C5X/J?:/V/1
NOAA TR NMFS CIRC-375
A UNITED STATES
DEPARTMENT OF
COMMERCE
PUBLICATION
*^°r^
:t:
NOAA Technical Report NMFS CIRC-375
U.S. DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
National Marine Fisheries Service
New Polychaeta from Beaufort,
with a Key to All Species
Recorded from North Carolina
JOHN H. DAY
SEATTLE, WA
JULY 1973
NOAA TECHNICAL REPORTS
National Marine Fisheries Service, Circulars
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315. Synopsis of biological data on the chum salmon,
Oncorhynchus keta (Walbaum) 1792. By Rich-
ard G. Bakkala. March 1970, iii + 89 pp., 15
figs., 51 tables.
319. Bureau of Commercial Fisheries Great Lakes
Fishery Laboratory, Ann Arbor, Michigan. By
Bureau of Commercial Fisheries. March 1970,
8 pp., 7 figs.
330. EASTROPAC Atlas: Vols. 4, 2. Catalog No.
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able from the Superintendent of Documents,
Washington, D.C. 20402.
331. Guidelines for the processing of hot-smoked chub.
By H. L. Seagran, J. T. Graikoski, and J. A.
Emerson. January 1970, iv + 23 pp., 8 figs.,
2 tables.
332. Pacific hake. (12 articles by 20 authors.) March
1970, iii + 152 pp., 72 figs., 47 tables.
333. Recommended practices for vessel sanitation and
fish handling. By Edgar W. Bowman and Alfred
Larsen. March 1970, iv + 27 pp., 6 figs.
335. Progress report of the Bureau of Commercial
Fisheries Center for Estuarine and Menhaden
Research, Pesticide Field Station, Gulf Breeze,
Fla., fiscal year 1969. By the Laboratory staff.
August 1970, iii + 33 pp., 29 figs., 12 tables.
336. The northern fur seal. By Ralph C. Baker, Ford
Wilke, and C. Howard Baltzo. April 1970, iii +
19 pp., 13 figs.
337. Program of Division of Economic Research,
Bureau of Commerecial Fisheries, fiscal year
1969. By Division of Economic Research. April
1970, iii + 29 pp., 12 figs., 7 tables.
338. Bureau of Commercial Fisheries Biological Lab-
oratory, Auke Bay, Alaska. By Bureau of Com-
mercial Fisheries. June 1970, 8 pp., 6 figs.
339. Salmon research at Ice Harbor Dam. By Wesley
J. Ebel. April 1970, 6 pp., 4 figs.
340. Bureau of Commercial Fisheries Technological
Laboratory, Gloucester, Massachusetts. By Bu-
reau of Commercial Fisheries. June 1970, 8 pp.,
8 figs.
341. Report of the Bureau of Commercial Fisheries
Biological Laboratory, Beaufort, N.C., for the
fiscal year ending June 30, 1968. By the Lab-
oratory staff. August 1970, iii -f- 24 pp., 11 figs.,
16 tables.
342. Report of the Bureau of Commercial Fisheries
Biological Laboratory, St. Petersburg Beach,
Florida, fiscal year 1969. By the Laboratory staff.
August 1970, iii + 22 pp.," 20 figs., 8 tables.
343. Report of the Bureau of Commercial Fisheries
Biological Laboratory, Galveston, Texas, fiscal
year 1969. By the Laboratory staff. August
1970, iii + 39 pp., 28 figs., 9 tables.
344. Bureau of Commercial Fisheries Tropical Atlan-
tic Biological Laboratory progress in research
1965-69, Miami, Florida. By Ann Weeks. Oc-
tober 1970, iv + 65 pp., 53 figs.
346. Sportsman's guide to handling, smoking, and pre-
serving Great Lakes coho salmon. By Shearon
Dudley, J. T. Graikoski, H. L. Seagran, and Paul
M. Earl. September 1970, iii + 28 pp., 15 figs.
347. Synopsis of biological data on Pacific ocean perch,
Sebastodes alutus. By Richard L. Major and
Herbert H. Shippen. December 1970, iii + 38
pp., 31 figs., 11 tables.
Continued on inside back cover.
^OATMOSP
'Went of °
U.S. DEPARTMENT OF COMMERCE
Peter G. Peterson, Secretary
NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION
Robert M. White, Administrator
NATIONAL MARINE FISHERIES SERVICE
NOAA Technical Report NMFS CIRC-375
New Polychaeta from Beaufort,
with a Key to All Species
Recorded from North Carolina
JOHN H. DAY
S
SEATTLE, WA
'JULY 1973
tr sale by the Superintendent of Documents. U.S. Government Printing Office
ashington, D.C. 20402 • $1.25
The National Marine Fisheries Service (NMFS) does not approve, rec-
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CONTENTS
Page
Introduction 1
Family Aphroditidae 4
Key to genera and species 4
Laetmonice filicornis Kinberg, 1855 4
Aphrodita sp 4
Aphrogenia sp 4
Family Polynoidae 4
Key to genera and species 4
Drieschia pellucida Moore, 1903 6
Lepidonotus sublevis Verrill, 1873 6
Lepidonotus variabilis Webster, 1879 6
Lepidametria commensalis Webster, 1879 6
Lepidasthenia sp 6
Subadyte pellucida (Ehlers, 1864) 6
Antinoella sarsi (Malmgren, 1865) 7
Harmothoe (Hermadion) acanellae (Verrill, 1881) 7
Gattyana cirrosa (Pallas, 1766) 7
Harmothoe imbricata (Linnaeus, 1767) 7
Harmothoe aculeata Andrews, 1891 7
Harmothoe trimaculata (Treadwell, 1924) 7
Harmothoe sp. A 7
Harmothoe sp. B 9
Family Polyodontidae 9
Key to genera and species 9
Polyodontes lupina (Stimpson, 1856) 9
Eupanthalis kinbergi (Mcintosh, 1876) 9
Family Sigalionidae 10
Key to genera and species 10
Pholoe minuta (Fabricius, 1780) 10
Sigalion arenicola Verrill, 1879 10
Leanira hystricis Ehlers, 1874 11
Psammolyce ctenidophora new species 11
Sthenelais boa (Johnston, 1833) 12
Sthenelais limicola (Ehlers, 1864) 12
Sthenelais sp 12
Sthenelais anocula new species 12
Family Palmyridae 14
Key to genera and species 14
Bhaivania goodei Webster, 1884 14
Paleanotus heteroseta Hartman, 1945 14
Family Pisionidae 14
Pisione remota (Southern, 1914) 14
Family Amphinomidae 15
Key to genera and species 15
Chloeia viridis Schmarda, 1861 15
Hipponoe gaudichaudi Audouin and Milne-Edwards, 1830 16
Amphinome rostrata (Pallas, 1776) 16
Paramphinome pulchella (Sars, 1872) 16
Pseudeurythoe ambigua (Monro, 1933) 16
iii
Family Phyllodocidae 16
Key to genera and species 16
Lug ia rarica Usehakov, 1958 18
Eteone lactea Claparede, 1868 19
Eteone heteropoda Hartman, 1951 19
Protomystides bidentata Langerhans, 1879 19
Hesionura elongata (Southern, 1914) 19
Euldlid (Pterocimis) macroceros Grube, 1860 20
E alalia (Eumida ) sat/guinea (Oersted, 1843) 20
Eulalia bilineata (Johnston, 1840) 20
E alalia viridis (Linnaeus, 1767) 20
Paranaitis kosteriensis (Malmgren, 1867) 20
Paranaitis speciosa (Webster, 1880) 22
Phyllodoce (Nereiphylla)fragilis Webster, 1879 22
Phyllodoce (Genetyllis) castaned (Marenzeller, 1879) 22
Phyllodoce (Anaitides) mucosa Oersted, 1843 22
Phyllodoce (Anaitides) groe?ilandica Oersted, 1843 22
Phyllodoce (Anaitides) longipes Kinberg, 1866 23
Phyllodoce (Anaitides) madeirensis Langerhans, 1880 23
Phyllodoce (Anaitides) arenae Webster, 1880 23
Phyllodoce (Anaitides) panamensis Treadwell, 1917 24
Family Pilargidae 24
Key to genera and species 24
Sigambra bassi (Hartman, 1945) 24
Synelmis albini (Langerhans, 1881) 24
Family Hesionidae 25
Key to genera and species 25
Gyptis vittata Webster and Benedict, 1887 25
Ophiodromus obscurus (Verrill, 1873) 25
Nerimyra punctata (Muller, 1776) » . . . 25
Parahesione luteola (Webster, 1880) 25
Family Syllidae 26
Key to genera and species 26
Trypanosyllia zebra (Grube, 1860) 28
Syllis (Haplosyllis) spongicold (Grube, 1855) 29
Syllis gracilis Grube, 1840 29
Syllis (Langerhatisia)ferrugina Langerhans, 1881 29
Syllis (Langerha nsia ) eornuta Rathke, 1843 29
Syllis (Typosyllis) hyalina Grube, 1863 29
Syllis (Typosyllis) regulata carolinae new subspecies 30
Syllis (Typosyllis) alternata Moore, 1908 30
Syllis (Typosyllis) prolifera Krohn, 1852 30
Amblyosyllis formosa Claparede, 1863 30
Sfreptosyllis arenae Webster and Benedict, 1884 30
Odontosyllis longiseta new species 32
Ensyllis lamelligera Marion and Bobretzky, 1875 32
Parapiono syllis longicirrata (Webster and Benedict, 1884) 32
Pionosyll is cf. uraga Imajima, 1966 33
Exogone gemmifera (Pagenstecher, 1862) 33
Exogone dispar (Webster, 1879) 33
Exogone verugera (Claparede, 1868) 34
Sphaerosyllis fortuita Webster, 1879 34
IV
Sphaerosyllis pirifera Claparede, 1868 34
Eurysyllis tuberculata Ehlers, 1864 34
Brania pusilla (Dujardin, 1839) 34
Brania clavata (Claparede, 1863) 35
Proceraea cornuta (Agassiz, 1863) 35
Proceraea fas data (Bosc, 1802) 35
- Autolytus prolifer (Muller, 1784) 35
Autolytus mbropunctatus (Grube, 1840) 35
Autolytus dentalius Imajima, 1966 35
Family Sphaerodoridae 36
Key to genera and species 36
Ephesiella claparedii (Greeff, 1866) 36
Family Nereidae 36
Key to genera and species 36
Lycastopsis pontica (Bobretzky, 1872) 38
Ceratocephale loveni Malmgren, 1867 38
Laeonereis culveri (Webster, 1879) 38
Websterinereis tridentata (Webster, 1880) 38
Ceratonereis irritablis (Webster, 1879) 38
Ceratouereis versipedata Ehlers, 1887 39
Platynereis dumerilii (Audouin and Milne-Edwards, 1833) 39
Nereis (Nereis) lamellosa Ehlers, 1868 39
Nereis (Nereis) grayi Pettibone, 1956 39
Nereis (Nereis) riisei Grube, 1856 39
Nereis (Nereis) falsa Quatrefages, 1865 41
Nereis (Neanthes) succinea Frey and Leuckart, 1847 41
Nereis (Neanthes) acuminata Ehlers, 1868 41
Family Nephtyidae 41
Key to subgenera and species of Nephtys 41
Nephtys (Inermonephtys) inermis Ehlers, 1887 42
Nephtys (Aglaophamus) verrilli Mcintosh, 1885 42
Nephtys (Aglaophamus) circinata Verrill, 1874 43
Nephtys pieta Ehlers, 1868 43
Nephtys squamosa Ehlers, 1887 43
Nephtys incisa Malmgren, 1865 43
Nephtys bucera Ehlers, 1868 43
Family Glyceridae 44
Key to genera and species 44
Hemipodus rosea s Quatrefages, 1865 44
Glycera dibrunehiata Ehlers, 1868 45
Glycera americana Leidy, 1855 45
Glyeera papillosa Grube, 1857 45
Glycera capitata Oersted, 1843 46
Glycera oxycephala Ehlers, 1887 46
Glycera tesselata Grube, 1863 46
Glycera asymmetrica new species 47
Family Goniadidae 47
Key to genera and species 47
Progoniada regularis Hartman, 1965 48
Goniadides carolinae new species 48
Glycinde solitaria (Webster, 1880) 49
Glyciude nordmanni (Malmgren, 1865) 49
Goniadella gracilis (Verrill, 1873) 51
Goniada teres Treadwell, 1931 51
Goniada maculata Oersted, 1843 51
Goniada littorea Hartman, 1950 51
Goniada brunnea Treadwell, 1906 51
Goniada uorvegica Oersted, 1845 51
Family Eunicidae 52
Key to genera and species 52
Lysidice ninetta ninetta Audouin and Milne-Edwards, 1833 52
Lysidice ninetta collaris Grube, 1870 52
Marphysa sanguinea (Montagu, 1815) 52
Eunice antennata (Savigny, 1820) 53
Eunice icebsteri Fauchald, 1969 53
Eunice filamoitosa Grube, 1856 53
Family Onuphidae 53
Key to genera and species 53
Hyalinoecia tubicola (Muller, 1776) 54
Diopatra cuprea cuprea (Bosc, 1802) 54
Diopatra cuprea spiribranchis Augener, 1906 55
Rhamphobrachium atlanticum new species 55
Onuphis (Nothria) conchylega Sars, 1835 56
Onuphis (Nothria) pallidula (Hartman, 1965) 57
Onuphis eremita Audouin and Milne- Edwards, 1833 57
Onuphis nebulosa Moore, 1911 57
Onuphis microcephala Hartman, 1944 57
Onuphis magna (Andrews, 1891) 57
Family Lumbrineridae 57
Key to genera and species 57
Lumbrineris inflata Moore, 1911 59
Lumbrineris coccinea (Renier, 1804) , . 59
Lumbrineris aberrans Day, 1963 59
Lumbrineris paradoxa Saint-Joseph, 1888 59
Lumbrineris tetraura (Schmarda, 1861) 59
Lumbrineris sp 60
Lumbrineris albidentata Ehlers, 1908 60
Lumbrineris latreilli (Audouin and Milne- Edwards, 1833) 60
Lumbrineris cruzensis Hartman, 1944 60
Lumbrineris brevipes (Mcintosh, 1903) 62
Lumbrineris frag ilia (Muller, 1776) 62
Lumbrineris januarii (Grube, 1878) 62
Lumbrineris tenuis Verrill, 1873 62
Family Arabellidae 63
Key to genera and species 63
Notocirrus spiniferus (Moore, 1906) 63
Drilonereis filum (Claparede, 1868) 63
Drilonereis longa Webster, 1879 63
Drilonereis magna Webster and Benedict, 1887 63
Arabella iricolor (Montagu, 1804) 64
Arabella mutans (Chamberlin, 1919) 64
Family Dorvilleidae 64
Key to genera and species 64
Ophryotrocha puerilis Claparede and Mecznikow, 1869 64
VI
Protodorvillea kefersteini (Mcintosh, 1869) 64
Dorvillea rudolphi (Delle Chiaje, 1828) 65
Dorvillea sociabilis (Webster, 1879) 65
Dorvillea caeca (Webster and Benedict, 1884) 65
Family Spionidae 65
Key to genera and species 65
Polydora commensalis Andrews, 1891 68
Polydora websteri Hartman, 1943 68
Polydora caeca (Oersted, 1843) 68
Polydora socialis (Schmarda, 1861) 69
Polydora hartmanae Blake, 1971 69
Polydora colonia Moore, 1907 69
Polydora tetrabranchia Hartman, 1945 69
Polydora ligni Webster, 1879 69
Spiophanes bombyx (Claparede, 1870) 69
Spiophanes wigleyi Pettibone, 1962 69
Scolecolepides viridis (Verrill, 1873) 70
Dispio uncinata Hartman, 1951 70
Spio cf. multioculata (Rioja, 1919) 70
Spio pettiboneae Foster, 1970 70
Spio setosa Verrill, 1873 72
Streblospio benedicti Webster, 1879 72
Paraprionospio pinnata (Ehlers, 1901) 72
Scolelepis squamata (Muller, 1806) 72
Aonides sp 72
Prionospio (Minuspio) cirrifera Wiren, 1883 72
Prionospio (Minuspio) cirrobranchiata Day, 1961 73
Prionospio heterobranchia Moore, 1907 73
Prionospio dayi (Foster, 1969) 73
Prionospio fallax Soderstrom, 1920 74
Prionospio cristata Foster, 1971 74
Prionospio steenstrupi Malmgren, 1867 74
Microspio pigmentata (Reish, 1959) 74
Laonice cirrata (Sars, 1851) 76
Ne7inides unidentata new species 76
Family Magelonidae 77
Key to species of Magelona 77
Magelona papillicornis Muller, 1858 77
Magelona phyllisae Jones, 1963 77
Magelona sp 78
Magelona rosea Moore, 1907 78
Family Poecilochaetidae 78
Poecilochaetus sp 78
Family Chaetopteridae 79
Key to genera and species 79
Phyllochaetopterus socialis Claparede, 1870 79
Spiochaetoptems costarum oculatus Webster, 1879 79
Chaetopterus variopedatus (Renier, 1804) 80
Mesochaetopteriis taylori Potts, 1914 80
Family Cirratulidae 80
Key to genera and species 80
Cirriformia grandis (Verrill, 1873) new combination 81
vu
Cimformia filigera (Delle Chiaje, 1828) 81
Dodecaceria corallii (Leidy, 1855) 82
Tharyx annulosus Hartman, 1965 82
Tharyx setigera Hartman, 1945 82
Tharyx marioni (Saint-Joseph, 1894) 82
Caulleriella killariensis (Southern, 1914) 82
Chaetozone gayheadia Hartman, 1965 83
Chaetozone setosa Malmgren, 1867 83
Family Orbiniidae 83
Notes on the genera of the subfamily Orbiniinae 83
Generic definitions 87
Naineris Blainville, 1828 87
Orbhtia Quatrefages, 1865 87
Phylo Kinberg, 1866 87
Scoloplos Blainville, 1828 87
Haploscoloplos Monro, 1933 87
Califia Hartman, 1957 87
Key to genera and subgenera of Orbiniinae and the species
recorded from North Carolina 88
Phylo ornatus (Verrill, 1873) 89
Phylo felix Kinberg, 1866 89
Orbinia americana new species 89
Orbinia riseri (Pettibone, 1957) new combination 90
Scoloplos (Scoloplos) capensis (Day, 1961) new combination 90
Scoloplos (Scoloplos) cf. acmeceps Chamberlin, 1919 91
Scoloplos (Scoloplos) armiger (Muller, 1776) 91
Scoloplos (Leodamas) rubra (Webster, 1879) 91
Haploscoloplos foliosus Hartman, 1951 91
Haploscoloplos fragilis (Verrill, 1873) 91
Haploscoloplos robustus (Verrill, 1873) 91
Family Paraonidae 92
Key to genera and species 92
Cirrophorus lyriformis (Annenkova, 1934) 92
Cirrophorus branchiatus Ehlers, 1908 93
Aricidea cerruti Laubier, 1967 93
Aricideafauveli Hartman, 1957 93
Aricidea fragilis Webster, 1879 93
Aricidea suecica Eliason, 1920 93
Aedicira albatrossae (Pettibone, 1957) 93
Aedicira belgicae (Fauvel, 1936) 94
Paraonis gracilis (Tauber, 1879) 94
Paraonis fulgens (Levinsen, 1883) 94
Family Opheliidae 94
Key to genera and species 94
Travisia parva new species 95
Ophelia denticulata Verrill, 1875 95
Armandia maculata (Webster, 1884) 95
Armandia agilis (Andrews, 1891) 95
Ophelina cylindricaudata (Hansen, 1879) 96
Family Scalibregmidae 96
Key to genera and species 96
Scalibregma inflation Rathke, 1843 96
Hyboscolex longiseta Schmarda, 1861 96
Family Arenicolidae 98
Arenicola cristata Stimpson, 1956 98
Family Capitellidae 98
Key to genera and species 98
Capitella capitata (Fabricius, 1780) 99
' Mediomastus californiensis Hartman, 1944 99
Leiochrides pallidior (Chamberlin, 1918) 99
Heteromastus filiformis (Claparede, 1864) 99
Notomastus lobatus Hartman, 1947 100
Notomastus latericeus Sars, 1851 100
Notomastus hemipodus Hartman, 1947 100
Notomastus americanus new species 100
Leiocapitclla glabra Hartman, 1947 100
Dasybranchus lumbricoides Grube, 1878 101
DasybrancJius lunulatus Ehlers, 1887 101
Family Maldanidae 102
Key to genera and species 102
Petaloproctus socialis Andrews, 1891 103
Nicomache trispinata Arwidsson, 1906 103
Praxillella sp 103
Clymenella torquata (Leidy, 1855) 103
Macroclymeiic zonalis (Verrill, 1874) 103
Axiothella mucosa (Andrews, 1891) 103
Branchioasyehix america?ia Hartman, 1945 104
Asyctiis carol hiac new species 104
Family Oweniidae 104
Oivenia fiisiformis Delle Chiaje, 1844 104
Family Sternaspidae 105
Steruaspis scutata (Ranzani, 1817) 105
Family Flabelligeridae 105
Key to genera and species 105
Diplocirrus capensis Day, 1961 105
Flabelligera sp 107
Pherusa inflata (Treadwell, 1914) 107
Pherusa ehlersi new species 107
Notes on the genera Piromis Kinberg, 1867 and Pherusa Oken, 1807 108
Piromis cruca (Claparede, 1869) new combination 108
Piromis cruca websteri new subspecies 109
Family Sabellariidae 109
Key to genera and species 109
Sabellaria bella Grube, 1870 110
Sabellaria floridensis Hartman, 1944 110
Sabellaria gracilis Hartman, 1944 110
Sabellaria vulgaris vulgaris Verrill, 1873 110
Sabellaria vulgaris beaufortensis Hartman, 1944 110
Family Pectinariidae Ill
Key to subgenera and species Ill
Pectinaria (Pectinaria) gouldii Verrill, 1873 Ill
Family Ampharetidae Ill
Key to genera and species Ill
Melinnopsis atlantica Mcintosh, 1885 112
l\
Melinna maculata Webster, 1879 113
Isolda pulchella Miiller, 1858 113
Auchenoplax crinita Ehlers, 1887 113
Samythella eliasoni new species 113
Amage auricula Malmgren, 1866 115
Amphicteis gunneri (Sars, 1835) 115
Ampharete acutifrons Grube, 1860 115
Ampharete parvidentata new species 115
Ampharete americana new species 116
Family Terebellidae 116
Key to genera and species 116
Terebellides stroemii Sars, 1835 118
Trichobranchus glacialis Malmgren, 1866 118
Thelepus setosus (Quatrefages, 1865) 118
Amphitrite ornata (Leidy, 1855) 119
Terebella pterochaeta Schmarda, 1861 119
Terebella lapidaria Linnaeus, 1767 119
Terebella rubra (Verrill, 1873) (Homonym) 119
Pista cristata (Miiller, 1776) 119
Pista palmata (Verrill, 1873) 119
Pista quadrilobata (Augener, 1918) 120
Loimia medusa (Savigny, 1818) 120
Loimia viridis Moore, 1903 120
Enoplobra)\chus sanguineus (Verrill, 1873) 120
Lysilla alba Webster, 1879 120
Amaeana trilobata (Sars, 1863) 122
Amaeana aecraoisis (Augener, 1918) 122
Polycirrus carolinensis new species 122
Polyeirrus eximius (Leidy, 1855) 123
Polycirrus eximius dubius new subspecies 123
Family Sabellidae 124
Key to genera and species 124
Branchiomma nigromaculata (Baird, 1865) 125
Hypsicomus phaeotaenia (Schmarda, 1861) 125
Potamilla (Potamethus) spathiferus (Ehlers, 1887) new combination 125
Potamilla reniformis (Leuckart, 1849) 126
Megalomma lobiferum (Ehlers, 1887) 126
Megalomma bioculatum (Ehlers, 1887) 126
Sabella melanostigma Schmarda, 1861 126
Sabella microphthalma Verrill, 1873 127
Jasmineira bilobata new species 127
? Oriopsis sp 127
Chone americana new species 129
Family Serpulidae 129
Key to genera and species 129
Filograna implexa Berkeley, 1828 131
Pomatoceros americanus new species 131
Serpula vermicularis granulosa Marenzeller, 1884 131
Spirobranchus giganteus (Pallas, 1766) 132
Hydroides crucigera Morch, 1863 132
Hydroides parva (Treadwell, 1901) 132
Hydroides sanctaecrucis (Morch, 1863) 132
Hydroides dianthus (Verrill, 1873) 132
Hydroides uncinata (Philippi, 1844) 132
Hydroides protulicola Benedict, 1887 134
Metavermilia multicristata (Philippi, 1844) 134
Vermiliopsis annulata (Schmarda, 1861) 134
Spirorbis (Circeis) spirillum Linnaeus, 1767 135
•Spirorbis (Janua) corrugatus (Montagu, 1803) 135
Literature Cited 135
Figures
Page
1. Aphrogenia sp. a, stouter type of notoseta; b, neuroseta. (Figure c and
d omitted.) Lepidasthenia sp. e, slender superior neuroseta; f, inferior
neuroseta. Subadyte pellucida g, neuroseta. Harmothoe sp A h,
notoseta; i, neuroseta. Harmothoe sp. B j, notoseta; k, neuroseta; 1,
anterior elytron. Psamuiolyce ctenidophora n. sp. m, head; n, 1st ely-
tron; o, 2d elytron; p, 6th elytron; q, superior falciger; r, middle
falciger; s, inferior falciger; t, anterior view of parapodium 8
2. Sthenelais sp. a, anterior elytron; b, posterior elytron; c, anterior view
of foot; d, simple falciger; e, pluriarticulate falciger. Sthenelais anoc-
ula n. sp. f, elytron; g, head; h, short-bladed falciger; i, pluriarticulate
falciger; j, anterior view of foot. Pisione remota k, 3d foot; 1, anterior
end; m, stout superior seta; n, slender superior seta; o, inferior seta. . 13
3. Protomystides bidentata a, anterior end; b, foot; c, superior view of
shaft -head with blade removed; d, lateral view of shaft -head ; e, infer-
ior seta; f, superior seta. Lugia rarica g, anterior end; h, foot; i, shaft-
head of seta; j, seta. Hesionura elongata k, anterior end; 1, foot; m,
seta. Phyllodoce panamensis n, foot; o, shaft-head of seta; p, anterior
end 21
4. Syllis regulata carolinae n. subsp. a, head; b, aciculum; c, posterior com-
pound seta; d, superior simple seta; e, inferior simple seta; f, foot.
Odontosyllis longiseta n. sp. g, head; h, pharynx slit dorsally and
flattened to show teeth; i, foot; j, seta. Pionosyllis cf. uraga k, foot
with longer dorsal cirrus; 1, superior seta; m, inferior seta. Syllis
(Langerhansia) ferrugina n, posterior foot; o, inferior seta of an an-
terior foot; p, inferior seta of a posterior foot; q, superior seta 31
5. Websteri)iereis tridentata a, head; b and c, dorsal and ventral views of
proboscis; d, anterior foot; e, posterior falciger; f, posterior foot of
juvenile. Nereis riisei g and h, dorsal and ventral views of proboscis;
i, anterior foot; j, notopodial falciger. Nereis lamellosa k and 1, dorsal
and ventral views of proboscis; m, anterior foot; n, notopodial falci-
ger; o, posterior foot. Nephtys (Aglaophamus) circinata p, anterior
end ; q, anterior view of foot 40
6. Hemipodus roseus a, jaw and support (black); b, proboscideal papillae;
c, posterior view of foot. Glycera asymmetrica n. sp. d, jaw and sup-
ports (black); e, proboscideal papillae; f, posterior view of anterior
foot; g, posterior view of posterior foot 46
7. Goniadides carolinae n. sp. a, head with proboscis partly extruded; b,
posterior view of anterior foot; c, posterior view of posterior foot; d,
end of proboscis with jaws; e, hooked posterior notoseta; f, spiniger-
ous seta; g, falcigerous seta; h, section of proboscis with papillae.
XI
Glycinde nordmanni i, anterior foot; j, posterior foot; k, acicular
notoseta. Goniada teres 1, proboscideal papilla; m, posterior view of
anterior foot; n, posterior view of posterior foot 50
8. Rhamphobrachium atlanticum n. sp. a, lateral view of anterior end; b,
posterior view of 1st foot; c, posterior view of 40th foot; d, comb-seta;
e, winged capillary seta; f, tip of enlarged aciculum of 2d foot; g,
pseudocompound seta of 2d foot; h, bidentate acicular seta 56
9. Lumbrineris cruzensis a, anterior end; b, maxillae; c, mandibles; d,
compound hook; e, simple hook; f, winged capillary seta; g, anterior
view of anterior foot; h, anterior view of middle foot; i, anterior view
of far posterior foot 61
10. Polydora caeca a, enlarged hook of setiger 5; b, winged capillary seta;
c, posterior notopodial spine; d, hooded hook. Spiophanes wigleyi e,
tridentate hooded hook from setiger 16 ; f, bidentate hooded hook from
a posterior foot; Prionospio cirrobranchiata g, anterior end; h, 8th
foot; j, 40th foot; k, hooded hook. Prionospio dayi 1, anterior end; m,
hooded hook; n, 5th foot with 4th gill; o, posterior foot; Microspio
pigmentata p, dorsal view of anterior end; q, ventral view of anterior
end; r, anterior view of 8th foot; s, anterior view of posterior foot;
t, hooded hook 71
11. Spio pettiboneae a and b, dorsal and ventral view of anterior end; c,
anterior view of anterior foot; d, hooded hook. Nerinides unidentata
n. sp. e, anterior end; f, anterior view of 6th foot; g, anterior view of
18th foot; h, inferior sabre seta; j, hooded hook. Magelona papilli-
cornis k, head; 1, anterior view of 6th foot; m, anterior view of abdomi-
nal foot; n, special seta from setiger 9; o and p, face view and profile
of abdominal hook. Magelona physillae q, head; r, anterior view of
6th foot; s, anterior view of abdominal foot; t, setae from setiger 9;
u and v, face view and profile of abdominal hook. Magelona sp. w,
head; x, anterior view of 5th foot; y, anterior view of abdominal foot; >
z, face view of abdominal hook 75
12. Dodecaceria corallii a, anterior end; b, spoon-shaped hook. Orbinia
americana n. sp. c, anterior end; d, posterior view of 10th foot; e,
thoracic hook; f, flail-seta from abdominal neuropodium; g, 6th ab-
dominal foot. Orbinia riseri h, anterior end; i, posterior view of 18th
foot; j, thoracic hook; k, flail-seta from abdominal neuropodium; 1,
posterior abdominal foot. Scoloplos capensis m, anterior end; n,
posterior view of 10th thoracic foot; o, thoracic hook; p, flail-seta
from abdominal neuropodium; q, 5th abdominal foot 85
13. Aricidea fauveli a, specialized posterior neuroseta. Aricidea cerrutii
b, specialized posterior neuroseta. Cirrophorus branchiatus c, special-
ized notoseta; Travisia parva n. sp. d, anterior segment; e, two pos-
terior segments; f, lateral view of body. Ophelina cylindricaudata g,
posterior end. Hyboscolex longiseta h, head; i, parapodium. Leio-
chrides pallidior j and k, face view and profile of hook. Notomastus
americanus n. sp. 1, cross-section of abdominal segment; m and n,
face view and profile of hook 97
14. Nicomache trispinata a, posterior end. Asyckis carolinae n. sp. b, head;
c, posterior end; d, hook from setiger 2; e and f, profile and face view
of posterior hook. Pherusa ehlersi n. sp. g, lateral view of entire worm;
h, lateral view of 8th segment showing papillae; i, neuropodial hook
from middle segment; j7 elongated neuropodial hook from setiger 4 . . . 106
Ml
15. Isolda pulchella a, lateral view of anterior end; b, dorsal view of bran-
chial region with gills cut short; c, notopodial hook of segment 4; d,
neuroseta of segment 3; e and f, edge-on and lateral views of thoracic
uncinus. Samythella eliasoni n. sp. g, anterior end; h, thoracic capil-
lary; i and j, edge-on and lateral view of thoracic uncinus; k, palea;
1, edge-on view of abdominal uncinus; m, posterior thoracic para-
podium. Ampharete parvidentata n. sp. n and o, edge-on and lateral
views of thoracic uncinus; p, abdominal uncinigerous pinnule; q and
r, edge-on and lateral views of abdominal uncinus; s, anterior end.
Ampharete americana n. sp. t, anterior end; u and v, edge-on and
lateral view of thoracic uncinus; w and x, edge-on and laterial view
of abdominal uncinus 114
16. Pista quadrilobata a, anterior end; b and c, edge-on and lateral views
of uncinus from first row. Amaeana trilobata d, thoracic capillary
seta; e, abdominal acicular seta; f, ventrolateral view of entire worm.
Amaeana accraensis g, plumose capillary seta. Polycimis eximius
h, ventral view of anterior end; i and j, edge-on and lateral view of
uncinus; k and 1, longer and shorter capillary setae. Polycirrus car-
olinensis n. sp. m and n, edge-on and lateral views of uncinus; o,
longer, smooth-bladed capillary; shorter plumose capillary. Polycirrus
eximius dubius n. subsp. q, profile of uncinus 121
17. Chone americana n. sp. a, dorsal view of entire worm; a', posterior end
of juvenile; b, ventral view of collar and lips; c and d, lateral and
edge-on views of thoracic hooks; e and f, edge-on and laterial view of
abdominal uncini; g, thoracic winged capillary; h, thoracic palea; j,
thoracic tapered capillary; k, winged capillary from anterior abdo-
men; 1, slender capillary from posterior abdomen. Jasmineira bilo-
bata n. sp. m and n, lateral and edge-on views of thoracic hook; o,
thoracic palea; p, thoracic winged capillary; q, abdominal capillary;
r, dorsal view of collar; s, ventral view of collar overlying ventral lips;
t, lateral view of worm; u and v, edge-on and lateral views of abdomi-
nal uncinus 128
18. Pomatoceros americanus n. sp. a, tube; b, flat form of operculum; c and
d, lateral and dorsal views of conical form of operculum; e, abdominal
neuroseta; f, thoracic uncinus. Serpula vermicularis granulosa g,
operculum. Hydroides uncinata h, operculum; i, lateral view of a
spine from the crown. Metavermelia multicristata j, dorsal view of
anterior end; k, operculum; 1, thoracic winged capillary; m, "seta
of Apomatus"; n and o, edge-on and lateral views of thoracic uncinus;
p, abdominal geniculate seta; q, edge-on view of abdominal uncinus . . 133
Table
Page
1. The environmental conditions at each station along the Beaufort Shelf
Transect 3
NEW POLYCHAETA FROM BEAUFORT,
WITH A KEY TO ALL SPECIES RECORDED FROM NORTH CAROLINA
By
JOHN H. DAY
Zoology Department, University of Cape Town
Rondebosch, South Africa
ABSTRACT
Over 6,000 polychaete worms belonging to 229 species were collected on a transect
running from the sandy shore near Beaufort. N.C., to the upper part of the continental
slope in 200 m. Eleven more species were collected from the shores of Beaufort Sound
and from grab samples in (00, 600, and 3,020 m off North Carolina. The whole collection
includes 19 new species, 2 new subspecies, and Hi new records for the United States.
These have been described. An examination of the literature revealed that a further 83
species had been recorded by earlier workers so that a total of 323 species of polychaete
worms are now known from North Carolina. Keys have been constructed to cover the
whole fauna, all original records have been listed, and references to good descriptions of
each species are given. During the course of the work several type specimens were
examined and this has resulted in certain changes in nomenclature and the redefinition
of certain genera in the families Orbiniidae, Flabelligeridae, and Ampharetidae.
INTRODUCTION
This study is based on material collected in
1965 while workingatthe Duke University Marine
Laboratory in Beaufort, N.C. The aim of the
main research work was to find the most suitable
method for analysing distribution patterns across
the continental shelf of North Carolina. The
results have since been published by Day, Field,
and Montgomery (1971). The data for the analysis
were obtained by sampling the benthic inverte-
brates along a line of 10 stations called the
Beaufort Shelf Transect, which ran from the
shore to 200 m on the continental slope. Over
15,000 specimens belonging to 619 species
were collected in this way and among them
were about 6,000 polychaete worms belonging
to 229 species. In order to identify them, all
the species that had previously been recorded
from North Carolina were listed and the litera-
ture was searched for good diagnostic descrip-
tions. It was soon found that more than half
the species from the transect were new to North
Carolina and quite a number of them were new
species. The new species and new records
were added to the list as they were identified
and eventually keys were constructed to cover
the whole fauna. As there is no comprehensive
work covering the warm water fauna of the
Atlantic coast of the United States, it is hoped
that this account of the fauna of North Carolina
may be of use to other workers.
A numbered list of papers containing original
records of Polychaeta from North Carolina is
given below and full references to these and
other papers consulted during the course of
the work will be found at the end of this report.
For the benefit of later workers it may be men-
tioned that several papers were found to be
particularly useful. An early paper by Andrews
(1891a) describes several new species. Hartman
(1945) describes many new species and gives
keys and ecological notes of these and the
other species recorded by earlier workers. In
all she deals with 104 species found on the
intertidal banks and shallow waters around
Beaufort. Hartman (1951). in her account of the
fauna of the Gulf of Mexico, describes a few
more species which occur in North Carolina.
Pettibone (1963a), in an account of the families
Aphroditidae through Trochochaetidae of New
England, lists several records from deeper
waters off North Carolina, and her keys and
descriptions are most useful. Most of the other
papers give lists of species without descriptions.
Clwo)ioIogical list of papa's containing original ra-nrds
of Pol gchacta fi'om North Carolina
1.
Stimpson. 18.76
13.
Hartman, 19.71
•5
Yerrill. 1878
14.
Pearse and Williams,
3.
Webster, 1879
19.71
4.
Wilson, 1882
15.
Wells, 1961
5.
Andrews. 1891a
16.
MaiiKLim, 1962
6.
Andrews, 1891b
17.
Pettibone, 1963a
i .
Wilson, 1900
18.
Wells, Wells, and
8.
Pearse, 1936
Cray, 1964
9.
Pearse, Humm, and
19.
Wells and Gray. 1964
Wharton, 1942
20.
McCloskey, 1970
(i.
Hartman. 1944a
21.
Day, Field, and
1.
Hartman, 194.7
Montgomery, 1971
■>
Hartman, 1947a
2'^
Foster, 1971
In the pages that follow, all the species re-
corded in these papers have been extracted
and added to the records obtained from the
present collections. For the sake of brevity, the
authority for each record is shown by a number
which refers to the list above. Thus a record by
Stimpson (1856) is shown by the figure 1, while
records of the 240 species obtained from the
present collection are shown by an asterisk. It
should be noted that such records were not all
made on the Beaufort Shelf Transect. In addi-
tion to the 229 species collected on the tran-
sect, I dredged a few species at the entrance to
Beaufort Inlet and collected others from the
shores of Pivers Island. I also wish to thank
Dr. Charlotte Mangum for the three species of
Maldanidae from the shoals in Beaufort Sound,
Dr. Larry McCloskey for many specimens from
corals in 6.5-18 m near Lookout Lighthouse,
Dr. Fred Grassle for specimens from 450 and
600 m on the continental slope, and Dr. Robert
Menzies for three interesting abyssal species
from 3,020 m off Beaufort. In all, 323 species
are now known from North Carolina; of these 19
are new species, 2 are new subspecies, 16 are
new records for the United States, and 105 are
new records for North Carolina.
The new taxa and new records for the United
States have been described and references to
one or more good descriptions have been given
for the rest. Some of the records are doubtful,
but it was felt that so long as the authority for
the record could be traced from the numbered
list of papers above, even these doubtful records
should be included for the sake of completeness.
For the nonspecialist, the most useful items in
this paper are the keys to the various families
and the distribution lists of the species. These
are as complete as possible. All the 323 re-
corded species have been included and a few
other species known from adjacent areas have
been added. They have been marked "no N.C.
record" but they will probably be found there
in the future. The keys make an initial separation
of the genera whose names are given in paren-
theses and the specific characters of the species
are then added. In some cases additional char-
acters have been added in square brackets to
exclude closely related species.
It is well known that the marine fauna of
North Carolina is subtropical and this is equally
true of the Polychaeta. Many tropical species
extend northward from Florida and the Carib-
bean and many Carol inean species extend
southward to the Gulf of Mexico. Then there
are a small number of cold-water Virginian
species which reach North Carolina. All of
these are what might be called North American
endemics, since many of the warmwater forms
also occur on the Pacific coasts of Mexico and
California. Polychaete worms are notoriously
widespread and, in addition to these endemics,
there are many species that extend across the
Atlantic to Europe and North West Africa. Others
are circumtropical or truly cosmopolitan.
Within the limits of North Carolina, there are
faunistic differences between the sounds and
the open sea and between different depth zones
on the continental shelf. In the shallow sounds
around Beaufort, the bottom varies from mud
to sand, there is little wave action and the
estuarine waters are more productive than those
of the open sea. The polychaete fauna of the
sounds includes many subtropical species which
occur at much deeper levels on the continental
shelf; more than half of them are restricted
to 10 m or more but here the specimens are
much smaller. It was at first thought that the
small specimens on the continental shelf were
juveniles but when ovigerous females of the
same size were found, it was realized that the
polychaete fauna of the continental shelf is
stunted. Whether this applies to other groups
beside the Polychaeta is uncertain but one
gains the impression that the water overlying
the continental shelf is not very productive.
The distribution at different depths was ob-
tained by an analysis of the records along the
Beaufort Shelf Transect. The 10 stations of
the transect were all sited on sand or sandy
mud at increasing depth intervals and run in a
straight line from the shore at Lookout Light-
house to the continental slope some 40 miles
out to sea. The environmental conditions at each
station are given in Table 1 and may be used
to supplement the depth range of the various
species marked with an asterisk in the systema-
tic section.
Polychaete worms represented 40% of the
whole benthic fauna. While a few species, such as
Nephtys picta were found at most of the stations,
the majority of the species are grouped in well-
marked zones at different depth intervals. Depth
itself is not thought to be the limiting factor but
rather the changes in other factors which are
correlated with depth. There is a well defined
but poor fauna on the open sandy shore; Scole-
lepis squamata is the dominant polychaete and
does not occur elsewhere either on the shel-
tered sand banks in the sounds or at deeper
levels in the open sea. The fauna of the Turbu-
lent Zone between 3 and 20 m included several
very common polychaetes such as Paleanotus
heteroseta, Goniadides carolinae, Magelona
papillicornis , and Macroclymene zonalis. The
fauna of the Outer Shelf between 40 and 120 m
included another group of common species,
the most abundant being Onuphis nebulosa and
Oioenia jusiformis. On the Upper Slope between
120 and 200 m the commonest polychaetes were
Lumbrineris cruzensis, Scoloplos capensis,
CJiaetozone setosa, and Notomastus latericeus.
Many scientists have helped me during the
course of this research. Apart from those that
I have mentioned earlier, I would like to thank
Dr. C. G. Bookhout, the Director of the Duke
University Marine Laboratory during 1965, my
assistant Mrs. Mary Potts Montgomery, and
many other friends in the laboratory. My par-
ticular thanks are due to Dr. Marian Pettibone
for advice during the writing of this report and
for the loan of many reprints and specimens
from the U.S. National Museum.
Dr. Nancy Foster gave me helpful advice in
advance of her publications on the Spionidae
and Dr. Olga Hartman sent me many specimens
for comparison. Further specimens were sent
by Dr. David George of the British Museum and
Table 1. — The environmental conditions at each station
along' the Beaufort Shelf Transect.
Bottom
Station
Loc
ation
Depth
(m)
Substrate
tempera-
No.
Lat N
Long W
ture (°C)
1
34° 37'
76°31'
(i
coS
8-27
■>
34° 37'
76° 31'
3
fS
8-24
3
34°36'
76° 30'
5
fS
8-24
4
34° 36'
76°29'
10
inS
9-23
5
34° 34'
76° 26'
20
coS
9-23
6
34° 27'
76°06'
Id
fS
11-25
7
34° 24'
75° 58'
so
fS
17-24
8
34° 23'
75° 55'
120
mS
14-21
9
34°23'
75° 53'
160
f$
14-22
10
34° 22'
75°52'
200
SM
12-21
1 The symbols used for the substrate types are: CO =
coarse, f = fine, m = medium, S = sand, M = mud.
Further details will hi' found in Day, Field, and Mont-
gomery (1971).
Dr. Helmet Zibrowius of Marseilles. To all these
workers I tender my thanks. The collections
were made during the tenure of a senior foreign
scientist's fellowship and I gratefully acknow-
ledge funds from grants 81-6264 and 81-6320
from the National Science Foundation.
Apart from certain reference specimens which
were sent to the Duke University Marine Labora-
tory, the whole collection has been donated to
the U.S. National Museum.
FAMILY APHRODITIDAE
Key to genera and species
1 Harpoon-shaped notosetae present. [Neurosetae spurred and
longer prong feathered on inner margin. Felt poorly devel-
oped. (Laetmonice). Fifteen pairs of elytra] L.filicornis
1 No harpoon setae 2
2 Neurosetae stout, acicular, not spurred. Felt well developed Aphrodita sp.
2' Neurosetae long and spurred, often with spines on longer
prong. Felt poorly developed. [Stouter notosetae not flat-
tened and serrated] Aphrogenia sp.
Laetmonice filicomis Kinberg, 1855
Aphrogenia sp.
Laetmonice filicorn is. -Fauvel, 1923: 36, Fig.
12 a-f. - Pettibone, 1963a: 11, Fig. 1. - Day,
1963b: 355.
Records. — Off Beaufort in 120-200 m (17,
21, *).
Distribution. — North Atlantic from Sweden
and Greenland south to the West Indies; ?
Australia; from the edge of the continental
shelf to abyssal depths.
Aphrodita sp.
Remarks. — The two juveniles collected may
belong to Aphrodita hastata Moore, reported by
Pettibone (1963a) from the Gulf of St. Lawrence
to Chesapeake Bay in 4-2,000 m.
Records. — Off Beaufort in 200 m (*).
Figiire la, b
Description. — Body 2 mm long, broadly oval.
Dorsal felt poorly developed. No eyes, no ocular
peduncles. Notosetae in a graded series; very
fine ones forming felt, stouter cylindrical forms
(Figure la) tapered to fine tips with a double
row of granules near end. No barbed setae nor
serrated sabre-setae. Neuroseta (Figure lb)
long, stout and spurred, with two to five spines
on longer limb.
Remarks. — The single specimen is a juvenile,
but the lack of barbed setae or sabre-setae
distinguishes it from both Hermonia and Ponto-
genia. It appears to be close to Aphrogenia
alba Kinberg, recorded from St. Thomas Island
in the West Indies, but it lacks the stout
notosetae with strongly curved tips figured
by Kinberg (1858) for a 15-mm specimen.
Record. — One juvenile off Beaufort in 200 m
(*).
FAMILY POLYNOIDAE
Key to genera and species
1 Lateral antennae inserted terminally at same level as median.
Presetal and postsetal lips of neuropodia subequal
1' Lateral antennae inserted ventrally below level of median.
Presetal lip of neuropodia longer than postsetal. [Fifteen
pairs of elytra covering most of short body]
2 Planktonic and probably a larval form. Cirrophores of dorsal
cirri elongated (Drieschia) D. pellucida
2' Benthonic forms. Cirrophores not elongated 3
3 Body with about 26 segments and 12 pairs of elytra. Notosetae
numerous (Lepidonbtus) 4
3' Body with more than 50 segments and 18 or more pairs of
elytra. Notosetae few or absent 5
4 Margins of elytra fringed and surface with minute rounded
microtubercles only L. sable vis
4' Margins of elytra fringed and surface with both macrotubercles
bearing 3-5 blunt projections and numerous microtubercles L. variabilis
5 A few fine notosetae. Neurosetae of middle segments include
1-2 giant setae. Elytra mottled, with a white central spot
and irregularly arranged on posterior segments Lepidametria commensalis
5' No notosetae at all. Neurosetae include fine blunt forms
superiorly and stouter bidentate forms interiorly, but no
giant setae. Elytra half brown, half white Lepidasthenia sp.
6 Neurosetae with a large spinous pocket at base of blade.
[(Subadyte). Notosetae stout with coarse serrations] S. pellucida
6' Neurosetae with basal serration smaller than distal ones 7
7 All neurosetae with unidentate tips 8
7' Most neurosetae with bidentate tips 10
8 Neurosetae mostly tapered to fine hairlike tips. [Body with
less than 40 segments (Antinoella) . Anterior pair of eyes
larger than posterior pair] A. sarsi
8' Neurosetae with strong, often curved tips 9
9 Body with 50-80 segments. Few (4-10) stout notosetae. Elytra
translucent, not fringed, but covered with conical micro-
tubercles Harmothoe (Hermadion) aeanellae
9' Body with 35-49 segments. Notosetae numerous and finer
than neurosetae. Elytra fringed with papillae and surface
covered with microtubercles Gattyana cirrosa
10 Elytra with a well developed marginal fringe 11
10' Elytra without a marginal fringe or with merely a few
small marginal papillae. [Surface covered with numerous
microtubercles and a few ovoid macrotubercles at posterior
margin. Eyes anteroventral] Harmothoe imbricata
11 Elytron surface divided into polygonal areas bearing large
pointed tubercles Harmothoe aculeata
11 ' Elytron surface not divided into polygonal areas and bears
only microtubercles. [Notosetae few and stout.] Harmothoe trimaculata
Drieschia pellucida Moore, 1903
Drieschia pellucida Moore, 1903: 794, pi. 55:
Fig. 1-12. - Pettibone, 1963a: 22. Fig. 5h-j.
Records. — Planktonic in the Gulf Stream (17).
Distribution. — Gulf Stream from Bermuda
to Massachusetts; 0-1,800 m.
Lepidonotus sublevis Verrill, 1873
Lepidonotus sublevis. - Hartman, 1942a: 22,
Fig. 7-12. - Pettibone, 1963a: 18, Fig. 3 e.
Records. — Several records between 0 and
100 m off North Carolina (5, 11, 15, 17, 18, 21, *).
Distribution. — Massachusetts to Florida and
the Gulf of Mexico; intertidal to 100 m. Often
commensal with hermit crabs.
Lepidonotus variabilis Webster, 1879
Lepidonotus variabilis Webster, 1879: 205. -
Hartman, 1951: 18 (notes).
Remarks. — This species is close to L. tenui-
setosus Gravier from the Red Sea.
ReeonU. — Many records from the shore to
18 m off North Carolina (3, 5, 11, 13, 15, 18, 20).
Distribution. — Virginia to the West Indies;
intertidal to a few meters.
Lepidametria commensalis Webster, 1879
Lepidametria commensalis. - Seidler, 1924: 148.
- Hartman, 1945: 10; 1951: 17. - Pettibone,
1963a: 19, Fig. 4 k.
Remarks. — In Day (1962: 634) I gave my
reasons for regarding Lepidametria as a syno-
nym of Lepidasthenia. Dr. Pettibone, who is
making an intensive study of the Polynoidae,
informs me that Lepidametria is a valid genus
and in deference to her opinion I have not
changed the name of Lepida m et ria commensalis.
Records. — Many records from the shore to
24 m off North Carolina (3, 5, 11, 13, 15, 17, 18).
Distribution. — Massachusetts to Florida and
the Gulf of Mexico; intertidal to 24 m.
Lepidasthenia sp.
Figure le, f
Description. — Lateral antennae about equal
to median and 1.7 times prostomial length.
Occipital flap semicircular with a smooth mar-
gin. Elytra smooth, half brown and half pale,
not large enough to cover middle of dorsum.
Notosetae absent. Superior neurosetae (Figure
le) slender, ending in blunt tips; inferior neuro-
setae (Figure If) stout and bidentate with a
small secondary tooth. Ventral margins of para-
podia without papillae.
Remarks. — Only a single incomplete speci-
men with 23 segments was obtained. It resem-
bles L. bruunea Day, from South Africa in the
pigment pattern on the elytra, in the absence
of notosetae and in the shape of the neurosetae.
It differs in having a well-developed occipital
flap and the lack of a row of papillae on the
ventral margins of the neuropodia. It may be
noted that Lepidametria bruunea Knox, (1960:
91, Fig. 58-63) is also similar. The description
of Knox's species was published later than that
of Day.
Records. — Off Beaufort in 120 m; one speci-
men (*).
Subadyte pellucida (Ehlers, 1864)
Figure lg
Scalisetosus pellucidus. - Fauvel, 1923: 74, Fig.
27 a-f.
Scalisetosus fragilis. - Day, 1967: 59, Fig. 1.7.
g-k.
Subadyte pellucida. - Pettibone, 1969: Fig. 4a-e.
Description. — Body 8-15 mm long with about
45 segments, very fragile and mottled with
greenish grey. Prostomium bilobed but without
anterolateral peaks. Lateral antennae inserted
ventrally at a lower level than median. All three
antennae much longer than prostomium. Fif-
teen pairs of large, delicate and deciduous elytra
covering entire body. Individual elytra with
minute marginal papillae and conical micro-
tubercles scattered over surface. Dorsal cirri
long with slender tips. Notosetae fairly stout
with a few coarse serrations and blunt tips.
Neurosetae (Figure lg) long and very trans-
parent with an enlarged serration forming a
pocket at base of blade, numerous faint serra-
tions along its length and a hooked tip with a
minute secondary tooth.
Records. — Five small specimens from 40 to
120 m off Beaufort (*). This is a new record for
the United States.
Distribution. — Eastern Atlantic from Scot-
land, Madeira Island, and Morocco to South
Africa; Mediterranean; Indian Ocean; inter-
tidal to 40 m.
Harmothoe imbricata (Linnaeus, 1767)
Harmothoe imbricata. - Fauvel, 1923: 55, Fig.
18 f-1. - Pettibone, 1963a: 36, Fig. 7 a-d.
Records. — Intertidal in the Cape Hatteras
area (18).
Distribution. — Arctic; North Atlantic from
Norway south to the Mediterranean and Labra-
dor to North Carolina; Pacific from the Bering
Sea to southern California and Japan to the
Yellow Sea; intertidal to 300 m.
Antinoella sarsi (Malmgren, 1865)
Antinoella sarsi. - Pettibone, 1963a: 30, Fig.
7e-j.
Records. — Two juveniles off Beaufort in 200
m (*).
Distribution. — Arctic and North Atlantic
from Norway to the British Isles and Labrador
to North Carolina; Pacific from the Bering Sea
to North Japan; 5-2,000 m.
Harmothoe (Hermadion) acanellae
(Verrill, 1881)
Harmothoe aeuleata Andrews, 1891
Harmothoe aeuleata. Andrews, 1891a: 278, pi.
12: Fig. 1-5. - Hartman, 1945: 10. -
1951: 19. - Nonato and Luna, 1970a: 67,
pi. 3: Fig. 26.
Records. — Several records from the shore
and shallow dredgings off North Carolina (5, 7,
11, 13, 14, 15).
