taxonID	type	description	language	source
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	(FIGS 4 D, 5) = Portunus (Achelous) De Haan, 1833 (type species Portunus spinimanus Latreille, 1819, by monotypy; gender masculine).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	Included species: Twenty-eight.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	= Lupea anceps de Saussure, 1858 = Lupea duchassagni Desbonne in Desbonne & Schramm, 1867 = Neptunus sulcatus A. Milne-Edwards, 1879 Achelous angustus (Rathbun, 1898)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	Achelous binoculus (Holthuis, 1969) = Portunus binoculus Holthuis, 1969 Achelous brevimanus Faxon, 1895 Achelous depressifrons (Stimson, 1859) = Amphitrite depressifrons Stimpson, 1859	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	= Lupa leachii De Haan, 1833 Achelous gibbesii (Stimpson, 1859) = Lupa gibbesii Stimpson, 1859	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	= Cancer ponticus Herbst, 1790 = Portunus dufourii Latreille, 1819 = Eriphia prismaticus Risso, 1827 = Neptunus hastatus var. rubromaculatus Steinitz, 1932 Achelous inaequalis (Miers, 1881) comb. nov. = Neptunus (Amphitrite) inaequalis Miers, 1881 Achelous iridescens (Rathbun, 1894) = Neptunu s (Hellenus) iridescens Rathbun, 1894 Achelous isolamargaritensis (Türkay, 1968) = Portunus (Achelous) floridanus isolamargaritensis Türkay, 1968	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	diagnosis	Diagnosis: Carapace (Fig. 5 A) approaching hexagonal shape,> 1.5 times as broad as long; dorsal surface finely granulate, with regions moderately demarcated, often with ridges or patches of granules in centre of regions; urogastric depression distinctly posterior to halflength of carapace. Front (Fig. 5 B) with four triangular or rounded lobes. Orbit ellipsoidal; supraorbital margin with relatively deep median and reduced lateral fissures. Inner supraorbital lobe often truncate or subdivided into two teeth or lobes. Infraorbital margin with broad notch. Anterolateral margin with nine spiniform teeth: anterior eight ones subequal in size; last tooth lateral, large, in most cases two or more times longer than other teeth. Posterolateral junction of carapace usually rounded. Sutures (Fig. 5 C) on thoracic sternum well developed; thoracic sternites partly granular. Chelipeds with merus bearing three to five spines on anterior border, unarmed or with one spine distally on posterior border; carpus with single spine on outer face and spine on inner face that can be extremely long; upper surface of palm with two or three teeth including usual tooth near articulation with carpus; chelae (Fig. 5 D) costate, moderately unequal and heterodontic; in larger chela, molariform tooth present proximally on cutting edge of dactylus. Dactyli of pereiopods 2 – 4 ensiform or cultriform, rarely lanceolate, usually markedly costate; setose on ventral margin. Merus of pereiopod 5 distinctly longer than broad, posterodistal spine present but may be obsolete. Male pleon (Fig. 5 C) narrowly triangular; crest on third pleomere moderately laminar; lateral margins of third pleomere straight or convex; terminal part of posterior thoracic episternite fills interspace between anterior margin of pleomere 3 and thoracic sternite 8; third to fifth terga fused but unclear sutures may remain, combined part usually with keels, subequal to sixth pleomere; sixth pleomere with lateral margins straight or sinuous, convergent distally. Telson elongately triangular. First male (Fig. 5 E) gonopod of moderate length, arched or sinuous; basal part robust, lying obliquely inwards; distal part moderately slender, curved anterolaterally and tapering distally to slender tip; pair of first gonopods not touching medially in pleonal cavity. Female vulva (Fig. 4 D) elongately drop-like, with long axis usually almost parallel to anterior margin of sternite. Systematic position: The clade of four Achelous species analysed in the present three-marker study (Fig. 1; as Achelous or Portunus) is in a sister position to the thalamitine genus Charybdis. Their combined clade is nested within the basal polytomy of the four clades [i. e. together with the clade of the lupocycline Lupocycloporus and the new genus Alionectes, the clade of the portunine Portunus (s. s.) and Callinectes] and with the joint clade of all remaining IWP genera previously included in Portunus (s. l.). The true phylogenetic relationship of those four clades thus remains unresolved at present, and the subfamily composition of the family Portunidae remains provisional owing to the paraphyly or polyphyly of the subfamily Portuninae itself. In none of the recent molecular phylogenetic reconstructions (Spiridonov et al., 2014; Evans, 2018; Mantelatto et al., 2018; present study) is Achelous shown as related to Portunus, the nominotypic genus of subfamily Portuninae. Achelous and Lupella (see discussion on the latter genus below) are distinguished by a peculiar set of morphological characters, with some apparent plesiomorphies (e. g. sutures still visible on fused male pleomeres 3 – 5) and apomorphies (e. g. a tendency for a long inner spine on the cheliped carpus). For these reasons, Spiridonov (2020) has recently separated these two genera into a new subfamily Achelouinae. This was also regarded as a step towards purifying the concept of Portuninae, which remains most probably paraphyletic, even with Achelous excluded. In the present 16 S analysis (Fig. 2), all Achelous species form a well-supported clade of mixed Atlantic and eastern Pacific representatives together with Lupella forceps and some Atlantic ‘ Portunus ’ species. This is consistent with the phylogenetic results in the studies of Evans (2018) and Mantelatto et al. (2007, 2009, 2018). The systematic position of those Lupella or ‘ Portunus ’ taxa, however, remained unresolved in these papers.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFD9A64FFC34FE6FFD7AFA06.taxon	description	A series of species up to now considered to be ‘ Portunus ’ or ‘ Portunus (Portunus) ’ [as listed by Mantelatto et al. (2018) or Evans (2018), respectively], and here transferred to Achelous (see below), are superficially similar to the genus Portunus (s. s.) in having a relatively broad carapace and a single tooth on the posterior border of the cheliped merus. Portunus (s. s.) differs from Achelous by the triangular pleon with a rounded telson apex (Fig. 15 C) vs. with a sharp telson apex (Fig. 5 C) and by the male gonopods, which are straight and threadlike (Fig. 15 E) vs. distinctly shorter, arched (Fig. 5 E). Most Achelous spp. also have a distinct tooth on the posterior margin of the merus of the last pereiopods (Fig. 5 A). Although reduced in some Achelous species, this character is shared by this genus and the taxa of the lupocycline clade in the present analyses consisting of Alionectes and Lupocycloporus (Figs 6 A, 7 A; see below). Remarks: The current composition of Achelous consists of species distributed in the Atlantic and the eastern Pacific. Among them, some species from the previous nominotypic subgenus Portunus (as listed by Ng et al., 2008) are also present. The support for separating most of them into a taxon distinct from the mostly IWP Portunus (Portunus) has already been highlighted by the molecular results of Mantelatto et al. (2007, 2009, 2018); the separate biogeographical affiliations also sustain such subdivision. Mantelatto et al. (2009) elevated six species from the subgenus Achelous to generic level. Three others were transferred there from the subgenus Portunus, and one from the genus Cronius Stimpson, 1860. Mantelatto et al. (2009) suggested the position of a group of four Atlantic Portunus species (i. e. Achelous anceps, Achelous floridanus, Achelous hastatus and Achelous ventralis) to be unclear owing to their basal separation from the main assemblage of species of their genus. In a subsequent study, Mantelatto et al. (2018) revealed an isolated position for Portunus anceps, whereas in the former study it was placed among the other Achelous species mentioned above in this paragraph. This is also the case for our 16 S analysis (Fig. 2), where these four species form an isolated but basally well-supported clade together with two other Atlantic species, Achelous inaequalis and Achelous forceps (= previously Lupella forceps). In the recent systematic account of the Brazilian portunids, Rodrigues et al. (2017) retained Achelous anceps, Achelous floridanus and Achelous ventralis in the genus Portunus based on a morphological distinction between Achelous and Portunus originally proposed by Verrill (1908): dactyli of pereiopods 2 – 4 ensiform or cultriform, markedly costate vs. relatively broad, lanceolate, leaf-like or cultriform, indistinctly costate. Given that there is no substantial morphological distinction between these three species and Achelous hastatus from the genus Achelous, we regard them as being congeneric. Four species of Portunus (Portunus) or Portunus (Achelous) [as listed by Ng et al., 2008; i. e. P. (P.) acuminatus, P. (P.) affinis, (P.) P. xantusii and P. (A.) floridanus] were originally affiliated with the genus Achelous (see: Stimpson, 1860, 1871; Faxon, 1893; Rathbun, 1930). Given that they agree with the main diagnostic characters of the genus, Mantelatto et al. (2018) resurrected their original generic name for the first three of these. Recently, Marco-Herrero et al. (2021) have also listed the remaining species, P. (A.) floridanus, under the generic name Achelous but left Achelous gibbesii under Portunus, despite its new affiliation already provided by Mantelatto et al. (2009). A further Portunus species, the eastern Pacific Achelous minimus, has never before been included in Achelous, but was regarded as a subspecies of Portunus xantusii by Garth & Stephenson (1966), which was originally in Achelous (see Stimpson, 1860; above). It is transferred here to Achelous based on the morphology of its male pleon and gonopods (see Garth & Stephenson, 1966). Size: Mostly medium-size portunids; maximum recorded size (CL × CW) ranges from 13.8 mm × 32.4 mm in Achelous tuberculatus to 65.0 mm × 110.0 mm in Achelous spinimanus (Verrill, 1908; Rathbun, 1930; Garth & Stephenson, 1966; Williams, 1984). Ecological notes: Achelous spp. generally occur from the upper subtidal zone to a depth of 200 m, with most species having a relatively broad depth range [Verrill, 1908; Rathbun, 1930; Garth & Stephenson, 1966; Williams, 1984; de Melo, 1996; Rodrigues et al., 2017; i. e. from 0 to 500 – 550 m in Achelous spinicarpus and Achelous spinimanus (see Holthuis, 1959; de Melo, 1996) or even to 640 m in Achelous floridanus (see Williams, 1984)]. They are not usually reported from estuaries, except for Achelous inaequalis in West Africa (Manning & Holthuis, 1981). These crabs prefer mostly soft, often mixed substrates and sometimes hard bottoms with algae or seagrass (in the upper subtidal zone) or sessile macrofauna (Garth & Stephenson, 1966; de Melo, 1996). Several species were found swimming in the water column or on flotsam (i. e. Sargasso seaweed; Verrill, 1908; Rathbun, 1930; Garth & Stephenson, 1966; Jerde, 1967), which implies an important role of swimming in their ecology. Geographical range: Tropical and subtropical West and East Atlantic, Mediterranean and Tropical East 142 M. KOCH ET AL. Pacific (Rathbun, 1930; Garth & Stephenson, 1966; Manning & Holthuis, 1981; d’Udekem d’Acoz, 1999).