Review of the sponge genus Penares (Demospongiae, Tetractinellida, Astrophorina) in the New Zealand EEZ, with descriptions of new species Author Sim-Smith, Carina Author Kelly, Michelle text Zootaxa 2019 2019-07-15 4638 1 1 56 journal article 26217 10.11646/zootaxa.4638.1.1 fb8c670e-e341-4236-9b05-402c7f92b8f1 1175-5326 3335260 E5A26EB4-1F98-4310-A8D7-A0F933E75D95

TABLE 12. Comparison of Penares species with respect to morphological characteristics and spicule dimensions, and revision of species names with respect to gender. Spicule dimensions are given as mean (minimum–maximum) length × mean (minimum–maximum) width (where available). OX = oxea; SOX = strongyloxea; TYLS = tylostyle; TYLST = tylostrongyle; STY = style; P = plagiotriaene rhabdome; D = dichotriaene rhabdome; OR: orthotriaene rhabdome; MOX = microxea; MR: microrhabd; O: oxyaster; TYL = tylaster; SP = spheraster; SPH = spherule; SA = strongylaster; A = aspidaster. Roman numerals (I, II) are used to differentiate between different size classes of the same spicule type.

Current name	New name	Distribution	Morphology	Skeleton	Megascleres (µm)	Microscleres (µm)
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Peneres helleri		Northeast	Irregularly massive, 20 cm;	Cortex dense crust of tangential	OX: 1300–1400 ×	MOX: 32–150 × 6
(Schmidt, 1864)		Atlantic;	smooth; oscules present;	microrhabds; oxeas tangential	20–40	O: 10–30
type		Mediterranean,	white to dark brown	in endocortex, loosely radial in	D: 350–450
		2–89 m		choanosome; triaene cladomes
				at surface
Penares alata	Penares alatus	South Africa,	Massive, lobed; 13 × 6 cm	Oxeas paratangential at the	OX: 1700–2050 ×	MOX: 16 – 140 × 2 – 7
(Lendenfeld, 1907)		84 m		surface, loosely radial in	50–60
				choanosome; triaene cladomes	ST: 1100–1800 × 60
				at surface	P: 130–1100
Penares aniosoxia		Brazil, 50 m	Spherical; 6 cm diameter;	Cortex separated from	OXI: 200–3500 × 15–28	MOX: 45–90 × 2–3
Boury-Esnault,			osculum, 10 mm, at apex;	choanosome by subdermal	OXII: 1100–1500 ×	O: 12–19
1973			surface covered with pebbles;	canals; choanosome is radial	20–45
			brown		ORI: 1280–2000
					ORII: 340–1060
Penares		Galapagos	Massive, lobed; 16 × 9 × 10	Cortex is a dense crust of	OX: 2496 (1426–3130)	MOX: 127 (68–169) × 7
apicospinatus		Islands	cm; smooth, oscules present	microrhabds and microxeas,	× 39 (21–48)	(4–9); microspined tips
Desqueyroux-			with raised rims; brown,	300–400 µm; palisade of triaene	OR: 1122 (712–1650)	MRI: 91 (67–120)
Faúndez & Soest,			interior orange-brown	cladomes below cortex; triaenes		MRII: 27 (19–42);
1997				and oxeas radial in choanosome		OI: 27 (17–37)
						OII: 9 (6–13)
Penares astronavis		New Zealand,	Thinly encrusting, 3 × 1 cm ×	Cortex is dense crust of	OX: 1269 (768–1692) ×	MR: 40 (16–82) × 13
sp. nov.		980 m	0.2 cm thick; greyish brown	microxeas; triaene cladomes	21 (13–27)	(6–20)
			in ethanol	below cortex; oxeas	D: 180 (117–255)	O: 54 (39–85)
				disorganised; oxyasters are
				abundant in choanosome.
