Timea clandestina, Leite, Dora M. B., Fonseca, Cássio A., Leal, Camille V. & Hajdu, Eduardo, 2015

Leite, Dora M. B., Fonseca, Cássio A., Leal, Camille V. & Hajdu, Eduardo, 2015, Two new species of Timea from the Southwest Atlantic (Timeidae, Demospongiae, Porifera), Zootaxa 4034 (1), pp. 161-181 : 166-178

Timea clandestina sp. nov.

( Fig. 3 ; Tab. 2)

Diagnosis. Finely encrusting Timea with straight or slightly curved (sub)tylostyle megascleres, each with an elongated tyle (310–435 Μm length, 2–10 Μm width, and 5–10 Μm tyle width), and a single but morphological variable category of microsclere, ranging from spherostrongylaster with a crown of thorns on the tip, to spined spheroxyaster measuring 6–20 Μm diameter.

Material examined. Holotype. MNRJ 18413 B, Hípica, Ilha do Papagaio (22°53'54.89"S - 41°58'41.95"W, Cabo Frio, RJ, Brazil), 11.1m depth, E. Hajdu coll., 0 9 May 2014.

Description. External Morphology. Thinly encrusting sponge, 0.2 mm thick on average. Total area of the specimen ca. 30 cm ². Color after fixation grayish beige ( Fig. 3 a). Surface microhispid ( Fig. 3 b). Oscula and ostia not detected; however, numerous small depressions on the surface were observed. The specimen was collected inadvertently, attached to the underside of the same pebble containing the holotype of T. berlincki sp.nov. (described above), and as such no in situ photo or observations were made.

Skeleton. Ectosomal skeleton a crust of asters. Choanosomal skeleton with dispersed asters and subtylostyles organized in bouquets piercing the ectosome and producing hispidation. Subtylostyles may also be found tangential to the substrate ( Fig. 3 c).

Spicules. Megascleres. (Sub)tylostyles, straight or slightly curved, with an elongated tyle and hastate apex ( Fig. 3 d). Microscleres. Spherostrongylasters to tylaster morphologies ( Fig. 3 e–f) with smooth conical rays either possessing a secondary furcation or ending in extremities with distally oriented spines, reminiscent of crowns. Rays rarely cylindrical. Slightly smaller and completely smooth forms were found and are thought to be growth forms of the same category ( Fig. 3 e, Tab. 1 ).

Ecology. Found in the sciophilous surface of the pebble on top of which the holotype of T. berlincki sp.nov. was collected. Water temperature at the time of collection was 21ºC.

Distribution. Known only from its type locality at Ilha dos Papagaios (Cabo Frio, RJ, Brazil), 11 m depth (fig. 1).

Etymology. The specific epithet is used as a noun in apposition. “ Clandestina ” refers to the clandestine, stealthy way by which this sponge appeared in the collecting bag where only the holotype of T. berlincki sp.nov. was supposed to have been collected.

Remarks. Table 2 lists many Timea spp. that appear only distantly related to the new species given obvious characters such as much larger, much smaller, or much stouter megascleres. There is nevertheless a group of 14 species that obliges a comparative analysis of microscleres in order to show their non-cospecificity to the new species. Thirteen of these still possess important distinctive features on the basis of categories of asters present, their diameter and micromorphology. These iclude T. anthastra , T. biphidostellata , T. crassa , T. curvistellifera , T. hallezi var. crassa , T. mixta , T. ornata , T. oxyasterina , T. parasitica , T. stellifasiata , T. stellivarians , T. trigonostellata and T. tristellata .

