Biemna humilis Thiele, 1903: 944
Desmacella tubulata sensu Dendy 1916: 116
Sigmatoxella humilis
Pulitzer-Finali 1982a: 107
Mycale aff. c ockburniana
Van Soest 1990: 305
Erhardt & Baensch 1998: 54
Desmacella humilis
Pulitzer-Finali 1993: 293
Kerasemna humilis
Richmond et al. 2011: 122
Mycale (Mycale) vansoesti Calcinai et al ., 2006: 197
Mycale (Mycale) corallina Calcinai et al. 2017
Mycale calcinaiae Van Soest & Hooper, 2020: 63
Mycale humilis
Mycale species of the tropical Indo-West Pacific (Porifera, Demospongiae, Poecilosclerida)
Van, Rob W. M.
Aryasari, Ratih
De, Nicole J.
Zootaxa
2021
2021-01-19
4912
1
1
212
86X4L
(Thiele, 1903)
Van & Aryasari & De
2021
Thiele
1903
[151,680,834,861]
Demospongiae
Mycalidae
Mycale
Animalia
Poecilosclerida
97
98
Porifera
species
humilis
comb. nov.
Kerasemna
Figs 61a–d, 62a–f, 63a–g, 64
Biemna humilis Thiele, 1903: 944, fig. 10. ? Desmacella tubulata sensu Dendy 1916: 116(not: Dendy 1905). Sigmatoxella humilis; Pulitzer-Finali 1982a: 107. Mycaleaff. c ockburniana; Van Soest 1990: 305; Erhardt & Baensch 1998: 54(not: Hentschel 1911) ? Desmacella humilis; Pulitzer-Finali 1993: 293. ? Kerasemna humilis; Richmond et al.2011: 122, fig. p. 123. Mycale (Mycale) vansoesti Calcinai et al., 2006: 197, figs 3A–C, 4A–F, 5A–J, 6A–F (not: Hajdu 1995). Mycale (Mycale) corallina Calcinai et al.2017: Supplementary file 2. Mycale calcinaiaeVan Soest & Hooper, 2020: 63. Mycale humilis; Van Soest & Hooper, 2020: 63.
Material examined. ZMB3203, two slides of holotypeof Biemna humilis(slides labeled as Desmacella humilis), Indonesia, Ternate, coll. W. K̹kenthal, 1893–1894; MSNG52803, paratypeof Mycale (Mycale) vansoesti Calcinai et al., 2006, Indonesia, N Sulawesi, Siladen Station, depth 17 m, 28 May 2002. ZMA Por.02905, Indonesia, Lesser Sunda Islands( Nusa Tenggara), Postillion Islands, anchorage off Pulau Sarassa, 7.1833°S 118.2°E, coral bottom, depth 36 m, dredge, coll. Siboga Expedition stat. 043, field nr. SE829I, 4 April 1899; ZMA Por.08028, Indonesia, Nusa Tenggara East, Komodo, NEcape, 8.4833°S 119.5683°E, depth 1–4 m, snorkeling, coll. R. W.M. van Soest, Indonesian-Dutch Snellius II Expeditionstat. 096, field nr. 096/ II/02, 19 September 1984(live color yellow); ZMA Por.08564, Indonesia, Sulawesi, SE Sulawesi, Taka Karlarang, reef, depth 6–8 m, SCUBA, coll. R. W.M. van Soest, Indonesian-Dutch Snellius II Expeditionstat. 147, field nr. 147/ III/11, 27 September 1984(yellow); ZMA Por.08953, Sulawesi, SE Sulawesi, SW Salayar, reef N of Pulau Bahuluang, 6.45°S 120.43°E, depth 10–15 m, SCUBA, coll. R. W.M. van Soest, Indonesian-Dutch Snellius II Expeditionstat. 