Marma Simon 1902
Creators: name: Salgado, Alexandre
name: Ruiz, Gustavo R. S.
Publisher: name: Zenodo
Year: 2020
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Araneae
Salticidae
Marma

description: Genus Marma Simon, 1902
Marma Simon, 1902: 376 (type species: Marma baeri Simon, 1902).
Thysema Mello-Leit „o, 1944: 390 (type species: Thysema dorae Mello-Leitão, 1944); synonymized by Galiano (1962: 36).
Paralophostica Soares & Camargo, 1948: 396 (type species: Paralophostica centralis Soares & Camargo, 1948); synonymized by Galiano (1962: 36).
Revised diagnosis. The Amphidraus-Marma clade (which also includes genera Nebridia and Yacuitella) is composed of small to medium-sized jumping spiders. The chelicerae in these genera have two promarginal teeth and one fissident retromarginal tooth with two or more cusps (Figs 51A, C; Galiano 1999: figs 6–7; Zhang & Maddison 2015: fig. 218; Salgado & Ruiz 2019: figs 28A–C). Furthermore, the male palp has an embolic disc with projections (Figs 5C, 7A, 9 A–C, 16A, 23A–C, 28A; Galiano 1999: figs 8–9, 15; Salgado & Ruiz 2019: fig. 3A), and the epigynal plate is sclerotized and lacks the typical euophryine window structure (a pair of rounded and less sclerotized regions that allow the visualization of the spermathecae; Figs 7G, 15F, 22F, 27F, 32F, 37H, 46C, 50F, 55F; see Zhang & Maddison 2015: figs 39, 46, 53, 60).
Males of Marma can be distinguished from related genera by having a sclerotized cymbial conductor (cc; Figs 36C, 40A), whereas it is membranous in the other members of the Amphidraus - Marma clade (see Salgado & Ruiz 2019: fig. 2E); by having part of embolus shaft hidden under the tegulum (Figs 7A, F, 9A, C, 15A, E), whereas it is typically exposed in the tribe (see Salgado & Ruiz 2019: figs 5A, 7C); and by not having the tegular lobe (Figs 13C, 15A, 25C, 27A), which is developed in Amphidraus and Nebridia (see Galiano 1963: Lam. XXVII, fig. 16 and Salgado & Ruiz 2019: fig. 3A) and slightly differentiated from the tegulum in Yacuitella (see Galiano 1999: fig. 9). In addition, males of Marma and Yacuitella have an embolus composed of a shaft alone (Figs 20C, 22A, 23A, 53C, 55A, 56C; see Galiano 1999: figs 8–9, 15), whereas in Amphidraus and Nebridia the embolus opening is extended, forming a tubular piece that is less sclerotized than the shaft (see Salgado & Ruiz 2019: figs 3A, 7C). Males of Marma can be distinguished from those of Yacuitella by having a single distal process on the embolic disc (Figs 23 A–C, 30C, 32A, 33A, D) and a single apophysis on the male palpal tibia (Figs 20D, 22B, 23 D–E, 30D, 32B, 33E) (Yacuitella has two processes on the embolic disc and three tibial apophyses; see Galiano 1999: figs 9–10). Females of Marma differ from those of Amphidraus by not having a coupling pocket in the epigynal plate (present in some species of Amphidraus; see Salgado & Ruiz 2019: fig. 27E). Also, the proximal copulatory duct is longer in Marma, which places the secondary spermatheca farther from the copulatory opening (Figs 6C, 7H, 10B, 18 C–D, 19A–B), whereas the secondary spermatheca is adjacent to the copulatory opening in Amphidraus and Yacuitella (see Salgado & Ruiz 2019: figs 17D, 25D; Galiano 1999: figs 12–13). The female of Nebridia remains unknown.
Description. Small to medium-sized (2.9–4.8 mm); standard jumping spider body (without striking modifications such as mimicry to ants, beetles, etc. or flattening of carapace; Figs 1, 2, 3, 5 A–B, 6A–B).
