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Abstract and Figures

The calcareous sponges collected during Indonesian-Dutch research projects, incorporated in the collections of the Naturalis Biodiversity Center (formerly the Rijksmuseum van Natuurlijke Historie and the Zoölogisch Museum of the University of Amsterdam), are described and discussed. A total of 37 species were distinguished, of which 16 are new to science, while several others are very poorly known. The new species are Clathrina purpurea sp.nov., Clathrina beckingae sp.nov., Clathrina sororcula sp.nov., Arthuria tubuloreticulosa sp.nov., Ernstia indonesiae sp.nov., Ernstia chrysops sp.nov., Ernstia klautauae sp.nov., Ernstia naturalis sp.nov., Ascandra kakaban sp.nov., Ascandra crewsi sp.nov., Ascaltis angusta sp.nov., Pericharax orientalis sp.nov., Sycetta vinitincta sp.nov., Anamixilla singaporensis sp.nov., Grantessa borojevici sp.nov. and Grantessa tenhoveni sp.nov. An additional six species reported from Indonesia, but not represented in our material, are briefly characterized.
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Accepted by M. Klautau: 9 Mar. 2015; published: 30 Apr. 2015
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ZOOTAXA
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Copyright © 2015 Magnolia Press
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Monograph
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ZOOTAXA
Calcareous sponges of Indonesia
ROB W.M. VAN SOEST* & NICOLE J. DE VOOGD
Naturalis Biodiversity Center, P.O. Box 9617, 2300 RA Leiden, The Netherlands,
E-mail: rob.vansoest@naturalis.nl; nicole.devoogd@naturalis.nl
*Corresponding author
Magnolia Press
Auckland, New Zealand
3951
VAN SOEST & DE VOOGD
2
·
Zootaxa 3951 (1) © 2015 Magnolia Press
ROB W.M. VAN SOEST & NICOLE J. DE VOOGD
Calcareous sponges of Indonesia
(Zootaxa 3951)
105 pp.; 30 cm.
30 Apr. 2015
ISBN 978-1-77557-683-9 (paperback)
ISBN 978-1-77557-684-6 (Online edition)
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UBLISHED I
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2015 B
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Magnolia Press
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© 2015 Magnolia Press
ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
Zootaxa 3951 (1) © 2015 Magnolia Press
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3
CALCAREOUS SPONGES OF INDONESIA
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Materials and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Systematic descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Phylum Porifera Grant, 1836. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Class Calcarea Bowerbank, 1864 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Subclass Calcinea Bidder, 1898. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Order Clathrinida Hartman, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Family Clathrinidae Minchin, 1900. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Genus Clathrina Gray, 1867 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Clathrina purpurea sp.nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Clathrina chrysea Borojevic & Klautau, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Clathrina heronensis Wörheide & Hooper, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Clathrina beckingae sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Clathrina aff. luteoculcitellaWörheide & Hooper, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Clathrina sororcula sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Clathrina stipitata (Dendy, 1891) comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Genus Arthuria Klautau, Azevedo, Cóndor-Luján, Rapp, Collins & Russo, 2013. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Arthuria tenuipilosa (Dendy, 1905) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Arthuria tubuloreticulosa sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Genus Ernstia Klautau, Azevedo, Cóndor-Luján, Rapp, Collins & Russo, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Ernstia indonesiae sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Ernstia chrysops sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Ernstia klautauae sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Ernstia naturalis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Family Levinellidae Borojevic & Boury-Esnault, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Genus Burtonulla Borojevic & Boury-Esnault, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Burtonulla sibogae Borojevic & Boury-Esnault, 1986. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Family Leucaltidae Dendy & Row, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Genus Ascandra Haeckel, 1872. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Ascandra kakaban sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Ascandra crewsi sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Genus Leucaltis Haeckel, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Leucaltis nodusgordii (Poléjaeff, 1883) comb. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Leucascidae Dendy, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Genus Ascaltis Haeckel, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Ascaltis angusta sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Genus Leucascus Dendy, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Leucascus flavus Cavalcanti, Rapp & Klautau, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Genus Ascoleucetta Dendy & Frederick, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Ascoleucetta sagittata Cavalcanti, Rapp & Klautau, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Family Leucettidae De Laubenfels, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Genus Leucetta Haeckel, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Leucetta chagosensis Dendy, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Leucetta microraphis Haeckel, 1872. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Genus Pericharax Poléjaeff, 1883. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Pericharax orientalis sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Order Murrayonida Vacelet, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Lelapiellidae Vacelet, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Genus Lelapiella Vacelet, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Lelapiella sphaerulifera Vacelet, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Subclass Calcaronea Bidder, 1898. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Order Leucosolenida Hartman, 1958. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Sycettidae Dendy, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Genus Sycetta Haeckel, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Sycetta vinitincta sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Genus Sycon Risso, 1827. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Sycon spec. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Family Grantiidae Dendy, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Genus Leucandra Haeckel, 1872. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Leucandra irregularis (Burton, 1930) comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
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Zootaxa 3951 (1) © 2015 Magnolia Press
Family Jenkinidae Borojevic, Boury-Esnault & Vacelet, 2000. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Genus Anamixilla Poléjaeff, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Anamixilla torresi Poléjaeff, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Anamixilla singaporensis sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Genus Uteopsis Dendy & Row, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Uteopsis argentea (Poléjaeff, 1883) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Family Heteropiidae Dendy, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Genus Sycettusa Haeckel, 1872. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Sycettusa sibogae (Burton, 1930) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Genus Grantessa Von Lendenfeld, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Grantessa borojevici sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Grantessa tenhoveni sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Genus Heteropia Carter, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Heteropia minor Burton, 1930 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Genus Vosmaeropsis Dendy, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Vosmaeropsis grisea Tanita, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Family Amphoriscidae Dendy, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Genus Amphoriscus Haeckel, 1872 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Amphoriscus semoni Breitfuss, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Genus Leucilla Haeckel, 1872. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Leucilla australiensis (Carter, 1886). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Additional Indonesian Calcarea not represented in the present collections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Arthuria darwinii (Haeckel, 1870) comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Leucosolenia sertularia (Haeckel, 1872) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Clathrina flexilis (Haeckel, 1872) comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Grantia capillosa var. longipilis sensu Breitfuss, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Aphroceras caespitosa (Haeckel, 1872) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Eilhardia schulzei Poléjaeff, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Abstract
The calcareous sponges collected during Indonesian-Dutch research projects, incorporated in the collections of the Natu-
ralis Biodiversity Center (formerly the Rijksmuseum van Natuurlijke Historie and the Zoölogisch Museum of the Univer-
sity of Amsterdam), are described and discussed. A total of 37 species were distinguished, of which 16 are new to science,
while several others are very poorly known. The new species are Clathrina purpurea sp.nov., Clathrina beckingae
sp.nov., Clathrina sororcula sp.nov., Arthuria tubuloreticulosa sp.nov., Ernstia indonesiae sp.nov., Ernstia chrysops
sp.nov., Ernstia klautauae sp.nov., Ernstia naturalis sp.nov., Ascandra kakaban sp.nov., Ascandra crewsi sp.nov., As
-
caltis angusta sp.nov., Pericharax orientalis sp.nov., Sycetta vinitincta s p.nov., Anamixilla singaporensis sp.nov., Grant-
essa borojevici sp.nov. and Grantessa tenhoveni sp.nov. An additional six species reported from Indonesia, but not
represented in our material, are briefly characterized.
Keywords: Porifera, Calcarea, new species,South East Asia, Indonesia
Introduction
Indonesia and surrounding region, often named the Coral Triangle (e.g. Hoeksema 2007; 2013), comprises a
generally recognized biodiversity hotspot. For most marine animal groups, the region shows enhanced numbers of
higher and lower taxa, and especially for sessile biota the area is the richest of all oceans (Briggs 1974). Sponges
are probably no exception, although this is obscured by a distinct research effort bias (Van Soest et al. 2013).
Whereas quite a lot of data has been published for the region on the largest sponge class, the Demospongiae Sollas
(1885) (cf. Van Soest 1989; Van Soest 1997; Hooper et al. 2002), the information on diversity in the Coral Triangle
of one of the smaller classes, the rather enigmatic Calcarea Bowerbank, 1864 is largely lacking. In fact, the World
Porifera Database (Van Soest et al. 2015) lists only 12 ‘accepted’ species from Indonesia, which comprise the
species originally described (‘endemics’) from Indonesia. Earlier, Haeckel (1872) described three species from
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Indonesia, Topsent (1897) reported one ‘European’ species, Breitfuss (1896a; 1896b; 1898) listed ten species, five
of which were ‘European’, and finally Burton (1930) listed twenty species, nine of which were ‘European’ names.
These low numbers would indicate that Calcarea are indeed rare and/or inconspicuous in Indonesian waters, as
Breitfuss (1898) concluded. However, from the many images of beautiful and intriguing calcareous sponges
appearing in regional underwater color guides (e.g. Colin & Arneson 1995; Gosliner et al. 1996; Erhardt &
Baensch 1998; Weinberg 2004) and on diver websites, we suspect that the diversity of Calcarea is not poor but
probably comparable to the other marine groups. So far, the identity of many of those beautiful species remains
largely undetermined until now, justifying a study focused on the Calcarea of Indonesia and adjacent countries.
Added inducements for us to attempt this enterprise are the absence of virtually any scientific studies on Indonesian
Calcarea since 1930 and the presence of large numbers of specimens in the collections of the Naturalis Biodiversity
Center.
The taxonomy of calcareous sponges (class Calcarea) has long suffered from inaccessibility to any but a
handful of informed specialists. With the advent of the Calcarea chapters of the Systema Porifera (Borojevic and
colleagues in Hooper & Van Soest 2002), the situation improved greatly by the establishment of a single ‘accepted’
classification down to the level of genera, in particular for subclass Calcinea/Order Clathrinida (Borojevic et al.
2002a) and subclass Calcaronea/Order Leucosolenida (Borojevic et al. 2002b). While recognizing that these
Calcarea chapters are a masterful and admirable effort by Radovan Borojevic and co-authors to bring Calcarea
systematics within reach of the wider sponge community, we also have to conclude that several problems still
remain. Due to the rather concise definitions, limited illustrations, and limited review of species assigned to or
recognized as belonging to the genera and families distinguished in the various groups, the Calcarea chapters do
not function as well as was hoped for. Still, the publication of the Systema Porifera resulted in an enhanced effort
on the taxonomy of calcareous sponges by taxonomists from Brazil-France, Germany-Australia, and Norway,
combining morphological and molecular tools to further disclose the biodiversity of this group (see e.g. Klautau &
Valentine 2003; Manuel et al. 2004; Dohrmann et al. 2008; Voigt et al. 2012; Klautau et al. 2013). In view of these
developments, we judged it timely to present here descriptions of all Indonesian calcareous sponges incorporated in
the collections of the Naturalis Biodiversity Center and to attempt to assign them to their proper affiliation.
Materials and methods
Our material consisted of a mix of old collection specimens, e.g. those of the Siboga Expedition (1899–1900)
described by Burton (1930), specimens collected by the first author during the Indonesian-Dutch Snellius II
Expedition (1984–85), and relatively recent (two-)yearly collections made by the second author (1997–2013).
During these recent fieldwork episodes, our own collecting activities were augmented by incidental samples
collected by our colleagues specialized in other sessile groups. Additionally, several voucher specimens or
fragments sent to us for identification by various natural products research groups were included in our material.
Specimens and fragments are all registered in the sponge collection of the Naturalis Biodiversity Center, either as
RMNH’s Por. four-digit registration number or ZMAs Por. five-digit registration number. The latter concern
specimens previously belonging to the Zoological Museum of Amsterdam. We also borrowed several slides from
the Natural History Museum in London (indicated as BMNH) to assist in our identifications. The combined
collections numbered 155 samples identified as belonging to the class Calcarea, predominantly from localities
within the EEZ of the Republic of Indonesia, but occasionally samples originated from adjacent states Timor Leste,
Singapore, Papua New Guinea, Philippines and Palau. Approximate locations from where the samples were
collected are presented in Fig. 1, demonstrating that the combined samples are representative of most geographic
parts of the Coral Triangle. Precise localities and further details of each specimen are provided in the Material
Examined section given with each species description. Geographic coordinates are in decimal degrees rounded off
to four decimals or less.
Depending of the taxonomic group and the state and size of the specimens, we made thick sections
perpendicularly and/or tangentially, either by hand directly from the alcohol preserved material, or we made
histological sections from a fragment that was first stained, using haemaluin-eosin or fuchsin stains, then was
transferred into paraffin, and subsequently sectioned by hand. Spicule suspensions were made by keeping a
fragment of the material in household bleach for a variable period depending of the thickness and state of
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preservation. The residual suspensions were washed five times in distilled water using a table centrifuge to
precipitate the spicules. Spicules were plated on microscope glass for light microscopy and on SEM stubs for
imaging, using a wide mouthed pipette to make sure all spicule types were included. Illustrations of sections were
made by a Leica DM5500 stacking microscope and of spicules by a JEOL Scanning Electron Microscope. In all
our preparations we made sure that parts of the sponge with different spicule types (cortical, choanosomal, atrial
and fringe skeletons) were represented, in order not to miss any spicule types. Measurements of the spicules were
made under light microscopy. Measurements included smallest—average—largest dimensions of each spicule type
of each specimen, based on 25 randomly selected spicules, unless otherwise indicated. In the case of equiactinal
spicules, measurements are provided of length and thickness of the actines of the triactines and the basal radiate
system of the tetractines, and additionally of the apical actines of the latter. In the case of sagittal spicules, unpaired
and one of the paired actines were measured separately. In the case of pseudosagittal or parasagittal spicules all
actines were measured separately. Apical actines of tetractines were sometimes less easy to measure in the
preparations, so occasionally we provide only the observed range of length and thickness. Terminology for the
skeletal structures and the spicules was taken from the Thesaurus of Sponge Morphology (Boury-Esnault &
Rützler 1997), occasionally expanded with adjectives to draw attention to special spicule features.
FIGURE 1. Map of Indonesia and adjacent countries showing approximate distribution of the sample location. Each white
square represents one or more localities, and one or more specimens. Detailed locality positions are provided in the material
examined data with each species (Free Edition map courtesy www.primap.com)
Results
The specimens available to us in the present study were assigned to a total of 37 species, 24 of which belonged to
the subclass Calcinea, 13 to the subclass Calcaronea. For the classification and the order in which higher taxa are
treated we follow the Systema Porifera chapters of Borojevic et al. (2002a; 2002b; 2002b).
Systematic Descriptions
Phylum Porifera Grant, 1836
Class Calcarea Bowerbank, 1864
Subclass Calcinea Bidder, 1898
Order Clathrinida Hartman, 1958
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Family Clathrinidae Minchin, 1900
Remarks. This family unites the Clathrinida species with an asconoid aquiferous system with simple unfolded
choanoderm. Most species have a cormus consisting of a network of thin-walled tubes lined by an unstructured
layer of mostly small triactines and/or tetractines, with occasional presence of diactines. The family was recently
revised using molecular sequence data (Klautau et al. 2013). This resulted in the erection of new genera
subdividing the former genus Clathrina Gray, 1867 s.l., some of which appeared to be affiliated to other families of
the Clathrinida. The present classification into genera Clathrina s.s., Ernstia Klautau et al., 2013, Arthuria Klautau
et al., 2013, Borojevia Klautau et al., 2013, Brattegardia Klautau et al., 2013, and a restricted Guancha Miklucho-
Maclay, 1868, appears not yet fully operational for classifying former Clathrina s.l. specimens because it is based
primarily on molecular analysis. Some former Clathrina species, are now assigned to Ascaltis Haeckel, 1872
(family Leucascidae). These changes mean in practice that Clathrina-like new species in this group cannot be
easily assigned to their proper genus unless sequence data are available, excepting Clathrina s.s., which remain
morphologically recognizable. Morphological differences of the various genera, such as proportion of triactines
and tetractines, or clathroid versus loosely anastomosed tubules, appear to be rather overlapping and not precise.
We will give informal diagnoses of the genera based on our perception of their variability. These diagnoses should
not be considered as formal definitions. For the classification of the known species we follow here the World
Porifera Database (Klautau in Van Soest et al. 2014), but for the new species the assignment to genera is tentative.
According to the molecular analysis of Klautau et al. (2013), Ascandra Haeckel, 1872 - in the Systema
Porifera assigned to family Leucaltidae Dendy & Row, 1913 - is very probably also a member of Clathrinidae.
Nevertheless, we here continue to assign it to Leucaltidae.
Genus Clathrina Gray, 1867 sensu Klautau, Azevedo, Cóndor-Luján, Rapp, Collins & Russo, 2013
Clathrinidae with a cormus of anastomosed tubes, asconoid aquiferous system, and lacking tetractine spicules
(after Klautau et al. 2013).
Clathrina purpurea sp. nov.
Figures 2a–e
Clathrina sp. Erhardt & Baensch 1998: 22.
Material examined. Holotype RMNH Por. 6625, Indonesia, N Sulawesi, Lembeh Strait, Teluk Makawide,
1.4847°N 125.2406°E, depth 15 m, SCUBA, coll. N.J. de Voogd, #LEM19/090202/064, 9 February 2012.
Description. A thick, seemingly conical (Fig. 2a) mass of thin, loosely anastomosed tubes of 2–3 mm in
diameter, forming an undulated uneven surface. There are frequent oscules (Fig. 2b), several of which are raised,
but water-collecting tubes with terminal oscules are lacking. Total size in life 15 x 10 cm; the preserved specimen
has shrunk to 5 x 5 x 0.5 cm (Fig. 2c). Color a distinctive reddish purple in life, dark red-brown and quite limp in
preserved condition. Consistency soft, easily torn.
Skeleton. The walls of the tubes (Fig. 2d) consist of a thin layer of triactines.
Spicules. Regular triactines only. Triactines (Fig. 2e) equiangular equiactinal, with thin cylindrical actines, 78–
120.3–153 x 5.5–6.1–7 µm.
Ecology. On reefs, at shallow depth.
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FIGURE 2. Clathrina purpurea sp. nov., holotype RMNH Por. 6625, a, habitus in situ at N Sulawesi (photo N.J. de Voogd), b,
detail of surface in situ showing distribution of oscules (photo N.J. de Voogd), c, preserved holotype (scale bar = 1 cm), d,
overview of skeleton (scale bar = 500 µm), e, SEM image of triactine.
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Remarks. This specimen is assigned to Clathrina in the restricted sense of Klautau et al. (2013) on account of
its habitus of anastomosed tubes and continuous choanoderm in combination with its lack of tetractines. Although
there are already numerous Clathrina species decribed, the present species stands out by its distinctive color and
large size, combined with numerous oscules and cylindrical triactines of intemediate size. Using the key of Klautau
& Valentine (2003) we were unable to find a matching species and assume here that it is undescribed. There is
some resemblance in habitus to Clathrina ceylonensis (Dendy, 1905), as redescribed by Klautau & Valentine
(2003: 16, fig. 10) but shape of the spicules of that species (conical actines) and length of the actines (67–96 µm)
are sufficiently distinct from our specimen to belong to a different species.
The photo on p. 22 of the Meeres Atlas 5 of Erhardt & Baensch is obviously this species; like our holotype it
was photographed in Lembeh Strait.
FIGURE 3. Clathrina chrysea Borojevic & Klautau (2000),ZMA Por. 16165 from SW Sulawesi, a, preserved habitus (scale
bar = 1 cm), b, overview of skeleton (scale bar = 1 mm), c, detail of surface skeleton (scale bar = 200 µm), d, SEM images of
spicules.
Clathrina chrysea Borojevic & Klautau, 2000
Figures 3a–d
Clathrina spec. Lévi, Laboute, Bargibant &Menou,1998: 74.
Clathrina chrysea Borojevic & Klautau, 2000: 189, fig. 1.
Material examined. ZMA Por. 16165, Indonesia, SW Sulawesi, Spermonde Archipelago, Samalona, 4.8747°S
119.3419°E, depth 6 m, SCUBA, coll. N.J. de Voogd, 20 April 1997.
Description. Cushion-shaped (Fig. 3a), slightly lobate, cormus of loosely anastomosed tubes. Water-collecting
tubes are lacking. Overall size 4.5 x 3 cm, height 0.5 cm, tubes 2–3 mm in diameter. Consistency soft, limp. Color
in life not noted (presumably yellow), in alcohol pale transparent white.
