U.S. Department of Commerce

September 2011

NOAA Professional Paper NMFS 12

A guide to the deep-water sponges 

of the Aleutian Island Archipelago

Robert P. Stone

Helmut Lehnert

Henry Reiswig
U.S. Department

of Commerce

Rebecca Blank

Acting Secretary of Commerce

National Oceanic

and Atmospheric

Administration

Jane Lubchenco, Ph.D.

Administrator of NOAA

National Marine 

Fisheries Service

Eric C. Schwaab

Assistant Administrator 

for Fisheries

The NOAA Professional Paper NMFS 

(ISSN 1931-4590) series is published 

by the Scientific Publications Office, 

National Marine Fisheries Service, 

NOAA, 7600 Sand Point Way NE, 

Seattle, WA 98115.

The Secretary of Commerce has

determined that the publication of 

this series is necessary in the transaction of the public business required by 

law of this Department. Use of funds 

for printing of this series has been approved by the Director of the Office 

of Management and Budget.

NOAA Professional 

Papers NMFS

Scientific Editor

Richard D. Brodeur, Ph.D.

Associate Editor

Julie Scheurer

National Marine Fisheries Service

Northwest Fisheries Science Center

2030 S. Marine Science Dr.

Newport, Oregon 97365-5296

Managing Editor

Shelley Arenas

National Marine Fisheries Service

Scientific Publications Office

7600 Sand Point Way NE

Seattle, Washington 98115

Editorial Committee

Ann C. Matarese, Ph.D. National Marine Fisheries Service

James W. Orr, Ph.D. National Marine Fisheries Service

Bruce L. Wing, Ph.D. National Marine Fisheries Service

The NOAA Professional Paper NMFS series carries peer-reviewed, lengthy original 

research reports, taxonomic keys, species synopses, flora and fauna studies, and data-intensive reports on investigations in fishery science, engineering, and economics. Copies 

of the NOAA Professional Paper NMFS series are available free in limited numbers to 

government agencies, both federal and state. They are also available in exchange for 

other scientific and technical publications in the marine sciences. Professional Papers 

are published online in PDF format at http://spo.nmfs.noaa.gov

NOTICE: This series was established in 2003 to replace the NOAA Technical Report 

NMFS series. 
NOAA Professional Paper NMFS 12

A guide to the deep-water sponges 

of the Aleutian Island Archipelago

Robert P. Stone

Helmut Lehnert

Henry Reiswig

September 2011

U.S. Department of Commerce

Seattle, Washington
Suggested reference

Stone, Robert P., Helmut Lehnert, and Henry Reiswig. 2011. A guide to the deepwater sponges of the Aleutian Island Archipelago. NOAA Professional Paper 

NMFS 12, 187 p.

Online dissemination

This report is posted online in PDF format at http://spo.nmfs.noaa.gov (click on 

Professional Papers link).

Copyright law

Although the contents of the Professional Papers have not been copyrighted and 

may be reprinted entirely, reference to source is appreciated.

Proprietary products

The National Marine Fisheries Service (NMFS) does not approve, recommend, 

or endorse any proprietary product or proprietary material mentioned in this 

publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or 

imply that NMFS approves, recommends, or endorses any proprietary product 

or proprietary material mentioned herein, or which has as its purpose an intent 

to cause directly or indirectly the advertised product to be used or purchased 

because of this NMFS publication.
iii

CONTENTS

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

About this guide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Zoogeography of sponges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Biology of sponges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Ecology of sponges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Importance as fish habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Vulnerability to disturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Monitoring bycatch of sponges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Collection and preservation of sponge specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Laboratory identification of sponge specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Calcareous sponges and demosponges........................................................... 9

Hexactinellid sponges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Class Calcarea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 1. Clathrina sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 2. Leucandra poculiformis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 3. Leucandra tuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Class Hexactinellida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 4. Farrea kurilensis ssp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

5. Farrea occa occa............................................................................ 17

 6. Farrea sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

 7. Family Euretidae; Genus nov., sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

 8. Tretodictyum sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

 9. Aphrocallistes vastus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

10. Heterochone calyx calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

11. Regadrella okinoseana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

12. Acanthascus (Acanthascus) profundum ssp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

13. Acanthascus (Rhabdocalyptus) dawsoni dawsoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

14. Acanthascus (Rhabdocalyptus) mirabilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

15. Acanthascus (Staurocalyptus) solidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

16. Acanthascus (Staurocalyptus) sp. nov. 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

17. Acanthascus (Staurocalyptus) sp. nov. 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

18. Aulosaccus pinularis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

19. Aulosaccus schulzei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

20. Bathydoris sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

21. Caulophacus (Caulophacus) sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

Class Demospongiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

22. Plakina atka . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

23. Plakina tanaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

24. Craniella arb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

25. Craniella sigmoancoratum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

26. Craniella spinosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

27. Craniella sputnika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

28. Erylus aleuticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

29. Geodia lendenfeldi nomen novum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

30. Poecillastra tenuilaminaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
iv

CONTENTS (continued)

31. Polymastia fluegeli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

32. Polymastia pacifica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

33. Stylocordyla borealis eous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

34. Aaptos kanuux. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

35. Rhizaxinella clavata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

36. Suberites excellens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

37. Suberites simplex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

38. Suberites sp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

39. Hemigellius porosus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

40. Cornulum clathriata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

41. Iophon piceum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

42. Iophon piceum abipocillus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

43. Megaciella anisochela. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

44. Megaciella spirinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

45. Clathria (Clathria) barleei. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

46. Clathria (Clathria) laevigata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62

47. Clathira (Axosuberites) lambei. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

48. Echinoclathria vasa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

49. Artemisina amlia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

50. Artemisina arcigera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

51. Artemisina stipitata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

52. Artemisina sp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

53. Coelosphaera oglalai. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

54. Inflatella globosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

55. Lissodendoryx (Lissodendoryx) behringi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

56. Lissodendoryx (Ectyodoryx) olgae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

57. Lissodendoryx (Lissodendoryx) oxeota. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

58. Lissodendoryx (Lissodendoryx) papillosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

59. Monanchora alaskensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75

60. Monanchora laminachela. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

61. Monanchora pulchra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

62. Crella brunnea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

63. Hymedesmia (Stylopus) dermata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

64. Hymedesmia (Hymedesmia) irregularis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

65. Kirkpatrickia borealis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

66. Phorbas paucistylifer. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

67. Melonanchora globogilva. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

68. Myxilla (Myxilla) behringensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

69. Myxilla (Ectyomyxilla) parasitica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

70. Myxilla (Burtonanchora) pedunculata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

71. Stelodoryx oxeata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

72. Stelodoryx toporoki. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

73. Stelodoryx vitiazi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

74. Echinostylinos hirsutus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
v

CONTENTS (continued)

75. Tedania (Tedania) dirhaphis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91

76. Tedania kagalaskai. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92

77. Asbestopluma ramosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

78. Cladorhiza bathycrinoides. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

79. Cladorhiza corona. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

80. Chondrocladia (Chondrocladia) concrescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

81. Biemna variantia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98

82. Euchelipluma elongata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99

83. Guitarra abbotti. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100

84. Guitarra fimbriata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

85. Amphilectus digitatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102

86. Esperiopsis flagrum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

87. Semisuberites cribrosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104

88. Mycale (Aegogropila) adhaerens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106

89. Mycale (Carmia) carlilei. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

90. Mycale (Mycale) jasoniae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108

91. Mycale (Mycale) loveni. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

92. Mycale (Mycale) tylota. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112

93. Latrunculia (Biannulata) oparinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

94. Latrunculia velera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114

95. Latrunculia sp. (undescribed). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

96. Axinella blanca. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

97. Axinella rugosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117

98. Bubaris vermiculata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118

99. Halichondria (Halichondria) colossea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

100. Halichondria (Halichondria) oblonga. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

101. Halichondria (Eumastia) sitiens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

102. Halichondria sp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122

103. Hymeniacidon assimilis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

104. Topsentia disparilis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

105. Cladocroce ventilabrum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126

106. Haliclona bucina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

107. Haliclona (Gellius) digitata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128

108. Haliclona (Gellius) primitiva. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

109. Haliclona tenuiderma. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130

110. Haliclona (Haliclona) urceolus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131

111. Haliclona sp. 1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132

112. Haliclona sp. 2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133

Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

Literature cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

Appendix I. Sponge species reported from Alaskan waters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Appendix II. Priorities for bycatch monitoring. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

Appendix III. Glossary of terms. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

Appendix IV. Spicule types. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

1

Introduction

The first dedicated collections of 

deep-water (>80 m) sponges from 

the central Aleutian Islands revealed 

a rich fauna comprising 102 species, 

including 28 species new to science 

and range extensions for 53 species. 

Based on these collections and the 

published literature, we now confirm the presence of 125 species (or 

subspecies) of deep-water sponges 

in the central Aleutian Islands. The 

inventory includes 10 species of calcareous sponges, 20 species of hexactinellid sponges, and 95 species of 

demosponges. Despite the initial collection efforts, the sponge fauna of 

this region of the North Pacific is still 

poorly known. Based on our extensive 

submersible observations, we estimate 

that there are several hundred sponge 

species yet to be inventoried or described from the region. This regional 

estimate (i.e., total percent known) 

is consistent with those made for all 

sponges worldwide (Hooper and Lévi, 

1994; Hooper and Van Soest, 2002).

The main purpose of this guide is to 

promote an awareness and appreciation of the importance of the sponge 

fauna in the North Pacific Ocean, 

particularly in the Aleutian Islands 

A guide to the deep-water sponges 

of the Aleutian Island Archipelago

Robert P. Stone (contact author)1

Helmut Lehnert2

Henry Reiswig3

1 Alaska Fisheries Science Center

National Marine Fisheries Service

National Oceanic and Atmospheric Administration

17109 Point Lena Loop Road

Juneau, Alaska 99801 

Email address for contact author: Bob.Stone@noaa.gov

2 Freelance Sponge Taxonomy

Eichenstr. 14, D-86507

Oberottmarshausen, Germany

3 Royal British Columbia Museum and University of Victoria (Biology)

675 Belleville Street

Victoria, British Columbia, Canada V8W 3N5

Abstract—The first dedicated collections of deep-water (>80 m) sponges from 

the central Aleutian Islands revealed a 

rich fauna including 28 novel species and 

geographical range extensions for 53 others. Based on these collections and the 

published literature, we now confirm the 

presence of 125 species (or subspecies) 

of deep-water sponges in the Aleutian 

Islands. Clearly the deep-water sponge 

fauna of the Aleutian Islands is extraordinarily rich and largely understudied. 

Submersible observations revealed that 

sponges, rather than deep-water corals, 

are the dominant feature shaping benthic 

habitats in the region and that they provide important refuge habitat for many 

species of fish and invertebrates including 

juvenile rockfish (Sebastes spp.) and king 

crabs (Lithodes sp). Examination of video 

footage collected along 127 km of the 

seafloor further indicate that there are 

likely hundreds of species still uncollected 

from the region, and many unknown 

to science. Furthermore, sponges are 

extremely fragile and easily damaged by 

contact with fishing gear. High rates of 

fishery bycatch clearly indicate a strong 

interaction between existing fisheries and 

sponge habitat. Bycatch in fisheries and 

fisheries-independent surveys can be a major source of information on the location 

of the sponge fauna, but current monitoring programs are greatly hampered by the 

inability of deck personnel to identify bycatch. This guide contains detailed species 

descriptions for 112 sponges collected in 

Alaska, principally in the central Aleutian 

Islands. It addresses bycatch identification 

challenges by providing fisheries observers and scientists with the information 

necessary to adequately identify sponge 

fauna.Using that identification data, areas of high abundance can be mapped 

and the locations of indicator species 

of vulnerable marine ecosystems can be 

determined. The guide is also designed 

for use by scientists making observations 

of the fauna in situ with submersibles, 

including remotely operated vehicles and 

autonomous underwater vehicles.

where the diversity and abundance of 

sponges appears to be extraordinary 

and bycatch in existing fisheries continues to be a major concern for resource managers. Bycatch in fisheries 

and fisheries-independent surveys is 

a major source of information on the 

location of the sponge fauna and also 

a source of specimens for study. This 

guide serves the additional purpose of 

providing fisheries observers and scientists with the information necessary 

to adequately identify sponge fauna 

so that the data can be included in 

existing databases. These data can be 

used to map areas of high abundance 

and the locations of indicator species 

of vulnerable marine ecosystems. The 

guide is also designed for use by scientists making observations of the fauna 

in situ with submersibles, including 

remotely operated vehicles (ROVs) 

and autonomous underwater vehicles 

(AUVs).

The protocols used to identify bycatch of sponges, both in commercial 

fishing operations and in fisheries surveys, have major shortcomings. They 

are largely restricted by the limited 

number of personnel dedicated to 

this task, a general lack of knowledge 

of the taxonomy of the sponge fauna, 

and the fact that sponges are particu-
2 Professional Paper NMFS 12

larly fragile and are often highly fragmented at the time 

of collection. 

Current practice aboard commercial fishing vessels 

in Alaska (note that not all vessels are required to have 

fishery observers onboard) calls for tasked personnel 

to identify sponge bycatch to the lowest practical taxa 

– essentially to Phylum Porifera. The Alaska Fisheries 

Science Center’s Fisheries Monitoring and Analysis 

Division (FMA) maintains all records, including depth 

and location data, through its North Pacific Groundfish 

Observer Program. The accuracy of those data depends 

foremost on the type of fishing gear, but also varies due 

to the spatial coverage of the gear (i.e., only start and 

end positions and depths are recorded). Specimens are 

typically not retained for specific scientific purposes. 

Current practice aboard NOAA survey vessels differs between the two principal surveys. Specimens are 

retained for further study upon request, but no formal 

program for the collection and preservation of specimens exists otherwise. The emphasis of both surveys 

is to gather information necessary for the sustainable 

management of groundfish species. Monitoring the 

bycatch of structure-forming invertebrates, such as 

sponges, has become more of a priority with recent 

emphasis on managing fisheries with an ecosystem approach. For the NMFS sablefish longline survey, sponges 

are only identified to the general level of “unidentified 

sponge.” For the NMFS groundfish trawl survey, wet 

weight of sponges is recorded either completely or by 

subsampling. Collection data are then coded (NMFS 

RACE Species Code Book – maintained by the Resource 

Assessment and Conservation Engineering Division’s 

Groundfish Assessment Program) and entered into a 

database. Sponges are identified from a guide that was 

developed specifically for the identification of benthic 

marine invertebrates collected along Alaska’s upper 

continental slope and shelf (Clark1

). This guide has 

been an important first step toward more adequately 

monitoring sponge bycatch, but it is largely incomplete, contains species that have never been confirmed 

to occur in Alaskan waters, and lists some species with 

inaccurate taxonomic nomenclature. Clearly there is a 

strong need for a guide dedicated to the identification 

of Alaskan sponges and a continued effort to properly 

identify the sponge fauna collected from the region. 

The sponges contained in this guide were collected 

principally during two expeditions to the central Aleutian Islands in 2004: one aboard the RV Velero IV and 

the other aboard the RV Roger Revelle. A few additional 

sponge specimens were collected during the pioneering 

work in the region aboard the RV Velero IV in 2002 and 

2003. The focus of the research supporting the cruises 

was on deep-water coral habitat; there was a dedicated 

effort to collect sponge fauna for formal identification 

only during the RV Velero IV cruise in 2004 and even then 

it was of secondary importance. We also include collections made in the eastern Gulf of Alaska aboard the RV 

Velero IV in 2005 and in the Bering Sea Canyons (Pribilof 

and Zhemchug) aboard the RV Esperanza in 2007.

We reviewed video footage of the seafloor collected 

during 31 dives with the submersible Delta and 13 dives 

with the ROV Jason II. A total of approximately 127 km 

of seafloor habitat was examined (35 km and 92 km, 

respectively) from that video footage for the presence 

of sponge fauna, with particular focus on taxa included 

in this guide. Habitat information including depth, 

substrate, and associated fauna were recorded for 

sponge observations, often made from multiple camera 

perspectives. 

About this guide

This guide contains detailed species descriptions for 112 

sponges collected in Alaska, principally in the central 

Aleutian Islands. Each species description begins with 

the scientific name using classical binominal nomenclature. The first name (always capitalized) is the genus. 

The second name (never capitalized) is the species. 

Some species may have a third name (never capitalized) for designated subspecies. And for some species 

a subgenus may have been designated and is placed in 

parentheses after the current genus. The name(s) of the 

author(s) of the species description (i.e., the person(s) 

who described the species) and year of description 

follow the scientific name. Parentheses placed around 

the author(s) name(s) indicates that there has been an 

accepted modification to genus assignment since the 

original description. For example, Acanthascus (Rhabdocalyptus) dawsoni dawsoni (Lambe, 1893) is represented 

as Genus (subgenus) species subspecies (Author, year). 

Each species description provides information on 

1) typical growth form(s), 2) surface morphology, including the presence and description of oscula, 3) consistency and texture, 4) known size range or dimensions, 

and 5) color in life and under various preservation 

methods. These characteristics, when used in conjunction with a confirmed photograph, can often provide 

a fairly accurate identification in the field. Definitive 

identification of most species, however, requires careful 

examination of the arrangement of microscopic skeletal 

structures, particularly the types, sizes, and location of 

spicules. Note that the identification of each species 

detailed in this guide has been confirmed by examination of microscopic features. We provide detailed information on the skeletal structure of each species so 

1 Clark, R. N. 2006. Unpubl. manuscript. Field guide to the 

benthic marine invertebrates of Alaska’s shelf and upper slope taken by NOAA/NMFS/AFSC/RACE Division trawl surveys. 302 p.
3

that readers can definitively confirm specimen identification. For most species, we provide photographs of 

specimens on deck (shortly after collection) or archived 

specimens and photographs of specimens in situ. The 

former will be most useful to fisheries observers and 

fishers to identify specimens on fishing and survey vessel 

decks shortly after collection. The latter will be useful 

to scientists attempting to identify and quantify sponge 

fauna in situ, with submersibles and remotely operated cameras. We provide the known zoogeographic 

range and information about the physical habitat and 

oceanographic habitat (i.e., temperature and salinity), 

if available (Koltun, 1959), of each species both within 

Alaskan waters and throughout its known range. Finally, 

we include for each species special remarks with regard 

to taxonomic history, biology, and ecology.

We used the following literature to construct the species lists and determine the zoogeography and other 

published information for each species: Austin (1985), 

Blake and Lissner (1994), Boury-Esnault and Rützler 

(1997), Brøndsted (1993), Burton (1934), Dickinson 

(1945), Hooper and Van Soest (2002), Koltun (1958, 

1959, 1970), Lamb and Hanby (2005), Lambe (1900), 

Laubenfels (1953), Sim and Kim (1988), Van Soest 

et al. (2008), and de Weerdt (1986a, 1986b). 

Appendix I provides a current and comprehensive 

taxonomic list of all sponges now known to occur in the 

deep waters (>80 m) of Alaska. Appendix II provides 

a ranking for all the species included in this guide in 

terms of their importance as fish habitat and vulnerability to disturbance from fishing activities. The average 

score for these two factors is used to prioritize species for 

monitoring as bycatch in commercial fisheries and stock 

assessment surveys. Appendix III includes a glossary of 

terms commonly used in the species descriptions. The 

terminology used in the section on skeletal structure is 

not included in Appendix III. We refer the reader to the 

Thesaurus of Sponge Morphology (Boury-Esnault and 

Rützler, 1997) for a comprehensive glossary of skeletal 

structure terminology. Appendix IV includes scanning 

electron microscopy (SEM) images of spicules from select calcareous, hexactinellid, and demosponges found 

in Alaskan waters. The purpose of Appendix IV is to 

provide readers with a representative collection of spicule images so that they may gain an understanding of 

the terms used in the guide and an appreciation for the 

variation between species and among spicule types. This 

collection also serves as a source of reference material 

for those who wish to microscopically examine sponges.

Zoogeography of sponges

All three major groups of sponges—Class Calcarea 

(calcareous sponges), Class Hexactinellida (hexactinellid or glass sponges), and Class Demospongiae (demosponges)—are well represented in Alaska. Only 12 

(6.1%) of the 196 species of sponges now known from 

Alaskan waters are calcareous sponges. They have skeletons composed entirely of calcium carbonate laid down 

as calcite and are consequently one of the faunal groups 

at high risk from increased acidification of North Pacific 

Ocean waters. They are principally found in shallow 

water and are very rare at depths below 250 m.

Glass sponges are represented by 52 species (26.5% 

of the total) in Alaskan waters. Glass sponges have a siliceous skeleton consisting of spicules with a hexactinal 

or 6-rayed pattern. They are generally more abundant 

in deeper water but have a very broad depth distribution 

in Alaska, ranging from 20 m in the fjords of Southeast 

Alaska to more than 2800 m on the Aleutian Island 

slope. Hexactinellid sponges are common throughout 

Alaska but form different habitats depending on geographical region. In the deeper slope habitats of the 

Aleutian Islands, we have observed large debris fields 

comprised mainly of glass sponge skeletons that provide 

attachment substrate for other invertebrates, including 

gorgonians and hydrocorals. In the Gulf of Alaska, glass 

sponges are generally solitary but do form low-diversity 

dense patches in areas of exposed hard substrate (Stone, 

unpubl. data, 2005). In southern Southeast Alaska, in 

Portland Canal along the border of Alaska and British 

Columbia, we have found small sponge reefs, similar in 

species composition to, but much smaller in size than, 

the massive bioherms reported farther south in British 

Columbia (Conway et al., 1991, 2001, 2005; Krautter et 

al., 2001). Small patches of hexactinellid reef, believed 

to be biohermal, have also been observed in northern 

Southeast Alaska near Juneau, possibly indicating that 

these structures are more common than originally 

hypothesized and may extend up through the inside 

waters of the Alexander Archipelago in areas where 

favorable conditions exist for reef formation.

Demosponges are by far the dominant group of 

sponges in Alaskan waters, with 132 documented species. Demosponges also have a siliceous skeleton but the 

spicules are not hexactinal in pattern and the skeleton 

may be replaced or largely supplemented with an organic collagenous network of spongin. They are found 

throughout Alaskan waters and over a very broad depth 

range (intertidal to more than 2800 m). High diversity 

sponge “gardens” recently discovered in the Aleutian 

Islands (Fig. 1) are dominated by demosponges but 

also include some hexactinellid and calcareous sponges.

A few deep-water sponges (e.g., Guitarra fimbriata,

Halichondria sitiens, and Hymeniacidon assimilis) are cosmopolitan in distribution (Dickinson, 1945), but recent 

research indicates that the occurrence of sibling species 

worldwide might also be common. Deep-water sponges 

are found throughout Alaska and have been reported 
4 Professional Paper NMFS 12 Figure 1 A study area in the central Aleutian Islands showing 15 sites where sponge gardens have been observed. Sponge gardens are areas of high species diversity and abundance and were located from detailed analysis of video footage collected with the Delta submersible in 2002–04 and and ROV Jason II in 2004. Depth contours are in meters.
5

as far north as the Beaufort Sea (de Laubenfels, 1953). 

Sponges inhabit a broad depth range and occur from 

the intertidal zone to the deep-ocean trenches. About 

24% (30 of 125 species) of the sponges known from the 

Aleutian Islands appear to be endemic to the region. 

Deep-water sponges, like deep-water corals, are exceptionally abundant and diverse in the Aleutian 

Islands (Stone, 2006) and not surprisingly, since both 

faunal groups have similar habitat requirements. Both 

groups require stable water currents for feeding and 

other metabolic processes and most require hard, 

exposed substrate for attachment. The geology and 

oceanography of the Aleutian Island Archipelago provide unique conditions to fulfill both requirements. 

The Archipelago contains more than 300 islands and 

extends over 1900 km from the Alaska Peninsula to the 

Kamchatka Peninsula in Russia. The Archipelago is supported by the Aleutian Ridge that forms a semi-porous 

boundary between the deep North Pacific Ocean to the 

south and the shallower Bering Sea in the north. The 

Aleutian Ridge is a volcanic arc, with more than 20 active volcanoes and frequent earthquake activity that was 

formed along zones of convergence between the North 

American Plate and other oceanic plates (Vallier et al., 

1994). The island arc shelf is very narrow in the Aleutian 

Islands and drops precipitously on the Pacific side to 

depths greater than 6000 m in some areas, such as the 

Aleutian Trench. Deep water flowing northward in the 

Pacific Ocean encounters the Aleutian Trench where it 

is forced up onto the Aleutian Ridge and into the Bering 

Sea through the many island passes (Johnson, 2003). 

Additionally, coastal water from the Alaska Stream enters through Unimak Pass in the eastern Aleutians and 

slowly flows northeastward along the Alaska Peninsula. 

The Aleutian North Slope Current flows eastward on 

the north side of the Aleutian Islands towards the inner 

continental shelf of the Bering Sea. This is a swift current and the steep continental slope forces much of the 

flow into the northwest-flowing Bering Slope Current 

(Johnson, 2003).

The collections made by Lambe (1900), de Laubenfels (1953), and Koltun (1958, 1959) provide a firm 

basis to closely examine the zoography of sponges from 

the region. The sponge fauna of the Aleutian Island 

Archipelago has strong taxonomic affinities with the 

sponge fauna of the Sea of Okhotsk (30% of species in 

common), seas of the Arctic Ocean (22% of species in 

common), the eastern (18% of species in common) and 

western Bering Sea (10% of species in common), and 

the Sea of Japan (17% of species in common). By comparison, only 12% of the sponge fauna from the Gulf of 

Alaska is common with that of the Aleutian Islands. This 

zoogeographic pattern has undoubtedly been influenced to some degree by historical sampling effort, but 

the taxonomic affinities of the Aleutian Island sponge 

fauna are clearly much greater with the sponge fauna 

to the west and north than they are with the eastern 

fauna. Burton (1934) suggested that earlier work by 

both Lambe and de Laubenfels indicated that the Arctic 

influence was apparent as far south as Vancouver Island. 

Several species are known from areas immediately adjacent to the Aleutian Island Archipelago and, although 

not yet reported from the region, likely occur there 

based on geographical proximity. These include species 

from the eastern Bering Sea (e.g., Aaptos kanuux) and 

the Commander Islands (Russia) in the western Bering Sea (e.g., Grantia monstruosa, Polymastia laganoides, 

Asbestopluma gracilis, and Axinella hispida).

Biology of sponges

Basic diagrams of sponge morphology, spicule types 

and skeletal structures, and a comprehensive glossary of 

terminology are available in the Thesaurus of Sponge 

Morphology by Boury-Esnault and Rützler (1997) and 

can be accessed via the World Porifera Database (Van 

Soest et al., 2008).2

Sponges are a primitive group of metazoans. They are 

sedentary animals, but a few species (e.g., Craniella spp.) 

may be free-living (unattached) during part of their life 

cycle (Lehnert and Stone, 2011). The dominant feature 

of the typical sponge body plan is the aquiferous system 

through which massive amounts of water are pumped 

(e.g., some large sponges are capable of filtering their 

own volume of water every 20 seconds). Water flow is 

unidirectional and maintained by flagellated cells (choanocytes) that are usually contained within chambers 

where oxygen and food particles are taken up by various 

cell types. Water flows in through inhalent pores or ostia 

and out through one or more larger exhalent openings 

or oscula. The oscula may open into a large cavity called 

an atrium or spongocoel (i.e., the large opening in tubeor vase-shaped sponges).

The sponge body consists of two distinct regions: 

the outer region (ectosome) and the central or inner 

region (choanosome) where the choanocyte chambers 

are located. Each region typically has distinct skeletal 

structures with a diagnostic complement of spicules. 

Spicules are grouped into two main categories: megascleres and microscleres. Megascleres are typically larger 

and provide the primary skeletal support. Microscleres 

are smaller (i.e., a microscope is required to see them) 

and generally function as packing and reinforcing 

2 The database can be found online at http://www.marinespecies.

org/porifera/. Click on “Sources”; type “Boury-Esnault” in “Sourcename”; type “publication” in “Sourcetype”; check the box to

“Limit to sources with full text”; and click on “Search” in lower right 

corner.
6 Professional Paper NMFS 12

structures. Many sponge cells are highly mobile and can 

move freely within the extracellular matrix. Some cells 

are also extremely pluripotent (i.e., capable of differentiating into other cell types) and sponges are capable 

of easily remodeling cell-cell junctions. These features 

probably allow sponges to adapt to diverse and extreme 

habitats and are largely responsible for the extreme phenotypic plasticity displayed by some sponges that makes 

identification from photographs alone so problematic. 

Sponges are generally nonselective filter feeders, 

feeding principally on bacteria, fungi, diatoms, dinoflagellates, and detritus (Bergquist, 1978; Pile et al., 

1996). A recent study, however, has shown that some 

hexactinellid species do exhibit size independent selective filtration of ultraplankton (Yahel et al., 2006). 

Carnivorous sponges were recently discovered that lack 

an aquiferous system altogether and possess structures 

modified to ensnare and capture larger prey such as 

zooplankton (Vacelet and Boury-Esnault, 1995; Watling, 

2007). Carnivorous species in Alaska are deep-water inhabitants and include Cladorhiza corona (Lehnert et al., 

2005), Cladorhiza bathycrinoides, Chondrocladia concrescens, 

and possibly Abestopluma ramosa. 

Few studies have been conducted on the growth rate 

and longevity of sponges, particularly those found in 

deep-water habitats and high-latitude ecosystems. No 

studies have been conducted on the growth of sponges 

in Alaska. In general, the growth rate of temperatewater sponges appears to be seasonal and relatively 

slow, occurring at rates comparable to those of deepwater corals (Ayling, 1983; Thomassen and Riisgård, 

1995; Fallon et al., 2010). Studies on the hexactinellid 

sponge Acanthascus (Rhabdocalyptus) dawsoni in British 

Columbia indicate a growth rate of 1.98 cm/yr and a life 

span of more than 200 years (Leys and Lauzon, 1998). 

Studies on the hexactinellid sponge Aphrocallistes vastus

in British Columbia indicate that it may grow considerably faster (10 cm/yr) but still live in excess of a century 

(Austin et al., 2007). We hypothesize that growth rates 

for sponges in Alaska are similar to those for British 

Columbia sponges but note that growth studies, particularly on demosponges, should be a high research priority in Alaska so that recovery rates from disturbance for 

sponge habitats can be estimated.

Deep-water sponges in the Aleutian Islands appear 

to have few predators. We have observed blood stars 

(Henricia spp.) displaying a typical feeding posture on 

several demosponges (e.g., Artemisina sp., Monanchora 

pulchra, Semisuberites cribrosa, and Haliclona sp.) at shallower depths (80 to 300 m). An earlier interpretation 

of this behavior, however, was that the sea stars were 

simply taking advantage of the sponges’ feeding currents (Anderson, 1960). Henricia were generally accepted as suspension feeders (Anderson, 1960), but 

our additional observation of large numbers of these 

sea stars present on dead sponges and decaying sponge 

fragments in debris “windrows” further implicate predation (or scavenging). In deeper water Hippasteria spp. 

sea stars appear to prey on several species of sponges. In 

the eastern Gulf of Alaska several sea stars (Hippasteria 

spp., Henricia longispina, and Poraniopsis inflata) prey on 

glass sponges (including Acanthascus dawsoni dawsoni, 

A. solidus, Aphrocallistes vastus, and Heterchone calyx) and 

several sea stars (Hippasteria spp., H. longispina, P. inflata, Pteraster tesselatus, and Ceramaster patagonicus) prey 

on demosponges (including Poecillastra tenuilaminaris, 

Halichondria sp., and Mycale loveni). The incidence of 

predation on deep-water sponges in Alaska, however, 

appears to be relatively low and limited to only a few 

species of sea stars. 

Advances are now being made in the study of the 

reproductive biology of sponges, but our current 

knowledge is based on studies of a small fraction of the 

species described to date worldwide; none from Alaska 

(Maldonado and Berquist, 2002). Sponges display 

highly diverse mechanisms of embryogenesis, larval 

differentiation, and reproduction that include both 

sexual and asexual processes (Berquist, 1978; Leys and 

Ereskovsky, 2006; Ereskovsky, 2010). Sexes are either 

temporarily or permanently separate and some species 

are hermaphroditic (Blake and Lissner, 1994). Many 

species are capable of regenerating viable adults from 

fragments, and additional asexual processes include 

the formation of gemmules and reduction bodies, 

budding, and possibly formation of asexual larvae 

(Maldonado and Berquist, 2002). Some species are 

oviparous while others are viviparous and brood larvae. 

Release of propagules (gametes, zygotes, or early embryos) is highly synchronous in oviparous species, but 

asynchronous in viviparous species that release fully 

brooded flagellated larvae (Maldonado and Berquist, 

2002). Several species of Geodiidae are gonochoristic 

and oviparous and this is assumed to be the general 

condition for the family (Cárdenas et al., 2009). Some 

species are viviparous, such as Stylocordyla, which has the 

added peculiarity that larvae are retained alive in the 

body until they have fully developed (Bergquist, 1972). 

Gemmules, reproductive structures that can survive 

adverse conditions such as desiccation or extreme cold, 

are not typically produced by marine sponges. Their 

common occurrence in hermit crab sponges may represent an adaptation to help counter the consequences 

of stranding on shore. 

Many sessile marine fauna, including sponges, have 

evolved the ability to produce or accumulate from associated microorganisms a diversity of unique chemical 

compounds or secondary metabolites that they utilize 

in predator defense, competition for resources, and 

as physiological adaptations to living in extreme environments (Haefner, 2003). More than 12,000 novel 
7

compounds have been isolated from sessile marine 

invertebrates, algae, and microorganisms worldwide 

(Faulkner, 2002). Many of the compounds are currently 

in early clinical or late preclinical development for use 

as treatments for cancer, tuberculosis, HIV, asthma, and 

many other diseases and ailments (Newman and Cragg, 

2004). Deep-water sponges show particular promise in 

this emerging research area, and several species collected from the Aleutian Islands as part of a pilot program 

in 2004 exhibited near 100% inhibition during primary 

screening for M. tuberculosis (Hamann3). Only a handful 

of sponge species from the Aleutian Islands have been 

examined for the presence of secondary metabolites 

(e.g., Na et al., 2010), but so far “hit rates” for biomedically active compounds are on the order of 10%, rather 

than 1% which is typical for samples collected elsewhere 

(Hamann3).

Ecology of sponges

Importance as fish habitat

Deep-water sponges provide important habitat to many 

species of fish and invertebrates, mostly as a source of 

refuge from predation and adverse conditions (e.g., 

strong currents) and as focal sites for foraging on prey 

species that aggregate in sponge habitat. Some fish 

use sponges (e.g., Aphrocallistes vastus and Acanthascus 

dawsoni dawsoni) as spawning substrate (Busby4) and 

others likely use sponge habitat as breeding sites. In 

Alaska, many commercial and non-commercial fisheries 

species are associated with deep-water sponges. Most 

associations are believed to be facultative rather than 

obligatory. Sponges provide important refuge habitat 

for juvenile rockfish (Sebastes spp.) in the Gulf of Alaska 

(Freese and Wing, 2003) and juvenile golden king crabs 

(Lithodes aequispina) in the central Aleutian Islands 

(Stone, 2006). Sponges also contribute to the biodiversity of deep-water habitats by providing spawning substrate for many species that are important trophic links 

to larger consumers (Fiore and Jutte, 2010). 

