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Fig. 1 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig. 1. Inner secondary deposits in zooidal tubes of fossil Rhabdopleuroidea. A–C. Rhabdopleura kozlowskii Kulicki, 1969; Bathonian and Callovian of Poland. After Kulicki 1971, modified; A, ZPAL Pb.II/8; B, ZPAL Pb.II/9; C, ZPAL Pb.II/10. D. Rhabdopleurites primaevus Kozłowski, 1967; Lower Ordovician of Öland, Sweden. After Skevington 1965, modified.

Fig.7. Rhabdopleura normani Allman,1869 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig.7. Rhabdopleura normani Allman,1869,Recent(ZPALPb.5/4).SEMmicrographs. A.Fragmentofcoeneciumcomposedofanerecttubeconnected withabranchedcreepingtube. B.Connectionofcreepingtubewitherecttube. C.Erecttube. D.Outersurfaceofcreepingtubedevoidofpellicle. E.Details ofbrokenperidermshowingarrangementoffusellarfibrils. F.Shortverticalfibrilsonperidermoferecttube. G.Peridermofthesameerecttubedevoidof verticalwrinkles.Abbreviations:b,band−likestructurecomposedofcorticalfibrils;c,creepingtube;co,fusellarcollar;e,erecttube;ff,fusellarfibrils;m, foreign material; o, oblique suture; v, wrinkle.

Fig. 4. Rhabdopleura compacta Hincks, 1880 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig. 4. Rhabdopleura compacta Hincks, 1880, Recent (ZPALPb.5/1). SEM micrographs. Micromorphology of outer surface of creeping tubes. A. Fragmentofcreepingtubewithprominentzig−zagsuture. B–D.Corticalfibrilsinfusellarcollars.Abbreviations:c,fusellarcollar;cf,corticalfibril;fm,foreign material; p, fusellar pellicle.

Fig. 6. Rhabdopleura compacta Hincks, 1880 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig. 6. Rhabdopleura compacta Hincks, 1880, Recent (ZPALPb.5/3). SEM micrographs. Secondary inner deposits in erect tubes. A. Growing end of an erect tube with inner cortical deposits. B. Detail of Fig. 5A showing an arrangement of cortical fibrils. C. Broken erect tube near its base showing membraneousinnerdeposits. D.Multilayeredstructureofinnermembraneousdeposits.Abbreviations:cf,corticalfibril;i,interfusellarsuture;id,innerdeposit; l, last fusellus; n, next to the last fusellus; o, outer edge of an erect tube.

Fig.8. Rhabdopleura kozlowskii Kulicki,1969 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig.8. Rhabdopleura kozlowskii Kulicki,1969;CallovianofPoland(ZPAL Pb.5/5). SEM micrographs. A. Fragment of erect tube with inner deposits. B.Fibrillarnatureoftheinnerdepositmaterial. C.Pitsinthefusellartissue. Abbreviations: f, fusellar layer; s, secondary inner deposits.

Fig.3. Rhabdopleura compacta Hincks,1888 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig.3. Rhabdopleura compacta Hincks,1888,Recent(ZPALPb.5/1).Surface micromorphology of prosicula. SEM micrographs. A. Distal part of prosicula connected with proximal part of metasicula. B. Outer surface of prosicula near boundary with metasicula. C. Arrangement of prosicular large pits. D–E. Details of foreiign material covering the outer surface of prosicula.F.Arragementofprosiculatinypitsonlargepitbottoms. G.Details of tiny pits. Abbreviations: c, creeping tube; e, erect tube; fm, foreign material; lp, prosicular large pit; m, metasicula; p, prosicula; r, rim of tiny pit; tp, prosicular tiny pit.

Fig. 2. Rhabdopleura compacta Hincks, 1888 in Cortical fibrils and secondary deposits in periderm of the hemichordate Rhabdopleura (Graptolithoidea)

Fig. 2. Rhabdopleura compacta Hincks, 1888, Recent. SEM micrographs of complete coenecia. A. Young coenecium composed of a sicula and two zooids (ZPAL Pb.5/1). B. Adult coenecium consisting of a sicula and a dozenorsozooids(ZPALPb.5/2). C.Youngcoeneciumbuiltofasiculaand three zooids (ZPALPb.5/3). Abbreviations: c, creeping tube; e, erect tube; m,metasicula;mm,marginalmembrane;p,prosicula;x,boundarybetween prosicula and metasicula.

FIGURE 12. Rhabdopleura emancipata 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 12. Rhabdopleura emancipata n. sp., A, B, D, E, paratype NIWA 161566; C, paratype 161211: A, B, distal part of a ringed tube, the bracketed section enlarged in B. C, transmitted-light view of a principal tube (stem) with openings of four ringed tubes all around; tentacles of one young zooid partly emergent; note how black stolon changes position within principal tube; cdc = cone of diaphragm complex. D, close-up of exterior surface of a principal tube showing parallel fusellar annuli marked by thin sutures. E, sclerotized tube, with terminal aperture at left, branching off a principal stolon. Scale bars: A, 200 μm; B, 100 μm; C, 400 μm; D, 100 μm; E, 1 mm.