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Skeleton. (Fig. 3b–c) Wall of tubes thin, at most two or three layers of triactines (Fig. 3c).
Spicules. Triactines only.
Triactines (Fig. 3d), regular equiangular, with conical actines, 75–117.2–144 x 7.5–9.1–11; a minority of the
spicules have one actine crooked, either upwards or downwards, and these are usually smaller than the regular
spicules (average actine length 82 µm).
Ecology. In reefs and reef lagoons, 6–28 m.
Distribution. Indonesia, New Caledonia.
Remarks. The identification of this material with Clathrina chrysea is based on shape of the specimen and
size and shape of the triactines. A similar species appears to be Clathrina heronensis Wörheide & Hooper, 1999
from the Great Barrier Reef, Australia, but its habitus is dissimilar in being flat, tubes are thinner (1 mm), and in
alcohol the color turns brown. Below, we assign a distinctly different specimen of our collection to C. heronensis.
Clathrina heronensis Wörheide & Hooper, 1999
Figures 4–e
Clathrina heronensis Wörheide & Hooper, 1999: 863, figs 3A–F; Klautau & Valentine, 2003: 28, fig. 21.
Material examined. RMNH 4499, Indonesia. Kalimantan, Berau region, Derawan Islands, Maratua Island, Haji
Buang Marine Lake, 2.2044°N 118.5987°E, depth 1–2 m, snorkeling, coll. L.E. Becking, # LE156, 11 September
2008.
Description. (Fig. 4a) Small, flatly encrusting loosely open network of thin tubes. Water collecting tubes or
oscules not apparent. In situ color is transparent white, discoloring to brown in preserved condition (Fig. 4b). Size
of cormus 1.5 x 2 cm, tubes thin, at most up to 2 mm in diameter, usually less than 1 mm. Consistency in preserved
state stiff, fragile.
Skeleton. (Figs 4c–d) Several layers of irregularly arranged triactines.
Spicules. Triactines only.
Triactines (Fig. 4e), mostly equiangular equiactinal, but some are faintly parasagittal, size of conico-cylindrical
actines 93–125.9–162 x 8–9.3–11.5 µm.
Eology. Shallow-water, in marine lake on mangrove roots. Elsewhere in caves.
Distribution. Indonesia, NE Australia.
Remarks. The specimen, which is now broken into two fragments, is closely similar to Australian material
according to descriptions of it in Wörheide & Hooper (1999) and Klautau & Valentine (2003). White color, brown
discoloration in alcohol, cormus shape and spicule size and form of our material fit well with the type.
Clathrina beckingae sp. nov.
Figures 5a–e
?Leucosolenia clathrus; Breitfuss, 1896a: 434; Breitfuss, 1898: 171 (not: Schmidt, 1864).
Material examined. Holotype RMNH Por. 1437, Indonesia, NE Kalimantan, Berau region, Derawan Islands,
Maratua Island, depth 1 m, snorkeling, coll. L. Colin, Kalimantan-Berau Expedition 2003, 23 October 2003.
Paratype RMNH Por. 4482, Indonesia, East Kalimantan Province, Berau Region, Maratua Island, Haji Buang
marine lake,
2.2044°N 118.5987°E, depth 1–2 m, snorkeling, coll. L.E. Becking, #LE 177, 11 September 2008.
Description. (Figs 5a, holotype, and 5b, paratypes). Cormus forming a conical mass of loosely anastomosing
tubes with several water collecting tubes ending in oscules elevated above the surface of the body. Color
transparent white in situ (verging to pale cream in the holotype specimen), becoming pale greyish yellow in
preserved condition (Fig. 5c). Size up to 2.5 x 2 x 2 cm, individual tubes 0.5–1 mm in diameter. Consistincy soft,
limp.
Skeleton. (Fig. 5d) Walls of tubes having only two or three layers of partly overlapping triactines, not densely
arranged.
Spicules. Triactines only.
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FIGURE 4. Clathrina heronensis Wörheide & Hooper (1999), RMNH Por. 4499 from Maratua, a, habitus in situ (photo L.E.
Becking), b, habitus in preserved state (scale bar = 1 cm), c, overview of tubar skeleton (scale bar = 500 µm), d, detail of tube
(scale bar = 200 µm), e, SEM images of spicules.
Triactines (Fig. 5e), equiangular equiactinal, with thin cylindrical actines, size of actines 48–84.9–106 x 4–
4.7–6 µm.
Ecology. Marine lake inhabitant.
Distribution. Indonesia.
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FIGURE 5. Clathrina beckingae sp. nov., a, holotype RMNH Por. 1437, photographed in situ in Maratua marine lake (photo
L. Colin), b, paratypes RMNH Por. 4482, ditto (photo L.E. Becking), c, preserved type material (scale bar = 1 cm), d, skeleton
of tubes (scale bar = 200 µm), e, SEM images of spicules of holotype (three on the left) and paratype (right).
Etymology. Named after our colleague Leontine E. Becking, expert on marine lake biota, to acknowledge her
assistance in the field and for collecting material of several of the species described in this paper, including the
paratype of the present species.
Remarks. By being white and having thin semitransparent tubes the new species resembles Clathrina
heronensis described above, and also occurring in the same habitat. However, there are several important and
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significant differences: (1) the actine length and thickness of the triactines is considerably smaller and the shape is
cylindrical, not conico-cylindrical, (2) distinct water-collecting tubes are present, elevated above the cormus, and
(3) the cormus is forming a conical mass elevated above the substratum, not flattened.
Although Breitfuss (1896a, 1898) did not provide adequate descriptive information or illustrations, the spicule
size data make it possible that his report of Clathrina (as Leucosolenia) clathrus from Ternate concerns the present
species. C. clathrus is restricted to European waters and it is highly unlikely that it would occur in Indonesia.
FIGURE 6. Clathrina aff. luteoculcitellaWörheide & Hooper, 1999, ZMA Por. 08567 from SW Salayar, a, preserved habitus
(scale bar = 1 cm), b, detail of surface skeleton (scale bar = 200 µm), c–d, SEM images of spicules, c, triactines, d, (pieces of
rare) diactines.
Clathrina aff. luteoculcitella Wörheide & Hooper, 1999
Figures 6a–d
? Clathrina luteoculcitellaWörheide & Hooper, 1999: 868, fig. 5; Klautau & Valentine, 2003: 30, fig. 23.
Clathrina luteoculcitella’Indonesia’; Klautau et al. 2013: 12.
Material examined. ZMA Por. 08657, Indonesia, SE Sulawesi, SW Salayar, NW coast of Pulau Guang, 6.35°S
120.45°E, depth 4–12 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 152/III/
40, 29 September 1984.
Description. Dense, partially digitate mass of tightly anastomosed thin tubes (Fig. 6a). Water-collecting tubes
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are lacking. Overall size 2.5 cm long, 1 cm wide, digitations 0.5–1 cm high. Live color noted as pale yellowish, in
alcohol it is beige-yellow.
Skeleton. (Fig. 6b) A dense layer of small triactines, with occasional diactines, but these are invariably broken
off.
Spicules. Triactines, possibly diactines.
Triactines, (Fig. 6c) equiangular equiactinal, with conical actines, 67–87.3–98 x 6–8.1–10 µm; rarely one of
the actines is crooked.
Diactines, (Fig. 6d) not certainly proper, few are present and these consist mostly of short broken pieces, 80
120 x 6 µm.
Ecology. Caves and overhangs in reef environment, 4–25 m.
Distribution. Indonesia, NE Australia.
Remarks. Klautau et al. (2013) listed this material as Clathrina luteoculcitella Wörheide & Hooper, 1999 on
account of the tightly anastomosed mass of fine tubes in combination with the small triactines with relatively
robust conical actines and a presence of diactines. The presence of diactines is a less convincing similarity with the
Australian material as these were only few and mostly broken in our specimen, whereas Wörheide & Hooper
pictured a specimen with lots of diactines sticking from its surface (their fig. 5B). Dr Michelle Klautau (personal
communication) used the present material in her phylogenetic analysis (see Klautau et al. 2013: fig. 1) and it
appeared in the same clade - but at some distance - as the Australian holotype of C. luteoculcitella. We assume here
that it is close to C. luteoculcitella but possibly distinct.
We note here that the specimen is also close to the Sri Lankan Clathrina ceylonensis (Dendy, 1905) (as
Leucosolenia coriacea var. ceylonensis Dendy, 1905: 226, pl.
XIII fig. 8). This also has a cormus of tightly
anastomosed tubes and triactines in exactly the same size and shape. Dendy (1905: 227) noticed diactines in some
of the specimens.
Clathrina sororcula sp. nov.
Figures 7a–c, 8a–e
Leucosolenia coriacea; Burton, 1930: 2, in part (not: Montagu, 1814)
? Clathrina sp. Colin & Arneson, 1996: 59, photo 227; Lim, De Voogd & Tan,2008: 162.
Material examined. Holotype RMNH Por. 2576, Singapore, Pulau Subar Darat (Little Sister), NW side, 2.2149°N
103.8318°E, depth 10 m, SCUBA, coll. N.J. de Voogd, #SIN16/010406/138, 1 April 2006,
Paratypes ZMA Por. 00135, Indonesia, Sulawesi, Salayar anchorage, 6.0963°S 120.4481°E, depth 0–36 m, trawl,
coll. Siboga Expedition stat. 213, 26 September 1899; ZMA Por. 00183a, Indonesia, Kalimantan, Karang Lintang,
Pulau Palabangan, Moearas Reef, 1.7714°N 118.9615°E, depth 0–54 m, trawl, hard coral sand, coll. Siboga
Expedition stat. 091, 22 June 1899; RMNH 2065, Indonesia, Java Sea, Kepulauan Seribu (Thousand Islands),
5.7606°S 106.7546°E, depth 12 m, SCUBA, coll. N.J. de Voogd, #SER.27, Kepulauan Seribu Expedition 2005, 17
September 2005.
Description. Rather loosely anastomosed cushion of white, semi-transparent tubes (Figs 7a, 8e), encrusting rocks
and mussels. Size up to 4 x 3 cm in lateral expansion, at least 1-3 cm in height in preserved condition. Several oscules
are visible in the in situ photo, but they are solitary and flush with the surface and do not appear to be distinct endings
of water-collecting tubes. Pale greyish brown to dirty white in preserved condition (Figs 7b, 8a). The preserved
holotype is broken into two equally sized fragments (Fig. 7b).
Skeleton. (Figs 8b–c) Walls made up of three or more layers of robust triactines.
Spicules. Triactines only.
Triactines (Figs 7c, 8d), equiangular equiactinal, cylindro-conical actines with rather abruptly sharp-pointed ends,
a small proportion are parasagittal, actines in a large size range, possibly divisible in two sizes, overall size 97–151.6
201 x 9–12.6–21 µm.
Ecology. Shallow-water.
Distribution. Singapore, Indonesia.
Etymology. The word sororcula (L.) means ‘little sister’, referring to the locality Pulau Subar Darat, Little Sister
Island.
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FIGURE 7. Clathrina sororcula sp.nov., holotype RMNH Por. 2576, a, photo in situ, from Singapore (photo N.J. de Voogd), b,
preserved fragments (scale bar = 1 cm), c, SEM images of the spicules.
Remarks. The new species differs from the Clathrina species reported here from Indonesian waters by the
combination of white semitransparent color and robust spicules. Using the key of Klautau & Valentine (2003) this
species keys out as the Northeast Australian Clathrina parva Wörheide & Hooper, 1999. The skeleton and the
spicules are indeed similar to the description of the type of that species, but the habitus and the size of the cormus
are quite different: cormus may reach 4 cm in lateral dimension (1 cm in C. parva), oscules are singly and flush (on
the top of fused tubes in C. parva), spicules are conico-cylindrical and sharply pointed (cylindrical and bluntly
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pointed in C. parva). Furthermore, we assume that photo’s in Colin & Arneson (1996: Clathrina sp. 227, Papua
New Guinea) and Lim et al. (2008, Singapore) are also referable to the present species.
FIGURE 8. Clathrina sororcula sp. nov., paratype ZMA Por. 00183a, a, preserved material collected by the Siboga Expedition
(scale bar = 1 cm), b, skeleton of tubes (scale bar = 500 µm), c, detail of the same (scale bar = 200 µm), d, SEM images of the
spicules.
Burton (1930) referred this (and other specimens of the Siboga Expedition) to the European species Clathrina
(as Leucosolenia) coriacea (Montagu, 1814), which is erroneous because that species has much more densely
anastomosed tubes and smaller spicules. Siboga specimens ZMA Por. 00136 and 00183b, identified by Burton
also as Clathrina (as Leucosolenia) coriacea possess tetractines, and are here assigned to Ernstia naturalis sp.nov.
(see below).
Clathrina stipitata (Dendy, 1891) comb. nov.
Figures 9a–b
Leucosolenia stipitata Dendy, 1891: 51, pl. I figs 4–6, pl. IV fig. 2, pl. IX fig. 5
Leucosolenia macleayi; Burton, 1930: 2 (not: Ascetta macleayi Von Lendenfeld, 1885)
Material examined. ZMA Por. 00134, Indonesia, Sulawesi, Karkaralong Islands, anchorage off Kawio and
Kamboling Islands, 4.672°N 125.4015°E, depth 23–31 m, dredged, coll. Siboga Expedition stat. 129, 22 July 1889.
Description. Stalked small sponge, dirty white in alcohol (Fig. 9a). Upper part consists of a tight mass of thin
tubes. Stalk approximately 6 mm long, body 8 x 6 mm. Consistency soft, easily damaged.
Skeleton. The wall of the tubes consist of several layers of triactines (Fig. 9b).
Spicules. (Fig. 9b) Triactines only, in two categories.
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Small equiangular equiactinal triactines, actines 69–76.5–84 x 5–5.9–7 µm.
Small parasagittal triactines (arrows in Fig. 9b), unpaired actines 123–135.8–151 x 66.6–7 µm, paired actines
66–73.8–85 x 5–5.4–6 µm.
Ecology. Deeper water down to 32 m.
Distribution. Indonesia, South East Australia.
Remarks. The identification is not entirely certain as Dendy cites slightly shorter unpaired actines for the type.
Clathrina macleayi (Von Lendenfeld, 1885) has much smaller spicules and is unlikely to be conspecific.
Remarkably, Burton (1963) in his list of specimens/slides in the collection of the Natural History Museum lists a
slide of the Siboga collection (BMNH 1928.6.18.3) as Leucosolenia stipitata, whereas he used the name L.
macleayi in the 1930 description.
Because of the stalked habitus of the present species, it would have been assigned to the genus Guancha in the
Systema Porifera classification (Borojevic et al. 2002a). However, in the recent revision of Klautau et al. (2013)
they demonstrated that the genus is polyphyletic and its type species falls in the same clade as Clathrina.
FIGURE 9. Clathrina stipitata (Dendy, 1891), ZMA Por. 00134, from N Sulawesi, a, preserved habitus (scale bar = 1 cm), b,
detail of surface skeleton showing regular and sagittal triactines (arrows) (scale bar = 100 µm).
Genus Arthuria Klautau, Azevedo, Cóndor-Luján, Rapp, Collins & Russo, 2013
Clathrinidae with asconoid aquiferous system possessing both triactines and tetractines, the latter in low proportion
(after Klautau et al. 2013).
Arthuria tenuipilosa (Dendy, 1905)
Figures 10a–f
Leucosolenia (Clathrina) tenuipilosa Dendy, 1905: 227, pl. XIII fig. 9.
Clathrina tenuipilosa; Klautau & Valentine, 2003: 40, fig. 32.
Arthuria tenuipilosa; Klautau et al. 2013: 12.
Material examined. ZMA Por. 15509b, Indonesia, Indonesia, SE Sulawesi, SW Salayar, NW coast of Pulau
Guang, 6.35°S 120.45°E, depth 3–4 m, SCUBA, coll. H.A. ten Hove, Indonesian-Dutch Snellius II Expedition stat.
152/II/Cave 1/A3, 28 September 1984.
Description. White mass of loosely anastomosed tubes (Fig. 10a), size 3 x 1.5 x 0.5 cm. Water-collecting tubes
are absent. Wall of tubes microscopically echinated by hairy-thin diactines (Figs 10b–c).
Skeleton. (Fig. 10b) The tube walls consist of a thin layer of triactines and tetractines.
Spicules. (Figs 10d–f) Triactines, tetractines, diactines.
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FIGURE 10. Arthuria tenuipilosa (Dendy, 1905), ZMA Por. 15509b.a, preserved habitus (scale bar 1 cm), b, overview of
cormus (scale bar = 500 µm). c, detail of surface with protruding diactines (scale bar = 200 µm), d–f, SEM images of the
spicules, d, triactine, e, tetractine, f, broken diactine, f1, detail of apex of diactine.
Triactines (Fig. 10d) equiangular equiactinal, 81–107.1–119 x 8–8.9–10.5 µm.
Tetractines (Fig. 10e), with equiangular equiactinal basal radiate system, actines 89–103.6–121 x 8–8.7–10
µm, apical actines 62–100 x 6–6.5 µm.
Diactines (Fig. 10f), with slightly swollen endings (Fig. 10f1) on one side and thinly pointed endings on the
opposite side, usually broken, 350–422 x 2.5–3 µm.
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Ecology. On the upper wall of a shallow-water cave.
Distribution. Indonesia, Sri Lanka.
Remarks. The spicule size data of our specimen conform closely to those given by Dendy (1905: 227) for the
type material, but differ from the measurements of Klautau & Valentine (2003: 41) taken from a paratype in having
on average longer and slightly thinner actines. Trichoxeas described by Klautau & Valentine (2003: 41) are thinner
(only 0.3 µm) than the diactines we measured in our material.
Clathrina luteoculcitella Wörheide & Hooper, 1999 also has a fringe of diactines sticking out of the tubewalls,
but that species has no tetractines and the live color is yellow.
Ernstia indonesiae sp.nov. described below has similar habitus and general spiculation, but the actines of both
triactines and tetractines are twice as long and thick, there is a complement of sagittal tri-and tetractines. The
diactines are also longer and thicker and do not stick out.
FIGURE 11. Arthuria tubuloreticulosa sp. nov., holotype RMNH Por. 5547, a, habitus in situ at Ternate (photo N.J. de Voogd),
b, preserved habitus (scale bar = 1 cm), c, surface view of skeleton (scale bar = 200 µm).
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FIGURE 12. Arthuria tubuloreticulosa sp. nov., holotype RMNH Por. 5547, SEM images of spicules, a, triactines, b,
tetractines.
Arthuria tubuloreticulosa sp. nov.
Figures 11a–c, 12a–b
Material examined. Holotype RMNH Por. 5547, Indonesia, Halmaheira, Ternate, Sulamadaha Bay, 0.8661°N
127.3316°E, depth 5–15 m, SCUBA, coll. N.J. de Voogd, #TER.05/261009/1, Ternate-Halmahera Expedition, 26
October 2009.
Description. An orange flattened mass of short oscular tubes (Fig. 11a), connected at the substratum by a basal
tubular network, growing on an encrusting octocoral (Briareum). The erect tubes maybe divided into one or two
side tubes. The preserved specimen (Fig. 11b) is fragmented, but there are several larger pieces that show the short
erect tubes forming the main basis of the specimen. Overall size estimated to be 5 x 3 cm, with small tubes about 2–
4 mm in diameter. Color beige in preserved condition.
Skeleton. (Fig. 11c) The walls of the tubes are thin, approximately 15 µm in thickness. The spicules are
dominated by triactines. The choanoderm lines all internal surfaces.
Spicules. (Figs 12a–b) Triactines, tetractines.
Triactines (Fig. 12a), equiangular, predominantly equiactinal, but a fair number are slightly parasagittal, with
the longer actines often slightly wobbled; most actines are cylindrical with abruptly pointed ends, but small
triactines may have their actines conical, size of actines 63–112.1–138 x 4–5.3–6.5 µm.
Tetractines (Fig. 12b), similar to triactines in size and shape, actines of basal triadiate system, 62–119.5–156 x
4–5.4–6 µm; apical actines 39–132 x 3.5–6.5 µm.
Ecology. Shallow-water reef.
Distribution. Known only from the type locality, Ternate.
Etymology. The name reflects the combination of a network of tubes on the substratum and erect tubes
protruding from it.
