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FIGURE 5 in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 5. Original illustrations of three described species of Rhabdopleura characterized by direct-frontal inception of ringed erect tubes (A‒C). Erect-tube inception in R. compacta (D‒F) appears to be a form of indirect erect-tube inception. Note that the direction of zooid budding is the same in A to C. A, Rhabdopleura mirabilis (from Sars 1872, pl. 1, fig. 5). B, Rhabdopleura annulata (from Norman 1921, fig. 4). C, Rhabdopleura recondita (after Beli et al. 2018, fig. 2C). D‒F, Rhabdopleura compacta (respectively after Hincks 1880, pl. 72, fig. 8; Stebbing 1970b, fig. 1; Stebbing 1970a, fig. 3).
FIGURE 6. Rhabdopleura grimaldii Jullien, 1890 in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 6. Rhabdopleura grimaldii Jullien, 1890 (A‒D, syntype MNHN-IB-2014-386) and R. manubialis Jullien & Calvet, 1903 (E, syntype MNHN-IB-2014-387). A, B, oblique views of two erect ringed tubes. C, adherent proximal part of a side branch (apsb), with a tapering pectocaulus (tp) (its side walls appearing as converging brown lines) and the broken base of an erect tube (bbet). D, similar to C, but also showing the creeping tube (ct) from which the broader side branch originated; note the faint outlines of oblique sutures on the creeping tube and side branch. E, erect ringed tube. Scale bars: A, B, 100 μm; C‒E, 200 μm. Images cropped from photos supplied by Pierre Lozuet, MNHN, Paris.
FIGURE 2. Interpretative images illustrating morphological terms and measured characters. A in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 2. Interpretative images illustrating morphological terms and measured characters. A, part of creeping tube (ct) of an unidentified Rhabdopleura (NIWA 158518) from Kermadec Ridge, with flat lateral margin (flm), faint oblique fusellar sutures and two ringed erect tubes (et) produced directly from the frontal surface of a creeping tube. B, distal end of an erect tube of Rhabdopleura francesca n. sp. (NIWA 158157), showing a vertical series of ring-like fuselli (f), the rim of each comprising a curled fusellar collar (fc). Tube diameter (td) is measured between fusellar collars; fusellus height (fh) is the distance between the bases of a pair of fusellar collars. C, transmitted-light image of part of a principal ('creeping') tube of R. emancipata n. sp. (NIWA 161211) showing a young zooid with contracted arm tentacles (at), stomach (st), rectum (r) and cephalic shield (cs), part of which is concealed by the reddish-brown stolon (black stolon or pectocaulus). D, aperture of an erect tube of R. francesca n. sp.; note the smooth-surfaced interior devoid of vertical fusellar fibrils. E, part of a creeping tube of an unidentified Rhabdopleura (NIWA 90267) from Cavalli Seamount with frontal fusellar sutures (fs, two arrowed). The white lines show the zigzag (zz) portion of the sutures (note the proximalwards shift to the left), with zigzags rendered as straight lines between points of intersection, regardless of suture curvature, to obtain measured angles (one example shown). F, close-up of part of R. francesca n. sp. (NIWA 158157) showing chains of dark 'dormant buds' (db) lying upon short stretches of the black stolon (bs); walls of enveloping creeping tubes almost invisible in image.
FIGURE 1 in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 1. Map showing the localities of four new Rhabdopleura taxa from New Zealand. From north to south these are R. emancipata n. sp., R. francesca n. sp., R. chathamica n. sp. and R. decipula n. sp. The boundary of the Extended Continental Shelf is shown surrounding New Zealand. The grey contour lines show the 1000 m depth interval. The map was generated using ArcGIS Pro.
FIGURE 8. Rhabdopleura francesca n in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 8. Rhabdopleura francesca n. sp., A‒E, holotype NIWA 158157; F‒H, paratype NIWA 162577: A, complete erect tube 2.5 mm long. B, holotype, possibly comprising more than one colony, on broken scallop shell. C, close-up of B showing that the black tubarium tracery in A is accentuated by continuous series of what look like dormant buds lying along black stolons. D, section of tube bracketed in A. E, apertural end of another erect tube. F, black stolon emergent from a 'dormant body' in part of a creeping tube. G, H, variations in creeping-tube width and fusellar patterning, the latter typically weak and thinly developed throughout colony; some point-to-point suture zigzags highlighted. Scale bars: A, 1 mm; B, 2 cm; C, 3 mm; D‒F, 100 μm; g, h, 200 μm.
