Craniella quirimure, Peixinho, Solange, Cosme, Bruno & Hajdu, Eduardo, 2005

Peixinho, Solange, Cosme, Bruno & Hajdu, Eduardo, 2005, Craniella quirimure sp. nov. from the mangroves of Bahia (Brazil) (Tetillidae, Spirophorida, Demospongiae), Zootaxa 1036, pp. 31-42 : 33-40

Craniella quirimure sp. nov.

Figs. 2 , 3, 4

Holotype — UFBA 1812­POR, (1) Barra do Jacuruna (ca. 13o02.857’S 38o48.857’W, Itaparica Channel, Jaguaripe, State of Bahia, Brazil) intertidal, mangrove, coll. S. Peixinho, i.1976.

Paratypes (1) Barra do Jacuruna (ca. 13o02.857’S 38o48.857’W, Itaparica Channel, Jaguaripe, State of Bahia, Brazil) intertidal, mangrove — UFBA 212­POR 15 specimens), coll. A. Sampaio, 02.xi.1975; UFBA 222­POR (7 specimens), coll. S.Peixinho, 24.viii.1975; UFBA 252­POR (9 specimens), coll. A. Sampaio, iv.1976; UFBA 254­POR (3 specimens), coll. S. Peixinho, i.1976; UFBA 262­POR, UFBA 161­ PGR, Coll. S.

Peixinho, 1976 (49 specimens), UFBA 1811­POR, coll. S. Peixinho, 1976; MNRJ 601 (1 specimen), coll. E. Hajdu, S. Peixinho & R. Desqueyroux­Faúndez, 22.i.1997; UFBA 1776­POR (8 specimens), coll. S. Peixinho, E. Hajdu, R.G.S. Berlinck, C.P. Santos & E.L. Esteves, 05.vi.2004; MNRJ 8417 (5 specimens), coll. S. Peixinho, E. Hajdu, R.G.S. Berlinck, C.P. Santos & E.L. Esteves, 05.vi.2004. (2) Itaparica Marina (S12º53,340' / W38º41,075', Itaparica Island, State of Bahia, Brazil) ­ UFBA 1888­POR (2 specimens), coll. B. Cosme, 30.i.2005. (3) Maraú River Delta (S13º58´/ W38º59´, Bay of Camamu, State of Bahia, Brazil), mangrove ­ UFBA 999­POR (3 specimens), coll. M.C. Guerrazzi et coll., leg. U.S. Pinheiro, 13.vi.2004.

Diagnosis

Craniella quirimure sp.nov. is the only Craniella in the Tropical Atlantic with cortical oxeas up to 1000 µm long and 28 µm thick, two categories of protriaenes, anatriaenes which can be larger than 9000 µm and sigmaspires smaller than 15 µm.

Specimens Oxeas Protriaenes Anatriaenes Sigmaspires

Holotype I. main choanosomal I. large rabdome rabdome 6,2­8,8 ­ 12,8 UFBA 0 1812 ­ 1671­2257 ­4551/9,9­21,1­28 1434­2148 ­3156/4,2­5,1­6,9>9600/4,1­6,2­8,6 POR II. cortical large cladi cladome

574­839,7­1166/14,5­18,7­28 21,4­40,4­48,8 18­25,6­31

III. anisoxeas large cladome 13­18,3­26

407­675,3­960/3,6­6,3­11,1 II. small rabdome

IV. rhizoidal 210­492­864/1­1,3­1,6

419­569­727/7­11,8­17 small cladi

25,3­46,4­64,7

small cladome 8­20,6­44

Paratype I. main choanosomal I. large rabdome Rabdome 7.7­9.8­11.6 UFBA 0 0 262 ­ 756.1­1877­2343.3/9.9­20.8­28.2 1497.5­1694.6­1882.8/4.2­5.9­7.7>9000/4.4­5.9­7.6 POR II. cortical large cladi cladome

