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Raninoidea. A-F, Lyreididae; A-C, Lysirude nitidus (=Lyreidus bairdii), USNM 66638; A, dorsal view of female; B, SEM image of small right eye in a stout podophthalmite; C, details of the cornea bearing hexagonal facets in hexagonal packing. D-F, Lysirude griffini, USNM 216726; D, dorsal view of male; E, SEM image of small right eye; C, details of the cornea bearing hexagonal facets in hexagonal packing. G-I, Raninidae: Cyrtorhininae: Cyrtorhina granulosa, MNHN-IU-2016-2020 (= MNHN-B16181); G, dorsal view of female; H, SEM image of small right eye; I, details of the cornea bearing hexagonal facets in hexagonal packing. J-L, Raninidae: Symethinae: Symethis sp., uncatalogued specimen; J, dorsal view of male; K, Confocal microscope image of small right eye showing the different shapes and sizes of facets through the cornea; L, details of the cornea bearing hexagonal facets in hexagonal packing. Figure by J. Luque.

Raninoidea. A-F, Lyreididae; A-C, Lysirude nitidus (=Lyreidus bairdii), USNM 66638; A, dorsal view of female; B, SEM image of small right eye in a stout podophthalmite; C, details of the cornea bearing hexagonal facets in hexagonal packing. D-F, Lysirude griffini, USNM 216726; D, dorsal view of male; E, SEM image of small right eye; C, details of the cornea bearing hexagonal facets in hexagonal packing. G-I, Raninidae: Cyrtorhininae: Cyrtorhina granulosa, MNHN-IU-2016-2020 (= MNHN-B16181); G, dorsal view of female; H, SEM image of small right eye; I, details of the cornea bearing hexagonal facets in hexagonal packing. J-L, Raninidae: Symethinae: Symethis sp., uncatalogued specimen; J, dorsal view of male; K, Confocal microscope image of small right eye showing the different shapes and sizes of facets through the cornea; L, details of the cornea bearing hexagonal facets in hexagonal packing. Figure by J. Luque.

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Image-forming compound eyes are such a valuable adaptation that similar visual systems have evolved independently across crustaceans. But if different compound eye types have evolved independently multiple times, how useful are eye structures and ommatidia morphology for resolving phylogenetic relationships? Crabs are ideal study organisms to explo...

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... nitidus (A. Milne-Edwards, 1880) (=Lyreidus bairdii) ( Fig. 8A-C) and Lysirude griffini 200! Goeke, 1985 ( Fig. 8D-F 8G-I), Symethis sp. (Fig. J-L), Cosmonotus grayi White, 1848 ( Fig. 9A-C), Notopus dorsipes (Linnaeus, 1758) ( Fig. 9D-F), Ranilia muricata H Milne Edwards, 1837 ( Fig. 9G-I), Ranina . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the ...
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... nitidus (A. Milne-Edwards, 1880) (=Lyreidus bairdii) ( Fig. 8A-C) and Lysirude griffini 200! Goeke, 1985 ( Fig. 8D-F 8G-I), Symethis sp. (Fig. J-L), Cosmonotus grayi White, 1848 ( Fig. 9A-C), Notopus dorsipes (Linnaeus, 1758) ( Fig. 9D-F), Ranilia muricata H Milne Edwards, 1837 ( Fig. 9G-I), Ranina . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to ...
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... nitidus (A. Milne-Edwards, 1880) (=Lyreidus bairdii) ( Fig. 8A-C) and Lysirude griffini 200! Goeke, 1985 ( Fig. 8D-F 8G-I), Symethis sp. (Fig. J-L), Cosmonotus grayi White, 1848 ( Fig. 9A-C), Notopus dorsipes (Linnaeus, 1758) ( Fig. 9D-F), Ranilia muricata H Milne Edwards, 1837 ( Fig. 9G-I), Ranina . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. ...
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... lyreidid crab Lyreidus nitidus ( Fig. 8A) has small sub-conical eyes resting in stout and much longer podophthalmite, nearly 66% larger than the corneal eye, and covered in fibrous setae ( Table 1). The studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly ...
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... lyreidid crab Lyreidus nitidus ( Fig. 8A) has small sub-conical eyes resting in stout and much longer podophthalmite, nearly 66% larger than the corneal eye, and covered in fibrous setae ( Table 1). The studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly twice as long as the corneal eye, and it is partly covered in small setae (Fig. 8E). Both eye and eyestalk are barely protected by a narrow orbit with one supraorbital fissure, and a short, 370! blunt, triangular ...