169, field nr. 169/ IV/24, 30 September 1984(light yellow); ZMA Por.09753, Papua New Guinea, no further data, dried, coll. M.C. Díaz, field nr. 90152 (orange-red); ZMA Por.12970, Indonesia, N Sulawesi, Bunaken Island, 1.6132°N 124.7797°E, reef drop-off, dried, coll. H. Erhardt, 22 September 1996(white-purple, photo in Erhardt & Baensch 1998); ZMA Por.13327, Indonesia, Sulawesi, SW Sulawesi, Spermonde Archipelago, Samalona, reef, depth 6 m, SCUBA, coll. N.J. de Voogd, field nr. SA/NV/060597/03, 6 May 1997(white-purple); ZMA Por.14523, Indonesia, N Sulawesi, Bunaken, ESE Siladen Island, 1.6317°S 124.8053°E, depth 18 m, coll. B.W. Hoeksema, SYMBIOSPONGE Expedition, field nr. 98/NS/MAY07/BH/077, 7 May 1998(white); ZMA Por.15829, Madagascar, Nosy B, Crater Bay, depth 4–10 m, muddy reef bottom, SCUBA, coll. J.H. Stock, 21 December 1963(orange); ZMA Por.16832, Indonesia, Maluku, Ambon, Tanjung Setan, depth 20 m, SCUBA, coll. S. Weinberg, 20 August 1995(orange); RMNH Por.1608, Palau, Koror, Ngerikuul Pass, E of Ngetekiou Island, pass between Ngeteklouand Ngermeuangel Island, 7.3167°N 134.5167°E, depth 8 m, SCUBA, coll. N.J, de Voogd, field nr. KOR02/120505/003, 12 May 2005(green); RMNH Por.1609, Palau, Koror, Ngerikuul Pass, E of Ngetekiou Island, pass between Ngeteklouand Ngermeuangel Island, 7.3167°N 134.5167°E, SCUBA, coll. N.J, de Voogd, field nr. KOR02/210505/095, 21 May 2005; RMNH Por.1610, Palau, Koror, Ngerikuul Pass, E of Ngetekiou Island, pass between Ngeteklouand Ngermeuangel Island, 7.3167°N 134.5167°E, SCUBA, coll. N.J, de Voogd, field nr. KOR02/210505/096, 21 May 2005; RMNH Por.2150, Bali, Nside of Nusa Penida, off Desa Buyuk, 8.6736°S 115.5436°E, depth 0–30 m, SCUBA, coll. N.J. de Voogd, Bali-Lombok Strait Expedition2001, field nr. BAL.34/210401/259, 21 April 2001(white); RMNH Por.2190, Bali, Padang Bai, Eside Tanjung Sari, 8.5294°S 115.5131°E, shallow slope, depth 0–30 m, SCUBA, coll. N.J. de Voogd, Bali-Lombok Strait Expedition2001, field nr. BAL.19/080401/097, 8 April 2001(white); RMNH Por.2608, Singapore, Pulau Tekukor( Monkey Island), NW side, 1.2308°N 103.8365°E, depth 12 m, SCUBA, coll. N.J. de Voogd, field nr. SIN.23/040406/170, 4 April 2006(yellow); RMNH Por.2617, Indonesia, Sulawesi, SW Sulawesi, Tanjung Bira, Pulau Liukan, 5.65°S 120.43°E, depth 25 m, SCUBA, coll. N.J. de Voogd, field nr. BIR05/230501/242, 23 May 2001; RMNH Por.2618, Indonesia, North Sulawesi, SW Siladen Island, 1.6167°N 124.7833°E, depth 15 m, SCUBA. coll. N.J. de Voogd, field nr. MD11/170502/079, 17 May 2002; RMNH Por.3054, Indonesia, Sulawesi, SW Sulawesi, Tanjung Bira, Pulau Liukan, 5.65°S 120.43°E, depth 28 m, SCUBA, coll. N.J. de Voogd, field nr. BIR05/230501/321, 23 May 2001; RMNH Por.5329, Indonesia, Halmahera, Maitara Maitara W, 0.7299°N 127.3624°E, depth 12 m, SCUBA, coll. N.J. de Voogd, Ternate-Halmahera Expedition2009, field nr. TER.09/291009/080, 29 October 2009; RMNH 8867, Taiwan, Dong YiJu, west side, outside fishing harbor, 23.2552°N 119.6671°E, SCUBA, coll. N.J. de Voogd, field nr. PES135, 28 July 2014(orange).