Penares aureus sp.		New Zealand,	Ridged, massive, 30 × 8 cm;	Crust of tangential microxeas	OX: 945 (573–1330) ×	MOX: 137 (70–230) ×
nov.		3–88 m	tan to charcoal grey; smooth;	and microrhabds in cortex;	21 (10–37)	7 (4–17)
			oscules 1–4 mm	triaene cladomes below cortex;	P: 395 (209–618)	MR: 38 (7–79) × 5
				oxeas are radial near the	D: 457 (355–533)	(2–9)
				surface, disorganised in the		O: 17 (8–42)
				interior
Penares candidata	Penares	Northeast	Encrusting; lobed surface; 3	Crust of microxeas in cortex	OX: 450–500 × 10	MOX: 50–100 × 4–5
(Schmidt, 1868)	candidatus	Atlantic;	cm diameter		D: short, length
		Mediterranean,			unknown;
		6–100 m
Penares chelotropa		Brazil, 42 m	2 cm; numerous pores; brown	Crust of tangential microxeas	OXI: 500–620 × 8–12	MOX: 50–65 × 1–3
Boury-Esnault,				and oxeas in cortex; triaene	OXII: 145–250 × 3–8	TYL: 9–19
1973				cladomes below cortex; oxeas	P: 160–250; rare
				disorganised in choanosome
Penares cortius		California; Costa	Irregularly massive, lobed; 12		OX: 950 × 22	MR: 50–160
de Laubenfels, de		Rica	× 7 × 6 cm; oscules on side of		D: 400	O: 9–25
1930			lobes; beige
Penares deformis		New Zealand,	Thinly encrusting; 3 × 3	Crust of microxeas; triaene	SOX: 1094 (656–1498)	MOX: 128 (92–144) ×
sp. nov.		1077–1408 m	cm; oscules 1 mm; brown in	cladomes at surface of sponge;	× 29 (21–40)	8 (7–10)
			ethanol	oxeas are disorganised;	P: 290 (64–489)	MR: 62 (32–108) × 7
				choanosome is densely packed	D: 332 (177–453);	(5–9)
				with oxyasters		O: 40 (22–78)
Penares dendyi		Indonesia, 16 m	Thinly encrusting; 5 × 0.1 cm	Crust of microxeas and oxeas in	O: 168–400 × 12–17	MOX: 56–88
(Hentschel, 1912)			thick; smooth; oscules 0.25	cortex; single layer of triaenes,	D: 200–240	SPH: 8–10
			mm diameter	cladomes at surface; oxeas
				radial in choanosome
Penares euastrum		Northeast	Bulbous; oscules 0.5 mmm;	Thin cortex	OX: 2500 × 35	MOXI: 125–180
(Schmidt, 1868)		Atlantic;	brown		D: 400–2000	MOXII: 45–50
		Mediterranean,				O: 7–10
		25–30 m				A: 120–175
Penares		Galapagos	Massive, no oscules	Crust of microxeas in cortex;	OX: 2300 × 70	MR: 35–160
foliaformis		Islands		single layer of triaenes,	D: 900	O: 16–24
(Wilson, 1904)				cladomes at surface; oxeas are
				loosely radial below triaenes
Penares		Caribbean	Mound-shaped;4 × 3 × 2 cm;	Crust of microxeas and oxeas	OX: 411–546 × 6–10	MOXI: 100–170 × 3–5
hartmeyeri			oscules scattered over surface	in cortex; triaene cladomes at	ORI: 180–220	MOXII: 40–60 × 1–3;
(Uliczka, 1929)				surface, also in choanosome	ORII: 100–120	O: 12–15
Penares		Japan	Irregularly massive, nodular	Crust of microxeas in cortex;	OX: 1000 ×36	MOXI: 140 × 7
hilgendorfi				triaene cladomes below cortex	D: 400; many	MOXII: 80 × 3–4
(Thiele, 1898)					malformed	MOXIII: 30
Penares		Korea	Irregularly massive, fistulose;	Crust of microxeas in cortex;	OX: 650–1300 × 9–30	MR: 50–160
hongdoensis (Jeon			8 × 5 × 6 cm; oscules at tips	triaene cladomes below	D: 560–850	O: 20–40
& Sim, 2009)			of fistules; dark grey	cortex; oxeas disorganised in
choanosome
Penares incrustans		Japan	Thickly encrusting; 7 × 5 × 2	Crust of microxeas in cortex;	OX: 500–820 ×10–15	MOX: 40–100 × 2–4
Tanita, 1963			cm; oscules 1–2 mm	triaene cladomes below cortex;	D: 670–710	O: 15–20
				very loose bundles of oxeas
				radiate towards surface
Penares intermedia	Penares	India; Sri Lanka;	Irregularly lobed; smooth;	Crust of tangential microxeas	OX: 1200 × 37	MOX: 180
(Dendy, 1905)	intermedius	East Africa, 183	oscules at tips of lobes; 2 ×	and oxeas in cortex; triaene	D: 370	O: 25
		m	3 cm	cladomes below cortex; oxeas
				in choanosome disorganised