A few species need a more thorough comparison. For instance, judging from its published description alone, T. centrifera differs from T. clandestina sp.nov. in the cylindrical rays and much smaller centri of its asters, as opposed to conical rays and large centri in the latter. Both species’ far apart occurrences add up on the improbability of cospecificity. Timea unistellata , on the other hand, is readily distinguished by the orange and red colors reported for specimens alive. In general, asters were reported to be bigger, reaching 25 µm in diameter in Topsent’s (1892) original description, 28 µm in Boury-Esnault & Lopes’s (1985) and 30 in Cruz’s (2002) description. Nonetheless, it may actually be the 6–7 µm exhibited by the smaller asters in the new species that better set it off T. unistellata ’s morphospace. Similarly to Timea unistellata , T. lowchoyi can also be distinguished by its dark color alive (red-brown), and asters that reach larger dimensions (28 µm), but do not reach the smallest diameters found in the new species. In addition, this further Australian species was described by Hooper (1986) with the typical pore-grooves that are so conspicuous to several species, among which T. ohuirae and T. berlincki sp. nov. (cf. above), but seem to be entirely absent from T. clandestina sp.nov. The latter is thus also deemed confidently set apart from its congeners and deserves the proposed status of new species.

TABLE 2. Comparative micrometric data on the spicules and overview of distribution of the known species of Timea . Values are in micrometres. Abbreviations are as follow: L, length; W, width; TW, tyle width, D, diameter; r, ray number

Spicules Complementary information Distribution

Species

Megascleres Microscleres

Timea berlincki sp. nov. * Subtylostyles with irregular Tylasters Color in life is orange-brown; after fixation Type locality: Cabo Frio (Rio de Holotype (MNRJ 18413a) elongated tyles. D: 8.5–12.3–15.3; r: 9–12 in ethanol, color is brownish beige. Janeiro, Brazil)

L: 204–454–630; W: 2.4–7.0–9.6; Spheroxyasters Additional records: Ilhabela (São TW: 4.8–8.0–9.6 D: 9.4–11.0– 13.4; r: 11–14 Paulo, Brazil)

Timea clandestina sp. nov. * Subtylostyles to tylostyles with an Spherostrongylasters After fixation, grayish beige color and Cabo Frio (Rio de Janeiro, Brazil) Holotype (MNRJ 18413b) elongated tyle. D: 6.0–12.3–20.4; r: 14–18 hispid surface. L: 310–382–436; W: 2.4–4.8–7.2;

TW: 4.8–6.9–9.6

Timea alba Bergquist, 1968 Tylostyles , may be somewhat Spherostrongylasters Color in spirit gray. New Zealand orig. descr.) irregular and bi- or 4-lobed. D: 34.8–44.6–49.2; r: 7–14, up L: 2970–3659–4018; W: 15.8– to 5.7 (width)

18.2–20.6; TW: 22–30.8

Timea anthastra Lévi, 1961 Tylostyles , elliptical tyle. Tylasters Purple-pink on the outside and greenish Seychelles orig. descr.) L: 350–400; W: 8–9; TW: 13–15 D: 12–14; r: 6–8 inside. There is a single orifice at the

summit of the projection.

Timea aspera ( Topsent, 1904) Tylostyles , marked elliptical tyle. Spherasters Gray at surface to dark yellow inside. Azores ( Portugal) orig. descr., as T. unistellata L: 660–1000; W: 12–20; TW: 5–12 D: 20–25

. a.)

Timea aurantiaca Bergquist, Tylostyles , oval tyle. Frequently Tylospherasters to Color in life, bright orange to red; gray in New Zealand 1968 (orig. descr.) subterminal. strongylospherasters to spirit. sciophilous.

Specimen 1 spherasters

L: 193–480–677; W: 2.3–4.8–6.0 Specimen 1

Specimen 2 D: 4.6–13.9–20

L: 217–340–400; W: 2.3–3.7–4.0 Specimen 2

D: 5.7–11.5–22.2; r: 8–20

……continued on the next page Species Spicules Complementary information Distribution

Megascleres Microscleres

Timea authia de Laubenfels, Tylostyles, oval tyle. Tylasters Shape massive, encrusting. Orange in life, Southern California ( Mexico) (orig. descr.) Tylostyles D: 23–6; r: <10–30 drab when preserved.