Description( Figs 61a–d, 63a). The species was recently described extensively by Calcinai et al.(2006)as Mycale (Mycale) vansoesti. We can confirm their excellent account of the species, also based on slides made from a fragment of the paratypesent to us by Dr. Barbara Calcinai. The specimens in their full-grown habitus exhibit wide-mouthed tubes, often two such tubes are found merged to a larger mass with two openings. Size may be considerable: Calcinai et al. (2006)report specimens of 40 cmhigh and 20 cmwide, with terminal vent up to 15 or 20 cm. The walls of the tube are thin (often less than 1 cmin thickness), web-like, supported by purple- or red-colored algal strands (genus Amphiroa). Juvenile or incipient growth forms may be much smaller and basically are encrusting. These may lack an upper large opening, in stead having lateral excurrent openings giving access to the hollow interior. The surface is irregularly conulose, caused by Amphiroastrands lifting up the surface tent-like. The skin between the strands, both on the outside and the inside of the tubes is punctured by many small incurrent and excurrent openings, of about 1 mmor less in diameter. The inner surface of the vent has also irregularly distributed larger openings, presumably oscules. Colors of larger specimens sometimes give an overall pinkish impression ( Figs 61a–b), due mostly to the algal strands, but the tissue inbetween is pale yellow, and yellow-orange colors ( Fig. 61c) are also reported frequently, especially in smaller encrusting specimens, occasionally white, rarely greenish. So far the colors are not clearly correlated to other features of the species, so we assume them to be variable. Preserved, colors are off-white or light beige ( Figs 61d, 63a). Consistency is fragile, easily damaged. Skeleton( Figs 62a–f). The algal strands (thickness 250–400 µm) form the framework of the skeleton, but spongin-encased spicule tracts of variable thickness (commonly 30–70 µmin diameter, occasionally as thick as 200 µm) and length (may be as long as 1 mmor more) connect the algal strands to form a secondary supporting structure. Between the algal strands and the spicule tracts there is an unorganized mass of toxodragmas, anisochelae, sigmas, rare trichodragmas and single tangentially arranged megascleres. No clear ectosomal skeleton on both sides of the tube walls. The toxodragmas may form vague clusters, but sigmas and anisochelae are scattered singly in the tissue, no rosettes. Spicules( Figs 63b–g). Mycalostyles, two categories of anisochelae, sigmas, toxas (mostly in toxodragmas), trichodragmas. Mycalostyles ( Fig. 63b,b 1), comparatively short and robust, with barely developed head, slightly fusiform, 207– 242.7–297 x 4– 6.7– 11 µm; typespecimen: 231– 267.3–297 x 4.5– 7.4– 11 µm. Anisochelae I ( Figs 63c), usually less common, narrow-shaped but otherwise normal, with both upper and lower alae developed, free part of the shaft about 40–50 %, size 17– 22.9– 30 µm; typespecimen: 17– 19.4– 22 µm. Anisochelae II ( Figs 63d), usually more common, comparatively slightly more reduced with upper and lower alae pointed towards each other, 10– 13.4– 19 µm; typespecimen: 10– 13.3– 16 µm. Sigmas ( Fig. 63e), usually very common, thin, asymmetrically curved, the endings look roughened under SEM, but no definite spines were observed, 16– 24.9– 30 µm; typespecimen: 19– 24.8– 30 µm. Toxodragmas ( Figs 63f,f 1), extremely common, consisting of an intertwined double-pack of individual thin raphide-like