Aplysina fulva (Pallas, 1766)

Pinheiro, Ulisses Dos S., Hajdu, Eduardo & Custódio, Márcio R., 2007, Aplysina Nardo (Porifera, Verongida, Aplysinidae) from the Brazilian coast with description of eight new species, Zootaxa 1609, pp. 1-51 : 14-21

publication ID	https://doi.org/ 10.5281/zenodo.178878
DOI	https://doi.org/10.5281/zenodo.6240783
persistent identifier	https://treatment.plazi.org/id/03E387FE-FFC9-FFF4-E3B5-FC9EFF3FB99B
treatment provided by	Plazi
scientific name	Aplysina fulva (Pallas, 1766)
status

Treatment

Aplysina fulva (Pallas, 1766) ( Figs. 3 View FIGURE 3 D, 7–8, Tab. IV)

Spongia fulva, Pallas (1766: 383) . Verongia fistularis forma fulva sensu Collete & Rützler (1977: 309) . Aplysina fistularis sensu Muricy (1989: 351) . Non Aplysina fistularis (Pallas, 1766; a valid species). Aplysina fulva sensu Muricy et al. (1991: 1187) , Muricy et al. (1993: 429), Pinheiro & Hajdu (2001: 149). Aplysina aff. cauliformis sensu Pinheiro & Hajdu (2001: 154) . Non Aplysina cauliformis (Carter, 1882, a valid species). For further synonymy cf. Wiedenmayer (1977: 66).

Type specimens status: As stated in the discussion on the type-specimen status for A. fistularis , the type material of Spongia fulva was also lost in the Seven Years´War. Aplysina fulva is a very inadequately diagnosed species amongst its congeners, as the debate upon its specific status lasted until the 1990s, when the species has been referred to A. fistularis forma fulva for the last time. In other words, doubts persisted on its specific status until very recently, and designation of a neotype would be most welcome. Surprisingly, this has never been done.

From Wiedenmayer’s (1977), van Soest’s (1978) and Zea’s (1987) descriptions of A. fulva , it appears that what seemed typical to them has some discrepancies to what appears typical, both from interpretation of Pallas´(1766) description and from extensive analysis of Brazilian populations. The few exceptions regarding A. fulva ´s morphospace according to our judgement are Wiedenmayer´s (1977) mention of "hollow cylinders" at the base of specimens, van Soest´s (1978) quotation of "large oscula with iris-type diaphragm" also on the massively encrusting base of specimens (his Plate XI, Fig. 2 View FIGURE 2 ), and Zea´s (1987) mention of oscula on top of tubular elevations up to 1–2 cm high. These features are absent from the Brazilian material studied here, and are suggestive to us that these authors might have been studying a complex of species, rather than A. fulva alone. Above all, these features do not match the description provided by Pallas (1766; translated in Wiedenmayer, 1977), nor the illustrations provided by Seba (1758) and reproduced in Wiedenmayer (1977; Pl. 6, figs. 2–3). None of these speak of, or picture any type of atrial cavity or large, iris-bearing oscula in A. fulva . Our choice of a neotype for A. fulva takes into consideration the description by Pallas (1766), the illustrations by Seba (1758), one of Wiedenmayer´s (1977) illustrated specimens (Plate 6, Figure 4 View FIGURE 4 ), Vacelet´s A. fulva (1990: colour photo on p. 31), and the colour pictures provided in Pinheiro & Hajdu (2001; figs. 3, 7–12).

The ICZN lists seven qualifying conditions for recognition of the validity of any neotype designation. We will argue how we match each of these.

Condition 75.3.1 determines that a neotype designation must be accompanied by a statement that it is selected with the express purpose of clarifying the taxonomic status or the type locality of a nominal taxon. This is so here. The arguments presented above clearly indicate how much confusion persists in the recent literature regarding the actual morphospace of A. fulva , above all through the realization that until very recently this species was considered only a variety of A. fistularis .

Condition 75.3.2. determines that a neotype designation must also be accompanied by a statement of the characters that the author regards as differentiating from other taxa the nominal species-group taxon for which the neotype is designated, or a bibliographic reference to such a statement. A diagnosis for the species is provided below.

