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FIGURE 3. Metacrimenes fenestra n. gen., n in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 3. Metacrimenes fenestra n. gen., n. sp., holotype, ovigerous female (cl 3.2 mm), CBM-ZC 17530; A, anterior part of carapace and cephalic appendages, lateral view; B, same, dorsal view [marginal setae omitted on right side]; C, telson, dorsal view; D, close up of posterior part of telson, dorsal view; E, lateral flagellum of left antennule, dorsolateral view; F, left antenna, ventral view (setae omitted); G, left uropod, dorsal view (marginal setae omitted).

FIGURE 7. Host crinoid Metacrinus rotundus Carpenter, 1885 in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 7. Host crinoid Metacrinus rotundus Carpenter, 1885 in situ. A, entire animal on the muddy bottom; B, holotype of Metacrimenes fenestra n. gen., n. sp. clinging to the arm of M. rotundus (blue arrow).

FIGURE 2. Metacrimenes fenestra n. gen., n in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 2. Metacrimenes fenestra n. gen., n. sp., holotype, ovigerous female (cl 3.2 mm), CBM-ZC 17530; A, carapace and cephalic appendages, lateral view; B, pleon, telson and pleonal appendages, lateral view.

FIGURE 1. Metacrimenes fenestra n. gen., n in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 1. Metacrimenes fenestra n. gen., n. sp., holotype, ovigerous female (cl 3.2 mm), CBM-ZC 17530, photographed shortly after capture in small aquarium, showing living colouration; A, habitus, lateral view; B, habitus, dorsal view.

FIGURE 6. Metacrimenes fenestra n. gen., n in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 6. Metacrimenes fenestra n. gen., n. sp., holotype, ovigerous female (cl 3.2 mm), CBM-ZC 17530; A, chela of left first pereopod, extensor view; B, chela and carpus of left first pereopod, mesial view; C, chela of left second pereopod, extensor view; D, dactylus and distal part of third pereopod, lateral view; E, dactylus and distal part of fifth pereopod, mesial view.

FIGURE 8 in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 8. Maximum-likelihood (ML) tree constructed from concatenated data sets of the mitochondrial COI and 16S rRNA genes of Metacrimenes fenestra n. gen., n. sp. and closely allied palaemonid taxa (log likelihood -10197.759547). Numbers at nodes are bootstrap values; only values>80% are shown. The outgroup (Brachycarpus biunguiculatus) is omitted.

FIGURE 5. Metacrimenes fenestra n. gen., n in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 5. Metacrimenes fenestra n. gen., n. sp., holotype, ovigerous female (cl 3.2 mm), CBM-ZC 17530; pereopods, lateral view. A, first pereopod; B, left second pereopod, C, right second pereopod; D, left third pereopod; E, left fourth pereopod; F, left fifth pereopod.

FIGURE 4. Metacrimenes fenestra n. gen., n in A new genus and new species of palaemonid shrimp (Decapoda: Caridea), associated with the deep-sea crinoid Metacrinus rotundus (Echinodermata: Isocrinidae), from Suruga Bay, Japan

FIGURE 4. Metacrimenes fenestra n. gen., n. sp., holotype, ovigerous female (cl 3.2 mm), CBM-ZC 17530; A, mandible, inner view; B, maxillule, outer view; C, maxilla, outer view; D, first maxilliped, outer view (endites broken during dissection); E, second maxilliped, outer view; F, third maxilliped, lateral view; G, same, coxa and antepenultimate article, ventral view.

Figure 10 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 10. Cladograms based on analysis of four genes of extant Isocrinida. A, maximum likelihood cladogram with bootstrap support; B, Bayesian inference cladogram with posterior probabilities; C, summary of results showing the main clades discussed in the text with support values shown at the nodes, bootstrap and posterior probability separated by a slash bar; approximate periods of divergence are indicated at nodes retrieved in ML and BI reconstructions.

Figure 9 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 9. Early ontogeny of first primibrachial articulations in extant isocrinids. A, B, early post-larval specimen of Endoxocrinus wyvillethomsoni; A, distal facet; B, proximal facet; C, juvenile specimen of Metacrinus sp., distal facet. al, adoral ligament area; ap, aboral ligament pit; ar, adoral axial ridge; fr, fulcral ridge; m, muscle area; mn, muscle nerve canal; n, brachial nerve canal; r, rudimentary fulcral ridge. Scale bar 100 µm.

Figure 8 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 8. Distal stalk articulations of Panglaocrinus isseliformis gen. nov., sp. nov.. A, D, distal synostosial facet of nodal 7 (cryptosymplexy); B, distal–lateral view of the same nodal; C, proximal synostosial facet of infranodal following N7 (cryptosymplexy); E, F, symplexial facet of distal internodal.