Penares		New Zealand,	Thinly encrusting or	Crust of tangential microxeas;	OX: 815 (562–1107) ×	MOX: 114 (34–220) ×
kermadecensis sp.		358–414 m	branched, 3 × 2 cm	triaene cladomes below cortex;	22 (13–33)	9 (4–17)
nov.				oxeas are tangential near	D: 197 (122–300)
				surface
Penares mastoidea	Penares	Caribbean;	Massive; hispid in places; 20	Crust of microxeas in cortex;	OX: 3000–3500 × 100	MOXI 100–170 × 7–12
(Schmidt, 1880)	mastoideus	Brazil, 153 m	cm; oscules 2 mm with raised	oxeas radial in choanosome	OR: 840–1150	OI: 8–20
			rim; whitish-grey with dark			OII: 35–45
			spots around oscules
Penares metastrosa	Penares	Japan, 286 m	Massive; oscules <1 mm	Crust of microxeas; triaene	OX: 450–820 × 10–24	MOX: 30–160 × 10
(Lebwohl, 1914)	metastrosus			cladomes below cortex; oxeas	D: 200–375	O: 30–50
				are disorganised in choanosome		SPH: 2–10
Penares micraster		New Caledonia,	Massive, 2 × 2 cm; oscules 1	Crust of microxeas in cortex;	OX: 1200–1500 ×	MOXI: 160–200 × 8–10
Lévi 1993		675–700 m	mm; white to orange	triaene cladomes in or under	10–20	MOXII: 60–70 × 5–6
				cortex	D: 250	SP: 6–8
Penares mollis sp.		New Zealand,	Lobed, 7 × 6 × 5 cm; oscules	Crust of microxeas; triaene	OX: 867 (573–1282) ×	MOX: 163 (84–356) ×
nov.		6–28 m	2–6 mm; soft; grey to black	cladomes below cortex; oxeas	17 (9–28)	8 (5–13)
				loosely radial in choanosome	OR: 380 (184–615)	MR: 60 (30–101) × 5
						(3–9)
						TYL: 11 (8–15)
Penares nux (de		Micronesia, 5 m	Massive, lobed; 6 × 8 × 5	Cortex separated from	OX: 830 × 12	MOXI: 100 × 2
Laubenfels, de			cm; oscules 3 mm; abundant	choanosome by subdermal	P: 250	MOXII: 25 × 0.5
1954)			pores; black	canals; crust of microxeas in		O: 15–36
				cortex; triaene cladomes in
				cortex; oxeas loosely radial in
				choanosome
Penares obtusus	Stelletta	South Africa,	Massive, hispid; red to brown	Crust of stronglyasters and	OX: 3500–5000 ×	MOX: 300–1000 × 7–11
Lendenfeld, 1907	obtusus	84 m	in ethanol; 5 × 3 × 3 cm	perpendicular microxeas;	80–100	OXI: 40–80
				centrotylote diactines in	STY: 2000–4500	OXII: 22–35
				choanosome; strong radial	D: 2600–3600	SA: 10–15
				organisation of megascleres
Penares ochracea	Penares	Australia	Irregularly cylindrical, 14 × 9	Crust of microxeas and	OX: 1016 × 25	MOX: 148 × 4
(Carter, 1886)	ochraceus		× 4 cm; oscules present; ochre	microrhabds in cortex; triaene	D: length not given	MR: <21
			yellow	cladomes below cortex
Penares okokewae		New Zealand,	Thickly encrusting, 10 cm ×	Crust of microrhabds, below	OX: 705 (418–909) × 13	MOX: 62 (34–104) × 5
sp. nov.		5 m	1 cm thick; oscules 2–4 mm;	which is a dense layer of	(9–16)	(3–7)
			cream	microxeas; triaene cladomes	D: 307 (213–384)	MR: 15 (6–26) × 3
				below cortex; oxeas are radially		(2–4)
				arranged; choanosome is		O: 32 (25–50)
				densely packed with oxyasters
Penares orbis sp.		New Zealand,	Massive, 4 × 3 × 2 cm;	Crust of tangential microxeas;	OX: 2378 (1117–3533)	MOX: 97 (33–207) × 6
nov.		406–1680 m	oscules 3 mm; cream	spherasters in a band just under	× 50 (16–143)	(3–9)
				the cortex; triaenes are above	TYLS: 12376 (553–	SP: 10 (7–13)
				and below cortex	3293) × 63 (21–123)	SPH: 11 (5–23)
					TYLST: 1997 (1175–
					2688) × 72 (38–110)
					D: 645 (390–984)
Penares		South Africa	Massive; smooth; no oscules;	Crust of tangential microrhabds	OX: 624 × 18	MR: 40–200 × 9
orthotriaena			foreign debris incorporated	in cortex; triaene cladomes	OR: 312; rare	TYL: 8–12
Burton, 1931				below cortex; oxeas
				disorganised in choanosome
Penares		New Caledonia,	Globular, 7 mm diameter; no	Crust of microxeas in cortex;	OX: 3000–4000 × 100	MOXI: 120–270 × 6
palmatoclada		570–610 m	oscules	triaene cladomes in or below	D: 750	MOXII: 50–80 × 3–4
Lévi, 1993				cortex.
