Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences

The question whether taxonomic descriptions naming new animal species without type specimen(s) deposited in collections should be accepted for publication by scientific journals and allowed by the Code has already been discussed in Zootaxa (Dubois & Nemésio 2007; Donegan 2008, 2009; Nemésio 2009a–b; Dubois 2009; Gentile & Snell 2009; Minelli 2009; Cianferoni & Bartolozzi 2016; Amorim et al. 2016). This question was again raised in a letter supported by 35 signatories published in the journal Nature (Pape et al. 2016) on 15 September 2016. On 25 September 2016, the following rebuttal (strictly limited to 300 words as per the editorial rules of Nature) was submitted to Nature, which on 18 October 2016 refused to publish it. As we think this problem is a very important one for zoological taxonomy, this text is published here exactly as submitted to Nature, followed by the list of the 493 taxonomists and collection-based researchers who signed it in the short time span from 20 September to 6 October 2016

Aplysina lacunosa Lamarck 1814

Aplysina lacunosa (Lamarck, 1814) (Figs. 11 B– 12, Tab. VI) Aplysina lacunosa, Hechtel (1983: 59), Muricy et al. (2006: 118); Verongia sp.a. sensu Hechtel (1976: 239). For further synonymy cf. van Soest (1978: 61). Lectotype: MNHN unregistered (designated by Topsent, 1932: 70, Pl. I, Fig. 5; cf. Wiedenmayer, 1977: 67) Studied material: Ceará - MNRJ 3046 (Canoa Quebrada). M. Guimarães coll, 25 m depth, II/ 2000. Bahia - MNRJ 3387, Bank Rodger, G. Nunan coll., 40–50 m depth, 6 /VI/ 1999. MNRJ 3402, Bank Rodger, G. Nunan coll., 40–50 m depth, 6 /VI/ 1999. MNRJ 3524 (Reserva Extrativista de Corumbau, Prado, 16 º 56`29 `` S - 39 º00`24 `` W), B. Segal and C.B. Castro coll., 18 /XI/ 1999. MNRJ 3557 (Reserva Extrativista de Corumbau, Prado) G. Muricy coll., 2 m depth, III/ 99. MNRJ 4436 (Programme REVIZEE Central V, Porto Seguro, 17 º06' 18 '' S - 36 º 44 ' 74 '' W), 50 m depth, 07/VII/ 2001. MNRJ 4652 (Programme REVIZEE Central V), 29 /VI/ 2001. Espírito Santo - MNRJ 4429 (20 º 30 ' 37.32 '' S 37 º 19 '05.39'' W), 89 m depth, 24 /IV/ 1996. MNRJ 4587 (Programme REVIZEE Central V., 19 º 48 ' 47 '' S - 037º 56 ' 33 '' W), 60 m depth, 18 /VII/ 2001. Diagnosis: Tubular sponge with a large apical pseudoscule, surface with irregular grooves (caliciform, meandriform) of variable depth. FIGURE 12: A–D. Aplysina lacunosa (Pallas, 1766). A–B. MNRJ 3387 (after preservation). C. Skeletal architecture (MNRJ 3046). D. Spongin fibres (MNRJ 3387). Scale bars = A–B. 1 cm, C. 500 µm, D. 50 µm. [The submitted file is of 72 dpi. Please resubmit a 300 dpi file]. Description The sponge consists of a single tube or clusters of up to 19 tubes, with a maximum length of 18.5 by 4.5 cm in diameter, and a large apical pseudoscule with 2.5 cm in diameter. The surface is characterized by irregular grooves (caliciform or meandriform) of variable depth. Specimens are finely conulose, small digitiform processes (ca. 0.5 cm in diameter) being observed in some specimens, projecting themselves from the rim of the pseudoscule. The colour is bright yellow in vivo, turning beige, purple or dark brown after preservation in alcohol. Consistency varies from hard to soft. Skeleton: Choanosome with a delicate and irregular network of spongin fibers with amber colour and thickness of 7–196 Μm (average 89 Μm). They have rather irregular (eccentric) black pith with thickness 7–37 Μm (average 16 Μm). Distribution: Tropical western Atlantic, Brazil (3–20 ºS): Ceará State, Bahia State, Reserva Extrativista de Corumbau, Espírito Santo State. World: Bahamas, Florida, Jamaica, Puerto Rico, Dominican Republic, Colombia, Bonaire, Curaçao. Ecology: The specimens were collected in warm and clear waters, 3–89 m deep. Remarks: Wiedenmayer (1977) commented that the fine pith was the character that differentiated A. lacunosa from its congeners. However, corroborating van Soest (1978), in this work we observed fibers in A. fistularis (MNRJ 5471) and A. fulva (MNRJ 5479) with fine pith and thick bark, and some specimens of A. lacunosa with thick pith and reduced bark (e.g. MNRJ 3387). These facts lead us to believe that the best diagnostic character for the species is the external morphology (surface with irregular, caliciform or meandriform grooves of variable depth), since this is not observed in any other species of the genus. van Soest (1978) and Zea (1987) raised the possibility that material identified as A. lacunosa from the Caribbean could comprise two species. Following this, Schmitt et al. (2005) differentiated a hard morphotype with reticulate skeleton of typically thinly pithed spongin fibers, and a soft morphotype with a more dendritic skeleton of heavily pithed fibers, reminiscent of the genus Suberea of the verongid family Aplysinellidae. The Brazilian material appears to us to belong to a single species, and having a typical reticulate skeleton, it conforms to the hard form of these authors. TABLE VI: Spongin fibres’ measurement data for Aplysina lacunosa (Lamarck, 1814) (in micrometers; S.D. = Standard Deviation and N= 30).Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 25-27, DOI: 10.5281/zenodo.17887