597.9­801.8­1041/9.9­18.4­33.3 29.6­36.4­42 21.7­28.2­33.2

III. anisoxeas large cladome 12.9­19.1­23.8

463.4­603.1­758.7/4.9­6.5­15.5 II. small rabdome

IV. rhizoidal 368.8­494.7­798.1/0.9­1.4­2.3

441.4­699.7­1124.8/9.7­15.4­28 small cladi

14.6­38.3­50.1

large cladome 6.2­22.6­33.8

Paratype I. main choanosomal I. large rabdome Rabdome 5.2­8.6­12.1 UFBA 0 0 254 ­ 1598.3­2067.7­2704.9/11.1­18.8­25.5 1433.7­2147.7­3156.5/4.2­5.1­6.9>7000/5.9­6.7­7.7 POR II. cortical large cladi cladome

521.1­842.5­1220.6/14.2­20.7­31.6 21.4­40.3­53.5 24.6­29.9­35.7

III. anisoxeas large cladome 21.1­27.4­33.4

380.5 ­ 594.2­808.3/3.9­5.3­8.5 II. small rabdome

IV. rhizoidal 209.8­491.6­864.2/1­1.3­1.6

520.4­718.2­881/13.8­17.4­25.3 small cladi

25.3­46.4­64.7

large cladome 4.9­7.3­11

Paratype I. main choanosomal I. large rabdome Rabdome 5.8­8.4­12.3 UFBA 0 1811 ­ 1362.8­1718.5­2489.8/7.4­16.3­21.2 1486.5­1713.9­1920.9/5.1­6.3­7.8>6000/4.3­5.9­6.8 POR II. cortical large cladi cladome

493.4­687.3­1019.2/11.8­16.1­24.7 32­37.7­41 24.9­31.3­41

III. anisoxeas large cladome 13.8­18.9­23.5

462.8­540.4­726.3/3.9­6.6­10.7 II. small rabdome

IV. rhizoidal 212.4­362.4­514.9/1.1­1.7­2.1

269­466.2­637.2/2.9­11.2­16.2 small cladi

331­36.7­42.8

large cladome 10.3­15.1­22.5

Description

External form irregularly ovoid, elliptical, barrel­shaped or nearly spherical. Specimens reach a maximum of 23mm in diameter and 25mm in height. Surface can be flat (more often) or slightly ridged, with or without perforations (inhalant apertures). Some specimens were seen to bear a crown of conulose­volcaniform projections on the rim of their apical side. Texture normally smooth. Consistency varies from easily compressible to nearly incompressible, what might be related to contraction after collection and handling. More often a single apical oscule occurs, also markedly contracted after collection. A few specimens possessed two or three oscula clustered. Aquiferous channels are clearly seen converging to the oscula in live specimens. In many preserved specimens oscula are not visible. The basal portion of the specimens bears rhizoidal extensions for anchoring in the soft sediment. These are variably intertwined, and can reach 73 mm (UFBA1812­POR) in length. In some specimens (ovoid and barrel­shaped) these extensions clearly project from several locations spread around the base of specimens. In others (elliptical), they appear to be more concentrated. In live specimens these extensions were either buried in the sediment, or entangled under the specimen, penetrating over 1­2 cm in the sediment. Live color is muddy­greenish and muddy­yellowish externally, turning brown in ethanol. Sectioned specimens are yellow inside, clearer on the central portion due to increased silica content, and have two conspicuous outer layers of distinct color. The outermost is sediment colored, reddish­brown, and is adjacent to a bright green sub­surface layer.

Skeleton

Nearly perfect radial arrangement of conspicuous spicule tracts of larger choanossomal oxeas spiraling from the base towards the surface of the sponge (ca. 3 tracts in 1 cm of surface area), terminating within a palisade­like layer (ca. 1080 mm thick) of smaller cortical oxeas adjacent to, and slightly piercing (ca. 40 µm) the surface. The radial arrangement is somewhat obscured by a dense criss­crossed reticulation of isolated spicules. Aquiferous channels are seen here and there in the choanosome (up to 280 µm across). Protriaenes are piercing the surface abundantly as commonly seen on other tetillids. Sigmaspires are distributed all over the sponge. The basal rhizoidal extensions have abundant intertwined anchoring anatriaenes.

