Titanoleptes, Kury & Bernabé, 2023

Kury, Adriano B. & Bernabé, Tiago, 2023, Description of two new genera and five new species of clade K 92 from Bahia and Espírito Santo, southeastern Brazil (Arachnida, Opiliones, Gonyleptidae), Zootaxa 5351 (4), pp. 401-434 : 415-418

Titanoleptes gen. nov.

urn:lsid:zoobank.org:act:C662C684-A0C1-459C-A8B9-905D9E91CA23

“Undescribed genera SOD-01, SOD-02”— Kury et al. 2022: 79.

Etymology. From the Greek Τῑτάν (one of the titans, divine beings descending from the primordial deities) + leptes (truncation of generic name Gonyleptes ), referring to the very large size of its species. Gender masculine.

Type species. Titanoleptes hyperion sp. nov.

Included species. Titanoleptes calcar ( Roewer, 1913) , Titanoleptes cupidensis ( Soares & Soares, 1946) , Titanoleptes eros sp. nov. and Titanoleptes hyperion sp. nov.

Diagnosis. Coxa IV of male without retrolateral apical apophysis ( Fig. 16c ), except in T. cupidensis ( Fig. 20a ). Trochanter IV of male armed with a strong, retrolateral, apical apophysis ( Figs. 21a, c ). Patella and tibia IV covered with yellow acuminate tubercles ( Figs. 20d , 24b ). Femur IV of male either substraight ( Fig. 19a ) or strongly curved to the midline on basal 1/ 4 in dorsal view ( Figs. 17a, c ). Tibia IV armed with rows of acuminate tubercles ( Figs. 20d , 24b ). VP of penis with ear-flaps small, triangular, ectal vertex connected but not fused with flange. Flabellum quadrangular, as a stylized shell, proximal sides gently arched, non-serrate, distal half extremely reduced, with distal sides coalesced into one, arched and deeply serrate ( Fig. 22e ).

Distribution ( Figs 14–15 ). Southern Bahia State and Espírito Santo State. All four species of Titanoleptes occur in the WWF ecoregion NT0103 (Bahia coastal forests) and two subsets of two species each are distributed on opposite margins of the Doce River.

Internal and external relationships of the four species of Titanoleptes . At a first glance, one could easily identify two forms in Titanoleptes gen. nov.: (1) T. eros sp. nov. and T. cupidensis are gracile and have a median apophysis on free tergite III in both sexes, substraight femur IV, and naked glans; (2) T. hyperion sp. nov. and T. calcar are robust and lack an apophysis on free tergite III of males, have a heavily curved femur IV, and have a unique scutum covering the glans. The parapatrid pattern of these two pairs— T. eros + T. cupidensis and T. hyperion + T. calcar are found on opposite margins of the Doce River ( Figs. 13–14 )—is highly suggestive of a sister relationship between each of the members of these pairs, regardless of whether the pairs themselves form a more inclusive clade or not. This hypothesis, though, is not corroborated by any of the analyses. The cladistic analysis of Kury et al. (2022) explored different parameters to assess the arrangement of the four Titanoleptes in the phylogeny of Sodreaninae. The favored analysis using the setk script (with implied weights and K value ca. 12) resolved Titanoleptes as a sister clade of Friburgoia / Urodiabunus , with an internal branching sequence of T. eros , T. cupidensis , and then T. hyperion + T. calcar . IE settings of K = 1 and 5, plus equal weights, yielded the same result. However, Bayesian inference did not recover Titanoleptes at all, finding only the clade T. hyperion + T. calcar and then recovering T. eros and T. cupidensis in a trichotomy with the rest of the Sodreaninae. We have decided not to arrange the four species of Titanoleptes into two genera, despite the weak support for their monophyly.

Key to the species of Titanoleptes

1. Ocularium armed with two small spherical tubercles of lighter color contrasting with background ( Figs. 16g , 27b ); femur IV of male strongly curved to the median line at proximal one quarter, armed at least with two strong retrolateral spines ( Figs. 17 , 28 ); free tergite III unarmed in males, either unarmed ( T. hyperion ) or with median apophysis ( T. calcar ) in females ( Figs. 16b, g , 27b ).............................................................................................. 2

1’. Ocularium with two high erect spines concolor with background ( Figs. 19f , 23c ); femur IV of male substraight, without prominent retrolateral spines ( Figs. 21a–d , 24a–d ); free tergite III armed with a median apophysis in both sexes ( Figs. 19b, h , 23 a, f ).............................................................................................. 3

2. Body color mahogany with black reticle ( Figs. 16b, g ); tubercles of scutal area III strong, acuminate and contrasting dark brown ( Figs. 16b, g ); RD6 of male Fe IV straight ( Fig. 16a ); FT II-III of female with yellow cones ( Figs 16g, h )................................................................................................ T. calcar ( Roewer, 1913)

2’. Body color dark brown, almost black; tubercles of scutal area III small, rounded and contrasting pale yellow ( Fig. 27b ); RD6 of male Fe IV strongly curved ( Fig. 27a ); FT II-III of female entirely unarmed ( Fig. 1f )............... T. hyperion sp. nov.