Aplysina

Aplysina in the Tropical western Atlantic Despite the extensive faunistic surveys on the Brazilian littoral zone and continental shelf undertaken between the 1960 ’s and the turn of the century, no other specimen of A. archeri (Higgins, 1875) was recollected. Therefore, we believe the record by Lendenfeld [1889; for Polejaeff’s (1884) A. tenuissima] to be a mistaken identification (probably A. lacunosa cf. Pinheiro & Hajdu, 2001) and consider this record invalid. We also invalidated A. capensis, following the steps of Carter (1881) and Bergquist (1980). We had the opportunity to reexamine several of Carter’s (1881), South African specimens (BMNH? – type, BMNH 1871.5. 12.1, BMNH 1938.3.26.82), which clearly contain abundant foreign inclusions in their fibres, thus being exclused from Aplysina ’s scope. De Laubenfels’ (1956) Brazilian record of the species is thus unreliable in view of the absence of a description, absence of a voucher specimen, and unrecognizable identification. Aplysina is one of a few sponge genera to present a larger diversity in the Atlantic (ca. twenty species), than the Indo-Pacific (fifteen species—many of which undoubtedly unrecognizable) oceans. In the Atlantic Ocean, the Caribbean region was the center of larger diversity, with seven species known: A. archeri, A. Bathyphila, A. cauliformis, A. fistularis, A. fulva, A. lacunosa and A. ocracea (sensu Maldonado & Young, 1998). In the present work we described eight additional new species and a new record for the Brazilian coast, raising the number of annotated species in this region from six (Pinheiro & Hajdu, 2001) to fifteen. Additional species for the Caribbean are also certain, as realized from a preliminary comparative assessment of Aplysina specimens in the BMNH and ZMA collections. These await a much needed revision of Caribbean specimens, which given the rationale above, is to be based on extensive field observations conducted on as many localities, as far apart from each other, as possible, as already argued for by Wiedenmayer (1977) for sponges in general. Considering Aplysina ’s conspicuousness, abundance and diversified chemistry associated to promising biological activities, it is a clear priority for a deeper, comprehensive taxonomic revision. Following is the amended list of species of Aplysina occurring in the South-western Atlantic, after which a tentative identification key for these species is offered: A. alcicornis sp.n., A. caissara, A. cauliformis, A. cristagallus sp.n., A. fistularis, A. fulva, A. insularis, A. lacunosa, A. lactuca sp.n., A. lingua sp.n., A. muricyana sp.n., A. orthoreticulata sp.n., A. pergamentacea, A. pseudolacunosa sp.n. and A. solangeae sp.n. In total, fifteen species were recognized.Published as part of 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, pp. 1-51 in Zootaxa 1609 on page 47, DOI: 10.5281/zenodo.17887

