New or little known demosponges ( Porifera ) from Espírito Santo coast and seamounts ( Brazil )

Phorbas capixaba sp. nov. is described from 54 m depth off Guarapari, diagnosed by its two clear-cut categories of isochelae from Phorbas spp. known from the Western Tropical Atlantic. Four nomina nova are proposed for homonyms in Phorbas spp.: P. bergmontae nom. nov. for P. areolatus Bergquist & Fromont, 1988, preoccupied by P. areolatus (Thiele, 1905); P. burtoni nom. nov. for P. arborescens sensu (Burton, 1956), preoccupied by P. arborescens (Ridley, 1884); P. hechteli nom. nov. for P. ramosus (Hechtel, 1983), preoccupied by P. ramosus (Lendenfeld, 1888; in part – composite species); P. tanitai nom. nov. for P. purpureus (Tanita, 1961), preoccupied by P. purpureus (Carter, 1886). Three little known species are redescribed: Aplysina alcicornis Pinheiro et al., 2007; Mycale (Aegogropila) escarlatei Hajdu et al., 1995 and Yucatania sphaeroidocladus (Hartman & Hubbard, 1999). All three are new records for Espírito Santo. The first and the last of these are range extensions, while the other fills a distribution gap. A list of sponges hitherto recorded from off Espírito Santo state, with indication of the bibliographic source of the record is given in appendix. In total, 118 species were compiled from 23 publications.