Remarks. The dominance of triactines over tetractines, and the normal-shaped apical actines of the tetractines
makes this Clathrina-like species a likely member of the recently erected genus Arthuria. The genus has a few
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Indo-Pacific species, but none of them seem to be close to our new species: South Australian A. dubia (Dendy,
1891) (as Leucosolenia) has much thicker actines (averaging 15–16 µm) and the tetractines are very rare (they were
not mentioned by Dendy, but subsequently reported by Klautau & Valentine, 2003: 26). Red Sea A. sueziana
Klautau & Valentine, 2003 has a different habitus and also much thicker actines. The above-described A.
tenuipilosa possesses diactines.
Haeckel’s (1872) description and figures of Ascaltis darwinii reminds rather strongly of the present species and
at first we assumed that it could be conspecific. However, shapes and sizes of the tri- and tetractines presented by
Haeckel consistently differ: actines are equiangular equiactinal, shorter (80–100 µm) but more robust and conical
(10–12 µm thick) than those of our new species. Clathrina darwinii should be transferred to Arthuria (see below).
Genus Ernstia Klautau, Azevedo, Cóndor-Luján, Rapp, Collins & Russo, 2013
Clathrinidae with asconoid aquiferous system possessing both triactines and tetractines in approximately equal
proportions or tetractines more frequently. The apical actine of the tetractines is long and thin (after Klautau et al.
2013).
Ernstia indonesiae sp. nov.
Figures 13a–c, 14a–d, 15a–e
? Clathrina sp. Colin & Anderson, 1995: 59, photo 228.
Material examined. Holotype ZMA Por. 16659, Indonesia, SW Sulawesi, Spermonde Archipelago, Langkai,
5.019°S 119.063°E, depth 6 m, SCUBA, coll. B.W. Hoeksema, #LK–186, 14 June 1997.
Paratypes ZMA Por. 07934, Indonesia, SE Sulawesi, Tukang Besi Islands, southern reef of Karang Kaledupa,
5.9333°S 123.8°E, depth 2–6 m, snorkeling, coll. J.C. den Hartog, Indonesian-Dutch Snellius II Expedition stat.
016/II/26, 8 September 1984; ZMA Por. 09107, Indonesia, SE Sulawesi, SW Salayar, NW coast of Pulau Guang,
6.35°S 120.45°E, depth 0–2 m, snorkeling, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat.
152/Cave 2/02, 28 September 1984; ZMA Por. 09486, Indonesia, SE Sulawesi, SW Salayar, NW coast of Pulau
Guang, 6.35°S, 120.45°E, depth 2–4 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition
stat. 152/Cave 1/06A, 28 September 1984; ZMA Por. 15509c, Indonesia, SE Sulawesi, SW Salayar, NW coast of
Pulau Guang, 6.35°S 120.45°E, depth 3–4 m, SCUBA, H.A. ten Hove, Indonesian-Dutch Snellius II Expedition
stat. 152/Cave 1/A3, 28 September 1984.
Description. Rounded mass of interwoven tubes (Fig. 13a), individual tubes 1–3 mm in diameter. In the one in
situ photo several oscules at the end of water-collecting tubes are apparent but these are not visible in the preserved
material (Figs 13b, 15a). Live color pinkish white, in alcohol pale orange. Overall size of specimens up to 5 x 4.5 x
2 cm.
Skeleton. (Figs 13c, 15b) Walls of tubes formed by a one or two layers of triactines and tetractines arranged
without apparent order.
Spicules. (Figs 14a–d, 15c–e) Triactines, tetractines, diactines.
Triactines (Figs 14a, 15d), equiangular equiactinal, in a large size range, but no clear division in larger and
smaller categories, 90206.7–269 x 10–17.3–22 µm
Tetractines (Figs 14b–d, 15c), less diverse in size, but divisible into those with regular equiangular equiactinal
basal radiate systems, actines 85–214.2–249 x 12–17.8–22 µm, apical actines long and sometimes wavy, 57–
124.1–204 x 6–10.7–14 µm; and parasagittal tetractines with unpaired actines, straight, 198–210 x 12–14 µm,
paired actines usually curved or wavy, 137–159 x 12–14 µm, apical actines 50–60 x 10 µm.
Diactines (Fig. 15e), uncommon, curved, usually with one end thicker and mucronate, (Fig. 15e1) the other
end tapering gradually into a thin straight or curved ending, usually broken into fragments, 200–345–600 x 2–4.6
9 µm.
Ecology. Reefs, shallow depth.
Distribution. Indonesia.
Etymology. Named after the country Indonesia.
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FIGURE 13. Ernstia indonesiae sp. nov., holotype ZMA Por. 16659 from Langkai, SW Sulawesi, a, habitus in situ (photo
B.W. Hoeksema), b, preserved holotype (scale bar = 1 cm), c, overview of skeleton (scale bar = 500 µm).
Remarks. In the key of Klautau & Valentine (2003), this material keys out as the Japanese Clathrina
sagamiana (Hôzawa, 1929) (p. 281, pl. XIII figs 1–2, textfig. 1), assigned to Ernstia in Klautau et al. 2013. There
are considerable differences between E. sagamiana and our specimens: the triactines of the former are smaller, the
apical actines of the tetractines are longer, and the diactines are also longer. Parasagittal tetractines were not
mentioned by Hôzawa. A further problem is that our material is all from shallow-water reefs and reef caves,
whereas the Japanese material was collected at 171 m depth.
It is likely that our specimens are conspecific with Clathrina sp. sensu Colin & Arneson (1996: photo 228)
from Papua New Guinea.
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FIGURE 14. Ernstia indonesiae sp. nov., holotype ZMA Por. 16659, SEM images of the spicules, a, triactine, b, tetractine, c,
detail of apical actine of tetractine, d, small tetractine.
Ernstia chrysops sp. nov.
Figures 16a–c, 17a–e
?Clathrina sp. Gosliner et al. 1996: 15, photo 1.
Material examined. Holotype RMNH Por. 1773, Indonesia, North Sulawesi, Manddin, between Bunaken and
Manado Tua, 1.612°N 124.7322°E, depth 20 m, SCUBA, coll. N.J. de Voogd, #MD04/180502/092, 18 May 2002.
Description. Mass of loosely anastomosed golden yellow tubes (Figs 16a–c) with few but prominent water-
collecting tubes (Figs 16a–b) some of which protrude 1 cm or more above the cormus. Size of cormus up to 5 x 6 x
1 cm. Individual tubes up to 5 mm in diameter. Oscules about as wide as the tubes (Fig. 16c), not flaring. In
preserved condition (Fig. 17a) oscules are not clearly visible and the specimens get a pale yellowish orange color
and have a ‘glassy’ outlook. Consistency soft.
Skeleton. (Fig. 17b) The walls of the tubes have a thin layer of triactines and tetractines, in equal proportion,
with apical actines of the larger tetractines projecting into the lumen of the tubes.
Spicules. (Figs 17c–e) Triactines, tetractines; a few small diactines were found.
Triactines (Fig. 17c) equiangular equiactinal, in a large size range, possibly divisible in smaller and larger,
overall actine sizes 75–210.3–330 x 9–13.8–18 µm (smaller (Fig. 17c1): 75–213 µm, larger (Fig. 17c): 255–330
µm); occasionally a triactine occurs with one of the actines angularly bent.
Tetractines (Fig. 17d), regular, on average larger than the triactines, likewise possibly divisible in smaller and
larger spicules, overall the upper sizes somewhat larger than the triactines, with actines of the basal triradiate
system 186–362.4–661 x 14–25.5–48 µm (smaller: Fig. 17d1): 186–276 µm, larger (Fig. 17d): 300–661 µm),
apical actines straight or more often curved (Fig. 17e), 18–45.9–122 x 4–10.1–18 µm.
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FIGURE 15. Ernstia indonesiae sp.nov.,paratype ZMA Por. 07934 from Karang Kaledupa, SE Sulawesi, a, preserved habitus
(scale bar = 1 cm), b, overview of skeleton (scale bar = 1 mm), c–e, SEM images of the spicules, c, various tetractines, d,
various triactines, e, diactines, e1, detail of apex.
Ecology. Deeper part of the reef.
Distribution. Indonesia; probably Philippines.
Etymology. The word chrysops (Gr.) means shining like gold, referring to the color.
Remarks. This species is close to Ernstia indonesiae sp.nov. in shape, but the color and the large size range in
smaller and larger spicules with upper size of the tetractines to reach 660 µm, the relatively wider tubes, which
have a ‘glassy’ outlook (preserved material), and the absence of diactines together appear solid indications of
distinctness. It is likely that the Philippine Clathrina sp. (photo 1) of Gosliner et al. 1996 belongs to the present
species.
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FIGURE 16. Ernstia chrysops sp. nov., holotype RMNH 1773 (photos N.J. de Voogd), a, in situ at N Sulawesi, b, ditto, c,
detail of cormus and oscule.
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FIGURE 17. Ernstia chrysops sp. nov., holotype RMNH Por. 1773, a, preserved holotype from N Sulawesi (scale bar = 1 cm),
b, surface skeleton showing large diversity in size of tetractines (scale bar = 200 µm), c–e, SEM images of spicules, c, large
triactine, c1, small triactine, d, large tetractine, d1, small tetractine, e, detail of large tetractine,.
Our collection also contained a flimsy fragment from deep water (240 m) off the coast of Sumba (ZMA Por.
09219, coll. Indonesian-Dutch Snellius II Expedition stat. 60), resembling in skeletal and spicular characters the
present new species: tetractines with basal triradiate system having actines up to 350 x 12 µm and strongly
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upcurved apical actines of approximately 100 µm, triactines on average smaller than the tetractines. In view of the
depth and the tiny size we hesitatingly assign this specimen to the present species.
FIGURE 18. Ernstia klautauae sp.nov., holotype ZMA Por. 08390 from Komodo, a, preserved habitus encrusting a dead coral
(scale bar = 1 cm), b, paratype RMNH Por. 9341, habitus in situ (Photo L. E. Becking), c, detail of cormus of holotype showing
apical actines of tetractines protruding in the atrial lumen (scale bar = 200 µm).
Ernstia klautauae sp. nov.
Figs 18a–c, 19a–c
Clathrina sp.nov. 13 ‘Indonesia’; Klautau et al. 2013: 5, table 2.
Material examined. Holotype ZMA Por. 08390, Indonesia, Nusa Tenggara, E of Komodo, Selat Linta, 8.5833°S
119.57°E, depth 4–11 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 079/III/
15, 18 September 1984.
Paratype RMNH Por. 9341, Indonesia, Papua, Misool, Raja Ampat, 1.984°S 130.5164°E. depth 0–2 m,
snorkeling, coll. L.E. Becking, # LE 652/MIS01/RIMG0679, 20 May 2011.
Description. Tightly anastomosed mass of thin tubes (Figs 18a–b). Tubes 1–2 mm in diameter. The in situ
photo (Fig. 18b) shows two water-collecting tubes with terminal oscules elevated above the cormus. Overall size 2
x 1 cm. Color (pale) yellow. Consistency soft.
Skeleton. Tetractines are dominating in the holotype, while tri- and tetractines are about equally present in the
paratype. Apical actines of tetractines protrude far and characteristically into the lumen between the tubes (Fig.
18c).
Spicules. (Figs 19a–c) Triactines, tetractines.
Triactines (Fig. 19a), with conical actines, in a wide range of thicknesses, overall size 93–108.4–125 x 6–8.8
11 µm.
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FIGURE 19. Ernstia klautauaesp.nov., holotype ZMA Por. 08390, SEM images of spicules, a, triactine, b, normal tetractine, c,
tetractines with long thin apical actines.
Tetractines, with conical actines of basal triadiate system 60–106.4–126 x 6–7.9–10 µm, apical actines
variable, 32–90.4–165 x 4–5.4–7 µm. On the basis of the apical actines these spicules are possibly divisible in two
types, with relatively thick straight apical actines (Fig. 19b), and with thin sharply pointed apical actines (Fig. 19c),
straight or slightly curved.
Ecology. Shallow coral reefs, marine lakes.
Distribution. Indonesia (Komodo, Papua).
Etymology. Named after Dr Michelle Klautau (Universidade Federal do Rio de Janeiro, Brazil) for her
pioneering efforts in calcarean taxonomy and phylogeny.
Remarks. The new species is assigned to Ernstia, because of proportion of tri- and tetractines (equal or more
tetractines) and the sharp thin apical actines of the tetractines. DNA of the present holotype material was extracted
by Klautau et al. (2013: table 2) and its sequence classified closely with Ernstia tetractina (Klautau & Borojevic,
2001) (Klautau et al. 2013: Fig. 1).
The habitus reminds of Norwegian Clathrina cribrata Rapp, Valentine & Klautau, 2001 but that species has
triactines only.
Ernstia naturalis sp. nov.
Figures 20a–d, 21a–d
Leucosolenia coriacea; Burton, 1930: 2, in part (not Montagu, 1814).
?Clathrina sp.; Erhardt & Baensch, 1998: 20.
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FIGURE 20. Ernstia naturalis sp. nov., a, holotype RMNH Por. 9342 in situ (photo L.E Becking), b, paratype RMNH Por.
5001 just after collecting (still under water) (photo L. E Becking, c, paratype RMNH Por. 5001, preserved fragments (scale bar
= 1 cm), d, paratype RMNH Por. 5001, overview of tubar skeleton (scale bar = 200 µm).
Material examined. Holotype RMNH Por. 9342, Indonesia, Papua, Misool, Raja Ampat, Panah Panah, marine
lake, 1.9386°S 130.3744°E, depth 0–2 m, snorkeling, coll. LE Becking, # LE745/MIS1/RIMG0679, 28 May 2011.
Paratypes ZMA Por. 00136, Indonesia, Nusa Tenggara, Sumbawa, Saleh Bay, anchorage E of Dangar Besar,
8.4254°S 117.7296°E, depth 0–36 m, trawl, coll. Siboga Expedition stat. 313, 14 February 1900; ZMA Por.
00183b, Indonesia, Papua, 1.7083°S 130.7916°E, depth 23 m, trawl, coll. Siboga Expedition stat. 164, 20 August
1899; RMNH 5000 Indonesia, East Kalimantan Province, Berau Region, Kakaban island, Kakaban marine
lake,
2.4133°N 118.5078°E, depth 0–2 m, snorkeling, coll. L.E. Becking #KKB/LE 536, 21/05/2009; RMNH
5001, Indonesia, East Kalimantan Province, Berau Region, Kakaban island, Kakaban marine lake, 2.4133°N
118.5078°E, depth 0–2 m, snorkeling, coll. L.E. Becking, #KKB/LE 537, 21 May 2009; RMNH Por. 9343, Papua
New Guinea, E of Wongat Island, Stat. PR53, 5.1353°S, 145.8228°E, depth 20 m, SCUBA, coll. L.E. Becking ,
#LE305/PB174146, 17 November 2012.
Description. (Figs 20a–c) Large, massive cormi made up of closely anastomed but individually distinct tubes.
Overall size up to 4–5 cm, tubes 0.3–1.5 mm in diameter. No distinct water collecting tubes, but several large flush
oscula are visible in the in situ photo of the holotype (Fig. 20a). Color in life, bright yellow (Fig. 20b), orange-
yellow or brownish yellow. Beige in preserved condition (Fig. 20c). Consistency soft.
Skeleton. (Fig. 20d) Layers of tri- and tetractines rather dense and thick. Triactines and tetractines in
approximately similar proportion (most specimens) or with more triactines (ZMA 00183b).
Spicules. (Figs 21a–d) Triactines and tetractines.
Triactines (Fig. 21a), with conical actines, 76–107.5–168 x 7.5–9.1–18 µm.
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FIGURE 21. Ernstia naturalis sp. nov., a, holotype RMNH Por. 9342, images of the spicules, a, triactine, b–c, tetractines, d,
detail of apical actine of tetractine.
Tetractines (Figs 21b–d), with actines of basal triaradiate system, 84–109.6–153 x 6–8.716 µm, apical
actines, thin and sharp (Figs 21c–d), 29–82.4–147 x 3–5.3–8 µm.
Ecology. In marine lakes and on reefs, from the surface down to 36 m.
Distribution. Indonesia, Papua New Guinea; possibly Palau (Erhardt & Baensch, 1998).
Etymology. Named after our host institute, Naturalis Biodiversity Center, Leiden.
Remarks. Among the Ernstia species reported from the Indo-West Pacific, Ernstia adusta (Wörheide &
Hooper, 1999 as Clathrina) is the most similar in shape and spicule sizes. However, that species is white, whereas
all our specimens of which color information was noted are yellow or yellow-brown in life.
Several specimens were found to contain isolated broken pieces of diactines, but these were insufficiently
common to consider them proper to the sponge.
Family Levinellidae Borojevic & Boury-Esnault, 1986
Genus Burtonulla Borojevic & Boury-Esnault, 1986
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Burtonulla sibogae Borojevic & Boury-Esnault, 1986
Figures 22a–c, 23a–f
Dendya prolifera; Burton, 1930: 2, figs 1–2; Colin & Arneson, 1995: 61, photo 234 (not: Dendy, 1913: 6)
Burtonulla sibogae Borojevic & Boury-Esnault, 1986: 447, text–fig. 2, pl. 2 figs A–D.
FIGURE 22. Burtonulla sibogae Borojevic & Boury-Esnault (1986), a, in situ image of RMNH Por. 1653 from the Palau
Islands (photo N.J. de Voogd), b, preserved holotype ZMA Por. 00146 (scale bar = 1 cm), c, in situ image of RMNH 1821 from
Maratua Island (photo B.W. Hoeksema).
Material examined. Holotype ZMA Por. 00146 (Fig. 22b), Indonesia, Lesser Sunda Islands, E coast of Roti
Island, Papela Bay, 10.63°S 123.42°E, depth 22 m, trawl, Siboga Expedition stat. 301, 30 January 1900.
Paratype ZMA Por. 00145, Indonesia, Irian Jaya, 1.71°S 130.79°E, depth 32 m, dredge, coll. Siboga
Expedition stat. 164, 20 August 1899.
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FIGURE 23. Burtonulla sibogae Borojevic& Boury-Esnault (1986), a, holotype ZMA Por. 00146, overview of skeleton (scale
bar= 500 µm), b, detail of skeleton of diverticula and tube wall (scale bar = 200 µm), c–f , SEM images of spicules, c, large
triactine of the tube wall, d, small triactines of the diverticula, e, large tetractine of the tube wall, f, small tetractines of the
diverticula.
Additional specimens. RMNH Por. 1653, Palao Islands, channel between Koror and Babeldaob, SW side near
Itelblong Island, 7.34°N 134.517°E, depth 12 m, SCUBA, coll. N.J. de Voogd, #KOR04/130505/029, 13 May
2005; RMNH Por. 1821, Indonesia, Kalimantan, Berau region, Maratua, depth not recorded, barrier reef, coll. L.
De Senerpont Domis, #BER28/171003/206; RMNH Por. 1824, Indonesia, Bali, Tulamben area, Temple Bay E,
8.27°S 115.6°E, depth 20 m, SCUBA, coll. N.J. de Voogd, #BAL24/140401/192, 14 April 2001; RMNH 1922,
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Indonesia, Nusa Tenggara, Bali, N side of Nusa Pendida, off Desa Byuk, 8.6736°S 115.5436°E, depth 15–20 m,
deep reef slope with patches of sand, SCUBA, coll. N.J. de Voogd, #BAL34/NV/210401/263, 21 April 2001;
RMNH Por. 1924, Indonesia, North Sulawesi, Manddin, between Bunaken and Manado Tua, depth 18 m, SCUBA,
coll. N.J. de Voogd, #MD04/190502/117, 19 May 2002; RMNH 2158, Indonesia, Sulawesi, Spermonde
Archipelago, Kundingareng Keke, 5.642°S 119.74°E, depth 10–15 m, SCUBA, coll. N.J. de Voogd, #UP/KK/
180500/054, 18 May 2000; RMNH 9195, Indonesia, Halmahera, Tidore Dea Tahua, 0.7528°N 127.392°E, depth
10–15 m, SCUBA, coll. N.J. de Voogd, #TER07/281009/, Ternate-Halmahera Expedition 2009, 28 October 2009;
RMNH Por. 9344 Indonesia, Papua, Raja Ampat, Pulu Wai East, reef, 0.6999°S 130.6666, depth 10–20 m,
SCUBA, coll. L.E. Becking, #RAJ61/LE222/245, Naturalis-LIPI 2007 Expedition, 11 December 2007; RMNH
Por. 9345, Indonesia, Papua, Raja Ampat, Pulu Wai East, reef, 0.6999°S 130.6666, depth 10–20 m, SCUBA, coll.