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... podophthalmite, nearly 66% larger than the corneal eye, and covered in fibrous setae ( Table 1). The studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly twice as long as the corneal eye, and it is partly covered in small setae (Fig. 8E). Both eye and eyestalk are barely protected by a narrow orbit with one supraorbital fissure, and a short, 370! blunt, triangular outer orbital spine directed anteriorly (Fig 8D). The cornea width is approximately 1.5% the length of the carapace, and it is constituted by a few hundred small hexagonal facets that are packed in hexagonal ...
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... studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly twice as long as the corneal eye, and it is partly covered in small setae (Fig. 8E). Both eye and eyestalk are barely protected by a narrow orbit with one supraorbital fissure, and a short, 370! blunt, triangular outer orbital spine directed anteriorly (Fig 8D). The cornea width is approximately 1.5% the length of the carapace, and it is constituted by a few hundred small hexagonal facets that are packed in hexagonal array, with an average facet diameter of 23 µm . ...
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... copyright holder for this preprint (which . http://dx.doi.org/10.1101/786087 doi: bioRxiv preprint first posted online Oct. 7, 2019; ( Fig. 8C; Table 1). Hexagonal facets in hexagonal array have been reported for Lyreidus tridentatus (see Scholtz and McLay, 2009), suggesting a shared lack of 'mirror' eyes among 375! crabs of the family ...
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... all the podotreme crabs studied, those of the family Raninidae have the broadest range of eye shapes, sizes, and orbital constructions. In the subfamily Cyrtorhininae, Cyrtorhina granulosa ( Fig. 8G) shows a considerable reduction of the corneal region compared with the rest of the eyestalk. Its cornea is sub-conical and dorsally truncated by an extension of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately ...
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... subfamily Cyrtorhininae, Cyrtorhina granulosa ( Fig. 8G) shows a considerable reduction of the corneal region compared with the rest of the eyestalk. Its cornea is sub-conical and dorsally truncated by an extension of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately 1.8% the length of the carapace, and it is constituted by small hexagonal facets in hexagonal array, with an average facet diameter of 35 µm ( Fig. 8H, I; Table 1 Fig. 8K, L; Table ...
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... truncated by an extension of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately 1.8% the length of the carapace, and it is constituted by small hexagonal facets in hexagonal array, with an average facet diameter of 35 µm ( Fig. 8H, I; Table 1 Fig. 8K, L; Table ...
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... of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately 1.8% the length of the carapace, and it is constituted by small hexagonal facets in hexagonal array, with an average facet diameter of 35 µm ( Fig. 8H, I; Table 1 Fig. 8K, L; Table ...
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... the substratum during the day, emerging at night to search for food (Skinner and Hill, 1986). Some taxa like Ranilia, Ranina, and 605! particularly Cosmonotus, have relatively large eyes covered in small facets of nearly the same size throughout the cornea, and have long podophthalmite eyestalks that can be held outside the sediment when buried (Fig. 8). Conversely, raninid crabs like Symethis (Fig. 7J) show an . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to display the preprint in ...
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... nitidus (A. Milne-Edwards, 1880) (=Lyreidus bairdii) ( Fig. 8A-C) and Lysirude griffini 200! Goeke, 1985 ( Fig. 8D-F 8G-I), Symethis sp. (Fig. J-L), Cosmonotus grayi White, 1848 ( Fig. 9A-C), Notopus dorsipes (Linnaeus, 1758) ( Fig. 9D-F), Ranilia muricata H Milne Edwards, 1837 ( Fig. 9G-I), Ranina . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the ...
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... nitidus (A. Milne-Edwards, 1880) (=Lyreidus bairdii) ( Fig. 8A-C) and Lysirude griffini 200! Goeke, 1985 ( Fig. 8D-F 8G-I), Symethis sp. (Fig. J-L), Cosmonotus grayi White, 1848 ( Fig. 9A-C), Notopus dorsipes (Linnaeus, 1758) ( Fig. 9D-F), Ranilia muricata H Milne Edwards, 1837 ( Fig. 9G-I), Ranina . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to ...
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... nitidus (A. Milne-Edwards, 1880) (=Lyreidus bairdii) ( Fig. 8A-C) and Lysirude griffini 200! Goeke, 1985 ( Fig. 8D-F 8G-I), Symethis sp. (Fig. J-L), Cosmonotus grayi White, 1848 ( Fig. 9A-C), Notopus dorsipes (Linnaeus, 1758) ( Fig. 9D-F), Ranilia muricata H Milne Edwards, 1837 ( Fig. 9G-I), Ranina . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. ...