Aplysina lingua Pinheiro, Hajdu & Custódio, 2007, sp.n.

Aplysina lingua sp.n. (Figs. 16 C, 17 B, 19, Tab. IX) Holotype: MNRJ 5476, west coral head (Parque Nacional Marinho dos Abrolhos, BA, 17 º 57 ' 703 '' S - 36 º 14 ' 795 '' W), 18 m depth, E. Esteves and G. Muricy coll., 03/III/ 2002. Paratype: MNRJ 5469, Parcel Paredes, south side, (Parque Nacional Marinho dos Abrolhos, BA, 17 º 53 ' 499 '' S - 38 º 58 '034'' W), 10 m depth, U.S. Pinheiro and G. Muricy coll., 05/III/ 2002. Diagnosis: Elongate lamellar form, similar to a tongue, Light yellow colour in vivo. Description of the species The specimen has an erect, tongue form (long, flattened, lamelar) with 18 cm length by 4 cm width and 2 cm thickness expanding toward the apex (Fig. 17 B, 19 A). The small oscula (1 mm) are spread over the entire surface, which is very finely conulose. The colour is a mixture of light yellowish-brown, light pinkish-brown and light-yellow in vivo, which turns into dark brownish-purple after preservation in ethanol. Consistency is flexible. Skeleton: Choanosome with a delicate and irregular network of spongin fibers (Fig. 19 B). Bark with amber or reddish colour, thickness of 37–192 Μm (average 117 Μm) and thick pith that can be black or amber with thickness varying between 10 and 35 Μm (average 31 Μm; Fig. 19 C). Distribution: Provisionally known only from the type locality off the State of Bahia (Brazil; Fig. 16 C). Ecology: The specimens were collected in a maximum of 18 m depth in shaded areas of coral heads. Etymology: The name of the species is a noum in apposition and relates to its morphology tongue-like (tongue = lingua in Latin). Remarks: A. lingua sp.n. approaches more closely the morphospace of A. fulva, known for its plasticity and undelimited diagnosis. Nevertheless, the hundreds to thousands of specimens of A. fulva observed at many localities along the Brazilian Coast only very seldom possessed a comparable mixture of colours when alive, and none of them were observed to comprise a single branch with a flattened, elongate, lamelar form. TABLE IX: Spongin fibres’ measurement data for Aplysina lingua sp.n. (in micrometers; S.D. = Standard Deviation and N= 30). Specimens Locality* Fibers Piths Thinnest Mean Thickest S.D. Thinnest Mean Thickest S.D. Holotype Abrolhos, BA 60.0 147.7 192.5 35.6 10.0 20.5 35.0 6.1 MNRJ 5476 Paratype Abrolhos, BA 37.5 88.3 175.0 25.1 10.0 16.4 27.5 4.8 MNRJ 5469 *BA, Bahia State.Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 35-36, DOI: 10.5281/zenodo.17887

Aplysina lactuca Pinheiro, Hajdu & Custódio, 2007, sp.n.