L.E. Becking, #RAJ61/LE233, Naturalis-LIPI 2007 Expedition, 11 December 2007.
Description (Figs 22a–c) Very characteristic yellow or pale light brown masses of small rounded globules
hiding for the most part a central tube, which usually protrudes slightly at the top. Basically, the cormus consist of
a single tube, ending in a small oscule with slightly raised rims, with side outgrowths (diverticulae or side-canals)
taking the form of clusters of globules reminding of the alveolae of human longs. We propose ‘alveolae’ as the term
for these globules. Inbetween the clusters of these alveolae the walls of the tube and its diverticulae are here and
there visible as a smooth membrane. Size of individuals may be up to 8 x 5 x 5 cm. Consistency soft.
Histology. Choanocytes are confined to the inner surfaces of the alveolar outgrowths, absent from the walls of
the tube (see extensive description of Borojevic & Boury-Enault, 1986).
Skeleton. (Figs 23a–b) The alveolae possess a skeleton consisting of a single layer of small equiangular and
equiactinal triactines and tetractines. The atrial wall possesses a single layer of large equiangular and sagittal
triactines and tetractines, the latter dominating.
Spicules. (Figs 23c–f) Large and small triactines and tetractines.
Large triactines (Fig. 23c), thin, cylindrical, slightly inequiactinal, but not truly sagittal, actines 186–226.4
306 x 7–7.8–9 µm
Large tetractines (Figs. 23e), thin, cylindrical, similarly equiactinal, except for apical actines, actines of the
basal triradiate system 192–270.3–390 x 6–7.410 µm, apical actines 84–330 µm x 4–6 µm.
Small triactines (Figs. 23d), usually slightly inequiactinal, 30–82.6–138 x 3.5–5.46.5 µm.
Small tetractines (Figs 23f), actines of the basal triradiate system 81–103.2–148 x 4–5.6–7 µm, apical actines
27–51.9–84 x 3–3.7–5 µm.
Ecology. Deeper parts of the reef.
Distribution. Indonesia, Palau.
Remarks. Burton (1930) in his work on the Calcarea of the Siboga Expedition assigned this species to Dendya
prolifera Dendy, 1913 (now Levinella prolifera), but this is clearly a different species with less prominent alveolar
diverticulae, and the choanocytes distributed all over the inner atrial surface. The present species was described by
Borojevic & Boury-Esnault (1986) on the basis of the misidentified Siboga specimens. It appears to be rather
common in Indonesia and surrounding regions, as it is pictured in several underwater guides.
Family Leucaltidae Dendy & Row, 1913
Genus Ascandra Haeckel, 1872
Remarks. The genus is assigned to Leucaltidae in the Systema Porifera, but it resembles loosely built Clathrinidae,
with the added peculiarity that the continuous choanoderm is folded over the long apical actines of the tetractines.
The latter occur usually in larger proportions than the triactines. Skeleton not differentiated in cortical and atrial
skeleton. In the molecular sequence analysis of Klautau et al. (2013) members of the genus appeared to end up with
the Clathrinidae clade.
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FIGURE 24. Ascandra kakaban sp. nov., a, holotype RMNH Por. 1696, in situ photo from Kakaban Island (photo N.J. de
Voogd), paratype RMNH Por. 4625, in situ photo from Kakaban Island (photo L.E. Becking), c, specimen not collected, in situ
photo from Kakaban Island (photo Lori Colin), d, holotype RMNH Por. 1696, reserved fragments (scale bar = 1 cm), e, largest
preserved fragment of holotype (scale bar = 1 cm).
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FIGURE 25. Ascandra kakaban sp. nov., a–b, paratype RMNH Por. 4625, a, light microscopy overview of skeleton of tube
(scale bar = 200 µm), b, detail of atrial surface showing apical actines of the tetractines (arrows) protruding in the atriual lumen
(scale bar = 100 µm), c-e, holotype RMNH Por. 1696, SEM images of the spicules, c, triactines, d, tetractines, e, detail of apical
actine of larger tetractine.
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Ascandra kakaban sp.nov.
Figures 24a–e, 25a–e
Material examined. Holotype RMNH Por. 1696, Indonesia, Kalimantan, Berau Islands, Kakaban island,
2.1409°N 118.5112°E, depth 0.5 m, snorkeling, coll. N.J. de Voogd, #BER08/121003/123, 12 October 2003.
Paratype RMNH 4625, Indonesia, East Kalimantan Province, Berau Region, kakaban island, Kakaban marine
lake, 2.4133°N 118.5078°E, depth 0–2 m, snorkeling, coll. L.E. Becking, #LE043, 31 August 2008.
Description. The cormus (Figs 24a–c) consists of a single (holotype, Fig. 24a) or several (paratype, Fig. 24b)
wide semi-transparent tubes, which are probably water collecting tubes, issuing from a network of relatively few
thinner tubes. In some, possibly ageing, individuals, like the holotype, these tubes at their base proliferate into a
cluster of intertwined ‘blind’ tubules, more brightly white colored, but in other individuals (paratype) this is less
developed. Size of the cormus in preserved condition (Figs 24d–e) up to 7.5 x 7 x 3 cm, largest fragment 4 x 3.5 x
3 cm (Fig. 24e).
Skeleton. (Figs 25a–b) The wall of the tubes is built of a thin layer of triactines and tetractines (Fig. 25a), with
the apical actines of the tetractines directed into the tubar lumen (Fig. 25b, arrows).
Spicules. (Figs 25c–e) Triactines, tetractines.
Triactines (Figs 25c), equiangular equiactinal, in a large size range, possibly divisible in two overlapping size
classes, actines thinly conical, tapering to sharp apices, overall length of actines, 57–154.0–220 x 4–9.4–12 µm.
Tetractines (Figs 25d), similar in shape to triactines but distinctly larger, actines of the basal triradiate system,
74–197.4–267 x 8.5–12.4–16 µm, apical actines curved, often wobbly (Fig. 25e), 35–110.3–237 x 4.5–10.3–16
µm.
Ecology. So far known only from a marine lake, at shallow depth on mangrove roots.
Distribution. So far known only from Kakaban.
Etymology. Named after the type locality, the island of Kakaban, one of the famous marine lakes of Indonesia.
Remarks. The new species is assigned to the genus Ascandra on account of the combination of independent
central tubes with proliferated ‘blind’ side tubes, and tetractines lining and echinating the inner tube surfaces with
their long curved apical actines. We were unable to demonstrate the presence of a folded choanoderm, but assume
the prominent apical actines protruding far into the tube lumen to be sufficient circumstantial evidence for
membership of Ascandra.We were able to compare the present species with a specimen of the type species of
Ascandra, A. falcata Haeckel, 1870 from NE Spain (ZMA Por. 14591). Shape, color and spiculation are different:
A. falcata forms more definitely a group of (yellow) tubes and possesses strongly curved diactines in addition to
the triactines and tetractines. It is similar to the present new species in being semitransparent and having the tri- and
tetractines approximately similar in size - although more robust - and likewise it has the tetractines larger than the
triactines.
The Indonesian species originally described by Haeckel (1872) as Ascandra sertularia, subsequently
reassigned to Leucosolenia by Dendy & Row (1913), on paper appears somewhat similar in habitus to our new
species. However, Haeckel clearly describes large, peculiarly ornamented diactines arranged perpendicular to the
surface, which are not present in our material.
The habitus of our new species is quite striking and distinct, but there is a strong resemblance to Ascandra
crewsi sp.nov. described below. The difference between the two is found mostly in the rarity or virtual absence of
triactines in A. crewsi sp.nov., whereas the two spicule types are in equal proportions in the present species.
Ascandra crewsi sp. nov.
Figures 26a–c, 27a–e
Leucosolenia sp.; Ralifo, et al., 2007: 7.
Material examined. Holotype ZMA Por. 17556, Papua New Guinea, Wahoo, 10.2518°S 150.758°E, depth 21 m,
in cave, SCUBA, coll. R. Sonnenschein #02130, 29 May 2002.
Description. (Figs 26a–b) The material consists of two samples showing thin-walled tubes with extensive
system of thin side branches forming a reticulation around the central tube. Size of the whole specimens in
preserved condition (Fig. 26b) up to 10 x 4 x 3 cm, central tubes less than 1 cm in diameter and branches of the
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network 1–2 mm in diameter. The network of side branches is frequently dead-ended, and these ends taper to a fine
point. The central tube ends in a wide oscule. Color snow-white, but the central tubes are semi-transparent.
Consistency soft, becoming limp out of the water, easily damaged.
FIGURE 26. Ascandra crewsi sp. nov., holotype ZMA Por. 17556 from Papua New Guinea, a, habitus in situ (photo L. E.
Becking), b, preserved holotype (scale bar = 1 cm), c, oblique tangential view of skeleton of atrial surface showing the
protruding apical actines of the tetractine spicules (scale bar = 500 µm).
Skeleton. The wall of the central tube and the side branches is formed by a single layer of tetractines, with the
long thin apical actines protruding far out into the tubar lumen (Fig. 26c).
Spicules. (Figs 27a–e) Predominantly tetractines, a few triactines.
Large tetractines (Figs. 27a) equiangular equiactinal, with actines of the basal triradiate system 159–206.4–246
x 1518.8–21 µm, apical actines usually curved at the end (Fig. 27d), 181–226.3–279 x 13–15.3–17 µm.
Small tetractines (Figs 27c), likewise equiangular equiactinal, with actines of the basal triadiate system 54–
90.2–117 x 7–7.3–8 µm, apical actines straight and needle-sharp (Fig. 27e), 62–95.8–114 x 2.5–3.1–3.5 µm.
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FIGURE 27. Ascandra crewsi sp.nov.,holotype ZMA Por. 17556, SEM images of the spicules, a, large tetractine, b, rare
triactine, c, small tetractine, d, apical actine of large tetractine, e, apical actine of small tetractine.
Small triactines (Fig. 29b), equiangular equiactinal, relatively robust, only a few were observed, actines 140–
150 x 10–12 µm.
Ecology. Deeper part of the reef.
Distribution. So far known only from Papua New Guinea.
Etymology. Named after Dr. Phil Crews of the University of California at Santa Cruz, who leads the research
group that was responsible for collecting the present specimens, and to acknowledge his great contributions to
sponge natural products detection and elucidation.
Remarks. The specimens from Papua New Guinea are considered to be a distinct species despite the overall
similarity to Ascandra kakaban sp.nov. described above. The present material has a larger cormus and there are
distinct differences in the spicules: triactines are virtually missing (only a few could be found and these were only
small ones), while the tetractines were more definitely in two size classes with the large tetractines having much
longer apical actines than those of A. kakaban sp.nov. Like in the Kakaban species, the present specimens look
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deceptively like Clathrinidae, but the central tube and reticulated side branches, the dominance of tetractines with
their long apical actines protruding far into the atrial lumen, and the paucity of triactines, are indicative of
Ascandra. There is also a resemblance with regional species of the genus Soleneiscus Borojevic, Boury-Esnault &
Vacelet, 1990 like S. radovani Wörheide & Hooper, 1999 and S. stolonifer (Dendy, 1891), from which the new
species differs in the lack of diactines and the reticulated habitus.
As discussed above, Ascandra sertularia Haeckel (1872) is distinct by having large diactines.
The present material was preliminarily identified as Leucosolenia sp. by one of us (RVS), under which
name the presence of novel secondary metabolites - leucosolenamides - were reported (Ralifo et al. 2007).
Genus Leucaltis Haeckel, 1872
Leucaltis nodusgordii (Poléjaeff, 1883) comb. nov.
Figures 28a–c, 29a–d, 30a–e
Heteropegma nodusgordii Poléjaeff, 1883 (in part, only the Torres Strait material): 45, pl. I fig. 7, pl. IV figs 1a–d; Dendy,
1905: 230;Jenkin, 1908: 453, fig. 103.
?Leucaltis bathybia var. mascarenica Ridley, 1884: 628, pl. LIV figs a, a’.
Leucaltis clathria sensu Dendy, 1913: 16, pl.2 figs 1–2; Hôzawa, 1940: 136, pl. VI fig. 3; Tanita, 1943: 394, pl. XIII fig. 27;
Borojevic & Klautau, 2000: 190, figs 2–3.
(not: Haeckel, 1872: 159, pl. 29 figs 3a–3c).
Leuconia paloensis; Colin & Arneson, 1995: 61, photo 235; Gosliner et al. 1996: 17, photo 8; Erhardt & Baensch, 1998 Atlas
4: 21, 24–25 (not: Tanita, 1943).
Material examined. RMNH Por. 1772, Indonesia, North Sulawesi, Negeri, Manado Tua South, depth 20 m,
SCUBA, coll. N.J. de Voogd, #MD08/160502/043, 16 May 2002 (several individuals); ZMA Por. 17557, Papua
New Guinea, Normanby Island, N point, 9.7328°S 150.7402°E, depth 21 m, SCUBA, coll. R. Schonnenschein,
#02136, 30 May 2002.
Examined for comparison. BMNH 1884.4.22.23a, two slides labeled ‘Type’ and ‘from type’, ‘Challenger
Torres Straits, coll. Brit.Mus. 27’ (subsequently labeled as Leucaltis clathria); ZMA Por. 12443, Seychelles,
Amirante Islands, Poivre Atoll, N rim, 5.7333°S 53.3167°E, depth 7–8 m, SCUBA, coll. R.W.M. van Soest,
Netherlands Indian Ocean Program E stat. 768/08, 31 December 1992; ZMA Por. 16248, Seychelles, Mahé, SE
coast, Anse Royale Bay, 4.7333°S 55.5167°E, depth 2–13 m, SCUBA, coll. R.W.M. van Soest, Netherlands Indian
Ocean Program E stat. 740/04, 24 December 1992; RMNH Por. 9314, off Guyana, 7.7°N 57.5°W, depth 65 m,
dredged, bottom muddy sand and shells, coll. ‘Luymes’ Guyana Shelf Expedition stat. 107, 5 September 1970.
Description. A clathrate mass of anastomosing tubes (Figs 28a, c), largest individual 12 x 6 x 6 cm, individual
tubes quite variable in length and diameter, undivided tube lengths up to 2.5 cm, diameter 2–8 mm. Tubes ending in
oscules, as wide as the tube (upright) or more often smaller (flush with the surface); oscules naked. Surface smooth,
consistency brittle but somewhat compressible. Color white or pinkish white, lavender-colored, becoming
yellowish white in preserved condition (Fig. 28b).
Histology. The choanocyte chambers (Figs 29b–d) are long, broad and branching, somewhat inbetween
syconoid and sylleibid.
Skeleton. (Figs 29a–d) Cortical skeleton formed by the basal triradiate system of giant tetractines mixed with
giant triactines (Figs 29a, c–d). Actines of the giant tetractines and triactines protrude into the choanomal skeleton.
Next to the actines of the giant tri- and tetractines, the choanosomal skeleton (Figs 29a–c) contains scattered
intermediate to small-sized regular triactines and tetractines, lining the choanocyte chambers, with free actines of
these spicules protruding into the lumen of the choanocyte chambers. The atrial skeleton (Figs 29a–b) consists of a
single layer of sagittal triactines, and less common tetractines, provided with characteristic abruptly curved paired
actines, the paired actines lined up in a single plane. There is a distinct complement of diactinal, triactine-derived,
spicules which may be the product of broken-off third actines, but the common occurrence indicates it may be a
reduced spicule form. Likewise, there are club-shaped irregular diactinal spicules, probably derived from the
abruptly curved sagittal triactines, scattered in the atrial region; occasional forms of these spicules occur with
reduced or absent unpaired actines.
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FIGURE 28. Leucaltis nodusgordii (Poléjaeff, 1883), a, habitus in situ of RMNH 1772 from Manado (photo N.J. de Voogd), b,
preserved specimen RMNH Por. 1772 (scale bar = 1 cm), c, habitus in situ of ZMA Por. 17557 from Papua New Guinea (photo
R. Sonnenschein).
Spicules. (Figs 30a–e) Giant triactines and tetractines, small regular triactines and tetractines, sagittal ‘abruptly
curved’ triactines and tetractines, reduced forms of the latter.
Giant triactines (Fig. 30a), equiangular, actines more or less equiactinal, 252–458.1–792 x 19–47.3–108 µm.
Giant tetractines (Fig. 30b), equiangular, actines more or less equiactinal, or occasionally with slightly longer
basal actine, 624–820.1–1110 x 66–85.2–97 µm.
Additionally, a single giant diactine, 846 x 36 µm, was observed. Possibly, it represented a reduced triactine.
Regular equiactinal triactines (Fig. 30d), occasionally tripod-shaped, 30–79.1–193 x 1.5–4.7–13 µm.
Regular tetractines (Fig. 30d), actines of the basal radiate system and apical actines more or less similar, 21–
83.9–138 x 2–4.4–7.5 µm.
Sagittal, abruptly angled triactines (Fig. 30e), unpaired actine 42–47.3–60 x 2.5–2.9–3.5 µm, paired actines,
42–62.1–72 x 3–3.8–4.5 µm.
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FIGURE 29. Leucaltis nodusgordii (Poléjaeff, 1883), a, histological cross section of tube wall of RMNH Por. 1772 showing
from left to right cortical skeleton of giant tri- and tetractines, equiangular triactines of the choanocyte chamber region, and
abruptly angled tri- and tetractines of the atrial skeleton (scale bar = 200 µm), b–d, cross sections of holotype BMNH
1884.4.22.23a, b, detail of atrial skeleton and subatrial chamber region (scale bar = 200 µm), c, cross section of chamber region
and cortex (scale bar = 500 µm), d, overview of cross section of tube wall (scale bar = 1 mm).
Sagittal, abruptly angled tetractines (Fig. 30f), unpaired actine 60–64.2–66 x 4–4.2–5 µm, paired actines 78–
79.3–87 x 6.5–6.8–7 µm, apical actine short, difficult to measure, approximately 10 x 3 µm.
Reduced diactinal modifications of sagittal triactines, size of actines 60–64.4–69 x 2.5–3.1–4 µm (not shown).
Ecology. Coral reefs, among living corals, 15–25 m.
Distribution. At least North Australia, New Caledonia, Indonesia, Papua New Guinea, Seychelles, Zanzibar;
possibly South Australia, Japan, Sri Lanka.
Remarks. The Papua New Guinea material (ZMA Por. 17757) included in the above given spicule data, had
somewhat smaller spicules: giant triactines 228–602 x 20–47 µm, giant tetractines 570–998 x 65–104 µm, small
triactines 54–66 x 2–3 µm, small tetractines paired & unpaired actines 54–66 x 1.5–2.5 µm and apical actines 8–12
x 2 µm, atrial triactines & tetractines with paired & unpaired actines 33–74 x 3–6 µm, apical actines 8–12 x 6 µm,
and diactines 50–60 x 4 µm.
We report here also two specimens belonging to this species from outside Indonesia, viz. the Seychelles (ZMA
Por. 12443 and 16248), with essentially similar habitus and spicule size data: giant triactines 111–610 x 18–54 µm,
giant tetractines 552–1230 x 90–156 µm, small triactines 48–126 x 2–7 µm, small tetractines with actines of the
basal triradiate system 66–115 x 2–5 and apical actines 10–35 x 2–5 µm, atrial triactines and tetractines with paired
and unpaired actines 45–60 x 2.5–6 µm and apical actines 10–15 x 6 µm, diactines 67–72 x 3–4 µm.
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FIGURE 30. Leucaltis nodusgordii (Poléjaeff, 1883), SEM images of the spicules of RMNH Por. 1772, a, giant and
intermediate sized triactines of the cortical region, b, giant and intermediate sized tetractines of the cortical region, c, small
regular-shaped triactines of the chamber layer, c1, small triactines seen from the side, d, regular equiangular tetractine of the
chamber layer, e, ‘abruptly angled’ triactines, f, ‘abruptly angled’ tetractines, both from the atrial region.
