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NOAA Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
A <strong>guide</strong> <strong>to</strong> <strong>the</strong> <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong><br />
<strong>of</strong> <strong>the</strong> Aleutian Island Archipelago<br />
Robert P. S<strong>to</strong>ne<br />
Helmut Lehnert<br />
Henry Reiswig<br />
U.S. Department <strong>of</strong> Commerce<br />
September 2011
U.S. Department<br />
<strong>of</strong> Commerce<br />
Rebecca Blank<br />
Acting Secretary <strong>of</strong> Commerce<br />
National Oceanic<br />
and Atmospheric<br />
Administration<br />
Jane Lubchenco, Ph.D.<br />
Administra<strong>to</strong>r <strong>of</strong> NOAA<br />
National Marine<br />
Fisheries Service<br />
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for Fisheries<br />
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NOAA Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
A <strong>guide</strong> <strong>to</strong> <strong>the</strong> <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong><br />
<strong>of</strong> <strong>the</strong> Aleutian Island Archipelago<br />
Robert P. S<strong>to</strong>ne<br />
Helmut Lehnert<br />
Henry Reiswig<br />
September 2011<br />
U.S. Department <strong>of</strong> Commerce<br />
Seattle, Washing<strong>to</strong>n
Suggested reference<br />
S<strong>to</strong>ne, Robert P., Helmut Lehnert, and Henry Reiswig. 2011. A <strong>guide</strong> <strong>to</strong> <strong>the</strong> <strong>deep</strong><strong>water</strong><br />
<strong>sponges</strong> <strong>of</strong> <strong>the</strong> Aleutian Island Archipelago. NOAA Pr<strong>of</strong>essional Paper<br />
<strong>NMFS</strong> 12, 187 p.<br />
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CONTENTS<br />
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1<br />
About this <strong>guide</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2<br />
Zoogeography <strong>of</strong> <strong>sponges</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3<br />
Biology <strong>of</strong> <strong>sponges</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5<br />
Ecology <strong>of</strong> <strong>sponges</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7<br />
Importance as fish habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7<br />
Vulnerability <strong>to</strong> disturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7<br />
Moni<strong>to</strong>ring bycatch <strong>of</strong> <strong>sponges</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8<br />
Collection and preservation <strong>of</strong> sponge specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9<br />
Labora<strong>to</strong>ry identification <strong>of</strong> sponge specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9<br />
Calcareous <strong>sponges</strong> and demo<strong>sponges</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9<br />
Hexactinellid <strong>sponges</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9<br />
Class Calcarea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11<br />
1 . Clathrina sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12<br />
2 . Leucandra poculiformis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13<br />
3 . Leucandra tuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14<br />
Class Hexactinellida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15<br />
4 . Farrea kurilensis ssp . nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16<br />
5 . Farrea occa occa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17<br />
6 . Farrea sp . nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18<br />
7 . Family Euretidae; Genus nov ., sp . nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19<br />
8 . Tre<strong>to</strong>dictyum sp . nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20<br />
9 . Aphrocallistes vastus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21<br />
10 . Heterochone calyx calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23<br />
11 . Regadrella okinoseana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25<br />
12 . Acanthascus (Acanthascus) pr<strong>of</strong>undum ssp . nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26<br />
13 . Acanthascus (Rhabdocalyptus) dawsoni dawsoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27<br />
14 . Acanthascus (Rhabdocalyptus) mirabilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28<br />
15 . Acanthascus (Staurocalyptus) solidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29<br />
16 . Acanthascus (Staurocalyptus) sp . nov . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30<br />
17 . Acanthascus (Staurocalyptus) sp . nov . 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31<br />
18 . Aulosaccus pinularis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32<br />
19 . Aulosaccus schulzei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33<br />
20 . Bathydoris sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34<br />
21 . Caulophacus (Caulophacus) sp . nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35<br />
Class Demospongiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37<br />
22 . Plakina atka . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38<br />
23 . Plakina tanaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39<br />
24 . Craniella arb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40<br />
25 . Craniella sigmoancoratum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41<br />
26 . Craniella spinosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42<br />
27 . Craniella sputnika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43<br />
28 . Erylus aleuticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44<br />
29 . Geodia lendenfeldi nomen novum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45<br />
30 . Poecillastra tenuilaminaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46<br />
iii
CONTENTS (continued)<br />
31. Polymastia fluegeli .......................................................................... 47<br />
32. Polymastia pacifica ......................................................................... 48<br />
33. Stylocordyla borealis eous ..................................................................... 49<br />
34. Aap<strong>to</strong>s kanuux ............................................................................ 50<br />
35. Rhizaxinella clavata ........................................................................ 51<br />
36. Suberites excellens .......................................................................... 52<br />
37. Suberites simplex ........................................................................... 53<br />
38. Suberites sp. .............................................................................. 54<br />
39. Hemigellius porosus ......................................................................... 55<br />
40. Cornulum clathriata ........................................................................ 56<br />
41. Iophon piceum ............................................................................. 57<br />
42. Iophon piceum abipocillus .................................................................... 58<br />
43. Megaciella anisochela ....................................................................... 59<br />
44. Megaciella spirinae ......................................................................... 60<br />
45. Clathria (Clathria) barleei. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61<br />
46. Clathria (Clathria) laevigata ................................................................. 62<br />
47. Clathira (Axosuberites) lambei ................................................................. 63<br />
48. Echinoclathria vasa ......................................................................... 64<br />
49. Artemisina amlia .......................................................................... 65<br />
50. Artemisina arcigera ......................................................................... 66<br />
51. Artemisina stipitata ......................................................................... 67<br />
52. Artemisina sp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68<br />
53. Coelosphaera oglalai ........................................................................ 69<br />
54. Inflatella globosa ........................................................................... 70<br />
55. Lissodendoryx (Lissodendoryx) behringi .......................................................... 71<br />
56. Lissodendoryx (Ectyodoryx) olgae ............................................................... 72<br />
57. Lissodendoryx (Lissodendoryx) oxeota ........................................................... 73<br />
58. Lissodendoryx (Lissodendoryx) papillosa ......................................................... 74<br />
59. Monanchora alaskensis ...................................................................... 75<br />
60. Monanchora laminachela .................................................................... 76<br />
61. Monanchora pulchra ........................................................................ 77<br />
62. Crella brunnea ............................................................................ 78<br />
63. Hymedesmia (Stylopus) dermata ............................................................... 79<br />
64. Hymedesmia (Hymedesmia) irregularis .......................................................... 80<br />
65. Kirkpatrickia borealis ........................................................................ 81<br />
66. Phorbas paucistylifer ........................................................................ 82<br />
67. Melonanchora globogilva ..................................................................... 83<br />
68. Myxilla (Myxilla) behringensis ................................................................ 84<br />
69. Myxilla (Ectyomyxilla) parasitica .............................................................. 85<br />
70. Myxilla (Bur<strong>to</strong>nanchora) pedunculata .......................................................... 86<br />
71. Stelodoryx oxeata ........................................................................... 87<br />
72. Stelodoryx <strong>to</strong>poroki .......................................................................... 88<br />
73. Stelodoryx vitiazi ........................................................................... 89<br />
74. Echinostylinos hirsutus ...................................................................... 90<br />
iv
CONTENTS (continued)<br />
75. Tedania (Tedania) dirhaphis ................................................................ 91<br />
76. Tedania kagalaskai ........................................................................ 92<br />
77. Asbes<strong>to</strong>pluma ramosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93<br />
78. Cladorhiza bathycrinoides ................................................................... 95<br />
79. Cladorhiza corona ......................................................................... 96<br />
80. Chondrocladia (Chondrocladia) concrescens ...................................................... 97<br />
81. Biemna variantia ......................................................................... 98<br />
82. Euchelipluma elongata ..................................................................... 99<br />
83. Guitarra abbotti .......................................................................... 100<br />
84. Guitarra fimbriata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101<br />
85. Amphilectus digitatus ...................................................................... 102<br />
86. Esperiopsis flagrum ........................................................................ 103<br />
87. Semisuberites cribrosa ....................................................................... 104<br />
88. Mycale (Aegogropila) adhaerens .............................................................. 106<br />
89. Mycale (Carmia) carlilei .................................................................... 107<br />
90. Mycale (Mycale) jasoniae ................................................................... 108<br />
91. Mycale (Mycale) loveni ..................................................................... 109<br />
92. Mycale (Mycale) tylota ..................................................................... 112<br />
93. Latrunculia (Biannulata) oparinae ........................................................... 113<br />
94. Latrunculia velera ......................................................................... 114<br />
95. Latrunculia sp. (undescribed) .............................................................. 115<br />
96. Axinella blanca ........................................................................... 116<br />
97. Axinella rugosa ........................................................................... 117<br />
98. Bubaris vermiculata ....................................................................... 118<br />
99. Halichondria (Halichondria) colossea .......................................................... 119<br />
100. Halichondria (Halichondria) oblonga .......................................................... 120<br />
101. Halichondria (Eumastia) sitiens .............................................................. 121<br />
102. Halichondria sp. .......................................................................... 122<br />
103. Hymeniacidon assimilis ..................................................................... 124<br />
104. Topsentia disparilis ........................................................................ 125<br />
105. Cladocroce ventilabrum ..................................................................... 126<br />
106. Haliclona bucina ......................................................................... 127<br />
107. Haliclona (Gellius) digitata ................................................................. 128<br />
108. Haliclona (Gellius) primitiva ................................................................ 129<br />
109. Haliclona tenuiderma ...................................................................... 130<br />
110. Haliclona (Haliclona) urceolus ............................................................... 131<br />
111. Haliclona sp. 1 ........................................................................... 132<br />
112. Haliclona sp. 2 ........................................................................... 133<br />
Acknowledgments ............................................................................. 134<br />
Literature cited ............................................................................... 134<br />
Appendix I. Sponge species reported from Alaskan <strong>water</strong>s ............................................ 137<br />
Appendix II. Priorities for bycatch moni<strong>to</strong>ring ...................................................... 143<br />
Appendix III. Glossary <strong>of</strong> terms .................................................................. 146<br />
Appendix IV. Spicule types ...................................................................... 147<br />
v
Abstract—The first dedicated collections<br />
<strong>of</strong> <strong>deep</strong>-<strong>water</strong> (>80 m) <strong>sponges</strong> from<br />
<strong>the</strong> central Aleutian Islands revealed a<br />
rich fauna including 28 novel species and<br />
geographical range extensions for 53 o<strong>the</strong>rs.<br />
Based on <strong>the</strong>se collections and <strong>the</strong><br />
published literature, we now confirm <strong>the</strong><br />
presence <strong>of</strong> 125 species (or subspecies)<br />
<strong>of</strong> <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong> in <strong>the</strong> Aleutian<br />
Islands. Clearly <strong>the</strong> <strong>deep</strong>-<strong>water</strong> sponge<br />
fauna <strong>of</strong> <strong>the</strong> Aleutian Islands is extraordinarily<br />
rich and largely understudied.<br />
Submersible observations revealed that<br />
<strong>sponges</strong>, ra<strong>the</strong>r than <strong>deep</strong>-<strong>water</strong> corals,<br />
are <strong>the</strong> dominant feature shaping benthic<br />
habitats in <strong>the</strong> region and that <strong>the</strong>y provide<br />
important refuge habitat for many<br />
species <strong>of</strong> fish and invertebrates including<br />
juvenile rockfish (Sebastes spp.) and king<br />
crabs (Lithodes sp). Examination <strong>of</strong> video<br />
footage collected along 127 km <strong>of</strong> <strong>the</strong><br />
seafloor fur<strong>the</strong>r indicate that <strong>the</strong>re are<br />
likely hundreds <strong>of</strong> species still uncollected<br />
from <strong>the</strong> region, and many unknown<br />
<strong>to</strong> science. Fur<strong>the</strong>rmore, <strong>sponges</strong> are<br />
extremely fragile and easily damaged by<br />
contact with fishing gear. High rates <strong>of</strong><br />
fishery bycatch clearly indicate a strong<br />
interaction between existing fisheries and<br />
sponge habitat. Bycatch in fisheries and<br />
fisheries-independent surveys can be a major<br />
source <strong>of</strong> information on <strong>the</strong> location<br />
<strong>of</strong> <strong>the</strong> sponge fauna, but current moni<strong>to</strong>ring<br />
programs are greatly hampered by <strong>the</strong><br />
inability <strong>of</strong> deck personnel <strong>to</strong> identify bycatch.<br />
This <strong>guide</strong> contains detailed species<br />
descriptions for 112 <strong>sponges</strong> collected in<br />
Alaska, principally in <strong>the</strong> central Aleutian<br />
Islands. It addresses bycatch identification<br />
challenges by providing fisheries observers<br />
and scientists with <strong>the</strong> information<br />
necessary <strong>to</strong> adequately identify sponge<br />
fauna.Using that identification data, areas<br />
<strong>of</strong> high abundance can be mapped<br />
and <strong>the</strong> locations <strong>of</strong> indica<strong>to</strong>r species<br />
<strong>of</strong> vulnerable marine ecosystems can be<br />
determined. The <strong>guide</strong> is also designed<br />
for use by scientists making observations<br />
<strong>of</strong> <strong>the</strong> fauna in situ with submersibles,<br />
including remotely operated vehicles and<br />
au<strong>to</strong>nomous under<strong>water</strong> vehicles.<br />
A <strong>guide</strong> <strong>to</strong> <strong>the</strong> <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong><br />
<strong>of</strong> <strong>the</strong> Aleutian Island Archipelago<br />
Robert P. S<strong>to</strong>ne (contact author) 1<br />
Helmut Lehnert 2<br />
Henry Reiswig 3<br />
1 Alaska Fisheries Science Center<br />
National Marine Fisheries Service<br />
National Oceanic and Atmospheric Administration<br />
17109 Point Lena Loop Road<br />
Juneau, Alaska 99801<br />
Email address for contact author: Bob.S<strong>to</strong>ne@noaa.gov<br />
2 Freelance Sponge Taxonomy<br />
Eichenstr. 14, D-86507<br />
Oberottmarshausen, Germany<br />
3 Royal British Columbia Museum and University <strong>of</strong> Vic<strong>to</strong>ria (Biology)<br />
675 Belleville Street<br />
Vic<strong>to</strong>ria, British Columbia, Canada V8W 3N5<br />
Introduction<br />
The first dedicated collections <strong>of</strong><br />
<strong>deep</strong>-<strong>water</strong> (>80 m) <strong>sponges</strong> from<br />
<strong>the</strong> central Aleutian Islands revealed<br />
a rich fauna comprising 102 species,<br />
including 28 species new <strong>to</strong> science<br />
and range extensions for 53 species.<br />
Based on <strong>the</strong>se collections and <strong>the</strong><br />
published literature, we now confirm<br />
<strong>the</strong> presence <strong>of</strong> 125 species (or<br />
subspecies) <strong>of</strong> <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong><br />
in <strong>the</strong> central Aleutian Islands. The<br />
inven<strong>to</strong>ry includes 10 species <strong>of</strong> calcareous<br />
<strong>sponges</strong>, 20 species <strong>of</strong> hexactinellid<br />
<strong>sponges</strong>, and 95 species <strong>of</strong><br />
demo<strong>sponges</strong>. Despite <strong>the</strong> initial collection<br />
efforts, <strong>the</strong> sponge fauna <strong>of</strong><br />
this region <strong>of</strong> <strong>the</strong> North Pacific is still<br />
poorly known. Based on our extensive<br />
submersible observations, we estimate<br />
that <strong>the</strong>re are several hundred sponge<br />
species yet <strong>to</strong> be inven<strong>to</strong>ried or described<br />
from <strong>the</strong> region. This regional<br />
estimate (i.e., <strong>to</strong>tal percent known)<br />
is consistent with those made for all<br />
<strong>sponges</strong> worldwide (Hooper and Lévi,<br />
1994; Hooper and Van Soest, 2002).<br />
The main purpose <strong>of</strong> this <strong>guide</strong> is <strong>to</strong><br />
promote an awareness and appreciation<br />
<strong>of</strong> <strong>the</strong> importance <strong>of</strong> <strong>the</strong> sponge<br />
fauna in <strong>the</strong> North Pacific Ocean,<br />
particularly in <strong>the</strong> Aleutian Islands<br />
1<br />
where <strong>the</strong> diversity and abundance <strong>of</strong><br />
<strong>sponges</strong> appears <strong>to</strong> be extraordinary<br />
and bycatch in existing fisheries continues<br />
<strong>to</strong> be a major concern for resource<br />
managers. Bycatch in fisheries<br />
and fisheries-independent surveys is<br />
a major source <strong>of</strong> information on <strong>the</strong><br />
location <strong>of</strong> <strong>the</strong> sponge fauna and also<br />
a source <strong>of</strong> specimens for study. This<br />
<strong>guide</strong> serves <strong>the</strong> additional purpose <strong>of</strong><br />
providing fisheries observers and scientists<br />
with <strong>the</strong> information necessary<br />
<strong>to</strong> adequately identify sponge fauna<br />
so that <strong>the</strong> data can be included in<br />
existing databases. These data can be<br />
used <strong>to</strong> map areas <strong>of</strong> high abundance<br />
and <strong>the</strong> locations <strong>of</strong> indica<strong>to</strong>r species<br />
<strong>of</strong> vulnerable marine ecosystems. The<br />
<strong>guide</strong> is also designed for use by scientists<br />
making observations <strong>of</strong> <strong>the</strong> fauna<br />
in situ with submersibles, including<br />
remotely operated vehicles (ROVs)<br />
and au<strong>to</strong>nomous under<strong>water</strong> vehicles<br />
(AUVs).<br />
The pro<strong>to</strong>cols used <strong>to</strong> identify bycatch<br />
<strong>of</strong> <strong>sponges</strong>, both in commercial<br />
fishing operations and in fisheries surveys,<br />
have major shortcomings. They<br />
are largely restricted by <strong>the</strong> limited<br />
number <strong>of</strong> personnel dedicated <strong>to</strong><br />
this task, a general lack <strong>of</strong> knowledge<br />
<strong>of</strong> <strong>the</strong> taxonomy <strong>of</strong> <strong>the</strong> sponge fauna,<br />
and <strong>the</strong> fact that <strong>sponges</strong> are particu-
2 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
larly fragile and are <strong>of</strong>ten highly fragmented at <strong>the</strong> time<br />
<strong>of</strong> collection.<br />
Current practice aboard commercial fishing vessels<br />
in Alaska (note that not all vessels are required <strong>to</strong> have<br />
fishery observers onboard) calls for tasked personnel<br />
<strong>to</strong> identify sponge bycatch <strong>to</strong> <strong>the</strong> lowest practical taxa<br />
– essentially <strong>to</strong> Phylum Porifera. The Alaska Fisheries<br />
Science Center’s Fisheries Moni<strong>to</strong>ring and Analysis<br />
Division (FMA) maintains all records, including depth<br />
and location data, through its North Pacific Groundfish<br />
Observer Program. The accuracy <strong>of</strong> those data depends<br />
foremost on <strong>the</strong> type <strong>of</strong> fishing gear, but also varies due<br />
<strong>to</strong> <strong>the</strong> spatial coverage <strong>of</strong> <strong>the</strong> gear (i.e., only start and<br />
end positions and depths are recorded). Specimens are<br />
typically not retained for specific scientific purposes.<br />
Current practice aboard NOAA survey vessels differs<br />
between <strong>the</strong> two principal surveys. Specimens are<br />
retained for fur<strong>the</strong>r study upon request, but no formal<br />
program for <strong>the</strong> collection and preservation <strong>of</strong> specimens<br />
exists o<strong>the</strong>rwise. The emphasis <strong>of</strong> both surveys<br />
is <strong>to</strong> ga<strong>the</strong>r information necessary for <strong>the</strong> sustainable<br />
management <strong>of</strong> groundfish species. Moni<strong>to</strong>ring <strong>the</strong><br />
bycatch <strong>of</strong> structure-forming invertebrates, such as<br />
<strong>sponges</strong>, has become more <strong>of</strong> a priority with recent<br />
emphasis on managing fisheries with an ecosystem approach.<br />
For <strong>the</strong> <strong>NMFS</strong> sablefish longline survey, <strong>sponges</strong><br />
are only identified <strong>to</strong> <strong>the</strong> general level <strong>of</strong> “unidentified<br />
sponge.” For <strong>the</strong> <strong>NMFS</strong> groundfish trawl survey, wet<br />
weight <strong>of</strong> <strong>sponges</strong> is recorded ei<strong>the</strong>r completely or by<br />
subsampling. Collection data are <strong>the</strong>n coded (<strong>NMFS</strong><br />
RACE Species Code Book – maintained by <strong>the</strong> Resource<br />
Assessment and Conservation Engineering Division’s<br />
Groundfish Assessment Program) and entered in<strong>to</strong> a<br />
database. Sponges are identified from a <strong>guide</strong> that was<br />
developed specifically for <strong>the</strong> identification <strong>of</strong> benthic<br />
marine invertebrates collected along Alaska’s upper<br />
continental slope and shelf (Clark 1 ). This <strong>guide</strong> has<br />
been an important first step <strong>to</strong>ward more adequately<br />
moni<strong>to</strong>ring sponge bycatch, but it is largely incomplete,<br />
contains species that have never been confirmed<br />
<strong>to</strong> occur in Alaskan <strong>water</strong>s, and lists some species with<br />
inaccurate taxonomic nomenclature. Clearly <strong>the</strong>re is a<br />
strong need for a <strong>guide</strong> dedicated <strong>to</strong> <strong>the</strong> identification<br />
<strong>of</strong> Alaskan <strong>sponges</strong> and a continued effort <strong>to</strong> properly<br />
identify <strong>the</strong> sponge fauna collected from <strong>the</strong> region.<br />
The <strong>sponges</strong> contained in this <strong>guide</strong> were collected<br />
principally during two expeditions <strong>to</strong> <strong>the</strong> central Aleutian<br />
Islands in 2004: one aboard <strong>the</strong> RV Velero IV and<br />
<strong>the</strong> o<strong>the</strong>r aboard <strong>the</strong> RV Roger Revelle. A few additional<br />
sponge specimens were collected during <strong>the</strong> pioneering<br />
work in <strong>the</strong> region aboard <strong>the</strong> RV Velero IV in 2002 and<br />
1 Clark, R. N. 2006. Unpubl. manuscript. Field <strong>guide</strong> <strong>to</strong> <strong>the</strong><br />
benthic marine invertebrates <strong>of</strong> Alaska’s shelf and upper slope taken<br />
by NOAA/<strong>NMFS</strong>/AFSC/RACE Division trawl surveys. 302 p.<br />
2003. The focus <strong>of</strong> <strong>the</strong> research supporting <strong>the</strong> cruises<br />
was on <strong>deep</strong>-<strong>water</strong> coral habitat; <strong>the</strong>re was a dedicated<br />
effort <strong>to</strong> collect sponge fauna for formal identification<br />
only during <strong>the</strong> RV Velero IV cruise in 2004 and even <strong>the</strong>n<br />
it was <strong>of</strong> secondary importance. We also include collections<br />
made in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska aboard <strong>the</strong> RV<br />
Velero IV in 2005 and in <strong>the</strong> Bering Sea Canyons (Pribil<strong>of</strong><br />
and Zhemchug) aboard <strong>the</strong> RV Esperanza in 2007.<br />
We reviewed video footage <strong>of</strong> <strong>the</strong> seafloor collected<br />
during 31 dives with <strong>the</strong> submersible Delta and 13 dives<br />
with <strong>the</strong> ROV Jason II. A <strong>to</strong>tal <strong>of</strong> approximately 127 km<br />
<strong>of</strong> seafloor habitat was examined (35 km and 92 km,<br />
respectively) from that video footage for <strong>the</strong> presence<br />
<strong>of</strong> sponge fauna, with particular focus on taxa included<br />
in this <strong>guide</strong>. Habitat information including depth,<br />
substrate, and associated fauna were recorded for<br />
sponge observations, <strong>of</strong>ten made from multiple camera<br />
perspectives.<br />
About this <strong>guide</strong><br />
This <strong>guide</strong> contains detailed species descriptions for 112<br />
<strong>sponges</strong> collected in Alaska, principally in <strong>the</strong> central<br />
Aleutian Islands. Each species description begins with<br />
<strong>the</strong> scientific name using classical binominal nomenclature.<br />
The first name (always capitalized) is <strong>the</strong> genus.<br />
The second name (never capitalized) is <strong>the</strong> species.<br />
Some species may have a third name (never capitalized)<br />
for designated subspecies. And for some species<br />
a subgenus may have been designated and is placed in<br />
paren<strong>the</strong>ses after <strong>the</strong> current genus. The name(s) <strong>of</strong> <strong>the</strong><br />
author(s) <strong>of</strong> <strong>the</strong> species description (i.e., <strong>the</strong> person(s)<br />
who described <strong>the</strong> species) and year <strong>of</strong> description<br />
follow <strong>the</strong> scientific name. Paren<strong>the</strong>ses placed around<br />
<strong>the</strong> author(s) name(s) indicates that <strong>the</strong>re has been an<br />
accepted modification <strong>to</strong> genus assignment since <strong>the</strong><br />
original description. For example, Acanthascus (Rhabdocalyptus)<br />
dawsoni dawsoni (Lambe, 1893) is represented<br />
as Genus (subgenus) species subspecies (Author, year).<br />
Each species description provides information on<br />
1) typical growth form(s), 2) surface morphology, including<br />
<strong>the</strong> presence and description <strong>of</strong> oscula, 3) consistency<br />
and texture, 4) known size range or dimensions,<br />
and 5) color in life and under various preservation<br />
methods. These characteristics, when used in conjunction<br />
with a confirmed pho<strong>to</strong>graph, can <strong>of</strong>ten provide<br />
a fairly accurate identification in <strong>the</strong> field. Definitive<br />
identification <strong>of</strong> most species, however, requires careful<br />
examination <strong>of</strong> <strong>the</strong> arrangement <strong>of</strong> microscopic skeletal<br />
structures, particularly <strong>the</strong> types, sizes, and location <strong>of</strong><br />
spicules. Note that <strong>the</strong> identification <strong>of</strong> each species<br />
detailed in this <strong>guide</strong> has been confirmed by examination<br />
<strong>of</strong> microscopic features. We provide detailed information<br />
on <strong>the</strong> skeletal structure <strong>of</strong> each species so
that readers can definitively confirm specimen identification.<br />
For most species, we provide pho<strong>to</strong>graphs <strong>of</strong><br />
specimens on deck (shortly after collection) or archived<br />
specimens and pho<strong>to</strong>graphs <strong>of</strong> specimens in situ. The<br />
former will be most useful <strong>to</strong> fisheries observers and<br />
fishers <strong>to</strong> identify specimens on fishing and survey vessel<br />
decks shortly after collection. The latter will be useful<br />
<strong>to</strong> scientists attempting <strong>to</strong> identify and quantify sponge<br />
fauna in situ, with submersibles and remotely operated<br />
cameras. We provide <strong>the</strong> known zoogeographic<br />
range and information about <strong>the</strong> physical habitat and<br />
oceanographic habitat (i.e., temperature and salinity),<br />
if available (Koltun, 1959), <strong>of</strong> each species both within<br />
Alaskan <strong>water</strong>s and throughout its known range. Finally,<br />
we include for each species special remarks with regard<br />
<strong>to</strong> taxonomic his<strong>to</strong>ry, biology, and ecology.<br />
We used <strong>the</strong> following literature <strong>to</strong> construct <strong>the</strong> species<br />
lists and determine <strong>the</strong> zoogeography and o<strong>the</strong>r<br />
published information for each species: Austin (1985),<br />
Blake and Lissner (1994), Boury-Esnault and Rützler<br />
(1997), Brøndsted (1993), Bur<strong>to</strong>n (1934), Dickinson<br />
(1945), Hooper and Van Soest (2002), Koltun (1958,<br />
1959, 1970), Lamb and Hanby (2005), Lambe (1900),<br />
Laubenfels (1953), Sim and Kim (1988), Van Soest<br />
et al. (2008), and de Weerdt (1986a, 1986b).<br />
Appendix I provides a current and comprehensive<br />
taxonomic list <strong>of</strong> all <strong>sponges</strong> now known <strong>to</strong> occur in <strong>the</strong><br />
<strong>deep</strong> <strong>water</strong>s (>80 m) <strong>of</strong> Alaska. Appendix II provides<br />
a ranking for all <strong>the</strong> species included in this <strong>guide</strong> in<br />
terms <strong>of</strong> <strong>the</strong>ir importance as fish habitat and vulnerability<br />
<strong>to</strong> disturbance from fishing activities. The average<br />
score for <strong>the</strong>se two fac<strong>to</strong>rs is used <strong>to</strong> prioritize species for<br />
moni<strong>to</strong>ring as bycatch in commercial fisheries and s<strong>to</strong>ck<br />
assessment surveys. Appendix III includes a glossary <strong>of</strong><br />
terms commonly used in <strong>the</strong> species descriptions. The<br />
terminology used in <strong>the</strong> section on skeletal structure is<br />
not included in Appendix III. We refer <strong>the</strong> reader <strong>to</strong> <strong>the</strong><br />
Thesaurus <strong>of</strong> Sponge Morphology (Boury-Esnault and<br />
Rützler, 1997) for a comprehensive glossary <strong>of</strong> skeletal<br />
structure terminology. Appendix IV includes scanning<br />
electron microscopy (SEM) images <strong>of</strong> spicules from select<br />
calcareous, hexactinellid, and demo<strong>sponges</strong> found<br />
in Alaskan <strong>water</strong>s. The purpose <strong>of</strong> Appendix IV is <strong>to</strong><br />
provide readers with a representative collection <strong>of</strong> spicule<br />
images so that <strong>the</strong>y may gain an understanding <strong>of</strong><br />
<strong>the</strong> terms used in <strong>the</strong> <strong>guide</strong> and an appreciation for <strong>the</strong><br />
variation between species and among spicule types. This<br />
collection also serves as a source <strong>of</strong> reference material<br />
for those who wish <strong>to</strong> microscopically examine <strong>sponges</strong>.<br />
Zoogeography <strong>of</strong> <strong>sponges</strong><br />
All three major groups <strong>of</strong> <strong>sponges</strong>—Class Calcarea<br />
(calcareous <strong>sponges</strong>), Class Hexactinellida (hexacti-<br />
nellid or glass <strong>sponges</strong>), and Class Demospongiae (demo<strong>sponges</strong>)—are<br />
well represented in Alaska. Only 12<br />
(6.1%) <strong>of</strong> <strong>the</strong> 196 species <strong>of</strong> <strong>sponges</strong> now known from<br />
Alaskan <strong>water</strong>s are calcareous <strong>sponges</strong>. They have skele<strong>to</strong>ns<br />
composed entirely <strong>of</strong> calcium carbonate laid down<br />
as calcite and are consequently one <strong>of</strong> <strong>the</strong> faunal groups<br />
at high risk from increased acidification <strong>of</strong> North Pacific<br />
Ocean <strong>water</strong>s. They are principally found in shallow<br />
<strong>water</strong> and are very rare at depths below 250 m.<br />
Glass <strong>sponges</strong> are represented by 52 species (26.5%<br />
<strong>of</strong> <strong>the</strong> <strong>to</strong>tal) in Alaskan <strong>water</strong>s. Glass <strong>sponges</strong> have a siliceous<br />
skele<strong>to</strong>n consisting <strong>of</strong> spicules with a hexactinal<br />
or 6-rayed pattern. They are generally more abundant<br />
in <strong>deep</strong>er <strong>water</strong> but have a very broad depth distribution<br />
in Alaska, ranging from 20 m in <strong>the</strong> fjords <strong>of</strong> Sou<strong>the</strong>ast<br />
Alaska <strong>to</strong> more than 2800 m on <strong>the</strong> Aleutian Island<br />
slope. Hexactinellid <strong>sponges</strong> are common throughout<br />
Alaska but form different habitats depending on geographical<br />
region. In <strong>the</strong> <strong>deep</strong>er slope habitats <strong>of</strong> <strong>the</strong><br />
Aleutian Islands, we have observed large debris fields<br />
comprised mainly <strong>of</strong> glass sponge skele<strong>to</strong>ns that provide<br />
attachment substrate for o<strong>the</strong>r invertebrates, including<br />
gorgonians and hydrocorals. In <strong>the</strong> Gulf <strong>of</strong> Alaska, glass<br />
<strong>sponges</strong> are generally solitary but do form low-diversity<br />
dense patches in areas <strong>of</strong> exposed hard substrate (S<strong>to</strong>ne,<br />
unpubl. data, 2005). In sou<strong>the</strong>rn Sou<strong>the</strong>ast Alaska, in<br />
Portland Canal along <strong>the</strong> border <strong>of</strong> Alaska and British<br />
Columbia, we have found small sponge reefs, similar in<br />
species composition <strong>to</strong>, but much smaller in size than,<br />
<strong>the</strong> massive bioherms reported far<strong>the</strong>r south in British<br />
Columbia (Conway et al., 1991, 2001, 2005; Krautter et<br />
al., 2001). Small patches <strong>of</strong> hexactinellid reef, believed<br />
<strong>to</strong> be biohermal, have also been observed in nor<strong>the</strong>rn<br />
Sou<strong>the</strong>ast Alaska near Juneau, possibly indicating that<br />
<strong>the</strong>se structures are more common than originally<br />
hypo<strong>the</strong>sized and may extend up through <strong>the</strong> inside<br />
<strong>water</strong>s <strong>of</strong> <strong>the</strong> Alexander Archipelago in areas where<br />
favorable conditions exist for reef formation.<br />
Demo<strong>sponges</strong> are by far <strong>the</strong> dominant group <strong>of</strong><br />
<strong>sponges</strong> in Alaskan <strong>water</strong>s, with 132 documented species.<br />
Demo<strong>sponges</strong> also have a siliceous skele<strong>to</strong>n but <strong>the</strong><br />
spicules are not hexactinal in pattern and <strong>the</strong> skele<strong>to</strong>n<br />
may be replaced or largely supplemented with an organic<br />
collagenous network <strong>of</strong> spongin. They are found<br />
throughout Alaskan <strong>water</strong>s and over a very broad depth<br />
range (intertidal <strong>to</strong> more than 2800 m). High diversity<br />
sponge “gardens” recently discovered in <strong>the</strong> Aleutian<br />
Islands (Fig. 1) are dominated by demo<strong>sponges</strong> but<br />
also include some hexactinellid and calcareous <strong>sponges</strong>.<br />
A few <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong> (e.g., Guitarra fimbriata,<br />
Halichondria sitiens, and Hymeniacidon assimilis) are cosmopolitan<br />
in distribution (Dickinson, 1945), but recent<br />
research indicates that <strong>the</strong> occurrence <strong>of</strong> sibling species<br />
worldwide might also be common. Deep-<strong>water</strong> <strong>sponges</strong><br />
are found throughout Alaska and have been reported<br />
3
4 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Figure 1<br />
A study area in <strong>the</strong> central Aleutian Islands showing 15 sites where sponge gardens have been observed. Sponge gardens are areas <strong>of</strong> high<br />
species diversity and abundance and were located from detailed analysis <strong>of</strong> video footage collected with <strong>the</strong> Delta submersible in 2002–04<br />
and and ROV Jason II in 2004. Depth con<strong>to</strong>urs are in meters.