Carapace: anterior region of carapace (near to anterior eyes) slightly narrower than widest section (Figs 8D, 13A); PME closer to PLE than to ALE (Fig. 8D); PME slightly more dislocated to lateral side of carapace than PLE (Fig. 8D); anterior eye row in frontal view wider than anterior section of carapace (Figs 3D, 47F); fovea placed slightly beyond PLE (Figs 6A, 20A, 21A); in lateral view the posterior third with abrupt slope (Figs 47B, E). Coloration: cephalic area black and thoracic area brown in both sexes (Figs 5A, 6A, 30A, 31A). Scale pattern: with blend of white and brown scales (Figs 2 A–D, 8D–E). Brown scales (Fig. 8E) are longer and slenderer than white scales (about the same width throughout its length) and bear lateral barbs. White scales (Figs 8A, E) are flattened and wide (at least twice the width of its base) and bear central ridge along their dorsal length; shallow, oblique grooves extending towards borders, with single lateral row of flattened barbs; venter of scales apparently also with barbs. White and brown scales are connected to the unmodified base (Figs 8A, C) and rest parallel to the surface. Cephalic region also with rigid, black protective bristles connected to round, well-delimited bases (Figs 8 A–B, F). Two different patterns of distribution of scales on carapace can be observed (best observed in live specimens): (1) presence of a distinct triangle of scales pointing backward, with scarce scales on sides (Figs 34 A–B, D–E, 47A–B, D–E); (2) homogeneous distribution of scales on carapace (Figs 1E, 29 A–F, 30A, 31A); anterior lateral eyes (ALE) and anterior median eyes (AME) surrounded by brown scales (Figs 1A, D, 3B, D, 12C, F, 34C, F, 47C, F).
Abdomen: dorsally variegated, with brown and white scales (Figs 5A, 6A); ventrally, it can be light brown with three longitudinal, narrow dark brown stripes (Figs 5B, 6B), or with pale border and thick dark brown longitudinal stripe (Fig. 30B), or without stripes (Figs 21D, 26B). Anal tubercle covered with white scales (yellow arrow in Figs 11A, G), like the ones present on carapace. Epiandrous region with a pair of conspicuous tufts of spigots, bearing about six long spigots each (Figs 11 H–I).
Chelicerae: with two promarginal teeth and one fissident retromarginal tooth with two or more cusps (Figs 51A, C; see Zhang & Maddison 2015: fig. 218); ventral keel with long (sensitive?) setae (Figs 51 A–B); frontal surface with sparse long and short setae (Figs 51 D–E), and long, barbed setae between teeth and fang (Fig. 51F).
Legs: in general, are pale with dark marks (small spots, or stripes that cover entire surface of an article, or rings that circulate an article; Figs 1A, 3C, 12A); femur I color pattern can vary within a species, being yellowish with dark marks, totally yellowish or totally dark in different males (compare Figs 12 A–C in M. linae sp. nov. and Figs 47A, 48E in M. rosea); color pattern for metatarsi is standard for all species and both sexes: metatarsi I–II with dark proximal ring, III–IV with proximal and distal dark ring (Figs 30B, 31B); all males with black tarsus on leg I (Figs 3A, 12 A–B, 29A, 34A–B).
Spinnerets (Figs 11 A–F; examined only in M. baeri male): spigots, in general, are reduced in number in all spinnerets. In the ALS there are only four piriform spigots forming an arch anteriorly (Figs 11 B–C); ALS also has a single major ampullate spigot (MAP) placed posteriorly, hidden in Fig. 11B and partially seen in Fig. 11C (females are expected to have two MAP in ALS). In the PMS there is an unusually elongate minor ampullate spigot (mAP) right behind a large tartipore, and possibly a single lateral aciniform spigot, which is broken off in both PMS photographed (Fig. 11F). In the PLS, among the tartipores, there are four unusually elongate aciniform spigots forming an arch facing mesally (Figs 11 D–E).
