FIGURE 1. Phylogenetic relationships among 38 meiuran families (36 Brachyura in ingroup), as represented by their oldest reliable vetted fossil species occurrences to date and their age brackets, used as calibration points for node dated molecular divergence time estimations of brachyuran crabs in Wolfe et al. (2023). Fossil species used for node dating are here represented as terminals. Base topology and full maximum clade credibility divergence time estimates (thin shaded yellow bars, representing 95% HPD) after Wolfe et al. (2023). Main brachyuran sections, and the family and superfamily ingroups focused on in this study, coloured as follows: Dromiacea (red), Homoloida (orange), Raninoida (yellow), Cyclodorippoida (green), heterotreme Eubrachyura (blue), and thoracotreme Eubrachyura (purple). Oldest crown Porcellanidae (Anomura, brown) and the extinct Eocarcinidae (black dotted line), as non-crown brachyuran outgroups. Fossil age ranges (thick dark grey bars) denote the maximum (left end) and minimum (right end) age brackets for the selected calibration points, based on available chronostratigraphic information derived from isotopic, magnetostratigraphic, or bio-stratigraphic data (see main text). Longer grey bars (e.g., Calappa zinsmeister) indicate a wider age gap between the tip maximum and minimum ages for a given taxon, and thus less age resolution for the relative age of the fossil, whereas thinner grey bars (e.g., pseudothelphusid sp.) indicate a narrower age gap between their tip maximum and minimum ages, and thus more resolution and preciseness of the relative age of the fossil. Phylogenetic placement of Joeranina kerri (Raninoida: Palaeocorystoidea: Palaeocorystidae) (yellow dotted line) after Luque et al. (2019a) and of Cretapsara athanata (Eubrachyura: Cretapsaridae) and Eogeryon elegius (Eubrachyura: Portunoidea: Eogeryonidae) (blue dotted lines) after Luque et al. (2021).
FIGURE 2. Meiura: A: Eocarcinus praecursor Withers, 1932, holotype NHM 18425, Lower Jurassic (Pliensbachian), UK. B: Anomura: B: Vibrissalana jurassica Robins and Klompmaker, 2019, holotype NHMW 2007z0149/0405, Upper Jurassic (Tithonian), Austria. C-G: Brachyura; C: Dromiacea: Dromioidea: Graptocarcinus muiri Stenzel, 1944, holotype BEG-21288 (=BEG00021288.000), Lower Cretaceous (Albian), San Luis Potosí, Mexico; D: Homoloida: Homoloidea: Homolidae: Doerflesia ornata Feldmann and Schweitzer, 2009, holotype 2007z0149/0015, uppermost Lower Cretaceous (upper Albian), Texas, USA; E-F: Raninoida; E: Palaeocorystoidea: Palaeocorystidae: Joeranina kerri (Luque, Feldmann, Schweitzer, Jaramillo, and Cameron, 2012, as Notopocorystes), holotype IGM p881128, upper Lower Cretaceous (Aptian), Colombia; F: Raninoidea: Lyreididae: Marylyreidus punctatus (Rathbun, 1935, as Notopocorystes), non-type specimen USNM 559038 Texas, USA; G: Cyclodorippoida: Cyclodorippoidea: Cymonomidae: Cymonomus primitivus Müller and Collins, 1991, holotype [HNHM M.91-135], Eocene (Priabonian), Budapest, Hungary. Photos by: Javier Luque (A, E, and F), Adiël Klompmaker (B), Ann Molineux, courtesy of Lisa Boucher and Adiël Klompmaker (C), Rodney Feldmann (D), and Matus Hyžný (G).
FIGURE 3. Crown Eubrachyura. A: Superfamily uncertain: Cretapsaridae: Cretapsara athanata Luque in Luque et al., 2021, holotype LYAM-9, lowermost Upper Cretaceous (lower Cenomanian), Burma. B: Potamoidea: Potamidae: Alontecarcinus buratoi De Angeli and Caporiondo, 2019, holotype IGVR 19.38, Eocene (Bartonian), Vicenza, Italy. C-J: Eubrachyura: Thoracotremana; C: Ocypodoidea: Ocypodidae: Afruca miocenica (Artal, 2008, as Uca), holotype MGSB 68653, Miocene (Langhian), Catalonia, NE Iberian Peninsula. D: Pinnotheroidea: Pinnotheridae: Pinnixa sp., specimen UF 115397, lower Miocene (Burdigalian), Panama Canal, Panama (Luque et al., 2017). E-J: Grapsoidea; E,F: Percnidae; E: Percnon santurbanensis Ceccon and De Angeli, 2019, holotype MCV.19/02, lower Oligocene (Rupelian), Vicenza, Italy; F: Percnon paleogenicus De Angeli, 2023. holotype MCV.23/738-22.341, upper Eocene (Priabonian) Vicenza, Italy. G: Grapsidae: Pachygrapsus hungaricus Müller, 1974, holotype HNHM 2004.163.1, middle Miocene (Badenian), Budapest area, Hungary. H: Varunidae: Brachynotus corallinus Beschin, Busulini, De Angeli and Tessier, 2007, holotype MCZ 1794, lower Eocene (Ypresian), Vicenza, Italy. I: Gecarcinidae: Cardisoma guanhumi Latreille, in Latreille, Le Peletier, Serville and Guérin, 1828, specimen USNM 618301, Pleistocene, Bermuda. J: Sesarmidae indet., specimen IHNFG-4991, lower Miocene (Aquitanian) of Chiapas, Mexico (Serrano-Sánchez et al., 2016). Photos by: Lida Xing (A), Antonio De Angeli (B, E, F), Pedro Artal (C), Javier Luque (D, I), Matúš Hyžný (G), Alessandra Busulini (H), and Francisco Vega (J).
