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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.

FIGURE 5 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific

FIGURE 5. Light micrographs of transverse sections of Saxipendium coronatum: A, Proboscis with heart–kidney complex. B, Anterior region of the proboscis neck. C, Posterior region of the proboscis neck showing the neuropore and skeleton plate. D, Anterior region of the collar. E, Posterior region of the collar showing the collar pores. F, Collar canal and pore. G, Posterior region of the collar showing the curved skeletal cornua. H, Anterior region of the trunk. Abbreviations: bp, branchial pore; bv, blood vessel; cc, collar canal; cl, collar lumen; clm, collar longitudinal muscles; cp, collar pore; cv, cardiac vesicle; dm, dorsal mesentery; g, glomerulus; gb, gill bar; go, gonad; gp, gonopore; 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 cornua; sp, skeletal plate; vm, ventral mesentery. Scale bars: A, C, D, G, H = 1000 µm; B, E = 500 µm; F = 200 µm.

FIGURE 2 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific

FIGURE 2. Drawings of live animals and photographs of fixed specimens of the family Harrimaniidae. A, Drawing of the dorsal side of Horstia kincaidi n. gen. et n. sp. B, Drawing of the lateral side of Mesoglossus intermedius n. gen. et n. sp. C, Drawing of the dorsal side of Mesoglossus intermedius. D, Drawing of the dorsal side of Stereobalanus willeyi n. sp. E, Drawing of the lateral side of Stereobalanus willeyi. F, Photograph of the dorsal side of Horstia kincaidi. G, Photograph of the lateral side of Ritteria ambigua n. gen. et n. sp. H, Photograph of the dorsal side of Mesoglossus intermedius. I, Photograph of the dorsal side of Stereobalanus willeyi. J, Photograph of the lateral side of Stereobalanus willey. Abbreviations: bp, branchial pore; c, collar; gr, genital ridge; p, proboscis; t, trunk. Scale bars = 2 mm

FIGURE 3 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific

FIGURE 3. Light micrographs of transverse sections of Stereobalanus willeyi n. sp.: A, Proboscis with heart–kidney complex. B, Posterior part of the proboscis showing the skeleton. C, Proboscis neck. D, Anterior region of the collar. E, Anterior pharyngeal region of the trunk. F, Genital region of the trunk. Abbreviations: cl, collar lumen; cv, cardiac vesicle; g, glomerulus; gb, gill bar; go, gonad; nc, nerve cord; np, neuropore; pbr, parabranchial ridge; pc, proboscis coelom; pl, pharynx lumen; plm, proboscis longitudinal muscles; pv, proboscis vesicle; s, stomochord; sb, skeletal body; sc, skeletal cornua; sl, stomochord lumen; vc, ventral caecum; vm, ventral mesentery. Scale bars: A, D, E, F = 1000 µm; B = 500 µm; (C) = 750 µm.

FIGURE 1 in A taxonomic revision of the family Harrimaniidae (Hemichordata: Enteropneusta) with descriptions of seven species from the Eastern Pacific

FIGURE 1. Light micrographs of transverse sections of the proboscis of: A, Saccoglosus pusillus, showing the arrangement of the proboscis longitudinal musculature in concentric rings. B, Protoglossus mackiei n. sp., showing the arrangement of the proboscis longitudinal musculature in radial plates. C, Mesoglossus macginitiei n. gen. et n. sp., showing the diffuse arrangement of the proboscis longitudinal musculature. Abbreviations: plm, proboscis longitudinal muscles. Scale bars = 500 µm.