Distribution. — North Carolina to Florida and
Brazil; intertidal to 69 m.
Harmothoe trimaculata (Treadwell, 1924)
Polynoe acanellae. — Hartman, 1942a: 27, Fig.
' 27-31.
Harmothoe (Hermadion) acanellae. - Pettibone,
1963a: 28, Fig. 6 1, m.
Records. — North Carolina, ? depth (17).
Distribution. — North Atlantic from Denmark
to West Greenland and south to North Carolina
in 4-2,000 m associated with the coral Acanella
normani.
Gattyana cirrosa (Pallas, 1766)
Harmothoe trimaculata Treadwell, 1924: 6. -
Hartman, 1938: 118, Fig. 38 a, 39 a, b;
1951: 19.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 70 m (18, 19).
Distribution. — North Carolina and West In-
dies; 0-70 m.
Harmothoe sp. A
Figure lh, i
Gattyana cirrosa. - Fauvel, 1923: 49, Fig. 17 a-f.
Pettibone, 1963a: 28, Fig. 5 b-d.
Records. — Beaufort, intertidal (5).
Distribution. — Arctic; North Atlantic from
Norway to France and Hudson Bay to South
Carolina; Pacific from the North Japan Sea
and Bering Straits to Washington; intertidal
to 1,153 m.
Description. — Body 3-4 mm long, with 21-23
segments and 10 pairs of elytra. Pigmentation
characteristic with dark brown blotches on
cirrophores of setigers 6 and 10. Prostomium
bilobed but without cephalic peaks and anterior
eyes well forward and ventral. Median antenna
three times prostomial length, laterals ventral
in origin and shorter than prostomium. An-
tennae and cirri not visibly papillose. Elytra
Figure 1. — Aphrogenia sp. a, stouter type of notoseta; b, neuroseta. (Fignre c and d omitted.) Lepidasthenia sp.
e, slender superior neuroseta; f, inferior neuroseta. Subadyte pellucida g, neuroseta. Harmothoe sp. A h, notoseta;
i, neuroseta. Harmothoe sp. B j, notoseta; k, neuroseta; 1, anterior elytron. Psammolyce ctenidophora n. sp.
m. head; n, 1st elytron; o, '2d elytron; p, Cth elytron; q, superior falciger; r, middle falciger; s, inferior falciger;
t, anterior view of parapodium.
delicate, without marginal fringes and surface
with only a few weak tubercles bearing four
to six spinules. Notosetae deciduous leaving
only a stout aciculum in the notopodia; when
present, notosetae very fine and serrated to
their hairlike tips (Figure lh). Neurosetae
(Figure li) slender, slightly stouter than noto-
setae and bearing long blades ending in naked
tips with two subequal teeth.
Remarks. — The size and number of segments
shows that all specimens are juveniles. The very
fine setae and pigmentation are quite distinctive.
Records. — Off Beaufort in 35 m (*).
Harmothoe sp. B
Figure lj-1
Description. — Body pale in alcohol, 4.5 mm
long with less than 25 setigers and 8-11 pairs
of elytra. Prostomium with small cephalic peaks
and anterior pair of eyes lateral. Lateral an-
tennae half as long as prostomium. Antennae
and dorsal cirri with short clavate papillae.
Elytra (Figure 11) mottled gray-green with a
central dark spot, not fringed but with a few
soft papillae and a scattering of hemispherical
weakly chitinized microtubercles on surface.
Notosetae (Figure lj), stout and strongly ser-
rated to their blunt, grooved ends. Neurosetae
(Figure Ik), more slender than notosetae and all
with bidentate tips.
Remarks. — These juvenile specimens may be
Harmothoe dearborni Pettibone, but the neuro-
setae have a much longer secondary tooth and
the body lacks a rusty red coloration.
Records. — Six juveniles off Beaufort in 35 m
(*).
Distribution. — (of H. dearborni). On Sargas-
sum weed floating off Massachusetts.
FAMILY POLYODONTIDAE
Key to genera and species
Anterior pair of eyes large and mounted on stalks. Three
antennae. Branchial vesicles between anterior parapodia.
Superior neurosetae not penicillate (paintbrush shaped)
(Polyodontes) P. lupina
Anterior pair of eyes sessile like posterior pair. Three an-
tennae. No branchial vesicles. Superior neurosetae not peni-
cillate (Eupanthalis) E. kinbergi
Polyodontes lupina (Stimpson, 1856)
Polyodontes lupina. — Hartman, 1945: 10; 1951:
19.
Records. — Beaufort, intertidal to 160 m (1,
5, 11, 13, *).
Distribution. — North Carolina; South Caro-
lina; Gulf of Mexico; intertidal to 160 m.
Eupanthalis kinbergi (Mcintosh, 1876)
Euarche tubifex Ehlers, 1887: 54, pi. 12: Fig.
1-7, pi. 13: Fig. 1.
E)ipanthalis kinbergi. - Fauvel, 1923: 100, Fig.
38 i-q. - Day, 1967: 94, Fig. 1.17. a-f.
Records. — One specimen from 450 m off
Beaufort (*).
Distribution. — Gulf of Mexico; North Atlan-
tic (Adventure Bank); off Angola; Mediterra-
nean; 64-1,000 m.
FAMILY SIGALIONIDAE
Key to genera and species
1 Body seldom longer than 10 mm. No cirriform branchiae on
elytrophores {Pholoe). [Elytra without concentric growth
lines. Forty -five segments or more] P.minuta
1 Body up to 200 mm long. Cirriform branchiae on elytrophores
after first few segments 2
2 No median antenna; lateral antennae papilliform (Sigalioii).
[Elytra with 8-13 branching papillae on outer margin; one
stylode on notopodium and two on neuropodium] S. arenicola
2' Median antenna present; lateral antennae indistinct and fused
to base of tentacular segment 3
3 Compound neurosetae spinigerous with laddered blades ending
in pointed tips. [Median antenna without ctenidia (Leanira).
No eyes, no simple neurosetae; elytra without marginal
papillae or lateral indentations] L. hystricis
3' Compound neurosetae mainly or entirely falcigerous with
simple or multiarticulate blades ending in bidentate tips 4
4 Median antenna with a large ceratophore. Elytra and dor-
sum encrusted with sand (Psammolyce). [Ctenidia on cera-
tophore of median antenna] P. ctenidophora
4' Median antenna with a short ceratophore. Elytra not en-
crusted with sand (Sthenelais) 5
5 Eyes present. Usually a few simple bipectinate setae at superior
edge of neuropodium 6
5' Eyes absent. No simple neurosetae. [Elytra without markings] S. anocula
6 External margins of elytra fringed with simple papillae, never
notched S. boa
6' External margins of anterior elytra with a few irregular
papillae and posterior ones with a notch S. limicola
Pholoe minuta (Fabricius, 1780) Sigalion arenicola Verrill, 1879
Pholoe minuta. - Fauvel, 1923: 120, Fig. 44 a-h. Sigalion arenicola. - Pettibone, 1963a: 48, Fig.
-Pettibone, 1963a: 46, Fig. 10 f, g. - Day, 11 a, b. - Nonato and Luna, 1970a: 72,
1967: 100, Fig. 1. 18. a-f. pi. 4: Fig. 46-48.
Records. — One specimen from 160 m off Records. — Three specimens from 20 m off
Beaufort (*). Beaufort (*).
Distribution. — Cosmopolitan; intertidal to Distribution. — Massachusetts to Georgia;
2,295 m. Brazil; intertidal to 37 m in sand.
10
Leanira hystricis Ehlers, 1874
Leanira hystricis. - Fauvel, 1923: 118, Fig.
43 h-m. - Pettibone, 1970a: 8, Fig. 4.
Description. — Median antenna short and with-
out ctenidia. Lateral antennae indistinct, prob-
ably fused to bases of tentacular segment. No
eyes. Palps long with basal sheaths. A cirro-
phore but no dorsal cirrus on segment 3. Elytra
without surface papillae, marginal fringes, or
lateral indentations. Cirriform branchiae from
segment 7 onwards. Notopodia with two long-
apical stylodes. Neuropodia with five long sty-
lodes, two arising from posteroventral bract.
No simple pectinate setae in neuropodia, all
neurosetae being compound with spinigerous
laddered blades.
Remarks. — Pettibone (1963a) and earlier
workers recorded Leanira hystricis from Massa-
chusetts to north of Puerto Rico. Pettibone
(1970a) has revised the synonymy of L. hys-
tricis and several other species and states that
the record from Massachusetts refers to L.
robusta Verrill and the record from north of
Puerto Rico refers to L. cirrata (Treadwell).
As noted above, the present specimens lack
simple neurosetae which agrees with Pettibone's
revised description of L. hystricis and dis-
tinguishes it from both L. robusta and L. cirrata.
On the other hand, it should be noted that the
presence or absence of labial lobes which Petti-
bone now regards as an important diagnostic
character was not recorded.
Records. — Seven specimens from 200 m off
Beaufort (*).
Distribution. — (According to Pettibone,
1970a) northeastern Atlantic; Iceland, United
Kingdom, Azores; 957-2,640 m.
Psammolyce ctenidophora New Species
Figure lm-t
Holotype.— USNM 43117.
Description. — Body incomplete with only 28
segments measuring 25 mm by 8 mm. Dorsum
covered with coarse sand or shell fragments
attached to elytra and mid-dorsum by branch-
ing adhesive papillae (Figure lm). Ventrum and
parapodial bases densely covered with segmental
bands of long slender papillae alternating with
narrower bands of small hemispherical papillae.
Prostomium (Figure lm) rounded but sunken
between anterior parapodia and dominated by
the large swollen ceratophore of median an-
tenna. Base of ceratophore with a pair of small
ctenidial flaps; ceratostyle missing. Anterior
pair of eyes much larger than posterior pair
and directed forwards. Lateral antennae as bi-
articulate conical projections arising from dor-
sal bases of tentacular lobes and not visibly
joined to prostomium. Tentacular lobes fused
basally below ceratophore of median antenna
and each bearing two tufts of slender, serrated
capillary setae protected on medial side by
cephalic sheaths. Dorsal tentacular cirri slightly
longer than ventral ones. A large pedunculate
facial tubercle above mouth and large prebuccal
flanges on either side; palps missing.
Elytra on setigers 2, 4, 5, 7, 9 . . . 27, and 28
(broken end) and dorsal tubercles on intervening
segments from setiger 3 onwards. First pair of
elytra (Figure In) large, oval, without an inci-
sion and covering head. Subsequent elytra small-
er and widely separated leaving dorsum bare
apart from sand grains. Second pair of elytra
(Figure 10) triangular and without lappets but
subsequent elytra (Figure lp) somewhat pro-
duced medially and with small lappets on pos-
terior margin. Exposed surfaces of all elytra
covered with long papillae and adherent sand
grains or shell fragments.
Parapodia (Figure It) similar throughout;
those of setigers 2 and 3 without specialized
features. Each notopodium short and stout with
a presetal flap dorsally. Neuropodium larger,
with scattered squat papillae and groups of long
filiform papillae; three groups around neuro-
setae and fourth group at base of ventral cirrus.
Notosetae slender and minutely serrated, many
directed downwards between parapodia. Neuro-
setae of three types; a superior group of stout
falcigers with well serrated shaft-heads (Figure
lq) and small blades with an indication of a
secondary tooth; a middle group of stout falci-
gers with almost smooth shaft-heads (Figure
lr); an inferior group of slender falcigers with
serrated shaft-heads and long bidentate blades
(Figure Is).
Remarks. — Species of the genus Psammolyce
have been distinguished mainly on the shape of
the elytra, some of which are incised while
11
others have long projecting lobes. As Willey
(1905) and Potts (1909) have remarked, these
features are variable for the elytra may change
along the length of the body in a single speci-
men. For this reason, P. ctenidophora is named
as a new species with hesitation. However, it
does not agree with the species described by
Fauvel (1923) from Europe, or those described
by Mcintosh (1885), Treadwell (1902), Augener
(1906). Hartman (1939), Hartman (1942b) or
Hartman (1965a) from the West Indies. The
presence of ctenidial flaps on the peduncle of
the median antenna of P. ctenidopliora appears
to be unique and the large pedunculate facial
tubercle has not been noted on any other species.
Records. — One specimen off Beaufort on rock
and sand in 20 m (*).
Sthenelais boa (Johnston, 1833)
Sthenelais boa. - Fauvel, 1923: 110, Fig. 41 a-1.
-Pettibone, 1963a: 50, Fig. 10 a-d. - Day,
1967: 109, Fig. 1.20.M.
Records. — Several records between Cape Hat-
teras and Beaufort from the shore to 100 m
(5,7, 11, 13, 17, 18).
Distribution. — Cosmopolitan in temperate
and tropical seas; intertidal to 150 m.
Sthenelais limicola (Ehlers, 1864)
Sthenelais limicola. - Fauvel, 1923: 113, Fig.
42 a-g. - Pettibone, 1963a: 51, Fig. 11 c-e. -
Day, 1967: 111, Fig. 1.20. m-r.
Records. — Common off Beaufort in 20-80 m
(17,*).
Distribution. — Atlantic from Norway to Spain
and the Gulf of St. Lawrence to North Carolina;
Mediterranean; South Africa; intertidal to
800 m.
Sthenelais sp.
Figure 2 a-e
Notes. — Apart from the typical specimens of
S. limicola recorded above, 10 aberrant speci-
mens were obtained which are closer to S. minor
Pruvot and Racovitza as described by Fauvel
(1923). In particular, there are no bipectinate
simple setae in the neuropodia. The anterior
elytra (Figure 2a) have no attached sand grains
but bear a few conical microtubercles on the
surface and a few simple papillae on the external
margin. Posterior elytra (Figure 2b) have a
small excision on the external margin. The feet
(Figure 2c) and the compound neurosetae (Fig-
ure 2d, e) are identical with those of S. limicola.
According to Fauvel, S. minor lacks simple
bipectinate neurosetae and the excision in the
posterior elytra, both of which are character-
istic of S. limicola. More material is required
to decide whether these American specimens
represent a new species or whether S. minor
is a synonym of S. limicola.
Records. — Ten specimens off Beaufort in
20-80 m (*).
Sthenelais anocula New Species
Figure 2f-j
Sthenelais anocula Day, Field, and Montgomery,
1971: 113 (Nomen nudum)
Holotype. — USNM 43128; 9 paratypes,
USNM 43129.
Description. — Body up to 35 mm long with
about 60 segments and uniformly pale in alcohol.
Prostomium (Figure 2g) rounded and without
eyes. Median antenna with a pair of small
bracts or ctenidia on ceratophore and an elon-
gated ceratostyle tapering to a slender terminal
filament. Lateral antennae not distinguished
but presumably fused to base of tentacular
segment which bears a ciliated cushion similar
to those on later notopodia. Superior tentacular
cirrus as long as median antenna but inferior
cirrus much shorter. Two setigerous lobes each
with a single terminal stylode. Palps missing
and their sheaths united to presetal bracts on
either side of mouth.
Setiger 3 with a dorsal tubercle but no
dorsal cirrus. Elytra on segments 2, 4, 5, 7,
9 . . . , and alternate segments to 27th and all
subsequent segments. Small cirriform branchiae
on all elytrophores and dorsal tubercles from
segment 4. Elytra broadly oval (Figure 2f),
with few cushion-shaped papillae on surface,
and about 15 simple digitiform marginal papil-
lae; no external notch even on posterior elytra.
12
Figure 2. — Sthenelais sp. a, anterior elytron; b, posterior elytron; c, anterior view of foot; d, simple falciger; e,
pluriarticulate falciger. Sthenelais anocula n. sp. f, elytron; g, head; h, short-bladed falciger; i, pluriarticulate
falciger; j, anterior view of foot. Pisione remota k, 3d foot; 1, anterior end; m, stout superior seta; n, slender
superior seta; o, inferior seta.
1::
Notopodia (Figure 2j) with three ciliated cush-
ions dorsally, three terminal stylodes on first
few feet but only one stylode from 10th foot.
Neuropodia with three small perisetal bracts;
one superior with an apical stylode, one antero-
inferior without a stylode and one postero-
inferior with an apical stylode. Third stylode
at apex of setigerous lobe. Sides of parapodia
without long papillae.
Notosetae as long, simple capillaries, mi-
nutely serrated on one margin. Neurosetae all
compound, simple bipectinate setae being
absent. Compound setae (Figure 2h, i) all with
smooth shaft -heads; a few with short bidentate
blades, many with long multiarticulate blades.
Remarks. — S. anocula is generally similar
to S. limicola but is distinguished from the
latter by the lack of eyes, the lack of simple
bipectinate neurosetae and the absence of an
incision on the posterior elytra.
Records. — Fifteen specimens on sandy mud
80-200 m off Beaufort (21, *).
FAMILY PALMRIDAE
Key to genera and species
Body elongate, completely covered by transverse rows of
paleae. Prostomium concealed under a fold of skin {Bhaica-
nia). Paleae with about 20 ribs including 5 major beaded
ones
Body rectangular with right and left groups of paleae distinct.
Prostomium visible between paleae (Paleanotus) . Paleae
with about 20 subequal beaded ribs
B. goodei
P. lieteroseta
Bhawania goodei Webster, 1884
Paleanotus lieteroseta Hartman, 1945
Bhawania goodei Webster, 1884: 308, pi. 7: Fig.
10-15. - Day, 1967: 118, Fig. 2.1. a-f.
Paleanotus lieteroseta Hartman, 1945: 12, pi. 1:
Fig. 1-6.
Records. — Common on corals in 5-20 m off
North Carolina (14,20, *).
Distribution. — Circumtropical; intertidal to
30 m in rock crevices and coral.
Records. — Common off Beaufort on sand at
4-20 m (11,21, *).
Distribution. — North Carolina in 4-20 m.
FAMILY PISIONIDAE
Only one genus and species known from
North Carolina.
Pisione remota (Southern, 1914)
Figure 2k-o
Praegeria remota Southern, 1914: 61, pi. 7,
pi. 8: Fig. 15 a-k. - Fauvel, 1923: 124,
Fig. 45 a-g.
Pisione remota. - Hartman, 1968: 181, Fig. 1-5.
I>i \crvption. — Juveniles up to 15 mm long
with slender bodies and long projecting para-
podia. Prostomium (Figure 21) inconspicuous
and embedded in large peristome between bases
of long, forwardly directed palps. Above them,
two pairs of biarticulate tentacular cirri; inner,
ventral pair minute and papilliform, outer dor-
sal pair long and slender. Peristomial acicula
long and stout, their flanged ends projecting
in front of mouth as supernumary jaws. Pro-
boscis eversible with 14 marginal papillae and
2 pairs of weakly chitinized true jaws normally
retracted back to setiger 4. Two fused pairs of
1 I
subdermal eyes at level of setiger 2. Setiger 1
with a small papilliform dorsal cirrus, a setig-
erous lobe and a long ventral cirrus directed
forwards. Dorsal cirrus of setiger 2 not en-
larged. Each normal foot (Figure 2k) with a
minute Particulate dorsal cirrus, a long setig-
erous lobe with two presetal lips and a ventral
cirrus similar to dorsal one but more distal
in origin. Setae of three types; superior one
(Figure 2m) stout, simple, and obliquely trun-
cate; second (Figure 2n) simple, with a slightly
curved and pointed end; below this three com-
pound setae (Figure 2o) with short falcigerous
blades.
Remarks. — P. remota differs from the type
species P. oerstedi Grube in lacking an elongate
dorsal cirrus on setiger 2 and differs from P.
africana Day in having all blades of the com-
pound setae short.
Records. — Juveniles common off Beaufort in
10-20 m (21, *). This is a new record for the
Atlantic coast of the United States.
Distribution. — Ireland; English Channel;
Mediterranean; southern California; Pacific
coast of Mexico; in 10-200 m.
FAMILY AMPHINOMIDAE
Key to genera and species
1 Notosetae in transverse palisades across dorsum. [Body stout
and oval. Caruncle linear (Euphrosine)] No N.C. record
1' Notosetae in compact tufts 2
2 Branchiae with regular bipinnate branches. [Body oval with
about 30 segments. Caruncle broad with pleated margins
(Chloeia). Conspicuous red bars on dorsum when adult] C. viridis
2' Branchiae as irregularly branching tufts 3
3 Neuropodium small and suckerlike with few short, hooked
setae (Hipponoe) H. gaudichaudi
3' Neuropodium well developed; setae numerous, not hooked 4
4 Neurosetae stout and acicular. Body large, stout, almost
quadrangular in section (Amphinome) . Caruncle small,
cordate A. rostrata
4' Neurosetae normal, spurred. Body small, less than 30 mm.
Ventrum flattened. Caruncle small or absent. [Branchiae
not present on posterior segments] 5
5 Notosetae of setiger 1 include stout hooks (not easily seen).
(Para m phi no m e pulchella) No N.C. record
5' Notosetae of setiger 1 without hooks. (Pseudeurythoe) . [Car-
uncle rudimentary. Branchiae from setiger 3] P. ambigua
Chloea viridis Schmarda, 1861
Chloeia euglochis Ehlers, 1887: 18, pi. 1: Fig. 1,
2, pi. 2: Fig. 1-8, pi. 3: Fig. 1-4.
Chloeia viridis. - Hartman, 1951: 29. - Nonato
and Luna, 1970b: 65, Fig. 1, 2.
Notes. — Body up to 117 mm long with 36-39
segments. Dorsum with a median dorsal stripe
in juveniles changing to violet brown segmental
bars in adults.
Records. — Cape Hatteras area on Sargassum
and off Beaufort in 40-120 m (18, *).
15
Distribution. — North Carolina; West Indies;
Gulf of Mexico; Brazil; low tide to 120 m.
Hipponoe gaudichaudi Audouin and
Milne-Edwards, 1830
Hipponoe gaudichaudi. - Fauvel, 1923: 132,
Fig. 47 1-p. - Pettibone. 1963a: 57, Fig.
13 a, b.
Records. — Two specimens on driftwood with
Lepas, off Beaufort (*).
Distribution. — Cosmopolitan in warm and
tropical seas.
Amphinome rostrata (Pallas, 1776)
Amphinome pallasii. - Fauvel, 1923: 127, Fig.
46.
Amphinome rostrata. - Hartman, 1951: 22,
pi. 4: Fig. 1. - Pettibone, 1963a: 59, Fig.
13 d, e. - Day, 1967: 123, Fig. 3.1. f-k.
Records. — Several records from driftwood
cast ashore in North Carolina (5, 7, 11, 13, 17,
18,*).
Distribution. — Cosmopolitan in warm and
tropical seas.
Paramphinome pulchella (Sars, 1872)
Paramphinome pulchella. - Pettibone, 1963a:
61, Fig. 13 f, g (with synonymy).
Paramphinome jeffreysii. - Hartman, 1965a:
58, pi. 1: Fig.b, c.
Records. — No North Carolina record, but
known from south of Long Island and the Gulf
of Mexico. (17).
Distribution. — North Atlantic from Norway
to Denmark and Iceland to the Gulf of Mexico;
from 37 to 5,500 m.
Pseudeurythoe ambigua (Monro, 1933)
Eurythoe ambigua Monro, 1933b: 6, Fig. 2.
Pseudeurythoe ambigua. - Hartman, 1945: 12.
Remarks. — Parewythoe Gust&fsona,ndPseud-
eurythoe Fauvel are very close. The main dis-
tinction is that in Pareurythoe the gills con-
tinue to the end of the body while in Pseudeury-
thoe they are limited to anterior segments.
This, however, may be a function of size and
most species of the Pseudeurythoe are small.
Records. — Beaufort, intertidal, and common
at 20 m (11,21, *).
Distribution. — Pacific coast of Panama; North
Carolina; intertidal to 20 m.
FAMILY PHYLLODOCIDAE
Key to genera and species
NOTE. — In the tentacular formula used below, the symbol 1 = tapered tentacular cirrus;
N = normal lamellar cirrus; 0 = absent (no cirrus or no setae); S = setae.
1 Two pairs of tentacular cirri. Only 4 antennae 2
1' Three pairs of tentacular cirri. Only 4 antennae 5
1" Four pairs of tentacular cirri. Four or 5 antennae 6
2 Two tentacular cirri on first segment. Prostomium flattened
and body usually white (Eteone) 3
2' One tentacular cirrus on first segment and one on second
which also bears setae. Prostomium not flattened and body
usually greenish (Lugia). [No dorsal cirrus on segment 3;
formula: 1 + S -W S ^r] L. rarica
N N
16
3 Setae present on segment 2; formula: 0— — I- S — 4
3' No setae on segment 2; formula: 0-j~ + 0— . [Dorsal cirri
asymmetrical and broader than long] Eteone lactea
4 Dorsal tentacular cirrus shorter than ventral one. Posterior
dorsal cirri asymmetrical, longer than broad Eteone heteropoda
4' Dorsal tentacular cirrus as long as ventral one. Dorsal cirri
almost symmetrical and as broad as long (E. longa) No N.C. record
5 Three tentacular cirri on three segments with setae from
second; formula: 1 + S — + S — . (Protomystides). Dorsal
N N
cirri long and hastate P. bidentata
5' Three tentacular cirri on two segments with setae from third;
formula: 1 + 0 — - + S— ■ (Hesionura). Dorsal cirri fusi-
1 N
form and much shorter than ventrals H. elongata
6 Four frontal antennae and a similar middorsal one (Eidalia) 7
6' Four frontal antennae but no middorsal one, a minute occipital
papilla sometimes present 10
7 Setae on both second and third tentacular segments; formula:
1 + S — + S — . Setigerous lobe bluntly rounded apically 8
7' No setae on any tentacular segment; formula: 1 + 0— : — h 0 — .
Setigerous lobe with a pointed superior projection apically . . . . E. (Ptero cirrus) macroceros
8 First tentacular segment not visible dorsally, second and third
distinct (subgenus Eumida). Proboscis almost smooth E. (Eumida) sanguined
8' All three tentacular segments distinct and separate (subgenus
Eidalia). Proboscis densely covered with papillae 9
9 Dorsum yellow with a pair of lateral stripes. Dorsal cirri
bluntly oval E. (Eidalia) bilineata
9' Dorsum uniformly green. Dorsal cirri hastate, pointed E. (Eidalia) viridis
10 Prostomium with a median posterior projection embraced by
broad "shoulders" formed of fused first and second tenta-
cular segments. [Setae from third tentacular segment;
formula: 1 + 0— + S— (Paranaitis)] 11
1 N
10' Prostomium truncate or notched posteriorly. First and second
tentacular segments not forming broad "shoulders". (Phyl-
lodoce) 12
11 Dorsal cirri reniform Paranaitis kosteriensis
11' Dorsal cirri asymmetrically oval Paranaitis speciosa
17
12
12'
13
13'
1 1
14'
15
15'
16
16'
17
17'
Prostomium truncate posteriorly without an occipital papilla.
Tentacular segments 1 and 2 fused; setae from second;
formula: 1 + S— + S— . [Dorsal cirri cordate] 13
Prostomium notched posteriorly with a minute occipital papilla.
Tentacular segments all separate though first obscure dor-
sally; no setae on any tentacular segment;
formula: 1 + 0-J-+ 0-^ 14
1 N
Body and dorsal cirri greenish yellow. Dorsal tentacular cirri
of second and third segments questionably flattened in
section Ph. (Nereiphylla) fragilis
Body and dorsal cirri red. All tentacular cirri rounded in
section Ph. (Gentyllis) castanea
Ventral cirri long and tapered to points.- Dorsum greenish
brown in adults and barred with brown in juveniles Pit. (Anaitides) mucosa
Ventral cirri oval with abruptly pointed tips. Dorsum greenish
blue PJi. (Anaitides) groenlandiea
Setigerous lobe pointed. "Neck" dusky. [Ventral cirri long,
slender, and pointed] Ph. (Anaitides) longipes
Setigerous lobe blunt. "Neck" not dusky 16
Ventral cirri long and tapered. Base of proboscis with numer-
ous papillae 17
Ventral cirri oval with blunt tips anteriorly, becoming abruptly
pointed posteriorly. Base of proboscis with six regular rows
of lateral papillae Pli. (Anaitides) madeirensis
Dorsum green with dark intersegmental cross bars PI/. (Anaitides) arenae
Dorsum green with median dark stripe Pit. (Anaitides) panamensis
Lugia rarica Uschakov, 1958
Figure 3g-j
Lugia rarica Uschakov, 1958: 204, Fig. 1 A-C
Description — Body vermiform, 18 mm long
by 1.5 mm wide for 84 segments. Eggs green;
dorsal cirri faintly red in alcohol. Prostomium
(Figure 3g) as broad as long, rounded in front,
with small lateral eyes but no occipital papilla.
Proboscis diffusely papillose. Tentacular seg-
ments well developed and separate. Tentacular
formula 1 + S tt + S — , there being ten-
N N
tacular cirri on segments 1 and 2 but none on
3. V2 and V3 both lamellar but V2 larger than V;i.
Dorsal cirri of normal body segments (Figure
3h) all oval but slightly longer posteriorly. Setig-
erous lobes blunt; ventral cirri similar to dor-
sal ones but smaller. Setae (Figure 3i, j) with
long, almost smooth blades and denticulate
shaft-heads bearing a curved tooth on one side.
Remarks. — Two specimens were obtained,
the larger being an ovigerous female. As com-
pared with Ushakov's original figures, the pro-
stomium is shorter and the second tentacular
cirrus (D2) is shorter. Again Ushakov's Figure
1 C shows a seta with a more coarsely ser-
rated blade and no tooth on the shaft-head.
However, this feature is only visible under oil-
immersion and the lack of a dorsal cirrus on
the third segment is characteristic and im-
mediately separates L. rarica from the type
species L. pterophora (Ehlers).
IX
Records. — Two specimens from muddy sand
in 200 m off Beaufort (*). This is a new record
for the United States.
Distribution. — Kamchatka in 5,070 m.
Eteone lactea Claparede, 1868
Eteone lactea. - Fauvel, 1923: 175, Fig. 63 a-d.
-Pettibone, 1963a: 70, Fig. 16 a-c.
Records. — Off North Carolina; intertidal to
28 m (11, 13, 17, 18).
Distribution. — Atlantic from the Shetland Is-
lands to the North Sea and the Gulf of St.
Lawrence to Florida; Mediterranean; intertidal
to 200 m.
Eteone heteropoda Hartman, 1951
Eteone heterpoda Hartman, 1951: 31, pi. 9:
Fig. 1-8. - Pettibone, 1963a: 72, Fig. 16 d.
Records. — Off Beaufort in 10-200 m (15, *).
Distribution. — Maine to North Carolina and
the Gulf of Mexico; intertidal to 18 m.
Protomystides bidentata Langerhans, 1879
Figure 3a-f
Mystides (Protomystides) bidentata. - Southern,
1914: 71, pi. 8: Fig. 17 a, b.
Protomystides bidentata. - Bergstrom, 1914: 184.
-Hartman, 1965a: 62.
Description. — Body threadlike, greenish yel-
low, about 12 mm long by 0.1 mm. Prostomium
(Figure 3a) elongate, slightly broader at trun-
cate posterior end. Four tapered and subequal
antennae, a pair of indistinct eyes, no occipital
papilla. Proboscis diffusely papillose. Three
well-developed and separate tentacular seg-
ments bearing three tentacular cirri and setae
from second. Formula 1 + S — + S — . Ventral
N N
cirrus of second segment (V2) lamellar but
tapered distally and longer than V3. Normal
parapodia (Figure 3b) with a long, blunt, setig-
erous t9J, an elongated oral dorsal cirrus
and a similar but smaller ventral cirrus. About
eight spinigerous setae per foot; shaft-heads
(Figure 3c, d) symmetrical and serrated; blades
(Figure 3e, f) grading in length.
Remarks. — As Southern (1914) has remarked,
the fact that tentacular cirrus V2 is longer and
more tapered than V3 makes it difficult to sepa-
rate Protomystides from Phyllodoce.
Records. — One specimen off Beaufort in 200
m (*).
Distribution. — Warm North and tropical At-
lantic; Mediterranean; in 10-4,950 m.
Hesionura elongata (Southern, 1914)
Figure 3k-m
Mystides (Mesomystides) elongata Southern,
1914: 74, pi. 5: Fig. 12.
Mystides (Pseudomystides) elongata. - Fauvel,
1923: 182, Fig. 66 d-g.
Mystides elongata. - Renaud, 1956: 10.
Eteonides elongata. - Hartmann-Schroder, 1963:
216, Fig. 21-23.
Hesionura elongata. - Hartman, 1965b: 18 (cata-
logue).
Description. — Body threadlike, very slender,
brownish green, about 15 mm long. Prostomium
(Figure 3k) about twice as long as broad, with
two pairs of frontal antennae and indistinct
brown eyespots. No median antenna. Proboscis
with numerous dark brown papillae. Three pairs
of tentacular cirri on first and second segments.
Both tentacular segments distinct, separate and
without setae. Third segment without a dor-
sal cirrus but with setae. Tentacular formula:
1 + 0~r+ S — . First tentacular cirrus cylindri-
cal and tapered, about 1.5 times segmental
breadth, second dorsal cirrus (D2) similar but
slightly longer, second ventral cirrus (V2) short,
only slightly longer than ventral cirri of subse-
quent segments. Normal parapodia (Figure 31)
with short, fusiform dorsal cirri, bluntly conical
setigerous lobes longer than dorsal cirri but
shorter than the very long fusiform ventral cirri.
Setae (Figure 3m) four or five in number with
bifid or even trifid shaft -heads and short, knife-
shaped blades with deeply serrated edges. Pygid-
ium with two very long slender anal cirri.
Remarks. — The blades of the setae fall off
easily and this has given rise to Southern's
statement, repeated by Fauvel, that simple setae
lit
are present. Hartmann-Schroder (1963) reviewed
the synonymy of the genus Mystides and its
subgenera and showed that these names could
not be used for Southern's species. She re-
ferred it to a Eteonides Hartmann-Schroder but
this in turn is a synonym of Hesiouura Hartmann-
Schroder.
Records. — Eleven specimens from 10 to 200
m off Beaufort (*).
Distribution. — Ireland; Bimini Islands; in 10-
20 m.
Eulalia (Pterocirrus) macroceros Grube, 1860
Eulalia (Pterocirrus) macroceros. -Fauvel, 1923:
167, Fig. 60 d-g (partim). - Day, 1960: 301,
Fig. 5 g-i; 1967: 152, Fig. 5.4. a-c.
Description. - Body up to 20 mm long, rather
broad, greenish. Prostomium bilobed poste-
riorly with a brownish swelling between the
lobes. Eyes large; median antenna long and
well forward. Proboscis with a narrow ring of
elongate papillae basally but mainly smooth
distally. First tentacular segment mainly fused
to prostomium; second and third segments
separate but without setae. Four pairs of ten-
tacular cirri; tentacular formula: 1 + 0 — h 0—
1 N
but second ventral cirrus (V2) flattened on one
margin. Normal body segments with cordate
dorsal cirri; superior part of setigerous lobes
slightly produced and ventral cirri slightly
pointed. Setae with markedly striate shaft-heads
and long blades.
Records. — Common on coral in 6.5-18 m off
Beaufort (20, *). This is a new record for the
Atlantic coast of the United States.
Distribution. — Mediterranean; Morocco to
Senegal; South Africa; Washington; 5 to 30 m.
Eulalia ( Eumida) sanguinea (Oersted, 1843)
Eulalia (Eumida) sanguinea. -Fauvel, 1923: 166,
Fig. 59 f-k. - Day, 1967: 155, Fig. 5.5. a-c.
Emu /da sanguiyiea. - Pettibone, 1963a: 88, Fig.
21 a, b. - Hartman, 1968: 275, Fig. 1-3.
Records. — Several records between Cape
Hatteras and Beaufort from the shore to 40 m
(3, 11, 13, 15, 17, 18,20,21, *).
Distribution. — Cosmopolitan from cold tem-
perate to subtropical seas; intertidal to 600 m.
Eulalia bilineata (Johnston, 1840)
Eulalia bilineata. - Fauvel, 1923: 162, Fig. 58
a-e. - Pettibone, 1963a: 86, Fig. 20. - Hart-
man, 1968: 261, Fig. 1,2.
Eulalia (Hypoeulalia) bilineata. - Day, 1967: 164,
Fig. 5.4. k-m.
Records. — North Carolina, intertidal (17, *).
Distribution. — Arctic; North Atlantic from
Norway to the English Channel and Nova Scotia
to North Carolina; South Africa; Pacific from
North Japan Sea to the Yellow Sea and Van-
couver Island to southern California; intertidal
to 2,000 m.
Eulalia viridis (Linnaeus, 1767)
Eulalia viridis. - Fauvel, 1923: 160, Fig. 57 a, b.
-Pettibone, 1963a: 85, Fig. 19. - Hartman,
1968: 267, Fig. 1-3.
Records. — Cape Hatteras to Beaufort, inter-
tidal (18).
Distribution. — Arctic; Atlantic from Norway
to Cape Verde Island and Iceland to North
Carolina; Pacific from northern Japan to China
and Alaska to Panama; ? Indian Ocean; inter-
tidal to 200 m.
Paranaitis kosteriensis (Malmgren, 1867)
Anaitis kosteriensis. - Bergstrom, 1914: 156,
pi. 1: Fig. 1, text Fig. 52a-c.
Phyllodoce (Anaitis) kosteriensis. -Fauvel, 1923:
157, Fig. 56 a-c.
Paranaitis kosteriensis. - Pettibone, 1963a: 77,
Fig. 17 d.
Records. — One small specimen from 160 m
off Beaufort (*).
Distribution. — North Atlantic from Sweden to
Ireland and Labrador to New England; 10-
2,000 m.
20
Figure B.—Protomystides bidentata a, anterior end; b, foot; c, superior view of shaft-head with blade removed'
d, lateral view of shaft-head; e, inferior seta; f, superior seta. Lugia rarica g, anterior end; h, foot; i, shaft-head of
seta; j, seta. Hesionura elongate, k, anterior end; 1, foot; m, seta. Phyllodoce panamensis n, foot; o, shaft-head
of seta; p, anterior end.
2\
Paranaitis speciosa (Webster, 1880)
Paranaitis speciosa. - Pettibone, 1963a: 75,
Fig. 17 a.
Records. — One small specimen from 20 m
off Beaufort (*).
Distribution. — Maine to Chesapeake Bay; in-
tertidal to 183 m.
Phyllodoee (Nereiphylla) fragilis Webster, 1879
Phyllodice [sic] fragilis Webster, 1879: 14, pi. 3:
Fig. 32-37.
Phyllodoee fragilis. - Hartman, 1942b: 111.
Nereiphylla fragilis. - Hartman, 1945: 14, pi. 2:
Fig. 1-4.
[A^oji] Nereiphylla fragilis. -Hartman, 1951: 34.
Remarks. — According to Bergstrom (1914:
102) , Nei'eiphylla BlamviUe and Ge » etyll isMalm-
gren are very similar, the main distinction being
that in Nereiphylla the dorsal tentacular cirri
of segments 2 and 3 are not rounded in section
as they are in Genetyllis but definitely flattened
so as to form slender lancetlike blades. When
describing Ph. (Nereiphylla) fragilis, neither
Webster nor Hartman mention that any of the
tentacular cirri are flattened and give the im-
pression that they are cirriform. While I do
not feel that this character, which may be due
to the method of preservation, is worthy of
generic status, I feel that further changes in
the name should be avoided until fresh or
preferably living specimens are examined.
Records. — Many records from the shore to
40 m between Cape Hatteras and South Carolina
(3,5, 11, 14, 15, 18, 19,20).
Distribution. — Virginiato South Carolina; in-
tertidal to 40 m.
Remarks. — As noted above, Genetyllis has
cylindrical tapered tentacular cirri. In compari-
son with the closely related Ph. (N.) fragilis,
Ph. (G.) cast an ea has a shorter, broader body
which is orange, not yellowish green and the
dorsal cirri are red. The tentacular cirri are
shorter and stouter and the dorsal cirri, though
cordate, are more pointed. Possibly the record
by Hartman (1951) of Nereiphylla fragilis from
Florida and the Gulf of Mexico with clavate to
cirriform tentacular cirri and dorsal cirri deep
purple when preserved should be referred to
Ph. (G.) castanea.
Records. — Two specimens from coral in 20 m
off Beaufort (*).
Distribution. — In warm and tropical waters
of all oceans; intertidal to 30 m.
Phyllodoee (Anaitides) mueosa Oersted, 1843
Phyllodoee (Anaitides) mucosa. - Fauvel, 1923:
152, Fig. 54 a-e. - Pettibone, 1963a: 81,
Fig. 18 f-g.
Anaitides mucosa. -Hartman, 1968: 235, Fig. 1,2.
Note. — The eight specimens from Beaufort
are all juveniles less than 30 mm long. They
agree with the descriptions of Fauvel and Petti-
bone in all respects except that the papillae
at the base of the proboscis are more numerous
with approximately 10 irregular rows on each
side instead of 6.
Records. — Off Beaufort and North Carolina
in 80-160 m (*).
Distribution. — Arctic and the North Atlantic
from Denmark to West Africa and Hudson Bay
to Mexico; California; intertidal to 400 m.
Phyllodoee (Anaitides) groenlandiea
Oersted, 1843
Phyllodoee (Genetyllis) castanea
(Marenzeller, 1879)
Genetyllis castenea. - Bergstrom, 1914: 158,
Fig. 53. - Hartman, 1968: 281, Fig. 1, 2.
[?] Nereiphylla fragilis. - Hartman, 1951: 34.
Phyllodoee (Genetyllis) castanea. - Day, 1967:
149 (non Fig. 5.3. d).
Phyllodoee (Anaitides) groenlandiea. - Fauvel,
1923: 153, Fig. 54 f-i. - Pettibone, 1963a:
80, Fig. 18 e.
Records. — Off North Carolina; intertidal to
1,585 m (17).
Distribution. — Arctic; North Atlantic from
Norway to the English Channel and Hudson
i-i
Bay to North Carolina; North Pacific from the
Bering Sea to northern Japan and southern
California; intertidal to 1,700 m.
Phyllodoce (Anaitides) longipes Kinberg, 1866
Phyllodoce (Anaitides) longipes. - Day, 1963a:
394, Fig. 3 d-f; 1967: 144, Fig." 5.2. a-c.
Anaitides longipes. - Hartman, 1968: 229, Fig.
1-3.
Description. — Body about 25 mm long; color
greenish ; a distinctive dusky "neck" immediately
behind the tentacular cirri and three rows of
fainter spots on subsequent segments. Pro-
stomium elongate-cordate with four frontal an-
tennae and a minute occipital papilla. One pair
of dark eyes. First tentacular segment not
visible dorsally, second and third distinct and
separate though without setae. Four pairs of
long cylindrical tentacular cirri; tentacular for-
mula: 1 + 0 — + 0 — . Dorsal cirri broadly
oval, almost circular anteriorly. Setigerous lobes
long, superior part of presetal lip pointed and
projecting well below inferior part. Ventral cirri
long, slender and pointed. Setae with shaft-
heads minutely striated.
Records. — Eight specimens on sand in 20-40
m off Beaufort (21, *). This is a new record for
the Atlantic coast of the United States.
Distribution. — North Carolina; South Africa;
California; Chile; intertidal to 40 m.
Phyllodoce (Anaitides) madeirensis
Langerhans, 1880
Phyllodoce (Anaitides) madeirensis. - Fauvel,
1923: 150, Fig. 23 d-h. - Day, 1967: 145,
Fig. 5.2. d-g.
Anaitides madeirensis. - Nonato and Luna,
1970b: 66, Fig. 5-8.
Description. — Body up to 100 mm long; bright
green when alive but fading in alcohol. Pro-
stomium cordate with a deep posterior notch
and an occipital papilla. Four frontal antennae
and a pair of dark eyes. Base of proboscis
with six lateral rows of papillae with about 11
papillae per row. Sometimes a median row of four
or five. Distal part of proboscis with six lumpy
ridges. First tentacular segment not visible
dorsally, second and third distinct and separate.
Four long, cylindrical tentacular cirri; tentacu
lar formula:
1 + 0 — - + 0 — • Dorsal cirri
1 N
hastate anteriorly, becoming rhomboidal on
middle segments. Setigerous lobes apical ly
blunt. Ventral cirri oval with blunt tips anteriorly
but pointed tips posteriorly. Setae with strongly
serrated shaft-heads.
Remarks. — The description of Phyllodoce
oculata by Ehlers (1887: 135, pi. 40: Fig. 4-6)
from Florida suggests that this species is a
synonym of Ph. madeirensis. The only doubtful
point is whether Ph. oculata lacks setae on the
third tentacular segment. Augener (1925), who
re-examined the type of Lopadorhynchus ery-
throphyllus Schmarda, from Jamaica, states
that it is identical with Ph. oculata although I
feel that the color as described and figured
by Schmarda is quite different from that of
Ph. oculata or Ph. madeirensis. If the three
species are really identical, Schmarda's name
would have priority. A reexamination of the
types is required.
Records. — Two specimens from coral in 10 m
off Beaufort (*).
Distribution. — Cosmopolitan in warm and
tropical seas; intertidal to 200 m.
Phyllodoce (Anaitides) arenae Webster, 1880
Phyllodoce arenae Webster, 1880: 105; 1886:
133, pi. 5: Fig. 10-12.
Phyllodoce (Anaitides) arenae. - Pettibone,
1963a: 82, Fig. 18 a-c.
Remarks. — This species is common in the
cold New England waters and is close to Ph.
panamensis from warmer waters farther south.
Both have ventral cirri pointed and longer than
the setigerous lobes but differ in the pigment
pattern; Ph. (A.) arenae has spindle-shaped
intersegmental crossbars while Ph. panamen-
sis has a dorsal stripe.
Records. — Six specimens in 20-200 m off
Beaufort (*).
Distribution. — Maine to New Jersey; inter-
tidal to 194 m.
23
Phyllodoce (Anaitides) panamensis
Treadwell, 1917
Fig'ure 3n-p
PlnjUodoce (Anaitides) panamensis. - Monro,
1933b: 24, Fig. 11a. b.
Description. — Body slender, up to 107 mm
long; color greenish with a dark median dorsal
stripe and a faint ventral one. Prostomium (Fig-
ure 3p) cordate with four frontal antennae and
a minute occipital papilla. Base of proboscis
completely covered with compressed papillae
irregularly arranged. First tentacular segment
not visible dorsally, second and third distinct
and separate. Four pairs of long cylindrical
tentacular cirri but no setae ; tentacular formula :
1 + 0~r+ Ott. D0l*sal cirri (Figure 3n) large
and oval; setigerous lobes blunt; ventral cirri
slender and pointed, far longer than setigerous
lobes. Setae (Figure 3o) with faintly striated
shaft -heads.
Remarks. — Ph. panamensis is very close to
Ph. arenae differing mainly in the color pattern;
possibly it is no more than a subspecies of
PJi. arenae.
Records. — Two specimens from 80 to 120 m
off Beaufort ( * ) .
Distribution. — Panama; 10 m.
FAMILY PILARGIDAE
Key to genera and species
1 Prostomium with three antennae. Notosetae always repre-
sented by a stout acicular seta 2
1' Prostomium with two antennae. Notosetae either absent or
represented by a stout hook 3
2 Notopodial acicular seta in the form of a hook. Antennae long.
(Sigambra). [Notopodial hook appearing on setiger 14-25] S. bassi
2' Notopodial acicular seta straight. Antennae short (Synelmis).
[Dorsal cirrus of first setiger twice as long as subsequent
ones] S. albini
3 Notosetae absent. (Pilargis) No N.C. record
3' Notosetae represented by a stout hook. [Dorsal and ventral
cirri small or absent {Cabira)] No N.C. record
Sigambra bassi (Hartman, 1945)
Ancistrosyllis bassi Hartman, 1945: 15; 1947b:
501, pi. 61: Fig. 1-7; 1951: 36, pi. 11: Fig. 1-
6.
Sigambra 6a.s.s/.-Pettibone, 1966: 186, Fig. 16.-
Hartman, 1968: 389, Fig. 1-5.
Records. — Beaufort Sound, intertidal to a few
meters (11, 12, 13, 18).
Distribution. — North Carolina to Florida;
central to southern California; intertidal to
33 m.
Synelmis albini (Langerhans, 1881)
Ancistrosyllis rigida. - Fauvel, 1919: 337, Fig.
1 a-e. - Hartman, 1947b: 498, pi. 62: Fig.
1-7. -Day, 1967: 215.
Synelmis albini. - Pettibone, 1966: 191, Fig.
19-21. - Hartman, 1968: 393, Fig. 1-5. -
Nonato and Luna, 1970b: 68, Fig. 10-14.
Records. — One juvenile off Beaufort in 80 m
(*).
Distribution. — Circumtropical and extends
from Brazil to North Carolina; southern Cali-
fornia; intertidal to 2,200 m.
24
FAMILY HESIONIDAE
Key to genera and species
1 Eight pairs of tentacular cirri; [3 antennae. Proboscis with
marginal papillae but no jaws. Notopodium vestigial with
few" setae (Gyptis)] G. vittata
V Six pairs of tentacular cirri 2
2 A median and two lateral antennae. [Proboscis with marginal
papillae (Ophiodromus)] O. obscurus
2' No median antenna, only two laterals 3
3 Palps biarticulate. Notopodium reduced to an aciculum in
dorsal cirrophore and usually a few setae (Nereimyra) N. punctata
3' Palps not articulated. Notopodium a small but distinct lobe
on dorsal cirrophore bearing several setae (Parahesione
luteola) No. N.C. record
Gyptis vittata Webster and Benedict, 1887
Gyptis vittata. - Pettibone, 1963a: 106, Fig. 28
c,d.
Records. — Cape Hatteras area, intertidal; off
Beaufort in 10-20 m (18, *).
Distribution. — Maine to North Carolina; inter-
tidal to 55 m.
Records. — Many records between Cape Hat-
teras and Beaufort; intertidal to 40 m (5, 11,
13, 15, 17, 18,20, *)
Distribution. — Massachusetts to the West In-
dies and the Gulf of Mexico; intertidal to 840 m.
Nereimyra punctata (Muller, 1776)
Ophiodromus obscurus (Verrill, 1873)
Podarke obscura. - Pettibone, 1963a: 104, Fig.
28 a, b.
Remarks. — Dr. Pettibone, in a personal com-
munication, maintains that Podarke Ehlers
should not be regarded as a synonym of Ophio-
dromus Sars since the type species of Podarke,
(P. agilis) has nearly uniramous parapodia,
while the type species of Ophiodromus, (O. vit-
tatus Sars = O. flexuousus Delia Chiaje) has
parapodia equally biramous. However both
genera have three antennae and six pairs of
tentacular cirri in contrast to Gyptis, Nereimyra ,
and Parahesione and, as noted earlier (Day,
1967), I agree with Dr. Hartman that Podarke
is a synonym of Ophiodromus. The number of
setae in the notopodia is, I feel, of specific but
not generic importance.
Castalia punctata. -Fauvel, 1923: 24, Fig. 89f-k.
Nereimyra punctata. - Pettibone, 1963a: 107,
Fig. 28 e.
Records. — Off Chesapeake Bay and off North
Carolina, ? depth (17).
Distribution. — Arctic, North Atlantic from
Norway to France and Hudson Bay to North
Carolina; Azores; Bering Sea; intertidal to
2,350 m.
Parahesione luteola (Webster, 1880)
Parahesione luteola. - Pettibone, 1956:
Fig. 1 a-e; 1963a: 108, Fig. 29 a-c.
281,
Records. — No record from North Carolina.
Distribution. — Massachusetts to New Jersey
and Georgia to the Gulf of Mexico; intertidal.
25
FAMILY SYLLIDAE
Key to genera and species
1 Ventral cirri distinct. Palps either separate or fused. Pharynx
straight, seldom coiled 2
1' Ventral cirri completely fused to setigerous lobes and in-
distinguishable. Palps completely fused. Pharynx long and
coiled. (Subfamily Autolytinae) 23
2 Palps quite separate. Always two pairs of tentacular cirri.
Antennae and dorsal cirri articulated. (Subfamily Syllinae) 3
2' Palps fused basally. One or two pairs of tentacular cirri.
Antennae and dorsal cirri seldom articulated (Subfamily
Eusyllinae) ' 11
2" Palps fused for more than half their length. One or two pairs
of tentacular cirri. Antennae and dorsal cirri not articulated
and often small (Subfamily Exogoninae) 16
3 Margin of pharynx with a circle of chitinous teeth (trepan)
and a small dorsal tooth as well. Body flattened (Trypcuio-
syllis). [Trepan with 10 teeth. Blades of setae bidentate.
Never more than one sexual stolon] T. zebra
3' Margin of pharynx with a single anterior dorsal tooth (Syllis) 4
4 Setae of anterior feet compound but setae of middle feet may
lose their blades and appear simple 5
4' All setae simple (subgenus Haplosyllis). [Setae shaped like
boathooks with a boss preceding the curved and bifid tip.
Dorsal cirri with 20-30 joints] S. (H.) spongicola
5 Setae of middle segments reduced to two large simple setae
with bluntly Y-shaped ends. [Dorsal cirri fusiform with about
10 joints] S. gracilis
5' Setae of middle segments fairly numerous and normally com-
pound 6
6 Two or three superior setae of middle segments with very
long tapered blades; inferior setae with much shorter fal-
cigerous blades (subgenus Langerhansia) 7
6' All setae of middle segments with falcigerous blades graded
in length (subgenus Typosyllis) 8
7 Dorsal cirri of posterior feet smooth or indistinctly articulated.
Blades of posterior setae with secondary tooth stouter than
apical one S. (L.)ferrugina
T Dorsal cirri of posterior feet distinctly articulated. Blades of
posterior setae with secondary tooth finer than apical one S. (L.) eornuta
26
8 Dorsal cirri of middle feet short and fusiform with about 15
joints. Setae obviously bidentate S. (T.) hyalina
8' Dorsal cirri of middle feet at least as long as segmental
breadth with more than 15 joints. Setae strongly bidentate 9
9 Setae of posterior feet with secondary tooth stronger than
apical one. Dorsal cirri with 25-40 joints S. (T.) regulata carolinae
9' Setae of posterior feet with secondary tooth not stronger than
apical one 10
10 Dorsal cirri of middle segments alternately with 15 or 20 joints.
Proventriculus extending over 7 to 9 segments S. (T.) alternata
10' Dorsal cirri of middle segments alternately with about 30 and
37 joints. Proventriculus extending over 5 to 6 segments S. (T.) prolifera
11 Pharynx coiled. Body short with about 13 setigers. Large
nuchal epaulettes (Anibhjosyllis). [Six triscuspid marginal
teeth] A. formosa
11' Pharynx straight. Body long with many setigers. No nuchal
epaulettes 12
12 Large knobbed acicula projecting from anterior feet. Pharynx
without teeth (StreptosyUis). [Setae with flanged shaft-
heads and blades] S. arenae
12' Acicula normal, not exposed. Pharynx with one or more teeth 13
13 A large occipital flap. Pharynx with a ventral semicircle of
teeth halfway back (Odontosyllis). [Setal blades very long
and strongly bidentate] O. longiseta
13' Occipital flap small or absent. Pharynx with a single anterior
dorsal tooth and a smooth or denticulate margin 14
14 Margin of pharynx denticulate (Eusyllis). [Dorsal cirri ex-
tremely long and first pair of ventral cirri lamellar] E. lamelligera
14' Margin of pharynx smooth 15
15 One pair of tentacular cirri (Parapionosyllis) P. longicirrata
15' Two pairs of tentacular cirri {Pionosyllis). [Superior setae
with elongated blades and inferior setae with short blades
with secondary tooth larger than apical one] P. cf. uraga
16 One pair of tentacular cirri 17
16' Two pairs of tentacular cirri 21
17 Dorsal cirri ovoid, minute; body surface without sticky papillae
or adherent silt. (Exogone) 18
17' Dorsal cirri bottle-shaped or onion-shaped; body surface with
sticky papillae and adherent silt (Sphaerosyllis) 20
18 Dorsal cirri on all setigers including second 19
18' No dorsal cirrus on setiger 2. [Median antenna hardly longer
than laterals] E. gemmifera
27
19 Median antenna much longer than laterals E. dispar
19' Median and lateral antennae all equally short E. verugera
20 Dorsal cirri on all setigers including second. No internal
capsules above parapodia {S. fortuita) No N.C. record
20' No dorsal cirrus on setiger 2. No internal capsules above
parapodia S. pirifera
21 Dorsum with six rows of large globular papillae across each
segment; antennae also globular (Eurysyllis) E. tuberculata
21' Dorsum without rows of globular papillae; [embryos carried
on backs of females (Brania)] 22
22 Dorsal cirri abruptly truncate and containing fibrillar struc-
tures. Setae with unidentate blades B. pusilla
22' Dorsal cirri normally tapered and lacking fibrillar structures.