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFDCA651FEBAFA2DFE24FDC8.taxon	description	(FIGS 3 A, 4 A, 6) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org: act: 5946851 F- 6789 - 4 AF 3 - 8 DB 9 - B 609146580 ED Type species: Neptunus (Hellenus) pulchricristatus Gordon, 1931, by present designation. Included species: Two. Alionectes pulchricristatus (Gordon, 1931) comb. nov. = Neptunus (Hellenus) pulchricristatus Gordon, 1931 = Neptunus (Hellenus) alcocki Gordon, 1930 [pre-occupied name, primary junior homonym of Neptunus (Hellenus) alcocki Nobili, 1906]	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFDCA651FEBAFA2DFE24FDC8.taxon	diagnosis	Diagnosis: Carapace (Fig. 6 A) broadly hexagonal, with outline of anterior half semicircular; width (without last lateral teeth) ~ 1.5 × length; dorsal surface convex; regions feebly developed. Front (Fig. 6 B) with four blunt triangular lobes projecting beyond tip of inner supraorbital lobe. Orbits large, circular, with inner supraorbital lobe rounded; upper border of orbit deep, without fissures; infraorbital margin with subrectangular shallow lateral notch. Anterolateral margin of carapace with nine teeth: eight anterior ones low, triangular or almost reduced, unequal in size; last tooth distinctly larger, lateral. Posterolateral junction of carapace subrectangular, slightly produced, pointed. Sutures of thoracic sternum well expressed; suture between thoracic sternum 6 and 7 developed in males and interrupted in females; suture 7 / 8 interrupted in males and almost fused in females. Thoracic sternites granular (Fig. 6 C). Merus of third maxilliped rounded distally, not produced anterolaterally. Chelipeds relatively robust; merus bearing three spines on anterior border and two spines distally on posterior border; carpus with outer spine. Chelae (Fig. 6 D) moderately unequal, heterodontic; palm with longitudinal serrated carinae, inner dorsal carina with one or two distal spines, proximal spine over articulation with carpus sharp or obsolete. In larger chela, well-developed molariform tooth present. Dactyli of pereiopods 2 – 4 cultriform, setose with fine plumose setae; merus of pereiopod 5 distinctly longer than broad; posterior margin with distal spine. Male pleon (Figs 3 A, 6 C) triangular, elongate, regularly tapering distally; pleomeres 2 and 3 forming low subequal crests, with at least second pleomere with crest marginally serrated. Lateral part of pleomere 3 in coaptation with terminal part of last thoracic episternite; third to fifth pleomere terga fused, without trace of sutures and keels; sixth pleomere elongate, tapering distally. First male (Fig. 6 E) gonopod relatively short and robust proximally; arched and steeply tapering distally to the slender tip; pair of first gonopods lying obliquely directed inwards, touching subdistal parts, with apices directed anterolaterally. Female vulva (examined in Alionectes pulchricristatus; Fig. 3 A) located in medial part of proximal portion of sternite, rounded, occupying more than one-third of sternite length, with long axis nearly perpendicular to anterior edge of sternite. Etymology: An arbitrary combination of the suffix of the name ‘ Xiphonectes ’ [from Greek, νεκτες, received] and the Latin alio, other, to point it out as ‘ another genus derived from Xiphonectes ’; gender masculine. Systematic position: The species of Alionectes were previously included in Xiphonectes, sharing with the latter, and with the new genera Eodemus, Incultus and Trionectes, a general body appearance with the carapace flattened and the similarly produced posterior anterolateral tooth, and the produced or subacute angle formed by the posterolateral and posterior margins. Alionectes is unique among these five genera mainly by the large circular orbits without dorsal fissures, by the distinct posterodistal tooth on the merus of the swimming leg and by the serrated margin of the transverse crest of the second (or also third) pleonal somites in males. The species of this genus, together with Eodemus, possess two posterodistal spines on the cheliped merus (vs. a single spine in Incultus and Xiphonectes, or one or two spines in Trionectes). The first male gonopods are, unlike those in the other four genera compared here, relatively short and proximally robust, and obliquely touch along their out-curved subdistal parts (for comparison of the gonopod shape and position, see the respective paragraph regarding Incultus). In the present phylogenetic analysis (Fig. 1), the representative of Alionectes occupies a position remote from the morphologically similar genera listed above, but in a sister position to Lupocycloporus. Both genera share a posterodistal spine on the merus of the last pereiopod (a character possessed by Lupocyclus Adams & White, 1848 and, despite being reduced, also by Achelous, along with the subfamily Thalamitinae; see Apel & Spiridonov, 1998), the subtriangular shape of the male pleon, the coaptation of the terminal part of the last thoracic episternite and the anterolateral margin of the third pleomere (shared also with Lupocyclus, but in a perfect state in Alionectes), and the stout, strongly hooked first male gonopods. Lupocycloporus is clearly distinguishable by its remarkably slender chelipeds and by the carapace bearing transverse ridges dorsally (instead of granulate patches on the dorsal regions in Alionectes) and rounded corners distolaterally (see also: Stephenson & Campbell, 1959; Spiridonov, 2020). Based on both morphological and molecular evidence, Alionectes is thus placed here in the subfamily Lupocyclinae, which previously contained only two genera, Lupocyclus and Lupocycloporus. Remarks: Although Alionectes spinipes was not included in the present molecular analysis, this species is easily included in the new genus because of its close morphological similarity to Alionectes pulchricristatus. Both species share orbits without any notch on the upper margin, the serrated margin of the blade-like ridge of the second pleonal somite, a spine on the merus of the swimming legs and almost identical gonopodes. Alionectes spinipes differs from Alionectes pulchricristatus by the triangular shape of the anterolateral teeth of the carapace (vs. reduced in size, tuberculiform), by the distinctly shorter lateral tooth (i. e. less than the width of the front; vs. distinctly longer than the width of the front in Alionectes pulchricristatus) and by a smooth, not serrated, ridge on the third pleonal somite (vs. serrated ridges on both the second and third pleomeres). Size: Relatively small crabs; the maximum reported size (CW) in Alionectes spinipes (male) is 26 mm (Stephenson & Rees, 1967 a) and in Alionectes pulchricristatus (female) 32 mm (Gordon, 1931). Ecological notes: The existing information for Alionectes pulchricristatus indicates it to be a subtidal species living mostly within the upper 100 m but known to a depth of ~ 290 m on various types of substrates (Stephenson & Rees, 1967 a; Apel & Spiridonov, 1998). Geographical range: Indo-West Pacific, so far known from the northern Indian Ocean (Gulf of Oman and eastwards) and in the western tropical Pacific from China and Vietnam to the Philippines, Indonesia and Australia (Stephenson, 1972 a, b; Apel & Spiridonov, 1998; present study).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC2A651FEEDFDAEFDADFD4C.taxon	description	(FIGS 4 E, 7)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC2A651FF4CFD79FBBBF9DB.taxon	materials_examined	Type species: Achelous whitei A. Milne-Edwards, 1861, by monotypy; gender masculine. Included species: Eight.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC2A651FF4CFD79FBBBF9DB.taxon	description	= Amphitrite gracilimanus Stimpson, 1858	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC2A651FF4CFD79FBBBF9DB.taxon	description	Rathbun, 1909	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC2A651FF4CFD79FBBBF9DB.taxon	description	= Portunus ponticus Fabricius, 1798 = Portunus ponticus Weber, 1795 (nomen nudum) Lupocycloporus sinuosodactylus (Stephenson, 1967) = Portunus sinuosodactylus Stephenson, 1967 Lupocycloporus wilsoni (Moosa, 1981) = Portunus (Lupocycloporus) wilsoni Moosa, 1981 Diagnosis: Carapace (Fig. 7 A) ~ 1.5 times as broad as long, approaching quasi-hexagonal shape, but with arcuate anterolateral margin, dorsally convex. Regions feebly developed, marked by granular ridges and patches. Front (Fig. 7 B) with four rounded or subtriangular lobes, median ones usually more producing than lateral ones; all lobes projecting beyond tips of inner supraorbital lobes, which are quasi-triangular in shape. Orbit nearly circular; supraorbital margin with two shallow, reduced fissures. Anterolateral margin armed with nine teeth: eight anterior ones increasing in size posteriorly or subequal, small, acute and projecting forwards; ninth tooth distinctly larger, lateral. Posterolateral angle of carapace broadly rounded. Sutures on thoracic sternum well developed (Fig. 7 C). Merus of third maxilliped with anteroexternal angle rounded or subrectangular, not produced laterally. Merus of chelipeds robust, with four to seven spines on anterior margin and two distal spines on posterior margin; carpus with spine on outer face. Chelae slender (Fig. 7 D), nearly equal, homoiodontic; palm with two subdistal spines on upper surface. Dactyli of pereiopods 2 – 4 cultriform, costate, weakly setose on ventral margin. Merus of pereiopod 5 distinctly longer than broad, with spine on posterior margin. Male pleon triangular (Figs 4 E, 7 C). Pleomeres 2 and 3 with low crests; lateral margins of pleomere 3 rounded, narrowing distally; posterior margin of last thoracic episternite touches most of anterolateral margin of pleomere 3. Pleomeres 3 – 5 fused, without sutures remaining but keeled. First male gonopod (Fig. 7 E) relatively short and robust proximally, curved and tapering distally; pair of first gonopods lying obliquely, touching subdistal parts, with apices directed anterolaterally or laterally. Female vulva (examined in Lupocycloporus gracilimanus) elongate, rounded on ends, located along most of the mesial margin of the sternite. Systematic position: Lupocycloporus has a distinct general morphology and extremely slender homoidontic chelae, similar to the chelae of Lupocyclus. Indeed, species of these genera exhibit sister-group relationships in recent phylogenetic reconstructions, and both genera were included in a separate subfamily Lupocyclinae Paulson, 1875 (Spiridonov et al., 2014; Evans, 2018). In the present phylogenetic reconstruction (Fig. 1), Lupocycloporus spp. form a basally unsupported clade with Alionectes pulchricristatus. Given that Lupocycloporus and Alionectes share common apomorphies [i. e. the swimming leg merus with a distinct posterodistal acute tooth (below), elongately subtrianglar male pleon and laterally bent gonopods with a swollen basal part], these and other characters (see discussion under Alionectes) might support their close relationship.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC1A654FCB6FDACFD59FCC1.taxon	description	Included species: Two. Arenaeus cribrarius (Lamarck, 1818) = Portunus cribrarius Lamarck, 1818 = Lupa maculata Say, 1818 = Arenaeus websteri H. G. Jones, 1968 Arenaeus mexicanus Gerstaecker, 1856 = Euctenota mexicana Gerstaecker, 1856 = Arenaeus bidens Smith, 1871 Diagnosis (modified from Williams, 1984): Carapace more than twice as wide as long, finely granulate. Front not so far advanced as outer orbital angles, with six teeth including inner orbitals; submedian pair of teeth at either side of central notch coalesced. Superior margin of orbit with two deep fissures dividing it into three lobes; inferior margin of orbit with wide external fissure, inner angle much advanced. Lower surface of carapace densely setose. Anterolateral margin with nine strong teeth, somewhat acuminate, heavily setose beneath; most posterior produced into strong lateral tooth. Chelipeds of moderate size; merus with three spines on anterior border and short tuberculiform one near distal end of posterior border; carpus with lateral ridges and inner spine, outer spine present or absent; palm with five longitudinal granulose ridges and two spines, one at articulation with carpus and another above base of dactylus. Walking legs rather short and broad, densely fringed with short setae. Swimming legs stout, unarmed. Pleomere 2 produced on each side into strong, sharp, slightly upcurved spine. Male pleon broadly triangular on segments 3 – 5, sixth segment elongate, telson narrowly triangular. First male pleopod long, slender, filiform, with short, swollen basal part. Remarks: No specimen was analysed in this study. Spiridonov et al. (2014) suggested Arenaeus to be the closest relative of Portunus (in the present restricted extent, below) and Callinectes, all relatively large swimming forms with a lack of conspicuous granular patches on the carapace, slender gonopods and leaf-like walking dactyli. Evans (2018) subsequently confirmed their close, but somewhat unresolved phylogenetic relationship. Size: Both males and females reach a width of 153 mm (Camp et al., 1977; Williams, 1984). G e o g r a p h i c a l r a n g e: We s t e r n A t l a n t i c, f r o m Massachusetts to Argentina; eastern Pacific, from Mexico to Peru (Rathbun, 1930; Zupolini et al., 2017).