Yucatania sphaeroidocladus (Hartman & Hubbard, 1999) Figs 2-19, Tab.I Diagnosis.The only Thrombidae with demotriaenes.Description.The largest specimen available, still an obvious fragment, is MNRJ 4557.It is cushion shaped, 12 x 9 x 5 cm (largest width, smallest width, thickness).Specimens can agglutinate abundant sediment (MNRJ 5872) and/or grow intermingled to abundant siliquariid gastropod shells (Siliquaria modesta; MNRJ 4557).Consistency of the sponge, where devoid of inclusions, is firm but compressible, somewhat friable.Texture is mostly rough also as a consequence of all the inclusions.Oscula were not seen, possibly due to the macerated condition of most samples.Colour alive (on deck) was reported as beige (MNRJ 4387D,4562,6192), white (MNRJ 4574) and lively yellow (MNRJ 5868B).Skeleton (Fig. 4).Confused architecture.Acanthodemotriaenes far outnumber acanthotriaenes.These spicules are densely packed both in the ectosome and choanosome, appearing at places to be somewhat juxtaposed in a palisade manner, while in other areas they are completely criss-crossing one another.Amphiasters were only spotted in dissociated spicules' mounts.Spicules (Tab.I).Megascleres.Remarks.The material from Espírito Santo studied here conforms in most characters to specimens previously described from Trinidad (type locality; HARTMAN & HUBBARD 1999, as Thrombus sphaeroidocladus), the Yucatan Península, Mexico (GÓMEZ 2006, as Y. clavus) andfrom Bahia, Brazil (MENEGOLA et al. 2009).Not only spicule shape and dimensions conform in general but also the remarkable association to siliquariid gastropods is maintained over this rather large distance (ca.8000 km).A single exception to this near-perfect match is the mean length of rhabdomes found by HARTMAN & HUBBARD (1999) in the demotriaenes (as acanthotriaenes) of the type material -405-416 µm, as opposed to 275-287 µm found here, and 305-321 µm found elsewhere (GÓMEZ 2006;MENEGOLA et al. 2009).Given the similarity of all other aspects observed, this is considered of minor importance.HARTMAN & HUBBARD (1999) studied fragments from a very large sponge, said to cover more than 3 m 2 in area.They remarked that species of Thrombus are apparently very rare as they are known from scattered records only.This may not hold true for Yucatania, dredged by the Program REVIZEE from three seamounts and two continental shelf locations, off Espírito Santo state.Albeit registered from neighboring areas in Bahia (MENEGOLA et al. 2009), the species does not appear to be equally common there.
Description.The single specimen is a small bush with two main, rather irregularly outlined branches, presumably previously attached to the substrate by a couple of short stolons about 1 cm long and 1 cm wide.The greatest diameter (ground projection) of the sponge is 7 cm, and the largest branch is 4 cm long and up to 2 cm thick.Consistency is compressible and slightly elastic, and texture a bit rough due to the rather irregular outline of the branches, in part a consequence of embedded detritus.Surface is microconulose upon closer inspection.No areolated pore fields were recognized in the preserved specimen, which might be due to considerable damage to its surface having been inflicted by the dredging.Nevertheless, a thin transparent membrane can be seen at places, partially reticulated, and interrupted by a couple of small, inconspicuous oscula (1-2 mm in diameter).Color in life was reported as orange, which turned to beige in ethanol.Skeleton .Ectosome with dense single layered crust of isochelae (Fig. 21).Uncommon, short subectosomal tracts of tornotes fan out underneath the surface, only slightly piercing it.The choanosome has a mainly plumose architecture, with sinuous, densely echinated tracts running towards the surface.These appear to be cored by foreign debris.Echinating acanthostyles  Acanthostyles I, more heavily spined in the basal half, spines mostly slightly curved toward the base, but for the basal 1/6-1/ 7 where they point to the apex slightly, basalmost spines frequently blunt (Fig. 26).Acanthostyles II (Figs 27 and 28), nearly equally spined all over, with spine orientation similar to that of acanthostyles I. Microscleres.Arcuate isochelae 1, stout axis and poorly developed alae (ca.25% the total height; Fig. 29).Arcuate isochelae 2, slender axis, more developed alae (ca.33% the total height; Fig. 30).
Ecology and Distribution: So far known only from the type locality, the south-eastern Brazilian continental shelf off Guarapari, at 54 m depth.The species was dredged from calcareous bottoms.
Etymology.The species name is a noun in apposition referring to the people born in Espírito Santo State, the 'capixabas'.
Remarks.There are 76 species listed for Phorbas in the World Porifera Database ( VAN SOEST et al. 2009), only three of which are known from Brazil (Tab.II): P. amaranthus Duchassaing & Michelotti, 1864; P. fusifer (Ridley & Dendy, 1887) and P. hechteli [new name for P. ramosus (Hechtel, 1983; as Anchinoe ramosus), preoccupied by P. ramosus (Lendenfeld, 1888; as Echinonema ramosum)].The new species clearly differs from all, as argued below.Phorbas amaranthus, according to VAN SOEST (1984), has larger tornotes and a single category of isochelae, intermediate in size between both categories here reported for the new species.Furthermore, its tornotes and acanthostyles are rather slender.Phorbas fusifer, as originally described, possesses smaller and slender tornotes, a single category of acanthostyles of intermediate dimensions, and a single category of isochelae, also intermediate between both categories occurring in the new species.Finally, Phorbas hechteli new name bears considerably thinner megascleres, where acanthostyles belong in a single size category.Once again, isochelae occur in a single category, intermediate in size to those of P. capixaba sp.nov.No further species of Phorbas is known from the Tropical western Atlantic, so the new species is here considered well differentiated from its congeners inhabiting the biogeographic region it pertains to.By analyzing the list of species in VAN SOEST et al. (2009) it became apparent that homonymy is a remarkable problem in Phorbas.Apart from the above-mentioned case, another three pairs of homonymous species exist, viz.Phorbas arborescens (Ridley, 1884; as Myxilla arborescens) and P. arborescens (Burton, 1956; as Hymedesmia arborescens), P. areolatus (Thiele, 1905; as Hymedesmia areolata) and P. areolatus Bergquist &Fromont, 1988, andP. purpureus (Carter, 1886; as Plumohalichondria purpurea) and P. purpureus (Tanita, 1961; as Anchinoe purpurea).Nomina nova are proposed here for all three cases pointed above, as follows.For clarity sake, P. hechteli nom.nov. is also repeated below.Description.The collected specimen consisted of two small fragments, each <1 cm in maximum diameter and ca. 1 mm thick.Virtually nothing remains apart from the preparations used to base this description.Their surface was slightly hispid.No oscules were apparent.Color in the preserved state was beige.Skeleton (Fig. 31).Ectosomal architecture an isotropic triangular reticulation formed by multispicular tracts of mycalostyles (4-8 spicules across).Meshes are 150-200 µm in diameter.Anisochelae I are organized in abundant rosettes, attached to the multispicular tracts at random, where a few sigmas I can also be found.Abundant anisochelae IV and rather fewer anisochelae II and sigmas II occur right in the middle of the meshes, side by side with pores (20-80 µm in diameter).Spicules (Tab.III, Figs 32-41).Megascleres.Mycalostyles (Fig. 32).Microscleres.Anisochelae I (Fig. 33 and 34), head about 50% total spicule height, large frontal alae slightly bent forward.Anisochelae II (Fig. 35), 'duck's bill'-like, head about 75% total spicule height, frontal alae pending down parallel to the axis.Anisochelae IV (Fig. 36), head about 70% total spicule height, both frontal alae markedly spurred apically, and nearly overpassing each other.Sigmas I (Fig. 37), smooth, stout, contorted, uncommon.Sigmas II (Fig. 38),smooth,slender,contorted,common. Toxas (Figs 39 and 40), rare, smooth, variably curved (only three seen).Micracanthoxeas (Fig. 41), straight, heavily spined (sanidaster-like, only one seen).
Remarks.The dimensions of the studied material precluded any deeper search for additional toxas and micracanthoxeas.For most characters, the Espírito Santo material reported here matches closely the micrometric and micromorphologic data presented in the original description of the species (HAJDU et al. 1995).Nevertheless, data presented by MURICY & HAJDU (2006) deviate quite substantially, especially as far as sigmas I, toxas and micracanthoxeas are concerned, all reaching quite larger dimensions (respectively, 105, 140 and 9 µm).As the only additional material studied by MURICY & HAJDU (2006) in contrast to HAJDU et al. (1995) was the São Paulo State (Ilhabela) specimen(s), these appear to be in need of a thorough revision.