Deep-water sponges are really no different than 

deep-water corals in the degree to which they provide 

structure; it depends on their maximum size, growth 

form, abundance, intraspecific fine-scale distribution 

(i.e., patch size and density), and interaction with 

other structure-forming invertebrates. In general, 

large arborescent species (e.g., Axinella blanca) and 

those with vase-like or barrel-like morphologies (e.g., 

Acanthascus dawsoni dawsoni and Mycale loveni) provide 

much surface area and consequently refuge space. 

Large sponges such as Mycale loveni have high value as 

fish habitat due to their size (up to 1 m high and wide) 

and abundance (up to 11 sponges per m2

 in the eastern Gulf of Alaska). Adult sharpchin rockfish (Sebastes 

zacentrus) and juvenile Sebastes spp. frequently use the 

cone-shaped sponges as perches in the eastern Gulf 

of Alaska. Smaller sponges at high densities may also 

provide important habitat. For example, finger sponges 

(Axinella rugosa) are small (maximum dimensions only 

2 cm wide by 17 cm high), but are used as perches 

by juvenile rockfish (Sebastes spp.) when present at 

high densities (up to 63 individuals per m2) in steep 

bedrock habitats (Stone, unpubl. data, 2005). Species 

such as Artemisina stipitata have several attributes that 

make them important as fish habitat, including large 

size, tendency to occur in dense patches (contagious 

distribution), and interaction with other emergent 

epifuana. Encrusting species such as Plakina tanaga, 

P. atka, and Bubaris vermiculata have little value as fish 

habitat. The degree to which sponges provide fish 

habitat also depends greatly on their depth distribution 

relative to that of shelter-seeking fishes. In the Aleutian 

Islands, for example, sponges found at depths greater 

than about 1200 m are less important as refuge habitat 

because most of the fish species at those depths appear 

to be less structure-oriented. 

In general, calcareous sponges are not of sufficient 

size or adequately abundant to provide important fish 

habitat. Leucandra tuba may be important for juvenile 

fish that use the easily accessible cavernous interior of 

this species to escape from larger predators. In contrast, 

demosponges are often quite large and form dense 

“gardens” in some areas of the Aleutian Islands (Fig. 1). 

Hexactinellid sponges are extremely important as fish 

habitat. Their skeletons are persistent after death and 

consequently provide the framework for the building 

of reefs or bioherms. Intact skeletons, and to some degree detached and fragmented skeletons, also provide 

attachment substrate for other sedentary structureproviding fauna, including hydrocorals, octocorals, and 

demosponges (Stone, unpubl. data, 2004). 

Vulnerability to disturbance

Bycatch observations in the Aleutian Islands and other 

regions of Alaska clearly support our submersible observations, indicating that sponges are far more abundant 

than corals in deep-water habitats. Sponges, particularly 

demosponges, appear to be the most abundant emergent epifauna in the Aleutian Islands. Approximately 

352 metric tons (t) of sponges are removed from the seafloor by commercial fishing in the Aleutian Islands and 

3 Hamann, M. 2009. Personal commun. Unpubl. data. University of Mississippi, P.O. Box 1848, University, MS 38677.

4 Busby, M. 2010. Personal commun. Alaska Fisheries Science 

Center, NOAA, NMFS, 7600 Sand Point Way N.E., Seattle, WA 

98115.
8 Professional Paper NMFS 12

Bering Sea each year – more than four times the weight 

of corals landed there as fisheries bycatch (NMFS5). 

All emergent epifauna are susceptible to disturbance 

from fishing gear (Stone, 2006) and most species of 

deep-water sponges are to some degree vulnerable to 

disturbance (also see Hogg et al., 2010, for a recent 

review). Fishing operations affect the seafloor from 

depths of 27 m to about 1200 m in the Aleutian Islands, 

with most effort at depths shallower than 200 m (Stone, 

2006). The degree to which a particular gear type affects sponge habitat depends on its configuration (i.e., 

physical area of contact), operation (i.e., physical forces 

on the seafloor), spatial and temporal intensity of operation, seafloor bathymetry and substratum type, and 

the resiliency of sponges to disturbance. Both direct 

and indirect effects from fishing activities on sponges 

likely occur. Direct effects include removal as bycatch, 

damage caused by physical contact, or detachment 

from the seafloor and translocation to unsuitable habitat. Indirect effects include increased vulnerability to 

predation, especially for sponges detached from the 

seafloor, and habitat alteration. Furthermore, there 

is some evidence that reproduction is suppressed in 

damaged shallow-water scleractinian corals due to a 

reallocation of energy reserves to tissue repair and 

regeneration (Wahle, 1983), and similar effects may 

occur in sponges. 

Most of the factors that control the degree to which 

sponges provide habitat structure are the same factors 

that control the degree to which they are vulnerable to 

disturbance. For example, large species with arborescent growth forms are more likely to be contacted by 

passing fishing gear. Species such as Plakina tanaga have 

low vulnerability to disturbance due to their encrusting 

habitus and absence in areas where bottom trawling is 

known to occur. Some species such as Poecillastra tenuilaminaris are highly vulnerable to disturbance due to 

their large size and high relative abundance. Artemisina 

stipitata is moderately vulnerable to disturbance due to 

its large size, erect form, and presence in areas where 

bottom trawling and longline fishing are known to occur. Some sponges, for example those with holdfast 

structures rather than root-like attachment systems, may 

be more vulnerable to detachment, and it is unknown 

if intact but otherwise undamaged sponges detached 

from the substrate (e.g., roller sponges; Reiswig, unpubl. data, 2010) can survive. 

The recovery rate of disturbed benthic ecosystems in 

Alaskan waters is of keen interest to fisheries managers 

but is still little more than conjecture due to limited 

5 National Marine Fisheries Service. 2004. Final programmatic 

supplemental groundfish environmental impact statement for Alaska groundfish fisheries. Alaska Region, NOAA, NMFS, P.O. Box 

21668, Juneau, AK 99802-1668.

knowledge of species distribution and basic life history 

processes. Rates of recovery will depend on growth rates 

and rates of sexual and asexual reproduction or speed of 

regeneration, which are unknown for most species. Two 

key components of ecosystem persistence are resistance 

(i.e., the ability to resist changes from disturbance) and 

resiliency (i.e., the ability to absorb changes and the 

speed with which it returns to its original condition) 

(Holling, 1973). In the case of sponges, resistance to 

disturbance depends largely on their physical size, morphology, and distribution relative to the disturbance. 

We know from studies in the Aleutian Islands and Gulf 

of Alaska that benthic emergent epifauna, especially 

sponges, have low resistance to fishing gear disturbance 

(Freese et al., 1999; Stone, 2006; Heifetz et al., 2009; 

Stone, unpubl. data, 2004–2005). We know little about 

the resiliency of emergent epifauna, however. 

Resiliency depends on several factors, including 

growth rate, recruitment rate, and reproductive ecology. Like deep-water corals, sponges have life history 

attributes such as slow growth rates and high longevity 

(Dayton, 1979), indicating that long periods of time 

would be necessary for habitats dominated by sponges 

to recover from disturbance (Freese, 2001). Growth 

rates for deep-water sponges are generally slow (see 

above) and therefore inherently difficult to study. 

Reference sites, such as undersea cables that provide 

a known time-line for recruitment and subsequent 

growth, show promise in studies of sponge habitat recovery (Levings and McDaniel, 1974) but have not yet 

been identified or targeted for such study in Alaska. 

The only known study to address the recovery dynamics of sponges in Alaska reported no visually detectable 

growth or regeneration one year post-disturbance with 

a bottom trawl (Freese, 2001). Due to the opportunistic 

nature of that study (observations were subjective, not 

quantifiable; not repeatable since specimens were not 

georeferenced; and made in an area subjected to high 

background disturbance), the findings are questioned. 

Contrastingly, a study conducted off the southeastern 

coast of the U.S. indicated that some sponges were 

capable of recovering completely within one year from 

damage caused by a research trawl (Van Dolah et al., 

1987).

Monitoring bycatch of sponges

Appendix II provides a list of all sponges for which we 

have provided species descriptions in this guide. For 

each species, the importance as fish habitat and the vulnerability to disturbance from fishing activities is ranked 

from low (1) to high (3). Ranks for the two measures 

are averaged to provide a score. We recommend that 

species with scores greater than 2.0 be considered a 
9

high priority for monitoring as bycatch in commercial 

fisheries and stock assessment surveys and that species 

lists used to record bycatch in fisheries and stock assessment surveys be appropriately updated to include 

these taxa. Subsequent mapping of these taxa should 

be undertaken in an effort to identify the location of 

vulnerable marine ecosystems. The actual or estimated 

weight of sponge bycatch should also be recorded since 

it is also a valuable indicator of rich sponge habitat.

Collection and preservation 

of sponge specimens

The collection protocols outlined by Etnoyer et al. 

(2006) for deep-water corals are generally appropriate 

for the collection of deep-water sponges. Shortly after 

collection, specimens should be photographed with 

an appropriate scale and the following information 

should be recorded: 1) growth form, 2) general surface 

information (roughness, presence, and description 

of oscula), 3) consistency, 4) size (dimensions), and 

5) color. Color should preferably be recorded immediately after collection since the color of some specimens 

may change shortly after removal from seawater and 

exposure to ambient light. Observations made in situ 

should pay particular attention to the presence and description of oscula, color, type of attachment (i.e., holdfast or root-like system), substrate type, the presence of 

associated fauna, and if collected, whether the specimen 

appears to be whole or only a fragment of the whole 

specimen. The specimen should be photographed or 

recorded on video prior to collection. Collection data 

should be written with pencil on waterproof paper that 

will be packaged with the specimen, and include date, 

location of collection (station number and GPS coordinates), name of collector, cruise number or vessel 

name, depth of collection, and gear type. Sponge specimens may be frozen (preferably at temperatures below 

–10°C) or stored in an 80–90% ethanol solution. Demosponges and hexactinellid sponges (but never calcareous sponges) to be used for histological purposes may 

be fixed in a buffered 5% formaldehyde solution for 24 

hours prior to storage in ethanol solution. Specimens 

to be used for biomedical research or molecular genetic 

analyses should preferably be frozen to –10°C or colder 

unless they will be processed otherwise immediately. 

For museum archiving, specimens should preferably 

be stored in an 80–90% ethanol solution. Very large 

specimens may be dried but a fragment of the specimen 

should be stored in ethanol solution. 

Laboratory identification of sponge specimens

Calcareous sponges and demosponges

For light microscopy and scanning electron microscopy 

(SEM) investigations of the spicules, small fragments of 

sponges (containing both ectosome and choanosome) 

are boiled in nitric acid or hypochlorite, and in several 

steps the spicules are sequentially transferred through 

distilled water to 95% ethanol with the aid of a centrifuge (Rützler, 1978; Lehnert and Van Soest, 1998). For 

light microscopy, small drops of the spicule-ethanol 

suspension are transferred onto glass slides and, after 

evaporation of the ethanol, embedded in Canada balsam. Semi-thin sections obtained with a razor blade are 

also embedded in balsam. For SEM studies, the spiculeethanol suspension is transferred directly onto stubs 

and coated with gold-palladium after ethanol evaporation (Lehnert and Van Soest, 1998). Finally, data of 

spicule types and their dimensions are compared to the 

literature descriptions to assign the specimens to their 

proper genus and species. 

Hexactinellid sponges

Individual specimens are first triaged to assign them 

to order and family by making temporary microscope 

preparations of surface spicules with commercial bleach. 

Specimens are then processed by family groups. Permanent picks or peels of surface spicules are mounted in 

Canada balsam and pieces (about 1 cm diameter) of 

body wall are digested in hot nitric acid to obtain a suspension of cleaned spicules. Smaller spicules are filtered 

onto 0.2 μm pore-size membrane filters and mounted 

in Canada balsam for microscopic examination. Very 

large spicules are picked from the suspension with 

forceps, cleaned, and mounted in balsam. Remaining 

medium-size spicules are washed, spread on slides, and 

mounted in balsam. Samples of individual spicule types 

are photographed by digital camera and, when necessary, dimensions (e.g., length, diameter) are measured 

by digitizer-interfaced microscopy. Data of spicule types 

and their dimensions are finally compared to the literature descriptions to assign the specimens to their proper 

genus and species.

11

CLASS CALCAREA
12 Professional Paper NMFS 12

Description. Conical tubes have thin walls laid in 

folds. Viewed with a hand-lens, the walls consist of a 

mesh of tissue with many open inhalant pores, round 

to oval, 300–600 μm in diameter. The osculum at the 

top of the cone has no spicule crown. Consistency is 

soft and elastic. Height of cones is to 3–5 cm; several 

basally fused cones may cover larger areas. Color in life 

is creamy white to pale yellow.

Skeletal structure. Triactines are arranged tangentially; large triactines (50–150 × 7–10 μm/ray), small 

triactines (20–30 × 3–6 μm/ray).

Zoogeographic distribution. Cosmopolitan. In Alaska 

– central Aleutian Islands. Elsewhere – the genus Clathrina presently contains 87 nominal species and has a 

worldwide distribution.

Habitat. In Alaska – sometimes grows around and partially encrusts the gorgonian Muriceides nigra at depths 

near 119 m.

Remarks. The genus Clathrina was erected by Gray 

(1867) and is in desperate need of revision at the species level.

Photos. Fragment of Clathrina sp. collected in the central Aleutian Islands at a depth of 119 m. Grid marks are 

1 cm2

. 2) The same specimen as in photo 1 (center) in 

situ encrusting the gorgonian Muriceides nigra.

1. Clathrina sp.
13

Description. Sponge is thickly encrusting. The surface 

is smooth to the unaided eye but rough to the touch due 

to large protruding spicules. Consistency is stiff, slightly 

compressible, and elastic. Individuals may reach a size 

of approximately 6 × 2 × 1 cm. Color in life is white to 

creamy white.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. There are large triactines (50–170 × 

5–15 μm/ray) and occasionally tetractines of the same 

dimensions. Small oxeas (60–85 × 3–6 μm) have un-

equal ends, most with a characteristic bend near one 

end. Small oxeas are only at the surface of inner and 

outer walls, equiangular and sagittal triactines are inbetween. Outer surfaces are covered by a mesh of small 

diactines, mesh-size approximately 25 μm in diameter; 

tracts form the mesh 20–40 μm in diameter. 

Zoogeographic distribution. Rare. In Alaska – central 

Aleutian Islands. Elsewhere – previously known only 

from the Sea of Japan.

Habitat. Grows around and encrusts Stylaster hydro-

corals at depths near 175 m. Species of Craniella may be 

attached to it (see photo 1).

Photos. 1) Whole specimen collected at a depth of 

175 m in the central Aleutian Islands. Grid marks are 1 

cm

2

. 2) The same specimen as in photo 1 in situ encrust-

ing a Stylaster hydrocoral. 

2. Leucandra poculiformis Hozawa, 1918 
14 Professional Paper NMFS 12

Description. Sponge has thin-walled (0.5 mm) tubes. 

Up to 17 individual tubes surround large internal 

areas. Surface is smooth to the unaided eye, irregularly 

undulating (i.e., wavy or sinuous) oxeas, and rough to 

the touch due to large protruding spicules. Walls are 

compressible and only slightly elastic. Diameter is to 30 

cm. Color in life is light grey to creamy white. 

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Walls consist of giant tri- and tetractines 

(200–1200 × 20–100 μm/ray), one ray usually shorter 

than the others. Irregularly distributed and undulating oxeas (95–140 × 3–5 μm) are very abundant and 

are scattered in between the giant tri- and tetractines 

without obvious orientation. Ectosomal areas with inhal-

ant pores have an outer layer of special spicules called 

pugioles (18–35 μm long) which form meshes, leaving 

open pores of about 50 μm diameter, with pugiole-tracts 

of approximately 25 μm in diameter. The mesh of the 

pugioles is underlain by a mesh of microdiactines of the 

same dimensions. 

Zoogeographic distribution. Locally common. In 

Alaska – Aleutian Islands and Bering Sea canyons. Else-

where – previously known only from the Sea of Japan.

Habitat. In the Aleutians Islands – depths between 

112 and 250 m on exposed bedrock, boulders, cobbles, 

and pebbles in generally low-relief (i.e., flat-bottomed) 

habitats. In the Bering Sea – known from Pribilof Can-

yon at depths between 203 to 220 m on pebbles and 

cobbles. Elsewhere – the holotype was described from 

the Sea of Japan near Okinoshima and collected at a 

depth of 106 m.

Photos. 1) Whole specimen collected at a depth of 

145 m in the central Aleutian Islands. Grid marks are 

1 cm

2

. 2) The same specimen as in photo 1 in situ. 

3) Specimen (center, top) at a depth of 120 m in the 

central Aleutian Islands.

3. Leucandra tuba Hozawa, 1918 
15

CLASS HEXACTINELLIDA
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4. Farrea kurilensis ssp. nov. Reiswig and Stone, in preparation 

Description. Lace-like mass has large-caliber (2-cm 

diameter), short, branching, and anastomosing tubes 

increasing in size (of the mass) by lateral growth, conforming, and attaching at multiple sites to the hard 

substrate and not growing erect. Surface of the mass 

is labyrinthic, but that of individual tube elements is 

smooth. Open terminal ends of tubes are effectively 

oscula. Consistency is flexible but brittle due to fusion 

of the very thin primary skeleton. Individuals observed 

in situ have a diameter of 170 cm. Color in life is translucent light blue to white.

Skeletal structure. The primary framework is a fused 

farreoid lattice of hexactins forming a network of square 

meshes with sides 439–750 μm long; it is continuous 

throughout the specimen. Loose megascleres include 

dermal and atrial pentactins, often with knobs or very 

short distal rays (170–375 μm tangential ray length, 

211–422 μm proximal ray length); very large anchorate clavules with completely smooth shafts (302–1243 

μm length); moderate-sized uncinates (1.03–2.99 mm 

length); very rare choanosomal hexactins (208–239 

μm ray length). Microscleres consist of two size classes 

of stellate discohexasters (30–89 μm and 91–293 μm in 

diameter).

Zoogeographic distribution. Locally common and 

abundant in some areas. In Alaska – central Aleutian 

Islands. Elsewhere – not reported.

Habitat. Attached to bedrock, mudstone, boulders, 

cobbles, and hexactinellid sponge skeletons at depths 

between 300 and 2249 m. 

Remarks. Farrea kurilensis presently has two subspecies: F. k. kurilensis, which occurs near the Kuril Islands 

and in the Sea of Okhotsk, has smooth pileate clavules, 

while F. k. beringiana, which also occurs near the Kuril 

Islands, has all clavules with spines. The new Aleutian 

subspecies differs from these in having anchorate clavules with smooth shafts and an alternate microsclere 

combination.

Photos. 1) Preserved (frozen then dried) fragment of 

a specimen collected at a depth of 2105 m in the central 

Aleutian Islands. Grid marks are 1 cm2

. 2) The same 

specimen as in photo 1 in situ. 
17

Description. Lace-like hemispheric mass has largecaliber (1.5 to 2-cm diameter), short, branching and 

anastomosing tubes increasing in size (of the mass) 

by lateral growth, attached to hard substrate at central 

origin and at multiple lateral contact sites. Surface of 

the mass is labyrinthic with open ends of tubes func-

tioning as oscula, but external and internal surfaces 

of individual tube elements are smooth. Consistency is 

flexible but brittle and fragile due to breakage of the 

very thin fused primary skeleton. An individual mass 

from a single settled larva may reach 3 m in diameter 

and over 1 m in height. Color in life is translucent white 

to pale yellow.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The primary framework is a fused farreoid lattice of hexactins forming a network of square 

meshes with sides 79–494 μm long; it is continuous 

throughout the specimen. Loose megascleres include 

dermal and atrial pentactins (190–305 μm tangential ray 

length, 78–280 μm proximal ray length); pileate and an-

chorate clavules usually with smooth shafts, rarely with 

one or two large spines (196–308 μm length); moderatesized uncinates (0.89–2.64 mm length). Microscleres 

are mainly oxyhexasters with relatively long primary 

rays (57–111 μm in diameter). There may be a very few 

discohexasters or onychohexasters. 

Zoogeographic distribution. Locally common and 

abundant in some areas. In Alaska – eastern Gulf of 

Alaska. Elsewhere – cosmopolitan, but the previous 

known northern limit of the subspecies was 55.4°N in 

northern British Columbia; the new Alaska collections 

extend the northern limit to 58.2°N.

Habitat. In Alaska – attached to bedrock, boulder, and 

cobbles at depths between 91 and 238 m. Elsewhere 

– attached to cobbles, bedrock, shell, coral skeletons, 

and hexactinellid sponge skeletons at depths between 

86 and 1360 m.

Remarks. This may be the largest species of sponge in 

the North Pacific Ocean, with a diameter reaching 3 m. 

This sponge provides much refuge habitat for small fish 

and micro-invertebrates that may be forage species for 

larger fish and crabs. 

Photos. 1) Fragment of a specimen collected at a 

depth of 168 m in the eastern Gulf of Alaska. Grid marks 

are 1 cm

2

. 2) Specimen at a depth of 165 m in the eastern Gulf of Alaska. 

5. Farrea occa occa Bowerbank, 1862
18 Professional Paper NMFS 12

Description. Erect bush of small-caliber (8-mm diameter), relatively long, branching tubes increasing in 

size (of the entire specimen, not the tube elements) by 

terminal growth of the constituent tubes; attached to 

hard substrate at a small original base. Surface of the 

mass consists of the terminal tube apertures as oscula, 

but both inner and outer surfaces of individual tube 

elements are smooth. Consistency is stiff and brittle 

due to fusion of the thin primary skeleton. Size of the 

original specimen from which the fragments were collected was not recorded, but it must have been at least 

10 cm tall. Color in life is white; preserved fragments 

are light brown.

Skeletal structure. The primary framework is a fused 

farreoid lattice of hexactins forming a network of rectangular meshes with longitudinal sides 269–720 μm 

long and lateral sides 534–778 μm long; it is continuous 

throughout the specimen. Loose megascleres include 

dermal and atrial pentactins, often with knobs as rudiments of the sixth distal rays (159–361 μm tangential 

ray length, 206–452 μm proximal ray length); pileate 

clavules without shaft thorns (333–490 μm length); 

moderate-size uncinates (1.02–2.98 mm length). Microscleres include oxyhexasters and hemioxyhexasters 

(62–117 μm in diameter); oxyhexactins (83–130 μm 

in diameter); discohexasters and hemidiscohexasters 

(48–76 μm in diameter); discohexactins (58–98 μm in 

diameter).

Zoogeographic distribution. In Alaska – a rare species, known only from a few locations in southern 

Amchitka Pass and near Bobrof Island in the central 

Aleutian Islands. Elsewhere – not reported.

Habitat. In Alaska – attached to small boulders, 

cobbles, and pebbles at depths between 529 and 905 m.

Remarks. Our Aleutian specimen was compared with 

the four other species of Farrea that lack anchorate 

clavules. It differs from each of them in its microsclere 

complement. Its description as a new species of Farrea

is now in progress.

Photos. 1) Preserved (frozen then dried) fragments of 

a specimen collected at 887 m in the central Aleutian 

Islands. 2) The same specimen as in photo 1 (indicated 

by the white arrow) in situ. The separation between the 

red laser marks is 10 cm.

6. Farrea sp. nov. Reiswig and Stone, in preparation
19

Description. Tall, circular, rigid funnel is attached 

to hard substrate, with small hollow digitate processes 

projecting from all lateral surfaces. In life the lateral 

processes are closed distally by tissues and loose spicules, 

but the fused skeletal framework is open at the ends. 

The single large circular osculum at the distal end has a 

crenulate margin. All surfaces are smooth to the naked 

eye, but under low magnification they are seen to be 

ornamented with fine hairs of projecting uncinate spicules. Consistency is hard and rigid. Height is to 29.2 cm 

and diameter to 13.7 cm; lateral processes 6–12 mm in 

diameter and up to 24 mm long are distributed rather 

evenly without pattern and commonly divide into two 

or three short branches in larger specimens. Color in 

life is yellow-orange to light orange.

Skeletal structure. The primary rigid skeleton of 

fused hexactine spicules has elongate meshes with 

longitudinal strands and radial septa, similar to the 

primary framework of Chonelasma. That framework 

curves smoothly out into and continues through the 

digitate processes. Slight indications of irregular dermal and atrial cortices are present, but they are usually 

only one dictyonalium in thickness. There are several 

types of loose megascleres: pinular hexactins on inner 

and outer surfaces of wall and processes, with thorned 

pinulus (67–287 μm long); tangential rays spined at the 

tips (88–196 μm long); proximal ray (79–954 μm long); 

simple hexactins and pentactins in subatrial position of 

wall and processes (106–518 μm ray length); tauactins 

only in spicule pads at tips of processes (158–479 μm ray 

length); diactins in atrial surface of wall and processes 

(88–196 μm ray length); discoscopules on both surfaces 

of wall and processes (234–480 μm long); tyloscopules 

only echinating atrial surface of processes (619–952 

μm long); uncinates echinating both surfaces of wall 

and processes (656–2785 μm long). Microscleres are 

mainly discohexactins (76%; 52–98 μm in diameter), 

moderately common oxyhexactins (20%; 34–94 μm 

in diameter), and few hemidiscohexasters (4%; 46–

100 μm in diameter).

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

7. Family Euretidae; Genus nov., sp. nov. Reiswig and Stone, in preparation

Habitat. Occurs singly or in small patches on exposed 

bedrock, mudstone, boulders, and cobbles at depths 

between 773 and 2084 m. 

Remarks. Juvenile Verrill’s Paralomis crabs (Paralomis 

verrilli) use the spongocoel as refuge habitat (Stone, 

unpubl. data, 2004).

Photos. 1) Whole specimen collected at a depth of 

1256 m in the central Aleutian Islands. Grid marks are 

1 cm2. 2) Same specimen as in photo 1 in situ.
20 Professional Paper NMFS 12

Description. This sponge has a thin-walled flaring 

funnel or trumpet that attaches to hard substrate by 

a short narrow stalk ending in a small basal disc. The 

outer dermal surface is smooth with a thin transparent 

spicule lattice covering a system of radiating ridges (average 1.1 mm wide) and grooves (average 0.8 mm wide) 

visible to the naked eye. The internal atrial surface, also 

smooth, is thoroughly and evenly pocked by openings 

of small exhalant canals (average 1.5 mm diameter). 

Under low magnification all surfaces appear furry due 

to projecting ends of uncinate and scopule spicules. 

Consistency is stony hard but crumbly and quite brittle. 

Only a single partial specimen was collected and measured 25.4 cm tall by 20.6 cm wide at the margin; wall 

thickness was 5–6 mm. Color in life is white; specimens 

dried or preserved in ethanol are brownish orange.

Skeletal structure. The main skeleton is a rigid framework of fused hexactins with grooves and septa typical of 

the genus. Loose megascleres include rough pentactins 

to hexactins with short distal ray (19–164 μm long), tapered tangential rays (106–473 μm long), and proximal 

rays (94–896 μm long); small rough regular hexactins 

(84–177 μm long rays); scopules with rounded tine tips 

(294–965 μm total length); uncinates (437–1480 μm 

long). Microscleres are mostly oxyhexasters and hemioxyhexasters (93%) with 1–4 nearly smooth, robust terminal rays (53–73 μm diameter); stellate discohexasters 

(7%) with 4–10 finely rough terminal rays (50–75 μm 

diameter); a very few oxyhexactins (>1%) similar in size 

to oxyhexasters.

Zoogeographic distribution. Locally abundant. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Occurs singly on bedrock, mudstone, boulders, and cobbles at depths between 704 and 1264 m. 

Photos. 1) Preserved (frozen) specimen collected at 

a depth of 866 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 in situ. 

The separation between the red laser marks is 10 cm.

8. Tretodictyum sp. nov. Reiswig and Stone, in preparation 
21

Description. Basic form is a hollow, thin-walled cone, 

but larger, older specimens add lateral mitten-like outgrowths becoming highly variable; overall it is similar to 

Heterochone calyx with which it is often confused. Surface 

is smooth, usually with a single large terminal osculum. 

Consistency is rigid and brittle. Size is up to 2 m high 

and 3 m laterally. Color in life varies from white to light 

yellow and orange.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Rigid skeleton of fused hexactinal spicules has distinctive honeycomb pattern of 1 mm wide 

channels. Several types of loose megascleres include 

dermal pinular hexactins with pinulus (109–185 μm 

long), tangential rays (38–142 μm long), proximal ray 

(53–232 μm long); small hexactins (85–201 μm/ray); 

scopules on both surfaces (262–497 μm long); atrial 

spiny diactins (262–722 μm long); uncinates (890–3440 

μm long). Microscleres are oxyhexasters (126–252 μm 

in diameter) and two types of discohexasters (16–25 and 

30–56 μm in diameter). 

Zoogeographic distribution. North Pacific Ocean; 

locally common and abundant in some areas. Alaska – 

Bering Sea to Southeast Alaska. Elsewhere – Japan to 

Baja Mexico. 

Habitat. Aleutian Islands – attached to cobbles and 

pebbles (low-relief habitat) and sometimes bedrock 

(high-relief habitat) at depths between about 100 and 

756 m. Bering Sea – attached to cobbles and pebbles at 

depths between 373 and 522 m. Gulf of Alaska – principally on bedrock, but also boulders and cobbles on the 

continental shelf and upper slope at depths between 

140 and at least 228 m. Observed in the glacial fiords 

9. Aphrocallistes vastus Schulze, 1886

of Southeast Alaska growing on bedrock at depths as 

shallow as 20 m. Elsewhere – Lamb and Hanby (2005) 

provide a worldwide depth range of 10 and 1600 m.

Remarks. This is one of the most ecologically important sponges in Alaska and perhaps the most thoroughly 

studied sponge species in the North Pacific Ocean. Due 

to its rigid skeleton, this species is an important structural component of the sponge reefs reported along 

the Pacific Coast of Canada (Conway et al., 1991, 2005; 

Krautter et al., 2001) and more recently in southern 

Southeast Alaska. Juvenile golden king crabs (Lithodes 

aequispina) use the spongocoel as refuge habitat in 

the Aleutian Islands (Stone, 2006) and in the Bering 

Sea Canyons (Stone, unpubl. data, 2007). The bigmouth sculpin (Hemitripterus bolini) deposits its eggs in 

the spongocoel in the Bering Sea and Gulf of Alaska 

(Busby3

). This species is preyed upon by the sea stars 

Hippasteria spp. and Poraniopsis inflata in the eastern 

Gulf of Alaska. Aphrocallistes vastus can be distinguished 

from the very similar Heterochone calyx calyx by the lack 

of pinular hexactins on the inner (atrial) surface and by 

the possession of very robust oxyhexasters with primary 

rays subsumed in the swollen centrum.

Photos. 1) Specimen collected at a depth of 168 m 

in the eastern Gulf of Alaska. Grid marks are 1 cm2. 2) 

Specimen with a juvenile golden king crab (Lithodes 

aequispina) in pre-molt condition at a depth of 190 

m in the central Aleutian Islands. 3) Specimen with a 

swarm of euphasiids at a depth of 170 m in the eastern 

Gulf of Alaska. Photo by J. Lincoln Freese (AFSC). 4)

Specimen at a depth of 190 m in the central Aleutian 

Islands. 5) Specimen at a depth of 20 m with Eualas sp. 

shrimp in Glacier Bay National Park, Southeast Alaska. 
22 Professional Paper NMFS 12

9. Aphrocallistes vastus Schulze, 1886 (continued)
23

Description. This sponge is polymorphic and similar 

to Aphrocallistes vastus, with which it is often confused. 

Sponge is cup or funnel shaped, and is up to at least 30 

cm in height and 40 cm in diameter. It forms bowls or 

plates in areas of low current. Lateral walls are often with 

hollow finger-shaped processes. Color is bright gold or 

pale yellow in the Aleutian Islands and Bering Sea. It 

displays two typical color morphs in the Gulf of Alaska 

– white and golden yellow.

Skeletal structure. It has a rigid skeleton of fused 

hexactinal spicules with a poorly delineated honeycomb pattern of 1 mm wide channels passing vertically 

through the walls. Several types of loose megascleres 

include pinular hexactins on inner and outer surfaces, 

with thorned pinulus (48–150 μm long), tangential rays 

spined at the tips (100–302 μm long), and proximal ray 

(69–1265 μm long); scopules on both surfaces (242–630 

μm long); spined hexactins (80–204 μm/ray); uncinates 

(500–1540 × 14–54 μm); rough centrotylote diactins 

(308–786 μm/ray) are apparently absent in some specimens. Microscleres are discohexactins and discohexasters (44–100 μm in diameter) with 1–4 secondaries and 

terminal discs; oxyhexactins and oxyhexasters (53–100 

μm in diameter).

Zoogeographic distribution. North Pacific Ocean; locally common and abundant in some areas. In Alaska – 

Bering Sea to Southeast Alaska. Elsewhere – Japan, Kuril 

Islands, Sea of Okhotsk, British Columbia to Panama.

Habitat. Aleutian Islands – attached to bedrock, 

cobbles, and pebbles, usually in low-relief habitats, at 

depths between 112 to 740 m. Bering Sea – attached to 

pebbles and hexactinellid skeletons at depths between 

375 and 522 m. Gulf of Alaska – on the continental shelf 

and upper slope at depths between 70 and at least 259 

m. Observed in the glacial fjords of Southeast Alaska 

growing on bedrock at depths as shallow as 21 m. Elsewhere – reported at depths between 23 and 1103 m.

Remarks. This is one of the most ecologically important sponges in Alaska. Juvenile golden king crabs 

(Lithodes aequispina) use the spongocoel as refuge habitat in the Aleutian Islands (Stone, 2006). Due to its rigid 

skeleton, this species is an important structural component of the sponge reefs reported along the Pacific 

coast of Canada (Conway et al., 1991, 2005; Krautter et 

al., 2001) and recently in southern Southeast Alaska. 

We have not been able to confirm the presence of the 

other subspecies (H. calyx schulzei) in Alaskan waters. 

This species may be preyed upon by the sea star Henricia 

longispina in the eastern Gulf of Alaska. Heterochone calyx 

calyx can be distinguished from the very similar Aphrocallistes vastus by the presence of pinular hexactins on the 

inner (atrial) surface and by the lack of robust oxyhexasters with primary rays subsumed in a swollen centrum.

Photos. 1) Specimen collected at a depth of 172 

m in the central Aleutian Islands. A small stalked demosponge grows from inside the specimen. Grid marks 

are 1 cm2

. 2) Specimen collected at a depth of 520 m 

in Zhemchug Canyon, Bering Sea. Grid marks are 1 

cm2. 3) Specimen at a depth of 180 m with a juvenile 

rosethorn rockfish (Sebastes helvomaculatus) and juvenile 

brown box crab (Lopholithodes foraminatus) in the eastern Gulf of Alaska. 4) Specimen at a depth of 181 m in 

the eastern Gulf of Alaska. 5) Specimen at a depth of 

190 m with a sharpchin rockfish (Sebastes zacentrus) and 

squat lobsters (Munida quadrispina) in the eastern Gulf 

of Alaska. Photo by J. Lincoln Freese (AFSC).

10. Heterochone calyx calyx Schulze, 1886
24 Professional Paper NMFS 12

10. Heterochone calyx calyx Schulze, 1886 (continued)
25

Description. This tube or sac sponge attaches basally 

to hard substrate. Lateral walls bear distinctive smooth 

depressions, each with a small (1–3 mm diameter) central hole (parietal osculum) connecting to the atrial cavity; edges around depressions often project several millimeters as parietal ledges; the large terminal osculum 

is normally covered by a coarse sieve plate and bordered 

by a flaring marginal cuff. Consistency is slightly compressible and rubbery. The partially collected Alaskan 

specimen lacked an oscular sieve plate, marginal cuff, 

and parietal ledges. Its atrial cavity was subdivided by 

longitudinal wall fusions. It is estimated to have been 25 

cm long by 13 cm diameter, but elsewhere incomplete 

specimens 48 cm long have been reported. Color in life 

is white; drab when dried or preserved in ethanol.