Setae with bidentate blades B. clavata
23 Superior simple setae with shafts as stout as those of com-
pound setae. Segments without ciliated bands. Reproduc-
tion by anterior scissiparity (Proceraea) 24
23' Superior simple setae with shafts more slender than those of
compound setae. Segments with ciliated bands. Reproduc-
tion by posterior scissiparity, gemmiparity or epigamy ( An to-
lytus) 25
24 Body colorless or with a pair of faint dorsolateral bands when
fresh. Nuchal epaulettes rudimentary. [Trepan with 18
teeth] P. corn u ta
24' Body with reddish bars on alternate segments when fresh.
Nuchal epaulettes reach setiger 1 P.fasciata
25 Nuchal epaulettes reach setiger 3-4. [Trepan with 30 teeth
including large laterals and small teeth in the dorsal and
ventral arcs. Body colorless or segments faintly banded
when fresh] A. dentalius
25' Nuchal epaulettes not reaching setiger 3 26
26 A chain of 2-8 sexual buds when mature. No color pattern.
Trepan with 24-36 subequal teeth A. prolifer
26' Sexual buds formed singly. Anterior segments with four red
spots when fresh. Trepan with 30-40 subequal teeth A. rubropunctatus
Trypanosyllis zebra (Grube, 1860) stomium broader than long, with large palps
well separated basally and two pairs of eyes.
Trypanosyllis zebra. - Fauvel, 1923: 269, Fig. Two pairs of tentacular cirri. Antennae and
101 a-e. - Day, 1967: 256, Fig. 12.6. a, b. dorsal cirri stout, with numerous (over 30) well-
marked joints. Margin of pharynx (trepan) with
Description. — Body markedly flattened, up to 10 equal teeth and a small dorsal tooth as well.
60 mm long, with two narrow purple lines across Setae all compound with bidentate blades, the
anterior segments and purple dorsal cirri. Pro- two teeth being subequal and close together
28
at tip of blade. No cluster of sexual buds below
pygidium.
Records. — Common on coral in 6.5-18 m off
Beaufort (20, *). This is a new record for the
United States.
Distribution. — English Channel; Mediterran-
ean; Indian Ocean; intertidal to 30 m.
Syllis ( Haplosyllis) spongicola (Grube, 1855)
Syllis (Haplosyllis) spongicola. - Fauvel, 1923:
257, Fig. 38 a-h. - Day, 1967: 240, Fig. 12.1.
e-i.
Haplosyllis spongicola. - Imajima, 1966d: 220,
Fig. 38 a-h. - Hartman, 1968: 433, Fig. 1-4.
Records. — Common from Cape Hatteras to
Beaufort; intertidal to 30 m (5, 11, 13, 15, 18,
20, *).
Distribution.— Cosmopolitan in warm and
tropical seas; intertidal to 100 m.
Syllis gracilis Grube, 1840
Syllis gracilis. - Fauvel, 1923: 259, Fig. 96 f, i. -
Pettibone, 1963a: 116, Fig. 32. - Imajima,
1966d: 248, Fig. 49 a-h.
Syllis (Syllis) gracilis. - Day, 1967: 241, Fig.
12.1. m-p.
Records. — Common from Cape Hatteras to
Beaufort; intertidal to 20 m (3, 11, 13, 17, 18,
20, *).
Distribution. — Cosmopolitan in temperate and
tropical seas; intertidal to 200 m.
basally; two pairs of eyes. Antennae and an-
terior dorsal cirri with 17-25 distinct joints;
posterior dorsal cirri (Figure 4n) with very in-
distinct joints. Setae all compound including
two or three superior ones with very long taper-
ing blades (Figure 4q) and several inferior
ones with bidentate blades of normal length
(Figure 4o). Posterior setae (Figure 4p) with
secondary tooth larger than apical one.
Records. — One specimen from 120 m off
Beaufort (*).
Distribution. — Eastern Atlantic from Ireland
and the Canary Islands to Angola and South
Africa; intertidal to 30 m.
Syllis (Langerhansia) cornuta Rathke, 1843
Syllis (Ehlersia) cornuta. - Fauvel, 1923: 267,
Fig. 100.
Syllis cornuta.- Pettibone, 1963a: 118, Fig. 31 i,j.
Langerhansia cornuta. - Imajima, 1966e: 256,
Fig. 51 a-o.
Syllis (Langerhansia) cornuta. - Day, 1967: 244,
Fig. 12.2. s-u.
Records. — Common from the Cape Hatteras
area to Beaufort; intertidal to 7 m (11, 17, 18,20).
Distribution. — Cosmopolitan from the Arctic
to the Antarctic; intertidal to over 2,000 m.
Syllis (Typosyllis) hyalina Grube, 1863
Syllis (Typosyllis) hyalina. -Fauvel, 1923: 262,
Fig. 98 a, b. - Day, 1967: 246, Fig. 12.2.
v-x.
Typosyllis hyalina. -Hartman, 1968: 487, Fig. 1-
3.
Syllis (Langerhansia) ferrugina
Langerhans, 1881
Figure 4n-q
Syllis (Ehlersia) ferrugina. - Fauvel, 1923: 269,
Fig. 100 k-u.
Syllis (Langerhansia) ferrugina. - Day, 1967:
244, Fig. 12.2. o-r.
Description. — Body about 10 mm long, with-
out color markings. Palps large and separate
Description. — Length 10-35 mm. Prostomium
with palps separate basally; three antennae,
two pairs of eyes, and anterior ocular specks.
Two pairs of tentacular cirri. Pharynx long with
anterior dorsal tooth. Antennae and dorsal cirri
distinctly jointed; dorsal cirri short and fusi-
form, those of middle segments having about
8-15 joints. Setae all compound with obviously
bidentate blades.
Records. — Two specimens from 20 and 160 m
off Beaufort (*).
Distribution. — Cosmopolitan in temperate and
tropical seas; intertidal to shallow depths.
29
Syllis (Typosyllis) regulata carolinae
New Subspecies
Figure 4a-f
Holotype. — USNM 43146; five paratypes,
USNM 43147.
Description. — Body 20 mm long, very slender
with rounded segments and long many-jointed
dorsal cirri. No color markings. Prostomium
(Figure 4a) broader than long with four small
eyes and elongate palps separated basal ly.
Median antenna much longer than laterals
and inserted between eyes. Pharynx extending
through 11 segments with a large anterior dorsal
tooth. Proventriculus with 32 rows of points
and extending through 5 or 6 segments. Dorsal
cirri alternating in length; longer ones of middle
segments (Figure 4f) exceeding twice segmental
breadth and having about 40 joints, shorter
ones equalling segmental breadth and having
about 25 joints. Setigerous lobes long and
conical, ventral cirri slender and pointed. Com-
pound setae similar throughout, with long spi-
nules on cutting edge of blade and strongly
bidentate ends, the secondary tooth becoming
stronger than apical one in posterior feet (Figure
4c). Two simple setae in posterior feet; superior
one (Figure 4d) truncate or emarginate and
inferior one (Figure 4e) with a definitely bi-
dentate tip.
Remarks. — Imajima (1966e) described Typo-
syllis regulata from Seto, Japan, and compared
it with Typosyllis truncata Haswell from Aus-
tralia and Typosyllis harti Berkeley and Berkeley
from Vancouver. All agree in having slender
bodies, long many-jointed dorsal cirri and
strongly bidentate compound setae, but differ
in the length of the proventriculus and details
of the acicula and posterior simple setae. Like
T. truncata, the Carolinean specimens have
the proventriculus extending through five or
six segments, but they differ in having biden-
tate inferior simple setae and not pointed ones.
Like T. regulata the Carolinean specimens differ
from T. harti in having slightly knobbed acicula
(Figure 4b) and setae with a strong secondary
tooth and long serrations on the cutting edge
of the blade. The Carolinean specimens differ
from T. regit la to in having a shorter proventric-
ulus, which extends through 5-6 segments and
not 11 and in having superior simple setae
whose ends are truncate to bilobed instead of
pointed. These differences are small and do not
warrant more than subspecific rank.
Records.— Off Beaufort in 20 m on shelly
sand (*).
Syllis (Typosyllis) alternata Moore, 1908
Typosyllis alternata. - Imajima, 1966e: 273, Fig.
58a-l.
Records. — Common on corals in 18 m off
Beaufort (20).
Distribution. — Alaska to California; Japan
and northwest Japan Sea; intertidal to 350 m.
Syllis (Typosyllis) prolif era Krohn, 1852
Syllis (Typosyllis) prolif era. - Fauvel, 1923: 261,
Fig. 97 a-g. - Day, 1967: 248, Fig. 12.3. g-i.
Typosyllis prolif era. - Imajima, 1966e: 292, Fig.
65 a-n.
Records. — On corals in 18 m off Beaufort (20).
Distribution. — English Channel; Mediterran-
ean; Indo-west-Pacific from South Africa to
Japan; Brazil; intertidal to 30 m.
Amblyosyllis formosa Claparede, 1863
Pterosyllis formosa. - Fauvel, 1923: 280, Fig.
105 h-m.
Amblyosyllis formosa. - Day, 1967: 259, Fig.
12.6. m-p.
Records. — On corals in 18 m off Beaufort (20).
Distribution. — North Atlantic from Plymouth
to Senegal; Mediterranean; intertidal to 30 m.
Streptosyllis arenae
Webster and Benedict, 1884
Streptosyllis arenae Webster and Benedict,
1884: 711, pi. 2: Fig. 17-21, pi. 3: Fig. 22,
23. - Pettibone, 1963a: 127, Fig. 31 1, m.
Records. — Two specimens from 5 to 10 m
off Beaufort (*).
Distribution. — Massachusetts; intertidal.
30
Figure 4. — Sl/llis regulata carolinae n. subsp. a, head; b, aciculum; c, posterior compound seta; d, superior simple
seta; e, inferior simple seta; f, foot. Odontosyllis loiigiseta n. sp. g\ head; h, pharynx slit dorsally and flattened
to show teeth; i, foot; j, seta. Pionosyllis cf. uraga k, foot with longer dorsal cirrus; 1, superior seta; m, inferior
seta. Syllis (Langerhansia) ferrugina n, posterior foot; o, inferior seta of an anterior foot; p, inferior seta of a
posterior foot; q, superior seta.
3 1
Odontosyllis longiseta, New Species
Figure 4g"-j.
Odontosyllis n. sp. Day, Field, and Montgomery,
1971: 121.
Holotype. — USNM 43120; 100+ paratypes,
USNM 43121.
Description. — Numerous specimens obtained
but all broken. Complete specimen probably
15-20 mm long by 1.5 mm wide for 60-70 seg-
ments. Body stout, strongly arched dorsally
but flattened ventral ly. No color pattern and
whole worm uniformly flesh pink in alcohol.
Prostomium (Figure 4g) much broader than
long and separated from tentacular segment
by a deep groove. Palps broad and flattened,
fused basally and bent ventrally. Two pairs of
large red eyes, three rather short antennae, and
a pair of curved nuchal ridges almost encircling
posterior pair of eyes. Tentacular segment very
short and fused dorsally to setiger 1, the large
circular occipital flap appearing to arise from
the latter. Two pairs of tentacular cirri slightly
longer than antennae. Mouth with well defined
lateral lips; buccal cavity with an internal dor-
sal projection at entrance to pharynx. Pharynx
short, broad and strongly chitinized with a ven-
tral arc of six recurved teeth and a cutting
plate on either side (Figure 4h). Pharynx ex-
tending from setiger 8 to 10 and followed by a
long muscular proventriculus with 60 rows of
points extending from setiger 10 to 18.
Antennae, tentacular cirri, and dorsal cirri
all smooth, tapered, and relatively short. An-
terior dorsal cirri barely three-quarters seg-
mental breadth, those of middle segments alter-
nately longer and equal to half segmental
breadth or shorter and equal to one third seg-
mental breadth. Parapodia (Figure 4i) with
setigerous lobes stout and obviously bilabiate
with the setae issuing from a slit; ventral cirri
stout with pointed tips. Setae (Figure 4j) all
compound, with shaft-heads well serrated and
blades unusually long and strongly bidentate;
secondary tooth as large as apical one. No simple
setae even in posterior feet.
Remarks. — As noted earlier, the buccal cavity
has an internal dorsal ridge which extends back
and becomes a digitiform lobe at the entrance
of the pharynx. A similar structure was noted
by Day (1967) in Odontosyllis poly cera (Schmar-
da) and Pharyngeovalvata natalensis Day. Both
also possess an occipital flap so that the two
genera are related although Pharyngeovalvata
lacks chitinous teeth.
In Odontosyllis longiseta the occipital flap
appears to arise from the first setiger but this is
due to the fact that the dorsal part of the ten-
tacular segment is fused to the first setiger.
In other species, where these two segments are
separate, the occipital flap definitely arises from
the tentacular segment.
O. longiseta is easily distinguished by the
long, strongly bidentate blades of the setae,
short dorsal cirri and the large occipital flap.
In O. fulgurans Claparede, recorded by Petti-
bone (1963a) from New England, the setal blades
are short, the dorsal cirri are about equal to
the segmental breadth and the occipital flap is
small. In O. ctenostoma Claparede, O. polycera
(Schmarda), and O. dugesiana Claparede the
setal blades are all short; in O. gibba Claparede
the blades are long but unidentate.
Records. — Common off Beaufort in 20-200 m
(21 *).
Eusyllis lamelligera
Marion and Bobretzky, 1875
Eusyllis lamelligera. - Fauvel," 1923: 294, Fig.
113 a-e. - Pettibone, 1963a: 120, Fig. 33,
34 a-d.
Records. — North Carolina in 7 m (3, 17).
Distribution. — North Atlantic from Massa-
chusetts to North Carolina and the English
Channel to Spain; Mediterranean; 7-37 m.
Parapionosyllis longicirrata
(Webster and Benedict, 1884)
Pionosyllis minuta. - Fauvel, 1923: 292, Fig.
Ill f.
Parapionosyllis longicirrata. - Pettibone, 1963a:
132, Fig. 35 e, f.
Records. — Thirteen specimens in 10-20 m off
Beaufort (21, *).
Distribution. — Massachusetts ; Mediterran-
ean; intertidal to 20 m.
32
Pionosyllis cf. uraga Imajima, 1966
Figure 4k-m
Description. — Body incomplete, with only 35
segments measuring 3 mm. No color markings.
Prostomium with two pairs of eyes and three
slender antennae; the median arising far back
and twice as long as laterals. Palps broad,
flattened, bent downwards, quite separate ba-
sally. Pharynx long with a smooth margin and
a small dorsal tooth a quarter the way back.
Proventriculus as long as pharynx with 30 rows
of points. Parapodia with stout, blunt setigerous
lobes bearing broad ventral cirri mainly fused
to ventral margin. Dorsal cirri alternately very
long and slender (Figure 4k), up to 1.5 times
segmental breadth or short and only half seg-
mental breadth. Setae characteristic and of
two types, 3 to 5 superior ones (Figure 41) with
very long blades tapering to fine tips and 7 to
10 inferior ones (Figure 4m) with broad, bi-
dentate chopper-shaped blades without mar-
ginal spinules. Apical tooth small and incon-
spicuous, secondary tooth larger and hooked.
Acicula with faintly knobbed tips.
Remarks. — The long tapered blades of the
superior setae are reminiscent of the subgenus
Langerhansia but the latter has articulated
dorsal cirri and here they are all quite smooth.
This single broken specimen from North Caro-
lina closely resembles Pionosyllis uraga as de-
scribed by Imajima, (1966c: 114, Fig. 37 a-g)
from Japan. However, the blades of the setae
lack spinules, the shaft-heads are not serrated
and the heads of the acicula lack a band of
microscopic spinules. More material is required
before the identification can be confirmed.
Records. — One specimen from 120 m off
Beaufort (*).
Exogone gemmifera (Pagenstecher, 1862)
Exogone gemmifera. - Fauvel, 1923: 305, Fig.
117 a-d. - Imajima, 1966a: 397, Fig. 2 a-h. -
Day, 1967: 274, Fig. 12.10. p-u.
Exogone naidina. - Pettibone, 1954: 258, Fig.
28 e.
Description. — Body 2-4 mm long with 24-33
setigers. Prostomium with two pairs of eyes
and three antennae, the median just longer than
the laterals. Palps short, completely fused and
rounded anteriorly. Pharynx with an anterior
dorsal tooth. Proventriculus extending through
two segments. One pair of small tentacular
cirri. Dorsal cirri small and ovoid and present
on all segments except setiger 2. Ventral cirri
distinct. Setae including a superior simple seta
with an obliquely truncate tip, one or two com-
pound setae with long, daggerlike blades and
four or five compound setae with swollen serrate
shaft-heads and minute bidentate blades with
the secondary tooth larger than the terminal
one. An inferior simple setae in posterior seg-
ments. Mature females carrying developing em-
bryos ventrally.
Remarks. — The description of Exogone nai-
dina Oersted given by Pettibone (1954) agrees
perfectly with that of E. gemmifera which she
includes in the synonymy of E. naidina. How-
ever, Fauvel and more recent workers do not
regard the two as synonymous and until the
types have been examined, I prefer to use the
better known name.
Records.— Off Beaufort in 18-40 m (20, *).
This is a new record for the Atlantic coast of
the United States.
Distribution. — Arctic Seas; North Atlantic
from France to North Carolina; Mediterranean;
South Africa; North Pacific from the Bering
Sea to Mexico and Japan to the Yellow Sea;
intertidal to 225 m.
Exogone dispar (Webster, 1879)
Exogone dispar. - Hartman, 1945: 16, pi. 2:
Fig. 7, 9, 10. - Pettibone, 1963a: 130, Fig.
35 d.
Exogone clavator Ehlers, 1913: 485, pi. 33: Fig.
1-6. - Day, 1967: 272, Fig. 12.10. a-f.
Exogone uniformis Hartman, 1961: 73, pi. 6:
Fig. 1, pi. 7: Fig. 1-4. - Imajima, 1966a:
400, Fig. 4 a-j. - McCloskey, 1970: 24.
Remarks. — A direct comparison of specimens
of E. clavator from South Africa with specimens
of E. dispar from North Carolina showed that
the two are identical. Imajima states that E.
uniformis differs from E. dispar in having fal-
cigerous setae with fewer teeth on the cutting
margin. His Figures 4 f, g, h show that the
:;::
number of teeth varies from anterior to poste-
rior feet and all other characters agree with
E. dispar. It is suspected that E. lourei Berkeley
is another synonym of E. dispar.
Records. — Common on sand and coral from
low tide to 20 m off North Carolina (3, 5, 11,
13.21.*).
Distribution. — Maine to Florida; Alaska to
the Pacific coast of Mexico; South Japan; South
Africa; intertidal to 130 m.
Exogone verugera (Claparede, 1868)
with short unidentate blades and, from setiger
2 onwards, one superior simple seta with a
smooth curved tip.
Remarks. — The specimen from Beaufort
agrees with Fauvel's description except that
the palps are shorter and the dorsal cirri are
smaller than the setigerous lobes.
Records. — One specimen from 20 m off Beau-
fort (*). This is a new record for the Atlantic
coast of the United States.
Distribution. — Mediterranean; British Co-
lumbia to southern California; 0-10 m.
Exogone verugera. - Fauvel, 1923: Fig. 117
m-r. - Pettibone, 1963a: 129, Fig. 31 a-d. -
Imajima. 1966a: 399, Fig. 3 a-h. - Day,
1967: 272, Fig. 12.10. g-1. - Hartman, 1968:
429, Fig. 1-4.
Records. — Three specimens in 10-20 m off
Beaufort (*).
Distribution. — Cosmopolitan in temperate
seas; intertidal to 150 m.
Sphaerosyllis fortuita Webster, 1879
Sphaerosyllis fortuita Webster. 1879: 221, pi.
4: Fig. 44-48.
Records. — No record from North Carolina.
Distribution. — Virginia; in 0 to 10 m.
Sphaerosyllis pirifera Claparede, 1868
Sphaerosyllis pirifera.
115 1-p.
Fauvel, 1923: 301, Fig.
Description. — Body 3 mm long with 36 seg-
ments. Surface with a few scattered adhesive
papillae. Prostomium with fused palps, four
eyes and three subequal bottleshaped antennae,
the median one inserted between posterior pair
of eyes. One pair of tentacular cirri. Dorsal cirri
similar to antennae and tentacular cirri and
present on all setigers except second. Setigerous
lobes bluntly conical. No sign of internal cap-
sules above parapodia. Five compound setae, all
Eurysyllis tubereulata Ehlers, 1864
Eurysyllis tubereulata. - Fauvel, 1923: 271,
Fig. 101 i-o.
Description. — Body 3 mm long with about
50 segments. Dorsum flattened and covered
with six rows of globular papillae, the outer-
most pair on each segment representing dorsal
cirri. Prostomium broader than long with three
globular antennae and four eyes. Palps bent
and united basally forming a hood in front of
mouth. Peristome with two globular papillae
and two pairs of globular tentacular cirri.
Pharynx with a trepan of 10 marginal teeth
plus a dorsal tooth. Proventriculus globular.
Parapodia each with a globular dorsal cirrus
and a short ventral cirrus separated from the
blunt setigerous lobe (not fused to it, as stated
by Fauvel). Setae compound and falcigerous
with unidentate blades of medium length; acic-
ula with swollen ends.
Records. — One specimen off Beaufort in 40
m (*). This is a new record for the United States.
Distribution. — Mediterranean; Madeira; En-
glish Channel; intertidal to a few meters.
Brania pusilla (Dujardin, 1839)
Grubea pusilla. - Fauvel, 1923: 299, Fig. 115 a-f.
Brania pusilla. - Day, 1967: 267, Fig. 12.9. d-f.
Records. — Abundant on corals off Beaufort,
in 6.5-18 m (20).
:;l
Distribution. — North Atlantic from the Eng-
lish Channel to Morocco; Mediterranean; South
Africa; intertidal to 30 m.
Distribution. — Massachusetts to Caribbean;
intertidal to 33 m.
Brania clavata (Claparede, 1863)
Grubea clavata. - Fauvel, 1923: 296, Fig. 114
a-e.
Brania clavata. - Hartman, 1944c: 338, pi. 24:
Fig. 5-8, pi. 25: Fig. 2. - Pettibone, 1963a:
133, Fig. 35 b. - Imajima, 1966a: 393,
Fig. 1 a-g.
Records. — Common on corals in 18 m off
Beaufort (20).
Distribution. — North Atlantic from Ireland
to France and Massachusetts to the Gulf of
Mexico; Mediterranean; North Pacific from the
Bering Sea to Japan and the Yellow Sea; inter-
tidal to 30 m.
Autolytus prolifer (Miiller, 1784)
Autolytus prolifer. - Fauvel, 1923: 311, Fig.
119. - ? Pettibone, 1963a: 145, Fig. 40. -
? Day, 1967: 284, Fig. 12.13. f-k. - Gidholm,
1967: 186, Fig. 14, 15, 7A, 8.
Note. — According to Gidholm (1967), many
of the earlier records of this species are doubtful.
Nonetheless, it should be noted that A. prolifer
has been reported from Beaufort Sound and on
corals in 6-18 m by several workers (5, 8, 11,
15, 17, 18, and 20). Apart from the type locality
(Norway) it has also been reported from several
localities in the North Atlantic, the Mediter-
ranean, and South Africa. Fresh material is
needed to confirm this distribution.
Proceraea cornuta (Agassiz, 1863)
Autolytus cornutus. - Pettibone, 1963a: 144,
Fig. 37 e. - Imajima, 1966b: 49, Fig. 13 a-i. -
McCloskey, 1970: 24.
Proceraea cornuta. - Gidholm, 1967: 205, Fig.
13 e, f, 28 a-c.
Records. — Beaufort, intertidal to a few meters
(3,7,20).
Distribution. — Arctic; Atlantic from Labra-
dor to Chesapeake Bay and Norway to English
Channel; Japan; intertidal to 140 m.
Autolytus rubropunctatus (Grube, 1840)
Proceraea rubropu aetata. - Andrews, 1891a:
283.
Autolytus rubropunctatus. - Fauvel, 1923: 314,
Fig. 120 e-i.
Records. — Bogue Sound (5); Andrews gives
no description and his specimens may belong
to Proceraea fasciata reported by many other
workers.
Distribution. — English Channel; Madeira;
Mediterranean; intertidal to 30 m.
Proceraea fasciata (Bosc, 1802)
Proceraea tardigrada Webster, 1879: 27. - An-
drews, 1891a: 282.
Autolytus fasciatus. - Pettibone, 1963a: 141,
Fig. 37 a, b, 38, 39.
Proceraea fasciata. - Gidholm, 1967: 203 (note
only).
Records. — Beaufort sound, intertidal to a
few meters (3, 5, 7, 8, 11, 17).
Autolytus dentalius Imajima, 1966
Autolytus alexandri. - Hartman, 1945: 17, pi. 2:
Fig. 11. (non Malmgren).
Autolytus dentalius Imajima, 1966b: 36, Fig.
7i-l.
Records. — Beaufort Sound, intertidal to a
few meters (11).
Distribution. — North Carolina; central Ja-
pan; intertidal to 10 m.
35
1
1'
2
9'
FAMILY SPHAERODORIDAE
Key to genera and species
Setae compound and falcigerous (EpJiesiella) 2
Setae simple, hooked (Spiiaerodorum) No N.C. record
Only 6 large papillae across dorsum of each segment E. claparedii
Ten to twelve large papillae across dorsum of each segment
(E. minutum) No N.C. record
Ephesiella claparedii (Greeff, 1866)
Spiiaerodorum claparedii. - Fauvel, 1923: 379,
Fig. 149 d,e.
Description. — Body stout and ovoid, 2 mm
long, with 18 segments. Prostomium indistinct,
papillose. Each segment with a transverse row
of six large papillae across dorsum and a band
of smaller papillae across ventrum. No small
papillae among large dorsal ones. Parapodia
uniramous and cylindrical with three oval pa-
pillae distally. About 10 falcigerous compound
setae per foot; blades faintly curved and uni-
dentate.
Remarks. — The single specimen obtained was
a female containing large eggs. It agrees very
well with Fauvel's description except it lacked
the small irregularly arranged papillae among
the large dorsal ones. According to Hartman
(1965b), this species should be referred to Sphae-
rodoridium Liitzen, but according to Pettibone
(1963a), it should be referred to Ephesiella
Chamberlin of which Sphaerodoridium is a
synonym.
Records. — One specimen from 200 m off
Beaufort (*). This is a new record for the
United States.
Distribution. — English Channel and Ireland;
intertidal to a few meters.
FAMILY NEREIDAE
Key to genera and species
1 Parapodia essentially uniramous throughout with one bundle
of setae. Proboscis without chitinous paragnaths or soft
papillae. Three pairs of tentacular cirri. (Lycastopsis) No N.C. record
1' Parapodia biramous after first two with 4-5 parapodial lobes
and two bundles of setae. Proboscis with chitinous parag-
naths or soft papillae. Four pairs of tentacular cirri 2
2 Parapodia with two ventral cirri. No falcigerous setae. [Pro-
boscis with soft papillae only. (Ceratocephale). Dorsal cirri
very long. Prostomium incised] C. loveni
2' Parapodia with one ventral cirrus. Falcigerous setae present
in neuropodia at least 3
3 Probiscis with soft papillae, a few on basal ring sometimes
brown and lightly chitinized. No falcigerous setae in pos-
terior notopodia 4
3' Proboscis with black chitinous paragnaths. Falcigerous setae
sometimes present in posterior notopodia 5
36
4 Proboscis with tufts of soft papillae on maxillary ring (Laeo-
nereis). Anterior parapodia with three notopodial lobes L. culvert
4' Proboscis with one row of soft or lightly chitinized papillae
on oral ring. (Websterinereis) Anterior parapodia with three
notopodial lobes. [Posterior parapodia with neuropodial
lobes well separated from notopodial lobes. No simple anky-
losed neurosetae] W. tridentata
5 Chitinous paragnaths restricted to maxillary ring (Cerato-
nereis). [No notopodial falcigers] 6
5' Chitinous paragnaths present on both basal and maxillary
rings 7
6 Anterior feet with two notopodial lobes. [Dorsal cirrus shorter
than superior lobe] C. irritabilis
6' Anterior feet with three notopodial lobes C. versipedata
7 Paragnaths on group VI as one or more conical points (Nereis) 8
7' Paragnaths on group VI as one or more plain transverse
bars (Perinereis) No N.C. record
7" Paragnaths on group VI as one or two small comblike bars
(Platy nereis). [Notopodial falcigers of posterior feet with
a minute terminal knob] P. dumerilii
8 Posterior notosetae include one or more falcigers (subgenus
Nereis) 9
8' Posterior notosetae all spinigerous (subgenus Neanthes). [An-
terior feet with three notopodial lobes] 11
9 Anterior feet with three notopodial lobes. [Proboscis with
groups VII and VIII forming a band of three to four rows.
Notopodial falcigers with long, lightly serrated blades] N. (Nereis) lamellosa
9' Anterior feet with two notopodial lobes 10
10 Group VI with a close-set group of 3-4 paragnaths; groups
VII and VIII with none. Notopodial falcigers of posterior
feet with short oval blades. [Dorsal cirri very short] N. (Nereis) grayi
10' Group VI with 4 paragnaths in a close-set group and groups
VII and VIII with 3-7 points in a single row. Notopodial
falcigers of posterior feet with long blades. [Two rows of
brown spots on anterior segments] N. (Nereis) riisei
10" Group VI with 4-6 paragnaths and groups VII and VIII with
numerous paragnaths in two to three irregular rows N. (Nereis) falsa
11 Paragnaths on basal ring of proboscis in distinct groups:
V = 1-4; VI = 6-8; VII and VIII = two or three irregular
rows. [Superior lobe of posterior feet lamellar] N. ( Neanthes J succinea
11' Paragnaths on basal ring of proboscis forming a broad and
continuous band of points N. (Neanthes) acuminata
37
Lycastopsis pontica (Bobretzky, 1872)
Lycastopsis tecolutlensis. - Hartman, 1951: 44.
Lycastopsis pontica. - Pettibone, 1963a: 150,
Fig. 41.
Records. — No North Carolina record.
Distribution. — Massachusetts to Virginia and
Mexico: West Indies and Brazil; Mediterranean
and Black Sea; Japan; central California; inter-
tidal and estuarine.
Ceratocephale loveni Malmgren, 1867
Chaunorhynchus loveni. - Hartman, 1942a: 49,
Fig. 83-84.
Ceratocephale loveni - Pettibone, 1963a: 152,
Fig. 42 a-b.
Records. — Five specimens from 40 m off
Beaufort (*).
Distribution. — North Atlantic from Norway
to the North Sea and Iceland to Virginia;
Okhotsk Sea; from 40 to 2,000 m.
Laeonereis culveri (Webster, 1879)
Nereis culveri. - Webster, 1886: pi. 6.: Fig.
25-30, pi. 7: Fig. 31-32.
Laeonereis culveri. - Hartman, 1945: 21. - Petti-
bone, 1971: 14, Fig. 5-7.
Records. — On the shores of Beaufort Sound
(8, 11, 13, 18).
Distribution. — Connecticut to Gulf of Mexico,
Florida, West Indies, Brazil and Uraguay ; inter-
tidal and estuarine in muddy sand.
Websterinereis tridentata (Webster, 1880)
Figure 5a-f
Nereis tridentata Webster, 1880: 113; 1886: 142,
pi. 7: Fig. 33-40.
Ceratonereis tridentata. - Hartman, 1945: 21,
pi. 3: Fig. 3, 4.
Laeonereis n. sp. - McCloskey, 1970: 26.
Websterinereis tridentata. - Pettibone, 1971: 21,
Fig. 8, 9.
Description. — Body slender, 20-30 mm long,
mainly pale but with brown marks at origin
of palps and brown spots above parapodia.
Prostomium (Figure 5a) oval with a slight
groove between antennae. Tentacular cirri very
short, seldom reaching setiger 1. Proboscis
(Figure 5b, c) without paragnaths or soft papil-
lae on maxillary ring but with soft papillae on
basal ring; area VI with one papilla; areas
VII and VIII with one row of seven minute
papillae, the middle three often chitinized and
brown. Anterior feet (Figure 5d) with three
notopodial lobes and a slightly longer dorsal
cirrus. Setigerous lobe of neuropodium with
well marked presetal and postsetal lips. Pos-
terior feet (Figure 5f) with all lobes more
pointed and dorsal cirrus small in juveniles but
longer than superior lobe in adults. Notosetae
as homogomph spinigers in all feet. Neurosetae
include homogomph and heterogomph spinigers
and heterogomph falcigers. Blades of anterior
falcigers fairly long, almost rectangular with
longer spinules distally; blades of posterior
falcigers (Figure 5e) shorter and more hooked
with tip attached back by a tendon.
Remarks. — The generic position of this spe-
cies has caused considerable difficulty. The pa-
pillae on the basal ring of the proboscis are
very small; in some specimens the whole pro-
boscis appears smooth; in others the three
papillae on area VII are brown and chitinized
like small paragnaths. Such specimens might
be refered to Eu nereis except that they lack
notopodial falcigers on posterior feet. A further
discussion will be found in Pettibone (1971:20)
who erected the genus Websterinereis with W.
trident a as the type-species.
Records.— Off Beaufort in 3-40 m (11, 13, 14,
18.20, *)
Distribution. — New Jersey to Florida and the
Gulf of Mexico; 3-40 m.
Ceratonereis irrilablis (Webster, 1879)
Nereis irritabilis Webster, 1879: 231, pi. 5: Fig.
56-64; pi. 6: Fig. 65-69.
Ceratonereis irritabilis. - Hartman, 1945: 20,
pi. 3: Fig. 7-9.
Records. — Common off the shores of Beaufort
Sound and offshore in 10-80 m (3, 5, 11, 13, 18,
19.21, *).
Distribution. — Virginia to North Carolina;
intertidal to 80 m.
:{«
Ceratonereis versipedata Ehlers, 1887
Nereis (Ceratonereis) versipedata Ehlers, 1887:
116, pi. 36: Fig. 5-10. - Monro, 1933b: 256.
Records. — On coral in 10-18 m off Beaufort
(14,20).
Distribution. — North Carolina to Florida and
the West Indies in 10-18 m.
Platynereis dumerilii
(Audouin and Milne-Edwards, 1833)
Platynereis dumerilii. - Fauvel, 1923: 359, Fig.
141 a-f. - Pettibone, 1963a: 154, Fig. 43. -
Day, 1967: 306, Fig. 14.4 d-k.
Records. — Common in mucus tubes attached
to weeds all along North Carolina; intertidal
to a few meters (3, 5, 11, 13, 14, 18).
Distribution. — Cosmopolitan in temperate
and tropical seas; intertidal to 10 m.
Nereis (Nereis) lamellosa Ehlers, 1868
Figure 5k-o
Nereis (Nereis) lamellosa. - Fauvel, 1936: 36. -
Day, 1967: 314, Fig. 14.7. a-t.
Description. — Prostomium broadly triangu-
lar with brownish marks laterally. Proboscis
(Figure 5k, 1) with group I = 1; II = an
oblique double row; III = an oval group of
about 10; IV = a wedge of numerous points;
V = 0-3; VI = a rosette of 8-10; VII and
VIII = three or four irregular rows. Anterior
feet (Figure 5m) with three pointed notopodial
lobes and a longer dorsal cirrus. Middle feet
with only two notopodial lobes; posterior feet
with superior notopodial lobe expanded and last
few feet (Figure 5o) with superior notopodial
lobe broad and lamellar bearing the small dorsal
cirrus at its apex. Anterior notosetae all homo-
gomph spinigers with rather short blades; pos-
terior notosetae mainly homogomph spinigers
but some feet with one or two homogomph fal-
cigers with rather long blades (Figure 5n).
Anterior and posterior neurosetae essentially
similar, including homogomph and heterogomph
spinigers with short blades and heterogomph
falcigers with rather straight blades.
Remarks. — Nereis (Nereis) lamellosa and
Nereis (NeantJies) sueeinea are very similar
apart from the presence of notopodial falcigers
in the former. They occur together in many
parts of the world although N. sueeinea extends
into estuaries while N. lamellosa does not. The
notopodial falcigers of N. lamellosa are not
numerous and readily lose their apices and
their shafts are no stouter than those of the
spinigers. As a result the complete absence of
notopodial falcigers and the separation of N.
sueeinea from N. lamellosa is not easy.
Records. — One specimen on coral in 10 m
off Beaufort (*). This is a new record for the
United States.
Distribution. — Mediterranean, Morocco, Sen-
egal, and South Africa; 10-150 m.
Nereis (Nereis) grayi Pettibone, 1956
Nereis (Nereis) grayi Pettibone, 1956: 282,
Fig. 3; 1963a: 183, Fig. 42 i.
Records. — Five specimens off Beaufort in
20-200 m (*).
Distribution. — Massachusetts; intertidal in
mud to 18 m.
Nereis (Nereis) riisei Grube, 1856
Fig-ure 5g-j
Nereis riisei. - Augener, 1922: 42; 1925: 6. -
Hartman, 1951: 46.
Description. — Body up to 30 mm long; fresh
specimens often with a brown bar across peri-
stome or setiger 2 and dorsolateral spots on
anterior segments. Tentacular cirri often reach-
ing setiger 6. Proboscis (Figure 5g, h) with
area I = 1-3 points in line; II = a double row;
III = an oval group; IV = a wedge; V = 0;
VI = a close-set group of 3-9 points; VII and
VIII = one row of 3-7. Anterior feet (Figure
5i) with two notopodial lobes and a slender
dorsal cirrus longer than short superior lobe.
Posterior feet essentially similar. Notosetae
of posterior feet include a few spinigers and
usually one large homogomph falciger (Figure
5j) with an almost straight, lightly serrated
blade.
39
Figure 5. — Websterinereis tridentata a, head; b and c, dorsal and ventral views of proboscis; d, anterior foot; e,
posterior falciger; f, posterior foot of juvenile. Nereis riisei g and h, dorsal and ventral views of proboscis; i,
anterior foot; j, notopodial falciger. Nereis lamellosa k and 1, dorsal and ventral views of proboscis; m, anterior
foot; n, notopodial falciger; o, posterior foot. Nephtys (Aglaophamus) circinata p, anterior ends; q, anterior view
of foot.
40
Records. — Eight specimens from 40 m off
Beaufort (*).
Distribution. — Florida and Mexico (Vera-
cruz) to the West Indies; on shallow reefs.
Nereis (Nereis) falsa Quatrefages, 1865
Nereis falsa. - Fauvel, 1923: 337, Fig. 129 e-m. -
Day, 1967: 317, Fig. 14.7 k-o (with syn-
onymy).
Nereis pelagica oeeidentalis Hartman, 1945:
20, pi. 4: Fig. 1-6: 1951: 46.
Nereis oeeidentalis. - McCloskey, 1970: 26.
Records. — Sounds and shallow reefs in 7 m
near Beaufort (5, 11, 13, 15, 18, 20, *).
Distribution. — Warm and tropical Atlantic
from France to West Africa and North Carolina
to the Gulf of Mexico; Mediterranean; South
Africa to Madagascar; intertidal to 30 m.
Nereis (Neanthes) succinea
Frey and Leuckart, 1847
Neanthes succinea. - Hartman, 1945: 17, pi. 3:
Fig. 1-2; 1968: 529, Fig. 1-5.
Nereis (Neanthes) succinea. - Pettibone, 1963a:
165, Fig. 44 a-e, 45 a-d. - Day, 1967: 321,
Fig. 14.9. a-e.
Records. — Common on the shores of Beaufort
Sound and offshore in 6.5-40 m (3, 5, 7, 11, 13,
15, 17, 18, 19,20,21, *).
Distribution. — Atlantic from the North Sea
to South Africa and Massachusetts to Uraguay;
Pacific Ocean from California to Panama; Indian
Ocean from Cape Point to Natal; estuarine and
intertidal to 40 m.
Nereis (Neanthes) acuminata Ehlers, 1868
Nereis (Neanthes) caudata. - Fauvel, 1923: 347,
Fig. 135 a-e. - Day, 1967: 321, Fig. 14.9. f-j. -
Hartman, 1968: 525, Fig. 1-5.
Nereis arenaceodentata Moore, 1903: 729, pi.
40: Fig. 1-10. - Pettibone, 1963a: 162, Fig.
44 i, 45 e.
Remarks. — Pettibone (1963a) states that
Nereis caudata Delle Chiaje is a homonym. The
next available name is Nereis acuminata Ehlers.
Records. — Seven specimens from 40 to 80 m
off Beaufort (*).
Distribution. — North Atlantic from the Eng-
lish Channel to Spain and Massachusetts to
Florida; Mediterranean; South Africa; south-
ern California to Mexico; southern Australia,
Tasmania, and New Zealand ; intertidal to 100 m.
FAMILY NEPHTYIDAE
Key to subgenera and species of Nephtys
1 Interramal gills long and involute, curving downwards, then
inwards 2
1' Interramal gills short and revolute, curving downwards, then
outwards (subgenus Nephtys) 4
1" No interramal gills (subgenus Micronephthys) No N.C. record
2 No proboscideal papillae. No ventral cirrus (subgenus Iner-
monephtys). [Dorsal cirri very long] N. (I.) inermis
2' Proboscideal papillae and ventral cirri present (subgenus
Aglaophamus). [Forked setae present] 3
3 Eyes present. Dorsal cirri digitiform. Neuropodium with a
straplike superior lamella and broad postsetal one N. (A.) verrilli
3' Eyes absent. Dorsal cirri flattened. Neuropodium without a
straplike superior lamella, only a large ligulate postsetal
one N. (A.) circinata
11
6
6'
Anterior dorsum with grey segmental bars. [A few short,
geniculate, postaeieular setae with coarse teeth at base of
blade] N.(N.) picta
Dorsum uniformly pale 5
Bases of parapodia with scalelike lamellae covering inter-
segmental junctions both dorsally and ventrally N. (N.) squamosa
Bases of parapodia without scales 6
A red spot in middle of prostomium. Shorter postaeieular
setae with denticles at base of blade N. (N.) bucera
No red spot on prostomium. Postaeieular setae without denti-
cles at base of blade N. (N.) i)icisa
Nephtys ( Inermonephtys) inermis
Ehlers, 1887
Nephthys (Aglaophamus) inermis Ehlers, 1887:
125, pi. 38: Fig. 1-6.
Nephthys inermis. - Fauvel, 1923: 375, Fig.
147. - Hartman, 1940: 234, pi. 39: Fig.
84-86; pi. 40: Fig. 95.
Inermonephtys inermis. - Fauchald, 1968: 16,
pi. 4: Fig. 31-35.
Description. — Body up to 50 mm long. Pro-
stomium bluntly rectangular in front with a
posterior prolongation. Two pairs of antennae
but neither pair obvious; anterior pair bent
down below margins of prostomium, posterior
pair on sides of nuchal grooves at level of sub-
dermal eyes. Proboscis very long and muscular
but without papillae. Jaws as lightly chitinized
ridges far back. First setiger small, bearing a
digitiform dorsal cirrus, a rounded postsetal
lamella on notopodium, and a ventral cirrus
similar to dorsal one. Second setiger similar
but larger. Third setiger with first branchia
originating from base of dorsal cirrus. Para-
podia fully developed at 10th segment. Noto-
podium formed by a blunt setigerous lobe bear-
ing a rudimentary presetal lamella, a large
dorsolateral postsetal lamella and below this,
a very long dorsal cirrus and an involute bran-
chia. Neuropodium formed of a bluntly conical
setigerous lobe bearing a rudimentary presetal
lamella, an even smaller postsetal lamella, and
a long, stout, and tapering ventrolateral cirrus.
Preacicular setae rather long but faintly barred;
postaeieular setae mainly with slender blades
minutely serrated but also a few forked setae.
Remarks. — Fauchald has erected a new genus
Inermonephtys with N. inermis as the type
species. The genus is distinguished by the lack
of proboscideal papillae, the character of the
jaws and a different interpretation of the pro-
stomial appendages. He regards the first pair
of antennae as missing, the second pair as ven-
tral and states that: "The nuchal organs are
very well developed and each is equipped with
one long, digitiform eversible process." His
Figure 34 shows these digitifprm processes as
separate from the prostomium, whereas my
specimens show that they arise from the mar-
gins of the prostium, close to the subdermal
eyes. For this reason I regard these processes
as the second pair of antennae, further back
than usual, but nonetheless homologous with
the second pair of antennae of other nephtyids.
Inermonephtys is obviously related to Agla-
ophamus and both are here regarded as sub-
genera of Nephtys.
Records.— Off Beaufort in 160-450 m (*).
Distribution. — Florida and tropical Atlantic;
Pacific; Mediterranean; 0-450 m.
Nephtys (Aglaophamus) verrilli
Mcintosh, 1885
Nephthys verrilli Mcintosh, 1885: 163, pi. 26:
Fig. 6, 7; pl.32A: Fig. 8.
Aglaophamus dicirris Hartman, 1945: 22;
1950; 122, pi. 18, Fig. 1-8.
i'Z
Aglaophamus verrilli. - Pettibone, 1963a: 190,
Fig. 48 c, d.
Nephtys dicirris. - Day, Field, and Montgomery,
1971: 121.
Remarks. — Hartman (1950: 121) referred N.
verrilli to Aglaophamus dibranchis (Grube) on
the grounds that they both have 14 rows of
papillae on the proboscis but Mcintosh clearly
stated that N. verrilli has 22 rows of papillae
as does A. dicirris.
Records. — Beaufort, common on the shores
of the sounds and offshore down to 200 m (11,
17,21, *).
Distribution. — Chesapeake Bay to the Gulf
of Mexico; California to the Pacific coast of
Mexico; New Zealand; intertidal to 200 m.
Nephtys (Aglaophamus) circinata
Verrill, 1874
Figure 5p, q
Aglaophamus circinata. - Pettibone, 1963a:
192, Fig. 48 a.
Description. — Length up to 50 mm. Pro-
stomium (Figure 5p) with well-developed an-
terior antennae and subequal, ventral ly situated
posterior ones. No eyes even in juveniles. Pro-
boscis with 14 rows of papillae. Setiger 1 with
a large ventral cirrus but no dorsal one. Bran-
chiae from setiger 2 or 3, becoming long and
involute (Figure 5q), with a large flattened and
distally pointed dorsal cirrus. Setigerous lobes
with pointed projections over tips of acicula.
Postsetal lamella of notopodium deeply notched
with superior part large and projecting dorso-
lateral^ and inferior portion small and oval.
Presetal lamella of neuropodium divided, form-
ing a rudimentary superior part and an oval
inferior part. Postsetal lamella large and ligu-
late. Ventral cirri flattened and spear-shaped.
Preacicular setae long and barred; postacicular
setae very long and minutely spinulose over
most of blade. No forked setae.
Remarks. — This species approaches N. (A.)
peruana Hartman but the latter lacks the di-
vided postsetal notopodial lamella.
Records. — Five specimens from 200 to 450
m off Beaufort ( * ) .
Distribution. — Gulf of St. Lawrence to Long
Island Sound; in 15-787 m.
Nephtys pitta Ehlers, 1868
Nephtys picta. - Hartman, 1945: 22; 1950: 103;
1951: 49, pi. 10: Fig. 4. - Pettibone, 1963a:
195, Fig. 49 c, 50 c-f.
Records. — Very common on protected shores
of North Carolina and in dredgings down to
200 m (3,5, 11, 13, 17, 18,21, *).
Distribution. — Massachusetts to Florida and
the Gulf of Mexico; intertidal to 200 m.
Nephtys squamosa Ehlers, 1887
Nephtys squamosa. - Hartman, 1940: 237, pi.
41: Fig. 98, 99; 1968: 597, Fig. 1, 2. - Petti-
bone, 1963a: 194, Fig. 47 e. - Nonato and
Luna, 1970b: 71, Fig. 27-31.
Records. — Common off Beaufort in 80-200 m
(21, *).
Distribution. — Massachusetts to Florida,
West Indies and Brazil; Morocco; southern
California; Atlantic and Pacific coasts of tropi-
cal America; 26-200 m.
Nephtys incisa Malmgren, 1865
Nephthys iucisia. - Fauvel, 1923: 369, Fig. 144
a, b.
Nepthys incisa. - Pettibone, 1963a: 198, Fig.
49 a, b, 51a.
Records. — Beaufort; intertidal (9).
Distribution. — North Atlantic from Sweden
to Portugal and Greenland to Chesapeake Bay;
Mediterranean; intertidal to 1,750 m.
Nephtys bueera Ehlers, 1868
Nephtys bueera. - Hartman, 1950: 105. - Petti-
bone, 1963a: 196, Fig. 49 d, 50 a, b, 51 d.
Records. — Common at low tide in Beaufort
Sound and offshore in 10-200 m (5, 8, 9, 11, 17,
18,21, *).
Distribution. — Massachusetts to North Caro-
lina; 0-200 m.
43
FAMILY GLYCERIDAE
Key to genera and species
1 Superior setae as simple capillaries and inferior ones com-
pound and spinigerous. Two presetal lobes (Ghjcera) 2
1 All setae compound and spinigerous. One presetal lobe (Hemi-
podus). [Proboscideal papillae include both oval and digiti-
form types] H. roseus
2 Gills present but sometimes small and retractile. Parapodia
with two postsetal lobes 3
2' Gills entirely absent. One or two postsetal lobes 4
3 Two separate cirriform gills, one on dorsal edge of para-
podium and one on ventral edge Ghjcera dibranchiata
3' One retractile branching gill arising from posterodorsal base
of foot; [postsetal lobes subequal] Ghjcera americana
4 Superior presetal lobe minute. [Papillae on proboscis long
and smooth] 5
4' Superior presetal lobe at least half as long as inferior one 6
5 Prongs of jaw support deeply notched, almost separate Glycera papillosa
5' Prongs of jaw support only slightly notched, almost fused Glycera capitata
6 Jaw support deeply forked, forming two slender divergent
prongs 7
6' Jaw support completely fused forming one asymmetrical piece.
[Papillae on proboscis with 8-10 rings] Ghjcera oxycephala
7 Papillae on proboscis smooth 8
7' Papillae on proboscis with 14-16 rings. [Parapodia with one
emarginate postsetal lobe (Glycera tennis)] No N.C. record
8 Two low postsetal lobes or one emarginate lobe on all feet Glycera tesselata
8' One low postsetal lobe on first few feet, dividing to form
a conical superior lobe and a low rounded median posterior
lobe on subsequent feet Glycera asymmetrica
Hemipodus roseus Quatrefages, 1865 Description.— Body slender, uniformly pale
in alcohol, 15-38 mm long by 0.8 mm wide with
Figure 6a-c 80-100 segments. Prostomium as a slender cone
with about 10 indistinct annulations and 4 small
Hemipodus roseus. -Arwidsson, 1899: 28, pi. 2: terminal antennae. No eyes. Proboscis with
Fig. 23, pi. 4: Fig. 58. - Hartman, 1950: 81 smooth papillae of two types (Figure 6b);
(table of characters). numerous shorter tongue-shaped forms and
Hemipodus borealis. - Hartman, 1950: 81; 1968: fewer digitiform papillae about five times as
637, Fig. 1, 2. long as broad. Four falcate jaws each with a
44
slender rodlike support (Figure 6a). All para-
podia uniramous bearing only spinigerous setae.
Anterior parapodia (Figure 6c) with an ovoid
dorsal cirrus well above the elongate setigerous
base, the conical ventral cirrus arising from
its ventral margin. Presetal lobe tapered and
about half as long as the setigerous base; post-
setal lobe low and rounded. Posterior feet with
a shorter presetal lobe and a longer, more
pointed ventral cirrus.
Remarks. — This is the first record of the
genus Hemipodus from the Atlantic since all
of the 20 species that have been described are
confined to the Pacific. However, the name
Hemipodus roseus has been used with consider-
able hesitation for the majority of the descrip-
tions are very similar and a reexamination of
the types is obviously necessary. A few species
appear to have characteristic proboscideal pa-
pillae but the rest are distinguished by dif-
ferences in the shape of the parapodia which
are known to change along the length of the
body even in a single specimen. The type species
of the genus is Hemipodus simplex (Grube) and
Ehlers (1901) regarded this as the only valid
species described before 1900. He included seven
species in the synonymy and among them was
H. roseus Quatrefages. Arwidsson (1899), how-
ever, argued that H. roseus was a valid species
and his figures of the presetal lobe of the para-
podia are closer to my specimens from North
Carolina than are those of Ehlers for H. sim-
plex. Knox (1960) has also figured the pro-
boscideal papillae of H. simplex as all flattened
and triangular in outline whereas my specimens
have papillae of two types as shown in Figure
6b. For these reasons I have used the name
H. roseus.
I have also compared the North Carolina
specimens with a specimen of H. borealis John-
son from Fox Island, Wash., which Dr. Hart-
man kindly sent to me. H. borealis is 38 mm
long and thus twice the size of the North
Carolina specimens but there are no other im-
portant differences either in the shape of the
parapodia or the proboscideal papillae or the
jaw supports. In brief, H. borealis appears to
be a synonym of H. roseus.
Records. — Common in 3-20 m of Beaufort
(21, *).
Distribution. — Pacific coast of the Americas
from Washington to Chile; intertidal to 18 m.