Spicules (measures in m): Megascleres Protriaenes I, associated to the main cortical/ choanossomal tracts, but easier to spot in the cortical region; eventually of prodiaene form; slender. Rhabdome, long, straight, sometimes with a sinuous base, always tapering gradualy; 1434­1925.9­3156/4.2­5.6­ 7.8 m. Cladome, straight and sharp, ocasionally asymetrical; 21­38.7­54/13­20.8 ­ 33 m.

Protrianes II, associated to the main cortical/choanossomal tracts, but easier to spot in the cortical region; eventually of prodiaene form; slender. Rhabdome, long, straight, sometimes with a sinuous base, always tapering gradualy; 210­473.1­908/0.8­1.4­ 2.2 m. Cladome, slender, straight or curved, and sharp, ocasionally asymetrical; 15­41.9­65/4.9­ 16.4­ 44 m.

Anatriaenes, rhizoidal, forming long tracts embedded in the soft sediment, the cladomes acting as hooks. Extremely long rhabdome, tapering gradually to a sharp end (like a long hair);>6000/4­6.1­ 8.5 m. Cladome, straight and sharp 17.6­28.7­ 41 m.

Oxeas I, main spicule of the choanosomal bundles; long, straight, stout in the central region and sharpening abruptly, 1363­1928.1­2705/7­17.5­ 26 m.

Oxeas II, cortical; short and robust, straight, sometimes curved, abruptly sharpened, ocasionally styloid, 493­764.8­1221/12­18.3­ 32 m.

Oxeas III, auxiliary spicules of the choanosomal bundles; anisoxeas, one end tapering gradually, the other abruptly, 381­567.2­808/3.9­5.9­ 11 m.

Oxeas IV, rhizoidal; similar to oxeas I, but shorter, 269­592.2­881/5.9­14.3­ 25 m.

Microscleres Sigmaspires, occurring both in the choanosome as well as in the rhizoidal extensions; small and acanthose, 5.7­9.0­ 12 m.

Ecology and distribution

The species is abundant at least on part of the intertidal mangrove area studied (Barra do Jacuruna), where it is found on muddy­sand substrate attached by its rhizoidal projections. Observed density was about 12 individuals per square meter. Clusters had at most 2­3 individuals. Water temperature may reach above 30ºC in summer months (December­February), and emmersion is calculated to be of no more than 1­2h on the lowest tydes. This is a delta area where salinity has been reported to vary from 3 to 31/ 1000, with an annual mean around 25/1000 ( Paredes et al., 1980). Other sponges occurring in the area are Mycale microsigmatosa Arndt, 1927 and Haliclona coerulea (Hechtel, 1965) . The Itaparica Marina is situated in a marine environment.

The Bay of Camamu is the target of an ongoing basic inventory of its benthic fauna (M.C. Guerrazzi et coll.). The species was seen on different locations in the bay, at varying distances from the entrance ( U.S. Pinheiro, pers. comm.), which suggests considerable adaptation to salinity fluctuations (not recorded).

Etymology

The species name, a name in apposition, is the Tupinambás Indians name for Todos os Santos Bay, its type locality, and where most of the specimens collected stem from.

Remarks

The very disparate sampling effort undertaken on the three collecting localities precludes any discussion on possible morphologic trends established on these populations. New collections on the Bay of Camamu and on Itaparica Island will allow a study of variability, which is beyond the scope of this contribution.

There are nearly 40 species of Craniella reported worldwide, seven of which from the Tropical Atlantic, viz.: C. carteri Sollas, 1886 ; C. cranium ( Müller, 1789) ; C. insidiosa Schmidt, 1870 ; C. laminaris ( George & Wilson, 1919) ; C. lens Schmidt, 1870 ; C. schmidti Sollas, 1888 and C. tethyoides Schmidt, 1870 .