Aplysina lactuca sp.n. (Figs. 16 B, 17 A, 18, Tab. VIII) Holotype: MNRJ 5477, Siriba Island, (Parque Nacional Marinho dos Abrolhos, BA), 10 m depth, U.S. Pinheiro and E. Esteves coll., 02/III/ 2002. Paratypes: MNRJ 5276, 5278, Pedra do Silva, south side, (Reserva Extrativista de Corumbau, Prado, BA, 16 º 53 ' 42.1 '' S - 39 º05' 31.2 '' W), 5 m depth, U.S. Pinheiro coll., 23 /I/ 2002. Additional material: MNRJ 8672, Parque Estadual da Pedra da Risca do Meio (03° 35 ' 889 S - 38 ° 23 ' 481 W, Fortaleza, CE, Brazil), 22 m depth, E. Hajdu coll., 14 /VII/ 2004. Diagnosis: It presents a centrally anastomosed lamellar form, which reminds the root buttresses of large tropical canopy trees. The species is yellowish-brown in vivo, turning purple after preservation in ethanol. Description of the species: The specimens besrs centrally anastomosed lamellas, giving the sponge a form which reminds the root buttresses of large tropical canopy trees (Figs. 17 A, 18 A–B). Some lamellas possess ridges and holes longitudinaly extending themselves from the base to the top, in only one of the sides. Small oscula (0.1 cm wide) are spread on all sides. The surface is finely conulose, with small depressions. The colour is yellowish-brown in vivo, turning purple after preservation in ethanol. Consistency is very soft and flexible. Skeleton: Choanosome with a delicate and irregular network of spongin fibers (Fig. 18 C). Bark with amber colour and thickness of 37–155 Μm (average 74 Μm). Thick pith can be black or amber with thickness of 7 –35 Μm (average 19 Μm; Fig. 18 D). Distribution: Provisionally endemic from the northeastern Brazilian coast (Fig. 16 B). The type locality is the Abrolhos region, located in the southern limit of the northeastern Brazilian sector. Ecology: The four collected specimens were photophilous, and always found on the upper part of rocks or coral heads. They were collected in depths that varied from 5 to 22 m. Etymology: The name of this species is related to its resemblance to a lettuce (lactuca in Latin). Remarks: Other species which bear some resemblance to A. lactuca sp.n. are A. alcicornis sp.n., A. cristagallus sp.n. and A. solangeae sp.n. However, although they all possess a lamellar form, none of them have centrally anastomosed lamellas as seen in A. lactuca sp.n. TABLE VIII: Spongin fibres’ measurement data for Aplysina lactuca sp.n. (in micrometers; S.D. = Standard Deviation and N= 30). Specimens Locality* Fibers Piths Thinnest Mean Thickest S.D. Thinnest Mean Thickest S.D. Holotype Abrolhos, BA 62.5 90.3 155.0 18.6 7.5 18.2 35.0 5.7 MNRJ 5477 Paratype Corumbau, BA 37.5 63.2 75.0 9.8 15.0 25.3 35.0 4.8 MNRJ 5276 Paratype Corumbau, BA 55.0 69.3 90.0 8.4 15.0 24.7 35.0 6.6 MNRJ 5278 * BA, Bahia State.Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 33-35, DOI: 10.5281/zenodo.17887

Aplysina solangeae Pinheiro, Hajdu & Custódio, 2007, sp.n.