as far north as <strong>the</strong> Beaufort Sea (de Laubenfels, 1953).<br />
Sponges inhabit a broad depth range and occur from<br />
<strong>the</strong> intertidal zone <strong>to</strong> <strong>the</strong> <strong>deep</strong>-ocean trenches. About<br />
24% (30 <strong>of</strong> 125 species) <strong>of</strong> <strong>the</strong> <strong>sponges</strong> known from <strong>the</strong><br />
Aleutian Islands appear <strong>to</strong> be endemic <strong>to</strong> <strong>the</strong> region.<br />
Deep-<strong>water</strong> <strong>sponges</strong>, like <strong>deep</strong>-<strong>water</strong> corals, are exceptionally<br />
abundant and diverse in <strong>the</strong> Aleutian<br />
Islands (S<strong>to</strong>ne, 2006) and not surprisingly, since both<br />
faunal groups have similar habitat requirements. Both<br />
groups require stable <strong>water</strong> currents for feeding and<br />
o<strong>the</strong>r metabolic processes and most require hard,<br />
exposed substrate for attachment. The geology and<br />
oceanography <strong>of</strong> <strong>the</strong> Aleutian Island Archipelago provide<br />
unique conditions <strong>to</strong> fulfill both requirements.<br />
The Archipelago contains more than 300 islands and<br />
extends over 1900 km from <strong>the</strong> Alaska Peninsula <strong>to</strong> <strong>the</strong><br />
Kamchatka Peninsula in Russia. The Archipelago is supported<br />
by <strong>the</strong> Aleutian Ridge that forms a semi-porous<br />
boundary between <strong>the</strong> <strong>deep</strong> North Pacific Ocean <strong>to</strong> <strong>the</strong><br />
south and <strong>the</strong> shallower Bering Sea in <strong>the</strong> north. The<br />
Aleutian Ridge is a volcanic arc, with more than 20 active<br />
volcanoes and frequent earthquake activity that was<br />
formed along zones <strong>of</strong> convergence between <strong>the</strong> North<br />
American Plate and o<strong>the</strong>r oceanic plates (Vallier et al.,<br />
1994). The island arc shelf is very narrow in <strong>the</strong> Aleutian<br />
Islands and drops precipi<strong>to</strong>usly on <strong>the</strong> Pacific side <strong>to</strong><br />
depths greater than 6000 m in some areas, such as <strong>the</strong><br />
Aleutian Trench. Deep <strong>water</strong> flowing northward in <strong>the</strong><br />
Pacific Ocean encounters <strong>the</strong> Aleutian Trench where it<br />
is forced up on<strong>to</strong> <strong>the</strong> Aleutian Ridge and in<strong>to</strong> <strong>the</strong> Bering<br />
Sea through <strong>the</strong> many island passes (Johnson, 2003).<br />
Additionally, coastal <strong>water</strong> from <strong>the</strong> Alaska Stream enters<br />
through Unimak Pass in <strong>the</strong> eastern Aleutians and<br />
slowly flows nor<strong>the</strong>astward along <strong>the</strong> Alaska Peninsula.<br />
The Aleutian North Slope Current flows eastward on<br />
<strong>the</strong> north side <strong>of</strong> <strong>the</strong> Aleutian Islands <strong>to</strong>wards <strong>the</strong> inner<br />
continental shelf <strong>of</strong> <strong>the</strong> Bering Sea. This is a swift current<br />
and <strong>the</strong> steep continental slope forces much <strong>of</strong> <strong>the</strong><br />
flow in<strong>to</strong> <strong>the</strong> northwest-flowing Bering Slope Current<br />
(Johnson, 2003).<br />
The collections made by Lambe (1900), de Laubenfels<br />
(1953), and Koltun (1958, 1959) provide a firm<br />
basis <strong>to</strong> closely examine <strong>the</strong> zoography <strong>of</strong> <strong>sponges</strong> from<br />
<strong>the</strong> region. The sponge fauna <strong>of</strong> <strong>the</strong> Aleutian Island<br />
Archipelago has strong taxonomic affinities with <strong>the</strong><br />
sponge fauna <strong>of</strong> <strong>the</strong> Sea <strong>of</strong> Okhotsk (30% <strong>of</strong> species in<br />
common), seas <strong>of</strong> <strong>the</strong> Arctic Ocean (22% <strong>of</strong> species in<br />
common), <strong>the</strong> eastern (18% <strong>of</strong> species in common) and<br />
western Bering Sea (10% <strong>of</strong> species in common), and<br />
<strong>the</strong> Sea <strong>of</strong> Japan (17% <strong>of</strong> species in common). By comparison,<br />
only 12% <strong>of</strong> <strong>the</strong> sponge fauna from <strong>the</strong> Gulf <strong>of</strong><br />
Alaska is common with that <strong>of</strong> <strong>the</strong> Aleutian Islands. This<br />
zoogeographic pattern has undoubtedly been influenced<br />
<strong>to</strong> some degree by his<strong>to</strong>rical sampling effort, but<br />
<strong>the</strong> taxonomic affinities <strong>of</strong> <strong>the</strong> Aleutian Island sponge<br />
fauna are clearly much greater with <strong>the</strong> sponge fauna<br />
<strong>to</strong> <strong>the</strong> west and north than <strong>the</strong>y are with <strong>the</strong> eastern<br />
fauna. Bur<strong>to</strong>n (1934) suggested that earlier work by<br />
both Lambe and de Laubenfels indicated that <strong>the</strong> Arctic<br />
influence was apparent as far south as Vancouver Island.<br />
Several species are known from areas immediately adjacent<br />
<strong>to</strong> <strong>the</strong> Aleutian Island Archipelago and, although<br />
not yet reported from <strong>the</strong> region, likely occur <strong>the</strong>re<br />
based on geographical proximity. These include species<br />
from <strong>the</strong> eastern Bering Sea (e.g., Aap<strong>to</strong>s kanuux) and<br />
<strong>the</strong> Commander Islands (Russia) in <strong>the</strong> western Bering<br />
Sea (e.g., Grantia monstruosa, Polymastia laganoides,<br />
Asbes<strong>to</strong>pluma gracilis, and Axinella hispida).<br />
Biology <strong>of</strong> <strong>sponges</strong><br />
Basic diagrams <strong>of</strong> sponge morphology, spicule types<br />
and skeletal structures, and a comprehensive glossary <strong>of</strong><br />
terminology are available in <strong>the</strong> Thesaurus <strong>of</strong> Sponge<br />
Morphology by Boury-Esnault and Rützler (1997) and<br />
can be accessed via <strong>the</strong> World Porifera Database (Van<br />
Soest et al., 2008). 2<br />
Sponges are a primitive group <strong>of</strong> metazoans. They are<br />
sedentary animals, but a few species (e.g., Craniella spp.)<br />
may be free-living (unattached) during part <strong>of</strong> <strong>the</strong>ir life<br />
cycle (Lehnert and S<strong>to</strong>ne, 2011). The dominant feature<br />
<strong>of</strong> <strong>the</strong> typical sponge body plan is <strong>the</strong> aquiferous system<br />
through which massive amounts <strong>of</strong> <strong>water</strong> are pumped<br />
(e.g., some large <strong>sponges</strong> are capable <strong>of</strong> filtering <strong>the</strong>ir<br />
own volume <strong>of</strong> <strong>water</strong> every 20 seconds). Water flow is<br />
unidirectional and maintained by flagellated cells (choanocytes)<br />
that are usually contained within chambers<br />
where oxygen and food particles are taken up by various<br />
cell types. Water flows in through inhalent pores or ostia<br />
and out through one or more larger exhalent openings<br />
or oscula. The oscula may open in<strong>to</strong> a large cavity called<br />
an atrium or spongocoel (i.e., <strong>the</strong> large opening in tube-<br />
or vase-shaped <strong>sponges</strong>).<br />
The sponge body consists <strong>of</strong> two distinct regions:<br />
<strong>the</strong> outer region (ec<strong>to</strong>some) and <strong>the</strong> central or inner<br />
region (choanosome) where <strong>the</strong> choanocyte chambers<br />
are located. Each region typically has distinct skeletal<br />
structures with a diagnostic complement <strong>of</strong> spicules.<br />
Spicules are grouped in<strong>to</strong> two main categories: megascleres<br />
and microscleres. Megascleres are typically larger<br />
and provide <strong>the</strong> primary skeletal support. Microscleres<br />
are smaller (i.e., a microscope is required <strong>to</strong> see <strong>the</strong>m)<br />
and generally function as packing and reinforcing<br />
2 The database can be found online at http://www.marinespecies.<br />
org/porifera/. Click on “Sources”; type “Boury-Esnault” in “Sourcename”;<br />
type “publication” in “Sourcetype”; check <strong>the</strong> box <strong>to</strong><br />
“Limit <strong>to</strong> sources with full text”; and click on “Search” in lower right<br />
corner.<br />
5
6 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
structures. Many sponge cells are highly mobile and can<br />
move freely within <strong>the</strong> extracellular matrix. Some cells<br />
are also extremely pluripotent (i.e., capable <strong>of</strong> differentiating<br />
in<strong>to</strong> o<strong>the</strong>r cell types) and <strong>sponges</strong> are capable<br />
<strong>of</strong> easily remodeling cell-cell junctions. These features<br />
probably allow <strong>sponges</strong> <strong>to</strong> adapt <strong>to</strong> diverse and extreme<br />
habitats and are largely responsible for <strong>the</strong> extreme phenotypic<br />
plasticity displayed by some <strong>sponges</strong> that makes<br />
identification from pho<strong>to</strong>graphs alone so problematic.<br />
Sponges are generally nonselective filter feeders,<br />
feeding principally on bacteria, fungi, dia<strong>to</strong>ms, din<strong>of</strong>lagellates,<br />
and detritus (Bergquist, 1978; Pile et al.,<br />
1996). A recent study, however, has shown that some<br />
hexactinellid species do exhibit size independent selective<br />
filtration <strong>of</strong> ultraplank<strong>to</strong>n (Yahel et al., 2006).<br />
Carnivorous <strong>sponges</strong> were recently discovered that lack<br />
an aquiferous system al<strong>to</strong>ge<strong>the</strong>r and possess structures<br />
modified <strong>to</strong> ensnare and capture larger prey such as<br />
zooplank<strong>to</strong>n (Vacelet and Boury-Esnault, 1995; Watling,<br />
2007). Carnivorous species in Alaska are <strong>deep</strong>-<strong>water</strong> inhabitants<br />
and include Cladorhiza corona (Lehnert et al.,<br />
2005), Cladorhiza bathycrinoides, Chondrocladia concrescens,<br />
and possibly Abes<strong>to</strong>pluma ramosa.<br />
Few studies have been conducted on <strong>the</strong> growth rate<br />
and longevity <strong>of</strong> <strong>sponges</strong>, particularly those found in<br />
<strong>deep</strong>-<strong>water</strong> habitats and high-latitude ecosystems. No<br />
studies have been conducted on <strong>the</strong> growth <strong>of</strong> <strong>sponges</strong><br />
in Alaska. In general, <strong>the</strong> growth rate <strong>of</strong> temperate<strong>water</strong><br />
<strong>sponges</strong> appears <strong>to</strong> be seasonal and relatively<br />
slow, occurring at rates comparable <strong>to</strong> those <strong>of</strong> <strong>deep</strong><strong>water</strong><br />
corals (Ayling, 1983; Thomassen and Riisgård,<br />
1995; Fallon et al., 2010). Studies on <strong>the</strong> hexactinellid<br />
sponge Acanthascus (Rhabdocalyptus) dawsoni in British<br />
Columbia indicate a growth rate <strong>of</strong> 1.98 cm/yr and a life<br />
span <strong>of</strong> more than 200 years (Leys and Lauzon, 1998).<br />
Studies on <strong>the</strong> hexactinellid sponge Aphrocallistes vastus<br />
in British Columbia indicate that it may grow considerably<br />
faster (10 cm/yr) but still live in excess <strong>of</strong> a century<br />
(Austin et al., 2007). We hypo<strong>the</strong>size that growth rates<br />
for <strong>sponges</strong> in Alaska are similar <strong>to</strong> those for British<br />
Columbia <strong>sponges</strong> but note that growth studies, particularly<br />
on demo<strong>sponges</strong>, should be a high research priority<br />
in Alaska so that recovery rates from disturbance for<br />
sponge habitats can be estimated.<br />
Deep-<strong>water</strong> <strong>sponges</strong> in <strong>the</strong> Aleutian Islands appear<br />
<strong>to</strong> have few preda<strong>to</strong>rs. We have observed blood stars<br />
(Henricia spp.) displaying a typical feeding posture on<br />
several demo<strong>sponges</strong> (e.g., Artemisina sp., Monanchora<br />
pulchra, Semisuberites cribrosa, and Haliclona sp.) at shallower<br />
depths (80 <strong>to</strong> 300 m). An earlier interpretation<br />
<strong>of</strong> this behavior, however, was that <strong>the</strong> sea stars were<br />
simply taking advantage <strong>of</strong> <strong>the</strong> <strong>sponges</strong>’ feeding currents<br />
(Anderson, 1960). Henricia were generally accepted<br />
as suspension feeders (Anderson, 1960), but<br />
our additional observation <strong>of</strong> large numbers <strong>of</strong> <strong>the</strong>se<br />
sea stars present on dead <strong>sponges</strong> and decaying sponge<br />
fragments in debris “windrows” fur<strong>the</strong>r implicate predation<br />
(or scavenging). In <strong>deep</strong>er <strong>water</strong> Hippasteria spp.<br />
sea stars appear <strong>to</strong> prey on several species <strong>of</strong> <strong>sponges</strong>. In<br />
<strong>the</strong> eastern Gulf <strong>of</strong> Alaska several sea stars (Hippasteria<br />
spp., Henricia longispina, and Poraniopsis inflata) prey on<br />
glass <strong>sponges</strong> (including Acanthascus dawsoni dawsoni,<br />
A. solidus, Aphrocallistes vastus, and Heterchone calyx) and<br />
several sea stars (Hippasteria spp., H. longispina, P. inflata,<br />
Pteraster tesselatus, and Ceramaster patagonicus) prey<br />
on demo<strong>sponges</strong> (including Poecillastra tenuilaminaris,<br />
Halichondria sp., and Mycale loveni). The incidence <strong>of</strong><br />
predation on <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong> in Alaska, however,<br />
appears <strong>to</strong> be relatively low and limited <strong>to</strong> only a few<br />
species <strong>of</strong> sea stars.<br />
Advances are now being made in <strong>the</strong> study <strong>of</strong> <strong>the</strong><br />
reproductive biology <strong>of</strong> <strong>sponges</strong>, but our current<br />
knowledge is based on studies <strong>of</strong> a small fraction <strong>of</strong> <strong>the</strong><br />
species described <strong>to</strong> date worldwide; none from Alaska<br />
(Maldonado and Berquist, 2002). Sponges display<br />
highly diverse mechanisms <strong>of</strong> embryogenesis, larval<br />
differentiation, and reproduction that include both<br />
sexual and asexual processes (Berquist, 1978; Leys and<br />
Ereskovsky, 2006; Ereskovsky, 2010). Sexes are ei<strong>the</strong>r<br />
temporarily or permanently separate and some species<br />
are hermaphroditic (Blake and Lissner, 1994). Many<br />
species are capable <strong>of</strong> regenerating viable adults from<br />
fragments, and additional asexual processes include<br />
<strong>the</strong> formation <strong>of</strong> gemmules and reduction bodies,<br />
budding, and possibly formation <strong>of</strong> asexual larvae<br />
(Maldonado and Berquist, 2002). Some species are<br />
oviparous while o<strong>the</strong>rs are viviparous and brood larvae.<br />
Release <strong>of</strong> propagules (gametes, zygotes, or early embryos)<br />
is highly synchronous in oviparous species, but<br />
asynchronous in viviparous species that release fully<br />
brooded flagellated larvae (Maldonado and Berquist,<br />
2002). Several species <strong>of</strong> Geodiidae are gonochoristic<br />
and oviparous and this is assumed <strong>to</strong> be <strong>the</strong> general<br />
condition for <strong>the</strong> family (Cárdenas et al., 2009). Some<br />
species are viviparous, such as Stylocordyla, which has <strong>the</strong><br />
added peculiarity that larvae are retained alive in <strong>the</strong><br />
body until <strong>the</strong>y have fully developed (Bergquist, 1972).<br />
Gemmules, reproductive structures that can survive<br />
adverse conditions such as desiccation or extreme cold,<br />
are not typically produced by marine <strong>sponges</strong>. Their<br />
common occurrence in hermit crab <strong>sponges</strong> may represent<br />
an adaptation <strong>to</strong> help counter <strong>the</strong> consequences<br />
<strong>of</strong> stranding on shore.<br />
Many sessile marine fauna, including <strong>sponges</strong>, have<br />
evolved <strong>the</strong> ability <strong>to</strong> produce or accumulate from associated<br />
microorganisms a diversity <strong>of</strong> unique chemical<br />
compounds or secondary metabolites that <strong>the</strong>y utilize<br />
in preda<strong>to</strong>r defense, competition for resources, and<br />
as physiological adaptations <strong>to</strong> living in extreme environments<br />
(Haefner, 2003). More than 12,000 novel
compounds have been isolated from sessile marine<br />
invertebrates, algae, and microorganisms worldwide<br />
(Faulkner, 2002). Many <strong>of</strong> <strong>the</strong> compounds are currently<br />
in early clinical or late preclinical development for use<br />
as treatments for cancer, tuberculosis, HIV, asthma, and<br />
many o<strong>the</strong>r diseases and ailments (Newman and Cragg,<br />
2004). Deep-<strong>water</strong> <strong>sponges</strong> show particular promise in<br />
this emerging research area, and several species collected<br />
from <strong>the</strong> Aleutian Islands as part <strong>of</strong> a pilot program<br />
in 2004 exhibited near 100% inhibition during primary<br />
screening for M. tuberculosis (Hamann 3 ). Only a handful<br />
<strong>of</strong> sponge species from <strong>the</strong> Aleutian Islands have been<br />
examined for <strong>the</strong> presence <strong>of</strong> secondary metabolites<br />
(e.g., Na et al., 2010), but so far “hit rates” for biomedically<br />
active compounds are on <strong>the</strong> order <strong>of</strong> 10%, ra<strong>the</strong>r<br />
than 1% which is typical for samples collected elsewhere<br />
(Hamann 3 ).<br />
Ecology <strong>of</strong> <strong>sponges</strong><br />
Importance as fish habitat<br />
Deep-<strong>water</strong> <strong>sponges</strong> provide important habitat <strong>to</strong> many<br />
species <strong>of</strong> fish and invertebrates, mostly as a source <strong>of</strong><br />
refuge from predation and adverse conditions (e.g.,<br />
strong currents) and as focal sites for foraging on prey<br />
species that aggregate in sponge habitat. Some fish<br />
use <strong>sponges</strong> (e.g., Aphrocallistes vastus and Acanthascus<br />
dawsoni dawsoni) as spawning substrate (Busby 4 ) and<br />
o<strong>the</strong>rs likely use sponge habitat as breeding sites. In<br />
Alaska, many commercial and non-commercial fisheries<br />
species are associated with <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong>. Most<br />
associations are believed <strong>to</strong> be facultative ra<strong>the</strong>r than<br />
obliga<strong>to</strong>ry. Sponges provide important refuge habitat<br />
for juvenile rockfish (Sebastes spp.) in <strong>the</strong> Gulf <strong>of</strong> Alaska<br />
(Freese and Wing, 2003) and juvenile golden king crabs<br />
(Lithodes aequispina) in <strong>the</strong> central Aleutian Islands<br />
(S<strong>to</strong>ne, 2006). Sponges also contribute <strong>to</strong> <strong>the</strong> biodiversity<br />
<strong>of</strong> <strong>deep</strong>-<strong>water</strong> habitats by providing spawning substrate<br />
for many species that are important trophic links<br />
<strong>to</strong> larger consumers (Fiore and Jutte, 2010).<br />
Deep-<strong>water</strong> <strong>sponges</strong> are really no different than<br />
<strong>deep</strong>-<strong>water</strong> corals in <strong>the</strong> degree <strong>to</strong> which <strong>the</strong>y provide<br />
structure; it depends on <strong>the</strong>ir maximum size, growth<br />
form, abundance, intraspecific fine-scale distribution<br />
(i.e., patch size and density), and interaction with<br />
o<strong>the</strong>r structure-forming invertebrates. In general,<br />
large arborescent species (e.g., Axinella blanca) and<br />
3 Hamann, M. 2009. Personal commun. Unpubl. data. University<br />
<strong>of</strong> Mississippi, P.O. Box 1848, University, MS 38677.<br />
4 Busby, M. 2010. Personal commun. Alaska Fisheries Science<br />
Center, NOAA, <strong>NMFS</strong>, 7600 Sand Point Way N.E., Seattle, WA<br />
98115.<br />
those with vase-like or barrel-like morphologies (e.g.,<br />
Acanthascus dawsoni dawsoni and Mycale loveni) provide<br />
much surface area and consequently refuge space.<br />
Large <strong>sponges</strong> such as Mycale loveni have high value as<br />
fish habitat due <strong>to</strong> <strong>the</strong>ir size (up <strong>to</strong> 1 m high and wide)<br />
and abundance (up <strong>to</strong> 11 <strong>sponges</strong> per m 2 in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska). Adult sharpchin rockfish (Sebastes<br />
zacentrus) and juvenile Sebastes spp. frequently use <strong>the</strong><br />
cone-shaped <strong>sponges</strong> as perches in <strong>the</strong> eastern Gulf<br />
<strong>of</strong> Alaska. Smaller <strong>sponges</strong> at high densities may also<br />
provide important habitat. For example, finger <strong>sponges</strong><br />
(Axinella rugosa) are small (maximum dimensions only<br />
2 cm wide by 17 cm high), but are used as perches<br />
by juvenile rockfish (Sebastes spp.) when present at<br />
high densities (up <strong>to</strong> 63 individuals per m 2 ) in steep<br />
bedrock habitats (S<strong>to</strong>ne, unpubl. data, 2005). Species<br />
such as Artemisina stipitata have several attributes that<br />
make <strong>the</strong>m important as fish habitat, including large<br />
size, tendency <strong>to</strong> occur in dense patches (contagious<br />
distribution), and interaction with o<strong>the</strong>r emergent<br />
epifuana. Encrusting species such as Plakina tanaga,<br />
P. atka, and Bubaris vermiculata have little value as fish<br />
habitat. The degree <strong>to</strong> which <strong>sponges</strong> provide fish<br />
habitat also depends greatly on <strong>the</strong>ir depth distribution<br />
relative <strong>to</strong> that <strong>of</strong> shelter-seeking fishes. In <strong>the</strong> Aleutian<br />
Islands, for example, <strong>sponges</strong> found at depths greater<br />
than about 1200 m are less important as refuge habitat<br />
because most <strong>of</strong> <strong>the</strong> fish species at those depths appear<br />
<strong>to</strong> be less structure-oriented.<br />
In general, calcareous <strong>sponges</strong> are not <strong>of</strong> sufficient<br />
size or adequately abundant <strong>to</strong> provide important fish<br />
habitat. Leucandra tuba may be important for juvenile<br />
fish that use <strong>the</strong> easily accessible cavernous interior <strong>of</strong><br />
this species <strong>to</strong> escape from larger preda<strong>to</strong>rs. In contrast,<br />
demo<strong>sponges</strong> are <strong>of</strong>ten quite large and form dense<br />
“gardens” in some areas <strong>of</strong> <strong>the</strong> Aleutian Islands (Fig. 1).<br />
Hexactinellid <strong>sponges</strong> are extremely important as fish<br />
habitat. Their skele<strong>to</strong>ns are persistent after death and<br />
consequently provide <strong>the</strong> framework for <strong>the</strong> building<br />
<strong>of</strong> reefs or bioherms. Intact skele<strong>to</strong>ns, and <strong>to</strong> some degree<br />
detached and fragmented skele<strong>to</strong>ns, also provide<br />
attachment substrate for o<strong>the</strong>r sedentary structureproviding<br />
fauna, including hydrocorals, oc<strong>to</strong>corals, and<br />
demo<strong>sponges</strong> (S<strong>to</strong>ne, unpubl. data, 2004).<br />
Vulnerability <strong>to</strong> disturbance<br />
Bycatch observations in <strong>the</strong> Aleutian Islands and o<strong>the</strong>r<br />
regions <strong>of</strong> Alaska clearly support our submersible observations,<br />
indicating that <strong>sponges</strong> are far more abundant<br />
than corals in <strong>deep</strong>-<strong>water</strong> habitats. Sponges, particularly<br />
demo<strong>sponges</strong>, appear <strong>to</strong> be <strong>the</strong> most abundant emergent<br />
epifauna in <strong>the</strong> Aleutian Islands. Approximately<br />
352 metric <strong>to</strong>ns (t) <strong>of</strong> <strong>sponges</strong> are removed from <strong>the</strong> seafloor<br />
by commercial fishing in <strong>the</strong> Aleutian Islands and<br />
7
8 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Bering Sea each year – more than four times <strong>the</strong> weight<br />
<strong>of</strong> corals landed <strong>the</strong>re as fisheries bycatch (<strong>NMFS</strong> 5 ).<br />
All emergent epifauna are susceptible <strong>to</strong> disturbance<br />
from fishing gear (S<strong>to</strong>ne, 2006) and most species <strong>of</strong><br />
<strong>deep</strong>-<strong>water</strong> <strong>sponges</strong> are <strong>to</strong> some degree vulnerable <strong>to</strong><br />
disturbance (also see Hogg et al., 2010, for a recent<br />
review). Fishing operations affect <strong>the</strong> seafloor from<br />
depths <strong>of</strong> 27 m <strong>to</strong> about 1200 m in <strong>the</strong> Aleutian Islands,<br />
with most effort at depths shallower than 200 m (S<strong>to</strong>ne,<br />
2006). The degree <strong>to</strong> which a particular gear type affects<br />
sponge habitat depends on its configuration (i.e.,<br />
physical area <strong>of</strong> contact), operation (i.e., physical forces<br />
on <strong>the</strong> seafloor), spatial and temporal intensity <strong>of</strong> operation,<br />
seafloor bathymetry and substratum type, and<br />
<strong>the</strong> resiliency <strong>of</strong> <strong>sponges</strong> <strong>to</strong> disturbance. Both direct<br />
and indirect effects from fishing activities on <strong>sponges</strong><br />
likely occur. Direct effects include removal as bycatch,<br />
damage caused by physical contact, or detachment<br />
from <strong>the</strong> seafloor and translocation <strong>to</strong> unsuitable habitat.<br />
Indirect effects include increased vulnerability <strong>to</strong><br />
predation, especially for <strong>sponges</strong> detached from <strong>the</strong><br />
seafloor, and habitat alteration. Fur<strong>the</strong>rmore, <strong>the</strong>re<br />
is some evidence that reproduction is suppressed in<br />
damaged shallow-<strong>water</strong> scleractinian corals due <strong>to</strong> a<br />
reallocation <strong>of</strong> energy reserves <strong>to</strong> tissue repair and<br />
regeneration (Wahle, 1983), and similar effects may<br />
occur in <strong>sponges</strong>.<br />
Most <strong>of</strong> <strong>the</strong> fac<strong>to</strong>rs that control <strong>the</strong> degree <strong>to</strong> which<br />
<strong>sponges</strong> provide habitat structure are <strong>the</strong> same fac<strong>to</strong>rs<br />
that control <strong>the</strong> degree <strong>to</strong> which <strong>the</strong>y are vulnerable <strong>to</strong><br />
disturbance. For example, large species with arborescent<br />
growth forms are more likely <strong>to</strong> be contacted by<br />
passing fishing gear. Species such as Plakina tanaga have<br />
low vulnerability <strong>to</strong> disturbance due <strong>to</strong> <strong>the</strong>ir encrusting<br />
habitus and absence in areas where bot<strong>to</strong>m trawling is<br />
known <strong>to</strong> occur. Some species such as Poecillastra tenuilaminaris<br />
are highly vulnerable <strong>to</strong> disturbance due <strong>to</strong><br />
<strong>the</strong>ir large size and high relative abundance. Artemisina<br />
stipitata is moderately vulnerable <strong>to</strong> disturbance due <strong>to</strong><br />
its large size, erect form, and presence in areas where<br />
bot<strong>to</strong>m trawling and longline fishing are known <strong>to</strong> occur.<br />
Some <strong>sponges</strong>, for example those with holdfast<br />
structures ra<strong>the</strong>r than root-like attachment systems, may<br />
be more vulnerable <strong>to</strong> detachment, and it is unknown<br />
if intact but o<strong>the</strong>rwise undamaged <strong>sponges</strong> detached<br />
from <strong>the</strong> substrate (e.g., roller <strong>sponges</strong>; Reiswig, unpubl.<br />
data, 2010) can survive.<br />
The recovery rate <strong>of</strong> disturbed benthic ecosystems in<br />
Alaskan <strong>water</strong>s is <strong>of</strong> keen interest <strong>to</strong> fisheries managers<br />
but is still little more than conjecture due <strong>to</strong> limited<br />
5 National Marine Fisheries Service. 2004. Final programmatic<br />
supplemental groundfish environmental impact statement for Alaska<br />
groundfish fisheries. Alaska Region, NOAA, <strong>NMFS</strong>, P.O. Box<br />
21668, Juneau, AK 99802-1668.<br />
knowledge <strong>of</strong> species distribution and basic life his<strong>to</strong>ry<br />
processes. Rates <strong>of</strong> recovery will depend on growth rates<br />
and rates <strong>of</strong> sexual and asexual reproduction or speed <strong>of</strong><br />
regeneration, which are unknown for most species. Two<br />
key components <strong>of</strong> ecosystem persistence are resistance<br />
(i.e., <strong>the</strong> ability <strong>to</strong> resist changes from disturbance) and<br />
resiliency (i.e., <strong>the</strong> ability <strong>to</strong> absorb changes and <strong>the</strong><br />
speed with which it returns <strong>to</strong> its original condition)<br />
(Holling, 1973). In <strong>the</strong> case <strong>of</strong> <strong>sponges</strong>, resistance <strong>to</strong><br />
disturbance depends largely on <strong>the</strong>ir physical size, morphology,<br />
and distribution relative <strong>to</strong> <strong>the</strong> disturbance.<br />
We know from studies in <strong>the</strong> Aleutian Islands and Gulf<br />
<strong>of</strong> Alaska that benthic emergent epifauna, especially<br />
<strong>sponges</strong>, have low resistance <strong>to</strong> fishing gear disturbance<br />
(Freese et al., 1999; S<strong>to</strong>ne, 2006; Heifetz et al., 2009;<br />
S<strong>to</strong>ne, unpubl. data, 2004–2005). We know little about<br />
<strong>the</strong> resiliency <strong>of</strong> emergent epifauna, however.<br />
Resiliency depends on several fac<strong>to</strong>rs, including<br />
growth rate, recruitment rate, and reproductive ecology.<br />
Like <strong>deep</strong>-<strong>water</strong> corals, <strong>sponges</strong> have life his<strong>to</strong>ry<br />
attributes such as slow growth rates and high longevity<br />
(Day<strong>to</strong>n, 1979), indicating that long periods <strong>of</strong> time<br />
would be necessary for habitats dominated by <strong>sponges</strong><br />
<strong>to</strong> recover from disturbance (Freese, 2001). Growth<br />
rates for <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong> are generally slow (see<br />
above) and <strong>the</strong>refore inherently difficult <strong>to</strong> study.<br />
Reference sites, such as undersea cables that provide<br />
a known time-line for recruitment and subsequent<br />
growth, show promise in studies <strong>of</strong> sponge habitat recovery<br />
(Levings and McDaniel, 1974) but have not yet<br />
been identified or targeted for such study in Alaska.<br />
The only known study <strong>to</strong> address <strong>the</strong> recovery dynamics<br />
<strong>of</strong> <strong>sponges</strong> in Alaska reported no visually detectable<br />
growth or regeneration one year post-disturbance with<br />
a bot<strong>to</strong>m trawl (Freese, 2001). Due <strong>to</strong> <strong>the</strong> opportunistic<br />
nature <strong>of</strong> that study (observations were subjective, not<br />
quantifiable; not repeatable since specimens were not<br />
georeferenced; and made in an area subjected <strong>to</strong> high<br />
background disturbance), <strong>the</strong> findings are questioned.<br />
Contrastingly, a study conducted <strong>of</strong>f <strong>the</strong> sou<strong>the</strong>astern<br />
coast <strong>of</strong> <strong>the</strong> U.S. indicated that some <strong>sponges</strong> were<br />
capable <strong>of</strong> recovering completely within one year from<br />
damage caused by a research trawl (Van Dolah et al.,<br />
1987).<br />
Moni<strong>to</strong>ring bycatch <strong>of</strong> <strong>sponges</strong><br />
Appendix II provides a list <strong>of</strong> all <strong>sponges</strong> for which we<br />
have provided species descriptions in this <strong>guide</strong>. For<br />
each species, <strong>the</strong> importance as fish habitat and <strong>the</strong> vulnerability<br />
<strong>to</strong> disturbance from fishing activities is ranked<br />
from low (1) <strong>to</strong> high (3). Ranks for <strong>the</strong> two measures<br />
are averaged <strong>to</strong> provide a score. We recommend that<br />
species with scores greater than 2.0 be considered a
high priority for moni<strong>to</strong>ring as bycatch in commercial<br />
fisheries and s<strong>to</strong>ck assessment surveys and that species<br />
lists used <strong>to</strong> record bycatch in fisheries and s<strong>to</strong>ck assessment<br />
surveys be appropriately updated <strong>to</strong> include<br />
<strong>the</strong>se taxa. Subsequent mapping <strong>of</strong> <strong>the</strong>se taxa should<br />
be undertaken in an effort <strong>to</strong> identify <strong>the</strong> location <strong>of</strong><br />
vulnerable marine ecosystems. The actual or estimated<br />
weight <strong>of</strong> sponge bycatch should also be recorded since<br />
it is also a valuable indica<strong>to</strong>r <strong>of</strong> rich sponge habitat.<br />
Collection and preservation<br />
<strong>of</strong> sponge specimens<br />
The collection pro<strong>to</strong>cols outlined by Etnoyer et al.<br />
(2006) for <strong>deep</strong>-<strong>water</strong> corals are generally appropriate<br />
for <strong>the</strong> collection <strong>of</strong> <strong>deep</strong>-<strong>water</strong> <strong>sponges</strong>. Shortly after<br />
collection, specimens should be pho<strong>to</strong>graphed with<br />
an appropriate scale and <strong>the</strong> following information<br />
should be recorded: 1) growth form, 2) general surface<br />
information (roughness, presence, and description<br />
<strong>of</strong> oscula), 3) consistency, 4) size (dimensions), and<br />
5) color. Color should preferably be recorded immediately<br />
after collection since <strong>the</strong> color <strong>of</strong> some specimens<br />
may change shortly after removal from sea<strong>water</strong> and<br />
exposure <strong>to</strong> ambient light. Observations made in situ<br />
should pay particular attention <strong>to</strong> <strong>the</strong> presence and description<br />
<strong>of</strong> oscula, color, type <strong>of</strong> attachment (i.e., holdfast<br />
or root-like system), substrate type, <strong>the</strong> presence <strong>of</strong><br />
associated fauna, and if collected, whe<strong>the</strong>r <strong>the</strong> specimen<br />
appears <strong>to</strong> be whole or only a fragment <strong>of</strong> <strong>the</strong> whole<br />
specimen. The specimen should be pho<strong>to</strong>graphed or<br />
recorded on video prior <strong>to</strong> collection. Collection data<br />
should be written with pencil on <strong>water</strong>pro<strong>of</strong> paper that<br />
will be packaged with <strong>the</strong> specimen, and include date,<br />
location <strong>of</strong> collection (station number and GPS coordinates),<br />
name <strong>of</strong> collec<strong>to</strong>r, cruise number or vessel<br />
name, depth <strong>of</strong> collection, and gear type. Sponge specimens<br />
may be frozen (preferably at temperatures below<br />
–10°C) or s<strong>to</strong>red in an 80–90% ethanol solution. Demo<strong>sponges</strong><br />
and hexactinellid <strong>sponges</strong> (but never calcareous<br />
<strong>sponges</strong>) <strong>to</strong> be used for his<strong>to</strong>logical purposes may<br />
be fixed in a buffered 5% formaldehyde solution for 24<br />
hours prior <strong>to</strong> s<strong>to</strong>rage in ethanol solution. Specimens<br />
<strong>to</strong> be used for biomedical research or molecular genetic<br />
analyses should preferably be frozen <strong>to</strong> –10°C or colder<br />
unless <strong>the</strong>y will be processed o<strong>the</strong>rwise immediately.<br />
For museum archiving, specimens should preferably<br />
be s<strong>to</strong>red in an 80–90% ethanol solution. Very large<br />
specimens may be dried but a fragment <strong>of</strong> <strong>the</strong> specimen<br />
should be s<strong>to</strong>red in ethanol solution.<br />
Labora<strong>to</strong>ry identification <strong>of</strong> sponge specimens<br />
Calcareous <strong>sponges</strong> and demo<strong>sponges</strong><br />
For light microscopy and scanning electron microscopy<br />
(SEM) investigations <strong>of</strong> <strong>the</strong> spicules, small fragments <strong>of</strong><br />
<strong>sponges</strong> (containing both ec<strong>to</strong>some and choanosome)<br />
are boiled in nitric acid or hypochlorite, and in several<br />
steps <strong>the</strong> spicules are sequentially transferred through<br />
distilled <strong>water</strong> <strong>to</strong> 95% ethanol with <strong>the</strong> aid <strong>of</strong> a centrifuge<br />
(Rützler, 1978; Lehnert and Van Soest, 1998). For<br />
light microscopy, small drops <strong>of</strong> <strong>the</strong> spicule-ethanol<br />
suspension are transferred on<strong>to</strong> glass slides and, after<br />
evaporation <strong>of</strong> <strong>the</strong> ethanol, embedded in Canada balsam.<br />
Semi-thin sections obtained with a razor blade are<br />
also embedded in balsam. For SEM studies, <strong>the</strong> spiculeethanol<br />
suspension is transferred directly on<strong>to</strong> stubs<br />
and coated with gold-palladium after ethanol evaporation<br />
(Lehnert and Van Soest, 1998). Finally, data <strong>of</strong><br />
spicule types and <strong>the</strong>ir dimensions are compared <strong>to</strong> <strong>the</strong><br />
literature descriptions <strong>to</strong> assign <strong>the</strong> specimens <strong>to</strong> <strong>the</strong>ir<br />
proper genus and species.<br />
Hexactinellid <strong>sponges</strong><br />
Individual specimens are first triaged <strong>to</strong> assign <strong>the</strong>m<br />
<strong>to</strong> order and family by making temporary microscope<br />
preparations <strong>of</strong> surface spicules with commercial bleach.<br />
Specimens are <strong>the</strong>n processed by family groups. Permanent<br />
picks or peels <strong>of</strong> surface spicules are mounted in<br />
Canada balsam and pieces (about 1 cm diameter) <strong>of</strong><br />
body wall are digested in hot nitric acid <strong>to</strong> obtain a suspension<br />
<strong>of</strong> cleaned spicules. Smaller spicules are filtered<br />
on<strong>to</strong> 0.2 µm pore-size membrane filters and mounted<br />
in Canada balsam for microscopic examination. Very<br />
large spicules are picked from <strong>the</strong> suspension with<br />
forceps, cleaned, and mounted in balsam. Remaining<br />
medium-size spicules are washed, spread on slides, and<br />
mounted in balsam. Samples <strong>of</strong> individual spicule types<br />
are pho<strong>to</strong>graphed by digital camera and, when necessary,<br />
dimensions (e.g., length, diameter) are measured<br />
by digitizer-interfaced microscopy. Data <strong>of</strong> spicule types<br />
and <strong>the</strong>ir dimensions are finally compared <strong>to</strong> <strong>the</strong> literature<br />
descriptions <strong>to</strong> assign <strong>the</strong> specimens <strong>to</strong> <strong>the</strong>ir proper<br />
genus and species.<br />
9
11<br />
CLASS CALCAREA
12 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
1. Clathrina sp.<br />
Description. Conical tubes have thin walls laid in<br />
folds. Viewed with a hand-lens, <strong>the</strong> walls consist <strong>of</strong> a<br />
mesh <strong>of</strong> tissue with many open inhalant pores, round<br />
<strong>to</strong> oval, 300–600 µm in diameter. The osculum at <strong>the</strong><br />
<strong>to</strong>p <strong>of</strong> <strong>the</strong> cone has no spicule crown. Consistency is<br />
s<strong>of</strong>t and elastic. Height <strong>of</strong> cones is <strong>to</strong> 3–5 cm; several<br />
basally fused cones may cover larger areas. Color in life<br />
is creamy white <strong>to</strong> pale yellow.<br />
Skeletal structure. Triactines are arranged tangentially;<br />
large triactines (50–150 × 7–10 µm/ray), small<br />
triactines (20–30 × 3–6 µm/ray).<br />
Zoogeographic distribution. Cosmopolitan. In Alaska<br />
– central Aleutian Islands. Elsewhere – <strong>the</strong> genus Clathrina<br />
presently contains 87 nominal species and has a<br />
worldwide distribution.<br />
Habitat. In Alaska – sometimes grows around and partially<br />
encrusts <strong>the</strong> gorgonian Muriceides nigra at depths<br />
near 119 m.<br />
Remarks. The genus Clathrina was erected by Gray<br />
(1867) and is in desperate need <strong>of</strong> revision at <strong>the</strong> species<br />
level.<br />
Pho<strong>to</strong>s. Fragment <strong>of</strong> Clathrina sp. collected in <strong>the</strong> central<br />
Aleutian Islands at a depth <strong>of</strong> 119 m. Grid marks are<br />
1 cm 2 . 2) The same specimen as in pho<strong>to</strong> 1 (center) in<br />
situ encrusting <strong>the</strong> gorgonian Muriceides nigra.
2. Leucandra poculiformis Hozawa, 1918<br />
Description. Sponge is thickly encrusting. The surface<br />
is smooth <strong>to</strong> <strong>the</strong> unaided eye but rough <strong>to</strong> <strong>the</strong> <strong>to</strong>uch due<br />
<strong>to</strong> large protruding spicules. Consistency is stiff, slightly<br />
compressible, and elastic. Individuals may reach a size<br />
<strong>of</strong> approximately 6 × 2 × 1 cm. Color in life is white <strong>to</strong><br />
creamy white.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. There are large triactines (50–170 ×<br />
5–15 µm/ray) and occasionally tetractines <strong>of</strong> <strong>the</strong> same<br />
dimensions. Small oxeas (60–85 × 3–6 µm) have unequal<br />
ends, most with a characteristic bend near one<br />
end. Small oxeas are only at <strong>the</strong> surface <strong>of</strong> inner and<br />
outer walls, equiangular and sagittal triactines are inbetween.<br />
Outer surfaces are covered by a mesh <strong>of</strong> small<br />
diactines, mesh-size approximately 25 µm in diameter;<br />
tracts form <strong>the</strong> mesh 20–40 µm in diameter.<br />
Zoogeographic distribution. Rare. In Alaska – central<br />
Aleutian Islands. Elsewhere – previously known only<br />
from <strong>the</strong> Sea <strong>of</strong> Japan.<br />
Habitat. Grows around and encrusts Stylaster hydrocorals<br />
at depths near 175 m. Species <strong>of</strong> Craniella may be<br />
attached <strong>to</strong> it (see pho<strong>to</strong> 1).<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
175 m in <strong>the</strong> central Aleutian Islands. Grid marks are 1<br />
cm 2 . 2) The same specimen as in pho<strong>to</strong> 1 in situ encrusting<br />
a Stylaster hydrocoral.<br />
13
14 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
3. Leucandra tuba Hozawa, 1918<br />
Description. Sponge has thin-walled (0.5 mm) tubes.<br />
Up <strong>to</strong> 17 individual tubes surround large internal<br />
areas. Surface is smooth <strong>to</strong> <strong>the</strong> unaided eye, irregularly<br />
undulating (i.e., wavy or sinuous) oxeas, and rough <strong>to</strong><br />
<strong>the</strong> <strong>to</strong>uch due <strong>to</strong> large protruding spicules. Walls are<br />
compressible and only slightly elastic. Diameter is <strong>to</strong> 30<br />
cm. Color in life is light grey <strong>to</strong> creamy white.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Walls consist <strong>of</strong> giant tri- and tetractines<br />
(200–1200 × 20–100 µm/ray), one ray usually shorter<br />
than <strong>the</strong> o<strong>the</strong>rs. Irregularly distributed and undulating<br />
oxeas (95–140 × 3–5 µm) are very abundant and<br />
are scattered in between <strong>the</strong> giant tri- and tetractines<br />
without obvious orientation. Ec<strong>to</strong>somal areas with inhalant<br />
pores have an outer layer <strong>of</strong> special spicules called<br />
pugioles (18–35 µm long) which form meshes, leaving<br />
open pores <strong>of</strong> about 50 µm diameter, with pugiole-tracts<br />
<strong>of</strong> approximately 25 µm in diameter. The mesh <strong>of</strong> <strong>the</strong><br />
pugioles is underlain by a mesh <strong>of</strong> microdiactines <strong>of</strong> <strong>the</strong><br />
same dimensions.<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – Aleutian Islands and Bering Sea canyons. Elsewhere<br />
– previously known only from <strong>the</strong> Sea <strong>of</strong> Japan.<br />
Habitat. In <strong>the</strong> Aleutians Islands – depths between<br />
112 and 250 m on exposed bedrock, boulders, cobbles,<br />
and pebbles in generally low-relief (i.e., flat-bot<strong>to</strong>med)<br />
habitats. In <strong>the</strong> Bering Sea – known from Pribil<strong>of</strong> Canyon<br />
at depths between 203 <strong>to</strong> 220 m on pebbles and<br />
cobbles. Elsewhere – <strong>the</strong> holotype was described from<br />
<strong>the</strong> Sea <strong>of</strong> Japan near Okinoshima and collected at a<br />
depth <strong>of</strong> 106 m.<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
145 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 . 2) The same specimen as in pho<strong>to</strong> 1 in situ.<br />
3) Specimen (center, <strong>to</strong>p) at a depth <strong>of</strong> 120 m in <strong>the</strong><br />
central Aleutian Islands.
15<br />
CLASS HEXACTINELLIDA
16 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
4. Farrea kurilensis ssp. nov. Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. Lace-like mass has large-caliber (2-cm<br />
diameter), short, branching, and anas<strong>to</strong>mosing tubes<br />
increasing in size (<strong>of</strong> <strong>the</strong> mass) by lateral growth, conforming,<br />
and attaching at multiple sites <strong>to</strong> <strong>the</strong> hard<br />
substrate and not growing erect. Surface <strong>of</strong> <strong>the</strong> mass<br />
is labyrinthic, but that <strong>of</strong> individual tube elements is<br />
smooth. Open terminal ends <strong>of</strong> tubes are effectively<br />
oscula. Consistency is flexible but brittle due <strong>to</strong> fusion<br />
<strong>of</strong> <strong>the</strong> very thin primary skele<strong>to</strong>n. Individuals observed<br />
in situ have a diameter <strong>of</strong> 170 cm. Color in life is translucent<br />
light blue <strong>to</strong> white.<br />
Skeletal structure. The primary framework is a fused<br />
farreoid lattice <strong>of</strong> hexactins forming a network <strong>of</strong> square<br />
meshes with sides 439–750 µm long; it is continuous<br />
throughout <strong>the</strong> specimen. Loose megascleres include<br />
dermal and atrial pentactins, <strong>of</strong>ten with knobs or very<br />
short distal rays (170–375 µm tangential ray length,<br />
211–422 µm proximal ray length); very large anchorate<br />
clavules with completely smooth shafts (302–1243<br />
µm length); moderate-sized uncinates (1.03–2.99 mm<br />
length); very rare choanosomal hexactins (208–239<br />
µm ray length). Microscleres consist <strong>of</strong> two size classes<br />
<strong>of</strong> stellate discohexasters (30–89 µm and 91–293 µm in<br />
diameter).<br />
Zoogeographic distribution. Locally common and<br />
abundant in some areas. In Alaska – central Aleutian<br />
Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> bedrock, muds<strong>to</strong>ne, boulders,<br />
cobbles, and hexactinellid sponge skele<strong>to</strong>ns at depths<br />
between 300 and 2249 m.<br />
Remarks. Farrea kurilensis presently has two subspecies:<br />
F. k. kurilensis, which occurs near <strong>the</strong> Kuril Islands<br />
and in <strong>the</strong> Sea <strong>of</strong> Okhotsk, has smooth pileate clavules,<br />
while F. k. beringiana, which also occurs near <strong>the</strong> Kuril<br />
Islands, has all clavules with spines. The new Aleutian<br />
subspecies differs from <strong>the</strong>se in having anchorate clavules<br />
with smooth shafts and an alternate microsclere<br />
combination.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen <strong>the</strong>n dried) fragment <strong>of</strong><br />
a specimen collected at a depth <strong>of</strong> 2105 m in <strong>the</strong> central<br />
Aleutian Islands. Grid marks are 1 cm 2 . 2) The same<br />
specimen as in pho<strong>to</strong> 1 in situ.
5. Farrea occa occa Bowerbank, 1862<br />
Description. Lace-like hemispheric mass has largecaliber<br />
(1.5 <strong>to</strong> 2-cm diameter), short, branching and<br />
anas<strong>to</strong>mosing tubes increasing in size (<strong>of</strong> <strong>the</strong> mass)<br />
by lateral growth, attached <strong>to</strong> hard substrate at central<br />
origin and at multiple lateral contact sites. Surface <strong>of</strong><br />
<strong>the</strong> mass is labyrinthic with open ends <strong>of</strong> tubes functioning<br />
as oscula, but external and internal surfaces<br />
<strong>of</strong> individual tube elements are smooth. Consistency is<br />
flexible but brittle and fragile due <strong>to</strong> breakage <strong>of</strong> <strong>the</strong><br />
very thin fused primary skele<strong>to</strong>n. An individual mass<br />
from a single settled larva may reach 3 m in diameter<br />
and over 1 m in height. Color in life is translucent white<br />
<strong>to</strong> pale yellow.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The primary framework is a fused farreoid<br />
lattice <strong>of</strong> hexactins forming a network <strong>of</strong> square<br />
meshes with sides 79–494 µm long; it is continuous<br />
throughout <strong>the</strong> specimen. Loose megascleres include<br />
dermal and atrial pentactins (190–305 µm tangential ray<br />
length, 78–280 µm proximal ray length); pileate and anchorate<br />
clavules usually with smooth shafts, rarely with<br />
one or two large spines (196–308 µm length); moderatesized<br />
uncinates (0.89–2.64 mm length). Microscleres<br />
are mainly oxyhexasters with relatively long primary<br />
rays (57–111 µm in diameter). There may be a very few<br />
discohexasters or onychohexasters.<br />
Zoogeographic distribution. Locally common and<br />
abundant in some areas. In Alaska – eastern Gulf <strong>of</strong><br />
Alaska. Elsewhere – cosmopolitan, but <strong>the</strong> previous<br />
known nor<strong>the</strong>rn limit <strong>of</strong> <strong>the</strong> subspecies was 55.4°N in<br />
nor<strong>the</strong>rn British Columbia; <strong>the</strong> new Alaska collections<br />
extend <strong>the</strong> nor<strong>the</strong>rn limit <strong>to</strong> 58.2°N.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, boulder, and<br />
cobbles at depths between 91 and 238 m. Elsewhere<br />
– attached <strong>to</strong> cobbles, bedrock, shell, coral skele<strong>to</strong>ns,<br />
and hexactinellid sponge skele<strong>to</strong>ns at depths between<br />
86 and 1360 m.<br />
Remarks. This may be <strong>the</strong> largest species <strong>of</strong> sponge in<br />
<strong>the</strong> North Pacific Ocean, with a diameter reaching 3 m.<br />
This sponge provides much refuge habitat for small fish<br />
and micro-invertebrates that may be forage species for<br />
larger fish and crabs.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> a specimen collected at a<br />
depth <strong>of</strong> 168 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks<br />
are 1 cm 2 . 2) Specimen at a depth <strong>of</strong> 165 m in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska.<br />
17
18 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
6. Farrea sp. nov. Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. Erect bush <strong>of</strong> small-caliber (8-mm diameter),<br />
relatively long, branching tubes increasing in<br />
size (<strong>of</strong> <strong>the</strong> entire specimen, not <strong>the</strong> tube elements) by<br />
terminal growth <strong>of</strong> <strong>the</strong> constituent tubes; attached <strong>to</strong><br />
hard substrate at a small original base. Surface <strong>of</strong> <strong>the</strong><br />
mass consists <strong>of</strong> <strong>the</strong> terminal tube apertures as oscula,<br />
but both inner and outer surfaces <strong>of</strong> individual tube<br />
elements are smooth. Consistency is stiff and brittle<br />
due <strong>to</strong> fusion <strong>of</strong> <strong>the</strong> thin primary skele<strong>to</strong>n. Size <strong>of</strong> <strong>the</strong><br />
original specimen from which <strong>the</strong> fragments were collected<br />
was not recorded, but it must have been at least<br />
10 cm tall. Color in life is white; preserved fragments<br />
are light brown.<br />
Skeletal structure. The primary framework is a fused<br />
farreoid lattice <strong>of</strong> hexactins forming a network <strong>of</strong> rectangular<br />
meshes with longitudinal sides 269–720 µm<br />
long and lateral sides 534–778 µm long; it is continuous<br />
throughout <strong>the</strong> specimen. Loose megascleres include<br />
dermal and atrial pentactins, <strong>of</strong>ten with knobs as rudiments<br />
<strong>of</strong> <strong>the</strong> sixth distal rays (159–361 µm tangential<br />
ray length, 206–452 µm proximal ray length); pileate<br />
clavules without shaft thorns (333–490 µm length);<br />
moderate-size uncinates (1.02–2.98 mm length). Microscleres<br />
include oxyhexasters and hemioxyhexasters<br />
(62–117 µm in diameter); oxyhexactins (83–130 µm<br />
in diameter); discohexasters and hemidiscohexasters<br />
(48–76 µm in diameter); discohexactins (58–98 µm in<br />
diameter).<br />
Zoogeographic distribution. In Alaska – a rare species,<br />
known only from a few locations in sou<strong>the</strong>rn<br />
Amchitka Pass and near Bobr<strong>of</strong> Island in <strong>the</strong> central<br />
Aleutian Islands. Elsewhere – not reported.<br />
Habitat. In Alaska – attached <strong>to</strong> small boulders,<br />
cobbles, and pebbles at depths between 529 and 905 m.<br />
Remarks. Our Aleutian specimen was compared with<br />
<strong>the</strong> four o<strong>the</strong>r species <strong>of</strong> Farrea that lack anchorate<br />
clavules. It differs from each <strong>of</strong> <strong>the</strong>m in its microsclere<br />
complement. Its description as a new species <strong>of</strong> Farrea<br />
is now in progress.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen <strong>the</strong>n dried) fragments <strong>of</strong><br />
a specimen collected at 887 m in <strong>the</strong> central Aleutian<br />
Islands. 2) The same specimen as in pho<strong>to</strong> 1 (indicated<br />
by <strong>the</strong> white arrow) in situ. The separation between <strong>the</strong><br />
red laser marks is 10 cm.
7. Family Euretidae; Genus nov., sp. nov. Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. Tall, circular, rigid funnel is attached<br />
<strong>to</strong> hard substrate, with small hollow digitate processes<br />
projecting from all lateral surfaces. In life <strong>the</strong> lateral<br />
processes are closed distally by tissues and loose spicules,<br />
but <strong>the</strong> fused skeletal framework is open at <strong>the</strong> ends.<br />
The single large circular osculum at <strong>the</strong> distal end has a<br />
crenulate margin. All surfaces are smooth <strong>to</strong> <strong>the</strong> naked<br />
eye, but under low magnification <strong>the</strong>y are seen <strong>to</strong> be<br />
ornamented with fine hairs <strong>of</strong> projecting uncinate spicules.<br />
Consistency is hard and rigid. Height is <strong>to</strong> 29.2 cm<br />
and diameter <strong>to</strong> 13.7 cm; lateral processes 6–12 mm in<br />
diameter and up <strong>to</strong> 24 mm long are distributed ra<strong>the</strong>r<br />
evenly without pattern and commonly divide in<strong>to</strong> two<br />
or three short branches in larger specimens. Color in<br />
life is yellow-orange <strong>to</strong> light orange.<br />
Skeletal structure. The primary rigid skele<strong>to</strong>n <strong>of</strong><br />
fused hexactine spicules has elongate meshes with<br />
longitudinal strands and radial septa, similar <strong>to</strong> <strong>the</strong><br />
primary framework <strong>of</strong> Chonelasma. That framework<br />
curves smoothly out in<strong>to</strong> and continues through <strong>the</strong><br />
digitate processes. Slight indications <strong>of</strong> irregular dermal<br />
and atrial cortices are present, but <strong>the</strong>y are usually<br />
only one dictyonalium in thickness. There are several<br />
types <strong>of</strong> loose megascleres: pinular hexactins on inner<br />
and outer surfaces <strong>of</strong> wall and processes, with thorned<br />
pinulus (67–287 µm long); tangential rays spined at <strong>the</strong><br />
tips (88–196 µm long); proximal ray (79–954 µm long);<br />
simple hexactins and pentactins in subatrial position <strong>of</strong><br />
wall and processes (106–518 µm ray length); tauactins<br />
only in spicule pads at tips <strong>of</strong> processes (158–479 µm ray<br />
length); diactins in atrial surface <strong>of</strong> wall and processes<br />
(88–196 µm ray length); discoscopules on both surfaces<br />
<strong>of</strong> wall and processes (234–480 µm long); tyloscopules<br />
only echinating atrial surface <strong>of</strong> processes (619–952<br />
µm long); uncinates echinating both surfaces <strong>of</strong> wall<br />
and processes (656–2785 µm long). Microscleres are<br />
mainly discohexactins (76%; 52–98 µm in diameter),<br />
moderately common oxyhexactins (20%; 34–94 µm<br />
in diameter), and few hemidiscohexasters (4%; 46–<br />
100 µm in diameter).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
19<br />
Habitat. Occurs singly or in small patches on exposed<br />
bedrock, muds<strong>to</strong>ne, boulders, and cobbles at depths<br />
between 773 and 2084 m.<br />
Remarks. Juvenile Verrill’s Paralomis crabs (Paralomis<br />
verrilli) use <strong>the</strong> spongocoel as refuge habitat (S<strong>to</strong>ne,<br />
unpubl. data, 2004).<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
1256 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ.
