cephalothorax; F – I, right legs. A, Dorsal. B, Lateral. C, Ventral. D, Frontal view. E, Lateral view. F, Prolateral. G, Retrolateral. H, Tibiae I and II, dorsal. I, Metatarsi I and II, dorsal. Scale bars, A – C, 1 mm; D – E, 500 µm; F – I, 2 mm.

FIGURE 7. Gelanor altithorax, male left palp (LB 308). A, Prolateral. B, Dorsal. C, Retrolateral. D, Ventral. Scale bars, 500 µm.

FIGURE 8. Gelanor altithorax, female (A – G, LB 177; H – L, LB 317); A – C, Cephalothorax; D – F, abdomen. A, Dorsal. B, Lateral. C, Ventral. D, Dorsal. E, Lateral. F, Ventral. G, Cephalothorax, Frontal. H, Carapace, Dorsal. I, Fovea, detail. J, Legs, prolateral. K, Detail of tibiae I and II, Prolateral view. L, Detail of metatarsus I and II, prolateral view. Scale bars, A – C, G, K, L, 1 mm; D – F, J, 2 mm; H, 200 µm; I, 20 µm.

FIGURE 10. Gelanor altithorax, female spinnerets (LB 317). A, left ALS. B, left PMS. C, left PLS. D, detail of Cy on PLS. Scale bars, A – C, G, 200 µm; F, H, I, L, 100 µm; E, A, C: 10 µm; B, 3 µm; D, 5 µm.

FIGURE 17. Gelanor consequus, male (LB 41), left palp, SEM. A, Dorsal. B, Apical. C, Ventral. D, Prolateral. E, Retrolateral view expanded. F, Apical, conductor detail. G, Tarsal organ. Scale bars, A – E, 100 µm; F, 30 µm; G, 10 µm.

FIGURE 18. Gelanor consequus. Male (LB 41). A, Fovea. B, Fang and peg teeth. C, Epiandrous spigots. D, Detail of epiandrous spigots. E, Spinnerets, general view. F, left ALS. G, ALS detail showing the MoPi (*). H, left PMS. I, left PLS.

FIGURE 21. Gelanor consequus, female (A – B, F, LB 49; C – E, G – H, LB 94); A – B, Chelicerae; C – H, Spinnerets and colulus. A – B, Cheliceral glands and tooth. C, Spinnerets, general view. D, Right ALS. E, Detail showing modified setae on ALS margin. F, Colulus. G, Right PLS. H, Left PMS. Scale bars, A, 20 µm; B, D – H, 10 µm; C, 100 µm.

FIGURE 22. Distribution map of records of Gelanor consequus and its junior synonyms. Red shapes represent type localities. Blue circles represent new records for the species. Red star, G. consequus; red hexagon, G. depressus (new synonymy); red square, G. gertschi (new synonymy); red triangle, G. heraldicus (new synonymy).

FIGURE 40. Gelanor latus, female (A – C, LB 88; D – FE, LB 27; G – H, LB 125); A – F, Genitalia; G – H, Tracheal system. A, Anterior. B, Ventral. C, Caudal. D – E, Spermathecae. D, Anterior. E, Ventral. F, Caudal. G, Tracheal system. H, Detail of the medium tracheal trunks showing their leaf – shaped tip. Scale bars, A – C, 200 µm; D – F, 100 µm.

FIGURE 42. Distribution records of G. latus and its junior synonyms. Red shapes represents type localities and blue circles other localities for the species. Red circle, G. latus; red star, G. mabelae (new synonymy); red square, G. mixtus (new synonymy); red triangle, G. ornatus (new synonymy); red hexagon, G. proximus (new synonymy).

FIGURE 57. Distribution records of Gelanor zonatus and its junior synonyms. Red shapes represent type localities and blue circles other localities for the species. Red star, G. distinctus (new synonymy); red square, G. insularis (new synonymy); red triangle, G. obscurus (new synonymy).