Aplysina solangeae sp.n. (Fig. 22 C, 23 C–E, 25, Tab. XII) Holotype: MNRJ 4173, Salvador´s Yacht Club (Salvador, BA, 12 º 59 ' 58.2 '' S - 38 º 31 ' 54.2 '' W), 3–5 m depth, E. Hajdu coll., 4 /VII/ 2001. Paratype: MNRJ 2578, Porto da Barra, Forte de Santa Maria, (Salvador, BA, 13 º03' S - 38 º 32 ' W); E. Hajdu coll., 30 /VII/ 1999. Additional material: MNRJ 5268–5274, Salvador´s Yacht Club (Salvador, BA, 12 º 59 ' 58.2 '' S - 38 º 31 ' 54.2 '' W), U.S. Pinheiro coll., 3 m depth, 29 /I/ 2002). MNRJ 8675, Parque Estadual da Pedra da Risca do Meio (Fortaleza, CE, 03º 35 ' 889 S- 38 º 23 ' 481 W), 22 m depth, E. Hajdu coll., 14 /VII/ 2004. Diagnosis: Predominantly lamellar form in a semi-radial arrangement. Lamellae moderately stout, with very short, slender digitiform projections in their apices, and oscula situated at the top of discrete volcanifom projections. Live colour yellow or yellow with purple stains, turning pinkish-brown or dark-brown after preservation in ethanol. Consistency very soft. Description of the species: The specimens have a predominantly lamellar form, with moderately stout lamellae (frequently ca. 1 cm thick) in a semi-radial arrangement, but digitiform processes anastomosed to the main lamella can also be observed. They have a maximum of 11 cm in height and 7 cm in width. In the apex of the lamellae there are several short, slender digitiform projections up to 0.4 cm in height, frequently in tufts, that are released from the sponge after preservation in alcohol (Figs. 23 D–E, 25 A–B). The small oscula of 0.15 cm in diameter are normally restricted to the upper part of the lamella on top of small volcaniform projections, but oscula of this type in the sides of the sponge are not rare. The surface is finely conulose, with small, irregular and shallow depressions. The colour is yellow or yellow with purple stains in vivo, turning pinkish-brown or dark-brown after preservation in ethanol. Consistency is very soft. Skeleton: Choanosome with a delicate and irregular network of spongin fibers (Fig. 25 C) with amber coloured bark 37–158 Μm thick (average 77 Μm) and a thick pith that can be black or amber and is 11–55 Μm thick (average 24 Μm; Fig. 25 D). Distribution: So far endemic from the northeastern Brazilian coast (Fig. 22 C). Ecological observations: Bahian specimens were collected in very shallow-waters, with a maximum of 4 m depth. The specimen from Ceará was collected with scuba diving at 22 m depth. In both cases, water temperature is warm, tropical, throughout the year. Etymology: The specific name solangeae is dedicated to Prof. Dr. Solange Peixinho, for her enthusiastic scientific supervision of EH’s first steps on the taxonomy of sponges, her always warm welcome and priceless logistic support to every collecting trip undertaken in the Salvador area since 1987, her over 30 years teaching Bahian sponges to hundreds of biology students, and a similar time dedicated to the study of varied aspects of sponge biology in the State of Bahia. TABLE XII: Spongin fibres’ measurement data for Aplysina solangeae sp.n. (in micrometers; S.D. = Standard Deviation and N= 30). Specimens Locality* Fibers Piths Thinnest Mean Thickest S.D. Thinnest Mean Thickest S.D. Remarks: Despite the large morphologic plasticity observed in the collected specimens, all share the diagnostic characters (relatively stout lamellar form; short, slender digitiform projections; small oscula spread all over the sponge and soft consistency). Aplysina solangeae sp.n. is the closest species to A. fulva as far as external morphology goes. Aplysina fulva is another species which shows large morphologic plasticity and may have a lamellar shape in a few cases. However, the latter differs from A. solangeae sp.n. because its consistency is only rarely soft, it is typically digitiform and without short, slender digitiform projections (even when lamellate), while the new species described here is predominantly lamellar with short, slender digitiform projections. Lamellate specimens of A. fulva do not show lamellae in a semi-radial arrangement, and their lamellae also tend to be more slender and smooth.Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 44-46, DOI: 10.5281/zenodo.17887

Aplysina pseudolacunosa Pinheiro, Hajdu & Custódio, 2007, sp.n.

