New shallow-water sponges (Porifera) from the Galápagos Islands Author Sim-Smith, Carina Author Hickman, Cleveland 0000-0002-2914-4687 hickman.c@rockbridge.net Author Kelly, Michelle 0000-0001-9673-0056 elly@niwa.co.nz text Zootaxa 2021 2021-08-02 5012 1 1 71 http://dx.doi.org/10.11646/zootaxa.5012.1.1 journal article 10.11646/zootaxa.5012.1.1 1175-5326 5158062 56C6852D-AAE0-4B6B-AB57-919CD62DAEC1 Dragmacidon raeae sp. nov. ( Fig. 12 ) Dragmacidon sp. Lizarazo Rodríguez et al. 2020 , 88–89, fig. 4, pl. 1 fig. 3. Material examined. Holotype — MCCDRS9411, Maria Elena Islet , Pinzón Island , 0.592° S , 90.684° W , 9 m , 20 Jan 2003 . Type locality. Maria Elena Islet , Pinzón Island ; 9 m . Habitat and distribution. Maria Elena Islet, Pinzón Island , Galápagos; and Punta Cruces, La Parguera-Piñas , Colombia . Found growing on rock substrate; 6–9 m . Description. Thickly encrusting to hemispherical sponge, 15 mm thick. Surface densely punctate, velvety to the touch. Small clusters of up to three oscules, each around 1 mm in diameter, are scattered over the sponge, surrounded by prominent asterose drainage canals. Texture is firm, barely compressible, but easily broken. Colour in life is bright red, colour in ethanol is golden brown ( Fig. 12A ). Skeleton. Choanosomal skeleton is plumoreticulate with clear, wavy multispicular primary tracts, and uni- to paucispicular secondary tracts, irregularly connecting the primary tracts. Tips of styles protrude slightly beyond the surface of the sponge. There is no special ectosome ( Fig. 12C ). Spicules . Megascleres— styles ( Fig. 12B ), short, stout and usually strongly bent near the blunt end, 368 (324– 397) × 19 (16–23) µm (n = 20). Etymology. Named for co-author Cleveland Hickman’s wife Rae, who accompanied him on numerous research trips to the Galápagos , where she additionally provided support and assistance to the library at the Charles Darwin Research Station. Remarks. Dragmacidon raeae sp. nov. is relatively unusual for the genus in that it only possesses styles instead of both oxeas and styles. In terms of morphology and skeletal architecture, D . raeae sp. nov. is strongly reminiscent of the southwest Pacific species, Dragmacidon australe ( Bergquist, 1970 ) , which also forms a bright red cushion, but which has oxeas and a larger size range of megascleres. Three other described species, D. coccineum ( Keller, 1891 ) , D. fibrosum ( Ridley & Dendy, 1886 ) and D. tumidum ( Dendy, 1897 ) , also only possess styles ( Table 4 ). All other characteristics of D. raeae sp. nov. are typical of some members of the genus (velvety surface, plumoreticulate skeleton, the lack of ectosome, and undifferentiated axial and extra-axial regions). TABLE 4. Comparison of current species of Dragmacidon . STY = styles; STR: strongyloxeas; OX = oxeas; TRD = trichodragmata; RHP = rhapides.

Species	Distribution	Morphology	Skeleton	Spicules (µm)
Dragmacidon raeae sp. nov.	Galápagos	Thickly encrusting to hemispheri-	Plumoreticulate, multispicular	STY: 368 (324–397) × 19 (16–23)
cal, firm, velvety, densely punctate,	primary tracts and irregular uni- to
prominent drainage canals around	paucispicular secondary tracts.
oscules, bright red.
Dragmacidon hendersoni	Galápagos	Thinly encrusting, minutely ridged,	Loosely plumoreticulate, multi-	STY I: 231 (209–248) × 6 (4–7)
sp. nov.	densely punctate, compressible,	spicular primarily tracts and vague	STY II: 197 (127–334) × 2 (1–3)
bright yellow.	secondary tracts. No ectosome.
Dragmacidon agariciforme	Sri Lanka, Arabian Sea, Mozam-	Massive, cushion shaped to cup-	Plumoreticulate, primary tracts	STY I: 213 (165–335) × 7 (5–10)
( Dendy, 1905 ), type	bique	shaped with a short stalk, minutely	terminating at conules, uni- to	STY II: 658 (650–750) × 6 (5–8),
conulose, grooved, dermal mem-	multispicular irregular second-	rare
brane, firm, compressible, brick red.	ary tracts. STY II are scattered	OX: 225 (185–298) × 8 (3–10)
between tracts. Spongin abundant.	TRD: present
No ectosome.
Dragmacidon alvarezae Zea	Caribbean, Florida, Gulf of Mexi-	Massive to encrusting, smooth to	Plumoreticulate, multispicular pri-	STY: 170–390 × 5–28
& Pulido, 2016	co, north coast of South America,	hirsute, oscules ≤ 5 mm, toughly	mary tracts terminate in brushes,	OX: 210–420 × 5–18
Bahamas.	compressible, abundant mucus, red	multispicular secondary tracts.
to bright orange.
Dragmacidon australe	Australia, NZ, New Caledonia	Massive to thickly encrusting,	Plumoreticulate to halicondroid,	STY I: 203–560 × 9–22
( Bergquist, 1970 )	conulose, ridged, hispid, dermal	plumose or plumo-echinated (core	STY II: 320–406 × 3–6
membrane between ridges, oscules ≤	of oxeas echinated by styles)	OX: 217–339 ×8–10
1 mm, abundant mucus, firm, slightly	multispicular primary tracts, pau-
compressible, bright scarlet.	cispicular secondary tracts, STY II
at surface, sparse spongin.
Dragmacidon clathriforme	Australia	Massive, lobose, oscules ≤ 10 mm,	Loose bundles of spicules, dense,	OX*: 330 × 11
( Von Lendenfeld, 1888 )	smooth, colour unknown.	confused mass.
Dragmacidon coccineum	Red Sea, Arabian Sea, Seychelles,	Hemispherical to spherical, firm,	Reticulated, no ectosome.	STY: 300–320 × 10
( Keller, 1891 )	Zanzibar, Kenya	ridged, bumpy, punctate, few oscules
3–5 mm, bright orange-red.