Verongida Bergquist, 1978
Aplysinidae Carter, 1875 Aplysina alcicornis Pinheiro, Hajdu & Custódio, 2007 Figs 42-46, Tab. IV Diagnosis.This is the only Aplysina in the Tropical western Atlantic with a flat lamellar body, relatively long, slender, irregularly digitiform fistules coloured as the sponge, and turning to a dark shade of brown in ethanol.
Description (Fig. 42-43).The single specimen is small, lamellate, ca. 8 cm high, 10 cm wide and 1 cm thick.Its apical portion is irregularly outlined, with short -lobate projections (ca.1-2 cm high) wherefrom fistules arise.Up to three fistules were seen originating from a single lobate projection.A few, thick, irregular digitiform processes are seen, either aligned to the main lamella (ca.8 cm high), or projecting from its side (ca. 2 cm high).The former bear apical fistules, while the latter do not.Surface is smooth, with a few oscules scattered on the apical portion of the lamella, 3-4 mm in diameter.Consistency was compressible, flexible.Live colour was homogeneously bright yellow (light coloured).Skeleton.A polygonal reticulation (Fig. 44) of spongin fibers (Figs 45 and 46) forming mostly three dimensional meshes of complex perimeter.Meshes are mostly broken up, but appear to be constrained between 400 and 1400 µm in diameter.
Ecology and distribution.The specimen was collected on rocky substrate, with an angle of about 60°, exposed to direct illumination.It had an epibiotic crinoid and many orange ophiuroids (Ophiotela danae, det.C.R.R. Ventura).
Remarks.Aplysina alcicornis is known from a single, recent description from the Abrolhos area (state of Bahia; PINHEIRO et al. 2007).This specimen was darker in color and its fistules could bifurcate.We do not judge the divergence observed in these characters when contrasting Bahia and Espírito Santo materials, sufficient to recognize yet another species in Aplysina, especially because so few specimens are available for study still.Similar color morphs occur in A. fistularis and A. fulva.The material described here brings new information, as this is the first time where a full specimen was photographed in situ.PINHEIRO et al. (2007) pointed out that anatomical (fiber) characters are quite inconclusive as diagnostic features.The fibers of the specimen studied here are slightly thinner than those reported from Bahia, albeit with thicker piths.These do not appear to be clear-cut diagnostic features, but a further analysis will be in order when additional A. alcicornis -like specimens are available.

DISCUSSION
The list of sponges from the state of Espírito Santo comprises now over 110 species, compiled from 23 publications (Appendix).Species reported from the Almirante Saldanha seamount were compiled here also.A large proportion of these species are known also from areas to the north of the state, and many frequently reach the Caribbean Sea, suggesting to a Tropical-western Atlantic affinity of the sponge fauna in this alleged biogeographic transition zone (SOLÉ-CAVA et al. 1981, PALACIO 1982).For example, 31 out of 47 fully-identified Demospongiae have known geographic ranges stretching all the way to the Central West Atlantic (see Appendix).
Among the remaining 62 species reported, 46 are identified only to the generic level (as "sp.", "cf.sp.X" or "aff.sp.X") and may prove to be conspecific to known Caribbean sponges or strengthen the group of provisional Brazilian endemics.The number of provisional endemic Brazilian demosponges described from the coast of the state of Espírito Santo is 15, including six provisional local endemic species -Cinachyra helena Rodriguez & Muricy, 2007;Erylus almirante Vieira et al., 2010;E. fluminense Vieira et al., 2010;E. revizee Vieira et al., 2010;Jaspis salvadori Boury-Esnault, 1973 and     (2006) suggest that these mountains act as a filter to sponge colonization of Trindade Island, which appears to be comparatively poor (23 species known up to now) in contrast to localities closer to the continent (over 100 species registered; Appendix).It seems premature to consider this a well-established general knowledge, because the taxonomic inventory of the sponge fauna of Trindade Island 'is still in its early steps.But, on the other hand, the impoverished insular fauna, when contrasted to localities on the continental shelf or closer to it, is the expected outcome of Island Biogeography Theory and a likely consequence of the comparatively smaller area and large distance of the island from the mainland.

