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Figures 1–3 in First record of the family Spengeliidae (Hemichordata: Enteropneusta) from Chinese waters, with description of a new species

Figures 1–3. Glandiceps qingdaoensis, new species, external form. (1) Fresh specimen (from dorsal side). (2) Preserved specimen (holotype). (3) Preserved specimen (paratype not dissected)

Figures 13–19 in First record of the family Spengeliidae (Hemichordata: Enteropneusta) from Chinese waters, with description of a new species

Figures 13–19. Glandiceps qingdaoensis, new species, anatomical and histological characteristics. (13) Transverse section through the median region of collar showing perihaemal coelom and mesentery. (14) Longitudinal section through median-posterior region of collar showing small disconnected medullary cavities arranged in a line and collar dorsal nerve strand. (15) Transverse section through the branchial region showing well-developed digestive portion and branchial portion. (16) Magnified upper part of Figure 15, showing dorsal septum, dorsal vessel, and splanchnic nerve-fibre layer. (17) Magnified lower part of Figure 15, showing ventral septum, ventral vessel, and nerve-fibre layer. (18) Longitudinal section through the branchial genital region showing gill bar, tongue bar, and distinct vesicle encircling every tongue bar. (19) Transverse section through intestine, showing the thin wall of digestive canal.

Figures 7–12 in First record of the family Spengeliidae (Hemichordata: Enteropneusta) from Chinese waters, with description of a new species

Figures 7–12. Glandiceps qingdaoensis, new species, histological characteristics. (7) Transverse section through base of proboscis passing through the pouched region of the stomochord, behind the glomerulus, showing the lateral pouch and median pouch of stomochord. The pericardium is above the median pouch. (8) Longitudinal section through median septum of the proboscis to show the vermiform process of the stomochord and its varied calibre (diameter of vermiform process tube). (9) Transverse section through anterior end of the stomochord showing small pericardial auricles and pericardium wedged in the longitudinal muscles. (10) Transverse section through commencement of the heart–kidney complex (include stomochord, glomerulus, and pericardium). The section passes immediately in front of the anterior end of the pericardium, and shows the right and left halves of the glomerulus on either side of the stomochord and distinct median glomerulus. (11) Longitudinal section through the heart–kidney complex showing large left half part of glomerulus, which envelops the anterior stomochord. (12) Transverse section through the junction between proboscis and collar showing chondroid tissue and nuchal skeleton which is strongly developed and occupying a large portion of the section.

Figures 4–6 in First record of the family Spengeliidae (Hemichordata: Enteropneusta) from Chinese waters, with description of a new species

Figures 4–6. Glandiceps qingdaoensis, new species, histological characteristics. (4) Longitudinal section through proboscis, collar, and anterior part of branchial-genital region of holotype, showing overall structure of proboscis. Circular musculature of the anterior part thicker than posterior part, dense and sparse longitudinal musculature, cornua of the skeleton extends to the median posterior portion of collar. (5) Transverse section through the anterior proboscis of paratype, showing the complete dorsal-ventral muscle septum. (6) Magnified photograph of centric portion of Figure 5, to show many irregular gland cells.

FIG. 7 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 7. — Bayesian cladogram based on enteropneust 18S and 16S rRNA sequences, rooted using an echinoid echinoderm (Echinocrepis rostrata Mironov, 1973). Numbers at nodes represent posterior probabilities (p). All nodes with p<0.95 were collapsed into polytomies.

FIG. 3. — Tergivelum baldwinae n. gen., n in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 3. — Tergivelum baldwinae n. gen., n. sp. in approximate dorsal view: A, surface details; B, locations of major nerves and muscles associated with proboscis and collar. Abbreviations for structural features: see text.

FIG. 8 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 8. — Single frame (at 5 min: 37 sec) of 8-min videotape of Tergivelum baldwinae n. gen., n. sp. paratype IV. Filled circles show path of a particle transported unobstructed about 5 cm above the bottom (from 5:47 to 5:53). Open circles show path of particle (small polychaete) drifting about 2 cm above the bottom and evidently getting stuck (at 5:36) in substantial mucus surrounding acorn worm;arrow indicates shadow cast on sea floor by the immobilized particle. Scale bar: 5 cm.