......continued on the next page ......continued on the next page ......continued on the next page TABLE 4. (Continued)

Species	Distribution	Morphology	Skeleton	Spicules (µm)
Dragmacidon condylia	New Caledonia	Thickly encrusting with bulbous	Plumoreticulate, multispicular	OX/STY**: 260 (208–289) × 12
( Hooper & Lévi, 1993 )	digitate projections, apical oscules,	primary tracts terminate in brushes	(10–14)
membranous surface, punctate, pale	protruding slightly beyond surface,
orange.	paucispicular secondary tracts,
sparse spongin, no ectosome.
Dragmacidon debitusae	New Caledonia	Massive, microconulose, dermal	Plumoreticulate, multispicular	OX/STY (rare)/STR (rare): 364
( Hooper & Lévi, 1993 )	membrane, fleshy surface, oscules	primary tracts terminate in sparse	(223–503) × 9 (2–15)
5–10 mm, compressive, orange to	brushes protruding slightly
beorange-yellow.	yond surface. Secondary uni- to
paucispicular tracts. Reticulate
spongin fibre skeleton also
present.
Dragmacidon decipiens	Australia	Massive, smooth to verrucose, mi-	Confused, a few tracts, reticulate	OX/STY/STR: 350 (278–483) ×
( Wiedenmayer, 1989 )	crohispid, oscules minute, very firm,	spongin fibre skeleton.	8 (4–11)
tan with greenish grey patches.	STR: 542–770 × 8–9, sinuous,
rare.
Dragmacidon durissimum	Sri Lanka, South India, Red	Massive, microconulose, dermal	Plumoreticulate, primary tracts	STY: ≤ 390 × ≤ 24
( Dendy, 1905 )	Sea, Arabian Sea, East Africa,	membrane, oscules small surrounded	terminate in conules, uni- to pau-	OX: ≤ 390 × ≤ 24
Seychelles, northern Australia,	by faint asterose drainage canals,	cispicular secondary tracts, free	TRD: hair-like
Indonesia	very hard, harsh to touch, yellow-	spicules scattered between tracts.
grey in alcohol.
Dragmacidon egregium	Chile	Branched, hispid, compressible,	Axinellid skeleton, fibre and spi-	STY: 770 × 20
( Ridley, 1881 )	elastic, yellowish brown.	cule skeleton.	OX: 300–400 × 15–22
Dragmacidon fibrosum	Patagonia	Massive, lobate, conulose, dermal	Plumoreticulate, fibre and spicule	STY: 630 × 15
( Ridley & Dendy, 1886 )	membrane, oscules small, soft, com-	skeleton, primary fibres, cored
pressible, greyish yellow in spirit.	by spicules terminate in tuffs that
protrude beyond the surface. No
ectosome.

TABLE 4. (Continued)