Aplysina pseudolacunosa sp.n. (Figs. 22 B, 23 A–B, 24, Tab. XI) Holotype: MNRJ 5478, western coral head (patch reef, Parque Nacional Marinho dos Abrolhos, BA, 17 º 57 ' 703 '' S - 36 º 14 ' 795 '' W), 18 m depth, E. Vilanova and G. Muricy coll., 03/III/ 2002. Paratypes: MNRJ 4581, stn. 34 (shallow, start: 20 º 24 ' 10 '' S - 39 º 55 ' 35 '' W, finish: 20 º 24 '08'' S - 39 º 55 ' 33 '' W, off Vitória, ES), 50 m depth, Programme REVIZEE Central V coll., 15 /VII/ 2001. MNRJ 4665, stn. 2 (13 º 38.98 ' S - 38 º 45.94 ' W, BA), 55 m depth, Programme REVIZEE Central V coll., 02/VII/ 2001. MNRJ 5461, coral head (patch reef, Parque Nacional Marinho dos Abrolhos, BA, 17 º 57 ' 703 '' S - 36 º 14 ' 795 '' W), 18 m depth, E. Esteves and G. Muricy coll., 03/III/ 2002. Additional material: MNRJ 1549, Pirapama´s Shipwreck (Recife, PE), 20 m depth, G. Muricy coll., 11 / II/ 1998. MNRJ 3048, Canoa Quebrada (CE), 25 m depth, M. Guimarães coll., II/ 2000. MNRJ 4299, stn. 7 (shallow, Royal Charlotte Bank, BA, start: 16 º 19 ' 55 '' S - 38 º 14 ' 39 '' W, finish: 16 º07'00'' S - 38 º 10 ' 20 '' W), 40 m depth, Programme REVIZEE Central V coll., 30 /VI/ 2001. MNRJ 4359, BA (15 º 34 '08'' S - 038º 49 ' 81 '' W), 20 m depth, Programme REVIZEE Central V coll., 01/VII/ 2001. MNRJ 4420, stn 2 (shallow, off Ponta de Castelhanos, SE of Morro de São Paulo, BA, 13 º 38 ' 98 '' S - 38 º 45 ' 94 '' W), 55 m depth, Programme REVIZEE Central V coll., 02/VII/ 2001. MNRJ 4669, stn 45 (shallow, ES, 20 º 40 ' 70 '' S - 34 º 35 '03'' W), 108 m depth, Programme REVIZEE Central V coll., 12 /VII/ 2001. Diagnosis: Specimens consist of tubes or small globules covered with pronounced ridges which render the surface very irregular. Abundant, variably large digital projections stem from nearly the entire surface of specimens. Specimens always turn beige or purple upon fixation, never black. Description of the species: Specimens vary from globose to tubular, with a maximum of 25 cm in height and 4 cm in width (holotype), covered by ridges, which render the surface very irregular (Figs. 23 A–B, 24 A). These ridges cover the entire surface of the specimens, are always short, and surround shallow valleys. Some small, slender digitiform projections are also observed. These projections can be anastomosed or not with other such ridges or the sponge main body. The sponges possess large apical pseudoscula, 1.5 cm in diameter, sometimes with an iristype diaphragm. Small oscula are also observed on the outer sides of the sponge. In some samples, small globes with apical oscula are present in the digitiform projections. The surface is finely conulose. The colour is bright yellow and beige in vivo, turning purple or brown after preservation in alcohol, never black. Consistency was firmly compressible. Skeleton: Choanosome with a delicate, irregular network of spongin fibers (Fig. 24 B) with amber coloured bark and width of 22–167 Μm (average 85 Μm), and a thick, black or amber pith with 8–47 Μm width (average 20 Μm; Figs. 24 C). The presence of spongin fibers wrapped and excavated by filamentous structures, possibly fungi, was observed in some specimens (MNRJ 4420, 4665) as in Aplysina muricyana sp.n.. Distribution: Brazilian Province (Fig. 22 B): Ceará State, Pernambuco State, Bahia State (Parque Nacional Marinho dos Abrolhos) and Espírito Santo State. Bathymetry: Found between 18 and 55 m depth. Etymology: The new species name, pseudolacunosa, stresses the superficial resemblance of its large inter-ridge depressions (valleys) to the surface lacunae of A. lacunosa. Remarks: The species most closely related to A. pseudolacunosa sp.n. is A. lacunosa, which also possesses tubes with a very marked relief. A. lacunosa has a surface characterized by irregular, reticulated ridges, producing caliciform depressions of variable depth, observed even in the smallest studied specimen (MNRJ 3557). We have analyzed a series of individuals (N = 10) of different sizes of A. pseudolacunosa sp.n. and they all possess a more open arrangement of ridges and valleys, as well as globular projections, instead of the narrow caliciform depressions formed amidst the reticulate arrangement of ridges in A. lacunosa. Both species have different spongin fibers too. A. lacunosa possesses a thinner (in average 16.6 Μm), irregular pith, while in A. pseudolacunosa sp.n. they are thicker (in average 20 Μm) and regular. Aplysina fistularis has also been observed to possess abundant projections of diverse morphologies. These are never similar to those reported here for A. pseudolacunosa sp.n., which cover the entire specimens, are always short, and produce shallow valleys. Specimen MNRJ 4669 was dredged at 108 m and presents a different morphology. It is globose, smaller and without projections, making its assignement to A. pseudolacunosa sp.n. rather uncertain. Thus, the occurrence of this species at such a depth is still doubtful. TABLE XI: Spongin fibres’ measurement data for Aplysina pseudolacunosa sp.n (in micrometers; S.D. = Standard Deviation and N= 30). Specimens Locality* Fibers Piths Thinnest Mean Thickest S.D. Thinnest Mean Thickest S.D . * BA, Bahia State; CE, Ceará State; PE, Pernambuco State; RN, Rio Grande do Norte State.Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 41-44, DOI: 10.5281/zenodo.17887