Skeletal structure. While the Aleutian specimen may 

have unusual morphology, its spiculation is typical for 

the species. Its skeleton consists mainly of loose spicules; 

spicule fusion is present only in the lower parts collected 

and is assumed to have been extensive in the basal part 

left on the attachment site. Megascleres include thick 

principal diactins with rounded tips (5–16 mm long); 

thin diactins with pointed tips (1.3–7.5 mm long); 

sword-shaped dermal hexactins with short, tapered, 

pointed distal rays (89–369 μm length), tapered and 

pointed tangential rays (156–318 μm length), and long 

tapered proximal rays (232–807 μm length); regular 

hexactins (134–292 μm ray length); atrial pentactins 

with sharp tapered tangential rays (122–295 μm ray 

length) and longer proximal rays (137–563 μm ray 

length); a few atrial triactins and stauractins of similar 

shape and size; short, thick atrial diactins (347–1047 μm 

length); small diactins around parietal oscula (42–197 

μm length). Microscleres include mainly oxystaurasters 

(61–107 μm diameter); a few oxyhexasters (70–129 μm 

diameter); dermal floricomes (78–113 μm diameter) 

with 10–14 terminal rays ending in heads with 2–3 teeth; 

remnants of graphiocomes are common, with central 

(25–43 μm diameter) and terminal rays as dispersed 

thin raphides (141–237 μm long).

Zoogeographic distribution. Widespread but uncommon. In Alaska – central Aleutian Islands. Elsewhere 

– reported from Indo-West Pacific Region including 

from near India, Northwest and South Australia, New 

Zealand, New Caledonia, Indonesia, Japan.

11. Regadrella okinoseana Ijima, 1896

Habitat. In Alaska – occurs singly on mudstone, bedrock, and possibly hexactinellid skeletons at depths between 1071 and 1395 m. Elsewhere – reported at depths 

between 390 and 1264 m.

Remarks. The new record of this species in the Aleutian Islands represents a range extension of over 3700 

km from the nearest previous known site near Japan.

Photos. 1) Fragment of preserved (frozen) specimen 

collected at a depth of 1386 m in the central Aleutian 

Islands. Grid marks are 1 cm2. 2) Same specimen as in 

photo 1 in situ. The separation between the red laser 

marks is 10 cm.
26 Professional Paper NMFS 12

Description. This soft barrel sponge, attached basally, 

has very large surface conules lacking prostal spicules. 

Occasionally two tubes are attached near the base. The 

outer surface is smooth, its large canals covered by a 

very delicate spicule lattice. The large atrial canals are 

uncovered and the atrial surface is reflected out of the 

large terminal osculum; there are no marginal spicules. 

Height and diameter is to 25 cm. Color in life is white 

to creamy white.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are thin diactins with rough 

tips (0.94–19.8 mm); dermal finely rough pentactins 

with cylindric rays ending in rounded tips (113–271 

μm ray length); similar but hexactine atrial spicules 

(125–239 μm ray length). Microscleres are discoctasters 

(119–234 μm diameter), each primary ray bearing 5–10 

slightly curved terminals; oxyhexactins, hemioxyhexasters, and oxyhexasters (98–199 μm diameter). Microdiscohexasters are absent.

Zoogeographic distribution. In Alaska – central Aleutian Islands. Known with certainty only from the single 

specimen collected southwest of Adak Island, but analysis of video records indicate that it is a relatively rare 

species in this area. Elsewhere – not reported.

Habitat. Attached to bedrock, mudstone, and large 

boulders at depths between 1446 and 2245 m.

Remarks. This specimen is in the process of being 

described so data are incomplete; it differs from all 

named species of the genus in several characters, and 

from A. platei in particular, in lack of both prostalia and 

microdiscohexasters, and lack of loose spicule lattice 

over atrial canal apertures.

12. Acanthascus (Acanthascus) profundum ssp. nov. Reiswig and Stone, in preparation

Photos. 1) Fragment of preserved (frozen) specimen 

collected at a depth of 2105 m in the central Aleutian 

Islands. Grid marks are 1 cm2. 2) Same specimen as in 

photo 1 in situ growing just behind Farrea kurilenis ssp. 

nov. 
27

Description. Soft straight or curved tube or barrellike sac, occasionally partially divided into two or three 

conjoined tubes, is attached basally to hard substrate. 

Surface is smooth but usually bearing a 1-cm tall veil of 

pentactins and diactins which may be clean but usually 

covered with small epizoans and sediment; large single 

terminal osculum has marginal fringe. Consistency is 

soft and compressible. Height is to 1 m and diameter to 

30 cm. Color in life is white but often coated with sediment, epizoic organisms, and flocculent material, giving 

it a brown to greenish brown appearance.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are thick prostal diactins to 

6 cm long; hypodermal pentactins with some smooth 

and some thorned tangential rays (0.9–4.1 mm long) 

and smooth proximal rays (0.9–4.1 mm long); principal diactins (4.2–11.4 mm long); thin short diactins 

(0.35–1.62 mm long); dermal stauractins (50–101 mm 

ray length), dermal pentactins with tangential rays 

(54–99 mm long) and proximal rays (44–100 mm long); 

atrial hexactins (55–177 mm ray length). Microscleres 

are oxyhemihexasters (48–104 mm in diameter); small 

discoctasters with straight terminals (56–82 mm in diameter); microdiscohexasters (17–28 mm in diameter).

Zoogeographic distribution. Locally common and 

abundant. In Alaska – eastern Gulf of Alaska. Elsewhere 

– British Columbia to southern California.

Habitat. In Alaska – attached to bedrock, cobbles, and 

pebbles at depths between 82 and 255 m. Elsewhere 

– attached to bedrock, cobbles, and pebbles on flat, 

inclined, or vertical surfaces (e.g., fjord walls), and following detachment may survive loose as roller sponges 

(Reiswig, unpubl. data, 2010) at depths between 10 and 

437 m.

Remarks. The species presently contains three subspecies: A. (R.) d. dawsoni (reviewed here); A. (R.) d. alascensis Wilson and Penney, 1930 (known only from Cape 

Spencer, Gulf of Alaska); and A. (R.) d. horridus Koltun, 

1967 (from the Bering Sea). A. (R.) d. dawsoni can be 

distinguished from A. solidus by the presence of at least 

some heavily thorned hypodermal pentactins; the latter 

species possesses only smooth hypodermal pentactins. 

13. Acanthascus (Rhabdocalyptus) dawsoni dawsoni (Lambe, 1893)

A. (R.) d. dawsoni is similar in appearance to A. mirabilis but likely differs in depth distribution and can be 

distinguished by its much smaller discoctasters (56–82 

vs. 144–180 μm diameter). We have observed a very 

similar species in the central Aleutian Islands at depths 

between about 400 and 1238 m attached to cobbles and 

hexactinellid skeletons. This species is preyed upon by 

the sea star Poraniopsis inflata and possibly by the sea star 

Henricia longispina in the eastern Gulf of Alaska. 

Photos. 1) Whole specimen collected at a depth of 

168 m in the eastern Gulf of Alaska. Grid marks are 1 

cm2. 2) Specimen (indicated by the white arrow) at a 

depth of 165 m in the eastern Gulf of Alaska. 
28 Professional Paper NMFS 12

Description. This species was known previously only 

from the holotype, which was the distal portion (about 

1/3 of total specimen) of a soft saccate sponge. Surface is overtly smooth but evenly covered with small 

conic protuberances and bears a veil of pentactins and 

diactins; large single terminal osculum with marginal 

fringe. Consistency is soft and compressible. From the 

holotype fragment, the original specimen was thought 

to be ca. 30 cm in length and 15 cm in diameter with a 

1-cm thick wall; a new Aleutian specimen is 15.5 cm in 

length and 10.1 cm in diameter. Color in life is white; 

light brown when preserved in ethanol.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are thick prostal diactins to 

2 cm long; hypodermal pentactins with mostly thorned 

paratropal tangential rays to 1 cm long; principal diactins (11.7–21.0 mm long); dermal diactins (plus a few 

stauractins and hexactins) (204–462 mm long); atrial 

hexactins (free ray 249–447 μm long, tangentials and 

parenchymal rays 153–316 mm long). Microscleres are 

oxyhexactins and a few oxyhexasters and hemioxyhexasters (72–232 mm in diameter with 33–87 mm long 

terminal rays); discoctasters (144–180 mm in diameter) with very short primary rays (20–28 mm long), 

each bearing 8–12 straight or s-curved, divergent terminal rays; microdiscohexasters mainly near or in the 

dermal surface (21–40 mm in diameter).

Zoogeographic distribution. Rare. Known only from 

Alaska – central Aleutian Islands and Gulf of Alaska. 

Elsewhere – not reported.

Habitat. Central Aleutian Islands – occurs singly on 

bedrock at depths between 1984 and 2790 m. Southern 

Alaska – attached to cobbles at depths near 1143 m.

Remarks. Our specimen recently collected in the 

Aleutian Islands is only the second known verified 

specimen of this species. It is very likely that A. (R.)

unguiculatus Ijima, 1904 is a junior synonym of A. (R.)

mirabilis. This species is similar in appearance to A.

dawsoni but likely differs in depth distribution and 

can be distinguished by its much larger discoctasters 

(144–180 vs 56–82 μm diameter). 

Photos. 1) Fragment of preserved (frozen) specimen 

collected at a depth of 2311 m in the central Aleutian 

Islands. Grid marks are 1 cm2

. 2) Same specimen as in 

photo 1 in situ. The separation between the red laser 

marks is 10 cm.

14. Acanthascus (Rhabdocalyptus) mirabilis (Schulze, 1899)
29

Description. This compressible but very spiny sac 

or vasiform sponge attaches basally to hard substrate. 

External surface is covered by a dense veil of projecting 

pentactins in amongst the long prostal diactin needles, 

with a single large terminal osculum bordered by a 

marginal fringe of diactins. Consistency is soft but spiky. 

Height is to 24 cm, diameter to 15 cm, and 23 mm in 

wall thickness. It may be found in clusters of up to nine 

individuals. Color in life is white but sometimes coated 

with sediment, epizoic organisms, and flocculent mate-

rial, giving it a brown to greenish brown appearance; 

drab when preserved.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are thick prostal diactins to 

4 cm long; hypodermal pentactins all have smooth or 

shagreened (never thorned) tangential rays (1.8–6.6 

mm long) and smooth proximal rays (5.5–9.6 mm 

long); principal diactins (2.0–8.4 mm long); dermal 

stauractins (75–180 mm ray length), dermal pentactins 

with tangential rays (88–193 mm long) and proximal 

rays (88–163 mm long); atrial hexactins (89–134 mm 

ray length). Microscleres are oxyhexasters and oxyhemi-

hexasters (113–179 mm in diameter); discoctasters with 

straight terminals (134–225 mm in diameter); microdis-

cohexasters (16–22 mm in diameter).

Zoogeographic distribution. North Pacific Ocean. 

Locally abundant. In Alaska – eastern Gulf of Alaska. 

Elsewhere – British Columbia to southern California 

(Santa Maria Basin). 

Habitat. In Alaska – attached to bedrock, cobbles, and 

pebbles at depths between 82 and 255 m. Elsewhere – 

reported at depths between 91 and 1373 m.

Remarks. Acanthascus solidus can be distinguished 

from the sometimes similar A. (R.) dawsoni dawsoni by 

its complete lack of thorned hypodermal pentactins 

that are always present and heavily thorned in the latter species. In the central Aleutian Islands a very similar 

species (possibly Acanthascus solidus but not confirmed) 

occurs at depths between 399 and 463 m. This species is 

preyed upon by the sea stars Hippasteria spp., Poraniopsis 

inflata, and possibly Henricia longispina in the eastern 

Gulf of Alaska.

Photos. 1) Mostly intact specimen collected at a depth 

of 167 m in the eastern Gulf of Alaska. Grid marks are 

1 cm

2

. 2) Specimen (indicated by the white arrow) at a 

depth of 165 m in the eastern Gulf of Alaska.

15. Acanthascus (Staurocalyptus) solidus (Schulze, 1899)
30 Professional Paper NMFS 12

Description. This species grows from a narrow attached base to either a thick massive or thick funnelshaped form. External surface has small rounded 

protuberances from each of which project several short 

diactin prostalia; there is a single large terminal osculum. Consistency is soft and easily torn. Height is up to 

24 cm and diameter to 17 cm. Color in life is brown to 

golden brown.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are thick prostal diactins 

(4.3–10.1 mm long), a few rough, but not thorned, 

hypodermal pentactins (0.21–1.09 mm tangential ray 

length); principal diactins (1.11–6.84 mm long); dermal hexactins with some pentactins and stauractins 

(40–139 mm ray length); atrial hexactins (43–218 mm 

ray length). Microscleres are oxyhexactins and a few 

hemioxyhexasters (86–175 mm in diameter); very small 

discoctasters with terminals strongly curved outward 

(56–87 mm in diameter); microdiscohexasters are 

absent.

Zoogeographic distribution. Locally abundant. 

In Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Attached to cobbles and pebbles at depths 

between 139 and 183 m.

Remarks. This species is distinguished from all other 

species of Acanthascus (Staurcalyptus) by the small size 

and shape of its discoctasters. A complete description 

is in preparation.

Photos. 1) Partially fragmented specimen collected 

with a trawl at a depth of 155 m in the central Aleutian 

Islands. Grid marks are 1 cm2

. 2) Specimen at a depth 

of 165 m in the central Aleutian Islands. 

16. Acanthascus (Staurocalyptus) sp. nov. 1 Reiswig and Stone, in preparation
31

Description. Soft tube or sac sponge attaches basally 

to hard substrate. Lateral surface has low conules 15 

mm apart and 3–5 mm high from which small groups 

of thin, short, prostal diactin spicules project; there is 

no veil of pentactins. The single terminal osculum is 

sharp-edged and without a marginal fringe. Both external and internal surfaces (and thus canal openings) are 

covered by tight spicule lattices. Consistency is soft and 

pliable; internal aspect is wooly. The type specimen is 30 

cm tall by 20 cm diameter (at the widest point). Color 

in life is white.

Skeletal structure. It is composed entirely of loose 

spicules excepting a small fused basal attachment 

structure. Megascleres are prostal diactins 9.3–16.2 

mm long; principal diactins 1.8–10.4 mm long; dermal pentactins with spiny tangential rays (72–201 μm 

long) and proximal rays (65–172 μm long); atrial spiny 

hexactins (65–172 μm ray length). Microscleres include 

two classes of discoctasters (124–382 and 112–167 μm 

diameter); stout oxyhexactins (94–139 μm diameter); 

oxy- and hemioxyhexasters (88–133 μm diameter); microdiscohexasters (13–22 μm diameter).

Zoogeographic distribution. Locally common. In 

Alaska – known only from the central Aleutian Islands. 

Elsewhere – not reported.

Habitat. Attached to cobbles in low-relief habitat at a 

depth of 711 m. Video records suggest it is also locally 

common on bedrock, mudstone, and cobbles in moderate-relief habitat at depths between 190 and 1556 m. 

Photos. 1) Whole specimen collected at a depth of 

711 m in the central Aleutian Islands. Grid marks are 1 

cm2

. 2) Same specimen as in photo 1 showing the detail 

of the osculum. 3) Same specimen as in photos 1 and 

2 in situ. The separation between the red laser marks 

is 10 cm.

17. Acanthascus (Staurocalyptus) sp. nov. 2 Reiswig and Stone, in preparation
32 Professional Paper NMFS 12

Description. Vase-shaped sponge attaches basally to 

solid substrate, thickest near its upper end. External surface is smooth and completely lacking large projecting 

spicules; both external and internal surfaces are lined by 

a lattice of loose spicules; there is a single large terminal 

osculum without a marginal spicule fringe. Consistency 

is very soft and easily torn. Height is to 24 cm, diameter 

to 15 cm, and 23 mm in wall thickness. Color in life is 

white; drab when preserved.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are thick diactins (4.4–5.6 

mm long); thin diactins (1.3–5.5 mm long); pinular dermal hexactins with projecting pinular ray (109–175 mm 

long), tangential rays (103–150 mm long) and proximal 

ray (86–140 mm long); pinular atrial hexactins with pro-

jecting pinular ray (93–300 mm long), tangential rays 

(109–214 mm long), and proximal rays (83–192 mm 

long). Microscleres include very large discasters, often 

called ”solasters” (193–438 mm in diameter) with fused 

primary rays recognizable as six hemispherical bosses; 

oxyhexasters and hemioxyhexasters (99–140 mm in di-

ameter); oxyhexactins (83–166 mm in diameter); small 

spherical discohexasters (26–36 mm in diameter).

Zoogeographic distribution. Rare. In Alaska – cen-

tral Aleutian Islands. Elsewhere – western Bering Sea, 

Kuril Islands and off the southern tip of the Kamchatka 

Peninsula.

Habitat. In Alaska – attached to bedrock, mudstone, 

or boulders at depths between 843 and 1715 m. Else-

where – reported at a depth of 117 m but most collec-

tions do not report depth.

Photos. 1) Mostly intact specimen collected at a depth 

of 843 m in the central Aleutian Islands. Grid marks 

are 1 cm

2

. 2) Same specimen as in photo 1 in situ. Note 

that the specimen has been torn on the left side (prior 

to collection) and the osculum is directed to the right.

18. Aulosaccus pinularis Okada, 1932
33

Description. Vase-shaped sponge attaches basally to 

solid substrate, thickest near its upper end. External 

surface is smooth, completely lacking large projecting 

spicules, and lined by a lattice of loose spicules. The 

internal atrial surface has a network of diactin bundles 

crossing the exhalant apertures, but a lattice of atrialia 

is absent. There is a single large terminal osculum lacking a marginal spicule fringe. Consistency is very soft 

and easily torn. Height is up to 45 cm, diameter to 22.5 

cm, and 34 mm in wall thickness. Color in life is creamy 

white; drab when preserved.

Skeletal structure. Skeleton is composed entirely of 

loose spicules. Megascleres are hypodermal pentactins 

and a few triactins and tetractins (0.6–3.4 mm long 

tangential rays; 0.9–4.4 mm long proximal rays); thick 

diactins (3.3–17.7 mm long); thin diactins (1.4–6.7 mm 

long); short atrial diactins (0.4–1.5 mm long); dermalia 

are a mixture of stauractins, pentactins, and hexactins 

(89–181 mm ray length); atrialia are mainly very large 

hexactins with some pentactins (224–1186 mm ray 

length). Microscleres include very large discasters, often called “solasters” (513–1389 mm in diameter), with 

primary rays fused into a slightly irregular sphere; very 

thin oxyhexactins, with irregular variants, and hemi-

oxyhexasters (87–165 mm in diameter); small spherical 

discohexasters (20–39 mm in diameter).

Zoogeographic distribution. Apparently a rare spe-

cies. In Alaska – central Aleutian Islands and Bering 

Sea (Pribilof Canyon). Elsewhere – Japan, Kuril Islands, 

Okhotsk Sea, and off southern California. 

Habitat. In Alaska – in the central Aleutian Islands 

it occurs on bedrock, boulders, and cobbles at depths 

between 1270 and 1350 m. In Pribilof Canyon it occurs 

at a depth of 300 m. Elsewhere – reported at depths 

between 117 and 419 m.

Remarks. Associated fauna include juvenile lithodid 

crabs (Paralomis verrilli and Lithodes couesi), pandalid shrimps, and the large ophiuroid Gorgonocephalus 

eucnemis.

Photos. 1) Whole specimen collected at a depth of 

300 m in Pribilof Canyon, Bering Sea. Grid marks are 1 

cm

2

. 2) Specimen collected at a depth of 1320 m in the 

central Aleutian Islands. 3) Same specimen as in photo 

2 in situ. Two pairs of red lasers each separated by 10 cm. 

19. Aulosaccus schulzei Ijima, 1896
34 Professional Paper NMFS 12

Description. Very small flattened ovoid sponge attaches to a cobble. External surface is smooth and covered 

by a lattice of mainly stauractin dermalia; about one-half 

of the surface is profusely spined by long diactins and 

sports a veil of raised pentactins. An osculum and atrial 

cavity could not be distinguished. Consistency is soft and 

delicate. The specimen is 11 by 8 by 2 mm. It is transparent and colorless.

Skeletal structure. Skeleton is composed entirely 

of loose spicules, but parts of the fused basidictyonal 

skeleton remain attached. Megascleres are hypodermal 

pentactins (0.26–1.4 mm long tangential rays; 0.47–1.55 

mm long proximal rays); thick prostal diactins (3.2–16.8 

mm long); thin diactins (0.96–6.82 mm long); dermal 

stauractins and diactins (81–381 mm ray length). Microscleres consist only of hemioxyhexasters (112–181 

mm in diameter) and oxyhexactins (124–204 mm in 

diameter).

Zoogeographic distribution. Rare. In Alaska – central Aleutian Islands. Elsewhere – small incompletely 

identifiable Bathydorus spp. have been reported worldwide.

20. Bathydorus sp., description by Reiswig and Stone, in preparation

Habitat. In Alaska – occurs on cobble but probably 

also on boulders and bedrock; collected from 494 m but 

may have a broad depth range. Elsewhere – small Bathydorus spp. are reported from depths of 446 to 1625 m.

Photo. 1) Whole specimen collected at a depth of 494 

m in the central Aleutian Islands.
35

Description. A very large mushroom-shaped body 

on a long cylindrical stalk attaches basally to hard substrate. The inhalant surface occupies the upper stalk 

and underside of the body; the exhalant surface on the 

upper body is reflected around the edges to the sharp 

margin on the under edges of the body; there is no distinct osculum. All surfaces are macroscopically smooth 

with large canals covered by lattices of loose spicules. 

Consistency of the body is compressible but firm; the 

stalk is stony hard due to fusion of the internal spicules and is hollow, with two main internal longitudinal 

canals. The specimen, measured dry, is 75 cm in total 

height, the body is 31 by 40 cm transversely, and the 

stalk is 68 cm long and 2.1 cm in diameter (narrowest 

point, 1.8 cm). Color in life is white to creamy white; 

drab when preserved.

Skeletal structure. Spicules in the main body remain 

loose, but the principal diactins in the stalk are fused 

to form a hard, rigid, and long-lasting structure. Megascleres include pinular hexactine dermalia (132–337 

μm pinular ray length, 90–163 μm tangential ray length, 

102–158 μm proximal ray length); supported by hypodermal pentactins (315–862 μm tangential ray length, 

312–1054 μm proximal ray length); atrialia are slightly 

thinner pinular hexactins (131–379 μm pinular ray 

length, 80–167 μm tangential ray length, 88–142 μm 

proximal ray length); supported by hypoatrial pentactins similar to those of the dermal side; choanosomal 

diactins are mainly in loose bundles (1.65–4.06 mm 

long); large choanosomal oxyhexactins are common 

(678–1579 μm ray length). Microscleres are all coarsely 

spined, including hemidiscohexasters and discohexasters (82–177 μm diameter) and discohexactins (90–185 

μm diameter).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – not reported.

Habitat. Occurs singly on bedrock and large boulders 

at depths between 1326 and 2680 m.

Remarks. The Aleutian Caulophacus (Caulophacus)

specimen has been compared to the 18 known species 

of the subgenus and found to be unassignable to any of 

them. Its description as a new species is in progress. It 

is used as a perch by several species of lithodid crabs, 

including Paralomis verrilli.

Photos. 1) Whole specimen collected at a depth of 

1806 m in the central Aleutian Islands. 2) Same specimen as in photo 1. 3) Same specimen as in photos 1 

and 2 in situ with Paralomis verrilli crabs. The separation 

between the red laser marks is 10 cm.

21. Caulophacus (Caulophacus) sp. nov. Reiswig and Stone, in preparation

37

CLASS DEMOSPONGIAE
38 Professional Paper NMFS 12

22. Plakina atka Lehnert, Stone and Heimler, 2005 

Description. Sponge is encrusting. Surface is convo-

luted but less distinctively so than Plakina tanaga. Single 

strands have a smooth surface, not microtuberculate. 

This sponge may cover large areas (up to 30 cm

2

) with 

a thickness of approximately 0.3 to 0.8 cm. The ecto-

some is reddish brown (Aleutian Islands), beige, or light 

brown (eastern Gulf of Alaska).

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Ectosomal dense spicule crust averages 100 μm in thickness. Choanosome is somewhat 

less densely packed with spicules, roughly arranged in 

tracts of varying orientation and with many spicules in 

between. Characteristic plakinid oxeas are slightly bent 

and thickest in the center (70–108 × 3–6 μm). A thicker 

category of diods, always with relatively long spines at 

the center, is probably derived from the spined category 

of triods (82–95 × 8–10 μm). Triods also occur in two 

distinct types: relatively rare simple triaxons (28–33 μm/

ray) and much more abundant, robust triaxons with a 

row of large spines on each ray close to the center of the 

spicule (23–40 × 3–6 μm/ray). Calthrops are rare and 

have a reduced fourth ray only. Tetralophose calthrops 

have tetrafurcate, occasionally pentafurcate rays, with 

microfurcate ends (18–23 μm in total length). 

Zoogeographic distribution. Locally abundant. In 

Alaska – central Aleutian Islands, eastern Gulf of Alaska 

(continental shelf off Cape Ommaney, Baranof Island), 

and southern Southeast Alaska (Portland Canal). Else-

where – northern British Columbia (Portland Canal).

Habitat. Encrusts bedrock, boulders, cobbles, pebbles, 

and hexactinellid skeletons at depths between 82–180 m 

(Aleutian Islands) and 95–253 m in the eastern Gulf of 

Alaska. May cover relatively large areas (up to 30 cm

2

).

Remarks. Plakina atka can be distinguished from 

the similar P. tanaga by its relatively smooth surface 

compared to the strongly convoluted and microtuberculated surface of the latter species, and by the presence of tetralophate lophocalthrops compared to the 

trilophose lophocalthrops of P. tanaga. 

Photos. 1) Specimen collected at a depth of 118 m in 

the central Alautian Islands. Grid marks are 1 cm

2

. 2) 

Specimen collected at a depth of 167 m in the eastern 

Gulf of Alaska. 3) Specimen at a depth of 118 m in the 

central Aleutian Islands.
39

Description. Sponge is encrusting. Convoluted surface has deep grooves between strands with a microtuberculated surface. Consistency is cheese-like. Circular 

oscula flush with the surface are visible in situ. This 

sponge may cover large areas (up to 1 m2) with a thickness of approximately 1 to 1.5 cm. Color in life is beige 

to light brown. 

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome is packed with lophocalthrops with a layer 20–250 μm thick. Spicules are 

characteristically bent diods, often centrotylote or with 

a reduced third ray in the form of a spine (85–97 × 

2–4 μm); two categories of triaxons, smooth, thinner 

triods (24–50 × 2–5 μm/ray) and thicker triods, often 

with one short, slender spine near the base of each ray 

(22–38 × 7–9 μm). Rare smooth calthrops have rays of 

the same dimensions as those of the triods. There are 

abundant trilophose calthrops with each ray tetrafurcate 

and, again microfurcate at the points, somewhat smaller 

than the triods and concentrated in the ectosome. Total 

length of lophocalthrops is 28–42 μm. Non-lophose rays 

are sometimes bifurcate. 

Zoogeographic distribution. Uncommon. In Alaska – 

known only from the type locality in Little Tanaga Strait, 

central Aleutian Islands. Elsewhere – not reported.

Habitat. Encrusts vertical and overhanging surfaces of 

bedrock and boulders at depths between 140 and 383 m. 

May cover relatively large areas (up to 1 m2

).

23. Plakina tanaga Lehnert, Stone and Heimler, 2005 

Remarks. Plakina tanaga can be distinguished from 

the similar P. atka by its strongly convoluted and microtuberculated surface compared to the relatively smooth 

surface of the latter species, and by the presence of trilophose lophocalthrops compared to the tetralophate 

lophocalthrops of P. atka.

Photo. 1) P. tanaga encrusting a bedrock scarp at a 

depth of 146 m in the central Aleutian Islands. 
40 Professional Paper NMFS 12

Description. Sponge is globular to subglobular. Oscula are occasionally present but not obvious. Spicules 

radiate from the center to the surface and protrude 

above it. Consistency is hard, cartilaginous, and only 

slightly elastic due to the high spicule density. With a 

diameter of 8 to 10 cm, this is one of the largest species of Craniella. The cortex is white; the interior of the 

sponge is orange-gold. 

Skeletal structure. Protriaenes may reach a length of 

more than 3 cm. Protriaenes and anatriaenes have relatively small clads; protriaenes, rhabd greater than 3 cm 

× 3–11 μm, clad 20–30 μm; anatriaenes, rhabd up to 10 

mm × 3–11 μm, clads, 40–85 μm, oxeas, up to 3 cm × 

10–40 μm, commata, 7–10 μm.

Zoogeographic distribution. Uncommon, but locally 

patchy. In Alaska – Aleutian Islands. Elsewhere – central 

California to the Gulf of California.

Habitat. In Alaska – found at depths between 88 and 

272 m in boulder and cobble habitats. Typically epizoic 

on calcareous sponges (e.g., Leucandra poculiformis) and 

various demosponges, including Tedania dirhaphis, 

Mycale jasoniae, and M. loveni. Elsewhere – reported at 

depths between 11 and 214 m.

Remarks. Reproduction in this group of sponges is 

oviparous without a larval stage, or viviparous with production of young adults within the parent (Van Soest 

and Rützler, 2002).

Photos. 1) Specimen collected at a depth of 100 m 

in the central Aleutian Islands. Specimen is attached 

to the demosponge Tedania dirhapsis. Grid marks are 

1 cm2

. 2) Same specimen as in photo 1 showing radial 

arrangement of large spicules in the interior and the 

presence of orange-colored eggs or young adults. 

3) Specimen at a depth of 170 m in the central Aleutian 

Islands.

24. Craniella arb (de Laubenfels, 1930)
41

Description. Body is globular with very long tracts 

of protruding spicules. Typically is attached to other 

sponges via these tracts which terminate in anchorlike, long-shafted triaenes. Internal bundles of spicules 

radiate from the center of the globular body to well 

beyond the surface of the sponge. The surface consists 

of a layer of dermal oxeas occurring only there and 

oriented perpendicular to the surface. The diameter 

of the body is 7 to 8 mm (without protruding spicules). 

Spicule tracts protrude 6–8 mm above the surface, giving the sponge a spiny appearance. Color in life is white 

or creamy white.

Skeletal structure. Large oxeas have unequal ends 

(3000–8000 × 45–72 μm), cortical oxeas (460–1340 

× 26–76 μm), protriaenes (1600–7300 μm long with 

clads 13–34 μm), anatriaenes have rhabds (2300–8700 

μm with clads 19–40 μm). The sigmaspires are unique 

among the known species of Craniella in that they are 

relatively large and resemble isochelae (22–34 μm). 

These sigmaspires are abundant and densely present in 

the cortical layer.

Zoogeographic distribution. Uncommon. In Alaska – 

central Aleutian Islands, Bering Sea (Pribilof Canyon). 

Elsewhere – previously known only from the original 

description in the Kuril Islands, Russia. 

Habitat. In Alaska – a cryptic species; epizoic on 

hexactinellid sponges and other demosponges, including Erylus aleuticus, at depths between 190 and 275 m. 

Elsewhere – no information available.

Remarks. Previously known as Tetilla sigmoancoratum 

Koltun, 1966, but recently transferred to the genus 

Craniella since it possesses both a cortex and the special 

cortical oxeas that are lacking in the genus Tetilla.

Photo. 1) Specimen collected at a depth of 208 m in 

Pribilof Canyon, Bering Sea. Specimen is growing on 

an unidentified hexactinellid sponge.

25. Craniella sigmoancoratum (Koltun, 1966)
42 Professional Paper NMFS 12

Description. Sponge is globular with many short, 

sharp spines. It typically attaches to other sponges via 

the spines. Characteristic of the genus, the internal arrangement of spicule bundles is strictly radial with bundles supporting the ectosomal conules, which are less 

than 1 mm in height. The surface is formed by a special 

cortex. Dermal oxeas form a palisade perpendicular 

to the surface. Spicule bundles protrude through the 

conules and diverge. Diameter is to 16–18 mm. Color 

in life is white or creamy white.

Skeletal structure. There are large oxeas (2300–4800 

× 23–42 μm) with unequal ends, both ends acute but 

one end filiform, cortical oxeas (685–1230 × 35–47 

μm), protriaenes (3400–5700 μm long), anatriaenes 

(up to 8000 μm long rhabdomes). Microscleres are 

c- and s-shaped sigmaspires with a maximum diameter 

of 11–14 μm.

Zoogeographic distribution. Uncommon. In Alaska – 

Bering Sea (Pribilof Canyon). Elsewhere – originally described from Vancouver Island. Reportedly occurs from 

the Aleutian Islands to British Columbia (Austin, 1985). 

Habitat. In Alaska – a cryptic species; epizoic on 

Myxilla pedunculata and probably other sponges at a 

depth of 236 m. Elsewhere – depths ranging from 36 to 

76 m (Vancouver Island).

Photo. 1) Specimen collected at a depth of 236 m in 

Pribilof Canyon, Bering Sea. Specimen is attached to 

the demosponge Myxilla pedunculata. 

26. Craniella spinosa Lambe, 1893
43

Description. Sponge is globular with numerous acute 

spines distributed over the surface. The surface between 

the spines is smooth, without any visible apertures. 

Characteristic of the genus, the internal arrangement of 

spicule bundles is strictly radial with bundles supporting 

the ectosomal conules. The sponge is rather hard, only 

slightly elastic. Diameter including the spines is to 30 

mm (16 mm without the spines). Color in life is white 

to creamy white.

Skeletal structure. Spicules are anatriaenes (3430–

8820 × 12–21 μm), clads (48–154 × 17–22 μm per ray), 

and protriaenes (4520–8960 × 12–40 μm). There are 

clads (50–250 × 5–17 μm per ray); choanosomal oxeas 

(4530–5425 × 50–75 μm); two categories of cortical ox-

eas: small, centrotylote oxeas (97–372 × 8–17 μm), tyle 

in the center more like a ring than a tyle, and larger 

ones (540-987 × 28–63 μm). Sigmaspires or similar sig-

moid spicules are absent.

Zoogeographic distribution. Uncommon. In Alaska 

– known only from the type locality in Amchitka Pass, 

central Aleutian Islands. Elsewhere – not reported.

Habitat. Patchy distribution. A cryptic species; epizoic 

on demosponges, including Myxilla sp., and occasion-

ally gorgonian corals (Primnoidae) in boulder, cobble, 

and sand habitats at depths between 115 and 199 m. 

Photos. 1) Specimens collected at a depth of 115 m 

in the central Aleutian Islands. 2) Two specimens (in-

dicated by the white arrows) at a depth of 191 m in the 

central Aleutian Islands. 