Condition 75.3.3. determines that a neotype designation must also be accompanied by data and description sufficient to ensure recognition of the specimen designated. Both a description of the neotype, including underwater photo of the live specimen, and a full description of the species on the basis of extensive evaluation of Brazilian populations are provided below.

Condition 75.3.4. determines that a neotype designation must include the author's reasons for believing the name-bearing type specimen(s) (i.e. holotype, or lectotype, or all syntypes, or prior neotype) to be lost or destroyed, and the steps that had been taken to trace it or them. The holotype of Aplysina fulva is Spongia fulva Pallas, 1766 as argued by Wiedenmayer (1977), who also pointed to the fact that this collection was entirely lost in a fire in Küstrin in the Seven Year´s War.

Condition 75.3.5. determines that a neotype designation must include evidence that the neotype is consistent with what is known of the former name-bearing type from the original description and from other sources. This has been extensively argued above. In spite of the few points of disagreement with the descriptions provided by Wiedenmayer (1977), van Soest (1978) and Zea (1987), the neotype proposed here would undoubtedly have been identified as A. fulva by all these authors. On top of that, our specimen appears to perfectly match Pallas´(1766) description. Our choice for a newly collected specimen stresses the importance of knowing the specimen´s live-colour, as seen in Fig. 7A View FIGURE 7 . The lighter extremities exhibited by the proposed neotype were reported upon by Wiedenmayer (1977) too, for specimens from the Bahamas.

Condition 75.3.6. determines that a neotype designation must include evidence that the neotype came as nearly as practicable from the original type locality. Pallas (1766) stated the original locality to be "Mare Americanum", which, according to our comprehension must be taken as comprising all the Tropical western Atlantic. Given that Peter Simon Pallas did his doctorate in Leiden in 1760 and subsequently spent some time in The Hague, where he published his Elenchus Zoophytorum , it appears quite conceivable to us that he might have had access to Brazilian materials. These could have been brought back to The Netherlands a century before during the time the Dutch had a colony in north-eastern Brazil, where A. fulva is abundant (even as debris at the many beaches), or subsequently, by any merchants or corsairs. Also, Pallas referred to pictures provided in Seba (1758), who lived in Amsterdam, and had been buying "curiosities" (such as sponges) at the docks of this port town, so that a Brazilian origin for his samples cannot be discarded either.

Condition 75.3.7. determines that a neotype designation must include a statement that the neotype is, or immediately upon publication has become, the property of a recognized scientific or educational institution, cited by name, that maintains a research collection, with proper facilities for preserving name-bearing types, and that makes them accessible for study. The natural history collections of Museu Nacional/ UFRJ are nearly 190 years old. The sponge collection comprises over 16000 specimens, over 150 of which are types or fragments from types.

Neotype: MNRJ 7975 ( Figs. 7A View FIGURE 7 , 8A View FIGURE 8 ), Tartarugas Beach, (Búzios, RJ), 5 m depth, U. S. Pinheiro and E. Hajdu coll., 15/v/2003. The Neotype consists of a cluster of twenty-seven digitiform processes (cylindrical or fusiform), always with irregular diameter. The specimen is 19 cm high and 18 cm wide. The small oscula (0.5–2 mm) appear randomly spread all over the surface, which can be smooth or finely conulose. The colour in vivo varies from yellow (mostly on the apices of the branches) to brown, becoming deep brownish-purple after preservation in alcohol. Without ectosomal specialization. Choanosome with an irregular network of smooth, straight, curved or undulating spongin fibers (fibres 39–68 µm in diameter, pith 34–58 µm in diameter). After preservation and preparation, fibers are yellow to dark-brown, nearly black. The same specimen has pith of both colours.