20 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
8. Tre<strong>to</strong>dictyum sp. nov. Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. This sponge has a thin-walled flaring<br />
funnel or trumpet that attaches <strong>to</strong> hard substrate by<br />
a short narrow stalk ending in a small basal disc. The<br />
outer dermal surface is smooth with a thin transparent<br />
spicule lattice covering a system <strong>of</strong> radiating ridges (average<br />
1.1 mm wide) and grooves (average 0.8 mm wide)<br />
visible <strong>to</strong> <strong>the</strong> naked eye. The internal atrial surface, also<br />
smooth, is thoroughly and evenly pocked by openings<br />
<strong>of</strong> small exhalant canals (average 1.5 mm diameter).<br />
Under low magnification all surfaces appear furry due<br />
<strong>to</strong> projecting ends <strong>of</strong> uncinate and scopule spicules.<br />
Consistency is s<strong>to</strong>ny hard but crumbly and quite brittle.<br />
Only a single partial specimen was collected and measured<br />
25.4 cm tall by 20.6 cm wide at <strong>the</strong> margin; wall<br />
thickness was 5–6 mm. Color in life is white; specimens<br />
dried or preserved in ethanol are brownish orange.<br />
Skeletal structure. The main skele<strong>to</strong>n is a rigid framework<br />
<strong>of</strong> fused hexactins with grooves and septa typical <strong>of</strong><br />
<strong>the</strong> genus. Loose megascleres include rough pentactins<br />
<strong>to</strong> hexactins with short distal ray (19–164 µm long), tapered<br />
tangential rays (106–473 µm long), and proximal<br />
rays (94–896 µm long); small rough regular hexactins<br />
(84–177 µm long rays); scopules with rounded tine tips<br />
(294–965 µm <strong>to</strong>tal length); uncinates (437–1480 µm<br />
long). Microscleres are mostly oxyhexasters and hemioxyhexasters<br />
(93%) with 1–4 nearly smooth, robust terminal<br />
rays (53–73 µm diameter); stellate discohexasters<br />
(7%) with 4–10 finely rough terminal rays (50–75 µm<br />
diameter); a very few oxyhexactins (>1%) similar in size<br />
<strong>to</strong> oxyhexasters.<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Occurs singly on bedrock, muds<strong>to</strong>ne, boulders,<br />
and cobbles at depths between 704 and 1264 m.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at<br />
a depth <strong>of</strong> 866 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ.<br />
The separation between <strong>the</strong> red laser marks is 10 cm.
9. Aphrocallistes vastus Schulze, 1886<br />
Description. Basic form is a hollow, thin-walled cone,<br />
but larger, older specimens add lateral mitten-like outgrowths<br />
becoming highly variable; overall it is similar <strong>to</strong><br />
Heterochone calyx with which it is <strong>of</strong>ten confused. Surface<br />
is smooth, usually with a single large terminal osculum.<br />
Consistency is rigid and brittle. Size is up <strong>to</strong> 2 m high<br />
and 3 m laterally. Color in life varies from white <strong>to</strong> light<br />
yellow and orange.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Rigid skele<strong>to</strong>n <strong>of</strong> fused hexactinal spicules<br />
has distinctive honeycomb pattern <strong>of</strong> 1 mm wide<br />
channels. Several types <strong>of</strong> loose megascleres include<br />
dermal pinular hexactins with pinulus (109–185 µm<br />
long), tangential rays (38–142 µm long), proximal ray<br />
(53–232 µm long); small hexactins (85–201 µm/ray);<br />
scopules on both surfaces (262–497 µm long); atrial<br />
spiny diactins (262–722 µm long); uncinates (890–3440<br />
µm long). Microscleres are oxyhexasters (126–252 µm<br />
in diameter) and two types <strong>of</strong> discohexasters (16–25 and<br />
30–56 µm in diameter).<br />
Zoogeographic distribution. North Pacific Ocean;<br />
locally common and abundant in some areas. Alaska –<br />
Bering Sea <strong>to</strong> Sou<strong>the</strong>ast Alaska. Elsewhere – Japan <strong>to</strong><br />
Baja Mexico.<br />
Habitat. Aleutian Islands – attached <strong>to</strong> cobbles and<br />
pebbles (low-relief habitat) and sometimes bedrock<br />
(high-relief habitat) at depths between about 100 and<br />
756 m. Bering Sea – attached <strong>to</strong> cobbles and pebbles at<br />
depths between 373 and 522 m. Gulf <strong>of</strong> Alaska – principally<br />
on bedrock, but also boulders and cobbles on <strong>the</strong><br />
continental shelf and upper slope at depths between<br />
140 and at least 228 m. Observed in <strong>the</strong> glacial fiords<br />
21<br />
<strong>of</strong> Sou<strong>the</strong>ast Alaska growing on bedrock at depths as<br />
shallow as 20 m. Elsewhere – Lamb and Hanby (2005)<br />
provide a worldwide depth range <strong>of</strong> 10 and 1600 m.<br />
Remarks. This is one <strong>of</strong> <strong>the</strong> most ecologically important<br />
<strong>sponges</strong> in Alaska and perhaps <strong>the</strong> most thoroughly<br />
studied sponge species in <strong>the</strong> North Pacific Ocean. Due<br />
<strong>to</strong> its rigid skele<strong>to</strong>n, this species is an important structural<br />
component <strong>of</strong> <strong>the</strong> sponge reefs reported along<br />
<strong>the</strong> Pacific Coast <strong>of</strong> Canada (Conway et al., 1991, 2005;<br />
Krautter et al., 2001) and more recently in sou<strong>the</strong>rn<br />
Sou<strong>the</strong>ast Alaska. Juvenile golden king crabs (Lithodes<br />
aequispina) use <strong>the</strong> spongocoel as refuge habitat in<br />
<strong>the</strong> Aleutian Islands (S<strong>to</strong>ne, 2006) and in <strong>the</strong> Bering<br />
Sea Canyons (S<strong>to</strong>ne, unpubl. data, 2007). The bigmouth<br />
sculpin (Hemitripterus bolini) deposits its eggs in<br />
<strong>the</strong> spongocoel in <strong>the</strong> Bering Sea and Gulf <strong>of</strong> Alaska<br />
(Busby 3 ). This species is preyed upon by <strong>the</strong> sea stars<br />
Hippasteria spp. and Poraniopsis inflata in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska. Aphrocallistes vastus can be distinguished<br />
from <strong>the</strong> very similar Heterochone calyx calyx by <strong>the</strong> lack<br />
<strong>of</strong> pinular hexactins on <strong>the</strong> inner (atrial) surface and by<br />
<strong>the</strong> possession <strong>of</strong> very robust oxyhexasters with primary<br />
rays subsumed in <strong>the</strong> swollen centrum.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 168 m<br />
in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks are 1 cm 2 . 2)<br />
Specimen with a juvenile golden king crab (Lithodes<br />
aequispina) in pre-molt condition at a depth <strong>of</strong> 190<br />
m in <strong>the</strong> central Aleutian Islands. 3) Specimen with a<br />
swarm <strong>of</strong> euphasiids at a depth <strong>of</strong> 170 m in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska. Pho<strong>to</strong> by J. Lincoln Freese (AFSC). 4)<br />
Specimen at a depth <strong>of</strong> 190 m in <strong>the</strong> central Aleutian<br />
Islands. 5) Specimen at a depth <strong>of</strong> 20 m with Eualas sp.<br />
shrimp in Glacier Bay National Park, Sou<strong>the</strong>ast Alaska.
22 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
9. Aphrocallistes vastus Schulze, 1886 (continued)
10. Heterochone calyx calyx Schulze, 1886<br />
Description. This sponge is polymorphic and similar<br />
<strong>to</strong> Aphrocallistes vastus, with which it is <strong>of</strong>ten confused.<br />
Sponge is cup or funnel shaped, and is up <strong>to</strong> at least 30<br />
cm in height and 40 cm in diameter. It forms bowls or<br />
plates in areas <strong>of</strong> low current. Lateral walls are <strong>of</strong>ten with<br />
hollow finger-shaped processes. Color is bright gold or<br />
pale yellow in <strong>the</strong> Aleutian Islands and Bering Sea. It<br />
displays two typical color morphs in <strong>the</strong> Gulf <strong>of</strong> Alaska<br />
– white and golden yellow.<br />
Skeletal structure. It has a rigid skele<strong>to</strong>n <strong>of</strong> fused<br />
hexactinal spicules with a poorly delineated honeycomb<br />
pattern <strong>of</strong> 1 mm wide channels passing vertically<br />
through <strong>the</strong> walls. Several types <strong>of</strong> loose megascleres<br />
include pinular hexactins on inner and outer surfaces,<br />
with thorned pinulus (48–150 µm long), tangential rays<br />
spined at <strong>the</strong> tips (100–302 µm long), and proximal ray<br />
(69–1265 µm long); scopules on both surfaces (242–630<br />
µm long); spined hexactins (80–204 µm/ray); uncinates<br />
(500–1540 × 14–54 µm); rough centrotylote diactins<br />
(308–786 µm/ray) are apparently absent in some specimens.<br />
Microscleres are discohexactins and discohexasters<br />
(44–100 µm in diameter) with 1–4 secondaries and<br />
terminal discs; oxyhexactins and oxyhexasters (53–100<br />
µm in diameter).<br />
Zoogeographic distribution. North Pacific Ocean; locally<br />
common and abundant in some areas. In Alaska –<br />
Bering Sea <strong>to</strong> Sou<strong>the</strong>ast Alaska. Elsewhere – Japan, Kuril<br />
Islands, Sea <strong>of</strong> Okhotsk, British Columbia <strong>to</strong> Panama.<br />
Habitat. Aleutian Islands – attached <strong>to</strong> bedrock,<br />
cobbles, and pebbles, usually in low-relief habitats, at<br />
depths between 112 <strong>to</strong> 740 m. Bering Sea – attached <strong>to</strong><br />
pebbles and hexactinellid skele<strong>to</strong>ns at depths between<br />
375 and 522 m. Gulf <strong>of</strong> Alaska – on <strong>the</strong> continental shelf<br />
23<br />
and upper slope at depths between 70 and at least 259<br />
m. Observed in <strong>the</strong> glacial fjords <strong>of</strong> Sou<strong>the</strong>ast Alaska<br />
growing on bedrock at depths as shallow as 21 m. Elsewhere<br />
– reported at depths between 23 and 1103 m.<br />
Remarks. This is one <strong>of</strong> <strong>the</strong> most ecologically important<br />
<strong>sponges</strong> in Alaska. Juvenile golden king crabs<br />
(Lithodes aequispina) use <strong>the</strong> spongocoel as refuge habitat<br />
in <strong>the</strong> Aleutian Islands (S<strong>to</strong>ne, 2006). Due <strong>to</strong> its rigid<br />
skele<strong>to</strong>n, this species is an important structural component<br />
<strong>of</strong> <strong>the</strong> sponge reefs reported along <strong>the</strong> Pacific<br />
coast <strong>of</strong> Canada (Conway et al., 1991, 2005; Krautter et<br />
al., 2001) and recently in sou<strong>the</strong>rn Sou<strong>the</strong>ast Alaska.<br />
We have not been able <strong>to</strong> confirm <strong>the</strong> presence <strong>of</strong> <strong>the</strong><br />
o<strong>the</strong>r subspecies (H. calyx schulzei) in Alaskan <strong>water</strong>s.<br />
This species may be preyed upon by <strong>the</strong> sea star Henricia<br />
longispina in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Heterochone calyx<br />
calyx can be distinguished from <strong>the</strong> very similar Aphrocallistes<br />
vastus by <strong>the</strong> presence <strong>of</strong> pinular hexactins on <strong>the</strong><br />
inner (atrial) surface and by <strong>the</strong> lack <strong>of</strong> robust oxyhexasters<br />
with primary rays subsumed in a swollen centrum.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 172<br />
m in <strong>the</strong> central Aleutian Islands. A small stalked demosponge<br />
grows from inside <strong>the</strong> specimen. Grid marks<br />
are 1 cm 2 . 2) Specimen collected at a depth <strong>of</strong> 520 m<br />
in Zhemchug Canyon, Bering Sea. Grid marks are 1<br />
cm 2 . 3) Specimen at a depth <strong>of</strong> 180 m with a juvenile<br />
rosethorn rockfish (Sebastes helvomaculatus) and juvenile<br />
brown box crab (Lopholithodes foraminatus) in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska. 4) Specimen at a depth <strong>of</strong> 181 m in<br />
<strong>the</strong> eastern Gulf <strong>of</strong> Alaska. 5) Specimen at a depth <strong>of</strong><br />
190 m with a sharpchin rockfish (Sebastes zacentrus) and<br />
squat lobsters (Munida quadrispina) in <strong>the</strong> eastern Gulf<br />
<strong>of</strong> Alaska. Pho<strong>to</strong> by J. Lincoln Freese (AFSC).
24 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
10. Heterochone calyx calyx Schulze, 1886 (continued)
11. Regadrella okinoseana Ijima, 1896<br />
Description. This tube or sac sponge attaches basally<br />
<strong>to</strong> hard substrate. Lateral walls bear distinctive smooth<br />
depressions, each with a small (1–3 mm diameter) central<br />
hole (parietal osculum) connecting <strong>to</strong> <strong>the</strong> atrial cavity;<br />
edges around depressions <strong>of</strong>ten project several millimeters<br />
as parietal ledges; <strong>the</strong> large terminal osculum<br />
is normally covered by a coarse sieve plate and bordered<br />
by a flaring marginal cuff. Consistency is slightly compressible<br />
and rubbery. The partially collected Alaskan<br />
specimen lacked an oscular sieve plate, marginal cuff,<br />
and parietal ledges. Its atrial cavity was subdivided by<br />
longitudinal wall fusions. It is estimated <strong>to</strong> have been 25<br />
cm long by 13 cm diameter, but elsewhere incomplete<br />
specimens 48 cm long have been reported. Color in life<br />
is white; drab when dried or preserved in ethanol.<br />
Skeletal structure. While <strong>the</strong> Aleutian specimen may<br />
have unusual morphology, its spiculation is typical for<br />
<strong>the</strong> species. Its skele<strong>to</strong>n consists mainly <strong>of</strong> loose spicules;<br />
spicule fusion is present only in <strong>the</strong> lower parts collected<br />
and is assumed <strong>to</strong> have been extensive in <strong>the</strong> basal part<br />
left on <strong>the</strong> attachment site. Megascleres include thick<br />
principal diactins with rounded tips (5–16 mm long);<br />
thin diactins with pointed tips (1.3–7.5 mm long);<br />
sword-shaped dermal hexactins with short, tapered,<br />
pointed distal rays (89–369 µm length), tapered and<br />
pointed tangential rays (156–318 µm length), and long<br />
tapered proximal rays (232–807 µm length); regular<br />
hexactins (134–292 µm ray length); atrial pentactins<br />
with sharp tapered tangential rays (122–295 µm ray<br />
length) and longer proximal rays (137–563 µm ray<br />
length); a few atrial triactins and stauractins <strong>of</strong> similar<br />
shape and size; short, thick atrial diactins (347–1047 µm<br />
length); small diactins around parietal oscula (42–197<br />
µm length). Microscleres include mainly oxystaurasters<br />
(61–107 µm diameter); a few oxyhexasters (70–129 µm<br />
diameter); dermal floricomes (78–113 µm diameter)<br />
with 10–14 terminal rays ending in heads with 2–3 teeth;<br />
remnants <strong>of</strong> graphiocomes are common, with central<br />
(25–43 µm diameter) and terminal rays as dispersed<br />
thin raphides (141–237 µm long).<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – central Aleutian Islands. Elsewhere<br />
– reported from Indo-West Pacific Region including<br />
from near India, Northwest and South Australia, New<br />
Zealand, New Caledonia, Indonesia, Japan.<br />
25<br />
Habitat. In Alaska – occurs singly on muds<strong>to</strong>ne, bedrock,<br />
and possibly hexactinellid skele<strong>to</strong>ns at depths between<br />
1071 and 1395 m. Elsewhere – reported at depths<br />
between 390 and 1264 m.<br />
Remarks. The new record <strong>of</strong> this species in <strong>the</strong> Aleutian<br />
Islands represents a range extension <strong>of</strong> over 3700<br />
km from <strong>the</strong> nearest previous known site near Japan.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> preserved (frozen) specimen<br />
collected at a depth <strong>of</strong> 1386 m in <strong>the</strong> central Aleutian<br />
Islands. Grid marks are 1 cm 2 . 2) Same specimen as in<br />
pho<strong>to</strong> 1 in situ. The separation between <strong>the</strong> red laser<br />
marks is 10 cm.
26 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
12. Acanthascus (Acanthascus) pr<strong>of</strong>undum ssp. nov. Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. This s<strong>of</strong>t barrel sponge, attached basally,<br />
has very large surface conules lacking prostal spicules.<br />
Occasionally two tubes are attached near <strong>the</strong> base. The<br />
outer surface is smooth, its large canals covered by a<br />
very delicate spicule lattice. The large atrial canals are<br />
uncovered and <strong>the</strong> atrial surface is reflected out <strong>of</strong> <strong>the</strong><br />
large terminal osculum; <strong>the</strong>re are no marginal spicules.<br />
Height and diameter is <strong>to</strong> 25 cm. Color in life is white<br />
<strong>to</strong> creamy white.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are thin diactins with rough<br />
tips (0.94–19.8 mm); dermal finely rough pentactins<br />
with cylindric rays ending in rounded tips (113–271<br />
µm ray length); similar but hexactine atrial spicules<br />
(125–239 µm ray length). Microscleres are discoctasters<br />
(119–234 µm diameter), each primary ray bearing 5–10<br />
slightly curved terminals; oxyhexactins, hemioxyhexasters,<br />
and oxyhexasters (98–199 µm diameter). Microdiscohexasters<br />
are absent.<br />
Zoogeographic distribution. In Alaska – central Aleutian<br />
Islands. Known with certainty only from <strong>the</strong> single<br />
specimen collected southwest <strong>of</strong> Adak Island, but analysis<br />
<strong>of</strong> video records indicate that it is a relatively rare<br />
species in this area. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> bedrock, muds<strong>to</strong>ne, and large<br />
boulders at depths between 1446 and 2245 m.<br />
Remarks. This specimen is in <strong>the</strong> process <strong>of</strong> being<br />
described so data are incomplete; it differs from all<br />
named species <strong>of</strong> <strong>the</strong> genus in several characters, and<br />
from A. platei in particular, in lack <strong>of</strong> both prostalia and<br />
microdiscohexasters, and lack <strong>of</strong> loose spicule lattice<br />
over atrial canal apertures.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> preserved (frozen) specimen<br />
collected at a depth <strong>of</strong> 2105 m in <strong>the</strong> central Aleutian<br />
Islands. Grid marks are 1 cm 2 . 2) Same specimen as in<br />
pho<strong>to</strong> 1 in situ growing just behind Farrea kurilenis ssp.<br />
nov.
13. Acanthascus (Rhabdocalyptus) dawsoni dawsoni (Lambe, 1893)<br />
Description. S<strong>of</strong>t straight or curved tube or barrellike<br />
sac, occasionally partially divided in<strong>to</strong> two or three<br />
conjoined tubes, is attached basally <strong>to</strong> hard substrate.<br />
Surface is smooth but usually bearing a 1-cm tall veil <strong>of</strong><br />
pentactins and diactins which may be clean but usually<br />
covered with small epizoans and sediment; large single<br />
terminal osculum has marginal fringe. Consistency is<br />
s<strong>of</strong>t and compressible. Height is <strong>to</strong> 1 m and diameter <strong>to</strong><br />
30 cm. Color in life is white but <strong>of</strong>ten coated with sediment,<br />
epizoic organisms, and flocculent material, giving<br />
it a brown <strong>to</strong> greenish brown appearance.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are thick prostal diactins <strong>to</strong><br />
6 cm long; hypodermal pentactins with some smooth<br />
and some thorned tangential rays (0.9–4.1 mm long)<br />
and smooth proximal rays (0.9–4.1 mm long); principal<br />
diactins (4.2–11.4 mm long); thin short diactins<br />
(0.35–1.62 mm long); dermal stauractins (50–101 mm<br />
ray length), dermal pentactins with tangential rays<br />
(54–99 mm long) and proximal rays (44–100 mm long);<br />
atrial hexactins (55–177 mm ray length). Microscleres<br />
are oxyhemihexasters (48–104 mm in diameter); small<br />
discoctasters with straight terminals (56–82 mm in diameter);<br />
microdiscohexasters (17–28 mm in diameter).<br />
Zoogeographic distribution. Locally common and<br />
abundant. In Alaska – eastern Gulf <strong>of</strong> Alaska. Elsewhere<br />
– British Columbia <strong>to</strong> sou<strong>the</strong>rn California.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, cobbles, and<br />
pebbles at depths between 82 and 255 m. Elsewhere<br />
– attached <strong>to</strong> bedrock, cobbles, and pebbles on flat,<br />
inclined, or vertical surfaces (e.g., fjord walls), and following<br />
detachment may survive loose as roller <strong>sponges</strong><br />
(Reiswig, unpubl. data, 2010) at depths between 10 and<br />
437 m.<br />
Remarks. The species presently contains three subspecies:<br />
A. (R.) d. dawsoni (reviewed here); A. (R.) d. alascensis<br />
Wilson and Penney, 1930 (known only from Cape<br />
Spencer, Gulf <strong>of</strong> Alaska); and A. (R.) d. horridus Koltun,<br />
1967 (from <strong>the</strong> Bering Sea). A. (R.) d. dawsoni can be<br />
distinguished from A. solidus by <strong>the</strong> presence <strong>of</strong> at least<br />
some heavily thorned hypodermal pentactins; <strong>the</strong> latter<br />
species possesses only smooth hypodermal pentactins.<br />
27<br />
A. (R.) d. dawsoni is similar in appearance <strong>to</strong> A. mirabilis<br />
but likely differs in depth distribution and can be<br />
distinguished by its much smaller discoctasters (56–82<br />
vs. 144–180 µm diameter). We have observed a very<br />
similar species in <strong>the</strong> central Aleutian Islands at depths<br />
between about 400 and 1238 m attached <strong>to</strong> cobbles and<br />
hexactinellid skele<strong>to</strong>ns. This species is preyed upon by<br />
<strong>the</strong> sea star Poraniopsis inflata and possibly by <strong>the</strong> sea star<br />
Henricia longispina in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska.<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
168 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks are 1<br />
cm 2 . 2) Specimen (indicated by <strong>the</strong> white arrow) at a<br />
depth <strong>of</strong> 165 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska.
28 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
14. Acanthascus (Rhabdocalyptus) mirabilis (Schulze, 1899)<br />
Description. This species was known previously only<br />
from <strong>the</strong> holotype, which was <strong>the</strong> distal portion (about<br />
1/3 <strong>of</strong> <strong>to</strong>tal specimen) <strong>of</strong> a s<strong>of</strong>t saccate sponge. Surface<br />
is overtly smooth but evenly covered with small<br />
conic protuberances and bears a veil <strong>of</strong> pentactins and<br />
diactins; large single terminal osculum with marginal<br />
fringe. Consistency is s<strong>of</strong>t and compressible. From <strong>the</strong><br />
holotype fragment, <strong>the</strong> original specimen was thought<br />
<strong>to</strong> be ca. 30 cm in length and 15 cm in diameter with a<br />
1-cm thick wall; a new Aleutian specimen is 15.5 cm in<br />
length and 10.1 cm in diameter. Color in life is white;<br />
light brown when preserved in ethanol.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are thick prostal diactins <strong>to</strong><br />
2 cm long; hypodermal pentactins with mostly thorned<br />
paratropal tangential rays <strong>to</strong> 1 cm long; principal diactins<br />
(11.7–21.0 mm long); dermal diactins (plus a few<br />
stauractins and hexactins) (204–462 mm long); atrial<br />
hexactins (free ray 249–447 µm long, tangentials and<br />
parenchymal rays 153–316 mm long). Microscleres are<br />
oxyhexactins and a few oxyhexasters and hemioxyhexasters<br />
(72–232 mm in diameter with 33–87 mm long<br />
terminal rays); discoctasters (144–180 mm in diameter)<br />
with very short primary rays (20–28 mm long),<br />
each bearing 8–12 straight or s-curved, divergent terminal<br />
rays; microdiscohexasters mainly near or in <strong>the</strong><br />
dermal surface (21–40 mm in diameter).<br />
Zoogeographic distribution. Rare. Known only from<br />
Alaska – central Aleutian Islands and Gulf <strong>of</strong> Alaska.<br />
Elsewhere – not reported.<br />
Habitat. Central Aleutian Islands – occurs singly on<br />
bedrock at depths between 1984 and 2790 m. Sou<strong>the</strong>rn<br />
Alaska – attached <strong>to</strong> cobbles at depths near 1143 m.<br />
Remarks. Our specimen recently collected in <strong>the</strong><br />
Aleutian Islands is only <strong>the</strong> second known verified<br />
specimen <strong>of</strong> this species. It is very likely that A. (R.)<br />
unguiculatus Ijima, 1904 is a junior synonym <strong>of</strong> A. (R.)<br />
mirabilis. This species is similar in appearance <strong>to</strong> A.<br />
dawsoni but likely differs in depth distribution and<br />
can be distinguished by its much larger discoctasters<br />
(144–180 vs 56–82 µm diameter).<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> preserved (frozen) specimen<br />
collected at a depth <strong>of</strong> 2311 m in <strong>the</strong> central Aleutian<br />
Islands. Grid marks are 1 cm 2 . 2) Same specimen as in<br />
pho<strong>to</strong> 1 in situ. The separation between <strong>the</strong> red laser<br />
marks is 10 cm.
15. Acanthascus (Staurocalyptus) solidus (Schulze, 1899)<br />
Description. This compressible but very spiny sac<br />
or vasiform sponge attaches basally <strong>to</strong> hard substrate.<br />
External surface is covered by a dense veil <strong>of</strong> projecting<br />
pentactins in amongst <strong>the</strong> long prostal diactin needles,<br />
with a single large terminal osculum bordered by a<br />
marginal fringe <strong>of</strong> diactins. Consistency is s<strong>of</strong>t but spiky.<br />
Height is <strong>to</strong> 24 cm, diameter <strong>to</strong> 15 cm, and 23 mm in<br />
wall thickness. It may be found in clusters <strong>of</strong> up <strong>to</strong> nine<br />
individuals. Color in life is white but sometimes coated<br />
with sediment, epizoic organisms, and flocculent material,<br />
giving it a brown <strong>to</strong> greenish brown appearance;<br />
drab when preserved.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are thick prostal diactins <strong>to</strong><br />
4 cm long; hypodermal pentactins all have smooth or<br />
shagreened (never thorned) tangential rays (1.8–6.6<br />
mm long) and smooth proximal rays (5.5–9.6 mm<br />
long); principal diactins (2.0–8.4 mm long); dermal<br />
stauractins (75–180 mm ray length), dermal pentactins<br />
with tangential rays (88–193 mm long) and proximal<br />
rays (88–163 mm long); atrial hexactins (89–134 mm<br />
ray length). Microscleres are oxyhexasters and oxyhemihexasters<br />
(113–179 mm in diameter); discoctasters with<br />
straight terminals (134–225 mm in diameter); microdiscohexasters<br />
(16–22 mm in diameter).<br />
Zoogeographic distribution. North Pacific Ocean.<br />
Locally abundant. In Alaska – eastern Gulf <strong>of</strong> Alaska.<br />
Elsewhere – British Columbia <strong>to</strong> sou<strong>the</strong>rn California<br />
(Santa Maria Basin).<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, cobbles, and<br />
pebbles at depths between 82 and 255 m. Elsewhere –<br />
reported at depths between 91 and 1373 m.<br />
Remarks. Acanthascus solidus can be distinguished<br />
from <strong>the</strong> sometimes similar A. (R.) dawsoni dawsoni by<br />
its complete lack <strong>of</strong> thorned hypodermal pentactins<br />
that are always present and heavily thorned in <strong>the</strong> latter<br />
species. In <strong>the</strong> central Aleutian Islands a very similar<br />
species (possibly Acanthascus solidus but not confirmed)<br />
occurs at depths between 399 and 463 m. This species is<br />
preyed upon by <strong>the</strong> sea stars Hippasteria spp., Poraniopsis<br />
inflata, and possibly Henricia longispina in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska.<br />
Pho<strong>to</strong>s. 1) Mostly intact specimen collected at a depth<br />
<strong>of</strong> 167 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks are<br />
1 cm 2 . 2) Specimen (indicated by <strong>the</strong> white arrow) at a<br />
depth <strong>of</strong> 165 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska.<br />
29
30 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
16. Acanthascus (Staurocalyptus) sp. nov. 1 Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. This species grows from a narrow attached<br />
base <strong>to</strong> ei<strong>the</strong>r a thick massive or thick funnelshaped<br />
form. External surface has small rounded<br />
protuberances from each <strong>of</strong> which project several short<br />
diactin prostalia; <strong>the</strong>re is a single large terminal osculum.<br />
Consistency is s<strong>of</strong>t and easily <strong>to</strong>rn. Height is up <strong>to</strong><br />
24 cm and diameter <strong>to</strong> 17 cm. Color in life is brown <strong>to</strong><br />
golden brown.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are thick prostal diactins<br />
(4.3–10.1 mm long), a few rough, but not thorned,<br />
hypodermal pentactins (0.21–1.09 mm tangential ray<br />
length); principal diactins (1.11–6.84 mm long); dermal<br />
hexactins with some pentactins and stauractins<br />
(40–139 mm ray length); atrial hexactins (43–218 mm<br />
ray length). Microscleres are oxyhexactins and a few<br />
hemioxyhexasters (86–175 mm in diameter); very small<br />
discoctasters with terminals strongly curved outward<br />
(56–87 mm in diameter); microdiscohexasters are<br />
absent.<br />
Zoogeographic distribution. Locally abundant.<br />
In Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Attached <strong>to</strong> cobbles and pebbles at depths<br />
between 139 and 183 m.<br />
Remarks. This species is distinguished from all o<strong>the</strong>r<br />
species <strong>of</strong> Acanthascus (Staurcalyptus) by <strong>the</strong> small size<br />
and shape <strong>of</strong> its discoctasters. A complete description<br />
is in preparation.<br />
Pho<strong>to</strong>s. 1) Partially fragmented specimen collected<br />
with a trawl at a depth <strong>of</strong> 155 m in <strong>the</strong> central Aleutian<br />
Islands. Grid marks are 1 cm 2 . 2) Specimen at a depth<br />
<strong>of</strong> 165 m in <strong>the</strong> central Aleutian Islands.
17. Acanthascus (Staurocalyptus) sp. nov. 2 Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. S<strong>of</strong>t tube or sac sponge attaches basally<br />
<strong>to</strong> hard substrate. Lateral surface has low conules 15<br />
mm apart and 3–5 mm high from which small groups<br />
<strong>of</strong> thin, short, prostal diactin spicules project; <strong>the</strong>re is<br />
no veil <strong>of</strong> pentactins. The single terminal osculum is<br />
sharp-edged and without a marginal fringe. Both external<br />
and internal surfaces (and thus canal openings) are<br />
covered by tight spicule lattices. Consistency is s<strong>of</strong>t and<br />
pliable; internal aspect is wooly. The type specimen is 30<br />
cm tall by 20 cm diameter (at <strong>the</strong> widest point). Color<br />
in life is white.<br />
Skeletal structure. It is composed entirely <strong>of</strong> loose<br />
spicules excepting a small fused basal attachment<br />
structure. Megascleres are prostal diactins 9.3–16.2<br />
mm long; principal diactins 1.8–10.4 mm long; dermal<br />
pentactins with spiny tangential rays (72–201 µm<br />
long) and proximal rays (65–172 µm long); atrial spiny<br />
hexactins (65–172 µm ray length). Microscleres include<br />
two classes <strong>of</strong> discoctasters (124–382 and 112–167 µm<br />
diameter); s<strong>to</strong>ut oxyhexactins (94–139 µm diameter);<br />
oxy- and hemioxyhexasters (88–133 µm diameter); microdiscohexasters<br />
(13–22 µm diameter).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – known only from <strong>the</strong> central Aleutian Islands.<br />
Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> cobbles in low-relief habitat at a<br />
depth <strong>of</strong> 711 m. Video records suggest it is also locally<br />
common on bedrock, muds<strong>to</strong>ne, and cobbles in moderate-relief<br />
habitat at depths between 190 and 1556 m.<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
711 m in <strong>the</strong> central Aleutian Islands. Grid marks are 1<br />
cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 showing <strong>the</strong> detail<br />
<strong>of</strong> <strong>the</strong> osculum. 3) Same specimen as in pho<strong>to</strong>s 1 and<br />
2 in situ. The separation between <strong>the</strong> red laser marks<br />
is 10 cm.<br />
31
32 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
18. Aulosaccus pinularis Okada, 1932<br />
Description. Vase-shaped sponge attaches basally <strong>to</strong><br />
solid substrate, thickest near its upper end. External surface<br />
is smooth and completely lacking large projecting<br />
spicules; both external and internal surfaces are lined by<br />
a lattice <strong>of</strong> loose spicules; <strong>the</strong>re is a single large terminal<br />
osculum without a marginal spicule fringe. Consistency<br />
is very s<strong>of</strong>t and easily <strong>to</strong>rn. Height is <strong>to</strong> 24 cm, diameter<br />
<strong>to</strong> 15 cm, and 23 mm in wall thickness. Color in life is<br />
white; drab when preserved.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are thick diactins (4.4–5.6<br />
mm long); thin diactins (1.3–5.5 mm long); pinular dermal<br />
hexactins with projecting pinular ray (109–175 mm<br />
long), tangential rays (103–150 mm long) and proximal<br />
ray (86–140 mm long); pinular atrial hexactins with projecting<br />
pinular ray (93–300 mm long), tangential rays<br />
(109–214 mm long), and proximal rays (83–192 mm<br />
long). Microscleres include very large discasters, <strong>of</strong>ten<br />
called ”solasters” (193–438 mm in diameter) with fused<br />
primary rays recognizable as six hemispherical bosses;<br />
oxyhexasters and hemioxyhexasters (99–140 mm in diameter);<br />
oxyhexactins (83–166 mm in diameter); small<br />
spherical discohexasters (26–36 mm in diameter).<br />
Zoogeographic distribution. Rare. In Alaska – central<br />
Aleutian Islands. Elsewhere – western Bering Sea,<br />
Kuril Islands and <strong>of</strong>f <strong>the</strong> sou<strong>the</strong>rn tip <strong>of</strong> <strong>the</strong> Kamchatka<br />
Peninsula.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, muds<strong>to</strong>ne,<br />
or boulders at depths between 843 and 1715 m. Elsewhere<br />
– reported at a depth <strong>of</strong> 117 m but most collections<br />
do not report depth.<br />
Pho<strong>to</strong>s. 1) Mostly intact specimen collected at a depth<br />
<strong>of</strong> 843 m in <strong>the</strong> central Aleutian Islands. Grid marks<br />
are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ. Note<br />
that <strong>the</strong> specimen has been <strong>to</strong>rn on <strong>the</strong> left side (prior<br />
<strong>to</strong> collection) and <strong>the</strong> osculum is directed <strong>to</strong> <strong>the</strong> right.
19. Aulosaccus schulzei Ijima, 1896<br />
Description. Vase-shaped sponge attaches basally <strong>to</strong><br />
solid substrate, thickest near its upper end. External<br />
surface is smooth, completely lacking large projecting<br />
spicules, and lined by a lattice <strong>of</strong> loose spicules. The<br />
internal atrial surface has a network <strong>of</strong> diactin bundles<br />
crossing <strong>the</strong> exhalant apertures, but a lattice <strong>of</strong> atrialia<br />
is absent. There is a single large terminal osculum lacking<br />
a marginal spicule fringe. Consistency is very s<strong>of</strong>t<br />
and easily <strong>to</strong>rn. Height is up <strong>to</strong> 45 cm, diameter <strong>to</strong> 22.5<br />
cm, and 34 mm in wall thickness. Color in life is creamy<br />
white; drab when preserved.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely <strong>of</strong><br />
loose spicules. Megascleres are hypodermal pentactins<br />
and a few triactins and tetractins (0.6–3.4 mm long<br />
tangential rays; 0.9–4.4 mm long proximal rays); thick<br />
diactins (3.3–17.7 mm long); thin diactins (1.4–6.7 mm<br />
long); short atrial diactins (0.4–1.5 mm long); dermalia<br />
are a mixture <strong>of</strong> stauractins, pentactins, and hexactins<br />
(89–181 mm ray length); atrialia are mainly very large<br />
hexactins with some pentactins (224–1186 mm ray<br />
length). Microscleres include very large discasters, <strong>of</strong>ten<br />
called “solasters” (513–1389 mm in diameter), with<br />
primary rays fused in<strong>to</strong> a slightly irregular sphere; very<br />
thin oxyhexactins, with irregular variants, and hemioxyhexasters<br />
(87–165 mm in diameter); small spherical<br />
discohexasters (20–39 mm in diameter).<br />
Zoogeographic distribution. Apparently a rare species.<br />
In Alaska – central Aleutian Islands and Bering<br />
Sea (Pribil<strong>of</strong> Canyon). Elsewhere – Japan, Kuril Islands,<br />
Okhotsk Sea, and <strong>of</strong>f sou<strong>the</strong>rn California.<br />
Habitat. In Alaska – in <strong>the</strong> central Aleutian Islands<br />
it occurs on bedrock, boulders, and cobbles at depths<br />
between 1270 and 1350 m. In Pribil<strong>of</strong> Canyon it occurs<br />
at a depth <strong>of</strong> 300 m. Elsewhere – reported at depths<br />
between 117 and 419 m.<br />
Remarks. Associated fauna include juvenile lithodid<br />
crabs (Paralomis verrilli and Lithodes couesi), panda-<br />
lid shrimps, and <strong>the</strong> large ophiuroid Gorgonocephalus<br />
eucnemis.<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
300 m in Pribil<strong>of</strong> Canyon, Bering Sea. Grid marks are 1<br />
cm 2 . 2) Specimen collected at a depth <strong>of</strong> 1320 m in <strong>the</strong><br />
central Aleutian Islands. 3) Same specimen as in pho<strong>to</strong><br />
2 in situ. Two pairs <strong>of</strong> red lasers each separated by 10 cm.<br />
33
34 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
20. Bathydorus sp., description by Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. Very small flattened ovoid sponge attaches<br />
<strong>to</strong> a cobble. External surface is smooth and covered<br />
by a lattice <strong>of</strong> mainly stauractin dermalia; about one-half<br />
<strong>of</strong> <strong>the</strong> surface is pr<strong>of</strong>usely spined by long diactins and<br />
sports a veil <strong>of</strong> raised pentactins. An osculum and atrial<br />
cavity could not be distinguished. Consistency is s<strong>of</strong>t and<br />
delicate. The specimen is 11 by 8 by 2 mm. It is transparent<br />
and colorless.<br />
Skeletal structure. Skele<strong>to</strong>n is composed entirely<br />
<strong>of</strong> loose spicules, but parts <strong>of</strong> <strong>the</strong> fused basidictyonal<br />
skele<strong>to</strong>n remain attached. Megascleres are hypodermal<br />
pentactins (0.26–1.4 mm long tangential rays; 0.47–1.55<br />
mm long proximal rays); thick prostal diactins (3.2–16.8<br />
mm long); thin diactins (0.96–6.82 mm long); dermal<br />
stauractins and diactins (81–381 mm ray length). Microscleres<br />
consist only <strong>of</strong> hemioxyhexasters (112–181<br />
mm in diameter) and oxyhexactins (124–204 mm in<br />
diameter).<br />
Zoogeographic distribution. Rare. In Alaska – central<br />
Aleutian Islands. Elsewhere – small incompletely<br />
identifiable Bathydorus spp. have been reported worldwide.<br />
Habitat. In Alaska – occurs on cobble but probably<br />
also on boulders and bedrock; collected from 494 m but<br />
may have a broad depth range. Elsewhere – small Bathydorus<br />
spp. are reported from depths <strong>of</strong> 446 <strong>to</strong> 1625 m.<br />
Pho<strong>to</strong>. 1) Whole specimen collected at a depth <strong>of</strong> 494<br />
m in <strong>the</strong> central Aleutian Islands.
21. Caulophacus (Caulophacus) sp. nov. Reiswig and S<strong>to</strong>ne, in preparation<br />
Description. A very large mushroom-shaped body<br />
on a long cylindrical stalk attaches basally <strong>to</strong> hard substrate.<br />
The inhalant surface occupies <strong>the</strong> upper stalk<br />
and underside <strong>of</strong> <strong>the</strong> body; <strong>the</strong> exhalant surface on <strong>the</strong><br />
upper body is reflected around <strong>the</strong> edges <strong>to</strong> <strong>the</strong> sharp<br />
margin on <strong>the</strong> under edges <strong>of</strong> <strong>the</strong> body; <strong>the</strong>re is no distinct<br />
osculum. All surfaces are macroscopically smooth<br />
with large canals covered by lattices <strong>of</strong> loose spicules.<br />
Consistency <strong>of</strong> <strong>the</strong> body is compressible but firm; <strong>the</strong><br />
stalk is s<strong>to</strong>ny hard due <strong>to</strong> fusion <strong>of</strong> <strong>the</strong> internal spicules<br />
and is hollow, with two main internal longitudinal<br />
canals. The specimen, measured dry, is 75 cm in <strong>to</strong>tal<br />
height, <strong>the</strong> body is 31 by 40 cm transversely, and <strong>the</strong><br />
stalk is 68 cm long and 2.1 cm in diameter (narrowest<br />
point, 1.8 cm). Color in life is white <strong>to</strong> creamy white;<br />
drab when preserved.<br />
Skeletal structure. Spicules in <strong>the</strong> main body remain<br />
loose, but <strong>the</strong> principal diactins in <strong>the</strong> stalk are fused<br />
<strong>to</strong> form a hard, rigid, and long-lasting structure. Megascleres<br />
include pinular hexactine dermalia (132–337<br />
µm pinular ray length, 90–163 µm tangential ray length,<br />
102–158 µm proximal ray length); supported by hypodermal<br />
pentactins (315–862 µm tangential ray length,<br />
312–1054 µm proximal ray length); atrialia are slightly<br />
thinner pinular hexactins (131–379 µm pinular ray<br />
length, 80–167 µm tangential ray length, 88–142 µm<br />
proximal ray length); supported by hypoatrial pentactins<br />
similar <strong>to</strong> those <strong>of</strong> <strong>the</strong> dermal side; choanosomal<br />
diactins are mainly in loose bundles (1.65–4.06 mm<br />
long); large choanosomal oxyhexactins are common<br />
(678–1579 µm ray length). Microscleres are all coarsely<br />
spined, including hemidiscohexasters and discohexasters<br />
(82–177 µm diameter) and discohexactins (90–185<br />
µm diameter).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Occurs singly on bedrock and large boulders<br />
at depths between 1326 and 2680 m.<br />
Remarks. The Aleutian Caulophacus (Caulophacus)<br />
specimen has been compared <strong>to</strong> <strong>the</strong> 18 known species<br />
<strong>of</strong> <strong>the</strong> subgenus and found <strong>to</strong> be unassignable <strong>to</strong> any <strong>of</strong><br />
<strong>the</strong>m. Its description as a new species is in progress. It<br />
is used as a perch by several species <strong>of</strong> lithodid crabs,<br />
including Paralomis verrilli.<br />
Pho<strong>to</strong>s. 1) Whole specimen collected at a depth <strong>of</strong><br />
1806 m in <strong>the</strong> central Aleutian Islands. 2) Same specimen<br />
as in pho<strong>to</strong> 1. 3) Same specimen as in pho<strong>to</strong>s 1<br />
and 2 in situ with Paralomis verrilli crabs. The separation<br />
between <strong>the</strong> red laser marks is 10 cm.<br />
35
37<br />
CLASS DEMOSPONGIAE
38 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
22. Plakina atka Lehnert, S<strong>to</strong>ne and Heimler, 2005<br />
Description. Sponge is encrusting. Surface is convoluted<br />
but less distinctively so than Plakina tanaga. Single<br />
strands have a smooth surface, not microtuberculate.<br />
This sponge may cover large areas (up <strong>to</strong> 30 cm 2 ) with<br />
a thickness <strong>of</strong> approximately 0.3 <strong>to</strong> 0.8 cm. The ec<strong>to</strong>some<br />
is reddish brown (Aleutian Islands), beige, or light<br />
brown (eastern Gulf <strong>of</strong> Alaska).<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Ec<strong>to</strong>somal dense spicule crust averages<br />
100 µm in thickness. Choanosome is somewhat<br />
less densely packed with spicules, roughly arranged in<br />
tracts <strong>of</strong> varying orientation and with many spicules in<br />
between. Characteristic plakinid oxeas are slightly bent<br />
and thickest in <strong>the</strong> center (70–108 × 3–6 µm). A thicker<br />
category <strong>of</strong> diods, always with relatively long spines at<br />
<strong>the</strong> center, is probably derived from <strong>the</strong> spined category<br />
<strong>of</strong> triods (82–95 × 8–10 µm). Triods also occur in two<br />
distinct types: relatively rare simple triaxons (28–33 µm/<br />
ray) and much more abundant, robust triaxons with a<br />
row <strong>of</strong> large spines on each ray close <strong>to</strong> <strong>the</strong> center <strong>of</strong> <strong>the</strong><br />
spicule (23–40 × 3–6 µm/ray). Calthrops are rare and<br />
have a reduced fourth ray only. Tetralophose calthrops<br />
have tetrafurcate, occasionally pentafurcate rays, with<br />
micr<strong>of</strong>urcate ends (18–23 µm in <strong>to</strong>tal length).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – central Aleutian Islands, eastern Gulf <strong>of</strong> Alaska<br />
(continental shelf <strong>of</strong>f Cape Ommaney, Baran<strong>of</strong> Island),<br />
and sou<strong>the</strong>rn Sou<strong>the</strong>ast Alaska (Portland Canal). Elsewhere<br />
– nor<strong>the</strong>rn British Columbia (Portland Canal).<br />
Habitat. Encrusts bedrock, boulders, cobbles, pebbles,<br />
and hexactinellid skele<strong>to</strong>ns at depths between 82–180 m<br />
(Aleutian Islands) and 95–253 m in <strong>the</strong> eastern Gulf <strong>of</strong><br />
Alaska. May cover relatively large areas (up <strong>to</strong> 30 cm 2 ).<br />
Remarks. Plakina atka can be distinguished from<br />
<strong>the</strong> similar P. tanaga by its relatively smooth surface<br />
compared <strong>to</strong> <strong>the</strong> strongly convoluted and microtuberculated<br />
surface <strong>of</strong> <strong>the</strong> latter species, and by <strong>the</strong> presence<br />
<strong>of</strong> tetralophate lophocalthrops compared <strong>to</strong> <strong>the</strong><br />
trilophose lophocalthrops <strong>of</strong> P. tanaga.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 118 m in<br />
<strong>the</strong> central Alautian Islands. Grid marks are 1 cm 2 . 2)<br />
Specimen collected at a depth <strong>of</strong> 167 m in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska. 3) Specimen at a depth <strong>of</strong> 118 m in <strong>the</strong><br />
central Aleutian Islands.
23. Plakina tanaga Lehnert, S<strong>to</strong>ne and Heimler, 2005<br />
Description. Sponge is encrusting. Convoluted surface<br />
has <strong>deep</strong> grooves between strands with a microtuberculated<br />
surface. Consistency is cheese-like. Circular<br />
oscula flush with <strong>the</strong> surface are visible in situ. This<br />
sponge may cover large areas (up <strong>to</strong> 1 m 2 ) with a thickness<br />
<strong>of</strong> approximately 1 <strong>to</strong> 1.5 cm. Color in life is beige<br />
<strong>to</strong> light brown.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some is packed with lophocalthrops<br />
with a layer 20–250 µm thick. Spicules are<br />
characteristically bent diods, <strong>of</strong>ten centrotylote or with<br />
a reduced third ray in <strong>the</strong> form <strong>of</strong> a spine (85–97 ×<br />
2–4 µm); two categories <strong>of</strong> triaxons, smooth, thinner<br />
triods (24–50 × 2–5 µm/ray) and thicker triods, <strong>of</strong>ten<br />
with one short, slender spine near <strong>the</strong> base <strong>of</strong> each ray<br />
(22–38 × 7–9 µm). Rare smooth calthrops have rays <strong>of</strong><br />
<strong>the</strong> same dimensions as those <strong>of</strong> <strong>the</strong> triods. There are<br />
abundant trilophose calthrops with each ray tetrafurcate<br />
and, again micr<strong>of</strong>urcate at <strong>the</strong> points, somewhat smaller<br />
than <strong>the</strong> triods and concentrated in <strong>the</strong> ec<strong>to</strong>some. Total<br />
length <strong>of</strong> lophocalthrops is 28–42 µm. Non-lophose rays<br />
are sometimes bifurcate.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
known only from <strong>the</strong> type locality in Little Tanaga Strait,<br />
central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Encrusts vertical and overhanging surfaces <strong>of</strong><br />
bedrock and boulders at depths between 140 and 383 m.<br />
May cover relatively large areas (up <strong>to</strong> 1 m 2 ).<br />
39<br />
Remarks. Plakina tanaga can be distinguished from<br />
<strong>the</strong> similar P. atka by its strongly convoluted and microtuberculated<br />
surface compared <strong>to</strong> <strong>the</strong> relatively smooth<br />
surface <strong>of</strong> <strong>the</strong> latter species, and by <strong>the</strong> presence <strong>of</strong> trilophose<br />
lophocalthrops compared <strong>to</strong> <strong>the</strong> tetralophate<br />
lophocalthrops <strong>of</strong> P. atka.<br />
Pho<strong>to</strong>. 1) P. tanaga encrusting a bedrock scarp at a<br />
depth <strong>of</strong> 146 m in <strong>the</strong> central Aleutian Islands.