toxas (less than 1 µmin thickness), leaving a characteristic eye-like central opening, 33– 42.5–51 x 4– 6.6– 10 µm; typespecimen: 34– 43.9–51 x 4– 7.4– 10 µm. FIGURE 61. Mycale (Kerasemna) humilis( Thiele, 1903), in situand preserved images from Indonesia, a–b, ZMA Por. 14523, from Bunaken Island, N Sulawesi, (photos B.W. Hoeksema), c, RMNH Por. 2617 from SW Sulawesi (photo N.J. de Voogd), d, ZMA Por. 08953 from Bahuluang Island, Salayar, preserved habitus (scale bar = 1 cm). FIGURE 62. Mycale (Kerasemna) humilis( Thiele, 1903), a–d, light microscopic images from histological slides of the holotype of Biemna humilis, ZMB3203, from Ternate, Indonesia, a, algal strand encrusted by skeletal tracts and microscleres, b, detail of a, c, massed presence of toxodragmas in the tissue, d, microscleres, including sigmas, toxodragmas, anisochelae I (arrow I) and anisochelae II (arrow II), e–f, light microscopic images of ZMA Por. 16382 from Ambon, Indonesia, e, skeletal structure of algal strands and interconnecting spicule tracts, f, detail of e. Trichodragmas ( Fig. 63g), not common, consisting of a small package of straight raphides, 7– 11.8–17 x 3– 6.6– 10 µm; typespecimen: 9– 11.9–15 x 3– 6.4– 9 µm.
Distribution and ecology( Fig. 64). Indonesia, Singapore, Papua New Guinea, Palau, Taiwan, Madagascar, Kenya,?Western India, on reefs down to 36 m.More ecological information is provided in Calcinai et al.(2006).
Remarks. We re-examined original slides of Thiele’s Biemna humilisand found abundant anisochelae I and II, apparently overlooked by Thiele. In most aspects, Biemna humilisconforms with what is so far known as Mycaleaff. cockburniana sensu Van Soest 1990and Mycale (Mycale) vansoesti Calcinai et al., 2006. Thiele’s material was a blade-like encrustation, not an elaborate wide-tubed specimen, but in spiculation and presence of Amphiroastrands there is complete conformity. Calcinai et al.’s name is a primary junior homonym of Mycale (Grapelia) vansoestiHajdu, 1995, which has priority. Calcinai et al. 2017proposed Mycale (Mycale) corallinato replace M. (M.) vansoestiCalcinai et al., because junior primary homonyms are permanently invalid (ICZN Art. 57.2). Van Soest & Hooper (2020: 63) were unaware of this replacement name (which was proposed in a Supplementary file, and at the time was not picked up by the World Porifera Database) and proposed Mycale calcinaiaeas a replacement name. Both names are junior synonyms of Mycale (Kerasemna) humilis( Thiele, 1903) comb.nov. FIGURE 64. Mycale (Kerasemna) humilis( Thiele, 1903), approximate distribution of specimens studied (green squares) and literature records (pale green dots). Squares and dots may represent several close collecting localities. Calcinai et al.’s description does not entirely conform to our above description as it omitted reporting the small trichodragmas, which were found in all our specimens, including Calcinai’s paratype. Because they are quite small and not at all very common, we assume that they were overlooked by Calcinai et al. Although Pulitzer-Finali (1993)reported the present species from Kenyaas Desmacella humilis, he pointed out in the discussion that it shared the toxodragmas with his Australian Kerasemna tenuityla. Subsequently, Kelly in Richmond et al. 2013, used the combination Kerasemna humilisfor our species. We confirm here the presence of the species in the Western Indian Ocean by reporting a specimen from Madagascar. The spicule shape and sizes (mycalostyles 244–249 x 6–7.5 µm, anisochelae