Aplysina cristagallus Pinheiro, Hajdu & Custódio, 2007, sp.n.

Aplysina cristagallus sp.n. (Fig. 9 C, 15, 16A) Holotype: MNRJ 3528, Pedra do Silva (Reserva Extrativista de Corumbau, Prado, BA), 8 m depth, G. Muricy coll., 14 /XI/ 1999. Diagnosis: Thinly lamellar habit, similar to a rooster’s crest, covered by rows of small apical digitiform projections bearing a darker colour on their apices. Bluish gray colour in vivo, turning into a light shade of brown in ethanol. Description: The sponge is lamellar, somewhat resembling a rooster’s crest. It is 0.3 cm thick, 7 cm long and 7 cm high. The apical portion of the crest has many small apical digitiform projections 0.6 cm high (more than 50) distributed in rows. The small apical digitiform projections remain in the preserved specimen and show their apical portion darker than the base, which has the same colour of the rest of the sponge (Figs. 15 A–B). In the upper part of the crest the sponge attains 1 cm in thickness. Small oscula appear on the apex and sides, where small projections are observed also bearing fistules on their top. The surface is very coarsely finely conulose. The colour is bluish gray in vivo (Fig. 9 C), which turns into a light shade of brown after preservation in ethanol. Consistency is very soft and flexible. Skeleton: Choanosome with a delicate and irregular network of spongin fibers (Fig. 15 C). Bark with amber colour and 46–232 Μm thickness (average 117 Μm). Thick, black or amber pith, 13–50 Μm thick (average 32 Μm; Fig. 15 D). Distribution: Provisionally known only from the type locality off the State of Bahia (Brazil; Fig. 16 A). Bathymetry: The specimen was collected at 8 m of depth. Etymology: The name of the species is a noun in apposition and recognizes its similarity with the rooster´s crest (crest = crista, and rooster = gallus in latin). Remarks: The species which appears closest to A. cristagallus sp.n. is A. alcicornis sp.n.. Both possess a lamelar habit, less than 1 cm thick, with small apical digitiform processes. However, A. alcicornis sp.n. is stouter and has a harder consistency, is yellowish-brown in vivo becoming dark brown when preserved, and its small digitiform processes are larger, frequently ramified and located solely on top of projections. Moreover, the oscula on A. alcicornis sp.n. are situated inside depressions and are surrounded by iris-type membranes. In contrast, A. cristagallus sp.n. is rather slender and possesses a very soft consistency, is bluish gray in vivo turning into a light shade of brown after preservation, and presents more numerous fistules which are organized in crests and bear darker apical portions. Both species are thus considered well distinguished from one another.Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 30-31, DOI: 10.5281/zenodo.17887