FIGURE 13 in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 13. Epibionts of Rhabdopleura emancipata n. sp.: A, at top, an ancestrula of Chaperia sp. (Bryozoa) and, at bottom, a juvenile of the homotrematid foraminiferan Sporadotrema cylindricum. B, Chaperiopsis n. sp. (Bryozoa). C, Celleporina n. sp. (Bryozoa). D, annelid tube (at) draped along a principal tube. E, Galeopsis brevissimus (Bryozoa). F, units of a chain-like unidentified epibiont on a principal tube. G, Strongylopora gracilis (Bryozoa). H, Microtylostylifer n. sp. (Porifera) at left and Tubulipora sp. (Bryozoa) at right. R = Rhabdopleura host. Scale bars: A, D‒G, 500 μm; B, C, H, 1 mm.
FIGURE 3 in Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa
FIGURE 3. Maximum likelihood phylogeny for Rhabdopleura reconstructed on: A, concatenated 16S+18S rRNA genes; B, 13 mitochondrial protein-coding genes. Numbers adjacent to nodes represent bootstrap support (≥ 50) /posterior probability (≥0.8) values.
FIGURE 4 in A revision of the genus Saccoglossus (Hemichordata: Enteropneusta: Harrimaniidae) with taxonomic descriptions of five new species from the Eastern Pacific
FIGURE 4. Light micrographs of transverse sections of Saccoglossus shumaginensis n. sp.: A, Proboscis. B, Proboscis with heart-kidney complex. C, Proboscis neck. D, Anterior region of the collar. E, Posterior region of the collar. Inset: Collar canal. F, Anterior pharyngeal region of the trunk. bs, branchial sac; bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; g, glomerulus; gb, gill bar; go, gonad; nc, nerve cord; pc, proboscis coelom; phd, periheamal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pv, proboscis vesicle; s, stomochord; sc, skeletal cornua; sp, skeletal plate; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars A, B, D, E, F = 500 µm; C = 200 µm; E inset = 100 µm.
FIGURE 1 in A revision of the genus Saccoglossus (Hemichordata: Enteropneusta: Harrimaniidae) with taxonomic descriptions of five new species from the Eastern Pacific
FIGURE 1. Photographs of fixed specimens from the NMNH (Smithsonian). With the exception of A and F, all of the specimens have lost the anterior end of the proboscis. A, Photograph of a coiled specimen of Saccoglossus porochordus n. sp. showing the proboscis and dorsal side. B, Photograph of the ventral side of S. porochordus n. sp. C, Photograph of the dorsal side of S. rhabdorhynchus n. sp. D, Photograph of the lateral side of S. rhabdorhynchus n. sp. E, Photograph of the right side of S. shumaginensis n. sp. F, Photograph of the left side of S. shumaginensis n. sp. G, Photograph of the right side of S. sonorensis n. sp. H, Photograph of the left side of S. sonorensis n. sp. c, collar; p, proboscis; t, trunk. Scale bars = 5 mm.
FIGURE 5 in A revision of the genus Saccoglossus (Hemichordata: Enteropneusta: Harrimaniidae) with taxonomic descriptions of five new species from the Eastern Pacific
FIGURE 5. Light micrographs of transverse sections of Saccoglossus sonorensis n. sp.: A, Proboscis with heart-kidney complex. Inset: Anterior region of the proboscis coelom. B, Proboscis neck showing the skeletal spine of the proboscis skeleton. C, Proboscis neck showing the proboscis vesicle. D, Anterior region of the collar. Inset: Collar nerve cord showing a nerve root. E, Posterior region of the collar. Inset: Collar canal. F, Anterior pharyngeal region of the trunk. ac, amoeboid cells; bp, branchial pore; bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; e, epithelia; g, glomerulus; gb, gill bar; go, gonad; nc, nerve cord; nr, nerve root; pc, proboscis coelom; phd, periheamal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pv, proboscis vesicle; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sl, stomochord lumen; sp, skeletal plate; ss, skeletal spine; tlm, trunk longitudinal muscles. Scale bars A, E, F, G = 500 µm; B and F inset = 100 µm; C, D, E inset = 200 µm.
FIGURE 3 in A revision of the genus Saccoglossus (Hemichordata: Enteropneusta: Harrimaniidae) with taxonomic descriptions of five new species from the Eastern Pacific
FIGURE 3. Light micrographs of transverse sections of Saccoglossus rhabdorhynchus n. sp.: A, Proboscis with heartkidney complex. B, Proboscis neck showing the spine of the proboscis skeleton. C, Proboscis neck showing the proboscis vesicle. D, Anterior region of the collar. E, Posterior region of the collar. Inset: Collar canal. F, Anterior pharyngeal region of the trunk. bp, branchial pore; bs, branchial sac; bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; nc, nerve cord; pc, proboscis coelom; np, neuropore; phd, periheamal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pv, proboscis vesicle; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sl, stomochord lumen; sp, skeletal plate; ss, skeletal spine; tl, trunk lumen; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars A, B, C, D, E, F = 500 µm; E inset = 100 µm.