27. Craniella sputnika Lehnert and Stone, 2011
44 Professional Paper NMFS 12

Description. Lobate sponge consists of several flattened lobes (oval to circular). The surface is hard due 

to an ectosomal crust of aspidasters about 0.5 mm in 

thickness. Numerous uniporal orifices are scattered 

over the surface at intervals of 0.5–1.0 mm. The orifices 

are circular, 0.5–1.0 mm in diameter, and slightly (<0.5 

mm) raised above the surface. Individual lobes measure as large as 17 × 12 × 3.5 cm. Color in life is creamy 

white.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The choanosome is softer and choanosomal megascleres are extremely variable with many 

malformed and distorted spicules, irregularly bent 

or straight. Spicule shapes range from oxeas, styles, 

tylostyles, strongyles to tylotes, some with irregular 

diameters, some club-shaped, size range 520–1728 × 

18–42 μm, orthotriaenes, also with variable rays, blunt 

or pointed, rays of cladome often bent in one direction, 

rhabds (615-790 × 23-42 μm), clads (270-380 × 24-35 

μm), aspidasters (200–225 × 160–175 × 28–62 μm), 

dermal centrotylote microstrongyles (30–47 × 4–6 μm), 

and oxyasters (9–12 μm in diameter). 

Zoogeographic distribution. Rare. In Alaska – known 

from only two locations north of Amlia and Atka Islands, 

central Aleutian Islands. Elsewhere – not reported.

Habitat. Attached to pebbles or shell hash in cobble, 

pebble, and sand habitats at depths between 127 and 

190 m.

Remarks. Some species of Erylus reportedly contain 

bioactive compounds with antitumor, antifungal, and 

antiviral properties, especially against retroviruses such 

as human immunodeficiency virus (HIV) (Adams and 

Hooper, 2001). E. aleuticus has not yet been examined 

for the presence of bioactive compounds and is a high 

priority for collection for that purpose.

Photos. 1) Specimen collected at a depth of 190 m in 

the central Aleutian Islands. 2) Specimen observed at a 

depth of 127 m in the central Aleutian Islands.

28. Erylus aleuticus Lehnert, Stone and Heimler, 2006
45

Description. Small, cylindrical sponge has an ectosomal hard cortex of sterrasters and a softer choanosome. 

Lendenfeld (1910) described the species as encrusting, 

cushion-shaped, or irregularly finger-shaped. The specimen collected in the Aleutian Islands was a single cylinder approximately 5 cm in height and 1 cm in width. 

Color in life is white to creamy white. 

Skeletal structure. There are sterrasters (170–200 × 

130-140 μm), choanosomal styles and oxeas (1200–1760 

× 30–40 μm), diaenes of the same size as oxeas, and 

tylasters (10 μm). Triaenes are absent. 

Zoogeographic distribution. Rare. In Alaska – Aleutian Islands and Southeast Alaska. Elsewhere – previously known from the west coast of North America from 

Southeast Alaska to southern California.

Habitat. In Alaska – found in cobble, pebble, and sand 

habitat at a depth of 190 m. Epizoic on the demosponge 

Erylus aleuticus. Elsewhere – no information available.

Remarks. Triaenes are absent in G. robusta, which 

prompted Lendenfeld (1910) to erect the genus Geod29. Geodia lendenfeldi nomen novum 

inella that is now regarded as synonymous with Geodia. 

This specimen is the same as Geodia robusta (Lendenfeld, 

1910). G. robusta (Lendenfeld, 1910) is a junior homonym of G. robusta (Lendenfeld, 1907) however, and 

according to the World Porifera Database (Van Soest et 

al., 2008) it has no valid name. Thus, we suggest renaming G. robusta (Lendenfeld, 1910) as Geodia lendenfeldi.

Photo. 1) Fragments of specimen collected at a depth 

of 190 m in the central Aleutian Islands. Grid marks are 

1 cm2. 
46 Professional Paper NMFS 12

Description. This sponge is polymorphic but typically 

has a plate-like or lamellar growth form. The surface of 

the sponge is smooth to the unaided eye but microscopically hispid. Small oscula occur on one side only. The 

consistency is firm, incompressible, and fragile. Size is 

highly variable but attains a length and height exceeding more than 1 m. Color in life is whitish to light brown, 

often with a dark fringe at the distal edge.

Skeletal structure. The choanosome consists of an 

unorganized mix of calthrops and oxeas: calthrops 

(320–540 × 35–48 μm per ray), occasionally triods with 

rays of the same size, oxeas (1240–2650 × 32–47 μm). 

Microscleres are streptasters (14–28 μm) and acanthose 

microxeas (117–157 × 3–5 μm) that are always bent in 

the middle.

Zoogeographic distribution. Common. In Alaska – 

Bering Sea (Zhemchug Canyon) and eastern Gulf of 

Alaska. May reach densities up to 19 per m2 in eastern 

Gulf of Alaska habitats. Elsewhere – holotype described 

from Sea of Japan, also reported from along the California Coast and the Gulf of California. Present records 

strongly suggest a circumboreal distribution in the 

North Pacific. However, morphological differences do 

exist between populations, suggesting some genetic 

isolation (see Remarks).

Habitat. In Alaska – attached to bedrock at depths 

between 149 and 486 m in the Bering Sea and attached 

to bedrock, boulders, and cobbles at depths between 

71 and 255 m in the eastern Gulf of Alaska. Elsewhere 

– reported at depths to 74 m. 

Remarks. Previously known as Normania tenuilaminaris

Sollas 1886, P. tenuilaminaris was originally described as 

having straight acanthose microxeas. The microxeas we 

examined from the Bering Sea specimens were consistently bent but this is the only difference between the 

specimens and in our opinion not sufficient to separate 

the specimens at the species level. However, it is probable that the Bering Sea population is genetically isolated and may have developed different morphological 

characters, such as the bent microxeas. This species is 

preyed upon by the sea stars Hippasteria spp., Poraniopsis inflata, Pteraster tesselatus, Ceramaster patagonicus, and 

possibly Henricia longispina. We have observed a very 

similar sponge in the central Aleutian Islands at depths 

between 142 and 1386 m but we would need to collect 

a specimen to confirm conspecificity.

Photos. 1) Fragments of specimen collected at a depth 

of 175 m in the eastern Gulf of Alaska. Note the hydroids 

30. Poecillastra tenuilaminaris (Sollas, 1886)

growing on the fringe. Grid marks are 1 cm2. 2) Same 

specimen as in photo 1 in situ. 3) Specimen collected 

at a depth of 486 m in Zhemchug Canyon, Bering Sea. 

Grid marks are 1 cm2. 
47

Description. This sponge is endopsammic and discshaped with numerous fistules on the upper surface. 

The surface is smooth with long protruding spicules 

scattered about. The cortical layer is cartilaginous and 

more resilient than the relatively soft choanosome. The 

bottom side of the sponge is very firm and slightly elastic 

and consists of a layer of spongin-cemented sediment. 

In situ the plate is buried in the sediment; only the 

fistules protrude. The fistules are cone-shaped with no 

obvious opening. The disc is up to 52 cm in diameter 

and about 1 cm in thickness (3–4 mm of the thickness 

is due to the basal sediment layer). Color in life is light 

yellow with bright yellow fistules; disc is light brown 

after freezing.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Polyspicular tracts arise from the basal 

sediment layer then widen, branch towards the surface, 

and fan out in the cortical layer. Polyspicular tracts are 

350–500 μm in diameter. The cortical layer is 500–700 

μm in thickness and, except for the ends of the ascend-

ing polyspicular tracts, consists of a mass of spicules 

without orientation. The outermost layer is a palisade 

of small tylostyles, oriented outward creating the mi-

croscopically hispid surface. The papillae also show the 

outermost palisade and the unorganized spicule mass 

below, but differ insofar as the polyspicular tracts run to 

the tip of the papillum. A central canal is visible in the 

center of the papillae but there is no opening visible. 

There are large fusiform tylostyles, in a wide size-range, 

longest ones often with one or several subterminal 

rings (180–1750 × 8–22 μm), and a small category of 

tylostyles (65–110 × 3–6 μm). 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not re-

ported.

Habitat. Patchily distributed at depths between 81 and 

338 m in generally low-relief (i.e., flat-bottomed) habi-

tats of small pebbles and coarse sand with moderate to 

high current. Found at maximum densities up to eight 

individuals per m2 in coarse sand habitats at 82 m depth.

Photos. 1) Specimen collected at a depth of 82 m in 

the central Aleutian Islands. Grid marks are 1 cm

2

. 2) 

Same specimen as in photo 1 in situ during collection. 

3) Specimen observed at a depth of 142 m in the central 

Aleutian Islands. A prowfish (Zapora silenus) lies in a den 

just below the specimen.

31. Polymastia fluegeli Lehnert, Stone and Heimler, 2005
48 Professional Paper NMFS 12

Description. This sponge is subglobular to cushionshaped. Surface is smooth to the unaided eye but 

microhispid with several papillae. Consistency is firm. 

Diameter is to at least 15 cm. Externally it is brownish orange in color; internally it is bright orange with lighter 

polyspicular tracts visible to the unaided eye.

Skeletal structure. Ectosomal palisade of small tylostyles are supported by intermediate tylostyles. Large 

tylostyles radiate in polyspicular tracts to the surface. 

There are large tylostyles (up to 4400 × 20 μm), intermediate tylostyles (200–600 × 12–16 μm), and small 

tylostyles (120–160 × 5–7 μm).

Zoogeographic distribution. Uncommon. North 

Pacific Ocean. In Alaska – central Aleutian Islands. 

Elsewhere – previously reported from Vancouver Island 

to California. 

Habitat. In Alaska – attached to pebbles at depths 

between 150 and 160 m. Elsewhere – 73 m depth (Vancouver Island). Lamb and Hanby (2005) report that 

32. Polymastia pacifica Lambe, 1893

this species occurs from Alaska to California at depths 

between the intertidal zone and 183 m.

Photo. 1) Fragment of a specimen collected at a depth 

of 155 m in the central Aleutian Islands. Grid marks are 

1 cm2.
49

Description. Thin stalk terminates in a globular or 

oval body with a smooth surface. Stalk is to 9 cm; body 

to approximately 2 cm

3

. Color in life is golden brown. 

Skeletal structure. Skeleton of the stalk is a tight mass 

of parallel spicules running into the body and radiating 

to the sides in tracts. Smaller oxeas form a palisade at 

the surface of the body. Microxeas form a tangential 

crust at the surface. Oxeas are exclusively centrotylote, 

sometimes with blunt ends. Oxeas occur in a wide 

size-range, possibly in three categories: 790–2460 × 

10–37 μm, 400–700 × 10–12 μm, and microxeas 70–110 × 

2–3 μm.

Zoogeographic distribution. Locally abundant in 

the North Pacific Ocean. In Alaska – central Aleutian 

Islands and Bering Sea (Pribilof Canyon). Elsewhere – 

Sea of Okhotsk. 

Habitat. In Alaska – attached to pebbles and emergent 

epifauna at depths between 125 and 307 m and generally in low-relief (i.e., flat-bottomed) habitats (central 

Aleutian Islands). Sponges with very similar morphol-

ogy have been observed in the Aleutian Islands at depths 

near 1100 m, but we would need to collect a specimen 

to confirm conspecificity. Elsewhere – reported at a 

depth of 200 m. 

Remarks. Stylocordyla borealis eous can be distinguished 

from the similar Rhizaxinella clavata by its thinner stalk 

and smooth oval body. Also, S. borealis eous has exclu-

sively centrolyte oxeas while R. clavata has tylostyles and 

subtylostyles. 

Photos. 1) Specimen collected at a depth of 127 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 (indicated by the white 

arrow) in situ. 3) Specimen collected at a depth of 208 

m in Pribilof Canyon, Bering Sea. Specimen is attached 

to an unknown hexactinellid sponge.

33. Stylocordyla borealis eous Koltun, 1966
50 Professional Paper NMFS 12

Description. Species is irregularly globular. Surface 

is almost smooth, microscopically slightly uneven. No 

oscula are visible. Consistency is firm, only slightly 

elastic. Diameter ranges from 9 to 19 mm. Color in life 

is mustard yellow.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Spicules are strictly radially arranged 

in the choanosome; spicule density is very high. Special 

dermal oxeas are densely arranged perpendicular to 

the surface and form a palisade. There are fusiform 

strongyloxeas (1795–2132 × 15–22 μm), subtylostyles 

(500–770 × 8–10 μm), and ectosomal tylostyles (104–215 

× 4–8 μm). 

Zoogeographic distribution. Locally common. In 

Alaska – known only from the type locality in Pribilof 

Canyon, Bering Sea. Elsewhere – not reported.

Habitat. Attached to pebbles in low-relief (i.e., flatbottomed) silt and sand habitat at depths between 203 

and 240 m. Often found in association with several 

unknown species of hydroids, zoanthids, and the demosponge Stylocordyla borealis eous.

Photos. 1) Specimen collected at a depth of 219 m in 

Pribilof Canyon, Bering Sea. 2) Same specimen as in 

photo 1 showing close-up detail of the radial arrangement of the choanosome. 3) Same specimen as in 

photos 1 and 2 in situ. The separation between the red 

laser marks is 10 cm.

34. Aaptos kanuux Lehnert, Hocevar, and Stone 2008
51

Description. Thin stalk terminates in a globular or 

oval body. Species is similar to Stylocordyla borealis eous. 

This species has a slightly thicker stalk which widens towards the body and a strongly hispid surface of the body 

compared to the smooth surface of S. borealis eous. Spicules are arranged radially in the interior of the sponge 

with dense spicule brushes at the surface. Length is up 

to 5.5 cm. Color in life is golden brown.

Skeletal structure. There are tylostyles to subtylostyles (1050–1780 × 15–32 μm) and smaller tylostyles 

(180–370 × 6–10 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– Bering Sea (Zhemchug Canyon). Elsewhere – the 

holotype was recorded from the Sea of Japan. 

Habitat. In Alaska – attached to hexactinellid sponge 

skeletons at a depth of 915 m. Elsewhere – reported on 

mud bottoms at a depth of 183 m.

Remarks. Rhizaxinella clavata can be distinguished 

from the similar Stylocordyla borealis eous by its thicker 

35. Rhizaxinella clavata (Thiele, 1898)

stalk and hispid oval body. Also, R. clavata has tylostyles 

and subtylostyles while S. borealis eous has exclusively 

centrolyte oxeas. 

Photo. 1) Specimen collected at a depth of 915 m in 

Zhemchug Canyon, Bering Sea. Specimen is attached to 

an unidentified hexactinellid sponge skeleton.
52 Professional Paper NMFS 12

Description. This massively lobate sponge has irregularly cylindrical lobes. Surface is smooth to the unaided 

eye but microscopically hispid. Circular oscula are near 

tips of lobes and slightly elevated above the surface. 

Consistency is firm but elastic. Height is to 14 cm. Color 

in life is yellowish brown.

Skeletal structure. Skeletal architecture is more or 

less confused in the interior but becomes radial towards 

the surface. The surface consists of a palisade of smaller 

tylostyles. There are thick tylostrongyles with round 

tyles at one end and only slightly tapering at the other 

(640–1890 × 17–46 μm), somewhat thinner and flexuous tylostrongyles (1140–2100 × 17–26 μm), and small 

tylostyles forming the palisade at the surface (175–520 

× 8–12 μm).

Zoogeographic distribution. Rare. In Alaska – Bering 

Sea (Zhemchug Canyon) and eastern Gulf of Alaska. 

Elsewhere – Holotype described from Sagami Bay, Japan; also recorded along Korean Coast (Sim and Kim, 

1988).

Habitat. In Alaska – attached to boulders and cobbles 

at depths between 390 and 601 m. Elsewhere – no information available.

Remarks. This was previously known as Rhizaxinella 

excellens Thiele, 1898. It typically harbors an abundant 

and diverse complement of ophiuroid associates.

Photos. 1) Fragments of specimen with ophiuroid 

associates collected at a depth of 390 m in Zhemchug 

Canyon, Bering Sea. 2) Close-up view of oscula atop 

the lobes.

36. Suberites excellens (Thiele, 1898)
53

Description. This small sponge has a subglobular to 

globular growth form. Surface is smooth to the unaided 

eye but microscopically hispid. Consistency is firm but 

elastic. Diameter is at least 5 cm. Color in life is golden 

brown to brown.

Skeletal structure. Ectosomal skeleton consists of bouquets of small tylostyles at the surface. Densely packed 

larger tylostyles in the interior are arranged without 

orientation. Tylostyles are large (620–1170 × 10–20 μm) 

and small (120–230 × 3–5 μm).

Zoogeographic distribution. Uncommon in the 

North Pacific Ocean. In Alaska – central Aleutian Islands. Elsewhere – Vancouver Island. 

Habitat. In Alaska – attached to pebbles at depths between 150 and 160 m. Elsewhere – reported at a depth 

of 73 m.

37. Suberites simplex Lambe, 1893

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
54 Professional Paper NMFS 12

Description. This “mobile” sponge lives as an irregularly globular encrustation on empty gastropod shells. 

Frequently lives in symbiosis with hermit crabs including Pagurus dalli. Surface is smooth to the unaided eye 

but microscopically hispid due to dense spicule brushes 

that leave open numerous small pores (50–100 μm diameter). Irregular short tracts in the choanosome form 

a vague reticulation. Diameter is at least 10 cm. Color in 

life is red, orange, or tan; pale yellow in ethanol.

Skeletal structure. Viewed under high magnification, 

the spicules are relatively dense without particular orientation. There are tylostyles to tylostrongyles (123–350 

× 7–12 μm).

Zoogeographic distribution. Locally abundant. In 

Alaska – Bering Sea to Southeast Alaska. 

Habitat. In Alaska – typically encrusts gastropod shells 

at depths between 20 and 165 m. 

Remarks. This species is indistinguishable from Suberites domuncula that is common to the Mediterranean Sea 

and Atlantic Ocean along the west coast of North Africa. 

However, we doubt conspecificity based on the disjunct 

zoogeography and leave the species assignment for this 

specimen as undecided. The species complex is in desperate need of taxonomic revision. There are probably 

several sibling species that live in symbiosis with hermit 

38. Suberites sp.

crabs (more than a dozen species reported worldwide). 

S. domuncula contains suberitine, a neurotoxin that can 

cause fatal hemolytic hemorrhaging in some animals 

including some reef fish. However, the hawksbill turtle

(Eretmochelys imbricata) does prey upon it in tropical waters (Meylan, 1988). No studies have been conducted 

on Alaskan specimens.

Photo. 1) Specimen collected at a depth of 84 m in 

the central Aleutian Islands. This is the ventral side of 

the sponge showing the cavity in which a hermit crab 

Pagurus dalli (completely retracted) is living. Grid marks 

are 1 cm2.
55

Description. This sponge is massively encrusting and 

very fragile. Surface is smooth to the unaided eye but 

microscopically hispid. No oscula are visible. Fristedt 

(1887) described this species as forming irregular 

knolls, 9 cm in longest dimension. Color in life is whitish 

yellow to creamy white.

Skeletal structure. It has an irregular unispicular or 

paucispicular mesh of oxeas (250–340 × 9–11 μm) and 

stigmata (40–60 μm).

Zoogeographic distribution. Widespread but uncommon. In Alaska – Bering Sea (Zhemchug Canyon). Elsewhere – North Pacific Ocean (Sea of Okhotsk and Sea 

of Japan), Arctic Ocean (Barents Sea, Kara Sea, Laptev 

Sea), North Atlantic Ocean (West of Spitzbergen, East 

of Greenland, Denmark Strait, Davis Strait, Gulf of St. 

Lawrence, between Iceland and the Faroe Islands).

Habitat. In Alaska – attached to cobbles at a depth of 

909 m. Elsewhere – reported at depths between 68 and 

256 m.

39. Hemigellius porosus (Fristedt, 1887)

Photo. 1) Fragments of specimen collected at a depth 

of 909 m in Zhemchug Canyon, Bering Sea.
56 Professional Paper NMFS 12

Description. This stalked fan-shaped sponge consists 

of a meshwork or net; not flat but slightly concave. 

Consistency is wiry due to the mesh of spicule tracts. It 

is relatively small; height to 11 cm. Color in life is light 

yellow.

Skeletal structure. Structure includes ectosomal 

tylotes with acanthaceous ends (280–420 × 6–8 μm); 

choanosomal styles with slightly acanthaceous blunt 

ends in a wide size range (215–2000 × 18–33 μm); and 

spherical isochelae (17–25 μm) and toxa in two size 

categories – large (55–220 μm) and small (10–18 μm).

Zoogeographic distribution. Uncommon. In Alaska – 

central Aleutian Islands. Elsewhere – previously known 

from only two records near the Commander Islands 

(Russia) in the western Bering Sea.

Habitat. In Alaska – attached to bedrock or mudstone 

at depths between 843 and 1720 m. Elsewhere – reported at a depth of 2440 m.

Remarks. Previously known as Melonchela clathriata

Koltun, 1955.

Photos. 1) Specimen collected at a depth of 1720 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ.

40. Cornulum clathriata (Koltun, 1955)
57

Description. This sponge is massively encrusting. 

According to Koltun (1959), growth form may also 

be massive-lobate, tabular, dactylate, ramified, or irregularly lobate. Oscula are on short elevations or flush 

with the surface. Dermal membrane is thin and pelli-

cular. Consistency is soft and easily torn. It may reach a 

height of 15 cm. Color in life is golden brown to brown.

Skeletal structure. Ectosomal tornotes (150–270 × 

6–10 μm) with micro-spined heads, are arranged in bun-

dles tangential to the surface. Choanosomal skeleton is 

reticulate with smooth styles or acanthostyles (195–435 × 

9–18 μm), anisochelae (18–37 μm), and bipocillae 

(8–14 μm).

Zoogeographic distribution. Widespread and locally 

common. In Alaska – central Aleutian Islands. Else-

where – North Pacific Ocean (Sea of Okhotsk and Sea 

of Japan), Arctic Ocean (Barents Sea including White 

Sea, Kara Sea – Vilkitsky Strait, and Greenland Sea), and 

North Atlantic Ocean (Norwegian Sea, Davis Strait, and 

Denmark Strait). 

Habitat. In Alaska – attached to pebbles and small 

cobbles at depths between 94 and 155 m. Elsewhere – 

eurybathic; reported at depths between 9 and 1785 m.

Remarks. This species often grows in association with, 

and encrusts, hydroids and bryozoans.

Photos. 1) Specimen collected at a depth of 94 m in 

the central Aleutian Islands. Specimen is encrusting the 

hydroid Abietinaria sp. (lower right). Grid marks are 1 

cm

2

. 2) Same specimen as in photo 1 (indicated by the 

white arrows) in situ with the plumose hydroids (Abieti-

naria sp.). 3) Specimen collected at a depth of 155 m 

in the central Aleutian Islands. Specimen is encrusting 

the bryozoan (Microporina cf. articulata). Grid marks 

are 1 cm

2

.

41. Iophon piceum (Vosmaer, 1882)
58 Professional Paper NMFS 12

Description. This sponge is massively encrusting. It 

is very similar to Iophon piceum except the ectosome 

is white rather than golden-brown and the oscula are 

distinctly conical. Diameter is to 25 cm. Color in life is 

white; turns completely brown with darker brown oscula 

in ethanol.

Skeletal structure. It is very similar to Iophon piceum

except that bipocillae are absent.

Zoogeographic distribution. Uncommon. In Alaska – 

central Aleutian Islands. Elsewhere – Sea of Okhotsk. 

Habitat. In Alaska – attached to boulders, cobbles, and 

pebbles at depths between 82 and 192 m. Elsewhere – 

reported at a depth of 1240 m.

Photos. 1) Specimen collected at a depth of 100 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ. 

42. Iophon piceum abipocillus Koltun, 1959
59

Description. This sponge has clusters of 4–8 tubes with 

a common stalk. The surface is finely hispid with no rec-

ognizable oscula. The consistency is soft and elastic. The 

dimensions of the short stalk are 2.0 × 0.5 cm and the 

tubes are approximately 11 × 5 cm. Color in life is light 

yellow; golden brown after freezing; beige in ethanol.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome is a thin translucent 

membrane with tangentially arranged tylotes, singly or 

in small bundles, facing in all directions within the tan-

gential plane. The ectosome contains many isochelae 

and is supported by underlying styles. The choanosome 

is a combination of a unispicular reticulation of single 

spicules and ascending paucispicular tracts connected 

by single spicules. Ascending tracts of styles penetrate 

the ectosomal membrane and cause the hispidation. 

Ectosomal tylotes (245–380 × 4–9 μm) have acanthose 

heads. Smooth choanosomal styles measure 490–615 × 

18–22 μm. Microscleres are palmate isochelae (13–17 

μm) with narrow extensions, a small category of palmate 

isochelae (6–8 μm), and a small category of distorted 

anisochelae (4–6 μm). 

Zoogeographic distribution. In Alaska – locally abun-

dant in central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Attached to bedrock, boulders, and cobbles 

at depths between 702 and 750 m.

Photos. 1) Specimen collected at a depth of 702 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ.

43. Megaciella anisochela Lehnert, Stone and Heimler, 2006
60 Professional Paper NMFS 12

Description. This irregularly massive-lobate sponge 

has an uneven, corrugated surface. Oscula are present 

on top of cylindrical elevations. Consistency is elastic, 

soft, and easily torn. Diameter is to 13 cm. Color in life 

is golden brown.

Skeletal structure. Ectosomal tornotes (160–210 × 

4-6 μm) have acanthose ends. There are choanosomal 

acanthostyles (175–212 × 12–16 μm), arcuate isochelae 

(28–32 μm), and toxa (90–150 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – eastern Tartar 

Strait, near the Pacific Coast of the southern Kuril Islands. 

Habitat. In Alaska – attached to small cobbles and pebbles in generally low-relief (i.e., flat-bottomed) habitat 

at depths between 150 and 160 m. Elsewhere – attached 

to pebbles in sandy habitat at depths between 71 and 

414 m and at temperatures between 2.3 and 6.6°C.

44. Megaciella spirinae (Koltun, 1958)

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
61

Description. This sponge is elongated and massive. It 

is irregularly ramified with a corrugated surface; oscula 

are flush with the surface. Diameter is to 15 cm. Color 

in life is orange-brown.

Skeletal structure. Dermal styles (270–360 × 3–6 μm) 

have acanthose heads. There are choanosomal smooth 

styles (420–540 × 18–35 μm), isochelae (16–20 μm), and 

toxa (130–720 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – Arctic Ocean 

(Barents Sea near Murmansk, Russia) and Northeast 

Atlantic Ocean (from Ireland to France and Norway). 

Habitat. In Alaska – attached to small cobbles and pebbles in generally low-relief (i.e., flat-bottomed) habitat 

at depths between 150 and 160 m. Elsewhere – reported 

at depths between 72 and 440 m and temperatures between 1 and 4.2° C.

Remarks. Specimens from the Northeast Atlantic 

are described as flabellate with a characteristic honeycombed surface of a different color (Van Soest and 

Stone, 1986). The spicule complement (types and size 

45. Clathria (Clathria) barleei (Bowerbank, 1866)

ranges) of this specimen conforms perfectly to Clathria 

barleei so we assume it to be another growth form of the 

same species in the Pacific Ocean. Van Soest and Stone 

(1986) report this species from the Arctic to the west 

coasts of Ireland and France so a circumpolar distribution seems possible. 

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
62 Professional Paper NMFS 12

Description. This sponge is massive, laterally compressed, and irregularly lobate. Consistency is soft and 

fragile with round, slightly elevated oscula. Diameter is 

to 30 cm. Color in life is golden brown. The ectosomal 

membrane may have a greyish tinge.

Skeletal structure. Structure includes ectosomal 

smooth small styles (230–290 × 6–10 μm); choanosomal 

smooth styles (440–560 × 20–30 μm) occasionally with 

acanthose heads; and small acanthostyles (90–240 × 

8–12 μm). Microscleres are palmate isochelae (20–25 

μm) and toxas (200–530 μm).

Zoogeographic distribution. Locally common in 

North Pacific Ocean. In Alaska – central Aleutian Islands. Elsewhere – known from Kuril Strait (Russia) and 

near Vancouver Island (Canada).

Habitat. In Alaska – attached to cobbles and pebbles 

at a depth of 167 m. Elsewhere – reported at depths 

between 72 and 138 m.

Photos. 1) Fragment of a specimen collected at a 

depth of 167 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 (indicated by the white arrow) in situ.

46. Clathria (Clathria) laevigata Lambe, 1893
63

Description. This sponge grows from an encrusting 

base to a mass of branches or lobes with an uneven 

surface. Consistency is soft and elastic. Diameter is to 

30 cm. Color in life is golden brown.

Skeletal structure. Ectosome consists of brushes of 

smaller subtylostyles and tylostyles with large protruding 

choanosomal styles. Choanosomal smooth styles, often 

with acanthaceous heads, are arranged in plumose 

tracts and echinated by small acanthostyles. There are 

eectosomal tylostyles and subtylostyles (520–1370 × 8–18 

μm); choanosomal large styles (540–1960 × 28–40 μm) 

with acanthose heads, acanthostyles (120–430 × 10–20 

μm), and palmate isochelae (18–25 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – southern Sea 

of Okhotsk and Sea of Japan (southern Kuril Islands).

Habitat. In Alaska – attached to small cobbles and 

pebbles in generally low-relief (i.e., flat-bottomed) 

habitat at depths between 150 and 160 m. Elsewhere – 

reported at depths between 91 and 550 m on oozy sand 

47. Clathria (Axosuberites) lambei (Koltun, 1955)

and pebbles and at temperatures between 1 and 1.3°C 

and a salinity of 34.18 psu.

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
64 Professional Paper NMFS 12

Description. This sponge has a stalked vase with a 

flared lip and a markedly concave surface; walls are 

5–10 mm in thickness. Consistency, except for the stalk, 

is very soft and elastic. In areas of water current this species typically bends over near the base and sometimes 

lies in contact with the seafloor. Height is to 13 cm and 

width to 3 cm. Color in life is light yellow to creamy 

white; golden brown after freezing.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome is a thin membrane 

packed with anchorate isochelas (21–27 μm). The choanosome consists of paucispicular tracts of thick styles 

(760–920 × 16–21 μm) and brushes of thin styles that fan 

out towards the surface with many isochelas in between. 

Thin styles (740–1230 × 4–6 μm) are slightly curved to 

sinuous and have finely acanthose heads. 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Occurring individually on exposed bedrock, 

cobbles, and pebbles at depths between 622 and 876 m 

and generally in low-relief (i.e., flat-bottomed) habitats.

Photos. 1) Preserved (frozen) specimen collected at 

a depth of 744 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 in situ 

during collection. 3) Same specimen as in photos 1 and 

2 in situ showing detail of the flared lip.

48. Echinoclathria vasa Lehnert, Stone and Heimler, 2006
65

Description. This sponge is stalked with a subhemi-

spherical or conical body. The stalk widens gradually 

from 4 to 25 mm over a distance of approximately 9 cm 

and is not sharply separated from the body. The consistency is soft, elastic, and easily torn. There are wart-like, 

slightly elevated oscula on the dorsal surface of the 

sponge only that are circular and 2 mm in diameter. Di-

ameter of conical body is to 20 cm; total height to 15 cm 

or more. Color in life is light orange to golden brown. In 

ethanol the body has a whitish, translucent ectosome on 

top with a yellowish, fibrous choanosome underneath.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome of the conical body is 

composed of a mesh-work of polyspicular tracts (55–175 

μm in diameter) of small styles with a mesh-size of 350–

750 μm. This large mesh is subdivided by a finer net of 

strands of spongin-embedded isochelae. This finer net 

has a mesh-size of 45–90 μm; single, translucent strands 

are 15–25 μm in diameter. The ectosome of the stalk 

is thinner and consists of a dense, unispicular layer of 

tangentially arranged, parallel-oriented thick styles. In 

the choanosome of the stalk, ascending polyspicular 

tracts of thick styles are connected by paucispicular 

tracts and single spicules, comparable to the choano-

some of the conical body. There are large, smooth styles 

(400–520 × 20–25 μm), small styles (330–550 × 10 μm) 

with acanthose heads and often with one prominent 

dent, isochelae (10–13 μm), and toxa (110–170 μm). 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Attached to bedrock, boulders, cobbles, and 

pebbles at depths between 97 and 253 m. Associated 

with the demosponge Mycale carlilei.

Photos. 1) Specimen collected at a depth of 119 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ showing dorsal 

surface with elevated oscula.

49. Artemisina amlia Lehnert, Stone and Heimler, 2006
66 Professional Paper NMFS 12

Description. This sponge is stalked and tube-shaped 

in the Aleutian Islands, but Koltun (1959) described 

this species as cushion-shaped, spherical, or somewhat 

elongated, and up to 5.5 cm in height. The surface of 

the sponge is smooth at the stalk and gradually changes 

from hispid to rugose in the lower half and is smooth 

again in the apical region. No oscula are visible. The 

consistency of the stalk is wiry and gets softer along its 

length. In areas of water current this species typically 

bends over near the base and sometimes lies in contact 

with the seafloor. Stalked forms are to 31 cm in height 

and 7 cm in width. Color in life is brown to golden 

brown.

Skeletal structure. There are ectosomal brushes of 

smaller subtylostyles (280–428 × 9–18 μm); choanosomal large subtylostyles (430–676 × 6–9 μm), occasionally 

with acanthaceous heads; palmate isochelae (6–15 μm); 

and toxas with acanthaceous ends (60–360 μm).

Zoogeographic distribution. Widespread and locally 

common. In Alaska – central Aleutian Islands. Elsewhere – widely distributed in the Arctic Ocean (Barents 

Sea – White Sea, Kara Sea – Vilkitsky Strait, Greenland 

Sea – Island of Spitsbergen) and Northeast Atlantic 

Ocean (Norwegian Sea, Denmark Strait, Davis Strait). 

Habitat. In Alaska – attached to cobbles and pebbles 

at depths between 170 and 436 m. Elsewhere – reported 

at depths between 14 and 1000 m and temperatures 

between 1.4 and 5.0°C.

Remarks. Specimens from the Aleutian Islands differ in form from previous records but the spicule 

complement (types and size ranges) of the specimens 

conform perfectly to A. arcigera so we assume they are 

a previously unreported growth form of the same species. The shrimp Eualus barbatus is often associated with 

this sponge.

Photos. 1) Specimen collected at a depth of 192 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ (center). 

3) Same specimen as in photos 1 and 2 showing details 

of the rugose surface and apical region.

50. Artemisina arcigera (Schmidt, 1870)
67

Description. This sponge is flabellate in the Aleutian Islands but spherical and lobate forms have been 

reported elsewhere. It is basally stalked and gradually 

tapers to a holdfast. Conspicuous oscula are present on 

dorsal surfaces. The stalk is cylindrical and rigid. The 

body is soft and elastic. Height and width are at least 50 

cm. The ectosome is yellow or light brown. The choanosome is orange-brown. 

Skeletal structure. Ectosomal skeleton has tangentially arranged small styles with acanthose heads (220–340 

× 6–10 μm). The reticulate choanosomal skeleton has 

fusiform styles with acanthose heads (460–510 × 18–25 

μm). Microscleres are palmate isochelae (10–19 μm).

Zoogeographic distribution. Locally abundant. In 

Alaska – central Aleutian Islands. Elsewhere – previously 

known from only two specimens collected in southern 

Kuril Strait until discovered in the central Aleutian 

Islands in 2004.

Habitat. In the Aleutian Islands – attached to bedrock 

outcrops, boulders, and cobbles at depths between 80 

and 239 m on shelf and upper slope habitats. Elsewhere 

– no information available.

Remarks. It is similar in appearance to Tedania kagalaskai, with which it co-occurs. The dorsal surface of A. 

stipitata is typically more rounded than T. kagalaskai and 

the body form of the latter species is much smoother 

and more distinctly triangular. A. stipitata is often associated with the gorgonian Fanellia compressa. 