Aplysina alcicornis Pinheiro, Hajdu & Custódio, 2007, sp.n.

Aplysina alcicornis sp.n. (Fig. 9 B, 11 D, 14, Tab. VII) Holotype: MNRJ 5473, Coral Head at Parque Nacional Marinho dos Abrolhos (BA), 12 m depth, G. Muricy and E. Esteves coll., 02/III/ 2002. Paratype: MNRJ 5472, 5474, southeast Patch Reef (Parque Nacional Marinho dos Abrolhos, BA), 12 m depth, G. Muricy and E. Esteves coll., 02/III/ 2002. Diagnosis: Lamellar habit with flattened digitiform branches, conspicuous small apical digitiform projections and oscula with iris-type diaphragms situated in depressions on the surface of the sponge. Specimens turn to dark-brown in ethanol. Description of the species: The specimens are lamellar, with 11 cm in width 10 cm in height and 1 cm in thickness. The general appearance somewhat resembles the fire coral Millepora alcicornis Linneus, 1758, larger digitiform projections being observed at the base and the apex of the specimens. In the upper part of these larger projections, several much smaller apical digitiform projections up to 1 cm long, can be observed (Fig. 9 B). These fistules are frequently branching dichotomously or trichotomously, and remained on specimens after preservation (Figs. 14 A–B). Oscula 0.3 cm in diameter, bearing iris-type diaphragms are situated in depressions which occur close to the apical portion of the lamellas. Other oscula, with 0.1 cm, open on the sides of the sponge. The surface is finely conulose. The colour is brown or yellow in vivo, turning to dark-brown after preservation in alcohol. Consistency is compressible. Skeleton: Choanosome with a delicate and irregular network of spongin fibers with amber coloured bark and thickness of 40–142 Μm (average 92 Μm) and a thick black or amber pith with thickness of 15–82 Μm (average 24 Μm; Figs. 14 C–D; Table VII). In the small apical digitiform projections, the fibers present a dendritic arrangement. In these structures the fibers are composed mostly by the pith, with only a thin layer of bark. TABLE VII: Spongin fibres’ measurement data for Aplysina alcicornis sp.n. (in micrometers; S.D. = Standard Deviation and N= 30). Specimens Locality* Fibers Piths Distribution: Provisionally known only from Parque Nacional Marinho dos Abrolhos, (Bahia State, Brazil; Fig. 11 D). Ecology: All the specimens were collected at 12 m depth, attached on the sides of coral heads. Etymology: The name of this species stresses its resemblance to the fire coral Millepora alcicornis. Remarks: The species which most closely resembles A. alcicornis sp.n. is A. cristagallus sp.n. which is described below. Aplysina insularis sensu Zea (1987; Pl. 6, fig. 5) bear some resemblance to A. alcicornis sp.n. (non A. insularis of authors). Nevertheless, Zea´s specimen is not considered conspecific on the basis of its markedly anastomosed cluster of otherwise cylindrical branches topped by oscula, which contrasts to the overall flattened habit of the new species.Published as part of 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, pp. 1-51 in Zootaxa 1609 on pages 29-30, DOI: 10.5281/zenodo.17887