FIGURE 11. Rhabdopleura emancipata 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 11. Rhabdopleura emancipata n. sp., A, B, holotype NIWA 13590; C, E, paratype 161211; D, paratype NIWA162566: A, B, close-up and whole colony (inset); principal tubes are identifiable in A by the dark-brown stolon that passes through them; abundant ringed tubes diverge almost at right angles from principal tubes. C, transmitted-light view of a principal tube (pt) from which two ringed tubes (rt) diverge and, between them, a branch of the principal tube with a bifurcation of the black stolon (bs); other abbreviations: cdc, cone of diaphragm complex; cs, contractile stalk (gymnocaulus) of zooid; fc, fusellar collar; s, septum. D, SEM of a principal tube (pt) (stem), with its aperture at left, and five ringed tubes in various stages of development. E, transmitted-light view of a principal tube (pt) (stem) and its black stolon (bs) in which are three young zooids (z), the middle one of which shows, from below, the opening of a potential ringed tube (ort). Scale bars: A, 3 mm; B, 3 cm; C‒E, 0.5 mm.

FIGURE 10. Rhabdopleura decipula 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 10. Rhabdopleura decipula n. sp., A, C, D, holotype 162592; B, E, paratype 3 NIWA 162591; F, paratype 4 NIWA 158517; G‒K, paratype 2 NIWA 161213. A, broken-tipped erect tube. B, distal end of young erect tube. C, D, subdistal and middle parts of same fully formed erect tube. E, F, parts of creeping tubes with some point-to-point suture zigzags highlighted. G, a prosicula (ps) and metasicula (ms); H‒K, reflected-light (H, I) and SEM (J, K) images of new colonies, with numbers indicating the inferred budding sequence (subsequent development seems to vary) of creeping tubes; the metasicula produces one or two metasiculae while one side of the prosicula is internally partitioned off to produce both a backwards-directed tube and an opposing creeping tube. Black stolons occur in H and a zooid in I. Scale bars: A, E‒H, 200 μm; B, 100 μm; C, D, K, 50 μm; I, J, 400 μm.

FIGURE 7. Rhabdopleura chathamica 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 7. Rhabdopleura chathamica n. sp., A‒C, F, paratype 90265; D, E, holotype 161214: A, part of colony showing three blind side branches, the proximal part of which is adherent; only one ringed erect tube (et) remains. B, close-up of middle side branch in A; note part of the pectocaulus/black stolon (bs) seen through a tear in the cuticle. C, the longest zooid seen, somewhat fouled and partly twisted in profile. D, non-fouled part of another erect tube showing well-developed fusellar collars. E, fusellar sutures on surface of creeping tube, with some point-to-point suture zigzags highlighted. F, close-up of fusellar sutures seen on adherent tube in B. Scale bars: A, 1 mm; B, 300 μm; C, F, 100 μm; D, 50 μm; E, 200 μm.

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.

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FIGURE 4 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 4. Original illustrations of described species of Rhabdopleura characterized by indirect inception of ringed erect tubes from creeping tubes via an intervening adnate portion that comprises a blind-ending branch. A, Rhabdopleura normani (from Allman 1869, pl. 8, fig. 1). Note that most of the colony is adherent; only the ringed tubes (rt) are erect, the proximal part of each side branch (ppsb) is adherent and the oblique sutures are not shown. B, schematic interpretative depiction of R. normani (after Andres 1977). C, D, Rhabdopleura grimaldii (from Jullien 1890, p. 181); note that ringed erect tubes (rt) are mostly lacking, with only the adherent proximal parts of side branches (ppsb) present. D is a cropped part of C to show the proximal parts of two side branches, with fusellar sutures and Jullien's putative 'very fine tubes' (vft)—actually the side walls of the tapering pectocaulus, the proximal end of which is transparent, hence the apparent gap between it and the pectocaulus of the creeping tube. E, F, Rhabdopleura striata (from Schepotieff 1909, pl. 7, figs 2, 7); E shows longitudinally striated erect parts of side branches; oblique sutures on prostrate parts omitted in E but shown in F, with characteristic web of fusellar sutures (wfs) in branch angles from main creeping tube. G, Rhabdopleura manubialis (from Jullien & Calvet, 1903, pl. 1, fig. 2) with dormant bodies (db).

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FIGURE 9. 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 9. Rhabdopleura francesca n. sp.: A, section of creeping tube of holotype with a broken erect tube; note the small erect-tube diameter relative to the width of the creeping tube. Rhabdopleura decipula n. sp.: B, holotype NIWA 162592 (left) and paratype 1 (NIWA 161212) colonies on broken shells; C, reflected-light image of part of a paratype 2 colony (NIWA 161213) seen in transparency on smooth shell, showing a loop of one creeping-tube branch curving back on itself, with a narrow crossconnection (cc) of the pectocaulus; erect tubes and zooids (z) and a dormant body (db) also seen. D, SEM image of same colony at lower magnification, with branch loop in C at top right. E, another part of same colony seen in transparency, showing a zooid inside a creeping tube to the left (distal) of a dark dormant body. Scale bars: A, 500 μm; B, 10 mm; C, 1 mm; D, 2 mm; E, 400 μm.