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FIGURE 31. Leucaltis clathria (Haeckel, 1872), RMNH Por. 9314, from the Guayana shelf, Atlantic Ocean, a, habitus of
preserved specimen (scale bar = 1 cm), b-g, SEM figures of the spicules, b, giant tetractine, c, giant triactines, d, large abruptly
angled triactines, e–e1, small abruptly angled tri- and tetractines, f–f1, regular tri- and tetractines, g, trichoxea.
We name this material Leucaltis nodusgordii, against the consensus in the literature that it should be named
Leucaltis clathria (Haeckel, 1872) (e.g. Dendy 1913; Lévi et al. 1998; Wörheide & Hooper 1999; Borojevic &
Klautau 2000). The treatment by previous authors of this and related specimens is rather frustrating. The
subsequent authors all refer to Poléjaeffs admittedly excellent description, but fail to compare it in detail with
Haeckel’s type of L. clathria. This should have been done, e.g. by Dendy (1913), who saw fragments of Haeckel’s
type brought to the Natural History Museum in London by Mr. R.W.H. Row. Dendy (1913: 17) stated that he
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examined the fragments carefully, and he gives some comments indicating that Haeckel’s description was
incomplete, but failed to provide measurements of the spicules. Unfortunately, we were not able to lay our hands on
a slide of Haeckel’s type still remaining in the collections in London, cited by Burton (1963: 598) and Wörheide &
Hooper (1999: 877). Both these sources, like Dendy (1913), did not describe the contents of the slide. The external
form is apparently so characteristic and convincing that proper description of the spicules and their variation is
largely neglected by most authors.
Haeckel (1872: 159, pl. 29 figs 3a–c) described a brownish (alcohol) clathrate mass of tubes, size 3–6 cm, from
a depth of 63 m off the coast of Florida, collected by A. Agassiz, as Leucaltis (or alternatively Artynas) clathria,
with the spicules described and measured as:
Triactines (‘mittelgross’ = middlesized) 400–600 x 30–50 µm. Tetractines (‘colossal’) 800–1200 x 100–150
µm, occasionally as long as 2000 µm. Sagittal tri- and tetractines (drawn with paired actines in 180° degree angle
and with bluntly rounded ends), paired actines 50–70 x 2–3 µm, unpaired actines 30–40 µm. Triactines more
common than tetractines. Occasional subregular triactines 400–700 x 1–2 µm. Although in his description he
mentions small choanosomal tri- and tetractines supporting the choanocyte chambers he does not provide
measurements. Dendy (1913) stated that these were amply present in the London fragments of the type.
We report here a specimen in the collections of the Naturalis Biodiversity Center, RMNH Por. 9314, from the
Guyana shelf (7.7°N 57.5°W, depth 65 m), which may be considered representative for the Western Atlantic
population to which Haeckel’s type is assumed to belong. The habitus (Fig. 31a) and the spiculation (Figs 31b–g)
(actines of giant tri- and tetractines: 210–530 x 15–63 µm, actines of small equiangular tri- and tetractines: 40–55 x
2–3 µm, abruptly angled tri- and tetractines with unpaired angles 40–45 x 3–4 µm, paired actines 50–65 x 3–4 µm)
of this specimen are indeed strikingly similar to the Indo-West Pacific specimens listed and described above.
However, there is one obvious difference in the spicule complement (see Fig. 31d), namely the presence in the
Guyana specimen of a second category of large ‘abruptly angled’ triactines with paired actines 122–141 x 8.5–10
µm and unpaired actines 111–126 x 6–10 µm, clearly twice as long and thick as the usual spicules reported by us
above. These spicules are not very common, so they might have been overlooked by Haeckel and Dendy. It appears
to us that this difference supports the specific distinctness of specimens from West Pacific and West Atlantic
localities. Klautau et al. (2013) found considerable DNA sequence distance between Leucaltis clathria from the
Caribbean and individuals assigned to L. clathria from Australia, and on that basis pleaded for making a specific
distinction. This confirms that both morphologically and genetically a separate species Leucaltis nodusgordii is
valid.
Family Leucascidae Dendy, 1893a
Genus Ascaltis Haeckel, 1872
Clathrina-like Leucascidae with pseudoatrium and tightly anastomosed asconoid aquiferous system covered by a
thin ‘skin’-like single layer of spicules. The anastomosed tubes form a cortex-like structure, obscuring individual
tubes. Spicules are small triactines and tetractines, and may occasionally include diactines. (after Klautau, et al.
2013).
Ascaltis angusta sp.nov.
Figures 32a–d, 33a–d
Material examined. Holotype ZMA Por. 09097, Indonesia, SE Sulawesi, Salayar, NW coast of Pulau Guang,
6.35°S 120.45°E, depth 2–4 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat.
152/II/Cave 1/06.
Paratypes ZMA Por. 08221a (ten individual cormi), Indonesia, SE Sulawesi, Tukang Besi Islands, southern
reef of Karang Kaledupa, 5.9333°S 123.8°E, depth 4–11 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch
Snellius II Expedition stat. 016/III/10, 8 September 1984.
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FIGURE 32. Ascaltis angusta sp. nov., a, preserved holotype ZMA Por. 09097 (arrow indicates slit-like pseudoatrium) (scale
bar = 1 cm), b, preserved paratypes ZMA Por. 08221 (scale bar = 1 cm), c, overview of skeleton of one of the paratypes (scale
bar = 200 µm), d, detail of surface skeleton (scale bar = 200 µm).
Description. (Figs 32a–b) Tight masses of thin tubes, surrounding a thin slit-like pseudoatrium (arrow in Fig.
32a). Individual specimens often occurring in small patches of a few cm
2
, connected by single tubes to form a
cormus-network. Size variable, the holotype (Fig. 32a) is 2 cm long and maximally 1.2 cm wide. Live color white,
but in alcohol this turns into yellow or beige. Oscules few, at the apices of single water-collecting tubes.
Skeleton. (Figs 32c–d) Tube system on the outside covered with a thin, often damaged membrane consisting
of a single spicule layer (Fig. 32c). Granular cells abound in the surface (Fig. 32d). Underneath, the tubes have thin
walls of usual two layers of spicules (Fig. 32d).
Spicules. (Figs 33a–d) Small triactines and tetractines of variable but similar shape, partially slightly or more
distinctly parasagittal.
Triactines, equiactinal, (Fig. 33a) length of actines 72–101.8–196 x 6–9.1–13 µm; parasagittal triactines (Figs
33c) with unpaired actines 180–198 µm, and shorter paired actines.
Tetractines, equiactinal (Fig. 33b) with actines of basal radiate system 80–103.4–126 x 6–8.6–11 µm, smooth
apical actines 45–105 x 4–6 µm; parasagittal tetractines (Fig. 33d) with unpaired actines 120–186 µm, paired
actines shorter.
Ecology. Shallow water reefs, often in caves.
Distribution. Indonesia.
Etymology. The word angustus (L.) means narrow, tight, referring to the tight anastomosis of the tubes of the
cormus in this species.
Remarks. Because the habitus of our new species with its pseudoatrium and its tight-meshed cormus matches
the definition of the genus Ascaltis, we believe it is a likely member of that genus. The thin membrane enclosing
the whole cormus is not always clearly visible and appears present only in places (preserved material). The new
species shows considerable resemblance to Ascaltis gardineri (Dendy, 1913) (as Leucosolenia), previously
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considered a member of Clathrina (cf. Klautau & Valentine 2003), but subsequently assigned to Ascaltis by
Klautau et al. 2013). However, there are distinct differences in the spicules. Klautau & Valentine (2013: 26) in their
redescription of the type material noticed the occurrence of two distinct size categories with differently shaped
apical actines of the tetractines. No parasagittal spicules occur in A. gardineri.
FIGURE 33. Ascaltis angusta sp. nov., holotype ZMA Por. 09097, SEM images of spicules a, regular equiangular equiactinal
triactine, b, regular equiangular equiactinal tetractines, c, sagittal triactines, d, sagittal tetractine.
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FIGURE 34. Leucascus flavusCavalcanti et al. (2013), a, habitus in situ of RMNH Por. 2279 from Bali (photo N.J. de Voogd),
b, preserved habitus of holotype ZMA Por. 13145 from SW Sulawesi (scale bar = 1 cm), c+d, in situ images of RMNH Por.
9346, from Ternate (photo N.J. de Voogd), e, tangential view of surface skeleton (scale bar = 500 µm), f, cross section (scale
bar = 500 µm).
Genus Leucascus Dendy, 1893a
Leucascus flavus Cavalcanti, Rapp & Klautau, 2013
Figures 34a–f, 35a–d
? Leucosolenia cerebrum; Breitfuss, 1896a: 434; Breitfuss, 1898: 172 (not Haeckel, 1872)
Leucascus flavus Calvalcanti et al., 2013: 297, Figs 14–15.
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FIGURE 35. Leucascus flavus Cavalcanti et al. (2013), SEM images of the spicules, a–c, holotype ZMA Por. 13145, a,
triactine, b, tetractine, c, detail of apical actine of tetractine showing the characteristic spination, d, ditto of RMNH Por. 9347
from Ternate.
Material examined. Holotype ZMA Por. 13145 ((Fig. 34b), Indonesia, SW Sulawesi, Spermonde
Archipelago, Bone Baku, 5.132°S 119.361°E, depth 12 m, SCUBA, coll. N.J. de Voogd #BB/NV/160597/27, 16
May 1997.
Additional specimens. RMNH 2279, Indonesia, Bali, Tulamben area, bay S of ‘Emerald Hotel’, 8.2847°S
115.6031°E, depth 20 m, SCUBA, coll. N.J. de Voogd, #BAL 23/120401/151, 12 April 2001 (Figs 34a, c–d);
RMNH 9346, Indonesia, Ternate, Halmahera, Sulamadaha Beach, 0.8632°N 127.3345°E, depth 15 m, SCUBA,
coll. N.J. de Voogd, #TER.04/2661009/NV017, Ternate-Halmahera Expedition 2009, 26 October 2009; RMNH
9347, Indonesia, Halmahera, Tidore Cobo, 0.7552°N 127.4066°E, depth 15 m, SCUBA, coll.
N.J. de Voogd,
#TER.15/0111009/, Ternate-Halmahera Expedition 2009, 1 November 2009.
Description. The description of the holotype (Fig. 34b) is here summarized from Cavalcanti et al. 2013:
Yellow massive sponge, with a thin skin (Fig. 34e) covering a tight mass of anastomosed tubes (Fig. 34f). Large
oscules, without membrane or spicule fringe. Spicules triactines 70–120 x 7.5–11 µm, tetractines with actines of
the basal triradiate system 75–115 x 7.5–10 µm, apical actines 41–61 x 4–5 µm. The apical actines are rugose-
spined (Fig. 35c) and protrude into the choanocyte tubes.
Description of additional specimens, also from in situ photos: lobate-folded massive sponges (Figs 34a–c) with
a pale-yellow or whitish yellow color in situ. Oscules wide, rounded or elongate, at the top of lobes.
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Spicules. (Figs 35a–d) Triactines and tetractines.
Triactines (Fig. 35a), equiangular equiactinal, occasionally parasagittal, 96–118.1–141 x 7.5–8.5–11 µm.
Tetractines (Fig. 35b), triradiate system equiangular and equiactinal, 90–114.3–127 x 7–9.2–10 µm, apical
actines spined (Figs 35c–d), 45–54.7–68 µm.
Ecology. Shallow-water reefs.
Distribution. So far known only from Indonesia, but widely distributed there (Sulawesi, Bali, Ternate).
Remarks. This species was recently described by Cavalcanti et al. (2013) on the basis of an Indonesian
specimen collected by one of us (NDV). Since then, the species was found several times in localities all over the
Indonesian archipelago. It may be mistaken for other yellowish massive calcineans, such as Ascoleucetta sagittata
Cavalcanti et al., 2013 and Leucetta chagosensis Dendy, 1913 (cf. below), but these differ in color (more definitely
brightly yellow), shape not so lobate but more massive and rounded, presence of large triactines and the lack of
spined apical actines of the tetractines. The latter is not easily seen with light microscopy but it is very obvious
under SEM.
There is a possibility that this species was previously reported by Breitfuss (1896a, 1898) from Ternate as
Leucosolenia cerebrum, a Mediterranean species now assigned to the genus Borojevia. Although his description is
quite concise and generally inadequate, he specifically mentions spined apical actines of the tetractines. It is also
possible that this concerns a so far undescribed species of Borojevia.
Genus Ascoleucetta Dendy & Frederick, 1924
Ascoleucetta sagittata Cavalcanti, Rapp & Klautau, 2013
Figures 36a–f
Ascoleucetta sagittata Cavalcanti et al., 2013: 308, Figs 21–22.
Material examined. Holotype ZMA Por. 13283, Indonesia, SW Sulawesi, Spermonde Archipelago, Kudingareng
Keke, depth 15 m, SCUBA, coll. N.J. de Voogd, #KK/NV/130497/32, 13 April 1997.
Description. The holotype description is here summarized from Cavalcanti et al. 2013: Yellow massive
sponge (Figs 36a–c) presumably consisting of a tight mass of tubes, but this is largely obscured. In stead there are
some folded portions and lacunae in the interior (Fig. 36d), which indicate the tubes. Presumably the aquiferous
system is solenoid, but this is not elaborated. The skeleton contains three categories of triactines, one large regular
(Fig. 36e), 300–550 x 20–55 µm, one small sagittal (Fig. 36g) with unpaired actines 85–144 x 5–10 µm and paired
actines 80–150 x 5–10 µm, one small regular (Fig. 36f), 110–160 x 12–15 µm, and small regular tetractines (Fig.
36h) with actines of the basal radiate system 110–150 x 10–12.5 µm and apical actines (Fig. 36h1) 37–65 x 5–7.5
µm. The latter are protruding into the lumen of the internal tubes.
Ecology. Shallow-water reefs.
Distribution. Indonesia.
Remarks. This species was recently described from a fragment, and we provide here images of the whole
specimen. The presence of sagittal triactines is not observed very frequently, and they are not associated with the
oscules, since there are no proper oscules in the holotype specimen. This species was earlier confused with
specimens assigned to Leucetta chagosensis because of similarity in spiculation with it (see below). By careful
comparison, we have become convinced that Ascoleucetta sagittata and the yellow globular-massive Leucetta-like
specimens, which are so commonly encountered in the shallow reefs of Indonesia and the adjacent regions, are
virtually indistinguishable and may very well be the same species. From molecular sequence results of Voigt et al.
(2012) there are indications that L. chagosensis is not closely related to other Leucetta species like Leucetta
microraphis Haeckel, 1872, which might support the idea that L. chagosensis is an Ascoleucetta, not a genuine
Leucetta, conspecific or very closely related to A. sagittata. This would require careful histological sections. We
refrain from making a definite conclusion on this species because of a forthcoming revision of Klautau et al. on
Leucetta.
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FIGURE 36. Ascoleucetta sagittata Cavalcanti et al. (2013), holotype ZMA Por. 13283, a–c, various images of preserved
holotype, showing surface and atrium (scale in a = 1 cm), d cross section of skeleton, showing layered structure (scale bar = 500
µm), c–f, SEM images of the spicules, e, large triactines of the surface, f, small triactine, e, sagittal triactine, f, tetractine, f1,
detail of tetractine.
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Family Leucettidae De Laubenfels, 1936
Genus Leucetta Haeckel, 1872
Remarks. The present collections contained two distinct species, (1) yellow or bright yellow, globular or with
rounded lobes, with large oscules often provided with a thin low rim, and (2) pink, massive with one or more,
frequently several, tubiform, smooth or grooved, elevations. Both species have a predominance of triactine spicules
in two distinct size categories, and rare small tetractine spicules. The spiculation of the two differs mostly in the
length and thickness of the large (’giant’) triactines, with the yellow lobes having the dimensions approximately
300–700 x 20–40 µm, and the pink tubes having these giant triactines with actine size 1000–1800 x 80–250 µm.
It is customary in the scientific literature to use the name Leucetta chagosensis Dendy, 1913 for the yellow
species (see e.g. Wörheide & Hooper 1999), but the pink species is only recently again named Leucetta
microraphis Haeckel, 1872, originally as L. primigenia var. microrhaphis), after periods in which the name L.
primigenia Haeckel, 1872 was used. These assignments are debatable and await proper revision of type material,
but for the time being we keep here the customary names. The status of L. chagosensis is also discussed above
under Ascoleucetta sagittata.
Leucetta chagosensis Dendy, 1913
Figures 37a–f, 38a–d
Leucetta chagosensis Dendy, 1913: 10, Pl. 1 fig. 6, Pl. 4 fig. 2; Gosliner et al., 1996: 16, photos 3 and 5; Lévi et al., 1998: 77;
Wörheide & Hooper, 1999: 882, figs 8H–M (with additional synonyms); Borojevic & Klautau, 2000: 194, fig. 6;
Wörheide et al., 2002: 1753; Baine & Harasti, 2007: 15.
Leucetta ‘lemon’ Colin & Arneson, 1995: 60, photo 230.
Material examined. ZMA Por. 08048, Indonesia, Nusa Tenggara, N of Sumbawa, Bay of Sanggar, 8.32°S
118.24°E, depth 2–4 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 114/II/05,
22 September 1984; ZMA Por. 08513, Indonesia, Nusa Tenggara, N of Sumbawa, Bay of Sanggar, 8.32°S
118.24°E, depth 5–7 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 114/III/16,
21 September 1984; ZMA Por. 08694, Indonesia, Maluku, Pulau Pulau Maisel, reef edge N of Mai, 5.4667°S
125.5167°E, depth 6–13 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 013/
IV/22, 4 September 1984; ZMA Por. 08923, Indonesia, SE Sulawesi, NE Take Bone Rate, S of Tarupa Kecil, 6.5°S
121.1333°E, depth 10–15 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 133/
IV/42, 25 September 1984; ZMA Por. 10846, Indonesia, Nusa Tenggara, Bali, Pulau Menjangan, 8.1167°S
114.5167°E, depth 4 m, coll. C.C. Hofman, #11, 30 July 1991; ZMA Por. 15947, Indonesia, Sumatera, Padang,
Pandan Island, 0.3833°S 100.2167°E, depth 10 m, coll. M. Pampus, 7 November 1995; ZMA Por. 16012,
Indonesia, Sulawesi, Spermonde Archipelago, Bone Baku, 5.132°S 119.361°E, depth 6 m, SCUBA, coll. B.W.
Hoeksema, 14 May 1997; ZMA Por. 17167, Indonesia, SW Sulawesi, Spermonde Archipelago, Kapoposang
Island, 4.6885°S 118.9367°E, depth 12.3 m, coll. R.A. Edrada, #TM84, 7 August 1997; ZMA Por. 17364,
Indonesia, SW Sulawesi, Spermonde Archipelago, Barong Lompo, depth 14.4 m, coll. R.A. Edrada, #TM14, 8
August 1997; ZMA Por. 17555, Papua New Guinea, 10.9561°S 150.4188°E, depth 12–18 m, SCUBA, coll. Rachel
Sonnenschein, #02126, 28 May 2002; ZMA Por. 17709, Papua New Guinea, Milne Bay, 9.1695°S 150.3012°E,
depth 9 m, SCUBA, coll. Karen Tenney, #03554, 17 December 2003; ZMA Por. 19879, Indonesia, North Sulawesi,
Siladen Island, 1.63°N 124.8°E, depth 10 m, coll. S. Tsukamoto, June 2006; RMNH Por. 1627, Palau, Koror, Siaes
Reef, western barrier, 7.3116°N 134.2268°E, depth 25 m, SCUBA, coll. N.J. de Voogd, #KOR07/170505/045, 17
May 2005; RMNH Por. 1628, Palau, Koror, Mutremdiu, W side Uchelbeluu Reef, 7.2737°N 134.5241°E, depth 20
m, SCUBA, coll. N.J. de Voogd, #KOR.03/210505/087, 21 May 2005; RMNH Por. 1652, Palau, Koror, Toagel
Mlungi Channel, western barrier reef off Babeldaob, 7.5425°N 134.4685°E, depth 27 m, SCUBA, coll. N.J. de
Voogd, #KOR09/180505/53B, 18 May 2005; RMNH 1792, Indonesia, NE Kalimantan, Berau Islands, NE-side
Maratua Island, 2.2802°N 118.6029°E, depth 0–30 m, SCUBA, coll. C. Fransen, #BER17/101003/092, East
Kalimantan Expedition 2003, 10 October 2003; RMNH Por. 1880, Indonesia, NE Kalimantan Islands, off Tanjung
Batu, 2.2396°N 118.0934°E, depth 0–30 m, SCUBA, coll. B.W. Hoeksema, #BER19/131002/151, East Kalimantan
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FIGURE 37. Leucetta chagosensis Dendy (1913), a, habitus in situ of ZMA Por. 16012 from SW Sulawesi (photo B.W.