Photos. 1) Specimen collected at a depth of 150 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ showing the 

51. Artemisina stipitata Koltun, 1958 

detail of the oscula on the dorsal surface. 3) Specimen 

observed in situ at a depth of 87 m in the central Aleutian Islands. 4) Specimen observed in situ at a depth of 

105 m in the central Aleutian Islands. A juvenile king 

crab uses the sponge as a perch. 
68 Professional Paper NMFS 12

Description. Aleutian Island specimens consist of 

many large rounded lobes growing from a narrow base. 

Large lobes often develop side-branches. Koltun (1959) 

described this same species as funnel-shaped, thickly 

tabular or massive. The surface is strongly rugose with 

a thin ectosomal membrane. Circular oscula are flush 

with the surface. It is soft and elastic but fragile. Size is 

large with height to 40 cm and width to 60 cm. Color 

in life varies from yellowish grey, light brown to orange; 

typically golden yellow color in situ.

Skeletal structure. Ectosomal thin styles form a uni-

spicular reticulation with microscleres scattered in 

between. Choanosomal multispicular tracts have large 

styles and single spicules. Styles are often acanthaceous 

at the base. There are large styles (490–840 × 15–30 μm), 

small styles (210–480 × 7–10 μm), palmate isochelae 

(12–22 μm), small toxa (30–120 μm), and large toxa 

(280–340 μm).

Zoogeographic distribution. Locally abundant. In 

Alaska – central Aleutian Islands. Elsewhere – Arctic 

Ocean (Barents, Greenland, Kara, and Laptev seas). 

Habitat. In Alaska – attached to bedrock, boulders, 

and cobbles at depths between 80 and 195 m. Elsewhere 

– reported at depths between 18 and 380 m.

Remarks. This is Artemisina apollinis sensu Koltun 

(1959). However, A. apollinis was originally described 

from near the Kerguelen Islands in the southern In-

dian Ocean so conspecificity is unlikely. It is similar in 

appearance to Mycale loveni with which it co-occurs, but 

this species is much more heavily lobed than M. loveni

and narrows to a stout and narrow base. M. loveni is more 

massive with a less defineable base. It may be preyed 

upon by the sea star Henricia sp. 

Photos. 1) Specimen collected at a depth of 195 m in 

the central Aleutian Islands. Grid marks are 1 cm

2

. 2) 

Specimen collected at a depth of 155 m in the central 

Aleutian Islands. Grid marks are 1 cm

2

. 3) Specimen 

observed at a depth of 122 m in the central Aleutian 

Islands.

52. Artemisina sp.
69

Description. This sponge is subglobular or massively 

encrusting. The smooth surface is covered irregularly 

with conical papillae of very different sizes, ranging 

from 2–20 mm in height and 1–18 mm in diameter. 

Individuals are to about 8 cm in diameter. Color in life 

is light orange; beige in ethanol.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome is a tangential arrangement of tylote bundles in some areas and a tangential 

arrangement of all spicule types in other areas. In the 

choanosome, strongyles are arranged halichondroid, 

relatively dense; polyspicular tracts are vaguely recognizable in some areas. The choanosomal megascleres vary 

from blunt ended (anis-) oxeas to strongyles (570–634 

× 27–32 μm), always with finely acanthose ends. Ectosomal tylotes (325–364 × 7–10 μm) have slightly acanthose 

ends. Microscleres are arcuate isochelae (47–52 μm). 

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – not reported.

Habitat. Attached to cobbles and pebbles at depths 

between 100 and 155 m.

53. Coelosphaera oglalai Lehnert, Stone and Heimler, 2006

Remarks. They may be overgrown with bryozoans and 

found in association with the demosponge Iophon piceum 

abipocillus.

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
70 Professional Paper NMFS 12

Description. This sponge is subglobular with a conulose surface. Tiny conules are distributed uniformly 

over the surface of the sponge, but oscula are atop small 

papillae that are restricted to the dorsal surface. The 

oscula contract upon fixation. Consistency is rather firm 

but elastic. Diameter is to approximately 50 cm. Color 

in life is orange, yellow, to creamy white; pale yellow in 

ethanol.

Skeletal structure. There is a thick dermal membrane, 

densely packed with tylotes (300–400 μm in thickness) 

without apparent organization. Tylotes (about 240–350 

× 5–10 μm) have distinctive swollen heads that form 

polyspicular tracts arranged in an irregular meshwork. 

There are no microscleres.

Zoogeographic distribution. Locally common in 

the North Pacific Ocean. In Alaska – central Aleutian 

Islands. Elsewhere – Bering Sea near Mednyi Island 

(Commander Islands), western Sea of Okhotsk, and 

Sea of Japan.

Habitat. In Alaska – attached to bedrock and cobbles 

at depths between 56 and 138 m. Elsewhere – on rock, 

gravel, and sand at depths between 6 and 299 m and a 

temperature of 2°C.

Remarks. The genus Inflatella typically has strongyles but Koltun (1959, fig. 59) described Inflatella 

globosa as having tylotes. So this species as described by 

Koltun might be transferred to another genus in the 

future. I. globosa is similar to Kirkpatrickia borealis but 

is more globular with oscules on top of small papillae, while the latter species is irregularly massive with 

relatively inconspicuous oscules. Also, I. globosa has 

tylotes only, while K. borealis has tylotes, styles, and a few 

strongyles. 

54. Inflatella globosa Koltun, 1955

Photos. 1) Specimen collected at a depth of 138 m in 

the central Aleutian Islands. Grid marks are 1 cm2. 2) 

Same specimen as in photo 1 in situ. 
71

Description. This sponge is globular. Surface is very 

uneven, covered with numerous wart-like papillae. 

Oscula atop the papillae contract upon collection. The 

contracted surface of the specimen on deck (see photo) 

suggests that it looks quite different in situ. Consistency 

is only slightly elastic, easy to tear. Ectosome is relatively 

thick and easily detachable. Size is to 30 cm (diameter) 

× 20 cm (height). Color in life ranges from yellow to 

light orange.

Skeletal structure. This species has ectosomal tornotes (380–750 × 10–18 μm) resembling strongyles and 

styles, choanosomal acanthostyles (290–680 × 20–40 

μm), and arcuate isochelae (42–47 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – known from the 

Bering Sea (Russia) and the Sea of Okhotsk.

Habitat. In Alaska – attached to cobbles and boulders 

at depths between 87 to 220 m. Elsewhere – reported at 

depths between 32 and 198 m.

Remarks. Koltun (1958) mentions strongylote ectosomal megascleres only in his description of this 

species, but we note some individual differences 

55. Lissodendoryx (Lissodendoryx) behringi Koltun, 1958

between specimens that we have examined. In some 

specimens strongylote forms dominate, while others 

have mainly stylote ectosomal tornotes. As all other 

characters and the dimensions of the spicules are within 

the range for L. behringi, we attribute these differences 

to intraspecific variations. Gross morphology and spicule complement are similar to Phorbas paucistyliferus, but 

the spicule architecture is quite different. 

Photo. 1) Specimen collected at a depth of 124 m in 

the central Aleutian Islands. Grid marks are 1 cm2. 
72 Professional Paper NMFS 12

Description. This sponge is massively encrusting. 

Oscula are in small depressions; surface is corrugated. 

Consistency is soft and very fragile. Diameter is to 15 cm. 

Color in life is brown to golden brown.

Skeletal structure. There are tangentially arranged 

ectosomal tylotes (240–290 × 4–8 μm), often strongylote. Choanosomal acanthostyles are in two size categories – large (235–310 × 12–18 μm), and smaller echinating acanthostyles (120–160 × 6–8 μm). There are large 

arcuate chelae (48–52 μm), small isochelae (15–20 μm), 

and sigmas (48–62 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – Arctic Ocean 

(western Barents Sea) and North Atlantic Ocean (near 

Norway).

Habitat. In Alaska – attached to small cobbles and 

pebbles in generally low-relief (i.e., flat-bottomed) habitat and at depths between 150 and 160 m. Elsewhere – 

reported at depths between 130 and 210 m.

56. Lissodendoryx (Ectyodoryx) olgae (Hentschel, 1929)

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands growing with the gorgonian 

Plumarella sp. Grid marks are 1 cm2.
73

Description. This sponge is massive-lobate and en-

crusting. Surface has small papillae. A thick ectosomal 

membrane covers an underlying honeycombed struc-

ture of the ectosome. Consistency is elastic and com-

pressible. Diameter is to 15 cm. Color in life is golden 

brown.

Skeletal structure. There are ectosomal oxeas and tylotes (239–322 × 9–12 μm), choanosomal acanthostyles 

(231–426 × 12–18 μm), and arcuate isochelae (16–

21 μm).

Zoogeographic distribution. Rare. In Alaska – central 

Aleutian Islands. Elsewhere – previously known from 

only two records in the Sea of Okhotsk (east of Sakhalin 

Island and in Kuril Strait). 

Habitat. In Alaska – encrusts other demosponges at 

depths between 758 and 955 m. Elsewhere – reported 

at depths between 100 and 110 m.

Remarks. Aleutian Island specimens encrust near the 

base of the demosponges Esperiopsis flagrum and Abesto-

pluma ramosa.

Photos. 1) Specimen collected at a depth of 954 m 

in the central Aleutian Islands. Specimen is encrusting the demosponge Esperiopsis flagrum near the base. 

Grid marks are 1 cm

2

. 2) Same specimen as in photo 1 

(indicated by the white arrow) in situ. The separation 

between the red laser marks is 10 cm.

57. Lissodendoryx (Lissodendoryx) oxeota Koltun, 1958
74 Professional Paper NMFS 12

Description. This species appears as described by 

Koltun (1958) as a globular or massive-lobate sponge, 

only slightly elastic and up to 6 cm in height. It bears 

closely spaced cone-shaped papillae and has an easily 

detachable dermal membrane. Color in life is gold to 

golden brown.

Skeletal structure. Spicules form an irregular mesh. 

Styles (495–1050 × 15–24 μm) are smooth, generally 

slightly acanthose at the blunt end. Ectosomal tylotes 

(275–415 × 8–10 μm) have acanthose ends. Microscleres 

are arcuate isochelae (25–38 μm).

Zoogeographic distribution. Rare. In Alaska – Bering 

Sea (Zhemchug Canyon). Elsewhere – previously known 

only from the Sea of Okhotsk.

Habitat. In Alaska – attached to hexactinellid sponge 

skeletons at a depth of 911 m. Elsewhere – no information available. 

58. Lissodendoryx (Lissodendoryx) papillosa Koltun, 1958

Photo. 1) Specimen collected at a depth of 911 m 

in Zhemchug Canyon, Bering Sea, with associated 

ophiuroids. 
75

Description. This stalked sponge widens from a 

relatively narrow peduncle to the appearance of many 

agglutinated tubes (up to eight tubes). Growth form 

may also be flabellate, dactylate, or irregularly lobate 

(Koltun, 1959). Surface is smooth; large oscula are arranged in rows on the apical surface. The consistency 

is elastic and compressible but difficult to tear due to 

the polyspicular tracts of the choanosome. Height is to 

at least 17 cm. Color in life is golden brown to brown.

Skeletal structure. There are ectosomal thin styles 

(120–200 × 7–9 μm), choanosomal styles (180–360 × 

13–20 μm) that form an irregular meshwork of polyspicular tracts, anchorate isochelae (70–90 μm), and 

small isochelae (30–40 μm).

Zoogeographic distribution. Widespread but uncommon in the North Pacific Ocean. In Alaska – central 

Aleutian Islands and Bering Sea. Elsewhere – Bering Sea 

(Russia) and Sea of Okhotsk. 

Habitat. In Alaska – attached to bedrock and cobbles 

at depths between 146 and 364 m. Elsewhere – on peb59. Monanchora alaskensis (Lambe, 1895)

bles, rocks, and sand at depths between 32 and 148 m 

and temperatures of 1.4 to 6.5°C.

Photo. 1) Specimen collected at a depth of 146 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
76 Professional Paper NMFS 12

Description. This sponge is stalked with a subglobular 

body and a corrugated surface in life which becomes 

conulose after freezing. Small circular oscula in depressions on the surface are only visible in live specimens. 

Consistency is elastic but firm. Total height is to 10 

cm; smooth stalk with a height of 4 cm. Color in life is 

golden yellow. 

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Ectosomal skeleton consists of a thick 

membrane of tangential brushes of smaller styles, 

fairly dense, and isochelae in between. Choanosome 

consists of thick, ascending polyspicular tracts of large, 

thick styles connected by shorter tracts of large styles. 

Ascending tracts (114–387 μm in diameter) support 

the surface tubercles and are visible to the unaided eye 

when dissecting the sponge. Spicules consist of megascleres – thin, small, ectosomal (sub)tylostyles (350–395 

× 9–11 μm), and choanosomal thick styles (840–1170 × 

34–42 μm). Two categories of microscleres are present – 

anchorate isochelae with a peculiar plate in the middle 

(22–25 μm) and thin sigmas (19–23 μm). 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Attached to boulders and cobbles at depths 

between 203 and 485 m.

Photos. 1) Preserved specimens (frozen) collected at 

a depth of 485 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimens as in photo 1 with 

other biota. 3) Same specimens as in photos 1 and 2 

(indicated by the white arrows) in situ.

60. Monanchora laminachela Lehnert, Stone and Heimler, 2006
77

Description. This sponge is flabellate with flattened, 

often coalesced branches. It is typically uniplanar in 

high current areas and multi-planed, even bushy, in 

areas of lower current. The surface is smooth and the 

consistency is elastic. Height and width are to at least 

50 cm. Color in life is orange, yellow, or brownish 

orange. 

Skeletal structure. Ectosomal subtylostyles are scattered without obvious orientation. There are ectosomal 

styles to subtylostyles (225–680 × 8–12 μm). Choano-

somal styles to subtylostyles are longer and thicker 

(670–990 × 28–40 μm). Isochelae are anchorate (18–22 

μm). Sigmas (16–20 μm) are typically present.

Zoogeographic distribution. Locally abundant. In 

Alaska – Aleutian Islands. Elsewhere – Pacific Coast of 

the Kuril Islands and Pacific Coast of Canada (British 

Columbia).

Habitat. In Alaska – attached to bedrock, boulders, 

cobbles, and pebbles at depths between 79 and 330 m. 

Elsewhere – attached to pebbles and rocks at depths 

between 87 and 232 m and at temperatures between 

1.8 and 5.3°C.

Remarks. May be preyed upon by the sea star Henricia

sp.

Photos. 1) Fragment of a specimen collected at a 

depth of 80 m in the central Aleutian Islands. Grid 

marks are 1 cm

2

. 2) Same specimen as in photo 1 in 

situ (center) with a lighter-colored specimen (right). 3) 

Specimen collected at a depth of 150 m in the central 

Aleutian Islands.

61. Monanchora pulchra (Lambe, 1894)
78 Professional Paper NMFS 12

Description. This sponge is polymorphic; may be tabular, irregularly compressed, occasionally massive-lobate 

or markedly lobate with an uneven, corrugated surface. 

The dermal membrane is thinly pellicular. Oscula are 

not visible on the surface. Dimensions are to 7 cm. Color 

in life is light to dark brown.

Skeletal structure. Ectosomal tornotes have 2–4 

pointed “teeth” at the ends (150–290 × 4–7 μm). There 

are choanosomal acanthostyles to acanthostrongyles 

(210–380 × 10–21 μm) and small anchorate isochelae 

(23–37 μm). Koltun (1959) additionally reports large 

anchorate isochelae (44–84 μm); these were lacking in 

specimens we examined from the Aleutian Islands.

Zoogeographic distribution. Uncommon. In Alaska – 

central Aleutian Islands. Elsewhere – Arctic Ocean (Barents Sea – White Sea, and Kara Sea) and North Atlantic 

Ocean (Norwegian Sea, Denmark Strait, Davis Strait). 

Habitat. In Alaska – attached to pebbles at a depth of 

190 m. Elsewhere – reported at depths between 20 and 

62. Crella brunnea (Hansen, 1885)

300 m, temperatures of 1.5 to 4.95°C, and salinities of 

27.68 to 35.01 psu.

Photo. 1) Fragmented specimen collected at a depth 

of 190 m in the central Aleutian Islands. Grid marks 

are 1 cm2.
79

Description. This sponge is small, thin, and encrusting. Oscula are not visible on the surface. Consistency 

is soft. Color in life is light orange; beige after preservation in ethanol. 

Skeletal structure. There are polytylote subtylostyles 

to subtylostrongyles (380–450 × 5–6 μm), erect on the 

substrate with the points facing out, and acanthostyles 

(430–470 × 10–12 μm).

Zoogeographic distribution. Widespread but uncom-

mon. In Alaska – Aleutian Islands. Elsewhere – Arctic 

Ocean (Barents Sea, Laptev Sea, Kara Sea – Vilkitsky 

Strait, and Greenland Sea) and North Atlantic Ocean 

(between Faroe Islands and Iceland). 

Habitat. In Alaska – attached to mollusk shells at a 

depth of 146 m. Elsewhere – attached to rocks and 

pebbles on sandy ooze at depths between 91 and 410 

m, water temperatures of 1.47 to 1.17°C, and salinities 

of 34.42 to 34.65 psu.

Photos. 1) Preserved specimen (in ethanol) attached 

to a green false jingle (Pododesmus macrochisma) shell 

collected at a depth of 146 m in the central Aleutian 

Islands. Grid marks are 1 cm

2

. 2) Close-up view of same 

specimen as in photo 1.

63. Hymedesmia (Stylopus) dermata Lundbeck, 1910
80 Professional Paper NMFS 12

Description. This sponge is thinly encrusting with a 

smooth or partially hispid surface. According to Koltun 

(1959) there are oscula atop broad papillae. Color in 

life is greyish yellow or bluish white.

Skeletal structure. There are acanthostyles (125–540 × 

16–32 μm). Ectosomal spicules are polytylote strongyles 

(280–410 × 6–10 μm) tangentially arranged in bundles. 

Microscleres are arcuate isochelae (32–48 μm).

Zoogeographic distribution. Widespread but uncommon. In Alaska – Bering Sea (Pribilof Canyon). Elsewhere – Arctic Ocean (White Sea) and North Atlantic 

Ocean (Davis Strait, Denmark Strait, southwest of Iceland near the Faroe Islands).

Habitat. In Alaska – grows on boulders and cobbles at 

depths around 300 m. Elsewhere – reported at depths 

between 293 and 1441 m.

64. Hymedesmia (Hymedesmia) irregularis Lundbeck, 1910

Photo. 1) Fragments of a preserved (frozen and then 

stored in ethanol) specimen collected at a depth of 300 

m in Pribilof Canyon, Bering Sea.
81

Description. In situ this species has a globular body 

covered with numerous papillae and possesses large 

circular oscula. On deck specimens typically collapse 

completely and appear to be irregularly massive-lobate 

without papillae and oscula. Surface is smooth and 

covered by a thin ectosomal membrane. Circular os-

cula on the surface are slightly elevated and close upon 

collection. The consistency is elastic and compressible. 

Diameter is to 10 cm. Color in life is yellow, orange, or 

golden brown. 

Skeletal structure. There are ectosomal tylotes (350–

440 × 4–7 μm); choanosomal styles include a few stron-

gyles (470–680 × 15–22 μm).

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – Northwest 

Pacific Ocean (previously known only from off Shikotan 

Island, Kuril Islands).

Habitat. In Alaska – attached to boulders and cobbles 

at depths between 82 and 426 m. Elsewhere – reported 

at depths between 472 and 479 m.

Remarks.K. borealis is similar to Inflatella globosa but 

is irregularly massive with relatively inconspicuous 

oscules, while the latter species is more globular with 

oscules on top of small papillae. Also, K. borealis has 

tylotes, styles, and a few strongyles while I. globosa has 

tylotes only. 

Photos. 1) Collapsed specimen collected at a depth of 

146 m in the central Aleutian Islands. Grid marks are 

1 cm

2

. 2) Same specimen as in photo 1 in situ.

65. Kirkpatrickia borealis Koltun, 1970
82 Professional Paper NMFS 12

Description. This sponge is globular with two distinct 

surface morphologies; some specimens are covered with 

round, crater-like depressions and round oscula that are 

flush with the surface, others have a surface covered 

with short papillae. Dermal membrane is thick. Consistency is soft and elastic. Diameter is to 20 cm. Color in 

life is bright orange (dominant phase) or yellow.

Skeletal structure. Anisotornotes (420–490 × 8–12 

μm) with long thin points form a dense ectosomal 

crust together with numerous isochelae. Choanosomal 

ascending plumose spicule tracts are of the same tornotes, cored and echinated by acanthostyles (280–540 

× 10–30 μm). Anchorate isochelae are strongly bent, 

the outline of a half circle and tridentate (32–40 μm). 

Koltun (1959) described somewhat smaller tornotes and 

acanthostyles.

Zoogeographic distribution. North Pacific Ocean. 

Locally common. In Alaska – central Aleutian Islands. 

Elsewhere – Sea of Okhotsk and Sea of Japan. 

Habitat. In Alaska – patchily distributed on bedrock 

and cobbles at depths between 56 and 120 m. Elsewhere 

– reported on rocks and ooze at depths between 3 and 

414 m and a temperature of 2.3°C.

Remarks. Gross morphology and spicule complement 

are similar to Lissodendoryx behringi, but the spicule architecture is quite different. 

Photos. 1) Specimen collected at a depth of 124 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ with the gorgonian Fanellia compressa. 3) Specimen collected at a depth 

of 110 m in the central Aleutian Islands. Grid marks are 

1 cm2. 4) Same specimen as in photo 3 in situ. 

66. Phorbas paucistylifer Koltun, 1958
83

Description. This sponge is globular. The surface is 

covered with numerous thin-walled, bulbous fistules, 

4–8 mm high and up to 4 mm in width; widest at the 

distal end. The ectosome is a translucent white layer, eas-

ily detachable, 400–650 μm thick. Diameter is to about 

10 cm. Color in life is light yellow with whitish, almost 

translucent fistules.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome has smooth tylotes 

partially arranged perpendicular to the surface but 

oriented in all directions to some degree. From the 

choanosome there are tracts of tylotes fanning out to-

wards the ectosome. Tylotes with acanthostyles are also 

found in the choanosome. Tylotes are 640–680 × 10–

12 μm. Choanosomal acanthostyles (660–670 × 20–

30 μm) are singly distributed without recognizable ori-

entation. Microscleres are three categories of isochelae: 

large isochelae with fimbriae (65–93 μm), large isoche-

lae of the same size but with dented outer margins, and 

small anchorate isochelae (23–25 μm). 

Zoogeographic distribution. Rare. In Alaska – central 

Aleutian Islands. Elsewhere – not reported.

Habitat. Attached to pebbles and shell hash at a depth 

of 173 to 190 m.

Photos. 1) Specimen collected at a depth of 190 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Specimen (indicated by the white arrow) observed at 

a depth of 179 m in the central Aleutian Islands.

67. Melonanchora globogilva Lehnert, Stone and Heimler, 2006
84 Professional Paper NMFS 12

Description. This sponge is massive and irregularly 

globular. Surface is smooth. Consistency is slightly elastic and rather firm due to the polyspicular tracts of the 

choansosome. Diameter is to at least 4–5 cm. Color in 

life is whitish to golden brown.

Skeletal structure. Tylotes (220–270 × 6–10 μm) are 

ectosomal. Choanosomal acanthostyles (250–380 × 

10–15 μm) occur in a meshwork of polyspicular tracts. 

There are anchorate large isochelae (70–80 μm), small 

isochelae (15–18 μm), large sigmas (65–78 μm), and 

small sigmas (27–32 μm).

Zoogeographic distribution. Widespread but rare in 

the North Pacific Ocean. In Alaska – central Aleutian 

Islands and Bering Sea. Elsewhere – Sea of Okhotsk, 

Sea of Japan, and British Columbia (Vancouver Island).

Habitat. In Alaska – encrusts other demosponges at 

depths between 190 and 195 m. Elsewhere – reported at 

depths between 32 and 104 m and a salinity of 33.95 psu. 

Remarks. In Alaska, the species is epizoic on the 

demosponge Halichondria oblonga. It was considered by 

Koltun (1958) as a subspecies of Myxilla incrustans. 

68. Myxilla (Myxilla) behringensis Lambe, 1895

Photo. 1) Specimen (center) collected at a depth of 

195 m in the central Aleutian Islands. Grid marks are 

1 cm2. Specimen is encrusting the demosponge Halichondria oblonga.
85

Description. This sponge is encrusting, typically on 

the shells of scallops (Chlamys sp.). Surface is finely 

structured with radiating canals leading to small oscula 

that are flush with the surface. Thin encrustation is 

less than 1 cm in height and typically covers the entire 

available shell surface. Color in life is light brown to 

golden brown.

Skeletal structure. Ectosomal tornotes have unequal 

ends and mammillate projections at these ends (160–

186 × 7–9 μm). Choanosomal acanthostyles are in an irregular meshwork (208–310 × 10–15 μm); microscleres 

are large isochelae (52–65 μm), small isochelae (15–20 

μm), and thin sigmata (17–25 μm).

Zoogeographic distribution. Locally abundant in the 

North Pacific Ocean. In Alaska – central Aleutian Is-

lands. Elsewhere – Bering Sea (Russia), Sea of Okhotsk 

(near the Kuril Islands) and British Columbia (near 

Vancouver Island). 

Habitat. In Alaska – encrusts scallop shells at depths 

between 98 and 250 m. Elsewhere – on rocks, pebbles, 

and sand at depths between 15 and 126 m.

Remarks. They are often very abundant in scallop 

beds.

Photos. 1) Specimen collected at a depth of 138 m 

in the central Aleutian Islands. Specimen completely 

encrusts a scallop (Chlamys sp.). Grid marks are 1 cm

2

. 

2) Specimen (center) collected at a depth of 98 m in 

the central Aleutian Islands. 3) A bed of scallops (Chla-

mys sp.), almost all of which are completed encrusted 

with Myxilla parasitica. The gorgonian Fanellia fraseri is 

at center. 

69. Myxilla (Ectyomyxilla) parasitica Lambe, 1893
86 Professional Paper NMFS 12

Description. This sponge is globular or lobate, usually 

with a stalk. The surface is slightly hispid. Irregularly 

scattered circular oscula are flush with the surface. Consistency is soft and elastic. Height and width are to 12 

cm. Color in life is yellow to bright yellow.

Skeletal structure. Spicules are arranged in long ascending fibers, connected by shorter spicule tracts and 

single spicules. There are smooth or slightly acanthose 

styles (270–551 × 10–20 μm), ectosomal tornotes (220–

380 × 6–10 μm), and microscleres that are anchorate 

isochelae (42–75 μm).

Zoogeographic distribution. Widespread but uncommon. In Alaska – Bering Sea (Pribilof Canyon). 

Elsewhere – Arctic Ocean (Kara Sea – Vilkitsky Strait, 

Laptev Sea) and North Atlantic Ocean (Iceland and 

Faroe Islands).

Habitat. In Alaska – attached to pebbles and cobbles 

at depths between 204 and 236 m. Elsewhere – reported 

at depths between 28 and 325 m.

Photos. 1) Specimen collected at a depth of 204 m 

in Pribilof Canyon, Bering Sea. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 showing the detail of 

the oscula. 3) Specimen in situ collected at a depth of 

236 m in Pribilof Canyon, Bering Sea. The demosponge 

Craniella spinosa is attached to the specimen. The separation between the red laser marks is 10 cm.

70. Myxilla (Burtonanchora) pedunculata Lundbeck, 1905
87

Description. This species exhibits two growth forms. 

One is a stalked, conical form with a ridged surface. 

The other is a massively encrusting, lobate sponge with 

a smooth surface. Consistency is rather hard and only 

slightly elastic, due to the densely packed spicules in 

the ectosome. The stalked specimen has a stalk of 2 × 

0.7 cm, and the body is 5 × 4 cm in greatest dimensions. 

Color in life for both growth forms is greenish yellow 

to light green.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Ectosome is densely packed with a tan-

gential arrangement of tornotes, peculiar shaped oxeas, 

and microscleres. The choanosome consists in places 

of an irregular reticulation of single spicules or short 

spicule tracts. Megascleres are oxeas (517–558 × 20–

30 μm); the points of oxeas have a ragged, dented 

outline. Tornotes have acanthose ends (230–270 × 9–

11 μm). Microscleres are large isochelae (54–110 μm), 

medium-sized isochelae (23–32 μm), small anchorate 

isochelae (9–13 μm), and thin centrotylote sigmas 

(8–12 μm). 

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – not reported.

Habitat. Attached to cobbles and pebbles at depths 

between 175 and 712 m.

Photos. 1) Specimen collected at a depth of 395 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Specimen (indicated by the white arrow) collected 

at a depth of 711 m in the central Aleutian Islands. 

The separation between the red laser marks is 10 cm. 

3) Specimen (top, center) collected at a depth of 175 m 

in the central Aleutian Islands. The separation between 

the red laser marks is 10 cm.

71. Stelodoryx oxeata Lehnert, Stone and Heimler, 2006
88 Professional Paper NMFS 12

Description. This sponge is stalked, flabellate, or a 

somewhat flattened funnel. Consistency is elastic with a 

thin ectosomal membrane. It is soft and floppy in situ. 

Height is to 17 cm and width to 12 cm. Color in life is 

light yellow; dark brown after freezing.

Skeletal structure. Ectosomal tylotes have acanthaceous ends (218–300 × 8–10 μm). Choanosomal styles 

(500–1140 × 21–30 μm) occasionally have acanthaceous 

heads, and anchorate isochelae are in two size categories – large (115–150 μm) and small (30–40 μm).

Zoogeographic distribution. Locally common in 

the North Pacific Ocean. In Alaska – central Aleutian 

Islands. Elsewhere – Sea of Okhotsk (near the Kuril 

Islands). 

Habitat. In Alaska – occurring in small patches (up to 

5 individuals per m2) on bedrock, mudstone, boulders, 

and cobbles at depths between 1714 and 2314 m. Elsewhere – found on sand, pebbles, and “reefs” at depths 

between 113 and 303 m and at temperatures between 

1.8 and 3.1°C.

Remarks. A sponge with similar appearance except 

larger (up to 50 cm diameter) and more robust (forming distinct bowls) occurs at depths between 1185 and 

1540 m. Collection of a specimen within this depth 

range and examination of the spicules is necessary to 

confirm conspecificity, however. 

Photos. 1) Preserved (frozen) specimen collected at 

a depth of 2176 m in the central Aleutian Islands. Grid 

marks are 1 cm2. 2) Same specimen as in photo 1 (lower 

center) in situ. 3) Close-up view of specimen in situ at a 

depth of 1505 m in the central Aleutian Islands.

72. Stelodoryx toporoki Koltun, 1958
89

Description. This sponge is tube-shaped, sometimes 

irregularly cylindrical, with a hollow center. It occurs 

in clusters of up to five individuals and is fragile and 

inelastic. Height is to 15 cm and width to 4 cm. Color 

in life is light yellow to golden brown.

Skeletal structure. Tangentially arranged tornotes 

have acanthaceous ends in the ectosome. There is an irregular meshwork of paucispicular tracts and individual 

acanthostyles in the choanosome. There are tornotes with 

acanthose ends (180–291 × 4–7 μm), acanthostyles (370–

520 × 20–29 μm), and anchorate isochelae (25–46 μm).

Zoogeographic distribution. Locally common in 

the North Pacific Ocean. In Alaska – central Aleutian 

Islands. Elsewhere – Bering Sea (near the Commander 

Islands) and the Sea of Okhotsk. 

Habitat. In Alaska – attached to cobbles and pebbles at 

depths between 155 m and 1009 m. Elsewhere – found 

on sand, pebbles, and “reefs” at depths between 115 

and 820 m.

Photos. 1) Specimen collected at a depth of 155 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Specimen collected at a depth of 1009 m in the cen-

tral Aleutian Islands. 3) Same specimen as in photo 2 

in situ.

73. Stelodoryx vitiazi (Koltun, 1959)
90 Professional Paper NMFS 12

Description. This sponge is short with a cylindrical 

body and hollow central cavity. Koltun (1970) reported 

that this species was stalked, but our observations indicate that it narrows near the base and does not necessarily have a true stalk. Surface is corrugated. Height is to 5 

cm. Color in life is light brown to light yellow.

Skeletal structure. There are two categories of styles: 

long, 1320 × 25 μm (Koltun: 660–1650 × 45–54 μm), and 

shorter, 350 × 9 μm (Koltun: 400–540 × 5–10 μm). It has 

isochelae (sigmancistras) 27–30 μm (Koltun: sigmoid 

arcuate chelae 18–27 μm) and sigmas 40 μm (Koltun: 

filiform sigmas 16–22 μm). We regard the differences in 

spicule sizes as intraspecific variation.

Zoogeographic distribution. Apparently a very rare 

species; known from only two locations. In Alaska – 

central Aleutian Islands. Elsewhere – Sea of Okhotsk. 

Habitat. In Alaska – occurring singly on cobbles and 

pebbles at depths between 665 and 711 m. Elsewhere – 

reported at depths between 1440 and 1540 m.

Photos. 1) Preserved (frozen) specimen collected at a 

depth of 711 m in the central Aleutian Islands. 2) Same 

specimen as in photo 1 (indicated by the white arrow) 

in situ on cobble.

74. Echinostylinos hirsutus Koltun, 1970
91

Description. This sponge is irregularly massive-lobate, 

often with dactylate processes bearing circular oscula on 

top. The surface is smooth. The consistency is soft and 

elastic. Height and width are to 20 cm. Color in life is 

yellow to golden yellow; brown after freezing. 

Skeletal structure. Styles are smooth (238–565 × 9–18 

μm). Ectosomal tylotes have slightly acanthose ends 

(215–487 × 5–8 μm). Microscleres are rhaphides in two 

size categories – large rhaphides (200–400 μm) and 

small rhaphides (45–180 μm).

Zoogeographic distribution. North and West Pacific 

Ocean. Uncommon. In Alaska – central Aleutian Islands 

and Bering Sea (Pribilof Canyon). Elsewhere – Sea of 

Okhotsk, Pacific Coast of Kuril Islands, and South China 

Sea.

Habitat. In Alaska – attached to cobbles at depths be-

tween 100 m and 341 m. Elsewhere – reported at depths 

between 4 and 550 m.

Remarks. Tedania dirhaphis was originally described 

by Hentschel (1912) from shallow Indonesian waters. 

Koltun (1959) reported the same species from the Sea 

of Okhotsk and the Kuril Islands in the North Pacific 

Ocean and the Alaskan specimens conform perfectly 

to Koltun’s decription. The Alaskan specimens also 

conform to the original description (p. 45 and pl. XIX, 

fig. 20) in spicule complement, growth form, and color 

(Hentschel, 1912) and differ from the Indonesian speci-

mens in size only (somewhat larger). Accordingly, we 

report this species as T. dirhapsis sensu Koltun, 1959, 

but caution that given the extremes in zoogeography, 

additional taxonomic work on this species is warranted. 

Photos. 1) Specimen collected at a depth of 100 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Preserved (frozen) specimen collected at a depth of 

341 m in Pribilof Canyon, Bering Sea. Grid marks are 

1 cm

2

. 3) Same specimen as in photo 1 in situ with a 

shortspine thornyhead (Sebastolobus alascanus) at left. 

The separation between the red laser marks is 10 cm.