L: 700; W: 10

Style

L: 200–840; W: 4–11

holotype reevaluated by Tylostyles, spherical to oval tyle, Strongylasters to oxyasters Color in life is yellowish orange, and beige Southern California ( Mexico) Carballo & Cruz-Barraza sometimes malformed. D: 3–14.7– 24 in alcohol. Surface hispid. 2006) Tylostyles

L: 207–477.5–915; W: 3.8–6.4–15;

TW: 3–8.2–15

Styles

L: 187–289.4–800; W: 2.5–3.8–5

Timea bifidostellata Pulitzer- Subtylostyles Spherasters Colorless according to field notes. Italy Finali, 1983 (orig. descr.) L: 230–600; W: 4.5–9.5 D: 11–16

Timea bioxyasterina Mothes Tylostyles , spherical to ovoid tyle, Oxyasters 1 Light-brown colored in ethanol. Surface Off Maranhão ( Brazil)

al., 2004 * (orig. descr.) sometimes bilobed. D: 32–62–85.8; r: 9–11 smooth. L: 304–579.5–769.5; W: 6.3–10.5– Oxyasters 2

15; TW: 2.5–12.3–17.5 D: 16.1–20.7–27.6; r: 9–11

Tylasters

D: 2.4–3.6–5; r: 9–10

Timea capitatostellifera Tylostyles with pronounced oval tyles. Spherotylasters Snow-white color. Gulf of Manaar ( India) Carter, 1880) (orig. descr., as L: 1128; W: 14 D: 50.8

Hymedesmia c.)

……continued on the next page TABLE 2. (Continued)

Species Spicules

Complementary information Distribution Megascleres Microscleres

Timea clippertoni Van Soest Tylostyles, subterminal tyle, Calthrops-like aster Pale beige in alcohol. Clipperton Isl. ( France) al., 2011 (orig. descr.) possibly in two categories, but with D: 21–42.2–78; r: 3–6

intermediates. Spherostrongylasters

L: 119–366.0–963; W: 2–4.6–9; D: 9–15.7–19; r: 8–16

TW: 3–6.8–11 Oxyspherasters

D: 5–7.1–9; r: 8–12

Timea crassa ( Topsent, 1900) Tylostyles to subtylostyles. Spherasters, sometimes Color pale yellow, sometimes yellowish Type locality: France (North Sea) orig. descr., as Hymedesmia malformed beige. Additional records: Azores ( Portugal), hallezi var. c.) D: 15 Canary Islands, France

(Mediterranean), Italy, Morroco, Senegal

sensu Topsent (1925) Tylostyles Spheroxyasters Color reddish. France (Mediterranean)

L: 245–875; W: 3–10 D: 15–25

Strongylasters

D: 8

sensu Lévi (1952) Tylostyles, straight, sharpening Strongylasters Color orangey. Senegal

gradually, regular round head. D: 6–22

L: 280–1000; W: 12

……continued on the next page Species Spicules Complementary information Distribution

Megascleres Microscleres

sensu Lévi (1952, as T. hallezi Tylostyles , ellipsoidal tyle, Spherasters Color yellow or brown. Senegal. crassa ) elongated, often irregular. D: 7–22

L: 200–600; W: 7–12; TW: 10–12

sensu Pulitzer-Finali (1983) Tylostyles, round tyle, not very Tylasters 1 Color orange. Italy

developed. D: 7–8

L: 580–990 Tylasters 2

W: 7–14 D: 12–22

Timea cumana Pulitzer-Finali, Tylostyles , round tyle. Strongylasters, variable Color dull yellow. Italy (orig. descr.) L: 210–1600; W: 4–20 centrum

D: 6–15; r: 8–13

Calthrops-like aster

D: 16–27; r: 4–6

Timea curacaoensis van Soest, Tylostyles, elongate tyles, often Branched asters Live color not noted, white in alcohol. Curaçao * (orig. descr.) style-like or subterminal. D: 14–19.2–23; r: 4–5

L: 299–834.2–1357; W: 2–8.2–14 Tylasters

D: 5.5–6.0–7.5; r: 8–9

Timea diplasterina Rützler et Tylostyles , spherical tyle. It may be Spheroxyasters 1 Live color ranging from drab to dull orange Belize, 2014 * (orig. descr.) subterminal. D: 38–48–59 brown.