Glycera dibranehiata Ehlers, 1868
Glycera dibranehiata. - Hartman, 1945: 23;
1950: 70, pi. 10: Fig. 9, 10; 1968: 621,
Fig. 1-4. - Pettibone, 1963a: 215, Fig. 56.
Records. — Common from Cape Hatteras to
Beaufort; intertidal to 20 m (5, 9, 11, 13, 17,
18,21, *).
Distribution. — Gulf of St. Lawrence to Flori-
da and the Gulf of Mexico; central California
to the Pacific coast of Mexico; intertidal to
400 m.
Glycera americana Leidy, 1855
Glycera americana. - Hartman, 1950: 73; 1968:
613, Fig. 1. - Pettibone, 1963a: 213, Fig.
54 a-e. - Nonato and Luna, 1970b: 71, Fig.
16.
Records. — Common from Cape Hatteras to
Beaufort; intertidal to 120 m (2, 3, 5, 7, 8, 9,
11, 13, 15, 17, 18,21, *).
Distribution. — Massachusetts to Argentina
and the Strait of Magellan; British Columbia
to Peru; South Australia and New Zealand;
intertidal to 310 m.
Glycera papillosa Grube, 1857
Glycera papillosa. - Day, 1967: 358, Fig. 16.1. j-1.
Description. — Length 20-30 mm. Prostomium
with eight rings. Proboscis with numerous long,
slender, and smooth papillae and few ovoid
forms. Jaw supports deeply forked; shorter
prong united to longer one by a deeply notched
pale area. Parapodia with two presetal lobes,
superior one very small and not immediately
evident; one low rounded postsetal lip. No
branchiae.
Remarks. — This species is generally similar
to G. capitata but may be distinguished by the
shape of the jaw supports.
Records. — Five specimens off Beaufort in
20-160 m (20, *). This is a new record for the
United States.
Distribution. — South Africa and Chile; inter-
tidal to 200 m.
45
Figure 6. — Hemipodus voseits a, jaw and support (black); b, proboscideal papillae; c, posterior view of foot. Glycera
asymmetrica n. sp. d, jaw and supports (black); e, proboscideal papillae; f, posterior view of anterior foot; g,
posterior view of posterior foot.
Glycera capitata Oersted, 1843
Glycera capitata. - Fauvel, 1923: 385, Fig. 151
a-e. - Hartman, 1950: 76, pi. 11: Fig. 1-4;
1968: 617, Fig. 1-4. - Pettibone, 1963a:
211, Fig. 53.
Records. — One small specimen from 120 m
off Beaufort (*).
Distribution. — Arctic; North Atlantic from
Greenland to North Carolina and Norway to
Madeira; Mediterranean; South Atlantic; Ant-
arctic ; Pacific from Alaska and Japan to Mexico ;
intertidal to 3,800 m.
Glycera oxycephala Ehlers, 1887
Glyct ra oxycephala. Hartman, 1940: 248, pi. 37:
Fig. 74, 75, pi. 43: Fig. 122-124, pi. 44:
Fig. 125; 1950: 70, pi. 10: Fig. 3, 4, text
Fig. 3; 1968: 625, Fig. 1-6.
Records. — Common off Beaufort in 3-120 m
(21,*).
Distribution. — Atlantic coast of United States
from North Carolina to the tropics and Pacific
Coast from Oregon to Panama; Galapagos
Islands; intertidal to 800 m.
Glycera tesselata Grube, 1863
Glycera tesselata. - Fauvel, 1923: 387, Fig.
152 a-c. - Hartman, 1950: 77, pi. 10: Fig.
11, 1968: 633, Fig. 1-3. - Day, 1967: 359,
Fig. 16.2. a-c.
Description. — Body up to 35 mm long. Pro-
boscis covered with long smooth papillae. Jaw
16
supports very deeply forked, the two slender
prongs being almost separate. Parapodia with
two presetal lobes with the superior one slightly
smaller. Either a single emarginate posterior
lip or two low, rounded lobes. No branchiae.
Records. — Seven specimens from 6.5-120 m
off Beaufort (20, *) This is a new record for
the Atlantic coast of the United States.
Distribution. — Cosmopolitan in temperate
and tropical seas; 5 to over 500 m.
Glycera asymmetrica New Species
Figure 6d-g'
Holotype. — USNM 43148; three paratypes,
USNM 43149.
Description. — Body rounded in section, ta-
pered at extremities and up to 50 mm long.
Prostomium elongate and tapered with about
15 indistinct rings but no visible eyes. Pro-
boscis covered with numerous long smooth
papillae and a few ovoid ones (Figure 6e). Jaw
supports (Figure 6d) formed of two unequal
rami quite separate except at point of contact
with jaw. Shorter ramus expanded basally,
longer ramus more slender. Parapodia without
gills. All feet with two digitiform presetal lobes,
the inferior one slightly longer. Postsetal lobes
changing along the body; first 20 feet with a
single truncate postsetal lobe (Figure 6f), pos-
terior feet with a small posterodorsal digitiform
lobe above a low rounded postsetal lobe (Figure
6g); middle feet changing from anterior to
posterior form.
Remarks. — The posterodorsal digitiform lobe
of the posterior feet is in the same position as
the gill of Glycera con valuta, but it is definitely
not a gill and is never as long as the presetal
lobes. Apart from this queer asymmetrical lobe
and rather shorter proboscideal papillae, this
species resembles G. tesselata. In particular the
form of the jaw supports is very similar. I am
indebted to Dr. Pettibone for informing me that
this species resembles G. sphyrabrancha
Schmarda from Jamaica the type of which was
redescribed by Augener, (1925: 29, Fig. 1). The
jaw supports, proboscideal papillae and gen-
eral appearance of the feet is similar. However
Augener describes a nonretractile gill arising
from the dorsal edge of the 30th and later feet
which becomes much longer and stouter than
the presetal lobes. It is possible, but unlikely,
that this is the same as the "small posterodor-
sal digitiform lobe" described above, which as
stated, is definitely not a gill.
Records. — Four specimens off Beaufort in
20 m (*).
FAMILY GONIADIDAE
Key to genera and species
1 Both anterior and posterior parapodia uniramous and bear
only compound setae. Chevrons present (Progoniada). [Body
mottled brown] P. regular is
1' Anterior parapodia uniramous, posterior ones with simple
notosetae and compound neurosetae. Chevrons present or
absent 2
2 A series of V-shaped chevrons at base of proboscis which is
covered with small scattered papillae 5
2' No V-shaped chevrons at base of proboscis; proboscideal
papillae arranged in regular rows with much longer ones
dorsolaterally 3
3 Compound setae either all falcigerous or some falcigerous
and some spinigerous. Notosetae of posterior segments
47
arising directly from body wall above dorsal cirrus (Gonia-
dides). [Notosetae as stout hooks. Eighteen or nineteen
anterior segments] G. carolinae
3' Compound setae all spinigerous. Notosetae of posterior seg-
ments arising from bilobed notopodia (Glycinda) 4
4 Twenty-four anterior segments without notosetae Glycinde solitaria
4' Thirty -seven anterior segments without notosetae Glycinde nordmanni
5 Superior compound setae spinigerous and inferior ones fal-
cigerous {Goniadella); [28-30 anterior uniramous segments] Goniadella gracilis
5' All compound setae spinigerous (Goniade) 6
6 Posterior notopodia bear 2-3 acicular notosetae. [Neuropodia
with 2 presetal lobes; jaws with an arc of 11 denticles above
and 9 below] Goniada teres
6' Posterior notopodia bear several capillary notosetae 7
7 Anterior feet (e.g., 15th) all with one presetal lobe. [Jaws with
few (3-5) denticles dorsally and the same number ventrally] Goniada maculata
T Anterior feet (e.g., 15th) with two presetal lobes 8
8 Papillae on proboscis with a prominent beak Goniada littorea
8' Papillae on proboscis squat and flanged, not beaked 9
9 Notosetae appear about segment 40-45. Jaws with 2-4 den-
ticles dorsally and about 10 ventrally Goniada brunnea
9' Notosetae appear about segment 30-37. Jaws with 15-25 den-
ticles dorsally and an equal number ventrally Gondki norvegica
Progoniada regularis Hartman, 1965
Progoniada regularis Hartman, 1965a: 100, pi.
16: Fig. a-f.
Records. — Two specimens in 450 m off Beau-
fort (*).
Distribution. — Western Atlantic from New
England to northwestern South America in
600-5,000 m.
Goniadides carolinae New Species
Fitrure 7a-h
Goniadides n. sp. Day, Field, and Montgomery,
1971: 121
Holotype.— USNM 43389; 40+ paratypes,
USNM 43390.
Description. — Body slender and threadlike,
16 mm long for 85 segments. Color pale with
light brown spots above and below parapodial
bases. Prostomium (Figure 7a) long and ta-
pered with eight distinct rings and four bi-
articulate terminal antennae. No eyes. Base of
proboscis smooth and without chevrons but
distal part with regular rows of dissimilar
papillae as in the genus Glycinde. As seen in
section (Figure 7h), middorsal row I absent;
dorsolateral band II formed of four alternating
rows of strongly chitinized and curved papillae;
lateral row III as a single series of low cones;
ventrolateral row IV similar to III; midventral
row V as a single series of minute cones each
slightly beaked. All papillae with apical pores.
End of proboscis (Figure 7d) armed with a pair
of ventrolateral macrognaths each bearing four
48
teeth, a single midventral micrognath, and a
dorsolateral arc of micrognaths.
Body divided into an anterior region of 18
uniramous segments and a posterior region of
numerous rather flattened and biramous seg-
ments. Distinction between regions not obvious.
Anterior parapodia (Figure 7b) with a tapered
dorsal cirrus, a long setigerous trunk bearing
a tapered presetal lobe and a low, rounded
postsetal lip and a tapered ventral cirrus arising
from base of trunk. Posterior parapodia (Figure
7c) characterized by presence of one or two
short hooked notosetae (Figure 7e) emerging
from medial margin of reduced dorsal cirrus
now representing the notopodium. Neuropodium
and ventral cirrus similar to setigerous trunk
and ventral cirrus of anterior segments. Setae
of anterior segments and neurosetae of posterior
segments identical; three superior ones being
spinigerous with heterogomph shaft-heads and
lightly serrated blades (Figure 7f) and one or
two inferior ones being falcigerous (Figure 7g)
with small blades ending in blunt tips.
Pygidium brownish with a pair of long anal
cirri.
Remarks. — The characters of the paratypes
were very constant, the only differences from
the holotype that were noted being the presence
of 5 micrognaths in the dorsolateral arc in-
stead of 9 and the anterior region consisting
of 19 segments instead of 18.
In general G. carolinae resembles a small
Glycinde but differs from that genus in the
possession of falcigerous setae, a reduced noto-
podium in the posterior region and possibly in
the proboscideal papillae although the range
of variation of these organs in Glycinde is un-
certain. The genus Goniadides was erected by
Hartmann-Schroder (1960) for G. aciculata
from the Red Sea. Regarding the proboscideal
papillae she stated : "Proboscidiale Organe wenig
unterschiedlich." In 1962 she described G. fal-
cigera from Peru with all neurosetae falcigerous
and proboscideal papillae of several types like
those of Glycinde although she does not men-
tion that genus. Goniadides carolinae is closer
to G. acicula but differs in the structure of the
notopodial hooks and the proboscideal papillae.
Records. — Common on sand in 10-20 m off
Beaufort (21, *). This is the first record of the
genus from the Atlantic and the coast of the
United States.
Glycinde solitaria (Webster, 1880)
Glycinde solitaria. - Hartman, 1950: 54, pi. 7:
Fig. 1-15. - Pettibone, 1963a: 222, Fig.
56h-n.
Records. — Cape Hatteras area to Beaufort,
intertidal (11, 18).
Distribution. — New Jersey to North Carolina
and Puerto Rico; intertidal to 47 m.
Glycinde nordmanni (Malmgren, 1865)
Figure 7i-k
Eone nordmanni. - Fauvel, 1923: 394, Fig.
155 h-n.
Glycinde nordmanni. — Arwidsson, 1899: 50, pi.
3: Fig. 45-47, pi. 4: Fig. 64, 65. - Hartman,
1950: 47 (key only).
Description. — Body about 30 mm long, olive
green in alcohol with midventral spots on ab-
dominal segments; prostomium long and tapered
with eyespots on both basal and distal rings.
Proboscis with the usual longitudinal rows of
dissimilar papillae; dorsolateral bands long,
clawlike and well chitinized, lateral and ventral
rows low and soft. Jaws with a pair of small
macrognaths ventrally and an arc of 15-25
micrognaths dorsally. No ventral micrognaths.
Body divided into an anterior region of 36-39
uniramous segments and a posterior region of
numerous, rather flattened biramous segments.
Anterior feet (Figure 7i) with a tapered dorsal
cirrus not incised basally, a parapodial trunk
bearing one presetal and one postsetal lobe of
equal length and a digitiform ventral cirrus.
Setae all compound and spinigerous and arise
between partly fused presetal and postsetal
lobes. Posterior feet (Figure 7j) with a small
notopodium and much larger neuropodium.
Notopodial lobes and dorsal cirrus subequal.
Notosetae (Figure 7k) short, stout, and acicular,
the bluntly hooked end surmounted by a dag-
ger-shaped guard. Neuropodia generally simi-
lar to setigerous lobes of anterior feet but pre-
setal and postsetal lobes shorter and more
pointed and ventral cirrus stouter and more
distal in origin. Neurosetae spinigerous like
those of anterior feet.
49
Figure Ir-Goniadides carolinae n. sp. a, head with proboscis partly extruded; b, posterior view of anterior foot;
c posterior view of posterior foot; d, end of proboscis with jaws; e, hooked posterior notoseta; f, spinigerous seta;
g falcigerous seta; h, section of proboscis with papillae. Glycinde nordmanni i, anterior foot; j, posterior toot;
k] acicular notoseta. Goniada teres 1. proboscidea] papilla; m, posterior view of anterior foot; n, posterior view ot
i-.- ti nor foot.
50
Record*. — Two specimens from 80 to 120 m
off Beaufort (*). This is a new record for the
United States.
Distribution. — Western Europe; 5-139 m.
Goniadella gracilis (Verrill, 1873)
Goniadella gracilis. - Hartman, 1950: 42, pi. 5:
Fig. 4-8. - Pettibone, 1963a: 220, Fig. 56
a-g. - Day, 1967: 368, Fig. 16.4. o-t.
Records. — Off Beaufort in 450 m (*).
Distribution. — Massachusetts to Rhode Is-
land; Irish Sea; South Africa; intertidal to
450 m.
(21, *). This is a new record for the United
States.
Distribution. — Jamaica.
Goniada maculata Oersted, 1843
Goniada maculata. - Fauvel, 1923: 392, Fig.
154 a-g. - Hartman, 1950: 20, pi. 1: Fig.
7, 8. - Pettibone, 1963a: 225, Fig. 58. -
Day, 1967: 367, Fig. 16.4. k-n.
Records. — Twelve specimens off Beaufort in
80-1,650 m (21, *).
Distribution. — Cosmopolitan from low tide
to 2,000 m.
Goniada teres Tread well, 1931
Figure 71 -n
Goniada teres. - Hartman, 1950: 33.
Description. — Body up to 80 mm long, olive
green in alcohol with midventral spots on ab-
dominal segments. Prostomium long and ta-
pered with 10 rings and 2 pairs of minute
terminal antennae. Proboscis very long and
covered with squat papillae (Figure 71) each
with a slit leading to a central pore and a horse-
shoe-shaped flange. Base of proboscis with 10
chevrons. Jaws include a pair of ventrolateral
macrognaths, a ventral arc of 5-9 micrognaths
and a dorsal arc of 10-15 micrognaths.
Body divided into an anterior region of 43
uniramous segments, 8-9 transitional segments
with bilobed notopodia but no notosetae, and a
posterior region of numerous biramous seg-
ments with notosetae. Sometimes 51 anterior
uniramous segments but no transitional seg-
ments. Anterior parapodia (Figure 7m) fully
developed from 15th segment. Each with a
tapered dorsal cirrus, a setigerous trunk bearing
two digitiform presetal lobes, and a single post-
setal lobe and below these a long ventral cirrus.
Setae spinigerous with long blades. Posterior
parapodia (Figure 7n) with a small bilobed
notopodium bearing two or three blunt acicular
setae and a neuropodium generally similar to
setigerous lobe of anterior feet but with shorter,
broader, and more pointed presetal and post-
setal lobes.
Records. — Common off Beaufort in 20-200 m
Goniada littorea Hartman, 1950
Goniada littorea Hartman, 1950: 23, pi. 3: Fig.
1-10.
Records. — Common off Beaufort in 3-160 m
(21, *). This is the first record from the Atlantic
coast of the United States.
Distribution. — Southern California; intertidal
to 160 m.
Goniada brunnea Treadwell, 1906
Goniada brunnea. - Hartman, 1950: 17, pi. 1:
Fig. 1-6, pi. 4, Fig. 1, text Fig. 1; 1968:
653, Fig. 1-4. - Pettibone, 1963a: 228, Fig.
57 a, b.
Records. — From the shore in the Cape Hat-
teras area and questionably in 200 m off Beau-
fort (18, ? *).
Distribution. — Massachusetts to North Caro-
lina and Alaska to southern California and
Hawaii; intertidal to 1,680 m.
Goniada norvegica Oersted, 1845
Goniada norvegica. - Fauvel, 1923: 393, Fig.
155 a-g. - Pettibone, 1963a: 227, Fig. 59.
Records. — One specimen from 200 m off
Beaufort (*).
Distribution. — North Atlantic from Norway
to West Africa and Iceland to the West Indies;
Mediterranean; from 40 to 900 m.
51
FAMILY EUNICIDAE
Key to genera and species
Three antennae. No tentacular cirrus. No branchiae. (Lysi-
dice). [Maxilla II with 4 teeth] 2
Five antennae. Tentacular cirri present or absent. Branchiae
present 3
Eyes oval L. ninetta ninetta
Eyes reniform L. ninetta collaris
Tentacular cirri present (Eunice) 4
Tentacular cirri absent (Marphysa). [Branchiae from about
20th foot to posterior end. Compound setae spinigerous.
Acicular setae bidentate] M. sanguined
Acicular setae tridentate. [Branchiae from setiger 4-7 to end
of body. Acicula with truncate or almost bilobed ends.
Antennae deeply annulated] E. antenuata
Acicular setae bidentate 5
Branchiae from setiger 3 to middle of body. Antennae and
dorsal cirri weakly annulated. Acicula bluntly pointed E. websteri
Branchiae from about setiger 25 to end of body. Antennae and
dorsal cirri smooth. Acicula with expanded, fist-shaped
ends E.filamentosa
Lysidice ninetta ninetta
Audouin and Milne-Edwards, 1833
Lysidice ninetta. - Fauvel, 1923: 411, Fig. 162
a-f. - Nonato and Luna, 1970a: 84. - Day,
1967: 403, Fig. 19.8. g-i.
Records. — Shallow reefs off North Carolina
(14).
Distribution. — English Channel; Mediterran-
ean; Angola; Brazil; ? Indian Ocean; intertidal
to 50 m.
Lysidice ninetta collaris Grube, 1870
noted, L. ninetta and L. collaris differ only in
the shape of the eyes which are reniform in
L. ninetta and oval in L. collaris. Fauchald states
that the degree of ocular pigmentation is a
function of size and regards the two as synony-
mous. My observations on worms of many sizes
does not agree with this. Moreover L. collaris
is restricted to subtropical or tropical waters
while L. ninetta is cosmopolitan. For these rea-
sons I have retained L. collaris as a subspecies
of L. ninetta.
Records. — Two specimens on coral in 18 m
off Beaufort (20, *).
Distribution. — Circumtropical; intertidal to
50 m.
Lysidice collaris Grube, 1870: 495. - Gravier,
1900: 272, pi. 14: Fig. 93-95, text Fig. 144-
147. - Day, 1967: 402, Fig. 17.8. a-f.
Lysidice ninetta. - Fauchald, 1970: 52.
Remarks — As many earlier workers have
Marphysa sanguinea (Montagu, 1815)
Marphysa sanguinea. - Fauvel, 1923: 408, Fig.
161 a-h. - Pettibone, 1963a: 236, Fig. 62. -
Day, 1967: 396, Fig. 17.5. u-y. - Hartman,
1968: 733, Fig. 1-5.
52
Records. — Common on intertidal mudbanks
from Cape Hatteras to Beaufort Sound (3, 5, 7,
11,13, 15, 17, 18,20, *).
Distribution. — Cosmopolitan in temperate and
subtropical waters. Mainly in intertidal mud
flats but may extend a few meters below.
Eunice antennata (Savigny, 1820)
Eunice antennata. - Fauvel, 1953: 240, Fig. 118
f-g. - Day, 1967: 384, Fig. 17.2. k-q. -
Hartman, 1968: 711, Fig. 1-5.
Eunice rubra. - Hartman, 1945: 24; 1951: 55. -
Nonato and Luna, 1970b: 81.
Remarks. — A careful comparison of E. an-
tennata from South Africa and E. rubra from
North Carolina showed that the two are identi-
cal. In both, the acicula are pale with bevelled
and truncate or almost bilobed ends.
Records. — Beaufort, intertidal to 40 m on
rock or coral (5, 11, 13, 20, *).
Distribution. — Circumtropical; intertidal to
over 50 m, often on coral.
Head notched in front; antennae with long,
poorly marked annulations; tentacular cirri and
anterior dorsal cirri also long and annulated.
Branchiae from setiger 3, at first with a single
filament but soon increasing to a maximum of
15 filaments and ending about middle of body
(setiger 53 in type). Acicular setae one or two
per foot, usually bidentate but occasionally
with an indication of a third tooth. Compound
setae falcigerous, strongly bidentate, with
bluntly ending guards.
Remarks. — Fauchald has shown that Web-
ster's original name E. lougicirrata is preoccu-
pied and that E. websteri belongs to a group of
closely related species whose differences he
tabulated although some of the characters men-
tioned are of minor significance. E. websteri is
also very close to E. pennata (Muller) from which
it differs in having blunt instead of pointed
guards to the falcigerous setae, in color mark-
ings, and in having articulated dorsal cirri.
Records. — Two specimens off Beaufort in
18-40 m (20 *).
Distribution. — Bermuda; intertidal.
Eunice websteri Fauchald, 1969
Eunice websteri Fauchald, 1969: 12, Fig. 6 (with
synonymy).
Eunice lougicirrata Webster, 1884: 31, pi. 12:
Fig. 74-80.
[Non] Eunice (Nicidion) lougicirrata Kinberg,
1865: 564.
Eunice pennata. - McCloskey, 1970: 24.
Description. — Body up to 120 mm long with
white bars on setigers 3 and 8 when fresh.
Eunice filamentosa Grube, 1856
Eunice filamentosa. - Monro, 1933b: 65, text
Fig. 27. - Hartman, 1944a: 107, pi. 6: Fig.
123-126. - Day, 1967: 392, Fig. 17.5. f-h.
Records. — Reefs and stones off Beaufort in
5-20 m (14,20, *).
Distribution. — Tropical Atlantic from North
Carolina and Florida to the West Indies and
Ghana and Angola on West Africa; Pacific coast
of Mexico and Galapagos Islands; South Africa
(Natal); intertidal to 120 m.
FAMILY ONUPHIDAE
Key to genera and species
1
r
2
2'
Tentacular cirri present 2
Tentacular cirri absent. [Gills as simple filaments, (Hyalinoe-
cia). Tube quill-like and translucent. Gills from setiger 24
onwards] H. tubicola
Gills from setiger 4 or 5 with filaments arranged in spirals on
a central axis. (Diopatra) 3
Gills as single filaments or pectinately branched 4
53
3 Body brown but without a color pattern. Pseudocompound
setae of setigers 1-3 bidentate. Comb-setae with about 20
teeth D. cuprea cuprea
3' Branchiferous segments with brown crossbars. Pseudocom-
pound setae of setigers 1-3 either bidentate or tridentate
Comb-setae about 25 teeth D. cuprea spiribranchis
4 Setae of first three feet very long, with spiny shafts and
hooked tips (Rhamphobranchium) . [Branchiae from setiger
6. No compound spinigerous setae] R. atlanticum
4 ' Setae of first three feet not much longer than subsequent ones
and never with spiny shafts (Onuphis) 5
5 Gills in middle of body as a single filament arising from the
dorsal cirrus (subgenus Nothria) 6
5' Gills in middle of body as two or more filaments arising from
the dorsal cirrus (subgenus Onuphis) 7
6 Body markedly flattened and first two feet elongated. Tube
flat with large shell fragments. Gills from setiger 11-13 O. (N.) conchylega
6' Body normal and first two feet not elongated. Tube not flat.
Gills from setiger 6-7. [2-3 spinigerous compound setae in
7th-10th foot. Segments barred in brown] O. (N.) pallidula
7 Gills start on setiger 1. Ceratophores of occipital antennae
with more than 11 rings O. (0.) eremita
T Gills start on setiger 6-8. Ceratophores of occipital antennae
with less than 10 rings 8
8 One or two spinigerous compound setae in 7th-12th foot.
[Anterior ventrum minutely speckled] 0. (O.) nebulosa
8' No spinigerous compound setae 9
9 Ventral cirri cirriform only on first two feet. Pseudocompound
setae tridentate 0. (O.) microcephala
9' Ventral cirri cirriform on first four of five feet. Pseudocom-
pound setae bidentate O. (0.) magna
Hyalinoeeia tubicola (Muller, 1776) Diopatra cuprea cuprea (Bosc, 1802)
Hyalinoecia tubicola. - Fauvel, 1923: 421, Fig. Diopatra cuprea. - Hartman, 1944a: 54, pi. 1:
166 i-q. - Pettibone, 1963a: 254, Fig. 65 d. - Fig. 9-14. - Pettibone, 1963a, 250, Fig. 66.
Day, 1967: 411, Fig. 17.9. 1-r. Diopatra cuprea cuprea. - Day, 1967: 417, Fig.
17.12. a-d.
Records. — Common on the continental slope
off Beaufort at 450-600 m (17, *). Records. — Common in Beaufort Sound on
Distribution. — Cosmopolitan below 200 m on intertidal banks and shallow dredgings offshore
sandy mud. to 20 m (2, 3, 4, 5, 7, 8, 9, 11, 13, 15, 17, 18, 21,
*).
Distribution. — Intertidal to 30 m on quiet,
54
sandy shores from Massachusetts to Brazil;
tropical West Africa; tropical Indian Ocean.
Diopatra cuprea spiribranchis Augener, 1906
Diopatra spiribranchis Augener, 1906: 145, pi.
5: Fig. 88-96. - Nonato and Luna, 1970b: 74.
Description. — Length up to 260 mm. Tube
muddy, fragile, without shell fragments. Body
pale with two brown bars close together on
posterior margins of branchiferous segments.
Ceratophores of occipital antennae with about
10 rings. Pseudocompound hooks of first three
feet strongly bidentate or even tridentate with
a slender third tooth. Comb-setae with 20-25
teeth. Ventral cirri cirriform on first four feet.
Remarks. — This subspecies resembles D. neo-
tridens Hartman in having pseudocompound
hooks with the third tooth slender and in having
comb-setae with numerous fine teeth, but the
pigment pattern is quite different and so is the
nature of the tube.
Records. — Ten small specimens in 40-160 m
off Beaufort (21, *).
Distribution. — West Indies and Brazil in 21-
200 m on muddy sand.
Rhamphobrachium atlanticum, New Species
Figure 8a-h
Holqtype. — USNM 43124; one paratype,
USNM 43125.
Description. — Both type specimens incom-
plete; holotype 55 mm long for 85 segments and
possibly 120 mm long when complete. Tube
constructed of mud and mucus with large shell
fragments plastered over anterior end. Anterior
end of body rounded (Figure 8a) but middle
segments flattened dorsally and 7 mm wide.
Color flesh brown, tentacles speckled, and
head flecked with dark pigment.
Occipital tentacles with short, 5-ringed cera-
tophores and rather long tapered ceratostyles.
Frontal tentacles ovoid and swollen. Tentacular
cirri dorsolateral and well developed. Mandibles
with well-developed cutting edges and pale
straight shafts. Maxillae rather soft and brown;
Mx. I = 1; Mx. II = 8 + 10; Mx. Ill = 9 + 0;
Mx. IV = 8 + 9; Mx.V = 1 + 1.
First three feet obviously capable of great
extension but retracted and wrinkled on holo-
type. Setae retracted but dissection revealed
very long setasacs extending back to segment
45. First three feet with well-developed dorsal
and ventral cirri (Figure 8b), but both cirri
reduced on subsequent segments. First gill as a
single filament on dorsal cirrus of sixth foot
but subsequent gills with a maximum of six
pinnately arranged filaments (Figure 8c). Ven-
tral cirri all cirriform to fifth foot but thereafter
as glandular cushions below setigerous lobes.
Setigerous lobes with a low presetal lip and a
conical postsetal lobe for first three feet. Post-
setal lobe reduced on fourth to tenth foot and
represented by a low postsetal boss on subse-
quent feet.
First three feet with three setal types: (a) six
to eight fine, pointed acicula extending into
base of dorsal cirrus; (b) about four stout,
greatly elongated acicula with curved tips (Fig-
ure 8f); and (c) about four much finer setae with
two rows of spines along the shaft and hooked
tips (Figure 8g). Setae of posterior feet include
three or four pointed acicula and numerous
winged capillaries (Figure 8e). Bidentate acicu-
lar setae (Figure 8h) and fine comb-setae with
15-20 teeth (Figure 8d) from about setiger 25-
30. No spinigerous compound setae present.
Remarks. — The length of the curved acicula
and spiny-shafted hooks of the first three feet
is remarkable. When these feet with their everted
setae are fully extended they must reach far
beyond the head and provide an efficient
means of grasping prey. The hooks in other
species of the genus are sometimes compound
but here they appear plain and the tips lack
sheaths.
Two species of Rhamphobrachium have been
recorded from the United States, namely R.
agassizi Ehlers from Florida to South America
in 770-805 m and R. longosetosum E. and C.
Berkeley from California to the Pacific coast
of Mexico in 18-740 m. R. longosetosum is im-
mediately distinguished from R. atlanticum by
the possession of spinigerous compound setae
in 5th to the 15th foot and branchiae from the
8th to 9th foot. R. agassizi (and R. chuni Ehlers
(1908) from the Indian Ocean) are closer since
they too lack spinigerous compound setae, but
again the gills start farther back. In R. agassizi
the first gill appears on the 11th to 17th foot
V,
Figure 8. — Rhamphobrachium atlanticum n. sp. a, lateral view of anterior end; b, posterior view of 1st foot; c, posterior
view of 40th foot; d, comb-seta; e, winged capillary seta; f, tip of enlarged aciculum of '2d foot; g, pseudocom-
pound seta of 2d foot; h, bidentate acicular seta.
and Ehlers' figures (pi. 17 Fig. 1-5 and pi. 18
Fig. 1-9) show the dorsal cirri of the first three
feet arising halfway along the setigerous lobes;
the maxillae have fewer teeth and the spinules
on the shafts of the pseudocompound hooks
are no longer than the thickness of the shaft
itself, whereas those of R. atlanticum are
double this. In R. chuni the gills start on the
12th foot, the first three feet do not extend
beyond the head and the ventral cirri become
stout glandular swellings on the 4th foot.
Records. — Off Beaufort in 20 and 120 m (*).
Onuphis (Nothria) conchylega Sars, 1835
Onuphis conchylega. - Fauvel, 1923: 415, Fig.
164.
Nothria conchylega. - Hartman, 1944a: 85, Fig.
105-112. -1968: 673, Fig. 1-6.
Onuphis (Nothria) conchylega. - Pettibone,
1963a: 246, Fig. 65a, pi. 17: Fig. 337-338. -
Day, 1967: 425, Fig. 17.13. k-p.
Remarks. — Pettibone (1970b, 251) has re-
defined Nothria Malmgren and has discussed
its controversial status and the characters
which distinguish it from Onuphis Sars, stress-
ing the modification of the first two feet in addi-
tion to the single branchial filaments. She has
assigned two species to Nothria as amended.
Unfortunately the modification of the anterior
feet in Onuphis varies from one species to
another; this applies to their size and the degree
to which they are rotated forwards, the number
of feet with cirriform ventral cirri and the de-
velopment of the hood over the pseudocom-
pound hooks. Similarly, some species which
have a pectinate series of branchial filaments
in the adult have only single filaments in juve-
niles. In view of these variations, I feel that it
is more correct as well as more convenient to
retain Nothria as a subgenus which usefully
56
splits up the many species of the large genus
Onwphis.
Records. — Beaufort on sheltered shores and
dredgings at 200 m (11, *).
Distribution. — Cosmopolitan from 10 m to
abyssal depths.
Onuphis (Nothria) pallidula (Hartman, 1965)
Nothria pallidula Hartman, 1965a: 105, pi. 17:
Fig. d-h.
Records. — Common off Beaufort in 10-200 m
(21, *).
Distribution. — New England to northeastern
South America from 10 to 805 m.
Onuphis eremita
Audouin and Milne-Edwards, 1833
Onuphis eremita. - Fauvel, 1923: 414, Fig. 163. -
Hartman, 1944a: 75; 1968: 691, Fig. 1-5.
Onuphis (Onuphis) eremita. - Pettibone, 1963a:
248, Fig. 65 c. - Day, 1967: 422, Fig. 17.12.
1-q.
Records. — Common off Beaufort in 3-120 m
(21,*).
Dis t rib ution. — Cosmopolitan in temperate and
tropical seas on sandy mud from 0 to 120 m.
Onuphis nebulosa Moore, 1911
Onuphis nebulosa. - Hartman, 1944a: 75, pi. 4:
Fig. 76-85; - 1945: 26; 1968: 699, Fig. 1-6.
Records. — Common in the sounds and off
Beaufort in 20-200 m (11, 21, *).
Distribution. — California to Panama and Gulf
of Mexico and North Carolina from 0 to 200 m.
Onuphis microcephala Hartman, 1944
Onuphis microcephala Hartman, 1944a: 78, pi.
3: Fig. 67-75; pi. 18: Fig. 339.
Records. — Cape Hatteras area and Beaufort,
intertidal (11, 18).
Distribution. — California; Gulf of Mexico;
Guatemala; North Carolina; intertidal.
Onuphis magna (Andrews, 1891)
Diopatra magna Andrews, 1891a: 286, pi. 14:
Fig. 14-20.
Onuphis magna. -Hartman, 1944a: 70; 1945: 26.
Records. — On intertidal and shallow banks
of Beaufort Sound (5, 7, 8, 9, 11, 13).
Distribution. — Pacific coast of Panama, Gulf
of Mexico and West Indies to North Carolina in
0-50 m.
FAMILY LUMBRINERIDAE
Key to genera and species
1 Parapodia with postsetal lobe subdivided to form digitiform
branchiae (Ni)ioe) No N.C. record
1' Parapodia with a single postsetal lobe (Lu mbrineris) 2
2 Prostomium rounded; body usually short (creeping forms).
[Anterior hooks compound] 3
2' Prostomium conical, sometimes elongated; body always long
(burrowing forms) 4
3 Jaws having Mx. Ill with three to four teeth and Mx. IV with
two teeth. Fresh specimens with brown patches on sides of
prostomium L. inflata
3' Jaws having Mx. Ill with two teeth and Mx. IV with one tooth.
Prostomium uniformly pale L. coccinea
57
4 Hooded hooks appear within first five feet 5
4 Hooded hooks appear after 8th foot 12
5 All hooded hooks simple 6
5' Anterior hooded hooks compound, subsequent ones simple 10
6 Prostomium very long, often pointed. First few feet very small 7
6' Prostomium broadly conical. First few feet normally developed.
[Apex of hooded hooks with a crest of several small denti-
cles above a larger tooth] 9
7 Hooded hooks with a crest of small denticles surmounting a
larger tooth. [Mx. II with three teeth. (L. acuta)] No N.C. record
7' Hooded hooks with two well-developed teeth 8
8 Teeth on hooded hooks at right angles. Mx. II with three teeth ;
maxillary supports short and broad L. aberrans
8' Teeth on hooded hooks almost parallel. Mx. II with five or six
teeth; maxillary supports long and slender L. paradoxa
9 Mx. Ill with two teeth. Postsetal lobe of parapodia always
markedly longer than presetal one L. tetraura
9' Mx. Ill with one tooth. Postsetal lobe of middle parapodia
hardly longer than presetal one L. sp.
10 Mx. II with three teeth; Mx. IV as large white plates with a
black margin. [Posterior parapodia with subequal lobes] L. albidentata
10' Mx. II with four or five teeth. Mx. IV completely dark 11
11 Mx. Ill with two adjacent teeth. Postsetal lobe obviously longer
than presetal in middle and posterior segments L. latreilli
11' Mx. Ill as a curved cutting plate with one tooth at the corner.
Postsetal and presetal lobes subequal and both short in
middle segments, but elongate and slender in far posterior
segments L. ciixzensis
12 Acicula black. [Simple hooded hooks appear after 20th foot] 13
12 Acicula pale 14
13 Mx. II with three teeth; Mx. Ill with one tooth. Anterior hooded
hooks with very long blades L. brevipes
13' Mx. II with four or five teeth; Mx. Ill with two teeth. Hooded
hooks always with short blades L. fragilis
14 Anterior setae include compound spinigerous forms; hooded
hooks appear about setiger 18, first few compound, remain-
der simple. Mx. Ill with two indistinct teeth L.januarii
1 i No compound spinigerous setae; hooded hooks always simple
and appear about 12th-17th foot. Mx. Ill with one tooth L. tenuis
58
Lumbrineris inflata Moore, 1911
Lumbrineris inflata Moore, 1911: 289, pi. 19:
Fig. 128-132, pi. 20: Fig. 133, 134. - Hart-
man, 1944a: 160; 1968: 757, Fig. 1-6. - Day,
1967: 435, Fig. 17.16. a-c.
Lumbrineris 'coccinea Pettibone, 1963a: 257,
Fig. 67 d-f (partim).
Remarks. — I do not agree with Dr. Pettibone's
suggestion that L. inflata is synonymous with
L. coccinea. The maxillae of L. inflata are quite
characteristic and L. inflata is restricted to
tropical and subtropical areas.
Records. — North Carolina; common on rocks
and coral from low tide to 18 m (14, 20, 21, *).
Distribution. — Circumtropical on rocky
shores; intertidal to 30 m.
Lumbrineris coccinea (Renier, 1804)
Lumbriconereis coccinea. - Fauvel, 1923: 432,
Fig. 172 g-n.
Lumbrinereis coccinea. - Pettibone, 1963a: 257,
Fig. 67 d-f (partim). - Day, 1967: 436, Fig.
17.16. i-m.
Records. — Fairly common on coral at 7-18 m
off Beaufort (20, *).
Distribution. — Temperate and tropical Atlan-
tic; Mediterranean; Indo-west-Pacific; intertidal
to 30 m on rock.
Lumbrineris aberrans Day, 1963
Lumbrineris aberrans Day, 1963a: 411,Fig.8a-f;
1967: 439, Fig. 17.17. a-c.
Lumbrineris crassicephala Hartman, 1965a: 117,
pi. 20: Fig. c-f.
Ln m brineris -plat ypy gos Fauchald, 1970: 106, pi.
18: Fig. a-d.
Description. — Body threadlike, up to 25 mm
long. Prostomium very long, highly contractile,
usually pointed, occasionally sausage-shaped.
Mandibles delicate and tapered to long slender
shafts in contact throughout. Dental formula:
Mx. I = (2-3) + (2-3), (main fangs indistinctly
bidentate or even tridentate); II = 3 + 3; III =
1 + 1; IV = 1 + 1 (large oval black plates).
Maxillary supports short and broad. First six to
eight parapodia very small or rudimentary, sub-
sequent feet larger, with conical postsetal lobes
obviously longer than the low presetal ones.
One to three broad-winged capillary setae from
first foot to posterior feet. One to two simple
hooks from setiger 4-6 to posterior end; each
hook bidentate with two stout teeth at right
angles to one another. Acicula pale.
Remarks — Fauchald's description of L. platy-
pygos is almost identical to that given above
but the examination of many specimens shows
that the shape of Mx. I is more variable than he
has indicated.
Records.— Off Beaufort in 5-20 m (21, *).
Distribution. — South Africa (26 m); Bermuda
(1,000 m) ; Pacific coast of Mexico.
Lumbrineris paradoxa Saint-Joseph, 1888
Lumbriconereis paradoxa. - Fauvel, 1923: 434,
Fig. 173 a-h.
Lumbrineris paradoxa. - Hartman, 1965a: 119,
pi. 20: Fig. a, b.
Lumbriconereis mucronata Ehlers, 1908: 95, pi.
12: Fig. 9-13.
Remarks. — The three species, L. acuta Verrill,
L. paradoxa Saint-Joseph and L. aberrans Day,
all with characteristically long prostomia, are
easily distinguished by the shape of the hooded
hooks. In L. acuta, as described by Hartman,
(1942a: 114, Fig. 10 d.), the hooks have a crest
of minute denticles above a larger tooth, as is
usual in the genus Lumbrineris. In L. paradoxa
there are two large teeth which are almost
parallel and in L. aberrans there are two very
stout teeth at right angles to one another. I
agree with Hartman that L. mucronata Ehlers
is a synonym of L. paradoxa, and it may also
be noted that the hooded hook figured by Petti-
bone, (1963a, Fig. 67 i) for L. acuta also refers
to L. paradoxa.
Records. — Four specimens off Beaufort in
160-200 m (*).
Distribution. — Azores; Bermuda; off Congo
River mouth and North Carolina in 44-1,700 m.
Lumbrineris tetraura (Schmarda, 1861)
Lu mbri)iconereis impatiens. -Fauvel, 1923: 429,
Fig. 171 a-i.
Lu mbrineris impatiens. - Pettibone, 1963a: 265,
Fig. 67 j.
59
Lu mbriconereis tetraura. - Day, 1953: 435 (with
synonymy).
Lumbrineris tetraura. - Day, 1967: 439, Fig.
17.16. u-w.
Records. — Virginia to Beaufort, intertidal (11,
17).
Distribution. — Cosmopolitan in temperate
and tropical seas; intertidal to 100 m.
Lumbrineris sp.
Description. — Length 10-15 mm. Prostomium
conical. Mandibular shafts in contact through-
out. Mx. I = 1 + 1; Mx. II = 4 + 4 (or 3 + 3 in
small, 10-mm specimens); Mx. Ill = 1 + 1 (cut-
ting plates smaller than Mx. IV); Mx.IV = 1 + 1.
Anterior feet with a low presetal lobe and a
longer postsetal one; middle feet small with
pre- and postsetal lobes subequal; posterior
feet with postsetal lobe definitely longer than
presetal one. Winged capillaries from the first
foot to middle of body. Simple hooks with very
elongate blades resembling broken-tipped cap-
illaries from third or fourth foot to about 20th
and short -bladed hooks thereafter. Acicula pale.
Remarks. — These small specimens do not
quite fit any known species and may be juve-
niles whose characters change later. In L. tenuis
the hooks appear in the 12th-17th foot. In L.
brivicirra Schmarda, recorded from South Africa,
Australia, New Zealand, Chile, and Japan, Mx.
II has five teeth but the other characters are
similar.
Records. — Twenty -three specimens in 40-200
m off Beaufort (21, *).
Lumbrineris albidentata Ehlers, 1908
Lu mbriconereis albidentata Ehlers, 1908: 97,
pi. 13: Fig. 7-13.
Lumbrineris albidentata. - Day, 1960: 357, Fig.
12 a-b; 1967: 434, Fig. 17.15. o-v.
Description. — Body fairly stout, up to 75 mm
long. Prostomium broadly conical. Mandibles
characteristic with broad divergent shafts form-
ing a rough X. Maxillae with Mx. I = 1 + 1;
II = 3 + 3 (large stout teeth); III = 1 + 1; IV =
1 + 1 (very largewhite plates with black margins).
Anterior feet with well-developed presetal lobes
and slightly longer and broader postsetal ones;
middle feet with subequal lobes; posterior feet
with equal tapered lobes curving upward but
not as long as setae. Winged capillaries from
first foot to middle of body. Four to five com-
pound hooks from first foot to 12th -17th and
simple hooks thereafter. Acicula pale.
Records. — Common off Beaufort in 40-200 m
(21, *); This is a new record for the United
States.
Distribution. — South Africa; 30-200 minsandy
mud.
Lumbrineris latreilli
(Audouin and Milne-Edwards, 1833)
Ln mbriconereis latreilli. - Fauvel, 1923: 431,
Fig. 171 m-r.
Lumbrineris latreilli. - Hartman, 1944a: 158,
pi. 9: Fig. 213-216. - Pettibone, 1963a: 258,
Fig. 67 a-c. - Day, 1967: 438, Fig. 17.16. p-t.
Records. — Off Beaufort in 10-40 m (14, 17,
21,*).
Distribution. — Cosmopolitan from low tide to
2,000 m.
Lumbrineris cruzensis Hartman, 1944
Figure 9a-i
Lumbrineris cruzensis Hartman, 1944a: 165, pi.
12: Fig. 263-269; 1968: 751, Fig. 1-6. - Fau-
chald, 1970: 83, pi. 12: Fig. g-j.
Description. — Pale, slender worms 35 mm
long by 0.7 mm for 150 segments. Prostomium
(Figure 9a) bluntly conical. Mandibles (Figure
9c) white, with flaring, well calcified cutting
plates and slender shafts in contact throughout.
Maxillae (Figure 9b) dark brown and well chi-
tinized ; Mx. I = 1 + 1; II = 4 + 4 (in exceptional
cases 3 + 3 or 5 + 5); III = 1 + 1 (curved cutting
plates) ; IV = 1 + 1 (larger plates than Mx. III).
Anterior feet (Figure 9g) well developed, each
with a low presetal lobe and a compressed, oval
postsetal one. Middle feet (Figure 9h) charac-
teristically small with presetal and postsetal
lobes conical and subequal. Far posterior feet
(Figure 9i) with slender elongated lobes slightly
shorter than setae and presetal lobe slightly
(in
longer than postsetal. Winged capillaries (Figure
9f) from first foot to middle of body and one or
two sometimes persisting in posterior feet.
Compound hooks (Figure 9d) from 3d or 4th
foot to 20th and simple hooks (Figure 9e) there-
after. All hooks with an oblique series of den-
ticles above larger first tooth. Acicula pale.
Remarks. — This species is one of the com-
monest worms on the continental shelf of North
Carolina. The specimens are almost always
broken and the characteristic terminal feet with
their subequal filamentous lobes are usually
missing. For this reason and because there are
several species with compound hooks ante-
riorly and closely related to L. cruzensis, the
characters of these North Carolina specimens
have been described in detail. The shape of
Mx. Ill is important but difficult to describe for
the statement that Mx. Ill has one tooth is not
really accurate; actually it has no teeth, only a
cutting edge with an angular corner. L. magal-
haenses Kinberg, L. gracilis Ehlers, and L. limi-
cola Hartman, are all said to have "one tooth"
on Mx. Ill and it is necessary to see the fila-
mentous subequal lobes on the far posterior
feet to distinguish L. cruzensis. The description
of L. cruzensis given by Hartman (1944a) fits
the North Carolina material in all respects ex-
cept that Mx. Ill are smaller than Mx. IV.
Records. — Abundant off Beaufort in 20-200
m (21, *). This is a new record for the Atlantic
coast.
Distribution. — British Columbia to Pacific
coast of Mexico; about 10 m to "slope depths".
"igure 9. — Lumbrineris cruzensis a, anterior end; b, maxillae; c, mandibles; d, compound hook; e, simple hook; f,
winged capillary seta; g\ anterior view of anterior foot; h, anterior view of middle foot; i, anterior view of far
posterior foot.
CI
Lumbrineris brevipes (Mcintosh, 1903)
Ninoe fusca Moore. 1911: 285. pi. 19: Fig. 110-
118.
Lumbrineris brevipes. - Pettibone, 1963a: 260,
Fig. 68 h.
Records.— Off North Carolina (17).
Distribution. — Massachusetts to North Caro-
lina; North West Spain; southern California;
Antarctica; in 100 to 3,000 m.
Lumbrineris fragilis (Muller, 1776)
Lumbriconereis fragilis. - Fauvel, 1923: 430,
Fig. 17 k-1. '
Lumbrineris fragilis. - Pettibone, 1963a: 262,
Fig. 69.
Records. — Common off Beaufort in 80-200 m
(21,*).
Distribution. — A cold-water species extend-
ing from Arctic southward in the Atlantic and
Pacific at deeper and deeper levels and reach-
ing North Carolina, Azores, and Mediterranean;
intertidal to 3,000 m.
Lumbrineris januarii (Grube, 1878)
Lumbrineris januarii. - Hartman, 1944a: 167,
pi. 13: Fig. 278-284.
Description. — Length up to 80 mm. Prosto-
mium conical; maxillae with Mx. I = 1 + 1;
II =5 + 5; III = 2 + 2 (second tooth indistinct);
IV = 1 + 1. Parapodia with postsetal lobe longer
than presetal but shorter than setae even in
posterior feet. Compound spinigerous setae as
well as winged capillaries from first few feet
to about 25th. Compound hooks from about
18th to 30th foot and simple hooks thereafter.
Acicula pale.
Records. — Off Beaufort in 40 m (*). This is
the first record from the United States.
Distribution. — West Indies to Brazil in 10-
40 m.
Lumbrineris tenuis Verrill, 1873
[?] Lumbriconereis atlantica Kinberg, 1869:
568; 1858-1910: 47, pi. 19: Fig. 43. - Hart-
man, 1948: 90, pi. 13: Fig. 1,2.
Lumbrineris tenuis.- Hartman, 1942a: 54.- Petti-
bone, 1963a: 264, Fig. 70 (with synonymy).
Lumbrineris atlantica. - Hartman, 1965a: 116,
pi. 19: Fig. A. * Day, Field, and Montgomery,
1971: 122.
Description. — Body slender, 30-60 mm long.
Prostomium bluntly conical. Mandibular shafts
delicate and in contact throughout. Maxillae
with Mx. I = 1 + 1; II = (4-6) + (4-6); III =
1 + 1 ; IV = 1+1. Mx. II usually with four or five
teeth but occasionally six (Hartman, 1942). Mx.
Ill as a cutting plate smaller than Mx. IV.
Anterior feet with a low presetal lobe and a
larger, compressed postsetal one. Middle feet
rather small with subequal presetal and post-
setal lobes. Posterior feet with longer lobes,
the presetal being obviously shorter than the
postsetal. Winged capillary setae from first foot
to middle feet. Hooded hooks from 12th-17th
foot or exceptionally from 9th (Pettibone, 1963a).
All hooks simple; anterior ones with long blades
but posterior ones with short blades and four
or more denticles above larger first tooth. Acicula
pale.
Remarks. — I am indebted to Dr. Pettibone for
specimens of L. tenuis and Dr. Hartman for
specimens of L. atlantica from 466 to 508 m off
Massachusetts, described in Hartman (1965a).
They appear to be identical .though Dr. Hart-
man's illustration of the maxillae (pi. 19: Fig. a)
represents a form with four teeth on Mx. II
whereas the specimen I dissected had five teeth.
The difficulty is to decide what name should be
used. L. tenuis Verrill has been well described.
L. atlantica Kinberg from La Plata has priority
but Kinberg's description was very brief; he
groups L. atlantica with others having Mx. Ill
unidentate but his figure does not indicate this
and when Hartman (1948) reexamined the type
the jaws had been removed. Again Kinberg did
not figure the far posterior feet. Since there are
several closely related species some doubt must
remain as to whether the specimens described
by Hartman (1965a) off Massachusetts should
really be referred to L. atlantica. In view of this
I have retained the well established name L.
tenuis.
Records. — Common off Beaufort in 20 m
(21,*).
Distribution. — Maine to Florida; intertidal to
abyssal depths.
62
1
1'
2
2'
3
3'
4'
5
5'
FAMILY ARABELLIDAE
Key to genera and species
Parapodia with one or more stout projecting acicula as well
as fine-tipped internal ones 2
Parapodia without a stout projecting aciculum {Arabella) 5
Maxilla I with first tooth greatly enlarged forming a pair of
fangs or pincers (Drilonereis) 3
Maxilla I with first tooth not much larger than others (Noto-
cirrus). [Mandibles present. Mx. I with 3-4 small denticles
at base] N. spiniferus
Mx. I with dentate bases 4
Mx. I with smooth bases. [Mandibles usually absent] D.filum
Parapodia inconspicuous on anterior segments but well de-
veloped and bilobed posteriorly. Mandibles very small, some-
times missing D. longa
Parapodia small on anterior segments, but larger posteriorly,
though always with a single (postsetal) lobe. Mandibles
obvious D. magna
Setae are all winged capillaries Arabella tricolor
Setae include winged capillaries and an inferior, blunt acicu-
lum with a small tapered blade Arabella mutans
Notocirrus spiniferus (Moore, 1906)
Drilonereis longa Webster, 1879
Notocirrus spiniferus. - Pettibone, 1963a: 275,
Fig. 73 a-i.
Records. — Chesapeake Bay, intertidal and
Beaufort in 5 m (17, *)•
Distribution. — Massachusetts to North Caro-
lina; intertidal to 16 m.
Drilonereis Mum (Claparede, 1868)
Drilonereis filum. - Fauvel, 1923: 436, Fig. 174
a-h. - Hartman, 1944a: 180; 1968: 799, Fig.
1,2.
Drilonereis longa. -Hartman, 1944a: 178; -1968:
801, Fig. 1. - Pettibone, 1963a: 272, Fig.
72 a-h.
Records. — North Carolina, intertidal (3, 5, 11,
17, 18).
Distribution. — Massachusetts to Georgia and
the West Indies; Washington; southern Cali-
fornia; intertidal to 2,452 m.
Drilonereis magna Webster and Benedict, 1887
Drilonereis magna. - Pettibone, 1963a: 273,
Fig. 71 h.
Records. — Cape Hatteras area, intertidal (18).
Distribution. — Mediterranean; Atlantic coast
of France; Florida; Panama; southern Cali-
fornia; low tide and shallow dredgings.
Records. — North Carolina, intertidal to 200 m
(11,13,17,18,21,*).
Distribution. — Newfoundland to South Caro-
lina and the Gulf of Mexico ; California to western
Mexico; low tide to 1,000 m.
63
Arabella tricolor (Montagu, 1804)
Arabella mutans (Chamberlin, 1919)
Arabella tricolor. - Fauvel, 1923: 438, Fig. 175
a-h. - Hartman, 1944a: 173; 1968: 789, Fig.
1-6. - Pettibone, 1963a: 269, Fig. 71 a-e. -
Day
1967: 446, Fig. 17.18. i-m.
Records. — North Carolina, intertidal to 80 m
(3.5.7.9, 11, 13. 18,20, *).
Distribution. — Cosmopolitan in temperate and
tropical seas extending from low tide to 83 m.
Cenothrix mutans. - Chamberlin, 1919: 329, pi.
61: Fig. 1-9, pi. 62: Fig. 1.
Arabella mutatis. - Hartman, 1944a: 173. - Day,
1967: 446, Fig. 17.18. f-h.