75. Tedania (Tedania) dirhaphis Hentschel, 1912
92 Professional Paper NMFS 12

Description. This large flabellate sponge is attached to 

substrate with a firm stalk that widens about 10 cm above 

the holdfast. Surface is very smooth; oscula are in several 

rows on the flattened apical surface. Body is soft; stalk 

is wiry. Maximum dimensions of the roughly triangular 

body are approximately 30 × 30 × 5 cm. Color in life is 

light brown to orange-brown.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Ectosomal tylotes (300–330 × 4–6 μm) 

have acanthose ends. There are choanosomal smooth 

styles (360–390 × 15–20 μm), and onychaetes possibly 

in two size categories – large (170–210 μm) and small 

(30–40 μm). 

Zoogeographic distribution. Locally common. In Alaska – central Aleutian Islands. Elsewhere – not reported.

Habitat. Attached to bedrock, boulders, and cobbles 

at depths between 59 and 170 m. Specimens with a similar gross morphology have been observed at depths near 

478 m, but collection of a specimen within this depth 

range and examination of the spicules is necessary to 

confirm conspecificity.

Remarks. Species is similar in appearance to Artimisina 

stipitata with which it co-occurs. The dorsal surface of 

T. kagalaskai is typically flatter and the body form more 

distinctly triangular and with a much smoother surface 

than A. stipitata. This is the only known species of Tedania that is stalked.

Photos. 1) Specimen collected at a depth of 146 m in 

the central Aleutian Islands. Grid marks are 1 cm2

. 2) 

Same specimen as in photo 1 in situ. 3) Specimen collected at a depth of 155 m in the central Aleutian Islands. 

Grid marks are 1 cm2. 4) Specimen in situ collected at a 

depth of 105 m in the central Aleutian Islands. 

76. Tedania kagalaskai Lehnert, Stone and Heimler, 2006
93

Description. This sponge is flagelliform. This spe-

cies appears as described by Koltun (1959) and often 

consists of numerous more or less cylindrical branches 

diverging from one point and arranged in one plane. 

We have observed up to five branches that diverge at 

multiple points, however. Branches may be bent or 

slightly curved. Occasionally the sponge is markedly 

elongated and has a ramified, stalk-like appearance 

with many closely spaced fine projections (according to 

Koltun). Height is to at least 72 cm with branches up to 

1.5 cm thick. Color in life is light yellow, light orange, 

or creamy white; the lower portion is often devoid of 

the fine lateral projections and consequently a darker 

color, often brown.

Skeletal structure. Styles are arranged in a central 

axis with side-tracts running into the lateral projec-

tions; styles are 238–1500 × 7–18 μm and anisochelae 

are 10–17 μm.

Zoogeographic distribution. Widespread and locally 

abundant. In Alaska – central Aleutian Islands, eastern 

Gulf of Alaska (near the Fairweather Ground). Else-

where – North Pacific Ocean (Kuril Islands, Vancouver 

Island), Arctic Ocean (Southwest Barents Sea, Kara 

Sea – Vilkitsky Strait, Laptev Sea – Shokalsky Strait, East 

Siberian Sea, Greenland Sea, and Baffin Bay), and 

North Atlantic Ocean (Faroe Islands). 

Habitat. In Alaska – attached with a holdfast to bed-

rock, mudstone, boulders, cobbles, and occasionally 

hexactinellid sponge skeletons at depths between 395 

and 1812 m; more abundant at depths shallower than 

1200 m. Elsewhere – reported on ooze and sandy-ooze 

bottoms at depths between 41 and 1134 m and temperatures of 1.64 to 4.8°C.

Remarks. This species occasionally has ophiuroid as-

sociates particularly at depths shallower than 500 m. It 

occurs within the same depth range as Esperiopsis flagrum

but attaches to hard substrate rather than anchoring in 

soft sediment. A. ramosa may have a carnivorous feeding 

habit like other Cladorhizidae so collected specimens 

may have small crustaceans trapped in their outer tissues. 

Photos. 1) Specimen collected at a depth of 1715 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Specimen collected at a depth of 1501 m in the 

eastern Gulf of Alaska. 3) Specimen observed in situ 

at a depth of 843 m in the central Aleutian Islands. 4) 

Close-up view of same specimen as in photo 3 showing 

detail of the closely spaced fine projections.

77. Asbestopluma ramosa Koltun, 1958
94 Professional Paper NMFS 12

77. Asbestopluma ramosa Koltun, 1958 (continued)
95

Description. This stalked sponge terminates in a partially inverted conical head. Basal root-like processes 

consist of glassy, polyspicular tracts (2.5 cm long). Polyspicular tracts of the root-like processes become united 

at the base of the cylindrical stalk and are coated by a 

thin ectosomal veneer. The circular upper plane of the 

head has a diameter of 1.4 cm and thin appendages 

(up to 12) are inserted along its margin. The appendages measure 1.9 cm × 0.8 mm, but the diameter of 

the appendages regularly varies like a string of pearls. 

Typical of the genus, the polyspicular tracts of the 

system of root-like processes are united at the base of 

the stalk, run in bundles through the stalk, and then 

diverge again to support the appendages. The height 

from distal ends of the root-like processes to the upper 

end of the head is up to at least 12 cm. The stalk is up 

to 7 cm long and 1–2 mm in diameter. Color in life is 

white or creamy white.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. Ectosomal veneer is packed with anisochelae. Fusiform styles (980–2100 × 12–40 μm) and 

tylostrongyles (350–900 × 2–10 μm) are most abundant 

at the distal parts of the root-like processes. There are 

large anisochelae (55–75 μm), rare small anisochelae 

(25–35 μm), large sigmas (95–110 μm), small sigmas 

(43–55 μm), and flattened sigmoids with claw-like appendages at the ends (42–45 μm). Large sigmas were 

typically damaged in our SEM preparations indicating 

that they are extremely thin-walled, hollow, and fragile. 

Zoogeographic distribution. North Pacific Ocean. 

Locally common. In Alaska – central Aleutian Islands; 

patchy distribution at low densities. Elsewhere – Sea of 

Okhotsk.

Habitat. In Alaska – on soft sediments (silt and sand) 

at depths between 1108 and 2854 m. Elsewhere – eurybathic; reported at depths between 150 and 3800 m.

Remarks. C. bathycrinoides probably has a carnivorous 

feeding habit like other Cladorhizidae. We observed 

one specimen in situ capture a small shrimp with one 

of its appendages (actually the shrimp swam into the 

appendage and appeared to adhere to it). Collected 

specimens may have small crustaceans trapped in their 

outer tissues.

78. Cladorhiza bathycrinoides Koltun, 1955

Photos. 1) Specimen collected at a depth of 1371 m in 

the central Aleutian Islands and held live in a shipboard 

aquarium. Note the basal root-like processes. 2) Same 

specimen as in photo 1 in situ. The separation between 

the red laser marks is 10 cm.
96 Professional Paper NMFS 12

Description. Body consists of a long cylindrical stalk 

with a basal plate and two sets of distal appendages: 

the basal set radiating in a full circle more or less in 

one plane, and the distal set forming a quarter circle 

of triangular-shaped structures oriented in a plane 

almost perpendicular to the basal appendages. Distal 

appendages are shorter than the basal and terminate 

in a spherical tyle. Root-like processes (common within 

the genus) are absent and the sponge attaches directly 

to hard substrate via the broadened basal plate. Consistency is wiry and elastic, with an easily detachable 

ectosomal membrane. Total length is to at least 33 cm. 

Color in life is light yellow or creamy white. 

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The three distinct parts of the sponge 

(basal plate, stalk with the basal appendages, and the 

crown) differ considerably in spicule types and their 

arrangement. The basal plate consists of fusiform 

styles and shorter anisoxeas with slightly unequal ends, 

packed in one plane parallel to the surface of the substrate. The basal plate has no ectosomal membrane and 

is devoid of microscleres. From the basal plate polyspicular tracts of long fusiform styles run in thick bundles 

through the center of the long cylindrical stalk, which 

is covered by an ectosomal membrane, densely packed 

with ancorate anisochelae. The basal appendages are 

constructed in a similar manner as the stalk. The polyspicular tracts of long fusiform styles reaching the crown 

fan out in one plane, leaving little space between the 

tracts. Single, thin (sub-) tylostyles are arranged more 

or less perpendicular to the thick fusiform styles. The 

ectosomal membrane of the crown is devoid of anisochelae but rather contains flattened sigmancistras with 

thin claw-like extensions at each end. These latter two 

spicule types occur exclusively in the crown. Megascleres include fusiform styles, almost anisoxeas (600–4260 

× 10–65 μm), present in all parts except the ectosomal 

membrane; (sub-) tylostyles (510–1650 × 8–20 μm) 

present in the ectosomal membrane of the crown; and 

thick, short anisoxeas with slightly unequal blunt ends 

(140–660 × 38–43 μm) present in the basal plate. Microscleres include anchorate anisochelae (30–42 μm), 

present in the ectosomal membrane of the stalk and 

the basal appendages; and flattened sigmancistras, each 

end with a thin pointed “claw” (35–42 μm), present in 

the ectosomal membrane of the crown. 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

79. Cladorhiza corona Lehnert, Watling and Stone 2005

Habitat. Occurs in small patches at depths between 

726 and 2077 m on bedrock, fragmented bedrock, 

mudstone, and cobbles, and at temperatures between 

1.8 and 3.1°C.

Remarks. C. corona is carnivorous and feeds mainly 

on calanoid copepods (Watling, 2007). Collected specimens may have small crustaceans trapped in their outer 

tissues.

Photos. 1) Specimen collected at a depth of 1720 m 

in the central Aleutian Islands. Grid marks are 1 cm2. 

2) Same specimen as in photo 1 in situ.
97

Description. This stalked sponge has numerous lateral 

processes with terminal swellings (as many as 80). Size 

and growth form of this species is variable. There are 

root-like tufts of polyspicular tracts fixing the sponge 

in the substrate. The consistency of the stalks is firm 

and wiry while the rest of the sponge is soft and fragile. 

Thick polyspicular tracts support the stalk; thinner poly-

spicular tracts support the branches. Height is to 30 cm. 

Color in life is golden brown to light brown and reddish 

brown in shallower water (<1000 m depth).

Skeletal structure. There are styles (950–4300 × 23–65 

μm), accessory styles (550–880 × 10–27 μm), ectosomal 

styles to tylostyles (220–270 × 1–4 μm), occasionally 

acanthose isochelae in two size categories – large (55–

130 μm) and small (18–45 μm), and sigmas in two size 

categories – large (70–165 μm) and small (16–50 μm).

Zoogeographic distribution. Widespread and locally 

common. In Alaska – central Aleutian Islands. Else-

where – North Pacific Ocean (Sea of Okhotsk, Kuril 

Islands), Arctic Ocean (Greenland Sea), and North 

Atlantic Ocean (Norwegian Sea). 

Habitat. In Alaska – occurs in generally flat, soft-sedi-

ment habitats but occasionally on silt-covered bedrock 

slopes at depths between 563 and 1984 m. In shallower 

water (<1000 m depth) this sponge appears to attach di-

rectly to hard substrate. Elsewhere – eurybathic; depths 

between 200 and 8660 m.

Remarks. Chondrocladia are carnivorous, but unlike 

other Cladorhizidae they possess a modified aquiferous system used to inflate the terminal swellings where 

they capture prey (Vacelet, 2008). Collected specimens 

may have small crustaceans trapped in their outer 

tissues.

Photos. 1) Specimen collected at a depth of 1341 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ. 3) Specimen 

observed at 1206 m in the central Aleutian Islands.

80. Chondrocladia (Chondrocladia) concrescens (Schmidt, 1880)
98 Professional Paper NMFS 12

Description. This sponge is subglobular. Surface is 

covered with numerous papillae. Oscula are at the top 

of the sponge in clusters. It is very soft, fragile, and easily 

torn. Diameter is to 10 cm. Color in life is greenish yellow to light brown; brown or dark brown after freezing.

Skeletal structure. A thin ectosomal membrane contains only microscleres. There are styles (420–1450 

× 18–35 μm), sigmas possibly in two size categories 

(40–310 μm), rhaphides (60–125 μm), and spherules 

(10 μm). Long ascending and branching polyspicular 

tracts of styles in the choanosome have many spicules 

scattered between the tracts. 

Zoogeographic distribution. Widespread but uncommon. In Alaska – central Aleutian Islands. Elsewhere 

– North Pacific Ocean (Bering Sea – Russia), Arctic 

Ocean (Barents Sea and Greenland Sea), and North 

Atlantic Ocean (eastern Scotian Shelf east to Iceland 

and south to the Canary Islands; western Mediterranean 

Sea).

Habitat. In Alaska – attached to cobbles and pebbles 

at depths between 155 and 489 m. Elsewhere – reported 

at depths between 62 and 1800 m depth.

Photos. 1) Specimen collected at a depth of 155 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Preserved specimen (frozen) collected at a depth of 

489 m in the central Aleutian Islands. Grid marks are 1 

cm2. 3) Same specimen as in photo 2 (indicated by the 

white arrow) in situ. 

81. Biemna variantia (Bowerbank, 1858)
99

Description. This sponge is flagelliform or whip-like 

and occasionally bifurcated. The species is similar to 

Asbestopluma ramosa and Esperiopsis flagrum (differentiation requires careful examination). Rigid long, thin 

stalk is covered with thin, relatively short processes. 

It is basally attached in soft substrates with a rigid 

root-like system. Length is to at least 35 cm, but few 

specimens attain this size. Color in life is light yellow 

to creamy white.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome consists of densely 

arranged isochelas, underlain by parallel oriented 

fusiform styles with blunt ends and smaller tylostyles. 

Single tylostyles are placed perpendicular to the orientation of the styles and tylostyles. The choanosome 

is dominated by ascending polyspicular tracts of styles, 

tylostyles, and isochelae. Megascleres are blunt-ended 

fusiform styles (1310–1510 × 40–55 μm). Tylostyles 

(620–660 × 9–13 μm) often have the tyle subterminal 

and occasionally polytylote. Microscleres are isochelae (80–95 μm), placochelae (70–88 μm), and sigmas 

(9–25 μm). 

Zoogeographic distribution. Locally abundant. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Attached with a root-like system in unconsolidated sediments at depths between 1525 and about 

2200 m. Fairly abundant in some areas, reaching densities near 15 individuals per m2

.

Remarks. There are only four known species of Euchelipluma worldwide. E. elongata is by far the largest species 

in the genus. This species does not appear to harbor 

the ophiuroid associates that are so common with the 

whip-like pennatulacean corals present in the same 

depth range. It may occasionally be preyed upon by the 

82. Euchelipluma elongata Lehnert, Stone and Heimler, 2006

sea star Hippasteria. It does not co-occur with Esperiopsis

flagrum but does overlap in depth range with Asbestopluma ramosa. Unlike A. ramosa, however, it anchors in 

soft-sediment rather than attaching to hard substrate. 

Photos. 1) Specimen collected at a depth of 2161 m 

in the central Aleutian Islands. Grid marks are 1 cm2. 

2) Same specimen as in photo 1 in situ (foreground).
100 Professional Paper NMFS 12

Description. This sponge is subglobular and massive. 

Consistency is elastic, soft, and easy to tear. Oscula are 

small, circular, and slightly elevated on short conical 

projections. The surface is smooth to the unaided eye 

but microscopically hispid or brain-like with narrow 

convolutions. Diameter is to 15 cm. Color in life is yel-

low, orange, or dark brown.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. There is no specialized ectosomal skel-

eton; the dermal membrane consists of spicule brushes 

at the ends of short polyspicular tracts, ascending only 

a short distance from the choanosome. Sections parallel 

to the surface show the regularly spaced spicule brushes 

up to 300 μm in diameter. Short polyspicular tracts of 

styles make up a relatively dense but irregular meshwork 

in the choanosome, mesh-size 100–200 μm, with single 

spicules scattered in between. The short length of the 

tracts gives a halichondroid appearance throughout 

much of the choanosome. Megascleres are fusiform 

styles (330–400 × 7–10 μm). There are seven types of 

microscleres: large placochelae (105–115 μm); small 

placochelae (30–50 μm); large biplacochelae (35–52 

μm); small and relatively rare biplacochelae (13–16 μm 

in diameter); a rare, large category of isochelae (58-

65 μm); small sigmoid isochelae (10–13 μm) almost 

closed; and small sigmoids (5–9 μm). All spicule-types 

are distributed without an obvious pattern throughout 

the sponge. 

Zoogeographic distribution. North Pacific Ocean. 

Locally common. In Alaska – central Aleutian Islands. 

Elsewhere – previously known only from Cordell Bank, 

California.

Habitat. In Alaska – attached to cobbles at depths be-

tween 100 and 146 m. Elsewhere – reported at depths 

between 35 and 46 m.

Photos. 1) Specimen collected at a depth of 138 

m in the central Aleutian Islands. Grid marks are 1 

cm

2

. 2) Same specimen as in photo 1 in situ (center). 

3) Specimen (center) collected at a depth of 146 m in 

the central Aleutian Islands.

83. Guitarra abbotti Lee, 1987
101

Description. This sponge is cushion-shaped with a tu-

berculate surface; no oscula visible. Consistency is only 

slightly compressible and not very elastic. Diameter is to 

20 cm. Color in life is yellow to orange-brown.

Skeletal structure. Spicules are anisotylotes (230–700 

× 5–15 μm) often one end conspicuously more inflated 

than the other, large placochelae (70–90 μm), small 

placochelae (35–55 μm), and bipocilli (6–15 μm).

Zoogeographic distribution. Cosmopolitan and lo-

cally common. In Alaska – central Aleutian Islands. Else-

where – Pacific coast of southern Kuril Islands, North 

Atlantic Ocean, Indian Ocean, and Antarctica. 

Habitat. In Alaska – attached to small boulders and 

cobbles at a depth of 146 m. Elsewhere – reported on 

rocky bottoms at depths between 28 and 188 m and at 

temperatures between 1.5 and 16.2°C.

Photos. 1) Partial specimen collected at a depth of 146 

m in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ (center).

84. Guitarra fimbriata Carter, 1874
102 Professional Paper NMFS 12

Description. This species appears as described by 

Koltun (1959) and may be massive-lobate, laminate, 

dactylate, funnel-shaped, or vase-like, and usually has a 

stalk. The surface is smooth. The consistency is elastic, 

easily compressible, but difficult to tear. Vase-like forms 

to 10 cm in height; funnel-shaped forms to more than 

60 cm in diameter and 30 cm in height. Color in life is 

light grey, creamy white, yellow or brown.

Skeletal structure. Spicules are styles (130–280 × 8–19 

μm) and palmate isochelae (12–28 μm).

Zoogeographic distribution. Widespread and locally 

common. In Alaska – central Aleutian Islands and eastern Gulf of Alaska. Elsewhere – North Pacific Ocean 

(Kamchatka Coast, Bering Sea, British Columbia), and 

Arctic Ocean (Chukchi Sea and East Siberian Sea).

Habitat. In Alaska – central Aleutian Islands (attached 

to pebbles at a depth of 100 m), eastern Gulf of Alaska 

(attached to bedrock, boulders, and cobbles at depths 

between 152 and 218 m). Elsewhere – reported at 

depths between 9 and 291 m.

Photos. 1) Specimen collected at a depth of 100 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ (center). 3) 

Specimen collected at a depth of 165 m in the eastern 

Gulf of Alaska. Grid marks are 1 cm2. 4) Same specimen 

as in photo 3 in situ. 

85. Amphilectus digitatus (Miklucho-Maclay, 1870)
103

Description. This sponge has a flagelliform or cylindrical growth form; similar to Asbestopluma ramosa and 

Euchelipluma elongata (differentiation requires careful 

examination). Occasionally it is bifurcated. A central 

long thin axis tapers near the tip and is covered with 

many thin processes. Processes are 3–5 mm in length, 

elastic but somewhat stiff. The central axis is up to 54 cm 

in length with a diameter of 4 mm at the base and 2.5 

mm at the tip. This cylindrical axis is compressible, elastic but resilient. Color in life is beige to creamy white. 

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. No special ectosome is developed. The 

central axis consists of interwoven polyspicular tracts of 

styles with many microscleres in between. The lateral 

processes are supported by single polyspicular tracts 

running into them. The outermost layer of the processes and the longitudinal axis consists of densely arranged 

microscleres. Megascleres are fusiform styles (980–1320 

× 20–28 μm). Microscleres are large palmate isochelae 

(98–112 μm), small palmate isochelae (28–43 μm), 

large sigmas (37–48 μm), and small sigmas (17–20 μm). 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not reported.

Habitat. Occurs at depths between about 700 and 

1389 m. Appears to bury with root-like processes in soft 

unconsolidated sediments.

Remarks. The lower portion of the stalk is often fouled 

with hydroids. The species occurs within the same depth 

range as Asbestopluma ramosa, but occurs in soft-sediment 

habitats rather than attaching to hard rock. It does not 

co-occur with Euchelipluma elongata.

86. Esperiopsis flagrum Lehnert, Stone and Heimler, 2006

Photos. 1) Mid-section of a specimen collected at a 

depth of 954 m in the central Aleutian Islands. Grid 

marks are 1 cm2. 2) Same specimen as in photo 1 in situ.
104 Professional Paper NMFS 12

Description. This stalked sponge has two distinct 

forms: 1) several small funnels or cups (up to at least 

20 cups per individual) that branch initially from a 

central stalk, and 2) a long, thin stalk (30 cm or more 

in length) that terminates in a single cup-shaped body, 

rarely two, with a maximum diameter of approximately 

25 cm. The stalk is wiry, firm but elastic; the body(s) of 

softer consistency. Surface is smooth to the unaided eye 

but microscopically hispid. Total height and width are 

to 70 cm (multi-cupped form). Color in life is golden 

brown to light brown. 

Skeletal structure. There is no specialized ectosomal 

skeleton; dermal membrane consists of the ends of ascending polyspicular tracts. Choanosomal polyspicular 

ascending tracts are quite conspicuous and cm-long 

fibers are easy drawn from the sponge. Tracts are connected by single spicules, styles in a very wide size range 

(75–650 × 6–15 μm).

Zoogeographic distribution. Widespread but uncommon. In Alaska – central Aleutian Islands, Bering Sea 

(Zhemchug Canyon), and Gulf of Alaska. Elsewhere 

– widely distributed along the north and east coasts 

of Russia (Bering and Chukchi Seas), Arctic Ocean 

(Greenland Sea), and North Atlantic Ocean (Norwegian Sea).

Habitat. In Alaska – the two forms appear to be ecomorphs. The multi-cupped form is attached primarily 

to bedrock (occasionally cobbles) at depths between 

80 and 270 m in low to moderate current areas. The 

single-cupped form is found in high current areas. 

Aleutian Islands – patchy distribution on moderately 

sloped sandy habitats; attached to pebbles with rootlike processes at depths between 99 and 306 m. Bering 

Sea – rare; pebble and sand slopes at depths around 

170 m. Elsewhere – reported at depths between 14 and 

325 m, temperatures between 1.9 and 7.0°C, and salinities between 29.81 and 35.23 psu.

Remarks. This species may be preyed upon by the 

blood star (Henricia sp.) and appears to be particularly 

fragile.

Photos. 1) Fragment of a specimen collected at a 

depth of 100 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 in situ 

(center). 3) Specimen collected at a depth of 97 m in 

the central Aleutian Islands. 4) Specimen collected at a 

depth of 170 m in Zhemchug Canyon, Bering Sea. Note 

that the specimen has a single large cup that was apparently two smaller cups now fused together. 5) Specimen 

87. Semisuberites cribrosa (Miklucho-Maclay, 1870)

collected at a depth of 139 m in the central Aleutian 

Islands. Note that the cup has been sliced open for 

examination. Grid marks are 1 cm2. 6) Same specimen 

as in photo 5 in situ (top half). 7) Same specimen as in 

photo 5 in situ (bottom half). 
105

87. Semisuberites cribrosa (Miklucho-Maclay, 1870) (continued)
106 Professional Paper NMFS 12

Description. This massive or thickly encrusting sponge 

has a bulbous or even fistulose surface. Oscula are 

not visible. Consistency is soft and elastic. Size is to 20 

cm in all dimensions. Color in life is yellow to golden 

brown.

Skeletal structure. It has ectosomal reticulation of 

polyspicular tracts with microscleres in-between tracts. 

There is a choanosomal mesh of polyspicular tracts, often branching. There are tylostyles (350–440 × 5–12 μm), 

anisochelae arranged differently in two size classes – 

large anisochelae in rosettes (75–100 μm) and small 

anisochelae (18–50 μm); sigmata (45–58 μm), and 

rhaphides single and in trichodragmata (45–90 μm).

Zoogeographic distribution. Widespread and common. In Alaska – central Aleutian Islands and Bering 

Sea. Elsewhere – North Pacific Ocean (Sea of Okhotsk, 

Sea of Japan, Vancouver Island) and Arctic Ocean 

(Greenland Sea). 

Habitat. In Alaska – encrusts all hard substrates including hydrocoral skeletons at depths between 104 and 

442 m. Generally found in rough, steep-sloped habitats. 

Elsewhere – on rocks, pebbles, and sand bottoms from 

the intertidal zone to a depth of 270 m.

Remarks. This species encrusts hydrocorals (Stylaster

sp. and Cyclohelia lamellata) and other sedentary biota 

in Aleutian Island coral gardens; inaccurately reported 

in Stone (2006) as Myxilla incrustans.

Photos. 1) Fragment of a specimen collected at a 

depth of 190 m in the central Aleutian Islands. Specimen is encrusting the hydrocoral Cyclohelia lamellata. 

Grid marks are 1 cm2

. 2) Same specimen as in photo 

1 (center) in situ. 3) Specimen collected at a depth of 

109 m in the central Aleutian Islands.

88. Mycale (Aegogropila) adhaerens (Lambe, 1893)
107

Description. This clavate sponge typically has a sin-

gle cylinder, occasionally several (up to four) smaller 

branches, atop a thinner well-defined stalk. One or 

more large oscula are atop each cylinder; several smaller 

oscula along sides are obvious in situ. Surface is smooth. 

Consistency is very soft, except for the wiry stalk. Height 

is to 50 cm or more. Color in life is golden brown.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The stalk consists of longitudinally 

arranged masses of tylostyles; individual tracts are not 

recognizable. A special ectosome is not developed. The 

choanosome consists of irregularly arranged pauci- and 

polyspicular tracts of tylostyles with masses of sigmas and 

very abundant anisochelae, both single and in rosettes. 

Spicules consist of tylostyles (470–520 × 10–14 μm), an-

isochelae (55–75 μm), and sigmas (65–80 μm). 

Zoogeographic distribution. Locally abundant (up 

to 8 individuals per m

2

). In Alaska – central Aleutian 

Islands. Elsewhere – not reported.

Habitat. Attached to bedrock, boulders, cobbles, and 

pebbles at depths between 82 and 360 m.

Remarks. Mycale loveni and M. bellabellensis are the 

only other species in the genus that exhibit a stalked 

growth form.

Photos. 1) Specimen collected at a depth of 150 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ. 3) Specimen 

with several M. loveni at a depth of 112 m. 4) Specimen 

with several Halichondria oblonga (upper right) at a 

depth of 119 m. 

89. Mycale (Carmia) carlilei Lehnert, Stone and Heimler, 2006
108 Professional Paper NMFS 12

Description. This massive sponge has several large 

tubes basally connected. In situ, several exhalent canals 

flow into large circular oscula surrounded by thin walls. 

These oscula collapse and are not readily visible on collected specimens. The surface is bulbous with many irregularly distributed conical processes. The consistency 

is rather soft, easily torn, and fibrous. Diameter is to 25 

cm. Color in life is yellow to light yellow.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome is a tangential arrangement of short spicule tracts and single spicules with 

many microscleres in between the tracts. The choanosome consists of rather short spicule tracts (60–95 μm 

in diameter) that frequently branch off side tracts in 

all directions. This pattern is obscured by the presence 

of many single mega- and microscleres in between the 

tracts and without obvious orientation. Megascleres are 

tylostyles (405–460 × 10–12 μm). Microscleres include 

large anisochelae (80–100 μm), small anisochelae 

(40–60 μm), and rhaphides (42–65 μm). 

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – not reported.

Habitat. Attached to cobbles at depths between 178 

and 340 m.

Remarks. M. jasoniae is quite similar to M. loveni but 

possesses rhaphides that the latter species lacks.

Photos. 1) Preserved (frozen) specimen collected at 

a depth of 208 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 in situ

with a sculpin (Malacocottus sp.). 3) Close-up of same 

specimen as in photos 1 and 2.

90. Mycale (Mycale) jasoniae Lehnert, Stone and Heimler, 2006
109

Synonym: Mycale (Carmia) bellabellensis (Lambe, 1905)

Description. This sponge is polymorphic; massively 

encrusting or stalked. The consistency is fragile, inelastic but difficult to tear against the direction of the long 

polyspicular tracts (laterally); easier to tear parallel to 

the tracts (top to bottom). Aleutian specimens may 

be stalked vases to 20 cm in height or massive forms 

to 1 m high and wide. Color in life varies from yellow 

to greenish yellow to brown. Gulf of Alaska specimens 

have a cone-shaped body with a slender stalk and are 

club-shaped until approximately 8 cm high or wide and 

then develop the characteristic cone shape. Cones may 

reach 1 m or more in height and diameter. Color in life 

varies from yellow to light yellow.

Skeletal structure. Very long ascending polyspicular 

tracts (visible to the unaided eye) lie below a reticulate 

dermal skeleton. Tracts are branching with thick tracts 

connected by shorter tracts. Tylostyles (370–495 × 10–15 

μm) often have the largest diameter just before the 

point. Anisochelae are large (80–110 μm) and small 

(30–42 μm).

Zoogeographic distribution. Widespread and locally abundant. In Alaska – Chukchi Sea, Bering Sea, 

Aleutian Islands, and Gulf of Alaska. Elsewhere – Sea of 

Okhotsk, Pacific Coast of the Kuril Islands, Chukchi Sea 

(Russia), Arctic Ocean (East Siberian Sea), and British 

Columbia.

Habitat. Central Aleutian Islands – typically attached 

to boulders, cobbles, and pebbles at depths between 56 

and 744 m (massive form) and 171 to 191 m (stalked 

form). Bering Sea (Pribilof Canyon) – uncommon; attached to cobbles and pebbles at depths between 260 

and 309 m. Eastern Gulf of Alaska – most specimens are 

attached to bedrock, boulders, and cobbles at depths 

between 143 and 289 m in the eastern Gulf of Alaska 

91. Mycale (Mycale) loveni (Fristedt, 1887)

(and presumably much deeper based on unconfirmed 

catches with longline gear). Elsewhere – no information 

is available.

Remarks. We consider M. loveni and M. bellabellensis to 

be the same species, the latter being a cone-shaped ecomorph adapted for low-current habitats in the Gulf of 

Alaska. The two species have the same spicule complement and arrangement and internal skeletal structure. 

In the eastern Gulf of Alaska the species is club-shaped 

until approximately 8 cm wide and then it develops 

the characteristic cone shape. The sharpchin rockfish 

(Sebastes zacentrus) and other rockfish species often use 

this sponge as perching habitat. This species is preyed 

upon by the sea star Ceramaster patagonicus. 

Photos. 1) Fragment of a specimen (massive form) 

collected at a depth of 309 m in Pribilof Canyon, Bering 

Sea. Grid marks are 1 cm2. 2) Several specimens (massive form) at a depth of 96 m in the central Aleutian 

Islands. 3) Several specimens (massive form) with a 

sharpchin rockfish (Sebastes zacentrus) at a depth of 119 

m in the central Aleutian Islands. 4) Preserved (frozen) 

specimen (stalked form) collected at a depth of 192 m 

in the central Aleutian Islands. Grid marks are 1 cm2. 

5) Same specimen as in photo 4 in situ. The separation 

between the red laser marks is 10 cm. 6) Specimen collected at a depth of 167 m in the eastern Gulf of Alaska. 

Grid marks are 1 cm2. 7) Juvenile specimen (with associated euphasid) collected at a depth of 165 m in the 

eastern Gulf of Alaska. Grid marks are 1 cm2. 8) Same 

specimen as in photo 7 in situ. 9) Specimen with a gravid 

sharpchin rockfish (S. zacentrus) at a depth of 170 m in 

the eastern Gulf of Alaska. Photo by J. Lincoln Freese 

(AFSC). 10) A more robust cone-shaped form with a 

sharpchin rockfish (S. zacentrus) in a higher current 

area at a depth of 179 m in the eastern Gulf of Alaska. 

Note the juvenile specimen at the upper left. Photo by 

J. Lincoln Freese (AFSC). 
110 Professional Paper NMFS 12

91. Mycale (Mycale) loveni (Fristedt, 1887) (continued)
111

91. Mycale (Mycale) loveni (Fristedt, 1887) (continued)
112 Professional Paper NMFS 12

Description. This sponge is massive with a smooth 

but convoluted surface. Circular oscula are on conical 

elevations. Consistency is inelastic and very easily torn. 

Diameter is to 10 cm. Color in life is yellow to golden 

brown.

Skeletal structure. There is a thick ectosomal crust of 

styles to tylotes that are scattered without obvious orientation and tangentially arranged with many microscleres 

in between. Choanosomal pauci- to polyspicular tracts 

are in triangular meshes with masses of sigmas in between. Rosettes of large anisochelae are less abundant. 

There are ectosomal styles to tylotes (415–570 × 10–16 

μm), choanosomal styles (670–890 × 15–25 μm), large 

anisochelae (95–112 μm), small anisochelae (15–35 

μm), large sigmas (82–105 μm), small sigmas (25–35 

μm), and rhaphides (70–110 μm).

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – southern 

Kuril Strait.

Habitat. In Alaska – attached to cobbles and pebbles 

at depths between 135 and 175 m. Elsewhere – reported 

at depths between 73 and 181 m.

Photos. 1) Specimen collected at a depth of 172 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 (indicated by the white 

arrow) in situ. 

92. Mycale (Mycale) tylota Koltun, 1958
113

Description. Globular sponge has numerous small oscula on short, wart-like elevations. Water jets are emitted 

through the oscula when the sponge is squeezed. Areolate pore-fields collapse upon collection and are only 

visible in situ. Consistency is rather firm, only slightly 

elastic and difficult to tear. Diameter is to at least 16 cm. 

Two distinct colors have been observed in situ – dark 

brown (dominant) and olive-green. 

Skeletal structure. Ectosome is thick and leathery, 

consisting of a cortex of tightly packed anisodiscorhabds. 

Fusiform, slightly sinuous styles have acanthose heads 

(320–525 × 9–15 μm) and anisodiscorhabds (42–50 μm).

Zoogeographic distribution. Locally common and 

abundant (densities up to 14 individuals per m2 in 

Alaska). In Alaska – central Aleutian Islands. Elsewhere 

– previously known only from the Kuril Islands in the 

Sea of Okhotsk.

Habitat. Attached to bedrock, boulders, cobbles, and 

pebbles at depths between 79 and 288 m (possibly to 

93. Latrunculia (Biannulata) oparinae Samaii and Krasokhin, 2002

438 m). Elsewhere – reported at depths between 176 

and 202 m.

Remarks. Sponges in the genus Latrunculia contain 

the cytotoxic discorhabdin class of pyrroloiminoquinone alkaloids that exhibit significant antiviral activity 

against hepatitis virus C (HCV), antimalarial activity 

against Plasmodium falciparum, and antimicrobial effects 

against the AIDS opportunistic pathogens methicillinresistant Staphylococcus aureus (MRSA), Mycobacterium 

intracellulare, and M. tuberculosis (Na et al., 2010). This 

species is associated with several gorgonians, including 

Thouarella spp. and Plumarella spp.