Additional material studied: Ceará State - MNRJ 689, Meirelles Beach (Fortaleza), A.L. Castro coll. MNRJ 3044, 3045, 3047 (Canoa Quebrada), M. Guimarães coll., 25 m depth, II/2000. UFRJPOR 4285, E. Araujo coll. Rio Grande do Norte State - UFRJPOR 3582 Urca do Tubarão, (04º50´S - 36º27´W), diver of Cooperbrasub coll. Pernambuco State - UFRJPOR 4804, Rata Island (Fernando de Noronha Archipelago), G. Muricy coll., 12 m depth, 14/II/1998. Alagoas State - MNRJ 1987, do Francês Beach (Marechal Deodoro), M.D. Correia coll., 1 m depth, 22/IX/1998. Bahia State - MNRJ 823, Coroa de Pedra, off Itaparica Island, Todos os Santos Bay (Itaparica), P.S. Young and M.B. Pereira coll., 03/II/1993. MNRJ 1486, 1487, Tacimirim Beach (Morro de São Paulo), R. Fernandes coll., 13 m depth, 11/I/1998. MNRJ 1506, Recife Califórnia, Programme REVIZEE coll., 20–30 m depth, 25/XII/1997. MNRJ 2599, Portinho de Itaparica (Itaparica, 12º53.44' S - 38º41.021' W), E. Hajdu coll., 2–4 m, 04/VIII/1999. MNRJ 2603, breakwater of the Harbour Authority, ( Salvador, 12º58.191' S - 38º31.237' W), E. Hajdu coll., 07/VIII/1999. MNRJ 3056, 3064 (Itacaré, 14º16.631´S - 38º58.888´W), A.C. da Silva coll., 15 m depth, 25/III/2000. MNRJ 3531, Pedra do Silva (Reserva Extrativista de Corumbau, Prado), G. Muricy coll., 1–3 m depth, 14/XI/1999. MNRJ 3541, 3547 (Reserva Extrativista de Corumbau, Prado, 16 º56'29'' S 39º00'24'' W), B. Segal and C.B. Castro coll., 14–22 m depth, 18/XI/1999. MNRJ 4168, 4171, 4177, 4178, 4180, 4181 Salvador Yacht Club ( Salvador), E. Hajdu coll., 3– 5 m. depth, 4/VII/2001. MNRJ 5479, Parcel Paredes, south side, (Parque Nacional Marinho dos Abrolhos, 17º53'499'' S - 38º58'034'' W), U.S. Pinheiro and G. Muricy coll., 10 m depth, 05/III/2002. UFRJPOR 3423, Calypso coll., station 67 (13º28´S - 38º48´W), 39 m depth. Espírito Santo State - MNRJ 2298 (Guarapari), A.M. Solé-Cava coll., III/1989. Rio de Janeiro State - MNRJ 83, João Fernandinho Beach (Armação dos Búzios), D. Pires coll., 26/VII/1983. MNRJ 1704, João Fernandinho Beach (Armação dos Búzios), E. Hajdu coll., 2–7 m depth, 09/V/1998. MNRJ 3554, Forno Beach (Arraial do Cabo), 2 m depth, 05/XII/ 1998. MNRJ 3957, Tartaruga Beach, (Armação dos Búzios), J. Creed coll., 02/IX/2000. MNRJ 3958, Azedinha Beach, (Armação dos Búzios), J. Creed coll., 07/IX/2000. MNRJ 3961, Canto Beach (Armação dos Búzios), J. Creed. coll., 30/IX/2000. MNRJ 3963, Caboclo Island (Armação dos Búzios), J. Creed. coll., 05/ IX/2000. MNRJ 4084, Ponta do Anequim (Cabo Frio Island, Arraial do Cabo), E. Hajdu and E. Vilanova coll., 6–10 m, 08/IV/2001. MNRJ 4152, Porcos Pequena Island (Angra dos Reis), G. Muricy and E. Vilanova coll., 2–5 m depth, 26/V/2001. MNRJ 4153, 4154, 4155, 4156, 4157, 4158, 4159, 4160, Porcos Pequena Island (Angra dos Reis), G. Muricy and E. Vilanova coll., 2–5 m depth, 26/V/2001. UFRJPOR 1204, Prainha Beach (Arraial do Cabo). UFRJPOR 1206, Boqueirão (Arraial do Cabo). UFRJPOR 2917, Pedra Vermelha (Cabo Frio Island, Arraial do Cabo), 10/V/1987. UFRJPOR 2940, Pedra Vermelha (Cabo Frio Island, Arraial do Cabo), N. Boury-Esnault coll., 14/VI/1987. UFRJPOR 2965, Pedra Vermelha (Cabo Frio Island, Arraial do Cabo), C. Russo coll., 24/VIII/1987. São Paulo State - MNRJ 357, Ponta do Jarobá, São Sebastião Channel (São Sebastião, 23º49.676' S - 45º25.278' W), E. Hajdu and G. Muricy coll., 0.5–2.5m depth, 26/I/1996. MNRJ 445, São Sebastião Channel (São Sebastião, 23º49.676' S - 45º25.278' W), E. Hajdu coll., 4 m depth, 19/II/1997. MNRJ 735, Barequeçaba, São Sebastião Channel (São Sebastião, 23º49.746' S - 45º26.478' W), E. Hajdu coll., 1 m depth, 08/I/1996. MNRJ 762, Saco do Poço, São Sebastião Island (Ilhabela, 23º45.658' S - 45º14.862' W), E. Hajdu coll., 8 m depth, 09/I/1996. MNRJ 1025, São Sebastião Channel (São Sebastião), R.G.S. Berlinck coll., I/1994. MNRJ 1302, Ponta do Jarobá, (São Sebastião), E. Hajdu coll., 2 m depth, 21/ XII/1997. MNRJ 1674, rocky coast between Prainha Beach and Toque-toque Beach (São Sebastião, 23º50.067' S - 45º29.449' W), E. Hajdu coll., 8 m depth, 24/IV/1998. MNRJ 1990, rocky coast between Prainha beach and Brava beach (Costão do Navio, São Sebastião, 23º49.607´S - 45º28.557´W), E. Hajdu coll., 6 m. depth, 29/I/1999. MNRJ 1991, Pedra Montada, Barequeçaba (São Sebastião, 23º49.746' S - 45º26.478' W), E. Hajdu coll., 4 m depth, 01/II/1999. MNRJ 3004, Curral Beach, São Sebastião Channel (Ilhabela), E. Hajdu coll., 10/I/2000. MNRJ 3561, between São Pedro and Veloso, (Ilhabela), E. Hajdu coll., 6 m depth, 10/XI/2000.