40 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
24. Craniella arb (de Laubenfels, 1930)<br />
Description. Sponge is globular <strong>to</strong> subglobular. Oscula<br />
are occasionally present but not obvious. Spicules<br />
radiate from <strong>the</strong> center <strong>to</strong> <strong>the</strong> surface and protrude<br />
above it. Consistency is hard, cartilaginous, and only<br />
slightly elastic due <strong>to</strong> <strong>the</strong> high spicule density. With a<br />
diameter <strong>of</strong> 8 <strong>to</strong> 10 cm, this is one <strong>of</strong> <strong>the</strong> largest species<br />
<strong>of</strong> Craniella. The cortex is white; <strong>the</strong> interior <strong>of</strong> <strong>the</strong><br />
sponge is orange-gold.<br />
Skeletal structure. Protriaenes may reach a length <strong>of</strong><br />
more than 3 cm. Protriaenes and anatriaenes have relatively<br />
small clads; protriaenes, rhabd greater than 3 cm<br />
× 3–11 µm, clad 20–30 µm; anatriaenes, rhabd up <strong>to</strong> 10<br />
mm × 3–11 µm, clads, 40–85 µm, oxeas, up <strong>to</strong> 3 cm ×<br />
10–40 µm, commata, 7–10 µm.<br />
Zoogeographic distribution. Uncommon, but locally<br />
patchy. In Alaska – Aleutian Islands. Elsewhere – central<br />
California <strong>to</strong> <strong>the</strong> Gulf <strong>of</strong> California.<br />
Habitat. In Alaska – found at depths between 88 and<br />
272 m in boulder and cobble habitats. Typically epizoic<br />
on calcareous <strong>sponges</strong> (e.g., Leucandra poculiformis) and<br />
various demo<strong>sponges</strong>, including Tedania dirhaphis,<br />
Mycale jasoniae, and M. loveni. Elsewhere – reported at<br />
depths between 11 and 214 m.<br />
Remarks. Reproduction in this group <strong>of</strong> <strong>sponges</strong> is<br />
oviparous without a larval stage, or viviparous with production<br />
<strong>of</strong> young adults within <strong>the</strong> parent (Van Soest<br />
and Rützler, 2002).<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 100 m<br />
in <strong>the</strong> central Aleutian Islands. Specimen is attached<br />
<strong>to</strong> <strong>the</strong> demosponge Tedania dirhapsis. Grid marks are<br />
1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 showing radial<br />
arrangement <strong>of</strong> large spicules in <strong>the</strong> interior and <strong>the</strong><br />
presence <strong>of</strong> orange-colored eggs or young adults.<br />
3) Specimen at a depth <strong>of</strong> 170 m in <strong>the</strong> central Aleutian<br />
Islands.
25. Craniella sigmoancoratum (Koltun, 1966)<br />
Description. Body is globular with very long tracts<br />
<strong>of</strong> protruding spicules. Typically is attached <strong>to</strong> o<strong>the</strong>r<br />
<strong>sponges</strong> via <strong>the</strong>se tracts which terminate in anchorlike,<br />
long-shafted triaenes. Internal bundles <strong>of</strong> spicules<br />
radiate from <strong>the</strong> center <strong>of</strong> <strong>the</strong> globular body <strong>to</strong> well<br />
beyond <strong>the</strong> surface <strong>of</strong> <strong>the</strong> sponge. The surface consists<br />
<strong>of</strong> a layer <strong>of</strong> dermal oxeas occurring only <strong>the</strong>re and<br />
oriented perpendicular <strong>to</strong> <strong>the</strong> surface. The diameter<br />
<strong>of</strong> <strong>the</strong> body is 7 <strong>to</strong> 8 mm (without protruding spicules).<br />
Spicule tracts protrude 6–8 mm above <strong>the</strong> surface, giving<br />
<strong>the</strong> sponge a spiny appearance. Color in life is white<br />
or creamy white.<br />
Skeletal structure. Large oxeas have unequal ends<br />
(3000–8000 × 45–72 µm), cortical oxeas (460–1340<br />
× 26–76 µm), protriaenes (1600–7300 µm long with<br />
clads 13–34 µm), anatriaenes have rhabds (2300–8700<br />
µm with clads 19–40 µm). The sigmaspires are unique<br />
among <strong>the</strong> known species <strong>of</strong> Craniella in that <strong>the</strong>y are<br />
relatively large and resemble isochelae (22–34 µm).<br />
These sigmaspires are abundant and densely present in<br />
<strong>the</strong> cortical layer.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
central Aleutian Islands, Bering Sea (Pribil<strong>of</strong> Canyon).<br />
Elsewhere – previously known only from <strong>the</strong> original<br />
description in <strong>the</strong> Kuril Islands, Russia.<br />
41<br />
Habitat. In Alaska – a cryptic species; epizoic on<br />
hexactinellid <strong>sponges</strong> and o<strong>the</strong>r demo<strong>sponges</strong>, including<br />
Erylus aleuticus, at depths between 190 and 275 m.<br />
Elsewhere – no information available.<br />
Remarks. Previously known as Tetilla sigmoancoratum<br />
Koltun, 1966, but recently transferred <strong>to</strong> <strong>the</strong> genus<br />
Craniella since it possesses both a cortex and <strong>the</strong> special<br />
cortical oxeas that are lacking in <strong>the</strong> genus Tetilla.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 208 m in<br />
Pribil<strong>of</strong> Canyon, Bering Sea. Specimen is growing on<br />
an unidentified hexactinellid sponge.
42 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
26. Craniella spinosa Lambe, 1893<br />
Description. Sponge is globular with many short,<br />
sharp spines. It typically attaches <strong>to</strong> o<strong>the</strong>r <strong>sponges</strong> via<br />
<strong>the</strong> spines. Characteristic <strong>of</strong> <strong>the</strong> genus, <strong>the</strong> internal arrangement<br />
<strong>of</strong> spicule bundles is strictly radial with bundles<br />
supporting <strong>the</strong> ec<strong>to</strong>somal conules, which are less<br />
than 1 mm in height. The surface is formed by a special<br />
cortex. Dermal oxeas form a palisade perpendicular<br />
<strong>to</strong> <strong>the</strong> surface. Spicule bundles protrude through <strong>the</strong><br />
conules and diverge. Diameter is <strong>to</strong> 16–18 mm. Color<br />
in life is white or creamy white.<br />
Skeletal structure. There are large oxeas (2300–4800<br />
× 23–42 µm) with unequal ends, both ends acute but<br />
one end filiform, cortical oxeas (685–1230 × 35–47<br />
µm), protriaenes (3400–5700 µm long), anatriaenes<br />
(up <strong>to</strong> 8000 µm long rhabdomes). Microscleres are<br />
c- and s-shaped sigmaspires with a maximum diameter<br />
<strong>of</strong> 11–14 µm.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
Bering Sea (Pribil<strong>of</strong> Canyon). Elsewhere – originally described<br />
from Vancouver Island. Reportedly occurs from<br />
<strong>the</strong> Aleutian Islands <strong>to</strong> British Columbia (Austin, 1985).<br />
Habitat. In Alaska – a cryptic species; epizoic on<br />
Myxilla pedunculata and probably o<strong>the</strong>r <strong>sponges</strong> at a<br />
depth <strong>of</strong> 236 m. Elsewhere – depths ranging from 36 <strong>to</strong><br />
76 m (Vancouver Island).<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 236 m in<br />
Pribil<strong>of</strong> Canyon, Bering Sea. Specimen is attached <strong>to</strong><br />
<strong>the</strong> demosponge Myxilla pedunculata.
27. Craniella sputnika Lehnert and S<strong>to</strong>ne, 2011<br />
Description. Sponge is globular with numerous acute<br />
spines distributed over <strong>the</strong> surface. The surface between<br />
<strong>the</strong> spines is smooth, without any visible apertures.<br />
Characteristic <strong>of</strong> <strong>the</strong> genus, <strong>the</strong> internal arrangement <strong>of</strong><br />
spicule bundles is strictly radial with bundles supporting<br />
<strong>the</strong> ec<strong>to</strong>somal conules. The sponge is ra<strong>the</strong>r hard, only<br />
slightly elastic. Diameter including <strong>the</strong> spines is <strong>to</strong> 30<br />
mm (16 mm without <strong>the</strong> spines). Color in life is white<br />
<strong>to</strong> creamy white.<br />
Skeletal structure. Spicules are anatriaenes (3430–<br />
8820 × 12–21 µm), clads (48–154 × 17–22 µm per ray),<br />
and protriaenes (4520–8960 × 12–40 µm). There are<br />
clads (50–250 × 5–17 µm per ray); choanosomal oxeas<br />
(4530–5425 × 50–75 µm); two categories <strong>of</strong> cortical oxeas:<br />
small, centrotylote oxeas (97–372 × 8–17 µm), tyle<br />
in <strong>the</strong> center more like a ring than a tyle, and larger<br />
ones (540-987 × 28–63 µm). Sigmaspires or similar sigmoid<br />
spicules are absent.<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– known only from <strong>the</strong> type locality in Amchitka Pass,<br />
central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Patchy distribution. A cryptic species; epizoic<br />
on demo<strong>sponges</strong>, including Myxilla sp., and occasionally<br />
gorgonian corals (Primnoidae) in boulder, cobble,<br />
and sand habitats at depths between 115 and 199 m.<br />
Pho<strong>to</strong>s. 1) Specimens collected at a depth <strong>of</strong> 115 m<br />
in <strong>the</strong> central Aleutian Islands. 2) Two specimens (indicated<br />
by <strong>the</strong> white arrows) at a depth <strong>of</strong> 191 m in <strong>the</strong><br />
central Aleutian Islands.<br />
43
44 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
28. Erylus aleuticus Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. Lobate sponge consists <strong>of</strong> several flattened<br />
lobes (oval <strong>to</strong> circular). The surface is hard due<br />
<strong>to</strong> an ec<strong>to</strong>somal crust <strong>of</strong> aspidasters about 0.5 mm in<br />
thickness. Numerous uniporal orifices are scattered<br />
over <strong>the</strong> surface at intervals <strong>of</strong> 0.5–1.0 mm. The orifices<br />
are circular, 0.5–1.0 mm in diameter, and slightly (
29. Geodia lendenfeldi nomen novum<br />
Description. Small, cylindrical sponge has an ec<strong>to</strong>somal<br />
hard cortex <strong>of</strong> sterrasters and a s<strong>of</strong>ter choanosome.<br />
Lendenfeld (1910) described <strong>the</strong> species as encrusting,<br />
cushion-shaped, or irregularly finger-shaped. The specimen<br />
collected in <strong>the</strong> Aleutian Islands was a single cylinder<br />
approximately 5 cm in height and 1 cm in width.<br />
Color in life is white <strong>to</strong> creamy white.<br />
Skeletal structure. There are sterrasters (170–200 ×<br />
130-140 µm), choanosomal styles and oxeas (1200–1760<br />
× 30–40 µm), diaenes <strong>of</strong> <strong>the</strong> same size as oxeas, and<br />
tylasters (10 µm). Triaenes are absent.<br />
Zoogeographic distribution. Rare. In Alaska – Aleutian<br />
Islands and Sou<strong>the</strong>ast Alaska. Elsewhere – previously<br />
known from <strong>the</strong> west coast <strong>of</strong> North America from<br />
Sou<strong>the</strong>ast Alaska <strong>to</strong> sou<strong>the</strong>rn California.<br />
Habitat. In Alaska – found in cobble, pebble, and sand<br />
habitat at a depth <strong>of</strong> 190 m. Epizoic on <strong>the</strong> demosponge<br />
Erylus aleuticus. Elsewhere – no information available.<br />
Remarks. Triaenes are absent in G. robusta, which<br />
prompted Lendenfeld (1910) <strong>to</strong> erect <strong>the</strong> genus Geod-<br />
45<br />
inella that is now regarded as synonymous with Geodia.<br />
This specimen is <strong>the</strong> same as Geodia robusta (Lendenfeld,<br />
1910). G. robusta (Lendenfeld, 1910) is a junior homonym<br />
<strong>of</strong> G. robusta (Lendenfeld, 1907) however, and<br />
according <strong>to</strong> <strong>the</strong> World Porifera Database (Van Soest et<br />
al., 2008) it has no valid name. Thus, we suggest renaming<br />
G. robusta (Lendenfeld, 1910) as Geodia lendenfeldi.<br />
Pho<strong>to</strong>. 1) Fragments <strong>of</strong> specimen collected at a depth<br />
<strong>of</strong> 190 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 .
46 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
30. Poecillastra tenuilaminaris (Sollas, 1886)<br />
Description. This sponge is polymorphic but typically<br />
has a plate-like or lamellar growth form. The surface <strong>of</strong><br />
<strong>the</strong> sponge is smooth <strong>to</strong> <strong>the</strong> unaided eye but microscopically<br />
hispid. Small oscula occur on one side only. The<br />
consistency is firm, incompressible, and fragile. Size is<br />
highly variable but attains a length and height exceeding<br />
more than 1 m. Color in life is whitish <strong>to</strong> light brown,<br />
<strong>of</strong>ten with a dark fringe at <strong>the</strong> distal edge.<br />
Skeletal structure. The choanosome consists <strong>of</strong> an<br />
unorganized mix <strong>of</strong> calthrops and oxeas: calthrops<br />
(320–540 × 35–48 µm per ray), occasionally triods with<br />
rays <strong>of</strong> <strong>the</strong> same size, oxeas (1240–2650 × 32–47 µm).<br />
Microscleres are streptasters (14–28 µm) and acanthose<br />
microxeas (117–157 × 3–5 µm) that are always bent in<br />
<strong>the</strong> middle.<br />
Zoogeographic distribution. Common. In Alaska –<br />
Bering Sea (Zhemchug Canyon) and eastern Gulf <strong>of</strong><br />
Alaska. May reach densities up <strong>to</strong> 19 per m 2 in eastern<br />
Gulf <strong>of</strong> Alaska habitats. Elsewhere – holotype described<br />
from Sea <strong>of</strong> Japan, also reported from along <strong>the</strong> California<br />
Coast and <strong>the</strong> Gulf <strong>of</strong> California. Present records<br />
strongly suggest a circumboreal distribution in <strong>the</strong><br />
North Pacific. However, morphological differences do<br />
exist between populations, suggesting some genetic<br />
isolation (see Remarks).<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock at depths<br />
between 149 and 486 m in <strong>the</strong> Bering Sea and attached<br />
<strong>to</strong> bedrock, boulders, and cobbles at depths between<br />
71 and 255 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Elsewhere<br />
– reported at depths <strong>to</strong> 74 m.<br />
Remarks. Previously known as Normania tenuilaminaris<br />
Sollas 1886, P. tenuilaminaris was originally described as<br />
having straight acanthose microxeas. The microxeas we<br />
examined from <strong>the</strong> Bering Sea specimens were consistently<br />
bent but this is <strong>the</strong> only difference between <strong>the</strong><br />
specimens and in our opinion not sufficient <strong>to</strong> separate<br />
<strong>the</strong> specimens at <strong>the</strong> species level. However, it is probable<br />
that <strong>the</strong> Bering Sea population is genetically isolated<br />
and may have developed different morphological<br />
characters, such as <strong>the</strong> bent microxeas. This species is<br />
preyed upon by <strong>the</strong> sea stars Hippasteria spp., Poraniopsis<br />
inflata, Pteraster tesselatus, Ceramaster patagonicus, and<br />
possibly Henricia longispina. We have observed a very<br />
similar sponge in <strong>the</strong> central Aleutian Islands at depths<br />
between 142 and 1386 m but we would need <strong>to</strong> collect<br />
a specimen <strong>to</strong> confirm conspecificity.<br />
Pho<strong>to</strong>s. 1) Fragments <strong>of</strong> specimen collected at a depth<br />
<strong>of</strong> 175 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Note <strong>the</strong> hydroids<br />
growing on <strong>the</strong> fringe. Grid marks are 1 cm 2 . 2) Same<br />
specimen as in pho<strong>to</strong> 1 in situ. 3) Specimen collected<br />
at a depth <strong>of</strong> 486 m in Zhemchug Canyon, Bering Sea.<br />
Grid marks are 1 cm 2 .
31. Polymastia fluegeli Lehnert, S<strong>to</strong>ne and Heimler, 2005<br />
Description. This sponge is endopsammic and discshaped<br />
with numerous fistules on <strong>the</strong> upper surface.<br />
The surface is smooth with long protruding spicules<br />
scattered about. The cortical layer is cartilaginous and<br />
more resilient than <strong>the</strong> relatively s<strong>of</strong>t choanosome. The<br />
bot<strong>to</strong>m side <strong>of</strong> <strong>the</strong> sponge is very firm and slightly elastic<br />
and consists <strong>of</strong> a layer <strong>of</strong> spongin-cemented sediment.<br />
In situ <strong>the</strong> plate is buried in <strong>the</strong> sediment; only <strong>the</strong><br />
fistules protrude. The fistules are cone-shaped with no<br />
obvious opening. The disc is up <strong>to</strong> 52 cm in diameter<br />
and about 1 cm in thickness (3–4 mm <strong>of</strong> <strong>the</strong> thickness<br />
is due <strong>to</strong> <strong>the</strong> basal sediment layer). Color in life is light<br />
yellow with bright yellow fistules; disc is light brown<br />
after freezing.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Polyspicular tracts arise from <strong>the</strong> basal<br />
sediment layer <strong>the</strong>n widen, branch <strong>to</strong>wards <strong>the</strong> surface,<br />
and fan out in <strong>the</strong> cortical layer. Polyspicular tracts are<br />
350–500 µm in diameter. The cortical layer is 500–700<br />
µm in thickness and, except for <strong>the</strong> ends <strong>of</strong> <strong>the</strong> ascending<br />
polyspicular tracts, consists <strong>of</strong> a mass <strong>of</strong> spicules<br />
without orientation. The outermost layer is a palisade<br />
<strong>of</strong> small tylostyles, oriented outward creating <strong>the</strong> microscopically<br />
hispid surface. The papillae also show <strong>the</strong><br />
outermost palisade and <strong>the</strong> unorganized spicule mass<br />
below, but differ ins<strong>of</strong>ar as <strong>the</strong> polyspicular tracts run <strong>to</strong><br />
<strong>the</strong> tip <strong>of</strong> <strong>the</strong> papillum. A central canal is visible in <strong>the</strong><br />
center <strong>of</strong> <strong>the</strong> papillae but <strong>the</strong>re is no opening visible.<br />
There are large fusiform tylostyles, in a wide size-range,<br />
longest ones <strong>of</strong>ten with one or several subterminal<br />
rings (180–1750 × 8–22 µm), and a small category <strong>of</strong><br />
tylostyles (65–110 × 3–6 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Patchily distributed at depths between 81 and<br />
338 m in generally low-relief (i.e., flat-bot<strong>to</strong>med) habitats<br />
<strong>of</strong> small pebbles and coarse sand with moderate <strong>to</strong><br />
high current. Found at maximum densities up <strong>to</strong> eight<br />
individuals per m 2 in coarse sand habitats at 82 m depth.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 82 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 . 2)<br />
Same specimen as in pho<strong>to</strong> 1 in situ during collection.<br />
3) Specimen observed at a depth <strong>of</strong> 142 m in <strong>the</strong> central<br />
Aleutian Islands. A prowfish (Zapora silenus) lies in a den<br />
just below <strong>the</strong> specimen.<br />
47
48 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
32. Polymastia pacifica Lambe, 1893<br />
Description. This sponge is subglobular <strong>to</strong> cushionshaped.<br />
Surface is smooth <strong>to</strong> <strong>the</strong> unaided eye but<br />
microhispid with several papillae. Consistency is firm.<br />
Diameter is <strong>to</strong> at least 15 cm. Externally it is brownish orange<br />
in color; internally it is bright orange with lighter<br />
polyspicular tracts visible <strong>to</strong> <strong>the</strong> unaided eye.<br />
Skeletal structure. Ec<strong>to</strong>somal palisade <strong>of</strong> small tylostyles<br />
are supported by intermediate tylostyles. Large<br />
tylostyles radiate in polyspicular tracts <strong>to</strong> <strong>the</strong> surface.<br />
There are large tylostyles (up <strong>to</strong> 4400 × 20 µm), intermediate<br />
tylostyles (200–600 × 12–16 µm), and small<br />
tylostyles (120–160 × 5–7 µm).<br />
Zoogeographic distribution. Uncommon. North<br />
Pacific Ocean. In Alaska – central Aleutian Islands.<br />
Elsewhere – previously reported from Vancouver Island<br />
<strong>to</strong> California.<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles at depths<br />
between 150 and 160 m. Elsewhere – 73 m depth (Vancouver<br />
Island). Lamb and Hanby (2005) report that<br />
this species occurs from Alaska <strong>to</strong> California at depths<br />
between <strong>the</strong> intertidal zone and 183 m.<br />
Pho<strong>to</strong>. 1) Fragment <strong>of</strong> a specimen collected at a depth<br />
<strong>of</strong> 155 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 .
33. Stylocordyla borealis eous Koltun, 1966<br />
Description. Thin stalk terminates in a globular or<br />
oval body with a smooth surface. Stalk is <strong>to</strong> 9 cm; body<br />
<strong>to</strong> approximately 2 cm 3 . Color in life is golden brown.<br />
Skeletal structure. Skele<strong>to</strong>n <strong>of</strong> <strong>the</strong> stalk is a tight mass<br />
<strong>of</strong> parallel spicules running in<strong>to</strong> <strong>the</strong> body and radiating<br />
<strong>to</strong> <strong>the</strong> sides in tracts. Smaller oxeas form a palisade at<br />
<strong>the</strong> surface <strong>of</strong> <strong>the</strong> body. Microxeas form a tangential<br />
crust at <strong>the</strong> surface. Oxeas are exclusively centrotylote,<br />
sometimes with blunt ends. Oxeas occur in a wide<br />
size-range, possibly in three categories: 790–2460 ×<br />
10–37 µm, 400–700 × 10–12 µm, and microxeas 70–110 ×<br />
2–3 µm.<br />
Zoogeographic distribution. Locally abundant in<br />
<strong>the</strong> North Pacific Ocean. In Alaska – central Aleutian<br />
Islands and Bering Sea (Pribil<strong>of</strong> Canyon). Elsewhere –<br />
Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles and emergent<br />
epifauna at depths between 125 and 307 m and generally<br />
in low-relief (i.e., flat-bot<strong>to</strong>med) habitats (central<br />
Aleutian Islands). Sponges with very similar morphology<br />
have been observed in <strong>the</strong> Aleutian Islands at depths<br />
near 1100 m, but we would need <strong>to</strong> collect a specimen<br />
<strong>to</strong> confirm conspecificity. Elsewhere – reported at a<br />
depth <strong>of</strong> 200 m.<br />
Remarks. Stylocordyla borealis eous can be distinguished<br />
from <strong>the</strong> similar Rhizaxinella clavata by its thinner stalk<br />
and smooth oval body. Also, S. borealis eous has exclusively<br />
centrolyte oxeas while R. clavata has tylostyles and<br />
subtylostyles.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 127 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 (indicated by <strong>the</strong> white<br />
arrow) in situ. 3) Specimen collected at a depth <strong>of</strong> 208<br />
m in Pribil<strong>of</strong> Canyon, Bering Sea. Specimen is attached<br />
<strong>to</strong> an unknown hexactinellid sponge.<br />
49
50 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
34. Aap<strong>to</strong>s kanuux Lehnert, Hocevar, and S<strong>to</strong>ne 2008<br />
Description. Species is irregularly globular. Surface<br />
is almost smooth, microscopically slightly uneven. No<br />
oscula are visible. Consistency is firm, only slightly<br />
elastic. Diameter ranges from 9 <strong>to</strong> 19 mm. Color in life<br />
is mustard yellow.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Spicules are strictly radially arranged<br />
in <strong>the</strong> choanosome; spicule density is very high. Special<br />
dermal oxeas are densely arranged perpendicular <strong>to</strong><br />
<strong>the</strong> surface and form a palisade. There are fusiform<br />
strongyloxeas (1795–2132 × 15–22 µm), subtylostyles<br />
(500–770 × 8–10 µm), and ec<strong>to</strong>somal tylostyles (104–215<br />
× 4–8 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – known only from <strong>the</strong> type locality in Pribil<strong>of</strong><br />
Canyon, Bering Sea. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> pebbles in low-relief (i.e., flatbot<strong>to</strong>med)<br />
silt and sand habitat at depths between 203<br />
and 240 m. Often found in association with several<br />
unknown species <strong>of</strong> hydroids, zoanthids, and <strong>the</strong> demosponge<br />
Stylocordyla borealis eous.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 219 m in<br />
Pribil<strong>of</strong> Canyon, Bering Sea. 2) Same specimen as in<br />
pho<strong>to</strong> 1 showing close-up detail <strong>of</strong> <strong>the</strong> radial arrangement<br />
<strong>of</strong> <strong>the</strong> choanosome. 3) Same specimen as in<br />
pho<strong>to</strong>s 1 and 2 in situ. The separation between <strong>the</strong> red<br />
laser marks is 10 cm.
35. Rhizaxinella clavata (Thiele, 1898)<br />
Description. Thin stalk terminates in a globular or<br />
oval body. Species is similar <strong>to</strong> Stylocordyla borealis eous.<br />
This species has a slightly thicker stalk which widens <strong>to</strong>wards<br />
<strong>the</strong> body and a strongly hispid surface <strong>of</strong> <strong>the</strong> body<br />
compared <strong>to</strong> <strong>the</strong> smooth surface <strong>of</strong> S. borealis eous. Spicules<br />
are arranged radially in <strong>the</strong> interior <strong>of</strong> <strong>the</strong> sponge<br />
with dense spicule brushes at <strong>the</strong> surface. Length is up<br />
<strong>to</strong> 5.5 cm. Color in life is golden brown.<br />
Skeletal structure. There are tylostyles <strong>to</strong> subtylostyles<br />
(1050–1780 × 15–32 µm) and smaller tylostyles<br />
(180–370 × 6–10 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– Bering Sea (Zhemchug Canyon). Elsewhere – <strong>the</strong><br />
holotype was recorded from <strong>the</strong> Sea <strong>of</strong> Japan.<br />
Habitat. In Alaska – attached <strong>to</strong> hexactinellid sponge<br />
skele<strong>to</strong>ns at a depth <strong>of</strong> 915 m. Elsewhere – reported on<br />
mud bot<strong>to</strong>ms at a depth <strong>of</strong> 183 m.<br />
Remarks. Rhizaxinella clavata can be distinguished<br />
from <strong>the</strong> similar Stylocordyla borealis eous by its thicker<br />
51<br />
stalk and hispid oval body. Also, R. clavata has tylostyles<br />
and subtylostyles while S. borealis eous has exclusively<br />
centrolyte oxeas.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 915 m in<br />
Zhemchug Canyon, Bering Sea. Specimen is attached <strong>to</strong><br />
an unidentified hexactinellid sponge skele<strong>to</strong>n.
52 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
36. Suberites excellens (Thiele, 1898)<br />
Description. This massively lobate sponge has irregularly<br />
cylindrical lobes. Surface is smooth <strong>to</strong> <strong>the</strong> unaided<br />
eye but microscopically hispid. Circular oscula are near<br />
tips <strong>of</strong> lobes and slightly elevated above <strong>the</strong> surface.<br />
Consistency is firm but elastic. Height is <strong>to</strong> 14 cm. Color<br />
in life is yellowish brown.<br />
Skeletal structure. Skeletal architecture is more or<br />
less confused in <strong>the</strong> interior but becomes radial <strong>to</strong>wards<br />
<strong>the</strong> surface. The surface consists <strong>of</strong> a palisade <strong>of</strong> smaller<br />
tylostyles. There are thick tylostrongyles with round<br />
tyles at one end and only slightly tapering at <strong>the</strong> o<strong>the</strong>r<br />
(640–1890 × 17–46 µm), somewhat thinner and flexuous<br />
tylostrongyles (1140–2100 × 17–26 µm), and small<br />
tylostyles forming <strong>the</strong> palisade at <strong>the</strong> surface (175–520<br />
× 8–12 µm).<br />
Zoogeographic distribution. Rare. In Alaska – Bering<br />
Sea (Zhemchug Canyon) and eastern Gulf <strong>of</strong> Alaska.<br />
Elsewhere – Holotype described from Sagami Bay, Japan;<br />
also recorded along Korean Coast (Sim and Kim,<br />
1988).<br />
Habitat. In Alaska – attached <strong>to</strong> boulders and cobbles<br />
at depths between 390 and 601 m. Elsewhere – no information<br />
available.<br />
Remarks. This was previously known as Rhizaxinella<br />
excellens Thiele, 1898. It typically harbors an abundant<br />
and diverse complement <strong>of</strong> ophiuroid associates.<br />
Pho<strong>to</strong>s. 1) Fragments <strong>of</strong> specimen with ophiuroid<br />
associates collected at a depth <strong>of</strong> 390 m in Zhemchug<br />
Canyon, Bering Sea. 2) Close-up view <strong>of</strong> oscula a<strong>to</strong>p<br />
<strong>the</strong> lobes.
37. Suberites simplex Lambe, 1893<br />
Description. This small sponge has a subglobular <strong>to</strong><br />
globular growth form. Surface is smooth <strong>to</strong> <strong>the</strong> unaided<br />
eye but microscopically hispid. Consistency is firm but<br />
elastic. Diameter is at least 5 cm. Color in life is golden<br />
brown <strong>to</strong> brown.<br />
Skeletal structure. Ec<strong>to</strong>somal skele<strong>to</strong>n consists <strong>of</strong> bouquets<br />
<strong>of</strong> small tylostyles at <strong>the</strong> surface. Densely packed<br />
larger tylostyles in <strong>the</strong> interior are arranged without<br />
orientation. Tylostyles are large (620–1170 × 10–20 µm)<br />
and small (120–230 × 3–5 µm).<br />
Zoogeographic distribution. Uncommon in <strong>the</strong><br />
North Pacific Ocean. In Alaska – central Aleutian Islands.<br />
Elsewhere – Vancouver Island.<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles at depths between<br />
150 and 160 m. Elsewhere – reported at a depth<br />
<strong>of</strong> 73 m.<br />
53<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
54 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
38. Suberites sp.<br />
Description. This “mobile” sponge lives as an irregularly<br />
globular encrustation on empty gastropod shells.<br />
Frequently lives in symbiosis with hermit crabs including<br />
Pagurus dalli. Surface is smooth <strong>to</strong> <strong>the</strong> unaided eye<br />
but microscopically hispid due <strong>to</strong> dense spicule brushes<br />
that leave open numerous small pores (50–100 µm diameter).<br />
Irregular short tracts in <strong>the</strong> choanosome form<br />
a vague reticulation. Diameter is at least 10 cm. Color in<br />
life is red, orange, or tan; pale yellow in ethanol.<br />
Skeletal structure. Viewed under high magnification,<br />
<strong>the</strong> spicules are relatively dense without particular orientation.<br />
There are tylostyles <strong>to</strong> tylostrongyles (123–350<br />
× 7–12 µm).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – Bering Sea <strong>to</strong> Sou<strong>the</strong>ast Alaska.<br />
Habitat. In Alaska – typically encrusts gastropod shells<br />
at depths between 20 and 165 m.<br />
Remarks. This species is indistinguishable from Suberites<br />
domuncula that is common <strong>to</strong> <strong>the</strong> Mediterranean Sea<br />
and Atlantic Ocean along <strong>the</strong> west coast <strong>of</strong> North Africa.<br />
However, we doubt conspecificity based on <strong>the</strong> disjunct<br />
zoogeography and leave <strong>the</strong> species assignment for this<br />
specimen as undecided. The species complex is in desperate<br />
need <strong>of</strong> taxonomic revision. There are probably<br />
several sibling species that live in symbiosis with hermit<br />
crabs (more than a dozen species reported worldwide).<br />
S. domuncula contains suberitine, a neuro<strong>to</strong>xin that can<br />
cause fatal hemolytic hemorrhaging in some animals<br />
including some reef fish. However, <strong>the</strong> hawksbill turtle<br />
(Eretmochelys imbricata) does prey upon it in tropical <strong>water</strong>s<br />
(Meylan, 1988). No studies have been conducted<br />
on Alaskan specimens.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 84 m in<br />
<strong>the</strong> central Aleutian Islands. This is <strong>the</strong> ventral side <strong>of</strong><br />
<strong>the</strong> sponge showing <strong>the</strong> cavity in which a hermit crab<br />
Pagurus dalli (completely retracted) is living. Grid marks<br />
are 1 cm 2 .
39. Hemigellius porosus (Fristedt, 1887)<br />
Description. This sponge is massively encrusting and<br />
very fragile. Surface is smooth <strong>to</strong> <strong>the</strong> unaided eye but<br />
microscopically hispid. No oscula are visible. Fristedt<br />
(1887) described this species as forming irregular<br />
knolls, 9 cm in longest dimension. Color in life is whitish<br />
yellow <strong>to</strong> creamy white.<br />
Skeletal structure. It has an irregular unispicular or<br />
paucispicular mesh <strong>of</strong> oxeas (250–340 × 9–11 µm) and<br />
stigmata (40–60 µm).<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – Bering Sea (Zhemchug Canyon). Elsewhere<br />
– North Pacific Ocean (Sea <strong>of</strong> Okhotsk and Sea<br />
<strong>of</strong> Japan), Arctic Ocean (Barents Sea, Kara Sea, Laptev<br />
Sea), North Atlantic Ocean (West <strong>of</strong> Spitzbergen, East<br />
<strong>of</strong> Greenland, Denmark Strait, Davis Strait, Gulf <strong>of</strong> St.<br />
Lawrence, between Iceland and <strong>the</strong> Faroe Islands).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles at a depth <strong>of</strong><br />
909 m. Elsewhere – reported at depths between 68 and<br />
256 m.<br />
55<br />
Pho<strong>to</strong>. 1) Fragments <strong>of</strong> specimen collected at a depth<br />
<strong>of</strong> 909 m in Zhemchug Canyon, Bering Sea.
56 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
40. Cornulum clathriata (Koltun, 1955)<br />
Description. This stalked fan-shaped sponge consists<br />
<strong>of</strong> a meshwork or net; not flat but slightly concave.<br />
Consistency is wiry due <strong>to</strong> <strong>the</strong> mesh <strong>of</strong> spicule tracts. It<br />
is relatively small; height <strong>to</strong> 11 cm. Color in life is light<br />
yellow.<br />
Skeletal structure. Structure includes ec<strong>to</strong>somal<br />
tylotes with acanthaceous ends (280–420 × 6–8 µm);<br />
choanosomal styles with slightly acanthaceous blunt<br />
ends in a wide size range (215–2000 × 18–33 µm); and<br />
spherical isochelae (17–25 µm) and <strong>to</strong>xa in two size<br />
categories – large (55–220 µm) and small (10–18 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
central Aleutian Islands. Elsewhere – previously known<br />
from only two records near <strong>the</strong> Commander Islands<br />
(Russia) in <strong>the</strong> western Bering Sea.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock or muds<strong>to</strong>ne<br />
at depths between 843 and 1720 m. Elsewhere – reported<br />
at a depth <strong>of</strong> 2440 m.<br />
Remarks. Previously known as Melonchela clathriata<br />
Koltun, 1955.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 1720 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ.
41. Iophon piceum (Vosmaer, 1882)<br />
Description. This sponge is massively encrusting.<br />
According <strong>to</strong> Koltun (1959), growth form may also<br />
be massive-lobate, tabular, dactylate, ramified, or irregularly<br />
lobate. Oscula are on short elevations or flush<br />
with <strong>the</strong> surface. Dermal membrane is thin and pellicular.<br />
Consistency is s<strong>of</strong>t and easily <strong>to</strong>rn. It may reach a<br />
height <strong>of</strong> 15 cm. Color in life is golden brown <strong>to</strong> brown.<br />
Skeletal structure. Ec<strong>to</strong>somal <strong>to</strong>rnotes (150–270 ×<br />
6–10 µm) with micro-spined heads, are arranged in bundles<br />
tangential <strong>to</strong> <strong>the</strong> surface. Choanosomal skele<strong>to</strong>n is<br />
reticulate with smooth styles or acanthostyles (195–435 ×<br />
9–18 µm), anisochelae (18–37 µm), and bipocillae<br />
(8–14 µm).<br />
Zoogeographic distribution. Widespread and locally<br />
common. In Alaska – central Aleutian Islands. Elsewhere<br />
– North Pacific Ocean (Sea <strong>of</strong> Okhotsk and Sea<br />
<strong>of</strong> Japan), Arctic Ocean (Barents Sea including White<br />
Sea, Kara Sea – Vilkitsky Strait, and Greenland Sea), and<br />
North Atlantic Ocean (Norwegian Sea, Davis Strait, and<br />
Denmark Strait).<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles and small<br />
cobbles at depths between 94 and 155 m. Elsewhere –<br />
eurybathic; reported at depths between 9 and 1785 m.<br />
Remarks. This species <strong>of</strong>ten grows in association with,<br />
and encrusts, hydroids and bryozoans.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 94 m in<br />
<strong>the</strong> central Aleutian Islands. Specimen is encrusting <strong>the</strong><br />
hydroid Abietinaria sp. (lower right). Grid marks are 1<br />
cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 (indicated by <strong>the</strong><br />
white arrows) in situ with <strong>the</strong> plumose hydroids (Abietinaria<br />
sp.). 3) Specimen collected at a depth <strong>of</strong> 155 m<br />
in <strong>the</strong> central Aleutian Islands. Specimen is encrusting<br />
<strong>the</strong> bryozoan (Microporina cf. articulata). Grid marks<br />
are 1 cm 2 .<br />
57
58 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
42. Iophon piceum abipocillus Koltun, 1959<br />
Description. This sponge is massively encrusting. It<br />
is very similar <strong>to</strong> Iophon piceum except <strong>the</strong> ec<strong>to</strong>some<br />
is white ra<strong>the</strong>r than golden-brown and <strong>the</strong> oscula are<br />
distinctly conical. Diameter is <strong>to</strong> 25 cm. Color in life is<br />
white; turns completely brown with darker brown oscula<br />
in ethanol.<br />
Skeletal structure. It is very similar <strong>to</strong> Iophon piceum<br />
except that bipocillae are absent.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
central Aleutian Islands. Elsewhere – Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – attached <strong>to</strong> boulders, cobbles, and<br />
pebbles at depths between 82 and 192 m. Elsewhere –<br />
reported at a depth <strong>of</strong> 1240 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 100 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ.
43. Megaciella anisochela Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge has clusters <strong>of</strong> 4–8 tubes with<br />
a common stalk. The surface is finely hispid with no recognizable<br />
oscula. The consistency is s<strong>of</strong>t and elastic. The<br />
dimensions <strong>of</strong> <strong>the</strong> short stalk are 2.0 × 0.5 cm and <strong>the</strong><br />
tubes are approximately 11 × 5 cm. Color in life is light<br />
yellow; golden brown after freezing; beige in ethanol.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some is a thin translucent<br />
membrane with tangentially arranged tylotes, singly or<br />
in small bundles, facing in all directions within <strong>the</strong> tangential<br />
plane. The ec<strong>to</strong>some contains many isochelae<br />
and is supported by underlying styles. The choanosome<br />
is a combination <strong>of</strong> a unispicular reticulation <strong>of</strong> single<br />
spicules and ascending paucispicular tracts connected<br />
by single spicules. Ascending tracts <strong>of</strong> styles penetrate<br />
<strong>the</strong> ec<strong>to</strong>somal membrane and cause <strong>the</strong> hispidation.<br />
Ec<strong>to</strong>somal tylotes (245–380 × 4–9 µm) have acanthose<br />
heads. Smooth choanosomal styles measure 490–615 ×<br />
18–22 µm. Microscleres are palmate isochelae (13–17<br />
µm) with narrow extensions, a small category <strong>of</strong> palmate<br />
isochelae (6–8 µm), and a small category <strong>of</strong> dis<strong>to</strong>rted<br />
anisochelae (4–6 µm).<br />
Zoogeographic distribution. In Alaska – locally abundant<br />
in central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Attached <strong>to</strong> bedrock, boulders, and cobbles<br />
at depths between 702 and 750 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 702 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ.<br />
59
60 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
44. Megaciella spirinae (Koltun, 1958)<br />
Description. This irregularly massive-lobate sponge<br />
has an uneven, corrugated surface. Oscula are present<br />
on <strong>to</strong>p <strong>of</strong> cylindrical elevations. Consistency is elastic,<br />
s<strong>of</strong>t, and easily <strong>to</strong>rn. Diameter is <strong>to</strong> 13 cm. Color in life<br />
is golden brown.<br />
Skeletal structure. Ec<strong>to</strong>somal <strong>to</strong>rnotes (160–210 ×<br />
4-6 µm) have acanthose ends. There are choanosomal<br />
acanthostyles (175–212 × 12–16 µm), arcuate isochelae<br />
(28–32 µm), and <strong>to</strong>xa (90–150 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – eastern Tartar<br />
Strait, near <strong>the</strong> Pacific Coast <strong>of</strong> <strong>the</strong> sou<strong>the</strong>rn Kuril Islands.<br />
Habitat. In Alaska – attached <strong>to</strong> small cobbles and pebbles<br />
in generally low-relief (i.e., flat-bot<strong>to</strong>med) habitat<br />
at depths between 150 and 160 m. Elsewhere – attached<br />
<strong>to</strong> pebbles in sandy habitat at depths between 71 and<br />
414 m and at temperatures between 2.3 and 6.6°C.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
45. Clathria (Clathria) barleei (Bowerbank, 1866)<br />
Description. This sponge is elongated and massive. It<br />
is irregularly ramified with a corrugated surface; oscula<br />
are flush with <strong>the</strong> surface. Diameter is <strong>to</strong> 15 cm. Color<br />
in life is orange-brown.<br />
Skeletal structure. Dermal styles (270–360 × 3–6 µm)<br />
have acanthose heads. There are choanosomal smooth<br />
styles (420–540 × 18–35 µm), isochelae (16–20 µm), and<br />
<strong>to</strong>xa (130–720 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – Arctic Ocean<br />
(Barents Sea near Murmansk, Russia) and Nor<strong>the</strong>ast<br />
Atlantic Ocean (from Ireland <strong>to</strong> France and Norway).<br />
Habitat. In Alaska – attached <strong>to</strong> small cobbles and pebbles<br />
in generally low-relief (i.e., flat-bot<strong>to</strong>med) habitat<br />
at depths between 150 and 160 m. Elsewhere – reported<br />
at depths between 72 and 440 m and temperatures between<br />
1 and 4.2° C.<br />
Remarks. Specimens from <strong>the</strong> Nor<strong>the</strong>ast Atlantic<br />
are described as flabellate with a characteristic honeycombed<br />
surface <strong>of</strong> a different color (Van Soest and<br />
S<strong>to</strong>ne, 1986). The spicule complement (types and size<br />
61<br />
ranges) <strong>of</strong> this specimen conforms perfectly <strong>to</strong> Clathria<br />
barleei so we assume it <strong>to</strong> be ano<strong>the</strong>r growth form <strong>of</strong> <strong>the</strong><br />
same species in <strong>the</strong> Pacific Ocean. Van Soest and S<strong>to</strong>ne<br />
(1986) report this species from <strong>the</strong> Arctic <strong>to</strong> <strong>the</strong> west<br />
coasts <strong>of</strong> Ireland and France so a circumpolar distribution<br />
seems possible.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
62 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
46. Clathria (Clathria) laevigata Lambe, 1893<br />
Description. This sponge is massive, laterally compressed,<br />
and irregularly lobate. Consistency is s<strong>of</strong>t and<br />
fragile with round, slightly elevated oscula. Diameter is<br />
<strong>to</strong> 30 cm. Color in life is golden brown. The ec<strong>to</strong>somal<br />
membrane may have a greyish tinge.<br />
Skeletal structure. Structure includes ec<strong>to</strong>somal<br />
smooth small styles (230–290 × 6–10 µm); choanosomal<br />
smooth styles (440–560 × 20–30 µm) occasionally with<br />
acanthose heads; and small acanthostyles (90–240 ×<br />
8–12 µm). Microscleres are palmate isochelae (20–25<br />
µm) and <strong>to</strong>xas (200–530 µm).<br />
Zoogeographic distribution. Locally common in<br />
North Pacific Ocean. In Alaska – central Aleutian Islands.<br />
Elsewhere – known from Kuril Strait (Russia) and<br />
near Vancouver Island (Canada).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles and pebbles<br />
at a depth <strong>of</strong> 167 m. Elsewhere – reported at depths<br />
between 72 and 138 m.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> a specimen collected at a<br />
depth <strong>of</strong> 167 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 (indicated<br />
by <strong>the</strong> white arrow) in situ.
47. Clathria (Axosuberites) lambei (Koltun, 1955)<br />
Description. This sponge grows from an encrusting<br />
base <strong>to</strong> a mass <strong>of</strong> branches or lobes with an uneven<br />
surface. Consistency is s<strong>of</strong>t and elastic. Diameter is <strong>to</strong><br />
30 cm. Color in life is golden brown.<br />
Skeletal structure. Ec<strong>to</strong>some consists <strong>of</strong> brushes <strong>of</strong><br />
smaller subtylostyles and tylostyles with large protruding<br />
choanosomal styles. Choanosomal smooth styles, <strong>of</strong>ten<br />
with acanthaceous heads, are arranged in plumose<br />
tracts and echinated by small acanthostyles. There are<br />
eec<strong>to</strong>somal tylostyles and subtylostyles (520–1370 × 8–18<br />
µm); choanosomal large styles (540–1960 × 28–40 µm)<br />
with acanthose heads, acanthostyles (120–430 × 10–20<br />
µm), and palmate isochelae (18–25 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – sou<strong>the</strong>rn Sea<br />
<strong>of</strong> Okhotsk and Sea <strong>of</strong> Japan (sou<strong>the</strong>rn Kuril Islands).<br />
Habitat. In Alaska – attached <strong>to</strong> small cobbles and<br />
pebbles in generally low-relief (i.e., flat-bot<strong>to</strong>med)<br />
habitat at depths between 150 and 160 m. Elsewhere –<br />
reported at depths between 91 and 550 m on oozy sand<br />
63<br />
and pebbles and at temperatures between 1 and 1.3°C<br />
and a salinity <strong>of</strong> 34.18 psu.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
64 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
48. Echinoclathria vasa Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge has a stalked vase with a<br />
flared lip and a markedly concave surface; walls are<br />
5–10 mm in thickness. Consistency, except for <strong>the</strong> stalk,<br />
is very s<strong>of</strong>t and elastic. In areas <strong>of</strong> <strong>water</strong> current this species<br />
typically bends over near <strong>the</strong> base and sometimes<br />
lies in contact with <strong>the</strong> seafloor. Height is <strong>to</strong> 13 cm and<br />
width <strong>to</strong> 3 cm. Color in life is light yellow <strong>to</strong> creamy<br />
white; golden brown after freezing.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some is a thin membrane<br />
packed with anchorate isochelas (21–27 µm). The choanosome<br />
consists <strong>of</strong> paucispicular tracts <strong>of</strong> thick styles<br />
(760–920 × 16–21 µm) and brushes <strong>of</strong> thin styles that fan<br />
out <strong>to</strong>wards <strong>the</strong> surface with many isochelas in between.<br />
Thin styles (740–1230 × 4–6 µm) are slightly curved <strong>to</strong><br />
sinuous and have finely acanthose heads.<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Occurring individually on exposed bedrock,<br />
cobbles, and pebbles at depths between 622 and 876 m<br />
and generally in low-relief (i.e., flat-bot<strong>to</strong>med) habitats.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at<br />
a depth <strong>of</strong> 744 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ<br />
during collection. 3) Same specimen as in pho<strong>to</strong>s 1 and<br />
2 in situ showing detail <strong>of</strong> <strong>the</strong> flared lip.
