Checklist, diversity descriptors and selected descriptions of a highly diverse intertidal sponge (Porifera) assemblage at Costa do Descobrimento (Bahia, Brazil) Author Bettcher, Larissa 0000-0002-6341-8207 Author Fernandez, Julio C. C. 0000-0002-6785-5041 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0000 - 0002 - 6785 - 5041 Author Gastaldi, Marianela 0000-0002-2494-016 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & Departamento de Biología, Escuela Superior de Ciencias Marinas, Universidad Nacional del Comahue, San Martín 247, 8520 San Antonio Oeste, Río Negro, República Argentina & CONICET-Centro de Investigación Aplicada y Transferencia Tecnológica en Recursos Marinos Almirante Storni, G ̧ emes, 1030 San Antonio Oeste, República Argentina. https: // orcid. org / 0000 - 0002 - 2494 - 0166 Author Bispo, André 0000-0003-4025-6839 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0000 - 0003 - 4025 - 6839 Author Leal, Camille V. 0000-0002-1175-6896 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & Departamento de Genética, Centro de Ciências da Saúde, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, 21941 - 902, Cidade Universitária, Rio de Janeiro, RJ, Brazil. https: // orcid. org / 0000 - 0002 - 1175 - 6896 Author Leite, Dora 0000-0003-3855-8192 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0000 - 0003 - 3855 - 8192 Author Avelino-Alves, Dhara 0000-0002-6850-741X Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0000 - 0002 - 6850 - 741 X Author Clerier, Pedro H. B. 0009-0001-5882-3208 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0009 - 0001 - 5882 - 3208 Author Rezende, Dafinny 0009-0008-6554-9754 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0009 - 0008 - 6554 - 9754 Author Gulart, Clara M. R. 0009-0003-8761-3669 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0009 - 0003 - 8761 - 3669 Author Pinheiro, Ulisses 0000-0003-3658-1372 https: // orcid. org / 0000 - 0003 - 3658 - 1372 Author Hajdu, Eduardo 0000-0002-8760-9403 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s / n, 20940 - 040, Rio de Janeiro, RJ, Brazil & https: // orcid. org / 0000 - 0002 - 8760 - 9403 text Zootaxa 2023 2023-05-03 5277 3 443 489 http://dx.doi.org/10.11646/zootaxa.5277.3.2 journal article 53292 10.11646/zootaxa.5277.3.2 b55f10ed-45cb-4496-9ec1-8cac7a557cea 1175-5326 10268325 ADA46B20-63F6-4AB7-8FE8-1D0989662E6B Acanthotetilla walteri Peixinho, Fernandez, Oliveira, Caíres & Hajdu, 2007 ( Tab. 1–2 , 10 ; Fig. 13A–G ) Studied material. MNRJ 21180 — P.M.M. Coroa Alta ( 16°14.056′ S , 38°57.252′ W , Santa Cruz Cabrália, BA , Brazil ), intertidal, coll. Leite, D. , 25 th March 2019 . Additional material. MNRJ 23318 — Recife do Mutá ( 16°34.49117′ S , 39°00.40300′ W ), Santa Cruz Cabrália, BA , Brazil ), intertidal, coll. Fernandez, J.C. & Clerier, P. , 12/xii/2020 . Description. Encrusting ( 1 cm x 2 cm x 0.1 cm ) to endolithic. Surface irregular, abundantly pierced by spicules. Consistency firm, but compressible. Color in vivo not recorded, and light beige in fixative ( Tab. 10 ). Skeleton. Ectosomal skeleton unspecialized, merely a radial arrangement of abundantly triaenes (protriaenes and prodiaenes) and oxeas piercing the surface ( Figs. 13A–B ). Sigmaspires concentrated slightly underneath the surface ( Fig. 13C ); some acanthoxeas scattered too. Choanosomal architecture radially organized by oxeas and rhabdome of protriaenes, with abundantly and scattered acanthoxeas and sigmaspires ( Figs. 13A, E ). Spicules. Megascleres ( Tab. 10 ; Figs. 13A–G ): protriaenes or prodiaenes ( Figs. 13A–B ), rhabdomes slightly stouter right beneath the cladome, gradually thinning to the apex (mostly broken),> 290–524.4–972 x 1–2.1–4 µm (n=7), cladi 30–44.7–59 x 1–2.0–3.8 µm (n=10); acanthoxeas ( Figs. 13C, F–G ), slightly curved centrally, sharp points, heavily spined, 233–272.6–371 x 9–15.4–21 µm; and oxeas ( Fig. 13F ), slightly curved, sharply pointed, 704–972.5– 1180 x 6 –9.3– 12 µm (n=9). Microscleres ( Tab. 10 ; Figs. 13C ): sigmaspires (13C), 9–12.0–14 µm. FIGURE 12. Haliclona ( Reniera ) tubifera ( George & Wilson, 1919 ) , MNRJ 22235: A , specimen in life, with arrow pointing to an oscule; B , another specimen (MNRJ 21160) in life; C , longitudinal section of the surface showing an iso- to subisodictyal reticulation in the ectosomal region; D , transverse section showing an irregular iso- to subisodictyal reticulation and rounded channels in the choanosomal region; E , oxeas. TABLE 9. Haliclona ( Reniera ) tubifera ( George & Wilson, 1919 ) : comparative data on morphology, spicules and known distribution. Spicule measurements in micrometers (µm) are presented as minimum–average–maximum (or minimum–maximum) of length x thickness. Data not reported are represented as n.r.

Sensu	Morphology features	Megascleres (oxeas)	Distribution (Depth)
George & Wilson, (1919) as Reniera tubifera	Habit irregular (reticular system of anastomosing cylindrical branches with oscules at their tips. Surface nr. Consistency quite fragile (not soft). Color pink or reddish purple, varying to brown in vivo , and fading quickly in (EtOH)	125‒170 x 3‒8	North Carolina (USA) / Atlantic (low tide)
Hechtel (1965) as H. hogarthi	Habit branched (numerous, slender, anastomosing branches). Surface microtuberculate (even but wrinkled) and punctiform. Oscules n.r. Consistency limp, soft, compressible and somewhat resilient. Light reddish-purple in vivo , and pale to dark drab in fixative (EtOH)	117‒157 x 5‒9	Jamaica, Caribbean / Atlantic (0.3‒0.6 m)
de Weerdt (2000)	Habit very variable, commonly cushion-shaped base with sev- eral oscular elevation (volcano or chimney). Surface smooth, even, slightly punctate. Consistency soft, compressible and fragile. Color different shades of purple and pink in vivo , and in fixative n.r.	ranging from 67.3‒90.5 x 2.8‒4.6 to 143.8‒178.6 x 6.9‒9.3 in different individuals	North and South Carolina (USA), Gulf of Mexico and Caribbean / Atlantic; and Panama / Pacific (0.5‒10 m)
MNRJ 20562; MNRJ 21160; MNRJ 22235	Habit thinly to thickly encrusting, occasion. like tubular struc- ture laying parallel to substrate, or lobate. Surface slightly rough and punctate. Oscules at the apex of the tubular struc- ture, or at the short lobate or volcano elevations (<1 cm high). Consistency soft and compressible. Color beige to light-purple in vivo , and light beige in fixative (EtOH).	respectively: 101.2‒126.5‒147 x 1.5‒5.5‒8; 90.8‒118.2‒137 x 2.4‒5.0‒8.2; 101‒128.1‒144.6 x 2.1‒5.1‒7.8	Costa do Descobrimento, Bahia, Brazil / Atlantic (intertidal)

Distribution. Only known from northeast Brazil : Camaçari ( type locality), the northern sector of BA, and Santa Cruz Cabrália, the southern sector of BA—new record. Ecology. The specimen was growing in a piece of hard coral ( viz. , not in live), which is associated with calcareous algae and hydrozoans. Remarks. Our material matches the original description of A. walteri by Peixinho et al. (2007) , as amended in Fernandez et al. (2012) . The species was originally described from northern Bahia ( 22 to 35 m depth) and had not yet been reported again. Here we are recording it from an intertidal area in