Diagnosis: Single branches or clusters of erect or repent projections, mostly digitiform of irregular diameter, but also lamellate, palmate and volcaniform, with small, scattered oscula. Live-colour mostly ochreyellow, but also brown, lemon-yellow or purplish-rose.

Description

The majority of specimens are composed of clusters of digitiform branches (cylindrical or fusiform), always with irregular diameter ( Fig. 7C View FIGURE 7 ). However, other morphologies are found, such as cushion-shaped, reptant (e.g. MNRJ 4177, 4178), lamellate (e.g. MNRJ 3056), palmate/pinnate (e.g. MNRJ 3963; Fig. 7B View FIGURE 7 ), ficiform/flabellate (e.g. MNRJ 2298) and volcaniform (e.g. MNRJ 4178). The size of specimens also varies considerably, with fused processes over 2 m high observed at Parque Nacional Marinho dos Abrolhos, while others are smaller than 3 cm in height, at the species known southern distribution limit (24º S). The processes can be found single or in groups of 2 to 80. The small oscula (0.5–2 mm) appear randomly spread all over the surface, most frequently, but some alignment is also visible. The surface can be smooth, finely conulose, rugose or marked by short irregular ridges and grooves. The colour in vivo is most often a brownish-yellow, but varies in diverse hues of yellow, green, lilac or brown, becoming purple, pink or dark brown after preservation in alcohol. Specimens in semi-obscure environments are often bright lemon-yellow, as well as small. The consistency is variable from extremely soft to considerably hard.