49. Artemisina amlia Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge is stalked with a subhemispherical<br />
or conical body. The stalk widens gradually<br />
from 4 <strong>to</strong> 25 mm over a distance <strong>of</strong> approximately 9 cm<br />
and is not sharply separated from <strong>the</strong> body. The consistency<br />
is s<strong>of</strong>t, elastic, and easily <strong>to</strong>rn. There are wart-like,<br />
slightly elevated oscula on <strong>the</strong> dorsal surface <strong>of</strong> <strong>the</strong><br />
sponge only that are circular and 2 mm in diameter. Diameter<br />
<strong>of</strong> conical body is <strong>to</strong> 20 cm; <strong>to</strong>tal height <strong>to</strong> 15 cm<br />
or more. Color in life is light orange <strong>to</strong> golden brown. In<br />
ethanol <strong>the</strong> body has a whitish, translucent ec<strong>to</strong>some on<br />
<strong>to</strong>p with a yellowish, fibrous choanosome underneath.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some <strong>of</strong> <strong>the</strong> conical body is<br />
composed <strong>of</strong> a mesh-work <strong>of</strong> polyspicular tracts (55–175<br />
µm in diameter) <strong>of</strong> small styles with a mesh-size <strong>of</strong> 350–<br />
750 µm. This large mesh is subdivided by a finer net <strong>of</strong><br />
strands <strong>of</strong> spongin-embedded isochelae. This finer net<br />
has a mesh-size <strong>of</strong> 45–90 µm; single, translucent strands<br />
are 15–25 µm in diameter. The ec<strong>to</strong>some <strong>of</strong> <strong>the</strong> stalk<br />
is thinner and consists <strong>of</strong> a dense, unispicular layer <strong>of</strong><br />
tangentially arranged, parallel-oriented thick styles. In<br />
<strong>the</strong> choanosome <strong>of</strong> <strong>the</strong> stalk, ascending polyspicular<br />
tracts <strong>of</strong> thick styles are connected by paucispicular<br />
tracts and single spicules, comparable <strong>to</strong> <strong>the</strong> choanosome<br />
<strong>of</strong> <strong>the</strong> conical body. There are large, smooth styles<br />
(400–520 × 20–25 µm), small styles (330–550 × 10 µm)<br />
with acanthose heads and <strong>of</strong>ten with one prominent<br />
dent, isochelae (10–13 µm), and <strong>to</strong>xa (110–170 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Attached <strong>to</strong> bedrock, boulders, cobbles, and<br />
pebbles at depths between 97 and 253 m. Associated<br />
with <strong>the</strong> demosponge Mycale carlilei.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 119 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ showing dorsal<br />
surface with elevated oscula.<br />
65
66 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
50. Artemisina arcigera (Schmidt, 1870)<br />
Description. This sponge is stalked and tube-shaped<br />
in <strong>the</strong> Aleutian Islands, but Koltun (1959) described<br />
this species as cushion-shaped, spherical, or somewhat<br />
elongated, and up <strong>to</strong> 5.5 cm in height. The surface <strong>of</strong><br />
<strong>the</strong> sponge is smooth at <strong>the</strong> stalk and gradually changes<br />
from hispid <strong>to</strong> rugose in <strong>the</strong> lower half and is smooth<br />
again in <strong>the</strong> apical region. No oscula are visible. The<br />
consistency <strong>of</strong> <strong>the</strong> stalk is wiry and gets s<strong>of</strong>ter along its<br />
length. In areas <strong>of</strong> <strong>water</strong> current this species typically<br />
bends over near <strong>the</strong> base and sometimes lies in contact<br />
with <strong>the</strong> seafloor. Stalked forms are <strong>to</strong> 31 cm in height<br />
and 7 cm in width. Color in life is brown <strong>to</strong> golden<br />
brown.<br />
Skeletal structure. There are ec<strong>to</strong>somal brushes <strong>of</strong><br />
smaller subtylostyles (280–428 × 9–18 µm); choanosomal<br />
large subtylostyles (430–676 × 6–9 µm), occasionally<br />
with acanthaceous heads; palmate isochelae (6–15 µm);<br />
and <strong>to</strong>xas with acanthaceous ends (60–360 µm).<br />
Zoogeographic distribution. Widespread and locally<br />
common. In Alaska – central Aleutian Islands. Elsewhere<br />
– widely distributed in <strong>the</strong> Arctic Ocean (Barents<br />
Sea – White Sea, Kara Sea – Vilkitsky Strait, Greenland<br />
Sea – Island <strong>of</strong> Spitsbergen) and Nor<strong>the</strong>ast Atlantic<br />
Ocean (Norwegian Sea, Denmark Strait, Davis Strait).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles and pebbles<br />
at depths between 170 and 436 m. Elsewhere – reported<br />
at depths between 14 and 1000 m and temperatures<br />
between 1.4 and 5.0°C.<br />
Remarks. Specimens from <strong>the</strong> Aleutian Islands differ<br />
in form from previous records but <strong>the</strong> spicule<br />
complement (types and size ranges) <strong>of</strong> <strong>the</strong> specimens<br />
conform perfectly <strong>to</strong> A. arcigera so we assume <strong>the</strong>y are<br />
a previously unreported growth form <strong>of</strong> <strong>the</strong> same species.<br />
The shrimp Eualus barbatus is <strong>of</strong>ten associated with<br />
this sponge.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 192 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ (center).<br />
3) Same specimen as in pho<strong>to</strong>s 1 and 2 showing details<br />
<strong>of</strong> <strong>the</strong> rugose surface and apical region.
51. Artemisina stipitata Koltun, 1958<br />
Description. This sponge is flabellate in <strong>the</strong> Aleutian<br />
Islands but spherical and lobate forms have been<br />
reported elsewhere. It is basally stalked and gradually<br />
tapers <strong>to</strong> a holdfast. Conspicuous oscula are present on<br />
dorsal surfaces. The stalk is cylindrical and rigid. The<br />
body is s<strong>of</strong>t and elastic. Height and width are at least 50<br />
cm. The ec<strong>to</strong>some is yellow or light brown. The choanosome<br />
is orange-brown.<br />
Skeletal structure. Ec<strong>to</strong>somal skele<strong>to</strong>n has tangentially<br />
arranged small styles with acanthose heads (220–340<br />
× 6–10 µm). The reticulate choanosomal skele<strong>to</strong>n has<br />
fusiform styles with acanthose heads (460–510 × 18–25<br />
µm). Microscleres are palmate isochelae (10–19 µm).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – central Aleutian Islands. Elsewhere – previously<br />
known from only two specimens collected in sou<strong>the</strong>rn<br />
Kuril Strait until discovered in <strong>the</strong> central Aleutian<br />
Islands in 2004.<br />
Habitat. In <strong>the</strong> Aleutian Islands – attached <strong>to</strong> bedrock<br />
outcrops, boulders, and cobbles at depths between 80<br />
and 239 m on shelf and upper slope habitats. Elsewhere<br />
– no information available.<br />
Remarks. It is similar in appearance <strong>to</strong> Tedania kagalaskai,<br />
with which it co-occurs. The dorsal surface <strong>of</strong> A.<br />
stipitata is typically more rounded than T. kagalaskai and<br />
<strong>the</strong> body form <strong>of</strong> <strong>the</strong> latter species is much smoo<strong>the</strong>r<br />
and more distinctly triangular. A. stipitata is <strong>of</strong>ten associated<br />
with <strong>the</strong> gorgonian Fanellia compressa.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 150 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ showing <strong>the</strong><br />
67<br />
detail <strong>of</strong> <strong>the</strong> oscula on <strong>the</strong> dorsal surface. 3) Specimen<br />
observed in situ at a depth <strong>of</strong> 87 m in <strong>the</strong> central Aleutian<br />
Islands. 4) Specimen observed in situ at a depth <strong>of</strong><br />
105 m in <strong>the</strong> central Aleutian Islands. A juvenile king<br />
crab uses <strong>the</strong> sponge as a perch.
68 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
52. Artemisina sp.<br />
Description. Aleutian Island specimens consist <strong>of</strong><br />
many large rounded lobes growing from a narrow base.<br />
Large lobes <strong>of</strong>ten develop side-branches. Koltun (1959)<br />
described this same species as funnel-shaped, thickly<br />
tabular or massive. The surface is strongly rugose with<br />
a thin ec<strong>to</strong>somal membrane. Circular oscula are flush<br />
with <strong>the</strong> surface. It is s<strong>of</strong>t and elastic but fragile. Size is<br />
large with height <strong>to</strong> 40 cm and width <strong>to</strong> 60 cm. Color<br />
in life varies from yellowish grey, light brown <strong>to</strong> orange;<br />
typically golden yellow color in situ.<br />
Skeletal structure. Ec<strong>to</strong>somal thin styles form a unispicular<br />
reticulation with microscleres scattered in<br />
between. Choanosomal multispicular tracts have large<br />
styles and single spicules. Styles are <strong>of</strong>ten acanthaceous<br />
at <strong>the</strong> base. There are large styles (490–840 × 15–30 µm),<br />
small styles (210–480 × 7–10 µm), palmate isochelae<br />
(12–22 µm), small <strong>to</strong>xa (30–120 µm), and large <strong>to</strong>xa<br />
(280–340 µm).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – central Aleutian Islands. Elsewhere – Arctic<br />
Ocean (Barents, Greenland, Kara, and Laptev seas).<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, boulders,<br />
and cobbles at depths between 80 and 195 m. Elsewhere<br />
– reported at depths between 18 and 380 m.<br />
Remarks. This is Artemisina apollinis sensu Koltun<br />
(1959). However, A. apollinis was originally described<br />
from near <strong>the</strong> Kerguelen Islands in <strong>the</strong> sou<strong>the</strong>rn Indian<br />
Ocean so conspecificity is unlikely. It is similar in<br />
appearance <strong>to</strong> Mycale loveni with which it co-occurs, but<br />
this species is much more heavily lobed than M. loveni<br />
and narrows <strong>to</strong> a s<strong>to</strong>ut and narrow base. M. loveni is more<br />
massive with a less defineable base. It may be preyed<br />
upon by <strong>the</strong> sea star Henricia sp.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 195 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 . 2)<br />
Specimen collected at a depth <strong>of</strong> 155 m in <strong>the</strong> central<br />
Aleutian Islands. Grid marks are 1 cm 2 . 3) Specimen<br />
observed at a depth <strong>of</strong> 122 m in <strong>the</strong> central Aleutian<br />
Islands.
53. Coelosphaera oglalai Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge is subglobular or massively<br />
encrusting. The smooth surface is covered irregularly<br />
with conical papillae <strong>of</strong> very different sizes, ranging<br />
from 2–20 mm in height and 1–18 mm in diameter.<br />
Individuals are <strong>to</strong> about 8 cm in diameter. Color in life<br />
is light orange; beige in ethanol.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some is a tangential arrangement<br />
<strong>of</strong> tylote bundles in some areas and a tangential<br />
arrangement <strong>of</strong> all spicule types in o<strong>the</strong>r areas. In <strong>the</strong><br />
choanosome, strongyles are arranged halichondroid,<br />
relatively dense; polyspicular tracts are vaguely recognizable<br />
in some areas. The choanosomal megascleres vary<br />
from blunt ended (anis-) oxeas <strong>to</strong> strongyles (570–634<br />
× 27–32 µm), always with finely acanthose ends. Ec<strong>to</strong>somal<br />
tylotes (325–364 × 7–10 µm) have slightly acanthose<br />
ends. Microscleres are arcuate isochelae (47–52 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> cobbles and pebbles at depths<br />
between 100 and 155 m.<br />
69<br />
Remarks. They may be overgrown with bryozoans and<br />
found in association with <strong>the</strong> demosponge Iophon piceum<br />
abipocillus.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
70 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
54. Inflatella globosa Koltun, 1955<br />
Description. This sponge is subglobular with a conulose<br />
surface. Tiny conules are distributed uniformly<br />
over <strong>the</strong> surface <strong>of</strong> <strong>the</strong> sponge, but oscula are a<strong>to</strong>p small<br />
papillae that are restricted <strong>to</strong> <strong>the</strong> dorsal surface. The<br />
oscula contract upon fixation. Consistency is ra<strong>the</strong>r firm<br />
but elastic. Diameter is <strong>to</strong> approximately 50 cm. Color<br />
in life is orange, yellow, <strong>to</strong> creamy white; pale yellow in<br />
ethanol.<br />
Skeletal structure. There is a thick dermal membrane,<br />
densely packed with tylotes (300–400 µm in thickness)<br />
without apparent organization. Tylotes (about 240–350<br />
× 5–10 µm) have distinctive swollen heads that form<br />
polyspicular tracts arranged in an irregular meshwork.<br />
There are no microscleres.<br />
Zoogeographic distribution. Locally common in<br />
<strong>the</strong> North Pacific Ocean. In Alaska – central Aleutian<br />
Islands. Elsewhere – Bering Sea near Mednyi Island<br />
(Commander Islands), western Sea <strong>of</strong> Okhotsk, and<br />
Sea <strong>of</strong> Japan.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock and cobbles<br />
at depths between 56 and 138 m. Elsewhere – on rock,<br />
gravel, and sand at depths between 6 and 299 m and a<br />
temperature <strong>of</strong> 2°C.<br />
Remarks. The genus Inflatella typically has strongyles<br />
but Koltun (1959, fig. 59) described Inflatella<br />
globosa as having tylotes. So this species as described by<br />
Koltun might be transferred <strong>to</strong> ano<strong>the</strong>r genus in <strong>the</strong><br />
future. I. globosa is similar <strong>to</strong> Kirkpatrickia borealis but<br />
is more globular with oscules on <strong>to</strong>p <strong>of</strong> small papillae,<br />
while <strong>the</strong> latter species is irregularly massive with<br />
relatively inconspicuous oscules. Also, I. globosa has<br />
tylotes only, while K. borealis has tylotes, styles, and a few<br />
strongyles.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 138 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 . 2)<br />
Same specimen as in pho<strong>to</strong> 1 in situ.
55. Lissodendoryx (Lissodendoryx) behringi Koltun, 1958<br />
Description. This sponge is globular. Surface is very<br />
uneven, covered with numerous wart-like papillae.<br />
Oscula a<strong>to</strong>p <strong>the</strong> papillae contract upon collection. The<br />
contracted surface <strong>of</strong> <strong>the</strong> specimen on deck (see pho<strong>to</strong>)<br />
suggests that it looks quite different in situ. Consistency<br />
is only slightly elastic, easy <strong>to</strong> tear. Ec<strong>to</strong>some is relatively<br />
thick and easily detachable. Size is <strong>to</strong> 30 cm (diameter)<br />
× 20 cm (height). Color in life ranges from yellow <strong>to</strong><br />
light orange.<br />
Skeletal structure. This species has ec<strong>to</strong>somal <strong>to</strong>rnotes<br />
(380–750 × 10–18 µm) resembling strongyles and<br />
styles, choanosomal acanthostyles (290–680 × 20–40<br />
µm), and arcuate isochelae (42–47 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – known from <strong>the</strong><br />
Bering Sea (Russia) and <strong>the</strong> Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles and boulders<br />
at depths between 87 <strong>to</strong> 220 m. Elsewhere – reported at<br />
depths between 32 and 198 m.<br />
Remarks. Koltun (1958) mentions strongylote ec<strong>to</strong>somal<br />
megascleres only in his description <strong>of</strong> this<br />
species, but we note some individual differences<br />
71<br />
between specimens that we have examined. In some<br />
specimens strongylote forms dominate, while o<strong>the</strong>rs<br />
have mainly stylote ec<strong>to</strong>somal <strong>to</strong>rnotes. As all o<strong>the</strong>r<br />
characters and <strong>the</strong> dimensions <strong>of</strong> <strong>the</strong> spicules are within<br />
<strong>the</strong> range for L. behringi, we attribute <strong>the</strong>se differences<br />
<strong>to</strong> intraspecific variations. Gross morphology and spicule<br />
complement are similar <strong>to</strong> Phorbas paucistyliferus, but<br />
<strong>the</strong> spicule architecture is quite different.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 124 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
72 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
56. Lissodendoryx (Ectyodoryx) olgae (Hentschel, 1929)<br />
Description. This sponge is massively encrusting.<br />
Oscula are in small depressions; surface is corrugated.<br />
Consistency is s<strong>of</strong>t and very fragile. Diameter is <strong>to</strong> 15 cm.<br />
Color in life is brown <strong>to</strong> golden brown.<br />
Skeletal structure. There are tangentially arranged<br />
ec<strong>to</strong>somal tylotes (240–290 × 4–8 µm), <strong>of</strong>ten strongylote.<br />
Choanosomal acanthostyles are in two size categories<br />
– large (235–310 × 12–18 µm), and smaller echinating<br />
acanthostyles (120–160 × 6–8 µm). There are large<br />
arcuate chelae (48–52 µm), small isochelae (15–20 µm),<br />
and sigmas (48–62 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – Arctic Ocean<br />
(western Barents Sea) and North Atlantic Ocean (near<br />
Norway).<br />
Habitat. In Alaska – attached <strong>to</strong> small cobbles and<br />
pebbles in generally low-relief (i.e., flat-bot<strong>to</strong>med) habitat<br />
and at depths between 150 and 160 m. Elsewhere –<br />
reported at depths between 130 and 210 m.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands growing with <strong>the</strong> gorgonian<br />
Plumarella sp. Grid marks are 1 cm 2 .
57. Lissodendoryx (Lissodendoryx) oxeota Koltun, 1958<br />
Description. This sponge is massive-lobate and encrusting.<br />
Surface has small papillae. A thick ec<strong>to</strong>somal<br />
membrane covers an underlying honeycombed structure<br />
<strong>of</strong> <strong>the</strong> ec<strong>to</strong>some. Consistency is elastic and compressible.<br />
Diameter is <strong>to</strong> 15 cm. Color in life is golden<br />
brown.<br />
Skeletal structure. There are ec<strong>to</strong>somal oxeas and tylotes<br />
(239–322 × 9–12 µm), choanosomal acanthostyles<br />
(231–426 × 12–18 µm), and arcuate isochelae (16–<br />
21 µm).<br />
Zoogeographic distribution. Rare. In Alaska – central<br />
Aleutian Islands. Elsewhere – previously known from<br />
only two records in <strong>the</strong> Sea <strong>of</strong> Okhotsk (east <strong>of</strong> Sakhalin<br />
Island and in Kuril Strait).<br />
Habitat. In Alaska – encrusts o<strong>the</strong>r demo<strong>sponges</strong> at<br />
depths between 758 and 955 m. Elsewhere – reported<br />
at depths between 100 and 110 m.<br />
Remarks. Aleutian Island specimens encrust near <strong>the</strong><br />
base <strong>of</strong> <strong>the</strong> demo<strong>sponges</strong> Esperiopsis flagrum and Abes<strong>to</strong>pluma<br />
ramosa.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 954 m<br />
in <strong>the</strong> central Aleutian Islands. Specimen is encrusting<br />
<strong>the</strong> demosponge Esperiopsis flagrum near <strong>the</strong> base.<br />
Grid marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1<br />
(indicated by <strong>the</strong> white arrow) in situ. The separation<br />
between <strong>the</strong> red laser marks is 10 cm.<br />
73
74 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
58. Lissodendoryx (Lissodendoryx) papillosa Koltun, 1958<br />
Description. This species appears as described by<br />
Koltun (1958) as a globular or massive-lobate sponge,<br />
only slightly elastic and up <strong>to</strong> 6 cm in height. It bears<br />
closely spaced cone-shaped papillae and has an easily<br />
detachable dermal membrane. Color in life is gold <strong>to</strong><br />
golden brown.<br />
Skeletal structure. Spicules form an irregular mesh.<br />
Styles (495–1050 × 15–24 µm) are smooth, generally<br />
slightly acanthose at <strong>the</strong> blunt end. Ec<strong>to</strong>somal tylotes<br />
(275–415 × 8–10 µm) have acanthose ends. Microscleres<br />
are arcuate isochelae (25–38 µm).<br />
Zoogeographic distribution. Rare. In Alaska – Bering<br />
Sea (Zhemchug Canyon). Elsewhere – previously known<br />
only from <strong>the</strong> Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – attached <strong>to</strong> hexactinellid sponge<br />
skele<strong>to</strong>ns at a depth <strong>of</strong> 911 m. Elsewhere – no information<br />
available.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 911 m<br />
in Zhemchug Canyon, Bering Sea, with associated<br />
ophiuroids.
59. Monanchora alaskensis (Lambe, 1895)<br />
Description. This stalked sponge widens from a<br />
relatively narrow peduncle <strong>to</strong> <strong>the</strong> appearance <strong>of</strong> many<br />
agglutinated tubes (up <strong>to</strong> eight tubes). Growth form<br />
may also be flabellate, dactylate, or irregularly lobate<br />
(Koltun, 1959). Surface is smooth; large oscula are arranged<br />
in rows on <strong>the</strong> apical surface. The consistency<br />
is elastic and compressible but difficult <strong>to</strong> tear due <strong>to</strong><br />
<strong>the</strong> polyspicular tracts <strong>of</strong> <strong>the</strong> choanosome. Height is <strong>to</strong><br />
at least 17 cm. Color in life is golden brown <strong>to</strong> brown.<br />
Skeletal structure. There are ec<strong>to</strong>somal thin styles<br />
(120–200 × 7–9 µm), choanosomal styles (180–360 ×<br />
13–20 µm) that form an irregular meshwork <strong>of</strong> polyspicular<br />
tracts, anchorate isochelae (70–90 µm), and<br />
small isochelae (30–40 µm).<br />
Zoogeographic distribution. Widespread but uncommon<br />
in <strong>the</strong> North Pacific Ocean. In Alaska – central<br />
Aleutian Islands and Bering Sea. Elsewhere – Bering Sea<br />
(Russia) and Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock and cobbles<br />
at depths between 146 and 364 m. Elsewhere – on peb-<br />
bles, rocks, and sand at depths between 32 and 148 m<br />
and temperatures <strong>of</strong> 1.4 <strong>to</strong> 6.5°C.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 146 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
75
76 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
60. Monanchora laminachela Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge is stalked with a subglobular<br />
body and a corrugated surface in life which becomes<br />
conulose after freezing. Small circular oscula in depressions<br />
on <strong>the</strong> surface are only visible in live specimens.<br />
Consistency is elastic but firm. Total height is <strong>to</strong> 10<br />
cm; smooth stalk with a height <strong>of</strong> 4 cm. Color in life is<br />
golden yellow.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Ec<strong>to</strong>somal skele<strong>to</strong>n consists <strong>of</strong> a thick<br />
membrane <strong>of</strong> tangential brushes <strong>of</strong> smaller styles,<br />
fairly dense, and isochelae in between. Choanosome<br />
consists <strong>of</strong> thick, ascending polyspicular tracts <strong>of</strong> large,<br />
thick styles connected by shorter tracts <strong>of</strong> large styles.<br />
Ascending tracts (114–387 µm in diameter) support<br />
<strong>the</strong> surface tubercles and are visible <strong>to</strong> <strong>the</strong> unaided eye<br />
when dissecting <strong>the</strong> sponge. Spicules consist <strong>of</strong> megascleres<br />
– thin, small, ec<strong>to</strong>somal (sub)tylostyles (350–395<br />
× 9–11 µm), and choanosomal thick styles (840–1170 ×<br />
34–42 µm). Two categories <strong>of</strong> microscleres are present –<br />
anchorate isochelae with a peculiar plate in <strong>the</strong> middle<br />
(22–25 µm) and thin sigmas (19–23 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Attached <strong>to</strong> boulders and cobbles at depths<br />
between 203 and 485 m.<br />
Pho<strong>to</strong>s. 1) Preserved specimens (frozen) collected at<br />
a depth <strong>of</strong> 485 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimens as in pho<strong>to</strong> 1 with<br />
o<strong>the</strong>r biota. 3) Same specimens as in pho<strong>to</strong>s 1 and 2<br />
(indicated by <strong>the</strong> white arrows) in situ.
61. Monanchora pulchra (Lambe, 1894)<br />
Description. This sponge is flabellate with flattened,<br />
<strong>of</strong>ten coalesced branches. It is typically uniplanar in<br />
high current areas and multi-planed, even bushy, in<br />
areas <strong>of</strong> lower current. The surface is smooth and <strong>the</strong><br />
consistency is elastic. Height and width are <strong>to</strong> at least<br />
50 cm. Color in life is orange, yellow, or brownish<br />
orange.<br />
Skeletal structure. Ec<strong>to</strong>somal subtylostyles are scattered<br />
without obvious orientation. There are ec<strong>to</strong>somal<br />
styles <strong>to</strong> subtylostyles (225–680 × 8–12 µm). Choanosomal<br />
styles <strong>to</strong> subtylostyles are longer and thicker<br />
(670–990 × 28–40 µm). Isochelae are anchorate (18–22<br />
µm). Sigmas (16–20 µm) are typically present.<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – Aleutian Islands. Elsewhere – Pacific Coast <strong>of</strong><br />
<strong>the</strong> Kuril Islands and Pacific Coast <strong>of</strong> Canada (British<br />
Columbia).<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, boulders,<br />
cobbles, and pebbles at depths between 79 and 330 m.<br />
Elsewhere – attached <strong>to</strong> pebbles and rocks at depths<br />
between 87 and 232 m and at temperatures between<br />
1.8 and 5.3°C.<br />
Remarks. May be preyed upon by <strong>the</strong> sea star Henricia<br />
sp.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> a specimen collected at a<br />
depth <strong>of</strong> 80 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in<br />
situ (center) with a lighter-colored specimen (right). 3)<br />
Specimen collected at a depth <strong>of</strong> 150 m in <strong>the</strong> central<br />
Aleutian Islands.<br />
77
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62. Crella brunnea (Hansen, 1885)<br />
Description. This sponge is polymorphic; may be tabular,<br />
irregularly compressed, occasionally massive-lobate<br />
or markedly lobate with an uneven, corrugated surface.<br />
The dermal membrane is thinly pellicular. Oscula are<br />
not visible on <strong>the</strong> surface. Dimensions are <strong>to</strong> 7 cm. Color<br />
in life is light <strong>to</strong> dark brown.<br />
Skeletal structure. Ec<strong>to</strong>somal <strong>to</strong>rnotes have 2–4<br />
pointed “teeth” at <strong>the</strong> ends (150–290 × 4–7 µm). There<br />
are choanosomal acanthostyles <strong>to</strong> acanthostrongyles<br />
(210–380 × 10–21 µm) and small anchorate isochelae<br />
(23–37 µm). Koltun (1959) additionally reports large<br />
anchorate isochelae (44–84 µm); <strong>the</strong>se were lacking in<br />
specimens we examined from <strong>the</strong> Aleutian Islands.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
central Aleutian Islands. Elsewhere – Arctic Ocean (Barents<br />
Sea – White Sea, and Kara Sea) and North Atlantic<br />
Ocean (Norwegian Sea, Denmark Strait, Davis Strait).<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles at a depth <strong>of</strong><br />
190 m. Elsewhere – reported at depths between 20 and<br />
300 m, temperatures <strong>of</strong> 1.5 <strong>to</strong> 4.95°C, and salinities <strong>of</strong><br />
27.68 <strong>to</strong> 35.01 psu.<br />
Pho<strong>to</strong>. 1) Fragmented specimen collected at a depth<br />
<strong>of</strong> 190 m in <strong>the</strong> central Aleutian Islands. Grid marks<br />
are 1 cm 2 .
63. Hymedesmia (Stylopus) dermata Lundbeck, 1910<br />
Description. This sponge is small, thin, and encrusting.<br />
Oscula are not visible on <strong>the</strong> surface. Consistency<br />
is s<strong>of</strong>t. Color in life is light orange; beige after preservation<br />
in ethanol.<br />
Skeletal structure. There are polytylote subtylostyles<br />
<strong>to</strong> subtylostrongyles (380–450 × 5–6 µm), erect on <strong>the</strong><br />
substrate with <strong>the</strong> points facing out, and acanthostyles<br />
(430–470 × 10–12 µm).<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – Aleutian Islands. Elsewhere – Arctic<br />
Ocean (Barents Sea, Laptev Sea, Kara Sea – Vilkitsky<br />
Strait, and Greenland Sea) and North Atlantic Ocean<br />
(between Faroe Islands and Iceland).<br />
Habitat. In Alaska – attached <strong>to</strong> mollusk shells at a<br />
depth <strong>of</strong> 146 m. Elsewhere – attached <strong>to</strong> rocks and<br />
pebbles on sandy ooze at depths between 91 and 410<br />
m, <strong>water</strong> temperatures <strong>of</strong> 1.47 <strong>to</strong> 1.17°C, and salinities<br />
<strong>of</strong> 34.42 <strong>to</strong> 34.65 psu.<br />
Pho<strong>to</strong>s. 1) Preserved specimen (in ethanol) attached<br />
<strong>to</strong> a green false jingle (Pododesmus macrochisma) shell<br />
collected at a depth <strong>of</strong> 146 m in <strong>the</strong> central Aleutian<br />
Islands. Grid marks are 1 cm 2 . 2) Close-up view <strong>of</strong> same<br />
specimen as in pho<strong>to</strong> 1.<br />
79
80 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
64. Hymedesmia (Hymedesmia) irregularis Lundbeck, 1910<br />
Description. This sponge is thinly encrusting with a<br />
smooth or partially hispid surface. According <strong>to</strong> Koltun<br />
(1959) <strong>the</strong>re are oscula a<strong>to</strong>p broad papillae. Color in<br />
life is greyish yellow or bluish white.<br />
Skeletal structure. There are acanthostyles (125–540 ×<br />
16–32 µm). Ec<strong>to</strong>somal spicules are polytylote strongyles<br />
(280–410 × 6–10 µm) tangentially arranged in bundles.<br />
Microscleres are arcuate isochelae (32–48 µm).<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – Bering Sea (Pribil<strong>of</strong> Canyon). Elsewhere<br />
– Arctic Ocean (White Sea) and North Atlantic<br />
Ocean (Davis Strait, Denmark Strait, southwest <strong>of</strong> Iceland<br />
near <strong>the</strong> Faroe Islands).<br />
Habitat. In Alaska – grows on boulders and cobbles at<br />
depths around 300 m. Elsewhere – reported at depths<br />
between 293 and 1441 m.<br />
Pho<strong>to</strong>. 1) Fragments <strong>of</strong> a preserved (frozen and <strong>the</strong>n<br />
s<strong>to</strong>red in ethanol) specimen collected at a depth <strong>of</strong> 300<br />
m in Pribil<strong>of</strong> Canyon, Bering Sea.
65. Kirkpatrickia borealis Koltun, 1970<br />
Description. In situ this species has a globular body<br />
covered with numerous papillae and possesses large<br />
circular oscula. On deck specimens typically collapse<br />
completely and appear <strong>to</strong> be irregularly massive-lobate<br />
without papillae and oscula. Surface is smooth and<br />
covered by a thin ec<strong>to</strong>somal membrane. Circular oscula<br />
on <strong>the</strong> surface are slightly elevated and close upon<br />
collection. The consistency is elastic and compressible.<br />
Diameter is <strong>to</strong> 10 cm. Color in life is yellow, orange, or<br />
golden brown.<br />
Skeletal structure. There are ec<strong>to</strong>somal tylotes (350–<br />
440 × 4–7 µm); choanosomal styles include a few strongyles<br />
(470–680 × 15–22 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – Northwest<br />
Pacific Ocean (previously known only from <strong>of</strong>f Shikotan<br />
Island, Kuril Islands).<br />
Habitat. In Alaska – attached <strong>to</strong> boulders and cobbles<br />
at depths between 82 and 426 m. Elsewhere – reported<br />
at depths between 472 and 479 m.<br />
Remarks. K. borealis is similar <strong>to</strong> Inflatella globosa but<br />
is irregularly massive with relatively inconspicuous<br />
oscules, while <strong>the</strong> latter species is more globular with<br />
oscules on <strong>to</strong>p <strong>of</strong> small papillae. Also, K. borealis has<br />
tylotes, styles, and a few strongyles while I. globosa has<br />
tylotes only.<br />
Pho<strong>to</strong>s. 1) Collapsed specimen collected at a depth <strong>of</strong><br />
146 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ.<br />
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66. Phorbas paucistylifer Koltun, 1958<br />
Description. This sponge is globular with two distinct<br />
surface morphologies; some specimens are covered with<br />
round, crater-like depressions and round oscula that are<br />
flush with <strong>the</strong> surface, o<strong>the</strong>rs have a surface covered<br />
with short papillae. Dermal membrane is thick. Consistency<br />
is s<strong>of</strong>t and elastic. Diameter is <strong>to</strong> 20 cm. Color in<br />
life is bright orange (dominant phase) or yellow.<br />
Skeletal structure. Aniso<strong>to</strong>rnotes (420–490 × 8–12<br />
µm) with long thin points form a dense ec<strong>to</strong>somal<br />
crust <strong>to</strong>ge<strong>the</strong>r with numerous isochelae. Choanosomal<br />
ascending plumose spicule tracts are <strong>of</strong> <strong>the</strong> same <strong>to</strong>rnotes,<br />
cored and echinated by acanthostyles (280–540<br />
× 10–30 µm). Anchorate isochelae are strongly bent,<br />
<strong>the</strong> outline <strong>of</strong> a half circle and tridentate (32–40 µm).<br />
Koltun (1959) described somewhat smaller <strong>to</strong>rnotes and<br />
acanthostyles.<br />
Zoogeographic distribution. North Pacific Ocean.<br />
Locally common. In Alaska – central Aleutian Islands.<br />
Elsewhere – Sea <strong>of</strong> Okhotsk and Sea <strong>of</strong> Japan.<br />
Habitat. In Alaska – patchily distributed on bedrock<br />
and cobbles at depths between 56 and 120 m. Elsewhere<br />
– reported on rocks and ooze at depths between 3 and<br />
414 m and a temperature <strong>of</strong> 2.3°C.<br />
Remarks. Gross morphology and spicule complement<br />
are similar <strong>to</strong> Lissodendoryx behringi, but <strong>the</strong> spicule architecture<br />
is quite different.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 124 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ with <strong>the</strong> gorgonian<br />
Fanellia compressa. 3) Specimen collected at a depth<br />
<strong>of</strong> 110 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 . 4) Same specimen as in pho<strong>to</strong> 3 in situ.
67. Melonanchora globogilva Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge is globular. The surface is<br />
covered with numerous thin-walled, bulbous fistules,<br />
4–8 mm high and up <strong>to</strong> 4 mm in width; widest at <strong>the</strong><br />
distal end. The ec<strong>to</strong>some is a translucent white layer, easily<br />
detachable, 400–650 µm thick. Diameter is <strong>to</strong> about<br />
10 cm. Color in life is light yellow with whitish, almost<br />
translucent fistules.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some has smooth tylotes<br />
partially arranged perpendicular <strong>to</strong> <strong>the</strong> surface but<br />
oriented in all directions <strong>to</strong> some degree. From <strong>the</strong><br />
choanosome <strong>the</strong>re are tracts <strong>of</strong> tylotes fanning out <strong>to</strong>wards<br />
<strong>the</strong> ec<strong>to</strong>some. Tylotes with acanthostyles are also<br />
found in <strong>the</strong> choanosome. Tylotes are 640–680 × 10–<br />
12 µm. Choanosomal acanthostyles (660–670 × 20–<br />
30 µm) are singly distributed without recognizable orientation.<br />
Microscleres are three categories <strong>of</strong> isochelae:<br />
large isochelae with fimbriae (65–93 µm), large isochelae<br />
<strong>of</strong> <strong>the</strong> same size but with dented outer margins, and<br />
small anchorate isochelae (23–25 µm).<br />
Zoogeographic distribution. Rare. In Alaska – central<br />
Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> pebbles and shell hash at a depth<br />
<strong>of</strong> 173 <strong>to</strong> 190 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 190 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Specimen (indicated by <strong>the</strong> white arrow) observed at<br />
a depth <strong>of</strong> 179 m in <strong>the</strong> central Aleutian Islands.<br />
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68. Myxilla (Myxilla) behringensis Lambe, 1895<br />
Description. This sponge is massive and irregularly<br />
globular. Surface is smooth. Consistency is slightly elastic<br />
and ra<strong>the</strong>r firm due <strong>to</strong> <strong>the</strong> polyspicular tracts <strong>of</strong> <strong>the</strong><br />
choansosome. Diameter is <strong>to</strong> at least 4–5 cm. Color in<br />
life is whitish <strong>to</strong> golden brown.<br />
Skeletal structure. Tylotes (220–270 × 6–10 µm) are<br />
ec<strong>to</strong>somal. Choanosomal acanthostyles (250–380 ×<br />
10–15 µm) occur in a meshwork <strong>of</strong> polyspicular tracts.<br />
There are anchorate large isochelae (70–80 µm), small<br />
isochelae (15–18 µm), large sigmas (65–78 µm), and<br />
small sigmas (27–32 µm).<br />
Zoogeographic distribution. Widespread but rare in<br />
<strong>the</strong> North Pacific Ocean. In Alaska – central Aleutian<br />
Islands and Bering Sea. Elsewhere – Sea <strong>of</strong> Okhotsk,<br />
Sea <strong>of</strong> Japan, and British Columbia (Vancouver Island).<br />
Habitat. In Alaska – encrusts o<strong>the</strong>r demo<strong>sponges</strong> at<br />
depths between 190 and 195 m. Elsewhere – reported at<br />
depths between 32 and 104 m and a salinity <strong>of</strong> 33.95 psu.<br />
Remarks. In Alaska, <strong>the</strong> species is epizoic on <strong>the</strong><br />
demosponge Halichondria oblonga. It was considered by<br />
Koltun (1958) as a subspecies <strong>of</strong> Myxilla incrustans.<br />
Pho<strong>to</strong>. 1) Specimen (center) collected at a depth <strong>of</strong><br />
195 m in <strong>the</strong> central Aleutian Islands. Grid marks are<br />
1 cm 2 . Specimen is encrusting <strong>the</strong> demosponge Halichondria<br />
oblonga.
69. Myxilla (Ectyomyxilla) parasitica Lambe, 1893<br />
Description. This sponge is encrusting, typically on<br />
<strong>the</strong> shells <strong>of</strong> scallops (Chlamys sp.). Surface is finely<br />
structured with radiating canals leading <strong>to</strong> small oscula<br />
that are flush with <strong>the</strong> surface. Thin encrustation is<br />
less than 1 cm in height and typically covers <strong>the</strong> entire<br />
available shell surface. Color in life is light brown <strong>to</strong><br />
golden brown.<br />
Skeletal structure. Ec<strong>to</strong>somal <strong>to</strong>rnotes have unequal<br />
ends and mammillate projections at <strong>the</strong>se ends (160–<br />
186 × 7–9 µm). Choanosomal acanthostyles are in an irregular<br />
meshwork (208–310 × 10–15 µm); microscleres<br />
are large isochelae (52–65 µm), small isochelae (15–20<br />
µm), and thin sigmata (17–25 µm).<br />
Zoogeographic distribution. Locally abundant in <strong>the</strong><br />
North Pacific Ocean. In Alaska – central Aleutian Islands.<br />
Elsewhere – Bering Sea (Russia), Sea <strong>of</strong> Okhotsk<br />
(near <strong>the</strong> Kuril Islands) and British Columbia (near<br />
Vancouver Island).<br />
Habitat. In Alaska – encrusts scallop shells at depths<br />
between 98 and 250 m. Elsewhere – on rocks, pebbles,<br />
and sand at depths between 15 and 126 m.<br />
Remarks. They are <strong>of</strong>ten very abundant in scallop<br />
beds.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 138 m<br />
in <strong>the</strong> central Aleutian Islands. Specimen completely<br />
encrusts a scallop (Chlamys sp.). Grid marks are 1 cm 2 .<br />
2) Specimen (center) collected at a depth <strong>of</strong> 98 m in<br />
<strong>the</strong> central Aleutian Islands. 3) A bed <strong>of</strong> scallops (Chlamys<br />
sp.), almost all <strong>of</strong> which are completed encrusted<br />
with Myxilla parasitica. The gorgonian Fanellia fraseri is<br />
at center.<br />
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70. Myxilla (Bur<strong>to</strong>nanchora) pedunculata Lundbeck, 1905<br />
Description. This sponge is globular or lobate, usually<br />
with a stalk. The surface is slightly hispid. Irregularly<br />
scattered circular oscula are flush with <strong>the</strong> surface. Consistency<br />
is s<strong>of</strong>t and elastic. Height and width are <strong>to</strong> 12<br />
cm. Color in life is yellow <strong>to</strong> bright yellow.<br />
Skeletal structure. Spicules are arranged in long ascending<br />
fibers, connected by shorter spicule tracts and<br />
single spicules. There are smooth or slightly acanthose<br />
styles (270–551 × 10–20 µm), ec<strong>to</strong>somal <strong>to</strong>rnotes (220–<br />
380 × 6–10 µm), and microscleres that are anchorate<br />
isochelae (42–75 µm).<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – Bering Sea (Pribil<strong>of</strong> Canyon).<br />
Elsewhere – Arctic Ocean (Kara Sea – Vilkitsky Strait,<br />
Laptev Sea) and North Atlantic Ocean (Iceland and<br />
Faroe Islands).<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles and cobbles<br />
at depths between 204 and 236 m. Elsewhere – reported<br />
at depths between 28 and 325 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 204 m<br />
in Pribil<strong>of</strong> Canyon, Bering Sea. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 showing <strong>the</strong> detail <strong>of</strong><br />
<strong>the</strong> oscula. 3) Specimen in situ collected at a depth <strong>of</strong><br />
236 m in Pribil<strong>of</strong> Canyon, Bering Sea. The demosponge<br />
Craniella spinosa is attached <strong>to</strong> <strong>the</strong> specimen. The separation<br />
between <strong>the</strong> red laser marks is 10 cm.
71. Stelodoryx oxeata Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This species exhibits two growth forms.<br />
One is a stalked, conical form with a ridged surface.<br />
The o<strong>the</strong>r is a massively encrusting, lobate sponge with<br />
a smooth surface. Consistency is ra<strong>the</strong>r hard and only<br />
slightly elastic, due <strong>to</strong> <strong>the</strong> densely packed spicules in<br />
<strong>the</strong> ec<strong>to</strong>some. The stalked specimen has a stalk <strong>of</strong> 2 ×<br />
0.7 cm, and <strong>the</strong> body is 5 × 4 cm in greatest dimensions.<br />
Color in life for both growth forms is greenish yellow<br />
<strong>to</strong> light green.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Ec<strong>to</strong>some is densely packed with a tangential<br />
arrangement <strong>of</strong> <strong>to</strong>rnotes, peculiar shaped oxeas,<br />
and microscleres. The choanosome consists in places<br />
<strong>of</strong> an irregular reticulation <strong>of</strong> single spicules or short<br />
spicule tracts. Megascleres are oxeas (517–558 × 20–<br />
30 µm); <strong>the</strong> points <strong>of</strong> oxeas have a ragged, dented<br />
outline. Tornotes have acanthose ends (230–270 × 9–<br />
11 µm). Microscleres are large isochelae (54–110 µm),<br />
medium-sized isochelae (23–32 µm), small anchorate<br />
isochelae (9–13 µm), and thin centrotylote sigmas<br />
(8–12 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> cobbles and pebbles at depths<br />
between 175 and 712 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 395 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Specimen (indicated by <strong>the</strong> white arrow) collected<br />
at a depth <strong>of</strong> 711 m in <strong>the</strong> central Aleutian Islands.<br />
The separation between <strong>the</strong> red laser marks is 10 cm.<br />
3) Specimen (<strong>to</strong>p, center) collected at a depth <strong>of</strong> 175 m<br />
in <strong>the</strong> central Aleutian Islands. The separation between<br />
<strong>the</strong> red laser marks is 10 cm.<br />
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72. Stelodoryx <strong>to</strong>poroki Koltun, 1958<br />
Description. This sponge is stalked, flabellate, or a<br />
somewhat flattened funnel. Consistency is elastic with a<br />
thin ec<strong>to</strong>somal membrane. It is s<strong>of</strong>t and floppy in situ.<br />
Height is <strong>to</strong> 17 cm and width <strong>to</strong> 12 cm. Color in life is<br />
light yellow; dark brown after freezing.<br />
Skeletal structure. Ec<strong>to</strong>somal tylotes have acanthaceous<br />
ends (218–300 × 8–10 µm). Choanosomal styles<br />
(500–1140 × 21–30 µm) occasionally have acanthaceous<br />
heads, and anchorate isochelae are in two size categories<br />
– large (115–150 µm) and small (30–40 µm).<br />
Zoogeographic distribution. Locally common in<br />
<strong>the</strong> North Pacific Ocean. In Alaska – central Aleutian<br />
Islands. Elsewhere – Sea <strong>of</strong> Okhotsk (near <strong>the</strong> Kuril<br />
Islands).<br />
Habitat. In Alaska – occurring in small patches (up <strong>to</strong><br />
5 individuals per m 2 ) on bedrock, muds<strong>to</strong>ne, boulders,<br />
and cobbles at depths between 1714 and 2314 m. Elsewhere<br />
– found on sand, pebbles, and “reefs” at depths<br />
between 113 and 303 m and at temperatures between<br />
1.8 and 3.1°C.<br />
Remarks. A sponge with similar appearance except<br />
larger (up <strong>to</strong> 50 cm diameter) and more robust (forming<br />
distinct bowls) occurs at depths between 1185 and<br />
1540 m. Collection <strong>of</strong> a specimen within this depth<br />
range and examination <strong>of</strong> <strong>the</strong> spicules is necessary <strong>to</strong><br />
confirm conspecificity, however.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at<br />
a depth <strong>of</strong> 2176 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 (lower<br />
center) in situ. 3) Close-up view <strong>of</strong> specimen in situ at a<br />
depth <strong>of</strong> 1505 m in <strong>the</strong> central Aleutian Islands.