Figure 7 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 7. External morphology and articulations of pinnule of Panglaocrinus isseliformis gen. nov., sp. nov.. A, general view; B, adoral–side view (cp: cover plates, lp: lateral plate); C, proximal–adoral view of Pn1; D, proximal–adoral of Pn2; E, proximal view of Pn3; F, proximal–aboral view of Pn5.

Figure 6 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 6. Articulations of secundibrachials of Panglaocrinus isseliformis gen. nov., sp. nov.. A, B, adoral view showing the pinnule socket; A, proximal–adoral view of IIBr4; B, distal–adoral view of IIBr5; C, proximal view of IIBr4; D, distal view of IIBr1; E, F, proximal view of IIBr12 (il: inner ligament area, m: muscular area).

Figure 5 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 5. Articulation facets of primibrachials of Panglaocrinus isseliformis gen. nov., sp. nov.. A, proximal–adoral view of IBr1 (muscular synarthry); B, distal-adoral view of IBr1 (synostosis, arrow: aboral ligament pit); C, D, distal muscular synarthries of IBr2ax, in distal and adoral view, respectively.

Figure 3 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 3. Anal cone of Panglaocrinus isseliformis gen. nov., sp. nov.. Arrow: IIBr2 bearing the first pinnule. C-ray and right secundibrachitaxis of D-ray removed.

Figure 12 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 12. Constraints to phylogenetic reconstruction and classification of stalked Isocrinida using molecular phylogeny, morphological characters and data from the fossil record. *Complete or at least proximal portion of crown known from the fossil record. The genus Isocrinus is here used 'sensu lato' (Isocrinus + Chladocrinus) as is Metacrinus (Metacrinus + Saracrinus); Cenocrinus is not mentioned because of its uncertain attribution to any of the subfamilies of Balanocrinidae. The Early Cretaceous genus Frasericrinus Ausich et al. 2015 is included within Balanocrinus 'sensu lato'. For the hypothetical attribution to Proisocrinidae of Angulocrinus and Millericrinus, reference is made to the text. Data derive from Hess (Hess 2011a, b), Améziane et al. (2021), and Roux and Philippe (2021). Colours used to differentiate Periods and Epochs are from the RGB Color Code according to the Commission for the Geological Map of the World (Gradstein et al. 2012).

Figure 1 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 1. The main morphological characters in extant Isocrinida. A, B, Metacrininae with long brachitaxes and abundant stout cirri; C, D, Diplocrininae with short brachitaxes and moderately developed cirri; E, F, Proisocrininae with well individualized aboral cup, short brachitaxes, and homeomorphic stalk with rudimentary cirri restricted proximally. A, large Saracrinus sp. on Tasmanian Seamount, courtesy of A. William, CSIRO; B, Metacrinus interruptus Carpenter, 1884 from Taiwan slope; C, Endoxocrinus parrae(Gervais in Guérin 1835) on Bahamas slope, courtesy of J.S. Miller, HBOI; D, E. alternicirrus Carpenter, 1882 from New Caledonia slope; E, F, Proisocrinus ruberrimus A.H.Clark, 1910; E, on Lifou slope (New Caledonia); F, from Tahiti slope.

Figure 4 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 4. Arm and stalk morphology of Panglaocrinus isseliformis gen. nov., sp. nov.. A, distal part of secundibrachitaxis with IIBr15ax. (arrow); B, distal part of proxistele with rudimentary cirrus beginning its growth in length; C, segment of distal stalk with the longest cirrus and maximum columnal height. N, nodal.

Figure 2 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 2. Crown morphology of Panglaocrinus isseliformis gen. nov., sp. nov.. A, general view (arrows: IIBrax); B, aboral cup and proximal arms (*arm trunk regenerating from IBrax upwards); C, inner view of crown (arrow: anal cone, C-ray removed).

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Figure 11 in Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific

Figure 11. Ancestry of the main morphological characters of Isocrinida. The list of characters taken into account is that of those identified in extant isocrinids. Abbreviations: Balanocr, balanicrinine type; Iso, isocrinine type; Diplo, diplocrinine type; No Cri, absence of inner crenularium; Distal, oriented downwards; Altern, alternicirration; Emb S, embayed synarthry; Syno2, secondary synostosis (cryptosynarthry); Sympl, marginal symplexial crenularium; D sm, deep symmorphy; Syno1, primary synostosis. Millericrinus and Angulocrinus are indicated for comparison with Proisocrinus (see text).