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC7A655FEDAFCD4FD65F925.taxon	description	(FIG. 8) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org: act: 98 CEC 882 - F 852 - 4 A 87 - 88 C 6 - 99 AB 5 FDC 0 C 51 Type species: Portunus lecromi Moosa, 1996, by present designation. Included species: Two. Allomonomia calla (Koch, Nguyen & ĎuriŠ, 2015) comb. nov.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC7A655FEDAFCD4FD65F925.taxon	diagnosis	Diagnosis: Carapace (Fig. 8 A) subhexagonal, with anterior outline semicircular; width (without lateral teeth) ~ 1.3 × length; dorsal surface with well-defined granulate regions. Front (Fig. 5 B) with four sharp triangular teeth, submedian teeth narrower than lateral teeth, all distinctly projecting beyond tip of inner supraorbital lobe. Epistome apophysis well developed and may be visible between median frontal lobes. Orbit relatively large, ellipsoidal, with supraorbital margin granulate, with two well-developed fissures in middle and on base of first anterolateral tooth; infraorbital margin strongly granulated, with narrow, deep lateral notch, with tooth-like elevation mesially and on ventral side of first anterolateral tooth. Anterolateral margin convex, longer than posterolateral margin, armed by nine teeth: eight anterior ones subequal, small, acutely forward projecting; ninth tooth distinctly larger, lateral. Posterolateral angle of carapace rounded. Sutures of thoracic sternum well expressed (Fig. 8 C); thoracic sternites partly granular. Merus of third maxilliped with anterolateral lobe subrectangular. Chelipeds moderately stout; merus with four spines on anterior and one spine distally on posterior border; carpus with usual sharp spine on outer face. Upper surface of palm with two granular crests, inner one ending by spine distally; outer surface with two granular crests ending on level of finger joint. Chelae (Fig. 8 D) nearly symmetrical but heterodontic; flattened molariform tooth present proximally at cutting edge of dactylus of one of the chelae. Dactyli of pereiopods 2 – 4 cultriform, costate, setose on ventral margin. Merus of pereiopod 5 a little longer than broad, without spine on posterior margin. Male pleon (Fig. 8 C) ‘ T’ - shaped. Posterior pleonal margin forming a pair of subparallel transverse laminar crests on both second and third pleomeres. Lateral margins of pleomere 3 straight, terminal part of posterior thoracic episternites narrow, fits in interspace between anterior margin of pleomere 3 and thoracic sternite 8. Pleomeres 3 – 5 fused, without sutures, but with elevated submarginal ridge posteriorly on fourth pleomere; telson narrowly triangular. First male (Fig. 8 E) gonopod moderately robust; distal half slender, held subparallel to each other; tip with wide opening. Female vulva remains undescribed. Etymology: Derived from the generic name Monomia [from Greek, meaning ‘ only one’], to which the new genus is generally similar, and the prefix allo- (Greek άλλο, other), alluding to the superficial morphological similarity but different systematic position of the new genus to the former; gender feminine. Systematic position: The two known species of Allomonomia were members of the subgenus / genus Monomia in earlier concepts (Ng et al., 2008; Koch et al., 2015 b). They share some general morphological characters (i. e. the carapace with a rounded posterolateral angle, the third meri of the maxillipeds with a triangular anterodistal projection, and the chelae with two teeth on the upper surface of the palm). Allomonomia differs from Monomia by the semicircular shape of the carapace (vs. more widened, semi-elliptic or hexagonal), the front with four subequal triangular teeth projecting distinctly beyond the inner supraorbital angle (vs. low and subequal ones with submedians lower than laterals, projecting slightly beyond the tip of the inner supraorbital angle), an indistinct median epistomial spine (vs. reaching distinctly beyond front), only one posterodistal spine on the cheliped merus (vs. predominantly two spines) and by the first male gonopods lying in a subparallel position, with broadened apical openings (vs. gonopods generally overlapping each other by bent region medially, with narrow terminal opening). In the present 16 S molecular analysis (Fig. 2), both species of Allomonomia are in a sister position to Incultus, whereas the three-marker analysis (Fig. 1) revealed paraphyly of Allomonomia, with the latter genus positioned as sister to Allomonomia calla. However, morphological characters provide proper support to an independent generic status of both genera. Both of them are very similar in the bent shape of the first male gonopods, which are lying in a pair with subparallel distal parts in Allomonomia (Fig. 8 E) but overlapping each other by the bent regions in Incultus (Fig. 13 E). Both these genera are also characterized by the presence of only a single posterodistal tooth on the cheliped merus. Together with this, Allomonomia species are different in the general appearance of the body from the species of Incultus, with a distinctly convex dorsal carapace with a semicircular anterior outline and rounded posterolateral angles (vs. hexagonal, deeply depressed, with elevated, swollen regions and with posterolateral angles acutely produced). The sister clade to the Allomonomia – Incultus clade in the present three-marker analysis (Fig. 1) is a branch composed of Xiphonectes and Trionectes (see below). The Allomonomia spp. were previously included in Monomia, whereas the species now included in Incultus and Trionectes were all previously included in the genus / subgenus Xiphonectes (as listed by Ng et al., 2008; Spiridonov et al., 2014). All species of the large, combined clade have the first male gonopod consisting of a robust basal part and bent distal part. The species of the Xiphonectes – Trionectes clade have the distal part short and hooked (Figs 16 E, 17 E), whereas species belonging to the Allomonomia – Incultus genera have the distal part of the first gonopod distinctly more sharply bent, more slender and longer (Figs 8 E, 13 E). Trionectes is also easily distinguished by having three frontal teeth on the carapace. Size: Although few specimens have been available for measurements, Allomonomia spp. appear to be medium-sized portunids, ranging from 20 mm × 30 mm in Allomonomia lecromi (Moosa, 1996) to a maximum CW of 38.4 mm in Allomonomia calla (Koch et al., 2015 b). Ecological notes: The two known species of the genus have been collected from relatively deep waters: Allomonomia lecromi at 120 – 150 m from the Chesterfield Islands (Moosa, 1996), Allomonomia calla at 106 – 111 m and from a tangle net deeper than 50 m, from the Philippines (Koch et al., 2015 b). Geographical range: Indo-West Pacific: known from Madagascar, the Philippines and New Caledonia.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC6A656FF4CF8DFFD4CF90C.taxon	description	Callinectes affinis Fausto-Filho, 1980 Callinectes amnicola Rochebrune, 1883 = Neptunus amnicola Rochebrune, 1883 = Neptunus edwardsi Rochebrune, 1883 = Neptunus marginatus var. truncata Aurivillius, 1898 = Callinectes latimanus Rathbun, 1897 Callinectes arcuatus Ordway, 1863 = Callinectes pleuriticus Ordway, 1863 = Callinectes dubia Kingsley, 1879 = Callinectes nitidus A. Milne-Edwards, 1879 Callinectes bellicosus Stimpson, 1859 = Callinectes ochoterenai Contreras, 1930 Callinectes bocourti A. Milne-Edwards, 1879 = Callinectes cayennensis A. Milne-Edwards, 1879 = Callinectes maracaiboensis Taissoun, 1972 Callinectes danae Smith, 1869 Callinectes exasperatus (Gerstaecker, 1856) = Lupea exasperata Gerstaecker, 1856 =? Lupa trispinosa Leach, 1816 = Callinectes tumidus Ordway, 1863 Callinectes gladiator Benedict, 1893 = Lupa smythiana White, 1847 (nomen nudum) Callinectes larvatus Ordway, 1863	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC6A656FF4CF8DFFD4CF90C.taxon	description	= Callinectes humphreyi H. G. Jones, 1968? Callinectes pallidus (Rochebrune, 1883) = Neptunus pallidus Rochebrune, 1883 Callinectes rathbunae Contreras, 1930 Callinectes sapidus Rathbun, 1896 [ICZN, 1964. Opinion 712] = Portunus diacantha Latreille, 1825 [name suppressed; ICZN, 1964. Opinion 712] = Callinectes sapidus acutidens Rathbun, 1896 Callinectes similis Williams, 1966 Callinectes toxotes Ordway, 1863 = Callinectes robustus A. Milne-Edwards, 1879 Diagnosis (modified from Rathbun, 1930): Similar to Portunus, distinguished by narrow, reversed ‘ T’ - shaped male pleon, and merus of third maxillipeds strongly produced outwards at anteroexternal angle. Body usually of large size; front formed by two or four teeth; dorsum of carapace with four dorsal carinae formed of single granules; two transverse, usually curved and subparallel lines on gastric region and one sinuous oblique line on branchial region leading inward from lateral tooth; nine strong anterolateral teeth, with posteriormost tooth at lateral angle; chelipeds stout; palm with five external carinae, with dorsodistal spine and proximal spine at articulation with carpus; carpus without inner spine, outer spine at widest part of article; merus with three (exceptionally four) stout spines on inner margin and small spine at posterodistal end; swimming legs without spines. Pleon of immature female triangular from fourth segment to telson. Remarks: As Spiridonov et al. (2014) noted, Callinectes is closely related to Portunus (in its herein redefined scope) and Atlantic Arenaeus, as is evident from their general appearance being relatively large swimming forms, sharing leaf-like walking dactyli, the lack of conspicuous granular patches on their carapace and slender gonopods. This view is confirmed by other recent cladistic (Karasawa et al., 2008) and molecular (Mantellato et al., 2009, 2018; Evans, 2018) studies of portunids. In the present study, a well-supported clade is formed by two species of Portunus and a single Callinectes (Arenaeus was not included in the present analysis). The infrageneric molecular diversity of American Callinectes analysed by Robles et al. (2007) revealed two major clades among 12 of 16 currently valid species included in that analysis: the ‘ bocourti ’ clade including Callinectes affinis, Callinectes bocourti, Callinectes rathbunae, Callinectes sapidus and Callinectes toxotes; and the ‘ danae ’ clade composed of Callinectes arcuatus, Callinectes bellicosus, Callinectes danae, Callinectes exasperatus, Callinectes larvatus, Callinectes ornatus and Callinectes similis. Morphological distinctions between these lineages are mainly in the relative length of the first gonopods (reaching the fourth thoracic sternite in the ‘ bocourti ’ group; see Williams, 1974). Size: Generally, in adult males and females the CW is between 100 and 150 mm (Rathbun, 1930); the largest males in low-salinity waters might reach even> 230 mm (Williams, 1965). Ecological notes: Species of Callinectes inhabit a variety of substrate types in estuaries and shallow oceanic waters, from the upper sublittoral zone to a depth of 40 m (Williams, 1965). Geographical range: The native range of the genus is along the Atlantic and Pacific coasts of America (Rathbun, 1930) and western Africa (Monod, 1956); recently introduced to European waters, including the Baltic, Mediterranean and Black seas (Nehring, 2011).