73. Stelodoryx vitiazi (Koltun, 1959)<br />
Description. This sponge is tube-shaped, sometimes<br />
irregularly cylindrical, with a hollow center. It occurs<br />
in clusters <strong>of</strong> up <strong>to</strong> five individuals and is fragile and<br />
inelastic. Height is <strong>to</strong> 15 cm and width <strong>to</strong> 4 cm. Color<br />
in life is light yellow <strong>to</strong> golden brown.<br />
Skeletal structure. Tangentially arranged <strong>to</strong>rnotes<br />
have acanthaceous ends in <strong>the</strong> ec<strong>to</strong>some. There is an irregular<br />
meshwork <strong>of</strong> paucispicular tracts and individual<br />
acanthostyles in <strong>the</strong> choanosome. There are <strong>to</strong>rnotes with<br />
acanthose ends (180–291 × 4–7 µm), acanthostyles (370–<br />
520 × 20–29 µm), and anchorate isochelae (25–46 µm).<br />
Zoogeographic distribution. Locally common in<br />
<strong>the</strong> North Pacific Ocean. In Alaska – central Aleutian<br />
Islands. Elsewhere – Bering Sea (near <strong>the</strong> Commander<br />
Islands) and <strong>the</strong> Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles and pebbles at<br />
depths between 155 m and 1009 m. Elsewhere – found<br />
on sand, pebbles, and “reefs” at depths between 115<br />
and 820 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 155 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Specimen collected at a depth <strong>of</strong> 1009 m in <strong>the</strong> central<br />
Aleutian Islands. 3) Same specimen as in pho<strong>to</strong> 2<br />
in situ.<br />
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74. Echinostylinos hirsutus Koltun, 1970<br />
Description. This sponge is short with a cylindrical<br />
body and hollow central cavity. Koltun (1970) reported<br />
that this species was stalked, but our observations indicate<br />
that it narrows near <strong>the</strong> base and does not necessarily<br />
have a true stalk. Surface is corrugated. Height is <strong>to</strong> 5<br />
cm. Color in life is light brown <strong>to</strong> light yellow.<br />
Skeletal structure. There are two categories <strong>of</strong> styles:<br />
long, 1320 × 25 µm (Koltun: 660–1650 × 45–54 µm), and<br />
shorter, 350 × 9 µm (Koltun: 400–540 × 5–10 µm). It has<br />
isochelae (sigmancistras) 27–30 µm (Koltun: sigmoid<br />
arcuate chelae 18–27 µm) and sigmas 40 µm (Koltun:<br />
filiform sigmas 16–22 µm). We regard <strong>the</strong> differences in<br />
spicule sizes as intraspecific variation.<br />
Zoogeographic distribution. Apparently a very rare<br />
species; known from only two locations. In Alaska –<br />
central Aleutian Islands. Elsewhere – Sea <strong>of</strong> Okhotsk.<br />
Habitat. In Alaska – occurring singly on cobbles and<br />
pebbles at depths between 665 and 711 m. Elsewhere –<br />
reported at depths between 1440 and 1540 m.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at a<br />
depth <strong>of</strong> 711 m in <strong>the</strong> central Aleutian Islands. 2) Same<br />
specimen as in pho<strong>to</strong> 1 (indicated by <strong>the</strong> white arrow)<br />
in situ on cobble.
75. Tedania (Tedania) dirhaphis Hentschel, 1912<br />
Description. This sponge is irregularly massive-lobate,<br />
<strong>of</strong>ten with dactylate processes bearing circular oscula on<br />
<strong>to</strong>p. The surface is smooth. The consistency is s<strong>of</strong>t and<br />
elastic. Height and width are <strong>to</strong> 20 cm. Color in life is<br />
yellow <strong>to</strong> golden yellow; brown after freezing.<br />
Skeletal structure. Styles are smooth (238–565 × 9–18<br />
µm). Ec<strong>to</strong>somal tylotes have slightly acanthose ends<br />
(215–487 × 5–8 µm). Microscleres are rhaphides in two<br />
size categories – large rhaphides (200–400 µm) and<br />
small rhaphides (45–180 µm).<br />
Zoogeographic distribution. North and West Pacific<br />
Ocean. Uncommon. In Alaska – central Aleutian Islands<br />
and Bering Sea (Pribil<strong>of</strong> Canyon). Elsewhere – Sea <strong>of</strong><br />
Okhotsk, Pacific Coast <strong>of</strong> Kuril Islands, and South China<br />
Sea.<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles at depths between<br />
100 m and 341 m. Elsewhere – reported at depths<br />
between 4 and 550 m.<br />
Remarks. Tedania dirhaphis was originally described<br />
by Hentschel (1912) from shallow Indonesian <strong>water</strong>s.<br />
Koltun (1959) reported <strong>the</strong> same species from <strong>the</strong> Sea<br />
<strong>of</strong> Okhotsk and <strong>the</strong> Kuril Islands in <strong>the</strong> North Pacific<br />
Ocean and <strong>the</strong> Alaskan specimens conform perfectly<br />
<strong>to</strong> Koltun’s decription. The Alaskan specimens also<br />
conform <strong>to</strong> <strong>the</strong> original description (p. 45 and pl. XIX,<br />
fig. 20) in spicule complement, growth form, and color<br />
(Hentschel, 1912) and differ from <strong>the</strong> Indonesian specimens<br />
in size only (somewhat larger). Accordingly, we<br />
report this species as T. dirhapsis sensu Koltun, 1959,<br />
but caution that given <strong>the</strong> extremes in zoogeography,<br />
additional taxonomic work on this species is warranted.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 100 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Preserved (frozen) specimen collected at a depth <strong>of</strong><br />
341 m in Pribil<strong>of</strong> Canyon, Bering Sea. Grid marks are<br />
1 cm 2 . 3) Same specimen as in pho<strong>to</strong> 1 in situ with a<br />
shortspine thornyhead (Sebas<strong>to</strong>lobus alascanus) at left.<br />
The separation between <strong>the</strong> red laser marks is 10 cm.<br />
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76. Tedania kagalaskai Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This large flabellate sponge is attached <strong>to</strong><br />
substrate with a firm stalk that widens about 10 cm above<br />
<strong>the</strong> holdfast. Surface is very smooth; oscula are in several<br />
rows on <strong>the</strong> flattened apical surface. Body is s<strong>of</strong>t; stalk<br />
is wiry. Maximum dimensions <strong>of</strong> <strong>the</strong> roughly triangular<br />
body are approximately 30 × 30 × 5 cm. Color in life is<br />
light brown <strong>to</strong> orange-brown.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Ec<strong>to</strong>somal tylotes (300–330 × 4–6 µm)<br />
have acanthose ends. There are choanosomal smooth<br />
styles (360–390 × 15–20 µm), and onychaetes possibly<br />
in two size categories – large (170–210 µm) and small<br />
(30–40 µm).<br />
Zoogeographic distribution. Locally common. In Alaska<br />
– central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> bedrock, boulders, and cobbles<br />
at depths between 59 and 170 m. Specimens with a similar<br />
gross morphology have been observed at depths near<br />
478 m, but collection <strong>of</strong> a specimen within this depth<br />
range and examination <strong>of</strong> <strong>the</strong> spicules is necessary <strong>to</strong><br />
confirm conspecificity.<br />
Remarks. Species is similar in appearance <strong>to</strong> Artimisina<br />
stipitata with which it co-occurs. The dorsal surface <strong>of</strong><br />
T. kagalaskai is typically flatter and <strong>the</strong> body form more<br />
distinctly triangular and with a much smoo<strong>the</strong>r surface<br />
than A. stipitata. This is <strong>the</strong> only known species <strong>of</strong> Tedania<br />
that is stalked.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 146 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 . 2)<br />
Same specimen as in pho<strong>to</strong> 1 in situ. 3) Specimen collected<br />
at a depth <strong>of</strong> 155 m in <strong>the</strong> central Aleutian Islands.<br />
Grid marks are 1 cm 2 . 4) Specimen in situ collected at a<br />
depth <strong>of</strong> 105 m in <strong>the</strong> central Aleutian Islands.
77. Asbes<strong>to</strong>pluma ramosa Koltun, 1958<br />
Description. This sponge is flagelliform. This species<br />
appears as described by Koltun (1959) and <strong>of</strong>ten<br />
consists <strong>of</strong> numerous more or less cylindrical branches<br />
diverging from one point and arranged in one plane.<br />
We have observed up <strong>to</strong> five branches that diverge at<br />
multiple points, however. Branches may be bent or<br />
slightly curved. Occasionally <strong>the</strong> sponge is markedly<br />
elongated and has a ramified, stalk-like appearance<br />
with many closely spaced fine projections (according <strong>to</strong><br />
Koltun). Height is <strong>to</strong> at least 72 cm with branches up <strong>to</strong><br />
1.5 cm thick. Color in life is light yellow, light orange,<br />
or creamy white; <strong>the</strong> lower portion is <strong>of</strong>ten devoid <strong>of</strong><br />
<strong>the</strong> fine lateral projections and consequently a darker<br />
color, <strong>of</strong>ten brown.<br />
Skeletal structure. Styles are arranged in a central<br />
axis with side-tracts running in<strong>to</strong> <strong>the</strong> lateral projections;<br />
styles are 238–1500 × 7–18 µm and anisochelae<br />
are 10–17 µm.<br />
Zoogeographic distribution. Widespread and locally<br />
abundant. In Alaska – central Aleutian Islands, eastern<br />
Gulf <strong>of</strong> Alaska (near <strong>the</strong> Fairwea<strong>the</strong>r Ground). Elsewhere<br />
– North Pacific Ocean (Kuril Islands, Vancouver<br />
Island), Arctic Ocean (Southwest Barents Sea, Kara<br />
Sea – Vilkitsky Strait, Laptev Sea – Shokalsky Strait, East<br />
Siberian Sea, Greenland Sea, and Baffin Bay), and<br />
North Atlantic Ocean (Faroe Islands).<br />
Habitat. In Alaska – attached with a holdfast <strong>to</strong> bedrock,<br />
muds<strong>to</strong>ne, boulders, cobbles, and occasionally<br />
hexactinellid sponge skele<strong>to</strong>ns at depths between 395<br />
and 1812 m; more abundant at depths shallower than<br />
1200 m. Elsewhere – reported on ooze and sandy-ooze<br />
bot<strong>to</strong>ms at depths between 41 and 1134 m and temperatures<br />
<strong>of</strong> 1.64 <strong>to</strong> 4.8°C.<br />
Remarks. This species occasionally has ophiuroid associates<br />
particularly at depths shallower than 500 m. It<br />
occurs within <strong>the</strong> same depth range as Esperiopsis flagrum<br />
but attaches <strong>to</strong> hard substrate ra<strong>the</strong>r than anchoring in<br />
s<strong>of</strong>t sediment. A. ramosa may have a carnivorous feeding<br />
habit like o<strong>the</strong>r Cladorhizidae so collected specimens<br />
may have small crustaceans trapped in <strong>the</strong>ir outer tissues.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 1715 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Specimen collected at a depth <strong>of</strong> 1501 m in <strong>the</strong><br />
eastern Gulf <strong>of</strong> Alaska. 3) Specimen observed in situ<br />
at a depth <strong>of</strong> 843 m in <strong>the</strong> central Aleutian Islands. 4)<br />
Close-up view <strong>of</strong> same specimen as in pho<strong>to</strong> 3 showing<br />
detail <strong>of</strong> <strong>the</strong> closely spaced fine projections.<br />
93
94 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
77. Asbes<strong>to</strong>pluma ramosa Koltun, 1958 (continued)
78. Cladorhiza bathycrinoides Koltun, 1955<br />
Description. This stalked sponge terminates in a partially<br />
inverted conical head. Basal root-like processes<br />
consist <strong>of</strong> glassy, polyspicular tracts (2.5 cm long). Polyspicular<br />
tracts <strong>of</strong> <strong>the</strong> root-like processes become united<br />
at <strong>the</strong> base <strong>of</strong> <strong>the</strong> cylindrical stalk and are coated by a<br />
thin ec<strong>to</strong>somal veneer. The circular upper plane <strong>of</strong> <strong>the</strong><br />
head has a diameter <strong>of</strong> 1.4 cm and thin appendages<br />
(up <strong>to</strong> 12) are inserted along its margin. The appendages<br />
measure 1.9 cm × 0.8 mm, but <strong>the</strong> diameter <strong>of</strong><br />
<strong>the</strong> appendages regularly varies like a string <strong>of</strong> pearls.<br />
Typical <strong>of</strong> <strong>the</strong> genus, <strong>the</strong> polyspicular tracts <strong>of</strong> <strong>the</strong><br />
system <strong>of</strong> root-like processes are united at <strong>the</strong> base <strong>of</strong><br />
<strong>the</strong> stalk, run in bundles through <strong>the</strong> stalk, and <strong>the</strong>n<br />
diverge again <strong>to</strong> support <strong>the</strong> appendages. The height<br />
from distal ends <strong>of</strong> <strong>the</strong> root-like processes <strong>to</strong> <strong>the</strong> upper<br />
end <strong>of</strong> <strong>the</strong> head is up <strong>to</strong> at least 12 cm. The stalk is up<br />
<strong>to</strong> 7 cm long and 1–2 mm in diameter. Color in life is<br />
white or creamy white.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. Ec<strong>to</strong>somal veneer is packed with anisochelae.<br />
Fusiform styles (980–2100 × 12–40 µm) and<br />
tylostrongyles (350–900 × 2–10 µm) are most abundant<br />
at <strong>the</strong> distal parts <strong>of</strong> <strong>the</strong> root-like processes. There are<br />
large anisochelae (55–75 µm), rare small anisochelae<br />
(25–35 µm), large sigmas (95–110 µm), small sigmas<br />
(43–55 µm), and flattened sigmoids with claw-like appendages<br />
at <strong>the</strong> ends (42–45 µm). Large sigmas were<br />
typically damaged in our SEM preparations indicating<br />
that <strong>the</strong>y are extremely thin-walled, hollow, and fragile.<br />
Zoogeographic distribution. North Pacific Ocean.<br />
Locally common. In Alaska – central Aleutian Islands;<br />
patchy distribution at low densities. Elsewhere – Sea <strong>of</strong><br />
Okhotsk.<br />
Habitat. In Alaska – on s<strong>of</strong>t sediments (silt and sand)<br />
at depths between 1108 and 2854 m. Elsewhere – eurybathic;<br />
reported at depths between 150 and 3800 m.<br />
Remarks. C. bathycrinoides probably has a carnivorous<br />
feeding habit like o<strong>the</strong>r Cladorhizidae. We observed<br />
one specimen in situ capture a small shrimp with one<br />
<strong>of</strong> its appendages (actually <strong>the</strong> shrimp swam in<strong>to</strong> <strong>the</strong><br />
appendage and appeared <strong>to</strong> adhere <strong>to</strong> it). Collected<br />
specimens may have small crustaceans trapped in <strong>the</strong>ir<br />
outer tissues.<br />
95<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 1371 m in<br />
<strong>the</strong> central Aleutian Islands and held live in a shipboard<br />
aquarium. Note <strong>the</strong> basal root-like processes. 2) Same<br />
specimen as in pho<strong>to</strong> 1 in situ. The separation between<br />
<strong>the</strong> red laser marks is 10 cm.
96 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
79. Cladorhiza corona Lehnert, Watling and S<strong>to</strong>ne 2005<br />
Description. Body consists <strong>of</strong> a long cylindrical stalk<br />
with a basal plate and two sets <strong>of</strong> distal appendages:<br />
<strong>the</strong> basal set radiating in a full circle more or less in<br />
one plane, and <strong>the</strong> distal set forming a quarter circle<br />
<strong>of</strong> triangular-shaped structures oriented in a plane<br />
almost perpendicular <strong>to</strong> <strong>the</strong> basal appendages. Distal<br />
appendages are shorter than <strong>the</strong> basal and terminate<br />
in a spherical tyle. Root-like processes (common within<br />
<strong>the</strong> genus) are absent and <strong>the</strong> sponge attaches directly<br />
<strong>to</strong> hard substrate via <strong>the</strong> broadened basal plate. Consistency<br />
is wiry and elastic, with an easily detachable<br />
ec<strong>to</strong>somal membrane. Total length is <strong>to</strong> at least 33 cm.<br />
Color in life is light yellow or creamy white.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The three distinct parts <strong>of</strong> <strong>the</strong> sponge<br />
(basal plate, stalk with <strong>the</strong> basal appendages, and <strong>the</strong><br />
crown) differ considerably in spicule types and <strong>the</strong>ir<br />
arrangement. The basal plate consists <strong>of</strong> fusiform<br />
styles and shorter anisoxeas with slightly unequal ends,<br />
packed in one plane parallel <strong>to</strong> <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substrate.<br />
The basal plate has no ec<strong>to</strong>somal membrane and<br />
is devoid <strong>of</strong> microscleres. From <strong>the</strong> basal plate polyspicular<br />
tracts <strong>of</strong> long fusiform styles run in thick bundles<br />
through <strong>the</strong> center <strong>of</strong> <strong>the</strong> long cylindrical stalk, which<br />
is covered by an ec<strong>to</strong>somal membrane, densely packed<br />
with ancorate anisochelae. The basal appendages are<br />
constructed in a similar manner as <strong>the</strong> stalk. The polyspicular<br />
tracts <strong>of</strong> long fusiform styles reaching <strong>the</strong> crown<br />
fan out in one plane, leaving little space between <strong>the</strong><br />
tracts. Single, thin (sub-) tylostyles are arranged more<br />
or less perpendicular <strong>to</strong> <strong>the</strong> thick fusiform styles. The<br />
ec<strong>to</strong>somal membrane <strong>of</strong> <strong>the</strong> crown is devoid <strong>of</strong> anisochelae<br />
but ra<strong>the</strong>r contains flattened sigmancistras with<br />
thin claw-like extensions at each end. These latter two<br />
spicule types occur exclusively in <strong>the</strong> crown. Megascleres<br />
include fusiform styles, almost anisoxeas (600–4260<br />
× 10–65 µm), present in all parts except <strong>the</strong> ec<strong>to</strong>somal<br />
membrane; (sub-) tylostyles (510–1650 × 8–20 µm)<br />
present in <strong>the</strong> ec<strong>to</strong>somal membrane <strong>of</strong> <strong>the</strong> crown; and<br />
thick, short anisoxeas with slightly unequal blunt ends<br />
(140–660 × 38–43 µm) present in <strong>the</strong> basal plate. Microscleres<br />
include anchorate anisochelae (30–42 µm),<br />
present in <strong>the</strong> ec<strong>to</strong>somal membrane <strong>of</strong> <strong>the</strong> stalk and<br />
<strong>the</strong> basal appendages; and flattened sigmancistras, each<br />
end with a thin pointed “claw” (35–42 µm), present in<br />
<strong>the</strong> ec<strong>to</strong>somal membrane <strong>of</strong> <strong>the</strong> crown.<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Occurs in small patches at depths between<br />
726 and 2077 m on bedrock, fragmented bedrock,<br />
muds<strong>to</strong>ne, and cobbles, and at temperatures between<br />
1.8 and 3.1°C.<br />
Remarks. C. corona is carnivorous and feeds mainly<br />
on calanoid copepods (Watling, 2007). Collected specimens<br />
may have small crustaceans trapped in <strong>the</strong>ir outer<br />
tissues.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 1720 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ.
80. Chondrocladia (Chondrocladia) concrescens (Schmidt, 1880)<br />
Description. This stalked sponge has numerous lateral<br />
processes with terminal swellings (as many as 80). Size<br />
and growth form <strong>of</strong> this species is variable. There are<br />
root-like tufts <strong>of</strong> polyspicular tracts fixing <strong>the</strong> sponge<br />
in <strong>the</strong> substrate. The consistency <strong>of</strong> <strong>the</strong> stalks is firm<br />
and wiry while <strong>the</strong> rest <strong>of</strong> <strong>the</strong> sponge is s<strong>of</strong>t and fragile.<br />
Thick polyspicular tracts support <strong>the</strong> stalk; thinner polyspicular<br />
tracts support <strong>the</strong> branches. Height is <strong>to</strong> 30 cm.<br />
Color in life is golden brown <strong>to</strong> light brown and reddish<br />
brown in shallower <strong>water</strong> (
98 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
81. Biemna variantia (Bowerbank, 1858)<br />
Description. This sponge is subglobular. Surface is<br />
covered with numerous papillae. Oscula are at <strong>the</strong> <strong>to</strong>p<br />
<strong>of</strong> <strong>the</strong> sponge in clusters. It is very s<strong>of</strong>t, fragile, and easily<br />
<strong>to</strong>rn. Diameter is <strong>to</strong> 10 cm. Color in life is greenish yellow<br />
<strong>to</strong> light brown; brown or dark brown after freezing.<br />
Skeletal structure. A thin ec<strong>to</strong>somal membrane contains<br />
only microscleres. There are styles (420–1450<br />
× 18–35 µm), sigmas possibly in two size categories<br />
(40–310 µm), rhaphides (60–125 µm), and spherules<br />
(10 µm). Long ascending and branching polyspicular<br />
tracts <strong>of</strong> styles in <strong>the</strong> choanosome have many spicules<br />
scattered between <strong>the</strong> tracts.<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – central Aleutian Islands. Elsewhere<br />
– North Pacific Ocean (Bering Sea – Russia), Arctic<br />
Ocean (Barents Sea and Greenland Sea), and North<br />
Atlantic Ocean (eastern Scotian Shelf east <strong>to</strong> Iceland<br />
and south <strong>to</strong> <strong>the</strong> Canary Islands; western Mediterranean<br />
Sea).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles and pebbles<br />
at depths between 155 and 489 m. Elsewhere – reported<br />
at depths between 62 and 1800 m depth.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 155 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Preserved specimen (frozen) collected at a depth <strong>of</strong><br />
489 m in <strong>the</strong> central Aleutian Islands. Grid marks are 1<br />
cm 2 . 3) Same specimen as in pho<strong>to</strong> 2 (indicated by <strong>the</strong><br />
white arrow) in situ.
82. Euchelipluma elongata Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge is flagelliform or whip-like<br />
and occasionally bifurcated. The species is similar <strong>to</strong><br />
Asbes<strong>to</strong>pluma ramosa and Esperiopsis flagrum (differentiation<br />
requires careful examination). Rigid long, thin<br />
stalk is covered with thin, relatively short processes.<br />
It is basally attached in s<strong>of</strong>t substrates with a rigid<br />
root-like system. Length is <strong>to</strong> at least 35 cm, but few<br />
specimens attain this size. Color in life is light yellow<br />
<strong>to</strong> creamy white.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some consists <strong>of</strong> densely<br />
arranged isochelas, underlain by parallel oriented<br />
fusiform styles with blunt ends and smaller tylostyles.<br />
Single tylostyles are placed perpendicular <strong>to</strong> <strong>the</strong> orientation<br />
<strong>of</strong> <strong>the</strong> styles and tylostyles. The choanosome<br />
is dominated by ascending polyspicular tracts <strong>of</strong> styles,<br />
tylostyles, and isochelae. Megascleres are blunt-ended<br />
fusiform styles (1310–1510 × 40–55 µm). Tylostyles<br />
(620–660 × 9–13 µm) <strong>of</strong>ten have <strong>the</strong> tyle subterminal<br />
and occasionally polytylote. Microscleres are isochelae<br />
(80–95 µm), placochelae (70–88 µm), and sigmas<br />
(9–25 µm).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Attached with a root-like system in unconsolidated<br />
sediments at depths between 1525 and about<br />
2200 m. Fairly abundant in some areas, reaching densities<br />
near 15 individuals per m 2 .<br />
Remarks. There are only four known species <strong>of</strong> Euchelipluma<br />
worldwide. E. elongata is by far <strong>the</strong> largest species<br />
in <strong>the</strong> genus. This species does not appear <strong>to</strong> harbor<br />
<strong>the</strong> ophiuroid associates that are so common with <strong>the</strong><br />
whip-like pennatulacean corals present in <strong>the</strong> same<br />
depth range. It may occasionally be preyed upon by <strong>the</strong><br />
99<br />
sea star Hippasteria. It does not co-occur with Esperiopsis<br />
flagrum but does overlap in depth range with Asbes<strong>to</strong>pluma<br />
ramosa. Unlike A. ramosa, however, it anchors in<br />
s<strong>of</strong>t-sediment ra<strong>the</strong>r than attaching <strong>to</strong> hard substrate.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 2161 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ (foreground).
100 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
83. Guitarra abbotti Lee, 1987<br />
Description. This sponge is subglobular and massive.<br />
Consistency is elastic, s<strong>of</strong>t, and easy <strong>to</strong> tear. Oscula are<br />
small, circular, and slightly elevated on short conical<br />
projections. The surface is smooth <strong>to</strong> <strong>the</strong> unaided eye<br />
but microscopically hispid or brain-like with narrow<br />
convolutions. Diameter is <strong>to</strong> 15 cm. Color in life is yellow,<br />
orange, or dark brown.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. There is no specialized ec<strong>to</strong>somal skele<strong>to</strong>n;<br />
<strong>the</strong> dermal membrane consists <strong>of</strong> spicule brushes<br />
at <strong>the</strong> ends <strong>of</strong> short polyspicular tracts, ascending only<br />
a short distance from <strong>the</strong> choanosome. Sections parallel<br />
<strong>to</strong> <strong>the</strong> surface show <strong>the</strong> regularly spaced spicule brushes<br />
up <strong>to</strong> 300 µm in diameter. Short polyspicular tracts <strong>of</strong><br />
styles make up a relatively dense but irregular meshwork<br />
in <strong>the</strong> choanosome, mesh-size 100–200 µm, with single<br />
spicules scattered in between. The short length <strong>of</strong> <strong>the</strong><br />
tracts gives a halichondroid appearance throughout<br />
much <strong>of</strong> <strong>the</strong> choanosome. Megascleres are fusiform<br />
styles (330–400 × 7–10 µm). There are seven types <strong>of</strong><br />
microscleres: large placochelae (105–115 µm); small<br />
placochelae (30–50 µm); large biplacochelae (35–52<br />
µm); small and relatively rare biplacochelae (13–16 µm<br />
in diameter); a rare, large category <strong>of</strong> isochelae (58-<br />
65 µm); small sigmoid isochelae (10–13 µm) almost<br />
closed; and small sigmoids (5–9 µm). All spicule-types<br />
are distributed without an obvious pattern throughout<br />
<strong>the</strong> sponge.<br />
Zoogeographic distribution. North Pacific Ocean.<br />
Locally common. In Alaska – central Aleutian Islands.<br />
Elsewhere – previously known only from Cordell Bank,<br />
California.<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles at depths between<br />
100 and 146 m. Elsewhere – reported at depths<br />
between 35 and 46 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 138<br />
m in <strong>the</strong> central Aleutian Islands. Grid marks are 1<br />
cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ (center).<br />
3) Specimen (center) collected at a depth <strong>of</strong> 146 m in<br />
<strong>the</strong> central Aleutian Islands.
84. Guitarra fimbriata Carter, 1874<br />
Description. This sponge is cushion-shaped with a tuberculate<br />
surface; no oscula visible. Consistency is only<br />
slightly compressible and not very elastic. Diameter is <strong>to</strong><br />
20 cm. Color in life is yellow <strong>to</strong> orange-brown.<br />
Skeletal structure. Spicules are anisotylotes (230–700<br />
× 5–15 µm) <strong>of</strong>ten one end conspicuously more inflated<br />
than <strong>the</strong> o<strong>the</strong>r, large placochelae (70–90 µm), small<br />
placochelae (35–55 µm), and bipocilli (6–15 µm).<br />
Zoogeographic distribution. Cosmopolitan and locally<br />
common. In Alaska – central Aleutian Islands. Elsewhere<br />
– Pacific coast <strong>of</strong> sou<strong>the</strong>rn Kuril Islands, North<br />
Atlantic Ocean, Indian Ocean, and Antarctica.<br />
Habitat. In Alaska – attached <strong>to</strong> small boulders and<br />
cobbles at a depth <strong>of</strong> 146 m. Elsewhere – reported on<br />
rocky bot<strong>to</strong>ms at depths between 28 and 188 m and at<br />
temperatures between 1.5 and 16.2°C.<br />
Pho<strong>to</strong>s. 1) Partial specimen collected at a depth <strong>of</strong> 146<br />
m in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ (center).<br />
101
102 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
85. Amphilectus digitatus (Miklucho-Maclay, 1870)<br />
Description. This species appears as described by<br />
Koltun (1959) and may be massive-lobate, laminate,<br />
dactylate, funnel-shaped, or vase-like, and usually has a<br />
stalk. The surface is smooth. The consistency is elastic,<br />
easily compressible, but difficult <strong>to</strong> tear. Vase-like forms<br />
<strong>to</strong> 10 cm in height; funnel-shaped forms <strong>to</strong> more than<br />
60 cm in diameter and 30 cm in height. Color in life is<br />
light grey, creamy white, yellow or brown.<br />
Skeletal structure. Spicules are styles (130–280 × 8–19<br />
µm) and palmate isochelae (12–28 µm).<br />
Zoogeographic distribution. Widespread and locally<br />
common. In Alaska – central Aleutian Islands and eastern<br />
Gulf <strong>of</strong> Alaska. Elsewhere – North Pacific Ocean<br />
(Kamchatka Coast, Bering Sea, British Columbia), and<br />
Arctic Ocean (Chukchi Sea and East Siberian Sea).<br />
Habitat. In Alaska – central Aleutian Islands (attached<br />
<strong>to</strong> pebbles at a depth <strong>of</strong> 100 m), eastern Gulf <strong>of</strong> Alaska<br />
(attached <strong>to</strong> bedrock, boulders, and cobbles at depths<br />
between 152 and 218 m). Elsewhere – reported at<br />
depths between 9 and 291 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 100 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ (center). 3)<br />
Specimen collected at a depth <strong>of</strong> 165 m in <strong>the</strong> eastern<br />
Gulf <strong>of</strong> Alaska. Grid marks are 1 cm 2 . 4) Same specimen<br />
as in pho<strong>to</strong> 3 in situ.
86. Esperiopsis flagrum Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This sponge has a flagelliform or cylindrical<br />
growth form; similar <strong>to</strong> Asbes<strong>to</strong>pluma ramosa and<br />
Euchelipluma elongata (differentiation requires careful<br />
examination). Occasionally it is bifurcated. A central<br />
long thin axis tapers near <strong>the</strong> tip and is covered with<br />
many thin processes. Processes are 3–5 mm in length,<br />
elastic but somewhat stiff. The central axis is up <strong>to</strong> 54 cm<br />
in length with a diameter <strong>of</strong> 4 mm at <strong>the</strong> base and 2.5<br />
mm at <strong>the</strong> tip. This cylindrical axis is compressible, elastic<br />
but resilient. Color in life is beige <strong>to</strong> creamy white.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. No special ec<strong>to</strong>some is developed. The<br />
central axis consists <strong>of</strong> interwoven polyspicular tracts <strong>of</strong><br />
styles with many microscleres in between. The lateral<br />
processes are supported by single polyspicular tracts<br />
running in<strong>to</strong> <strong>the</strong>m. The outermost layer <strong>of</strong> <strong>the</strong> processes<br />
and <strong>the</strong> longitudinal axis consists <strong>of</strong> densely arranged<br />
microscleres. Megascleres are fusiform styles (980–1320<br />
× 20–28 µm). Microscleres are large palmate isochelae<br />
(98–112 µm), small palmate isochelae (28–43 µm),<br />
large sigmas (37–48 µm), and small sigmas (17–20 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Occurs at depths between about 700 and<br />
1389 m. Appears <strong>to</strong> bury with root-like processes in s<strong>of</strong>t<br />
unconsolidated sediments.<br />
Remarks. The lower portion <strong>of</strong> <strong>the</strong> stalk is <strong>of</strong>ten fouled<br />
with hydroids. The species occurs within <strong>the</strong> same depth<br />
range as Asbes<strong>to</strong>pluma ramosa, but occurs in s<strong>of</strong>t-sediment<br />
habitats ra<strong>the</strong>r than attaching <strong>to</strong> hard rock. It does not<br />
co-occur with Euchelipluma elongata.<br />
103<br />
Pho<strong>to</strong>s. 1) Mid-section <strong>of</strong> a specimen collected at a<br />
depth <strong>of</strong> 954 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ.
104 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
87. Semisuberites cribrosa (Miklucho-Maclay, 1870)<br />
Description. This stalked sponge has two distinct<br />
forms: 1) several small funnels or cups (up <strong>to</strong> at least<br />
20 cups per individual) that branch initially from a<br />
central stalk, and 2) a long, thin stalk (30 cm or more<br />
in length) that terminates in a single cup-shaped body,<br />
rarely two, with a maximum diameter <strong>of</strong> approximately<br />
25 cm. The stalk is wiry, firm but elastic; <strong>the</strong> body(s) <strong>of</strong><br />
s<strong>of</strong>ter consistency. Surface is smooth <strong>to</strong> <strong>the</strong> unaided eye<br />
but microscopically hispid. Total height and width are<br />
<strong>to</strong> 70 cm (multi-cupped form). Color in life is golden<br />
brown <strong>to</strong> light brown.<br />
Skeletal structure. There is no specialized ec<strong>to</strong>somal<br />
skele<strong>to</strong>n; dermal membrane consists <strong>of</strong> <strong>the</strong> ends <strong>of</strong> ascending<br />
polyspicular tracts. Choanosomal polyspicular<br />
ascending tracts are quite conspicuous and cm-long<br />
fibers are easy drawn from <strong>the</strong> sponge. Tracts are connected<br />
by single spicules, styles in a very wide size range<br />
(75–650 × 6–15 µm).<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – central Aleutian Islands, Bering Sea<br />
(Zhemchug Canyon), and Gulf <strong>of</strong> Alaska. Elsewhere<br />
– widely distributed along <strong>the</strong> north and east coasts<br />
<strong>of</strong> Russia (Bering and Chukchi Seas), Arctic Ocean<br />
(Greenland Sea), and North Atlantic Ocean (Norwegian<br />
Sea).<br />
Habitat. In Alaska – <strong>the</strong> two forms appear <strong>to</strong> be ecomorphs.<br />
The multi-cupped form is attached primarily<br />
<strong>to</strong> bedrock (occasionally cobbles) at depths between<br />
80 and 270 m in low <strong>to</strong> moderate current areas. The<br />
single-cupped form is found in high current areas.<br />
Aleutian Islands – patchy distribution on moderately<br />
sloped sandy habitats; attached <strong>to</strong> pebbles with rootlike<br />
processes at depths between 99 and 306 m. Bering<br />
Sea – rare; pebble and sand slopes at depths around<br />
170 m. Elsewhere – reported at depths between 14 and<br />
325 m, temperatures between 1.9 and 7.0°C, and salinities<br />
between 29.81 and 35.23 psu.<br />
Remarks. This species may be preyed upon by <strong>the</strong><br />
blood star (Henricia sp.) and appears <strong>to</strong> be particularly<br />
fragile.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> a specimen collected at a<br />
depth <strong>of</strong> 100 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ<br />
(center). 3) Specimen collected at a depth <strong>of</strong> 97 m in<br />
<strong>the</strong> central Aleutian Islands. 4) Specimen collected at a<br />
depth <strong>of</strong> 170 m in Zhemchug Canyon, Bering Sea. Note<br />
that <strong>the</strong> specimen has a single large cup that was apparently<br />
two smaller cups now fused <strong>to</strong>ge<strong>the</strong>r. 5) Specimen<br />
collected at a depth <strong>of</strong> 139 m in <strong>the</strong> central Aleutian<br />
Islands. Note that <strong>the</strong> cup has been sliced open for<br />
examination. Grid marks are 1 cm 2 . 6) Same specimen<br />
as in pho<strong>to</strong> 5 in situ (<strong>to</strong>p half). 7) Same specimen as in<br />
pho<strong>to</strong> 5 in situ (bot<strong>to</strong>m half).
87. Semisuberites cribrosa (Miklucho-Maclay, 1870) (continued)<br />
105
106 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
88. Mycale (Aegogropila) adhaerens (Lambe, 1893)<br />
Description. This massive or thickly encrusting sponge<br />
has a bulbous or even fistulose surface. Oscula are<br />
not visible. Consistency is s<strong>of</strong>t and elastic. Size is <strong>to</strong> 20<br />
cm in all dimensions. Color in life is yellow <strong>to</strong> golden<br />
brown.<br />
Skeletal structure. It has ec<strong>to</strong>somal reticulation <strong>of</strong><br />
polyspicular tracts with microscleres in-between tracts.<br />
There is a choanosomal mesh <strong>of</strong> polyspicular tracts, <strong>of</strong>ten<br />
branching. There are tylostyles (350–440 × 5–12 µm),<br />
anisochelae arranged differently in two size classes –<br />
large anisochelae in rosettes (75–100 µm) and small<br />
anisochelae (18–50 µm); sigmata (45–58 µm), and<br />
rhaphides single and in trichodragmata (45–90 µm).<br />
Zoogeographic distribution. Widespread and common.<br />
In Alaska – central Aleutian Islands and Bering<br />
Sea. Elsewhere – North Pacific Ocean (Sea <strong>of</strong> Okhotsk,<br />
Sea <strong>of</strong> Japan, Vancouver Island) and Arctic Ocean<br />
(Greenland Sea).<br />
Habitat. In Alaska – encrusts all hard substrates including<br />
hydrocoral skele<strong>to</strong>ns at depths between 104 and<br />
442 m. Generally found in rough, steep-sloped habitats.<br />
Elsewhere – on rocks, pebbles, and sand bot<strong>to</strong>ms from<br />
<strong>the</strong> intertidal zone <strong>to</strong> a depth <strong>of</strong> 270 m.<br />
Remarks. This species encrusts hydrocorals (Stylaster<br />
sp. and Cyclohelia lamellata) and o<strong>the</strong>r sedentary biota<br />
in Aleutian Island coral gardens; inaccurately reported<br />
in S<strong>to</strong>ne (2006) as Myxilla incrustans.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> a specimen collected at a<br />
depth <strong>of</strong> 190 m in <strong>the</strong> central Aleutian Islands. Specimen<br />
is encrusting <strong>the</strong> hydrocoral Cyclohelia lamellata.<br />
Grid marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong><br />
1 (center) in situ. 3) Specimen collected at a depth <strong>of</strong><br />
109 m in <strong>the</strong> central Aleutian Islands.
89. Mycale (Carmia) carlilei Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This clavate sponge typically has a single<br />
cylinder, occasionally several (up <strong>to</strong> four) smaller<br />
branches, a<strong>to</strong>p a thinner well-defined stalk. One or<br />
more large oscula are a<strong>to</strong>p each cylinder; several smaller<br />
oscula along sides are obvious in situ. Surface is smooth.<br />
Consistency is very s<strong>of</strong>t, except for <strong>the</strong> wiry stalk. Height<br />
is <strong>to</strong> 50 cm or more. Color in life is golden brown.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The stalk consists <strong>of</strong> longitudinally<br />
arranged masses <strong>of</strong> tylostyles; individual tracts are not<br />
recognizable. A special ec<strong>to</strong>some is not developed. The<br />
choanosome consists <strong>of</strong> irregularly arranged pauci- and<br />
polyspicular tracts <strong>of</strong> tylostyles with masses <strong>of</strong> sigmas and<br />
very abundant anisochelae, both single and in rosettes.<br />
Spicules consist <strong>of</strong> tylostyles (470–520 × 10–14 µm), anisochelae<br />
(55–75 µm), and sigmas (65–80 µm).<br />
Zoogeographic distribution. Locally abundant (up<br />
<strong>to</strong> 8 individuals per m 2 ). In Alaska – central Aleutian<br />
Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> bedrock, boulders, cobbles, and<br />
pebbles at depths between 82 and 360 m.<br />
Remarks. Mycale loveni and M. bellabellensis are <strong>the</strong><br />
only o<strong>the</strong>r species in <strong>the</strong> genus that exhibit a stalked<br />
growth form.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 150 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ. 3) Specimen<br />
with several M. loveni at a depth <strong>of</strong> 112 m. 4) Specimen<br />
with several Halichondria oblonga (upper right) at a<br />
depth <strong>of</strong> 119 m.<br />
107
108 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
90. Mycale (Mycale) jasoniae Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This massive sponge has several large<br />
tubes basally connected. In situ, several exhalent canals<br />
flow in<strong>to</strong> large circular oscula surrounded by thin walls.<br />
These oscula collapse and are not readily visible on collected<br />
specimens. The surface is bulbous with many irregularly<br />
distributed conical processes. The consistency<br />
is ra<strong>the</strong>r s<strong>of</strong>t, easily <strong>to</strong>rn, and fibrous. Diameter is <strong>to</strong> 25<br />
cm. Color in life is yellow <strong>to</strong> light yellow.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some is a tangential arrangement<br />
<strong>of</strong> short spicule tracts and single spicules with<br />
many microscleres in between <strong>the</strong> tracts. The choanosome<br />
consists <strong>of</strong> ra<strong>the</strong>r short spicule tracts (60–95 µm<br />
in diameter) that frequently branch <strong>of</strong>f side tracts in<br />
all directions. This pattern is obscured by <strong>the</strong> presence<br />
<strong>of</strong> many single mega- and microscleres in between <strong>the</strong><br />
tracts and without obvious orientation. Megascleres are<br />
tylostyles (405–460 × 10–12 µm). Microscleres include<br />
large anisochelae (80–100 µm), small anisochelae<br />
(40–60 µm), and rhaphides (42–65 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – not reported.<br />
Habitat. Attached <strong>to</strong> cobbles at depths between 178<br />
and 340 m.<br />
Remarks. M. jasoniae is quite similar <strong>to</strong> M. loveni but<br />
possesses rhaphides that <strong>the</strong> latter species lacks.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at<br />
a depth <strong>of</strong> 208 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 in situ<br />
with a sculpin (Malacocottus sp.). 3) Close-up <strong>of</strong> same<br />
specimen as in pho<strong>to</strong>s 1 and 2.
91. Mycale (Mycale) loveni (Fristedt, 1887)<br />
Synonym: Mycale (Carmia) bellabellensis (Lambe, 1905)<br />
Description. This sponge is polymorphic; massively<br />
encrusting or stalked. The consistency is fragile, inelastic<br />
but difficult <strong>to</strong> tear against <strong>the</strong> direction <strong>of</strong> <strong>the</strong> long<br />
polyspicular tracts (laterally); easier <strong>to</strong> tear parallel <strong>to</strong><br />
<strong>the</strong> tracts (<strong>to</strong>p <strong>to</strong> bot<strong>to</strong>m). Aleutian specimens may<br />
be stalked vases <strong>to</strong> 20 cm in height or massive forms<br />
<strong>to</strong> 1 m high and wide. Color in life varies from yellow<br />
<strong>to</strong> greenish yellow <strong>to</strong> brown. Gulf <strong>of</strong> Alaska specimens<br />
have a cone-shaped body with a slender stalk and are<br />
club-shaped until approximately 8 cm high or wide and<br />
<strong>the</strong>n develop <strong>the</strong> characteristic cone shape. Cones may<br />
reach 1 m or more in height and diameter. Color in life<br />
varies from yellow <strong>to</strong> light yellow.<br />
Skeletal structure. Very long ascending polyspicular<br />
tracts (visible <strong>to</strong> <strong>the</strong> unaided eye) lie below a reticulate<br />
dermal skele<strong>to</strong>n. Tracts are branching with thick tracts<br />
connected by shorter tracts. Tylostyles (370–495 × 10–15<br />
µm) <strong>of</strong>ten have <strong>the</strong> largest diameter just before <strong>the</strong><br />
point. Anisochelae are large (80–110 µm) and small<br />
(30–42 µm).<br />
Zoogeographic distribution. Widespread and locally<br />
abundant. In Alaska – Chukchi Sea, Bering Sea,<br />
Aleutian Islands, and Gulf <strong>of</strong> Alaska. Elsewhere – Sea <strong>of</strong><br />
Okhotsk, Pacific Coast <strong>of</strong> <strong>the</strong> Kuril Islands, Chukchi Sea<br />
(Russia), Arctic Ocean (East Siberian Sea), and British<br />
Columbia.<br />
Habitat. Central Aleutian Islands – typically attached<br />
<strong>to</strong> boulders, cobbles, and pebbles at depths between 56<br />
and 744 m (massive form) and 171 <strong>to</strong> 191 m (stalked<br />
form). Bering Sea (Pribil<strong>of</strong> Canyon) – uncommon; attached<br />
<strong>to</strong> cobbles and pebbles at depths between 260<br />
and 309 m. Eastern Gulf <strong>of</strong> Alaska – most specimens are<br />
attached <strong>to</strong> bedrock, boulders, and cobbles at depths<br />
between 143 and 289 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska<br />
109<br />
(and presumably much <strong>deep</strong>er based on unconfirmed<br />
catches with longline gear). Elsewhere – no information<br />
is available.<br />
Remarks. We consider M. loveni and M. bellabellensis <strong>to</strong><br />
be <strong>the</strong> same species, <strong>the</strong> latter being a cone-shaped ecomorph<br />
adapted for low-current habitats in <strong>the</strong> Gulf <strong>of</strong><br />
Alaska. The two species have <strong>the</strong> same spicule complement<br />
and arrangement and internal skeletal structure.<br />
In <strong>the</strong> eastern Gulf <strong>of</strong> Alaska <strong>the</strong> species is club-shaped<br />
until approximately 8 cm wide and <strong>the</strong>n it develops<br />
<strong>the</strong> characteristic cone shape. The sharpchin rockfish<br />
(Sebastes zacentrus) and o<strong>the</strong>r rockfish species <strong>of</strong>ten use<br />
this sponge as perching habitat. This species is preyed<br />
upon by <strong>the</strong> sea star Ceramaster patagonicus.<br />
Pho<strong>to</strong>s. 1) Fragment <strong>of</strong> a specimen (massive form)<br />
collected at a depth <strong>of</strong> 309 m in Pribil<strong>of</strong> Canyon, Bering<br />
Sea. Grid marks are 1 cm 2 . 2) Several specimens (massive<br />
form) at a depth <strong>of</strong> 96 m in <strong>the</strong> central Aleutian<br />
Islands. 3) Several specimens (massive form) with a<br />
sharpchin rockfish (Sebastes zacentrus) at a depth <strong>of</strong> 119<br />
m in <strong>the</strong> central Aleutian Islands. 4) Preserved (frozen)<br />
specimen (stalked form) collected at a depth <strong>of</strong> 192 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
5) Same specimen as in pho<strong>to</strong> 4 in situ. The separation<br />
between <strong>the</strong> red laser marks is 10 cm. 6) Specimen collected<br />
at a depth <strong>of</strong> 167 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska.<br />
Grid marks are 1 cm 2 . 7) Juvenile specimen (with associated<br />
euphasid) collected at a depth <strong>of</strong> 165 m in <strong>the</strong><br />
eastern Gulf <strong>of</strong> Alaska. Grid marks are 1 cm 2 . 8) Same<br />
specimen as in pho<strong>to</strong> 7 in situ. 9) Specimen with a gravid<br />
sharpchin rockfish (S. zacentrus) at a depth <strong>of</strong> 170 m in<br />
<strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Pho<strong>to</strong> by J. Lincoln Freese<br />
(AFSC). 10) A more robust cone-shaped form with a<br />
sharpchin rockfish (S. zacentrus) in a higher current<br />
area at a depth <strong>of</strong> 179 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska.<br />
Note <strong>the</strong> juvenile specimen at <strong>the</strong> upper left. Pho<strong>to</strong> by<br />
J. Lincoln Freese (AFSC).
110 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
91. Mycale (Mycale) loveni (Fristedt, 1887) (continued)
91. Mycale (Mycale) loveni (Fristedt, 1887) (continued)<br />
111
112 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
92. Mycale (Mycale) tylota Koltun, 1958<br />
Description. This sponge is massive with a smooth<br />
but convoluted surface. Circular oscula are on conical<br />
elevations. Consistency is inelastic and very easily <strong>to</strong>rn.<br />
Diameter is <strong>to</strong> 10 cm. Color in life is yellow <strong>to</strong> golden<br />
brown.<br />
Skeletal structure. There is a thick ec<strong>to</strong>somal crust <strong>of</strong><br />
styles <strong>to</strong> tylotes that are scattered without obvious orientation<br />
and tangentially arranged with many microscleres<br />
in between. Choanosomal pauci- <strong>to</strong> polyspicular tracts<br />
are in triangular meshes with masses <strong>of</strong> sigmas in between.<br />
Rosettes <strong>of</strong> large anisochelae are less abundant.<br />
There are ec<strong>to</strong>somal styles <strong>to</strong> tylotes (415–570 × 10–16<br />
µm), choanosomal styles (670–890 × 15–25 µm), large<br />
anisochelae (95–112 µm), small anisochelae (15–35<br />
µm), large sigmas (82–105 µm), small sigmas (25–35<br />
µm), and rhaphides (70–110 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – sou<strong>the</strong>rn<br />
Kuril Strait.<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles and pebbles<br />
at depths between 135 and 175 m. Elsewhere – reported<br />
at depths between 73 and 181 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 172 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 (indicated by <strong>the</strong> white<br />
arrow) in situ.