Records. — Off Beaufort in 5-160 m (*).
Distribution. — Pacific coasts of tropical Amer-
ica; Easter Island; Galapagos Islands; Florida;
Cape Verde Islands; South Africa to tropical
East Africa; intertidal to 160 m.
FAMILY DORVILLEIDAE
Key to genera and species
Dorsal cirri ovoid, without cirrophores or internal acicula and
arise from ends of parapodia 2
Dorsal cirri elongate, mounted on cirrophores with internal
acicula and arise from bases of parapodia (DorviUea) 3
Palps as mere papillae; antennae similar. No forked setae
(Ophryotrocha) O. puerilis
Palps elongate with terminal ovoid palpostyles. Superior forked
setae present (Protodorvillea). [A dorsal cirrus on setiger 1.
Antenna two-jointed] P. kefersteini
No dorsal cirrus on setiger 1 4
A dorsal cirrus on setiger 1. [Superior forked setae present
with slightly unequal prongs] D. rudolphi
No superior forked setae D. sociabilis
Superior forked setae present with very unequal prongs D. caeca
Ophryotrocha puerilis
Claparede and Mecznikow, 1869
Ophryotrocha puerilis. - Fauvel, 1923: 450, Fig.
180 a-h. - Hartman, 1944a: 191, pi. 15: Fig.
325-330; 1968: 823, Fig. 1-6. - Day, 1967:
452, Fig. 17.20. a-f.
Records. — Beaufort, intertidal (11).
Distribution. — Cosmopolitan in warm and
tropical seas occurring in aquaria and on pro-
tected shores.
Protodorvillea kefersteini (Mcintosh, 1869)
Protodorvillea biarticulata. Day, 1963a: 414,
Fig. 8 g-1; 1967: 452, Fig. 17.20. g-1.
Protodorvillea kefersteini. - Hobson, 1971: 542,
Fig. 8 a-d.
Protodorvillea biuret ieulata (sic). - Day, Field,
and Montgomery, 1971: 122.
Description. — Length 5 mm for 50-100 seg-
ments. Prostomium rounded in front with an
annulus in front of antennae. Anterior pair of
eyes minute, often faded, posterior pair larger.
Antennae small, club-shaped, often weakly bi-
articulate. Palps long, wrinkled but terminal
palpostyles distinct and oval. Parapodia long
64
with a retractile presetal lip; an oval dorsal
cirrus at distal end of first and all subsequent
feet; ventral cirri similar to dorsal ones. Superior
setae include one or two long tapered capil-
laries serrated at base of blade and one or two
shorter forked setae with subequal prongs
flanged on inner margins. Inferior setae com-
pound with bidentate falcigerous blades varying
in length.
Remarks. — The only other species of Proto-
dorvillea recorded from the Atlantic coast of
United States is P. minuta Hartman from deep
waters off New England. It is easily distin-
guished by its minute papilliform palps. Hobson
(1971) has redescribed the type of Stauroce-
phalus kefersteini Mcintosh from Scotland and
her account shows that P. biarticulata Day
from South Africa is synonymous. P. gracilis
(Hartman) from California and P. recuperata
Banse and Nichols from Puget Sound are prob-
ably synonyms of P. kefersteini too but Mrs.
Hobson has provisionally retained the name
P. gracilis to cover both.
Records. — Thirteen specimens in 10-120 m
off Beaufort (21, *).
Distribution. — North Atlantic from Scotland
to Ireland and Massachusetts; South Africa;
intertidal to 120 m.
Dorvillea rudolphi. - Hartman, 1945: 27, pi. 5:
Fig. 2, 6. - Day, 1967: 457, Fig. 17.21. d-j.
Stauronereis rudolphi. - Pettibone, 1963a: 231,
Fig. 60.
Records. — North Carolina, intertidal to 120 m
(3,9, 11, 13, 17, 18,20,21, *).
Distribution. — Mediterranean; temperate and
tropical Atlantic from Norway to South Africa
and Massachusetts to the West Indies; Pacific
from British Columbia to Chile; intertidal to
265 m.
Dorvillea socictbilis (Webster, 1879)
Dorvillea sociabilis. - Hartman, 1945: 27, pi. 5:
Fig. 1, 4, 5; 1951: 66, pi. 8: Fig. 3, 5.
Records. — North Carolina, common from low
tide to 160 m (3, 5, 11, 13, 14, 15, 18, 20, *).
Distribution. — Virginia to Florida; inter-
tidal to 160 m.
Dorvillea caeca (Webster and Benedict, 1884)
Stauroneris caecus. - Pettibone, 1963a: 233,
Fig. 61.
Dorvillea rudolphi (Delle Chiaje, 1828)
Staurocephalus rudolphi. - Fauvel, 1923: 446,
Fig. 178 a-p.
Records. — Two specimens off Beaufort on
coral in 10 m (*).
Distribution. — North Japan Sea; Washing-
ton; Gulf of St. Lawrence to Massachusetts;
intertidal to 154 m.
FAMILY SPIONIDAE
Key to genera and species
1 Setiger 5 enlarged and bears stout burrowing hooks laterally 2
1' Neither setiger 4 nor 5 enlarged; no stout burrowing hooks 10
2 Branchiae start on setiger 2 (Boccardia) No N.C. record
2' Branchiae start on setiger 6-11 (Polydora) 3
3 Fifth hooks without accessory teeth but sometimes with a
twisted sheath on concave side of apex 4
3' Fifth hooks with an accessory tooth 7
4 Fifth hooks flanged or with a twisted sheath. [No specialized
posterior notosetae] 5
4' Fifth hooks quite plain. [Hooded hooks from setiger 7] 6
65
5 Fifth hooks with a lateral flange. Hooded hooks from setiger
10-17. Pygidium with several small papillae. [Lives in shells
of hermit crabs] P. commensalis
5' Fifth hooks with a twisted sheath or ridge on concave side
of apex. Hooded hooks from setiger 7. Pygidium saucer-
shaped. [No notosetae on first foot] P. websteri
6 Notosetae present on first foot. Posterior notosetae include
2-3 short spines slightly stouter than the long capillaries P. caeca
6' No notosetae on first foot. No specialized posterior notosetae P. socialis
7 Fifth hooks with a hairy tuft between main and accessory
tooth. [No eyes. Notosetae present on first foot. No special-
ized posterior notosetae] P. hartmanae
7' Fifth hooks without hairs between main and accessory tooth 8
8 Fifth hooks with two teeth partly encircled by a shelf or flange.
Posterior notosetae include large hooked spines P. colonia
8' Fifth hooks normally bidentate without an encircling flange.
No specialized posterior notosetae 9
9 Only four pairs of gills on setigers 7 to 10. Notosetae absent
from first foot. [Body very small] P. tetrabranchia
9' About 14 pairs of gills. No notosetae on first foot. An occipital
tentacle on prostomium. [Bifid, bristled setae accompany
fifth hooks] P. Ugni
10 Branchiae entirely absent. Neurosetae of first foot include a
stout hook-shaped seta. (Spiophanes) 11
10' Branchiae present. No specialized seta on first foot 12
11 Prostomium with pointed lateral projections. No occipital
tentacle. Hooded hooks bidentate throughout. [Notopodial
lobe of setiger 1 well developed] S. bombyx
11 ' Prostomium without lateral projections or occipital tentacle.
Anterior hooded hooks minutely tridentate, posterior ones
bidentate S. wigleyi
12 Branchiae from setiger 1 13
12' Branchiae from setiger 2 20
13 Branchiae present on 20 setigers or more 14
13' Branchiae on setiger 1 or 1-3 only 19
14 Posterior notosetae include hooded hooks. Branchiae absent
from posterior segments. [Prostomium bilobed in front.
(Scolecolepides)] S. viridis
14' Posterior notosetae without hooded hooks. Branchiae continue
to posterior segments 15
66
15 Middle parapodia with small accessory branchiae. Anterior
branchiae fused to notopodial lamellae [Prostomium pointed.
(Dispio)] D. uncinata
15' Middle parapodia without accessory branchiae. Anterior bran-
chiae partly fused or free from notopodial lamellae 16
16 Prostomium rounded or expanded in front (Spio) 17
16' Prostomium with distinct lateral peaks (Malacoceros) No N.C. record
17 Prostomium expanded in front and bears several eyespots.
No occipital tentacle. Hooded hooks in neuropodia from
setiger 28-32 S. multioculata
17' Prostomium rounded in front and bears 4 eyespots. Occipital
tentacle present or absent. Hooded hooks in neuropodia
from setiger 10-11 18
18 No occipital tentacle. Head with brown markings. Hooks bi-
dentate to tridentate, 6-9 per foot S. pettiboneae
18' Occipital tentacle present. Head without brown markings.
Hooks always bidentate, 15-20 per foot S. setosa
19 One pair of smooth branchiae on setiger 1. A dorsal crest
across setiger 2. (Streblospio) S. benedicti
19' Three pairs of pinnate branchiae (or scars thereof) on setigers
1 to 3. A dorsal crest across setiger 1. [Peristome folded
round sides of prostomium (Paraprionospio)] P. pinnata
20 Posterior notopodia with hooded hooks 21
20' Posterior notopodia without hooded hooks 29
21 Branchiae fused to notopodial lamellae anteriorly, becoming
separate later and continuing to posterior segments. Pygid-
ium with a cushion below anus (Scolelepis) S. squamata
21' Branchiae free from notopodial lamellae anteriorly and con-
fined to anterior part of body. Pygidium with anal cirri 22
22 Prostomium with an occipital tentacle. Branchiae never pin-
nate (Aonides) [Hooded hooks bidentate] A. sp.
22' Prostomium without an occipital tentacle. Branchiae either
pinnate, smooth or both (Prionospio) 23
23 All branchiae smooth and number 4 to 40 pairs (subgenus
Minuspio) 24
23' At least one pair of branchiae pinnate and number 4 or 5 pairs 25
24 About 6 pairs of long branchiae. Hooded hooks multidentate.
Genital pouches between middle neuropodia of adults P. (M.) cirrifera
24' About 11 pairs of normal branchiae. Hooks with one tooth
above main fang. No genital pouches P. (M.) cirrobranchiata
25 Five pairs of branchiae with 1st, 4th, and 5th pinnate. [No
dorsal crest across setiger 7] P. heterobranchia
25' Four pairs of branchiae 26
67
26
26'
27
27'
28
28'
Only the 4th pair of branchiae pinnate. Hooded hooks biden-
tate. Prostomium square in front P. dayi
First and 4th pair of branchiae pinnate. Hooded hooks multi-
dentate. Prostomium rounded in front 27
A dorsal crest across setiger 7 or setigers 7 and 9 28
No dorsal crest across setiger 7 or 9 P. steenstrwpi
A dorsal crest across setiger 7 only P.fallax
A dorsal crest across setiger 7, a low ridge across setiger 8,
and a crest across setiger 9 P. cristata
29 Prostomium without an occipital tentacle. [Branchiae free
from notopodial lamellae and continue to posterior seg-
ments (Microspio). Prostomium with a brown patch. Hooded
hooks from setiger 12 with 3-5 denticles above main fang] M. pigmentata
29' Prostium with an occipital tentacle 30
30 Prostomium rounded. Branchiae free from notopodial lamellae
and absent from posterior segments. Membranous genital
pockets between neuropodia of middle segments (Laonice) L. cirrata
30' Prostomium pointed. Branchiae fused to notopodial lamellae
in anterior segments but free later and continue to posterior
segments. No pockets between neuropodia (Nerinides).
[Hooded hooks unidentate] N. unidentata
Polydora commensalis Andrews, 1891
Polydora commensalis Andrews, 1891b: 291, pi.
15: Fig. 27. - Hartman, 1945: 32; 1969: 133,
Fig. 1-4. -Blake, 1969: 815, Fig. 3; 1971: 17,
Fig. 11. - Foster, 1971: 20, Fig. 1-12.
Records. — Cape Hatteras area and Beaufort,
intertidal to a few meters (5, 6, 11, 18, 22)
Distribution. — Massachusetts to North Caro-
lina; Caribbean; Pacific Siberia to California and
western Mexico; intertidal to 30 m; commensal
with hermit crabs.
Polydora websteri Hartman, 1943
Polydora caeca. - Webster, 1879: 252, pi. 9:
Fig. 119-122 (non Oersted).
Polydora websteri Hartman, 1945: 33; 1951: 81;
1969: 151, Fig. 1-5. - Blake, 1969: 814, Fig.
2; 1971: 6, Fig. 3. - Foster, 1971: 26, Fig.
30-36.
Records. — Cape Hatteras to Beaufort, inter-
tidal and dredged (3, 5, 11, 13, 15, 18, 19, ? *).
Distribution. — Newfoundland to Gulf of Mexi-
co ; Oregon to Southern California ; Hawaii ; inter-
tidal to 100 m.
Polydora caeca (Oersted, 1843)
Figure lOa-d
Polydora caeca. - Fauvel, 1927: 52, Fig. 18 a-k. -
Day, 1967: 469, Fig. 18.3. e-h.
Description. — Length up to 20 mm. Prosto-
mium notched in front and produced back as a
ridge to setiger 2. Four eyes. Setiger 1 with
notosetae. Fifth hooks (Figure 10a) plain and
unidentate. Gills from setiger 7-9 to middle of
body at least. Hooded hooks (Figure lOd) from
setiger 7. Specialized posterior notosetae (Fig-
ure 10c) as two to four straight spines only
slightly stouter than accompanying capillaries
(Figure 10b). Pygidium saucer-shaped. Juveniles
with scattered pigment flecks on anterior seg-
ments.
Remarks. — P. caeca is closely related to
P. socialis and the small differences between
158
them have been discussed by Mesnil (1896) and
Hartman (1941a) among others. The most useful
distinction is the presence of notosetae in setiger
1 of P. caeca. In P. socialis the ridge from the
prostomium reaches setiger 4 and the gills are
still present on posterior segments. Further,
P. socialis lacks spines in the posterior noto-
podia but, as these are not distinct in P. caeca,
this is not an obvious character.
Records. — Off Beaufort in 5-80 m (*). This is
a new record for the United States.
Distribution. — Arctic; Atlantic from Green-
land to South Africa; Indian Ocean; Mediter-
ranean; intertidal to 30 m.
Polydora socialis (Schmarda, 1861)
Polydora socialis. - Hartman, 1941a: 310, pi. 48:
Fig. 41, 42; 1945: 33; 1969: 147, Fig. 1, 2. -
Blake, 1969: 816, Fig. 5; 1971: 20, Fig. 13,
14.
Records. — Cape Hatteras area and Beaufort;
intertidal to a few meters (11, 13, 18).
Distribution. — California to Chile; North
Carolina and Gulf of Mexico; intertidal to 10 m.
Polydora hartmanae Blake, 1971
Polydora anoculata. - Hartman, 1945: 33 (non
P. anoculata Moore, 1907).
Polydora hartmanae Blake, 1971: 25, Fig. 16.
Records. — Shallow dredging in Beaufort
Sound (11).
Distribution. — North Carolina; shallow
dredgings.
Polydora colonia Moore, 1907
Polydora tetrabranchia Hartman, 1945
Polydora tetrabranchia Hartman, 1945: 34, pi. 1:
Fig. 7-10. - Blake, 1971: 10, Fig. 7.
Records. — Dredged from Bogue Sound, N.C.
(11).
Distribution. — North Carolina; 3 m.
Polydora ligni Webster, 1879
Polydora ligni. - Hartman, 1941a: 309, pi. 48:
Fig. 47-49; 1945: 32; 1969; 137, Fig. 1-6. -
Blake, 1971: 5, Fig. 1, 2. - Foster, 1971: 22,
Fig. 13-21.
Records. — Cape Hatteras to Beaufort; inter-
tidal to a few meters. (11, 13, 18, *).
Distribution. — Both sides of United States in
temperate seas; Gulf of Mexico; northern Eu-
rope; estuarine, intertidal, and shallow dredg-
ings.
Spiophanes bombyx (Claparede, 1870)
Spiophanes bombyx. - Fauvel, 1927: 41, Fig.
14 a-i. - Hartman, 1951: 85; 1969; 181, Fig.
1-5. - Day, 1967: 474, Fig. 18.5. a-e. - Foster,
1971: 40, Fig. 66-75.
Remarks.— In Day (1967), I described S.
bombyx as having an occipital tentacle; this
is an error since this species lacks an occipital
tentacle.
Records. — Cape Hatteras to Beaufort from
low tidemark on sheltered beaches to dredgings
in 200 m (11, 13, 18,21, *).
Distribution. — Atlantic from Sweden and New
England to the Falkland Islands and South
Africa; eastern Pacific from Canada to Cali-
fornia; intertidal to 200 m.
Polydora colonia Moore, 1907: 199, pi. 15: Fig.
18-23. - Hartman, 1945: 32. - Blake, 1971:
15, Fig. 10.
Polydora h opium inhaca Day, 1957: 99, Fig. 6k-i;
1967: 468, Fig. 18.2. n.
Records. — Beaufort Sound, intertidal (11).
Distribution. — North West Pacific; Massachu-
setts to North Carolina and Jamaica; Mozam-
bique; intertidal.
Spiophanes ivigleyi Pettibone, 1962
Figure 10e, f
Spiophanes wigleyi Pettibone, 1962: 83, Fig. 5,
6. - Hartman, 1965a: 153, pi. 28: Fig. e, f. -
Foster. 1971: 43, Fig. 76-85.
Remarks. — The original description of S. wig-
leyi states that the hooded hooks are bidentate.
My specimens from North Carolina showed a
69
small third tooth above two large ones. Dr.
Pettibone kindly sent me a specimen of S.
ivigleyi from Massachusetts and this resolved
the discrepancy. The first few hooks on setigers
15-18 (Figure lOe) have a vertical series of three
teeth ; in the following segments the small upper-
most tooth becomes minute and in posterior
segments it is absent (Figure lOf). This has also
been noted by Foster (1971).
Records.— Off Beaufort in 40-200 m (21, *).
Distribution. — Massachusetts; Gulf of Mex-
ico; South Africa; 0-200 m.
Scolecolepides viridis (Verrill, 1873)
Scolecolepides viridis. - George, 1966: 76, Fig.
1 a-f. - Foster, 1971: 37, Fig. 57-65.
Records. — Cape Hatteras area, intertidal (18).
Distribution. — Newfoundland to South Caro-
lina; in mudbanks of estuaries to 37 m.
Dispio uncinata Hartman, 1951
Dispio uncinata Hartman, 1951: 87, pi. 22: Fig.
1-5, pi. 23: Fig. 1-4; 1969: 105, Fig. 1-4. -
Foster, 1971: 73, Fig. 161-174.
Records. — Off Beaufort in 3-40 m (22, *).
Distribution. — Massachusetts to the West In-
dies and the Gulf of Mexico; southern Cali-
fornia; intertidal to 40 m.
Spio cf. multioculata (Rioja, 1919)
(?) Spio multioculata. - Fauvel, 1927: 44, Fig.
15 h-o.
Description. — Only two fragmentary speci-
mens, the larger 1.7 mm for 18 segments. No
color markings. Prostomium T-shaped with blunt
anterolateral projections; several eyespots; no
occipital papilla. Gills from setiger 1 to end of
fragment (setiger 18); all gills larger than noto-
podial lamellae and separated from them. Neuro-
podial lamellae broadly oval throughout. Only
capillary setae in notopodia. Neurosetae mainly
capillaries with sabre-setae from 8th foot. No
hooded hooks before end of fragment.
Remarks. — So far as can be seen from these
small fragmentary specimens, the characters
agree perfectly with those of S. multioculata.
The absence of neuropodial hooks is not sur-
prising as they do not occur on S. multioculata
before setiger 28-32. However this species has
not been recorded from the United States and
larger and better preserved specimens are re-
quired to confirm the record.
Records. — Two specimens off Beaufort in
120 m (*).
Distribution. — (of S. multioculata) Bay of
Biscay; intertidal.
Spio pettiboneae Foster, 1970
Figure lla-d
Spio (Spio) pettiboneae Foster, 1971: 35, Fig.
48-56.
Spio filicornis var. nov. Day, Field, and Mont-
gomery, 1971: 122.
Description. — Body up to 11 mm long; head
(Figure 11a, b) and anterior segments flecked
with brown both dorsally and ventrally. Pro-
stomium rounded in front and extending as a
blunt keel to setiger 2. Four subdermal eyes but
no occipital tentacle. Peristome broad and
partly fused to setiger 1. Setiger 1 with small
notopodial and neuropodial lobes and a large
gill. Subsequent parapodia (Figure lie) similar
but larger, each with a well-marked presetal
lobe, a short, blunt postsetal lobe and a large
straplike gill arching over dorsum. Anterior gills
fused basally to postsetal lamellae but pos-
terior ones almost separate. Neuropodia with
small, oval postsetal lobes throughout. Noto-
setae as capillaries with finely punctate blades.
Anterior neurosetae similar but shorter. Hooded
hooks from setiger 11, six or seven per foot.
Anterior hooks bidentate (Figure lid), subse-
quent ones with superior tooth cleft forming
tridentate hooks.
Remarks. — All the specimens from Beaufort
were broken with the largest anterior fragment
having 24 segments. They were originally named
a new variety of S. filicornis and I am indebted
to Dr. Foster for informing me that she had
found the same form in the Gulf of Mexico and
named it S. (S.) pettiboneae. The description
given above is based on the Beaufort material
but it agrees with that of Foster (1971) with
two exceptions. My specimens, which were
presumably younger than those of Dr. Foster,
70
Figure 10. — Polydora caeca a, enlarged hook of setiger 5; b, winged capillary seta; c, posterior notopodial spine;
d, hooded hook. Spiophanes wigleyi e, tridentate hooded hook from setiger 16; f, bidentate hooded hook from a
posterior foot. Prionospio cirrobranchiata g, anterior end; h, 8th foot; j, 40th foot; k, hooded hook. Prionospio
dayi 1, anterior end; m, hooded hook; n, 5th foot with 4th Kill; o, posterior foot. Microspio pig mciitata p, dorsal
view of anterior end; q, ventral view of anterior end; r, anterior view of 8th foot; s, anterior view of posterior
foot; t, hooded hook.
71
were more strongly pigmented and the hooded
hooks appeared effectively bidentate with only
the faintest sign of a third tooth. When fresh,
the pigmentation was striking and character-
istic. Most of the head was brown, there were
always a pair of spots on either side of the mid-
ventral line of anterior segments and often
flecks on the tentacles, the dorsum and the
parapodia.
Records. — Common off Beaufort in 3-120 m
(21.*).
Distribution. — Gulf of Mexico; intertidal.
Spio setosa Verrill, 1873
Spio setosa. - Hartman, 1942a: 63, Fig. 119,
120; 1945: 31, pi. 6: Fig. 1,2.
Records. — Cape Hatteras area and Beaufort
Sound, intertidal (11, 18).
Distribution. — Massachusetts to North Caro-
lina on intertidal sandbanks.
doubts expressed by Sdderstrom (1920). The
possession of a well-developed first setiger,
three pairs of pinnate gills from the first foot
and the large winglike expansions of the peri-
stome which enfold the pointed prostomium
are all distinctive. Dr. Foster has shown that
eight taxa are synonyms of P. pinnata.
Records. — Common off Beaufort in 1-200 m
(21, *).
Distribution. — Cosmopolitan in temperate and
tropical seas from 1 to 500 m.
Scolelepis squamata (Muller, 1806)
Neri)ie cirratulus. - Fauvel, 1927: 36, Fig. 11
g-n. - Hartman, 1969: 115, Fig. 1-5.
Neri)ie agilis. - Hartman, 1945: 31.
Scolelepis (Scolelepis) squaniata. - Pettibone,
1963b: 90 (synonymy). - Foster, 1971: 59,
Fig. 118-131.
Scolelepis squamata. - Day, 1967: 483, Fig.
18.7. c-h.
Streblospio benedicti Webster, 1879
Streblospio benedicti. - Webster, 1886: 149,
pi. 8: Fig. 48-50. - Hartman, 1945: 34, pi. 6:
Fig. 4; 1969: 189, Fig. 1, 2. - Foster, 1971:
112, Fig. 276-283.
Records. — Cape Hatteras area and Beaufort,
intertidal and shallow dredgings in the sound
(11, 15, 18).
Distribution. — Maine to Florida and the Gulf
of Mexico; Denmark and France; Washington
to California; estuarine and intertidal to 10 m
in sandy mud.
Paraprionospio pinnata (Ehlers, 1901)
Prionospio pinnataEhlers, 1901: 163. -Hartman,
1960: 114, pi. 9: Fig. 1-3; 1969: 161, Fig.
1-4. - Day, 1967, 488, Fig. 18.8. i-1.
Prionospio tenuis. - Hartman, 1945: 32 {non
Verrill).
Paraprionspio pinnata. - Foster, 1969: 389, Fig.
12-21 (with synonymy); 1971: 102, Fig. 237-
246.
Remarks. — Dr. Foster has resurrected Caul-
lery's genus Paraprionospio in spite of the
Records. — Cape Hatteras to South Carolina,
common on wave-washed sandy shores (3, 5,
11,13,18,21,*).
Distribution. — Cosmopolitan in temperate and
tropical seas; intertidal in sand.
Aonides sp.
Remark*. — Two small specimens were ob-
tained which were identified by me as juveniles
of Aonides oxycephala (Sars). The specimens
were sent to the U.S. National Museum with the
rest of the collection. Dr. Pettibone has recently
informed me that the larger specimen has now
been identified by Dr. Nancy Foster as Aonides
mayaguezensis Foster (1969: 393, Fig. 22, 33;
1971: 66, Fig. 43-154) originally described from
Puerto Rico in 3 m. A. mayaguezensis is a
small species only 6.6 mm long with fewer
branchiae than A. oxycephala (15-16 pairs in-
stead of 20-30) and neuropodial hooks from
setiger 19-23 instead of setiger 32-35. Possibly
these are juvenile characters.
Records. — Off Beaufort in 20 m (*).
Prionospio (Minuspio) cirrifera Wiren, 1883
Prionospio cirrifera. - Fauvel, 1927: 62, Fig. 21
72
k-n. - Hartman, 1965a: 150; 1969: 155, Fig.
1, 2. - Day, 1967: 486, Fig. 18.8. a-d.
Prionospio delta Hartman, 1965a: 46.
Minuspio cirrifera. - Foster, 1971: 108, Fig.
262-275.
Remarks. —Foster (1971) has erected a new
genus Minuspio, with Prionospio cirrifera as the
type species, which very conveniently includes
all species with four or more pairs of gills none
of which are pinnate. While I do not feel that
this grouping is worthy of generic status it is
most useful as a subgenus.
Records.— Off Beaufort in 10 m (22, *).
Distribution. — Arctic; Atlantic from Sweden
to South Africa and Greenland to South America ;
Bering Sea to southern California; 10-2,500 m.
Prionospio (Minuspio) cirrobranchiata
Day, 1961
Figure lOg-k
Prionospio ? cirrifera. - Hartman, 1951: 84 (non
Wiren).
Prionospio cirrobranchiata Day, 1961: 488, Fig.
4 a-d; 1967: 488, Fig. 18.8. e-h.
Description. — Body up to 20 mm long. Pro-
stomium (Figure lOg) flattened, spade-shaped,
and square in front with four eyes. About 10-12
pairs of smooth cirriform gills starting on
setiger 2; all gills about twice length of noto-
podial lamellae. Anterior notopodial lamellae
(Figure lOh) tapered and pointed; subsequent
ones (Figure lOh) shorter and directed laterally.
Neuropodial lamellae longer than broad even
on posterior feet. No lateral pouches between
neuropodia. Hooded hooks in neuropodia from
setiger 17-19, numbering 4-5 per foot. Individual
hooks (Figure 10k) with a single tooth above
main fang.
Remarks. — Hartman (1951) gives a similar
description of Prionospio ? cirrifera from Florida
but does not mention the structure of the hooks.
Laubier ( 1962) also describes a form from Venice
under the name of P. cirrifera with 10-11 pairs
of smooth cirriform branchiae but again he
does not describe the structure of the hooks.
P. cirrifera Wiren has five or six pairs of smooth
branchiae, genital pouches between the neuro-
podia and hooks with four pairs of teeth above
the main fang.
Records. — Off Beaufort in 80-200 m (21, *).
Distribution. — North Carolina to the Gulf of
Mexico; South Africa; intertidal to 300 m.
Prionospio heterobranchia Moore, 1907
Prionospio heterobranchia Moore, 1907: 195,
pi. 15: Fig. 1-6. - Foster, 1971: 90, Fig. 199-
212.
Prionospio heterobranchia texana. - Hartman,
1951: 85.
Records. — Cape Hatteras area, intertidal (18).
Distribution. — Massachusetts to the West In-
dies and the Gulf of Mexico; intertidal to 10 m.
Prionospio dayi (Foster, 1969)
FiKOire lOl-o
Apoprionospio dayi Foster, 1969: 383, Fig.
1-11; 1971: 97, Fig. 226-236.
Description. — Body up to 30 mm long. Pro-
stomium (Figure 101) square in front with four
eyes. Setiger 1 reduced and fused to peristome.
Four pairs of branchiae on setigers 2 to 5; first
pair small, smooth and cirriform; second and
third pairs broad, smooth and compressed;
fourth pair (Figure lOn) largest with a double
series of papillae on inner margin. Notopodial
lamellae pointed and medial to notosetae; fourth
lamella largest, subsequent ones low and oval.
A membranous crest across setiger 7. Neuro-
podial lamellae prominent and rounded, that of
setiger 2 particularly enlarged. Posterior feet
(Figure lOo) with similar postsetal lamellae in
both rami. No genital pockets between neuro-
podia. Hooded hooks in neuropodia from setiger
16-18 with 8-10 per foot. Notopodial hooded
hooks from about setiger 40. Individual hooks
(Figure 10m) with three pairs of denticles above
main fang. Pygidium with three anal cirri, median
one long and lateral ones short.
Remarks. — Foster (1969: 388) has discussed
the affinities of this species and has given a
key which neatly separates P. dayi from P.
pygmaea Hartman, P. saldanha Day, and P.
caspersi Laubier.
P. dayi and P. pygmaea both have multi-
dentate hooded hooks while P. saldanha and
P. caspersi have a single tooth above the main
73
fang. Further P. dayi has a membranous crest
across setiger 7 while P. pygmaea has not.
In a partial revision of the genus Prionospio
to which numerous taxa have been assigned.
Dr. Foster has erected a new genus Apopriouo-
spio to include those species with four pairs
of gills starting on setiger 2 and with only the
fourth pair pinnate. While it would be most
helpful to divide the large genus Prionospio,
I believe that such a narrow division as that
suggested would demand the erection of too
many genera or subgenera. P. ehlersi Fauvel
has four pairs of gills with only the first pair
pinnate; P. steenstrupi Malmgren (the type
species of the genus Prionospio), P. fallax
Soderstrom, and P. bocki Soderstrom have four
pairs of gills of which the first and the fourth
are pinnate; P. plumosa Sars has four pairs
with all except the third pair pinnate. P. dayi
and the three species allied to it have four
pairs with only the fourth pinnate; P. hetero-
branchia Moore has five pairs with the first,
fourth, and fifth pinnate. Beyond this there are
many species with more pairs of gills. If a
subgenus were proposed which included the
type species P. steenstrupi and all others with
four pairs of gills starting on setiger 2 and one
or more of the gills pinnate it would include
many closely related species but it is felt that
the genus Apoprionospio as presently defined
creates too many difficulties.
Records. — Common off Beaufort in 3-200 m
(21,22, *).
Distribution. — North Carolina and the Gulf of
Mexico; intertidal to 200 m.
Prionospio fallax Soderstrom, 1920
(?) Prionospio malmgreni Claparede, 1869: 73,
pi. 22: Fig. 3.
Prionospio fallax Soderstrom, 1920: 235, Fig.
135, 144, 145.
Prionospio malmgreni. - Fauvel, 1927: 61, Fig.
21 a-e. - Day, 1963a: 418; 1967: 492, Fig.
18.9. a-c. - Hartman, 1969: 159, Fig. 1-4. -
Day, Field, and Montgomery, 1971: 122.
Remarks. — Foster (1971: 82) has drawn at-
tention to the fact that Claparede's original de-
scription of P. malmgreni is so vague and con-
tradictory that it is impossible to be certain of
the gill arrangement. Thus the species is in-
determinate. Nonetheless, the description of
P. malmgreni given by Fauvel (1927) and later
workers agrees perfectly with the original de-
scription of P. fallax Soderstrom. Soderstrom
also suggested that the two species are identical.
Records. — Off Beaufort in 10-200 m (21, *).
Distribution. — Atlantic from Sweden to Ma-
deira; Mediterranean; South Africa; southern
California; 10-200 m.
Prionospio cristata Foster, 1971
Prionospio cristata Foster, 1971: 87, Fig. 186-
199.
Records. — Beaufort in sand at 0.5-32 m (22).
Distribution. — North Carolina, Gulf of Mexi-
co, and the West Indies; 0.5-32 m.
Prionospio steenstrupi Malmgren, 1867
Prionospio steenstrupi. - Fauvel, 1927: 60, Fig.
21 f-i. - Hartman, 1965a: 152; 1969: 165,
Fig. 1, 2. - Day, 1967: 489, Fig. 18.9. o-r. -
Foster, 1971: 84, Fig. 175-185.
Records. — Common off Beaufort in 160-200 m
(21,22, *).
Distribution. — North Atlantic from Norway
to Greenland and New Brunswick to Florida;
Alaska to southern California; Japan; South
Africa; intertidal to 1,745 m.
Microspio pigmentata (Reish, 1959)
Figure lOp-t
Spiophanes pigmentata Reish, 1959: 11, pi. 6:
Fig. 1-4.
Nerinides pigmentata. - Hartman, 1961: 92.
Spio (Microspio) pigmentata. - Foster, 1971: 35
(list only).
Description. — Body about 10 mm long. Pig-
mentation characteristic, including a square
brown patch on prostomium, lateral and ventral
marks on peristome, and midventral spots and
fainter marks on parapodia of anterior seg-
ments (Figure lOp, q). Prostomium rounded
anteriorly and extending back as a low ridge
to setiger 2. Four eyes but no occipital tentacle.
Setiger 1 small, partly fused to peristome, with-
out branchiae but with distinct parapodial lobes
71
Figure 11. — Spio pettiboneae a and b, dorsal and ventral views of anterior end; c, anterior view of anterior foot;
d, hooded hook. Nerinides unidentata n. sp. e, anterior end; f, anterior view of 6th foot; g\ anterior
view of 18th foot; h, inferior sabre seta; j, hooded hook. Magelona papillicornis k, head; 1, anterior view of 6th
foot; m, anterior view of abdominal foot; n, special seta from setiger 9; o and p, face view and profile of abdominal
hook. Magelona physillae q, head; r, anterior view of 6th foot; s, anterior view of abdominal foot; t, setae from
setiger 9; u and v, face view and profile of abdominal hook. Magelona sp. w, head; x, anterior view of 5th foot;
y, anterior view of abdominal foot; z, face view of abdominal hook.
75
bearing both notosetae and neurosetae. Bran-
chiae from setiger 2 to posterior segments,
each gill straplike and separate from notopodial
lobes. Notopodia of anterior feet (Figure lOr)
with distinct presetal lamellae, and larger
rounded postsetal lamellae; neuropodia similar
but smaller. All lamellae of posterior feet (Fig-
ure 10s) smaller, the postsetal lamellae of neuro-
podia being mere ridges. No sign of genital
pockets between neuropodia of middle segments.
Pygidium with four anal cirri. Notosetae as
narrow-bladed capillaries throughout, there
being no notopodial hooks. Neurosetae as capil-
laries in anterior feet but capillaries mainly
replaced by 8-12 hooded hooks from 12th foot.
Individual hooks (Figure lOt) with a series of
four denticles above main fang.
Remarks. — The genus Microspio Mesnil with
its type species Spio mecznikoivianus Claparede,
has had a checkered history and even now its
status is controversial. Discussions will be found
in Soderstrom (1920), Fauvel (1927), Holmquist
(1967), and Foster (1971). Microspio has bran-
chiae from setiger 2, while Spio has branchiae
from setiger 1. According to Soderstrom, Micro-
spio may also be distinguished from Spio by the
possession of only two dorsal ciliated organs
per segment instead of four, but this distinction
demands staining and sectioning. Unfortunately
Soderstrom confused these clear distinctions
by including other species in the genus Micro-
spio which have gills from setiger 1. Holmquist
has shown that these should be transferred to
other genera. Foster gives a useful list of the
species which may be included in Microspio
which she regards as a subgenus of Spio. Hart-
man (1959a) listed Microspio as a synonym of
Paraspio Czerniavsky but the original diagnosis
states that Paraspio has branchiae on all setig-
erous segments and, as Holmquist states, it
is a synonym of Spio.
Spiophanes pigmentata Reish was trans-
ferred to the genus Nerinides by Hartman ( 1961)
but the latter genus has the anterior branchiae
fused to the notopodial lamellae and the pro-
stomium has a well-developed occipital tentacle.
Hartman (1969: 91) also described another
species, Nerinides maculata from southern Cali-
fornia, which Foster (1971) refers to as Spio
(Microspio) mac/data. It is closely allied to
Spiophanes pigmentata but it lacks the brown
patch on the prostomium, the hooks appear in
the neuropodium of setiger 11 not 12, and they
have two small teeth in tandem above the main
fang instead of three to five.
Records.— Off Beaufort in 40-80 m (*).
Distribution. — Southern California "in shelf
depths"
Laonice cirrata (Sars, 1851)
Laonice cirrata. - Fauvel, 1927: 38, Fig. 12 a-e. -
Day, 1967: 480, Fig. 18.6. h-k. - Hartman,
1969: 107, Fig. 1-4. - Foster, 1971:
69, Fig. 155-160.
Records.— Off Beaufort in 80-120 m (*).
Distribution. — Cosmopolitan from the Arctic
to the Antarctic from 30 m to abyssal depths.
Nerinides unidentata New Species
KiniJre lle-j
Holotype.— USNM 43150.
Description. — Holotype incomplete with only
24 segments measuring 6 mm. No color mark-
ings. Prostomium (Figure lie) sharply pointed
anteriorly and produced as a blunt keel to seti-
ger 1. Two pairs of eyes and a large, erect
occipital tentacle. Setiger 1 well developed with
both notosetae and neurosetae. Dorsum flat-
tened and without membranous crests. Gills
from setiger 2 to end of fragment (setiger 24).
Anterior gills (Figure llf) completely fused to
notopodial lamellae, but 8th and succeeding
feet with shorter postsetal lamellae and gills
free and well tapered (Figure llg). Neuropodial
lamellae prominent, oval, never bilobed. No
genital pockets. Notosetae as limbate capillaries,
often with punctate shafts. Anterior neurosetae
similar. Inferior sabre-setae with punctate
blades (Figure llh) from setiger 12. Hooded
hooks from setiger 20, each with a blunt uni-
dentate tip (Figure llj).
Remarks. — This species agrees with N. knight-
jonesi de Silva from Ceylon in having uniden-
tate hooded hooks but differs in the shape of
the prostomium, in having the branchiae com-
pletely fused to the dorsal lamella on the first
few feet and in having hooded hooks in the
neuropodia of setiger 20, not setiger 43.
Records. — One specimen off Beaufort in 10 m
(*)•
76
FAMILY MAGELONIDAE
Key to species of Magelona
1 Setiger 9 with specialized setae having a subterminal expan-
sion [Prostomium spatulate and smoothly curved anteriorly.
Hooded hooks with a pair of denticles above main fang] M. papillicornis
V Setiger 9 without specialized setae 2
2 Prostomium with anterolateral angles or "horns" 3
2' Prostomium smoothly curved in front. [Hooded hooks with
a pair of denticles above main fang. Parapodia of anterior
region with a small lobe medial to notosetae] M. rosea
3 Hooded hooks with a single denticle above main fang. Noto-
podia of anterior region without a small lobe medial to
notosetae M. phyllisae
3' Hooded hooks with a pair of denticles above main fang. Noto-
podia of anterior region with a small lobe medial to noto-
setae M. sp.
Magelona papillicornis Muller, 1858
Figure llk-p
Magelona papillicornis. - Fauvel, 1927: 64,
Fig. 22 a-h. - Day, 1967: 495, Fig. 19.1.
a-d. -Jones, 1963: 23 (key only).
Magelona rosea. - Wells and Gray, 1964: 73
(non M. rosea Moore, 1907).
Magelona sp. -Jones, 1968: 272, Fig. 1-33.
[?:
[?
Descriptlo)i. — Body white, seldom more than
50 mm long in North Carolina, up to 170 mm
in Europe. Prostomium (Figure Ilk) spatulate
with anterior margin smoothly rounded. Palps
with four rows of adhesive papillae. Body clearly
divided; anterior region with nine setigers and
posterior region with numerous setigers. Noto-
podia of setigers 1-8 (Figure 111) with a small
papilla medial to notosetae and a tongue-shaped
postsetal lamella; neuropodia with a similar
but smaller postsetal lamella. Setiger 9 without
medial papillae but with small presetal as well
as postsetal lamellae in both rami. Abdominal
parapodia (Figure 11m) with medial papillae
both dorsal ly and ventral ly and equal, oval
notopodial and neuropodial lamellae curving
towards one another. Lateral membranous
pockets between one parapodium and the next.
Setigers 1-8 with long bilimbate capillaries;
setiger 9 with fans of specialized setae bearing
spatulate blades ending in mucronated tips
(Figure lln). Abdominal setae as short hooded
hooks, bidentate in profile but actually with a
pair of denticles above the main fang. (Figure
llo,p).
Remarks. — Dr. Jones informs me that M.
papillicornis is probably the same as Magelona
sp. Jones (1968) from Woods Hole; M. riojai
Jones is very close but, according to Jones,
there are differences in the shape of the pro-
stomium and in the setae of the 9th foot.
Records. — Pamlico Sound, intertidal and
abundant off Beaufort in 3-10 m ( 19, 21, *).
Distribution. — Atlantic from Scotland to
South Africa and Brazil; ? Massachusetts;
North Carolina; Mediterranean; Madagascar;
intertidal to 100 m.
Magelona phyllisae Jones, 1963
Figure llq-v
Magelona piiyllisae Jones, 1963: 2, Fig. 1-11.
Description. — Body colorless, up to 20 mm
long. Prostomium (Figure llq) flattened, not
much broader than body and anterolateral
angles or "horns" well marked. Palps long,
77
with four irregular rows of papillae. Parapodia
of setigers 1-8 (Figure llr) without papillae
medial to notosetae but with elongated post-
setal lamellae in notopodia and shorter but
similar lamellae in neuropodia Setiger 9 with
subequal lamellae, bearing bilimbate setae simi-
lar to those of setigers 1-8 (Figure lit). Ab-
domen with postsetal lamellae of both rami
rather small and expanded distal ly (Figure
lis). Relatively large papillae medial to hook-
rows both dorsally and ventrally. Individual
hooded hooks with only one large denticle above
main fang (Figure llu, v).
Remarks. — As noted by Jones (1963: 25)
this species is close to M. longicornis Johnson.
It may be distinguished by the absence of medial
papillae above the thoracic notosetae and the
possession of larger medial papillae on abdomi-
nal segments. Dr. Jones informs me that these
specimens from Beaufort appear to be the same
as specimens from Sapelo Island, Ga., and
Port Aransas, Tex.
Records. — Off Beaufort in 10 m (*). This is a
new record for the United States.
Distribution. — Peru in 181 m; ? Georgia and
Texas.
Magelona sp.
Figure llw-z
15 mm. They were obviously different from the
other species recorded here and for this reason
the characters are summarized below.
Description. — No color markings. Prosto-
mium (Figure llw) almost as broad as long
with distinct anterolateral angles. Anterior
parapodia (Figure llx) with medial papillae
above notosetae and postsetal lamellae of both
rami flattened and ligulate. Setiger 9 with
postsetal lamellae only, and with bilimbate
capillaries similar to those of setigers 1-8.
Abdominal parapodia (Figure lly) with broad
postsetal lamellae in both rami and papillae
medial to rows of hooks both dorsally and
ventrally. Individual hooks (Figure llz) with
two denticles side by side above main fang.
Remarks. — This species is allied to both M.
filiformis Wilson and M. cornuta Wesenberg-
Lund. A specimen of M. filiformis, which Dr.
Wilson kindly sent to me, had much narrower
parapodial lamellae on the thorax. Possibly
these North Carolina specimens are closer to
M. co run to but more material is required to
confirm the record.
Records.— Off Beaufort in 80-120 m (*).
Magelona rosea Moore, 1907
Magelona rosea Moore, 1907: 201, pi. 16: Fig.
24-30. -Jones, 1963: 23 (key only).
Material examined. — Three anterior frag-
ments were collected of which the longest was
Records. — Off Beaufort in 80 m (*).
Distribution. — Massachusetts; intertidal.
FAMILY POECILOCHAETIDAE
Poeciloehaetus sp.
Remarks. — Only two anterior fragments were
obtained. They were obviously juveniles, the
larger measuring 4 mm for 22 segments. Final
identification is impossible, since the posterior
segments with their specialized setae are miss-
ing. However, the other characters agree with
P. serpens. The dorsum is smooth, not papillose,
though one specimen has a chitinized projection
on segment 9, rather like that described by
Hartman (1939) for P. johnsoni. The nuchal
organ has three free lobes with fuzzy edges,
and it is difficult to be certain whether the ends
are broken. The middle lobe reaches the pos-
terior margin of setiger 2 and the lateral lobes
are only slightly longer than broad. As usual
in the genus there are curved spines in the
neuropodia of setigers 2 and 3 and long, flask-
shaped parapodial lobes with knobbed ends on
setigers 7 to 13. Plumose setae appear among
the capillaries on setiger 20. The smooth dor-
sum excludes P. fulgoris and the three-lobed
nuchal organ suggests P. serpens.
Records.— Off Beaufort in 80 m (*).
78
FAMILY CHAETOPTERIDAE
Key to genera and species
1 Notopodia of middle region bilobed or trilobed. Tube often
horny and ringed 2
1' Notopodia of middle region never bilobed. Tube neither horny
nor ringed 3
2 A pair of minute tentacular cirri as well as large grooved
palps (Phyllochaetopterus). Middle region of 7 or more
segments. Tube horny, often branching P. socialis
2' No tentacular cirri, only a pair of grooved palps. Tube horny,
ringed, solitary (Spiochaetopterus). Middle region with
about 20 segments. [Eyes present] S. costarum oculatus
3 Middle region of five segments, the last three with notopodia
fused to form large paddles. Palps much shorter than
anterior region (Chaetopterus) C. variopedatus
3' Middle region of less than five segments, bearing separate,
fingerlike notopodia, very like those of posterior region.
Palps long (Mesochaetopterus). [Middle region of three long
segments] M. taylori
Phyllochaetopterus socialis Claparede, 1870
Phyllochaetopterus socialis. - Fauvel, 1927: 84,
Fig. 30 a-1. - Day, 1967: 525, Fig. 22.1. h-r.
Records. — Off Beaufort in 20 m (*).
Distribution. — Cosmopolitan in temperate
and tropical seas; intertidal to 100 m.
Spiochaetopterus costarum oculatus
Webster, 1879
Spiochaetopterus oculatus Webster, 1879: 47,
pi. 8: Fig. 98-102. - Hartman, 1945: 35. -
Barnes, 1964: 397, Fig. 1-4.
Spiochaetopterus costarum oculatus. - Gitay,
1969: 15.
Description. — Body slender, up to 60 mm
long; dark ventral patch from setiger 6 to 7,
white patch from setiger 7 to 9. Prostomium
oval, eyes dark, conspicuous. Buccal segment
large, fleshy, collarlike; palps long, colorless.
No tentacular cirri. Anterior region of nine
uniramous flattened setigers. A single stout
brown cutting seta in fourth foot. Middle region
of about 20 segments each with simple club-
shaped notopodia. Tube long, slender, translu-
cent, annulated. Animal solitary, living in
sandy mud.
Remarks. — All the specimens dredged off
Beaufort were juveniles, with very delicate
transparent tubes which lacked annuli, so that
it would appear that these develop with age, as
the tube thickens. Juveniles as small as 5 mm
already had the characteristic dark patch on
the ventrum of setiger 6. All specimens except
one had conspicuous eyes. The smallest speci-
mens only had three or four segments in the
middle region, but the number increases rapidly
with the length of the worm.
According to Barnes, the main difference
between S. costarum costarum and S. costarum
oculatus is that S. costarum costarum uses
only one mucus bag to collect food particles,
while S. costarum oculatus uses eight or more.
Records. — Cape Hatteras area to Beaufort
on sheltered banks and below low tide (3, 11, 13,
18, *).
Distribution. — Massachusetts to the Gulf of
Mexico; intertidal and shallow dredgings.
79
Chaetopterus variopedatus (Renier. 1804)
Chaetopterus variopedatus. - Fauvel, 1927: 77,
Fig. 26 a-n. - Day, 1967: 529, Fig. 22.2. a-g.
- Hartman. 1969: 209, Fig. 1-3.
Rt cord*. — Cape Hatteras to Beaufort, inter-
tidal to 30 m (4, 5, 7. 8, 9, 11, 13. 18).
Distribution. — Cosmopolitan in temperate
and tropical seas; intertidal to 100 m.
Mesochaetopterus taylori Potts, 1914
Mesochaetopterus taylori Potts, 1914: 958, pi.
1: Fig. 1-3, pi. 3: Fig. 5. 6. 9, text Fig. 1-5. -
Hartman, 1969: 213, Fig. 1-4.
Description. — Body up to 100 mm long. Pro-
stomium small, oval, without eyes in adult.
Buccal segment large, swollen, and collarlike
with a pair of long grooved palps. No tentacular
cirrus. Anterior region of nine uniramous seg-
ments. Several stout brown cutting setae in
fourth foot. Middle region of three long seg-
ments with simple notopodial lobes. Cup-shaped
organs on second and third segment of mid-
region. Posterior region not clearly distinguished
from middle region; notopodia similar, but seg-
ments progressively shorter and notopodia more
conical. Tube fragile and usually covered with
sand.
Records. — Cape Hatteras area and Beaufort
on intertidal mudbanks (18, *).
Distribution. — Western Canada to northern
California and North Carolina; intertidal.
FAMILY CIRRATULIDAE
Key to genera and species
1 Several grooved tentacular filaments (or scars showing their
origin) above first few setigers 2
1' One pair of grooved tentacular filaments or palps at junction
of setiger 1 and last annulus of peristome 4
2 Tentacular filaments and gills arise on same segment (Cirra-
tulus) No N.C. record
2' Tentacular filaments arise posterior to first gill filament which
appears on setiger 1. [Sigmoid acicular hooks present
(Cirriformia)] 3
3 Tentacular filaments in a row between setigers 1 and 2. Gill
filaments of middle segments arise immediately above noto-
setae C. graudis
3' Tentacular filaments arise above setiger 4. Gill filaments of
middle segments arise farther above notosetae than distance
between notosetae and neurosetae C.filigera
4 Never more than 10 pairs of gill filaments. Acicular setae
with spoon-shaped ends. (Dodecaceria). [Body dark; 9-10
pairs of gills] D. corollii
4' Many pairs of gills. Acicular setae when present, lack spoon-
shaped ends 5
5 No acicular setae even in posterior segments, only tapered
capillaries ( Tharyx) 6
5' Acicular setae present as well as capillaries, though former
sometimes restricted to posterior segments 8
80
6 Capillary setae of long posterior segments are short with saw-
edged blades T. annulosus
6' Capillary setae with smooth or finely spinulose blades through-
out 7
7 Prostomium with eyespots. Only a few capillary setae per
parapodium T. setigera
T Prostomium without eyespots in adult. Numerous capillary
setae [possibly adult of T. setigera] T. marioni
8 Acicular setae of terminal segments very prominent and
numerous, almost encircling body (Chaetozone) 9
8' Acicular setae of terminal segments not obviously different
from preceding ones (Caulleriella). [Acicular setae short,
with faintly bidentate ends without hoods] C. killariensis
9 Acicular spines present in neuropodia from first setiger on-
wards Ch. gayheadia
9' Acicular spines appear in neuropodia of middle segments Ch. setosa
Cirriformia grand is
(Verrill, 1873) new combination
Cirratulus grandis Verrill, 1873a: 606, pi. 15:
Fig. 80, 81. - Hartman, 1942b: 126.
Description. — Body yellowish green, up to
150 mm long. Segments short, rounded dor-
sally, and flattened ventrally. Prostomium
bluntly conical, without eyes but with diffuse
dark pigment. Peristome fused to prostomium,
rather long and irregularly wrinkled but not
segmented. A transverse row of about 20 grooved
tentacular filaments above junction of setiger
1 to 2. A median gap in tentacular row
separating right and left groups. Cylindrical
(not grooved) gill filaments from setiger 1 (thus
anterior to tentacular filaments) to posterior
part of body. Gill filaments, even of middle seg-
ments, arise immediately above notosetae. Cap-
illary setae in both rami of all segments.
Yellow acicular spines quite distinct in both
rami of posterior segments but longer and more
like capillaries in anterior feet; spines distinct
in juveniles from notopodium of setiger 35 and
neuropodium of setiger 19 but not distinct in
adults in anterior third of body.
Remarks. — Verrill's description is not clear
regarding the origin of the first gill filaments and
the tentacular filaments. As noted by Day (1967:
500) the tentacular filaments arise from the
posterior dorsal margin of the peristome. In
Cirriformia this peristomial projection extends
further back than the first setiger which bears
the first pair of gill filaments; in Cirratulus the
first pair of gill filaments arises at the same
level as the tentacular filaments. In Cirriformia
grandis the tentacular filaments arise just pos-
terior to the gill filaments on setiger 1 so that
this species is a link between the two genera.
In Cirratulus cirratus (Muller) the tentacular
filaments arise above the gill filaments on setiger
1. Thus Cirriformia grandis and Cirratulus
cirratus are very alike in this respect, but
C. cirratus can easily be distinguished by the
fact that in the middle of the body the gill
filaments arise at least as far above the noto-
setae as the distance between the notosetae
and neurosetae.
Records. — Off Beaufort in 10-80 m (3, *).
Distribution. — Massachusetts to North Caro-
lina; intertidal to 80 m.
Cirriformia ftligera (Delle Chiaje, 1828)
Audouiuia filigera. - Fauvel, 1927: 92, Fig.
32 h-m.
Cirriformia filigera. - Hartman, 1951: 94. -
Day, 1967: 518, Fig. 20.4. p-q.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 50 m (11, 13, 18, 19, 20).
81
Distribution. — Cosmopolitan in warm and
tropical seas; intertidal to 50 m.
Dodecaceria corallii (Leidy, 1855)
Figure 12a, b
Naraganseta corallii Leidy, 1855: 494. - Miner,
1950: 344.
Dodecaceria near concharum. - Hartman, 1951:
94.