southern Bahia . The endolithic/cryptic habit of A. walteri likely explains the few records of the species this far, which is absent from large taxonomic inventories such as those by Muricy et al. (2006 , 2011 ), Moraes (2011) and Moura et al. (2016) . FIGURE 13. Acanthotetilla walteri Peixinho, Fernandez, Oliveira, Caires & Hajdu, 2007 , MNRJ 21180: A , transverse sections showing megascleres (protriaenes and oxeas) radially and piercing to the surface; B , detail of protriaenes; C , ectosomal region with several sigmaspires; D , a reproductive bud on the surface and E , several acanthoxeas scattered throughout the choanosomal region; F , a large smooth oxea and acanthoxeas; G , an acanthoxeas in detail. TABLE 10. Acanthotetilla walteri Peixinho, Fernandez, Oliveira, Caires & Hajdu, 2007 : comparative data on morphology, spicules and known distribution. Spicule measurements in micrometers (µm) are presented as minimum–average–maximum (or minimum–maximum) of length x thickness. Only length is presented for sigmaspires. Spicules

sensu	Morphology features	Megascleres	Microscleres (sigmaspires)	Distribution (Depth)
Fernandez et al. (2012) ; review of the type material	Habit encrusting (up to 3.5 mm thick) to endolithic. Surface irregu- lar and smooth to slightly hispid. Consistency slightly compressible. Color in vivo n.r., and dark to light grey in fixative (EtOH)	Oxeas, 742‒995.1‒1232 x 5.4‒13.0‒21 Acanthoxeas, 238‒297.1‒378 x 28‒28.0‒28 Protriaenes (rhabdome; cladi) 308‒722.7‒1372 x 3.6‒3.8‒7.2; 18‒40.5‒58 x 1.8‒3.6‒7.2	7‒10.2‒18	Camaçari (northern Bahia), Brazil / Atlantic (22‒35 m)
MNRJ 21180	Habit encrusting (up to 1 mm thick) to endolithic. Consistency firm, but compressible. Surface irregular and mostly pierced by megascleres. Color in vivo not ob- served, and light beige in fixative (EtOH)	Oxeas, 704‒972.5‒1180 x 6‒9.6‒13 Acanthoxeas: 233‒272.8‒371 x 8.7‒15.7‒21.4 Protriaenes (rhabdome, cladi), 290‒540.1‒972 x 1‒2.1‒4; 30‒44.7‒59 x 1‒2.0‒3.8	9‒12.0‒14	Costa do Descobrimento, Bahia, Brazil / Atlantic (intertidal)

A bud made of spicules and spongin was observed ( Fig. 13D ). So far, a similar reproductive specialisation has not been reported from other known species of Acanthotetilla (cf. van Soest 1977 , Peixinho et al . 2007 , Fernandez et al. 2012 ). However, surface buds were reported from other tetillid sponges, for instance: Cinachyrella apion ( Uliczka, 1929 ) from the western Atlantic ( Ŗtzler & Smith 1992 ), C. australiensis ( Carter, 1886 ) from the Indopacific (cf. Ŗtzler & Smith 1992 ) and C. anatriaenilla Fernandez, Kelly & Bell, 2017 from the west Pacific ( Fernandez et al. 2017 ). The latter two species share spined microxeas, which resemble acanthoxeas of Acanthotetilla species. Although acanthoxeas and microxeas are acanthose monaxonic spicules ( Fernandez et al. 2012 ), the origin of both (as homologous or not) has still not been determined by phylogenetic analysis ( Szitenberg et al. 2013 ). Some important observations about spicules of A. walteri should be made here, a consequence of imprecisions in the original description of the species. In regard to protriaenes, a single category is present, with rhabdome thickness of 3.6–3.8–7.2 µm, as already pointed out in Fernandez et al. (2012) . As for the acanthoxeas, also only a single category is present, the smaller ones shown in Peixinho et al. (2007) being misinterpreted due to the angle of the spicules in the SEM image. Micrometric values for these spicules are 238–297.1–378/28–28.0–28 µm as mentioned in Fernandez et al. (2012) , and reproduced in Table 10 .