FIG. 6 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 6. — Cross sections (B-J) of Tergivelum baldwinae n. gen., n. sp. at levels indicated in 4B: A, parasagittal section in region of gill pores showing primary gill bar with skeleton (arrowed) and secondary gill bar with skeleton (arrowheads) and coelomic space (asterisk); B, cross section of trunk near anterior extremity of ventral nerve cord (in rectangle); C, detail from rectangle in 6B with ventral nerve cord underlain by ventral blood vessel (arrowhead); D, cross section of anterior trunk showing dorsal protuberances and back veils (the left one broken off); dark spheres are testes; E-J, intestinal region of trunk; asterisks mark lateroventral folds); dorsal and ventral nerve cords indicated, respectively, by arrows and arrowheads; K, section through three testes; L, detail of testis with spermatozoafilled lumen toward top right; M, medium-sized oocytes; N, large oocyte. Abbreviations for structural features: see text. Scale bars: A, 100 μm; B, D, 1 mm; C, 50 μm; E-J, N, 500 μm; K, M, 200 μm; L, 20 μm.

FIG. 5 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 5. — Cross sections of Tergivelum baldwinae n. gen., n. sp. at levels indicated in Figure 4B: A, near anterior tip of proboscis, arrow indicates tract of longitudinal muscle fibers; B, enlargement of rectangle in 5A showing epidermis (toward right) underlain by basal concentration (arrowed) of diffuse intraepidermal nervous system; proboscis coelom contains meshwork of connective tissue and muscle cells; C, more posterior section of proboscis; D, section through collar showing anterior end of right buccal muscle; specimen oriented with left side somewhat in advance of right, so left buccal muscle not yet in view; E, detail of the center rectangle in 5D; showing lumen of collar cord separating dorsal non-neural and ventral neural regions; perihaemal coeloms (marked by asterisks) lie on either side of dorsal blood vessel; F, detail of left-hand rectangle in 5D. Collar-trunk septum separates trunk coelom (above) from collar coelom (below); G, section through level of the mouth; lateral lips comprise ventral region of collar; posteroventral extremity of proboscis forms anterior margin of mouth; H, detail in the rectangle in 5G; beneath dorsal nerve cord is dorsal blood vessel with parts of trunk coelom (asterisks) on either side; I, section at level of posterior rim of mouth; buccal muscles approaching each other and will merge a few sections more posteriorly; pharynx comprising ventral digestive region (arrowheads) and dorsal branchial region (obliquely sectioned) penetrated by gill pores. Abbreviations for structural features: see text. Scale bars: A, C, D, G, I, 1 mm; B, 50 μm; E, 100 μm; F, 300 μm; H, 200 μm.

FIG. 4 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 4. — Fixed specimens of Tergivelum baldwinae n. gen., n. sp.: A, holotype (ventral view) which broke into three parts when fixed – namely the left back veil (lbv), most of intestinal region of trunk (at right), and anterior body (at left, comprising proboscis, collar, and anterior extremity of trunk); B, holotype (dorsal view, excluding left back veil) with numbered letters indicating levels of cross section in Figures 5 and 6; C, enlargement of anterior portion of holotype in ventral view; lateral lips of collar (arrowheads) curl dorsally as a preservation artifact; D, enlargement of anterior portion of the holotype in dorsal view; arrows indicate dorsal protuberances; E, enlargement of anterior portion of holotype in left side view with arrow indicating fossa of left proboscis nerve; F, dorsal view of anterior region of trunk of paratype II (corresponding to area in rectangle in panel 4D); back veils and dorsal protuberances dissected away revealing dorsal nerve cord (arrowed) flanked by slot-shaped gill pores. Abbreviations for structural features: see text. Scale bars: A, B, 1 cm; C-F, 2 mm.

FIG. 2 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 2. — Single video frames of living Tergivelum baldwinae n. gen., n. sp.: A, anterior of paratype IV, white arrowhead indicates fossa of left proboscis nerve; B, holotype laying down fecal trail; suction sampler in background; C, paratype II laying down fecal trail, 29 cm between red laser dots. Abbreviations: co, collar; lbv, left back veil; pr, proboscis; rlf, right lateroventral fold. Scale bars: A, 2 cm; B, C, 3 cm.

FIG. 1 in A new deep-sea species of epibenthic acorn worm (Hemichordata, Enteropneusta)

FIG. 1. — Collection site (depth: 3952-3953 m) for holotype and paratypes I-VI of Tergivelum baldwinae n. gen., n. sp.