Type 1 Spheroxyasters 2

L: 325–700; W: 14–23 D: 8–17–30

Type 2 Diplasters (spirasters-like):

L: 129–350; W: 5–13 D: 13–20–27

……continued on the next page Species Spicules Complementaryinformation Distribution

Megascleres Microscleres Timea floridusa Carballo & Tylostyles to subtylostyles, Tylasters 1 Ochre, light brown or light orange, in life Mexican (Pacific Ocean) Cruz-Barraza, 2006 (orig. sometimes subterminal. Number of spines: 4–8 and pale brown in alcohol. descr.) L: 130–420–1120; W: 2.5–5.6–15; r: 6

TW: 2.5–6.7–16.3 Tylasters 2

D: 8–12–17

Timea geministellata Pulitzer- Tylostyles, elongated tyle, often Strongylasters to oxyasters Italy Finali, 1978 (orig. descr.) trilobated and sometimes subterminal. D: 9–11; r: 7–10

L: 400–650; W: 5–7; TW: 6–8 Branched asters (anfiasters)

D:15; r: 3–4

sensu Voultsiadou & Vafidis Tylostyles, trilobate and sometimes Strongylasters to oxyasters. Greece 2004) subterminal tyle. Branched asters

L: 190–750 D: 6.8–20 (both categories)

Timea granulata Bergquist, Tylostyles , round or subterminal Tylasters 1 Color in alcohol, pale brown. Republic of Palau 1965 (orig. descr.) ovate tyle. D: 13–14.8–16.5

L: 170–401–687; W: 1.5–4–8 Tylasters 2 (rare)

D: 4–4.8–5.8

sensu Uriz (1988) Tylostyles, round tyle, ovoid, or Spherasters (massive centrum) Color greenish brown in alcohol.

subterminal swelling. D: 5–12

L: 204–650; W: 4–7

……continued on the next page TABLE 2. (Continued)

Species Spicules Complementaryinformation Distribution

Megascleres Microscleres

Timea hechteli Lehnert & Tylostyles to styles-like variation. Oxyspherasters Type locality: Florida ( U.S.A., Gulf of Heimler, 2001 (sensu Little, L: 211–569.1–1015; W: 3–7.1–12 D: 18–26.4–33 Mexico) as Halicometes stellata ) Strongylasters Additional records: Barbados

D: 7–12.8–22

Timea intermedia ( Lévi, 1958) (Sub) (tylo)styles; tyle Tylasters Yellow color. No visible pores. A slightly Red Sea orig. descr., as Timeopsis i.) mammiliform/ bulbous ovoid, D: 9–11; r: 6, 4–6 terminal elevated osculum was found in one of the

subterminal or absent. spines. specimens.

Tylostyles

L: 300; W: 7

Subtylostyles

L: 775; W: 10–15

Timea irregularis Sarà & Tylostyles to subtylostyles Spheroxyasters Color yellow ochre, or yellow-brownish Mediterranean Siribelli, 1960 (orig. descr.) L: 252–1020; W: 3–8 D: 24–35

(Spher)oxyasters (branched) D:14–21

(Spher)oxyasters

D:7–10

Timea lophastraea (Hentschel, (Subtylo)styles, irregular and/or Tylasters Color grayish yellow in alcohol. Southwestern Australia) (orig. descr., as subterminal tyle. D: 11–14; Axis: 2–6; r: 4–6 Hymedesmia l.) Subtylostyles

L: 336–696; W: 2–7

Oxeas

L: 68–155; W: 1–2.5

……continued on the next page Timea micraster Lehnert & Tylostyles , round tyle. Spherasters Bright orange colored, still orange in Jamaica Heimler, 2001 * (orig. descr.) L: 325–758; W: 7–10 D: 15–37 ethanol. Spherasters (massive centrum):

D: 2–3

Timea mixta ( Topsent, 1896) Tylostyles. Spherasters Ochre-yellow. Type locality: France (Mediterranean) sensu. Topsent, 1900; as L: 275–>1000; W: 3–9 D: 15–35 (larger ones deeper in Additional records: Canary Islands, Hymedesmia m.) the sponge) Aegean Sea Tylasters

5–6

Spherasters

Tylostyles. 29–40.0–46 (±3.9)

sensu Hechtel (1976) * (YPM L: 310–536–1001 (±171.6) Spherostrongylasters n.r. NortheasternBrazil 8954, new data) W: 2.4–7.8–14.4 (±3.1) 14.4–18.2–21.6 (±2.0)