Species	Distribution	Morphology	Skeleton	Spicules (µm)
Dragmacidon grayi (Wells,	North Carolina, Florida	Massive, punctate, prominent	Plumose primary tracts protrude	STY: 240–300 × 9–15
Wells & Gray, 1960 )	asterose drainage canals around	beyond surface, secondary tracts	OX: 360–460 × 5–10
oscules, oscules 1 mm, stiff, yellow	only near surface.
in alcohol.
Dragmacidon incrustans	Tuvalu	Thickly encrusting, microconulose,	Spicule and fibre skeleton, plu-	STY: 40–200 × 10
( Whitelegge, 1897 )	dermal membrane, punctate, oscules	mose, few secondary tracts, scat-	OX: 350 × 8
2.5 mm, soft, fragile, cream in	tered free spicules, sparse spongin.
formalin.	No ectosome.
Dragmacidon kishinense	Northeast Pacific	Thickly encrusting, lobate, oscules	Weakly plumoreticulate near	STY/OX: 1037 (550–1960) × 18
Austin, Ott, Reiswig, Roma-	≤ 4 mm, asterose drainage canals,	surface, confused interior, no ecto-	(7–33)
gosa & McDaniel, 2013	compressible, tough, orange.	some.
Dragmacidon lunaecharta	West Africa	Massive, hemispherical, smooth,	Spicule and fibre skeleton, loosely	STY: 400 × 14
( Ridley & Dendy, 1886 )	oscules minute, firm, compressible,	reticulate, secondary tracts irregu-	OX: 350 × 13
pale yellow in alcohol.	lar, no ectosome.
Dragmacidon mexicanum	California, Mexican Pacific	Thinly encrusting, oscules 1 mm,	Spicule and fibre skeleton, plu-	STY: 300–400 × 20–24
( De Laubenfels, 1935 )	compressible, fragile, beige.	mose.	OX: 400–560 × 16–27
Dragmacidon mutans (Sarà,	Chile	Flabellate, looks fibrous, friable,	Spicule and fibre skeleton, plu-	STY/OX I: 100–220 × 10–12
1978)	oscules minute.	moreticulate, low spongin.	STY/OX II: 200 × 1–2
Dragmacidon ophisclera De	California, Mexican Pacific	Massive, smooth, dermal membrane,	Spicule and fibre skeleton, con-	STY: 1200 × 25
Laubenfels, 1935	soft, compressible, beige in alcohol.	fused reticulation, perpendicular at	OX: 650 × 23
surface.	TRD: present;
RHP: often sinuous
Dragmacidon oxeon (Dick-	Galápagos, California, Mexican	Massive with horn-like protuber-	Unknown.	STY/OX: 800 (600–1150) ×
inson, 1945)	Pacific	ances, detachable dermal membrane,	30–50
beige in alcohol.

TABLE 4. (Continued)

Species	Distribution	Morphology	Skeleton	Spicules (µm)
Dragmacidon reticulatum	Caribbean, Bermuda, Florida,	Thickly encrusting to massive, lo-	Plumoreticulate, multispicular	STY: 250–540 × 8–23
( Ridley & Dendy, 1886 )	Brazil	bate, rugose, microconulose, hispid,	dense primary tracts, spicules	OX: 219–520 × 8–20
dermal membrane a reticulation	protrude beyond surface, irregular,
of fibrous tissue, oscules 2–5 mm,	confused secondary tracts.
firm, barely compressible, abundant
mucus, bright red-orange.
Dragmacidon sanguineum	Natal	Massive, microhispid, numerous	Plumoreticulate, multispicular	STY: 140 × 7
( Burton, 1933 )	oscules 1–2 mm, firm, compressible,	primary tracts, unispicular second-	OX: 211 × 11
dark red dried.	ary tracts, abundant spongin, no
ectosome.
Dragmacidon tuberosum	Cape Verde	Massive, oscules 2–3 mm, punctate,	Plumoreticulate, multispicular	STY: 315–420 × 14–24
Topsent, 1928	finely hispid, firm.	primary tracts of styles, unispicu-	OX: 370–420 ×15–24
lar secondary tracts of oxeas, scat-	TRD: present.
tered free spicules, sparse spongin,
no ectosome.
Dragmacidon tumidum	Australia	Massive, lobose, rugose, oscules	Loosely reticulate, multispicu-	STY: 180 × 6
( Dendy, 1897 )	abundant on margins, very small,	lar primary tracts terminate in
very compressible, greenish-yellow	brushes, abundant free spicules, no
in spirit.	spongin.

* described as oxystrongyles. ** combined into a single category FIGURE 12. Dragmacidon raeae sp. nov. , holotype MCCDRS9411: A. In situ . B. Vertical cross section showing the irregular plumoreticulate skeleton. C. Styles. Only one species of Dragmacidon has been previously described from the Galápagos Islands— D. oxeon ( Dickinson, 1945 ) . Dragmacidon oxeon differs considerably from D. raeae sp. nov. by morphology (it is covered with “hornlike protuberances”) and by the possession of much longer oxeas and styles. Three other species of Dragmacidon have been described from the tropical/subtropical eastern Pacific: D. mexicanum ( De Laubenfels, 1935 ) and D. ophisclera De Laubenfels, 1935 from California, and D. mutans ( Sarà, 1978 ) from Chile . All three species differ from D. raeae sp. nov. by the possession of both oxeas and styles ( Table 4 ). A new, unnamed species of Dragmacidon was described from the shallow waters of the Colombian Pacific by Lizarazo Rodríguez et al. (2020) . This specimen matches D. raeae sp. nov. in colour, texture, morphology, size of styles 302 (152–374) × 19 (9–26) µm, and skeletal morphology. We propose that this specimen is also D. raeae sp. nov. , extending the range of the species to include the Colombian Pacific.