Figs 20 -
Figs 20-30, Tab.II Holotype.BRAZIL, Espírito Santo: Off Guarapari (Vitória-Trindade Seamounts Chain, Program REVIZEE, R/V Astro Garoupa, Central VI cruise, stn.#y4, 20.633°S-40.025°W,54 m depth), 14/vi/2002, H.P. Lavrado leg., MNRJ 6176.Diagnosis.This is the only Phorbas in the Tropical western Atlantic with two clear cut categories of arcuate isochelae.Description.The single specimen is a small bush with two main, rather irregularly outlined branches, presumably previously attached to the substrate by a couple of short stolons about 1 cm long and 1 cm wide.The greatest diameter (ground projection) of the sponge is 7 cm, and the largest branch is 4 cm long and up to 2 cm thick.Consistency is compressible and slightly elastic, and texture a bit rough due to the rather irregular outline of the branches, in part a consequence of embedded detritus.Surface is microconulose upon closer inspection.No areolated pore fields were recognized in the preserved specimen, which might be due to considerable damage to its surface having been inflicted by the dredging.Nevertheless, a thin transparent membrane can be seen at places, partially reticulated, and interrupted by a couple of small, inconspicuous oscula (1-2 mm in diameter).Color in life was reported as orange, which turned to beige in ethanol.Skeleton.Ectosome with dense single layered crust of isochelae (Fig.21).Uncommon, short subectosomal tracts of tornotes fan out underneath the surface, only slightly piercing it.The choanosome has a mainly plumose architecture, with sinuous, densely echinated tracts running towards the surface.These appear to be cored by foreign debris.Echinating acanthostyles
2):233-246, April, 2011    Phorbas capixaba sp.nov.The 13 Hexactinellida present another three provisional endemic species -Hyalonema conquerorTabachnick et al., 2009; Hyalonema dufresnei Tabachnick et al.,  2009 and Lophocalyx brasiliensis Menshenina et al., 2007.Of the 12 Calcarea reported herein, three are provisional endemics -Paraleucilla sp.(referred as 'sp.nov.' in MORAES et al. 2006); Sycettusa pelagica(Ridley, 1881)  and Vosmaeropsis sericatum (Ridley, 1881).The latter two are also provisional endemics from the state of Espírito Santo.Only two species are provisional disjunct records from non-adjacent areas, both in need of detailed morphologic comparisons with topotypical specimens -Dysidea avara (Schmidt, 1862) and Timea cumana Pulitzer-Finali, 1977.The MNRJ and UFRJPOR collections together hold over 1000 sponge samples originating from the coast of the state of Espírito Santo, which are bound to reveal important answers to questions related to marine sponge biodiversity in this biogeographic transitional zone.An important research venue regards the level of population connectivity along the line of stepping-stones represented by the seamounts stretching from the border of the continental shelf off Vitória, to Trindade Island and the Archipelago of Martim Vaz, ca.1160 km to the east.This issue might be studied through a comprehensive evaluation of genetic homogeneity (percentage of haplotypes) and distances between distinct populations of the same species, occurring along this latitude.Data presented in MORAES et

Table II .
Comparative table of spicule dimensions of Tropical western Atlantic Phorbas.Values are in micrometers and are expressed as follows: smaller length -mean -larger length/smaller width -mean -larger width (N, when distinct from 25 spicules).(n.r.)Not recorded.

Table III .
Comparative table of spicule dimensions of published records of Mycale (Aegogropila) escarlatei type series, contrasted to those observed here for the Espírito Santo specimen.Values are in micrometers and are expressed as follows: smaller length -mean -larger length/smaller width -mean -larger width (N, when distinct from 20 spicules).(n.f.)Not found.

Table IV
. Comparative table of spongin fibers' dimensions of Aplysina alcicornis type series, contrasted to those observed here for the Espírito Santo specimen.Values are in micrometers, expressed as follows: smaller width -mean -larger width.