TW: 4.8–9.0–14.4 (±2.9) Micrasters

2.4–4.0–7.2 (±1.4)

Timea moorei ( Carter, 1880) Tylostyles , spherical tyle. Possibly Spherasters 1, spherasters 2 Color glistening white. Gulf of Manaar ( India) orig. descr., as Hymedesmia also with oxeas. and asters with abruptly pointed

.) L: 875; W: 28.2 ray (all categories)

D: 21.2

Timea ohuirae Carballo & (Sub)tylostyles with generally well- Oxyasters The color in life is pale orange, and beige Mexican (Pacific Ocean) Cruz-Barraza, 2006 (orig. formed spherical or oval tyle. Some D: 16–42; r: 8–14 or light yellow in alcohol. Oscula rose descr.) have malformed tyle and annular Strongylasters slightly in thicker areas of the sponge, or

swellings. D: 15–25; r: 8–13 flush with the surface.

L: 169–990; W: 2.5–17.5; TW: 3.8– Tylasters to strongylasters

20 D: 5–8; r: 6–9

……continued on the next page Species Spicules Complementaryinformation Distribution

Megascleres Microscleres

Timea ornata Lévi & Lévi, Tylostyles , ovoid tyle. Spheroxyasters Philippines (orig. descr.) L: 330–600; W: 5–9 D: 40–55

Timea oxyasterina Rützler et Tylostyles , tyle not well marked, Oxyasters Live color deep red. Numerous circular Belize, 2014 * (orig. descr.) sometimes stylotes or subterminal D: 25–32–37 aquiferous openings. (mucronate).

L: 152–320; W: 3–11

Timea parasitica (Higgin, Tylostyles, large subterminal tyle. Spheroxyasters Bahamas) * (orig. descr., as L: 508; W: 7.3 D:>26

Donatia p.) Tylasters

D: ~13

Timea perastra (de Tylostyles Tylasters of about 12 rays Color yellow in life and pale gray Dry Tortugas (Florida, Gulf of Laubenfels, 1936) * (orig. L: 540–690–1888; W: 7–9–13 each. Another sort of stubby preserved in alcohol. Mexico, USA) descr., as Halicometes p.) armed asters also occurs.

Tylasters

D: 12

Armed Asters

D: 8

Timea secirm Moraes, 2011 * Tylostyles, round or oval tyle, Oxyspherasters Color yellow in life and white preserved in Saint Peter and Saint Paul’s orig. descr.) sometimes subterminal. D: 12–18–28; r: 18 alcohol. Archipelago ( Brazil)

L: 200–429–715; W: 1–4–7 Strongylasters

D: 4–7–11; r: 14

Timea simplistellata Pulitzer- Tylostyles, round, subterminal Spheroxyasters The color is a very light violet. Type locality: Italy Finali, 1983 (orig. descr.) and/or bilobed tyle. D: 27–44; r: 12–14 Additional records: Cape Verde

L: 1600–2500; W: 12–21

……continued on the next page Timea stellata (Bowerbank, Tylostyles, subterminal tyle. Strongylasters Type locality: Guernsey (United 1866) sensu Rützler (2002, Tylostyles 1 D: 8–10 Kingdom)

holotype reevaluated) L: 90–120; W: 2.5–3.5 Tylasters to oxyasters Additional records: Cape Verde, Tylostyles 2 D: 14–22 Mediterranean ( Croatia, France, L: 190–230; W: 2.5–4.5 Greece, Italy, Monaco, Tunisia, Turkey)

sensu Pulitzer-Finali (1983) Tylostyles Strongylasters to rarely Color lemon yellow. Italy

Length: 135–900; Width: 2.5–9 oxyasters

D: 8–13

Timea stellifasciata Sarà & Tylostyles , well-marked tyle, Oxyasters Color brownish or brownish-yellow. Type locality: Italy Siribelli, 1960 (orig. descr.) sometimes trilobed. Sometimes Calthrops-like aster (rare) Additional records: Cape Verde

subterminal. Branched asters

L: 125–595; W: 3–8 D: 7–21 (collectively)

sensu Boury-Esnault (1973) * Tylostyles Tetraradiate asters n.r. NortheasternBrazil MNHN.LBI.NBE 959, own L: 223– 458–944 (±188.2) 9.6–14.6–19.2 (±2.4)

measurements) W: 2.4–5.2–14.4 (±2.5) Strongylaster

TW: 2.4–5.5–12.0 (±2.1) 4.8–7.4–16.8 (±1.6)