Hoeksema), b, habitus in situ of ZMA Por. 17709 from Papua New Guinea (photo K. Tenney), c, habitus ‘on deck’ of samples
ZMA Por. 17167 from SW Sulawesi, d, tangential view of surface skeleton (scale bar = 500 µm), e, cross section of peripheral
region (scale bar = 500 µm), f, oscular rim of ZMA Por. 17167 showing concentration of sagittal triactines (arrows) (scale bar
= 200 µm).
Expedition 2003, 13 October 2003; RMNH Por. 1923, Indonesia, Nusa Tenggara, Bali, SE-end of Tulamben beach,
8.2778°S 115.5959°E, depth 18 m, SCUBA, coll. N.J. de Voogd, #BAL22/NV/130401/183, Bali Lombok Strait
Expedition 2001, 13 April 2001; RMNH Por. 2910, Indonesia, Nusa Tenggara, Bali, NE side of Pulau Serangan,
off lighthouse, 8.7214°S 115.2586°E, depth 0–17 m, SCUBA, coll. N.J. de Voogd, #BAL14/050401/073, Bali
Lombok Strait Expedition 2001, 5 April 2001; RMNH Por. 9196 Indonesia, Halmahera, Maitara Maitara W,
0.7299°N 127.3624°E, depth 0–30 m, SCUBA, coll. N.J. de Voogd, #TER09/291009/, Ternate-Halmahera
Expedition 2009, 29 October 2009; RMNH Por. 9197, Indonesia, Halmahera, off Hiri Pulau Maka, 0.9119°N
127.3091°E, depth 0–30 m, SCUBA, coll. N.J. de Voogd, #TER13/311009/, Ternate-Halmahera Expedition 2009,
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31 October 2009; RMNH Por. 9348, Indonesia, off Halmahera mainland, Teluk Dodinga, Karang Galiasa Kecil W,
0.8525°N 127.5888°E, depth 0–30 m, SCUBA, coll.
N.J. de Voogd, #TER39/141109/, Ternate-Halmahera
Expedition 2009, 14 November 2009.
FIGURE 38. Leucetta chagosensis Dendy (1913), SEM images of the spicules of ZMA Por. 08694 from the Maisel
Archipelago (Banda Sea), a, large triactine from the surface skeleton, b, small triactine, c, tetractine, d, sagittal triactines.
Description. (Figs 37a–c) Yellow globular, semiglobular or massively lobate sponge with one or more
prominent oscules with slightly raised rim. There is usually an ‘atrium’ or large internal cavity, while the
choanosome is provided with irregularly elongate radiate canals. Size up to 12 x 8 x 8 cm. Surface optically
smooth, but harsh to the touch. Consistency toughly compressible.
Histology. Leuconoid aquiferous system.
Skeleton. Ectosomal skeleton (Fig. 37d) a thin layer of small triactines forming small rounded (‘alveolar’)
meshes, carried by subectosomal large triactines. Choanosomal skeleton lacunose (Fig. 37e), in places similar to
ectosomal arrangement of small alveolae, with large lacunae. These lacunae are bordered by rare tetractines,
sticking into the lumen with their apical actines. Oscular rims have a layer of sagittal triactines (Fig. 37f arrows).
Spicules. (Figs 38a–d) Large triactines, small triactines, tetractines.
Large triactines (Fig. 38a), equiangular equiactinal, 274–480.1–720 x 20–32.3–60 µm.
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Small triactines (Fig. 38b), equiangular equiactinal, 102–138.9–186 x 8–10.9–17 µm, rarely some triactines
have crooked ‘paired’ or single actines and these may then appear sagittal (Fig. 38d); these are particularly
concentrated in the rims of the oscules (see arrows in Fig. 37f) and are of the same general size as the regular small
triactines.
Small tetractines (Fig. 38c), rare, with equiangular equiactinal basal radiate system, actines 104–126.2–135 x
9–11.5–13 µm, with small apical actines, 12–15 x 6–8 µm.
Ecology. Shallow-water reefs.
Distribution. Indonesia, Papua New Guinea, Palau Islands. Elsewhere reported from a.o. Northeast Australia,
New Caledonia, Vanuatu, French Polynesia, the Western Indian Ocean, and the Red Sea (Wörheide et al. 2008).
Remarks. By the bright yellow color our specimens differ clearly from the pink or whitish Leucetta
microraphis (cf. below), and also by the much larger size of the triactines of that species.
L. chagosensis may be confused with small light-colored Pericharax orientalis sp.nov., but the surface of that
species is supported by a layer of special cortical sagittal triactines, lacking in the present species. The oscular
sagittal triactines of L. chagosensis bear a resemblance to these cortical triactines of P. heteroraphis.
Furthermore, there is considerable resemblance to Ascoleucetta sagittata (cf. above), a yellow massive-
globular species with a central atrium and strongly folded choanosome. If this species is genuinely different from
L. chagosensis then they can be kept apart by the lack of oscules and oscular rims with concentrations of sagittal
triactines in A. sagittata. We will await further research before it can be decided that the two must be merged.
Over its large distribution, this species was demonstrated to have considerable genetic structure with
recognizable discrete geographic populations (see Wörheide et al., 2002).
Leucetta microraphis Haeckel, 1872
Figures 39a–d, 40a–d
Leucetta primigenia var. microraphis Haeckel, 1872: 119, pl. 21 Figs 10–17.
Leucetta primigenia; Lendenfeld, 1885: 1117; De Laubenfels, 1954: 253, fig. 179; Borojevic, 1967b: 3, fig. 2; Borojevic &
Klautau, 2000: 193, fig. 5; Colin & Arneson, 1995: 60, photo 229; Gosliner et al., 1996: 16, photo 4; Erhardt & Baensch,
1998: 22.
?Leucetta solida; Breitfuss, 1896: 434; 1898: 169 (not: Schmidt, 1862).
Material examined. ZMA Por. 00152, Indonesia, Nusa Tenggara, E coast of Sumbawa, Sapeh Bay, 8.5839°S
119.0324°E, depth 0–38 m, dredge and reef exploration, Siboga Expedition stat. 311, 12 February 1900; ZMA Por.
08108, Indonesia, Nusa Tenggara, N of Sumbawa, Bay of Sanggar, 8.3417°S 118.262°E, depth 1–4 m, coastal reef
with sea grass, snorkeling, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 120/II/06, 21
September 1984; ZMA Por. 08510, Indonesia, Nusa Tenggara, N of Sumbawa, Bay of Sanggar, 8.32°S, 118.24°E,
depth 5–7 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition 1984 stat. 114/III/13, 21
September 1984; ZMA Por. 08649, Indonesia, SE Sulawesi, SW Salayar, NW coast of Pulau Guang, 6.35°S,
120.45°E, depth 4–12 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition 1984 stat. 152/
III/32, 28 September 1984; ZMA Por. 09051, Indonesia, Nusa Tenggara, NE coast of Sumba, 9.95°S 120.8°E,
depth 50 m, dredge, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition 1984 stat. 68/V/17, 16
September 1984; ZMA Por. 14536, Indonesia, North Sulawesi, Bunaken, SW Nain Island, 1.7681°N 124.7671°E,
depth 18 m, SCUBA, coll. B.W. Hoeksema, #SYMBIOSPONGE 98/NS/MAY08/BH090, 8 May 1998; ZMA Por.
15917, Indonesia, Sumatera, W, Padang, Pandan Island, W, 0.3833°S 100.2167°E, depth 5 m, SCUBA, coll. M.
Pampus, #PA B13, 7 November 1995; ZMA Por. 15936, Indonesia, Sumatera, Padang, Pisang Island, 0.95°S
100.1667°E, depth 5 m, SCUBA, coll. M. Pampus, # PP1, 1 December 1995; ZMA Por. 16011, Indonesia, SW
Sulawesi, Spermonde Archipelago, Samalona W, 5.125°S 119.342°E, depth 5–10 m, SCUBA, coll. B.W.
Hoeksema, #2, 8 May 1997; ZMA Por. 16682, Indonesia, SW Sulawesi, Spermonde Archipelago, Lae-lae,
5.1336°S 119.3381°E, depth 5–10 m, SCUBA, coll. N.J. de Voogd, #LL/NV/240597/D, 24 May 1997; ZMA Por.
17740, Indonesia, Kalimantan, Berau Islands, SE-side Kakaban Island, 2.1261°N 118.5517°E, depth 0–30 m,
SCUBA, coll. R.G. Moolenbeek, #BER03/36; ZMA Por. 20126, Indonesia, North Sulawesi, S of Siladen Island,
1.63°N 124.8°E, depth 10 m, coll. S. Tsukamoto, #06M039, 6 June 2006; RMNH Por. 1777, Indonesia, NE
Kalimantan, Berau Islands, NE-side Maratua Island, 2.2802°N 118.6029°E, depth 5 m, SCUBA, coll. K. van
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Egmond, #BER17/101003/093, East Kalimantan Expedition 2003, 10 October 2003; RMNH Por.1809, Indonesia,
Nusa Tenggara, Bali, Tulamben area, bay S of ‘Emerald’ Hotel, 8.2847°S 115.6031°E, depth 30 m, SCUBA, coll.
N.J. de Voogd, #BAL23/NV/140401/205, Bali Lombok Strait Expedition 2001, 14 April 2001; RMNH Por. 1883,
Indonesia, Sulawesi, Spermonde Archipelago, Bone Lola, 5.0°S 119.35°E, depth 6 m, SCUBA, coll. N.J. de
Voogd, 29 October 2002; RMNH Por. 2129, Indonesia, Java Sea, off Jakarta, Kepulauan Seribu, Payung Besar
Island, E side, 5.8219°S 106.5631°E, depth 10–15 m, SCUBA, coll. N.J. de Voogd, #SER43/260905/167,
Kepulauan Seribu Expedition 2005, 26 September 2005; RMNH Por. 6610, Indonesia, North Sulawesi, Lembeh
Strait, SW Sarena Kecil, 1.4555°N 125.2236°E, depth 25 m, SCUBA, coll. K. van Egmond, #LEM15/070212, 7
February 2012.
FIGURE 39. Leucetta microraphis Haeckel (1872), a, habitus ‘on deck’ of ZMA Por. 20126 from N Sulawesi (scale bar = 1
cm) b, habitus in situ of ZMA Por. 16011 from SW Sulawesi (photo B.W. Hoeksema), c, habitus in situ (not collected), photo
B.W. Hoeksema, d, overview of spicules of ZMA Por. 08649 from Salayar (scale bar = 1 mm).
Description. (Figs 39a–c) Dark pink, brownish pink or pinkish-white sponges with variable shape, lobate,
flabellate or massive, with raised or terminal oscules of 0.5–2 cm diameter, atrial lumen up to 2 cm in diameter.
Size up to 11 x 10 x 6 cm.
Histology. Leuconoid aquiferous system.
Skeleton. Essentially ‘alveolar’, small rounded meshes encircling the choanocyte chambers. Just below the
surface there are giant triactines (Fig. 39d).
Spicules. (Figs 40a–d) Giant triactines, small triactines, tripods, small tetractines.
Giant triactines (Fig. 40a), equiangular equiactinal, with conical actines, 1175–1536.3–1980 x 132–169.3–216
µm.
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FIGURE 40. Leucetta microraphis Haeckel (1872), SEM images of the spicules of ZMA Por. 16682 from SW Sulawesi, a,
giant triactine, b, intermediate-sized triactine, c, small triactine, d, tetractine.
Intermediate (Fig. 40b) and small (Fig. 40c) triactines, equiangular equiactinal, 99–175.7–270 x 8–15.8–33
µm; rare tripods with undulating paired actines 150–200 µm
Small tetractines (Fig. 40d), rare, actines of basal radiate system 135–150 x 9–12 µm, apical actines 50–70 µm.
Ecology. Shallow-water reefs down to 30 m.
Distribution. Indonesia, Papua New Guinea, New Caledonia, tropical Australia.
Remarks. This species is in urgent need of revision. Haeckel’s treatment of the name L. microraphis makes the
species unrecognizable and its type locality unknown. No original specimens have ever been redescribed. The use
of the name is entirely based on ‘tradition’. Pinkish brown specimens identified by us under this name can be
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confused with smaller brownish specimens of Pericharax orientalis sp.nov., but the sharp ridges and yellow
interior color of that species do not occur in L. microraphis.
Breitfuss’ (1896, 1898) record of the Mediterranean Leucetta solida (Schmidt, 1862 as Grantia) from Ternate
quite possibly concerns the present species as large and small triactines were mentioned. No detailed spicule size
data were provided by Breitfuss.
Genus Pericharax Poléjaeff, 1883
Remark. The commonly reported tropical shallow-water Pericharax heteroraphis of authors is here given a new
name for reasons presented below.
Pericharax orientalis sp. nov.
Figures 41a–e, 42a–e
Pericharax heteroraphis; Dendy, 1913: 13; Colin & Arneson, 1995: 60, photo 232, (?also photo 231); Gosliner et al, 1996: 16,
photo 6; Allen & Steene, 1996: 30–31; Wörheide & Hooper, 1999: 886, figs 9G–M; Borojevic & Klautau, 2000: 195;
Baine & Harasti, 2007: 15 (not: Pericharax carteri var. heteroraphis Poléjaeff, 1883: 66, pl. II fig. 5, pl. VII fig.8 = P.
carteri)
Material examined. Holotype RMNH Por. 5259, Indonesia, Halmahera, restaurant Floridas, 0.7599°N
127.3571°E, depth 12 m, SCUBA, coll. N.J. de Voogd, #TER03/251009/010, Ternate Halmahera Expedition 2009,
25 October 2009.
Paratypes ZMA Por. 08220, Indonesia, SE Sulawesi, Tukang Besi Islands, southern reef of Karang Kaledupa,
east entrance, 5.9333° S 123.8° E, depth 4–11 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II
Expedition stat. 016/III/08, 6 September 1984; ZMA Por. 17280, Indonesia, Sumatera, Painan coast, Aua Island, 60
km from Padang, 1.3667°S 100.5667°E, depth 5–10 m, SCUBA, coll. R.A. Edrada, #DH77, 13 July 2002; RMNH
Por. 8551, Indonesia, Nusa Tenggara, Bali, Tulamben area, bay S of Emerald Hotel, 8.2847°S 115.6031°E, depth
20 m, SCUBA, coll. N.J. de Voogd, #BAL23/NV/150401/207, Bali Lombok Strait Expedition 2001, 15 April 2001.
Additional specimens. ZMA Por. 00149, Indonesia, Irian Jaya, 1.7083° S 130.7916° E, depth 32 m, trawled,
coll. Siboga Expedition stat. 164, 20 August 1899; ZMA Por. 00150, Indonesia, Irian Jaya, 1.7083° S 130.7916° E,
depth 32 m, trawled, coll. Siboga Expedition stat. 164, 20 August 1899; ZMA Por. 00151, Indonesia, Irian Jaya,
1.7083° S 130.7916° E, depth 32 m, trawled, coll. Siboga Expedition stat. 164, 20 August 1899; ZMA Por. 00181a,
Indonesia, S Sulawesi, anchorage off S point Kabaena Island, 5.5165° S 121.9502° E, depth 22 m, trawled, coll.
Siboga Expedition stat. 209, 23 September 1899; ZMA Por. 00181b, Philippines, Sulu Islands, anchorage off N
Ubian, 6.126° N 120.4333° E, depth 16–23 m, trawled, coll. Siboga Expedition stat. 099, 28 June 1899; ZMA Por.
06526, Indonesia, Nusa Tenggara, N of Sumbawa, Bay of Sanggar, 8.3417°S 118.2617°E, depth 0–1 m, snorkeling,
coll. J. Brouns, Indonesian-Dutch Snellius II Expedition 1984 stat. 120/A2, 21 September 1984; ZMA Por. 08561,
Indonesia, Sulawesi, SE, Take Bone Rate, NE, western edge of reef Taka Garlarang, 6.45°S 121.2083°E, depth 6–8
m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition 1984 stat. 147/III/08, 27 September
1984; ZMA Por. 08627, Indonesia, Sulawesi, SW, Salayar, SW, Pulau Guang, NW coast, 6.35°S 120.45°E, depth
4–12 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition 1984 stat. 152/III/09, 28
September 1984; ZMA Por. 08628, Indonesia, SE Sulawesi, Salayar SW, Pulau Guang, NW coast, 6.35°S
120.45°E, depth 4–12 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 152/III/
10, 28 September 1984; ZMA Por. 08786, Palau Islands, Argulpelu Reef, depth 15 m, SCUBA, coll. M.K. Harper,
#DJF 81–133, 1981; ZMA Por. 16010, Indonesia, SW Sulawesi, Spermonde Archipelago, Kudinggareng Keke,
5.102°S 119.286°E, depth 10–15 m, SCUBA, coll. B.W. Hoeksema, #4, 24 April 1997; ZMA Por. 17168,
Indonesia, SW Sulawesi, Spermonde Archipelago, Kapoposang, 4.6885°S 118.9367°E, depth 13.2 m, SCUBA,
coll. R.A. Edrada, #TM87, 8 August 1997; ZMA Por. 17364, Indonesia, Sulawesi, Spermonde Archipelago,
Barang Lompo, 5.05°S 119.3°E, depth 14.4 m, SCUBA, coll. R.A. Edrada, 25 July 1997; ZMA Por. 17412,
Indonesia, SW Sulawesi, Spermonde Archipelago, Bone Lola, 5.0°S 119.35°E, depth 6 m, SCUBA, coll. N.J. de
Voogd, #NV/BL/291002/349, 29 October 2002; ZMA Por. 17558, Papua New Guinea, Solomon Sea, 10.22°S
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150.87°E, depth 18–24 m, SCUBA, coll. R. Sonnenschein, #02160, 2 June 2002; ZMA Por. 17708, Papua New
Guinea, Milne Bay, 9.62°S 150.9556°E, depth 6–21 m, SCUBA, coll. R. Sonnenschein, #03513, 12 December
2003; ZMA Por. 17740, Indonesia, Kalimantan, Berau Islands, SE side Kakaban Island, 2.1261°N 118.5517°E,
depth 25–30 m, SCUBA, coll. R.G. Moolenbeek, #BER03/36, 2003; RMNH Por. 1633, Palau, Koror, W of Ulong
Island, Ngerumekaul Channel, 7.2819°N 134.2454°E, depth 27 m, SCUBA, coll. N.J. de Voogd, #KOR08/170505/
053A, 17 May 2005; RMNH Por. 1935, Indonesia, Manddin, between Bunaken and Manado Tua, depth 17 m,
SCUBA, coll. N.J. de Voogd, #MD4/180502/093, 18 May 2002; RMNH Por. 9351, Indonesia, Sulawesi, Kabaena
Island, anchorage off S point, 5.5165°S 121.9502°E, depth 22 m, dredge, hard bottom, coll. Siboga Expedition stat.
209, 23 September 1899; RMNH Por. 9352 Philippines, Sulu Islands, anchorage off North Ubian, 6.125°N
120.4333°E, depth 16–23 m, dredge, hard bottom, coll. Siboga Expedition stat. 099, 30 June 1899.
Description. (Figs 41a–c) Upright thick-walled tubular individuals with central vent, occasionally several
individuals are grouped together (Fig. 41a). In mature condition the sides of the tubes are sharply ridged (Fig. 41a,
c), and colors are a mottled reddish brown, green-brown, grey-green and greyish yellow (Fig. 41a). In young
specimens, which are usually lighter colored, the walls often lack ridges and at most show rounded folds (Fig. 41b).
Color of the atrial wall (inside the vent) is yellow or limegreen. Size in mature condition may be considerable. The
preserved holotype is 11 cm high and 15 cm wide (the preserved part is one half of the in situ specimen), with vents
approximately 4 x 1.5 cm; one of the paratypes is even 15 cm high. Overall, specimens may reach 15–30 cm in
height, and 8–20 cm in widest expansion. Surface smooth, but harsh to the touch, prickly because of large
triactines. Consistency firm to hard, usually somewhat compressible.
Histology. Leuconoid aquiferous system.