FIGURE 6 in A revision of the genus Saccoglossus (Hemichordata: Enteropneusta: Harrimaniidae) with taxonomic descriptions of five new species from the Eastern Pacific
FIGURE 6. Light micrographs of transverse sections of Saccoglossus palmeri n. sp.: A, Anterior region of the proboscis. B, Proboscis with heart-kidney complex. C, Proboscis neck. D, Anterior region of the collar. E, Anterior pharyngeal region of the trunk. bv, blood vessel; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; go, gonad; nc, nerve cord; pc, proboscis coelom; phd, periheamal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pv, proboscis vesicle; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sp, skeletal plate; tlm, trunk longitudinal muscles. Scale bars A, B, D, E = 250 µm; C = 150 µm.
FIGURE 2 in A revision of the genus Saccoglossus (Hemichordata: Enteropneusta: Harrimaniidae) with taxonomic descriptions of five new species from the Eastern Pacific
FIGURE 2. Light micrographs of transverse sections of Saccoglossus porochordus n. sp.: A, Proboscis with heartkidney complex. B, Heart-kidney complex. C, Proboscis neck. D, Anterior region of the collar. Inset: Dorsal collar nerve cord and crest. E, Posterior region of the collar. F, Collar canal. G, Dorsal groove of the collar. H, Anterior pharyngeal region of the trunk. a, axons; bs, branchial sac; bv, blood vessel; cb, cell bodies; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; dg, dorsal groove; g, glomerulus; gb, gill bar; go, gonad; mc, mucous cells; nc, nerve cord; ncc, nerve cord crest; pc, proboscis coelom; phd, periheamal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pp, proboscis pore; pv, proboscis vesicle; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sp, skeletal plate; ss, skeletal spine; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars A, D and F = 750 µm; B and C = 300 µm; D inset and E inset = 200 µm.
FIGURE 11. A in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 11. A phylogenetic hypothesis for the family Harrimaniidae that was constructed using the tree of Cannon et al. (2009) as a backbone, to which were added the new genera Horstia, Ritteria and Mesoglossus. A, Proboscis longitudinal musculature in radial bundles. B, Direct development, absence of branchial synapticules, hepatic caeca, genital wings, trunk lateral septa, parabranchial ridges and circular muscles in the trunk. C, Four tubular genital ridges restricted to the branchial region, reduced proboscis canals and pores. D, No individual character is known to be unique to this group. E, Dorsal and ventral proboscis septa. F, Parabuccal ridges. G, Prominent dorsal proboscis groove, proboscis short and conical, anterior edge of the collar is ruffled, anterior and posterior neuropores, parabuccal ridges extend to the posterior part of the collar, dorsal and ventral septa in the proboscis and collar. H, Collar broader than long, no peribuccal diverticula, proboscis conical, a little longer than broad. I, No genital ridges, gill pores on an elevated ridge, proboscis round and as long as broad, trunk very narrow. J, Loss of radial proboscis longitudinal musculature. K, Proboscis longitudinal musculature is diffuse. L, Live in deep sea, coronate proboscis skeleton. M, Absence of proboscis neck, two pairs of genital ridges, proboscis round, proboscis and collar broader than long, proboscis pore opens into the neuropore. N, Collar pores, cardiac vesicle attached to dorsal the wall of the proboscis, externally visible gonopores, gill pores not visible externally. O, Proboscis at least twice as long as broad, diffuse proboscis longitudinal musculature. P, Proboscis at least twice as long as broad, proboscis longitudinal musculature in concentric rings.
FIGURE 10. A in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 10. A map of the geographic distribution of harrimaniid enteropneusts on the west coast of North America: 1, Stereobalanus willeyi n. sp. Newport Bay, CA. 2, Stereobalanus canadensis San Diego, CA (also found in the Lower Estuary and Gulf of St. Lawrence, including east of Prince Edward Island; New England). 3, Ritteria ambigua n. gen. et n. sp. Monterey, CA. 4, Horstia kincaidi n. gen. et n. sp. Whidbey Island, WA. 5, Saccoglossus pusillus Barkley Sound, BC; Cape Arago, OR; San Pedro, San Diego, La Jolla, Anaheim, Newport Bay, CA; Ensenada, Mexico. 6, Saccoglossus bromophenolosus Padilla Bay, Willapa Bay, WA. 7, Mesoglossus intermedius n. gen. et n. sp. Shelter Cove, Moss Beach, CA. 8, Mesoglossus macginitiei n. sp. Newport Bay, CA. 9, Harrimania maculosa Prince William Sound, Kodiak, AK; Providence Bay, Siberia. 10, Harrimania planktophilus Barkley Sound, BC. 11, Protoglossus mackiei n. sp. Moss Beach, CA.