Photos. 1) Specimen collected at a depth of 150 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 (right) in situ with the 

olive-green morph (left). 3) Specimen collected at a 

depth of 146 m in the central Aleutian Islands. Grid 

marks are 1 cm2. 4) Same specimen as in photo 3 (center) in situ.
114 Professional Paper NMFS 12

Description. This cone-shaped sponge has a flattened 

smooth top that is typically circular but occasionally 

kidney-shaped. On deck, specimens appear subglobular because they have been laterally compressed. This 

species has a smooth uneven surface, thick and leathery, only slightly elastic, and easily torn. The interior is 

markedly fibrous, somewhat similar to L. oparinae but 

differing slightly in shape and clearly in the absence 

of areolate pore-fields and the form of the anisodiscorhabds. Diameter is to about 12 cm. Color in life is 

dull brown; dark brown on deck.

Skeletal structure. SEM images of spicules are shown 

in Appendix IV. The ectosome is a unispicular layer of 

discorhabds, all arranged with their longitudinal axis 

perpendicular to the surface. The choanosome is a 

reticulation of polyspicular tracts of styles with some discorhabds in between. Megascleres are styles with slightly 

acanthose heads (500–540 × 9–11 μm). Microscleres are 

relatively smooth anisodiscorhabds (37–43 μm). 

Zoogeographic distribution. Locally common. In 

Alaska – central Aleutian Islands. Elsewhere – not 

reported.

Habitat. Attached to bedrock, boulders, and cobbles 

at depths between 412 and 1009 m, but relatively rare 

at depths shallower than 600 m.

Remarks. There appears to be another species of 

Latrunculia in the central Aleutian Islands at intermediate depths (i.e., slightly overlapping the depth ranges 

of both L. oparinae and L. velera). Latrunculia specimens 

in the depth range of 200 to 500 m should be a priority 

for collection.

Photos. 1) Preserved (frozen) specimen collected at 

a depth of 1009 m in the central Aleutian Islands. Note 

that the specimen has been laterally compressed. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 (far 

left) in situ. 3) Specimen observed in situ at a depth of 

929 m. The separation between the red laser marks is 

10 cm.

94. Latrunculia velera Lehnert, Stone and Heimler, 2006
115

Description. This globular sponge has broad cratershaped pore fields; these are prominent structures 

on the sponge surface. Surface is uneven and slimy. 

Diameter is to about 8 cm. Color in life is khaki-green 

to olive-green.

Skeletal structure. The majority of megascleres are 

anisostyles, fusiform, not terminally spined, rarely 

polytylote, with only slight differentiation between 

the two ends (325–397 × 7–10 μm). Microscleres are 

anisodiscorhabds (49–66 × 4–7 μm). The manubrium, 

a base of six short, downward pointing smooth spines, 

is closely followed by the basal whorl, which consists of 

a ring of horizontally aligned, smooth spines. The me-

dian whorl (21–31 μm in diameter) is located midway 

along the shaft and composed of undulate petals with 

denticulate margins, and some sculpted regions. The 

subsidiary whorl is composed of similar undulating petals with denticulate margins and sculpted sections, but 

the petals slant upwards and are located just underneath 

the apical whorl. The apical whorl is formed of fused 

undulating petals with denticulate margins forming a 

beautiful corona.

Zoogeographic distribution. Locally abundant. In 

Alaska – continental shelf off Cape Ommaney, Baranof 

Island, eastern Gulf of Alaska. Elsewhere – a sponge, 

believed to be this species, has also been reported from 

British Columbia and northern Washington State.

Habitat. In Alaska – attached to bedrock at depths 

between 69 and 210 m, temperatures between 5.5 and 

7.6°C, and salinities between 32.2 and 33.7 psu. Else-

where – no information available.

Photos. 1) Specimen collected at a depth of 102 m 

in the eastern Gulf of Alaska. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ with the hydro-

coral Distichopora borealis (right).

95. Latrunculia sp. (undescribed) 
116 Professional Paper NMFS 12

Description. This sponge has irregular branching 

and is extremely arborescent. Consistency is moderately firm and elastic. Surface of the side branches is 

smooth but microscopically hispid. Size is to at least 

50 cm × 50 cm. Color in life is highly variable ranging 

from light brown, light grey, light yellow, golden-yellow 

to light orange. Light brown is the dominant color 

in waters deeper than 120 m in the central Aleutian 

Islands; golden yellow and light orange are dominant 

in shallower water.

Skeletal structure. Skeletal architecture consists of 

ascending spicule tracts; spicules protruding from these 

tracts cause the hispid surface of the ectosome. Spicules 

are long oxeas (850–1350 × 18–35 μm) and short styles 

(320–530 × 8–15 μm).

Zoogeographic distribution. Locally abundant. In 

Alaska – central Aleutian Islands and Bering Sea. Elsewhere – Bering Sea near Mednyi Island (Commander 

Islands, Russia).

Habitat. In Alaska – forms dense fields almost completely covering the seafloor in areas of bedrock, boulders, and cobbles on steep slope habitat and at depths 

between 80 and 269 m. Elsewhere – reported at depths 

to 160 m.

Photos. 1) Specimen collected at a depth of 140 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 in situ with the gorgonian Plumarella sp. (right). 3) Specimen observed in situ 

at a depth of 92 m in the central Aleutian Islands.

96. Axinella blanca Koltun, 1959
117

Description. This sponge is clavate. Branching, fanshaped, or single tubes (occasionally two tubes) are to 

17 cm in height. Consistency is firm but elastic, with a 

hispid surface. Color in life is grey, light yellow, golden 

brown to brownish red.

Skeletal structure. Axinellid spicule tracts consist of 

styles, strongyles, and oxeas. All spicules have a large 

variation in length up to 1750 μm.

Zoogeographic distribution. Widespread and locally 

abundant. In Alaska – central Aleutian Islands, Bering 

Sea, and eastern Gulf of Alaska. Elsewhere – North 

Pacific Ocean (Sea of Japan), Arctic Ocean (Barents 

Sea and Greenland Sea), and North Atlantic Ocean 

(Norwegian Sea).

Habitat. In Alaska – attached to hard substrate at 

depths between 87 and 712 m. Densities to 63 individu-

als per m

2

 in the eastern Gulf of Alaska. Elsewhere – re-

ported at depths between 90 and 320 m, temperatures 

between 2 and 5°C, and salinities between 34.20 and 

35.01 psu.

Remarks. Though superficially similar to Mycale 

carlilei, A. rugosa is a hollow tube whereas M. carlilei is a 

solid tube with a flat top and one or more prominent 

oscula. The Alaskan specimens are the first tube forms 

reported for the species. 

Photos. 1) Specimen collected at a depth of 172 m 

in the eastern Gulf of Alaska. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 in situ. 3) Specimen (in-

dicated by the white arrow) collected in situ at a depth 

of 712 m in the central Aleutian Islands. 

97. Axinella rugosa (Bowerbank, 1866)
118 Professional Paper NMFS 12

Description. This sponge is encrusting, elongated, foliate or lobate. Consistency is soft with a hispid surface. 

Dimensions are to 15 cm2. Color in life is bright red to 

golden brown.

Skeletal structure. It has characteristic axinellid spicule tracts. Styles are 426–4550 × 10–38 μm; strongyles 

and oxeas are irregularly curved or distorted (208–588 

× 6–15 μm).

Zoogeographic distribution. Widespread and locally 

common. In Alaska – central Aleutian Islands. Elsewhere – Arctic Ocean (Barents and Greenland Seas), 

North Atlantic Ocean (Norwegian Sea to the Azores), 

and Mediterranean Sea.

Habitat. In Alaska – encrusts mudstone, cobbles, and 

pebbles at depths between 165 and 888 m. Elsewhere – 

reported at depths between 9 and 1360 m.

Remarks. This is the same species as Axinella vermiculata sensu Koltun, 1959. 

Photos. 1) Specimen (center) collected with the 

gorgonian Plumarella aleutiana at a depth of 165 m in 

the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Specimen (right) collected on mudstone at a depth 

of 888 m in the central Aleutian Islands. 

98. Bubaris vermiculata (Bowerbank, 1866)
119

Description. This species appears as originally described and is irregularly cup-shaped and somewhat 

compressed. Large osculum is on the inside of the cup. 

The surface is smooth on the outside and hispid on the 

inside of the cup. The consistency is firm. Width is to 50 

cm and height to 30 cm. Color in life is greyish brown 

to creamy white. 

Skeletal structure. Smaller oxeas have tangential 

arrangement in the ectosome and larger oxeas are in 

characteristic confused “halichondroid” arrangement 

in the choanosome. Oxeas in two size categories range 

in length from 140–2000 μm.

Zoogeographic distribution. Rare. In Alaska – Bering Sea (Pribilof Canyon). Elsewhere – North Atlantic 

Ocean (Denmark Strait). 

Habitat. In Alaska – attached to cobbles at a depth of 

300 m. Elsewhere – reported at a depth of 1039 m.

Remarks. The specimen collected in Pribilof Canyon 

is only the second record for the species worldwide 

and the first from the Pacific Ocean. Several specimens 

collected from the Atlantic Coast of the U.S. (Massa99. Halichondria (Halichondria) colossea Lundbeck, 1902

6 Van Soest, Rob. 2008. Personal commun. Zoological Museum, University of Amsterdam, Amsterdam, The Netherlands 1090 GT.

chusetts) have been tentatively identified as H. colossea

(Van Soest6). 

Photo. 1) Fragment of a specimen collected at a depth 

of 300 m in Pribilof Canyon, Bering Sea. 
120 Professional Paper NMFS 12

Description. This cylindrical, tube-shaped sponge 

has a narrow stalk. It may branch near the base of the 

stalk so that several tubes (up to 12) may be connected 

basally. Consistency is only slightly elastic and stiff. Near 

the base the tubes are 3–4 mm in diameter and widen 

to a maximum diameter of 2 cm. Maximum height is to 

18 cm. Color in life is golden brown with a light, almost 

whitish tip. 

Skeletal structure. The brown part of the sponge 

is more stiff and covered with an ectosomal layer of 

densely packed, parallel arranged tangential oxeas. The 

whitish tip of the sponge is covered by a thin dermal 

membrane without spicules. Choanosomal ascending 

paucispicular tracts are connected by single spicules. 

Oxeas are 320–480 × 14–20 μm.

Zoogeographic distribution. Widespread and locally abundant. In Alaska – central Aleutian Islands. 

Elsewhere – Arctic Ocean (Barents Sea and Kara Sea) 

and North Atlantic Ocean (Greenland and the Faroe 

Islands).

Habitat. In Alaska – patchy distribution; attached 

to hard substrate (generally cobbles and pebbles) at 

depths between 115 and 305 m. Elsewhere – reported 

at depths between 18 and 823 m.

Photos. 1) Specimens collected with the demosponge 

Myxilla behringensis (left center) at a depth of 195 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimens as in photo 1 in situ. 3) Cluster of 

specimens (lower center) below the golden king crab 

(Lithodes aequispina) at a depth of 272 m in the central 

Aleutian Islands. 

100. Halichondria (Halichondria) oblonga (Hansen, 1885)
121

Description. This sponge has a short stalk that typi-

cally branches into several agglutinated tubes. Consistency is inelastic and stiff due to a combination of high 

spicule density and lack of spongin, but easy to tear. 

Oscula are not obvious. Height is to at least 15 cm. Color 

in life is light brown, golden brown, or creamy white.

Skeletal structure. Spicules are arranged tangentially 

in the ectosome as is typical for the genus. The choa-

nosome is constructed by polyspicular tracts and many 

spicules without orientation in between. Oxeas are of 

a wide size range (145–1200 × 5–20 μm).

Zoogeographic distribution. Cosmopolitan and lo-

cally common. In Alaska – central Aleutian Islands, 

Bering Sea (Pribilof Canyon), and Arctic Ocean. Else-

where – North Pacific Ocean, Arctic Ocean, and North 

Atlantic Ocean.

Habitat. In Alaska – attached to boulders and cobbles 

at depths between 97 and 167 m (central Aleutian 

Islands). In the Bering Sea it attaches to cobbles and 

pebbles at depths between 208 and 300 m. Elsewhere – 

reported at depths between 6 and 220 m.

Remarks. Koltun (1959) described a cushion-shaped 

sponge with a surface of numerous elongated papillae 

with oscula on top. The agglutinated small tubes of the 

Aleutian Island specimens are the equivalent of these 

papillae in a fully grown sponge. The very long oxeas 

in halichondroid confused arrangement occur in both 

specimens and are regarded as diagnostic. 

Photos. 1) Specimen collected at a depth of 167 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 (center) in situ growing 

in a cluster of the same species. 3) Fragmented speci-

men collected at a depth of 300 m in Pribilof Canyon, 

Bering Sea. 

101. Halichondria (Eumastia) sitiens (Schmidt, 1870)
122 Professional Paper NMFS 12

Description. This sponge is extremely polymorphic; 

reported growth forms include encrusting, massivelobate, tubular, lobate, and vase-shaped. Oscula may 

be flush with the surface, slightly elevated, or on larger 

cone-shaped elevations. Consistency is only slightly 

elastic but easily fragmented. Size is to at least 30 cm × 

30 cm. Color in life ranges from brown, grey, orange, 

green, yellow, to creamy white.

Skeletal structure. Ectosomal spicules are tangentially arranged. Choanosomal arrangement of spicules 

is mostly confused but in places may be a unispicular 

reticulation or meshes of polyspicular tracts. Spicules 

are oxeas (220–335 × 15–24 μm).

Zoogeographic distribution. Locally abundant. In 

Alaska – Aleutian Islands, Bering Sea (Pribilof Canyon), 

and eastern Gulf of Alaska. Elsewhere – North Pacific 

Ocean.

Habitat. In Alaska – attached to hard substrate. Depths 

between 155 and 208 m (Aleutian Islands); depths between 71 and 255 m (eastern Gulf of Alaska); Bering 

Sea (Pribilof Canyon) – locally abundant; attached to 

bedrock, cobbles, pebbles, and encrusting the gorgonian coral Plumarella echinata at depths between 208 and 

309 m. Elsewhere – no information available.

Remarks. This species might represent an undescribed species. The Alaskan specimens are similar to 

Halichondria panicea (Pallas, 1766) that is common to 

the North Atlantic Ocean and was recently introduced 

to San Francisco Bay. Alaskan specimens have a slightly 

different external appearance and the spicules are 

shorter and more robust than those of H. panicea. There 

are ongoing discussions about whether there are several sibling species or whether the Atlantic and Pacific 

Ocean populations are conspecific (Erpenbeck and Van 

Soest, 2002). Eggs and hatching larvae similar to those 

of the snailfish (Careproctus sp.) were found in flabellate forms of this species in Pribilof Canyon, Bering Sea 

(Busby4

). This species is preyed upon by the sea stars 

Hippasteria spp., Pteraster tesselatus, Ceramaster patagonicus, 

and possibly Henricia longispina.

Photos. 1) Specimen collected at a depth of 175 m 

in the central Aleutian Islands. Grid marks are 1 cm2. 

2) Same specimen as in photo 1 in situ. The specimen 

was detached and lying on the seafloor in an area that 

had been trawled. 3) A fragment of a specimen collected at 152 m in the eastern Gulf of Alaska. Grid marks 

are 1 cm2. 4) Same specimen as in photo 3 (lower right) 

in situ. Crinoids (Florometra serratissima) use the sponge 

102. Halichondria sp. 

as an elevated perch. 5) Specimen collected at a depth 

of 310 m in Pribilof Canyon, Bering Sea. This specimen 

has completely encrusted the gorgonian Plumarella

echinata (tips exposed at right). Grid marks are 1 cm2. 

6) Same specimen as in photo 5 (indicated by the white 

arrow) in situ. 7) Specimen collected at a depth of 155 

m in the central Aleutian Islands. Grid marks are 1 cm2.
123

102. Halichondria sp. (continued)
124 Professional Paper NMFS 12

Description. This sponge is polymorphic; massivelobate, massively cylindrical, ramified, semiglobular, or 

even encrusting. Some Aleutian specimens are highly 

ramified with some branches coalesced together. Rows 

of small oscula are visible in situ. The consistency is 

only slightly elastic, stiff, and easily torn. It appears to 

be particularly fragile. Height of highly ramified form 

is to 1 m or more. Color in life is yellow, light brown, 

golden-brown or creamy-white.

Skeletal structure. Ectosomal spicules are tangen-

tially arranged without further orientation. There is 

choanosomal reticulation of polyspicular tracts and 

in other parts short tracts and single spicules without 

orientation. Styles (135–560 × 6–23 μm) probably occur 

in two size categories.

Zoogeographic distribution. Cosmopolitan and lo-

cally common. In Alaska – central Aleutian Islands, 

Bering Sea, and Chukchi Sea. Elsewhere – North Pacific 

Ocean (Sea of Okhotsk and Sea of Japan), Arctic Ocean 

(Barents Sea, East Siberian Sea, and Kara Sea), and 

North Atlantic Ocean.

Habitat. In Alaska – attached to cobbles at depths 

between 119 and 253 m; also encrusts the gorgonian 

Muriceides nigra. Elsewhere – reported at depths be-

tween 15 and 110 m.

Photos. 1) Partial specimen (ramified form) collected at a depth of 119 m in the central Aleutian Islands. 

Grid marks are 1 cm

2

. 2) Same specimen as in photo 

1 (center) in situ. 3) Close-up view of same specimen 

as in photos 1 and 2 showing rows of small oscula. 

4) Fragment of specimen (encrusting form) collected 

at a depth of 160 m in the central Aleutian Islands. 

Specimen encrusts the gorgonian Muriceides nigra. 

103. Hymeniacidon assimilis Levinsen, 1887
125

Description. On deck this species appears a massive 

sponge, but in situ it forms stout hollow tubes with 

spicule tracts protruding far above the surface. The 

consistency is fragile, inelastic, and easily torn. Height 

is to 7 cm and width to 5 cm. Color in life is light brown 

to creamy white.

Skeletal structure. Skeleton is confused and hali-

chondroid. Large oxeas are 438–1400 × 13–21 μm; 

small oxeas (50–150 × 4–6 μm) are concentrated in the 

ectosome.

Zoogeographic distribution. Uncommon. In Alaska – 

central Aleutian Islands, Bering Sea, and Arctic Ocean 

(Beaufort Sea – Point Barrow). Elsewhere – Vancouver 

Island, British Columbia. 

Habitat. In Alaska – attached to mudstone on steep 

canyon habitat at a depth of 2828 m. Grows in small 

clusters of up to eight individuals. Elsewhere – reported 

at depths between 80 and 110 m.

Remarks. This species was previously known as 

Halichondria disparilis Lambe, 1893. It appears to be 

extremely eurybathic. 

Photos. 1) Preserved (frozen then stored in ethanol) 

specimen collected at a depth of 2828 m in the central 

Aleutian Islands. Grid marks are 1 cm

2

. 2) Same speci-

men as in photo 1 (lower left) in situ with a cluster of 

the same species.

104. Topsentia disparilis (Lambe, 1893)
126 Professional Paper NMFS 12

Description. Two growth forms exist: 1) massive to 

hemispherical, and 2) fan-shaped and stalked. The fan 

may be thin-bladed or form an ovoid tube with a large 

osculum on top. The surface is irregularly corrugated 

with crater-like oscula. Both growth forms narrow somewhat to a basal attachment or short stalk. Consistency 

is soft, elastic, and easily torn. Size is to 10 cm × 10 cm. 

Color in life is reddish brown or golden brown. 

Skeletal structure. There is an ectosomal unispicular 

reticulation of tangentially arranged oxeas (160–180 × 

5–12 μm). Where oxeas are connected, one to four are 

arranged perpendicular to the surface. There is a choanosomal unispicular reticulation; in places paucispicular tracts are connected by single spicules.

Zoogeographic distribution. Widespread but uncommon. In Alaska – central Aleutian Islands. Elsewhere 

– North Pacific Ocean (Sea of Japan), Arctic Ocean 

(Barents Sea and Greenland Sea), and North Atlantic 

Ocean (Davis Strait).

Habitat. In Alaska – attached to cobbles, pebbles, 

and bivalve shells at depths between 155 and 235 m. 

Elsewhere – reported at depths between 40 and 718 m.

Photos. 1) Specimen collected at a depth of 190 m 

in the central Aleutian Islands. Grid marks are 1 cm2

. 

2) Same specimen as in photo 1 (center) in situ. 

3) Specimen attached to a scallop shell (Chlamys sp.) 

collected at a depth of 155 m in the central Aleutian 

Islands. Grid marks are 1 cm2. 4) Specimen attached 

to a pebble collected at a depth of 155 m in the central 

Aleutian Islands. Grid marks are 1 cm2.

105. Cladocroce ventilabrum (Fristedt, 1887)
127

Description. This small vase-shaped sponge has a 

thin fistular stalk or basal fistular stolons. Due to the 

reticulation of single spicules these sponges are only 

slightly elastic and consequently very fragile. Height is 

to 3 cm and width to 1 cm. Color in life is light brown 

to golden brown.

Skeletal structure. There is a unispicular reticulation 

of oxeas (165–195 × 7–9 μm), slightly longer than those 

originally described.

Zoogeographic distribution. Rare. In Alaska – central 

Aleutian Islands. Elsewhere – known only from Sagami 

Bay, Japan.

Habitat. In Alaska – attached to the calcareous sponge 

Leucandra tuba at a depth of 145 m. Elsewhere – depths 

between 80 and 95 m.

106. Haliclona bucina Tanita and Hoshino, 1989

Photo. 1) Specimen (center) attached to the calcareous sponge Leucandra tuba collected at a depth of 145 

m in the central Aleutian Islands. Grid marks are 1 cm2.
128 Professional Paper NMFS 12

Description. Koltun (1959) described this species as 

“elongated, thinly tabular (foliate), or rolled up and intergrown in such a manner that it acquires an irregular 

shape ... hollow dactylate projections....” Aleutian Island specimens are typically stalked and flabellate. The 

surface is smooth. The oscula are circular, chimney-like, 

on top of oblique tubes. The consistency is only slightly 

elastic and easy to tear. Size is to at least 15 cm in height 

and width. Color in life is light brown to golden brown; 

appears characteristically yellow in situ. 

Skeletal structure. The choanosomal skeleton consists 

of long polyspicular tracts connected by single spicules, 

oxeas (320–370 × 12–18 μm), and thin sigmas (18–

25 μm).

Zoogeographic distribution. Uncommon. In Alaska 

– central Aleutian Islands. Elsewhere – North Pacific 

Ocean (Sea of Okhotsk near the Pacific coast of the 

southern Kuril Islands).

Habitat. In Alaska – attached to bedrock, boulders, cobbles, and pebbles at depths between 96 and 

107. Haliclona (Gellius) digitata (Koltun, 1958)

258 m. Elsewhere – found on sand and gravel at depths 

between 285 and 287 m and at a temperature of 1.7°C.

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2. 
129

Description. According to Koltun (1959) this species 

is massive-lobate, or cushion-shaped, up to 10.5 cm in 

height, and often forms dactylate, lobate or other types 

of projections. Aleutian Island specimens are dactylate 

with hollow centers; up to three fingers are basally attached. Surface is corrugated. Color in life is light yellow, brown, or golden brown.

Skeletal structure. The ectosomal skeleton is a tan-

gential, unispicular reticulation of single spicules. 

Conspicuous polyspicular tracts, several mm in length, 

are connected by a reticulation of single spicules in the 

choansosome. Oxeas are 140–175 × 8–13 μm and toxa 

are 45–135 μm.

Zoogeographic distribution. Widespread but uncom-

mon. In Alaska – central Aleutian Islands and Bering 

Sea. Elsewhere – North Pacific Ocean (Sea of Okhotsk, 

Sea of Japan, Pacific coast of the Kuril Islands), Arctic 

Ocean (White Sea), and North Atlantic Ocean (west of 

Greenland).

Habitat. In Alaska – attached to pebbles at a depth 

of 138 m. Elsewhere – found on sand, rock, and ooze 

at depths between 27 and 200 m, at temperatures be-

tween 0.1 and 3.4°C, and at salinities between 27.0 and 

33.9 psu.

Photos. 1) Partial specimen collected at a depth of 138 

m in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Same specimen as in photo 1 (indicated by the white 

arrows) in situ covered with ophiuroids.

108. Haliclona (Gellius) primitiva (Lundbeck, 1902)
130 Professional Paper NMFS 12

Description. This sponge is cushion-shaped with a 

finely hispid surface. Consistency is very delicate; only 

slightly elastic and easily torn. Circular oscula are slightly elevated from the surface and not visible on collected 

specimens. Height is to 15 cm and width to 8 cm. Color 

in life is light grey to creamy white. 

Skeletal structure. Choanosomal paucispicular tracts 

are connected by single spicules and short tracts; oxeas 

(330–430 × 13–15 μm).

Zoogeographic range. Widespread but rare. In Alaska 

– central Aleutian Islands. Elsewhere – North Pacific 

Ocean (Sea of Okhotsk and Sea of Japan), Arctic Ocean 

(Barents Sea and Greenland Sea), and North Atlantic 

Ocean. 

Habitat. In Alaska – attached to bedrock, boulders, 

and mudstone at depths between 974 and 1706 m. 

Elsewhere – reported at depths between 0 and 15 m 

(Barents Sea) and at a depth of 887 m (North Atlantic 

Ocean).

Photos. 1) Fragments of preserved (frozen then ethanol) specimen collected at a depth of 1352 m in the 

central Aleutian Islands. Grid marks are 1 cm2

. 2) Same 

specimen as in photo 1 in situ. 3) Close-up view of same 

specimen as in photo 2 and 3.

109. Haliclona tenuiderma (Lundbeck, 1902)
131

Description. This sponge is stalked and oviform or 

stalked tubes (de Weerdt, 1986a). Aleutian Island specimens are stalked tubes; stalk approximately 1 cm long, 

overall length to about 10 cm. Color in life is light yellow 

with a darker stalk.

Skeletal structure. There is a unispicular reticulation 

of oxeas (178–280 μm).

Zoogeographic range: Uncommon. In Alaska – 

central Aleutian Islands (first record in the North Pacific Ocean). Elsewhere – Arctic Ocean (Kara Sea) and 

North Atlantic Ocean (North Sea).

Habitat. In Alaska – attached to cobbles at depths near 

490 m. Elsewhere – found on sediment covered stones 

in sheltered habitats at depths between 5 and l000 m.

Photos. 1) Preserved (frozen) specimen collected at 

a depth of 490 m in the central Aleutian Islands. Grid 

marks are 1 cm2

. 2) Same specimen as in photo 1 (right) 

in situ.

110. Haliclona (Haliclona) urceolus (Rathke and Vahl, 1806)
132 Professional Paper NMFS 12

Description. This sponge is massively encrusting or 

cushion-shaped. Oscula are deep and crater-shaped. 

Surface is slightly rough; consistency is soft and fragile. 

Diameter is to about 10 cm. Color in life is light brown 

or golden brown.

Skeletal structure. There is a choanosomal arrangement of oxeas (160–220 × 10–15 μm) in paucispicular 

tracts connected by single spicules. 

Zoogeographic distribution. Uncommon. In Alaska – 

central Aleutian Islands and Bering Sea. 

Habitat. Attached to cobbles and pebbles at depths 

between 150 and 160 m. 

Remarks. This species might represent an undescribed species.

111. Haliclona sp. 1 

Photo. 1) Specimen collected at a depth of 155 m in 

the central Aleutian Islands. Grid marks are 1 cm2.
133

Description. This sponge is flabellate; single- or multiplaned. Oscula are conspicuous, slightly elevated and 

scattered over the entire surface. It is extremely slimy, 

even in preservative. Consistency is very soft and elastic. 

Height and width are to 20 cm. Color in life is yellowish brown to light brown, but appears characteristically 

white in situ.

Skeletal structure. Ectosomal tangential unispicu-

lar reticulation is strengthened by large quantities of 

spongin which covers the sponge like a perforated 

plate. Choanosomal paucispicular tracts are connected 

by single spicules to unispicular reticulation of oxeas 

(180–230 × 5–12 μm).

Zoogeographic distribution. In Alaska – locally abun-

dant in central Aleutian Islands. 

Habitat. Attached principally to bedrock, but occa-

sionally to boulders and cobbles at depths between 74 

and 195 m; more common at depths shallower than 

120 m. 

Remarks. This species might represent an unde-

scribed species. It is similar in growth form to Haliclona 

cinerea and displays the slime strands previously regard-

ed as diagnostic for the species. However, H. cinerea has 

smaller oxeas than this species and is known only from 

shallow waters of the Celtic Seas Region of the North 

Atlantic Ocean. This species may be preyed upon by the 

blood star (Henricia sp.).

Photos. 1) Specimen collected at a depth of 80 m 

in the central Aleutian Islands. Grid marks are 1 cm

2

. 

2) Specimen (right) observed at 92 m in the central 

Aleutian Islands.

112. Haliclona sp. 2
134 Professional Paper NMFS 12

Acknowledgments

We thank E. Brown, D. Carlile, S. France, L. Watling, 

S. Rooney, P. Malecha, T. Marshall, K. Lowyck, J. Hocevar, M. Ridgway, A. Andrews, and J. Mondragon for 

assistance with the collection of specimens. We thank 

John Hocevar of Greenpeace International for allowing 

us to include in this guide specimens collected during 

their 2007 Bering Sea Canyons Expedition. T. Warshaw 

provided the topside photographs of the Bering Sea 

specimens; B. Masuda helped with much of the photo 

editing. M. Kelly (NIWA) provided the skeletal structure 

description for the undescribed species of Latrunculia. 

The field collections were supported by the Alaska Fisheries Science Center of the National Marine Fisheries 

Service, West Coast and Polar Regions Undersea Research Center of NOAA’s National Undersea Research 

Program, and the North Pacific Research Board. We 

thank three anonymous reviewers for helpful comments 

on an earlier version of this manuscript. Funding for the 

completion of this guide was provided by the National 

Marine Fisheries Service, Office of Habitat Conservation, as part of the Deep Sea Coral Research and Technology Program. 

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137

PHYLUM PORIFERA

CLASS CALCAREA

Order Baerida

 Family Baeriidae 

 Leuconia alaskensis de Laubenfels, 1953; (Arctic Ocean – Point Barrow, Beaufort Sea)

 Leucopsila cf. stylifera (Borojevic, Boury-Esnault and Vacelet, 2000); (Aleutian Islands)

Order Clathrinida

 Family Clathrinidae 

 Clathrina sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 12

Order Leucosolenida

 Family Amphoriscidae 

 Leucilla nuttingi (Urban, 1902); (Aleutian Islands)

 Family Grantiidae 

 Leucandra ananas (Montagu, 1818); (Arctic Ocean – Point Barrow, Beaufort Sea)

 Leucandra heathi Urban, 1906; (Aleutian Islands)

 Leucandra poculiformis Hozawa, 1918. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 13

 Leucandra pyriformis (Lambe, 1893); (Aleutian Islands)

 Leucandra taylori Lambe, 1900; (Aleutian Islands)

 Leucandra tuba Hozawa, 1918. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 14

 Family Leucosoleniidae

 Leucosolenia eleanor Urban, 1905; (Aleutian Islands, Gulf of Alaska)

 Family Sycettidae 

 Sycon compactum Lambe, 1893; (Aleutian Islands)

CLASS HEXACTINELLIDA

Subclass AMPHIDISCOPHORA

Order Amphidiscosida 

 Family Hyalonematidae 

 Hyalonema (Cyliconema) apertum apertum Schulze, 1886; (western Aleutian Islands)

 Hyalonema (Cyliconema) apertum simplex Koltun, 1967; (Bering Sea)

 Hyalonema (Cyliconema) hozawai vicarium Koltun, 1967; (Bering Sea)*

 Hyalonema (Leptonema) ovuliferum Schulze, 1899; (southern Gulf of Alaska)

 Hyalonema (Cyliconema) tenerum vitiazi Koltun, 1967; (Bering Sea)*

Subclass HEXASTEROPHORA

Order Hexactinosida

 Family Farreidae

 Farrea kurilensis ssp. nov. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 16

 Farrea occa occa Bowerbank, 1862. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 17

 Farrea watasei Okada, 1932; (Bering Sea)*

 Farrea sp. Okada, 1932; (Bering Sea)

 Farrea sp. nov. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 18

Appendix I. Sponge species reported from Alaskan waters

This list includes the sponges currently known to 

occur in the deep waters (>80 m) of Alaska. The list 

has been drawn from the published literature (mostly 

species descriptions) and from recent collections of 

specimens that have been definitively identified by us 

through examination of microscopic characters. We 

provide a complete list of all species reported from 

the area, but only provide detailed species descriptions in this guide for those species that we collected 

and examined ourselves. For all other species we provide the zoogeographic ranges, but refer the reader to 

the primary literature for those species descriptions. 