Description. — Body dark green to black, 8-12
mm long, rounded anteriorly, rather flattened
posteriorly. Burrows common in encrusting
corallines or corals, e.g., Astrangia danae. Pro-
stomium (Figure 12a) as an oval hood over-
hanging mouth; eyes indistinguishable. Peri-
stome long, annulated, fused to prostomium
anteriorly, and bearing a pair of cylindrical
gills and a pair of grooved palps posteriorly
at junction with setiger 1. Similar gill filaments
of decreasing length present above notosetae
of setigers 1-6 or 1-9; thus 7-10 pairs in all.
Segments short and without parapodial promi-
nences, the setae arising directly from body wall.
Anterior segments bearing only limbate capil-
laries with minutely serrated margins. First
hooks in neuropodium of setiger 8-12 and in
notopodium of setiger 10-13; middle segments
with hooks but few capillaries; posterior seg-
ments with hooks and more capillaries. Indi-
vidual hooks (Figure 12b) with slightly sigmoid
shafts and excavated ends preceded by a boss
thus resembling spoons with a lump before the
bowl.
Remarks. — D. corallii differs from D. con-
charum Oersted in having more pairs of gills
and more pointed bowls to the spoon-shaped
hooks.
Records. — Common in corals in 6.5-20 m off
Beaufort (20, *).
Distribution. — Massachusetts to the Gulf of
Mexico in 0-20 m.
Tharyx annulosus Hartman, 1965
Tharyx annulosus Hartman, 1965a: 167, pi. 34:
Fig. a-e.
[?] Caulleriella annulosa. - Banse and Hobson,
1968: 31, Fig. 7 a.
Re murks. — The specimens from North Caro-
lina agree very well with Dr. Hartman's descrip-
tion. The acicular spines reported by Banse and
Hobson (1968) were not seen and possibly belong
to a different species with many similar char-
acters.
Records. — Off Beaufort in 80-200 m (21, *).
Distribution. — New England to tropical South
America; South Africa; 80-4,540 m.
Tharyx setigera Hartman, 1945
Tharyx setigera Hartman, 1945: 35, pi. 7: Fig.
1-3.
Note. — T. setigera differs from T. marioni in
possessing eyespots and in having fewer setae;
these may be juvenile characters.
Records. — Cape Hatteras area to Beaufort,
intertidal (11, 15, 18).
Distribution. — North Carolina; intertidal.
Tharyx marioni (Saint-Joseph, 1894)
Tharyx marioni. - Fauvel, 1927: 100, Fig. 35
a-b. - Hartman, 1965a: 169. - Day, 1967:
505, Fig. 20.2. a-c.
Remarks. — Anterior fragments of T. marioni
are difficult to distinguish from those of Chaeto-
zone setosa since the acicular setae of the latter
are not well differentiated from capillaries in
anterior segments.
Records.— Off Beaufort in 18-80 m (20, *).
Distribution. — Temperate North and South
Atlantic from the English Channel, eastern
Canada to North Carolina, northern South
America, South Africa; intertidal to 1,000 m.
Caulleriella killariensis (Southern, 1914)
Heterocirrus killariensis. - Fauvel, 1927: 97,
Fig. 34 d-h.
Description. — Body threadlike, 8-12 mm long.
Prostomium conical and acutely pointed but
without eyes. Peristome faintly annulated, with
a pair of long grooved palps at junction with
setiger 1. Filiform branchiae from setiger 1 to
near end of body; anterior branchial filaments
long, arising immediately above notosetae; pos-
terior filaments short, arising well above noto-
82
setae. Few setae per parapodium; four to six
short capillaries per ramus up to setiger 7 and
one or two fine capillaries plus one or two short
acicular hooks per ramus in subsequent seg-
ments. Individual hooks minutely bidentate and
without a hood or sheath.
Records. — One specimen from 10 m off Beau-
fort (*). This is a new record for the United
States.
Distribution. — Ireland; in 10-20 m.
Chaetozone gayheaciia Hartman, 1965
Chaetozone gayheadia Hartman, 1965a: 166.
Chaetozone setosa. - Day, 1967: 510, Fig. 20.1.
1-p (non Malmgren).
flfr<»-d.s.— Off Beaufort in 40-160 m (*).
Distribution. — New England to North Caro-
lina in 40-300 m; South Africa (95 m).
Chaetozone setosa Malmgren, 1867
Chaetozone setosa. - Fauvel, 1927: 101, Fig.
35 d-k. - Hartman 1965a: 166; 1969: 241,
Fig. 1-3.
Records. — Common off Beaufort in 40-200 m
(21, *).
Distribution. — Arctic and in temperate waters
of the North and South Atlantic; southern
California; Mediterranean; probably cosmopoli-
tan; from 40 to 4,436 m.
FAMILY ORBINIIDAE
Notes on the Genera of the
Subfamily Orbiniinae
While the definitions of genera of the sub-
family Orbiniinae published by different workers
readily distinguish typical species, they are not
in absolute agreement and "difficult" species
may be referred to different genera or sub-
genera according to which authority is con-
sulted. The orbiniids recorded from North Caro-
lina include several of these difficult species,
and it was thus necessary to consider the ge-
neric definitions very carefully before the col-
lection could be identified. As the work pro-
ceeded it appeared worthwhile to redefine all
genera of the subfamily Orbiniinae.
Useful discussions of the whole family Orbi-
niidae will be found in Eisig (1914), Fauvel
(1927), Hartman (1957), Pettibone (1957), and
Day (1967). Hartman (1957: 242) divided the
family Orbiniidae into two subfamilies. The
Protoariciinae includes genera with two achae-
tous segments behind the prostomium. We are
not concerned with this subfamily here and for
further details the reader is referred to Hart-
man's account. The subfamily Orbiniinae in-
cludes all the larger orbiniids with one achae-
tous (peristomial) segment behind the pro-
stomium, an eversible epithelial proboscis,
well-developed parapodia and branchiae on
many segments. The Orbiniinae include the
genera Orbinia Quatrefages, Phylo Kinberg,
Scoloplos (Scoloplos) Blainville, Scoloplos (Leo-
da mas) Kinberg, Scolaricia Eisig, Haploscolo-
plos Monro, Califia Hartman, and Naineris
Blainville (with the subgenus Polynaineris
Pettibone).
These genera are distinguished by different
combinations of characters. Generic definitions
will be given later, but meanwhile it may be
noted that Naineris is easily separated by the
possession of a bluntly rounded to square pro-
stomium; all other genera have pointed conical
prostomia.
Scoloplos occupies a central position in the
subfamily. It agrees with Haploscoloplos and
Califia in having none to two foot-papillae ( =
postsetal papillae, podial lobes or podial fringe)
on the posterior thoracic neuropodia and none
to two stomach -papillae ( = subpodial papillae,
ventral papillae or ventral fringe) below the
neuropodia. In distinction to this, typical species
of Orbinia and Phylo have five or more foot-
papillae and numerous stomach-papillae. In-
evitably, "difficult" species occur; Orbinia john-
soni (Moore) has only 1 foot-papilla but a maxi-
mum of 3 stomach -papillae; Orbinia dubia Day
has a maximum of 3 foot-papillae and up to
12 stomach-papillae; Phylo norvegictis (Sars)
and Orbi>iia exarmata (Fauvel), have more than
10 foot-papillae but no stomach-papillae; Scolo-
plos (Scoloplos) riseri Pettibone has a maxi-
mum of 3 foot-papillae and up to 9 stomach-
papillae. It may be noted too, that it is sometimes
83
difficult to distinguish between the foot-papillae
on the lower edge of the neuropodium and the
stomach-papillae on the ventrum immediately
below. For these reasons it is suggested that
the distinction between Orbinia and Phylo on
the one hand, and Scoloplos, Haploscoloplos,
and Califia on the other, be based on the total
number of papillae behind and below the pos-
terior thoracic neurosetae. According to Eisig
(1914), certain specimens of Scoloplos armiger
which is the type species of Scoloplos have a
maximum of two foot-papillae and a maximum of
two stomach-papillae giving a total of four
papillae of both types. On this basis all species
with a total of five or more foot-papillae plus
stomach-papillae would be included in Orbinia
or Phylo and those with four or less would be
referred to Scoloplos, Haploscoloplos, or Califia.
Orbinia, the type genus of the family is closely
related to Phylo. The latter is regarded as a
valid genus by Hartman (1957) and Day (1967)
but as a subgenus of Orbinia by Pettibone
(1957, 1963). Phylo is distinguished from Orbinia
by the possession of heavy spines or "spear-
headed spines" in the posterior neuropodia.
These form the anterior row of neuropodial
setae but the inferior ones are not very distinct
and only the superior ones of adult specimens
are greatly elongated so that they project well
above the dorsum. In P. ornatus (Verrill) the
spines are less distinct than usual so that Petti-
bone (1963a) has referred it to Orbinia. In most
species, however, the spines with their spear-
shaped or arrowshaped ends are very distinc-
tive and Phylo is accepted here as a valid genus.
As mentioned earlier, the remaining genera
with four or fewer foot-papillae or stomach-
papillae are all related to Scoloplos. They are
distinguished from one another by the seg-
mental position of the first pair of branchiae,
the structure of the neuropodium in both the
thorax and abdomen and the nature of the
neuropodial setae.
Scoloplos has been accepted as a valid genus
by all modern workers. It is commonly divided
into two subgenera — Scoloplos (Scoloplos) with
the type species Scoloplos armiger (Muller) and
Scoloplos (Leodamas) with the type species
Leodamas verax Kinberg. Pettibone (1957), in
her diagnosis of the two subgenera, stresses
the fact that in Leodamas (sic), the thoracic
neuropodial lobes are low, rounded, and with-
out papillae while in Scoloplos sensu strictu,
the thoracic neuropodial lobes are provided
"with papilla in middle of lobe, with or without
1 or 2 additional papillae on lower part." Hart-
man (1957), in her diagnosis of the two sub-
genera, stresses the presence of two or more
acicula in the abdominal neuropodia and bran-
chiae starting on setiger 10 or not until setiger
26 in Scoloplos sensu strictu and a single heavy
aciculum in abdominal neuropodia and bran-
chiae from setiger 5 or 6 in Scoloplos (Leo-
damas). Afrer consulting the descriptions of
many species of Scoloplos, it would appear that
the most useful criterion for the distinction of
the two subgenera is the appearance of the
first pair of branchiae on setiger 5-6 in Scolo-
plos (Leodamas) and the appearance of bran-
chiae on setiger 8-10 or some subsequent seg-
ment in Scoloplos (Scoloplos). The number of
acicula in the abdominal neuropodia does not
appear to be constant and is not stated in the
descriptions of many species.
Scolaricia has been accepted as a valid genus
although few species have been assigned to it.
Since different workers have used different
combinations of characters to distinguish Sco-
laricia from Scoloplos, Eisig's original descrip-
tion of the type species Scolaricia typica was
consulted and an attempt was made to examine
the type specimen from Italy., This could not be
traced in the Paris museum but eventually three
specimens from Marseilles, identified by Dr.
G. Bellan, were obtained with the help of Dr.
H. Zibrowius. The following diagnosis was ex-
tracted from Eisig's original account and am-
plified from the Marseilles specimens.
Scolaricia typica Eisig, 1914 from Marseilles.
Description. — Length up to 150 mm for
250 segments. Prostomium pointed. Thorax flat-
tened with 18-21 setigers. Transition to abdomen
abrupt. Branchiae narrow and lanceolate, pre-
sent from setiger 15 or 16. Postsetal lobe of
notopodium short and tapered in thorax, be-
coming weakly scalpel-shaped and as long as
branchiae in anterior abdomen. No interramal
cirri. Thoracic neuropodia as vertical halfmoon-
shaped ridges, with a median notch but without
a foot-papilla in setigers 1-7, becoming oval
with a more dorsal notch and a single foot-
papilla on last three to six thoracic setigers.
Stomach -papillae entirely absent. Abdominal
84
Figure 12. — Dodecaceria corallii a, anterior end; b, spoon-shaped hook. Ovbinia cuiiericcuia n. sp. c, anterior end;
d, posterior view of 10th foot; e, thoracic hook; f, flail-seta from abdominal neuropodium; g, 6th abdominal foot.
Orbinia riseri h, anterior end; i, posterior view of 18th foot; j, thoracic hook; k, flail-seta from abdominal
neuropodium; 1, posterior abdominal foot. Scoloplos capensis m, anterior end; n, posterior view of 10th thoracic-
foot; o, thoracic hook; p, flail-seta from abdominal neuropodium; q, 5th abdominal foot.
85
neuropodia with a broad basal flange and a
bilobed dorsal projection. Inner (dorsal) lobe
longer and stouter than outer lobe. No ventral
cirri.
Thoracic notosetae as numerous crenulate
capillaries; abdominal notosetae similar but with
a few short forked setae in addition. Thoracic
neurosetae as four or five rows of completely
smooth, slightly bent hooks with guards plus
small tufts of crenulate capillaries at the median
notch of the neuropodium and at the upper
margin. Abdominal neurosetae include three or
four acicula with projecting ends, two or three
normally tapered crenulate capillaries and five
or six flail-setae with stout, lightly serrated shafts
and tapered tips sometimes bent at an angle.
Remarks. — In discussing the diagnostic
characters of his new genus, Eisig (1914: 427)
does not mention the flail-setae but stresses the
notch in the thoracic neuropodia, the absence
of stomach-papillae, the presence of only one
foot-papilla on the last few thoracic segments
and the lamellar expansion at the base of the
abdominal neuropodia.
Fauvel (1927: 7) in his introduction to the
family Ariciidae, states that flail-setae are pe-
culiar to the genus Scolaricia and uses the
notch in the thoracic neuropodia in his generic
key. Hartman (1957: 295) uses the flail-setae to
distinguish Scolaricia from Scoloplos. Pettibone
(1957) does not mention Scolaricia since it was
not represented in the collections of the U.S.
National Museum. Day (1957: 547) distinguishes
Scolaricia from Scoloplos by the possession of
flail-setae, the notched thoracic neuropodia,
and the lamellar base of the abdominal neuro-
podia. Unfortunately later studies have shown
that none of these characters are peculiar to
Scolaricia. The examination of Scolaricia typica
showed that the notches in the thoracic neuro-
podia are neither obvious nor deep but are
mere depressions on either side of the origin
of the single foot-papilla. They may be seen
in species of Scoloplos, as illustrated for Scolo-
plos armiger, the type species of Scoloplos, by
Hartman (1957, pi. 29: Fig. 2.) Again the flail-
setae are differentiated from normal crenulate
capillaries to varying degrees and are present
in Scoloplos riseri Pettibone, Scoloplos acme-
ceps Chamberlin, and even to some degree in
Scoloplos armiger. The lamellar base to the
abdominal neuropodia is also developed to
varying degrees; to a slight extent in Scoloplos
armiger and to a greater extent in Haploscolo-
plos pugettensis (Pettibone) and Scoloplos
acmeceps as illustrated by Hartman (1957: pi.
26: Fig. 3, pi. 30: Fig. 2.) In Orbinia dubia Day,
the lamellar lobe is very large and even notched
in the posterior abdomen. As this species has a
row of 20 + 20 stomach-papillae on the pos-
terior thorax it obviously does not fit in the genus
Scolaricia. In the face of all this evidence it will
be obvious that Scolaricia must become a syn-
onym of Scoloplos.
The genus Haploscoloplos was erected by
Monro (1933a) with Scoloplos cylindrifer Ehlers
as the type species. It is generally similar to
Scoloplos but is distinguished by the absence
of hooks in the thoracic neuropodia, the thora-
cic neurosetae being all crenulate capillaries.
It may be noted, however, that the development
of neuropodial hooks is very variable in Scolo-
plos, some species having very few hooks, and
it is suspected that very juvenile specimens of
Scoloplos lack hooks in most or all thoracic
segments. Haploscoloplos is accepted as a valid
genus by Hartman (1957) but is regarded as a
synonym of Scoloplos sensu strict!/ by Pettibone
(1957). She writes: "Until it can be established
how much the abrasive action of certain sub-
strata has to do with the formation of certain
types of crotchets [here termed hooks] from
capillaries, the character does not seem to be
a good one." To me there seems no possibility
that the normal hooks with rounded ends and
guards could have been formed from broken
or abraded crenulate capillaries. While I recog-
nize that juvenile specimens of some species
of Scolojylos may be wrongly assigned to Haplo-
scoloplos, I agree with Hartman and Monro in
recognizing it as a valid genus.
The genus Califia was erected by Hartman
(1957), with C. calida Hartman as the type spe-
cies. It differs from Scoloplos in having brush-
tipped hooks, as well as crenulate capillaries
in the anterior thoracic neuropodia. Pettibone
(1957) described Scoloplos (Scoloplos) schniitti
with similar characters but with normal hooks
as well as brush-tipped ones. It is obvious that
both should be included in Califia and the only
question that remains is whether Califia should
retain its generic rank or is better regarded as
a subgenus of Scoloplos. Pettibone (1963a) re-
gards it as a subgenus of Scoloplos. I agree
86
with Hartman in recognizing Califia as a valid
genus.
Generic definitions
Having considered the main distinguishing
features of the various genera and subgenera
of the subfamily Orbiniinae, their diagnostic
characters are set out below. It will be noted
that certain characters are not mentioned since
they are regarded as being of specific impor-
tance only. Among these are the notch in the
neuropodial lobe of the thorax, the presence of
a lamellar base or ventral cirri on the abdominal
neuropodia, the presence or absence of flail-
setae, and the number of acicula in abdominal
neuropodia. Again, the presence of only one
achaetous segment behind the prostomium is
omitted for this applies to all genera of the
Orbiniinae.
Naineris Blainville, 1828
Orbiniinae with prostomium rounded to square
in front. First pair of branchiae starting on any
thoracic setiger from 2 to 23. Thoracic neuro-
podia with none to two foot-papillae ; no stomach-
papillae. Thoracic neurosetae include crenulate
capillaries, hooks and sometimes subuluncini
(intermediate forms) as well.
Type species: Nais quadricuspida Fabricius,
1780.
Orbinia Quatrefages, 1865
Orbiniinae with prostomium pointed . First pair
of branchiae on setiger 5-9. Posterior thoracic
segments usually with several foot-papillae and
numerous stomach-papillae but with at least
five papillae of both types combined on some
thoracic segment. Thoracic neurosetae include
blunt hooks (exceptionally replaced by subu-
luncini in 0. exarmata) and usually crenulate
capillaries; heavy spear-tipped spines or brush-
tipped setae absent.
Type species: Aricia sertulata Savigny, 1820
(includes Aricia cuvieri Audouin and Milne-
Edwards, 1833).
Phylo Kinberg, 1866
Orbiniinae with prostomium pointed. First
pair of branchiae on setiger 5-7. Resembles
Orbinia in having posterior thoracic segments
usually with several foot-papillae and numerous
stomach-papillae but with at least five papillae
of both types combined on some thoracic seg-
ment. Thoracic neurosetae include heavy spear-
tipped spines as well as blunt hooks and crenu-
late capillaries.
Type species: Phylo fell x Kinberg, 1866.
Scoloplos Blainville, 1828
Orbiniinae with prostomium pointed. First pair
of branchiae on setiger 5 or some subsequent
thoracic segment. Posterior thoracic segments
with few or no foot-papillae and stomach-papillae
and never more than four papillae of both types
combined. Thoracic neurosetae include blunt
hooks and usually crenulate capillaries; heavy
spear-tipped spines and brush-tipped setae
absent.
Type species: Lumbricus armiger Muller,
1776.
Subgenus Scoloplos (Scoloplos). - Scoloplos
species with first pair of branchiae on setiger
8-10 or some subsequent thoracic segment.
Subgenus Scoloplos (Leodamas). - Scoloplos
species with first pair of branchiae on setiger
5-6.
Type species: Leodamas verax Kinberg, 1886
Haploscoloplos Monro, 1833
Orbiniinae with prostomium pointed. First
pair of branchiae on setiger 9 or subsequent
thoracic segment. Posterior thoracic segments
with few foot-papillae and stomach-papillae
and never more than four papillae of both types
combined. Thoracic neurosetae include crenu-
late capillaries only.
Type species: Scoloplos cylindrifer Ehlers,
1905.
Califia Hartman, 1957
Orbiniinae with prostomium pointed. First pair
of branchiae on setiger 8 or subsequent thoracic
segment. Posterior thoracic neuropodia with not
more than two foot-papillae. No stomach-papil-
la. Thoracic neurosetae of anterior segments
include brush-tipped setae, crenulate capil-
laries and sometimes blunt hooks.
Type species: Califia calida Hartman, 1957.
In accordance with these amended generic
definitions certain species will have to be trans-
XT
ferred to different genera. Among these are: Scoloplos (Scoloplos) risen Pettibone, 1957
Scolaricia typica Eisig. 1914 becomes Scolo- becomes Orbinia riseri (Pettibone) n. comb.
plos (Scoloplos) typica (Eisig). n. comb. Scoloplos (Scoloplos) schmitti Pettibone, 1957
Scolarcia haasi Monro. 1937 becomes Scolo- becomes Cal ifia schmitti (Pettibone)
plos (Scoloplos) haasi (Monro), n. comb. Scoloplos (Leodamas) fimbriatus Hartman,
Scolaricia capensis Day, 1961 becomes Scolo- 1957 becomes Orbinia finibriata (Hartman)
plos (Scoloplos) capensis (Day) n. comb. n. comb.
Key to genera and subgenera of Orbiniinae
and the species recorded from North Carolina
1 Prostomium rounded to square in front. (0-2 foot-papillae; no
stomach-papillae. (Naineris) No N.C. record
1' Prostomium conical and pointed 2
2 At least 5 papillae (foot-papillae plus stomach-papillae) on
one or more posterior thoracic segments 3
2' Not more than 4 papillae (foot papillae plus stomach-papillae
on any posterior thoracic segment 6
3 Neurosetae of posterior thoracic segments include one or a
series of heavy spear-tipped spines as well as blunt hooks.
(Phylo) 4
3' Neurosetae of posterior thoracic segments lack heavy spear-
tipped spines. (Orbinia) 5
4 No interramal cirrus between notopodia and neuropodia of
abdominal segments. Heavy spear-tipped spines not always
distinct Phylo ornatus
4' A long interramal cirrus on anterior abdominal segments.
Heavy spines with arrow-shaped tips Phylo felix
5 Branchiae from setiger 6. No interramal cirrus on abdominal
segments Orbinia anwricana
5' Branchiae from setiger 8-10. An interramal cirrus on abdomi-
nal segments Orbi)iia riseri
6 Anterior thoracic neurosetae include brush-tipped setae as
well as crenulate capillaries and sometimes blunt hooks.
(Califia ) No N.C. record
6' Anterior thoracic neurosetae without brush-tipped setae 7
7 Thoracic neurosetae of adults include only crenulate capil-
laries. (Haploscoloplos) 8
7' Thoracic neurosetae of adults include rows of hooks as well
as crenulate capillaries. (Scoloplos) 10
8 An interramal cirrus between abdominal notopodia and neuro-
podia 9
8' No interramal cirrus. [Abdominal neuropodia with lobes longer
than branchiae; no ventral cirrus] Haploscoloplos foliosus
9 A ventral cirrus below anterior abdominal neuropodia Haploscoloplos fragilis
9' No ventral cirrus Haploscoloplos robiistiis
10
10'
11
11'
112
12'
First pair of branchiae on setiger 5-6. Scoloplos (Leodamas).
[Abdpminal neuropodia with inner lobe much shorter than
outer; no ventral cirrus] Scoloplos (L.) rubra
First pair of branchiae on setiger 8-10 on some subsequent
segment Scoloplos (Scoloplos) 11
One or two foot-papillae on last thoracic neuropodia; 1-
ventral cirri on anterior abdominal segments Scoloplos (S.) acmeceps
One foot-papilla on last thoracic neuropodia; no ventral cirri
on anterior abdominal segments 12
Eighteen or nineteen thoracic setigers with branchiae from
setiger 17-18. No brown bars on posterior thorax Scoloplos (S.) acmeceps
Seventeen or eighteen thoracic setigers with branchiae from
setiger 13-14. Brown bars across thorax from setiger 9 Scoloplos (S.) capensis
Phyla ornatus (Verrill, 1873)
Orbinia ornata. - Hartman, 1945: 28. -Pettibone,
1963a:285, Fig. 75 a-b.
Phylo ornatus. - Hartman, 1957: 265, pi. 24:
Fig. 1-10; - 1969: 41, Fig. 1-5.
Records. — Cape Hatteras area to Beaufort,
intertidal to 10 m (5, 9, 11, 13, 18).
Distribution. — Massachusetts to the Gulf of
Mexico; California to Pacific coast of Mexico;
intertidal to 32 m.
Phylo felix Kinberg, 1866
Phylo felix. - Hartman, 1957: 262, pi. 23: Fig.
1-7.
Re cords. — Off Beaufort in 120-200 m (*).
Distribution. — Brazil south to Antarctica;
California to Pacific coast of Mexico; in shallow
dredgings to 200 m.
Orbinia americana New Species
Figure 12c-g
Holotype.— USNM 43138.
Description. — Holotype incomplete, and 10
mm long for 38 segments. Prostomium acutely
pointed and without eyes. Thorax (Figure 12c)
of 21 flattened setigers. Branchiae from setiger
5. Notopodial lobes evident from setiger 1.
Thoracic neuropodia (Figure 12d) as lateral
ridges with an increasing number of foot-
papillae; setiger 3 with three; setiger 4 with
four; setiger 18 with seven. Stomach-papillae
from setiger 17 to 22 with a maximum of seven
stomach-papillae on setiger 20. Thoracic noto-
setae as tufts of crenulate capillaries plus two
or three forked setae with unequal prongs.
Thoracic neurosetae as three rows of hooks
with a few crenulate capillaries behind them at
upper and lower margins. No enlarged spines.
Individual hooks (Figure 12e) with smooth,
curved ends.
Abdominal parapodia dorsal in origin, with
two or three foot-papillae below neuropodia of
first few. Branchiae larger than slender post-
setal lobes of notopodia. No interramal cirri.
Neuropodia (Figure 12 g) bilobed with inner
lobe shorter and stouter than outer. First five
abdominal neuropodia with two cirri basally;
sixth and subsequent neuropodia with only the
superior one persisting as a ventral cirrus.
Abdominal notosetae include numerous capil-
laries and three or four short forked setae.
Abdominal neurosetae include two acicula, one
or two long flail-setae with faintly serrated
shafts and abruptly tapered tips (Figure 12f),
and two or three shorter, evenly tapered crenu-
late capillaries.
Remarks. — This species is allied to Orbinia
sioani Pettibone, but there are fewer thoracic
89
segments, fewer foot-papillae and fewer stom-
ach-papillae.
Rieords. — One specimen off Beaufort in 122
m i*).
Orbinia riseri (Pettibone, 1957)
New Combination
Fisoire 12h-l
Scoloplos (Scoloplos) riseri Pettibone, 1957:
163. Fig. 2 a-d; 1963a: 288, Fig. 74 e-f.
Description. — Body without color markings,
50-60 mm long. Prostomium (Figure 12h) sharply
conical. Thorax of 18-19 setigers with branchiae
from 10th setiger (or 8th in juveniles). Noto-
podial postsetal lobe well developed from setiger
1. Thoracic neuropodia as lateral ridges with
one large foot-papilla from setiger 3, two on
10th foot and three on 16th. Stomach-papillae
increasing from 1 below 16th foot to 14-18 at
junction of thorax and abdomen (Figure 12i).
Abdominal notopodia as chopper-shaped lobes
smaller than the branchiae. A well-developed
interramal cirrus (Figure 121). Abdominal neuro-
podia bilobed; several stomach-papillae below
first few abdominal neuropodia decreasing to
2 on setiger 26 (7th abdominal segment) and
thereafter flattened to form two lamellar expan-
sions at base of neuropodium.
Thoracic notosetae as crenulate capillaries.
Abdominal notosetae with three or four forked
setae among capillaries. Thoracic neurosetae
in four or five vertical rows; an anterior row of
crenulate capillaries with blades broadened
basal ly, then two or three rows of hooks and a
posterior row of fine capillaries. Individual hooks
(Figure 12j) blunt and well serrated, with deli-
cate sheaths. Abdominal neurosetae with two
acicula, two or three fine crenulate capillaries,
and one or two long flail-setae (Figure 12k) with
serrated shafts and abruptly tapered tips.
Remarks. — This species has been described
in some detail as the original description was
based on a specimen only 25 mm long and pre-
sumably a juvenile with branchiae from setiger
8. It has been transferred to the genus Orbinia
since it possesses many stomach-papillae.
Records. — Off Beaufort in 120-160 mm (*).
Distribution. — Massachusetts to the Gulf of
Mexico; intertidal to 160 m.
Scoloplos (Scoloplos) capensis (Day, 1961)
New Combination
Figure 12m-q
Scolaricia capensis Day, 1961: 480, Fig. 1 p-s;
1967: 549, Fig. 23.5. a-d; - Day, Field, and
Montgomery, 1971: 122.
Description. — Body up to 25 mm long for 105
segments with brown bars across thorax from
setiger 9 to 17 when fresh. Prostomium (Figure
12m) sharply pointed, longer than broad. Thorax
with 16-17 setigers and branchiae from setiger
13-14. Notopodia with postsetal lobes obvious
from setiger 1, slender anteriorly, longer and
stouter in abdomen but never as stout as the
branchiae. Thoracic neuropodia (Figure 12n)
as low lateral ridges with a single median foot-
papilla from setiger 4, increasing in length and
moving to a superior position towards end of
thorax. No stomach-papilla. Abdominal seg-
ments without interramal cirri. Abdominal neuro-
podia bilobed with inner lobe rather longer than
outer and base expanded to form a broad brown
unnotched lateral lamella. No ventral cirri.
Notosetae of thorax and abdomen as crenu-
late capillaries, there being no forked setae.
Thoracic neurosetae in five vertical rows; an
anterior row of crenulate capillaries, then
three rows of hooks and finally a posterior row
of crenulate capillaries. Individual hooks (Figure
12o) almost straight, blunt and serrated, with
delicate guards. Abdominal neurosetae sup-
ported by one aciculum and include a few small
crenulate capillaries and one or two longer and
stouter flail-setae (Figure 12p).
Remarks. — In accordance with the generic
revision described above, Scolaricia capoisis
becomes Scoloplos (Scoloplos) capensis. The
specimens from North Carolina agree with the
holotype from South Africa apart from the fact
that the fresh specimens had well-marked brown
bars across posterior thoracic segments which
were not observed on the holotype. S. (S.) capen-
sis is close to S. (S.) acmeceps Chamberlin; but
it has fewer thoracic segments, the gills appear
more anteriorly, and there are no forked setae.
No brown bars were noted on S. (S.) acmeceps.
Records. — Common off Beaufort in 40-200 m
(21, *). This is a new record for the United
States.
Distribution. — South Africa in 86 m.
90
Scoloplos (Scoloplos) cf. acmeceps
Chamberlin, 1919
[?] Scoloplos acmeceps. - Hartman, 1957: 282,
pi. 30: Fig. 1-7; 1969: 43, Fig. 1-5.
Description. — Body about 20 mm long and
without color markings. Thorax of 18-19 setigers
with branchiae from setiger 17 or 18. Postsetal
lobe of notopodium short and stout; neuropodia
with a single small foot-papilla from setiger 4.
Thoracic neurosetae include numerous broad
crenulate capillaries and a few serrated and
bent hooks inferiorly. Abdomen with notopodia
broadly chopper-shaped. No interramal cirri.
Neuropodia bilobed with inner ramus longer
than outer, and base forming a broad, unnotched
flange in place of ventral cirri. Abdominal noto-
setae as crenulate capillaries only; neurosetae
as a few smoothly tapered capillaries and four
or five flail-setae with abruptly tapered tips.
Remarks. — Two incomplete specimens were
obtained. They agree with Dr. Hartman's de-
scription except that they lack forked setae
among the abdominal notosetae.
Records. — Off Beaufort in 120-200 m (*).
Distribution. — (of S. acmeceps) Alaska south
to western Mexico; intertidal.
Scoloplos (Scoloplos) armiger (Muiler, 1776)
Scoloplos armiger. - Hartman, 1957: 280, pi. 29:
Fig. 1-7. - Pettibone, 1963a: 292, Fig. 76 h-i. -
Day, 1967: 544, Fig. 23.6. k-n.
Records.— Off Beaufort in 120-200 m (21, *).
Distribution. — Reportedly cosmopolitan in
intertidal to shelf depths but many records
doubtful.
Scoloplos (Leodamas) rubra (Webster, 1879)
Scoloplos (Leodamas) rubra. - Hartman, 1951:
74, pi. 20: Fig. 1-6; 1957: 291, pi. 32: Fig. 1-6.
Remarks. — The only points worth adding to
Dr. Hartman's description are that the thorax
may extend over 24-28 segments, that while
most of the thoracic neuropodia lack foot-papil-
lae, one may occur on the last one or two seg-
ments and that the abdominal neurosetae are
all smoothly tapered crenulate capillaries, flail-
setae being absent.
Records. — Cape Hatteras area to Beaufort,
intertidal to 200 m (3, 5, 11, 13, 18, *).
Distribution. — North Carolina to Florida;
intertidal to 200 m.
Haploscoloplos foliosus Hartman, 1951
Haploscoloplos foliosus Hartman, 1951: 78.
Remarks. — Only two juveniles measuring 8
mm were obtained. Their characters agree with
Dr. Hartman's description but adult specimens
are required to confirm the record.
Records. — Cape Hatteras area to Beaufort,
intertidal and 200 m (18, *).
Distribution. — North Carolina to the Gulf of
Mexico; intertidal to 200 m.
Haploscoloplos fragilis (Verrill, 1873)
Haploscoloplos frag His. - Hartman, 1951: 76, pi.
21: Fig. 1-3; 1957: 271, pi. 25: Fig. 1-3.
Scoloplos (Scoloplos) fragilis. - Pettibone,
1963a: 290, Fig. 76 a-f.
Records. — Cape Hatteras area to Beaufort,
intertidal to a few meters (3, 5, 9, 11, 13, 17, 18,
*).
Distribution. — Gulf of St. Lawrence to Florida
and the Gulf of Mexico; intertidal to 100 m.
Haploscoloplos robustus (Verrill, 1873)
Haploscoloplos bustoris. - Hartman, 1945: 30.
Haploscoloplos robustus. - Hartman, 1951: 78,
pi. 21: Fig. 4-6; 1957: 272, pi. 25: Fig. 4-6.
Scoloplos (Scoloplos) robustus. - Pettibone,
1963a: 288, Fig. 76 g.
Records. — Cape Hatteras area to Beaufort;
intertidal (2, 3, 5, 9, 11, 13, 17, 18).
Distribution. — Gulf of St. Lawrence to North
Carolina and the Gulf of Mexico; intertidal to
57 m.
91
FAMILY PARAONIDAE
Key to genera and species
1 Prostomium with a median dorsal antenna (may be lost) 2
1 Prostomium without an antenna 8
2 Posterior notosetae include specialized forms as well as
capillaries; posterior neurosetae as capillaries only (Ci)'ro-
phorus) 3
2' Posterior notosetae are all capillaries. Posterior neurosetae
include specialized forms as well as capillaries (Aricidea) 4
2" No specialized setae in notopodia or neuropodia of posterior
segments, only capillaries (Aedicira) 7
3 Specialized notosetae forked. [Body reddish] C. lyriformis
3' Specialized notosetae stout, pointed and acicular with a fine
subterminal filament C. branchiatus
4 Specialized neurosetae with hoods or guards 5
4' Specialized neurosetae without hoods but sometimes acicular
or abruptly tapered to a slender filament 6
5 Specialized neurosetae with a rounded spioniform hood en-
closing a bent unidentate end Ar. cerrutii
5' Specialized neurosetae with a long pointed guard covering
convex side of rostrum Ar.fau veli
6 Specialized neurosetae with a stout shaft tapering to a slender
blade with an oblique breaking plane at junction Ar. fragilis
6' Specialized neurosetae in anterior abdomen with a stout shaft
abruptly tapering to a slender filament; in posterior seg-
ments only the stout sigmoid shaft persists Ar. suecica
7 Setigers 1-3 with a cirriform postsetal lobe on neuropodium Ae. albatrossae
7' Merely a minute postsetal papilla on neuropodium of anterior
segments Ae. belgicae
8 Posterior notosetae include specialized forms (Paraonides) No N.C. record
8' Posterior neurosetae include specialized forms (Paraonis).
[Specialized neurosetae as sigmoid acicular hooks] 9
9 Branchiae cirriform beginning on setiger 6-8 and numbering
9-16 pairs P. gracilis
9' Branchiae compressed and lamelliform beginning on setiger 4
and numbering 16-25 pairs P.fulgens
Cirrophorus lyriformis (Annenkova, 1934) Cirrophorus lyriformis. - Hartman, 1965a: 138.
Cirrophorus Jurcatus. -Hartman, 1969: 69, Fig. 1.
Aricidea (Cirrophorus) furcata Hartman, 1957:
324, pi. 43: Fig. 6. Records.— Off Beaufort in 10-120 m (21, *).
92
Distribution. — Arctic; off California; New
England; Mediterranean; 10 to 500 m.
Cirrophorus branchiatus Ehlers, 1908
Figure 13c
Cirrophorus branchiatus. - Day, 1963a: 423,
Fig. 9 i-o ; 1967: 563, Fig. 24.3. a-e. - Laubier,
1965: 469, Fig. 1.
Description. — Body about 25 mm long for
120 segments. Prostomium bluntly conical with
a short median antenna. Branchiae from setiger
5 to 20-25. Postsetal lobes of notopodia well
developed on anterior segments, short on middle
segments but long on terminal segments. A
heavy spine among notosetae from setiger 13;
each spine (Figure 13c) short, bluntly pointed
with a fine subterminal filament. No specialized
neurosetae.
Remarks. — Cirrophorus aciculatus Hartman
from deep water off southern California and off
Dutch Guinea is very similar but the heavy
notopodial spines are not reported to have a
subterminal filament.
Records.— Off Beaufort in 20 m (*).
Distribution. — Irish Sea; South Africa; Medi-
terranean; British Columbia; 20-200 m.
Aricidea eerruti Laubier, 1967
Figure 13b
Aricidea jeffreysii. - Fauvel, 1927: 75, Fig. 25
a-e. - Hartman, 1957: 322, pi. 43: Fig. 2. -
Day, 1967: 558, Fig. 24.1.j-m.
Aricidea ( Aricidea ) jeffreysii. - Pettibone, 1963a:
305, Fig. 80 a-e. '
Aiicidea eerruti Laubier, 1967: 102, Fig. 1.
Remarks. — The specialized notosetae of pos-
terior segments are sigmoid hooks with spioni-
form hoods (Figure 13b). Pettibone's Figure
80 e does not show this but the shape is well
illustrated by Laubier, Hartman, and Day. Lau-
bier has also shown that the Mediterranean
specimens, doubtfully assigned by Cerruti (1909)
to Scolecolepis ? jeffreysii Mcintosh from Green-
land, are, in fact, distinct.
Records. — Off Beaufort in 20 m (*).
Distribution. — North Atlantic from Ireland
and the Gulf of St. Lawrence to Massachusetts;
Mediterranean and Black Sea; South Africa;
western Canada; in 1-2,000 m.
Aricidea fauveli Hartman, 1957
Figure 13a
Aricidea fragilis. - Fauvel, 1936: 65, Fig. 6, 7
(uou Webster).
Aricidea fauveli Hartman, 1957: 318, pi. 43:
Fig. 1 (synonymy). - Day, 1967: 560, Fig.
24.2. a-d.
Remarks. — This species is characterized by
the special neurosetae of posterior segments
(Figure 13a) which are unidentate with a long
hood on the convex side of the apex.
Records.— Off Beaufort in 20-160 m (*). This
is a new record for the United States.
Distribution. — Morocco and tropical West
Africa to South Africa; 20-200 m.
Aricidea fragilis Webster, 1879
Aricidea fragilis. - Hartman, 1945: 30, pi. 6: Fig.
3; 1957: 317, pi. 43: Fig. 3.
Aricidea (Aricidea) fragilis. - Pettibone, 1965:
129, Fig. 1-3.
Records. — Cape Hatteras area to Beaufort,
intertidal to 200 m (3, 10, 11, 18, 21, *).
Distribution. — Chesapeake Bay to Gulf of
Mexico; intertidal.
Aricidea suecica Eliason, 1920
Aricidea suecica Eliason 1920: 52, Fig. 14-15. -
Hartman, 1957: 318; 1969: 65, Fig. 1-5.
Aricidea (Aricidea) suecica. - Pettibone, 1963a:
307, Fig. 80 f, g.
Records.— Off Beaufort in 120 m (*).
Distribution. — Arctic and North Atlantic to
the British Isles and North Carolina; southern
California; in 5-2,000 m.
Aedicira albatrossae (Pettibone, 1957)
Aricidea (Aricidea) albatrossae
1963a: 305, Fig. 81 a-f.
Pettibone,
Remarks. — Only a single anterior fragment
with 22 segments was obtained, but the first few
93
parapodia are so characteristic that the identity
is certain without the posterior region. This
fresh specimen is slightly different from Dr.
Pettibone's description of the type which had
been in the museum since 1883.
The anterior dorsum is speckled with dark
pigment between the bases of the gills and the
postsetal lobes of the notopodia; the latter are
shown in Pettibone's Figure 81 e as equal in
thickness to the gills, but in this specimen they
are only one-third the thickness. A. albatrossae
is unique among paraonids in having well-
developed postsetal lobes on the anterior neuro-
podia. Those of setigers 1-3 have a broad base
narrowing to a cirriform projection but in subse-
quent segments in the branchial region, only
the broad base remains. Towards the end of
the branchial region the base flattens to form a
small postsetal lamella.
Records. — One specimen off Beaufort in 200
m (*).
Distribution. — Massachusetts; in 150-2,500m.
Aedicira belgicae (Fauvel, 1936)
Aricidea (Aedicira) belgicae. - Hartman, 1957:
327.
Aedicira belgicae. - Day, 1963a: 424; 1967:
563, Fig. 24.3. f-i. - Hartman, 1965a: 133.
Remarks. — While A. albatrossae is easily dis-
tinguished by the possession of cirriform post-
setal lobes on the neuropodia of the first three
setigers, A. belgicae merely has a minute papilla
on all neuropodia up to the middle of the bran-
chial region. Such papillae were not noted in
earlier descriptions and indeed they are easily
overlooked, but they were found in South Afri-
can specimens of A. belgicae. Similar papillae
were found in Aricidea fau veli but not in Aricidea
suecica. It is possible that they occur in several
other paraonids. It may be noted that anterior
fragments of A. belgicae cannot be identified
as the special setae of Aricidea spp. are often
confined to far posterior segments.
Records. — Off Beaufort in 120-200 m (*).
Distribution. — Atlantic from Greenland to
Uruguay, South Africa, and Antarctica; in 30-
4,950 m.
Paraonis gracilis (Tauber, 1879)
Paraonis gracilis. - Hartman, 1957: 330, pi. 44:
Fig. 4, 5; 1969: 75, Fig. 1-3.
Paraonis (Paraonis) gracilis. - Pettibone, 1963a:
301, Fig. 79 a-d.
Paraonis gracilis gracilis. - Day, 1967: 566, Fig.
24.4. a, b.
Records.— Off Beaufort in 120-200 m (21, *).
Distribution. — Atlantic from Greenland and
Denmark to Antarctica and South Africa ; Bering
Sea; southern California; depth 5-2,000 m.
Paraonis fulgens (Levinsen, 1883)
Paraonis julg ens. - Fauvel, 1927: 71, Fig. 24 g-1.
Paraonis (Paraonis) fulgens. - Pettibone, 1963a:
302, Fig. 79 e-f. '
Records. — Cape Hatteras area, intertidal (18).
Distribution. — North Atlantic from Denmark
to the English Channel and Maine to Massa-
chusetts; intertidal to 10 m.
FAMILY OPHELIIDAE
Key to genera and species
1 Body stout and maggot-shaped, not grooved ventrally. [Cirri-
form branchiae on all setigers from second onwards. Lateral
swellings above and below posterior parapodia (Travisia).
Twenty setigers and 3-4 achaetous preanal segments]
1' Body fusiform or slender, grooved ventrally either from second
or 8th-10th setiger onwards
2 Ventral groove and branchiae start on 8th-10th setiger (Ophe-
lia). [Body with 32 setigers with branchiae from 10th to 27th] .
Travisia parva
2
Ophelia denticulate.
94
2' Ventral groove and branchiae start on setiger 2. [Pygidium
tubular]
3 Small lateral eyespots between parapodia from setiger 6-7
(Armandia)
3' No lateral eyespots (Ophelina). [Twenty-seven or twenty-eight
setigers with branchiae from 2nd to 24th, but those on middle
segments reduced and often missing]
4 Twenty-nine setigers, with branchiae from 2nd to 26th. Para-
podial lobes all short and globular
4' Thirty-six or more setigers, with branchiae from 2nd to last.
Anterior feet with elongate and pointed presetal lobes
Ophelina
cylindricaudata
Armandia
maculata
Armandia agilis
Travisia parva New Species
Figure 13d-f
Holotype. — USNM 43126; one paratype,
USNM 43127
Description. — Body of holotype fusiform (Fig-
ure 13f), 12 mm long with 24 segments including
20 setigers and 4 achaetous preanal segments.
Paratype 6 mm long with 20 setigers and 3
achaetous preanal segments. Prostomium sharp-
ly conical; anterior segments triannulate, pos-
terior segments biannulate; no sign of tessela-
tion or papillae. Cirriform branchiae behind
notosetae from setiger 2 to 20 but none on
achaetous preanals. Anterior segments rounded
in section (Figure 13d), 14th and succeeding
segments (Figure 13e) with stout lateral swell-
ings above and below setae. Pygidium with
about eight short blunt lobes encircling anus.
Remarks. — The other species of Travisia re-
corded from the United States namely T. carnea
Verrill, T. profundi Chamberlin, T. grauieri
Mcintosh, T. brevis Moore, T. granulata Moore,
and T. pupa Moore, all have 24 or more setigers;
they differ also in the distribution of the bran-
chiae and the presence of granules or pustules
on the skin.
Records. — Two specimens off Beaufort in
20-80 m (*).
Ophelia denticulata Verrill, 1875
Ophelia denticulata Verrill, 1875: 39. - Tebble,
1953: 362.
Ophelia neglecta Schneider, 1892: 1, pi. 14. -
Fauvel, 1927: 132, Fig. 46 g-h.
Ophelia limacina.
Rathke).
Hartman, 1942a: 130 (non
Records. — Cape Hatteras area to Beaufort,
intertidal to 20 m (18, 21, *).
Distribution. — Maine to North Carolina; Eng-
lish Channel and Atlantic coast of France; inter-
tidal to 20 m.
Armandia maculata (Webster, 1884)
Ophelina maculata Webster, 1884: 322, pi. 11:
Fig. 54, 55.
Armandia maculata. - Hartman, 1942b: 129, Fig.
14 a.
Records.— Oft Beaufort, in 10-40 m (21, *).
Distribution. — Bermuda; intertidal to 40 m.
Armandia agilis (Andrews, 1891)
Ophelina agilis Andrews, 1891a: 289, pi. 15:
Fig. 21-26, 28.
Armandia agilis. - Hartman, 1945: 37; 1951: 97.
Remarks. — A. agilis and A. longicaudata
(Caullery) from the Indian Ocean are very alike.
Both are large species with a long tapered
presetal lobe on the anterior parapodia and gills
from the second to the last setiger. The main
difference is that A. agilis has 36-52 segments
while A. longicaudata has only 30-32.
Records. — Cape Hatteras area to Beaufort,
intertidal to 40 m (5, 9, 11, 13, 15, 18, *).
Distribution. — North Carolina and the Gulf of
Mexico; intertidal to 40 m.
95
Ophelina cylindricaudata (Hansen, 1879)
Figure 13g
Ophelina cylindricaudata. - Stop-Bowitz, 1945:
49. Fig. 5.
[?] Ammotrypane chaetifera Hartman, 1965a:
187. pi. 43.
Description. — Body slender, tapered at both
ends. 8-19 mm long for 26-28 setigers. No
achaetous preanal segments. Prostomium coni-
cal and pointed. A ventral groove starting from
setiger 2. Parapodial lobes uniformly small and
button-shaped. Cirriform branchiae from setiger
2 to 24, followed by 3-4 abranchiate setigers
(Figure 13g). Branchiae of middle segments
small or occasionally absent. No lateral eye-
spots between parapodia. Pygidium cylindrical
and about equal to length of last five or six
setigers. Superior anal cirri as mere crenula-
tions on dorsal lobe over anus; ventral cirrus
stout, annulated and digitiform. Anterior setae
fairly long, those of middle segments progres-
sively shorter and those posterior abranchiate
segments stout, almost acicular.
Remarks. — As originally described by Han-
sen, Ammotrypane cylindricaudata was said to
have 34 setigers, but Stop-Bowitz (1945) states
that the type has 28 setigers and that the bran-
chiae in the middle of the body are very variable;
in some they are slightly reduced, in others
very small or even absent. Possibly A. chaetifera
Hartman, which has gills on a few anterior and
a few posterior segments, but none over most
of the body, is synonymous. Stop-Bowitz has
shown that Ophelina Oersted antedates Ammo-
trypane Rathke, by a few months.
Records. — Common off Beaufort in 120-200
m (21, *).
Distribution. — Norway; Greenland; Canada;
Mediterranean; 30-911 m.
FAMILY SCALIBREGMIDAE
Key to genera and species
1 Anterior setigers with branched gills and posterior setigers
with dorsal and ventral cirri. No acicular setae (Scalibregma).
1' Body without gills or any parapodial projections. No acicular
setae. {Hyboscolex)
S. inflation
H{ longiseta
Scalibregma inflatum Rathke, 1843
Scalibregma inflatum. - Fauvel, 1927: 123, Fig.
44 a-f. - Day, 1967: 590, Fig. 27.2. e-j. -
Hartman, 1969: 313, Fig. 1-4.
Records. — Cape Hatteras area and Beau-
fort, intertidal and 160 m in mud (18, *).
Distribution. — Cosmopolitan from the Arctic
to the Antarctic; intertidal to 160 m in mud.
Hyboscolex longiseta Sehmarda, 1861
Figure 13h, i
Hyboscolex longiseta. - Day, 1967: 584 (synon-
ymy), 588, Fig. 27.2. a-d.
Oncoscolex pacificus. - Hartman, 1969: 311,
Fig. 1, 2.
Description. — Body arenicoliform, up to 20
mm long for 60 segments; color pale to black.
Prostomium (Figure 13h) with stout lateral pro-
jections and thus broadly T-shaped. Eyes visi-
ble if prostomium extended. Buccal segment
very short and achaetous. Anterior segments
(Figure 13i) with four annuli; posterior ones
with two or one. No parapodial projections, the
setae projecting directly from body wall. No
gills or cirri. Setae as smooth capillaries plus a
few short forked setae feathered on inner mar-
gins; no acicular setae.
Remarks. — The synonymy of the genera
Hyboscolex, Oncoscolex, Eumenia, and Poly-
physia is confused. Discussion will be found in
Day (1961: 216; 1967: 584).
Records. — On corals off Beaufort in 18 m
(20,*).
Distribution. — -South Africa and western Ca-
nada to Pacific coast of Mexico; intertidal to
10 m.
96
Figure 13. — Aricidea fauveli a, specialized posterior neuroseta. Aricidea cemitii b, specialized posterior neuroseta.
Cirropkonis branchiatus c, specialized notoseta. Travisia parva n. sp. d, anterior segment; e, two posterior-
segments; f, lateral view of body. Ophelina cylindricandata g, posterior end. Hyboscolex longiseta h, head; i,
parapodium. Leiochrides pallidior j and k, face view and profile of hook. Notomastus americanus n. sp. 1,
cross-section of abdominal segment; m and n, face view and profile of hook.
97
FAMILY ARENICOLIDAE
Only one genus and species known from Records. — Cape Hatteras to Beaufort (1, 5,
North' Carolina. 7, 11, 13, 18, *).
. , . . _. ,__„ Distribution. — Massachusetts to the West In-
F dies, Gulf of Mexico, and California; burrows in
Arenicola cristata. - Wells, 1961: 10. pi. 1-4; muddy sand on sheltered beaches.
1962: 333, pi. 1, 2. - Hartman, 1969: 415,
Fig. 1.
FAMILY CAPITELLIDAE
Key to genera and species
Xot( . — Generic distinctions are based mainly on the distribution of setal types in the thorax. In
the following key this is shown by a thoracic formula where:
C = capillary seta P = peristome
O = without seta H = hooded hook
Notosetae are shown above the line and neurosetae below. The total number of thoracic segments
includes the peristome.
1 Thorax with 9 setigerous segments including the setigerous
peristome. [First 5-7 segments with capillary setae and
remainder with hooks in female but notosetae of setigers 8
and 9 as stout genital setae in male (Capitella). No branchiae
or parapodial projections on abdominal segments] 2
1' Thorax with 10 or more setigerous segments following the
achaetous peristome 2
2 Ten thoracic setigers following peristome.
[Formula: P + — — — ■ = 11. Abdominal neuropodia with
4C + 6H '
short rows of hooks. No gills (Mediomastus)] M. californiensis
2' Eleven thoracic setigers following peristome 3
2" Twelve thoracic setigers following peristome.
12C
[Formula: P + — — - = 13 (Leiochrides)] L. pallidior
2'" Thirteen to fifteen thoracic setigers following peristome 7
3 Last six thoracic setigers with hooded hooks.
5C -1- 6H
[Formula: P -f '— — — = 12. Saclike gills above neuro-
5C + 6H 6
podia of terminal campanulate abdominal segments (Hetero-
mastus)] H. filiform is
3' Thorax without hooks or only in last one or two setigers
(Notomastus) 4
4 All 11 thoracic setigers with capillaries in both rami;
formula: P + — — = 12. Abdominal neuropodia with super-
ior branchial lobes 5
4' First thoracic setiger without neurosetae; last setiger some-
times with hooks. No branchiae 6
98
5
5'
6'
7'
S
8'
Abdominal neuropodia with obvious straplike branchiae
Abdominal neuropodia with small superior branchial lobes . . . .
All thoracic neuropodia with capillaries; formula:
11C
P + t X77 = 12. Abdominal hooks with a single arc of
0 + IOC
5 to 6 denticles above main fang
Last thoracic neuropodium with hooks; formula:
P + — — — — — = 12. Hooks with two arcs of 4 and 2
0 + 9C + 1H
denticles above main fang
Peristome followed by 13 thoracic setigers bearing capillaries;
13C*
formula: P + = 14. Abdominal neuropodia with retrac-
tile gill filaments (Dasybranchus)
Peristome followed by 14-15 thoracic setigers; formula:
P +— 14C or 15C = 1516 Abdominal
(CorO) + (11C-13C) + lHor2H
neuropodia without gills {Leio capitella)
Neuropodial gills with about 30 filaments
Neuropodial gills with few filaments
N. lobatus
N. latericeus
N. he mi pod us
N. am erica nus
L. glabra
D. lumbricoides
D. lunulatus
Capitella capitata (Fabricius, 1780)
Capitella capitata. - Fauvel, 1927: 154, Fig. 55
a-h.-Hartman, 1947a: 404, pi. 43: Fig. 1,2.-
Day, 1967: 595, Fig. 28.2. i-m.