FIGURE 4 in Three new species of Glossobalanus (Hemichordata: Enteropneusta: Ptychoderidae) from western North America

FIGURE 4. Light micrographs of transverse sections of Glossobalanus barnharti. A, Proboscis with massive radial muscle bundles. B, Proboscis heart-kidney complex with lateral and dorsal glomerulus. C, Peduncle showing the large lumen of the stomochord. D, Peduncle with deep and narrow keel. E, Collar. F, Collar cord with dorsal nerve roots. G, Anterior pharynx large dorsal branchial lumen. H, Intestinal region with lateral septum and median gonads. Abbreviations: bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cv, cardiac vesicle; ds, dorsal septum; ep, epidermis g, glomerulus; gb, gill bar; mg, median gonad; lg, lateral gonad; ls, lateral septum; mg, median gonad; nc, nerve cord; nr, nerve roots; pbd, peribuccal diverticula; pc, proboscis coelom; phd, perihaemal diverticula; pl, pharynx lumen; plm, proboscis longitudinal muscles; s, stomochord; sk, skeletal keel; tc, trunk coelom; vp, vascular plexus. Scale bars: A, E, F, G, H = 500 µm; B, C = 200 µm; (D) = 250 µm.

FIGURE 3 in Three new species of Glossobalanus (Hemichordata: Enteropneusta: Ptychoderidae) from western North America

FIGURE 3. Light micrographs of transverse sections of Glossobalanus hartmanae. A, Proboscis with conspicuous circularmuscle layer, radial muscle bundles, heart-kidney complex and ventral septum. B, Proboscis peduncle showing the ventrolateral pouches of the stomochord and short skeletal keel. C, Collar showing a midventral vascular plexus. D, Dorsal nerve cord of collar with dorsal roots. E, Posterior collar showing canals. F, Anterior pharynx with plicate gill-bar epithelium. G, Intestinal region with lateral septum and median gonads. Abbreviations: a, atrium; bv, blood vessel; cc, collar canal; cg, ciliated groove; cl, collar lumen; cv, cardiac vesicle; ds, dorsal septum; ep, epidermis; g, glomerulus; gb, gill bar; gbe, gill-bar epithelia; gd, gonad; gp, gill pore; mg, median gonad; lg, lateral gonad, ls, lateral septum; nc, nerve cord; nr, nerve roots; pbd, peribuccal diverticula; pc, proboscis coelom; pcm, proboscis circular muscles; phd, perihaemal diverticula; pl, pharynx lumen; plm, proboscis longitudinal muscles; pr, parabranchial ridges; s, stomochord; sc, skeletal cornua; sk, skeletal keel; tlm, trunk longitudinal muscles; vp, vascular plexus. Scale bars: A, C, F, G = 500 μm; B = 200 μm; D, E = 250 μm.

FIGURE 2 in Three new species of Glossobalanus (Hemichordata: Enteropneusta: Ptychoderidae) from western North America

FIGURE 2. Light micrographs of transverse sections of Glossobalanus williami. A, Proboscis with radial muscle bundles, heart-kidney complex, and ventral septum. B, Proboscis heart-kidney complex with a distinct lumen and poorly developed glomerulus. C, Proboscis peduncle showing the proboscis canal and obtuse skeletal keel. D, Anterior collar showing only a dorsal septum. E, Mid collar with parabuccal diverticula and lateral skeletal cornua. F, Posterior collar showing canals to the first pair of gill atria. G, Pharynx with conspicuous gill slits. H, Pharynx showing only primary gonads. Abbreviations: cc, collar canal; cl, collar lumen; cv, cardiac vesicle; dep; dorsal epidermal pocket; ds, dorsal septum; ep, epidermis g, glomerulus; gb, gill bar; go, gonad; gp, gill pore; nc, nerve cord; nfl, nerve-fiber layer; pbd, peribuccal diverticula; pc, proboscis coelom; pcm, proboscis circular muscles; pcp, proboscis canal (and pore) of the peduncle; phd, perihaemal diverticula; pl, pharynx lumen; plm, proboscis longitudinal muscles; pn, anterior neuropore; pr, parabranchial ridges; s, stomochord; sb, skeletal body; sc, skeletal cornua; sk, skeletal keel; tc, trunck coelom; tlm, trunk longitudinal muscles; vs, ventral septum. Scale bars: A, E, F = 500 μm; B, C = 100 μm; D, G, H = 250 μm.

FIGURE 1 in Three new species of Glossobalanus (Hemichordata: Enteropneusta: Ptychoderidae) from western North America

FIGURE 1. Photograph of fixed specimens of A, Glossobalanus hartmanae n. sp.; B, G. b a r n h a r t i n. sp.; C, G. m i n u t u s. Abbeviations: bg, branchiogenital region of trunk; c, collar; ct, caudal region of trunk; ht, hepatic trunk; p, proboscis. Scale bar = 5 mm.

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.