Collodinae Stimpson, 1871

Guinot, Danièle & Bakel, Barry Van, 2020, Extraordinary majoid crabs: the genus Esopus A. Milne-Edwards, 1875 in the new subfamily Esopinae subfam. nov., and erection of Paulitinae subfam. nov. (Crustacea, Decapoda, Brachyura, Majoidea, Inachoididae Dana, 1851), Zootaxa 4766 (1), pp. 101-127 : 113

Subfamily Collodinae Stimpson , 18714

Collodini Stimpson, 1871: 119.

Microrhynchinae Miers, 1879: 651 .

Collodini Števčić 2005: 99.

Colloidini [sic] Števčić 2013: 185.

Type genus. Collodes Stimpson, 1860 (type species by monotypy: Collodes granosus Stimpson, 1860 ).

Description (based on the type species of the type genus, C. granosus ). Carapace subovate-subpyriform, resting on setting gutter. Dorsal surface with tubercles and spines ( Rathbun 1925: pl. 36, fig. 1, pl. 217, fig. 1, Collodes granosus ; Garth 1958: pl. 3, fig. 5, C. granosus ; Hendrickx 1999: fig. 31A; Santana 2008: figs. 12C, 13A, C. granosus ). Rostrum short, tip bifid ( Rathbun 1925: fig. 31, pl. 217, fig. 1, C granosus ). Preorbital tooth small. Postorbital tooth well developed, not detached, laterally directed, inner margin concave, curving around eye. Antenna: articles 1, 2+3 fused to epistome, urinary orifice sunken in epistome; basal article with entire carina on internal margin, external margin coarsaly granulate ( Garth 1958: fig. 4A, C. granosus ); flagellum exposed in dorsal view. Proepistome long, laterally compressed, separated by small gap from spiniform process of epistome. Epistome approximatively as wide as long. Mxp3: merus expanded at its anteroexternal angle; crista dentata with small, acute teeth. Thoracic sternum flat, except the first three inflated sternites. Sternum/pterygostome junction incomplete, sternite 4 extended only on a short distance; Milne-Edwards openings not separated from chelipeds. Exposed pleurites 5–8 forming rather narrow collar, with crenulate margin. Male chelipeds robust, palm high; fingers gaping in a broad oval for basal three-fifths ( Hendrickx 1999: fig. 31B). Female chelipeds weak ( Rathbun 1925, pl. 36, fig. 2, C. granosus ). Ambulatory legs rather depressed, hairy, dactyli with setae. Pleon: first somite dorsal, produced into small spine. Male gonopore partially condylar, partially coxal ( Guinot et al. 2013: 127). Male pleon with somites 1–5 free, somite 6 being fused to telson (pleotelson); female pleon with somites 1–4 free, somites 5, 6 being fused to telson (pleotelson) ( Rathbun 1925: pl. 36, fig. 2; Santana 2008: fig. 15B, C. granosus ). G1: variously developed lobe with subapical aperture, cluster of long spines near tip ( Garth 1958: pl. E, fig. 2, C. granosus ). Pleonal-locking mechanism: button on sternite 5; socket on pleotelson.

Remarks. The genus Collodes that includes numerous species showing a variety of morphological forms is clearly paraphyletic. It is why our description of the Collodinae is essentially based on the type species of the genus, C. granosus from the Pacific coast of the Americas ( Hendrickx 1999: fig. 32), and will apply to it and only close genera.

The cladistic analysis of Santana (2008: 225, figs. 65, 66) did not recover a monophyletic group. Five species form a basal complex: Collodes armatus Rathbun, 1878 ( Melo 1996: fig. p. 199; Santana 2008: figs. 11A, 12A); C. inermis A. Milne-Edwards, 1878 ( Melo 1996: fig. p. 200; Santana 2008: figs. 12D, 16A; Santana & Tavares 2017: figs. 1C, 2C); C. leptocheles Rathbun, 1894 ( Santana 2008: figs. 17A, B, 18A, 19A; Alves-Júnior et al. 2019: fig. 4); C. levis Rathbun, 1901 ( Santana 2008: figs. 19C, 20A, C); C. tumidus Rathbun, 1898 ( Santana 2008: figs. 26C, 28A, C). Most of the ohers species form a polytomy: Collodes anartius Colavite, Windsor & Santana, 2019 ; C. gibbosus (Bell, 1835) ( Santana 2008: figs. 12B, 13A, 14B); C. granosus Stimpson, 1860 ( Hendrickx 1999: fig. 31; Santana 2008: figs. 12C, 15A); C. obesus A. Milne-Edwards, 1878 ( Santana 2008: figs. 19D, 21A); C. robustus Smith, 1883 ( Drach & Guinot 1983: fig. 3; Guinot & Richer de Forges 1997: fig. 12A; Santana 2008: figs. 22B, C, 23A); C. rostratus A. Milne-Edwards, 1879 ( Melo 1996: fig. p. 201; Santana 2008: figs. 23C, 24A); C. tenuirostris Rathbun, 1894 ( Santana 2008: figs. 25A, C, 26A, B); C. trispinosus Stimpson, 1871 ( Melo 1996: fig. p. 202; Santana 2008: fig. 27A, B); C. tuerkayi Santana & Tavares, 2017 ( Santana & Tavares 2017: figs. 1A, 2A). See also the molecular phylogenetic tree represented as maximum likelihood topology of three mitochondrial and two nuclear genes in seven of the 15 described species of Collodes ( Colavite et al. 2019: fig. 1).

Other included genera. Probably, there may be several, close to C. granosus , to be added in the future (see below, Status of other inachoidid genera).