I 22–25 µm, anisochelae II 17–19 µm, sigmas 21–29 µm, toxodragmas 45–50 µm, and trichodragmas 12–17 µm) conform to those of the numerous specimens we report from the Western Pacific. Apparently orange colored specimens are more common there, as both our specimen, Pulitzer-Finali’s specimens and Richmond’s specimen are reported to have that color. Still, orange and yellow specimens were also reported from Indonesia(see Fig. 61c), Singapore, Taiwanand Papua New Guinea. The coralline algae found associated with the species recognized so far appear to be mostly Amphiroaspec., at least in the pink, purple, or pale yellow-white forms. Possibly, the association is thus obligatory, but this may be only for these forms. Yellow-orange specimens may have a different algal associate (cf. Calcinai et al.2006). Pulitzer-Finali’s (1993) Kenyaand Zanzibarrecords mention the presence of toxodragmas and sigmas, but do not report anisochelae. It is possible that the specimens do not contain anisochelae and belong to M. (K.) tenuiylarather than to M. (K.) humilis. Dendy’s (1916)report of a deviating Desmacella tubulataappears on paper to be a likely record of the present species from Western India, as Dendy mentions ‘toxa arranged in a sheaf or toxodragma’. This will need to be reexamined for certainty. Mycale cockburniana Hentschel, 1911and its variety albanensis Hentschel, 1911, generally shares the spicule complement with the present species: mycalostyles 192–256 x 3–4 µm, two size categories of anisochelae 21–27 µmand 13–16 µm, thin sigmas 22–25 µm, and trichodragmas 25–30 µm. The habitus is encrusting and contains ingrown ‘Pflanzentheile’, adding to the similarity. The main difference is the absence of the peculiar toxodragmas. The skeleton is described as conforming to subgenus Carmia(cf. above). Kerasemna tenuitylaPulitzer-Finali, 1982(see below) is a close relative, differing mostly in the absence of anisochelae. We maintain this as a separate species for the time being.
3026466452
ZMA
Indonesia
Por.
36
-7.1833
Pulau Sarassa
5511
118.2
Postillion Islands
97
98
1
Nusa Tenggara Timur
3026466765
ZMA
Komodo, NE
Indonesia
3
-8.4833
East
7
119.5683
Por.
97
98
1
Nusa Tenggara Timur
3026466489
[429,950,1501,1528]
R
W. M. van Soest & Indonesian-Dutch Snellius
Indonesia
97
98
1
Nusa Tenggara Timur
3026466716
[959,1368,1501,1528]
II
Expedition
Indonesia
97
98
1
Nusa Tenggara Timur
3026466400
1984-09-19
II
Indonesia
97
98
II/02
1
Nusa Tenggara Timur
3026466855
ZMA
Indonesia
7
Taka Karlarang
Por.
97
98
1
Sulawesi Utara
3026466680
[480,1005,1573,1600]
R
W. M. van Soest & Indonesian-Dutch Snellius
Indonesia
97
98
1
Sulawesi Utara
3026466408
[1014,1427,1573,1600]
II
Expedition
Indonesia
97
98
1
Sulawesi Utara
3026466722
[151,553,1609,1636]
1984-09-27
III
Indonesia
97
98
III/11
1
Sulawesi Utara
3026466538
ZMA
SW Salayar
Indonesia
13
-6.45
Por.
782
120.43
97
98
1
Sulawesi Utara
3026466444
[769,1282,1645,1672]
R
W. M. van Soest & Indonesian-Dutch Snellius
Indonesia
97
98
1
Sulawesi Utara
3026466455
II
Expedition
Indonesia
97
98
1
Sulawesi Utara
3026466644
[408,873,1681,1708]
1984-09-30
IV
Indonesia
97
98
IV/24
1
Sulawesi Utara
3026466652
ZMA
M. C. Diaz
Papua New Guinea
Por.
97
98
1
3026466472
1996-09-22
ZMA
H. Erhardt
Indonesia
1.6132
Bunaken Island
7
124.7797
Por.
97
98
1
Sulawesi Utara
3026466414
ZMA
Indonesia
Por.
6
Samalona
Spermonde Archipelago
97
98
1
Sulawesi Utara
3026466807
[700,1231,1825,1852]
1997-05-06
SA
Indonesia
97
98
1
Sulawesi Utara
3026466922
ZMA
Indonesia
Por.