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC5A658FF18F8EBFDCBFA6C.taxon	description	(FIG. 9)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC5A658FF18F8EBFDCBFA6C.taxon	description	Cavoportunus dubius (Laurie, 1906) = Neptunus (Achelous) dubius Laurie, 1906 = Portunus (Cycloachelous) yoronensis Sakai, 1974 Diagnosis (after Nguyen & Ng, 2010): Carapace hexagonal in shape, width ~ 1.3 × length, dorsal surface glabrous; frontal margin slightly upturned; surface microscopically granular with distinct regions; metagastric ridge distinctly visible, other ridges indistinct, dorsal surface with numerous small granules. Front with four teeth, slightly projecting beyond tip of internal supraorbital angle. Orbit relatively large, with moderate dorsal inclination; inner supraorbital margin rounded; upper border of orbit with single fissure. Infraorbital margin straight, with forward-directed tooth. Anterolateral margin with nine teeth: first tooth largest; second to ninth teeth smaller, subequal in size. Posterolateral junction of carapace rounded. Basal antennal article with lobe-like process. Merus of third maxilliped produced anterolaterally. Cheliped length ~ 1.8 × width of carapace; manus slightly less massive than merus; fingers relatively long, slender. Adult male with two prominent deep depressions on sternite eight inside sternopleonal cavity, anterior being larger, in which first gonopod sits. First male gonopod short, relatively stout, ‘ S’ - shaped; basal part relatively slender; subdistal part prominently dilated, lobiform; distal part with distinct folds on tip. Second gonopod elongate, tapering, about four-fifths length of first gonopod. Penis coxal, long, slender, proximally lodged between two raised processes of sternite 8. Female vulva mostly surrounded by thick, whitish, crescent-shaped rim; salient, short sternal cover on outer margin; separate, small, round operculum. Systematic position: In the present three-marker phylogenetic reconstruction (Fig. 1), Cavoportunus is resolved as an independent branch within a large clade of IWP genera previously belonging to Portunus (s. l.), in a sister position to two multigenera branches (i. e. the Allomonomia – Incultus – Trionectes – Xiphonectes clade and the Cycloachelous – Eodemus – Monomia clade). As pointed out by Nguyen & Ng (2010), Cavoportunus is morphologically most similar to Cycloachelous. Both taxa share some important synapomorphies, in particular a tendency to a decreasing carapace widthto-length ratio and approaching a circular carapace shape (Figs 9 A, 10 A), the secondary symmetrization and homoiodonty of the chelipeds (Spiridonov et al., 2014), and a short and stout first gonopod (Fig. 9 E), but which is morphologically complex and exhibits coaptation of the median groove. The deep depression 150 M. KOCH ET AL. (Fig. 9 E) on each side on the eighth thoracic sternite (to accommodate the lateral flange of the gonopod apex) is a unique autapomorphy for Cavoportunus (Nguyen & Ng, 2010). Cavoportunus can also be distinguished easily from Cycloachelous by the presence of a single fissure (vs. two shallow ones) on the supraorbital margin of the carapace and by a single distal spine (vs. two spines) on the posterior margin of the cheliped merus (Figs 9 A, 10 A). Remarks: Currently, Cavoportunus is monotypic, containing only the species Cavoportunus dubius. In the present study, the genus is represented by two specimens: a male Cavoportunus aff. dubius (MNHN-IU- 2008 - 12565) from the Marquesas Islands, and an ovigerous female Cavoportunus dubius (MNHN-IU- 2014 - 4099) from the Philippines. The size of the female (CW 15 mm) is close to the upper limit known for Cavoportunus dubius (see Nguyen & Ng, 2010), but except for sexual dimorphism there are no noticeable differences between the specimens. Those two specimens seem genetically different [3.0 % (COI) or 5.2 % (16 S)], hence the former might represent an undescribed species. S i z e: S m a l l c r a b s; m a x i m u m r e p o r t e d s i z e o f Cavoportunus dubius (female) is 14.3 mm × 18.5 mm (Nguyen & Ng, 2010). Ecological notes: The single representative of the genus is a low subtidal species occurring between 20 – 30 and 245 m (Spiridonov & Zhadan, 1999; Nguyen & Ng, 2010). Geographical range: Indo-West Pacific: from the Gulf of Aden (Neumann & Spiridonov, 1999) and Madagascar (present study, MNHN) to Japan, the Philippines, New Caledonia and French Polynesia (Spiridonov & Zhadan, 1999: fig. 1 A; Nguyen & Ng, 2010).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCBA65AFEEDFA07FD4EF91E.taxon	description	(FIGS 4 F, 10, 11)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCBA65AFEEDFA07FD4EF91E.taxon	description	= Neptunus (Achelous) octodentatus Gordon, 1938 Cycloachelous orbicularis (Richters, 1880) = Achelous orbicularis Richters, 1880 Cycloachelous orbitosinus (Rathbun, 1911) = Portunus (Achelous) orbitosinus Rathbun, 1911 Cycloachelous suborbicularis (Stephenson, 1975)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCBA65AFEEDFA07FD4EF91E.taxon	diagnosis	Diagnosis: Carapace (Fig. 10 A) quasi-hexagonal or almost circular in shape, usually <1.5 times broader than long, flattened, with granular ridges and / or patches or entirely covered with granules. Regions moderately developed. Orbit ellipsoidal, with two shallow fissures on supraorbital margin. Infraorbital margin visible in dorsal view, with pronounced notch; outer infraorbital lobe appressed to ventral side of outer supraorbital lobe (= first anterolateral tooth). Frontal margin (Fig. 10 B) consisting of four low lobes. Nine well-developed anterolateral teeth: first to eighth subequal; ninth somewhat longer than others or of similar size. Posterolateral junction of carapace rounded. Sutures on thoracic sternum distinct (Fig. 10 C); suture between thoracic sternites 6 and 7 nearly complete in males and interrupted in females; thoracic sternites usually smooth. Anterolateral angle of maxilliped 3 merus strongly produced laterally or auriculate. Chelipeds relatively slender; merus with three or four spines on anterior margin and two distal spines on posterior margin; carpus with single spine on anterior margin of outer face. Chelae (Fig. 10 D) usually of nearly equal size and homoiodontic, without proximal molariform tooth on cutting edge of dactylus; palm with two carinae and single spine on upper face (excluding spine at articulation with carpus) and three carinae on outer face. Dactyli of pereiopods 2 – 4 cultriform, costate, weakly setose on ventral margin. Merus of pereiopod 5 distinctly longer than broad, without spine on posterior margin. Male pleon (Fig. 10 C) quasi-triangular or with lateral margin of sixth pleomere markedly convex. Lateral margins of pleomere 3 straight or slightly concave; terminal part of posterior thoracic episternite fills interspace between anterior margin of pleomere 3 and thoracic sternite 8. Pleonal terga 3 – 5 fused, without remaining sutures; keels usually absent. Gonopod 1 (Fig. 10 E) short and stout, distally hooked, or moderately slender, curved. Female vulva (Fig. 4 F) a narrow, oblique fissure, relatively distant from margins. Systematic position: In our three-marker phylogenetic reconstruction (Fig. 1), Cycloachelous forms a sister group to Monomia and Eodemus, with unsupported internal relationships. Cycloachelous, together with Cavoportunus (see above), shares with these genera some morphological apomorphies, such as the female vulva in the form of a narrow slit, and (mostly) two distal spines on the posterior margin of the cheliped merus. However, Cycloachelous spp. are distinct in overall appearance from other genera of the former Portunus (s. l.) owing to their carapace shape and symmetrical chelipeds. Remarks: Cycloachelous granulatus has been considered to comprise two subspecies: the nominotypic one and Cycloachelous granulatus unispinosus, originally Achelous granulatus var. unispinosus Miers, 1884. The latter subspecies was described based upon a single type specimen from Prince of Wales Channel, Queensland, Australia (Miers, 1884: 230, pl. 23 B). Later, Miers (1886: 180) considered it as a separate species, Neptunus (Achelous) unispinosus. Miers (1884: 230) suggested it could be distinguished from Cycloachelous granulatus ‘ in wanting the submedian spine of the posterior margin of the arm of the chelipedes’, meaning by a single spine on the posterior margin of the cheliped merus, vs. two spines (a generic character of Cycloachelous). In the subsequent description, he added an important character of the frontal teeth or lobes: ‘ the median teeth smaller, but not less prominent than submedian teeth’ (Miers, 1886: 180). However, in Cycloachelous granulatus the median frontal teeth are distinctly less prominent than submedians (laterals in the present terminology). This indicates that Miers’ variety unispinosus is indeed a separate species, probably not even belonging to Cycloachelous. We (M. K. and Z. Ď.) examined the holotype specimen of Achelous granulatus var. unispinosus Miers, 1886 (NHM 82.7); the distinctions mentioned by Miers (above) were confirmed. In addition, the male pleons also differ in both the crabs, in that the penultimate somite is shorter, but wider anteriorly in Cycloachelous granulatus (vs. longer, but narrower distally) and the telson is respectively larger (vs. smaller). The morphology of P. (Monomia) euglyphus (as listed by Ng et al., 2008) shows some characters typical for Cycloachelous, the most significant being the unique shape of the male gonopods, which are similar to those of the type species, Cycloachelous granulatus (compare: Serène, 1969: fig. 6; Nguyen & Ng, 2010: fig. 5 D). The present analysis confirms the inclusion of Neptunus (Amphitrite) euglyphus Laurie, 1906 in Cycloachelous.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC9A65CFC14FD27FADDFA00.taxon	description	(FIGS 4 H, 12) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org: act: 0676 E 050 - 17 EF- 4030 - 8605 - BB 6 D 580 EC 7 D 7 Type species: Portunus pseudohastatoides Yang & Tang, 2006, by present designation. Included species: Six. Eodemus arabicus (Nobili, 1905) comb. nov. = Neptunus (Hellenus) arabicus (Nobili, 1905) = Portunus acerbiterminalis Stephenson & Rees, 1967 Eodemus hastatoides (Fabricius, 1798) comb. nov. = Portunus hastatoides Fabricius, 1798 = Portunus hastatoides Weber, 1795 (nomen nudum) E o d e mu s p s e u d o h a s t a t o i d e s (Yang & Tang, 2006) comb. nov. = Portunus pseudohastatoides Yang & Tang, 2006	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFC9A65CFC14FD27FADDFA00.taxon	diagnosis	Diagnosis: Cephalothorax and chelipeds with fine, hair-like marginal setae. Carapace (Fig. 12 A) hexagonal, more than two times as broad as long (including lateral teeth); dorsal surface tomentose, regions feebly defined by patches of fine granules. Front (Fig. 12 B) with three or four lobes slightly projecting beyond tip of inner orbital lobe; median tooth or pair distinctly smaller than laterals. Orbit relatively large, approaching circular; inner supraorbital lobe rounded; supraorbital margin with two narrow fissures with appressed walls; infraorbital margin with broad, ‘ Y’ - shaped lateral notch. Anterolateral margin with nine sharp teeth: first to eighth teeth subequal in size; eighth tooth arising anteriorly from base of ninth tooth; ninth tooth large, directed laterally. Posterolateral junction of carapace produced to distally upturned spiniform tooth. Sutures of thoracic sternum well expressed (Fig. 12 C); thoracic sternites usually smooth or sparsely granular. Merus of third maxilliped anterolaterally produced to elongate projection. Chelipeds relatively robust; merus with four spines on anterior and two spines distally on posterior border; carpus with usual outer spine. Chelae (Fig. 12 D) slightly unequal, heterodontic; in larger chela, molariform tooth present proximally at cutting edge of dactylus; fingers densely setose distomedially. Dactyli of pereiopods 2 – 4 cultriform, costate, setose on ventral margin. Merus of pereiopod 5 distinctly longer than broad, without spine on posterior margin. Male pleon ‘ T’ - shaped (Fig. 12 C), with sixth pleomere elongate, narrow. Lateral margins of pleomere 3 markedly concave; pleomere terga 3 – 5 fused, without remaining sutures. First gonopod (Fig. 12 E) relatively long, arched, tapering distally; pair of first gonopods not overlapping, only touching each other in median body plane, with distal parts directed anteriorly. Female vulva (Fig. 4 H) in form of a narrow slit located close to mesial margin of sternite. Etymology: Derived from Latin eodem, ‘ to the same place’, alluding to the similar outer appearance of included species; gender masculine. Systematic position: The species of the portunine Eodemus and of the newly established genera Incultus and Trionectes, plus the lupocycline Alionectes, were previously affiliated in the genus (or