The taxonomic names are all modern, systematically 

arranged to family, and alphabetically arranged by 

genus and species within family. (*) indicates that the 

species was reported without precise coordinates but 

we believe that the collection was made in Alaskan 

waters.
138 Professional Paper NMFS 12

 Family Euretidae

 Chonelasma sp. Schulze, 1899; (northern Gulf of Alaska)

 Eurete irregularis Bowerbank, 1876; (Bering Sea)*

 Genus nov., sp. nov. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 19

 Family Tretodictyidae

 Tretodictyum sp. nov. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 20

 Family Aphrocallistidae

 Aphrocallistes vastus Schulze, 1886. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 21

 Heterochone aleutiana (Okada, 1932); (western Aleutian Islands)

 Heterochone calyx calyx Schulze, 1886. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 23

 Heterochone calyx schulzei Koltun, 1967; (Bering Sea)

 Heterochone tenera Schulze, 1899; (Bering Sea)* 

 Heterochone n. sp. A Schuchert and Reiswig, 2006; (Gulf of Alaska)

Order Lyssacinosida 

 Family Euplectellidae 

 Euplectella oweni Herklots and Marshall, 1868; (Bering Sea)*

 Euplectella sp. Koltun, 1967; (Bering Sea)*

 Holascus undulatus Schulze, 1899 (southern Gulf of Alaska)

 Regadrella okinoseana Ijima, 1896. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 25

 Family Rossellidae 

 Acanthascus (Acanthascus) alani profundum Koltun, 1967; (Bering Sea)*

 Acanthascus (Rhabdocalyptus) australis Topsent, 1901; (Bering Sea)

 Acanthascus (Rhabdocalyptus) borealis Okada, 1932; (Bering Sea)

 Acanthascus (Rhabdocalyptus) dawsoni alascensis Wilson and Penney, 1930 (Bering Sea)

 Acanthascus (Rhabdocalyptus) dawsoni dawsoni (Lambe, 1893) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 27

 Acanthascus (Rhabdocalyptus) dawsoni horridus Koltun, 1967; (Bering Sea)*

 Acanthascus (Staurocalyptus) dowlingi (Lambe, 1894); (Gulf of Alaska)

 Acanthascus (Rhabdocalyptus) heteraster Okada, 1932; (Bering Sea)

 Acanthascus (Rhabdocalyptus) mirabilis (Schulze, 1899) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 28

 Acanthascus (Acanthascus) mitis Koltun, 1967; (Bering Sea)*

 Acanthascus (Acanthascus) profundum ssp. nov. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . Page 26

 Acanthascus (Staurocalyptus) rugocruciatus Okada, 1932; (Bering Sea)

 Acanthascus (Staurocalyptus) solidus (Schulze, 1899) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 29

 Acanthascus (Rhabdocalyptus) unguiculatus (Ijima, 1904); (western Aleutian Islands)

 Acanthascus (Staurocalyptus) sp. nov. 1 Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . Page 30

 Acanthascus (Staurocalyptus) sp. nov. 2 Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . Page 31

 Aulosaccus fissuratus Okada, 1932; (Bering Sea, western Aleutian Islands)

 Aulosaccus ijimai (Schulze, 1899); (southern Gulf of Alaska)

 Aulosaccus pinularis Okada, 1932. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 32

 Aulosaccus schulzei Ijima, 1896. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 33

 Bathydorus echinus Koltun, 1967; (Bering Sea)*

 Bathydorus laevis spinosus Wilson, 1904; (Bering Sea)*

 Bathydorus sp. Okada, 1932; (Bering Sea)

 Bathydorus sp. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 34

 Caulophacus (Caulophacus) elegans Schulze, 1885; (Bering Sea)*

 Caulophacus (Caulophacus) schulzei hyperboreus Koltun, 1967; (Bering Sea)*

 Caulophacus (Caulophacus) sp. nov. Reiswig and Stone, in preparation. . . . . . . . . . . . . . . . . . . . . . . . . . . Page 35

 Scyphidium tuberculata (Okada, 1932); (Bering Sea)

CLASS DEMOSPONGIAE

Order Dendroceratida

 Family Darwinellidae

 Aplysilla glacialis (Merejkowski, 1877); (Arctic Ocean)
139

Order Homosclerophorida

 Family Plakinidae

 Plakina atka Lehnert, Stone and Heimler, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 38

 Plakina tanaga Lehnert, Stone and Heimler, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 39

Order Spirophorida

 Family Tetillidae

 Craniella arb (de Laubenfels, 1930) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 40

 Craniella craniana de Laubenfels, 1953; (Arctic Ocean – Point Barrow, Beaufort Sea)

 Craniella sigmoancoratum (Koltun, 1966) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 41

 Craniella spinosa Lambe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 42

 Craniella sputnika Lehnert, Stone and Heimler, 2011 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 43

 Craniella villosa Lambe, 1893; (Aleutian Islands, Gulf of Alaska)

Order Astrophorida

 Family Ancorinidae

 Penares cortius de Laubenfels, 1930; (Gulf of Alaska)

 Stelletta validissima Thiele, 1898; (Bering Sea)

 Family Geodiidae

 Erylus aleuticus Lehnert, Stone and Heimler, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 44

 Geodia lendenfeldi nomen novum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 45

 Geodia mesotriaena Lendenfeld, 1910; (southern Gulf of Alaska)

 Family Pachastrellidae

 Poecillastra japonica (Thiele, 1898); (Bering Sea)

 Poecillastra tenuilaminaris (Sollas, 1886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 46

Order Hadromerida

 Family Polymastiidae

 Polymastia andrica de Laubenfels, 1949; (Arctic Ocean – Point Barrow, Beaufort Sea)

 Polymastia fluegeli Lehnert, Stone and Heimler, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 47

 Polymastia kurilensis Koltun, 1962; (Gulf of Alaska, Bering Sea)

 Polymastia pacifica Lambe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 48

 Radiella sol Schmidt, 1870; (eastern Gulf of Alaska)

 Family Stylocordylidae

 Stylocordyla borealis eous Koltun, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 49

 Family Suberitidae

 Aaptos kanuux Lehnert, Hocevar and Stone 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 50

 Pseudosuberites montiniger (Carter, 1880); (eastern Gulf of Alaska – Cross Sound)

 Rhizaxinella clavata (Thiele, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 51

 Suberites concinnus Lambe, 1895; (Gulf of Alaska, Bering Sea, Arctic Ocean)

 Suberites excellens (Thiele, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 52

 Suberites montalbidus Carter, 1880; (eastern Aleutian Islands)

 Suberites simplex Lambe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 53

 Suberites suberia (Montagu, 1818); (Gulf of Alaska, Bering Sea)

 Suberites virgultosus (Johnston, 1842); (Arctic Ocean – Point Barrow, Beaufort Sea)

 Suberites sp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 54

Order Haplosclerida (Suborder Petrosina)

 Family Petrosiidae

 Petrosia (Petrosia) borealis (Lambe, 1895); (western Aleutian Islands)

Order Haplosclerida (Suborder Haplosclerina)

 Family Niphatidae

 Hemigellius porosus (Fristedt, 1887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 55

Order Poecilosclerida (Suborder Microcionina)

 Family Acarnidae

 Cornulum clathriata (Koltun, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 56

 Iophon piceum (Vosmaer, 1882) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 57
140 Professional Paper NMFS 12

 Family Acarnidae (continued)

 Iophon piceum abipocillus Koltun, 1959. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 58

 Megaciella anisochela Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 59

 Megaciella spirinae (Koltun, 1958) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 60

 Wigginsia wigginsi de Laubenfels, 1953; (Arctic Ocean – Point Barrow, Beaufort Sea)

 Family Microcionidae

 Clathria (Clathria) barleei (Bowerbank, 1866) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 61

 Clathria (Clathria) laevigata Lambe, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 62

 Clathria (Axosuberites) lambei (Koltun, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 63

 Echinoclathria beringensis (Hentschel, 1929); (Arctic Ocean – Point Barrow, Beaufort Sea)

 Echinoclathria vasa Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 64

 Artemisina amlia Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 65

 Artemisina arcigera (Schmidt, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 66

 Artemisina stipitata Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 67

 Artemisina sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 68

Order Poecilosclerida (Suborder Myxillina)

 Family Coelosphaeridae

 Coelosphaera oglalai Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 69

 Inflatella globosa (Koltun, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 70

 Lissodendoryx (Lissodendoryx) amaknakensis (Lambe, 1895); (Gulf of Alaska, Bering Sea)

 Lissodendoryx (Lissodendoryx) behringi Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 71

 Lissodendoryx (Lissodendoryx) firma (Lambe, 1895); (western Aleutian Islands)

 Lissodendoryx (Ectyodoryx) olgae (Hentschel, 1929). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 72

 Lissodendoryx (Lissodendoryx) oxeota Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 73

 Lissodendoryx (Lissodendoryx) papillosa Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 74

 Family Crambeidae

 Monanchora alaskensis (Lambe, 1895) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 75

 Monanchora laminachela Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 76

 Monanchora pulchra (Lambe, 1894) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 77

 Family Crellidae

 Crella brunnea (Hansen, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 78 

 Family Hymedesmiidae

 Hymedesmia (Stylopus) dermata Lundbeck, 1910. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 79

 Hymedesmia (Hymedesmia) irregularis Lundbeck, 1910. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 80

 Hymedesmia (Stylopus) longurius Lundbeck, 1910; (Aleutian Islands)

 Kirkpatrickia borealis Koltun, 1970. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 81

 Phorbas paucistylifer Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 82

 Family Iotrochotidae

 Iotroata magna (Lambe, 1900); (western Aleutian Islands)

 Family Myxillidae

 Melonanchora globogilva Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 83

 Myxilla (Myxilla) barentsi Vosmaer, 1885; (Gulf of Alaska, Bering Sea, Arctic Ocean)

 Myxilla (Myxilla) behringensis Lambe, 1895. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 84

 Myxilla (Myxilla) incrustans (Johnston, 1842); (Arctic Ocean – Point Barrow, Beaufort Sea; 

 Gulf of Alaska)

 Myxilla (Burtonanchora) lacunosa Lambe, 1893; (Aleutian Islands, Gulf of Alaska)

 Myxilla (Ectyomyxilla) parasitica Lambe, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 85

 Myxilla (Burtonanchora) pedunculata Lundbeck, 1905. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 86

 Stelodoryx oxeata Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 87

 Stelodoryx toporoki Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 88

 Stelodoryx vitiazi (Koltun, 1959) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 89

 Family Phellodermidae

 Echinostylinos hirsutus Koltun, 1970. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 90

 Family Tedaniidae

 Tedania (Tedania) dirhaphis Hentschel, 1912. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 91
141

 Family Tedaniidae (continued)

 Tedania (Tedania) fragilis Baer, 1906; (eastern Aleutian Islands) 

 Tedania kagalaskai Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 92

Order Poecilosclerida (Suborder Mycalina)

 Family Cladorhizidae

 Abyssocladia bruuni Lévi, 1964; (southern Gulf of Alaska) 

 Asbestopluma occidentalis (Lambe, 1893); (Bering Sea)

 Asbestopluma ramosa Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 93

 Cladorhiza bathycrinoides Koltun, 1955. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 95

 Cladorhiza corona Lehnert, Watling and Stone, 2005. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 96

 Cladorhiza longipinna Ridley and Dendy, 1886 (southern Gulf of Alaska)

 Chondrocladia (Chondrocladia) concrescens (Schmidt, 1880) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 97

 Family Desmacellidae

 Biemna rhadia de Laubenfels, 1930; (southern Gulf of Alaska)

 Biemna variantia (Bowerbank, 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 98

 Family Guitarridae

 Euchelipluma elongata Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 99

 Guitarra abbotti Lee, 1987. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 100

 Guitarra fimbriata Carter, 1874. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 101

 Family Esperiopsidae

 Amphilectus digitatus (Miklucho-Maclay, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 102

 Amphilectus lobatus (Montagu, 1818); (Alaska)

 Esperiopsis flagrum Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 103

 Semisuberites cribrosa (Miklucho-Maclay, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 104

 Family Mycalidae

 Mycale (Aegogropila) adhaerens (Lambe, 1893) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 106

 Mycale (Carmia) carlilei Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 107

 Mycale (Carmia) helios (Fristedt, 1887); (Bering Sea, Arctic Ocean)

 Mycale (Mycale) hispida (Lambe, 1893); (Gulf of Alaska, Aleutian Islands)

 Mycale (Mycale) jasoniae Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 108

 Mycale (Mycale) lingua (Bowerbank, 1866); (central Aleutian Islands)

 Mycale (Mycale) loveni (Fristedt, 1887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 109

 Mycale (Mycale) modesta (Lambe, 1894); (Gulf of Alaska, Bering Sea) 

 Mycale (Mycale) toporoki Koltun, 1958; (Bering Sea)

 Mycale (Mycale) tylota Koltun, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 112

 Family Isodictyidae

 Isodictya quatsinoensis (Lambe, 1892); (Gulf of Alaska, Bering Sea)

Order Poecilosclerida (Suborder Latrunculia)

 Family Latrunculiidae

 Latrunculia occulta Lehnert, Stone and Heimler, 2006; (central Aleutian Islands)

 Latrunculia (Biannulata) oparinae Samaii and Krasokhin, 2002. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 113

 Latrunculia velera Lehnert, Stone and Heimler, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 114

 Latrunculia (undescribed species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 115

Order Halichondrida

 Family Axinellidae

 Axinella blanca Koltun, 1959. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 116

 Axinella rugosa (Bowerbank, 1866) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 117

 Family Bubaridae

 Bubaris vermiculata (Bowerbank, 1866). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 118

 Family Halichondriidae

 Halichondria (Halichondria) colossea Lundbeck, 1902. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 119

 Halichondria (Halichondria) lambei Brøndsted, 1933; (Arctic Ocean – Point Barrow, Beaufort Sea)

 Halichondria (Halichondria) oblonga (Hansen, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 120

 Halichondria (Eumastia) sitiens (Schmidt, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 121

 Halichondria sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 122
142 Professional Paper NMFS 12

 Family Halichondriidae (continued)

 Hymeniacidon assimilis Levinsen, 1887. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 124

 Topsentia disparilis (Lambe, 1893) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 125

Order Haplosclerida

 Family Chalinidae

 Cladocroce ventilabrum (Fristedt, 1887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 126

 Haliclona bucina Tanita and Hoshino, 1989. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 127

 Haliclona (Gellius) digitata (Koltun, 1958) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 128

 Haliclona (Gellius) primitiva (Lundbeck, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 129

 Haliclona (Rhizoniera) rufescens (Lambe, 1892); (Gulf of Alaska, Bering Sea, Arctic Ocean) 

 Haliclona tenuiderma (Lundbeck, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 130

 Haliclona (Haliclona) urceolus (Rathke and Vahl, 1806) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 131

 Haliclona sp. 1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 132

 Haliclona sp. 2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 133

 Family Petrosiidae

 Xestospongia hispida (Ridley and Dendy, 1886); (Gulf of Alaska)
143

Importance as Vulnerability to

Species fish habitat disturbance Score

CLASS CALCAREA

Clathrina sp. 1 2 1.5

Leucandra poculiformis 1 1 1.0

Leucandra tuba 2 2 2.0

CLASS HEXACTINELLIDA

Farrea kurilensis ssp. nov. 3 2 2.5

Farrea occa occa 3 3 3.0

Farrea sp. nov. 1 1 1.0

Genus nov., sp. nov. 2 1 1.5

Tretodictyum sp. nov. 1 2 1.5

Aphrocallistes vastus 3 3 3.0

Heterochone calyx calyx 3 3 3.0

Regadrella okinoseana 1 1 1.0

Acanthascus (Acanthascus) profundum ssp. nov. 1 1 1.0

Acanthascus (Rhabdocalyptus) dawsoni dawsoni 3 2 2.5

Acanthascus (Rhabdocalyptus) mirabilis 1 1 1.0

Acanthascus (Staurocalyptus) solidus 2 2 2.0

Acanthascus (Staurocalyptus) sp. nov. 1 2 3 2.0

Acanthascus (Staurocalyptus) sp. nov. 2 2 2 2.0

Aulosaccus pinularis 1 1 1.0

Aulosaccus schulzei 1 2 1.5

Bathydorus sp. 1 1 1.0

Caulophacus (Caulophacus) sp. nov. 2 1 1.5

CLASS DEMOSPONGIAE

Plakina atka 1 1 1.0

Plakina tanaga 1 1 1.0

Craniella arb 1 2 1.5

Craniella sigmoancoratum 1 1 1.0

Craniella spinosa 1 1 1.0

Craniella sputnika 1 1 1.0

Erylus aleuticus 1 1 1.0

Geodia lendenfeldi 1 1 1.0

Poecillastra tenuilaminaris 3 3 3.0

Polymastia fluegeli 1 1 1.0

Polymastia pacifica 1 1 1.0

Stylocordyla borealis eous 1 2 1.5

Aaptos kanuux 1 2 1.5

Rhizaxinella clavata 1 2 1.5

Suberites excellens 2 1 1.5

Suberites simplex 1 1 1.0

This list includes the deep-water (>80 m) sponges presented in this guide. Each species is ranked from low 

(1) to high (3) for its importance as fish habitat and 

its vulnerability to disturbance from fishing activities. 

Ranks for these two measures are averaged to provide 

Appendix II. Priorities for bycatch monitoring

a score. Species with scores greater than 2.0 should 

rank as a high priority for monitoring as bycatch in 

commercial fisheries and stock assessment surveys. 

The scores for high priority species are presented in 

bold lettering.
144 Professional Paper NMFS 12

Importance as Vulnerability to

Species fish habitat disturbance Score

CLASS DEMOSPONGIAE (continued)

Suberites sp. 1 1 1.0

Hemigellius porosus 1 2 1.5

Cornulum clathriata 1 1 1.0

Iophon piceum 1 2 1.5

Iophon piceum abipocillus 1 2 1.5

Megaciella anisochela 1 1 1.0

Megaciella spirinae 1 2 1.5

Clathria (Clathria) barleei 1 2 1.5

Clathria (Clathria) laevigata 1 2 1.5

Clathria (Axosuberites) lambei 1 2 1.5

Echinoclathria vasa 1 1 1.0

Artemisina amlia 1 2 1.5

Artemisina arcigera 1 2 1.5

Artemisina stipitata 2 3 2.5

Artemisina sp. 2 3 2.5

Coelosphaera oglalai 1 2 1.5

Inflatella globosa 1 2 1.5

Lissodendoryx (Lissodendoryx) behringi 1 2 1.5

Lissodendoryx (Ectyodoryx) olgae 1 1 1.0

Lissodendoryx (Lissodendoryx) oxeota 1 1 1.0

Lissodendoryx (Lissodendoryx) papillosa 1 1 1.0

Monanchora alaskensis 2 2 2.0

Monanchora laminachela 1 2 1.5

Monanchora pulchra 3 2 2.5

Crella brunnea 1 1 1.0

Hymedesmia (Stylopus) dermata 1 1 1.0

Hymedesmia (Hymedesmia) irregularis 1 1 1.0

Kirkpatrickia borealis 1 2 1.5

Phorbas paucistylifer 1 2 1.5

Melonanchora globogilva 1 1 1.0

Myxilla (Myxilla) behringensis 1 1 1.0

Myxilla (Ectyomyxilla) parasitica 1 1 1.0

Myxilla (Burtonanchora) pedunculata 1 2 1.5

Stelodoryx oxeata 1 2 1.5

Stelodoryx toporoki 2 1 1.5

Stelodoryx vitiazi 1 1 1.0

Echinostylinos hirsutus 1 1 1.0

Tedania (Tedania) dirhaphis 1 2 1.5

Tedania kagalaskai 2 2 2.0

Asbestopluma ramosa 2 2 2.0

Cladorhiza bathycrinoides 1 1 1.0

Cladorhiza corona 1 2 1.5

Chondrocladia (Chondrocladia) concrescens 1 2 1.5

Biemna variantia 1 2 1.5

Euchelipluma elongata 1 1 1.0

Guitarra abbotti 2 1 1.5

Guitarra fimbriata 2 1 1.5

Amphilectus digitatus 2 2 2.0

Esperiopsis flagrum 2 1 1.5

Semisuberites cribrosa 2 2 2.0
145

Importance as Vulnerability to

Species fish habitat disturbance Score

CLASS DEMOSPONGIAE (continued)

Mycale (Aegogropila) adhaerens 2 3 2.5

Mycale (Carmia) carlilei 2 2 2.0

Mycale (Mycale) jasoniae 2 2 2.0

Mycale (Mycale) loveni 3 3 3.0

Mycale (Mycale) tylota 1 2 1.5

Latrunculia (Biannulata) oparinae 2 2 2.0

Latrunculia velera 2 1 1.5

Latrunculia sp. (undescribed) 1 1 1.0

Axinella blanca 3 3 3.0

Axinella rugosa 2 2 2.0

Bubaris vermiculata 1 1 1.0

Halichondria (Halichondria) colossea 2 1 1.5

Halichondria (Halichondria) oblonga 1 2 1.5

Halichondria (Eumastia) sitiens 2 2 2.0

Halichondria sp. 2 3 2.5

Hymeniacidon assimilis 2 2 2.0

Topsentia disparilis 1 1 1.0

Cladocroce ventilabrum 1 2 1.5

Haliclona bucina 1 1 1.0

Haliclona (Gellius) digitata 1 2 1.5

Haliclona (Gellius) primitiva 1 2 1.5

Haliclona tenuiderma 1 1 1.0

Haliclona (Haliclona) urceolus 1 2 1.5

Haliclona sp. 1 1 2 1.5

Haliclona sp. 2 2 1 1.5
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Appendix III. Glossary of terms

Acanthose: spined

Anchorate: anchored or anchor-like

Apical: located at the apex; distal 

Arborescent: highly branched

Areolate: surface covered with numerous circular areas 

Choanosome: interior tissue

Clavate: club-shaped

Conspecificity: belonging to the same species

Conules: cone-shaped projections

Conulose: surface with numerous conules raised by the 

underlying skeleton

Crenulate: surface with minutely notched or scalloped 

projections

Dactylate: finger-like projections

Digitate: deeply divided finger-like projections

Ecomorph: growth form adapted for different ecological regime

Ectosome: exterior surface

Emergent epifauna: fauna having most of its parts elevated above the level of the surrounding seafloor

Endopsammic: main part of the body buried in sand

Epizoic: living attached to the body of an animal; a 

non-parasitic animal that lives attached to the outer 

surface of another animal

Eurybathic: broadly distributed with depth

Filiform: shaped like a thread

Fistules: cone-shaped elevations or tube-like protuberances

Flabellate: fan-shaped

Flagelliform: shaped as a single, very long, erect branch

Foliate: shaped like a leaf

Globular: ball-shaped or spherical

Gonochoristic: having male and female individuals in 

the same population

Hispid: covered with bristly hairs

Holotype: the single specimen used as the basis for the 

first published description of a species and designated 

as the type specimen

Labyrinthic: an intricate structure of interconnecting 

passages

Lobate: having lobes or rounded projections

Mammillate: shaped like a nipple

Massive: large and compact but amorphous or without 

a definable shape (note that the term “lumpy” is used 

in some of the sponge literature)

Megascleres: large structural spicules

Microscleres: small to minute reinforcing or packing 

spicules

Osculum (a): the opening through which water is expelled from the sponge

Ostium (a): the opening through which water enters 

the sponge

Oviparous: producing eggs that are laid and hatch 

externally

Papillae: Nipple-like protuberances projecting from the 

sponge surface that bear either ostia, oscula, or both

Pellicular: covered with a liquid film or organic membrane

Polymorphic: occurring in different shapes 

Polyspicular: composed of multiple spicule types

Saccate: shaped like a sac or pouch

Sibling species: a species that closely resembles another 

in appearance and other characteristics but cannot 

interbreed with it

Spicules: small needle-like structures made of silica or 

calcium carbonate that support the soft tissues of 

sponges

Spongin: the fibrous framework of collagen that forms 

the sponge’s organic skeleton

Spongocoel (= atrium): the central exhalant cavity of 

the sponge

Stolon: a creeping or prostrate rope-like structure

Subglobular: deviating slightly from globular

Subspecies: a morphologically and genetically distinct 

population within a species that inhabits a separate 

geographic area or depth range

Tabular: having a plane surface or flat

Tuberculate: warty or verrucose

Uncinate: large straight diactin spicules covered with 

barbs and brackets all inclined at one end

Viviparous: producing live offspring from within the 

body of the parent
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Appendix IV. Spicule types

This appendix includes scanning electron microscopy (SEM) images of spicules from select calcareous sponges and demosponges collected during the 

2004 Aleutian Island expeditions. It also includes 

several SEM images of spicules from a few hexactinellid sponge species that are common in Alaskan 

waters. The purpose of the appendix is to provide 

the reader with a representative collection of spicule 

images so that they may gain an understanding of the 

terms used in the guide and an appreciation for the 

variation between species and among spicule types. 

This collection also serves as a source of reference 

material for those who wish to microscopically examine sponges. We refer the reader to the Thesaurus 

of Sponge Morphology (Boury-Esnault and Rützler, 

1997) for a comprehensive glossary of terminology 

and a pictorial guide to spicule types and skeletal 

structures. The species in this appendix have been 

arranged systematically to family and alphabetically 

arranged within family. 

Class Calcarea, Family Grantiidae, Leucandra poculiformis Hozawa, 1918. 

(A) Overview of occurring spicules: large tri- and tetractines and small oxeas; (B) large triactine; (C) a rare large 

tetractine; and (D) acanthose microxeas with characteristic bend called a “lance-head” by Hozawa.
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Leucandra poculiformis Hozawa, 1918 (continued)

Class Calcarea, Family Grantiidae, Leucandra tuba Hozawa, 1918. 

(A) Pugiole with unusual biforked end; (B) pugioles, small and large oxeas; (C) typical pugiole with normal points; 

(D) giant tetractines among small category of triactines; and (E) triactines among minute oxeote spicule types.
149

Leucandra poculiformis Hozawa, 1918 (continued)
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Class Hexactinellida, Family Farreidae, Farrea occa occa Bowerbank, 1862. 

(A) An anchorate clavule; (B) a pileate clavule; (C) a dermal pentactin; (D) dictyonal framework; and (E) an oxyhexaster.
151

Farrea occa occa Bowerbank, 1862 (continued)

Class Hexactinellida, Family Aphrocallistidae, Aphrocallistes vastus Schulze, 1886.

(A) Dictyonal framework (scale bar equals 500 μm); (B) a pentactin from a juvenile specimen; (C) a scopule from a juvenile specimen; (D) a scopule head; (E) a spiny oxyhexaster; and (F) a stout oxyhexaster.
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Aphrocallistes vastus Schulze, 1886 (continued)
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Class Demospongiae, Family Plakinidae, Plakina atka Lehnert, Stone and Heimler, 2005.

(A) Spined calthrops with reduced 4th ray, all rays with basal spines; (B) microfurcate end of ray from a tetraloph, 

tetrafurcate lophocalthrops; (C) tetralophose calthrops, all rays tetrafurcate, points of rays microfurcate; (D) smooth 

diods, smooth triods, and thicker triods with strongly spined bases of rays; and (E) thick category of diods with central 

spines. Probably derived from spined triods through reduction of one ray. (Figures A–E reproduced with permission 

from Zootaxa, Magnolia Press.)
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Plakina atka Lehnert, Stone and Heimler, 2005 (continued)
155

Class Demospongiae, Family Plakinidae, Plakina tanaga Lehnert, Stone and Heimler, 2005.

(A) Smooth oxeas, triods, and trilophose calthrops; (B) weakly spined triods and trilophose calthrops; and (C) 

trilophose calthrops with tetrafurcate lophose rays and biforked non-lophose ray. (Figures A–C reproduced with 

permission from Zootaxa, Magnolia Press.)
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Class Demospongiae, Family Geodiidae, Erylus aleuticus Lehnert, Stone and Heimler, 2006.

(A) An anisotylote (one of several unusual spicule types occurring in this species); (B) a juvenile apidaster with different surface structure; (C) adult apidasters; (D) a centrolyte microstrongyle on the surface of an apidaster; (E) 

cladome of triaene with bent rays; (F) star-shaped surface structure on adult aspidaster; (G) oxyaster; and (H) surface structures on aspidasters may vary in geometry. (Figures B, C, D, E, and F reproduced with permission from the 

Journal of the Marine Biological Association of the United Kingdom.)
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Erylus aleuticus Lehnert, Stone and Heimler, 2006 (continued)
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Class Demospongiae, Family Polymastiidae, Polymastia fluegeli Lehnert, Stone and Heimler, 2005. 

(A) Small category of tylostyle and tyle of large tylostyle (note the radiolarian skeleton at lower right); (B) large and 

small tylostyles; and (C) small category of tylostyles on large tylostyles. (Figures A and B reproduced with permission 

from Facies, Springer Verlag.)
159

Class Demospongiae, Family Suberitidae, Aaptos kanuux Lehnert, Hocevar and Stone, 2008. 

(A) Overview of spicules; large strongyloxeas, medium-size subtylostyles, and short ectosomal tylostyles; (B) mediumsized subtylostyles and ectosomal tylostyles; (C) ectosomal tylostyles and ends of medium-sized spicules. (Figures A–C 

reproduced with permission from Zootaxa, Magnolia Press.) 
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Class Demospongiae, Family Acarnidae, Megaciella anisochela Lehnert, Stone and Heimler, 2006. 

(A) Category of thick styles among smaller tylotes and microscleres; (B) small category of palmate isochela; (C) large 

category of palmate isochela; (D) acanthose end of tylote (close-up view); (E) a distorted, tiny anisochela (this is the 

only species of Megaciella with anisochelae); and (F) an anisochela. (Figures A–D reproduced with permission from 

Zootaxa, Magnolia Press.) 
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Megaciella anisochela Lehnert, Stone and Heimler, 2006 (continued)
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Class Demospongiae, Family Microcionidae, Echinoclathria vasa Lehnert, Stone and Heimler, 2006. 

(A) A palmate isochela with narrow alae (note the rounded ends of alae); (B) long ectosomal thin styles among thick 

styles; (C) a palmate isochela, both alae with two points (acanthose end of thin style at upper right); (D) thick styles; 

(E) reduced palmate isochela (note different points on the alae); and (F) acanthose end of thin style. (Figures A–F 

reproduced with permission from Zootaxa, Magnolia Press.)
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Echinoclathria vasa Lehnert, Stone and Heimler (continued)

Class Demospongiae, Family Microcionidae, Artemisina amlia Lehnert, Stone and Heimler, 2006. 

(A) A smooth toxon; (B) a large, smooth style; (C) a thin style with prominent tooth (upper left to lower right); (D) 

an isochela; and (E) a close-up view of prominent tooth on thin category of style. (Figures A–E reproduced with 

permission from Zootaxa, Magnolia Press.)
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Artemisina amlia Lehnert, Stone and Heimler, 2006 (continued)
165

Class Demospongiae, Family Coelosphaeridae, Coelosphaera oglalai Lehnert, Stone and Heimler, 2006. 

(A) A choanosomal strongyle among microscleres; (B) microacanthose end of a choanosomal strongyle (close-up 

view); (C) acanthose end of tylote (close-up view); (D) isochelae (right) and acanthose end of a strongyle (left); and 

(E) an ectosomal tylote (center). (Figures A–E reproduced with permission from Zootaxa, Magnolia Press.)
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Coelosphaera oglalai Lehnert, Stone and Heimler, 2006 (continued)

Class Demospongiae, Family Crambeidae, Monanchora laminachela Lehnert, Stone and Heimler, 2006. 

(A) Choanosomal thick style among thin styles; (B) an ectosomal thin style; (C) an anchorate isochela with characteristic central plate; (D) variation of isochela with fused alae; (E) an isochela (rear view); and (F) a sigma. (Figures 

A–D and F reproduced with permission from Zootaxa, Magnolia Press.) 
167

Monanchora laminachela Lehnert, Stone and Heimler, 2006 (continued)
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Class Demospongiae, Family Myxillidae, Melonanchora globogilva Lehnert, Stone and Heimler, 2006. 

(A) An ectosomal tylote among microscleres; (B) smooth end of tylote and pointed end of acanthostyle (close-up 

view); (C) acanthostyles; (D) a large isochelae (shaft of inner margins of alae with fimbriae); (E) fimbriate inner 

margins of alae of isochelae (close-up view); (F) second type of large isochela with different shape and dented outer 

margins; (G) small category of isochelae; and (H) dented margins of second type of isochela (close-up view). (Figures 

A–H reproduced with permission from Zootaxa, Magnolia Press.)
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Melonanchora globogilva Lehnert, Stone and Heimler, 2006 (continued)
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Class Demospongiae, Family Myxillidae, Stelodoryx oxeata Lehnert, Stone and Heimler, 2006. 

(A) A large oxea with acanthose ends among tornotes with acanthose ends; (B) acanthose end of tornote and a sigma; 

(C) acanthose end of oxea (close-up view); (D) large category of anchorate isochela; (E) medium-sized category 

of anchorate, polydentate isochela; (F) small category of anchorate isochelae and centrolyte sigmas; (G) acanthose 

end of ectosomal tornote, centrotylote sigma (above) and medium-sized isochela (below); and (H) large category of 

isochela (side view). (Figures A–F reproduced with permission from Zootaxa, Magnolia Press.)
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Stelodoryx oxeata Lehnert, Stone and Heimler (continued)
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Class Demospongiae, Family Tedaniidae, Tedania kagalaskai Lehnert, Stone and Heimler, 2006. 

(A) Thick choanosomal styles and ectosomal tylotes; (B) acanthose end of tylote (close-up view); (C) large category 

of onychaete (lower left to upper right) among tylotes; and (D) small category of onychaete (note the unequal ends). 

(Figures A–D reproduced with permission from Zootaxa, Magnolia Press.)
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Class Demospongiae, Family Cladorhizidae, Cladorhiza bathycrinoides Koltun, 1955. 

(A) An anisochela (rear view) with sigma (below); (B) a sigma; (C) a sigmancistra; (D) an anisochela; (E) an anisochela (rear view); (F) small end of anisochela with sharp claws (possibly adapted to capture prey) and pointed end 

of sigmancistra (above); (G) tylostrongyles and anisochelae; and (H) tylostrongyle and anisochelae.
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Cladorhiza bathycrinoides Koltun, 1955 (continued)
175

Class Demospongiae, Family Cladorhizidae, Cladorhiza corona Lehnert, Watling and Stone, 2005. 

(A) Claw-like appendages at the small end of anisochelae; (B) a sigmancistra; (C) a tylostyle among microscleres; 

(D) a large fusiform style and short oxeas; (E) an anisochela with claw-like appendages; (F) a small (anis-)oxea (a 

spicule type present in the basal plate); and (G) large fusiform styles and small (anis-)oxea. (Figure A reproduced 

with permission from the Journal of the Marine Biological Association of the United Kingdom.)
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Cladorhiza corona Lehnert, Watling and Stone, 2005 (continued)
177

Class Demospongiae, Family Guitarridae, Euchelipluma elongata Lehnert, Stone and Heimler, 2006. 

(A) Fusiform choanosomal style; (B) an isochela and sigmas; (C) ectosomal tylostyle, slightly polytylote among microscleres; (D) characteristically thin placochela; (E) developmental stage of polytylote tylostyle and two placochelae; 

(F) a sigma; (G) an isochela, placochela and sigmas; and (H) long fusiform styles, shorter ectosomal tylostyles and 

microscleres. (Figures A–F reproduced with permission from Zootaxa, Magnolia Press.)
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Euchelipluma elongata Lehnert, Stone and Heimler, 2006 (continued)
179

Class Demospongiae, Family Guitarridae, Guitarra abbotti Lee, 1987. 

(A) Fusiform styles; (B) a large placochela among styles; (C) biplacochelae; (D) fimbriae of biplacochela (closeup view); (E) tiny, spined isochela; (F) a biplacochela showing frontal process; and (G) fimbriae of a biplacochela 

(close-up view).
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Guitarra abbotti Lee, 1987 (continued)
181

Class Demospongiae, Family Esperiopsidae, Esperiopsis flagrum Lehnert, Stone and Heimler, 2006. 

(A) Microscleres (large and small categories of isochelae and sigmas); (B) a small isochela; (C) long fusiform styles 

and microscleres; (D) large and small sigmas; (E) a small isochela (side view); (F) a large isochela (rear view); and 

(G) large isochelae (front and rear views). (Figures A–F reproduced with permission from Zootaxa, Magnolia Press.)
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Esperiopsis flagrum Lehnert, Stone and Heimler, 2006 (continued)
183

Class Demospongiae, Family Mycalidae, Mycale (Carmia) carlilei Lehnert, Stone and Heimler, 2006. 

(A) Spicule overview (styles, anisochelae and sigmas); (B) an anisoschela (top view); (C) an anisochela (side view); 

and (D) a sigma. (Figures A–D reproduced with permission from Zootaxa, Magnolia Press.)
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Class Demospongiae, Family Mycalidae, Mycale (Mycale) jasoniae Lehnert, Stone and Heimler, 2006. 

(A) Spicule overview (subtylostyles, large and small anisochelae, and rhaphids); (B) large anisochelae (front, side 

and rear views); (C) side view of a large anisochela (note top view of broken end of small anisochela); (D) small 

category of anisochela and rhaphids; and (E) a small anisochela. (Figures A–D reproduced with permission from 

Zootaxa, Magnolia Press.)
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Class Demospongiae, Family Latrunculiidae, Latrunculia velera Lehnert, Stone and Heimler, 2006. 

(A) Spicule overview (styles and anisodiscorhabds); (B) acanthose end of style; (C) a rare polytylote tylostyle; (D) an 

anisodiscorhabd; (E) an anisodiscorhabd (apex view); (F) apex of anisodiscorhabd (side view); (G) early developmental stage of anisodiscorhabd on pointed end of style (acanthose pointed ends, as shown here, are rare); (H) developmental stage of anisdiscorhabd between two fully developed ones; (I) smoother variation of an anisodiscorhabd; 

and (J) anisodiscorhabds. (Figures A–I reproduced with permission from Zootaxa, Magnolia Press.)

Mycale (Mycale) jasoniae Lehnert, Stone and Heimler, 2006 (continued)
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Latrunculia velera Lehnert, Stone and Heimler, 2006 (continued)
187

Latrunculia velera Lehnert, Stone and Heimler, 2006 (continued)