93. Latrunculia (Biannulata) oparinae Samaii and Krasokhin, 2002<br />
Description. Globular sponge has numerous small oscula<br />
on short, wart-like elevations. Water jets are emitted<br />
through <strong>the</strong> oscula when <strong>the</strong> sponge is squeezed. Areolate<br />
pore-fields collapse upon collection and are only<br />
visible in situ. Consistency is ra<strong>the</strong>r firm, only slightly<br />
elastic and difficult <strong>to</strong> tear. Diameter is <strong>to</strong> at least 16 cm.<br />
Two distinct colors have been observed in situ – dark<br />
brown (dominant) and olive-green.<br />
Skeletal structure. Ec<strong>to</strong>some is thick and lea<strong>the</strong>ry,<br />
consisting <strong>of</strong> a cortex <strong>of</strong> tightly packed anisodiscorhabds.<br />
Fusiform, slightly sinuous styles have acanthose heads<br />
(320–525 × 9–15 µm) and anisodiscorhabds (42–50 µm).<br />
Zoogeographic distribution. Locally common and<br />
abundant (densities up <strong>to</strong> 14 individuals per m 2 in<br />
Alaska). In Alaska – central Aleutian Islands. Elsewhere<br />
– previously known only from <strong>the</strong> Kuril Islands in <strong>the</strong><br />
Sea <strong>of</strong> Okhotsk.<br />
Habitat. Attached <strong>to</strong> bedrock, boulders, cobbles, and<br />
pebbles at depths between 79 and 288 m (possibly <strong>to</strong><br />
113<br />
438 m). Elsewhere – reported at depths between 176<br />
and 202 m.<br />
Remarks. Sponges in <strong>the</strong> genus Latrunculia contain<br />
<strong>the</strong> cy<strong>to</strong><strong>to</strong>xic discorhabdin class <strong>of</strong> pyrroloiminoquinone<br />
alkaloids that exhibit significant antiviral activity<br />
against hepatitis virus C (HCV), antimalarial activity<br />
against Plasmodium falciparum, and antimicrobial effects<br />
against <strong>the</strong> AIDS opportunistic pathogens methicillinresistant<br />
Staphylococcus aureus (MRSA), Mycobacterium<br />
intracellulare, and M. tuberculosis (Na et al., 2010). This<br />
species is associated with several gorgonians, including<br />
Thouarella spp. and Plumarella spp.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 150 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 (right) in situ with <strong>the</strong><br />
olive-green morph (left). 3) Specimen collected at a<br />
depth <strong>of</strong> 146 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 4) Same specimen as in pho<strong>to</strong> 3 (center)<br />
in situ.
114 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
94. Latrunculia velera Lehnert, S<strong>to</strong>ne and Heimler, 2006<br />
Description. This cone-shaped sponge has a flattened<br />
smooth <strong>to</strong>p that is typically circular but occasionally<br />
kidney-shaped. On deck, specimens appear subglobular<br />
because <strong>the</strong>y have been laterally compressed. This<br />
species has a smooth uneven surface, thick and lea<strong>the</strong>ry,<br />
only slightly elastic, and easily <strong>to</strong>rn. The interior is<br />
markedly fibrous, somewhat similar <strong>to</strong> L. oparinae but<br />
differing slightly in shape and clearly in <strong>the</strong> absence<br />
<strong>of</strong> areolate pore-fields and <strong>the</strong> form <strong>of</strong> <strong>the</strong> anisodiscorhabds.<br />
Diameter is <strong>to</strong> about 12 cm. Color in life is<br />
dull brown; dark brown on deck.<br />
Skeletal structure. SEM images <strong>of</strong> spicules are shown<br />
in Appendix IV. The ec<strong>to</strong>some is a unispicular layer <strong>of</strong><br />
discorhabds, all arranged with <strong>the</strong>ir longitudinal axis<br />
perpendicular <strong>to</strong> <strong>the</strong> surface. The choanosome is a<br />
reticulation <strong>of</strong> polyspicular tracts <strong>of</strong> styles with some discorhabds<br />
in between. Megascleres are styles with slightly<br />
acanthose heads (500–540 × 9–11 µm). Microscleres are<br />
relatively smooth anisodiscorhabds (37–43 µm).<br />
Zoogeographic distribution. Locally common. In<br />
Alaska – central Aleutian Islands. Elsewhere – not<br />
reported.<br />
Habitat. Attached <strong>to</strong> bedrock, boulders, and cobbles<br />
at depths between 412 and 1009 m, but relatively rare<br />
at depths shallower than 600 m.<br />
Remarks. There appears <strong>to</strong> be ano<strong>the</strong>r species <strong>of</strong><br />
Latrunculia in <strong>the</strong> central Aleutian Islands at intermediate<br />
depths (i.e., slightly overlapping <strong>the</strong> depth ranges<br />
<strong>of</strong> both L. oparinae and L. velera). Latrunculia specimens<br />
in <strong>the</strong> depth range <strong>of</strong> 200 <strong>to</strong> 500 m should be a priority<br />
for collection.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at<br />
a depth <strong>of</strong> 1009 m in <strong>the</strong> central Aleutian Islands. Note<br />
that <strong>the</strong> specimen has been laterally compressed. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 (far<br />
left) in situ. 3) Specimen observed in situ at a depth <strong>of</strong><br />
929 m. The separation between <strong>the</strong> red laser marks is<br />
10 cm.
95. Latrunculia sp. (undescribed)<br />
Description. This globular sponge has broad cratershaped<br />
pore fields; <strong>the</strong>se are prominent structures<br />
on <strong>the</strong> sponge surface. Surface is uneven and slimy.<br />
Diameter is <strong>to</strong> about 8 cm. Color in life is khaki-green<br />
<strong>to</strong> olive-green.<br />
Skeletal structure. The majority <strong>of</strong> megascleres are<br />
anisostyles, fusiform, not terminally spined, rarely<br />
polytylote, with only slight differentiation between<br />
<strong>the</strong> two ends (325–397 × 7–10 µm). Microscleres are<br />
anisodiscorhabds (49–66 × 4–7 µm). The manubrium,<br />
a base <strong>of</strong> six short, downward pointing smooth spines,<br />
is closely followed by <strong>the</strong> basal whorl, which consists <strong>of</strong><br />
a ring <strong>of</strong> horizontally aligned, smooth spines. The median<br />
whorl (21–31 µm in diameter) is located midway<br />
along <strong>the</strong> shaft and composed <strong>of</strong> undulate petals with<br />
denticulate margins, and some sculpted regions. The<br />
subsidiary whorl is composed <strong>of</strong> similar undulating petals<br />
with denticulate margins and sculpted sections, but<br />
<strong>the</strong> petals slant upwards and are located just underneath<br />
<strong>the</strong> apical whorl. The apical whorl is formed <strong>of</strong> fused<br />
undulating petals with denticulate margins forming a<br />
beautiful corona.<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – continental shelf <strong>of</strong>f Cape Ommaney, Baran<strong>of</strong><br />
Island, eastern Gulf <strong>of</strong> Alaska. Elsewhere – a sponge,<br />
believed <strong>to</strong> be this species, has also been reported from<br />
British Columbia and nor<strong>the</strong>rn Washing<strong>to</strong>n State.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock at depths<br />
between 69 and 210 m, temperatures between 5.5 and<br />
7.6°C, and salinities between 32.2 and 33.7 psu. Elsewhere<br />
– no information available.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 102 m<br />
in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ with <strong>the</strong> hydrocoral<br />
Distichopora borealis (right).<br />
115
116 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
96. Axinella blanca Koltun, 1959<br />
Description. This sponge has irregular branching<br />
and is extremely arborescent. Consistency is moderately<br />
firm and elastic. Surface <strong>of</strong> <strong>the</strong> side branches is<br />
smooth but microscopically hispid. Size is <strong>to</strong> at least<br />
50 cm × 50 cm. Color in life is highly variable ranging<br />
from light brown, light grey, light yellow, golden-yellow<br />
<strong>to</strong> light orange. Light brown is <strong>the</strong> dominant color<br />
in <strong>water</strong>s <strong>deep</strong>er than 120 m in <strong>the</strong> central Aleutian<br />
Islands; golden yellow and light orange are dominant<br />
in shallower <strong>water</strong>.<br />
Skeletal structure. Skeletal architecture consists <strong>of</strong><br />
ascending spicule tracts; spicules protruding from <strong>the</strong>se<br />
tracts cause <strong>the</strong> hispid surface <strong>of</strong> <strong>the</strong> ec<strong>to</strong>some. Spicules<br />
are long oxeas (850–1350 × 18–35 µm) and short styles<br />
(320–530 × 8–15 µm).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – central Aleutian Islands and Bering Sea. Elsewhere<br />
– Bering Sea near Mednyi Island (Commander<br />
Islands, Russia).<br />
Habitat. In Alaska – forms dense fields almost completely<br />
covering <strong>the</strong> seafloor in areas <strong>of</strong> bedrock, boulders,<br />
and cobbles on steep slope habitat and at depths<br />
between 80 and 269 m. Elsewhere – reported at depths<br />
<strong>to</strong> 160 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 140 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ with <strong>the</strong> gorgonian<br />
Plumarella sp. (right). 3) Specimen observed in situ<br />
at a depth <strong>of</strong> 92 m in <strong>the</strong> central Aleutian Islands.
97. Axinella rugosa (Bowerbank, 1866)<br />
Description. This sponge is clavate. Branching, fanshaped,<br />
or single tubes (occasionally two tubes) are <strong>to</strong><br />
17 cm in height. Consistency is firm but elastic, with a<br />
hispid surface. Color in life is grey, light yellow, golden<br />
brown <strong>to</strong> brownish red.<br />
Skeletal structure. Axinellid spicule tracts consist <strong>of</strong><br />
styles, strongyles, and oxeas. All spicules have a large<br />
variation in length up <strong>to</strong> 1750 µm.<br />
Zoogeographic distribution. Widespread and locally<br />
abundant. In Alaska – central Aleutian Islands, Bering<br />
Sea, and eastern Gulf <strong>of</strong> Alaska. Elsewhere – North<br />
Pacific Ocean (Sea <strong>of</strong> Japan), Arctic Ocean (Barents<br />
Sea and Greenland Sea), and North Atlantic Ocean<br />
(Norwegian Sea).<br />
Habitat. In Alaska – attached <strong>to</strong> hard substrate at<br />
depths between 87 and 712 m. Densities <strong>to</strong> 63 individuals<br />
per m 2 in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Elsewhere – reported<br />
at depths between 90 and 320 m, temperatures<br />
between 2 and 5°C, and salinities between 34.20 and<br />
35.01 psu.<br />
Remarks. Though superficially similar <strong>to</strong> Mycale<br />
carlilei, A. rugosa is a hollow tube whereas M. carlilei is a<br />
solid tube with a flat <strong>to</strong>p and one or more prominent<br />
oscula. The Alaskan specimens are <strong>the</strong> first tube forms<br />
reported for <strong>the</strong> species.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 172 m<br />
in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ. 3) Specimen (indicated<br />
by <strong>the</strong> white arrow) collected in situ at a depth<br />
<strong>of</strong> 712 m in <strong>the</strong> central Aleutian Islands.<br />
117
118 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
98. Bubaris vermiculata (Bowerbank, 1866)<br />
Description. This sponge is encrusting, elongated, foliate<br />
or lobate. Consistency is s<strong>of</strong>t with a hispid surface.<br />
Dimensions are <strong>to</strong> 15 cm 2 . Color in life is bright red <strong>to</strong><br />
golden brown.<br />
Skeletal structure. It has characteristic axinellid spicule<br />
tracts. Styles are 426–4550 × 10–38 µm; strongyles<br />
and oxeas are irregularly curved or dis<strong>to</strong>rted (208–588<br />
× 6–15 µm).<br />
Zoogeographic distribution. Widespread and locally<br />
common. In Alaska – central Aleutian Islands. Elsewhere<br />
– Arctic Ocean (Barents and Greenland Seas),<br />
North Atlantic Ocean (Norwegian Sea <strong>to</strong> <strong>the</strong> Azores),<br />
and Mediterranean Sea.<br />
Habitat. In Alaska – encrusts muds<strong>to</strong>ne, cobbles, and<br />
pebbles at depths between 165 and 888 m. Elsewhere –<br />
reported at depths between 9 and 1360 m.<br />
Remarks. This is <strong>the</strong> same species as Axinella vermiculata<br />
sensu Koltun, 1959.<br />
Pho<strong>to</strong>s. 1) Specimen (center) collected with <strong>the</strong><br />
gorgonian Plumarella aleutiana at a depth <strong>of</strong> 165 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Specimen (right) collected on muds<strong>to</strong>ne at a depth<br />
<strong>of</strong> 888 m in <strong>the</strong> central Aleutian Islands.
99. Halichondria (Halichondria) colossea Lundbeck, 1902<br />
Description. This species appears as originally described<br />
and is irregularly cup-shaped and somewhat<br />
compressed. Large osculum is on <strong>the</strong> inside <strong>of</strong> <strong>the</strong> cup.<br />
The surface is smooth on <strong>the</strong> outside and hispid on <strong>the</strong><br />
inside <strong>of</strong> <strong>the</strong> cup. The consistency is firm. Width is <strong>to</strong> 50<br />
cm and height <strong>to</strong> 30 cm. Color in life is greyish brown<br />
<strong>to</strong> creamy white.<br />
Skeletal structure. Smaller oxeas have tangential<br />
arrangement in <strong>the</strong> ec<strong>to</strong>some and larger oxeas are in<br />
characteristic confused “halichondroid” arrangement<br />
in <strong>the</strong> choanosome. Oxeas in two size categories range<br />
in length from 140–2000 µm.<br />
Zoogeographic distribution. Rare. In Alaska – Bering<br />
Sea (Pribil<strong>of</strong> Canyon). Elsewhere – North Atlantic<br />
Ocean (Denmark Strait).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles at a depth <strong>of</strong><br />
300 m. Elsewhere – reported at a depth <strong>of</strong> 1039 m.<br />
Remarks. The specimen collected in Pribil<strong>of</strong> Canyon<br />
is only <strong>the</strong> second record for <strong>the</strong> species worldwide<br />
and <strong>the</strong> first from <strong>the</strong> Pacific Ocean. Several specimens<br />
collected from <strong>the</strong> Atlantic Coast <strong>of</strong> <strong>the</strong> U.S. (Massa-<br />
119<br />
chusetts) have been tentatively identified as H. colossea<br />
(Van Soest 6 ).<br />
Pho<strong>to</strong>. 1) Fragment <strong>of</strong> a specimen collected at a depth<br />
<strong>of</strong> 300 m in Pribil<strong>of</strong> Canyon, Bering Sea.<br />
6 Van Soest, Rob. 2008. Personal commun. Zoological Museum, University<br />
<strong>of</strong> Amsterdam, Amsterdam, The Ne<strong>the</strong>rlands 1090 GT.
120 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
100. Halichondria (Halichondria) oblonga (Hansen, 1885)<br />
Description. This cylindrical, tube-shaped sponge<br />
has a narrow stalk. It may branch near <strong>the</strong> base <strong>of</strong> <strong>the</strong><br />
stalk so that several tubes (up <strong>to</strong> 12) may be connected<br />
basally. Consistency is only slightly elastic and stiff. Near<br />
<strong>the</strong> base <strong>the</strong> tubes are 3–4 mm in diameter and widen<br />
<strong>to</strong> a maximum diameter <strong>of</strong> 2 cm. Maximum height is <strong>to</strong><br />
18 cm. Color in life is golden brown with a light, almost<br />
whitish tip.<br />
Skeletal structure. The brown part <strong>of</strong> <strong>the</strong> sponge<br />
is more stiff and covered with an ec<strong>to</strong>somal layer <strong>of</strong><br />
densely packed, parallel arranged tangential oxeas. The<br />
whitish tip <strong>of</strong> <strong>the</strong> sponge is covered by a thin dermal<br />
membrane without spicules. Choanosomal ascending<br />
paucispicular tracts are connected by single spicules.<br />
Oxeas are 320–480 × 14–20 µm.<br />
Zoogeographic distribution. Widespread and locally<br />
abundant. In Alaska – central Aleutian Islands.<br />
Elsewhere – Arctic Ocean (Barents Sea and Kara Sea)<br />
and North Atlantic Ocean (Greenland and <strong>the</strong> Faroe<br />
Islands).<br />
Habitat. In Alaska – patchy distribution; attached<br />
<strong>to</strong> hard substrate (generally cobbles and pebbles) at<br />
depths between 115 and 305 m. Elsewhere – reported<br />
at depths between 18 and 823 m.<br />
Pho<strong>to</strong>s. 1) Specimens collected with <strong>the</strong> demosponge<br />
Myxilla behringensis (left center) at a depth <strong>of</strong> 195 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimens as in pho<strong>to</strong> 1 in situ. 3) Cluster <strong>of</strong><br />
specimens (lower center) below <strong>the</strong> golden king crab<br />
(Lithodes aequispina) at a depth <strong>of</strong> 272 m in <strong>the</strong> central<br />
Aleutian Islands.
101. Halichondria (Eumastia) sitiens (Schmidt, 1870)<br />
Description. This sponge has a short stalk that typically<br />
branches in<strong>to</strong> several agglutinated tubes. Consistency<br />
is inelastic and stiff due <strong>to</strong> a combination <strong>of</strong> high<br />
spicule density and lack <strong>of</strong> spongin, but easy <strong>to</strong> tear.<br />
Oscula are not obvious. Height is <strong>to</strong> at least 15 cm. Color<br />
in life is light brown, golden brown, or creamy white.<br />
Skeletal structure. Spicules are arranged tangentially<br />
in <strong>the</strong> ec<strong>to</strong>some as is typical for <strong>the</strong> genus. The choanosome<br />
is constructed by polyspicular tracts and many<br />
spicules without orientation in between. Oxeas are <strong>of</strong><br />
a wide size range (145–1200 × 5–20 µm).<br />
Zoogeographic distribution. Cosmopolitan and locally<br />
common. In Alaska – central Aleutian Islands,<br />
Bering Sea (Pribil<strong>of</strong> Canyon), and Arctic Ocean. Elsewhere<br />
– North Pacific Ocean, Arctic Ocean, and North<br />
Atlantic Ocean.<br />
Habitat. In Alaska – attached <strong>to</strong> boulders and cobbles<br />
at depths between 97 and 167 m (central Aleutian<br />
Islands). In <strong>the</strong> Bering Sea it attaches <strong>to</strong> cobbles and<br />
pebbles at depths between 208 and 300 m. Elsewhere –<br />
reported at depths between 6 and 220 m.<br />
Remarks. Koltun (1959) described a cushion-shaped<br />
sponge with a surface <strong>of</strong> numerous elongated papillae<br />
with oscula on <strong>to</strong>p. The agglutinated small tubes <strong>of</strong> <strong>the</strong><br />
Aleutian Island specimens are <strong>the</strong> equivalent <strong>of</strong> <strong>the</strong>se<br />
papillae in a fully grown sponge. The very long oxeas<br />
in halichondroid confused arrangement occur in both<br />
specimens and are regarded as diagnostic.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 167 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 (center) in situ growing<br />
in a cluster <strong>of</strong> <strong>the</strong> same species. 3) Fragmented specimen<br />
collected at a depth <strong>of</strong> 300 m in Pribil<strong>of</strong> Canyon,<br />
Bering Sea.<br />
121
122 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
102. Halichondria sp.<br />
Description. This sponge is extremely polymorphic;<br />
reported growth forms include encrusting, massivelobate,<br />
tubular, lobate, and vase-shaped. Oscula may<br />
be flush with <strong>the</strong> surface, slightly elevated, or on larger<br />
cone-shaped elevations. Consistency is only slightly<br />
elastic but easily fragmented. Size is <strong>to</strong> at least 30 cm ×<br />
30 cm. Color in life ranges from brown, grey, orange,<br />
green, yellow, <strong>to</strong> creamy white.<br />
Skeletal structure. Ec<strong>to</strong>somal spicules are tangentially<br />
arranged. Choanosomal arrangement <strong>of</strong> spicules<br />
is mostly confused but in places may be a unispicular<br />
reticulation or meshes <strong>of</strong> polyspicular tracts. Spicules<br />
are oxeas (220–335 × 15–24 µm).<br />
Zoogeographic distribution. Locally abundant. In<br />
Alaska – Aleutian Islands, Bering Sea (Pribil<strong>of</strong> Canyon),<br />
and eastern Gulf <strong>of</strong> Alaska. Elsewhere – North Pacific<br />
Ocean.<br />
Habitat. In Alaska – attached <strong>to</strong> hard substrate. Depths<br />
between 155 and 208 m (Aleutian Islands); depths between<br />
71 and 255 m (eastern Gulf <strong>of</strong> Alaska); Bering<br />
Sea (Pribil<strong>of</strong> Canyon) – locally abundant; attached <strong>to</strong><br />
bedrock, cobbles, pebbles, and encrusting <strong>the</strong> gorgonian<br />
coral Plumarella echinata at depths between 208 and<br />
309 m. Elsewhere – no information available.<br />
Remarks. This species might represent an undescribed<br />
species. The Alaskan specimens are similar <strong>to</strong><br />
Halichondria panicea (Pallas, 1766) that is common <strong>to</strong><br />
<strong>the</strong> North Atlantic Ocean and was recently introduced<br />
<strong>to</strong> San Francisco Bay. Alaskan specimens have a slightly<br />
different external appearance and <strong>the</strong> spicules are<br />
shorter and more robust than those <strong>of</strong> H. panicea. There<br />
are ongoing discussions about whe<strong>the</strong>r <strong>the</strong>re are several<br />
sibling species or whe<strong>the</strong>r <strong>the</strong> Atlantic and Pacific<br />
Ocean populations are conspecific (Erpenbeck and Van<br />
Soest, 2002). Eggs and hatching larvae similar <strong>to</strong> those<br />
<strong>of</strong> <strong>the</strong> snailfish (Careproctus sp.) were found in flabellate<br />
forms <strong>of</strong> this species in Pribil<strong>of</strong> Canyon, Bering Sea<br />
(Busby 4 ). This species is preyed upon by <strong>the</strong> sea stars<br />
Hippasteria spp., Pteraster tesselatus, Ceramaster patagonicus,<br />
and possibly Henricia longispina.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 175 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 in situ. The specimen<br />
was detached and lying on <strong>the</strong> seafloor in an area that<br />
had been trawled. 3) A fragment <strong>of</strong> a specimen collected<br />
at 152 m in <strong>the</strong> eastern Gulf <strong>of</strong> Alaska. Grid marks<br />
are 1 cm 2 . 4) Same specimen as in pho<strong>to</strong> 3 (lower right)<br />
in situ. Crinoids (Florometra serratissima) use <strong>the</strong> sponge<br />
as an elevated perch. 5) Specimen collected at a depth<br />
<strong>of</strong> 310 m in Pribil<strong>of</strong> Canyon, Bering Sea. This specimen<br />
has completely encrusted <strong>the</strong> gorgonian Plumarella<br />
echinata (tips exposed at right). Grid marks are 1 cm 2 .<br />
6) Same specimen as in pho<strong>to</strong> 5 (indicated by <strong>the</strong> white<br />
arrow) in situ. 7) Specimen collected at a depth <strong>of</strong> 155<br />
m in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
102. Halichondria sp. (continued)<br />
123
124 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
103. Hymeniacidon assimilis Levinsen, 1887<br />
Description. This sponge is polymorphic; massivelobate,<br />
massively cylindrical, ramified, semiglobular, or<br />
even encrusting. Some Aleutian specimens are highly<br />
ramified with some branches coalesced <strong>to</strong>ge<strong>the</strong>r. Rows<br />
<strong>of</strong> small oscula are visible in situ. The consistency is<br />
only slightly elastic, stiff, and easily <strong>to</strong>rn. It appears <strong>to</strong><br />
be particularly fragile. Height <strong>of</strong> highly ramified form<br />
is <strong>to</strong> 1 m or more. Color in life is yellow, light brown,<br />
golden-brown or creamy-white.<br />
Skeletal structure. Ec<strong>to</strong>somal spicules are tangentially<br />
arranged without fur<strong>the</strong>r orientation. There is<br />
choanosomal reticulation <strong>of</strong> polyspicular tracts and<br />
in o<strong>the</strong>r parts short tracts and single spicules without<br />
orientation. Styles (135–560 × 6–23 µm) probably occur<br />
in two size categories.<br />
Zoogeographic distribution. Cosmopolitan and locally<br />
common. In Alaska – central Aleutian Islands,<br />
Bering Sea, and Chukchi Sea. Elsewhere – North Pacific<br />
Ocean (Sea <strong>of</strong> Okhotsk and Sea <strong>of</strong> Japan), Arctic Ocean<br />
(Barents Sea, East Siberian Sea, and Kara Sea), and<br />
North Atlantic Ocean.<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles at depths<br />
between 119 and 253 m; also encrusts <strong>the</strong> gorgonian<br />
Muriceides nigra. Elsewhere – reported at depths between<br />
15 and 110 m.<br />
Pho<strong>to</strong>s. 1) Partial specimen (ramified form) collected<br />
at a depth <strong>of</strong> 119 m in <strong>the</strong> central Aleutian Islands.<br />
Grid marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong><br />
1 (center) in situ. 3) Close-up view <strong>of</strong> same specimen<br />
as in pho<strong>to</strong>s 1 and 2 showing rows <strong>of</strong> small oscula.<br />
4) Fragment <strong>of</strong> specimen (encrusting form) collected<br />
at a depth <strong>of</strong> 160 m in <strong>the</strong> central Aleutian Islands.<br />
Specimen encrusts <strong>the</strong> gorgonian Muriceides nigra.
104. Topsentia disparilis (Lambe, 1893)<br />
Description. On deck this species appears a massive<br />
sponge, but in situ it forms s<strong>to</strong>ut hollow tubes with<br />
spicule tracts protruding far above <strong>the</strong> surface. The<br />
consistency is fragile, inelastic, and easily <strong>to</strong>rn. Height<br />
is <strong>to</strong> 7 cm and width <strong>to</strong> 5 cm. Color in life is light brown<br />
<strong>to</strong> creamy white.<br />
Skeletal structure. Skele<strong>to</strong>n is confused and halichondroid.<br />
Large oxeas are 438–1400 × 13–21 µm;<br />
small oxeas (50–150 × 4–6 µm) are concentrated in <strong>the</strong><br />
ec<strong>to</strong>some.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
central Aleutian Islands, Bering Sea, and Arctic Ocean<br />
(Beaufort Sea – Point Barrow). Elsewhere – Vancouver<br />
Island, British Columbia.<br />
Habitat. In Alaska – attached <strong>to</strong> muds<strong>to</strong>ne on steep<br />
canyon habitat at a depth <strong>of</strong> 2828 m. Grows in small<br />
clusters <strong>of</strong> up <strong>to</strong> eight individuals. Elsewhere – reported<br />
at depths between 80 and 110 m.<br />
Remarks. This species was previously known as<br />
Halichondria disparilis Lambe, 1893. It appears <strong>to</strong> be<br />
extremely eurybathic.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen <strong>the</strong>n s<strong>to</strong>red in ethanol)<br />
specimen collected at a depth <strong>of</strong> 2828 m in <strong>the</strong> central<br />
Aleutian Islands. Grid marks are 1 cm 2 . 2) Same specimen<br />
as in pho<strong>to</strong> 1 (lower left) in situ with a cluster <strong>of</strong><br />
<strong>the</strong> same species.<br />
125
126 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
105. Cladocroce ventilabrum (Fristedt, 1887)<br />
Description. Two growth forms exist: 1) massive <strong>to</strong><br />
hemispherical, and 2) fan-shaped and stalked. The fan<br />
may be thin-bladed or form an ovoid tube with a large<br />
osculum on <strong>to</strong>p. The surface is irregularly corrugated<br />
with crater-like oscula. Both growth forms narrow somewhat<br />
<strong>to</strong> a basal attachment or short stalk. Consistency<br />
is s<strong>of</strong>t, elastic, and easily <strong>to</strong>rn. Size is <strong>to</strong> 10 cm × 10 cm.<br />
Color in life is reddish brown or golden brown.<br />
Skeletal structure. There is an ec<strong>to</strong>somal unispicular<br />
reticulation <strong>of</strong> tangentially arranged oxeas (160–180 ×<br />
5–12 µm). Where oxeas are connected, one <strong>to</strong> four are<br />
arranged perpendicular <strong>to</strong> <strong>the</strong> surface. There is a choanosomal<br />
unispicular reticulation; in places paucispicular<br />
tracts are connected by single spicules.<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – central Aleutian Islands. Elsewhere<br />
– North Pacific Ocean (Sea <strong>of</strong> Japan), Arctic Ocean<br />
(Barents Sea and Greenland Sea), and North Atlantic<br />
Ocean (Davis Strait).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles, pebbles,<br />
and bivalve shells at depths between 155 and 235 m.<br />
Elsewhere – reported at depths between 40 and 718 m.<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 190 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 (center) in situ.<br />
3) Specimen attached <strong>to</strong> a scallop shell (Chlamys sp.)<br />
collected at a depth <strong>of</strong> 155 m in <strong>the</strong> central Aleutian<br />
Islands. Grid marks are 1 cm 2 . 4) Specimen attached<br />
<strong>to</strong> a pebble collected at a depth <strong>of</strong> 155 m in <strong>the</strong> central<br />
Aleutian Islands. Grid marks are 1 cm 2 .
106. Haliclona bucina Tanita and Hoshino, 1989<br />
Description. This small vase-shaped sponge has a<br />
thin fistular stalk or basal fistular s<strong>to</strong>lons. Due <strong>to</strong> <strong>the</strong><br />
reticulation <strong>of</strong> single spicules <strong>the</strong>se <strong>sponges</strong> are only<br />
slightly elastic and consequently very fragile. Height is<br />
<strong>to</strong> 3 cm and width <strong>to</strong> 1 cm. Color in life is light brown<br />
<strong>to</strong> golden brown.<br />
Skeletal structure. There is a unispicular reticulation<br />
<strong>of</strong> oxeas (165–195 × 7–9 µm), slightly longer than those<br />
originally described.<br />
Zoogeographic distribution. Rare. In Alaska – central<br />
Aleutian Islands. Elsewhere – known only from Sagami<br />
Bay, Japan.<br />
Habitat. In Alaska – attached <strong>to</strong> <strong>the</strong> calcareous sponge<br />
Leucandra tuba at a depth <strong>of</strong> 145 m. Elsewhere – depths<br />
between 80 and 95 m.<br />
127<br />
Pho<strong>to</strong>. 1) Specimen (center) attached <strong>to</strong> <strong>the</strong> calcareous<br />
sponge Leucandra tuba collected at a depth <strong>of</strong> 145<br />
m in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
128 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
107. Haliclona (Gellius) digitata (Koltun, 1958)<br />
Description. Koltun (1959) described this species as<br />
“elongated, thinly tabular (foliate), or rolled up and intergrown<br />
in such a manner that it acquires an irregular<br />
shape … hollow dactylate projections….” Aleutian Island<br />
specimens are typically stalked and flabellate. The<br />
surface is smooth. The oscula are circular, chimney-like,<br />
on <strong>to</strong>p <strong>of</strong> oblique tubes. The consistency is only slightly<br />
elastic and easy <strong>to</strong> tear. Size is <strong>to</strong> at least 15 cm in height<br />
and width. Color in life is light brown <strong>to</strong> golden brown;<br />
appears characteristically yellow in situ.<br />
Skeletal structure. The choanosomal skele<strong>to</strong>n consists<br />
<strong>of</strong> long polyspicular tracts connected by single spicules,<br />
oxeas (320–370 × 12–18 µm), and thin sigmas (18–<br />
25 µm).<br />
Zoogeographic distribution. Uncommon. In Alaska<br />
– central Aleutian Islands. Elsewhere – North Pacific<br />
Ocean (Sea <strong>of</strong> Okhotsk near <strong>the</strong> Pacific coast <strong>of</strong> <strong>the</strong><br />
sou<strong>the</strong>rn Kuril Islands).<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, boulders,<br />
cobbles, and pebbles at depths between 96 and<br />
258 m. Elsewhere – found on sand and gravel at depths<br />
between 285 and 287 m and at a temperature <strong>of</strong> 1.7°C.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
108. Haliclona (Gellius) primitiva (Lundbeck, 1902)<br />
Description. According <strong>to</strong> Koltun (1959) this species<br />
is massive-lobate, or cushion-shaped, up <strong>to</strong> 10.5 cm in<br />
height, and <strong>of</strong>ten forms dactylate, lobate or o<strong>the</strong>r types<br />
<strong>of</strong> projections. Aleutian Island specimens are dactylate<br />
with hollow centers; up <strong>to</strong> three fingers are basally attached.<br />
Surface is corrugated. Color in life is light yellow,<br />
brown, or golden brown.<br />
Skeletal structure. The ec<strong>to</strong>somal skele<strong>to</strong>n is a tangential,<br />
unispicular reticulation <strong>of</strong> single spicules.<br />
Conspicuous polyspicular tracts, several mm in length,<br />
are connected by a reticulation <strong>of</strong> single spicules in <strong>the</strong><br />
choansosome. Oxeas are 140–175 × 8–13 µm and <strong>to</strong>xa<br />
are 45–135 µm.<br />
Zoogeographic distribution. Widespread but uncommon.<br />
In Alaska – central Aleutian Islands and Bering<br />
Sea. Elsewhere – North Pacific Ocean (Sea <strong>of</strong> Okhotsk,<br />
Sea <strong>of</strong> Japan, Pacific coast <strong>of</strong> <strong>the</strong> Kuril Islands), Arctic<br />
Ocean (White Sea), and North Atlantic Ocean (west <strong>of</strong><br />
Greenland).<br />
Habitat. In Alaska – attached <strong>to</strong> pebbles at a depth<br />
<strong>of</strong> 138 m. Elsewhere – found on sand, rock, and ooze<br />
at depths between 27 and 200 m, at temperatures between<br />
0.1 and 3.4°C, and at salinities between 27.0 and<br />
33.9 psu.<br />
Pho<strong>to</strong>s. 1) Partial specimen collected at a depth <strong>of</strong> 138<br />
m in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Same specimen as in pho<strong>to</strong> 1 (indicated by <strong>the</strong> white<br />
arrows) in situ covered with ophiuroids.<br />
129
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109. Haliclona tenuiderma (Lundbeck, 1902)<br />
Description. This sponge is cushion-shaped with a<br />
finely hispid surface. Consistency is very delicate; only<br />
slightly elastic and easily <strong>to</strong>rn. Circular oscula are slightly<br />
elevated from <strong>the</strong> surface and not visible on collected<br />
specimens. Height is <strong>to</strong> 15 cm and width <strong>to</strong> 8 cm. Color<br />
in life is light grey <strong>to</strong> creamy white.<br />
Skeletal structure. Choanosomal paucispicular tracts<br />
are connected by single spicules and short tracts; oxeas<br />
(330–430 × 13–15 µm).<br />
Zoogeographic range. Widespread but rare. In Alaska<br />
– central Aleutian Islands. Elsewhere – North Pacific<br />
Ocean (Sea <strong>of</strong> Okhotsk and Sea <strong>of</strong> Japan), Arctic Ocean<br />
(Barents Sea and Greenland Sea), and North Atlantic<br />
Ocean.<br />
Habitat. In Alaska – attached <strong>to</strong> bedrock, boulders,<br />
and muds<strong>to</strong>ne at depths between 974 and 1706 m.<br />
Elsewhere – reported at depths between 0 and 15 m<br />
(Barents Sea) and at a depth <strong>of</strong> 887 m (North Atlantic<br />
Ocean).<br />
Pho<strong>to</strong>s. 1) Fragments <strong>of</strong> preserved (frozen <strong>the</strong>n ethanol)<br />
specimen collected at a depth <strong>of</strong> 1352 m in <strong>the</strong><br />
central Aleutian Islands. Grid marks are 1 cm 2 . 2) Same<br />
specimen as in pho<strong>to</strong> 1 in situ. 3) Close-up view <strong>of</strong> same<br />
specimen as in pho<strong>to</strong> 2 and 3.
110. Haliclona (Haliclona) urceolus (Rathke and Vahl, 1806)<br />
Description. This sponge is stalked and oviform or<br />
stalked tubes (de Weerdt, 1986a). Aleutian Island specimens<br />
are stalked tubes; stalk approximately 1 cm long,<br />
overall length <strong>to</strong> about 10 cm. Color in life is light yellow<br />
with a darker stalk.<br />
Skeletal structure. There is a unispicular reticulation<br />
<strong>of</strong> oxeas (178–280 µm).<br />
Zoogeographic range: Uncommon. In Alaska –<br />
central Aleutian Islands (first record in <strong>the</strong> North Pacific<br />
Ocean). Elsewhere – Arctic Ocean (Kara Sea) and<br />
North Atlantic Ocean (North Sea).<br />
Habitat. In Alaska – attached <strong>to</strong> cobbles at depths near<br />
490 m. Elsewhere – found on sediment covered s<strong>to</strong>nes<br />
in sheltered habitats at depths between 5 and l000 m.<br />
Pho<strong>to</strong>s. 1) Preserved (frozen) specimen collected at<br />
a depth <strong>of</strong> 490 m in <strong>the</strong> central Aleutian Islands. Grid<br />
marks are 1 cm 2 . 2) Same specimen as in pho<strong>to</strong> 1 (right)<br />
in situ.<br />
131
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111. Haliclona sp. 1<br />
Description. This sponge is massively encrusting or<br />
cushion-shaped. Oscula are <strong>deep</strong> and crater-shaped.<br />
Surface is slightly rough; consistency is s<strong>of</strong>t and fragile.<br />
Diameter is <strong>to</strong> about 10 cm. Color in life is light brown<br />
or golden brown.<br />
Skeletal structure. There is a choanosomal arrangement<br />
<strong>of</strong> oxeas (160–220 × 10–15 µm) in paucispicular<br />
tracts connected by single spicules.<br />
Zoogeographic distribution. Uncommon. In Alaska –<br />
central Aleutian Islands and Bering Sea.<br />
Habitat. Attached <strong>to</strong> cobbles and pebbles at depths<br />
between 150 and 160 m.<br />
Remarks. This species might represent an undescribed<br />
species.<br />
Pho<strong>to</strong>. 1) Specimen collected at a depth <strong>of</strong> 155 m in<br />
<strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .
112. Haliclona sp. 2<br />
Description. This sponge is flabellate; single- or multiplaned.<br />
Oscula are conspicuous, slightly elevated and<br />
scattered over <strong>the</strong> entire surface. It is extremely slimy,<br />
even in preservative. Consistency is very s<strong>of</strong>t and elastic.<br />
Height and width are <strong>to</strong> 20 cm. Color in life is yellowish<br />
brown <strong>to</strong> light brown, but appears characteristically<br />
white in situ.<br />
Skeletal structure. Ec<strong>to</strong>somal tangential unispicular<br />
reticulation is streng<strong>the</strong>ned by large quantities <strong>of</strong><br />
spongin which covers <strong>the</strong> sponge like a perforated<br />
plate. Choanosomal paucispicular tracts are connected<br />
by single spicules <strong>to</strong> unispicular reticulation <strong>of</strong> oxeas<br />
(180–230 × 5–12 µm).<br />
Zoogeographic distribution. In Alaska – locally abundant<br />
in central Aleutian Islands.<br />
Habitat. Attached principally <strong>to</strong> bedrock, but occasionally<br />
<strong>to</strong> boulders and cobbles at depths between 74<br />
and 195 m; more common at depths shallower than<br />
120 m.<br />
Remarks. This species might represent an undescribed<br />
species. It is similar in growth form <strong>to</strong> Haliclona<br />
cinerea and displays <strong>the</strong> slime strands previously regarded<br />
as diagnostic for <strong>the</strong> species. However, H. cinerea has<br />
smaller oxeas than this species and is known only from<br />
shallow <strong>water</strong>s <strong>of</strong> <strong>the</strong> Celtic Seas Region <strong>of</strong> <strong>the</strong> North<br />
Atlantic Ocean. This species may be preyed upon by <strong>the</strong><br />
blood star (Henricia sp.).<br />
Pho<strong>to</strong>s. 1) Specimen collected at a depth <strong>of</strong> 80 m<br />
in <strong>the</strong> central Aleutian Islands. Grid marks are 1 cm 2 .<br />
2) Specimen (right) observed at 92 m in <strong>the</strong> central<br />
Aleutian Islands.<br />
133
134 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Acknowledgments<br />
We thank E. Brown, D. Carlile, S. France, L. Watling,<br />
S. Rooney, P. Malecha, T. Marshall, K. Lowyck, J. Hocevar,<br />
M. Ridgway, A. Andrews, and J. Mondragon for<br />
assistance with <strong>the</strong> collection <strong>of</strong> specimens. We thank<br />
John Hocevar <strong>of</strong> Greenpeace International for allowing<br />
us <strong>to</strong> include in this <strong>guide</strong> specimens collected during<br />
<strong>the</strong>ir 2007 Bering Sea Canyons Expedition. T. Warshaw<br />
provided <strong>the</strong> <strong>to</strong>pside pho<strong>to</strong>graphs <strong>of</strong> <strong>the</strong> Bering Sea<br />
specimens; B. Masuda helped with much <strong>of</strong> <strong>the</strong> pho<strong>to</strong><br />
editing. M. Kelly (NIWA) provided <strong>the</strong> skeletal structure<br />
description for <strong>the</strong> undescribed species <strong>of</strong> Latrunculia.<br />
The field collections were supported by <strong>the</strong> Alaska Fisheries<br />
Science Center <strong>of</strong> <strong>the</strong> National Marine Fisheries<br />
Service, West Coast and Polar Regions Undersea Research<br />
Center <strong>of</strong> NOAA’s National Undersea Research<br />
Program, and <strong>the</strong> North Pacific Research Board. We<br />
thank three anonymous reviewers for helpful comments<br />
on an earlier version <strong>of</strong> this manuscript. Funding for <strong>the</strong><br />
completion <strong>of</strong> this <strong>guide</strong> was provided by <strong>the</strong> National<br />
Marine Fisheries Service, Office <strong>of</strong> Habitat Conservation,<br />
as part <strong>of</strong> <strong>the</strong> Deep Sea Coral Research and Technology<br />
Program.<br />
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Publishing, Madeira Park, BC, 398 p.<br />
Lambe, L. M.<br />
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Alaska. The Ottawa Naturalist, Volume XIV (9):152–172.<br />
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Assoc. U.K. 85:1359–1366.<br />
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Appendix I. Sponge species reported from Alaskan <strong>water</strong>s<br />
This list includes <strong>the</strong> <strong>sponges</strong> currently known <strong>to</strong><br />
occur in <strong>the</strong> <strong>deep</strong> <strong>water</strong>s (>80 m) <strong>of</strong> Alaska. The list<br />
has been drawn from <strong>the</strong> published literature (mostly<br />
species descriptions) and from recent collections <strong>of</strong><br />
specimens that have been definitively identified by us<br />
through examination <strong>of</strong> microscopic characters. We<br />
provide a complete list <strong>of</strong> all species reported from<br />
<strong>the</strong> area, but only provide detailed species descriptions<br />
in this <strong>guide</strong> for those species that we collected<br />
PHYLUM PORIFERA<br />
CLASS CALCAREA<br />
Order Baerida<br />
Family Baeriidae<br />
Leuconia alaskensis de Laubenfels, 1953; (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Leucopsila cf. stylifera (Borojevic, Boury-Esnault and Vacelet, 2000); (Aleutian Islands)<br />
Order Clathrinida<br />
Family Clathrinidae<br />
Clathrina sp. ........................................................................Page 12<br />
Order Leucosolenida<br />
Family Amphoriscidae<br />
Leucilla nuttingi (Urban, 1902); (Aleutian Islands)<br />
Family Grantiidae<br />
Leucandra ananas (Montagu, 1818); (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Leucandra heathi Urban, 1906; (Aleutian Islands)<br />
Leucandra poculiformis Hozawa, 1918 .....................................................Page 13<br />
Leucandra pyriformis (Lambe, 1893); (Aleutian Islands)<br />
Leucandra taylori Lambe, 1900; (Aleutian Islands)<br />
Leucandra tuba Hozawa, 1918 ...........................................................Page 14<br />
Family Leucosoleniidae<br />
Leucosolenia eleanor Urban, 1905; (Aleutian Islands, Gulf <strong>of</strong> Alaska)<br />
Family Sycettidae<br />
Sycon compactum Lambe, 1893; (Aleutian Islands)<br />
CLASS HEXACTINELLIDA<br />
Subclass AMPHIDISCOPHORA<br />
Order Amphidiscosida<br />
Family Hyalonematidae<br />
Hyalonema (Cyliconema) apertum apertum Schulze, 1886; (western Aleutian Islands)<br />
Hyalonema (Cyliconema) apertum simplex Koltun, 1967; (Bering Sea)<br />
Hyalonema (Cyliconema) hozawai vicarium Koltun, 1967; (Bering Sea)*<br />
Hyalonema (Lep<strong>to</strong>nema) ovuliferum Schulze, 1899; (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Hyalonema (Cyliconema) tenerum vitiazi Koltun, 1967; (Bering Sea)*<br />
Subclass HEXASTEROPHORA<br />
Order Hexactinosida<br />
Family Farreidae<br />
Farrea kurilensis ssp. nov. Reiswig and S<strong>to</strong>ne, in preparation ..................................Page 16<br />
Farrea occa occa Bowerbank, 1862 ........................................................Page 17<br />
Farrea watasei Okada, 1932; (Bering Sea)*<br />
Farrea sp. Okada, 1932; (Bering Sea)<br />
Farrea sp. nov. Reiswig and S<strong>to</strong>ne, in preparation ..........................................Page 18<br />
137<br />
and examined ourselves. For all o<strong>the</strong>r species we provide<br />
<strong>the</strong> zoogeographic ranges, but refer <strong>the</strong> reader <strong>to</strong><br />
<strong>the</strong> primary literature for those species descriptions.<br />
The taxonomic names are all modern, systematically<br />
arranged <strong>to</strong> family, and alphabetically arranged by<br />
genus and species within family. (*) indicates that <strong>the</strong><br />
species was reported without precise coordinates but<br />
we believe that <strong>the</strong> collection was made in Alaskan<br />
<strong>water</strong>s.