Skeleton: Choanosome formed by an irregular polygonal reticulation of spongin fibers, with a diameter of 21–275 Μm (average 75 Μm; Fig. 8B–C View FIGURE 8 ). The bark is amber in colour, with the exception of specimens with stouter fibers, which have a reddish bark. The pith is black, with diameter of 10–72 Μm (average 27 Μm; Fig. 8D View FIGURE 8 ).

TABLE IV: Spongin fibres’ measurement data for Aplysina fulva (Pallas, 1766) (in micrometers; S.D. = Standard Deviation and N=30).

Specimens Locality* Fibers Piths

Thinnest Mean Thickest S.D. Thinnest Mean Thickest S.D.

...... continued (next page) *AL, Algoas State; BA, Bahia State; CE, Ceará State; ES, Espirito Santo State; PE, Pernambuco State; RJ, Rio de Janeiro State; RN, Rio Grande do Norte State; SP, São Paulo State.

Distribution: Tropical western Atlantic: Brazil (3–24º S, Fig. 3 View FIGURE 3 D): Ceará, Rio Grande do Norte, Pernambuco (Fernando de Noronha Archipelago), Alagoas, Bahia ( Salvador, Parque Nacional Marinho dos Abrolhos), Espírito Santo, Rio de Janeiro, São Paulo. World: Colombia, Cuba, Curaçao, Bahamas, Barbados, Bonaire, Florida, U.S. Virgin Islands and Virgin Island.

Ecology: Aplysina fulva is found on diverse environments. Habitats can have high or low hydrodynamic conditions, be exposed or protected from light, and range from 1 to 40 m in depht. Specimens living in highenergy environments were seen to be creeping, forming an anastomosing system of ridges topped by a dense series of small oscula. Aplysina fulva ´s population can be scarce, with only some individuals, or one of the most conspicuous components of all benthic fauna of the locality (e.g. Porcos Island, Arraial do Cabo; Tartaruga Beach, Búzios, RJ). This sponge also presents a substantial aggregated fauna, such as polychaetes, ophiuroids, amphipods and decapods. We observed the seastar Oreaster reticulatus feeding upon Aplysina fulva at Búzios (RJ). The specimen had turned black at the point of contact with its predator.

Remarks: Our knowledge of Aplysina fulva ’s morphologic plasticity is based on nearly two decades of in situ observation of variably dense populations, mostly from São Paulo, Rio de Janeiro and Bahia States. These were coupled to isolated observations of other Brazilian populations, frequent underwater photography, some videos, and laboratory work on the collections of Museu Nacional. Our conclusion is that a typical Brazilian A. fulva will be composed of an erect cluster of solid, irregular, digitiform branches, mostly 10–30 cm high and 1–4 cm thick, and will be of a brownish-yellow live-colour. Nevertheless, variability is the rule for A. fulva , and no clear morphologic gaps appear to allow recognition of additional species. Specifically, a doubt persists on how much flat palmate A. fulva can be. This is a crucial point to evaluate its status in face of A. pergamentacea , and some new, mainly lamelate species described below. The specimens of A. fulva studied here which showed some sort of considerable lateral compression of their branches (e.g. Fig. 7B View FIGURE 7 ), were referred to A. fulva instead of A. pergamentacea , because they possessed cylindrical branches in addition, and/ or turned to very dark shades of purplish-brown in the fixative, nearly black if formerly exposed to the air. Contrastingly, as shown below, A. pergamentacea remains lighter coloured in the fixative.

Further variability is seen in the number of digits, which spread from a single to over 80 in a cluster. Their height may be consistently smaller than 10 cm in an entire population (São Sebastião Channel; cf. Pinheiro & Hajdu, 2001), or reach over 1 m (Parque Nacional Marinho dos Abrolhos), as illustrated by Humann (1992) for the West Indies (det. S. Pomponi).

An investigation of the cellular morphology of A. fulva showed that despite the external morphologic variability, the cellular morphotypes from distant localities (ca. 400 km) do not present significant differences ( Pinheiro et al, 2004). However, complementary studies, perhaps with the use of molecular techniques, will still be necessary to clear the status of the species along its large distribution range on the Tropical western Atlantic.