FIGURE 4. Crown Eubrachyura, cont. A: Pseudothelphusioidea: Pseudothelphusidae indet., specimen UF 354202, Early Miocene (Burdigalian), Panama Canal, Panama (Luque et al., 2019b). B: Trichodactyloidea: Trichodactylidae indet., specimen CTA 47, Eocene (Barrancan), Contamana, Peru. C: Majoidea: Epialtidae + Mithracidae: Micippa antiqua Beschin, De Angeli, and Checchi, 2001, holotype I.G. 286477, lower Oligocene (Rupelian), Vicenza, Italy. D-G: Portunoidea; D: Eogeryonidae: Eogeryon elegius Ossó, 2021, holotype MGB 69151, lower Upper Cretaceous (Cenomanian), Northern Guadalajara Province, Spain (Ossó, 2016); E: Geryonidae: Chaceon helmstedtense (Bachmayer and Mundlos, 1968) (as Coeloma ?), holotype NHMW 1968/0773/0002, lower Oligocene (Rupelian), Northwestern Germany; F: Portunidae: Thalamitinae: Lessinithalamita gioiae De Angeli and Ceccon, 2015, holotype MCV14/15, lower Eocene (Ypresian), Vicenza, Italy; G: Polybiidae: Liocarcinus heintzi Schweitzer and Feldmann, 2010b, holotype MNHN R03778, lower Oligocene (Rupelian), Western Aquitaine, France. H: Parthenopoidea: Parthenopidae + Dairoididae: Aragolambrus collinsi Ferratges, Zamora, and Aurell, 2019, holotype MPZ-2019/211, lower Eocene (Ypresian-Ilerdian), Huesca province, Spain. I: Calappoidea: Calappidae: Calappa zinsmeisteri Feldmann and Wilson, 1988, holotype USNM 404877, upper Eocene, Seymour Island, Antarctica. J: Cancroidea: Cancridae: Anatolikos undecimspinosus Schweitzer, Feldmann, González-Barba, and Ćosović, 2006, holotype MHN-UABCS/Ba12-9, middle Eocene, Baja California Sur, Mexico. Photos by: Javier Luque (A), Pierre Olivier Antoine (B), Antonio De Angeli (C and F), Àlex Ossó (D), Barry van Bakel (E), Rodney Feldmann (G, I-K), and Fernando Ferratges (H).
FIGURE 5. Crown Eubrachyura, cont. A: Dorippoidea: Bartethusa hepatica Quayle and Collins, 1981, holotype BM In.61704, middle Eocene (Bartonian), Isle of Wight, England. B: Leucosioidea: Leucosiidae: Typilobus alponensis Beschin, De Angeli, and Zorzin, 2009, holotype IGVR78539, lower Eocene (Ypresian), Verona, Italy. C: Goneplacoidea: Euryplacidae: Chirinocarcinus wichmanni (Feldmann, Casadío, Chirino-Galvez, and Aguirre-Urreta, 1995) (as Glyphithyreus), holotype GHUNLPam 7015, lowermost Paleocene (lower Danian), Neuquén Province, Argentina. D-F: Eriphioidea; D-E: Eriphiidae: Eriphia verrucosa (Forskal, 1775) (as Cancer), in Betancort et al. (2014), specimen number indet., upper Miocene, Islas Canarias, Spain. F: Eriphia cocchi Ristori, 1886, specimen C-021-2.1, lower Pliocene (Zanclean-Piacentian), Italy. G: Trapezioidea: Trapeziidae: Archaeotetra lessinea De Angeli and Ceccon, 2013, holotype MCV 12/05-I.G.360311, lower Eocene (Ypresian), Monte Magré, Lessini Mounts, Italy. H: Eriphioidea: Oziidae: Ozius collinsi Karasawa, 1992, holotype MFM39001, lowermost Middle Miocene, Okayama Prefecture, Japan, Japan. I: Pilumnoidea: Pilumnidae: Glabropilumnus trispinosus Beschin, Busulini, and Tessier, in Beschin et al., 2016b, holotype VR 94508, lower Eocene (Ypresian), Verona, Italy. J: Goneplacoidea: Goneplacidae: Carcinoplax temikoensis Feldmann and Maxwell, 1990, holotype AR 1943, upper Eocene (Kaiatan or Runangan), Westland, New Zealand. K: Xanthoidea: Pseudorhombilidae: Pseudorhombila patagonica Glaessner, 1933, holotype In. 28031, Miocene indet., Santa Cruz Province, Argentina. L: Xanthoidea: Panopeidae: Panopeus incisus Beschin, Busulini, De Angeli, and Tessier, 2007, holotype MCZ 2009, lower Eocene, Vicenza, Italy. M: Xanthoidea: Xanthidae: Phlyctenodes edwardsi Beschin, Busulini, Tessier, and Zorzin, 2016b, holotype VR 94284, lower Eocene (Ypresian), Verona, Italy. Photos by: Richard Howard (A, K), Antonio De Angeli (B, G), Rodney Feldmann (C, J), Juan Francisco Betancort (D, E), Àlex Ossó (F), Hiroaki Karasawa (H), Alessandra Busulini (I, L, M).