subgenus) Xiphonectes (as listed by Ng et al., 2008; Spiridonov et al., 2014). These genera are similar in their overall appearance, with a more or less strongly depressed carapace, usually with a long pair of lateral teeth and elevated patches of granules on the dorsum, and with produced or subacute posterolateral angles. Their morphological distinction, together with Xiphonectes in its present reduced extent (below), is discussed under those genera. Eodemus can be distinguished easily by the more distinct ‘ T’ - shape of the male pleon, concave lateral margins of the third male pleomere and the distinctly laterally produced anterolateral lobe of the third maxilliped merus. Based on our three-marker analysis, Eodemus is closely related to the species of Monomia (s. s., below) and Cycloachelous. Those genera share, among other features, the overall shape of the carapace, with granular patches on regions of the dorsum, and four frontal teeth, with the submedian pair smaller (in some Eodemus spp., the submedians are fused to a single tooth; see also Wong et al., 2010). Other potential synapomorphies are the produced anterolateral lobe of the third maxilliped merus, two posterodistal spines on the cheliped merus, the ‘ T’ - shaped male pleon, concave lateral margins of the third pleomere (at least in most Monomia) and a slit-shaped female vulva located close to the mesial margin of the sternite. The first gonopods are relatively long, usually bent and distally slender in Eodemus and Monomia (s. s.), whereas they are stout or moderately slender and curved in Cycloachelous. All examined specimens of Eodemus are relatively small (CW usually ≤ 41 mm), whereas the species of Monomia are represented by distinctly larger specimens (see below). The posterior lateral tooth of the carapace is of variable length in both genera, but generally longer in Eodemus. The posterolateral junction of the carapace is produced to a distally upturned spine in the new genus, whereas it is greatly rounded in Monomia spp. In the latter, the epistome is medially produced to a long median spine distinctly overreaching the front, whereas such a structure is not recognizable from the dorsal view in Eodemus spp. The first male gonopods, despite the generally similar shape in both genera, touch in the median body plane along their bent parts in Eodemus spp., but in the majority of Monomia spp. they overlap each other (except for some Australian species with subparallel, not overlapping, distal parts of the gonopods). Remarks: For Monomia argentata, the iridescence of the cuticle had been incorporated as a typical character to its specific name by White (1847; as Amphitrite argentata; a nomen nudum). The name was subsequently also accepted by A. Milne-Edwards (1861; as Neptunus argentatus). This remarkable phenomenon of iridescence was also observed by us in E. pseudohastatoides (see Koch et al., 2015 a), which also contributes to the hypothetical relationship of Eodemus and Monomia indicated by the molecular analysis (Fig. 1). Among all other species of the genus, E. arabicus (types examined, MNHN B 5926, B 5927) can be distinguished by the posterolateral junction of the carapace not as upturned as in the former two species. Neptunus (Hellenus) tweediei, P. trilobatus and P. dayawanensis have recently been synonymized with Xiphonectes unidens (now E. unidens) by Nguyen & Ng (2021). Two new Xiphonectes species described in the latter study, X. subtilis and X. vassilyi, as formerly belonging to the X. hastatoides species complex (Nguyen & Ng, 2021), are now also affiliated in Eodemus. Size: Maximum reported size ranges from CW 25 mm [E. unidens, in Stephenson & Rees, 1967 a (as Portunus tweediei)] to 19.4 mm × 41.0 mm [E. hastatoides, in Apel & Spiridonov, 1998 (as Portunus hastatoides)]. Ecological notes: Some species appear to be mostly shallow-water living species, such as E. arabicus recorded in the intertidal to the upper subtidal zone down to 55 m (Red Sea, ZMMU Ma 3295) or E. hastatoides with a depth range from 7 to 100 m (Stephenson & Rees, 1967 a; Stephenson, 1972 a, b; Moosa, 1981 a; Apel & Spiridonov, 1998; Yang et al., 2012; Nguyen & Ng, 2021). Geographical range: Indo-West Pacific: from the Red Sea and the western Indian Ocean to Japan, the Philippines, Indonesia, Australia, the Chesterfield Islands and New Caledonia (Crosnier, 1962; Stephenson, 1972 a, b; Moosa, 1996; Apel & Spiridonov, 1998; Neumann & Spiridonov, 1999; Poupin, 2010).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCFA65FFC15F9F0FB04FEDF.taxon	description	(FIGS 3 A, 4 E, 13) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org: act: 2 B 1 B 6 DEC- 6146 - 4 A 5 B-A 77 B- 4340 C 140 CDF 5 Type species: Neptunus tuberculosus A. Milne-Edwards, 1861, by present designation. Included species: Three. Incultus alcocki (Nobili, 1905) comb. nov. = Neptunus (Hellenus) alcocki Nobili, 1906 Incultus brockii (De Man, 1887) comb. nov. = Neptunus brockii De Man, 1888	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCFA65FFC15F9F0FB04FEDF.taxon	diagnosis	Diagnosis: Carapace (Fig. 13 A) flattened, broadly hexagonal,> 1.5 times as long as broad (including lateral teeth). Dorsal surface granulated; regions conspicuously elevated, with pair of closely set cardiac elevations most prominent; distinct wide depressions present behind orbits. Front (Fig. 13 B) with three or four subequal low or obsolete lobes distinctly projecting beyond inner supraorbital lobe. Orbit relatively large, ellipsoidal, with inner supraorbital lobe rounded; upper border of orbit with two short fissures. Infraorbital margin granulated, with broad lateral notch. Anterolateral margin with nine blunt or lobiform teeth; ninth distinctly larger, lateral. Posterolateral angles of carapace produced, distinctly upturned over posterior margin. Sutures (Fig. 13 C) and grooves of thoracic sternum poorly to moderately distinct; thoracic sternites partly granular. Antennal flagellum long, more than twice orbital width. Merus of third maxilliped elongate, anterolaterally produced to rounded lobe. Chelipeds stout; merus with three or four spines on anterior border, one distal spine on posterior border, followed more basally with distinct, obliquely elevated sulcus forming small tuberculate lobe and continuing as transverse groove or crossing distal merus. Carpus with short spine on outer face. Chelae (Fig. 13 D) moderately unequal, heterodontic, with flattened molariform tooth located proximally at cutting edge of dactylus; palm unarmed or with small spine distally over dactylar articulation, two distinct longitudinal crests on outer surfact and two on upper surfaces of palm; fingers distinctly shorter than palm. Dactyli of pereiopods 2 – 4 cultriform, costate, setose on ventral margin. Merus of pereiopod 5 distinctly longer than broad, without spine on posterior margin. Male pleon (Fig. 13 C) quasi-triangular; crests on second and third pleomeres low. Lateral margins of third pleomere straight; apex of thoracic episternite 7 fills in interspace between anterior margin of pleomere 3 and episternite 8; third to fifth somites fused, without remaining sutures, crests or keels on them may be present; last pleomere elongate. First male gonopod (Fig. 13 E) stout or moderately slender distally, arched or almost rectangularly bent; tip rounded, with moderately wide subterminal opening; gonopods in pair overlapping each other by bent regions (not obvious in I. alcocki possessing relatively short gonopods). Female vulva (Fig. 4 E) elliptic, located at mesial or posteromesial margin of sternite. Etymology: The genus name is derived from the Latin adjective ‘ incultus ’, meaning rough, unkempt, but is here treated as a masculine noun in apposition. It alludes to the somewhat untidy external appearance of the known species; gender masculine. Systematic position: All three included species were listed in the subgenus Xiphonectes by Ng et al. (2008), but later accepted at generic level following Spiridonov et al. (2014). The included species are similar to Xiphonectes (as restricted in this study) and to Eodemus, Trionectes and the lupocycline Alionectes, in general appearance of the body. The carapace in all these genera is flattened, with a similarly long posterior tooth of the anterolateral series and with an angulate junction of the posterolateral and posterior margins. Incultus bears a morphologically close relationship to Xiphonectes and Trionectes. It differs from these genera by: (1) distinctly elevated dorsal regions on the carapace (vs. feebly demarcated in both); (2) lobate anterolateral teeth on the carapace (vs. predominantly sharply triangular in both); (3) the front with three or four low and often indistinct subequal teeth (vs. four distinct triangular or rounded teeth, with submedian ones smaller in Xiphonectes, and three subequal triangular or blunt teeth, with the median one more slender in Trionectes); (4) the infraorbital margin with a broad lateral notch (vs. narrow in Xiphonectes and deep, ‘ V’ - shaped in Trionectes); (5) the male pleon with second and third somites each bearing a low transverse crest (vs. second high, laminar and third shallow; the latter might also be high in some Xiphonectes spp.); (6) the male pleon with the sixth somite elongately trapezoid (vs. similar in Trionectes, but distinctly constricted subdistally in Xiphonectes); (7) the stout chelipeds (vs. comparatively slender in Xiphonectes), with fingers distinctly shorter than the palm (vs. subequal in both other genera); and (8) the first male gonopod with a rounded tip, being relatively long, almost rectangularly bent at the midlength at least in I. brockii and I. tuberculosus; the pair of gonopods overlap each other by the bent regions (vs. gonopods slender, smoothly out-curved distally, not meeting medially in Xiphonectes; similar but shorter and stouter in Trionectes).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCFA65FFC15F9F0FB04FEDF.taxon	description	Eodemus is distinguishable from Incultus by deep orbits, with two incisions with fully appressed walls, and with the inferior margin bearing a broad ‘ Y’ - shaped lateral notch. The cheliped merus bears two spines distally on the posterior border. The male pleon is almost ‘ T’ - shaped, with the sixth somite elongate, narrow (vs. pleon subtriangular in Incultus, with the sixth pleomere trapezoid). The first gonopods are relatively long, arched, distally slender; the pair of first gonopods are touching in the median body plane at the bent parts. Based on the present molecular analyses, Incultus is closely related to Allomonomia. Species of both taxa have the first male gonopods bent in a similar way; those lie in a pair overlapping each other by their bent regions in Incultus gen. nov., while being subparallel with distal parts in Allomonomia. Both genera are also characterized by the presence of only a single posterodistal tooth on the cheliped merus. Additionally, Incultus is different from Allomonomia in having a hexagonal and deeply depressed carapace with elevated swollen regions and the posterolateral angles acutely produced; whereas in Allomonomia, the carapace is distinctly convex dorsally, with a semicircular anterior outline and with rounded postrolateral angles. In addition to the general shape of the body mentioned above, Alionectes differs from Incultus mainly by large circular orbits without dorsal fissures (vs. with small, ‘ Y’ - shaped fissures in Incultus) and by a distinct posterodistal tooth on the merus of the swimming legs. The cheliped merus bears two spines distally on the posterior border (vs. a single distal spine and an oblique keel more proximally). The male pleon has the second and third somites with subequally low crests, marginally serrated on the second pleomere. The first male gonopods are relatively short and robust proximally, tapering distally to the out-curved tip; the pair of first gonopods lies obliquely inwards, touching at their subdistal parts. Remarks: Neptunus (Hellenus) alcocki Nobili, 1905 (= Portunus alcocki; see: Stephenson, 1972 a, b; Neumann & Spiridonov, 1999) has until recently been included in the subgenus Portunus (Xiphonectes). It is known from the holotype male and a single female from the Gulf of Aden (Neumann & Spiridonov, 1999); it has also been recorded in the Red Sea [Laurie, 1915; and present material, SMF (V. A. S.)]