Timea stelligera (Carter, Tylostyles, mammiliform or pear- Probably two categories are Color, in its apparently washed-out state, Type locality: Honduras (? Caribbean) 1882) * (orig. descr., as shaped tyle. May be subterminal. present. light grey. Massive, conoidal, lobate, erect, Additional records: Azores (?), Cliona s.) L: 1121; W: 21.2 Spherasters and tylasters somewhat compressed near the base, Seychelles (?)

D: 17; r: 8–10 expanded towards the center.

Timea stellivarians (Carter, Tylostyles, oval tyle. Spherasters Color yellow. Gulf of Manaar ( India) 1880) (orig. descr., as L: 452; W: 14.1 D: 42.3

Hymedesmia s.)

……continued on the next page Species Spicules Complementaryinformation Distribution

Megascleres Microscleres

Timea stenosclera Hechtel, Tylostyles , elongate/oval tyle. Oxyspherasters Color is orange in life, grayish-brown in Type locality: Barbados

* (orig. descr.) L: 173–210–252; W: 1.7–2.1–2.4; D: 11.8–19.7–25.9 alcohol. TW: 3.5–3.6–4.7 Strongylospherasters to

oxyasters

D: 4.7–9.2–11.8

sensu Rützler et al. (2014) Tylostyles, small mostly spherical Spheroxyasters to Yellow-orange to ochre live color with Belize

tyle. spherostrongylasters sinuous exhalant canals. L: 176–287–310; W: 2–5–7 D: 19–22–25

Spheroxyasters (rare)

D: 15–17–25

Timea tethya (de Laubenfels, Styles Oxyspherasters (massive Pale orange color. Hawaii ( USA)) (orig. descr., as Kotimea L: 700; W: 14 centrum, rare)

D: 38

Oxyasters (not illustrated) D: 20

Tylasters

D: 5–7

Timea tethyoides Burton, 1959 Tylostyles , round tyle. Spherasters 1 (choanosomal) Light greyish-brown color after Zanzibar (Tanzania) orig. descr.) L: 400–800; W: 22 D: 80 preservation. Spherasters 2 (ectosomal)

D: 20

Timea tetractis Hentschel, Tylostyles , round tyle. Strongylasters Gray color. Indonesia (Arafura Sea) (orig. descr.) L: 184–520; W: 2–7 D: 7–12; r: 8–13

Calthrops-like asters

D: 15–31; r: 4

……continued on the next page TABLE 2. (Contninued)

Species Spicules Complementaryinformation Distribution

Megascleres Microscleres

Timea trigonostellata (Carter, Tylostyles, small round tyle. Oxeas Calthrops-like asters Snow-white, glistening color. Gulf of Manaar ( India) 1880) (orig. descr., as acuate, long, thin and smooth. D: 12.7 Hymedesmia t.) Tylostyles:

L: 352.8; W: 14.1

Oxeas:

L: 705.5; W: 7

Timea tristellata (Topsent, Tylostyles Triple asters (sic) Brick red color. Gulf of Manaar ( India) 1892) (orig. descr.) L: 370–400 (?) D: 17–20

Timea unistellata (Topsent, Tylostyles Spherasters Salmon-colored. Type locality: France (Mediterranean) 1892) (orig. descr., as L: 370–400 D: 20–25 Additional records: Azores, Canary Hymedesmia u.) Islands, Iceland, Italy, Madagascar (?), Mauritius (?), Monaco, Spain

(Mediterranean), Tunisia

Figures

FIGURE 3. Timea clandestina sp. nov. (MNRJ 18413 B, holotype). (A) Specimen after fixation; (B) Detail of a cross section showing the bouquets formed by tylostyles; (C) Cross section of specimen; (D) Tylostyles; (E) Variation of the spherostrongylasters to tylasters of the same category, including growth forms (gf); (F) Extremes of the variation in the same size category.