Tullgrenella Mello-Leitão, 1941

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Tullgrenella Mello-Leitão, 1941: 191 (type species: Euophrys morenensis Tullgren, 1905; gender feminine, by original designation); Galiano, 1970: 324.

Akeloides Mello-Leitão, 1944: 374 (type species: Akeloides quadripunctatus Mello-Leitão, 1944). Synonymized by Galiano (1970: 324).

Diagnosis. Males of Tullgrenella resemble those of other related Freyina genera, such as: Aphirape C. L. Kock, 1850 by having a hook-shaped RTA retro-dorsally (see Galiano 1981b: 6, figs 21, 23); Sumampattus Galiano, 1983 by the presence of endites with distal projections (see Galiano 1970: fig. 15; Galiano 1981a: figs 1–2, 9; Galiano 1983: fig. 8), tegular membranous process (Figs 2D–2F) and pars pendula (see Galiano 1983: figs 6–7; Fig. 4F); Hisukattus Galiano, 1987 and Akela Peckham & Peckham, 1896 by having proximal retrolateral lobe on the tegulum (see Galiano 1999: fig. 2E). Females of Tullgrenella resemble those of other freyines, such as: Aphirape and Sumampattus by the window in ventral epigyne and the presence of posteriorly-opening coupling pocket (Figs 3C–D, 3F); Hisukattus and Akela by the copulatory ducts in concentric loops (Figs 3E–F). However, the genus Tullgrenella can be distinguished from the aforementioned genera by males having a cradle-shaped conductor (Figs 2F, 4A–F) and a tegular membranous process upside to the embolar base (Figs 4A, 4F, 8C), and females having the window on the ventral epigyne (Figs 5A–D) and monoplanar concentric copulatory ducts (Figs 7A–Q).

Description. Length 4.0–9.0 mm. Dark, reddish–brown salticids (Figs 1A–F, 20A–H, 32A–F, 45A–F). Carapace reddish-brown, cephalic region smooth and dark, thoracic region with slope and two perpendicular bands of white setae to each other on thoracic slope (Figs 8A, 9A). Anterior half of thoracic region at the same level as cephalic region. Chelicerae robust and vertical in both sexes (not dimorphic), promargin with two teeth (proximal reduced), and retromargin with one tooth (Figs 44A–B). Legs brown and yellow, with brown spots on distal portion and tarsus always lighter than other articles of legs. Legs formula: 4312/ 4132 in both sexes. Tarsal claws small, with five inferior teeth, not surpassing subungueal sulci (Figs 44C–D), and the distal one larger in females. Claw tufts poorly developed (Figs 44C–D, see arrow). Abdomen: dark or brownish–yellow, with white spots on dorsal posterior abdomen (Figs 8A, 10A, 13A–F, 20A–H, 23A, 32A–F, 33A); venter usually with a median longitudinal brown stripe of irregular contours (Figs 8B – 17B); sides with white longitudinal band at tip of anterior abdomen. Coloration of spinnerets as in legs. Male palp: femur macrosetae d1–1–1. Femur with ventral projection (Figs 43C–D, see arrow). Tibia with modest RTA with ventral sinuose or acute prong (Figs 8D, 23D) and usually incipient hook-like projection between the RTA and prolatero-dorsal tibial apophysis (Figs 8D, 10D, 14D, see red arrow). Usually with a small prolatero-dorsal tibial apophysis (Figs 8D, 14D). Cymbium apex suboval or sickle-shaped (Figs 4A–F). Tegulum oblique in relation to cymbium apex and central depression (Figs 14C, 21C, 25C, 29C, 33C, 40C, 48C, see red arrow) with inverted U-shaped spermophore visible (Figs 8C, 10C, 12C, 23C, 33C, 40C, 46C, 48C); tegular distal portion with developed proximal retrolateral lobe of tegulum (Figs 8C, 10C, 12C, 14C). Tegulum with two projections: membranous process of tegulum (globose or elongated) upside to embolar base (Figs 6A, 6C, 8C, 10C, 38C, 46C, 48C, see purple arrow) and one conductor (Figs 2F, 4A–F, 6A, 6C, 8C, 10C, 21C, 23C, 27C, 33C, 38C, 46C, 48C). Embolar base projecting dorso-prolaterally (Figs 6C, 6F, 6I). Embolus sclerotized and slender, embolus way starting at 8–9 o’ clock (left palp; Figs 4A–F, 6A–I, 8C, 21C, 27C, 33C, 46C). Epigynum: epigynal plate with well-developed posteriorly-opening coupling pocket (Figs 5A–D, 7A–Q) and usually with a window circling around the copulatory opening, where the membranous copulatory ducts below could inflate during the movement of copulation (Figs 5A–C, 7A–Q). Copulatory opening median-posteriorly located, sometimes forming a septum or semi-septum, normally facing up (Figs 5A–C, 9C, 11C, 15C, 47C, 49C). In dorsal view, the spiral copulatory ducts with one, two, or three loops, and usually dorso-posteriorly situated from spermathecae (Figs 7A–Q). Spermathecae circular to ovoid, with the base of the fertilization ducts placed above the spermathecae at the median-posterior region of the epigynum (Figs 7A–Q).

Note. Species of Tullgrenella (Figs 1A–F, 20A–H, 32A–F, 45A–F) have bright colors, as well as a slightly elongated body, as in most genera of the subtribe e.g., Phiale C. L. Koch, 1846, Aphirape, Sumampattus, Akela and Hisukattus . These features suggest possible mimetic relationships with other aposematically colored arthropods (Edwards 2015).

Distribution. Bolivia, Chile, Brazil, Paraguay, Uruguay, and Argentina (Figs 50–54).

Natural history. Most individuals were collected on outcrops. Their habitat can be considered the soil or epigeous shrubs that occur in southern South America (grasslands and rocks), such as Chacoan subregion (Brazil, Uruguay, and Argentina), Brazilian subregion (Northern Brazil), and Andean region (Chile). Few individuals were recorded in the South American Transition Zone (Figs 50–54). The altitudinal occurrence of these species varies from 4 to 4776 m a. s. l. (Figs 50–54).

Species groups. Based on the morphological variation of genital structures (palp and epigynum), as well as the geographic regionalization and altitudinal occurrence, we propose five species groups for the genus Tullgrenella: morenensis group (with six species, including the type species); quadripunctata group (three species); melanica group (two species); serrana group (six species, included four new species) and peniaflorensis group (two species). Putative synapomorphies of genital structures for each species group are suggested below.