Records. — Beaufort Sound, intertidal and
dredged in muddy sand (11, 13, 15, 18, *).
Distribution. — Cosmopolitan in black mud of
estuaries and protected harbors from 0 to 30 m.
Mediomastus californiensis Hartman, 1944
Mediomastus californiensis Hartman, 1947a:
408, pi. 46: Fig. 3, 4; 1969: 387, Fig. 1-4.
Records. — Common off Beaufort in 10-20 m
(20,21, *).
Distribution. — California; intertidal and es-
tuarine.
Description. — Body about 20 mm long. Pro-
stomium bluntly conical with ventral eyespots.
Thorax including an achaetous peristome and
12 setigerous segments with capillary setae in
both rami. Abdomen of numerous thin-walled
segments with fused notopodial tori and noto-
podial hook-rows continuous across middorsal
line. Interramal organs as prominent papillae
between notopodia and neuropodia from last
thoracic segment onwards. Abdominal neuro-
podia with a small superior branchial lobe and
long rows of hooks almost meeting on mid-
ventral line. Individual hooks (Figure 13j, k)
with short hoods and a crest of seven to nine
subequal denticles above main fang.
Records. — Four specimens off Beaufort in
160 m (*).
Distribution. — British Columbia and Cali-
fornia; intertidal to 89 m.
Leiochrides pallidior (Chamberlin, 1918)
Figure 13j, k
Le ioch rides pallidio r Hartman , 1947a : 429 ; 1969 :
383.
Heteromastus filiformis (Claparede, 1864)
Heteromastus filiformis. - Fauvel, 1927: 150,
Fig. 53 a-i. - Hartman, 1947a: 427, pi. 52:
Fig. 1-4; 1969: 377, Fig. 1-5. - Day, 1967:
601, Fig. 28.3. a-d.
99
Rt cords. — Common in Beaufort Sound on
intertidal mudbanks (11, 12, 13, 15, 18, 20, *).
Distribution. — North Atlantic from Sweden
and Greenland south to Morocco and the Gulf
of Mexico; Mediterranean; South Africa; North
Pacific from Japan to southern California; 0-
100 m.
Notomastus lobatus Hartman, 1947
Notomastus lobatus Hartman, 1947a: 415, pi. 51:
Fig. 1-5; 1969: 399, Fig. 1-5.
Remarks. — A large 150-mm specimen was
obtained from burrows in intertidal mudbanks
near Cape Fear by members of the Zoology
Department, Chapel Hill University. Apart from
the straplike branchiae, the flattened ventrum is
characteristic. Two commensals were found in
the burrows, a polynoid polychaete Lepidas-
thenia sp. and a sluglike bivalve mollusk of the
family Montacutidae which creeps over the body
of the Notomastus.
Records. — North Carolina in intertidal mud
Distribution. — Southern Californiato Mexico;
intertidal to 500 m.
Notomastus latericeus Sars, 1851
Notomastus latericeus. - Fauvel, 1927: 143, Fig.
49 a-h. - Hartman, 1947a: 411. - Day, 1967:
599, Fig. 28.2. a-d.
Records. — Cape Hatteras area and Beaufort,
intertidal to 200 m (5, 13, 18, 21, *).
Distribution. — Cosmopolitan, intertidal to
4,360 m.
Notomastus hemipodus Hartman, 1947
N< it a m a st us (CI is to mastus)hemipodus Hartman,
1947a: 424, pi. 48: Fig. 1-3; 1951: 103, pi.
24: Fig. 1-3; 1969: 393, Fig. 1-5.
Remarks. — N. hemipodus Hartman and N.
aberans Day are closely related. The individual
hooks of N. hemipodus have a single arc of five
or six denticles above the main fang. In N.
aberans there are two arcs of denticles, the first
arc with four or five larger denticles and the
second arc with five to seven smaller denticles.
Records. — Off Beaufort in 120 m (21, *).
Distribution. — Florida and southern Cali-
fornia; intertidal to 120 m.
Notomastus americanus New Species
Figure 131-n
Notomastus n. sp. Day, Field, and Montgomery,
1971: 123.
Holotype. — USNM 43118; 14 paratypes,
USNM 43119.
Description. — Holotype almost complete,
measuring 18 mm for 55 segments. Color yel-
lowish brown with dark faecal pellets visible
through abdominal wall. Prostomium broadly
conical with subdermal eyespots at base. An
achaetous peristome followed by 11 setigerous
thoracic segments. Capillary setae present in
all notopodia but absent in first neuropodium,
present in setiger 2 to 10 but replaced by hooks
11C
in 11th. Formula: P +
12. Ab-
0 + 9C + IH
domen not clearly differentiated but posterior
segments with hook-rows on well-marked ridges.
Nephridial papillae obscure and branchial pro-
jections absent. Hook-rows short throughout;
notopodial rows (Figure 131) separate medially,
neuropodial rows separated by a midventral
gap longer than a hook-row. Individual hooks
(Figure 13m, n) with two arcs of denticles above
main fang, first arc with four denticles, second
with two; formula: MF : 4 : 2.
Remarks. — N. americanus resembles N. teres
Hartman in lacking neurosetae in the first seti-
ger and in having hooks in the neuropodia of
the posterior thorax. It differs in having the
hooks confined to the last thoracic segment and
in having the abdominal hook-rows all short
and widely separated ventrally.
Records. — Fifteen specimens in 80-200 m off
Beaufort (21, *).
Leiocapitella glabra Hartman, 1947
Leiocapitella glabra Hartman, 1947a: 438, pi. 54:
Fig. 1-3.'
Leiocapitella atlantica Hartman, 1965a: 193.
Description. — Prostomium broadly conical,
depressed; eyespots indistinct. All segments
100
short and biannulate and thorax distinguished
from abdomen only by nature of setae. An
achaetous peristome followed by 14-15 seti-
gerous segments bearing capillaries only or
capillaries and hooks. Abdominal segments
without branchial projections, and all bearing
hooded hooks; hook-rows all short and well
separated. Individual hooks with hoods as
broad as long and with three denticles in a
triangle above main fang.
Remarks. — The material from North Carolina
differs in some respects from Hartman's original
description and appears to provide a link be-
tween L. glabra and L. atlantica. The distribu-
tion of the thoracic setae differs in all four
specimens. The first neuropodium may bear
capillary setae on one side and not on the
other, or the first two neuropodia may lack
setae on one side or all the anterior setae may
be present. Similarly, hooks may be restricted
to the last thoracic neuropodium or the last
two thoracic neuropodia. The four formulae are
given below:
P +
14C + (C and H)
0 + 12C + 2H
= 16.
P +
P +
P +
P +
(C
or 0)
+
(C
orO) + 12C
15C
+
1H
13C
+
(C
or H) + (C or
14C
H)
(C
01
0)
+ 12C + 1H
14C
= 16;
= 16;
15;
13C + 1H = 15-
Hartman's original description of L. glabra
was based on two specimens. Both lacked
setae in the first neuropodium on both right
and left sides, but one had capillaries in the
notopodia of 14 segments behind the peristome,
while the other had both capillaries and hooks
in the 15th segment behind the peristome.
The thoracic formulae are thus:
14C
Hartman's description of L. atlantica from
slope depths off Massachusetts gives the fol-
lowing formula:
P +
14C
= 15.
P +
0 + 12C + 1H
15 and
0 + 11C + 2H
If all the specimens are considered together
it becomes obvious that the distribution of setae
on the thorax is very variable; both capillaries
and hooks may occur in the same fascicle, setae
may be present on one side and not the other
or hooks may replace capillaries in the neuro-
podia of one or two segments at the end of the
thorax. It is best to consider the whole group
as one variable species with 15 or 16 thoracic
segments, with neurosetae usually lacking
from the first setiger and with hooks replacing
the capillaries in the last one or two neuropodia.
Records . — Off Beaufort in 40-80 m (*).
Distribution. — Southern California to the
Pacific coast of Mexico; Massachusetts; in
40-200 m.
Dasybranchus lumbricoides Grube, 1878
Dasybranchus lumbricoides. - Hartman, 1947a:
431, pi. 56: Fig. 3, 4; 1951: 103; 1969: 373,
Fig. 1-3.
Records. — Cape Hatteras to Beaufort, inter-
tidal in muddy sand (5, 11, 12, 13, 18, *).
Distribution. — Philippine Islands; Galapagos
Islands; California to Mexico and Florida to
North Carolina in intertidal mud.
Dasybranchus lunulatus Ehlers, 1887
Dasybranchus lunulatus. - Hartman, 1947a: 432,
pi. 56: Fig. 1,2; 1951: 103.
Records. — Beaufort, intertidal (12, 13).
Distribution. — North Carolina, Florida, and
Puerto Rico; intertidal to 4 m in muddy sand.
101
FAMILY MALDANIDAE
Key to genera and species
1 Head with a prominent cephalic keel but not surrounded by a
flattened plate with a raised margin
1' Head with an inclined dorsal plate surrounded by a raised
margin
2 Pygidium petaloid with a central anus (Petaloproetus). [Body
including 21 setigers and 2 achaetous preanal segments]
2' Pygidium funnel-shaped with margin encircled by cirri (Nico-
mache). [A deep pocket above anus]
Neuropodial spines or hooks present in setiger 1. Pygidium
encircled by anal cirri
No neurosetae in setiger 1. No anal cirri. Pygidium as a flat
or concave plate with a dorsal anus above
4 Anal cone protruding beyond ring of long anal cirri and pro-
vided with a stout ventral valve (Praxillella). [Setiger 1 with
2 neuropodial spines. Five achaetous preanal segments] . . . .
4' Anus sunk in a pygidial funnel rimmed with anal cirri
5 Setiger 4 with an anterior membranous collar (Clymenella).
[Eighteen setigers plus 2 achaetous preanal segments. Seti-
ger 1 with about 5 neurosetae]
5' Setiger 4 without a membranous collar
6 Twenty-five or more setigers. No achaetous preanal segments
(Macroclymene). [Eye spots on prostomium. Setigers 1-3
with 1-3 neuropodial spines]
6' Twenty-one or fewer setigers. Achaetous preanal segments
present
7 Setiger 1 with 4 or more neuropodial hooks similar to those
in setiger 4 (Axiothella) . [18 setigers and 2 achaetous pre-
anal segments]
7' Setigers 1-3 with 1-3 neuropodial spines bearing vestigial
denticles or none and quite unlike hooks of setiger 4
(Euclymene)
8 Segments 7-11 dorsally covered with numerous short gill
filaments (Branchioasychis) . [19 setigers]
8' Segments 7-11 without gill filaments. [Cephalic ridge low,
cephalic plate deeply incised laterally (Asychis). Margin of
cephalic plate smooth. Pygidium with a ventral pocket]
9 Body of 19 setigers with numerous blisters or pustules from
6th onwards. Cephalic rim low laterally and forming a
shallow pocket posteriorly (A. elongata)
9' Body of 18 setigers without blisters or pustules. Cephalic rim
high laterally and forming a very deep pocket posteriorly
2
3
P. socialis
N. tri&pinata
Praxillella sp.
5
C. torquata
6
M. zonalis
7
A. mucosa
No N.C. record
B. americana
9
No N.C. record
A. carolinae
102
Petaloproctus socialis Andrews, 189 1
Petaloproctus socialis Andrews, 1891a: 295, pi.
17: Fig. 36-41. - Hartman, 1945: 40, pi. 8:
Fig. 3, 4.
Remarks.— As in the widely distributed spe-
cies P. terricola Quatrefages, there is a ten-
dency for the last few setigers to develop dorsal
fleshy lobes which slant backwards over the
next segment. However, the two species are
easily distinguished for P. terricola has 22 seti-
gers while P. socialis has 19. According to
Arwidsson (1906: 118), Petaloproctus filifer
(Verrill) from Massachusetts has 21 setigers.
Records. — On sheltered sandbanks in Beau-
fort Sound and in dredgings offshore in 20 m
(5,7, 11, 15,21, *).
Distribution. — North Carolina; intertidal to
20 m.
Nicomache trispinata Arwidsson, 1906
Figure 14a
Nicomache trispinata Arwidsson, 1906: 104, pi.
2: Fig. 74-77, pi. 3: Fig. 78, 79, pi. 8: Fig.
257-261, pi. 11: Fig. 349.
Description. — Body broken and number of
segments uncertain. Head speckled with brown;
two groups of ocelli anteriorly; cephalic crest
high, nuchal grooves curved; no cephalic plate
or raised margin. Setigers 1 to 3 with a single
stout, smooth spine in each neuropodium. Seti-
gers 4 and 5 with 10 or more neuropodial hooks.
Individual hooks with poorly marked tendons
and a vertical series of four teeth above main
fang. Posterior end (Figure 14a) with a short,
poorly defined achaetous preanal segment.
Pygidium slanting, with a central anus below
a deep pocket extending forward to level of
achaetous preanal segment. Pygidial funnel
rimmed with 24 subequal triangular cirri.
Remarks. — The material is fragmentary so
that final identification is uncertain. The ob-
served characters agree with Arwidsson 's de-
scription of specimens from Norway which have
23 setigers and 1 achaetous preanal segment.
Records. — Fragments of five specimens from
sand at 20 m off Beaufort (*).
Distribution. — Norway and Greenland; 50-
350 m.
Praxillella sp.
Description. — All specimens broken thus total
length and number of setigers unknown. Pro-
stomium bluntly triangular with numerous eye-
specks. Cephalic plate oval with rim high,
smooth and without obvious lateral or posterior
notches. Nuchal grooves straight, % length of
plate. Setigers 1, 2, and 3 with 2-3-3 neuropodial
spines, each with three vestigial denticles above
rostrum and a whisp of tendon below. Posterior
end with five achaetous preanal segments pre-
ceding pygidial ring. First preanal as long as
last setiger, second slightly shorter, third half
the length of last setiger and fourth and fifth
rudimentary and together equal to length of
third preanal. Pygidium with a circular ridge
bearing a ring of 10 long anal cirri and a pro-
truding anal cone. Anus with a large ventral
valve. Individual hooks with a compact vertical
series of five teeth above main fang and two or
three tendons below.
Remarks. — As far as I am aware, no species
of Praxillella with five achaetous preanal seg-
ments has been recorded from the United States,
but until the number of setigers can be deter-
mined from a complete, unbroken worm, no
specific identification is possible.
Records.— Off Beaufort in 80-200 m (*).
Clymenella torquata (Leidy, 1855)
Clymenella torquata. - Hartman, 1945: 40, pi. 8:
Fig. 1, 2.
Records. — Cape Hatteras to Beaufort; inter-
tidal to 50 m (3, 5, 8, 11, 15, 18, *).
Distribution. — Massachusetts to North Caro-
lina; intertidal to 50 m in sandbanks.
Macroclymene zonalis (Verrill, 1874)
Clymenella zonalis. - Magnum 1962: 7.
Records. — Abundant at Beaufort, intertidal to
40 m (16,21, *).
Distribution. — North Carolina; intertidal to
40 m.
Axiothella mucosa (Andrews, 1891)
Axiothea mucosa Andrews, 1891a: 294, pi. 16:
Fig. 29-35.
103
Axiothella mucosa. - Hartman, 1945: 38, pi. 8:
Fig. 3. G; 1951: 104, pi. 1.
Clymenella mucosa. - Mangum, 1962: 5.
Records. — Beaufort Sound, intertidal (5, 7, 11,
13. 15. 16, *).
Distribution. — North Carolina to Florida;
forms massed tubes in intertidal sandbanks.
Branehioasychis americana Hartman, 1945
Maldane elo)igata. - Andrews, 1891a: 294 (non
Verrill).
BrcuicJiioasychis americana Hartman, 1945: 40,
pi. 9: Fig. 1-4; 1951: 105.
Records. — Beaufort Sound, intertidal (5, 11,
13).
Distribution. — North Carolina to Florida and
the Gulf of Mexico; intertidal to 5 m in soft black
mud.
Asychis carolinae New Species
Figure 14b-f
Asychis carolinae Day, Field, and Montgomery,
1971: 123 (nomen nudum).
Holotype. — USNM 43139; nine paratypes,
USNM 43140.
Description. — Holotype complete, 22 mm long
by 0.5 mm. Body encased in a slender mud tube
and pale in alcohol apart from black flecks on
head and first two segments. Prostomium (Figure
14b) broad, flattened, and smoothly curved in
front. No eyespots. Cephalic plate oval; raised
margin with one deep lateral incision but other-
wise smooth; posterior part forming a deep
pocket extending back to end of head. Cephalic
ridge low, broad, and indistinct. Nuchal grooves
strongly curved. Body with 18 setigerous seg-
ments. Anterior segments increasingly long;
seventh and eighth 8 times longer than broad;
ninth and succeeding ones progressively shorter
and setiger 18 broader than long. No achaetous
preanal segment. Pygidium (Figure 14c) elon-
gated, only slightly shorter than setigers 17 and
18 combined. Anus dorsal with a grooved plate
beyond it overhanging a very deep ventral
pocket. Setiger 1 without neurosetae; neuro-
podia of setigers 2 to 4 with 3-5 neuropodial
hooks; setiger 5 and subsequent segments with
a row of 12-15 hooks. Hooks of setigers 2-4
(Figure 14d) with an arc of 3 denticles above
main fang; hooks of subsequent segments (Fig-
ure 14e, f) with a close-set cap of 10 teeth above
main fang and well-developed tendons below.
Remarks. — Most species of Asychis, including
the type species A. biceps, have a toothed mar-
gin around the cephalic plate. A. capoisis Day
from South Africa has a cephalic margin similar
to that of A. carolinae but it has 19 setigers, 2
preanal segments, and the pygidial plate has
no ventral pocket. Maldane cuculigera Ehlers,
from deep water off Florida has a similar head
but it too has 19 setigers and the pygidium is
truncate. Possibly the closest is Asychis elon-
gata (Verrill), originally named Maldane elonga-
ta but transferred by Verrill (1900) to the genus
Maldanopsis, which is now regarded as a sy-
nonym of Asychis. Dr. Pettibone kindly sent
me a specimen identified by Verrill for compari-
son with A. carolinae. The two species are very
alike in regard to the head and pygidium but
A. elongata has 19 setigers and from setiger 6
onwards the body has a scattering of small
blisters or pustules. In this respect A. elongata
is closer to Branehioasychis americana Hartman
but it lacks the simple branchial filaments
which characterize the latter.
Records. — Common off Beaufort in 120-200
m (21, *).
FAMILY OWENIIDAE
Only one genus and species known from
North Carolina.
Owenia fusiformis Delle Chiaje, 1844
Records. — Beaufort, intertidal to 200 m (5, 11,
13,21,*).
Distribution. — Cosmopolitan in temperate and
tropical seas from 0 to 200 m.
O ire nia fusiformis. - Fauvel, 1927: 203, Fig. 71
a-f. - Day, 1967: 649, Fig. 31.1. e-j.
104
FAMILY STERNASPIDAE
Only one genus and species known from Records. — Off Beaufort in 600 m (*).
North Carolina. Distribution. — Cosmopolitan on stiff mud or
clay bottoms in 0-600 m.
Sternaspis scutata (Ranzani, 1817)
Sternaspis scutata. - Fauvel, 1927: 216, Fig. 76
a-g. - Day, 1967: 648, Fig. 31.1. a-d.
FAMILY FLABELLIGERIDAE
Key to genera and species
1 Neurosetae as annulated capillaries, very similar to notosetae.
Cephalic cage poorly developed. [Eight branchial filaments
(JHplo cirrus) . Branchial filaments equal in thickness; neuro-
setae with a minute terminal hook] D. capensis
1' Neurosetae as falcate hooks, very different from notosetae.
Cephalic cage well developed 2
2 Neurosetae as articulated hooks. Body covered with very long
clavate papillae embedded in a mucilaginous sheath (Fla-
belligera) Flabelligera sp.
2' Neurosetae as simple hooks often with barred shafts. Surface
with short papillae but no mucilaginous sheath 3
3 Neuropodial hooks with unidentate tips. Branchial filaments
arranged in a horseshoe-shaped arc. Body with skin papillae
numerous but not arranged in longitudinal rows (Pherusa) 4
3' Neuropodial hooks with bidentate tips. Branchial filaments
arranged in multiple series on a tongue-shaped lobe. Body
with skin papillae arranged in a few longitudinal rows on
dorsum and ventrum and whole surface encrusted with sand
(Piromis). [Two rows of papillae on dorsum and two on
ventrum] 5
4 Skin papillae encircling anterior margins of segments. Bran-
chial filaments of two sizes Pherusa inflata
4' Skin papillae irregularly scattered. Branchial filaments all
similar Pherusa ehlersi
5 Neuropodial hooks start on setiger 3 (Piromis eruea) No N.C. record
5' Neuropodial hooks start on setiger 4 Piromis eruea websteri
Diplocirrus capensis Day, 1961 r . L „,, , „
form, about 15 mm long. Buccal apparatus re-
Diplocirrus capensis Day, 1961: 509, Fig. 9 a-f; tractile, consisting of an indistinct prostomial
1967: 666, Fig. 32.4. e-j. ridge with ocular pigment, a pair of grooved
palps and eight subequal branchial filaments.
Description. — Body muddy brown, arenicoli- Setiger 1 with two or three elongated notosetae
105
^.•£0 ■/.;-•;.■/ z:';.;;,';-/; '•^;^-:;^-'J"
Hi ViWw.
Figure 14. — Nicomache trispinata a, posterior end. Asychis carolinae n. sp. b, head; c, posterior end; d, hook from
setiger 2; e and f, profile and face view of posterior hook. Pheriisa ehlersi n. sp. g, lateral view of entire worm;
h, lateral view of 8th segment showing papillae; i, neuropodial hook from middle segment; j, elongated neuropodial
hook from setiger 4.
directed forwards and forming a rudimentary
cephalic cage; subsequent segments with shor-
ter setae directed laterally; notosetae include
about five barred capillaries; neurosetae as
three or four shorter, stouter barred setae each
with the curved terminal joint forming a minute
hooklet. Body surface densely covered with
flask-shaped adhesive papillae. Anterior 9-11
segments swollen and without obvious seg-
mental constrictions; posterior 20 segments
forming a narrower "tail" with obvious seg-
mental constrictions.
Remarks. — D. capensis differs from D. glau-
cus (Malmgren), the type species of Diplocirrus
Haase, by having all eight branchial filaments
of equal thickness and in other minor characters
concerning the setae. In Day (1961: 510), I
suggested that the definition of Diplocirrus
be amended to include D. capensis. The amended
definition would include Ilyphagus Chamberlin.
106
Hartman (1965a: 178) described Ilyphagus
octobranchus from slope depths off the coast
of New England which thus becomes Diplocir-
rus octobranchus (Hartman). As Dr. Hartman
has noted, it differs from D. capensis in lacking-
eyes and in having neurosetae which taper
to fine tips instead of ending in minute hooklets.
Records.— Off Beaufort in 120-200 m (21, *).
This is a new record for the United States.
Distribution. — South Africa and North Caro-
lina in 120-200 m.
Flabelligera sp.
Remarks. — Over 36 post-larval specimens
3-4 mm long for 13-17 segments were obtained.
They all contain orange yolk granules in the
midgut. Most of the characters suggest Flabel-
ligera affinis Sars. Each neuropodium has a
single-jointed falcate hook and the surface is
flabby with scattered groups of long club-shaped
papillae. The mucilaginous sheath has not de-
veloped and there is only a single arc of 8 + 8
branchial filaments on each side of the cephalic
ridge.
Records. — Off Beaufort in 40 m (*).
Pherusa infiata (Treadwell, 1914)
Trophonia inflata Treadwell, 1914: 213, pi. 12:
Fig. 33.
Stylarioides inflata. - Hartman, 1951: 98.
Pherusa inflata. - Wells and Gray, 1964: 74. -
Hartman, 1969: 297, Fig. 1-5.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 50 m (15, 16).
Distribution. — Atlantic from North Carolina
to Florida and Pacific from Oregon to Mexico;
intertidal to 50 m.
Pherusa ehlersi New Species
Figure 14g-j
Siphonostomum cariboum. - Ehlers, 1887: 158,
pi. 42: Fig. 6-9, pi. 43: Fig. 1 (jpartim, non
Grube 1859).
Pherusa n. sp. McCloskey, 1970: 26.
Holotype.— USNM 43133.
Description. — Holotype (Figure 14g), 12 mm
long for about 40 segments; maximum breadth
2 mm. Surface with small, cylindrical papillae
and attached sand grains. Papillae (Figure 14h),
longer and less numerous dorsally, shorter and
more numerous ventrally. Cephalic cage well
developed but poorly defined. Setigers 1-3 with
barred capillaries in both rami, all directed for-
wards. Notopodia of subsequent segments
with three of four shorter barred capillaries
directed laterally. Neurosetae of setiger 4 (Fig-
ure 14j) still slender, barred, and directed for-
wards but tips definitely curved; neurosetae of
setiger 5 as five short simple unidentate hooks;
following neurosetae (Figure 14i) similar, but
numbers decreasing to one on middle segments
and increasing again on "tail" segments. Buccal
apparatus including a bulbous lower lip, a pair
of stout grooved palps, a prostomial ridge with
four eyes, and a semicircular cephalic hood
with a single marginal arc of about 20 branchial
filaments of uniform thickness.
Remarks. — The holotype is one of a few
specimens collected by Dr. L. McCloskey from
a coral head growing in 18 m off Beaufort. The
sandy crust on the dorsal surface of the body is
thin and quite different from the hard head
shield of Ph. parmata (Grube) or Ph. inflata
(Treadwell) as redescribed by Hartman (1951).
Nonetheless there are several resemblances to
the latter species. The cephalic cage is similar
and in both, the first neuropodial hooks occur
on setiger 4. However, there are also important
differences. The thinner sandy crust has been
noted. The skin papillae are scattered and do
not form rings encircling the anterior margins
of the segments, as they do in Ph. inflata. The
latter species is also reported to have two sizes
of branchial filaments, 6 larger ones and 13
smaller.
I believe Ph. ehlersi may well be the same as
some, but not all of the specimens from Key
West described by Ehlers (1887) under the
name of Siphonostomum cariboum Grube. All
the characters are the same except what Ehlers
terms the "Kiemfaden tragenden Blatte" here
termed the cephalic hood. Ehlers' description
and his figure of the branchial apparatus (pi.
42, Fig. 7) was based on "einem anderem"
specimen. It shows a tongue-shaped lobe with
numerous branchial filaments which is charac-
teristic of the genus Piromis to which S. cari-
boum has since been referred. However, other
species of Piromis (including the type species
107
P. arenosus Kinberg and P. eruca websteri
described below) have all the neuropodial hooks
annulated and at least the anterior ones bi-
dentate. The skin papillae are in longitudinal
rows, and the whole surface is encrusted with
sand. I believe that Ehlers' "other specimen"
does not refer to the same species as the rest
of his description of S. caribou m but only a
reexamination of Ehlers' specimens can settle
this. In the meantime, it is safer to give the
Beaufort specimen a new name.
Records. — On corals off Beaufort in 18 m
(20. *).
Distribution. — Florida (Key West).
Notes on the genera Piromis Kinberg, 1867
and Pherusa Oken, 1807
Since Pherusa eruca has been recorded from
Cape Hatteras by Wells and Gray (1964) and
this species has many characters which suggest
that it is allied to Trophonia arenosa Webster,
from Virginia and Piromis arenosus Kinberg,
the type species of the genus from South Africa,
it was necessary to examine the three more
carefully. Dr. David George of the British Mu-
seum kindly sent me specimens of Trophonia
eruca Claparede from Naples, which is the type
locality, and Dr. Marian Pettibone sent me the
specimen from Pamlico Sound identified by
Wells as Pherusa eruca, as well as the two
syntypes of Trophonia arenosa Webster from
Virginia. Many specimens of Piromis arenosa
Kinberg were available in my own collections
in the University of Cape Town.
It may be said at once that Wells' specimen
from Pamlico Sound is not a flabelligerid at all.
It has had its head removed, but obviously
belongs to the family Poecilochaetidae.
The genus Piromis (synonym: Semiodera
Chamberlin), is characterized by having the
branchial filaments arising from the surface of
a tongue-shaped lobe in two series each with
many irregular rows, whereas Pherusa has one
series of branchial filaments arising from the
edge of a semicircular hood above the cephalic
ridge. In addition, Piromis arenosus has biden-
tate neuropodial hooks, a sandy crust covering
the body, and relatively few skin papillae, those
on the dorsum and ventrum (but not the para-
podia) being arranged in a few longitudinal
rows. These characters are shared by Trophonia
eruca Claparede and Trophonia arenosa Web-
ster, both of which have been referred to the
genus Pherusa, while many other species of
Pherusa have unidentate neuropodial hooks,
numerous scattered skin papillae, and no sandy
crust. As will be shown, both Trophonia eruca
and T. arenosa, when dissected, proved to have
the branchial filaments arranged in multiserial
rows on a tongue-shaped lobe. They are in fact
species of Piromis. Thus the genera Piromis and
Pherusa may be distinguished not only on the
origin of the branchial filaments, but also on
the bidentate or unidentate neuropodial hooks
and the arrangements of the skin papillae.
Further, Piromis is covered by a sandy crust,
while most species of Pherusa are not; Pherusa
ehlersi described above is intermediate.
Piromis eruca (Claparede, 1869)
New Combination
Trophonia eruca Claparede, 1869: 105, pi. 15:
Fig. 2.
Stylarioides eruca. - Fauvel, 1927: 119, Fig. 42
h-1.
(Nou) Pherusa eruca. - Wells and Gray, 1964:
74.
Material examined. — The following descrip-
tion is based on two specimens of Trophonia
eruca kindly sent me by Dr. 'David George of
the British Museum. They were collected at
Naples (the type locality of T. eruca) and identi-
fied by Mcintosh and the registration number is
1921:5:1:2651/2. Both were brown in alcohol,
the palps and branchial filaments are missing,
and many of the setae are broken.
Description. — Body up to 60 mm long with
73 segments. Surface covered with a sandy
crust, well marked dorsally but indistinct ven-
trally. Skin papillae long, knobbed, and pro-
jecting through the sand. Papillae arranged in
longitudinal rows, two rows dorsally and two
ventrally and groups of about six papillae around
each bundle of setae, those around neurosetae
extending ventrally. No scattered papillae. Buc-
cal apparatus including a pleated lip around
mouth, a pair of large grooved palps, and a
dorsal tongue-shaped branchial lobe with an
indistinct median cephalic ridge. Four close-set
eyes. Branchial filaments in irregular rows in
each group. Cephalic cage poorly defined but
including forwardly directed capillaries of seti-
108
gers 1 and 2. Setae of following segments
shorter and more laterally directed. Notosetae
as five to seven barred capillaries. Neurosetae
as five to seven bidentate hooks from setiger
3 onwards.
Remarks. -r-The buccal apparatus was re-
tracted into a membranous sheath and the
structures recorded above were observed after
dissection. Many of the branchial filaments were
missing but the scars showed that they had been
arranged in two multiserial groups on a tongue-
shaped lobe as is usual in the genus Piromis.
The number of filaments was estimated at 30 on
each side of the cephalic ridge but Fauvel
states that juveniles may have as few as 8-10
filaments. The first hook on setiger 3 was much
longer than those on subsequent neuropodia,
but is otherwise similar to that shown in Fauvel
(1927: Fig. 42 k).
Claparede's Trophonia eruca is a typical
member of the genus Piromis, as shown by the
arrangement of the branchial filaments, the dis-
tribution of the skin papillae, the sandy dorsal
crust, and the bidentate neuropodial hooks. It
differs from the type species, Piromis arenosus
Kinberg, in having only two dorsal and two
ventral rows of skin papillae instead of four
dorsal and four ventral rows.
As noted earlier the record of Wells and Gray
(1964) is incorrect so that P. eruca is not known
from the United States. However, as shown
below, Trophonia arenosa Webster is very close.
Distribution. — North Atlantic from the Eng-
lish Channel to southern France; Mediterra-
nean; intertidal to 10 m.
Piromis eruca websteri New Subspecies
Trophonia arenosa Webster, 1879: 245, pi. 7:
Fig. 92-97. -(Non) Piromisarenosa Kinberg,
1867.
Stylarioides arenosa. - Miner, 1950: 372, pi. 118.
Remarks. — Two syntypes of Trophonia are-
nosa Webster were kindly sent to me by Dr.
Pettibone of the U.S. National Museum, Wash-
ington, D.C. The head of one syntype had been
removed but the other syntype, when dissected,
proved to have a similar buccal apparatus to
that described above for P. eruca. The other
characters, including the arrangement of the
skin papillae and the sandy crust, were also
identical. In fact the only difference observed
is that the first hook appears in the neuropodium
of setiger 4, not 3. This agrees with Webster's
original description. This is a small difference
from the stem form and possibly further collect-
ing may show that the first hooks may appear
on either setiger 3 or 4. However it seems best
to consider Webster's species distinct. Webster's
original name becomes a junior homonym of
P. arenosa Kinberg so I have designated it
P. eruca websteri.
Piromis roberti (Hartman 1951), originally
described as Semiodera roberti may be con-
specific but since the distribution of the skin
papillae and the neuropodial hooks were not
described, the question must be left open.
Distribution. — Virginia, intertidal.
FAMILY SABELLARIIDAE
Key to genera and species
1 Opercular lobes with stout dorsal hooks at base of operular
peduncles; two rows of opercular paleae
1' Opercular lobes without hooks; three rows of paleae. [Middle
row of paleae not forming a cone concealing inner row.
Three parathoracic segments bearing stout oar-shaped
setae (Sabellaria)]
2 Four parathoracic segments. Outer row of paleae with smooth
margins (Lygdamis)
2' Three parathoracic segments. Outer row of paleae with bi-
pinnate lateral projections (Idanthyrsus)
No N.C. record
No N.C. record
109
Middle row of paleae alternately long and short 4
Middle row of paleae all of equal length 5
Innermost row of paleae produced into short points. Ends of
outer paleae with a median denticulate tooth between 2-3
short lateral teeth Sabellaria bella
Innermost row of paleae produced into long spikes resem-
bling long ones of middle row. Outer paleae ending in a long
barbed median spike between two short lateral teeth Sabellaria floridensis
Middle paleae all produced into long, erect spikes Sabellaria gracilis
Middle paleae all curved or hooked 6
Middle paleae long, curved and pointed Sabellaria vulgaris vulgaris
Middle paleae short, hooked and blunt Sabellaria vulgaris beaufortensis
Sabellaria bella Grube, 1870
Sabellaria vulgaris vulgaris Verrill, 1873
Sabellaria bella Hartman, 1944b: 342, pi. 33:
Fig. 53-66.
Records. — Beaufort, intertidal (10).
Distribution. — North Carolina to South
America; intertidal.
Sabellaria floridensis Hartman, 1944
Sabellaria floridensis Hartman, 1944b: 345, pi.
31: Fig. 37-41; 1951: 107.
Records. — Cape Hatteras to South Carolina,
intertidal to 30 m; common on corals (13, 14,
18, 19,20, *).
Distribution. — Florida and North Carolina;
intertidal to 30 m.
Sabellaria gracilis Hartman, 1944
Sabellaria gracilis Hartman, 1944b: 343, pi. 34:
Fig. 66-72; 1969: 507, Fig. 1-5.
Ri cords. — Cape Hatteras area, intertidal (18).
Il< marks. — David W. Kirtley in a personal
communication states that this is a very doubtful
record.
Distribution. — Southern California; ? North
Carolina; intertidal.
Sabellaria vulgaris Hartman, 1944b: 341, pi. 32:
Fig. 42-44. - McCloskey, 1970: 26.
Sabellaria cementarium. -Wells and Gray, 1964:
74 (non Moore).
Remarks. — I am indebted to David W. Kirtley
for informing me that Sabellaria cementarium
recorded by Wells and Gray should be referred
to Sabellaria vulgaris.
Record. — Cape Hatteras to Beaufort, inter-
tidal to 40 m; common on corals and scallops
(2, 3, 5, 7, 10, 11, 13, 15, 18, 20, 21, *).
Distribution. — Massachusetts to Georgia; in-
tertidal to 40 m.
Sabellaria vulgaris beaufortensis
Hartman, 1944
Sabellaria vulgaris beaufortensis Hartman,
1944b: 342, pi. 32: Fig. 45-47; 1945: 43.
Remarks. — Generally similar to S. vulgaris
vulgaris except for the paleae of the middle row
which are shorter, curled inwards, and very
blunt, almost truncate at the end. The two sub-
species are found together; typical specimens
are easily identified but some intermediates
occur.
Records. — Beaufort, intertidal to 40 m (11,
18,*).
Distribution. — North Carolina, 0-40 m.
110
FAMILY PECTINARIIDAE
Key to subgenera and species
1 Cephalic veil quite free from operculum 2
1' Caphalic veil completely or partly fused to operculum. [Oper-
cular rim smooth. Fifteen setigers of which 12 have uncini.
(subgenus Lagis)] No N.C. record
2 Opercular rim dentate (subgenus Amphictene) No N.C. record
2' Opercular rim smooth (subgenus Pectinaria) [Twelve seg-
ments with uncini; uncini with about 7 major teeth; about
15 pairs of scaphal hooks] P. (Pectinaria) gouldii
Pectinaria (Pectinaria) gouldii
Verrill, 1873
Pectinaria (Cistenides) gouldii. - Hartman,
1941b: 328, pi. 50: Fig. 11, 17, pi. 52: Fig. 20.
Cistenides gouldii. - Hartman, 1942a: 74, Fig.
130, 135, 138.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 120 m (2, 3, 5, 7, 9, 11, 13, 15, 18, *).
Distribution. — Massachusetts to Florida and
the West Indies; intertidal to 120 m.
FAMILY AMPHARETIDAE
Key to genera and species
1 Small acicular neurosetae embedded laterally on segments
3-6. Short, stout notopodial hooks sometimes present behind
gills (Melinninae)
1' No neurosetae laterally on segments 3-6. No hooks behind
gills. [Large paleae sometimes present on segment 3 (Ara-
pkaretinae)]
2 Postbranchial hooks present
2' No postbranchial hooks. [Thirteen uncinigerous thoracic seg-
ments. Ridge across segment 6 dentate]
3 Fourteen uncinigerous thoracic segments. Ridge across seg-
ment 6 dentate. All 4 pairs of gills smooth
3' Thirteen uncinigerous thoracic segments. Ridge across seg-
ment 6 smooth-edged. Four pairs of gills; two pairs smooth
and two pairs pinnate
4 Two pairs of gills; [12 uncinigerous thoracic segments. First
two rows on uncini long and ventrally placed]
4' Three pairs of gills; [12 uncinigerous thoracic segments.
Buccal tentacles smooth. No glandular ridges on pro-
stomium]
4" Four pairs of gills
1
3
Melinnopsis atlantica
Melinna metadata
Isolda pulchella
Auchenoplax crinita
Samijthella eliasoni
111
6
6'
Two diverging glandular ridges on prostomium 6
No glandular ridges on prostomium. [Twelve uncinigerous
thoracic segments] 7
Eleven uncinigerous thoracic segments. Small capillaries on
segment 3 Amage auricula
Fourteen uncinigerous thoracic segments. Large paleae on
segment3. [Notopodia with papilliform dorsal cirri] Amphicteis gunneri
Large paleae on segment 3 Amparete acutifrons
Either small capillaries or no setae on segment 3 8
Innermost gill of each group much shorter than others; a wide
median gap between the two groups of gills. Thoracic uncini
with two rows of three teeth Ampharete parvidentata
Gills not very different in length; median gap between groups
of gills not obvious. Thoracic uncini with two rows of five
teeth Ampharete americana
Melinnopsis atlantica Mcintosh, 1885
Melinnopsis atlantica Mcintosh, 1885: 441, 27A:
Fig. 18.
Sytitypes. — Three fragmentary syntypes from
3,110 m off Chesapeake Bay; British Museum
Catalogue No. 1885: 12: 1:330.
Description. — Types broken and poorly pre-
served thus several features uncertain. Pro-
stomium normal but detailed shape doubtful.
Buccal tentacles grooved. An oblique fleshy
ridge slanting up from behind mouth and form-
ing a dorsal crest across segment 6. Presence
or absence of toothed margin to dorsal crest
uncertain due to poor preservation. Segment 3
with a branchial ridge bearing two groups of
four gills arranged three in front and one behind.
Gills long smooth and tapered. Fine acicular
neurosetae embedded in sides of segments 3,
4, 5, and 6. No notosetae on segments 3 and 4
but small tufts of capillaries on segments 5 and
6. Segment 7 and subsequent thoracic seg-
ments with winged capillaries on notopodia and
single row of uncini in neuropodia. Posterior
end of thorax disintegrated and number of
uncinigerous thoracic segments uncertain al-
though 10 recorded in original description.
Thoracic notopodia and neuropodia without
cirriform projections. Abdomen broken and
number of segments now uncertain although
25-35 implied in original description. Unciniger-
ous pennules, rectangular, without superior
papillae. Rudimentary notopodial lobes present
on some abdominal segments. Thoracic uncini
with a single vertical series of four or five teeth
above small rostral point and basal prow.
Abdominal uncini with seven or eight teeth
arranged in two irregular rows above very small
rostral point and basal prow.
Remarks. — Since the original description
given by Mcintosh was incomplete, Dr. David
George of the British Museum kindly sent me
the type material to check certain points. The
emended description given above records for
the first time the presence of a fleshy ridge
across segment 6 and acicular neurosetae on
the side of the same segment. Again, there may
have been more than the 10 uncinigerous thora-
cic segments mentioned in the original descrip-
tion.
A fresh specimen of Melinnopsis was dredged
in 3,020 m at lat 36°02'30"N, long 73°51'W by
RV Eastward (Eastward Station 2756, Septem-
ber 15, 1965). Unfortunately the specimen is now
missing but the following details show that it
was close to, if not conspecific with M. atlantica
which was found further north but at the same
abyssal depth.
The head, gills, and setation of segments
3-6 agree with M. atlantica but in the fresh
specimen it was possible to see that the dorsal
ridge across segment 6 was incised to form
16 triangular lobes. The thorax was complete
112
and had 13 segments with both notosetae and
uncini. The thoracic uncini had four or five
teeth above the rostral point. The abdomen was
broken at the 13th segment; the uncinigerous
pinnules were rectangular as in M. atlantica and
there were rudimentary notopodial lobes above
the first three uncinigerous pinnules.
In view of the fact that all the characters
which can be checked on the type material
agree with those recorded on the fresh speci-
men and that the only points of difference con-
cern features which are probably due to the
poor condition of the syntypes, it is proposed
that Melinnopsis atlantica be defined as having
a ridge across segment 6 divided into 16 tri-
angular lobes and possessing 13 uncinigerous
thoracic segments starting from segment 7.
As shown by Day (1964), the related genera
of the subfamily Melinninae with two groups
of four gills are Melinnexis Annekova, with 14
uncinigerous thoracic segments and Melinnides
Wesenberg-Lund, with 12. Both of these genera
have an enlarged tentacular process.
Records. — One specimen off Beaufort in
3,020 m (*).
Distribution. — Off Chesapeake Bay in 3,110
m.
Melinna maculata Webster, 1879
Melinna maculata Webster, 1879: 261, pi. 10:
Fig. 145-147. - Hartman, 1951: 108, pi. 27:
Fig. 1,2.
Melinna cristata.- Hartman, 1945: 43 (non Sars).
of mouth. Oblique lateral folds extending from
ventrum of segment 3 around sides of branchial
region and uniting with a transverse ridge
across segment 6. Margin of transverse ridge
smooth (Figure 15b). Two groups of four gills
on segment 3; gills in each group fused basal ly;
outer two gills smooth and cirriform, inner two
gills pinnate. Notosetae of branchial segments
include a stout hook on segment 4 (Figure 15c)
and small tufts of winged capillaries on seg-
ments 5 and 6. Neurosetae of branchial seg-
ments as fine acicular setae (Figure 15d) deeply
embedded on sides of segments 3, 4, and 5 but
not 6. Thirteen uncinigerous thoracic segments
starting from segment 7. About 32 abdominal
segments with square uncinigerous pinnules;
no vestigial notopodia. Thoracic uncini (Figure
15e, f) with a single vertical series of five or six
teeth above small rostrum and basal prow.
Records. — A single specimen from 20 m off
Beaufort (*). This is a new record for the United
States.
Distribution. — Brazil and tropical West
Africa; intertidal to 20 m.
Auchenoplax crinita Ehlers, 1887
Auchenoplax crinita Ehlers, 1887: 209, pi. 44:
Fig. 10-16. - Hartman, 1965a: 216, pi. 47:
Fig. a-d.
Records. — Off Beaufort in 600 m (*).
Distribution. — Massachusetts to South Amer-
ica and Morocco in 200-1,500 m.
Records. — Cape Hatteras to Beaufort, inter-
tidal (3, 11, 13, 18).
Distribution. — Virginia to the Gulf of Mexico;
intertidal to 10 m.
Isolda pulchella Muller, 1858
Figure 15a-f
Isolda pulchella. - Augener, 1918: 517, pi. 7:
Fig. 229, text Fig. 88. - Day, 1967: 691, Fig.
35.1. k-n.
Descriptio)!. — Body up to 45 mm long, pale
brown in color with barred tentacles. Prostom-
ium (Figure 15a) a curved hood pinched in
laterally. Numerous minute eyespots. Buccal
tentacles grooved and arise from a shelf in roof
Samythella eliasoni New Species
Figure 15g-m
Eclysippe vanelli. - Eliason, 1955: 10, Fig. 3 a-1.
- (No)i) Lysippe vanelli Fauvel, 1936: 96,
Fig. 13 a-d.
Holotype.— USNM 43145.
Description. — Holotype colorless in alcohol,
10 mm long, and encased in a friable mud tube.
Prostomium (Figure 15g) as a flattened triangu-
lar lobe overhanging mouth; no glandular pros-
tomial ridges but one pair of small eyes. Short
grooved buccal tentacles protruding from mouth.
Branchial ridge above segment 3 well developed
with right and left groups of three gills nar-
rowly separated in median line. Gills all very
113
Figure 15. — Isolda pulchella a, lateral view of anterior end; b, dorsal view of branchial region with gills cut short;
c, notopodial hook of segment 4; d, neuroseta of segment 3; e and f, edge-on and lateral views of thoracic uncinus.
Sdiiiythella elianoii n. sp. g, anterior end; h, thoracic capillary; i and j, edge-on and lateral view of thoracic
uncinus; k. palea; I. edge-on view of abdominal uncinus; m, posterior thoracic parapodium. Ampharete parvi-
dentata n. sp. n and o, edge-on and lateral views of thoracic uncinus; p, abdominal uncinigerous pinnule; q and r,
edge-on and lateral views of abdominal uncinus; s, anterior end. Ampharete americana n. sp. t, anterior end;
u and v, edge-on and lateral view of thoracic uncinus; w and x, edge-on and lateral view of abdominal uncinus.
114
long, outermost obviously tapered and more
slender than other two. Branchial segments
3-6, all with notosetae but without neurosetae.
Notosetae of segment 3 as long tapering paleae
( Figure 15k) reaching tip of prostomium. Noto-
setae of segment 4 as minute winged capil-
laries, those of segments 5 and 6 longer but
still shorter than those of uncinigerous seg-
ments. Twelve uncinigerous thoracic segments
from segment 7. Notosetae (Figure 15h) as
bilimbate capillaries; notopodial lobes of pos-
terior thoracic segments with a pair of small
cirriform projections (Figure 15m). Thoracic
uncini (Figure 15i, j) with two rows of five teeth
above rostral point and basal prow. Abdomen
of 13-14 segments bearing oval uncinigerous
pinnules without superior papillae. No vestigial
notopodia. Anal cirri not evident. Abdominal
uncini (Figure 151) with an irregular double
series of four teeth above rostral point.
Remarks. — The genus Samythella was erected
by Verrill (1873a) for S. elongata, which lacks
paleae, has 3 pairs of gills, 12 uncinigerous
thoracic segments, uncini with a single row of
teeth. As originally defined, Samythella was
closely related to Eusamytha Mcintosh and
Eclysippe Eliason. Samythella was redefined by
Day (1964) to include all three genera and in
the emended definition paleae may be present
or absent. The type species of Eclysippe is
Lysippe vanelli Fauvel, with paleae smaller
than the notosetae of the uncinigerous thoracic
segments. Fauvel's description left it uncertain
whether there were three or four pairs of gills.
Eliason's description of Eclysippe vanelli is
clear, and the species he examined is certainly
the same as the one described above from
Beaufort. I doubt, however, that it is the same
as Fauvel's Lysippe vanelli and, for this reason,
I have renamed it Samythella eliason i.
Records. — One specimen off Beaufort in
200 m (*).
Distribution. — Sweden; in 15 m.
Amage auricula Malmgren, 1866
Amage auricula. - Wollebaek, 1912: 65, pi. 13:
Fig. 1-6. - Hessle, 1917: 120. - Hartman,
1945: 44.
Distribution. — Arctic and cold North Pacific
and Atlantic south to Sweden and North Caro-
lina; in 0-200 m.
Amphicteis gunneri (Sars, 1835)
Amphicteis gunneri. - Hessle, 1917: 116, pi. 1:
Fig. 9. - Fauvel, 1927: 231, Fig. 80 a-k. -
Day, 1967: 695, Fig. 35.2. g-n.
Records. — Common off Beaufort in 40-120
m (21, *).
Distribution. — Cosmopolitan; from intertidal
to abyssal depths.
Ampharete acutifrons Grube, 1860
Ampharete acutifrons. - Hessle, 1917: 96. - Day,
1967: 699, Fig. 35.4. a-f. - Hartman, 1969:
537, Fig. 1-4.
Ampharete grubei. - Fauvel, 1927: 227, Fig. 79
a-p.
Description. — Body tapered, 10-35 mm long.
Prostomium as a bluntly triangular hood over
mouth; one pair of small eyespots. Buccal cirri
papillose. Two groups of four cirriform gills on
branchial ridge across segment 3. A bundle
of 12 large paleae on segment 3, and bundles
of small notopodial capillaries on segments 5
and 6. Segment 4 achaetous and partly fused
to segment 3. No neurosetae on segments 3 to
6. Twelve segments with winged capillaries in
notopodia and uncini in neuropodia from seg-
ment 7 onwards. About 12 abdominal segments
with a cirriform papilla at upper edge of each
uncinigerous pinnule. Pygidium with a pair of
anal cirri. Thoracic uncini with two vertical rows
of four or five teeth above a small rostrum and
basal prow.
Records. — Common off Beaufort in 20-160 m
(21, *).
Distribution. — Arctic; eastern Atlantic from
Norway to South Africa; California; 5-1,380 m.
Ampharete parvidentata New Species
Figure 15n-s
Ampharete n. sp. Day, Field, and Montgomery,
1971: 123.
Records. — Beaufort, in shallow dredgings
(11).
Holotype.— USNM 43143; 19 paratypes,
USNM 43144.
115
Description. — Holotype 12 mm long and en-
cased in a friable mud tube. Prostomium (Figure
15s) as a rectangular lobe above the extruded
buccal tentacles; no glandular ridges but two
pairs of eyespots. Buccal tentacles stout and
papillose. Branchial ridge on segment 3 well
developed with two groups of four gills sepa-
rated by a broad median gap. All gills long and
cylindrical, with outer ones markedly longer
than inner ones. Small capillary notosetae on
branchial segments 3, 5 and 6. Segment 4 fused
to segment 3 and achaetous. No neurosetae on
segments 3-6. Twelve segments with well-
developed notosetae and uncini starting from
segment 7. Abdomen with 12 uncinigerous seg-
ments without rudimentary notopodia but with
uncinigerous pinnules with well-marked super-
ior papillae (Figure 15p). Pygidium with one
pair of long anal cirri. Thoracic uncini (Figure
15n. o) with two vertical rows of three teeth
above a medial rostral point and blunt basal
prow. Abdominal uncini (Figure 15q, r) small,
with six or seven teeth in a double row.
Remarks. — The shape of the prostomium in
the Ampharetidae varies with the opening of
the mouth. In the holotype, the mouth is ex-
panded for the extrusion of the buccal cirri and
the prostomium is roughly square; in many of
the paratypes the mouth is closed and the pros-
tomium is bluntly pointed in front and pinched
in at the sides.
Most species of Ampharete have well-
developed notosetae forming large paleae on
segment 3; here they are very small and often
difficult to see at the base of the outermost gill.
Apart from this character, the most distinctive
features are the broad gap between right and
left groups of gills and the small number of
teeth on the thoracic uncini; hence the specific
name Ampharete parvidentata.
Records. — Fairly common off Beaufort in
35-120 m (21, *).
Ampharete amerieana New Species
Figure 15t-x
Holotype. — USNM 43141; seven paratypes,
USNM 43142
Description. — Body 10 mm long and encased
in a fragile mud tube. Prostomium (Figure 15t)
bluntly triangular and folded in laterally; one
pair of small lateral eyespots but no glandular
ridges. Buccal tentacles papillose (observed on
a paratype). Branchial ridge well marked with
two groups of four long, subequal, cirriform gills
narrowly separated in median line. Each group
of gills with three in line and one slightly pos-
terior in origin. A bundle of small notosetae on
segment 3 and bundles of slightly larger noto-
setae on segments 5 and 6. Segment 4 achaetous
and completely fused to segment 3. Twelve seg-
ments with well-developed notosetae and uncini
starting from segment 7. Abdomen with 12 un-
cinigerous segments without vestigial notopodia
and without superior papillae on the oval
uncinigerous tori. Thoracic notosetae as broad -
bladed bilimbate capillaries. Thoracic uncini
10 per row; individual uncini (Figure 15u, v)
with two vertical rows of five teeth above the
small median rostral point and blunt basal
prow. Abdominal uncini (Figure 15w, x) shorter
and broader than thoracic ones and with two
irregular rows of four teeth.
Remarks. — When first examined, these small
specimens were thought to be juveniles until
it was noticed that one 10-mm specimen con-
tained large eggs. The most characteristic fea-
ture is the small size of the notosetae on seg-
ment 3. The lack of superior papilla on the
abdominal pinnules is unusual in Ampharete
but these structures are often small and easily
overlooked.
Records. — Fifteen specimens off Beaufort
in 10-20 m (*).
FAMILY TEREBELLIDAE
Key to genera and species
1 Gills present on segments 2-4
1 Gills absent from segments 2-4. [Tentacular lobe large and
frilly. Median ventral glandular pads reduced and ventro-
lateral areas of neuropodia swollen. No tube]
13
116
2 Thoracic neurosetae as long-shafted hooks starting on seg-
ment 6. Tentacular lobe frilly 3
2' Thoracic neurosetae as avicular uncini starting on segment
5. Tentacular lobe collar-shaped 4
3 A single median gill with four partly fused lamellated lobes
(Tereb <e Hides). Eighteen bundles of notosetae T. stroemii
3' Two or three pairs of simple cirriform gills (Trichobranchus) .