3. Cx IV of male with strong inner apical apophysis ( Figs. 19b, c , 20a ); Tr IV with strong apical inner apophysis ( Fig. 20a ).......................................................................... T. cupidensis ( Soares & Soares, 1946)

3’. Cx IV of male unarmed internally ( Figs. 23a, b ); Tr IV with small apical inner apophysis................. T. eros sp. nov.

Figures

FIGURE 1. Examples in vivo of some of the species treated in this paper: a–c. Cajango camacanensis sp. nov., a. Alpha male, from Una, Bahia State, Brazil; b. Beta male from São José da Vitoria, Bahia State, Brazil; c. Female from Una, Bahia State, Brazil; d. Titanoleptes calcar (Roewer, 1913), male from Prado, Bahia State, Brazil; e–f. Titanoleptes hyperion sp. nov. from Braço do Sul, Espirito Santo State, Brazil, e. Male, f. Female. Photos by A. Anker & P. Martins (a–d) and A.B. Kury (e–f). All used with permission.

FIGURE 13. Cajango vindicator sp. nov., male holotype (MNRJ 9053), from Camacan, genital morphology: a. Penis, distal part, dorsal view; b. Same, sinistrolateral view; c. Same, ventral view.

FIGURE 14. Eastern Brazil, showing the distribution of the two robust species of Titanoleptes gen. nov., which are allopatrid, distributed in opposite margins of the Doce river.

FIGURE 15. Eastern Brazil, showing the distribution of the two gracile species of Titanoleptes gen. nov. which are allopatrid, distributed in opposite margins of the Doce River. These two species are less common in collections. The extension of the map is the same as that of the previous one for comparison.

FIGURE 16. Titanoleptes calcar (Roewer, 1913), (MNRJ 17818) from Sooretama: a. Male, habitus, dorsal view; b. Same, detail of dorsal scutum; c. Male, coxae I–IV, stigmatic area and free sternites, ventral view; d. Male, body, sinistrolateral view; e. Same, frontal view; f. Same, posterior view; g. Female, dorsal scutum, dorsal view; h. Same, sinistrolateral view. Scale bars = 1 mm.

FIGURE 17. Titanoleptes calcar (Roewer, 1913), male (MNRJ 17818) from Sooretama: a. Left femur IV, dorsal view; b. Same, prolateral view; c. Same, ventral view; d. Same, retrolateral view. Scale bars = 1 mm.

FIGURE 19. Titanoleptes cupidensis (Soares & Soares, 1946), (MNRJ 17869), male (A–F), and female (G, H), from Sooretama: a. Male, habitus, dorsal view; b. Same, detail of dorsal scutum; c. Same, ventral view; d. Same, dextrolateral view; e. Same, posterior view; f. Same, frontal view; g. Female, habitus, dextrolateral view; h. Same, dorsal view. Scale bars = 1 mm.

FIGURE 20. Titanoleptes cupidensis (Soares & Soares, 1946), male (MNRJ 17869), from Sooretama:a. Left coxa andtrochanter IV, dorsal view; b. Left patella IV, prolateral view; c. Left coxa to proximal femur IV, prolateral view; d. Left leg IV, ventral view. Scale bars = 1mm.

FIGURE 21. Titanoleptes cupidensis (Soares & Soares, 1946), male (MNRJ 17869), from Sooretama, left Fe IV: a. Dorsal view; b. Retrolateral (external) view; c. Ventral view; d. Prolateral (internal) view. Anterior direction upwards in a to c. Scale bars = 5 mm.

FIGURE 22. Titanoleptes cupidensis (Soares & Soares, 1946), (MNRJ 17869), from Sooretama, male genitalia, distal part: a. Dorsal view; b. Ventral view; c. Dextrolateral view; d. Same, panoramic; e. Latero-apical view; f. Dorso-apical view; g. Glans, dextrolateral view. Scale bars = 100 μm (a, b, d, e), 50 μm (c, f, g).

FIGURE 23. Titanoleptes eros sp. nov., male/female paratypes (MNRJ 7077) from ES, Santa Teresa: a. Male, habitus, dorsal view; b. Same, ventral view; c. Same, frontal view; d. Same, posterior view; e. Same, dextrolateral view; f. Female, habitus, dorsal view; g. Female, habitus, dextrolateral view. Scale bars = 1 mm.

FIGURE 24. Titanoleptes eros sp. nov., male paratype (MNRJ 7077) from ES, Santa Teresa, free tergites and proximal left leg IV: a. Dorsal view; b. Prolateral view; c. Ventral view; d. Retrolateral view. Scale bars = 1 mm.

FIGURE 27. Titanoleptes hyperion sp. nov., male paratype (MNRJ 18669) ES, Santa Teresa: a. Habitus, panoramic, dorsal view; b. Habitus, dorsal view; c. Same, ventral view; d. Same, sinistrolateral view; e. Same, frontal view; f. Same, posterior view. Scale bars = 1mm.

FIGURE 28. Titanoleptes hyperion sp. nov., male holotype (MNRJ 9076), from Domingos Martins, femur IV (each figure is rotated 90° along the proximal-distal axis relative to each adjacent one). From the top: retrolateral view. ventral view, prolateral view and dorsal view. All scale bars = 5 mm.