. The species was not included in the present molecular analyses. Based on the holotype, illustrated by Guinot (1957: figs 3, 4, 9, 10) and examined by us (M. K. and Z. Ď.), the species possesses three low frontal lobes, a character typical for Trionectes or some species of Eodemus and Xiphonectes. However, it shows a closer morphological affinity to Incultus rather than to other portunine genera. Its median frontal tooth is evidently derived from the fusion of two small submedian teeth and it has low lateral frontal teeth, similar to I. brockii, with four obsolescent teeth forming a shallowly undulating margin (in I. tuberculosus, the two submedians are distinct but small). In contrast, in the other three genera the frontal teeth are triangular. The stout arched first male gonopods are also distinct from those of Trionectes, which distally are strongly bent and slender, with a minute terminal opening, whereas in I. alcocki they only feebly taper distally to a moderately wide opening (see Guinot, 1957: figs 8 – 10). Furthermore, I. alocki shares with other Incultus spp. the unique morphology of the rounded female vulva located at the mesial or posteromesial margin of the sternite (compare Neumann & Spiridonov, 1999: fig. 2 C; Apel & Spiridonov, 1998: fig. 117 d) and is herein thus transferred to Incultus. Size: Small to medium-sized species; maximal recorded size range from 7.8 mm × 16.3 mm in I. alcocki (Neumann & Spiridonov, 1999) to 20.5 mm × 35.0 mm in I. brockii (Yang et al., 2012) and 29.0 mm × 49.0 mm in I. tuberculosus (Stephenson & Rees, 1967 a). Ecological notes: The included species show different depth ranges, with I. brockii being an intertidal to upper subtidal species (Spiridonov, 1999), whereas I. tuberculosus occurs on various substrates from the upper subtidal to continental slope, ranging from 7 – 9 to 580 m in depth (Stephenson, 1972 a, b; Apel & Spiridonov, 1998; Yang et al., 2012). Incultus alcocki appears to be an upper subtidal species occurring mostly on sand (Laurie, 1915; Neumann & Spiridonov, 1999; present material, SMF). Geographical range: Indo-West Pacific: I. tuberculosus has the broadest range, from the Red Sea and East Africa to the Hawaiian Islands (Stephenson, 1972 a, b; Apel & Spiridonov, 1998); I. brockii is restricted to the eastern Indian Ocean and the western Pacific; and I. alcocki is currently known only from the Red Sea and the Gulf of Aden (see above).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCCA662FC11FE81FF61FEDF.taxon	description	(FIGS 3 D, 4 G, 14) = Portunus (Amphitrite) De Haan, 1833 {type species Portunus gladiator Fabricius, 1798, subsequent designation by Miers (1886); name pre-occupied by Amphitrite Müller, 1771 [Polychaeta]; gender feminine. = Portunus (Monomia) Gistel, 1848 (replacement name for Amphitrite De Haan, 1833; gender feminine). Included species: Ten.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCCA662FC11FE81FF61FEDF.taxon	description	Monomia gladiator (Fabricius, 1798) = Portunus gladiator Fabricius, 1798 = Cancer menestho Herbst, 1803	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFCCA662FC11FE81FF61FEDF.taxon	diagnosis	Diagnosis: Carapace (Fig. 14 A) quasi-hexagonal; ~ 1.6 – 1.8 times as broad as long; margins setose, dorsal surface tomentose, with well-defined granulate regions. Front (Fig. 14 B) with four subtriangular blunt teeth, submedian teeth usually distinctly lower than lateral ones, all teeth projecting slightly beyond tip of inner orbital lobe; median epistomial apophysis well developed, reaching distinctly beyond front. Orbit relatively large, ellipsoidal, with upper margin granulate and two well-developed fissures, in middle and near base of first anterolateral tooth; lower orbital margin with large lateral notch, one or two strong obtuse mesial teeth, and outer infraorbital lobe appressed to ventral side of outer supraorbital lobe (= first anterolateral tooth). Anterolateral margin of carapace convex, slightly longer than posterolateral margin, armed by nine teeth: eight anterior ones subequal, small, acute and projecting forwards; ninth tooth distinctly larger, lateral. Posterolateral angle of carapace broadly rounded. Sutures on thoracic sternum well developed (Fig. 14 C). Merus of third maxilliped with anterolateral lobe subtriangular. Chelipeds moderately stout; merus with four or five spines on anterior border and two (rarely one) spine on posterior border; carpus with spine on outer face, which may be reduced; upper surface of palm with two granular crests, inner one strong, ending by spine distally; outer surface with two granular crests ending at level of finger joint, lower arched, creating outer ventral, sharply produced margin. Chelae (Fig. 14 D) slightly unequal, weakly heterodontic, with molariform tooth on cutting edge of dactylus (this molariform tooth may be reduced in large specimens). Dactyli of pereiopods 2 – 4 styliform, densely setose on ventral margin. Merus of pereiopod 5 a little longer than broad, without spine on posterior margin. Male pleon ‘ T’ - shaped (Figs 3 D, 14 C). Pleomeres 2 and 3 forming transverse laminar crests; lateral margins of pleomere 3 usually distinctly concave; apex of posterior thoracic episternite fills interspace between anterior margin of pleomere 3 and thoracic sternite 8. Pleomeres 3 – 5 fused, but a groove remains at place of suture between pleomeres 3 and 4; usually keels and crests present on these pleomere terga. Pleomere 6 as long as broad or longer, sometimes with a small median spine directing backwards on distal part; with telson narrowly triangular. First male gonopods (Fig. 14 E) long, bent; gonopods in pair overlapping each other by bent regions, with distal parts directed anterolaterally, or subparallel and touching only medially. Female vulva (Fig. 4 G) slit-shaped, located at anteromesial margin of sternite, often covered with cuticular cup. Systematic position: The genus is morphologically similar to the newly established genus Allomonomia (see discussion under that genus). They share the following morphological features: rounded posterolateral angles of the carapace, third maxilliped merus with a triangular anterodistal projection and the chelae with two teeth on the upper surface of the palm. Monomia differs from Allomonomia by the front having four low and subequal teeth, with the submedians lower than the laterals, projecting slightly beyond the tip of the inner supraorbital angle, the median epistomial spine reaching distinctly beyond the front, predominantly two posterodistal spines on the cheliped merus and by the first male gonopods in a pair usually overlapping each other along their bent parts. The three-marker phylogenetic analysis (Fig. 1) shows Monomia as most closely related to Cycloachelous and Eodemus (see above), together forming a basal well-supported clade. As already discussed, these three genera are similar in the presence of granular patches on the dorsal regions of the carapace, four frontal teeth with the submedian pair smaller, a produced anterolateral lobe of the third maxilliped merus, two posterodistal spines on the cheliped merus (rarely single in Monomia spp.), the ‘ T’ - shaped male pleon, concave lateral margins of the third male pleomere, and the first gonopods relatively long, usually bent, distally slender, and a slit-shaped female vulva. They differ, as discussed above for Eodemus, in mostly larger body sizes in Monomia, usually longer posterior lateral tooth and acutely produced posterolateral angle of the carapace in Eodemus (latter rounded in Monomia) and the medially produced epistome in Monomia. The first male gonopods overlap each other along their bent parts in the majority of Monomia spp. (except for some Australian spp. with subparallel, not overlapping, distal parts of the gonopods), whereas they only touch at their bent parts in Eodemus. Cycloachelous shares with the aforementioned taxa some apparent morphological apomorphies, such as a slit-shaped female vulva and (generally) two distal spines on the posterior margin of the cheliped merus. However, Cycloachelous spp. are distinct in their general appearance from other genera owing to their circular carapace shape, stout gonopods with unique shapes and symmetrical chelipeds (see Remarks for the respective genera). Remarks: Monomia now contains ten species. Previously, 14 species were included in Monomia (see Ng et al., 2008; Koch et al., 2015 b; Koch & ĎuriŠ, 2018). Four species are herein removed and transferred to different genera: two species, P. (M.) lecromi and M. calla, are placed in the new genus Allomonomia; P. (M.) euglyphus is transferred to Cycloachelous; and P. (M.) ponticus is transferred to Lupocycloporus. There has been substantial confusion regarding the nomenclature of the type species of the genus Monomia, Portunus gladiator Fabricius, 1798. It has been considered as a secondary homonym of Cancer gladiator Fabricius, 1793, a junior subjective synonym of P. sanguinolentus (Stephenson & Cook, 1973; Chertoprud et al., 2012). Stephenson & Cook (1973) then suggested Amphitrite haani Stimpson, 1858 as the next available synonym of P. gladiator. However, there is current agreement that this suggestion of homonymy is based on a misinterpretation of the International Code of Zoological Nomenclature (Ng et al., 2008; Spiridonov et al., 2014; Windsor et al., 2019). Furthermore, both morphological and molecular genetic evidence (Windsor et al., 2019) clearly indicate that M. gladiator and M. haani are distinct species and that M. pseudoargentata (Stephenson, 1961) is a junior subjective synonym of M. haani. The morphological examination of the Australian species, M. curvipenis (one male, AM P. 80096) and M. rubromarginata (1 female, MV J 45390; four males, MV J 45533) showed some characteristics such as the small median spine directed backwards on the distal part of the sixth male pleomere and a different shape of the first gonopods compared with the rest of the species. The generic affiliation of these species needs to be confirmed by further molecular analyses. Size: These are medium to large portunid crabs, with the maximum size (CL × CW) ranging from 22.0 mm × 38.0 mm (M. argentata; Yang et al., 2012) to 61.0 mm × 111.6 mm (M. haani, this study; ZIN RAN male). Ecological notes: This group comprises predominantly lower subtidal crabs occurring to depths not exceeding 100 m, with soft, mainly sandy bottoms (Sakai, 1939, 1976; Crosnier, 1962; Dai & Yang, 1991; Apel & Spiridonov, 1998; Chertoprud et al., 2012), but M. argentata seems to be a deepwater species known from depths down to 400 – 402 m (four males, MNHN-IU- 2018 - 4942). Geographical range: Indo-West Pacific: from the Red Sea and south-eastern Africa to Japan, the Philippines, Australia, New Caledonia and Samoa (Sakai, 1939, 1976; Stephenson, 1972 a, b; Apel & Spiridonov, 1998; Poupin, 2010); there is a questionable record of M. argentata from the Hawaiian Islands (Castro, 2011).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	(FIGS 3 C, 4 B, 15) = Portunus Weber, 1795 (type species Cancer pelagicus Linnaeus, 1758, designation by Rathbun, 1926; gender masculine) [Opinion 394]; see Holthuis (1952). Full synonymy: see Ng et al. (2008): 152 [Portunus (Portunus)]. Included species: Thirteen.