138 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Family Euretidae<br />
Chonelasma sp. Schulze, 1899; (nor<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Eurete irregularis Bowerbank, 1876; (Bering Sea)*<br />
Genus nov., sp. nov. Reiswig and S<strong>to</strong>ne, in preparation .....................................Page 19<br />
Family Tre<strong>to</strong>dictyidae<br />
Tre<strong>to</strong>dictyum sp. nov. Reiswig and S<strong>to</strong>ne, in preparation ......................................Page 20<br />
Family Aphrocallistidae<br />
Aphrocallistes vastus Schulze, 1886 .......................................................Page 21<br />
Heterochone aleutiana (Okada, 1932); (western Aleutian Islands)<br />
Heterochone calyx calyx Schulze, 1886 .....................................................Page 23<br />
Heterochone calyx schulzei Koltun, 1967; (Bering Sea)<br />
Heterochone tenera Schulze, 1899; (Bering Sea)*<br />
Heterochone n. sp. A Schuchert and Reiswig, 2006; (Gulf <strong>of</strong> Alaska)<br />
Order Lyssacinosida<br />
Family Euplectellidae<br />
Euplectella oweni Herklots and Marshall, 1868; (Bering Sea)*<br />
Euplectella sp. Koltun, 1967; (Bering Sea)*<br />
Holascus undulatus Schulze, 1899 (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Regadrella okinoseana Ijima, 1896 ........................................................Page 25<br />
Family Rossellidae<br />
Acanthascus (Acanthascus) alani pr<strong>of</strong>undum Koltun, 1967; (Bering Sea)*<br />
Acanthascus (Rhabdocalyptus) australis Topsent, 1901; (Bering Sea)<br />
Acanthascus (Rhabdocalyptus) borealis Okada, 1932; (Bering Sea)<br />
Acanthascus (Rhabdocalyptus) dawsoni alascensis Wilson and Penney, 1930 (Bering Sea)<br />
Acanthascus (Rhabdocalyptus) dawsoni dawsoni (Lambe, 1893) .................................Page 27<br />
Acanthascus (Rhabdocalyptus) dawsoni horridus Koltun, 1967; (Bering Sea)*<br />
Acanthascus (Staurocalyptus) dowlingi (Lambe, 1894); (Gulf <strong>of</strong> Alaska)<br />
Acanthascus (Rhabdocalyptus) heteraster Okada, 1932; (Bering Sea)<br />
Acanthascus (Rhabdocalyptus) mirabilis (Schulze, 1899) ......................................Page 28<br />
Acanthascus (Acanthascus) mitis Koltun, 1967; (Bering Sea)*<br />
Acanthascus (Acanthascus) pr<strong>of</strong>undum ssp. nov. Reiswig and S<strong>to</strong>ne, in preparation .................Page 26<br />
Acanthascus (Staurocalyptus) rugocruciatus Okada, 1932; (Bering Sea)<br />
Acanthascus (Staurocalyptus) solidus (Schulze, 1899) .........................................Page 29<br />
Acanthascus (Rhabdocalyptus) unguiculatus (Ijima, 1904); (western Aleutian Islands)<br />
Acanthascus (Staurocalyptus) sp. nov. 1 Reiswig and S<strong>to</strong>ne, in preparation .......................Page 30<br />
Acanthascus (Staurocalyptus) sp. nov. 2 Reiswig and S<strong>to</strong>ne, in preparation .......................Page 31<br />
Aulosaccus fissuratus Okada, 1932; (Bering Sea, western Aleutian Islands)<br />
Aulosaccus ijimai (Schulze, 1899); (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Aulosaccus pinularis Okada, 1932 ........................................................Page 32<br />
Aulosaccus schulzei Ijima, 1896 ..........................................................Page 33<br />
Bathydorus echinus Koltun, 1967; (Bering Sea)*<br />
Bathydorus laevis spinosus Wilson, 1904; (Bering Sea)*<br />
Bathydorus sp. Okada, 1932; (Bering Sea)<br />
Bathydorus sp. Reiswig and S<strong>to</strong>ne, in preparation ...........................................Page 34<br />
Caulophacus (Caulophacus) elegans Schulze, 1885; (Bering Sea)*<br />
Caulophacus (Caulophacus) schulzei hyperboreus Koltun, 1967; (Bering Sea)*<br />
Caulophacus (Caulophacus) sp. nov. Reiswig and S<strong>to</strong>ne, in preparation ..........................Page 35<br />
Scyphidium tuberculata (Okada, 1932); (Bering Sea)<br />
CLASS DEMOSPONGIAE<br />
Order Dendroceratida<br />
Family Darwinellidae<br />
Aplysilla glacialis (Merejkowski, 1877); (Arctic Ocean)
Order Homosclerophorida<br />
Family Plakinidae<br />
Plakina atka Lehnert, S<strong>to</strong>ne and Heimler, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 38<br />
Plakina tanaga Lehnert, S<strong>to</strong>ne and Heimler, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 39<br />
Order Spirophorida<br />
Family Tetillidae<br />
Craniella arb (de Laubenfels, 1930) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 40<br />
Craniella craniana de Laubenfels, 1953; (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Craniella sigmoancoratum (Koltun, 1966) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 41<br />
Craniella spinosa Lambe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 42<br />
Craniella sputnika Lehnert, S<strong>to</strong>ne and Heimler, 2011 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 43<br />
Craniella villosa Lambe, 1893; (Aleutian Islands, Gulf <strong>of</strong> Alaska)<br />
Order Astrophorida<br />
Family Ancorinidae<br />
Penares cortius de Laubenfels, 1930; (Gulf <strong>of</strong> Alaska)<br />
Stelletta validissima Thiele, 1898; (Bering Sea)<br />
Family Geodiidae<br />
Erylus aleuticus Lehnert, S<strong>to</strong>ne and Heimler, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 44<br />
Geodia lendenfeldi nomen novum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 45<br />
Geodia mesotriaena Lendenfeld, 1910; (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Family Pachastrellidae<br />
Poecillastra japonica (Thiele, 1898); (Bering Sea)<br />
Poecillastra tenuilaminaris (Sollas, 1886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 46<br />
Order Hadromerida<br />
Family Polymastiidae<br />
Polymastia andrica de Laubenfels, 1949; (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Polymastia fluegeli Lehnert, S<strong>to</strong>ne and Heimler, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 47<br />
Polymastia kurilensis Koltun, 1962; (Gulf <strong>of</strong> Alaska, Bering Sea)<br />
Polymastia pacifica Lambe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 48<br />
Radiella sol Schmidt, 1870; (eastern Gulf <strong>of</strong> Alaska)<br />
Family Stylocordylidae<br />
Stylocordyla borealis eous Koltun, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 49<br />
Family Suberitidae<br />
Aap<strong>to</strong>s kanuux Lehnert, Hocevar and S<strong>to</strong>ne 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 50<br />
Pseudosuberites montiniger (Carter, 1880); (eastern Gulf <strong>of</strong> Alaska – Cross Sound)<br />
Rhizaxinella clavata (Thiele, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 51<br />
Suberites concinnus Lambe, 1895; (Gulf <strong>of</strong> Alaska, Bering Sea, Arctic Ocean)<br />
Suberites excellens (Thiele, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 52<br />
Suberites montalbidus Carter, 1880; (eastern Aleutian Islands)<br />
Suberites simplex Lambe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 53<br />
Suberites suberia (Montagu, 1818); (Gulf <strong>of</strong> Alaska, Bering Sea)<br />
Suberites virgul<strong>to</strong>sus (Johns<strong>to</strong>n, 1842); (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Suberites sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 54<br />
Order Haplosclerida (Suborder Petrosina)<br />
Family Petrosiidae<br />
Petrosia (Petrosia) borealis (Lambe, 1895); (western Aleutian Islands)<br />
Order Haplosclerida (Suborder Haplosclerina)<br />
Family Niphatidae<br />
Hemigellius porosus (Fristedt, 1887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 55<br />
Order Poecilosclerida (Suborder Microcionina)<br />
Family Acarnidae<br />
Cornulum clathriata (Koltun, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 56<br />
Iophon piceum (Vosmaer, 1882) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Page 57<br />
139
140 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Family Acarnidae (continued)<br />
Iophon piceum abipocillus Koltun, 1959 ....................................................Page 58<br />
Megaciella anisochela Lehnert, S<strong>to</strong>ne and Heimler, 2006 .....................................Page 59<br />
Megaciella spirinae (Koltun, 1958) .......................................................Page 60<br />
Wigginsia wigginsi de Laubenfels, 1953; (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Family Microcionidae<br />
Clathria (Clathria) barleei (Bowerbank, 1866) ..............................................Page 61<br />
Clathria (Clathria) laevigata Lambe, 1893 .................................................Page 62<br />
Clathria (Axosuberites) lambei (Koltun, 1955) ..............................................Page 63<br />
Echinoclathria beringensis (Hentschel, 1929); (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Echinoclathria vasa Lehnert, S<strong>to</strong>ne and Heimler, 2006 .......................................Page 64<br />
Artemisina amlia Lehnert, S<strong>to</strong>ne and Heimler, 2006 ........................................Page 65<br />
Artemisina arcigera (Schmidt, 1870) .....................................................Page 66<br />
Artemisina stipitata Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 67<br />
Artemisina sp. .......................................................................Page 68<br />
Order Poecilosclerida (Suborder Myxillina)<br />
Family Coelosphaeridae<br />
Coelosphaera oglalai Lehnert, S<strong>to</strong>ne and Heimler, 2006 ......................................Page 69<br />
Inflatella globosa (Koltun, 1955) ........................................................Page 70<br />
Lissodendoryx (Lissodendoryx) amaknakensis (Lambe, 1895); (Gulf <strong>of</strong> Alaska, Bering Sea)<br />
Lissodendoryx (Lissodendoryx) behringi Koltun, 1958 .........................................Page 71<br />
Lissodendoryx (Lissodendoryx) firma (Lambe, 1895); (western Aleutian Islands)<br />
Lissodendoryx (Ectyodoryx) olgae (Hentschel, 1929) ..........................................Page 72<br />
Lissodendoryx (Lissodendoryx) oxeota Koltun, 1958 ...........................................Page 73<br />
Lissodendoryx (Lissodendoryx) papillosa Koltun, 1958 .........................................Page 74<br />
Family Crambeidae<br />
Monanchora alaskensis (Lambe, 1895) ....................................................Page 75<br />
Monanchora laminachela Lehnert, S<strong>to</strong>ne and Heimler, 2006 ..................................Page 76<br />
Monanchora pulchra (Lambe, 1894) .....................................................Page 77<br />
Family Crellidae<br />
Crella brunnea (Hansen, 1885) .........................................................Page 78<br />
Family Hymedesmiidae<br />
Hymedesmia (Stylopus) dermata Lundbeck, 1910 ............................................Page 79<br />
Hymedesmia (Hymedesmia) irregularis Lundbeck, 1910 .......................................Page 80<br />
Hymedesmia (Stylopus) longurius Lundbeck, 1910; (Aleutian Islands)<br />
Kirkpatrickia borealis Koltun, 1970 .......................................................Page 81<br />
Phorbas paucistylifer Koltun, 1958 ........................................................Page 82<br />
Family Iotrochotidae<br />
Iotroata magna (Lambe, 1900); (western Aleutian Islands)<br />
Family Myxillidae<br />
Melonanchora globogilva Lehnert, S<strong>to</strong>ne and Heimler, 2006 ...................................Page 83<br />
Myxilla (Myxilla) barentsi Vosmaer, 1885; (Gulf <strong>of</strong> Alaska, Bering Sea, Arctic Ocean)<br />
Myxilla (Myxilla) behringensis Lambe, 1895 ................................................Page 84<br />
Myxilla (Myxilla) incrustans (Johns<strong>to</strong>n, 1842); (Arctic Ocean – Point Barrow, Beaufort Sea;<br />
Gulf <strong>of</strong> Alaska)<br />
Myxilla (Bur<strong>to</strong>nanchora) lacunosa Lambe, 1893; (Aleutian Islands, Gulf <strong>of</strong> Alaska)<br />
Myxilla (Ectyomyxilla) parasitica Lambe, 1893 ..............................................Page 85<br />
Myxilla (Bur<strong>to</strong>nanchora) pedunculata Lundbeck, 1905 .......................................Page 86<br />
Stelodoryx oxeata Lehnert, S<strong>to</strong>ne and Heimler, 2006 .........................................Page 87<br />
Stelodoryx <strong>to</strong>poroki Koltun, 1958 ..........................................................Page 88<br />
Stelodoryx vitiazi (Koltun, 1959) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 89<br />
Family Phellodermidae<br />
Echinostylinos hirsutus Koltun, 1970 ......................................................Page 90<br />
Family Tedaniidae<br />
Tedania (Tedania) dirhaphis Hentschel, 1912 ..............................................Page 91
Family Tedaniidae (continued)<br />
Tedania (Tedania) fragilis Baer, 1906; (eastern Aleutian Islands)<br />
Tedania kagalaskai Lehnert, S<strong>to</strong>ne and Heimler, 2006 ...................................... Page 92<br />
Order Poecilosclerida (Suborder Mycalina)<br />
Family Cladorhizidae<br />
Abyssocladia bruuni Lévi, 1964; (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Asbes<strong>to</strong>pluma occidentalis (Lambe, 1893); (Bering Sea)<br />
Asbes<strong>to</strong>pluma ramosa Koltun, 1958 ...................................................... Page 93<br />
Cladorhiza bathycrinoides Koltun, 1955 ................................................... Page 95<br />
Cladorhiza corona Lehnert, Watling and S<strong>to</strong>ne, 2005 ....................................... Page 96<br />
Cladorhiza longipinna Ridley and Dendy, 1886 (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Chondrocladia (Chondrocladia) concrescens (Schmidt, 1880) .................................. Page 97<br />
Family Desmacellidae<br />
Biemna rhadia de Laubenfels, 1930; (sou<strong>the</strong>rn Gulf <strong>of</strong> Alaska)<br />
Biemna variantia (Bowerbank, 1858) ................................................... Page 98<br />
Family Guitarridae<br />
Euchelipluma elongata Lehnert, S<strong>to</strong>ne and Heimler, 2006 ................................... Page 99<br />
Guitarra abbotti Lee, 1987 ............................................................. Page 100<br />
Guitarra fimbriata Carter, 1874 ......................................................... Page 101<br />
Family Esperiopsidae<br />
Amphilectus digitatus (Miklucho-Maclay, 1870) ............................................ Page 102<br />
Amphilectus lobatus (Montagu, 1818); (Alaska)<br />
Esperiopsis flagrum Lehnert, S<strong>to</strong>ne and Heimler, 2006 ...................................... Page 103<br />
Semisuberites cribrosa (Miklucho-Maclay, 1870) ............................................ Page 104<br />
Family Mycalidae<br />
Mycale (Aegogropila) adhaerens (Lambe, 1893) ............................................ Page 106<br />
Mycale (Carmia) carlilei Lehnert, S<strong>to</strong>ne and Heimler, 2006 .................................. Page 107<br />
Mycale (Carmia) helios (Fristedt, 1887); (Bering Sea, Arctic Ocean)<br />
Mycale (Mycale) hispida (Lambe, 1893); (Gulf <strong>of</strong> Alaska, Aleutian Islands)<br />
Mycale (Mycale) jasoniae Lehnert, S<strong>to</strong>ne and Heimler, 2006 ................................. Page 108<br />
Mycale (Mycale) lingua (Bowerbank, 1866); (central Aleutian Islands)<br />
Mycale (Mycale) loveni (Fristedt, 1887) .................................................. Page 109<br />
Mycale (Mycale) modesta (Lambe, 1894); (Gulf <strong>of</strong> Alaska, Bering Sea)<br />
Mycale (Mycale) <strong>to</strong>poroki Koltun, 1958; (Bering Sea)<br />
Mycale (Mycale) tylota Koltun, 1958 ..................................................... Page 112<br />
Family Isodictyidae<br />
Isodictya quatsinoensis (Lambe, 1892); (Gulf <strong>of</strong> Alaska, Bering Sea)<br />
Order Poecilosclerida (Suborder Latrunculia)<br />
Family Latrunculiidae<br />
Latrunculia occulta Lehnert, S<strong>to</strong>ne and Heimler, 2006; (central Aleutian Islands)<br />
Latrunculia (Biannulata) oparinae Samaii and Krasokhin, 2002 ............................... Page 113<br />
Latrunculia velera Lehnert, S<strong>to</strong>ne and Heimler, 2006 ...................................... Page 114<br />
Latrunculia (undescribed species) ..................................................... Page 115<br />
Order Halichondrida<br />
Family Axinellidae<br />
Axinella blanca Koltun, 1959 ........................................................... Page 116<br />
Axinella rugosa (Bowerbank, 1866) ..................................................... Page 117<br />
Family Bubaridae<br />
Bubaris vermiculata (Bowerbank, 1866) .................................................. Page 118<br />
Family Halichondriidae<br />
Halichondria (Halichondria) colossea Lundbeck, 1902 ....................................... Page 119<br />
Halichondria (Halichondria) lambei Brøndsted, 1933; (Arctic Ocean – Point Barrow, Beaufort Sea)<br />
Halichondria (Halichondria) oblonga (Hansen, 1885) ....................................... Page 120<br />
Halichondria (Eumastia) sitiens (Schmidt, 1870) .......................................... Page 121<br />
Halichondria sp. .................................................................... Page 122<br />
141
142 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Family Halichondriidae (continued)<br />
Hymeniacidon assimilis Levinsen, 1887 ....................................................Page 124<br />
Topsentia disparilis (Lambe, 1893) .......................................................Page 125<br />
Order Haplosclerida<br />
Family Chalinidae<br />
Cladocroce ventilabrum (Fristedt, 1887) ...................................................Page 126<br />
Haliclona bucina Tanita and Hoshino, 1989 ...............................................Page 127<br />
Haliclona (Gellius) digitata (Koltun, 1958) ................................................Page 128<br />
Haliclona (Gellius) primitiva (Lundbeck, 1902) ............................................Page 129<br />
Haliclona (Rhizoniera) rufescens (Lambe, 1892); (Gulf <strong>of</strong> Alaska, Bering Sea, Arctic Ocean)<br />
Haliclona tenuiderma (Lundbeck, 1902) ..................................................Page 130<br />
Haliclona (Haliclona) urceolus (Rathke and Vahl, 1806) .....................................Page 131<br />
Haliclona sp. 1 .......................................................................Page 132<br />
Haliclona sp. 2. ......................................................................Page 133<br />
Family Petrosiidae<br />
Xes<strong>to</strong>spongia hispida (Ridley and Dendy, 1886); (Gulf <strong>of</strong> Alaska)
Appendix II. Priorities for bycatch moni<strong>to</strong>ring<br />
This list includes <strong>the</strong> <strong>deep</strong>-<strong>water</strong> (>80 m) <strong>sponges</strong> presented<br />
in this <strong>guide</strong>. Each species is ranked from low<br />
(1) <strong>to</strong> high (3) for its importance as fish habitat and<br />
its vulnerability <strong>to</strong> disturbance from fishing activities.<br />
Ranks for <strong>the</strong>se two measures are averaged <strong>to</strong> provide<br />
Importance as Vulnerability <strong>to</strong><br />
Species fish habitat disturbance Score<br />
CLASS CALCAREA<br />
Clathrina sp. 1 2 1.5<br />
Leucandra poculiformis 1 1 1.0<br />
Leucandra tuba 2 2 2.0<br />
CLASS HEXACTINELLIDA<br />
Farrea kurilensis ssp. nov. 3 2 2.5<br />
Farrea occa occa 3 3 3.0<br />
Farrea sp. nov. 1 1 1.0<br />
Genus nov., sp. nov. 2 1 1.5<br />
Tre<strong>to</strong>dictyum sp. nov. 1 2 1.5<br />
Aphrocallistes vastus 3 3 3.0<br />
Heterochone calyx calyx 3 3 3.0<br />
Regadrella okinoseana 1 1 1.0<br />
Acanthascus (Acanthascus) pr<strong>of</strong>undum ssp. nov. 1 1 1.0<br />
Acanthascus (Rhabdocalyptus) dawsoni dawsoni 3 2 2.5<br />
Acanthascus (Rhabdocalyptus) mirabilis 1 1 1.0<br />
Acanthascus (Staurocalyptus) solidus 2 2 2.0<br />
Acanthascus (Staurocalyptus) sp. nov. 1 2 3 2.0<br />
Acanthascus (Staurocalyptus) sp. nov. 2 2 2 2.0<br />
Aulosaccus pinularis 1 1 1.0<br />
Aulosaccus schulzei 1 2 1.5<br />
Bathydorus sp. 1 1 1.0<br />
Caulophacus (Caulophacus) sp. nov. 2 1 1.5<br />
CLASS DEMOSPONGIAE<br />
Plakina atka 1 1 1.0<br />
Plakina tanaga 1 1 1.0<br />
Craniella arb 1 2 1.5<br />
Craniella sigmoancoratum 1 1 1.0<br />
Craniella spinosa 1 1 1.0<br />
Craniella sputnika 1 1 1.0<br />
Erylus aleuticus 1 1 1.0<br />
Geodia lendenfeldi 1 1 1.0<br />
Poecillastra tenuilaminaris 3 3 3.0<br />
Polymastia fluegeli 1 1 1.0<br />
Polymastia pacifica 1 1 1.0<br />
Stylocordyla borealis eous 1 2 1.5<br />
Aap<strong>to</strong>s kanuux 1 2 1.5<br />
Rhizaxinella clavata 1 2 1.5<br />
Suberites excellens 2 1 1.5<br />
Suberites simplex 1 1 1.0<br />
143<br />
a score. Species with scores greater than 2.0 should<br />
rank as a high priority for moni<strong>to</strong>ring as bycatch in<br />
commercial fisheries and s<strong>to</strong>ck assessment surveys.<br />
The scores for high priority species are presented in<br />
bold lettering.
144 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Importance as Vulnerability <strong>to</strong><br />
Species fish habitat disturbance Score<br />
CLASS DEMOSPONGIAE (continued)<br />
Suberites sp. 1 1 1.0<br />
Hemigellius porosus 1 2 1.5<br />
Cornulum clathriata 1 1 1.0<br />
Iophon piceum 1 2 1.5<br />
Iophon piceum abipocillus 1 2 1.5<br />
Megaciella anisochela 1 1 1.0<br />
Megaciella spirinae 1 2 1.5<br />
Clathria (Clathria) barleei 1 2 1.5<br />
Clathria (Clathria) laevigata 1 2 1.5<br />
Clathria (Axosuberites) lambei 1 2 1.5<br />
Echinoclathria vasa 1 1 1.0<br />
Artemisina amlia 1 2 1.5<br />
Artemisina arcigera 1 2 1.5<br />
Artemisina stipitata 2 3 2.5<br />
Artemisina sp. 2 3 2.5<br />
Coelosphaera oglalai 1 2 1.5<br />
Inflatella globosa 1 2 1.5<br />
Lissodendoryx (Lissodendoryx) behringi 1 2 1.5<br />
Lissodendoryx (Ectyodoryx) olgae 1 1 1.0<br />
Lissodendoryx (Lissodendoryx) oxeota 1 1 1.0<br />
Lissodendoryx (Lissodendoryx) papillosa 1 1 1.0<br />
Monanchora alaskensis 2 2 2.0<br />
Monanchora laminachela 1 2 1.5<br />
Monanchora pulchra 3 2 2.5<br />
Crella brunnea 1 1 1.0<br />
Hymedesmia (Stylopus) dermata 1 1 1.0<br />
Hymedesmia (Hymedesmia) irregularis 1 1 1.0<br />
Kirkpatrickia borealis 1 2 1.5<br />
Phorbas paucistylifer 1 2 1.5<br />
Melonanchora globogilva 1 1 1.0<br />
Myxilla (Myxilla) behringensis 1 1 1.0<br />
Myxilla (Ectyomyxilla) parasitica 1 1 1.0<br />
Myxilla (Bur<strong>to</strong>nanchora) pedunculata 1 2 1.5<br />
Stelodoryx oxeata 1 2 1.5<br />
Stelodoryx <strong>to</strong>poroki 2 1 1.5<br />
Stelodoryx vitiazi 1 1 1.0<br />
Echinostylinos hirsutus 1 1 1.0<br />
Tedania (Tedania) dirhaphis 1 2 1.5<br />
Tedania kagalaskai 2 2 2.0<br />
Asbes<strong>to</strong>pluma ramosa 2 2 2.0<br />
Cladorhiza bathycrinoides 1 1 1.0<br />
Cladorhiza corona 1 2 1.5<br />
Chondrocladia (Chondrocladia) concrescens 1 2 1.5<br />
Biemna variantia 1 2 1.5<br />
Euchelipluma elongata 1 1 1.0<br />
Guitarra abbotti 2 1 1.5<br />
Guitarra fimbriata 2 1 1.5<br />
Amphilectus digitatus 2 2 2.0<br />
Esperiopsis flagrum 2 1 1.5<br />
Semisuberites cribrosa 2 2 2.0
Importance as Vulnerability <strong>to</strong><br />
Species fish habitat disturbance Score<br />
CLASS DEMOSPONGIAE (continued)<br />
Mycale (Aegogropila) adhaerens 2 3 2.5<br />
Mycale (Carmia) carlilei 2 2 2.0<br />
Mycale (Mycale) jasoniae 2 2 2.0<br />
Mycale (Mycale) loveni 3 3 3.0<br />
Mycale (Mycale) tylota 1 2 1.5<br />
Latrunculia (Biannulata) oparinae 2 2 2.0<br />
Latrunculia velera 2 1 1.5<br />
Latrunculia sp. (undescribed) 1 1 1.0<br />
Axinella blanca 3 3 3.0<br />
Axinella rugosa 2 2 2.0<br />
Bubaris vermiculata 1 1 1.0<br />
Halichondria (Halichondria) colossea 2 1 1.5<br />
Halichondria (Halichondria) oblonga 1 2 1.5<br />
Halichondria (Eumastia) sitiens 2 2 2.0<br />
Halichondria sp. 2 3 2.5<br />
Hymeniacidon assimilis 2 2 2.0<br />
Topsentia disparilis 1 1 1.0<br />
Cladocroce ventilabrum 1 2 1.5<br />
Haliclona bucina 1 1 1.0<br />
Haliclona (Gellius) digitata 1 2 1.5<br />
Haliclona (Gellius) primitiva 1 2 1.5<br />
Haliclona tenuiderma 1 1 1.0<br />
Haliclona (Haliclona) urceolus 1 2 1.5<br />
Haliclona sp. 1 1 2 1.5<br />
Haliclona sp. 2 2 1 1.5<br />
145
146 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Appendix III. Glossary <strong>of</strong> terms<br />
Acanthose: spined<br />
Anchorate: anchored or anchor-like<br />
Apical: located at <strong>the</strong> apex; distal<br />
Arborescent: highly branched<br />
Areolate: surface covered with numerous circular areas<br />
Choanosome: interior tissue<br />
Clavate: club-shaped<br />
Conspecificity: belonging <strong>to</strong> <strong>the</strong> same species<br />
Conules: cone-shaped projections<br />
Conulose: surface with numerous conules raised by <strong>the</strong><br />
underlying skele<strong>to</strong>n<br />
Crenulate: surface with minutely notched or scalloped<br />
projections<br />
Dactylate: finger-like projections<br />
Digitate: <strong>deep</strong>ly divided finger-like projections<br />
Ecomorph: growth form adapted for different ecological<br />
regime<br />
Ec<strong>to</strong>some: exterior surface<br />
Emergent epifauna: fauna having most <strong>of</strong> its parts elevated<br />
above <strong>the</strong> level <strong>of</strong> <strong>the</strong> surrounding seafloor<br />
Endopsammic: main part <strong>of</strong> <strong>the</strong> body buried in sand<br />
Epizoic: living attached <strong>to</strong> <strong>the</strong> body <strong>of</strong> an animal; a<br />
non-parasitic animal that lives attached <strong>to</strong> <strong>the</strong> outer<br />
surface <strong>of</strong> ano<strong>the</strong>r animal<br />
Eurybathic: broadly distributed with depth<br />
Filiform: shaped like a thread<br />
Fistules: cone-shaped elevations or tube-like protuberances<br />
Flabellate: fan-shaped<br />
Flagelliform: shaped as a single, very long, erect branch<br />
Foliate: shaped like a leaf<br />
Globular: ball-shaped or spherical<br />
Gonochoristic: having male and female individuals in<br />
<strong>the</strong> same population<br />
Hispid: covered with bristly hairs<br />
Holotype: <strong>the</strong> single specimen used as <strong>the</strong> basis for <strong>the</strong><br />
first published description <strong>of</strong> a species and designated<br />
as <strong>the</strong> type specimen<br />
Labyrinthic: an intricate structure <strong>of</strong> interconnecting<br />
passages<br />
Lobate: having lobes or rounded projections<br />
Mammillate: shaped like a nipple<br />
Massive: large and compact but amorphous or without<br />
a definable shape (note that <strong>the</strong> term “lumpy” is used<br />
in some <strong>of</strong> <strong>the</strong> sponge literature)<br />
Megascleres: large structural spicules<br />
Microscleres: small <strong>to</strong> minute reinforcing or packing<br />
spicules<br />
Osculum (a): <strong>the</strong> opening through which <strong>water</strong> is expelled<br />
from <strong>the</strong> sponge<br />
Ostium (a): <strong>the</strong> opening through which <strong>water</strong> enters<br />
<strong>the</strong> sponge<br />
Oviparous: producing eggs that are laid and hatch<br />
externally<br />
Papillae: Nipple-like protuberances projecting from <strong>the</strong><br />
sponge surface that bear ei<strong>the</strong>r ostia, oscula, or both<br />
Pellicular: covered with a liquid film or organic membrane<br />
Polymorphic: occurring in different shapes<br />
Polyspicular: composed <strong>of</strong> multiple spicule types<br />
Saccate: shaped like a sac or pouch<br />
Sibling species: a species that closely resembles ano<strong>the</strong>r<br />
in appearance and o<strong>the</strong>r characteristics but cannot<br />
interbreed with it<br />
Spicules: small needle-like structures made <strong>of</strong> silica or<br />
calcium carbonate that support <strong>the</strong> s<strong>of</strong>t tissues <strong>of</strong><br />
<strong>sponges</strong><br />
Spongin: <strong>the</strong> fibrous framework <strong>of</strong> collagen that forms<br />
<strong>the</strong> sponge’s organic skele<strong>to</strong>n<br />
Spongocoel (= atrium): <strong>the</strong> central exhalant cavity <strong>of</strong><br />
<strong>the</strong> sponge<br />
S<strong>to</strong>lon: a creeping or prostrate rope-like structure<br />
Subglobular: deviating slightly from globular<br />
Subspecies: a morphologically and genetically distinct<br />
population within a species that inhabits a separate<br />
geographic area or depth range<br />
Tabular: having a plane surface or flat<br />
Tuberculate: warty or verrucose<br />
Uncinate: large straight diactin spicules covered with<br />
barbs and brackets all inclined at one end<br />
Viviparous: producing live <strong>of</strong>fspring from within <strong>the</strong><br />
body <strong>of</strong> <strong>the</strong> parent
Appendix IV. Spicule types<br />
This appendix includes scanning electron microscopy<br />
(SEM) images <strong>of</strong> spicules from select calcareous<br />
<strong>sponges</strong> and demo<strong>sponges</strong> collected during <strong>the</strong><br />
2004 Aleutian Island expeditions. It also includes<br />
several SEM images <strong>of</strong> spicules from a few hexactinellid<br />
sponge species that are common in Alaskan<br />
<strong>water</strong>s. The purpose <strong>of</strong> <strong>the</strong> appendix is <strong>to</strong> provide<br />
<strong>the</strong> reader with a representative collection <strong>of</strong> spicule<br />
images so that <strong>the</strong>y may gain an understanding <strong>of</strong> <strong>the</strong><br />
terms used in <strong>the</strong> <strong>guide</strong> and an appreciation for <strong>the</strong><br />
Class Calcarea, Family Grantiidae, Leucandra poculiformis Hozawa, 1918.<br />
147<br />
variation between species and among spicule types.<br />
This collection also serves as a source <strong>of</strong> reference<br />
material for those who wish <strong>to</strong> microscopically examine<br />
<strong>sponges</strong>. We refer <strong>the</strong> reader <strong>to</strong> <strong>the</strong> Thesaurus<br />
<strong>of</strong> Sponge Morphology (Boury-Esnault and Rützler,<br />
1997) for a comprehensive glossary <strong>of</strong> terminology<br />
and a pic<strong>to</strong>rial <strong>guide</strong> <strong>to</strong> spicule types and skeletal<br />
structures. The species in this appendix have been<br />
arranged systematically <strong>to</strong> family and alphabetically<br />
arranged within family.<br />
(A) Overview <strong>of</strong> occurring spicules: large tri- and tetractines and small oxeas; (B) large triactine; (C) a rare large<br />
tetractine; and (D) acanthose microxeas with characteristic bend called a “lance-head” by Hozawa.
148 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Leucandra poculiformis Hozawa, 1918 (continued)<br />
Class Calcarea, Family Grantiidae, Leucandra tuba Hozawa, 1918.<br />
(A) Pugiole with unusual biforked end; (B) pugioles, small and large oxeas; (C) typical pugiole with normal points;<br />
(D) giant tetractines among small category <strong>of</strong> triactines; and (E) triactines among minute oxeote spicule types.
Leucandra poculiformis Hozawa, 1918 (continued)<br />
149
150 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Class Hexactinellida, Family Farreidae, Farrea occa occa Bowerbank, 1862.<br />
(A) An anchorate clavule; (B) a pileate clavule; (C) a dermal pentactin; (D) dictyonal framework; and (E) an oxyhexaster.
Farrea occa occa Bowerbank, 1862 (continued)<br />
Class Hexactinellida, Family Aphrocallistidae, Aphrocallistes vastus Schulze, 1886.<br />
(A) Dictyonal framework (scale bar equals 500 µm); (B) a pentactin from a juvenile specimen; (C) a scopule<br />
from a juvenile specimen; (D) a scopule head; (E) a spiny oxyhexaster; and (F) a s<strong>to</strong>ut oxyhexaster.<br />
151
152 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Aphrocallistes vastus Schulze, 1886 (continued)
Class Demospongiae, Family Plakinidae, Plakina atka Lehnert, S<strong>to</strong>ne and Heimler, 2005.<br />
(A) Spined calthrops with reduced 4 th ray, all rays with basal spines; (B) micr<strong>of</strong>urcate end <strong>of</strong> ray from a tetraloph,<br />
tetrafurcate lophocalthrops; (C) tetralophose calthrops, all rays tetrafurcate, points <strong>of</strong> rays micr<strong>of</strong>urcate; (D) smooth<br />
diods, smooth triods, and thicker triods with strongly spined bases <strong>of</strong> rays; and (E) thick category <strong>of</strong> diods with central<br />
spines. Probably derived from spined triods through reduction <strong>of</strong> one ray. (Figures A–E reproduced with permission<br />
from Zootaxa, Magnolia Press.)<br />
153
154 Pr<strong>of</strong>essional Paper <strong>NMFS</strong> 12<br />
Plakina atka Lehnert, S<strong>to</strong>ne and Heimler, 2005 (continued)
Class Demospongiae, Family Plakinidae, Plakina tanaga Lehnert, S<strong>to</strong>ne and Heimler, 2005.<br />
(A) Smooth oxeas, triods, and trilophose calthrops; (B) weakly spined triods and trilophose calthrops; and (C)<br />
trilophose calthrops with tetrafurcate lophose rays and biforked non-lophose ray. (Figures A–C reproduced with<br />
permission from Zootaxa, Magnolia Press.)<br />
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Class Demospongiae, Family Geodiidae, Erylus aleuticus Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) An anisotylote (one <strong>of</strong> several unusual spicule types occurring in this species); (B) a juvenile apidaster with different<br />
surface structure; (C) adult apidasters; (D) a centrolyte microstrongyle on <strong>the</strong> surface <strong>of</strong> an apidaster; (E)<br />
cladome <strong>of</strong> triaene with bent rays; (F) star-shaped surface structure on adult aspidaster; (G) oxyaster; and (H) surface<br />
structures on aspidasters may vary in geometry. (Figures B, C, D, E, and F reproduced with permission from <strong>the</strong><br />
Journal <strong>of</strong> <strong>the</strong> Marine Biological Association <strong>of</strong> <strong>the</strong> United Kingdom.)
Erylus aleuticus Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
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Class Demospongiae, Family Polymastiidae, Polymastia fluegeli Lehnert, S<strong>to</strong>ne and Heimler, 2005.<br />
(A) Small category <strong>of</strong> tylostyle and tyle <strong>of</strong> large tylostyle (note <strong>the</strong> radiolarian skele<strong>to</strong>n at lower right); (B) large and<br />
small tylostyles; and (C) small category <strong>of</strong> tylostyles on large tylostyles. (Figures A and B reproduced with permission<br />
from Facies, Springer Verlag.)
Class Demospongiae, Family Suberitidae, Aap<strong>to</strong>s kanuux Lehnert, Hocevar and S<strong>to</strong>ne, 2008.<br />
(A) Overview <strong>of</strong> spicules; large strongyloxeas, medium-size subtylostyles, and short ec<strong>to</strong>somal tylostyles; (B) mediumsized<br />
subtylostyles and ec<strong>to</strong>somal tylostyles; (C) ec<strong>to</strong>somal tylostyles and ends <strong>of</strong> medium-sized spicules. (Figures A–C<br />
reproduced with permission from Zootaxa, Magnolia Press.)<br />
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Class Demospongiae, Family Acarnidae, Megaciella anisochela Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Category <strong>of</strong> thick styles among smaller tylotes and microscleres; (B) small category <strong>of</strong> palmate isochela; (C) large<br />
category <strong>of</strong> palmate isochela; (D) acanthose end <strong>of</strong> tylote (close-up view); (E) a dis<strong>to</strong>rted, tiny anisochela (this is <strong>the</strong><br />
only species <strong>of</strong> Megaciella with anisochelae); and (F) an anisochela. (Figures A–D reproduced with permission from<br />
Zootaxa, Magnolia Press.)
Megaciella anisochela Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
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Class Demospongiae, Family Microcionidae, Echinoclathria vasa Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) A palmate isochela with narrow alae (note <strong>the</strong> rounded ends <strong>of</strong> alae); (B) long ec<strong>to</strong>somal thin styles among thick<br />
styles; (C) a palmate isochela, both alae with two points (acanthose end <strong>of</strong> thin style at upper right); (D) thick styles;<br />
(E) reduced palmate isochela (note different points on <strong>the</strong> alae); and (F) acanthose end <strong>of</strong> thin style. (Figures A–F<br />
reproduced with permission from Zootaxa, Magnolia Press.)
Echinoclathria vasa Lehnert, S<strong>to</strong>ne and Heimler (continued)<br />
Class Demospongiae, Family Microcionidae, Artemisina amlia Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) A smooth <strong>to</strong>xon; (B) a large, smooth style; (C) a thin style with prominent <strong>to</strong>oth (upper left <strong>to</strong> lower right); (D)<br />
an isochela; and (E) a close-up view <strong>of</strong> prominent <strong>to</strong>oth on thin category <strong>of</strong> style. (Figures A–E reproduced with<br />
permission from Zootaxa, Magnolia Press.)<br />
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Artemisina amlia Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)
Class Demospongiae, Family Coelosphaeridae, Coelosphaera oglalai Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) A choanosomal strongyle among microscleres; (B) microacanthose end <strong>of</strong> a choanosomal strongyle (close-up<br />
view); (C) acanthose end <strong>of</strong> tylote (close-up view); (D) isochelae (right) and acanthose end <strong>of</strong> a strongyle (left); and<br />
(E) an ec<strong>to</strong>somal tylote (center). (Figures A–E reproduced with permission from Zootaxa, Magnolia Press.)<br />
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Coelosphaera oglalai Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
Class Demospongiae, Family Crambeidae, Monanchora laminachela Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Choanosomal thick style among thin styles; (B) an ec<strong>to</strong>somal thin style; (C) an anchorate isochela with characteristic<br />
central plate; (D) variation <strong>of</strong> isochela with fused alae; (E) an isochela (rear view); and (F) a sigma. (Figures<br />
A–D and F reproduced with permission from Zootaxa, Magnolia Press.)
Monanchora laminachela Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
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Class Demospongiae, Family Myxillidae, Melonanchora globogilva Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) An ec<strong>to</strong>somal tylote among microscleres; (B) smooth end <strong>of</strong> tylote and pointed end <strong>of</strong> acanthostyle (close-up<br />
view); (C) acanthostyles; (D) a large isochelae (shaft <strong>of</strong> inner margins <strong>of</strong> alae with fimbriae); (E) fimbriate inner<br />
margins <strong>of</strong> alae <strong>of</strong> isochelae (close-up view); (F) second type <strong>of</strong> large isochela with different shape and dented outer<br />
margins; (G) small category <strong>of</strong> isochelae; and (H) dented margins <strong>of</strong> second type <strong>of</strong> isochela (close-up view). (Figures<br />
A–H reproduced with permission from Zootaxa, Magnolia Press.)
Melonanchora globogilva Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
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Class Demospongiae, Family Myxillidae, Stelodoryx oxeata Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) A large oxea with acanthose ends among <strong>to</strong>rnotes with acanthose ends; (B) acanthose end <strong>of</strong> <strong>to</strong>rnote and a sigma;<br />
(C) acanthose end <strong>of</strong> oxea (close-up view); (D) large category <strong>of</strong> anchorate isochela; (E) medium-sized category<br />
<strong>of</strong> anchorate, polydentate isochela; (F) small category <strong>of</strong> anchorate isochelae and centrolyte sigmas; (G) acanthose<br />
end <strong>of</strong> ec<strong>to</strong>somal <strong>to</strong>rnote, centrotylote sigma (above) and medium-sized isochela (below); and (H) large category <strong>of</strong><br />
isochela (side view). (Figures A–F reproduced with permission from Zootaxa, Magnolia Press.)
Stelodoryx oxeata Lehnert, S<strong>to</strong>ne and Heimler (continued)<br />
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Class Demospongiae, Family Tedaniidae, Tedania kagalaskai Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Thick choanosomal styles and ec<strong>to</strong>somal tylotes; (B) acanthose end <strong>of</strong> tylote (close-up view); (C) large category<br />
<strong>of</strong> onychaete (lower left <strong>to</strong> upper right) among tylotes; and (D) small category <strong>of</strong> onychaete (note <strong>the</strong> unequal ends).<br />
(Figures A–D reproduced with permission from Zootaxa, Magnolia Press.)
Class Demospongiae, Family Cladorhizidae, Cladorhiza bathycrinoides Koltun, 1955.<br />
(A) An anisochela (rear view) with sigma (below); (B) a sigma; (C) a sigmancistra; (D) an anisochela; (E) an anisochela<br />
(rear view); (F) small end <strong>of</strong> anisochela with sharp claws (possibly adapted <strong>to</strong> capture prey) and pointed end<br />
<strong>of</strong> sigmancistra (above); (G) tylostrongyles and anisochelae; and (H) tylostrongyle and anisochelae.<br />
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Cladorhiza bathycrinoides Koltun, 1955 (continued)
Class Demospongiae, Family Cladorhizidae, Cladorhiza corona Lehnert, Watling and S<strong>to</strong>ne, 2005.<br />
(A) Claw-like appendages at <strong>the</strong> small end <strong>of</strong> anisochelae; (B) a sigmancistra; (C) a tylostyle among microscleres;<br />
(D) a large fusiform style and short oxeas; (E) an anisochela with claw-like appendages; (F) a small (anis-)oxea (a<br />
spicule type present in <strong>the</strong> basal plate); and (G) large fusiform styles and small (anis-)oxea. (Figure A reproduced<br />
with permission from <strong>the</strong> Journal <strong>of</strong> <strong>the</strong> Marine Biological Association <strong>of</strong> <strong>the</strong> United Kingdom.)<br />
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Cladorhiza corona Lehnert, Watling and S<strong>to</strong>ne, 2005 (continued)
Class Demospongiae, Family Guitarridae, Euchelipluma elongata Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Fusiform choanosomal style; (B) an isochela and sigmas; (C) ec<strong>to</strong>somal tylostyle, slightly polytylote among microscleres;<br />
(D) characteristically thin placochela; (E) developmental stage <strong>of</strong> polytylote tylostyle and two placochelae;<br />
(F) a sigma; (G) an isochela, placochela and sigmas; and (H) long fusiform styles, shorter ec<strong>to</strong>somal tylostyles and<br />
microscleres. (Figures A–F reproduced with permission from Zootaxa, Magnolia Press.)<br />
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Euchelipluma elongata Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)
Class Demospongiae, Family Guitarridae, Guitarra abbotti Lee, 1987.<br />
(A) Fusiform styles; (B) a large placochela among styles; (C) biplacochelae; (D) fimbriae <strong>of</strong> biplacochela (closeup<br />
view); (E) tiny, spined isochela; (F) a biplacochela showing frontal process; and (G) fimbriae <strong>of</strong> a biplacochela<br />
(close-up view).<br />
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Guitarra abbotti Lee, 1987 (continued)
Class Demospongiae, Family Esperiopsidae, Esperiopsis flagrum Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Microscleres (large and small categories <strong>of</strong> isochelae and sigmas); (B) a small isochela; (C) long fusiform styles<br />
and microscleres; (D) large and small sigmas; (E) a small isochela (side view); (F) a large isochela (rear view); and<br />
(G) large isochelae (front and rear views). (Figures A–F reproduced with permission from Zootaxa, Magnolia Press.)<br />
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Esperiopsis flagrum Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)
Class Demospongiae, Family Mycalidae, Mycale (Carmia) carlilei Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Spicule overview (styles, anisochelae and sigmas); (B) an anisoschela (<strong>to</strong>p view); (C) an anisochela (side view);<br />
and (D) a sigma. (Figures A–D reproduced with permission from Zootaxa, Magnolia Press.)<br />
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Class Demospongiae, Family Mycalidae, Mycale (Mycale) jasoniae Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Spicule overview (subtylostyles, large and small anisochelae, and rhaphids); (B) large anisochelae (front, side<br />
and rear views); (C) side view <strong>of</strong> a large anisochela (note <strong>to</strong>p view <strong>of</strong> broken end <strong>of</strong> small anisochela); (D) small<br />
category <strong>of</strong> anisochela and rhaphids; and (E) a small anisochela. (Figures A–D reproduced with permission from<br />
Zootaxa, Magnolia Press.)
Mycale (Mycale) jasoniae Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
Class Demospongiae, Family Latrunculiidae, Latrunculia velera Lehnert, S<strong>to</strong>ne and Heimler, 2006.<br />
(A) Spicule overview (styles and anisodiscorhabds); (B) acanthose end <strong>of</strong> style; (C) a rare polytylote tylostyle; (D) an<br />
anisodiscorhabd; (E) an anisodiscorhabd (apex view); (F) apex <strong>of</strong> anisodiscorhabd (side view); (G) early developmental<br />
stage <strong>of</strong> anisodiscorhabd on pointed end <strong>of</strong> style (acanthose pointed ends, as shown here, are rare); (H) developmental<br />
stage <strong>of</strong> anisdiscorhabd between two fully developed ones; (I) smoo<strong>the</strong>r variation <strong>of</strong> an anisodiscorhabd;<br />
and (J) anisodiscorhabds. (Figures A–I reproduced with permission from Zootaxa, Magnolia Press.)<br />
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Latrunculia velera Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)
Latrunculia velera Lehnert, S<strong>to</strong>ne and Heimler, 2006 (continued)<br />
187