Skeleton. At the surface (Fig. 41e) there is a cortex of small, predominantly slightly sagittal triactines,
overlying a layer of subdermal rounded lacunae supported by subcortical giant triactines (Fig. 41d). The
choanosomal skeleton is basically alveolar but this is often obscured (Fig. 41d), consisting of small regular
triactines surrounding the choanocyte chambers. The atrial wall (Fig. 41f) shows a predominance of small
tetractines, with their apical actines protruding in the atrium.
Spicules. (Figs 42a–e) Giant triactines, small regular triactines, small sagittal triactines, small tetractines.
Giant triactines (Fig. 42a), equiangular equiactinal, tapering to conical ends, 360–834.2–1560 x 25–67.9–132 µm.
Small regular triactines (Fig. 42c), 60–159.3–228 x 7–12.2–18 µm.
Ectosomal sagittal triactines (Fig. 42b), unpaired actines 61–95.1–132 x 6–8.6–13 µm, paired actines 60–83.2
138 x 5–8.112 µm; rarely the sagittal triactines develop a fourth actine (Fig. 42d).
Atrial small regular tetractines (Fig. 42e), actines of basal triradiate system 126–177.8–228 x 7–9.914 µm,
apical actines curved, 57–87.4–111 x 6–7.4–9 µm.
Ecology. Coral reefs, 4–36 m.
Distribution. Indonesia, Papua New Guinea, NE Australia, New Caledonia, Philippines, Seychelles, Southern
Japan (not: Tristan da Cunha).
Remarks. This is a very common species, as is evident from the frequency with which it appears in
underwater color guides. In many ways it can be called iconic: it is almost certainly one of the largest calcareous
sponges, and has a distinctive mottled brown-yellow appearance and hard consistency, quite the opposite of
mainstream calcareous sponges which tend to be small, whitish and soft. Despite its common occurrence its usually
employed name, Pericharax heteroraphis turned out to be problematic and we were forced to replace it for reasons
given below.
Our extensive material generally fits the description of a specimen from the Chagos Archipelago, Western
Indian Ocean, named as Pericharax heteroraphis Poléjaeff, 1883 by Dendy (1913). Spicule sizes and shapes given
by him match closely with ours. Dendy compared the Tristan da Cunha deep-water type material of Poléjaeffs
Pericharax carteri var. homoraphis and P. carteri var. heteroraphis with his specimen. Unfortunately, he did not
emphasize several observed differences with his own specimen (e.g. size of the giant triactines and shape of the
apical actines of the tetractines) and failed to erect a new species for his shallow-water tropical material. It is highly
unlikely that specimens from shallow-water coral reefs (4–36 m) as treated here and by Dendy (Salomon Reef, 25
m) are conspecific with the type material from 108 m deep in the cold South Atlantic waters off Tristan da Cunha.
We derive this hypothesis from research into the allegedly cosmopolitan distribution of two sponge species in
Calcarea (see
Solé-Cava et al. 1991), which demonstrated that such species consist of several genetically different
cryptic species with limited geographic distributions.
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FIGURE 41. Pericharax orientalis sp. nov., a, habitus in situ of holotype RMNH Por. 5259 from Ternate (photo N.J. de
Voogd), b, habitus in situ of RMNH 1633 from Palau (photo N.J. de Voogd), c, habitus of paratype ZMA Por. 08220 ‘on deck’
(scale bar = 1 cm), d, cross section of wall just below the oscules of holotype ZMA Por. 5259 showing cortical skeleton (left)
choanosomal skeleton with giant, inytermediate and small regular triactines, and atrial skeleton (scale bar = 200 µm), e,
tangential view of surface skeleton of holotypeshowing cortical skeleton of sagittal triactines(scale bar = 200 µm), e, detail of
atrial skeleton of holotype with curved apical actines of tetractines protruding in the atrial lumen (scale bar = 100 µm).
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FIGURE 42. Pericharax orientalis sp. nov., SEM figures of paratype ZMA Por. 08220, a, giant triactines, b, small sagittal
triactine, c, small regular triactine, d, rare sagittal tetractine, e, regular tetractine.
A further problem is that Dendy (1913) as a first reviser failed to choose one of the two varieties described by
Poléjaeff as the typical variety, which - as the nominotypical taxon in the sense of the Code - has to be called
Pericharax carteri carteri, leaving one of the two other names as name for the second variety. According to Dendy
(1913) and Dendy & Row (1913), the variety homoraphis is not a proper Pericharax, but a Leucetta. If we wish to
save the genus name Pericharax from becoming a synonym of Leucetta, it is imperative that the variety
heteroraphis is chosen as the typical variety, because from Poléjaeffs text and his Plate II fig. 5, and from Dendy’s
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(1913) comparisons, it is clear that it is the variety that has the characters of the genus Pericharax (e.g. the cortical
skeleton). This means without any doubt (see ICZN articles 45.6.4 and 47) that the name heteroraphis has to be
replaced by carteri. Hence, from now onwards the Tristan da Cunha Pericharax material is to be called Pericharax
carteri Poléjaeff, 1883 with P. heteroraphis as an objective junior synonym. The species to which Dendy’s and our
specimens belong obviously needs a different name.
A candidate for it is Pericharax peziza Dendy, 1913 from 23 m depth off Cargados Carajos. Its skeleton is
generally similar to our specimens and also the spicules do not differ significantly. However, the shape is a wide
shallow cup, unlike our specimens and unlike all the specimens depicted in many color guides. Also, Dendy
himself considered it a different species from his P. ‘heteroraphis’.
Pericharax canaliculata Burton & Rao, 1932 from the Birmese Mergui Archipelago differs strongly in shape
and spicule sizes, and might not even be a proper Pericharax.
For these reasons, we decided to propose the new name Pericharax orientalis sp.nov.
Order Murrayonida Vacelet, 1981
Family Lelapiellidae Vacelet, 1977
Genus LelapiellaVacelet, 1977
Lelapiella sphaerulifera Vacelet, 1977
Figures 43a–g
Lelapiella incrustans subsp. sphaerulifera Vacelet, 1977: 362, fig. 7.
Material examined. ZMA Por. 09087, Indonesia, SE Sulawesi, NE Take Bone Rate, western edge of reef Taka
Garlarang, 6.45°S 121.2083°E, depth 35 m, SCUBA, coll. H.A. ten Hove, Indonesian-Dutch Snellius II Expedition
stat. 114/V/05, 27 September 1984.
Description. White encrusting sponge (Fig. 43a) on dead coral, size 1.5 x 0.5 x 0.3 cm. Surface irregular, with
thin grooves and small pits; harsh to the touch.
Skeleton. (Fig. 43b) The cortex consists of a dense mass of spherules and tripods, carried over a large
subcortical space by tracts of diactines. The basal skeleton is a mass of ‘hockey-stick’ diactine spicules and
spherules.
Spicules. (Figs 43c–g) Tripods, tetractines, diactines, ‘hockey-stick’ diactines, spherules.
Tripods (Fig. 43c), usually inequiactinal, 171–226.2–285 x 36–46.0–57 µm.
Tetractines (Fig. 43d), rare, actines of the basal radiate system 78–100 x 9, short apical actine, 10–20 µm.
Diactines (Fig. 43g) slightly crooked in the middle, tapering to bluntly rounded endings, 126–193.2–297 x 13–
17.2–27 µm
‘Hockey-stick’ diactines (Fig. 43f), club-shaped, 84–95.8–106 x 18–19.1–21 µm (shaft), thickened part 30–33
µm in diameter.
Spherules (Fig. 43e), bean-shaped, often asymmetrical, 26–30.2–36 x 17–18.2–20 µm.
Ecology. Deeper water under overhangs, in reefs.
Distribution. Indonesia, New Caledonia.
Remark. The present material matches the type in great detail.
Subclass Calcaronea Bidder, 1898
Order Leucosolenida Hartman, 1858
Family Sycettidae Dendy, 1893a
Genus Sycetta Haeckel, 1872
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FIGURE 43. Lelapiella sphaerulifera Vacelet (1977), ZMA Por. 09087, a, preserved fragments (scale bar = 1 cm), b, cross
section of cortex and subcortical space showing cortical tripods, spherules and tracts of diactines (scale bar = 200 µm), c–g,
SEM images of the spicules, c, tripod, d, tetractine, e, spherule, f, hockeystick spicule, g, diactine.
Sycetta vinitincta sp. nov.
Figures 44a–e, 45a–e, 46a–h
Material examined. Holotype RMNH Por. 1873, Indonsesia, NE Kalimantan, Berau Islands, Samama Island,
2.1254°N 118.336°E, depth 2 m, SCUBA, coll. B.W. Hoeksema, #BER10/071003/053, Berau 2003 Expedition, 7
October 2003.
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FIGURE 44. Sycetta vinitincta sp. nov., a, habitus in situ of the holotype RMNH Por. 1873 from the Berau region (photo B.W.
Hoeksema), b, detail of holotype to show papillar surface, c, a second specimen from the same locality (not collected), d,
preserved paratype ZMA Por. 08633 from Salayar (scale bar = 1 cm), e, overview of spicules (scale bar = 200 µm).
Paratype ZMA Por. 08633, Indonesia, SE Sulawesi, Salayar, SW coast, off the NW coast of Pulau Guang,
6.35°S 120.45°E, depth 4–12 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat.
152/III/15, 29 September 1984.
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FIGURE 45. Sycetta vinitincta sp. nov., holotype RMNH Por. 1873, light microscopic images, a, overview of peripheral
protrusion showing the papillar surface (scale bar = 500 µm), b, cross section of the tubar wall to show atrial and papillar
skeleton (scale bar = 100 µm), c, detail of a single papilla showing ‘inarticulated’ tubar skeleton, d, tangential view of outer
surface skeleton (scale bar = 200 µm), e, tangential view of atrial surface (scale bar = 200 µm).
Additional material. ZMA Por. 08221b, Indonesia, SE Sulawesi, Tukang Besi Islands, southern reef of Karang
Kaledupa, 5.9333°S 123.8°E, depth 4–11 m, SCUBA, coll. R.W.M. van Soest, Indonesian-Dutch Snellius II
Expedition stat. 016/III/10, 8 September 1984.
Description. Wine-red irregular broadly sac-shaped thin-walled tubes (Figs 44a–c), with irregular-shaped
knob-like side-tubes (diverticula ?). In alcohol the specimens become white (Fig. 44d). Surface of tube and side
projections is covered in tiny rounded papillae of up to 0.5 mm in height and width (Fig. 44b). Size of entire
specimens up to 4 cm high, individual side-tubes, up to 3 cm long and 8 mm in diameter, overall the sponge may be
4–5 cm wide, atrial lumen up to approximately 2 cm in diameter. Consistency soft and easily damaged.
Histology. No histological section was made, but thin hand sections of the skeleton indicate that it has a
probable syconoid aquiferous system (Fig. 45a–c), but with choanocyte chambers shallow and closely adjacent to
each other in the papillae. See also the Remarks below.
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FIGURE 46. Sycetta vinitincta sp. nov.,SEM images of the spicules of holotype RMNH Por. 1873, a, large slightly sagittal
triactines of the atrial skeleton, b, large slightly sagittal tetractines of the atrial skeleton, c, oxhorn triactine spicules of the
papillar surface, d, oxhorn tetractine spicules of the papillar surface, e, subatrial sagittal triactine, f, subatrial sagittal tetractine,
g, atrial triactine with short unpaired actine, h, atrial tetractine with short unpaired actine.
Skeleton. (Figs 45a–e) The walls of the atrial tube and the side-tubes bear continuous close-set semi-globular
(‘alveolar’) papillae (Figs 45a–b, d). Spicules of these alveolar papillae are predominantly characteristic ‘oxhorn-
shaped’ small triactines and tetractines (with upturned paired actines) in a single layer, but near the atrial lumen
there is an additional subatrial smaller sagittal triactine spicule located (Fig. 45b–c), differing from the alveolar
oxhorn-shaped spicules in having the paired actines recurved. The atrial wall (Fig. 45c, e) is supported by regular
larger tri- and tetractines, predominantly tetractines and smaller sagittal tri- and tetractines with paired actines
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almost in one plane and characteristic short unpaired actines. The skeletal structure of this species might be
interpreted as inarticulate.
Spicules. (Figs 44e, 46a–h) Regular equiangular equiactinal triactines and tetractines in two size categories,
oxhorn-shaped sagittal triactines and tetractines, small sagittal tri- and tetractines in two categories.
Atrial triactines (Fig. 46a) with unpaired actine up to 330–444.3–530 x 8–10.3–13 µm, paired actines up to
190–378.8–489 x 9–10.5–13 µm.
Atrial tetractines (Figs 46b, f) often with shorter unpaired actine, up to 210–359.2–479 x 9–11.8–14 µm, paired
actines up to 240–396.6–426 x 9–10.7–12 µm, and apical actines 18–21.8–34 x 4–6.6–10 µm.
Triactines with recurved actines (Fig. 46e) , unpaired actines 149–176.8–242 x 7.5–8.3–9 µm, paired actines
108-126.4-135 x 6.5-7.6-9 mm.
Triactines of the alveolar papillae, oxhorn-shaped (Fig. 46c), with unpaired actines 28–68.0–123 x 5–6.2–8
µm, paired actines 91–110.2–129 x 5–5.9–8 µm.
Tetractines of the alveolar papillae, oxhorn-shaped (Fig. 46d), with unpaired actine up to 48–106.1–180 x 8
µm, paired actines up to 93–120.8–135 x 6–6.8–8 µm and apical actine approximately 20–36.5–61 x 4.5–5.6–7 µm.
Atrial triactines, sagittal, with short unpaired actines (Fig. 46g), and paired actines almost in one plane,
unpaired actines 68–89.3–112 x 7–8.3–10 mm, paired actines 153–174.3–193 x 8–9.2–11 mm.
Atrial tetractines, sagittal, with short unpaired actines (Fig. 46h) and paired actines almost in one plane,
unpaired actines 18–33.2–53 x 8–8.6–11 mm, paired actines 88–172.0361 x 8–8.8–10 mm, apical actines 25–
47.4–95 x 7–8.1–9 mm.
Ecology. Shallow-reefs.
Distribution. So far known only from Indonesia, at Salayar and the Berau region.
Etymology. The word vinum (L.) means wine, tinctus (L.) means colored, referring to the wine-red color of the
species.
Remarks. The genus assignment is tentative. Among Calcaronea, the papillate surface and a wall of alveolar
choanocyte chambers only occur in the poorly known genus Sycetta, with type species the Indian Ocean S.
sagittifera (Haeckel, 1872), and other species the Mediterranean S. conifera (Haeckel, 1872), and two Antarctic
species, S. antarctica Brøndsted, 1931 and S. primitiva (Brøndsted, 1931) (originally as Tenthrenodes). The spicule
shapes and sizes of these species are clearly different from those of the present species, especially the Indian Ocean
S. sagittifera, which lacks tetractines. Oliver Voigt (personal communication) made histological sections of the
paratype, ZMA Por. 08633, and this confirmed that the alveolar structures are indeed syconoid choanocyte
chambers, with choanocytes lining the entire internal walls. He pointed out that there is a great similarity in skeletal
structure and histology with the South Australian ‘Sycon’ carteri Dendy, 1893. The two are probably closely
related, differing in habitus (‘S.’ carteri is a strongly divided bush of thin orange colored tubes) and spiculation
(‘S.’ carteri has brushes of short diactines on the papillae). Molecular work (Voigt et al. 2012) also demonstrated
that ‘S.’carteri is in its turn closely related to Leucascandra caveolata Borojevic & Klautau, 2000 (described also
in Lévi et al. (1998: 79). This species is rather more distinct in habitus and skeletal features: it is a dense mass of
orange–brown, smooth, delicate tubes provided with thinner side tubes, the walls of the tubes are ‘caveolate’, i.e.
rounded-syconoid (our interpretation), and there is a distinct cortex. There is no papillate surface as in Sycetta
vinitincta sp.nov. and ‘Sycon’ carteri, but the three species share similar oval syconoid choanocyte chambers and
the inarticulate skeleton. Leucascandra is at present assigned to the family Jenkinidae, which is, however,
postulated to be polyphyletic (Voigt et al. 2012; Dohrmann et al. 2006).
Genus Sycon Risso, 1827
Sycon spec.
Figures 47a–d, 48a–e
?Sycon raphanus var. tergestinum; Breitfuss, 1896: 434; Breitfuss, 1898: 173 (not: Haeckel, 1872)
Material examined. ZMA Por. 15509d, Indonesia, SE Sulawesi, SW Salayar, NW coast of Pulau Guang, 6.35°S
120.45°E, depth 3–4 m, SCUBA, H.A. ten Hove, Indonesian-Dutch Snellius II Expedition stat. 152/Cave 1/A3, 28
September 1984.
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FIGURE 47. Sycon spec., ZMA Por. 15509d, from Salayar, a, preserved habitus (scale bar = 2 mm), b, microscopic overview
of upper part and fringe (scale bar = 1 mm), c, cross section of tube wall showing palisade of small diactines at the distal cones,
and tubar skeleton of T-shape triactines (scale bar = 200 µm), d, overview of spicules (scale bar = 500 µm).
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FIGURE 48. Sycon spec., ZMA Por. 15509d, from Salayar, SEM images of spicules, a, thin diactine from the oscular fringe,
a1, detail of apex, b, large diactine, b1, detail of apex, c, small diactine of distal cone, d, T-shaped triactine from the tubar
skeleton, e, T-shaped triactine of the atrial skeleton, f, bouquets of diactiines of the distal cones.
Description. Tiny sponge (Fig. 47a), 4 mm high, 3 mm in diameter. Color cream in alcohol. Oscules provided
with silvery white-opaque stiff dense fringe, underneath which there is a loose collar of widespread diactinal
spicules (Fig. 47b, arrows). Surface of main body microconulose-spinose.
Skeleton. Typical radial arrangement (Fig. 47c) of tubar spicules lining the syconoid chambers, with distal
cones crowned by bouquets of diactines (Fig. 48f); atrial skeleton with several layers of triactines; central tube
crowned by peripheral fringe of long thin diactines (Fig. 47b), at the base of which there is a circle of thick
diactines forming a collar. These diactines are also found spread over the body.
Spicules. (Figs 47d, 48a–f) Tubar triactines, atrial triactines, fringe diactines, large collar diactines, small
diactines of the distal cones.
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Diactines of the oscular fringe (Figs 48a, a1), thin, but elongately fusiform, 1000–1500 x 6–10 µm.
Diactines of the collar (Figs 48b, b1), thick, fusiform, 500–650 x 15–22 µm.
Diactines of the distal cones curved (Figs 48c, f), at both ends tapering to sharp points, 120–220 x 5–6 µm.
Tubar triactines (Fig. 48d), sagittal, T-shaped, with unpaired actines longer than paired actines, unpaired
actines sharply pointed; unpaired actines 90–110 x 5–6 µm, paired actines 45–60 x 4–6 µm.
Triactines of the atrium (Fig. 48e), sagittal, T-shaped, with unpaired actines shorter than paired actines,
unpaired actines sharply pointed, 45–80 x 6 µm, paired actines 90–100 x 6 µm.
Ecology. Shallow-water reef cave.
Distribution. Indonesia.
Remarks. We consider this description as preliminary and e refrain from naming the present species, because
the single specimen was tiny and most of the material was used up in making a few inadequate preparations. The
habitus reminds of the Mediterranean Sycon humboldti (Risso, 1827), type species of the genus, but that has
tetractines. Only few Sycon species lacking tetractines have been described, e.g. Grantia cupola (Haeckel, 1872)
from Japan (it has no diactines), and Sycon grantioides Dendy, 1916 from the Indian Ocean (tubar triactines with
unpaired actines twice as long).
Breitfuss’ (1896, 1898) report of the Mediterranean (sub)species Sycon raphanus tergestinum from
Ternate may possibly concern our present species. No spicule data were provided by Breitfuss, and, of course, the
occurrence of a Mediterranean Sycon species in Indonesian waters is highly unlikely.
New Caledonian Sycon gelatinosum (Blainville, 1834), as redescribed by Borojevic & Klautau (2000) and
Northeast Australian Sycon capricorn Wörheide & Hooper, 2003 differ strongly from the present material by
having a branched tubular habitus, relatively smooth surface, possessing atrial tetractines, and differently shaped
spicules.