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	= Cancer pelagicus Forskål, 1775 [pre-occupied name, primary junior homonym of Cancer pelagicus Linnaeus, 1758] = Cancer cedonulli Herbst, 1794 = Portunus denticulatus Marion de Procé, 1822 = Portunus (Portunus) pelagicus var. sinensis Shen, 1932	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	Portunus reticulatus (Herbst, 1799) = Cancer reticulatus Herbst, 1799	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	= Cancer sanguinolentus Herbst, 1783 = Cancer gladiator Fabricius, 1793 = Callinectes alexandri Rathbun, 1907 =? Cancer raihoae Curtiss, 1938	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	= Portunus tropicalis Marion de Procé, 1822 = Lupea pudica Gerstaecker, 1856 = Lupea parvula Desbonne, in Desbonne & Schramm, 1867 Portunus segnis (Forskål, 1775) = Cancer segnis Forskål, 1775	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	= Neptunus trituberculatus Miers, 1876 Diagnosis: Carapace (Fig. 15 A) usually broadly hexagonal, much broader than long, width (without lateral teeth) - to-length ratio> 1.5. Dorsal surface finely granulated, with regions feebly demarcated, sometimes with narrow rows of granules in their centres. Urogastric depression distinctly anterior (rarely slightly posterior) to half length of carapace. Front (Fig. 15 B) with four triangular or rounded teeth; median ones may be extremely reduced. Supraorbital margin with two short fissures; infraorbital margin with shallow lateral notch. Epistome apophysis well developed, may be producing medially as prominent spine far overreaching submedian frontal teeth. Anterolateral margin with nine spiniform teeth; teeth of anterior series (except last tooth) short, subequal in size; last tooth distinctly larger, lateral. Posterolateral junction of carapace rounded. Most of the sutures on thoracic sternum indistinct (Fig. 15 C), thoracic sternites generally smooth. Merus of third maxilliped with anteroexternal angle rounded or subrectangular, not produced laterally. Chelipeds usually stout; merus with posterior border subparallel to anterior (or convex), bearing three or four spines on anterior border, and unarmed or with one spine distally on posterior border. Carpus with a spine and carinae on outer face; in a few cases, carina ends in an additional spinule. Chelae (Fig. 15 D) slightly inaequal, heterodontic upper surface of palm with two distal teeth; molariform tooth present proximally at cutting edge of dactylys of larger chela. Dactyli of pereiopods 2 – 4 relatively broad, lanceolate, leaf-like or cultriform, indistinctly costate, densely setose on ventral margin. Merus of pereiopod 5 subquadrate, only little longer than broad or, more rarely, elongate, distinctly longer than broad, without a spine on posterior margin. Male pleon (Figs 3 C, 15 C) narrowly triangular, almost smooth. Pleomeres 2 and 3 with low keels, in some species not visible in ventral view; lateral margins of pleomere 3 straight or concave; terminal part of posterior thoracic episternite fills interspace between anterior margin of pleomere 3 and thoracic sternite 8; third to fifth pleomeres fused, without remaining sutures but with indistinct transverse keels; combined part distinctly longer than sixth pleomere, the latter with lateral margins convergent distally; telson triangular and rounded, subequal or shorter than sixth pleomere. First male gonopod (Fig. 15 E) with short and robust transverse base and long, string-like distal part, basally curved and broadly arching anteriorly, touching medially; apices straight, reaching telson basis. Female vulva (Fig. 4 B) in form of a transverse slit or rounded, located in medial part of proximal portion of sternite, with long axis subparallel or oblique to anterior margin of sternite. Systematic position: Based on the present molecular results and a morphological comparison, Portunus is closely related to Callinectes. Spiridonov et al. (2014) presented Portunus, Callinectes and Atlantic Arenaeus as key members of the subfamily Portunidae, with unresolved positions for other species of Portunus (s. l.) diagnosed by being relatively large swimming forms sharing leaf-like walking dactyli, the lack of conspicuous granular patches on the carapace and slender gonopods. The other ‘ Portunus ’ species were separated into multiple subgenera (see listing by Ng et al., 2008), some of which have since been raised to generic level (Mantelatto et al., 2009; Nguyen & Ng, 2010; Chertoprud et al., 2012; Spiridonov et al., 2014). Here, they are all nested inside a major monophyly as part of an unresolved basal polytomy together with Portunus – Callinectes, Achelous, Lupocyclinae and the Thalamitinae clades (Fig. 1). Remarks: From 24 species of Portunus (Portunus) as listed by Ng et al. (2008), the present generic reassignment reduces this to 13 species. Portunus mauritianus Ward, 1942 had already been synonymized with P. (P.) segnis by Ng et al. (2008); three species (P. asper, P. gibbessi and P. rufiremus) were transferred to Achelous by Mantelatto et al. (2009) and four others (P. acuminatus, P. affinis, P. minimus and P. xantusii) by Mantelatto (2018); and P. floridanus has recently been transferred to Achelous by Marco-Herrero et al. (2021). The remaining four species (P. anceps, P. hastatus, P. inaequalis and P. ventralis) are assigned to Achelous in this study (see above). Portunus serratifrons (Montrouzier, 1865) was already suggested to be a juvenile Scylla serrata by A. Milne-Edwards (1873), but it is still listed herein as a valid species of Portunus, albeit rather provisionally. The generic affiliation of P. madagascariensis (Hoffmann, 1877) needs to be re-examined owing to its close similarity to P. sanguinolentus. The species is thus here provisionally regarded as valid.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF1A664FEC0FEB7FBF2FED0.taxon	description	Size: The genus includes the group of medium-sized species P. pubescens (maximum size CL × CW 25.0 mm × 45.0 mm; Crosnier, 1962), P. convexux (30.4 mm × 51.6 mm; our data, SMF male), P. rufiarcus (CW 27.1 mm, the only known specimen, holotype; Davie, 1987) and P. sayi (37.1 mm × 76.1 mm; Rathbun, 1930). The maximum known sizes of the remaining larger species reach distinctly higher v a l u e s, e. g. C L × C W: 6 1.5 m m × 1 3 6.0 m m i n P. sanguinolentus (see Apel & Spiridonov, 1998), 77.2 mm × 163.4 mm in P. reticulatus (see Lai et al., 2010), 82.0 mm × 188.0 mm in P. trituberculatus (see Yang et al., 2012) and CW 173 mm in P. armatus (female MNHN-IU- 2018 - 1413). Ecological notes: Portunus spp. are generally shallow-water dwellers, confined mostly to the intertidal and upper subtidal zones, preferring sandy substrates but also occurring on other types of substrates, notably seagrass meadows; larger-sized species are known for their complex migration pattern and frequent occurrence in the outer parts of estuaries. Portunus sanguinolentus and especially P. sayi are commonly associated with flotsam, thus travelling large distances over deep waters (reviewed by Spiridonov, 2013). Portunus sayi is likely to reproduce when living on floating sargassum (Verrill, 1908; Rathbun, 1930). Geographical range: Indo-West Pacific (majority o f s p p.); t r o p i c a l w e s t e r n A t l a n t i c (P. s a y i); Mediterranean Sea (P. segnis, Lessepsian migrant from the Red Sea, now common Mediterranean species; Lai et al., 2010).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF7A666FC25FE80FB25FB44.taxon	description	(FIGS 3 B, 4 C, 16) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org: act: BCC 68580 - 6 F 16 - 44 F 1 - 8 DDC-D 41241 E 34 F 7 B Type species: Portunus (Amphitrite) tenuipes De Haan, 1835, by present designation. Included species: Seven. Trionectes andersoni (De Man, 1887) comb. nov. = Neptunus (Hellenus) andersoni De Man, 1887 Trionectes mariei (Guinot, 1967) comb. nov.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF7A666FC25FE80FB25FB44.taxon	description	= Portunus spiniferus Stephenson & Rees, 1967 Trionectes tenuipes (De Haan, 1835) comb. nov. = Amphitrite tenuipes De Haan, 1835) Trionectes tridentatus (Yang, Dai & Son, 1979) comb. nov. = Portunus tridentatus Yang, Dai & Song, 1979 Diagnosis: Carapace (Fig. 16 A) broadly hexagonal, width ~ 1.5 times as long as broad (without lateral teeth); dorsal surface variously granulate, with regions moderately demarcated. Front (Fig. 16 B) with three distinct triangular or rounded lobes, median lobe subequal but more slender than lateral ones, all three lobes distinctly projecting beyond tip of inner supraorbital lobe. Orbit with inner supraorbital angle truncated, with oblique distal margin; dorsal orbital margin with two short fissures; infraorbital margin with deep ‘ V’ - shaped lateral notch. Anterolateral margin with nine spiniform, triangular or lobiform teeth, irregular in size; last tooth distinctly larger, lateral. Posterolateral junction of carapace angular, pointed, often distinctly upturned, sharp. Sutures of thoracic sternum unclear (Fig. 16 C); thoracic sternites may be granular. Merus of third maxilliped elongate, distally markedly produced to triangular, trapezoidal or rounded lobe. Chelipeds robust, with merus bearing three or four spines on anterior border and one or two spines distally on posterior border; carpus with single spine on outer face, which, in some species (T. pseudotenuipes), is obsolete; upper surface of palm with one or two distal teeth. Chelae (Fig. 16 D) slightly inaequal, heterodontic, with moderate molariform tooth at cutting edge of dactylus in larger chela. Dactyli of pereiopods 2 – 4 cultriform or styliform, costate, with short setose emargination on ventral margin. Merus of pereiopod 5 a little longer than broad, without posterodistal spine. Male pleon (Figs 3 B, 16 C) quasi-triangular. Pleomeres 2 and 3 with laminar crests. Lateral margins of pleomere 3 straight or convex; terminal part of posterior thoracic episternite fills interspace between anterior margin of pleomere 3 and thoracic sternite 8. Pleomeres 3 – 5 fused without remaining sutures, but indistinct keels may be present. Pleomere 6 with lateral margins sinuate. First male gonopod (Fig. 16 E) short, curved, tapering distally to slender tip; pair of first gonopods lying obliquely directed inwards but isolated, not touching medially, with apices curved anterolaterally. Female vulva (Fig. 4 C) (examined in T. mariei and T. tenuipes) in form of an oblique widened slit, located in medial part of proximal portion of sternite. Etymology: Given that species were previously included in Xiphonectes, the new generic name is a combination of the Greek ‘ τρί’, three (pointing on trilobed front) and ‘ νήκτης’, swimmer, thus a ‘ trilobed swimmer’; gender masculine. Systematic position: All species of Trionectes, together with the new genera Eodemus, Incultus and the lupocycline genus Alionectes, were previously considered members of Xiphonectes (see listing by Ng et al., 2008). All these taxa are similar in overall appearance, with a more or less strongly depressed carapace, usually with a long pair of lateral teeth and elevated patches of granules on the dorsum and with a produced or subacute posterolateral angle. Based on the morphological comparison and the present molecular analysis, Trionectes is most closely related to Xiphonectes (s. s.; present paper), and Incultus is more basal (see remarks for Xiphonectes, below). Trionectes differs from these genera by: (1) the front with three subequal triangular or blunt teeth in Trionectes (vs. four distinct triangular or rounded teeth, with submedian ones smaller in Xiphonectes, or four shallow subequal teeth in Incultus); (2) feebly demarcated dorsal regions of the carapace (vs. distinctly elevated in Incultus); (3) predominantly sharply triangular anterolateral teeth on the carapace (vs. lobate in Incultus); (4) the infraorbital margin deep, ‘ V’ - shaped, in Trionectes (vs. with narrow lateral notch in Xiphonectes, or with broad lateral notch in Incultus); (5) the male pleon elongately trapezoid (vs. same in Incultus, but with the sixth pleomere distinctly constricted subdistally in Xiphonectes); (6) the chelipeds stout (vs. same in Incultus, but comparatively slender in Xiphonectes), with fingers subequal to palm length (vs. same in Xiphonectes but distinctly shorter in Incultus); (7) the first male gonopods short, stout, subdistally bent, not meeting medially (vs. similar but more slender, smoothly out-curved in midlength in Xiphonectes, but relatively long, bent in midlength, with pair of gonopods overlapping each other medially in Incultus); and (8) the female vulva, which is located at the mesial or posteromesial margin of the sternite in Incultus and has a different position in Trionectes and Xiphonectes. From the lupocycline Alionectes, Trionectes can be distinguished easily by the tridentate front, the presence of a dorsally incised orbital margin, which is entire in Alionectes, the posterodistally unarmed merus of the swimming leg, and by the smooth, entire margin of the transverse crest on the second pleonal somite in males (vs. serrated on the second pleomere in