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... lyreidid crab Lyreidus nitidus ( Fig. 8A) has small sub-conical eyes resting in stout and much longer podophthalmite, nearly 66% larger than the corneal eye, and covered in fibrous setae ( Table 1). The studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly ...
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... lyreidid crab Lyreidus nitidus ( Fig. 8A) has small sub-conical eyes resting in stout and much longer podophthalmite, nearly 66% larger than the corneal eye, and covered in fibrous setae ( Table 1). The studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly twice as long as the corneal eye, and it is partly covered in small setae (Fig. 8E). Both eye and eyestalk are barely protected by a narrow orbit with one supraorbital fissure, and a short, 370! blunt, triangular ...
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... podophthalmite, nearly 66% larger than the corneal eye, and covered in fibrous setae ( Table 1). The studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly twice as long as the corneal eye, and it is partly covered in small setae (Fig. 8E). Both eye and eyestalk are barely protected by a narrow orbit with one supraorbital fissure, and a short, 370! blunt, triangular outer orbital spine directed anteriorly (Fig 8D). The cornea width is approximately 1.5% the length of the carapace, and it is constituted by a few hundred small hexagonal facets that are packed in hexagonal ...
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... studied specimen of Lysirude griffini ( Fig. 8D) has even more reduced sub-conical eyes and a longer and broader podophthalmite than L. nitidus; the podophthalmite is nearly twice as long as the corneal eye, and it is partly covered in small setae (Fig. 8E). Both eye and eyestalk are barely protected by a narrow orbit with one supraorbital fissure, and a short, 370! blunt, triangular outer orbital spine directed anteriorly (Fig 8D). The cornea width is approximately 1.5% the length of the carapace, and it is constituted by a few hundred small hexagonal facets that are packed in hexagonal array, with an average facet diameter of 23 µm . ...
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... copyright holder for this preprint (which . http://dx.doi.org/10.1101/786087 doi: bioRxiv preprint first posted online Oct. 7, 2019; ( Fig. 8C; Table 1). Hexagonal facets in hexagonal array have been reported for Lyreidus tridentatus (see Scholtz and McLay, 2009), suggesting a shared lack of 'mirror' eyes among 375! crabs of the family ...
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... all the podotreme crabs studied, those of the family Raninidae have the broadest range of eye shapes, sizes, and orbital constructions. In the subfamily Cyrtorhininae, Cyrtorhina granulosa ( Fig. 8G) shows a considerable reduction of the corneal region compared with the rest of the eyestalk. Its cornea is sub-conical and dorsally truncated by an extension of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately ...
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... subfamily Cyrtorhininae, Cyrtorhina granulosa ( Fig. 8G) shows a considerable reduction of the corneal region compared with the rest of the eyestalk. Its cornea is sub-conical and dorsally truncated by an extension of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately 1.8% the length of the carapace, and it is constituted by small hexagonal facets in hexagonal array, with an average facet diameter of 35 µm ( Fig. 8H, I; Table 1 Fig. 8K, L; Table ...
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... truncated by an extension of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately 1.8% the length of the carapace, and it is constituted by small hexagonal facets in hexagonal array, with an average facet diameter of 35 µm ( Fig. 8H, I; Table 1 Fig. 8K, L; Table ...
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... of the cuticle of 380! the podophthalmite that extends towards the pole of the eye, further reducing the area occupied by the cornea (Fig. 8H). The cornea width is approximately 1.8% the length of the carapace, and it is constituted by small hexagonal facets in hexagonal array, with an average facet diameter of 35 µm ( Fig. 8H, I; Table 1 Fig. 8K, L; Table ...
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... the substratum during the day, emerging at night to search for food (Skinner and Hill, 1986). Some taxa like Ranilia, Ranina, and 605! particularly Cosmonotus, have relatively large eyes covered in small facets of nearly the same size throughout the cornea, and have long podophthalmite eyestalks that can be held outside the sediment when buried (Fig. 8). Conversely, raninid crabs like Symethis (Fig. 7J) show an . CC-BY-NC-ND 4.0 International license It is made available under a was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to display the preprint in ...

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... However, the compound eyes of slipper lobsters with their superposition optical structures may improve vision under dimlight conditions (reviewed in Meyer-Rochow, 2001;Cronin and Porter, 2008). Such refracting superposition eyes have also been reported for other benthic decapods in deep-water systems, such as the crab superfamilies Homolodromioidea, Dromioidea, and Homoloidea (Luque et al., 2019), and this type of eye is associated with some nocturnal arthropods (Meyer-Rochow and Gál, 2004). Given their possession of these specialized eye structures, we infer that these slipper lobsters have excellent dim-light vision, which would assist them in their nocturnal activity patterns Goldstein et al., 2015) and facilitate adaptation to habitats of different depths. ...