Family Grantiidae Dendy, 1893b
Genus Leucandra Haeckel, 1872
Leucandra irregularis (Burton, 1930) comb. nov.
Figures 49a–b, 50a–e
Anamixilla irregularis Burton, 1930: 6, fig. 5.
Not: Hôzawa, 1940: 155, pl. VII fig. 12.
Material examined. Holotype ZMA Por. 00144, Indonesia, Nusa Tenggara, Sumbawa, Bay of Bima near South
Fort, 8.505°S 118.695°E, depth 55 m, trawled, bottom mud with patches of coral sand, coll. Siboga Expedition stat.
047, 8 April 1899; BMNH 1929.8.30.6, five slides made from the holotype.
Description. Hispid thick-walled pear-shaped tube (Fig. 49a), size approximately 2.5 cm high, 1 cm in widest
diameter, dirty white in color (alcohol), with rather soft consistency. The original specimen had a narrow apical
opening with naked rim, but this has been cut off and mounted in one of the slides, kept in the Natural History
Museum.
Skeleton. (Fig. 49b) Due to the poor preservation, histological sections show a jumbled skeleton, but here and
there rounded choanocyte chambers are apparent indicating a leuconoid aquiferous system. There is a cortex of
smaller triactines (Fig. 49b top) overlying a choanosomal skeleton of confusedly arranged giant triactines, with
occasional giant diactines at sharp angles to the surface. The atrial skeleton (Fig. 49b bottom) is predominantly
formed by tetractines. The specimen orginally was in the possession of a naked osculum, supported by tangential
lengthwise arranged diactines, but this has not survived in the present remains. One of the slides made by Burton
and kept in the Natural History Museum collection shows the oscular skeleton to be formed by a dense mass of
tetractines similar to those of the atrial skeleton.
Spicules. (Figs 50a–e) Giant triactines, intermediate and smaller triactines, smaller tetractines, giant diactines.
Cortical triactines (Fig. 50d), actines relatively thin, usually sagittal or parasagittal, rarely regular equiangular
and equiactinal, paired actines not infrequently somewhat wobbly, unpaired actines 84–221.2–311 x 7–10.7–15
µm, paired actines, 123–263.2–510 x 6–10.4–15 µm.
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FIGURE 49. Leucandra irregularis (Burton, 1930), holotype ZMA Por. 00144 from Sumbawa, a, preserved holotype (scale
bar = 1 cm), b, cross section of the holotype BMNH 1929.8.30.6, showing an upper cortical skeleton of small triactines, a
choanosomal skeleton of giant triactines and a few oscular diactines, and an atrial skeleton of tetractines (scale bar = 500 µm).
Choanosomal giant or large triactines (Figs 50a–b), usually slightly sagittal, but with actines almost invariably
strongly different in length, even when equiangular. The shape somewhat resembles pseudosagittal triactines. Size
quite variable but recognizable by relatively thick actines, 270–597.1–1080 x 23–38.4–52 µm.
Atrial tetractines (Figs 50c), rather irregular in shape, apical actine prominent, unpaired actines, 174–293.3
420 x 5–10.3–13 µm, paired actines 120–287.2–438 x 4–10.8–17 µm, apical actines 20–60.8–120 x 4–6.8–10 µm.
Oscular diactines (Fig. 50e), occasionally also in the tubar skeleton, 820–1242.9–2100 x 45–55.5–63 µm.
Ecology. On soft bottom at 55 m.
Distribution. Indonesia, known only from the type locality off the island of Sumbawa.
Remarks. Burton assigned this species to Anamixilla probably because of the similarity of the complement of
giant triactines, which also often show a parasagittal shape with unequal actine lengths. However, the structure of
the skeleton and the aquiferous system differ clearly from Anamixilla torresi Poléjaeff, 1883 (see below).
Remarkably, at a later date (1963) Burton omitted to report the presence of giant diactines in this species, which is
one of the distinguishing features.
There is a compelling general similarity with Sri Lankan Leucandra donnani Dendy, 1905, which shares the
shape, skeletal organization and spiculation. Differences are found in the sizes of the various spicule types, with
overall smaller lengths and widths.
It also shows similarity with Japanese Leucandra rigida Hôzawa, 1940, which shares the giant triactines and
diactines, differing in the presence of small choanosomal diactines. Northeast Australian Leucandra sphaeracella
Wörheide & Hooper, 2003 has giant triactines and diactines, differing likewise from L. irregularis by having small
diactines, but these are found in the ectosomal cortex. Both species have the spicule sizes in the same range as
those of L. irregularis, so they are to be regarded as members of a regional complex of closely similar species.
Leucandra nicolae Wörheide & Hooper, 2003 is also similar in spicule sizes, but it lacks the giant diactines, and
has two types of microdiactines. A further close form probably is Leucandra tropica Tanita, 1943 from Palau (also
reported by De Laubenfels, 1954). It differs by the presence of cortical tetractines, in addition to atrial tetractines.
Hôzawa (1940) recorded this species from Haiti (Caribbean), but provided no description of the spicules.
The habitus figure he provided differs in shape from the holotype. Its identity is unclear, but in view of the
localities at opposite parts of the globe it is not likely conspecific with the Indonesian specimen.
The resemblance of the choanosomal triactines to pseudosagittal spicules is perhaps attributable to the possible
polyphyletic nature of these spicules, occurring in Grantiidae and Heteropiidae as discussed in Voigt et al. (2012).
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FIGURE 50. Leucandra irregularis (Burton, 1930), holotype ZMA Por. 00144, SEM images of the spicules, a, giant triactines,
b, intermediate-sized triactines of the choanosome, c, atrial tetractines, d, cortical triactines, e, oscular diactine.
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FIGURE 51. Anamixilla torresi Poléjaeff (1883), RMNH Por. 6572, a, habitus in situ in North Sulawesi (scale bar = 1 cm,
photo N.J. de Voogd).b, tangential view of surface skeleton (scale bar = 1 mm), c, histological cross section showing cortical
and choanosomal giant and smaller triactines, subatrial triactines and atrial tetractines (scale bar = 1 mm).
Family Jenkinidae Borojevic, Boury-Esnault & Vacelet, 2000
Genus Anamixilla Poléjaeff, 1883
Anamixilla torresi Poléjaeff, 1883
Figures 51a–c, 52a–c
Anamixilla torresi Poléjaeff, 1883: 50, pl. IV figs 2a–c; Dendy, 1893a: 97; Burton, 1930: 5, fig. 4; Borojevic, 1967b: 9; Erhardt
& Baensch, 1998: 73.
Material examined. ZMA Por. 00139, Indonesia, Irian Jaya, 1.71°S 130.79°E, depth 32 m, trawled, coll. Siboga
Expedition stat. 164, 20 August 1899; ZMA Por. 00140, Indonesia, Nusa Tenggara, Timor, Samau Island,
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Haingsisi, 10.1358°S 123.6297°E, depth 0–38 m, trawled, coll. Siboga Expedition stat. 303, 2 February 1900;
ZMA Por. 00141, Indonesia, Maluku, Banda anchorage, 4.5398°S 129.9084°E, depth 9–45 m, trawled, coll. Siboga
Expedition stat. 240, 22 November 1899; ZMA Por. 00142, Indonesia, Ambon anchorage, 3.7°S 128.15°E, depth
40 m, trawled, coll. Siboga Expedition stat. 231, 14 November 1899; ZMA Por. 08006, Indonesia, Nusa Tenggara,
E of Komodo, Teluk Slawi, 8.8°S 119.52°E, underneath coral head, depth 1–4 m, snorkeling, coll. R.W.M. van
Soest, Indonesian-Dutch Snellius II Expedition stat. 069/II/13, 17 September 1984; ZMA Por. 16351, Indonesia,
North Sulawesi, Bunaken Island, in cave, 1.6132°N 124.7797°E, depth 32 m, SCUBA, coll. H. Erhardt, 17
September 1996; RMNH Por. 6572, Indonesia, North Sulawesi, Lembeh Strait, Teluk Makawide, 1.48°N
125.24°E, depth 12 m, SCUBA, coll. N.J. de Voogd, #LEM19/090212/062, 9 February 2012; RMNH Por. 6612,
Indonesia, North Sulawesi, Lembeh Strait, Tanjung Pandea, 1.4°N 125.17°E, depth 14 m, SCUBA, coll. N.J. de
Voogd, #LEM24/110212/126, 11 February 2012.
FIGURE 52. Anamixilla torresi Poléjaeff (1883), RMNH Por. 6572, SEM images of spicules, a, equiangular parasagittal giant,
intermediate and small triactines, b, subatrial sagittal triactines, c, atrial tetractines (trichoxeas from the fringe not shown).
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Description. Wide irregular-shaped tubes (Fig. 51a) of up to 2.5 cm diameter and with short fringe at the rim
of flaring, trumpet-like tube-endings. Overall size of cormus up to 10 cm or more in length. Color white, with
blueish fringe in larger tubes. Consistency brittle, but limp when taken out of the water.
Histology. Syconoid aquiferous system.
Skeleton. Inarticulate, with a cortex formed by tangential giant and medium sized triactines (Fig. 51b), a
choanosomal skeleton (Fig. 51c) formed by similar giant triactines and the unpaired actines of subatrial triactines.
Atrial skeleton (Fig. 51c) consists of tetractines.
Spicules. (Figs 52a–c) Giant triactines, medium sized triactines, small triactines, medium sized and small
tetractines; these spicules are all parasagittal. There are also - mostly broken - trichoxeas, uncommon, which likely
represent spicules from the tubar fringe.
Giant triactines (Fig. 52a), equiangular with unequal length actines (parasagittal), 990–1640–2010 x 66–84.2
120 µm.
Medium-sized triactines (Fig. 52b) with unequal length actines (parasagittal), usually with an unpaired actine
longer than both unequal length paired actines, 330–471.3–602 x 13–24.7–35 µm.
Tetractines (Fig. 52c), with unpaired actines usually shorter than the paired actines, in a large size range from
giant-sized to small, 30–394.1–690 x 2–16.6–24 µm, paired actines 50–590.6–900 x 2–16.1–20 µm, apical actines
approximately 31–53.8–90 x 7–11.3–19 µm.
Trichoxeas, in bundles or loose, mostly broken, up to at least 300 x 2.5 µm (only observed in RMNH Por.
6572).
Ecology. Mostly in the deeper reef environment, 20–40 m.
Distribution. Indonesia, elsewhere North Australia and New Caledonia.
Remarks. This material generally conforms closely to Poléjaeff's description, and to brief characterizations of
Burton and Borojevic. The species is apparently not uncommon.
Anamixilla singaporensis sp. nov.
Figures 53a–c, 54a–d
?Leucosolenia sp. Lim et al.2008: 163.
Material examined. Holotype RMNH Por. 9350, Singapore, Pulau Subar Laut, NW, 1.2°N 103.83°E, depth 6 m,
SCUBA, coll. N.J. de Voogd, #SIN22/040406/160, 4 April 2006.
Description. Small group of whitish or pale beige tubes; a single basal tube may divide into two or three (Fig.
53a), ending in wide oscules, with frayed rim or coarse fringe. Diameter of tubes less than 1 cm, length up to 3 cm,
usually slightly constricted at the open end. Consistency brittle, easily damaged.
Skeleton. (Figs 53b–c) Inarticulate, with a cortical thick layer of giant and intermediate-sized triactines.
Choanosomal skeleton (Fig. 53b) formed by the unpaired actines of subatrial sagittal triactines. Atrial skeleton
(Fig. 53c) thin, built by tetractines, with their apical actines protruding into the atrial lumen. The oscular rim has a
thin spread of perpendicular trichoxeas.
Spicules. (Figs 54a–d) Giant and intermediate triactines, sagittal triactines, tetractines, thin diactines.
Equiangular equiactinal triactines (Fig. 54a) of the cortical region, in a wide size range from giant to
intermediate sizes, 165–331.1–905 x 7–26.2–78 µm.
Sagittal triactines (Fig. 54b), usually with long unpaired actine and wide-angled to almost straight paired
actines, unpaired actines 88–288.3–434 x 5–8.1–10 µm, paired actines 72–129.2–172 x 6–7.6–9 µm.
Tetractines (Fig. 54c), predominantly regular, but with apical actine often curved, unpaired actines 180–277.3
342 x 8–10.7–12 µm, paired actines 182–204.2–222 x 8–8.8–10 µm, apical actines 42–58.390 x 6–7.4–8 µm.
Thin diactines (Fig. 54d), usually broken, approximately 75–210 x 2–3 µm.
Ecology. Shallow water amongst filamentous algae.
Distribution. Known only from the type locality off the coast of Singapore.
Etymology. The name refers to the type locality.
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FIGURE 53. Anamixilla singaporensis sp. nov., holotype RMNH 9350 from Singapore, a, habitus in situ at 6 m depth (scale
bar = 1 cm, photo N.J. de Voogd), b, histological cross section showing syconoid aquiferous system and inarticulate skeleton of
giant triactines (scale bar = 500 µm); upper part: cortical skeleton of giant and intermediate-sized triactines, lower part: atrial
skeleton of tetractines, c, tangential view of atrial surface (scale bar = 200 µm), showing atrial tetractines and subatrial
triactines.
Remarks. The species is close to Anamixilla torresi in skeletal structure and spicular composition, but the
habitus of the new species is simply tubular without the flaring trumpet ending of A. torresi, and two spicule types
are distinctly different in shape and size. The actines of the giant triactines are only up to 905 µm in the present
species, against up to 2000 µ or more in A. torresi. The sagittal triactines of the choanosomal skeleton in our new
species have short straight paired actines; the unpaired actines are only up to 434 x 10 µm against up to 602 x 35
µm in A. torresi, which also has the angle of the paired actines more equiangular. Especially the atrial tetractines
show a striking difference in that in A. torresi the paired actines of these spicules are very often much longer than
the unpaired actines, whereas those of the new species are almost equal in length. Also, the unpaired actines of the
new species are only half the length of those of A. torresi (up to 342 x 12 against 690 x 24 µm in A. torresi), and the
paired actines only a third in length (up to 222 x 10 vs. up to 900 x 23 µm in A. torresi).
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FIGURE 54. Anamixilla singaporensis sp. nov., holotype RMNH 9350, SEM images of spicules, a, cortical and choanosomal
equiangular equiactinal triactines, b, subatrial sagittal triactine, c, atrial sagittal tetractine, d, trichoxea.
Genus Uteopsis Dendy & Row, 1913
Uteopsis argentea (Poléjaeff, 1883)
Figures 55a–b, 56a–c, 57a–g
Ute argentea Poléjaeff, 1983: 43, pl. I fig.3, pl. IV fig.3, pl. V figs 1a–p; Dendy, 1893a: 92.
Uteopsis argentea; Dendy & Row, 1913: 769; Burton, 1930: 5, fig. 3; Burton, 1963: 530.
Sycon sp.; Colin & Arneson, 1995: 61, photo 238.
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FIGURE 55. Uteopsis argentea (Poléjaeff, 1883), RMNH Por. 6593 from N Sulawesi, a, habitus in situ (photo B.W.
Hoeksema), b, preserved specimen (scale bar = 1 cm).
Material examined. ZMA Por. 00155, Indonesia, Lesser Sunda Islands, West Timor, Samau Island, Haingsisi,
10.205°S 123.4591°E, depth 23 m, trawled, coll. Siboga Expedition stat. 060, 27 April 1899; ZMA Por. 00182,
Indonesia, Irian Jaya, 1.7083°S 130.7916°E, depth 32 m, trawled, coll. Siboga Expedition stat. 164, 20 August
1899; RMNH Por. 1871, Indonesia, NE Kalimantan, Berau Islands, off Tanjung Batu, 2.2396°N 118.0934°E, 10–
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15 m, SCUBA, coll. B.W. Hoeksema, #BER.19/091003/158, East Kalimantan-Berau Expedition 2003, 9 October
2003; RMNH Por. 2598, Singapore, Pulau Subar Laut, NW, 1.2°N 103.83°E, depth 6 m, SCUBA, coll. N.J. de
Voogd, #SIN22/040406/160, 4 April 2006; RMNH Por. 6593, Indonesia, North Sulawesi, Lembeh Strait, Pulau
Abadi, 1.4336°N 125.2062°E, depth 30 m, SCUBA, coll. B.W.Hoeksema, #LEM22/100212/104, 10 February
2012.
FIGURE 56. Uteopsis argentea (Poléjaeff, 1883), RMNH Por. 6593, histological sections, a, lengthwise section through entire
tube showing cortical oxeas, inarticulate choanosomal skeleton formed by the unpaired actines of subatrial triactines, and atrial
skeleton of tetractines (scale bar = 500 µm), b, detail of cross section, showing from top downwards cortical triactines, giant
diactine, syconoid choanocyte chambers supported by subatrial sagittal triactines and containing numerous embryos, and atrial
skeleton of tetractines and triactines (scale bar = 200 µm), c, detail of choanosome showing amphiblastula embryos (scale bar =
50 µm).
Description. Bush of thin tubes (Figs 55a–b). Individual tubes up to 8 cm high, 3–4 mm in diameter,
maintaining this diameter over most or all their length. Tubes may divide into two at a short distance above the
substratum, but are unbranched for most of their length. Terminal oscules provided with a very short lighter colored
fringe. Smooth surface, tough-hard consistency. Color pale brown.
Histology. Syconoid aquiferous system (Fig. 56b), with oval-rounded choanocyte chambers.
Skeleton. (Figs 56a–c) Strong, dense cortex of aligned tangential giant diactines (Fig. 56a), covered by a thin
layer of small equiangular or sagittal triactines and scattered small diactines. Choanosomal skeleton inarticulate
(Fig. 56b), thin and predominantly formed by the unpaired actines of subatrial sagittal triactines, with a minority of
similar tetractines. Atrial skeleton of small sagittal tetractines and triactines, with scattered small diactines.
Spicules. (Figs 57a–g) Giant diactines, small diactines, triactines, tetractines.
Giant diactines (Fig. 57a) from the cortex, 12002170.03600 x 36–60.6–78 µm.
Small diactines (Fig. 57b), 105–211.0–360 x 1.5–3.4–8 µm.
Triactines from the cortical skeleton (Fig. 57c), unpaired actines 96–104.7–126 x 6–6.1–6.5 µm, paired actines
60–75.2–93 x 6–6.7–7 µm.
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FIGURE 57. Uteopsis argentea (Poléjaeff, 1883), RMNH Por. 6593, SEM images of the spicules, a, giant diactines, b, small
diactines, c, cortical triactine, d,choanosomal triactine, e, subatrial triactines, f, subatrial tetractine, g, atrial tetractines.
Triactines from the choanosome (Fig. 57d), unpaired actines 252–430.4–576 x 5–7.8–11 µm, paired actines
96–180.6–258 x 5–8.6–9 µm.
Triactines from the atrial skeleton (Fig. 57e), unpaired actines 99–121.2–135 x 4–5.1–7 µm, paired actines
142–155.3–165 x 6–7.0–9 µm.
Tetractines from the choanosome (Fig. 57f), unpaired actines 250–301.7–384 x 8–8.1–8.5 µm, paired actines
126–159.0–181 x 6–8.3–11 µm, apical actines 45–58 x 6 µm.
Tetractines from the atrial skeleton (Fig. 57g), unpaired actines 96–185.3–312 x 5–5.7–6 µm, paired actines
96–114.0–129 x 5–6.7–8 µm, apical actines 20–24 x 4 µm.
Ecology. Deeper parts of the reefs.
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Distribution. Indonesia, elsewhere South Australia.
Remarks. The type of Uteopsis argentea from the SE coast of Australia is similar in shape and overall
spiculation, but there are a few discrepancies. Atrial tetractines of the type have much shorter unpaired actines, and
the unpaired actine of the choanosomal triactines may be up to 750 µm in length. These differences are considered
too small for specific separation. Future records of the species may find consistent differences among the widely
separated populations.
FIGURE 58. Sycettusa sibogae (Burton, 1930), holotype ZMA Por. 00148 from Kalimantan, a, habitus of preserved holotype
(scale bar = 1 cm), b–d, BMNH 1929.8.30.2a slides of holotype, b, cross section showing inarticulate skeleton made up of the
paired actines of pseudosagittal triactines on the outside covered by the paired actines of regular giant triactines, on the atrial
side with a cover of atrial sagittal triactines (scale bar = 500 µm), c, tangential view of the skeleton of the oscule showing a <