FIGURE 8 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 8. Light micrographs of transverse sections of Ritteria ambigua: A, Proboscis with heart-kidney complex. B, Junction of the proboscis and collar. C, Anterior region of the collar. D, Anterior pharyngeal region of the trunk. E, Genital region of the trunk. F, Proboscis skeleton, neuropore and proboscis pore. G, Collar canal. H, Collar nerve cord showing the nerve root. Abbreviations: bs, branchial sac; bv; blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; e, epithelia; g, glomerulus; gb, gill bar; go, gonad; m, mouth; nc, nerve cord; nf, nerve fiber layer; np, neuropore; nr, nerve root; pbr, parabuccal ridge; pc, proboscis coelom; phd, perihaemal diverticulum; plm, proboscis longitudinal muscles; pp, proboscis pore; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sp, skeletal plate; tlm, trunk longitudinal muscles. Scale bars: A, B, C, D, E = 500 µm; F, G, H = 200 µm.
FIGURE 9 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 9. Light micrographs of transverse sections of Horstia kincaidi n. gen. et n. sp.: A, Proboscis with heartkidney complex. B, Proboscis neck. C, Collar. Inset: Collar nerve cord and stomochord. D, Anterior pharyngeal region of the trunk. Inset: Collar canal. E, Pharyngeal region of the trunk. F, Genital region of the trunk. Abbreviations: bp, branchial pore; bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; go, gonad; nc, nerve cord; np, neuropore; pc, proboscis coelom; phd, perihaemal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pv, proboscis vesicle; s, stomochord; sc, skeletal plate; sk, skeletal keel; sl, stomochord lumen; sp, skeletal plate; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars: A, C, E, G = 500 µm; B = 200 µm; D, F = 100 µm; H = 300 µm.
FIGURE 7 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 7. Light micrographs of transverse sections of Mesoglossus macginitiei n. gen. et n. sp.: A, Proboscis with heart–kidney complex. B, Proboscis coelomic cavity showing amoeboid-like cells. C, Proboscis neck. D, Anterior region of the collar showing the neuropore. E, Anterior region of the collar showing the dorsal nerve-cord crest. Inset: Dorsal nerve cord crest. F, Posterior region of the collar showing the peribuccal diverticula. G, Anterior pharyngeal region of the trunk. Inset: Collar canal. Abbreviations: ac, amoeboid-like cells; bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; go, gonad; nc, nerve cord; ncc, nerve cord crest; np, neuropore; pbd, peribuccal diverticula; pc, proboscis coelom; phd, perihaemal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; proboscis vesicle; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sp, skeletal plate; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars: A, D, G, H = 500 µm; B = 50 µm; C, I = 200 µm; F = 300 µm.
FIGURE 6 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 6. Light micrographs of transverse sections of Mesoglossus intermedius n. gen. et n. sp.: A, Proboscis with heart–kidney complex. B, Proboscis neck. C, Proboscis neck showing the neuropore. D, Collar. E, Posterior region of the collar, (inset) collar canal. F, Anterior pharyngeal region of the trunk. Abbreviations: bp, branchial pore; bv, blood vessel; cc, collar canal; clm, collar longitudinal muscles; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; nc, nerve cord; np, neuropore; pc; proboscis coelom; phd, perihaemal diverticulum; pl, pharynx lumen; plm, proboscis longitudinal muscles; pp, proboscis pore; s, stomochord; sc, skeletal cornua; sk, skeletal keel; sp, skeletal plate; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars: A, D, E, G = 500 µm; B = 100 µm; C, F = 200 µm.
FIGURE 4 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific
FIGURE 4. Light micrographs of transverse sections of Protoglossus mackiei n. sp.: A, Proboscis with heart–kidney complex. Inset: Proboscis complex. B, Proboscis neck. C, Proboscis neck showing the neuropore. D, Collar. Inset: Collar nerve cord. E, Anterior pharyngeal region of the trunk. Inset: Collar canal. F, Pharyngeal region of the trunk. Abbreviations: bp, branchial pore; bs, branchial sac; bv, blood vessel; cc, collar canal; clm, collar longitudinal muscles; cm, circular muscle layer; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; nc, nerve cord; ncl, nerve cord lacunae; nf, nerve fiber layer; np, neuropore; pbr, parabuccal ridge; pc, proboscis coelom; pl, pharynx lumen; plm, proboscis longitudinal muscles; phd, perihaemal diverticulum; pp, proboscis pore; pv, proboscis vesicle; s, stomochord, sc, skeletal cornua; sk, skeletal keel; sp, skeletal plate; tlm, trunk longitudinal muscles; vm, ventral mesentery. Scale bars: A, E, G, I = 500 µm; B, C, D, H = 100 µm; F = 300 µm.
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