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Amber fossils provide snapshots of the anatomy, biology, and ecology of extinct organisms that are otherwise inaccessible. The best-known fossils in amber are terrestrial arthropods—principally insects—whereas aquatic organisms are rarely represented. Here we present the first record of true crabs (Brachyura) in amber—from the Cretaceous of Myanmar (~100–99 Ma). The new fossil preserves large compound eyes, delicate mouthparts, and even gills. This modern-looking crab is nested within crown Eubrachyura, or ‘higher’ true crabs, which includes the majority of brachyuran species living today. The fossil appears to have been trapped in a brackish or freshwater setting near a coastal to fluvio-estuarine environment, bridging the gap between the predicted molecular divergence of non-marine crabs (~130 Ma) and their younger fossil record (latest Cretaceous and Paleogene, ~75 to 50 Ma) while providing a reliable calibration point for molecular divergence time estimates for ‘higher’ crown eubrachyurans.
... Anatomically, these crabs lie in between Anomura and Eubrachyura, and all current data strongly support a paraphyletic podotreme grade with brachyuran affinity. [31] Analysis of eight Sanger sequenced genes including 58 of ∼100 brachyuran families, [27] analyses of mitogenomes, [30,32] and a recent transcriptomic analysis [33] each recovered podotreme paraphyly (the former with weak support). Relationships recovered among podotremes were entirely contradictory between those analyses. ...
... Meanwhile, morphological trees, including fossils, have sampled more extensively from podotreme lineages. [2,31] Thus, a major goal of future research should represent all meiuran families with morphological data, and all extant families with strongly supported phylogenomic data, for a well-resolved total evidence phylogeny. ...
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A fundamental question in biology is whether phenotypes can be predicted by ecological or genomic rules. At least five cases of convergent evolution of the crab-like body plan (with a wide and flattened shape, and a bent abdomen) are known in decapod crustaceans, and have, for over 140 years, been known as "carcinization." The repeated loss of this body plan has been identified as "decarcinization." In reviewing the field, we offer phylogenetic strategies to include poorly known groups, and direct evidence from fossils, that will resolve the history of crab evolution and the degree of pheno-typic variation within crabs. Proposed ecological advantages of the crab body are summarized into a hypothesis of phenotypic integration suggesting correlated evolution of the carapace shape and abdomen. Our premise provides fertile ground for future studies of the genomic and developmental basis, and the predictability, of the crab-like body form.
... As of yet, molecular phylogenetics has been insufficient to resolve the puzzle of podotremes, therefore our depiction of their extant relationships in Figure 2 relies on morphological data. Anatomically, these crabs lie in between Anomura and Eubrachyura, though all current data strongly support a paraphyletic podotreme grade with brachyuran affinity (Luque et al. 2019a). Analysis of eight Sanger sequenced genes including 58 of ~100 brachyuran families (Tsang et al. 2014), analyses of mitogenomes (Tan et al. 2018b(Tan et al. , 2019, and a recent transcriptomic analysis ) each recovered podotreme paraphyly (the former with weak support). ...
... Of 11 extant podotreme families, however, over one third lack molecular data: no sequences have been published for Poupiniidae, Lyreididae, and Phyllotymolinidae, and only a single 18S sequence is available for Homolodromiidae. Meanwhile, morphological trees, including fossils, have sampled more extensively from podotreme lineages (Luque et al. 2019a(Luque et al. , 2019b. Thus, a major goal of future research should represent all meiuran families with morphological data, and all extant families with strongly supported phylogenomic data, for a well-resolved total evidence phylogeny. ...
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A fundamental question in biology is whether phenotypes can be predicted by ecological or genomic rules. For over 140 years, convergent evolution of the crab-like body plan (with a wide and flattened shape, and a bent abdomen) at least five times in decapod crustaceans has been known as 'carcinization'. The repeated loss of this body plan has been identified as 'decarcinization'. We offer phylogenetic strategies to include poorly known groups, and direct evidence from fossils, that will resolve the pattern of crab evolution and the degree of phenotypic variation within crabs. Proposed ecological advantages of the crab body are summarized into a hypothesis of phenotypic integration suggesting correlated evolution of the carapace shape and abdomen. Our premise provides fertile ground for future studies of the genomic and developmental basis, and the predictability, of the crab-like body form.
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