Fifteen bundles of notosetae. [Three pairs of gills] T. glacialis
4 Gills as numerous simple filaments (Thelepus). [Gill filaments
on three segments. Base on uncini produced forwards with
a terminal attachment button and a small notch below it] T. setosus
4' Gills branched 5
5 Notosetae with denticulate tips. Lateral lobes either present
or absent on segments 2-4. Over 20 segments with notosetae 6
5' Notosetae with smooth tips. Lateral lobes present on seg-
ments 2-4. About 17-18 segments with notosetae 9
6 Gills with short basal stumps and long terminal branches;
notosetae with short denticulate tips (Amphitrite). Small
lateral lobes on segments 1, 2, and 3. [Over 35 segments
with notosetae] A. ornata
6' Gills with well-developed trunks and short terminal branches.
Notosetae with long denticulate tips in posterior segments.
No lateral lobes on segments 2-4 (Terebella) 7
7 Two pairs of gills. Only posterior third of body without noto-
setae T. pterochaeta
7' Three pairs of gills. Notosetae continue to near end of
abdomen 8
8 Posterior notosetae with a spur preceding the denticulate
blade T. lapidaria
8' Posterior notosetae without a spur preceding the denticulate
blade T. rubra
9 Two pairs of gills (one often missing). Uncini avicular, those
of first row often with long basal shafts (Pista) 10
9' Three pairs of gills. Uncini pectinate, with a single vertical
series of teeth and lack basal shafts (Loimia) 12
10 Gill filaments spirally branched, forming a compact tuft or
"pom-pon" at end of trunk. [Uncini of first row with long,
narrow shafts] Pista eristata
10' Gill filaments dendritically branched 11
11 Second pair of lateral lobes divided to form two tapered pro-
jections. Uncini of first row without long necks Pista palmata
11' Second pair of lateral lobes not divided. Uncini of first row
with long necks and tapered shafts Pista quadrilobata
117
12 Uncini with 5-6 teeth. Tubes constructed of sand and/or shells Loimia medusa
12 Uncini with 6-8 teeth. Tubes constructed of mud and sand Loimia viridis
13 Notopodia of middle segments vascular and divided to form a
tuft of branchial filaments with minute spinulose capillaries
at their tips (Enoplobranchus) E. sanguineus
13' Notopodia never vascular or divided 14
14 No uncini even on posterior neuropodia and ventrum of thorax
diffusely glandular. Notosetae present on thoracic segments
at least 15
14' Uncini present from 7th or more posterior segment. No achae-
tous middle region and capillary notosetae present from
segment 2 (Polycirrus) 17
15 No achaetous middle region. Notosetae restricted to 6-12
anterior segments (Lysilla). [Notosetae minute and their
distribution uncertain] L. alba
15' An achaetous middle region. Notosetae as capillaries on 9-13
anterior segments and acicular after the achaetous middle
region (Amaeana) 16
16 Capillary notosetae all with smooth wings. Body purple A. trilobata
16' Capillary notosetae including winged forms and others with
plumose blades like an ear of wheat. Body pale A. accraensis
17 Notosetae including plumose forms with blades formed of a
series of overlapping cones as well as larger, smooth-
winged forms P. carolinensis
17' Notosetae not including plumose forms 18>
18 Notosetae on 16-20 segments, all with minutely spinulose
blades. Three large pairs of nephridia P. eximius
18' Notosetae on 25-32 segments, all with smooth to faintly hispid
blades. Six large pairs of nephridia P. eximius dubius
Terebellides stroemii Sars, 1835 Trichobranchus glacialis Malmgren, 1866
Terebellides stroemi. - Fauvel, 1927: 291, Fig. Trichobranchus glacialis. - Fauvel, 1927: 288,
100 i-q. - Day, 1967: 713, Fig. 36.1. f-j. Fig. 100 a-h. - Miner, 1950: 350, pi. 113. -
Terebellides stroemii. - Hartman, 1969: 653, Day, 1967: 711, Fig. 36.1. a-e.
Fig. 1-7.
Records. — Off Beaufort in 160-200 m (*).
Records. — Off Beaufort in 40-200 m (18, 21, Distribution. — Cosmopolitan; rarely inter-
*). tidal but common in deeper water down to 200 m.
Distribution. — Cosmopolitan from the Arctic
to the Subantarctic ; occasionally intertidal but „, , . ,A , ,0,>-^
„ . , ,, , . A„„ Thelepus setosus (Quatrefages, 1865)
usually at depths down to 4,000 m. r
Thelepus setosus. - Fauvel, 1927: 273, Fig. 95
a-h. - Hartman, 1951: 113; 1969: 649, Fig.
1-6. - Day, 1967: 729, Fig. 36.6. a.
118
Record*. — Cape Hatteras to Beaufort, inter-
tidal (15, 18,20).
Distribution. — Cosmopolitan in all warm tem-
perate areas; intertidal to about 100 m.
Amphitrite ornata (Leidy, 1855)
Amphitrite ornata. - Verrill, 1873a: 320, pi. 16:
Fig. 82. - Hartman, 1945: 44.
Records. — Cape Hatteras to Beaufort Sound,
intertidal to 18 m (3, 5, 7, 8, 11, 15, 18, 20).
Distribution. — Massachusetts to North Caro-
lina; intertidal to a few meters in soft mud.
Terebella pterochaeta Schmarda, 1861
Terebella bruneo-comata Ehlers, 1887: 237, pi.
51: Fig. 1-5.
Amphitritides bruneo-comata. - Pearse and Wil-
liams, 1951: 139. - Hartman, 1959: 499.
Terebella pterochaeta. - Day, 1967: 747, Fig.
36.10. a-f.
Remarks. — Terebella bruneo-comata has
been well described and illustrated by Ehlers
(1887) and his description agrees in detail with
the description of Terebella pterochaeta
Schmarda, given by Day (1967). Both have 2
pairs of branched gills, notosetae with narrow
wings and denticulate tips, and 16 ventral pads.
T. bruneo-comata has 27 segments with noto-
setae and uncini with three arcs of close-set
denticles while T. pterochaeta has 28-33 seg-
ments with notosetae and uncini with a crest of
close-set denticles. Both characters are variable
in the genus and the number of arcs of denticles,
which are always irregular and integrated, is
never clear. The two appear conspecific and
Schmarda's name has priority. As Hessle (1917)
has defined the genus Terebella with two or
three pairs of branching gills, I see no point in
referring this species to Amphitritides Augener.
Records. — On shallow reefs off North Caro-
lina (14).
Distributio)). — South Africa; tropical Indo-
Pacific from the Red Sea to Indo-China; ? West
Africa; North Carolina to Florida; intertidal to
50 m.
Terebella lapidaria Linnaeus, 1767
Terebella lapidaria. - Fauvel, 1927: 254, Fig. 87
f-1.
Records. — Cape Hatteras area, intertidal (18).
Distribution. — Warm North Atlantic from the
English Channel to Morocco; Mediterranean;
Massachusetts to North Carolina; intertidal to
30 m.
Terebella rubra (Verrill, 1873) (Homonym)
Lepraea rubra Verrill, 1873a: 321 (non Terebella
rubra Risso, 1826).
Terebella rubra. -Hartman, 1945: 44; 1951: 112.-
McCloskey, 1970: 28.
Remarks. — Hartman, (1959a) reported that
Terebella rubra (Verrill) is a junior homonym.
I have not seen a specimen but the published
descriptions state that it has three pairs of gills,
that the notosetae continue to near the end of
the abdomen, and that individual notosetae do
not possess a spur at the base of the denticu-
lated blade. These characters suggest that
Verrill's species must be close to T. ehrenbergi
Grube from the Indo-Pacific and T. schmardaei
Day from South Africa. Dr. Pettibone has sug-
gested that the type be examined before it is
renamed.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 7 m (3, 5, 11, 13, 15, 18, 20).
Distribution. — Massachusetts to North Caro-
lina; intertidal to a few meters.
Pista cristata (Muller, 1776)
Pista cristata. - Fauvel, 1927: 266, Fig. 93 a-g. -
Hartman, 1945: 44; 1951: 113; 1969: 615,
Fig. 1-3. - Day, 1967: 738, Fig. 36.7. h-j.
Recoi-ds. — North Carolina, intertidal to 20 m
(3, 11, 13,21, *).
Distribution. — Arctic and throughout the At-
lantic to South Africa; Mediterranean; North
Pacific; intertidal to 200 m.
Pista palmata (Verrill, 1873)
Scionopsis palmata Verrill, 1873a: 614, pi. 11:
Fig. 3. - Miner, 1950: 349, pi. 113.
Pista palmata. - Hartman, 1951: 112.
119
Ri cords. — Cape Hatteras to Beaufort Sound;
intertidal to 7 m (3, 11. 13. 15, 18. 20).
Distribution. — Massachusetts to Florida and
the Gulf of Mexico: intertidal to a few meters.
Pista quadrilobata (Augener, 1918)
Figure 16a-c
Nicolea quadrilobata. - Augener. 1918: 532, pi.
6: Fig. 183. pi. 7: Fig. 226-227. text Fig. 90.
Pista quadrilobata. - Day 1967: 740, Fig. 36.8.
a-e.
Description. — Tentacular lobe collar-shaped,
with orange tentacles and numerous eyespots.
Buccal segment (Figure 16a) with large, wing-
like lateral lobes united basally to form a sheath
at base of tentacles; second segment short
with rudimentary ventrolateral lobes; third seg-
ment with large lateral lobes. Two pairs of
dendritically branched gills (one gill often miss-
ing). Smooth-tipped winged capillaries on 18
segments starting from segment 4. Rows of
avicular uncini from segment 5. Uncini of first
and second rows (Figure 16b, c) with a long
neck below rostrum and base extending back
as a short, tapered shaft; uncini of subsequent
segments with shorter necks and without shafts.
Remarks. — This species is close to Pista
pahnata but the latter is reported to have the
lateral lobes of segment 3 divided to form
tapered projections and the uncini of the first
row differ in shape.
Records. — Beaufort Sound between tide
marks and on coral in 18 m off Beaufort (20, *).
This is a new record for the United States.
Distribution. — South and South West Africa;
intertidal to 20 m.
Loitnia medusa (Savigny, 1818)
Loimia turgida. - Andrews, 1891a: 298, pi. 18:
Fig. 46-49.
Lunula medusa. - Hartman, 1945: 46, pi. 10:
Fig. 2, 3; 1951: 111; 1969: 601, Fig. 1-3. -
Day, 1967: 743, Fig. 36.9. a-e.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 20 m (5, 7, 11, 13, 15, 18, *).
Distribution. — In warm to tropical waters of
all oceans from 0 to 100 m.
Loimia viridis Moore, 1903
Loimia viridis Moore, 1903: 723, Fig. 11-14. -
Hartman, 1945: 46, pi. 10: Fig. 4, 5; 1951:
111.
Remarks. — This species is closely related to
L. medusa if not merely a variety of it. According
to Hartman (1945) "L. viridis differs from
L. medusa [see above] since it constructs mud-
covered tubes instead of coarse shell-covered
ones; ventral thoracic gland shields are notably
broader and thicker and uncini have teeth more
closely spaced." The uncini have six or seven
(or eight) teeth whereas there are four to six
in L. medusa.
Recoi'ds. — Cape Hatteras to Beaufort, inter-
tidal (11, 13, 18).
Distribution. — Massachusetts to the Gulf of
Mexico; intertidal.
Enoplobranchus sanguineus (Verrill, 1873)
Chaetobranchus sanguineus. - Verrill, 1873a:
616.
Enoplobranchus sanguineus. - Hartman. 1942a:
75, pi. 113; 1944c: pi. 54: Fig. 8; 1945: 47.
Description. — Body markedly tapered, 100-
350 mm long, red when alive. Tentacular lobe
large and frilly, bearing numerous long con-
tractile tentacles. Normal gills absent but noto-
podia branched and vascular from about seg-
ment 9. Vascular branches with small spinulose
capillaries at their ends. Posterior notopodia
not vascular and without notosetae. Neurosetae
absent. Ventrum glandular and midventral
pads greatly reduced. No tube.
Records. — Cape Hatteras to Beaufort on
intertidal mudbanks and on coral at 6.5 m (3,
11, 13, 18,20, *).
Distribution. — Gulf of St. Lawrence to North
Carolina; intertidal to 6.5 m.
Lysilla alba Webster, 1879
Lysilla alba Webster, 1879: 63, pi. 10: Fig. 148. -
Hartman, 1945: 47.
Records. — Beaufort, on sheltered shores (3,
11).
120
Figure 16. — Pista quadrilobata a, anterior end; b and c, edg'e-on and lateral views of uncinus from first row. Amaeana
trilobata d, thoracic capillary seta; e, abdominal acicular seta; f, ventrolateral view of entire worm. Amaeana
accraensis g, plumose capillary seta. Poly cirrus eximius h, ventral view of anterior end; i and j, edge-on and lateral
view of uncinus; k and 1, longer and shorter capillary setae. Polycirrus carolinensis n. sp. m and n, edge-on and
lateral views of uncinus; o, longer, smooth-bladed capillary; p, shorter pulmose capillary. Polycirrus eximius dubius
n. subsp. q, profile of uncinus.
121
Distribution. — Massachusetts to North Caro-
lina; intertidal.
Amaeana trilobata (Sars, 1863)
Figure 16d-f
Amaea trilobata. - Fauvel, 1927: 285, Fig. 99a-e.
Amaeana trilobata. - Day, 1967: 718, Fig. 36.3.
e-h.
Description. — Body purple when fresh, up to
20 mm long, swollen anteriorly, slender pos-
teriorly. Tentacular lobe large, folded, often
trefoil-shaped, bearing numerous short tentacles.
No eyespots. No branchiae. Notosetae from
segment 3, and setae sometimes retracted into
notopodial lobes. First 10-11 setigers with
small smooth-winged capillaries (Figure 16d)
followed by 5 achaetous segments at junction
of thorax and abdomen; finally about 30 ab-
dominal segments bearing a few blunt acicular
setae (Figure 16e) in notopodia. Neuropodia
and neurosetae entirely absent. Ventrum of
thorax (Figure 16f) swollen, glandular and
tessellated, with a median groove concealing
a row of nine poorly defined ventral pads.
Records. — Common off Beaufort in 80-200 m
(21.*).
Distribution. — Arctic; Norway; New Eng-
land; Mediterranean; South Africa; Japan; 50-
500 m.
Amaeana accraensis (Augener, 1918)
Fi giire 16f>'
Amaea accraensis. - Augener, 1918: 561, pi. 7;
Fig. 246. text Fig. 98. - Kirkegaard, 1959:
89, Fig. 22.
Description. — Generally similar to A. trilo-
bata described above, with following exceptions:
Body pale not purple; 11-13 thoracic segments
with notosetae, not 10-11; notosetae of two
types, longer ones with smooth narrow wings
resembling those of A. trilobata and shorter
ones (Figure 16g) with completely spinulose
blades.
Remarks. — All 11 specimens collected were
broken between the 9th and 11th setigerous
segments, and no abdominal segments with
acicular notosetae were obtained.
Records.— Off Beaufort in 80-160 m (21, *).
This is a new record for the United States.
Distribution. — Ghana; ? South Africa; inter-
tidal.
Polycirrus carolinensis New Species
Figure 16m-p
Polycirrus n. sp. - McCloskey, 1970: 26 (record
only).
Holotype. — USNM 43122; four paratypes,
USNM 43123.
Description. — Body uniformly pale in alcohol,
30 mm long for 80 segments, swollen anteriorly
and constricted between setigers 10 and 15.
Tentacular lobe large, irregularly folded, roughly
trefoil-shaped, with numerous contractile ten-
tacles. Notosetae from segment 2 and present
on 33 segments in holotype (paratypes with
25, 27, 38, and 48 segments with notosetae).
Notopodia without terminal free lobes. Neuro-
setae from 7th setigerous segment. Mouth with
lower lip as a large ventral cushion. Ventral
surface of thorax from segment 2 to 10 with
glandular swellings. Ventral pads small and
sunken in a midventral groove. Large nephridial
papillae below notopodia of setigers 4, 5, and
6 and a doubtful one on setiger 3. Five or six
pairs of large nephridia revealed by dissection.
Notosetae of two types; four or five longer
capillaries with smooth blades (Figure 16o) and
three or four shorter capillaries with plumose
blades (Figure 16p) formed by a series of over-
lapping cones on a slender axis. Neuropodial
uncini small and arranged in a single row. In-
dividual uncini (Figure 16m, n) with one large
tooth and an arc of 8-10 small teeth above main
fang; formula: MF: 1 : 8-10. Base of uncinus
striated.
Remarks. — The type specimens were col-
lected off Beaufort in 20 m on sand; Dr. Mc-
Closkey's specimens were collected off Beaufort
on coral in 18.3 m. The plumose setae of P.
carolinensis shows that it is related to P. plnmo-
sus Wollebaek, P. medius Hessle, and P. coc-
ci nea Grube but it differs from them in the num-
ber of segments bearing notosetae, the seg-
ment on which the first row of uncini occurs,
and in the shape of the uncini themselves.
Records. — On coral and sand in 18.3-20 m
off Beaufort (20, *).
122
Polycirrus eximius (Leidy, 1855)
Figure 16h-l
Torquea eximia Leidy, 1855: 146, pi. 11; Fig.
51, 52.
Material examined. — Three specimens iden-
tified by Verrill in 1881 from Station 1003 in
Vineyard Sound, Mass. (USNM 8165). Six speci-
mens dredged off Beaufort in 5-80 m.
Description. — Body up to 9 mm long for 52
segments. Tentacular lobe (Figure 16h) trefoil-
shaped, with folded margins bearing numerous
tentacles. No gills. Notosetae on 16-18 seg-
ments starting from second and all notopodia
with terminal fleshy tags. Notosetae of two
lengths (Figure 16k, 1); both types bilimbate
with minutely spinulose or hispid blades. Ven-
trum with a narrow medial groove separating
10 glandular ventrolateral swellings. Setigers
1-3 with obvious nephridial papillae; three pairs
of large nephridia revealed by dissection. Uncini
from setiger 7 ( = segment 8). Individual uncini
(Figure 16i, j) with a cap of about seven long
teeth above main fang.
Remarks. — The description given above is
based on the fresh specimens dredged off
Beaufort. The three specimens identified by
Verrill are similar apart from the following de-
tails. Notosetae are present on segments 18-20
instead of segments 16-18. Nephridial papillae
were seen on setigers 1-6 instead of setigers
1-3 but dissection still revealed only three
large pairs of nephridia although small ones
may be present on the other three segments.
The uncini were generally similar but the details
were clearer; above the main fang there was a
median tooth surmounted by an arc of six to
eight teeth giving the formula MF : 1 : 6-8.
Leidy's type of Torquaea eximia has been
lost and his description is inadequate. He com-
ments on the red color of the body and the
numerous extensible tentacles. The length of
the worm was half an inch (? 12 mm) for 40-60
segments. The number of segments with noto-
setae was not stated but it was noted that uncini
commence on segment 8, i.e., setiger 7. The
figures of the notosetae and uncini are too small
to show specific characters.
To summarize, it may be stated that the
characters of the North Carolina specimens
agree with the characters of specimens identi-
fied by Verrill and are consistent with Leidy's
original description as far as it goes. It is worth
noting too, that the above description of P.
eximius agrees with that given by Fauvel (1927),
for Polycirrus tenuisetis Langerhans from Ma-
deira, with the exception that P. eximius is red
in life while P. tenuisetis is colorless (? in life).
Later descriptions of P. eximius by Verrill
and Smith (1874) and by Miner (1950) state
that there are more segments with notosetae
and that the uncini start after setiger 7. Possibly
these descriptions refer to the different form
described below.
Records.— Off Beaufort in 5-80 m (20, 21,
*). The intertidal records by Webster (1879),
Andrews (1891a), Hartman (1945), and Wells
and Gray (1964) do not give diagnostic charac-
ters and are thus doubtful.
Distribution. — Massachusetts to North Caro-
lina; ? Madeira; intertidal to 80 m on sandy mud.
Polycirrus eximius dubius New Subspecies
Figure 16q
Polycirrus eximius. - Verrill, 1873a: 616, pi. 16:
Fig. 85. - McCloskey, 1970: 26.
Holotype.— USNM 43130.
Description. — Body pink in alcohol when
fresh, about 20 mm long for 80 segments. Ten-
tacular lobe large and frilly with numerous
tentacles. No gills. Notosetae on 31 segments
starting from segment 2; individual notopodia
with terminal fleshy tags. Nephridial papillae
obvious below notopodia of setigers 1-6, the
first three larger than others. Six pairs of large
nephridia revealed by dissection. Ventrum with
a median groove separating 10 pairs of ventro-
lateral glandular swellings. A short row of
uncini on setiger 9 and longer rows on subse-
quent segments. Notosetae of two lengths;
longer ones with broader blades and shorter
ones with narrower blades; both types with
blades apparently striated under low magnifica-
tion but finely spinulose when seen under oil-
immersion. Uncini (Figure 16q) with an arc of
about five long teeth above main fang.
Remarks. — The holotype was found on coral
at 18 m off Beaufort by Dr. L. McCloskey. This
subspecies has setae very similar to those of
P. eximius Leidy described above but there are
more segments with notosetae, the uncini first
appear on setiger 9 and there are six large
123
pairs of nephridia not three large and the rest
small. Verrill (1873a) describes P. cximius as
having 25 segments with notosetae and uncini
from setiger 8 and this description is repeated
by Miner (1950). It is possible that P. eximius
is a very variable species with 16-31 segments
bearing notosetae, uncini commencing on seti-
ger 7-9 and three to six pairs of large nephridia.
However this range of variation is wider than is
usual in the genus Polycimis and until more
specimens have been studied in detail it is
better to treat these larger specimens with
25-31 bundles of notosetae as a separate sub-
species.
Records. — On coral at 18 m off Beaufort
(20, *).
Distribution. — Probably New England to
North Carolina but records confused.
FAMILY SABELLIDAE
Key to genera and species
1 Thoracic neurosetae as a row of avicular uncini and often a
row of pickaxe setae as well. Radioles usually with eyespots
1' Thoracic neurosetae as a row of long-shafted hooks. Radioles
without eyespots
2 A row of pickaxe setae as well as avicular uncini on thorax.
No stylodes on radioles
2' No pickaxe setae. Radioles with paired stylodes externally
(Branchiomma). [Uncini with two arcs of denticles. Stylodes
slender. Dark flecks on body]
3 Collar setae in compact bundles and consist of long-winged
capillaries
3' Collar setae in long double rows and consist of stout capil-
laries with short blades just projecting from body (Hypsi-
comus)
4 Collar produced ventrally as a sheath to branchial lobes (H.
circumspiciens)
4' Collar uniformly short, not produced ventrally
5 Thoracic notosetae include both winged capillaries and paleae
with very short broad blades. [Dorsal radioles with eye-
spots halfway along (Pota nulla)]
5' Thoracic notosetae as winged capillaries with broader or
narrower wings; no paleae
6 Collar prolonged ventrally forming a basal sheath for branchial
lobes
6' Collar not prolonged ventrally
7 Two or more radioles, each with one large subterminal eye
(Megalomma)
7' Dorsal radioles with a series of small lateral eyespots (Sabella) .
Subterminal eyes on most radioles. [Collar not notched dorso-
lateral^]
8' Subterminal eye restricted to two dorsal radioles
2
10
B. nigromaculata
No N.C. record
H. phaeotaeiiia
P. spathiferus
P. reniformis
M. lobiferum
M. bioculatum
124
9 Eyespots paired and regularly arranged along radioles
9' Eyespots irregularly scattered along radioles
10 Abdominal segments with rows of minute uneini almost en-
circling body. Thoracic notosetae all winged capillaries.
[Radioles united by a web (Myxicola)]
10' Abdominal segments with short compact rows of uneini. Thora-
cic notosetae include both winged capillaries and paleae . .
11 Radioles without lateral flanges and always separate. Abdomi-
nal notosetae avicular, with broad tails
11' Radioles flanged, sometimes partly united. Abdominal noto-
setae quadrangular, without tails
12 Radioles 3 pairs, not united by web. (Oriopsis)
12' Radioles 6-10 pairs, united by web for half their length or more
(Chone). [Collar not notched ventrally. Tips of radioles
tapered. Juveniles with a caudal filament]
S. melanostigma
S. microphthalma
No N.C. record
11
Jasmineira bilobata
12
Oriopsis sp.
C. americana
Branchiomma nigromaculata (Baird, 1865)
Branchiomma nigromaculata. - Hartman, 1951:
114. - Day, 1967: 770, Fig. 37.4. m-p.
Records. — Off Beaufort, intertidal to 20m (13).
Distribution. — North Carolina to the West In-
dies; Cape Verde Islands; South Africa; Red
Sea; tropical Indian Ocean; Japan; intertidal
to about 50 m.
Hypsicomus phaeotaenia (Schmarda, 1861)
Protulides elegans Webster 1884: 325, pi. 11:
Fig. 63-74.
Hypsicomus torquatus. - Hartman, 1945: 47.
Hypsicomus elegans. - Hartman, 1951: 115.
Hypsicomus phaeotaenia. - Day, 1967: 761,
Fig. 37.2. i-n.
Re ma rks. — Webster's original description and
figures of Protulides elegans agree very closely
with specimens of Hypsicomus phaeotaenia
from the Indian Ocean examined by me and with
the present specimen from Beaufort. The name
H. phaeotaenia Schmarda has priority. Hartman
(1945) reported a specimen from Beaufort as
H. torquatus (Grube) and agreed with Augener
(1922) that Protulides elegans Webster was a
synonym. Since I have not seen Grube's original
description I cannot comment on this synonymy
but I do not agree with Hartman (1951) that
H. circumspiciens Ehlers is a synonym of Pro-
tulides elegans Webster and thus a synonym of
H. phaeotaenia. Ehlers' description and figures
are very clear and the ventrally elongated collar
of his H. circumspeciens is quite different from
the short collar of H. phaeotaenia.
Records. — Cape Hatteras to South Carolina;
intertidal to 20 m (5, 11, 13, 15, 18, 20, *).
Distribution. — North Carolina to the West
Indies and the Gulf of Mexico; circumtropical;
intertidal to 30 m.
Potamilla (Potamethus) spathiferus
(Ehlers, 1887) New Combination
Potamis spathiferus. - Ehlers, 1887: 278, pi. 54:
Fig. 7-11, pi. 55: Fig. 1-4.
Description. — Body very slender, up to 48 mm
long and encased in a sandy tube. No color
markings. Branchial lobes long and slanting,
each bearing a semicircle of six to nine separate,
elongated radioles ending in very long tapered
filaments. Collar with four lobes all projecting
forwards; paired dorsal lobes smaller and
narrower; paired ventrolateral lobes slanting,
their ventral ends forming a sheath supporting
the branchial base. Thorax with eight setigerous
segments. Notosetae of setigers 2-8 include
two or three winged capillaries and three to
five paleae with oval blades and small tapered
tips. Neurosetae include a row of pickaxe setae
125
with delicate symmetrical blades tapering to
tine tips and a row of avicular uncini with
striated crests, long arched necks and rather
long shafts. Abdomen with about 70 segments;
pygidium blunt. Abdominal notosetae as avicu-
lar uncini similar to those on thorax. Abdominal
neurosetae as small winged capillaries with
blades broad basal ly tapering to long slender
tips.
Remarks. — As noted above, only one type of
abdominal neuroseta was found in the speci-
mens from Beaufort, namely capillaries with
wings broad at the base and then narrowing to
very long slender tips. These may be inter-
preted as modified paleae or winged capillaries.
When partially retracted, only the long slender
tip shows like a narrow-winged capillary, and
it is possible that this is the reason why Ehlers
describes (but does not figure) two types of
abdominal neurosetae. Ehlers referred his speci-
mens to Potamis but as this was preoccupied
in the Lepidoptera, Chamberlin (1919) proposed
a new name Potamethus. This appears so close
to Potamilla that I have accorded it the rank of
a subgenus.
Records. — Three specimens off Beaufort in
18 m on coral heads (20, *).
Distribution. — Florida and West Indies; in
10 to 500 m.
Potamilla reniformis (Leuckart, 1849)
Potamilla oculifera Verrill, 1873a: 322, pi. 17:
Fig. 86.
Potamilla tortuosa Webster, 1879: 265, pi. 10:
Fig. 149-153.
Put mn ilia reniformis. - Fauvel, 1927; 309, Fig.
107 a-1. - Johansson, 1927: 142. - Day, 1967:
764, Fig. 37.3. a-f.
Pseudopotamilla reniformis. - Hartman, 1945:
47; 1959a: 557.
Potamilla (Pseudopotamilla) oculifera. - Miner,
1950: 119.
Remarks. — I agree with Johansson that Pseu-
dopotamilla Bush is not separable from Pota-
milla Malmgren.
Records. — Cape Hatteras to Beaufort, inter-
tidal to a few meters (3, 11, 15, 18).
Distribution. — Cosmopolitan; intertidal to
LOO m.
Megalomma lobiferum (Ehlers, 1887)
Brancliiomma lobiferum Ehlers, 1887: 254, pi.
53: Fig. 10-15'.
Megalomma lobiferum. - Hartman, 1951: 115.
Remarks. — This species is close to M. vesicu-
losu m (Montagu) but differs in having a straight
unnotched collar and pickaxe setae with sym-
metrical blades.
Records. — On shallow reefs off North Caro-
lina (14).
Distribution. — North Carolina and Florida;
intertidal to 4 m.
Megalomma bioculatum (Ehlers, 1887)
Branchiomma bioculatum Ehlers, 1887: 260, pi.
53: Fig. 1-9.
Megalomma bioculatum. - Hartman, 1951: 115.
Description. — Body up to 73 mm long with
brown bars across radioles and sometimes
brown marks on ventral surface of thorax. Tube
covered with sand or gravel. Branchial lobes
short, each with a semicircle of about 16 radioles.
Palps short, broad, faintly spiral. Large sub-
terminal eyes on dorsal pair of radioles pre-
ceding the very short naked tips. No eyes on
other radioles. Collar low, widely divided dor-
sally, not notched dorsolateral^ but slanting
forward and forming paired triangular ventral
lappets. Thorax of eight setigers with broad-
winged capillaries of varying length in noto-
podia. Notosetae including an anterior row of
pickaxe setae with plain delicate blades taper-
ing symmetrically to fine tips and a posterior
row of avicular uncini. Individual uncini with
striated crests, short necks, and elongated
bases. Abdomen with notosetae similar to avicu-
lar uncini of thorax; neurosetae as broad-winged
capillaries.
Records.— Off Beaufort in 18-40 m (20, 21,
*).
Distribution. — Florida and the Gulf of Mexico;
in 20-200 m.
Sabella melanostigma Schmarda, 1861
Sabella melanostigma. - Ehlers 1887: 263. -
Johansson, 1927: 121, text fig. 15.
126
Records. — Cape Hatteras to South Carolina,
intertidal to about 50 m (14, 15, 18, 19).
Distribution. — North Carolina, Bermuda, and
the West Indies; intertidal to 50 m.
Sabella microphthalma Verrill, 1873
Sabella microphthalma. - Hartman, 1942a: 80;
1945: 47; 1951: 117.
Records. — Cape Hatteras to South Carolina,
intertidal to 40 m (2, 3, 5, 7, 11, 13, 14, 15, 18,
20, *).
Distribution. — Massachusetts to Florida and
the Gulf of Mexico; intertidal to 40 m.
Jasmineira bilobata New Species
FiRiire 17m-v
Holotype. — USNM 43134; one paratype,
USNM 43135.
Description. — Holotype incomplete, very slen-
der, measuring 9 mm by 0.5 mm for 19 seg-
ments. Branchial lobes (Figure 17t) long and
slanting, each with eight radioles united only
at base. Each radiole with paired pinnules in-
creasing in length distally, tip slender, hardly
distinguishable from terminal pinnules. Ventral-
most radiole much shorter than others with
only two or three stumpy pinnules basally.
Palps short and grooved. Lips with baggy ven-
tral pouches (Figure 17s) partly covered by
notched ventral projections of collar. Collar
with two closely apposed dorsal lobes (Figure
17v), low lateral parts and ventral lobes pro-
duced forwards as notched lappets; dorsolateral
notches between dorsal lobes and low lateral
parts of collar. Thorax with eight setigers; first
short and united to collar, remainder slightly
broader than long. Setiger 1 with winged capil-
laries only; setigers 2-8 with one or two winged
capillaries (Figure 17p) and four paleae (Figure
17o) having oval blades and short, finely pointed
tips. Thoracic neurosetae as a row of five long-
shafted hooks (Figure 17m, n) with a swelling
at origin of shaft and a rostrum capped with a
close-set series of denticles and a faint indica-
tion of a hood.
Abdomen broken after segment 9. Abdominal
neurosetae include three winged capillaries
(Figure 17q) and shorter forms with hastate
blades embedded in the flesh. Abdominal
notosetae as avicular uncini (Figure 17u, v)
with four or five close-set arcs of denticles
above rostrum and short, laterally compressed
bases.
Remarks. — The shape of the collar immedi-
ately distinguishes J. bilobata from the well-
known European species J. elegans Saint-
Joseph, J. caudata Langerhans, and J. candela
(Grube). Possibly the most closely related spe-
cies is J.filiformis Hartman (1965a) from deep
water off Brazil. It too, has a collar with elon-
gated ventral lobes and thoracic hooks with a
swelling at the origin of the shaft. Hartman's
description is very brief but her figure does not
show dorsal lobes to the collar or notches in
the elongated ventral lobes. Moreover there are
only four pairs of radioles and there is no men-
tion of paleae among the thoracic notosetae.
Records. — Two specimens off Beaufort in
123 m (*).
? Oriopsis sp.
Description. — Length 3.5 mm including bran-
chiae; 11 thoracic and 19 abdominal segments.
Three pairs of flanged radioles. Collar low dor-
sally, widening laterally and forming two united
triangular lobes ventrally; margin of collar
smooth. Thoracic notopodia with three winged
capillaries and two paleae bearing oval blades
and fine tapering tips. Thoracic neurosetae as a
row of four or five long-shafted hooks each with
one large tooth and a close-set cap of small
denticles above rostrum. Abdominal uncini
almost square and arranged in rows of six or
seven per notopodium. Individual uncini with a
recurved basal prow and main fang surmounted
by three or four rows of smaller teeth with four
teeth per row.
Remarks. — The presence of paleae among
the thoracic notosetae and the numerous ab-
dominal segments are unusual in the genus
Oriopsis. Oridia crenicollis Annenkova, shares
these characters and is doubtfully referred to
Oriopsis by Banse (1957). It differs from the
Beaufort material in having a scalloped edge
to the collar.
Records.— Off Beaufort in 40-160 m (*).
127
Figure 17. — Chone americana n. sp. a, dorsal view of entire worm; a', posterior end of juvenile; b, ventral view of collar
and lips; c and d, lateral and edge-on views of thoracic hooks; e and f, edge-on and lateral view of abdominal
uncini; g, thoracic winged capillary; h, thoracic palea; j, thoracic tapered capillary; k, winged capillary from
anterior abdomen; 1, slender capillary from posterior abdomen. Jasmineira bilobata n. sp. m and n, lateral and
edge-on views of thoracic hook; o, thoracic palea; p, thoracic winged capillary; q, abdominal capillary; r, dorsal
view of collar; s, ventral view of collar overlying ventral lips; t, lateral view of worm; u and v, edge-on and
lateral views of abdominal uncinus.
128
Chone americana New Species
Figure 17a-l
Chone n. sp. - Day, Field, and Montgomery,
1971: 123.
Holotype.— USNM 43134; 50 paratypes,
USNM 43135.
Description. — Holotype (Figure 17a), uni-
formly pale in alcohol, 26 mm long including
branchiae; 8 thoracic and 43 abdominal setigers.
Branchial lobes semicircular, each with 10
radioles united by a web for half their length.
Tips of radioles very long and tapered. Ventral-
most radioles short with few pinnules or none.
Palps short, triangular, abruptly tapered. Lips
(Figure 17b) fused ventrally to form a notched
triangle. Collar narrowly divided dorsally, very
low and smooth-edged laterally, slightly wider
ventrally but not notched or divided in mid-
ventral line. Thorax of eight biannulate setigers.
Abdomen tapered with 43 setigers; pygidium
pointed.
Thoracic notosetae of three types: five or six
narrow-winged capillaries (Figure 17g), six to
eight paleae with rounded blades and short
pointed tips (Figure 17h), and a few slender,
deeply embedded setae with fine tips barely
projecting through skin. Thoracic neurosetae
as a row of about five long-shafted hooks (Figure
17c, d) with a close-set cap of denticles and a
delicate hood over the main fang. Anterior ab-
dominal segments with normal winged capil-
laries (Figure 17k) in neuropodia and uncini in
notopodia. Individual uncini (Figure 17e, f) with
short square bases and three rows of four or
five teeth above rostrum. Posterior abdominal
neuropodia with long slender-winged capillaries
(Figure 171).
Remarks. — Juvenile paratypes may have as
few as 6 radioles and the number of abdominal
segments may be as low as 30. In these young
specimens the pygidium ends in a well-developed
caudal filament (Figure 17a) but intermediates
between this and the adult type occur.
Chone americana is allied to C. filicaudata
Southern; the juveniles with their long caudal
filaments were at first thought to belong to this
species although the paleae and abdominal
uncini differ. The adult form is closer to C. duneri
but the latter has a better developed collar.
Records. — Common off Beaufort in 19 to 160
m (*).
FAMILY SERPULIDAE
Key to genera and species
1 Thorax symmetrical with 5-12 setigers
1' Thorax asymmetrical with 3-4 setigers. [Shell small and spirally
coiled (Spirorbinae)]
2 Operculum always present; no pinnules on opercular stalk.
Thorax with 6-7 setigers (Serpulinae)
2' Operculum absent or poorly developed; pinnules present on
opercular stalk. Thorax with 5-12 setigers. [Collar setae
with a separate toothed lobe at base of blade (Filograna)] .
3 Opercular stalk with a pair of small wings below operculum . .
3' Opercular stalk wingless, but sometimes wrinkled or annulated
4 Collar setae as a few winged capillaries. Operculum as a flat
or conical calcareous plate (Pomatoceros) . [Operculum
without three-pronged projections and usually conical with
a dorsal talon]
4' Collar setae with a spinulose lobe at base of blade. Operculum
as a calcareous plate with branching projections (Spiro-
branchus)
2
11
F. implexa
4
5
P. americanus
S. giganteus
129
5 Collar setae include stout bayonet-setae with conical bosses
at base of smooth blade 6
5' Collar setae all winged capillaries with smooth or serrated
blades. [Operculum chitinous, either conical or cylindrical] 13
6 Operculum a simple funnel formed of fused radii. (Serpula)
[Inner surface of funnel minutely granular] S. vermicularis granulosa
6 Operculum of two parts: a basal funnel of fused radii and an
upper crown of 8-14 horny spines. (Hydroides) [Tips of radii
ending in simple points] 7
7 Opercular spines with one or more pairs of lateral spinules;
[tips curved inwards] 8
7' Opercular spines without lateral spinules 10
8 Opercular spines with 2-3 pairs of lateral spinules. Bayonet-
setae with denticulate shaft-heads. Tube attached to floating
objects (H. elegans) No N.C. record
8' Opercular spines with one pair of lateral spinules. Bayonet-
setae with shaft-heads smooth apart from the two conical
bosses 9
9 Opercular spines equal in size H. crucigera
9' Opercular spines unequal; [often knobbed at point of in-
flection] H. parva
10 Tips of opercular spines all curved inwards toward center 11
10' Tips of opercular spines not all curved inwards toward center 12
11 Opercular spines graded in length. Many spines with an exter-
nal conical spinule near base H. sanctaecrucis
11 ' Opercular spines equal in length. All spines with an inner
hooked spinule near base H. uncinatus
12 Tips of spines curved in same direction, some pointing in-
wards, some sideways, and some outwards H. dianthus
12' Tips of spines curved outwards. [Tips of radii curved out-
wards] H. protulicola
13 Opercular stalk annulated, replacing second dorsal radiole on
left side. Operculum without internal septa. Gauge of
uncinus simple {Metavermilia) M. multicristata
13' Opercular stalk wrinkled, replacing first dorsal radiole on
right side. Operculum truncate with internal septa and
external rings. Gauge of uncinus bifurcate (Vermiliopsis) V. annulata
14 Shell coiled anticlockwise when seen from above. Incuba-
tion in tube. Collar setae without a fin at base of blade Spirorbis (Circeis) spirillum
14' Shell coiled anticlockwise when seen from above. Incuba-
tion in operculum. Collar setae with a fin at base of blade .... Spirorbis (Janua) corrugatus
130
Filograna implexa Berkeley, 1828
Filograna implexa. - Fauvel, 1927: 376, Fig. 129
a, b. - Day, 1967: 817, Fig. 38.7. a-h. -
Zibrowius, 1968: 179, pi. 10: Fig. 1-12.
Salmacina dysteri. - Fauvel, 1927: 377, Fig. 129
c-k.-Hartman, 1951: 120.
Record*. — Cape Hatteras to South Carolina,
intertidal to 20 m (11, 13, 14, 18, *).
Distribution. — Cosmopolitan in temperate and
tropical seas; intertidal to over 100 m.
Pomatoceros americanus New Species
Figure 18a-f
Pomatoceros caeruleus. -Wells and Gray, 1964:
74 (non Schmarda).
Pomatoceros triqneter. - McCloskey, 1970: 26
(non Linnaeus).
Pomatoceros n. sp. - Day, Field, and Mont-
gomery, 1971: 123.
Holotype. — USNM 43131; ca. 100 paratypes,
USNM 43132.
Description. — Tube (Figure 18a) pinkish
white, triangular in section; median ridge
smooth and projecting as a tooth over aperture.
Body up to 18 mm long. Branchial crown formed
of two arcs of 12-15 radioles united by a web for
one-third their length. Radioles with two blue
bands when fresh. Opercular stalk blue, triangu-
lar basally but flattened and broader distally,
with a pair of smooth narrow wings below oper-
culum. Operculum (Figure 18c, d) swollen, apex
conical, and calcareous resembling a Phrygian
cap; some paratypes with operculum covered
by a flattened calcareous plate (Figure 18b).
Collar incised ventrolateral ly forming paired
lateral lobes and a median ventral lobe with
small fillets in incisions. Lateral lobes extending
back as thoracic membranes to end of thorax
and united ventrally as a small apron at origin
of abdomen. Collar setae as a few, small limbate
capillaries. Notosetae of setigers 2 to 7 as larger
limbate capillaries. Thoracic neurosetae as long
rows of uncini; individual uncini (Figure 18f)
with one row of about 12 teeth preceding a broad
emarginate gouge. Abdominal neurosetae elon-
gated on posterior segments but all with a den-
ticulate apex produced as a spike on one side
(Figure 18e).
Re ma rks. — P<> m a toceros a m erica nus differs
from the European P. triqneter mainly in the
shape of the operculum. Tricorn projections
which often occur on the opercular plate of
P. triqneter, are absent in P. americanus, and
conical cap with its rectangular talon, when
well developed, is very characteristic. Speci-
mens of P. triqneter, kindly loaned to me by
Dr. George of the British Museum, do not show
this character.
P. americanus has been confused with P.
caeruleus (Schmarda), originally described
as Placostegus caeruleus from the Cape of
Good Hope and New Zealand. As shown by
Day (1955), the Cape form is a distinct species
which lacks collar setae and is now known as
Pomatoleios kraussii (Baird); the New Zealand
form of P. caeruleus has an operculum with two
tiers of plates and has not been recorded with
certainty from America. P. americanus has also
been confused with Pomatoleios caerulescens
Augener, from the Gulf of Mexico and Pomato-
ceros miuutus Rioja recorded from both coasts
of Mexico and by Zibrowius (1969) from Brazil.
A discussion of these species will be found in
Zibrowius (1970b: 15). P. miuutus differs from
P. americanus in having a tube with three ridges,
in its smaller size (maximum length 6 mm), only
five or six radioles to each branchial lobe, an
operculum with a bilobed calcareous plate,
and thoracic uncini with a bifurcated gouge.
Records. — Abundant on corals and Pecten
shells in 18-40 m off Beaufort (19, 20, 21, *).
Distribution. — I am indebted to Dr. H. Zibrow-
ius for the following extension of range: off
Sapelo Island, Ga., in 34 m; south of Tortugas,
Fla., in 75 m; Gulf of Mexico, lat 29°25'N, long
88°40'W in 40 m.
Serpula vermicularis granulosa
Marenzeller, 1884
Figure 18g
Serpula granulosa. - Marenzeller, 1884: 19, pi.
4: Fig. 1.
Serpula vermicularis. - McCloskey, 1970: 26.
Description. — Tube pinkish white, circular in
section, faintly ridged. Body 20 mm long. Oper-
cular stalk smooth and wingless; operculum
(Figure 18g) as a shallow funnel formed of 20-40
radii with blunt tips; inner surface of funnel with
131
numerous granules. Collar incised, forming a
median ventral lobe and paired lateral lobes.
Lateral lobes continuous with thoracic mem-
branes reaching setiger 7 and uniting ventrally
as a short apron at origin of abdomen. Collar
setae include limbate capillaries and stout
bayonet-setae with two bosses at base of blade.
Thoracic uncini with one row of six teeth graded
in size. Abdominal uncini with four to eight
teeth but otherwise similar to those of thorax.
Abdominal neurosetae changing from T-shaped
forms, with a denticulate blade at right angles
to shaft, to slender, wingless capillaries near
posterior end of abdomen.
Remarks. — As shown by the above descrip-
tion, the subspecies granulosa is similar to
S. vermicularis apart from the granules on the
operculum. I do not feel that it merits specific
rank.
Records. — On coral off Beaufort in 18 m
(20,*).
Hydroides ( Ei< 'po mat us ) parvus. -Augener, 1933:
366.
Hydroides parvus. - Zibrowius, 1970: 6, pi. 1:
Fig. 5.6.
Records. — Cape Hatteras area, intertidal (18).
Distribution. — North Carolina to the West In-
dies; Gulf of Mexico and Columbia; intertidal.
Hydroides sanetaecrucis (Morch, 1863)
Hydroides sanctae-crucis. - Fauvel, 1919a: 478,
Fig. 23.
Eupomatus floridamus. -Wells and Gray, 1964:
74.
Records. — North Carolina, Gulf of Mexico
and French Guiana; intertidal to a few meters.
Hydroides dianthus (Verrill, 1873)
Spirobranehus giganteus (Pallas, 1766)
Spirobranchus giganteus. - Ehlers, 1887: 286,
pi. 57: Fig. 1-7. - Pixell, 1913: 80, pi. 8: Fig.
6. - Day, 1967: 803, Fig. 38.3. h-k. - ten
Hove, 1970: 14, pi. 2: A, B, Fig. 35-63. -
Zibrowius, 1970: 14, pi. 3: Fig. 1-10.
Records. — Off Beaufort on corals in 18 m
(20,*).
Distribution. — Circumtropical in 0-50 m.
Hydroides crueigera Morch, 1863
Hydroides crueigera. - Monro, 1933c: 1083, Fig.
26.
Hydroides bispinosa. - Bush, 1910: 496. - Hart-
man, 1942a: 88.
Records. — Cape Hatteras area, intertidal, and
off Beaufort on coral in 18 m (18, 20, *).
Distribution. — South America (Punta Are-
nas); Burmuda; North Carolina; Gulf of Pana-
ma; Hawaii; intertidal to 30 m.
Hydroides parva (Tread well, 1901)
Eupomatus parvus Treadwell, 1901: 210, Fig.
79,80.
Eupomatus dianthus. - Hartman, 1945: 48, pi.
10: Fig. 1; 1951: 118. - Rioja, 1957: 260,
Fig. 15.
Hydroides dianthus. - Zibrowius, 1971a: 697,
Figs. 1-5.
Records. — Cape Hatteras to South Carolina,
intertidal to about 30 m (5, 8, 9', 11, 13, 14, 15,
18, 19,20, *).
Distribution. — Massachusetts to the West In-
dies and the Gulf of Mexico; Mediterranean;
intertidal to 30 m.
Hydroides uncinata (Philippi, 1844)
Figure 18 h, i
Eupomatus uncinatus. - Ehlers, 1887: 285, pi.
58: Fig. 6-11. - McCloskey, 1970: 26.
Hydroides uncinata. - Fauvel, 1927: 357, Fig.
122 a-h. - Zibrowius, 1968: 109, pi. 13:
Fig. 28.
Description. — Length up to 60 mm. Tube
stout, rugose, adnate, often irregularly coiled.
Opercular stalk without wings. Opercular fun-
nel (Figure 18h) radially symmetrical and formed
of about 30 radii ending in tapered points.
Opercular crown of 10-11 equal horny spines
curving inwards and ending in sharp points;
a faint lateral flange proximally, but no lateral
132
Figure 18. — Pomatoceros americanus n. sp. a, tube; 1>, flat form of operculum; c and d, lateral and dorsal views of
conical form of operculum; e, abdominal neuroseta; f, thoracic uncinus. Scrp/ilu vermicularis granulosa g,
operculum. Hydroides uncinata h, operculum; i, lateral view of a spine from the crown. Metavermelia nutlti-
o'istata j, dorsal view of anterior end; k, operculum; 1, thoracic winged capillary; m, "seta of Aponmtus"; n and
o, edge-on and lateral views of thoracic uncinus; p. abdominal geniculate seta; q, edge-on view of abdominal
uncinus.
133
spinules; each spine (Figure 18i) with an in-
wardly directed hook at its base. Seven thoracic
setigers. Thoracic membranes extending to end
of thorax. Bayonet-setae with a pair of smooth,
conical bosses at base of smooth blade. Thora-
cic uncinigerous rows black; individual uncini
with six or seven teeth.
Remarks. — H. uncinata is distinguished from
H. pseudouncinata and H. gairacoisis by the
simple points to the radii. Zibrowius (1970a:
693) reports that the worldwide records of
H. uncinata are based on several closely allied
but distinct species. He regards Philippi's origi-
nal description of H. uncinata as insufficient
and feels the name should be dropped. These
specimens from North Carolina belong to a
group or possibly one species including H.
spo)igicola Benedict, H. elegaiitulus Bush, H.
decora Treadwell, and H. alatalateralis (Jones).
I do not have the specimens to sort out this
tangle. I report the name H. uncinata and the
above description and leave other workers
with the necessary material to investigate
further.
Record*.— Off Beaufort in 6.5-18 m (20, *).
Hydroides protulicola Benedict, 1887
Hydroides protulicola Benedict, 1887: 550, pi.
20: Fig. 71, pi. 21: Fig. 18-23.
Eupomatus protulicola. - Rioja, 1946: 199, Fig.
10-13. - Hartman, 1951: 119.
Records. — Cape Hatteras to Beaufort, inter-
tidal to 40 m (13, 18, *).
Distribution. — Cape Hatteras and North Car-
olina to the Gulf of Mexico; intertidal to 40 m.
Metavermilia multicristata (Philippi, 1844)
Figure 18j-q
Y< rmiliopsis multicristata. - Zibrowius, 1968:
128, pi. 3: Fig. 25-34, pi. 14: Fig. g.
Vermiliopsis occidentalis. - McCloskey, 1970:
28 {non Mcintosh).
Metavermilia multicristata. - Zibrowius, 1971b:
1375, Fig. 1.
Description. — Tube adnate, with three ridges
including a low, regularly scalloped lateral pair,
and a smooth median keel ; no transverse ridges
formed by earlier apertures. Body slender, 11
mm long, uniformly pale in alcohol. Branchial
crown (Figure 18j) formed by two slanting bran-
chial lobes each bearing seven radioles. Oper-
cular stalk formed from long and annulated
second dorsal radiole; no wings. Operculum
(Figure 18k), fig-shaped with base soft, swollen,
and white; heavily chitinized distal part brown,
with a faint rim around truncate end; no internal
septa. Collar frilly and incised, forming paired
lateral lobes and a single ventral lobe; lateral
lobes continuous with wide thoracic membranes
to setiger 3 then abruptly narrowed but reaching
end of thorax. Collar setae as slender-winged
capillaries; notosetae of setigers 2 to 7 with
many winged capillaries (Figure 181) and a few
"setae of Apomatus" (Figure 18m) with most of
blade finely dentate. Thoracic uncini (Figure
18n,o) with a single or double row of 12 teeth
preceding the large truncate gouge. Abdominal
uncini (Figure 18q) with two or three rows of
10 teeth; abdominal capillaries (Figure 18p)
geniculate, with a tapering denticulate blade
almost at right angles to shaft.
Remarks. — V. multicristata is unusual in that
the opercular stalk replaces the second and
not the first radiole of the left branchial lobe,
further the opercular stalk is annulated and
the operculum itself lacks internal septa. Fur-
ther remarks will be found in Zibrowius (1968).
It will be noted that the tube described above
differs from that described by Zibrowius which
usually has five to seven dentate ridges.
Records. — On corals in 18 m off Beaufort
(20, *). This is a new record for the United
States.
Distribution. — Mediterranean and warm east-
ern Atlantic from the Bay of Biscay to Madeira,
Canary Islands, and Ghana; from shallow water
to 943 m.
Vermiliopsis annulata (Sehmarda, 1861)
Vcrniilia annulata Sehmarda, 1861: 28, pi. 21:
Fig. 176. - Ehlers, 1887: 308, pi. 58: Fig.
12-16; pi. 59: Fig. 1-3.
Records. — On coral reefs in North Carolina
in a few meters (14).
Distribution. — North Carolina to Florida and
the West Indies; intertidal to 4 m.
1.34
Spir or bis (Circeis) spirillum Linnaeus, 1767
Spirorbis (Dexiospira) spirillum. - Fauvel, 1927:
392, Fig. 132, f-p. - Bergen, 1953: 41, Fig.
6 a-c, pi. 1: Fig. h-i. - Gee, 1964: 417. Fig.
6a-f.
Spirorbis (Circeis) spirillum. - Bailey, 1969: 401
(list only).
Remarks. — This is a very doubtful record.
Records. — On Pecten shells off Beaufort;
intertidal to 20 m (15).
Distribution. — Arctic; North Pacific; North
Atlantic; intertidal to 20 m.
Spirorbis (Januo) corriigatus (Montagu, 1803)
Spirorbis (Dexiospira) corrugatus. - Fauvel,
1927: 393, Fig. 133 h-p. - Hartman, 1951:
121. - Zibrowius, 1968: 203. pi. 13: Fig.
16-27.
Spirorbis (Janua ) corrugatus. -Bailey, 1969: 401
(list only).
Remarks. — Professor Knight-Jones, in a per-
sonal communication, suggests that the record
from Sargassum may refer to the allied species
S. (J.)formosus Bush.
Records. — On Sargass/i m cast ashore near
Cape Hatteras (18).
Distribution . — Mediterranean; warm and
tropical Atlantic to the Gulf of Mexico; inter-
tidal to a few meters.
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