Male palp: in general, brown with some dark spots (Figs 30 C–D); short trochanter (Figs 30D, 37B); femur without modifications (Figs 32B, 37B); short patella without modifications (Fig. 7C) and generally with white scales concentrated dorsally (orange scales in M. wesolowskae sp. nov.; Figs 30D, 33E, G–H); tibia with the same length as the patella and with a single apophysis placed retrolaterally (RTA) (Figs 5D, 7C, 13D, 15B, 20D, 22B, 25D, 27B, 30D, 32B); dorsally with a smooth single dark scale (without shallow, oblique grooves; Figs 33E, I); the retrolateral tibial apophysis (RTA) is generally finger-shaped with dark tip (Figs 36D, 44D, 48D, 53D); tip of RTA with parallel ridges (Figs 23 D–E, 28D, 33F, 52E); cymbium with tip narrower than the proximal region (Figs 7E, 15D, 22D, 27D, 32D, 37D, 50D, 55D); sometimes with apical dark spot (Figs 30D, 37B); with a small proximal retroventral projection (possibly part of paracymbium; indicated by arrows in Figs 9F, 16D, 28B, 40B); with sclerotized cymbial conductor (cc, a modification of cymbial groove) placed on retrolateral ventrodistal portion (Figs 5C, 9E, 13C, 20C, 25C, 30C, 40A); cymbial conductor with a groove on which the embolus tip rests (Figs 9E, 16C, 28B, 33B, 40A; white arrow in Fig. 56D); subtegulum with a prolateral lobe (Figs 23B, 56 A–B); tegulum without proximal lobe and with large retrolateral “shoulder” (Figs 9A, 23A, 33A, C); tegular shoulder with a groove on which the embolus shaft rests (black arrow in Fig. 33C); typical euophryine retrolateral sperm duct loop is absent (Figs 7F, 15E, 22E, 27E, 32E, 37G, 50E, 55E); embolic disc with a pointed process (PED) projected to apical portion of the palp (Figs 5C, 9 A–B, 33D, 40C, 52A); embolic disc with a dorsal groove (indicated by black arrow in Figs 9C, 16A, 23C, 28A, 33D, 40C, 52A, 56C); base of embolus shaft with a projection of granulated appearance (Figs 16A, 28A, 33D, 40C, 52 A–B, 56C); part of embolus shaft hidden under the tegulum (Figs 7A, F, 9A, C); embolus shaft with small spikes, sometimes forming a spiral row (Figs 9D, 16B, 23C, 28C, 33D, 40D, 52C, 56C). Female with unmodified palp (Fig. 3D).
Expansion of palp and copulation (Figs 38–39): the male grabs the female’s abdomen with his leg I before the expansion of the palp (Figs 39 A–D); the bulb rotates to the retrolateral side when the basal hematodocha is filled with hemolymph (Figs 38 A–B). Later, the distal hematodocha is filled and the embolic disc rotates to the prolateral side (Figs 38 A–B, D–E). The subtegulum and the tegulum remain coupled during the expansion (Figs 38 D–E).
Epigyne: epigynal plate generally with two distinct copulatory openings (Figs 6C, 7 G–H, 10A, 17A, 41A) (M. sinuosa sp. nov. has the copulatory openings fused in a single opening; Figs 21E, 22 F–G, 24A, C); copulatory ducts with glands on their initial portion (near to copulatory openings), independent from those of secondary spermathecae (Figs 10D, 43 C–D); copulatory ducts length ranges from short (e.g. M. linae sp. nov.; Figs 14 C–D) to very long (e.g. M. argentina; Figs 45C, F); internal tegument of copulatory ducts is smooth (Fig. 42D); the secondary spermathecae are reduced and generally placed far from the copulatory openings (Figs 10B, 17 B–D, 19A–B, 24B, F, 28E–F, 33J, 41B, 43A–B); primary spermathecae with homogeneous diameter in some species (Figs 14 C–D, 31C–D); in other cases, with their initial portion dilated, tapering towards the fertilization ducts (Figs 21 E–F, 24D, 26C–D); internally, spermathecae have spikes on the tegument (ets; Figs 41 C–D, 42A–B); Bennett’s glands (BG) externally appear as a circular hole (Figs 10C, 19 C–D, 24E); internally they are projected into the middle of the spermathecae and have a spiky appearance (spikes probably bear openings of individual glands; Figs 41C, 42 A–C); fertilization ducts are laterally projected (Figs 10C, 14C, 15G).
Remarks about crypsis: Members of Marma are specialized in hunting on the ground, rock surfaces, urban constructions and tree trunks. They are not found inhabiting intertwined tree branches or leaves. The preference for open environments as those mentioned above is related to the cryptic coloration in shades of brown that provide them efficient camouflage with the substrate (Figs 1, 2, 3, 12, 29, 34, 39, 47). Also, no bright colors are observed in males (Figs 2 A–B, 3A–B, 12A–C, 29A–C, 34A–C, 47A–C).
description: Published as part of Salgado, Alexandre & Ruiz, Gustavo R. S., 2020, Taxonomic revision of Marma Simon, 1902 (Araneae: Salticidae: Euophryini), pp. 287-353 in Zootaxa 4899 (1) on pages 289-292, DOI: 10.11646/zootaxa.4899.1.16, http://zenodo.org/record/4456831