The new species described here differs from C. carteri and C. lens because both latter species do not possess sigmaspires. Additionally, none of these were reported to possess two categories of protriaenes, and the main choanosomal oxeas of both seem to be always much stouter (always over 20ìm thick) than those of the new species (sometimes as small as 7 m thick). Of those Tropical Western Atlantic Craniella species with sigmaspires, C. schmidti and C. tethyoides have these microscleres much larger (respectively ca. 20 and 35 m in largest diameter). Both species are deep­water inhabitants being thus well differentiated on ecological grounds too, when compared to the new species apparent shallow, warm, mangrove restricted distribution. One further diagnostic character stems from C. schmidti s much stouter choanosomal oxeas.

Craniella cranium is a problematic species in need of revision. Originally described from the Boreal Eastern Atlantic, the species has subsequently been reported from several other, mostly Boreal/Temperate localities, on many distinct biogeographic provinces. On an exclusively biogeographical probability rationale, Boreal records on both the Atlantic and the Pacific are likely, Southern Temperate records and a postulated anti­tropical distribution are less likely, and Tropical records and their implied cosmopolitanism are much less likely. With this in mind, it is very improbable on biogeographical grounds alone that C. cranium will be found on warm, shallow Tropical Brazil. Consequently, Carters (1890) record of C. cranium for the Fernando de Noronha Arquipelago, despite not revised here, is most probably equivocal. We have taken Van Soest et alls (2000) description as a recent revision of Boreal Western Atlantic C. cranium , and used it for a comparison with C. quirimure sp.nov. The Boreal species has much stouter cortical (up to 55 m thick) as well as choanosomal oxeas (up to 70 m thick). Its triaenes, both pro­ and ana­, were reported to be much longer than those in the new species (respectively, up to 8500ìm and 20000ìm). Sigmaspires in C. cranium may reach 20 m in diameter while in C. quirimure sp.nov. these are smaller than 15ìm. Given all these morphological differences, and keeping in mind both species apparent very low biogeographic relatedness, C. quirimure sp.nov. is considered very well differentiated from C. cranium .

The two remaining Craniella reported from the Tropical (or nearly so) Atlantic are C. insidiosa and C. laminaris . The first one was poorly described by Schmidt (1870) and was considered unrecognizable already by Sollas (1888). No further comments can be made here on it status. C. laminaris , in spite of originally described from Newport River (eastern U.S.A.), a locality nearly 9000 km away from Bahia, appears to be the morphologically closest to the new species. Both share the very shallow water habitat, as well as comparably large cortical oxeas and sigmaspires, and possess additional categories of protrianes (cf. George & Wilson, 1919). A series of traits can be used to separate them. Cortical oxeas can be much stouter (up to 28 m thick) in C. quirimure sp.nov., choanosomal oxeas can be much longer (up to 4551 m) and stouter (up to 28 m thick), and anatriaenes can be over 4x larger (up to 9600 m). The new species is thus considered well distinguished from other congeners in the Tropical west Atlantic.

Figures

FIGURE 2. Craniella quirimure sp. nov. A. Two specimens alive in an aquarium. Scale bar 2 cm. B. Specimen sectioned sagitally to show inner color pattern. Scale bar 2 cm

FIGURE 4 Schematic representation of the megasclere complement and SEM of the microscleres of the holotype. A. Protriaene I. B. Cladome of the prodiaene I. C. Protriaene II. D. Cladome of the prodiaene II. E. Oxea I (main, choanosomal). F. Oxea II (cortical). G. Oxea III (anisoxea, choanosomal). H. Oxea IV (rhizoidal). I. Anatriaene. J. Sigmaspire. Scale bar in m: A­D, I = 25; E = 250; F­H = 100 and J = 2.