Penares saccharis		Galapagos	Massive to encrusting, lobed,	Crust of tangential microxeas in	OX: 4641 (67–900) ×	MOX: 140 (66–170) ×
(de Laubenfels, de		Islands;	6 × 4 × 2 cm; encrusted by	cortex; triaene cladomes below	12–28	3–8
1930)		California	algae; smooth with hispid	cortex; oxeas and triaenes	D: 300	MR: 22–35–50
			patches; oscules present;	loosely radial in choanosome
			mustard yellow
Penares scabiosus		Galapagos	Massive, 2 × 3 × 3 cm;	Crust of microxeas and	OX: 2760–3200 ×	MOX: 75 (44–106) × 6
Desqueyroux-		Islands	smooth	microrhabds in cortex; single	40–80	(4–10)
Faúndez & Soest,				layer of triaene cladomes below	D: 620–1100	MRI: 43–60
1997				cortex; asters abundant below		MRII: 21–23
				triaenes; oxeas disorganised in		MRIII: 15
				choanosome		O: 20–38
Penares schulzei		India; Sri Lanka;	Massive, lobed; 5 × 5 × 6	Crust of microxeas in	OX: 976 (658–1171) ×	MOX: 109 (41–274) ×
(Dendy, 1905)		New Caledonia;	cm; oscules <3mm at tips of	cortex; oxeas disorganised in	32 (23–45)	8 (4–16)
		New Zealand,	lobes; purple	choanosome	D: 160 (127–183)	O: 19 (12–29)
		106–430 m
Penares sclerobesa	Penares	Azores;	Thickly encrusting, lobed;	Oxeas and triaenes project well	OX:2000–7000 × 50–70
Topsent, 1904	sclerobesus	northeast	hispid in places; oscules	beyond surface	D: 100–115
		Atlantic, 1022–	present
		1165 m
Penares sineastra		Suriname; South	Branching; 5 × 1 cm thick; no	Crust of microxeas forming	OXI: 700–1263	MOX: 42–246 × 3–12
Soest, 2017		America, 100 m	oscules	a dense reticulation around	OXII: 354–700
				apertures; triaene cladomes	D: 321 (282–336)
				below cortex; bundles of oxeas
				present in choanosome
Penares sollasi		Indonesia	Massive, 6 × 5 × 4 cm; black;	Crust of microxeas in cortex;	OX: 1000 × 20	MOXI: 70–90
Thiele, 1900			smooth; oscules 2–3 mm	triaene cladomes below cortex;	D: 250; rarely OR	MOXII: 26–35
				oxeas are radial near the		OXI: 35–40
				surface, disorganised in the		OXII: 12–17
				interior
Penares sphaera		South Africa,	Massive, 8 × 4 × 3 cm;	Crust of microxeas in cortex;	OX: 1250–1700 ×	MOX: 60–185 × 4–11
(Lendenfeld, 1907)		500 m	oscules 4 mm	subdermal canals under cortex;	35–60	SPH: 10–25
				triaene cladomes below cortex;	D: 350–470
				oxeas are radial near the
				surface, disorganised in the
				interior
Penares		New Zealand,	Massive, 12 × 9 ×7 cm;	Crust of tangential microxeas	OX: 1426 (647–2373) ×	MOX: 129 (52–250) ×
turmericolor sp.		350–1680 m	oscules 1–2 mm; yellow	and microrhabds; triaene	38 (11–130)	10 (6–16)
nov.			orange	cladomes below cortex; oxeas	D: 463 (219–807)	MR: 31 (12–78) × 5
				are loosely radial		(3–8)
						O: 19 (6–31)
Penares tylotaster		New Zealand,	Massive, 8 × 4 cm; smooth,	Crust of tangential microxeas	OX: 1212 (978–1404) ×	MOX: 198 (85–287) ×
Dendy, 1924		128 m	oscules 3–4 mm	and microrhabds; triaene	33 (22–45)	14 (8–20)
				cladomes below cortex; oxeas	D: 543 (390–637)	MR: 55 (36–79) × 6
				are radial near the surface,		(4–8)
				disorganised in the interior		TYL: 9 (7–12)
Penares		New Zealand,	Thickly encrusting to	Crust of microxeas in cortex;	OX: 835 (580–1081) ×	MOX: 116 (29–299) ×
vermiculatus sp.		1–75 m	massive, 30 × 10 cm; oscules	oxeas are loosely radial;	21 (10–36)	6 (3–15)
nov.			2–4 mm; yellow brown	triaenes are uncommon	P: 215 (159–340)	O: 19 (9–28)
					D: 278 (208–391)