18
-1.6317
Siladen Island
7
124.8053
Bunaken
97
98
1
Sulawesi Utara
3026466441
ZMA
Nosy B
Madagascar
7
Crater Bay
Por.
97
98
1
3026466518
[823,1432,1933,1960]
1963-12-21
SCUBA
J. H. Stock
Madagascar
97
98
1
3026466707
[151,989,1969,1996]
ZMA
Indonesia
20
Tanjung Setan
Por.
97
98
1
Maluku Utara
3026466448
1995-08-20
SCUBA
S. Weinberg
Indonesia
97
98
1
Maluku Utara
3026466606
RMNH
Palau
Por.
8
98
99
7.3167
Ngeteklou
7
134.5167
Ngerikuul Pass
97
98
1
Koror
3026466493
[783,1191,151,177]
SCUBA
Voogd
Palau
98
99
1
Koror
3026466527
2005-05-12
KOR
Palau
98
99
KOR02
1
Koror
3026466478
RMNH
Palau
Por.
7.3167
Ngeteklou
7
134.5167
Ngerikuul Pass
98
99
1
Koror
3026466601
[764,1189,223,249]
SCUBA
Voogd
Palau
98
99
1
Koror
3026466702
2005-05-21
KOR
Palau
98
99
KOR02
1
Koror
3026466314
RMNH
Palau
Por.
7.3167
Ngeteklou
7
134.5167
Ngerikuul Pass
98
99
1
Koror
3026466723
[724,1152,295,321]
SCUBA
Voogd
Palau
98
99
1
Koror
3026466877
2005-05-21
KOR
Palau
98
99
KOR02
1
Koror
3026466459
[281,1431,330,357]
RMNH
Bali, N
Palau
Por.
15
-8.6736
Desa Buyuk
7
115.5436
Nusa Penida
98
99
1
Koror
3026466754
2001-04-21
SCUBA
N. J. de Voogd & Bali-Lombok Strait Expedition
Palau
98
99
1
Koror
3026466827
RMNH
Padang Bai, E
Palau
Por.
15
-8.5294
Tanjung Sari
7
115.5131
Bali
98
99
1
Koror
3026466357
2001-04-08
SCUBA
N. J. de Voogd & Bali-Lombok Strait Expedition
Palau
98
99
1
Koror
3026466491
RMNH
Singapore
Por.
12
1.2308
Monkey Island
7
103.8365
Pulau Tekukor
98
99
1
3026466921
[294,1216,511,538]
2006-04-04
SCUBA
N. J. de Voogd
Singapore
98
99
1
3026466554
RMNH
Indonesia
Por.
25
-5.65
Pulau Liukan
783
120.43
Tanjung Bira
98
99
1
Sulawesi Utara
3026466879
2001-05-23
SCUBA
N. J. de Voogd
Indonesia
98
99
1
Sulawesi Utara
3026466651
RMNH
SW Siladen Island
Indonesia
15
1.6167
Por.
7
124.7833
98
99
1
Sulawesi Utara
3026466790
[1054,1432,618,645]
2002-05-17
MD
Indonesia
98
99
MD11
1
Sulawesi Utara
3026466334
[151,1431,654,681]
RMNH
Indonesia
Por.
28
-5.65
Pulau Liukan
783
120.43
Tanjung Bira
98
99
1
Sulawesi Utara
3026466509
[151,961,690,717]
2001-05-23
SCUBA
N. J. de Voogd
Indonesia
98
99
1
Sulawesi Utara
3026466374
RMNH
Maitara Maitara W
Indonesia
12
0.7299
Halmahera
7
127.3624
Por.
98
99
1
3026466364
2009-10-29
SCUBA
N. J. de Voogd & Ternate-Halmahera Expedition
Indonesia
98
99
1
3026466742
RMNH
Taiwan
23.2552
Dong YiJu
7
119.6671
98
99
RMNH 8867
1
3026466339
[598,1381,798,825]
2014-07-28
SCUBA
N. J. de Voogd
Taiwan
98
99
1