Alionectes). Trionectes is also easily distinguished from Eodemus by the tridentate front, but also by the subtriangular male pleon (vs. a ‘ T’ - shaped pleon) and by the distally produced triangular or circular rounded, not distolaterally produced, third maxilliped merus (vs. a distinctly laterally produced anterolateral lobe). The genera also differ by the position and shape of the female vulva, which is in the form of a widened slit in Trionectes and somewhat distant from the margin of the sternite, and chinking located close to the mesial margin in Eodemus. Remarks: Three species differ from the remaining ones in the genus. The examined MNHN and ZMMU specimens of T. mariei and MNHN specimens of T. spiniferus and the species T. tridentatus, based on published reports (Dai & Yang, 1991; Yang et al., 2012), all have distinct unequal and sharp anterolateral teeth on the carapace, the third maxillipeds with a rounded distal projection, more robust chelipeds and three spines on the upper surface of the chela palm, but they all share the trilobed front, and the pleon bears a distinct laminar crest on the second pleomere typical for Trionectes. Only T. spiniferus was available for the present molecular analysis but, based on the morphological similarity of all three species and its deeply nested position in the Trionectes clade (Fig. 3), the inclusion of T. mariei and T. tridentatus in the genus appears to be well founded. Size: Small to medium-sized crabs; maximum known size ranges from CW 16 mm in T. rugosus (A. Milne-Edwards, 1861) to CL × CW: 24.5 mm × 53.3 mm in T. tridentatus (Yang et al., 2012). Ecological notes: The species with better-documented records (i. e. T. tenuipes and T. pseudotenuipes), occur mostly in the upper subtidal zone, to a depth of 35 m, on sand or muddy sand (Ramadan, 1936; Monod, 1938; Spiridonov, 1999). The few known records of T. mariei are also from upper subtidal sand patches on reefs (Guinot, 1957; present study: ZMMU Ma 3408 and SMF 47928 specimens). Geographical range: Indo-West Pacific: from the Red Sea (Ramadan, 1936; Monod, 1938; present study) and the western Indian Ocean to the Philippines and New Caledonia (Stephenson, 1972 a, b; Spiridonov, 1999; Poupin, 2010; Yang et al., 2012).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	description	(FIG. 17)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	description	Xiphonectes aculeatus Koch & ĎuriŠ, 2019 Xiphonectes bidens (Laurie, 1906) = Neptunus (Hellenus) bidens Laurie, 1906 Xiphonectes gracillimus (Stimpson, 1858) = Amphitrite gracillimus Stimpson, 1858 Xiphonectes guinotae (Stephenson & Rees, 1961) = Portunus guinotae Stephenson & Rees, 1961 Xiphonectes hainanensis (H. - L. Chen, 1986)	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	description	= Portunus (Xiphonectes) macrophthalmus Rathbun, 1906	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	description	= Portunus stephensoni Moosa, 1981	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	description	= Portunus tenuicaudatus Stephenson, 1961	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	diagnosis	Diagnosis: Carapace (Fig. 17 A) broadly hexagonal, about twice as broad as long when lateral spines are included, width (without lateral teeth) ~ 1.5 × length; dorsal surface variously granulate, with regions moderately demarcated, with patches of granules, sometimes in some species projecting to centre, forming elevated tubercles. Front (Fig. 17 B) with four triangular or rounded lobes; submedian lobes distinctly smaller than laterals, sometimes fused, projecting beyond tip of inner supraorbital lobe. Orbit nearly circular, with inner supraorbital lobe small, angulate, often merged to inner supraorbital margin, latter with two short fissures; infraorbital margin clearly visible in dorsal view, with narrow lateral notch. Anterolateral margin with six to nine spiniform teeth; particular teeth may have a tendency to reduction; teeth of anterior series (except last tooth) short, irregular in size, last tooth distinctly larger, lateral. Posterolateral junction of carapace angular, pointed, often distinctly upturned, sharp. Thoracic sternites partly granular, with sutures (Fig. 17 C) well demarcated. Merus of third maxilliped elongate, distally produced to rounded lobe, rarely angulate. Chelipeds usually relatively slender; with merus bearing three to five spines on anterior border and one spine or obtuse lobe distally on posterior border; carpus with a sharp spine on outer face; upper surface of palm with two (rarely one) distal teeth. Chelae (Fig. 17 D) slightly inaequal, heterodontic, with a moderate molariform tooth at cutting edge of dactylus in larger chela. Dactyli of pereiopods 2 – 4 cultriform, costate, with short, hairy emargination on ventral margin. Merus of pereiopod 5 a little longer than broad. Male pleon quasi-triangular (Fig. 17 C). Pleomeres 2 and 3 with low transverse smooth crests, that on pleomere 2 higher; lateral margins of pleomere 3 convex or acuminated; terminal part of posterior thoracic episternite fills interspace between anterior margin of pleomere 3 and thoracic sternite 8. Pleomeres 3 – 5 fused without apparent sutures, but feeble keels may be present. Pleomere 6 with lateral margins sinuate, constricted subdistally, rarely elongate, simple. First male gonopod (Fig. 17 E) in most cases elongate, curved, tapering distally to slender tip; pair of first gonopods lying obliquely directed inwards but not touching medially, apices curved anterolaterally. Female vulva relatively large, located in medial part of proximal portion of sternite, drop-shaped or ovoid, with long axis subparallel or oblique to anterior margin of sternite. Systematic position: Xiphonectes (as listed by Ng et al., 2008; Spiridonov et al., 2014) is subdivided here, based on the present molecular analyses, into five genera, of which four (i. e. Eodemus, Incultus, Trionectes and the lupocycline Alionectes] are newly established. The four new genera are nested inside three unrelated clades, and two of them occupy sister positions with other currently known or new portuniine genera. All these taxa are similar by their overall appearance, with a more or less strongly depressed carapace, usually with a pair of long lateral teeth, elevated patches of granules on the dorsum, and produced or subacute posterolateral angles. Of the above new taxa, Trionectes is sister positioned in the present molecular analysis to the newly restricted Xiphonectes, whereas Incultus is more basal. Xiphonectes differs from these by: (1) feebly demarcated dorsal regions of the carapace (vs. distinctly elevated in Incultus); (2) predominantly sharply triangular anterolateral teeth on the carapace (vs. lobate in Incultus); (3) frequent cases of reductions in the number of anterolateral teeth (vs. a full set of nine teeth in Incultus and Trionectes); (4) the front with four distinct triangular or rounded teeth, with submedian ones smaller (vs. four shallow subequal teeth in Incultus and three subequal triangular or blunt teeth in Trionectes); (5) the infraorbital margin with a narrow lateral notch (vs. a broad lateral notch in Incultus and deep, ‘ V’ - shaped notch in Trionectes); (6) the male pleon with the sixth somite distinctly constricted subdistally (vs. elongately trapezoid in Incultus and Trionectes); (7) the chelipeds comparatively slender in Xiphonectes (vs. stout in the other two), with fingers subequal to the palm length (vs. distinctly shorter in Incultus); (8) the first male gonopods slender, smoothly out-curved distally, not meeting medially in Xiphonectes (vs. short and stout in Trionectes, but relatively long, bent in midlength, with the pair of gonopods overlapping each other medially in Incultus); and (9) by the female vulva being located relatively distant from the sternite margins, vs. located at the mesial margin or posteromesial in Incultus. Xiphonectes is distinguishable from Alionectes by the presence of a dorsally incised orbital margin, which is entire in Alionectes, by a single posterodistal spine on the cheliped merus (vs. two spines in Alionectes) and by a smooth laminar crest on the second pleonal somite of males, whereas the margin is serrated in the latter genus.	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
A600031FFFF5A669FC81FB55FEF0F8A4.taxon	description	Eodemus is distinguishable easily from Xiphonectes by the ‘ T’ - shaped male pleon (vs. subtriangular) and by the distinctly laterally produced anterolateral lobe of the third maxilliped merus (vs. distally produced, rounded or angular, not distolaterally produced) and by a slit-like female vulva. Remarks: Since Xiphonectes was raised to full generic status by Spiridonov et al. (2014), it has become one of the most species-rich portunid genera, containing, according to Ng et al. (2008), 28 species, with X. longispinosus composed of four subspecies. Ng et al. (2008) include X. longispinosus longimerus Spiridonov, 1994, which they consider to be an available name for the species. This name was originally introduced as Portunus longispinosus forma longimera by one of us (Spiridonov, 1994). When introducing this name, Spiridonov (1994: fig. 5) used an infrasubspecific rank with the status of ‘ forma’, intentionally making this name unavailable (ICZN, 1999: Articles 10.2 and 15.2); the reason for this was to introduce the forms of X. longispinosus as a preliminary step towards the revision of this species complex. Recently, this species was described by Spiridonov (2016) as X. tuerkayi Spiridonov, 2016. The list by Ng et al. (2008) does not include X. leptocheles. This species was described and illustrated by A. Milne-Edwards (1873: 159, pl. 4, fig. 1), but was synonymized, without particular discussion, with X. longispinosus by Stephenson & Campbell (1959). Later, this name disappeared even from major portunid accounts (i. e. Stephenson, 1972 a, b; Sakai, 1939, 1976; Crosnier, 2002), but was listed in Xiphonectes (s. l.) in the studies by Spiridonov (2016) and Nguyen & Ng (2021) as a synonym of X. longispinosus. The photograph of the type specimen of X. longispinosus from HMCZ was previously kindly sent by A. Baldinger to one of us for another study (Spiridonov, 2016: fig. 1). Its current re-examination and comparison with the description and illustration of X. leptocheles confirms a separate status of the latter species. As a result of the present revision, only 14 species currently remain in Xiphonectes, with the four former X. longispinosus subspecies all elevated to species level. The remaining species are herein transferred to the newly established Eodemus (six species), Incultus (three species), Trionectes (eight species) and to the new lupocycline Alionectes (two species). Two species that were not included in our molecular analysis, X. latibrachium and X. paralatibrachium, show some morphological differences; they have the median frontal teeth projecting beyond the lateral ones, indistinct supraorbital notches and more robust chelipeds. Their inclusion in Xiphonectes should be confirmed by future genetic analyses. During the examination of the MNHN material in Paris, additional morphotypes of the X. longispinosus complex, with some potentially new species, were recognized by one of us (M. K.). A further revision of this group may therefore be necessary. Size: Small crabs; maximum reported size ranges from CL × CW: 8.8 mm × 17.0 mm in X. guinotae (Apel & Spiridonov, 1998) to 14.0 mm × 30.3 mm in X. iranjae (Yang et al., 2012). Ecological notes: The genus includes both upper subtidal species (e. g. X. guinotae, X. iranjae and X. longispinosus; see Apel & Spiridonov, 1998; Spiridonov, 1999, 2016) and lower subtidal species (e. g. X. tuerkayi from the western Indian Ocean; Spiridonov, 2016) occurring mostly on sandy substrates. Geographical range: Indo-West Pacific: from the Red Sea and the western Indian Ocean to Japan, Australia, the Hawaiian Islands and French Polynesia (Stephenson, 1972 a, b; Poupin, 2010; Castro, 2011).	en	Koch, Milan, Spiridonov, Vassily A., Ďuriš, Zdeněk (2023): Revision of the generic system for the swimming crab subfamily Portuninae (Decapoda: Brachyura: Portunidae) based on molecular and morphological analyses. Zoological Journal of the Linnean Society 197: 127-175
